..

Bulletin of the SCANDINA VIAN SOCIETY

FOR PARASITOLOGY

WITH PROCEEDINGS FROM THE SYMPOSIUM ON ECOLOGY OF BIRD PARASITES, VILNIUS, LITHUANIA, 25-28 JUNE, 1998 Vol. 8 No. 2 1998 HUIJ.,ETI (JF TilE 'lET F(

Tlw Bnlldin is a mernbership journal of' the Scandinavian Society f()l· Parasitology, Besides mcinhership information, it also presents articles on all aspects of parasitology, with priority given to contributors Jhm1 the Nordic countries and other ECO: members of the Society. lt will include review artieks, short articles/communications. ( :omments on any topic within the field of parasitology may be presented as Letters to the Fdilor. The Bulletin is also open f�)r a short presentation of new projects. All contributions should be written in English. Review articles are commissioned by the editor, however, suggestions f�lr reviews are welcomed. Subscriptions arc available to non-members upon request from the Publisher. The subscription rate is SEK 300 per year (two issues annually). Subscriptions should be paid to the treasurer ofthe SSP:

Tor Atk Mo Na1ional Veterinary Institute P.O. Box 8156 Dep. N oo:n Oslo, NORWAY, e-mail: [email protected] Pos1al giro account number: 0814 3937489

Scandinavinn Society for Parasitology (Nordisk Forening fOrParasitologi) Society Board:

President: E. Tcllcrvo Valtonen (Finland) Vice-President: Karl Skirnisson (Iceland) General-Secretary: Maria Vang Johansen (Denmark) Treasurer: Tor Atle Mo (Norway) Board Member: Ingela Krantz (Sweden) Suppleants: Eskild Petersen (Denmark), Katarina Gustavsson (Sweden)

Cover: In Norse mythology, the giant ash tree - Yggdrasill - spreads its limbs over the entire mankind. The ash has three roots, each of them sucking water from its own spnng. The first spring- Hvergelmir- is found in the ice cold North; next to the spring, the serpent Niohoggr is ceaselessly gnawing at the roots of the ash. The second spring­ Mimisbrunnr - is the source of wisdom and is guarded by Mimir. The third spring­ Uroarbrunnr - is guarded by three women, the Noms, which mete out man's thread of life. Editors: Gedimim

Organh Lithuania� Telephom

Printed in Norway by HS-Trykk A/S ISSN 0803-4907 EMO PROCEEDINGS A AAMATUKOGU of the symposium on

ECOLOGY OF BIRD-PARASITE INTERACTIONS

arranged on behalf of the

I 11hl1 lt ·r. Baltic Society for Parasitology and the 111 la uld Scandinavian Society for Parasitology

in Vilnius, Lithuania

25 - 28 June, 1998

!lsitologi)

>s over nits own ng, the 5pring­ pnng­ thread of

Editors: Gediminas Valkiiinas and Hans-Peter Fagerholm

Organizing and Scientific Secretariat: Lithuanian Academy of Sciences, Gedimino ave. 3, Vilnius 2600, Lithuania. Telephone: + 370 2 614011, telefax: + 370 2 618464, e-mail: [email protected]

803-4907 11

Scientific Programme and Organizing Committee: PREl Hm1s-Petcr Fagerholm, Abo Akademi University, Finland

Lars Gustafsson, University of Uppsala, Sweden Tl was b Toivo Jarvis, Estonian Agricultural University, Estonia for Pa th Vytautas Kontrimavicius, Lithuanian Academy of Sciences, Lithuania 25 at the S Peter Nansen, The Danish Centre for Experimental Parasitology, Denmark Zoone

Karl Skfrnisson, University of Iceland, Iceland Tl streng Arne Skorping, University of Bergen, Norway parasi Gediminas Valkiiinas, Institute of Ecology, Lithuania (Chairman) ecolog discus co-op( focusi Local Organizing Committee: opport attract Vytautas KontrimaviCius, Lithuanian Academy of Sciences, Lithuania apperu Gediminas Valkiiinas, Institute of Ecology, Lithuania (Chairman) bird iJ spectn Juozas Virbickas, Institute of Ecology, Lithuania popuh Birute Vosylyte, Lithuanian Academy of Sciences, Lithuania exam]: impac1 etc. Tl of ora hope tl ornith< Sponsors: scieno The Nordic Academy for Advanced Studies (NorFA) Ot thanks Lithuanian Academy of Sciences the su Institute of Ecology (Lithuania) sponsc from tJ The Open Society Fund of Lithuania issue c Lithuanian State Science and Studies Foundation TI the ine The Scandinavian Society for Parasitology (SSP)

The British Society for Parasitology 28 Sef The Baltic Society for Parasitology (BSP) Gedin Lithuanian Exhibition Centre "Litexpo" Hans-: 111

PREFACE

The Baltic-Scandinavian symposium on "Ecology of Bird-Parasite Interactions" was held on behalf of the Baltic Society for Parasitology and the Scandinavian Society for Parasitology at the Lithuanian Academy of Sciences in Vilnius, Lithuania between 25th and 28th June 1998. It was the second parasitological meeting jointly organized by the Societies. The first Baltic-Scandinavian parasitological meeting "Parasitic Zoonoses and the Ecology of Parasites" was held in Lithuania in September 1994. The Symposium on "Ecology of Bird-Parasite Interactions" was organized to strengthen the professional links, and to exchange ideas and information between parasitologists, ecologists, evolutionary biologists and ornithologists in the field of ecology of bird-parasite interactions. The purpose of the meeting was to present and discuss results and to stimulate future research in the area as well as to develop such co-operative projects in bird ecology that take parasitism into consideration. Owing to focusing on a single taxonomic host group, the symposium provided a unique opportunity for extensive interdisciplinary discussions on the subject. The meeting attracted more than 60 persons from 18 European countries, USA and Australia, as will appear from the present proceedings. During the symposium, the fascinating parasite­ bird interactions were analysed ranging from a molecular level to the cdl:nmunity spectrum with most participants interested in interactions on individual level and in populations. Several general problems of ecological parasitology were discussed. For example, migration and parasite dispersal, host fitness and population consequences, impact of parasites on bird population dynamics, bias of sampling in bird parasitology, etc. The papers based on plenary lectures given by invited speakers as well as abstracts of oral and poster presentations, are published in this issue of the SSP Bulletin. We hope the contributions will be stimulating for the advancement of ecology, parasitology, ornithology and evolutionary biology as well as for the interaction of these biological sciences. On behalf of the organizing committee, we are pleased to express our deepest thanks to the invited lectures and other participants for their valuable contributions to the success of the symposium. We wish to record special thanks to the symposium sponsors: without this support the symposium would never have taken place. Support from the Scandinavian Society of Parasitology made it possible to produce the present issue of the Bulletin. Through an editorial process, some linguistic corrections have been introduced, but the individual authors are responsible for thepresented terminology, interpretations, etc.

28 September 1998

GediminasValkiiinas

Hans-Peter Fagerholm iv

CONTENTS Char seas< Belo. INVITED LECTURES Para: Ti cks and mites as permanent and temporary bird ectoparasites, pathogensvectors and Bick1 disseminators

Alekseev, Andrey N ...... 1 Thei Bone Parasites as regulators of natural populations: Fashion as a confounding variable

Ashfo rd, Richard W...... 9 Heln origi Longstanding associations between bird hosts and nematode parasites as Bom manifested by conservative structural features in some ascaridoid nematodes

Fagerholm, Hans-Peter ...... 15 Para: Fred Parasites in relation to life history and secondary sexual traits

Gustafsson, Lars ...... 17 Anal ente1 The consequences of parasite induced effects on the population dynamics of birds Dalii Hudson, Peter J ?...... 18 Spec Aquatic birds as agents of parasite dispersal: A field test of the effectiveness of, plat) helminth colonisation strategies By eh Kennedy, Clive R...... 23 Para: Bird-parasite-viruses: interpopulational interactions By en Lvov, Dmitri K...... 29 Expe Darwinian medicine? Lessons from avian blood parasite ecology (Ces1 Read, Andrew F. & Shut/er, Dave ...... 30 Ciel1 Macroparasites as selective agents in birds Gryt

Skorping, Arne ...... 38 Hem Haematozoa of wild birds: Peculiarities in their distribution and pathogenecity chicl

Valkiiinas, Gediminas ...... 39 Cod1

The Cure

SUBMITTED ABSTRACTS OF ORAL AND POSTER PRESENTATIONS Com D. rt A vi an parasite databases and collections: Current status of the international Czat reference centre for avian haematozoa (IRCAH)

Adlard, Robert D...... 47 Dior COffiJ Wsevolod Borisovich Dubinin (1913-1958) as a researcher of bird parasites Czat Alimov, Aleksander F. & Dubinina, Helen V...... 48 V

Changes in population structure of an argasid tick Argas arboreus during nesting season of its host, the cattleegret Bubulcus ibis, in South Africa

Belozerov, Valentin N. & Kopij, Grzegorz ...... 48

Parasite infection in the great tit )rs and Bickle, Angharad & Clark, F ...... 49

...... 1 The interdmediate host as ecological determinant of tapeworm fauna of gulls

Bondarenko, Svetlana ...... , ...... 50 le ...... 9 Helminth faunae of Charadriiformes in Alaska: Peculiarities of zoo geography and origin Bondarenko, Svetlana & Kontrimaviaus, Vytautas ...... , ...... 51

...... 15 Parasitic infections of birds of prey in the Netherlands

Fred H.M. Borgsteede and Pedro E.F. Zoun...... 51

...... 17 Analysis of retaliatory reactions in domestic quails after challenge by Salmonella enteritidis ds Dalius Butkauskas and Jonas Babonas ...... 18 . . .51 Special features of formation of fauna of helminths in urban and wild mallard {\nas platyrhynchos L. populations Bychkova, Elizabeta ...... 23 ...... , ...... 52 Parasites of falcons in Belarus

Bychkova, Elizabeta Vintchevski, Alexandr ...... 29 & ...... , ...... 53 Experimental trials to infett domestic ducks withcoot diorchids ...... 30 (Cestoda, Hymenolepididae) Cielecka, Danuta, Czapliiiski, Bogdan, Olszewska, Gizela &

Grytner-Zi�cina, Barbara ...... , ...... 53 ...... 38 Hematological and biochemical investigations on experimental coccidiosis in meat chickens

...... 39 Codreanu-Balcescu, Doina & Curca , Dumitru ...... 54

The ascorbic acid, stress and poultry reactivity in parasitic diseases

Curca,Dumitru & Codreanu-Balcescu, Doina ...... 55

'IS Consequenses of a higly probable error in labelling Diorchis injlatus and D. ransomi fromAythya nyroca Gilldenstadt Czapliiiski, Bogdan & Cielecka, Danuta...... 56

...... 47 Diorchis acuminatus (Clerc, 1902) Clerc, 1903 (Cestoda, Hymenolepididae)- A complicated type of the genus Diorchis ...... 48 Czapliliski, Bogdan & Cielecka, Danuta ...... 56 vi

Host speci:ficity of Cladogynia pittalugai (Lopez-Neyra, 1932) Czapliiiski, Bogdan, Czapliiiska, Urszula , Grytner-Zi�cina,Barbara &

Gierczak, Anna ...... 51

Isospora coccodia (Protozoa, Eimeriidae) of passerine birds on the Courish spit

Dolnik, Olga ...... 58

Borrelia genospecies prevalence as a marker of the possible role of birds in the distributionof tick-borne spirochetosesin the baltic region

Dubinina, Helen V. & Alekseev, Andrey N...... 58

Ticks (Ixodoidea, Ixodidae) of migratory birds on the Kurish spit

Efremova, Galina ...... 59

Parasitic arthropods of birds and their role in maintenance of natural foci of Arbovirus diseases in Belarus

Efremova, Galina. and Gembitsky, A...... 60

Blood protozoan parasites (Protozoa: Kinetoplastida and Haemosporida) of wild birds from Bulgaria

Golemansky, V., Pilarska, D., Zehtindjiev, Pavel, Valkiiinas, G. & Iezhova, T ...... 61

The prevalence of Sarcocystis in some wild birds and poultryin Lithuania and in neighbouring territories

Grikienienci Jadvyga & lezhova, Tatjana ...... 61

Occurrence of Borrelia burgdoiferisensu lato Spirochetes in passerine birds

Gryczynska, Alicja & Zgodka, Andrzej ...... 62

On the speci:ficity of Fimbriaria sarcinalis (Cestoda: Hymenolepididae) in relation to hosts

Grytner-Zit;.cina, Barbara & Cielecka, Danuta ...... 63

The ecology of the haematozoan parasites of European blackbirds Turdus merula, and their effects on host fitness

Hatchwell, B.J., Anwar, M., Wood, M.J. Chamberlain, D. & Perrins C.M...... 63

A new nasal bird schistosome fromEurope

Horak, Petr, Kolarovd, Libuse & Dvorak, Jan ...... 64

Haematozoa as biological tags in bird subspecies studies

Iezhova, Tatjana & Valkiiinas, Gediminas ...... 64

Investigations into bird parasitodes in Estonia

Jiirvis, Toivo ...... 65 viii

Chromosome studies on some cyclophyllidean cestodes, the parasites of birds of northwestern Chukotka

...... 57 PetkeviCiilte, Romualda & Regel, Kira V...... 7

Developmental stages of bird trematodes in populations of the pond snail Lymnaea stagnalis (L.) inhabiting anthropogenic water environments in the upper Silesian ...... 58 industrial region (Southern Poland)

Pokora, Zbigniew ...... 7

Ability of host to recover from blood parasite infections - a longitudinal study in ...... 58 the black grouse Ratti, Osmo, Alatalo, Rauno V. &Rintamaki, Pekka T...... 7

...... 59 Pisci vorous birds in the helminths development cycle of the Gulf of Gdansk (the Baltic Sea)

Rokicki, Jerzy, Rolbiecki, Leszek & Okulewicz, Anna ...... 7

...... 60 Parasites of the rock ptarmigan (Lagopus mutus) in Iceland

Skfmisson, Karl ...... 7 j Specific features of anti-salmonela immunity developed by "TALOVAC 1p9 SE" ...... 61 and "SG VACCINE NOBILIS" vaccines Slavenaite, Sigita, Sruoga, Aniolas, Butkauskas, Dalius & Skonsmanas, Jonas ...... 8 n Evolutionary relationships among avian and mammalian blood flukes (Digenea: ...... 61 Schistosomatidae)

Snyder, Scott D...... 8

...... 62 Allergenic acarofauna from nests ofse lected species of synanthropicbirds inPoland Solarz, Krzysztof, Szilman, Piotr & Szilman, Ewa ...... 8

New data on the karyotypes of genus Ichthyocotylurus odening, 1969 (Trematoda, ...... 63 Strigeidae) StaneviCiilte, Graiina & Kiseliene, Valerija ...... 8 la, Some peculiarities of the distribution of rhinonyssid mites (Acari, Gamasina, ...... 63 Rhinonyssidae) of birds Stanyukovich, Maria K. & Butenko, O.M...... 8

...... 64 Frenkelia, buzzards and rodents: Multiple mechanisms of parasite survival strategy Svobodova, Milena, Votjpka, fan & Vofisek, Petr ...... 8

...... 64 Conformity of phenologies of mallard Anas platyrhynchosL. and insect vectors Tereshkina, Nadezhda & Kozulin, Alexandr ...... 8

...... 65 Investigating the regulation of parasitic nematode fecundity Tompkins, Daniel M...... 8 viii

Rate« Chromosome studies on some cyclophyllidean cestodes, vacci1 the parasites of birds of northwestern Chukotka Vank1 Petkeviaiite, Romualda & Regel, Kira V...... 76 Preva Developmental stages of bird trematodesin populations of the pond snail Lymnaea stagnalis (L.) Czecl inhabiting anthropogenic water environments in the upper Silesian Volf, industrial region (Southern Poland)

Pokora, Zbigniew ...... 77 Blocx Repul Ability of host to recover from blood parasite infections - a longitudinal study in Vo tjr the black grouse Vo lf, Ratti, Osmo, Alatalo, Rauno V. & Rintamaki, Pekka T...... 78

Pisci vorous birds in the helminths development cycle of the Gulf of Gdansk (the Baltic Sea)

Rokicki, Jerzy, Rolbiecki, Leszek & Okulewicz, Anna ...... 78

Parasites of the rock ptarmigan (Lagopus mutus) in Iceland

Skfrnisson, Karl ...... 79

Specific features of anti-salmonela immunity developed by "TALOV AC 109 SE" and "SG VACCINE NOBILlS" vaccines

Slav&zaite, Sigita, Sruoga, Aniolas, Butkauskas, Dalius & Skonsmanas, Jonas ...... 80

Evolutionary relationships among avian and mammalian blood flukes (Digenea: Schistosomatidae)

Snyder, Scott D...... 80

Allergenic acarofauna fromnests of selectedspecies of synanthropic birds inPoland

Solarz, Krzysztoj, Szilman, Piotr & Szilman, Ewa ...... 81

New data on the karyotypes of genus Ichthyocotylurusodening, 1969 (Trematoda, Strigeidae)

Staneviaiite, Graiina & Kiseliene, Valerija ...... 82

Some peculiarities of the distribution of rhinonyssid mites (Acari, Gamasina, Rhinonyssidae) of birds

Stanyukovich, Maria K. & Butenko, O.M...... 83

Frenkelia, buzzards and rodents: Multiplemechanisms of parasite survival strategy Svobodova, Milena, Votjpka, fan & Vofisek, Petr ...... 83

Conformity of phenologies of mallard Anas platyrhynchosL. and insect vectors

Tereshkina, Nadezhda & Kozulin, Alexandr ...... 84

Investigating the regulation of parasitic nematode fecundity Tompkins, Daniel M...... 85 ix

Rate of antibody titer after challenge by virulent Salmonella enteritidis culture in vaccinated hens

Vankeviaene, Birute, & Butkauskas, Dalius ...... 85 ...... 76 �a ea Prevalence of coccidia of the genus Caryospora in captive birds of prey in the Czech Republic lll Volf,]ifi & Modry, David ...... 86

...... 77 Blood parasites and haematophagous insects of raptors (Falconiformes) in the Czech Republic n Vorypka, fan, Svobodowi, Milena, Vofisek, Petr, Peske, LuboS, Lacina, David &

Volf, Petr ...... 87 ...... 78

...... 78

...... 79

E"

> ...... 80

...... 80

J ...... 81 Dda,

...... 82

...... 83

tlcgy ...... 83

s ...... 84

...... 85 Bull Scand Soc Parasito/1998; 8 (2): 1-8

TICKS AND MITES AS PERMANENT AND TEMPORARY BIRD ECTOPARASITES, VECTORS AND DISSEMINATORS OF PATHOGENS

Andrey N. Alekseev

Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia

Mites and ticks were frrst named by 2. Temporary parasites - inhabitants of Aristotle, given the name t1x it Q11 � bird nests, transmitted mainly hori­ (Acaris - in Latin) that literally means zontally (11). 3. Temporary parasites, obligate blood­ "very short, small". The first description suckers, which use ground-feeding and of a bird mite, most probably Pteronyssus domestic birds as hosts (III). ' truncatus- according W. Dubinin (1951), th Pathogen reservoirs and disseminators was made at the end of the XVII- cen­ of disease agents are found mainly, if not tury. Now there are nearly fifty mite and exclusively, in the last two groups. There tick families known to be parasites of is a widely held opinion and increasing birds (Fig. 1) •. empirical evidence that vertically trans­ All representatives of the families can mitted parasites, including mites, are less be divided into three large ecological virulent than horizontally transmitted groups: ones. A significant number of exceptions 1. Permanent bird parasites: mites, which to this rule will be examined here. The are transmitted as a rule vertically - principal scheme of mite development from parents to nestling (I in Fig. 1). includes adults, ova, larvae and up to 3 nymphal stages. All, apart from the ova, may include parasitic stages. Amongst the bloodsucking ticks, ixodids have only active stages, whereas argasids have * This scheme was made on the basis of three the well known system constructed by one larval and a variable number of Krantz (1978) and fulfilled and corrected nymphal stages. by using the data of Russian scientists Freyanoidea mites - the typical repre­ (Bregetova, 1956; Dubinin, 1951, 1953, sentatives of the frrst ecological group - 1957; Filippova, 1966, 1977; Gilarov, live on the barbs of the feathers mainly of Bregetova, 1975, 1977; Volgin, 1969) aquatic birds. Their detrimental effects and other acarologists (Kethley, 1982; have not been adequately investigated. O'Connor, 1982), whose systems were The majority of Pterolichoidea species supported by theRussian taxonomists. live as external commensals or 2

Fi�

Supercohort parasi Order Suborder Cohort Superfamily family Subfatily Localilation Bol Subcohort paras it Mi1 xodidae-,Ixodinae II,III b,v x s:------Ixodoidea 4\ablyOHioae III b,v skin; s rgasidaa-p\rgasioae II, b,v ofpatl: t 4lrnithodorinae II, b,v PILIOACARIHA Nuttalliellida11

odide parasi tIxodida) but se Gausillit----Galasoi Rhinonyssidae I, rt harmfi ARASITIFORHES �ejina liponyssi dae II, b,v ercoaegistina /facrinyssidae) obtust esostigaata ntennophorina lOer•anyssidae II, b,v (Gaaasida) ropodina 10 weigh1 ill-tim 3 faailies I ( pionacaridae I, fq birds. raoglyphidae I, fq nalgoidea pideraoptidaefa�ilies I, s other 1 caridides ne�idocoptidae I, s,sc 4 inhabi ainosioptidae I, sc,ff reyanoidea--3 faailies I, f flight l Psoropti

ytoditoidea---tytodi tidae I, rt parasi1 soroptoidea cause istrophoroidea biloba arcoptoidea freynetidaig peleogna�inae I, rt the na ndeost igM tides CARIFORHES abidos toaMa tin a -iTydaeidae hen sb reynetinae . �ydeoidea (Dubir Troabidifoms upod1na �a"rencannae (Actinedida, Iol inidae (Fain, Prostigmata) upodoidea of the delloidea alacaroidea tacheyletiinae I, fq leg" di Cheyletida nithocheyletiinae I, s nystae distrib1 t I, fq,pr birds. [Jdj--"'1'"'"'"''''"'"aphignathoidea I Naphigllitha Cheyl11toide arpi rhynchidae I, se docop1 Syringophilidae I, fq CloacaridaerPneuaophaginaeeyletinae I, rt fingert lCI Cheletosoaa tin i) subfati ies dae, D inhabit flaking Lee1111enhoekiidaeoaca ri nae 11, s Parasitengolli Tra.bidioi�Trombiculidaefailies ll, s,b,v and ea Tetranychoidea 5 riophyoidea their fe 1aliptostoaatoidea probab (dogs z families rythraeoidea ydrachnidia ribatida (Cri ptos ti gaata) 3

Fig. 1 (previous page). Scheme of phylogenetic relationships of mite and tick bird parasites. ion Bold letters - bird parasites on ly ; cursive bold letter - taxo nomic groups in which bird parasites are present . I- first, 11- 2-nd and Ill- 3- rd ecological group of acari-bird parasites. Mite parasites of: f-feather; ff - feather fo llicles; fq - feather quill; rt - respiratory tract; s - b,v b,v skin ; se- subcutaneous tissues ; a! - mites as source of allergens; b - bloodsuckers; v - vector b,v of pathogens; pr - predacious mites, inhabitants in the feather quill. b,v

parasites on the wing and tail feathers, inhabit the follicles of developing feathers but some species are known to be very or subcutaneous tissues of birds. rt harmful bird parasites. Pterolichus Many mite families have a specific b,v obtusus, for example, causes agitation, phoretic stage of their development - the b,v weight loss, reduced egg production and deutonymph, also called the hypopus. In ill-timed moult of wild and domestic the suborder Sarcoptiformes, there is a fq birds. Syringobiidae mites differ from the very interesting family Hypoderidae fq 5 other Pterolichoidea in that the species all (Fain, 1967) which inhabit nests of birds s,sc inhabit the space within the quills on the of many orders; their deutonymphs are se,ff f flight feathers. parasitic in the subcutaneous, and more f,fq,sc sc,ff Six out of 13 families of Analgoidea rarely the visceral, tissues of theihost and s,sc fq mites inhabit wingfeathers and do not increase their body size 4- tb 5-fold rt damage their host seriously, whereas all during development. Deutonymphs of this representatives of Epidermoptidae are mite have neither a mouth nor functional

rt parasites of the bird skin, often being a gut. The life cycle of Hypoderoidea is cause of serious diseases. Epidermoptes closely tied to those of their hosts, with bilobatus, the "hen scabies" agent, injures new deutonymphs being produced near rt the nape, throat and breast parts of the the hatching time of the hosts. hen skin. Heavy invasion kills the host The group of Pyroglyphidae mites (Dubinin, 1953). Knemidocoptidae mites includes free living forms, nest inhabi­ (Fain, Elsen, 1967) cause serious injuries tants and parasites of bird skin and inner­ of the bird legs (Dubinin, 1953), "scaly­ skin layers (Gaud, 1968). Representatives

fq leg" disease. Knemidocoptiasis is widely of two first subgroups can serve as a s distributed among wild and domestic strong source of allergens, from which fq,pr birds. When heavily infested by Knemi­ not only birds suffer: inhabitants of se docoptes mutans, birds sometimes lose synanthropic bird nests also cause aller­ fq rt fingertips or a whole finger. Apionacari­ gies in man. dae, Dermoglyphidae mites (Analgoidea) Trombidioidea (Trombidiformes) inhabit the space within the quills, feed on mites (Vercammen-Grandjean, 1968; flaking tissues at the base of the quills Vercammen-Grandjean & Langston, 5 s,b,v and cause the host to react by pulling out 1971-1976, in: Krantz, 1978) are bio­ their feathers. Laminosioptidae were most logically distinct from the Sarcoptiformes probably first parasites of the Mammalia bird-parasite. As a rule, their larval stage (dogs and rodents) (Dubinin, 1951) which is parasitic but the deutonymph or adult are not. Some of them are even able to 4

kill their hosts, e.g. Apolonia tigipioensis of the same suborder, inhabit the mid to spend (Leeuwenhoekiidae) is fatal to chickens in lower respiratory tract of birds. Speleog­ rial, ( Brazil (Torres & Braga, 1939: in Krantz, nathid mites (suborder Trombidiformes), blood 1978). Some other Trombiculidae larvae being much more mobile and pathogenic, belly use many species of birds as a source of are distributed along the whole respira­ brood blood (Tamiya, 1962, in: Kulagin & tory tract and often cause bird death. uneas Tarasevich, 1972). These bloodsuckers Rhinonyssid mites are the only eodo­ weigh include one of the main vectors of scrub parasitic group among Gamasoidea death typhus (tsutsugamushi fever), Lepto­ (Parasitiformes). Their feeding stage is effect trombidium akamushi. It seems very the protonymph. They inhabit the bird Tomr

probable that Leptotrombidium deliense - nostrils (Fain, 1957), and lungs (Butenko, gall in the major vector of scrub typhus in the 1984) and sometimes cause a lethal spam Asian and Pacific region (Kethley, 1982) pneumonia. Many species of Rhinonyssi­ disea� - uses migratory birds as hosts and in this dae are bloodsucking mites. acari:: way can serve to disseminate the scrub Clayton & Tompkins (1995) wrote to sur typhus agent. Harpirinchidae mites that the virulence of ectoparasites (i.e. the 9 mo (Cheyletoidea) are skin parasites found on harm they cause) is linked to mode of specie the surface of, or in, the skin of the transmission, e.g. horizontally transmitted chetol throat, neck, breast and underside of the Mesostigmatid mites are much more res en wings. Injuries are sometimes heavy: harmful in comparison with vertically patho1 severe host tissue reaction appears, transmitted Analgesoides mites (Dubinin, So resulting in greatly enlarged tumorlike 1951). Horizontally transmitted Gama­ being growths. Some species of Cheyletidae and soidea, being bloodsucking mites, very many all Syringophilidae live inside the quills of often have as hosts not only different sid n feathers and feed on subcutaneous fluids birds, but also mammals and reptiles, and stages by piercing the wall of the quill. A single can serve as the reservoirs, vectors and many bird may host many mite species simulta­ disseminators of many pathogens such as state t neously, with each species inhabiting viruses, rickettsiae, bacteriae, spirochetes uted 1 different feather tracts. Inside the feather and protozoans (Bregetova, 1956). Or­ mate, quill, predacious cheyletids nithonyssus sylviarum (Liponyssidae) is are a (Cheletosomatini) make a living by known as a parasite of wild migratory can d hunting on the other parasite mites birds, but in USA and Canada is a prob­ effect1 (Volgin, 1969). lem in the poultry houses where it can vector The whole bird respiratory tract, being bite man and is suspected as an Ornitho­ the bi: a rich source of easily available food sis vector. The most dangerous bird pullo1 (mucus, frail cells, blood under the thin parasites are in the Dermanyssidae fam­ They, vessel walls) serves as a niche for many ily. Three stages of their development - of wh endoparasitic mite species of Acarifor­ adults, proto- and deutonymphs - are breab mes. Turbinoptidae (Sarcoptiformes) are bloodsuckers. They can survive being (Tadji small and sluggish mites that inhabit bird frozen, their reproductive ability is great Gissat nostrils and sometimes affect also the and their distribution is world-wide. bites 1 deeper parts of the nasal cavities (Fain, Dermanyssus J?allinae is a well known garis, 1957). Cytoditidae, another representative pest in t11e poultry houses. These mites prove< 5

e mid to spend most of their time in the nest mate­ rus can be harboured and reproduced in Speleog­ rial, emerging periodically to feed on the the A. persicus body (Nemova et al, iformes), blood either of nestlings or of the lower 1990). thogenic, belly of adults that are incubating eggs or Hard ticks (Ixodidae) belong to the respira­ brooding the young. They cause bird second and third ecological groups. The d death. uneasiness, feather loss, decrease of latter plays the most important role as ly eodo­ weight and oviposition, and sometimes pathogen reservoirs and vectors. Ixodids masoidea death of chickens. These are typical have to be subdivided on 3 ecological stage is effects on wild birds also (Clayton & subgroups. In the first subgroup are the the bird Tompkins, 1995). Being synanthropic, D. permanent nest parasites such as Ixodes Butenko, gallinae mites, parasites of pigeons and lividus. They can be reservoirs of many a lethal sparrows, often bite man and cause pathogens, from arboviruses to proto­ inonyssi- disease which even has the specific term zoans, but can rarely serve as the dis­ acariasis (Zemskaya, 1973). Their ability seminators of pathogens, which are 5) wrote to survive in a hungry state for more than instead transferred mainly by their bird � (i.e. the 9 months makes this widely distributed hosts (Filippova, 1977). Ixodes uriae, mode of species not only a vector of hen spiro­ typical seabird parasite that inhabit cliff­ msmitted chetosis and Asian bird plague, but a nesting seabird colonies, belongs to the �h more reservoir and a disseminator of these second subgroup. Its life is only partly vertically pathogens. connected with bird nests as it moults and Dubinin, Soft ticks (Argasidae, Ixodoidea), survives off its hosts within the cracks in l Gama­ being inhabitants of nests and holes, have the rock face rather than in the nests tes, very many features in common with the gama­ themselves. They feed on the chicks of different sid mites (2nd ecological group). All seabirds (e.g. on kittiwake during 3-6 tiles, and stages are bloodsuckers. They can survive days [Barton et al, 1995]) and can serve :tors and many months (even years) in a hungry as the main pathogen reservoir. Migra­ �such as state (Filippova, 1966). They are distrib­ tory seabirds transport many pathogens ,irochetes uted mainly in countries with a hot cli­ (Lvov & Jliychev, 1979), whose survival 56). Or­ mate, are very resistant to the dryness and and transmission cycles within foci are :sidae) is are a great pest in poultry houses. Birds effected by I. uriae. nigratory can die through anaemia or from the Ixodid representatives of the third � a prob­ effects of salivary toxins. Argasids are subgroup are parasites of ground-feeding re it can vectors of Borrelia anserina, the cause of birds. They are represented mostly by Ornitho­ the bird spirochetosis, and Aegyptionella larvae and nymphs caught on the host. )US bird pullorum, bird hemosporidiosis agent. The cause of such predominance might be dae fam­ They can also harbour arboviruses, some due less to any preferential ability of host )pment - of which are pathogenic for man. Out­ attachment by the pre-imaginal stages 1S - are breaks of fevers in Dushanbe (adult females are much more mobile and ve being (Tadjikistan) caused by Issyk-Kul and aggressive [Alekseev et al, in press]), but ' is great Gissar arboviruses were connected with the result of avian tick paralysis caused rid-wide. bites of Argas persicus and Argas vul­ by engorged adult females, e.g. Ixodes 11 known garis, parasites of synanthropic birds. We brunneus engorged females kill passer­ �se mites proved experimentally that Gissar arbovi- ines (Luttrell et al, 1996). 6

collecte The representatives of the third group 1998). These results of our investigation Baltic l of ticks are 2- or 3-host parasites and, concur with the data of Olsen et al. 1998; 3 being the most ancient bloodsucking (1995). According to Postic et al. (1997) arthropcxls, can transmit pathogens of and our data (Alekseev et al, 1998), Aleksee many taxonomic groups excluding Plas­ Borrelia garinii were found mainly in I. Makrou of imag modia (Alekseev, 1985). Parasites of persulcatus. This phenomen may serve as persulc. ground-feeding birds (ticks as well as proof that the interface in the triad: bird - Parazit< mites) can create extensive contacts I. persulcatus - B. garinii is much more between different reservoirs of pathogens: important than has hitherto been consid­ Anders< larvae engorged on rodents may continue ered. Ticks transferred by birds can re­ voirs f Acad s, their life cycle feeding on birds, emerged establish natural foci of diseases on the adults can transmit pathogen to man. territories where they were previously Anders• Birds are a known reservoir of the Borre­ eradicated, e.g. by DDT (Chunikhin, KC. Bi mit lia garinii - agent of neuroborreliosis 1966). Bo (Van Dam et al, 1993) and sometimes Our survey allows us to conclude that bamsteJ serve as a source of more than one gena­ mites and ticks, creatures named by Bar ton species for the ticks that fed on them Aristotle "short, small", play not a small attachn (Humair et al, 1996). The aggregated but a very significant role as bird para­ Ixodes spatial pattern of preimaginal tick stages sites, vectors and disseminators of bird Rissa t; feeding on birds (Barton et al, 1995) and man pathogens, and probably also 1995; 1 allows the direct exchange of pathogens host population regulators. Bregetc between cofeeding ticks (see survey: (Gamm Randolf et al, 1996). The role of migra­ Acknowledgements stvo "N tory birds and their parasitic ticks as I wish express my gratitude to Dr. tion of disseminators of borreliae is now well H.V. Dubinina, Dr. S.V. Mironov and Butenk proven (Anderson et al, 1988, 1990; Dr. A. V. Bochkov (Zoological Institute, serine 1 Miyamoto et al, 1993; Nicholls & Cal­ Russian Academy of Sciences, St.­ stvo M< 1996 lister, and other). Ticks that feed on Petersburg, Russia) for her help and Dr. Claytor birds transfer different borrelia species S. Randolph (Oxford Tick Research effects from North to South (Olsen et al, 1995), Group, Department of Zoology, Oxford, success playing a role of pathogen mixers in the UK) for her valuable advices and im­ Parasit• natural foci of borrCliosis. Ixodes ricinus provement of the English version of the Chunil; persulcatus and I. collected by us on the manuscript. borne ( West-Europe migratory routes of birds Mosc01 (Finnish Gulf beach, Kurish Spit) demon­ References Russiat strated a marked difference in the borrelia Alekseev AN. The theory of connections of species with which they are infected: I. feeding types and digestion of blood-sucking Dubini: ricinus were infected mostly by Borrelia arthropods with their ability to be specific I. Intro afzelii, whereas I. persulcatus by Borre­ vectors of transmissive diseases agents. dae an• dae fat lia garinii. I. ricinus collected on the Parazitologia 1985; 19: 3-7 (in Russian) "Naukz Kurish Spit in the fall contain 8 times Alekseev AN, Dubinina HV, Antykova LP et USSRJ more unidentified spirochetes than ticks al. Tick-borne borreliosis patlwgcn identifi­ (in Rus collected in the spring (Al ekseev et al, cation in Ixodes ticks (Acarina, Ixodidae) 7

