Biology

Endoparasitic Trematodes, as fully developed organisms, occur in vertebrate animals only, with very few exceptions; they inhabit almost all the organs (with the exception of the nervous and osseous systems and the male genitalia), but by preference the intestine in all its extent from the oral cavity to the anus; and, further, certain species or groups inhabit only quite restricted parts of the intestine. Besides in the intestine other species live in the liver, or in the bile-ducts, or in the gall-bladder; other accessory organs of the intestine, such as the pancreas, bursa Fabricii (of birds), are only infected by a few species. Many inhabit the lungs, or the air sacs in fowls, a few the trachea. Trematodes have also been known to occur in the urinary bladder, the urethra and the kidneys of all classes of vertebrates; they are also present in the vascular system of a few tortoises, birds and mammals; in birds they even penetrate from the cloaca into the oviducts, and are occasionally found enclosed in the laid eggs; one species is known to occur in the cavum tympani and in the Eustachian tube of a mammal (Dugong), another in the frontal sinus of the polecat; several species infest the conjunctival sac under the membrana nictitans of birds, one species even lives in cysts in the skin of song-birds. In an analogous manner the ectoparasitic Trematodes are not entirely confined to the surface of the body or the trachea of the lower vertebrate animals; a few species appear exclusively in the urinary bladder, in the œsophagus, and in the case of sharks in an accessory gland of the rectum.

Trematodes live free and active within the organs attacked, though they may attach themselves by suction for a longer or shorter period; in other cases, however, they bore more or less deeply into the intestinal wall with their anterior end, or lie in cysts of the intestinal wall which only communicate with the lumen through a small opening; in those species living in the lungs of mammals the host likewise produces a cyst, which usually encloses two specimens; such association of a pair is also observed in other situations, and, though this is the rule in species sexually distinct, it is not entirely confined to these.

As regards the AGE attained by endoparasitic Trematodes, there are but few reliable records, and these differ considerably; the overwhelming majority of species certainly live about a year, or perhaps a little longer, but there are some whose term of life extends to several or many years.

Trematodes are but rarely found encysted in the higher vertebrate animals; the condition, however, is more frequent in amphibians, and especially in fishes, as well as in numerous invertebrate animals.

Classification of the Trematodes of Man.

The following classification, partly artificial, partly natural, embraces only the flukes found in man:—

Order. Digenea, v. Beneden, 1858.

Anterior sucker single and median, present. Eggs few. The (specialized) terminal portion of the uterus serves as a vagina. Development indirect, i.e., an intermediate host is required.

Sub-order. Prostomata, Odhner, 1905.

Mouth surrounded by the anterior sucker.

Group. Amphistomata, Rudolphi, 1801, ep., Nitzsch, 1819.

Gut forked, two suckers, the posterior sucker (acetabulum) terminal or ventro-terminal behind the genitalia, or at most embraced by the vitellaria. Skin with no spines. Excretory bladder a simple sac opening dorsally near hind end. Testes in front of ovary. Genital pore, median in anterior third of body. Thick flukes, almost circular in cross section.

Family. Paramphistomidæ, Fischoeder, 1901.

Amphistomata: Body not divided into a conical anterior portion and disc-like caudal portion. Ventral pouch absent.

Sub-family. Paramphistominæ, Fisch., 1901.

Paramphistomidæ: Oral sucker without evaginations. Not in man.

Sub-family. Cladorchiinæ, Fisch., 1901.

Paramphistomidæ: Oral sucker with evaginations; testes, two, deeply cleft (fig. ). Genera: Watsonius, Cladorchis, etc.

Family. Gastrodisciidæ, Stiles and Goldberger, 1910.

Amphistomata: With body divided into a conical cephalic and disc-like caudal portion (fig. ). Posterior sucker ventro-terminal. Oral sucker with evaginations. Genera: Gastrodiscus and Homalogaster.

Group. Distomata, Retzius, 1782.

Gut forked, two suckers, the posterior sucker (acetabulum) ventral. It is always separated from the hind end by at least a part of the genitalia.

Family. Fasciolidæ, Railliet, 1895.

Large flat forms, genital pore in front of ventral sucker, the latter powerful. Vitellariæ of numerous follicles, united by branching vitellarian ducts, at the sides of the body meeting posteriorly and extending ventrally and dorsally. Cirrus and vagina without spines. No crown of strong spines around sucker. Testes much branched. Uterus not well developed. Excretory bladder much branched. Eggs large.

Sub-family. Fasciolinæ, Odhner, 1910.

Large or median forms, gut much branched. Body has a shoulder separating head from body. Receptaculum seminis absent. Ovary branched, ventral sucker in anterior part of body. Genus: Fasciola.

Sub-family. Fasciolopsinæ, Odhner, 1910.

Shoulder absent. Receptaculum seminis present. Ovary branched, gut takes a zig-zag course with kinks on it, ventral sucker in anterior part of body. Genus: Fasciolopsis.

Family. Opisthorchiidæ, Braun, 1901, emend. auctor.

Ovary in front of testes. Small to medium flukes, very transparent, tapering anteriorly. Vitellaria moderately developed not extending in front of sucker. Cirrus absent. Seminal vesicle a twisted tube free in parenchyma. Testes near hind end one behind the other, lobed or branched, but not dendritically. Excretory bladder Y-shaped, the two limbs short, the stem S-shaped passing between the testes. Receptaculum seminis well developed. Laurer’s canal present. Uterine coils transverse, numerous. Eggs small.

Sub-family. Opisthorchiinæ, Looss, 1899, emend. auctor.

Opisthorchiidæ in which the excretory pore is terminal. Excretory bladder long, dorsal to testes. Uterine coils not overlapping gut forks. Genera: Opisthorchis, Paropisthorchis, Clonorchis, Amphimerus, etc.

Sub-family. Metorchiinæ, Lühe, 1909.

Opisthorchiidæ in which the excretory pore is ventral. Excretory bladder short, ventral to testes. Uterine coils partly overlapping gut forks and extend anteriorly beyond the sucker. Vitellaria compressed on the sides of the body. Genus: Metorchis.

Family. Dicrocœliidæ, Odhner, 1910.

Ovary behind testes. Testes behind the ventral sucker, between it and the ovary. Body thin and transparent. Cirrus sac encloses the pars prostatica and seminal vesicle. Skin smooth. Gut forks do not reach posterior end. Receptaculum seminis and Laurer’s canal present. Vitellaria, moderate, lateral in mid-body slightly overlapping the gut. Uterus with an ascending and descending branch and numerous transverse coils extending to hind end. Eggs dark brown, 25 µ to 60 µ. Excretory bladder tubular in posterior third or half of body. Parasitic in bile-ducts of mammals and birds. Genus: Dicrocœlium.

Family. Heterophyiidæ, Odhner, 1914.

Ovary in front of testes. Genital pore behind or on a level with ventral sucker. Genital pore surrounded by a pseudo-sucker (i.e., its muscle is not sharply separated from but blends with the body muscles). Cirrus sac absent, consequently vesicula seminalis and pars prostatica lie free. Vagina and ejaculatory duct unite into a common duct before opening. Small and very small forms. Body covered with scales. Genera: Heterophyes, Metagonimus, etc.

Family. Troglotremidæ, Odhner, 1914.

More or less flattened Distomes of compact form, 2 to 13 mm. long. Ventral surface flat or somewhat hollowed, dorsal surface arched. Skin completely covered with pointed spines. Musculature weakly developed also in the suckers in those forms that inhabit cysts. Gut with pharynx and a not very long œsophagus and cæca, which end more or less shortly before the hind end. Excretory bladder Y-shaped or tubular. Pars prostatica and seminal vesicle always distinct. Testes elongated, symmetrically placed in or behind the middle of the body. Ovary directly in front of the testes, right-sided, generally much lobed. Receptaculum seminis and Laurer’s canal present. Vitellaria generally well developed, exclusively or for the most part confined to the dorsal surface, leaving only a median band unoccupied. Uterus either very long, coiling here and there, or shorter and more convoluted. Eggs in first case small 17 µ to 25 µ, in the second much larger 63 µ to 85 µ or even 120 µ (?) long. Parasitic in carnivora or birds, generally occurring in pairs in cyst-like cavities. Genera: Paragonimus, Pholeter, Collyriclum, Troglotrema.

Family. Echinostomidæ, Looss, 1902.

More or less elongated flukes, small or very large, much flattened anteriorly, less so posteriorly, or even round. Suckers near one another, the anterior small and weak, the posterior large and powerful directed obliquely backwards. Surrounding the oral sucker dorsally and laterally but not ventrally is a fold or “collar” bearing a row or rows of pointed spines which are continued round laterally on to the ventral corners, the number being constant for each species, the corner spines large or specialized, skin anteriorly scaled or spiny. Alimentary canal consists of a pharynx, epithelial “pseudo-œsophagus” and gut cæca reaching to posterior end. Testes behind one another in hind body. Ovary on right side or median directly in front of the testes. Vitellaria lateral, usually extending to the hind end and not beyond the ventral sucker anteriorly. Genital pore just in front of ventral sucker. Uterus in transverse loops. Genital sinus absent or present. Receptaculum seminis and Laurer’s canal present. Eggs thin shelled and large, bright yellow, 65 µ to 120 µ long. Excretory bladder Y-shaped. Parasitic in gut of vertebrates, especially birds.

Sub-family. Echinostominæ, Looss, 1899.

Cirrus sac usually reaching to centre of ventral sucker, but not beyond. Cirrus long, usually without spines, coiled when retracted. Seminal vesicle tubular, twisted. On the head a ventral uniting ridge between the angles of the collar. Dorsal circlet of spines, single or double, not interrupted unless the collar itself is dorsally divided. Genera: Echinostoma, etc.

Sub-family. Himasthlinæ, Odhner, 1910.

Cirrus sac reaching far beyond ventral sucker. Cirrus armed with strong rose-thorn-shaped hooks. Vesicula seminalis tubular not coiled. Cervical collar not continued across ventral aspect. Spines on collar in one row. Body armed with fine needle-shaped spines.

Family. Schistosomidæ, Looss, 1899.

Sexes separate. Genital pore behind the ventral sucker. Ventral sucker elevated above the surface. Pharynx absent. Gut forks reunite to form a single stem. In ♂ four or more testicular follicles. In ♀ a single ovary, just in front of the union of the gut forks. Vitellaria on either side of the united gut stem.

The Trematodes Observed in Man.

Family. Paramphistomidæ, Stiles and Goldberger, emend. 1910.

Sub-family. Cladorchiinæ, Fisch., 1901.

Genus. Watsonius, Stiles and Goldberger, 1910.

Cladorchinæ.—Body pyriform. Ventral pouch absent. Acetabulum ventral or (?) ventro-subterminal, very large, margins projecting, aperture small. Genital pore in front of bifurcation of gut, not surrounded by a sucker; ductus hermaphroditicus apparently absent. Excretory pore at posterior end of excretory vesicle, behind Laurer’s canal. Oral sucker with a pair of irregularly globular suctorial pouches; œsophagus thickened distally; cæca long, not wavy; end in acetabular region.

Male Organs.—Testes two lobed, smaller than acetabulum; longitudinally, nearly or quite coinciding; transversely they abut or slightly overlap; preovarial in equatorial and caudal thirds. Pars musculosa not largely developed; cirrus pouch absent.

Female Organs.—Ovary and shell gland post-testicular. Vitellaria extend from gut fork to slightly beyond gut ending; uterus intercæcal, partly post-testicular. Laurer’s canal in front of excretory vesicle.

Type Species.—Watsonius watsoni, Conyngham, 1904.

Watsonius watsoni, Stiles and Goldberger, 1910.

