Classification

Phylum Nemata

Order Spirurida

Introduction

The representatives of the order Spirurida Chitwood, 1933 are nematode parasites of fishes, amphibians, reptiles, and mammals. They are some of the most common parasites found in vertebrates. Morphologically and phylogenetically diverse, the order includes 2 suborders, with 12 superfamilies with a large number of families, subfamilies, species, and subspecies that often inhabit a unique site of localization in the host, such as the esophagus, stomach, body cavities, blood vessels, and so on.

Recently their study has involved modern methods, such as scanning electron microscopy, molecular biology, and other techniques. As a result, new conceptions about their classification and complex life cycles are presented here.

Morphology and Locations within the Host

Spirurids are parasites having typical morphological features clearly distinguishing them from other nematodes. Their body is spindle-shaped. The front and back edges can be narrow or tapered. They possess an anterior extremity which is bilaterally symmetrical and they lack lateral, external labial papillae. In some members, the sexual dimorphism between males and females is very pronounced. Typically, the cuticle of the spiruridis has clear ornamentation. Their body surface may be transverse grooved, having different forms—spikes, teeth, edges, wart-like formations, wrinkles, and others. The mouth opening usually is surrounded by 2 lateral 3-section lips. In some cases, they possess additional dorsal and ventral lips. Some spirurids have a clearly-differentiated buccal cavity or stoma which leads into the pharynx, which can have different forms. The esophagus is divided into 2 parts: The anterior, which is muscular and shorter, and the posterior, which is glandular and longer. Males are without a genital bursa, but sometimes they have tail-cuticular wings. The caudal papillae are always ventral or ventrolateral in position. There is no pre-anal sucker. There are usually 2 spicules, different in shape and length from one another. Normally the right one is shorter and wider. The female genital opening is variable in its distance from the anterior end depending on the species (see Figures 1 and 2).

Figure 1. Whole body view of a species of Pterygodermatites from a bat. Showing: The mouth is at the anterior end (A), a double row of spines runs the length of body terminating just before the tail (B), and the tail (C).
(Source: K. Cajiao Mora, 2022. License: CC BY.)

Figure 2. Anterior end of a species of spirurid Pterygodermatites from a bat of the genus Myotis collected in eastern Colombia. Small hooks of the cuticle can be seen around the mouth and thin spinose parts of the cuticle can be seen running posteriad. In this species, the spines occur down the body and terminate before the tail.
(Source: K. Cajiao Mora, 2022. License: CC BY.)

Females lay eggs (Figure 3) with juveniles already formed. The juvenile stages, which are preinfective in the final or definitive host, develop entirely within an intermediate host, which may be crustaceans, beetles, coprophages, and other insects. Adults are parasites in the host’s gastrointestinal tract, nasal cavity, blood vessels, eyes, and conjunctival sacs, under the skin, and in different tissues or body cavities of fishes, birds, and mammals.

Figure 3. Spirurid egg from a short-eared owl Asio flammeus.
(Source: T. Pennycott, Edinburgh DataShare. License: CC BY.)

Overview of the Superfamilies

See Table 1 for an aggregated, selective implementation of the higher level taxonomy for this group with hosts and sites of localization noted (see also Kanchev et al., 2016; Vassilev et al., 1986).

Table 1. Selected Superfamilies of Nematodes with Hosts and Sites of Localization

Superfamily Acuarioidea

Acuarioidea tend to inhabit the upper alimentary tract or the muscles of the gizzard in birds, or may occur in the stomach of mammals, and are sometimes pathogenic. According to Anderson (2000), most adult acuarioids occur in the gizzard of birds, while a few species are found in both the proventriculus and in the posterior half of the esophagus. Most acuarioids occur in birds living in aquatic habitats and relatively few in birds associated with terrestrial habitats.

Acuarioids can move from one attachment site to another, leaving behind lesions devoid of worms. Cram (1931) reported openings through the gizzard lining associated with Acuaria hamulosa. Alicata (1938) found adult A. hamulosa mainly in tissues of the gizzard at its junction with the intestine. All acuarioids produce oval, smooth, thick-shelled eggs, each of which contains a small first-stage juvenile (J₁).

Hamann (1893) gave an account of how transmission probably occurs. Piana (1897) showed that Dispharynx nasuta of the proventriculus of gallinaceous and passerine birds developed in terrestrial isopods. Cram (1931; 1934), Cuvillier (1934), and Alicata (1938) conducted experiments to investigate the transmission and development of some acuarioids in terrestrial hosts and showed the importance of various insects and isopods as intermediate hosts. Garkavi (1956) investigated the development and transmission of Streptocara crassicauda.

Species of acuarioids in terrestrial hosts develop successfully in a great variety of arthropod intermediate hosts, including isopods, grasshoppers, beetles, and even diplopods. Acuarioids which parasitize aquatic hosts develop to the third stage in the hemocoel of aquatic crustaceans or in amphipods. Third-stage juveniles (J3) vary considerably in morphology especially in the posterior quarter or fifth of the body, which is bent dorsally, and the tail is always armed with spines or tubercles. In other species the tail is unarmed and generally conical. Infections come after ingesting arthropods containing infective third-stage juveniles (J₃). In piscivorous birds, such as cormorants, transmission depends on frog and fish paratenic hosts (Anderson, 2000).

Superfamily Aproctoidea

Aproctoidea include parasites in air sacs, nasal cavities, subcutaneous tissues of the head and neck, and orbits of birds (Anderson and Bain, 1976). Their eggs are thick-shelled and include a fully developed first-stage juvenile (J1). Little is known about the transmission of any of the species in the Aproctoidea (Anderson, 2000).

Superfamily Camallanoidea

Camallanoidea are parasites of the stomach and intestines of lower predaceous vertebrates (Chabaud, 1975) (see the chapter by Choudhury for a more in-depth summary of the Camallanoidea). Some of them occur in amphibians and reptiles, especially turtles (Baker, 1987), but also in marine, estuarine, and freshwater fishes (Ivashkin et al., 1971). One of the first demonstrations of heteroxeny in the Nemata was by Metchnikoff (1866) and Leuckart (1876) concerning the development of Camallanus lacustris of European fishes in copepods. Camallanoids are viviparous nematodes. Their intermediate hosts are crustaceans (Kupriyanova, 1954). Juveniles enter the haemocoel and develop into the third infective stage and they are armed with a few terminal spines. In the paratenic hosts (planktonivorous fishes) juveniles may grow to the fourth stage or become encapsulated in the tissues. Jackson and Tinsley (1998) found juveniles in aquatic toads (Pipidae) in Africa. The paratenic hosts move the juveniles in the food chain. The predator (piscivorous) definitive host can become infected by ingesting copepods or paratenic hosts with juveniles. Linstow (1909) described the juveniles of Camallanus lacustris in the isopod Asellus aquaticus and Fusco (1980) reported that some juveniles of Spirocamallanus cricotus developed successfully in white shrimp (Penaeus setiferus). Invasion of predaceous vertebrates is carried out after eating of intermediate or paratenic hosts which contain infective juveniles of camallanids (Anderson, 2000).

Superfamily Diplotriaenoidea

Diplotriaenoidea include spirurids in the air sacs of reptiles and birds (Anderson and Bain, 1976). Chabaud (1955) noted that eggs passed through the respiratory system and out via the feces. Anderson (1957) confirmed these observations experimentally. The female produces oval, smooth, thick-shelled eggs containing a fully developed first-stage juvenile (J1). Eggs hatch in the gut of their intermediate hosts (grasshoppers and locusts). Their anterior end is surrounded by rows of spines and the tail tip is rounded and also encircled by a row of spines.

Superfamily Dracunculoidea

Dracunculoidea consist of Spiruridae species that occur in tissues and serous cavities mainly of fishes, reptiles, birds, mammals (Chabaud, 1975), and rarely amphibians (Petter and Planelles, 1986) (see the chapter by Choudhury for a more in-depth summary of the Dracunculoidea). According to Anderson (2000), after insemination, the female grows large numbers of first-stage juveniles (J1). They must be dispersed into the environment where an available copepod may be colonized; these serve as intermediate hosts. In many species, the fully gravid female must be immersed in fresh water, which causes her to burst, thus releasing the juveniles into the environment. The female elicits a skin lesion or migrates into the rectum and protrudes from the body of the host. In some species, juveniles released within the host make their way to the tissues, including the blood. Most dracunculoids occur in hosts which have contact with fresh water.

Superfamily Filarioidea

Filarioidea contain parasites of the tissues and tissue spaces of all classes of vertebrates other than fishes (Anderson and Bain, 1976) (see the chapter by Notarnicola for a more in-depth summary of the Filarioidea). They are all transmitted by haematophagous arthropods. Members of the Filariidae family cause skin lesions and release eggs and/or juveniles in the host. They attract arthropod vectors, mainly individuals in the Muscidae family. The cephalic structures are rather simple. Pseudolabia are absent, but in some groups there may be cuticular elevations or spines. The cephalic papillae are well developed. The buccal cavity usually is considerably reduced. Spicules are variable in length and dissimilar in morphology (Anderson, 2000). Anderson (1957) suggests that the specialized life cycles of onchocercids evolved from those of the orbit-inhabiting Thelazia and the subcutaneous filariids (Filaria and Parafilaria). Other authors have suggested a relationship between some onchocercids and habronematoids like Draschia and Habronema (Chandler et al., 1941; Bain, 1981).

Superfamily Gnathostomatoidea

Gnathostomatoidea constitute spirurid nematodes characterized by massive, complex pseudolabia, and often spinous cephalic inflations (Chabaud, 1975). They are parasites in gastric mucosa of turtles in eastern North America (Hedrick, 1935). Some species of Gnathostoma have been well studied because of their significance to human and animal health. Members of the Gnathostomatoidea separate eggs in an undeveloped state, embryonate to second-stage juveniles (J2), and hatch in water. Intermediate hosts are copepods or insects of various crustaceans other than copepods (Anderson, 2000).

