Bulletin of the
SCANDINAVIAN SOCIETY
FOR PARASITOLOGY
• , · . ,.._ WITH PROCEEDINGS OF THE SYMPOSIUM ON ECOLOGICAL , . ;'PARASITOLOGY ON THE TURN OF MILLENIUM, ST. PETERSBURG,
• 'i: RUSSIA, 1-7 JULY, 2000 ({ • . o<> ��V vol. 10 No. 2 2ooo ;:;:y BULLETIN OF THE SCANDINA VIAN SOCIETY FOR P ARASITOLGY
The Bulletin is a membership journal of the Scandinavian Society for Parasitology. Besides membership information, it also presents articles on all aspects of parasitology, with priority given to contributors fi-om the Nordic countries and other members of the Society. It will include review articles, short articles/communications. Comments on any topic within the field of parasitology may be presented as Letters to the Editor. The Bulletin is also open for a short presentation of new projects. All contributions should be written in English. Review articles are commissioned by the editor, however, suggestions for reviews are welcomed. Subscriptions are available to non-members upon request from the Publisher. The subscription rate is 40 EURO per year (two issues annually). Subscriptions should be paid to the treasurer of the SSP:
Tor Atle Mo National Veterinary Institute P.O. Box 8156 Dep. N-0033 Oslo, NORWAY, e-mail: [email protected] Postal giro account number: 0814 3937489
Scandinavian Society for Parasitology (Nonlisk Ftircning fOrParasitologi) Society Board:
President: E. Tellervo Valtonen (Finland) Vice-President: Karl Skirnisson (Iceland) General-Secretary: Maria Vang Johanscn (Denmark) Treasurer: Tor Atle Mo (Norway) Board Member: Ingela Krantz (Sweden) Suppleants: Eskild Petersen (Denmark), Mats Wahlgrcn (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 spring. 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 oflifc.
Printed in Norway by HS-Trykk NS JSSN 0803-4907 n 8 O>ln1 Vl/\J vv t1 PROCEEDINGS Ol/\J3 of the symposium on
ECOLOGICAL PARASITOLOGY ON THE TURN OF MILLENNIUM
aiTanged on behalf of the
Russian Parasitological Society
and the Scandinavian Society for Parasitology m St. Petersburg, Russia
1-7 July, 2000
Editors: Kirill V. Galaktionov, Oleg Pugachev, Ken MacKenzie and Karl Skirnisson
Symposium secretariat: K. Galaktionov, 0. Pugachev and I. Podvyaznaya, Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab., 1, 199034 St. Petersburg, Russia. Fax:+ 812 3282941, e-mail: [email protected] ii
PREFACE
In 1994 the Scandinavian Society for Parasitology (SSP) arranged for the first time a special symposium on Parasites of Biological and Economical Significance in the Aquatic Environment in Heimaey, Iceland (see the SSP Bulletin 4.2). Based on the same idea another special symposium was arranged on behalf of the SSP in Stykkish6lmur, Iceland in 1996 that dealt with Parasites and Ecology ofMarine and Coastal Birds (SSP Bulletin 6.2). In 1998 the Baltic Society for Parasitology and the SSP arranged a joint symposium on Ecology of Bird-Parasite Interactions in Vilnius, Lithuania (SSP Bulletin 8.2) and this issue is a result of a recently organized joint symposium of the Russian Parasitological Society and the SSP in St. Petersburg, Russia on Ecological Parasitology on the Turn of Millennium. These four meetings as well as the Baltic-Scandinavian Symposium on Parasitic Zoonoses and Ecology of Parasites held in 1994 in Vilnius, Lithuania (SSP Bulletin 5.1) have brought parasitologists in this regions together and promoted a very valuable exchange of information and initiated various collaboration among Scandinavian, Baltic and Russian parasitologists. The Bulletin of the SSP has been an important forum to publish invited papers and submitted abstracts which have been presented by hundreds of parasitologists during the above mentioned meetings. Several very valuable review articles, based on invited lectures given at the meetings, have been prepared and published in the Bulletin. There is no doubt that the organisation of the special meetings has increased the attention and the scientific value of the journal.
Due to various reasons St. Petersburg was chosen to be the venue of a joint symposium of the Russian Parasitological Society and the SSP on ecological parasitology. First to mention is the fact that in the 20's and 30's Professor Valentin A. Dogiel (1882-1955) founded in the city the Russian school of ecological parasitology. His books on ecological parasitology and books on the subject written by his Russian followers were translated to English and subsequently also stimulated the development of ecological parasitology in other countries of the world. Also, a considerable number of parasitologists is working on parasites at different institutions in St. Petersburg, e.g. the Zoological Institute, the St. Petersburg State University and the Institute of Lake and R i vcr F ishcry (GosNIORKh). Many of them have continued Dogiel's tradition on vrul(l)',ical parasitology. Furthermore, close relationships exists between p:uwdlolP)',ists working in St. Petersburg and colleagues in the states of' the l\11 ll!l'l 1. I SS I\, especially, however, of those in NW Russia and in the Haiti(; wunl I PI tiKJll, as well as for western parasitologists, it was considered to Ill: caw I!> nH1w i!l St.
Petersburg. During the "Cold War" the western and eastern llfillltlw-, 1d t'1 come for the scientists of these countries to meet to present and discuss research data, exchange ideas and develop collaboration. Altogether Ill scientists from 25 countries took part in the meeting. Eleven plenary lecturers and 49 submitted oral presentations were given and altogether 73 posters were shown. Papers or abstracts based on plenary lectures and abstracts of the submitted presentations are printed in this issue. As the readers will see the presentations mainly covered various aspects of modern ecological parasitology, e.g. patterns of parasite communities, strategies of parasite life-cycles and life histories, parasite manipulations with host behaviour, parasites and wildlife management and vector's ecology. Two workshops were organised during the symposium, on "Ecological parasitological problems of NW Europe" and "Parasites of marine and coastal birds in Arctic and subarctic regions". On behalf of the Local and Scientific Organising Committees we would like to express our sincere thanks to the invited speakers, to the participants and to all other persons who contributed to the great success of the meeting. Last but not least we are very grateful to the sponsors listed below who kindly supported the sympostum. September 30, 2000 Kirill V. Galaktionov Karl Skirnisson Scientific Organizing Committee: Andrew N. Alekseev, Zoological Institute, Russia E. Tellervo Valtonen, University of JyvaskyHi, Finland Oleg N. Pugachev, Zoological Institute, Russia Kirill V. Galaktionov, Zoological Institute, Russia Karl Skirnisson, Institute for Experimental Pathology, University of Iceland Tor Atle Mo, National Veterinary Institute, Norway Kurt Buchmann, Royal Veterinary and Agricultural University, Denmark Clive Kennedy, University of Exeter, UK Gediminas Valkiunas, Institute of Ecology, Lithuania iv Local Organizing Committee: Kirill V. Galaktionov, Zoological Institute, St. Petersburg Oleg N. Pugachev, Zoological Institute, St. Petersburg Irina M. Podvyaznaya, Zoological Institute, St. Petersburg Helena V. Dubinina, Zoological Institute, St. Petersburg Andrew A. Dobrovolskij, St. Petersburg State University Sponsors: The generous support from the following sponsors is gratefully acknowledged: - Russian Academy of Sciences -Nordic Academy for Advanced Studies (NorFA) - Russian Foundation for Basic Research - "EkteparetSoerlies Fond for Viltlevende Fulger", Norway -Zoological Institute of the Russian Academy of Sciences -St. Petersburg State University - "Nevatrans ", Russia - "IRVI" Ltd, Russia V CONTENTS INVITED LECTURES Vector (tick) as ecological niche of different parasitic organisms I A. N. Alekseev & H. V Dubinina...... Selective pressures and parasite strategies 6 C. Combes ...... Ecology and genetics of Daphnia-microparasites infections: which factors determine parasite distribution? D. Ebert ...... 12 An ecological approach to the analysis of trematode life-cycles 13 K. V Galaktionov & A.A. Dobrovolskij ...... Phylogeny, history and biodiversity: understanding fauna! structure and biogeography in the marine realm 19 E. P. Hoberg & A. Adams ...... Parasite application in conservation and pest control 38 PJ. Hudson ...... Freshwater fishparasites and habitat change 39 CR. Kennedy ...... The evolutionary ecology of parasite-induced changes in host phenotype R. Poulin ...... 44 Infracommunities: structure and composition 49 O.N. Pugachev ...... What determines the longevity of mammalian nematodes? 55 A. Skorping & A. F. Read ...... Getting to the core of the parasite communitites: separating reality fromnoise 61 E. T. Valtonen, K. Pulkkinen & M Julkunen ...... SUBMITTED PAPERS - ORAL PRESENTATIONS Significanceof the amoebocyte-producting organ ofBiom phalaria glabrata snails (strainsselected for susceptibility/resistance) in cellular responseto Echimostoma caproni mother sporocyst infection 65 G.L. Ataev, A.A. Dobrovolskij, A. V Avan essian & C. Coustau ...... The main features and factors limiting Metazoan parasite commw1ities in fish of the Baltic Sea (Lithuanian coastal zone) and Curonian Lagoon 66 E. Bacevicius ...... The commwlity ofMyxosporea fiom the kidney of the roach Rutilus rutilus lacustris 67 MD Badmaeva, NM Pronin & S. V Pronina ...... Comparision of inlmune response mechanisms in rainbow trout against gyrodactylids and ciliates 67 K. Buchmann, J Sigh, C. V Nielsen, M Dalgaard, T. Lindenstro m & A. Li ...... vi Parasitic gastr·oenteritis of cattle in Sweden: importance of the ecology ofthe free-living stages S.O. Dimander, J. Ho glund & P.J. Wa ller ...... 68 Host specificity and dispersal str·ategy of gy:rodactylids in the Black Sea coastal biocenosis E. Dmitrieva ...... 68 Is ospora (Protozoa, Sporozoa) infection in passerine birds of various fe eding habits 0. V Dolnik ...... 69 Parasite community structure as an index of environmental conditions G.N Dorovskikh ...... 70 Parasitological monitoring in evacuation zone near Chemobyl NPP G. Efr emova ...... 70 The eel swimbladder nematode Anguillicola crassus (Nematoda: Dracunculoidea) in Irish eels D. W Evans & MA . Matthews ...... 7 I Fish anisakidein Khuzestan province of South West Iran A. Farahnak...... 71 The ecology ofixodidae ticks in Novgorod fo rest region under the condition of anthropogenic loading VG. Fyodorova & Z.S Lalaeva...... 72 Comparative analysis of the monogenean faunas of the Black and Baltic Seas A. Gaevskay a & E. Dmitr ieva...... 72 «Core and satellite» hypothesis: an example from fishparas ite communities P. Call i & G. Crosa ...... 73 Diversity of fish parasites in relation to environmental stress M Gelnm� S Sebelova, L. Dusek, J. Jarkovskf;, L. Machala, B. Koubkova & P. Jurajda ...... 74 Communityecology of three different monogenean parasite-host systems M Gelnar, L Matej usova, A. Simko va & S Morand ...... 7 4 Species life-cycle, developmental cycle and life scheme in parasitology (definition and correlationof terms) L.A. Ghitchenok...... 75 A study of the parasite fa una of Phox inus phoxinus L. J:i'om the small rivers of NorthernDvina basin E. Golicova ...... 75 Exploitation of gastropods by trematodes: host morphology and parasite strategy A.M Gorbushin ...... 76 Parasitic systems oftr·ematodes and the mechanisms of their regulation A.L Granovitch ...... 76 Population dynamics ofArgul us coregoni Thorell(Cmstacea: Branchiura) T Hakalahti & E. T Valtonen ...... 77 A seroepidemiological survey of Dictyocaulus vivipar us, lungworm infection of cattle in Sweden J. Ho glund, S.O. Dimander & P.J Wai ler ...... 78 vii Ecological characteiistics offish parasite faunas in north-eastem Azerbaijan rivers SR. lbrahimov & B.S. Agaeva ...... 78 Conservative flagging samples - a simple method to obtain detailed infmmation on the host-seeking activityof Ixo des ricinus PM Jensen...... 79 Variability of the parasite fauna of oceanic fish species dming different peiiods of a migration cycle (by example of blue whiting of the Northwest Atlantic) A.B. Karasev ...... 80 Genetic heterogeneity of trematode species reflects the structmeof their natmal populations K V Kha! turin, NA. Mikhailova & A.! Granovich ...... 80 The effect of host density on ectoparasite distribution B. Krasnov & I Khokhlova ...... 81 Mortalityof Atlantic salmon parr due to high intensities of eyeflukes T Larsen & F.R. Lund ...... 