I. Kanev

Life-cycle, delimitation and redescription of Echinostoma revolutum (Froelich, 1802) (Trematoda: Echinostomatidae)

The life-cycle of Echinostoma revolutum (Froelich, 1802) Dietz, 1909 has been completed experimentally beginning with infected snails collected at the type-locality, near Erlangen, Germany. Based on the specimens obtained, each stage of the life-cycle has been redescribed. Important taxonomic features are discussed and hitherto unknown characteristics are described. Synonyms for E. revolutum are: Fasciola revoluta Froelich, 1802; Echinostoma paraulum Dietz, 1909; E. audyi Lie & Umathevy, 1965; and E. ivaniosi Mohandas, 1973. Adults and larvae described as E. revolutum in other works are found to be identical with Echinostoma echinatum (Zeder, 1803), E. trivolvis (Cort, 1914), E. jurini (Skvortsov, 1924), E. caproni Richard, 1964, Moliniella anceps (Molin, 1859), Echinochasmus beleocephalus (Linstow, 1873) and other echinostome species. For nearly a century, incorrect morphological, biological, life-cycle and host information has been attributed to E. revolutum, and at times these data have contributed to the diagnoses of the species. Occasionally, authors actually working with E. revolutum have ascribed their results to other species. Based on extensive experimental life-cycle studies beginning with infected snails from type-localities, it is shown that (1) the first intermediate host is a lymnaeid snail; (2) the second intermediate hosts are various pulmonate and prosobranch snails, mussels, frogs and freshwater turtles; (3) the final hosts are birds; (4) E. revolutum cercariae and adults have 37 collar spines; (5) the species occurs only in Europe and Asia; (6) Cercaria echinata Siebold, 1937, Echinostoma echinatum (Zeder, 1803) and E. jurini (Skvortsov, 1924) are the closely related 37-spined allies in Europe; and (7) species specific characteristics are expressed only in the larvae and the host-parasite relationships. The adults of E. revolutum cannot be identified using morphological criteria and it is proposed that worms with 37 collar spines belonging to the genus Echinostoma and occurring in naturally infected birds in Europe and Asia be referred to an “E. revolutum group.”

REDESCRIPTION OF ECHINOSTOMA TRIVOLVIS (CORT, 1914) (TREMATODA: ECHINOSTOMATIDAE) WITH A DISCUSSION ON ITS IDENTITY

The life-cycle of Echinostoma trivolvis (Cort, 1914) has been completed experimentally and the validity and identity of this species are discussed. Synonyms for cercariae and adults of E. trivolvis are as follows: Cercaria trivolvis Cort, 1914, C. trisolenata Faust, 1917, C. acanthostoma Faust, 1918, C. complexa Faust, 1919; Distoma echinatum Zeder, 1803, of Leidy (1888, 1904) and Hassall (1896); E. echinatum (Zeder, 1803) of Hassall (1896), Stiles & Hassall (1895), Barker & Laughlin (1911), Barker (1916) and Swales (1933); E. revolutum (Frölich, 1802) Dietz, 1909 of Johnson (1920), Fallis (1934), Beaver (1937) and Fried & co-workers (1968–1989); E. armigerum Barker & Irvine, 1915; E. coalitum Barker & Beaver, 1915; E. callawayensis Barker & Noll, 1915; E. paraulum Dietz, 1909 of Miller (1937); E. multispinosum Vigueras, 1944; and Echinoparyphium contiguum Barker & Bastron, 1915. The first intermediate host is the planorbid snail Helisoma trivolvis. Second intermediate hosts are various pulmonate and prosobranch snails, mussels, planarians, fishes, frogs, tadpoles and freshwater turtles. Final hosts are various birds and mammals. E. trivolvis occurs only in North America.

The occurrence and identification of Echinostoma revolutum from North American Lymnaea elodes snails.

Lymnaea elodes snails collected in northern Indiana, U.S.A., were infected with larval stages of an echinostome bearing 37 collar spines and resembling members of the Echinostoma group. The taxonomic status of this digenean was determined through experimental infections of various definitive and first-intermediate hosts. In addition, characteristics of the penetration and paraesophageal glands in cercariae from this echinostomatid were compared with those from E. revolutum and E. trivolvis. Results indicate that this recently discovered 37-collar-spined echinostome parasitizing lymnaeid snails is E. revolutum, making this the first clear report of this trematode in North America.

