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avian trichomoniasis

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avian trichomoniasis

Summary

  • Last modified
  • 03 January 2018
  • Datasheet Type(s)
  • Animal Disease
  • Preferred Scientific Name
  • avian trichomoniasis
  • Overview

  • Trichomonas gallinae (Rivolta, 1878) is a flagellate protozoan that infects a variety of birds, especially columbids (doves and pigeon...

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Pictures

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PictureTitleCaptionCopyright
Interaction between Trichomonas gallinae and erythrocytes. Figs.a), b) show the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Figs. c) and d) show the adherence between parasites and red blood cells after 90 min. E: erythrocyte; T: trichomonad; AF: anterior flagella; AX: axostyle; UM: undulating membrane. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
TitleInteraction between Trichomonas gallinae and erythrocytes
CaptionInteraction between Trichomonas gallinae and erythrocytes. Figs.a), b) show the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Figs. c) and d) show the adherence between parasites and red blood cells after 90 min. E: erythrocyte; T: trichomonad; AF: anterior flagella; AX: axostyle; UM: undulating membrane. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
Copyright2002 Elsevier BV
Interaction between Trichomonas gallinae and erythrocytes. Figs.a), b) show the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Figs. c) and d) show the adherence between parasites and red blood cells after 90 min. E: erythrocyte; T: trichomonad; AF: anterior flagella; AX: axostyle; UM: undulating membrane. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
Interaction between Trichomonas gallinae and erythrocytesInteraction between Trichomonas gallinae and erythrocytes. Figs.a), b) show the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Figs. c) and d) show the adherence between parasites and red blood cells after 90 min. E: erythrocyte; T: trichomonad; AF: anterior flagella; AX: axostyle; UM: undulating membrane. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).2002 Elsevier BV
Interaction between Trichomonas gallinae and erythrocytes. Fig. a) shows the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Fig. b) shows adherence between both cells after 120 min. E: erythrocyte; T: trichomonad; AF: anterior flagellum. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
TitleInteraction between Trichomonas gallinae and erythrocytes
CaptionInteraction between Trichomonas gallinae and erythrocytes. Fig. a) shows the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Fig. b) shows adherence between both cells after 120 min. E: erythrocyte; T: trichomonad; AF: anterior flagellum. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
Copyright2002 Elsevier BV
Interaction between Trichomonas gallinae and erythrocytes. Fig. a) shows the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Fig. b) shows adherence between both cells after 120 min. E: erythrocyte; T: trichomonad; AF: anterior flagellum. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).
Interaction between Trichomonas gallinae and erythrocytesInteraction between Trichomonas gallinae and erythrocytes. Fig. a) shows the contact between the T. gallinae isolate (TG9) and erythrocytes after 60 min. Fig. b) shows adherence between both cells after 120 min. E: erythrocyte; T: trichomonad; AF: anterior flagellum. Arrows show the closed adherence between parasites and erythrocytes suggesting erythrophagocytosis (From De Carli GA, Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 87:277-283, with permission from Elsevier).2002 Elsevier BV

Identity

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Preferred Scientific Name

  • avian trichomoniasis

International Common Names

  • English: avian trichomonosis; canker; frounce; roup; Tichomonas gallinae infection in birds; trichomoniasis, trichomonas gallinae, tetratrichomonas gallinarum, in birds; trichomonosis, avian; Upper digestive tract trichomoniasis

Overview

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Trichomonas gallinae (Rivolta, 1878) is a flagellate protozoan that infects a variety of birds, especially columbids (doves and pigeons) worldwide (BonDurant and Honigberg, 1994). This trichomonad species occurs in the upper digestive tract, and some virulent isolates can spread to other parts of the body. T. gallinae is the organism originally described by Rivolta (1878) as Cercomonas gallinae from the mouth and crop of a pigeon. The taxonomic position of the genus Trichomonas was reported by Levine et al. (1980):

Kingdom: Protista
Subkingdom: Protozoa
Phylum: Sarcomastigophora Honigberg and Balamuth, 1963
Subphylum: Mastigophora Diesing, 1866
Class: Zoomastigophorea Calkins, 1909
Order: Trichomonadida Kirby, 1947 emend. Honigberg, in Camp, Mattern and Honigberg, 1974
Family: Trichomonadidae Honigberg, 1963
Subfamily: Trichomonadinae Honigberg, 1963

T. gallinae, a very common parasite of pigeons, causes serious losses in these birds, and also in turkeys. In columbids, T. gallinae is typically transferred in pigeon ‘milk’ from the crop of an infected parent bird to the offspring within a short time after the latter have hatched from the eggs. All the offspring become infected. Once acquired, the infection can be very long-lasting, some infections persisting for up to 2 years (BonDurant and Honigberg, 1994).