82 Interrelationships of crabs, leeches, fishand trypanosomes in coastal waters of North Norway K MacKenz ie, W Hemmingsen & P.A. Jansen ...... 82 Notes on mesoparasitism A. Marchenkov ...... 8 3 Patternsof similaiity in the parasitefau nas ofLar us gulls J Mellen ...... 83 How does Argulus fo liaceus L. (Crustacea: Branchiura) match withits hosts at the behavioural and ecological levels? V Mikh eev, A. Pasternak & E. T Valtonen ...... 84 Seasonal dynamics of the feather mite Monojoubertia microphylla (Astigmata: Analgoidea: Proctophyllodidae) on thechaffinch Fri ngilla coelebs S V Mironov ...... 8 5 Feeding relationships of intermediate hosts affect transmission success of larval cestodes A. Pasternak, K Pulkkinen, V Mikheev & E. T Valtonen ...... 86 Kmyotypicaldifferentiation of the species of Eubothrium (Cestoda: Pseudophyllidae) R. PetkeviCillte ...... 86 Changing risk of tick-borne encephalitis in Scandinavia with predicted climate change SE. Randolph & D.J Rogers ...... 87 Nematode fauna of sheep with emphasize to zoonoses in the west ofiran SM Sadjjadi ...... 88 Electrophoretypes, genotypes and Borrelia agents of Ix odes persulcatus ticks A. V Semenov, A.N Alekseev & H V Dubinina ...... 88 Morphophysiological and ecological analysis of ontogenesis in terrestrial parasitengona (Acm· iformes) A. B. Shatrov ...... 89 Vlll Morph ol og ical and ka riol ogical studies ofEchinochasmus sp. cercariae (Trematoda, Echinochasmidae) G. StaneviCiiite, V Kiseliene & R. PetkeviCiu te ...... 89 The ecolog ic al value of acanthocephalans for detecting traffic related pollution B. Sures ...... 90 First studies on the stress resp onse of mammals following helm inth infections and heavy metal exp osure & H ...... B. Sures, G. Scheej W Kloas Taraschewski ...... 91 Estonian Oesophagostomum spp. is olates inpi gs H Talvik, CM Christensen & T Jarvis ...... 92 Sexual ornaments signal parasite resistance in amale fish J Taskinen, R. Kortet & T Sinisalo ...... 92 Parasites in hosts with special life histories & T ...... G. Valkiu nas Iez hova ...... 93 Environmental impact assessment of parasite control measures in cattle P.J Wai ler, SO. Dimander, G. W Yeates & J Ho glund...... 93 SUBMITTED PAPERS - POSTER PRESENTATIONS Sp or oz oans (Sporozoa, Apic omplexa) - blood parasites of tundra water-swamp birds, spending winter in Azerbaijan M M & ...... Aliyev, G. Gaibova, Musayev E. Sultanov ...... 95 Comparative analysis of the helminthocen oses of native riparian mustelids (Lutra lutra, Mustella lutreola) in relation to width of food spectra E. Anisimova & V Sidorovich ...... 95 The significance of molluscs in conservation of trematode life -cyc les G.L. Ataev, E. V Koz minsky & A .A. Dobrovolskij...... 96 Parasite systems with acanthocephalans ofthe genus Filicolis Luhe, 1911 (Acanthocephala) and their geographical modifications G. I Atrashkevich...... 96 Hist ol og ical and histh oh emical changes in Tox ocara canis tissues under the action of levamisole R. Aukstikalniene, 0. Kublickiene & A. Vysniaus kas...... 97 Ecological analysis of fishparasi te fa una in Kaliningradreg ion reservoirs E. Avdeeva & E. Evdokimova ...... ··················· . . . 98 Haemosp or id ian parasites of birds from Belarus A. Babushnikova...... 98 Pomphorhynchus laevis (Muller, 1776) (Acanthocephala) remains in mesenkrium tissue of flounder (Platichthys jlesus L. ) : morphol ogy, localization and origin E. Bacevicius ...... 99 ix Gammarids of Lake Baikal basin as intennediate hosts of helminths DR. Baldanova, UA. Kritskay a & NM Pronin ...... 99 Comparative analysis of the parasite faunas of brown trout prur (Salmo trutta L.) fi-om some water bodies of the Paan�arvi National Pru·k and the River Oulru1ka Y Y Barskaya, B.S. Shulman & E.P. leshko ...... I 00 Importance of ecol ogically dete nnined characters in the tax on omy of cestodes S. Bondarenko & V Kontrimavichus ...... I 0 I Parasite communities of small rodents inthe Paanajarvi and Oulanka National Parks S. V Bugmyrin, E.P. leshko, VS.Ani kanova & L.A. Bespyatova ...... I 0 I Special fe atures of the fonnati on of rodent helminthoc oenoses in urban landscapes of Belruus E. Bychkova ...... I 02 Role of horseflies (Diptera, Tabanidae) inthe ecological balance of bi ota P.A. Chirov & A.M Peterson ...... I03 House dust mites in four Vilnius pre-schools A. Dautartiene & A. Jurkuvenaite ...... I 03 Differentiation and placental-like organization of the uterus in Nippotaenia mogurndae (Cestoda) V G. Davydov & J V Korn eva ...... I 04 End op arasites of juvenile (age 0+, 1+) cod, Gadus morhua, in Icelandic waters preliminruyresults M Ey dal, A. Kristmundsson, S. H Bambir, R. Magnusd6ttir & S. Helgason ...... I 04 How Nematodes communicate, -with a note on phasmid ultrastructure H-P. Fagerholm, M Brunaska, A. Roepstorpj G. Jungersen & L. Eriksen ...... I 05 Interc ommunications between zone of mite bite (genus Ix odes) localizati on ru1d the hwnan bioenergetical system V Fyodorova & V Kosmenco ...... I 06 Coccidia (Sp or oz oa, Apicomplexa) ofwater and marsh fowl ofthe tw1dra overwintering in Azerbaijan G. Gaibova, M Aliyev, M Musayev, E. Sultanov...... I 07 Cestodes ofthe eider duck population inhabiting Spitzbergen ru1d Franz JosefLand (preliminruy communicati on) A.K Galkin, K V Galakt ionov & SF Marasaev ...... I 07 Parasite invasion as a risk fa ctor in the development of inflammatory diseases of the pelvis organs TA. Gasanova, A .V Mikhailov & P.A. Chirov ...... I 08 Eudiploz oon nipponicum (Diplozoidae, Monogenea) - comparative analysis of microec ol ogy and habitat specificity in relation to parasite ontogenetic development M Gelnar, I Hodowi, B. Koubkowi, A. Simkova & TH Zurawski ...... I 09 Ta xonomy ru1d systematics ofthe mon ogenean frunily Dipl oz oidae in the light of recent knowledge M. Gelnar, B. Koubkova, I Hricova, M Dlouha & M Kli mova ...... I I 0 X Some characteristics of the distribution of Tetr aonchus monenteron (Mon ogenea) and Ergasilus sieboldi (Crustacea) on gi lls of pike Esox lucius P.l Gerasev ...... 110 Comparative analysis of monogenean fa unas and populations from several beloni fonn fishes L.A. Ghitchenok...... Ill The lab orat my rat as de finitive and inte nnediate host for Sarcocystis rodentifelis (Protista: Coccidia) from the black rat J Grikieniene & L. Kutkiene ...... Ill The helminth fauna of the common shrew, Sore:xaraneus L., from two sites near Vienna, Austria C. Hoerweg...... 112 Are pop ulati on dynamics of willow pta rmigan affected by parasites? P. Holmstad, P. Hudson & A. Skorping...... 113 Ec ological analysis of the fish myxosporean fauna of Azerbaijan water bodies in the light of new data ab out the biology of these parasites S.R. lbrahimov & R.K. Kerimova ...... 113 On the fonnati on of infection foc uses of echinost om osis in sea-birds in the Barents Sea D. lshkulov ...... 114 The trichinellosis situati on in Estonia T Jarvis, l Miller & E. Pozio ...... 115 Efficacy of some integrated and non-chemotherapeutic control method s against lamb monieziosis in Estonia A. Kaarma & E. Ma gi...... :. 115 Several aspects of the influence of abnonnal light conditions on the preimaginal stages of blood-s ucking mosquitoes S. Karpova ...... 116 The helminths of reptiles from South Urals V Khabibullin ...... 116 Parasites of the eel in Latvia M Kirj usina & K. Vismanis ...... 117 On the is opod fish parasite fa una of the Atlantic coast ofnmih-westem Europe A.F. Kononenko ...... 118 Isopoda parasites ofNorth Sea fishes A.F. Kononenko ...... 118 The parasite capacity of theho st populati on E. V Koz minsky...... 119 Ectoparasites of juvenile (age 0+, 1+) cod, Gadus morhua, in Icelandic waters prel iminary res ults A. Kristmundsson, M Ey dal, S.H Bambir, R. Magnusd6ttir & S. Helgason ...... 119 Parasitism as a bioindicator of ecological and seas onal characteristics of seabird populati ons (by the example of the murres fr om Seven Islands arc hipelago) V V Kukl in ...... 120 XI Assessment of the pathogenicity of seabird helminths us ing biochemical testing MM Kukl ina & VV Kukl in ...... 121 Coprophagy and transplacental transmission as possible ways of distribution of some sarc osp orid ian (Protista: Coccid ia) species in rodents L. Kutkiene & J Grikieniene ...... 121 A new disease in perch (Percajluviatilis): occtmence of lymph ocystis in Finnish lakes KM Leskis enoja, E. T Valtonen & R. W Ho.ffinann ...... 122 New data on theepi demiol ogy of bird haemoprote ids on the Curonian spit G. LiutkeviCius ...... 123 Effects of different natural products against sarcoptic mange mites in swine E. Ma gi & A. Kaarma ...... 123 Studies on the female reproductive system of Diphyllobothrium latum L. G. Poddubnaya ...... 124 Influence of environmental factors on survival of trematode cercariae fi'om littoral prosobranchs of the White and the Barents Seas: an experimental study V V Prokoji€N...... 12 4 Platelet activation markers in parasit ic diseases D. Prokopowicz, J Matowicka-Karn a & A. Panas iuk...... 12 5 A PCR - based method for the detection of Tetrahymena corlissi contamination of Ichthy ophthirius multif iliis in in vitro culture D. Pugovkin, T Wahli & R. Felleis en ...... 126 Dependence of Teleos te i helminth fauna Jl'om Atlantic Antarctic waters on their fe eding peculiarities G. Rocljuk...... 12 7 The in fluence of magnetized water on the larvae of Culex pipiens 0. Rogachy ova ...... 127 Prelim in my data on nucleotide sequences of 18S rDNA of two species of cmstac ean paras ites fi·om Lake Baikal 0. Rus inek, K. Kuz nedelov & E. Rusinek...... 128 Comparison of the helminth faunas ofBu fo viridis Laurenti, 1768 and B. regularis Reuss, 1828 collected in Egypt D. Schneider & H Sattmann ...... 129 Some results of investigation of "b lack spot" disease of brea in(Abram is brama) fi'om the Curonian Lagoon (s outh-east partof the Baltic Sea) 0. Shuhgalter ...... 129 Why do br own rats Rattus norvegicus liv ing in the sewer system of the old town of Reykjavik have so fe w parasites? K. Skfrnisson & E. J6hanns d6ttir ...... 130 Host specificity of gainasid mites (Ga inasina: Spintumicidae, Macronyssidae) fi·om bats (Chiroptera) MK. Stany ukovich...... 131 xii Is the monogcncan Amyrocephalusparadoxus capable of regulating the population structure ofO+ class of St iz os tedion lucioperca? VK St arovoitov & P.J . Gerasev ...... I 3 I Twnour-like infections of cyprinid fish(Cyprinidae) fromCuronian Lagoon (the Baltic Sea) RV Sudarev & VK. Starovoitov ...... 132 Distribution of mosquitoes (Diptera: Culicidae) in different habitats of urban territory N Tereshkina & M Trukh an ...... I 33 Dynamics ofabundance ofHy sterothylacium aduncum (Nematoda), a parasite ofsprat in the Black Sea L. Tkachuk ...... I 3 3 The influence of an environmental factor (pollution) on abundance of the dominant parasite species of shrimp Palaemon elegans Rathke in the Black Sea L. Tkachuk ...... 13 4 Tendency fo r a polycyclic seasonal development of Proteocephalus cernuae in the Rybinsk reservoir A. V Tyutin ...... 