Syn.: Amphistomum watsoni, Conyngham, 1904; Cladorchis watsoni, Shipley, 1905.

Body, 8 to 10 mm. long, by 4 to 5 mm. broad, by 4 mm. thick; tapers anteriorly to 2·5 mm. Caudal extremity bluntly rounded, venter surrounded by an elevated ridge, surface with transverse ridges best defined ventrally. Genital pore median about one-quarter of body length from anterior end at level of suctorial pouches. Acetabulum 1 mm. in diameter, margin projecting, aperture small. Mouth in a groove with digitate papillæ. Oral sucker very large, one-fifth of length of body, with a pair of irregularly globular pouches. Œsophagus somewhat longer than sucker. Excretory pore at the level of the acetabular aperture. The vesicle extends from the plane of the transverse vitelline ducts to centre of acetabulum.

Male Organs.—Testes deeply notched adjoining one another. Vesicula seminalis much coiled and dilated, pars musculosa not coiled. Pars prostatica (?) dilated, ejaculatory duct long and narrow, opening on a papilla; genital atrium papillated.

Female Organs.—Ovary dorso-posterior of posterior testis. Shell gland dorsal to ovary. Vitellaria ventral and lateral to gut cæca extending from gut fork to equator of acetabulum. Uterus dorsal to testes, ductus hermaphroditicus absent. Laurer’s canal opens in dorso-median line slightly behind anterior border of sucker.

Eggs.—123 µ to 133 µ long by 75 µ to 80 µ broad.

Habitat.—Jejunum and duodenum of man, German West Africa. The parasite has only been found once in man. The patient, a negro from German West Africa, died at Zola, Northern Nigeria. The symptoms were persistent watery diarrhœa without blood or mucus. The parasites were also passed in the stools. It occurs also in monkeys.

Family. Gastrodisciidæ.

Genus. Gastrodiscus, Lkt., 1877.

Acetabulum small, caudal and ventral margin raised, aperture relatively large. Genital pore without sucker. Excretory pore post-vesicular, posterior to opening of Laurer’s canal. Œsophagus with muscular thickening; cæca not wavy, long, end post-equatorial and post-testicular.

Male Genitalia.—Testes two, branched pre-ovarial.

Female genitalia.—Ovary and shell gland post-testicular. Vitellaria extracæcal; uterus intercæcal; Laurer’s canal entirely prevesicular.

Type.—Gastrodiscus ægyptiacus, Cobbold, 1876.

Gastrodiscus hominis, Lewis and McConnell, 1876.

Syn.: Amphistomum hominis, Lew. and McConn.

Body, reddish in the fresh, 5 to 8 mm. long; posteriorly, 3 to 4 mm. broad. The disc has incurved edges which are interrupted in front where it joins the anterior cylindrical portion and posteriorly behind the ventral sucker. The disc itself and ventral surface are covered with a number of (microscopic) papillæ. Pharynx provided with two diverticula or pouches. The bifurcation of the gut lies sometimes above, sometimes below the level of the genital pore. The gut cæca end about the level of the centre of the acetabulum.

Genital Pore.—About the middle of the conical anterior portion. (It appears to be surrounded by a muscular sucker.) Leiper (1913) describes the ducts as discharging at the tip of a large fleshy papilla, the surface of which bears cuticular bosses.

Testes much lobed, the anterior is smaller than the posterior and lies at about the level where the anterior conical portion joins the disc. The posterior testis just in front of the anterior margin of the acetabulum separated from it by the ovary. The ovary, somewhat oval in shape or slightly constricted in the middle, lies slightly to the right of the median line. Dorsal to it lies the well-developed shell gland, Laurer’s canal opening in front of the excretory bladder. The excretory bladder is a long sac with its opening at its posterior extremity about the level of the middle of the acetabulum. The vitellaria are restricted in extent. They do not extend forward beyond the anterior border of the posterior testis. They are best developed in the area between the acetabulum and the termination of the gut cæca.

The eggs are oval and measure 150 µ in length by 72 µ in breadth.

Habitat.—Cæcum and large intestine of man. Also in the pig (5 per cent.) in Annam.

Distribution.—This parasite has been recorded from Assam (not uncommon), British Guiana (Indian immigrants), and Cochin China.

Gastrodiscus ægyptiacus, Cobbold, 1876, and G. secundus, Looss, 1907, occur in the horse; G. minor, Leiper, 1913, in the pig in Nigeria and Uganda.

Family. Fasciolidæ, Raill., 1895.

Sub-family. Fasciolinæ, Odhner, 1910.

Genus. Fasciola, L., 1758.

The ventral sucker is situated at the level of the junction of the cone with the body, viz., at the level of the “shoulder,” and is large and powerful. The cuticle is covered with strong spines; the gut cæca run in the mid-line to the hind end, and are provided with numerous long lateral and fewer and shorter median branches. The ovary lies on one side in front of the transverse vitelline duct; the testes lie obliquely one behind the other. The uterus, in the shape of a rosette, lies in front of the genitalia. Laurer’s canal is present; the vesicula seminalis lies in the cirrus pouch; the ova are large, not very numerous, and only develop after they have been deposited. Parasites of the biliary ducts of herbivorous animals.

Fasciola hepatica, L., 1758.

Syn.: Distomum hepaticum, Retz., 1786; Fasciola Humana, Gmel., 1789; Distomum caviæ, Sons., 1890; Cladocœlium hepaticum, Stoss., 1892.

Length 20 to 30 mm., breadth 8 to 13 mm., cephalic cone 4 to 5 mm. in length and sharply differentiated from the body by a shoulder on each side. Spines in alternating transverse rows and extending on the ventral surface to the posterior border of the testes, and on the dorsal surface not quite so far. The spines are smaller on the cephalic cone than on the posterior part of the body, where they are discernible with the naked eye. The suckers are hemispherical, and near each other; the oral sucker is about 1 mm. and the ventral sucker about 1·6 mm. in diameter. The pharynx, which includes almost the entire œsophagus, measures 0·7 mm. in length and 0·4 mm. in breadth. The intestine bifurcates at the limit of the cephalic cone and the branches are even here furnished with diverticula directed outwardly. The ovary is ramified and situated in front of the transverse vitelline duct, usually on the right side; the shell gland lies near the ovary in the median line; posterior to the transverse vitelline ducts are the greatly ramified testes, which occupy the greater portion of the posterior part of the body, with the exception of the lateral and posterior border; the long vasa efferentia only unite as they enter the cirrus pouch. The vitellaria occupy the sides of the posterior part of the body, commencing at the level of the ventral sucker and uniting behind the testes. The ova are yellowish-brown, oval, operculated, 130 µ to 145 µ in length, 70 µ to 90 µ in breadth (average size 132 µ by 70 µ).

The Liver Fluke inhabits the bile-ducts of numerous herbivorous mammals (sheep, ox, goat, horse, ass, rabbit, guinea-pig, squirrel, beaver, deer, roe, antelope, camel, kangaroo, and others), and is distributed over the whole of Europe, though not to an equal extent. It is further known in North Africa, in North and South America, as well as in Australia; it is also found in Asia, as it has been reported from Japan, China, and Tonkin (Gaide, two cases in man). In some districts of Germany it is very frequent, and the slaughter-house statistics of various places show that it is of daily occurrence. Fasciola magna occurs in herbivora in America.

The liver fluke, however, is by no means a harmless parasite, for it produces in domestic animals, more especially in sheep, a disease of the liver that appears epidemically in certain years and districts, and commits great ravages amongst the flocks.

[The following records show the enormous loss caused in sheep by this parasite. In 1812, in the Midi, principally in the Departments of the Rhône, Herault, and Gard, the disease was rampant; 300,000 sheep perished in the Arles territory, and 90,000 in the Arrondissements of Nîmes and Montpellier. In 1829 and 1830, in the Department of the Meuse and near localities, not only sheep but oxen died in enormous numbers; for instance, in the Arrondissement of Verdun out of 50,000 sheep 20,000 died, and out of 20,000 cattle 2,200 died. In England, in 1830, 2,000,000 sheep were carried off; whilst in 1862 60 per cent. of the sheep died in Ireland; and in 1879 over 300,000 were lost in England; whilst as late as 1891 one owner in the same country lost over 10,000 sheep (Live Stock Journal, October 30, 1891).—F. V. T.]

The disease usually commences towards the end of summer with an enlargement of the liver, induced by the invasion of numerous young flukes; in the autumn and winter the animals suffer from the consequences of disordered biliary secretion; they become feverish, emaciated, and anæmic, and lose their appetite. In consequence of the consecutive atrophy of the liver, œdema and ascites set in, and many animals succumb to this “liver rot.” On examination the liver is found to be shrunken, the bile-ducts are enormously dilated and in parts saccular and full of flukes. Should the animals survive this stage, spontaneous recovery ensues in consequence of the flukes commencing to leave the liver in the spring, but the liver remains changed and its sale is prohibited when the changes are extensive.

[The following stages may be noticed in sheep suffering from fascioliasis. Gerlach recognized four stages, based on the varied relations that the flukes contract with the liver of their host. These periods are sometimes very marked, but at others, owing to subsequent infections, the features become merged and so obliterated. But when a single infestation occurs they are very marked.

[The first period is called the PERIOD OF IMMIGRATION. This occurs at the fall of the year and generally passes unperceived, as the young flukes do little harm to the liver. It varies from four to thirteen weeks. Gerlach has remarked upon cases of death from apoplexy at this period.

[The second period is the PERIOD OF ANÆMIA. This occurs in November and December. The sheep at first fatten rapidly, but later the mucous membranes become pale and of a yellowish hue, and the sheep become sluggish and cease to feed. The fæces are normal, but may contain fluke ova.

[The third period is the PERIOD OF WASTING. This corresponds with the beginning of January—about three months after the entry of the larvæ. Emaciation now becomes very marked, the skin and mucous membranes blanched, temperature variable and marked by an irregular curve; respiration laboured and quick; appetite regular; abortion frequently occurs in pregnant ewes; pressure on the back causes the animals to fall; local œdemas occur, the most perceptible in the submaxillary space, extending below the larynx and over the cheeks and parotids (called “bourse,” “boule” in France; “watery poke” or “cockered” in England). Death usually occurs at this period, but a fourth stage may occur.

[The fourth period is the PERIOD OF MIGRATION OF THE FLUKES. This is a period of convalescence and recovery, generally in May and June.—F. V. T.]

Oxen suffer less in general, but even in these animals “stray” hepatic flukes are occasionally found in the lungs, enclosed in thick-walled cysts.

Pathological Anatomy.—The bile-ducts are conspicuous on the surface of the liver. They are thickened and much dilated and in parts saccular, and considerable atrophy of the liver cells accompanies the condition. Histologically there is immense proliferation of the epithelium of the bile-ducts leading to “adenomata.”

The LIFE-HISTORY of the liver fluke was discovered by R. Leuckart and P. Thomas. According to these investigators the elongated miracidium (fig. , a) ciliated all over develops from the eggs a few weeks after the latter (fig. 142) have reached the water, and after it has become free the embryo penetrates and becomes a sporocyst (fig. , b) in a water-snail (Limnæus truncatulus, Müll. = L. minutus, Drap.) that is common in fresh water, and can live in the smallest collection of water as well as in fields that have been flooded. The sporocyst first of all produces rediæ, which remain in the same host (and under certain circumstances, e.g. in summer, these develop a second generation of rediæ), and these finally form cercariæ (fig. ). The latter become encysted on blades of grass and are taken up by the respective hosts with their food; this takes place towards the end of summer, while the sheep feeding on the pasture land in the spring spread the eggs of the fluke, and sometimes the fluke itself, by passing them with their fæces.