Experimental infections have been used in pathogenic studies, especially subcutaneous mouse assay (Honigberg, 1961; Frost and Honigberg, 1962). A number of birds have been infected experimentally. They include the bobwhite quail (Colinus virginianus), canary (Serinus canaria), English sparrow (Passer domesticus), barn swallow (Hirundo rustica), goldfinch and song sparrow (Melospiza melodia), Tovi parakeet (Brotogeris jugularis) and cardinal (Cardinalis cardinalis) (Levine, 1973). Parenteral infections have also been produced experimentally in rats and kittens (Levine, 1973). T. gallinae caused abscesses following subcutaneous injection in mice (Frost and Honigberg, 1962).


Transmission


T. gallinae is an exceptionally hardy parasite, and can survive sub-optimal environmental conditions without a cyst form (indeed, no such form is known in the genus). Survival in tap water for short periods ensures that drinking water can be a source of infection for domestic fowl. An individual bird may lose the parasite, but since pigeons live in groups, the close contact among them ensures reinfection. The behaviour of pigeons is conducive to spread of the parasite. The behaviours important in transmitting infection are billing during courtship and feeding (involving ‘pigeon milk’) of the young by infected parent birds. However, adults can also be reinfected from water sources frequented by infected pigeons. An infection may also be established in a raptor that has fed on an infected prey bird (BonDurant and Honigberg, 1994).

The spread of T. gallinae to domestic fowls such as turkeys and chickens, is dependent primarily on obtaining the parasite from drinking water from sources visited by feral pigeons. It seems that, in view of the feeding habits of domestic fowls, in these birds epizootics of T. gallinae are dependent on the presence of pigeons as the reservoir hosts. In chickens and turkeys, the organism may be transmitted in drinking water, but not from one generation to the next as in the Columbiformes.

Since T. gallinae is not found in the intestinal tract beyond the muscular stomach, the parasite cannot be transmitted in faecal droppings. The routes of escape can only be the nares, eyes and mouth, the last being the most important portal of exit. In all birds other than columbids, water is the main medium of infection (BonDurant and Honigberg, 1994).


Age predisposition


In the pigeon, trichomonosis is mainly a disease of young birds; 80-90% of the adults are infected but show no signs of disease (Levine, 1973).

Distribution

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Trichomonas gallinae infects a variety of birds all over the world. The domestic pigeon, Columba livia, is the primary host of this flagellate. Other columbiform hosts have been found to harbour the parasite, as have domestic fowls (especially turkeys), Java sparrows, various raptors, and sea gulls. Experimental infections have been established in many kinds of passerine birds.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

BahrainPresentSamour et al., 1995
ChinaPresentZhang et al., 1982
IndiaPresentBhattacharya et al., 1997
IranPresentBozorgmehri-Fard and Moeinvaziri, 1985
Saudi ArabiaPresentSilvanose et al., 1998; Samour and Naldo, 2003
TurkeyPresentAydin et al., 2000
United Arab EmiratesPresentSilvanose et al., 1998

Africa

EgyptPresentAbd-El-Motelib and Galal, 1994
South AfricaPresentHuchzermeyer, 1978; Pepler and Oettlé, 1992

North America

CanadaPresentPresent based on regional distribution.
-British ColumbiaPresentRosenfield et al., 2002
MexicoPresentMoedano and Moreno, 1976
USAPresentPresent based on regional distribution.
-ArizonaPresentBoal et al., 1998; Rosenfield et al., 2002
-CaliforniaPresentRupiper and Harmon, 1988; Willoughby et al., 1995
-ColoradoPresentStabler, 1954
-ConnecticutPresentSasseville et al., 1988
-FloridaPresentConti and Forrester, 1981
-HawaiiPresentKocan and Banko, 1974
-New YorkPresentTangredi, 1978; Stone and Nye, 1981
-North DakotaPresentRosenfield et al., 2002
-TexasPresentGlass et al., 2001
-UtahPresentOstrand et al., 1995
-WisconsinPresentRosenfield et al., 2002