135 Regularities in the distribution oflethal haemosporidian parasites ofpoultiy G. Valkiilnas & T Iez hova ...... I 35 Distribution ofParacoenogonimus ovatus Katsurada, 1914 in fishes of Russia V Voronin, R Kudenz ova, Y Str elko v, N Chernyshova, E. Golubeva, A. Zhokh ov, M Safronov & V Fetis ov...... I 36 Occurrence ofrnicrosporidia and Schistace phalus spp. in sticklebacks ofthe Gulf of Finland near St. Petersburg: results oflong-term observations VN Voronin & ME. Safronov ...... I 36 The platyhelminths ofinsectivores in Lithuania R. Zqsityte...... I 3 7 Anthropogenic pressure influences on Ix odes tick populations S.D. Zharkov, P.M Jens en, H V Dubinina & A.N Alekseev...... I 37 Bull Seand Soc Parasitol 2000; 10 (2): 1-5 INVITED LECTURES VECTOR (TICK) AS ECOLOGICAL NICHE OF DIFFERENT PARASITIC ORGANISMS A.N. Alekseev and H.V. Dubinina Zool og ical Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia E-mail: [email protected] The main purpose of our communica organisms. Different types of interac tion is to test Academician Eugeny Pav tions, however, serve as important links lovsky's (1934) postulate that any living in the chain of complicated parasitic organism represents some kind of "parasi systems, including vector-borne disease tocosmos". We humans are able to live systems. As we have stated previously because, under normal circumstances, the (Aiekseev, 1993), mixed infections of "microcosm" of our microflora is the basis vectors in natural fo ci of diseases is of our normal metabolism. Nevetiheless most probably the rule rather than the the same commensals or foreign flora exception. We agree with the basic become deadly dangerous when this bal statement of P.A. Petristcheva (1972), ance is disturbed. The appearance of that between diffe rent pathogens AIDS served as a trigger, stimulating the transmitted by the same vector there discovery of a legion of so-called oppor exists a very complicated interface, tunistic infections. These may consist of which determines the successful sur our own previously commensal micro vival, development and transmission of flora, or might be representatives of exotic one or another parasitic organism to the agents such as flea-borne microsporidia. vertebrate host. In the past, imagining ourselves to be There exists, however, an opposite the "kings of nature", we hardly permitted opinion: based on observations that the thought that the same processes are different pathogens are able to coexist common and typical of all other organ simultaneously in the same individual isms on the lower steps of the "ladder" of ixodid tick, some authors portray the life fo rms. In the study of parasitic sys tick as a kind of a «cabinet», containing tems most investigators have typically different «shelves» (i.e. ecological neglected the interaction between compo niches) on which different pathogens nents of the "parasitocosmos" inside host live and multiply practically independ- 2 ent of each other. This concept ignores the isms or parasites. It was shown previ type of pathogen, the time and stage at ously (Alekseev, 1996; Alekseev et al, which one or another pathogen infests the 1996) that when Ixodes persulcatus invertebrate host (vector) and ignores the ticks already naturally infected by host reaction to the invasion . According to Borrelia burgdorje ri sensu lata were our definition of biological vector speci superinfected with TBE virus, infection ficity (Alekseev, 1989), any pathogen that was much less successful than in previ is specific is able to multiply and enlarge ously uninfected ticks. its biomass to be transmitted to another In our study we are considering I host. Only at the end of these processes persulcatus ticks as an ecological niche might a vector be considered as a «filled for different groups of pathogens. cabinet». In nature, however, simultane While studying the population of taiga ous infections of a single vector is a very tick in the vicinity of the megapolis of rare phenomenon: symbionts (e.g. Wolba St. Petersburg, we discovered that this chia) and some pathogens (e.g. tick-borne population is not homogenous but encephalitis virus) may be transmitted represents two sub-populations. We transovarially, while others, patiicularly called the larger group "normal", and bacteria ( spirochetes, rickettsiae, piro the other one "anomalous" because of plasmidae) infect the tick chiefly via an anomalies in their exoskeleton struc infected blood meal; helminths, fo r exam ture, apparently a result of anthropo ple, are acquired exclusively with the genic pressure (Alekseev, 1996). The blood meal. The manner of penetration by responses of both sub-populations to these parasites and their interaction inside pathogens and to imp01iant abiotic the vector are very important factors in factors (Alekseev & Dubinina, 2000) their survival and multiplication. When were different. This permitted us to simultaneously infected by Leishmania analyse the I persulcatus population major and Leishmania gymnodactyli, not as one, but as two different eco Phlebotomus papatasii sand-flies died logical niches. Vector response to (Alekseev et al, 1975). Trematode super Borrelia and Ehrlichia species was invasion of snails, which were previously quite different, depending on the group naturally infected by another trematode, to which the host belonged. may kill the host. The trematode-infected Responses of uninfected and in host subjected to super-invasion either fected ticks to a temperature gradient fully or partially suppressed the develop (Alekseev & Dubinina, 2000), and to ment of the newcomer maritae. All such humidity and gravity were used as tested pairs of parasites belonged to dif markers of the two sub-populations' ferent families (Ataev, 2000). In sand fl ies characteristics. All these markers, as successfully infected by two species of well as measurements of tick activity Leishmania, the first-comer had an advan parameters using the authors' own tage in development in the host gut «ticks-drome» (Alekseev, 1996; Alek (Safjanova et al, 1976). Ixodes ticks are seev et al, 2000, in press) were used to not an exception, being host to many confirm the existence of two sub different groups of symbiotic and patho populations. The same methods were genic (for vertebrate hosts) microorgan- 3 used to reveal differences in the interac It was established that infected tion of the pathogens within the tick-host specimens appeared from the litter interfa ce. Negative hydro- and geo-taxes mainly when there was a positive tem determined the activity of ticks moving up perature gradient (temperature of sur vegetation stems. In the normal popula face minus temperature in the litter). tion, similar numbers of uninfected and The threshold gradient at which ticks infected ticks showed both taxes, whereas emerged from the litter is here called in the anomalous population, more in the TSG. Our earlier report of a differ fected than uninfected ticks showed nega ence between TSG values of uninfected tive taxes. There was a greater total num and infected ticks (Alekseev & Dubin ber of active ticks among anomalous ina, 2000) was based on this phenome infected ticks. This was especially clearly non. The decrease in the number of demonstrated in males, because the ana anomalous ticks under a negative TSG tomical anomalies were detected more was twice as large as that of normal precisely in this sex. Infection by any ones. Using the PCR method to detect pathogen decreased activity parameters infection, we compared the behaviour (speed, height and activity index) in both of mono-, dual- and triple-infected groups of ticks, but the relative decrease specimens, and showed that the reac in anomalous specimens was less than in tion to abiotic factors of such infected normal ones. normal and anomalous ticks was some- 3 2,5 U) CV 2 -:a n:s > 1,5 � U) 1- 1 0,5 0 .... - ..... - ...... - - ...... ui CIJ - :::: 0 CIJ ui .s e QJ CIJ .s a.; ..... J::; a.; la '5 c::: � � c::: '5 la � 0 rtl la 0 � Cl\ :::J . :::J Cl\ .Q � a a .� . ai ui ai .Q I:Q ai � � ai Type of infection Fig. 1. Temperature gradient (TSG) values, at which nonnal (A) and anomalous (B) infe cted ticks appeared from the litter. 4 times quite opposite. Borrelia garznu structure of tick sub-populations, which infected normal ticks showed a TSG of serve as an ecological niche for differ 0.82° C compared with 2.63° C for simi ent pathogens. larly infected anomalous ticks. The oppo site reaction was established among Bor relia afzelii infected specimens: B. afzelii Acknowledgements infected anomalous specimens had a TSG I wish to express my gratitude to Dr. of 1° C compared with 2.01° C for normal S. Randolph (Oxford Tick Research specimens (Fig. 1). Most of the ticks with Group, Department of Zoology, anomalies infected by two pathogens Oxford, UK) for her valuable advice simultaneously (whatever the type of and improvement of the English infection) had lower TSG values that did version of the manuscript. The research normal ticks. Only dual infection by a described in this publication was made mixture of B. ajzelii and B. garinii, both possible in pmi by Grant 98-04-49899 highly pathogenic for humans, increased from the Russian Basic Research the TSG of both types of ticks. Foundation and in pmi by Grant Activity indices were also dependent 9600864 Danish Research Council on the type of infection, with observed synergistic or antagonistic effects of dif References ferent pathogens eo-inhabiting the same Alekseev AN. Composition of invertebrate host. The more intensively infected were host "p arasit ocosmos" and its imp ortance I persulcatus ticks, the more often were for transmission of transmissive infecti on Borrelia found in their salivary glands agents. Parasitol Rev (Leningrad: Izdatel (Moskvitina et al, 1995). Furthermore, stvo «N auka») 1989; 36: 5-20 (in Russian) active multiplication of any pathogen Alekseev AN. Tick-borne path ogen syste m enhances its quality as a parasite and and its emergent qualities. St.Petersburg: increases its transmissiOn probability Zool ogical Institute RAS, Inc. 1993 (In (Mackinnon & Read, 1999). We found a Russian) higher prevalence of high infection inten Alekseev AN. 1996. Tick path ogen interac sity (Borrelia No per 250 darkfields) in tions: Behaviour of infected and uninfected anomalous ticks than in normal ones. ticks (Ixodidae). In Acarol ogy IX. R. Also, amongst ticks more intensively Mitchell, D.J. Horn, G.R. Needham, W.C. infected by Borrelia, anomalous speci Welbourn (eds.) Columbus, Ohio 1996; 1: mens were the most active ones. Dual 113-5 infection of B. aftelii and B. garinii en Alekseev AN, Buren kova LA, Vasilyeva IS, hanced activity of both groups of ticks, Dubinina HV, Chunikhin SP. The function but more so in anomalous ticks than in ing of mixed tick-b orne infection foci in normal ones. Borrelia-Ehrlichia eo Russia. Med. parazitol. (Moscow) 1996; 4: inhabitants had a synergistic effect, caus 9-16 (In Russian) ing a decrease in TSG and an increase in Alekseev AN, Dubinina HV. Abi otic pa infected tick activity, most marked rameters and die! and seasonal activity of amongst anomalous ticks. All these data Borrelia--infected and uninfected Ix odes stress the importance of monitoring not pers ulcatus (Acarina, Ixodidae). J Med only the type of infection, but also of the Ent omol 2000; 37: 9-15 5 Aleks eev AN, Jensen PM, Dubinina HV, Smirnova LA, Makr ouchina NA, Zharkov SD. Peculiarities of behavi our of taiga (Ixodes persulcatus) and sheep (Ixodes ricinus) ticks (Acarina, Ixodidae) dete nnined by diffe rent methods. Folia Parasitol 2000; 47: 00-00 (In Press) Alekseev AN, Safjanova VM, Karapetian AB. On the viability of sand flies (Diptera, Phy ch odidae, Phleb otominae) at the infection with pr omastigotes of the diffe rent species of Leishmania. Parazitologia 1975; 9: 271-7 (In Russian) Ataev GL. Development of the Tremat oda patthenites. Diss. Dr. Sci. St. Petersburg 2000 (In Russian) Mackitm on MJ, Read AF. Genetic relation ships between parasite virulence and transmis sion in the rodent malaria Plasmodium cha baudi. Ev oluti on 1999; 53: 689-703 Moskvitina GG, Korenberg EI, Spielman A, Schchyogoleva TV. On frequencies of general ised infecti on in unfed adult ticks of the genus Ix odes in Russian and American foci of the borrelioses. Parazitol ogia 1995; 29: 353-9 (In Russian) Safjanova VM, Alekseev AN, Karapetj an AB. The fate of promastigotes of Leishmania tropica major and L. gymnodactyli in the or ganism of Phlebotomus papatasi at the mixed infection. Parazitologia 1976; 10 : 78-83 (In Russian) Pavl ov sky EN. Host organism as a parasite habitat. Nature 1934; 1: 80-9 1 (In Russian) Petristcheva P A. 1972. Biozen otical bands of bloodsucking arthrop ods with vertebrates and disease agents. Main results and perspectives of development of the theory of the natural focality of the human diseases. Moscow 1972: 108-23 (In Russian) Bull ScandSac Parasito/ 2000; 10 (2): 6-11 SELECTIVE PRESSURES AND PARASITE STRATEGIES C. Combes Centre de Biologie et Ecologie Tropicale, Universite, 66860 Perpignan, France Phone: 33 4 68 66 20 50, Fax : 33 4 68 66 21 81, e-mail: [email protected] "[Host-parasite relationships] are extremely interesting also from the point of view of theoretical considerations" (V. A. Dogiel, 1947) Sewall Wright's adaptive surface Sewall Wright (1932) described the bolizes an individual. Natural selection relationships between gene combinations tends to gather genotypes towards sum and fitnessas a landscape with peaks and mits because individuals close to a sum valleys. The landscape was represented mit have a reproductive success better as a two-dimensional figure (Fig. 1 ). In than individuals situated on slopes or spite of Wright writing that this "dia valleys. Conversely, mutation tends to grammatic representation of the field of disperse individuals away from the peak. gene combinations in two di + ..· mensions instead of many thou _ ,',: sands" was a "very inadequate representation" of reality, Wright's proposition of adaptive peaks had an immense success: "Sometimes in science a simple phrase or diagram appears to be so cogent an encapsulation of an important truth that it takes on almost mythological propor tions" (Lewin, 1988). A population can be repre sented on Wright's landscape as Fig. 1. The original figure of an adaptive landscape as a more or less tightly clustered symbolized by S. Wright (1932). Dotted lines are lines cloud of points. Each point sym- of equal fitness. 