collabüratürs - taxonomy · collabüratürs flies e. paul catts university of delaware newark,...
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COLLABüRATüRS
FLIES
E. PAUL CATTS University of Delaware Newark, Delaware
LICE
KARY C. EMERSON Arlington, Virginia
PENTASTOMIDS
ALEX FAIN Institute de Médecine Tropicale
Prince Léopold Antwerp, Belgium
NEMATODES, ACANTHOCEPHALANS,
LE.ECHES
DONALD HEYNEMAN University of California San Francisco, California
PARASITES OF FISHES
GLENN L. HOFFMAN Eastern Fish Disease Laboratory Leetown, "Vest Virginia
TICKS
HARRY HOOGSTRAAL U.S. Naval Medical Research Unit
No. 3 Cairo, U.A.R.
PARASITES OF AMPHIBIANS
AND REPTILES
HAROLD M. KAPL\N Southern Illinois University Carbondale, Illinois
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LICE
KE CHUNG KIM Pennsylvania State University University Park, Pennsylvania
PROTOZOANS
NORMAN D. LEVINE University of Illinois Urbana, Illinois
FLEAS
ROBERT E. LEWIS Iowa State University Ames, Iowa
LICE
ROGER D. PRICE University of Minnesota St. Paul, Minnesota
BUGS
RAy:yrOND E. RYCK:YL\N Loma Linda University Loma Linda, California
TREMATODES, CESTODES
MARIETTA VOGE University of California Los Angeles, California
MITES
CONRAD YUNKER Rocky Mountain Laboratory Hamilton, Montana
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Chapter 16
PENT ASTOMIDS
by Alex Fain
PENTASTOMIDS wmtitute a highly aberrant group of arthropods (Fain 1961; Heymons 1935). They have a wormlike and generally annulated appearance. The body is white. legless, and either cylindrical or flat.
These arthropods are typically heteroxenous parasites. In the most evolved species, the adults live in the respiratory tract of carnivorous animaIs, usually snakes, and the larvae develop in the tissues of various animaIs, usually mammals.
The intermediate host becomes infected by drinking water or eating food contaminated by fecal material or by mucus from the respiratory tract of an animaI harboring adult pentastomids; the definitive host becomes infected by eating animaIs or viscera containing nymphs.
The development in the intennediate host gener:llly takes several months. The development:ll stages include a prim:lry or migr:lting bna; a secondary or resting larva which molts sever:ll times; and a tertiary larva or nymph which encysts in the tissues of the host, generally in the peritoneal cavity. The terti:lry larva usuaUy remains encysted until ingested by the definitive hast. but sometimes. for reasons not dearly understood. it escapes from its cystic envelope and migra tes through the tissues of the intermediate host, causing acute peritonitis and possibly death (Chalmers 1899).
493
Natural infections have been reported in sorne endothermal species occasionally used in the laboratory. Experimental infections have also been produced, The pentastomids that may be encountered in endothermal laboratory species are listed in Table 16.1. The most important species are described below.
Linguatula sel'rata (Tongue Worm)
Linguatula serrata, a relatively benign parasite, is found throughout the world, but its exact incidence is unknown. It is commonest in Europe, especially eastern Europe (Heymons 1942). and has been re· ported from the United States (Stiles 1895), South America (Gelormini and Roveda 1938). South Africa (OrtIepp 1934), Asia (Faust 1927; Heymons 1935), the Philippine Islands (Tubangui :lnd :\lasiluilgan 19:16), ,-\ustralia (Pullar 1936), and ~ ew Ze:lland (Gurr 1953).
.-\dults accur in the nasal passages of the dog and other canids, and rarely in dames tic farm species :lncl man (Heymons 1942). The nymph sometimes occurs in the wild ~orway rat, black rat, guinea pig, rabbi t, cat, titi monkey, gelada baboon, and man (Bochefontaine 1876; Heymons 1942; Kuntz, Myers. and Vice 1967; Strong. Shattuck, and Wheeler 1926). lt is common in domestic farm species. A report of canine nymphal linguaculosis in Japan ,
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494 PARASITES
is of doubtful validity (Yamashita and Ohbayashi 1954). The nymph was prohabl)' ATmillifer moniliformis.
The incidence of L. sen-ata in laboratory specimens is unknown. It is not likely to be encountered except in dogs obtained from pounds or in rodents obtained from their natural habitat. The mouse, guinea pig, and rock squirrel (Otospermophilus beecheyi) have been infected experimen
. tally (Heymons 194~; Hobmaier and Hob· maier 1940; Koch 190i).
