Essentials of Nematodology Edited by К. [. Skrjabin Vol. ХХИ ИМ. Ivashkin, А. A. Sobolev, and L. A. Khromova Camallanata of Animals and Man and Diseases Caused by Them Translated from Russian AKADEMIY А NAUK SSSR - GEL'MINTOLOGICHESKAY A LABORATORIY А OSNOVY NEMATODOLOGII, TOM ХХП Academy of Sciences of the USSR - Helminthological Laboratory ESSENTIALS OF NEMATODOLOGY, Volume XXII Edited by Academician К.1. Skrjabin И, М. lvashkin, A. A. Sobolev, and L. A. Khromova Camallanata of Animals and Man and Diseases Caused by Them (Kamallanaty zhivotnykh i cheloveka i vyzyvaemye imi zabolevaniya) Izdatel'stvo “Nauka” Moskva 1971 Translated from Russian Israel Program for Scientific Translations Jerusalem 1977 ТТ 76-50005 The Publication of this Translation was Supported by THE UNITED STATES — ISRAEL BINATIONAL SCIENCE FOUNDATION Copyright © 1977 Keter Publishing House Jerusalem Ltd. Cat. No. 61223 1 ISBN 0 7065 1572 2 Translated by Hilary Hardin Printed and bound by Keterpress Enterprises Jerusalem, Israel Available from the U.S. DEPARTMENT OF COMMERCE National Technical Information Service Springfield, Va. 22151 CONTENTS ОИ Gees ios В В hoa axel Gud’ o paket sien sli Bieeus forse ie 1 General Part Morphological and Anatomical Description of Camallanata ................. 3 BOE alld зе АЕ eS WOU Vaden: tie ead acne оке сене 3 (Gi CUS rene ERROR RC Iscal > ©: HATARSUAT MRE aA: ere Ae Bel eran Se eee 3 БИС ое OF te серва Chem ое о а еее 4 ПЕНИЯ ЕО О О О С о О ее «5 7 Ве а а Е а ви 10 ВБ оБуог тие у Of the\Gamiall lama taeiy. iii sioyer о Ноа, 11 Phylogenetic relationships of the Camallanata and their reflection in Ме стана Or EME пота ег ее о ен 15 Biological characteristics of the suborder Camallanata Chitwood,1936 .... 16 Ontogenetic development in the superfamily Camallanoidea ........... 16 Ontogenetic development in the superfamily Dracunculoidea ......... 18 Ontogenetic development in the superfamily Anguillicoloidea ......... 21 Systematic Part Subprder Camallanata, Chitwood ; LOSGI ее че 23 Supetiamily, Camallanoidea) Giavassos,, VOZORRN В.А. CO. we ole 23 Family Camallanidac Raillier andi Elenny,)) VOLS rae): и, 5 24 Subfamily Canrallaninaeey cli; побои Meee METS aoe. SN 26 Genus Camallanus (Railliet and Henry, 1915) Yeh, 1960 ..... 26 Genus Camallanides Baylis and Daubney, 1922 ............ 85 Genus Paracamallanus Yorke and Maplestone, 1926 ....... 95 Gents Руслана mice lays ба ее а 97 CEenusySieTpliMemMay, ей а 1960. соевое. tye -elore ЕО 98 GeniseZ ее тан е ск Ване tale 114 SublamulysProcamallaninae МЕ LOCO о о ны 130 Cems РТо а аа ts! Ва У ТВ ее 130 ВЕ Орто а аа РИ "Обе WOOL еее 164 Нега Ао еоГотаед п. SUPCMAM, „еее ео езеевье 209 База узде сомаае Nama Sti y Та, не ее 210 Сени да софа Уашмарие и 193550 x \sjarcpepeud «rege «fellate 3 210 Family Philyctainophomdae Romans; 1960, о На ыы о На 214 ill Genus Phliyctainophora Steiner, 1921 ....:.... ее еченые 214 Family Skrjabillanidae Schigin and Schigina, 1958 .................. PAY Genus Skrjabillanus Schigin and Schigina, 1958 .......... 218 Genus Agrachanus Tichomirova in litt. .......ssseseeees 220 Genus MWolimartay WoraveCn LOGGUe «cis... ее ео 223 Family Tetanonematidae Skrjabin and Schikhobalova, 1948 .......... 228 Genus Tietanonemia Steiner, 19371... оса ве 228 Superfamily Dracunculoidea Cameron, 1934 ............ еее конноь 231 Family Dracunculidae Leiper, 1912 —........ еее внаеюно 232 Subfamily Dracunculinae Stiles, 1907 ......... STA AE, 232 Genus’ Dra cunmewlus Веера 109 ооо р Subfamily Avioserpentinae Wehr and Chitwood, 1934 emend. Chitwood: T9Sa) “ee ov es ae en os + ie eres Sis a овес 252 Genus’ Avioserpen’s “Webriand ‘Chitwood, 1954 ^^... вай 253 Subfamily Micropleurinae Baylis and Daubney, 1922 ............ 270 Gents Mictroplewra Linstow,: 1900 7 oo ot. oe. sic os cee 270 Family Philométridae Baylis and Daubney, 1926 -...... о. сосвааа 277 Subfamily Philometrinae Yamaguti, 1935 ........... В - 277 Genus’ P hiliome tia "Costa аа 279 Subgenus Philometra Rasheed, 1963 ......... oe > 279 Subgenus Alinem/a) Rasheed, 1309 «лье ere ey 308 Subgenus Ranjhinema Rasheed, 1963 Ls oboe es _-. 309 Genus “Buckley ella Rasheed 1903 а MRS oc 811 Genis Tchtihyotilaria Yamaguti, 1935... рой 312 Genus Nitomwema ‘Khalil, 1960 :.... се. co vue sie cen eee 314 Genus Philometroides: Yamagutl, 1985 .-..... ой SiT Genus Pseudophilometroides Parukhin, 1966 ....... 335 Genus Ruimai . Travassos, 1960 80.. peel eya tied ier leet sence eee 336 Genus Thwaitia Rasheed, 1968.17. cabianclame.. 31000 337 Subfamily Philoneminae n..subfama,: :9% На золы SE 348 Genus Philonema Kuitunen-Ekbaum, 1988 1... vilewaidud.... 348 BiBILOPTA DIY "ао ото ис оне 359 indexes. Of WEITETA ана реет а... осень ie osc shes oe 377 5 FOREWORD Volume XXII of "Еззепа1$ of Nematodology'"' contains descriptions of all species of the suborder Camallanata Chitwood, 1936, which parasitize in man and animals throughout the world. By the collection and study of the large body of data published in the Soviet and world literature during the last 150 years and its analysis, the authors have obtained a radical revision of the classification of these interesting helminths. The large superfamily Cucullanoidea has usually been included in the sub- order Camallanata although the species of this superfamily show morphologi- cal characters which are not characteristic for typical Camallanata. V. М. Ivashkin and myself (1968) removed the Cucullanoidea from the suborder Camallanata and established for them the new suborder Cucullanata with the two superfamilies Cucullanoidea Ivaschkin, 1962 and Gnathostomatoidea Skrjabin and Ivaschkin, 1968. The classification of the Camallanata underwent another revision when the present authors established the new superfamily Anguillicoloidea which, together with the superfamilies Camallanoidea Travassos, 1920 and Dracunculoidea Cameron, 1934, form the suborder Camallanata. The new superfamily contains four families, including the family Skrjabillanidae, established in 1958 by A. A. Shigin and N. G. Shigina, who placed it in the superfamily Dracunculoidea. The family Skrjabillanidae originally contained only the single genus Skrjabillanus, but the authors of this work included also the genera Agrachanus Tichomirova п. в. and Molnaria Moravec, 1968. The authors consider the new superfamily Anguillicoloidea, the species of which show a combination of the characters of such different superfamilies as the Camallanoidea and the Dracunculoidea, as transitional between these two superfamilies. In the superfamily Dracunculidae Cameron, 1934 the authors place only the families Dracunculoidea Leiper, 1912 and Philometridae Baylis and Daubney, 1926. The first family preserves its former structure, but the Philometridae, which previously contained only the subfamily Philometrinae Yamaguti, 1935, is considered also to contain the new subfamily Philoneminae, with the single genus Philonema Ekbaum, 1933 because of the characteristics 6 of the life cycle of the type species of the genus, Ph. oncorhynchus, particularly the long time (about 3 years) of maturation in the definitive host, and also the presence of polynuclear esophageal glands. Of the life cycles of species of Camallanata, the ontogeny of only 20 species has so far been studied. None of the species studied are strictly specific for their intermediate hosts or for their localization. The species of this suborder develop (to third-stage larvae) in the body cavity of lower crustaceans: Cyclops, Diaptomus, and Branchiura. This monograph on the Camallanata, the first and only in the literature intended for biologists and helminthologists in the medical and veterinarian field, will enable specialists to determine all species of Camallanata pa- rasitizing in animals throughout the world. K. I. Skrjabin CGE EB РЗАЗЕИРАВЕУТ MORPHOLOGICAL AND ANATOMICAL DESCRIPTION OF CAMALLANATA Form and size of the body The form and size of the body of Camallanata vary markedly and cannot be used as typical characters of the suborder. In some groups the length of the body is not over 22—23 mm in males and 25—28 mm in females (superfamily Camallanoidea), while in others the body is very long and thin; for example, females of Dracunculus medinensis reach a length of 1.5m (superfamily Dracunculoidea). The body of Camallanata is of more or less uniform width most of its length, narrowing toward the ends. Sexual dimorphism is generally ex- pressed inthe difference in size between males and females, whichis sometimes very marked (family Dracunculidae). Cuticle The cuticle of Camallanata is usually smooth and without any differentia- tions. Transverse striation is generally present and sometimes well developed. Tornquist (1931) stated that in the Camallanidae the cuticle consists of six layers, which are clearly distinguishable at the caudal end. The forms which are characteristically localized in the inner parenchymatous organs, connective tissue, and blood vessels have sometimes more or less de- veloped cuticular tubercles on the surface. Rasheed (1963) distinguished the following types of cuticle in the family Philometridae (Figure 1): 1. The cuticle is smooth, the most common type of cuticle, present in species of most genera. It has a smooth surface without ornamentation or thickenings (Figure 1,a)(Philometra Costa, 1845). 2. The cuticle has convexities or swellings in some places, "tubercles" and "'spines'' (Figure 1,b), which may be scattered over the whole surface or are concentrated in some areas (Philometroides Yamaguti, 1935). 3. The cuticle has dense, shining convexities on the body: "рШагз" and "сопез" which are formed by the cortical, fibrillar, and basal layers of the cuticle (Figure 1,c—h); they vary in form and size: sometimes needle- shaped as in Buckleyella Rasheed, 1963, sometimes cone-shaped as in Nilonema Khalil, 1960. In species of the genus Tetanonema from the vascular system of Cyclostomata cuticular tubercles are weakly developed. The females of most species of Camallanata have three teatlike forma- tions at the end of the tail; there are six of them in the females of Spiro- 8camallanus xenopodis. The larger the female, the less numerous are these formations, the largest specimens having two or one, or some- times none at all. The same is observed in the infective larvae. Lateral wings are not always present. 0.2mm FIGURE 1. Types of cuticle in the Philometridae: a — smooth (Philometra — general view); b — with “tubercles” and "spines" (Philometroides — general view); с — e — with “warts” and "rods" (с, d— Buckleyella — general view; е — part of cross section, enlarged); f — В —with"cones"(f— Nilonema gymnarchi — general view; 8 — Nilonema senticosa, magnified; h—same, Nilonema gymnarchi) (after Rasheed, 1963). Hematophagous Camallanata are red during life; fixed specimens are usually whitish yellow or brownish. Structure of the cephalic end Chitwood and Wehr (1934) and Chitwood and Chitwood (1950) summarized the data on the structure of the cephalic end of nematodes, particularly of the Camallanata. They noted that in the suborder Camallanata there are no ventrolateral papillae or lips, but there may be rudiments of lips or lateral valves; the latter are present in most species of the superfamily Camallanoidea. The mouth is hexagonal in the Procamallaninae. The outer ring consists of four papillae; the papillae of the inner ring are very small. Species of the superfamily Dracunculoidea have neither lips nor valves; the round mouth is enclosed in a thin circumoral membrane, on the outside of which cuticularized protuberances may be present (Dracunculus, 9Avioserpens) ог absent (Philometra, Micropleura) (Figure2). The inner ring of papillae is well developed; the outer ring consists of eight papillae. In Micropleura and Philometra all the papillae are dis- tinctly developed, while in Dracunculus and Avioserpens the median papillae of the outer ring are partly fused. In Dracunculus the interoventral and interodorsal papillae become fused in the female during development, but in the male the papillae maintain the normal arrangement. Species of Dracunculidae have a thickened cuticular shield which projects anteriorly, forming a circumoral protuberance, and it surrounds the anterior end of the esophagus posteriorly. The genus Dracunculus differs from Avioserpens inthe presence of a cuticular ring around the mouth on the cephalic shield characteristic for both genera. FIGURE 2. Arrangement of the cephalic papillae in the Philometridae (apical): a—Philometra (Philometra); b— Philometra (Ranjhinema); c—Philometra sydneyi Rasheed, 1963; d, e— Buckleyella buckleyi Rasheed, 1963; f— Nilonema senticosa (Baylis, 1927); g— Thwaitia Rasheed, 1963; В — Philometroides anguillae (Ishii, 1916); i—Philo- metra (Alinema) (after Rasheed, 1968). The superfamily Camallanoidea is characterized by а well-developed stoma in most forms and at least a distinct vestibule in others (Figures 3, 4). The superfamily Dracunculoidea has a rudimentary stoma. Amphids, 10 described for some genera of Camallanata, are situated near the lateral cephalic papillae; they are particularly well developed in the genera Dracunculus and Avios’erpens:. i) FIGURE 3. Structure of the buccal capsule of Camallanidae: | a a — bivalvate (apical and ventral); b — entire (apical and ventral) (after Yeh, 1960). The difficulty in the study of the cephalic end of species of Philometridae resulted in that about 30 species were described without mention of the cephalic papillae. In1963, during a study of the cephalic end of Philometridae in lactophenol, Rasheed found that cephalic papillae are always present in this group. She did not find other structures on the cephalic end of Philometridae. FIGURE 4. Structure of the cephalic end in the Camallanidae: a—Procamallanus; b = Spirocamallanus; с — Piscilania; 4 — Paracamallanus; е — Camallanus; f — Camallanides; g — Zeylanema; В — Serpinema (after Yeh, 1960b). Пл The number, size, form, and structure of the papillae vary considerably. For example, the cephalic papillae are so large in some genera that they form lobelike structures (Buckleyella and Thwaitia). These papillae are strongly muscular; nerve endings are not recognizable. In other forms the papillae are very small, and only the nerve endings are recognizable; the cuticular processes surrounding them are very small (Alinema). The cervical papillae are never very well developed in the Camallanata. They are situated near the nerve ring, slightly before or behind it. In the genera Avioserpens and Dracunculus the cervical papillae are also situated slightly behind the nerve ring and are difficult to distinguish, The cervical papillae are not mentioned indescriptions of other genera. Intestinal tract The mouth is slitlike, rounded or triagonal, and directed dorsoventrally. The buccal capsule of Camallanata shows a great diversity of structure in the different genera and is an important character for their determination. Higher taxa differ markedly in this character. The Camallanidae have a strongly developed chitinous armature of the buccal capsule and its adjacent parts. In the genus Camallanus the walls of the capsule form strong reddish brown ''valves"' with longitudinal ribs. Tornquist (1931) showed that there is no reason to consider these valves as a characteristic formation that would differentiate these species from others, as authors have done in the past. This is only a strong development of the walls of the buccal capsule, and not a differentiated morphological element. Such thickenings of the lateral walls of the buccal capsule are present also in other genera of Camallanidae and particularly well deveioped in the genus Camallanides. Species of three genera of Camallanidae have a cuticular ring between the buccal capsule and the esophagus; other genera show characteristic trident formations on the dorsal and ventral side of the body. All these formations are yellow or reddish brown. Chitwood and Wehr (1934) consider the genus Procamallanus, which has a hexagonal mouth and no cuticular ''valves,'' as the most primitive genus of Camallanidae. As regards the morphology of the stoma in the Camallanata, the authors note that despite its distinctive structure, divisions may be found in species of Camallanidae which resemble the divisions of the stoma of other genera of this family. The primitive character of the genera of Procamallaninae is expressed in the absence of ''valves"' and in that the stoma, which is rounded in cross section and not dorsoventrally compressed, is usually less developed than in other genera of this subfamily. A prostoma and telostoma can be distinguished, althoughthe various elements of the protorhabdion are not distinct. The stoma of the genera of Proca- mallaninae resembles in general that of the Thelaziidae (Spirurata). Chit- wood and Wehr think that the stoma of Paracamallanus,in whicha prorhabdion (from which the ''valves'' develop), mesorhabdion, and telo- rhabdion are distinctly developed, is the original type of the buccal section in genera of the subfamily Camallaninae. 12 The telostoma in the genus Camallanus is surrounded by а chitinous ring (or collar), which is homologous to the telorhabdion, Chitinous append- ages (tridents) are developed at the base of the prorhabdion. In the genera Camallanus and Paracamallanus two pairs of winglike formations, lateral to the mouth, are present оп the outside. The stoma is not divided in the genus Anguillicola, for which Yamaguti (1935) established а separate family, and in the genus Skrjabilla- nus of the family Skrjabillanidae Schigin and Schigina, 1958. The present authors and У. А. Tikhomirova, who studied the morphology and biology of the family Skrjabillanidae and clarified the life cycle of Agrachanus scardinii and Molnaria erythrophthalmi, place these two families in the new superfamily Anguillicoloidea. The stoma of the Dracunculidae is rudimentary. The suborder Camallanata contains forms in which the esophageal glands are primarily mononuclear; forms with polynuclear glands apparently ap- peared secondarily (Chitwood and Chitwood, 1950). The superfamily Camallanoidea is characterized by a relatively well- developed musculature and three esophageal glands, one of which is situated dorsally and the other two subventrally. The dorsal gland extends through both parts of the esophagus, and its duct opens before the nerve ring. The subventral glands are situated in the posterior part of the esophagus, and their ducts open into its anterior part. The nuclei of the glands are situated at the same distance from the anterior end, in the posterior part of the glandular part of the esophagus. The family Dracunculidae has strongly developed esophageal glands, the nuclei of which are long and narrow. The ducts open in the region of the esophagus. The esophagus of Camallanata was studied by Jagerskidld (1909), Magath (1919b), Tornguist (1931), and Hsu (1933а). It is not of uniform structure in the different groups of this suborder. In the Camallanidae the anterior part is muscular and the posterior part glandular. The an- terior part is slightly shorter than the posterior part. The posterior part projects slightly into the intestine. Magath (1919b) mentions three valve- like formations at the end of the esophagus. Tornquist (1931), who also studied Serpinema microcephalus, stated, however, that valves are absent at the end of the esophagus. Serpinema microcephalus has a divided esophagus. The dorsal opening of the gland is situated slightly behind the base of the stoma; the subventral openings are situated at the anterior end of the posterior part of the esophagus. There are 18 radial and six marginal nucleiinthe anterior part of the esophagus. The radial nuclei are arranged in four groups: two groups of three nuclei before the opening of the dorsal gland and two groups of six nuclei behind it. The marginal nuclei form two groups, one near the opening of the dorsal gland and the other between the third and fourth radial group. The.posterior part of the esophagus also has 18 radial nuclei in four groups and six marginal nuclei in two groups. The first group of six radial nuclei is situated near the anterior end of this part of the esophagus at the level of the openings of the subventral glands, the second group of six radial nuclei is situated slightly before the middle of this part of the esophagus, and the third and fourth groups of three radial nuclei are situated near the base of the esophagus. One marginal group is situated directly behind the openings of the subventral glands, the other near the third group of radial nuclei. The esophagus of the Dracunculidae was studied by Jagerskiold (1893), Strassen (1907), Mirza (1929), Hsu (1933b), and Yamaguti (1935). It may be cylindrical,as in Micropleura, club-shaped or barrel-shaped, as in Philometra, or divided into a short, narrow anterior muscular part and a long, wide posterior glandular part,as in Philonema,Dracunculus, and Avioserpens: in Dracunculus and Avioserpens the glandular part narrows at the nerve ring. The anterior part of the esophagus of Dracunculus, Avioserpens, and Philometra forms a hemi- spherical swelling; this is not the case in Philonema and Micropleura. In the genus Philometra the esophagus is usually widened around the mouth and has the form of a bulb; in some species the three lobes of the esophagus project from the mouth or become cuticularized and pointed, forming esophageal teeth. The lumen of the esophagus is usually wide in its anterior part, which in some species forms a cavity surrounded by the lobes of the esophagus. The esophagus of Nilonema is cylindrical and is of almost the same width its whole length. The genus Rumai hasa club-shaped esophagus; the anterior part is cylindrical, the posterior part gradually increases in width. In Philometra parasiluri the esopha- gus is entirely muscular. The other genera of Dracunculidae have either an esophagus divided into an anterior muscular and a posterior glandular part or glandular tissue situated in the dorsal part of the esophagus. The radial and marginal nucleiin Dracunculus, Avioserpens, and Micropleura have the same arrangement as in Camallanus. All these nuclei also show a uniform arrangement relative to the openings of the esophageal glands. The region behind the narrowing at the nerve ringin Dracunculus and Avioserpens corresponds to the posterior part of the esophagus of Camallanus. The dorsal esophageal gland and its nucleus are very large. The fourth and fifth groups of radial nuclei in Philometra are divided into two subgroups, but the marginal and radial nuclei have the same arrangement as in Dracunculus; in the latter genus, however, the subventral glands are very much smaller. Yamaguti described the genus Ichthyofilaria, the esophagus of which has a posterior glandular appendage as in Contracaecum. However, no histological study of this structure has been made. Chitwood and Chitwood (1950) stated that the genus Philonema shows an extreme development of the esophageal gland in the Dracunculoidea. This genus has a typical spiruroid-filarioid esophagus, and it is doubtful whether it should be placed in the suborder Camallanata. The three open- ings of the esophageal glands are situated in the posterior, markedly wider glandular part of the esophagus, and the esophageal glands are mononuclear. The radial nuclei in all six groups are distributed in groups of three, i.e., all six groups are subdivided. Since there are three groups of radial nuclei in the anterior muscular part of the esophagus, this part may be homologous to the anterior part of the esophagus in the Camallaninae; the position of the openings of the subventral glands support this view. The opening of the dorsal gland is displaced much farther posteriorly in these forms. Each esophageal gland contains several hundred nuclei of varying size which are often arranged in ''constellations, "а result of nuclear budding caused by gigantism. This may be analogous rather than homologous to the case of polynuclear glands in the Spiruridae. In the latter the many nuclei of the glands apparently developed by division as they are all of more or less the 14 1 сл same size. И Philonema developed independently, the gigantism of these glands in Dracunculus and Philometra may be correlated with the nonuniform nuclear division (amitosis?) in the glands of Philonema. Hsii(1933b) described "simple tubular''glandsin Philometra and Dracunculus, stating that other nematodes have "branched tubular'' glands. It is easy to make a mistake during the examination of these structures, espe- cially when the glands end in single or branched terminal lobules. For example, in Dracunculus oesophagea the subventral glands are complex ra- mified alveolar formations, while the dorsal gland is apparently a simple branched alveolar gland. The intestine of Camallanata is usually a simple tube ending in an anus, which is obliterated in adult females of Dracunculidae. Genitalia Males. The male genitalia consist of a single testis, which passes into the vas deferens and ends in the ejaculatory duct, which opens into the cloaca. The elements of the copulatory apparatus vary considerably in structure in the different families. The spicules are of different length in the Camallanidae. The left spicule is always shorter, with rounded section, and wings are absent. Itisless wide than the right spicule. The right spicule has usually, but not always, wings, and the left wing is wider than the right. The right spicule ends ina hookin Camallanides. Both spicules are pointed. The species of the family Dracunculidae have thin spicules pointed at the distal end. They are of equal or nearly equal length. Spicules are absent in the superfamily Anguillicoloidea. A chitinized plate is present in some forms. A gubernaculum is not present in all genera, It is nearly triangular in the Camallanidae. In the Dracunculidae the presence of a gubernaculum has been established in the genera Dracunculus, Avioserpens, Philometra, and Micropleura. A gubernaculum is absent in the Anguillicoloidea. The caudal wings of males are never very large. They are best developed in the family Camallanidae, in which they are relatively narrow. Caudal wings are absent in the family Dracunculidae except in some species of the genus Micropleura, which have only a right wing which is partly inter- rupted. There are two liplike processes at the cloaca in the genus Phil omietinias The genital papillae on the tail of males usually form two longitudinal subventral rows, some of the papillae being preanal, others adanal, and still others postanal. Pedunculate papillae are present in the Camallanidae, as these also have caudal wings; in other families there are indistinct sessile papillae. The arrangement of the caudal papillae inthe genus Dracunculus is characteristic. The preanal papillae are arranged in two rows, often densely; these rows are situated anteriorly, close to- gether; they then diverge, forming sides of an equilateral triangle, the base of which is the cloaca, The angle of divergence of the rows of papillae is about 90° in Dracunculus medinensis, but it is acute in ШО. оезо- phagea. The first two pairs of postanal papillae also have a distinctive 10 arrangement. The first pair is situated close together, but the papillae of the second pair are more widely spaced than the distance from the papillae of the corresponding side of the right pairs. The caudal part has apparently become shorter during evolution, resulting in this arrangement of the papillae. The caudal end of males of Camallanata may be moderately long, rounded, moderately pointed (Camallanidae), or cone-shaped (Dracunculus). The caudal end has the form of a truncated cone in the genus Philo- metra (there may or may not be papillae on the caudal end). It is conical in the genus Rumai. The males of the genus Philonema have a long, thin tail which tapers to a point. Females. The structure of the female genitalia shows marked dif- ferences in the different genera and families. The species of the family Camallanidae have one anterior ovary. This is followed by the oviduct, which passes into the uterus, ovejector, and the vagina, which is lined with cuticle. The vulva is usually situated in the middle of the body, but it may be displaced slightly anteriorly or posteriorly. The vulva of the genus Camallanus has distinctly projecting lips, one of which may project more markedly. The vulva forms a tubelike appendage in the genus Camallanides which is slightly dorsoventrally compressed. It projects above the surface and is directed posteriorly. The posterior uterus is either bluntly rounded at the end or it ends in a rudimentary ovary. In the genus Dracunculus the vulva is situated before the middle of the body only in young specimens; after fertilization it is closed by a plug of mucus and is eventually obliterated. The uterus is very large, saclike, and fills the body cavity in adults. There are two ovaries, which undergo reduction in adults. The female genitalia of the genus Philo- metra are of similar structure. All species of the suborder Camallanata are viviparous. HISTORY OF STUDY OF THE CAMALLANATA The suborder Camallanata was established by Chitwood in 1936, but in 1937 he proposed new names for some suborders of nematodes, including the Camallanata, which he renamed Camallanina (after Chitwood, 1937). Skrjabin and Shul'ts (1940) reinstated the original name, which we have also accepted here. Chitwood included two superfamilies in the suborder Camallanata: Camallanoidea Travassos, 1920 and Dracunculoidea Cameron, 1934. The original diagnosis of the suborder is as follows: Spirurida. Pseu- dolabia absent; lips also usually absent. Six rudimentary lips sometimes present. Stoma sometimes forming two lateral valves. Esophageal glands mononuclear, dorsal gland opening anterior to the third group of radial nuclei (Chitwood, 1937). The superfamily Camallanoidea was established by Travassos in 1920, with the following families: Camallanidae Railliet and Henry, 1915, Cucullanidae Cobbold, 1864, and Dacnitoididae Travassos, 1920. 11 16 т? Baylis (1923a) did not consider the superfamily Camallanoidea and the family Dacnitoididae sufficiently substantiated. He placed the families Camallanidae and Cucullanidae in the superfamily Spiruroidea, and did not consider them to be more closely related to each other than to other families of the superfamily. Yorke and Maplestone (1926) used the same arrange- ment. Superfamilies are not mentioned in the review of Baylis and Daubney which also appeared in 1926. The order Filarioidea is stated to include the families Filariidae, Philometridae (=Dracunculidae), Spiruridae, Camal- lanidae, Cucullanidae, and Gnathostomatidae. Travassos! work was thus not corrected, and later authors did not consider the families Camallanidae and Cucullanidae as closely related. Tornquist (1931) published a monograph on these two families, in which he gave a detailed analysis of their history of study and systematics and carried out important morphological and histological studies. Tornquist's work constitutes the culmination of the increasing tendency to dissociate the families Camallanidae and Cucullanidae. Tornquist gives the following points of similarity between the families: the same number of cephalic (oral, in his terminology) papillae, the structure of the nervous system, polymyarian musculature, the number of phagocytic organs, the position of the vulva, the composition of hosts, and the type of localization. However, he also gives a number of characters which differentiate these families: the difference in size between the sexes, the form of the cervical papillae, the number of cells of the excretory duct, the form of the posterior esophagus, the arrangement and structure of the esophageal glands, the length of the spicules, the mode of reproduction (the Camallanidae are viviparous, the Cucullanidae oviparous), and the presence (Camallanidae) or absence (Cucullanidae) of a caudal bursa. We have stressed the charac- ters which, according to Tornquist,, have the greatest systematic importance, As to the systematic position of the two families, Tornquist concluded that the Cucullanidae should be placed in the superfamily Strongyloidea Weinland, 1858. He suggests that the Camallanidae should be retained in the super- family Spiruroidea Railliet and Henry, 1915. Tornquist's proposals were not accepted. Chitwood and Wehr (1934) published a work in which the superfamily Camallanoidea is not mentioned, but he concluded that the Camallanidae and Cucullanidae may be connected through a more primitive form related to the Thelaziidae. In Chitwood's classification the superfamily Camallanoidea established by Travassos was re-instated and placed near the Dracunculoi- dea. The latter superfamily was first mentioned by Cameron (1934) and only the family Dracunculidae was originally included in it. The Dracun- culoidea resemble the Camallanoidea in that they have mononuclear eso- phageal glands and the larvae have large pocket-shaped phasmids. The species of both superfamilies parasitize vertebrates in the adult stage and crustaceans in the larval stages. In the "Кеу to Parasitic Nematodes, '' Volume IV (Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954) the family Cucullanidae Cobbold, 1864 was placed inthe superfamily Camallanoidea Travassos, 1920, and it was con- sidered, together with the superfamily Dracunculoidea Cameron, 1934, to belong to the suborder Callamanata Chitwood, 1936. The authors thought that a phylogenetic classification of the suborder Camallanata could not yet be constructed because of the absence of data on the development and morphology of many species of the group. They gave the following classifi- cation of the suborder: 12 Suborder Camallanata Chitwood, 1936 Superfamily Camallanoidea Travassos, 1920 Family Camallanidae Railliet and Henry, 1915 Family Cucullanidae Cobbold, 1864 Subfamily Cucullaninae Yorke and Maplestone, 1926 Tribe Cucullaninea Sobolev, 1952 Tribe Dichelininea Sobolev, 1952 Subfamily Seuratinae Hall, 1916 Superfamily Dracunculoidea Cameron, 1934 Family Dracunculidae Leiper, 1912 Subfamily Dracunculinae Stiles, 1907 Subfamily Avioserpensinae Wehr and Chitwood, 1934 Subfamily Philometrinae Yamaguti, 1935 Subfamily Micropleurinae Baylis and Daubney, 1922 Family Tetanonematidae Skrjabin and Schikhobalova, 1948 Family Anguillicolidae Yamaguti, 1935 Yamaguti (1961) gave a classification in which the families Camallanidae Railliet and Henry, 1915 and Cucullanidae Cobbold, 1864 were considered as belonging to the order Spiruridea Diesing, 1861. In his view the family Camallanidae contains five and the Cucullanidae seven genera, in two sub- families. For the other large group of nematodes which, according to the classifica- tion of Soviet authors (1954), are included in the suborder Camallanata and in the superfamily Dracunculoidea Cameron, 1934, Yamaguti established the new order Philometridea Yamaguti, 1961, in which he placed the families Philometridae, Anguillicolidae, Dracunculidae, Robertodollfusidae, and Crassicaudidae; for the family Tetanonematidae with one genus and one species he established the new order Tetanonematidea Yamaguti, 1961. The classification of this group, according to Yamaguti, is thus as follows: { Order Philometridea Yamaguti, 1961 Family Philometridae Baylis and Daubney, 1926 Subfamily Philometrinae Yamaguti, 1935 Genus Philometra Costa, 1845 Genus Clavinema Yamaguti, 1935 Genus Coregonema Bauer, 1946 Genus Ichthyofilaria Yamaguti, 1935 Genus Philometroides Yamaguti, 1935 Genus Philonema Kuitunen-Ekbaum, 1933 Family Anguillicolidae Yamaguti, 1935 Genus Anguillicola Yamaguti, 1935 Family Dracunculidae Leiper, 1912 Subfamily Dracunculinae Stiles, 1907 Genus Dracunculus (Linnaeus, 1758) Subfamily Avioserpentinae Chitwood, 1935 Genus Avioserpens Wehr and Chitwood, 1934 Subfamily Chelonidracunculinae Yamaguti, 1961 Genus Chelonidracunculus Yamaguti, 1961 Subfamily Micropleurinae Baylis and Daubney, 1922 18 Genus Micropleura Linstow, 1906 Genus Ophiodracunculus Yamaguti, 1961 13 19 Family Robertodollfusidae Chabaud and Campana, 1950 Genus Robertodollfusa Chabaud and Campana, 1950 Family Crassicaudidae Skrjabin and Andreewa, 1934 Genus Crassicauda Leiper and Atkinson, 1914 Genus Placentonema Gubanov, 1951 Order Tetanonematidea Yamaguti, 1961 Family Tetanonematidae Skrjabin and Schikhobalova, 1948 Genus Tetanonema Steiner, 1921 Ivashkin (1962) removed the family Cucullanidae Cobbold, 1864 from the superfamily Camallanoidea Travassos, 1920 and established for it the new superfamily Cucullanoidea Ivaschkin, 1962, including it in the suborder Camallanata Chitwood, 1936. Inglis (1967) placed the family Cucullanidae in the superfamily Seuratoidea Chabaud, Campana-Rouget, and Brygoo, 1959, suborder Ascaridata Skrjabin, 1915. Skrjabin and Ivashkin (1968) established the suborder Cucullanata Skrjabin and Ivaschkin, 1968, with the superfamilies Cucullanoidea Ivaschkin, 1962 and Gnathostomatoidea Skrjabin and Ivaschkin, 1968. Le-Van-Hoa and Fan-Ngok-Khue (1968), without mentioning the work of Ivashkin (1962) and Skrjabin and Ivashkin (1968a), established the super- family Cucullanoidea, instead of the superfamily Seuratoidea. Shigin and Shigina (1958) described a new nematode, 5. tincae,from the renal serosa of tench, which they placed in the new genus Skrjabil- lanus and new family Skrjabillanidae. The authors placed the new family in the superfamily Dracunculoidea together with the family Anguillicolidae, in which they found a number of similar characters: a well developed stoma, absence of spicules, esophagus divided into amuscular and a glandular part, vivipary. Sobolev (1952) placed the family Anguillicolidae in the superfamily Dracunculoidea, since it has to be considered as part of the suborder Camallanata in all characters. Chitwood (1937) placed in the subfamily Philometrinae the characteristic genus Phlyctainophora Steiner, 1921. Sobolev (1952) agreed with Chitwood and left it in this subfamily, but Roman (1960) established for this genus the new family Phlyctainophoridae Roman,1960. In our opinion, the families Anguillicolidae Yamaguti, 1935, Skrjabillanidae Schigin and Schigina, 1958, Phlyctainophoridae Roman, 1960, and Tetanonematidae Skrjabin and Schikhobalova, 1948 should be re- moved from the superfamily Dracunculoidea and placed in the superfamily Anguillicoloidea Sobolev, Ivaschkin, Tichomirova, and Khromova. Chitwood also thinks that the positionofthe genus Philonema Kuitunen- Ekbaum, 1933 in the classification of the suborder is not clear because there are mononuclear esophageal glands in its species. Sobolev (1952) placed this genus in the subfamily Philometrinae. Because of the life cycle of the type species of this genus, Philonema oncorhynchi, particularly the long period of maturation in the definitive host (about 3 years), as well as the morphological characters of the species of the genus, we consider it justified to establish for the genus Philonema the new subfamily Philoneminae. Skrjabin and Ivashkin (1968a) consider that the suborder Camallanata Chitwood, 1936 contains two superfamilies: Camallanoidea Travassos, 1920 with one family, Camallanidae Railliet and Henry, 1915, and the Dracun- culoidea Cameron, 1934 with the families Dracunculidae Leiper, 1912, Anguillicolidae Yamaguti, 1935, and Tetanonematidae Skrjabin and Schikho- balova, 1948. 14 20 The present authors consider the Camallanata to contain three super- families: Camallanoidea Travassos, 1920 with one family, Camallanidae Railliet and Henry, 1915, Anguillicoloidea Sobolev, Ivaschkin, Tichomirova, and Khromova n. superfam. with the families Anguillicolidae Yamaguti, 1935, Phlyctainophoridae Roman, 1960, Skrjabillanidae Schigin and Schigina, 1958, and Tetanonematidae Skrjabin and Schikhobalova, 1948, and Dracun- culoidea Cameron, 1934 with the families Dracunculidae Leiper, 1912 and Philometridae* Baylis and Daubney, 1926. PHYLOGENETIC RELATIONSHIPS OF THE CAMALLANATA AND THEIR REFLECTION IN THE CLASSIFICATION OF THE SUBORDER The superfamily Camallanoidea contains only one family, Camallanidae, in which the genera are clearly differentiated. Yeh (1960b) divided this family into two subfamilies, Camallaninae Yeh, 1960 and Procamallaninae Yeh, 1960, as follows: Camallaninae: Camallanus, Camallanides, ака сана Тана ь Ре Тата oer plume may Ziey lam emia: Procamallaninae: Procamallanus, Spirocamallanus. The subfamily Procamallaninae is a homogeneous group. Olsen (1952b) divided the species into two groups according to the structure of the buccal capsule; the species of the subfamily were placed in a single genus. The most primitive genus is Procamallanus. The subfamily Camallaninae is also a fairly homogeneous group; its most primitive genus is Para - camallanus. Of great interest are the data of Moorthy (1938b) on the development of Zeylanema sweeti and the data of Kupriyanova (1954), Campana-Rouget (1961b), and Moravec (1969) on the development of Camallanus lacustris. The stoma of third-stage larvae is of the Paracamallanus type,in C. lacustris it becomes transformed into а camallanoid type during the third molt, and ш Z. sweeti into a zey- lanemoid type during the fourth molt. These data suggest that Para - camallanus ** is the most ancient, primitive group in the Camallaninae. The structure of the buccal capsule of Camallanides is intermediate between the buccal organs of Procamallaninae and Camallaninae, but it still resembles the Camallaninae type because of the vertical ribs, lateral valves, and the almost round buccal capsule. All genera of Camallaninae, except Camallanides, have the distinct characters of the family. The genus Camallanides does not show marked differences from them, but its species are characterized by strongly developed lateral walls (valves) of the buccal capsule with marked reduction of the tridents, which are well developed in the other genera of Camallaninae (substitution of functions). * Rasheed (1963) studied the literature and extensive material, analyzed and redistributed the species of the genera of the family Philometridae and gave more specific diagnoses of the genera. Her classification of the family Philometridae has been accepted here. ** Further evidence of the primitiveness of Paracamallanus is that the posterior part of the buccal capsule of adult specimens shows a formation which is found only in the larvae of other genera of Camallaninae. 15 We consider the superfamily Anguillicoloidea, with four families, аз а transitional group which combines characters of such different superfamilies as the Camallanoidea and Dracunculoidea. It is difficult to construct a classification of the Anguillicoloidea, since a number of the families contain monotypic, incompletely known genera. In the superfamily Dracunculoidea Cameron, 1934 we retain the families Dracunculidae Leiper, 1912, with the subfamilies Dracunculinae Stiles, 1907, Avioserpentinae Wehr and Chitwood, 1934 emend. Chitwood, 1935, Micro- pleurinae Baylis and Daubney, 1922, and Philometridae Baylis and Daubney, 1926, with the subfamilies Philometrinae Yamaguti, 1935 and Philoneminae п. subfam. BIOLOGICAL CHARACTERISTICS OF THE SUBORDER CAMALLANATA CHITWOOD, 1936 The suborder Camallanata contains three superfamilies: Camallanoidea Travassos, 1920, Anguillicoloidea п. superfam., and Dracunculoidea Cameron, 1934. The biology of the species of these superfamilies will be described separately. Ontogenetic development in the superfamily Camallanoidea Published data on the biology of the Camallanoidea are few and concern only species of the genera Camallanus Railliet and Henry, 1915, Procamallanus Olsen,1952, and Zeylanema Yeh,1960. The development of C. lacustris (Zoega,1776) was studied by Mechnikov (Mecznikow, 1866), who found two larvae of different stages of development in Cyclops sp. Leuckart (1876) thought that an intermediate host, Cyclops, has a part in the development of С. lacustris,and he described larvae at various stages. Linstow (1909) found larvae which he determined as C. lacustris in Cyclops andinAsellus aquaticus, and Leiper (1910a) found them in Cyclops sp. The life cycle of this parasite was studied in detail by Kupriyanova (1954), Campana-Rouget (1961b), and Moravec (1969). Camallanus lacustris parasitizes in the intestine of freshwater fish. The females are viviparous, and the larvae are released into the water with the feces of the fish. Kupriyanova found that larvae discharged from a mature female into the water survive for 11—12 days at 9—10°С. The intermediate hosts of С. lacustris are Mesocyclops.leuckarti and Acantho= cyclops viridis. The first larval molt in the host was observed to take place after 24—48 hours at 19—21°. After the molt the old cuticle adheres closely to the new one and is therefore difficult to see. After molting, the larvae resemble larvae removed from females, except that the caudal end is more rounded, Their length is 0.445—0.446 mm and their maximum width 0.014—0.015 mm. The second molt takes place 5—6 days later. A distinct chitinous capsule appears at the cephalic end. The intestinal tract is divided into esophagus (muscular and glandular) and intestine. 16 21 The larvae are now 0.496—0.520 mm long and their maximum width is 0.017 mm. Kupriyanova found that the life cycle of С. lacustris in- volves an accessory host, possibly fish. The accessory host may be omitted from the cycle and is possibly a reservoir. Campana-Rouget did not mention Kupriyanova's studies but considers Leuckart's work as an accurate study of the life cycle, with respect to biological and morphological details. Campana-Rouget made another study of the development of this parasite in the intermediate host, with particular attention to the cephalic structures, which are important for the phylogeny of the group. According to her data, the intermediate hosts of С. lacustris are Acanthocy- clops viridis and Macrocyclops sp. Larvae emerging from the female attach themselves to the substrate with the caudal end and survive for 3 days at 20—24°. Campana-Rouget observed the first molt in the in- termediate host after 4—5 days. The second-stage larva has a buccal capsule, the nerve ring is situated 0.75 mm from the anterior end, and the esophagus becomes divided into a glandular and a muscular part. The larva is 0.700 mm long and 0.038 mm wide. The second molt takes place 8—10 days later. In the third-stage larva the buccal capsule is distinct and the eso- phagus is divided into two parts. The larva is now 0.880 mm long and 0.049 mm wide; the nerve ring is situated 0.095 mm from the anterior end and the anus 0.075 mm from the posterior end. The tail has the form ofa truncated cone with three distinct denticles. Four or five days after the second molt the larvae coil into a spiral, but they do not become encapsulated. The larvae do not develop further in Cyclops, but they remain alive as long as the host. As determined by Kupriyanova (1954), C.truncatus develops in the same intermediate hosts and has the same development as C. lacustris. Li (1935) established that the development of Spirocamallanus fulyvidraconis).(—Procamallanusyfulvidrac oni's: №, 1935) takes place in the body cavity of Cyclops. The first molt takes place 8—9 days after infection at room temperature; further development in the host was not observed. According to Pereira et al. (1936), Spirocamallanus cearensis (=Procamallanus cearensis Pereira, Vianna, Diaz, and Azevedo, 1936) develops in Diaptomus cearensis, and is localized in the body cavity. The authors did not observe the molt of the larvae in the inter- mediate host, but according to differences in the morphological characters, they think that the first molt takes place about 3 days after entry intothe host. After the molt the larvae are 0.66 —0.77 mm long and 0.03—0.04 mm wide. The buccal capsule has the same structure as in adult specimens. The esophagus is divided into a muscular (0.14 mm long) and a glandular (0.12 —0.13 mm long) part. The anus is situated 0.073 mm from the posterior end, The tail is short and ends in three or four denticles. These larvae are infective for the second intermediate host, a fish, in which the parasite completes the second molt but does not develop further than the third stage. The authors think that the definitive host possibly becomes infected directly when it eats the crustaceans, so that the life cycle of the parasite may con- tinue without a second intermediate host. They assume that only one molt takes place in the crustacean, but there are probably two molts, as the first molt is difficult to observe because it takes place soon after the larva enters the intermediate host, and the differences between the first- and 22 second-stage larvae are very small. 17 23 The development of Zeylanema sweeti (=Camallanus sweeti Moorthy, 1937), a parasite of the intestine of Ophicephalus gachua, was studied in detail by Moorthy (1938b). The intermediate host of Z. sweeti 15 Mesocyclops leuckarti. The first molt is observed after 24—36 hours, the second after 5—7 days. The definitive host may become infected by eating Mesocyclops, but Moorthy thinks that in nature infection of the definitive host does not occur without a second in- termediate host. It appears from the above that the first-stage larvae of all Camallanoidea grow in the external environment and mature slowly in the intermediate host. The first molt takes place soon after infection of the crustacean, 24—48 hours later in Camallanus lacustris and C.truncatus and immediately after entry into the body cavity of Mesocyclops in Zeylanema sweeti. Some authors (Li,1935; Pereira, et al. , 1936) probably overlooked the first molt and thought that the development of Camallanus in the crustacean ceases at the second larval stage. The second-stage larvae are infective, in their view; however, the description of these larvae corresponds to that of third-stage larvae given by Moorthy (1938b). The third-stage larvae are infective and pass from the intermediate host directly into the definitive host, or they enter a reservoir host (small fish) and then enter the definitive host together with it. Ontogenetic development in the superfamily Dracunculoidea The superfamily Dracunculoidea also includes viviparous forms, and since the vulva of adult females is usually obliterated, the larvae emerge through a rupture of the uterus and cuticle when the female emerges from the definitive host into the water and becomes subjected to a different osmotic pressure. Before entering the intermediate host, the larvae of Dracunculoidea sometimes spend some time in the water and remain infective. Fedchenko (1870), Fairley and Linston (1924), Isaev (1934a, b), and Craig and Faust (1937) established that Dracunculus medinensis develops in Cyclops for 12—15 days, but Manson (1893) and Leiper (1907) mentioned longer periods (to5—6 weeks), Isaev observed two molts of О. medinensis in Cyclops. Craig and Faust maintain that the larvae of this species de- velop in the body cavity of Cyclops with one molt, possibly two. Moorthy (1938a) gives a detailed description of the life cycle of D. medinensis. The larvae develop in the body cavity of Mesocyclops leuckarti and M.hyalinus. The first molt takes place after 5—7 days and the second 8—12 days after infection at 32—39° and after 8—10 and 13—36 days, respectively, at 12—21°. The second molt was found to take place before the cuticle of the first molt had been shed. Onabamiro (1954) recorded Thermocyclops niserianus as the intermediate host of О. medinensis. Desportes (1938) showed that Mesocyclops fuscus is the inter- mediate host of Dracunculus oesophagea; the first larval molt takes | place 10 days after infection. 18 24 Brackett (1938) described a new species from snakes, Dracunculus ophidensis, and established that Cyclops viridis is its intermediate host. The larva molts in Cyclops 12—15 days after infection and the second-stage larva becomes infective (the author apparently overlooked the first molt). If the high rate of infection of snakes and the relatively weak infection of Cyclops by D. ophidensis are compared, the author as- sumes that there should be an animal which "accumulates invasion." It was found by experiment that tadpoles may play the role of a carrier host, i.e. the parasitic larvae live in them for some time without further development. Chabaud and Campana (1949) described a new species of Dracunculidae from birds: Avioserpens galliardi. The intermediate host was found to be a Cyclops,in which molting was observed 12 hours after infection. The authors evidently observed the second molt of A. galliardi and overlooked the first molt. According to their data, the cyclops contained a second-stage embryo 15 days after infection, the posterior end of which showed denticles which are characteristic for third-stage larvae. An interesting study of the life cycle of Avioserpens mosgovoyi, a parasite of the subcutaneous tissue of Fulica atra and Podiceps Sets Tosh we. muta collins. gand и.о соед ema) ваз dbecnymadesby, Supryaga (1965а, b,1967a,1969a). Females of A. mosgovoyi, passing into the outer environment through the ulcerated skin of the bird, discharge larvae into the water, where they are swallowed by Cyclops. After a few minutes, the larvae penetrate into the body cavity, where they develop to the third stage. At a water temperature of 25—30°, the first molt takes place 55—10 days after infection of the Cyclops; the second molt takes place 2—3 days after the first or 7—12 days after infection. After the first molt the larvae did not shed the larval sheath, and both sheaths were shed to- gether after the second molt. At a water temperature of 20—25° the first molt takes place after 16—19 days and the second after 19—26 days. When the water temperature is 15° or below, the larvae do not develop further and do not molt, but they are still capable of further development. When the infected Cyclops were trans- ferred to an optimal temperature (24—26°), the larvae developed and became infective. In a case of superinvasion, when larvae of A. mosgovoyi of the first, second, or third stage were already present in the body cavity of the crustacean, the larvae of the first and the second infection became infective; however, it took much longer for the larvae of the second infection to de- velop. The same was observed when the Cyclops were infected witha large number of larvae of a particular stage (7—23 specimens). However, at such an intensity of infection, all the larvae became infective. Supryaga (1967a, b) established that reservoir hosts, vertebrates (fish, Amphibia) and invertebrates (larvae of Odonata), may take part in the life cycle of A. mosgovoyi. The larvae grow slightly in size in the reser- voir host but do not molt. In the definitive host, third-stage larvae of A. mosgovoyi pass from the intestine into the body cavity. They then pass along the serosa of the internal organs and the mesentery to the air sacs and reach the subcutaneous tissue after 8—12 days. The third molt takes place in the body cavity of the bird 4—5 days after infection; the fourth molt is completed in the subcutaneous tissue after 10 days in male larvae and after 13—14 days in female larvae. 19 25 Fertilization of females takes расе 15—16 days after infection and the vulva then begins to be obliterated. Fertilized females of A. mosgovoyi migrate into the submaxillary space of the bird, where a tumorlike formation appears after 18—20 days; ulcers are formed in this formation after 25—27 days. Females of A. mosgovoyi begin to discharge larvae 28—29 days after infection. This continues for 10—11 days, and then the females begin to die. The life span of females of A. mosgovoyi inthe definitive host is 39—44 days, that of males more than 9 months. The life cycle of Philometroides sanguinea (= Philometra sanguinea Rudolphi, 1819), which parasitizes on the fins and in the body cavity of crucian carp, was studied by Nybelin (1931) and Wierzbicki (1960). Nybelin established that the intermediate hosts of this species are Cyclops strenuus; Leptocyclops agilis, and Pachcyclops sp. Wierzbicki observed that in May—June, when the water temperature is sufficiently high, mature females of Ph. sanguinea fall from the fins into the water where they split open and release a large number of larvae. The free-living larvae grow to some extent in the water. For example, on the first day after they emerged from the female they were 0.37—0.40 mm long and 0.010—0.013 mm wide; 5 days later they were 0.41—0.42 mm long and 0.013—0.016 mm wide. In the intermediate host the larvae penetrate into the body cavity, where they become infective after 4—5 days. Although the larvae sometimes live for as long as two months in the crustaceans, they remain infective only 8—10 days. The infective larvae enter the intestine together with the Cyclops and migrate under the peritoneum into the region of the kidneys, swim bladder, and gonads, where they grow and develop to maturity. After mating, the fertilized females leave the body cavity and pass to the fins, where they remain for some time. In spring or early summer, females containing larvae enter the water. The life cycle of Ph. sanguinea thus lasts a year. However, Wierzbicki found in experiments that the life cycle may be shortened if the ambient temperature is higher. Vismanis (1964, 1966, 1967) studied the life cycle of Philometroides lusiana Visman, 1966, а parasite of carp, and found that the intermediate hosts of this species are Copepoda: Macrocyclops albidus, Eu- eyclops serrulatus; Mesocyclops leuckarti, Cyclops strenuus, Acanthocyclops viridis, and others. Larval development in the intermediate host lasts 6—7 days. When they enter the intestine, the infected crustaceans are digested, and the larvae of Philometroides released penetrate through the wall of the intestine into the body cavity of the fish, where they develop further. After fertiliza- tion of the females, the males of Ph. lusiana settle in the wall of the swim bladder, where they apparently live until the host dies, while the fe- males migrate to their permanent location, i.e. the scale pouches. They are about 4 mm long at this time. After they become attached under the scales, they grow rapidly. If the fish are infected in June, females of Ph. lusiana are 30 mm long at the beginning of August. During the winter, at low water temperatures (1—3°) the development of the nematodes ceases and begins again in spring, when the water temperature rises to 11°. At this time the females are 100—110 mm long and contain a large number of larvae which are 0.40—0.47 mm long. The larvae are released into the water at the end of May and in the first half of June at a water temperature of 15—20°. The life cycle of Ph. lusiana lasts about a year (from June to May—June the next year). 20 Kovaleva and Khromova (1967) made some observations of the biology of Philometra globiceps (Rudolphi, 1819), which parasitizes in the sex glands of Uranoscopus scaber. They found that the fish become in- fected after spawning in summer-autumn; further development of the para- sites and of the genital products of the fish apparently proceed at the same time. When the fish reach stages IV—V of gonad maturity, the females contain a large number of fully formed larvae. In the spawning period of the fish (June, July) the walls of the uterus and cuticle of females of Ph. globiceps rupture, and numerous larvae are discharged into the water together with the genital products of the fish. The life cycle of Ph. globiceps lasts about one year. Molnar (1966a, b, c) made some observations of the development of Philometra ovata (Zeder, 1803), a parasite of the body cavity of bream and roach,Thwaitia abdominalis (= Philometra abdominalis Nybelin, 1928), a parasite of the body cavity of minnow and gudgeon, and Ба аллее фа | (= Philometra, ciiseihta, Skrjabin, 1917),.a pas rasite of the opercula and subcutaneous tissue of the head of bleak. The author states that the fish become infected in June and July when they swal- low infected Cyclops. The females of Philometridae reach their maximum size only in May—June of the following year, and release a large number of larvae into the outer environment. The above data show that the species of the subfamily Philometrinae, like the genera of the subfamily Dracunculinae, have an annual life cycle. Ontogenetic development in the superfamily Anguillicoloidea The life cycle has been studied only in the species Agrachanus scardinii and Molnaria erythrophthalmi of the family Skrjabillanidae. Tikhomirova (published here the first time) showed that the development of these species takes place with an intermediate host which she determined as species of the genus Argulus: A foliaceus and A. coregoni. In contrast to other Camallanata, the larvae of these species are not released into the outer environment. From the uterus they enter the tissue surround- ing the parasites, and then they are carried by the bloodstream to the mus- cles and skin. Carp lice parasitizing on fish swallow the nematode larvae, which pass rapidly from the intestine into the body cavity and from there to the legs. The larvae grow and molt twice in the carp lice. The first molt 26 takes place after 4 days and the second after 8—9 days. The fish become infected when they feed on carp lice containing infective larvae. According to the life cycle of A. scardinii, the species of the Anguillicoloidea are, in their biological characteristics, more specialized than the species of other superfamilies of the suborder Camallanata. Twenty life cycles of species of Camallanata have been studied more or less completely. The species of the superfamilies Camallanoidea and Dracunculoidea studied do not show a strict specificity for their inter- mediate host or their localization. Their development to the third larval stage takes place in the body cavity of Cyclops and Diaptomus. 21 A free-living larval stage is characteristic for these species, but while growth of the larvae in the external environment is very distinct in the Camallanoidea, it is very minor in the Dracunculoidea (apparently in all species). The larvae of species of Anguillicoloidea are very delicate and die rapidly in the environment; from the definitive host they immediately enter the intermediate host, and from this again the definitive host. Reservoir parasitism in the suborder Camallanata has been little studied so far. We think that in the Camallanoidea, the most primitive of the Spirurata, reservoir parasitism of type II (after Ivashkin,1962) is present, i.e. the larvae grow in the reservoir host but do not undergo qualitative changes and do not molt. This type of reservoir parasitism is evidently present in Spirocamallanus cearensis, in which Pereira et al. (1936) observed the growth of larvae in small fish which are not the defini- tive hosts of this parasite. They think that the fish are a "superfluous in- termediate host'' in the life cycle of 5. cearensis, although such an intermediate host has a valuable part in "concentrating the parasites. " Reservoir parasitism was observed by Kupriyanova(1954)for Camallanus lacustris and: C.truneatus’. Brackett (1938) stated that tadpoles may be experimental "carrier hosts" of Dracunculus. The above data of Supryaga on reservoir parasitism in Avioserpens mosgovoyi deserve special attention. 22 21 SYSTEMATIC. PAR T: SUBORDER CAMALLANATA CHITWOOD, 1936 Diagnosis: Spirurida. Esophageal glands mononuclear, rarely with nuclear budding. Lips absent or rudimentary. Pseudolabia always absent. Stoma well developed or rudimentary. Esophagus divided into two parts: a muscular anterior and a glandular posterior part. Viviparous, de- velopment heteroecious. Intermediate hosts Crustacea. Larvae with large pocket-shaped phasmids. Parasites of vertebrates, mainly fish. Type superfamily: Camallanoidea Travassos, 1920. Key to the superfamilies of the suborder Camallanata 1 (2). Stoma well developed or reduced. Males without spicules........ О RG ae Cae a superfamily Anguillicoloidea n. superfam. 2 ). Stoma well developed or reduced. Males with spicules. SCE MSvOlMeWiell” GevViclOpeG а Е к Abo ew Sees ow И ey aii Ue A superfamily Camallanoidea Travassos, 1920. ПЗ Stomea: weakly developed Пиво оны, ое р emcee wea sits SUPERFAMILY CAMALLANOIDEA TRAVASSOS, 1920 Diagnosis. Camallanata. Eight cephalic papillae in the outer ring. Stoma well developed. Walls of stoma usually thick, sometimes forming chitinous ''valves.'' Mouth slitlike, hexagonal. Lips absent or rudimentary. Pseudolabia absent. Anterior part of esophagus muscular, posterior part glandular. Spicules present. Gubernaculum present or absent. Vulva situated in middle of body. Viviparous. Type family: Camallanidae Railliet and Henry, 1915. 23 28 FAMILY CAMALLANIDAE RAILLIET AND HENRY, 1915 Synonym: Cucullanidae Cobbold, 1884 Historical review The family Camallanidae was established by Railliet and Henry in1915 for the single genus Camallanus. Baylis and Daubney (1922 )described anew spe- cies of this genus, С. prashadi, whichdiffered from previously described species ofthe genus Camallanus inthatthe outer surface of the lateral walls of the buccal capsule shows thickenings of a reddishbrown color separated Буа longitudinal groove through the middle of the lateral wall. The authors consid- ered this character as soimportant that they established the new genus Camal- lanides Baylis and Daubney, 1922. Yorke and Maplestone (1926) stated that the buccalcapsule of C. cyathopharynx Baylis, 1923 is divided intoan anterior and a posterior part of almost equal size. As such a division of the buccal capsule is not present in other species of the genus, the authors established because of this character the new genus Paracamallanus Yorke and Maplestone, 1926, which they placed in the family Camallanidae together with the genera Camallanus, Camallanides, and Pro- camallanus. After a study of the buccal capsule in species of the genus Proca- mallanus (which at that time contained 24 species), Olsen (1952b) placed the species of this genus in two genera: Procamallanus and Spiro- camallanus. Olsen placed the species in which the inner surface of the buccal capsule was smooth in the first genus, and the species in which this surface showed spiral thickenings in the second genus. Ali (1956) described the new species C. singhi,and because of the absence of ''tridents or rods" established for it the genus Neocamalla- nus Ali, 1956. Yeh (1960b, c) used the structure of the buccal capsule and spicules and the host composition of the forms of Camallanus as a basis for establish- ing four genera: Camallanus (Railliet and Henry, 1915) Yeh, 1960, Piscilania Yeh,1960, Serpinema Yeh, 1960, and Zeylanema Yeh, 1960. He considered the genus Neocamallanus Ali,1956 asa synonym of the genus Camallanus. Since the genera Camallanus, Camallanides, Paracamallanus,and Piscilania have a buccal capsule consisting of two lateral valves, while in the species of the genera Procamallanus and Spirocamallanus the buccal capsule is entire and tridents are always absent, Yeh placed these groups of genera in two subfamilies, Camallaninae and Procamallaninae of the family Camallanidae. Chakravarty and Majumdar (1960) published a communication on the classification of nematodes of the family Camallanidae, in which they divided the family into two subfamilies: Camallaninae and Neocamallaninae. The 24 29 first subfamily contains the genera in which there is a pair of buccal valves and tridents are present or absent (Camallanus, Camallanides, and Paracamallanus) and the second the genera with an entire buccal capsule and without tridents (Procamallanus and Indocamallanus (=Neocamallanus)).* Thus, Yeh and Chakravarty et al. divided the family Camallanidae into two subfamilies on the basis of the same characters, but since Yeh's work was published first, the name Neocamallaninae Chakravarty and Majumdar, 1960 should be considered as a synonym of Procamallaninae Yeh, 1960. The family Camallanidae has the following composition: Family Camallanidae Railliet and Henry, 1915 Sy rae Camallaninae Yeh, 1960 ‘Genus Camallanus (Railliet and Henry,1915) Yeh, 1960 Genus Camallanides Baylis and Daubney, 1922 Genus Paracamallanus Yorke and Maplestone, 1926 Genus Piscilania Yeh, 1960 Genus Serpinema Yeh, 1960 Genus Zeylanema Yeh, 1960 Subfamily Procamallaninae Yeh, 1960 Genus Procamallanus Yeh, 1960 Genus Spirocamallanus Olsen, 1922 Diagnosis. Camallanoidea. Mouth dorsoventral, slitlike or hexa- gonal. Lips absent or rudimentary. Pseudolabia absent. Outer ring of cephalic papillae consisting of four large and four rudimentary papillae; inner ring formed by six very small papillae. Stoma strongly chitinized, either laterally compressed, forming cuticular lateral ''valves"', or barrel- shaped (Procamallanus). Esophagus divided into a muscular anterior and a glandular posterior part. The males have caudal wings and pedunculate papillae. Spicules different. Gubernaculum present or absent. Vulva situated in middle of body. Vivi- parous. Type subfamily: Camallaninae Yeh, 1960. Key to the subfamilies of the family Camallanidae He) )| Wuccall capsuledconsisting*er twalateralvvalvesy 2122 У. . PEARY ©). ВАР ХА NO TS A SRW A Camallaninae Yeh, 1960. Petey “Buccal capsule’ entirety а yb as le ie Procamallaninae Yeh, 1960. * The genus Neocamallanus was established by Chakravarty, Majumdar, and Sain together with the record of a new species, heteropneusti, the description of which was published in 1961. The authors probably did not know that the name Neocamallanus was preoccupied by Ali (1956), and therefore published a note in 1963 on the change of the name Neocamallanus to Indocamallanus, with the single species I. heteropneusti Chakravarty, Majumdar, and Sain, 1961. 25 30 SUBFAMILY CAMALLANINAE YEH, 1960 Diagnosis. Camallanidae. Buccal capsule with lateral valves, mouth slitlike, tridents present or absent. Type genus: Camallanus (Railliet and Henry, 1915). Key to the genera of the subfamily Camallaninae 1 (2). Buccal capsule divided into almost equal anterior and posterior PAIGE НИНЕ Paracamallanus Yorke and Maplestone, 1926. 2 (1). Buccal capsule without a posterior part or this markedly smaller thar the aitemerpart С китов ао sued). 2 eee (3). 3 (4). Buccal valves with deep lateral grooves, tridents replaced by rodlike FOPMAtLOnSs cis. ыы. = Camallanides Baylis and Daubney, 1922. 4 (3). Buccal valves without lateraliarooves 1102... що (5). 5 (8). Buccal capsule with longitudinal thickenings forming teeth....... ее а asus, Eo Ла. ORR Le Zeylanema Yeh, 1960. 6 (7). Longitudinal thickenings without armature .............. [7h 7 (6). Buccal capsule with short thickenings anteriorly and spines posteri- ОРТ ль ФМ! eel. Aon as са Piscilania Yeh, 1960. 8 (5). Buccal capsule with smooth, undivided thickenings ~ оо eee (9). 9 (10). Buccal thickenings divided laterally into a dorsal and a ventral Breupy). „лева Або el vente Serpinema Yeh, 1960. 10 , (9). Buccal thiekenings 5 A 3 AB “(QS6T ‘ПУ Joye) [elo1e] ‘этеш JO pus JoMojUe — р :[е1э3ет ‘ques — о Чемиэлозлор ‘auies — а ‘[eorde ‘pus o1jeydeo — ев “Ge AUN SIA :096Т ‘Чэл (9S6T ‘ПУ) TYSUTS зпиерешео = = = = Е AS) sa ffl 71 papillae extend into the caudal wings. There are also two pairs of circum- anal papillae near the cloaca. Five pairs of other papillae are preanal and 4 pairs postanal. The distance between the first and second preanal pairs and between the second and third pairs is about twice the distance separating the third, fourth, and fifth pairs. The first three pairs of postanal papillae are displaced, and the last pair is situated slightly separately, in the middle between the third pair and the end of the tail which is 0.10—0.14 mm long, Female. Length 6.35—6.48 mm, maximum width 0.14—0.19 mm. Dor- soventral width of cephalic end 0.073 mm, Excretory pore situated 0.20— 0.24 mm from anterior end. Buccal capsule (without basal ring) 0.073 mm long, 0.076 mm wide. The single ovary extends into the anterior third of the body and then turns posteriorly and reaches almost to the vulva. Posterior branch of uterus blind, situated in the anal region. Both branches of uterus 77 opening together into the muscular ovejector which is 0.43 mm long. Vulva 7 со with two distinct lips. Anterior lip larger and slightly overlapping the posterior lip. Vulva situated 2.9—3.34 mm from cephalic end. Uterus containing embryos 0.19 mm long and 0.011 mm wide. Tail 0.18—0.26 mm long, with a pair of phasmids 0.026 mm from the end. End of tail with 3 pa- pillae, two lateral and one terminal. References: Ali, 1956, pp.19—21; Yeh, 1960a; pp. 103—106. Camallanus thapari Gupta, 1959 (Figure 37) Host:; hana tigrina. Localization: small intestine. Distribution: Pakistan. Description (after Gupta, 1959). Male. Length 9.27—13.1 mm, width 0.25—0.29mm. Cuticle thin, with delicate striation. Mouth surrounded by 2 lateral and 4 submedian papillae. Dorsoventral width of cephalic end 0.1—0.13 mm. Buccal capsule formed by two valves 0.076—0.09 mm long (without posterior ring) and 0.10—0.11mm wide. Valves with 9—11 crests. Alarge chitinized ring 0.075—0.080 mm wide at the connection between valves and esophagus. Tridents well developed; median branch 0.040—0.048 mm long, lateral branches 0.050—0.060 mm. Club-shaped muscular anterior part of esophagus 0.46—0.50 mm long and 0.09—0.1 mm wide, cylindrical glandular posterior part 0.40—0.44 mm long and 0.09—0.10 mm wide. Nerve ring situated 0.14—0.17 mm, excretory pore 0.36 mm from cephalic end, Tail 0.14—0.15 mm long, with bifid tip. Caudal wings long, narrow, begin- ning from the cuticular swelling on the ventral side, maximum width about 0.03 mm. Six pairs of preanal,2 pairs of small, pedunculate adanal, and 5 pairs of postanal papillae. The two posterior pairs of the postanal papillae are pedunculate and isolated; the three anterior pairs are pedunculate and situated close together. The three posterior pairs of precloacal papillae are situated close together; the other precloacal papillae are isolated. Right spicule relatively thick, about 0.40—0.46 mm long, with a lateral process or tooth 0.04 mm from the end. Left spicule thinner, slightly widened anteriorly, 0.28—0.32 mm long. 72 (77) FIGURE 37. Camallanus thapari Gupta, 1959: a — anterior end, lateral; b — caudal end of male, ventral; c — same, lateral; d — caudal end of female, lateral (after Gupta, 1959). Female. Length 20.2—27.9 mm, width 0.45—0.60 mm. Cuticle thin, with delicate striation. Dorsoventral width of head 0.18—0.20mm. Mouth sur- rounded by 2 lateral and 4 submedian papillae. Buccal capsule with two valves 0.12—0.15 mm long (without posterior ring) and 0.15—0.16 mm wide. Valves with 11 ribs. A large, chitinized ring 0.09—0.1 mm wide at the con- nection between valves and esophagus. Tridents well developed: median branch 0.035—0.050 mm, lateral branches 0.05—0.06 mm long. Club-shaped muscular anterior part of esophagus 0.63—0.69 mm long and 0.12—0.15 mm wide, cylindrical glandular posterior part 0.59—0.61 mm long and 0.13—0.15 mm wide. Nerve ring situated 0.18—0.2 mm from cephalic end. Tail0.135—0.165 mm long, finger-shaped, with two small cuticular processes at the end. na specimen about 20.2 mm long the vulva is situated before the middle of the body, 8.2 mm from the cephalic end. Lip of vulva lobelike. tmemedie mee) Gupta, 1959, p. 774. Camallanus thaparus Sahay and Narayan, 1968 (Figure 38) Host: Channa (Ophiocephalus) striatus. Localization: intestine. вето ion’: india. Description (after Sahay and Narayan,1968). Thin, threadlike forms, dark red in life, without recognizable cuticular striation. Valves of buccal capsule wider than long in females, but this difference is small in males. Buccal capsule consisting of two valves with two outer thickenings and 9 longitudinal ribs. Cuticular ring situated at connection between valves and esophagus. Esophagus divided into a muscular and a glandular part, the glandular part shorter. Valves are present where the glandular part passes into the intestine. 73 Tridents distinct, with three branches of different length, their end bluntly pointed. Male. Length 7.00—7.44 mm, width 0.20—0.22 mm. Buccal valves 79 without the cuticular ring 0.095—0.115 mm long, 0.09—0.110 mm wide. 80 Cuticular ring 0.08—0.09mm wide. Length of muscular part of esophagus 0.51—0.52 mm, of glandular part 0.41—0.43 mm. Lateral branches of trident 0.06—0.07 mm long, median branch 0.025mm. Nerve ring situated 0.16—0.18 mm from cephalic end. FIGURE 38. Camallanus thaparus Sahay and Narayan, 1968: a — anterior end; b — cephalic end, lateral; c — trident; d — caudal end of female, lateral; e — caudal end of male, ventral (after Sahay and Narayan, 1968). Caudal end curved ventrally. Tail 0.170—0.183 mm long, conical, ending in two spines. Thirteen pairs of caudal papillae, 7 pairs precloacal, 2 adanal, and 4 pairs postcloacal. The first and second of the 4 postanal pairs are situated behind the cloaca, close together; the other two pairs are spaced at different intervals. Almost all 7 pairs of precloacal papillae are equally spaced except the fifth pair, which is situated 0.10—0.12 mm from the fourth pair. The caudal wings extend slightly beyond the proximal end of the large spicule. Left spicule weakly chitinized, 0.20 —0.225 mm long, the right spicule 0.425 mm long. Both spicules pointed. 74 81 Female. Length 16.6mm, maximum width 0.237 шт. Length of buccal valves without cuticular ring 0.13 mm, width 0.16 mm. Cuticular ring 0.115mm wide. Length of muscular part of esophagus 0.725 mm, of glandular part 0.525 пт. Lateral branches of trident 0.06 mm long, median branch 0.035 mm. Nerve ring situated 0.23 mm from cephalic end. Tail tapering, with rounded end with two thin, spinelike processes 0.15mm long. Vulva situated behind middle of body, 5.5mm from the caudal end, di- viding the body at the ratio of 3.2:1. Vagina long and muscular. Oviparous. Eggs present in posterior part of vagina. Reference: Sahay and Narayan, 1968, pp.118—124. Camallanus trichogasterae Pearse, 1933 (Figure 39) MOS urvchogasteratrichopterwts. Mocaliz ation: imtestine, Пьет вы Ето я: Asia. Description (after Pearse, 1933). Dark yellowish brown, opaque forms, with distinct transverse striation. Chitinized valves of buccal capsule with 14—20 ribs. FIGURE 39. Camallanus trichogasterae Pearse, 1933: a — anterior end; b — posterior end of male, lateral; с — posterior end of female, lateral (after Pearse, 1933). Male. Length3.55mm,. Тай, 0.07 mm long, blunt, curved ventrally. Five pairs of large postanal papillae and 2 pairs of small papillae at the end of the tail. Four preanal papillae. Length of right spicule 0.36 mm. Female. Length 7.2 mm. Depth of chitinized buccal capsule 0.09 mm. Length of buccal valves 0.06 mm, of chitinized pharynx 0.03 mm. Length of muscular esophagus 0.4mm, of glandular part 0.68mm. Tail 0.12 mm long, blunt, ending in a pair of small spines. Vulva situated at anterior third of body. 75 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.29; Pearse, 1933, рр. 184—185. Camallanus tridentatus (Drasche, 1884) (Figure 40) Host: Arapaima gigas. Localization: intestine. Distribution: Brazil (Amazon Basin). Description (after Travassos, Artigas, and Pereira, 1928). Male not known. FIGURE 40. Camallanus tridentatus (Drasche, 1884): ‚а — cephalic end, lateral; b — same, dorsoventral (after Travassos, Artigas, and Pereira, 1928). Female. Length7.7—10mm. Buccal capsule with two chitinized valves which are 0.17 mm wide and 0.12 mm long. Valves with 30 ribs. Tridents 0.1mm long, branches rodlike. Length of esophagus 0.65mm,; length of ventricle 0.75mm. Vulva situated 3.4 mm from posterior end. Tail 0.15 mm long. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.29; Travassos, Artigas, and Pereira, 1928, p.20. Camallanus truncatus (Rudolphi, 1814) (Figure 41) Synonyms: Cucullanus truncatus Rudolphi, 1814; Camalla- nus lacustris Yorke and Maplestone, 1926, пес Zoega, 1776 Hosts: definitive— Perca fluviatilis, Caspialosa вова niykovi,..C:oregonus lavaretus,oxyrhynchus, Osmerue eperlanus, .E.sox-luciugs, Silurus elanie, Коба Lucioperca lucioperca, Lucioperea volgensis, Acerina cernua, Rhombus maximus; intermediate — Mesocyclops letekarti,,.Acanthoeyclops,viridiss Localization: intestine. Distribution: Western Europe, USSR. Description (after Mészaros, 1967). Branches of tridents extending beyond the middle between buccal capsule and nerve ring. 76 82 Male. Length 2.7—4.8mm. Length of buccal capsule 0.078—0.083 mm. Median branch of tridents 0.097—0.10mm long. Muscular part of esophagus 0.48—0.52 mm long, glandular part 0.45—0.58 mm. Female. Length 3.80—10.60mm. Length of buccal capsule 0.077— 0.130mm. Median branch of tridents 0.077—0.13 mm long. Muscular part of esophagus 0.58—0.62 mm long, glandular part 0.62—0.82 mm. The development of Camallanus truncatus was studied by Kupriyanova (1954). The females areviviparous. Larvae discharged intothe : water with the feces of fish are colorless, 0.429—0.430 mm long, and 0.012— 0.013 mm wide. There is no chitinized buccal capsule and the intestinal tract is weakly developed and not differentiated. The tail is long and pointed. Larvae of С. truncatus survive 11—12 days in water at 9—10°. 02 mm 0.730. mm FIGURE 41. Camallanus truncatus (Rudolphi, 1814): a — cephalic end, ventral; b — same, lateral; с — larva from an adult female (а, b — after Mészdros, 1967; с — after Kupriyanova, 1954). At 18—20°, larvae swallowed by Cyclops enter the body cavity after 40 minutes. At 19—21°, larvae of С. truncatus undergo the first molt in Cyclops after 24—48 hours. The old cuticle is very thin and adheres closely to the new one. Second-stage larvae are 0.430—0.432 mm long and 0.013 mm wide. The caudal end becomes more rounded. The second molt takes place after 5—6 days. The larvae now have a distinctly chitinized buccal capsule, but tridents are still absent, and they are probably not developed at all in larvae in Cyclops, as none were observed even after 32 days inthehost. The esophagus is divided into a muscular and a glandular part. The caudal end becomes much shorter and the larvae turn yellowish. Length 0.478—0.489 mm, width 0.015 mm. If the temperature falls to 7—9°, development of the larvae is markedly retarded: the first molt takes place after 2—3 days and the second after 15 days. 83 Kupriyanova used perch as definitive host. During the first two weeks of development in the perch the larvae increase in body mass without dis- tinct morphological changes. After 10 days larvae of С. truncatus are 1.084 mm long and 0.039 mm wide; depth of buccal capsule 0.041 mm, width 0.026 mm; length of anterior part of esophagus 0.182 mm, of posterior part 0.176 mm, 77 84 After 20—25 days of development in the definitive host the tridents begin to develop and their development is completed 5—7 days later. The larvae are now 0.117 mm long and 0.010—0.011 mm wide. Kupriyanova assumed that further development of C. truncatus may take place either with an intermediate host or with an intermediate and a reservoir host. In the reservoir host (juveniles of Leuciscus idus, L,leuciseus, Abramis brama, "Регса fluviatilis) the larvae settle in the intestine, usually in the anterior part; however, they apparently do not develop there. References: Ivashkin and Khromova, 1964, pp.98—104; Kupriyanova, 1954, рр.373—377; Sobolev, 1962, pp.270—274; Mészaros, 1967, р. 159. Camallanus wolgensis Lewaschow, 1929 (Figure 42) Hostz eve cus semi rat s:: Localization: intestine. Distribution: USSR (Lower Volga). Description (after Lewaschoff, 1929). Buccal capsule with dark lateral valves, with 15—17 longitudinal ribs. After a short constriction, the posterior end of the valves forms a widening (thickening). The two thickenings form a chitinized ring. 0.05 mm wide before the esophagus. Tridents strongly chitinized, 0.065 mm long. Four well-developed sub- median papillae at the anterior end. Esophagus divided into a muscular anterior and a glandular posterior part. The muscular part of the esopha- gus was 0.52 mm long and the glandular 0.45 mm in one specimen. Three large nuclei present in the posterior part of the esophagus. Several large cells with distinct nuclei around the rectum. (83 ) FIGURE 42. Camallanus wolgensis Lewaschow, 1929: a — cephalic end, lateral; b — posterior end of male, lateral (after Lewaschoff, 1929). Male. Length 1.58 mm, width of cephalic end in the middle of the valves 0.075 mm, maximum width 0.077 mm, at the anus 0.047 mm. Tail 0.083 mm long. Caudal wings present; 6 pairs of preanal, 2 pairs of paranal, and 78 3 pairs of postanal papillae. Spicules 0.14 and 0.08 mm long. The spicules have a sheath. Gubernaculum absent. Female. Length of immature specimen 2.78 mm, width at the buccal capsule 0.127 mm, at the vulva 0.067 mm. Length of esophagus 0.97 mm. Tail 0.15 mm long, its end pointed and skittle-shaped. Vulva situated 1.77 mm from cephalic end. Tridents, which are present in adults, are absent in young specimens. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.29; Lewaschoff, 1929, pp.123—126. Camallanus yehi Fernando and Furtado, 1963 (Figure 43) Eton pC Dammam @lol oie pihialius ) эсставые. Moca 1 х азов sIntvestime: Distribution: Singapore. Description (after Fernando and Furtado, 1963a). Lateral valves of buccal capsule with 18 longitudinal ribs without armature. Buccal capsule sclerotized, pharynx also sclerotized. Glandular esophagus straight. There is a pair of small, rudimentary tridents in males; in females they are rep- resented by a small rod. FIGURE 43. Camallanus yehi Fernando and Furtado, 1963: a — cephalic end, lateral; b — caudal end of male, lateral; с — caudal end of female, lateral (after Fernando and Furtado, 1963а). Male. Length 5.76—6.10 mm, width 0.120 mm. Buccal capsule 0.056 mm long and 0.085 mm wide. Pharynx 0.022 mm long, maximum width 0.059 mm. Muscular part of esophagus 0.252 mm long, 0.096 mm wide, glandular part of 79 85 esophagus 0.720 mm 1оп$, 0.160 mm wide. Nerve ring surrounding muscular part of esophagus 0.096 mm from its beginning, 0.163 mm from the anterior end. Tail conical, curved ventrally, ending in a pointed process. Cloaca situated 0.096 mm from end of tail. Six pairs of preanal papillae (one pair . slightly before the cloaca) and 7 pairs of postanal papillae, one pair slightly behind the cloaca. Two equal, weakly chitinized spicules 0.110 mm long. Female. Length 7.2—10.5 mm, width 0.240 mm. Buccal capsule 0.056— 0.059 mm long, 0.074—0.082 mm wide. Sclerotized pharynx 0.026 mm long and 0.059 mm wide. Muscular part of esophagus 0.320 mm long, 0.096 mm wide, glandular part of esophagus 0.640—0.800 mm long, 0.122—0.128 mm wide. Nerve ring surrounding muscular part of esophagus 0.085—0.093 mm from its beginning, 0.166—0.185 mm fromthe cephalicend. Tail conical, with rounded end. Anus situated 0.240—0.315 mm from end of tail. Vulva situated slightly before middle of body, dividing it at the ratio of 1:1.1—1:1.5. Reference: Fernando and Furtado, 1963a, рр. 59—61. Camallanus zacconis Li, 1941 (Figure 44) Hos t204 ae cio, tiem mince ki, Localization: large intestine. Distribution: ‘China, Description (after Li, 1941). Cuticle smooth, Buccal capsule con- sisting of two valves with 17—18 longitudinal ribs. Cuticular ring situated at the connection between the buccal capsule andthe esophagus. Tridents with three branches of equal length. There are two small tetragonal chitinized plates at the anterior margin of the buccal valves. Cephalic papillae forming two rings. Inner ring consisting of 6 small papillae, the outer of 4 papillae and 2 amphids. Glandular part of esophagus slightly longer than muscular part; nerve ring situated at the anterior quarter of the muscular part of the esophagus. a FIGURE 44. Camallanus zacconis Li, 1941: a — anterior end, lateral; b) caudal end of male, lateral (after Li, 1941). 80 Male. Length 3.15—3.35 mm, width 0.16—0.18 mm. Buccal capsule 0.09—0.11 mm long, 0.1—0.11 mm wide. Cuticular ring 0.5—0.6 mm wide. Nerve ring situated 0.145—0.170 mm from cephalic end. Muscular part of esophagus 0.315—0.330 mm long, glandular part 0.35—0.37 mm. Caudal wings narrow. Right spicule markedly longer and thicker than the left, 0.12—0.145 mm long and 0.007 mm wide. Left spicule 0.08—0.093 mm 86long, 0.004 mm wide. Seven pairs of precloacal,2 pairs of paracloacal, and 6 pairs of postcloacal papillae. The postcloacal papillae are sometimes div- ided into three groups: an anterior group of three pairs, a middle group of two pairs, and the posterior pair is isolated. Tail short, 0.081—0.086 mm long, with blunt end. Female. Length 3.65—3.75 mm, width 0.14—0.16 mm. Buccal capsule 0.12 mm long, 0.12—0.13 mm wide. Cuticular ring 0.080—0.13 mm wide. Nerve ring situated 0.19—0.24 mm from cephalic end. Muscular part of esophagus 0.41—0.44 mm long, glandular part 0.45—0.47 mm. Vulva situated in posterior part of body, 1.52 mm from the caudal end, protruding above the sur- face, dividing the body at the ratio of 1:0.71 or 1.4:1. The uterus contained eggs at the morula stage. Tail 0.55 mm long, thin, with blunt end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.29; Li, 1941, pp.195—199. Camallanus sp. Chatterji, 1936 Host: Morenia ocellata. ШосаТтла 108: intestine. Distribution: Burma, Description (after Baylis, 1939). Male not known. Female. Length11.2 mm. Buccal valves wider than long, with10 longitu- dinalribs. Vulva situated before middle of body, with protruding anterior lip. Note. The description is made of one immature female. According to Yeh (1960b), reptiles are not typical hosts of the genus Camallanus. However, the genus of this form cannot be determined because of the in- adequate description. We place it therefore provisionally in the genus Camallanus, in which it was described. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.28; Baylis, 1939, p.214; Chatterji, 1936, р. 81. Camallanus sp. Kung, 1948 (Figure 45) Host: bull frog. Localization: intestine. Юве wit? om: South, Africa; Description (after Kung, 1948), Male not known, Female (young specimen). Length 2.72 mm, width 0.184 mm. Body narrowing posteriorly from the middle, cuticle distinctly transversely striated. Buccal capsule 0.121 mm deep, maximum width 0.172 mm. Lateral valves 81 87 88 with 29—32 longitudinal ribs. Mouth surrounded by 6 cephalic papillae, four submedian and two lateral. Tridents about 0.079 mm long, not reaching anterior part of esophagus. Nerve ring situated 0.20 mm, cervical papillae 0.32 mm from cephalic end. Length of esophagus 0.86 mm, length of mus- cular anterior part 0.39 mm. Vulva protruding above the surface, situated 1.69 mm from the cephalic end. Vagina about 0.248 mm long. Uterus in- completely filled with larvae. Tail 0.135 mm long, ending in three processes. AID, @ un и FIGURE 45. Camallanus sp. Kung, 1948: a,b — cephalic end, different aspects; с — caudal end of female, lateral, а — region of vulva, lateral (after Kung, 1948). Reference: Kung, 1948, рр. 103—106. Camallanus sp. Liu and Wu, 1941(Figure 46) Host: Macrone Longirostris. Localization: stomach. Di stribution¢e China, Description (after Liu and Wu, 1941). Male not known. Female. Length 4.88 mm, width 0.2 шт. Length of esophagus 1.1 mm, width 0.1mm. Head oblong, mouth dorsoventral, slitlike. Two lateral amphids and 4 pairs of submedian papillae, the two pairs near the corners of the mouth larger. Valves of buccal capsule with 15—17 longitudinal ribs. Dorsal and ventral to each amphid is an area of thickened cuticle, the anterior part of which passes into the valve; its posterior margin is dis- tinctly separated from the valve. The buccal capsule is not chitinized behind the valves. A cuticular collar around the connection between buccal capsule and esophagus. Length of buccal valves 0.131 mm, Intervals be- tween longitudinal ribs 0.007 mm. Intervals of striation of cuticle 0.0048 mm. Esophagus consisting of two parts, the glandular part longer. Trident well developed. Median branch 0.112 mm long. Vulva situated slightly before posterior third of body, 1.87 mm from posterior end. Tail 0.69 mm long, with rounded end. Larvae hatching in the uterus. 82 (87) FIGURE 46. Camallanus sp. Liu and Wu, 1941. Cephalic end, apical (after Liu and Wu, 1941). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.28; Liu and Wu, 1941, рр. 67—72. Camallanus sp. Moorthy, 1938 (Figure 47) ее Васоыз puckelii, Lepidocephalicht hy s (век = malis, Ophiocephalus gachua., Localization: body cavity, intestine (free and encysted). Destribution: India. 205mm FIGURE 47. Camallanus sp. Moorthy, 1938: a — encysted larva from intestine of О. gachua; b — caudal end of encysted larva; c — cephalic end of encysted larva (after Moorthy, 1938b). Description (after Moorthy, 1938b). Length of larva 1.1—1.6 mm, width 0.05—0.06 mm. Stoma 0.035—0.048 mm deep. Buccal capsule narrow, 83 89 90 half as long as in the third-stage larva; longitudinal ribs absent. Two pairs of papillae in the outer ring. Amphids present at anterior margin of body, between two pairs of chitinized tetragonal plates. Esophagus con- sisting of a muscular anterior part 0.19—0.22 mm long and a glandular posterior part 0.16—0.22 mm long. Nerve ring situated 0.09—0.11 mm from cephalic end. Excretory pore situated slightly behind the nerve ring. Ш- testine consisting of 100—120 cells arranged in transverse rows, 3—4 cells in each row. Anus situated 1.0—1.5 mm from anterior end. Tail 0.063— 0.081 mm long, conical, without processes. References: Baylis, 1939, p.214; Moorthy, 1938b, pp. 335—336. Camallanus sp. Moorthy, 1938 Host: “(Mes oeyelops lewekarti. Localization: body cavity. Distribution: India. Description (after Moorthy, 1938b), Larva 1.09 mm long and 0.07 mm wide. Stoma 0.046 mm deep, consisting of an almost completely chitinized buccal capsule and distinct buccal valves with 14 longitudinal ribs of dif- ferent length. They extend from the cephalic end to the chitinous ring behind the buccal capsule. Two pairs of papillae in the outer ring. Amphids situa- ted between two pairs of chitinized tetragonal plates. Cuticle deeply trans- versely striated with intervals of about0.004mm. Muscular anterior part of esophagus 0.26 mm long, glandular posterior part0.14mm; boundary between the two parts distinct. Nerve ring situated 0.13 mm from cephalic end. Excretory pore situated slightly behind the nerve ring. Nucleus of ex- cretory cell situated at the base of the muscular part of the esophagus. Intestine consisting of 65 cells arranged in transverse rows,3—4 cells in each row. Genital primordium situated 0.72 mm from anterior end, consisting of 4 longitudinally arranged cells. Anus situated 1.05 mm from anterior end. Phasmids rounded, situated slightly behind the anus. Tail 0.46 mm long. Three spines of equal length at end of tail. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.28; Baylis,1939, p.214; Moorthy, 1938b, p.323. Camallanus sp. Vuylsteke, 1964 (Figure 48) Host: fish. Localization: intestine. Distribution: Central Africa, Description (after Vuylsteke, 1964). Male not known. Female. Length 10 mm, maximum width 0.315 mm. Buccal capsule 0.092 mm long. Length of esophagus 0.700 mm. Genital opening situated 4.200 mm from cephalic end. Vagina directed posteriorly, uterus diverging. Larvae, twisted, were found free at the end of the body. Tail with three small pointed processes at the end. 84 TT TA р ТТУ АТАКИ! ms 0./mam. “ii A FIGURE 48. Camallanus sp. Vuylsteke, 1964: a — cephalic end; b — anterior end; c — posterior end of female (after Vuylsteke, 1964). Reference: Vuylsteke, 1964, p.58. Genus Camallanides Baylis and Daubney, 1922 Historical review The genus Camallanides was established for the species Camal- lanides prashadi, which was the only species of the genus for a long time. Khera (1954) described С. piscatori and С. ptyas, and estab- lished the two subgenera Camallanides and Procamallanides. The validity of the new subgenera because of the presence or absence of dorsal and ventral chitinoid bodies in the buccal capsule is doubtful. Campana-Rouget (1961a) and Deshmukh (1968) think that division of the genus Camallanides into subgenera is not justified. Gupta (1959) found one female of Camallanides in Pakistan, the de- scription of whichwe give as Camallanides sp. Yamaguti (1967b) placed "Camallanus bungari'" MacCallum, 1918 in this genus with a question mark, Diagnosis. Camallaninae. Buccal capsule with thickened lateral walls, with ribs on the inside. The lateral walls of the buccal capsule have two longitudinal massive reddish brown thickenings on the outside which are sepa- rated by a longitudinal groove in the middle. Outside the dorsal and ventral 91 part of the buccal capsule are rodlike formations which may be homologues of the tridents characteristic of the genus Camallanus. 85 There is а cuticular ring at the base of the buccal capsule. The esopha- gus consists of a muscular anterior part and a glandular posterior part. Cervical papillae small, setalike. Male. Caudal wings present, supported by 14 pairs of pedunculate papillae, seven pairs of them preanal. Right spicule longer and more mas- sive than the left; it has wings and a hooklike distal end. Gubernaculum present. Female. Vulva with a tubular, posteriorly directed appendage situated before middle of body. Posterior uterus without an ovary. Parasites of the intestinal tract of snakes and frogs. Type species: Camallanides prashadi Baylis and Daubney, ifs bapa Camallanides prashadi Baylis and Daubney, 1922 (Figure 49) Hosts: Naja hannah, Bungarus fasciatus, Ptyas (жалей тв”) mucosus) Rana tigrina ; Localization: intestine. Distribution: India. Description (after Baylis, 1939). Cuticular striation very fine and indistinct, with intervals of 0.003—0.004 mm. The buccal valves appear like four separate cuticular formations. Each valve has two wide outer thickenings witha longitudinal groove; 14 longitudinal ribs onthe inside which end anteriorly in toothlike appendages. Tridents absent, but there are yellow rodlike formations which are sometimes of irregular form, They are connected with the chitinous base on the dorsal and ventral sides at the margins of the valves. Male. Length 5.8—6.6 mm, width 0.21—0.25 mm. Dorsoventral width of cephalic end 0.08—0.09 mm. Length of buccal valves 0.06 mm, width 0.075 mm. Length of rodlike formations 0.06 mm. Posterior ring of buccal ‘apparatus 0.033 mm wide. Anterior part of esophagus 0.38 mm long, length of esophagus 0.8 mm. Nerve ring situated 0.15 mm, excretory pore0.25 mm, and the small, seta-shaped cervical papillae 0.28 mm from the cephalic end, Caudal wings well developed. Tail about 0.06 mm long, pointed and usu- ally curved ventrally, its end covered by the wings. Caudal papillae pedun- culate, decreasing in size posteriorly. The 7 pairs of preanal and the 2 pairs of adanal papillae are curved ventrally, while the 5 pairs of postanal papillae are situated laterally. The first three pairs are larger and situated close together. Right spicule massive, 0.24 mm long, with wings almost its entire length. Its end is hooked. Left spicule without wings, thin and pointed, about 0.14 mm long. Gubernaculum nearly triangular, about 0.025 mm long. 93 Female. Length 14.2—17.7 mm, width 0.40—0.47 mm. Dorsoventral width of cephalic end 0.12—0.13 mm. Length of buccal valves 0.09—0.1 mm, width 0.1 mm. Length of rodlike formations 0.07—0.1 mm. Posterior ring of buccal apparatus 0.045 mm wide. Anterior part of esophagus 0.47— 0.50 mm long, length of esophagus 1.0mm. Nerve ring situated 0.19 mm, ex- cretory pore 0.31 mm, and the small, seta-shaped cervical papillae 0.35 mm from the cephalic end. 86 (92) FIGURE 49. Camallanides prashadi Baylis and Daubney, 1922: a — anterior end of male, lateral; b — cephalic end, ventral; c — posterior end of male, lateral; d — region of vulva, lateral; e — caudal end of female, lateral; f,g — cephalic end, different aspects; Н — region of vulva, lateral; i — caudal end of male, lateral (a — e — after Agrawal, 1967; f —i— after Baylis and Daubney, 1922). The lips of the vulva form a tubular appendage which is sometimes dorsoventrally flattened and projects free posteriorly for 0.3—0.4mm. The appendage is situated at the anterior third of the body, 5.9—6.1 mm from the cephalic end. The vulva is situated on the ventral surface of the appendage, near its end. Vagina narrow, extending posteriorly for about 2 mm. Poste- rior branch of uterus without an ovary. Tail 0.4—0.6 mm long, tapering, slightly thickened before the conical end. 87 Description (after Аргаума1, 1967). Nematodes of medium size, slender, markedly truncate at both ends. Cuticle thin, with delicate striation. Mouth dorsoventral, surrounded by 6 papillae. Buccal capsule consisting of two valves, which consist of two chitinoid formations separated by a wide longitudinal groove. Valves with 10—12 complete and incomplete longitu- dinal ribs in both sexes. A large, chitinized ring at the connection between valves and esophagus. Tridents simple, pointed, yellow, situated opposite the corners of the buccal valves. Dorsal and ventral chitinoid formations 0.10 —0.15 mm long. Male. Length 6.74—7.29 mm, width 0.24—0.28 mm. Dorsoventral width of cephalic end 0.08—0.09 mm. Length of buccal capsule 0.06—0.065 mm, width 0.080—0.085 mm. Chitinoid ring 0.040—0.042 mm wide. Tridents 0.065—0.075 mm long. Length of club-shaped muscular anterior part of esophagus 0.29—0.30 mm, width 0.075—0.080 mm, length of cylindrical glandular posterior part 0.42 mm, width 0.09—0.095 mm, Length of esoph- agus 0.71—0.72 mm. Nerve ring surrounding anterior part of esophagus, situated 0.14 mm, excretory pore 0.19—0.22 mm from cephalic end. Tail conical, pointed, curved ventrally, forming a ring, 0.06—0.09 mm long. Caudal wings beginning at the cuticular thickening on the ventral side and extending to end of tail. Fourteen pairs of caudal papillae: 7 preanal, 2 paracloacal, and 5 postanal pairs. The papillae are pedunculate, riblike, and decrease in size posteriorly. Preanal papillae almost equally spaced. Paracloacal papillae curved in- ward and ventrally. Postanal papillae forming three groups. Right spicule 0.29—0.36 mm long, its end curved in а hook. Left spicule, slender, markedly 94 truncate, 0.175—0.190 mm long. Gubernaculum yellow, triangular, 0.028— 0.029 mm long. Female. Length 13.56—19.88 mm, width 0.40—0.48 mm. Dorsoventral width of cephalic end 0.11—0.14 пт. Length of buccal capsule 0.08—0.115 mm, width 0.11—0.14 mm. Chitinoid ring 0.055—0.070 mm wide. Length of muscular anterior part of esophagus 0.31—0.40 mm, width 0.10—0.15 mm, length of glandular posterior part 0.51— 0.63 mm, width 0.10—0.15 mm. ‘Length of esophagus 0.88—0.99 mm. Nerve ring situated 0.18—0.26 mm, excretory pore 0.32—0.56 mm, cervical papillae 0.52—0.86 mm from cephalic end. Tail 0.29—0.36 mm long, tapering, with a slight widening before the end. Vulva forming a tubular process which is dorsoventrally flattened and projects free, situated 6.1—8.6 mm from the cephalic end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.35; Agrawal, 1967, pp.338—341; Baylis, 1939, p.274; Baylis and Daubney, 1922, pp. 263—347. Camallanides bungari (MacCallum 1918) Yamaguti, 1961 (Figure 50) Host: Bungatrus candidus. Localization not given. Distribution: Java. Description (after MacCallum, 1918). Male not known. 88 Female. Mouth wide, subterminal. Opening of esophagus with two rings, the outer ring with small teeth. The rounded anterior margin of the esoph- agus is covered with small denticles. Esophagus bottle-shaped and mus- cular. Vulva situated at posterior third of body. Tail pointed, with three spines. FIGURE 50. Camallanides bungari (MacCallum, 1918): a — anterior end; b — posterior end of female, lateral (after MacCallum, 1918). References: MacCallum, 1918, рр. 123—124; Yamaguti, 1961а, рр.217—228. 95 Camallanides dhamini Deshmukh, 1968 (Figure 51) Host: Ptyas mucosus. Localization: intestine. Distribution: India. Description (after Deshmukh, 1968). Thin, red nematodes. Cuticle not striated. Head with 6 papillae, four submedian and two lateral. A group of esophageal glands at the base of the buccal capsule. Anterior part of esophagus club-shaped and muscular. Male. Length 5.5 mm, width 0.22 mm. Buccal capsule 0.086 X0.071 mm large. Capsule with 14 ribs. Length of posteriorly directed rods 0.064 mm. Buccal ring 0.043 mm wide. Nerve ring situated 0.15 mm from cephalic end. Length of anterior part of esophagus 0.29 mm, of posterior part 0.37 mm. 89 Caudal wings 1.10 mm long and 0.24 mm wide. Fifteen pairs of pedun- culate caudal papillae, 7 pairs precloacal, regularly distributed. The 5 pairs of postcloacal papillae are situated just behind the cloaca. Behind this group is a pair of double papillae, two papillae situated almost on the same base. Another pair of papillae situated in the middle between end of tail and the double pair of papillae. Spicules 0.31 and 0.26 mm long. Guber- naculum 0.017 mm long and 0.016 mm wide. Tail 0.13 mm long. 0.7 шт FIGURE 51. Camallanides dhamini Deshmukh, 1968: a — anterior end, lateral; b — region of vulva, lateral; c — caudal end of male, ventral; d — caudal end of female, lateral (after Deshmukh, 1968). Female. Length 11.3—14.76 mm, width 0.30—0.39 mm. Buccal capsule 0.105 X0.090 mm large, with 16 ribs. Length of posteriorly directed rods 0.086 mm. Buccal ring 0.048 mm wide. Nerve ring situated 0.15—0.17 mm from cephalic end. Length of anterior part of esophagus 0.28—0.31 mm, of posterior part 0.43—0.46 mm. The tubular vagina projects posteriorly in the form of an appendage 0.33 mm long. This appendage is situated 4.93—5.74 mm from the cephalic end. Larvae 0.49 mm long and 0.018 mm wide. Reference: Deshmukh, 1968, рр. 119—122. 96 Camallanides hemidenta Majumdar, 1965 (Figure 52) Host: Channa (Ophiocephalus) striatus. Localization: “intestine: Distribution: India. Description (after Majumdar, 1965). Cuticle finely transversely striated. Buccal capsule formed by two lateral valves, each valve with 11 90 97 longitudinal ribs. Tridents reducedtorods. Cephalic and cervical papillae absent. Esophagus divided into along, muscular, club-shaped anterior part and a shorter glandular posterior part. Male. Length 2.62—3.48 mm, width 0.13—0.16 mm. Length of buccal valves 0.03—0.04 mm, width 0.05 mm. Length of trident 0.013 mm. Nerve ring situated 0.10—0.11 mm from cephalic end. Length of muscular part of esophagus 0.24—0.28 mm, width 0.06 mm, length of glandular part 0.40— 0.53 mm, width 0.09—0.10 mm. Tail curved ventrally, 0.06—0.09 mm long. Caudal wings extending to end of tail. Six pairs of caudal papillae, five pairs postanal and one pair preanal. Left spicule 0.13—0.18 mm long, right spicule 0.11—0.13 mm long. Gubernaculum absent. FIGURE 52. Camallanides hemidenta Majumdar, 1965: a — anterior end; b — caudal end of male, ventral (after Majumdar, 1965). Female. Length 5.92 mm, width 0.22 mm. Length of buccal valves 0.05 mm, width 0.06 mm. Length of trident 0.013 mm. Nerve ring situated 0.22 mm from cephalic end. Length of muscular part of esophagus 0.29 mm, width 0.09 mm, length of glandular part 0.55 mm, width 0.11 mm. Tail long, 0.29 mm, conical. Vulva protruding above the surface, situated in anterior half of body, 3.45 mm from caudal end. Vagina directed anteriorly. Uterus opposite, filled with larvae. Larvae 0.104 mm long. imeme Cmeece Мариадаг, 1965, Бр: 222—225. Camallanides piscatori Khera, 1954 (Figure 53) In; ORES IN EWP ise ОИ Е, Localization: intestine. Drstribution; India. Description (after Khera, 1954). Nematodes of medium length. Cuticle finely striated with intervals of 0.0035—0.006 mm. Mouth dorso- ventral, surrounded by 6 papillae. Buccal capsule consisting of two valves, appearing like four separate reddish brown formations. Eachvalve consists of 91 two chitinoid formations separated by а wide longitudinal groove. Valves with 12 complete and incomplete longitudinal ribs. Tridents represented by simple, yellow, pointed chitinoid rods which are sometimes of irregular form; they are attached to the valves. Esophagus divided into a club-shaped mus- cular anterior part and a cylindrical glandular posterior part. Three small valves at the connection of the esophagus with the intestine. Intestine wider than the posterior part of the esophagus and with a wide lumen. Cells of in- testine with a distinct nucleus and granular protoplasm. Rectum small, narrow and lined with cuticle. Three oval rectal glandular cells with large nuclei at the connection between intestine and rectum. 2.7 mm FIGURE 53. Camallanides piscatori Khera, 1954: а — anterior end, ventral; b — same, lateral; с — caudal end of male, lateral; 4 — caudal end of female, lateral; e — genital system of female (after Khera, 1954). Male. Length 5—7 mm, width 0.18 mm. Dorsoventral width of cephalic end at the anterior corners0.77 mm. Lengthof buccal valves 0.075 mm, width 0.093 mm. Lengthoftridents 0.048mm. Posterior ring of buccal apparatus 0.09mm wide. Length of esophagus 0.71 mm (anterior part 0.31 mm, 98 posterior part 0.4mm). Nerve ring situated 0.115 mm, excretory pore 0.3 mm, and the small, seta-shaped cervical papillae 0.28 mm from the cephalic end. Caudal wings well developed. Tail conical, pointed, 0.12 mm long. Caudal papillae pedunculate and mushroom-shaped, decreasing in size to- ward end of tail. Fourteen pairs of caudal papillae, 5 pairs precloacal and 9 pairs postcloacal. The anterior pair of precloacal papillae is situated near the beginning of the caudal wings. The four posterior pairs of post- cloacal papillae are equally spaced. The postcloacal papillae form three groups of three papillae. All caudal papillae are ventral except the third pair from the posterior end, which is lateral, Right spicule thick and with a wing its entire length,0.32 mm long. Its proximal end is conical and ends in a pointed process, Left spicule without a wing, thin and pointed,0.22 mm long. Gubernaculum triangular, 0.025 mm long. 92 Female. Length 15—17.1 mm, width near the vulva 0.5—0.55 mm. Dorsoventral width of cephalic end at the anterior corners 0.114 mm. Length of buccal valves 0.1—0.11 mm, width 0.135—0.142 mm. Tridents 0.096 mm long. Posterior ring of buccal apparatus 0.04mm wide. Length of esophagus 0.93—0.99 mm (anterior part 0.36—0.39 mm, posterior part 0.58—0.62 mm). Rectum 0.08mm long. Nerve ring situated 0.245 mm, excretory pore 0.38 mm, and the small, seta-shaped cervical papillae 0.35 mm from the cephalic end. Tail 0.42—0.48 mm long, tapering, with a small thickening before the conical end. The lips of the vulva formatubular process which is dorso- ventrally flattened and projects posteriorly for 0.12—0.16mm. Ina spe- cimen 15.6 mm long this process is situated 6.5 mm from the cephalic end. The muscular vagina opens at the base of the process and then extends posteriorly for 0.9 mm along the ventral wall of the body. Uteri opposite. Posterior branch of uterus without an ovary. The single ovary is situated near the anterior end of the intestine. Viviparous. Uteri filled with larvae, waveh are 01 12—0.15 mm long. References: Campana-Rouget, 196la, р.28; Khera, 1954, pp. 55—58. Camallanides ptyas Khera, 1954 (Figure 54) Host: Ptyas mucosus. Localization: intestine. Distribution: India. Description (after Khera, 1954). Medium-sized nematodes. Cuticle with delicate striation. Mouth rounded, with 6 papillae: four submedian and two lateral. Buccal apparatus consisting of two reddish brown chitinoid 99 valves separated by a wide longitudinal groove. Valves with 10—12 complete and incomplete longitudinal ribs (12 in females, 10—12 in males). The ante- rior end of the ribs is markedly widened in females. Tridents or "rods" yellowish brown, simple, straight, chitinoid, rounded or blunt at both ends. They are attached to the corners of the valves, where there are no dorsal and ventral chitinoid bodies. Esophagus divided into a club-shaped mus- cular anterior part and a cylindrical glandular posterior part. Esophagus connected with the intestine by three small valves. Intestine wider than the posterior part of the esophagus, with a wide lumen. Cells of intestine with granular protoplasm and small nuclei. Rectum small, 0.1 mm long (in fe- male), very narrow. Three nuclear rectal glandular cells at the connection between intestine and rectum. Male. Length 8.5—8.8 mm, width 0.25—0.29 mm. Dorsoventral width of cephalic end 0.07—0.085 mm. Length of buccal valves 0.062—0.066 mm, maximum width 0.07—0.075 mm. Length of tridents 0.055—0.058 mm. Posterior ring of buccal capsule 0.028 mm wide. Length of esophagus 0.725—0.76 mm (anterior part 0.255—0.27 mm, posterior part 0.47—0.49 mm), Cuticle striated with intervals of 0.002—0.0035 mm. Nerve ring surrounding anterior part of esophagus 0.13—0.15 mm, excretory pore situated 0.3 mm, and the small cervical papillae 0.24 mm from the cephalic end. 93 FIGURE 54. Camallanides ptyas Khera, 1954: а — anterior end, ventral; b — same, lateral; с — caudal end of male, lateral; d — larvae; е — genitalia of female (after Khera, 1954). Caudal wings well developed. Tail conical, pointed, 0.048—0.050 mm long, curved ventrally. Caudal papillae large and pedunculate. Five pairs 100 of precloacal papillae; the first three pairs are lateral. The two posterior pairs of precloacal papillae and all postcloacal papillae are ventral. All papillae of the same size. Right spicule thick, with wings on a large part, 0.19—0.21 mm long and with hooked end. Left spicule weakly chitinized, threadlike, without wings, 0.09—0.1 mm long. Gubernaculum L-shaped, 0.028 mm long. Female. Length 23—25 mm, width 0.42—0.46mm. Width of cephalic ‘end at the anterior corners 0.09—0.1 mm. Length of buccal valves 0.09 mm, maximum width 0.11—0.125 mm. Length of tridents 0.075 mm. Posterior ring of buccal capsule 0.045 mm wide. Length of esophagus 1.06—1.16 mm (anterior part 0.36—0.395 mm, posterior part 0.7—0.765 mm). Nerve ring surrounding anterior part of esophagus 0.16—0.17 mm, excretory pore sit- uated 0.42 mm, and the small cervical papillae 0.4 mm from the cephalic end. Cuticle striated with intervals of 0.003—0.005 mm. Tail 0.46—0.5 mm long, tapering, with a small thickening before the end. Lips of vulva forming a tubular process which is dorsoventrally flat- tened and extends posteriorly. Length of process 0.3 mm. In a specimen 25 mm long the process is situated 11mm from the cephalic end. Vulva situated on the ventral surface of the process, near its end. Vagina nar- row, extending to the base of the process and then posteriorly for 1.4mm. Uteri opposite. A single ovary, situated near the anterior end of the in- testine. Posterior branch of uterus without an ovary. Viviparous. Uteri filled with small larvae with pointed tail. Larvae 0.10—0.12 mm long. Reference: Khera, 1954, рр. 110—114. 94 Camallanides sp. Gupta, 1959(Figure 55) Hostel ydrophis, cy anoci ne tu sy, Localization: small intestine. Distribution: Pakistan. Description (after Gupta, 1959). Male not known. Female (one specimen). Length 10.4 mm, width0.4 mm, Cuticle thin, with delicate striation. Mouth dorsoventral, with 3 papillae on each side. Buccal capsule with two valves, which consist of two chitinized masses зер- arated by a wide longitudinalgroove. Length of buccal capsule 0.1 mm, width 0.11 mm. Valves with 12 complete and incomplete longitudinal ribs. Tridents represented by simple, pointed, chitinized rods of irregular form, with a dorsal and a ventral chitinoid body situated opposite the anterior margin of the buccal valves. Length of rod 0.065—0.09 mm. Posterior ring of buccal capsule 0.045 mm wide. Dorsoventral width of cephalic end at the anterior corners 0.13 mm. 101 Length of esophagus 0.92 mm, width 0.12 mm; esophagus divided into a club-shaped anterior part 0.42 mm long and 0.06 mm wide and a cylindrical glandular posterior part 0.5 mm long and 0.12 mm wide. Nerve ring situated 0.19 mm, excretory pore 0.32 mm from cephalic end. FIGURE 55. Camallanides sp. Gupta, 1959: a — cephalic end of female, lateral; b — caudal end of female, lateral; c — region of vulva (after Gupta, 1959). Tail conical, 0.33 mm long. Lips of vulva forming a tubular appendage which is dorsoventrally flattened and projects posteriorly for 0.04 mm, This appendage is situated slightly before middle of body, 4.6 mm from the cephalic end. Vulva situated on the ventral side of the appendage, near its end. The vagina extends anteriorly to the opening of the appendage. Reference: Gupta, 1959,'pp. 778—779. Genus Paracamallanus Yorke and Maplestone, 1926 Synonym: Camallanus Railliet and Henry, 1915 (in part) Diagnosis. Camallaninae, characterized by a markedly widened 95 102 103 connection between buccal capsule and esophagus. This part of the body is not surrounded by a cuticular ring and is almost funnel-shaped. Buccal capsule markedly laterally compressed, especially anteriorly. The slitlike mouth extends far posteriorly on the dorsal and ventral side. The capsule is much wider than long laterally. There are a few longitudinal ribs on the inside of the buccal capsule. The ribs of the right and left side are situated opposite each other and are short, extending only for about half the length of the capsule. Tridents present. Lips of vulva weakly developed. Type species: Paracamallanus cyathopharynx (Baylis, 1923) Yorke and Maplestone, 1926. Paracamallanus cyathopharynx (Baylis, 1923) Yorke and Maplestone, 1926 (Figure 56) Synonym: Camallanus cyathopharynx Baylis, 1923 Hest si MmeterobranichivsVanbuillaris; ‘Cla'rtras lazer a. Localization: intestine and pyloric caeca. Distr атоме ey pt. Historical review Paracamallanus cyathopharynx was described by Baylis (1923a) from Heterobranchus anguillaris inEgypt. The main characters the species 15 а large, chitinized "buccal cavity'' or 'pharynx'' behind and in ad- dition tothe paired buccalvalves. This structure separates the valves from the esophagus. Baylis was not sure that spicules were present and he found only 5 preanal papillae. Inthe same hostandalsoin Clarias lazera, Campana-Rouget (1961а) found specimens which were identical with the species of Baylis. Campana- Rouget transferred Camallanus sweeti Moorthy, 1937 and Neo- camallanus singhi Ali, 1956 tothe genus Paracamallanus. She probably did not know Yeh's study (Yeh, 1960b), in which the genus Zey - -lanema Yeh, 1960 was established because of differences in the struc- ture of the buccal capsule in species of Camallanus. Camallanus sweeti Moorthy, 1937 is asynonym of Zeylanema sweeti (Moorthy, 1937) Yeh, 1960, and Neocamallanus singhi is a synonym of Ca- mallanus.)sinpihi,(Aliy. 1956), Хей; «1960. Description (after Baylis, 1923а). Striation of cuticle with intervals of 0.006 mm. Length of chitinous buccal apparatus, including valves and pharynx 0.1—0.13 mm. Buccal valves with 10—12 longitudinal ribs of dif- ferent length; tridents absent. The small cervical papillae are situated just behind the nerve ring and the excretory pore slightly before the pos- terior end of the muscular part of the esophagus. Male. Length of largest specimen 5.9 mm, maximum width 0.12 mm. Length of esophagus 0.93—1.1 mm, muscular part 0.44—0.56 mm long. Tail 0.05—0.07 mm long. Five pairs of preanal papillae, situated at the sides, widely spaced. Spicules not found; the males were probably not mature. Campana-Rouget (1961а) gives the following information on the male. Right spicule chitinized, 0.210—0.230 mm long, ending in a small pointed process, without a wing and with a wide sheath. Left spicule more strongly chitinized, very small, 0.025—0.030 mm long, resembling а gubernaculum. The vesicular caudal wings are united before the cloaca and connected 96 (102) 00f/™mm а 21: Ух ви FIGURE 56. Paracamallanus cyathopharynx (Baylis, 1923): a,b,c — cephalic end, different aspects; d — cephalic end of female, dorsal; e — same, lateral; f — posterior end of male (a — after Yorke and Maplestone, 1926; b, с — after Tornquist, 1931; d, e — after Baylis, 1923; f — after Campana- Rouget, 1961). ventrally. Papillae difficult to distinguish, 5 pairs of preanal and 3 pairs of subventral postanal papillae were found. There may be 2 further pairs of lateral papillae, one in the middle of the tail and the other nearer its end. Except for the first postanal pair, all papillae are very small. Female. Length of largest specimen 9.2 mm, width 0.18 mm. Length of esophagus 1.28—1.32 mm, muscular part 0.65—0.67 mm long. Tail 0.35—0.40 mm long. Vulva not protruding, situated slightly behind middle of body (4.3 mm from the posterior end in a specimen 9.2 mm long). The vagina extends posteriorly for 0.8 mm, Viviparous. References: Baylis, 1923a, pp.1—38; Campana-Rouget, 1961а, pp. 25—27; Tornquist, 1931, p. 234; Yeh, 1960b, pp.117—124; Yorke and Maplestone, 1926, рр. 378—379. Genus Piscilania Yeh, 1960 Synonym: Camallanus Railliet and Henry, 1915 Diagnosis. Camallaninae. Buccal capsule with inner longitudinal ridges which are short anteriorly and spinelike posteriorly. Parasites of marine fishes. Type species: Piscilania melanocephalus (Rudolphi, 1819) Yeh, 1960. 97 104 Piscilania melanocephalus (Rudolphi, 1819) Yeh, 1960 (Figure 57) Synonyms: Cucullanus melanocephalus Rudolphi, 1819; Camallanus melanocephalus Rudolphi, 1819 Hosts: Pelam ys sp-,-Auxis Se... Stan brus s'arda, 5. rochei; Si сое. Товарное Localization: intestine. Distribution: Mediterranean, North Sea. Description (after Tornquist, 1931). Cuticle transversely striated. Ribs on valves of buccal capsule not extending their whole length, only about half the length of the capsule. Cephalic end slightly narrowing at the con- nection of the two parts of the esophagus. FIGURE 57. Piscilania melanocephalus (Rudolphi, 1819) Yeh, 1960: a,b — cephalic end, different aspects (after Tornquist, 1931). Male. Length 12.59 mm, width 0.296 mm. Length of buccal capsule 0.109 mm. Chitinized ring at the base of the capsule 0.039 mm long, 0.078 mm wide. Branches of tridents extending from the large cuticular plate, immediately behind the mouth. Lateral branches of tridents markedly longer and thicker than median branch, extending 0.218 mm from the ce- phalic end. Lateral branches 0.078 mm long, median branch 0.0312 mm. Anterior part of esophagus 1.482 mm long, posterior part 0.936 mm. Tail with rounded end, 0.109 mm long. Spicules indistinct, right spicule 0.687 mm long, left spicule 0.608 mm. Number and arrangement of papillae not determined. Reference: Tornguist, 1931, р, 341. Genus Serpinema Yeh, 1960 Synonym: Camallanus Railliet and Henry, 1915 (in part) Diagnosis. СатаПаптае. Buccal ridges smooth, divided into dorsal and ventral groups. 98 105 Parasites of reptiles. Type species: Serpinema intermedius (Hsu and Hoeppli, 1931) Yeh, 1960. Serpinema intermedius (Hsu and Hoeppli, 1931) Yeh, 1960 (Figure 58) Synonym: Camallanus intermedius Hsu and Hoeppli, 1931 rior t<)) (Geo clemysm mee vie Sit. Localization: small intestine and rectum. Distribution: ‘China’, Description (after Hsti and Hoeppli, 1931). Cuticle finely transversely striated. Buccal valves slightly wider than long, with 10 longitudinal ribs and with subventral and subdorsal papillae. At the anterior part of the valves are formations which are difficult to see, probably amphids. Tridents well developed. Winglike formations around the mouth in apical view, two on each lip. Esophagus divided into a muscular anterior part and a glandular posterior part. Nerve ring situated near base of buccal capsule. Cervical papillae very small. Е] a К) ( \“ Ue Ks : rad < A 4 9 С] х FIGURE 58. Serpinema intermedius (Hsti and Hoeppli, 1931): а — cephalic end, lateral; b — same, ventral; с — same, apical; а — caudal end of male, ventral; е — same, lateral (after Нзй and Hoeppli, 1981). Male. Length 5.65—7.90 mm, width 0.20—0.23 mm. Dorsoventral width of cephalic end 0.14—0.16 mm, lateral width 0.13—0.15 mm. Dorsoventral 99 106 width of cuticular ring 0.035—0.090 mm. Median branch of trident 0.060— 0.085 mm long. Nerve ring situated 0.16—0.165 mm, excretory pore 0.26— 0.28 mm, cervical papillae 0.31—0.41 mm from cephalic end. Muscular part of esophagus 0.45—0.51 mm long, glandular part 0.49—0.55 mm. Caudal wings well developed; 7 pairs of riblike preanal papillae, 2 pairs of small adanal papillae, and 5 pairs of postanal papillae, which form two groups on each side: one group of three and the other of two pairs of papillae. Near the end of the tail are two openings of caudal glands. Tail with bifid end, 0.55—0.56 mm long. Right spicule thick, 0.49—0.63 mm long, with bifid end. The two ends of the spicule are of different length, the shorter about 2 /3 as long as the longer. Left spicule thin, 0.20—0.26 mm long, ending in a fine process. The preanal papillae, except the pair at the anus, form the line from which muscles resembling a sucker radiate to both sides. Female. Length 11.05—12.1 mm, width 0.275—0.345 mm. Dorsoventral width of cephalic end 0.165—0.175 mm, lateral width 0.11—0.13 mm. Dor- soventral width of cuticular ring 0.082—0.14 mm. Median branch of trident 0.10—0.12 mm long. Nerve ring situated 0.17—0.215 mm, excretory pore 0.27—0.315 mm, cervical papillae 0.41—0.535 mm from cephalic end. Mus- cular part of esophagus 0.52—0.56 mm long, glandular part 0.60—0.69 mm. Tail trifid, 0.17—0.21 mm long. Vulva situated slightly behind middle of body, 0.525—0.600 mm from end of tail. Lips of vulva indistinct. Vagina long, with strongly muscular walls. Two opposite uteri, posterior branch ending blind, without an ovary. Viviparous. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.26; Hsu and Hoeppli, 1931, pp.567—588; Yeh, 1960, pp: 107—116. Serpinema amazonicus (Ribeiro, 1941) Yeh, 1960 (Figure 59) Synonyms) Саша Павия ama zionmcus, Ribeiro, 1941 Host: Pod ocenemis, -oxpam sya. Lo@adli zation: intestine. Dist#eibutLonk,«Brazil, Description (after Ribeiro, 1941). Cuticle transversely striated with intervals of 0.008 mm. Cephalic end truncate in both sexes. Tail ta- pering in the female. Buccal capsule with two valves, two tridents and separated ventrally or dorsally from the body by cuticular grooves. Buccal valves lateral, curved inward. Each valve with 17—20 longitudinal ribs on the inner surface. They have four thickenings on the outer surface, two anterior transverse and two median in the form of a crescent. The tridents, are situated at the connection of the valves. The tridents consist of a main body 0.087 mm long and 0.069 mm wide, inthe form of adisk, withthree di- verging branches of about the same length. The outer margin of the median branch shows distinct serration, and there is a curve along the posterior margin of the cuticular groove at the distal end. A chitinized ring at the connection between valves and esophagus. Cephalic papillae difficult to dis- tinguish, two pairs present, one laterodorsal, the other lateroventral. 100 Esophagus divided into an anterior club-shaped part and a posterior tubular part, with a fold in the anterior third; at the posterior end of the esophagus are three valves which project into the intestine. 107 ЕДА у ТИ FIGURE 59. Serpinema amazonicus (Ribeiro, 1944): a,b — cephalic end, lateral; c — cephalic end, dorsal; d — trident; e — anterior end, lateral; {— same, dorsal; g — distal end of large spicule; В — caudal end of male, lateral; i — end of tail of female; j — caudal end of female, lateral; К — region of vulva (after Ribeiro, 1941). 101 108 109 Cervical papillae and excretory pore not recognizable. Intestine narrower than esophagus. Male. Length 11.95 mm, width 0.40 mm. Buccal valves 0.17 mm long, 0.18 mm wide. Median branch of trident 0.069 mm long. Chitinized ring 0.043 mm long and 0.104 mm wide. Length of esophagus 1.14 mm, anterior part 0.61 mm long, posterior part 0.53 mm. Nerve ring situated 0.40 mm from cephalic end. Tail curved ventrally, with transverse musculature. Caudal wings well developed, fused anteriorly, extending to end of tail. Caudal papillae arranged as follows: 7 pairs preanal, long and pedunculate, one pair pedunculate, postanal, and one pair of postanal sessile papillae near end of tail. Spicules threadlike, of different length. Larger spicule 0.57 mm long, with wider base and bifid end. Smaller spicule more weakly chitinized, about 0.24 mm long. Gubernaculum absent. End of tail conical, not bifid. Anus situated 0.13 mm from posterior end. Female. Length 15.08—17.92 mm, width 0.53—0.68 mm. Buccal valves 0.204—0.208 mm long and 0.226—0.239 mm wide. Median branch of trident 0.078— 0.087 mm long. Chitinized ring 0.056 mm long and 0.139 mm wide. Length of esophagus 1.38—1.40 mm, anterior part 0.73—0.75 mm long, posterior part 0.63—0.65 mm long. Nerve ring situated 0.426 mm from cephalic end. Uteri didelphic, prodelphic. Vulva with lips protruding to different length in different specimens, situated 8.64—9.17 mm from anterior end. Ovejector divided into a widened, strongly muscular anterior part 0.078—0.096 mm long and 0.096—0.10 mm wide and a narrow posterior part 0.304—0.391 mm long and 0.052 mm wide. [If the anterior lip of the vulva protrudes, the vagina is directed obliquely anteriorly; if the posterior lip protrudes, the vagina is directed obliquely posteriorly. Uteri divergent, posterior branch without an ovary, ending in a blind sac about 0.938 mm from the posterior end. Anterior uterus passing into the ovary at about the connection between esophagus and intestine. Uterus filled with larvae, which are comma-shaped, 0.252 mm long and 0.008 mm wide, with conical end. Tail ending in three processes, one dorsal and two lateroventral. Rectum 0.18 mm long. Anus with indistinct lips, situated 0.21 mm from posterior end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.25; Diaz-Ungria, Gallardo, and Manuel, 1968, рр. 550—570; Ribeiro, 1941, pp.723—727; Yeh, 1960b, рр. 117—124. Serpinema kachugae (Baylis and Daubney, 1922) Yeh, 1960 (Figure 60) Synonym: Camallanus kachugae Baylis and Daubney, 1922 Host: Kachiusda iit. Localization: intestine. Distribution: Japan. Description (after Baylis, 1939). Striation of cuticle with intervals of 0.005 mm. Dorsoventral width of cephalic end at its anterior corners 0.13— 0.15 mm in males and 0.17 mm in females. Buccal valves slightly wider than long, 0.11—0.13 mm long without the posterior ring and 0.14—0.16 mm wide. Valves with 8 or 10 longitudinal ribs, ten ribs present only in large 102 110 specimens. Width of posterior ring of buccal apparatus 0.1 mm. Median branch of tridents 0.08—0.1 mm long. Length of esophagus, measured from the cephalic end, 1.18—1.55 mm, anterior part,0.54—0.66 mm. Nerve ring situated 0.20—0.23 mm and the small seta-shaped cervical papillae 0.50— 0.55 mm from the anterior end. Excretory pore situated at the level of the papillae. Intestine markedly narrower than esophagus. < = < > NTS ААА FIGURE 60. Serpinema kachugae (Baylis and Daubney, 1922): a — anterior end of female, lateral; b — caudal end of male, lateral; с — region of vulva, lateral (after Baylis and Daubney, 1922). Male. Length 10.9—14.5 mm, width 0.3—0.37 mm. Body slightly thicker in the region of the caudal wings than further anteriorly. The ventral region between the wings can apparently be depressed by the caudal muscles, form- ing a suckerlike organ. Seven pairs of pedunculate preanal papillae, 2 pairs of adanal papillae, and 6 pairs of postanal papillae. The first two pairs of postanal papillae are separated; the third, fourth, and fifth pairs form a group. All papillae are lateral, but the sixth pair is situated more ventrally, slightly behind the cloaca. Right spicule relatively thick, about 0.97 mm long. Its distal end is apparently simple and finely pointed. Left spicule very thin, 0.43 mm long. Female. Length 20.8—22 mm, width 0.45—0.5 mm. Tail finger-shaped, blunt and with slightly bifid end. Tail about 0.3 mm long; a pair of papillae 0.17 mm from the end, situated in small depressions of the cuticle. Vulva situated before middle of body, 9.1—10.4 mm from anterior end. It has strongly protruding lips, anterior lip wider than posterior lip and overlapping it. Each lip consists of a cuticular swelling with granular content and is divided inside into several parts by cuticular septa. Vagina narrow, extend- ing posteriorly for about 2 mm, gradually widening, and passing into the short common duct of the uterus, which extends posteriorly. 103 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.26; Baylis, 1939, pp.207—210; Baylis and Daubney, 1922, pp. 322—325; Yeh,” 1960b, ‘pp. 107-116, Serpinema magathi (Sprehn, 1932) Yeh, 1960 (Figure 61) Synonyms: Camallanus magathi Sprehn, 1932; С. parvus Caballero, 1939 Hosts: Clinostersnums cor pidcides, GAIplLegrumy Cs hiranpe'se Localization: intestine. Distributions Bolivaa, Mexico, Description (after Sprehn, 1932). Cuticle finely transversely striated with intervals of 0.002 mm. Median rib of buccal capsule with four ends on each side. Cervical papillae situated immediately behind the nerve ring, 0.65 mm from the cephalic end. Excretory pore situated 0.767 mm from the cephalic end. Male. Length 7.65 mm, width 0.38 mm. Length of valves 0.12 mm, width 0.138 mm. Length of muscular part of esophagus 0.39 mm, width 0.13 mm, length of glandular part 0.585 mm, width 0.156 mm. Three valves about 0.065 mm long at the connection between esophagus and intestine. Tail with blunt end, 0.113 mm long. Spicule 0.65 mm long, width at base of terminal process 0.010 mm. Female. Length 8.8—10.5mm, width 0.5 mm. Length of valves 0.14mm, width 0.165 mm. Muscular part of esophagus 0.487 mm long, 0.156 mm wide, glandular part 0.78 mm long, 0.218 mm wide. Vulva situated 5.1—5.3 mm from cephalic end, with two lips. Vagina extending posteriorly. Tail with blunt end. Tail 0.212 mm long, indistinctly bifurcate. Size of eggs 0.03 X0.027 mm. Description (after Caballero, 1939a). Cuticle transversely striated. Maximum width at anterior part of the esophagus. Buccal capsule with two valves, with 11 straight ribs which do not extend to the chitinized ring and form an angle. A short rib is also present in the middle of the buccal capsule, at the level of the median papilla. Male. Length 3.783 mm, width 0.292 mm, Length of buccal capsule 0.107 mm, width 0.115 mm. Width of chitinized ring at base of buccal cap- sule 0.086 mm. Length of lateral branches of trident 0.086 mm, of median 111branch 0.061 mm; branches flat and wide. Anterior part of esophagus 0.328 mm long, 0.066 mm wide anteriorly and 0.11 mm posteriorly. Nerve ring situated 0.172 mm, cervical papillae 0.273 mm from cephalic end, Anus situated 0.092 mm from end of tail. Tail with two wings. Seven pairs of preanal lateral papillae with a long stalk and 4 pairs of postanal papillae, three of them behind the cloaca and a smaller one at the end. Caudal wings 0.273 mm long. Larger spicule 0.585 mm long and 0.016 mm wide, with a hairlike process 0.041 mm long at the distal end. Female. Length 7.15—7.90 mm, width 0.37—0.39 mm. Length of buccal capsule 0.115— 0.135 mm, width 0.131 mm. Chitinized ring 0.094—0.098 mm wide, Lateral branches of trident 0.090 mm long, median branch0.074 mm. Length of anterior part of esophagus 0.370 mm, width0.082 mm at its anterior end 104 and 0.131—0.152 mm at its posterior end. Glandular part of esophagus 0.585 mm long and 0.131—0.135 mm wide. Nerve ring situated 0.173 - 0.213 mm from cephalic end. Cervical papillae and excretory pore not found. FIGURE 61. Serpinema magathi (Sprehn, 1932): а — cephalic end, lateral; b — caudal end of male, lateral; с — caudal end of female, lateral (after Sprehn, 1932). Vulva with strongly developed lips, situated in posterior half of body, 3.35—3.80 mm from end of tail. The two uteri are opposite. Anus situated 0.253 mm from end of tail which is finger-shaped and ends in three papillae. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.27; Caballero, 1939а, pp.448—450; Sprehn, 1932, Dowco 2e4, Е. 119505. эр. Об. Serpinema microcephalus (Dujardin, 1845) Yeh, 1960 (Figures 62—64) оу ше uC ama alan nis) mune o ce para liuis (Юшасаш, 1845); ©. beuspinosus (Meidy, 1861); С. commusts Ва Це? and Henry, 1915; Cre mel digae Nace luna, mone ie ome .c epi alus МаесСа ао, Poe п еезаля МаеСа юз, 1908: © 7 at Vorcid iamare дас Саи, IONS] ©. зеабкае 'Мас@а она, 11978: ©. telstuidin1 ssWae@alluma, ВОО Capa eran Cran usual aca tion 99: ис е мибальт ааа аи, Oi: Сен lan советов о ров 118 9: Geo аси ос ле эа, lus ПЕН. whoo Сп Cuneta Векещес. 1871 tle, LEO dees Нм с, tiem la. Mes Om pike Ml ae Sue (ое muy So Cea = Piper BC. le our оса Cul кавер Guan iC 36 re OO t 12 С а ету ака, Берет вая: Ска ем уса ca bien el осла Chie mer Cuma aie Ся S Cie роа Nia lao = eran Gay Ss) езде Ue А волос ве Lys) Odor a tus ah Де 1 Om vex Вт иены ue ANU Cui, Due i матем у с, аа. 105 Localization: intestine. Europe, Algeria, North America. Distrib ооо Historical review Dujardin(1845) describedanematodefrom Emys orbicularis as Cucullanus microcephalus. Perrier (1871) gaveadescription of Cu- cullanus dumerillii after a specimen from a tortoise in the Zoological Garden in Paris. Linstow described ш 1897 Cucullanus dumerillii Perrier, 1871 from Emys sp.? in the Zoological Garden in Hamburg. Railliet and Henry (1915) thought that the specimen described by Linstow is different from that of Perrier, and they renamed it Cucullanus con- fusus. Seurat (1915) described a species of the genus Camallanus from Clemmys leprosa as Cucullanus microcephalus Duj. Magath (1919a) stated that Dujardin's description of a specimen from Clemmys seurati, although he could not determine was inaccurate, and named it C. the species according to Dujardin's description. This inconsistency was pointed out by Baylis and Daubney in 1922. ) Ne if НИ FIGURE 62. Serpinema microcephalus (Dujardin, 1845): a — posterior end of male, ventral; b — anterior end, ventral; c — region of vulva, lateral; d — terminal part of uterus, oviduct, and ovary (diagrammatic) (after Seurat, 1915). Tornquist(1931)described Camallanus microcephalus from material of the Zoological Museum in Berlin from Emys sp. and from Clemmys leprosa (in Tornquist's view, this is not leprosa, since there were two labels on the jar, on one of them a question mark after the word leprosa). Tornquist noted that the specimens from Emys and Thus,Camallanus confusus from Clemmys were identical. Railliet and Henry, 1915 and Camallanus seurati Magath,1919 are the same species. 106 Linstow determined the specimens in the Berlin museum from Emys sp. as Cucullanus dumerillii Perrier. There is no information as to whether these specimens were used for the description of this species in 1131897, although Tornquist thought that this is possible. The host is the same 114 in the description and on the label, but its locality is not mentioned. Tornquist did not find it justified to separate the species described by Linstow in 1897 from that described by Perrier in 1871. He also thought that it was not justified for Magath to consider the forms of Camallanus from Emys orbicularis andfrom Clemmys leprosa as different species merely according to Dujardin's description (1845). Tornquist also thought that all North American species of the genus Camallanus from Chelonia are synonyms of C. microcephalus, and considered only С. scabrae MacCallum, 1918 as a valid species. Caballero (1943) gave a detailed description and drawings of this species, which we give below under Serpinema microcephalus, since we agree with Chitwood (1933b), who considered С. scabrae and the other North American species of the genus as synonyms of С. microcephalus. Yeh (1960b) divided the genus Camallanus into several genera and transferred the species Camallanus microcephalus tothe genus Serpinema. Description (after Tornquist, 1931). Cuticle transversely striated. Six oral papillae, Each valve of buccal capsule with 10 ribs which form two groups, an upper and a lower group; the ribs are curved and extend obliquely posteriorly. Short ribs are absent behind the mouth and behind the chitinized ring. A strong thickening extends in the midline, almost through the whole buccal capsule. Branches of tridents of equal length, the outer branches wider than the median branch, which is more pointed. Behind the buccal capsule begins the anterior part of the esophagus, which is here half as wide as in its posterior part. The cephalic end becomes narrower at the anterior part of the posterior part of the esophagus; this narrowing is small in the male but more dis- tinct in the female. Male. Length 6.93—10.84 mm, width 0.265 mm. Thickness of cuticle at the nerve ring and at the cloaca 0.0025 mm, in the middle of the body 0.0028 mm. Depth of buccal capsule 0.109 mm. Anterior end of esophagus situated 0.468 mm from the cephalic end. Length of posterior part of esophagus 0.780 mm. Opening of dorsal esophageal gland situated 0.040 mm behind the buccal capsule, openings of subventral glands situated 0.035 mm from beginning of posterior. part of esophagus. Nerve ring situated 0.203 mm from cephalic end. Excretory pore situated 0.190 mm behind the nerve ring; length of efferent duct 0.125 mm. Excretory organ U-shaped. Cer- vical papilla on the right side situated 0.135 mm behind the nerve ring, left papilla 0.160 mm behind it. The papillae are slightly raised above the surface. The boundaries between the cells are clearly visible in the anterior part of the intestine, but the nuclei are not recognizable. Rectum 0.130 mm long, cloaca 0.085 mm. The testis ends 0.750 mm behind the connection of the esophagus with the intestine. Anterior part of testis coiled, the anterior coil situated 0.460 mm from the connection between esophagus and intestine; posterior part of testis straight. The narrow transitional part leading to the vas deferens is about 0.050 mm long. Vas deferens 1.180 mm long; it becomes narrower proximally, and the duct becomes markedly wide 0.750 mm from its beginning (seminal vesicle). Transitional part leading to ejaculatory duct 0.060 mm long. Length of ejaculatory duct 1.270 mm. 107 | 8 FIGURE 63. Serpinema microcephalus (Dujardin, 1845): a—d— anterior end, different aspects; e, f — caudal end of male, lateral; р — caudal end of female, lateral; h — region of vulva, lateral (after Tornquist, 1931). Length of tail 0.062 mm. Seven pairs of precloacal and 5 pairs of postcloacal papillae. Precloacal papillae situated subventrally. The three anterior pairs of postcloacal papillae are also subventral, situated close together, and larger than the other two pairs. Dis- tance between second and third pair of postcloacal papillae (counting from end of tail) smaller than that between the sccond and first pairs, which are situated near the end of the tail. The postcloacal papillae at the end of the tail are lateral. Spicules of different length, right spicule 0.800 mm long, 115 left spicule 0.380 mm. Larger spicule with a small wing on the left side. Both spicules with rounded cross section. Female. Length 11.684—12.730 mm, width 0.390 mm. Thickness of cuticle at the nerve ring 0.0028 mm, at the vulva 0.0042 mm, and at the anus 0.0025 mm. Depth of buccal capsule 0.125 mm. Anterior end of esophagus situated 0.562 mm from cephalic end. Posterior part of esopha- gus 0.687 mm long. 108 Vulva situated 5.647 mm from the cephalic end, with strong lips, the anterior lip markedly thicker than the posterior and directed obliquely posteriorly. The duct from the vulva is curved broadly anteriorly (about 0.030 mm). Only this U-shaped part of the duct is covered with cuticle. The posterior branch of the uterus is curved anteriorly for 0.230 mm. This curve is situated 0.250 mm from the rectum. The anterior branch extends to the point where the two branches unite and then turns posteriorly. Oviduct 1.430 mm long, without loops. Seminal vesicle beginning 0.430 mm from the uterus; it is 0.500 mm long. Ovary extending posteriorly for 1.270 mm and then turning anteriorly. The part extending anteriorly is about 0.750 mm long, and there is another part extending posteriorly for about 0.400 mm. Length of ovary about 2.420 mm. Posterior part of body tapering to end of tail. Tail 0.172 mm long, pointed. Description (after Caballero, 1943). Coloration reddish. Females much larger than males. Cuticle transparent, with transverse striation which is more distinct than the longitudinal striation, with intervals of 0.012—0.024 mm. Anterior end rounded, wider than the posterior end, with 3 pairs of papillae: a mediolateral, a ventrolateral, and a dorsolateral pair. Valves of buccal capsule lateral, convex outside and concave inside, yellowish red, with chitinized walls, each with 15 chitinized ribs which are wide an- teriorly and pointed where they converge near the esophagus. Median dorsal rib short, the lateral ribs longer, extending to the cuticular ring at the base of the valves. Ribs attached anteriorly to two chitinized lateral plates, which curve at a right or acute angle to the dorsal and ventral surface of the cap- sule. Posterior part of buccal capsule attached to the cuticular ring. At the level of the ring are tridents dorsally and ventrally, their branches with sinuous or serrate margin; their ends often widen inthe form ofa hook, and their middle bears a disk which has a notch anteriorly. Esophagus divided into a club-shaped muscular anterior part which begins at the ring; the posterior part is cylindrical and glandular. Valves are present at the connection of the muscular and glandular esophagus and of the glandular part with the intestine. Intestine cylindrical with thick walls. Male. Length 9—9.217 mm, width 0.236—0.273 mm. Length of buccal valves 0.112—0.116 mm, width 0.136—0.140 mm; thickness of valves at the base 0.015—0.020 mm. Length of ribs 0.065 mm, width 0.004—0.005 mm, length of smaller ribs 0.008 mm. Length of cuticular ring 0.013—0.023 mm, width 0.086 mm. Length of tridents 0.100 mm, space between lateral branches 0.144 mm. Length of disk 0.044 mm, width 0.044—0.056 mm. Median branch of trident 0.078 mm long, 0.010 mm wide; right branch 0.096 mm long, 0.020 mm wide, left branch 0.092 mm long, 0.024 mm wide. Muscular part of esophagus 0.436 mm long, 0.127—0.136 mm wide; glandular part 0.582— 0.618 mm long, 0.100—0.145 mm wide. Intestine 0.084—0.088 mm wide. Anus situated 0.152—0.196 mm from end of tail. Nerve ring situated 0.160—0.184 mm, excretory pore 0.512 mm, cervical papillae 0.352—0.392 mm from the cephalic end. Tail finger-shaped, with two small lateral papillae at the end. Caudal wings finely transversely striated; the lateral muscles in this region converge radially to the opening of the intestine and the spicules. The caudal wings are supported by 10 pairs of pedunculate lateral papillae: 7 pairs preanal and 3 pairs postanal; the two pairs near the cloaca are situated close together. Cloaca with 2 pairs of sessile ventral papillae. 109 Spicules of different length, narrow and weakly chitinized, ending in a pointed 117 process. Larger spicule 0.772—0.880 mm long 0.016—0.018 mm wide at the manubrium; smaller spicule 0.360—0.368 mm long, 0.008 mm wide at the manubrium. Gubernaculum absent. 0240mm FIGURE 64. Serpinema microcephalus (Dujardin, 1845): a — anterior end, lateral; b — caudal end of male, lateral; c — caudal end of female, lateral; d,e — cephalic end in different positions; f — wident; g — caudal end of male, lateral (а — с — after MacCallum, 1918; d — g — after Caballero, 1943). Female. Length 13.853—14.598 mm, width 0.345—0.400 mm. Length of buccal capsule 0.120—0.132 mm, width 0.168—0.176 mm, thickness of valves 0.024 mm. Large ribs 0.091—0.099 mm long, 0.005—0.008 mm wide; small ribs 0.015—0.023 mm long. Cuticular ring at base of buccal capsule 0.021—0.026 mm long, 0.112—0.114 mm wide. Tridents 0.095—0.104 mm long, space between lateral branches 0.164—0.176 mm. Median branch of trident 0.056—0.060 mm long, 0.016—0.032 mm wide; right branch 0.096— 0.104 mm long, 0.020—0.028 mm wide, left branch 0.104—0.108 mm long, 0.028 mm wide. Middle part of trident 0.044 mm long, 0.056 mm wide. Muscular anterior part of esophagus 0.436—0.491 mm long, 0.136—0.145 mm wide, glandular part 0.563—0.673 mm long, 0.100—0.116 mm wide. Intestine straight, with 4—5 large rectal cells, which are 0.100—0.160 mm long. Anus situated 0.260—0.291 mm from end of tail. Nerve ring situated 0.252—0.296 mm, cervical papillae 0.404 mm from cephalic end, Tail tapering and ending in a process with 3 tubercles. Genitalia amphi- delphic, occupying greater part of posterior half of body. Vulva situated 110 behind middle of body, 7.199—7.544 mm from end of tail. Lips of vulva large, protruding above the surface, 0.309—0.318 mm long and 0.109—0.145 mm wide. Vagina large, cylindrical, muscular, extending anteriorly for 0.460— 0.500 mm, 0.044—0.052 mm wide. Twouterine branches. Viviparous. Length of larvae in uterus 0.301 mm, width 0.012 mm. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.27; Baylis and Daubney, 1922, pp.263—347; Caballero, 1943, pp.195—200; Chitwood, 1933b, p.242; Dujardin, 1845, p. 654; Linstow, 1897, pp.608—632; MacCallum, 1918, pp.123—124; Magath, а Do. 40 — OU: Perrier, Lol) pp. 937—339: Rallliet and Нейку, Г9ЭТ5, pp. 146—252; Seurat, 1915, рр. 423—426; Tornguist; 1931, pp. 1— 441; ЕН L260. рЬ. 10—16 Serpinema octorugatus (Baylis, 1933) Yeh, 1960 (Figure 65) Synonym: Camallanus octorugatus Baylis, 1933 Host: Heosemys grandis. Localization: stomach, small intestine. Distribution: Malaysia. Description (after Baylis, 1933). Cuticle transversely striated with intervals of 0.005—0.006 mm. Valves of buccal capsule, including posterior ring, about as long as wide. Anterior margin of valves with two inner chitinoid thickenings. Eight longitudinal ribs onthe inner surface of each valve. Tridents with three branches of almost equal length. Median branch narrow and thin laterally; its end is not completely chitinized. Lateral branches thick, wider at the end. Anterior margin of base of trident concave and serrated. Nerve ring situated 0.25—0.40 mm, excretory pore 1180.4 mm, cervical papillae indistinct, situated 0.44 mm from the cephalic end. FIGURE 65. Serpinema octorugatus (Baylis, 1933): a — cephalic end of female, lateral; b — trident, dorsoventral; c — caudal end of male, ventral; 4 — distal end of right spicule, lateral (after Baylis, 1933). 111 Male. Length 8.6—11.63 mm, maximum width 0.38—0.43 шт. Valves of buccal capsule 0.14—0.15 mm wide. Length of esophagus 1.28—1.3 mm, of muscular anterior part 0.44—0.45 mm, of glandular posterior part 0.68—0.7 mm. Maximum width of esophagus 0.14—0.16 mm. Tail of the form typical for the genus. Wings fleshy and muscular, extending to the ventral surface anteriorly, curved inward ventrally, forming a deep longi- tudinal groove, at the posterior end of which the cloaca is situated. Tail about 0.13—0.14 mm long. There are usually 7 pairs of pedunculate pre- anal papillae, 4 pairs of postanal papillae situated close together, and one pair of small, sessile lateral papillae at the end of the tail. Spicules of markedly different length and form. Left spicule weakly chitinized, slender, about 0.5 mm long. Right spicule thicker and strongly chitinized, 0.75— 0.76 mm long without the lateral process. It is tubular, with membranous wings at the distal end. A process about 0.14 mm long laterally at its end, bent almost at a right angle to the longitudinal axis. The process is narrow; it becomes thicker distally and has membranous wings. Female. Length 15.6—19 mm, maximum width 0.55—0.62 mm. Valves of buccal capsule 0.17—0.18 mm wide. Length of esophagus 1.57 mm, of muscular anterior part 0.51—0.6 mm, of glandular posterior part 0.87— 0.88 mm. Maximum width of esophagus 0.19—0.2 mm. Tail 0.24—0.34 mm long, with a pair of indistinct papillae 0.14—0.19mm 119from the end. Vulva situated 6.8—7.8 mm from the anterior end, with pro- truding anterior lip. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.26; Baylis, 1933, рр. 628—632;. Yeh, 1960b,, pp.107— IGE Serpinema trispinosa (Leidy, 1852) Yeh, 1960 (Figure 66) Synonyms: Cucullanus trispinosus Leidy, 1852; Camatitiames trispinosus (Leidy, 1852) Railliet and Henry, 1915; C. chelydrae MacCallum, 1918; С. cyathocephalus MacCallum, 1918; С. ele- gans MacCallum, 1918; С. floridianae. Mec€allum, 1918; ©. scabrae MacCallum, 1918: С. troosti MacCallum! 918; Сова ricanus Magath, 1919; C. microcephalus (in part) sensu Chitwood, 1932; C. microcephalus sensu Rausch, 1947 Host: Pseudemys decussata. Localization: stomach and small intestine. Distribution: Cuba. Description (after Barus and Moravec, 1967). Medium-sized forms with transversely striated cuticle. Anterior end slightly flattened, narrow- ing toward the buccal capsule. Buccal capsule large, strongly pseudochi- tinized, consisting of two lateral valves on which there are two lateral groups of 5 ribs extending from the apex to the base of the inner surface. Between these two lateral groups are 4—5 short ribs near the anterior margin of the capsule. Each valve with two transversely striated plates. A strongly pseu- dochitinized ring at the base of the buccal capsule, which is slightly narrower than the valve. Tridents almost reaching the chitinized ring. Four sub- apical oral papillae, two dorsolateral and two ventrolateral. A pair of small 112 lateral cervical papillae 0.002—0.003 mm long. Esophagus consisting of a relatively short, bottle-shaped muscular part which is wider posteriorly and a cylindrical glandular part of about the same length. Muscular part of esophagus lined with cuticle. Nerve ring surrounding muscular part of esoph- agus near the buccal capsule. Intestine straight, reddish brown. Small unicellular rectal glands around the rectum. Male. Length 6.93—7.45 mm, maximum width 0.27—0.28 mm. Length of buccal capsule 0.129—0.135 mm, width 0.150—0.171 mm. Tridents 0.075—0.081 mm long. Length of muscular part of esophagus 0.44—0.46 mm, width of its widened part 0.15 mm. Glandular part of esophagus 0.50— 0.54 mm long. The small cervical papillae are situated 0.36 mm from the anterior end. Nerve ring surrounding muscular part of esophagus at its anterior end, 0.20—0.23 mm from the cephalic end. Tail curved ventrally, with subventral caudal wings; 12 pairs of pedun- culate caudal papillae, 7 pairs preanal and 5 pairs postanal. The first, second, third, and fourth pairs of postanal papillae are situated close to- 120 gether. Spicules of different length. Longer spicule slightly widening at the proximal end, 0.788—0.816 mm long, smaller spicule 0.360—0.385 mm. Tail 0.163—0.174 mm long. 02 mm © FIGURE 66. Serpinema trispinosa (Leidy, 1852): а — cephalic end, lateral; b — same, ventral; с — caudal end of male, lateral; а — end of tail of female; e — caudal end of female, lateral (after Baru§ and Moravec, 1967). Female. Length 10.60—13.00 mm, maximum width 0.27—0.28 mm. Buccal capsule reddish brown, 0.159 mm long, 0.180—0.195 mm wide. Tridents 0.075—0.111 mm long. Muscular part of esophagus 0.48—0.55 mm long, 0.149—0.163 mm wide. Glandular part of esophagus about as long as the muscular part,0.43—0.61 mm. The small cervical papillae are situated 0.38 mm, nerve ring situated 0.24—0.25 mm from the cephalic end. Tail conical, distinctly thinner than the body, 0.16—0.27 mm long. It ends in three cuticular processes. Vulva situated 5.16—5.65 mm from the posterior end, its anterior lip protruding, posterior lip small. Vagina relatively short, directed anteriorly. Larvae present in the uterus. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.29; Barus and Moravec, 1967, pp.318—320; Yeh, 1960b, pp. 117—124. 113 те] Genus Zeylanema Yeh, 1960 Synonym: Camallanus Railliet and Henry, 1915 (in part) Diagnosis. СатаПаптае. Buccal capsule with two valves with inner longitudinal denticulate ribs, ending in a chitinized ring before the connection with the esophagus. Tridents present or absent. Male with small caudal wings and two simple spicules of different length. Parasites of fish. Type species: Zeylanema anabantis (Pearse, 1933) Yeh, 1960. Zeylanema anabantis (Pearse, 1933) Yeh, 1960 (Figures 67, 68) Synonyms: Camallanus anabantis Pearse, 1933 (in part);.C; unsipiculus Khera, 1954 Hosts: Anabas testudineus, Puntius filamentosus, Rasbora daniconius. Localization: intestine. Distribution: Thailand, Ceylon, India. Historical review Pearse (1933) described this species from Anabas in Bangkok. Chakravarty (1939) gave a redescription and a drawing of it from the typical host from India. Kulasiri and Fernando (1956) found itinCeylon. There are inconsistencies in the original description and drawings of Pearse, as noted by Yeh (1960b). Pearse stated that the males have tridents and the females, which were much smaller, had none. Yehconcluded that Pearse had apparently examined two different species, one with, the other without tridents. The presence of two species of the same family inthe same host is a mixed infection, whichis quite common, Yehexamined the material of Pearse which consisted ofa single female with10 longitudinal buccal ribs anda pair of tridents; this . corresponds to the description of the female by Pearse. The buccal valves, how- ever, corresponded to Pearse's description of the male, and there is no doubt that the female and male do not belong to the same species. On the basis of the description and drawing of male and female from the collection of Pearse, and also according to the data of the two above publi- cations, Yeh retained the species described by Pearse, and also gave a de- scription of his own material given below. Yeh renamed the other species described by Pearse Zeylanema pearsei Yeh,1960 (Camallanus anabantis Pearse,1933,in part) Camallanus unispiculus Khera, 1954 is considered a synonym of Zeylanema anabantis. Below we give the description after Pearse, Chakravarty, Yeh, and Khera. Description (after Pearse, 1933), Cuticle transversely striated. Buccal capsule with two valves with 7—10 ribs. Male. Length 6.7 mm, width 0.1 mm. Tridents situated above and below the buccal valves. Chitinized buccal capsule 0.09 mm long. Muscular part of esophagus 0.33 mm long, glandular part 0.54 mm. Nerve ring situated 0.07—0.09 mm behind the buccal capsule. Tail 0.063 mm long, conical, curved ventrally, pointed, with two small spines at the end. Four pairs of preanal papillae,5 pairs of postanal papillae, and 3 pairs of discoid processes 114 122 123 at the end of the tail. Right spicule 0.59 mm long, left spicule 0.14 mm. А depression on the dorsal surface before the anus. Description (after Chakravarty, 1939). Threadlike, transversely striated forms. Ventral pairs of cephalic papillae present. Buccal capsule with two chitinized valves, with 9 pairs of moniliform longitudinal ribs which are shorter on the dorsal and ventral side. Tridents present in both sexes. A chitinized ring at the beginning of the esophagus. Fsophagus divided into a muscular and a glandular part. Male. Length 6.15mm, width 0.15mm. Length of muscular part of esophagus 0.4 mm, of glandular part 0.55 mm. Chitinized buccal capsule 0.1mm long. Nerve ring situated 0.2 mm from the cephalic end. Tail 0.12 mm long, curved ventrally, ending in a spine. Spicules of different length, right spicule longer. Short precloacal wings present. Papillae: 5 pairs preanal, 5 pairs postanal, one pair adanal, and 3 pairs at end of tail. Female. Length 21.5 mm, width 0.45 mm. Length of muscular part of esophagus 0.45 mm, of glandular part 0.96 mm. Chitinized buccal capsule 0.15 mm long. Nerve ring situated 0.25 mm from cephalic end. Tail 0.25 mm long, bifid. Viviparous. Description (after Yeh, 1960b). Buccal valves with 9 longitudinal rows of teeth in both sexes. Five median rows always present on the entire buccal capsule; two outer lateral rows are often shorter. The second outer row is usually short. Tridents distinct. Male. Length 2.9—4 mm, width 0.10—0.13 mm. Length of buccal cavity 0.07 mm, width 0.08 mm. Muscular anterior part of esophagus 0.29—0.31 mm long, posterior part 0.38—0.46 mm. Tail 0.09 mm long, with two small finger-shaped processes at the end. Six pairs of precloacal papillae, one pair situated slightly before the cloaca, and 5 pairs of postcloacal papillae, the first pair situated immediately be- hind the cloaca. A pair of phasmids present. Spicules weakly sclerotized and of different length. Left spicule 0.11 mm long, right spicule 0.58—0.65 mm long. Hemale. Lengthy .2—loe4 mm, width ©. 16—0.3 mm. Buccal cavity 0.10—0.11 mm long and 0.13—0.17 mm wide (dorsoventrally). Muscular anterior part of esophagus 0.36—0.40 mm long, posterior part 0.65—0.78 mm. Nerve ring situated 0.23—0.34 mm from the cephalic end. Tail conical, 0.15 mm long, with three small finger-shaped processes at the end. Vulva situated slightly before middle of body, dividing the body at the ratio Oi Lol ol... 3% Description (after Khera, 1954). Nematodes of medium size. Cuticle thin, with weak transverse striation. The chitinized ring situated between the valves of the buccal capsule and the esophagus large and thick. Two tetragonal outer thickenings and 9 longitudinal ribs on each valve. All longitudinal ribs have a thickening and on each thickening is a small but distinct spine. Tridents well developed, but the median branch very small. Ends of branches pointed. Three valves at the connection of esophagus and intestine. Intestine as wide as posterior part of esophagus. Rectum 0.05 mm long inthe female. Three rectal glandular cells at the connection between intestine and rectum. Nerve ring surrounding anterior part of esophagus in its narrowest part. Lateral cervical papillae small, thin, and pointed, situated at the level of the anterior part of the esophagus. 115 FIGURE 67. Zeylanema anabantis (Pearse, 1933): а — cephalic end, lateral; b,c — same, ventral; 4 — same, apical; е — caudal end of male, lateral; f— same, ventral; g — caudal end of female, lateral; В — processes on tail of females (after Yeh, 1960c). Male. Length 10.9 mm, width 0.124 mm. Cephalic end blunt, caudal end pointed. Valves of buccal capsule 0.09 mm long,0.1 mm wide. Chitinized ring 0.07 mm wide. Median branch of trident 0.029 mm long, lateral bran- ches 0.079 mm. Length of esophagus 0.85 mm, club-shaped muscular ante- rior part 0.73 mm long, cylindrical glandular posterior part 0.48 mm. Nerve 124ring situated 0.17 mm, cervical papillae 0.21 mm from cephalic end. Tail curved ventrally. A small depression (which may be the outline of the developing sucker) situated 0.43 mm from the posterior end. Caudal wings absent. Tail 0.058 mm long. Lips of cloaca protruding. Sixteen pairs 1250f sessile caudal papillae, 14 pairs precloacal and 2 pairs postcloacal. Only the right spicule present, 0.63 mm long, weakly chitinized, thin, and difficult to see; its proximal end wide and bifid, the distal end thin and pointed. The single testis is curved around the anterior end of the intestine. 116 (124) i AN, ie а — anterior end of male, lateral; b — cephalic end of female, lateral; с — trident; 4 — caudal end of male, lateral; е — caudal end of female, lateral; f — region of vulva, lateral; g — cephalic end, lateral; В, i — caudal end of male, lateral; } — caudal end of female, lateral (а — f — after Khera, 1954; р —j — after Fernando and Furtado, 1963b). FIGURE 68. Zeylanema anabantis (Pearse, 1933): Female. Length 16.3—17.2 mm, width 0.2—0.225 mm. Cephalic end blunt, caudal end pointed. Valves of buccal capsule 0.080—0.085 mm long, 0.137—0.14mm wide. Chitinized ring 0.082—0.090 mm wide. Median branch of trident 0.020—0.024 mm long, lateral branches 0.086—0.093 mm, Length of esophagus 1.45—1.47 mm, its club-shaped muscular anterior part 0.57 mm long, cylindrical glandular posterior part 0.88—0.90 mm. Nerve ring situated 0.18—0.217 mm, cervical papillae 0.23 mm from cephalic end. Tail 0.186—0.190 mm long, tapering, pointed. Vulva situated before middle of body, 6.77 mm from the anterior end in a specimen 16.3 mm long. Its lips indistinct. Ovejector situated near the vulva, large, with thick 117 muscular walls,0.24 mm long. Two opposite uteri. A single anterior ovary, curved around the end of the intestine. Posterior part of uterus with blind end. Buccal valves with 9 longitudinal denticulate ribs, the median ribs extend- ing along the whole valves. One or two of the lateral ribs shorter than the others. Pharynx short, sclerotized. Esophagus divided into a muscular part and a glandular part. Tridents distinct. Male. Length 5.94—6.72 mm, width 0.208—0.216 mm. Buccal capsule 0.072—0.079 mm long and 0.101 mm wide. Pharynx 0.021 mm long,0.072 mm wide. Muscular part of esophagus 0.352 mm long, 0.108—0.112 mm wide; glandular part 0.608 mm long,0.112 mm wide. Nerve ring surrounding mus- cular part of esophagus 0.126—0.162 mm from its anterior end, 0.216— 0.270 mm from the cephalic end. Tail conical, curved ventrally, with pointed end. Cloaca situated 0.101— 0.108 mm from end of tail. Six pairs of precloacal and 8 pairs of postcloacal papillae, one pair immediately before and one pair immediately behind the cloaca. Spicules of different length, weakly sclerotized, 0.88—1.08 and 0.252 mm long. The number of postcloacal papillae is larger than described by Yeh, but their arrangement is the same. Two pairs of small processes at the posterior end which were not mentioned by Yeh (1960). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.25; Chakravarty, 1939, pp.317—318; Fernando and Furtado, 1963, pp.64, 153; Khera, 1954, pp.100—103; Kulasiri and Fernando, 1956, pp.420—424; Pearse, 1933, pp.182—183; Yeh, 1960b, pp. 107—110; 1960c, ‘pp. 1174-124. 126 Zeylanema atridentus (Khera, 1954) Yeh, 1960 (Figure 69) Synonym: Camallanus atridentus Khera, 1954 Host: Ophiocephalus punctatus. Localization: intestine. Distribution: India. Description (after Khera, 1954). Cuticle with fine transverse striation. Buccal capsule wider than long, widest slightly behind its middle. Two lateral valves with two outer thickenings and 20 longitudinal ribs on the inside. Longitudinal ribs simple, smooth, and very thin. A ring of fine denticles at the base of the chitinized buccal capsule. Tridents absent. Esophagus divided into a shorter club-shaped muscular anterior part and a cylindrical glandular posterior part. Three valves at the connection between esophagus and intestine. Intestine as wide as posterior part of esophagus. Lumen of intestine narrow. Rectum short and narrow. Three rectal glands at the connection between the intestine and the rectum. Male. Length 4.2—4.9 mm, width 0.077—0.108 mm, Anterior end blunt, posterior end pointed. Valves of buccal capsule 0.031—0.0345 mm long, 0.039—0.048 mm wide. A large, thick, chitinized ring 0.035—0.045 mm wide at the connection between buccal capsule and esophagus. Length of esophagus 0.64—0.71 mm, length of muscular part 0.22—0.24 mm, of glandular part 0.41—0.48 mm. 118 12 т FIGURE 69. Zeylanema atridentus (Khera, 1954): a — anterior end; b — region of vulva, lateral; c — caudal end of male, lateral (after Khera, 1954). Nerve ring surrounding anterior part of esophagus 0.083—0.086 mm from the cephalic end. Cervical papillae situated 0.096—0.1 mm behind the nerve ring. Excretory pore medioventral, situated at level of cervical papillae. Posterior end curved ventrally. Tail 0.065—0.068 mm long, tapering. Caudal wings and sucker absent. Ten pairs of ventral pedunculate caudal papillae, 5 pairs precloacal, 2 pairs cloacal, and 3 pairs postcloacal. The third and fourth pairs of precloacal papillae are situated close together. The two pairs of cloacal papillae are also situated close together. Spicules weakly and not completely chitinized. Right spicule longer than left, simple and thin, 0.156 mm long. Left spicule thick, divided at the base and widening before the distal end,0.07 mm long. The single testis has the form of an '8" and is situated around the anterior end of the intestine. Female. Length 7.6—9.1 mm, width 0.11—0.124 mm. Anterior end blunt, posterior end very thin. Valves of buccal capsule 0.040—0.045 mm long, 0.063—0.068 mm wide. A large, thick, chitinized ring 0.052 mm wide at the connection between buccal capsule and esophagus. Length of esopha- gus 0.74—0.85 mm, length of muscular part 0.21—0.27 mm, of glandular part 0.53—0.58 mm. Nerve ring surrounding anterior part of esophagus 0.097—0.126 mm from the cephalic end. Cervical papillae situated behind the nerve ring 0.11— 0.135 mm from the cephalic end. Excretory pore medioventral, situated at level of cervical papillae. Tail 0.22—0.30 mm long, tapering, its end very thin. Vulva situated before middle of body, 3.5 mm from the cephalic end in a specimen 8 mm long. Vulva with protruding lips. Vagina narrow, muscular, 0.46—0.55 mm long, extending posteriorly. The two uteri are opposite and filled with larvae. Posterior branch of uterus ending blind. There is a single anterior ovary coiled around the anterior end of the intestine. Larvae thick, 0.124—0.140 mm long. References: Khera, 1954, pp.103—106; Yeh, 1960b, pp.117—120. 119 Zeylanema fernandoi Yeh, 1960 (Figure 70) Hosts: Ophiocephadds pometatuas OO. striatus. Localization: intestine. Distribution: Ceylon. Description (after Yeh, 1960b). Three females examined. They are rather thick and short. Buccal valves with 9 longitudinal ribs. The 7 median 128 ribs have a tooth posteriorly. The posterior part of the buccal capsule паг- rows rapidly and then widens before the connection with the esophagus. Tridents well developed. Cervical papillae very small. (127) 0/ шт FIGURE 70. Zeylanema fernandoi Yeh, 1960: a — cephalic end, lateral; b — same, ventral; c — caudal end of female, lateral (after Yeh, 1960b). Male unknown. Female. Length 2.9—5.4 mm, width 0.14—0.27 mm. Length of buccal capsule 0.10—0.11 mm, dorsoventral width 0.10—0.12 mm. Length of mus- cular anterior part of esophagus 0.36—0.40 mm, of posterior part 0.52— 0.65 mm. Cervical papillae situated 0.16—0.18 mm, nerve ring 0.19— 0.21 mm from the cephalic end. Vulva situated slightly behind middle of body. Tail 0.1 mm long. Reference: Yeh, 1960b, рр. 114—115. Zeylanema kulasirii Yeh, 1960 (Figure 71) Hosts; Anabas testudineus, Ophiocephalus punetatue: Localization, intestine. 