In districts where Limnæus truncatulus is absent, analogous species act as the intermediary hosts, of which one example according to Lutz is Limnæus oahuensis in the Sandwich Islands.

[The host in Europe is Limnæus truncatulus. This snail extends from Siberia to Sicily and Algeria, and according to Captain Hutton is a native of Afghanistan. It also occurs in Thibet, Amoor, Morocco, Tunis, Canary Islands and the Faroe Islands. It deposits its eggs or spawn upon the mud around ponds, ditches and streams. The eggs are laid in batches of thirty to a hundred, each snail laying as many as 1,500 eggs; they are united into strips of a gelatinous substance. In about two weeks young snails appear. It is amphibious, being more frequently met with out of the water than in it. It occurs in elevated spots as well as in low-lying districts. Moquin-Tandon found it at 4,000 feet in the Pyrenees. In the allied species, L. peregra, the fluke will develop up to a certain stage, but never completes all its varied phases.

[In South America the host is probably Limnæus viator, Orb., and in North America Limnæus humilis, Say.—F. V. T.]

In human beings as well as in some of the mammals quoted above, the liver fluke is only a casual parasite, and hitherto only twenty-eight cases have been observed in man; the infection was mostly a mild one and there were no symptoms, or only very trifling ones; a few isolated cases were only discovered post mortem. Occasionally, however, even when the infection was inconsiderable, severe symptoms were set up, which in isolated cases led to death. The symptoms (enlargement and painfulness of the liver, icterus) merely pointed to a disease of the liver.

Diagnosis can only be established by finding eggs in the fæces. Care should be taken not to confuse them with those of Dibothriocephalus latus.

Halzoun.

In North Lebanon, the liver fluke is, according to A. Khouri, a frequent parasite of man, not in the liver, however, but in the pharynx. The occurrence in this unusual site is effected by the eating of raw infected livers, especially those of goats (Capra hircus). The flukes thus taken in do not all reach the stomach, where they would be soon killed, but some of them attach themselves to the pharyngeal mucosa and to the adjoining parts, and there cause inflammation and swelling, which lead to dyspnœa, dysphagia, dysphonia and congestion of the head, sometimes even to still more severe symptoms, and even death. The affection termed “Halzoun” lasts some hours or several days, and after vomiting recovery sets in. In other cases man becomes infected in the usual way by ingesting cysts attached to grass or the underside of leaves of plants (e.g., Rumex sp.), where they are overlooked from their scanty size (0·2 to 0·3 mm.).

As the liver fluke feeds on blood it is possible that it also reaches, particularly when young, the circulatory system, and cases have been known in which it has been carried by the blood into organs far from its original situation. Such cases also have been repeatedly observed in men. Probably the parasite described by Treutler, 1793, as Hexathyridium venarum, which protruded from the ruptured anterior tibial vein of a man, was a young liver fluke. A few adult specimens were found by Duval in the portal and other veins post mortem at Rennes (1842) in a man, aged 49, and a similar statement is reported by Vital from Constantine (1874). Giesker, in 1850, found two hepatic flukes in a swelling on the sole of the foot of a woman. Penn Harris states that he observed six specimens in Liverpool in a spontaneously ruptured abscess of the occiput of a two months old infant. Another case which, like the previous one, is reported by Lankester, relates to a sailor who suffered from an abscess behind the ear, and from which a liver fluke was expelled. Finally, Dionis de Carrières reports the case of a man, aged 35, in whose right hypochondriac region a tumour the size of a pigeon’s egg had formed, and from which a young liver fluke was extracted.

From such records it is not impossible that Distomum oculi humani, Ammon, 1833, as well as Monostomum lentis, v. Nordm., 1832, may have been very young hepatic flukes that had strayed. Ammon found four specimens (length 0·5 to 1 mm.) of his species (named Distomum ophthalmobium by Diesing in 1850) between the opaque lens and the capsule of a five months old child in Dresden, and von Nordmann discovered his Monostomum lentis to the number of eight specimens (only 0·3 mm. in length) in the opaque lens of an old woman. Minute white bodies which Greef found in the cortex of the lens of a fisherman, aged 55, removed on account of cataract, were with some reserve regarded as Trematode larvæ. The fact that Ammon found that the intestinal cæca of the worm discovered by him had no lateral branches does not negative the above opinion, as in the liver fluke the intestinal cæca are originally unbranched, and according to Lutz they only develop lateral ramifications later, between the twelfth and twenty-second day of infection (fig. 144).

Fasciola gigantica, Cobbold, 1856.

Syn.: Distomum giganteum, Diesing, 1858; Fasciola gigantea, Cobbold, 1858; Cladocœlium giganteum, Stoss., 1892; Fasciola hepatica var. angusta, Raill., 1895; Fasciola hepatica var. ægyptiaca, Looss, 1896.

This species is closely allied to Fasciola hepatica, but is distinguished by its elongated body, short cephalic cone, almost parallel sides, larger ventral sucker, which is also closer to the oral sucker, and by its larger eggs. Length up to 75 mm., width up to 12 mm. Oral sucker 1 to 1·2 mm., ventral sucker up to 1·7 mm. in diameter. Eggs 150 µ to 190 µ long by 75 µ to 90 µ broad.

Habitat.—Bile-ducts of Camelopardalis giraffa, Bos taurus, Bos indicus, Bos bubalis, Ovis aries and Capra hircus.

Distribution.—Africa.

This species has once been observed in man by Gouvea, in Rio de Janeiro, in a French naval officer who became ill with fever, cough and slight blood-spitting. The lungs were normal except for a sharply circumscribed spot at the base of the left lung. Twenty days later during a fit of coughing the patient spat up a fluke 25 mm. long, characterized by its slender aspect and by the size of its ventral sucker, and its close proximity to the oral sucker. Considering the fact that Gouvea’s patient had spent many weeks in July of the same year in Dakar (Senegambia), where according to Railliet Fasciola gigantica is common in slaughtered animals, and considering also the characters of the fluke, Railliet rightly assumes that one had to do with the African giant fluke and that the patient had infected himself in Dakar.

Sub-family. Fasciolopsinæ, Odhner, 1910.

Genus. Fasciolopsis, Looss, 1898.

Ventral sucker large, and elongated posteriorly into a sac. Cirrus pouch long and cylindrical, its greatest length being occupied by the sinuous tubular seminal vesicle, on which exists a peculiar cæcal appendage. Laurer’s canal present.

Fasciolopsis buski, Lank., 1857.

Syn.: Distomum buski, Lank., 1857; Dist. crassum, Cobbold, 1860, nec v. Sieb., 1836.

The length of the body varies; it may measure 24 to 37 or even attain 70 mm.; the breadth is from 5·5 to 12 to 14 mm. In the pig the fresh parasites measure, smallest, 12 to 8 mm.; largest, 35 to 16 mm. (Mathis and Léger). Skin without spines, but according to Heanly always present in man and pig specimens. The oral sucker measures 0·5 mm. in diameter; the ventral sucker is three to four times as large; the pharynx is globular, 0·7 mm. in diameter; the prepharynx is provided with a sphincter; the intestinal cæca extend to the posterior border with two characteristic curves, one at the anterior border of the anterior testis, the other between the two testes. The genital pore is at the anterior border of the ventral sucker; the cylindrical cirrus pouch extends from behind the ventral sucker to half-way to the shell gland. The seminal vesicle extends forwards within the cirrus pouch as a convoluted tube. From its anterior portion is given off the cæcal appendage, which has itself short lateral diverticula. It runs backwards, ending blindly about 0·5 mm. from the posterior end of the cirrus sac. The seminal vesicle is continued as the pars prostatica (?) 0·5 mm. long, and this by the very short ejaculatory duct (13 µ), and finally by the fairly long cirrus, which is beset with very fine spines except at either extremity. The ovary and shell gland are situated at about the middle of the body with the testes behind them, and the uterus in front. The vitellaria extend from the ventral sucker to the posterior border. The eggs measure 120 µ to 130 µ in length and 77 µ to 80 µ in breadth, and resemble those of Echinochasma sp. in dogs. The larval stages are said to occur in shrimps.

Habitat.—Intestine of pig and man.

Distribution.—In man: India, Siam, China, Assam, Sumatra. It is common in Cochin China (16 out of 133 Annamites, Noc.), in Tonkin very rare. Dr. J. Bell has sent me [J. W. W. S.] human specimens from Hong Kong. In pigs: very common in South China (Heanly). Common in pigs in Hong Kong. Sixteen out of 248 pigs (i.e., 6 per cent.) infected in Hanoi.

Fasciolopsis rathouisi, Ward, 1903.

Syn.: Distomum rathouisi, Poirier, 1887.

Fifteen to 19 mm. long by 8·5 to 10·5 mm. broad by about 3 mm. thick. Skin with spines (Leiper). Bluntly oval or elliptical with short cephalic cone which is absent in Fasciolopsis buski. Oral sucker subterminal, 0·25 to 0·29 mm. broad by 0·2 mm. in antero-posterior diameter. Distant from ventral sucker by about twice its diameter. Ventral sucker 1·32 to 1·38 mm. broad by 0·68 to 0·7 mm. in antero-posterior diameter. Œsophagus extremely short. Cirrus sac not conspicuous and straight as in Fasciolopsis buski, but is convoluted. Testes one behind the other (according to Poirier they lie beside one another), more compactly branched, broader and denser than in Fasciolopsis buski. Ovary on right side, small, coarsely branched. Uterus in broad, closely grouped coils, packed with ova anterior to ovary. Vitellarian acini more numerous and somewhat differently distributed. Eggs 150 µ by 80 µ, thin shelled. [H. B. Ward, who has examined this species, and from whose account the above is mainly taken, considers that it is a good species, although the differences between it and Fasciolopsis buski are slight, while Odhner, who examined the original species, is of the opposite opinion.—J. W. W. S.] The parasite appears to cause diarrhœa, wasting and occasionally jaundice.

Habitat.—Intestine of man.

Distribution.—China, common in some parts (Goddard).

Fasciolopsis goddardi, Ward, 1910.

Twenty-one to 22 mm. long, 9 mm. broad. Skin with spines (Leiper). Uterus very closely coiled, most striking character is the large size of the vitelline acini. Imperfectly known.

Distribution.—China (Shanghai).

Fasciolopsis fülleborni, Rodenwaldt, 1909.

The fully extended fluke is tongue-shaped, 50 by 14 mm.; two contracted specimens measured 40 by 15 mm. and 30 by 16 mm. respectively. Skin without spines, with according to Leiper cephalic cone not clearly defined. Oral sucker circular, 0·75 mm. in diameter, slightly larger than that of Fasciolopsis buski. Ventral sucker 2·6 mm. in diameter (that of Fasciolopsis buski 1·6 to 2 mm.). Length 2·9 mm. (as in Fasciolopsis rathouisi), the excess of length over breadth being due to the posterior elongated sac-like prolongation of the sucker. Prepharyngeal sphincter present. Pharynx 0·7 mm. in diameter. Œsophagus practically absent. Gut cæca similar to those of Fasciolopsis buski.

Testes—regularly branched, separated by an incurving of the cæca, the anterior occupying a smaller area than the posterior.

Ovary—very small, as in Fasciolopsis buski, on the right side.

Shell Gland—almond-shaped, 2·3 by 1·2 mm. In Fasciolopsis buski it is round and smaller, 1 to 1·5 mm. in diameter.

Vitellaria—similar in distribution to those of Fasciolopsis buski, but the acini are strikingly small.

