CESTODES

1 are extremely common Cestodes or tapeworms (class Cestoda, phylum Platy- helminthes)
parasites of birds. Most species infect the intestine, but a few can be found in the ceca or under the gizzard lining. hey are readily distinguished from other worm parasites (trematodes, nematodes, and acanthocephalans) by their segmented appearance. !irds ha"e the most di"erse cestode fauna of any "ertebrate group. #ild birds are often infected with large numbers of cestodes and a"erage pre"alence can be $uite high. %epending on host species, apparently healthy birds may be infected with tens, hundreds, or, in some cases, thousands of cestodes SYNONYMS Cestodiasis, hymenolepididiasis, drepanidotaeniasis, fimbriariasis, gastrotaeniasis. ETIOLOGY Cestodes belong to the phylum Platyhelminthes. Most species infect the intestine, a few species infect the ceca, and Gastrotaenia infects the gizzard. &ccasionally, cestodes in"ade abnormal sites including the ureters ) and the gizzard muscle. 'dult cestodes are white and translucent when ali"e. hey range from (-) mm to ( m long , but many are less than (* cm. Cestode bodies consist of a holdfast (scolex), a short nec+, and a body (strobila) made up of repeated units (proglottids) that gi"e it a segmented appearance. ' mature strobila consists of three zones, a zone of immature proglottids posterior to the scolex, a zone of sexually mature proglottids with functional reproducti"e systems, and a postreproduc- ti"e (gra"id) zone consisting of proglottids that contain eggs ready for dispersal from the host. hree of the (- orders recognized '.% are represented in the cestode fauna of birds. 'bout /* species belong to the orders etrabothri- idea and Pseudophyllidea and the remainder belong to the Cyclophyllidea . Most cyclophyl- lidean species that infect birds are found in the families 0ymenolepididae, %ilepididae, and %a"aineidae. Molecular e"idence indicates that the Cyclophyl- lidea is the most highly deri"ed order. The Tetrabothriidea is its closest relati"e and the Pseudophyllidea occupies a more basal position. he Pseudophyllidea is polyphyletic with only one family that is found in birds, the %iphyllobothriidae. 1ecent e"idence indicates that the %iphyllobothriidae is independent and ancestral to the Pseudophyllidea, but formal classification of the cestodes has yet to reflect molecular results. hese three groups can be distinguished by morphology of the scolex and mature proglottids . Cyclophyllidean scolices ha"e four muscular suc+ers. Most species also ha"e a rostellum (a muscular organ within the scolex) that can be pro2ected from its apex. he rostellum is usually armed with hoo+s 1

and the number, shape, and size of these are of taxo- nomic importance. etrabothriidean scolices ha"e

2 four large leafli+e suc+ers called bothridia and lac+ a rostellum. 3colices of the diphyllobothriids ha"e
one dorsal and one "entral groo"e (bothria) instead of suc+ers or bothridia.

MORP OLOGIC!L

COMP!RISON

O"

SCOLICES

!ND

M!T#RE

PROGLOTTIDS

O"

!D#LT

DIP YLLO$OT RIID% TETR!$OT RIID% !ND CYCLOP YLLIDE!N CESTODES& Cestodes ha"e complex reproducti"e systems. Mature proglottids of these three groups can be distinguished by the type and location of the "itelline (yol+) glands, the structure of the uterus, the position of the genital pore, and by the presence or absence of a uterine pore

! partial list o' cestode species that ha(e bee) reported to cause patholo*y% disease% or +ortality i) ,ild birds.
Parasite !D#LT CESTODES ORDER CYCLOP YLLIDE! "a+ily y+e)olepididae Gastrotaenia cygni Gastrotaenia dogieli Distributio) ost order Sourc e

.', 3' 4u, 's

' '

# # 4 4

Microsomacanthus parvula Dicranotaenia coronula Hispaniolepis falcata "a+ily Da(ai)eidae Otiditaenia conoideis Otiditaenia macqueeni Raillietina sp. "a+ily Dilepididae Choanotaenia infundibulum Parorchites zederi "a+ily Gryporhy)chidaeParadilepis delachau i