Superfamily Habronematoidea

Habronematoidea are nematodes with typical head structures. The pseudolabia are not large and median lips are present (Chabaud, 1975). It includes economically important and well-studied groups such as the tetramerids (including Tetrameres spp.) of the proventriculus of birds and noted for their peculiar sexual dimorphism, as well as the habronematids (including Habronema, Draschia, and Parabronema) which are transmitted by adult muscid dipterans to horses, certain ruminants, poultry, and other draft animals. They are localized in the stomach of horses and certain ruminants, including camels and elephants. Females, which occur in small tumors in the stomach wall, deposit oval, thin-shelled eggs. The latter usually hatch in the stomach releasing small, poorly differentiated juveniles with an anterior spine-like tooth. Juveniles pass out with the feces of the host. The superfamily also includes aberrant genera such as Hedruris (Anderson, 2000). In the United States, Ransom (1913) first discovered that the juveniles of horses developed in juveniles of muscid flies inhabiting nearby dung.

Superfamily Physalopteroidea

Physalopteroidea are parasites in the stomach and intestines of vertebrates. The mouth is encircled by large triangular lips having 1 or more teeth. A buccal capsule is absent. Males include a caudal alae. They usually meet ventrally in front of the cloaca and are supported by at least 4 papillae. The spicules are equal, subequal, or unequal. The female genital atrium is near then anterior or posterior half of body near the anus (Cheng, 1973).

Superfamily Rictularioidea

Rictularioidea consist of many species divided into several genera and subgenera (Quentin, 1969; Chabaud, 1975). They have no pseudolabia and have a denticulate, hexagonal oral opening and a sizeable buccal cavity with teeth. The presence of numerous large body spines is also diagnostic. The eggs are oval, with smooth, thick shells and each contains a fully developed first-stage juvenile (J₁). Eggs hatch in the gut of the insect intermediate host. These worms are parasites in the lumen of the intestine or firmly attached to the mucosa (ileum and in the region immediately posterior to its junction with the Malpighian tubules (Seureau, 1973). Witenberg (1928) fed young dogs the viscera of reptiles in which he had found rictularioid juveniles. The juvenile provokes the formation of a syncytium of epithelial cells which becomes surrounded by a fibrous capsule, which lies between the circular muscles and the epithelium of the ileum (Seureau, 1973).

Superfamily Spiruroidea

Spiruroidea include thelazioids, gnathostomatoids, habronematoids, rictularioids and physalopteroids (Chitwood and Chitwood, 1950). According to Chabaud (1975) the removal and elevation to superfamily status of several groups reduced Spiruroidea to 4 small families.

Spirurids are parasites in the stomach. They hatch thick-shelled eggs containing a fully differentiated first-stage juvenile (J₁) having a cephalic hook and rows of minute spines around the rather blunt anterior end. The tail of the first-stage juvenile (J₁) is often blunt and surrounded by a circlet of minute spines. Paratenesis is a common phenomenon in the transmission of spiruroids and the third-stage juveniles (J₃) of several species have been found in tissues of a variety of vertebrates which ingest infected insects, such as dung beetles. Third-stage juveniles (J₃) are generally large and possess some of the cephalic characteristics of adults. Their caudal extremities in some species possess terminal spines or tubercles, but in other species the terminal end is rounded and unornamented at the caudal extremity (Anderson, 2000).

Superfamily Thelazioidea

Thelazioidea consist of families united mainly on the basis of cephalic structures (Chabaud, 1975). The members of Thelazioidea are ovoviviparous and oviparous eyeworms of birds and mammals and the rhabdochonids (Rhabdochona) of fishes and primates. Intermediate hosts are different species of muscids.

Phylogenetics, Key Points

Within the order Spirurida, the superfamilies Habronematoidea and Thelazioidea are well established groups. Representatives of the families Cystidicolidae and Rhabdochonidae are widespread and show great diversity, especially in North America, but their phylogenetic relationships remain largely unexplored and the family Hedruridae appears to be an early branching line of the spirurids (Choudhury and Nadler, 2018).

Wu and colleagues (2008) explored the intra- and interspecific evolutionary variation among species of Camallanus collected from various fish species in China. Phylogenetic analyses of the nematodes suggested that there are 2 main clades, corresponding to different individuals of C. cotti and C. hypophthalmichthys from various fish species, although the interior nodes of each clade received poor support.

Černotíková and colleagues (2011) have worked out the phylogenetic relationships of 38 orders, including many families among the order Spirurida (namely, Camallanidae, Cystidicolidae, Daniconematidae, Philometridae, Physalopteridae, Rhabdochonidae, and Skrjabillanidae) and some among the order Ascaridida. The nematode species the authors examined are mostly parasites of marine and freshwater fishes from various locations in New Caledonia, as well as various locations in Africa, Asia, North America, South America, and Europe. Well supported trees allowing the study of phylogenetic relationships among some spirurine nematodes support the placement of Cucullanidae at the base of the suborder Spirurina, but the validity of the genera Afrophilometra and Caranginema is not supported. It is apparent that geographical isolation is not the cause of speciation in this parasite group and there is no evidence of coevolution with fish hosts (Černotíková et al., 2011).

Taxonomy

Following are details of taxonomy, type species, and distribution of representatives from the order that were proposed by Gibbons (2010). They differ somewhat from previous views, for instance, those presented in Chabaud (1975) and Anderson (2000), and Hodda’s (2022) more recent treatment. Though this chapter generally follows Gibbons (2010), the conflicting data in the literature indicate the necessity of taxonomic revisions among the order Spirurida.

Included are nominally narrative descriptions of the groups with their authority names. Some groups included here are covered additionally in other chapters within this book.

Suborder Camallanina Chitwood, 1936

The suborder Camallanina Chitwood, 1936 are parasites of the stomach and intestines of lower predaceous vertebrates (Chabaud, 1975). According to Gibbons (2010), Camallanina included the following superfamilies, families, subfamilies, genera, and subgenera, listed with some notes.

Superfamily Camallanoidea Travassos, 1920

Family Camallanidae

Subfamily Camallaninae Railliet and Henry, 1915

Type genus Camallanus Railliet and Henry, 1915 (= Zeylanema Yeh, 1960). These are parasites of fishes and amphibians (Chabaud, 1975), reptiles (Baker, 1987), and estuarine and freshwater fishes (Ivashkin et al., 1971).

Subgenus Zeylanema (Yeh, 1960) Moravec and Scholz, 1991

Type species is C. (Zeylanema) anabantis Pearse, 1933. These live in the intestine of the freshwater fish in the families Anabantidae, Cyprinidae, Belontiidae, and Clariidae from India.

Genus Neocamallanus Ali, 1957

Type species N. singhi Ali, 1957. These live in the intestines of Channa striata, Hampala dispar, and Xenetodon cancila from Laos.

Genus Neoparacamallanus Bilqees and Akram, 1982

Type species N. sweeti (Moorthy, 1937) Bilqees and Akram, 1982. These are parasites of freshwater fishes.

Subfamily Procamallaninae Yeh, 1960. Petter (1979) lists species in this subfamily.

Type genus Procamallanus Baylis, 1923

Type species Procamallanus laeviconchus (Wedl, 1862) Railliet and Henry, 1915. These live in the stomach and intestine of fishes and amphibians.

Species Procamallanus spiculogubernaculus. Sinha (1988) proposed a life cycle of P. spiculogubernaculus, which parasitizes fishes.

Subgenus Denticamallanus Moravec and Thatcher, 1997

Type species Procamallanus (Denticamallanus) dentatus Moravec and Thatcher, 1997. These live in the intestine of characid fish Bryconops alhurnoides from the Uburu River, Amazonas State, Brazil

Subgenus Isospiculus Ali, 1957

Subgenus Spirocamallanus Olsen, 1952

Species Procamallanus (Spirocamallanus) hilarii Vaz & Pereira. These live in the intestines of Acestrorhynchus microlepis (unspecified), Astyanax bimaculatus (adult), As. fasciatus (adult), As. parahybae (adult), Hoplias lacerdae (adult), H. malabaricus (adult), Oligosarcus macrolepis (adult), Rhamdia quelen (adult), Salminus hilarii (adult), Steindachnerina elegans (adult and juvenile), and Trichomycterus piurae (unspecified), all from Brazil (Luque et al., 2011).

Species Procamallanus (Spirocamallanus) fulvidraconis. Moraveč and colleagues (2003) redescribed P. (S.) fulvidraconis from central China.

Subgenus Monospiculus Ali, 1957 has no designated type species

Species ?Procamallanus (Monospiculus) parasiluri Fujita, 1927

Subgenus Procamallanus (Baylis, 1923 genus) Ali, 1957. This does not yet have a type species proven.

Species ?Procamallanus (Procamallanus) laeviconchus Baylis, 1923. These are common parasites of African freshwater fishes.

Subgenus Punctocamallanus Moravec and Scholz, 1991

Type species Procamallanus (Punctocamallanus) punctatus Moravec and Scholz, 1991. These are parasites that live in the stomach of freshwater fishes in Laos.

Subgenus Spirocamallanoides Moravec and Sey, 1988

Type species Procamallanus (Spirocamallanoides) siluri Osmanov, 1964

Subgenus Spirocamallanus (Olsen, 1952 genus) Moravec and Sey, 1988

Type species Procamallanus (Spirocamallanus) spiralis (Baylis, 1923). These are parasites that live in the intestine of fishes and amphibians.

Genus Batrachocamallanus Jackson and Tinsley, 1995

Type species Batrachocamallanus xenopodis (Baylis, 1929) Jackson and Tinsley, 1995. These are parasites of African amphibians.

Genus Xenopus (Jackson and Tinsley, 1995)

Genus Malayocamallanus Jothy and Fernando, 1970

Type species Malayocamallanus intermedius Jothy and Fernando, 1970. These are parasites in Fluta alba in Malaysia (Jothy and Fernando, 1970).

Genus Onchocamallanus Petter, 1979

Type species Onchocamallanus bagarii (Karve and Naik, 1951) Petter, 1979. These live in the intestine of Bagarius bagarius with an intermediate host of cyclopoid copepods Mesocyclops leuckarti and M. crassus (De and Maity, 1999) in India.

Genus Platocamallanus Bilqees and Akram, 1982

Type species Platocamallanus mehrii (Agrawal, 1930) Bilqees and Akram, 1982. These are parasites of freshwater fishes.