Cirrus Sac—is the most characteristic feature of this species. It is a powerfully built, convoluted sac standing out clearly on the body. It is not a uniform, straight cylinder 0·25 to 0·33 mm. in diameter, as in Fasciolopsis buski, but even in fully extended flukes is typically convoluted. It is 1 mm. thick in the middle, but in other parts varies much from this. The posterior end of the cirrus sac is at two-thirds or more of the distance from ventral sucker to shell gland. In the case of Fasciolopsis buski the posterior end of the sac only extends half-way.

Seminal Vesicle—has a peculiar convoluted, saccular and angular course, but the cæcal appendage characteristic of the genus appears to be absent!

Excretory System.—The main stem gives off very regular transverse branches which are well seen posteriorly.

Eggs.—100 µ by 73 µ. Thin shelled.

Habitat.—Intestine. Mahommedan from Calcutta.

[It is evident that a re-examination of fresh material is required before the validity of all these species can be accepted.—J. W. W. S.]

Family. Troglotremidæ, Odhner, 1914.

Genus. Paragonimus, Braun, 1899.

Body egg-shaped or somewhat elongated, generally more broadly rounded in front than behind. Covered all over with spear-shaped spines arranged in groups. Gut cæca winding with dilatations or constrictions in parts. Ventral sucker in or in front of the middle of the body. Excretory bladder cylindrical, very long and broad, reaching in front to the bifurcation of the gut. The lateral excretory canals join the bladder only a little in front of the excretory pore. Genital pore median just behind the ventral sucker. Genital sinus duct-like. Cirrus sac absent. Male terminal organs very small. Ejaculatory duct present. Testes and ovary deeply lobed, the testes in or just behind the middle, the ovary somewhat laterally placed just behind the ventral sucker. Uterus forms a coil behind the ventral sucker. Eggs rather large, thin shelled, the ovarian cell still unsegmented on deposition. Receptaculum seminis, small.

Parasitic in the lungs of mammals, enclosed in cyst-like cavities, generally in pairs.

Type Species.—P. westermanii in the tiger.

Paragonimus ringeri, Cobb., 1880.

Syn.: Distoma ringeri, Cobb., 1880; Distoma pulmonale, Baelz, 1883; Distoma pulmonis, Suga, 1883.

The body is of a faint reddish-brown colour and plump oval shape. The ventral surface a little flattened; 7·5 to 12 mm. in length, 4 to 6 mm. in breadth, and 3·5 to 5 mm. thick (in man). The oral sucker (0·75 mm.) is subterminal; the ventral sucker (0·8 mm.) somewhat in front of the middle of the body. Pharynx spherical, 0·3 mm. in diameter, or 0·4 by 0·3 mm.; œsophagus, 0·02 mm.; intestinal cæca convoluted, asymmetrical, the first part having the same structure as the œsophagus. The cuticle is covered with spines in groups; the excretory pore opens at the posterior end rather on the ventral surface, the excretory ducts open into the elongated bladder at the hind end near the pore. Genital pore behind the ventral sucker and median. Genital sinus 0·2 mm. long with thick wall, ejaculatory duct 0·13 mm., pars prostatica 0·2 mm., seminal vesicle duct-like of irregular outline. Behind the sucker the ovary on the left, and the closely packed uterine coil on the right (though amphitypy of these two organs is common); the two irregularly lobed testes lie side by side posteriorly. Vitellaria extensive, leaving only a median dorsal and ventral space free. Seminal receptacle probably absent; Laurer’s canal present. The eggs are oval, brownish-yellow, fairly thin shelled, and measure on an average 81·2 µ by 49·2 µ.

The following species are also known:—P. westermanii, Kerb., 1878, in the tiger, and P. kellicotti, Ward, 1908, in the pig, dog, and cat (N. America). Ward and Hirsch give the following differences between the spines of the three forms:—

P. ringeri.

P. westermanii.

P. kellicotti.

Shape Chisel-shaped, mod-
erately heavy. Lancet-shaped,
very slender. Chisel-shaped,
heavy.

Distribution Circular rows, in
groups. Circular rows,
in groups. Circular rows,
singly.

Two other species, P. rudis, Diesing, 1850, in a Brazilian otter (Lutra brasiliensis) and P. compactus, Cobbold, 1859, in the Indian ichneumon, are but little known.

Habitat.—Lungs, pleuræ, and especially the bronchi of man and dog. The alleged occurrence (of eggs) in other organs may be due to confusion with those of Schistosoma japonicum.

Distribution.—China, Korea, and especially in Japan, where, according to Katsurada, there are no districts that are entirely free from pulmonary flukes. The mountainous provinces of Okayama, Kumamoto, Nagano and Tokushima are the principal centres.

Pathology.—The number present in the lung varies from two to twenty, about. Usually one cyst contains one worm, but in the dog each cyst contains two. The cysts admit the tip of the finger, and have a fibrous wall 1 mm. thick. They originate partly from dilatation of bronchi and bronchioles. Others arise from the inflammatory reaction of lung tissue into which the worms have wandered. The worms and their eggs cause bronchitis and peribronchitis, catarrhal, hæmorrhagic, or purulent, and areas of consolidation. Areas containing eggs in their centre resembling tubercle nodules are not uncommon, and extensive cirrhosis of the lung may be found. As a result of these changes, emphysema and bronchiectasis also occur.

As to the development, only the following details are known: that the eggs, which before segmentation of the ovum reach the open in the sputum and through being swallowed also in the fæces, develop in water into a miracidium ciliated all over, which hatches and swims about freely. According to Manson this takes place in four to six weeks.

Sub-family. Opisthorchiinæ, Looss, 1899.

Genus. Opisthorchis, R. Blanch., 1845.

Opisthorchiinæ with lobed testes. Laurer’s canal present. Parasitic in the bile-ducts of mammals and birds.

Opisthorchis felineus, Riv., 1885.

Syn.: Distoma conus, Gurlt, 1831 (nec Creplin, 1825); Distoma lanceolatum, v. Sieb., 1836, v. Tright, 1889 (nec Mehlis, 1825 = Fasciolo lanceolata, Rud., 1803); Distoma sibiricum, Winogr., 1892; Distoma tenuicolle, Mühl., 1896.

This parasite is yellowish-red in the fresh condition, and almost transparent. The body is flat, with a conical neck at the level of the ventral sucker marked by a shallow constriction; this, however, is only noticeable in fresh and somewhat contracted specimens. Posteriorly to the ventral sucker the lateral borders run fairly parallel; the posterior end is either pointed or rounded off. The length and breadth vary according to the contraction, being usually 8 to 11 mm. by 1·5 to 2 mm. The suckers are about one-fifth to one-sixth of the length of the body distant from each other, and of about equal size (0·23 to 0·25 mm.). The œsophagus is hardly any longer than the pharynx, which lies close behind the oral sucker; the intestinal cæca reach almost to the posterior border and are often filled with blood. The excretory pore is at the posterior extremity, and the excretory bladder forks in front of the anterior testis. The testes in the posterior fourth of the body lie obliquely one behind the other; the anterior one has four lobes, the posterior one five lobes; the ovary is in the median line transversely, simple or slightly lobed; behind it lies the large pear- or retort-shaped receptaculum seminis and Laurer’s canal. The uterus is in the median field. The vitellaria occupy the fairly broad lateral areas, in about the central third of the body, beginning behind the ventral sucker and terminating at about the level of the ovary; the acini are small and arranged in groups of seven to eight, separated by interstices. The genital pore is close in front of the ventral sucker. The eggs are oval with sharply defined operculum at the pointed pole, 30 µ, by 11 µ.

This species, which is frequently confused with others, inhabits the gall-bladder and bile-ducts of the domestic cat especially; but is also found in the dog, in the fox, and in the glutton (Gulo borealis). It has been observed in France, Holland, North Germany (being particularly frequent in East Prussia), in Russia, Scandinavia, Siberia, Japan, Tonkin, Hungary, and Italy. The North American form (from cats and Canis latrans) is a distinct species (Opisthorchis pseudofelineus).

In man this species was first found by Winogradoff in Tomsk (nine cases), then by Kholodkowsky in a peasant from the neighbourhood of Petrograd who had travelled a great deal in Siberia, and finally by Askanazy in five persons who were natives of the East Prussian district of Heydekrug. In Tomsk, Opisthorchis felineus is the most frequent parasite of man that comes under observation at post mortem (6·45 per cent.), whereas Tænia saginata has only been found in 3·2 per cent., Echinococcus in 2·4 per cent., Ascaris lumbricoides in 1·6 per cent., and Oxyuris vermicularis in 0·8 per cent. of the autopsies. In the district of Heydekrug, however, the species in question is also frequent, as in a few years five cases came to our knowledge (of which three were diagnosed by the discovery of the eggs in the fæces).

In none of Winogradoff’s nine cases had the death of the patient been caused direct by the parasites, yet more or less extensive changes in the liver were found in all of them; such as dilatation of the bile-ducts with inflammation and thickening of their walls, and foci of inflammation or atrophy in the liver substance; icterus was present five times and atrophy of the liver an equal number of times; ascites was observed three times, and in two cases, probably of recent date, the organ was enlarged. The number of parasites found fluctuated between a few and several hundreds.

In two of Askanazy’s cases, which he examined more closely, carcinoma which had developed at the places most invaded by flukes was found at the post-mortem, so that perhaps there may be grounds for the connection which the author seeks to establish between cancer of the liver and the changes induced by the parasites; these changes consist of numerous and even ramified proliferations of the epithelium of the biliary duct into the connective tissue, which is likewise proliferated. The number of worms found in one case amounted to over 100; in a second case, in which the parasites had also invaded the pancreatic duct, their number was even larger.

Winogradoff as well as Askanazy found isolated flukes in the intestine also.

Unfortunately, nothing much is known of the history of the development of Opisthorchis felineus; we only know that when deposited the eggs already contain a ciliated miracidium, which, however, according to my experience, does not hatch out in water, but only after the entry of the eggs into the intestine of young Limnæus stagnalis; no further development, however, occurs. Winogradoff states that he has seen the miracidia hatch after the eggs had been kept in water for a month at 37° C.; and has even observed free miracidia in the bile of man and of a dog respectively. Although the whole post-embryonal development of the cat fluke remains yet to be investigated, Askanazy by a series of experiments on cats and dogs has discovered the mode of infection. The intermediate hosts are fish, and mainly the ide, in this country called Tapar (Idus melanotus, H. and Kr.), and of subsidiary importance the roach (Leuciscus rutilus). Both species of fish as well as others are readily eaten raw by man on the Courland lagoon (Baltic). It is, moreover, significant that those persons whom Askanazy found infected with the cat fluke were also infected with Dibothriocephalus latus, the intermediate host of which is also fish (Lota sp., Esox sp., Perca sp.).

In one of his nine cases Winogradoff also saw a small fluke covered all over with spines, which he conjectured to be the young stage of Opisthorchis felineus; as, however, according to my experience, this species, even in smaller specimens, is always without spines, the above hypothesis cannot be accepted. It is much more probable that one of the other species that also invade the liver of cats may accidentally be introduced into man; we know, in fact, that Metorchis albidus, Braun, and Metorchis truncatus, Rud., are both covered with spines. As, however, the spines of the first-named species are rather apt to fall off, and also as it possesses a different shape (spatula-shaped), it may be assumed that probably Winogradoff had found Metorchis truncatus, Rud., 1819, in his patient.

Genus. Paropisthorchis, Stephens, 1912.

Structure as in Opisthorchis, except that the ventral sucker and genital pore occur on the apex of a process or pedicle projecting from the anterior portion of the body. This process is about 1/2 mm. long, and is retractile.

Paropisthorchis caninus, Barker, 1912.