129 Прето (Gevon. Description (after Yeh, 1960b). Buccal valves with 9 longitudinal rows of denticles only in the anterior 2/3 of the valves. A pair of rudi- mentary tridents present. Male. Length 1.8—2.6 mm, width 0.07 mm. Length and dorsoventral width of buccal cavity 0.05 mm. Muscular anterior part of esophagus 0.23— 0.26 mm long, posterior part 0.27—0.34 mm. The thin, ventrally curved tail ends in two finger-shaped processes. Caudal wings distinct. Six pairs of precloacal papillae, one median pair just before the cloaca, 6 pairs of post- cloacal papillae, the first pair situated just behind the cloaca. A pair of phasmids present. Spicules weakly sclerotized (difficult to measure). Left spicule 0.15—0.19 mm long, right spicule 0.40—0.42 mm long. 120 (128) FIGURE 71. Zeylanema kulasirii Yeh, 1960: a — cephalic end, lateral; b — same, ventral; с — same, apical; d — caudal end of male, ventral; e — same, lateral (after Yeh, 1960c). Reference: Yeh, 1960b, pp.112—114. Zeylanema mastacembeli Sahay and Sinha, 1966 (Figure 72) Роз Wess chee manloye Luis. ale maa tse Localization: intestine. Dis te vb ud wome) ата. Description (after Sahay and Sinha, 1966b). Thin, threadlike, medium-sized nematodes, blood-red in life, cuticle not striated. The buccal valves are wider than long in males, as wide as long in females, Valves with two outer thickenings and 9 longitudinal moniliform ribs. Tridents distinct, the three branches of almost the same length, with blunt end. Male. Length 11.60 mm, width 0.208 mm. Size of buccal valves, without the posterior chitinized ring, 0.088—0.096X0.112—0.128 mm. Chitinized ring situated at the connection between valves and esophagus, 0.072— 0.088 mm wide. Club-shaped muscular anterior part of esophagus 0.528—0.560mm long. Median branch of tridents 0.036—0.05 mm long, lateral branches 1300 .058—0.084 mm. Nerve ring situated 0.32—0.336 mm from the cephalic end. Tail, conical, blunt, pointed, curved ventrally, 0.096 mm long. 121 (129) 0.29 mm FIGURE 72. Zeylanema mastacembeli Sahay and Sinha, 1966: a — anterior end, lateral; b — trident; c — caudal end of male, lateral (after Sahay and Sinha, 1966b). Eleven pairs of pedunculate papillae in the region of the cloaca,7 pairs precloacal, 2 pairs on the protruding lips of the cloaca, andthe other 2 pairs postcloacal. A protruding cuticular tubercle is situated before the cloaca, 0.384 mm from end of tail. Caudal wings present. Only the right spicule recognizable, weakly chitinized, difficult to see, 0.336 mm long. Female. Length 19.2—26.28 mm, width 0.224—0.272 mm. Size of ‘buccal valves, without the posterior chitinized ring, 0.103—0.112X 0.123 — 0.128 mm. The chitinized ring is situated at the connection between valves and esophagus, 0.075—0.084 mm wide. Club-shaped muscular anterior part of esophagus 0.576 mm long. Median branch of trident 0.084 mm long, lateral branches 0.083—0.089 mm. Nerve ring situated 0.232—0.238 mm from the cephalic end. Tail tapering, with bluntly pointed end, 0.21—0.22 mm long. Vulva situated 6.768—9.28 mm from the anterior end (i.e. at the anterior third of the body). Vagina muscular, long, uterus with two opposite branches. Viviparous. Reference: Sahay and Sinha, 1966b, pp. 247—252. Zeylanema pearsei Yeh, 1960 (Figure 73) Synonym: Camallanus anabantis Pearse, 1933 (in part) Hosts: Rasbora daniconius, Anabas testudinews. Localization: intestine. 122 Distribution: Ceylon, Singapore. Historical revew (see Z. anabantis). Description (after Yeh, 1960). Buccal valves with 7 rows of teeth. Tridents absent. Female. Length 5.1 mm, width 0.15 mm. Length of buccal cavity 0.08 mm, width (dorsoventral) 0.007 mm. Length of muscular anterior part of esophagus 0.36 mm, of glandular posterior part 0.52 mm. Tail 0.11 mm long. Tail with three distinct terminal spines. Vulva situated slightly before middle of body, dividing the body at the ratio of 1:1.2. Description (after Pearse, 1933). Body transversely striated. Buccal capsule with a pair of valves, each with 7—10 ribs. Female. Length 4.1 mm, width 0.09 mm. Tail conical, 0.12 mm long, with two or three small spines. Vulva situated slightly behind middle of body. The uterus of some specimens contained larvae. Description (after Fernando and Furtado, 1963a,b). Relatively large forms. Buccal valves sclerotized in both sexes, with 7 longitudinal denticulate ribs. Pharynx weakly chitinized; behind it extends the muscular esophagus, 131 which passes into the glandular esophagus. Tridents absent. 2/™mm FIGURE 13. Zeylanema pearsei Yeh, 1960: a — cephalic end, lateral; b — caudal end of male, lateroventral; c — region of vulva, lateral; 4 — caudal end of female, lateral; e,f, 5 — cephalic end, different aspects; В — posterior end of female, lateral (a — Я — after Fernando and Furtado, 19636; e —h — after Yeh, 1960b). Male (one damaged specimen). Tail conical, curved ventrally, with pointed end. A small tubercle near end of tail. Cloaca situated 0.100 mm 123 132 from end of tail. Eight pairs of precloacal and 8 pairs of postcloacal ра- pillae, one pair situated just before and one pair just behind the cloaca. The second pair of precloacal papillae is very small. Spicules weakly chitinized of different length. Left spicule 0.155 mm long, right spicule 0.880 mm. Female. Length 3.680 mm, width 0.096 mm. Length of buccal capsule 0.063 mm, width (dorsoventral) 0.074 mm. Length of pharynx 0.022 mm, width 0.056 mm. Length of muscular part of esophagus 0.304 mm, width 0.056 mm. Length of glandular part of esophagus 0.480 mm, width 0.056 mm. Nerve ring surrounding muscular part of esophagus 0.130 mm from its anterior end or 0.222 mm from the cephalic end. Tail conical, ending in three finger-shaped processes. Anus situated 0.096 mm from end of tail. Vulva situated slightly behind middle of body, dividing the body at the ratio of 119055 References: Fernando and Furtado, 1963a, pp. 62—64; 1963b, pp. 141—163; Pearse, 1933, рр.182-— 1833 Yeh,1960b, pp, iL) — hia ? Zeylanema spinosa Furtado, 1965 (Figure 74) Host Вега ртека. FIGURE 74. Zeylanema spinosa Furtado, 1965: а, Б — cephalic end; с — caudal end of female, lateral; 4 — region of vulva, lateral (after Furtado, 1965). 124 Mocali zation: intestine. Distribution: Malaysia. Description (after Furtado, 1965). Large, spindle-shaped nematodes, with finely transversely striated cuticle. Cuticle with 4 longitudinal rows of cuticular spines with irregular intervals, two rows on each side. Buccal capsule with two strongly sclerotized lateral valves, with 9 longitudinal 133 denticulate ribs (each rib with about 7 hooks). The median ribs extend along the whole buccal valve; the lateral ribs are markedly shorter. Pharynx strongly sclerotized; behind it extend the straight, club-shaped muscular part and the straight, cylindrical glandular part of the esophagus. A pair of tridents present, about 0.03 mm long. Male unknown. Female. Length 6.996 mm, width 0.231 mm. Buccal capsule 0.054 mm long and 0.057 mm wide (dorsoventral). Length of pharynx 0.012 mm, width 0.048 mm. Length of muscular part of esophagus 0.231 mm, width 0.066 mm. Glandular part of esophagus 0.891 mm long and 0.072 mm wide. Nerve ring surrounding muscular esophagus 0.180 mm from its anterior end or0.240mm from the cephalic end. Tail conical, short, ending in three processes. Anus situated 0.099—0.132 mm from end of tail. Vulva situated before middle of body, dividing the body at the ratio of 1.0:1.4. Lips of vulva protruding. Reference: Furtado, 1965, pp.677—680. Zeylanema sweeti (Moorthy, 1937) Yeh, 1960 (Figures 75,76) Synonyms: Camallanus sweeti Moorthy, 1937; Paracamalla- Hos ophiocepha li Karve, 1941 Hosts: definitive — Ophiocephalus gachua; intermediate — Cyelops (— Mesocyelopsimleuckanin, ©.) hyalimmus; reser- Voirs— Barbus роке в” ticto; Lepidocephalichthys thermalis, Gambusia sp. Localization: intestine (in definitive host), wall of intestine (in reservoirs), body cavity (in intermediate hosts). ее vow ton: Мата. Description (after Moorthy, 1937b). Stoma 0.06—0.07 mm long, with two valves. A wide chitinized ring separates the valves from the esophagus. The 9 longitudinal moniliform ribs are of about the same length. They extend from the anterior end to the chitinized ring. Mouth wide, surrounded by an inner ring of 2 pairs of small papillae and an outer ring of 2 pairs of larger papillae. Amphids indistinct. Tridents present, the median branch about 0.046 mm long, the two submedian branches 0.015mm long. Two quadrilateral chitinized plates at the anterior margin of the buccal valves. Male. Length 3.0—3.9 mm, width 0.13 mm. Muscular anterior part of esophagus 0.31—0.38 mm long, glandular posterior part 0.38—0.44 mm. Tail 0.05—0.063 mm long; in immature forms there are tubercles at the end of tail as in the females; in mature males there is only one ventrolateral, median tubercle which is markedly reduced and resembles a small teatlike process. Thirteen to 15 pairs of genital papillae; 4—6 pairs preanal (5 pairs in most specimens), 2 pairs circumanal, and 7 pairs postanal. The first two pairs of postanal papillae are lateral and form a group, the third pair is 125 135 134 ventrolateral and isolated, and the sixth and seventh pairs are ventrolateral and arranged in a group; the circumanal papillae are isolated. The position of the preanal papillae varies from subventral to ventrolateral. Spicules of different length: right spicule 0.21—0.30 mm long, left spicule thinner, 0.1—0.18 mm long. A suckerlike organ situated 0.32 mm from end of tail. cy N ` 6 CIT А Caan FIGURE 75. Zeylanema sweeti (Moorthy, 1937): a — anterior end, lateral; b,c,d — cephalic end, different aspects; e — region of vulva, lateral; f,h — caudal end of female, different aspects; g, i — caudal end of male, dif- ferent aspects (after Moorthy, 1937). Female. Length 3.2—5.5 mm, width 0.15 mm. Cuticle finely striated withintervals of 0.004mm. Muscular anterior part of esophagus 0.32—0.42 mm long, glandular posterior part 0.4—0.52 mm. Nerve ring situated between anterior and middle third of muscular part of esophagus, excretory pore situated 0.14 mm from the cephalic end. Vulva situated 2.0—3.0 mm from the anterior end, with distinct, protruding lips. Posterior lip displaced laterally by an anteriorly directed process; such a process is usually not present in young, unfertilized females. Vagina narrow, extending posteriorly for about 0.3 mm. Tail with three tubercles at the end, 0.05—0.07 mm long; the tuber- cles are distinct in immature forms, the mediodorsal process markedly longer and thicker than the two ventrolateral processes. In mature specimens the processes are less distinct and of about the same length. 126 Description (after Baylis, 1939). Cuticle transversely striated with intervals of 0.004 mm. Buccal valves 0.06—0.07 mm long (including basal ring). Valves with 9 inner longitudinal denticulate ribs and two rectangular outer thickenings. Median branch of tridents about 0.046 mm long, lateral branches about 0.025 mm. Anterior part of esophagus 0.31— 0.42 mm long, posterior part 0.38—0.52 mm. Nerve ring situated at anterior third of anterior part of esophagus, the excretory pore (in females) about 0.16 mm from the anterior end. Male. Length 3.0—3.9 mm, width 0.13 mm. Tail 0.050—0.063 mm long. Three almost equal terminal spines in immature specimens. In mature males the median spine is very small and the two ventrolateral spines are larger. Thirteen to 15 pairs of caudal papillae, 4—6 pairs preanal (5 pairs in most specimens), one pair adanal, and 8 pairs postanal. The first and second pairs from the posterior end are lateral and arranged in a group, the third pair is ventrolateral and isolated, the fourth and fifth pairs are ventrolateral in a group, the sixth and seventh pairs are ventrolateral ina group, and the eighth and ninth pairs are circumanal, isolated. The position of the preanal papillae varies from subventral to ventrolateral. A sucker- like organ present about 0.32 mm from the posterior end. Right spicule 0.21—0.3 mm long. Left spicule markedly thinner, 0.1—0.18 mm long. Female. Length 3.2—3.5 mm, width 0.15 mm. Tail 0.05—0.07 mm long, with three spines at the end. Vulva situated 2—3 mm from the anterior end, with protruding lips. In fertilized females the anterior lip is displaced laterally by the overlapping posterior lip and forms a lateral process. Vagina narrow, extending posteriorly for about 0.3 mm. According to Moorthy (1937b, 1938b), the first intermediate host of this species is Cyclops leuckarti or C. hyalinus, thesecond intermediate host small fish (Barbus puckelli, В. ticto, Lepidocephalich- thys thermalis) which are eaten by the definitive host. Larvae were also found in Gambusia sp., which had been introduced from Italy for the control of Cyclops. The larvae are encysted in the wall of the in- testine in the second host. Thedefinitivehost, Ophiocephalus gachua, has an infection rate of 95% in India. The larvae are released into the intestine of the definitive host and then discharged into the water and swallowed by Cyclops. Passage of the larvae into the body cavity of Cyclops takes place 2—3 hours after in- fection. Twenty-four to 36 hours later these larvae undergo the first molt. The second molt takes place 5—12 days later, depending on the temperature. Cyclops swallowed by fish are killed by the digestive juices, particularly bile, and the larvae become active. The third molt apparently takes place 136at this time. The larvae later enter the definitive host, which eats the fish that feed on Cyclops. The fourth molt takes place in the intestine. When the parasites become mature, copulation takes place probably in the anterior part of the intestine, where the largest concentration of males and females is found. Fertilized females migrate to the posterior part of the intestine, and the larvae hatch here and are discharged into the water with the feces. They are then swallowed by Cyclops. The presence of third-stage larvae in the intestine of the definitive host (Ophiocephalus) suggests that direct infection may have taken place, although the larvae may have reached the intestine together with fish. Adult nematodes were found only in Ophio- cephalus: 127 Biology (after Moorthy, 1938Ъ). Mesocyclops leuckarti and М. hyalinus were used in experiments; adult females of М. leuckarti were most easily infected. The infection rate reached 75% in one experi- ment. Four 10 6 hours after the larvae are swallowed, they penetrate into the body cavity, where they develop further. One Cyclops may contain 12—15 larvae. Such specimens are sluggish and tend to sink to the bottom of the vessel. Infected Cyclops survived to 52 days in the laboratory at 50—56.5° and to 70 days at 30.5—38.5°. Fresh bile of Ophiocephalus kills Cyclops in8 minutes and activates the larvae. The larvae leave the Cyclops through the antennae, legs, and the boundary of cephalothorax and abdomen. After they reach the furca or oral segment, the larvae perforate the cuticle by rotatory movements of the cephalic end. Only in 3 out of 108 cases did larvae emerge from the anal segment. The larvae emerged after 50 minutes to 2 hours. Larvae which did not emerge died after 3—4 hours of active movement. Hydrochlo- ric acid in a solution of 0.1—1.0% killed the Cyclops in 2—3 minutes, but activity of the larvae was weak. The yield of larvae under laboratory con- ditions was small: only 7 larvae were obtained from 140 infected Cyclops. Third-stage larvae were found in the following naturally infected fish: Barbus puckelli, 20% B. ticto, 15% Lepidocephalichthys thermalis, 90% Gambusia sp., 5%. The larvae found in B. ticto and in Gambusia sp. were identical with the larvae of Z. sweeti from experimentally infected Cyclops. Moorthy gives the following description of the different stages of develop- ment of Z. sweeti: First-stage larva. Length 0.19—0.28 mm, width 0.12—0.018 mm. Adorsal triangular denticle present at the anterior end. Amphids in the form of openings. Nerve ring situated 0.025—0.045 mm and genital primordium 0.11—0.17 mm from the cephalic end. Phasmids large, saclike, situated slightly behind the anus. Tail 0.033—0.046 mm long, with blunt end. Im- mediately after the first molt, which takes place 24—36 hours after entry into the body cavity of Cyclops, the larvae lose the triangular denticle. Second-stage larva. The stoma becomes distinct and rectangular at this stage. Excretory pore distinct, situated slightly behind the nerve ring. After the second molt, which takes place after 5—7 days at high tem- peratures and after 8—12 days at moderate temperatures (see above), the stoma becomes partly chitinized and differentiated. The outer ring of papil- lae, consisting of two pairs, now becomes recognizable. The genital pri- mordium consists, as at the preceding stage, of four cells. Its length re- mains unchanged but it is twice as wide. Three characteristic pointed processes appear at the end of the tail (under the exuvia). 137 Third-stage larva (from experimentally infected Cyclops). Buccal capsule partly chitinized and incompletely differentiated from the esophagus by chitinized valves. Eight to 11 partly chitinized moniliform ribs of almost equal length extend from the anterior margin of the buccal valve. Thegenital primordium consists of 8 cells. The mediodorsal caudal process is markedly more massive than the others. 138 Larvae of this stage from the intestine of naturally infected fish (B. puckelli and L. thermalis) are,ata later stage of development, al- most one and a half times as long as specimens obtained from Cyclops. Chitinization of the buccal apparatus is complete. 128 (137) FIGURE 76. Zeylanema sweeti (Moorthy, 1937): a — first-stage larva, general view; b — anterior end of first-stage larva; c — anterior end of second-stage larva; Ч — same, posterior end; e — anterior end of larva during first molt (after Moorthy, 1938). Larvae were not examined during the third molt. Fourth-stage larva (after the fourth molt, from the intestine of naturally infected Ophiocephalus. Thefourthstage was determined by the degree of development of the genital primordium and the presence of a vulva. The stoma resembles that of the adult. The sex is easily determined. Fifth-stage larva (female), Anterior and posterior parts of uterus developed. Anterior part ending in a long tubular ovary, posterior part with a small, rudimentary ovary. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.28; Baylis, 1939, pp.212—214; Karve, 1941, pp. 34-37; Moorthy, 1937, pp.302—306; Moorthy, 1938, рр.323—342; Yamaguti, 1961b, p.43; Yeh, 1960b, pp.107—116. 129 SUBFAMILY PROCAMALLANINAE YEH, 1960 Synonym: Neocamallaninae Chakravarty and Majumdar, 1960 Diagnosis. Camallanidae. Buccal capsule entire, with or without thickenings; tridents absent. Type genus: Procamallanus Baylis, 1923. Key to the genera of the subfamily Procamallaninae ВИРА <, ЧН hain i bios, A ле Spirocamallanus Olsen, 1952. 2 (1)... Buceaiieapsule smooth, without‘spiralthickenings}tte ое +; be) nda RE oc Nas ates а И. 2 Procamallanus Baylis, 1923. Genus Procamallanus Baylis, 1923 Synonyms: Cucullanus Muller, 1777 (in part); Camallanus Railliet and Henry, 1915 (in part); Thelazo Pearse, 1933 (?); Indo- camallanus Chakravarty, Majumdar and Sain, 1963 Diagnosis. Procamallaninae. Buccal capsule with rounded cross section, usually longer than wide, widest in the middle. Capsule sometimes as long as wide. Inner surface of capsule smooth. Mouth usually hexagonal, with 6 rudimentary processes at its margin. Cephalic end with 4 large sub- median papillae and amphids which are situated slightly nearer the mouth than the papillae. Esophagus divided into a muscular anterior and a glandular posterior part. Male. Tail conical; caudal wings present. Nine pairs of riblike pre- anal papillae. Adanal and postanal papillae present. Right spicule some- ‘times with wings. Gubernaculum usually absent. Female. Vulva situated before middle of body. Posterior ovary absent. Viviparous. A group of teatlike appendages of varying number at the end of the tail. Parasites of the stomach and intestine of fish and amphibians. Larvae found in Copepoda. Type species: Procamallanus laeviconchus (Wedl, 1862) Railliet and Henry, 1915. 139 Procamallanus laeviconchus (Wedl, 1862) (Figure 77) synonyms: Cucullanus laeviconchus Wedl, 1862; Cama ame laeviconchus (\еа1, 1862) Railliet and Henry, 1915 Hosts: Synodontis schaal, Bagrus bayad. Localization: intestine. Distribution: North Africa. 130 140 Description (after Baylis, 1923а). Cuticle coarsely striated with intervals of 0.020 mm. Width of cephalic end at the widest part of the buccal capsule 0.06—0.08 mm. Buccal capsule 0.0675—0.07 mm long, 0.0425— 0.06 mm wide. Esophagus consisting of a muscular anterior part and a long glandular posterior part. Nerve ring situated 0.175—0.2 mm, excretory pore 0.43 mm (female) from the cephalic end. s > tare | ea КИТ «ШИ FIGURE 77. Procamallanus laeviconchus (Wedl, 1862): a — cephalic end; b — caudal end of male, lateral; c — anterior end; d — caudal end of female, lateral; е — region of vulva, lateral; f — anterior end; g — caudal end of female, lateral; h — caudal end of male, ventral (a — after Yorke and Maplestone, 1926; b — e — after Tornquist, 1931; f—h — after Campana-Rouget, 1961a). Male. Length 3.65 mm, width 0.11 mm. Distance from cephalic end to posterior end of muscular part of esophagus 0.4 mm; length of esophagus 1mm. Tail 0.037 mm long. Nine pairs of riblike preanal papillae (some- times 8 or 10 papillae on one side and 9 on the other). One pair of papillae similar to the preanal papillae lateral to the cloaca. There are also ap- parently two small subventral pairs at the sides of the cloaca, one pair behind the other, and three small pairs of postanal papillae which are some- times indistinct. Left spicule 0.05 mm long, right spicule 0.15 mm. Gubernaculum not found. Female. Length 15.5 mm, width 0.35 mm. Distance from cephalic end to posterior end of muscular part of esophagus 0.47 mm, length of esophagus Тута. Tail conical, tapering, ending in three short, blunt processes. Tail 0.15 mm long. Vulva with protruding anterior lip, situated slightly before middle of body (7 mm from anterior end in a specimen 15.5 mm long). Vagina at first narrow, winding posteriorly for about 1.1 mm before passing into the uterus, then gradually widening. Only one ovary. Posterior branch of uterus ending blind, about 1.4 mm from the posterior end. The loops of the ovary extend to 0.4—0.9 mm behind the connection between esophagus and intestine. Viviparous. 131 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.30; Baylis, 1923a, pp.24—38; Yamaguti, 1961b, р. 46. Procamallanus armatus Campana-Rouget and Therezien, 1965 (Figure 78) Host: A@geupilla austradis. Localization: stomach. Distribution: Madagascar. Description (after Campana-Rouget and Therezien, 1965). Body tapering at both ends. Buccal capsule oblong, with smooth walls. Esophagus divided into a muscular and a glandular part. Tail conical, with processes in the female, short and forming a spiral in the male. Excretory pore and deirids situated at about the same level, slightly behind the nerve ring. (141) 0/00 пит 0100 пт FIGURE 78. Procamallanus armatus Campana-Rouget and Therezien, 1965: a — cephalic end, dorsoventral; b — same, lateral; c — same, apical; d — caudal end of female, lateral; е — caudal end of male, ventral; f — larva from uterus (after Campana-Rouget and Therezien, 1965). 132 141 Male. Length 5.65 mm, width 0.23 mm. Buccal capsule 0.120 mm long, 0.072 mm wide. Anterior part of esophagus 0.500 mm long, posterior part 0.665 mm. Nerve ring situated 0.210 mm, deirids 0.272 mm, excretory pore 0.318 mm from cephalic end. Eight pairs of preanal papillae almost equally spaced, their stalks embedded in the cuticle; 2 pairs of small adanal papillae near the cloaca; a group of postanal papillae consisting of a pair which re- sembles the preanal papillae, then smaller lateroventral papillae, the first pair distinctly separated from the others, and two or three indistinct pairs. Right spicule 0.190 mm long, thin, strongly chitinized, widened in the form of a hook with a blunt process at the distal end so that the spicule looks like a harpoon. Left spicule absent. Female. Length 8.8—13.25. Description of a specimen 12.93 mm long. Width at the vulva 0.35 mm. Buccal capsule 0.120 mm long and 0.092 mm wide, round in apical view, but appears like six regularly arranged festoons. Mouth round, surrounded by 6 small chitinoid formations, two median, two lateroventral, and two laterodorsal. Outside the capsule there are four sub- median papillae in the outer ring (papillae of inner ring not recognizable) and two lateral amphids. Inner opening of buccal capsule slightly triangular. Length of muscular part of esophagus 0.590 mm, of glandular part 0.800 mm. Anus situated in a posteriorly directed slit 0.150 mm from the conical end of the tail which bears three blunt processes. Nerve ring situated 0.230 mm, deirids 0.260 mm, excretory роге 0.310 mm from cephalic end. Vulva situated slightly behind middle of body (7 mm from cephalic end), slightly protruding. Vagina short, directed anteriorly and joined by the two opposite branches of the uterus with thin walls, extend- ing to the esophagus and the rectum, respectively, and containing numerous embryos with a thin tail, 0.400—0.410 mm long. Reference: Campana-Rouget and Therezien, 1965, pp.165—170. 142 Procamallanus aspiculus Khera, 1955 (Figure 79) Host: Bagarius bagarius. Localization: intestine. Distribution: India. Description (after Khera, 1955). Male. Length 4.8 mm, width 0.12 mm. Cuticle thin, transversely striated with intervals of 0.004—0.007 mm. Buccal capsule smooth except for platelike processes at the base. A crescent-shaped cuticular process on the inner surface of the dorsal side of the buccal capsule. Cuticular ring represented by two spinelike formations at the base of the buccal capsule. Two processes in the cavity of the capsule. Buccal capsule spherical, 0.075 Ж0.075 mm. Esophagus divided into a club-shaped anterior part 0.27 mm long and a glandular posterior part also 0.27 mm long. In- testine with a narrow lumen, Nerve ring situated 0.15 mm, the small lateral cervical papillae 0.3 mm, and the excretory pore 0.32 mm from the cephalic end. Tail curved ventrally, with conical end, 0.41 mm long. Caudal wings well developed. Six pairs of precloacal and 2 pairs of postcloacal pedunculate papillae. The peduncles of the precloacal papillae are larger than those of the postcloacal papillae. Spicules and gubernaculum absent. The single 133 testis is situated 0.25 mm behind the glandular part of the esophagus. It resembles a figure 8. НН ED) his if Ц 27mm FIGURE 79. Procamallanus aspiculus Khera, 1955: a — anterior end, lateral; b — cephalic end; c — caudal end of male, lateral (after Khera, 1955). Female not known. Reference: Khera, 1955, рр. 243— 252. 143 Procamallanus attui Pande, Bhatia, and Rai, 1963 (Figure 80) Host: Wallago авт. Localization: body cavity. Distributvony,y Ша: Description (after Pande, Bhatia, and Rai, 1963b). Long, milk-white nematodes. Buccal capsule slightly yellowish brown, with smooth walls and with 4 oval chitinized plates, each with a fringe of 10—12 leaflike processes. Tail ending in 3 finger-shaped processes in both sexes. Male. Length 7—22 mm, width 0.247—0.399 mm. Buccal capsule cup- shaped, 0.13—0.133 mm long and 0.095—0.152 mm wide. Length of esophagus 1.197—2.166 mm, length of muscular anterior part 0.627—1.026 mm, of glandular posterior part 0.57—1.14 mm. Nerve ring situated 0.34—0.361 mm, excretory pore 0.55—0.77 mm from cephalic end. Tail curved ventrally, with wide caudal wings which extend 2.18—2.32 mm anteriorly from the posterior end. Spicules of different length and form: right spicule longer and thicker, its anterior part wider and the posterior part narrowing, 0.27—0.28 mm long; left spicule needle-shaped, 0.1—0.17 mm long. Fifteen pairs of caudal papillae: 10 longer, riblike preanal pairs equally spaced and 5 postanal pairs, the last pair situated above the trifid end of the tail. Female. Buccal capsule barrel-shaped, 0.247—0.25 mm long, 0.052— 0.073 mm wide. Length of esophagus 1.11—3.34 mm, length of muscular anterior part 0.6—1.33 mm, of glandular posterior part 0.51—1.91 mm. 134 0.9mm FIGURE 80. Procamallanus attui Pande, Bhatia, and Rai, 1963: a — cephalic end of male; b — caudal end of male; c — cephalic end of female, lateral; d — caudal end of female (after Pande, Bhatia, and Rai, 1963а). 144 Nerve ring situated 0.23—0.49 mm, excretory pore 0.29—0.95 mm from cephalic end. Genital opening without a swelling near the vulva, situated Ze lO mm perore the caudal end. Tail 0.10.38 0mm long, ending in 3 pointed processes. Reference: Pande, Bhatia, and Rai, 1963b, рр. 105—118. Procamallanus brevis Kung, 1948 (Figure 81) ог: Or. ое виа етом: iabestine. ЕЕ Ото: Бон Ее геа. Description (after Kung, 1948). Body narrowing in the posterior third, curved ventrally. Cuticle with distinct transverse striation. Mouth surrounded by 6 small papillae, four submedian and two lateral. The barrel- shaped buccal capsule is oblong and laterally compressed. Its walls are strongly chitinized, without ribs. The wall of the lower part of the capsule is thicker than that of the upper anterior part, and its strongly thickened poste- rior marginforms a ringlike connection with the anterior end of the esophagus. Cup-shaped cuticular processes project into the buccal capsule from its base. Male. Length 1.75—1.85 mm, width 0.083—0.092 mm. Buccal capsule 0.09 mm long and 0.065 mm wide. Muscular anterior part of esophagus club-shaped, 0.165—0.170 mm long. Nerve ring situated 0.132 mm, excre- tory pore 0.172 mm from cephalic end. Tail.blunt, curved ventrally. Caudal 145 wings thin, about 0.32 mm long. Eight or 9 pairs of equally shaped preanal papillae. One pair of adanal papillae, 3 pairs of small circumanal papillae, and 4 pairs of postanal papillae, the last pair the largest, situated near the medioventral line. Between this last pair is a single sessile papilla. Right spicule 0.079—0.089 mm long, left spicule weakly chitinized, 0.039 mm long. 135 (144) 2.7mm FIGURE 81. Procamallanus brevis Kung, 1948: а — anterior end, lateral; b — caudal end of male, lateral; с — same, ventral; d — caudal end of female, lateral (after Kung, 1948). Female. Length 1.95—2.1 mm, width 0.15—0.175 mm. Length of club- shaped muscular anterior part of esophagus 0.198—0.22 mm. Nerve ring situated 0.016 mm from cephalic end, excretory pore situated behind it. Tail 0.016 mm long, ending in 5—7 spinelike processes. Vulva situated behind middle of body, usually 0.83—0.904 mm from posterior end. Vagina 0.51 mm long, extending posteriorly almost to the tail. Two branches of uterus, filled with numerous larvae. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.30; Kung, 1948, pp.146—148. Procamallanus chakravartyi (Chakravarty, Majumdar, and Sain, 1961) Fernando and Furtado, 1963 (Figure 82) Synonyms: Neocamallanus heteropneusti Chakravarty, Majumdar, and Sain,1961; Indocamallanus heteropneusti Chak- ravarty, Majumdar, and Sain, 1963 Host: Heteropneustes fossilis. LocalizatLren: «stomach: DVSt Proton se nea: Historical review This species was describedfrom Heteropneustes fossilis as Neo- camallanus heteropneusti. However,the name Neocamallanus is preoccupied by Ali (1956), and the authors therefore proposed т 1963 to name it Indocamallanus andthenewspecies Indocamallanus hetero- pneusti. Theauthors comparedthe genus Indocamallanus withthe ge- nus Procamallanus, from whichit differs only inthe presence of a guber- naculum; this seems insufficient to establisha genus inthe Camallanidae, since the size and form of the spicules and gubernaculum vary inthis family according to the degree of chitinization (Yeh, 1960b; Fernandoand Furtado,1963b). 136 Fernando and Furtado (1963) placedthis speciesinthe genus Procamalla- nus, butsincethereisaspecies P. heteropneusti Ali, 1956, they named it Procamallanus chakravartyi. Theauthors noted that three species of Procamallanus havebeendescribedfrom Heteropneustes fos- Tels SPpLeCuULOouUpernaculus Agrawal, 936, Pe eonius us Fernando and Furtado, 1963, and Р. chakravartyi Fernando and Furtado, 1963; these are probably the same species, but they are considered as valid species until further material becomes available. Description (after Chakravarty, Majumdar, and Sain, 1961). Cuticle coarsely striated. Males smaller than females. Mouth forming a dorsoven- tral slit surrounded by a chitinized buccal capsule. Cephalic papillae and tridents absent. Esophagus divided into a club-shaped muscular anterior 146 part and a longer glandular part. Male. Length 3.9—4.0 mm, width 0.120 mm. Length of buccal capsule 0.048—0.060 mm, width 0.036—0.048 mm. Esophagus divided into a mus- cular and a glandular part: length of muscular part 0.204—0.228 mm, width 0.072 mm; length of glandular part 0.36—0.44 mm, width 0.72 mm. Nerve ring situated 0.120—0.144 mm from cephalic end. The curved, conical tail has caudal wings. Tail 0.036—0.048 mm long. Fifteen pairs of pedunculate caudal papillae, one pair postanal. Only the right whiplike spicule present, 0.216—0.252 mm long. Gubernaculum consisting of three parts, two of them shorter than the third. The shorter parts apparently originate on the main stem; short parts 0.048 and 0.039 mm long, longer part 0.072 mm. ES COO tam egg vasa vane 02 mm FIGURE 82. Procamallanus chakravartyi (Chakravarty, Majumdar, and Sain, 1961): a — anterior end; b — caudal end of male (after Chakravarty, Majumdar, and Sain, 1961). Female. Length 4.8—7.2 mm, width 0.156—0.168 mm. Length of buccal capsule 0.084—0.092 mm, width 0.060—0.068 mm. Esophagus divided into a muscular and a glandular part: length of muscular part 0.30—0.324 mm, width 0.048—0.072 mm; length of glandular part 0.336—0.540 mm, width 0.048—0.072 mm. Nerve ring situated 0.168—0.192 mm from cephalic end. Tail conical, without papillae, 0.096—0.132 mm long. Vulva situated near 137 middle of body, 2.4—3.3 mm from end of tail. Uteri opposite. Viviparous. Length of larvae 0.264—0.272 mm, width 0.024—0.028 mm. References: Chakravarty, Majumdar and Sain, 1961, pp. 221—224; Chakravarty, Majumdar, and Sain, 1963, pp. 415—416; Fernando and Furtado, 1963, 2.156. Procamallanus clarius Ali, 1956 (Figure 83) Host; Claritas. batrachus. Localization: stomach. Distribution: India, Singapore. 147 j FIGURE 83. Procamallanus clarius Ali, 1956: а — cephalic end, apical; b — same, lateral; с — anterior end of female, lateral; а — caudal end of female, lateral; е — caudal end of male, ventral; f — region of vulva, lateral; g — spicules, ventral; В — cephalic end, lateral; i — caudal end of male, lateral; j — region of vulva, lateral; К — caudal end of female, lateral (a —g — after Ali, 1956; h—k — after Fernando and Furtado, 1963b). 138 148 Historical review Procamallanus clarius wasdescribedfrom Clarias batrachus from India. The species was recorded by Fernando and Furtado (1963a) from the same host in Singapore; their material agrees with the description of Ali but their specimens are slightly larger. Description (after Ali, 1956). Livenematodes reddish. Females thicker and longer thanmales. Cuticle finely transversely striated with intervals of 0.007 mm. The body narrows inthe region of the esophagus inbothsexes; ce- phalic endtruncate. Tailshort, conical. Buccal capsule without spiral thicken- ings. Two pairs of submedian cephalic papillae and a pair of amphids. Male. Length 2.44—3.13 mm, maximum width 0.089—0.093 mm. Width of cephalic end 0.043—0.045 mm. Excretory pore situated 0.21—0.26 mm from cephalic end. Buccal capsule inthe form of avase, 0.053—0.055 mm long and 0.055—0.058 mm wide. Nerve ring situated 0.12—0.16 mm from cephalic end, surrounding the muscular part of the esophagus. Muscular anterior part of esophagus 0.23—0.28 mm long, glandular posterior part 0.32—0.38mm long. Length of esophagus 1/4 of length of body. One threadlike testis which extends to the esophagus, where it describes loops and ends in a posteriorly directed part. Vas deferens as wide as the testis; seminal vesicle opening into the cloaca through the short ejaculatory duct. Tail broadly curved, its thin, pointed distal part included in the caudal wings which are very narrow and difficult to see except at the end. Fourteen pairs of caudal papillae, 10 pairs preanal and 4 pairs postanal. The first three pairs of postanal papillae form a group; the last pair is isolated at the end of the tail. Immediately before the last pair of postanal papillae is a pair of transparent cuticular processes which are characteristic for the genus. Spicules thin and weakly chitinized, of similar form but of different length. Right spicule 0.085—0.087 mm long, left spicule 0.029—0.030 mm long. Both spicules are widened in the proximal part and pointed in the distal part. Tail 0.046 — 0.047 mm long, about 1/60 of length of body. Female. Length 5.69—6.02 mm, width 0.133—0.149 mm. Width of cephalic end 0.074—0.080 mm. Excretory pore situated 0.330—0.336 mm from the cephalic end. Buccal capsule 0.058—0.065 mm long and 0.065— 0.073 mm wide. Posterior end narrowing a short distance before the anus, end of tail blunt. Vulva with protruding lips, situated between third and last quarter of body, 1.67 mm from end of tail. Vagina 0.35 mm long, directed posteriorly and then turning anteriorly. Two branches of uterus, the pos- terior branch ending blind, the anterior branch passing into the ovary through the narrow oviduct. Ovary reaching glandular part of esophagus, but its terminal part is directed posteriorly. Seminal vesicle with an inflated part at the connection between oviduct and uterus. Tail 0.037 mm long, i.e. about 1/150 of length of body. Viviparous. Description (after Fernando and Furtado, 1963). Females longer and thicker than males. Buccal capsule barrel-shaped, weakly sclerotized, without thickenings. Pharynx opening into the straight muscular part of the esophagus, which is followed by the straight glandular part. Male. Length 4.24 mm, width 0.096 mm (one specimen). Length of buc- cal capsule 0.047 mm, width 0.040 mm. Length of pharynx 0.011 mm, width 0.025 mm. Length of muscular part of esophagus 0.360 mm, width 0.054mm; length of glandular part 0.576 mm, width 0.054 mm. Nerve ring surrounding muscular part of esophagus 0.126 mm from its anterior end ог 0.180 mm from the cephalic end. 139 Tail conical, curved ventrally, with pointed end. Cloaca situated 0.047 mm 149 from end of tail. Ten pairs of preanal and 4 pairs of postanal papillae, one pair at the anus. Spicules of different length, straight, one spicule three times as long as the other; length 0.123 and 0.043 mm, respectively. Female. Length 5.6—6.1 mm, width 0.144—0.176 mm (two specimens). Buccal capsule 0.076—0.080 mm long and 0.058—0.062 mm wide. Length of pharynx 0.010 mm, width 0.036 mm. Length of muscular part of esophagus 0.370—0.400 mm, width 0.080 mm; length of glandular part of esophagus 0.560—0.610 mm, width 0.064—0.080 mm. Nerve ring surrounding muscular part of esophagus 0.090—0.108 mm from its anterior end or 0.180—0.208 mm from the cephalic end. Tail conical, ending in two finger-shaped processes. Anus situated 0.108—0.140 mm from end of tail. Vulva situated behind mid- dle of body, dividing the body at the ratio of 1.2:1—1.4:1. Viviparous. References: Ali, 1956, pp.6—8; Fernando and Furtado, 1963a, pp. 64—65. Procamallanus confusus Fernando and Furtado, 1963 (Figure 84) Host: Heteropneustes fossilis. Localization: intestine. Distribution :. Ceylon. Description (after Fernando and Furtado, 1963b). Male. Length 3.15—5.0 mm, width 0.045—0.085 mm. Length of muscu- lar part of esophagus 0.235 mm, width 0.039 mm; length of glandular part 0.270 mm, width 0.030 mm. Nerve ring situated 0.114 mm from the ce- phalic end, 0.066 mm from anterior end of esophagus. Tail strongly curved ventrally; caudal wings weakly developed. Eight pairs of preanal and 8 pairs of postanal papillae. Two pairs of papillae situated around the cloaca. Spicules of different length and form. The smaller left spicule is wide and slipper-shaped, 0.045 mm long and 0.009 mm wide; right spicule sometimes thickened, 0.255 mm long. FIGURE 84. Procamallanus confusus Fernando and Furtado, 1963: a — anterior end; b — caudal end of male, lateral; с — caudal end of female, lateral (after Fernando and Furtado, 1963Ь). 140 150 Female. Length 4.2—4.8 mm, width 0.085 mm. Length of buccal capsule 0.06 mm, width 0.045 mm. Muscular part of esophagus 0.270— 0.300 mm long, 0.045 mm wide; glandular part 0.330—0.375 mm long, 0.030 mm wide. Nerve ring situated 0.150 mm from cephalic end, 0.090 mm from anterior end of esophagus. Vulva situated behind middle of body, divid- ing the body at the ratio of 1.2:1. Tail conical, 0.150 mm long. Reference: Fernando and Furtado, 1963, pp.153—156. Procamallanus daccai Gupta, 1959 (Figure 85) Host: catfish (species not determined). Localization: small intestine. Distribution: East Pakistan. Description (after Gupta, 1959). Cuticle more strongly striated in females than in males. Mouth surrounded by 2 lateral and 4 submedian papillae. The yellowish brown buccal capsule has a smooth lining, without ribs or leaflike processes, barrel-shaped. Basal ring of buccal apparatus 151 represented by two club-shaped structures; lower part of wall of buccal capsule thicker than upper anterior part, the posterior margin strongly thickened and forming a rim which is connected with the anterior end of the esophagus. (150) FIGURE 85. Procamallanus daccai Gupta, 1959: a — anterior end, lateral; b, d — caudal end of male, lateral; с — same, ventral; e — end of tail of female; f — caudal end of female, lateral (after Gupta, 1959). 141 Male. Length 3.55—4.5 mm, width 0.1—0.125 mm. Striation of cuticle with intervals of 0.013—0.016 mm. Dorsoventral width of cephalic end 0.035—0.045 mm. Esophagus divided into a club-shaped muscular anterior part 0.325—0.625 mm long, 0.042—0.05 mm wide. Nerve ring situated 0.125— 0.13 mm, excretory pore 0.18—0.22 mm from cephalic end. Tail curved ventrally, with rounded end. Tail 0.05—0.07 mm long. Caudal wings well developed. Seven or 8 pairs of long preanal papillae, one pair of adanal papillae, and 9 pairs of postanal papillae; all papillae pedunculate. Three postanal pairs are very small and are situated sub- ventrally at the sides of the cloaca, one behind the other, then follow five pairs of postanal papillae grouped together, and the small posterior pair is isolated. Preanal papillae equally spaced, adanal pair situated laterally at the cloaca. Right spicule with wide proximal end, narrow and pointed at the distal end, 0.36—0.48 mm long. Left spicule absent. Gubernaculum 0.06—0.07 mm long. Female. Length 4.76—7.39 mm, width 0.12—0.13 mm. Striation of cuticle with intervals of 0.01—0.02 mm. Esophagus divided into a club- shaped muscular anterior part 0.26—0.35 mm long, 0.03—0.04 mm wide. Nerve ring situated 0.12—0.13 mm, excretory pore 0.204—0.26 mm from cephalic end. Tail with rounded end, with two spinelike processes at the end. Tail 0.12—0.18 mm long. Vulva situated behind middle of body, 2.6—3.96 mm from cephalic end. Uterus distended by larvae. Eggs in uterus at various stages of development. Reference: Gupta, 1959, pp.775—778. Procamallanus devendri Sinha and Sahay 1966 (Figure 86) Host: Heteropneustes fossilis. Localization» stomach. Distributions India: Description (after Sinha and Sahay, 1966). Small nematodes, blood- red in life. Cuticle coarsely striated. Buccal capsule chitinized, without teeth, tridents, rods, papillae, or spiral striation. Esophagus divided into a club-shaped muscular anterior part and a longer glandular part; ratio of length of the two parts 1:1.3. Male. Length 3.2—3.5 mm, width 0.05—0.08 mm. Length of buccal capsule 0.032—0.036 mm, width 0.028—0.038 mm. Nerve ring situated 0.074—0.076 mm from cephalic end. Esophagus divided into a muscular part 0.144—0.158 mm long and a glandular part 0.21—0.24 mm long. Tail 152 conical, curved ventrally, with caudal wings, 0.03—0.04mm long. Thirteen pairs of pedunculate caudal papillae, eight pairs preanal and five pairs postanal, A winglike process situated between the third and fourth pair of postanal papillae on each side. The papillae of the fifth, last, pair are situa- ted one on each side at the end of the tail. Only right spicule present, whip- shaped, with wide proximal end, tapering distally, 0.288—0.3 mm long. Gubernaculum strongly chitinized, 0.02—0.024 mm wide in the middle, maximum width 0.024—0.028 mm, minimum width 0.01—0.014 mm. Female. Length 6.4—6.8 mm, width 0.13—0.14 mm. Length of buccal capsule 0.048—0.05 mm, width 0.03—0.032 mm. Nerve ring situated 0.062 mm from cephalic end. Esophagus divided into a muscular part 142 FIGURE 86. Procamallanus devendri Sinha and Sahay; 1966: a — anterior end of female; b — caudal end of male, ventral; c — caudal end of female, lateral (after Sinha and Sahay, 1966). 0.2—0.202 mm long and a glandular part 0.27—0.274 mm long. Tail conical, without papillae, with two short processes at the end. Tail 0.08—0.082 mm long. Vulva situated behind middle of body, 3.47—3.62 mm from cephalic end, with protruding lips. The vagina extends far posteriorly. Uterus with two opposite branches. Viviparous. Reference: Sinha and Sahay, 1966, pp. 384—388. Procamallanus glossogobii (Pearse, 1933) Southwell and Kirschner, 1937 (Figure 87) Synonym: Thelazo glossogobii Pearse, 1933 Host: Glossogobius giurus. Localization: intestine. Distribution: Thailand. Historical review Pearse (1933) found four nematodes in the intestine of Gloss ogobius giurus, for which he established the genus Thelazo with the single species Th. glossogobii Pearse,1933. His diagnosis of the genus is as follows: Buccal capsule chitinized, barrel-shaped, without valves or lips. Posterior end of tail twisted. Gubernaculum present. Spicules 143 153 of different length, caudal wings absent. Vulva situated near the anterior end. Southwell and Kirschner (1937) described Procamallanus slomei n.sp. Theydiscussedthegenus Thelazo andcametothecon- clusion that the buccal capsule in Pearse's drawing is typical for Proca- mallanus and that the nematode should be placed in that genus. Ina revision of the genus Procamallanus, Annereaux (1946) concluded that Southwell and Kirschner were wrong. He stated that the presence of a gu- bernaculum and the absence of caudal wings are sufficient reasons to maintain the genus Thelazo. However, Annereaux did not determine the systematic position of the genus Thelazo andhis argument is weakened by his statement that the genus Thelazo resembles the genus Procamallanus. This is a typical example of the establishment of new systematic units without a detailed study of the forms. The diagnosis of Pearse is too short and does not determine the systematic position of his species. We therefore consider Thelazo glossogobii asa species of the genus Procamal- lanus. FIGURE 87. Procamallanus glossogobii (Pearse, 1933) Southwell and Kirschner, 1937: a — cephalic end; b — anterior end; c — caudal end of female, lateral; d — caudal end of male (after Pearse, 1933). Description (after Pearse, 1933). Cuticle thin and transparent. Buc- cal capsule reddish brown, barrel-shaped, its anterior margin thin, thicker posteriorly especially near the base. Muscular part of esophagus slightly widened posteriorly. 144 Male. Length 2.9 mm, width 0.08 mm. Tail 0.04 mm long, conical, with rounded end. Gubernaculum 0.06 mm long, almost straight. Spicules inthe form ofa coil, right spicule 0.45 mm long, left spicule 0.17 mm long. Six sessile preanal papillae situated close together. Caudal wings absent. Female. Length3.1 mm, width 0.1 mm. Length of buccal capsule 0.07 mm. Muscular part of esophagus 0.275 mm long. Vulva situated at end of anterior part of body. Eggs and larvae in the uterus arranged ina row. Tail 0.13 mm long, conical, ending in two short, divergent spines. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.[?!; Annereaux, 1946, pp.299—303; Pearse, 1933, p.179—191;-Southwell and Kirschner, 1937, pp. 245—265. 154 Procamallanus heteropneustus Ali, 1956 (Figure 88) Host: Heteropneustes fossilis. Localization: stomach. Distribution: India. Description (after Ali, 1956). Body thin, cylindrical, slightly narrower in the region of the esophagus. Cuticle thick, transverse striation with in- tervals of 0.006 mm. Excretory pore situated slightly behind the connection between the muscular and glandular parts of the esophagus. Nerve ring sur- rounding muscular part of esophagus near its beginning. Buccal capsule widest in the middle; basal and anterior parts of the same width. Wall of buccal capsule without spiral thickenings, thicker at the base than at the apex. Mouth oval in apical view. Two pairs of submedian cephalic papillae. A pair of lateral amphids. Cephalic end truncate. 77 mm FIGURE 88. Procamallanus heteropneustus Ali, 1956: а — cephalic end, apical; b — same, lateral; с — anterior end of female, lateral; d — caudal end of female, lateral; e — posterior end of male, ventral; f — region of vulva, lateral; g — spicules, ventral (after Ali, 1956). 145 155 Male. Length 4.51—4.56 mm, width 0.09—0.10 mm. Dorsoventral width of cephalic end 0.051 mm. Excretory pore situated 0.37 mm, nerve ring 0.13 mm from cephalic end, Length of buccal capsule 0.063 mm, width 0.047 mm, anterior and posterior parts 0.029 mm wide. Length of esophagus 0.83 mm, i.e. about 1/5 of length of body, muscular part 0.27 mm long, glandular part 0.56 mm. The single testis extends to the muscular part of the esophagus. Tail with narrow caudal wings which are contiguous posteri- orly. Thirteen pairs of long, riblike caudal papillae, eight pairs preanal and five pairs postanal. The postanal papillae begin immediately behind the cloaca; three pairs are situated close together and the fifth pair at the end of the tail. Between the last two pairs of postanal papillae are two transpa- rent, blunt, rounded processes which are characteristic for the genus. Spicules weakly chitinized, of different length but of similar form, widened proximally and pointed distally; right spicule 0.073 mm long, left spicule 0.043 mm. Female. Length 6.68—7.70 mm, width 0.130—0.135 mm. Dorsoventral width of cephalic end 0.068 mm. Excretory pore situated 0.46 mm, nerve ring 0.16 mm from cephalic end. Buccal capsule 0.071 mm long, 0.059 mm wide, anterior and posterior parts 0.036 mm wide. Mouth 0.034 mm long and 0.014 mm wide. Esophagus 1.08 mm long,i.e. 1/7 of length of body; an- terior part 0.39 mm long, posterior part 0.69 mm. Vulva with distinct lips, situated 2.74 mm from the posterior end, dividing the body at the ratio of 3:1. Length of ovejector 0.57 mm. Uteri diverging. One anterior ovary; oviduct narrow. A distinct seminal vesicle present at the connection of the uteri. Ovary sinuous, its end directed anteriorly. In mature females the uteri are filled with larvae about 0.407 mm long and 0.016 mm wide. The genital primordium of the larvae adheres closely to the intestine, situated 0.133 mm from the cephalic end. Tail of females narrowing, with blunt end, 0.038 mm long. Reference: Ali, 1956, pp.3—6. Procamallanus hindenensis Lal, 1965 (Figure 89) Host: HeteropmetsStes Поз. Localization: “stomach, Distribution: ав. Description,.(after Lal, 1965) Male. Length 2.7—4.0 mm, width 0.05—0.11 mm. Buccal capsule, 0.053—0.059 mm long and 0.032 mm wide. Length of esophagus 0.493— 0.648 mm, muscular anterior part 0.166—0.298 mm long, glandular posterior part 0.255—0.426 mm. Nerve ring situated 0.138—0.170 mm from the ce- phalic end. Tail curved ventrally, with rounded end. Caudal wings well de- veloped. Thirteen pairs of pedunculate genital papillae, nine pairs preanal and four pairs postanal. Only the right spicule present, 0.227—0.514 mm long. Gubernaculum bifid anteriorly, pointed posteri- orly. 146 Female. Length 5.1—7.1 mm, width 0.11—0.17 mm. Buccal capsule 0.064 mm long, 0.043—0.059 mm wide. Length of esophagus 0.588—0.916 mm, muscular anterior part 0.268—0.357 mm long, glandular posterior part 0.341—0.639 mm. Nerve ring situated 0.192—0.224 mm from the cephalic end. Tail pointed, bifid, 0.085—0.149 mm long. One ovary. Vulva situated behind middle of body, 3.0—4.7 mm from cephalic end. FIGURE 89. Procamallanus hindenensis Lal, 1965: а — anterior end; b — caudal end of male, lateral; с — region of vulva, lateral; 9 — caudal end of female, lateral; е — spicules (after Lal, 1965). Reference: Lal, 1965, pp.199—200. Procamallanus lonis Yamaguti, 1941 (Figure 90) host?) Lovuninvaeula tus": Localization: small intestine. Distribution: Japan. Description (after Yamaguti, 1941). Male. Length 13.6—14.75 mm, width 0.32—0.35 mm. Cuticle about 0.012 mm thick, finely transversely striated at the cephalic end. Nerve ring situated 0.3—0.36 mm from cephalic end. Buccal capsule with smooth walls about 0.010 mm thick, 0.110—0.117 mm wide and 0.150—0.165 mm long, 147 including the basal ring which is 0.060—0.066 mm wide. Muscular anterior part of esophagus 0.55—0.59 mm long and 0.11 mm wide, glandular posterior part 0.85—0.90 mm long and 0.099—0.110 mm wide. 157 Tail 0.096—0.105 mm long. Caudal wings connected anteriorly, about 0.45 mm long, supported by 3 long, thin preanal papillae and 6 shorter but similar postanal papillae on each side. Right spicule 0.5 mm long, some- times sigmoid at the distal end, with narrow wings; left spicule 0.22 mm long, weakly chitinized,with almost straight distal end. a FIGURE 90. Procamallanus lonis Yamaguti, 1941: a — anterior end; b — caudal end of male (after Yamaguti, 1941). Female. Length 23.8—26.6 mm, body of almost uniform width its entire length, 0.65—0.70 mm wide, except at the ends. Cuticle 0.016 mm thick, transversely striated except at the cephalic end. Nerve ring situated 0.34—0.35 mm from cephalic end. Buccal capsule 0.13—0.15 mm wide, 0.19—0.20 mm long including the basal ring which is 0.075—0.084 mm wide. Wall of basal ring 0.013—0.014 mm thick. Muscular anterior part of esoph- agus 0.60—0.63 mm long and 0.12—0.14 mm wide, glandular posterior part 0.85—1.1 mm long and 0.11—0.13 mm wide. Tail blunt-conical, 0.16—0.18 mm long. The uterus extends to the posterior end of the muscular part of the esophagus if it contains embryos, ending 1.0—1.18 mm from end of tail. Vulva situated 8.5 mm from cephalic end, dividing the body at the ratio of 1:1.8—2.23. Larvae 0.52—0.57 mm long and 0.018—0.021 mm wide. Reference: Yamaguti, 1941, pp.343—396. 148 158 Procamallanus magurii Lal, 1965 (Figure 91) Most: Charias“batrachus*: Localization: stomach. Distribution: India. Description (after Lal, 1965). Male. Length 3.36—5.29 mm, width 0.096—0.149 mm. Buccal capsule 0.053—0.075 mm long and 0.043—0.053 mm wide. Length of esophagus 0.658—0.97 mm, muscular anterior part 0.245—0.371 mm long, glandular posterior part 0.413—0.599 mm. Nerve ring situated 0.149—0.213 mm from cephalic end. с FIGURE 91. Procamallanus Magurii Lal, 1965: a — anterior end; b — caudal end of male, ventral; с — caudal end of female, lateral; d — spicules (after Lal, 1965). Tail curved ventrally, with rounded end. Caudal wings well developed. Fourteen pairs of caudal papillae, nine pairs preanal and five pairs postanal. One spicule, 0.371—0.436 mm long. Gubernaculum bifid anteriorly, pointed posteriorly. | Female. Length 7.14—11.53 mm, width 0.149—0.256 mm. Buccal capsule 0.085—0.107 mm long and 0.048—0.064 mm wide. Length of esoph- agus 0.85—1.123 mm, muscular anterior part 0.330—0.658 mm long, 149 159 glandular posterior part 0.511—0.720 mm. Nerve ring situated 0.180— 0.213 mm from cephalic end. Tail pointed, distinctly bifid, 0.107—0.170 mm long. One ovary. Vulva situated behind middle of body, 3.427—6.141 mm from cephalic end. Referenee: Lal 1965. pp. 199—200. Procamallanus malacensis Fernando and Furtado, 1963 (Figure 92) Hoist: Channa Тастие. Localization: intestine. Distribution? dadonesia. Description (after Fernando and Furtado, 1963a). Relatively small nematodes. Female larger than male (one specimen of each sex examined), Buccal capsule weakly sclerotized, pharynx weakly sclerotized, opening into the muscular part of the esophagus; then follows the glandular part. FIGURE 92. Procamallanus malacensis Fernando and Furtado, 1963: a — cephalic end, lateral; b — caudal end of male, lateral; c — region of vulva, lateral; d — caudal end of female, lateral (after Fernando and Furtado, 1963а). Male. Length 2.640 mm, width 0.096 mm. Buccal capsule 0.040 mm long, 0.033 mm wide. Length of pharynx 0.015 mm, width 0.026 mm. Length of glandular esophagus 0.449 mm, width 0.032 mm. Nerve ring surrounding muscular part of esophagus 0.130 mm from its anterior end or 0.174 mm from the cephalic end. Tail conical, curved ventrally, with pointed end. Cloaca situated 0.040 mm from end of tail. Nine pairs of preanal and 2 pairs of postanal papillae, one pair just before the cloaca and one pair behind the anus. Spicules weakly sclerotized, of different length: larger spicule 0.130 mm long, smaller, thick- er spicule 0.050 mm, width 0.075 mm. Female. Length 4.494 mm, width 0.150 mm. Buccal capsule 0.056 mm long and 0.045 mm wide. Length of pharynx 0.019 mm, width 0.037 mm. 160 Muscular esophagus 0.352 mm long, 0.048 mm wide; length of glandular esophagus 0.528 mm, width 0.040 mm. Nerve ring surrounding muscular 150 part of esophagus 0.133 mm from its anterior end or 0.207 mm from the cephalic end. Tail conical, ending in three finger-shaped processes. Anus situated 0.089 mm from end of tail. Vulva situated just behind middle of body, dividing the body at the ratio of 1.25:1. Reference: Fernando and Furtado, 1963, рр. 65—67. Procamallanus mathurai Pande, Bhatia, and Rai, 1963 (Figures 93, 94) Host: Heteropneustes fossilis. LOCAIIZATIOM:S SromZzcia, Юаня liachiey, (161) 0.0%. mm FIGURE 98. Procamallanus mathurai Pande, Bhatia, and Rai, 1963: а — cephalic end; b — anterior end of adult female, lateral; с — caudal end of male, dorsoventral; d—right spicule; е — gubernaculum, different aspects; f — caudal end of female, ventral (after Pande, Bhatia, and Rai, 1963Ь). 151 (162) FIGURE 94. Procamallanus mathurai Pande, Bhatia, and Rai, 1963: а — first-stage larva, general view; b — anterior end of first-stage larva; с — cephalic end of third-stage larva; Я — caudal end of third-stage larva, lateral; e — third-stage larva, general view; f — anterior end of fourth-stage larva; g — posterior part of body of young female (after Pande, Bhatia, and Rai, 1963b). Description (after Pande, Bhatia, and Rai, 1963b). Light pink nematodes, with finely striated, thick cuticle. Cephalic end truncate, with 4 submedian cephalic papillae on the lower margin of the weakly chitinized rim and with a pair of lateral amphids. Buccal capsule barrel-shaped, with smooth, yellowish brown, chitinized wall. Esophagus consisting of two dis- tinct parts. Tail with markedly projecting lateral processes at the end. Male. Length 3.2—4.1 mm, width 0.092—0.11 mm. Tail curved ven- trally. Buccal capsule 0.048—0.051 mm long and 0.032—0.038 mm wide. Length of esophagus 0.49—0.62 mm, muscular anterior part 0.23—0.29 mm long, glandular posterior part 0.26—0.33 mm. Nerve ring situated 0.12— 0.13 mm, excretory pore 0.24—0.27 mm from the cephalic end. Tail with narrow caudal wings, with 12 pairs of long, riblike papillae. The eight anterior pairs are widely spaced. Seven pairs of preanal papillae, 152 161 one pair of adanal papillae, and a group of 4 postanal pairs, the second pair the longest. Only the right spicule present, 0.25—0.27 mm long, its distal end markedly thinner. Gubernaculum slightly Y-shaped, the two proximal parts of different length: longer part 0.036—0.04 mm long, shorter part 0.044—0.05 mm, Female. Length 5.0—7.8 mm, width 0.13—0.15 mm. Buccal capsule 0.054—0.064 mm long and 0.04 mm wide. Length of esophagus 0.67—0.88 mm, muscular anterior part 0.3—0.4 mm long, glandular posterior part 0.37— 0.48 mm. Nerve ring situated 0.15—0.16 mm, excretory pore 0.2—0.29 mm from the cephalic end. Vulva situated behind middle of body, 2.46—3.52 mm from end of tail which is 0.09—0.112 mm long. First-stage larva taken from a female 0.5—0.54 mm long and 0.026—0.028 mm wide. Cuticle transversely striated. A spinelike process situated laterally at the cephalic end. Buccal capsule rudimentary, 0.0043 mm wide. Length of esophagus 0.08—0.084 mm; Nerve ring situated 0.043 mm from the cephalic end. Anus situated 0.2—0.25 mm from end of tail. Third-stage larva 3.08 mm long. Cephalic end conical, buccal capsule rudimentary, esophagus simple, club-shaped,0.1 mm long. Tail 0.01 mm long. Fourth-stage larva (female). Length 3.7 mm, width 0.8 mm. Buccal capsule 0.044 mm long and 0.044 mm wide. Length of esophagus 0.49 mm. Nerve ring situated 0.12 mm, excretory pore 0.17 mm from the cephalic end. Vulva situated 1.42 mm from end of tail. Vagina muscular,0.56 mm long. Tail 0.045 mm long. Fifth-stage larva (female). Length 5.135 mm, width 0.086 mm. Buccal capsule 0.056 mm long and 0.052 mm wide. Length of esophagus 0.62 mm, muscular anterior part 0.27 mm long, glandular posterior part 1620.35 mm. Nerve ring situated 0.14 mm, excretory pore 0.3 mm from the 163 cephalic end. Vulva situated 2.185 mm from end of tail; uterus with a few eggs. Tail 0.05 mm long. Reference: Pande, Bhatia, and Rai, 1963b, pp.105—118. Procamallanus mehrii Agarwal, 1930 (Figure 95) Host: Wallago attu. Localization: mainly posterior part of body cavity; one specimen found in the swim bladder. Distribution: India. Description (after Baylis, 1939). Cuticular striation with intervals of 0.0036 mm. Buccal capsule of different form in the two sexes. In the male it is cuplike; in the female it is more oblong and barrel-shaped. Its walls without spiral thickenings. At the anterior end of the buccal cavity there are 4 chitinized platelike structures, the two median structures mar- kedly larger. They are weakly separated and covered laterally with groups of filaments which are processes of the wall of the capsule. Together with those of the median plates, there are 9—10 filaments in the maleand 12 in the female; nearer to the outer plates there are 6—7 filaments in the male and 10 in the female. Their form is also different in the two sexes: they are more or less conical in the male, oblong and more or less spindle- shaped in thefemale (Agarwal, 1930). 153 164. FIGURE 95. Procamallanus mehrii Agarwal, 1930: a — cephalic end of male; b — cephalic end of female; c — anterior end of female, lateral; d — chitinous plate with 12 leaflike processes; e — caudal end of female, lateral; f — cephalic end; р — region of vulva, lateral (a,b — after Agarwal, 1930; с--р— after Khera, 1955). Male. Length 16—19 mm, width 0.32—0.36 mm. Depth and width of buccal capsule 0.17 mm. Anterior part of esophagus 0.92—1.14 mm long, posterior part 0.8—1.2 mm. Nerve ring situated about 0.27—0.32 mm, excretory pore 0.48—0.5 mm from the cephalic end. Tail about 0.1 mm long. Caudal wings 0.94—1.2 mm long. Spicules of the same length, 0.285 mm. Female. Length 33 mm, width 0.425 mm, Length of buccal capsule 0.35 mm, width 0.2 mm. Anterior part of esophagus 1.61 mm long, posterior part 1.41 mm. Nerve ring situated 0.53 mm, excretory pore 0.96 mm from the cephalic end. Tail 0.23 mm long. Vulva situated about 10 mm from posterior end. The cuticle is thickened near the vulva and forms а ''slitlike '' swelling ventrally in front of the vulva and a larger swelling on the dorsal surface of the body. Description (after Khera, 1955). Male not described. Female. Length 40 mm, width 0.51 mm, Cuticle striated with intervals of 0.0035—0.006 mm. Buccal capsule chitinized, barrel-shaped, 0.355 mm long and 0.2 mm wide; its walls without spiral thickenings. At the anterior end of the buccal capsule are 4 chitinized platelike formations of equal size with 12 leaflike processes. Esophagus divided into a club-shaped muscular anterior part which is longer than the cylindrical glandular posterior part. Length of muscular part 1.71 mm, of glandular part 1.5 mm. Intestine wide, with large lumen, Rectum short and narrow,0.12 mm long. Three large glandular cells at the connection between intestine and rectum. Nerve ring and cervical papillae situated at the level of the anterior part of the 154 esophagus, 0.54 and 0.93 mm, respectively, from the cephalic end. Cervical papillae large and projecting. Excretory pore situated 1.2 mm from the cephalic end. Tail finger-shaped, 0.238 mm long. Vulva situated in posterior half of body, 14.8 mm from end of tail. Cuticle thickened ventrally just before the vulva. The large thickening on the dorsal surface described by Agarwal (1930) is absent. Vagina muscular, curved, then extending straight poste- riorly for 1.4mm. Common duct of uterus short, dividing intotwo branches, one extending anteriorly, the other posteriorly. The anterior branch of the uterus ends in an ovary, 2.1 mm before the posterior end of the esophagus. Posterior branch of uterus ending blind, 1.5 mm before the anus. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Agarwal, 1930, pp.59—64; Baylis, 1939, рр. 218— 219; Khera, 1955, pp. 243—244. Procamallanus muelleri Agrawal, 1966 (Figure 96) Host: Heteropneustes fossilis. Localization: stomach, Distribution: India. Description (after Agrawal, 1966b). Males and females of slightly different size. Mouth surrounded by 2 lateral and 4 submedian papillae. Buccal capsule barrel-shaped, smooth, without crests or groups of leaves. Basal part of capsule thickened, forming a circular connection with the esophagus. Cuticle more strongly striated in females than in males. In- tervals of striation 0.012—0.015 mm in males and 0.015—0.018 mm in females. Male. Length 4.64 mm, width 0.11 mm. Width of head 0.05 mm. Buccal capsule 0.07 mm long and 0.04 mm wide. Esophagus divided into a club- shaped muscular anterior part 0.27 mm long and 0.04 mm wide and a gland- ular posterior part 0.24 mm long and 0.03 mm wide. Length of esophagus 0.51 ата. Nerve ring situated 0.12 mm, excretory роге 0.18 mm from the cephalic end. Tail with rounded end, 0.13—0.22 mm long. Caudal end curved ventrally, with one spiral coil. Caudal wings well de- veloped, extending to end of tail, 0.35—0.51 mm long. Fourteen to 18 pairs of anal papillae, eight or nine pairs preanal, one pair adanal, and five to seven pairs postanal. The preanal papillae are pedunculate, almost equally spaced. The single pair of pedunculate adanal papillae is situated near the anus. Two pairs of small sessile papillae surround the anus; 3 to 5 pairs of the postanal papillae are long, pedunculate, grouped together; the papillae of the posterior pair are small, sessile, isolated. Phasmids present at end 16601 tail. Spicules tubular, of the same length and similar in form, 1.40— 1.41 mm long, widening at the anterior end and tapering posteriorly. Gubernaculum absent. Female. Length 3.56—6.54 mm, width 0.11—0.17 mm. Width of head 0.05—0.06 mm. Buccal capsule 0.057—0.07 mm long, 0.04—0.05 mm wide. Length of muscular anterior part of esophagus 0.28—0.31 mm, width 0.04—0.06 mm, glandular posterior part 0.31—0.39 mm long, 0.05—0.07 mm wide. Length of esophagus 0.59—0.69 mm. Nerve ring situated 0.13— 0.16 mm, excretory pore 0.17—0.21 from the cephalic end. Tail short, 165 155 rounded, ending in two small spinelike processes; tail 0.10—0.15 mm long. Vulva situated behind middle of body, 1.98—3.61 mm from the cephalic end. Uterus filled with larvae and eggs. (165) 0.? mm Е Е a Е Е = ce =\— CAA = 9 SS ий о FIGURE 96. Procamallanus muelleri Agrawal, 1966: a,b — anterior end, lateral; c, d — caudal end of male, lateral; e — caudal end of female, lateral (after Agrawal, 1966b). Reference: Agrawal, 1966b, рр. 204—208. Procamallanus planoratus Kulkarni, 1935 (Figure 97) Host: Claritas batrachus:. Localization: intestine. Distrib wtton ? “dis: Description (after Kulkarni, 1935), Cephalic end narrowing, rounded. Cuticle smooth, without striation. Buccal capsule 0.082 mm long and 0.061 mm wide, with a smooth, chitinized lining without leaves or crests. Two club- shaped structures at the base of the buccal capsule, Length of esophagus 0.41 mm. Nerve ring situated 0.24 mm, excretory роге 0.31 mm from the cephalic end. 156 io) Л с FIGURE 97. Procamallanus planoratus Kulkarni, 1935: a — anterior end; b — region of vulva, lateral; c,d — caudal end of male, lateral; e — caudal end of female (after Kulkarni, 1935). Male. Length 3.85 mm, width 0.061 mm. Tail slightly curved ventrally, with bluntly pointed end. Caudal wings well developed. Seven pairs of pre- anal and one pair of postanal pedunculate papillae. The stalks of the pre- anal papillae are longer than those of the postanal papillae. Spicules of the same length, 0.12 mm. Gubernaculum absent. Female. Length 5.07—7.26 mm, width 0.079 mm. Tail bluntly pointed, 0.11 mm long. Vulva situated in about middle of body, 1.4—3.38 mm before the caudal end. A distinct lobelike structure forms the anterior lip of the vulva. Vagina directed posteriorly. Description (after Baylis, 1939). Cuticle without striation. Buccal capsule 0.082 mm long and 0.061mm wide, without crests or leaves. Basal ring of buccal apparatus represented by two chitinoid formations of spine- like form. Esophagus, measured from the cephalic end, 0.41 mm long. Nerve ring situated 0.24 mm, excretory pore 0.31 mm from cephalic end. Male. Length 3.85 mm, maximum width 0.061 mm. Caudal end moderately curved ventrally with rounded end. Eight pairs of pedunculate papillae, seven pairs preanal and one pair postanal. Spicules of the same lemma, Ogi7 iaawaaie Female. Length 5.0—7.3 mm, maximum width 0.079 mm. Vulva situated 1.4—3.38 mm from end of tail. Its anterior lip forms a projecting lobe. Tail 0.11 mm, long with blunt end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Baylis, 1939, pp.219—220; Kulkarni, 1935, рр. 29—32; Yamaguti, 1961a, p. 45. 157 Procamallanus sigani Yamaguti, 1935 (Figure 98) Host: Siganus fuscéestens. Localization: small intestine. DistPibution: Papan, Description (after Yamaguti, 1935). Cuticle 0.006 mm thick, finely transversely striated. Buccal capsule with smooth outer and inner surface, with a distinct cuticular ring at the base. Male. Length 10—14 mm. Length of buccal capsule 0.093—0.105 mm, width 0.030—0.114 mm. Nerve ring situated 0.22—0.26 mm from cephalic end. Muscular anterior part of esophagus 0.32—0.41 mm long, 0.07—0.08mm wide; glandular posterior part 0.58—0.68 mm long, 0.066—0.088 mm wide. \\s 42 mm FIGURE 98. Procamallanus sigani Yamaguti, 1935: a — anterior end of female, lateral; b — posterior end of male, lateral (after Yamaguti, 1935). Testis beginning 1.8—2.7 mm from the cephalic end, widening posteriorly until it narrows abruptly at the connection with the relatively short, curved vas deferens. Seminal vesicle about 4 mm long, markedly narrower at both 168 ends. The small ejaculatory duct is lined with high cylindrical epithelium, about 1.5 mm long. The long spicule is distinctly curved, about 0.4 mm long. Short spicule 0.12—0.15 mm long. Caudal wings connected anteriorly and supported on each side by 3 long, thin preanal and 3 shorter postanal papillae. Behind the anus are 3 further outer papillae on each side: one inside and two 158 outside the caudal wings. Many subcuticular muscle fibers present in the preanal region of the caudal wings. Tail 0.15—0.20 mm long. Female. Length19—27 mm, maximum width 0.55 mm. Length of buc- cal capsule 0.11—0.12 mm, width 0.11—0.138 mm. Nerve ring situated 0.28—0.33 from cephalic end. Muscular anterior part of esophagus 0.40—0.51 mm long and 0.100— 0.135 mm wide, glandular posterior part 0.63—0.91 mm long and 0.11— 0.13 mm wide. The anterior end of the ovary is situated at different levels in different specimens. In a mature specimen 21.5 mm long it is situated 1.34 mm behind the vulva, in others at the same level or anterior to it. In a specimen 26 mm long it is situated 4.1 mm from the cephalic end. Oviduct forming a figure 8. The simple, tubular uterus begins 1.4—2.4 mm from the cephalic end and ends 0.75—2.8 mm from the posterior end ina vermiform appendage which is directed posteriorly or anteriorly. Vagina muscular, 0.75—1.2 mm long, straight, sometimes curved in the proximal part. Vulva situated 7.3—11.3 mm from the cephalic end, dividing the body at the ratio of 1:1.3—1.75; its anterior lip about 0.060 mm long with a thin cuticular membrane, the teatlike posterior lip is 0.018 mm long and 0.030 mm wide. Fully developed larvae in the uterus with a thin tail, 0.52 mm long and 0.015 mm wide. Length of tail of female 0.30—0.38 mm, with pointed end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.34; Yamaguti, 1935, pp.378—380; 1961b, р. 45. Procamallanus slomei Southwell and Kirschner, 1937 (Figure 99) Host: Xenopus laevis. Localization: stomach. Distributton:s Ateiea: Description (after Southwell and Kirschner, 1937). VWiawhes ensthilio—o2 ит, МЛЯ 0.09—0.15 mm.) Cutrele less strongly striated than in the female. Excretory pore situated in the region of the nerve ring. Mouth with 6 papillae: two lateral and four median, The brownish yellow, barrel-shaped buccal capsule has a hexagonal anterior opening; it is 0.087—0.097 mm long and 0.065—0.075 mm wide. Inner sur- face of buccal capsule smooth. Esophagus divided into a muscular anterior part, swollen posteriorly, 0.2—0.24 mm long; glandular posterior part 0.22—0.29 mm long. Nerve ring surrounding muscular part of esophagus between its anterior and middle third. Testis forming a long, thin, sinuous tube situated in the region of the glandular esophagus. It passes into the long, wide vas deferens, the anterior end of which is widened and forms the seminal vesicle. The vas deferens then narrows and passes into the ejac- 169ulatory duct. Only one spicule, 0.087—0.1 mm long. Caudal wings present. Twelve pairs of caudal papillae: eight pairs preanal, one pair adanal and three pairs postanal. All papillae pedunculate. Tail curved ventrally, with rounded end, 0.06 mm long. Female. Length 2.5—2.9mm, width 0.22mm. Buccal capsule 0.135 mm long and 0.097 mm wide. Muscular part of esophagus 0.26 mm long, glandular part 0.29 mm. Vulva situated before middle of body. Tail with rounded end 159 and with a crown of 8—10 spines. Ovary beginning т the region of the mus- cular part of the esophagus and passing into the oviduct; uterus wide, saclike, double, extending almost to the end of the tail. Walls of uterus wrinkled. Ovejector beginning at the connection of the two uteri, forming a long tube which extends to the vulva and ends in the short vagina, which is reddish brown. Viviparous. Length of larvae 0.25 mm, width 0.014 mm. =, Е] | ep, Е З NS © а FIGURE 99. Procamallanus slomei Southwell and Kirschner, 1937: a — anterior end, lateral; b — caudal end of male, lateral; c — same, ventral; Ч — caudal end of female, showing spines (after Southwell and Kirschner, 1937). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.34; Southwell and Kirschner, 1937, рр. 245—265. -Procamallanus sphaeroconchus Tornquist, 1931 (Figure 100) Hosts 2serranwsspe, reulthyrs “sp: Localization: small intestine. Distributvon? Gull of Suez: Description (after Tornquist, 1931). Red to reddish brown forms in life. The body narrows more markedly in the female than in the male from the anterior part of the posterior part of the esophagus. It then widens slightly again at the buccal capsule. Buccal capsule rounded. Mouth with 6 170 cuticular thickenings inthe form of small rectangles inside the mouth, parallel to its long side. Excretory system in the form of the letter H. Male. Measurements not given. Length of buccal capsule 0.094 mm. Nerve ring situated 0.25 mm from the cephalic end. Anterior part of esoph- agus 0.437—0.484 mm long, posterior part 0.484—0.530 mm long. Posterior end with 7 pairs of subventral preanal papillae. The space between the second and third pairs, counting from the cloaca, is the largest, that between the pairs near the cloaca the smallest. Five pairs of postanal caudal papillae. The second and fourth pairs are subventral and extend onto the caudal wings. The other papillae are lateral. The first pair, counting from the cloaca, is situated slightly before the second, the third slightly 160 before the fourth. The most widely spaced caudal papillae are the second and third pairs. Right spicule 0.320 mm long, left spicule 0.133 mm. Right spicule with small wings, slightly flattened. Female. Length 10.873—21.341 mm, maximum width 0.219—0.200 mm. Depth of buccal capsule 0.109—0.140 mm. Nerve ring situated 0.265— 0.343 mm from the cephalic end. Anterior part of esophagus 0.533—0.640 mm long, posterior part 0.577—0.702 mm. Vulva situated 4.134—8.112 mm from the cephalic end, without lips. Wall of buccal capsule 0.011 mm thick. Wall of basal ring 0.017 mm thick. Vagina 0.100 mm long. Ovejector 1.36 mm long, connected with a ventrally directed part of the uterus, 1.2 mm long. Uterus ending 0.350 mm before the rectum, situated 1.3mm from the cephalic end. Oviduct directed posteriorly 1.55 mm long. Ovary 4.33 mm long. Tail 0.234—0.312 mm long. Viviparous. all FIGURE 100. Procamallanus sphaeroconchus Tornqujst, 1931: a — anterior end; b — caudal end of male, lateral; c — caudal end of female, lateral; а — region of vulva, lateral (after Tornquist, 1931). References: Skrjabin, Shikhobalova, Soboley, Paramonov, and Sudarikov, 1954, р.34; Tornquist, 1931, pp.306—312; Yamaguti, 1961b, p. 46. Procamallanus spiculogubernaculus Agarwal, 1958 (Figure 101) Host: Heteropneustes fossilis. Localization: intestine. Distribution: India. 171 Description (after Agarwal, 1958). Male. Length 3.68 mm, width 0.088 mm; length of buccal capsule 0.06 mm, width 0.036 mm; anterior part of esophagus 0.24 mm long, 0.044 mm wide, posterior part 0.3 mm long, width 0.036 mm wide. ring situated 0.12 mm from cephalic end. Tail blunt, conical. Seven pairs Nerve 161 of caudal papillae, four pairs preanal and three pairs postanal, Right spicule 0.344 mm long, left spicule 0.05 mm. Gubernaculum 0.05 mm long, incom- pletely fused with the smaller spicule. Caudal wings absent. FIGURE 101. Procamallanus spiculogubernaculus Agarwal, 1958. Caudal end of male, lateral (after Agarwal, 1958). Female. Length 4.84 mm, width 0.088 mm; length of buccal capsule 0.08 mm, width 0.048 mm; anterior part of esophagus 0.32 mm long, 0.06 mm wide, posterior part 0.046 mm long,0.048 mm wide. Rectum 0.05 mm long. Nerve ring situated 0.142 mm, excretory pore 0.232 mm from cephalic end. Тай 0.1mm long, tapering, pointed, with bifid end. Vulva situated behind middle of body, 2.64 mm from the cephalic end, distinctly raised above the surface. Length of vagina 0.608 mm. Reference: Agarwal, 1958, рр. 348—349. Procamallanus sp. Ali, 1956 Host: Ophiacephalus pachua, Localization: stomach. Distributron: Тата: Description (after Ali, 1956). Male unknown. Female. Length 5.542 mm, maximum width 0.23 mm. Buccal capsule barrel-shaped, 0.061 mm long and 0.085 mm wide. Nerve ring situated 0.13 mm, excretory pore 0.22 mm from the cephalic end. Esophagus 0.475 mm long, i.e. 1/12 of length of body. Muscular part of esophagus 0.186 mm long, glandular part 0.289 mm. Vulva situated in middle of body, 2.5 mm from cephalic end. Tail 0.065 mm long. Two appendages at the end of the tail. Buccal capsule with a spiral thickening which does not extend to its anterior end. References: Ali, 1956, pp. 15—16; Yamagutr,” 1961, бр: 172 Procamallanus sp. Vuylsteke, 1964 (Figure 102) Hos te tish., Localization; intestine. 162 FIGURE 102. Procamallanus sp. Vuylsteke, 1964: а — anterior end; b — cephalic end (after Vuylsteke, 1964). Ето: Ара“ Description (after Vuylsteke, 1964). One specimen in bad condition. Transverse striation of cuticle with intervals of 0.012 mm at the posterior end of the esophagus. Length of esophagus 0.490 mm. Reference: Vuylsteke, 1964, рр. 64—66. Procamallanus sp. Mawson, 1957 (Figure 103) Host: Sigmanum nebulosus. Localization: intestine. Distribution: Australia. Description (after Mawson, 1957b). Male unknown. FIGURE 103. Procamallanus sp. Mawson, 1957 Anterior end of female, lateral (after Mawson, 1957b). 163 173 Female. Length19 mm. Buccal capsule 0.12 mm 1оп8, 0.11 mm wide, including the thickness of the walls (0.015 mm). Excretory pore situated 0.4 mm from the cephalic end. Anterior part of esophagus 0.43 mm long, posterior part 0.75 mm. Tail conical, 0.28 mm long, with simple, rounded end. Vulva not raised above the surface, situated almost a third of the length of the body from the cephalic end. Uterus filled with spirally coiled larvae 0.45 mm long, with a long tail. Reference: Mawson, 1957b, p.178. Genus Spirocamallanus Olsen, 1952 Diagnosis. Procamallaninae. Buccal capsule not divided into lateral valves; inner wall of capsule with spiral thickenings which are either con- tinuous or consist of interrupted ridges. Esophagus divided into a muscular anterior andaglandular posterior part; posterior part usually longer. Caudal wings present in males; to 14 pairs of caudal papillae. Gubernaculum absent. Two spicules, of the same or of different length, of similar or dif- ferent form. Numerous spine-shaped processes present or absent on the tail of females. Vulva situated in posterior half of body. Viviparous. Parasites of fish and amphibians. Type species: Spirocamallanus spiralis (Baylis, 1923) Olsen, 1952. Spirocamallanus spiralis (Baylis, 1923) Olsen, 1952 (Figure 104) Synonym: Procamallanus spiralis Baylis, 1923 Hosts: Heterobranchus anguillaris, Synodontts ewpterus,. Localization: stomach and intestine. Distribution.) Мос Ареса: Historical review The description of the species was given intwo works (1923a,b): inthe first Baylis described a male and inthe second males andafemale. Tornquist (1931) gave a description of the species based partly on Baylis! material (from the se- cond host) and partly on collections from Sparidae and Labridae from the Gulf of Suez. This material was caught by the expedition led by Jagerskiold (1909). In Volume IV of the ''Key to Parasitic Nematodes '(1954)a list of the hosts of P. s pi- ralis is givenaccording to the above two authors, but the locality is mentioned only for Jagerskidld's collection. A year before Volume IV of the "Кеу" was published, a work by Winter (1953) appeared which was not known to the authors of the above survey. Winter recorded the species (which he placed in the genus Spirocamallanus, in agreement with Olsen, 1952), in marine fishes caught near the coast of Mexico. He gives a description (based on females), stating that the Mexican specimens are identical in all important characters with Spirocamallanus spiralis (Baylis, 1923) Olsen, 1952. The identification is based on the structure of the buccal capsule and of the anterior and posterior end of the body. The species 164 174 is widely distributed, occurring in fish which are quite unrelated systemat- ically and in their ecology (Winter, 1953, р. 138). 0 0.5 mm FIGURE 104. Spirocamallanus spiralis (Baylis, 1928). Cephalic end, lateral (after Baylis, 1928b). Campana-Rouget (1961a) mentioned that the number of species of Spiro- camallanus parasitizing in marine fishes is very small compared with the number of species in freshwater fish; there are only a few species. It is unlikely that the same species is found in hosts with a different biology. Tornquist's description does not exactly agree with the description of Baylis: the buccal capsule contains 9 spiral ribs instead of 12, the proportions of the two parts of the esophagus are different, and the males are markedly larger. Campana-Rouget therefore proposed to remove Tornquist's specimens from S. spiralis and named them Spirocamallanus tornquisti (=Pro- camallanus spiralis sensu Tornquist, 1931). Winter's specimens from Mexican fish also belong to S. tornquisti. Description (after Baylis, 1923b). Buccal capsule not divided into two lateral valves. Tridents absent. Inner side of buccal capsule with spirally arranged riblike thickenings. Cephalic end slightly laterally com- pressed. Male. Length 7 mm, maximum width 0.16 mm. Length of buccal capsule 0.07 mm. Distance from cephalic end to end of muscular anterior part of esophagus 0.45 mm; length of esophagus 0.78 mm. Right spicule 0.15 mm long, left spicule about 0.10 mm, Seven pairs of riblike papillae in the caudal wings, 2 pairs of short pe- dunculate papillae situated ventrally, and one pair at the anterior lip and one pair at the posterior lip of the cloaca. There are apparently some further small papillae near the end of the tail. Female. Length 22.25mm, maximum width 0.40mm. Length of buccal capsule 0.09 mm. Distance from anterior end of body to nerve ring 0.27 mm, to end of muscular part of esophagus 0.58 mm, to end of glandular part1.10 mm, and to vulva 9.25 mm. Tail 0.16 mm long. Only one ovary; posterior branch of uterus ending blind. Vagina extending straight posteriorly from the vulva. Viviparous. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.34; Baylis, 1923a, pp.27—28; 1923b, pp.137—139: Campana-Rouget. 1961, pp.24—25; Jagerskidld, 1909, p.66; Tornquist, 1931, p.400; Winter, 1953, pp.137—139; Yamaguti, 1961b, p. 46. 165 Spirocamallanus amarali (Vaz and Pereira, 1934) Olsen, 1952 (Figure 105) Synonym: Procamallanus amarali Vaz and Pereira, 1934 Host: Leporinus sp. Localization: small intestine. Distribution’ Braz Description (after Vaz and Pereira, 1934). Male (one specimen), Length 9.25 mm, width 0.28 mm. Mouth rounded, buccal capsule large, more or less spherical, with strongly chitinized walls and with 8 chitinized ribs. A chitinous ring between buccal capsule and muscular part of esophagus. Buccal capsule 0.07 mm long and 0.062 mm wide. Muscular part of esophagus club-shaped, gradually widening posterior- ly, 0.31 mm long. It is followed by a ventricle about 0.6 mm long. Four papillae around the mouth, two median and two lateral. Nerve ring surrounding anterior part of esophagus slightly behind its middle,0.2 mm from the cephalic end. Posterior end curved ventrally, with narrow caudal 175 wings. Spicules of different length, strongly chitinized, tapering distally; larger spicule 0.44 mm long, smaller spicule 0.24 mm. Fourteen pairs of caudal papillae: 8 pairs preanal,3 pairs adanal, and 3 pairs postanal, the last two postanal pairs situated immediately behind the anus and the other pair near theend of the tail. Anus situated 0.14 mm from the posterior end. FIGURE 105. Spirocamallanus spiralis (Vaz and Pereira, 1934): a,b — anterior end, different aspects; c — caudal end of male, ventral (after Vaz and Pereira, 1934). Female not known, References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.30; Vaz and Pereira, 1934, рр. 96—97. Spirocamallanus bagarii (Karve and Naik, 1951) Olsen, 1952 (Figure 106) Synonym: Procamallanus bagarii Karve and Naik, 1951 166 Hosts Эазагти-, Бабаев! 05 (— Bapammncmyarr el 11), Са - liehrous bimaculatwus. hocalizat loam: stomach: Distribution: India. Description (after Karve and Naik, 1951). Striation of cuticle distinct, annulation of body coarse, more distinct in the anterior half. Ce- phalic end truncate and slightly wider than the other part of the body. Buc- cal capsule identical in both sexes, strongly chitinized, wider than long, usually cup-shaped but more or less spherical in some specimens, Pro- jecting, interrupted ribs on the inner surface of the capsule. At the bottom of the capsule are three cuticular folded processes directed anteriorly, one dorsal and two subventral. They correspond to the three sectors of the esophagus. Mouth large, elliptical or rounded, surrounded by three pairs of cephalic papillae. Muscular anterior part of esophagus club-shaped posteriorly. The esophagus opens into the intestine through a triradial opening which is bordered by three valves that project into the intestine. Excretory pore large, situated slightly behind the connection of the two parts of the esophagus. Cervical papillae situated at the level of the nerve ring or slightly before it. 176 Male. Length 4.95—9.5 mm, width 0.15—0.25 mm. Transverse striation of cuticle with intervals of 0.0025 mm. Width of cephalic end 0.13—0.17 mm. Buccal capsule 0.092—0.112 mm long and 0.11—0.135 mm wide. Muscular anterior part of esophagus 0.48—0.7 mm long, glandular posterior part 0.49—0.77 mm. Nerve ring surrounding anterior part of esophagus 0.195— 177 0.265 mm from the cephalic end. Cervical papillae situated 0.18—0.25 mm from the cephalic end. Caudal region markedly thicker than preceding part of body and strongly curved ventrally. Caudal wings and parallel caudal muscles well developed. Tail 0.055—0.07 mm long. Seven pairs of long, pedunculate, riblike preanal and at least 6 pairs of postanal pairs of papillae. Counted from the posterior end, the first pair is lateral, the second subdorsal, and the third, fourth, and sixth pairs are situated close together. Two spicules of different length but difficult to see, particularly the left. Right spicule thicker, 0.15— 0.165 mm long. Left spicule weakly chitinized, thin, 0.13—0.145 mm long. Both spicules with thin end. Gubernaculum chitinized, 0.033—0.0475 mm long. Female. Length 14.1—16.8 mm, width 0.3—0.35 mm. Transverse striation of cuticle with intervals of 0.003 mm. Width of cephalic end 0.16—0.19 mm. Buccal capsule 0.11—0.13 mm long and 0.135—0.17 mm wide. Muscular anterior part of esophagus 0.83—0.95 mm long, glandular posterior part 0.71—0.92 mm. Nerve ring surrounding anterior part of esophagus 0.26—0.3 mm from the cephalic end, cervical papillae situated 0.255— 0.275 mm from the cephalic end. Tail 0.11—0.15 mm long, with three spines at the end. A single pair of caudal papillae, situated about 0.055 mm from end of tail. Vulva situated 7.85—9.6 mm from cephalic end, passing into a small, oblique, tubular part which continues posteriorly into the muscular vagina which is 0.55—0.61 mm long. The vagina extends posteriorly for some distance, parallel to the ventral body wall and then diverges, runs paral- lel to the dorsal wall for a short distance and then unites with the two op- posite branches of the uterus. The muscular vagina is surrounded by uni- cellular glands at its beginning. Cuticle swollen in the region of the vulva. Viviparous, 167 (176) FIGURE 106. Spirocamallanus bagarii (Karve and Naik, 1951): a — anterior end of female, dorsal; b, c — caudal end of male, lateral; d — same, ventral; e — region of vulva; f— tail of female, lateral (after Karve and Naik, 1951). Reference: Karve and Naik, 1951, pp. 30—34. Spirocamallanus barroslimai (Pereira, 1935) Olsen, 1952 (Figure 107) Synonym: Procamallanus barroslimai Pereira, 1935 Boosting BATE Les Localization: small intestine. DiS) tr, Uta. Onn ear az ails Description (after Pereira, 1935). Small reddish nematodes. 168 Male. Length 3.1 mm, width 0.14 mm; cuticle with fine transverse striation. Mouth rounded, buccal capsule large, almost spherical, with about 20 chitinized lines which are wider at the base and become gradually narrower anteriorly and to theleft, most of them interrupted. They form the wall of the buccal capsule, which is 0.03 mm long and 0.04 mm wide. Esophagus club-shaped, 0.28 mm long, maximum width 0.07 mm; glandular part about 0.5 mm long. Caudal end forming a spiral, tapering; caudal wings 178 weakly developed, beginning 0.15 mm from posterior end. Tail 0.10 mm long. The ventral side of the tail shows distinct, oblique muscles and 6 pairs of subventral papillae which form two rows, three pairs preanal and three pairs postanal. Spicules small, almost of the same length,0.05 and 0.06 mm. 27mm jets OCF mm И а b FIGURE 107. Spirocamallanus barroslimai Pereira, 19365: a — anterior end; b — posterior end of male, lateral (after Pereira, 1935). Female unknown. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.30; Pereira, 1935, pp.60—61. Spirocamallanus cearensis (Pereira, Vianna Diaz, and Azevedo, 1936) Olsen, 1952 (Figure 108) Synonym: Procamallanus cearensis Pereira, Vianna Diaz, and Azevedo, 1936 Hosts: definitive —Astyanax bimaculatus vittatus; inter- mediate — Diaptomus cearensis and D. azevedoi; reservoir — Curimatus elegans. Lotalyzation: wsmall intestine. Dis ri butt om: MBrazal. Description (after Pereira, Vianna Diaz, and Azevedo, 1936). Small, reddish nematodes. Male. Length about 4 mm, width about 0.18mm. Cuticle with moderately distinct transverse striation. Mouth rounded, buccal capsule large, spheri- cal, with 18 chitinized lines which are wider at the base and become thinner anteriorly and toward the left and disappear at the margin of the mouth. Width of buccal capsule about 0.04 mm. Esophagus club-shaped, 0.27 mm long. Between esophagus and buccal capsule is a vestibulum intheform ofa chitinized ring on which the 0.008-mm-long buccal capsule is situated. 169 Glandular part of esophagus about 0.03 mm long; two large nuclei near its posterior end. Nerve ring situated about 0.16 mm, excretory pore about 0.20 mm from the cephalic end. Posterior end curved ventrally, strongly tapering, with very narrow caudal wings. Cloaca situated 0.15 mm from posterior end. The ventral surface shows thin oblique muscles and 7 pairs of papillae arranged in two subventral rows, four pairs preanal and three pairs postanal. Spicules short, chitinized, almost of the same length, about 0.057 mm long. 179 FIGURE 108. Spirocamallanus cearensis (Pereira, Vianna Diaz, and Azevedo, 1936): а — male, general view; b — caudal end of male, ventral; с — larva from uterus of mature female; 4 — first-stage larva from body cavity of Diaptomus; е— second-stage larva from body cavity of Diaptomus cearensis; Г, р — third-stage larvae (after Pereira, Vianna Diaz, and Azevedo, 1936). 170 180 181 Female. Length 10.3—14 mm, width 0.28—0.42 mm. Cuticle with moderately distinct transverse striation. Width of buccal capsule 0.04— 0.05 mm; walls of capsule with 18 chitinized lines. Esophagus 1.75— 1.93 mm long. Nerve ring situated 0.12—0.16 mm, excretory pore 0.21— 0.26 mm from the cephalic end. Anus situated 0.08—0.14 mm from end of tail. Tail markedly tapering. Vulva situated 6.3—6.4 mm from end of tail. Uteri diverging; posterior ovary absent. Viviparous. Development. The adult females parasitize in the small intestine of fish: Astyanax bimaculatus vittatus (Tetragonopterinae; Cha- racididae). Eggs develop into larvae in the uterus. The larvae move in all directions in the uterus and collect in the posterior branch, including the ovejector. The uterus becomes filled with coiled masses or long strings of larvae. The fully developed uterus extends to the posterior end of the esoph- agus anteriorly and to the anus posteriorly. Length of larvae 0.46—0.56 mm, width 0.03 mm, Cuticle with fine trans- verse striation. Cephalic end truncate, mouth simple. There is a distinctly chitinized conical process dorsally at the cephalic end. Esophagus narrow, cylindrical, 0.10—0.11 mm long. Anus situated 0.21—0.22 mm from end of tail. Tail relatively long, about 40—45% of the length of the body, tapering and ending in a thin pointed process. Larvae taken from a female survive in fresh water for many days. When placed in water, the larvae sink slowly, often adhering to the bottom with the tail. Diaptomus cearensis and D. azevedoi were experimentally infected. The larvae of Spirocamallanus penetrated the wall of the intestine and settled in the body cavity without becoming encysted. After about 3 days the first molt takes place. The larvae may remain in the body cavity of the intermediate host for a long time. Diaptomus apparently easily tolerates infection with this parasite. The larvae move freely in the body cavity. We always found only a single larva of Spirocamallanus ina specimen of Diaptomus. Description of larvae considered to be second-stage larvae from the body cavity of Diaptomus. Length 0.66—0.77 mm, width 0.03— 0.04 mm. Buccal capsule of the same structure as in the adult, except that there are only 9—10 chitinized diagonal thickenings. Width of buccal capsule 0.016—0.020 mm, length 0.012 mm. Muscular part of esophagus 0.14 mm long, glandular part 0.12—0.13 mm. Intestine straight, rectum short, straight, anus situated 0.073 mm from end of tail. Tail short, ending in 3 or 4 small denticles. The authors consider this larva infective for fingerlings of Curimatus elegans, the reservoir host. After entering the fish, the larvae settle in the beginning of the small intestine, where the second molt takes place and become third-stage larvae. The genital primordium now develops. In a female larva 1.0 mm long the genital primordium consists of a row (0.04 mm) of cells situated 0.32 mm from end of tail. The oviduct is recognizable in a larva 1.9 mm long; it is situated 0.84 mm from end of tail. The genital primordium has a distinct outline in a female larva 2.4 mm long. The buccal capsule has now the typical striation with 8—11 lines. It is assumed that the definitive host may become infected directly by eating Diaptomus,ji.e. the life cycle of the parasite may be completed 171 182 without a reservoir host. The authors assume that only one molt takes place in the crustacean, although there are probably two molts, but the first is difficult to observe because it is completed soon after the larvae enter the intermediate host, and the differences between first- and second-stage lar- vae are very small. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р.32; Leuckart, 1876, рр. 109—112; Pereira, Vianna Diaz, and Azevedo, 1936, pp. 209—226. Spirocamallanus chimuzensis Freitas and Ibanez, 1968 Host: Pygidium punctulatum. Localization: intestine. Distributvon?~ Рева: Description (after Freitas and Ibanez, 1968). Male. Length 12—13 mm, width 0.24—0.25 mm. Cuticle longitudinally striated, body tapering at both ends. Mouth round, buccal capsule strongly chitinized, its posterior part inthe form ofa ring, capsule 0.078—0.083 mm long and 0.058—0.067 mm wide. Nine spiral ribs (4 complete and 5 incomplete) on the inside of the buccal capsule. Esophagus club-shaped; muscular part 0.40—0.42 mm long, 0.12—0.13 mm wide. Glandular part well developed, 0.57—0.58 mm long and 0.12—0.13 mm wide. Intestine more or less straight. Nerve ring situated 0.18 mm, ex- cretory pore 0.20 mm from the cephalic end. Larger spicule 0.22—0.23 mm, smaller spicule 0.16—0.17 mm long. Gubernaculum absent. Anus situated 0.13 mm from end of tail. Caudal wings absent. Caudal papillae small, 6 pairs, three preanal and three postanal. Genital duct directed anteriorly. Testis more or less sinuous, not extending to the beginning of the intestine. Female. Length 34—37 mm, width 0.63—0.67 mm. Buccal capsule 0.093 mm long and 0.073—0.080 mm wide. Inner surface of buccal capsule with 4 spiral ribs (three complete and one incomplete). Muscular part of esophagus 0.58—0.62 mm long, 0.15—0.17 mm wide; glandular part 0.78— 0.85 mm long, 0.15—0.17 mm wide. Nerve ring situated 0.23—0.25 mm, excretory pore 0.27 mm from the cephalic end. Uteri didelphic, amphidelphic. Lips of vulva not protruding, situated in anterior half of body, 20.26 mm from end of tail. Ovejector directed poste- riorly, 1.46 mmlong. Anterior half of duct situated before the vulva, posterior half behind the vulva. Rectum 0.18 mm long. Anus situated 0.13—0.18 mm from end of tail. Viviparous. Reference: Freitas and Ibanez, рр. 146—148. Spirocamallanus fariasi (Pereira, 1935) Olsen, 1952 (Figure 109) Synonym: Procamallanus fariasi Pereira, 1935 Hosts: Leporinus sp., Pygocentris sp. Localization: small intestine. Юве Бои Bracil: Description (after Pereira, 1935). Small, reddish nematodes. 172 Male. Length 3.5—5.0 mm, width 0.18—0.27 mm. Cuticle with fine transverse striation. Mouth rounded, buccal capsule large, barrel-shaped, with relatively thick walls and with 20 chitinized lines which are wider at the base and become narrower anteriorly and to the left; many of them are interrupted. These formations occupy about the two posterior thirds of the capsule; length of buccal capsule 0.09 mm, width 0.04—0.06 mm. FIGURE 109. Spirocamallanus fariasi (Pereira, 1985): а — cephalic end, lateral; b — caudal end of male, lateral (after Pereira, 1935). Esophagus club-shaped, 0.27—0.31 mm long, maximum width 0.07— 0.09 mm; glandular part 0.5 mm long. Nerve ring situated 0.16—0.17 mm from the cephalic end. Caudal end curved ventrally, tapering, 0.11—0.13 mm long. Spicules small, of almost the same length, smaller spicule 0.08— 0.09 mm long, larger spicule 0.09—0.10 mm long. Ventral side with distinct oblique muscles and 9 pairs of papillae in two subventral rows, three pairs precloacal and six pairs postcloacal. Female unknown. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.32; Pereira, 1935, рр. 61—62. Spirocamallanus fulvidraconis (Li, 1935) Olsen, 1952 (Figures 110, 111) Beton wis), Procamallanws fLulyidraconis Та, 1935; Pro - camallanus siluri Osmanov, 1964 Hosts: definitive — Pseudobagrus fulvidraco, Parasilurus 183 asotus, Liocassis ussuriensis, Leuciscus lehmanni, Barbus brachycephalus, Pelecus cultratus, Aspiolucius esocinus, Glyptosternum reticulatum, Pseudoscaphirhyn- chus kaufmanni; intermediate — Cyclops. Localization: esophagus and intestine. 184 Distribution: China, USSR (Far East, Middle Asia). Historical review Osmanov (1954)described Procamallanus sp. fromnumerous hosts in a study of the helminth fauna of fish in Uzbekistan. Osmanov (1964)named the 173 (183) FIGURE 110. Spirocamallanus fulvidraconis (Li, 1935): a — anterior end of male; b — posterior end of male, lateral; c — anterior end of female; Ч — posterior end of female; e, f — anterior end, lateral; g — cephalic end; h — caudal end of male, lateral; i— caudal end of female, lateral (a—d — after Osmanov, 1964; e—i — after Skryabina, 1969). parasite Procamallanus siluri. Onastudyofaspeciesof Procamal- lanus from Pseudoscaphirhynchus kaufmanni, Skryabina (1969) concludedthat P. siluri Osmanov isidenticalwith Spirocamallanus fulvidraconis <(Li,1935)- Description (after Li, 1935). Cephalic end truncate, mouth rounded, surrounded by 6 small papillae, four submedian and two amphids. Buccal capsule barrel-shaped, its walls with continuous spiral thickenings. Esoph- agus divided into a muscular anterior part and a longer glandular part. Cuticle thick, coarsely striated. Male. Length 3.95—5.35 mm, width 0.05—0.1 mm, Length of buccal capsule 0.04—0.05 mm, width 0.07—0.075 mm. Nerve ring situated 174 0.135—0.160, excretory роге 0.270—0.280 mm from the cephalic end. Muscular part of esophagus 0.320—0.340 mm long, glandular part 0.360— 0.430 mm. ; Spicules of markedly different length and form: right spicule thinner and 0.180—0.240 mm long; left spicule wide, resembling a gubernaculum, 0.025—0.032 mm long. Tail 0.040—0.055 mm long, slightly curved ventrally; the muscles in the curved part are well developed and resemble a pseudo- sucker. Caudal wings symmetrical, supported by 9 pairs of pedunculate papillae. Pemale:) Lensth 9.8—12.25 mm, width 0.15— 0.18 mm. Length of buccal capsule 0.05—0.055 mm, width 0.08—0.09 mm. Nerve ring situated 0.165— 0.175 mm, excretory pore 0.3—0.33 mm from the cephalic end. Muscular part of esophagus 0.4—0.41 mm long, glandular part 0.42—0.53 mm. Vulva situated at about the last fifth of the body, 2.2—2.6 mm from end of tail. Branches of uterus diverging, posterior branch ending blind, without an ovary; viviparous. Tail conical, ending in three short, blunt processes. Tail 0.110—0.120 mm long. Description (after Osmanov, 1964,as Procamallanus siluri). Small, viviparous forms. Mouth without lips. Buccal capsule usually longer than wide in males; this character is variable in females, but the capsule is wider usually than long. Male. Length 3.14—4.8 mm, width 0.084—0.147 mm. Buccal capsule 0.058—0.074 mm long and 0.051—0.066 mm wide. Glandular part of esopha- gus 0.4—0.53 mm long, muscular part0.29—0.34mm. Larger spicule 0.12— 0.168 mm long, smaller spicule 0.025—0.030 mm. Eight or 9 pairs of pre- anal papillae and 5 pairs of postanal papillae. Caudal wings present. Female. Length 5.2—9.5 mm, width 0.109—0.231 mm. Buccal capsule 0.074—0.110 mm long and 0.085—0.131 mm wide. Muscular part of esopha- gus about 0.562 mm long, glandular part 0.804 mm. Vulva situated in posterior half of body. End of tail usually rounded. Description (after Skryabina, 1969). Cuticle thick, striated. Buccal capsule usually as wide as long, but slightly longer than wide in some 185 specimens; spiral thickenings on the capsule not present in all specimens. Esophagus divided into a muscular and a glandular part. Excretory pore situated below the nerve ring. Cervical papillae situated above the connec- tion between muscular and glandular part of esophagus. 186 Male. Length 3.40 (2.80—3.40) mm; width of cephalic end at the buccal capsule 0.068 (0.064—0.070) mm, width in middle of body 0.13 (0.11—0.15)mm. Length of buccal capsule 0.064 (0.064—0.069)mm, width0.052 (0.052—0.060)mm. Nerve ring situated 0.15 (0.15—0.17) mm, excretory pore 0.28 (0.27—0.28 )mm from the cephalic end. Muscular part of esophagus 0.26 (0.26—0.40) mm long, glandular part 0.43 (0.36—0.45) mm. Tail (from the cloaca) 0.052 (0.043— 0.052) mm long, Larger spicule 0.17 (0.11—0.17) mm long, smaller spicule 0.021 (0.021—0.026) mm. Nine pairs of preanal papillae and 5 pairs of post- anal papillae. Small caudal wings present. Tail with processes at the end. Female. Length 5.80 (3.80—5.90) mm, width at cephalic end 0.12 (0.086—0.12)mm, near the vulva 0.13 (0.11—0.15) mm, and near the anus 0.052 (0.047—0.056) mm. Buccal capsule 0.10 (0.077—0.101) mm long, 0.12 (0.077—0.12) mm wide. Nerve ring situated 0.21 (0.13—0.21) mm, excretory pore 0.26 (0.23—0.26) mm, cervical papillae 0.56 mm from the cephalic end. Vulva situated behind middle of body, 3.18 (2.10—3.18) mm from the ce- phalic end. 175 (185) FIGURE 111. Spirocamallanus fulvidraconis (Li, 1935): a — cephalic end of female, apical; b — same, lateral; c — caudal end of male, ventral; d — same, lateral; е — caudal end of female, lateral; f — larva from uterus; р — cephalic end of larva before first molt, buccal capsule not yet developed; h — cephalic end of larva immediately after first molt, buccal capsule already developed, remains of the shed cuticle visible; i — 6-day-old larva with small buccal capsule, simple esophagus, and thin tail; j — 14-day-old larva (5 days after first molt) with well-developed buccal capsule, differentiated esophagus, and with three processes at end of tail (after Li, 1935). Muscular part of esophagus 0.48 (0.38—0.48) mm long, glandular part 0.063 (0.47—0.63)mm. Tail 0.10 (0.064—0.10) mm long. Tail with two processes at the end. Development. Li (1935) found that Spirocamallanus fulvi- draconis develops in the body cavity of Cyclops serratus, C. vicinus, andC. magnus. A larva which enters the water remains at the bottom, winding into a coil and unwinding regularly. Cyclops swallows 176 187 {пе1агуае. Linotedthat many larvae die when swallowed. After they reach the body cavity, the larvae begin to move and remain active until after the molt, when they coil and remain motionless. Li observed only one larval molt in Cycl ops, which he considered as the first, at room temperature 8—9 days after infection, He did not observe further molts during 5 weeks of observation. However, he noted morpho- logical changes after 5—6 days (disappearance of the dorsal tooth, develop- ment of the buccal capsule, increase in the number of genital cells), which are characteristic for the second-stage larva. Li did not observe the first molt, and what he considered as the second stage was in fact the third. Li gives a description of the first-stage larva and the development of its organs. The cephalic end bears a dorsal spine for piercing the intestinal wall of Cyclops. The wall of the esophagus is very thin, and three nuclei of esophageal glands are visible at the base of the esophagus. Cell nuclei, renettes, are situated in the anterior third of the intestine. The intestine contains small granules mainly in the posterior part. The rectal glands are already well developed. The wall of the rectum forms а thin slit. The genital pri- mordium is unicellular and situated behind the middle of the body. Tail long and tapering. Cuticle with distinct transverse striation. The cuticle of a larva which has just passed out of the uterus is thin and coarsely striated. The only changes from the first stage are increase in thickness and in length. After the first molt the cuticle becomes very thick with distinct striation. The buccal capsule changes markedly. In a larva which has just entered the water the cephalic end bears a dorsal spine; this is lost on the 5th day (probably during the first molt, when a small but distinct buccal capsule develops). After 8—9 days, before the second molt, the capsule becomes oblong with a narrow lumen. [f pressure is applied to the cover glass, the inner lining of the capsule is displaced, while the buccal capsule widens to a form resembling that of the adult, but the inner surface is smooth, without thickenings. In a newly hatched larva the nerve ring is situated in about the middle of the esophagus; when the glandular part develops, it is gradually displaced anteriorly until it is situated at the anterior fifth of the esophagus after the second molt. The excretory pore is at first situated behind the middle of the esophagus, During development of the glandular part of the esophagus it is displaced to before the middle of the esophagus. From the pore posteriorly extends a long duct which ends in a large, club-shaped cell with a large nucleus. The nu- cleus of this cell is situated at the anterior third of the intestine in a newly hatched larva, but it moves anteriorly until it is situated in the middle of the posterior part of the esophagus in the mature larva. Changes in the esophagus are marked during development. In the newly hatched larva the wall of the esophagus is very thin, particularly near its anterior end, with a wide lumen. After 2 or 3 days inthe Cyclops, the wall of the esophagus becomes thicker and the lumen narrower. The division of the esophagus into two parts takes place at the second molt, and there is a distinct muscular anterior and a glandular posterior part. The esophagus increases to 3 times its original length in 3 weeks. 177 The cells of the intestine become larger and more numerous and contain pigment granules. Thecontents of the intestine of the newly hatched larva are small granules; after 2 or 3 days in the body cavity of Cyclops, the intestine becomes filled with yellow particles of irregular form. These particles increase in size but are discharged during the second molt or just after it, and the intestine contains then little or practically no solid matter. The reddish brown color of the intestine after the second molt is due to pigment granules concentrated around the nuclei of the cells of the intestine. In a newly hatched larva the genital primordium is unicellular and 188 situated behind the middle of the body. After 6 days (after the molt) it consists of 4 cells, after 14 days of more than 10 cells, and from then on it develops little. The long, thin tail gradually decreases in size during growth, leaving an empty sheath. Before the second molt, a short, trifid tail, resembling that of the adult female, develops inside the old cuticle. Larval growth in the body cavity of Cyclops (mm) (after Li, 1935) Number of days Length 1.015 Width 0.044 Distance of nerve ring from cephalic end 0.088 Distance of excretory pore from cephalic end 0.146 Length of esophagus 0.320 (2 parts) | (2 parts) | (2 parts) Length of intestine 0.400 0.422 0.596 Length of tail 0.045 0.055 0.055 References: Ivashkin and Khromova, 1964, рр. 98—104; Osmanov, 1964, p.39; Skryabina, 1969, pp.146—151; Campana-Rouget, 1961, pp..425—433;, Li, 1935, рр, 103—113: .О15еа,. 1952, . pp, 73-215. Spirocamallanus globoconchus (Ali, 1960) (Figure 112) Synonym: Procamallanus globoconchus Ali, 1960 Host: Rita has#etai. Localization: gills. Пизе а butions india. Description (after Ali, 1960). Male. Reddish in life, thin, particularly the anterior part, with trun- cate cephalic end. Posterior end pointed and with wide caudal wings. Length 6.3—7.4 mm, maximum width 0.12—0.14mm. Dorsoventral width of cephalic end 0.08—0.10 mm. Cuticle 0.032 mm thick, transverse striation with inter- vals of 0.013 mm. There is also a very fine striation in the intervals. Nerve 178 ring surrounding anterior part of esophagus 0.23—0.28 mm from the cephalic end; excretory pore situated 0.30—0.36 mm from the cephalic end. Buccal capsule nearly spherical, almost as wide as long (0.14—0.15 mm long and 0.12—0.14 mm wide). There is a characteristic thick-walled, hood- shaped formation at the apex and three shell-like formations and a pair of nodular chitinous formations at the base of the capsule, and several trans- verse linear thickenings on the wall of the capsule. Two pairs of submedian 189 cephalic papillae and a pair of lateral amphids. Esophagus 1.23—1.48 mm long, 1/5 of length of body. Muscular part of esophagus 0.50—0.54 mm long, glandular part 0.73—0.94 mm long. FIGURE 112. Spirocamallanus globoconchus (Ali, 1960): a — anterior end; b — cephalic end; с — caudal end of male, ventral; 4 — spicules; e — ar- rangement of papillae around the cloaca (after Ali, 1960). Tail pointed, with wide, asymmetrical wings. Left wing wider and longer than the right, 1.11 and 0.65 mm long, respectively. Thirteen pairs of pe- dunculate caudal papillae, 8 pairs preanal and 5 pairs postanal. The preanal papillae are equally spaced and larger than the postanal papillae. The first four pairs of postanal papillae form a group; the last pair is isolated at the end of the tail. Three pairs of small, sessile papillae situated around the anus; two of the pairs are larger than the third, which is situated medio- ventrally before the cloaca. The other two pairs are lateral, one before, 179 the other behind the cloaca. Spicules of different length and form. Right spicule longer, more or less needle-shaped, with widened anterior and pointed posterior end. Left spicule boat-shaped and more strongly chitinized. 190 Right spicule 0.15—0.18 mm long, left spicule 0.030—0.038 mm. Ratio of length of spicules about 5:1, Tail 0.032—0.036 mm long. Female unknown. References: Ali, 1960, pp.131—132; Olsen, 1952, pp.173—215. Spirocamallanus gubernaculus (Khera, 1955) (Figure 113) Synonym: Procamallanus gubernaculus Khera, 1955 Host: Hita \eita. Localization: ° intestme, Distribution: India. Description (after Khera, 1955). Threadlike nematodes, tapering at both ends. Cuticle thin, delicately striated with intervals of 0.003—0.004 mm. Mouth surrounded by 6 papillae, two lateral and four submedian. Buccal capsule chitinized, barrel-shaped. Its walls with 16—18 interrupted spiral thickenings. A pair of finger-shaped thickenings parallel to the longitudinal axis of the buccal capsule. Esophagus divided into a club-shaped muscular anterior part and a longer, cylindrical glandular posterior part. Intestine wider than posterior part of esophagus, with wide lumen. Three small rectal gland cells at the connection between intestine and rectum. Nerve ring sur- rounding anterior part of esophagus. Cervical papillae very small. Male. Length 8.5 mm, width 0.109 mm. Length of buccal capsule 0.062 mm, width 0.039 mm. Anterior part of esophagus 0.41 mm long, posterior part 0.645 mm. Nerve ring situated 0.155 mm, cervical papillae 0.39 mm from the cephalic end. Tail tapering, blunt, conical. Caudal wings well developed. Eleven pairs of caudal papillae, 5 pairs preanal and 6 pairs 'postanal. The five pairs of preanal papillae are equally spaced, riblike, pedunculate. The first pair of postanal papillae is ventral, pedunculate. The last two pairs are lateral, sessile, widely spaced. Left spicule absent, right spicule colorless, 0.29 mm long. Dorsal wall of cloaca thickened. Gubernaculum strongly chitinized, knife-shaped, brown to dark brown, with a convexity 0.066 mm long in the middle of the ventral surface. Female. Length 18.25 mm, width 0.141 mm. Length of buccal capsule 0.062 mm, width 0.042 mm. Anterior part of esophagus 0.28 mm long, posterior part 0.925 mm. Rectum 0.07 mm long. Nerve ring situated 0.113 mm, cervical papillae 0.3 mm, excretory pore 0.2 mm from the ce- phalic end. Tail 0.175 mm long,a pair of papillae on the ventral side at the end. Anterior lip of anus directed posteriorly, closing the anus. Lips of vulva forming а small, projecting appendage directed posteriorly, 0.05 mm long. Vulva situated in posterior half of body, 10.25 mm from the cephalic end. The muscular vagina extends to the base of the appendage and then turns posteriorly, extending for 0.4 mm, where it passes into the short com- mon duct of the uterus, which has two branches, an anterior and a posterior. 191The anterior branch ends in an ovary 2.6 mm behind the esophagus. The posterior branch ends blind 1.3 mm before the anus. 180 АА аня мага зака =U FIGURE 113. Spirocamallanus gubernaculus (Khera, 1955): a — anterior end, lateral; b — caudal end of female, lateral; с — caudal end of male, lateral; Я — region of vulva, lateral; e — anterior end of male, lateral; f — caudal end of female, lateral; р — caudal end of male, lateral (a—d — after Khera, 1955; e—g — after Sood, 1968). Description (after Sood, 1968). Cuticle finely transversely striated with intervals of 0.003—0.006 mm. Body with tapering ends. Mouth sur- rounded by 6 papillae, two lateral and four submedian. Buccal capsule barrel-shaped, with spiral thickenings. Esophagus divided into a muscular anterior and a glandular posterior part. Male. Length 4.25—4.31 mm, width 0.09—0.1mm. Dorsoventral width of cephalic end 0.021—0.028 mm. Length of buccal capsule 0.049—0.051 mm, width 0.025—0.038 mm. Muscular anterior part of esophagus 0.27—0.32 mm 192 long, 0.047—0.05 mm wide; glandular posterior part 0.39—0.44 mm long, 0.041— 0.05 mm wide. Nerve ring situated 0.11—0.13 mm, excretory pore 0.16 mm from the cephalic end. Caudal wings well developed. Tail 0.025—0.03 mm long. Twelve pairs of caudal papillae, 7 pairs preanal and 5 pairs postanal. The first pair of post- anal papillae is displaced toward the cloaca; the other four pairs are situated close together. Only right spicule present, weakly chitinized, widened an- teriorly and tapering posteriorly, 0.21—0.22 mm long. Gubernaculum dark, 0.065—0.07 mm long. 181 Female. Length 7.59—9.06 mm, width 0.19—0.20 mm. Dorsoventral width of cephalic end 0.025—0.032 mm. Length of buccal capsule 0.06— 0.07 mm, width 0.03—0.032 mm. Muscular anterior part of esophagus 0.31—0.39 mm long, 0.03—0.05 mm wide; glandular posterior part 0.49— 0.60 mm long, 0.049—0.05 mm wide. Nerve ring situated 0.13—0.145 mm, excretory pore 0.17—0.19 mm from the cephalic end. Tail 0.07—0.081 mm long, with two processes at the end. Vulva situated in posterior half of body, 4.81—5.70 mm from the cephalic end. Viviparous. References: Khera, 1955, pp.245—247; Sood, 1968, рр. 94—96. \ Spirocamallanus hilarii (Vaz and Pereira, 1934) Olsen, 1952 (Figure 114) Synonym: Procamallanus hilarii Vaz and Pereira, 1934 Hosite) Salminu sigvilarnyri: Localization: small intestine. Distzr i bution: Brazil: Description (after Pereira, 1934). Anterior end curved dorsally, whitish, other part of body dark. Cuticle transversely striated. Mouth rounded, buccal capsule with chitinized walls, and with about 16 ribs from one side of the capsule to the other; some of the smaller (second-order) ribs are connected anteriorly. The muscular part of the esophagus is separated from the buccal! capsule by a strongly chitinized ring; it gradually becomes wider posteriorly, about 0.41 mm long and 0.1 mm wide. Glandular part of esophagus very long. (193) FIGURE 114. Spirocamallanus hilarii (Vaz and Pereira, 1934): а — anterior end; b — cephalic end; с — caudal end of female, lateral; d —caudal end of male, ventral; e — same, lateral (after Vaz and Pereira, 1934). 182 Three pairs of small cephalic papillae; one lateral,two median. Nerve ring situated about 0.2 mm, excretory pore 0.36 mm from the cephalic end. Male. Length 6 mm, width 0.2 mm. Buccal capsule 0.057 mm long, 0.057 mm wide. Glandular part of esophagus 1.5 mm long. Tail curved ventrally, with oblique muscle fibers directed to the middle of the ventral surface. Spicules of almost the same length, very small, 0.082 mm and 0.062 mm long; smaller spicule slightly wider at the proximal end. Tail with 8 pairs of widely spaced pedunculate ventral papillae, 3 pairs preanal, 2 pairs adanal, and 3 pairs postanal. Anus situated 0.16 mm from end of tail. Tail tapering sharply behind the last pair of papillae. 193 Female: Length 14mm, width 0.36mm. Buccal capsule 0.062 mm long and 0.062 mm wide. Glandular part of esophagus 2.02 mm long. Viviparous. Body almost entirely filled with embryos. Vulva situated in middle of body. Intestine forming a loop about 0.5mm from the posterior end. Anus situated 0.12 mm from end of tail. Tail blunt, with a spine about 0.028 mm long at the end. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Vaz and Pereira, 1934, pp.97—99. Spirocamallanus hyderabadensis (Ali, 1956) (Figure 115) Synonym: Procamallanus hyderabadensis Ali, 1956 EOS hls); LS сеенонат а, Localization: stomach. Dist but on: india: Description (after Ali, 1956). Body reddish in life, very thin, with slightly narrowing ends, cephalic end truncate, tail pointed in both sexes. Cuticle transversely striated. Intervals of cuticular striation about 0.004mm in middle of body. Cephalic end with one pair of submedian papillae and lateral amphids. Walls of buccal capsule thickened at the base and gradually becoming thinner anteriorly. Buccal capsule nearly cylindrical. Male. Length 3.03—3.13 mm, width 0.077 mm. Excretory pore situated 1940.20 mm, nerve ring 0.066—0.069 mm from the cephalic end. Length of buccal capsule 0.04—0.05 mm, width 0.025—0.030 mm. Muscular part of esophagus 0.24—0.31 mm long, glandular part 0.35—0.42 mm. Length of esophagus 0.59—0.73 mm, about 1/5 of length of body. Testis situated in anterior third of body, extending to base of muscular part of esophagus, where it turns posteriorly. Ejaculatory duct short and narrow. Caudal wings well developed, 0.290—0.329 mm long. Ten pairs of riblike caudal papillae, 6 pairs preanal and 4 pairs postanal. A pair of cuticular processes, characteristic for the genus, situated imme- diately behind the third pair of postanal papillae, Last pair of postanal papil- lae situated near end of tail. Spicules of different length and form, right spicule longer, 0.18—0.20 and 0.042 —0.050 mm long, respectively. Left spicule widest in the middle, with hooklike end and wide proximal part. Right spicule of uniform width most of its length, slightly wider at proximal end and pointed at distal end. Tail 0.068 mm long. 183 FIGURE 115. Spirocamallanus hyderabadensis Ali, 1956: a — cephalic end, apical; b — same, lateral; с — anterior end of female, lateral; d — posterior end of female, lateral; e — caudal end of male, ventral; f — region of vulva, lateral; 5 — spicules, ventral (after Ali, 1956). Female. Length 5.58—6.01 mm, width 0.130 шт. Excretory pore situated 0.26 mm, nerve ring 0.130—0.138 mm from the cephalic end. Buccal capsule 0.073—0.075 mm long and 0.039—0.040 mm wide. Muscular part of esophagus 0.269—0.370 mm long, glandular part 0.370—0.400 mm, Length of esophagus 0.63—0.77 mm, about 1/9 of length of body. One ovary, situated in region of esophagus. Oviduct passing intotheseminalvesicle. Vagina long, extending straight to the vulva, which protrudes and has distinct lips. Pos- terior uterus ending in a blind зас. Viviparous. Tail 0.036 mm long. Reference: Ali, 1956, pp.11—13. 195 Spirocamallanus iheringi (Travassos, Artigas, and Pereira, 1928) Olsen, 1952 (Figure 116) Synonym: Procamallanus iheringi Travassos, Artigas, and Pereira, 1928 | Hosts: Salminus hilarii, Hoplias,sp.y, Tetragonopteras sp., Leporinus sp., Anostomatinae sp. Localization: intestine and pyloric caeca. Distriputions, Brazil. 184 Description (after Travassos, Artigas, and Pereira, 1928). Cuticle with transverse striation. Mouth rounded, buccal capsule large, with a chitinized lining inthe form of spiralribs; 4—5 large ribs in males, 14—16 smaller ribs in females. Anterior end of esophagus with a chitinous ring which separates it from the buccal capsule; esophagus club-shaped, glandu- lar part long. Intestine forming one or two coils. Nerve ring situated 0.20—0.28 mm from the cephalic end, in about middle of esophagus; excre- tory pore situated 0.4 mm from the cephalic end. a у с d f FIGURE 116. Spirocamallanus iheringi (Travassos, Artigas, and Pereira, 1928): a — anterior end; b — cephalic end of male; с — cephalic end of female; d — caudal end of female, lateral; е — caudal end of male, lateral; f — ovejector (after Travassos, Artigas, and Pereira, 1928). Male. Length 12mm, width 0.3mm. Buccal capsule 0.072—0.080 mm long, 0.056—0.064 mm wide. Muscular part of esophagus 0.31—0.37 mm long, 0.08 mm wide; glandular part 0.75 mm long. Tail curved ventrally, with narrow caudal wings which extend 0.5 mm before the anus; oblique muscles on the ventral surface of the caudal end, from the cloaca to the end 196 of the caudal wings. Five pairs of caudal papillae, one pair preanal, one pair adanal, three pairs postanal. Cloaca situated about 0.10—0.12 mm from end of tail; tail conical. Spicules 0.24—0.25 and 0.52—0.53 mm long, re- Spectively. Female. Length 16—21 mm, width 0.5—0.8 mm. Buccal capsule 0.088—0.096 mm long and 0.072—0.080 mm wide. Muscular part of esopha- gus 0.41—0.45 mm long, 0.10—0.11 mm wide; glandular part 0.80—0.84 mm long. Vulva situated in about middle of body; ovejector very long (1.6 mm) and thin (0.065 mm). Branches of uterus diverging. Viviparous. Anus situated 0.16—0.24 mm from end of body, rectum 0.20 mm long. Tail conical, 0.08—0.10 mm long, ending in a finger-shaped process. 185 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Travassos, Artigas, and Pereira, 1928, p.21. Spirocamallanus inopinatus (Travassos, Artigas, and Pereira, 1928) Olsen, 1952 (Figure 117) Synonym: Procamallanus inopinatus Travassos, Artigas, and Pereira, 1928 с Host: Leporinws Эр. Localization: intestine. DIStributron: Вега 2.7 mm res 42mm 2/ шт act 2/ mm ony 22 mm SS) FIGURE 117. Spirocamallanus inopinatus (Travassos, Artigas, and Pereira, 1928): а — anterior end; b — cephalic end of female; с — cephalic end of male; d— caudal end of female, lateral; е — caudal end of male, lateral; f — ovejector (after Travassos, Artigas, and Pereira, 1928). Description (after Travassos, Artigas, and Pereira, 1928). Cuticle 197 striated. Excretory pore situated 0.32 mm, nerve ring 0.25—0.32 mm from cephalic end, around middle of esophagus. Mouth rounded, buccal capsule strongly chitinized; inner surface of capsule with small ribs arranged ina spiral, about 15 ribs in both sexes; the ribs do not extend to the margins of the capsule, occupying about 2/3 of its length and of characteristic appear- ance. Esophagus club-shaped. Male. Length 5.12 mm, width 0.24 пт. Length of buccal capsule 0.09mm. Muscular part of esophagus 0.36 mm long, 0.10 mm wide; gland- ular part 0.56 mm long,0.10 mm wide. Tail curved ventrally, the narrow wings with 8 pairs of papillae, 2 pairs preanal and 6 pairs postanal, the first 186 five pairs situated closer to the end of the tail and close together, the last pair slightly more widely separated. Spicules of almost the same length, about 0.120mm; cloaca situated about 0.20 mm from end of tail which is finger -shaped. Female. Length 16—30 mm, width 0.8 mm. Length of buccal capsule 0.12mm. Muscular part of esophagus 0.45 mm long, 0.13 mm wide; gland- ular part 0.76 mm long,0.16 mm wide. Viviparous. Vulva situated near middle of body, ovejector long and thin, its single part about 1.4mm long, 0.056 mm wide. Rectum about 0.24 mm long; anus situated 0.20 mm from posterior end. Tail conical, blunt. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р.33; Travassos, Artigas, and Pereira, 1928, pp.20—21. Spirocamallanus istiblenni Noble, 1966 (Figure 118) ое ISi MOUS mews vs Glo eh. Loca limarlon:, (intestine: его: \Намат. Description (after Noble, 1966). Mouth round, without lips. In apical view the anterior part of the buccal capsule has 8 lateral processes and is surrounded by weak folds. The orange-brown buccal capsule is lined with 13 or 14 spiral thickenings. Between the buccal capsule and the body wall are 4 elongate buccal sinuses, two dorsal and two ventral. They are apparently part of the osmoregulatory system. Two excretory ducts extend along the entire body and unite at the level of the glandular part of the esophagus, forming an excretory sinus (with a large nucleus), which opens to the outside through a short duct. Posterior part of intestine and rectal gland supported by distinct muscle fibers. Male. Length 14.9 mm, width 0.221 шт. Length of buccal capsule 0.075 mm, width 0.072 mm. Muscular part of esophagus 0.325 mm long, glandular part 0.485 mm. Excretory pore situated 0.400 mm, nerve ring 0.208 mm from cephalic end. Six pairs of preanal and 3 pairs of postanal caudal papillae. Caudal wings curved anteriorly and contiguous ventrally. Ratio of length of spicules 3:2. Right spicule 0.274 mm long, left spicule 0.184 mm. Anus surrounded by 4 199small papillae. Tail finger-shaped, sometimes with three small processes. Tail 0.166 mm long. Female. Length 21.50 mm, width 0.336 mm, Length of buccal capsule 0.077 mm, width 0.077 mm. Muscular part of esophagus 0.397 mm long, glandular part 0.488 mm. Excretory pore situated 0.400, nerve ring 0.220mm from cephalic end. Vulva situated slightly before middle of body, 8.200 mm from cephalic end. Vagina extending posteriorly for about 1.3mm. Distal part of vagina 0.4 mm long, with a thick muscular wall. Posterior ovary usually much shorter than the anterior ovary, probably rudimentary, 0.850 mm long, containing cells resembling immature eggs of the anterior Ovary; eggs in posterior ovary 0.008 X0.010 mm large. The large anterior ovary is coiled into rings and together with the oviduct is about 10 mm long, or half the length of the body; ripe eggs in anterior ovary 0.011 x 0.015 mm large. The oviduct narrows before the seminal vesicle, and the seminal vesicle narrows before the wide, thin-walled uterus, the walls of which con- tain gland cells. Anus situated 0.177 mm from end of tail. 187 (198) 1966: FIGURE 118. Spirocamallanus istiblenni Noble, apical; d — posterior lateral; b — cephalic end, lateral; c — same a — anterior end f — caudal end of male, lateral; e — caudal end of female, end of female, lateral; ventral (after Noble, 1966). g — same, lateral; ‚ pp. 360—366. 1966 Noble, Reference: 188 Spirocamallanus kerri (Pearse, 1933) Olsen, 1952 (Figure 119) Synonym: Procamallanus kerri’ Pearse, 1933 Host: Glossogobius giurus. Localization: intestine. Distribution: Thailand. Description (after Pearse, 1933). Male unknown. Female. Length 5.45 mm, width 0.08 mm. Length of buccal capsule 0.04 mm. Muscular part of esophagus 0.29 mm long, glandular part 0.32 mm. Tail 0.1mm long. Buccal capsule chitinized, barrel-shaped, with thick walls strongly striated on the inner side. Tail narrowed in the middle, curved ventrally, sometimes pointed, with two small spines at the end. Vulva sit- uated between the first and second thirds of the body. 2 = UD Laat ee FIGURE 119. Spirocamallanus kerri (Pearse, 1983): a,b — cephalic end of female; с — caudal end of female, lateral (after Pearse, 1933). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and sudarikov, 1954, p.33; Pearse, 1933, pp.185—186; Olsen, 1952, pp.173— РИ. 200 Spirocamallanus mazabukae Yeh, 1957 (Figure 120) Host: freshwater fish. Localization: intestine. DiS Cig thi tomas Wvnal@z, Description (after Yeh, 1957). Slender nematodes, with finely transversely striated cuticle. Buccal capsule with 12 complete oblique ribs in both sexes. Esophagus divided into a muscular and a glandular part. Male. Length 11.8—14.9 mm, width 0.16—0.18 mm. Cervical papillae situated 0.14—0.18 mm, excretory pore 0.44—0.47 mm from cephalic end. Nerve ring situated in about middle of muscular part of esophagus, 0.22— 0.24 mm from cephalic end. Buccal capsule 0.09 mm long, 0.06 mm wide. 189 Muscular anterior part of esophagus 0.39—0.42 mm long, the longer gland- ular part 0.95—1.05 mm. Cloaca situated 0.15—0.18 mm from end of tail. Caudal wings well developed, supported by pedunculate papillae. The first of the 3 pairs of preanal papillae is separated from the others; the6 pairs of postanal papillae are almost equally spaced. On each side of the cloaca are 2 pairs of large papillae. The longer, thick, right spicule is 0.470— 0.473 mm long, the thinner, less strongly chitinized left spicule has a bifid end and is 0.272—0.283 mm long. Gubernaculum absent. FIGURE 120. Spirocamallanus mazabukae Yeh, 1957: a — anterior end, ventral; b — cephalic end, lateral, showing the buccal capsule; c — caudal end of male, ventral; d — caudal end of female, lateral; e — caudal end of male, lateral, left spicule intact, right spicule torn off (after Yeh, 1957). Female. Length 16.2 mm, maximum width 0.18 mm. Cervical papillae situated 0.16 mm, excretory pore 0.46 mm, nerve ring 0.27 mm from ce- phalic end. Length of buccal capsule 0.10 mm, width 0.075 mm, Muscular part of esophagus 0.46 mm long, glandular part 0.85 mm. Tail 0.19 mm long, markedly narrowing in the middle and tapering at the end. Vulva situated slightly before middle of body, 7.5 mm from cephalic end. Vagina at first directed posteriorly and then turning anteriorly. References: Campana-Rouget, 196la, pp.24—25; Yeh, 195: pp, Пава 128. 201 Spirocamallanus monotaxis Olsen, 1952 (Figure 121) Host: "Monotax is -“erandoculis. Localization: intestine. 190 Distribution: Hawaii. Description (after Olsen, 1952b), Cephalic end smooth, rounded, papillae indistinct. Cuticle thin, transversely striated with intervals of 0.004 mm. Buccal capsule barrel-shaped, with spiral thickenings. A ring- shaped thickening at the base of the capsule. Muscular anterior part of esophagus club-shaped, glandular posterior part also club-shaped. A short valve apparatus between glandular part of esophagus and intestine. Male. Length 15.6—17.7 mm, maximum width 0.26—0.29 mm. Posterior end sharply curved ventrally. Length of buccal capsule 0.11 mm, width 0.07 mm, its walls with 16 spiral thickenings. Anterior part of esophagus 0.39—0.44 mm long, maximum width 0.08—0.09 mm, Nerve ring situated 0.28—0.30 from cephalic end. Excretory pore situated 0.19 mm behind the nerve ring ina male 15.8 mm long. Posterior part of esophagus 0.76 mm long, maximum width 0.11—0.12 mm. Intestine 0.08 mm wide. Cire ЕН ПО АЦ СО u ts OD Ц FIGURE 121. Spirocamallanus monotaxis Olsen, 1952: a — cephalic end, lateral; b — anterior end, lateral; c — region of vulva; d — caudal end of female, lateral; e — caudal end of male, lateral; f —same, ventral (after Olsen, 1952b). In a specimen 17.7 mm long the testis begins 1.46 mm from the cephalic end and extends posteriorly for 7.0 mm. It narrows and passes into the seminal vesicle, which extends posteriorly for 7.1 mm. After a second narrowing, it passes into the ejaculatory duct which is 1.7 mm long. Caudal wings 0.67 mm long, beginning on the ventral surface, where they are united. Seven pairs of pedunculate caudal papillae, 3 pairs preanal, one pair adanal, and 3 pairs postanal. Spicules of similar form but of different length: right spicule 0.29—0.32 mm long, left spicule 0.23—0.24 mm. Tail 0.20 mm long. Female. Length 34.0 mm, maximum width 0.55 mm, Buccal capsule 0.11 mm long and 0.08 mm wide, with 10 spiral thickenings on the walls. 191 Anterior part of esophagus 0.59 mm long, maximum width 0.12 mm. Nerve 202 ring situated 0.37 mm from cephalic end. Posterior part of esophagus 1.11 mm long, maximum width 0.13 mm. Intestine 0.12 mm wide. Vulva situated 13.6 mm from cephalic end, i.e. at about 3/8 of the length of the body. Vagina directed posteriorly, its distal part short (0.07 mm), with thin wall near the vulva, which is followed by a swollen muscular part about 0.72 mm long. Then follows a narrow tube to 3.9 mm behind the vulva. Uterus very wide,filling the body cavity. It is united anteriorly with the ovary, which extends to the esophagus anteriorly and beyond the vulva pos- teriorly. Posterior branch of uterus blind. Embryos inuterus about0.34mm long and 0.018 mm wide. Rectum 0.31 mm long. There are numerous muscle bundles near the anus extending across the body cavity. Tail with small conical end 0.04mm long, with two small, teatlike processes visible only under high magnification, Tail 0.24 mm long. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Suda- rikov, 1954, p.34; Olsen, 1952b, pp.197—198; Yamaguti, 1961b, р. 45. Spirocamallanus murrayensis (Johnston and Mawson, 1940) Olsen, 1952 (Figure 122) Synonym: Procamallanus murrayensis Johnston and Mawson, 1940 Hosts: Pseudaphritis urvillei, Percalates collonormnm: Phectirpplites.ambiguus, Localization: intestine (?) — not given. Distribution: Australia. Description (after Johnston and Mawson, 1940). Buccal capsule with spiral thickenings not laterally flattened, 0.070 mm long and 0.070 mm wide in the middle, its walls very thick, its margins with six lobes. FIGURE 122. Spirocamallanus murrayensis (Johnston and Mawson, 1940): a — cephalic end; b — caudal end of male (after Johnston and Mawson, 1940). Male. Length 4—5 mm, maximum width 0.13 mm. Muscular anterior part of esophagus ending 0.4mm from the cephalic end, glandular posterior part ending0.9mmfromthis end. Both parts moreor less curved anteriorly. Nerve ring situated0.2 mm fromcephalic end. Caudal wings united anteriorly, 192 Г = q : 0.33 mm long. Three pairs of pedunculate preanal papillae, 2 pairs post- 203 anal, and 2 pairs of sessile papillae adanal. Spicules 0.29 and 0.20 mm long, simple, pointed. Tail 0.15 mm long. Female. Length 8—10 mm, maximum width 0.25—0.30 mm. Vulva forming a transverse slit before middle of body; branches of uterus op- posite. Tail 0.1 mm long, its narrow distal part 0.030 mm long and 0.010 mm wide. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Johnston and Mawson, 1940, p.347; Yamaguti, 1961b, р. 45. Spirocamallanus olseni Campana-Rouget and Razarihelissoa, 1965 (Figure 123) Moss: сене“ Nave raves i. litt ya mus анаест na lin cant uss Lutjanus sp. Localization: stomach. Distribution: Madagascar. Description (after Campana-Rouget and Razarihelissoa, 1965). Small nematodes, tapering slightly at both ends. Buccal capsule oblong, with thick- ened walls, inner surface with 8 or 9 spiral ribs; capsule 0.085—0.095 mm long and 0.062—0.072 mm wide. Mouth rounded, its inner diameter as long as the width of the buccal capsule; 4 submedian papillae in the outer ring and 2 distinct amphids laterally. Buccal capsule surrounded by 8 longitu- dinal muscle stripes with equal intervals. Anterior part of esophagus club- shaped, muscular, posterior part slightly longer, glandular. Nerve ring situated in middle of first part of esophagus, deirids slightly before it, excretory pore slightly behind it, near the connection of the two parts of the esophagus. 0200mm FIGURE 128. Spirocamallanus olseni Campana-Rouget and Razarihelissoa, 1965: a — cephalic end, dorsoventral; b — same, apical; c — caudal end of male, ventral; d — caudal end of female, lateral (after Campana-Rouget and Razarihelissoa, 1965). 193 4 Male. Length 9—15 mm, width 0.25 mm. Nerve ring situated 0.280 mm, 204 deirids 0.140 mm, excretory роге 0.390 mm from cephalic end. Muscular part of esophagus 0.38 mm long, glandular part 0.56 mm. Right spicule thick, gutter-shaped, 0.330—0.380 mm long; left spicule thinner and less strongly chitinized, 0.180—0.220 mm long. Caudal wings well developed, united anteriorly on the ventral surface, where the body wall is strengthened by oblique muscles. All papillae pedunculate,8 pairs: 3 pairs preanal, one pair adanal, and 4 pairs postanal; their arrangement varies slightly in dif- ferent specimens and is often asymmetrical, and the last pair, which is situated more ventrally than the preceding pair, is sometimes situated at the same level as this, posteriorly. Two pairs of teatlike, chitinized, finger- shaped formations at the cloaca. Female. Length 12—20 mm, width0.5 mm. Nerve ring situated 0.285 mm, deirids 0.175 mm, excretory pore 0.615 mm from the cephalic end. Muscular part of esophagus 0.55 mm long, glandular part 0.72 mm, Tail round, slightly tapering, witha finger-shaped appendage 0.040—0.042 mm long and with two small spines. In a specimen 14.5 mm long, the vulva is situated in the middle of the body, in other, larger individuals it is situated slightly behind the middle and in smaller specimens (12 mm) before the middle; the vulva is displaced therefore further posteriorly in the larger specimens. It protrudes weakly. Vagina directed posteriorly, branches of uterus opposite, distended, with thin wall, containing eggs and larvae. Reference: Campana-Rouget and Razarihelissoa, 1965, pp.171—176. Spirocamallanus ophiocephalus (Ali, 1960) (Figure 124) Synonym: Procamallanus .ophiocephalus, Ali, 1960 Host Ophiocephalus ‘punctatus’. LocoMe atone, gills: (7); Distribution: India. Description (after Ali, 1960). Male. Body thin, cylindrical, narrowing posteriorly and ending ina pointed tail. Cephalic end widened at the buccal capsule. Cuticle 0.037 mm thick in middle of body, transversely striated with intervals of 0.011 mm. Length 5.8 mm, width 0.19 mm. Dorsoventral width of cephalic end 0.13 mm, Excretory pore situated 0.33 mm, nerve ring 0.24 mm from cephalic end. Buccal capsule spherical, with sickle-shaped formations with 'manubria" arranged obliquely at the base; ''cutting edges'' form the margins of the capsule, their sharp ends contiguous immediately below the amphids. Five to 7 transverse ribs in the buccal capsule. Four cephalic papillae. Esoph- agus 1.67 mm long,1/4 of length of body, divided into two parts 0.66 and 1.01 mm long. Tail conical, with wide, symmetrical caudal wings 0.48 mm long and 0.26 mm wide. Eight pairs of riblike preanal papillae forming two groups of four. Postanal papillae also forming two groups. The first group consists of 4 pairs situated close together; a fifth pair is displaced to the 205 end of the tail. Two pairs of papillae situated around the cloaca: one pair 194 slightly before and one pair slightly behindit. Spicules of different length and form. Right spicule 5 times as long as the left,0.15 mm long. Left spicule 0.034 mm long. Right spicule with wide, rectangular anterior end and pointed posterior end. Left spicule wider in the middle, with narrow anterior end and inward-curved posterior end. Tail 0.2 mm long. Female unknown. FIGURE 124. Spirocamallanus ophiocephalus (Ali, 1960): a — anterior end, lateral; b — cephalic end, lateral; c — spicules; d — posterior end of male, ventral; e — arrangement of papillae around the cloaca in the male (after Ali, 1960). Reference: Ali, 1960, pp. 132—134. Spirocamallanus parasiluri (Fujita, 1927) Olsen, 1952 (Figure 125) Synonym: Procamallanus parasiluri Fujita, 1927 Host: Parasilurus asotus. Localization: intestine. Distribution: Japan. Description (after Fujita, 1927). Description based only on females. Body uniformly thin, tapering from the anus and ending in three small spines. Length 6 mm, width in the middle 0.14 mm. Except in the cephalic region, the cuticle is about 0.014 mm thick, with fine transverse striation with inter- 206vals of 0.004 mm. Buccal capsule formed by dark brown valves, truncate anteriorly and narrowing posteriorly. A wide, thick ring at the posterior end of the capsule. Outer surface of buccal capsule smooth, inner surface with 195 oblique parallel striation, Capsule 0.12 mm long and 0.07 mm wide. Tridents absent. Esophagus 0.43 mm long and 0.05 mm wide, its anterior part muscular. Intestine 4.68 mm long and 0.05 mm wide. Nerve ring situated 0.19 mm, excretory pore 0.16 mm from the cephalic end. Vulva situated 5.0mm from the cephalic end, i.e. at more than 2/3 of the body. One ovary, 1.45 mm long and 0.046 mm wide, situated in anterior half of body, extending anteriorly to 2.5 mm from the cephalic end. Uterus double: anterior part 1.11 mm long, larger than the posterior part, which is 0.65mm long and extends posteriorly almost to the anus. Vagina with thick wall, 0.41 mm long, directed obliquely posteriorly. All specimens were young, without ripe eggs. Tail 0.12 mm long, 29 mm =— о № b FIGURE 125. Spirocamallanus parasiluri (Fujita, 1927): a — buccal capsule; b — caudal end of male, lateral (a — after Fujita, 1927; b — after Yamaguti, 1935). Description (after Yamaguti, 1935). Yamaguti found four mature females and one immature male of this species and gave a brief description and a drawing of the caudal end of the male. Mouth wide, with membranous margin which is supported by regularly spaced spines in grooves on the thickened anterior margin of the buccal capsule, and with delicate muscle fibers attached to the spines which extend across the smooth outer surface of the capsule posteriorly. Inner surface of buccal capsule with continuous 207 spiral ribs. Cuticle to 0.030 mm thick at the nerve ring, which is situated 0.23—0.25 mm from the cephalic end. Male. Length 4.44 mm, width 0.11 mm, Buccal capsule 0.090 mm long and 0.075 mm wide. Muscular anterior part of esophagus 0.35 mm long and 0.05 mm wide, posterior glandular part 0.338 mm long and 0.05 mm wide. Yamaguti found only one spicule inthe form of anelongate horn. The subventral caudal wings do not project much. Eight pairs of caudal papillae, the preanal 196 208 209 и 9” papillae long and pedunculate, with intervals which decrease posteriorly; the three pairs of postanal papillae are similar, situated close together. Female. Length12.3 mm. Buccal capsule 0.108 —0.114 mm long and 0.090—0.096 mm wide. Muscular anterior part of esophagus 0.475—0.51 mm long and 0.06—0.08 mm wide, glandular posterior part 0.51—0.6 mm long and 0.050—0.070 mm wide. Vulva without protruding lips, situated at about 1/5 of the length of the body from the caudal end. Eggs almost round, 0.024—0.027 mm wide; eggs segmented, without developed larvae. Reference Ss: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Spudemikov 1 954, p.35; Fujita, 1927, рр. 169—176; Yamaguti, 1935, рр.347—386. Spirocamallanus реге!га1 (Annereaux, 1946) Olsen,1952 (Figure 126) Synonym: Procamallanus pereirai Annereaux, 1946 Hosts: Micropodon undulatuss Leprocottius: ar malt us), Обе Еву ое mirabilis, ~“Huntdulus parvupuninis 7 Gree lila Pee ея ms. | At hier TM O;pis abit ims Localization: intestine. ие: Во тоя: USA, Historical review Thedescriptionof Procamallanus pereirai (Annereaux, 1946)is the first record of the genus in North America. Sogandares- Bernal (1955)found P. pereirai in Micropodon (Sciaenidae). The parasite was originally found not in Perciformes butin Atherinops, which belongs to the Mugilidae. Noble and King (1959) found the species in Gillichthys mirabilis. They gave a description of P. pereirai and the drawings are reproduced here. They found that 800 out of 810 fish examined were infected with the parasite. Young specimens were usually localized in the anterior part of the intestine, larger specimens in the posterior part. All nematodes were at- tached to the mucosa by the buccal capsule. If there was food in the intestine, the nematodes concentrated around it. (Noble and King, 1959, рр. 681—683.) Noble and King (1959) found P. pereirai also in the following hosts: Leptocottus armatus (Cottidae), Atherinops affinis and Fundulus parvipinnis (Mugiliformes), and Girella nigricans (Perciformes). The infection rate of these hosts was high. The average number of specimens of P. pereirai was proportional to the size of the fish. The salinity of the water had no effect on the infection. Description (after Annereaux, 1946). Reddish in life. Body thin, narrower at both ends. Cephalic end truncate, mouth rounded, terminal. Buccal capsule barrel-shaped, with 14 continuous spiral thickenings in both sexes. Cuticle thick, finely transversely striated in the cephalic region; striation coarser in posterior part of body. Male. Length 6.8mm, width 0.189 mm. Posterior end slightly curved ventrally. Buccal capsule 0.07 mm long and 0.06 mm wide. Muscular part of esophagus 0.33 mm long, glandular part 0.43 mm. Caudal wings sym- metrical, connected ventrally 0.357 mm from end of tail, supported by 9 pairs 197 of riblike рарШае, 3 pairs preanal and 6 pairs postanal. Spicules of similar form but of different length: right spicule 0.43 mm long, left spicule 0.21 mm, Tail 0.138 mm long, with two small, rounded processes at the end. (208) FIGURE 126. Spirocamallanus pereirai (Аппегезих, 1946): a — anterior end; b — caudal end of male, ventral; c — caudal end of female, lateral; 9 — buccal capsule; е — cloaca of male, lateral; f — end of tail of female; g — caudal end of male, lateral (a—c — after Annereaux, 1946; d—g — after Noble and King, 1959). Female. Length 8.73—10.00 mm, width 0.232—0.258 mm. Length of buccal capsule 0.080—0.086 mm, width 0.07—0.08 mm. Muscular part of esophagus 0.39 mm, glandular part 0.51—0.53 mm. Excretory pore situated about 0.40 mm from the cephalic end. Vulva situated slightly behind middle of body. Tail tapering, constricted at the anus and ending in a finger-shaped process which is directed ventrally, with short terminal spines. Uterus containing eggs at different stages of development, but no larvae. Description (after Noble and King, 1959). Red in life, intestine blackish red, 198 Male (25 specimens examined). Length 10.80—17.65 mm, width 0.125—0.280 mm. Length of buccal capsule 0.052—0.079 mm, width 0.048— 0.076 mm. Muscular part of esophagus 0.280—0.500 mm long, glandular part 0.400—0.775 mm. Caudal end with symmetrical wings. Nine pairs of caudal papillae, 3 pairs preanal and 6 pairs postanal. All papillae with flattened apex, from which extend threadlike formations, which apparently connect them with the wings. Two pairs of small modified papillae, situated at the cloaca. These papillae were not mentioned by Annereaux. Spicules of similar form but of different length: right spicule about 0.43 mm long, left spicule 0.20 mm. Tail with two small, kidneylike appendages at the end. Female (25 specimens examined). Length 13.85—26.30 mm, width 0.235—0.460 mm. Length of buccal capsule 0.058—0.083 mm, width 0.052— 0.075 mm. Muscular part of esophagus 0.350—0.525 mm long, glandular Ва 0925 —0.897 тата. Vulva usually situated slightly behind middle of body. Uterus usually filled with eggs and embryos. Posterior end pointed and narrowing behind the anus with a narrow ventral finger-shaped process, which ends in two small terminal appendages like those in the male. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Annereaux, 1946, pp.299—303; Noble and King, 1959, pp.679—685; Sogandares-Bernal, 1955, pp.587—594; Yamaguti, 1961b, p. 45. Spirocamallanus rarus (Travassos, Artigas, and Pereira, 1928) Olsen 1952 (Figure 127) Synonym: Procamallanus rarus Travassos, Artigas, and Pereira, 1928 (210) ?2./mm — FIGURE 127. Spirocamallanus rarus (Travassos, Artigas, and Pereira, 1928): a — anterior end of male; b — cephalic end of male; с — caudal end of male (after Travassos, Artigas, and Pereira, 1928). 199 210 Hosts: Pimelodella lateristriga, Rhynodoras dorbig- nyi. Localization: intestine. Distribution: Brazil. Description (after Travassos, Artigas, and Pereira, 1928). Male (one specimen), Length 5.3 mm, width 0.11 mm. Excretory pore situated about 0.38 mm, nerve ring 0.25 mm from the cephalic end. Buccal capsule wide, with 3—4 spiral ribs which project markedly; length of capsule 0.072 mm, maximum width 0.056 mm. Muscular part of esopha- gus separated from the capsule by a chitinized ring, about 0.68 mm long, 0.04 mm wide; glandular part cylindrical, 0.64 mm long and 0.048 mm wide, Caudal end with narrow wings and 8 pairs of papillae, 4 pairs preanal and 4 pairs postanal. Spicules 0.4 and 0.17 mm long. Tail 0.12 mm long, conical, Female unknown. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.33; Travassos, Artigas, and Pereira, 1928, pp.5—68. Spirocamallanus singhi (Ali, 1956) Olsen, 1952 (Figure 128) Synonym: Procamallanus singhi Ali, 1956 Host: Callhichrous: bimaculatus. Localization: stomach. Distribution: India. Description (after Ali, 1956). Reddish in life. Cuticle transversely striated with intervals of 0.005 mm. Mouth oval. Cephalic end with 2 pairs of submedian papillae and a pair of lateralamphids. Esophagus divided into two parts. Body cylindrical, narrower anteriorly; tail short, pointed in both sexes. Buccal capsule with double spiral thickenings. Male. Length 5.9—6.68 mm, thickness 0.133—0.143 mm. Excretory pore ‘situated 0.31 mm, nerve ring 0.105—0.107 mm from the cephalic end. Buc- В | cal capsule 0.059—0.065 mm long and 0.053—0.059 mm wide. Length of esophagus 0.68—0.71 mm, i.e. 1/8 of length of body. Muscular part of esoph- agus 0.305—0.316 mm long, glandular part 0.38—0.40mm. The coiled, single testis is situated in the esophageal region. It passes in the middle of the body into the vas deferens, which has the same width, and then into the seminal vesicle, which passes into the ejaculatory duct. Caudal end nar- rowing before the cloaca, Caudal wings 0.39—0.41 mm long. Eleven pairs of caudal papillae, 7 pairs preanal and 4 pairs postanal. Lateral cuticular processes present. Spicules of different length and form: right spicule 0.200—0.205 mm long, left spicule 0.042—0.044 mm. The short spicule is widened in the middle, then becomes narrower and then widens again, The long spicule is almost of the same width throughout, with pointed distal end and slightly widened proximally. Tail 0.049—0.052 mm long. Female. Length 6.152—6.92 mm, width 0.166—0.170 mm. . Ехсгеогу pore situated 0.45 mm, nerve ring 0.168—0.180 mm from the cephalic end. 200 FIGURE 128. Spirocamallanus singhi (Ali, 1956): a — cephalic end, apical; b — same, lateral; c — anterior end of female, lateral; d — posterior end of female, lateral; e — posterior end of male, ventral; f— region of vulva, lateral; g — spicules (after Ali, 1956). Buccal capsule 0.076—0.083 mm long and 0.059—0.066 mm wide. Length of esophagus 1.05—1.12 mm,i.e. 1/6 of length of body. Muscular part of esoph- agus 0.44—0.48 mm long, glandular part 0.61—0.64 mm. Oviduct narrow, ovary threadlike, forming a loop and extending anteriorly to the esophagus. Seminal vesicle swollen. Uterus extending posteriorly; vagina also connected with the blind opposite uterine sac. Vagina about 2.85—3.3 mm long, slightly sinuous, vulva protruding, with distinct lips. Tail 0.025 mm long, conical, with two small processes at the end. еее ее 1956, ро: 95—11. Уамабыы. 19615: р. 46. 212 Spirocamallanus tornquisti (Baylis, 1923) Campana-Rouget, 1961 (Figure 129) Synonym: Procamallanus spiralis Baylis, 1923 sensu Tornquist, 1931 Ect Брава © mena Gems Or aki iy Sao US sim ones Labridae —Cheilinus trilobatus; Soleidae —Achinus sp.; Sciaeni- dae. Localization: stomach and intestine. Distribution: Red Sea, Gulf of Suez, Gulf of Mexico, Pacific coast of Mexico. 201 Historical review Campana -Rouget (1961a) noted discrepancies between the description of Procamallanus spiralis of Tornquist(1931)and that of Baylis: the buc- cal capsule contains 9 spiral ribs instead of 12, the proportions of the two parts of the esophagus are different, andthe males are markedly larger; she there- fore suggested that Tornquist's specimens are а different species and named them Spirocamallanus tornquisti. Campana-Rouget stated that the number of species of Spirocamallanus parasitizing in marine fishes is very small compared with the number in freshwater fishes, and that it is unlikely that the same species would be found inhosts witha different biology. Sheis therefore of the opinion that Winter's specimens (1953 ) from Mexican fish are во овала ti. Е а de TILL FIGURE 129. Spirocamallanus tornquisti (Baylis, 1928): a — anterior end; b — caudal end of male, lateral; c, d — caudal end of female, lateral (after Tornquist, 1931). Description (after Tornquist, 1931). Inner surface of buccal capsule with spiral ridges, outer surface with longitudinal ridges. Excretory system of the type "Р". Male. Length 10.924—13.505 mm. Nerve ring situated 0.172—0.234 mm from the cephalic end. Posterior part of esophagus 0.561—0.672 mm long, maximum width 0.156—0.234 mm. Tail 0.062 mm long. Buccal capsule 0.062—0.094 mm long. Female. Length 13.026—27.160 mm. Length of buccal capsule 0.078—0:094 mm. Nerve ring situated 0.250—0.265 mm from the cephalic end. Length of posterior part of esophagus 0.561—0.655 mm. Vulva situated 5.507—12.48 mm from anterior end. Tail 0.172—0.187 mm long. 202 213 Description (after Winter, 1953). Male not described. Female. Redinlife. Length 17.930—21.663 mm, maximum width 0.232—0.564 mm. Cephalic end with 6 weakly projecting lobes. Two pairs of cephalic papillae and 2 amphids. Mouth rounded. Cuticle with distinct transverse striation at the ends of the body, without striation in the middle. Tail conical, truncate, with a narrow process with two small teatlike appendages. Buccal capsule barrel-shaped, 0.076—0.106 mm long and 0.068—0.087 mm wide. Its amber-colored lining has a spiral thickening with 9 turns. At the base of the capsule is a ring before the esophagus. Club-shaped anterior part of esophagus 0.498—0.664 mm long and 0.065—0.106 mm wide. Gland- ular posterior part also club-shaped, 0.566—0.863 mm long and 0.061— 0.114 mm wide. It projects into the intestine with a short valve apparatus. Intestine 0.053—0.103 mm wide. Anus situated 0.082—0.217 mm from posterior end. There is a distinct musculature in the region of the anus. Nerve ring situated in the posterior part of the glandular part of the esophagus, 0.274—0.361 mm from the cephalic end. Cervical papillae small, situated 0.144—0.152 mm, excretory pore 0.304—0.329 mm from the cephalic end. Vulva situated in middle of body, 8.964—9.960 mm from posterior end. Vagina directed posteriorly, the part near the vulva with thick muscular walls; vagina 0.057—0.076 mm long, extending posteriorly for 0.498— 1.240 mm. Uterus filling the greater part of the body cavity, containing embryos 0.399 mm long and 0.015—0.019 mm wide. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р.34; Campana-Rouget, 1961а, рр. 24—25; Tornquist, ISIS, 300; NVateies 1953.1 рр. 137—139 Уажмазаы, 19615, jo, 26. Spirocamallanus vachai Sinha and Sahay, 1965 (Figure 130) ЕЕ Eauutte O\pgiac вв ув. уасва. Localization: intestine. Di Sues Lo tt On = Шота. Description (after Sinha and Sahay, 1965). Small nematodes, blood-red in life. Cuticle thick, with fine transverse striation. Buccal capsule chitin- ized, without teeth, tridents, or papillae; it contains two systems of obliquely intersecting spiral thickenings. Esophagus divided into a club-shaped mus- cular anterior part and a longer glandular posterior part; ratio of length of these parts 1:1.5. Male. Length 4.64—7.308 mm, maximum width 0.05—0.084 mm. Length of buccal capsule 0.0 4—0.05 mm, width 0.042 mm. Muscular part of esoph- 214 agus 0.34—0.40 mm long, glandular part 0.42—0.52 mm. Nerve ring situated 0.124—0.148 mm from the cephalic end. Tail conical, 0.04—0.042 mm long, curved ventrally, with wings. Six pairs of preanal and 5 pairs of post- anal pedunculate papillae; a winglike cuticular process on each side of the body behind the last postanal pair of papillae. Spicules weakly chitinized, bifid proximally, distal end pointed. Right spicule 0.182 mm long, left spicule 0.056 mm. 203 215 Male. nea EL ТАТА ИТУ И | a т 4 arr FIGURE 180. Spirocamallanus vachai Sinha and Sahay, 1965: а — anterior end; b — caudal end of male, ventral; с — caudal end of female, lateral (after Sinha and Sahay, 1965). Female. Length 8.9—12.72 mm, width 0.1—0.164 mm. Length of buccal capsule 0.056—0.064 mm, width 0.046—0.056 mm. Muscular part of esopha- gus 0.45—0.52 mm long, glandular part 0.58—0.72 mm. Nerve ring situated 0.12—0.14 mm from the cephalic end. Tail conical, 0.2 mm long, without papillae, ending in two short processes. Vulva situated behind middle of body, 3.28—3.68 mm from end of tail; vagina muscular and sinuous. parous. Vivi- Reference: Sinha and Sahay, 1965, рр. 49—53. Spirocamallanus viviparus (Ali, 1956) Olsen, 1952 (Figure 131) Synonym: Procamallanus viviparus Ali, 1956 Host: Mystus microphthalmus. Localization: stomach, Drstributvon: India: Description (after Ali, 1956). Cuticle thick, transversely striated with intervals of 0.005 mm in middle of body. Body narrowing anteriorly and posteriorly. Two submedian pairs of cephalic papillae and 2 lateral amphids. Reddish in life. Buccal capsule barrel-shaped. Length 2.3—3.19 mm, maximum width 0.083—0.092 mm. Nerve ring surrounding muscular part of esophagus 0.098—0.110 mm from the cephalic end. Excretory pore situated 0.26 mm from the cephalic end. Buc- cal capsule 0.051—0.082 mm long and 0.047—0.048 mm wide. Anterior part of esophagus 0.20—0.23 mm long, posterior part 0.25—0.29 mm. Length of esophagus 0.45—0.52 mm, i.e. about 1/6 of length of body. Testis thread- like, extending to the anterior end of the intestine. Caudal wings wide anteriorly, narrower posteriorly. Spicules of different length and form, right spicule 0.186 mm long, left spicule 0.069 mm, Ratio of length of spicules =3:1. Right spicule with wide proximal part, curved inward at the 204 distal end. Left spicule slightly widened in the middle and curved like а hook at the distal end. Caudal wings 0.47 mm long and 0.15 mm wide, their anterior border passing across the ventral surface. Ten pairs of caudal papillae, 6 pairs preanal and 4 pairs postanal. Cuticular processes present between the third and last postanal pair of papillae. Tail 0.043—0.054 mm long. FIGURE 131. Spirocaimallanus viviparus (Ali, 1956): а — cephalic end, apical; b — same, lateral; с — anterior end of female, lateral; d — posterior end of female, lateral; e — posterior end of male, ventral; Г — region of vulva, lateral; р — spicules (after Ali, 1956). Female. Length 4.98—5.27 mm, maximum width 0.183—0.211 mm, Nerve ring situated 0.133—0.138 mm, excretory pore 0.29 mm from the cephalic end. Tail with three small processes at the end. Buccal capsule 0.059—0.061 mm long and 0.054—0.057 mm wide. Muscular anterior part of esophagus 0.29—0.31 mm long, glandular posterior part 0.41—0.43 mm. Length of esophagus 0.70—0.74 mm, about 1/7 of length of body. One ovary, extending anteriorly to the anterior third of the body. The oviduct surrounds the intestine, and passes into the seminal vesicle at the connection between esophagus and intestine. Vulva situated between third and last quarter of body. The posterior blind branch of the uterus extends to the anus. Vagina to 0.35 mm long. Vulva with protruding lips. Tail 0.038 mm long. hehe remces “Ali, 1955, рр. 13—15; Yamacutl, 1961 b, в.46. 205 216 Spirocamallanus wrighti (Pereira, 1935) Olsen, 1952 (Figure 132) Synonym: Procamallanus wrighti Pereira, 1935 Hosts: Astyanax sp., Hoplias malabaricus, Leporinus Sp: )-Py pocentnus’ sp: Localization: small intestine. Distribution: ‘Brazil. Description (after Pereira, 1935). Small, reddish nematodes. Male. Length 3.86—4.6 mm, width 0. 18—0.27 mm. Cuticle with fine transverse striation. Mouth rounded. Buccal capsule large, spherical. Its walls with 12—16 chitinized ribs which are wide at the base, becoming narrower anteriorly and to the right, ending before the margin of the mouth, Width of buccal capsule 0.04—0.06 mm. Muscular part of esophagus club- shaped, 0.27—0.34 mm long, maximum width 0.08 mm; between esophagus and buccal capsule is a chitinized ring 0.008 mm high. Glandular part of esophagus about 0.47 mm long. Nerve ring situated 0.18 mm, excretory pore 0.17—0.18 mm from the cephalic end. Wer a е FIGURE 132. Spirocamallanus wrighti (Pereira, 1935): a — anterior end, lateral; b — cephalic end, lateral; с — caudal end of male, ventral; d — caudal end of female, lateral; e — distal end of spicules (after Pereira, 1935). 206 ee oe Tail curved ventrally, with caudal wings which begin 0.25 mm from end of tail; anus situated 0.13—0.15 mm from end of tail. Spicules short, weakly chitinized, about 0.045 mm long. The ventral surface of the posterior end contains distinct oblique muscles. There are eight pairs of papillae in two subventral rows, 4 pairs preanal and 4 pairs postanal. ы Female. Length 8.2—10.6 mm, width 0.30—0.38 mm. Cuticle with fine transverse striation. Buccal capsule 0.049—0.11 mm wide, with 20 chitinized ribs. Muscular part of esophagus 0.32—0.42 mm long, glandular part 0.5— 0.70 mm. Nerve ring situated 0.25—0.29 mm, excretory pore 0.21—0.42 mm 217from the cephalic end. Anus situated 0.12—0.23 mm from the posterior end. Tail tapering. Vulva situated 3.4—4.5 mm from end of tail; branches of uterus diverging. Viviparous. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.34; Pereira, 1935, рр. 53—62. Spirocamallanus xenopodis (Baylis, 1929) Olsen, 1952 (Figure 133) Synonym: Procamallanus xenopodis Baylis, 1929 Host: Xenopus muelleri. Localization: intestine (?). Diisie wb uit wom: Асса» Description (after Baylis, 1929b). Small nematodes. Buccal capsule barrel-shaped, its walls with irregularly interrupted ribs which form a spiral or are arranged diagonally. Striation of cuticle coarser in males than in females. Male. Length 0.60—1.37 mm, maximum width 0.12—0.15 mm. Cuticular striation with intervals of 0.004 mm. Length of buccal capsule 0.08—0.10 mm. Muscular part of esophagus 0.26—0.30 mm long. Posterior end of glandular part of esophagus situated 0.46—0.50 mm from the cephalic end, nerve ring situated 0.13—0.15 mm from the cephalic end. FIGURE 133. Spirocamallanus xenopodis (Baylis, 1929): a — anterior end; b — caudal end of female, lateral (after Baylis, 1929b). 207 Nine pairs of preanal pedunculate papillae and 4 pairs of postanal papillae, the three anterior pairs with long peduncles and situated subventrally; the last pair, large and sessile, is lateral. Spicules of different length, left spicule shorter and markedly thicker than the right. Right spicule 0.10— 0.11 mm long. Tail with blunt end, smooth, 0.03 mm long. Female (immature). Length 0.74—0.78 mm, maximum width 0.09— 0.10 mm. Length of buccal capsule 0.070—0.075 mm. Distance from ce- phalic end to end of muscular part of esophagus 0.22—0.24 mm, to end of glandular part 0.37—0.38 mm. Nerve ring situated 0.11—0.12 mm from the cephalic end. Vulva situated 0.31—0.41 mm from end of tail which bears 7 spines, one terminal and median, the other six forming a ring; the number of spines is constant. Tail.0.04—0.05 mm long. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.34; Baylis, 1929b, pp.378—380; Yamaguti, 1961b, p.113. 218 Spirocamallanus sp. Campana-Rouget, 1961 (Figure 134) Host. synodontts зева, Localization not given. Прето стом" Africa, Description (after Campana-Rouget, 1961а). Length of buccal capsule 0.09 mm, width 0.08 mm; capsule with 10 spiral ribs on the inner surface. Outer surface of capsule with 8 finger-shaped formations which begin at the Раз ` and end in about the middle of the capsule; they are weakly chitinized. Tornquist thinks that they form the outer layer of the buccal capsule. FIGURE 134. Spirocamallanus sp. Campana-Rouget, 1961. Cephalic end (after Campana-Rouget, 1961a). Length of muscular part of esophagus 0.51 mm, of glandular part 0.43 mm. Nerve ring situated 0.28—0.30 mm, deirids 0.174mm from the cephalic end. In an immature female the vulva was situated 6.74 mm from the cephalic end. Right spicule 0.170 mm long, left spicule 0.122 mm. Three pairs of pre- anal papillae, 4 or 5 pairs of postanal papillae, and 2 pairs of adanal papillae. Reference: Campana-Rouget, 1961а, pp. 23—24. 208 SUPERFAMILY ANGUILLICOLOIDEA N. SUPERFAM. Historical review Sobolev (1952), in his classification of the suborder Camallanata, in- cluded the family Anguillicolidae Yamaguti, 1935 in the superfamily Dracunculoidea. Shigin and Shigina (1958) described the new species Skrjabillanus tincae, for which they established a new genus. This differs so markedly from all other genera of the family that the authors placed it in a separate family, Skrjabillanidae, which they considered related to the family Anguillicolidae. Campana-Rouget (1964) considered the family Anguillicolidae as related to the family Tetanonematidae Skrjabin and Schikhobalova, 1948, which Skrjabin and Shikhobalova (1948) had established for the genus Tetanone- ma Steiner, 1937, in contrast to the author, who had placed the genus pro- visionally in the subfamily Aproctinae. Yamaguti (1961b) placed the family Tetanonematidae in the order Tetano- nematidea Yamaguti, 1961. We think the families Anguillicolidae Yamaguti, 1935, Tetanonematidae Skrjabin and Schikhobalova, 1948, and Skrjabillanidae Schigin and Schigina, 1958 should be placed in a new superfamily which we name Anguillicoloidea Sobolev, Ivaschkin, Tichomirova, and Khromova п. superfam. We also place the family Phlyctainophoridae Roman, 1960 provisionally in this superfamily. Diagnosis. Camallanata. Body threadlike or cylindrical. Males smaller than females. Cephalic papillae present. Stoma developed or re- 219duced. Esophagus well developed or reduced. Anus present or absent. Spicules and gubernaculum absent in males, or a chitinized plate is present. Caudal papillae present. Vulva present. Parasites of the connective tissue, body cavity, and blood sinuses of fish and Cyclostomata. Type family: Anguillicolidae Yamaguti, 1935. Key to the families of the superfamily Anguillicoloidea PS VOL LICACVECLODESE sass ress ane se с а ome erate со ат hohe ee 2 ес” ея ол ен о: еее ее ai 3. 