Subfamily Paracamallaninae Stromberg and Crites, 1974

Type genus Paracamallanus Yorke and Maplestone, 1928

Subgenus Dentocamallanus Moravec and Scholz, 1991

Type species Paracamallanus (Dentocamallanus) sweeti (Moorthy, 1937), These live in teeth on the ribs of the buccal capsule of fishes.

Superfamily Dracunculoidea

Representatives of this superfamily are parasites in tissues and serous cavities mainly of fishes, reptiles, birds, mammals (Chabaud, 1975), and sometimes in amphibians (Petter and Planelles, 1986). Intermediate hosts are copepods (Anderson, 2000).

Genus Lockenloia Adamson and Caira, 1991. This genus is not assigned to a family. It consists of parasites that live in the heart of the shark species Ginglymostoma cirratum.

Type species Lockenloia sanguineus Adamson and Caira, 1991

Family Dracunculidae (Stiles, 1907 subfamily) Leiper, 1912. These are parasites of reptiles, birds, and mammals.

Genus Fuellebornius Leiper, 1926

Type species Fuellebornius medinensis (Linnaeus, 1758) Leiper, 1926. These are parasites of humans Homo sapiens affecting the subcutaneous connective tissues, which then move to the surface of the skin and provoke the formation of a blister which bursts causing the anterior end of the worm to be exposed.

Family Anguillicolidae Yamaguti, 1935, These live in the swimbladder of eels (Laetsch et al., 2012).

Genus Anguillicola Yamaguti, 1935

Subgenus Anguillicola (Yamaguti, 1935) Moravec and Taraschewski, 1988

Type species Anguillicola (Anguillicola) globiceps Yamaguti, 1935. The definitive hosts are eels of the genus Anguilla and the intermediate hosts are planktonic copepods.

Genus Anguillicoloides Moravec and Taraschewski, 1988 (Moravec and Taraschewski, 1988)

Type species Anguillicoloides crassus (Kuwahara et al., 1974) Moravec and Taraschewski, 1988. These are parasites of the swimbladder of eels.

Family Skrjabillanidae Shigin and Shigina, 1958. These generally live in the peritoneal cavity of freshwater fishes. The occurrence of these nematodes in their final and intermediate host (Argulus foliaceus) in Hungary has been observed by Molnár and Szekely (1998).

Subfamily Skrjabillaninae (Shigin and Shigina, 1958 family) Chabaud, 1965

Genus Kalmanmolnaria Sokolov, 2006 (= Molnaria Moravec, 1968). These are parasites in the subcutaneous tissues of freshwater fishes (Scardinius erythrophthalmus) from Lake Balaton, Kis-Balaton, Fish Farms in Hungary. According to Anderson (2000), parasites of this genus can be found also in the serosa of the swimbladder, kidneys, and intestine, as well as on the mesentery of S. erythrophthalmus in the Commonwelath of Independent States. Intermediate hosts are crustaceans.

Genus Sinoichthyonema Wu, 1965

Type species Sinoichthyonema amuri (Garkavi, 1972) Moravec, 1982. According to Zhokhov and Molodozhnikova (2008), S. amuri have been introduced into the Volga Basin (Russia) occasionally during the process of introduction of fishes from the Amur River. This species also has been registered in Hungary by Molnár (1989).

Species Sinoichthyonema itenopharyngodoni Wu, 1973. The systematic status of this species was put forward by Moraveč (1982) including determining that this species is identical to S. amuri. Its definitive hosts are Rutilus rutilus and Scardinius erythrophthalmus.

Genus Garkavillanus Lomakin and Chernova, 1980

Type species Garkavillanus amuri (Garkavi, 1972) Lomakin and Chernova, 1980

Subfamily Esocineminae Moravec, 2006

Type genus Esocinema Moravec, 1977 (monotypic)

Type species Esocinema bohemicum Moravec, 1977. These live under the serosa of the air bladder of the pike Esox lucius in North Bohemia, Czechia.

Family Guyanemidae Petter, 1975. These include parasites that live in the peritoneal cavity and tissues of fish.

Genus Pseudodelphis Adamson and Roth, 1990

Type species Pseudodelphis oligocotti Adamson and Roth, 1990. These live in the peritoneal cavity and mesenteries surrounding the intestine of a marine fish species Oligocottus maculosus in coastal waters of British Columbia, Canada.

Genus Histodytes Aragort et al., 2002

Type species Histodytes microocellatus Aragort et al., 2002. These live in the gill, heart, kidney, spleen, and gonad tissues of the elasmobranch Raja microocellata. They were described based on material obtained from specimens from the continental shelf of the estuary of Muros y Noia, Spain (off the northwestern costs of the Iberian Peninsula) and are the only guyanemid genus described since the first was found on the European Atlantic coast (Aragort et al., 2002).

Genus Moravecia Ribu and Lester, 2004. These live in the gill filaments of green porcupine fish or may be found in the blood vessels and body cavity. This genus was described based on materials obtained from Tragulichthys jaculiferus found in Moreton Bay, Queensland, Australia (Ribu and Lester, 2004).

Type species Moravecia australiensis Ribu and Lester, 2004

Species Moravecia argentinensis. These were described by Braicovich and colleagues (2007) and are found in the blood vessels and body cavity of the Brazilian flathead Percophis brasiliensis. This is the first species of the genus reported from South American waters.

Subfamily Travassosneminae Moravec, 2006

Genus Travassosnema De Araujo Costa et al., 1991

Type species Travassosnema travassosi De Araujo Costa et al., 1991. These are viviparous parasites that live in tissues behind the eyes of Acestrorhynchus lacustris which are found in the Tres Marias Reservoir, Mina Gerais State, Brazil.

Subspecies Travassosnema travassosi paranaensis. These live in the body cavity of the characid fish Acestrorhynchus lacustris from the Paraná River near Guaira in southern Brazil (Moraveč et al., 1993; Silva-Souza and Saraiva, 2002).

Family Philometridae Baylis and Daubney, 1926

Genus Afrophilometra Moravec et al., 2009. This genus includes some species which parasitize Hydrocynus forskahlii from Lake Turkana, northwestern Kenya (Moraveč et al., 2009).

Type species Afrophilometra hydrocyoni (Fahmy et al., 1976) Moravec et al., 2009

Subfamily Philometrinae (Baylis and Daubney, 1926)

Genus Ichthyonema Diesing, 1861 (syn. Philometra Costa, 1845)

The type species is not clearly determined, but it may be Ichthyonema fuscum Diesing, 1861. These are parasites of the body cavity of marine fish.

Genus Paraphilometroides Moravec and Shaharom-Harrison, 1989

Type species Paraphilometroides nemipteri Moravec and Shaharom-Harrison, 1989. These are found in the dorsal fin and operculum of the marine perciform fish Nemipterus peronii from the coastal waters off Kuala Terengganu, Malaysia. Moraveč (2010) has imaged a gravid female of a paratype specimen of P. nemipteri using scanning electron microscopy after which he observed a unique cephalic structure, which clearly distinguishes Paraphilometroides from other philometrids.

Genus Margolisianum Blaylock and Overstreet, 1999

Type species Margolisianum bulbosum Blaylock and Overstreet, 1999. These are found in the southern flounder Paralichthys lethostigma from Ocean Springs, Mississippi Sound, Mississippi and Galveston Bay, Texas, United States. During their maturation, these spirurids have a different localization. Immature females are parasites of the eye, while mature and gravid females can be found in the subcutaneous tissues of the mouth and head, and males may be found in the muscle adjacent to the dorsal fin just posterior to the head (Gibbons, 2010).

Genus Dentiphilometra Moravec and Gui Tang Wang, 2002

Type species Dentiphilometra monopteri Moravec and Gui Tang Wang, 2002. These are found in the abdominal cavity of the ricefield eel Monopterus albus from Hubei Province in central China. This is the second philometrid species recorded from fishes of the order Synbranchiformes (Moraveč and Wang, 2002).

Genus Caranginema Moravec et al., 2008

Type species Caranginema americanum Moravec et al., 2008. These are found in the subcutaneous tissue of the crevalle jack Caranx hippos from southern Gulf of Mexico. This is the seventh species of Philometrinae recorded from marine and brackish water fishes in Mexico (Moraveč et al., 2008).

Subfamily Alineminae Moravec, 2006

Type genus Alinema Rasheed, 1963 (Gibbons, 2010)

Subfamily Neophilometroidinae Moravec et al., 2002

Type genus Neophilometroides Moravec et al., 2002

Type species Neophilometroides caudatus (Moravec et al., 1995) Moravec et al., 2002. These live in the swimbladder of Neotropical freshwater catfish and the pimelodid catfish Rhamdia guatemalensis from the Papaloapan River in Tlacotalpan, State of Veracruz, Mexico (Moraveč et al., 2002).

Subfamily Phlyctainophorinae (Roman, 1965) (Gibbons, 2010)

Genus Phlyctainophora Steiner, 1921

Type species Phlyctainophora lamnae Steiner, 1921. These live in the subcutaneous tissue of Lamna nasus from the North Atlantic Ocean. Jones and Delahunt (1995) found the same species in tumor-like lesions on the tail fin and which provoke a chronic inflammatory response in the host, the dogfish Squalus acanthias. This is the first record for a member of the genus established in New Zealand and the first record of Phlyctainophora adults from the Southern Hemisphere.

Species Phlyctainophora squali. This species was obtained from Squalls acanthias in eastern Pacific Ocean off Los Angeles, California, United States, at a depth of 200 m (Dwight and Murrady, 1969).

Family Micropleuridae (Baylis and Daubney, 1926) Travassos, 1960

Subfamily Micropleurinae Baylis and Daubney, 1926. These are parasites found in fish, amphibians, and reptiles.

Genus Protenema Petter and Planelles, 1986

Type species Protenema longispicula Petter and Planelles, 1986. These are found in the amphibians Necturus maculosus (family Proteidae) found in the lakes of Minnesota, United States.

Genus Granulinema Moravec and Little, 1988

Type species Granulinema carcharini Moravec and Little, 1988. These are parasites of the bull shark Carcharhinus leucas found in Lake Borgne, Louisiana, United States. The site of localization in the host is unknown (probably the abdominal cavity).