�stigation collected in St. Petersburg and Kaliningrad Fain A. Les hypopes parasites des tissus �n et al. Baltic Regions of Russia. J Med Entomol cellulaires des oiseaux (Hypodectidae: 1998; 35: 136-42 Sarcoptiformes). Bull Inst r Sci nat Belg :!. (1997) 1967; 43 r, 1998), Alekseev AN, Dubinina HV, Smimova LA, inly in /. Makrouchina NA. Differences in behaviour Fain A, Elsen P. Les acariens de la fa mille of imaginal and preimaginal stages of Ixodes r serve as Knemidokoptidae producteurs de gale chez ad: bird- persulcatus and Ixodes ricinus ticks. Med les oiseaux (Sarcoptiformes). Acta Zoo! Parazitol Moscow (in Russian). In press Pathol Antverp 1967; 45 1ch more n. consid­ Anderson JF. Mammalian and Avian Reser­ Filippova NA. Argasid ticks (Argasidae). � can re­ voirs for Borrelia burgdoiferi. Ann NY Leningrad, lzdatelstvo "Nauka", 1966 (In �s on the Acad Sci 1988; 539: 180-9 1 series: The USSR fauna. Paukoobraznii. T. 4, N 3) (in Russian) reviously Anderson JF, Magnarelli LA, Stafford Ill hunikhin, KC. Bird-feeding ticks transstadially trans­ Filippova NA. Ixodid ticks subfamily Ixodi­ mit Borrelia burgdoiferi that infect Syrian nae. Leningrad, Izdatelstvo "Nauka", 1977 :lude that hamsters. J Wild! Dis 1990; 26: 1-10 (In series: The USSR fauna. Paukoobraznii. T. 4, N 4) (in Russian) 1med by Barton TR, Harris MP, Wanless S. Natural t a small attachment duration of nymphs of the tick Gaud J. Acariens de la sous-famille des ird para­ Ixodes uriae (Acari: Ixodidae) on kittiwake Dermatophagoidinae recoltes clans les ; of bird Rissa tridactyla nestlings. Exp Appl Acarol plumes des oiseaux. Acarologia 1968; 10: 1bly also 1995; 19: 499-509 292-312 Bregetova NG. The Gamasid mites Gilarov MS, Bregetova NG (Eds.). Classifi­ (Gamasoidea). Moscow-Leningrad, Izdatel­ cation of soil inhabitant mites: I. Sarcopti­ stvo "Nauka" Sci 1956 (In series: Classifica­ formes; Ill. Mesostigmata. Leningrad: e to Dr. tion of USSR fauna, 61) (in Russian) Izdatelstvo "Nauka", 1975, 1977 (in Rus­ sian) mov and Butenko OM. Rhinonyssid mites of nonpas­ Institute, serine birds of the USSR. Moscow: Izdatel­ Humair PF, Wallich R, Gem L. 1996. :es, St.­ stvo Moscow St Univ, 1984 (in Russian) Reservoir competence of blackbirds (Turdus r and Dr. merula) for the Lyme disease spirochetes. Clayton DH, Tompkins DM. Comparative Research VII Int Cong Lyme Borreliosis, San Fran­ effects of mites and lice on the reproductive cisco, California. USA, 1996: A037 Oxford, success of rock doves (Calumba livia). and im­ Parasitology 1995; 110: 195-206 Kethley J. Acariformes. In: Parker SP, ed. m of the Synopsis and classification of living organ­ Chunikhin SP. Role of birds in the tick­ isms. New York: McGraw-Hill, Inc. 1982; 2: borne encephalitis foci treated by acaricides. 117-46 Moscow: A vtoreferat Kand Diss, 1966 (in Russian) Krantz GW. A manual of acarology. Second ections of Edition. Corvallis: Oregon State University cl-sucking Dubinin WB. Feather mites (Analgesoidea). Book Stores, Inc; 1978 e specific I. Introduction in the study; 11. Epidermopti­ Kulagin SM, Tarasevich IV. Tsutsugamushi ; agents. dae and Freyanidae families; Ill. Pterolichi­ fever. Moscow: Izdatelsto "Medicina", 1972 sian) dae family. Moscow-Leningrad: Izdatelstvo (in Russian) "Nauka", 1951, 1953, 1957 (In series: The ova LP et USSR fauna. Paukoobraznii. T. 6, N 5, 6, 7) 1 identifi­ (in Russian) Ixodidae) 8 Bull Sc1 Lvov DK, niychev GD. Bird migrations and Zemskaya AA. Parasitic gamasid mites and disease agents transmission. Moscow: their medical importance. Moscow: Izdatel­ Izdatelstvo "Nauka", 1979 (in Russian) stvo "Medicina", 1973 (in Russian)

Luttrell MP, Creekmore LH, Mertins JW. Avian tick paralysis caused by Ixodes brun­ neus in the Southeastern United States. J POP Wildl Dis 1996; 32: 133-6

Miyamoto K, Nakao M, Fujita H, Sato F. The ixodid ticks on migratory birds in Japan and the isolation of Lyme disease spirochetes from bird-feeding ticks. Jpn J Sanit Zool 1993; 44: 315-26

Nemova NV, Alekseev AN, Kostyukov MA Fashi< et al. Experimental study of Gissar virus We Argas (Bunyaviridae) reproduction in 1960'� persicus ticks. Med Parazitol, Moscow 1990; "The 1 1: 35-6 (in Russian) come! Nicholls TH, Callister SM. Lyme disease Warm spirochetes in ticks collected from birds in enviro midwestern United States. J Med Entomo1 faced" 1996; 33: 379-84 exagg( O'Connor BM. Astigmata. In: Parker SP, ed. climat1 Synopsis and classification of living organ­ researc isms. New York: McGraw-Hill, Inc. 1982; 2: energe 146-69 who aJ Olsen B, Jaenson TOT, Bergstrom S. cost o Prevalence of Borrelia burgdorferi sensu overwl lata-infected ticks on migrating birds. App which Environ Microbiol 1995; 61: 3082-7 would Randol ph SE, Gem l, Nuttall PA. eo­ feeding ticks: epidemiological significance Fashh for tick-borne pathogen transmission. Thc Parasitol Today 1996; 12: 472-9 of fasl Van Dam AP, Kuiper H, Vos K, Widjo­ affecti jokusumo A et al. Different genospecies of argue, Borrelia burgdorferi are associated with In: distinct clinical manifestation of Lyme monog borreliosis. Clin Infect Dis 1993; 17: 708-17 Manse Volgin VI. Mites of the family Cheyletidae doubt of the world. Leningrad: Izdatelstvo ofhun "Nauka", 1969 (In series: Classification of the so USSR fauna, 101) (in Russian) meclici Bull Scand Soc Parasito/ 1998; 8 (2): 9- 14 mites and v: Izdatel­ )

PARASITES AS REGULATORS OF NATURAL POPULATIONS: FASHION AS A CONFOUNDING VARIABLE

Richard W. Ashford

Liverpool School of TropicalMedicine, Pembroke Place, LiverpoolL3 5QA, UK

Fashion in science We have come a long way since, in the monograph as providing a sound 1960's, we were repeatedly warned that biological record for when the subject "The Ice Age cometh". In fact, we have would be taken over completely by come half-circle to the extent that "Global biochemists and mathematical tpodellers. Warming" is said to be "the greatest Apart from the facts th�t some environmental threat with which we are biochemists now call themselves faced". The fr rst of these fashionable molecular biologists, modellers call exaggerations was promoted by a themselves ecologists, and both are still in climatologist who wanted to establish a the ascendancy, nothing much has research institute. The second is changed in 66 years.) Hookworms are as energetically advocated by politicians abundant todayas they ever were, except who are anxious to divert attention, at the in Europe and USA, yet they are no cost of a few millions, away from the longer perceived as a major problem. overwhelming real environmental issues What has changed? Fashion! Hookworms which actually face us today, and which have gone out of fashion. would cost untold millions to deal with. Today it's malaria which is "the world's most important infectious Fashion in parasitology disease", though for Croll (1973), Though these examples of the effects malaria had ceded fr rst place to of fashion are more dramatic than those schistosomosis. This was in the brief affecting parasitology, our subject, I period between the euphoria of malaria argue, is not immune to fashion. eradication and the discovery of circum­ In a review of Clayton Lane's (1932) sporozoite antigens as candidate vaccines. monograph on bookworms Philip Malaria is, of course, promoted by Manson-Bahr opened with: "Without organisations which need high-proflle doubt bookworms are the greatest source projects to justify their existence. In the of human misery". This, remember, from same way that the Rockefeller the son-in-law of the 'father of tropical Foundation needed bookworms, WHO medicine'. (By the way, Lane justifies his and others need malaria. Desowitz 10

(1991), in "The Malaria Capers", is interpretation and control of a parasitic The particularly eloquent on this point. disease. two m: Molecules are also in fashion, so an hand, enormous slice of malaria money goes Fashion in parasites and host Anders towards technological innovation while, populations develo] in actual fact, the malaria problem is So malaria is today's fashion in dynam almost wholly socio-economic. Malaria is human parasitology, but what of fashion 1977), generally easily diagnosed, easily, quickly in the study of the effects of parasites on adv ane and cheaply treated, very rarely fatal natural populations of hosts? field st (except to visitors - hence our heightened In the early days of ecology parasites having perception of it as a problem) ; its and disease were regarded as among the one o1 epidemiology is well understood, and it is potentially important factors limiting altruis1 generally easily controlled. I've had population size: parasites were the otJ malaria three times and influenza is much fashionable in ecology from the outset. questic more unpleasant. Comprehensive studies such as those questic It cannot be denied that malaria does of Findlay & Middleton (1932) on & Zu1 sometimes cause serious intractable parasites as determinants of cyclic vole correla disease, but not nearly as frequently as populations were at first very promising, with tl1 we are led to believe by the figures of with the discovery of the brain parasite, ThE one, sometimes two, sometimes three M organism later known as Toxoplasma Anden million deaths it is supposed to cause microti, then Frenkelia microti. More sions, annually. The malaria PR machine rarely detailed studies, however, failed to import reminds us that the vast majority of these demonstrate significant effects on is imp cases are easily avoidable nor that only individuals, and certainly showed no started one of the four malaria parasites regularly measurable effect on populations (Elton, fashior causes fatal disease, that nearly all deaths 1942). Similarly, early suggestions evoluti are in Africa, or that the problem is most (Seligman & Sambon, 1907) that Scottish ThE serious in areas were the disease is red grouse Lagopus scoticus populations largely epidemic, seasonal, or sporadic. For were regulated by Leucocytozoon lovati flood I villagers living in holoendemic areas, who could not be substantiated. paras it comprise the majority of cases, malaria is These and other examples led Lack popula just one among numerous problems. (1954) among others to propose that to tes To some degree our fe ar of malaria parasites have little if any effect on selecti< was determined 100 years ago, when populations. Fashion turned against interes Ross himself, desperately campaigning parasites as ecological regulators and for individ for support in his search for the life cycle, some 25 years it became difficult to find an evO: 'confirmed' that the deadly kala-azar, more than a page or two on parasites in now known to be quite a different disease, ecological text books. Some visceral leishmaniosis, was a form of Brilliant writers such as Baer (1952) popuh malaria (Ross, 1899). and Esch (1977) kept alive the study of Chl Malaria provides an excellent example parasite ecology in its own right, but this import of how fashion rather than reason has was rarely related to host population popula guided our approach to the study, dynamics or to general ecology. Scottis 11

parasitic Then in the late '70s and early '80s Grouse Commission (Lovat, 1911), it two major forces came to bear. On one was claimed that grouse populations are hand, the intellectual duo of May and regulated by the nematode Tricho­ d host Anderson translated the recently strongylus tenuis. Krebs (1978), in his developed models of insect population excellent resume of grouse population shion in dynamics into parasite models (Anderson, theories barely mentions parasites: ,f fa shion 1977), opening the doors both for populations were supposed to be ·asites on advanced video-games, and for focu sed regulated by the interacting effects of field studies. On the other hand Hamilton, food supply and territory size. More parasites having made his major contribution to recently, as shown by Hudson (1998) at mong the one of the great Darwinian anomalies, this meeting , parasites have again limiting altruism, fixed his considerable mind on become important. Even if we accept that were the other, sexual selection. Using very grouse populations are regulated by mtset. questionable methods and even more parasites, grouse live at grossly inflated as those questionable data he and Zuk (Hamilton population densities in a semi-natural 932) on & Zuk, 1982) actually came up with a habitat managed specifically to encourage clic vole correlation which, surprisingly, agreed them. On a well-managed grouse-moor romising, with their prediction. the only predation is by shooting (game­ parasite, The intrinsic value of May and keepers, encouraged by landowl)ers, are eoplasma Anderson's early parasitological excur­ notorious in their flouting of wildlife ti. More sions, or Hamilton's bizarre ideas is not legislation, and remove natural pre­ ailed to important to the present argument. What dators), food is maintained at artificially :cts on is important is that, between them, they high levels by management, and nesting lWed no started a fashion. Parasites became sites are only limited by territory size. s (Elton, fashionable both in ecology and Grouse scarcely provide a good test of �gestions evolutionary biology. the effects of parasites on natural Scottish The new epidemiological field studies populations. �ulations largely stimulated by Anderson led to a If parasites really were important in m lovati flood of excellent data on dynamics of regulating populations, it would be parasite populations in relation to host expected that systematic natural changes ed Lack populations. Other field studies designed in bird distribution or numbers would ose that to test Hamilton's ideas on sexual correlate with parasitological changes. A ffect on selection have led to a reawakening of few examples of recent systematic against interest in the effects of parasites on changes in bird populations include the ; and for individuals, populations and species, from invasion of Europe by the collared dove t to find an evolutionary perspective. Streptopelia decaocto, the decline in the asites in British Isles of the house sparrow Passer Some comments on parasites in bird domesticus, tree sparrow Passer r (1952) populations montanus, song thrush Turdus study of Changing fashion regarding the philomelos, skylark Alauda arvensis. and but this importance of parasites in bird the increase in siskin Carduelis spinus. lpulation populations is reflected in studies on the None of these changes is fully Scottish red grouse. In the report of the understood, but changing habitat and 12

food supply are thought to be responsible: reproductive excess is frequently largely of the t there is no reason to suppose that stochastic (weather etc .) or due to we ha parasites have had any influence. starvation, lack of habitat, or predation. candid It is quite clear that parasites may Some of these factors are not density­ with e< under certain circumstances have a dependent, so are additive. Others are at sor disastrous effect on individuals and even, density-dependent, so are not additive, nations more rarely, on populations. Good but can replace one another. I suggest, it parasit examples are with malaria and is only when all these parameters are nisus trypanosomosis in Europeans and cattle ineffective that parasites become limiting. goshav trying to invade Africa or, possibly, the Obviously, with r-selected hosts in Ashfor Hawaiian honeycreeper story (Warner, unpredictable habitats, where there is a (Ashfo 1968). The effect of parasites on founder great reproductive excess, . many studies populations, either by protecting or individuals can be killed without reducing urbica. eliminating the indigenous competitors is the subsequent numbers of breeders. With owl S undeniable. I have called this the Wells K-selected species in stable habitats , on rodent� effect: the protection of populations by the other hand, loss of even a few parasit their parasites (Ashford, 1997). The individuals may cause serious reduction we fou effect of parasites on host individuals is in subsequent density. Presumably, that t equally undeniable, though I would therefore, parasites can 'afford' to be sufficit suggest that the broad assumption that more pathogenic in r-selected than in K­ on pop any infection is harmful is a gross selected hosts. As Moreau (1966) Ont oversimplification. It is arguable that a pointed out long before the K-r spectrum have f� light hookworm infection for example, had been described, tropical passerines variabl may even be beneficial in maintaining are astonishingly K-selected, so they perhap normal homeostatic balances. should be excellent subjects for found 1 The most obvious instance where the comparative study of any effects of was m factors which normally limit populations parasites on populations. larger are inactive is in domestic birds or other infecti< animals The apparent importance of Field studies Howev parasitic disease in domestic birds What we are interested to know, then, the ne compared with natural populations is not so much the effect of parasites on infecti< (Bennett et al, 1991) is surely partially the mystical figure, Ro, which indicates birds 1 explained by the careful management of the rate of increase of a population, but infecti< their habitat such that population which factors limit the final, or be bee: densities can be increased by orders of equilibrium population density, that is, lost the magnitude. which factors determine the carrying In All species produce young in excess of capacity of the environment. (unput those required to maintain a stable We have, over the past few years, obesus population close to the carrying capacity, attempted to find population studies on looked or equilibrium level of the habitat. It is hosts, which were sufficiently detailed to tapewe only if parasites cause a population to be allow the grafting of a parasite study to give a • cropped excessively that they will reduce see if, indeed, parasites are important. produc carrying capacity. The removal of the Birds, of course, are the subjects of some 13 ltly largely of the best population studies but, to date, each sampling session, and superimposed r due to we have been unable to find a good infection rates. Bartonella infection was predation. candidate. We have looked, in association found to increase in prevalence with age, )t density­ with ecological studies of varying depth, as was Nuttalia meri and an unidentified )thers are at some twenty host-parasite combi­ spirochete. Only the last parasite, which t additive, nations . These studies cover haemo­ is the rarest, is absent fr om the oldest suggest, it parasites in birds: sparrowhawk Accipiter animals, indicating that the oldest animals neters are nisus (Ashford et al, 1990, 1991) are cured or that the infected ones are e limiting. goshawk Accipiter gentilis (Toyne & dead. There was no evidence of I hosts in Ashford, 1997), swift Apus apus pathogenicity in the commoner parasites, there is a (Ashford et al, 1994), and unpublished but without precise measures of age, the ;, many studies on the house martin Delichon results would have been very misleading .. 1t reducing urbica, dipper Cinclus cinclus and tawny With the tapeworms, there was close ders. With owl Strix aluco; tapeworm larvae in agreement between parasite load and age :tbitats, on rodents, and both blood and intestinal which, since older animals are heavier, �n a few parasites of gerbils. In no instance have gives the misleading impression that body reduction we found any of the predicted indicators weight is increased by infection. ·esumably, that the parasites affect individuals rd' to be sufficiently to have any measurable effect References than in K­ on populations. Anderson RM. The regulation of host lU (1966) One of the most serious problems we population growth by parasites. Parasitology ·spectrum have faced is that of age as a confounding 1978; 76: 119-58 passerines variable (even more than fashion, Ashford RW. The leishmaniases as model so they perhaps!). In our sparrowhawk studies we zoonoses. Ann Trop Med Parasitol 1997; 91: found that Leucocytozoon toddi infection iects for 693-701 �ffects of was much higher in small broods than in larger ones indicating, possibly, that Ashford RW, Cezilly F. Hafner H, Gory G. Scarcity of haematozoa in some colonial infection of adults reduced brood size. birds in southern France. Ecologija 1994: However, since most infection occurs at 33-5 now, then, the nest and birds gradually lose their trasites on infection as they age and, further, older Ashford RW, Green EE, Holmes PR, Lucas AJ. Leucocytozoon toddi in British 1 indicates birds produce larger broods, the lower sparrowhawks Accipiter nisus: patterns of lation, but infection rate in larger broods may simply infection in nestlings. J Nat Hist 1991; 25: final, or be because the parents are older and have 269-77 r, that is, lost their infection. carrying In our recent much more detailed Ashford RW, Wyllie I, Newton I. (unpublished) studies on Psammomys Leucocytozoon toddi in British sparrowhawks Accipiter nisus: observations ew years, obesus, the fat sand rat, in Tunisia, we on the dynamics of infection. J Nat Hist ;tudies on looked at three blood parasites and a 1990; 24: 1101-7 jetailed to tapeworm. Using the eye-lens weight to � study to give a reliable relative measure of age, we Baer JG. Ecology of animal parasites. Urbana: University of Illinois Press, 1952 important. produced precise population pyramids for ts of some 14

Bennett GF, Peirce MH, Ashford RW. Avim1 Southwood, TRE. Ecological Methods. Bull Scm haematozoa: mortality and pathogenicity. J London: Chapman and Hall, 1966 Nat Hist 1991; 27: 993-1001 Toyne EP, Ashford RW. Blood parasites of Croll NA. Parasitism and other associations. nestling goshawks. J Raptor Res 1997; 31: London: Pitman, 1973 81-3 LON Desowitz RS. The malaria capers. New Warner RE. The role of introduced diseases York: WW Norton, 1991 in the extinction of the endemic Hawaiian avifauna. Condor 1968; 70: 101-20 Elton C Voles mice and lemmings, problems in population dynamics. Oxford: Clarendon Press, 1942

Esch GF, ed. Regulation of parasite populations. New York: Academic Press, lnstitu 1977

Findlay GM, Middleton AD. Epidemic disease among voles, with special reference to Toxoplasma. J Anim Ecol 1932; 3: 150-60 Spre Hamilton WD, Zuk M. Heritable true fitness classific and bright birds: a role for parasites? the in Science 1982; 384-7 investig;

Hudson PJ. The consequences of parasite studies < induced effects on the population dynamics Gerhard of birds. Bull Scand Soc Parasitol 1998; 8: 1998, I 18-22 different enigmati Krebs, CJ. Ecology The experimental Henry, analysis of distribution and abundance. New York: Harper International, 1978 occur m in the Lack D. The natural regulation of animal introduc populations. Oxford: Clarendon Press, 1954 system, Lane C. Hookworm infections. London: as a tool 1932 also rev Lovat Lord (ed.). The grouse in health and Contrac disease. London: 1911 microce, parasite, Moreau RE. The bird faunas of Africa and its islands. New York: Academic Press, 1966 Recent electropl Ross R. Report on the nature of kala azar. addresse Calcutta: Government of lndia, 1899 group Seligman CG, Sambon LW. Preliminary Fagerho note on a Leucocytozoon found in the blood Cauc of the red grouse Lagopus scoticus. Lancet tradition 1907; 173: 829-30 delimitin Methods. Bull Scand Soc Parasitol 1998; 8 (2): 15-17

1arasites of 1997; 31 : LONG-STANDING ASSOCIATIONS BETWEEN BIRD HOSTS �diseases AND NEMATODE PARASITES AS MANIFESTED IN . Hawaiian CONSERVATIVE STRUCTURAL FEATURES IN SOME ::> ASCARIDOID NEMATODES

Hans-Peter Fagerholm

Institute of Parasitology, Department of Biology, Abo Akademi University, BioCity, 20520 Abo, Finland

Sprent (1983), review ing the recognizes an ordered distribution of the classification of ascaridoids, ap tl y attributed sense organs into distinct ', groups the initiation of critical systematic (Fagerholm, 1991) it appears • that the investigations on this nematode group to the numerous species of Contracaecum from studies ofHartwich (1954, 1964, 1974). Dr. birds, al though occurring in different host Gerhard Hartw ich, who died in January species and regions of the world, are 1998, made important investigations on structurally isolated from species occurring different ascaridoids, incl uding the in other host categories. enigmatic genus Contracaecum Railliet and How ever, it has been noted that this Henry, 1912. The species of this genus apparent strict pattern is broken by the occur mainly in bird- and mammalian hosts occurrence of one species, C. ogmorhini in the aquatic environment. Hartwich Johnston & Maw son, 1941, in otariid seals. introduced the use of the "excretory" This parasite, which has been show n to be system, as ob served by JagerskiOld (1893), found off al l continents mainly in the as a tool for classification. Hartwich (1964) Southern Hemisphere, (also the Pacific also revised the cl assificat ion of species of coas t of S. Am: new data) is structurally Contracaecum and established C. similar to the species frombi rds (Fagerholm microcephalum (Rudolphi, 1809), a bird & Gibson, 1987). It is thus likely that the parasite, as the type species of the genus. species has been secondarily adapted to Recent studies using isozyme otariid seal s from a bird host, and is not a electrophoresis and structu ral data have "primary exploitation" sensu Sprent (1986) addres sed the concept of species within the (although mammalian origin the species group (See Paggi . and Bullini ,1994; occurring in birds can not be excluted). The Fager holm 1988, 1991). hosts involved (only otariid seal s) could Caudal structu res of the male have indicate that this is a case of host­ traditionally been show n to be important in succession-extension, seals feeding on bird delimiting species within this genus. If one prey, although fish vectors co uld also have 16 been directly involved (host-range References Jagerskio Nematod expansion), although the latter alternative Fagerholm, H-P. Patterns of caudal papillae has sofar not been substantiated (Still to be in Contracaecum osculatum (Nematoda) and Olafsdott analysed is the systematic position of the some related species from different regions (Nemato< bird parasite C. multipapillatum (Drasche, of the world. Int J Parasitol 1988; 18: 1039- (Halicho, 1882) in which the proximal sense organs in 51 1997; 22: the male may reach posterior to the cloaca Fagerholm H-P. Systematic implications of Paggi P, (Fagerholm 1988), a feature, as a rule, not caudal morphology in ascaridoid nematode anisakids occurring in species found in birds). Host­ parasites. Syst Parasitol 1991; 19: 215-29 25-39 range-expansion of anisakids between Fagerholm H-P. Nematode parasites of Sprent J different animal phyla is a rare phenomenon marine- and shore birds, and their role as of ascar in spite of existing trophic links. Instead, pathogens. Bull Scand SocParasitol 1996; 6: species tend generally to be host specific as 16-30 is exemplified in a recent study by Fagerholm H-P, Gibson Dl. A redescription Olafsdottir and Hauksson (1997) on of the pinniped parasite Contracaecum parasites of the gray seal .. Althoug rather ogomorhini (Nematoda, Ascaridoidea), with host specific, detrimental effect on an assessment of its antiboreal circumpolar individual birds by these often very common distribution. Zool Scr 1987; 16:19-24 parasites can be demonstrated (e.g. in C. Fagerholm H-P, Overstreet RM, Humphery­ Bull Scan magnipapillatum Chapin, 1925; see Smith I. Contracaecum magnipapillatum Fagerholm et al . 1996). However, a (Nematoda, Ascaridoidea): Resurrection and potential to regulate host populations pathogenic effects of a common parasite remains to be shown (Fagerholm, 1996). from the proventriculus of Anous minutus The present observations are supportive from the Great Barrier Reef, with a note on F of a hypothesis of a long-standing eo­ C. variegatum. Helminthologia 1996; 33: evolutionary process of birds and the 195-207 parasites of the genus Contracaecum. This Hartwich G. Die Vorderdannstrukturen, das can be inferred from the specific structural Excretionssystem sovi der Kopfbau der caudal features which are different from Ascariden und ihre taxonomishe Bedeutung. those found in species from phocid seals .. Wissenshaftliche Zeitschrifl, Martin- Luther The occurrence of a "bird-type" Contra­ Universitat. Halle Wittenbcrg 1954; 3: 1171- caecum-species in otariid seals represents a 212 Field rare exception. Specific cladistic studies to Hartwich G. Revision dcr vogclparasitischen catcher suggest early events in the evolution of both Nematoden Mitteleuropas-II. Die Gattung trade-off: the hosts and their parasites are needed Contracaecum Raillict & Henry, 1912 parasite ; Homologous structural fe atures also in (Ascaridoidea). Mittcil 7-A X)I Museum Berlin ons may ascaridoid species of other families are 1964; 40: 15-53 bility exi suggested to reveal exisisting morpho­ Hartwich G. Keys to tlle genera of tive effOI clades. However, much additional basic Ascaridoidea. In:Anderson, R.C., Chabaud, infectioru descriptive work of numerous nematode A.G. and Wilmolt, S. (Eds) CUI keys to the collared species is needed to facilitate such studies ncm,atodes of vertebrates. Farnham Royal, (Fagerholm, 1991). Commonwealth AgricuHmal Bure

Jagerskiold AA. Beitrage zur Kenntniss der nematode systematics. A. R Stone, H. M. Platt Nematoden . Zool Jabrb, Anat 1893; 7: 499-532 an d L. F. Kbalil (eds.). Systematic Association, u papillae Special Volume. London, Academic Press, ltoda) and Olafsdottir D, Hauksson E. Anisakid U.K. 22: 283-90. 1983 nt regions (Nematoda) infestations in Icelandic Grey Seals 18: 1039- (Halichoerus grypus Fabr.) J Nortbw Fish Sci Sprent JFA. Parasitological reminiscences: 1997; 22: 259-69 An address delivered to the members of the Australian Society for Parasitology on 18 cations of Paggi P, Bullini L. Molecular taxonomy in May 1982. In: Cremin M, Dobson C, Moor­ nematode anisakids. Bull Scand Soc Parasitol 1994; 4: house E, ed. Parasite Lives. Papers on para­ 215-29 25-39 sites, their hosts and their associations. To :asites of Sprent J F A. Observations on the systematics honour J. F. A. Sprent. London . Univ. ir role as of ascaridoid nematodes. In Concepts in Queensl. Press. 1986: 3-11 )1 1996; 6:

lescription �racaecum dea), with rcumpolar 24

lumphery­ Bull Scand Soc Parasitol 1998; 8 (2): 17 mp illatum �tion and 1 parasite s minutus PARASITES IN RELATION TO LIFE HISTORY AND a note on 1996; 33: SECONDARY SEXUAL TRAITS

turen, das Lars Gustafsson 1fbau der Uppsala University, Uppsala, Sweden 3edeutung. n- Luther ; 3: 1171-

Field experiments on the collared fly­ effects on immune functions and resulted in rasitischen catcher have demoll$trated a number of increased intensity of haematozoan parasites Gattung trade-offs between life history traits. Anti­ at the end of the breeding season and increa­ ry, 1912 parasite adaptations such as immune functi­ sed prevalence the subsequent breeding urn Berlin ons may also be costly. Therefore the possi­ season. Secondary sexual traits may also be bility exists of trade-offs between reproduc­ costly. In male collared flycatcher the size :enera of tive effort and the ability to resist parasitic of such a character also relates to immune Cbabaud, infections. Brood size manipulations of function and degree of parasitism :eys to the collared flycatcher fe males had negative un Royal, ux, No 2., EM U.. R� M ATUKOGt r Bull Scand Soc Parasitol l998; 8 (2): 18-22 on the of birds avian p• This THE CONSEQUENCES OF PARASITE INDUCED EFFECTS number tionshi� ON THEPOPULATION DYNAMICS OF BIRDS papers I vi duals Peter J. Hudson Specifi' are the Unit of Wildlife Epidemiology, Institute of Biological Sciences, University of Stirling, Stirling FK9 4LA, UK effects examim minor q 1. Do Historically, the discipline of popula­ hosts could withstand. As such, ecolo­ dan tion ecology has had an interesting rela­ gists believed parasites and their hosts 2. Do tionship with that of parasitology and had evolved a balance with parasites 3. Do nowhere has this been better illustrated reducing their virulence and the host de­ I answe than in the relationship between studies of veloping defence mechanisms to maintain ise that birds and their parasites. While most a balance. experim birds carry parasites, and there have been In a series of papers, Anderson and ductive many studies that have listed the parasites May (1978, 1991) synthesised the disci­ ease dy of birds, relatively few have examined plines of population biology and parasi­ sider bo what impact parasites have on the bird's tology and demonstrated that parasites sites an population dynamics. Indeed, not until may operate in a density-dependent man­ tively m recently have field and experimental ner and are thus quite capable of regu­ on game studies been undertaken which allow us to lating a host population. In particular, test the statement made by David Lack, they identified the conditions when we Parasit4 more than forty years ago (Lack, 1954), could expect parasites to stabilise a host Regl.i that "While further evidence is needed, it population and when the parasite could arise wl seems unlikely that disease is an impor­ destabilise host numbers to an extent that increase tant factor regulating the numbers of most could lead to cyclic or even chaotic host feC' wild birds". changes in host abundance. Exploration density The failure of ecologists to study the of the characteristics of parasites indi­ and an impact of parasites on the demography of cated that attenuated virulence is not which iJ bird populations was principally because necessarily an advantage since this could host abl.i parasites were seen as benign symbionts compromise transmission, so we should of Ande that would have little impact on the host expect virulence to he at a level that induced population. Ecological texts, such as maximises transmission. As a conse­ aggrega1 Ricklefs (1973), considered parasites to quence of these plenary studies, the pat­ within t be a special form of predator that did not terns and processes 1 lf several parasite­ host nur lead to the death of its prey but one that host systems have hccn examined to de­ developr adjusted its food supply to a level the termine whether parasites have an impact fecundit: 19

on the survival and breeding production parasites within the host population will of birds and thus have a consequence for tend to destabilise the dynamics (May & avian population dynamics. Anderson, 1978). The outcome for any This paper will attempt to examine a system, given the constraints of other 'ECTS number of fe atures of parasite-bird rela­ ecological processes, will then be a con­ tionships at the population level. Previous sequence of the tension between these papers have examined the impact on indi­ various components. viduals (Hudson & Dobson, 1995, 1997). Demonstrating regulation at the Specifically, I address the question "What population level is not a simple task, and are the consequences of parasite induced the only real approach is to manipulate, ng, Stirling effects at the population level?" and I in an experimentally controlled manner examine this question by addressing three the relative abundance of host or parasite minor questions: and quantitatively predict changes in host 1. Do parasites regulate host abun- abundance. Such an approach is rela­ eh, ecolo­ dance? tively simple when you can predict when :heir hosts 2. Do parasites cause population sinks? and where parasites are going to have a parasites 3. Do parasites exclude host species? large impact on their host abundance and te host de­ I answer these questions using the prem­ if experiments can be replicated both o maintain ise that a combination of monitoring, within and between populations. In this experimentation and modelling is a pro­ respect, investigating the role of parasites lerson and ductive approach to understanding dis­ in cyclic species can be productive since l the disci­ ease dynamics. Within this paper I con­ we can expect the parasites to have a .nd parasi­ sider both macroparasites and micropara­ large impact during the decline phase of : parasites sites and make no apologies for selec­ cyclic fluctuations and we can replicate 1dent man­ tively using examples from my own work the experimental removal of parasites e of regu­ on gamebirds to explore these questions. within and between populations just prior particular, to the cyclic crashes. when we Parasites and hostre gulation Detailed monitoring of grouse demog­ lise a host Regulation of host abundance will raphy coupled with field experiments and tsite could arise when there is a density dependent mathematical modelling have demon­ extent that increase in parasite .induced reduction in strated that the parasitic nematode �n chaotic host fecundity or survival. Increased host Trichostrongylus tenuis reduces the �xploration density leads to higher transmission rates fecundity and survival of red grouse and 1sites indi­ and an increased impact of parasites these effects are sufficient to cause the tce is not which if sufficiently large may regulate cyclic fluctuations observed in red grouse this could host abundance. We know from the work (Hudson et al, 1992, Hudson, 1992, we should of Anderson & May (1978) that parasite Dobson & Hudson, 1992). Experimental level that induced reduction in host survival and an treatment of a large proportion of grouse a conse- aggregated distribution of parasites in 6 populations has prevented popula­ s, the pat- within the host population will stabilise tions of grouse undergoing cyclic declines 1 parasite­ host numbers. In contrast, time delays in in abundance and demonstrated that para­ ned to de­ development, parasite reduction in host sites are not only sufficient but are neces­ an impact fecundity and a random distribution of sary cause of population cycles of red 20

a seco grouse (Hudson & Dobson, 1997, Hud­ ing population sinks. Such patterns con­ for tht son et a!, in ms). trasts with the areas without louping-ill borne where winter losses are consistently ers; se Parasites and population sinks higher than breeding losses (Hudson et al, sponse Directly transmitted microparasites 1995, 1998). Levels of infection with cation that induce a lifelong immune response in louping- ill are independent of grouse no w their host may die out in small popula­ density and consequently will not regulate Howe' tions after the number of susceptible indi­ grouse populations and disease could the vc viduals in the host population falls and wipe out in local pockets . These can be popula any remaining susceptibles are protected considered true population sinks, not a sist. p, fr om infection through herd immunity. pseudo-sink where immigrants artificially tern, th Persistence depends on the influx of more increase the population above a density not an susceptible individuals, recruited to the dependent regulated density. import population either by birth or immigration pathog fr om uninfected areas. Indeed, virulent Parasite mediated competition ticks pathogens can persist within a spatially Competition, mediated through a persist structured metapopulation when hosts shared natural enemy, is known as cus) ( di sperse between sub-populations, even "apparent competition" and could be an compel when coexistence would not occur at the important force determining community effects local population scale. The pattern of structure and biodiversity. Parasite medi­ may nt disease flow through the metapopulation ated competition is particularly signifi­ vims. ' will depend specifically on the type of cant, given the rise in emerging diseases petitior system. Directly transmitted diseases may and the opportunity that pathogens have of the spread tl1rough the host population in a to reduce host abundance. would series of waves but tick borne diseases Ticks and other invertebrate vectors are relatively sedentary since ticks are take their blood meal from a range of Conclt restricted to specific habitats and can vertebrate hosts and provide an opportune Det survive for a year or more in the absence route for between-species transmission. vi duals of hosts. In this respect, tick borne dis­ The bacterium Borrelia burgdoiferi, that can ree ease may generate "host population causes lyme borreliosis in humans, has ducti on sinks" by significantly reducing a local increased dramatically in North America field eJ group of hosts, leaving a suitable and as tick numbers have multiplied following strate t empty habitat for immigrants, which in the expansion of white-tailed deer in popula! turn subsequently become infected and rough scrub. The pathogen normally cir­ studies die. Such areas are identified by immi­ culates in populations of the white-footed the eye grants as uninhabited suitable habitat, mouse, but changes in the acorn crop of red particularly since immigration by suscep­ have altered the movement of mice and, populat tibles may occur at a time of year when together with other ecological factors, ondary ticks are not active. resulted in the increased passage of the can effi Preliminary population data fr om pathogen from mouse to tick to humans have a estates with the tick borne disease, loup­ (Jones et al, 1997). populat ing-ill indicates that the heavy mortality Interestingly, parasite-mediated com­ cies. caused by the pathogen is in effect caus- petition can occur through the presence of 21

a second species that need not be a host References erns con­ for the pathogen. For example, in tick Duping-ill Anderson RM , May RM . Regulation and borne pathogens not all hosts are amplifi­ nsistently stability of host-parasite population interac­ ers ; some produce a strong immune re­ lson et al, tions I. Regulatory processes. J Anim Ecol sponse so there is insignificant multipli­ 1978; 47: 219-47 jon with cation of the pathogen within the host and lf grouse Anderson RM , May RM . Infectious diseases no transmission fr om host to vector. t regulate of humans. Dynamics an d con trol. Oxford However, by providing a blood meal for se could University Press. 1991 the vector, they maintain the vector ;e can be AP, population and help the pathogen to per­ Dobson Hudson PJ. Regulation and (S, not a stability of a free-living host-parasite system, sist. For example, in the louping-ill sys­ lliifi allci y Trichostrongylus tenuis in red grouse. Il: tem, the mountain hare (Lepus timidus) is a density Population models. J Anim Ecol 1992; 61: not an amplifier of the virus but is an 487-98 important host for the ticks that carry the pathogen. As such the hares multiply the Hudson PJ. The effect of a parasitic nema­ tode on the breeding production of red tick'> and allow both ticks and virus to rough a grouse. J Anim Eco1 1986; 55: 85-94 persist in red grouse (Lagopus l. scoti­ nown as cus) (Hudson et al, 1995). Therefore, Hudson PJ. Grouse in Space and Time. .Jld be an competition is being mediated through the Game Conservancy Trust: Fordiqgbridge, Jmmunity 1992 effects of a pathogen even though the host site medi­ may not play any role in amplifying the Hudson PJ, Dobson AP. Host-parasite proc­ Y signifi­ virus. This is still parasite-mediated com­ esses and demographic consequences. In: : diseases petition, however, because in the absence Clayton DL, Moore J, eds. (Host Parasite �ens have of the hares the impact on the grouse Evolution) 1996. 128-54 would be reduced. e vectors Hudson PJ, Gould EA, Laurenson MK, Gaunt M, Reid HW, Jones LD, Norman R, range of Conclusion MacGuire K, Newborn D. The epidemiol­ Jpportune Detailed experimental studies of indi­ ogy of louping-ill, a tick borne viral infec­ ISmis sion. viduals have demonstrated that parasites tion of grouse and sheep. Parassitologia ife ri, that 1998; 39: 319-23 can reduce the survival and breeding pro­ nans, has duction of individual hosts. More recently Hudson PJ, Greenman JV. Parasite mediated 1 America field experiments are starting to demon­ competition . Biological and theoretical prog­ following strate that parasites have an impact at the ress. Trends in Ecology & Evolution 1998: deer in population level. Some of our recent in press mally cir­ studies have shown that parasites cause AP. �te-footed Hudson PJ, Newborn D, Dobson Regu­ the cyclic population crashed in numbers lation and stability of a free-living host­ :orn crop of red grouse, other parasites can cause parasite system, Trichostrongylus tenuis in mice and, population sinks and the presence of sec­ red grouse. 1: Monitoring and parasite re­ l factors, ondary hosts can act as a reservoir that duction experiments. J Anim Ecol 1992; 61: ge of the 477-86 can effectively eliminate a host. Parasites ) humans have an important role to play in the Hudson PJ, Norman R, Laurenson MK, population dynamics of some bird spe­ Newborn D, Gaunt M, Reid H, Gould E, Jed com­ cies. Bowers R, Dobson AP. Persistence and :esence of 22 transmission of tick-borne viruses: Ixodes Bull Se ricinus and louping-ill virus in red grouse populations. Parasitology 1995; 111, 49-58