7 Wright's diagram gives a picture of "how a) in the parasite genomes, the prob a species' fitness varies as a result of ability of meeting the host; modifications in its genetic make-up" b) in the host genomes, the probabil (Lewin, 1988). Wright used his repre ity of avoiding infective stages of the sentation to illustrate how a population parasites. can move fr om one peak to another (the This is what I called the "encounter "shifting balance" model). Wright links arms race". It implies principally behav evolution to adaptive peaks in the fo l ioural adaptations and counter lowing terms: "The problem of evolution adaptations. as I see it is that of a mechanism by which the species may continually find Then, at a second level (which corre its way from lower to higher peaks in sponds to "post-infective" events), natu such a fi eld. In order that this may occur, ral selection favours gene combinations there must be some trial and error which improve: mechanism on a grand scale by which c) in the host genomes, the probabil the species may explore the region sur ity of killing the parasite; rounding the small portion of the field d) in the parasite genomes, the prob which it occupies". ability of surviving in the hostile envi ronment created by the host. Encounter and compatibility filters This is what I called the "compatibil The notion of adaptive landscape can ity arms race". It implies principally be applied to parasite species without defence and evasion mechanisms. particular alterations. However, the fitness of free living Selective pressures in parasite-host species depends only on their own gene systems could thus be symbolized by combinations. On the contrary, the fit two filters, an encounter filter and a ness of parasite species depends selection of both on its genes and on the genes host's genes. A parasite must be to meet adapted not just to an environ- :>- - (j) ment, but to a living environment c ::r .£ z _ peak :0 A . --- -- _ ·� o_ E ... ,� ., 0 0 £ -� CA )t � ;:�:-�-�-� - ' CJ -�-:b::, (j) , ' '"- ...... ,, Cil - "(5 . 0 (/) Kl ea (/) (j) c valley (j) Ol 8 :: oc t) X 0 --+------��------X .c Fig. 3. A peak of parasite high fitness (the peak is supposed to have a round shape, to simplify). A is an individual host close to the summit, B is another individual, close to the bottom (sup posedly having genes conferring low compatibility), C is another individual in a valley (unsuit able for the parasite). At right, 3-D view of a peak. 9 tes eB, CB ( e and c mean encounter and with compatibility are likely to be the compatibility, respectively). Because A main targets of natural selection ("hide is closer to the summit than B, the prob or fight", Hochberg, 1998). Because of ability of having its fitness reduced by the high costs which are usually associ the parasite is greater for host A than for ated with behavioural changes in most host B. This means that there exists an free-living animals, it can be expected "inversed landscape" if we consider host that natural selection often invests in fitness: where there is a peak of parasite closing the compatibility filter ("fight") fitness, there is a valley of host fitness. rather than in closing the encounter filter ("hide"). This scheme allows one to discuss various evolutive aspects of parasite-host 4) The model describes how a para relationships. site species can move from one adaptive peak to another. Since adaptive peaks are 1) If hosts produce mutations which mobile in the landscape, it can happen disperse individuals away from the peak, that two host species have (at a cettain it is those individuals which are away moment of evolution) neighbouring co from the summit which have in average ordinates. If, at the same time, mutations the best fitness because they have a in parasites make certain individuals better chance to escape parasitism (and wandering around their usual peak (let thus transmit their genes). This means us remember Wrigth's expression "trial that selective pressures arising from and error mechanism"), they may exploit parasites result in host populations a new peak, i. e. a new host species. "moving the peaks", either by changing Transfer implies only sufficient overlap behaviour (moving along x in Fig. 3) or ping of both encounter and compatibility by improving immune defence (moving co-ordinates (Fig. 4). Parasites "explore" along y), or both. potential host species in the same way 2) Such changes in the adaptive land free-living species explore the surround scape which is offered to parasites mod ing region in Wright's landscape. Large ify the pressures exetied on them. It can host-spectra mean that several host thus be expected that counter-strategies species constitute a single adaptive peak will be selected in the parasite gene pool. for a parasite (one may imagine a more These continuous selections of measures complicated representation associating and counter-measures are nothing else several peaks of different altitudes if not than a Red Queen process (V an V alen, all the host species provide the same 1973). Here the Red Queen is symbol fitness to the parasite). ized by parasites provoking shifts of host 5) The model can be useful to under peaks and hosts provoking shifts of stand why speciation often occurs when parasite peaks. It is clear that where a parasite exploits several host species, there is a peak of parasite fitness,there is either after transfers or after host speci a valley of host fitness, and reciprocally. ation. Alloxenic speciation (Euzet & 3) It is possible to speculate under Combes, 1980) occurs when adaptive what conditions strategies associated peaks are no longer overlapping. The with encounter or strategies associated isolating mechanism can be associated 10 Discussion y Various criti cisms have been : transfer addressed to g Wright's adaptive ..0 landscape (see 1B 0... Provine, 1986), the E 0 main one being that () � � ·::·�<·.<··-6?': : ' £ x and y-axes are · � discrete entities "0 QJ �·>::������r:::·:::'// (gene combinations) Cii '6 and not continuos 0 (/) barrier to "2r�- -· (/) variables, in such a ro gene flow -- (/) way that the adaptive QJ c surface does not QJ 0> exist in mathematical 1ii : \ 0 terms. Wright de ..c ��2)�'lf '' fe nded himself by B saying that "an intel L------X ligible representation host genes associated with encounter depends on some enormous simplifica tion" and by insisting Fig. 4. Overlapping peaks allowing transfer of parasite P from host species A to host species B. After transfer, displacement of peaks on the symbolic may provoke isolation of parasite gene pools. It is supposed that character of the host B moved to position B' because of selection of genes associ adaptive landscape ated with encounter (arrow 1), or to position B" because of selec (see Lewis, 1988). tion of genes associated with compatibility (arrow 2). Any inter The representa mediate position is possible. Pressures other than those arising tion of host species from the parasite-host system can "move the peaks". as adaptive peaks fo r with the encounter filter or with the parasites is even more symbolic because compatibility filter. These processes it uses gene combinations of the host and have been described by Combes & seems to neglect gene combinations of Theron (2000) as "polymorphism of the parasite. However, as shown above, habitat encounter" and "polymorphism visualizing adaptive peaks in this way of habitat compatibility". Similar reason allows one to investigate several aspects ing consists of considering that the adap of the evolution of parasite-host systems. tive peak is defined at the microhabitat If one wish to represent all the gene scale instead of the host species scale combinations involved in encounter and and leads to the concept of synxenic compatibility of a parasite-host system, it speciation (Euzet & Combes, 1980) with is not fo rbidden to consider that x axis homologous isolating mechanisms. symbolizes all genes controlling encoun ter, whatever they belong to the parasite 11 or the host, and similarly for y axis and genes controlling compatibility. In this case, two adaptive surfaces should be designed on a single 3-axis system, one fo r the parasite, one fo r the host. Be cause parasite and host compete for the same resource, peaks should never coin cide. If, in a given system, peaks coin cide, this means that the system is mutu alistic, not parasitic. References Combes C, Theron A. Metazoan parasites and resource heterogeneity: constraints and benefits. Int J Parasitol 2000; 30: 299-304 Combes C. Interactions Durables. Ecologie et Evolution du Parasitisme. Masson, Paris, 1995. Combes C. Fitness of Parasites: pathology and selection. Int J Parasitol 1997; 27: 1-10 Euzet L, Combes C. Les problemes de l'espece chez les animaux parasites. Mem. Soc Zoo! France 1980; 40: 239-85 Lewin R. The uncertain perils of an invisible landscape. Science 1988; 240: 1405-06 Provine WP. Sewall Wright and Evolutionary Biology. Univ Chicago Press, Chicago, 1986 Van Valen L. A new evolutionary law. Evo lution Theory 1973; 1: 1-30 Wright S. The roles of mutation, inbreeding, crossbreeding, and selection in evolution. Proc Vlth Intern Congress Genetics 1932; 1: 356-66 Bull Seand Soc Parasitol 2000; 10 (2): 12 ECOLOGY AND GENETICS OF DAPHNIA-MICROPARASITE INTERRACTIONS: WHICH FACTORS DETERMINE PARASITE DISTRIBUTION? D. Ebert Zoologisches Institut, Univsitat Base!, Base!, Switzerland Phone: 41 61 267 3488; Fax: 41 61 267 3457, E-mail: [email protected] There is hardly any multicellular competition among parasites) and spatial organism that is not parasitized during and temporal structure of the host some stage of its life. But not every host (meta)-population. population harbours every potential I present the results of a number of parasite species and within a population studies which were designed to under not every host is parasitized with the stand which factors govern the pres same parasites. Most host populations ence/absence patterns as well as the coexist with only a small set of their diversity of parasites in experimental and potential parasites and most host indi natural populations of the planctonic viduals carry only a small subset of the crustacean Daphnia magna. I will focus parasites in a population. A number of on three examples. I. A study of pres models have been suggested to explain ence/absence patterns of parasites in this diversity. These models are based on rock pool Daphnia populations. II. Test epidemiological factors (e.g. density ing models using experimental epidemi dependent transmission, population size, ology. Ill. The impact of genetic factors number of host species), biological in determining parasite success. factors (host quality and host genotype, Bull Seand Sac Parasito/ 2000; 10 (2): 13-18 AN ECOLOGICAL APPROACH TO THE ANALYSIS OF TREMATODE LIFE-CYCLE EVOLUTION K.V. Galaktionov1 and A.A. Dobrovolskif 1Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia, Fax: +8 12 - 1140444, e-mail: [email protected] 2Faculty of Biological and Soil Sciences, St. Petersburg State University, St. Petersburg, Russia Trematode life-cycles are amazingly Trematodes are distributed from the complex and diverse. They are character Arctic to the Antarctic and their life ized by changes of generations (parthe cycles are completed in different ecosys nogenetic and hermaphroditic) and ani tems (fresh water, terrestrial, marine mal hosts (intermediates and final). Each littoral, pelagic, benthic, etc). What is generation must develop from an imma the reason of their obvious success? Are ture state to a mature fo rm. Each phase there any universal features of the life in a life-cycle must be capable of carry cycle structure promoting their success ing out its particular function. Free liv ful transmission in practically all eco ing larvae must be required to penetrate systems? Is it possible that they have a host, parasitic larvae must survive in evolved some specific adaptations which the body of an intermediate host, adults provide life-cycles suitable for specific should be capable of reproduction. To ecological systems? To answer these ensure that these tasks may be completed questions, we should consider initial successfully various adaptations (mor steps of trematode evolution. phological and behavioural) have been developed in the course of evolution. Formation of life-cycle These