2. Caudal wings present in males. Vulva situated in anterior half of body И, ede cae Skrjabillanidae Schigin and Schigina, 1958. Caudal wings absent. Vulva situated in posterior part of body ....... Ро aA aula ck eel «Wa cash Slate oda о Anguillicolidae Yamaguti, 1935, 3. Воду long, threadlike. Cuticle with more or less irregularly distrib- Mie OUD CROLE SHIM aMLerlOr ране ово ею О Ser ae eR Tetanonematidae Skrjabin and Schikhobalova, 1948. Body cylindrical, curved ventrally. One row of symmetrical rounded processes on each side situated subdorsally, 15 processes in a row Beart ie Gace ee iawn cco) io Reema За Phlyctainophoridae Roman,1960. 209 219 FAMILY ANGUILLICOLIDAE YAMAGUTI, 1935 Diagnosis. Anguillicoloidea. Slender nematodes. Stoma developed. Esophagus consisting of a muscular anterior swelling and a cylindrical muscular-glandular posterior part. Anus absent. Male without spicules. Caudal papillae present. Vulva situated in posterior half of body. Vivi- parous. Type genus: Anguillicola Yamaguti, 1935, Genus Anguillicola Yamaguti, 1935 Diagnosis. Anguillicolidae. Buccal cavity wide. Esophagus divided into a muscular anterior swelling with three projecting lobes and a cylind- rical muscular-glandular posterior part. The posterior end of the eso- phagus forms a glandular ventricle which projects into the intestine. In- testine swollen, dark. Caudal glandular cells project into the ejaculatory duct. Male. Spicules absent. Testis beginning at the caudal end. Seminal vesicle distinct. Ejaculatory duct opening on a projecting papilla. Female. Vulva situated in posterior half of body, on the apex of a projecting cone. Vagina short, surrounded by dense glandular cells. Branches of uterus opposite. Viviparous. Parasites of the swim bladder of freshwater fishes. Type -speciecsi А. clobiceps—Yamasutl, 1955. Anguillicola globiceps Yamaguti, 1935 (Figure 135) Host: Anguilla, japonica. Localization: swim bladder. Distribution: Japan. Description (after Yamaguti, 1935). Male. Length 26 mm, width 0.8 mm. Length of buccal capsule 0.06 mm. 221 Anterior swelling of esophagus 0.38 mm wide. Nerve ring situated 0.6mm from the cephalic end. Six pairs of caudal papillae. Female. Length 60 mm, width 1.8 mm. Length of buccal capsule 0.1 mm. Anterior swelling of esophagus 0.7 mm wide. Nerve ring situated 0.6 mm from the cephalic end. Vulva situated about 5 times farther from the cephalic end than from end of tail. Larvae in uterus 0.28 mm long. 210 (220) 7 1500 TM АИ тата а — Не = 5 FIGURE 135. Anguillicola globiceps Yamaguti, 1935: a,b — cephalic end, different aspects; c, d, e — caudal end; f, h, i — anterior end; g, m — caudal end; j — cephalic end; k — region of vulva; 1 — egg with larva and larva; п — female, general view (a—e — after Yamaguti, 1935; Е-п — after Wu, 1956). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.90; Yamaguti, 1935, pp. 347—386; 1961b, pp. 79—80; Wu, 1956, рр. 99— 106. 211 Anguillicola australiensis Johnston and Mawson, 1940 (Figure 136) Host: Anguilla reinhartri. Localization: swim bladder. Distribution: Australia. Description (after Johnston and Mawson, 1940). Cephalic end bulb- shaped, neck distinctly constricted. Body tapering posteriorly, tail pointed. The anterior bulb-shaped widening is much wider dorsoventrally than late- rally. Mouth with 6 small papillae; buccal cavity wider at the base than anteriorly, with toothed inner margin which appears like a crown of leaves. Ratio of length of esophagus to length of body 1:30; esophagus strongly mus- cular, widening proximally. At the anterior end of the esophagus are 6 lobes which project into the buccal cavity. Nerve ring situated behind the cephalic swelling. Excretory pore situated on an elevation near the posterior end of the esophagus. Intestine wide, with dark content; the dark intestinal contents are not visible at the posterior end, and the rectum is probably a narrow tube. ty GH ПОТА : ди on Ивеко meu f FIGURE 186. Anguillicola australiensis Johnston and Mawson, 1940: а — cephalic end, lateral; b — same, ventral; с — caudal end of female; Я — caudal end of male, lateral; е — region of vulva; f — anterior end, lateral (after Johnston and Mawson, 1940). 222 Male. Length 40 mm, width 1mm. Two pairs of preanal and 2 pairs of postanal papillae. Spicules absent. Female. Length 60—70 mm, width 1.5 mm (young females 25—30 mm long,0.5 mm wide). In a specimen 25 mm long the swollen cephalic end is 0.14 mm long, 0.22 mm wide dorsoventrally, and 0.13 mm wide laterally. Length of buccal capsule 0.010 mm, width in the middle 0.028 mm, Esopha- gus 0.82 mm long. Nerve ring situated 0.18 mm from the cephalic end. An anus was not found in females, but 0.4 mm from the end of the tail in a spe- cimen 25 mm long and at corresponding points in larger specimens there is a small incision before which are situated four large glandular masses; before these, 1.1mm from end of tail, the dark intestinal content is not visible; 212 the rectum is probably a narrow tube. Vulva with protruding lips, situated on a round elevation which is situated 1/6 of the length of the body from the posterior end. Ovaries extending into the anterior and caudal regions. Eggs with thin shell, 0.012—0.015 X0.025—0.026 mm large. Reference: Johnston and Mawson, 1940, р.351. 213 222 FAMILY PHLYCTAINOPHORIDAE ROMAN, 1960 Historical review Steiner (1921 )described a unique nematode from a single female which he named Phlyctainophora lamnae. The form ofthe body was not des- cribed, as it was studied only from sections. Yorke and Maplestone (1926) con- sidered the systematic position of this genus as uncertain, while Chitwood (1937) placed it in the subfamily Philometrinae. Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov (1954) also retained the genus in the subfamily Philometrinae, but Ruyck and Chabaud (1960) did not agree; they found lar- vae of this nematode on the fins of fish and because of their possible direct development, they assumed that the genus is closely related to the group of tissue Aphasmidia: Muspicea and Robertdollfusa. Roman (1960) established for this genus the family Phlyctainophoridae Roman, 1960, which she placed in the order Enoplida of the subclass Adenophorea. Until a more detailed study is made of the species of this family, we place it provisionally in the superfamily Anguillicoloidea. Diagnosis. Anguillicoloidea. Cuticle smooth, but with slight striation in some places. Intestine narrow compared with the size of the parasite, Uterus filled with embryos; vulva not found. Adults and young forms parasitize in the tissue of fish. type genus: Philyctainoephora. Steiner, 1921. Genus Phlyctainophora Steiner, 1921 Diagnosis. Phlyctainophoridae. Body curved ventrally. A row of symmetrical rounded protuberances on each side, subdorsally 15 in a row. Anterior end with one such swelling, posterior end with two swellings. Cuticle smooth. Intestine narrow. Viviparous. 223 Parasites of Euselachiae. Type species: Phlyctainophora lamnae’ Эешее, 192 Phlyctainophora lamnae Steiner, 1921 (Figure 137) Hoste: Wamin al corn b iea: Массе мыслей. Localization: between mandibular cartilage and skull. Distribution: Atlantic Ocean (?). Description (after Janicki and Rasin, 1930). Male unknown, 214 C/00™mm Ss FIGURE 137. Phlyctainophora lamnae Steiner, 1921: a — genetal view; b — anterior end; с, Я — cross sections; e — anterior end of larva; f— larva; р — anterior end of larva; В — larva; i — caudal end of larva (a—f — after Steiner, 1921; g—i — after Ruyck and Chabaud, 1960). 224 Female. Length 17 mm. Body curved ventrally, consisting of a tube filled with embryos. On the outside are 15 round protuberances 0.009 mm wide situated dorsally and submedially. Musculature absent. The thin- walled intestine follows the curvature of the body. Mouth and anus not recognizable, Uterus large, filled with larvae. Larvae slender, with a tooth- like appendage at the cephalic end. : The life cycle has not been studied, but in 1960, Ruyck and Chabaud found and described larvae of nematodes which they identified as Phlyc- tainophora lamnae in swellings at the base of the ventral surface of the abdominal fins of Mustelus mustelus. Description of larvae. Length 0.520—0.600 mm, width 0.030 mm. Cephalic end with a distinct dorsal tooth. A larva 0.600 mm long showed the following characters: esophagus cylindrical, widened posteriorly, 0.080 mm long. Nerve ring situated 0.030 mm, excretory pore 0.110 mm from the anterior end. The intestine occupied almost the whole body cavity. Rectum 215 0.070 mm long. Genital opening ventral, 0.025 mm long, situated 0.140 mm before the anus. Тай 0.120 mm long, tapering and with pointed end. Phas- mids not found. The general appearance of the larvae and the simple esophagus probably indicate that they live in the tissues and cannot survive in the outer en- vironment; the large size of the larvae makes it impossible for them to be transmitted by hematophagous vectors. Their life cycle is therefore prob- ably the same as in the tissue Aphasmidia, or perhaps as follows: An adult female, dying in the host, releases a large number of larvae which grow and disperse throughout the tissues. The body of such a shark infected with larvae may be eaten by other sharks, so that infection is probably transmitted by cannibalism. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.85; Janicki and Rasin, 1930, p.33, Ruyck and Chabaud, 1960; pp. 386—389; Steiner, 1921, рр. 591—595. 216 oe Sree e FAMILY SKRJABILLANIDAE SCHIGIN AND SCHIGINA, 1958 Historical review The family Skrjabillanidae was established for the new genus and species Skrjabillanus tinca from Tinca tinca. Molnar (1966b) described two further species of the genus: 5. scardinii and 5. erythrophthal- mi from Scardinius erythrophthalmus in Hungary. In 1966, during a study of the helminth fauna of fish in Czechoslovakia, Moravec (1968) found these three species in the typical hosts and made a comparative study of the morphology of the 3 species. He found that S. erythrophthalmi differs markedly from the other two species in the form of the cephalic end and type of buccal capsule, and considered that these differences justified the estab- lishment of а new genus. He also noted that some characters of S. eryth- rophthalmi do not agree with the original diagnosis of the genus Skrja- billanus but were added by Molnar in 1966. Moravec therefore proposes 225the new genus Molnaria for erythrophthalmi, with the type species бе ста erythropht maim: Hetransferred hihi omiet ga vin testinalis (Dogiel and Bychowsky, 1934) Moravec, 1968 and Philo- metra leucisci (Agapova, 1963) Moravec, 1968 to the genus Molnaria. Tikhomirova (1968—1969) (published here for the first time) made a detailed study of the morphology of 5. tincae апа 5. scardinii and concluded that the buccal capsule of S. tincae differs markedly from ina Onto. tineae. The buccal capsule of (Ss. Scardinit isihexagonal; that of S. tincae is cup-shaped. The outer width of the capsule is 0.013 mmin 5. tincae, 0.018 юм ш 5. scardinii. At the base of the capsule of S. scardinii there are 6 chitinous teeth (two teeth on each sector of the esophagus). The teeth are almost of the same size and have the form of a truncate pyramid. The buccal capsule of S. tincae does not have such chitinized formations. There are 4 tubercles (of the embedded type) at the cephalic end ш 5. scardinii, but such structures areabsentin 5. tincae. Malesof 5. tincae have 6 pairs of papillae at the caudal end, while in males of 5. scardinii there are only 5 pairs of caudal papillae; a pair of sessile preanal papillae is absent. S. scardinii parasitizes in the serosa of the swim bladder and in the eyes; S. tincae is a parasite of the serosa of the kidneys. Tikhomirova decided therefore to establish the new genus Agrachanus for S. scardinii with the type species A. scardinii (Molnar, 1966) Tichomirova п. comb. There are thus three genera in the family Skrjabillanidae: Skrjabilla- nus Schigin and Schigina, 1958; Agrachanus Tichomirovan. gen. ; Molnaria Moravec, 1968. 217 Diagnosis. Anguillicoloidea. Body threadlike. Cephalic end with or without tubercles. Buccal capsule strongly or weakly developed. Esophagus muscular or muscular-glandular. Amphids very large. Males markedly smaller than females. Anus obliterated in adults. Caudal end of males with wings supported by pedunculate and sessile papillae. The sessile papillae are preanal and postanal or only postanal. Spicules and gubernaculum absent; a chitinous rod or. chitinous plate is present. Genital opening sit- uated on a strongly projecting genital papilla. Vulva situated in anterior part of body. Caudal end of females bluntly rounded and with three pro- cesses. Type genus: Skrjabillanus Schigin and Schigina, 1958. Key to the genera of the family Skrjabillanidae (after Tikhomirova) 1. Buccal capsule smooth, without chitinized teeth at the base........... Seas ЕЕ RIOR ae Skrjabillanus Schigin and Schigina, 1958. Chitinized teeth present at the base of the buccal capsule ......... а; 2. Three chitinized teeth at the base of the buccal capsule ........... Е ty uA Е ee i al ion Molnaria Moravec,’ 1968. Six chitinized*teeth at the base of the buccal сари ее. eset tees BNET За ИИ, пос. Бам Agrachanus Tichomirova п. gen. Genus Skrjabillanus Schigin and Schigina, 1958 Diagnosis. Skrjabillanidae. Threadlike nematodes with rounded anterior end and pointed posterior end. Cephalic end club-shaped. Buccal 226 capsule cup-shaped. Intestine straight, ending blind before the posterior end. Amphids large, bean-shaped. Females more than twice as large as males. ' Males with 6 pairs of genital papillae, three pairs long, pedunculate, post- anal. Sessile papillae both preanal and postanal. Genital opening situated on a markedly projecting papilla. A chitinized rod bent at a right angle projects partly outside. The vulva is obliterated in mature females; its rudiment is visible in the region of the esophagus. The greater part of the body cavity of the female is occupied by the large uterus, which is filled with larvae. Parasites of fish. Type species: Skrjabillanus tincae Schigin and Schigina, 1958. Skrjabillanus tincae Schigin and Schigina, 1958 (Figure 138) Вос" п eine о ти са,. Localization: under serosa of kidneys. Distribution: USSR, Czechoslovakia. Description (after Shigin and Shigina, 1958). Delicate, very thin, threadlike nematodes. Females more than twice as large as males. Males burst when placed in water. Cuticle delicately transversely striated with 218 intervals of 0.001—0.0012 mm; the cuticle of the anterior part of the body has ringlike girdles with intervals of 0.010—0.033 mm. Cephalic end slightly club-shaped. Mouth rounded, buccal capsule cup- shaped with thick, chitinized walls. Esophagus consisting of a muscular and a muscular-glandular part; glandular tissue situated on the dorsal side of the esophagus. Cephalic papillae arranged in two rings; inner ring consist- ing of large amphids and paired (dorsal and ventral), weakly developed papil- lae; outer circle with 8 papillae arranged in pairs. Male. Length 6.1—7.45 mm, maximum width 0.022—0.023 mm. Body widest in the middle and at the cephalic end. Mouth 0.005—0.006 mm wide. Inner width of buccal capsule 0.006—0.0065 mm, outer width 0.008—0.0085mm. Length of buccal capsule 0.0045 mm. Esophagus 1.047 mm long; muscular anterior part 0.275 mm long; the other part of the esophagus contains gland- ular tissue on the dorsal side. Intestine ending blind, 0.547 mm from the caudal end. (227) FIGURE 138. Skrjabillanus tincae Schigin and Schigina, 1958: a — cephalic end, lateral; b —same, apical; c, d — caudal end of male, different aspects; е — cephalic end; f — caudal end of male, ventral; р — caudal end of female, lateral (a—d — after Shigin and Shigina, 1958; e—g — after Ergens, 1960). Tail 0.032 mm long, with thin caudal wings which begin 0.015—0.017 mm above the genital opening and extend in wide ribbons to the end of the tail. The caudal wings are supported by three pairs of postanal pedunculate papillae situated in the anterior half of the tail. There are also 3 pairs of 219 sessile papillae, one pair preanal, situated slightly behind the beginning of the caudal wings, the other two pairs situated at the level of the second and third pairs of pedunculate papillae. Genital opening situated on a markedly projecting papilla. Gubernaculum and spicules absent. A chitinous 227 rod 0.008—0.010 mm long, bent at a right angle, projects from the anterior wall of the end of the ejaculatory duct. The distal end of this rod always projects halfway outside, so that it probably has the function of the absent spicules. Female. Length 16.5—22.0 mm, maximum width 0.047—0.065 mm. Inner width of buccal capsule 0.010 mm, outer width 0.013 mm, length 0.008 mm. Mouth 0.0067 mm wide. Caudal end bluntly rounded, with two ventrolateral processes, Vulva not found, but according to the position of the uterus, it is situated near the end of the esophagus. Viviparous. Almost the entire body cavity is occupied by the large uterus, which is filled with larvae 0.216 mm long and 0.0065 mm wide. Excretory pore situated 0.331 mm from the cephalic end. Description (after Ergens, 1960). Medium-sized, thin nematodes, cuticle with fine transverse striation. The cephalic end is the widest part of the body. Mouth passing into the buccal capsule, which is strongly chitinized. Esophagus consisting of a muscular and a glandular part. Vi- viparous. 228 Male. Length 7.0 mm, width 0.019 mm. Posterior end with well-de- veloped caudal wings with thin papillae. Three pairs of papillae pedunculate and postanal, the following 3 pairs short and sessile, the first pair pre- anal and the others postanal. Genital opening situated on a tubercle in the middle of the caudal wings. Gubernaculum and spicules absent, their function apparently taken by a rectangular, curved plate. Female, Length 15.4—19.8 mm, width 0.062 mm. Buccal capsule 0.007—0.008 mm long,0.009 mm wide. Tail blunt, with two small processes at the end. Vulva not found; the greater part of the uterus filled with larvae. References: Shigin and Shigina, 1958, pp.395—399; Ergens, 1960, _ pp. 230—231; Molnar, 1966b, pp.143—158; Moravec, 1968, р.322. Genus Артасйапиз$ Tichomirova in litt. Synonym: Skrjabillanus Schigin and Schigina, 1958 (in part) Diagnosis. Skrjabillanidae. Body threadlike. Cephalic end rounded and slightly widened. Buccal capsule hexagonal, with strongly chitinized walls and very strong base. Six teeth at the base of the capsule. Amphids large, rounded. Esophagus entirely muscular. Females more than twice as large as males. Tail of males with 5 pairs of papillae, four pairs pedun- culate and one pair sessile. Gubernaculum and spicules absent. A chitinized plate present. Vulva situated in anterior part of body. Viviparous. Para- sites of fish. Type species: Agrachanus scardinii (Molnar, 1966) Tichomirova in litt. 220 Agrachanus scardinii (Molnar, 1966) Tichomirova in litt. (Figure 139) Synonym: Skrjabillanus scardinii Molnar, 1966 Ost Nea mom nuts) ев у якорь i had mu sh. Localization: serosa of posterior part of swim bladder and kidneys. Юте остов: Hungary, USSR. Description (after Molnar, 1966b). Male. Transparent, threadlike nematodes. Length 3.2—4.4 mm. Body eylindrical, of the same width its entire length. Cuticle smooth. Cephalic end rounded, with 4 depressed papillae and 2 amphids. Buccal capsule chitinized, 0.006—0.007 mm wide and 0.004 mm long. Esophagus not divided, cylindrical, 0.025—0.026 mm long, 0.007—0.008 mm wide. An esophageal gland extends along the esophagus. Intestine of the same width its entire length. Bursalike caudal wings present at the caudal end, one wing dorsal, the other two lateral. The lateral wings are curved ventrally, forming a sac. Caudal end strongly curved ventrally; on the dorsal side it is grooved almost 229 to the end of the dorsal wing. At the level of the last pair of papillae two riblike formations extend to the posterior end of the lobe. The part covered with these formations refracts light strongly, thus differing from the other parts of the dorsal lobe. Caudal end with 4 pairs of pedunculate and one pair of sessile papillae. The last pair of pedunculate papillae is mushroom- shaped. Sessile papillae situated between the second and third pair of pe- dunculate papillae. Two equal short spicules, 0.006—0.007 mm long. They project into the transversely oval subterminal cloaca. Female. Transparent, threadlike nematodes. Length 8—13 mm, body of uniform width, 0.049—0.070 mm wide. Cuticle smooth. Four elongate papillae and 2 large amphids at the cephalic end. Buccal capsule 0.12 mm wide and 0.008 mm long, with 5 anteriorly directed teeth at its base. Eso- phagus of the same width throughout,opening into wide intestine through three 230 valves. Width of esophagus 0.012—0.014 mm. Intestine uniformly wide. A large esophageal gland begins below the buccal capsule; it gradually widens and extends posteriorly. At the valves of the esophagus it becomes markedly wider and extends along the intestine for double the length of the esophagus. The other esophageal gland is much narrower and extends from the middle of the esophagus to the vulva. Rectum gradually narrowing,passing into the subterminal anus. Vulva situated in anterior part of body, sometimes behind the anterior part of the uterus. Vagina short, weakly chitinized, thin, passing into the markedly widened ovejector which extends posteriorly for a short distance, then turns anteriorly to the anterior end of the uterus and makes a loop, becomes gradually wider, and passes into the uterus. The posterior end of the uterus is rounded and passes into the narrow, spirally coiled oviduct and the single ovary, the narrowed distal part of which is bent like a hook at the rectum. Tail tapering, with two dorsolateral and one ventral teatlike processes 0.003—0.004 mm long. According to the stage of de- velopment, the larvae are 0.090 to 0.140 mm long. Description (after Tikhomirova). Thin, delicate forms. Cephalic end slightly widened. Four cephalic tubercles of the embedded type situated submedially, thin and not always distinct. Buccal capsule hexagonal. The walls of the capsule, particularly the base, are strongly chitinized. Base of buccal capsule with 6 chitinous teeth inthe form of a truncate py- ramid; teeth of almost the same size except one, which is apparently reduced. 221 (229) 002 тит ПЕ FIGURE 139. Agrachanus scardinii (Molnar, 1966): а — cephalic end, apical; b — same, different aspect; с — caudal end of male, ventral; 4 — anterior end; e — caudal end of female, lateral; f — cephalic end; g — same, apical; h — caudal end of male, ventral (a—e — after Molnar, 1966b; f-h — after Tikhomirova). Esophagus entirely muscular, at first widened and then of uniform width its entire length. Esophagus surrounded by distinct glands. Four cephalic papillae. Two large, rounded amphids. Male. Length 4.5—5.0 mm, width 0.02—0.03 mm. Inner width of buccal capsule 0.010—0.012 mm, outer width 0.012—0.014 mm; length 0.005 mm. Esophagus 0.3—0.4 mm long. Caudal end with thin wings which begin slightly before the genital opening. The wings are supported by 5 pairs of papillae, four pairs pedunculate and one pair sessile, situated between the second and third pair of pedunculate papillae. Length of pedunculate papillae varying. Tail of varying form. Genital opening situated on a strongly projecting papilla. Spicules and gubernaculum absent; there is a chitinized plate 0.005 mm long, of varying thickness. Female. Length 14.0—16.0 mm, width 0.05—0.07 mm. Inner width of buccal capsule 0.010—0.012 mm, outer width 0.018 mm; length of capsule 0.005—0.007 mm. Lateral walls of capsule 0.0018 mm thick, base 0.0036 mm thick. Mouth rounded, 0.0072 mm wide. Cephalic tubercles 0.0072 X 0.0108 mm large. Esophagus 0.34—0.53 mm long. Nerve ring situated 0.27 mm from the cephalic end. Vulva situated in anterior part of body; vagina short and narrow. Ovejector well developed. Uterus filled with larvae which are, according to their age, 0.07—0.13 mm long. Tail bluntly rounded, with three processes which are 0.003—0.004 mm long. 222 The life cycle was studied by Tikhomirova. She noted that the uterus 231 of mature females is filled with larvae which are, according to their stage of development, 0.12 to 0.14 mm long. Older larvae in the uterus are very mobile. Hatching of larvae takes place periodically, growth in the uterus about 6 days. The larvae enter the muscles with the bloodstream and later the skin of the host, where they form accumulations (usually 5—6, but some- times as many as 50 larvae are visible in a field of vision of the micro- scope). Larvae from the blood differ from larvae from the uterus in their large vacuoles. The larvae from the blood are more resistant, preserving their motility in water for 1—5 hours and for several hours in a drop of saline. Tikhomirova recorded carp lice of the genus Argulus as inter- mediate host,in the body cavity of which she found a larva of Agrachanus scardinii ina study of natural infections. In experimental infection of carp lice, she found that the larvae migrate from the intestine to the body cavity in several minutes. The larvae settle in the legs. Young specimens of Argulus are more susceptible to infection than adults. Usually 2—5, maximum 17 larvae were found in experimentally infected Argulus. During the first days in the body cavity the larvae are active and grow. On the 2nd day in the body cavity they are 0.172 mm long and after 5 days they are twice as large as larvae from the uterus. Two molts were observed in the intermediate host, 4 days and 8—9 days after infection. After they have reached the infective stage, the larvae enter the definitive host when the carp lice feed on fish. Tikhomirova used Scardinia erythrophthal- mus. as definitive host. Riemerences: Molnar, 19666, pe. 150—157. Moravec, 1969, р. 924. Genus Molnaria Moravec, 1968 Synonym: Skrjabillanus _ Schigin and Schigina, 1958 (in part) Diagnosis. Skrjabillanidae. Body threadlike. Cephalic end with 4 submedian tubercles directed anteriorly. Buccal capsule oval, with chiti- nized walls, weakly developed. Buccal cavity practically absent. Three chitinized teeth at the base of the capsule. Esophagus entirely muscular. Females more than twice as large as males. Caudal end of male with 5 pairs of papillae. Gubernaculum and spicules absent. A chitinized plate present. Vulva situated in anterior part of body. Viviparous. Parasites of fish. Type species: Molnaria erythrophthalmi (Molnar, 1966) Moravec, 1968. Molnaria erythrophthalmi (Molnar, 1966) Moravec, 1968 (Figure 140) Sywonym skrmjabillanus erythrophthalm1 Molnar, 19/66 Hoists) Ica Pdi niwis (ery the opiht hadim urs): Localization: mesentery, serosa of intestine. Distribution: Hungary, USSR. 223 Description (after Molnar,1966b). Transparent forms, tapering at 232 both ends. Cuticle smooth. Cephalic end with 4 protruding hemispherical papillae and 2 distinct amphids. Width of cephalic papillae 0.012 mm at the base and 0.008 mm at the apex. Three lips surround the chitinized buccal capsule which are 0.002 mm thick and 0.007—0.009 mm wide. Esophagus cylindrical, 0.29—0.31 mm long. An esophageal gland extends along the eso- phagus to the intestine. Male. Length 3.3—5 mm, width 0.049—0.057 mm. Caudal end with well- developed bursalike wings consisting of longitudinal dorsal and two lateral lobes. The lateral lobes are curved ventrally, forming а sac. The dorsal part of their posterior end is markedly thinner and continues almost to the end of the dorsal lobe. Parallel to the narrow part of the caudal end extend two riblike formations dorsally to the distal end of the lobe. The part bordered by these formations refracts light differently. Two indistinct riblike formations extend laterally from the curved tail, united with the lateral lobes. Tail with 5 pairs of papillae, four pairs pedunculate. The pedunculate papillae of the first and fourth pair are mushroom-shaped. Sessile papillae are situated at the base of the third pair of pedunculate papillae. Two equal, setalike spicules 0.008—0.009 mm long. Cloaca oval, subterminal, Female. Length 11—13.3 mm, width 0.1—0.3 mm. Mature females transparent. Caudal end with 3 oblong teatlike processes 0.008 mm long. One of the caudal processes is ventral, the other two dorsoventral. The end of the dorsoventral papillae is divided by a groove. Anus subterminal. Body cavity occupied entirely by the uterus, which is filled with larvae anteriorly and with eggs posteriorly. Larvae 0.110—0.120 mm long. Young females 0.3—0.9 mm long. At this stage the papillae both at the cephalic and at the caudal end project more markedly. Esophagus not divided, sometimes wider than the buccal capsule, 0.30—0.36 mm long and 0.016—0.018 mm wide. Behind the buccal capsule begins a large esophageal gland which gradually widens distally, increases markedly in size above the valves of the esophagus, and continues along the intestine for twice the ‘length of the esophagus. Another much narrower esophageal gland extends from the middle of the esophagus to the vulva. The esophagus ends in three larve valves at the connection with the widened part of the intestine which becomes narrower and ends in the anus. Vulva situated in anterior part of body, slightly behind the anterior end of the uterus. Vagina thin, short (0.022 mm), indistinct, ovejector markedly widened. The ovejector extends at first posteriorly and then turns anteriorly and widens at the intestinal valves, turns posteriorly and passes into the uterus. Uterus of the same width its entire length, extending posteriorly and then curving and passing into the thin, spirally coiled oviduct, which then passes into the wider ovary. The narrow distal end of the ovary turns posteriorly like a hook. Description (after Tikhomirova). Thin, transparent nematodes. Females more than twice as large as males. Body tapering at both ends. Cuticle smooth. Cephalic end with a small elliptical buccal capsule. Capsule chitinized. Inside the capsule are three chitinous teeth of almost 234the same size. Cephalic end with 4 round cephalic tubercles situated submedially and directed anteriorly. Each tubercle bears two papillae. There are 8 cephalic papillae and two large, rounded amphids. A cuticular ring present between buccal capsule and esophagus. 224 (233) FIGURE 140. Molnaria erythrophthalmi (Molnar, 1966): а — anterior end; b — caudal end of female; с — caudal end of male, ventral; d — cephalic end; e—same, apical; f— caudal end of male, ventral; g — same, lateral; h — caudal end of female; i— larva (a—c — after Molnar, 1966b; d—i — after Tikhomirova). Male. Length 4.5—6.0 mm, width 0.05—0.08 mm. Width of buccal capsule 0.008—0.009 mm, length 0.003 mm. Cephalic tubercles 0.012— 0.014 mm wide and 0.010—0.012 mm high. Esophagus muscular, 0.3—0.4mm long, ending in three valves. Esophageal glands distinct. Caudal wings thin, beginning 0.015 mm before the genital opening. The wings are supported by 5 pairs of papillae, four pairs pedunculate and one pair sessile, situated opposite the third pair of pedunculate papillae. Three postanal pairs and 225 one pair of terminal pedunculate papillae are of different length: the first pair extends to the margin of the wing, the second pair only 2/3 of its width, and the third pair a third of its width; the papillae of the terminal pair are mushroom-shaped and have short stalks. Processes of the hypodermis extend to the wings but are without papillae; they also support the wings when these are extended. The tail may be rounded, pointed, or swollen at the end. Spicules and gubernaculum absent. There is a chitinous plate, which always projects outside. Length of plate 0.006 mm. Its thickness varies. The plate is apparently a chitinization of the lower lip of the cloaca. Female. Length 8.0—16.2 mm, width 0.2—0.4 mm. Cephalic end with 4 well-developed submedian tubercles 0.018—0.021 mm wide and 0.015— 0.018 mm high. Width of buccal capsule 0.0108 mm, length 0.003 mm. Esophagus 0.35—0.5 mm long, of almost the same width throughout. Large esophageal glands present. Intestine ending 0.085 mm from end of tail. Nerve ring situated 0.266 mm from the cephalic end. Tail bluntly rounded, with three processes 0.005—0.007 mm long; two of the processes are bifid. The greater part of the genital duct extends in the long axis of the body. Vulva situated slightly below end of esophagus; vagina short and narrow, ovejector well developed. Uterus filled with larvae 0.07—0.12 mm long. It passes into the oviduct and then into the ovary with thinner walls; the distal end of the ovary is curved in a hook, ; Life cycle (after Tikhomirova), The uterus of mature females is filled with larvae, the length of which depends on their stage of development, 0.07 to 0.12 mm. The intermediate host is Argulus, in the body cavity of which the larvae parasitize. In experimental infections of Ar - gulus the larvae migrated from the intestine into the body cavity in several minutes. They usually become localized in the legs. Two molts were ob- served in the intermediate host, 4 days and 8—9 days after entry into the host. When Argulus feeds оп fish, larvae of the infective stage enter the definitive host: Scardinius erythrophthalmus. References: Molnar, 1966b, pp.150—157; Moravec, 1968, p.322. 235 Molnaria intestinalis (Dogiel and Bychowsky, 1934) Moravec, 1968 ( Figure 141) Synonym: Philometra intestinalis Dogiel and Bychowsky, 1934 Host: Scardinius erythrophthalmus. Localization: intestine. Distribution: USSR, Hungary, Czechoslovakia, Description (after Dogel' and Bykhovskii, 1934). Male unknown. Female. Length 10 mm, width 0.166 mm. Body threadlike. Cephalic end with 4 short, rounded papillae around the mouth. Esophagus straight, its anterior end widened like a funnel; buccal capsule absent. Length of esophagus 0.4 mm. Posterior end of body rounded, with a small tubercle. Anus and vulva absent. Almost the entire body cavity is occupied by the uterus, which is filled with embryos; two ovaries, at opposite ends of the body. Anterior end of uterus extending to 0.5 mm of the cephalic end, posterior end 0.2 mm before the end of the tail. 226 FIGURE 141. Molnaria intestinalis (Dogiel and Bychowsky, 1934): а — general view; b — cephalic end (after Розе!" and Bykhovskii, 1934). References: Agapova, 1966, p.218; Dogel' and Bykhovskii, 1934; pp. 310—311; Molnar, 1966b, p.157; Moravec, 1968, p.322. Molnaria leucisci (Agapova, 1963) Moravec, 1968 иному: Ро 1Тотжеска Leucis e1 Agapova, 1963: Ph. эр. Asapovas 1956 EOS a Lielle LSie@ Uks) 1аи5. Localization: intestine. Dish ei owt von: USSE.. Description (after Agapova, 1963). Male unknown. Female. Length 5.5—6.7 mm, width of cephalic end 0.042—0.045 mm, of posterior end 0.015 mm. Females white, threadlike. Cephalic end with 4 short, rounded, distinct papillae around the mouth, Lips absent. Esopha- gus straight, slightly widened at the anterior end, 0.296—0.359 mm long. 236 Posterior end of body with three distinct processes 0.0076 mm long. Al- most the entire body cavity is occupied by the uterus, which is filled with larvae. Vulva and anus absent. Viviparous. Везетемсез: jpAgapova,, 1956, рр. 5—60; 1963); pp-137—1383 1966, p.218; Molnar, 1966b, p.157; Moravec, 1968, p.322. 227 FAMILY TETANONEMATIDAE SKRJABIN AND SCHIKHOBALOVA, 1948 Historical review Steiner (1937) described Tetanonema strongylurus from the blood sinuses of Bdellostoma heptatrema and established for it the new genus Tetanonema Steiner, 1937, which he provisionally placed in the subfamily Aproctinae. However, Skrjabin and Shikhobalova (1948) did not agree, and because of the presence of papillae irregularly distributed on the whole body, the reduction of the esophagus, which apparently cannot receive food, the large number of papillae on the caudal end of females (Steiner considered them a character of intersexuality but he probably examined both females and males, which he considered as females because of the absence of spicules), localization in the blood of Bdellostoma (Cyclostomata), and other characters, they removed the genus Tetanonema from the subfamily Aproctinae and established the new family Tetanone- matidae Skrjabin and Schikhobalova, 1948, which they placed in the superfamily Dracunculoidea Cameron, 1934, suborder Camallanata Chitwood, 1936 together with the Dracunculidae and Cystoopsidae. Diagnosis. Anguillicoloidea. Body threadlike, with slightly pointed anterior end; posterior end blunt. Eight cephalic papillae inthe outer ring, arranged in pairs. Cuticle of anterior part of body with irregularly arranged tubercles. Intestine reduced. Male. Caudal wings absent. Female. Vulva situated slightly before anterior third of body. Parasites of the circulatory system of Cyclostomata. Type genus: Tetanonema.~ Steiner, 1937. Genus Tetanonema Steiner, 1937 Diagnosis. Tetanonematidae. Body threadlike. Cephalic end slightly pointed, posterior end blunt. Cephalic end with 4 pairs of submedian papillae in the outer ring, inner ring consisting of not always distinct lateral papillae. The region behind the cephalic end has numerous irregularly distributed tubercles. Cuticle with fine intercrossing striation. Esophagus short, re- duced, consisting of barely recognizable duct and wrinkled tissue without musculature around it. Tail blunt, rounded. Male. Caudal wings absent. Female. Vulva situated slightly before anterior third of body. Rectum nearly obliterated. 228 237 Parasites of the circulatory system of Cyclostomata. Type and only species: Tetanonema strongylurus Steiner, 1937. Tetanonema strongylurus Steiner, 1937 (Figure 142) Eros) seid’ liors)tom-yarthe pitialtime mar Localization: subcutaneous blood sinuses. Distribution: Atlantic Ocean. Description (after Steiner, 1937). Body threadlike; cephalic end slightly pointed, caudal end blunt. Cuticle 0.005 mm thick, with intercrossing striation. Cuticle consisting of three layers: surface layer thin, 0.0013 mm thick, staining more strongly with acid carmine than the other two layers. Hight longitudinal fields, the ventrosubmedian and dorsosubmedian fields thin and continuous, distinct only in some places; dorsal field with narrow base, consisting of a simple row of cells. Lateral fields rectangular, multicellular (12—23 cells), with wide base which widens toward the eso- phagus or the intestine. The lateral, dorsal, and ventral fields are more distinct in the region of the esophagus. FIGURE 142. Tetanonema strongylurus Steiner, 1937: a — anterior end; b — cephalic end, lateral; с — same, apical; d = region of vulva, lateral; e — caudal end of female, ventral; f — caudal end of male, lateral (after Steiner, 1937). 229 238 The four submedian pairs of cephalic papillae are weakly developed; they do not protrude above the surface and form the outer ring. A simple interolateral papilla often present in the inner ring. Numerous papillae are irregularly distributed behind the "head'' and also at the level of the posterior part of the esophagus and in some places on the other parts of the body. Amphids small, inthe form of an oval sac, their opening small, rounded, situated near the four submedian cephalic papillae. Mouth narrow. Buccal capsule absent. Lumen of esophagus reduced, probably not absorbing food; esophageal tissue completely reduced in some places, forming compressed, irregular triangular prisms. There are nu- merous nuclei and also apparently three reduced esophageal glands behind the nerve ring. Intestine consisting anteriorly of 18 rectangular cells which gradually pass posteriorly into prismatic cells and increase to 29. A lumen is present only in the anterior part of the intestine, and it is widest dorso- ventrally. Intestinal contents not visible, except anteriorly, where dark co- agulated masses, possibly exudative, are visible. Cells of intestine without distinct membranes, cell content light, finely granular, except for larger dark orange inclusions at the inner wall. Male (immature). Length 166—210 mm. Female. Length 255—275 mm. Rectum more or less obliterated. Uterus amphidelphic. References: Skrjabin and Shikhobalova, 1948, p.100; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.93; Campana- Rouget, 1964, pp.564—565; Steiner, 1937, рр. 681—688. 230 SUPERFAMILY DRACUNCULOIDEA CAMERON, 1934 Historical review In Volume IV of the ''Key to Parasitic Nematodes" Skrjabin et al. (1954) placed three families in the superfamily Dracunculoidea: Dracunculidae, Anguillicolidae, and Tetanonematidae. The latter two families contain only one genus and species, but the family Dracunculidae has four subfamilies: Dracunculinae, Avioserpentinae, Micropleurinae, and Philometrinae. The first 3 subfamilies contain only one genus, but the subfamily Philomet- rinae contains 12 genera. We agree with Baylis and Daubney (1926), who considered all philometrids as a homogeneous group in the family Philo- metridae. Other authors (Travassos, 1960; Yamaguti, 1961b; Rasheed, 1963) placed all philometrids in this family; Yamaguti even established the new order Philometridea in 1961. However, we think that itis more correct to place all these forms in the family Philometridae of the superfamily Dracunculoidea. We place the families Anguillicolidae, Phlyctainophoridae, Skrjabilla- nidae, and Tetanonematidae in the superfamily Anguillicoloidea, and the superfamily Dracunculoidea has therefore the following composition: Superfamily Dracunculoidea Cameron, 1934 Family Dracunculidae Leiper, 1912 Subfamily Dracunculinae Stiles, 1907 Subfamily Avioserpentinae Wehr and Chitwood, 1934 Subfamily Micropleurinae Baylis and Daubney, 1922 Family Philometridae Baylis and Daubney, 1926 Subfamily Philometrinae Yamaguti, 1935 Subfamily Philoneminae n. subfam. Diagnosis. Camallanata. Body thin. Sexual dimorphism sometimes present; males markedly smaller than females, Cephalic end rounded; stoma usually weakly developed. Anus and vulva obliterated in adult females, vulva sometimes weakly marked. Spicules present. Viviparous. Parasites of the connective tissue, body cavity, and closed cavities and organs of mammals, fish, birds, and reptiles. Type family: Dracunculidae Leiper, 1912. Key to the families of the superfamily Dracunculoidea 1 (2). Cephalic end with a cuticular shield... Dracunculidae ILjSnyoreveny WS 2 (1). Cuticular shield at cephalic end absent осо с SON Ce Ol OO COMO nb Circ soa о бла 231 FAMILY DRACUNCULIDAE LEIPER, 1912 Diagnosis. Dracunculoidea. Body long and thin, females markedly longer than males. Cephalic end rounded, with a cuticular shield. Lips or pseudolabia absent. Esophagus consisting of a muscular anterior part and a glandular posterior part, or entirely muscular. Anus and vulva obliterated or rudimentary in adults. Viviparous. Gubernaculum present or absent. Type subfamily: Dracunculinae Stiles, 1907. Key to the subfamilies of the family Dracunculidae 12) Males with only: a right caudal. wing и: « «a8 isn» eae ee И ated-sus.Ariteme sheqel sige Micropleurinae Baylis and Daubney, 1922. 2 (1). Caudal wings absent. 3 (4). Circumoral ring absent on cephalic shield О ae et eae а Avioserpentinae Wehr and Chitwood, 1934. 4.(3)., _Cireumoral ring present on,.cephalic shield. .)...2.. зи ры И С Е Dracunculinae Stiles, 1907. ео он SUBFAMILY DRACUNCULINAE STILES, 1907 Diagnosis. Dracunculidae, with a distinct circumoral ring on the cephalic shield. Inner ring of cephalic papillae with 4—6 papillae, outer ring with 4 double papillae. Esophagus divided into a short muscular part and a long glandular part. Spicules and gubernaculum present. Vulva and anus obliterated in females. Parasites of reptiles and mammals. Туре серая Dracuncudus, Reichard, 1559, Genus Отасипсии$ Reichard, 1759 Synonyms: Furia Linnaeus, 1858; Gordius Linnaeus, 1758 (partim); Vena Gallandat, 1773; Filaria Muller, 1787; Vermiculus Dunglison, 1858; Fuellebornius Leiper, 1926 Historical review Dracunculus is the oldest genus of the suborder Camallanata. 240 Skrjabin and Shul'ts (1940) noted that the "ге snakes'' mentioned in sources for at least 600 years apparently refer to the guinea worm, D. medinensis. 232 Neumann (1895) recorded the species from Python natalensis and named it D.dahomensis. Yorke and Maplestone (1926) considered 16а Us medinenis iis. Turkhud (1920) recorded it from Naia tripudians in India and named iuoracunculus medinens i's . Mackin (1927) recorded it from Chelydra serpentina and named it Dracunculus globocephalus, although he stated that except for several minor details of morphology, size, and host, D. globocephalus and О. medinensis are almost identical. Нза (1933b) described р. houdemeri from Natrix piscator and found that the head is se- parated from the body by a marked constriction. This character was also noted by Mackinin D. globocephalus, obtained from aturtle. Hst mentioned that during a study of Mackin's material, he found that the number and arrangement of cephalic papillae in ШО. globocephalus are the same as in D. houdemeri. He also found the openings of two subventral glands in D. globocephalus inthe same position as in D. houdemeri. Benbrook (1932) recorded D. medinensis from a silver fox (Iowa, USA). He found D.medinensis also ina dog and mink in 1940. Hsu and Watt (1933) recorded the guinea worm in two dogs in Peiping and determined them as О. medinensis. Travassos (1934) recorded D. fuelleborni in an opossum in South America. Mirza and Basir (1937) described D. medinensis from Varanus and noted that Dracunculus fron: man occurs also in animals. Brackett (1938) recorded the guineawormfrom Thamnophis sirta- lis and named it О. ophidensis. He found variation of the species, е. 5.11 the number and arrangement of the genital papillae in males and in the length of the spicules. Goble (1942) recorded О. medinensis,a specimen in each hind leg of Mustela cicognanii from a swamp near Diana (Lewis County, New York State). Chandler (1942) found it in six out of nine raccoons examined and identified itas О. insignis Leidy, 1858. He noted that the arrangement of the papillae resembles that in the des- cription and drawing of Travassos but that the embryos are identical with those of О. medinensis. Chabaud (1960b) found a male in Acranto- phis madagascariensis and described itas Dracunculus doi. Deshmukh (1969) described the new species Dracunculus alii and О. coluberensis. The first species was described from several males in the body cavity and mesentery of Natrix piscator (ШО. houdemeri Hsu, 1933 had already been described from the same host, but Deshmukh did not compare his new species with it); the description of the second species was based on a single male from the lungs of Coluber helena; it is probably a species of Filariata, not Camallanata, and we therefore do not describe this species here, Thus, 10 species of this genus have been described so far, three from mammals and six from reptiles. 241 The systematic position of the genus was determined only recently, al- though the life cycle of D. medinensis was described by Fedchenko backin1872 and much has been published onthe genus Dracunculus since. Some authors place the genus even now in the Filariata. A serious obstacle to the study of the species of this genus is the dif- ficulty in finding males. There are even remarks which doubt the existence of males. The reason is the biology of the parasites; the long females are 233 242 found easily under the skin, but the males have been overlooked. Moorthy (1937) and Onabamiro (1956) mentioned other factors like the early copula- tion of the species, the slow movement of the males after copulation, their localization in the tissues in which copulation takes place and their shorter life span. . An important description of the male of О. medinensis was given in 1937 by Moorthy. Brackett (1938) gave a new generic diagnosis of Dracunculus according to Moorthy's data and his own from a study of D. ophidensis Brackett, 1938, the life cycle of which he described. Comparing the species of the genus Dracunculus, Brackett and Moorthy placed great importance on the caudal armature of the male, considering it as the most important character for differentiation. Desportes (1938) and Chabaud and Campana (1949) were doubtful about this criterion. Chabaud and Campana stated that the differences are not decisive for all dracunculids. Desportes showed the importance of individual variations of the tail of males, and realized that it is impossible to base the diagnosis only on these charac- ters, as Moorthy and Brackett had done. These authors refused to give a classification and diagnosis based on a few characters alone, but that the caudal armature of the male is important emerges from the key to the species of the genus Dracunculus given by Chabaud and Campana, The form of the spicules, gubernaculum and genital papillae have an important place in this key. Chitwood (1933a) identified a form found in North Ameri- can carnivores as D. medinensis, but in 1950, after description of the male, he identified itas D. insignis (Leidy, 1858), noting, however, that this description differs only slightly from the description of Moorthy (1937a) and that when further material has been examined, the characters of the two species may overlap. Chitwood stated that the identification of D.fuelleborni is doubtful. Chandler (1942) stated that it was probably identical with D. insignis. Mirza and Roberts (1957), on the basis of material from the subcutaneous tissue of Natrix sipedon, described structures which had been found in other species of Dracunculus but not mentioned by some authors or had been shown in drawings but not mentioned in the text; they came to the conclusion that the number and arrangement of the genital papillae were the same in their specimens as in D. medinensis, D. houdemeri, D. globocephalus, and О. ophidensis. They also noted that the mouth of males is surrounded by a cuticular crest, from which extend 4 posterior spinelike processes, and that these, according to published data (Hsu, 19335; Travassos, 1934; Brackett, 1938 ; Moorthy, 1938a), are present in the males of Dracunculus but may or may not be present in females. As there are many variations in the same species, the authors concluded that their specimens from Natrix sipedon belong to the species described by Mackin (1927) and Brackett (1938). One male in the material from Natrix sipedon had spicules of al- most the same length; two others had spicules of different length. The spicules cannot therefore be used to distinguish species of Dracunculus. Mirza and Roberts doubt the validity of О. houdemeri, since Hsu (1933a) distinguished his species from the other species because the head is smaller than the body and separated from it byaconstriction. However, this’ cannot be used as a specific character, since it was based on a single female, which may have been damaged or deformed because of age or bad fixation. 234 Mirza (1957) studied a large material from almost all animals recorded as definitive hosts of the species of Dracunculus. Ina study of numerous females of D. medinensis he found that the number and arrangement of the cephalic papillae vary according to the age of the nematode. Mirza concluded that there are only two species, one species which parasitizes mammals and another which parasitizes reptiles. Chitwood (1933b) had stated that wild mammals in North America are infected by Dracunculus medinensis, whichis widely known from man in the Old World. He wrote that the guinea worm occurs only in animals probably because of sanitation conditions in the United States. This could be a physiological variety which hasnot become adapted to man, or the life conditions are the reason that so far no natural infections of man by this parasite have been found in America. In other countries, where conditions are different, this is probably a com- mon parasite of man and only occasionally found in animals. D. medinen- sis has been recorded in dogs and wild animals in the USSR, where the parasite has not been found in man since the guinea worm was eradicated in Middle Asia by improved sanitation. The following authors have recorded this species from animals: Isaev (1934b) from dogs, Petrov and Chertkova (1946, quoted in Murtazaev, 1965) from badgers, Chernyshev (1954) from jackals, badgers, and jungle cats, Dubnitskii (1959) from red fox, Petrov and Chubabriya (1955) from the domestic cat, and Moskvin (1958), Ch'ung-Hsiung (1958), and Murtazaev (1965) from dogs. Воеу (1964) wrote: ''There is no doubt that in Middle Asia and in the south of Kazakhstan there are natural 243 foci of Dracunculus infections where the parasite apparently circulates between carnivores and Cyclops." If it is remembered that in all experiments studying the life cycle of D. medinensis dogs and cats are used, although the intensity of infection is lower, and that most cases of natural infection of dogs have been recorded in areas where the human population is not affected: China, Cairo, Uganda, Argentina, and North America (R. Muller, 1968), Mirza's view (1957) that there is only one species of Dracunculus parasitizing mammals and man is apparently correct. Yamaguti (1961b) established the new genus Chelonidracunculus for Dracunculus globocephalus andthegenus Ophiodracunculus for Dracunculus dahomensis, О. ophidensis, and hypothetically for О. houdemeri. He considered the differences between these genera as so marked that he even placed his new genera in different subfamilies, although he probably knew the studies of Mirza and Mirza and Roberts and was aware of the minor differences between the species of Dracunculus as given by these authors in their descriptions. According to the above, the genus Dracunculus contains at present only two species: D. medinensis (Linnaeus, 1758), a parasite of mam- mals,and D. oesophagea (Polonio,1859),a parasite of reptiles. Dracunculus insignis (Leidy, 1858) and О. fuelleborni Tra- vassos, 1934 are synonyms of Dracunculus medinensis, and О. dahomensis (Neumann,1895), О. ophidensis Brackett, 1938, D. Бщоросерна1 5. Mackin, 1927, О. houdemeri Hsu, 1933, 5.’ diol Chabaud, 1960, and D. alii Deshmukh, 1969 are synonyms of Dracun- culus oesophagea (Polonio, 1859). 235 244 Diagnosis. Dracunculinae. Anterior end with a circumoral elevation which usually projects strongly and is of variable form. Circumoral ring present. Mouth small. Inner ring of cephalic papillae consisting of 4—6 papillae (the interolateral papillae always separated, the interodorsal papillae sometimes partly fused). Outer ring consisting of 4 double papillae. Amphids present behind the lateral papillae of the outer ring. Cervical papillae situated slightly behind the nerve ring. Esophagus consisting of a muscular anterior part and a glandular posterior part. The muscular anterior part is very short. The glandular part is long and divided by a constriction into two parts, a short anterior part and a longer posterior part. Nerve ring surrounding esophagus in the region of its constriction. Intestine straight. Phasmids distinct. Female much larger than male. Male with 10—13 pairs of genital papillae, some of them preanal and some postanal, their arrangement varying. Spicules usually of the same or nearly the same length. Gubernaculum present. Genitalia forming a straight duct; testis and ejaculatory duct indistinctly connected. Tail forming a tight spiral. Vulva situated in anterior part of body, near the cephalic end; vulva atrophied or closed in adult females. Vagina extending anteriorly for a short distance. Two branches of uterus, united in a thin-walled duct be- fore and behind the vulva. In mature females the uterus occupies the entire body cavity. Ovaries straight or curved at the end. Viviparous. Larvae which have emerged in the uterus have a long, thin tail, but their tail is short and bilobed in the infective stage. Adult specimens parasitize in the connective tissue of mammals and reptiles, larvae in Copepoda. Type species: Dracunculus medinensis (Linnaeus, 1758). Dracunculus medinensis (Linnaeus, 1758) {Figures 143, 144) Synonyms: Gordius medinensis L., 1758; Vena medinemeage Gallandat, 1773; Dracunculus graecorum Gruner,1777; Fitaria medinensis Gmelin,1790; Furia vena medinensis Modeer, 1795; Filaria tropica Rudolphi,1809; Е. aethiopica Valenciennes, 1856: Е, insignis» Leidy, 1858; Dracunculwus’ aethi Opa came Diesing, 1861; Filaria guinensis Dunglison, 1893; Vermiculus capillaris Dunglison, 1895; Fuellebornius medinensis Faust, 1929; Dracuneulus insignis (Leidy, 1858); D.° fuelleboris Travassos, 1934 Hosts: definitive — man, Canis familiaris, ‘Procyon le es Putorius vison, Vulpes fulva, Mustela ermineay mmr canadensis, Ondatra zibeticus, Didelphis aurita;) immer mediate'— Cyclops ‘coronatus, С. quadricornis, С. ООО me Vie te Ti wes. Е: Shane 1S Сарата ves еее Macrocyclops fus¢us, Mesocyclops leuckarti; Bucwolmes serratulus, Acanthocyclops bicuspidatus; Tropocyclene prasinus, Orthonoides sp. Localization: connective tissue and subcutaneous fat in definitive hosts; body cavity in intermediate hosts. 236 | . 246 Distribution: Africa, tropical and subtropical Asia, South, Central, and North America. Description (after Moorthy, 1937a). Male. Length 12—29 mm, width 0.4 mm. Mouth small, surrounded by two rings of papillae. Inner ring consisting of 4—6 distinct papillae, the interolateral papillae always distinctly separated, the interodorsal and intero- ventral papillae sometimes partly fused. Outer ring always consisting of 4 double papillae. Amphids situated behind the interolateral papillae. Cer- vical papillae situated slightly behind the nerve ring. Length of esophagus 9.6 mm ina specimen 26 mm long. The esophagus consists of a short muscular anterior part and a long glandular posterior part which is con- stricted near the nerve ring. Anus situated 0.25 mm from end of tail. Ten pairs of caudal papillae, four pairs preanal and six pairs postanal. Counting from the caudal end, the first pair of papillae is ventrolateral, the second and third pairs subventral, not separated, the fourth pair ventral, situated near the third, and the fifth and sixth pairs some distance before the fourth, the fifth pair ventrolateral and the sixth subventral. The four pairs of pre- anal papillae form two rows which begin ventrolaterally and are directed subventrally. The first pair is adanal or situated slightly before the anus; the other three pairs are preanal. Phasmids lateral, situated behind the fifth pair of papillae. Spicules of almost the same length, 0.49—0.73 mm; guber- naculum 0.2 mm long. ‘ Female. Length 1.5—53.0 cm, width 1mm. Rectangular circumoral elevation present. Mouth surrounded by two rings of papillae. Inner ring consisting of 4 papillae; interolateral papillae separated, interodorsal and interoventral papillae partly fused. Outer ring consisting of 4 double papillae. Amphids situated behind the interolateral papillae. Length of esophagus 10mm ш а specimen 24 mm long. Intestine pigmented in young individuals; the pigmentation gradually disappears in old specimens. Anus situated 0.25 mm from end of tail in a female 24 mm long and 0.9mm from this end in a female 50 cm long. The end of the tail of young females has four caudal spines. Vulva situated 10.3 mm from the cephalic end ina female 24 mm long. Description (after Travassos, 1934, as D. fuelleborni). Male. Length 27—29 mm, width 0.29—0.30 mm. Caudal end spirally coiled. Cephalic end with a chitinized shield with two not protruding lips which barely accommodate a distinct pair of papillae. Cephalic shield 0.100—0.117 mm wide. Buccal capsule reduced, esophagus rudimentary. Under the cephalic shield is a chitinized strengthening in the form of a funnel between the buccal cavity and the cuticle. Esophagus 0.57—0.61 mm long, weakly developed, represented in the posterior part by a nontransparent cell membrane which apparently has a glandular function; membrane 0.28 X0.18 mm large. Tail with markedly reduced lateral wings, conical, with distinct, slightly asymmetrical papillae. Papillae arranged on the ventral side as follows: 3—4 pairs preanal, one pair adanal, 4—6 pairs postanal, more or less equally spaced. Cloaca situated 0.38—0.42 mm from end of tail. Spicules of almost the same length, 0.38—0.42 mm long. Gubernaculum weakly chitinized, with indistinct outline, 0.88—0.100 mm long, 0.032 mm wide. Genital duct almost straight, its anterior end folded double, its anterior part situated 0.85 mm from the cephalie end. 237 (245) 0/20 mm FIGURE 148. Dracunculus medinensis (Linnaeus, 1758). Cephalic ends: a—e — after Moorthy, 1937; Е, р — after Skrjabin and Shul'ts, 1931; В — after Chitwood and Chitwood, 1950; i-m — after Travassos, 1934. Caudal ends: n,o — after Moorthy, 1937; q,r — after Chitwood and Chitwood, 1950; s,t — after Travassos, 1934; p — region of cloaca — after Chitwood and Chitwood, 1950. Female. Length 465—490 mm, width 1.5mm. Body cylindrical, cuticle with fine transverse striation. Cephalic end with two small protruding lips and strengthened by a chitinized shield which is slightly yellowish and about 0.400 mm wide. Lips situated in the midline, dorsally and ventrally. The rounded mouth has 3 pairs of papillae: a pair of lateral, a pair of subventral, and a pair of subdorsal papillae. Buccal cavity very small, with a chitinized strengthening between it and the cuticle. Esophagus rudimentary, about 1mm 238 q Te) long, without distinct transition to the intestine. Lateral fields light and transparent, median fields not transparent. Musculature polymyarian, consisting of cells of a pear-shaped protoplasmic body protruding into the cavity. Tail curved ventrally; anus situated 1.2 mm from end of tail. Rectum rudimentary. Genitalia amphidelphic, with diverging straight uteri. Ovaries folded double above the uteri, relatively short and thin. The posterior uterus extends beyond the anus; the anterior uterus ends 20 mm from the cephalic end. The position of the vulva was not determined, but it is probably situated in the middle of the body. The larvae have a primordium of the intestine and a distinct anus. A pair of teatlike processes situated laterally slightly before the anus. Length of larvae 0.300—0.429 mm, width 0.06 mm. Anus situated 0.096 mm from end of tail, papillae 0.100 mm from this end. | Description (after Chitwood, 1950, аз О. insignis). 247 Male. Length 17—22 mm, width 0.240 mm. Length of esophagus 13— 14 mm, of muscular anterior part 0.3 mm. Nerve ring situated 0.62 mm, cervical papillae 0.70 mm, excretory pore 0.80 mm from the cephalic end. Tail 0.35 mm long. Spicules of the same length, 0.46—0.495 mm long; zubernaculum 0.119 mm long. Five pairs of obliquely arranged preanal papillae. The postanal papillae are arranged in a transverse row, two pairs subventral and one pair ventrolateral, then follow two pairs of sub- ventral papillae and behind them paired lateral phasmids. The first study of the life cycle of D. medinensis was published by A. P. Fedchenko (1870—1872), who determined that Cyclops is the intermediate host. Fedchenko studied the larvae in the body cavity of the intermediate host, and described the mode of entry of the larvae of D. medinensis into Cyclops. He thought that the larvae attach themselves to the legs of the crustacean and then penetrate into the body through the abdominal integument. That the larvae actively penetrate into Cyclops was also stated by Blanchard (1890), Manson (1893), and Wenyon (1908). Ikov (1886) stated that the larvae enter Cyclops passively, and this was ac- cepted by Leiper (1907), Roubaud (1913), and Chatton (1918). Massino (1926) observed larvae penetrating into the body cavity of Cyclops through a damaged intersegmental slit. This author acknowled- ged the passive entry of the larvae, but did not deny possible active penetra- tion. The most thorough study of this problem was made by Isaev (1934 a—c), who contributed much to the eradication of dracunculiasis inthe USSR. Isaev (1924—1931) came to the conclusion that the penetration of larvae of D. medinensis into the intermediate host takes place passively, when they are swallowed by Cyclops. Golvan and Lancastre (1968) think that Cyclops deliberately swallow the larvae. Data on the metamorphosis of larvae of D. medinensis inthe intermediate host are very contradictory. Fedchenko (1871) observed larvae in infected Cyclops and established that they became infective after 12 days. Manson (1893) stated that molting of larvae of О. medinensis is delayed for six weeks. Leiper (1907) observed that embryos shed the cuticle after 8 days. After 2 days the larvae lose the delicate transparent membrane; and growth and differentiation of the internal organs take place in the fifth week, when they apparently become mature (after Moorthy, 1938). According to Fairley and Linston (1924), the 239 larvae of О. medinensis become infective about 15 days after infection Ob Gye hor Sy Isaev (1934a) established that the larvae become infective 13 days after infection of the intermediate host and that two molts take place in the body cavity of Cyclops. Manson-Bahr (1935) observed that the larvae shed the cuticle two or three times. Craig and Faust (1937) stated that the metamorphosis of the larvae in the 249 body cavity of Cyclops involves one, possibly two molts and that the lar- vae sometimes become infective after 10—12 days. (248) 005 mm SS FIGURE 144. Dracunculus medinensis (Linnaeus, 1758). Larvae at different stages of development: a — anterior half of first-stage larva, lateral; b — region of phasmids of first-stage larva, ventral; с — cephalic end of first-stage larva, apical; 4 — posterior half of first-stage larva, lateral; e — cephalic end of larva during first molt in Cyclops; f— caudal end of larva during first molt in Cyclops; g — cephalic end of larva during second molt in Cyclops; В — caudal end of larva during second molt in Cyclops (after Moorthy, 1938a). 240 — se. Moorthy and Sweet (1936a, b) gave a brief description of two types of second-stage larvae of D. medinensis, assuming that they are probably different sexes. However, Moorthy (1937a) disclaimed this view. In the same paper he gave a description of the later stages of development of the larvae in Cyclops, noting that two molts take place: after 5—7 days and after 8—12 days at 32—33° and after 8—12 days and after 13—16 days at 12 — 20°. According to Golvan and Lancastre (1968), the first molt in Cyclops takes place after 8 days and the second after 22 days. Infection with larvae of Dracunculus causes retardation of the de- velopment of the intermediate host: infected immature Cyclops do not become mature, and infected adult females do not form egg sacs. With a low intensity of infection (to 4 specimens) all the larvae complete their develop- ment in Cyclops at the same time, ina higher intensity of infection, the first or second molt was observed only in a few larvae. Complete develop- ment of the larvae in the host is protracted for 7—10 days under these con- ditions. Moorthy (1938a) stated that if Cyclops infected with 2—3 larvae are superinfected with a new quantity of larvae 2—3 weeks later, i.e., after the first larvae have completed their development, 60—70% of the Cyclops die in 24 hours. If there are more than 5 larvae in the second infection, they show the same delay in development as in a normal infection. Isaev (1934b) infected dogs experimentally with D. medinensis by feeding them Cyclops infected with larvae of Dracunculus and found that the development of D. medinensis inthe dogs lasted 9—10 months. He noted that the infection of Cyclops and of the dogs was equally suc- cessful when it was carried out with larvae from man and with larvae from experimentally infected dogs. Experiments to infect dogs with Dracunculus were also made by Moorthy and Sweet (1936a, b; 1938) and Onabamiro (1956). They used dogs one month or 1—5 months old. Moorthy and Sweet (1938) stated that all the dogs which were successfully infected had earlier been heavily infected with Acanthocephala. They therefore assumed that such dogs became in- fected more easily than healthy dogs, since ulceration of the intestine facili- tated the penetration of the larvae, or the resistance of the animals was perhaps lowered. Onabamiro (1956) used therefore only month-old dogs in his experiments, since they are usually still not infected with helminths. He used infective larvae taken from Cyclops, or infected Cyclops. The experiments of Moorthy and Sweet (1938) to introduce larvae of D. medin- ensis under the skin of dogs and intravenously were negative. There is no definite knowledge of the development of D. medinensis in the definitive host. It is assumed that after they are released in the in- testine after the Cyclops are digested, the larvae penetrate through the walls of the stomach and intestine and develop further in the mesentery, reaching their final location through the lymphatic system. Supryaga 250(1967a) found for Avioserpens mosgovoyi and Adams (1969) for Philonema oncorhynchi that the parasites move actively through the tissues to their definitive location. Since these species are closely related to О. medinensis, it may be assumed that the migration of D. medin- ensis inthe definitive host is similar. Onabamiro (1956) studied the development of D. medinensis in the definitive host, but could not follow 241 the development of the worms to 40 days after infection; 43 to 48 days after infection the nematodes were found mainly in the region of the axillary and inguinal lymph glands, so that Onabamiro concluded that the larvae migrate first along the lymphatic channels. Miller (1968) assumed that the third molt probably takes place 15 to 22 days after infection. Onabamiro (1956) thought that the third molt takes place during the migration of the larvae into the deeper tissues. He observed the fourth molt of D. medinensis when he dissected a dog 43 days after infection. Males of О. medinensis were not found in experimentally infected animals 6 months after infection. According to Supryaga (1967a), males of A. mosgovoyi may live for two years in the definitive host. Dracunculiasis is an ancient and endemic parasitic disease in tropical and subtropical countries in the Orient. In the opinion of Isaev, it first appeared in Africa as a disease of natural foci of wild animals living near small water bodies. Primary foci of dracunculiasis of man are known from Africa, Arabia, and India. Negro slaves apparently introduced the disease to America, where it became endemic in South America. Isolated cases have been recorded in persons not of local origin in Europe. In the Soviet Union endemic foci of dracunculiasis existed in Uzbekistan, where the entire population of towns and villages was sometimes infected. The people of Bukhara, Dzhizak, Karshi, and Samarkand suffered particu- larly from the disease; the most ancient and persistent focus was in Bukhara. Under the supervision of Isaev, dracunculiasis was eradicated in the USSR 1051931. Dracunculiasis is found in persons of all ages but usually in people 20 to 40 years old. Lesions may be single or multiple; one to 50 or more para- sites may be found ш а patient. The parts of the body affected are those which come most frequently in contact with water, and therefore 86% of all infections are in the legs. Females complete their development in the definitive host in 9—14 months without any outward symptoms of diease. Dracunculiasis is mani- fested clinically only a few hours before local changes of the skin are found, beginning with short-term, sometimes severe prodromal symptoms accom- panied by itching, erythema, and urticaria. Vertigo, syncopes, irritability, even delirium and dyspnea are sometimes present. These prodromal symp- toms continue 2—3 days. Then local symptoms appear in the form of a cha- racteristic inflammatory infiltration. The skin above the cephalic end of the parasite turns red, the inflammatory focus becomes raised, to the size of a walnut, on the apex of which a blister is formed. After sometime the blister bursts and discharges a milky-white fluid containing a large number 251 of larvae. The cephalic end of the parasite projects through the opening and the symptoms of the prodromal period of the disease disappear. The necrotic whitish mass surrounding the cephalic end of the parasite and visible when the blister bursts usually becomes detached of its own accord after 11 days but it may be removed earlier by operation. The dangers of dracunculiasis are its complications: a parasite which penetrates into the joints may cause ankylosis. The disease may develop complications after secondary infections, and abscesses, phlegmonae, gangrene, and also arthritis, synovitis, and epididymo-orchitis may occur. 242 The diagnosis of the disease is easy, since the parasite is localized in the subcutaneous tissue and is often visible under the skin. Treatment. The prodromal symptoms disappear completely after administration of adrenalin. The popular treatment of dracunculiasis, the most ancient and the most effective, is removal of the parasite. The pro- truding cephalic end is carefully drawn out and wound around a piece of gauze so that it cannot be retracted, and the wound is bandaged. This is repeated for several days until the whole worm has been removed; it is not recommended to extract the parasite rapidly, since it may break, and treatment is then more complicated and recovery delayed. In Dzhizak and Samarkand the usual practice is to remove the worm in one operation by opening the skin blister. Epidemiology and prophylaxis. The source of human infection with dracunculiasis are water bodies containing infected Cyclops. In Old Bukhara such water sources were cisterns which were filled with water from irrigation canals. The water from the cisterns was brought to houses by water carriers. Water bodies became infected when ceremonial ablutions were made and water carriers were therefore often infected with D. medinensis. Control of the parasite directed by Prof. Isaev consisted in breaking the contact between the parasite and man, by eradicating Cyclops in water bodies, and in treating patients and improving the water supply. Isaevy suggested the following measures for the control of dracun- culiasis: 1. Detection of infected persons, a census, and rendering the parasites harmless. 2. Elimination of infected Cyclops incisterns; cleaning of all urban cisterns; draining of all water bodies and refilling them with fresh water. 3. A campaign to acquaint the population with measures of control of dracunculiasis through social and individual prophylaxis (a precautionary measure is to drink only boiled water and to wash fruit and vegetables). References: Boev,1964, pp.4—6; Dubnitskii, 1950, pp. 323—328; НЕЧеНЕЮК о, 1870, р. 71; 1872, ppil—= 24311873 Бр. 51—68; Шоу, 1886, р. 1: 155ем.19.26. pp. 12—16: 1930, pp. 51—53 1934а, в. 212; ЭЗАЬ, р. 231; L934 e; р.238; Litvinov, 1968, рр.347—350; Massino, 1926, рр. 639—644; Moskvin, 1958, рр. 124—125; Murtazaev, 1965, рр. 99— 101; Petrov and Chubabriya, 1955, рр. 231—233; Pod''yapol'skaya and Kapustin, 1958, р. 366; Skrjabin and Shikhobalova, 1948, р. 607; Skrjabin and Shul'ts, 1931, р.768; 1940, р. 470; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.64; Bastian, 1863, pp.101—134; Baylis,1939, pp.63—66; Benbrook, 2521932, рр. 821—824; 1940, pp. 260—262; Blanchard, 1890, pp.9—10; Brackett, 1938, pp. 353—361; Chabaud,1960a, рр. 100—103; Chabaud and Campana, 1949, pp.67—76; Chandler, 1941, pp.153—157; pp. 255—268; Chatton, 1918, pp. 338—348; Cheatem and Cook, 1948, pp. 421—423; Chitwood, 1933a, pp. 802—804; 1950, рр.14—15; Cinotti, 1906, pp.466—470; Craig and Faust, 1937, pp.329—332; Deshmukh, 1969, pp.105—110; Desportes, 1938, рр. 305— 326; Dickmans, 1948, pp.39—40; Donges, 1966, pp.252—256; Erickson 1946, рр. 494—509; Ewing and Hibbs, 1966, pp.515—519; Fairley and Linston, 1924, pp.93—103; Farmer and Witter, 1952, p.174; Goble, 1941, p.221; Golvan and Lancastre, 1968; pp.705—710; Helmoldt and Jungherr, 1955, рр. 463—469; Hinz, 1965, pp.90—103; Hsu, 1933, pp.101—118; Hsu and Hoepply, 1931, pp.567—588; Hsu and Watt, 1933, p.1326; Hugghins, 243 1958, рр. 40—46; Kothari, Pardnani, Menta and Anand, 1968, pp. 435—436; Layne, Birkenholz and Griffo, 1960, p.685; Leiper, 1907, рр. 129—132; 1910b, pp. 65—66; 1913, p.265; Leuckart, 1876, pp.109—112; Mackin, 1927, pp. 91—94; Manson, 1893, pp.947—961; Manson-Bahr, 1935, pp. 893—961; Mirza, 1929, pp.129—156; 1957, pp.44—47; Mirza and Basir, 1937, pp. 26— 32; Mirza and Roberts, 1957, pp. 40—43; Moorthy, 1937, pp. 220—224; 1938, рр. 437—460; Moorthy and Sweet, 1936, рр. 437—442; 1936, р. 568; 1938, p.301; Moravec, 1963, pp.378—379; 1966, pp.281—283; Muller, 1966, рр. 431—432; 1967, рр. 747— 750; 1968, pp. 331—338; Neumann, 1895, рр. 123— 127; Onabamiro,1950, ‘p.31; 1954, pi180;° 1956) pp. 157=166; “Rates 1965, p.350; Reddy and Valli,1967, pp.23—25; Roubaud, 1913, pp. 281—288; Schwabe, 1956, p.651; Siegler, 1946, p.179; Strassen, 1907, рр. 110—129; Travassos, 1934, pp.235—238; Turk, 1950, рр.215—216; Turkhud,1920, p. 727; Valenciennes, 1856, p.259; Wenyon, 1908, рр. 432—135; Wilson, 1958, p.256; Yamaguti, 1961, pp.677—678; Yorke and Maplestone, 1926, p. 442. Dracunculus oesophagea (Polonio, 1859) (Figures 145—147) Synonyms: Filaria oesophagea Polonio, 1859; Е. dahomensis (Neumann 1895); Dracunculus dahomensis (Neumann, 1895); D. globocephalus Mackin, 1927; О. houdemeri Hsu, 1939; Doe phagea (Polonio,1859); D. ophidensis Brackett, 1938; О. doi Chabaud, 1960; D. alii Deshmukh, 1969 Hosts: definitive —Thamnophis sirtalis, Natrix sipeguen Wy. pascator, Tropidonotus natrix регва, 1. пе Natrix, Tr. viperinus; Python natalensis, Psa tna sibilans, Acrantopsis madagascariensis, Chely@paaaea. pentina;. intermediate — Cyclops viridis; reservoirs — вена backs. Localization: connective tissue in definitive hosts, body cavity in intermediate hosts and reservoirs. Distribution: North America, Indochina, Africa, Madagascar, Italy. Description (after Neumann,1895, as D. dahomensis). Male. Length 4.8 cm, width0.34 mm. Body gray, tapering only poster- iorly. Tail forming a spiral of four coils. Cuticle 0.003—0.004 mm thick, with marked longitudinal striation; transverse striation fine and dense, with intervals of 0.0015 mm, Lateral lines wide, each of them occupying about 1/5 of the circumference of the body, Cephalic end with 8 papillae. Tail conical, with recurved end. Spicules of almost the same length, 0.400—0.425 mm. One pair of preanal papillae, two pairs of postanal papillae, the last pair near the end of the tail. Female. Length 50—80 cm, width 1—1.25 mm. Cuticle 0.006—0.010mm thick, transparent, homogeneous, with transverse striation with intervals of 0.0025—0.006 mm, Lateral lines very wide, occupying a quarter of the circumference of the body, consisting of a granular substance in which are scattered rounded or oval cells, 0.018—0.022 mm wide. Median lines dif- 254 ficult to see, even in microscopic sections. Muscles arranged as in О. medinensis. 244 (253) > $< BRK 2/00 mm 2050 тата FIGURE 145. Dracunculus oesophagea (Polonio, 1859). Cephalic end, different aspects: b—d — after Mackin, 1927; a, $ —i — after Chabaud, 1960; j,1,t—w — after Hsti, 1933b; К, ш — after Brackett, 1938; п-г — after Desportes, 1938 ев after Deshmukh, 1969. 245 255 Cephalic end slightly narrowing anteriorly, mouth surrounded by a ring- shaped lip. A thick, rounded sensory papilla, slightly depressed at the apex, on each median line. Between these two papillae are two others, one right and one left, which are less high and more conical, their apex usually trun- cate and divided into two or three secondary processes. Between each median papilla and each of the two lateral papillae but slightly farther pos- teriorly is another papilla resembling the latter papillae. Mouth surrounded by 8 papillae, in an arrangement as in О. medinensis. A distinct cephalic shield absent. Tail conical, curved ventrally, ending in a short teatlike process. Anus probably situated near end of tail; intestine reduced, a posterior opening probably absent. The intestine is flattened and displaced toward the uterus, which is filled with embryos. Embryos 0.400—0.425 mm long and 0.012—0.015 mm wide. Cephalic end curved ventrally, slightly tapering and with a plane surface in the middle of which the buccal funnel is situated. The body becomes conical in its poste- rior third and ends in a strongly tapering whiplike tail about 0.100 mm long. Cuticle 0.00175—0.002 mm thick, with very fine, indistinct striation with intervals of 0.00175 mm; striation not recognizable on the tail. Description (after Mackin,1927,as О. globocephalus). Body threadlike, of uniform width throughout. Posterior part of body slightly tapering, ending in a conical spine in both sexes. A small constriction around the cephalic end appears like a neck. Anterior part bluntly rounded, without lips. Eight cephalic papillae symmetrically arranged around the mouth; the largest pair situated dorsoventrally, and the ends of the papillae are directed anteriorly. The next largest pair is lateral, and each papilla of this pair is double, cone-shaped, and contains two nerve endings; the other four papillae are submedian. A pair of cervical papillae about 0.9 mm from the cephalic end on the lateral lines. Lateral fields wide, occupying 1/8 to 1/6 of the circumference of the body. They are not recognizable in total preparations. Median lines markedly reduced. Male. Length 16—20 mm, width 0.170—0.220 mm. Caudal papillae .consisting of a small preanal papilla and a pair of cup-shaped postanal papillae situated ventrolaterally, one before the other. Tail in fixed speci- mens coiled into a spiral of one to four loops. Genitalia forming a straight duct which begins a quarter of the length of the body from the cephalic end and extends posteriorly to the cloaca. The testis passes gradually into the vas deferens, which fills almost the entire body cavity at the cloaca and then becomes abruptly narrower. Right spicule in the form of a long, narrow needle without wings, 0.8 mm long and 0.005 mm thick, always protruding from the genital cone for more than a third of its length. Left spicule 0.2 mm long and 0.010 mm thick, its proximal third forming a hollow tube widening at the end and the two distal thirds in the form of two tubules connected laterally, the left side united with the proximal end and with the right branch. These two branches become fused into a thin process distally. This spicule was extruded in most specimens. Numerous unicellular glands behind and along the sides of the cloaca, their ducts opening into the cloaca. Female. Length variable, 30 to 133 mm, width 0.280 to 0.680 mm. The females apparently increase in size as the embryos develop. The intestinal tract consists of the short muscular part of the esophagus, a much longer glandular part, and the intestine. Buccal cavity or pharynx absent. Mouth more or less triangular, opening directly into the esophagus. 