Jones C,G. et al. Chain reactions linking acorn s to gypsy moth outbreaks and lyme AQ disease risk, Science 1998; 279: 1023-6 AJ Lack D. The natural regulation of animal numbers. Oxford University press. 1954

May RM , Anderson RM . Regulation and stability of host-parasite population interac­ tions II. Destabilizing processes. J Anim Ecol 1978; 47: 248-67

Ricklefs, R.E.. Ecology. Second Edition . University of Pennsylvania, Nelson. 1973 Tht parasit siderat helmin ments bility ' ity to helmin in fish inhosp isolati< localiti (1988) specie� aquati< sation that th' and ra indirec munitit differe1 were s larity v allogen genic c These 1 the hyt: no din present Bull Scand Soc Parasitol l998; 8 (2): 23-28

AQUATIC BIRDS AS AGENTS OF PARASITE DISPERSAL: A FIELD TEST OF THEEFFECTIVENESS OF HELMINTH COLONISATION STRATEGIES

Clive R. Kennedy

Department of Biological Sciences, University of Exeter, Exeter, EX4 4PS, UK

The distribution of freshwater fish changes in the helminth fauna of fishin a parasites is notoriously patchy, and con­ small lake over 28 years. The population siderable differences may exist in the crash and almost complete disappearance helminth fauna of neighbouring catch­ of the fish populations provided ',a unique ments (Kennedy, 1981, 1985). The mo­ opportunity to study the disappearance of bility and vagility of birds and their abil­ helminth species and their recolonisation ity to serve as definitive hosts for many of the lake, and so a field test of the rela­ helminths whose larval stages are found tive effectiveness of autogenic and allo­ in fish gives them the potential to cross genic colonisation strategies. inhospitable barriers, breakdown habitat The lake under study is Slapton Ley, a isolation and introduce new species into small (0.7 km2) shallow (maximum depth localities (Kennedy, 1981). Esch et al, 2.5m), isolated, coastal, freshwater la­ (1988) therefore suggested that allogenic goon in Devon of recent origin (c. 1000 species, and especially those maturing in yrs) (Kennedy et al, 1994: Kennedy, aquatic birds, should have greater coloni­ 1996). The lake and its catchment have sation potential than autogenic species in been the subject of investigation for many that they could be dispersed more widely years and the fish and their helminth and rapidly. They tested this hypothesis fauna have been studied regularly since indirectly by comparing helminth com­ 1970 (Kennedy, 1996). Attention has munities in selected species of fish and in focused on the helminths of roach Rutilus different localities, and found that there rutilus, rudd Scardinius erythrophthal­ were significantly higher levels of simi­ mus, perch Perca fluviatilis and pike larity within and between localities due to Esox lucius. Eel Anguilla anguilla are allogenic species when compared to auto­ also present in the lake, but studies on genic ones in cyprinid and percid species. their parasite fauna did not commence These findings were considered to support until 1993. Information on the composi­ the hypothesis, but there has hitherto been tion of the parasite fauna up to 1973 was no direct test of the hypothesis. In the summarised by Canning et al, (1973) and present paper, an account is given of the up to 1975 by Kennedy (1975). After 24

1973 samples of all species except eel aquatic bird also declined between 1985 were taken regularly, at least three times and 1990. Table a year and in some years monthly, by the Prior to 1973, 9 species of helminth author and examined for parasites by were reported from the 4 species of fish standard methcxls. Over the period the in the lake (Canning et al, 1973). Only 2 lake has become progressively more eu­ of these species were allogenic: Diplos­ Parasi trophic and by the end of the 1970s it tomum spathaceum and D. gasterostei. approached the hypereutrophic state The 7 autogenic species comprised 3 (Johnes & Wilson, 1996). During the species of monogenean, 2 species of severe winter of 1984/85 the lake froze cestode and 2 species of acanthocephalan, Dactyl over, and this produced the ideal condi­ (Table 1). Overall, the helminth fauna Neoda tions for winterkill of the fish (Kennedy, was species poor: the absence of proteo­

1996). Populations of roach, rudd and cephalid cestodes, nematodes and adult Tetrao perch especially fell to levels at which digeneans from fish and the paucity of only a few, or even single specimens, dactylogyrid species on roach and rudd were caught in the samples (Kennedy et are all believed to reflect the isolation of CaryoJ al, 1994). A nucleus of species remained the lake and the difficulty of autogenic Ligula in the region of the main inflow river to species in colonising it. Between 1975 Dip!os the lake, and this formed the basis for the and 1984 the autogenic species Acan­ subsequent re-colonisation of the lake. thocephalus lucii became locally extinct Re-colonisation by rudd was first detected and this was followed by A. clavula after D.gast in 1987, but by roach and perch not until 1987. However, Podiceps cristatus had Ty lode 1990. Pike were always catchable introduced the allogenic species Ligula throughout the period, although they intestinalis and Tylodelphys clavata into T.podi. tended to be emaciated and in poor con­ the lake, both species being recorded for dition. Eel numbers appear to have been the first time in 1973, and later a third Acanth unaffected. species, T. podicipina, in 1977 (Kennedy A. clavi The lake has always been attractive to & Burrough, 1981: Kennedy, 1981). aquatic birds: flocks of Larus ridibundus Thus by 1987 the composition of the regularly roost there, and both Phalacro­ helminth fauna had changed significantly corax carbo and Ardea cinerea are from 7 autogenic species and 2 allogenic 2.= regularly to be found fishing there. The ones pre-1973 to 5 autogenic species and expansion of the roach population in the 5 allogenic ones. early 1970s, however, attracted Podiceps Fish mortality following the 1984/85 Au =; cristatus to the lake for the first time in severe winter, and the only slightly less Ro = over 30 years. One pair bred in 1973, and severe 1985/86 winter, was catastrophic. the breeding population increased to be­ The number per "standard" day's netting tween 5 and 8 pairs in the late 1970s. fell from 91.4 in 1984 to 1.6 in 1985 and Following the crash of the fish popula­ 1.3 in 1986 (Kennedy et al, 1994). Re­ tions, numbers declined and only one covery and re-colonisation was very slow breeding pair was present in 1985, but by and it was not until 1990 that populations 1995 this had increased again to 9 or 10 again reached normal levels. pairs (Elphick, 1996). Other species of 25 veen 1985 Table 1 Changes in the composition of the helminth parasite fauna of fishin Slapton Ley before and after the host population crash f helminth ies of fish 3). Only 2 :::: Diplos­ Parasite species Year asterostei. Status Pre 1973 1977 1984 // 1990 1991 1992 1993 1994 1996 Host nprised 3 1973 :pecies of xephalan, Dactylogyrus vistulae Au p p p p A A A A A A Ro, Ru nth fauna Neodactylogyrus sp. Au p p p p A A A A A A Ro, Ru of proteo­

and adult Tetraonchus monenteron Au p p p p p p p p p p Pi Jaucity of Caryophyllaeus laticeps Au p p p A A A A A A A Ro and rudd ;olation of Caryophy/laeides fe nnica Au p p p p A A A A A A Ro, Ru

autogenic Ligula intestinalis Al A p p p p p p p p p Ro reen 1975 Diplostomum spathacewn Al p p p p p p p p p ,P Ro, Ru ies Acan­ Pe, Pi lly extinct D.gasterostei Al p p p p A p A A p A Pe zvula after

>tatus had Ty lodelphys clava/a Al A p p p p p p p p p Ro, Ru es Ligula Pe, Pi avata into T.podicipina Al A A p p A p A A p p Pi :::orded for Au p p A A A A A A A A Ee er a third Acanthocephalus lucii ' ' (Kennedy A.clavula Au p p p P * A* A A A A A Pe, Ee y, 1981). cm of the *l gnificantly = last recorded in perch in 1980 and from eels in 1987

� allogenic 2. = Eels are not included in the above table as they were not surveyed thoroughly until l995. By that time pecies and Pseudodactylogyrus spp. (Av) and Anguillicola crassus (Av) were present. Neither taxon had been recorded in samples taken in 1987 or earlier years. e 1984/85 Au = autogenic; Al =allogenic ightly less Ro = roach, Ru - rudd, Pe = perch, Pi =pike, Ee = eel. lastrophic. y's netting 1985 and 994). Re­ very slow opulations 26

When regular sampling became pro­ into the lake by P. cristatus, as were T. ductive again in 1990, it was found that podicipina and D. gasterostei. all the autogenic species with the excep­ The impact of L. intestinalis on the tion of Tetraonchus monenteron on pike fish populations has been major. It is had disappeared fr om the lake (Table 1). specific to roach in the lake, and by 1975 The most plausible explanation of this is it had reduced the roach population to a that the populations of roach, rudd and level at which intra-specific competition perch had fallen below the threshold level had declined and this in turn had relieved at which the helminth populations could the inter-specific competition pressure on survive and transmit i.e. below the break rudd, the population of which increased point, and so the parasite populations again (Kennedy & Burrough, 1981: became locally extinct. The pike popula­ Wyatt & Kennedy, 1998). Thereafter, as tion, which remained at a higher level the transmission window for L. intestina­ over the same period, was clearly large lis narrowed (Wyatt & Kennedy, 1989), enough to permit the survival of T. so the population declined until in 1984 it monenteron. To-date, none of the auto­ was barely detectable in the samples. The genic species that went extinct have re­ second cycle of infection commenced in colonised the lake. By contrast, all the 1991, and events were repeated. The allogenic species have survived or re­ expansion of the roach population after colonised (Table 2). It seems probable the population crash was checked that D. spathaceum and T. clavata, the (Kennedy et al, 1994), permitting a more most prevalent and abundant and least equitable balance between roach and rudd specific of the allogenic species, survived numbers. By 1997 the L. intestinalis in the nucleus of the fish population: both population had again fallen to barely species occurred in pike and in the few detectable levels, and the species again roach and perch caught between 1985 and appeared to be on the verge of local ex­ 1989, and both species occurred at high tinction. Thus, one allogenic species has levels of prevalence and abundance in had a profound effect on the relative 1990. By contrast, D. gasterostei and T. abundance and population dynamics of podicipina, the most specific and least the fish species (Kennedy, 1996). common of the digeneans, did not re­ At the time of writing, little seems to appear in numbers until 1994 although a have changed since 1995 and 1996. Only very few specimens of each were found in 6 species are present' in the 4 species of 1991. No roach infected with Ligula in­ fish compared to the 11 recorded before testinalis were found from 1985 to 1989 the population crash, but 5 of these spe­ inclusive (Kennedy et al, 1984) and only cies are allogenic. Only one autogenic a single specimen in an old fish was species has survived the crash: the others found in 1990. By 1991, however, infec­ disappeared and have not re-colonised. tion levels had risen to epidemic propor­ Indeed, very few autogenic species ap­ tions It seems very likely, therefore, that pear to have been able to colonise the L. intestinalis did not survive the roach lake in its 1000 years of existence, let population crash, but was re-introduced alone any re-colonise it. By contrast all 5 '"' 1-• , V M- ... - ;:> � - en 0-- g. � ...... � .... l:i ...... � (i) -::r ...... (I) q (t-- - !5 1-J.o?)• � 2. � ��a?5::ers (')�- 0 00 � (JQ 0.. :::::1 ::I c er '"' oo� :::::� e ...... 0 ...... r/) '-'' (') v,t:l � .�� - gi2 ...... 00 � (') � '""" 0.. � :::;" :::::1. (')W_. (1:) '"' 0 � (') o o � ...... § ::::P � @'g §« · � � r/) cr · :::::( ('). @...... \Ot!)v, �_.� ,.., _. ::::;:' e w r/) i]e:,.�CT' a � ...... ::I ' 9' g_, I] 0 i';J � oo .•'"' \0�� c @ oo � :::;" � ��� � oo ::i' oorn < C·o � � :... g.� c. � � ;::;-:2· 2 §'"' .J::.. \0� 0 g. a � e. ?5 � 0 e...... 0 < � :>< 1-· ...... � (P � ::r ::r ��...... (P · V\��'9• r/) (') I � >-+,'<: 0 ....., � r/) I ::I '< "'-� o.. ,.., � :::.....· � t"""+-...... � ..__, 1 oo •• o.. �..... a§ � V\ t;; � :--3

Table 2 Infection levels of selected helminth species in fish in Slapton Ley before and afterthe host population crash

Parasite species and infection level Year

1979 1981 1983 1984 If 1990 1991 1992 1993 1994 1996 a) Perch Ty lodelphys clavata Prevalence 100 98.2 100 100 100 100 100 100 100 80.0 Abundance 52.7 11.9 75.3 39.4 72.1 42.2 32.3 30.1 43.9 12.8 ±SD 23.4 9.4 37.9 13.3 41.3 37.6 29.4 17.4 16.8 11.9

Tpodicipina Prevalence 100 98.2 100 84.8 0 11.1 0 0 5.2 40.0 Abundance 7.2 8.8 6.7 2.9 0 0.05 0 0 0.05 0.4 ±SD 3.3 3.6 2.8 2.2 0 0.2 0 0 0.2 0.2

Diplostomum gasterostei Prevalence 16.9 3.5 9.1 2.0 0 18.0 0 0 10.5 0 Abundance 0.04 0.003 0.12 0.02 0 0. 18 0 0 0.23 0 ±SD 0.001 0.06 0.3 0.001 0 0.04 0 0 0.4 0 b) Roach Ligula intestinalis Prevalence 16.3 11.4 2.1 3.1 1.4 71.4 69.7 45.6 16.7 2.1 Max. intensity 2 2 1 1 1 8 21 4 1

D.spathaceum Prevalence 77. 1 88.2 N N lOO 100 100 94. 1 lOO lOO Abundance 5.9 13.3 12.9 17.1 15.5 12.0 17.0 7.9 ±SD 2.9 17.2 27.1 7.2 7.1 11.0 10.0 6.2 N -..l 28 Bull S allogenic species are still present, and 3 Kennedy CR. The fish of Slapton Ley. Fld are believed to have re-colonised since Stud 1996; 8: 685-97

1994. Allogenic species are thus clearly Kennedy CR, Burrough RJ. The establish­ the more successful colonists and the role ment and subsequent history of a population of birds as the agent of their dispersal and of Ligula intestinalis in roach Rutilus rutilus colonisation is of major importance. The (L.) J Fish Bioi 1981; 19: 105-26 et hypothesis of Esch al, (1988) is now Kennedy CR, Wyatt RJ & Starr K. The de­ supported not only by indirect evidence, cline and natural recovery of an unmanaged but also by a direct test comparing the coarse fishery in relation to changes in land effectiveness of autogenic and allogenic use and attendant eutrophication . In: Cowx colonisation strategies. IG, ed. Rehabilitation of Freshwater Fisher­ ies. Oxford: Fishing News Books 1994: 366- References 75 01 Wyatt RJ, Kennedy CR. The effects of a Canning EU, Cox FEG, Croll NA, Lyons cient change in the growth rate of roach, Rutilus KM. The natural history of Slapton Ley farnil rutilus (L.), on the population biology of the Nature Reserve: VI. Studies on the parasites. fish tapeworm Ligula intestinalis (L.). J Fish mine� Fld Stud 1973; 3: 68 1-718 Bioi 1988; 33: 45-57 ics. ! Elphick D. A review of 35 years of bird­ distril Wyatt RJ, Kennedy CR. Host-constrained ringing at Slapton Ley (1961-1995) together introc epidemiology of the fish tapeworm Ligula with a brief historical review of ornithologi­ M intestinalis (L.). J Fish Bioi 1989; 35: 215- cal observations. Fld Stud 1996; 8: 699-725 27 last 1 Esch GW, Kennedy CR, Bush AO, Aho JM. 10 m Patterns in helminth communities in fresh­ cludi1 water fish in Great Britain: alternative taiga, strategies for colonisation. Parasitology erts b 1988; 96: 1-14 R< Johnes PJ, Wilson HI\1. The limnology of were Slapton Ley. Fld Stud 1996; 8: 585-612 The f tion Kennedy CR. The natural history of Slapton Ley Nature Reserve VIII. The parasites of lead t fish, with special references to their use as a epide source of information about the aquatic community. Fld Stud 1975; 4: 177-89

Kennedy CR. Parasitocoenoses dynamics in freshwater ecosystems in Britain. Trudy Zool Inst AN USSR 1981; 108: 9-22

Kennedy CR. Interactions of fish and para­ site populations: to perpetuate or pioneer? In Rollinson D & Anderson RM , eds. Ecology and Genetics of Host-Parasite Interactions, London : Academic Press, 1985, Linn Soc Symp Series 11. 1-20 Bull Scand Soc Parasitol l998; 8 (2): 29 1 Ley. Fld

establish­ population flus rutilus BIRDS-PARASITES-VIRUSES: INTERPOPULATIONAL INTERACTIONS C. The de­ lnmanaged Dmitri Lvov �es in land In: Cowx The D.l. Ivanovsky Institute of Virology RAMS, Moscow, Russia ter Fisher- 1994: 366-

Objective: The birds are the most an­ was examined in the following models: 1) 'fe cts of a cient reservoir of hundred viruses of 14 transmitted by mosquitoes of the viruses :h, Rutilus logy of the families. High population density deter­ of Karelia fever in eastern Fennoscandia :L.). J Fish mines the development of mass epizoot­ Geta fever in eastern Siberia (Toga­ ics. Seasonal migrations promote virus viridae, Alfavirus), West Nile virus of distribution for long distances and their encephalitis in the South of Rw�.sian Plain :on strained introduction in various ecosystems. (Flaviviridae, Flavivirus); 2) the group of rm Ligula 1; 35: 215- Materials and Methods: During the viruses from Flaviviridae, Bunyaviridae, last 10 years ecological sounding of over Reoviridae families the obligated hosts of 10 million km2 of Northern Eurasia in­ which are birds and ticks Ixodes uriae; 3) cluding Arctic, Northern-Middle Southern influenza viruses. taiga, leaf-bearing forests, steppes, des­ Conclusions: For the purpose of re­ erts belts was carried out. vealing the emerging and re-emerging Results: Among isolated 69 viruses 25 infections and prognosis of epidemic were described as new for the science. situations it is necessary to carry out the The features of interpopulational interac­ monitoring in key points of Northern tion of birds-arthropods viruses which Eurasia as a part of international pro­ lead to the development of epizootics and gram. epidemics were studied. Such situation Bull Scand Soc Parasitol l998; 8 (2): 30-32 paras domir favou ratios preva DARWINIAN MEDICINE? LESSONS FROM AVIAN BLOOD will t PARASITE ECOLOGY more stanc< Andrew F. Read 1 and Dave Shutler 2 garnet (Read 1 Institute of Cell, Animal and Population Biology, University of Edinburgh, Edinburgh EH9 D� 3JT, Scotland, ln<; 2 avian Biology Department, Acadia University, Wolfville, Nova Scotia. Canada BOP lXO confo ship 1 appar The application of evolutionary biol­ is little reason to expect that optimality do no ogy to infectious disease research - what mcxlels will successfully predict evolution of ga is optimistically called Darwinian Medi­ of traits we understand even less well, tios il cine (Williams & Nesse, 1991; Nesse & such as parasite virulence. Let . us con­ a rei< Williams, 1994; Ewald, 1994) - has gen­ sider how the former theory has fared in and 0 erated much excitement, particularly predicting sex ratios of blocxl parasites of is clo: among evolutionary biologists (e.g., the protozoan suborder Haemosporina. theor) Westoby, 1994; Futuyma, 1995; Stearns, Each of the genera we consider prcxluces length 1999). In the context of infectious dis­ sexually dimorphic transmission stages parity ease, the enterprise mostly involves appli­ (gametocytes) in the peripheral blocxl of of th< cation of optimality mcxlels to pathogenic their vertebrate host. As we have argued being organisms. Optimality mcxlels normally elsewhere (Read et al, 1992, Shutler & fied. assume population dynamic equilibria, Read, in press), gametocyte sex ratios releva yet this is not an obvious feature of infec­ should be under intense natural selection. systeu tious diseases, where rich epidemiological Simple mcxlels can be used to predict a nus h dynamics are expected and frequently relationship between gametocyte preva­ for e observed (Anderson & May, 1991; lence (proportion of infectious hosts) in a readil: Nowak & May, 1994). Hence, the appli­ host population and the sex ratio that Leuco cability of optimality mcxlels to infectious should be favoured by natural selection transn disease remains to be demonstrated em­ (Hamilton, 1967; Read et al, 1992; Read ratio pirically (Read et al, 1999). et al, 1995). In populations where game­ be the Sex allocation theory, when applied to tocyte prevalence (and hence proportion birds; free-living organisms, has provided some of hosts which are infectious) is low, cyte 1 of the best evidence in favour of the opti­ transmission rates and hence frequency of sex ra mality approach to trait evolution multiple infections will be low. In such even ' (Charnov, 1982; Frank, 1990). If this circumstances, gametes contributing to few vl bcxly of theory can not be readily trans­ the mating pool within a vector blocxl garnet ferred to disease-causing organisms, there meal will likely come from the same never 31

parasite clone. Self fertilisation will unfertilised female gametes. This idea dominate, with natural selection therefore requires that Haemoproteus microgame­ favouring female-biased gametocyte sex tocytes release fewer microgametes at ratios. In contrast, where gametocyte exflagellation than do Leucocytozoon prevalence is high, multiple infections microgametocytes. Finally, Haemopro­ will be common, and hence outcrossing teus parasites in the populations consid­ more frequent. Under these circum­ ered by Shutler et al. (1995) are probably stances, natural selection will favour all vectored by midges; Leucocytozoon gametocyte sex ratios closer to unity parasites are vectored by larger-bodied (Read et al, 1995). black flies (Simuliidae). If gamete num­ tugh EH9 Data from various populations of the bers in a blood meal are more often lim­ lXO avian protozoan genus Leucocytozoon iting for Haemoproteus, sex ratios closer conform remarkably well to this relation­ to 1:1 would be favoured to ensure that ship (Read et al, 1995); data from the fertilisation occurred. apparently similar genus Haemoproteus In principle, these ideas could be )ptimality do not (Shutler et al, 1995). Irrespective readily addressed with the right data: evolution of gametocyte prevalence, mean sex ra­ • Are Haemoproteus infections more less well, tios in Haemoproteus populations fall in readily controlled than Leucocytozoon : us con­ a relatively narrow band between 0.30 infections? How do Mst age­ � fared in and 0.42. In most instances, the sex ratio prevalence proftles compare for the rasites of is closer to 1:1 than to that predicted by two genera? 10sporina. theory. Elsewhere we have considered at • Are there differences in the number of produces length possible explanations for this dis­ viable gametes per gametocyte at ex­ m stages parity (Shutler & Read, in press). Three flagellation? It may be possible to de­ blood of of these hypotheses have the merits of termine this on morphological grounds re argued being plausible and relatively easily falsi­ alone. :hutler & fied. First, the gametocyte prevalences • Are gamete numbers more often lim­ ex ratios relevant to sex ratio evolution might be iting in blood meals within which selection. systematically underestimated in the ge­ Haemoproteus fertilisation occurs? It predict a nus Haemoproteus. This could occur if, may be possible to assess this directly; te preva­ for example, Haemoproteus is more of great interest would be Haemopro­ osts) in a readily cleared by host immunity than is teus sex ratios in populations where ·atio that Leucocytozoon. This would mean that substantially larger hippoboscid flies selection transmission patterns responsible for sex are the principle vector. 92; Read ratio evolution in Haemoproteus would Finally, it would also be of consider­ :re game­ be those predominantly found in younger able interest to have sex ratios of Hae­ roportion birds; the key parameter is thus gameto­ moproteus and Leucocytozoon parasites , is low, cyte prevalence in those birds. Second, drawn from the same bird population }Uency of sex ratios close to 1:1 would be favoured (better yet, the same birds). This would In such even where selfing rates are high if rather confirm that the different sex ratios re­ mting to few viable gametes are released per male ported by Read et al. (1995) and Shutler or blood gametocyte: natural selection should et al. (1995) reflect biological differences he same never favour sex ratios which result in between the two genera. 32

There are at least three reasons why Nesse RM , Williams GC. Why We Get Sick. gathering such data would be a worth­ New York: Time Books, 1994 while exercise. First, at first glance (and Nowak MA, May RM. Superinfection and even after serious consideration), the the evolution of parasite virulence. Proc R Haemoproteus data fly in the face of Soc Loud B 1994; 255: 81-9 otherwise successful sex ratio theory; Read AF, Aaby P, Antia R, Ebert D, Ewald violations of theory may point to inter­ PW, Gupta S, Holmes EC, Sasaki A, Shields esting (and currently unrecognised) bio­ DC, Taddei F, Moxon RE. What can evolu­ logical factors which violate assumptions tionary biology contribute to understanding of the model. Second, gametocyte sex virulence? In: Steams, SC, ed. Evolution in ratio is potentially a very cheap way of Health and Disease. Oxford: Oxford Univer­ assessing population genetic structure in sity Press, 1999, in press malaria populations (Read et al, 1992); Read AF, Anwar M, Sbutler D, Nee S. Sex Ta we need to know if and when it is accu­ allocation an d population structure in ma­ fully, rate. Third, if sex allocation theory can laria and related parasitic protozoa. Proc macrc not make quantitatively successful pre­ Roy Soc B 1995; 260: 359-63 worm dictions about infectious disease agents, Read AF, Narara A, Nee S, Keymer AE, are v< the prognosis for Darwinian medicine, Day KP. Gametocyte sex ratios as indirect extrac which concerns traits such as virulence measures of outcrossing rates in malaria. often for which we know substantially less Parasitology 1992; 104: 387-95 bleedi about fitness consequences than we do in Sbutler D, Bennett GF, Mullie A. Sex pro­ Intuiti the case of sex ratio, is rather gloomy. portions of Haemoproteus blood parasites limitet Our data thus provide both a stimulus and local mate competition. Proc Natl Acad leave and a note of caution with which to tem­ Sci USA 1995; 92: 6748-52 eratioJ per the euphoric rush of optimality theo­ Sbutler D, Read AF. Local mate competi­ sites 1 rists into infectious disease biology. tion , and extraordinary and ordinary blood despit parasite sex ratios. Oikos 1998; 83: in press intew References few st Steams SC, ed. Evolution in Health and Anderson RM , May RM . Infectious Diseases Disease. Oxford; Oxford University Press, macro of Humans. Dynamics and Control. Oxford: 1999 Fn Oxford University Press, 1991 easy 1 Westoby M. Adaptive thinking in medicine. Cbarnov EL. The Theory of Sex Allocation . have 1 Tr Ecol Evol 1994; 9: 1-2 Princeton: Princeton University Press, 1982 their r Williams GC, Nesse RM . The dawn of dar­ zoans Ewald PW. Evolution of Infectious Diseases. winian medicine. Q Rev Biol 1991; 66: 1-22 samplt Oxford: Oxford University Press, 1994 sites li Frank SA. Sex allocation theory for birds by iru and mammals. Ann Rev Ecol Syst 1990; 21 : birds ; 13-55 infecti' Futuyma DJ. The uses of evolutionary biol­ mined ogy. Science 1995; 267: 41-2 cons m

Hamilton WD. Extraordinary sex ratios. Our k Science 1967; 156: 477-88 ------·------ll:ki

Bull Scand Soc Parasitol 1998; 8 (2): 33-38 � Get Sick.

3Ction and :e. Proc R MACROPARASITES AS SELECTIVE AGENTS IN BIRDS

D, Ewald Arne A, Shields Skorping can evolu­ Department of Zoology, University of Bergen, Realfagbygget, Allegaten 41, erstanding 5007 Bergen, Norway volution in 1rd Univer-

\fee S. Sex Take any kind of bird, dissect it care­ and population dynamic effects of 1re in ma­ fully, and the chances of finding macroparasites in wild birds is therefore )zoa. Proc macroparasites such as flukes, tape­ scant. The few good studies that exist worms, acanthocephalans or nematodes typically come from game birds where �ymer AE, are very high. These worms will always large samples can easily be obtained (e.g. as indirect extract resources from the host, and will Hudson, 1986). Still, birds have probably r1 malaria. often induce disease symptoms such as been challenged by endoparasitic bleeding, immunopathology or diarrhoea. helminths throughout their evolutionary

.. Sex pro­ Intuitively, in a world where resources are history, and these parasites can be found ! parasites limited and individuals are competing to in almost every organ. If we are going to Natl Acad leave most descendants in the next gen­ understand how parasites may affect eration, adaptations to resist macropara­ important aspects of bird biology, such as

3 competi- sites must be of some importance. Still, life histories, mating systems or foraging 1ary blood despite the almost epidemic rise in the behaviour, we certainly need more studies .: in press interest of parasites, there are surprisingly on macroparasites. few studies on the evolutionary effects of Iealth and macroparasites in birds. Selective pressure of macroparasites is sity Press, From a practical point of view it is density-dependent easy to understand why most workers Macroparasites are usually much medicine. have chosen other kinds of parasites as larger than microparasites and can only their models. Prevalences of blood proto­ infect the host via transmission stages. wn of dar- zoans can be estimated by taking blood The number of parasites within a bird at 1, ; 66: 1-22 samples and infection levels of ectopara­ a particular time is therefore a direct sites like mites or lice can be determined consequence of the number of successful by inspection. Most macroparasites of infections. Usually, both the negative birds are endoparasites, and the level of effect of a macroparasite on its host and infection can only be accurately deter­ the host-response to the parasite is den­ mined by killing the bird and time­ sity-dependent (Behnke & Barnard, consuming dissection in the laboratory. 1992). This means that the selective pres­ Our knowledge of both the evolutionary sure on the host to develop efficient re- 34

If : sistance mechanisms is also density­ must recently have been considerably to eva dependent. If a particular parasite is to relaxed. this tJ play any role as a selective agent on the Observed infection patterns: Exposure parasi host, the rate of exposure must therefore or susceptibility? Althot be sufficiently high over several genera­ Goater & Holmes (1997) suggested tions to affect host fitness. We do not affect three requirements that must be met if know whether this is a common phe­ menta parasites should play any role in the nomenon or not, since long-termed studies demor evolution of host traits: 1. Parasite num­ of exposure rates of macroparasites are Here, bers must vary within a host population; extremely rare. We do know however that be de 2. This variation must be associated with many macroparasites may show large SUSCeJ variation in host fitness; and, 3 . . The traits variations in infection levels over time. In factor: selected for (e.g. resistance) will have to the example given in Fig. 1, the two parasi covary with parasite numbers. The evi­ nematodes, Sp lendidofilaria papillocer­ not on dence for the two first requirements are : cus and Capillaria caudinflata, show (Fig. overwhelming; almost all macroparasites marked temporal variations in prevalence numb< are nonrandomly distributed among their in willow ptarmigan (Lagopus lagopus) tion o host individuals and in most cases where fr om an area in northern Norway during a variac fitness effects on the host have been seven year period. During the last years inclivic looked for experimentally, it has been both nematodes have been rare, and any becau: found. The third requirement is harder to selective pressure exerted during earlier place confirm. years, when they where more common, parasi have 2 para.'>i sarily 80 ences 70 11 Splendidofilaria pappilocerca 60

Cl) 11 Capillaria caudinflata (.) 50 s:: Cl) iii 40

...� ' c.. 30 ' I ' 20 1 i 10 ' c 0 1992 1993 1994 1995 1996 1997

Year

Fig. 1. Prevalence of two nematodes in willow ptarmigan from samples shot in the autumn each year in the period 1992 - 1997.

Fig. 2 from ( 35

It might be argued that although expo­ 1Siderably If a host trait, like resistance, is going to evolve, host individuals which develop sure is highly variable and that hosts may this trait must on average have lower acquire high burdens just because of bad Exposure parasite burdens than those that do not. luck, individuals that develop some re­ Although we know that many traits may sistance will still, on average, have lower suggested affect the number of parasites in experi­ parasite burdens. This is probably true, Je met if mental hosts, it is much more difficult to but given the large stochasticity in expo­ le in the demonstrate this fr om natural situations . sure, the effect of having different sus­ .site num­ Here, variations in parasite numbers will ceptibility on the realised parasite burden ::>pulation; be determined both fr om variability in may be quite small. How can we separate iated with susceptibility and by external, ecological the effect of exposure versus susceptibil­ The traits factors related to exposure. Many macro­ ity in natural populations? Experimental ll have to parasites tend to show marked variations laboratory infections are of limited value The evi­ not only in time (Fig. 1) but also in space because hosts are not subjected to natural ments are (Fig. 2). Much of the variation in parasite variability in exposure. Experimental ::>parasites numbers observed within a host popula­ field studies where hosts can be individu­ 1ong their tion could therefore be caused by highly ally tagged and recaptured, or correlative ses where variable exposure rates where some host studies between parent and offspring 1ave been individuals have high infections simply parasite loads (see for example Boulinier n has bee because they happened to be at the wrong et at, 1997) would be better, but are lim­ harder to place at the wrong time, while others are ited to parasites that can be quantified parasite fr ee because they accidentally without killing the host. The alternative is have avoided being exposed. Variations in to do a cross-sectional study on the eo­ parasite numbers do not therefore neces­ variation of parasites with different sarily tell us anything about host differ­ routes of transmission. ences in susceptibility.