Syn.: Distoma conjunctum, Lewis and Cunningham, 1872; Opisthorchis noverca, M. Braun, 1903 (pro parte); Opisthorchis caninus, Barker, 1912 (?).

Length varies from 2·75 to 5·75 mm. in preserved specimens, average 3·6 to 5·2 mm. Body uniformly spinose, though as a rule spines are not present on the pedicle. Body slightly concavo-convex, the concavity being ventral. Oral sucker 0·28 mm. Pharynx 0·224 by 0·184 mm. Œsophagus 0·04 mm. Ventral sucker 0·176 mm. in diameter. Pedicle about 1/2 mm. long, may be completely retracted.

Genital Pore—opens on the apex of the pedicle in front of the ventral sucker. Its exact position varies with the state of contraction of the parts. In certain cases it actually opens within the cuticular border of the sucker, in other cases it opens externally to the sucker and anterior to it. The opening is covered with scales. The vas deferens and uterus run alongside one another until they merge near the apex of the pedicle into a common sinus.

Vitellaria—consist of eight acini on each side, extending from slightly behind the base of the pedicle to the anterior border of the ovary, or as far back as a line separating the posterior border of the ovary from the anterior border of the anterior testis.

Testes.—Anterior testis 0·496 by 0·44 mm.; posterior testis 0·52 by 0·48 mm., usually ovoid, though both may be regularly lobed. The anterior testis is usually on the left side.

Ovary—multilobular, the lobes 6 to 8 being irregular in size and shape.

Shell Gland—extensive and diffuse, occupying an area which approximately corresponds with the loop of the transverse vitelline ducts.

Seminal Receptacle—globular, to the right of and dorsal to the posterior lobe of the ovary.

Laurer’s Canal—generally runs from the end of the receptacle with a single curve medially and backwards.

Uterine Coils—form loosely packed transverse coils terminating slightly in front of the level of the first vitelline acini. From here the uterus passes forwards into the pedicle to the left and ventral to the seminal vesicle.

Seminal Vesicle—commences about the level of the first vitelline acini. The coils displace the uterus ventrally and to the left. In the pedicle the vesicle diminishes in extent and lies in its dorsal (anterior) side.

Habitat.—Liver of pariah dogs, India. In North-Western Provinces about 40 per cent. are infected. This fluke appears to be different from Amphimerus (Opisthorchis) noverca in man, as the latter has not the pedicle on the summit of which lie the sucker and common genital pore.

Genus. Amphimerus, Barker, 1912 (?).

Structure as in Opisthorchis, except that the vitellaria are separated into two portions, an ant-ovarial and a post-ovarial.

Amphimerus noverca, Barker, 1912 (?).

Syn.: Distomum conjunctum, McConnell, 1876 (nec Cobbold, 1859); Opisthorchis noverca, M. Braun, 1903 pro parte.

At the autopsy of two Mahommedans who died in Calcutta, McConnell found a large number of Distomata in the thickened and dilated bile-ducts. The worms were lancet-shaped, covered with spines, and measured 9·5 to 12·7 mm. in length and 2·5 mm. in breadth. The two suckers lie very close to one another, the anterior one being larger than the ventral; the genital pore opens immediately in front of the ventral sucker; pharynx spherical; intestinal cæca extending far back. At the commencement of the posterior third of the body the two testes, somewhat apart, the anterior one roundish, the posterior one distinctly lobed. The transverse and slightly lobed ovary in front of the bifurcation of the Y-shaped excretory bladder, whence the uterus, in convolutions barely spreading beyond the central field, extends to the pore; the vitellaria in the lateral areas commence behind the ventral sucker and extend to the testes. Cirrus pouch absent. Eggs oval, 34 µ by 21 µ.

Genus. Clonorchis, Looss, 1907.

Structure as in Opisthorchis, distinguished, however, by the branched testes situated one behind the other, the branches of which ventrally encroach upon the gut forks; dorsal to the testes the S-shaped excretory bladder, the main branches of which, arising at the level of the bifurcation of the gut, open into the bladder below its anterior end. Parasitic in the bile-ducts of mammals and man.

Clonorchis sinensis, Cobbold, 1875.

Syn.: Distoma sinense, Cobbold, 1875; Distoma spathulatum, R. Leuckart, 1876 (nec Rudolphi, 1819); Distoma hepatis innocuum, Baelz, 1883.

In shape resembles Opisthorchis felineus, 13 to 19 mm. long, 3 to 4 mm. broad, at the beginning of sexual maturity 12 to 13 mm. long, 2·5 to 3 mm. broad. Oral sticker 0·58 to 0·62 mm., ventral sucker 0·45 to 0·49 mm. in transverse diameter. In the parenchyma numerous yellowish or brownish granules, especially behind the oral sucker and at the posterior end. Testicular branches very long, in the anterior testis often four, in the posterior testis five branches. Ovary generally with three large lobes and a smaller lobe. Vitellaria not always symmetrical, generally extending laterally from the ventral sucker to the ovary, interrupted in parts.

Eggs 26 µ to 30 µ by 15 µ to 17 µ. Average 29 µ by 16 µ.

This (?) species was discovered in 1874 by McConnell, in Calcutta, in the bile-ducts of a Chinaman who died shortly after being admitted into hospital.

Habitat.—Bile-ducts of man, dog and cat.

Distribution.—Especially in China, apparently rare in Japan.

Clonorchis endemicus, Baelz, 1883.

Syn.: Distoma sinense s. spathulatum p.p.; Distoma hepatis endemicum s. perniciosum, Baelz, 1883; Distoma japonicum, R. Blanchard, 1886.

Very similar to the previous species and consequently generally confused with it. Length between 6 and 13 mm., width varying between 1·8 and 2·6 mm. Oral sucker 0·37 to 0·5 mm., usually 0·43 to 0·45 mm. in transverse diameter; ventral sucker 0·33 to 0·45 mm., usually 0·37 to 0·40 mm. No pigment in parenchyma; anterior testis with four, posterior testis with five branches. Vitellaria continuous, ova 26 µ by 13 µ to 16 µ.

Habitat.—Bile-ducts of man, dog, cat and pig.

Distribution.—This species occurs very frequently in man, in certain districts of Japan, especially in the province of Okayama, Central Japan, in particular localities of which above 60 per cent. of the population are infected. The worms are sometimes found in enormous numbers in the liver (upwards of 4,000), also in the pancreas and rarely in the duodenum. It is common in Tonkin and Indo-China. Léger in Tonkin found 50 per cent. of people apparently in normal health infected, so that probably symptoms only arise when the infection is intense. [The exact distribution of these two species is, however, not precisely defined at present, as commonly no distinction is made between them.—J. W. W. S.]

Verdun and Bruyant deny, in opposition to Looss, the possibility of being able to distinguish within the genus Clonorchis the two species described, but they admit the justification for the new genus. They also report the occurrence of Opisthorchis felineus in man in Tonkin (Compt. Rend. Soc. de Biol., lxii, 1907).

Pathology.—Both species of Clonorchis give rise to grave symptoms. The liver is generally enlarged, though when the infection has lasted some time it begins to contract. The surface of the organ is studded with white vesicles, and on cutting into it one sees numerous cavities with thickened walls (distended bile-ducts) filled with a brownish fluid containing innumerable eggs, which cause its colour. Microscopically, the epithelium of the bile-ducts is either (1) entirely destroyed, or (2) actively proliferates, forming an adenomatous outgrowth. Occasionally this proliferation is not limited by the wall of the bile-duct but penetrates it and leads to a growth of numerous new ducts, forming a malignant biliary adenoma. The bile-ducts have their connective tissue wall greatly sclerosed. These fuse with one another, forming areas of sclerosis devoid of liver tissue. As a result of these changes the liver cells atrophy and undergo fatty pigmentary and granular degeneration. Besides these changes, due probably to the toxic action of the flukes, mechanical obstruction due to the actual plugging of the ducts by the flukes causes retention of bile and icterus, and through pressure on veins, ascites and hypertrophy of the spleen.

To what extent blood or bile respectively forms the food of the flukes is uncertain.

Life-history.—(Kobayashi, 1911, Mitteilungen aus dem kaiserlichen Institut für Infektions-Krankheiten zu Tokio, pp. 58–62.)

It results from the work of Kobayashi in Japan that fresh-water fish form the second intermediate host for Clonorchis endemicus. He fed cats with encysted flukes (cercariæ) from various fish and easily succeeded in infecting them, e.g. a kitten, proved to be uninfected by repeated examination of its fæces, was fed on infected fish; a month later innumerable flukes were found in the bile-ducts, gall-bladder, pancreas and even in the duodenum. The fish infected were Leucogobis güntheri, Pseudorasbora parva, and to a less extent Acheclognathus lanceolata, Acheclognathus limbata, Paracheclognathus rhombea, Pseudoperilampus typus, Abbottina psegma, Biwia zezera and Sarcocheilichthys variegatus. The cysts occur throughout the muscles and subcutaneous tissue of the fish. Length 0·13 mm., breadth 0·1 mm. The cercaria lies folded in the cyst, length 0·5 mm. breadth 0·1 mm. It tapers posteriorly. Skin at first covered with fine spines, disappearing as they grow older. Body dotted with fine pigment.

The first intermediate host is still unknown.

Sub-family. Metorchiinæ, Lühe, 1909.

Genus. Metorchis, Looss, 1899, emend. auctor.

Hind end rounded. Gut forks reach extreme end. Testes only slightly lobed, filling the hind end.

Metorchis truncatus, Rud., 1819.

This species, which attains a length of 2 mm., is slender and conical, the anterior end is pointed and the posterior truncated, and provided with a muscular tuberosity that resembles a terminal sucker; for this reason the discoverer of the species (Rudolphi) classed it with the Amphistomes. The cuticle in the young, as well as in the adult specimens, is entirely and closely covered with spines. Suckers about equal in size (0·134 to 0·172 mm.); the ventral sticker lies somewhat in front of the middle of the body. The pharynx is small (0·09 mm.), the œsophagus minute, the intestinal cæca reach to the posterior extremity. Between them, and in front of their blind ends, lie the two elliptical testes, one generally a little in front of the other. In front of them, either in the median line or somewhat laterally, the spheroidal ovary is situated; in front, again, is the uterus, the coils of which usually extend beyond the median field. The vitellaria are at the sides of the central third of the body, thus commencing in front of the ventral sucker; cirrus pouch absent; the genital pore is close in front of the acetabulum. The excretory pore is terminal (?). Eggs 29 µ by 11 µ.

Metorchis truncatus lives in the bile-ducts of the seal, cat, dog, fox, and glutton (Gulo borealis). The source of infection is unknown, although one would suspect fish. Askanazy did not succeed in getting this fluke in his feeding experiments, but another species, Metorchis albidus, not uncommon in cats by feeding them on roach (Leuciscus rutilus).

Family. Heterophyiidæ, Odhner, 1914.

Genus. Heterophyes, Cobbold, 1866.

Syn.: Cotylogonimus, Lühe, 1899; Cænogonimus, Looss, 1899.

No crown of spines on head. Body divided into a narrow, movable, anterior part (neck), and a broader, less movable, posterior portion, which contains the genitalia. The suckers separated from one another by a space equal to half the length of the body or more; the pharynx is close behind the oral sucker; the œsophagus is long; the intestinal cæca extend to the posterior border; the genital pore is placed laterally, and behind the ventral sucker. Genital sucker provided with a circlet of chitinous rodlets, shaped like stags’ horns. The testes are at the posterior end, the ovary in a median position in front of them. Laurer’s canal with receptaculum seminis present; the small vitellaria are at the sides of the posterior part of the body. Parasitic in the intestine of mammals and birds.

Heterophyes heterophyes, v. Sieb., 1852.