MORPHOLOGY
The adults have a transparent, tongueshaped body with approximately 90 annuli (Fig. 16.1) (Sambon 1922). The anterior end has two pairs of simpIe retractile hooks. The female is 80 to 130 mm long and 10 mm wide, and reddish orange eggs are visible along the median line of the
FIG. 16. J. Linguatula serra ta. (Le!t) Male. (Righi) Female. (Courtesy of A. Fain, Institut de Médecine Tropicale Prince Léopold.)
OF ENDOTHERMAL LABORATORY ANIMALS
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5mm FIG. 16.2. Linguatula serrata nymphs. (Cour. tesy of A. Fain, Institut de Médecine Tropi· cale Prince Léopold.)
body; the male is 20 mm long and 3 to 4 mm wide. The nymph, sometimes called Pentastomum denticulatum, is 4 to 6 mm long and 1 mm wide (Fig. 16.2). It has spinous body rings and two pairs of binate hooks (Fig. 16.3). The egg is oval, about iO to 90 p. in diameter, and is individ ually enclosed in a thin bladderlike envelope containing a clear fluid. It has a thick chi tinous shell containing an embryo with rudimentary mouthparts and four short legs, each bearing two clawlike hooks. On the back of the em brvo is the so-called dorsal organ or facette.
LIFF. CYCLE
Eggs are expelled from the definitive host in the nasal mucus or are swallowed and passed in the feces (Hobmaier and Hobmaier 1940). When ingested by an intermediate host, thev hatch in the intestine. The resultant hrvae migrate to the internaI organs, usually the mesenteric lymph nodes, and after about 6 months and nine molts, they develop into infective nymphs. These nymphs remain viable in
PENTASTOMIDS
FIG. 16.3. Linguatula sena la nymph. Note spi nous body rings and the two pairs of binate hooks. (Courtesy of A. Fain, Institut de Méde· cine Tropicale Prince Léopold.)
the intermediate hast for over 2 vears. The definitive hast becomes infected' by ingesting viscera containing the infective stage, but the method by which nymphs get ta the nasal cavities is unknown. This may occur while contaminated food is being masticated or possib!y later during emesis. Adults survive about 2 years in the defini· tive hast. Thev feed on nasal mucus and secretions and ~ccasionally on blood (Hey· mans 1942).
PATHOLOCIC EFFECTS
Vsually there are no signs of infection, but a severe catarrhal or suppurative rhinitis and epistaxis sometimes occur (Enigk and Düwel 1957; Heymans 1942). Restlessness, sneezing, and difficult breathing are occasionally seen. The sense of smell is ohen reduced or abolished.
The nymph generally does not produce signs and is an inciden tal necropsy finding, appearing as a small fibrous or calcified tuberde in the viscera. A massive infection has camed peritonitis in an experimentally infected guinea pig (Koch 1907).
DJAC;-.lOSIS
Diagnosis is based on clinical signs and the presence of L. serrata eggs in the feces or nasal mucus.
CONTROL
Newly acquired dogs showing signs of upper respira tory disease should be exam·
495
ined for this parasite. Infected dogs can be treated by spraying the nasal passages with an aerosol containing ascaridol (the active ingredient in chenopodium oil) (Enigk and Düwel 1957), or the parasites can be removed surgically (Olt and Strôse 1914).
There is no treatment for nymphal infections.
PUBLIC HEALTH CONSIDERATIONS
This parasite is not an important public health problem (Fain 1960). The nymph and, very rarely, the adults have been reported in man.
Porocephalus
Porocephalus crotali occurs in North and South America, P. clavatlls occurs in South America, and P. subulifer is con· fined ta tropical Africa (Fain 1961, 1966; Heymans 1935). Porocephalus nymphs are occasionally found in the viscera of sorne endothermal laboratary species. They are relatively benign parasites. Adults usually live in the lung of large snakes.
The P. crotali nymph has been found in deer mice and the cotton rat in the Vnited States (Layne 1967; Self and McMurry 1948), and there is a doubtful record in a marmoset (Sagllinus) (Heymons 1935). The nymph of P. clavatus has been reported from the common marmoset (Heymons 1935), from laboratory tam:lfins (Saguin1ls nigricollis) in the United States (Cosgrove, Nelson, and Gengozian 1968; Nelson, Cosgrove, and Gengozian 1966), and, erroneously, from African primates and man (Fiennes 1967). The nymph of P. subulifer has been found in a guenon (Heymons 1935) and in a ~a1ago (Fain 1961), and nymphs of an unidentified species of Porocepha llis have been recO\'ered from a squirre1 monkey (A. Fain, unpublished data).