4u, 's, .' 4u, 's, 'f, 'f, 's 'f, 4u, 's 'f, 's .' C 'ntarctica 'f, 's

' 5 '6, Ch, 7al, 7r C, %5 7r # 7r 7r 7al ', Co, 8, 7al6, 7r, Pa, 3t Pr Pe C C # C, %5 # # #

Parasite "a+ily Diphyllobothriidae !chistocephalus solidus

%istribution C

0ost order

3ource

', Ch6, Ci, Co, # 8, 7al, 7a", Paradilepis scolecina C Pe 7r, Pa, Pe, Paradilepis sp. 's Pe Po, Pr "a+ily !+abiliidae # Po !chistotaenia scolopendra 3' # Po !chistotaenia srivastavi .' %igula intestinalis !chistotaenia C tenuicirrus ', Ci,.' Ch6, 8, # Po Pe6, Po6, # "a+ily Paruteri)idae "scometra choriotidis 's Pa 7r 7a"6, Metroliasthes lucida .' 7al L!R.!L CESTODES "a+ily Mesocestoididae .' "ia 'f # Ph Mesocestoides sp.Cyclophyllidean C sp. Mammals ORDER TETR!$OT RIIDE! # #etrabothrius s$oogi 's Pr # #etrabothrius sp. 'u Pr "a+ily Diphyllobothriidae %igula intestinalis C 3ee abo"e C !pirometra sp. C Mammals #

# # # # #, 4 C # C # #

0ost orders, ', 'nseriformes9 Ca, Caprimulgiformes9 Ch, Charadriiformes9 Ci, Ciconiiformes9 Co, Columbiformes9 8, 8alconiformes9 7al, 7alliformes9 7a", 7a"iiformes9 7r, 7ruiformes9 Pa, Passeriformes9 Pe, Pelecaniformes9 Ph, Phoenicopteriformes9 Pi, Piciformes9 Po, Podicipediformes9 Pr, Procellariiformes9 3tr, 3trigiformes9. 'steris+s when present indicate the ma2or host order(s) of a particular species. 3ource of material, #, wild9 4, experimental9 C, capti"e9 %, domestic9 5, source un+nown. %istribution, .', .orth 'merica9 3', 3outh 'merica9 4u, 4urope9 's, 'sia9 C, Cosmopolitan9 'f, 'frica.
DLP YLLO$OT RIID!E

wo intermediate hosts are re$uired to complete %i- phyllobothriid life cycles . he first host is a copepod and the second is a "ertebrate, usually a fish. he exception is !pirometra whose species use all "ertebrates except fish as second intermediate hosts.

Represe)tati(e li'e cycles o' cyclophyllidea) a)d diphyllobothriid cestodes& Lar(al sta*es o' the Gryporhy)chidae