Syn.: Distomum heterophyes, v. Siebold, 1852; Heterophyes ægyptica, Cobbold, 1866; Mesogonimus heterophyes, Railliet, 1890; Cœnogonimus heterophyes, Looss 1900; Cotylogonimus heterophyes, Braun, 1901.

Length up to 2 mm., breadth 0·4 mm.; the neck not sharply defined; in life it stretches to double the length of the hind body. The scales are rectangular, 5 µ to 6 µ by 4 µ, their posterior margin serrate with seven to nine teeth. Cuticular glands are numerous on the ventral surface, especially in the fore part of the body, and partly discharge at the anterior border of the oral sucker. The oral sucker is 0·09 mm., the ventral sucker 0·23 mm. in diameter; the pharynx measures 0·05 to 0·07 mm. in length; the œsophagus is about three times as long; posteriorly the intestinal cæca are directed one towards the other and terminate beside the excretory bladder. Close in front of the posterior ends of the intestinal branches are the two elliptical testes, which are not exactly on the same level. In the middle in front of them is the receptaculum seminis, and in front of the latter lies the spherical or elliptical ovary. The two vasa efferentia unite to form the vas deferens, which after a short course passes over into the angularly bent seminal vesicle; after the entry of the prostatic glands it becomes united with the metraterm (vagina), and the common duct opens into the genital sucker. The latter is somewhat smaller than the ventral sucker, lateral to and close (0·15 mm.) behind it, and bears a not entirely closed ring of from seventy-five to eighty chitinous rods (20 µ in length). The vitellaria on either side consist of about fourteen acini. The uterus is spread almost throughout the entire posterior part of the body. The eggs have thick shells with a knob resembling that of Clonorchis eggs but not so prominent, and measure 30 µ by 17 µ; they contain a completely ciliated miracidium with a rudimentary intestinal sac.

This species was discovered in 1851 by Bilharz in the intestine of a boy who died in Cairo; a second case was only found in 1891 and published by R. Blanchard, so that it appeared as if the species were very scarce. According to Looss, this is, however, not the case, but the species easily escapes notice on account of its small size. Looss found it in Alexandria twice in nine autopsies, and once in Cairo, and has recently stated that in man “it is not at all uncommon to meet with the parasite in cadavers, and the eggs of the worm in the stools of the patients.” Leiper records one case from Japan and one from China. The parasites occupy the middle third of the small intestine, and even when present in large numbers appear to be harmless.

This small species, according to Looss, frequently occurs in Egyptian dogs, less so in cats, and has also been found in the fox, as well as once in Milvus parasiticus; Janson also reports this species from the intestine of the dog in Japan.

Metagonimus, Katsurada, 1913; Yokogawa, Leiper, 1913.

Resembles in general structure Heterophyes. In the arrangement of its ventral genital suckers resembles but differs from that of Tocotrema, Looss. The ventral and genital suckers lie laterally and on the right.

Metagonimus yokogawai. Katsurada, 1913.

Syn.: Yokogawa yokogawai, Leiper, 1913.

One to 1·5 mm. long, seldom 2·5 mm., and 0·4 to 0·7 mm. broad; elliptical in shape. The body is thickly covered with nail-shaped spines about 10 µ long. Oral sucker 77 µ, to 85 µ in diameter. Ventral sucker characteristic and peculiar 0·12 to 0·14 mm. by 0·08 to 1 mm. It is a sac-like organ placed deeply in the body, but does not open as in other flukes on the ventral surface. Testes elliptical, not quite symmetrically placed at the hind end of the body. Vesicula seminalis retort-shaped, situated transversely, internal to the ventral sucker. Pars prostatica present. Ejaculatory duct opens with the uterus into a genital sinus, which, together with the internal opening of the ventral sucker, opens into a pit at the front of the ventral sucker. The opening of the genital sinus and that of the ventral sucker are furnished with a complex muscular apparatus. Ovary spherical, 0·12 to 0·13 mm. in diameter, lies in the middle of the hind body. Receptaculum seminis and Laurer’s canal present. Vitellaria in the hind half of the body, consisting of about ten acini on each side. Shell gland to the left of the ovary. Uterus forms three to four transverse coils. Eggs elliptical, double contoured, yellowish-brown in colour. There is no shoulder below the operculum as in the eggs of Cl. sinensis. At the rounder end there is a thickening or knob different from the spine-like or hook-like process seen in Cl. sinensis. Dimensions 28 µ by 16 µ.

Habitat.—Mainly in upper or middle portion of jejunum, rarely in cæcum. They penetrate deep into the mucosa, but not into the submucosa, and post mortem appear as a number of small brown points. They frequently occur in the solitary glands, which they destroy. They cause chronic catarrh of the gut. Parasitic in man and mammals.

Geographical Distribution.—Japan.

Life-history.—The cercarial stage occurs in a trout (Plecoglossus altivelis) and seldom in Crassius sp. and Cyprinus sp. Infection takes place through the eating of the fish raw. Seven to sixteen days later eggs appear in the fæces (of dog).

Family. Dicrocœliidæ, Odhner, 1910.

Genus. Dicrocœlium, Dujardin.

Dicrocœliidæ, with leaf-shaped bodies, pointed posteriorly and anteriorly. Greatest width behind the mid-line. Vitellaria double. The testes smooth or indented, lying symmetrically or obliquely beside or behind the ventral sucker. The ovary approaches the median line behind one testis. Parasitic in the liver and gall-bladder (rarely in the intestine) of members of all classes of vertebrate animals—by preference in birds and mammals.

Dicrocœlium dendriticum, Rud., 1819.

Syn.: Dicrocœlium lanceatum, Stil. and Hass., 1896; Fasciola lanceolata, Rud., 1803 (nec Schrank, 1790); Distomum lanceolatum, Mehlis, 1825; Dicrocœlium lanceolatum, Dujardin, 1845.

Body lancet-shaped, narrowing especially at the anterior extremity; length 8 to 10 mm., breadth 1·5 to 2·5 mm., the greatest breadth usually behind the middle of the body. Suckers distant from each other by about one-fifth the length of the body; oral sucker about 0·5 mm., ventral sucker about 0·6 mm. Pharynx globular, adjoining the oral sucker; œsophagus 0·6 mm. in length; intestinal cæca reach to four-fifths of the body length. Genital pore at the level of the bifurcation of the intestine; cirrus pouch small and slender. The large, slightly lobed testes lie obliquely one behind the other behind the ventral sucker; the ovary, which is considerably smaller, is placed behind the posterior one; the vitellaria, commencing at the level of the posterior testis, terminate far before the cæca. The uterus, situated behind the ovary, extends throughout the posterior end, not confined to the central field, but overlapping the lateral fields with its transverse coils; at the posterior edge of the body it turns back again and winds forwards to the ovary in transverse loops, then between the testes, and finally, dorsal to the ventral sucker, terminates in the genital pore. The thick-shelled eggs when young are yellowish, when older dark brown. They measure 38 µ to 45 µ by 22 µ to 30 µ. They contain an oval or roundish miracidium, only the anterior part of which is ciliated, and which possesses a rudimentary intestinal sac with a boring spine. The miracidia do not hatch out in water spontaneously, but, according to Leuckart, in the intestines of slugs (Limax, Avion), but they do not develop either in these (slugs) or in water-snails.

The lancet fluke inhabits the biliary duct of herbivorous and omnivorous mammals (sheep, ox, goat, ass, horse, deer, hare, rabbit, pig), and is often found associated with the liver fluke; it is not, however, so common nor so widely disseminated, nevertheless, it has been met with outside of Europe, namely, in Algeria, Egypt, Siberia, Turkestan, and North and South America.

In man it is still more uncommon than the liver fluke, and has hitherto only been observed seven times (Germany, Bohemia, Italy, France, and Egypt); it may, however, have occurred more frequently, and have been overlooked, as in slight infections it produces no special symptoms.

The intermediate host is still unknown. Leuckart for some time held the opinion that small species of Planorbis from fresh water, which contain encysted Distomata, were to blame, and he supported his views by a feeding experiment which seemingly yielded positive results; this, however, is not definitely proved. Piana’s statement that small land snails are the intermediate hosts has also not been proved.

Family. Echinostomidæ, Looss, 1902.

Sub-family. Echinostominæ, Looss, 1899.

Genus. Echinostoma, Rud. 1809; Dietz, 1910.

Fore-body not bulging. Greatest width at or behind the ventral sucker. Oral sucker not atrophied. Collar kidney-shaped with a double dorsally unbroken row of spines, terminating in four to five angle spines. The border spines of the aboral series not larger than the oral. Skin spined or smooth. Body elongated. Uterus long with numerous transverse coils. Ventral sucker in the anterior quarter of body. Cirrus sac small, almost completely in front of the ventral sucker. Testes round or oval, smooth incurved or lobed, in the hinder half of body. Ovary median or lateral in front of testes. Vitellaria from hinder margin of ventral sucker to end of body. Eggs oval, 84 µ to 126 µ by 48 µ to 82 µ.

The spines placed most ventrally, or those placed most medially on ventral surface, are from differences of position or form termed “angle” spines, the rest “border” spines.

Type.—Echinostoma echinatum, Rud.

Echinostoma ilocanum, Garrison, 1908.

Length 4 to 5 mm., breadth 1 to 1·35 mm., thickness 0·5 to 0·6 mm. The circum-oral disc 0·3 mm. broad, separated by a shallow groove from the body. Crown of forty-nine spines and five to six angle spines on each side continuous with an irregularly alternating series of fourteen spines on the dorsum. Largest spines are 34 µ long, 8 µ thick at the base. The remainder of the dorsal spines are 24 µ by 6 µ. Skin thickly covered with scales on the margins of the body as far back as the level of the hind testis. Oral sucker, 0·18 mm.; ventral sucker, 0·4 to 0·46 mm. Its anterior border about 0·07 mm. from the anterior end. Pharynx 0·17 mm. long, 0·11 mm. broad. Testes about mid-line of the body, much lobed; the lobes of the anterior testis run transversely, while the axis of the posterior testis is longitudinal, as often occurs in the Echinostomidæ. Cirrus sac reaches to the centre of the ventral sucker. Ovary transversely oval in front of the testes. Vitellaria commence about half-way between the ventral sucker and ovary and extend to the posterior end. Eggs numerous, 92 µ to 114 µ by 53 µ to 82 µ.

Average.—99·5 µ by 56 µ.

Habitat.—Gut of man (Filipinos), Philippine Islands.

Echinostoma malayanum, Leiper, 1911.

Twelve millimetres long, 3 mm. broad, 1·3 mm. thick. Ends bluntly rounded. At the anterior end a ventral furrow on either side, one-third the width of the body, marking off the circum-oral collar. Along its edge is a row of forty-three spines extending across the middle line dorsally but not ventrally. The spines vary in size from 0·07 mm. in length (ventrally) to 0·05 to 0·016 mm. (dorsally). Cuticular spines also exist on the ventral side as far back as posterior end of body, but dorsally limited to a triangular area ending in front of the ventral sucker. Oral sucker 0·07 mm. thick, occupying the middle third of the circum-oral disc; pharynx 0·25 mm. in diameter; œsophagus 0·04 mm. long; gut cæca simple, extending to end of body; ventral sucker 0·9 mm. long by 0·75 mm. broad by 0·7 mm. deep; wall about 0·25 mm. thick. The sucker is inclined at an angle of 40° to the ventral surface. Testes lobed, one behind the other, behind the ventral sucker. Cirrus pouch well developed, reaching to the posterior edge of the sucker. Genital pore in the angle between neck and anterior lip of ventral sucker. Ovary smooth, 0·3 mm. in diameter, 0·85 mm. behind ventral sucker. Vitellaria very numerous, extending from posterior margin of sucker to posterior end of body, where they intermingle. Eggs few in number, brown and large.