Central America and Caribbean

Trinidad and TobagoPresentKaminjolo et al., 1988

South America

BrazilPresentPresent based on regional distribution.
-Rio Grande do SulPresentTasca and Carli, 1999a; De et al., 1979
-Sao PauloPresentPereira and Almeida, 1943
ChilePresentToro et al., 1999

Europe

CroatiaPresentGreguric et al., 1986
GreecePresentGithkopoulos and Liakos, 1987
ItalyPresentCatelli et al., 1999
MacedoniaPresentKulisic et al., 1996
PolandPresentNativeWieliczko et al., 2003
PortugalPresentPonce et al., 2002
Russian FederationPresentGreguric et al., 1986
SloveniaPresentDobeic, 2003; Dovc et al., 2004
SpainPresentMartínez-Moreno et al., 1989; Ponce et al., 2002
UKPresentCooper and Petty, 1988; Cousquer, 2003
Yugoslavia (former)PresentKulisic et al., 1996

Oceania

AustraliaPresentMcKeon et al., 1997

Pathology

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Lesions of T. gallinae trichomonosis appear as firm, yellow, caseous areas. In the brains and liver they may replace considerable amounts of the organ involved. In the upper digestive tract they may appear as huge, firmly attached masses (mouth) or as disc-shaped lesions -‘yellow buttons’- with central, spur-like projections (oesophagus and crop). Massive lumps may occur in the tissues of the neck, or encircling caseous regions may involve the mucosa of the oesophagus, crop, or proventriculus (Samour et al., 1995).

Certain isolates of the parasite appear to have a predilection for some tissues. The Jones’ Barn isolate almost invariably attacks the liver of infected pigeons, only moderately involving the regions of the head and neck. The Mirza isolate, on the other hand, causes extensive damage to the tissues of the head and the mucosa of the upper digestive tract, only rarely entering the abdominal cavity (Perez-Mesa et al., 1961; BonDurant and Honigberg, 1994).

The lining of the mouth, pharynx, oesophagus crop, and glandular stomach have commonly been involved. The head sinuses, orbital regions, and brain are sometimes infected. Caseous masses in the neck tissues are common. The most frequently involved visceral organ is the liver. The liver surface may become almost entirely covered with caseous lesions which, by contact extension, may involved the intestinal and gizzard surfaces, sub-sternal membranes, air sacs, lungs and pericardium. The myocardium may become caseous, apparently as an extension of processes in the pericardium. The pancreas is also sometimes infected. In young squabs the navel region is sometimes so thoroughly involved that the skin and adjacent tissues, as well as the local organs, are cemented in one huge caseous mass. The caseous lesions of the upper digestive tract may become so extensive that they completely occlude the lumen, making passage of food an impossibility. Large quantities of fluid frequently accumulate in the crop.

The more rarely involved sites include spleen, kidney, trachea and upper respiratory tract, bone marrow, and the middle ear. It has been noted by several observers that lesions of the upper digestive tract frequently end abruptly at the glandular stomach (Samour et al., 1995).

The histopathology of infection with T. gallinae has not been extensively treated, although some observations have been made (Stabler, 1954).


Factors affecting pathogenicity


In vitro cultivation of T. gallinae affects inherent pathogenicity levels in infections of mice (Honigberg, 1961) and pigeons (Stabler et al., 1964). In mice there is a progressive decrease in the mean volume of subcutaneous abscesses with cultivation time (Honigberg, 1961). In avian hosts, after between 17 and 21 weeks of cultivation, virulence for pigeons was lost by the JB isolate, although it could be restored after at least up to 28 weeks of cultivation, by serial passages in non-immune pigeon squabs (Stabler et al., 1964).

It was also demonstrated by Stabler et al. (1964) that the presence of antibiotics, such as penicillin and streptomycin, in the culture medium results in more rapid attenuation of virulence. When isolated in the presence of these antibiotics, the JB isolated became avirulent within 7.5 to 9 weeks of in vitro culture.

Diagnosis

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Clinical examination


Upper digestive tract trichomonosis (oral cavity, choanae, tongue, pharynx, crop, upper oesophagus and infraorbital sinuses) is readily diagnosed by observation of the lesions together with finding of the protozoa (Levine, 1973; Samour et al., 1995).


Laboratory examination


The throats of pigeons are swabbed and the swabbings used to inoculate trypticase-yeast extract-maltose (TYM) medium for incubation. Samples are cultured axenically in vitro,without antibiotics, in a TYM medium without agar; pH 7.2, supplemented with 10% (v/v) heat inactivated horse serum, at a temperature of 37°C (±0.5). Some cultures can be established without the use of antibiotics, others are isolated with 5000 units of penicillin and 1000 µg of dihydrostreptomycin per ml of the original culture. Dying of parasites is strongly recommended to observe morphological features. Parasites from a 24-h culture in TYM medium are stained by the Giemsa method (Carli et al., 1979; Tasca and Carli, 1999).

