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Home , Acanthocephala, Acanthocephalus lucii, Andracantha, Archiacanthocephala, Arhythmorhynchus, Filicollis

Key to Reported in Waterfowl

,s . 9]-4. .~ UNITED STATES DEPARTMENT OF THE INTERIOR . A3 and Wildlife Service I Resource Publication 173 no . 173 Resource Publication

This publication of the Fish and Wildlife ervice is on of a eri e of emitechni cal or instructional materials dealing with investigations related to wildlife and fish. Each i published as a eparate paper. The Service distributes a limited number of these reports fo r the u e ofF deral and tate agencies and cooperators. A list of recent i ues appears on inside back cover.

U.S. FISH & vJ L .t..Jlf'E SE VICE Natleaal Wetl ~ n." s " eaearcb Coat. NASA • Slieoll Co ..p .. ter Coaploa 1010 Gauae Beule•af"d s&Well, LA 70411

opies of this publication may b obtained from the Publication nit, U. . Fish and Wildlife ervice, 1atomic Building, Room 14 , Wa hington, DC 20240, or may be purchased from the ational T chnical Information Service ( TIS), 52 5 Port Royal Road, Springfield , VA 22161.

Library of Congress Cataloging-in-Publication Data

McDonald, Malcolm Edwin, 1915- Key to Acanthocephala reported in waterfowl.

(Resource publication I nited State Department of the Interior, Fish and Wildlife Service ; 173) Bibliography: p. upt. of Docs. no.: I 49.66:173 1. Acanthocephala-Identification. 2. Waterfowl­ Parasites-Identification. 3. - Parasited-Identification. I. Title. II. Series: Resource publication (U.S. Fish and Wildlife ervice) ; 173. 914.A3 no. 173 333.95'4'0973 8-600312 [QL39l.A2) [639. 9'7841) Key to Acanthocephala Reported in Waterfowl

By Malcolm E. McDonald

l ...... H ~ ll & "ll UII I-1- ~ 1-H\ ' IC t-

T OF THE INTERIOR E Resourc Publication 173 Wa hington, D. . • 1988

Contents

Pag

Introduction 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Checklist of Acanthocephala 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 • 0 0 0 0 3

Key to Families and Genera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 • 0 0 0 0 • 0 0 0 0 0 0 • 0 0

Guide to Identification of 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9

Keys to Species 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 • 0 • 0 • 0 0 0 0 0 0 0 0 0 0 0 0 • 0 • 0 0 0 0 • 0 0

References 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0. 0 0 0 0 44

Ill

Key to Acanthocephala Reported m Waterfowl

b

Malcolm E. McDonald 1

U.S. Fish and Wildlif e e1--vice ational Wildlife Health Re earch Cente?· 6006 ch?-o der Road Madison, Wi consin 53711

Introduction even spherical. The probo ci ma b w II n in a narrow or wide band; expansion may produce a p ar­ This is the third part of a continuing series on shaped (pyriform) structur , of which th narrow helminths reported in waterfowl (McDonald 1974, end may be point d or rounded, hort or long ( 1981). Coots and moorhens (in Family Rallidae, Acanthocephalan Anatomy). An important f ature Order Gruiformes) are incl ud ed with the Anatidae of the keys is the numb r of rows of hook on the of Anseriformes. The goal of these studies i com­ and the numb r of hook in each row. A plete coverage of waterfowl helminths of the world, coll ections have increa ed in th numb r of p ci ­ although the original incentive-inadequate knowl­ mens examin ed, they have also increas d in th edge of the parasites of orth American water­ amount of variation record d, o that th rang in fowl-is less true now. World coverage is desirable the number of hook ha increa d. Th hooks ar because the world distribution of the family, tribes, always numbered from the anterior nd· b cause th and even many species of waterfowl often results arrangement of row generally shows no differenc , in world distribution of parasites. it is rarely indicated. Length of a hook i indicat d The format of this key follows that of the other from the anterior tip to the posterior nd; th ho k in the series: a checklist of species, a general key is external on the proboscis, th root i int rna! and to families and genera, a guide to identification of is giv en s parat ly. pecies, and keys to species of each that has Eggs, which ar routin ly used to identify speci more than one species reported in waterfowl. Thi in the k ys, are r ally developing . Thr J group is the smallest of the series, with 52 pecie , membranes, referred to a 'shell s," ar pr sent· of which 11 may be considered accidental, normally the middle shell in egg of Polymorphu i thickest, belonging in birds of other order (two also normally ha an elongate hape, and how extension at each mature onl y in marine -although recorded pole of the embryo. o taxonomic significance is in many birds-and one in freshwater rodents). One pre ently known for the fibrils on the outer surface species is reported only from domestic waterfowl of eggs that were recently recorded in everal and one on ly experimentally from ­ species; hypothetically, they rmght serve to entangle lings, whereas ix other specie have been reported the eggs among the vegetation, where they are more from both wild and domestic waterfowl. The fre­ likely to be ingested. quency of occurrence and status of hosts in the Intermediate hosts of Acanthocephala have been checkli st have been based on the literature. All id entified as crustacea of only a few orders: Amphi­ Acantho ephala in waterfowl are found in the small poda, , and Decapoda; they are almost totally intestine (usually in the po terior portion) and the aquatic, intermediate in size, and feed on dead large intestine. and r mains. everal use fish as transport Probosces usually are ovoid or sometime ellip­ or paratenic hosts; one is known to u e nakes and tical. They may be elongate-narrow or wide, or frogs as transport hosts to birds. · Many species (perhaps a majority) are distin­ 1 Retired; presenl addre : 1017 Magnoli a Lane, 1adi on. \~ i s. gujshed by a yell ow or orange color and may readily 5371 3. be een with the unaided eye as orange bodies in 1 the intermediate ; however, at least one com­ pronounced pathological ffect on their intestinal mon acanthocephalan of waterfowl in Eurasia (Fili­ wall .) Few of the previously cited paper· included coUis anati in Asellus communis) appear as a whit detailed studies of the dead birds, and the causes of body. everal species cause a distinct change in th death were u ually assumed. In a study of parasite behavior of the intermediate host which render it numbers in in Scotland in which Acantho­ more vulnerable to predation by the final host cephala regularly occurred, Thompson (19 5) com­ (Bethel and Holmes 1973, 1974, 1977; Holmes and mented, "In this study there is no evidence to show Bethel 1972). cycles generally require at least that P. botulus increa ed the mortality rate of 2 months for completion. The intermediate ho t eiders." AI though he collected dead eiders regular­ species are preferred foods of many juvenile and ly, no determination of the cause of death wa adult waterfowl. attempted. An experimental study of antagoni tic Van Cleave (1918) believed it was ignificant that reactions between clas e of helminths during multi­ only one species of Acanthocephala had ever been ple infections (Petrov and Egizbaeva 1972) showed found in an individual host, and there were no significant reduction in the ize and number of records of the occurrence of more than one genu hymenolepidids when pr sent with polymorphid in in any host species. This is no longer true. Coryno­ the small intestine. soma and , for example, repeatedly As with other groups of helminth of waterfowl occur in the same , and several species of Poly­ the recent major references to Acanthocephala morphu have been recorded for the same host (at originated in Rus ia: Petrochenko (195 , 1971b least three are recorded in the same individual). [English translation]) and Petrochenko and A few members of Acanthocephala cause consider­ Kotel'nikov (1962). Then a erie of tudie by able damage and even death in waterfowl. Two Khokhlova (1966a, 1966b, 1971, and 1977) species in particular, Polymorphus minutus and terminated in a partial update of Petrochenko anatis, have repeatedly been cited in (Khokhlova 1978), actuall y using the data of Schmidt Europe and Asia, in domestic waterfowl as well a (cited below). Studies of the and life hi torie in wild birds (Petrochenko 1958, 197lb; McDonald of these forms up to that time are li sted in McDonald 1969b; Macdonald eta!. 1978); The pathogenic ef­ (1969a, 1969b). The of portions of the fects were described by Petrochenko and others group has been revised by Schmidt (1972, 1973, (McDonald 1969b). Polymorphus minutus is the only 1975) and Schmidt and Kuntz (1967); Amin (19 2) species that definitely occurs in North America. recently reviewed the entire . Denny (1969) Filicollis species of Eurasia (reported also in and Podesta and Holmes (1970) discus ed the cranes in Alaska, perhaps because of the number of biology of Acanthocephala and other helminth cranes that cross the Bering trait between the two carried by the intermediate host , Gammaridae, in continents) are particularly noteworthy. The probos­ Canada. Many papers on the same topic for Europe cis of the female perforates the intestine wall and and Asia are included in the literature (McDonald enlarges to form a pea-sized bulb lying against the 1969a). Kontrimavichus and Atrashkevich (1982) outside of the intestine in the body cavity; the body described parasitic systems and their po ition in of the remains within the intestinal lumen. the study of population biology of helminths. The bulb ultimately becomes embedded in connec­ Bullock (1969) gave an excellent account of th tive tissue. Excellent accounts are provided in techniques of preservation and mounting on Petrochenko (195 , 197lb) and Petrochenko and microscope slides. Koteh'likov (1962). I could detect no distinct differences between Eiders are the only wild birds in which repeated Coryno oma sudsuche Belopol' kaya 1959 and Cory­ outbreaks of djsease and death from Acanthocephala nosoma mandarinca Oshmarin 1963, both reported have been reported (Grenquist 1951, 1970; Thorn from ducks of Primorsk, and I finall y eliminated the and Garden 1955; Clark et al. 1958; Garden et a!. second as a synonym. The same thing wa true for 1964), but dead individual birds of other species have iheringi Machado Filho 1961a, Gory­ b n r corded, and at least one lake and marsh area no oma longilemniscatu Machado Filho 1961b, and in the United States routinely shows heavy infec­ Corynosoma moififernandesi Machado Filho 1962a, tions with some lo ses. (I observed from 1,500 to all reduced to synonyms of Coryno oma enri ttii 10,000 Acanthocephala in dead swans, and noted Molfi and Fernandes 1953, of Brazil. Polymorphus magnus Skrjabin 1913 has been The form listed as Polymo1-phus trochus of merged with P. minutu Goeze 1782, following the Khokhlova is distinct from P. t1·ochus Van leave lead of Bezubik 1957a. Bezubik howed that speci­ 1945. P. trochus Van Cleave i almost universal men of P. minutus described by Petrochenko were in Fulica americana, and rarely occurs in duck actually P. contortus; the true P. minutus then in­ in orth America, except occa ionally wh n they clud d P. magnus. 2-12 mm long showed are severely ill (twice in mallards). The form of only one peak of variation in length for each sex, in­ the proboscis is strikingly different in the two dicating only one species. Females 4-7 mm long mor exes. P. trochu of Khokhlova ha b n re­ frequently pos essed mature eggs than lacked them ported repeatedly in wild ducks in Asia, but (precisely t he range that Petrochenko said lacked has not been recorded in th Europ an coot. Th them)-too large fo r P. minutus (actually P. conlor­ form of the probo ci show li ttle difference tu ) and too young for P. magnus. Only researchers between males and female . I have refrain d in England (Crompton and Harrison 1965) have fol­ from renaming thi s form, but a new nam lowed Bezubik's lead; P. magnus has been recorded neces ary. repeatedly since 1957, 34 times in wild ducks and Common nam s of ho ts (Tab! 1) u ed in thi 40 times in domestic duck , mostly in . publication follow thos of cott (1977).