Habitat.—Gut of man (Tamils), Malay States.

Sub-family. Himasthlinæ, Odhner, 1910.

Genus. Artyfechinostomum, Clayton-Lane, 1915.

Crown of thirty-nine spines, continuous over dorsum. Two corner spines long. Vitellaria extend from posterior margin of sucker to posterior end of fluke. Eggs without filament. [Although the possession of strong rose-thorn hooks is given by Odhner as a sub-family characteristic, yet in this genus assigned to this sub-family they have not been seen.—J. W. W. S.]

Artyfechinostomum sufrartyfex, Clayton-Lane, 1915.

Spirit specimens: 9 by 2·5 by 0·8 mm. thick. Ventral sucker conspicuous, 1 mm. in diameter. Cirrus sac 2 mm. long. Testes lobed, about 1·5 mm. in diameter. Posterior border of posterior testes 1 mm. from posterior end. Vitellaria meet posteriorly behind the posterior testis.

Family. Schistosomidæ, Looss, 1899.

Genus. Schistosoma, Weinl, 1858.

Syn.: Gynæcophorus, Dies., 1858; Bilharzia, Cobb., 1859; Thecosoma, Moq. Tandon, 1860.

The males have bodies that widen out considerably behind the ventral sucker, the lateral parts of which in-roll ventrally, forming the almost completely closed canalis gynæcophorus, within which the female is enclosed. There is no cirrus pouch. The male has five or six testes, the females are filiform; the uterus is long. There is no Laurer’s canal. The ova almost equally attenuated at either extremity; they have a small terminal spine, and are not provided with a lid. They contain a miracidium, ciliated on all sides, which is characterized by the possession of two large glandular cells, which discharge anteriorly beside the gastric sac. They live in the vascular system of mammals. (An allied genus [Bilharziella] lives in the blood-vessels of birds.)

Schistosoma hæmatobium, Bilharz, 1852.

Syn.: Distoma hæmatobium, Bilh.; Distoma capense, Harley, 1864.

The Male is whitish, 12 to 14 mm. in length, but is already mature when 4 mm. long. The anterior end is 0·6 mm. or a little over in length. The suckers are near each other, the oral sucker is infundibular, and the dorsal lip is longer than the ventral one. The ventral sucker is a little larger, 0·28 mm., and is pedunculated. A little behind the ventral sucker the body broadens to a width of 1 mm., decreasing, however, in thickness; the lateral edges in-roll ventrally, so that the posterior part of the body appears almost cylindrical, 0·4 to 0·5 mm. in diameter; the posterior extremity is somewhat more attenuated. The dorsal surface of the posterior part of the body is covered with spinous papillæ. There are delicate spines on the suckers, and larger ones invest the entire internal surface of the gynæcophoric canal, as well as a longitudinal zone at the edge of that side of the external surface that is covered by the other side rolling over it. The œsophagus is beset with numerous glandular cells (fig. 173), and presents two dilatations; the intestinal bifurcation is close in front of the ventral sucker, the two branches uniting sooner or later behind the testes into a median trunk, which may again divide at short intervals. The excretory pore is at the posterior end, but placed somewhat dorsally; the genital pore is at the beginning of the gynæcophoric canal, thus behind the ventral sucker; into it opens the vas deferens which, posteriorly, broadens into the seminal vesicle and then continues as the vasa efferentia of the four or five testes (fig. 173).

The Female—filiform, about 20 mm. in length, pointed at each end, and measuring 0·25 mm. in diameter in the middle. Their colour varies according to the condition of the contents of the intestine. (Posteriorly they are dark brown or blackish.) The cuticle is smooth except in the sucker, where there are very delicate spines, and at the posterior end, where there are other larger spines. The oral sucker is a little larger than the pedunculated ventral sucker (0·07 and 0·059 mm. respectively). The anterior part of the body, 0·2 to 0·3 mm. in length; the œsophagus is as in the male. The intestinal bifurcation is in front of the ventral sucker, the two branches uniting behind the ovary and the trunk running in a zigzag manner to the posterior border. There are indications of diverticula at the flexures. The ovary is median. In young females it is of an elongated oval shape; in older females the posterior end becomes club-shaped, whereas the anterior end becomes attenuated; the oviduct originates at the posterior end, but immediately turns forwards and joins the parallel vitelline duct in front of the ovary (fig. 174), where the shell gland cells open; the common canal becomes dilated to form the oötype, and then proceeds as the uterus, with only slight convolutions, along the central field to the genital pore, which lies in the middle line immediately behind the ventral sucker. The single vitellarium starts behind the ovary and extends to the posterior end. The acini are situated at the sides of the excretory duct, which runs a median course. The eggs are compact spindles, much dilated in the middle; they have no lid, and are provided with a terminal spine (rudimentary filament) at the posterior end, measuring 120 µ to 150 µ in length and 40 µ to 60 µ in breadth, but vary in size and shape (fig. ).

Distribution.—In order to understand the distribution of the worms and eggs in the body, it may be well to recall the blood supply of the abdominal and pelvic organs. It is generally assumed that the early life (? cercarial stage) of the worms occurs in the liver, and that the young worms travel from here, where they are invariably found, to their various sites along the portal vein and its tributaries and so against the blood stream. The tributaries of the portal vein are:—

(1) Superior mesenteric, the tributaries of which are: (a) the veins of the small intestine; (b) ileo-colic; (c) right colic; (d) middle colic; (e) right gastro-epiploic; and (f) inferior pancreatic. By these paths infection of the small intestine, ascending and transverse colon and pancreas would occur.

(2) Splenic. (Ova have been recorded by Symmers in the spleen.)

(3) Inferior mesenteric, the tributaries of which are (a) superior hæmorrhoidal veins from the upper part of the hæmorrhoidal plexus; (b) sigmoid veins from sigmoid flexure and lower portion of descending colon; (c) left colic vein draining descending colon.

The superior hæmorrhoidal veins form a rich plexus in the rectum, and below this level in the upper and middle parts of the anal canal. The plexus forms two networks, an internal plexus in the submucosa and an external on the outer surface. The internal plexus opens at the anal orifice into: (a) branches of the inferior hæmorrhoidal vein (from the pudic); (b) the external plexus. The external plexus gives off: (a) inferior hæmorrhoidal opening into internal pudic (of internal iliac vein); (b) mid-hæmorrhoidal into internal iliac or its branches; and (c) superior hæmorrhoidal opening into inferior mesenteric. The external plexus further communicates with the vesico-prostatic plexus. The vesico-prostatic (vaginal) plexus opens into the vesical veins, which drain into the interior iliac vein. This plexus also receives afferents from the pudendal plexus, the chief tributary of which is the dorsal vein of the penis. The pudendal plexus also receives branches from the inferior pudic and the anterior surface of the bladder.

There is thus a communication between the portal vein and the vena cava by means of these plexuses, viz., through the inferior and middle hæmorrhoidals, and by the inferior hæmorrhoidals to the bladder and thence by the vesical veins or the pudic to the caval system (interior iliac).

It is thus by the inferior mesenteric and its tributaries that the worms reach the descending colon, rectum, anal canal, and eventually the bladder, and in some cases the caval system.

Before considering what is actually found post mortem in these veins and the organs drained by them, we may further recall the fact that the calibre of “medium” veins is 4 to 8 mm., “small” veins less than 40 µ in diameter and capillaries 8 µ to 20 µ. Further, the maximum diameter of the male worm is 1 mm., that of the female 280 µ and eggs in utero 80 µ to 90 µ long by 30 µ to 40 µ.

Liver and Portal Vein.—Here worms are most easily found post mortem. Often only males are found and these of the same size, and if females occur only a few worms are found in copulâ. The worms are frequently not full size and the males may contain no free spermatozoa in their testes, and as regards the females some may be fertilized, others not, as shown by the presence or absence of spermatozoa in the seminal receptacle or uterus. In either case they may contain eggs—lateral-spined—usually one, less often two, but there may be as many as five or six. These eggs may also show some abnormality, which takes the form of: (1) abnormal contents, viz., disintegrating yolk cells with or without an ovarian cell; (2) abnormal shape but with normal contents and probably represented by the collapsed and empty egg-shells which are found in the tissues.

As to the interpretation of these facts, Looss believes that these lateral-spined eggs are products of young females whose egg-laying is not at first properly regulated. The shape that the eggs take, viz., with a lateral spine, is determined by an excess of material—ovarian and yolk cells—being present in the oötype. The shape of eggs depends upon the position they have in the oötype during their formation. In young females an excess of cells—yolk cells especially—accumulates, distending not only the dorsal wall but a portion also of the short duct joining the oötype to the uterus. The result of this is that the axis of the oötype and egg is almost transverse to the body, and the posterior funnel-shaped portion of the oötype, instead of being terminal, has now a lateral or rather a ventral position, so that the spine which occupies this portion, instead of being terminal, is now lateral. It is noteworthy that these lateral-spined eggs are thicker, owing to the excess of material present, and not uncommonly have a curved anterior border, due to a projection of the anterior end into the anterior opening of the oötype.

As these eggs are being laid by females in the portal vein they are carried back to the liver by the blood stream. The liver is one of the commonest sites for these eggs; also terminal-spined eggs may be found here for the same reason.

Hæmorrhoidal Veins.—Mature worms, generally in copulâ, are usually found here, though young not fully grown females may also occur. The tissues of the rectal wall (or colon) show, as a rule, large quantities of lateral-spined eggs, though less often only terminal-spined eggs may be found.

Vesico-prostatic Plexus.—Worms in copulâ are found in the veins of the submucosa in the bladder, and the eggs in the mucosa, and those voided are usually terminal-spined, though lateral-spined eggs are not so rare as generally thought. The problem next arises as to how the eggs get to the lumen of the gut or bladder.

The female worm is 280 µ in diameter. Veins in the submucosa of the rectum less than 178 µ in diameter are not affected with endophlebitis. It is probable that the female even by stretching could not penetrate much beyond this. Eggs are probably then laid in the submucosa as near the muscularis mucosa as possible. Now if the eggs are laid in a vein of larger calibre than the worm fills, the eggs would be carried back to the inferior mesenteric vein, so that presumably the worm must succeed in blocking the vein already narrowed by endophlebitis, so that by the stasis which ensues the eggs may escape from the veins. How this occurs is not exactly known; it is not necessarily due to the spine, as the same escape into the tissues occurs in spineless eggs, such as those of Schistosoma japonicum. The eggs, then, pass as foreign bodies through the tissues. Another hypothesis is that the worms leave the veins in order to lay their eggs, but the evidence is against this.

Caval System.—Occasionally worms that have passed through the vesical plexus may be found in the iliac vein, inferior vena cava, and even the lungs. If the worms are young they contain a lateral-spined egg; if adult, numerous (50 to 100) terminal-spined eggs.

Lungs.—When the liver is strongly infected with (terminal-spined) eggs it is possible that by passive movements some may pass into the intralobular veins, and thence by the inferior vena cava to the lungs.

Gall-bladder.—Similarly terminal-spined eggs pass into the bile-capillaries and gall-bladder (where they may be abundant), and so into the fæces.

Detection of Eggs.—Occasionally eggs may be found in various other parts of the body. They are best detected by macerating pieces of the tissue in question in about 1/4 per cent. hydrochloric acid at 50 to 60°C. (Looss).

Pathological changes:—

Rectum.—These have been studied thoroughly by Letulle in the case of an apparently pure infection of the rectum. They take the form of a chronic diffuse inflammation, which may result in—(1) ulceration, or (2) hyperplasia of the mucosa, producing adenomata.