List of Symptoms/Signs

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SignLife StagesType
Digestive Signs / Anorexia, loss or decreased appetite, not nursing, off feed Sign
Digestive Signs / Congestion oral mucous membranes, erythema, redness oral mucosa Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Dysphagia, difficulty swallowing Sign
Digestive Signs / Erythema, redness of pharynx Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Esophageal obstruction Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Excessive salivation, frothing at the mouth, ptyalism Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Inability to open (trismus) and / or close jaw, mouth Sign
Digestive Signs / Oral mucosal ulcers, vesicles, plaques, pustules, erosions, tears Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Pharyngeal ulcers, vesicles, erosion, papules, sores pharynx Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Digestive Signs / Tongue protrusion Sign
Digestive Signs / Tongue ulcers, vesicles, erosions, sores, blisters, cuts, tears Sign
General Signs / Ataxia, incoordination, staggering, falling Sign
General Signs / Dysmetria, hypermetria, hypometria Sign
General Signs / Increased mortality in flocks of birds Sign
General Signs / Lack of growth or weight gain, retarded, stunted growth Sign
General Signs / Laryngeal, tracheal, pharyngeal swelling, mass larynx, trachea, pharynx Sign
General Signs / Neck swelling, mass cervical region Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Diagnosis
General Signs / Oral cavity, tongue swelling, mass in mouth Sign
General Signs / Orbital, periorbital, periocular, conjunctival swelling, eyeball mass Sign
General Signs / Sudden death, found dead Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
General Signs / Underweight, poor condition, thin, emaciated, unthriftiness, ill thrift Sign
General Signs / Weight loss Sign
Nervous Signs / Dullness, depression, lethargy, depressed, lethargic, listless Sign
Ophthalmology Signs / Blindness Sign
Ophthalmology Signs / Lacrimation, tearing, serous ocular discharge, watery eyes Sign
Ophthalmology Signs / Purulent discharge from eye Sign
Pain / Discomfort Signs / Mouth, oral mucosal or tongue pain Sign
Respiratory Signs / Abnormal breathing sounds of the upper airway, airflow obstruction, stertor, snoring Sign
Respiratory Signs / Dyspnea, difficult, open mouth breathing, grunt, gasping Sign
Respiratory Signs / Epistaxis, nosebleed, nasal haemorrhage, bleeding Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Diagnosis
Respiratory Signs / Mucoid nasal discharge, serous, watery Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign
Respiratory Signs / Nasal mucosal ulcers, vesicles, erosions, cuts, tears, papules, pustules Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Diagnosis
Respiratory Signs / Purulent nasal discharge Poultry:Day-old chick,Poultry:Young poultry,Poultry:Mature female,Poultry:Cockerel,Poultry:Mature male Sign

Disease Course

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Trichomonas gallinae causes a variety of pathological manifestations depending on the isolate of the parasite and species of bird infected (Jessup, 1980; Baker, 1986; Cooper and Petty, 1988; Henderson et al., 1988). The normal sites of this parasite are the mouth, pharynx, oesophagus and crop, with most of the isolates never leaving the upper digestive tract. The virulent isolates may cause lesions in the upper digestive tract of birds; some of these isolates spread to other parts of the body. The parasite initially infects the upper digestive tract, causing the formation of ulcers in the mouth, sinuses, orbital region, pharynx, oesophagus, crop and even the proventriculus (Levine, 1973), which allow the pathogen to enter the circulatory system. The trichomonads later gain access to the liver, and cause the formation of caseous lesions. Non-immune pigeons may die of liver dysfunction within 14 to 17 days after inoculation. Other virulent isolates include those that invade the head and neck sinuses and from there enter the eye regions and brain. Other virulent T. gallinae establish in the organs of the thoracic and abdominal cavities, such as, lungs, heart, liver, and pancreas. Transmission of T. gallinae is horizontal (BonDurant and Honigberg, 1994).