Table 1. o.f hosts.

Common na me

American coot Fulica ameTicana (order Gruiformes) Baha ma pinta il Anas bahomen ·is (order Gaviiform Bay duck Tribe Aythyini Mall a r A nas platyThyncho. Brazilian teal A nw zonetla b1·asi li nsis Manda rin duck A ix gal 1·iculata Brown dipper Cinclus pallasii ommon moorh n Gallinula chloTopus (order Passeriforme ") (ord r Gruiform ) oot Ge nu Fu.lica Mu k du k Bizium Lobato (order Gruiforme ) orthern pin tail Anos ct uta Dome Lie duck Anas platy1·hynchos (dom.) ommon pocha rd A ythya j'e?·ina Dome t ic muscovy Cairina mo chala (d om.) Puddle duck Trib Anatini Eide r Tribe omateriini Rosybill llo peposoco ulc~ bassanus a du k Gen ra 'Vfelonill o. fl isll·iunitu:;. (order P I caniforme ) langulo ommon goldeneye Bucephala clangulo wan Ge nus Cygnus Greater caup A ythy l mar-ila Tuft d duck Aythyo fuligula Ha rl equin duck His/1·ionicus histrionicus Waterfowl Fa mil y Anatid ae Hooded me rgan r M ergn · cucullo Ius Checklist of Acanthocephala Reported m Waterfowl

Frequency of Statu of ho t pecies records (wi ld = W)

Class

Family Oligacanthorthynchidae N eoncicola avicula (Travassos 1917) Rar w

ORDER PALAEACA THOCEPHALA

Family anguillae (Mull er 1789) Accidental w (Muller 1776) Accidental w Acanthocephalus ranae (S chrank 17 Accidental w

Family Centrorhynchus aluconis (Muller 1780) A cidental w Centrorhynchus teres (Westrumb 1 21) Acc id ental w

Family mergi (Lundstrom 1942) Infrequent w Andracantha phalacrocoraci (Yamaguti 1939) Accid ental w frassoni (Molin 1858) Accidental w Arhythmorhynchus invaginabili (Lin tow 1902) Accidental w Arhythmorhynchu tere Van Cleave 1929 Accidental w Corynosoma anatarium Van Cleave 1945 Rare w Corynosoma con ·trictum Van Cleave 191 haracteri stic w oryno. oma ~nrie ttii Molfi a nd F ernand e 1953 Infreq u nt W, Domesti c oryno. oma peposacae (Porta 1914) Rare w oryno. ·oma . · me1·m (For sell 1904) Infrequent w or yno.·oma ·trumosum (Rudolphi 1 02) Infrequent w CO'Iynosoma sudsuche B I pol' kaya 1959 Rare w orynosoma tunitae Weiss 1914 Rare w Filicollis mati · (S chrank 17 ) Characteri stic W, Domestic Pilicollis trophimenkoi Atrashk evich 1982 Infreq uent w Hexaglandula paucihamatu · (Heinze 1936) Ac idental w Polymot·phw,; aclugan nsis Petrochenk o 1949 Frequent w Polymorphns acutis Van leave and Starrett 1940 Infrequent w Polymo1-phus biziurae J ohn ston and Edmond 1948 Rare w Polymorphu. cincli B I p I' kaya 1959 Experim ental Domestic Polymo1·phus cant rlu-> (Bremser 1821) om mon w Polynwrphu · 01-ynoid · krjabin 1913 Infrequent w Polymorphus cucullatus Van leave and Starrett 1940 Rare w Polymorph~~· diploinjZatu Lundstrom 1942 Frequent W, Domestic P lymor-phus ko tylewi Petrochenk o 1949 Infrequent W, Dome tic P lyrnorphus mariti· Van I av 1939 Frequent w Frequency of tatu of ho t Species record (wild = W)

Polym01·phus mathevo sianae Petrochenko 1949 Frequent v Polymorphu meyeri Lundstrom 1942 Rare w Polymorphu miniatus (Linstow 1896) Rar w Polymorphus minutu (Goeze 1782) Characteristic W, D me tic Polymo1·phus obtusus Van Cleave 191 Infrequent w Polymorphus paradoxus Connell and Corner 1957 Infrequen w Polymorphus phippsi Kostylev 1922 Frequent w Polym01·phus pupa (Linstow 1905) Frequent Polym01·phu triatus (Goeze 17 2) Frequent 'W, Domesti Polymorphus strumosoide Lundstrom 1942 Frequent w Polymorphus swartzi Schmidt 1965 Rare w Polymorphus trochus Van Cleave 1945 ommon W, coot Polym'Jrphu (trochu of Khokhlova 1966) Frequent w altmani (Perry 1942) Rare w Profilicolli anticus (Van Cleave 1920) Infrequent w Profilicolli botulus (Van Cleave 1916) Frequ ent w Profilicolli forma us (Schmidt and Kuntz 1967) Rare Dome ti Profilicollis major (Lund trom 1942) Infrequent w

Family cylindraceus (Schrank 17 Accidental w Plagiorhynchus gracilis Petrochenko 195 Infrequent w Acanthocephalan Anatomy