Species Granulinema simile Moravec and Little, 1988. The site of localization in the host is also unknown (Moraveč and Little, 1988).

Genus Kamegainema Hasegawa et al., 2000

Type species Kamegainema cingulum (Linstow, 1902) Hasegawa et al., 2000. These live in the subcutaneous tissue of amphibians.

Family Daniconematidae Moravec and Køie, 1987. This family includes viviparous parasites of fish.

Type genus Daniconema Moravec and Køie, 1987

Type species Daniconema anguillae Moravec and Køie, 1987. These live under the serosa of the swimbladder and intestine of eels Anguilla anguilla found in Lake Esrum, northern Zealand, Denmark. A new family Daniconematidae was established to accommodate it (Moraveč and Køie, 1987).

Genus Mexiconema Moravec et al., 1992

Type species Mexiconema cichlasomae Moravec et al., 1992. These live in the abdominal cavity or viscera, or (rarely) the skin of species cichlids of the genus Cichlasoma in the coastal lagoons of Celestun, North Yucatán, Mexico. They live in the mesentery, swimbladder, liver, spleen, kidney, intestinal lumen, serosal cover of the intestine, or (rarely) in the skin of their hosts, such as C. helleri, C. motaguense, and C. pearsei (Moravec et al., 1992). Other habitats in Campeche, Mexico are El Vapor (a freshwater lagoon adjacent to Terminos Lagoon), Palizada, Santa Gertrudis, El Cayo (a saltwater portion within Terminos Lagoon), Pargos, and Rio Champoton. Habitats in Quintana Roo, Mexico include Rio Lagartos (a coastal lagoon) and Noh Век (a lake). El Vapor, Palizada, Santa Gertrudis, Rio Lagartos, and Noh Век are truly freshwater localities; all the remaining sites are saltwater or marine localities (Moraveč et al., 1992).

Genus Syngnathinema Moravec et al., 2001

Type species Syngnathinema californiense Moravec et al., 2001. These live in the vascular system of the Bay pipefish Syngnathus leptorhynchus in California, United States. Based on histological studies, they have been found also in the circulatory system including the sinus venosus, atrium, and renal and hepatic veins.

Family Lucionematidae Moravec et al., 1998. These are viviparous parasites of fish.

Type genus Lucionema Moravec et al., 1998

Type species Lucionema balatonense Moravec et al., 1998. These live in the swimbladder of the European pikeperch Stizostedion lucioperca from Lake Balaton, Hungary (Moraveč et al., 1998).

Suborder Spirurina

Moraveč (2007) reviewed the spirurines of fish belonging to approximately 300 species in 4 superfamilies, namely Gnathostomatoidea, Habronematoidea, Physalopteroidea, and Thelazioidea. He has suggested that the classification and taxonomy of species of this suborder in fish requires reevaluation using new techniques, such as scanning electron microscopy and molecular biology (Moraveč, 2007).

Superfamily Acuarioidea

Family Acuariidae (Railliet et al., 1912)

Subfamily Acuariinae Railliet et al., 1912

Genus Antechiniella Quentin and Beveridge, 1986. These live in Australian marsupials.

Type species Antechiniella suffodiax (Beveridge and Barker, 1975) Quentin and Beveridge, 1986

Genus Chandleronema Little and Ali, 1980. These live in the stomach of raccoons Procyon lotor and muskrats from the United States.

Type species Chandleronema longigutturata (Chandler, 1942) Little and Ali, 1980

Genus Cordonema Schmidt and Kuntz, 1972. These are parasites of birds.

Type species Cordonema venusta Schmidt and Kuntz, 1972

Genus Molinacuaria Wong and Lankester, 1985. These live under the gizzard lining of birds of the species Dendragapus obscurus fuliginosus, Gallinula chloropus indica, and Alcippe brunnea brunnea from Vancouver Island, Canada; China; and Taiwan, respectively.

Type species Molinacuaria bendelli (Adams and Gibson, 1969) Wong and Lankester, 1985

Genus Syncuaria Gilbert, 1927. These live in the gizzard of grebes, storks, and cormorants.

Type species Syncuaria ciconiae Gilbert, 1927

Genus Tikusnema Hasegawa et al., 1992. These live in the stomach and small intestine of the ricefield rat Rattus argentiventer in Indonesia.

Type species Tikusnema javaense Hasegawa et al., 1992

Genus Voguracuaria Wong and Anderson, 1993. These live in the esophagus of the whimbrel Numenius phaeopus phaeopus in Vogur, Iceland.

Type species Voguracuaria lankesteri Wong and Anderson, 1993

Genus Willmottia Mawson, 1982. These are parasites of birds, Malurus cyaneus, from Tasmania.

Type species Willmottia australis Mawson, 1982

Genus Xenocordott Mawson, 1982. These live in the gizzard of Australian birds, Phylidonyris novaehollandiae and Gymnorhina tibicen.

Type species Xenocordott patonae Mawson, 1982

Subfamily Schistorophinae

Genus Deliria Vicente et al., 1980. These live in the stomach of birds, Pitangus sulphuratus, found in Rio de Janeiro State, Brazil.

Type species Deliria gomesae Vincente et al., 1980

Genus Paracuaria Rao, 1951. These live in the submucosa of crop in seabirds or the stomach of insectivorous mammals.

Type species Paracuaria adutica (Creplin, 1946)

Genus Pseudoaviculariella Gupta and Kazim, 1978. These live in the gizzard of the cattle egret Egretta garzetta from Lucknow, India.

Type species Pseudoaviculariella srivastavai Gupta and Kazim, 1978

Subfamily Schistorophinae Travassos, 1918

Genus Quasithelazia Maplestone, 1932

Genus Schistogendra Chabaud and Rousselot, 1956

Genus Sobolevicephalus Parukhin, 1964. These live under the gizzard of birds. This genus was listed by Anderson and colleagues (2009) as synonym of Hadjelia Seurat, 1916.

Type species Sobolevicephalus chalycyonis Parukhin, 1964

Superfamily Filarioidea

Genus Avifilaris Saunders, 1955. These live in the blood of Rhodothraupis, Passerina, Pitangus, and Empidonax.

Type species Avifilaris fringillidarum Saunders, 1955

Family Filariidae (Weinland, 1858) Cobbold, 1879. This represents a collective group for agamic forms named “Agamofilaria” Stiles, 1907.

Subfamily Filariinae Weinland, 1858

Genus Cystofilaria Skrjabin and Shikhobalova, 1948. These are parasites of which the adults may be found in cysts under the muscular layer of the esophagus in dogs.

Type species is C. balkanica Skrjabin and Schikhobalova, 1948

Genus Paracanthocheilonema Vladimirov, 1959 in Buliginskaya et al., 1959. These are parasites in Rhombomys opimus, Meriones meridianus, and M. erytbrourus found in the Kashkadarinsk region of Uzbekistan.

Type species Paracanthocheilonema vite (Krepkogorskaya, 1933) Vladimirov, 1959 in Buliginskaya et al., 1959

Family Onchocercidae (Leiper, 1911)

Subfamily Onchocercinae Leiper, 1911

Genus Bisbalia Bain and Guerrero, 2003. These are are parasites found in the membranous pocket in the pleural cavity of Heteromys anomalus (order Rodentia: Geomyoidea) in northern Venezuela.

Type species Bisbalia vossi Bain and Guerrero, 2003

Genus Cherylia Bain et al., 1985. These are parasites of the ventral subcutaneous and perimuscular tissues of the South American marsupial Metachirops opossum found in French Guiana.

Type species Cherylia guyanensis Bain et al., 1985

Subgenus Cercopithifilaria (Eberhard, 1980). These are parasites transmitted by ticks and are found in primates, ruminants, carnivores, marsupials, and monotremes.

Type species Cercopithifilaria kenyensis Eberhard, 1980

Genus Chabfilaria Bain et al., 1983. These are parasites of Xenarthra found in French Guiana and Guyana.

Type species Chabfilaria jonathani Bain et al., 1983

Genus Cruorifilaria Eberhard et al., 1976. These are parasites that live in the renal and pulmonary blood vessels, and (rarely) the coronary vessels of the capybara Hydrochoerus hydrochaeris in Colombia.

Type species Cruorifilaria tuberocauda Eberhard et al., 1976

Subgenus Dasypafilaria (Eberhard, 1982). These are parasites that live in the omentum of the family Dasypodidae (including the 9-banded armadillo Dasypus novemcinctus) found in southern Louisiana, United States.

Type species Dasypafilaria averyi Eberhard, 1982

Genus Josefilaria Moorhouse et al., 1979. These are parasites of the ghost bat Macroderma gigas found in Australia.

Type species Josefilaria mackerrasae Moorhouse et al., 1979

Genus Loxodontofilaria Berghe and Gillain, 1939. These are parasites of elephants in Africa and Burma, Caprinae and Bovidae in Japan, and hippopotamus in Africa.

Type species Loxodontofilaria loxodontis Berghe and Gillain, 1939

Genus Mansonella Faust, 1929. These are parasites that develop in the subcutaneous tissues of their hosts and may be found in the Caribbean region, Central America, South America, and Africa.

Type species Mansonella ozzardi (Manson, 1897) Faust, 1929

Subgenus Cutifilaria (Bain and Schulz-Key, 1974 genus) Uni et al., 2004. These are parasites in Cervidae in Europe and Japan.

Type species Mansonella (Cutifilaria) wenki (Bain and Schulz-Key, 1974)

Subgenus Esslingeria (Chabaud and Bain, 1976) Eberhard and Orihel, 1984. These are parasites of humans Homo spaiens, African anthropoid apes, and South American rodents.

Type species Mansonella (Esslingeria) perstans (Manson, 1891) Eberhard and Orihel, 1984

Subgenus Mansonella (Faust, 1929) Eberhard and Orihel, 1984. These are parasites of humans Homo sapiens, rodents, and carnivores.

Type species Mansonella (Mansonella) ozzardi (Manson, 1897) Faust, 1929

Subgenus Sandnema (Chabaud and Bain, 1976) Eberhard and Orihel, 1984. These are parasites of Asian primates and insectivores.

