Members of the Philometridae represent the most important group of dracunculoid nematodes parasitizing fishes. In his monograph treating the Dracunculoidea, Moravec (2006) reported a total of 11 genera and 105 species of philometrids parasitizing freshwater, brackish-water and marine fishes. However, during the last six years (2007-2012), an additional 42 new species of Philometridae have been described, representing a 40% increase of the number of nominal species. Most of these species (30) belong to Philometra Costa, 1845, mainly represented by parasites of marine fishes, a few others (8) to Philometroides Yamaguti, 1935, and a single one to each of the following genera: Caranginema Moravec, Montoya-Mendoza et Salgado-Maldonado, 2008, Dentiphilometra Moravec et Wang, 2002, Dentirumai Quiazon et Moravec, 2013* and Spirophilometra Parukhin, 1971. Moreover, three new genera, Afrophilometra Moravec, Charo-Karisa et Jirků, 2009, Caranginema and Dentirumai, were erected. Representatives of seven genera, Afrophilometra, Buckleyella Rasheed, 1963, Caranginema, Dentiphilometra, Dentirumai, Paraphilometroides Moravec et Shaharom-Harrison, 1989 and Rumai Travassos, 1960, were studied using scanning electron microscopy (SEM) for the first time. Thirteen known but poorly described philometrid species were redescribed and, in some species of Caranginema and Philometra, previously unknown conspecific males were discovered and described. The male surface ultrastructure studied by SEM provided new taxonomically important features for species distinction. Gene sequencing was used in several recent studies and advanced our understanding of phylogenetic interrelationships among representatives of seven genera (Afrophilometra, Alinema Rasheed, 1963, Caranginema, Nilonema Khalil, 1960, Philometra, Philometroides and Rumai) and of the extent of the biodiversity of philometrids. New data were obtained on the biology and pathogenicity of several species of Nilonema, Philometra, Philometroides and Rumai. The need to carry out surveys in order to find males and to use SEM and gene sequencing to identify philometrids is emphasized. Appropriate quantitative methods to determine the impact of philometrids in ovarian tissue on host fecundity are recommended. Further detailed studies on philometrids would be significant not only from the theoretical viewpoint, but also because of their practical implications. A list of philometrid nematode species by continents is provided.
Examinations of nematodes collected from some marine fishes off the southwestern coast of Java, Indonesia in 2000 and 2001 revealed the presence of the following six species: ascaridoids Ichthyascaris grandis sp. n. from the intestine of Lophiomus setigerus (Vahl), I. cf. longispicula Li, Liu, Liu et Zhang, 2012 from the intestine of Conger cinereus Rüppel, Ichthyascaris sp. from the body cavity of Lobotes surinamensis (Bloch), and Raphidascaroides halieutaeae Yin, 1983 from the intestine of Halieutaea stellata (Vahl), and philometrids Philometra ivaschkini Parukhin, 1976 from the stomach wall of Trichiurus lepturus Linnaeus and P. psettoditis Moravec, Walter et Yuniar, 2012 from the body cavity (liver) of Psettodes erumei (Bloch et Schneider). Descriptions of these nematodes based on light and scanning electron microscopical studies are provided. The new species I. grandis sp. n. is mainly characterised by large body measurements (males and females up to 41.8 mm and 73.6 mm long, respectively), the length of spicules (0.99-1.05 mm), the tail tip usually without rudimentary spines and by the presence of 44-53 pairs of caudal papillae, eight to twelve of which being postanals. In addition to new data on the morphology of R. halieutaeae and other nematodes recorded, the 11 species of Raphidascaroides Yamaguti, 1941 poorly described from marine fishes in South Asia and reviewed in the monograph of Sood (2017) are considered species inquirendae and incertae sedis.
Based on light and scanning electron microscopical studies, the following five species of the Philometridae (Nematoda: Dracunculoidea) are described from female specimens collected from marine fishes off the southwestern coast of Java, Indonesia: Philometra lobotidis sp. n. from the abdominal cavity of the Atlantic tripletail Lobotes surinamensis (Bloch) (Lobotidae, Perciformes); Philometra javaensis sp. n. from the abdominal cavity of the immaculate puffer Arothron immaculatus (Bloch et Schneider) (Tetraodontidae, Tetraodontiformes); Philometra psettoditis sp. n. from the musculature of the Indian spiny turbot Psettodes erumei (Bloch et Schneider) (Psettodidae, Pleuronectiformes); Philometroides indonesiensis sp. n. from the musculature of the hound needlefish Tylosurus crocodilus crocodilus (Péron et Lesueur) (Belonidae, Beloniformes); and Philometroides trichiuri sp. n. from the dorsal fin of the largehead hairtail Trichiurus lepturus Linnaeus (type host) and the savalai hairtail Lepturacanthus savala (Cuvier) (both Trichiuridae, Perciformes). All these new species are distinguished from their congeners parasitizing marine fishes by morphological (mainly the shape and structure of the cephalic and caudal ends and of the oesophagus) and biometrical features. Besides previously known Philometra pellucida (Jägerskiöld, 1893) and Philometra ocularis Moravec, Ogawa, Suzuki, Miyazaki et Donai, 2002, they are the only nominal philometrid species recorded from Indonesian waters.