adaptations are specific and de It is generally accepted that early pend on the life-cycle phase and ecologi trematodes (protrematodes) evolved in cal conditions of the environment. coastal zones and marine prosobranches Therefore, it is impossible to discuss became their first (primary) host (Gi evolution of life-cycles without consid netzinskaja, 1968, English translation - erating the ecological conditions that 1988; Pearson, 1972; Cable, 1965, 1974; influence these life-cycles. This is the Gibson, 1987; Galaktionov, Dobrovol essence of the ecological approach skij, 1998). The first two-host life-cycle which we use in our studies. arose between the triassic and jurassic 14 periods and it was associated with the aquatic ecosystems. Such "trans wide distribution of Teleostei. These ecosystem transfer", no doubt, proved events could only have taken place in a the basis for biological spread and estab restricted area that was characterized by lishment by trematodes and promoted a high diversity and density of animals. their expansion into new ecosystems, The emergence of second intermedi sometimes very different from their ate hosts and, consequently, ofthree-host original site of origin. life-cycles, became a turning point in the conquest of new environments by trema Patterns of Strigeidida and Plagior todes. Of crucial importance is the fact chiida transmission that distribution of infection passed from It should be stressed that although the a small short-living larva (cercaria) with three-host scheme of life-cycles is pre I imited locomotion ability to a second dominant, it is not absolute. The evolu intermediate host, a much bigger animal tion of life-cycle phases and complete occupying a much larger living space. life-cycles are different in different These were new opportunities for the trematode groups. This is well illustrated transmiSSion of life-cycles, enabling by the most specialized trematodes their further development in depths of which are found in the higher orders of water inhabited by different organisms. Strigeidida and Plagiorchiida. In these It is obvious that the three-host systems organisms the three-host scheme is pre became the more universal form of dominant. However, some specific fe a trematode life-cycle and permitted tures of strigeidids life-cycles, such as spread of these parasites into ecosystems embryonation of eggs in water, free of diffe rent types. swimming miracidia, long and complex Lecithodendriidae sensu lata (a suc development of metacercariae, long cessful group of higher trematodes in maturation of adults, relatively slow egg cluded in the Plagiorchiida branch) is a production etc., practically prevent life good example of a three-host scheme. cycle completion in ecosystems with The aquatic larvae of invertebrates com frequent and sharp changes in ecological pleting their development on land were conditions (for example, marine littoral incorporated as second intermediate zone). Eggs and miracidia are especially hosts into the prolecithodendriid' s life vulnerable as their functions are carried cycles. As a result, three groups of in out in the environment. The strategy of sect-eating animals (evolutionary unre slow egg production does not provide a lated) took the place of their previous high density of eggs in the environment final hosts. The new hosts were am and this is also unfavourable. At the phibians (Anura), insectivorous birds same time these specific fe atures of and bats. In every group of hosts specific Strigeidida life-cycles (unfavourable species and genera of trematodes under littoral conditions) are very effec emerged, and sometimes divergence tive in fresh water ecosystems in which reached the level of the family. So, due Strigeidida evolved and prospered. to their second intermediate hosts, The evolution of Plagiorchiida life lecithodendriids left the boundaries of cycles took a different direction. Mirac- 15 idia only hatch from eggs in the intestine cercaria is subjected to unfavourable of a molluscan host. Parthenogenetic environmental conditions. Dicrocoelids, generations are characterized by ex which have relatively recently success tremely high productivity, formation of fully inhabited terrestrial habitats, have definitive organ systems takes place in still retain the structure of their life the metacercariae, and therefore adult cycle. They have solved the problem of maturation in the definitive host occurs cercariae distribution in a manner which quickly and adults rapidly produce large protects them from the environment. numbers of eggs. This direction of Dicrocoelium dendriticum cercariae, evolution has resulted in life-cycles that which still present tails, enter the mol are more universal than those of lusc mantle cavity and form so called strigeidids. Two of the most specializied "slug balls", or mucoid globules there. Plagiorchiida groups are the These are excreted into environment and microphallids and lecithodendriids. The eaten by ants (the second intermediate microphallids are able to inhabit hosts). ecosystems with very variable conditions Representatives of the genera Cor such as littoral and terrestrial rigia and Euritrema are more special environments and Lecithodendriidae izied in this respect. Not cercariae, but prosper in fresh water ecosystems in thick-walled daughter sporocysts con which Strigeidida life-cycles are also taining larvae with rudimentary tails, common. leave molluscs and pass into the envi Life-cycles in marine littoral and ronment. In all three examples the proc terrestrial ecosystems ess of infection of the second intermedi Transmission of trematodes in littoral ate hosts has changed. It has now be and terrestrial ecosystems became possi come passive as they are ingested by the ble only after some adaptations arose host. This is similar to the situation that permitted the trematodes to be more already described for miracidia. or less independent from the environ Another terrestrial trematode, ment. Of those, probably "passive" infec Brachylaimidae, adapted to the condi tion of molluscs is the most impmiant. tions of dry land independently from This arose in trematodes that completed Dicrocoeliidae. It chose a different ap their life-cycles in water and eventually proach. Isolation of the cercariae from became widespread. In these cases the the environment was achieved by the miracidia do not leave the eggs when fact that in most brachylaimids cercaria they are free in environment. Instead are not released into the environment at they hatch only after they have been all. The life-cycle has secondarily be ingested by the molluscan host. come two-host one in the genera Leu A situation with the second free cochloridium, Hasstilesia etc, as the living larva (cercaria) is more compli molluscs have combined the functions of cated. On one hand, it is very impotiant both firstand second intermediate hosts. for transmission and provides for pene The emergence of two-host life tration into the second intermediate host. cycles that has become the main trend in On the other hand a free-swimming the evolution of trematodes adapted to 16 ecosystems with extreme environmental environment. This results in a drastic conditions. Two-host life-cycles are decrease in density of aquatic animals, widely distributed within another trema and consequently in less probability of tode group which has successfully successful infection. Biological adapta adapted to the littoral zone of northern tions in these conditions are obviously seas. This group is the Microphallidae directed towards maximal longevity of which demonstrates a range of "transi invasive larvae in the environment. This tory" fo rms ranged from those with free has been achieved in a number of ways living cercariae to those in the "pyg including the passive suspension in maeus" group. Metacercaria morpho seawater. Hemiurids that have developed genesis takes place within daughter complex tails prove an excellent exam sporocysts in this group (Deblock, 1977; ple of this (Gibson, Bray, 1979; K0ie, Galaktionov, 1996). 1990). The life-cycle of microphallids in the It is our opinion that the increase in ''pygmaeus" group is undoubtedly highly spatial scale of ocean systems required a adaptive to ecosystems with extreme new strategy in a life-cycle transmission. conditions. These trematodes are clearly Dispersion abilities of cercariae and predominant in littoral molluscs on the second intermediate hosts (mainly inver coasts of the northern seas (Norwegian, tebrates) became insufficiently to ensure Barents and White Seas) (Galaktionov, location and infection of final hosts such Bustnes, 1999). A representative of this as pelagic carnivores fishes. The adop group, Microphallus pseudopygmaeus, is tion of an additional host (a third inter the only species of seabird trematodes mediate host) became necessary. Initially that completes its life-cycle in the high this host main functions were transport Arctic shore ecosystems of Franz Josef and protection of the parasite. On one Land (Galaktionov, 1996). In boreal hand, it guaranteed the prolongation of regions trematodes with three-host life juvenile parasite stages and, on the other, cycles are predominant in littoral ecosys it provided transport between the site of tems. Probably, under "milder" ecologi parasite dispersion and the location of cal conditions the possibility of metacer final hosts. This gave rise to fo ur-host cariae become widely distributed be cycles. They were originally facultative cause of the activity of the second inter and then became obligatory. mediate host outweighs the advantages Four host life-cycles are characteris of complete independence from the tic for at least some didimozoids. Young environment achieved by the two-host specimens of these trematodes (at the life-cycle. initial stages of adult morphogenesis) have been recorded in the intestines of a Life-cycles in pelagic ecosystems number of small fishes. This has led to Ability to exploit on the offshore the possible conclusion by Nikolaeva shelf and pelagic regions of oceans is (1965) and Pearson (1972) that juvenile impossible without specific adaptations Didimozoidae are "transported" from and changes in life-cycle structure. This crustaceans to large pelagic carnivores is determined by the enormously of such as tuna and sierra in the intestines 17 of small fishes. This mode of transport is host) appears to be much more probable a fe ature of all species in this taxon than completion of a life-cycle involving inhabiting pelagic waters. several host animals (even two of them). Although the authenticity of fo ur-host The purely quantitative losses associ life-cycles in Didimozoidae may still be ated with complex life-cycles are com in question, they are obligatory in some pensated by the acquisition of very spe Hemiuridae. In these cases planktonic cialized adaptations (morphological, crustaceans act as hosts fo r mesocer physiological, behavioural, etc.) devel cariae (an intermediate fo rm between oped by the life-cycle phases in the cercaria and metacercaria). The metacer course of the evolution. The already carial stage has been transferred to mentioned advantages of the incorpora fi shes, thereby permitting the use of tion of intermediate hosts, and the fact these large and more long ranged carni that intermediate hosts are usually in vores as final hosts. The introduction of cluded in the trophic chain of the final a mesocercaria provides fo r additional hosts is of special importance. Complex morphogenetic changes to take place in life-cycles provide for adaptations that their development. ensure successful transmission in all Four-host cycles have also arisen types of ecosystems. In terms of biologi independently in some freshwater trema cal progress, complex life-cycles have todes, such as Strigea and A/aria. Ini been favoured and they have provided tially this was probably associated with fo r the universal success of parasites. their invasion and establishment in large This is why complex life-cycles are so fr eshwater basins. Adoption of new frequently encountered among parasites hosts (which is common in trematodes) and have resulted in their widespread promoted incorporation of large carni distribution in the biosphere. vores into the life-cycles. Many of these hosts such as foxes, dogs and wolves, Acknowledgements were only peripherally connected with We wish to thank Dr. S.W.B. Irwin the fr esh water basins that were essential fo r correction of the English. This study fo r the evolution of these parasites. was supported by RFBR (grant N 