''-_____ Neck

~~~---- Cerebral ganglion ~...,._. ____ Proboscis receptacle

'~~~--- Lemnisci ""'""""--Trunk ---f;~~--- Testis

r---- Constriction

Ovum with Embryo (Polymorphus)

!~-+------Middle membrane (shell) (Fertilization membrane)

'<'<~+------Inner membrane (shell) -+++t------Acanthor t------Outer membrane (shell) Variations of Proboscis

Cylindrical ovoid E longate ovoid

Pyriform Spherical Key to Families and Genera (Acanthocephala Reported in Waterfowl)

1. Body (trunk) without spine ...... 2 Trunk with at least a few spines usually on anterior portion of body .... (F. Polymorphidae)-5 2. Male with less than eight cement g lands; probosci h ath sac-like, with two muscula r layers ...... 3 Male with eight cement glands, not definitely paired; probo cis sheath a single-wall d ac, not stout; probo cis hooks in 5-6 transver e circle , very powerful, roots in first (apical) r w of hooks bifur- cated; male 12 mm long, female 400 mm long; gg oval; accid ntal in waterfowl ...... (F. )- eoncicola 3. Mal s with ix c ment glands, of hig hly variabi fo rm (Fig. 1.1); sometime with lateral or ant - riorl y dir cted proce es from main root of proboscis hook (Fig. 1.2); accidental in waterfowl, normally in fish or ...... (F. Echinorhynchidae)-Acanlhoc phalus Males with three very long narrow cement g la nd ...... 4 4. Probosci h ath attach d near middle of probo ci wall; probosci po terior to attachm nt bear­ ing simple thornlike pines, anteriad it bear trong hook with recurved root (Fig. 1.3); eggs oval ...... (F. C ntrorhynchidae)-Centrorhynchu Proboscis sheath attach d at base of cylindrical or club- hap d pr bo cis; probo cis hook uniform in shape (Fig. 1.4); eggs oval ...... (F. Plagiorhynchidae)-Plagio?·hynchu Anterior trunk pin in two horizontal zone , ·eparated by small gap (Fi ·. 1. 5) ... A ndmcantha Anterior trunk pine in on -zone ...... 6 6. pines not limited to anterior region of t runk, som (genital) occurring at posterior end; anterior trunk pines often extending more posteriad on ventral surface (Fig. 1. 6) ...... Andracanlha pin limit d to anterior, none at po terior end ...... 7 7. Hook on midv ntral urfac of probo cis often distinctly la rger a nd heavier Lhan other hook at corre pe nding level on dor al surface; prob ci u ually enlarged (swollen) near center , spindle­ haped (Fig . 1.7) ...... orynosoma Hook of dorsal and ventral sides of probo cis imilar in ize (Fig. 1. ) .... A1·hythmorhynchus Body mark dly xually dimorphic; probosci off mal spherical, with reduced hooks in tar- haped radiating rows at tip (Fig. 1.10; actual! on probosci portion of sphere formed from inflated proboscis and ant rior portion of neck), pher u ually lying outside inte tina] wall due t per­ foration, with body of worm till in side inte tin ; proboscis of male nearly orbicular, with 1 longitudinal row of 10- 11 hook ach, 21 - 31 1-1m long, anterior with roots; 6 compact r ni fo rm · m nt glands ...... Fi~icollis Pr bosci. spherical to cy lindrical, imilar in both . exe , row of hooks extend ent ire length of probo ci , probo ci entir ly within inte tine of host ...... 9 ~. I· our · m nt gland in male, elongate, narrow, lying parallel in on bundl ; proboscis spherical to cy lindrical , con tri tion in body u ually evident ...... 10 Six ·ement g lands in mal , int tine- hap d, in two groups of three each (Fig. 1.11); probosci ovoid , with I v n longitudinal row of hook ; constriction on trunk slight (females fall into Pr /ymorphus key) ...... Hexaglandula 10. 8gg val; prob scis pherical, rarely slightly ovoid (Fig. 1.12) ...... Profilicollis Eggs elongate, middle she ll thi ck, extend d and narrowed at poles; probo cis ovoid , pyriform, or cylindrical (Fig . 1. 9) ...... Polymorphus Guide to Identification of Species (Acanthocephala Reported in Waterfowl)

ote: Includes all genera reported in waterfowl in recent literature.

Genus Species reported in waterfowl

A canthocephalus 3 species in waterfowl, of perhaps 35 in genus; all accid A ndracantha 2 species in waterfowl, of 3 in genus. See key to pecie . A rhythmorhynchus 3 species, of perhaps 25 in genus. See key to specie . Centrorhynchus 2 species, of perhaps 60 in genus. See key to specie . Corynosoma 8 species, of perhaps 30 in genus. ee key to species. Falsifilicollis Synonym of Profilicollis. Filicollis 2 species, all in genus. See key to species. H exaglandula H. paucihamatus, of 4 species in genus. Perhap accidental, type host a gull. G nu perhaps belongs in Polymo1·phus; see Petrochenko 195 or 1971b for de cription . Neoncicola . avicula, of 2 in genus; rare; South America; see Petroch nko 195 or 1971b for description, under Prosthenonhi . Parafilicollis Synonym of Profilicollis. Polymorphus 24 species, of about 26 in genus. ee key to specie . Profilicolli 5 species, of in genus. See key to pecie . Plagiorhynchus 2 species, of about 22 in genus. See key to species. Fig. 1. Acanthocephalan Genera

1.2 Proboscis hook with root and lateral processes

Cement glands .l.------:

1.1 Acanthocephalus

Receptacle attachment

1.3 Centrorhynchus Probo cis receptacle (one layer) /Z-.f(--{j_~--a--r~-a-a..... ' t. .:.:: ...... · · · .) ·········· ··········

··· ·· ···· · ..

1.4 Plagiorhynchu 1.5 Po t rior nd with hook

1.6 Andracantha 1.7 Coryno oma

1.9 Polymorphu Ovum

1.8 A rhythmorhynchu 1.10 Filico lli proboscis

Keys to Species

Genus A canthocephalu

ote: Infection obtained by inge tion of freshwater isopod ru tacea.

1. Probo cis hooks with two lateral proce es at top of roots; hooks in 10 longitudinal rows. with 6- 7 hoo k in each row; probo ci club- haped, wid th about 1/3 of length (Fi ·. 2.1 ); accid ntal, normall y in fi sh; orth America, Eura ia ...... A. anguilla Probosci hooks without lateral processes on roots ...... 2

2. Female 8-17 mm long, male 5-12 mm long cylindrical; probo cis hooks in 12-16 longitudinal r ws of 7-9 hooks each; proboscis 500-700 ~-tm long, 250-300 ~-t m wide; lemnisci about 1,600 1-1111 long (Fig. 2.2); accidental, normally in fi h; Europe ...... A. lucii Female 20-60 mm long, male 5-12 mm long, cylindrical; proboscis hook mo tly in 16 (12-20) longitudinal rows, with 4-6 hooks in each row; probo ci 450-500 ~-tm long, 300 ~-tm wide; I m­ ni sci 600-800 ~-tm long (Fig. 2.3); accid ental normall y in Amphibia; orth Ameri ca, Europ ...... A . rana

Descriptions: Petrochenko 1956, 1971a.

Genus Andracantha

ote: Sourc of infection unknown; probably marine.

1. Posterior field of trunk spine continues down ventral side nearly to posterior tip; f mal 4.6-6 mm long, male 4-4.25 mm long; probosci hook in 16- 19 longitudinal row with 10- 14 in ach row, hooks increase in size to 9th hook (w ith roots), Ia t 5 without root ; 9th hook 66 J..l m long in male, 105 J.J m long in female; cement glands, arrang d irregula rl y; male without genital pin , oft n ab ental o in female; eggs 8 -96 J.Jm by 42-50 J.Jm , oval (Fig. 3. 1); inf requ nt, in fi h-eating birds; orth Ameri ca, Asia ...... A. phalac?·ocoTacis Po terior field of trunk pines does not continue down ventral sid e; male 3-4 mm long, f mal e 4.7- 4.9 mm long; probosci with 15-17longitudinal rows of hooks, with 9- 10 in each row fir t 7 hooks increase in siz to 60 J.Jm long in male, 70 J.Jm long in femal , with root re t without roots; often without g ni ta! pines; 6 cement glands, in pairs; egg 44-50 J.Jm by 16- 24 J.Jm , oval (Fig. 3.2); infrequent, in fi h- ating bi rd ; orth Am rica, Eurasia ...... A. rn 1·gi