The rat, mouse, and hamster have been experimentally infected with eggs of P. crotali (Esslinger 1962a, b), ancl the mouse, rat, and guinea pig have been experimentally infected with eggs of P. clavatus (da Fonseca 1939).
Porocephalus cla<mtllS is apparently common in laboratorv tamarins obtained from their natural h~bitat, and an incidence of 29% has been reported (Self and
496 PARASITES OF ENDOTHERMAL LABORATORY ANIMALS
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FIc. 16.4. Porocephalus nymph. (Courtesy of A. Fain, Institut de Médecine Tropicale Prince Léopold.)
Cosgrove 1968). The incidence of P. crotali and P. subulifer in laboratory specimens is unknown. They are not likely to be encountered except in sorne primates and wild rodents obtained from endemic areas.
I\fORPHOLOCY
The Porocephalus nymph has a cylin· drical, smoothly annulated body which is often club shaped (Fig. 16.4) (Heymons 1935). Nymphs of P. crotali and P. da· vatus are about 8 to 14 mm long; the nymph of P. subulifer is 7 to 15 mm long and 1.2 to 1.5 mm wide. Each nymph has approximately 30 to 45 annuli. Two un· equal pa~rs of hooks are located at the anterior end around the mouth. The inner pair is simple; each of the outer hooks has an accessory spine. The adults are similar, only larger.
LlFE CYCLE
The life cvcle resembles that of Linguatula sen'ata' except that the adults oc· cur in snakes instead of in the dog and other canids. Adults of P. crotali are corn· mon in rattlesnakes (Crotalus) and also occur in the cottonmouth water moccasin (Aneistrodon piscivorus); adults of P. clavatus occur in boas (Boa, Epicrates and Eunectes); and those of P. stlbulifer occur only in file snakes (Mehelya) (Fain 1961,
1966; Heymons 1935; Penn 1942; Self and McMurry 1948).
PATHOLOCIC EFFECTS
In deer mice and the cotton rat, the nymph locates in the viscera, mesentery, and abdominal and thoracic walls (Layne 1967). It produces no signs or serious pa· thology.
In primates, the nymph encysts in many tissues (Fig. 16.5. 16.6), including the liver. lungs. peritoneum. and meninges, but produces little or no injury (A. Fain. unpublished data; Nelson, Cosgrove, and Gengozian 1966). Inflammatory reaction is minimal (Fig. 16.7) unless the nymph dies; then a foreign body reaction and graduaI resorption occur (Fig. 16.8).
DIAGNOSIS
Since This parasite usually does not produce signs or lesions, diagnosis is made by finding the nymph at necrops)'.
CONTROL
No special procedures other than routine sanitation are necessan. The Porocephalus nymph can occur only in labora· tory species permined to ingest food contaminated with feces of infected snakes. There is no treatment.
PUBLIC HEALTH CONSIDERATIONS
POl'Ocephalus is of no known public health importance. AU reports of the nymph in man are of doubtful \'alidity (Fain 1960).
Arrnillifer ar711illatlls
This rela tively benign parasi te occurs naturally only in tropical A[rica, where it is common. The nymphs develop in var· ious endothermal species and are frequently found in the rhesus monkey, other macaques, galagos, guenons, mangabeys, baboons, the chimpanzee, dog, and man (Fain 1960, 1961, 1966; Fiennes 1967; Hey. mons 1935). There is a report of the occurrence of the nymphs in a New World monkey, a capu chin, from a European zoo (Desportes and Roth 1943), but This infection was probably acquired in the zoo. Adults of A"millifer armillatus live in the lung of large snak·es.
The incidence of A. arrnillatus in lab
FIG. 16.5. Porocephalus clavatus nymphs (arrows) in the liver and lungs of a tamarin (Saguinus nigricollis). (From Nelson, Cosgrove, and Gengozian 1966. Courtesy of American Association for Laboratory Animal Science.)
FIG. 16.6. Porocephalus nymph on the liver of a squirrel monkey. (Courtesy of A. Fain, InStitut de Médecine Tropicale Prince Léopold.)
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FIG. 16.7. Histologie section of encysted Porocephalus clavatus nymph in the liver of a tamarin. (A) Liver tissue. (B) Parasite. Note absence of inflammation. (From Nelson, Cosgrove, and Gengozian 1966. Courtesy of Ameri· can Association for Laboratory Animal Science.)