98-04- 49706) and INTAS (grant N 97-1 0224). Concluding remarks The data presented has demon References strated that complex life-cycles have Cable RM . "Thereby hangs a tail". J Parasi enabled parasites to utilized new groups tol l965; 51: 2-12 of hosts and expand into ecosystems of Cable RM . Phylogeny and taxonomy of diffe rent types. The new opportunities trematodes with reference to marine species. gained, probably outbalanced the inevi In: Vernberg WB, ed. Synbiosis in the sea. table difficulties connected with the Columbia, 1974: 173-93 extension of host chains and transmis Deblock S. De l'abregement du cycle evolutif sion of infection. This has occurred in chez les u·ematodes digenes microphallides. spite of the fact that completion of a Libro homenaje a! Dr Eduardo Caballero y simple life-cycle (host -free-living larva - EMU RAA MATUKOGU 18 Caballero. Instituto de Biologia, publicatio nes especiales. Mexico, 1977; 4: 151-60 Galaktionov KV. Life-cycle and distribution of seabird helminths in Arctic and sub-Arctic regions. Bull Scand Soc Parasitol 1996; 6: 31-49 Galaktionov KV, Bustnes JO. Distribution patterns of marine bird digenean larvae in periwinkles along the southern Barents Sea coast. Dis Aquat Org 1999; 37: 221-30 Galaktionov KV, Dobrovolskiy AA. The origin and evolution of trematode life-cycles. St. Petersburg: Izdatelstvo Nauka, 1998 (In Russian) Gibson Dl. Questions in digenean systemat ics and evolution. Parasitology 1987; 95: 429-60 Gibson DI, Bray RA. The Hemiuroidea: terminology, systematics and evolution. Bull Brit Mus (Natur Hist) 1979; Zoo! ser 36: 35- 146 Ginetsinskaya TA. Trematodes, their life cycles, biology and evolution. New Delhi, 1988 K0ie M. On the morphology and life history of Hemiurus luehei Odhner, 1905 (Digenea: Hemiuridae). J Helminthol 1990; 64: 193- 202 Nikolaeva VM. On the developmental cycle of trematodes in the family Didymozoidae (Monticelli, 1888) Poche, 1907 Zoo! Szurn 1965; 44: 1317-27 (In Russian) Pearson JC. A phylogeny of life-cycle pat terns of the Digenea. Adv Parasitol 1972; 10: 153-89 Bu!!Scand Soc Parasito/ 2000; 10 (2): 19-37 PHYLOGENY, HISTORY AND BIODIVERSITY: UNDERSTANDING FAUNAL STRUCTURE AND BIOGEOGRAPHY IN THE MARINE REALM E.P. Hoberg1 and A. Adams2 1Biosystematics and National Parasite Collection Unit, United States Department of Agricul ture, Agricultural Research Service, BARC East No. 1180, 10300 Baltimore Avenue, Belts ville, Maryland, USA 20705. 2US Food and Drug Administration, 11630 West 20th Street, PO Box 15905, Lenexa, Kansas, USA 66285 "The past is the key to the present, where historical reconstruction involving parasites contributes to a predictive fr amework fo r discovering the interaction of biotic communities, environment and climate. " Hoberg (1997) Abstract Helminth faunas in pinnipeds and Relatively archaic and widespread dis seabirds have been assembled across tributions fo r host-parasite assemblages widely disparate temporal and spatial appear indicated by some campulid scales. Structure in these systems is digeneans, including species of Orthos revealed by the branching order of phy planchnus, various diphyllobothriid logenies fo r parasites and hosts that cestodes, some anisakine nematodes in allows us to examine the history of phocids and otariids and acanthocepha diversification for host-parasite assem lans in cetaceans and pinnipeds. In con blages and to identify the range of de trast, cestodes of the genus Anophryo terminants that have influenced ecologi cephalus, and some species of anisakine cal and biogeographic patterns. Defined nematodes appear to have common across a spatial continuum, faunas at regional histories fo r diversification in their broadest extent may have distribu pinnipeds across northern seas during tions that are bipolar and global in scale the past 3-5 million years. These histo or may be geographically delimited ries remain compatible with new calibra within regional zones. Geographic scale tions fo r the initial opening of Bering may be linked to the relative age fo r Strait and communication between the initial association of parasite and host North Pacific and North Atlantic prior to taxa, vagility of the assemblage, and the the Pliocene/Pleistocene boundary. duration fo r their history of coevolution. Generalities developed in the Arctic 20 Refugium Hypothesis to describe the McLennan, 1991, 1993; Hoberg, 1996, history fo r this regional fauna continue 1997; Brooks and Hoberg, 2000). to have substantial explanatory power Parasite fa unas in avian and mammal and the significance of climatological ian hosts in the marine realm have re determinants, and environmental pertur ceived minimal attention with respect to bation fo r fauna! diversification is em studies of cospeciation and historical phasized. Definitive hosts in these sys biogeography (Hoberg, 1997). Elucidat tems are postulated as the primary driv ing the patterns and history fo r biodiver ers of isolation and diversification fo r sity, and the determinants of continuity parasites. Phylogenetic studies among and structure fo r such complex assem tetrabothriid tapeworms, campulid dige blages requires detailed information fo r neans, and ascaridoid nematodes have phylogeny of both hosts and parasites. revealed complex histories involving Some components of the parasite faunas coevolution and extensive colonization in pinnipeds and seabirds have been that describes a tapestry to elucidate examined in detail based on phyloge general-level determinants for diversifi netic methods (e.g., Hoberg, 1986, 1992, cation in marine faunas. Many parasite 1995; Hoberg et al. , 1997a). Complete groups, however, remain poorly studied, assessment of these systems, however, and have yet to be evaluated based on had until recently been limited by the phylogenetic methods. Phylogenetic availability of well resolved phylogenies studies of parasites and hosts provide fo r particular mammalian or avian host critical context fo r understanding pat groups (e.g., fo r pinnipeds, Arnason et terns in biodiversity, fauna! structure and al. , 1995; Berta and Wyss, 1994; Berta both contemporary and historical and Sumich, 1999; fo r alcids, Friesen et biogeography. al. , 1996a). Helminth fa unas in pinnipeds and Phylogeny, history and biodiversity seabirds have been assembled across Biodiversity can be assessed across a widely disparate temporal and spatial number of fu nctional hierarchies in a scales. Fmiher, host and geographic continuum linking populations, species, ranges of parasites are historically con ecosystems and communities at local, strained by genealogical and ecological regional and global scales. Established associations (e.g., Brooks and McLen within a phylogenetic context, biodiver nan, 1993 ). Structure in these systems, sity results from an intricate interaction revealed by the branching order of phy of history, ecology and geography as the logenies fo r parasites and hosts, allows determinants of organismal evolution us to examine the history of diversifica · and distribution. A fo undation fo r under tion fo r host-parasite assemblages. Al standing patterns and processes in the ternative hypotheses for history involv structure of biotic associations at any ing varying degrees of coevolution or level emanates from phylogenetic and colonization are the cornerstone fo r historical reconstruction, and this is elucidating origin, temporal continuity particularly evident in studies of com and biogeography. Consequently, para plex host-parasite systems (Brooks and sites serve as models to articulate a 21 comprehensive understanding of the and duration of the host group; (5.) history and structure of the biosphere fa unas of low diversity that are depau (Hoberg, 1997; Brooks and Hoberg, perate in contrast to relictual; and (6.) 2000). associations of variable temporal dura Coevolution, encompassing both co tion and varying degrees of cospecia speciation and coadaptation, is consis tion/coadaptation depending on time tent with association by descent and a fr ame fo r colonization of host clade. In a protracted history fo r components of an context of coevolution or colonization, assemblage (e.g., Brooks and McLennan, the temporal duration of an assemblage 1991, 1993; Hoberg et al. , 1997a). A is elucidated by interpretation of data coevolutionary history is corroborated encompassing: (1.) host- parasite distri through examination and interpretation butions; (2.) historical biogeography of of host-parasite associatiOns which hosts; (3 .) regional history; and ( 4.) demonstrate: ( 1.) consistency and con physical geology (Brooks, 1985; Brooks gruence in host-parasite phylogenies or and McLennan, 1991; Hoberg, 1986, area relationships; (2.) a high degree of 1992, 199 5, 1997; Hoberg and Adams, cospeciation or coadaptation; (3.) recog 1992). nition of numerical or phylogenetic These alternatives set a hypothesis relicts; and ( 4.) often widespread geo driven fr amework and a series of predic graphic distributions that in the case of tions and expectations fo r identification marine systems may be global or bipolar of the range of complex determinants of in extent. Additionally, general congru genealogical and ecological diversity and ence in biogeographic patterns among fauna! structure. Represented is a power complex host-parasite assemblages are fu l tool to explore the history of marine characteristic of faunas that have been and other biotas. We apply this fo unda influenced by coincidental physical and tion and fo cus on the helminth fa unas biotic processes as determinants of dis characteristic in pinnipeds and within a tribution (Hoberg, 1986, 1992, 1997). In context defined by: (1.) phylogeny and such instances, geographic scale may be history fo r hosts (2.) phylogenetic data linked to the relative age for initial asso fo r parasites; and (3.) patterns of host ciation of parasite and host taxa, vagility association. Specifically we explore the of the assemblage, and the duration fo r overlying generalities and contrasting their history of coevolution. patterns fo r fauna! structure (e.g., "ar Components of a biota that have been chaic" and widespread, versus "recent" structured by colonization contrast with and regional), host distribution and coevolutionary systems and demonstrate: historical biogeography while articulat (1.) incongruent and inconsistent phy ing hypotheses fo r the driving mecha logenies for parasites and hosts; (2.) nisms that influence diversity in marine similarities in host trophic ecology; (3 .) systems. fa unas that are geographically or region ally delimited; ( 4.) parasite faunas in A brief history fo r pinniped hosts: which diversification is temporally cir Pinnipeds are a monophyletic group cumscribed in the context of the origin of marine mammals with origins in 22 Boreal-SubArctic seas of the North associatiOns with specific host groups. Pacific 25 million years ago (MYA) Geographic scale may be linked to age, (Berta and Wyss, 1994; Berta and Sum duration and vagility of the assemblage; ich, 1999). Among the contemporary patterns may also be influenced by colo fauna, 3 extant families are recognized nization events. Typically bipolar distri including Otariidae (Berta and Demere, butions have been demonstrated across 1986), Odobenidae (Berta and Wyss, all groups of helminths in pinnipeds 1994; Demere, 1994; Arnason et al. , including acanthocephalans such as 1995 ) and Phocidae (Ray, 1976; Repen Corynosoma (e.g., Golvan, 1959; Zdzi ning et al. , 1979; Arnason et al. , 1995; towiecki, 1986), anisakine nematodes Bininda-Emonds and Russell, 1996; such as Contracaecum and Pseudoter Stanley et al. , 1996; Berta and Sumich, ranova (Deliamure, 1955; Fagerholm 1999). The history fo r extant higher taxa and Gibson, 1987; Bullini et al. , 1994, extends to the middle Miocene, 14-16 1997; Paggi and Bullini, 1994; Arduino MY A, with centers of diversification fo r et al. , 1995), diphyllobothriid tapeworms otariids and odobenids in the N01ih including Diphyllobothrium spp. Pacific basin and fo r phocids in the (Deliamure, 1955; Iurakhno, 1991; Wo N01ih Atlantic. Phocids and otariids are jciechowska and Zdzitowiecki, 1995), secondarily fo und in the Southern Hemi and the campulid digeneans (Deliamure, sphere and a minimum of 3 independent 1955; Adams and Rausch, 1989). Al invasions into the Southern Ocean that though broad scale patterns areapparent, influenced the contemporary fauna are few taxa have been examined phyloge recognized since the Miocene/Pliocene netically, a requisite first step in devel transition, 5-6 MYA (e.g., Repenning et