Description : Schmidt 1975.

Fig. 3. Genus Andracantha

Ovum

3.1 A. phalacrocoracis 3.2 A . mergi Genus A rhythmorhynchu

ote: One species obtained by inge tion of marine decapod Cru tacea; fi h erve as transport ho ts.

1. Hooks of midventral side of proboscis 2-3 time larger than the rest; anterior body ovate, 2.3-2.6 mm long, eparated by constriction from remaining 617 of body, po terior end widened; neck very short· proboscis with 18 longitudinal rows of 20 hooks each; body 30- 0 mm long (Fig. 4.1); accidental in mergan er , normall y in other aquatic bird ; Eurasia, Africa, Australia ...... A. frassoni Hook of midventral side of proboscis only sl ightly enlarged ...... 2

2. Proboscis with 22- 24 longitudinal rows of 1 -19 hooks each; neck long; hook at least 45-51 Jim long; lemnisci long, reach beyond armed trunk; body -35 mm long (Fig. 4.2); accidental in mergan er , normally in shorebirds; Europe, orth America ... . ~ ...... A. invaginabilis Probo ci with 18 longitudinal row of 14-16 hook each, h ks u ually 60 Jim long; accidental in merganser , normally in shorebird and gulls; Eura ia, outh America ...... A. ter s

Descriptions: Petrochenko 1958, 1971b. Fig. 4. Genus Arhythmorhynchus

f/1p

4.1 A . fras oni Rt (:::' (:) ?{] P' Proboscis hooks ff1 (one row) ~ ~ ?t:l ?:D p> ~ ~ 4.2 A. invaginabilis Genus Centrorhynchu

ote: ource of infections unknown; genus mostly in terrestrial birds. orynosoma aluconi.· in ·nakes and as transport hosts.

1. Body 6-45 mm long (male 6-25 mm , female 11-45 mm); proboscis 1,000-1,500 11m long, with 30-32 longitudinal row of 15 (16-1 reported) hooks each; anterior 6- 7 hooks with well-developed roots (hooks and roots both 55 11m long), po terior hook pinelike without root (Fig. 5. 1); accidental in mergansers, normally in birds of prey; Eurasia ...... C. aluconi · Body 14-20 mm long; proboscis 650-700 11m by 400 11 m , cylindrical or anteri or portion nearly orbicular, narrowed toward base, with 32-34 longitudinal rows of 11-12 hook each, anterior 5-6 hooks large (36-38 11m long, roots 4 -5 11m long), posterior 5-6 hooks spineli ke, numbers 6-10 with anterior process of root (Fig. 5.2); accidental in pochards, normall y in corvid pa serine birds; Europe ...... C. tere.

Description : Petroch nko 195 , 197lb.

Fig. 5. Genus Centrorhynchus

~ ~ ~ (} (jV ~ ~ ~ ~ ~ ~ ;: Proboscis hooks (one row)

5.1 C. aluconis Proboscis hooks

4

5

~~ 6 ~ 9 ~ 10 5.2 C. teres Genu Coryno oma

ote: Infection obtained by inge tion of fre hwater (on ·peci s) or marin (two p cie ) · :~mphipod and isopod ru tacea, or by inge tion of marine fi h rving a transport ho ts (two p cie ).

1. Triangular hape, showing spine facing 1 mrn, on !ant ...... 2 Anterior end !anted to ventral ide, to middle or be ond ...... 3

2. Triangular hape, front 1 mm wid ; body 5 rnm long; pine in ant ri or part not xt nding a far a middle area; genital spine around genital por at po terior nd; with 1 longitudinal row of 10 hooks each (Fig. 6.1 ); orth America, orth frica; in eid r (a nd n rth rn gannet) ...... lunitae Forebody swollen, posterior taper d ...... 4

3. Body 5-7 rnm long, occasionall y 9 rnrn; anteri or part inflated, about 1/4-1/3 of ntir trunk, ·pine extend on ventral id e to middle area; genital ·p ine around po terior tip in mal ; pr bo ci with 1 longitudinal row of 10-11 hook each; large t hook number 6 and 7, or number 7 and , 64- 0 1-1rn long, root longer than hook; 1 mnisci broad and fl at. hotter than proboscis sheath; te tes oval, more r le ·s opposite· cement gland 6, pyriform. two g roup f three, longate; eggs 79-101 !-lrn long by 19 ~J ill wide (Fig. 6.2); accidental, norma ll y in marine ·, fre­ quent in marine fish- ating bird but rarely mature; coasts of orth Am ri ca, northern Europe ...... ·t?-umo.·um Trunk spine extend ventrall y to posterior end, mixing with genital pines; body small , 3 rnm long; anterior part enlarged, inflated, probo cis cylindrical with light swelling at 2/3 of length, with 22- 24 longitudinal row of hook of 12-13 hook each; large t hooks number 7- , with recurved roots, about 67 1-1m long, nding with 4-5 hooks without root·; t tes round, on b hind the other; lemnisci slightly shorter than probosci sheath; egg 79-101!-lm long by 16-29 ~J ill wide (Fig. 6.3); a ·cidental, normally in inarine mammal (Pinnipedia), common in fi h-eating birds, but rarely mature ...... ·em.erme

4. Forebody swoll en, po t rior taper d; spines spar e around po terior nd of body; body 3.9-5.2 rnm long; for body covered with hook ; probo ci about - 50 ~Jill long, with 1 longitudinal row of 12 hook each; largest hook in rniddl of probo cis, number 7, 64 1-1 m long; hooks toward anteri or end are larger, become smaller toward rear; testes oval, id e by si de; lern ni ci shorter than proboscis sheath; 6 cement glands in two group of three; eggs 6 ~J ill long by 25 ~Jill wide (Fig. 6.4); in manda rin and harlequin duck ; Asia (Primor ·k) ...... sud ·uche

o. Body wo ll en at forebody; con t ri cted below fo reb dy ...... 6 F orebody not swoll en, body not constri cted ...... 7

6. F orebody wollen , strong constri ction below forebody; body 2.2 -6.0 mrn long; proboscis cylin­ drical, woll en below middle, largest hooks at about swoll n area, 41-47 1-1m long, other hooks 30-41 1-1rn long at tip and 35-41 1-1rn long at base of probosci ·; te te oval, overlapping about 1/3; egg 0-10 1-1m long by 12-16 1-1m wide (Fig. 6.5); common in duck ; orth America ...... C. con ·trictum Body elongate, broadening anteriorly, light constriction; body 8-11 rnm long; body club-shaped, widened fo rward, anteri or part with strong co ni cal spines; males with spines present sparingly at posterior end; proboscis 340-430 ~Jill long, with 14-1 longitudinal rows of 12-14 hooks each; large t hooks number 4; lemnisci cylindrical, about twice as long as proboscis sheath; eggs 64-6 1-1rn long by 17- 1 ~Jill wide; rare, in rosybill s; South America (Brazil, Argentina) ...... C. pepo ·acae 7. Body relatively small 4.2- .6 mm long; with small number of uticular pin irr gular!) cat­ tered in both sexes at geni tal pores; proboscis about 550 Jim long and 2 0-290 Ji m in diam t r, armed with 14 longitudinal rows of 8 or 9 hook each· large t hook about J.lm Ion ·about at the middle of the proboscis, hook near the ba e about 47-59 J.l m long, tho e near th tip 59- 2 J.lm long; egg 100- ll2 J.lm long b 20-23 J.lm wide (Fig. 6.6); in duck ; orth m ri a (U ..) ...... C. anatm·ium Body fairly small , middle ection large, one worm with for body bent over at ant ri or; probo i with 20 longitudinal row , with hooks in rows each; hook in thre b It , wo ll en b low he middle; largest hooks numbers 4 and 5, 46 Jim long (root 56 Jim long); I mnisci long, inuou , about 3 time the length of proboscis sheath; te te oval, about 1/3 overlappin ·; g · 9 - 100 Jim long by 19-21 J.lm wid e (Fig. 6.7)· rare, in dome tic muscovys, Bahama pin tails, and Brazilian teals; South America (Brazil) ...... n riel/ i i