he eggs are thic+ shelled, operculate, and re$uire a period of de"elopment in water before hatching. he egg e"entually releases a coracidium lar"a, which 4 is essentially an oncosphere surrounded by a ciliated co"ering. he oncosphere penetrates the gut of the copepod host and de"elops into a procercoid lar"a in the hemo- coel. #hen eaten by a fish, the procercoid penetrates the host gut, resumes de"elopment in the body ca"ity, "isceral organs or musculature, and transforms into the pleurocercoid stage. his stage is infecti"e to the a"ian host. CLINIC!L SIGNS Cestode infections in wild birds are normally asymptomatic, but when clinical signs are present, they are nonspecific and similar to those reported in poultry . 4maciation, wea+ness, and occasionally diarrhea and hemorrhage may be accompanied by changes in posture or locomotory and feeding beha"ior. 'mong naturally infected wild birds, emaciation has been obser"ed in common eiders infected with !chis& tocephalus solidus and %ateriporus sp. in a :ong-tailed %uc+ 'Clangula hyemalis( infected with G) cygni , in 3hort-tailed 3hearwaters 'Puffinus tenuirostris( and :ittle Penguins '*udyptula minor) infected with #etrabothrius spp., and in an 'rctic :oon infected with %igula in& testinalis . ;n the latter case, the loon also displayed generalized wea+ness and diarrhea. ' few species of cestodes in duc+s and grebes can cause "arying degrees of hemorrhage where they attach to the intestinal mucosa that might be detected as bloody feces. DI!GNOSIS he presence of eggs, gra"id proglottids, or cestode fragments in the feces of the host is diagnostic for cestode infection. Cyclophyllidean and tetrabothriidean cestodes can be distinguished from di- phyllobothriid cestodes on the basis of egg morphology and proglottid structure . %i- phyllobothriid eggs ha"e hard operculate shells , but may be difficult to distinguish from eggs produced by trematodes. he gra"id proglottids of some families are sufficiently distinct to permit identification to that le"el. ;dentification of cestodes to order, family, and generic le"els re$uires microscopic study of adult specimens and e"aluation of the morphology of the scolex and reproducti"e systems . ;dentification to species le"el re$uires e"aluation of the presence or absence of the rostellar hoo+s on the scolex and their number, shape, and size. he position, size, and shape of components of the male and female reproducti"e systems are also important, including the number of testes and their spatial relationships within the mature proglottid.. P#$LIC E!LT CONCERNS/ 'dult cestodes found in birds cannot be transmitted directly to humans and do not pose a health threat. 0owe"er, the lar"al stages of Mesocestoides and !pirometra can infect humans when consumed in raw or undercoo+ed meat . DOMESTIC !NIM!L E!LT CONCERNS ransmission of cestodes from wild to domestic birds re$uires a wild reser"oir host to pro"ide a source of cestode eggs and intermediate hosts to support de"elopment of lar"al stages of the parasites. he practice of raising duc+s and geese on reser"oirs and natural wetlands fre$uented by wild waterfowl can lead to outbrea+s of cestode infection, disease, and losses in domestic waterfowl . :osses of domestic waterfowl as a result of infection with G) dogieli ha"e been documented in 4astern 4urope and the for-

mer <331 . 3imilarly, contamination of local ponds by wild anatids can also lead to outbrea+s in capti"e waterfowl in zoological collections . Cestode di"ersity in capti"e birds at the #arsaw =oo was less than that in wild birds, presumably because a suitable range of intermediate hosts was not present. 0ILDLI"E POP#L!TION IMP!CTS he potential influence of parasites on host population dynamics is difficult to assess. 'dult cestodes are not generally considered pathogenic or a threat to a"ian populations under normal conditions . 'bsence of clinical signs ma+es it difficult to detect infected birds. 3ic+ birds usually die unnoticed and, if they are found, they are usually infected with a "ariety of parasites, ma+ing it impossible to attribute the condition to a specific agent. 1eports of mortality in the field need to be interpreted with caution, particularly if mixed infections are in"ol"ed. 8or example, cestodes ha"e been associated with emaciation and star"ation of large numbers of birds during sudden cold snaps ,but theirrole in these deaths remains unresol"ed. TRE!TMENT !ND CONTROL #ild birds brought in from the field for propagation or relocation programs are usually infected with cestodes. 3tress associated with capture or confinement may exacerbate the effects of cestode infections. reatment recommendations "ary but in general )iclosa+ide has been recommended for species of 7ruiformes , 'nseriformes (except for geese), 8alconiformes, and 3trigiformes . Pra1i2ua)tel is effecti"e in species of Columbiformes, starlings and other 3turnidae , and in bustards. !oth niclosamide and fluben- dazole are effecti"e in controlling infection in capti"e floc+s of bustards. &striches infected with Houttuynia struthionis was successfully treated with 'e)be)da1ole& here is no practical way to control cestode infections in wild birds. Control measures re$uire a disruption in life cycles by reducing or eliminating contact with potential intermediate hosts. his may be possible on a limited scale for capti"e floc+s but is not feasible on a scale that would affect wild populations.
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