oping hypotheses fo r the history of this al. , 1979). fauna. Phylogeny and historical biogeogra phy fo r pinnipeds establishes the spatial History fo r some Diphyllobothriidae and temporal limits on the distribution of (Eucestoda): the phylogenetically disparate elements Diphyllobothriids are among the of the helminth parasite faunas in basal groups of tapeworms and their otariids, odobenids and phocids. Diffe r occurrence in mammalian hosts from ences in specific phylogenetic hypothe marine systems may be consistent ini ses fo r the pinnipeds do not substantially tially with a history of colonization and alter the fo llowing hypotheses articu secondary coevolution (e.g., Bray et al. , lated fo r the evolution of host-parasite 1999; Hoberg et al. , 2000). It has yet to associations, fa una! history, distribution be completely resolved whether or not and structure. those taxa typical in cetaceans and pin nipeds, respectively represent subclades "Archaic" associations- the global and within the diphyllobothriids, or if there bipolar patterns has been an extensive history fo r host Geographically widespread faunas switching (Iurakhno, 1991). encompassing global or antitropical Diphyllobothriids are broadly distrib distributions are indicative of early uted among northern and southern pin- 23 nipeds. Elements of the fauna in otariids, among the genera endemic to Antarctica monachines and phocines appear to be and other diphyllobothriids are also distinct (Markowski, 1952a, 1952b; unresolved. Interestingly, such species as Wojciechowska and Zdzitowiecki, D. pacificum may have extensive ranges 1995), suggesting extended and diver in both notihern and southern otariids gent evolutionary trajectories for these including Eumetopias, Otaria and Arcto lineages of tapeworms. Patterns of dis cephalus; alternatively a complex of tribution compatible with a protracted poorly diffe rentiated species could also coevolutionary history in pinnipeds, are be represented (R.L. Rausch, Pers. evidenced by such genera as Glandi Comm., 2000). A clear understanding of cephalus, Baylisia, and Baylisiella in the history of this diverse assemblage of Mirounga and the lobodontines. Moder hosts and parasites is hindered by the ate generic-level diversity fo r diphyllo absence of phylogenetic studies among bothriids is observed in the ,...----- Cal/orhinus Lobodontini and the dis- ,----- Arctocepha/us tinctive nature of the cir- r---- Za lophus Eumetopias cumantarctic fauna would Neophoca have been established since Phocarctos :::::5-6 MY A. Otaria r-----'"""-'"-W<"'-"'-'---- Odobenus * 2 4 56 In contrast, Diphyllo ,------Monachus Oan Leptonychotes * 7 bothrium has a bipolar Ommatophoca distribution in northern and Lobodon Hydrurga southern hemisphere Pho Mirounga cidae and Otariidae, proba ..----'"""-"'-'---- Erignathus* 56 ,------Cystophora bly having become estab Phoca fa scia fa * 5 lished independently Phoca groenlandica * 5 a Phoca hispida * 3 5 through separate invasions Halichoerus of the Southern Hemi Phoca largha sphere by Phocidae (5-6 Phoca vitulina vitulina Phoca vitulina concolor MYA), "Arctocephalinae" Phoca vitulina stejnegeri (5MYA) and Otariinae Phoca vitulina richardsi (3MYA) (Repenning et al. , Fig. 1. Phylogeny of the Pinnipedia showing distribution and 1979; Berta and Sumich, history for species of Orthosplanchnus. Relictual distribu 1999). Relationships tions (stars) are indicated for 0. oculatus (Oo), 0. rossicus among the species distrib (Or), 0. arcticus (Oa) and 0. fr aterculus (Of) in odobenids uted in the Southern Hemi and for 0. antarcticus (Oan) in Leptonychotes vveddelli. sphere and Antarctica are Colonization (asterisks) explains associations for 0. pyg (Op) and other species among phocines; distributions currently unknown and it is maeus in Enhydra lutra and Eschrichtius gibbosus are not shown. uncertain if the fauna is Numbers show the sequence of speciation among species of phylogenetically parti- Orthosplanchnus. Phylogeny for the pinnipeds is modified tioned among phocids, from Amason et al. (1995), Be1ia and Wyss (1994) and otariids and cetaceans; Be1ia and Sumich (1999); fmiher alternativehypotheses are putative relationships presented in Bininda-Emonds and Russell (1996) 24 the genera and species of the Diphyllo tion of toothed-whales from manne bothriidae. fishes (Fermindez et al. 1998a, 1998b). Further instances of host-switching by History fo r some Campulidae (D igenea): campulids from odontocetes to pinnipeds Digeneans of the family Campulidae are represented by Zalophotrema in sea are primarily parasites in odontocete lions (Fermindez et al. , 1998a) and Or cetaceans (Deliamure, 1955) and appear thosplanchnus in odobenids (Figs. 1 and to be a group that originated by coloniza- 2). HOST(S) unordered I!I======::JoCampula c=J Odontoceti Dl CI:J Odobenus rosmarus rJ111!1B!IIIIIlllll!llllll!llllllllll!llllll!lllllllll!llllllll!l0. albamarinus � Leptonychotes weddelli ....·. ·.·.· o 0. ocu!atus mJ Erignathus barbatus miJ Phoca hispida r;==::::::::z� 1111 Phoca fasciata IIIIHRmJ Phoca groenlandica 11!1 Eschrichtius gibbosus 1111 Delphinaplerus leucas - Enhydra lutra mmJ polymorphic � equivocal Lw GEOGRAPHY unordered r;::::====:::::======:JIDCampul a c=J Holarctic r;===:::z::==:::·=:..=.. =.. ::: ..·= ..=" ·::J· .· o 0. albamarinus E:J Arctic Atlantic la Bering Sea 0. oculatus - Antarctica pygmaeus E;3 equivocal 0. r;::::===�o 0. rossicus o 0. fra terculus Fig. 2. Phylogeny for species of Orthosplanchnus showing host associations and biogeographic history (see Appendix 1). Host and geographic distribution were mapped and optimized on this tree using MacClade 3.05 (Maddison and Maddison, 1992). Fig. 2a. Host associations indicate a basal relationship with odobenids subsequent to colonization from odontocetes. Orthos planchnus exhibits a relictual distribution in Odobenus rosmarus (Or) and Leptonychotes weddelli (Lw); other associations in phocines, Erignathus barbatus (Eb), Phoca hispida (Ph), P. fa sciata (Pf), and P. groenlandica (Pg), and other marine mammals, Enhydra lutra (El) and Eschrichtius gibbosus (Eg) are attributed to colonization. Fig. 2b. Biogeographic history for Orthosplanchnus is largely confined to northern seas, with evidence of vicariance, or a single invasion into the SouthernOcean represented by 0. antarcticus 25 Comparative studies based on the of hosts or parasites (e.g., Deliamure, phylogeny of Orthosplanchnus suggest 1955; Adams and Rausch, 1989; that species of this genus coevolved with Fernandez et al. , 1998a). It was gener odobenids and monachines. Distribu ally accepted that Orthosplanchnus spp. tions fo r contemporary species of Or were primarily associated with phocine thosplanchnus are relictual, indicated by seals from northern seas in the Pleisto basal associations fo r 4 species in Odo cene and that the occurrence in walruses benus rosmarus and the highly derived reflected recent colonization; the distri position fo r 0. antarcticus in Leptony bution fo r 0. antarcticus was thus seen chotes (Fig. 2a). Secondarily, sequential as enigmatic. Critical examination of colonization fr om odobenids to phocids relationships with pinniped hosts and and other marine mammals accounts fo r Orthosplanchnus (Figs. 1 and 2) indi contemporary distributions of 0. pyg cates associations with odobenids and maeus, 0. arcticus and 0. fraterculus. lobodontines that extend at a minimum Temporal limits and historical biogeog to near the Miocene-Pliocene transition, raphy fo r odobenines and monachines, > 5MYA. however, suggests an alternative (Re penning et al. , 1979; Berta and Sumich, Regional Holarctic faunas- the Berin 1999). Colonization of the latter group gian connection fr om walruses could be postulated dur Regionally or geographically delim ing a period of fauna! interchange in the ited faunas are often associated with Central American Seaway, adj acent to specific host-groups and definable tem the Pacific/Caribbean (5-8 MY A), prior poral limits fo r assemblages that are to establishment of the monachines and linked to a history of host-switching by diversification of the Lobodontini in the parasites. These explain a certain range Southern Hemisphere. In either situation, of patterns in biogeography and host all associations with phocids and other associations fo r helminths in northern marine mammals could represent coloni phocines and alcid seabirds. Synchronic zation, both temporally deep, consistent and congruent patterns fo r speciation with distribution in weddell seals, and and historical biogeography have been relatively recent to contemporary, con postulated fo r phylogenetically disparate sistent with distributions in sea otter, tetrabothriid and dilepidid tapeworms gray whales, and notihern phocines. and some anisakine nematodes in marine Relictual distributions revealed birds or pinnipeds across high latitudes through phylogenetic studies of cam of the Holarctic (Hoberg, 1986, 1992, pulids and Orthosplanchnus in pinnipeds 1995; Hoberg and A dams, 1992; Hoberg demonstrate that the history does not et al. , 1997a; Bullini et al. 1994, 1997; parallel Anophryocephalus and Alcatae Mattiucci et al. , 1998; Paggi et al. , nia in the Holarctic (Figs. 2a and 2b ). 2000). Prior assumptions about host associa tions and the history fo r Orthosplanch History fo r tapeworms in pinnipeds: nus were posed outside of the context Acquisition of Anophryocephalus by provided by phylogeny fo r either species pinnipeds from odontocetes (about 2.5-3 26 MYA) occurred subsequent to major antarctic and arctocephalines and otari segregation of northern and southern ines in the Southern Hemisphere ocean fa unas .::;5 MY A (Repenning et al. , (Hoberg and Adams, 1992). Coloniza 1979; Arnason et al. , 1995; Berta and tion of phocines by Anophryocephalus Sumich, 1999), accounting fo r absence occurred after cladogenesis of the phocid of these cestodes in true seals of the subfamilies and subsequent to the estab- lishment of monachines in the ..------Callorhinus Southern Hemisphere (Fig. 3). Arctocephalus ..------Basal species of Anophryo ..------Za lophus Eumetopias* 6 7 cephalus (A. anophrys, A. inuito Neophoca rum, and A. arcticensis ), except A. Phocarctos Otaria skrjabini, are associated with the high latitude seas of the Atlantic or ..------Monachus with the Arctic basin; all are spe Leptonychotes Ommatophoca cifically linked to a history of Lobodon cospeciation or coadaptation in Hydrurga ringed seals, Phoca hisp ida (Figs. 3 L---- Mirounga ..------Erignathus and 4). An association between ..------Cystophora Anophryocephalus, phocines and As Phoca fa sciata*3 P. hispida extends no deeper than .-----lA Phoca groen/andica * 4 �.!'-�lA�.Aar Phoca hispida * 1 2 3 4 2.5-3.0 MYA in the Atlantic and Halichoerus Arctic basin, consistent with the Phoca largha * 3 5 Phoca vitulina vitulina origin of this phocine (Arnason et Phoca vitulina concolor al. , 1995) and host and geographic Phoca vitu!ina stejnegeri distributions fo r these cestodes s Phoca vitulina richardsi * 3 (Hoberg and Adams, 1992, Hoberg, 1992, 1995). Patterns of distribu Fig. 3. Phylogeny of the Pinnipedia showing tion fo r 2 of these species, A. distribution and history for spp. Anophryocephalus skrjabini and A. arcticensis in other A basal association (star) is recognized between phocines are compatible largely Phoca hispida and 4 species, A. anophrys (Aa), A. inuitorum (Ai), A. sktjabini (As), and A. arcticen with a history of colonization. sis (Aar). Colonization (asterisks) is consistent Assuming that the basal species with distributions of species in other phocines are historically linked to the Arctic including A. nunivakensis (An) in which speciation basin, then 2 independent invasions is associated with the host-switch. Species in of the North Pacific are postulated Eumetopias, A. eumetopii (Ae) and A. ochotensis (Figs. 3 and 4); compare to Hoberg (Ao ), resulted from colonization of this otariid fromphocines and subsequent cospeciation. Num ( 1992, 1995) and Hoberg and bers show the sequence of speciation among spe Adams (1992). Anophryocephalus cies of Anophryocephalus. Phylogeny for the would have been widespread in the pinnipeds is modified fiom Arnason et al. ( 1995), region, entering the North Pacific Berta and Wyss (1994) and Berta and Sumich from the Arctic coincidental with 2 (1999); further altemative hypotheses are pre sequential openings of Bering sented in Bininda-Emonds and Russell (1996). Strait (concurrent with 2 interstadi- 27 als) in the Pleistocene. Although Bering intense periods of isolation leading to Strait was open