Recent examinations of some marine fishes from off the southern coast of Iraq revealed the presence of five species of Philometra Costa, 1845 (Nematoda: Philometridae): Philometra arabiensis sp. n. (males and females) from the ovary of the shrimp scad Alepes djedaba (Forsskål) (Carangidae, Carangiformes), Philometra psettoditis Moravec, Walter et Yuniar, 2012 (females) from the body cavity of the Indian halibut Psettodes erumei (Bloch et Schneider) (Psettodidae, Pleuronectiformes), Philometra terapontis Moravec, Gopalakrishnan, Rajkumar, Saravanakumar et Kaliyamoorthy, 2011 (female) from the ovary of the jarbua terapon Terapon jarbua (Forsskål) (Terapontidae, Centrarchiformes), Philometra sp. (females) from the ovary of the Arabian blackspot threadfin Polydactylus mullani (Hora) (Polynemidae, Carangariformes) and Philometra sp. 2 of Moravec et al. (2016a) (females) from the ovary and body cavity of the bartail flathead Platycephalus indicus (Linnaeus) (Platycephalidae, Perciformes). Philometra arabiensis sp. n. is mainly characterised by the length of spicules (198-243 µm) and gubernaculum (75-99 µm), the gubernaculum/spicule length ratio (1 : 2.33-2.79), the structure of the gubernaculum distal portion and the male caudal end, and the body length of males (1.86-2.73 mm). The present findings of P. psettoditis and P. terapontis in fishes of the Arabian Gulf represent new geographical records for these parasites.
Based on light and scanning electron microscopical studies, the following five gonad-infecting species of the Philometridae (Nematoda: Dracunculoidea) are described from marine perciform fishes off the eastern coast of India (Bay of Bengal): Philometra sphyraenae sp. n. (males and females) from the pickhandle barracuda Sphyraena jello Cuvier (Sphyraenidae), Philometra gerrei sp. n. (males and females) from the whipfin silver-biddy Gerres filamentosus Cuvier (Gerreidae), Philometra otolithi sp. n. (single female) from the tigertooth croaker Otolithes ruber (Bloch et Schneider) (Sciaenidae), Philometra sp. (females) from the Belanger's croaker Johnius belangerii (Cuvier) (Sciaenidae), and Philometroides eleutheronemae sp. n. (females) from the fourfinger threadfin Eleutheronema tetradactylum (Shaw) (Polynemidae). All new species are distinguished from their congeners parasitizing gonads of marine fishes by morphological (mainly the gubernaculum structure in males and the shape and structure of the cephalic and caudal ends and of the oesophagus in females) and biometrical features. Philometra rajani Mukherjee, 1963 is considered a species inquirenda.
Specimens of three little-known species of Rhabdochona (Nematoda: Rhabdochonidae) were collected during occasional examinations of some freshwater fishes in India: R. (Rhabdochona) hellichi turkestanica (Skryabin, 1917) in Schizothorax sp. (Cyprinidae, Cypriniformes) from the Lodhomakhola and Rangit Rivers, West Bengal and Sikkim, respectively; R. (R.) hospeti Thapar, 1950 in Tor sp. (Cyprinidae) from the Rangit River; and R. (Globochona) mazeedi Prasad et Sahay, 1965 in Clupisoma garua (Hamilton) (Schilbeidae, Siluriformes) from the Farakka Dam Lake, West Bengal. Their detailed light and electron microscopical studies revealed some taxonomically important, previously not observed features and made possible their redescription. Fourth-stage larvae of R. hospeti are described for the first time. Rhabdochona hellichi turkestanica (syns. R. denudata filamentosa Bykhovskaya-Pavlovskaya, 1936, R. kashmirensis Thapar, 1950, R. schizothoracis Siddiqi et Khattak, 1984) is proposed as a subspecies, differing from the nominotypical subspecies R. hellichi hellichi (Šrámek, 1901) mainly in the shape of the distal end of the left spicule, molecular data and geographical distribution. Rhabdochona moraveci Katoch et Kalia, 1991 (a homonym to R. moraveci Duggal et Kaur, 1987) is renamed R. indica nom. n. The following six species are considered new junior synonyms of R. hospeti: Comephronema [sic] mackiewiczi Malhotra et Rautela, 1984, Rhabdochona moraveci Duggal et Kaur, 1987, R. bifidum Kakar et Bilqees, 2007, R. uvaginus Kakar et Bilqees, 2007, R. bolani Kakar, Bilqees et Ahmad, 2008 and R. cephalodiverticula Kakar, Bilqees et Ahmad, 2008. Rhabdochona edentati Paul et Majumdar, 1994 is considered a species incertae sedis.