Type species Mansonella (Sandnema) digitata (Chandler, 1929) Eberhard and Orihel, 1984

Subgenus Tetrapetalonema (Faust, 1935) Eberhard and Orihel, 1984. These are parasites of platyrrhine primates.

Type species Mansonella (Tetrapetalonema) marmosetae (Faust, 1935) Eberhard and Orihel, 1984

Subgenus Tupainema Eberhard and Orihel, 1984. These are parasites of tree shrews in Southeast Asia.

Type species Mansonella (Tupainema) dunni (Mullin and Orihel, 1972) Eberhard and Orihel, 1984

Genus Molossinema Georgi et al., 1987. These are parasites of the cerebral ventricular system of the bat Molossus ater in Trinidad.

Type species Molossinema wimsatti Georgi et al., 1987

Genus Strianema Eberhard et al., 1993. These are parasites that live in the subcutaneous tissues of Venezuelan armadillos Dasypus species.

Type species Strianema venezuelensis Eberhard et al., 1993

Genus Struthiofilaria Noda and Nagata, 1976. These are parasites which live in the body cavity of the ostrich Sruthio camelus found in in Misaki Park Zoo, Osaka Prefecture, Japan.

Type species Struthiofilaria megalocephala Noda and Nagata, 1976

Genus Yatesia Bain et al., 1982. These live in the skeletal muscle fascia of capybara Hydrochoerus hydrochaeris in Colombia.

Type species Yatesia hydrochoerus (Yates, 1980) Bain et al., 1982

Subfamily Waltonellinae Bain and Prod’hon, 1974

Genus Edesonfilaria Yeh, 1960

Species Edesonfilaria malayensis. These live in the subserosal connective tissues of the abdominal and thoracic cavities of cynomolgus monkeys Macaca fascicularis from Indonesia (Nonoyama et al., 1984).

Genus Foleyella Seurat, 1917. These live in the subcutaneous and intermuscular connective tissues and body cavities in chameleonid reptiles.

Type species Foleyella candezei (Fraipont, 1882) Seurat, 1917

Genus Foleyellides Caballero, 1935. These are parasites of anuran amphibians, mainly in the family Ranidae.

Type species Foleyellides striatus (Ochoterena and Caballero, 1932) Caballero, 1935

Genus Loaina Eberhard and Orihel, 1984. These are parasites of North American rabbits.

Type species Loaina uniformis (Price, 1957) Eberhard and Orihel, 1984.

Genus Ochoterenella Caballero, 1944. These live in the body cavity of anuran amphibians, mainly of the family Bufonidae in the Neotropics.

Type species Ochoterenelladigiticauda Caballero, 1944

Genus Paramadochotera Esslinger, 1986. These are parasites of Mantidactylus redimitus, a racophorid frog in Madagascar.

Type species Paramadochotera guibei (Bain and Prod’hon, 1974) Esslinger, 1986

Subfamily Dirofilariinae Sandground, 1921

Genus Pelecitus Railliet and Henry, 1910. These live in the tendons, muscles, and (rarely) wings of birds and mammals.

Type species Pelecitus helicinus (Molin, 1860)

Subfamily Splendidofilariinae Chabaud and Choquet, 1953

Genus Splendidofilaria Skrjabin, 1923

Subgenus Amfilaria Lopez Caballero and Jimenez Millan, 1979

Type species Splendidofilaria (Avifilaria) mavis (Leiper, 1909) Anderson, 1961

Subgenus Arteriofilaria Lopez Caballero and Jimenez Millan, 1979

Type species Splendidofilaria (Arteriofilaria) algonquinensis (Anderson, 1955) Anderson, 1961

Subgenus Soninella Lopez Caballero and Jimenez Millan, 1979

Type species Splendidofilaria (Soninella) verrucosa Oschmarin, 1950

Subgenus Splendidofilaria (Skrjabin, 1923 genus) Lopez Caballero and Jimenez Millan, 1979

Type species Splendidofilaria (Splendidofilaria) pawloski Skrjabin, 1923

Genus Andersonfilaria Bartlett and Bain, 1987. These live in the fossa of the dorsal wall of the pelvic girdle of the common waxbill Estrilda astrild (order Passeriformes) in Africa.

Type species Andersonfilaria africanus Bartlett and Bain, 1987

Genus Dessetfilaria Bartlett and Bain, 1987. These live in the capsule of the outer wall of the aorta in the heart of toucans in French Guiana and Brazil.

Type species Dessetfilaria guianensis Bartlett and Bain, 1987

Genus Rumenfilaria Lankester and Snider, 1982. These live in the subserosal connective tissue between the folds of the ruminal wall of moose Alces alces from northwestern Ontario, Canada.

Type species Rumenfilaria andersoni Lankester and Snider, 1982

Genus Serofilaria Wu and Yun, 1979 (in Wu et al., 1979). These live in the lymphatic vessels of the serous membrane covering the internal organs of pigs in China.

Type species Serofilaria suis Wu and Yun, 1979 (in Wu et al., 1979)

Genus Splendidofilarioides Texeira de Freitas and Nicanor Ibafiez, 1968. These are parasites of the birds Mimus longicaudatus in Peru.

Type species Splendidofilarioides pachacuteci Texeira de Freitas and Nicanor Ibanez, 1968

Genus Eulimdana Founikoff, 1934. These are parasites of birds.

Type species Eulimdana clava (Wedl, 1856)

Subfamily Lemdaninae Lopez-Neyra, 1956q

Genus Lemdana Seurat, 1917. These live in the subcutaneous connective tissue of the head, neck in the vicinity of the trachea, the esophagus, and crop of birds.

Type species Lemdana marthae Seurat, 1917

Genus Makifilaria Krishnasamy et al., 1981. These live in the peritoneal cavity of the island flying fox Pteropus hypomelanus found in Pulau Langkawi, Malaysia.

Type species Makifilaria inderi Krishnasamy et al., 1981.

Superfamily Aproctoidea

Family Aproctidae (Yorke and Maplestone, 1926 subfamily) Skrjabin and Shikhobalova, 1945 (monotypic)

Genus Hovorkonema Jurasek, 1977. These live in the stomach of the Carpathian wild boar Sus scrofa atilla in Lucenec, Slovakia.

Type species Hovorkonema gastrofilana Jurasek, 1977

Subfamily Aproctinae Yorke and Maplestone, 1926

Genus Desmidocercella Yorke and Maplestone, 1926

Type species Desmidocercella (Desmidocercella) numidica (Seurat, 1920)

Subgenus Skrjabinocercella Gushanskaya, 1953

Type species Desmidocercella (Skrjabinocercella) incognita Solonitzin, 1932

Genus Lissonema Linstow, 1903. These live in the abdominal cavity of Otus sunia from eastern Asia.

Type species Lissonema rotunda Linstow 1903

Genus Parasaurositus Gupta and Johri, 1989. These live in the intrahepatic spaces of the Indian soft shell turtle Aspideretes gangeticus in India.

Type species Parasaurositus yamagutii Gupta and Johri, 1989

Genus Pseudodiomedenema Gupta and Johri, 1988. These are parasites of the pleural cavity of hoopoe Upupa epops found in Lucknow, India.

Type species Pseudodiomedenema cameroni Gupta and Johri, 1988

Genus Squatnoftlaria Schmerling, 1925

Superfamily Diplotriaenoidea

Family Diplotriaenidae (Skjrabin, 1916 subfamily) Anderson, 1958

Subfamily Diplotriaeninae Skrjabin, 1916

Genus Spinodiplotriaena Kalyankar and Pallawadar, 1989. These live in the body cavity of the common mynah bird Acridotheres tristis in India.

Type species Spinodiplotriaena urmilii Kalyankar and Pallawadar, 1989

Genus Versternema Bain et al., 1992. These live in the body cavity of the ostrich Struthio camelus in Botswana.

Type species Vesternema struthionis Bain et al., 1992.

Superfamily Gnatohostomatoidea

Family Gnathostomatidae Railliet, 1895

Subfamily Ancyracanthinae Yorke and Maplestone, 1926

Genus Elaphocephalus Molin, 1860. These live in the feet of birds Psittacus macao.

Type species Elaphocephalus octocornutus Molin, 1860

Genus Metaleptus Machida et al., 1982

Species Metaleptus rabuka. These live in the stomach of Mustelus griseus in the north Pacific Ocean off Honshu, Japan (Moraveč and Nagasawa, 2000)..

Species Metaleptus manazo. These also have been recorded in the north Pacific Ocean off Honshu, Japan (Moraveč and Nagasawa, 2000).

Superfamily Habronematoidea

Family Habronematidae (Chitwood and Wehr, 1932 subfamily) Ivaschkin, 1961

Subfamily Habronematinae Chitwood and Wehr, 1932

Genus Dermofilaria Rivolta, 1884. These are parasites of equids and bovines.

Type species Dermofilaria irritans Rivolta, 1884

Subfamily Histiocephalinae Gendre, 1922

Genus Sobolevicephalus Parukhin, 1964

Type species Sobolevicephalus chalcyonis Parukhin, 1964

Family Tetrameridae Travassos, 1914

Subfamily Tetramerinae (Travassos, 1914)

Genus Acanthophorus von Linstow, 1876 (syn. Tetrameres)

Genus Ascarophis van Beneden, 1871. These live in the gastrointestinal tract of marine fish.

Type species Ascarophis morrhuae Beneden, 1871 (Gibbons, 2010). Intermediate hosts are decapods (Enalus gaimardi, Eupagurus pubescens, Hetairus polaris, Pagurus pubescens, Pandalus borealis, and Spirontocaris spinus) from the Bering Sea (Uspenskaya, 1953; 1954), lobster (Homarus americana) in North America (Uzmann, 1967), crab Carcinus maenas from off the coast of Brittany in France (Petter, 1970), crustaceans (Anisogammarus kygi, A. ochotensis, A. tiuschovi, Idothea ochotensis, and Pagurus middendorffii) from the littoral zone of Big Shantar Island in the Okhotsk Sea (Tsimbalyuk et al., 1970), shore crabs (Hemigrapsus oregonensis), porcelain crabs (Pachycheles rudis) in California, United States (Poinar and Kuris, 1975), and Callianassa californiensis, Pagurus samuelis, Pag. granosimanus, Pachycheles pubescens, and Pugettia producta (Poinar and Thomas, 1976).