20 18 16 14 Q) (,) 12 � � 10 2! 8 Q. 6 4 2 0 2 3 4 5 6 Area hot in the

Fig. 2. Variation in prevalence of the fllarian nematode Splendidofilaria papillocerca from 6 different areas in Troms0 county, Norway. 36

A cross-sectional study in willow ptar­ Within the whole sample we found a sig­ Apaniu migan nificant agreement in how different para­ to redu If differences in host susceptibility has site species ranked the birds. Birds with switchi an important impact on the distributional high numbers of one parasite species transmi pattern of parasites, and if susceptibility tended to have high numbers of other search is determined by some general host char­ parasites as well. This study therefore intake. acteristic (e.g. nutritional level), we suggests that variations in host suscepti­ expose1 would expect that intensities of different bility do affect infection patterns, despite velopin parasite species will covary. We should a large stochasticity in exposure. It there­ make therefore look at the distributional pattern fore supports one of the basic conditions (Wakel - not of one parasite - but of many differ­ necessary if parasites are to play any role One ent species with different routes of infec­ in the evolution of host traits. these p: tion. The typical overdispersed distribu­ respon< tion of one parasite species tells us that it Benefits of host adaptations must ex­ The id< tends to accumulate faster in some host ceed costs sets of individuals then in others. If we look at If variability in susceptibility does just th� another parasite species this would play a role in the parasite burden ob­ kinds < probably also show an overdispersed served in a host individual, should we reprodt distribution, and so would a third and a then expect hosts to evolve resistance ecolog) fourth and so on. Then we could ask the mechanisms? This question is related to Davies, question: Do the different parasite species another requirement that should be added been st tend to accumulate in the same host indi­ to the list of Goater & Holmes (1997). If of a he viduals? If various parasite species, with host traits are to evolve as a selective off ag: different transmission biology, tend to response to parasitism host individuals Barnar1 covary in intensities, it must be because who develop the trait must on average 1996). they evaluate the host susceptibility have higher fi tness than hosts that do site ma similarly - some host individuals are gen­ not. If the advantage of having some trait den, b1 erally good hosts and others are bad. that confers resistance is small, because differ With this problem in mind we shot a much of the variability is determined by factors sample of 55 willow ptarmigan (Lagopus the lottery of exposure, and the evolved (Skorpi lagopus) at a small locality within a few trait in addition is costly, hosts that do 1995, : days (Holmstad & Skorping, 1998 ). The not evolve the trait may have higher fit­ develor birds were infected with 8 different spe­ ness. Both theoretical and empirical con­ effects cies of parasites. Some, like blood proto­ siderations suggest that most resistance reprodt zoans and the ftlarian nematode Splen­ mechanisms are expensive. Immune reac­ host-rel didofilaria papillocerca, were transmit­ tions require substantial metabolic in­ should ted by insect vectors, others, like coccidi­ vestment that otherwise could have been Forbes, ans and Trichostrongylus tenuis were used for other purposes (Wakel in & 1997). monoxenous. We ranked the host indi­ Apanius, 1997). An immune response Thi� viduals with respect to the numbers they will also have pathological consequences mal irr contained of each parasite species. Then for the host - indeed immunopathological with m we performed a non-parametric anova to damage may be more harmful to the host tion of test if the ranking deviated from random. than the parasite itself (see Wakelin & stress a ��-�------------

37

1und a sig­ Apanius, 1997). Behavioural adaptations the observed difficulties of inducing ster­ �rent para­ to reduce exposure may also be costly - ile immunity against macroparasites by Birds with switching to another prey to avoid a food­ vaccination (Behnke & Barnard, 1992). te species transmitted parasite, could require longer So far, however, it remains a fascinating ; of other search time and lower rates of energy idea. Developing testable models on im­ therefore intake. Finally, since hosts are usually munological behaviour under specified t suscepti­ exposed to many different parasites, de­ ecological conditions is therefore a major ns, despite veloping an immune response to one may challenge for the future. e. It there­ make it more vulnerable to others (Wakelin Apanius, 1997). conditions & References lY any role One way that hosts could get around these problems is to develop the ability to Behnke JM, BarnardCJ, Wa.kelin D. Under­ standing chronic nematode infections: Evo­ respond to parasites in a flexible manner. lutionary considerations, current hypotheses The idea that animals are equipped with must ex- and the way forward. Int J Parasitol 1992; sets of rules that that can be used to ad­ 22: 861-907 1ility does just the investment optimally in different Boulinier T, Sorci G, Monnat JY, Danchin urden ob­ kinds of activities such as foraging or E. Parent-offspring regression suggests should we reproduction has dominated evolutionary heritable susceptibility in a natural popula­ ecology for several decades (e.g. Krebs & resistance tion of kittywa.ke Rissa tridactyla. J Evol Davies, 1997). Within parasitology it has related to Biol l997; 10: 77-85 i be added been suggested that the immune response Forbes MRL. Parasitism and host reproduc­ (1997). of a host must be flexible and be traded If tive effort. Oikos 1993; 67: 444-50 z selective off against other demands (Behnke & ndividuals Barnard, 1992, Sheldon & Verhulst, Goater CP, Holmes JC. Parasite-mediated n average 1996). The fitness cost of having a para­ natural selection. In: Clayton DH, Moore J, �s that do site may vary, not only with parasite bur­ eds. Host-parasite evolution . General princi­ ples and avian models. Oxford, New York, some trait den, but also because parasite species Tokyo: Oxford University Press, 1997: 9-29 l, because differ in pathogenicity, depending on rmined by factors such as size, fecundity or location Holmstad PR, Skorping A. Covariation of 1e evolved (Skorping et al, 1991, Read & Skorping, parasite intensities of willow ptarmigan, Lagopus lagopus L. Can J of Zool 1998; in ts that do 1995, Skorping & Read, 1998). Recent press higher fit­ developments suggest that the fitness irical con­ effects of parasites may also depend on Hudson PJ. The effect of a parasitic nema­ resistance reproductive investment, age or other tode on the breeding production of red nune reac­ host -related factors and that the host grouse. J Anim Ecol 1986; 55: 85-92 abolic in­ should flexibly respond accordingly (e.g. Krebs, J. R and Davies, N. B. Behavioural have been Forbes, 1993, Skorping, 1996, M0ller, ecology. An evolutionary approach .. Oxford, 'akelin & 1997). Blackwell Science Ltd. 1997 This idea of the evolution of an opti­ response Lloyd S. Environmental influences on host mal immunological behaviour fits well 1sequences immunity. In: Grenfell BT, Dobson AP, eds. tthological with numerous observations on modula­ Ecology of infectious diseases in natural lo the host tion of immunity by host factors such as populations. Cambridge: Cambridge Univer­ Vakelin & stress or nutrition (Lloyd, 1995), and with sity Press, 1995: 327-61 38

Bull Sew M�ller AP. Parasitism and the evolution of Skorping A. Why should marine and coastal host life history. In: Clayton DH, Moore J, bird ecologists bother about parasites? Bul­ eds. Host-parasite evolution. General princi­ letin of the Scandinavian Society for Para­ pals and avian models. Oxford, New York, sitology 1996; 6: 98-102 Tokyo: Oxford University Press, 1997: 105- H 27 Skorping A, Read AF. Drugs and parasites: global experiments in life history evolution. Read AF, Skorping A. The evolution of Ecology Letters 1998; 1: 10-3 tissue migration by parasitic nematode lar­ vae. Parasitology 1995; Ill: 359-71 Wakelin D, Apanius V. Immune defence: genetic control. In: Clayton DH, Moore J, Sheldon BC, Verhulst S. Ecological immu­ Im eds. Host-parasite evolution. General princi­ nology: Costly parasite defences and trade­ ples and avian models. Oxford, New York, offs in evolutionary ecology. TREE 1996; Tokyo: Oxford University Press, 1997: 30-58 11: 317-21

Skorping A, Read AF, Keymer AE. Life Abstra history covariation in intestinal nematodes of Cer1 mammals. Oikos 1991; 60: 365-72 mospor mospor researc biology the dift of infe sampli1 shootin migrat< prevale (iv) the ficatior

Key "' genicit: proteu; Trypan

Mu evoluti, the res The ir harmle opporo both et parasit attracti investi; Bull Scand Soc Parasitol 1998; 8 (2): 39-46 and coastal tSites? Bul­ V for Para-

HAEMATOZOA OF WILD BIRDS: PECULIARITIES IN i parasites: ' evolution. THEIR DISTRIBUTION AND PATHOGENICITY

Gediminas V alkiunas te defence: , Moore J, Institute of Ecology, Lithuanian Academy of Sciences, Akademijos 2, Vilnius 2600, �ral princi­ Lithuania New York, 997: 30-58

Abstract Certain aspects of the ecology of hae­ tial theoretical traps because of the com­ mosporidian parasites (Sporozoa: Hae­ plicated life histories of these haemato­ mosporida) of birds which await future zoa, the epidemiology of the diseases and research in ecology and evolutionary the migratory behaviour of their avian biology are highlighted. These include: (i) hosts. The main aim of this paper is to the difficulty of estimating the true level highlight some important aspects of the of infection by blood-film surveys, (ii) ecology of haemosporidian parasites sampling bias in netting, trapping and [phylum Sporozoa (=Apicomplexa), or­ shooting of birds, (iii) the influence of der Haemosporida, genera Haemopro­ migratory behaviour of birds on infection teus, Plasmodium and Leucocytozoon] prevalence at a place of investigation, and that may await future research. These (iv) the problem of parasite species identi­ groups of bird haematozoa have been fication. investigated particularly well and have often been used in research on ecology Key words: haematozoa, birds, patho­ and evolutionary biology. genicity, evolutionary biology, Haemo­ proteus, Plasmodium, Leucocytozoon, Haemosporida of birds are obligate­ Trypanosoma heteroxenous parasites which use blood­ sucking Diptera (families Culicidae, Much theoretical work in ecology and Ceratopogonidae, Simuliidae and Hip­ evolutionary biology has been based on poboscidae) as final hosts and vectors. the results of surveys of blood parasites. The life cycles of the parasites are re­ The investigation of avian blood using viewed by Atkinson and van Riper (1991) harmless for host methods gives a perfect and Valkifulas (1997). opportunity to take large samples from both common and protected birds. These The difficulty of estimating the true parasitological methods are particularly level of infection by blood film surveys attractive to ornithologists. However, the An advantage of ecological research investigations have also presented poten- using haemosporidian parasites as a 40 model is that, once infected, birds remain chronic parasitemia even during the sea­ traps, infected either for life or many years (see son of active transmission. An extensive the Cu Valkiiinas, 1997). This allows a long­ examination of blood ftlms is necessary; relativ period control of infected ringed bird however, even this is not always enough been 1 specimens in natural populations. How­ to detect a Plasmodium infection. It avail a! ever, the period of patent infection is should be mentioned that even when re­ meth01 seasonally restricted and totally different corded, most of these chronic infections weak in species of Plasmodium, Haemoproteus of Plasmodium cannot be identified to sampl1 and Leucocytozoon. In the northern part species, even by experts, because of low thresh of the Holarctic, most species of malarian intensities and common mixed infections (Valki parasites (genus Plasmodium) cannot be (Laird, 1998). Experimental inoculation parasi found in the blood in early spring and of blood from wild birds into susceptible of Ht during the autumnal migration. The pe­ captive hosts gives better results but is birds, riod of chronic parasitemia of Haemo­ expensive. As a rule, searching blood and L proteus is longer than Plasmodium; how­ films reveals only a part of the real distri­ the he ever, gametocytes of most species of bution of malaria. It is important to note rarely Haemoproteus are usually still absent in that the same is true for Trypanosoma as nettin! the blood in the early spring migration, the intensity of Trypanosoma parasitemia is con and they disappear from the blood during is usually low. The microscopy of blood and n the autumnal migration. Extensive search smears is an insensitive method to deter­ specia of blood smears may reveal a few game­ mine the prevalence of Trypanosoma here. 1 tocytes of Leucocytozoon in the early infection in birds. The method of cultiva­ of pat spring migration as well as during the late tion of the parasites is at least 5-7 times lae d< autumnal migration and even in winter. more sensitive (see Kirkpatric, Lauer, cytes : As a rule, birds infected with Plasmo­ 1985; Valkiiinas, 1997). Thus, data on juveni dium and Haemoproteus cannot be re­ prevalence of Plasmodium and Trypano­ corded during early spring or late autum­ soma infections obtained by the blood­ nal migration by blood film survey, but smear technique does not reflect the para­ Leucocytozoon is often registered. That is sitological situation in free-living popula­ why the data on prevalence of infection, tions. Such data cannot be used to illus­ which are based on blood film micros­ trate theories in evolutionary biology. copy, should be critically used in ecologi­ Immunological methods are applicable Catchi and biJ cal investigations of Haemosporida. One for the estimation of the prevalence of should observe that absence of parasites infection (Graczyk et al, 1994) and Shooti in the blood does not necessarily mean promising on the generic level but are still that the investigated birds are not in­ inaccurate for parasite species determi­ Killed fected. The best season for blood investi­ nation. road gation using a microscope is May-July in Taken the northern part of the Holarctic. During Sampling bias in netting, trapping and kept a1 this period, most infections are patent. shooting of birds (95% ( Moreover, for Plasmodium the period of The majority of ecological investiga­ acute infection is very short with few tions on Haemosporida are based on birds Table parasites present in the blood during the caught in mist nets or special stationary pendi1 41

?: the sea­ traps, for example big Rybachy traps on that have been caught in mist nets and big extensive the Curonian Spit in the Baltic Sea. Only Rybachy traps (Table 1). The maximum 1ecessary; relatively healthy specimens that have level of the parasitemia in adult birds rs enough been leading an active life in nature are may be higher, but it usually does not �ction. It available for investigation using these exceed 10 %. Exceptions are extremely when re- method of netting and trapping, but birds rare and have been recorded only in adult infections weak by parasitic infections are under birds (Bennett et al, 1993). However, the ntified to sampled. All active birds are below a primary parasitemia significantly exceeds se of low threshold level of parasitemia in the wild the above-mentioned level, and may reach infections (Valkiiinas, 1997). That is about 40-50 25 % at the top of parasitemia in sponta­ loculation parasites per 1000 erythrocytes in species neously infected juvenile chaffinches that 1sceptible of Haemoproteus in young Guvenile) have been monitored in the laboratory. Its but is birds, and it is even less for Plasmodium Moreover, heavily infected young chat­ ng blood and Leucocytozoon. This explains why finches were shot, and also recorded as eal distri­ the heavy parasitemia has been extremely road killers (Table 1). It is interesting to tlt to note rarely registered in birds captured by note that one male great tit Parus major osoma as netting even on endemic territories, but it heavily infected with Haemoproteus ma­ Lrasitemia is common in experimental captive birds joris was recorded by the author in the of blood and may be detected in the wild using north of Lithuania. This bird was caught to deter­ special methods. One example is analysed by a domestic cat. In this case the inten­ 'anosoma here. On the Curonian Spit, the intensity sity of the parasitemia exceeded 30 % of 1f cultiva- of parasitemia of Haemoproteus fringil­ erythrocytes infected. These facts can be 5-7 times lae does not exceed 5 % of the erythro explained in the following way. It has :, Lauer, cytes and it is usually considerably less in been shown that infected young chat­ data on juvenile chaffinches Fringilla coelebs finches become less mobile at the top of Trypano­ le blood­ Intensity of parasitemia the para­ Number of parasites per 1000 erythrocytes g popula- Method Infected 1 to illus- minimum maximum biology. pplicable Catching by mist nets 674 <1 34 lience of and big Rybachi traps (24-48) 194) and Shooting 49 <1 122 1t are still (100-141) determi- Killed by cars on the 68 <1 69 road (55-87) Taken from nests and 5 <1 250* ping and kept at the laboratory (233-278) (95% confidence limits are given in parentheses) *data are given at the peak of parasitemia nvestiga­ l on birds Table 1. Intensity of Haemoproteus fringillae parasitemia in juvenile chaffinches de­ tationary pending on methodof investigation (June -August 1983-1992, 1994) 42 parasitemia (Valkiiinas, 1997). Reduced already passed through a heavy phase of Probabl mobility during the heavy phase of infec­ the infection and have acquired premuni­ been el tion makes the birds more vulnerable to tion (Ahmed, Mohammed, 1978; Garn­ specime predators and other unfavourable condi­ ham, 1980; Valkiiinas, 1997). Further­ (Valkiii tions. It implies a reduced ability to com­ more, the method does not provide infor­ that, in pete and an increased mortality rate in mation on the fate of birds from the group of the 1 nature. If birds survive the primary acute of non-infected specimens which have of infec attack of the parasites, then they may be caught the parasites between two periods be care caught in mist nets. This situation creates of recapture. Data on recapture of chaf­ of dise two important conclusions. Firstly, only a finches, either non-infected or infected Ideally, relatively healthy part of a bird popula­ (low chronic parasitemia) with Haemo­ plemen tion is usually available both for orni­ proteus fringillae, on the Curonian Spit It is thologists and parasitologists that have are shown in Table 2. Based on ringing res ear cl been been using netting and trapping for studies, the proportion of chronically bi catching birds. So, to measure the real infected and non infected birds during the birds o! impact of the parasites, special methods first year of life were almost identical. season of investigation should be designed. These Surprisingly, the data shows that signifi­ nonbre< methods must allow the observer to fol ­ cantly fever non-infected than infected chronic low the fate of birds during the critical birds were recaptured in the successive of recc phase of infection. Secondly, data on year. One may speculate that some non­ shot ga recaptures of wild ringed infected and infected birds have subsequently gained sequen1 non-infected birds, that are based on the the infection and have not survived the intensit netting and trapping, do not necessarily primary attack of the parasites. Accord­ be car contribute to the understanding of the ing to our data, over 20 % of non-infected ecolog) vitality of the individuals because the chaffinches gain the infection in the sec­ importl infected birds are specimens that have ond year of life on the Curonian Spit. data, tt of erytJ Ficedu Recaptured major sample in the year of birth in subsequent years Status Examined 1997). (age up to 4 months) (age up to 15 months) paras in n % n % exampl ods of to be a Non-infected 502 26 5.2 4.0 20 part of (3.4-7.4) (2.4-5.9)

Infected 299 17 5.7 28 9.4 The ir (3.3-8.6) (6.3-13.0) of birt (95 % confidence limits aregiven in parentheses) place ( Mi Table 2. Recaptures of chaffinches, either non-infected or infected with Haemoproteus also b( fringillae, on the Curonian Spit (breeding season in 1983-1992, 1994, 1996) sitolog 43 y phase of Probably, some of the infected birds have analysed here. A set sequence of nligra­ I premuni- been elinlinated as is found in juvenile tion of different populations of birds oc­ 78; Garn­ specimens during the first year of life curs during the seasonal nligrations . Further­ (Valkiunas, 1997). These examples show (Payevsky, 1985). As a result, data on vide infor- that, in order to measure the real impact infection of birds belonging to different 1 the group of the parasites, the results on recapture populations are available at each place of llich have of infected and non-infected birds should investigation. Mixing of data from differ­ vo periods be carefully interpreted taking all aspects ent populations causes difficulties in in­ e of chaf­ of disease epidenliology into account. terpretation of the parasitological situa­ r infected Ideally, field studied should also be sup­ tion at any site of investigation. Infection 1 Haemo­ plemented by experimental work of birds with Leucocytozoon on the Cu­ )nian Spit It is important to note that much of ronian Spit is an illustration (Fig. 1). In )n ringing research in ecological parasitology has summer, only breeding chaffinches of the hronically been based on blood parasites of game local Curonian population are present, during the birds obtained by shooting. The shooting and birds infected with Leucoc_ytozoon identical. season is usually strictly restricted to a are extremely rare. This is explained by tat signifi- nonbreeding period when parasitenlias are (i) the absence of transnlission of the 1 infected chronic or latent. This explains the rarity parasites on the Spit due to lack of vec­ mccessive of recordings of heavy parasitenlias in tors (there is no breeding places of Si­ ;ome non­ shot gamebirds (Valkiiinas, 1997). Con­ miliidae and, as a result, the young gen­ Jy gained sequently, the shooting data on infection eration of chaffinches is non-infected), ·vived the intensities in gamebird populations should (ii) rarity of infection of chaffinches at . Accord­ be carefully interpreted when used in wintering places and nligratory routes n-infected ecology and evolutionary biology. It is and (ill) relative stability of the Curonian n the sec­ important to note that, according to our population of chaffinches (the local birds Spit. data, the heavily infected (more than 5 % arrive to breed only on the Spit and the of erythrocytes infected) pied flycatchers number of inhabited infected chaffinches Ficedula hypoleuca and great tits Parus from populations nligrating over the ter­ major have never been recorded in bird ritory is quite negligible) (Sokolov, 1991, samples from nest-boxes (see Valkilinas, Valkiiinas, 1997). A number of infected years 1997). It is likely that birds with heavy young and adult chaffinches were re­ onths) parasitenlia do not enter the boxes. These corded on the Spit during nligration of % examples further show that special meth­ birds from northern populations ods of investigations should be designed (Karelian, Finnish, etc.) in spring and to be able to monitor the heavily infected autumn. It is worth noting that especially 4.0 part of the bird population. high prevalence of the infection has been 4-5.9) recorded in April and October when the 9.4 The influence of nligratory behaviour birds of the northern populations nligrate 1-13.0) of birds on infection prevalence at a over the Spit. According to the ringing place of investigation data, in autumn, the local population of zoproteus Migratory behaviour of birds should chaffinches leave for the wintering places also be taken into consideration in para­ before the nligration of birds of the north­ sitological investigations. One example is ern populations starts. Conversely, in 44

...... , . . An r- -, ... studies I graphic \2 :1 the reh I migran1 I \ juvenil< I I 70 tions. lJ . I into la . · . I l numbe1 � signific r. data. 11 I parasitt I birds. ' 30 terpret: tion at I prevale I breedir: .I 10 (preval % corr scapus VII VIII IV V VI IX X Sylvia Fig. 1. Infection of adult chaffinches, trapped on the Curonian Spit, in relation to the the Cu affiliation of the bird population interch separat I - cbaffinches from the northern populations migrating over the Spit; 2 - chaffinches from the local Curonian population (according to Sokolov, 1982); 3 - prevalence of Leucocytozoon, (Sokol1 4-6 - several cases of Leucocytozoon infection recorded in birds of the Curonian population. was e:x The abscissa is the calendar (months) and the ordinate: on the right - the number of trapped parasit birds (percentage); on the left - prevalence of infection (percentage). Vertical lines are 95-% creates confidence limits. parasit parisot spring, the northern populations migrate analysed with reference to the ornitho­ bird pc through the Curonian Spit before the logical situation in the area of investiga­ territor local population arrives from their win­ tion. Secondly, prevalence of infection in tion oJ tering grounds. This well explains the nonringed birds reflects the local parasi­ popula dynamics of infection of chaf:finches with tological situation only during a very Sor Leucocytozoon on the Spit (Fig. 1). The short period in summer when migrating are ve1 prevalence of infection strongly depends birds are absent from the region (for ex­ are ne' on the number of investigated birds be­ ample, from 10 June until 10 August on the Ne longing to the northern populations. This the Curonian Spit in the Baltic Sea); gain tl: situation also leads to important conclu­ however, the period might be prolonged on mi. sions. Firstly, to estimate the actual fre­ for about a month by investigating only prevalc quency of infection in a particular bird ringed specimens and birds caught on the very c: population, the data must be carefully nests. connec ------�

45

An important fe ature of ecological breeding season. As a result, the heavy studies on bird parasites in every geo­ parasitemias are totally absent in the 90 graphical area is the necessity to estimate study area and only low chronic para­ the relative number of long-distance im­ sitemias may be recorded. For example, migrants inhabiting the territory following Haemoproteus pallidus is very common juvenile dispersal and seasonal migra­ in Ficedula hypoleuca and F. albicollis 70 tions. Immigrants may introduce parasites in North Europe, but a great majority of into local populations. Sometimes, the the birds are infected in over-wintering number of immigrants is large enough to sites in Africa (Valkiiinas, 1997). significantly influence the prevalence data. It is important to note that unusual The problem of parasite species identi­ parasites may be brought in by immigrant fication birds. That causes difficulties in the in­ It is important briefly to note that the terpretation of the epidemiology of infec­ identification of species of bird Hae­ tion at the study site. For example, high mosporida is either rarely performed or prevalence of Leucocytozoon in the not always correct in studies on evolu­ breeding populations of Parus major tionary biology. Some misidentifications 10 (prevalence of the infection is 14.3 %, 95- are common. For example, H. pallidus % confidence limit is 7.6-23.7), Phyllo­ parasitizing Ficedula hypoleuca and F. scopus sibilatrix (14.2 %, 4.7-27.9) and albicollis has rarely been distinguished Sylvia atricapilla (47.6 %, 38.9-61.1) on fr om H. balmorali. Correct identification ion to the the Curonian Spit 1s a result of active is important because the life histories and interchange observed between widely virulence of different species are different .cbes from separated populations of these birds and unknown for many species of bird Jcytozoon, (Sokolov, 1991; Valkiiinas, 1996). As it parasites. Papers on evolutionary biology 10pulation. was explained above, the transmission of that are based only on generic identifica­ of trapped parasites does not occur on the Spit. This tion do not correspond to demands of are 95-% creates a situation where only low chronic current parasitology. parasitemias (relatively benign in com­ To stimulate the progress in ecology parison to heavy ones) are present in the and evolutionary biology using parasites ornitho­ bird populations. Without question, such as a model , joint projects on parasitology, investiga­ territories are not suitable for investiga­ ornithology and evolutionary biology are fection in tion of parasite influence on fr ee-living to be recommended. The participation of 11 parasi­ populations. parasitologists is important not only dur­ � a very Some species of bird Haemosporida ing the phase of investigation of blood migrating are very common in Palearctic birds but smears and identification of species of 1 (for ex­ are never (or infrequently) transmitted in parasites (as usually is the case), but also .ugust on the Northern Palearctic. As a rule, birds during the phases of planning and data tic Sea); gain the infection at wintering places and analysis. This would reduce the epidemi­ >rolonged on migratory path ways. The data on ological mistakes (see Valkiiinas, 1997) ting only prevalence of such infections should be that so often occur in ecological and :ht on the very carefully interpreted as they are not evolutionary studies using parasitological connected with transmissions during the data. 46

Acknowledgements morphology and metabolism. Academic Bull Sea I am grateful to the staff of Biological Press: New York et al ., 1980: 95-144 station of the Zoological Institute (the Graczyk TK, Cranfield MR, Skjoldager Russian Academy of Sciences) for help in ML, Shaw ML. An ELISA for detecting the field and advice in ornithology. I am anti-Plasmodium spp. antibodies in Afri­ also indebted to Dr. P. Henderson can black-footed penguins (Spheniscus (Department of Zoology, University of demersus). J Parasitol 1994; 80: 60-6 Oxford) for correcting the language. Laird M. Avi an malaria in the asian AVIAr References tropical subregion. Springer: Singapore, 1998 AND Ahmed FE, Mohammed A.-HH. Haemo­ STATl proteus columbae: course of infection, Payevsky VA. Demography of birds. NAL relapse and immunity to reinfection in the Leningrad: Nauka, 1985 (in Russian) AVIAr pigeon. Z Parasitenkd 1978; 57: 229-36 Sokolov LV. Postbreeding movements Robert 1 Atkinson CT, van Riper Ill C. Patho­ and fidelity to birth regions of the chat­ Protozo: genicity and epizootiology of avian hae­ finches at the Kurische Nehrung. In: Dol­ South E matozoa: Plasmodium, Leucocytozoon, nik VR, ed. Population ecology of the chaffinch (Fringilla coelebs). Leningrad: and Haemoproteus. In: Loye JE, Zuk M The m (eds). Bird-parasite interactions: Ecology, Nauka, 1982: 215-28 (in Russian) source evolution, and behaviour. Oxford Univer­ Sokolov LV. Philopatry and dispersal of specim< sity Press. 1991: 19-48 birds. Leningrad: Nauka, 1991 (in Rus­ stained Bennett GF, Peirce MA, Ashford RW. sian) begun · Avian haematozoa: mortality and patho­ asia in Valkiiinas G. Ecological implications of genicity. J Nat Hist 1993; 27: 993-1001 examin hematozoa in birds. Bull Scand SocPara­ Laird ' Garnham PCC. Malaria in its various sitol 1996; 6: 103-13 U.S. Al vertebrate hosts. In: Kreier JP (ed). Ma­ Valkiiinas G. Bird Haemosporida. Vil­ at Kua laria; 1. Epidemiology, chemotherapy, nius, 1997 (in Russian) expand< of the 1 ment C Waiter (Washi1 World United tional 1 larial P Newfou ship of C. Grei reorgan more at Centre ------m;

Academic Bull Scand Soc Parasitol 1998; 8 (2): 47-87 15-144 Skjoldager r detecting es in Afri­ ABSTRACTS OF SUBMITTED PAPERS: )pheniscus ORAL AND POSTER PRESENTATIONS ): 60-6 the asian Singapore, AVIAN PARASITE DATABASES under the direction of Professor Gordon AND COLLECTIONS: CURRENT F. Bennett. In 1995, on Gordon Bennett's STATUS OF THE INTERNATIO­ retirement, the IRCAH was donated to of birds. NAL REFERENCE CENTRE FOR the Queensland Museum in Brisbane, ssian) AVIAN HAEMATOZOA (IRCAH) where it now resides. The IRCAH collec­ novements tion has now been incorporated into the 'the chaf­ Robert D. Adlard collections of the Queensland Museum Protozoa Section, Queensland Museum, g. In: Dol­ and each specimen has been issued with a South Brisbane, Queensland, Australia 'gy of the museum registration number in addition Leningrad: to its former IRCAH accession number. The IRCAH collection is a unique re­ The collection will retain its integrity, be m) source which consists of over 64,000 promoted as the recognised global re­ tspersal of specimens of avian blood parasites in pository for specimens of avian haemato­ (in Rus- stained blood films. The collection was zoa, and its future protected by the policy begun with samples collected in Malay­ of the Queensland Museum to preserve asia in 1959 by H. Elliot McClure, and cations of specimens in perpetuity. Although Aus­ examined for parasites by Dr. Marshall Sac Para- tralia now houses the IRCAH, there are Laird with financial support from the records of haematozoa from only 16 U.S. Army Medical Research Unit based species of Australian birds from a total of rida. Vil- at Kuala Lumpur. This collection was 770 species recorded. Research to docu­ expanded in 1963 under the joint auspices ment the blood-borne parasites of birds of the U.S. Army Research & Develop­ has commenced in a collaborative pro­ ment Command (Tokyo, Japan) and the gram between the Queensland Museum Waiter Reed Army Institute of Research and the Currumbin Sanctuary, Gold (Washington, D.C., U.S.A.). In 1968, the Coast, Queensland. Preliminary results World Health Organisation (WHO, show relatively high intensities of hae­ United Nations) established the Interna­ matozoa in species of the avian families tional Reference Centre for Avian Ma­ Cracticidae and Meliphagidae/Oriolidae. larial Parasites at Memorial University, Newfoundland, Canada under the leader­ ship of Dr. Marshall Laird and Dr. Ellis C. Greiner. In 1975, the collection was reorganized and its title amended to the more appropriate, International Reference Centre for Avian Haematozoa (IRCAH), 48 Egypt WSEVOLOD BOIUSOVICH DU­ hensive study of this group of bird para­ BININ (1913-1958) AS A RE­ sites in the world. Hafez SEARCHER OF BIRD PARASITES W.B. Dubinin created a new system of data 01 Acari as a whole and determined the de si rat Aleksander F. Alimov and Helen V. Dubin­ position of Acari within the system of Mater ina Chelicerata (1956, 1957). Classification A. arb �logical Institute, Russian Academy of of separate groups of mites of superfami­ Egret Sctences, St. Petersburg, Russia lies Analgesoidea, Demodicoidea and Orang( Cheyletoidea was elaborated in detail made a Wsevolod Borisovich Dubinin (1913- (1952-1954, 1956, 1957). Many repre­ to leru 1958), being a researcher of the Zoologi­ sentatives of these superfamilies are bird popula cal Institute and Deputy Director for the parasites. Result Zoological Museum, worked in different In a study of parasitic Acari of bird� bird CO areas: ecological-parasitological studies and mammals W.B. Dubinin elaborated At tl of different vertebrate animals, a study of and widely used zonal-microclimate (Noven nest parasites and other inhabitants of principle of the study of habitat of these compot burrows and nests, broad zoogeographic parasites, principles of simultaneous engorg1 studies. A particularly convenient object integrated study of specific traits of Acari Ill (36� of this research were feather mites. The structure (their morpho-functional char­ laid eg; study of these mites permitted him to acteristics) and their biological functions. consist: reveal a number of patterns in distribution Such approach permitted to give up which ; and relationships of Tubinares (1949, considering many Acari as "aberrant" essenti� 1953), Anseriformes (1950, 1953), species and show origin of separate appear; Galliformes (1950, 1956), Alcidae species and parasitic fauna of the host as (66% I (1952), Pelicaniformes (1953) and ratite a whole (1947, 1951, 1953, 1956). decreas birds (1956). The important role of and ad Antarctica (Archigea) in the Eocene and CHANGES IN POPULATION nesting Miocene for the settlement of many birds STRUCTURE OF AN ARGASID tick poJ from Indo-Malayan and Australian region TICK ARGAS ARBOREUS DURING strong 1 to South America and much later invasion fr om 7 NESTING SEASON OF ITS HOST' to Africa, Asia, Europe, and North THE CATTLE EGRET BUBULCUS number America (1950-1956) was shown on the IBIS, IN SOUTH AFRICA The se> basis of the vast parasitological material. acterise A thorough study was conducted of 2 Valentin N. Belozerov1 and Grzegorz Kopij decreas1 feather mites. This permitted him to 1 Biological Research Institute, St. Petersburg 52.6% create a three-volume monograph of the State University, Russia; 2University of the increase series Fauna of the USSR (1951, 1953, Orange Free State, Blocmfontcin, South Conclru 1956). It is a comprehensive study of Africa structur feather mites including not only fauna, Cattle E Objective: The life cycle of Argas ar­ morphology, taxonomy of this group of similar boreus Kaiser et al ., a common parasite parasites, but also data on their ecology, chronise host specificity, formation of parasite inhabiting rookeries of the Cattle Egret host. 11 Bubulcus ibis (L.) all over the African fauna, zoogeography, and paleontology. this tick Up to now it remains the most compre- continent, was studied earlier only in 49

Jird para- Egypt (Kaiser, 1966; Guirgis, 1971; lengthening due to some adaptive sea­ Hafez et al, 1971,1972). Comparative sonal delays of development at nymphal system of data on its life cycle from other areas are stage and of reproduction at adult stage. nined the desirable. ;ystem of Materials and Methods: Collections of PARASITE INFECTION IN THE �sification A. arboreus from a rookery of the Cattle GREAT TIT uperfami­ Egret during its nesting season in the Orange Free State (South Africa) were Angharad Bickle and F. Clark idea and Biology Department, Leicester University, made and treated afterwards in laboratory in detail Leicester, UK. ny repre­ to learn seasonal changes in the tick population. s are bird The parasite fauna of the great tit (Parus Results:The nesting season in the studied major L.) population on Gotland, Sweden i of birds bird colony lasts from October till March. has been studied for the last seven years. �laborated At the beginning of the season This study, from 1996-1998, focuses on :roclimate (November) the population of ticks is the sexual signal, parasite status and t of these composed of overwintered, freshly­ immune response. tultaneous engorged adult ticks (64%) and nymphs Fifty fields of a thin blood smear, Ill (36%), as well as of numerous freshly­ s of Acari taken from the brachial vein (at various nal char­ laid eggs. The only change in December times of the day), were examined using a functions. consists in the hatching of larval ticks x 100 oil immersion objective. Parasite give up which actively attack the nestlings. More and leukocyte counts taken from two 'aberrant" essential changes occur in January due to separate smears of the same blood sample separate appearance of numerous nymphs I-ll were repeatable. 1e host as (66% of the whole population) and a The two most common blood parasites decrease in number of nymphs Ill (9%) 6). identified: Haemoproteus majoris and and adult ticks (25%). At the end of Hepatozoon parus were less prevalent nesting season (February) the changes in LATION than in the years examined by Allander & tick population had opposite direction (a RGASID Bennett(1994). Isospora sp. were identi­ [)URING strong decrease in number of all nymphs fied in faecal samples. There was no ; HOST, from 75 to 44%, with an increase in significant difference between the leuko­ rJULCUS number of adult ticks from 25% to 56%). cyte counts of chicks at day 14 then The sex ratio among adult ticks is char­ resampled as breeding adults. Whether acterised by a male dominance with its the leukocyte count of individuals is Jrz Kopip decrease from 61.3% (November) to significant in terms of fitness is still to be 52.6% (January) and with a further Petersburg demonstrated in this population. Inter­ ;ity of the increase up to 60.2% (February). estingly initial analyses shows that the in, South Conclusions: The changes in population chin area (ranked) is negatively correlated structure of the tick A. arboreus in the with the total white blood cell count (r2= Cattle Egret rookeries in South Africa are lrgas ar- -0.31, p<0.05, n=53), however; the similar with those in Egypt being syn­ sample can be increased and work still 1 parasite chronised with seasonal rhythms of the continues. !tiC Egret host. The usual univoltine life cycle of � African this tick can be complicated partly by its only in 50

Reference and dilepidids in seawater use polychaets, HELM RADRJ Allander K. & Bennett G.F. Prevalence and gammaridian amphipods and euphausi­ PE CUI intensity of hematozoon infection in a acean crustacean as intermediate hosts. GRAPl population of great tits Parus major from Tetrabotriids use different species of fish 25: Gotland, Sweden. J Avian Bioi 1994; 69- as paratenic vertebrate hosts. The fresh Svetlana 74 Hymenolepis water species of (sensu maviCim lato) (Hymenolepedidae) of gulls use only THE INTERMEDIATE HOST AS Institute branchyopodien artropods, Anostraca, ECOLOGICAL DETERMINANT OF Notostraca, and Conchostraca, as inter­ The ori TAPEWORM FAUNA OF GULLS mediate hosts. It is known from the erature Svetlana Bondarenko paleontological data that (i) just Poly­ (except Institute of Ecology, Vilnius, Lithuania chaeta may have given rise of Branchio­ ised. Th poda, (ii) the distribution latter in sea of helm Objective: The role of intermediate host tookplace during Cambrian time and (iii) bigger 1 in the forming of cestoda fauna of birds is than Branchiopoda have been found only hosts. J discussed. in inland waters. Current tapeworm fauna faunae ' Material and Methods: The life cycles of birds fe eding on branchiopods is Alaska of 26 species of cestodes were used for determined by the species of cestoda ancient analysis, including our own data con­ which has historically developed in these territory cerning the life cycles of 8 species from invertebrates. Freshwater oligochaets are Problem tundra of West Chukotka and 3 species intermediate hosts for 4 species of Aplo­ new ter from the Northern Sea of Okhotsk. Four­ paraksis of gulls which are usually places tJ teen species of gulls from the North parasites of shorebirds. Dilepidids use tionship Pacific basin, Alaska and Antarctic as gammarids and dipteran larvae ecosyste well as their cestodes were divided into (chironomids) as intermediate host. The hosts, ar, two ecological groups with respect to life cycle of Wardium fryei give the association in the breeding season with opportunity to understand the role of the PARAS marine or inland fresh or brackish-waters. intermediate host in evolution of life BIRDS The first group of gulls includes 4 spe­ cycles. Comparison of the data dealing LANDS cies, the second one consists of 10 spe­ with the process of host-s peciation for cies. Respectively, the first group of large gulls, the main hosts for W. fryei, Fred H.M cestodes includes 8 species: Hymeno­ with the data on the distribution of para­ Institute 1 DLO , lepididae -2, Dilepididae -4, Tetrabothrii­ site clearly has confrrmed the correctness ) Le dae -2. The composition of the second of hypothesis that tapeworm phylogeny From 19 group is: Hymenolepididae -10 species, does not mirror vertebrate host phy­ Wild Bil Dilepididae -3 and Schistocephalinae-1. logeny. At the same time this comparison and fro Only 4 species of Pseudophyllidae occur­ has corroborated the theory that in these investiga ring in both groups indicates the connec­ cestodes evolutionary history of the birds of tion of birds with both fresh-water and cestode-invertebrate relationships was Strigifon littoral feeding sites. more archaic. (17.4%) Results and Conclusions: The cestoda 470 (57 fauna show clear subdivision of seawater parasites and freshwater species. Hymenolepidids 51