De cription : Petrochenko 195 1971b; Belopol' kaya 1959; Machado Filho 196la, 196lb, 1 chmidl 1965b· Oshmarin 1963; Bourgeoi and Threlfall 19 2.

Fig. 6. Genus Corynosoma

6.1 C. tunitae

6.2 C. strumosum 6.3 C. semerme

6.4 C. udsuche

6.5 C. constrictum 6.6 C. anatariurn

6. 7 . enriettii Genus Filicollis

Family Polymorphidae

ote: Infections obtained by ingestion of freshwater isopod Crustacea. Salamanders serve experimen­ tally as transport hosts in one species (F. trophimenkoi).

1. Proboscis hooks in male up to 41 J.lm long, reduced in female; body 7.6- 12 mm long in male, 24.9-36.4 mm long in female, long, fusiform in female; eggs 58-71. 8 J.lm long by 17-22.2 J.lm wide, oval , without projection of middle shell ; mature fe male cylindrical, band-shaped, with bul b at anterior end , outside intestine (Fig. 7 .1); characteristic, in all waterfowl, typically in northern pintails and other puddle ducks; Eurasia ...... F. anatis Proboscis larger, hooks up to 4 Ji m long in male, reduced in female; body 6-7.4 mm long in male, 14.3-21 mm long in female (reported 2/3 as large as F. anatis); eggs 84-118 Jim by 25.6-39.3 Jim , with surface covered by fine fib ril s (reported 1.5 times larger than F. anatis), middle shell weakly projecting at poles; matur female flat-ovate, tablet-shaped with bulb on ventral side posterior of anterior end (bulb 2.275-3.212 mm diameter; Fig. 7.2); infrequent, in long-tailed ducks, greater scaups, and ; Asia (Chukotka) ...... F . trophimenkoi

Descriptions: Petrochenko 195 , 197lb; Atrashkevich 19 2. Fig. 7. Genus Filicollis

~ 0) ~ fD cJ ~ ~ ~P:J

Female in situ in intestine of host

7.1 F. anatis 7.2 F. trophimenkoi Genus Polymorphus

Family Polymorphidae

ote: Infections obtained by ingestion of freshwater (seven species) or marine (one species) amphipod Crustacea. One species also uses fish as transport hosts.

1. Proboscis markedly sexually dimorphic, strongly inflated pear-shaped in female, ovoid without inflation in male; in female posterior of proboscis 380-560 lim long, 310-490 lim in diameter, anterior end nipplelike, 100-175 lim in diameter; in male proboscis 385-560 lim long, 210 lim in diameter, elliptic; with 20 longitudinal rows of hooks, alternately 11 and 12 hooks each, larg st 46-53 lim long (numbers 3 and 4; Fig. 8.1); infrequent or accidental in puddle ducks, common and normal in American coots; orth America ...... P. trochus Van Cleave Proboscis not sexually dimorphic, same in both sexes ...... 2

2. Eggs wide-oval, inner shell without elongate polar prolongations...... 3 Eggs elongate, middle shell particularly spindle-shaped, with polar prolongations often enlarged at tips (not mentioned further) ...... 5

3. Proboscis cylindrical, slightly swollen in rear 1/4; proboscis hook in 21-22 longitudinal rows of hooks, with 9-11 hooks in each, first 5 hooks large, with roots, largest about 90 lim long, re­ mainder small; body 6.2-18 mm long (shape of eggs indicates species of genus Profilicollis but shape of proboscis is not indicative; Fig. 8.2); rare, in musk ducks; Australia ...... P. biziume Proboscis wide-ovoid, almost spherical in appearance ...... 4

4. Eggs oblong, 66-80 lim long by 19-231im wide, with very slight, hort polar projections of middle shell; proboscis nearly spherical, with 14-16longitudinal rows of hooks, 7-10 hooks in ach row, hooks 66-73 1im long, with roots 66-791im long; testes long-oval, tandem but slightly overlapping; body 8-16 mm long, cylindrical (Fig. 8.3; eggs do not quite fit any genus); infrequent, in hooded mergansers, ducks, and loons; orth America, Asia ...... P. obtu us Eggs elongate, 124-1481im long by 34-40 lim wide, with short wide prolongations of pole of mid­ dle shell; proboscis with 18-20 longitudinal rows of hooks, 7- hooks in each row; largest hooks 93-1081im long (numbers 4 and 5), roots 108-130 lim long; testes oval, tandem; body cylindrical, 13-25 mm long (Fig. 8.4; eggs not quite matching others of genus); infrequent, in eid ers; Arctic Asia ...... P. pupa

5. Proboscis with 6 hooks in each row ...... 6 Proboscis with more than 6 hooks in each row ...... 8

6. Body 8- .5 mm long, thick in middle, constriction at anterior and posterior 1/5; mall pines on trunk; proboscis with 12 longitudinal rows of hooks, with 6 hooks in each row, size unknown; lemnisci club-shaped, 1.1-2 mm long; testes long-oval, tandem, but overlapping 1/2; cement gland in two groups of three each, groups tandem but slightly overlapping, intestiniform; eggs 94-96 lim long by 22- 26 lim wide (Fig. 8.5); accidental, normally in gulls; Europe ( R) ...... Hexaglandula paucihamatus Proboscis with 16 or 17 longitudinal rows of hooks...... 7

7. Proboscis ovoid, 470-700 lim long, with 16longitudinal rows of hooks; 6 hooks in each row; largest hook 71.4 lim long (number 3); forebody with spines extending more posteriad on ventral side; body 15 mm long (female); eggs 118-132 lim long by 22-25 lim wide; frequent, in eid ers; Arctic Eurasia ...... P. phippsi Probosci almost as broad as long, 300 1-1m long by 300 1-1m wide in male 400 1-1m long by 300 1-1m wide in female; with 16-17 longitudinal rows of hooks; 6 hooks in each, largest hooks 63-69 1-1m long (number 3); lemnisci club-shaped; body elongate, 7-14 mm long; eggs 96- 10 1-1m long by 18 1-1m wide; rare, in mallards and tufted ducks; Europe ...... P. meyeri

Body less than 6 mm long ...... 9 Body more than 6 mm long ...... 1

9. Each longitudinal row of proboscis hooks with 7 (to or 9) hooks ...... 10 Each longitudinal row of proboscis hooks with (to 9 or 10) hooks ...... 12 Each longitudinal row of probosci hooks with 9 (to 10) hooks ...... 13 Each longitudinal row of proboscis hook with 10 (to 11 or 12) hooks ...... 14

10. Proboscis ovoid, slightly wollen, with 20 longitudinal rows of hooks (occasionally 1 or 22 row ), with 7 or 8 hook in each row; largest hooks 40 1-1m long (number 6), root 52!-lm long; fir t 4 hook large, with long roots, remainder small; testes round to oval, opposite; body 1.83-3.8 mm long; eggs 105-112 1-1m long by 19-20 1-1m wide, surface covered by network of fine fibrils in loops (Fig. .6); frequent in duck , particularly in northern pintail ; Arctic A ia ...... P. sp. (P. t1·o hus of Khokhlova) Body hort, plump ...... 1 1

11. Body short, 4.6- 7. mm long; proboscis ovoid, with 14 longitudinal rows of hooks, 7-9 hooks in each row; largest hook 69 1-1m long (number 3); lemnisci about 11/a the length of probo ci sheath; te tes round, oblique to each oth r; egg 90-105 1-1m long by 18 1-1m wide, urfa cover d with fine fibril (Fig. .7); frequent, particularly in scot rs; Eurasia ...... P. diploinjlatu. Body short, plump, strongly bent over, 1-2. mm long; proboscis with 15- 1 longitudinal row , each with 6- hook , 3-4 with roots, 40-47 1-1m long; testes oval; lemnis i club-shaped, short; eggs 80-130 1-1m long by 15- 19 1-1m wide; common, in puddl duck ; orth Ameri a, Eur p ...... P. conloTtu;