Based on light and scanning electron microscopical studies, the following nine species of Philometridae (Nematoda: Dracunculoidea) are described from female worms parasitizing marine perciform fishes belonging to six families off the northern coast Australia (near Darwin): Philometra australiensis sp. n. from the swimbladder of the king threadfin Polydactylus macrochir (Günther) (Polynemidae); P. epinepheli Dewi et Palm, 2013 from the operculum of the orange-spotted grouper Epinephelus coioides (Hamilton) (Serranidae); Philometra johnii Moravec et Ali, 2013 from the gonad of the croaker Johnius sp. (Sciaenidae); P. macrochiri sp. n. from the sensory fin of P. macrochir; P. zabidii sp. n. from the ovary of the ninespine batfish Zabidius novemaculeatus (McCulloch) (Ephippidae); Philometra sp. 1 and Philometra sp. 2 from the ovary of the Spanish flag snapper Lutjanus carponotatus (Richardson) (Lutjanidae) and the silver grunt Pomadasys argenteus (Forsskål) (Haemulidae), respectively; Philometroides eleutheronemae Moravec et Manoharan, 2013 from the ovary of the fourfinger threadfin Eleutheronema tetradactylum (Shaw) (Polynemidae); and Spirophilometra endangae Dewi et Palm, 2013 from the pectoral fins of E. coioides. The new species P. australiensis is characterized mainly by the structure of the cephalic end, 14 minute cephalic papillae, absence of caudal projections and body length of gravid female (67 mm), P. macrochiri by the presence of a conspicuously large anterior oesophageal bulb, 14 very small cephalic papillae and the truncated posterior end of body without any caudal projections, whereas P. zabidii is characterized by the presence of distinct caudal projections, the number (14) and larger size and arrangement of cephalic papillae, a poorly developed anterior oesophageal inflation, the body length (114 mm) and the host family (Ephippidae). All above-mentioned species were recorded from Australian waters for the first time.
Small subunit rRNA sequences were obtained from 38 representatives mainly of the nematode orders Spirurida (Camallanidae, Cystidicolidae, Daniconematidae, Philometridae, Physalopteridae, Rhabdochonidae, Skrjabillanidae) and, in part, Ascaridida (Anisakidae, Cucullanidae, Quimperiidae). The examined nematodes are predominantly parasites of fishes. Their analyses provided well-supported trees allowing the study of phylogenetic relationships among some spirurine nematodes. The present results support the placement of Cucullanidae at the base of the suborder Spirurina and, based on the position of the genus Philonema (subfamily Philoneminae) forming a sister group to Skrjabillanidae (thus Philoneminae should be elevated to Philonemidae), the paraphyly of the Philometridae. Comparison of a large number of sequences of representatives of the latter family supports the paraphyly of the genera Philometra, Philometroides and Dentiphilometra. The validity of the newly included genera Afrophilometra and Caranginema is not supported. These results indicate geographical isolation has not been the cause of speciation in this parasite group and no coevolution with fish hosts is apparent. On the contrary, the group of South-American species of Alinema, Nilonema and Rumai is placed in an independent branch, thus markedly separated from other family members. Molecular data indicate that the skrjabillanid subfamily Esocineminae (represented by Esocinema bohemicum) should be either elevated to the rank of an independent family or Daniconematidae (Mexiconema africanum) should be decreased to Daniconematinae and transferred to the family Skrjabillanidae. Camallanid genera Camallanus and Procamallanus, as well as the subgenera Procamallanus and Spirocamallanus are confirmed to be paraphyletic. Paraphyly has also been found within Filarioidea, Habronematoidea and Thelazioidea and in Cystidicolidae, Physalopteridae and Thelaziidae. The results of the analyses also show that Neoascarophis, Spinitectus and Rhabdochona are monophyletic, in contrast to the paraphyletic genus Ascarophis. They further confirm the independence of two subgenera, Rhabdochona and Globochona, in the genus Rhabdochona. The necessity of further studies of fish-parasitizing representatives of additional nematode families not yet studied by molecular methods, such as Guyanemidae, Lucionematidae or Tetanonematidae, is underscored.