HELMINTH FAUNAE OF CHA­ Buteo buteo (n=2027, 278 died of illness, polychaets, RADRIIFORMES IN ALASKA: of which 127 due to parasites), Accipiter euphausi­ PECULIARITIES OF ZOOGEO- gentilis (341, 44, 28), Accipiter nisus liate hosts. GRAPHY AND ORIGIN (462, 62, 44), Falco tinnunculus (603, cies of fish 181, 111), Tyto alba (283, 38, 31), Strix The fresh Svetlana Bondarenko and Vytautas Kontri­ aluco (187, 49, 34), Athene noctua (176, pis (sensu mavicius 43, 18) and Asio otus (361, 85, 62). Owls lls use only Institute of Ecology, Vilnius, Lithuania died more frequently fr om parasites (67% Anostraca, vs 54%) than Falconiformes. In Falconi­ 1, as inter- The original investigations and the lit­ formes, coccidiosis was common in B. from the erature on helminths of Charadriiformes buteo and F. tinnunculus. Infections with just Poly­ (except Alcidae) in Alaska are general­ Capillaria spp. and ascarids were com­ " Branchio­ ised. The number of the Holarctic species mon in all above mentioned bird species. tter in sea of helminths in Alaska is significantly Infections with spirurids were commonly ne and (iii) bigger than the number of their final observed in B. buteo, F. tinnunculus, T. found only hosts. Most probably that the current alba and A. otus, occasionally in the vorm fauna faunae of Charadriiformes helminths in other mentioned species. Strigeid trema­ hiopods is Alaska maintains the features of the tode infections were frequent in B. buteo, of cestoda ancient fauna of the single Pleistocene less in F. tinnunculus, A. gentilis and the >ed in these territory of the East-Siberia and Alaska. owls, but absent in A. nisus. Cestode xhaets are Problems of (i) the helminths settling in infections were common in B. buteo only. �s of Aplo­ new territories during settling in new The airsac worm Cyathostoma spp. was re usually places their final hosts and (ii) the rela­ found in B. buteo, A. gentilis and all :pidids use tionship of parasitic worms with the owls. Acanthocephalan infections were 1 larvae ecosystems through their intermediate only common in S. aluco. host. The hosts, are discussed. i give the ANALYSIS OF RETALIATORY role of the PARASITIC INFECTIONS IN REACTIONS IN DOMESTIC :m of life BIRDS OF PREY IN THE NETHER­ QUAILS AFTER CHALLENGE BY 1ta dealing LANDS SALMONELLA ENTERITIDIS ciation for r W. fryei, Fred H.M. Borgsteede and Pedro E.F. Zoun Dalius Butkauskas and Jonas Babonas Institute for Animal Science and Health (ID­ )n of para­ Institute of Ecology, Vilnius, Lithuania correctness DLO), Lelystad, the Netherlands phylogeny Objective: Domestic quails ( Cotumix From 1975-1990, the Working Group on host phy­ cotumix japonica ) differ in shorter time Wild Bird Mortality in the Netherlands, �omparison of growth and higher intensity of meta­ and fr om 1991-1995 ID-DLO have tat in these bolic processes comparing with hens and investigated the cause of death of 4720 y of the turkeys. Intensity of metabolic processes birds of prey (3685 Falconi- and 1035 :hips was causes quails resistance to contamination Strigiformes). Of these birds, 820 with salmonellas. However, this question (17.4%) died of illness. Of the ill birds, has no widespread investigation. In 470 (57.3%) died of infections with reference to it we decided to evaluate parasites. The most common birds were 52 lewsky, reactions of quails after challenge by S. concentration of pyruvates was ascer­ (Frohlic enteritidis wild strain. For this aim tained at the same ti me. populati several indexes were analysed: 1) death Conclusions: Inactivated vaccine dominar rate; 2) changes in concentration of "TALOV AC 109 SE" prevents quails the urb carbohydrates; 3) dynamics of egg laying; from mortality and breaking down of egg accordir 4) eggs and droppings contamination with laying after challenge by virulent S. commor S. enteritidis . enteritidis culture. Challenge disturbed (32.0%) Materials and Methods: Forty quails metabolic processes of carbohydrates dominar were separated into two groups each of both in groups V and N, however, con­ was for 20 birds. First group (V) consisted of centration of glucose in group V recovers dularis birds which were vaccinated twice (on in significantly shorter time. No salmo­ nutus (I' 30th and 45th days of life ) with inacti­ nella could be isolated from droppings differen' vated "T ALOVAC 109 SE" vaccine. and eggs one month postchallenge. females· Second group (N) was formed of nonvac­ tions are cinated quails. BothV and N groups were SPECIAL FEATURES OF FORMA­ 46.2% a challenged orally by virulent S .enteritidis TION OF FAUNA OF HELMINTHS 1.85, P: culture of one day growing in broth IN URBAN AND WILD MALLARD populati medium in dose 0.2 mllbird. Registration ANAS PLA TYRHYNCHOS L. POPU­ was 3 ti of death cases and dynamics of egg laying LATIONS differem was started at first day after challenge. young rr Blood samples were obtained from wing Elizabeta B ychkova are insi, vein in such order: one week before, one Institute of ZOOlogy, National Academy of 1.97, p: day post and one week post challenge. Sciences, Minsk, Belarus Thus Peculiarities of metabolism of carbohy­ Civilization process causes formation of assemble drates were evaluated according to con­ urban populations of wild birds near big and the centrations of glucose, lactates and cities and health-resort areas. Mallard with CC( pyruvates. Droppings and eggs were shows a clear tendency for location near with its � examined for presence of S. enteritidis big cities. The high densityof mallards in one month afterchallenge. urban populations and special features of PARAS Results: During firstweek after challenge ecology influence the helminthfauna of LARUS 25 eggs in group V and 5 eggs in group N this bird. In 1995- 1997, 106 mallards were collected. Further laying in group N from an urban (Minsk) and a wild (Minsk Elizabeta was disturbed entirely and didn't recover and Neswizh districts) populations were tchevskf at all. Two weeks after challenge dying of 1 dissected and examined for helminth Institute birds in group N began. Mortality rose up Sciences, parasites. Examinations showed the to 20% . There was no dying in group V. Universit following results. The infestation was Presence of salmonellas in droppings and higher in the wild population of Mallard eggs shell, yolk and albumen was exam­ The fau than in the urban one (75.6%, 53.8%; 2 ined by traditional cultural methods. No x Accipitri =5.08, P=0.03). However, the abundance serotype of salmonellas was isolated one Belarus 1 of certain species of helminths month after challenge. Breaking down of are undeJ [Echinoparyphium recurvatum (Linstow, amount of glucose and lactates in quails 1873), Bilharziella polonica (Kowa- blood samples was significant. Arising of 53

lewsky, 1895), Echinostoma revolutum 79 birds from eight species of falcons as ascer- (Frohlich, 1802)] was higher in the urban collected in Belarus were examined for population. The trematodes were the helminth parasites between 1986-1996. vaccine dominant parasites in both the wild and A total of 14 helminth species (8 tts quails the urban populations (32.0%, 74.0% nematodes, 5 trematodes, 1 cestode) were wn of egg accordingly). The acanthocephales were recorded from birds of family Accipitri­ rulent S. common only in the wild population dae. Helminths were found in 49 birds disturbed (32.0%). In the wild population, the (62.0%). 30 birds were free of helminth ohydrates dominant complex of parasitic worms parasites. Eight helminth species were wer, con­ was formed by Plagiorchis multiglan­ classified as the host specialists of fal­ r recovers dularis Semenov, 1927, Cotylurus cor­ cons, and 6 species were noted to be the lo salmo­ nutus (Rudolphi, 1808), B. polonica. The generalists in raptors and other bird droppings differences of infestation of males and species. Several widespread species tge. females in the urban and the wild popula­ [Porrocaecum depressum (Zeder, 1800), tions are insignificant (88.2% and 66.7%; P. angusticolle (Molin, 1860), Syn­ li'ORMA­ 2 2 46.2% and 63.4%; X = 2.49, P=0. 1; X = himantus laticeps (Rudolphi, 1819)] are VIINTHS 1.85, P=0.25 accordingly). In the urban the basis of helminthfauna of falcons. lLLARD population, the infestation of adult birds The high intensity of infection of juvenile ,. POPU- was 3 times higher than young ones . The forms of P. depressum and Cladotaenia differences of infestation of adult and globifera (Batsch, 1786) was recorded young mallards from the wild population from April to December in Talpa eu­ 2 �ademy of are insignificant (82.4%, 70.8%; X = ropaea L. (1-222) and Sorex araneus L. 1.97' p =0.25). (1-18). It shows that these species of Thus the differences in helminth helminths are widespread on territory of mation of assemblages of mallards from the urban Belarus. The prevalence of the infection : near big and the wild populations are connected was especially high in Buzzard (75.8%) Mallard with ecological conditions, first of all and Marsh Harrier (66.6%). ttion near with its environment. Thus, the falcons are heavily infested tallards in with helminths. The fauna of the parasites �atures of PARASITES OF FALCONS IN BE­ includes both stenoxenic and euryxenic tfauna of LARUS species. mallards 1 Id (Minsk Elizabeta Bychkova and Alexandr Vin­ EXPERIMENTAL TRIALS TO 2 ions were tchevski INFECT DOMESTIC DUCKS WITH 1Institute of Zoology, National Academy of helminth COOT DIORCHIDS (CESTODA, Sciences, Minsk, Belarus; 2Grodno State wed the HYMENOLEPIDIDAE) University, Grodno, Belarus tion was f Mallard Danuta Cielecka, Bogdan Czaplinski, Gizela 2 The fauna of bird parasites of family Olszewska and Barbara Grytner-Zivcina i3.8%; X Accipitridae has not been studied in Department of General Biology and Parasi­ bundance Belarus because many species of raptors tology, The Medical University of Warsaw, ths helmin are under protection of law. Warsaw, Poland (Linstow, (Kowa- 54

Many authors report about common coot HEMATOLOGICAL AND BIO­ showed diorchids parasitizing numerous species CHEMICAL INVESTIGATIONS ON group : of Anseriformes. In order to verify if the EXPERIMENTAL COCCIDIOSIS IN was am domestic ducks may play the role of the MEAT CHICKENS as wel final host of Diorchis brevis Rybicka, untreatc 1 1957, of D. inflatus (Rud., 1819) and of Doina Codreanu-Balcescu and Dumitru also d 2 D. ransomi Johri, 1939, sixty birds of Curca chicken 1 different age (from one day, one week, Institute of Biology, Romanian Academy, The lo\ Bucharest, Romania; 2Faculty of Veterinary one month, two months to adults) have ascorbi Medicine, Bucharest, Romania been exposed to experimental infestation. groups Invasive oncospheres have been taken were t Objective: The aim of this study was to from cestodes found in Fulica atra L. and chicken establish hematological and biochemical contacted with Heterocypris incongruens Conclu modifications in Eimeria tenella experi­ (Ramd.), Cypridopsis vidua (O.F.M.), C. biochen mentally infested meat chickens, follow­ newtoni Brady et Rob., Cyclocypris serious ing different treatments with the Roma­ laevis (O.F.M.), and Notodromas experin nian anticoccidial Colidiniu. monacha (O.F.M.), in which invasive They pl Materials and Methods: The experi­ cysticercoides have been obtained. The lactic e ments were carried out as follows: group experiments were carried out from June cidial c 1 - control ; group 2 - 25,000 E. tenella to December. The following numbers of action oocysts were administrated to 30 days old ducks have been exposed to infestation drinkint chicken; group 3 - 25,000 E. tenella with cysticercoids of: to deter oocysts were administrated to 30 days old prevent chicken and in the 5th day p. i., when the D. brevis 6 ses. D. injlatus- 23 clinical signs have been appeared, 120 D. ransomi 22 mgll Colidiniu in drinking water was THE D. brevis + D. injlatus 2 administrated and continued for 5 days; AND D. brevis + D. injlatus + D. ransomi 7 group 4 -25,000 E. tenella oocysts and PARA� 120 mgll Colidiniu in drinking water were After infestation faeces were examined simultaneously administrated to 30 days Dumitru th th old chicken; the anticoccidial administra­ Balcesct every day, and from 14 to 18 day, 1 tion was continued for 10 days. Blood Departt sometimes later) and afterwards post­ was collected by cubital vein puncture Veterim mortem examinations were done. In each 2 fr om the 40 days old chickens and hema­ lnstitutl case negative results have been obtained. Buchare tologic (hemoglobinemia, hematocrit, Therefore, we agree with the opinion of mean corpuscular hemoglobin concentra­ Tolkacheva (1991) who examined rich Objecti tion (MCHC) as well as biochemical Russian and Asian collections of dior­ of hem (glycemia, proteinemia, ascorbic acid and chids and came to conclusion that there is haviour calcemia) parameters were determined by no natural exchange of these tapeworms were pu current methods of the clinical laboratory. between coots and ducks. Of course, with eh Results: It was emphasized a marked exceptions cannot be excluded, but they ment of lowering of hemoglohinemia in infected need careful and exact confirmation. the AA untreated chickens. The hematocrit mental < 55

showed a marked decreasing tendency in Materials and Methods: We made the BIO­ group 2 compared with group 3, which experiments on 10 days old broilers )NS ON was ameliorated for group 3. The MCHC (chickens), supplementary fed with 2 g )SIS IN as well as glycemia decreased in the AA/kg fodder, monitored in comparison untreated infested chickens. The glycemia with some broilers left as control. Both Dumitru also diminished in a hypoponderable groups of broilers were subjected to heat chickens group infested with E. tenella. stress; the environmental temperature was ° \.cademy, The lowest values of the proteinemia and 28-32 C, the normal one for this age 'eterinary ascorbinemia were recorded for the category being 22-26°C. Another experi­ groups 2 and 3. Higher levels of calcemia ment, the cockerel Leghorns were infected were registered in untreated infested with 1,200 Ascaridia galli eggs. Blood y was to chickens compared with the treated ones. was collected by puncture in the cubital chemical Conclusions: The hematological and vein, using heparin as an anticoagulant. 1 experi­ biochemical parameters emphasised The AA content in the whole blood and , follow­ serious perturbations in the meat chicken tissue was determined according to the � Roma- experimentally infested with E. tenella. technical indications described by Roe They proved the therapeutic and prophy­ and Kuether. experi­ lactic effects of the Romanian anticoc­ Results: After 4 weeks of breeding at ° •s: group cidial Colidiniu. It has no toxic or adverse temperature 28-32 C, the weight in­ ·. tenella action in concentration of 120 mg/l in creases, the hematologic and biochemical days old drinking water. Further studies are needed investigations have made evident a series tenella to determine the therapeutical schedule to of modifications as: lowering of hema­ days old prevent or to treat the chicken coccidio­ tocrit and pyruvic acid, as well as of AA �¥hen the ses. in bursa Fabricius; increase of mean �ed. 120 corpuscular haemoglobin concentration lter was THE ASCORBIC ACID, STRESS (MCHC) and of protein concentration, of 5 days; AND POULTRY REACTIVITY IN turbidity with zinc sulphate and of AA ysts and PARASITIC DISEASES content in: adrenal glands, kidneys, liver, 1ter were spleen and total blood. The experimental 1 30 days Dumitru Curca and Doina Codreanu­ infestation with A. galli eggs revealed 2 ministra­ Balcescu decrease of the AA concentration in 1 s. Blood Department of Physiopathology, Faculty of adrenal glands, starting the 9th day, Veterinary Medicine, Bucharest, Romania; puncture leukocytosis, heterophilia and eosino­ 2Institute of Biology, Romanian Academy, 1d hema­ penia; lymphopenia was reported only Bucharest, Romania matocrit, during the fr rst days after infection, mcentra­ Objective: This paper presents the results subsequently followed by eosinophilia chemical of hematological, biochemical and be­ and a slight lymphocytosis. acid and haviour investigations in chickens which Conclusions: The pathogenic action of nined by were put to a caloric stress in comparison the higher environmental temperature as JOratory. with chickens which received a supple­ the parasitic infection are permanent marked ment of ascorbic acid (AA) in food, also stressing factors, which affect the host's infected the AA status in chickens with experi­ life as well as the evolution of the rela­ :matocrit mental ascaridiasis. tionships between the organism and the 56

diorchic parasites, including the haematological numerous incorrect determinations of the D. injl and immunological restructuring and also cestodes found by many authors in ducks. probabJ the morphological and physiopathological It is so not only because of the wrong recurva modifications. synonyms, but also because of identical possibil number and similarity of shape and size becaus< CONSEQUENCE OF A HIGHLY of the rostellar hooks of following tape­ the bla< PROBABLE ERROR IN LABELING worms: 1) Diorchis ransomi from coots scolex · DIORCHIS INFLA TUS AND D. with D. parvogenitalis from ducks, 2) D. toid be< RANSOM/ FROM AYTHYA NYROCA injlatus, D. americanus and D. brevis long ha GULDENSTADT from coots with D. stefanskii as one of D. elis, the most common parasites of ducks. parasiti Bogdan Czaplitiski and Danuta Cielecka Czaplitiski B., Szelenbaum D. (1974) and reporte1 Department of General Biology and Parasi­ Tolkacheva (1991) indicate how to tology, The Medical University of Warsaw, probabl distinguish each of the above mentioned Warsaw, Poland becaus< tapeworm species. In our opinion the corresp common coot diorchids are, as a rule, Czaplinski (1956) noticed and figured acumin unable to parasitize ducks. Diorchis injlatus and D. ransomi in 2 out referrin of 23 examined Aythya nyroca Gilld., DIORCHIS ACUMINA TUS (CLERC, be out determined at that time as Nyroca rufa 1902) CLERC, 1903 (CESTODA, sider L (L.), shot at the Druzno Lake (Mazurian HYMENOLEPIDIDAE) - A COM­ acumin Lakeland in Poland). A re-examination of PLICATED TYPE OF THE GENUS genus this material revealed that both white­ DIORCHIS gether ' eyed ducks were infected exclusively by take, 01 these two species of tapeworms: 2 and Bogdan Czaplitiski and Danuta Cielecka perime1 176 specimens of D. injlatus and 1 and Department of General Biology and Parasi­ 141 specimens of D. ransomi. Because tology, The Medical University of Warsaw, no tapeworm species characteristic of Warsaw, Poland Table 1 Anatidae have been simultaneously fo und, and no similar result has been The present authors, thanks to Dr. C. Region repeated in over one thousand anseriform Vaucher, were able to study a preparation birds post-mortem examinations, one deposited in the Museum of Natural Sacbalit explanation seems acceptable: the tape­ History with inscription: "(T) Institut de Kamcba worms were erroneously labeled as Zoologie, Diorchis acuminata (Clerc, Poland coming fr om white-eyed ducks instead of 1902), Fulica atra, Oural, M.M.V. 6/64 from coots. The relatively good figures of 13176 Universite de Neuchatel". This Slovaki: both mentioned tapeworms caused their preparation contains 15 tapeworm fr ag­ Turkme ments stained in the state of strong con­ citations by many authors and led to Chany l erroneous conclusions, among others, to traction not useful for exact morphologi­ Kazakbl the wrong synonymy of D. parvogenitalis cal studies. However, there are among Tadjikis with D. ransomi as it has been done e. g. them two fragments with scolices: the in two known monographs by Spassky first with 10 diorchoid hooks 68 jl,m long, Canada (1963), and by Spasskaya (1966) and belongs to one of the three possible coot Canada ------$'

57

diorchids: D. americanus, D. brevis and HOST SPECIFICITY OF CLADO­ ons of the D. inflatus, the second being the most GYNIA PITTALUGAI (LOPEZ­ ; in ducks. probable because of the characteristic NEYRA, the wrong 1932) recurvation of the blade. The fourth f identical Bogdan Czaplitiski\ Urszula Czaplitiska2, possibility - D. stefanskii is excluded � and size 1 because of the lack of a small incision at Barbara Grytner-Zivcina and ving tape­ 1 the blade base. The second fragment with Anna Gierczak ram coots 1 scolex with 10 diorchoid (close to arcua­ Department of General Biology and Para- cks, 2) D. sitology, The Medical University of Warsaw, D. brevis toid because of short guard and blade and Warsaw, Poland; 2W. Stefatiski Institute of as one of long handle) hooks 28 �-tm long belongs to D. elisae, which is a typical tapeworm Parasitology, Polish Academy of Sciences, of ducks. Warsaw, Poland 1974) and parasitizing Anatidae, until now not reported from Fulica atra. D. brevis was how to Cladogynia pittalugai has been described probably not noticed by Clerc (1903), mentioned for the first time from Anas platyrhyn­ because its length of hooks does not 1inion the chos L. coming from Grenada (Spain). correspond with his own description. D. ts a rule, Two specimens of this species were found acuminatus is, as a matter of fact, a name for the second time by Krotov in one referring to several species and begins to Anas platyrhynchos dom in Sachalin and be out of use in recent papers. We con­ 'CLERC, described as a new species Dicranotaenia sider D. elisae to be a synonym of D. :STODA, bisacculata Krotov, 1949. It is notewor­ acuminatus, as the type species of the � COM- thy that Ryzhikov (1963) summarising genus Diorchis, albeit its presence to­ GENUS results of helminthological research done gether with D. brevis is probably a mis­ in the Far East, indicates among potential take, or a very rare case that needs ex­ final hosts of this tapeworm 1202 speci- �lecka perimental confirmation. nd Parasi­ f Warsaw, Table 1. Records of C. pittalugai in representatives of the genus Aythya

o Dr. C. Region Host No of birds: Intensity Author(s) ·eparation infected examined ' Natural Sachalin A. marila 1 2 Krotov 1952 nstitut de Kamchatka A. marila 1 24 >200 Spassky, Bobova 1952 1 (Clerc, if.V. 6/64 Poland A.fuligula 1 130 8 our data �1". This Slovakia A. fuligula "rare" Macko, Birova 1985 )rm fr ag­ Turkmenia A. fuligula 1 13 16 Golovkova 1972 �ong con­ Chany Lake A.ferina 1 14 2 Krotov 1949 ,rphologi­ Kazakhstan A. ferina 5 32 1- 1237 Maksimova 1967 ·e among !ices: the Tadjikistan A. ferina 1 12 35 Borgarenko 1981 �-tm long, Canada A. affi nis 25 30 adult 1 - 7282 Hair 1975 ;ible coot Canada A. affi nis 12 52juv. 1 - 77 Hair 1975 58

times mens of the genus Anas and 195 speci­ counted after flotation by standard simult mens of the genus Aythya dissected in method. that region, in which only 3 times single Results: The peak of oocyst production is Ameri 1996) hosts were infected with C. pittalugai in the afternoon, between 16 and 19 (including the above mentioned report of o'clock in summer and between 17 and 19 infecte Krotov (1949), the remaining two see o'clock in autumn. The intensity of infec­ Swedi below). Until now nobody has confirmed tion is higher in the species of birds that suggel North the occurrence of C. pittalugai in the collect food on the ground than in those exclus final host of the genus Anas, but in Table collecting it in the air. Young birds have ticks, 1 authors who have found this tapeworm higher intensity and prevalence of infec­ patho! several times in representatives of the tion (some species with "island inhabita­ our o� genus Aythya are presented. tions" are an exception). At the beginning found According to the data above, Aythyini of autumnal migration, the infection ern b spp. are the most common hosts of C. prevalence and intensity are greatly contai pittalugai,and most intensively infected increased in all investigated species of dat< hosts are adults of A. affinis in Canada birds, and these parameters become lower y i at the end of the migration. In summer, dual ISOSPORA COCCIDIA (PROTO­ birds with a low level of fat are especially (12.5- ZOA, EIMERIIDAE) OF PASSER­ heavily infected, and in autumn, the birds ticks INE BIRDS ON THE COURISH with an average level of fatness have the Spit) afzelii SPIT highest intensity of infe ction. ricinu Conclusions: It seems that Isospora have Bo Olga Dolnik no great pathogenic influence on wild Zoological Institute, Russian Academy of COOOel passerine birds in nature, because a great Sciences, St. Petersburg, Russia migra1 number of these parasites in birds doesn't betwe� prevent them fr om accumulating fat and Objective: We studied the level of Iso­ lia ge also fr om migrating in the propitious spora infection in different species of durin� period. wild passerine birds in nature during of tic breeding period and autumnal migration. BORRELIA GENOSPECIES PREV A­ darkfi We had no possibility to dissect the hosts, LENCE AS A MARKER OF THE we e1 but we tried to estimate the harm of the POSSIBLE ROLE OF BIRDS IN very 1 infection for these birds in nature by THE DISTRIBUTION OF TICK­ of spiJ indirect data. BORNE SPIROCHETOSES IN THE cies p Materials and Methods: 518 passerine BALTIC REGION oppos birds of 50 species were trapped in mist were nets on the Courish Spit (=Curonian Spit) Helen V. Dubinina and Andrey N. Alekseev (25.2� in the Baltic Sea during summer and Zoological Institute, Russian Academy of the p: autumn 1995-1997. Droppings fr om Sciences, St. Petersburg, Russia garim every individual bird were collected at the mainl: moment of capture. Later the droppings Migratory birds were proven to be host­ activit were checked for Isospora oocysts. The reservoirs of the tick-borne borrelioses the sr number of oocysts in each sample was agents (Humair et al, 1993), and some- pl aine ·------·"''"

59

standard times even of more than one genospecies of pathogens, which can be transferred by simultaneously (Humair et al, 1997). birds during the spring and fall waves of Kluction is American authors (Nicholls, Callister, their migration. i and 19 1996) found 1 Borrelia burgdoiferi 17 and 19 infected Ixodes scapularis 8 years ago. TICKS (IXODOIDEA, IXODIDAE) r of infec­ Swedish authors (Olsen et al , 1995) OF MIGRATORY BIRDS ON THE birds that suggested that birds migrating from the KURISH SPIT 11 in those North were transferring mainly, if not Galina Efremova )irds have exclusively, B. garinii infected I. ricinus Institute of Zoology, Minsk, Belarus of infec­ ticks, whereas ticks had spirochetes inhabita­ pathogenic for human beings. Analysing Objective: To investigate the role of beginning our own results (Alekseev et al, 1998) we migratory birds in the carrying and infection found that I. persu!catus from the north­ feeding of Ixodidae ticks. � greatly ern beach of the Finnish Gulf mostly Materials and Methods: 14501 speci­ pecies of contained B. garinii (70-81 %, 1995-1997 mens of migratory birds (81 species) )fie lower y data) with a very high percentage of were examined on the Kurish Spit summer, dual infection: B. garinii and B. afzelii (=Curonian Spit) in the course of a seven especially (12.5-52%), whereas in the I. ricinus year period investigation in spring and in the birds ticks from the Kurish Spit (=Curonian autumn. 412 specimens of birds were : have the Spit) (Kalinigrad Region) mainly B. afzelii were found. Dual infection in I. infested with 1421 larvae and nymphs of Ixodes ricinus L. Jora have ricinus ticks was very rare. Results: The number of birds with ticks on wild Both localities of the survey are was higher in spring (2.9% of infected se a great connected by Italian-Spanish way of bird birds) than in autumn (0.8%). In spring, :Is doesn't migration. We stated the great difference the main hosts of I. ricinus were g fat and between I. ricinus tick infection by borre­ Fringilla coelebs L. (carried 47.8% of )ropitious lia genospecies in the spring and fall during the firstand second seasonal peaks collected ticks), Coccothraustes cocco­ of tick abundance: in the spring using thraustes (L.) (11.5%), Sylvia borin (Bodd.) (6.2%), and in autumn - Parus PREVA­ darkfleld technique of borrelia detection major L.(44.9%), Erithacus rubecula )F THE we estimated spirochete prevalence as (L.) (20.4%) and Turdus philomelos IDS IN very low, i.e. 7.3±0.9%, whereas 75.8% Brehm.(8.2%). The damage of legs of TICK­ of spirochetes were identified as genospe­ Fringilla coelebs Knemidocoptes cies pathogenic for human being. Just the by IN THE jamaicensis opposite: in the fall spirochete prevalence was recorded. Virologists were 3.5 higher than in the spring have isolated four strains of TBE virus from the blood serum of P. major and Alekseev (25.2%), but ticks contained only 9.5% of Turdus merula ademy of the pathogenic borrelia genospecies. B. L. as well as from the internal organs of Phoenicurus phoenicu­ garinii prevailed in I. persulcatus (North) rus mainly in the second half of their seasonal (L.). Conclusions: be host­ activity whereas in I. ricinus (South) - in The discovery of arbovi­ )ffelioses the spring. All these facts might be ex­ ruses in the blood and internal organs of 1d some- plained by the differences of the sources migratory birds in spring testify to the 60 BLOC possibility of carrying the infection from L., Gamasina (5 species ) and I. lividus. (PRO wintering grounds. ricinus, carried by Antihemagglutinins to the West Nile, I. AND migratory birds, may be responsible for Sindbis, Uukuniemy and TBE viruses BIRD some fe atures in the distribution of arbo­ were discovered in migratory birds in virus infections. It should be taken into spring (Gembitsky, 1989). V. ( consideration that the period of the activ­ Conclusions: The discovery of the arbo­ Zehtin ity of I. ricinus coincides with the period viruses and antibodies to the viruses in 1Institl of bird's seasonal migrations. As the migratory birds in spring is the witness of Scienc preimaginal stages of I. ricinus parasitize the possibility of carrying infection from Ecolog on the birds, they can be carried in other wintering grounds. The activity of para­ Vilniu regions. In connection with the discovery sites of pasture and the bird's nest-burrow of TBE virus in migratory birds, this fact type begin exactly at this time, and the Obje( indicates the possibility of introducing transmission of the arboviruses from record arboviruses in the local biocenoses as infected birds to blood-sucking arthro­ the av well as the formation of new and the pods is possible. The nest-burrow para­ far, th maintenance of existing natural foci of the sites of synurbic and colonial species of of the arbovirus infections. birds, which repeatedly settle the nests, preser are particular dangerous. The greatest and fa PARASITIC ARTHROPODS OF quantity of blood-sucking arthropods birds 1 BIRDS AND THEIR ROLE IN were registered in the nests of Sturnus Mate1 MAINTENANCE OF NATURAL vulgaris L., Calumba livia L., Delichon birds ! FOCI OF ARBOVIRUS DISEASES urbica (L.), Riparia riparia (L.).The invest IN BELARUS isolation TBE virus from Gamasina ticks were (in summer) and the isolation of the virus menta Galina Efremova andA. Gembitsky seven years later from I. lividus, which of Z Institute of Zoology, Minsk, Belarus was collected from the nests of R. riparia Scienc of the same settlement, indicate the during Objective: To investigate blood-sucking stability of natural foci of TBE virus. The Septet arthropods of bird in Belarus. discovery of TBE virus in spring before releas1 Materials and Methods: In 1973-1996, the arrival of birds showed that the clippi1 parasitic arthropods from 1590 bird nests infection reserved in the diapaused Ixodi­ air-dri were collected and studied. 2195 speci­ dae ticks. The results show that the bird stainet mens of birds were examined, and 435 ectoparasites have the potential possibil­ of 10< specimens of the birds were recorded to ity for long-term preservation and dis­ low (> be infested with 1714 specimens of blood­ semination of the infection agents in the tives. sucking arthropods. natural foci of diseases in Belarus. Resul Results: Ixodidae ticks (Ixodes ricinus ex ami L., I. arboricola Schulze and Schlottke cent) • and I. lividus Koch ) were prevailing or me among the ectoparasites (9 1.2% of col­ least, ' lected arthropods). Virologists have to th isolated the strains of TBE virus from Haem Lullula arborea (L.) Coracias garrulus cytozc 61

BLOOD PROTOZOAN PARASITES Kinetoplastida (Trypanosoma) have been lividus. (PROTOZOA: KINETOPLASTIDA recorded. Mixed infections of birds with st Nile, AND HAEMOSPORIDA) OF WILD two or more genera of the protozoans viruses BIRDS FROM BULGARIA were common. Most common blood birds in protozoans were Haemoproteus belopol­ 1 1 V. Golemansky , D. Pilarska , Pavel skyi, Plasmodium (Haemamoeba) sp. he arbo­ 1 2 2 Zehtindjiev , G. Valkiunas and T. Iezhova and Trypanosoma everetti. Our prelimi­ ruses in 'Institute of Zoology, Bulgarian Academy of nary findings suggest a similarity in itness of Sciences, Sofia, Bulgaria; 2Institute of fauna of avian blood protozoans in Bul­ on from Ecology, Lithuanian Academy of Sciences, garia and in adjacent territories of of para­ Vilnius, Lithuania Europe. However, large samples are .-burrow needed before conclusions can be drawn and the Objective: Relatively little has been regarding ecology and fauna of all genera :s from recorded concerning the haematozoa of of the parasites in the country. arthro­ the avifauna of the Balkan Peninsula. So w para- far, there is no information on distribution THE PREVALENCE OF SARCO­ 1ecies of of the parasites in Bulgaria. This paper CYSTIS IN SOME WILD BIRDS 1e nests, presents results of a study on prevalence AND POULTRY IN LITHUANIA greatest and fauna of blood protozoan parasites of AND IN NEIGHBOURING TERRI­ hropods birds from the Northern Bulgaria. TORIES Stumus Materials and Methods: A total of 63 birds of 11 species of Passeriformes were >elichon Jadvyga Grikieniene and Tatjana Iezhova :L.).The investigated for blood parasites. The birds Institute of Ecology, Vilnius, Lithuania na ticks were sampled at the Biological Experi­ mental Station "Kalimok" of the Institute he virus Objective: No published records are of Zoology (Bulgarian Academy of , which available of Sarcocystis in muscles from Sciences), Nova Cherna, Tutrakan, riparia wild birds or poultry in Lithuania or during the period from June, 1993 to ate the neighbouring countries. Our objective September, 1994. Captured birds were us. The was to determine the prevalence of Sar­ released after blood was obtained by : before cocystis spp. infection among some wild clipping a claw. All blood smears were hat the birds and poultry in Lithuania and in air-dried, fixed in absolute methanol, j Ixodi­ South Curonian Spit (Russia). stained with Giemsa' s stain. A minimum :he bird Materials and Methods: Three hundred of I 00 fields were examined under both IOSSibil­ and seventy five wild birds of 45 species low (x25) and high (x 1 power objec­ nd dis­ 00) belonging to 5 orders (Passeriformes, tives. ; in the Piciformes, Charadriiformes, Columbi­ Results and Conclusions: A total of 17 formes and Strigiformes) and 97 poultry examined passeriform birds (27 .0 per (76 hens and 21 turkeys, Galliformes) cent) of 6 species were infected with one were examined for the presence of sarco­ or more species of blood Protozoa. At sporidia in muscles. Birds were obtained least, 9 species of the parasites belonging from a number of sources including birds to the orders Haemosporida (genera collected for other studies by various Haemoproteus, Plasmodium Leuco­ and researchers at the Institute of Ecology cytozoon), Adeleida (Hepatozoon) and 62 ON 1 during the period 1986-1997. In many on a wide range of mammals, birds and BRIAJ cases, frozen carcasses were used. reptiles. Passerine birds are important HYMl Skeletal muscle samples from pectoral, hosts for immature Ixodes ricinus (L.). In TION wing, leg or neck muscles were examined order to determine the possible role of microscopically in a compressor after avian hosts in the maintenance of this Barbar; staining them with methylene blue. Some pathogen in nature, we examined birds Cielec� of the samples were examined by means blood for the presence of Borrelia Departl of squash preparation. 16 passerine birds burgdoiferi. tology, were investigated histologically. Materials and Methods: Birds were Warsa\ Results and Conclusions: The preva­ captured on April, May, June, August lence of Sarcocyctis spp. infection among and September 1996, in the Masurian The f< investigated wild birds and poultry was Lake region (North-East Poland), proved genus ferina, low. Sarcocysts were detected only in 3 to be an area where Lyme disease is passerine birds of 339 investigated - 1 of endemic. Small volumes of birds' blood nest, a 58 robin Erithacus rubecula, 1 of 6 pied were obtained from brachial vein and on pas flycatcher Ficedula hypoleuca, and 1 of 2 analysed for the presence of Borrelia of the. and A. fieldfare Turdus pilaris. All the infected burgdoiferi spirochetes by the nested­ birds were collected in South Curonian PCR technique (a DNA fragment con­ Cielecl Spit. The species of Sarcocystis in pas­ taining the hbb gene of Borrelia specie1 serine birds were not identified as no burgdoiferi sensu lato was amplified). The et cystozoites were examined. Out of 76 Results: A different tick infestation rate are pa investigated fowl, sarcocysts were found was observed in birds in dependence of oncosr only in one free-ranging cock A radially species. The most parasitized by ticks from 1 specie1 striated cyst wall, 2.5-3 J.!m thick, and the bird species belonged to three families Howe' banana-shaped cystozoites, 9-11 J..Lm in Turdidae, Fringillidae and Troglodytidae presef1 size, were morphologically similar to were chosen to investigations. A total of and m those of the species S. horvathi known for 152 blood samples were analysed and becom the fowl. This is the first record of muscle positive results were obtained in 6 cases. identif cysts of the genus Sarcocystis in birds in All the birds in which Borrelia burgdor­ cases s Lithuania and in South Curonian Spit. fe ri sensu lato spirochetes were detected belong to species chaffinch (Fringilla In t all fou OCCURRENCE OF BORRELIA coelebs). There was only 24 individuals hosts BURGDORFERI SENSU LATO (15,8%) of this species, and infection rate materi: SPIROCHETES IN PASSERINE was 25%. derive< BIRDS Conclusions: Statement of Borrelia rian burgdoiferi presence in passerine birds I Alicja Gryczynska and Andrzej Zgodka blood confirmed that birds can serve as Slonsk Uniwersity of Warsaw, Warsaw, Poland natural reservoir of spirochetes on the Fish pc Th( study area. They can be a bacteria source Objective: In Europe, the etiological for ticks engorging on them and play an specie1 agent of Lyme disease - Borrelia burg­ important role in Lyme disease dissemi­ has be doiferi, is transmitted primarily by the nation. contac tick Ixodes ricinus (L.) which may fe ed cyclop ·�------'*'"