12. Body 5.5-10.5 mm long; constriction rather evident, in middle of body; n k conical, twice as long as probosci ; probosci ellipso id , with 17- 1 (rarely 1 6) longitudinal rows of hook , 9 (rar ly ) hooks in each row; largest hook 63- 6 1-1m long (number 4); lemni ci about twic a. long as probo cis sheath; teste oval, opposite or oblique; egg 102- 123 1-1m long by 19-21 1-1m wid (Fig. 8.8); infrequent in ducks, normally in aquatic rod nt ; orth America ( anada) ...... P. pn rado:ws Body 2- 12.5 mm long, constriction evident; prob ci ovoid, lightly woll n; with 16 (s metim s 18, 20, or 22) longitudinal row of hook , 8-10 hooks in each row; largest hook 67-79 1-1m long (numb r 4); fir t 4 with trong hook and roots, r mainder much reduced; lemnisci long r than probo cis sheath; testes oval, obl ique to each other; egg 73-149 1-1m long by 17-23 1-1m wid (Fig. 8.9); characteri tic and common in Eura ia, infrequent in To rth America ... . P. minutu.

13. Body 3. 5-4.33 mm long, spindle-shap d, length of exe id entical; probo ci long-oval, swollen, with 12 longitudinal rows of hooks, 9-10 hooks in each row; large t hooks 53-71!-lmlong (number 5); long roots on fir t hook , last four with r duced roots; I mni ci equal length of probo ci sheath; testes round, opposite; eggs 125 1-1m long by 17. 1-1m wide (Fig. .10); infrequent, in puddle ducks and sea ducks; Asia ...... P. kostyl wi Body 2.87 mm long; proboscis cylindrical, with 12-14 longitudinal row of hook , 9-10 hooks in each row; largest hooks 64 1-1m long (number 2); lemni ci hotter than probo ci h ath; te tes round, oblique to each other; eggs 107- 124!-lmlong by 21-22 1-~m wid (Fig.. 11); experimental in domestic duckling , normally in brown dippers in tream; Asia ...... P. cincli 14. Body short, swollen at anterior, 3.5-5.5 mm long, pear-shaped· spines on forebody extend 3 tim s as far ventrally as dorsally; proboscis wide cylindric, with 16-1 longitudinal row of hooks, 9-11 hook in each row; largest hooks 40-52 J..tm long, about middle of probo cis on wollen area; lemnisci about the length of proboscis sheath; testes round, opposite; eggs 102-111 J..tm long by 1 -21 J..tm wide (Fig. 8.12); infrequent, in ea ducks; Eurasia ...... P. strumosoirles Additional forms with 10 (to 11 or 12) hooks in each row ...... 15

15. Body elongate, forebody long swollen, spiny; body 4.75 mm long (femal ); con triction below forebody at 113 of the length; proboscis cylindrical, inflat d 2/3 from anterior nd; with 18 longitudinal rows of hooks, 10-11 hooks in ach row; longest h oks 60 J..t m long (number 5-6), last 3 hooks without roots; lemni ci club-shaped, lightly longer than proboscis sheath; eggs 78-86 J..tm long by 15-16 J..tm wide (Fig. 8.13); infrequent, in common goldeneyes; orth America (Ala ka) ...... P . .·wa1 ·tzi Body 3 mm long, anterior swollen, tapered to rear; proboscis club- hap d, with 10 longitudinal rows of hooks, 10-12 hooks in each row; anterior hook with root , iz unknown; te te large, oval; eggs 88-96 J..tm long by 14. JAm wide (Fig. 8. 14); infr quent. in mallard and long-tail ed ducks; Asia ...... P. cO?ynoide::;

16. Each longitudinal row of probosci hook with 11 (and 12) alternating or total pr s nt ...... 17

17. Body 2.63-4.46 mm long; anterior wollen, no con triction, tapering to rear; with 20 longitudinal rows of hooks on probosci , 11 and 12 hook alt rnating in rows, largest hook 49 J..tm 10ng (numbers 7 or ), first 7 or hook well developed, with roots, remaining 4 without root ; teste round, opposite; lemnisci shorter than probo ci sheath; eggs 102-109 J..tm long b. 17. - 23 J..lffi wide (Fig. 8.15); frequent, in puddle ducks and ea duck·; Arctic A ia .... . P. maihevossianae Body plump toward rear, tapered anteriorly, 4-6.5 mm long; neck very long, tapered; proboscis with 16 longitudinal row of. hook , with 11 -12 hook in ach row; large t hooks 40-45 J..tm long at tip, remainder 32-42 J..tm long; lemni ci mor than twice the length of probo cis sheath; teste oval, tandem, but overlapping (Fig. .16); infr quent, in ducks; orth Am rica, A ia ...... P. ucuh·. .

1 . Body with evident con triction ...... 19 o constri ction pr ent ...... 20

19. Forebody spiny, \vith constriction before end of spiny area; body 10.3 mm long, uniform width; proboscis long-oval, with 16 (sometimes up to 18) longitudina l rows of hook·, with 7 (som tim s 6) hooks in each row; rar , in Ana· p.; Magellan Straits, outh Am ri ca ...... P. miniatus Body 10.26-12.49 mm long; body with anterior constriction at about 1/3 the length of piny ar a; probo cis long-oval, with 1 longitudinal row of hooks, with hooks in each row; large ·tho ks at anterior (number 1-4; 71 -75.6 J..tm long), cond 4 hook small r; neck long; lemnisci longer than proboscis heath; body 10.26-12.49 mm long; egg 107-120 J..tm long by 1 J..tm wide (Fig. . 1 7); frequent, in puddle duck ; Asia ...... P. actuganensi8

20. Body 6- 15 mm long, swoll n in foreb dy, tapered toward rear; probosci ·with 16longitudinal rows of hooks, with 12-16 hooks in ea h row; large t h ok in middle of row ; eggs 95-99 J..lm long by 27 J..lm wid e (Fig. .1 ); frequent, in ducks and wan ; Europe ...... P. st ,·iCLtus Body cylindrical, long ...... 21

21. Bod 10-13 mm long; proboscis ovoid, swollen below middle; with 22 longitudinal rows of hook , 12 hook. in each row; largest hook at tip, 73- 5 J..tm long; lemnisci twice as long a probosci h ath; test s long-oval, tandem, slightly overlapping (Fig. .19); rare, in hooded me rgan e rs; 'orth Ameri ·a ...... P. cucullalu::; Body 8-14 mm long, cylindrical; proboscis long-ovoid; with 16-17 longitudinal rows of hook , 7-8 hooks in each row; longest hooks in middle, 57-661im long; lemni ci twice as long a proboscis sheath; testes oval, tandem; eggs 102-120 lim long by 12-17 lim wide (Fig. .20); frequent, in aythyine ducks; orth America, Asia ...... P. ma1·ili

Descriptions: Petrochenko 1958, 1971b; Johnston and Edmonds 194 ; Bezubik 1957a 1957b; onnell and Corner 1957; Schmidt 1965a; Khokhlova 1966a, 1966b, 1971 1977; Denny 1969; Podesta and Holm 1970; Bourgeois and Threlfall 1982.

Fig. 8. Genus Polymorphus

8.1 P. trochus

0 Ovum 8.2 P. biziurae

. 7 P. diploinflatu

P. paradoxu

Ovum

8.11 P. cincli

8.12 P. strumosoides 8.1 3 P. wartzi

8.1 4 P. corynoides 8.15 P. mathevossianae

8.16 P. acutis c

8_17 P. actuganens~s

8.18 P. triatu

8 _19 p. cucullatus

Genus Profilicollis

ote: Infections obtain ed by ingestion of decapod Crustae a, one in a freshwater pecie · and thr e in marine forms.

See also Polymorphu biziurae, a species whose proboscis is o ate or cylindrical, but who ·g ar oval, without polar prolongations of the middle membrane, and who e in term diate host may b ·1 fr h­ water decapod .

1. Proboscis with over 20 longitudinal rows of hook ...... 2 Proboscis with 20 or fewer (12 to 20) longitudinal row of hooks ...... 3