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498
FIG. 16.8. Histologie section of the liver of a tamarin containing degenerating Porocephalus clavatus nymph. Note inflammatory reaction. (From Nelson, Cosgro\'e. and Gengozian 1966. Counes)' of American Association for Labora· tory Animal Science.)
oratory specimens is unknown. It is nOl likely to be encountered except in primates obtained from tropical Africa.
MORPHOLOGY
The nymphs have a cylindrical, annulated body about 13 to 23 mm long (Fig. 16.9) (Fain 1961). The annuli are thick and projecting, and their number varies from 15 to 19 in the male nymph and from 18 to 22 in the female nymph. The hooks are simple. Adults closely re· semble the nymphs but are larger.
LIFE CYCLE
The adults live in the lung of large African snakes (Python, Bitis) (Broden and Rodhain 1907, 1908-1909, 1910). The in· termediate host becomes infected by swallowing food contaminated by snake feces or saliva.
PATHOLOGIC EFFECTS
Encysted nymphs are commonly found in the peritoneal cavity (Fig. 16.10) (Fain
PARASITES OF ENDOTHERMAL LABOR\\TORY ANIMALS
FIG~ 16.9. Armillife,' annillatus nymphs. (From Fain 1961. Courtes)' of Musée royal de l'Afrique Centrale.)
1961). Often they are located beneath the capsule or are embedded in the superficial layers of the liver. They usually cause little or no reaction in the host even at high levels of infection, but there is one report of peritonitis and death caused by Armillifcr nymphs in 7 of 24 laboratory mangabeys (Whitney and Kruckenberg 1967).
DIAGNOSIS
Diagnosis is based on finding nymphs at necropsy.
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499 r PENTASTOMIDS
CONTROL
Other than routine sanitation, no special control procedures are necessary. In· fection can only occur if laboratory species are permitted ta ingest eggs passed in the feces or saliva of infected snakes. There is no treatment.
PUBLIC HEAI.TH CONSIDERATIONS
In sorne parts of tropical Africa man is frequently infected (Bouckaert and Fain 1959; Cannon 1942), but the parasite in endothermal laboratory species is of no public health importance. Man can only be infected by ingesting eggs passed in snake feces or saliva.
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502 PARASITES OF ENDOTHERMAL LABORATORY ANIMALS
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Gurr, L. 1953. A note on the occurrence oE Linguatula serrata (Frohlich, 1789) in the wild rabbit, Oryctolagus CUlljeulus, in New Zealand. New Zealand J. Sei. Technol. 35: 49-50.
Heymons, R. 1935. Pentastomida, pp. 1-268. In H. G. Bronn's h.lassen und Ordnungen des Tierreichs. Vol. 5, Sect. 4, Bk. 1. Akademische VerlagsgesellschaEt M.B.H., Leipzig.
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Koch, M. 1907. Zur h.ennmis des Parasitismus der Pentastomen. VerhandI. Deut. Ges. Pathol. 17:265-79.
Kuntz, R. E., Betty J. Myers, and T. E. Vice. 1967. Intestinal protozoans and parasites oE the gelada baboon (Theropithecus gelada Rüppel, 1835). Proc. Helminthol. Soc. Wash. D.C. 34:65-66.
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PENTASTOMIDS
Linguatula seT1'ata (Frohlich, 1789) in AusnaUa. Australian Veto J. 12:61-64.
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Sti1es, C. W. 1895. On the recent occurrence of Linguatula rhinaria, Railliet, 1886, and Taenia echinococcus, V. Siebold, 1853, in the United States of North America. Trans. Fint Pan-Am. Med. Congr. Wash., D.C. (Part 2): Il63-65.
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5°3
of the Hamilton Rice Expedition to the Amazon. Harvard Univ. Press, Cambridge, Mass.
Tubangui, M. A., and Victoria A. Masiluiigan. 1936. Notes on Philippine linguatulids (Arthropoda: Pemastomida). Philippine J. Sei. 60:399-403.
Whitney, R. A., Jr., and S. M. Kruckenberg. 1967. Pentastomid infection associated with peritonitis in mangabey monk.eys. J. Am. Veto Med. Assoc. 151:907-8.
Worms, M. J. 1967. Parasites in newly imported animaIs. J. Inst. Animal Tech. 18: 39-47.
Yamashita, J., and M. Ohbayashi. 1954. On a tongue worm, Linguatula SeT1'ata Frolich, 1789, showing the sarne parasitism as in the intermediate hast within a dog body (in Japanese). Mem. Fac. Agr. Hokkaido Univ. 2:146-48.