63

ON THE SPECIFICITY OF FIM­ and Macrocyclops albidus, or the ostra­ birds and BRIARIA SARCINA LIS (CESTODA: cod Heterocypris incongruens. It is important HYMENOLEPIDIDAE) IN RELA­ probable that the packets of eggs of F. rs (L.). In TION TO HOSTS sarcinalis are too large to be ingested by � role of small crustaceans of this kind. In line � of this Barbara Grytner-Zi�cina and Danuta with this, potential intermediate hosts led birds Cielecka should perhaps be sought among much Borrelia Department of General Biology and Parasi­ larger representatives of the Amphipoda tology, The Medical University of Warsaw, or Oligochaeta, which deep-diving Aythya ·ds were Warsaw, Poland ducks would be capable of taking, along , August with silt from the bottom of a body of \1asurian The four species of diving duck of the water. ), proved genus Aythya occurring in Poland are A. F. sarcinalis has not yet been re­ isease is fe rina, A. nyroca and A. fuligula, which corded from ducks other than Aythya ls' blood nest, and A. marila, which is only found species. vein and on passage and in winter. It was from two Borrelia of these duck species, Aythya fu ligula THE ECOLOGY OF THE HAEMA­ : nested­ and A. nyroca, that Grytner-Zi<::cina and TOZOAN PARASITES OF EURO­ lent con­ Cielecka described a new Fimbriaria PEAN BLACKBIRDS TURDUS Borrelia species, named F. sarcinalis, in 1994. MERULA, AND THEIR EFFECTS fled). The characteristic features of the species ON HOST FITNESS tion rate are packets of eggs containing up to 30 oncospheres, which differentiate it clearly dence of B.J. Hatchwell1, M. Anwar2, M.J. Wood1, D. F. fa sciolaris Fimbriaria by ticks from - the only Chamberlain2 and C.M. Perrins2 families species known for nearly 200 years. 1Department of Animal and Plant Sciences, lodytidae However, on the basis of analysis of The University of Sheffield, U.K.; 2Edward . total of preserved material collected in Poland Grey Institute, Department of Zoology, The 'Sed and and made available by other authors, it University of Oxford, U.K. 6 cases. becomes clear that tapeworms originally )urgdor­ identified as F. fa sciolaris are in many The haematozoan parasites of a rural detected cases specimens of F. sarcinalis. population of European blackbirds Tur­ r

soma, Atoxoplasma and Microfilaria INVESTIGATIONS INTO BIRD ons from were recorded in birds. M. f. flava can be PARASITOSES IN ESTONIA There are easily distinguished from the other two the mor­ subspecies by a significantly higher Toivo Jarvis , different prevalence of Haemoproteus (48.9 %, Estonian Agricultural University, Tartu, misdeter­ Estonia 95-% confidence limit is 34.3-63.6; the ;ycle and same parameters are 12. 1 %, 3.1-24.2 in l. Moreo­ Objective: To present a review about M. f. fe ldegg and 17.0 %, 8.5-30.0 in M. :mtion of investigations on bird parasitoses in f. th unbergi). Moreover, Haemoproteus eflect the Estonia. anthi clearly dominates in M. f.flava, and :tcter) of Results: The most thoroughly investi­ it is rare in other investigated subspecies. cation in gated bird parasitosis in Estonia is M. f. thunbergi has an exceptionally high contact chicken eimeriosis. This research has prevalence of Leucocytozoon (22.6%, been carried out during many years under 13.1-37.0; the same parameters are 2.2, nt of the the leadership of J. Parre. Studies on 0.0-8.7 in M. f. flava and 0.0, 0.0-2.8 in IGA MZ eimeriosis in chicken have been carried M. f. fe ldegg). Species of Plasmodium r Grant out by E. Olkonen, T. Schattschneider, are especially rare in M. f. thunbergi (5.7 H.-E. Simovart a. o. in the years 1968- %, 1.1-14.4) and most common in M. f. 1993. The following problems have been fe ldegg (42.4 %, 24.3-56.8). >GICAL treated: 1. Haematologic changes after Conclusions: During late stages of spring PECIES invasion with Eimeria tenella in nonim­ migration, subspecies of M. flava can be munized and immunized chicks. 2. Influ­ distinguished by prevalence of infection ence of superinvasion on the protein lkiunas and some peculiarities of the fauna of composition of the blcxxl serum in chicks ani a haematozoa. The investigated birds have in the case of eimeriosis caused by E. similar wintering places (Africa and tenella. 3. Changes in lymphoid organs of ew signs South Asia), but different breeding areas. chicken in the case of infection with E. pecies is Prevalence of Plasmodium infection is tenella. 4. Anticoccidial immunity in � parasi- especially high in birds breeding in the poultry flocks. 5. The manufacture and for this South Palearctic (M. f. fe ldegg), Leuco­ use of chicken eimeriosis polyvaccine. 6. rk is to cytozoon in birds breeding in the North Elaboration of control measures against ection of Palearctic (M. f. thunbergi), and Haemo­ chicken eimeriosis. The helminthofauna Wagtail proteus in birds with an intermediate of domestic duck and goose has been a during position of the breeding areas in compari­ thoroughly investigated by L. Veldemann son to those mentioned above. Among the (1950-1 960). 73% of ducks and 48% of Jecimens investigated subspecies, M. f. flava is the geese were invaded with parasites. 29 tbspecies main host for H. anthi. The mass investi­ species of parasites have been diagnosed ration in gation of birds during late stages of in ducks, 15 species have been found in ay 1986 spring migration is a promising technique geese. The most important parasitoses trs tech­ for bird subspecies studies. "eldegg ­ were ducks tetramerosis, geese amidos­ tomosis, ducks notocotylosis, geese and -gi - 53. ducks ganguleterakidosis, geese and 'Proteus, ducks eimeriosis. "rypano- 66 CER( The results of the investigations into Materials and Methods: Adult female EFFH flukes (Trematcxla) of water and shore mites were stimulated with controlled IDEN birds on the western coast of Estonia have vibrations, both with and without prior BILH, been presented by V. Jogis (1959). 52 stimulation with C02• Experiments were birds belonging to 27 species (5 orders) conducted in the laboratory under differ­ Sonja l were examined and 57 species of flukes, ent light intensities. Depart belonging to 19 families, were recorded. Results: Under "night" conditions (light Kiel, G Observations on the epidemiology of intensity40-50 lux), stimulation with nitrate the avian tapeworm Schistocephalus C02 will not act as an activator but rather (SEM) solidus in the dog (Jarvis, 1994). as a primer so that a subsequent stimula­ their n Conclusion: For the most part the inves­ tion with low-level vibrations will acti­ Result tigations into bird parasitoses in Estonia vate the mites but only for the duration of and T. have resulted from practical need<; the vibrations. Mites already active will son of (economic interest) and are related to respond to C02 with a "stopping" reac­ with tJ domestic birds. tion; the mites stop and remain motionless and, a: unless further stimulated with vibrations. related BEHAVIOURAL RESPONSE OF This behaviour is interpreted as a defence signifi< THE CHICKEN MITE, DERMA­ against being eaten by the potential host. distrib1 NYSSUS GALLINAE, TO HOST­ A blow of C02 could mean that the bird is papilla RELATED STIMULI close enough to see the mite moving nitrate whereas vibrations indicate that the bird light m Ole Kilpinen is moving and thus its attention is not Conch Danish Pest Infestation Laboratory, Lyngby, focused on the mite. cate th Denmark Conclusion: D. gallinae can detect sents a potential hosts by means of expired COz related Objective: Dermanyssus gallinae is a and vibrations of the substrate induced by inadeq1 well-known ectoparasite of domesticated host movements, but the behavioural closely poultry and a wide range of wild birds. It response depends on thelight intensity. studyir spends most of the time hidden in cracks neither and crevices, and will only come out to evaluat feed when a potential host is near. The fixatioJ question addressed in this study was how Thus, 1 the mite detects the presence of a poten­ tions c tial host. 67

CERCARIAL CHAETOTAXY - AN ensure the collection of data that is com­ tlt female EFFICIENT MEANS FOR THE parable between studies, even though controlled IDENTIFICATION OF TRICHO­ some argentophilic structures are not tout prior BILHARZIA SPECIES? visible in SEM. ents were !er differ- Sonja Kock and Wolfgang Boeckeler ONTOPHYLOGENETIC DIFFER- Department of Zoology, University of Kiel, ENTIATION OF CYSTICERCOIDS :ms (light Kiel, Germany llinae is Georgiy Krasnoshchekov and Nina Tomi­ 1, and a Objective: In order to evaluate the im­ lovskaya jon will portance of cercaria! chaetotaxy in the Institute of Ecology of the Volga River t the best identification of Trichobilharzia species, Basin, RAS, Togliatti, Russia i is 5-10 papillary patterns of the European avian litial C02 schistosomatids T. ocellata and T. franki Objective: The study of ontophylogenetic to be 50- were examined and compared. adaptations of larval cestodes of birds. tivate the Materials and Methods: Sensory papil­ Material and Methods: Investigations t1S (light lae were revealed by staining with silver were conducted on seven types of cys­ ion with nitrate. Scanning electron nlicroscopy ticercoids from the families of Hymeno­ mt rather (SEM) was also employed to confirm lepididae (14 species) and Dilepididae (6 : stimula­ their number and distribution. species). Larvae (mature, developing and will acti­ Results: The chaetotaxy of T. ocellata transplanted into an intermediate host) Iration of and T. jranki were identical. A compari­ were studied based on electron nlicro­ �tive will son of the papillary pattern of T. ocellata graphs and histochemical techniques. 1g" reac­ with the pattern of some non-European Results: At the early stages of develop­ totionless and, as such, presumably more distantly ment, cysticercoids are covered with ibrations. related Trichobilharzia species, revealed primary tegument. It is characterised by 1 defence significant differences in the number and the presence of a nlicrovilli border and tial host. distribution of sensilla. Some sensory lack of specific inclusions. Mature larvae ie bird is papillae stained insufficiently with silver retain such a tegument on their tails only. moving nitrate or were difficult to exanline using The tail structure is essentially distinct in the bird light microscopy. different types of cysticercoids. It is most 1n is not Conclusions: The obtained results indi­ simply arranged in essentially distinct in cate that papillary pattern analysis repre­ different types of cysticercoids. It is most 1 detect sents a method of discrinlinating distantly simply arranged in monocercus which ired C02 related species of Trichobilharzia but is follicles are formed by the tegument only. juced by inadequate for the discrinlination of The hypertrophy of microvilli and intensi­ 1avioural closely related species. However, when fication of the membrane secretion from :nsity. studying the chaetotaxy of cercariae their tops take place after the transplan­ neither the possibility of subjective tation. The basic function of the tail is to evaluations nor the nlisleading effects of protect larvae from the host's hemocytes. fixation artefacts can be totally excluded. It is effected by the cytolytic properties of Thus, the use of SEM in future exanlina­ tegumental membranes. The cyst struc­ tions of Trichobilharzia chaetotaxy will ture is most versatile. Specific distinc- 68 helmit tions show up in the tegumental structure, capillaria eggs in the faeces of birds. In brane amount and thickness of fibrous layers, the intermediate host - the earthworm - number of muscle and secretory cells, larvaehatch and moult once. In the intes­ T1 presence of a pseudo-myeline layer. The tine of a definitive host ( rook ) larvae gation basic fu nctions of the cyst include the moult three times and in about 30 days ne mat protection of larvae in the digestive tract, become mature. C. corvorum parasitises prono activation and evagination of the scolex. in the small intestine of a definitive host necrot Differentiation of the scolex tegument for about 7-9 months and produces eggs. celluh occurs after the invagination. It is fol­ After that, the helminths are eliminated of the lowed by the replacement of the microvilli fromthe intestine of birds. for microtriches, as well as by the accu­ 38 two-three-week-old rooks (C. ON 1 COCJ mulation of inclusions in the syncytium frugilegus L.) were used in the experi­ matrix. Thus, the cyst ensures the condi­ ment. The rooks were infected with the IN L1 tions for progressive evolution of the tail nematode corvorum L. by feeding C. Liuda and its transformation into an additional them on the earthworms. The experi­ Aniol< protective envelope. mental birds were decapitated every day Paulat Conclusion: The comparative study of over the period of 7-31 days after the 1Instit1 cysticercoids has shown that they are a infection. Some pieces of the small intes­ 2Vytat progressively developing type of larval tine were taken for a micromorphological Lithua cestodes of birds. The intensification of investigation, and histological prepara­ the protective function of larval elements tions were made in an ordinary way. Objec can be associated with a migrant way life At the early stages (on the 7th and birds of the definitive hosts. 11th days) of infection the catarrhal type invest of inflammation in thetissues of the small remai MICROMORPHOLOGICAL ALTE­ intestines was observed whereas from the specie RATIONS IN THE ROOK (CORVUS 12th to the 25th day a strongly pro­ was FRUGILEGUS L.) INTESTINE DUR­ nounced cellular infiltration on the mu­ Sarco ING EXPERIMENTAL CAPIL­ cous membrane of the small intestine was Anser LARIOSIS noticed. Lymphocytes, histiocytes and nets i eosinophils were predominating. In many Cur or Ona Kublickiene places there was a distinct epithelial 1997 Vilnius University, Vilnius, Lithuania desquamation; the epithet cells were Mate; peeled off and localized in the lumen of micro The purpose of this work was to study the the intestine. At this period of infection, belon1 developmental dynamics of micromor­ C. corvorum was localised deeply in the white phological changes in the rook intestine mucosa of the intestine wall, which bean 1 during experimental capillariasis. The caused distinct inflammatory breaches (Mer£ dynamics of the alterations in the small and petechiae. penelj intestine of experimental rook infected On the 30th day after the infection tufted with Capillaria corvorum (Rudolphi, helminths reached their puberty and were (Tadc 1819) Travassos, 1915 was investigated. localised in the opening of the small platyt It was established (Slikas Kasper­ & intestine of the bird. However, the were sone, 1978) that larvae are formed in the deterr 69 f birds. In helnliths fastened to the mucous mem­ sarcocysts in 28 muscle sections stained :ihworm - brane with their front end. with methylene blue. The morphometric the intes­ The results of the histological investi­ analysis of cyst walls and cystozoites was gations have shown that the infection with performed in native preparations. < ) larvae t 30 days nematoda Capillafi a corvorum has a Results and Conclusions: Sarcocysts Jarasitises pronounced destructive effect causing were detected in 10 (40%) of investigated litive host necrotic-degenerative processes and birds, i.e. in 5 white - fronted geese, in 4 1ces eggs. cellular infiltration in the small intestine mallards and in 1 shelduck. On the basis �linlinated of therook. of morphometric analysis of cyst walls and cystozoites two sarcocyst types were SAR­ )()ks (c. ON THE DISTRIBUTION OF detected in both geese and ducks. The COCYSTIS 1e experi­ IN WATERFOWL BIRDS obtained data testify to high infection . with the IN LITHUANIA level of waterfowl birds. y feeding l l Liuda Kutkiene , Jadvyga Grikieniene , DISTRIBUTION OF CHAFFINCH e experi­ 1 2 Aniolas Sruoga · and Algimantas HAEMOPROTEIDS AND THEIR �very day Paulauskas 1'2 POSSIBLE VECTORS ON THE after the 1Institute of Ecology, Vilnius, Lithuania; CURONIAN SPIT aall intes­ 2Vytautas Magnus University, Kaunas, )hological Lithuania GediminasLiutkevieius prepara­ Institute of Ecology, Vilnius, Lithuania vay. Objective: Sarcosporidia in waterfowl : 7th and birds in Europe are not yet sufficiently Objective: Haemoproteids of chaffinch Thai type investigated. Also, some uncertainties still Fringilla coelebs L. are unique objects the small Sarcocystis remain in differentiating for investigations of vectors of Haemo­ from the species. The objective of tllis research proteus on the Curonian Spit. The ecol­ 1gly pro­ was to investigate the occurrence of ogy of chaffinch and the fauna of the the mu­ Sarcocystis in waterfowl birds (order haemoproteids in tllis bird have been stine was Anseriformes) accidentally stuck in fish­ investigated exceptionally well here. ytes and nets in the Nemunas delta and in the Prelinlinary data on the distribution of In many Curonian Bay (Lithuania) in the period of chaffinch Haemoproteus and their possi­ epithelial 1997 - 1998. ble vectors are given in tllis presentation. Us were Materials and Methods: Under the light Materials and Methods: 146 specimens lumen of nlicroscope we have investigated 25 birds of chaffinch were investigated by blood infection, belonging to the following eight species: 7 smear technique on the Curonian Spit in ,ly in the (Anser albifrons), 1 white - fronted geese the Baltic Sea during May - August, which bean goose (Anser fa balis), 3 goosanders l, 1997. Ceratopogonids (genus Culicoides) breaches (Mergus merganser), 2 wigeons (Anas were collected several times per week penelope), 1 pochard (Aythya fe rina), 1 using standard methods. infection tufted duck (Aythya fuligula), 1 shelduck Results: Species of Haemoproteus, and were (Tadoma tadoma) and 9 mallards (Anas Trypanosoma and Microfilaria were 1e small platyrhynchos). Head and neck muscles recorded in chaffinches. Haemoproteids ver, the were exanlined. Infection intensity was were donlinant parasites. Haemoproteus deternlined in compressor by counting 70 values fringillae, H. dolniki and H. magnus vaccination of Japanese quails (Coturnix ganisn were recorded. The former two species coturnix japonica) on antibiotic and quails were most common. The overall preva­ adhesive activity of microflora of the erably lence of Haemoproteus infection was digestive tract of Japanese quails have microf 36.2 % (95 %-confidence limit is 26.1 - been investigated. MAl 48.9) in adult birds and 14.3 % (8.0 - Materials and Methods: Two groups of quails 23.3) in juveniles. This testifies the active J apancse quails were compared. The first nated < transmission of the parasites on the study group was vaccinated twice with inacti­ digesti site. Six species of Culicoides (C. arcan­ vated Salmonella enteritidis Talovac 109 (35 .Oo/. tidus, C. obsoletus, C. punctatus, C. SE vaccine, and the second (control) do mine pinctipennis, C. segnis and C. impunc­ group was unvaccinated. The composi­ in 50. tatus) were recorded feeding on birds. C. tion of bacteria of the digestive tract of mediut impunctatus was the most abundant Japanese quails was determined by mediut species (over 90 % of all collected speci­ method of Bergy, and the antibiotic MAl V mens of Culicoides). The highest density activity was investigated by method of MAl "' of ceratopogonids was in June. In July, Ayzeman and Egorov. Adheric features Conch the density of Culicoides decreased of microorganisms of digestive tract of tract o dramatically, and only C. impunctatus quails were investigated using express­ otic an was recorded in the end of July. Culicoi­ method. Adherence was regarded to be nil tion of des were not found feeding on birds in when mean adherence index (MAl) is 0- enterid August. 1.0; weak - MAl = 1.01-2.0; medium - activit) Conclusions: Especially active transmis­ MAl = 2.01-4.0, strong - MAl=> 4. immun sion of Haemoproteus takes place in June Results:Antibiotic activity of microflora on the Curonian Spit. Experimental work of the digestive tract was very high. on vectors of the parasites should be Antagonistic activity was 78.26% of the LOW recommended during this period of sum­ bacteria in unvaccinated birds, and this INFES mer. C. impunctatus is incriminated to be parameter was 91.66% in vaccinated TIT P} one of the possible important vectors of birds. The majority of the investigated species Haemoproteus on the Spit. bacteria were capable to inhibit the Tomasz development of several bacteria. A zone K�dra2 RELATIONSHIP BETWEEN IM­ of inhibition was 20-30 mm. In the vacci­ 1 DepartJ MUNITY OF JAPANESE QUAILS nated quails in comparison to unvacci­ AND ACTIVITY OF MICROFLORA nated ones, the antagonistically active Warsav,. Parasito OF THE DIGESTIVE TRACT bacteria were especially common, the Warsav,. zone of inhibition was considerably Vanda Lubianskiene, RataJankauskiene and higher and the spectrum of the inhibitory Objectl Vesta Skrodenyte activity was wider. The higher antagonis­ expecte Institute of Ecology, Vilnius, Lithuania tic bacteria activity against Staphylococ­ contani cus aureus have been recorded in vacci­ Objective: spring, Due to antibiotic and adhesive nated quails. However, the bacteria breedin; activity, microflora of the digestive tract nearly completely inhibited the develop­ low pre can influence the processes of macroor­ ment of Candida albicans and Bacillus ganism immunity. The impact of the subtilis in unvaccinated quails. Mean 71

; (Cotumix values of adherence of intestinal microor­ Materials and Methods: New nestboxes ganisms in vaccinated and unvaccinated were used on two forest areas biotic and quails indicate that vaccination consid­ (Kampinoski National Park in 1995 and )fa of the erably increase the ability of normal Mlociny Forest Park in 1996) near War­ tuails have microflora to adhere the intestinal walls. saw, Poland. During breeding season MAI in the intestine of unvaccinated nesting species were identified, and on the >groups of quails was 1.65, and it was 2.84 in vacci­ day of fledging, nests were collected and l The first nated ones. Among microorganisms of the analysed for the occurrence of ectopara­ vith inacti­ digestive tract of unvaccinated quails, nil sites. alovac 109 (35.0%) and weak (35.0%) MAI were Results: Examination of nests of hole­ i (control) dominate, and strong MAI was recorded nesting species on both study areas � composi­ in 50.0% of intestinal microflora and revealed that prevalence of adult fleas ve tract of medium in 25.0%. In vaccinated quails, (Ceratophyllus gallinae) in Blue Tits mined by medium (42. 1%) and strong (36.8%) nest is always lowest (KPN: P. caeruleus antibiotic MAI were most common, whereas weak - 20%, Great Tit P. major - 43%, Pied method of MAl was recorded in 21.0% of bacteria. Flycatcher F hypoleuca - 32%; P. ic features MFP: Conclusions: Microflora of the digestive caeruleus - 13%, P. major 25%, Starling re tract of tract of Japanese quails has high antibi­ S. vulgaris 82% ). The average number of g express­ otic and adhesive activity. The vaccina­ adult fleas in Blue Tits nests was similar xito be nil tion of Japanese quail s with Salmonella to that of Great Tit (KPN: Blue Tits - 13, vfAI) is 0- enteridis effects both the microflora Great Tits - 14.5; MFP: Blue Tits - 7, medium - =>4. activity of the digestive tract and the Great Tits - 3) and either greater than (in immunity of birds. KPN Pied Flycatchers - 2.5) or lower microflora than (in MFP Starlings - 27.5) other very high. migratory species. 6% of the LOW PREVALENCE OF FLEA Conclusions: The intensity of infestation , and thi s INFESTATION IN NESTS OF BLUE with fleas was similar in both species of vaccinated TIT PARUS CAERULEUS Tits, but the prevalence of fleas was tvestigated lower in Blue Tits nests. Apparently, 1hibit the 1 Tomasz D. Mazgajski _and Aleksander H. when Great Tits are more numerous than a. A zone K�dra2 Blue Tits, the latter has limited access to the vacci­ I Department of Ecology, University of areas with old nesting sites - where the > unvacci­ 2 spring infections with after-wintered adult lly active Warsaw, Warsaw, Poland; Institute of Parasitology, Polish Academy of Sciences, fleas occur. The average number of adult lffion, the Warsaw, Poland fleas in Blue Tits nests depended on the nsiderably presence of particular migratory species inhibitory Objective: Blue Tits Parus caeruleus are of hole-nesters - Pied Flycatchers in KPN antagonis­ expected to be particularly exposed to phylococ- started breeding later than Tits and were contamination of fleas during early exposed to fleas in much lower degree, in vacci- spring, mostly because of their early but Starlings in MFP started breeding at bacteria breeding activity, but many studies reveal the same time as Tits, and - due to their : develop­ low prevalence of fleas in Blue Tits' nest. ! Bacillus dominance, and aggressive behaviour - limited Tits mobility or territory selec- Is. Mean 72 tion, and consequently chances of infec­ Results: The average level of coccidian to COli and t\\' tions with fleas. infections of nestlings didn't affect the average weight of the fledging nestlings Mater PARENTAL FEEDING EFFORT (Rs=-0.20, n= 19, p=0.40). However, were 1 AFFECTS BOTH NESTLING parental feeding effort, correlated posi­ (in th QUALITY AND ENDOPARASITE tively with average weight of nestlings before before INFECTION LEVELS (Rs=0.53, n= 19, p=0.03), and negatively collect with average level of coccidian infection sites. , Tomasz D. Mazgajsld1 and Aleksander H. (Rs=-0.50, n= 19, p=0.04). 2 and in Kydra Conclusions: The quality of nestlings is I built affected by the quantity of food they Department of Ecology, University of nestbo 2 receive, but surprisingly, the infection Warsaw, Warsaw, Poland; Institute of level with orally transmitted I. lacazei is Result Parasitology, Polish Academy of Sciences, fleas , Warsaw, Poland decreased by the increase of the parental feeding effort. We suggest that expected nests i that c Objective: Besides investigation of the increase in I. lacazei infections could have been masked by better immune fleas relationship between Starling Stumus A ea rh response of better-fed nestlings. Addi­ vulgaris nestlings' weight and levels of group� infection with endoparasite Isospora tionally, as parental feeding effort seems to be the causative factor here, it should old ne lacazei (Coccidia: Apicomplexa), the phaga be taken into account in further studies of effect of the third factor - parental feeding Thl endoparasite effect on nestlings - just like effort was analysed. The food compensa­ was g it was suggested in ectoparasite studies. tion of the effect of ectoparasites was boxes previously suggested, but in the case of Conell orally transmitted endoparasites, any ECTOPARASITE DIVERSITY IN ence tl increase in feeding effort may also bear STARLINGS' NESTS IN RELA­ nests 1 increased risk of transmission. TION TO THE AGE OF THE NEST­ SITE were r Materials and Methods: Starling nes­ at the tlings were weighed on the 17th day of I were 1 life. At the same time, samples of excre­ Tomasz D. Mazgajsld , Aleksander H. 2 I when 1 ment were taken from most nestlings - Kydra and KrzysztofKowalczyk I materi and promptly preserved in 2% K CrO . 2 7 Department of Ecology, University of winter 2 Shortly afterwards, samples were ana­ Warsaw, Warsaw, Poland; Institute of and ir lysed under light microscope (x40). Parasitology, Polish Academy of Sciences, crease� Average levels of coccidian infections Warsaw, Poland materi were calculated for each nest. Parental sites. � feeding effort - the average number of Objective: As studies of the composition nestsit• feeding trips per nestling per hour - was of ectoparasite fauna in nests of Starling greatl) estimated using two subsequent, 30- Stumus vulgaris didn't take into account causin, minutes observations of feeding trips on neither the age of the nest-site nor the both s< the 15thday of nestlings' life. time of nest collection, our study attempt 73

to compare the infestation of the one-year FEATHER MITES: GENERAL coccidian and two-year-old nestboxes. MORPHOLOGICAL ADAPTA- affect the New nestboxes AND nestlings Materials and Methods: TIONS, PHYLOGENY were placed in the Mlociny Forest Park However, COEVOLUTIONARY RELATION­ (in the vicinity of Warsaw, Poland) tted posi­ SHIPS WITH BIRDS before the 1996 breeding season. Just nestlings before Starlings fledged, nests were Sergei V. Mironov 1egatively collected and analysed for the ectopara­ Zoological Institute, Russian Academy of infection Sciences, Saint Petersburg, Russia sites. Some nestboxes were not cleaned, and in 1997 we collected nests that were �stlings is Feather mites (FM) are highly specialised built both in cleaned and not-cleaned 'ood they permanent obligatory ectoparasites nestboxes. infection associated with almost all recent orders of Results: In one-year-old nestboxes mostly lacazei is birds (Aves). According to recent concept fleas Siphonaptera were present (83% �parental FM are a paraphyletic group consisting nests infested). In two-year-old nestboxes expected of 3 superfamilies Analgoidea, Pteroli­ that contain no old nest material both tns could choidea, Freyanoidea of the mite suborder fleas (100% nests), mites and ticks immune Astigmata. FM are variously adopted for Acarina were found. In addition to these �s. Addi­ inhabiting different types of microhabitats groups , in two-year-old nestboxes with ort seems on a bird body. They live on different old nest material also feather lice Mallo­ it should types of plumages and even specialised to phaga were found. studies of certain structure elements of feathers, on The intensity of infestation with fleas - just like a skin of birds , in nasal cavities and air was greater in these two-year-old nest­ �tudies. sacs. boxes that contain old nest material. A comparative morphological studies Conclusions: Age of the nestsite influ­ [TY IN of different groups of all recent subfami­ ence the diversity of the ectoparasites in RELA- lies show that representatives of different nests of Starlings. In 1996, where there 8: NEST- taxa of higher (familial) rank adapted to were no nestboxes with old nest material the same microhabitats display similar at the start of breeding season, only fleas main morphological adaptations even if were present in Starlings nest. In 1997, mder H. they are rather distantly related one to when some nestboxes containing old nest another. Five general morphotypes could material allowed for the development and be recognised. It is concluded, that devel­ ·ersity of wintering of ectoparasites, both diversity opment of certain adequate morphotypes ;titute of and intensity of ectoparasites were in­ was independent in several different Sciences, creased. The presence of the old nest phyletic lines. A phylogenetic hypothesis II_laterial influences intensity of ectopara­ elaborated by cladistic methods for the sttes. Starlings ' behaviour (high mobility, superfamily Analgoidea allowed to pro­ nposition nestsite usurpation, brood parasitism) pose a hypothesis of the evolution of ' Starling greatly facilitates fleas transmission rates general morphological adaptations of FM >account causing the prevalence to remain high o� during the pioneering and settling in the : nor the both seasons. different microhabitats. f attempt 74

A cladistic analysis of phylogenetic Capillaria carbonis (Rud.,) on one ined relationships of the FM families Aven­ cormorant in the Milicz area in Lower tions, zoariidae and Alloptidae based on mor­ Silesia. This nematode is hosted by: rural: phological characters and subsequent Podiceps ruficollis, Pelecanus crispus, Resu comparative analysis of obtained hy­ P. onocrotalus, P. carbo, Phalacroco­ affec1 potheses for FM and recent phylogenetic rax. pygmaeus, Ardea purpurea, Egretta hosts hypotheses of aquatic birds indicate a alba, Mergus squamatus, Stercorarius lympl basic pattern of coevolutionary relation­ pomarinus. The distribution: Europe plasn ships of feather mites with their hosts. (England, Yugoslavia, Bulgaria, Roma­ that The clear pattern of phylogenetic paral­ nia), Asia (Kazakhstan and republics of immu lelism, which could be expected, is dis­ Middle Asia, Far East, Afghanistan). Besid turbed by different evolutionary events as Outside the Palearctic Region in North medi< a host shift, incoherent evolutionary rate America, Africa, Australia. The biology SeX-Si in mite lineages within same host phyletic of C. micropapillatum was described by peripi branch, and probably by extinction of Semenova (1971) focusing on its devel­ in ma certain phyletic lines of FM. opment in different specie of Copepoda of res and fry. Infective larvae may also con­ as c< CONTRACAECUM MICROPAPIL­ centrate in paratenic hosts: dragon-flies, hemo LATUM (STOSSICH, 1890) (NEMA­ tadpoles and differentfish species. levels TODA) - NEW SPECIES TO THE infect PARASITOFAUNA IN POLAND HEALTH IMPACT OF BLOOD stage: PARASITES IN BREEDING GREAT cause 1 2 Anna Okulewicz and Jerzy Rokicki TITS an em 'Department of Parasitology, Microbiologi­ were cal Institute, Wroclaw University, Wroclaw, Indrek Ots1 and Peeter H5rak2 urban 2 Poland; Department of Invertebrate Zool­ 'Institute of Zoology and Hydrobiology, sugge 2 ogy, Gdansk University, Gdynia, Poland Tartu University, Tartu, Estonia; Estonian breed Institute of Zoology and Botany, Tartu, mass The paper deals with the results of para­ Estonia parasi sitological autopsy of cormorants col­ Cone. lected in the area of the Vistula Lagoon Objective: Hypotheses of hemoparasite­ sui tat and the Gulf of 'Gdansk in May 1995 and mediated sexual selection and reproduc­ testinl August 1997. Seven birds were infected tive costs rely on the properly untested select with Contracaecum micropapillatum; the assumption that avian blood parasite of rep nematode location on the host were infections are harmful to their hosts. We hypothesised that if blood parasites proventriculus and gizzard. The intensity THE impose any serious health impact on their was from 2 to 35 specimens (males, MOR avian hosts, then infected individuals females and larvae). Description and INTE measurement concerning the present must differ from uninfected ones in material of Phalacrocorax carbo corre­ respect to hemato-serological general Algim sponded to those published by Barus et health and immune parameters. Institu al. (1978). Information on the nematodes Material and methods: Health impact of Vytaut of P. carbo in Poland is poor. Up to now Haemoproteus blood parasites on their Lithua only Okulewicz (1989) has recorded great tit (Parus major) hosts was exam- 75

on one ined during three years in two popula­ The studies have revealed that different in Lower tions, breeding in contrasting (urban and population and species of both the para­ )Sted by: rural) habitats in south-east Estonia. sites and the hosts may have different crisp us, Results: Haemoproteus blood parasites genetic polymorphism and heterozygos­ alacroco­ affected the health state of their avian ity. Our population dynamic studies hosts. Infected individuals had elevated which consider genetic polymorphism of z, Egretta rcorarius lymphocyte hemoconcentration and helminths and birds have shown fluctua­ Europe plasma gamma-globulin levels, indicating tions of different genotypes in space and that both cell-mediated and humoral time. We studied the genetic variation of 1, Roma­ N. mblics of immunity are involved in host defence. trematodes attenuatus and N. ephem­ hanistan). Besides, the effect of parasites on cell­ era (multiple-host parasite withlow host in North mediated immunity was both age- and specificity) and their intermediate e biology sex-specific, as infection status affected (molluscs) and definitive (birds) hosts. cribed by peripheral blood lymphocyte counts only The investigation of genetic variation its devel­ in males, and among these, the magnitude within and between 7 populations of these ::opepoda of response was greater in old individuals trematodes showed that levels of genetic also con­ as compared to yearlings. Heterophile variation had different values and ranged .gon-flies, hemoconcentration, plasma albumin from P=0.33, H=0.059 to P=0.61, �s. levels and hematocrit were not affected by H=0. 177 for N. attenuatus and from infection status, suggesting that blood P=0.47, H=0.134 to P=0.59, H=0.23 for BLOOD stages of Haemoproteus infection do not N. ephemera. On the other hand the GREAT cause severe inflammatory response or genetic polymorphism among host species anemia. In two years, infected individuals and population also different: for inter­ were heavier than uninfected ones in the mediate host (for example molluscs) urban but not in the rural study area. This ranged from P=O.O, H=O.O (A rion sp.) to robiology, suggests, that under certain circumstances P=l .OO, H=0.63 (Cerithium caerulleum), 2Estonian breeding great tits may avoid losing body and for definitive host - birds: from y, Tartu, mass in order to save resources for anti­ P=0.06 (Larus marinus) to P=0.54, parasite immune response. H=0. 169 (Anas platyrhynchos). Genetic Conclusion: Our results support the variation of parasite between populations )parasite­ suitability of using hemoparasites for from different species of host and be­ ·eproduc­ testing hypotheses regarding sexual tween populations from the same host untested selection and physiological mechanisms species in different localities is different. parasite of reproductive cost. In the case of maximum specificity for a 1osts. We host species and population the parasite parasites THE ROLE OF GENETIC POLY­ genome is strictly bound to one host ton their MORPHISM IN BIRD-PARASITE genome and must imperatively find it dividuals INTERACTIONS from generation to generation. Evolution ones in of the host polymorphism is thus determi­ general Algimantas Paulauskas nant with regard to that of the parasite. Institute of Ecology, Vilnius; Lithuania; Parasite strains which are adapted to their mpact of Vytautas Magnus University, Kaunas, local host population are more virulent Lithuania on their than parasite strains introduced into host lS exam- 76 populations which previously had no Results: Cell divisions were mainly DEVE experience with this parasite strain. The observed in developing embryos. The BIRD existence of such large polymorphism in karyotypes of five studied species of TIOM parasite and host populations gives an genus Aploparaksis, A. fu rcigera, A. LYMN opportunity for the parasites to be brachyphallos, A. retroversa, A. occi­ ITING adapted to the environmental changes. dentalis and A. fi liformis, contained 6 ENVII The rate at which such high heterozygos­ pairs (2n= 12) of small chromosomes, SILES ity occurs in natural helminth population ranging in size from 1.5 to 4.0 Jlm. The (SOUl defines the genetic structure of the first pair of homologues was larger than helminth population and can influence the the remaining elements, which decrease in Zbignie Departt ability of the helminth to respond to size fairly gradually. The site of cen­ Silesian selection pressure. tromere localisation in chromosomes Katowi• could not be established exactly. Such CHROMOSOME STUDIES ON general peculiarities of chromosome set Object SOME CYCLOPHYLLIDEAN CES­ structure are also characteristic for liminar TODES, THE PARASITES OF Dicranotaenia fallax, Dicranotaenia sp., todes 1 BIRDS OF NORTHWESTERN Retinometra giranensis, Anatinella commc CHUKOTKA spinulosa, Wardium retracta. The inhabit karyotypes of Sobolevicanthus mastigo­ situate< Romualda Petkeviciiite1 and Kira V. RegeF praedita and S. spasskii consisted of 4 Region 1lnstitute of Ecology, Vilnius, Lithuania; pairs of subtelocentric, 1 pair of subtelo­ 2Institute of the Biological Problems of the submetacenrtic and 1 pair of small meta­ North, Magadan, Russia re centric homologues. Fimbriaria sp. XIF possessed symmetric chromosome set, *C Objective: Only scattered data exist on composed of 5 pairs (2n=l0) of meta­ (sy the karyology of cyclophyllidean cesto­ centric chromosomes. The karyotype of FUJ des, undoubtedly due to the small chro­ D *C Dichoanotaenia bacilligera (fam. mosomes and the difficulties in obtaining *C Dilepididae) consisted of 8 pairs of = good cytological preparations. A kary­ ( i biarmed chromosomes fairly gradually Scl ological approach can contribute to a decreasing in size from 4.0 to 2.1 flill. ECJ better knowledge of the group. We have, *C Conclusions: Cyclophyllidean cestodes therefore, studied the chromosome sets of (=1 are karyotypically conservative. There 14 species belonging to the families Eel was noted a tendency to maintain a Hymenolepididae and Dilepididae. FUJ constant chromosome number and mor­ *C Materials and Methods: The study was phology within genus. According to the d (=J carried out on strobilae of mature hel­ Scl existing data, diploid chromosome num­ minths, obtained at the dissection of natu­ MC bers in fam. Hymenolepididae ranges rally infected birds (Anatidae and Scolo­ *C pacidae). The birds were collected from from 2n=6 to 2n= 12. The most species (=N Chauna lowland (Northwestern Chukot­ have 12 chromosomes in their diploid sets ka). Chromosome materials were prepar­ and this can be considered the basic Explant number for the family. The most studied d-smal ed from cell suspension by using an air­ 1 dried method and stained with Giemsa. representatives of fam. Dilepididae have colleC! 16 chromosomes in their diploid sets. 77