246 At the connection of the muscular and the glandular part of the esophagus are two wide glands, appearing like two large sacs attached to the outer side. These sacs are partly hollow. Ducts connect the esophagus withthe glands. A third, larger, gland is situated at the opposite side of the esopha- gus, extending posteriorly from a point 0.5 mm from the cephalic end and ending at the connection between the glandular part of the esophagus and the intestine. These three glands are very similar: they have a large nucleus, which is more than 3 mm long in the largest gland. The lumen of the pos- terior part of the esophagus is almost closed in its greater part and forms a narrow slit. Intestine ending blind in mature females. The posterior end of the intestine in the curve of the tail is attached to the ventral body wall by a band of tissue. Excretory pore situated 1—1.5 mm from the cephalic end, according to the size of the specimen. Two short ovaries, one at each end of the body; one is wound around the glandular part of the esophagus and extends anteriorly 2 mm from the cephalic end, where it passes into the anterior uterus. The posterior ovary is wound around the posterior end of the intestine and extends to the anus. Fertilization apparently takes place throughout the uterus. The entire body cavity is occupied by the uteri, which are more or less filled with de- veloping embryos. The wall of the uterus consists of two layers, a thin outer layer and a nuclear inner layer about 0.003 mm thick. Vulva situated slightly behind middle of body. Vagina at first directed dorsally toward the left lateral line and then turning anteriorly into the angle between the intestine and the uterus: Vagina and vulva atrophied in females with fully developed larvae; only a short part of the vagina, connec- ted with the vulva, is present. Description (after Hst,1933b,as О. houdemeri). Cuticle with- out transverse striation. Cephalic end narrowing and separated from the body by a constriction. Mouth oval, surrounded by two rudimentary lips which are hemispherical in apical view and triangular in lateral view; they project dorsally and ventrally and have a hemispherical outer margin. Finger-shaped appendages are present on each side of the cephalic end. Lateral amphids are visible in apical view, one dorsal and one ventral, and 256two lateral and four submedian papillae. The dorsal and the ventral papilla are small, weakly protruding, and with double nerve endings. The mouth is followed directly by the muscular part of the esophagus. Its anterior part 257 15 enclosed in cuticular outer integument which extends dorsoventrally. The muscular part of the esophagus is very short and is followed by a long glandular part which begins with a widening of the dorsal esophageal gland and then continues into a narrower part. Width of esophagus 0.290 mm. Intestine 0.070 mm wide, distinctly demarcated from the esophagus. Nerve ring situated immediately behind the anterior widened part of the glandular part of the esophagus. Female. Length 213 mm, width 0.85 mm. Nerve ring situated 0.93 mm from the cephalic end. Length of esophagus 48.10 mm, muscular part 0.38 mm long, glandular part 47.72 mm. Anterior widened part of glandular part of esophagus 0.05 mm long. Rectum 0.011 mm long. Intestine short, 0.5 mm. Tail conical, pointed,0.28 mm long. A pair of lateral papillae оп the tail. Anus recognizable, but very small. Two thin ovaries; anterior part of anterior ovary beginning about 2.1mm from the cephalic end, about 7.1mm long; posterior end of posterior ovary beginning 10.44 mm from end 247 of tail, about 7.75 mm long. Vagina not recognizable. Branches of uterus containing developed larvae, occupying the greater part of the body cavity. Larvae in the uterus 0.330—0.363 mm long, 0.015—0.018 mm wide; tail 0.28 mm long. (256) FIGURE 146. Dracunculus oesophagea (Polonio, 1859). Caudal ends: a — after Moorthy, 1937; b,c — after Brackett, 1938; d,e — after Mackin, 1927; f-h — after Desportes, 1938; k,1 — after Chabaud, 1960b; m, п — after Deshmukh, 1969. Spicules: i — after Desportes, 1938; j — after Hsu, 1933b. Description (after Chabaud,1960b, as D. doi). Male. Length 28 mm, width 0.250 mm, Tail with three coils of a spiral. Cuticle thin and transparent, with transverse striation with intervals of 248 0.0025 mm. Lateral fields 0.070 mm wide. Cephalic end rounded, with two small lateral processes. Mouth small and rounded, surrounded by an inner ring of 6 large papillae: two single ventral papillae and two double dorsal ‚ рарШае. Outer ring consisting of 8 large papillae arranged in pairs and two amphids. Behind the mouth is a chitinized buccal capsule about 0.040 mm long in the form of a cap which envelops the anterior part of the esophagus. The cross section of the buccal capsule is pentagonal, with one dorsal, two lateral, and two lateroventral sides. At the same level is situated the transition from the triradial symmetry of the esophagus (one dorsal lobe and two lateroventral lobes) to bilateral symmetry. Muscular part of esophagus 0.330 mm long, glandular part with a markedly widened anterior part 0.300 mm long. Excretory pore situated 0.700 mm, nerve ring 0.710 mm, deirids 0.800 mm from the cephalic end. The first postanal pair of papillae is reduced; the second postanal pair is situated distinctly posteriorly; the terminal subdorsal pair is bifid; each of the group of papillae on the subventral line is situated on a small cuti- cular ring. The 8 small preanal papillae are arranged in two small longi- tudinal rows. Tail 0.240 mm long. Spicules simple, ending in a point, about 0.460 mm long. Gubernaculum weakly chitinized, of irregular form, 0.130 mm long. Description (after Brackett, 1938,as О. ophidensis). Mouth 258 surrounded by an inner ring of 6 papillae, the inner lateral papillae most strongly protruding, and an outer ring of double papillae. Esophagus di- vided into a short muscular part and a long glandular part. Nerve ring surrounding esophagus at the constriction. Cervical papillae situated behind the nerve ring. 259 Male. Length 16 mm, width 0.17—0.20 mm. Length of esophagus 7.7 mm. Nine pairs of caudal papillae, 5 pairs preanal and 4 pairs postanal. The first and second pairs are subventral, the third and fourth pairs ventral, and the fifth to ninth pairs are arranged around the anus. Tail 0.17 mm long. Spi- cules of almost the same length, 0.554 and 0.523 mm. Anterior end of testis 13.8 mm long, ending 2.7 mm before the cephalic end. Female. Length to 250 mm. Vulva situated in middle or behind middle of body. In a specimen 24.6 mm long, the esophagus was 9.9mm long, the anterior ovary 5.1mm, and the vagina 1.4mm long. Vulva situated 15.1 mm from the cephalic end. Viviparous. The larvae are released through the duct of the uterus and through the body wall at the cephalic end. First-stage larva: length 0.43—0.45 mm, width 0.016 mm. Cuticle striated; tail long and thin. pecond-stage larva: length 0.62—0.65 mm. Cuticle smooth; tail short and with lobes situated laterally; several unicellular glands at the cephalic end and around the rectum and tail. Description (after Deshmukh, 1969, as D. alii). Cylindrical, milky white nematodes. Cephalic end blunt and rounded, tail curved ventrally, ending in a point. Cuticle smooth. Body uniformly wide except for the tail. Mouth surrounded by a cuticularrim. Two rings of papillae, an outer and an inner,visible in apical view. The two pairs of outer papillae are sub- median, of complicated structure. The inner papillae are simple, arranged in three pairs. Amphids situated laterally. 249 (258) FIGURE 147. Dracunculus oesophagea (Polonio, 1859): а — larva, general view, lateral; b — abnormal process on the tail of a larva; с — posterior end of a 14-day-old larva, lateral; 4 — Cyclops with larvae 14 days after experimental infection; e — first-stage larva, general view (a—d — after Desportes, 1938; e — after Mackin, 1927). Male. Length 13.09—24.40 mm, maximum width 0.17—0.28 mm, Width of cephalic end 0.03—0.04 mm. Nerve ring situated 0.34—0.58 mm from the cephalic end. Esophagus consisting of a muscular and a glandular part, the posterior part much wider than the anterior. Length 0.17—0.28 and 0.18— 0.29 mm, respectively. Tail curved ventrally, without caudal wings. Cloaca situated 0.15—0.24mm from end of tail. Twelve pairs of caudal papillae. Six pairs preanal, one pair situated slightly before the cloaca. Four pairs of papillae 250 arranged in two groups on each side of the cloaca. The papillae diverge from the medioventral to the ventrolateral position. A further pair of pa- pillae is situated slightly before these groups and lies medioventrally. Six pairs of postanal papillae: one pair medioventral, situated slightly behind the cloaca, the second pair at the first third of the tail, situated laterally, two further medioventral pairs, situated together, between the second pair and the last two pairs, and the last two pairs situated laterally close together. Spicules of similar form and of almost the same length, 0.23—0.30 and 0.22—0.29 mm long. Gubernaculum distinctly divided into two branches 0.05—0.07 mm long and 0.01—0.02 mm wide. Description (after Desportes, 1938, аз D.oesophagea). Body cylindrical, tapering at both ends. Cuticle with fine transverse striation with intervals of 0.0015 mm. Cephalic end hemispherical, with a round cuticular shield which projects less in young specimens. Mouth with a single lip, almost rounded. Lumen of muscular part of esophagus triradial, 260 one of the sectors ventral; the cuticle surrounding it strongly thickened, forming a ring which is wide in large specimens. The cephalic shield bears 8 papillae in two concentric rings. The first ring consists of four papillae, two median and two lateral; the papillae are low, conical, inclined toward the mouth. The outer ring consists of four large, kidney-shaped, bifid papillae. Amphids present. Cervical papillae situated slightly behind the nerve ring, 0.91 mm from the cephalic end in a female 60 mm long. Male. Length 11.7—20 mm, width 0.2 mm. Cervical papillae situated 0.65 mm from the cephalic end in a specimen 14.5 mm long, slightly behind the nerve ring. Length of esophagus 9—10.2 mm, muscular anterior part 0.2 mm long. Tail pointed, twisted, with a variable arrangement of papillae. Five pairs of preanal papillae form two lines curved in opposite directions. A pair of large papillae is situated behind the cloaca. Then follow 4 further pairs of distinct postanal papillae. The first of these pairs consists of two swollen ventrolateral papillae, slightly behind and lateral to which are the openings of the phasmids. The following two pairs of papillae are usually situated close together. The last pair of papillae, which are small and ventrolateral, is situated near the end of the tail. Caudal wings absent. Genitalia represented by a long testis, which at first surrounds the esopha- gus and continues into the swollen ejaculatory duct. Spicules of almost the same length: left spicule 0.283—0.29 mm long, right spicule 0.297—0.30 mm. Gubernaculum weakly chitinized, pyramidal, 0.065 mm long. Female. Length 280—437 mm, width 0.3—0.8 mm. Length of esophagus 12—23 mm. Cephalic shield protruding, distinct in lateral view. Circumoral ring distinct. Lateral papillae high, median papillae sometimes double. Cervical papillae situated behind the nerve ring, 1 mm from the cephalic end. Genitalia consisting of two branches of the uterus fused into a single duct, which tills the entire body cavity. The vulva disappears in mature females; only an indistinct spot is visible 182 mm from the cephalic end. Large females are filled with eggs or with motile larvae. Larvae. Length 0.475 mm, width 0.018 mm, body tapering at the cephalic and particularly at the posterior end, ending in a long tail. Cuticle 0.0017 mm thick, with fine transverse striation with intervals of 0.0016 mm. Distinct papillae present at the anterior end: lateral papillae low, submedian papillae comma-shaped. Dorsally behind the mouth is a distinct spine. Nerve ring 251 261 situated 0.065 mm from the cephalic end; а thin duct passes ventrally, ending in the excretory pore. Length of esophagus 0.115 mm, its lumen wide and with hyaline content. Length of intestine 0.160 mm. Rectum a 0.022 mm long, anus situated ventrally, 0.3 mm from the cephalic end. . Saclike phasmids situated laterally in posterior third of Боду, 0.012 mm behind the anus. The tail forms about the last third of the body, its end about 0.065 mm long. Mackin (1927) noted that larvae from the uterus lived for about 7 days in saline, but were sluggish. If kept in tapwater or pond water, the larvae were very active for a few days after the females i had died. Their activity decreased after 9—10 days, and all died after 12 days. Mackin fed numerous larvae soon after they emerged from the female to various Crustacea: Ostracoda, Copepoda, Cladocera. Cyclops bicuspidatus readily swallowed the larvae, which rapidly passed through the wall of the intestine into the body cavity. Other crustaceans did not become infected. Cyclops did not suffer from the presence of the nema- todes, but if there was a severe infection, the intestine ruptured and the Cyclops died. Such a severe infection probably occurs rarely, if at all in nature. : Brackett (1938) also found that larvae from mature females, after entering the water, are in constant motion, remain suspended without sinking to the bottom and remain active for several days. First-stage larvae are cha- racterized by the very long, thin tail. The larvae develop in Cyclops vi- ridis. Twelve to 15 days after entering Cyclops, the larvae become infective; they now lose the long, thin tail and grow to a length of 0.62 mm, Their activity decreases but the larvae preserve mobility. The reservoir hosts may be sticklebacks, in which the larvae are present free in the body cavity. Brackett did not observe further development or growth in sticklebacks. Infection of the definitive host, a snake, takes place when it swallows infected Cyclops. Desportes (1938) mentioned that the larvae are very active in water in the first few hours. They develop in Macrocy- clops fuscus. The first molt takes place after 10 days, and the second- stage larva has a very short tail. After 14 days the larvae are 0.510 mm long and the tail 0.040 mm; four pointed appendages are present on the tail. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.69; Brackett,1938, pp.353—361; Chabaud, 1960a, рр. 100—103; Deshmukh, 1969, рр.105—110; Desportes, 1938, pp. 305—326; Hsu, 1933, pp.101—118; Mackin, 1927, pp.91—94; Mirza, 1929, рр. 129—156; 1957, рр. 44—47; Mirza and Basir, 1938, pp.26—32; Mirza and Roberts, 1957, pp.40—43; Moravec, 1963, pp.378—379; 1966; рр. 281—283; Neumann, 1895, pp.123—127; Travassos, 1934, рр. 235—237. SUBFAMILY AVIOSERPENTINAE WEHR AND CHITWOOD, 1934 EMEND. CHITWOOD, 1935 Synonym: Avioserpensinae Wehr and Chitwood, 1934 Historical review The subfamily was established in 1934, although the authors only had fragments of females. The authors established a new genus and the new 252 a subfamily, Avioserpensinae, because of the absence of the circumoral chitinized ring which is characteristic for Dracunculus, to which the new species is closely related. Chitwood (1935) changed the name Avioserpensinae to Avioserpentinae Wehr and Chitwood, 1934, but did not change the diagnosis. Yamaguti (1961b) recognized the name Avioserpentinae Chitwood, 1935 and made the name Avioserpensinae a synonym. 262 Skrjabin, Shikhobalova, Sobolev, Paramonov and Sudarikov (1954) disregarded the change of name and named the subfamily Avioserpensinae Wehr and Chitwood, 1934. To avoid confusion, Supryaga (1967a) suggested using the name Avioserpentinae Wehr and Chitwood, 1934, emend. Chitwood, 1935. Diagnosis. Dracunculidae. Cephalic end with 14 papillae arranged in two concentric rings. Amphids situated between the rings. Cephalic shield present, but a distinct circumoral ring absent. Cervical papillae oblong. Esophagus consisting of a muscular anterior and a glandular posterior part. Anterior part markedly shorter than posterior part. Spicules well developed. Gubernaculum strongly chitinized. Anus and vulva absent in the adult female. Viviparous. Type genus: Avioserpens Wehr and Chitwood, 1934. Genus Avioserpens Wehr and Chitwood, 1934 Synonym: Oshimaia Sugimoto, 1934 Historical review The genus Avioserpens was established for a species from the subcutaneous tissue of the neck andthe oral mucosaof Egretta garzetta in North America; the authors named it AVioserpens denticulophas- ma. Although they had only fragments of females 590.0 mm long (a cephalic and two caudal ends), the authors established for the genus the new subfamily Avioserpensinae. Wehr (1934) recorded Anas fuligula from Florida as the definitive host of this species. Sugimoto (19340) established the genus Oshimaia for the species OsShimaia taiwana, which he had described in 1919 from a submaxillary tumor and the subcutaneous tissue of the femur of ducklings in Taiwan. Sugimoto placed the new genus in the family Philometridae. In 1935, apparently because of Wehr's statement that the parasite from A. fuligula is identical with Avioserpens denti- culophasma from Egretta, Chitwood made A. denticulophasma а Synonym of Avioserpens taiwana (= Oshimaia taiwana (Sugimoto, 1919)) without a differential diagnosis, and made Aviose rpens denticulophasma andthe genus Oshimaia Sugimoto, 1934 syno- nyms of Avioserpens taiwana. However, the names Oshimaia taiwana (Sugimoto, 1919) and Oshimaia Sugimoto, 1934 appeared in the literature independently until 1950. Chabaud, Campana, and Truong-Tang-Ngok (1950) described nematodes from tumors in the submaxillary region of young domestic ducks from 253 263 South Vietnam which they named Oshimaia taiwana (Sugimoto, 1919). After comparison of these forms with Avioserpens galliardi Chabaud and Campana, 1949, the authors concluded that they belong to the same genus. Chabaud et al. (1950) established that the genera Avioserpens Wehr and Chitwood, 1934 and Oshima ia Sugimoto, 1934 are identical and that they belong to the family Dracunculidae Leiper, 1912. Since the genus Avioserpens was described before the genus Oshimaia, the name Avioserpens Wehr and Chitwood, 1934 has priority. Although Chabaud, Campana, and Truong-Tang-Ngok (1950) had come to the same conclusion as Chitwood (1935), later authors (Skrjabin et al., 1954; Yamaguti, 1961b) quoted only Chabaud et al. (1950) on the identity of the general Avioserpens and Oshimaia. The authors designated A. denticulophasma as the type species of the genus Avioserpens Wehr and Chitwood,1934, retaining A. taiwana in the genus among other species. Yamaguti (1961b), however, stated that А. denticulophasma 15а зупопут of А. taiwana. They apparently considered Chitwood's conclusion that the two species are identical as not sufficiently substantiated. After an analysis of the composition of the genus Avioserpens. and of the morphology of the parasite, the definitive hosts, the localization, and the geographical distribution, Supryaga (1967a) agreed with Chitwood that А. denticulophasma and A.taiwana are identical and made A. denticulophasma asynonym of A. taiwana. Chabaud and Campana (1949) described Avioserpens galliardi, which they found in an aponeurosis of the upper part of the esophagus of Egretta garzetta inthe Eastern Pyrenees. The species was des- cribed from one male and one female. Singh (1949b) described Avioserpens multipapillosa from the subcutaneous tissue of the neck of Ardeola grayi from India. He had only one female. Singh did not know the description of A. galliardi Chabaud and Campana, 1949, and he therefore compared A. multipapillosa only with A. denticulophasma and A, taiwana. Hedistinguish=-d A. multipapillosa from А. taiwana by its smaller size and by the ventrally curved tail. However, the length of the female of A. multipapillosa cannot be used as a specific character because the specimen was immature, and a curved tail is present also in other species of Avioserpens. Supryaga (1967b) reviewed the descriptions of A. galliardi and A. multipapillosa and because of the absence of distinct differences between these species, their similar localization (subcutaneous tissue of the neck), the same definitive hosts, and the wide distribution of А. galliardi in many Ciconiiformes in Europe and Asia, concluded that A. galliardi andA. multipapil- losa are identical. The descriptions of both species were published in 1949, but Chabaud and Campana's study appeared a few months earlier, and the authors had males, whileSingh had described his species from a single immature female. Supryaga (1967b) therefore suggested retaining А. galliardi and making А. multipapillosa a synonym of it. His arguments seem convincing. Avioserpens bifidus was described by Olsen (1952a) from Clangula islandica in North America. The 264 nematodes were found at the base of the tongue, only fragments of mature females (one cephalic end and two caudal ends)590 mm long; a male was not 254 ] found. Olsen distinguished his species from А. denticulophasma and A. multipapillosa by the form of the mouth, which is a trans- verse slitin A. bifidus, while it is rounded in the other species, and also by the form of the phasmids and the size of the larvae. After examination of the form of the mouth ш A. galliardi and A. mosgovoyi, Supryaga (1967b) found that this is not a constant specific character; Turemuratov (1965) gave a drawing of the cephalic end of A. galliardi, in which the mouth is shown as a transverse slit. Supryaga (1967b) concluded that the characters used by Olsen are not sufficient for the establishment of a new species. Furthermore, Olsen did not compare his species with the descriptions of A. galliardi and А. taiwana. The size of the larvae of A. bifidus can also not be used as a specific character, since the length of the larvae does not differ much from that of other species of the genus. Because of the above, the localization and the definitive host, and the almost identical size of the females of А. bifidus and А. taiwana, Supryaga (1967b) concluded that they are identical; А. taiwana was published first and зо ре made A. bifidus a synonym of it. Mawson (1957) described a male and a female of a new species Avioserpens, A. nana, found in the subcutaneous tissue of Ardea herodias from Canada. He distinguished it from other species of the genus by the smaller size of the female. However, since this female was immature, its length cannot be used as a specific character. Supryaya (1967b) stated that because of the localization (subcutaneous tissue of the neck), its definitive host (Ardea), and its geographical distribution, A. папа is closely related to A. galliardi. The length of the body, tail, and gubernaculum is very similar in the two species. Supryaga (1967b) considered as a specific difference the different length of the spicules in the two species: 0.240 mm in A. nana and 0.330—0.446 mm in А. galliardi although Mawson's male specimen was probably immature. Mawson did not mention the pigmentation or the length of the intestine of the male of A. nana, according to which its age could have been determined. Supryaga assumed that the validity of A. nana is doubtful and that А. nana is closely related to А. galliardi. We agree with his arguments and consider Avioserpens nana as a synonym of и ата rid. During a study of the helminth fauna of birds in the Azov estuaries in the Krasnodar Territory in 1965—1967, Supryaga found in 12 out of 148 Fulica atra specimens of Avioserpens which he described as Avioserpens mosgovoyi Supriaga, 1965. A. mosgovoyi differs distinctly from A. galliardi inthe structure of the gubernaculum, the form and smaller size of the spicules, and in its hosts (A. mosgo- voyi parasitizes in Rallidae and Colymbiformes,A. galliardi in Ciconiiformes). In its localization, A. mosgovoyi closely resembles 265A. taiwana, a parasite of domestic ducks in South Vietnam, which also causes tumors in the submaxillary region. However, according to experi- mental data, (3 five-day-old birds became infected out of 12 mallards of different age), Supryaga considered the mallard as only a facultative host ofA mos fSoOv oO yi. In the original description (1965a) the author gives only minor charac- ters which distinguish his species from A. taiwana: the greater length 255 266 of the larvae (0.448—0.500 пт in А. mosgovoyi апа 0.420 mm in A. taiwana) and the greater size of the female (88.0 cmin A. mos- govoyi and19.5cmin А. taiwana). Supryaga (1967b) stated that ''the length of the body is apparently a definite character for the distinction of species of the genus. However, the length of females of A. mosgovoyi increases 30—45 times from their fertilization to maturation of the larvae inthe uterus and their discharge. A comparison of the size of first-stage larvae of the species of Avioserpens shows that their length cannot be used for the distinction of species of the genus." In view of the above, A. mosgovoyi is probably not a valid species, but the author thinks that only comparison of material from Vietnam and the USSR will make it possible to decide this question, and we therefore consider A. mosgovoyi provisionally as a valid species. Yamaguti (1961b) recognized six species in the genus Avioserpens: the type species A. denticulophasma Wehr and Chitwood, 1934, A. bifidus Olsen, 1952, A. galliardi Chabaud and Campana, 1949, A. multipapillosa Singh, 1949, A. nana Mawson, 1957, and A. taiwana (Sugimoto, 1919), We recognize only three species in the genus Avioserpens: A. palliarai, В, taiwana, and A. mos govoy4. Diagnosis. Avioserpentinae. Cephalic end with a shield, without a distinct circumoral chitinized ring. Fourteen pairs of cephalic papillae arranged in two concentric rings. Inner ring with two interolateral papillae; outer ring with four dorsoventral double papillae and eight paired ventral, lateroventral, and dorsal papillae. Amphids situated between the two rings of papillae, immediately behind the interolateral papillae. Mouth round or oval. Esophagus consisting of a short muscular anterior part and a long glandular posterior part. The glandular part has a swollen anterior part separated from the other part by the nerve ring. Cervical papillae weakly projecting. Intestine tubular. Mature females several tens of times larger than males. Male. Genital papillae postanal. Tail curved ventrally. Spicules ` of equal length, strongly chitinized. The narrowed distal end of the guber- naculum has spines on the lower side; gubernaculum chitinized. Genital duct single; testis and ejaculatory duct gradually merging. Female. Vulva situated slightly behind middle of body, atrophied in adults. Two uteri, amphidelphic in adults and sometimes opistodelphic in young females. In mature females the uterus is saclike and fills the entire body cavity. The larvae in the uterus of mature females havea long, awl-shaped tail. Infective larvae have a short tail with three spines. Mature specimens parasitize in the subcutaneous tissue or in an aponeurosis in the upper part of the esophagus of birds. The larvae de- velop in the body cavity of Copepoda. Reservoir hosts sometimes have a part in the life cycle. Type species: Avioserpens taiwana (Sugimoto, 1919) Chitwood, 1935. Avioserpens taiwana (Sugimoto, 1919) Chitwood, 1935 (Figure 148) Synonyms: Filaria taiwana Sugimoto, 1919; Oshimaia taiwa- na (Sugimoto, 1919); Avioserpens denticulophasma Wehr and Chitwood, 1934; A. bifidus Olsen, 1952 256 : Hoscis: ‘Anas platyrhyncha, A. fuligula, “Anhinga anhinga, Clangula (= Glaucionetta) islandica. Localization: subcutaneous tissue; tumors under the lower jaw and under the mucosa of mouth and pharynx. Distribution: Taiwan, Vietnam, USA. Description (after Sugimoto, 1934a, as Oshimaia taiwana). Male unknown. Female. Length 100—240 mm, width 0.8 mm. Cuticle smooth. A short, narrow vestibule between esophagus and mouth. Length of cylind- rical part of esophagus 1.3mm. Nerve ring surrounding posterior part of esophagus 1 mm from the cephalic end. Anus atrophied. Embryos thin, white, 0.390—0.420 mm long and 0.156—0.020 mm wide. After passing into the outer environment the embryos are 0.420—0.510 mm long and 0.021 mm wide. Tail long. 0079 mm 005mm а FIGURE 148. Avioserpens taiwana (Sugimoto, 1919): a — cephalic end; b—caudal end; с — cephalic end of female, lateral; d — same, apical; е — posterior end of female, lateral; f — first-stage larva; р — anterior end of larva; В — phasmids of first-stage larva, ventral (a, b — after Sugimoto, 1934b; c—h — after Olsen, 1952a). 267 Description (after Wehr and Chitwood,1934, as A. denticulo- phasma). The authors only had fragments of females (one cephalic and two caudal ends). Female. Width0.750 mm. Mouth rounded, enclosed in a delicate membrane with radial fibers. Esophagus consisting of a short, narrow muscular anterior part and a relatively large glandular part; boundary between the two parts indistinct. Intestine narrow, ending blind; anus rudimentary. Tail conical, curved dorsally. Vulva not found; two ovaries, one at each end of the body. Viviparous. Larvae from the uterus. Length 0.400—0.450 mm, width 0.143— 0.16 mm. Head small, conical, with dorsal processes. Tail pointed, curved ventrally. Phasmids large, postanal, with serrated margin. 257 Description (after Olsen, 1952a, as А. bifidus). The author only had fragments of two mature females 700 mm long, including one cephalic end. Female. Width 0.580 mm atthe anterior glandular widening of the esophagus and 0.835 mm near the posterior end of the second part of the esophagus. Cuticle with delicate transverse striation on the whole body, with 4 or 5 wide transverse stripes covering part of the body from both sides near the posterior end of the esophagus. Excretory pore not found. Width of head 0.313 mm, dorsoventral width 0.348 mm. Mouth forming a transverse slit 0.068 Х 0.023 mm large, situated on a square area 0.103 mm long, and with oval parts 0.070 X 0.117 mm contiguous dorsally and ventrally. Fourteen cephalic papillae. The interodorsal and interoventral papillae are projecting, with a double inner part with a striated stalk. Muscular anterior part of esophagus about 0.174 mm long and 0.150 mm wide; anterior glandular swelling 0.916 mm long and 0.406 mm wide; glandular posterior part of esophagus about 0.0075 mm long. Intestine not recognizaple. Tail sharply curved,with a distinct fold at the curvature. Anus not found and the direction of the curvature of the tail could therefore not be determined. End of tail oblique, with a narrowing of the parenchyma before the end. Tail markedly tapering from a point 0.116 mm from the end. Vagina and vulva not found. An ovary present in the posterior part of the body, its end 0.882 mm from end of tail. Uterus filled with larvae. Viviparous. First-stage larvae from the uterus 0.465—0.512 mm long and 0.013— 0.0143 mm wide; tail 0.132—0.156 mm long, pointed, straight. Phasmids large, postanal, with smooth margins. Cephalic end with a conical dorsal spine; 4 cephalic papillae. Esophagus with the anterior and posterior bulbs characteristic for the genus. Life cycle not established. Sugimoto (1934a) stated only that he found larvae of A. taiwana inthe body cavity of Cyclops sp., but did not describe them. 268 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.70; Chabaud and Campana, 1949, рр. 67—76; Chabaud, Campana and Truong-Tang-Ngok, 1950, pp. 335—339; Chitwood, 1935, рр. 51—54; 93—96; Olsen, 1952а, рр. 150—153; Sugimoto, 1919; 1934а, рр.261—266; Wehr, 1934, p.11; Wehr and Chitwood, 1934, рр. 10—11; Yamaguti, 1961b, p.328. Avioserpens galliardi Chabaud and Campana, 1949 (Figures 149—151) Synonyms: Avioserpens multipapillosa Singh, 1949; A. nana Mawson, 1957 Hests: Egretta garzietta, Ardeola greyil, А. ШЕЕ Localization: aponeurosis in upper part of esophagus, under the skin, in the neck. 258 Distribution: Europe, Canada, India, USSR. Historical review Avioserpens galliardi was described from a male and a female found in an aponeurosis in the upper part of the esophagus of Egretta garzetta inthe Eastern Pyrenees. A. galliardi is widely distributed in the USSR, and helminthologists confirmed the validity of the species and considerably increased the range of its definitive hosts. Shakhtakhtinskaya (1951) described Petroviprac- ta vigissi from the thoracic cavity of a mallard (Azerbaidzhan), for which she established a new genus. Chabaud and Campana-Rouget (1952) compared the posterior end of the male of P. vigissi with that of the specimen of A. galliardi described in 1949 and concluded that the two species are identical. Supryaga (1967b) agreed. Shakhtakhtinskaya thus first recorded A. galliardi inthe USSR. Ina study of parasites of fish-eating birds in the Astrakhan reserve, Kosupko (1963) found two males of A. galliardi, in Egretta garzetta onthe esophagus and in a mallard on the trachea. The wide distribution of A. galliardi in the Aral area is reported by Turemuratov (1965), who found this species in Ardea cinerea, A. purpurea, Egretta alba, в. garzetta, and Botaurus stellaris. Ina study of birds in the lower reaches of the Ob, Daiya (1967) found a male of A. galliardi ве 5 Mic mem can ет! and another in (Gay vats tel lata =the nematodes were localized in the subcutaneous tissue of the neck. \In the Azov estuaries in the Krasnodar Territory Supryaga (1967b) found this species in the subcutaneous tissue of the neck in Ardea pur- ВЕ а. ок ера‘ Warzetta, and) Botauris ys tellaris,/so that birds of three families are definitive hosts of A. galliardi: Ardeidae, Anatidae, and Gaviidae. Description (after Chabaud and Campana, 1949). Male. Length 8.7mm, width0.18 mm. Body forming two coils of а spiral. Transverse striation present on the whole body, very fine and dense. Cervical papillae very small, situated 0.48 mm from the cephalic end, 0.09 mm behind the nerve ring. Length of muscular part of esophagus 2690.125 mm, of glandular part 0.22 mm in the widened anterior part and 3.2 mm in the other part. Tail narrow, 0.3 mm long. Because of bad fixation, the phasmids and caudal papillae were not visible. Spicules 0.445 and 0.410 mm long. Gubernaculum strongly chitinized, smooth, 0.15 mm long. Female. Length 210 mm, width 0.68 mm. Cuticle with fine transverse striation which is almost absent in the middle of the body but distinct in the posterior part, where the intervals are 0.001—0.002 mm. Cephalic end separated from the body by a marked constriction situated 0.15 mm from the cephalic end. Mouth rounded, small, slightly protruding. Circumoral ring absent. Lumen of esophagus triangular, its apex ventral. Cephalic shield with 14 papillae in two concentric rings. Two amphids situated laterally between the two rings of papillae, forming two distinct 270 oval parts; the inner of these parts bearing an interolateral papilla. Median papillae large. Ventral and dorsal papillae double. They are situated on elevations which are clearly visible in lateral view. The lateral 259 papillae of the outer ring form two pairs on each side; the lateral papilla is slightly larger than the other papilla. Cervical papillae hemispherical and weakly protruding, situated 1.6mm from the cephalic end. Tail ending 271in a small, pointed appendage about 0.01 mm long. Length of tail 0.75 mm. (269) FIGURE 149. Avioserpens galliardi Chabaud and Campana, 1949: а — cephalic end, lateral; b — same, apical; с — anterior end; 4 — caudal end of female, lateral; e — region of vulva, lateral (after Chabaud and Campana, 1949). A simple papilla situated at the anus. Phasmids not found. Length of muscular part of esophagus 0.35 mm, Glandular part divided by the nerve ring into a swollen anterior part, which is oval and 0.85 mm long, and a posterior part 62 mm long. The intestine is recognizable only in the posterior region, where it narrows sharply before the anus. Position of vulva not determined. Uterus, filled with larvae, occupying almost the entire body cavity; it curves slightly above the anus in the caudal part and ends 3.8 mm before the cephalic end. Description (after Singh, 1949b, as A. multipapillosa). Female (immature). Length 105 mm, width0.12 mm. The widening of the glandular part of the esophagus distends the cephalic end in the form of a cupola. Cephalic end with 7 pairs of distinct papillae, two pairs 260 — „В submedian, one pair lateral, and two pairs оп each side between the lateral and submedian papillae. Esophagus divided into a short muscular anterior part 0.186 mm long and a glandular swelling 0.688 mm long and 0.658 mm wide. Intestine thin, rectum short. Tail 0.68 mm long, tapering, curved ventrally. Vulva indistinct, situated 12.25 mm from the cephalic end, dividing the body at the ratio of 1:7.5. (270) FIGURE 150. Avioserpens galliardi Chabaud and Campana, 1949. Caudal end of male (after Chabaud and Campana, 1949). Description (after Mawson, 1957a, as A. nana). Cuticle with fine annulation. Cephalic papillae more distinct in females, two situated dorsally and two ventrally near the mouth; outer ring consisting of a pair of lateral papillae and 4 pairs of submedian papillae. Amphids not found. Esophagus consisting of a short anterior and a longer, widened posterior part. Male. Length 7 mm. Length of anterior part of esophagus 0.140 mm, of posterior part 0.700 mm. Tail long, tapering sharply to a point. At its narrowing there are 5 ventral spines, and two accessory spines at the end. Tail 0.300 mm long. Analand caudal papillae absent. Bothspicules 0.240 mm long, their proximal third wider. Gubernaculum 0.150 mm long, with widened proximal part. Female. Length27.5 mm. Length of anterior part of esophagus 0.150 mm, of posterior part 0.500 mm. Vulva and larvae not found. Tail long, tapering, then narrowing abruptly, ending ш а point. Tail 0400 mm long. Life cycle (after Chabaud and Campana, 1949). Larvae from the uterus of A. galliardi placed in physiological saline and in fresh water were very active and remained alive for 2 days. First-stage larva 261 0.540 mm long, 0.018 mm wide. Length of esophagus 0.16 пил; ratio of length of esophagus to length of larva 3:4. Nerve ring situated 0.077 mm, anus 0.37 mm from the cephalic end. Twelve hours after the larvae had been taken from the uterus, they were placed together with Cyclops; 24 hours later, three of the Cyclops examined contained a larva with a pointed tail, which is characteristic for A. galliardi. Twelve hours after infection, one out of 6 Cyclops infected contained two larvae which had molted (the long thin tail was absent). (270) aes | 0020 mm Е Е | S м \ S Fil {4s | S a “aR сс 2 fd i Е } 5 | 8], d e FIGURE 151. Avioserpens galliardi Chabaud and Campana, 1949: a — anterior end of male; b — anterior end of female; с — posterior end of male, lateral; d — cephalic end of female; e — caudal end of male; f — caudal end of female, lateral; g — cephalic end; В — anterior end (a—e — after Mawson, 1957а; Е — after Singh, 1949b). ; 272 A larva taken from a Cyclops after 15 days was 0.480 mm long, and the tail showed the characteristic spines. The esophagus was 0.24mm long. References: Daiya, 1967, pp.142—145; Kosupko, 1963, рр. 158—159; Shakhtakhtinskaya, 1951, p.163; Skrjabin, Shikhobalova, Sobolev, Para- monov, and Sudarikov, 1954, p.70; Supryaga, 1967b, pp.116—165; Ture- muratov, 1965, p.102; Chabaud and Campana, 1949, pp.67—76; 1952, р. 482; Chabaud, Campana and Truong-Tang-Ngok, 1950, pp. 335—339; Mawson, 1957a, pp.213—219; Singh, 1949b, pp.54—55; Yamaguti, 1961b, р. 929. | Avioserpens mosgovoyi Suprjaga, 1965 (Figures 152—156) Ho'sits: | Fulicavatra; Podiceps erisitatus, “Ph Ик весе PY errs ета ету ах. Localization: subcutaneous tissue. Distr rb wiron: USSR. 262 Description (after Supryaga, 1965а). Large nematodes, slightly thinner toward the cephalic and caudal end. Body threadlike, white. Cuticle thin, transparent, with delicate transverse striation. Mouth simple, without a cuticular ring or other formations, rounded, or short-oval. (273) ПЕНЕЙ И он РОТ ИО г: FIGURE 152. Avioserpens mosgovoyi Suprjaga, 1965: a — anterior end of male, lateral; b — caudal end of male, lateral; c — spicules and gubernaculum (after Supryaga, 1967a). 263 (274) GAL хх oN FENN ~ ИАА ] ee Na С FIGURE 153. Avioserpens mosgovoyi Suprjaga, 1965: a — cephalic end, apical; b — anterior endof female, lateral; с — caudal end of fernale (after Supryaga, 1967a). Fourteen cephalic papillae in two concentric rings: inner ring consisting of two interolateral papillae, outer ring of four double dorsoventral papillae and eight paired ventral, lateroventral, laterodorsal, and dorsal papillae. Amphids situated between the two rings of papillae, behind the interolateral papillae. Esophagus consisting of a short muscular anterior part and a long glandular posterior part. Glandular part divided by the nerve ring into a swollen anterior and an elongate, narrower posterior part. Intestine tubular, long and narrow. Male. Length 6.8—14.0 mm, maximum width 0.135—0.209 mm. Mouth passing into the distinct buccal cavity, which is 0.010 mm long and 0.015 mm wide. Muscular part of esophagus 0.085—0.132 mm long, with thick walls and a distinct sinuous lumen. Width of muscular part of esophagus 0.040—0.052 mm. Glandular part 0.110—0.220 mm long and 0.070—0.145 mm wide. Posterior elongate part of glandular part of esophagus 1.40—2.31 mm long. 264 Nerve ring surrounding constricted part of glandular part of esophagus, situated obliquely, 0.250—0.350 mm from the cephalic end. Intestine 5.7 to 11.3 mm long, according to the length of the body; its width and pigmentation decrease with age, and it is difficult to see on the background of the genital duct, around which it is wound inthe form of a nar- row light brown ribbon. Genital duct single, 5.45—11.331mm long. Spicules reddish brown, strongly chitinized, curved ventrally. Proximal end of spicules widened. 273 From this end extend two dark brown stylets, which end in small processes at the distal end. Between the stylets extends a light brown, delicate chitinized membrane which forms a canal along the spicules, narrowing 274 distally. Spicules of almost the same length, right spicule projecting slightly anteriorly, 0.170—0.190 mm long, left spicule 0.165—0.185 mm long. Maximum width of spicules at the proximal end 0.024 mm. The Spicules narrow 0.04 mm from the proximal end, forming a constriction 0.016 mm wide, and then widen again to 0.020 mm. Width of spicules at the distal end 0.004 mm. Gubernaculum reddish brown, strongly chitinized,-0.080—0.108 mm long, its proximal end 0.020—0.030 mm wide; the gubernaculum narrows to 0.10—0.15 mm distally. The narrow part of the gubernaculum bears distinct spines on the lower surface. Tail conical, curved ventrally, ending in three spines; the two ventral spines 0.004 mm long, the dorsal spine 0.008 mm. Each spine ends in papillae which are supplied with nerve endings. Four pairs of postanal sessile papillae. Female. Length 880.0—1130.0mm, maximum width 0.8—1.0 mm. Muscular part of esophagus 0.198—0.286 mm long, 0.015—0.050 mm wide. The glandular part of the esophagus consists of a swollen anterior and an elongate, narrower posterior part separated by the nerve ring. Length of swollen glandular part 0.470—0.960 mm, width 0.430—0.600 mm. Nerve ring situated 0.743—1.950 mm, cervical papillae 1.14 mm from the cephalic end. The entire body cavity is occupied by the wide, saclike uterus filled with larvae, which are 0.450—0.500 mm long. Life cycle, after Supryaga (1965b). The intermediate hosts of А. mosgovoyi are species of Cyclops and Diaptomus. Larvae which have passed out of the uterus have a long awl-shaped process at the posterior end (0.048—0.064 mm) and a distinctly transversely 275 striated cuticle. Length 0.440—0.510 mm, maximum width 0.012—0.016 mm. Length of esophagus 0.120 mm, of intestine 0.140 mm. Cephalic end rounded, mouth terminal. Length of tail0.094 mm. The larvae are active in water, remaining mobile for 4—6 days at 20—25°. A few hours after entering the intestine of Cyclops, the larvae pass into the body cavity, where they develop further. At a water temperature of 22—29° the first molt takes place 6—8 days after infection of Cyclops; the larvae do not discard the sheath. Second- stage larvae are little active; cuticle delicate, smooth; the awl-shaped process at the tail is lost; their size decreases slightly: length 0.439— 0.499 mm, width 0.016—0.022 mm. Length of esophagus 0.135—0.220 mm, length of intestine 0.1800—0.2025 mm. Nerve ring and excretory pore distinct, situated 0.0728—0.0810 and 0.0810—0.0864 mm, respectively, from the cephalic end. 265 FIGURE 154. Avioserpens mosgovoyi Suprjaga, 1965: а — 1агуае in Cyclops; b-— first-stage larva; с — second-stage larva; d — third-stage larva from Cyclops; е — third-stage larva from the reservoir host 74 days after infection, (after Supryaga, 1967a). The second molt takes place after 2—3 days. The third-stage larvae are active and grow rapidly without any morphological changes; their 276 distinguishing character is the presence of three spines (two lateral and one dorsal) at the end of the tail. Third-stage larvae remain viable for 4 months in Cyclops (Supryaga,1967a). At this stage the larvae are 0.389— 0.478 mm long and 0.019—0.022 mm wide. Cuticle with delicate transverse striation. Cephalic end with two amphids and four double papillae. Nerve ring situated 0.078—0.081 mm from the cephalic end. Length of esophagus 0.162—0.262 mm. At the base of the esophagus are three long glands which open into the esophagus near the nerve ring. Tail 0.040—0.067 mm long. Genital primordium oval, 0.013—0.019 mm long, with a large nucleus, situated 0.04—0.054 mm behind the end of the esophagus. Supryaga (1967b) found that the life cycle of A. mosgovoyi includes reservoir hosts: fingerlings of roach, gobies, and sticklebacks and dragonfly larvae, in which the larvae do not change morphologically but remain viable for 2—2.5 months. After entering the stomach of the definitive host, the larvae of A. mosgovoyi migrate into the serosa and mesentery, where the third molt takes place 4—5 days after infection. The fourth-stage larvae differ 266 according to their sex. Length of male larva 0.970 mm, of female larva 1.710 mm. The esophagus consists of a muscular and a glandular part, and a glandular mass is formed above the nerve ring. The tail ends in five spines. The male larvae have a distinct preanal widening in which the spicules are situated, which are curved ventrally and 0.035—0.040 mm long. Gubernaculum 0.020—0.025 mm long. FIGURE 155. Avioserpens mosgovoyi Suprjaga, 1965: a — anterior and posterior end of fourth-stage larva (male) 7 days after infection of the definitive host; b — anterior and posterior end of fourth- stage larva (female) 9 days after infection of the definitive host (after Supryaga, 1967a). The larvae later migrate through the air sacs into the subcutaneous tissue, where the fourth molt and copulation take place after 12 days in males and after 13—14 days in females. After copulation, the vulva, vagina, and anus of females become obliterated; the intestine atrophies and is pressed against the body wall by the strongly developed uterus, which is filled at first with eggs and then with larvae; four weeks after infection of the definitive host, A. mosgovoyi begins to discharge larvae. Under optimal conditions the entire life cycle in the intermediate and definitive hosts lasts 36 — 53 days. Supryaga (1969b) established experimentally that the males of A. mosgovoyi remain viable for a long time and actively emerge from 277 the tissues of the bird 2—3 days after the death of the host; the life of males in the definitive host lasts markedly longer than of females, as long as 260 days. The author assumed that repeated fertilization of females during a second infection may occur, even if a second infection takes place after a long time. 267 Avioserpensiasis of aquatic birds (after Supryaga, 1967a, 1969а). Avioserpens mosgovoyi causes a disease which is widely distributed in the flooded Azov limans in the Krasnodar Territory, in- fecting to 35% of coots and to 20—25% of grebes. Mature females are localized mainly in the subcutaneous tissue of the submaxillary space, forming large tumors. The size and weight of the head of the bird are often doubled by the tumors. At a high intensity of infection (more than 15 specimens), mature females of Avioserpens also become localized in the subcutaneous tissue of the head, neck, trunk, legs and wings and in the serosa of the intestine and around the cloaca. In places where tumors are formed the skin becomes very thin and without feathers; the bulging parasite is distinctly visible through the skin. In any localization, the cephalic end of females reaches an area of skin without feathers and the most frequent contact with water, i.e., the submaxillary region, around the eyes, the articulation of the wings, the knees and calcanean joints, the cloaca. The surface of the skin where perforations are made develops ulcers, each caused by a female. The ulcers increase in size and the area becomes covered with sores, which then dry and form scabs. The tumors are hard to the touch. The skin around them and subcutaneous fat are inflamed; The inflammation sometimes spreads, causing death of the bird. The helminths are situated under the skin in a thin serous capsule in a dense coil. The nematodes constantly contract, and when the tumors are large and the skin is markedly stretched, the movements frequently become visible through the skin while the bird is still alive. If the females are localized in the serosa of the large intestine and in other places, smaller masses are formed and it is easier to remove them. When infected areas are opened, after a female has died and all larvae are discharged, a mass of necrotic tissue and nematodes are found. This entire mass is enclosed in the capsule; it is reddish brown and of loose consistency. The most favorable conditions for the spread of the infection are present in stagnant water bodies, which dry up and contain large concen- trations of Cyclops and Diaptomus (the intermediate hosts of А. mosgovoyi) and also fish, amphibians, and dragonfly larvae (reservoirs), which the birds eat. This is probably the reason that the highest rate of infection (to 35%) and intensity (more than 10 females) is found in such water bodies from July to September. The mortality of coots and grebes caused by Avioserpens is also highest in such water bodies. 279 Avioserpensiasis is a highly pathogenic disease. It is particularly severe in birds infected with 5 — 10 or more females of A. mosgovoyi; the birds lose weight (40—60%), become lethargic, and often die. Supryaga stated that control is possible only by protecting water bodies against the entry of first-stage larvae. He suggests that at the beginning of June, when tumors are formed but the larvae have not yet been dis- charged, the birds should be shot. References: Supryaga, 1965a, pp.272—275; 1965b, pp. 275—277; 1967a, pp.199—202; 1967b, pp. 243—244; 1969a, pp.65—67; 1969b, pp. 45—246. 268 FIGURE 156. Avioserpens mosgovoyi Suprjaga, 1965: a — tumor in the submaxillary region of a соот, caused by A. mosgovoyi; = A. mosgovoyi under the skin of the submaxillary region of Podiceps cristatus; ¢— A. mosgovoyi under the skin in the femoral region of Fulica atra (after Supryaga, 1967). 269 SUBFAMILY MICROPLEURINAE BAYLIS AND DAUBNEY, 1922 Diagnosis. Dracunculidae. Mouth rounded, with 4 (double?) papillae in the outer ring.. Esophagus divided into a muscular and a glandular part or not divided. Males with pointed tail; caudal wings absent, or only a right wing present. Caudal papillae present. Spicules developed. Vulva situated at the anterior end or in middle of body. Type genus: Micropleura Linstow, 1906. Genus Micropleura Linstow, 1906 Diagnosis. Micropleurinae. Cuticle with fine transverse striation and irregular rows of small tubercles. Esophagus divided into a short anterior and a long, wide posterior part. Male, Caudal wings present. End of tail conical, pointed. Three pairs of preanal and 4 pairs of postanal papillae. Spicules, short, very thin and pointed. Gubernaculum well developed. Female. Tail bluntly rounded. A pair of large, protruding subventral papillae. Vulva indistinct, situated slightly before middle of body. Vagina and vulva without musculature, Viviparous. Parasites of the body cavity and serosa of reptiles. Type species: Micropleura vivipara Linstow, 1906. Micropleura vivipara Linstow, 1906 (Figure 157) Host: Gavialis gangeticus:. Localization: body cavity, serosa. Distribution: Zoological Garden in Calcutta. Description (after Linstow, 1906c). Cephalic end without teeth or lips; anterior end rounded, with 6 papillae arranged in a ring and only 280 slightly protruding; mouth small and round; cuticle smooth; lateral lines low and narrow, without a canal; excretory pore absent; end of tail rounded. Male. Length 35 mm, width 0.72 mm. A thickening on each side which ends in a papilla. Musculature strongly developed; lateral lines 0.03 mm wide, wider laterally. Length of esophagus 4.009 mm. Nerve ring situated 1.002 mm from the cephalic end. Tail 0.062 mm long. On each side of the tail are three small papillae arranged in an arc on the ventral side; then follows a postanal papilla on a circular elevation on each side and behind them a small papilla on one side of the short tail. Both spicules 0.47 mm long. Female. Length37 mm, width0.79 mm. Length of esophagus 5mm, Nerve ring situated 1.25 mm from the cephalic end. Vulva situated slightly before middle of body, dividing the body at the ratio of 5:6. The branches of the uterus pass into the ovaries anteriorly and posteriorly. Ovaries 2.31 mm long. Тай 0.207 mm long. Viviparous. Larvae in the uterus 0.57 mm long, 0.017 mm wide. Cephalic end rounded. Tail long, pointed. Cuticle with distinct transverse striation. 270 FIGURE 157. Micropleura vivipara Linstow, 1906: а — anterior end; b — cephalic end; с — caudal end of male, lateral; d — caudal end of female, ventral (a, d — after Baylis and Daubney, 1922; b — after Baylis, 1939; c — after Baylis, 1924). References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p. 86; Linstow, 1906c, рр. 269—270; Yamaguti, 1961b, p.194; Yorke and Maplestone, 1926, р. 436. 281 Micropleura indica Khera, 1951 (Figures 158,159) Hosts: definitive — Prionyx ganpeticus, Lissemys pune - fata imtermediate — Cy clopis : Localization: body cavity. Distribution: india: Description (after Khera, 1951). Medium-sized nematodes. Cuticle thin, delicate. Mouth surrounded by 6 papillae, two lateral, two subdorsal, and two subventral. Male. Length 5—7 mm, width0.41 mm. Width at the cephalic papillae 0.09mm. Cuticular striation very fine. Length of esophagus 1.08—1.55 mm. Esophegus divided into a muscular anterior part 0.26— 0.4 mm long and a glandular posterior part. Tail pointed, 0.245 mm long. Five pairs of postanal and 3 pairs of preanal papillae; all papillae pedun- culate. Postanal papillae slightly smaller than the preanal papillae. Left spicule 0.172—0.2 mm long, right spicule 0.14—0.16 mm. Gubernaculum strongly chitinized, 0.052 mm long. Female. Body blunt at both ends. Length 16—23 mm, width 0.84 mm. Width of head at the cephalic papillae 0.19 mm. Cuticle smooth, but with 2—6 rows of small papillae which protrude markedly in the region of the 282 esophagus and project less in the posterior part of the body. Length of esophagus 1.9—2.02 mm, length of muscular anterior part 0.585 mm. 271 (281) FIGURE 158. Micropleura indica Khera, 1951: а — anterior end of female, lateral; b — cephalic end of female, apical; с — tail of female, lateral; 4 — female genitalia; е — caudal end of male, lateral; f — first-stage larva; g — anterior end of first-stage larva, lateral; h — anal region of first-stage larva, lateral; 1 — anterior end; j — caudal end of male, lateral (a—h — after Siddiqi and Jairajpuri, 1963; i, j — after Khera, 1951). Nerve ring situated 0.501 mm, excretory роге 0.9—1.02 mm from the cephalic end. Tail 0.28 mm long. Vulva situated before middle of body, 3.1mm from the cephalic end in a young female 7.2 mm long. Vulva atrophied in adult females. Vagina narrow, without musculature. Branches of uterus opposite. Eggs 0.05—0.080 X 0.040 mm large. Length of larvae 0.72—0.87 mm, width 0.020 mm. Length of esophagus 0.17 mm, Length of tail 0.39 mm. Description (after Siddiqi and Jairajpuri, 1963). Body compact, slightly narrowing toward the rounded cephalic end and the conical caudal end. Cuticle thin and transparent, without striation or small papillae. Mouth terminal, without cuticularization, surrounded by a double ring of papillae. Six large papillae in the outer ring and 6 papillae in the inner ring. Amphids large, protruding, opening in pores which are situated laterally. Esophagus divided into a muscular anterior part and a glandular posterior part. Nerve ring situated at the connection between the two parts. Protruding muscles absent at the level of the nerve ring. Longitudinal nerve fibers well developed. Male. Length 4.54—6.12 mm, width 0.27—0.36 mm. Length of esopha- gus 0.90—1.14 mm, length of the short muscular part 0.18—0.23 mm; ratio of length of muscular and glandular parts 1:4. Tail conical, tapering, 272 0.19—0.22 mm long; 3 pairs of preanal and 5 pairs of postanal papillae. Smaller spicule 0.11—0.14 mm long, larger spicule 0.12—0.17 mm. Gubernaculum strongly projecting. Female. Length 25.0 mm, width 0.87 mm. Length of muscular anterior part of esophagus 0.68 mm, of glandular posterior part 2.12 mm; this part slightly wider than the anterior part. Ratio of length of the two parts about 1:3. Uterus didelphic, amphidelphic, large, with twisted ovaries. Vagina small, narrow. Vulva not functional, atrophied, situated in about middle of body. In mature females the uterus contains numerous first-stage larvae and fills the entire body cavity. Life cycle (after 5199191 and Jairajpuri, 1963). The larvae of М. indica are of typical dracunculid structure. The larvae are discharged into the water. When larvae are kept together with Cyclops, these be- come infected in 2 hours. The larvae molt twice inthe Cyclops and develop into third-stage larvae after about 6 days. First-stage larvae thin, very active, free-swimming; when swallowed by Cyclops, they enter the body cavity through the intestine after 8—12 hours. The larvae remain active in the intestine and also in the body cavity, where they feed on the fluid. Infected Cyclops usually contain 4—6 larvae, sometimes to 14. Length of larvae 0.86— 0.89 mm, width 0.025—0.026 mm. Cuticle thin and delicately striated. Length of esophagus 0.16—0.17 mm. Nerve ring situated 0.068—0.089 mm, 283 excretory pore 0.075—0.090 mm from the cephalic end. Phasmids large, protruding, saclike, situated slightly behind the anus. Tail 0.35—0.38 mm long, threadlike, with pointed end. FIGURE 159. Micropleura indica Khera, 1951: a — anterior end of first-stage larva at the molt; b — posterior end of first-stage larva at the molt; с — second-stage larva; d — anterior end of second-stage larva; e — posterior end of second-stage larva; f — anterior end of second-stage larva at the molt; р — posterior end of second-stage larva at the molt; В — third-stage larva; 1 — anterior end of third-stage larva; j — posterior end of third-stage larva (after Siddiqi and Jairajpuri, 1963). 273 284 Second-stage larva. First-stage larvae molt 36—48 hours after they have been swallowedby Cyclops. Thesecond-stage larvae remain about 12—24 hours in the body cavity of the host. They are 0.52—0.6mm long and 0.027—0.032 mm wide. Intestinal tract divided into an esophagus 0.18—0.19mm long, an-intestine 0.25—0.26 mm long, and a rectum 0.025—0.032 mm long. Esophagus not yet divided into a muscular and a glandular part. Nerve ring situated 0.057—0.064 mm, excretory pore 0.079—0.081 mm from the cephalic end. Tail short, pointed, 0.074— 0.095 mm long. Third-stage larva. The second-stage larvae shrink and remain inside the sheath until the internal organs become differentiated; this takes place after 40—50 hours. The host is exhausted and sluggish at this time. Some Cyclops _ infected with numerous larvae die; they decompose, and the larvae pass into the water, remain active for some time and then die, Third-stage larvae resemble the second-stage larvae in size and general appearance, differing only in a few details. Length 0.53—0.71 mm, width 0.025—0.038 mm. Length of esophagus 0.18—0.32 mm; length of muscular anterior part 0.062—0.10 mm, of glandular posterior part 0.12—0.22 mm. Intestine 0.22—0.29 mm long; rectum 0.32—0.043 mm long. Nerve ring situated at the connection between muscular and glandular part of eso- phagus. Excretory pore situated 0.085—0.14mm from the cephalic end. Tail blunt, 0.043—0.072 mm long. Infection of Lissemys punctata was not performed. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.86; Khera, 1951, pp.55—58; Siddiqi and Jairajpuri, 1963, рр. 99—105; Yamaguti, 19615, р. 195. Micropleura vazi Travassos, 1933 505 6-1 С атлааю Sere ew oupis Localization: abdominal cavity. Distribution: South America. Description (after Travassos, 1933). Cuticle smooth, with longitudinal rows of cones and with transverse striation on the whole body with intervals of 0.040—0.064 mm. Musculature polymyarian, with narrow lateral fields. Sexual dimorphism marked. Mouth simple, apparently with two primitive lips and with three pairs of markedly reduced papillae. Excretory pore situated at the middle of the esophagus, consisting of a large cell with 4 processes, two anterior and two posterior, which are situated obliquely to the lateral fields. Esophagus divided into a muscular and a glandular part. Each consists of two almost equal parts. Nerve ring situated at the connection between anterior and posterior part of esophagus. Male. Length6 mm, width0.7 mm. Length of esophagus 1.2mm, length of thinner anterior part 0.33 mm, of posterior part 0.93 mm. Excretory pore situated 0.28 mm from the cephalic end. Tail coiled into a spiral, with lateral wings supported by 7 pairs of papillae; four more or less equally spaced postanal pairs and three preanal pairs, also equally spaced. 274 A single papilla before the cloaca. Two rows of cuticular cones оп the ventral side. Spicules thin, threadlike, both about 0.168 mm long. Guber- naculum wedge-shaped, about 0.056 mm long. Cloaca situated 0.29 mm from end of tail. Genitalia consisting of an ejaculatory duct about 0.73 mm long and an almost straight vas deferens which ends before the end of the intestine. Female. Length 25 mm, width1mm. Length of esophagus 2.2 mm, length of anterior part 0.60 mm, of posterior part 1.6mm. Excretory pore situated 0.9mm from the cephalic end. Tail conical, anus situated 0.5mm from end of tail; vulva point-shaped, situated in middle of body. Ovejector reduced, atrophied in gravid females. The uterus consists of a large funnel-shaped sac which extends to the middle of the oviduct without forming loops. Oviducts and ovaries short. Ovaries situated at the ends of the body (amphidelphic), ending in a large, light-refracting cell. The 285 uterus contains larvae at all stages of development, without eggs. Fully developed larvae tapering posteriorly; length about 0.37 mm, maximum width 0.013 mm. The esophagus is constricted and already divided into an anterior part about 0.11 mm long and a posterior part about 0.13 mm long. Anus situated0.10—0.14 mm from the end of the awl-shaped tail. In the middle of the esophagus, 0.067 mm from the cephalic end, there is a short transverse ridge which is of light color and does not stain, probably the primordium of the excretory cell. The larvae pass out through ruptures of the uterus and the body wall. References: Skrjabin, Shikhobalova, Sobolev, Paramonov,and Sudarikov, 1954, p.90; Travassos, 1933, pp.161—163; Yamaguti, 1961b, ва 95 Micropleura trionyxi Agrawal, 1966 (Figure 160) БЕ On yi xa pram pe ti cus). Localization: intestine. Description (after Agrawal,1966a). Medium-sized nematodes; body tapering at both ends. Cuticle thin, delicately striated. Cephalic end with 6 papillae, two lateral and 4 submedian. Esophagus divided into a short, narrow, muscular anterior part and a longer, wider, glandular posterior part. Male. Length 3.5—3.76 mm, width 0.18—0.26 mm; width of cephalic end 0.05—0.08mm. Buccal capsule very short. Length of muscular part of esophagus 0.15—0.25 mm, width 0.04—0.06 mm; length of glandular part 0.65—0.98 mm, width 0.06—0.08 mm; length of esophagus 0.80—1.13 mm. Nerve ring situated 0.16—0.23 mm, excretory pore 0.67—0.74 mm from the cephalic end. Eight pairs of caudal papillae: four pairs preanal, one pair adanal, three pairs postanal; a pair of phasmids at the end of the tail. Both spicules tubular, 0.14—0.18 mm long; gubernaculum long, wedge-shaped, 0.04 mm long. Female. Length 4.62—6.92 mm, width 0.53—0.71 mm; width of 286 cephalic end 0.05—0.07 mm. Length of muscular part of esophagus 0.21— 0.31 mm, width 0.06—0.08 mm; length of glandular part 0.95—0.25 mm, width 0.09—0.10 mm. Nerve ring situated 0.19—0.28 mm, excretory pore 275 0.53—0.56 mmfromthe cephalic end. Tail conical,0.10—0.15 mm long. Vulva situated nearly in middle of body, 2.72—3.66 mm from the cephalic end. Viviparous. (285) IM NNT И ААА АДАМОВА ll t a FIGURE 160. Micropleura trionyxi Agrawal, 1966: a — anterior end of male, lateral; b — caudal end of male, lateral; c — caudal end of female, ventral (after Agrawal, 1966a). Reference: Agrawal, 1966a, pp.107—114. 276 FAMILY PHILOMETRIDAE BAYLIS AND DAUBNEY, 1926 Diagnosis. Dracunculoidea. Length varying. Cephalic end rounded. Mouth simple, without lips, surrounded by 6 or 8 papillae. Anus sometimes absent in adults. Male much smaller than female; spicules of the same length, thin, pointed; gubernaculum present or absent. Female. Vulva indistinct or absent. Vagina rudimentary or absent in females filled with larvae. Branches of uterus opposite, tubular. Ovaries short, situated at ends of body. Viviparous. Parasites of the body cavity, serosa, and connective tissue of fish. Type subfamily: Philometrinae Yamaguti, 1935. Key to the subfamilies of the family Philometridae Esophagus divided into a muscular anterior part with two swellings and a short posterior part with a glandular appendage projecting into the еее MSoolmevereenul Тая sp raaloyavoyouekcleeet s Bait G о а Им Philometrinae Yamaguti, 1935, Esophagus divided into a narrow muscular anterior part and a widened glandular posterior part. Esophageal glands polynuclear ............. Philoneminae п. subfam . SUBFAMILY PHILOMETRINAE УАМАСОТТ, 1935 Historical review Yamaguti (1935) established the subfamily Philometrinae Yamaguti, 1935, with five genera: Philometra Costa,1845; Philometroides Yamaguti, 1935; Clavinema Yamaguti,1935; Ichthyofilaria Yama- guti, 1935; Sanguinofilaria Yamaguti, 1935. Skrjabin et al. (1954) added the genera Philonema Kuitunen-Ekbaum, 1933; Coregonema Bauer,1946; Phlyctainophora Steiner, 1921, but only provisionally, since the subfamily was in need of revision; the authors stated that the genera described by Yamaguti in 1935 (Philo- Metin Odes, “Clavimemaz.lchthyofiltaria! and: Sanguinofidla - ria) are known only from females. Rasheed (1963) studied the species of the family Philometridae from a large material and recognized eight genera and three subgenera in the 277 287 genus Philometra: Philometra, Alinema, and Ranjhinema.: Two of these genera have been established recently: Nilonema Khalil, 1960 and Rumai Travassos, 1960, and two were established by the author, Buckleyella Rasheed,1963 and Thwaitia Rasheed,1963, Rasheed retained Philometra Costa,1845, Ichthyofilaria Yamaguti, 1935, Philonema Kuitunen-Ekbaum, 1933, and Philometroides Yamaguti, 1935, but she considered Clavinema Yamaguti, 1935 as a synonym of Philometra and does not discuss the genus Coregonema, since she considered some of its characters as doubtful; she does not mention the genus Phlyctainophora. After a detailed study of the cephalic ends and the cuticle of the species of Philometridae and the discovery of cephalic papillae and cuticular processes of varying form and structure, Rasheed (1963) redistributed the species in the genera of the family; we have accepted her classification. Diagnosis. Philometridae. Esophagus short, entirely muscular or divided into a short muscular anterior part and a relatively long gland- ular posterior part. Anus and vulva atrophied. Body long and threadlike (the females are markedly longer than the males in this case) or relatively short and thick. Viviparous. Type север hrlomietra; (Costa, 0845. Key to the genera of the subfamily Philometrinae (after Rasheed, 1963) СОН СТе ООН 6. is cle боись его yar ape taller Ga el ae I. Cuticle with processes wi. чо Maelo dee. ole tebe, ЗОО a 1. Muscular anterior part of esophagus elongate, with two swellings se- parated by the nerve ring, with a free posterior glandular process at the connection of the two parts ....... Ichthyofilaria Yamaguti, 1935. Esophagus. without the.above: characters) <2. f.i1. .7 sli) 0. (eee ae 2.’ .Four large; lobe-=shaped, cephalic: papillae 5). 1: гие See 3. More than four cephalic papillae, papillae indistinct, small ог ENG. Pi а о оо RCS О eS 4. 3. Esophagus swollen or widened near the mouth, Tail of female blunt GE COMME. hn 5 arated custo So Se ое Thwaitia Rasheed, 1963 . Esophagus not swollen around the mouth, cylindrical. Tail of female А ease RRR aaNet ce Вита! Travassos, 1960. 4. Tail of female blunt, rounded, or slightly tapering. Tail of male blunt or rounded, with or without lobes; cloaca\terminal ...... )n)eleeenns Е sis as whe RSIS: BALSA Philometra Costa, 1845. 5. Cuticle with cone-shaped processes; esophagus cylindrical, not widened around the mouth. Tail of female pointed ... Nilonema Khalil, 1960. Cuticle without cones; esophagus bulb-shaped or slightly widened around the mouth. Tail of female rounded or blunt..........+..... 6. 6. Cuticle with finger-shaped cuticularized structures arranged in a cer- tain order; cephalic papillae forming large, flattened lobes. Cuticu- larized teeth present in the esophagus ... Buckleyella Rasheed, 1963 . Cuticle with small, irregularly arranged cones; cephalic papillae small. Teeth absent in esophagus „ни Philometroides Yamaguti, 1935. 278 Key to the subgenera of the genus Philometra 1. Cephalic papillae of outer ring large, lobe-shaped, arranged in double О OR Se Abd fal aa valine ners Seen и eS a аи, Ranjhinema Rasheed, 1963. Cephalic papillae of outer ring smaller, sometimes fused but not ar- варенье Pair Suns. die Ecur анте mse Tal aie, Е о л.н Dis 2. Slit of mouth covered with small structures appearing like beads; cephalic papillae small, difficult to see .... Alinema Rasheed, 1963. Slit of mouth simple, without ornamentation; cephalic papillae of both NRCS EMEC. о ме ры Philometra Rasheed, 1963. 288 Genus Philometra Costa, 1845 Synonyms: Filaria Mueller, 1787 (in part); Ichthyonema Diesing, 1861; Sanguinofilaria Yamaguti, 1941 Historical review Philometra reticulatum Costa, 1845 was described from Uranoscopus scaber. Railliet (1916) compared the species with Filaria globiceps Rudolphi, 1819, which had been transferred by Diesing (1861) to the genus Ichthyonema, and considered them identical and the names Filaria and Ichthyonema as synonyms of the genus Pimetonmletoayaviith the type species Ph. \ 5 lobieeps , making Ph. reticulatum asynonym. The number of species in this genus is rapidly increasing, but the validity of many species is doubtful. Skrjabin et al. (1954) list 37 species in the genus, Yamaguti (1961) 44 species, and Rasheed (1963), who revised the genus, about 50 species. Diagnosis. Philometrinae. Body cylindrical, with blunt, rounded, or slightly tapering ends. Cuticle smooth. Cephalic papillae present: outer papillae arranged in pairs, fused or double; inner circle, if present, consisting of 4 submedian papillae. Esophagus with an anterior widening, with an esophageal gland and aventricle or without them. Intestine ending blind. Anus and vulva absent. Ovaries opposite, uterus compact. Viviparous. Males with spicules of the same or of nearly the same length. Gubernaculum present or absent. Cloaca terminal. Parasites of marine and freshwater fishes. iywe spectre s: SP hwlo mein a. этот ер 5. (Вы, 1819) Ве тее 1916. Subgenus Philometra Rasheed, 1963 Philometra globiceps (Rudolphi, 1819) Railliet, 1916 (Figures 161—163) Synonyms: Filaria globiceps Rudolphi,1819; Philometra reticaudata Costa, 1845 279 Hosts: Blennius phycis, Uranoscopus scaber. Localization: body cavity, gonads, Distribution: Mediterranean, Bermuda, Black Sea. Description (after Willemoes-Suhm, 1871). Live nematodes blood-red, with dark intestine, which extends close to the body wall almost to the posterior end. Musculo-cutaneous sac 0.014mm thick. Anterior end of esophagus slightly widened, mouth funnel-shaped, with- out lips, with 4 papillae. Esophagus very short,consisting of radial muscle fibers with a structureless membrane outside and lined with the continuation of the cuticle. Esophagus rounded posteriorly, intestine much wider, with thin walls, wide lumen, ending blind before the posterior end, attached to the caudal muscles only by thin strands. Tail short, rounded. Anus not found. 289 fae eax > ТВ FIGURE 161. Philometra globiceps (Rudolphi, 1819): a — anterior end of male; b — posterior end of male; c,d — anterior end of female; e — posterior end of female; f — beginning of uterus; g — development of eggs and first-stage larva (after Willemoes-Suhm, 1871). Male. Length 6 mm, width 0.1—0.29 mm. Genitalia represented by testes and vas deferens. One spicule, with bifid end. 280 Female. Length 200 mm, width 1—2 шт. Two ovaries and a duct connecting them which functions as a uterus. One ovary situated at the cephalic and the other at the posterior end, the first forming loops around the esophagus and the second around the intestine. The uterus of mature females is filled with larvae with a long, thin tail which tapers to a point. Length of larvae 0.56 mm, width 0.02 mm. Mouth absent, but at the anterior end is a very narrow tube followed by a granulate primordium of the in- testine. In the uterus of a female more than 20mm long eggs at all stages of development may be present. Description (after Linton, 1901). Blood-red nematodes, with dark brown intestine. Uterus filled with active larvae with some granulated 290 speckles in the middle of the body. Female. Length 55mm, width uniform, 0.45—0.70mm. Width of cephalic end 0.19 mm. FIGURE 162. Philometra globiceps (Rudolphi, 1819): a — anterior end of specimen from Lobotes surinamensis; b-— larva from uterus; с — anterior end of specimen from Pomatomus saltatrix; а — beginning of in- testinal tract; е — connection of esophagus and intestine; f — posterior end; g — part of wall of intestine at the posterior end, characteristic reticulation visible; h — longi- tudinal section of part in middle of body of specimen from Tarfon atlanticus; i — eggs (after Linton, 1901). 281 Description (after Kovaleva and Khromova, 1967). Large nematodes; mature specimens reddish brown, juveniles light yellow, body long, thread- like, of uniform width, except at the rounded ends, which are slightly nar- rower. Cuticle thin and smooth. Mouth simple. Cephalic end with 8 papil- lae, four in the outer ring and four in the inner ring; papillae distinct in apical view. Only the four papillae of one ring are visible in dorsal view, so that Willemoes-Suhm mentioned only four papillae. One pair of 291 lateral amphids. Esophagus straight, muscular, widened anteriorly; it ends in a ventricle consisting of several large cells projecting into the intestine. Intestine wide, dark reddish brown. It becomes markedly narrower posteriorly and ends blind, adhering to the body wall near the posterior end. Nerve ring distinct. Uterus occupying nearly the whole body cavity, filled with larvae; it extends slightly anteriorly beyond the posterior end of the esophagus. Vulva and anus absent. SOY FIGURE 163. Philometra globiceps (Rudolphi, 1819): a — cephalic end, apical; b — general view; c — posterior end of male (b,c — after Yorke and Maplestone, 1926). Male. Length1.67—4.6mm, Measurements are of amale 4.6 mm long. Width 0.08 mm. Length of esophagus 0.32 mm, width 0.088 mm. Nerve ring situated 0.104 mm from the cephalic end. Spicules of the same length, needle-shaped, 0.137 mm long; gubernaculum 0.044 mm long. 282 292 Female. Length23.017to60 mm. Measurements of specimens 60 mm long. Width 0.60 mm. Length of esophagus 0.9mm, width 0.15 mm. Nerve ring surrounding esophagus 0.214 mm from the cephalic end. Life cycle not studied. The literature mentions only a few failures to infect Cyclops and Decapoda with larvae of Ph. globiceps. Dissection of Uranoscopus scaber from the Black Sea (Kovaleva and Khromova (1967)) showed a high infection rate with this parasite (35.1—86.5%); the intensity of infection varied from 2 to 9 specimens. The fish became infected after spawning in summer-autumn; later de- velopment of the nematodes and of the genital products apparently takes place at the same time. When the fish begin to spawn, the larvae pass out of the body through a rupture in the body wall and are discharged together with the genital products of the fish into the water, where they live free for 8—9 days at 22—24°. The experiments of Kovaleva and Khromova to infect crustaceans with larvae of Ph. globiceps were negative. The life cycle of the parasite is completed ina year. Adults do not leave their localization in the host, and the dark, dried cuticle of nematodes of earlier infections are therefore often found during dissection. References: Kovaleva and Khromova, 1967, pp.472—473; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р. 73; Linton, WIG pp. 141—481; 1967, рр. 985—126: Willemoes-Suhm, 1871, pp: 175 203: Ааа, 19616, р. 75. Philometra abramidis Osmanov, 1964 (Figure 164) EMO Sits Abia mus bie am, a. Localization: kidneys. Distribution: USSR (Syr-Darya River). Description (after Osmanov, 1964). Body threadlike. Mouth surrounded by papillae. FIGURE 164. Philometra abramidis Osmanov, 1964. Posterior end of male (after Osmanov, 1964). 283 Male. Length 2.00 mm, width 0.05 mm. Spicules of different length. Larger spicule 0.249mm long, smaller spicule 0.156 mm. Gubernaculum present, 0.066 mm long. Cloaca situated at the posterior end, with two liplike processes. Female unknown. | Reference: Osmanov, 1964, pp. 38—42. Philometra amazonica Travassos, 1960 (Figure 165) Host: Calophysus macropterus. Localization: body cavity. Distribution: Amazon River. Description (after Travassos, 1960). Male unknown, 293 Female. Coloration pink, body cylindrical. Length 250—950 mm, width 0.50—1.0 mm. Cephalic end blunt, without formations. Posterior end conical, blunt. Cuticle smooth. Mouth triangular, passing directly into the esophagus. Esophagus club-shaped, with widened anterior end, forming a pseudobulb, and a narrower part where the nerve ring is situated. Posterior part of esophagus gradually widening. Length of esophagus 1.20—1.40mm. Nerve ring situated in anterior quarter of esophagus. Intestine cylindrical, dark, rectum narrow, transparent. Anus subterminal. Genitalia amphidelphic. Anterior ovary situated near esophagus, posterior ovary at the posterior end. In mature females the uterus is filled with larvae and occupies the entire body cavity. Larvae 0.416—0.445 mm long, maximum width 0.014 mm. Vulva not recognizable. J NY FIGURE 165. Philometra amazonica Travassos, 1960: a — cephalic end, apical; b — same, lateral; c,d — anterior end, lateral; e, f — posterior end (after Travassos, 1960). Reference: Travassos, 1960, pp.15—20. 284 Philometra baylisi Vaz and Pereira, 1934 (Figure 166) оО Piamieliod us elmer ars) ; Localization: abdominal cavity. Distribution; Brazil: Description (after Vaz and Pereira, 1934). Cuticle transversely striated, without spines, papillae, or other formations. Mouth rounded, with 3 lips: two larger lateral lips and a smaller dorsal lip. The lips bear 3 small papillae, one of them median and two lateral. Esophagus beginning at the mouth; it is narrower at both ends and wider in the middle, 1.11 mm long. 294 FIGURE 166. Philometra baylisi Vaz and Pereira, 1934: a — anterior end, lateral; b — same, dorsolateral; с — same, showing esophagus and intestine; d — tail of female, lateral (after Vaz and Pereira, 1934). Nerve ring situated 0.24 mm, excretory pore 0.51 mm from the cephalic end, Intestine atrophied, pressed against a side of the body by the uterus, forming a loop behind the esophagus. Vulva not recognizable. Anus situated 0.16 mm from posterior end. Tail ending in a spine 0.012 mm long. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.73; Vaz and Pereira, 1934, pp.87—103; Yamaguti, 1961b, р. 76. Philometra biglobocerca Belouss, 1965 (Figure 167) Ros tin Мо №. (Ss. oy. Localization: body cavity. Distribution: USSR (Suifun River). Description (after Belous, 1965). Male unknown, 285 FIGURE 167. Philometra biglobocerca Belouss, 1965: a — anterior end, lateral; b — posterior end, lateral (after Belous, 1965). Female. Length 48mm, width 1.2mm, Cuticle with weak transverse striation. Posterior end slightly curved ventrally, with two spherical lateral processes 0.160 mm wide. Mouth surrounded by 6 papillae. Length of esophagus 1.24mm, width 0.070 mm. Anterior end of esophagus with a club-shaped widening 0.100 mm wide. Intestine widest near the esophagus, 295 then tapering posteriorly and ending blind. Ovaries threadlike, one ex- tending from the anterior and one from the posterior end of the uterus. Uterus filled with larvae 0.350 mm long and 0.016 mm wide. Reference: Belous, 1965, pp.57—58. Philometra fujimotoi Furuyama, 1932 (Figure 168) Host<:nOphiocephakhws angus Localization: fins, body cavity. Distribution: Japan. Description (after Furuyama, 1934). Males markedly smaller than females. Male colorless. Live mature females from the fins of the host are pinkish red. Cuticle smooth, body threadlike. Mouth without lips or papillae. Esophagus cylindrical, short, consisting of muscular and glandular formations; esophageal gland with one large nucleus; intestine about half as long as the body, ending blind, rectum atrophied. Male. Length 3.99 mm, width near posterior епа 0.964 пт. Cephalic end rounded, without lips or papillae. Posterior end truncate, witha membranous process. Two spicules of the same length, needle-shaped, about 0.09mm long, maximum width 0.007 mm. Female. Lengthabout 40 mm, maximum width1.5—2.0mm. Body tapering anteriorly and posteriorly, cephalic end rounded, posterior end pointed. Esophagus cylindrical, about 2.5 mm long. Intestine forming a yellowish brown tube, almost half as long as the body. 286 296 ve | | FIGURE 168. Philometra fujimotoi Furuyama, 1932: a — anterior end; b — posterior end; c — region of vulva (after Furuyama, 1934). In mature females the uterus contains only fully developed embryos, occupying a large part of the body, from one end to the other, ending at both ends in an ovary which is turned in the opposite direction. Vulva and vagina not found in mature females but visible in the middle and posterior third of the body in young specimens. Viviparous; uterus of mature females filled withlarvae, younger females filled with eggs at different stages of development. The eggs are at first spherical and later ovoid, with thin, hyaline shell. Larvae which have left the uterus are filarioid, 0.43 mm long and 0.015 mm wide, tapering posteriorly; tailthreadlike, а third as long as the body. Cuticle smooth, with fine transverse striation. Head rounded, mouth small, barely visible, without lips or papillae. The esophagus, in which a lumen is already visible, occupies the anterior third of the body. Intestine distinct because of the presence of granules which refract light strongly. Anus situated 0.09 mm from the posterior end. The life cycle was studied in 1934 by Furuyama. Mature females burst when placed in tapwater. Thousands of larvae are released into the water and remain active for about a week. Their activity later decreases, and they die after 2 weeks. Humuivama assumed that Cyclops*leuckarti, С. serrulatus, Cwesvematis, С. strenwus, “and С. эр. arejintermediate hosts) of Ph. fujimotoi. The larvae enter Cyclops and develop in the body eavity. Growth of the larvae apparently ceases after about a week, but the larvae remain alive for a long time in the host. Larvae 0.52—0.58 mm long, 0.013—0.015 mm wide. Furuyama fed fully developed larvae from Cyclops to Ophioce- phalus argus. Young adult females 11.5—13.5 mm long and 0.26— 0.48 mm wide were found in the fins after 4 months. The female later grows larger without morphological changes. The rate of growth of the nematodes in the definitive host depends on the water temperature. 287 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.74; Furuyama, 1932, p.17; 1934, pp.15—17; Yamaguti, 1961b, p. 76. Philometra hyderabadensis Rasheed, 1963 (Figure 169) Host: Wallageo attur Localization: mesentery. Distribution: india: Description (after Rasheed, 1963). Male unknown. Female. Length 23mm, width 0.03 mm. Body tapering slightly at both ends. Cuticle smooth, lateral fields distinct, with wide glandular areas. Mouth simple, cephalic papillae typical for Philometridae. Papillae arranged in two rings: an outer ring with 8 papillae in pairs and an inner ring of 4 small submedian papillae. A pair of amphids situated laterally. Esophagus beginning at the mouth, forming a bulb and with wide lumen anteriorly. It becomes narrower at the nerve ring, which is situated 0.19mm from the cephalic end. Esophagus 1/7 of the length of the body, 3.28 mm long and 0.85 mm wide at the anterior end. An esophageal gland with a large nucleus is situated in the middle, the excretory pore before it behind the nerve ring; the gland extends posteriorly to the end of the esophagus. A fleshy ventricle opens into the intestine, which ends blind before the end of the tail. 297 FIGURE 169. Philometra hyderabadensis Rasheed, 1963: a — anterior end, lateral; b — cephalic end; c — posterior end of esophagus; 4 — cephalic end, papillae shown; e — posterior end of female (after Rasheed, 1963). 