The Taxonomy and Biology of Philophthalmid Eyeflukes

Publisher Summary This chapter discusses the stages of the life cycle of eyeflukes. Eyeflukes in the genus Philophthalmus are found world-wide and occur in both marine and fresh-water habitats. Most live in the orbit of birds in various microhabitats, but a few can be found in the intestine and at least one species in the mouth cavity. Significant information is available on the mating behavior of adults, the host-finding behavior of miracidia, and cyst formation by cercariae. The adult stages are good research subjects because they are readily accessible in the orbit of birds. Light microscopy studies indicate the presence of various surface features on eye fluke rediae, such as sensory papillae, a birth pore, and posterior appendages. Metacercariae are infective immediately upon encystment and rotation within the cyst. Eyefluke metacercarial cysts have their narrow end open to the environment, and they do not survive for long periods of time. Transplant studies are especially facilitated because major surgery is not required to transfer worms from host to host.

An Overview of the Biology of Echinostomes

Echinostomes are digenean parasites in the class Trematoda. The adult worms are parasites of fishes, reptiles, birds and mammals, including humans. Their characteristic morphological feature is a crown of spines and a collar-like tegumentary flap surrounding the oral sucker. Larval stages develop in freshwater and marine snails, mussels, insects, amphibians and reptiles. Adult and larval echinostomes are used widely as models for laboratory work for the following reasons: 1). They are wide spred and common and some species are cosmopolitan. Other species are found in several geographical regions. Relatively few species are restricted locally. 2). They are found in many wild and domestic birds and mammals, including humans. Many echinostomes are hematophagous, ingesting lymph and blood from their hosts. Therefore, they are parasites with significant health and economic importance. Some species such as Echinostoma lindoense Sandground & Bonne, 1940 (known after Kanev, 1985 as E. echinatum Zeder, 1803) and E. ilocanum (Garrison, 1908) cause severe health problems in humans in Southeast Asia. 3). Echinostomes infect most organs and sites in the abdominal cavity of birds and mammals, including the alimentary, excretory and reproductive systems. 4). Larval and adult echinostomes provide good material for biological, physiological, biochemical, immunological and other studies in modern biology. 5). Echinostomes can be maintained easily and inexpensively in the laboratory.

LIFE HISTORY AND IDENTIFICATION OF PHILOPHTHALMUS LUCIPETUS FROM ISRAEL

A Philophthalmus species whose larval stages have been isolated from Melanopsis praemorsa snails collected in Israel and formerly identified as probably belonging to Philophthalmus palpebrarum has now been reevaluated as belonging to Philophthalmus lucipetus. The present determination is based on a detailed study of all the stages of the parasites life cycle--eggs, intramolluscan stages, cercariae, and adults. They were compared to the original species P. lucipetus, recently reexamined and redescribed from source material in Vienna, Austria. The identity of the eyefluke from Israel and P. lucipetus was further confirmed by successful cross-infections employing miracidia of the Israeli isolate in European (Bulgarian) Fagotia acicularis acicularis snails on the one hand and miracidia of P. lucipetus from Bulgaria in Israeli Melanopsis praemorsa on the other hand, but none infected Melanoïdes tuberculata snails from Israel.

LIFE CYCLE AND IDENTIFICATION OF AN EYEFLUKE FROM ISRAEL TRANSMITTED BY MELANOIDES TUBERCULATA (MÜLLER, 1774)

A philophthalmid species from Israel using the freshwater snail Melanoides tuberculata as intermediate host was studied. The biological and morphological characteristics of all developmental stages of the life cycle of this philophthalmid were described, and compared to those of Philophthalmus lucipetus Rudolphi, 1819 from Israel, Philophthalmus gralli Mathis and Leger, 1910 from Jordan, Philophthalmus palpebrarum Looss, 1899, Philophthalmus nocturnus Looss, 1907, Cercaria distomatosa Looss 1896 from Egypt, and Philophthalmus lucknowensis Baugh, 1962 from India. The possible identity with 1 of these species is discussed. On the basis of comparative analysis of the data for all parasite stages in the life cycle, geographical distribution, snail hosts, and snail host specificity, we propose to designate the Israeli Melanoides tuberculata-transmitted eye fluke to Philophthalmus distomatosa n. comb. (Looss, 1896), (Digenea: Philophthalmidae).

Comparative observations on cercariae and metacercariae of Echinostoma trivolvis and Echinoparyphium sp.