:re mainly DEVELOPMENTAL STAGES OF the direct result of coal-mining activity. >ryos. The BIRD TREMATODES IN POPULA­ Usually they are shallow and without­ species of TIONS OF THE POND SNAIL flow ponds, with muddy bottom and cigera, A. LYMNAEA STAGNALIS (L.) INHAB­ strong mineralization of water, what is :, A. occi­ ITING ANTHROPOGENIC WATER related to their high trophy. Sand-pits ontained 6 ENVIRONMENTS IN THE UPPER originated from the inflow of atmospheric omosomes, SILESIAN INDUSTRIAL REGION or ground water due to sand-excavations. 0 J.lm. The (SOUTHERN POLAND) The degree of water pollution in sand-pits larger than in connection with their flow-through and decrease in Zbigniew Pokora usually considerably depth, is smaller in Department of Biology and Parasitology, te of cen­ comparison to sinkholes. Silesian Medical Academy, Materials and Methods: The material in romosomes Katowice, Poland .ctly. Such selected anthropogenic water reservoirs 1osome set was collected during the years 1996- Objective: In this paper results of pre­ �ristic for 1997, fromApril to October. liminary investigations on larval trema­ 'taenia sp., Results: In the examined reservoirs 254 todes parasitizing in populations of the Anatinella adult snails were collected (mean shell common pond snail (Lymnaea stagnalis), !Cta. The height 42 mm). Results of the parasi­ inhabiting typical sinkholes and sand-pits s mastigo­ tological examinations are presented in situated in the Upper Silesian Industrial ;isted of 4 the table below. Region are presented. Sinkholes were of subtelo­ mall meta­ c c ·iaria sp. XIPHIDIOCERCARIAE: 1· 2 >some set, * Cercaria pseudogracilis Zdun ) of meta­ (syn. Cercaria gracilis Wes.-Lund) FURCOCERCARIAE: ryotype of 1• 2 D * Cercaria longiremis Wes.-Lund cz (fam. * Cercaria ocellata La V al. pairs of (=Trichobilharzia ocellata (La V al.) Brumpt cere.; 1 gradually Schistosomatidae) X �.1 f.lill. ECHINOSTOMATA: * Cercaria echinata Siebold 1 cestodes (=Echinostoma revolutum (Frolich) Looss cere.; .2 ve. There Echinostomatidae) XI naintain a FURCOCERCARIAE: XIPHIDIOCERCARIAE: and mor­ * Cercaria ocellata La V al. * Cercaria gibba Sin. ling to the d (=Trichobilharzia ocellata (La V al.) Brumpt cere.; (syn. Cercaria tenuispina Luhe 2 Schistosomatidae) X =Opisthioglyphe ranae (Frolich) Looss cere.; :ome num­ 1· 2 MONOSTOMATA: Plagiorchiidae) ae ranges * Cercaria monostomi Linstow FURCOCERCARIAE: 2 )St species (=Notocotylus seineti Fuhr. cere .; Notocotylidae) X * Cercaria cristata La V al. 2 liploid sets ! (=Sanguinicola sp. cere.; San,gunicolidae) the basic Exp lanations: C-common 'species', c-rare 'species', D-numerous 'species', )St studied d-small 'species'; X-cercariae known as larvae of trematodesparasitizing in birds, 1 2 iidae have collected in examined sinkhole-ponds, collected in examined sand-pits. j sets. 78

Conclusions: In L. stagnalis collected in Results: The most common blood para­ phys c both types of examined anthropogenic sites in studied black grouse population Ligula water reservoirs, the dominant larvae of were Leucocytozoon lovati and microfi­ solidu bird trematodes were cercariae of E. laria. The probability of an individual to tracae revolutum, while cercariae of T. ocellata have a Leucocytozoon infection decreased 1890); were common and numerous only in with increasing age and was highest in strum( examined sinkhole-ponds. yearlings (50%). The incidence of micro­ Th� ftlaria was lowest among yearlings (12%) Dip[Oj ABILITY OF HOST TO RECOVER but increased with age. cephai FROM BLOOD PARASITE INFEC­ Conclusions: Repeated examination of paras it TIONS - A LONGITUDINAL STUDY haematozoan infection status revealed occasi1 IN THE BLACK GROUSE that black grouse are often able to re­ of the cover from Leucocytozoon infection. ring in 1 2 Osmo Ratti , Rauno V. Alatalo and Pekka Leucocyto­ and so 3 There was no evidence that T. Rintamaki zoon affected survival of black grouse. a basi• 1Arctic Centre, University of Lapland, Contrary, black grouse seldom recovered most z Rovaniemi, Finland; 2Department of Bio­ from microftlaria infection. Moreover, are the logical and Environmental Science, Univer­ carbo sity of Jyvaskyla, Jyvaskyla, Finland; microftlaria infection seemed also to 3Department of Zoology, Uppsala University, reduce life-span of male black grouse. Uppsala, Sweden These findings suggest that black grouse PARA immune system is able to successfully PTAR Objective:. Sampling each host individ­ eliminate or at least control Leucocyto­ IN ICJ ual only once gives us momentary infor­ zoon infection but not microftlaria infec­ mation on blood parasite infection status tion. Karl Sl lnstitut of individuals. However, the understand­ Keldur, PISCIVOROUS BIRDS IN THE ing of parasite-host interaction demands Iceland also knowledge on dynamics of infections HELMINTHS DEVELOPMENT CYCLE OF THE GULF OF within individual hosts. Very little is Object known about ability of free living hosts to GDANSK (THE BALTIC SEA) of exaJ recover from different blood parasite 1 the rcx infections. Jerzy Rokicki , Leszek Rolbiecki 1 and Anna Okulewicz 2 Icelanc Materials and Methods: We studied a 1Department of Invertebrate Zoology, Materi (Tetrao tetrix) black grouse population in Gdansk University, Gdynia, Poland; De­ In 199, 1988-1994. central Finland during We partment of Parasitology, Microbiological 88 ptaJ sampled blood from 284 black grouses Institute, Wroclaw University, Wroclaw, were e 391 times (up to 5 times/individual) for Poland Also, examination of haematozoan parasites. Blood smears were air-dried and fixed were Birds are an important link in the devel­ with 100 % methanol and sent to the ethylac International Reference Centre for Avi an opment of some parasites occurring in the presem Haematozoa (Memorial University of Gulf of Gdansk. There are seven species eggs. ] Newfoundland) where they were stained of them : Digenea: Cryptocotyle conca­ nation with Giemsa stain and examined by G. F. vum (Creplin, 1825); Diplostomum didofilt Bennett. spathaceum (Rudolhp, 1813); Tylodel- 79 lood para­ phys clavata (Nordmann, 932); Cestcxla: on 12 ptarmigans by examining the population Ligula intestinalis (L.); Schistocephalus oesophagus, trachea, crop and the con­ d microfi­ solidus (MUller, 1776); Nematcxla: Con­ nective tissue and fat deposits surround­ lividual to tracaecum micropapllatum (Stossick, ing these organs. Ectoparasites: Eight decreased 1890); Acanthocephala: Corynosoma birds were examined in 1998 for feather highest in strumosum (Rudolphi, 1802). lice. In September 1997 louse flies were of micro­ The most often occurring species are sampled, or their occurrence noted, on ngs (12%) Diplostomum spathaceum and Schisto­ 175 ptarmigans which were life-captured cephalus solidus. Most of the noted and ringed in Hrisey, N-Iceland. Hae­ ination of parasites were found on fish and only matozoans: Blood smears were collected revealed occasionally on birds or molluscs. Most and stained from the 175 life-captured ble to re- of the 30 piscivorous bird species occur­ ptarmigans in Hrisey as well as from infection. ring in the Gulf of Gdansk are migrating further 94 birds ringed in NE-Iceland in �ucocyto­ and sometimes wintering ones. Fish make August 1997. However, as only few of :k grouse. a basic food only for some of them. The the blood smears have been examined, the recovered most abundant, typical piscivorous birds results on the occurrence of haematozo­ Moreover, are the following species: Phalacrocorax ans have to be regarded as preliminary. also to carbo and Podiceps cristatus. Results: Endoparasites: Two different ok grouse. sized Eimeria oocysts were found in the .ck grouse PARASITES OF THE ROCK faecal samples. A relatively large species tccessfully PTARMIGAN (LAGOPUS MUTUS) (oocysts measuring 25.2 x 17.7f.lm, range eucocyto­ IN ICELAND 21.3-31.0 x 12.4-21.7f.lm, n=114) was lria infec- found in 46 birds (prevalence 52.9%). A Karl Skimisson relatively small species (oocysts measur­ Institute for Experimental Pathology, ing 15.0 x 13.1f.lm, range 12.5-17. 1 x Keldur, University of Iceland, Reykjavik, N THE 12.4-14.0f.lm, n=9) was found in one bird Iceland >PMENT (prevalence 1.1%). Two intestinal nema­ tcxles have been found: Capillaria LF OF Objective: To present preliminary results caudinflata (prevalence 34. 1 %, mean A) of examinations on the parasite fauna of intensity 37, range 1-340) was found in the rock ptarmigan (Lagopus mutus) in cki 1 and the small intestine and caeca and Iceland. Trichostrongylus tenuis (prevalence Materials and Methods: Endoparasites: Zoology, 11.8%, mean intensity 3.4, range 1-9) land; De­ In 1994 and 1995 the small intestines of was exclusively found in the caeca. )biological 88 ptarmigans and the caeca of 93 birds Tissue parasite: S. papillocerca was not Wroclaw, were examined for helminth parasites. found. Ectoparasites: Two mallophagans Also, 87 faecal samples from rectum were found; Goniodes lagopi (prevalence were examined by the formalin­ 75%) and Lagopoecus affinis (prevalence the devel­ ethylacetate concentration methcxl for the 63%). During the ringing activities the ·ing in the presence of protozoans and helminth louse fly Omithomya chloropus was �n species eggs. Tissue parasite: In 1998 an exami­ noted on approximately every fourth le conca­ nation for the ftlarial nematcxle Splen­ ptarmigan and could be sampled from lostomum didofilaria papillocerca was carried out Tylodel- 14,3% of the individuals. Haematozoans: 80

So far, preliminary results on the exami­ using qualitati ve methods of blood drop tosoma nation of the blood smears have not agglutination with Kirzajev antigen and parasiti revealed any parasites. double diffusion into agar gel as well as m amm< Conclusions: Parasites of the rock ptar­ quantitative immunoenzymatic method compri: migan in Iceland have not been systemati­ ELISA . which I cally surveyed before. Although relatively Results and Conclusions: Estimation of study s few birds have been examined already antibody titer has shown that revaccina­ monopl seven species have been found. T. tenuis tion of Hisex brown breed hens with schisto! and the Eimeria spp. are reported for the "TALOV AC 109 SE" has not only in­ ated ill< firsttime from Icelandic ptarmigans. creased antibody titer and stability the pm against Salmonella enteritidis( after branch SPECIFIC FEATURES OF ANTI­ vaccination and revaccination average this fan SALMONELLA IMMUNITY DE­ antibody titer in the first experimental Materi: VELOPED BY "TALOV AC 109 SE" group was S/N=0.52, S/N=0.122 and in preservi AND "SG VACCINE NOBILIS" the second group was S/N=0.64, around . VACCINES S/N=0.166, respectively) but also devel­ from th oped immunity against other pathogenic mitochc I Sigita Slavenaite', Aniolas Sruoga , Dalius bacteria of Salmonella genus (S. ty­ the pd I 2 Butkauskas and Jonas Skonsmanas phimurium, S. pullorum, S. gallinarum). sequenc I sequenc Institute of Ecology, Vilnius, Lithuania, It has also been determined that "TALO­ 2 VAC 109 SE" vaccine developed higher aligned Stock Company "Vievio paukstynas", using P, Vievis, Lithuania level of immunity than "SG VACCINE NOBILIS", however both vaccines ensure Results transmi1 Objective: Vaccination represents one of immunity against S. enteritidis and S. dritobil. current measures to combat with salmo­ gallinarum. Higher stability of the im­ bilharzi nellosis, therefore vaccination of Hisex mune system response to vaccination has Howeve brown and Hisex white breed hens using been stated in Hisex white breed hens in bilharzi "TALOV AC 109 SE" and "SG VAC­ comparison with Hisex brown breed hens. mitted t CINE NOBILIS" vaccines has been evolutio carried out in the Vievis Poultry Yard in EVOLUTIONARY RELATION­ malian 1996. The objective of our work was to SHIPS AMONG AVIAN AND MAM­ the othe determine effectiveness of the above MALIAN BLOOD FLUKES (DIGE­ Conclw vaccines in the development of immunity NEA: SCHISTOSOMATIDAE) systema againstSalmonella enteritidis. Scott D. Snyder matid t Materials and Methods: Hisex brown Department of Biology, University of New host affi and Hisex white breed hens vaccinated Mexico, Albuquerque, New Mexico, USA spond \\ with inactivated "TALOV AC 109 SE" a of the 1 product of German company "TAD" and Objective: This study sought to use switchin live "SG VACCINE NOBILIS" by Neth­ ribosomal and mitochondrial DNA se­ matidae erlands company "INTERVET" have quence information to construct an hy­ definitiv been used for an experiment. An antibody pothesis of phylogenetic relationships mammal titer has been determined in the blood of among the genera comprising the Schis- from the vaccinated and revaccinated hens 81 blood drop tosomatidae. The Schistosomatidae (Supported by NSF/Sloan Grant BIR- .ntigen and parasitize the vascular systems of birds, 9626072) as well as mammals, and croccxlilians. The family is le method comprised of 13 distinct genera, seven of ALLERGENIC ACAROFAUNA which parasitize birds. The results of this FROM NESTS OF SELECTED timation of study should demonstrate if the family is SPECIES OF SYNAN-THROPIC revaccina­ monophyletic, if avian and mammalian BIRDS IN POLAND hens with schistosomatid lineages arose and radi­ Krzysztof Solarz, Piotr Szilman and Ewa )t only in- ated independently of one another, and if Szilman 1 stability the parasitism of pulmonate or proso­ branch snails is the ancestral condition in �p�ent of Biology and Parasitology of dis( after Stlestan Medtcal. Academy, Katowice, Poland this farnily. ill average Materials and Methods: Ethanol­ lperimental Obj ective: The possible sources of preserved parasites were obtained from 122 and in allergenic mites i.n dwelli.ngs and stored around the world. DNA was extracted SIN=0.64, products are nests of synanthropic birds, from these parasites, and ribosomal and also devel­ especially sparrows and swallows. The aim mitochondrial genes were amplified using pathogenic of this study was to i.nvestigate the possible the polymerase chain reaction. DNA lS (S. ty- occurrence of these mites (pyroglyphid dust sequences were obtained using automated 1llinarum). mites, acarid and glycyphagid storage mites sequencing technology. Sequences were at "TALO­ or others) i.n nests of domestic sparrows aligned using ClustalW and analysed ped higher (Passer domesticus) [P D] and swallows using PAUP*. VACCINE (Delichon urbica [D U] and Hirundo Results: The four avian, pulmonate snail­ ines ensure rustica [H R]) fromsouth part of Poland transmitted genera (Bilharziella, Den­ tis and S. Materials and Methods: The study was dritobilharzia, Gigantobilharzia, Tricho­ of the im­ carried out from January 1997 to March bilharzia) form a monophyletic group. ination has 1998. A total of 25 nests were examined However, Austrobilharzia and Omitho­ :eel hens in i.ncludi.ng 11 nests of P D, 7 nests of D U bilharzia, avian schistosomatids trans­ xeed hens. and 7 nests of H R. Majority of these nests mitted by prosobranch snails, share closer was collected i.n Solarnia near Lubliniec, in evolutionary relationships to some mam­ LATION­ Tychy and vicinity and i.n Laziska Gorne malian schistosomatid parasites than to ID MAM­ (all localities from Upper Silesia i.n Poland). the other avian schistosomatids. S (DIGE­ The mites were extracted using the Berlese Conclusions: These data indicate that �E) method and preserved i.n 70% ethanol. For systematic schemes grouping schistoso­ identification the mites were mounted in matid parasites according to definitive Hoyer's mediumon microscope slides. ity of New host affinity need to be modified to corre­ Results: Twenty four of 25 nests contained eo, USA spond with the evolutionary relationships mites (96% ). A total of 10 007 mite of the genera. It is apparent that host specimens were isolated i.ncluding 9 852 ht to use switching has occurred in the Schistoso­ mites from the order Astigmata (98%). DNA se­ matidae at both the intermediate and Among the astigmatid mites (13 species) the let an hy­ definitive host levels and that some most abundant were Hirstia passericola [H lationships mammalian parasite genera have arisen P] (82.35% of the total count), the Schis- from ancestors parasitic in birds. 82

Glycyphagus domesticus [G D] (9.61%), chia) were collected in the lake Asveja in SOMJ Dermatophagoides evansi (3.17%) and summer of 1994-1997. The study of DISTl Tyrophagus similis (2.56% ). So, H P have chromosomes was carried out on somatic MITE been found as numerically dominant, cells of parthenites using air-drying NON, especially in the nests of D U (99. 25%) and techniques. H R (93.17%). G D, the allergenic mite Results: The diploid number of chromo­ Maria l 1Zooloj species, was found only in 2 nests of P D. G somes in metaphase plates of both inves­ 20kskii D was the dominant in a total count of mites tigated species was found to have a mode from the nests of P D (48.3%). Other of 20. Mitotic complement of platy­ I. The rh astigmatid species were distinctly less cephalus consisted of metacentric (pairs ous ar numerous and formed (altogether) only 1, 9, 10), submetacentric (pairs 6-8) and of g1 0.76% of all mites. subtelocentric (pairs 2 - 5) chromosomes. Mesos Conclusions: Nests of both species of The chromosome complement of varie­ I. cavitie swallows are most frequently dominated by gatus was found consist of subtelocentric More H P. The potential sources of some aller­ (pairs 1 - 5), submetacentric (pairs 6 - 8) are knc genic mites (as G D or Tyrophagus spp.) and metacentric (pairs 9, 10) chromo­ bution are the nests of domestic sparrows (P D). somes. The mitotic cells of variegatus I. differe: collected from 6 molluscs contained an factors additional 21st chromosome. The mor­ NEW DATA ON THE KARYO­ season TYPES OF GENUS ICHTHYO­ phology of this chromosome was different num be COTYLURUS in metaphase plates from various indi­ ODENING, 1969 possibi viduals. The small subtelocentric to (TREMATODA, STRIGEIDAE) differeJ submetacentric additional chromosome with tJ Grazina StaneviCiute and Valerij a Kiseliene was found in chromosome sets of I. victim Instituteof Ecology, Vilnius, Lithuania variegatus from four snails. The larger species submetacentric additional chromosome, rhinon: Objective: In the course of karyological which most probably is B-chromosome, birds i: research of trematodes, the chromosome was found in metaphase plates of I. Ichthyocotylurus platycephalus gation. sets of variegatus from two snails. (Creplin, 1825) Odening 1969 and analys( I. Conclusions: Karyotypes of I. platy­ variegatus (Creplin, 1825) Odening species cephalus and I. variegatus display a 1969, a parasite of piscivorous birds, sentati' typical strigeid pattern with 20 chromo­ belonging to the family Laridae, Gavii­ materi< somes. Comparative karyological analy­ dae, Podicipedidae, Phalacrocoracidae, anatid l sis showed that principal morphological of hunt was investigated. The aim of this study changes in karyotypes during the specia­ was also to determine the usefulness of Okskii tion in this group was caused by a Reserv1 karyological data as new characteristic smaller or larger pericentric inversions. together with traditional taxonomic informl methods in a comparative study of tre­ rhinon) matodes. (Troue: Materials and Methods: Naturally materia infected first intermediate hosts - the flocks 1 molluscs Va lvata piscinalis (Prosobran- juvenile 83

: Asveja in SOME PECULIARITIES OF THE ones. There are specimens in a popula­ study of DISTRIBUTION OF RHINONYSSID tion, which serve as a reservoir of para­ Jn somatic MITES (ACARI, GAMASINA, RHI­ sites (to 90 mites on the bird). The data air-drying NONYSSIDAE) OF BIRDS on the interrelations connected with sexual, age, reproductive activity, list of 1 2 Maria K. Stanyukovich andO.M. Butenko species, specificity to bird-hosts in sev­ )f chromo­ 1 Zoological Institute, St Petersburg, Russia; eral regions of Russia (for example, in Joth inves­ 20kskii State Biosphere Reserve, Russia tve a mode Ryazan and in Novosibirsk Oblasts) are I. platy­ given. f The rhinonyssid mites is the most numer­ ttric (pairs ous and complicated in collecting group FRENKELIA , and BUZZARDS AND 6-8) of gamasids (Acari, Parasitiformes, Jmosomes. RODENTS: MULTIPLE MECHA­ Mesostigmata) parasiting in the nasal Jf I. varie­ NISMS OF PARASITE SURVIVAL cavities, tracheae and lungs of birds. ltelocentric STRATEGY More than 400 species of rhinonyssids 1airs 6 - 8) are known in the world fauna. The distri­ Milena Svobodova1, Jan Votypka1 and Petr ) chromo­ bution of these mites on the hosts is rather Votisek2 variegatus different. It depends from the many 1Department of Parasitology, Charles Uni­ ntained an factors: species of bird, age, sex, locality, versity, 2Czech Society for Ornithology, The mor- season, etc . The birds can have different Prague, Czech Republic ts different number of rhinonyssids species. The rious indi­ possibility of exchange between birds of Frenkelia spp. (Apicomplexa: Sarcocys­ centric to different species with rhinonyssids is rare tidae) are dixenous coccidian parasites of rromosome with the exception of contacts between buzzard (Buteo buteo) and its rodent I. sets of victim and prey or birds of different prey. Sporocysts are excreted in buz­ The larger species in the places of feeding. The zards' faeces (final host), while tissue romosome, rhinonyssids from nasal cavities of anatid cysts develop in the brain of rodents romosome, birds is the convenient group for investi­ (intermediate hosts). The prevalence of F. ates of I. gation. It's possible to compare and glareoli and F. microti was studied in analyse rhinonyssids from different Czech Rep. Buzzard faeces were col­ I. platy­ species of ducks, geese and other repre­ lected on nests with young, rodents (field a display sentatives of the family. The analysis of vole Microtus arvalis, wood mice Apo­ 0 chromo- material collected from nasal cavities of demus sylvaticus and A. jlavicollis, bank 1cal analy­ anatid birds in spring and autumn seasons vole Clethrionomys glareolus) were ·phological of hunting (1960-1990) on the territory of snap-trapped. The total prevalence of the specia­ Okskii State Biosphere sporocysts in buzzards was 72%. The sed by a Reserve (Russia) gave the concrete prevalence increases with age of nes­ ersions. information concerning the distribution of tlings, 75% of samples being positive rhinonyssids from genus Rhinonyssus before fledging. The prevalence of F. (Troues.). Some results based on the glareoli in bank vole is 38%, of F. mi­ material collected could be given. In croti in field vole 5%. The main buzzard flocks the quantity of parasiting mites on prey, however, is the field vole. The small juvenile ducks is more than on the old number of bank vole in buzzard diet is therefore compensated by 10 fold higher 84 anotl prevalence of the parasite in the interme­ The study of breeding urban and wild­ than diate host. The prevalence also differs nesting Mallard Anas platyrhynchos between habitats of rcxients trapping, populations has been carried out in INVJ being the highest in ecotones. Frenkelia Minsk, Belarus since 1989. In 1996, the TIOI sporocysts probably cumulate there, investigations of mallard Haemosporida because buzzard use ecotones as perching and vectors of the parasites began. The FEC sites. This makes the bank vole more aim of this paper is to present the differ­ Danie vulnerable to infection. ence of vector's fauna and phenology at Depru Diet composition significantly affects urban and wild nature territories and its Scien, prevalence in buzzards. Frenkelia sp. relation with phenology of mallard. U.K. sporocysts were found in faecal samples Samples of Diptera were taken by from 57% of nests in 1993, and 90% in means of an entomological sweep-net and Obje 1994. The proportion of field vole in the light traps. The preimaginal stages of paras diet was 24% and 52%, resp. Higher Culicidae and Simuliidae were taken at extret prevalences in rcxients are also recorded their habitats located near mallard's of e: in spring than in autumn, when the pro­ nesting places. infect portion of adult individuals in rcxient Collections of blocxi-sucking Diptera high 1 population is the highest. from urban areas are presented by 19 infect The prevalence of F. microti in field species of Culicidae, 21 - of Ceratopo­ maxir voles collected on nests and snap-trapped gonidae and 6 - of Simuliidae. At the are of differed significantly (8% and 2% re­ territory of Belarus, 39 species of Culici­ same spectively.), suggesting that infection dae, 28 - of Ceratopogonidae and 38 -of host i increases the risk of predation by the final Simuliidae are known. Among them the cause: host. Experiment with related coccidian S. mosquitoes Culex pipiens pipiens L., Mate1 dispersa confirmed that infected mice are Culiseta morsitans Theob. and the black­ ne mat more susceptible to predation by the final flies Simulium truncatum Lundstr., S. remov host, the long eared owl (Asia otus). austeni Edv. have been recorded as 171 I Multiple mechanisms of parasite survival vectors of bird Haemosporida. At urbi­ colchi strategy therefore include 1. high envi­ coenoses, only C. p. pipiens was col­ contai ronmental resistance of infective stages; lected. Phenological indices of mallard as an 2. locally increased prevalence in inter­ life cycle are different at urban and egg v mediate host; 3. manipulation of the host nature areas. The urban nestlings hatch embry behaviour increasing the risk of preda­ out three weeks later than the nestlings of stage . tion. wild-nesting mallards. The hatching and c at 21 nestling pericxi in wild nature is at the basis) CONFORMITY OF PHENOLOGIES time of the most abundance of all groups worm OF MALLARD ANAS PLA TYRHYN­ of vectors. In urbicoenoses, the nestlings Resul1 CHOS L. AND INSECT VECTORS hatch out at the end of June, when abun­ follOW! dance of Culicidae and Ceratopogonidae was n Nadezhda Tereshkina and Alexandr Kozulin reduces about 1.5 times , Simuliidae - 20 negati' Institute of Zoology, National Academy of times. At this time, the structure of Sciences, Minsk, Belarus mean < vector's faunistic complex is changed, 0.32 (I 85

another species became more abundant correlation between the log-transformed md wild­ than in May and in the beginning of June. mean number of viable eggs per female •rhynchos worm and mean female worm length was l out in INVESTIGATING THE REGULA­ observed (p = 0.02). Furthermore, at 1996, the TION OF PARASITIC NEMATODE counts below 80 worms, female worm zosporida FECUNDITY length increased with increasing worm �gan. The count (p < 0.001) whilst, at counts above he differ­ Daniel M. Tompkins 80, female worm length decreased with nology at Department of Biological and Molecular increasing worm count (p = 0.03). �s and its Sciences, University of Stirling, Stirling, Conclusions: Influences on H. gallina­ rd. U.K. rum fecundity appear to occur via effects taken by on worm size. The non-linear relationship :p-net and Objective: A primary characteristic of between worm count and worm length stages of parasitic nematode life-histories is their implies that variations in the host immune taken at extreme fecundity. Since the probability system may be the primary determinant mallard's of each egg/larva produced actually of worm fecundity at low counts, whilst infecting a new host is often very low, the density-dependent competition among g Diptera high fecundity is necessary to ensure that worms may be the primary determinant of xi by 19 infections persist and parasite fitness is worm fecundity at high counts. Ceratopo­ maximised. However, large differences �. At the are often observed in the fecundity of the RATE OF ANTIBODY TITER AF­ of Culici­ same nematode species infecting different TER CHALLENGE BY VIRULENT ld 38 -of host individuals. Here I investigate the SALMONELLA ENTERITIDIS CUL­ them the causes of such variation. TURE IN VACCINATED HENS rJiens L., Materials and Methods: The parasitic the black­ nematode Heterakis gallinarum was I Birute VankeviCiene and Dalius But- ndstr., S. removed from one half of the viscera of 2 orded as 171 ring-necked pheasants (Phasianus kauskas I At urbi­ colchicus). The number of viable eggs Vievis poultry factory, Vievis, Lithuania; 2 was col­ contained within female worms was used Institute of Ecology, Vilnius, Lithuania f mallard as an index of nematode fecundity. An rban and egg was assumed to be viable if the Objective: Trial is devoted to evaluate ngs hatch embryo developed through to the infective influence of virulent Salmonella enteriti­ �stlings of stage after being maintained for 21 days dis culture on titers of specific antibodies �hing and ° in Hisex white (W) and Hisex brown (B) at 21 C. Worm fecundity (on a per host is at the breed hens vaccinated with inactivated basis) was analysed with respect to both groups vaccine "TALOVAC 109 SE". ill worm count and worm length. : nestlings Materials and Methods: Trial was Results: H. gallinarum worm counts hen abun­ carried out on laying hens of W and B followed an aggregated distribution that pogonidae breeds (from 280 to 510 days of life at was not significantly different from a iidae - 20 challenge moment). The birds were negative binomial with an arithmetic Licture of vaccinated twice with "TALOV AC 109 mean of 25.64 and a positive exponent of changed, SE" and challenged orally by virulent S. 0.32 (p = 0.25). Upon analysis, a positive enteritidis broth culture (0.2 ml!bird) on 86

14th day after second vaccination. Sam­ PREVALENCE OF COCCIDIA OF Caryo ples of blood were obtained on 35 day GENUS CARYOSPORA IN CAPTIVE sus w post challenge. Antibody titers were BIRDS OF PREY IN THE CZECH 25.0 > determined by using quantitative immu­ REPUBLIC wall. ] nocapture assay ELISA. micro1 1 L2 Results and Conclusions: Figures of Jiri Volf and David ModfY cal tc 1 antibody titers (IDEXX ELISA S/N) in Dept. of Parasitology, University of Veteri­ (15.07 nary and Pharmaceutical Sciences Bmo, blood samples of treated hens showed 2 substi1 Bmo, Czech Republic; lnstitute of Parasi­ high similarity (from 0.32 to 0.27 S/N). cysts tology, Academy of Sciences of the Czech Only in one case SIN quantity found Republic, Ceske Budejovice, Czech Repub­ chrysc equal to 0.57. It corresponds to the lowest lic (40.0- titer of specific antibodies. Laying of eggs thick hasn't been disturbed on post challenge Objective: The purpose of this study was uum time (hens were observed two months ). to describe the spectrum of coccidia of sporO< According to our earlier trial single, genus Caryospora in birds of prey in the X 23.; samples with low titer of antibodies were Czech Republic. Stieda found with 2.5% frequency on the con­ Materials and Methods: Since Septem­ absent trary to high level in majority cases ber 1996 to March 1998 the faecal sam­ Concl ascertained among twice vaccinated hens. ples of 397 birds of prey were examined statut� Due to experiences in Bavaria, where for parasites using faecal flotation the si: nearly 600.000 broiler breeders were method. The examined birds came from identi< vaccinated in last two years, vaccination raptor rehabilitation centers, Zoos and occurs program reduced S . enteritidis isolation private raptor keepers. Revealed coccidia status rate from 40% - before start of the vacci­ were allowed to sporulate in 2.5% (w/v) nation schedule - to 2.5%. Complete potassium dichromate solution and oo­ BLOC eradication of S. enteritidis can not be cysts were measured and determined MAT1 achieved by vaccination and probiotics using light and NIC microscopy. RAP1 treatment only. One of the reasons leading Results: The coprological examinations THE to failure of current measures to combat revealed four species of genus with salmonellas may be constant pres­ Caryospora in hosts of order Falconifor­ Jan \ ence of some percentage of immunosup­ mes. In the faecal samples of 4 of 47 Vonse pressed birds in breeders flocks. Moreo­ (8.51 %) examined Falco cherrug, 3 of Petr y, 1Depar ver, hens with such weak immunity 26 (11 .54%) F. peregrinus and 11 of 47 versity system create the main source of S. (23.40%) F. tinnunculus oocysts of 3Depar enteritidis shedding. Caryospora kutzeri were found. Oocysts Rep ubi of Caryospora neofalconis were found in 1 of 47 (2.13%) of F. cherrug, 4 of 26 Rapto: (15.38%) of F. peregrinus and 3 of 47 of blo (6.38%) of F. tinnunculus. 1 of 10 ex­ cocytc amined Aquila chrysaetos and 1 of 13 soma, examined Circus aeruginosus were found toes (1 to be passing oocysts of not yet described ozoon. species. The sporulated oocysts of protez� 87

IDIA OF Caryospora sp. 1 from Circus aerugino­ (blackfly, mosquito or hippoboscid fly) is ambiguous. :APTIVE sus widely oval 24.5 x 21.8 �m (23.0- In 1996 and 1997, we investigated the , CZECH 25.0 x 21.0-24.0 �m) with 2.0 �m thick wall. Polar granule, oocyst residuum and occurrence of blood parasites of raptors. Blood was collected from nestlings and micropyle absent. The sporocysts spheri­ adults of common buzzard (Buteo buteo, cal to subspherical, 16.2 x 15.6 �m n= 102), kestrel (Falco tinnunculus, r of Veteri­ (15.0-17.0 x 15.0-17.0 �m). Stieda and :lces Bmo, n=68) and sparrowhawk (Accipiter nisus, substieda bodies were absent. The oo­ n=324). Blood films were Giemsa­ · of Parasi­ cysts of Caryospora sp. 2 from Aquila . the Czech stained, cultivation was done on blood chrysaetos x ech Repub- subspherical 43.0 37.5 �m agar. (40.0-49.0 x 34.0-39.0 �) with 2.2 �m High prevalences of Leucocytozoon, thick wall. Polar granule, oocyst resid­ Trypanosoma and Haemoproteus were study was uum and micropyle were absent. The found in adult buzzards and sparrow­ :occidia of sporocysts spherical to subspherical, 23.8 hawks. All these parasites were found in young buzzards and sparrowhawks, too, prey in the x 23.3 �m (23.0-25.0 x 22.0-25.0 �m). Stieda and substieda bodies also were but the prevalence was markedly lower. Trypanosomes occurred in young and :e Septem­ absent. adult sparrowhawks and buzzards. No aecal sam­ Conclusions: While the new species parasites were found in the blood of � examined statute of Caryospora sp. 1 is apparent, nestling kestrels. Plasmodium was not the size of Caryospora sp. 2 is almost flotation found at all. C. megafalconis, which �ame from identical to In 1996, 1997 and 1998 haemato­ Zoos and occurs in hosts of family Falconidae. Its phagous insects feeding on nesting birds xi coccidia status should be an object of future study. were collected using the sticky bands, �.5% (w/v) which were exposed twice for 6 - 8 days n and oo­ BLOOD PARASITES AND HAE­ and blood-sucking insects were deter­ determined MATOPHAGOUS INSECTS OF mined. In total, sticky bands were in­ '(. RAPTORS (FALCONIFORMES) IN stalled on 18 buzzard, 17 sparrowhawk aminations THE CZECH REPUBLIC and 12 kestrel nests. )f genus In total, we captured 4 species of 1 Falconifor­ Jan Vot.YPka , Milena Svobodova\ Petr blackfly (the most common being Eusim­ Voi'isek2, Lubos Peske, David Lacina3 and ,f 4 of 47 ulium securiforme), 12 species of biting PetrVolf 1 midges (their respective representation rrug, 3 of 1Department of Parasitology, Charles Uni­ depending on the observed locality and d 11 of 47 2 versity; Czech Society for Ornithology; host species), 2 species of mosquito, and IOCysts of 3 Department of Zoology, Prague, Czech frequently the hippoboscid fly Omitho­ d. Oocysts Republic myia avicularia. re found in Our results show, that an important 4 of 26 g, Raptors are known as hosts of 4 genera part of populations of three raptor species td 3 of 47 of blood protozoans: Plasmodium, Leu­ in three different localities are infected of 10 ex­ cocytozoon, Haemoproteus and Trypano­ with blood parasites. Vectors of different d 1 of 13 soma, which are transmitted by mosqui­ blood parasites in respective hosts are were found toes (Plasmodium), blackflies (Leucocyt­ being sought for. t described ozoon) and biting midges (Haemo­ ocysts of proteus). The vector of trypanosomes Bull Scand Sac Parasito/ 1998; 8 (2): 88-91 68 I. hum a (HGf stricb FIRST DETERMINATION OF EHRLICHIA INFECTED TICKS w AMONG THE PRIMARY VECTORS OF THE TICK-BORNE ricint ENCEPHALITIS AND BORRELIOSIS IN THE RUSSIAN in the BALTIC REGION lining HGE 1997

A. Alekseev1, H. Dubinina I, L. Schouls2 flagg1 DF tt 1Zoological Institute, Russian Academy of Sciences, Universitetskaja emb., 1, by a St. Petersburg 199034, Russia, antigc 2Research Laboratory for Infectious Diseases, N ationa1 Institute of Public Health and fixed Environmental Protection, PO Box 1, Antonie van Leeuwenhoeklaan 9, gated 3720 BA, Bilthoven, The Netherlands retia 1996) Abstract Tl: This is the first communication of fr om the region surrounding St. Peters­ know Ehrlichia spp. detected in Ix odes persul­ burg or Kaliningrad. However, recently (Zlob catus Schulze and Ix odes ricinus (L.) dogs fr om St. Petersburg region were 1989) ticks collected in the Russian territory. fo und to show clinical signs similar to vecto; We tested a small number of ticks fo r the the symptoms described fo r severe Sprin; presence of Ehrlichia and Borrelia under ehrlichiosis (0. Arestov, personal com­ activi a darkfield microscope (DF). One of 10 munication). nymp I persulcatus was Ehrlichia spp. and In the USA (Ahkee & Ramirez, 1996; invest Borrelia garinii positive. One I ricinus Goodman et al., 1996; Barlough et al., Be out of 12 investigated by DF contained 1997) clinically and serologically con­ In bo1 not only spirochetes, but also Ehrlichia firmed cases of human ehrlichiosis have of the spp. been described fr equently, and in Europe Borre some of cases have been suspected al. , 1 � The north beach of the Finnish Gulf according to the serological data tions (Morskaj a, Lisy Nos) is an area of recre­ (Petrovec et al. , 1997; Dumler et al. , but SL ation for the inhabitants of St. Peters­ 1997). often burg. In this area I persulcatus, the pri­ Investigation of patients have provi­ three . mary vector of tick-borne encephalitis ded evidence fo r mixed infections with fe cted (TBE) and borreliosis, is abundant, and tick-borne pathogens such as Ehrlichia sprroc subsequently there is fo cus of both dise­ spp., Borrelia burgdorfe ri sensu lato in antige ases (Antykova et al. , 1998). addition to Babesia microti (Mitchell et contai Until now, no information about the al. , 1996). Infection of Ix odes ticks with ajzeli1 presence of rickettsia-like diseases such multiple pathogens seems to be typical, recent as granulocytic ehrlichiosis is available e.g. Pancholi et al. (1995) found 2 out of ricinu 89