2. Proboscis with 221ongitudinal rows of hoo k , 7-8 in each row; large t hooks are fir t 3, 9- 11 lim long; eggs 126- 155 lim by 30-41 JAm , elliptical (Fig. 9.1); infrequ nt, in eid ers; orth · me ri ·a ...... P. aYe! icws Proboscis with 25-30 longitudinal rows of hooks, 9- 12 in each row; larg st hooks (befor middl ) 50-70 lim long; eggs 60-70 lim by 22-30 Jim , oval (Fig. 9.2); rar , in ider ; rth Am ri ca ...... P. aU nLani

3. Proboscis with 16-20 longitudinal rows of hoo k ...... 4 Proboscis with 12- 15 longi tudinal row of 7-9 hooks each; 2nd and 3rd hooks 5-11 7 JAITI I ng ; lemni ci very long, ilat, 5-5.2 mm long; egg 62-65 lim by 23 lim (Fig. 9.3); infr qu nt, in dom ti duck ; Taiwan ...... P. formosu.s

4. Probo cis with 16-18 longitudinal row of 7- hooks in each; larg st pr bo cis hoo ks (3rd in row ) 80-96 lim long; lemni ci perhaps lanceolate, 1. 38-3.66 mm I ng; eg s 6 - 10 lim by 2 - 40 JA ITI (Fig . 9.4); frequent, in eid ers a nd oth r ea ducks; orth Ameri ca, Eura ia ...... P. botuLus Proboscis with 16- 20 longitudina l rows of 7-9 hook each; larg t probosci h ok (3 rd and 4th hook ) 105-108 lim long (root mea urement); lemni sci long, wide t in middl , 2.6- 3.2 mm long; egg 90-96 JAITI by 30-39 JA ITI , elongat d ovoid (Fig. 9.5); in frequent, in a and bay du ks; • urop ...... P. maJor

D cription : P trochenko 195 , 197lb; chmidt and Kuntz 19 "7; Kh ok hlova 1977; Bourg oi and Thr !fall ] 9 2. F tg.. 9 . Genus Profilicollis

Ovum Ovum 9.1 P. arcticu 9.2 P. altmani

9.3 P. formosus

Genus Plagiorhynchus

ote: Infections obtained in one species by ingestion of terrestrial isopod crustacea.

l. Proboscis with 16-18longitudinal rows of 11-15 hooks each, hooks 71-83 Jim long in first 9 rows, smaller toward posterior; proboscis cylindrical; eggs 48-60 Ji m by 18-29 Jim , oval; body 9.5-10 mm long, elliptical (Fig. 10.1); accidental, in probably sick mallard, normally in terrestrial passerine birds; orth America ...... P. cylindraceus Proboscis with 20 longitudinal rows of hooks, 16 in each row; proboscis cylindrical, hooks 64-71 Jim in first 9 rows, small er toward posterior; eggs 65 Jim by 26 Jim , oval; body 6-11.2 mm long, cylin ­ drical (Fig. 10.2); rare in ducks, also in common moorhens and shorebirds; eastern Europe (USSR) ...... P. gracilis

0 scriptions: Petrochenko 1958a, 1971b.

Grenquist, R. 1951. On the recent fluctuation in number References of waterfowl in the Finnish archipelago. Pages 494-496 Amin, 0. M. 1982. Acanthocephala. Pages 933-940 in in Proc. 10. Int. Congr. Ornithol. , pp ala, weden, S. P. Parker, ed. ynopsis and classification of living 1950. organisms. Vol. 1. McGraw-Hill Book Co., ew York. Grenquist, R. 1970. On mortality of the eid r duck ( oma­ Atrashkevich, G. I. 1982. Filicollis tr-ophimenkoi sp. n. te'ria molli sima) caused by acanthocephalan parasites (Acanthocephala; Polymorphidae) from Anatidae in [in Finnish]. Suom. Riista 20:24-34. north-western Chukotka [in Rus ian]. Parazitologiya Holmes, J. C., and W. M. Bethel. 1972. Modification of (Leningr.) 16:1 02-106. intermediate host behavior by parasites. Pages 123-149 Belopol'skaya, M. M. 1959 . The parasite fauna of bird in E . V. Canning and C. A. Wright, eds. Behaviow-al of the udzukhinsk Preserve (Primorsk). II. Acantho­ aspects of parasite transmi sion. Zoo!. J. Linn. Soc. 51 cephala [in Russian]. Parazitol. Sb. 18:304-320. uppl. 1, Academic Pre s, London. Bethel, W. M., and J. C. Holmes. 1973. Altered eva ive Johnston, T. H., and . J. Edmonds. 194 . Au tralian behavior and responses to light in amphipods harboring Acanthocephala. o. 7. Trans. R. Soc. . Aust. 72:69- 76. acanthocephalan cystacanths. J. Parasitol. 59:945-956. Kh okhlova, I. G. 1966a. Acanthocephala of birds of Bethel, W. M., and J. C. Holmes. 1974. Correlation of Chukotka [in Russian]. Tr. Gel'mintol. Lab. 17:245-259. development of altered evasive behavior in Gamma1-us Kh okhlova, I. G. 1966b. On the fauna and morphology of lacustTis (Amphipoda) harboring cy tacanths of Poly­ Acanthocephala of birds of the lower Yenisei and oril k morphus pamdoxus (Acanthocephala) with the infectiv­ lakes [in Rus ian]. Tr. Gel'mintol. Lab. 17:260-276. ity to the definitive host. J. Parasitol. 60:272-274. Khokhlova, I. G. 1971. Acanthocephala of birds of Yakutsk Bethel, W. M., and J . C. Hol mes. 1977. Increased vulner­ AS R [in Ru s ian]. Tr. Gel'mintol. Lab. 22:215-223 . abili ty of amphipod to predation owing to altered Khokhlova, I. G. 1977. Revision of the genus P?-ofilicollis behavior induced by larval acanthocephalans. Can. J. Meyer, 193 1 = syn. Falsifilicollis Webster, 1948 = syn. Zoo!. 55: 11 0-115. Parafilicolli Petrochenko, 1956 (Acanthocephala: Bezubik, B. 1957a. Studies on Polymorphus minutus Filicollidae) [in Russian]. Helminthologia (Brati I.) (Goeze, 17 2)-syn. Polymorphus .magnus krjabin, 15(1974):803- 10 . 1913. Acta Parasitol. Pol. 5:1-8. Khokhlova, I. G. 1978. A taxonomi c review of acantho­ Bezubik, B. 1957b. Supplement to the study on the cephalans of birds in the US R [in Russian]. Tr. helminth fauna of wi ld ducks (subfamily Anatinae). Acta Gel'mintol. Lab. 28:1 21-166. Parasitol. Pol. 5: 177- 179. Kontrimavichus, V. L. , and G. I. Atrashkevich. 1982. Bourgeois, C. E., and W. Threlfall. 19 2. Metazoan para­ Parasitic ystems and their role in the population biology sites of three species of (Anatidae). Can. J. Zoo!. of helminths [in Russian]. Parazytologiya (Leningr.) 60 :2253-2257. 16:177- 187. Bullock, W. L. 1969. Morphological features as tools and McDonald, M. E . 1969a. Annotated bibliography of hel­ as pitfalls in acanthocephalan systematics. Page 9-43 minths of waterfowl (Anatidae). . . Fish Wild!. erv., in G. D. chmidt, ed . Problems in systematics of para- Spec. Sci. Rep.- Wild!. 125. 333 pp. it . ni ver ity Park Press, Baltimore, Md. McDonald, M. E. 1969b. Catalogue of helminth of water­ lark, G. M., D. O'Meara, and J. W. Van Weelden. 195 . fowl (Anatidae). U.S. Fi h Wildl. erv., Spec. Sci . An epizootic among ider ducks in volving an acantho­ Rep.-Wild!. 126. 692 pp. cephalid worm. J . Wild!. Manage. 22:204-205. McDonald, M. E. 1974. Key to reported in Co nnell, R. , and A. H. orner. 1957. Polymorphus para­ waterfow l. Bur. Sport Fish. Wild!. , Resour. Pub!. 122. do:~-us sp. nov . (Acanthocephala) parasitizing beavers and 44 pp. mu. krats in Alb rta, Canada. Can. J. Zoo!. 35:525-533. McDonald, M. E. 1981. Key to trematodes reported in rompton, D. W. T., a nd J . G. Harriso n. 1965. Observa­ waterfowl. Fish Wildl. Serv., Resour. Pub!. 142. 156 pp. tion on Polymorphus minutus (Goeze, 1782) (Acantho­ Macdo nald, J. W. , D. Lea, and G. A. Hamilton. 1978. c phala) from a wi ldfowl reserve in Kent. Parasitology Parasitic worms causing deaths of mute swans. Br. 55:345-355. Birds 71:358-359. D nny , M. 1969. Life-eye! of helminth parasites using Machado Filho, D. A. 1961a. ova contribuit;ao para o con­ amnta?"US lacustTi · a an intermed iate host in a Cana­ heci mento do genero "Corynosoma" Liihe, 1904 dian lak . Parasitology 59:795- 27. (Metacanthocephala, , Polymor­ Ga rd n, E . A., . Rayski, and V. M. Thorn. 