Epidemiology

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Trichomonas gallinae infects pigeons and a variety of birds, including turkeys, and has caused important economic losses in this species as well as in chickens and pet birds (BonDurant and Honigberg, 1994). In pigeons and mourning doves, trichomonosis is transmitted from adults to the squabs in the pigeon milk, which is produced in the crop. The squabs are infected within minutes after hatching. Hawks and other wild raptors become infected by eating infected birds. Turkeys and chickens are infected through contaminated drinking water. Feral pigeons and other columbid birds are usually the original source of infection (Levine, 1973).

Impact: Economic

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Trichomonas gallinae is very common in domestic pigeons, in which it often causes serious losses. Present data regarding economic losses are not available. In 1954 the US Department of Agriculture estimated that the parasite causes an annual loss of US $47,000 in turkeys; in 1965, it was estimated that the flagellate caused an annual loss of US $23,000 in chickens and US $1,186,000 in turkeys. T. gallinae is common in mourning doves, and may cause serious losses in these birds (Levine, 1973).

Disease Treatment

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Treatment is only feasible in captive birds because the drugs used for treatment must be administered orally, either by force feeding or by treating the food and/or water. Antiprotozoal medications that have been used are dimetridazole, metronidazole, copper sulfate, quaternary ammonia, carnidazole, 2-amino-5-nitrothiazole and aminonitrothiazole. Some of these medications are under review and some can be specifically used only on non-food birds (metronidazole) (Frank, 2004). Ronidazole presents excellent activity against T. gallinae with high safety margin (Samour and Naldo, 2003). Dimetridazole was formulated for drinking water medication and as a prolonged-release tablet. To suppress T. gallinae infection, successfully, medicated drinking water containing dimetridazole (400 mg/L) has to be administered for at least 3 days (Inghelbrecht et al., 1996).

Treatment of wild birds is difficult because of the availability of natural food and water sources.

Resistance of T. gallinae isolates to the nitroimidazole drugs ronidazole, carnidazole and metronidazole has been reported (Franssen and Lumeij, 1992; Munoz et al., 1998). Although ronidazole showed a greater potency than the other nitroimidazole derivatives, 8-22 times more drug was necessary for efficacy compared with the others (Munoz et al., 1998).

Prevention and Control

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Effective treatment of a population of susceptible birds involves removal of the source of infective organisms and treatment of treatable infected individuals. Source removal often requires the destruction of chronically infected birds and protection of water sources from wild birds. Drugs have been employed, either in drinking water or topically in the mouth and throats of the birds. Older remedies included anise and aniline oil, tartar emetic, acriflavin, iodine with or without glycerine, carbolic acid with glycerine, ‘sulfoliquid’, sulfathizole and sulfanilamide, nitrate stick, powdered alum, and writers ink (Stabler, 1954; BonDurant and Honigberg, 1994).

In addition, the following preventive measures are advised: practice a high standard of sanitation at all times; do not add birds to an established flock without quarantine for 30 days; and provide a source of clean, fresh water and eliminate all sources of stagnant water (Butcher, 2003). In captive birds, cull or treat carrier birds, cull adults whose offspring become infected, regularly disinfect food and water sources with 10% bleach solution, screen out (exclude) wild birds to protect from contamination by wild pigeons and other birds, segregate young birds from adults, and segregate susceptible birds from recovered or carrier birds. To minimize infections in wild birds, minimize dove and pigeon concentrations at feeders (or do not feed) and at water receptacles. Severely affected captive or wild birds should be killed.

References

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Abd-El-Motelib TY; Galal BEl-G, 1994. Some studies on Trichomonas gallinae infection in pigeons. Assiut Veterinary Medical Journal, 30(59):277-288; 19 ref.

Abraham R; Honigberg; BM, 1964. Structure of Trichomonas gallinae (Rivolta). Journal of Parasitology, 50:608-619.

Almond C; Dowda H, 1976. Trichomonas gallinae: characterization and regulatory properties of lactic dehydrogenase. Experimental Parasitolology, 40:25-32.

Aydin L; Coskun S; Gülegen E, 2000. Efficacy of carnidazole (Spatrix) and metronidazole (Flagyl) against Trichomonas gallinae in naturally infected pigeons. Acta Parasitologica Turcica, 24(1):65-66; 12 ref.

Baker JR, 1986. Trichomoniasis, a major cause of vomiting in budgerigars. Veterinary Record, 118(16):447-449; 6 ref.

Benchimol M; Leal D; Mattos A; Diniz JAP, 1997. Fine structure of Trichomonas gallinae. Biocell, 21:47-58.

Betterton H; Almond C; Dowda H, 1976. Trichomonas gallinae: characterization and regulatory properties of lactic dehydrogenase. Experimental Parasitology, 40:25-32.