About 300 species belonging to four superfamilies (Gnathostomatoidea, Habronematoidea, Physalopteroidea and Thelazioidea) of the nematode suborder Spirurina are known as the adult parasites of freshwater, brackish-water and marine fishes. They are placed in four families, of which the Gnathostomatidae, including Echinocephalus with a few species and the monotypic Metaleptus, are parasites of elasmobranchs, whereas Ancyracanthus contains one species in teleosts; the Physalopteridae is represented in fish by four genera, Bulbocephalus, Heliconema, Paraleptus and Proleptus, each with several species in both elasmobranchs and teleosts. The majority of fish spirurines belongs to the Rhabdochonidae, which includes 10 genera (Beaninema, Fellicola, Hepatinema, Heptochona, Johnstonmawsonia, Megachona, Pancreatonema, Prosungulonema, Rhabdochona and Vasorhabdochona) of species parasitizing mainly teleosts, rarely elasmobranchs, and the Cystidicolidae with about 23 genera (Ascarophis, Caballeronema, Capillospirura, Comephoronema, Crenatobronema, Cristitectus, Ctenascarophis, Cyclozone, Cystidicola, Cystidicoloides, Johnstonmawsonoides, Metabronema, Moravecnema, Neoascarophis, Parascarophis, Prospinitectus, Pseudascarophis, Pseudoproleptus, Salvelinema, Similascarophis, Spinitectoides, Spinitectus, Sterliadochona), with many species parasitic in teleosts only. Because of difficulties in studying fish spirurines, associated with their morphological and biological peculiarities, most species of these parasites are poorly known. It is apparent that their present classification system does not reflect phylogenetic relationships and a taxonomic revision of this nematode group, based on detailed morphological (including SEM and TEM), life history and molecular studies of individual species, is quite necessary. In Cystidicolidae, several genera have been based on details in the cephalic structures visible only with the aid of SEM, but it will be evident whether or not these tiny features are of generic importance only when more cystidicolids are described using SEM and comparative molecular data become available. Data on the biology of fish spirurines are scarce. In known cases, their life cycles involve aquatic arthropods (crustaceans or insects) as intermediate hosts, in which, sometimes, the larvae undergo a precocious development and may even attain adulthood and become gravid in these invertebrates; sometimes, fish paratenic hosts are known to occur in cystidicolids parasitizing as adults piscivorous definitive hosts. Some spirurine species are pathogenic and are known as causative agents of serious fish diseases. and Consequently, further detailed studies on fish spirurines are significant not only from the theoretical viewpoint, but they may also have practical implications.
The nematode superfamily Dracunculoidea includes 166 recognized species, of which 150 (90%) are parasitic in about 300 species of freshwater, brackish-water and marine fishes. Fish dracunculoids are placed in 31 genera (86% of all dracunculoid genera) belonging to eight of the nine dracunculoid families: Anguillicolidae, Daniconematidae, Guyanemidae, Lucionematidae, Micropleuridae, Philometridae, Skrjabillanidae, and Tetanonematidae; the genus Lockenloia is considered incertae sedis. Because of difficulties in studying fish dracunculoids, associated with their morphological and biological peculiarities, most species of these largely histozoic parasites are poorly known and males of the majority of species and of eight genera have not yet been discovered. It is apparent that the present classification system of dracunculoids as a whole does not reflect phylogenetic relationships and a taxonomic revision of this nematode group, based on detailed morphological (including SEM and TEM), life history and molecular studies of individual species, is quite necessary. Data on the biology of fish dracunculoids is scarce. In known cases, their life cycles involve copepods, ostracods or branchiurids as intermediate hosts and, sometimes, fish paratenic hosts are known to occur in dracunculoid species parasitizing as adults piscivorous definitive hosts. However, nothing is known about the life cycles of representatives of 20 genera. Some species of dracunculoids, particularly of philometrids, are highly pathogenic and are known as agents of serious fish diseases. During recent years, especially the importance of Philometra spp. parasitizing the gonads of many species of marine fishes has increased due in particular to the rapid development of marine aquaculture, because they may significantly decrease fish reproduction or even cause full parasitic castration. Therefore, further detailed studies on fish dracunculoids are significant not only from the theoretical viewpoint, but they may also have practical implications.