Ulcerative Form.—The mucosa is transformed into a mass of vascular connective tissue. The connective tissue spaces next become invaded by numerous mononuclear cells. The tissue itself undergoes diffuse sclerosis, becoming hard and fibroid. Eventually ulcerative necrosis sets in. During these changes the Lieberkühn glands are destroyed. The process does not extend to the submucosa, in this respect differing from that in chronic dysentery.

Hyperplastic Form.—The Lieberkühn glands of the mucosa at first hypertrophy; then there is an actual hyperplasia resulting in adenomata. The interstitial tissue of the glands is also greatly hypertrophied, giving rise to very vascular granulations. These growths are often hollow and contain worms. Many eggs are found in the mucosa on their way to the lumen of the gut.

The muscularis mucosa is thickened up to twice or even ten times the normal. Its vessels are dilated (36 µ to 80 µ), but they do not allow of the passage of worms.

The submucosa is profoundly changed; rigid and hard instead of supple. It is here that the greatest number of eggs occur. A remarkable condition of endophlebitis exists in the veins of the submucosa, not only in the smaller ones but also in the larger ones (370 µ by 270 µ). This endophlebitis results in a more or less complete occlusion of the vessels of the lumen.

The muscular coats are free from change, also their veins.

The Serous Coats.—The veins about 1,900 µ, also show endophlebitis. Besides the rectum, in extreme cases even the transverse colon, the cæcum and small intestine may be affected.

Bladder.—In the early stages the mucosa is deep red and swollen like velvet, or there may be localized patches of hyperæmia or extravasation. The subsequent changes take two chief forms:—

(1) Sandy Patches.—The mucosa looks as if it were impregnated with a fine brownish or yellowish powder (myriads of ova). This is accompanied by a gradual hypertrophy and new formation of connective tissue, so that dry, hard or plate-like patches with this sandy appearance arise; the thickening eventually affects all the coats of the bladder. In the older patches many of the eggs are calcified. These patches sooner or later break down, ulcerate and necrose. Phosphatic deposits are abundant and stone is common. These patches are not found in the rectum.

(2) Papillomata.—Where the inflammatory change produced by the eggs gives rise to hypertrophy and hyperplasia of the mucosa, papillomata result, the axis of which is formed by connective tissue of the submucosa. These are most variable in shape and form and bleed readily, and sometimes contain cavities of extravasated blood.

As in the rectum, it is in the submucosa that eggs are most abundant, and worms in copulâ occur in the veins of this layer, but endophlebitis is not as general as described in the rectum. Malignant disease of the bladder is not an uncommon sequela of bilharziasis. Besides the bladder, the ureters and kidneys may in advanced cases be involved. The prostate and vesiculæ seminales are commonly diseased. Eggs have been recorded in the semen. The urethra is frequently attacked; the vagina in the female.

Eggs also occur in the lymphatic glands of the gut.

Geographical Distribution.—East Africa: Nile Valley, Red Sea Coast, Zanzibar, Portuguese East Africa, Delagoa Bay, Natal, Port Elizabeth.

South Africa: Cape Colony, Orange Free State, Transvaal, Mauritius, Bourbon, Madagascar.

West Africa: Angola, Cameroons, Gold Coast, Gambia, Senegal, Sierra Leone, Lagos, Nigeria.

North Africa: Tripoli, Tunis, Algeria, parts of the Sahara.

Central Africa: Sudan, various portions. Uganda, Nyasaland.

It occurs with varying frequency in these regions. It is probably more widely spread than this list implies, as undoubtedly many cases are seen which are not recorded.

Isolated cases have been recorded from Arabia, India, Greece, Cyprus.

The means by which infection is brought about are still uncertain; we only know that the miracidia (fig. 175) enclosed in the discharged eggs do not hatch if the eggs remain in the urine, but after cooling perish. As soon, however, as the urine is diluted with water the shell swells, generally bursting lengthways, and releases the miracidium from its investing membrane, so that it can swim about with the aid of its cilia. In its structure it differs but little from the miracidium of Fasciola hepatica, as, for instance, in the lack of eyes; the two large gland cells situated on either side of the intestinal sac are also present in the miracidia of Fasciola hepatica.

Sarcode Globules.—This is a term applied to certain globules which at times appear in the miracidium and are later ejected. Some authors consider them as indicative that the miracidium has developed into a sporocyst, but Looss considers them to be degeneration products.

The Bilharzia mission, under R. T. Leiper, sent to Egypt by the War Office early in 1915, reports that cercariæ of bilharzia type were recognized in four of the commonest fresh-water molluscs around Cairo.

With material obtained from naturally infected Planorbis boissyi acute bilharziosis was experimentally produced in rats, mice, and monkeys. Infection takes place experimentally through the skin and also through the mucous membrane of the mouth and œsophagus. The miracidium, after entering the mollusc, develops into a sporocyst. This gives rise not to rediæ, but to secondary sporocysts, which, in turn, produce cercariæ. These, like the adult worm, differ from other distomes in lacking a muscular pharynx.

Schistosoma mansoni, Sambon, 1907.

According to Manson, Sambon and others, the eggs with lateral spines belong to a species different from Schistosoma hæmatobium. Infections with this species only are said to occur in the Congo, Southern States of North America, West Indies (Guadeloupe) and Brazil (Bahia). The following characters, according to Flu, differentiate this species: (1) In the male the transition from the anterior portion of the worm to the lateral fields (the infolded portions which form the gynæcophoric canal) is not a gradual one as in Schistosoma hæmatobium, but in this case the lateral fields rise suddenly, almost at right angles to the anterior portion. (2) The ovaries have a well-marked convoluted course as in no other schistosome. (3) The oötype is symmetrical in reference to the long axis of the body, its duct being lateral on the ventral side (Looss’ explanation of this we have already given). (4) The worms live exclusively in portal vein and tract. (As lateral-spined eggs occur also in the bladder, this is not exactly true.)

Schistosoma hæmatobium, Bilharz, 1852.

Male, four or five large testes. Gut forks unite late, so that the single gut stem is short. Female, ovary in posterior half of body. Uterus very long, voluminous, with many terminal-spined eggs, some lying in pairs. Vitellaria in posterior fourth of body. Cercariæ in Bullinus contortus and Bullinus dybowski (syn.: Physa alexandrina) in Egypt.

Schistosoma mansoni, Sambon, 1907.

Male, eight small testes. Gut forks unite early, so that the single gut stem is very long. Females, ovary in anterior half of body. Uterus very short; usually only one lateral-spined egg at a time in utero. Vitellaria occupy posterior two-thirds of body. Cercariæ in Planorbis boissyi in Egypt.

The above morphological descriptions are founded on worms of each species, derived from experimentally infected mice (Leiper, R. T., Brit. Med. Journ., March 18, 1916, p. 411).

Schistosoma japonicum, Katsurada, 1904.

Syn.: S. cattoi, Blanchard, 1905.

Male.—Eight to 19 mm., but extreme limits are 5 to 22·5 mm. Consists of a short fore-body, separated by the ventral sucker from the hind-body. The ventral sucker is stalked and somewhat larger than the oral sucker. Both suckers are larger than the corresponding ones in S. hæmatobium. Body usually smooth, but in the fresh state numerous fairly evident spines along the margin of the canal. Œsophagus: two bulbs. The junction of the gut forks more posterior than in S. hæmatobium, the median united gut stem occupying a quarter to one-fifth to one-sixth of the body length. An excretory canal runs along each side of the body, opening into the dorsal excretory pore. Testes irregularly elliptical, six to eight in number, in the anterior part of hind-body. The vasa efferentia unite into a common vas deferens which opens directly behind the ventral sucker. The seminal vesicle lies just behind this.

Female.—Up to 26 mm., generally thinner than the male. Surface smooth. Suckers armed with fine spines. Ventral sucker larger than oral. Body thicker behind the region of the ovary. The gut forks unite immediately behind the ovary. The united gut much thicker than in S. hæmatobium. Ovary elliptical, almost in the mid-body, its hinder portion dilated. The oviduct arises from its posterior end and then runs sinuously forward, where it is joined by the vitellarian duct; the vitellarium well developed, extending from behind the ovary almost but not quite to the posterior end as in S. hæmatobium. Shell gland ducts enter at the junction point of oviduct and vitelline duct. The canal here forms an oötype and then proceeds as the uterus to open directly behind the ventral sucker. The uterus occupies almost half the hind-body. In S. hæmatobium this is not so. The uterine canal is cleft-like, i.e., its dorso-ventral diameter is much greater than its lateral diameter. The number of eggs varies from about 50 to 300 from observations made in various hosts.

Eggs.—In utero assume various shapes, as they are soft; the lumen of the uterus is narrow. Outside they are oval, faint yellow, double contoured. In fæces the eggs measure 83·5 µ, by 62·5 µ (man); 85 µ by 61·5 µ (cattle); 98·2 µ by 73·8 µ (dog). The eggs have either small lateral spines or thickenings, and Looss at the opposite side has described cap-like thickenings. The eggs in the tissues undergo various deformities, and may contain a miracidium, as also the eggs in fæces do; or the contents may consist of granular matter or amorphous masses or they may be calcified. Lymphocytes and giant cells may also invade the eggs.

Mode of Infection.—The miracidia hatch in water in as little as fifteen minutes, but the majority in one to three hours. They will live in water for about twenty-four hours. In water they undergo a transformation into “larvæ,” which then penetrate the skin, as has been shown by Japanese writers to hold good for man, cattle, dog and cat. The penetration of the skin is attended with an eruption on the legs, “Kabure.” The exact route by which the worms reach the portal vein is uncertain. Infection in Japan takes place from spring to autumn, especially May to July, when the soil is contaminated with manure of cattle infected with S. japonicum. They also appear to develop in molluscs. Leiper and Atkinson found cercariæ (in sporocysts) in the liver of a mollusc, Katayama nosophora. They infected mice by immersing them in water containing liver emulsion and so free cercariæ, thus confirming the similar results of Miyairi and Suzuki.

Habitat.—The worm occurs in Japan, China, and the Philippines. The normal host is man and mammals. Cattle, dog and cat are often found naturally infected. Mice can also be experimentally infected. Their seat of election is the portal vein and its branches, especially the mesenteric veins. They either swim free in the blood or remain fixed by their suckers to the intima of the vessels. They have also been found in the vena cava and right heart of a cat, but not so far in the vesical plexus.

Eggs are found in the submucosa and mucosa of the gut, especially the colon, and at times in the serosa and subserosa of the small intestine, where they give rise to new growths. Occasionally eggs are found in the brain. The life of the worms is at least two years.

Pathogenic Effects.—Anæmia through loss of blood due to worms; enlarged spleen, toxic in origin (?); phlebitis, thrombosis, due to portal stasis; the eggs, however, cause the greatest mischief. They are carried by the circulation to various organs where they produce inflammation, granulation tissue, and later connective tissue.

Liver.—The eggs reaching this organ give rise to granulomata and hence enlarged liver, and later, when connective tissue is formed, to contraction. The surface is rough and irregularly granular, “parasitic embolic cirrhosis” of Yamagiwa.

Gut.—The eggs in the mucosa and submucosa cause catarrh and destruction of tissue or new growth. In the small intestine the eggs are mainly in the serosa and subserosa, where they give rise to polypoid or branched growths.

Spleen.—Enlarged, at first due to toxin (?) and later due to portal stasis. Eggs in the spleen are uncommon.

Ascites also arises from the portal stasis, and is generally present in advanced cases.

Eggs may be found in many other situations: glands (numerous), mesentery, stomach, pancreas, kidney, etc. The bladder remains free.