Betterton H; Dowda H, 1976. An analysis of oxidative metabolism in Trichomonas gallinae. Comparative Biochemistry and Physiology, Part B, Biochemistry and Molecular Biology, 54:163-165.

Bhattacharya HM; Laha R; Bhattacharya D, 1997. Trichomonosis in pigeons. Indian Veterinary Journal, 74(4):341; 5 ref.

Blood DC; Studdert VP, 1988. Baillière's comprehensive veterinary dictionary. Baillière's comprehensive veterinary dictionary., xii + 1124pp.; [52000 entries].

Boal CW; Mannan RW; Hudelson KS, 1998. Trichomoniasis in Cooper's hawks from Arizona. Journal of Wildlife Diseases, 34(3):590-593; 21 ref.

Bozorgmehri-Fard MH; Moeinvaziri M, 1985. Report of an outbreak of trichomoniasis in domestic pigeons (Columba livia). Journal of Veterinary Faculty, University of Tehran, 40(2/4):73-79; 7 ref.

Butcher GD, 2003. Pigeon Canker. University of Florida. IFAS Extension. Online at (http://edis.ifas.ufl.edu).

Carli GAde; Brasseur P; Rott MB; Silva ACda; Wendorff A; Benchimol M, 1994. Determination of hemolytic activity of different strains of trichomonads of genus Trichomonas Donné, 1836 and Tritrichomonas Kofoid, 1920. Journal of Protozoology Research, 4(4):158-163; 28 ref.

Carli GAde; Tasca T, 2002. Trichomonas gallinae: a possible contact-dependent mechanism in the hemolytic activity. Veterinary Parasitology, 106(4):277-283; 27 ref.

Catelli E; Poglayen G; Terregino C; Orlando C; Tonelli A; Issa Gadale O; Roda R; Agnoletti A, 1999. Survey of endoparasites of the digestive tract of Columba livia (Gmelin, 1789) in Florence. Selezione Veterinaria, No. 2:75-85; 34 ref.

Conti JA; Forrester DJ, 1981. Interrelationships of parasites of white-winged doves and mourning doves in Florida. Journal of Wildlife Diseases, 17:529-536.

Cooper JE; Petty SJ, 1988. Trichomoniasis in free-living goshawks (Accipiter gentilis gentilis) from Great Britain. Journal of Wildlife Diseases, 24(1):80-87; 19 ref.

Cousquer G, 2003. Trichomoniasis in tawny owls (Strix aluco) from the south west of England. Journal of Wildlife Diseases, 39:11-12.

Daly JJ, 1970. The maltose metabolism of Trichomonas gallinae (Rivolta, 1878). I. Growth studies. Journal Parasitology, 56:883-888.

Daly JJ, 1971. The maltose metabolism of Trichomonas gallinae (Rivolla, 1878). II. Journal Parasitology, 57:370-374.

Daly JJ; Mathews HM; Koch WC; Bost JW, 1974. The effect of different carbohydrates on the glycogen content and growth of Trichomonas gallinae. Comparative Biochemistry and Physiology, Part B, Biochemistry and Molecular Biology, 48:651-659.

De Carli GA; Pansera MCG; Guerrero J, 1979. Trichomonas gallinae (Rivolta, 1878) Stabler, 1938, no trato digestivo superior de pombos domésticos, Columba livia, no Rio Grande do Sul – Primeiro Registro. Acta Biológica Leopoldensia, 1:85-95.

Diamond LS, 1954. A comparative study of 28 culture media for Trichomonas gallinae. Experimental Parasitology, 3:251-258.

Diamond LS, 1957. The establishment of various trichomonads of animals and man in axenic cultures. Journal of Parasitology, 43:48-490.

Diamond LS, 1964. Freeze-preservation of protozoa. Cryobiology, 1:95-102.

Dobeic M, 2003. Regulation of population size of street pigeons in Ljubljana, Slovenia. Acta Vererinaria Beograd, 53:171-182.

Dovc A; Zorman-Rojs O; Vergles Rataj A; Bole-Hribovsek V; Krapez U; Dobeic M, 2004. Health status of free-living pigeons (Columba livia domestica) in the city of Ljubljana. Acta Veterinaria Hungarica, 52:219-26.

Dowda H Jr; Betterton HO, 1974. Purification and properties of malic dehydrogenase from Trichomonas gallinae. Experimental Parasitology, 36:415-423.

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