Species Ascarophis mexicana. These were described by Moraveč and colleagues (1995) from the stomach of Epinephelus morio and E. adscensionis from the Gulf of Mexico and southeastern Mexico in the states of Yucatán and Veracruz. This is the second Ascarophis species known to parasitize fishes of the genus Epinephelus (Moravec et al., 1995).

Genus Caballeronema Margolis, 1977. These live in the alimentary canal of the marine fish, Scorpaenichthys marmoratus, found off the Pacific coast of Canada.

Type species Caballeronema wardlei (Smedley, 1934) Margolis, 1977

Genus Capillospirura Skrjabin, 1924. These are parasites of the digestive tract of sturgeons. Based on the characteristics of the cephalic structure of specimens from Old World sturgeons, Capillospirura Skrjabin, 1924 (phylum Nemata: family Cystidicolidae) has been redefined.

Type species Capillospirura ovotrichuria Skrjabin, 1924

Species C. ovotrichuria Skrjabin, 1924 (as determined by Appy and Dadswell, 1978).

Species C. argumentosa (Skrjabina, 1966) (= Ascarophis argumentosus). These are from Old World sturgeons (Appy and Dadswell, 1978).

Species C. pseudoargumentosa (= Caballeronema pseudoargumentosus). These are from a New World sturgeon (Appy and Anderson, 1982).

Genus Comephoronema Layman, 1933. These live in the alimentary tract of freshwater fish.

Type species Comephoronema werestschagini Layman, 1933

Species Comephoronema multipapillatum. These are from the anterior intestine and cecum of the squirrelfish, Holocentrus adscensionis (Pereira et al., 2014). This is the fifth nominal species of Comephoronema and the first nematode registered in H. adscensionis and the first species of the genus in the Neotropical part of the Atlantic Ocean (Pereira et al., 1993).

Genus Crenatobronema Solov’eva, 1987. These are parasites in fish from the Pacific Ocean. In his review concerning to the suborder Spirurina, Moraveč (2007) considers this genus “inadequately known.”

Type species Crenatobronema guentheri (Baylis, 1929) Solov’eva, 1987.

Genus Cystidicoloides Skinker, 1931 (syn. Sterliadochona Skrjabin, 1948. These are parasites of South American freshwater fish. Rasheed (1965) and Moraveč (1967) have synonymized Sterliadochona Skrjabin, 1946 with Cystidicoloides.

Type species Cystidicoloides fischeri (Travassos et al., 1928) Skinker, 1931. These were redescribed by Moraveč and colleagues (2008) noting the localization in the stomach of Pygocentrus piraya and Serrasalmus brandtii from Três Marias Reservoir, Upper São Francisco River, Minas Gerais state, Brazil. Based on morphological features, the authors (Moraveč et al., 2008) accomplished several taxonomic transformations, such as: Heliconema izecksohni Fabio, 1982 is transferred to Cystidicoloides as C. izecksohni (Fabio, 1982). Cystidicoloides uniseriata Valovaya and Valter, 1988 is considered a species inquirenda. It has been proposed as a newly erected genus, Salmonema, with the type species S. ephemeridarum. Cystidicoloides prevosti (Choquette, 1951) has been transferred to Salmonema as S. prevosti (Choquette, 1951). Sterliadochona savini Skryabin, 1948 and Sterliadochona Skryabin, 1948 are considered as species inquirenda and genus inquirenda, respectively.

Genus Echinurioides Thwaite, 1926. These are parasites of the spurwinged goose, Plectropterus spp., in northern Nigeria.

Type species Echinurioides plectropteri Thwaite, 1926 (Gibbons, 2010). Skrjabin and Sobolev (1963) list Echinurioides as a synonym of Tetrameres Creplin, 1846. Alexander and McLaughlin (1997) report the type species as Tetrameres plectropteri (Thwaite, 1926), with host Plectropterus gambensis in Nigeria.

Genus Gubernaculomeres Oshmarin and Parukhin, 1963. These are parasites that live in the proventriculus of Astur gentilis and Aquila clanga.

Type species Gubernaculomeres tubocloacis (Oshmarin, 1956) Oshmarin and Parukhin, 1963

Genus Moravecnema Justine et al., 2002 is considered to be a parasite of the deep sea hydrothermal fish Pachycara thermophilum from the Mid-Atlantic Ridge.

Type species Moravecnema segonzaci Justine et al., 2002. This is the first species of parasitic nematode described from a fish endemic to hydrothermal deep sea vents.

Genus Prospinitectus Petter, 1979. These live in the intestine of the fish Euthynnus affirtis off Kuala Lumpur, Malaysia and in the China Sea.

Type species Prospinitectus mollis (Mameav, 1968) Petter, 1979

Genus Pseudascarophis Ko et al., 1985. These live in stomach of the fish Kyphosus cinerascens from off the southeastern coast of Japan.

Type species Pseudascarophis kyphosi Ko et al., 1985

Species Pseudascarophis brasiliensis. These are found in the stomach of Kyphosus sectatrix from off Rio de Janeiro, southeastern Brazil (Pereira et al., 2013).

Genus Salmonema Moravec et al., 2008. These live in the digestive tract of freshwater fish.

Type species Salmonema ephemeridarum (Linstow, 1872) Moravec et al., 2008

Genus Similascarophis Munoz et al., 2004. These are parasites of the digestive tract of marine fish off the Chilean coast.

Type species Similascarophis maulensis Munoz et al., 2004

Genus Cristitectus Petter, 1970. Discovered in the mid-1990s.

Genus Salvelinema Trofimenko, 1962. Discovered in the mid-1990s.

Genus Ctenascarophis Mamaev, 1968 and 1967 Petter, 1969. Discovered in the mid-1990s.

Genus Tetrameres Creplin, 1846. This genus is remarkable for the fact that the mature female is almost spherical in shape, blood red in color, and lies embedded in the proventricular glands of birds.

Species Tetrameres americana Cram, 1927. These occur in the proventriculus of fowl and turkeys. The definitive hosts of T. americana are the grasshoppers Scyllina cyanipes in Puerto Rico and Melanoplus femurrubrum and M. differentialis in mainland United States, and have been recorded elsewhere from the United States and in South Africa. Intermediate hosts are M. femurrubrum, M. differentialis, and Blatella germanica.

Species Tetrameres fissispina (Diesing, 1861). These occur in the duck, pigeon, fowl, turkey, and wild aquatic birds, and has a wide distribution. Intermediate hosts are the water crustacean Daphnia pulex and Gammarus pulex.

Species Tetrameres crami Swales, 1933. These occur in domestic and wild ducks in North America. Their intermediate hosts are the amphipods Gammarus fasciatus and Hvalella knickerbockeri.

Species Tetrameres confusa Travassos, 1919. These occur in the proventriculus of fowl pigeon and other birds in Brazil. Their intermediate hosts are probably similar to those for T. fissispina.

Species Tetrameres mohtedai Bahlerao and Rao, 1944. These occur in fowl in India and Southeast Asia. Their intermediate hosts are cockroaches and grasshoppers, such as Spathosternum praszniferum and Oxya nitidula

Species Tetrameres pattersoni (Cram, 1933). These occur in quail. Their intermediate hosts are grasshoppers and cockroaches (Soulsby, 1982), Chortophaga viridifasciata and Melanoplus femurrubrum, and its definitive host is Colinus virginianus (Anderson, 2000).

Species Tetrameres cardinalis Quentin and Barre, 1976. These have been found in the northern cardinal (Cardinalis cardinalis (syn. Richmondia cardinalis)) in Mexico. Its development occurs in Locusta migratoria.

Species Tetrameres coccinea (Seurat, 1914) Travassos, 1914. These are from the Phoenicopterus ruber, Bubulcus ibis, and Platalea leucorodia Linnaeus, 1758 (Junker and Boomker, 2007).

Species Tetrameres lhuillieri (Seurat, 1918). These are found in Alectoris graeca (Meisner, 1804) and Columba oenas Linnaeus, 1758 from Algeria.

Species Tetrameres nouveli (Seurat, 1914) Travassos, 1914. These are found in the black winged stilt, Himantopus himantopus (Linnaeus, 1758) in Algeria and Nigeria.

Species Tetrameres plectropteri Thwaite 1926. These are found in Plectropterus gambensis.

Species Tetrameres paradisea Ortlepp, 1932. These have been recovered from Anthropoides paradisea (Lichtenstein, 1793) and have been described from South African hosts.

Species Tetrameres prozeskyi (Ortlepp, 1964). These have been described from South African hosts. They occur in Tockus erythrorhynchus and To. leucomelas.

Species Tetrameres numida Junker and Boomker, 2007. These occur in Numida meleagris from Musina (also known as Messina), Limpopo Province, South Africa.

Superfamily Physalopteroidea

Family Physalopteridae (Railliet, 1893 subfamily) Leiper, 1908. These are parasites of the alimentary canal (as well as the esophageal, gastric, or aortic walls) of the selachian Chlamydoselachus anguineus from the Pacific coast of central Honshu, Japan.

Type species Metaleptus rabuka Machida et al., 1982

Subfamily Physalopterinae Railliet, 1893

Genus Kreisiella Jones, 1985. These live in the stomach of the Australian lizard Egernia inornate.

Type species Kreisiella chrysocampa Jones, 1985. Goldberg and colleagues (2008) report finding Kreisiella chrysocampa in Emoia (family Scincidae) from Papua New Guinea.

Genus Leptosoma Travassos, 1920

Type species Leptosoma leptosoma (Gervais, 1848). Leptosoma leptosoma adults live in the stomach or intestine of mammals, birds, reptiles, and amphibians.

Genus Paraphysaloptera Gupta and Kazim, 1979. These are parasites of the gizzard lining of the hoopoe Upupa epops.

Type species Paraphysaloptera alii (Gupta and Kazim, 1978) Gupta and Kazim, 1979. These are found in the gizzard of the hoopoe Upupa epops (Gupta and Kazim, 1979; Martín-Vivaldi et al., 2014).