288 The thin anterior ovary begins near the posterior end of the esophagus; the posterior ovary begins at the end of the intestine. Uterus filled with larvae 0.18—0.25 mm long and 0.015—0.017 mm wide. Cephalic end of larva rounded and apparently with a toothlike process near the mouth. Tail distinctly tapering. End of tail rounded, slightly tapering, with two small papillae at the end, each with double nerve endings. Reference: Rasheed,1963, pp.89—130. Philometra inimici Yamaguti, 1941 Synonym: Phe icryptocentri Yamaguti, 1961 РА = minnie as Таро тие ry proce mt Guise wil Iitaie te Localization: outer surface of stomach, body cavity. Wwe rb ution : Japan. Description (after Yamaguti, 1941). Male unknown. Female. Length 140mm, width1.3mm. Body pointed at the ends but of uniform width otherwise. Cephalic and caudal papillae absent. Cuticle thin, wrinkled. Nerve ring situated 0.24 mm from the cephalic end. Eso- phagus 1.06 mm long and 0.17 mm wide in the anterior part; an esophageal gland is present in about the middle. Ventricle indistinct, projecting into the intestine. Intestine dark, 0.5mm wide at the beginning and tapering 298 posteriorly, extending to the posterior end. Anterior ovary oval, 0.1mm wide. Uterus extending to about the middle of the esophagus. Posterior ovary elliptical, 0.31 X0.1 mm large, situated obliquely, behind the posterior end of the uterus, its distal end 0.35 mm before the posterior end; embryonal duct 0.15 mm long, widened at the connection with the uterus, which ends about 0.5mm before the posterior end. Uterus long, cylindrical, with rounded ends, filled with embryos 0.31—0.36 mm long and 0.012—0.015 mm wide. еее: югом (atter Vamaputi, 1961 а аз Ph. ciryptocentri)- Female. Length 93mm, width 0.8 пт. Body threadlike, tapering at both ends. Cuticle thin and smooth. Cephalic end not pointed, without papillae. Esophagus 0.7mm long, its anterior part widened, 0.078 mm wide, the other part cylindrical; ventricle about 0.950 Х0.080 mm, ending in valves. Intestine at first markedly widened, becoming thinner and obliterated pos- teriorly. The anterior ovary begins at the anterior end of the intestine, forms a double loop, and opens into the uterus 0.8 mm from the blunt pos- terior end, where it is curved before entering the uterus. Larvae 0.28— 0.35 mm long and 0.010 mm wide. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and pudamikov, 1954, p.77; Yamagcuti, 1941, p.393; 1961а, р. 218. Philometra lateobracis Yamaguti, 1935 (Figure 170) Synonyms: Sanguinofilaria lateolabracis Yamaguti, 1935; во те реа 5 Om beromor kh Уатта, 1995; Ph: sie laemae Yamaguti, 1935 289 299 Hosts: Lateolabrax japonicus, Parapristipoma trili- neatum, Epinephelus akaara, Sciaena schlegeli, Scom- beromorus chinensis, Otolithus ruber, Hemisphamus геот 511, Lates.calcariter, Johna@ms sina. Localization: ovary and oviduct. Distribution: Japan, Pakistan, India. Historical review The species was designated as the type species of the genus Sanguino- filaria,in which theauthoralsoplaced S. pinnicola Yamaguti, 1935 and S. scomberomori Yamaguti,1935. He did not describe the genus very convincingly. He stated that it differs from Philometra in the absence of cephalic and caudal papillae and of a differentiated stomach (Yamaguti, 1935). Yamaguti (1941) transferred it later to Philometra, but considered the absence of cephalic and caudal papillae as the main characters. Rasheed (1963) studied the specimens in the British Museum and specimens from the ovary of Otolithus ruber on the coast of West Pakistan and stated that cephalic papillae are present in all specimens. Caudal papillae are absent. The glands of the ventricle are distinct. She also considered Ph. scomberomori and Ph. sciaenae as synonyms of this species. Rasheed (1965) recorded two new hosts of the species in West Pakistan. Description (after Yamaguti, 1935). Воду of uniform width its entire length, with rounded ends. Male unknown, Female. Length 23 mm, width0.9mm. Cuticle thin and smooth, Cephalic and caudal papillae absent. Mouth narrow, leading directly into the esophagus. Length of esophagus 1mm, width 0.08—0.1mm. The eso- phagus is divided into a slightly swollen muscular anterior part and a cylindrical posterior part which is connected with the esophageal gland on the dorsal side. The posterior end of the esophagus projects into the intestine with a small glandular appendage. Between the intestine and the esophagus is a rudimentary stomach consisting of a few cells which are compressed anteroposteriorly. Intestine wide, narrowing posteriorly into a thin canal which adheres to the body wall near its end. Intestine with dark brown contents. Nerve ring situated in the narrowest part of the esophagus, about 0.21 mm from the cephalic end. One cylindrical ovary at each end of the body; it turns toward the middle of the body after the beginning of each end of the uterus. Length of ovary 2.7—2.8mm. Uterus filled with threadlike larvae which are 0.45mm long and 0.018 mm wide. Vulva and vagina absent. Description (after Yamaguti, 1941). Male unknown. Female. Length to 100 mm, width 0.55 mm. Body dark red, very wide, with bluntly pointed ends. Cephalic papillae absent or rudimentary. Caudal papillae absent. Nerve ring situated about 0.2mm from the cephalic end. Esophagus to1.06 mm long, 0.090 mm wide inthe widened anterior part. Intes- tine to 0.42 mm wideat first, narrowing posteriorly; anus absent. Anterior ovary to 2.1 mm long, including the embryonal duct, which begins 2.65 mm 300 from the cephalic end. Uterus extending almost to the cephalic end; 290 posterior ovary forming a loop at its posterior end. Larva with threadlike tail, length 0.47 mm, width 0.015 mm, genital opening situated at a point dividing the body at a ratio of 2.2—3.0:1. (2 9 9 ) ТИ 70 mm 70mm о o 9 Г.) Fy ° Pe oO Г. b | 7355 FIGURE 170. Philometra lateobracis Yamaguti, 1935: a — posterior end, lateral; b — anterior end, lateral; c — same, showing esophagus; 4 — caudal end of female, lateral; e — transition of esophagus into intestine; glandular stomach and process visible (a, b — after Yamaguti, 1935; c—e — after Rasheed, 1963). Description (after Rasheed, 1963). Female. Length 130—200 mm, width 0.8—0.9mm. Body of uniform width its entire length except at the bluntly pointed ends, which are slightly narrower. Cuticle thin and smooth. Cephalic papillae arranged in two rings: 4 papillae in the inner ring and 8 in the outer, the latter arranged in four pairs, the inner papilla of each pair situated submedially. There is also a pair of lateral amphids. All papillae are clearly visible in apical view. Esophagus widened at the cephalic end, entirely muscular, 0.9—1.4mm long and 0.09—0.1 mm wide in the widest part, around the mouth. The ventricle and process contain distinct gland cells; they open into the intes- tine, which is very wide and occupies almost the entire width of the body. It ends blind near the posterior end and adheres to the body wall. Nerve ring surrounding esophagus 0.2—0.25 mm from the cephalic end. Ovaries thin. Uterus of mature females filled with larvae; eggs at different stages of development present in young forms. Length of larvae 0.47 mm, width 0.019 mm. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, рр. 78,86; Rasheed, 1963, рр. 124—126; 1965, рр. 349—350; хамасие, 1935, рр. 958—360; 1941, р.392; 19616, р. 76. Philometra managatuwo Yamaguti, 1941 Host: Stromateoides argenteus. Localization: ovaries. 291 Distribution: Japan. Description (after Yamaguti, 1941). Male unknown, Female. Length 460 mm, width 1.30 mm. Body dark red, with conical ends. Cephalic papillae absent or rudimentary. Caudal papillae absent. Nerve ring situated 0.2—0.25 mm from the cephalic end. Esophagus 0.85— 1.075 mm long, 0.12—0.132 mm wide at the anterior swelling, with a gland- ular formation dorsally. Ventricle small, 0.13—0.35 mm. Intestine 0.5mm wide at the beginning, reaching to the posterior end. Anterior ovary about 3.0 mm wide, beginning about 3.0 mm from the cephalic end; it crosses the esophagus where it turns posteriorly and passes into an embryonal duct 0.1mm wide. Posterior ovary curved at the posterior end for about 1.5mm. Uterus extending to the posterior end of the esophagus or slightly beyond it. Larvae 0.45—0.6 mm long and 0.015 mm wide. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov; 1954, p..7;. Yamaguti, 1941, р. 393; 19615, p76. 301 Philometra mariae Layman, 1930 (Figure 171) Hosts: Protopsetta_ herzensteini,, Limanda, senha eee Sebastodes schlegelii-. Localization: subcutaneous tissue. Distribwtions, USSR(-Far. Kast). Description (after Lyaiman, 1930). Male unknown, FIGURE 171. Philometra mariae Layman, 1930: а — anterior end; b — caudal end (after Lyaiman, 1930). Female. Length 1.6—3.2 mm, width 0.73—0.81 mm. Mouth surrounded by a characteristic buccal capsule 0.22 mm long and 0.24mm wide. Eso- phagus 0.73 mm long. Intestine ending in a rudimentary anus. Uterus 292 a ПАРЕ ee ЗИРИИИИРОЗИИИ 302 with an anterior and a posterior branch. Nerve ring distinct, forming processes to the sides of the body. Genital opening absent. Viviparous. References: Lyaiman, 1930, p.120; Yamaguti, 1961b, p. 76. Philometra ovata (Zeder, 1803) (Figure 172) Synonym: Filaria: ovata Zeder, 1803 HOGESS ANP EMS lj Ame, vrs мп. Ij Sblens eus верная, Goebiotluviratilis, »Rutwlus* caspicus) В. frisai сити, Abramis ballenrusy, beuciscus) Neweiscus,) Ly dus; Vem virmba. “Asipius asipius, bilieca) Б]огЕпа, Phoxinus laevis. Localization: body cavity, occasionally intestine (females); males found on the serosa of the posterior sac of the swim bladder, rarely in the body cavity. Distribution: Western Europe, USSR. Description (after Skrjabin, 1923а). Male. Length 6 mm, width 0.1 mm. Female. Yellowish white, length 125 mm, width1.5mm. Larvae in uterus to 0.47 mm long and with a long tail (1/4 of length of body). Description (after Kosareva, published for the first time). Male not described. Female. Length 9—9.3 mm, width 0.6 —1.5 mm. Coloration yellowish white. Cephalic end rounded, with 4 weakly developed tubercles. Mouth triangular, bordered by three lobes in dorsal view. Buccal capsule absent. Esophagus slightly widened anteriorly,1.5 to2 mm long. Intestine ending blind. Uterus large, occupying entire body cavity, one ovary at each end. The size ofthe uterus changes with age. Vulva and vagina absent. Caudal end blunt, with distinct lateral tubercles which vary in size in different specimens. Viviparous. Description (after Molnar, 1966a). Male. Length 1.7—2.3 mm, width 0.045—0.053 mm. Transparent forms; cuticle smooth, witn fine transverse striation. Cephalic end rounded, with 4 elongate papillae arranged around the periphery. Body almost uniformly wide its entire length, slightly narrowing at the cephalic end. Caudal end truncate, with two slightly projecting lateral teatlike processes which are connected by distinct narrow lobes. Two spicules of different length, with a short, thick stem and long, thin, tapering apical part. Larger spicule 0.246—0.325 mm long, smaller spicule 0.136—0.204 mm. Stem 0.007—0.009 mm wide, apical part at first 0.005 mm, average width 0.0025 mm. Gubernaculum 0.053—0.074 mm long, forming a curved chitinized plate with lanceolate apex. There is a hornlike process in its distal part, above which the gubernaculum is curved dorsally and anteriorly. Males are localized in the peritoneum and on the serosa of the posterior sac of the swim bladder, rarely in the body cavity. Female. Length 100—120 mm, width 0.8—1.2mm. Mature females red, rarely yellowish red. Cuticle smooth. Cephalic end rounded, with 4 distinctly protruding papillae with wide base and with three lips. Eso- phagus and intestine pressed against the body wall. Posterior end with 303 two distinct, flattened teatlike processes. Anus subterminal, Body cavity 293 303 occupied entirely by the uterus, which is filled with larvae. Length of larvae 0.4—0.5mm. Young females yellowish red or red. Length 3.0— 20.0 mm. Cuticle smooth, with fine vertical striation. Body cylindrical. (302) FIGURE 172. Philometra ovata (Zeder, 1803): а — anterior end, lateral; b — posterior end; с — part of female; d — cephalic end of male; e, f — caudal end of male, different as- pects (after Molnar, 1966a). Esophagus not divided and, except for the bulblike widening at the mouth, of uniform width its entire length. One side of the esophagus is serrated and bears an esophageal gland which opens into the esophagus below the anterior widening and continues until it is twice as wide as the esophagus in its middle. The esophagus is separated from the wider intestine by 3 valves. Intestine containing reddish brown pigment. Its width is uniform but it narrows markedly before the tail and continues to the subterminal anus. Nerve ring situated below the esophageal bulb; distinct ganglia extend from it to the cuticle. The tail ends in two large teatlike processes which project in small specimens. Vulva rounded, situated in the posterior third of the body; vagina with chitinized walls, extending anteriorly for 0.180—0.220 mm, Uterus filled with eggs, occupying the greater part of the body cavity. In young specimens the anterior end of the uterus gradually narrows and ends at the posterior end of the esophagus; in more mature females it is markedly curved and ends near the esophagus. From the anterior end of the uterus begins the long, cylindrical anterior ovary which forms one or two loops near the esophagus, then turns and ends behind the anterior end of the uterus. The posterior ovary extends to the posterior end of the uterus and forms several loops and ends abruptly. The two ovaries are connected with the uterus in the same manner, Females move freely in the body cavity of bream and roach, 294 The life cycle of the species was studied by Molnar (1966a), who stated that in Lake Balaton mainly bream and roach older than one year infected with Ligula are severely infected with Ph. ovata. The rate of infection in both hosts is 100%. Infection with Philometra was not observed in fish less than a year old. Molnar noted that females of Ph. ovata complete their life cycle only when the fish are also infected with Ligula intestinalis or Dia- gramma interrupta. This agrees with the data of Kreplin (1825), who found Filaria cyprini rutilii (probably identical with Ph. ovata) ina roach infected with Ligula intestinalis. In breams not infected with Ligula and more than a year old the rate of infection is 100%; males and females of Ph. ovata are recorded throughout the year, but the intensity of infection is lower and only males become mature. The males are also localized in the wall of the swim bladder; there is thus no migration of females into the abdominal cavity. Such females never grow longer than 3 mm. Infection of bream and roach in Lake Balaton takes place in the second half of June, and young males 0.96—1.01 mm long and 0.027—0.047 mm wide and young females 0.54—0.87 mm long and 0.016—0.045 mm wide were found at this time. There were also specimens from the abdominal cavity which were larvae with narrow tails. Specimens longer than 1mm had already lost the long tail characteristic for the larvae and had rudi- 304 ments of spicules or of a chitinized vaginal opening. The chitinized parts of the spicules of these specimens were never longer than 0.040 mm, and in males less than 1.4mm long spicules were not found. In July, growing males and females were found only in the swim bladder. Length of males 1.02—2.10 mm, width 0.032—0.061 mm, length of females 0.95—2.10 mm, width 0.032—0.059 mm. In August, the males had grown to their full size. Their chitinized organs did not change further. Length of large spicule 0.246—0.325 mm, of small spicule 0.136—0.204 mm; length of gubernaculum 0.053—0.078 mm, After August females in fish containing plerocercoids and in fish without them developed differently. In fish not infected with Ligula males are 1.70—2.20 mm long and 0.041—0.053 mm wide and females 1.66—2.36 mm long and 0.039—0.053 mm wide. They do not grow further and are localized in July under the serosa of the posterior part of the swim bladder. The chitinized part of the vagina of these females is 0.080—0.100 mm long. In fish infected with plerocercoids the males also do not grow further but the females continue to grow and migrate from the air sac into the abdominal cavity. They are now 1.50—11.50 mm long and 0.029—0.034mm wide. Developing eggs 0.004—0.006 mm wide are present in the uterus. Some females develop slowly and may continue to develop to the beginning of the next cycle under favorable conditions. When the adult worms migrate, males and females may be found in the swim bladder, less frequently in the body cavity. After the females have left the host, the males disappear; at the end of July males and females of the same size which have not completed their growth may be found in the swim bladder. In September, females which have migrated into the body cavity are 2.90—25.00 mm long; males which have remained in the swim bladder are 295 305 1.8—2.0 шт 1005. Together with the males there are often smaller fe- males 1.4—5.6mm long. Some of these probably migrate later. According to the above data, copulation takes place in July or in early August. In March, the yellowishred or red females are actively moving in the body cavity of bream; they are 13.7—51.2 mm long and 0.36—0.68 mm wide. The uterus is filled with rounded, nonsegmented, thin-walled eggs 0.030—0.035 mm wide. In April the color of females varies from bright red to dark red. They are 45—70 mm long, and the uterus contains eggs 0.035—0.040 mm wide with 8—32 blastomeres. At the beginning of May the eggs become oblong, 0.040—0.052 mm large, with sickle-shaped embryos. The females are 70—90 mm long. In the second half of May all females are 9—12 cm long. They are bright red. The uterus is filled with numerous free first-stage larvae. At the end of May — early June fully grown females leave the host, enter the water, and burst; the larvae which are discharged settle on the bottom. The free first-stage larva is 0.350—0. 510 mm long and 0.012—0.022 mm wide inthe middle. The cephalic end is rounded, the posterior end tapering. Molnar found that some species of Cyclops common in Lake Balaton (Cyclops strenuus, Macrocyclops albidus, and Acantho- cyclops viridis) swallow larvae inthe water. The larvae of Ph. ovata pass through the wall of the intestine of the intermediate host and were observed mobile in the body cavity 15 days after infection. Death of the host stopped further development. The life cycle of Ph. ovata is almost identical in roach. References: Skrjabin, 1923a, pp.1—98; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p. 76; Kreplin, 1825, p. 86; Molnar, 1966a, pp.227—242; Nybelin, 1928, pp.1—4; 1931, рр. 58—64; Yamaguti, 19615, p.77. Philometra parasiluri Yamaguti, 1935 (Figure 173) Synonyms: Ph. opsalichthydis Yamaguti, 1935; Clavinema parasiluri Yamaguti, 1935 Hosts: Opsalichthys uncirostris, Parasilurus азовев Mogurnda obscura, Zacco platypus. Localization: eye orbit, body cavity. DistriputLon: Uso (Amur Basin), Japan, Historical review Yamaguti (1935) published a study of the helminth fauna of fish in which he described new species of Philometra, Ph. parasiluri and Pay opsaliteht hyd sy He also established the new genus Clavinema with the single species C. parasiluri, which is found in the same host as Ph. parasiluri. Rasheed (1963) stated that the only character distinguishing Ph. parasiluri and Ph. opsalichthydis is the 296 306 position of the esophageal gland and the granulate appearance of the esophagus in Ph. opsalichthydis; a granulate esophagus has not been found in other species of Philometra, possibly because the large granulate esophageal gland covers the entire esophagus. There is no great difference in the position of the opening of the esophageal gland: in one species it opens before the nerve ring, in the other below the nerve ring, which is probably a variation and not a specific character, and Rasheed therefore considered Ph. opsalichthydis as a synonym of Phe wparas ilar i. Rasheed also stated that the genus Clavinema Yamaguti, 1935 was described from unfertilized females. The generic diagnosis is based on their small size, the absence of cephalic and caudal papillae, and the presence of a bulb-shaped anterior part of the esophagus which continues anteriorly in three flattened lobes projecting slightly from the mouth, and with a small esophageal gland situated in about the middle of the esophagus. These characters are not very important if the parasite is not mature, and they are therefore not very useful as diagnostic criteria. Philometra parasiluri Yamaguti, 1935 was described from the orbit of Parasilurus asotus, the typical host of Clavinema, which was found on the inside of the operculum and in the submaxillary connective tissue. The difference between the two genera is the absence of cephalic and caudal papillae and the presence of a small gland in the middle of the esophagus in Clavinema. Cephalic and caudal papillae are present in Ph. parasiluri and the esophageal gland is very large. Cephalic papillae may be absent, asinother speciesof Philometra, but the other characters depend on the length of the body, which depends on the develop- ment of the eggs and larvae inthe uterus. The genus Clavinema cannot therefore be considered valid until mature specimens are found, and Rasheed suggests that itis a synonym of Philometra and Cla- VMitcomida parasiluri asynonym ol Ph> jparasiluri:) (We apree with her. © Description (after Yamaguti, 1935). Male unknown. Female. Lengthto 34mm, width0.5mm. Body threadlike, with almost truncate cephalic end. The head bears 8 large papillae with wide base, equally spaced at the anterior margin. Several indistinct accessory papillae are present on each side of the whole body. The conical muscular anterior part of the esophagus is 0.075—0.096 mm wide; its three anterior lobes project slightly from the mouth, which is 0.024—0.027 mm wide; the thin, cylindrical posterior part of the esophagus is 1.0—1.4 mm long, and its dorsal gland is well developed. Between the esophagus and the intestine is a small muscular ventricle, at the posterior end of which a lobed gran- ular process projects into the intestine which ends posteriorly in a compact strand which is attached to the wall of the posterior end. Anus absent. Two distinct papillae 0.027—0.031 mm wide are present on the blunt tail. The cylindrical, turned anterior and posterior ovaries are sometimes curved. The anterior ovary extends anteriorly to the nerve ring, which is situated about 0.12—0.2 mmfrom the cephalic end. The anterior end of the uterus is situated 0.27—0.31 mm from the cephalic end and its posterior end 0.34—0.8 mm from the end of the tail. Larvae in the uterus are 0.36—038 mm long and 0.09—0.012 mm wide. Vulva absent. 297 FIGURE 173. Philometra parasiluri Yamaguti, 1935: a — anterior end of female, lateral; b — posterior end of female, lateral (after Yamaguti, 1935). Yamaguti (1941) found in the orbital cavity of Parasilurus asotus a single mature female of Philometra 53 mm long and 0.75 mm wide in which he found, in addition to the two symmetrical terminal papillae, another 7 papillae at the posterior end (one dorsal, two subventral, and four lateral) and longitudinal rows of 2—5 smaller papillae; however, these are probably not always present. 307 Description (after Yamaguti,1935,as Ph. opsalichthydis). Female. Length 35—43 mm, width 0.5—0.53 mm. Eight cephalic papillae which are slightly flattened and 0.018—0.024 mm wide at the base. Two protruding caudal papillae 0.024—0.036 mm wide. Mouth 0.030—0.033 mm wide. The thickened muscular anterior part of the eso- phagus is 0.090—0.111 mm wide; its three convex papillae project slightly from the mouth; the cylindrical glandular posterior part of the esophagus is 0.83—1.32 mm long. Dorsal esophageal gland beginning behind the nerve ring. A muscular ventricle and its process present. Nerve ring situated 0.22—0.25 mm from the cephalic end. The intestine ends ina firm strand attached to the ventral body wall at the posterior end. The cylindrical, sinuous ovaries, one at each end of the body, differ markedly in position: the anterior ovary usually begins before the uterus (in not fully mature females), the anterior end of which is situated 0.37— 0.7 mm from the cephalic end; the posterior ovary sometimes extends al- most to the end of the tail or begins before the posterior end of the uterus, which is situated 0.43—0.66 mm from the posterior end. The uterus contains numerous fully developed embryos with a long, thin tail which are 0.39—0.42 mm long and 0.009—0.012 mm wide. References: Belous,1965, pp.48—65, Roitman, 1963b, pp. 263—312; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p. 78; Yamaguti, 1935, рр. 352—356; 1941, p.391; 1961а, pp. 217—228. 298 Philometra pellucida (Jagerskidld, 1893) (Figure 174) Synonym: Ph. sebastisci Yamaguti, 1941 Hosts: marine fishes Tetradon stellatus, T. hispidum, PVC Oars mMacropterus, | Belone,liwra, “sebastisecus marmoratus. Localization: testes. Distribution: Australia, Japan. Description (after Rasheed, 1965). Male unknown. Female. Length 45—85 mm, maximum width 0.6—1.0mm. Body tapering at both ends, which are not pointed. Weakly developed papillae visible in apical view. Esophagus with wide lumen, its walls slightly wrinkled. Lumen of the esophagus widening, forming a ''cavity'' in its anterior, bulb-shaped part. Esophagus with strong muscles anteriorly which pass through the bulb. Length of esophagus 1.4—1.56 mm. Tail rounded, with two very small papillae. The ovaries begin one at each end of the body. Uterus filled with developing eggs in young specimens, with larvae in mature forms. Larvae 033—0.4 mm long,0.007 mm wide. The larvae are coiled into a ring. (308) 02.7 тт FIGURE 174. Philometra pellucida (Jagerskiold, 1893): а, b, с, d, g, j — cephalic end of female, different aspects; i — anterior end of female, lateral; e, Е, h, К — posterior end of female, different aspects (after Rasheed, 1963, 1965). 299 Rasheed's description is the same as that of Jagerskiold, except for 308 the smaller size, which Rasheed attributes to age differences and the im- maturity of the specimens she examined. Description (after Yamaguti,1941,as Ph. sebatisci). Male unknown. Female. Length to 135 mm, width 2.0mm. Coloration dark red, ends of body blunt. Cephalic and caudal papillae absent. Cuticle thin and wrinkled. Nerve ring situated 0.25—0.3 mm from the cephalic end. Esophagus 1.25—1.45 mm long and 0.12—0.17 mm wide in the widened anterior part. Esophageal gland and ventricle weakly developed. Anus absent. Ovaries 0.12—0.32 mm wide, extending anteriorly to the anterior bulb of the esophagus, beginning near the posterior end. Uterus filled with larvae insheaths. Length of larvae 0.32—0.35 mm, width 0.015— 0.020 mm, with a caudal spine 0.006—0.008 mm long. Nerve ring situated 0.048—0.057 mm from the cephalic end, esophagus and intestine differen- tiating, anus dividing the body at a ratio of 6—6.1:1. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.79; Jagerskidld, 1893, p.86; Rasheed, 1963, pp.-89—130; .1965,..pp, 352—356; . Yamaguti,, 1941, р.393; 19615, в 309 Philometra percalates Johnston and Mawson, 1940 (Figure 175) host: Per cava tesmc 0] олова т, Localization: body cavity. Distribution: Australia. Description (after Johnston and Mawson, 1940). Male. Length 2.6 см. Cephalic end rounded, with 8 small papillae. Esophagus 0.25 mm long, with widened anterior end, narrowed near the nerve ring, widened posteriorly. Nerve ring situated 0.15 mm from the cephalic end. Spicules 0.105 mm long, with narrow wings; gubernaculum 0.04 mm long, with wide apex. Tail truncate, with 4 lobes, the ventral pair of lobes longer and the dorsal pair more distinct. FIGURE 175. Philometra percalates Johnston and Mawson, 1940: a — cephalic end; b — posterior end of male (after Johnston and Mawson, 1940). 300 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.78; Johnston and Mawson, 1940, p.349; Yamaguti, 961, ptt. Philometra pinnicola (Yamaguti, 1935) Yamaguti, 1941 (Figure 176) Symenyms: Sanguinofilaria pinnieola, Yamaguti, 1935; Philo- metra serranelli-cabrillae Janiszewska, 1949 Hoste: £Mpinephelus akkara, Serranelius cabrilla. Localization: fins, ovaries. Distribution: Japan, Hurope. Historical review Ph. pinnicola was described inthe genus Sanguinofilaria Yamaguti, 1935, which the author later (1941) considered as a synonym one halonveLra. Rasheed (1963) compared the description of Ph. pinnicola and Ph. serranelli-cabrillae Janiszewska, 1949, and stated that these species are very similar and that their hosts all belong to the family Serranidae and their distribution is also similar. She therefore made Pheeserranelli-=cabrillae а synonym of Ph. pinnicola. Description (after Yamaguti, 1935, 1941). Body cylindrical, dark red in life. Intestine narrow, without dark pigment granules in the epi- thelium. Male unknown, Female. Length 20—40mm, width 0.65—1.25 mm. Nerve ring situated 0.15 mm from the cephalic end. Length of esophagus 1.0—1.25 mm, width at the anterior swelling 0.080—0.090 mm. Ovaries 0.055—0.11 mm wide, twisted around the ends of the uterus. Embryos with a sheath, with a brown 310 tooth at the cephalic end; length of embryos 0.32—0.37 mm, width 0.02— 0.025 mm. Description (after Janiszewska, 1949, as Ph. serranelli- cabrillae). Male unknown. me my le’. Body threadlike, reddish to reddish brown, tapering at both ends. Length 32—60mm. Cephalic and posterior end blunt; sensory papillae absent. Cuticle smooth. Width of body 1 mm ina large female. Esophagus cylindrical, 1.30 mm long and 0.15 mm wide at the widest point in females 60 mm long. ''Pharynx'' with a gland on the dorsal side which extends to half its length. Esophagus slightly widened anteriorly, passing through a small ventricle, which consists of glandular cells, and then with a small glandular process into the intestine. The intestine ends blunt in the posterior part, where it narrows and adheres to the body wall. It consists of large oblong cells and is filled with small reddish brown pigment granules. The intestine is markedly darker in adult than in young females. Nerve ring situated 0.37 mm from the cephalic end. The uterus forms a broad duct which occupies the entire width of the body cavity; it extends almost to the nerve ring and ends at the end of the esophagus. 301 а FIGURE 176. Philometra pinnicola (Yamaguti, 1935): а — anterior end, lateral; b — posterior end, lateral; с — region of connection between esophagus and intestine; d — anterior end (a—c — after Janiszewska, 1949; d — after Yamaguti, 1935). It is entirely filled with embryos. In mature females the uterus contains filarioid embryos 0.35 mm long and 0.07 mm wide. On each side of the uterus is an ovary which forms a more or less winding tube. Females taken from the definitive host and placed in fresh water or in physiological saline break up rapidly. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.78; Janiszewska, 1949, pp.7—20; Yamaguti, 1935, 2.360 1941, 0.393; 19615, pi 77. 311 Philometra scomberesoxis Nikolaeva and Naidenova, 1964 (Figure 177) Hosts Scomberesox saurus’. Localization: body cavity. Distribution: Ющап and Tyrrhenian seas. Description (after Nikolaeva and Naidenova, 1964). Male unknown, Female. Body thick, slightly tapering at the ends. Both ends of body blunt, without processes or sensory papillae. Cuticle smooth, with numerous small oval cells with large rounded nuclei below it. Such cells 302 — Ss are present throughout the body but they are particularly numerous at the ends. The reddish brown color of the body is due to a dark brown pigment in the wall of the intestine. FIGURE 177. Philometra scomberesoxis Nikolaeva and Naidenova, 1964: a — anterior end, lateral; b — posterior end (after Nikolaeva and Naidenova, 1964). Pharynx small, with a large gland. Esophagus cylindrical, without widenings, distinct only in young females. Ventricle indistinct, but the esophagus passes into a glandular appendage which consists of several cells at the beginning of the intestine. Intestine wide, ending blind in the posterior part. Length 93.382 (77.00—105.240) mm. Width at the pharynx 0.396 (0.300—0.420) mm, at the nerve ring 0.555 (0.435—0.630) mm. Maximum width 1.812 (1.500—2.190) mm. Nerve ring situated 0.225 (0.120— 0.315) mm from the cephalic end. Esophagus 0.900 mm long. 312 Uterus wide, occupying the entire body cavity. It begins below the end of the esophagus and ends posteriorly at the end of the intestine. Ovaries situated at both ends of the uterus. At the cephalic end the ovary makes a turn below the nerve ring and is directed posteriorly. In the posterior part of the tail it turns sharply anteriorly. Uterus large, filled with filarioid embryos (larvae). Length of embryos 0.470 (0.375—0.527) mm, width 0.018 (0.016—0.019) mm. Cuticle of mature females thin and loose; it ruptures easily and the larvae released move actively in sea water. Young females. Coloration lighter than in adults, cream and slightly yellowish to light reddish brown. Length 38.0 (34.48—41.15) mm. Width at end of pharynx 0.256 (0.170—0.333) mm, at the nerve ring 0.428 (0.248—0.540) mm. Maximum width 0.794 (0.430—1.245) mm. Nerve ring situated 0.154 (0.084—0.255) mm from the cephalic end. Esophagus 0.861 (0.630—0.990) mm long and 0.090 (0.074—0.11) mm wide. The entire body of young females is filled with blastomeres (eggs at various stages of cleavage). Embryos still absent. Vulva already atrophied. Ovary 1.590 mm long. Reference: Nikolaeva and Naidenova, 1964, pp.126—168. 303 Philometra sebastodis Yamaguti, 1941 Host:, Sebastodes joyneri. Localization: pectoral fin. Distribution: Japan. Description (after Yamaguti, 1941). Male unknown. Female. Length 5.45—5.54 mm, maximum width 0.3—0.37 mm in mid- dle of body with blunt ends. Cephalic and caudal papillae absent. Nerve ring situated 0.12—0.13 mm from the cephalic end. Esophagus cylindrical, 0.24—0.27 mm long and 0.022—0.030 mm wide at the anterior widening. Ventricle cylindrical, 0.045 X0.018—0.021 mm large, with a relatively large process projecting into the intestine. The intestine narrows into a short thread and extends to the posterior end. Anterior ovary threadlike, 0.070— 0.090 mm long at the ventricle or beginning of the intestine. Posterior ovary 0.3—0.38 mm long, with a funnel-shaped thickening 0.032—0.045 mm wide around its narrower proximal end, which is directed anteriorly or poste- riorly at the posterior end of the uterus. Uterus filled with embryos, narrowing slightly at both ends, extending from the anterior end of the intestine to near the posterior end. Older embryos are markedly narrower and longer than younger ones, 0.17—0.19 mm long and 0.003 mm wide. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Ssudarikov, 1954, p.79; Yamaguti, 1941, р. 394; 1961, р. 7%. 313 Philometra spari Yamaguti, 1961 (Figure 178) Host. Siparus “miacrocepha ис”. Localization: swim bladder. DATS Сич Otte sia pale Description (after Yamaguti, 1961а). Male unknown. FIGURE 178. Philometra spari Yamaguti, 1961: a — anterior end of female, lateral; b — caudal end of female, ventrolateral (after Yamaguti, 1961a). 304 Female. Length 110 тата, maximum width 1.4mm at the cephalic end; body tapering at both ends. Cuticle with fine transverse striation. Head broadly rounded, without papillae. Esophagus 0.83 mm long, with a nar- row dorsal gland which extends its whole length and apparently continues into the ventricular part, which is situated between esophagus and intestine; the bulblike anterior widening of the esophagus is 0.12 mm wide. Ventricle 0.065 mm long and0.11 mm wide, consisting of gland cells filled with granules and ending in 3 valves which project into the intestine. Intestine narrowing posteriorly and ending blind, without an anus. Tail truncate, with 4 indistinct terminal papillae. The anterior ovary begins 0.22 mm from the cephalic end, turns posteriorly, winding before entering the uterus, which begins 1.5mm behind the cephalic end and is filled with rounded eggs of varying size (0.013—0.026 mm). The posterior ovary makes two loops at the posterior end. Reterence: Yamaguti, 1961а, ‘рр. 217—228. Philometra sydneyi Rasheed, 1963 (Figure 179) Host: "large white fish." Localization: subcutaneous tissue. Distribution: Australia. Description (after Rasheed, 1963). Male unknown. | Female. Length 30—150mm, width 0.35 mm. The cephalic end bears 8 papillae arranged in pairs in the outer ring and 4 papillae on the sub- 314 median line in the inner ring. Amphids protruding, situated laterally. Cuticle smooth. Mouth wide, opening directly into the esophagus. Near FIGURE 179. Philometra sydneyi Rasheed, 1963: а — anterior end; b— cephalic end, showing esophageal teeth; c — end of esophagus; d — posterior end of female, lateral (after Rasheed, 1963). 305 the mouth the esophagus forms а bulb and continues anteriorly in three sharp teeth which are strongly chitinized. Length of esophagus 4.92 mm, width 0.15mm at the anterior bulb. The large esophageal gland opens above the nerve ring and continues posteriorly to the end of the esophagus. Ventricle and process distinct. Nerve ring situated 0.3 mm from the cephalic end. Intestine of almost the same width as the posterior part of the esophagus, ending blind near the end of the tail. Two long, thin ovaries, extending in opposite directions and uniting in the common duct of the uterus. Vulva absent. Tail slightly tapering. Larvae 0.36 mm long,0.01 mm wide, with rounded head and pointed tail. Reference: Rasheed, 1963, pp.89—130. Philometra tauridica Ivaschkin, Kovaleva, and Khromova п. sp. (Figure 180) Hosts: Atherina mochon pontica, Trachurus) еде raneus ponte ws FIGURE 180. Philometra tauridica Ivaschkin, Kovaleva, and Khromova п. sp: а — cephalic end, apical; b — anterior end; с — caudal end of male. 306 315 Localization: body cavity. Dis triubwti on:)" Black Sea: Description. Light yellow nematodes. Cuticle smooth. Cephalic end with papillae, visible in apical view: 8 papillae in the outer ring, ar- ranged in pairs, and 16 papillae in the inner ring, arranged in groups of four. Esophagus entirely muscular, with an anterior widening. Nerve ring distinct. Uterus strongly developed, occupying the entire body cavity. Male. Length 2.981 mm, width 0.054 mm. Length of esophagus 0.075 mm, width 0.011 mm. Nerve ring situated 0.034 mm from the се- phalic end. Both spicules 0.088 mm long; gubernaculum 0.016 mm long. Female (not fully mature specimens). Length 11.032—30.228 mm. Measurements of a female 17.032 mm long: width 0.416 mm, length of esophagus 0.731 mm, width 0.067 mm; length of esophageal swelling 0.101 mm; nerve ring situated 0.127 mm from the cephalic end. Philometra zebrini Yamaguti, 1961 (Figure 181) Host: Zebrini zebrinus. Localization: subcutaneous connective tissue. Distribution: Japan. Description (after Yamaguti, 1961а). Male unknown. FIGURE 181. Philometra zebrini Yamaguti, 1961: a — anterior end of female, lateral; b — tail of female, lateral; c — transverse section of esophagus at the thickening; the single dorsal and the paired subventral esophageal glands are shown; d — transverse section of connection between esophagus and intestine (after Yamaguti, 1961a). 307 317 Female. Length 30—35 тата, width1.5—2mm. Воду cylindrical, blood-red in life. Cuticle smooth, transversely striated. The cephalic end usually projects markedly because the esophageal bulb projects anteriorly. Posterior end sometimes bilobed in mature specimens, but tapering in young specimens. Anterior esophageal bulb spherical, with triradial lumen, 0.3—0.5 mm wide; esophagus 0.1—0.16 mm wide, slightly widened behind its middle, where the lumen is flattened and displaced by the thick esophageal gland; subventral glands reduced at this point. The posterior end of the esophagus projects into the intestine and consists of a spongy parenchyma with large cells, appearing like a rudimentary ventricle. In the wall of the esophagus are two small, distinct vesicular structures which are probably swollen ducts of the esophageal glands, but they cannot be traced to the swollen part of the dorsal esophageal gland. At this part of the gland is a narrow duct with indefinite outline adhering to the nucleus; a distinct duct is not recognizable in the subventral gland. Intestine wide, ending blind near the caudal end. The tubular anterior ovary 0.05—0.08 mm wide extends straight posteriorly from the esophageal bulb to the anterior end of the uterus. The uterus begins slightly behind the anterior end of the intestine in young specimens but extends to the esophageal bulb in mature females. The posterior ovary is tubular, 0.09—0.13 mm wide and curves posteriorly around the posterior end of the uterus. Vulva absent. Length of larvae in uterus 0.41mm, width 0.017 mm, Reference: Yamaguti, 1961а, рр. 217—228. Subgenus Alinema Rasheed, 1963 Diagnosis. Philometra. Mouth surrounded by a row of cuticularized moniliform structures. The wide lumen of the anterior part of the esophagus is surrounded by 3 large lobes which occupy almost the entire width of its anterior part. Tail slightly pointed. Parasites of freshwater fish. Type species: Philometra (Alinema) alii Rasheed, 1963. Philometra (Alinema) alii Rasheed, 1963 (Figure 182) Hosts: Pimelodus sp., P.ipati, WPirinampus shee pirinampus, Localization: mesentery, intestine. Вов сое Brazil: Description (after Rasheed, 1963). Male unknown. Female. Length 150mm, width0.37 mm. Cephalic papillae in- distinct. An inner ring of submedian papillae is apparently absent. Mouth wide, the entire slit covered with moniliform cuticularized formations. The mouth passes directly into the wide esophagus; its three lobes occupy 308 the whole anterior part of the esophagus and project markedly. Nerve ring surrounding esophagus 0.37—0.4 mm from the cephalic end. Esophagus completely muscular, 0.95—1.8mm long, its posterior end with a muscular ventricle with a small process into the intestine. Intestine ending blind at the posterior end. Tail tapering. An ovary at each end of the body which is connected with the blind tube of the uterus; mature females have no oviduct or vulva. Uterus distended with larvae, which have a rounded head and pointed tail. Length of larvae 0.38—0.40 mm, width 0.020—0.023 mm. Description (after Inglis and Ogden, 1964). Male. Length 6.43—7.26 mm, width 0.10—0.16mm. Width of head 0.044—0.053 mm. Cephalic papillae very small, cuticle smooth. Length of esophagus 0.40 mm. Tail rounded, with two pairs of protruding papillae and a pair of very small papillae. One pair of the protruding papillae is situated behind the cloaca; the other pair is situated almost at the end of the tail. The pair of small papillae is situated between and slightly before them. Spicules thin and simple, right spicule twice as long as the left. Right spicule 0.69—0.83 mm long, left spicule 0.36—0.41 mm. Guberna- culum long, 0.10—0.12 mm long, a distinct denticle at the posterior end. 318 (317) FIGURE 182. Philometra (Alinema) alii Rasheed, 1963: а — posterior end of male, ventral; b — same, lateral, left side; с — same, lateral, showing the left spicule; d — anterior end of female; e — cephalic end, lateral; f — same, dorsoventral; g — caudal end of female; h — larva (a—c — after Inglis and Ogden, 1964; d—h — after Rasheed, 1963). Female. Length 46.8—93.2 mm, width 0.44—0.53 mm. Width of head 0.09—0.12mm. Length of esophagus 1.23—1.49 mm. References: Inglis and Ogden, 1964, рр. 523—525; Rasheed, 1963, pp. 389—130, Subgenus Ranjhinema Rasheed, 1963 Diagnosis. Philometra. Eight fleshy lobes arranged in four pairs on the submedian line; each lobe with two papillae of the same size 309 with small nerve endings. А pair of lateral amphids. The anterior part of the esophagus resembles an inverted flask with wide bottom. Tail with two large papillae at the end. Males unknown, Parasites of marine fish. Type species: Philometra (Ranjhinema) polynemii Rasheed, 1963. Philometra (Ranjhinema) polynemii Rasheed, 1963 (Figure 183) Host: Polynemus tetradactytlus:. Localization: body cavity. Distribution: Pakistan: Description (after Rasheed, 1963). Male unknown. (319) FIGURE 188. Philometra (Ranjhinema) polynemii Rasheed, 1963: a — anterior end of female, lateral; b — cephalic end, dorsoventral; c — same, lateral; 4 — caudal end of female, lateral; е — same, ventral; f — larva; g — posterior end of female, lateral; h — details of esophagus (after Rasheed, 1963). Female. Length 50—60 mm, width 0.9—1.2 mm. Cuticle smooth, but transverse fibers are distinct at some points and form bundles so that they appear like transverse bands below the cuticle. 310 —————— ТОНИ У Cephalic end rounded, width0.21—0.26 mm, with 8 fleshy papillae situated 319 submedially in four pairs and a pair of lateral amphids. The papillae have two lobes, and since they are fleshy, the nerve endings are difficult to see. Nerve ring surrounding the esophagus 0.23—0.26 mm from the anterior end. The esophagus has the form of a flask, with the base around the mouth; it then becomes slightly narrower and its width remains almost the same its entire length except at the posterior end, where it becomes again wider; it is widest in the anterior part. Length of esophagus 2.04—2.08 mm, width 0.16—0.18 mm. There is a large esophageal gland which extends to the end of the esophagus and opens near the nerve ring. Ventricle small, weakly muscular; intestine ending blind at the posterior end. Tail rounded, with two large papillae at the end. Anus and vulva atro- phied. Ovaries opposite, thin and long, uterus filled with larvae 0.38— 0.39 mm long, 0.015—0.016 mm wide. Reference: Rasheed, 1963, pp.89—130. 320 Genus Buckleyella Rasheed, 1963 Diagnosis. Philometrinae. Body cylindrical, with blunt, narrowing ends. Head with large lobelike papillae and a pair of amphids. Cuticle with rodlike structures arranged in a definite order, with grooves below them. Esophageal gland with а membrane. Ventricle and process present. Esophagus bulb-shaped at the anterior end, the three lobes projecting into chitinized teeth. Intestine ending blind. Ovaries opposite. Anus and vulva atrophied. Tail blunt, rounded, with two lateral papillae. Males unknown. Parasites of marine fish. Type species: Buckleyella buckleyi Rasheed, 1963. Buckleyella buckleyi Rasheed, 1963 (Figure 184) Eos © nor Iinrem us тата. Localization: mesentery. Distribution: Pakistan. Description (after Rasheed, 1963). Male unknown, Female. Length 46—110mm, width 0.3—0.5 mm. Body of uniform width its entire length, with tapering ends. There are sharp, needle- shaped or rodlike structures arranged in a definite order projecting from the cuticle. Below these structures are fibrillar formations in the form of warts which become more distinct when the body contracts and are difficult to see in extended specimens and in females distended with larvae; on the other hand, the ''rods'' are more conspicuous in females filled with larvae. 321 The cephalic end bears 8 oblong, flat, lobe-shaped papillae and a pair of amphids which are characteristic for this genus. Width of cephalic end 0.17—0.19 mm. Nerve ring situated 0.25—0.3 mm from the cephalic end, The esophagus begins at the mouth; it has a bulb-shaped widening 311 at the anterior end and continues anteriorly in three cuticularized teeth which project from the slit-shaped mouth. These teeth are pointed and are distinct in young forms; they are used for piercing blood vessels and tissues. (320) FIGURE 184. Buckleyella buckleyi Rasheed, 1963: a — cephalic end, showing the slightly retracted esophageal teeth, dorsoventral; Ь — same, another specimen; с — tail of female, lateral; 4 — same, dorsoventral; e — section of esophagus of a young female, showing esophageal teeth; f — posterior end of female, showing the “warts” (contracted specimen); р — larva; В — tail of female, dorsoventral, showing the rodlike formations (after Rasheed, 1963). Esophagus entirely muscular, 3.96—5.1 mm long, width 0.14—0.15 mm at the anterior bulb. The esophageal gland is not very distinct and has a membrane; ventricle indistinct. The intestine has a reticulate epithelium and ends blind at the posterior end. Anus absent. Tail rounded, with two large lateral papillae so that the tail appears bifid in some specimens, Two ovaries, situated near the caudal and cephalic end, passing into the uterus, which is filled with larvae. Larvae large, with rounded cephalic end and sharply tapering tail, 0.39—0.48 mm long, maximum width 0.0125—0.015 mm. Reference: Rasheed, 1963, pp.89—130. Genus Ichthyofilaria Yamaguti, 1935 Diagnosis. Philometrinae. Body cylindrical, with pointed ends. Mouth simple. Esophagus divided into a muscular anterior part with two swellings separated by the nerve ring and a short posterior part with a glandular appendage projecting into the intestine. Between the two parts of the esophagus is a long, vermiform gland which ends in the anterior part of the intestine. This becomes narrower posteriorly, where it is not functional, continuing in a tube to the end of the body. Anterior ovary rudimentary; posterior ovary cylindrical, extending to the posterior end of the body and returning to the uterus. Uterus containing larvae. 312 Parasites of the body cavity of fish. Туре and-only species: Ichthyofilaria dasycotti Yamaguti, 1935. Ichthyofilaria dasycotti Yamaguti, 1935 (Figure 185) Host. Wasycottustsetiper. mocalixnat ron: body cavity. Distribution: Japan. Description (after Yamaguti, 1935). Male unknown. Female. Length 30.43—38 mm, width 0.5—0.8 mm. Body cylindrical, pointed at both ends. Nerve ring situated obliquely, 0.18—0.246 mm from 322 the cephalic end. Mouth simple, Esophagus divided into a muscular anterior part and a short posterior part which is 0.29—0.348 mm long. The esophagus has two thickenings which are divided by the nerve ring and which are of varying width (0.12—0.18 mm) according to the contraction of the body. The posterior part of the esophagus is narrower than the anterior part and forms a small glandular appendage which projects into the intestine. Dorsal to the connection of the two parts of the esophagus 15 a long, vermiform gland containing a few nuclei. The irregularly widened intestine contains amorphous reddish brown inclusions and forms posteriorly a narrow, nonfunctional tube which ends blind in an awl-shaped swelling at the end of the tail. Anterior ovary atrophied into a vermiform process 0.15mm long, posterior ovary cylindrical, 2.94 mm long and 0.19mm wide. The wide, tubular uterus begins 3.31 mm from the cephalic end and 3.35 mm from the posterior end. Uterus filled with larvae 0.160—0.180 mm long and 0.003 mm wide. Vagina and vulva not recognizable. тат /0 тат а b FIGURE 185. Ichthyofilaria dasycotti Yamaguti, 1935: a — anterior end; b — posterior end; с — cephalic end (after Yamaguti, 1935). 313 References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.80; Yamaguti,1935, рр.357—358; 1961b, p.78. Genus Nilonema Khalil, 1960 Diagnosis. Philometrinae. Large nematodes, body with tapering, pointed ends. Cuticle with large conical processes. Esophagus cylindrical, without an anterior widening. Cephalic papillae small. Ovaries opposite. Anus and vulva atrophied. Males unknown. Parasites of freshwater fish. Type species: Nilonema gymnarchi Khalil, 1960. Nilonema gymnarchi Khalil, 1960 (Figure 186) Host: Gymnanchus wilotieus: Localization: swim bladder. (323) FIGURE 186. Nilonema gymnarchi Khalil, 1960: a — anterior end of female, lateral; b — tail of female, lateral; c — adult female, lateral; d — cuticular conical process; e — transverse section of posterior part of female; f— same, in middle of body; g — larva (after Khalil, 1960). 314 Distribution: Sudan. Description (after Khalil, 1960). Male unknown. Female. Length of largest specimen 115 mm, width 1.2mm. Body of almost uniform width its entire length, with pointed ends. Cuticle relatively thick, with numerous irregularly distributed conical, teatlike 323processes. The number of these structures is very variable and depends to some extent on the size of the specimen. There were 75 on the largest specimen, The anterior processes are situated near the cephalic end and are scattered irregularly on the dorsal and ventral side of the body. They are more numerous and are arranged more densely in the caudal region to the end of the tail. Their size decreases toward the end of the tail, but it is more or less constant in the middle of the body, where they are 0.16—0.27 mm high and 0.15—0.3 mm wide at the base. Mouth simple, opening directly into the short, cylindrical, muscular esophagus which is 0.57—0.81 mm long and 0.05—0.06 mm wide. Esopha- geal gland absent. Nerve ring situated 0.18—0.27 mm from the cephalic end. Intestine with relatively thick walls, wider than the esophagus. Its lumen becomes gradually narrower posteriorly and finally disappears, the 324 intestine forming a cord which is attached to the body wall. The long, cylindrical, curved ovaries are situated at each end of the body. Posterior ovary usually sinuous. The two ovaries are connected with the uterus by a narrow oviduct. Uterus large, with thin walls, occupying almost the entire body cavity. Vagina and vulva absent. Length of larvae 0.48—0.54 mm, width 0.014 mm. Cephalic end blunt, tail long and pointed; intestine, esophagus and anal pore are visible. me neGence: Khalil, 1960, pp. 55—58. Nilonema senticosa (Baylis, 1927) Rasheed, 1963 (Figure 187) Synonyms: Philometra senticosa Baylis,1927; Philometroides senticosa (Baylis, 1927) Travassos, 1960 Host: Arapaima gigas. Localization: body cavity. Distribution: South America. Historical review Nilonema senticosa was described from the swim bladder of Arapaima gigas in Brazil and was placed in the genus Philometra. Travassos (1960) transferred the species to the genus Philometroides Yamaguti, 1935. Rasheed (1963), after a study of Baylis' types in the British Museum and specimens of Nilonema gymnarchi Khalil, 1960, showed that the characters of Philometra senticosa resembled those of Nilonema more than those of Philometroides. Rasheed thought that the general habitus, the conical tubercles on the cuticle, the cylindrical esophagus, and the absence of a large esophageal gland prove that Baylis' species belongs to Nilonema and not to Philometroides. 315 Description (after Baylis, 1927). Male unknown. Female. Length 90—125 mm, width 0.62—0.85 mm. Cephalic end markedly more pointed than posterior end. Cuticle thick, apparently consisting of two layers. It bears a row of spinelike processes with a refracting surface. Under high magnification, these uneven outgrowths all over the body, including the anterior part of the esophagus, make the nematode look like a stem of bramble. Cephalic end rounded [sic], mouth simple, without lips. Cephalic papillae indistinct. Esophagus indistinctly divided into a muscular and a glandular part, very narrow anteriorly, except for a slight widening around the mouth before the nerve ring. Behind the nerve ring the esophagus gradually widens into a club. A narrow ''neck'' separates the esophagus from the intestine. Three valves between esophagus and intestine. The esophagus is apparently enclosed in a granulate membrane. Length of esophagus 1.37—1.5 шп. Nerve ring situated 0.35—0.4 mm from the cephalic end; slightly behind it is the small, semitransparent excretory pore, which is apparently connected with a group of cells in the body wall. The intestine extends almost through the whole body in the form of a wide tube formed by 325large cells. The intestine apparently ends blind, adhering to the body wall. Vulva absent. Branches of uterus opposite, forming a continuation of the tube which extends almost from one end of the body to the other and is connected at each end with the short, coiled ovaries. Viviparous. Uterus filled with larvae with blunt cephalic end and markedly pointed, conical tail. 0.3 mm FIGURE 187. Nilonema senticosa (Baylis, 1927): a — anterior end of female, lateral; b — posterior end of female, lateral; с — cephalic end; 4 — anterior end; e, f — posterior end, ventral (a, b — after Baylis, 1927b; c—f — after Travassos, 1960). Description (after Travassos, 1960). Male unknown. Female. Length 60—130 mm, width 0.41—0.71 mm, Cephalic end blunt, without ornamentation. Posterior end conical. Cuticle with distinct transverse striation and large conical spines of varying size (0.043—0.073mm) 316 irregularly distributed from the end of the esophagus to the anus. Muscula- ture polymyarian, body with narrow, indistinct longitudinal fields. The esophagus begins at the mouth, which is a triangular slit without formations. Esophagus club-shaped, consisting of a cylindrical anterior part and a widened posterior part. Length of esophagus 1.2—1.4 mm, width 0.12— 0.157 mm. Nerve ring situated in the cylindrical part, at its anterior third. Intestine cylindrical,dark. It ends in a narrow but permeable rectum. Anus situated 0.063—0.081 mm from the posterior end. Genitalia amphi- delphic. The anterior ovary forms several loops at the end of the eso- phagus. The posterior ovary extends to the anus. Uterus large, saclike, occupying the entire body cavity, pressing the intestine to the side and the ovaries to the ends of the bodv. Females filled with first-stage larvae. Vulva and ovejector not found. Larvae in uterus about 0.300 mm long, width 0.014 mm. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.79; Baylis,1927b, pp.35—47; Rasheed, 1963, рр. 89— 130; Travassos,1960, pp.15—20. 326 Genus Philometroides Yamaguti, 1935 Diagnosis. Philometrinae. Body cylindrical, long, with blunt ends and numerous cuticular thickenings on the cuticle. Parietal muscle cells strongly developed on the dorsal and ventral side and distinctly flattened in the peripheral zone. Cephalic and posterior end without papillae. Mouth simple. Esophagus partly swollen anteriorly, with a well-developed dorsal esophageal gland and a muscular ventricle, and also with a glandular appendage which projects into the intestine. Intestine, ovary, uterus, and larvae resembling those of Philometra. Viviparous. Parasites of marine fish. Type species: Philometroides seriolae (Ishii, 1931). у Philometroides seriolae (Ishii, 1931) (Figure 188) Synonym: Filaria seriolae Ishii, 1931 Hosts: marine fishes. Localization not given. Wrstripution: ‹ Japan. Description (after Yamaguti, 1935). Male unknown, Female. Length 38 mm, width1.7mm. Body of about the same width its entire length, blunt at the ends, cuticle thick, with irregularly scattered flattened, teatlike thickenings. The muscle cells of the surface are strongly developed, particularly on the dorsal and ventral side; their peripheral compressed part is platelike and stains strongly with eosin. The two granular midlines are narrow in dorsoventral view and elongate in trans- 327verse section. The flattened wide lateral fields consist of fine fibrils with 317 numerous nuclei. Mouth simple, opening into the slightly swollen, cylindrical muscular anterior part of the esophagus which is 4.6 mm long, with a large, dorsal gland beginning at the nerve ring, 0.4mm from the cephalic end, (326) 20mm | | i Zi | МАХ) FIGURE 188. Philometroides seriolae (Ishii, 1931): a — anterior end; b — posterior end; с — transverse section (after Yamaguti, 1935). Length of gland 3.6mm, width 0.4mm. The muscular ventricle with a small glandular appendage projects into the intestine. Ovary long and thin, ex- tending to the ends of the body. The uterus contains embryos 0.6 mm long, with thin tail. Vagina and vulva absent. Description (after Yamaguti, 1941). Body cylindrical, long, with blunt ends and numerous cuticular thickenings. Cephalic and posterior end without papillae. Mouth simple. Esophagus partly swollen anteriorly, with a well-developed dorsal gland and a muscular ventricle, and also with a glandular appendage which projects into the intestine. ~ Male unknown. Female. Length 38 mm, width1.7mm. Body of about the same width its entire length, blunt, cuticle relatively thick, with irregularly scattered, flattened, teatlike thickenings. Length of esophagus 4.6mm, Nerve ring situated 0.4mm from the cephalic end. Ovary long and thin, extending to the ends of the body. The uterus contains embryos 0.6mm long. Vagina and vulva absent. Viviparous. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov,1954, p.79; Ishii, 1931, p.207; Yamaguti, 1935, pp. 355—356; 1941, pp. 343—396. Philometroides anguillae (Ishii, 1916) Rasheed, 1963 (Figure 189) Boasts; Anguilla rain hoardtit. Aa. aD OBL C ae Localization: body cavity. Distribution: Queensland. 318 Historical review The nematode from the eye orbit of Anguilla japonica, described as Filaria anguillae Ishii, 1916, was transferred by Yamaguti (1935) to the genus Philometra. Rasheed, who knew the description of Ishii and the illustrations, found that her specimens and Ishii's species were similar. Yamaguti stated that there were 6 cephalic papillae, two lateral and two on the dorsal and two on the ventral side. Rasheed found 8 distinct cephalic papillae and a pair of amphids. Ishii illustrated two protruding papillae at the posterior end, but in Rasheed's specimens the tail was almost rounded; only in one specimen she found two papillae on the ventral surface; the papillae were indistinct in the other two specimens. Except for the difference in the number of papillae, the species of Ishii resembles Ph. anguillae ша! respects. The genus of host is also the same. Description (after Rasheed, 1963). Male unknown. Female. Length 88—100 mm, width 0.7—0.8mm. Cuticle smooth, but under high magnification several tubercles are visible near the tail and 328 numerous tubercles scattered on other parts of the body. Body of uniform width its entire length except at the ends, where it tapers slightly. FIGURE 189. Philometroides anguillae (Ishii, 1916) Rasheed, 1963: a — anterior end of female, lateral; b — cephalic end of female, lateral; с — posterior end of female, dorsoventral (after Rasheed, 1963). Cephalic end with 8 papillae and a pair of amphids. Length of esophagus 1.59—1.62 mm, width of the swollen anterior part 0.17—0.19mm. There is a small ventricle with a process projecting into the intestine. Nerve ring situated 0.18—0.20 mm from the cephalic end. Anus absent, intestine ending blind near the posterior end. The anterior ovary forms loops above the esophagus; posterior ovary situated near the posterior end. Uterus filled with fully developed eggs and embryos, which are 0.43—0.47 mm long and 0.010—0.012 mm wide. Tail rounded, with two papillae situated ventrally, near the end of the tail. References: Rasheed, 1963, pp.89—130; Yamaguti, 1935, pp. 347—386. 319 Philometroides denticulatus Rasheed, 1965 (Figure 190) Hosts:.Pristipoma hasta, Оо Швы. ruber, Localization: body cavity, swim bladder. Distribution: Pakistan. Description (after Rasheed,1965). Coloration yellowish white. Body tapering at both ends. Cuticle thin, with prominent warts on the whole body which are formed only by the cortical layer of the cuticle. Under the cortical layer is a small point or nucleus in the wart which sometimes appears like the pore of an amphid. Male unknown. (329) FIGURE 190. Philometroides denticulatus Rasheed, 1965: а — anterior end, lateral; b — esophagus, removed from body; с — cephalic end, apical; d — cephalic end; е — posterior end of female, ventral; f — papillae on part of the cuticle; g — anterior end of larva; h — posterior end of larva (after Rasheed, 1965). Female. Length 250—320 mm, width 0.52—0.67 mm. Esophagus thickened anteriorly in the form of a bulb, which projects from the mouth in three small parts resembling teeth. On each side of the anterior part of the esophagus there are muscles which probably move the esophagus. Cephalic papillae very small and difficult to see. Length of esophagus 1—2.15 mm. Esophageal gland saclike, with a large nucleus in the middle, 329 extending along the entire esophagus and opening slightly before its widening, about 0.060 mm from the cephalic end. The esophagus is not divided posteriorly and is connected with the intestine by small valves. Nerve ring surrounding esophagus 0.38—0.45 mm from the cephalic end. 320 The ovaries begin at the ends of the body and continue into the curved, saclike uterus, which is filled with larvae 0.63—0.73 mm long and 0.009— 0.011 mm wide. Some larvae which have been discharged from the fe- male already have a rudimentary genital duct and an anus situated 0.1mm from the end of the tail. Toothlike formations and two laterally protruding papillae are present at the cephalic end. Tail rounded; two small papillae present at its end. Anus absent. Reference: Rasheed, 1965, pp. 349—362. 330 Philometroides lusiana (Vismanis, 1966), n. comb. (Figures 191,192) оу mis: Ри Помеека Таз Visman,. 19625 3b bid ome tira lusiana Vismanis, 1966 EWOrsntac © yapioia W'S)! саврио. Localization: under scales, in scale pouches, body cavity, wall of swim bladder. Bisel wih ion: Ussik. Historical review This species was described in 1962 аз Philometra lusii Visman, 1962, but since there were mistakes in the original description of the female, the author redescribed the species and, in accordance with the rules of nomenclature, renamed it Ph. lusiana Vismanis, 1966. Description (after Vismanis,1967, as Philometra lusiana),. Males markedly shorter than mature females (ratio about 1:50). Male. Length 2.9—3.5mm, width 0.038—0.045 mm. Body almost of uniform width its entire length, tapering at the ends. Anterior end narrower than posterior end. Width of anterior end 0.015—0.016 mm at the pharynx, width of posterior end 0.024—0.027 mm at the terminal widening. Cuticle smooth, thin. Cephalic end rounded. Mouth apical, Posterior end slightly curved ventrally and widened, forming collarlike formations in two spongelike lobes. Intestine opening at the posterior end, and the pointed spicules projecting subterminally. Spicules0.19to0.23 mmlong; their width in the middle about 0.0027 mm. Gubernaculum 0.051—0.062 mm long, slightly curved, with a characteristic tooth at the distal end which resembles a pointed arrow. Length of tooth 0.0079—0.0084 mm. Female. Length115 mm, width 1mm. Females are pink or red in life, threadlike. Cuticle with numerous papillae 0.014—0.025 mm long scattered irregularly on the whole body. There are about 2000 papillae on a female 70 mm long. Body tapering at both ends. Anterior end with 4 cephalic papillae. Mouth trihedral, leading into the esophagus, which is straight with a slight widening anteriorly. In a female 9 cm long and 0.8mm wide, the eso- phagus is 2 mm long and 0.07—0.09 mm wide. Intestine brownish red, narrowing posteriorly and ending blind. Anus and vulva atrophied in adults. 321 (331) Е Е >» | $ FIGURE 191 Philometroides lusiana (Vismanis, 1966): a — cephalic end; b — anterior end, lateral; с — posterior end of female; 4 — posterior end of female; е — 1агуа from Cyclops during first molt; f — same, during second molt; р — fourth-stage larva from the swim bladder of carp 23 days after infection (a—d — after Vismanis, 1966; e—g — after Vasil'kov, published for the first time). 322 332 Posterior end with 4 paired processes of different size and form. Two of them are distinct processes of the body at the end of the tail. They are about 0.04 mm long. Between them are two smaller cuticular lateral processes, one of which is sometimes small. Nerve ring situated 0.165— 0.242 mm from the cephalic end in females 6—9 cm long. The entire body cavity of mature females is occupied by the uterus, which contains numerous rounded eggs, 0.034 to 0.040 mm wide at the end of winter. The eggs have a thin shell. The larvae develop in May. Ovaries small, cylindrical, with smooth surface, situated at both ends of the body. Viviparous. Ee Re Oe ge < ago pee вы ково == Нов, SS ewe к. ЕЕ FIGURE 192. Philometroides lusiana (Vismanis, 1966): а — anterior end of female; b — posterior end of female; с — posterior end of male; 9 — carp infected with Ph. lusiana (after Vasil'kov, published for the first time) 323 333 Young, unfertilized females and males are found in the body cavity and in the wall of the swim bladder during the copulation period. The females closely resemble the males in size. Young females 2.3 to 3.5 mm long, width at the genital pore 0.0297—0.0375 mm. Surface smooth, without papillae. Ends of body rounded, cephalic end slightly more pointed than the posterior end. In young females the intestine is well developed and opens at the posterior end. After fertilization, the body of the female becomes covered with small papillae, cephalic and caudal papillae develop, the genital pore and anus atrophy. The larvae grow in the uterus in spring. When they come into contact with water, mature females break up and the larvae are released. Length of larvae 0.40—0.47 mm, width 0.015—0.019 mm. The larvae are threadlike, with thin, smooth cuticle. Their cephalic end is blunt and rounded, the tail pointed. Biology. Vismanis (1964) studied the life cycle of the parasite. He found that it lasts a year,from June to May. Intermediate hosts may be any of the following: Macrocyclops albidus, Eucyclops ser- ги] ао ©. macruroides, Мевосус1орз' Ченскава ана Cyclops strenuus. The larvae grow in the body cavity of the host. They molt after 3—4 days; development in the intermediate host lasts 6—7 days, and the larvae then become infective. Carp become infected in June when they swallow infected water fleas. The copepods are digested and the larvae are released, penetrate through the wall of the intestine and enter the body cavity. Five days later they concentrate around the swim bladder, gonads, and kidneys and grow and develop into males and females. This lasts about one month, and the females are then fertilized. At this time the young forms are of about the same length: females 2.3 to 3.5mm, males 2.9 to 3.5 mm. After fertilization of the females, the males penetrate into the wall of the swim bladder; the females leave the body cavity and move to their permanent location in the scale pouches, where they are found at the end of June or in early July. The females grow rapidly: they are 4—8 mm long at the end of June — early July, 30 mm in early August, 60 mm in October—November, and 100—110 mm in the following spring. In the middle of May, when the water temperature is 11°C, larvae begin to develop and become 0.4—0.45 mm long. In Latvia, the larvae are released into the water at the end of May —early June when the water temperature is 15—20°C. Vasil'kov (1968b) gave data on the life cycle of the species in the central regions of the RSFSR; he used in his experiments Acantho- eyclops Virnidis, “A.yvernalis,(Eucyeclops macruroioeen Cyclops strenuus, and Mesocyclops leuckarti as experimental intermediate hosts. The larvae molt first after 3—4 days; the second molt takes place after 7—8 days, and they are then 0.60—0.65 mm long. In the intestine of the definitive host, they penetrate into the wall of the intestine and then into the body cavity; 5—8 days after infection, the larvae are localized on the serosa and after 14—15 days in the liver and kidneys, where they molt the first time in the definitive host. After 18—21 days the larvae are found in the wall of the swim bladder and molt the second time; after 30—40 days sexual differentiation has ended, and the larvae are 2.9—4.5 mm long. Fertilization now takes place. 8 324 334 At the end of August and in September the females move into the scale pouches, where they remain throughout autumn, winter and spring (8—9 months). The males remain in the swim bladder. Vasil'kov found that the life of females lasts 11—12 months and of males 13—14 months. Vismanis stated that Ph. lusiana is at present widely distributed in the European Soviet Union; infections have been recorded in pond fisheries in Latvia, Belorussia, European Russia, andthe Ukraine. Vasil'kov (1964, 1967a, b, 1968) found the disease in carp in the Kaliningrad, Minsk, Ternopol, and Moscow regions. The clinical symptoms of the disease appear when the parasites enter the scale pouches and are as follows: small tubercles with slight hemor - rhages appear on the anterior part of the body. Damaged, swollen scales often form a kind of collar around the head. In places where the nematodes come into contact with the scales the pigment of the scales is partly des- troyed, forming a kind of imprint of the worm. When the larvae enter the water in spring, the points of localization become red, so that they appear like ulcers observed in erysipelas. The nematodes have a toxic effect on the carp and apparently feed on the blood of the host (Vasil'kov, 1964). All ages of carp contract the disease, but yearlings and two-year-olds are most heavily infected (to 70—90%). The intensity of infection is also high, 25—35 specimens (to 50—75). If a young fish contains more than 3 nematodes it dies; it probably cannot tolerate the migration of fertilized females from the body cavity to their permanent location in the scale pouches. Carp one and two years old infected with Philometroides have a low commercial value and are usually rejected. During the migration, the larvae increase in size and molt; hemorrhages are found in the intestine, liver, and swim bladder. Infected fish do not attain standard weight and are usually sluggish. Diagnosis is made by finding the parasites in the scale pouches. Where tubercles and raised scales are found the scales are taken with a forceps and the parasites are found coiled under them, singly or sometimes in twos or threes. Adult females are found fromthe hemorrhages, swollen scales, imprints of the parasites on the scales, and their visibility through the scales. On the venter, where the scales are white, the nematodes are clearly recognizable. The parasites are usually found on the head and opercula, sometimes in the tissue of the operculum and in the eye orbit. Vismanis and Vasil'kov recommend the following measures of prevention and control: 1. Transport of carp for breeding purposes from farms where the disease is prevalent should be forbidden. 335 2. Rearing and foraging ponds in which infected fish are found should be cleaned and kept without water during the winter. 3. In main ponds, which supply water to other ponds, older carp should not be released to feed. 4, To prevent infection of young fish, stock from the spawning ponds should be removed immediately after spawning. 5. Young fish should be removed from the spawning ponds not later than 6 days after spawning, because the larvae of Ph. lusiana are not infective for 6 days. Vasil'kov obtained good results with the following anthelmintics: diethylcarbamazine citrate and ditrazine base (30 or 40% aqueous soluticn) 325 for the treatment of spawners and young stock. Не recommends doses of 0.3—0.4 g/kg of weight; the fish are treated in spring, 2—3 weeks before spawning, or in autumn, during transfer to the wintering ponds, at a water temperature of 12—14°C; the preparation is less effective at lower tempera- Lures: References: Ivashkin and Khromova, 1965, pp.98—99; Kuz'movich, 1964, pp.66—67; Vasil'kov, 1967a, рр. 449—452; 1967b, pp. 62—64; 1968a, pp. 53—56; 1968b, pp.28—30; Vismanis, 1962, рр. 93—96, 1964, рр. 192—193; 1966, рр.9—10; 1967, рр. 759—761. Philometroides maplestonei (Travassos, Artigas, and Pereira, 1928) Rasheed, 1963 (Figure 193) Synonym: Philometra maplestonei Travassos, Artigas, and Pereira, 1928 Ро uo rablmeinandn Geli aare 0 Localization: body cavity. овен ето: Brazil: Description (after Travassos, Artigas, and Pereira, 1928, аз Philometra maplestonei). Male unknown. FIGURE 193. Philometroides maplestonei (Travassos, Artigas, and Pereira, 1928): a — anterior end of female, lateral; b — detail of cuticle; c — tail of female, lateral; d — larva (after Travassos, Artigas, and Pereira, 1928). Female. Length 144mm, width 0.22 mm; cuticle with small processes 336irregularly distributed on the whole body. Mouth rounded, without an armature; esophagus divided intoathinner anterior part about 0.28 mm long and 0.05 mm wide and a posterior part 0.43 mm long and 0.10 mm wide. 326 Anus very small, situated 0.048 mm from the posterior end; rectum very thin, difficult to find. Tail finger-shaped, posterior end tapering from the end of the genital ducts, about 0.65 mm from the posterior end. Vulva not found; loops of genitalia extending to the end of the esophagus anteriorly. Nerve ring situated inthe posterior part of the anterior part of the esophagus. Embryos about 0.320 mm long and 0.18 mm wide, esophagus already differentiated, about 0.100 mm long, tail ending ina widening inthe form ofa racket, so that it appears bifid. Reference: Travassos, Artigas, and Pereira,1928, p.27. Philometroides masu (Fujita, 1940) Rasheed, 1940 (Figure 194) Synonym: Philometra masu Fujita, 1940 Most Олеогнуневиь шасон.. moc Wirz tone. body саму. Distribution: Japan. 0.70 mm 0/0 mm FIGURE 194. Philometroides masu (Fujita, 1940): a — cephalic end, dorsal; b — posterior end; c — larva; d — eggs and larvae (after Fujita, 1940). Soi DWeseription (after Nujita, 1940, as Philometra мази). Male unknown. 927 Female. Length 16.5—63.0 mm, width 0.32—0.85 mm. Cuticle thin, with numerous small papillae irregularly scattered on the body. Cephalic end rounded, mouth simple, without lips. Length of pharynx 0.11 mm, width 0.08 mm ina female 16.50 mm long. Pharynx constricted in the middle and longitudinally striated on the whole surface. Anterior part of esophagus 0.46 mm long, posterior part 0.99 mm, width 0.17 mm. Intestine. 14.68 mm long, 0.13 mm wide anteriorly, ending blunt 0.26 mm from the posterior end. Rectum and anus atrophied. Nerve ring situated 0.34 mm from the cephalic end. Vulva and vagina atrophied in mature females. The vulva may be found with difficulty in the anterior third of the body in some immature specimens. Uterus single, with thin wall, wide and straight, 13.12 mm long and 0.28 mm wide. Ovaries paired, opposite; anterior ovary 2.50 mm long, turned posteriorly before the posterior part of the esophagus; posterior ovary 2.25 mm long, ending 0.18 mm before the end of the tail. Eggs oblong, 0.040 mm wide. Larvae threadlike, 0.75 mm long and 0.03 mm wide, with a distinct thick-walled esophagus and anterior part of the intestine. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.77; Fujita, 1940, pp.389—390; Yamaguti, 1941, p. 393; 1961а, pp. 217-218, 1961b, р. 76. Philometroides nodulosa (Thomas, 1929) Dailey, 1967 (Figure 195) Synonym: Philometra nodulosa Thomas, 1929 Hosts: Catostomus commersoni, Huro salmoudes, Localization: body cavity. Distribution: North America. Historical review Thomas found a single mature female of this species in 1927 and studied experimentally some aspects of development of the nematode. Dailey (1967) recorded the species from the headof Catostomus commersoni, He described the male (description not available). Dailey identified it as Philometroides nodulosa (Thomas, 1929) Dailey, 1967. Description (after Thomas,1929,as Philometra nodulosa), Male unknown, Female. Length 28 mm, width 0.456 mm. Body brilliant white in life. The body tapers from the middle toward both ends, cephalic end rounded, posterior end truncate. High, light-refracting processes on the cuticle. They are uniformly distributed on the entire surface so that they form rows of incomplete rings; distance between rings 0.004—0.124mm. Height of processes 0.006 mm, length 0.415 mm. Cuticle transversely 338 striated. Three layers of the cuticle can be distinguished after triple staining after Mallory: an outer layer less than 0.001 mm thick and two layers 0.002 mm thick together. Mouth terminal, surrounded by 6 small lips situated in a shallow depres- sion. The dorsal and ventral lips are sometimes turned back and slightly larger than the lateral lips. 328 Esophagus with muscular anterior part in the form of a bulb which extends for 0.010 mm. This widening is 0.049 mm wide and behind it the esophagus is 0.040 mm wide and weakly muscular. Before and dorsal to the widening there is a small glandlike body. The esophagus is connected with the wide, degenerating intestine 0.15 mm wide which gradually narrows and extends near the body wall. Anus absent. 0 00 thy 090 1000 | (i WLU UM ПИ FIGURE 195. Philometroides nodulosa (Thomas, 1929): a — anterior end of female, lateral; b — posterior end of female, ventral; с — cephalic end; d—f — eggs from uterus at different stages of development; g — larva from uterus; В — Cyclops brevispinosus infected with larvae (after Thomas, 1929) The shorter anterior ovary is 0.332 mm wide. Posterior ovary rudi- mentary. Uterus filled with mobile larvae and eggs at different stages of development. Eggs with thin hyaline shell, ovoid in the early stages, 0.012 X 0.014 mm large. Lateral lines wide and distinct, about 0.014 mm wide in middle of body. About 54 muscle fibrils on each side of the dorsal or ventral nerve extend into the body cavity, forming muscular margins at the sides. The muscle strands extend almost to the midline of the lateral fields and are visible 339 after modified triple staining after Mallory. Description of larvae from the uterus. Length 0.316 mm. All larvae of the same size. Width 0.006 mm 0.002 mm from the cephalic end. The body becomes conical and pointed, tail threadlike, with small papillae atthe end; ‘Cuticle finely striated, 329 Anus situated оп the ventral surface, 0.048 mm from the posterior end. p A convexity is present around the anus. | Two lips around the mouth, one dorsal and one ventral. Esophagus thin, occupying anterior third of body. Intestine wide, distinct because of the presence of strongly refracting granules. Large cells present which may be developing gonads, Two indistinct dark brown bodies in the middle of the intestine. Segmented glands at the posterior end, opening 0.040 mm from end of tail. | Life cycle. Thomas (1929) observed that when the cuticle of a female bursts in the water, the larvae are released; they are very mobile; some adhere to the bottom with the tail, coiling into a spiral. Under experimental conditions, the larvae survive 1—3 days in water. Thomas observed that Cyclops brevispinosus swallowed the larvae, which increase in size in the host and remain viable 6—7 days, until the Cyclops dies. Experi- ments to infect the definitive host with these parasites were negative, probably because the larvae did not become infective inthe Cyclops. Dailey (1967) reported successful infection of Cyclops vernalis and С. thomasi with larvae of Ph. nodulosa; the larvae reached the third stage after 20 days inthe host. Dailey also successfully infected the definitive host. He found third-stage larvae in the swim bladder and kidneys 9 days after infection. The larvae moved at first ventrally and then anteriorly, to the head during the fourth and fifth stages. Copulation 2 was observed at the early fifth stage in females and during the anterior migration in mature males. The males then died, but the fertilized females continued to grow. The entire development lasts 146—365 days, depending on the temperature. Dailey stated that the parasites are only slightly pathogenic for the definitive host. References: Dailey, 1967, рр. 1266—1267; Thomas, 1929, рр. 193—199. Philometroides plectroplites (Johnston and Mawson, 1940) Rasheed, 1963 (Figure 196) Synonym: Philometra plectroplites Johnston and Mawson, 1940 HoOsteiPlLecraoephites anvb ie wits, Localization: body cavity. Distribution: Australia. 340 Description (after Johnston and Mawson, 1940, аа Philometra pleetroplites).: Male unknown, Female. Length over 105 mm. Cuticle with numerous irregularly scattered warts. Cephalic end rounded, without lips or papillae. Length of esophagus 1.05 mm, width 0.09 mm; its widened anterior part contains a small, almost hemispherical vestibule 0.048 mm long and 0.056 mm wide. Nerve ring situated 0.2 mm from the cephalic end. Vulva and anus atrophied. The large uterus occupies almost the entire body cavity; it passes into the oviduct about 0.5mm from the cephalic end. Length of larvae in the uterus _ 0.47 mm, width about 0.022 mm; they are coiled into two spiral turns. Cephalic end of larvae rounded, posterior end pointed. 330 2 b FIGURE 196. Philometroides plectroplites (Johnston and Mawson, 1940) Rasheed, 1963: а — anterior end; b — same, lateral; с — posterior end of female, lateral (a — after Johnston and Mawson, 1940; b,c — after Rasheed, 1963). Description (after Rasheed, 1963). Male unknown, Female. Length 105 mm, cuticle with numerous irregularly scattered tubercles. Cephalic papillae present. Esophagus widened anteriorly, 1.2 mm long and 0.2 mm wide in the widened part. Nerve ring situated 0.2mm from the cephalic end. Uterus filled with larvae coiled into a spiral; larvae 0.45—0.47 mm long and 0.022—0.023 mm wide. Tail rounded. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р. 78; Johnston and Mawson, 1940, p.349; Rasheed, 1963, pp. 89—130. Philometroides sanguinea (Rudolphi, 1819) Rasheed, 1963 (Figure 197) Synonyms: Filaria sanguinea Rudolphi,1819; Philometra sanguinea (Rudolphi,1819); Philometra trilabiata Belouss, 1965; Phiemwearas sin Ishii, 1933 Oss: Garassius ‘Carassius, С. auratws ic ibelmor ое а та тон” ас. Distribution: Europe, Asia. Historical review This species was described by Rudolphi (1819) as Philometra sanguinea, which he found under the skin of the caudal fin of Cyprinus gibelio (=Carassius auratus gibelio). Rudolphi's description was not very detailed; he only stated that the body is rounded, blood-red, both ends blunt, and that the posterior part is more slender in the females. 341 Later authors therefore recorded red nematodes found in varous freshwater fishes as Philometra sanguinea, although they were localized in the body cavity. 331 = у D “ii —_ —А,—А, a тниныя а FIGURE 197. Philometroides sanguinea (Rudolphi, 1819): a — anterior end; b — posterior end of female, rudiment of vagina visible; с — transverse section at the rudimentary vagina; d — genital pore of young female, lateral; e — posterior end of male; f — anterior part of a young form (a—c — after Mirza, 1929; d—f — after Wierzbicki, 1960). Linstow (1902, 1909) gavea long list of the hosts of this species: Caras - sius vulgaris, Leuciscus rutilus, Abramis Бгаюа. ВЫ Vimba, Blicca bjorkna, Osmerus eperlanus, Апр vulgaris, Cottus gobio; as localizations he gave the body cavity, but for Abramis vimba under the skin of the operculum, Linstow gave a more detailed and specific description of Ph. sanguinea: 342 coloration red, esophagus very short, cephalic end with 4 papillae. Length of male 2.3mm, width 0.036 mm, Length of female 40 mm, width 1 mm, Length of spicules 0.24 mm: Length of larvae in uterus 0.5mm, width. 0.026 mm. In an examination of specimens of Carassius, Nybelin (1928) found between the rays of the caudal fin females 40 mm long which he determined as Philometra sanguinea. He stated that the cuticle is covered with 332 a SS er ee numerous papillae. Ш 1931, after examination of the original material of Rudolphi in the Berlin Museum, he found thateven badly preserved specimens show the cuticular papillae characteristic for the species on the whole body and stated that Ph. sanguinea isa specific parasite of Carassius. Nybelin established that Rudolphi's description applies only to parasites localized in the fins, and he proposed naming nematodes from the body cavity of species of Cyprinidae Philometra abdominalis Nybelin, 1928. In a study of the helminth fauna of freshwater fishes in Sweden in 1928, Nybelin found in the pectoral finsof Abramis farenus and under the skin of the opercula in A. blicca nematodes which he did not identify as Ph. sanguinea because the cuticle was not quite smooth and the posterior end, unlike in Philometra ovata (Zeder), was bifid, not rounded, andthe cephalic end bore 4 well-developed papillae. In 1931, convinced that these nematodes are distinct from Ph. sanguinea and Ph. ovata, Nybelin named them Philometra opercularis Nybelin,1931. He also stated that four species of Philometra occur in European freshwater fishes and gave the following key for their determination. A. Cuticle with numerous papillae. Localization in the caudal fin, Evelyn bie=dorsal LinnOre’ air ersiSiinuls ee wees ED ee ee, OR oe Bea A eS eS a pec a han Ee es CMa RN CR -Philometra sanguinea Rudolphi, 1819. Ee «Cuticle smooth. Posterior end bifid. Localization under the skin in the anterior part ОЕ оу, ресъога fins" Opercullay TOUS А. с Иа. Philometra opercularis Nybelin, 1931. 2. Posterior end rounded. Parasites of the body cavity aecnotheabout 40—60) mim-s Coloration red: iat. i 5 ol ie sete eae ира... - Philometra abdominalis Nybelin, 1928 (= sanguinea auct. ). DP) wiltenethatori2 so! nm-ryColoration yellowish white”). oc. Suess yea То А EIS | ЗЕ, Le at ek Philometra ovata (Zeder). — A high rate of infection of Carassius with Philometra sanguinea was observed by Wierzbicki (1960) in Poland, by Cakay (1957) in Czechoslo- vakia, Molnar (1966b) in Hungary, and by Vismanis and Nikulina (1968) in the USSR; all these authors mention the strict specificity of the parasite for Carassius, since in lakes where Carassius were found infected other species of fish were free of infection. Vismanis and Nikulina (1968) examined the original material of итога sSianeulnea’ from Carassius auratus) in ‘ponds in the Shipun District of the Altai Territory and from Poland, compared the GesenipronsioL Phy vsancuiniea,. Ph. trilabiata’ Bellouss; 1965, and Ph. carassii Ishii, 1933, and came to the conclusion that all these Species are identical and made Philometra carassii Ishi, 1933 and 343 Philometra trilabiata Belouss, 1965 synonyms of Philometra Sanguinea Rudolphi,1819. We agree and, in accordance with Rasheed (1963), transfer this species to the genus Philometroides, Yamaguti, EOS Description (after Skrjabin, 1923a,as Philometra sanguinea). Male. Length 2.3mm, width 0.036 mm. Tail ending in two lobes; spicules 0.24 and 0.197 mm long; end of spicules pointed. 333 Female. Coloration blood-red. Length 40mm, width1.0mm. Length of esophagus 1.56 mm. Intestine ending blind near the posterior end, Larvae in the uterus 0.5mm long and 0.26 mm wide. | Description (after Dogel' and АКБтегоу, 1959, аа Philometra sanguinea). Male not described. ~ Female. Length 10—35 mm. Cephalic end with a hemispherical process, at the apex of which the mouth is situated. The process has a strong musculature and around its base is a muscular collar which is apparently formed by four modified cephalic tubercles which have become fused. The cephalic part narrows slightly posteriorly, forming a short neck, and then begin small papillae, more than 150, which are irregularly distributed but show a tendency to form two lateral rows, particularly toward the ends of the body. РарШае pedunculate, partly embedded. The body is widest behind the head and tapers posteriorly. Length of : esophagus 1.33 mm. In young forms the sides of the posterior end bear a paired process which resembles a caudal fin with a crescent-shaped notch between the two halves. Wierzbicki (1960) described young forms of Ph. sanguinea. Young females are very small during the copulation period. Unlike in the adults, the cuticle is smooth in young males and females. There is a well-developed intestine, which later atrophies in females: the intestine becomes narrower and the anus closed completely. ; Male. Length 3.1—3.3mm. Posterior end with two spicules and a gubernaculum., Е Female. Length2.1—2.7mm. At the posterior third of the body is a single tubercle, where the genital pore is situated; this atrophies after copulation. Wierzbicki (1960) established that the life cycle of Ph. sanguinea lasts one year in nature. He found females in Carassius throughout the year but, according to the season, they were at different stages of develop- ment. The youngest females were found in the body cavity of the fish in the summer from May to October. -Fertilized females leave the body cavity. They do not appear in the fins until September; their number in the fins increases in autumn and winter. Almost all the parasites leave the fins in June. Larvae which had left the uterus remained alive in water for 9 days and remained infective for 5 days. They were 0.37—0.40 mm long. They 344 develop further in the body cavity of the intermediate host (Cyclops kolensis Lill.), After entering the water and then the intermediate and the definitive host, the larvae grow to 0.41— 0.42 mm after 5 days, 0.44—0.46 mm after 10 days, 0.53—0.57 mm after 20 days, 0.65 mm after 33 days, and 1.02 mm after two months. The proportions of the body change with growth. The tail becomes shorter: it is 0.070—0.075 mm long when the larva enters the water, after 4 days it is 0.066—0.071 mm long, after 6 days 0.063—0.067 mm, and after a month in the body cavity of Cyclops the tail disappears. The width of the larva, on the other hand, increases withage. Onthe first day whenthey leave the female, the larvae are 0.001—0.013 mm wide, after 5 days 0.013—0.016 mm, and after 10 days 0.015—0.017 mm; the larvae of Ph. sanguinea are infective but they lose their infectivity after 14 days. | 334 After the infective larvae enter the intestine of the definitive host, they penetrate its wall and move into the body cavity. References: Belous, 1965, рр. 48—65; Dogel' and Akhmerov, 1959, pp. 287—304; Skrjabin, 1923a, pp.1—98; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, pp.1—927; Vismanis and Nikulina, 1968, pp. 514—519; Cakay, M957, юр, 909—912: Linstow,) 1902, вр. 210—232: 1909: рр. 47—83; Molnar, 1966, рр. 227—241; Nybelin, 1928, рр. 1—4; 1931, рр. 58— 64; Rasheed, 1963, pp. 89—130; Rudolphi, 1819, pp. 1—811; Wierzbicki, 1960, eno On Vi, 1956, pp. 99-196. Genus Pseudophilometroides Parukhin, 1966 Diagnosis. Philometrinae. Body cylindrical, long, with projecting anterior and rounded posterior end. Cuticle with numerous thickenings. Head with 8 papillae, tail without papillae. Mouth simple. Esophagus without an anterior swelling, with a well-developed, cylindrical dorsal gland. Anus and vulva atrophied. Viviparous. Parasites of marine fish. Туюе SPeCleSss вс ее kom et rowdes ato pl, Раба: 1966. Pseudophilometroides atropi Parukhin, 1966 (Figure 198) Eos VA tO, puis, ait Opus - (345) FIGURE 198. Pseudophilometroides atropi Parukhin, 1966: а — anterior end of female; b — posterior end of female; с — larva; d — esophagus and esophageal gland; e — cephalic end, apical (after Parukhin, 1966). 335 Localization: body cavity. Distribution: Gulf of Tonkin. Description (after Parukhin, 1966). Male unknown. Female. Length 19.5—36.0 mm, width 0.645—1.125 mm. The entire body, except the anterior part, is densely covered with cuticular thickenings in the form of papillae. Cephalic region markedly projecting. Length of projecting part 0.20 mm, width 0.20 mm. Mouth simple. Cephalic end with 8 papillae and 2 amphids. Esophagus without an anterior swelling, 1.2—1.65 mm long, 0.062— 0.065 mm wide. There is an esophageal valve 0.093 mm long and 0.087 mm wide. On the dorsal side of the esophagus is a large cylindrical digestive gland 0.765 mm long and 0.18 mm wide; the gland begins in a slight inden- 345tation of the anterior margin of the esophagus and extends to the esophageal 346 valve. Nerve ring situated 0.177 mm from the cephalic end. Uterus filled with larvae 0.495 mm long and 0.012 mm wide. Posterior end rounded, Anus and vulva atrophied. References: Parukhin, 1966, pp. 766—767; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, pp. 72—79; Rasheed, 1963, pp. 89— 130. Genus Rumai Travassos, 1960 Diagnosis. Philometrinae. Body cylindrical, with tapering ends. Head with apical processes with a concave surface between them and with 4 papillae. Cuticle smooth. Esophagus club-shaped, not swollen anteriorly. Tail conical. Males unknown. Parasites of freshwater fish. Type species: Rumai rumai Travassos, 1960. Rumai rumai Travassos, 1960 (Figure 199) Host: Arapaima gigas. Localization: body cawity: Distribution: Amazon’ River. Description (after Travassos, 1960). Male unknown. Female. Length 40 mm, width 0.64 mm. Cephalic end with a cuticular formation which appears like a pyramid in profile, while it is slightly concave in apical view. Four papillae are situated on this formation. Average width of cephalic formation 0.16 mm. Length in lateral part 0.087 mm, at the mouth 0.075 mm. Cuticle smooth. Mouth leading directly into the esophagus. Esophagus large, club-shaped, its posterior end wider, with a wide lumen. Its anterior part is almost cylindrical; nerve ring situated near the mouth. Length of esophagus 2.51 mm, width in the posterior part 0.36 mm, in the anterior part, at the nerve ring,0.144 mm. Nerve ring situated 0.315 mm from the cephalic end. Intestine apparently atrophied. Anus absent. Genitalia amphidelphic. Anterior ovary situated near the esophagus, opposite the uterus. Posterior ovary extending anteriorly from 336 the posterior end and partly concealed by the uterus. Uterus wide and , saclike, occupying almost the entire body cavity. It is filled with first— stage larvae. Posterior end of body conical. 0/mm d FIGURE 199. Rumai rumai Travassos, 1960: a — anterior end, lateral; b,c — cephalic end, different aspects; d — posterior end (after Travassos, 1960). Reference: Travassos, 1960, pp. 15—20. Genus Thwaitia Rasheed, 1963 Historical review Thwaite (1927) described a nematode from the eye of Balistes sp. from Ceylonas 'Philometra взр." Although he had only a few fragments of the cephalic and the posterior end, they permitted the description of the most important characters. Rasheed was interested in this material be- cause of the strange form of the cephalic papillae. The material was placed at her disposal by Prof. Cashew of the University of Liverpool, where the type of the species described by Thwaite was deposited. It proved identical with the specimen in the British Museum. After a detailed study of the material, Rasheed came to the conclusion that these nematodes belong to a new genus for which 4 cephalic papillae are specific, which are present only in a few genera and species of Philometridae. Linton (1901) described a nematode with 4 papillae as Ichthyonema sanguineum. Four cephalic papillae are also present in Philometra abdominalis Nybelin, 1928 (=Ichthyonema 337 347sanguinea 1л1пзфо\, 1874), Ph. rischta Skrjabin, 1917 (|B; opercularis Nybelin,1931), Ph. intestinalis Dogiel and Bychowsky, 1934, Ph. translucida Walton,1927, and Ph. americana Kuitunen- Ekbaum, 1933 in which the papillae are weakly developed. Rasheed placed all these forms in the new genus Thwaitia Rasheed, 1963. Molnar (1966b) thinks that Ph. intestinalis belongs to the family Skrjabillanidae Schigin and Schigina, 1958, although he accepts the classification of the family Philometridae proposed by Rasheed (Molnar, 1969). Moravec (1968) established the new genus Molnaria Moravec, 1968 of the family Skrjabillanidae in which he placed M. intestinalis (Dogiel and Bychowsky, 1934) Moravec, 1968 (=РЬ. intestinalis Dogiel and Bychowsky, 1934). Wealso place М. intestinalis in the genus Molnaria Moravec, 1968. Diagnosis. Philometrinae. Head with 4 large papillae in the outer ring. Inner ring indistinct or absent. Esophagus widened anteriorly. Ventricle present or absent. Tail blunt or rounded, with or without papillae. Males unknown. Parasites of freshwater and marine fish. Type species: Thwaitia balistii Rasheed, 1963. Thwaitia balistii Rasheed, 1963 (Figure 200) Synanym::Philemetra spe Thwatte; "£927 Host: Balistes sp. hocalization: eyes: Dicie i pilt tons” Geylon, Description (after Rasheed, 1963). Male unknown. Female. Length to 210 mm, width 0.54—0.6 mm. фито FIGURE 200. Thwaitia balistii Rasheed, 1963: a — larva; b — cephalic end of female, lateral; с — same, dorsoventral; 4 — anterior end of female, esophagus, ovary, and uterus visible; e — posterior end of female, dorsoventral; f — same, lateral (after Rasheed, 1963). 338 348 Cephalic end blunt, appearing square in dorsoventral view; it bears large papillae which project laterally; width 0.18—0.19 mm. There are 4 flat lobes, each of which bears two fleshy papillae with indistinct nerve endings. These double papillae are situated on the submedian line; there is also а pair of small lateral amphids. The anterior end of the esophagus is widened, narrowing to the nerve ring, and then of almost uniform widthits entirelength. Esophageal gland less distinct than in Philometra. Nerve ring surrounding esophagus about 0.14—0.15 mm from the cephalic end. Esophagus muscular its entire length, which is 2.04—2.34 mm; its width near the mouth is 0.09— 0.17 mm. An esophageal ventricle or a process are absent. Intestine slightly wider than the posterior end of the esophagus, ending blind near the posterior end. Anus absent. Tail rounded, with two wide lobes projecting laterally, each lobe with two papillae. Two ovaries, one situated in the anterior, the other in the posterior part of the body. Their position is not constant and probably depends on the movements, state of preservation, or age of the worm. Uterus distended with larvae, which are very large: length 0.36—0.4 mm, width 0.017—0.019 mm. Their cephalic end is rounded and apparently bears two papillae. Cuticle with fine transverse striation; mouth opening into a wide intestine. Anus situated about 0.084 mm from the end of the pointed tail. References: Shigin and Shigina, 1958, pp.395—399; Linton, 1901, pp. 441—481; Molnar, 1966, p.157; 1969, p.137; Moravec, 1968, р. 322; Rasheed, 1963, рр. 89—130; Thwaite, 1927, pp. 225—242. Thwaitia abdominalis (Nybelin, 1928) Rasheed, 1963 (Figure 201) Synonym: Philometra abdominalis Nybelin, 1928 Moss iG ONO Зоо ВОТ Ее, —eucisceus aiduse Localization: body cavity. Distribution: USSR, Western Europe. Historical review The name Philometra abdominalis was given by Nybelin (1931) to separate mixed material which was placed under a single label together with Philometra sanguinea and had been identified as this species by Kreplin (1825), Diesing (1861), Linstow (1874), and other authors who had found these forms in various Cyprinidae. Nybelin did not give distinct differences between Ph. abdominalis and Ph. ovata, distinguishing them only according to color and size. Molnar (1967) studied typical specimens of Ph. abdominalis from Gobio gobio, and found that the males of these species are clearly distinguished by the form and size of the spicules and gubernaculum. ae Rasheed (1963) transferred the species Philometra abdominalis to the genus Thwaitia. Description (after Molnar,1967,as Philometra abdominalis ). 339 Male. Length 2.1—3.7 mm, width 0.037—0.057 mm. Cuticle smooth, 349 cephalic end rounded, with 4 oblong papillae. Body of uniform width its entire length, except for a slightly narrower cephalic end. Posterior end blunt, with two lateral teatlike processes which are indistinctly connected dorsally. р Two spicules of almost the same length, moved by the gubernaculum. One spicule 0.127—0.172 mm long, the other 0.120—0.164 mm. Width of spicules 0.007 mm at the slightly thickened gubernaculum, 0.005 mm at the beginning, and 0.003 mm in the middle. Gubernaculum 0.53—0.65 mm long in the form of a curved chitinized plate with lanceolate end. Thereis a dorsal invagination in the form of a fringe in its distal part (however, this invagination is not as prominent as in Ph. ovata). Immediately after this invagination the gubernaculum is curved sharply dorsally and an- teriorly. FIGURE 201. Thwaitia abdominalis (Nybelin, 1928): a — cephalic end of female 3 months old; b — posterior end of female 3 months old; c — posterior end of male (after Molnar, 1967). Female. Length 100—120 mm, width 0.9—1 mm. Adults red or yellow- ish red. Body cavity filled by the uterus. Cuticle smooth, Cephalic end rounded, with 4 slightly protruding papillae and 3 lips. Body cylindrical, of the same width its entire length. Caudal end rounded; two indistinct cuticular processes are visible on it in dorsoventral view. These pro- cesses are more distinct in young females 20—40 mm long. The life cycle is completed in a year in the temperate zone, and the size of females in different seasons can therefore be determined with accuracy. According to Molnar, obligatory hosts are only Gobio gobio and Phoxinus phoxinus, in which all females attain maturity. Life cycle. Thw. abdominalis was studied by seasonal dis- sections of Ph. phoxinus andG. gobio. The fish become infected when they swallow Cyclops containing larvae of the nematode in June 350and July. Juvenile parasites, with a pointed tail like the larvae, were found at this time in the abdominal cavity of the hosts. During the summer, 340 developing nematodes were found under the mucosa of the swim bladder, where copulation takes place. Developing females usually remained in the abdominal cavity in August and September. Males and unfertilized females were found during the whole year under the mucosa of the swim bladder; they do not grow. Females which passed into the abdominal cavity were 22—66 mm long in autumn and 60—72 mm in the following spring. The first larvae appear at the beginning of May, and the uterus of females is filled with fully de- veloped larvae 77—110 mm long in the middle of May. At the end of June the females leave the host actively through the anus. They break up in the water; the larvae are released and survive for 3—5 days, attracting the at- tention of Cyclops by their active movements. Cyclops is the inter- mediate host, in which the larvae develop to the infective stage. Cyclops collected where fish were caught for study became infected with larvae of Thw. abdominalis under laboratory conditions. Mature females could sometimes not leave the abdominal cavity of the host and died in it. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, p.73; Kreplin, 1825, p.95; Diesing, 1861, p.42; Linstow, 1874, рр. 122—134; Molnar, 1967, pp.293—300; 1969, рр. 137—143; Nybelin 1928, pp.1—4; 1931, pp. 58—64; Rasheed, 1963, pp. 89—130. Thwaitia bagri Khalil, 1965 (Figure 202) Host: Bagrus bayad. Localization: under the skin lateral to the mouth. Distribution: Sudan (Nile). Description (after Khalil, 1965). Male unknown. Female. Length of mature female 20—31 mm. Body of almost uniform width its entire length, width 0.48— 0.68 mm, slightly narrowing at the ends. Cuticle smooth and relatively thick. Cephalic end rounded, with 4 large, thick papillae situated on the submedian line; there is also a pair of small amphids which are difficult to see. Width of cephalic end 0.11—0.17 mm. Mouth simple, surrounded by 3 elevations which form a triangular cone. Mouth opening directly into a short muscular esophagus which is 1.48— 1.65 mm long and 0.05—0.08 mm wide. The esophagus is widened anteriorly and then gradually narrows and becomes threadlike. The large esophageal gland opens into the esophagus before the nerve ring and extends along the whole esophagus laterally, ending blind atthe end of the esophagus. Nerve ring situated 0.2—0.26 mm from the cephalic end. Intestine relatively thick-walled, wider than the esophagus; its lumen gradually narrows and disappears at the end. The intestine adheres to the wall of the posterior end of the body. Anus absent. The long, cylindrical, reverted ovaries 351аге situated at the ends of the body and vary markedly in position. The large uterus has thin walls and extends through the entire body. It is filled with larvae. Vagina and vulva absent. Tail blunt, square, with two large, conical lobes, 0.34—0.39 mm wide. ‘Larvae 0.24—0.31 mm long, 0.016—0.019 mm wide, with blunt cephalic and pointed posterior end. Cuticle with fine striation. Esophagus 0.099— 0.143 mm long; intestine containing large granules. Anus situated 0.06— 0.066 mm from the posterior end. 341 352 М рб м а FIGURE 202. Thwaitia bagri Khalil, 1965: a — anterior end of female, lateral; b — posterior end of female, lateral; с — tail of female, ventral; 4 — cephalic end, apical; e — larva from uterus (after Khalil, 1965). Because of their bright red coloration, the worms are visible under the skin of the fish. When placed in physiological saline, they straighten and some of them live for 2 days at room temperature and slightly longer in the refrigerator. When adult worms enter the water, they straighten and burst in the region of the esophagus, releasing the larvae into the water. After 2—3 hours, all the larvae are discharged and swim free in the water. They survive for about 3 days in tapwater at room temperature. The female then soon dies. Most of the females found under the skin of the fish lateral to the mouth were mature, and only a few did not contain larvae. Reference: Khalil, 1965, pp. 309—312. Thwaitia kotlani Molnar, 1969 (Figure 203) Host: Aspius aspius- Localization: body cavity. Distribution: Hungary. Description (after Molnar, 1969). Male. Length 2.1—2.6mm, width 0.041—0.062 mm. Cuticle transparent; body tapering slightly at both ends. Cephalic end rounded, without papillae. Caudal end blunt, with two teatlike processes which are connected by a membrane. Two narrow, short spicules of almost the same length, 0.090— 0.10 and 0.082—0.098 mm. Gubernaculum forming a curved chitinized plate with ring-shaped end; it is curved dorsally in about the middle and then dorso-anteriorly; it is 0.053—0.066 mm long. Female. Length 61 mm, width 1mm. Red nematodes. Body cylindrical, tapering slightly at bothends. Cephalic end rounded, with 4 indistinct papillae and 3 lips. End of tail rounded; two small teatlike processes on both sides in dorsoventral view. Body cavity entirely occupied by the uterus, which is filled with larvae. Length of larvae 0.44—0.46 mm; cephalic end rounded, posterior end tapering. 342 2.5 LS SOLOS Perry rr Ke, HEE : Rate FIGURE 203. Thwaitia kotlani Molnar, 1969: a — anterior end of female, lateral; b — posterior end of female, lateral; с — posterior end of male (after Molnar, 1969). Immature females 20—30 mm long have teatlike processes at the posterior end. Molnar studied the life cycle and gives the following details. Thw. kotlani has a one-year cycle. Mature females filled with larvae were found only in May. The discharged larvae enter the water and continue to developin Cyclops. Aspius aspius became infected in summer, probably when they swallowed infected Cyclops or smaller plankton- feeding fish. After copulation, the males of Тим. kotlani remain in ' the swim bladder throughout the year, and fertilized females settle in the abdominal cavity. It is not known when the females move from the swim bladder into the abdominal cavity, but probably in early autumn, since females 15—20 mm long were found in the body cavity in early spring. The uterus of such females was filled with eggs in the early stages of development. At the beginning of April the females were 25—30 mm long and the eggs were already ripe. Older larvae hatch in May. Reference: Molnar, 1969, pp. 137—143. 353 Тома Ша rischta (Skrjabin, 1917 ){ Figure 204) Ppa oOmMyt Роме са s,riceht a. Skrjabiny 1917 Hosts: Pseudaspits leptocephalus, Aspius aspius. Localization: under the skin of the head. Distribution; Europe, Asia. 343 354 FIGURE 204. Thwaitia rischta (Skrjabin, 1917): a — anterior end of female; b — posterior end of female; c — posterior end of male; d — fragment of female; e — anterior end, lateral; f — cephalic end, apical; g — larva (a—d — after Molnar, 1966b; e—g — after Lewaschoff, 1929). Description (after Skrjabin,1923b,as Philometra rischta). Male not described. Female. Length17 to20 mm. Measurements of a female 19 mm long and 0.6 mm wide in the middle of the body. Cephalic end 0.1mm wide, posterior end 0.25 mm wide. Width at level of posterior part of esophagus 0.39 mm. Cephalic end slightly blunt, with 4 distinct, hemispherical, convex tubercles arranged symmetrically on the lateral lines and on the dorsal and ventral line. In the middle of the rectangular field formed by these tuber- cles is the trihedral mouth, which leads into the strongly muscular eso- phagus. The first part of the esophagus is slightly swollen and separated from the other part by a constriction; however, the esophagus does not form a funnel-shaped invagination anteriorly. Length of esophagus 1.345 mm, width 0.12 mm in the posterior part. On the dorsal side, in the wall of the esophagus, there is a large gland with a large nucleus, In- testine extending almost to the end of the body, ending blind. The posterior end has a distinctive structure; it consists of two lateral teatlike processes directed obliquely laterally, situated 0.25 mm from each other. Between the lateral processes the posterior margin forms a more or less deep crescent-shaped invagination. 344 The anterior end of the large uterus is situated 0.255 mm from the cephalic end, the posterior end 0.73 mm from the posterior end. At the posterior end of the uterus begins the small, slightly twisted ovary, which extends posteriorly inthe form ofathin, cylindrical appendage. The uterus is filled with developed larvae which are 0.357 mm long, 0.017 mm wide at their thickened cephalic end. Description (after Lewaschoff,1929,as Philometra rischta). Cuticle thin, smooth. Cephalic end with 4 large conical processes situated submedially. The anterior end is slightly laterally flattened so that these processes are arranged in pairs slightly more close together on the dorsal and ventral side. Mouth forming a triangular slit. Beginning of esophagus swollen like a club, strongly muscular. This swelling passes abruptly into the other part of the esophagus, which is narrower and long 0.05 mm from the anterior end. Male not described. Female. Length 20.66—31.11mm. Width 0.49—0.70 mm, but varying in different parts of the body: 0.12—0.13 mm at the end of the "head, "' 0.45—0.61 mm at the posterior end of the esophagus, and 0.18—0.29 mm at the posterior end. Length of esophagus 2.16—2.45 mm, width in the posterior part 0.01—0.07 mm. Vulva and anus not found. Uterus with thin walls and so distended with larvae that the intestine is pressed against the body wall. The larvae are situated 0.74 mm from the cephalic end and 0.13 mm from the caudal end. Length of larvae 0.369 mm, width at the cephalic end 0.013 mm. Description (after Molnar, 196ба). Male. Length 1.68—2.00 mm, width 0.022—0.039 mm. Cuticle trans- parent, smooth. Cephalic end rounded, with 4 circumoral papillae. It narrows toward a slightly constricted 'neck.'' Behind the neck the body is almost of the same width as the cephalic end. Posterior end with lateral teatlike processes which are connected dorsally by a narrow lobe. 355 Spicules of almost the same length, 0.045—0.057 and 0.045—0.053 mm. Width of spicules at the end 0.003 mm. Gubernaculum 0.031—0.041 mm long, forming a curved chitinized plate. Female. Mature females yellowish red, length 30—32 mm, width 0.8—1.0 mm. Body tapering slightly at both ends. Cuticle smooth. Cephalic end with 4 distinct papillae and 3 lips. Posterior end with two distinct, lateral, teatlike processes. Uterus filled with larvae, occupying almost the entire body cavity, pressing the esophagus and intestine to the body wall. Anus subterminal. Length of young females 5—7 mm. Coloration yellowish red or red. Cuticle smooth. Body of almost uniform width its entire length, tapering slightly at the ends. Posterior end with two teatlike processes. Cephalic end with 4 distinct papillae and 3 lips. Esophagus cylindrical, forming a bulb anteriorly. Its posterior end is connected with the wide intestine by valves. The esophagus is serrated from the bulb to the valves; it contains an esophageal gland which becomes gradually thicker, being twice as wide as the esophagus in the middle, and then gradually narrows again, ending in the esophageal valves. Intestine of uniform width its entire length, narrowing posteriorly and ending in the subterminal anus. Nerve ring situated behind the esophageal bulb. Vulva rounded, situated in the posterior third of the body; vagina chitinized, 0.004 mm wide, extending anteriorly for 0.019—0.021 mm. Uterus forming a sac, filled with eggs, 345 356 narrowing at both ends in young specimens; in older specimens it passes into the tubular ovaries. The anterior ovary begins from the uterus near the esophageal valves and makes several loops at the level of the esophagus, turns and ends at the anterior end of the uterus. The posterior ovary forms loops near the rectum and turns, ending around the uterus. The life cycle was studied by Molnar (1966a), who found that Thw. rischta has an annual life cycle. Adult females of Thw. rischta 35—42 mm long were found at the end of May and in early June on the inner surface of the operculum, sometimes under the skin of the head, in the orbit, rarely at the base of the anterior fin. One to 35 specimens were usually found in a host. Males were found only in summer. There were usually 2 specimens under the serosa of the posterior part of the swim bladder; one male was found on the caudal fin. A single male was found in the swim bladder of Aspius aspius in May; females were not found in this specimen. Fish 2 years old and more become infected with larvae of Thw. rischta atthe end of June. Males 1.53—1.92 mm long and 0.024—0.031 mm wide and females 1.04—1.057 mm long and 0.031—0.037 mm wide were first found in the swim bladder in early July. In the middle of July only males 1.78—2.73 mm long and 0.029—0.037 mm wide were found in the swim bladder of several spe- cimens of Alburnus. In the middle of August females and males disappeared from the swim bladder, and several juvenile females 1.30—2.73 mm long and 0.033— 0.105 mm wide were found on the inside of the operculum and in other parts of the head. In heavily infected fish (over 25 specimens) immature females 3.56—6.57 mm long and 0.114—0.217 mm wide were found in September in the muscles near the pectoral fin. In autumn the females continue to de- velop and become 5.6—6.8 mm long and 0.20—0.26 mm wide at the end of October. In early April, the females are 8.21—22.50 mm long and 0.26— 0.47 mm wide and contain nonsegmented eggs 0.014—0.019 mm wide. Segmentation of the eggs continues in early May and reaches the morula stage; the slightly oval eggs are 0.038—0.042 mm wide. From 10 to 20 May an elongate developing embryo is visible, and at the end of May the uterus is filled with mobile first-stage larvae. Mature females 35—42 mm long leave the host at the end of May or in early June. Fish examined after 10 June showed only tissue lesions caused by Thw. rischta. When they enter the water,females of Thw. rischta burst, and the first-stage larvae are released. They are 0.280—0.430 mm long at this time. Moln&r stated that Cyclops strenuus, Macro- cyclops albidus, and Acanthocyclops viridis are very sus- ceptible to infection with this parasite; however, 100% infection may take place only in young population of crustaceans. Only a third of adults become infected. References: Skrjabin, 1923b, pp.1—9; Skrjabin, Shikhobalov, Sobolev, Paramonov, and Sudarikov, 1954, p.78; Lewaschoff, 1929, рр. 121—128; Molnar 1966a, рр.227—241; 1966b, рр. 147—149. 346 Тома! На sp. (Linton, 1910) Rasheed, 1963 Synonym: Ichthyonema sanguinea Linton, 1901 Host: fish. Localization: body cavity. Distribution: Europe. Description (after Linton, 1910). Male unknown. Female. Length 30 mm, width 0.81—1.0mm; width of cephalic end 0.23 mm. Adults long and narrow, slightly truncate at the cephalic end. Four lips, each with two papillae. Esophagus at first slender, gradually widening; length 1.14 mm, width 0.1 mm at the beginning and 0.17 mm at the end. Intestine at first slightly wider than esophagus. Uterus large, occupying the greater part of the body cavity and filled with embryos. References: Linton,1901, pp.441—481; Rasheed, 1963, pp. 89—130. Thwaitia translucida (Walton, 1927) Rasheed, 1963 (Figure 205) Synonym: Philometra translucida Walton, 1927 ось: sox lucius:. Localization: abdominal cavity. 357 Distribution: North America. Description (after Walton, 1927, аа Philometra translucida). Male unknown. FIGURE 205. Thwaitia translucida (Walton, 1927). Anterior end (after Walton, 1927). Female. Length 75 mm, width 0.75 mm. Cephalic and caudal end truncate. Body whitish, transparent, delicate but resilient. Lateral lines wide and transparent. Mouth surrounded by 4 small but distinct papillae, 347 esophagus with а distinct bulb anteriorly. Length of esophagus 4 mm, width 0.18 mm. Intestine oval,0.3mm wide. Esophageal gland small and opaque. The opposite uteri extend from the connection between the eso- phagus and the intestine for 6/7 of the length of the body. The two ovaries consist of short, winding tubes which extend from each end of the uterus. Vulva and vagina not recognizable. Eggs spherical, 0.040—0.050 mm wide. Reference: Walton, 1927, р. 89. SUBFAMILY PHILONEMINAE N. SUBFAM. Historical review Chitwood (1937) placed the genus Philonema in the subfamily Philometrinae, but stated that the position of the genus in this subfamily is doubtful, since the esophageal glands are polynuclear but the larvae are of the dracunculoid type. Chitwood and Chitwood (1950) also stated that Philonema shows an extreme development of the esophageal gland in the Dracunculoidea; the species of this genus have a typical spiruroid- filarioid esophagus. Skrjabin et al. (1954) provisionally kept the genus Philonema inthe subfamily Philometrinae. A study of the life cycle of Philonema oncorhynchi (Platzer, 1964; Platzer and James, 1967; Adams, 1969) showed that development in the definitive host may last as long as 3 years. Because of the above morphological and biological characteristics of the species of Philonema, we consider it justified to establish the sub- family Philoneminae n. subfam. for this genus. Diagnosis. Philometridae. Esophagus divided ито а narrow muscular anterior part and a widened glandular posterior part. Esophageal gland polynuclear. Posterior end of males long and thin, tapering. Type genus: Philonema Kuitunen-Ekbaum, 1933. 358 Genus Philonema Kuitunen-Ekbaum, 1933 Synonym: Coregonema Bauer, 1946 Historical review The genus was established when Kuitunen-Ekbaum described a new nematode from Salmonidae, Philonema oncorhynchi. Simon and Simon (1936) and Fujita (1939, 1940) described six further species of the genus. After a study of specimens of Philonema from fish near Kamchatka and comparison with the descriptions of species of the genus Philonema and Coregonema Bauer, 1946, Akhmerov (1955) identified his speci- mens with Philonema oncorhynchi Kuitunen-Ekbaum, 1933 and questioned the validity of the species of the genus Philonema. He 348 considered Philonema agubernaculum Simon and Simon, 1936 and Ph. elongata Fujita, 1940 as synonyms of Ph. oncorhynchi and assumed (he did not know Fujita's work of 1939) that the specific characters of Philonema kondai, Ph. salvelini, Ph. tenui- cauda, and Ph. ochotense (Fujita, 1939) are not convincing and that- they are probably also synonyms of Ph. oncorhynchi. Не also stated that Ph. oncorhynchi and Coregonema sibirica Bauer, 1946 are indistinguishable, but their hosts and ecology are completely different: the hosts of Ph. oncorhynchi are Salmonidae, the hosts of C. sibirica Coregonidae; the Salmonidae are anadromous fishes and their adults live in the sea, while the Coregonidae live in fresh water. These ecological characters also refer to the nematodes parasitizing them: Ph. oncorhyn- ehi is a marine species, С. Sibirica afreshwater species. The dif- ferences in size of these species do not exceed the boundaries of genus and species, and therefore Akhmerov made С. sibirica Bauer, 1946 а Synonym or a subspecies of Ph. oncorhynchi and the genus Core- gonema .a synonym of the genus Philonema. In a study of the migrations of Salmonidae in the Far East, Mamaev et al. (1959) established that Ph. oncorhynchi is a freshwater form, and they therefore proposed that the species and the subspecies Ph. oncorhynchi sibirica should be considered as a freshwater species. Rumyantsev (1965), who first recorded Ph. sibirica from the European USSR (Karelia), also considered Coregonema as a synonym of Philonema, but he thinks that sibirica is a valid species, quite ее ео скота Pho oncorhynchi.’ As to. Ph. elongata, Hujita, 1940, Rumyantsev agreed with Akhmerov (1955) that it is a synonym of Ph. oncorhynchi and suggested, quoting Tamao (1960) and Meyer (1960), Mitte hbekondal. Phe, salvelini,and Ph. tenuilcauda,described by Fujita (1939), are also synonyms of Ph. oncorhynchi. Rasheed (1963) placed 6 species in the genus Philonema: Ph. Бонни hing Ро: ap iiberimaculum. РЯ. elongata.) РВ. Во. маме тии, and) ihe tem wile a uid а. After reading the descriptions of Fujita (1939, 1940), we agree with Akhmerov and Rumyantsev and consider the species described by him as synonyms of Ph. oncorhynchi. Bashirullah (1967) published a short communication on the development of the species of the genus Philonema in Salmonidae with different life cycles. The study was made to establish the identity of Philonema oncorhynchi from the anadromous Omecormhynehws песка and Ph} asubernaculum: from freshwater 359 trout, and because of the differences in the life cycles, cross infections, and the protein composition of the parasites, he considered Ph. oncorhyn- сари. agubernaculuim аз different Species.) He also trans planted adult Ph. oncorhynchi (but without larvae) from almost mature sockeye salmon into immature trout. The larvae did not develop in trout. We therefore recognize 3 species inthe genus Philonema: Ph. ое умен. Ри: “agubernaculum, and Ph. sibirica. Diagnosis. Philoneminae. Females markedly larger than males; body threadlike; anterior end rounded, posterior end elongate; mouth with lips or papillae or without them; esophagus cylindrical, esophageal gland polynuclear, 349 Male. Posterior end elongate and spirally twisted; anus subterminal; spicules of the same length; gubernaculum absent. Female. Anus and vulva atrophied in adults; body cavity occupied by the uterus, which contains developed larvae. Two short ovaries situated at the ends of the body. - Parasites of the body cavity and, possibly, of the connective tissue of fish. Type species: Philonema oncorhynchi Kuitunen-Ekbaum, 1933. Philonema oncorhynchi Kuitunen-Ekbaum, 1933 (Figure 206) Synonyms: Philonema kondai Fujita, 1939; Ph. ochotense Виа, 2939: Ph. salvelini, Fujita, 19399 Ph. tenuicauda Fujita, 1939; Ph. elongata Fujita, 1939 Hosts: Oncorhynchus nerka,. О.“ kheta, -SJalvelings leucomaenis. Localization: body cavity, connective tissue. Distribution: British Columbia. Description (after Kuitunen-Ekbaum, 1933a). Male. Length 35 mm, width 0.35 mm. Mouth simple. Esophagus consisting of an anterior part and a longer posterior part. Length of eso- phagus 2.4mm. Posterior end tapering and spirally coiled. Anus sub- terminal, situated 0.35 mm from end of tail. Several pairs of postanal papillae present. Spicules of the same length, 0.4 mm long, 0.007 mm wide, curved. Gubernaculum absent. Female. Length 360 mm, width1.3mm. Body brilliant white, live specimens semitransparent. Cephalic end rounded, mouth simple, without lips or papillae. Esophagus cylindrical,2.4mm long. Intestine extending as a flat, dark ribbon throughout the entire body and ending blind, adhering to the body wall with a muscle cord. Anus and vulva atrophied in mature females; the body cavity is almost completely occupied by the uterus, which is filled with larvae; there are two short ovaries, one at each end of the body. Posterior end tapering, slightly twisted. Length of larvae in the uterus 0.54 mm, width 0.03 mm. The body of the larvae tapers posteriorly and ends in a long, thin tail which is as long as the body. Mouth with two small papillae. A thin esophagus and the intestine are visible. Kuitunen-Ekbaum stated that females of Ph. oncorhynchi placed in seawater discharge numerous larvae which die in 2 days. 360 Description (after Fujita, 1940). Body thin, cephalic end widened, posterior end elongate. Cuticle thick, striated with intervals of 0.008 mm on the cephalic and posterior parts. РарШае indistinct. Mouth narrow, slitlike. Length of short, narrow muscular part of esophagus 0.9 mm, width 0.09 mm. Anterior part of esophagus widened to 0.15mm, witha 361thick wall. Posterior part of esophagus 1.6mm long and 0.16 mm wide. Intestine narrow, to 22 mm long and 0.09 mm wide. Rectum and anus usually atrophied; rectum 0.25 mm long in young forms, anus situated 0.063 mm from the posterior end. Nerve ring situated 0.5mm from the cephalic end; excretory pore not recognizable. 350 —— ss ST И (360) FIGURE 206. Philonema oncorhynchi Kuitunen-Ekbaum, 1933: a,b — anterior end; с — posterior end of female; d — larva; e,f— posterior end of male, different aspects; g — first-stage larva, lateral; h — second-stage larva, cephalic end; i— cephalic end of young female, apical; j — anterior end of third-stage larva; k — posterior end of third-stage larva; 1 — posterior end of young male, ventral (a—f — after Kuitunen-Ekbaum, 1933; р—1 — after Platzer and Adams, 1967). Male. Length 35mm, width 0.3mm. Two spicules, 0.39 mm long, with pointed end. Caudal papillae rounded and sessile; 4 pairs of preanal papil- lae arranged in two groups; six pairs of postanal papillae in one group. Caudal wings dark. Female. Length 100 mm, width0.9mm. Genitalia paired. Vulva and vagina atrophied in adults. Vulva of immature females situated slightly before middle of body. Vagina 1 mm long, directed posteriorly; uterus thin, markedly widening. 351 The life cycle of Ph. oncorhynchi was studied by Platzer (1964) and Platzer and James (1967) in Lake Cultus (Canada), where the intermediate host is Cyclops bicuspidatus. Development in the definitive host was observed in naturally infected young sockeye salmon which had been kept in fresh water for 2 years. Mature females of Ph. oncorhynchi burst in fresh water after a minute. The larvae discharged remain suspended in the water for several days; some first-stage larvae may live for 17 days in lake water at 8°C. Cyclops swallow suspended larvae and larvae lying on the bottom. There were usually one or two larvae in the body cavity of a Cyclops, but sometimes as many as seven. Differences in the behavior of infected and uninfected Cyclops were not observed. The first-stage larvae have a long, threadlike tail which tapers sharply. Mouth surrounded by papillae, with a triangular denticle dorsally. Esophagus apparently not divided, with numerous refracting particles. Nerve ring situated almost in middle of esophagus. Intestine consisting of 18—30 cells with a distinct nucleus, not united with the proctodeum. The genital primordium consists of 4 cells, the nucleus of which contains a large quantity of confluent chromatin, Rectum with thin wall, anus distinct. Second-stage larva. After 25 days at 8°C and 17 days at 12°C infected Cyclops contained molted larvae, but the cuticle of the first- stage larva had not been shed. The second-stage larva is shorter than the first-stage larva and shows the following differences: dorsal tooth absent, intestine narrow, nuclei protruding above the surface of the intestine, rectum thicker, tail shorter and sharply tapering; genital primordium not visible; the larvae did not move actively in the hemocoel but made slow bending movements in response to the movements of the host. At the end of the second stage the larvae were as long as the first-stage larvae, and the esophagus was already divided into a narrower muscular part and a thicker glandular part. Third-stage larvae were found insome Cyclops after 17 days at 12°C. They differ from the late second-stage larvae as follows: glandular part of the esophagus distinct, occupying half the length of the larva and filling the pseudocoel; larvae very mobile, both in the body cavity and 362 outside it; rectum narrow, with thin wall; distinct esophageal-intestinal valves projecting into the intestine. The body cavity of Cyclops is almost entirely filled with second- stage larvae. Third-stage larvae from an experimentally infected fish were identical with larvae from Cyclops. However, larvae removed from fish 4 and 5 days after infection were smaller than third-stage larvae from Cyclops. Larvae taken from fish 9—10 days after infection were of the same size as the larvae from Cyclops. Younger larvae are distributed mainly in the peritoneum near the body wall and in the serosa of the swim bladder; older larvae are localized mainly in the wall of the swim bladder. Fourth-stage larvae appear after 15 months. Average length of female larva 4.80 mm, of male larva 3.90 mm. Uterus of female didelphic, vulya not fully developed. The genital duct of the male is connected with the:ventral wall of the cloaca, but the spicules are not yet fully developed. 352 This stage was later found in the body cavity together with the subimagines, which appear after 20 months. Several subimagines were found in the fish earlier than fourth-stage larvae. Subimaginal females 42 mm long had a large uterus, ovaries, and a vulva. A subimaginal male 18 mm long had spicules, anal papillae, a tridentate gubernaculum, and genital ducts with regional differentiation and mobile sperm. The cephalic and caudal papillae are indistinct in adults, and have therefore not been described. They are distinct in immature live speci- mens or in specimens which have just died but have not yet been fixed. The preanal papillae are sessile; the postanal papillae project above the cuticle. The papillae of both types contain nerve endings. The papillae are arranged in pairs and singly, and their number is not constant. There are 10 pairs of preanal papillae, 2 pairs in the depression of the cloaca, and 10 pairs of postanal papillae. The mouth is slitlike and surrounded by an inner ring of 6 sickle-shaped papillae, an outer ring of 4 pairs of papillae, and a pair of amphids behind the interolateral papillae. The authors noted the long period of development in the definitive host, probably caused either by the low water temperature required by the host (Salmonidae) or by adaptation of the parasite: the protracted development of the nematodes coincides with the development of the host. Females of Ph. oncorhynchi are usually filled with developed first- stage larvae when the salmon spawns. The fish probably become infected in January—February, since young salmon do not leave the lake for the sea before April and can therefore contain a large number of infective larvae in the wall of the swim bladder. The authors observed that the larvae begin to develop in the swim bladder only at the age of 18 months, that is, about 6 months after the migration of the salmon to the sea. At the age of 32 months, when the fish usually begin to migrate to fresh water for spawning, almost adult nema- todes are present in the body cavity. During the migration, the gonads of the fish increase in size, and the parasites grow and mature accordingly. Fish ready to spawn contain nematodes, the uterus of which is filled with 363 developed larvae. When the fish arrive at the spawning grounds, the larvae are ready to leave the uterus. The authors assumed that such an exact correlation between the de- velopment of larvae of Ph. oncorhynchi and the maturation and spawn- ing of their host is determined by hormones of the salmon. Bashirullah (1967) experimentally established that the rapid development of Ph. oncorhynchi is parallel to the rapid development of the gonads of the fish during the last 6 months of its life: immature salmon contained only immature nematodes, but females of Ph. oncorhynchi containing larvae were found only in mature fish aged 2,3, and 4 years. References: Akhmeroy, 1955, pp.99—137; Rumyantsev, 1965, рр. 1082—1083; Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, т954, р: 80; Bashirullah, 1967, p.3332; Fujita, 1939, pp. 260—262; 1940, рр. 389—390; Kuitunen-Ekbaum, 1933a, рр. 71—75; Platzer, 1964, р. 561; Platzer and Adams, 1967, pp. 41—43; Rasheed, 1963, рр.89—130; Smedley, 1933, рр. 169—179. 353 Philonema agubernaculum Simon and Simon, 1936 (Figure 207) Hosts: Salmonidae — Prosopium williamsoni, Salvelinus Гоа а Эа имо assh. a. Settare Localization: body cavity. FIGURE 207. Philonema agubernaculum Simon and Simon, 1936: a — cephalic end; b — larva; c,d — posterior end of male, lateral; 9 — anterior end; e — posterior end of female (a—c — after Fujita, 1940; d—f — after Simon and Simon, 1936). 364 Distribution: North America. Description (after Simon and Simon, 1936). Male. Length 12—23 mm, width 0.296 mm. Esophagus muscular- glandular, cylindrical, 1.526 mm long,0.189 mm wide. Length of muscular anterior part 0.404 mm. Anus situated 0.276 mm from end of tail. Spi- cules of the same length, markedly curved, weakly chitinized, 0.284 mm long on the outside of the curve. Gubernaculum absent. Female. Length 49—140 mm, width 0.771 шт. Length of esophagus 1.725 mm, width 0.220 mm; length of anterior part 0.553 mm. Anus situated 0.446 mm from end of tail. In females with eggs the anus is sometimes atrophied, and vulva and vagina are completely atrophied. The uterus of mature females is filled with embryos and occupies almost the entire body cavity. The uterus of young females contains eggs at different stages of de- velopment, 0.032 Х 0.046 mm large. Length of embryo 0.343 mm. 354 Meyer (1958, 1960) and Vik (quoted in Platzer and Adams, 1967) studied the development of Ph. agubernaculum in Salvelinus fontinalis and Salmo salar. They assumed that adult females leave the fish when they spawn, and the larvae are released when the females burst in the water. Meyer and Vik infected Cyclops sp. withlarvae of Ph. agubernacu- lum. However, when they fed the infected Cyclops to the trout, it did not become infected. Vik obtained larvae of Ph. agubernaculum from the tissues of OSmerus mordax by pepsin digestion and infected trout with them. However, he thought that further experiments are neces- sary to determine the part played by Osmerus in the life cycle of Bae asubernacululm. References: Skrjabin, Shikhobalova, Sobolev, Paramonov, and Sudarikov, 1954, р.80; Meyer,1958, p.42; 1960; Simon, 1936, рр. 440—442. Philonema sibirica (Bauer, 1946) Rumyantsev, 1965 (Figure 208) Eaomym: Coregonema Srvbirica Bauer, 1946 Hosts: Coregonidae. Localization: body cavity. Distribution: water bodies in the north of the USSR. Historical review The original description of the species (1946) was very brief and a drawing was not given. Bauer established the new genus Coregonema, but the diagnosis was not sufficiently detailed and the key given by the author gave only differences of two genera: Philometra and Dracunculus. Several studies had appeared by 1948 in which records of the species were given and the list of its hosts was enlarged, proving that the species is widely distributed in Siberia. Akhmerov (1955) made Coregonema a synonym of the genus Pimvonmenva, and ©, sibirica a.subspecies of Ph, oncorhyn eis Rumyantsev stated in 1965 that Coregonema is a synonym of the genus Philonema after a study of the parasites of Coregonus albula. 365 Description (after Bauer,1946,as Coregonema sibirica). Lips absent. Mouth without appendages. Lumen of esophagus widening like a funnel at the beginning. Anterior end rounded, posterior end pointed. Males 10—15 times shorter than females. Male. Lengthto16 mm. Length of esophagus 2mm. Glandular part of esophagus 1.35 mm long, muscular part 0.65 mm. Spicules of the same length, 0.350—0.360 mm. Length of gubernaculum 0.120 mm. Female. Lengthto150 mm. Length of glandular part of esophagus 0.9mm, of muscular part 0.5mm. Anus and vulva atrophied in mature females. Viviparous. Description (after Rumyantsev, 1965). Male. Length 11.2mm, width 0.16 пт. Length of esophagus 1.04 mm, width 0.11mm. Muscular part of esophagus shorter than glandular part, 0.43 mm long. Ratio of length of muscular to glandular part = 1:1.5. 355 FIGURE 208. Philonema sibirica (Bauer, 1946): a — cephalic end, apical; b — anterior end; c — posterior end of male; d — anterior end; е — cephalic end, apical; f — posterior end of male (a—c — after Bauer, 1946; d—f— after Rumyantsev, 1965). Tail pointed and spirally twisted. Spicules of the same length, distinctly curved, 0.25 mm long and 0.006 mm wide. Gubernaculum absent. Female. Length 30 mm, width 0.36 mm. Cephalic end rounded, with a simple mouth, Length of esophagus 1.17 mm, width 0.19 mm. Length | of muscular part of esophagus 0.49 mm. Lumen of esophagus slightly Pe widened at the beginning. Posterior end narrowing to a straight or slightly к. curved, pointed process. 4 366 References: Akhmerov, 1955, рр. 97—187;. Bauer, 1946, ‘р. 161960 рр. 97—156; 1968b, pp.157—174; Bauer and Greze, 1948, рр. 186—194; Bauer y and Nikol'skaya, 1948, рр. 175—176; Petrushevskii and Bauer, 1948, рр. 217— 231; Rumyantsev, 1965, pp. 1082—1083. Г aw Se Ee a Philonema sp. Naidenova and Nikolaeva, 1968 (Figure 209) Aon Роны маты”. pint Localization: body cavity. Distribution: Aegean Sea. Description (after Naidenova and Nikolaeva, 1968). Cephalic end rounded, posterior end pointed. Mouth simple, without lips but with papillae. Male unknown. ; Female. Length 2.0—2.5 mm, width 0.264—0.279 mm. Esophagus a. slightly widened anteriorly and posteriorly, cylindrical, 0.238—0.248 mm long 356 and 0.009—0.016 mm wide. Anus and vulva atrophied. The two short ovaries are situated at both ends of the body. The loop of the ovary in the anterior part of the body does not extend beyond the connection between esophagus and intestine. Body cavity of females occupied by the uterus with developing embryos 0.120—0.150 mm long and 0.006—0.009 mm wide. а b FIGURE 209. Philonema sp. Naidenova and Nikolaeva, 1968: а — anterior end; b — posterior end of female (after Naidenova and Nikolaeva, 1968). Reference: Naidenova and Nikolaeva, 1968, p. 74. 357 BIBLIOGRAPHY Publications in Russian Agapova, А.Т. Parazity ryb vodoemov Zapadnogo Kazakhstana (Parasites of Fish from the Water Bodies in Kazakhstan). — Trudy Instituta Zoologii AN KazakhSSR, \01.5:5—60. Alma-Ata. 1956. Agapova, А.Т. Philometra leucisci —novyi vid nematody и губ Kazakhstana (Philometra leucisci, a New Species of Nematode in Fish of Kazakhstan). — In: “Parazity dikikh zhivotnykh Kazakhstana,” рр.137—139. Alma-Ata,Izdatel'stvo AN KazakhSSR. 1963. Agapova, A.I. Parazity ryb vodoemov Kazakhstana (Parasites of Fish from the Water Bodies of Kazakhstan), рр.1—342. Alma-Ata. 1966. Agapova,A.I. and А.Р. Maksimova. Parazity ryb vodoemov Yuzhnogo Kazakhstana. — Parazity zhivothnykh Kazakhstana (Parasites of Fish from the Water Bodies of South Kazakhstan — Parasites of Animals in Kazakhstan).— Trudy Instituta Zoologii AN KazakhSSR. Vol.14:71—87. Alma-Ata. 1960. Akhmerov, A. Kh. Parazitofauna ryb reki Kamchatki (The Parasitic Fauna of Fish from the River Kamchatka). — Izvestiya Tikhookeanskogo Instituta Rybnogo Khozyaistva i Okeanografii, Vol.43:99— 137. 1955. Bauer, O.N. Parazitofauna ryapushki iz razlichnykh vodoemov SSSR (The Parasitic Fauna of Coregonus from Water Bodies in the USSR). — Trudy Leningradskogo Obshchestva Estestvoispytatelei, 69, No.4:7—21. 1946. Bauer, O.N. Parazity ryb reki Eniseya (Parasites of Fish from the Yenisei River). — Izvestiya Vsesoyuznogo Instituta Ozernogo i Rechnogo Rybnogo Khozyaistva, Vol.27:97—156. 1948а. Bauer, О.М. Parazity ryb reki Leny (Parasites of Fish from the Lena River). — Ibid., Vol.27:157—174. 1948b. Bauer, О.М. and У.1. Greze. Parazity ryb ozera Taimyr (Parasites of Fish from Lake Taimyr). — Ibid., Vol.27:186—194. 1948. Bauer, О.М. and М.Р. Nikol'skaya. К poznaniyu parazitov ryb reki Anadyr’ (Contribution to the Study of Parasites from the Anadyr River). — Ibid., Vol.27:175—-176. 1948. Belous, E.V. Paraziticheskie chervi presnovodnykh pozvonochnykh Primorskogo kraya (Parasitic Worms in Freshwater Vertebrates of the Maritime Territory). Candidate Thesis. — Vsesoyuznyi Institut Gel'mintologii im. akad. К.Т. Skryabina. Moskva. 1952. Belous, E.V. Nematody presnovodnykh ryb Primorskogo kraya (Nematodes of Freshwater Fish from the Maritime Territory). — In: "Paraziticheskie chervi domashnikh i dikikh zhivotnykh,” рр.48—65. Vladivostok. 1965. Boev, S.N. Issledovaniya po prirodnoi ochagovosti gel’'mintozov у Kazakhstane (Studies on the Natural Focality of Helminthiases in Kazakhstan). — Trudy Instituta Zoologii AN KazakhSSR, Vol.22:4. 1964. Bykhovskaya-Pavlovskaya. I.E. Vliyanie vozrasta na izmenenie parazitofauny u okunya (The Influence of Age on Changes of the Parasitic Fauna of Perch). — Parazitologicheskii Sbornik Zoologicheskogo Instituta AN SSSR, Vol. 8:99— 130. 1940. Bykhovskaya-Pavlovskaya, I.E. and В.Е. Bykhovskii. Parazitofauna ryb Akhtarinskikh limanov (Azovskoe more, del'ta reki Kubani) (Thr Parasitic Fauna of Fish from the Akhtar Limans (Sea of Azov, Kuban River Delta)). — Ibid., Vol. 8:131—161. 1940. Bykhovskaya-Pavlovskaya, Т.Е., A.V. Gusev, М.М. Dubinina, T.S. Smirnova, L.L. Sokolovskaya, С.А. Shtein, $.5. Shul'man, and V.M. Epshtein. Opredelitel’ parazitov presnovodnykh ryb SSSR (Key to Parasites of Freshwater Fish in the USSR), рр.1—776. — Moskva-Leningrad, Izdatel'stvo AN SSSR. 1962. Bykhovskii, B.E. Parazitologicheskie issledovaniya na Barabinskikh ozerakh (Parasitological Studies on the Baraba Lakes). — Parazitologicheskii Sbornik Zoologicheskogo Instituta AN SSSR, Vol.6:4387— 482. 1936. 359 Bykova, E.V. К poznaniyu parazitofauny promyslovykh ryb ozera Chelkar (Contribution to the Study of the Parasitic Fauna of Commercial Fish from Lake Chelkar). — Uchenye Zapiski Leningradskogo Gosudarstvennogo Universiteta, Seriya Biologicheskikh Nauk, 43, №.11:33—43. 1939. Chernyshev,V.I. К ekologii i parazitofaune shakala iz Tadzhikistana (On the Ecology and Parasitic Fauna of the Jackal from Tadzhikistan). — Trudy Instituta Zoologii i Parazitologii AN TadzhSsR. Vol.21:151—165. 1954. Ch'ung-Hsiung, Е. Drakunkulez и sobak у Kazakhstane (Dracunculiasis in Dogs of Kazakhstan). — Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 27, No.2:219. 1958. Daiya, G.G. Nematody gusinykh ptits nizov'ya reki Obi (Nematodes of Anseriformes from the Lower Reaches of the Ob River). — Problemy Parazitologii, рр.142—145. Kiev, "Naukova Dumka.” 1967. Роре!', V.A. Problemy issledovaniya parazitologii ryb (Studies of the Parasitology of Fish). — Trudy Leningradskogo Obshchestva Estestvoispytatelei, 62, No.3:247—268. 1933. Dogel', V.A. and A. Kh. Akhmerov. Nematody ryb reki Amur (Nematodes of Fish from the Amur River). — Acta hydrobiol. sin., Vol.3:287—304. 1959. ‘ Роре! ', V.A. and В.Е. Bykhovskii. Parazity ryb Kaspiiskogo morya (Parasites of Fish from the Caspian Sea). — Moskva, Izdatel'stvo АМ SSSR. 1934. 151 pp. Dubinin, V.B. Vliyanie osoloneniya reki Malyi Uzen' na parazitofaunu naselyayushchikh ee ryb (The Effect of Salinization of ше Malyi Uzen' River on the Parasitic Fauna of Fish). — Zoologicheskii Zhurnal, 27, No.4:335—342. 1948. Dubnitskii, A.L. Pervyi v SSSR sluchai obnaruzheniya drakunkuleza u lisits (The First Record of Dracunculiasis of Foxes in the USSR). — Trudy Tsentral'noi Nauchno-Issledovatel'skoi Laboratorii Pushnogo Zverevodstva, Vol. 6:323—328. 1950. Fedchenko, А.Р. О stroenii i razmnozhenii rishty (On the Structure and Reproduction of the Guinea Worm Dracunculus medinensis). — Izvestiya Obshchestva Lyubitelei-Estestvoispytatelei, 8, No.1:71—78. 1870. Fedchenko, A.P. Zoologicheskie zametki k anatomii kruglykh chervei (Zoological Notes on the Anatomy of Nematodes). — Izvestiya Obshchestva Lyubitelei-Estestvoispytatelei, 10, No.11:51—68. 1873. Gavrilova, N.G. Zoogeograficheskii analiz parazitov ryb Amudar'i i Syrdar'i у predelakh Turkmenskoi provintsii (Zoogeographical Analysis of Parasites of Fish of the Amu Darya and Syr Darya Rivers in Turkmenia). — Parazitologiya, 2, No.3:232—236. 1968. Gnedina, М.Р. and N.V. Savina. 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О sposobe proniknoveniya lichinok rishty у tsiklopov (On the Penetration of Guinea Worm Larvae into Cyclops).—Meditsinskaya Parazitologiya i Parazitnarnye Bolezni, 3, No. 3:212;230. 1934a. Isaev, L.M. Eksperimental'naya rishta u sobak (Experimental Infection of Guinea Worm in Dogs). — Ibid., 3, №.3:231—238. 1934b. Isaev, L.M. Prostoi sposob massovogo issledovaniya rakoobraznykh na lichinki paraziticheskikh chervei (A Simple Method of Studying Larvae of Parasitic Worms in Crustacea). — Ibid., 3, No. 3:238—240. 1934c. Ivashkin, V.M. Osvyazi mezhdu tolshchinoi skorlupy уаз spirurat i gruppoi ikh promezhutochnykh khozyaev (On the Relationship between the Thickness of the Egg Shell of Spirurata and their Intermediate Hosts). — Tezisy dokladov nauchnoi konferentsii Vsesoyuznogo obshchestva - gel'mintologov, p.56. Moskva. 1958. Ivashkin, V.M. 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K epidemiologii drakunkuleza v Gane (On the Epidemiology of Dracunculiasis in Ghana). — Meditsinskaya Parazitologiya i Parazitarnye Bolezni, 37, No. 7:347—350. 1968. Lyaiman, E.M. Paraziticheskie chervi ryb zaliva Petra Velikogo (Parasitic Worms in Fish from Peter the Great Bay). — Izvestiya Tikhookeanskoi Nauchno-promyslovoi Stantsii, 3, No. 6:120. Vladivostok. 1930. Lyaiman, Е.М. Bolezniryb, prichinyaemye paraziticheskimi chervyami (Diseases of Fish Caused by Parasitic Worms). — Moskva-Leningrad, Pishchepromizdat. 1934. Lyaiman, Е.М. Kurs boleznei ryb (A Course in Diseases of Fish). — Moskva, Pishchepromizdat. 1949. ‚ Матает, Yu.L., A.M. Parukhin, О.М. Baeva, andP.G. Oshmarin. Gel'mintofauna dal'nevostoch- nykh lososevykh у svyazi $ voprosom о lokal'nykh stadakh i putyakh migratsii etikh ryb (Helminth Fauna of Pacific Salmonidae in Relation to the Local Populations and their Migrations). Vladivostok. 1959. 74 pp. Markevich, A.P. 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Publications in Other Languages* Adams J. В. 1969. Migration route of invasive juvenile Philonema oncorhynchi (Nema- toda: Philometridae) in young salmon.— J. Fish. Res. Board Canada, 26, Ne 4, p. 941—946. Agarwal М. P. 1930. А new nematode Procamallanus тейгй п. sp., from а loca! siluroid fish — Wallago attu.— Allahabad Univ. Studies, 6, № 2, р. 59—64. Agarwal S. C. 1958. On a new species of Procamallanus Baylis 1923 (Nematoda) Procamallanus spiculogubernaculus. — Current Sci., 27, p. 348—349. Agrawal V., 1966a. Three new reptilian nematodes from Lucknow. — Trans. Amer. Microscop. Soc., 85, Ne 1, p. 107—114. Agrawal V. 1966b. On a new nematode Procamallanus muelleri n. sp. from the stomach of a freshwater fish Heteropneustes fossilis. — Proc. Helminthol. Soc. Washington, 33, № 2, р. 204—208. АА : Agrawal V. 1967. Some new Camallanoidea (Spirurida) nematodes from fishes, am- phibias and reptiles. — Ann. parasitol. humaine et compar., 42, Ne 3, p. 327—342. Ali $ М 1956. Studies on the nematode parasites of fishes found in Hyderabad state. — idiat inthol., 8, р. 1—83. | Che aM SM Hee Goon г species of Procamallanus Baylis, 1923 from India with a key to the species. — Indian J. Helminthol. 34, № 1/2, р. 129—138. et Anantaraman М. 1966. Dracuntiasis in animals. — Proc. 1 Internat. Parasitol. Congr. г . 798—799. Mode НЕ, г 1946. А new nematode, Procamallanus регеги with a key to the genus. — Trans. Amer. Microscop. Soc., 65, № 4, р. 299—303. 4 ' а Annigeri G. С. 1962. А viviparous nematode, Philometra sp. in the ее (9) lithus argenteus (Cuvier).— J. Marine Biol. Assoc. India, 4, Ne 2, p. 263—265. * [Reproduced from the Russian original.] 364 Вагиз У., Могауес Е. 1967. А survey of helminths from the Cuban turtle Pseudemys decussata Gray (Emydidea).— Vestn. Ceskosl. spol. zool., 31, Ne 4, str. 313—394. BaruS V., Могауес Е. 1969. Three interesting nematodes from Aonyx cinerea (Carni- vora) from Malaya.— Folia parasitol. (Praha), 16, Ne 3, str. 235—236. 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Helminthol., 34, Ne 1—2, p. 103—106. Yeh Г. $. 1960b. On the reconstruction of the genus Camallanus Railliet et Henry, 1915. — J. Helminthol., 34, Ne 1—2, p. 107—116. Yeh L. S. 1960c. On a collection of Camallanid nematodes from fresh water fishes in Ceylon. — J. Helminthol., 34, Ne 1—2, p. 117—124. Yorke W., Maplestone P. A. 1926. The nematode parasites of vertebrates. London. Zeder J. G. H. 1803. Anleitung zur Naturgeschichte der Eingeweidewurmer. Bamberg. Zschokke Е., Heitz A. 1914. Endoparasiten aus Salmoniden von Kamtschatka. — Rev. suisse Zool., 22, Ne 8, p. 205—217. 376 INDEX OF СЕМЕРА* A Agrachanus Tichomirova, in litt. 228 Anguillicola Yamaguti, 1935 219 Avioserpens Wehr et Chitwood, 1934 262 B Buckleyella Rasheed, 1963 320 С Camallanides Baylis et Daubney, 1922, 90 Camallanus (Railliet et Henry, 1915) 29 Camallanus Railliet et Henry, 1915 (in part.) 101 Camallanus Railliet et Henry, 1915 (in part.) 104 Camailanus Railliet et Henry, 1915 (in part.) Camalianus Railliet et Henry, 1915 (in part.) 138 Coregonema Bauer, 1946 358 Cucullanus Mueller, 1777 (in part.) 138 D Dracunculus Reichard, 1759 239 F Filaria Mueller, 1787 (in part.) 239 Filaria Mueller, 1787 (in part.) 287 Fuellebornius Leiper, 1926 239 Furia Linnaeus, 1758 239 G Gordius Linnaeus, 1758 (in part.) 239 Ichthyofilaria Yamaguti, 1935 321 Ichthyonema Diesing, 1861 287 Indocamallanus Chakravarty, Majumdar, Sain, 1963 138 M Micropieura Linstow, 1906 279 Molnaria Moravec, 1948 231 N Neocamallanus Ali, 1956 29 Nilonema Khalil, 1960 322 О Oshimaia Sugimoto. 1934 262 P Paracamallanus Yorke et Maplestone. 1926 101 Philometra Costa, 1945 287 Philometroides Yamaguti, 1935 326 Philonema Kuitunen-Ekbaum, 1933 358 Phlyctainophora Steiner, 1921 222 Piscilania Yeh, 1960 103 Procamallanus Baylis, 1923 138 Pseudophilomtetroides Parukhin, 1966 344 R 1960 345 $ Sanguinofilaria Yamaguti, 1941 287 Serpinema Yeh, 1960 104 Skrjabillanus Schigin et Schigina, 1958 225 Skrjabillanus Schigin et Schigina. 1958 (in part.) 228 Spirocamallanus Rumai Travassos, Olsen, 1952 173 T Tetanonema Steiner, 1937 236 Thelazo Pearse, 1933 138 Thwaitia Rasheed, 1963 346 У Vena Gallandat, 1773 239 Vermiculus Dunglinson, 1898 239 Z Zeylanema Yeh, 1960 120 INDEX OF SPECIES A abdominalis Nybelin, 1928 Philometra 348 abdominalis (Nybelin, 1928) Rasheed, 1963 Thwaitia 348 abramidis Osmanov, 1964 Philometra 292 aethiopica Valenciennes, 1856 Filaria 244 aethiopicus Diesing, 1861 Dracunculus 244 agubernaculum Simon et Simon, 1936 Philo- nema 363 alatus Rudolphi, 1802 Cucullanus 31 alii Deshmukh, 1969 Dracunculus 252 alii Rasheed, 1963 Philometra (Alinema) 317 amarali Vaz et Pereira, 1934 Procamallanus 174 * [Reproduced from the Russian original. Page numbers refer to those in the Russian book, which appear in the left-hand margin of the translation.] 377 amarali (Vaz et Pereira, 1934) Olsen, 1952 Spirocamallanus 174 amazonica Travassos, 1960 Philometra 292 amazonicus Ribeiro, 1941 Camallanus 106 amazonicus (Ribeiro, 1941) Yeh, 1960 Serpi- nema 106 americanus Magath, 1919 Camallanus 111 americanus Magath, 1919 Camallanus (in part.) 119 amabantis Pearse, 1933 Camallanus (in vart.) 12] anabantis Pearse, 1933 Camallanus (in part.) 130 anabantis (Pearse, 1933) Yeh, 1960 Zeylane- ma 121 anguillae (Ishii, 1916) Rasheed, 1963 Philo- metroides 327 anguillae Schrank, 1798 Cucullanus 31 armatus Campana-Rouget et Therezien, 1965 Procamallanus 140 uspicuius Khera, 1955 Procamallanus 142 atridentus Khera, 1954 Camallanus 126 utridentus (Khera, 1954) Yeh, 1960 Zeyla- nema 126 utropi Parukhin, 1966 Pseudophilometroides 344 attui Pande, Bhatia, Rai, 1963 Procamallanus 143 australiensis Johnston et Mawson, 1940 An- guillicola 221 bagarii Karve et Naik, 1951 Procamallanus 175 bagarii (Karve et Naik, 1951) Olsen, 1952 Spirocamallanus 175 bagri Khalil, 1965 Thwaitia 350 balistii Rasheed. 1963 Thwaitia 347 barroslimai Pereira, 1935 Procamallanus 177 barroslimai (Pereira, 1935) Olsen, 1935 Spi- rocamallanus 1777 baylisi Karve, 1930 Camallanus 43 baylisi Vaz et Pereira, 1934 Philometra 293 bifidus Olsen, 1952 Avioserpens 266 biglobocerca Belouss, 1965 Philometra 294 brevis Kung, 1948 Procamallanus 144 buckleyi Rasheed, 1963 Buckleyella 320 bufonis Agrawal, 1967 Camallanus 44 bungari (MacCallum, 1918) Yamaguti, 1961 Camallanides 94 С capillaris Denglison, 1895 Vermiculus 244 carangis Olsen, 1954 Camallanus 45 carassti Ishii, 1933 Philometra 340 cearensis Pereira, Vianna Dias, Azevedo, 1936 Procamallanus 178 cearensis (Pereira, Vianna Dias, Azevedo, 1936) Olsen, 1952 Spirocamallanus 178 cernuae Gmelin, 1790 Cucullanus 31 ceylonensis Fernando et Furtado, 1963 Ca- mallanus 47 chakravartyi (Chakravarty, Majumdar, Sain, 1961) Fernando et Furtado. 1963 Procamal- lanus 145 chelydrae MacCallum. 1918 Camallanus 111 chelydrae MacCallum, 1918 Camallanus (т part.) 119 378 chimuzensis Freitas et Ibanez, 1968 Spiroca- mallanus 181 clarius Ali, 1956 Procamallanus 146 confusus Railliet et Henry, 1915 Camallanus confusus Fernando et Furtado, 1963 Proca- mallanus 149 coronatus Zeder, 1800 Cucullanus 31 cotti Fujita, 1927 Camallanus 48 cryptocentri Yamaguti, 1961 Philometra 297 cyathocephalus MacCallum, 1918 Camallanus 111 cyathocephalus MacCallum, 1918 Camallanus 119 cyathopharinx Baylis, 1923 Camallanus 102 cyathopharynx (Baylis, 1923) Yorke et Maple- stone, 1926 Paracamallanus 102 cynophylectis Sahay, 1966 Camallanus 50 D daccai Gupta, 1959 Procamallanus 150 dahomensis Neuman, 1895 Dracunculus 252 dahomensis (Neuman, 1895) Filaria 252 dasycotti Yamaguti, 1935, Ichthyofilaria 321 denticulatus Rasheed, 1965 Philometroides 328 denticulophasma Wehr et Chitwood, 1934 Avioserpens 266 devendri Sinha et Sahay, 1966 Procamallanus 151 dhamini Deshmukh, 1968 Camallanides 95 doi Chabaud, 1960 Dracunculus 252 dumerillii Perrier, 1871 Cucullanus 111 E elegans MacCallum, 1918 Camallanus 111 elegans MacCallum, 1918 Camallanus 119 elegans Zeder, 1800 Cucullanus 31 elongata Fujita, 1939 Philonema 359 equispiculus Sood, 1968 Camallanus 51 erythrophthalmi (Molnar, 1966) Moravec, 1968 Molnaria 231 erythrophthalmi Molnar, 1966 Skrjabillanus 231 F fariasi Pereira, 1935 Procamallanus 182 fariasi (Pereira, 1935) Olsen, 1952 Spiroca- mallanus 182 fernandoi Yeh, 1960 Zeylanema 127 floridianae MacCallum, 1918 Camallanus 111, 119 fuelleborni Travassos, 1934 Dracunculus 244 fujimotoi Furuvama, 1932 Philometra 295 fulvidraconis Li. 1935 Procamallanus 182 fulvidraconis (Li, 1935) Olsen, 1952 Spiro- camallanus 182 G galliardi Chabaud et Campana, 1949 Avioser- pens 268 globiceps Yamaguti, 1935 Anguillicola 219 globiceps Rudolphi, 1819 Filaria 288 globiceps (Rudolphi, 1819) Railliet, 1916 Philometra 288 globocephalus Mackin, 1927 Dracunculus 252 globoconchus Ali, 1960 Procamallanus 188 globoconchus (Ali, 1960) Spirocamallanus 188 glossogobii Pearse, 1933 Thelazo 152 glossogobii (Pearse, 1933) Southwell et Kirschner, 1937 Procamallanus 152 graecorum Grtiner, 1777 Dracunculus 244 gubernaculus Khera, 1955 Procamallanus 190 gubernaculus (Khera, 1955) Spirocamallanus ~190 guinensis Dunglison, 1893 Filaria 244 gymnarchi Khalil, 1960 Nilonema 322 H hemidenta Majumdar, 1965 Camallanides 96 heteropneusti Chakravarty, Majumdar, Sain, 1961 /ndocamallanus 145 heteropneusti Chakravarty, Majumdar, Sain, 1961 Neocamallanus 145 heteropneustus Ali, 1956 Procamallanus 154 hilarii Vaz et Pereira, 1934 Procamallanus 192 hilarii (Vaz et Pereira, 1934) Olsen, 1952 Spirocamallanus 192 hindenensis Lal, 1965 Procamallanus 155 houdemeri Нза, 1933 Dracunculus 252 hyderabadensis Rasheed, 1963 Philometra 296 hyderabadensis Ali, 1956 Procamallanus 193 hyderabadensis (Ali, 1956) Spirocamallanus 193 hypophthalmichthys Achmerov, 1954 Camal- lanus 52 пейте? Travassos, Artigas et Pereira, 1928 Procamallanus 195 theringi (Travassos, Artigas et Pereira, 1928) Olsen, 1952 Spirocamallanus 195 indica Khera, 1951 Micropleura 281 inglisi Agrawal, 1967 Camallanus 54 inimici Yamaguti, 1941 Philometra 297 inopinatus Travassos, Artigas et Pereira, 1928 Procamallanus 196 inopinatus (Travassos, Artigas et Pereira, 1928) Olsen, 1952 Spirocamallanus 196 insignis (Leidy, 1858) Dracunculus 244 insignis Leidy, 1858 Filaria 244 intestinalis (Dogiel et Bychowsky. 1934) Mo- гауес, 1968 Molnaria 235 intestinalis Dogiel et Bychovsky, 1934 Phi- lometra 235 intermedius Hsti et Hoeppli, 1935 Camallanus 104 intermedius (Нзй et Hoeppli, 1931) Yeh, 1960 Serpinema 104 istiblenni Noble, 1966 Spirocamallanus 197 J johni Yeh, 1960 Camallanus 52 K kaapstaadi Southwell et Kirschner, 1937, Ca- mallanus 55 kachugae_ Baylis et Daubney, 1922 Camal- lanus 108 ; kachugae (Baylis et Daubney, 1922) Yeh, 1960 Serpinema 108 kerri Pearse, 1933 Procamallanus 199 kerri (Pearse, 1933) Olsen, 1952 Spirocamal- ‘lanus 199 kirandensis Baylis, 1928 Camallanus 56 kondai Fujita, 1939 Philonema 359 kotlani Molnar, 1969 Thwaitia 351 kulasirii Yeh, 1960 Zeylanema 128 379 L lacustris Yorke et Maplestone, 1926 nec Zoega, 1776 Camallanus 81 lacustris (Zoega, 1776) Camallanus 3! lacustris (Zoega, 1776) Mueller, 1779 Cucul- lanus 31 lacustris Zoega, 1776 Echinorhynchus 31 laeviconchus (Wedl, 1862) Railliet et Henry, 1915 Camallanus 139 laeviconchus Wedl, 1862 Cucullanus 139 laeviconchus (Wedl, 1862) Procamallanus 139 lamnae Steiner, 1921 Phlyctainophora 223 lateolabracis Yamaguti, 1935 Philometra 298 lateolabracis Yamaguti, 1935 Sanguinofilaria 298 leucisci (Agapova, 1963) Moravec, 1968 Mol- naria 235 leucisci Agapova, 1963 Philometra 235 lissemysus Gupta et Singh, 1959 Camallanus Jl longitridentatus Fernando et Furtado, 1963 Camallanus 59 lonis Yamaguti, 1941 Procamallanus 156 lucii Viborg, 1795 Cucullanus 31 lusiana Vismanis, 1966 Philometra 330 lusiana (Vismanis, 1966) comb. п. Philo- metroides 330 lusii Visman, 1962 Philometra 330 M magathi Sprehn, 1932 Camalianus 110 magathi (Sprehn, 1932) Yeh, 1960 Serpinema 110 magnorugosus Caballero, 1939 Camallanus 60 magurii Lal, 1965 Procamallanus 158 malacensis Fernando et Furtado, 1963 Pro- camallanus 159 managatuvo Yamaguti, 1941 Philometra 300 maplestonei Travassos, Artigas et Pereira, 1928 Philometra 335 maplestonei (Travassos, Artigas et Pereira, 1928) Rasheed, 1963 Philometroides 335 mariae Layman, 1930 Philometra 301 marinus Schmidt et Kuntz, 1969 Camallanus 61 mastacembeli Agrawal, 1967 Camallanus 62 mastacembeli Sahay et Sinha, 1966 Zeylanema 129 masu Fujita, 1940 Philometra 336 masu (Fujita, 1940) Rasheed, metroides 336 mathurai Pande, Bhatia, Rai, 1963 Procamal- lanus 160 mazabukae Kung, 1948 Camallanus 65 mazabukae Yeh, 1957 Spirocamallanus 200 medinensis (Linnaeus. 1758) Dracunculus 244 medinensis Gmelin, 1790 Filaria 244 medinensis Faust, 1929 Fuellebornius 244 medinensis Linnaeus, 1958 Gordius 244 medinensis Gallandat, 1773 Vena 244 mehrii Agarwal, 1930 Procamallanus 162 melanocephalus Rudolphi, 1819 Camallanus 103 melanocephalus 103 1963 Philo- Rudolphi, 1819 Cucullanus melanocephalus (Rudolphi, 1819) Yeh, 1960 Piscilania 103 microcephalus (in part.) sensu Chitwood, 1932 Camallanus 119 microcephalus (Dujardin, 1845) Camallanus 111 microcephalus Dujardin, 1945 Cucullanus 111 microcephalus (Dujardin, 1845) Yeh, 1960 Serpinema 111 microcephalus sensu Rausch, 1947 Camal- lanus 119 monotaxis Olsen, 1952 Spirocamallanus mosgovoyi Supriaga, 1965 Avioserpens muelleri Agrawal, 1966 Procamallanus multilineatus Kung, 1948 Camallanus 66 multipapillosa Singh, 1949 Avioserpens multiruga Walton. 1932 Camallanus 67 murrayensis Johnston et Mawson, 1940 Pro- camailanus 202 murrayensis (Johnston et Mawson, Olsen, 1952 Spirocamallanus 202 N nana Mawson, 1957 Avioserpens 268 nodulosus Gupta, 1959 Camallanus 68 nodulosa Thomas, 1929 Philometra 337 nodulosa (Thomas, 1929) Dailey, 1967 Phi- lometroides 337 201 272 164 268 1940) о oachotense Fujita, 1939 Philonema 359 octorugatus Baylis, 1933 Camallanus 117 octorugatus (Baylis, 1933) Yeh, 1960 Serpine- ma 117 oesophagea (Polonio, 1859) Dracunculus 252 oesophagea Polonio, 1859 Filaria 252 olseni Campana-Rouget et Razarihelissoa, 1965 Spirocamallanus 203 oncorhynchi Kuitunen-Ekbaum, nema 359 ophidensis Brackett, 1938 Dracunculus 252 ophiocephali Pearse, 1933 Camallanus 70 ophiocephali Karve, 1941 Paracamallanus 133 ophiocephalus Ali, 1960 Procamallanus 204 ophiocephalus (Ali, 1960) Spirocamallanus 204 opsalichthydis Yamaguti, 1935 Philometra 305 ovata Zeder, 1803 Filaria 301 ovata (Zeder, 1803) Philometra 301 oxycephalus Ward et Magath, 1916 Camal- lanus 71 1933 Philo- P papillosus Zeder, 1800 Cucullanus 31 parasiluri Yamaguti, 1935 Clavinema 305 parasiluri Yamaguti, 1935 Philometra 305 parasiluri Fujita, 1927 Procamallanus 205 parasiluri (Fujita, 1927) Olsen, 1952 Spiro- camallanus 205 parvus Caballero, 1939 Camallanus 110 pearsei Yeh, 1960 Zeylanema 130 pellucida (Jagerskisld, 1893) Philometra 307 percae Schrank, 1788 Cucullanus 31 percalates Johnston et Mawson, 1940 Philo- metra 309 pereirai Annereaux, 1946 Procamallanus 207 pereirai (Annereaux, 1946) Olsen, 1952 Spiro- camallanus 207 pinnicola (Yamaguti, 1935) Philometra 309 Yamaguti, 1941 380 pinnicola Yamaguti, 1935 Sanguinofilaria 309 pipientis Walton, 1935 Camallanus 72 piscatori Khera, 1954 Camallanides 97 planoratus Kulkarni, 1935 Procamallanus 166 plectroplites Johnsion et Mawson, 1940 Phi- lometra 339 plectroplites (Johnston et Mawson, 1940) Rasheed, 1963 Philometroides 339 polynemii Rasheed, 1963 Philometra (Ranjhi- nema) 318 prashadi Baylis et Daubney, 1922 Camallani- des 91 ptyas Khera, 1954 Camallanides 98 R ranae Khera, 1954 Camallanus 73 rarus Travassos. Artigas et Pereira, 1928 Procamallanus 209 гагиз5 (Travassos, Artigas et Pereira, 1928) Olsen, 1952 Spirocamallanus 209, reticaudata Costa, 1845 Philometra 288 rischta Skrjabin, 1917 Philometra 353 rischta (Skrjabin, 1917) Thwaitia 353 rumai Travassos, 1960 Rumai 345 S salmonae Chakravarty, 1942 Camallanus 75 salvelini Fujita, 1939 Philonema 359 sanguinea Rudolphi, 1819 Filaria 340 sanguinea Linton, 1901 Ichthyonema 356 sanguinea (Rudolphi, 1819) Philometra 340 sanguinea (Rudolphi, 1819) Rasheed, 1963 Philometroides 340 scabrae MacCallum, 1918 Camallanus 111 scabrae MacCallum, 1918 Camallanus 119 scardinii (Molnar, 1966) Tichomirova in litt. Agrachanus 228 scardinii Molnar, 1966 Skrjabillanus 228 sciaenae Yamaguti, 1935 Philometra 298 scomberesoxis Nikolaeva et Naidenova, 1964 Philometra 311 scomberomori Yamaguti, 1935 Philometra 298 sebastisci Yamaguti, 1941 Philometra 307 sebastodis Yamaguti, 1941 Philometra 312 senticosa (Baylis, 1927) Rasheed, 1963 Nilo- nema 324 senticosa Baylis, 1927 Philometra 324 senticosa (Baylis, 1927) Travassos, 1966 Phi- lometroides 324 seriolae Ishii, 1931 Filaria 326 seriolae (Ishii, 1931) Philometroides 326 serranelli-cabrillae Janiszewska, 1949 Philo- metra 309 seurati Magath, 1919 Camallanus 111 sibirica Bauer, 1946 Coregonema 364 sibirica (Bauer, 1946) Rumyantzev, 1965 Phi- lonema 364 sigani Yamaguti, 1935 Procamallanus 167 siluri Osmanov. 1964 Procamallanus 182 singhi (Ali, 1956) Yeh, 1960 Camallanus 75 singhi Ali, 1956 Neocamallanus 75 singhi Ali, 1956 Procamallanus 210 singhi (Ai, 1956) Olsen, 1952 Spirocamalla- nus 210 slomei Southwell et Kirschner, 1937 Proca- mallanus 168 spari Yamaguti, 1961 Philometra 313 species Agapova, 1956 Philometra 235 species Ali, 1956 Procamallanus pi species Campana-Rouget, 1961 Spirocamal- lanus 218 / Sphaeroconchus Tornquist, 1931 / Procamal- lanus 169 / spiculogubernaculus Agarwal, 1958 Procamal- lanus 170 7 spinosa Furtado, 1965 Zeylanema 132 spiralis Baylis, 1923 Procamallanus 173 spiralis Baylis, 1923 sensu Tornquist, 1931 Procamallanus 212 spiralis (Baylis, 1923) Olsen, 1952 Spiroca- mallanus 173 strongylurus Steiner, 1937 Tetanonema 237 sweeti Moorthy, 1937 Camallanus 133 sweeti (Moorthy, 1937) Yeh, 1960 Zeylanema 133 sydneyi Rasheed, 1963 Philometra 313 T taiwana (Sugimoto, 1919) Avioserpens 266 taiwana Sugimoto, 1919 Filaria 266 taiwana (Sugimoto, 1919) Oshimaia 266 tauridica Ivashkin, Khromova, Kovaleva sp. п. Philometra 314 tenuicauda Fujita, 1939 Philonema 359 testudinis MacCallum, 1918 Camallanus 111 testudinis MacCallum, 1918 Cucullanus 111 thapari Gupta, 1959 Camallanus 77 thaparus Sahay et Narayan, 1968 Camallanus 78 tincae Rudolphi, 1918, larvae Cucullanus 31 tincae Schigin et Schigina, 1958 Skrjabillanus 226 tornquisti (Baylis, 1923) Campana-Rouget, 1961 Spirocamallanus 212 translucida Walton, 1927 Philometra 356 translucida (Walton, 1927) Rasheed, 1963 Thowaitia 356 trichogasterae Pearse, 1933 Camallanus 80 tridentatus (Drasche, 1884) Camallanus 80 trilabiata Belouss, 1965 Philometra 340 trionyxi Agrawal, 1966 Micropleura 285 trisponosa (Leidy, 1852) Yeh, 1960 Serpinema 119 trispinosus_ (Leidy, 1852) Railliet et Henry, 1915 Camallanus 119 381 trispinosus (Leidy, 1861) Camallanus 111 trispinosus Leidy, 1852 Cucullanus 119 troosti MacCallum, 1918 Camallanus 119 tropica Rudolphi, 1809 Filaria 244 truncatus (Rudolphi, 1814) Camallanus 81 truncatus Rudolphi, 1814 Cucullanus 81 U unispiculus Khera, 1954 Camallanus 121 У а Sinha et Sahay, 1965 Spirocamallanus 1 vazi Travassos, 1933 Micropleura 284 velocissima Diesing, 1851 Ascaris 31 velocissima Nordmann, 1832 (?) Oxyuris 31 vena medinensis Modeer, 1795 Furia 244 vivipara Linstow, 1906 Micropleura 279 viviparus Ali, 1956 Procamallanus 2\4 viviparus (Ali, 1956) Olsen, 1952 Spirocamal- lanus 214 viviparus Bloch, 1782 nec Linstow, 1906 Cu- cullanus 31 W wolgensis Lewaschoffi, 1929 Camallanus 83 wrighti Pereira, 1930 Procamallanus 216 wrighti (Pereira, 1935) Olsen, 1952 Spiroca- mallanus 216 Хх xenopodis Baylis, 1929 Procamallanus 217 xenopodis (Baylis, 1929) Olsen, 1952 Spiro- camallanus 217 Y yehi Fernando et Furtado, 1963 Camailanus 84 Z zacconis Li, 1941 Camallanus 85 zebrini Yamaguti, 1961 Philometra 315 “ГУ = hy tag iu) ae Ат ру. ч ум а i Mile PF О и ух Pe i) * Ё А “Any 4 НАЯ eer * An A? as a fe yea ear