68 I. scapularis ticks infected with the Stedingk et al. , 1997), and the presence human granulocytic ehrlichiosis agent of Ehrlichia species in I. persulcatus is (HGE) and Borrelia burgdorfe ri sensu not known in Europe. £CKS stricto. We tested a small number of ticks fo r We investigated I. persulcatus and I. the presence of Ehrlichia among DF fflE ricinus (L.) tick populations fr om areas Borrelia positive I. persulcatus and I. ,N in the vicinity of St. Petersburg and Ka­ ricinus ticks collected in areas of the liningrad fo r mixed infections with TBE, Russian Baltic region (see above). One HGE and B. burgdorfe ri s.l. During the of 10 I. persulcatus was positive in a 1997 season 579 ticks were collected by PCR based, reverse line blot hybridisati­ flagging and then investigated using the on assay (Schouls et al. , 1998, in press). DF technique. All ticks were examined This specimen was also infected with B. 1, by a direct IFA method to detect TBE garinii. One I. ricinus tick out of 12 in­ antigens. DF positive specimens were vestigated contained not only spiroche­ hand fixed in 70% ethanol and then investi­ tes detected by the DF method, but gated using a PCR method to detect Bor­ Ehrlichia spp. as well. relia genospecies (Rijpkema et al. , Although we fo und only two ticks 1996). positive fo r Ehrlichia, these data suggest The Kaliningrad region was also that infections with multiple pathogens .t. Peters­ known as a fo cus of TBE and borreliosis are not only present in the American I. ' recently (Zlobin et al. , 1996; Rej tchuk et al. , scapularis, but do also occur in Eastern ion were 1989) with I. ricinus as the primary European, i.e. - I. ricinus, and in Euro­ ;imilar to vector for these pathogens. Within the Asian I. persulcatus. Ticks fr om the r severe Spring and Summer peaks of I. ricinus Baltic regions may harbour at least three nal corn- activity more than a total of 900 larvae, different pathogenic genera: the TBE nymphs and adults were collected and virus, three Borrelia species and Ehr­ ·ez, 1996; investigated using the same methods. lichia species. gh et al., Both tick populations are allopatric. The determination of this new patho­ ally con­ In both regions mentioned, the presence gen circulating in Russian territories is osis have of the TBE virus and of two species of of great importance fr om epidemiologi­ n Europe Borrelia was demonstrated (Alekseev et cal and clinical points of view empha­ mspected al. , 1998). Double and even triple infec­ sizing the necessity of differential dia­ �al data tions of both vector ticks were common, gnosis between different forms of infec­ :r et al. , but such mixed infections occurred most tion. Magnarelli et al. (1995) stated that often in I. persulcatus ticks. Two out of very often cases of serologically confir­ ve provl­ three I. persulcatus ticks which were in­ med Lyme borreliosis are not investiga­ ons with fe cted with TBE also contained Borrelia ted fo r Babesia or Ehrlichia antibodies. Fihrlichia spirochetes. One tick contained the TBE They concluded - and we agree with u lato in antigen and Borrelia garinii, and the one them - that multiple tick-associated ill­ itchell et contained TBE, B. garinii and Borrelia nesses have to be suspected in endemic [cks with afzelii. Ehrlichia species have not until zones, and thus tick-bitten patients : typical, recently been detected in European I. should undergo a more extensive labo­ l2 out of ricinus ticks (Cinco et al. , 1997, 1998; ratory examination. 90

Scho Acknowledgements tectic The research described in this publi­ demicity in Italy. J Microbiol Infect Dis 1998 the (In press) cation was made possible in part by Micr Grant "Biological Diversity-97" (2-69) Dumler JS, Dotevall L, Gustafson R, Von The State Science and Technology Pro­ Granstrom M. A population-based seroepi­ HS, demiologic study of human granulocytic gramme, in part by a Danish Grant CPR Gran 210474-3636, in part by Grant N 98-04- ehrlichiosis and Lyme borreliosis on the ehrli 49899 from the Russian Basic Research West Coast of Sweden. J Infect Dis 1997; Micr 175: 720-2 Foundation. Zlob Goodman JL, Nelson C, Vitale B et al. Di­ Kozl rect cultivation of the causative agent of hu­ References ceph man granulocytic ehrlichiosis. New England Ahkee S, Ramirez J. A cause of concurrent conf J Med 1996; 334: 209- 15 Lyme meningitis with ehrlichiosis. Scand J rial i Magnarelli LA, Dumler JS, Anderson et Micr Infect Dis 1996; 28: 527-8 JF al. Coexistence of antibodies to tick-borne Alekseev AN, Dubinina HV, Antykova LP et pathogens of babesiosis, ehrlichiosis, and al. Tick-borne borreliosis pathogen identifi­ Lyme borreliosis in human sera. J Clin cation in Ix odes ticks (Acarina, Ixodidae) Microbiol 1995; 33: 3054-7 collected in St. Petersburg and Kaliningrad Baltic Regions of Russia. J Med Entomol Mitchell PD, Reed KD, Hofkes JM. Im­ 1998; 35: 136-42 munoserologic evidence of coinfection with Borrelia burgdorfe ri, Babesia microti, and Antykova LP, Vershinsky BV, Pyzina LV et human granulocytic Ehrlichia species in re­ al. Monitoring of fo ci of tick-borne en­ sidents of Wisconsin and Minnesota. J Clin cephalitis and tick-borne Lyme borreliosis in Microbiol 1996; 34: 724-7 St. Petersburg: map modelling of its results Pancholi P, Kolbert CP, Mitchell PD et al. for the purpose of ecological-epidemio­ Ixodes dammini logical assessment of the territory (In Russi­ as a potential vector of hu­ man granulocytic ehrlichiosis. J Infect Dis an). In: Alimov AF, ed. Ecological monito­ 1995; 172: 1007-12 ring of parasites. St. Petersburg: Zool Inst RAS, 1998: 18-20 Petrovec M, Fur1an SL, Zupanc TA et al.

Barlough JE, Madigan JE, Kramer VL et al. Human disease in Europe caused by a gra­ Ehrlichia Ehrlichia phagocytophila genogroup rick­ nulocytic species. J Clin Microbiol ettsiae in ixodid ticks fr om California col­ 1997; 35: 1556-9 lected in 1995 and 1996. J Clin Microbiol Rejtchuk EA, Chernukha YuG, Petrov EM. 1997; 35: 2018-2 1 Lyme-disease in Kaliningrad region. XII

Cinco M, Padovan D, Murgia R et al. Coex­ Conf. on Natural Nidality of Diseaset. istence of Ehrlichia phagocytophila and Bor­ Moscow: Ministry of Health USSR, 1989: relia burgdorfe ri sensu lato in Ixodes ricinus 23-4 (in Russian) fr om Italy as determined by 16S rDNA sequ­ Rijpkema S, Golubic D, Molkenboer M et al. encing. J Clin Microbiol l997; 35: 3365-6 Identification of fo ur genomic groups of Borrelia burgdorfe ri Ix odes ri­ Cinco M, Padovan D, Murgia R. et al. Infec­ sensu lato in cinus tion rate of !. ricinus (Acarina: Ixodidae) collected in a Lyme borreliosis ende­ with Borrelia burgdorfe ri sensu stricto, Bor­ mic region in northern Croatia. Exp Appl relia garinii, Borrelia aft elii and group VS Acarol 1996; 20: 23-30 I I 6 in two areas of low and high rate of en- 91

Schouls LM, Van de Pol lE, Schot CS. De­ tection of Ehrlichia and Borrrelia Species in 1998 ct Dis the Dutch Ixodes ricinus Ticks. J Clin Microbiol l998 (in press)

;tafson R, V on Stedingk LV, Gurtelschmid M, Hanson �d seroepi­ HS, Gustafson R, Dotevall L, Engvall EO, ranulocytic Granstrom M. The human granulocytic ;is on the ehrlichiosis (HE) agent in Swedish ticks. Clin Dis 1997; Micro bioi Infect 1997; 3: 573-4

Zlobin VI, Mamaev LV, Dzhioev YuP, I et al. Di­ Kozlova IV. Genetic types of tick-borne en­ gent of hu­ cephalitis virus. In: Zlobin VI, ed. Intern sci w England conf "Tick-borne viral, rickettsial and bacte­ rial infections". Irkutsk: Inst. Epiderniol. and �rson JF et Microbiol., 1996: 39-40 (in Russian) tick-borne 1iosis, and ra. J Clin

; JM. Im­ �ction with icroti, and :cies in re­ ota. J Clin

PD et al. ;tor of hu­ Infect Dis

TA et al. by a gra­ Microbiol

etrov EM. gion. XII Diseaset. SR, 1989:

er M et al. groups of Ixodes ri­ osis ende­ Exp Appl Bull Scand Soc Parasitol 1998; 8 (2): 92-96

PARASITES OF ATLANTIC SALMON (SALMO SALAR) AND BROWN TROUT (SALMO TR UTTA) FROM THERIVER AKERSELV A, OSLO, NORWAY

* Even Thoen1, 0yvind Haugland1 and Erik Sterud2'

1Norwegian College of Veterinary Medicine, P. 0. Box 8146 Dep., N-0033 Oslo, Norway 2National Veterinary Institute, P. 0. Box 8156 Dep., N-0033 Oslo, Norway

Abstract Parr of Atlantic salmon and brown way, by examining parr of these species trout from the River Akerselva, Oslo, from different seasons in the River Akers­ Norway, were examined for parasites. A elva, Oslo. total of fifteen parasite species were found. Of these were 5 ciliates, 1 myxo­ Materials and methods sporidian, 2 monogeneans, 3 trematodes The fish were caught August 28, 1997 and 4 nematodes. (water temperature: 22 °C), November 3, 1997 (4 °C) and May 22 1998 (12 °C) by Introduction electro-fishing in the River Akerselva, The parasite fauna of Norwegian sal­ Oslo. The fish were kept alive (species monids is fairy well known. There are separated) in river water in the laboratory numerous works dealing with individual until examination. They were killed by a parasite species (Huitfeldt-Kaas, 1927; sharp blow to the head. All external and Vik, 1954, 1959a,b, 1962, 1964; Hal­ internal organs were examined for para­ vorsen, 1970; Halvorsen & MacDonald, sites by the aid of dissecting- and phase 1972; Lien & Borgstry>�m, 1973). Included contrast light microscopes. If possible, here are also many works on Gyrodacty­ the number of each parasite species was lus safaris (see references in Mo, 1994). determined, and this is presented as inten­ However, only few works present data on sity range in Table 1. Ecological terms the total parasite fauna of salmonids follow Bush et al. (1997). (Kennedy, 1977, 1978; Bristow, 1991 and The species determinations are Bristow & Berland ,1991a,b). The aim of based upon: Bykhovskaya-Pavlovskaya the present study was to further increase et al. (1962) (trematodes), Lom & Dyk­ the knowledge of the freshwater parasite ova (1992) (protozoans, myxosporidi­ fauna of Atlantic salmon Salmo salar L. ans), Malmberg (1970) (Gyrodactylus), and brown trout Salmo trutta L. in Nor- andMoravec (1994) (nematodes).

* Corresponding author

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93

Table 1. List of parasites found on Salmo salar and Salmo trutta from the River Akerselva.

No. inf = number infected, Int = intensity range (? = int. not recorded). Abbreviations: BC = body cavity, CV = corpus vitreum, F = fins GA = gill arches, GF = gill filaments, I= intestine, L = lens, LI = liver, S = skin, ST = stomach,

Season Summer Autumn Spring month (water temperature) August (22 oq November (3 oq May (12 °C) Host species (number of specimens examined) Sa!mon(10) Trout(?) Salmon(lO) Trout(10) Salmon(4) Trout(8) Weight (range, in g.) 13-55 9-72 10-41 14-167 12-32 10-29 Length (range, in mm) 106-162 99-187 96-156 116-244 98-141 105-134 Parasite species Site No. Int. No. Int. No. Int. No. Int. No. Int. No. Int. inf inf inf inf inf inf PROTOZOA Ep istylis lwoffi Faure-Frerniet, 1943 S,F,GA 4 ? 1 ? Apiosoma sp. s 1 ? 1 ? 1 ? 1 ? 5 ? Riboschyphidia sp. s 5 ? 6 ? 2 ? Trichodina sp. S,GF 1 ? 2 ? 1 ? 1 ? Chilodonella piscicola (Zacharias, 1894) GF 1 ? MYXOZOA Chloromyxum truttae Leger, 1906 LI 1 ? MONOGENEA Gyrodactylus derjavini Mikhailov, 1975 S, F 1 4 1 1 6 1-10 9 3-25 3 3-12 8 3-70 Gyrodactylus magnificus Malmberg, 1957 S, F 1 1 TREMATODA Crepidostomum metoecus (Braun, 1900) I 2 8-25 2 1-2 4 2-8 Diplostomum sp. L, CV 9 10-20 6 10-40 10 6-35 9 20-50 4 5-9 7 20-80 Ichthyocotylurus sp. BC, PC 1 2 1 1 2 1-3 NEMATODA Cystidicoloides ephemeridarum (Linstow, 1872) ST 1 1 1 1 1 1 I 2 1 1 Pseudocapillaria salvelini (Polyansky, 1952) I 1 1 Rhabdocona oncorhynchi (Fujita, 1921) I 4 1-4 7 1-30 5 2-35 4 4-40 7 2-30 Raphidascaris acus (Bloch, 1779) I 1 1 5 1-3 94

Results and Discussion the lens and corpus vitreum in 92 % of than The results are presented in Table 1 the fish (n= 49). The inten-.ity was nor­ there and discussed below: mally high (range: 5-80). Owen et al. cies f "" Five ciliate species were found on the (1992) found that low level infection (4-7 C; external surfaces of the fish. Chilo­ metacercariae per eye) significantly re­ found donella piscicola was found only on the duces the vision and thereby the prey trout. salmon, and only trout harboured capture ability of sticklebacks Gastero­ found accor Ep istylis lwoffi ; a colonial ciliate recog­ steus aculeatus. McKeown & Irwin nised by the horse-shoe shaped macronu­ (1996) showed that Diplostomum spp. salmc cleus of the zooids. Ep istylis sp. indirectly caused mortality in roach Ru­ can S( (probably E. lwoffi ) has been found on tilus rutilus, due to increased susceptibil­ several occasions in Norway before, but ity to predation by bird and fish. This is Ackn We t has not been determined to species level. supported by various authors (Crowden Popp( For ciliates in general we found lower & Broom, 1980; Field & Irwin, 1994). In with ' prevalences in the summer sample. High light of these investigations, Diplosto­ water temperatures and low water level mum could have a significant negative normally stress fish and favour ciliate re­ effect on the populations of salmon and Refer production. The present results do not trout in the RiverAkerselva. Bristo: reflect these facts. The reason for this is Four species of intestinal nematodes salmo1 unknown. were found in the fish. The most common The 3 Gyrodactylus derjavini is commonly species was Rhabdocona oncorhynchi. Parasi found on salmon and brown trout in riv­ This finding was presented in a separate Augus ers draining to the Oslo fjord" In our communication by Sterud et a/.(1998) Bristo· study it was present in all samples of both and represented then the first record of some salmon and trout with a marked decrease this species from brown trout in Europe. salmot in both prevalence and intensity in the Adult males and females of Rhaphidas­ of NOI summer sample. According to Mo (1997), caris acus were found in the intestine of tions 1991; j this is a probable result of temperature both salmon and trout. To the best of our induced immunity. One individual of G. knowledge, this nematode is not previ­ Bristo' magnificus was found on a fin of a spring ously known from salmonids in Norway. sites o sampled trout. The minnow Phoxinus Brown trout is regarded as a common fi­ Norwa phoxinus is the type host of G. magnifi­ nal host for R. acus under European con­ 15th s ety for cus, which has never previously been ditions, while Atlantic salmon is not on sal a, ' found on trout in Norway. However, it is the list of final hosts (see Moravec, 1991; suspected that the monogenean could 1994). The nematode was commonly have been transferred via the net used found in the spring sample (5 of 8 trout Bush , AW, P when electro-fishing instead of an natural infected). An occurrence only in the terms: host-switch. This was shown to be possi­ spring is in agreement with previous 1997; l ble for other Gyrodactylus species under European findings (Moravec, 1994). One experimental conditions by Mo (1987). adult female of Pseudocapillaria salve­ Bykho• The most common parasite in the pre­ lini was found in an autumn sampled Dubini sent survey was Diplostomum sp. The trout. Although this is a holoarctic spe­ freshw: metacercariae of this fluke were found in cies commonly found in other salmonids 95

1 92 % of than its type host Salvelinus alpinus, SSSR. Zool Inst, No. 80, Leningrad, 1962. (Translated from Russian by the Israel Progr was nor­ there are no published records of this spe­ Scien Transl, Jerusalem, 1964) ren et al. cies fr om Norwegian trout. :ction (4-7 Cystidicoloides ephemeridarum was .1 Crowden AB, Broom DM. Effects of the eye­ :::antly re- found in t11e stomach of both salmon and ' fluke, Diplostomum sp athaceum, on the be­ the prey trout. C. ephemeridarum has never been haviour of dace (Leuciscus leuciscus). Anim Gastero­ found in salmon in Norway before, but Behav 1980; 28: 287-94 according to Moravec (1994), Atlantic & Irwin Field JS, Irwin SWB. The epidemiology, num spp. salmon is one of many salmonids which treatment and control of diplostomiasis on a ·oach Ru­ can serve as host for this nematode. fish fa rm in Northern Ireland. In: Pike A W, Lewis eds. Parasitic Diseases of Fish. lsceptibil­ JW, Acknowledgements Dyfed: Samara Publishing Ltd, 1994: 87-100 h. This is We thank Drs. Tor Atle Mo, Trygve (Crowden Halvorsen 0. Studies on the helminth fauna Poppe and Chris Appleby for their help 1994). In of Norway XV: On the taxonomy and biolo­ Diplosto- with various parts of this study. gy of plerocercoids of Diphyllobothrium Cobbold, 1858 (Cestoda, Pseudophyllidea) negative from North West Europe. Nytt Mag Zool lmon and References 1970; 18: 113-74 Bristow GAB. On the myth of protozoan free 1ematocles salmonids in Norway's freshwater systems. Halvorsen 0, Macdonald S. Studies on the common The 3rd Int. Sump. on Problems in Fish helminth fauna of Norway XXVI: The dis­ :Jrhynchi. Parasitology, Abstracts of Reports, 14-21 tribution of Cyathocephalus truncatus (Pallas) in the intestine of brown trout separate August 1991. Petrozavodsk, Russia, p.18 (Salmo trutta L.). Norw J Zoo! 1972; 20: al. (l998) Bristow GA, Berland B (a). A report on 265-72 �ecord of some metazoan parasites of wild marine Huitfeldt-Kaas H. Cyathocephalus truncatus t Europe. salmon (Salmo salar L.) from the west coast :aphidas­ of Norway with comments on their interac­ Pall. als Ursache von Fisch-Epizootien. Nytt Mag Naturvidensk B 1927; 65: 145-51 :estine of tions with fa rmed salmon. Aquaculture 1991; 98: 311-8 �st of our Kennedy CR. Distribution and zoogeo­ Jt previ­ Bristow GA, Berland B (b). On some para­ graphical characteristics of the parasite Norway. sites of salmonids from a tarn in western fauna of Char Salvelinus alpinus in Arctic Norway, including Spitsbergen and Jan nmon fi­ Norway. In: Uggla A, ed. Proceedings of the Mayen islands. Astarte 1977; 10: 49-55 'ean con­ 15th Symposium of the Scandinavian Soci­ ety for Parasitology, 4-5 October 1991, Upp­ s not on Kennedy CR Studies on the biology of sala, Sweden . Bull Scand Soc Parasitol vloravec, Eubothrium salvelini and E. crassum in 199 I; I: 71 )ffiffiOnly resident and migratory Salvelinus alpinus and Salmo trutta and in S. salar in North f 8 trout Bush AO, Lafferty KD, Lotz JM, Shostak Norway and the islands of Spitsbergen and in the A W. Parasitology meets ecology on its own terms: Margolis et al. revisited. J Parasitol Jan Mayen. J Fish Biol 1978; 12: 147-62 previous 1997; 83: 575-83 �4). One Lien L, Borgstr0m R. Studies on the a salve­ Bykhovskaya-Pavlovskaya IE, Gusev A V, helminth fauna of Norway XXXI: Distribu­ tion and seasonal occurrence of Proto­ sampled Dubinia MN, et al. Key to parasites of cephalus sp. Weinland, 1858 (Cestoda: Pro- �tic spe­ freshwater fish of the U.S.S.R.. Acad Nauk llmonids 96 teocepbala) in brown trout, 5'almo trutta L., Vik R. Investigations on the pseudophylli­ from southern Norway. Norw J ZOOl 1973; dean cestooes of fish, birds and mammals in A 21: 293-7 the n0ya water system in Tr0ndelag. Part I. SCAr Cyathocephalus truncatus and Schisto­ Lom J, Dykova I. Protozoan parasites of cephalus solidus. Nytt Mag Zool 1954; 2: 5- Minul fishes.Amsterdam: Elsevier, 1992 51 Prese1 Malmberg G. The excretory systems and the Vik R. Studies on the helminth fauna of marginal books asa basis for the systematics Norway. Ill. Occurrence and distribution of of Gyrodactylus (Trematoda, Monogenea). Trianophorus robustus Olsson, 1892 and T. Agene Ark Zool, Ser 2 1970; 23 : 1-235 nodulosus (Pallas, 1760) (Cestoda) in Nor­ McKeown CA, Irwin SWB. Accumulation of way. Nytt Mag Zool l959; 8: 64-73 Diplostomum spp. (Digenea: Diplostomati­ Vik R. Bendelorm hos ferskvqnnsfisk i dae) Metacercariae in the eyes of 0+ and 1+ Norge. Norges Jeger- og Fiskerforbund, Jo­ roach (Rutilus rutilus). Int J Parasitol 1997; hansen & Nielsen Boktrykkeri, Oslo, Nor­ 27: 377-80 way 1959: 55 Mo TA. Seasonal occurrence of Gyrodac­ Vik R. B0rsteiglen funnet i Norge. Fauna tylus derjavini (Monogenea) on Brown trout, 1962; 15: 31-6 Salmo trutta, and Atlantic salmon, S. salar, in the Sandvikselva river, Norway. J Parasi­ Vik R. Studies of the helminth fauna of tol 1997; 83: 1025-9 Norway. V. Plerocercoids of Diphyllo­ Re O) ' bothrium spp. from the R0ssaga water sys­ Mo TA. Status of Gyrodactylus safaris Re 1) tem, Nordland County. Nytt Mag Zool 1964; problems and research in Norway. In: Pike proble 5: 25-93 A W, Lewis JW, eds. Parasitic Diseases of memb Fish. Dyfed: Samara Publishing Ltd, 1994: Re 2) 43-56 memb( Mo TA. Taxonomiske og biologiske under­ numbe s0kelser. Virksomheten i 1986 og forslag til virksombeten i 1987. Rapport nr. 2. Gyro­ dactylusunders0kelsene ved Zoologisk Mu­ seum, Universitetet i Oslo 1987.

Moravec F. Parasitic nematodes of freshwa­ ter fishes of Europe. Dordrecbt: Kluwer Academic Publishers, 1994 It was Owen SF, Barber I, Hart PJB. Low-level in­ soon a1 fection by eye fluke, Diplostomum spp, af­ change fects the vision of 3-spined sticklebacks, further Gasterosteus-aculeatus. J Fish Bioi 1993; 42: compla 803-6 send Ol Sterud E, Tboen E, Haugland 0. First record Re 3) 1 of Rhabdocona oncorhynchi (Nematoda: student Rbabdoconidae) from brown trout in Europe. Bull Bur Ass Fish Patbol l998; In press was ag should 97 S()ClE'fY N.EWS �udoph ylli­ ammals.�n Iag. i>art' I. SCANDINAVIAN SOCIETY FOR PARASITOLOGY Schisto- 1954; 2: 5- Minutes from 1he BmmJ telephone meeting 06. October, 1998: ' Present: Tcllcrvo Yaltonen, Jorun Tharaldsen, Ingela Krantz, KarJ, Skirnisson, Tor Atle 1 fauna of Mo, Katarina Gustafsson and Maria Vang Johansen ibution of Agenda for the meeting: �92 and T. 0. Comments to the minutes from the last meeting. a) in Nor- 3 1. Latest news from ICOPA in Japan (including sorting out the problems - Federation.cotrespondence) and news from the Polish jubilee meeting. annsfisk 'i. 2. Question of collecting the fees and acquiring new members. (Tor-Atle and 1rbund, Jo­ the local treasures from each country should tell about the local situations). Jslo, Nor- 3 New members, acceptance of applicants. 4. The next meeting in Iceland. (Karl will info rm). ·ge. Fauna 5. Latest news on the WAAVP in Copenhagen (MVJ will brief). 6. Other issues. fauna of Re 0) The minutes from October 29, 1997 were accepted without comments. Diphyllo­ water sys­ Re 1) Tellervo attended the ICOPA meeting in Japan, where she sorted out the Z0011964; problems with Professor van Knapen, World Federation for Parasitology, and paid our membership fee and exchanged new addresses.

Re 2) Norway, Finland, Iceland and Denmark have sent out payment forms to their members. Sweden is still waiting to get the forms fr om the giro office. Until now, the number of people who has paid is: 8 non-Scandinavians; 25 in Norway, (18 missing); 9 in Finland, (? missing); 18 in Denmark (19 missing); 5 in Iceland, (all have paid). Ingela informed that Sweden has about 70 members. It was agreed that all local treasures should send an updated membership list to Tor as soon as possible and keep him informed about changes. It was further suggested that we change to Euro accounts, which will be possible from next year. This needs to be further discussed. All local treasures could further inform that no members had complained about the paying procedure, but it was agreed that we from now on shall send out payment forms every year.

Re 3) Five new members fr om Finland and one fr om Denmark were approved. Ph.D. students are not regarded as students, hence they have to pay full membership fee. It was agreed to make an active move towards getting new members to SSP. Everyone should advocate for the society and the meeting next year. . 98 Re 4) Prior to the meeting Karl had sent out information about the Iceland meeting. So far ·40 ·people have responded to the first announcement. The Second Announcement will be sent out within the next month to all members. New members, although not ' ' formally accepted by theBo ard, will receive the announcement as well. Pro It was further agreed that the announcement should be sent to the Russian, the Polish, Sep and the Baltic Societies. Parasitology Today as well as the European and World soq Federation for Parasitology should also be informed. Ch; Re 5) Maria had nothing to report. Dcx Re 6) Jorun asked for inputs to the Bulletin, which she has not gotten from any of the Uni members. Advertisements of meetings, Ph.D. defences, awards, employments as well as scientific inputs are highly wanted. The next number of the Bulletin will be sent out Dar shortly and one more will come before Christmas. So please, help Jorun with pan1 information. Due to trouble sending mails, Maria will send out a new revised e-mail address list. Ph .I Roy Please remember to inform the secretary when addresses are changed.

------0------Pitt a 41 07. 10.98 Maria Vang Johansen ncnt

A d Pari Hcln Tan! :'c Dan} intc� Schlj DisC; Heal Wol'l

Dani "Myi fOCU! partil Eriks 99 �ting. So tncement front I h•1nnark mgh not Prolt'',""i f't'IN Nansen, Danish Centre for Experimental Parasitology, was in e Polish, St'plt'lllh;'t 1'1\JH ckcled Honorary Member of thePolish Society for Parasitology at the 1 World st wkty\ '\!1Vt'lUS anniversary.

Clmrlollt

Ph.D. has been awarded a 5-years research professorship at the ress list. Stig Milan Thamsborg Royal Veterinary and Agricultural University in the area organic husbandry systems.

Ph.D. Michael Larsen, Danish Centre for Experimental Parasitology, has been granted a 2-years senior research fellowship from the Lundbeck Foundation - to study nematode-killing microfungi.

A group of researchers (headed by Peter Nansen) from Danish Centre for Experimental Parasitology has been granted an exten..o;;ion (1998-2001) of the ongoing Ruminant Helminth Research Project. The project is supported by Danida and includes Kenya, Tanzania, Zambia and Zimbabwe. The coordinator is Ph.D. Lee Willingham.

Danish Centre for Experimental Parasitology (DCEP) took the initiative to an international workshop: "The epidemiology and host-parasite relationships of Schistosoma japonicum ", September 8-10, 1998, Jiangsu Institute of Parasitic Diseases, Wuxi, Peaples Republic of China (PRC). Sponsors were: CEP, Ministry of Health (PRC), SIDA, WHOffDR, FAO. Totally 62 participants from 11 countries. Workshop coordinator: Dr. Lee Willingham.

Danish Centre for Experimental Parasitology arranged an international workshop on "Mysteries of Ascaris: What they are, how they behave and what to do about them. A focus on Ascaris in pig and man" October 8-11, 1998, Copenhagen. Totally 52 participants from 18 countries. Workshop coordinators Allan Roepstorff and Lis Eriksen. 100 198 autl and GUIDELINES FOR CONTRIBUTORS to tJ may If tl All contributions should be submitted as word-processed manuscripts on floppy eo m disk, accompanied by two exactly matching print-outs of good reading-quality. The inse preferred storage medium is a 3lh inch disk in MS-DOS or Windows compatible format. The text should be written in Word or WordPerfect or other word processing l programs convertible to these. With a Macintosh computer, save the file in the MS­ Stm DOS compatible option. Please indicate the word processor (and version) used to generate the file, the type of computer, the operating system, and the formatted capacity Anc of the diskette. Hor The articles/communications should normally not exceed 4 printed pages, including utpr tables, figures, and references, and may contain a maximum of 2000 words if there are Lun no figures or tables. The first page should show the title of the article, and the name(s) som of the author(s). The authors' addresses should be given, and the complete correspon­ dence address with telephone and telefax number (if available). The text should follow, Boo without subheadings, but a short summary, maximum 100 words, may be included. Aus The text should be typed unjustified (unaligned right margins), without hyphenation Chit (except for compound words), and at 1 Y2 line spacing. Do not type page numbers. Md Label the hard copies by hand at the bottom of the page. Please ensure that the digit 1 CE< and the letter 'I' have been used properly, likewise with the digit 0 and the letter '0'. Do vete not use decorative formatting, such as boldface and centred headings, or underlining of Sosi titles or subheads. fag� Authors are obliged to follow the rules governing biological nomenclatures, as laid 1) down in e.g. the International Code of Zoological Nomenclature. Disease names Tho should follow the principles of Standardized Nomenclature of Parasitic Diseases Nev (SNOPAD). Figure legends must be included on the diskette, but the figures will be handled 1 conventionally. They should be marked on the back with the title of the article and yoUJ name of the (first) author. autt the Line drawings should be provided as good quality hard copies suitable for reproduction as submitted. ace( Photographs must be provided as glossy prints, and be of sufficiently high quality to allow reproduction on standard (not glossy) paper. Colour plates will not be printed. References in the text shold be stated by giving in brackets the name of the author and the year of publication, e.g. (Thornhill, 1987) or (Austin & Austin, 1987). If there In t are more than two authors, only the first name plus et al. is given (Lund-Larsen et al, imp 1977). The reference list should be in alphabetical order, and follow the style set forth in Uniform Requirements to Manuscripts Submitted to Biomedical Journals, Br Med J 101

1988; 29<1: 110 1 '1 1\dnnH'!'S to journals should contain names and initials of the

authors , ariH'k titk, tlw abbreviated name of the journal, year of publication, volume, and Jirst and la•;t pitl�t· numbers of the paper. Journals should be abbreviated according to t he "J .ist ul jumnals indexed in Index Medicus". Authors without access to this list may type the litll name of the journal, and the Editor will take care of the abbreviations. If there arc more than six authors, list only the first three and add 'et al'. Personal 1 floppy communications and unpublished data should not be used as references, but may be ity. The inserted in tlh� text (within parenthesis marks). mpatible ocessing Ex amples of correct forms of references are given below: the MS- Standard journal article: used to capacity Anonymous. Some facts on small animal practice. Vet Rec 1987; 120: 73 Horsberg TE, Berge GN, H0y T et al. Diklorvos som avlusningsmiddel for fisk: klinisk - ncluding utpr0ving og tok.;;isitetstesting. Nor Vet Tidsskr 1987; 99: 611 5 there are Lund-Larsen TR, Sundby A, Kruse V, Velle W. Relation between growth rate, serum name(s) somatomedin and plasma testosterone in young bulls. J Anim Sci 1977; 44: 189-94 1rrespon­ i follow, Books and other monographs: kd. Austin B, Austin DA. Bacterial fish pathogens : disease in farmed and wild fish. henation Chichester: Ellis Horwood, 1987 mmbers. McFerran JB , McNulty MS, eds. Acute virus infections of poultry: a seminar in the e digit 1 CEC programme, Brussels 1985. Dordrecht Martinus Nijhoff, 1986. (Current topics in :'0'. Do veterinary medicine and animal science 37) lining of Sosialdepartementet. Tsjernobyl-ulykken: Rapport fr a Helsedirektoratets ractgivende faggruppe. Oslo: Universitetsforlaget, 1987 (Norges offentlige utredninger NOU 1987: , as laid 1) � names Thornl1ill JA Renal endocrinology. In: Drazner FH, ed. Small animal endocrinology. f"Jiseases New York: Churchill Livingstone, 1987: 315-39 handled The manuscript (diskette and paper copies) should be sent to one of the editors in kle and your country, see inside of back cover. Label the diskette with the name of the (first) author. Manuscripts are accepted for publication after review and recommendation by the Editorial Board. Authors will be notified by the Editor-in-Chief about final 1ble for acceptance and expected time of publication. uality to rlted. REPRINTS WILL NOT BE A V AILABLE. e author If there In the interest of speed, no proofs will be sent to authors. It is therefore of vital en et al, importance that the manuscripts are carefully checked before submission. set forth tr Med J !I

DcnHHH it FknlHllllF, h Vel, a11d forlu ll!, hhl lni h nl ;i,ll•l Molec 1\ml"f'V, l U, DK I H /!l hnl11blwl I (Tel: 11\'J l I hn· i"! WF!h•H: 145 \'l ZH)/14 ) li;h;l!l lli'>1dund c-nmil: cco)(nlk vl.dk h·ehmd: Nntwnal Vet lns!.lSwedish SJgun'luJ Rich!cr, tJnivtJsity I Jniv Agric. �-;dt•n!.,l kpl

Maria Vang .Johanscn, oflccland, I nst . forExp. ol l'amsitoL, 1' .0. Box 707.\, Danish Bilharzia0is Lab. , Pathol. Kcldur, P.O. Box �; 7 SO 07 \ Jppsala, Jregcrsborg Allc 1 D, 8540, JS-112 Reykjavik (Tcl : 14(1 I X674 1 5(J, Fax: DK-2920 Charlottcnlund (Tcl: +354 5674700, Fax: I 4(J I x:109 ! 62) e-mail: (Tel: +45 77327732, Fax: +354 5673979) Johan.I loglund(a)sva.se +45 77327733) e-mail: e-mail: [email protected] mvj @bilharziasis.dk Lars-Ake Nilsson, Karl Skirnisson, University University of Gotcborg, Inst. Eskild Petersen, Statens of Iceland, Inst. fo r Exp. of Med. Micro bioi. & Seruminstitut, Lab. of Pathol., Keldur, IS-112 Immunol., Guldhedsgatan Parasitology, DK-2300 Reykjavik 10, S-413 46 Goteborg Copenhagen S (Tel +354 5674700, Fax: (Tel: 46 31 6047 17, Fax +46 (Tel: +45 32683223, Fax: +354 5673979) 31 604688) +45 32683033) e-mail: e-mail: [email protected] [email protected] Jan Thulin Norway: National Board of Fisheries, Finland: Tor A Bakke, Zoological Inst of Marine Research, Margaretha Gustafsson Museum, University of P.O. Box 4, S-453 21 Abo Akademi, Dept. of Oslo, Sarsgt. 1, N-0562 Oslo Lysekil (Tel: +46 52314180, Biol., BIOCITY, Artillerigt. (Tel: +47 2285 1 678, Fax: Fax: +46 523 13977) e-mail: 6, FIN-20520 Abo (Tel : + +47 2285 1 837) e-mail: [email protected] 358 2 12654603, Fax: +358 t.a. bakke@toyen. uio.no 212654748) e-mail: Editor of Baltic News: [email protected] Bj0rnGj erde, Norwegian Peter Nansen, Danish Ctr. of Coll. Vet. Med., Dept. of Exp. Parasitol, Royal Vet. Hannu Kyronseppa, Parasitol. P.O. Box 8146 and Agric. Univ., Auroran Sairaala, Dep., N-0033 Oslo (Tel: Biilowsvej 13, DK- 1870 Nordenskioldsgt. 20, +47 22 964969, Fax: +47 Fredriksberg C, (Tel: +45 FIN-00250 Helsinki 22964965) e-mail: 3528 2780, Fax: +45 3528 (Tel: +358 9 4715983, Fax: bj [email protected] 2774) e-mail: +3 58 9 471 5900) [email protected] BULLETIN OF THE SCANDINA VIAN SOCIETY FOR PARASITOLOGY

VOL. 8 No. 2 CONTENTS December 1998

Proceedings of the Baltic-Scandinavian Symposium on Ecology of Bird-Parasite Interactions, Vilnius, Lithuania, 25-28 June, 1998

Gediminas Va lkiiinasand Hans-Peter Fagerholm (Editors) ...... ]

First determination of Ehrlichia infected ticks among the primary vectors of the tick­ borne encephalitis and borreliosis in the Russian Baltic region

A. Alekseev, H. Dubinina and L. Schouls ...... 88

Parasites of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) from the river Akerselva, Oslo, Norway

E. Th oen, @. Haugland and E. Sterud ...... 92

SOCIETY NEWS

Minutes from Board meeting 6 Oct. 1998 ...... 97

News from Denmark ...... 99

Guidelines for contributors ...... I 00