Species Paraphysaloptera indica. These are found in the intestine of the hoopoe Upupa epops (Gupta and Johri, 1985; Martín-Vivaldi et al., 2014).

Subgenus Chlamydonema (Hegt, 1910 genus) Gupta and Johri, 1987

Type species Physaloptera (Chlamydonema) praeputiale (Linstow, 1888) Travassos, 1917. These live in the stomach of Canis latrans, Felis catus domesticus, F. pardus, and Ca. familiaris and are found in Asia, Africa, Europe, North America, and South America.

Species Physaloptera praeputialis. Described by Linstow in 1888 from a wild cat (Felis catus) from Brazil. Later, Walton (1927) assigned this specimen to the group Mammal. This is probably the first record of P. praeputialis in North America. In the stomach of lynx (Lynx rufus texensis) and ocelot (Felis pardalis) from Mexico the same species has been reported also by Caballero y Caballero and Peregrina (1938).

Subgenus Physaloptera (Rudolphi, 1819 genus) Gupta and John, 1987

Type genus Physaloptera Rudolphi, 1819. Physaloptera are common nematodes found in the stomach and muscles of mammals (such as cats, dogs, and humans), reptiles, amphibians, and birds. Physalopterids attach to the walls of the duodenum and stomach (Naem and Asadi, 2013) and are known to have pathological consequences such as catarrhal gastritis, gastrointestinal upset, erosion of the mucosa, ulcers, and vomiting (Soulsby, 1965).

Figure 4. Views of Physaloptera rara (phylum Nemata: order Spirurida: family Physalopteridae). A) Anterior end of P. rara from a domestic dog obtained from Iowa, United States. Note the 2 large lips each with 3 small anteriorly-directed teeth (which is typical of Physaloptera spp.); B) lateroventral view of the cloacal area showing spicules of P. rara from a bobcat in Nebraska, United States; C) ventral view of rays and associated velum of posterior end of P. rara from Iowa, United States. Note that the scale bar is the same for both C and D; D) ventral view of a dissected specimen of P. rara. Note that this is the same individual as is shown intact in figure C.
(Source: S. L. Gardner, HWML. License: CC BY.)

Genus Skrjabinoptera Shulz, 1927. These are found in reptiles

Species Skrjabinoptera phrynosoma (Ortlepp, 1922). These are a common stomach worm of reptiles that live in Texas, United States, as well as horned toads Phrynosoma cornutum (Anderson, 2000). The intermediate hosts are the ants Pogonomyrmex barbatus var. molefaciens (Lee, 1957).

Subfamily Proleptinae (Schulz, 1927)

Genus Neoleptus Ubelaker and Dailey, 1975. These are parasites found in the fish Heterodontus philippi and Mustelus antarticus (Specian et al., 1975).

Type species Neoleptus australis (Johnston and Mawson, 1943) Specian et al., 1975

Subfamily Mirzalopterinae Wason and Johnson, 1977

Type genus Mirzaloptera Wason and Johnson, 1977. These live in the stomach of the bat Rhinopoma microphyllum in Jodhpur, India.

Type species Mirzaloptera barbari Watson and Johnson, 1977

Family Rictulariidae (Hall, 1915 subfamily) Railliet, 1916

Genus Quentius Chabaud and Bain, 1981. These live in the duodenum and small intestine of Neotropical marsupials (Marmosa spp.) in Cali, Colombia (Chabaud and Bain, 1981).

Type species Quentius kozeki Chabaud and Bain, 1981

Genus Shamimana Gupta and Masoodi, 1990. These live in the intestine of the marine fish Plotosus arab off the Trivandrum coast near Kerala, India

Type species Shamimana durdanae Gupta and Masoodi, 1990

Superfamily Spiruroidea

The superfamily Spiruroidea combines the family Spiruridae Oerley, 1885.

Family Spiruridae Oerley, 1885

Genus Gastronodus Singh, 1934. These live in nodules on the stomach wall of the muskrat Crocidura coerulea in Hyderabad State, India.

Type species Gastronodus strasseni Singh, 1934

Genus Dollfusnema Caballero, 1974. These live in the intestine of the marine fish Paralabrax clathratus from Mexico.

Type species Dollfusnema piscicola Caballero, 1974

Genus Isospirura Sood and Parshad, 1972. These live in the stomach of Millardia meltada, Mus musculus bactrianes, and Mus booduga in Ludhiana, India.

Type species Isospirura meltadi Sood and Parshad, 1972

Genus Paracymea Gupta and Jaiswal, 1987. These live in the intestine of the birds Anser indicus in the Prince of Wales Zoological Gardens, Lucknow, India.

Type species Paracymea yamagutii Gupta and Jaiswal, 1987.

Family Gongylonematidae (Hall, 1916 subfamily) Sobolev, 1949

Genus Gongylonema Molin, 1857. These embed in the mucosa and submucosa of the anterior region of the gut of birds and mammals. Usually, the final hosts are sheep and goats, and sometimes also horses, cattle, swine, poultry, dogs, cats, and numerous other wild and domestic mammals and birds. As such, according to Soulsby (1982), some gongylonemids can affect the health of humans and domestic animals, for example, G. pulchrum Molin, 1857 which can be found in most parts of the world. This parasite species occurs in sheep, goats, cattle, pigs, zebu, buffalo, and (less frequently) horses, camels, donkeys, and wild boar. It may also develop in humans, particularly in the oral epithelium, but also subcutaneously (see Figure 5). The site of localization in non-human animals is the esophagus where G. pulchrum embeds in a zigzag pattern in the mucosa or submucosa. In ruminants, it may also appear in the rumen. The intermediate hosts are coprophagous beetles of the genera Aphodius, Onthophagus, Blaps, Caccobius, and others (over 70 species). Migrating juveniles root in the wall of the gastroesophageal region. They excyst in the stomach and then migrate anteriorly to the oral cavity and finally reach the wall of the esophagus. The species G. verrucosum (Giles, 1892) may be present in the rumen of sheep, goat, cattle, deer, and zebu in India, the United States, and South Africa. Gonglyonema monnigi Baylis, 1926 develops in the rumen of sheep and goats in South Africa. Gonglyonema ingluvicola Ransom, 1904 and G. crami Smit, 1927 occur in fowl in North America, India, the Philippines, Taiwan, Europe, and Australia. Gonglyonema sumani Bhalerao, 1933 occurs in the crop of domestic fowl in Uttar Pradesh State, India. The cockroach Blatella germanica may be infected with this worm. Gongylonema verrucosum embeds in the epithelium, causing just a slight chronic inflammatory reaction with hypertrophy and cornification, but G. ingluvicola may burrow into the crop and cause severe lesions in heavy infections.

Figure 5. Gongylonema spp. life cycle. Gongylonema is a genus of spirurid nematodes which includes the veterinary parasite G. pulchrum (also called the gullet worm or stitch worm) along with several other parasites of mammals and birds. Incidental human infections with Gongylonema are rare, and species-level identifications are difficult and seldom confirmed. The life cycle diagram of Gongylonema spp. shows: Adult Gongylonema inhabit the upper gastrointestinal tract of the definitive host in sites such as the mouth, esophagus, rumen, and stomach (1). The long, thin adults are found in shallow tunnels in the squamous epithelial surfaces of these tissues; the female produces thick-shelled, embryonated eggs containing first-stage (J₁) juveniles. Expelled eggs are released from the tunnels during epithelial desquamation and are carried down the gastrointestinal tract and shed in the feces (2). Intermediate host insects become infected after ingesting eggs in host feces (3). Juveniles develop in the hemocoel of the intermediate host, eventually becoming encapsulated as infective third-stage (J₃) juveniles in the thoracic muscles (4). Suitable definitive hosts become infected after ingesting infected intermediate hosts (5). Juveniles are released in the stomach, which embed in the gastric or duodenal mucosa, and eventually migrate to the upper gastrointestinal tract after 2–3 months (6). Migration of juveniles often creates characteristic zig-zag or sinusoidal tracks in the affected epithelial tissues. Maturation is completed in the upper gastrointestinal tract. Human infections occur following the ingestion of intermediate host arthropods (7), either intentionally or accidentally, in contaminated food or water. In these cases, worms have been found in the mucosal tissues of the lips, cheek, tongue, tonsils, gums, and occasionally esophagus. A few cases of spurious egg passage have been documented, which may be due to the inadvertent consumption of adult Gongylonema in certain types of meat (for example, chicken gizzards or pork tongue).
(Source: Adapted from Division of Parasitic Diseases and Malaria, United States Centers for Disease Control and Prevention, 2019. Public domain.)

Bickova and colleagues (2017) report some gongylonematid species that occur in Belarus, such as: Gongylonema neoplasticum (Fibiger et Ditlevsen), which occurs in the European water vole Arvicola amphibius, forest dormouse Dryomys nitedula, and common dormouse Muscardinus avellanarius, all from the Brest and Gomel regions (Luninety District). Gongylonema sorici Fain, 1955 is found in the common shrew Sorex araneus from NP “Belovezhskaya Pushcha” in Belarus. Kinsella and colleagues (2016) describe G. archboldi found in tunnels in the gastric mucosa of the cotton rat Sigmodon hispidus from Highlands County, Florida, United States. Measurements are also given for specimens from the cotton mice Peromyscus gossypinus, oldfield mice Pe. polionotus, Florida mice Podomys floridanus, and golden mice Ochrotomys nuttalli from the same locality. Additional specimens have been collected from the cotton rat and the rice rat Oryzomys palustris from Berry Island, San Patricio County, Texas, United States.

Subgenus Progongylonema Hernandez-Rodriguez and Gutierrez-Palomino, 1992. These live in the mucosa under the tongue of Pica pica, Garrulus glandarius, Cyanopica cyanus, and Corvus monedula (order Passeriformes, family Corvidae) in Córdoba Province, southern Spain.

Type species Gongylonema (Progongylonema) pacoi Hernandez-Rodriguez and Gutierrez- Palomino, 1992

Genus Chlamydoprocta Chandler, 1954. These are parasites of the skunk Mephitis mephitis in Minnesota, United States.