Comparative observations were made on cercariae of Echinoparyphium sp. from Physa gyrina in Charlies pond, Stokes County, North Carolina and cercariae of Echinostoma trivolvis from Helisoma trivolvis in Northampton County, Pennsylvania. The cercaria of Echinoparyphium sp. has 43 collar spines, lacks penetration and paraesophageal glands, and has a conical tail without fin folds. The cercaria of E. trivolvis has 37 collar spines, penetration and paraesophageal glands, a finger-like process at the tip of the tail and fin folds. The length of the cercarial body and tail of E. trivolvis was significantly greater than that of Echinoparyphium sp. Cercariae of both species encysted in Biomphalaria glabrata snails in single and concurrent infections. In concurrent infections with a single cercaria of each species, 2 encysted metacercariae were adjacent to each other in the saccular kidney of the snail at 24 hr postinfection. The diameter of encysted metacercariae of E. trivolvis was significantly greater than that of Echinoparyphium sp. Echinoparyphium sp. metacercariae excysted at 39 C in an alkaline trypsin-bile salts medium used previously to excyst E. trivolvis. The length of excysted metacercariae of E. trivolvis was significantly greater than that of Echinoparyphium sp.

Life-cycle, delimitation and redescription of Catatropis verrucosa (Frölich, 1789) Odhner, 1905 (Trematoda: Notocotylidae)

The life-cycle of Catatropis verrucosa (Frölich, 1789) Odhner, 1905 has been completed experimentally starting from infected snails collected along the River Danube in Europe. Each stage of the life-cycle is redescribed. Taxonomic problems are discussed and the main features of the species are listed. Synonyms for C. verrucosa are Fasciola verrucosa Frölich, 1789, F. anseris Gmelin, 1790, Monostoma verrucosa (Frölich, 1789) Zeder, 1800, and Catatropis charadrii Skrjabin, 1915. Other names, such as Notocotylus triserialis Diesing, 1839, Notocotyle triseriale (Diesing, 1839) Diesing, 1850, Monostoma verrugueux Dujardin, 1845, “Monostoma sp. du canard” of Blanchard (1847), Notocotyle verrucosum (Frölich, 1789) Monticelli, 1892, N. verruqueux Railliet, 1895, and Distoma verrucosum (Frölich, 1789) Wolffhugel, 1900, were found to represent adults and/or larvae of C. verrucosa. Conversely, but less often, adults and larvae of other species were found described and illustrated as C. verrucosa. One of these, C. verrucosa of Joyeux (1922), was renamed Pseudocatatropis joyeuxi Kanev & Vassilev, 1986. Occasionally, authors actually working with C. verrucosa ascribed their results to different species. Based on experimental life-cycle studies, the following facts were demonstrated. (1) The first intermediate hosts are the prosobranch freshwater snails Bithynia tentaculata (Linnaeus, 1758) and B. leachi (Leach, 1818). (2) The same snails are also first intermediate hosts for Notocotylus imbricatus (Looss, 1893) Szidat, 1935, N. parviovatus Yamaguti, 1934, and N. ponticus Tschiaberaschvili, 1966. In all these species, the species characteristics are expressed by the adult morphology only, and the larvae cannot be identified by morphological criteria. It is proposed that tri-oculate monostome cercariae found in naturally infected B. tentaculata and B. leachi be referred to as “Cercaria imbricata group”. These cercariae include Cercaria imbricata Looss, 1893, C. helvetica I Dubois, 1928, C. triophthalmia Faust, 1930, C. fennica I Wikgren, 1956; C. ephemera of Lutta (1934); C. monostomi of Mathias (1925), Lutta (1934) and Zdun (1961), Cercaria Notocotylus attenuatus of Francalanci & Manfredini (1969), and Monostome cercaria I Emmel, 1943. (3) There is no second intermediate snail host in the life-cycle of C. verrucosa. (4) The final hosts are birds. (5) The adult worms possess, on the ventral body surface, a median ridge and two lateral rows of 12 (range 11–14) papillae per row.

Argentophilic Structures of the Miracidium of Echinostoma trivolvis (Cort, 1914) (Trematoda: Echinostomatidae)

Argentophilic structures of the miracidium of Echinostoma trivolvis were described from 80 specimens reared from material originally collected in eastern Pennsylvania, U.S.A. Miracidia were impregnated with 0.5% aqueous silver nitrate solution. The miracidium has 18 epidermal plates arranged in 4 rows of 6 + 6 + 4 + 2 = 18. Up to 20 papillalike structures on the terebratorium were arranged along 3 axes and in 5 groups. A single papilla was located at the base of each of the 6 epidermal plates of the first row. The eyespots were located posterior to the first row of plates and 2 excretory pores were located anterior to the last row of plates.