1964. A para­ ph idae). Rev. Bras. Bioi. 21 :249-251. itic di a in eid r du ks. Bird Study 11:280-287. Machado Filho, D. A. 1961b. Contribuit;ao para o con­ Goez , J. 17 2. \ er u<: h ein r aturgeschichte der Einge­ hecimento do genero Corynosoma Liihe, 1904 (Metacan­ weid wurm r thierisch r Korper. Blankenburg, East thocephala, Palaeacanthocephala, Polymorphidae). G rmany. Pag s 139- 16 . Mem . Inst. Oswaldo Cruz Ri o J. 59: 1 1- 184. Machado Filho, D. A. 1962. Sobre "C01yno oma enriet­ chm id t, G. D. 1965a. Polymorphus swo1·tzi sp. n. , and tii" Molfi e Ferna ndes, 1953 (Metacanthocephala, other Acanthocephala of Al a kan du ·k . . .J. Parasilol. Palaeacanthocephala, Polymorphidae). Rev. Bra . Bioi. 51: 09- 13. 22:143-151. chmidt, G. D. 1965b. oryno ·oma bipapillum. p. n. from Molfi, A., and B. de Freita Fernand e . 1953. " o?yno­ Bonapart ' gull Laru · ph1'lad lphia in Al aska, with a oma enriettii" n. p., paras ita de patos e marreco note on C. const1·ictum Van Cleav , 19 l . J. Parasitol. domesticos (Palaeacanthocephala: Polymorphidae). Arq. 51 : 14- 16. Bioi. Tecnol. (Curitiba) :3-6. chmidt, G. D. 1972. Revi ion of th class Arch iacantho­ 0 hmarin, P. G. 1963. Para itic worms of mammal and cephala Meyer, 1931 (phylum A anthoc phala), with m­ birds in the Maritim Territory. [Rus ian text] Izd . phasi on Oligacanthorynchida , outh\ II et Ma fi , Akad. auk SSR, Mo cow. 322 pp. 1925. J. Para it I. 5 :290-297. Petrochenko, V. I. 1956. Acanthocephala (thornyheads) chmidt, G. D. 1973. Re urrection of 011lh'U'ellina Wi en­ of domestic and wild . Vol. I. [Russian text] lzd. berg, 1932 with a description of outhwellina dimo1·pha Akad. auk USSR, Moscow. 435 pp. sp. n. and a key to gen ra in Polymorphidae (Acantho­ Petrochenko, V. I. 195 . Acanthocephala (thornyhead ) cephala). J . Parasitol. 59:299-306. of domestic and wild animals. Vol. II. [Russian text] Izd . chmidt, G. D. 1975. Andracantha, a new g nu f Acan­ Akad. auk USSR, Moscow. 458 pp. thocephala (Polymorphidae) from fi h- ating bi rd , with Petrochenko, V. I. 197la. English tran lation of Petro­ description ofthr e pecie. J. Para ito!. 61:6 15- 620. chenko 1956. Israel Progr. Sci. Trans!., Jeru alem; .S. Schmidt, G. D., and R. E. Kun tz. 1967. ot s nth lif Department of Commerce, a tiona! Technical Informa­ eye! of Polymm·phus (Profilicollis).fonnosus p. n., and tion Service, Springfield, Va. records of Ar-hythmorhynchu · hispidns an I ave, Petrochenko, V. I. 197lb. English translation of Petro­ 1925 (Acanthocephala) from Taiwan. J. Para itol. chenko 1958. I rae! Prog r. ci. Tran I. J erusalem. .S. 53: 05- 09. Department of ommerce, ational Technical Informa­ Scott, P. 1977. A coloured key to the wildfowl of lh world. tion ervice, Springfield , Va. The Wildfowl Tru t, limbridg ; W. R. Royle and ons, Petrochenko, V.I., and G. A. Kotel'nikov. 1962. tili za­ London. 96 pp. tion of pond s for raising waterfowl and prophylaxis of Skrjabin, K. J. 1913. Zw· Acanthoc phalen-Fauna Ru sisch helminthiases. [Russian text] Izd. Min. Sel'khoz. RSFSR, Turke tan . A. Acanthocephalen d r umpf- und \ a - Moscow. 139 pp. ervog I. Zoo!. Jahrb. Abt. y t. Oekol. Ge r. Tiere Petrov. Yu. F., and Kh. I. Egizbaeva. 1972. The problem 35:403-414. of antagonistic in terrelations between hymenolepidids, Thom, V. M., and E. A. arden. 1955. A heavy morlalit polymorphids, and spirurata in gastro-in testinal tracts among eider duck . Fair I ·Ie Bird Ob rv. Bull. 2:32". of domestic ducks. [Russian text] Izv. Akad. auk Kaz. Thompson, A. B. 19 5. Pro.filicollis botulus (Acantho­ SR, Ser. Bioi. auk 3:48-51. cephala) abundance in the id r· duck ( omo le1·ia mollis­ Podesta, R. B. , and J. C. Holmes. 1970. The life cycle sima) on the Ythan stuary. Aberde n hir . Para it­ of three polymorphid s (Acanthocephala) occurring as ology 91:563-575. juvenile in Hyalella azteca (Amphipoda) at Cooking Van Cleave, H. J . 191 . The Acanthoc phala of orth Lake, Alberta. J . Parasitol. 56: 111 -1123. American bird . Trans. Am . 1icrosc. oc. 37:19-47. A list of current Resource Publications follows.

157. The Breeding Bird Survey: Its First Fifteen Years, 1965-1979, by ChandlerS. Robbins, Danny Bystrak, and Paul H. Geissler. 1986. 196 pp. 158. Techniques for Studying Nest Success of Ducks in Upland Habitats in the Prairie Pothole Region, by Albert T. Klett, Harold F. Duebbert, Craig A. Faanes, and Kenneth F. Higgins. 1986. 24 pp. 159. Research and Development Series: An Annotated Bibliography, 1889-1985, compiled by Thomas J. Cortese and Barbara A. Groshek. 1987. 160. Manual of Acute Toxicity: Interpretation and Data Base for 410 Chemicals and 66 Species of Freshwater Animals, by Foster L. Mayer and Mark R. Ellersieck. 1986. 579 pp. 161. Interpretation and Compendium of Historical Fire Accounts in the Northern Great Plains, by Kenneth F. Higgins. 1986. 39 pp. 162. Population Ecology of the Mallard. VIII. Winter Distribution Patterns and Survival Rates of Winter­ Banded Mallards, by James D. Nichols and James E. Hines. 1987. 154 pp. 163. Forested Wetlands of the Southeast: Review of Major Characteristics and Role in Maintaining Water Quality, by Parley V. Winger. 1986. 16 pp. 164. Effects of Contaminants on Naiad Mollusks (Unionidae): A Review, by Marian E. Havlik and Lei£ L. Marking. 1987. 20 pp. 165. Marking and Tagging of Aquatic Animals: An Indexed Bibliography, by Lee Emery and Richard Wydoski. 1987. 57 pp. 166. Checklist of of the United States, the U.S. Territories, and Canada, by Richard C. Banks, Roy W . McDiarmid, and Alfred L. Gardner. 1987. 79 pp. 167. Field Guide to Wildlife Diseases. Vol. 1. General Field Procedures and Diseases of Migratory Birds, by Milton Friend, Cynthia J. Laitman, and Randy Stothard Kampen. 1987. 225 pp. 168. Mourning Dove Nesting: Seasonal Patterns and Effects of September Hunting, by Paul H. Geissler, David D. Dolton, Rebecca Field, Richard A. Coon, H. Franklin Percival, Don W. Hayne, Lawrence D. Soileau, Ronnie R. George, James H. Dunks, and S. Dwight Bunnell. 1987. 33 pp. 169. Saltcedar Control for Wildlife Habitat Improvement in the Southwestern United States, by Theodore A. Kerpez and Norman S. Smith. 1987. 16 pp. 170. Pesticide Use and Toxicology in Relation to Wildlife: Organophosphorus and Carbamate Compounds, by Gregory J. Smith. 1987. 171 pp. 171. Sand and Gravel Pits as Fish and Wildlife Habitat in the Southwest, by William J. Matter and R. William Mannan. 1988. 11 pp. 172. Satellite 'Telemetry: A New Tool for Wildlife Research and Management, by Steven G. Fancy, Larry F. Pank, David C. Douglas, Catherine H. Curby, Gerald W. Garner, Steven C. Amstrup, and Wayne L. Regelin. 1988. 54 pp.

NOTE: Use of trade names does not imply U.S. Government endorsement of commercial products. 1111111 ~ ~~~~~ ~ ~nr' [l ~lili ~~ 11m~~ ijlrnfllll\1 11 1~1 11 111 e Nation's principal conservation agency, the Department of the Interior has 3 1822 00009991 1 )Sibility for most of our nationally owned public lands and natural resources. TillS mcludes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for the enjoyment of life through outdoor recrea­ tion. The Department assesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for peo­ ple who live in island territories under U.S. administration.

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