Type species Chlamydoprocta itascensis Chandler, 1954

Genus Mastigonema Dailey and Perrin, 1973. These are oviparous parasites of the forestomach of Cetacea, Stenella graffmani and S. longirostris, that are found in the eastern tropical Pacific Ocean.

Type species Mastigonema stellae Dailey and Perrin, 1973

Genus Mazzia Khalil and Vogelsang, 1932. These are found in dasypodid mammals in Argentina.

Type species Mazzia mazzia Khalil and Vogelsang, 1932

Genus Paraspiralatus Gibbons et al., 2004. This genus is morphologically more specialized than other Neotropical genera that parasitize paleoendemic mammals (Chabaud et al., 1983).

Type species Paraspiralatus sakeri. This was found recently in the stomach of a wild-caught, female saker falcon in Saudi Arabia (Gibbons et al., 2004).

Species Mazzia bialata. These parasitize dasypodid mammals (such as Chaetophractus villosus) from Buenos Aires, Argentina (Chabaud et al., 1983).

Genus Spirobakerus Chabaud and Bain, 1981. These are parasites of the cricetid Zygodontomys brevicauda of Colombia.

Type species Spirobakerus weitzeli Chabaud and Bain, 1981

Genus Spirosprattus Smales, 2004. These are found in cysts in the stomach wall of Australian rodents, such as the Cape York rat Rattus leucopus (family Muridae).

Type species Spirosprattus scyphiformis Smales, 2004

Family Pneumospiruridae Wu and Hu, 1938

Genus Pneumospirura Wu and Hu, 1938

Type species Pneumospirura hainanensis Wu and Hu, 1938?. These are parasites of birds and mammals, including some carnivores, including the bobcat Felis rufus in North America (as redescribed by Pence and Stone, 1977).

Species Pneumospirura bassarisci. These are localized on the bronchioles of the ringtail Bassariscus astutus (Pence and Stone, 1977).

Species Pneumospirura capsulata. These are parasites in the common badger.

Species Pneumospirura rodentium. Wertheim and Giladi (1977) described these as a lung parasite of Gerbillus dasyurus and Meriones crassus.

Superfamily Thelazioidea

Family Thelaziidae Skrjabin, 1915

Genus Thelazo Pearse, 1933. According to Pearse (1933), Thelazo is erected for T. glossogobii described from the definitive host, the tank goby Glossogobius giurus. The diagnosis is based on the work of Pearse (1933) who placed the genus in the family Thelaziidae.

Type species is Thelazo glossogobii Pearse, 1933. These are found in marine and brackish waters from the Red Sea, East Africa, South Asia, the Indian Ocean, China, Australia, and the islands of the Pacific Ocean.

Subfamily Thelaziinae (Skrjabin, 1915 family) Baylis and Daubney, 1926

Genus Thelazia Bosc, 1819

Species Thelazia skrjabini Ershov, 1928. These are parasites of the orbits (including under the lids, conjunctiva, and nictitating membrane, and in the lachrymal glands and ducts) of birds and mammals, including cattle in Europe, Asia, and North America.

Species Thelazia rhodesii (Desmarest, 1828). These occur primarily in cattle, sheep, goats, and buffaloes, and its habitat is cosmopolitan (Soulsby, 1982). The intermediate hosts are Musca larvipara, M. convexifrons, and M. amica.

Species Thelazia gulosa Railliet and Henry, 1910. These appear in cattle in most parts of the world.

Species Thelazia alfortensis Railliet and Henry, 1910. These occur in cattle in Europe.

Species Thelazia lacrymalis (Gurlt, 1831). These develop in the horse in most parts of the world. Musca oseris transmits T. lachrymalis in regions delineated by the former Soviet Union, while M. autumnalis appears to be an important vector in the United States (Soulsby, 1982).

Species Thelazia callipaeda Railliet and Henry, 1910. These develop in the fat body (Anderson, 2000) and live under the nictitating membrane of the dog in East Asia and has been reported from rabbits and humans.

Species Thelazia californiensis Price, 1930. These occur in sheep, deer, cats, dogs, and humans in the United States.

Species Thelazia leesei Railliet and Henry, 1910. These have been reported from dromedary camels in the former Soviet Union and elsewhere in Asia.

Subgenus Isothela Railliet, 1925. These are viviparous parasites occurring in birds.

Subgenus Pericyema Railliet, 1925. These are ovoviviparous parasites in mammals.

Type species Thelazia (Pericyema) callipaeda Railliet and Henry, 1910.

Subgenus Thelazia (Bosc, 1819 genus). These are viviparous parasites that occur in mammals.

Type species Thelazia (Thelazia) rhodesi (Desmarest, 1827) Railliet and Henry, 1910

Family Rhabdochonidae (Travassos, Artigas and Pereira, 1928 subfamily) Skrjabin, 1946. These are parasites that live in the gallbladder of freshwater fish.

Genus Beaninema Caspeta-Mandujano et al., 2001. Caspeta-Mandujano and colleagues (2001) re-erected this genus and described a new species and new genus. The members of this genus are parasites in the gallbladder of the freshwater fish Cichlasoma hearli from the Santiago River, Tepic, Nayarít, Mexico.

Type species Beaninema nayaritense Caspeta-Mandujano et al., 2001.

Genus Fellicola Petter and Køie, 1993. These are parasites that live in the gallbladder of the marine fish Coryphaenoides rupestris (a ray-finned fish) from the North Atlantic off the Faroe Islands (Petter and Køie, 1993). According to Petter and Køie (1993), the new genus is close to the genera Johnstonmawsonia, Vasorhabdochona, and Pancreatonema but differs from these genera in having longitudinal thickenings in the anterior dilated part of the pharynx.

Type species Fellicola longispiculus Petter and Køie, 1993 (Gibbons, 2010).

Genus Megachona Mejía-Madrid and Pérez-Ponce de León, 2007. Megachona most closely resembles Beaninema Caspeta-Mandujano et al., 2001, F. Petter and Køie, 1993, and Rhabdochona Railliet, 1916.

Type species Megachona chamelensis Mejía-Madrid and Pérez-Ponce de León, 2007. These are from the intestinal cecae of the blue striped chub Sectator ocyurus (Kyphosidae, Perciformes) from Chamela Bay, Mexico.

Genus Rhabdochona Railliet, 1916. The reconstruction of this genus was suggested by Moraveč (1975). These live in the intestines of fish.

Subgenus Afrochona Puylaert, 1973. These live in the intestine of the fish Aphyosemion cameronensis in Olounou, Cameroon (Gibbons, 2010).

Type species is Rhabdochona (Afrochona) camerounensis Puylaert, 1973

Subgenus Globochonoides Moravec, 1975. These live in the intestine of freshwater fishes.

Type species Rhabdochona (Globochonoides) coronacauda Belouss, 1965

Species Rhabdochona (Globochonoides) pseudomysti. This is 1 of 2 recently species of rhabdochonid nematodes that live in the intestines of freshwater fishes in Chiang Mai Province, northern Thailand as recorded by Moraveč and Yooyen (2011). It is from the catfish Pseudomystus siamensis (Regan) (Bagridae, Siluriformes) from Fang Brook, a tributary of the Kok River in the Mekong River basin, Fang District, Thailand.

Species Rhabdochona (Globochona) thaiensis is also recently described and is from the cyprinid Mystacoleucus marginatus (Valenciennes) (Cyprinidae, Cypriniformes) in the Ping River in the Chao Phraya River basin, Muang District, Thailand. These are the first nominal species of Rhabdochona reported from Thailand (Moraveč and Yooyen, 2011).

Species Rhabdochona (Globochona) rasborae. These are found in the intestine of the freshwater cyprinid fish, sidestripe rasbora Rasbora paviana from Tirant in the Bangbaimai Subdistrict, Muang District, Surat Thani Province, southern Thailand. This is the third nominal species of Rhabdochona Railliet, 1916, and the second species of the subgenus Globochona reported from fishes in Thailand (Moraveč and Kanda, 2012).

Genus Johnstonmawsonoides Machida, 1975. These live in the intestine of the marine teleost fishes Nemichthys scolopaceus in Suruga Bay, Japan.

Type species Johnstonmawsonoides nemichthyos Machida, 1975. These are among the known helminths of meso- and bathypelagic fishes of Norfolk Submarine Canyon in the western North Atlantic (Gartner and Zwerner, 1989).

Genus Neoascarophis Machida, 1976. These live in the intestine of the marine teleost fishes Coelorhynchus multispinulosus and Bathygadus garretti in Suruga Bay, Japan.

Type species Neoascaropbis yarihige Machida, 1976

Subfamily Prosungulonematinae Skrjabin et al., 1967. This subfamily was presented by Chabaud (1975) as a synonym of Rhabdochonidae.

Type genus Prosungulonema Roitman, 1963. McVicar and Gibson (1975) supported the validity of the genus Prosungulonema. Later, Caspeta-Mandujano and colleagues (2001) did not list the genus as valid in the family Rhabdochonidae. These are parasites of freshwater teleost fishes.

Type species P. siniperca (Dogiel and Akhmerov, 1959)

Genus Pancreatonema McVicar and Gibson, 1975 (according to Chabaud, 1975)

Type species Pancreatonema torriensis McVicar and Gibson, 1975

Species Pancreatonema torriensis. These are from the pancreatic duct of Raja naevus from off the coast of Aberdeen in northwest Scotland, as described and discussed by McVicar and Gibson (1975).

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Supplemental Reading

Chabaud, A. G. 1954. Sur le cycle évolutif des Spirurides et de Trématodes ayant une biologie comparable: Valeur systématique des caractères biologiques (suite et fin). [In French.] Annales de parasitologie humaine et comparée 29: 358–425.

De, N. C., and R. N. Maity. 1999. Larval development of Onchocamallanus bagarii (Nematoda: Camallanidae) in copepods. Folia Parasitologica 46: 53–58.

Poinar, G. O., Jr., and G. M. Thomas. 1976. Occurrence of Ascarophis (Nematoda: Spiruridea) in Callianassa californiensis Dana and other decapod crustaceans. Proceedings of the Helminthological Society of Washington 43: 28–33.

Yorke, W., and P. A. Mapelstone. 1926. The Nematode Parasites of Vertebrates. Churchill, London, United Kingdom, 536 p.