at least by the Mio speciation fo r Anophryocephalus. Thus, cene/Piiocene transition (Marincovich the first invasion resulted in A. skrjabini and Gladenkov, 1999), it may not have and the second led to A. nunivakensis, A. become significant fo r phocid biogeog eumetopii and A. ochotensis (Fig. 4). The raphy until later in the Pliocene. Range geographic distribution of A. skrjabini in extension fo r hosts and parasites into the P. hisp ida includes the Arctic basin Bering Sea, fo llowed by emergence of adj acent to Bering Strait (Hoberg and Beringia and closure of Bering Strait Adams, 1992), and rather than a secon during glacial maxima resulted m dary dispersal from the North Pacific, m=:=:=:=:=:=:=:=:=:=:::Jo Trigonocotyle HOSTS anophrys unordered fP��=====:=ZJoA c::::::J Odontoceti oA inuitorum CB Phoca hispida r;==::==.:=::==.:::::==.:o � Phoca fasciata r;=':±t±:ti:bz::JEJAskr jabini f:aD Phoca vitu!ina richardsi r;z:::!:ib::::=:::JaA arcticensis lllllllllll!l Phoca largha rmJ Phoca groenlandica rJ1ili!li!iai!!111111ii!Anun ivakensis - Eumetopias jubatus llliiTl polymorphic 111A. ochotensis � equivocal 111A eumetopii IJllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllll Trigonocoty/e GEOORAPHY unordered r;::::=:=:=:=:=:=:=:::Jo A. anophrys c::::::J Arctic Atlantic * oA inuitorum 1111!11111 North Pacific * r;:::======::::J 1111: Cosmopolitan '1!11111111111111111111111111111111111111111111111!111111A. skrjabini � equivocal m====::::JoA arcticensis IJIIillllllllllllllllllll!illl!!IA nunivakensis A. ochotensis A. eumetopii Fig. 4. Phylogeny for Anoph1yocephalus spp. showing host associations for pinnipeds and historical biogeography across the Holarctic region. The phylogenetic tree for Anoph!yocepha lus is based on the hypothesis presented by Hoberg (1995). Host and geographic distribution were mapped and optimized on this tree using MacClade 3.05 (Maddison and Maddison, 1992). Fig. 4a. Host associations are compatible with origin of Anophtyocephalus by coloniza tion from odontocetes (see also Hoberg and Adams, 1992). Basal species are associated with a history of cospeciation in Phoca hispida (Ph). Occurrence of A. sk1jabini in P. fa sciata (Pf), P. largha (PI) and P. vitulina richardsi (Pvr) may be consistent with colonization. Species in Eumetopias jubatus (Ej ) developed by cospeciation fo llowing colonization. Fig. 4b. Bio geographic patterns are consistent with a history linked to the Arctic-Atlantic basin (asterisk), and 2 independent invasions of the North Pacific basin (stars) in the Pleistocene coinciding with sequential openings of Be ring Strait 28 this may represent a relictual distribution lineage leading to Eumetop ias from fo r this species in its typical host. A Ne ophoca + Phocarctos and Otaria, and limited distribution fo r A. arcticensis in thus fo llowed the invasion of otariines the eastern Canadian Arctic is consistent into the Southern Hemisphere about with speciation by peripheral isolation of 3MYA (Fig. 3) (see Repenning et al. , this widespread ancestral population of 1979; Berta and Demere, 1986; Berta parasites in ringed seals and colonization and Sumich, 1999). of P. groenlandica. Overall the patterns Biogeographic and historical studies of distribution and diversification fo r of Anophryocephalus in northern pin Anophryocephalus are consistent with nipeds and Alcataenia in Alcidae have models fo r rapid speciation (Hoberg, revealed a recent history fo r these as 1995). semblages and a pattern of diversifica It is apparent that A. skrjabini and A. tion driven by climatological events nunivakensis are linked to 2 successive since the Pliocene (Hoberg and Adams, marine transgressions and independent 1992; Hoberg 1986, 1992, 1995). These openings of Bering Strait during the unrelated faunas originated by host Pleistocene (Fig. 4a, and 4b) . Remarka switching and were largely structured by bly, these parasites are exquisite indica sequential colonization during periods of tors of environmental conditions and extreme climatic variation and habitat fa una! continuity in peripheral seas of pe1iurbation that characterized the past Beringia during the Quaternary and can 2.5-3.0 MY. reveal considerable insights about the biogeographic history fo r phocines in the History fo r some anisakines in pin region (Hoberg, 1995). Given the hy nipeds: pothesis for the timing of arrival of Diversification among certain ani ringed seals in the N01ih Pacific basin, sakines appears to have paralleled, at conceivably the dating for these subse least in pmi, the history fo r Alcataenia quent isolation events might be fmiher and Anophryocephalus across the narrowed. Along with other species of Holarctic (Paggi and Bullini, 1994; Anophryocephalus, these cestodes are Bullini et al. 1994, 1997; Hoberg, 1995). direct indicators of (1.) complex bio Diversification within Contracaecum geographic events; and (2.) cryptic histo osculatum in the boreal and arctic zones ries of isolation at the intraspecific level of the Atlantic occurred within the past fo r their phocine hosts (Hoberg, 1995). 3.0 MY; striking host associations in Anophryocephalus eumetopii and A. Erignathus, P. groenlandica, and Hali ochotensis developed in Eumetopias choerus are evident. Diffe rentiation jubatus fo llowing colonization in the within Pseudoterranova decipiens oc N01ih Pacific :S2 MYA (see Hoberg and curred primarily among phocines over Adams, 1992, Hoberg, 1992). The asso the past 3.5 MY in the Atlantic, Arctic, ciation of Anophryocephalus and Eume and N01ih Pacific (Bullini et al. , 1994). topias was established subsequent to the The presence of 4 cryptic species among divergence of the Otariinae and "Arcto the Pseudoterranova decipiens - com cephalinae", after divergence of the plex in phocines and Eumetopias may 29 indicate a history of colonization in the marine systems across high latitudes of North Pacific ::;; 2 MYA (Bullini et al. , the Holarctic spanning the late Tertiary 1994, 1997; Mattiucci et al. , 1998; Paggi and Quaternary (Hoberg, 1986, 1992, et al. , 2000) in parallel to Anophryo 1995, 1996, 1997; Hoberg and Adams, cephalus spp. in seals and Steller's Sea 1992). The development of the ARH Lion. Additionally, speciation within resulted from empirical and phylogenetic Phocascaris resulting in P. phocae in studies of two highly disparate assem harp and gray seals and P. cystophorae blages of parasites and homeothermic in hooded and gray seals is related to hosts. This interdisciplinary hypothesis isolation and divergence in the North has become a fo cal point fo r discussions Atlantic about .5MYA (Paggi and within parasitology (Brooks and McLen Bullini, 1994). nan, 1993 ), and conceptually has become In these instances, differentiation an integral cornerstone of historical among a complex of morpho and sibling biogeographic studies of seabirds in the species is postulated to have been driven Notih Pacific and Okhotsk Sea (Siegel by isolation in Pleistocene refugia, and Causey, 1991; Friesen et al. , 1996b; D. via segregation in specificpinniped hosts Siegel-Causey, Pers. Comm., 2000) and (Paggi et al. 1991; Nascetti et al. 1993; pinnipeds (Betia and Sumich, 1999). Orecchia et al. , 1993; Bullini et al. , The ARH is a synthesis integrating 1994, 1997; Paggi and Bullini, 1994). A phylogeny, ecology, and geological fu rther generality fo r anisakines in pin history in a macroevolutionary frame nipeds appears to be a history of peri work addressing the apparent congruent patric speciation with isolation and and synchronic histories fo r marine host speciation strictly linked to geographic parasite assemblages across the Holarc isolation of the definitive hosts, high tic. Demonstrated is the power of phy lighting the impotiance of small effec logenetic approaches in elucidating large tive population size on diversification of scale geographic and historical patterns anisakines within the northern fauna in the structure and continuity of biotas (Arduino et al. 1995). (Hoberg 1995, 1997): "Speciation of hosts and parasites Arctic Refugium Hypothesis: coincided with patiitioning of the Notih Radiation of hosts and parasites in Pacific basin (and high latitudes of the Subarctic to Arctic refugia during the Holarctic Region) into refugial zones late Pliocene into the Quaternary appears during glacial maxima coinciding with as a general conceptual model where periods of maximum eustatic reduction refugial effects, isolation and habitat in sea level. Range contraction, vicari fragmentation have been significant ance, isolation and speciation occurred determinants for fauna! diversity during stadials. Range expansion, from (Hoberg, 1986, 1992). The Arctic Refu refugial centers and colonization oc gium Hypothesis (ARH) was formally curred during interstadials. These alter presented and named in 1992, and repre nating sequences of climatic extremes sents a synoptic outline for speciation constituted the driving mechanism of processes, and historical biogeography in diversification rather than extinction 30 among disparate taxa in marine envi lighted. Recent hypotheses fo r an earlier ronments at high boreal latitudes of the opening of Bering Strait now suggest a Holarctic Region during the late Plio potentially longer period of marine cene and Pleistocene epochs ....." exchange, prior to inception of periodic (Hoberg, 1992, p. 614). and cyclical isolation events ( eustatic Additional salient points were out transgression/ regression cycles) that lined (see Hoberg, 1992; Hoberg and characterized the late Pliocene and Pleis Adams, 1992), and have since been tocene (Sher, 1999; Marincovich and expanded in detail (Hoberg, 1995). The Gladenkov, 1999). Although Bering central points of the hypothesis include: Strait was open possibly fo r the first (1.) general replicated patterns fo r speci time near the Miocene- Pliocene transi ation of phylogenetically disparate hosts tion 4.8-5.5 MYA, the impact of this and parasites- indicative of common connection between the Arctic and Pa causality; (2.) small effective popula cific basins on the biogeographic history tions of hosts and parasites; (3.) cyclical fo r pinnipeds has yet to be elucidated climatic variation with eustatic reduction (e.g., Repenning et al. , 1979; Berta and in sea level; ( 4.) habitat disruption, and Sumich, 1999). The chronology for some vicariance of host-parasite populations; parasite-host assemblages across Holarc (5.) range restriction in ephemeral and tic seas could have a longer temporal marginal refugia and micro-refugia duration, but such would have been during stadials leading to cospeciation dependent on the specific environmental and coadaptation; (6.) range expansion conditions and biotic provinces that during interstadials leading to coloniza characterized the Arctic basin in the late tion (host switching); and (7.) a specific Miocene. time frame during the late Pliocene and Pleistocene. The ARH serves as a model Host switching and diversification in or paradigm fo r speciation processes and marine systems historical biogeography across the A dominant and recurring theme or Holarctic since the Pliocene (Hoberg general trend in diversification of the 1995) and has received substantial cor helminth faunas in marine homeotherms roboration through studies of anisakine including cetaceans, pinnipeds and sea nematodes (e.g., Paggi et al. , 1991; birds has been colonization (Hoberg, Nascetti et al. , 1993; Orrechia et al. , 1986, 1996, 1997; Hoberg et al. , 1997a; 1994; Bullini et al. , 1994, 1997). As a Hoberg et al. , 1999a; Brooks and valid generality, the ARH has been rein Hoberg, 2000). Few taxa of helminths fo rced by recent studies on phylogeny are indicators of historical coevolution and biogeography of marine birds in ary linkages, or association by descent, cluding alcids (Friesen et al. , 1996a,b ), between marine and terrestrial faunas and shags (e.g., Siegel-Causey, 1991). (Deliamure, 1955). The concept of "hos The critical significance of the Berin tal radiation" (e.g., Iurakhno, 1991) has gian nexus as a physical determinant of defined ecologically linked switches marine biogeography during the Pliocene among phylogenetically unrelated hosts; and Pleistocene continues to be high- e.g., among marine mammals, and 31 among marine mammals and avian hosts. among phocines, whales and sea otters. Instances of colonization of poikilo Thus, the hist01y of a group may involve therms from a homeotherm-source, an extended association with its hosts, however, have not been identified. but also may be structured by both eo Host-switching has occurred on vary evolution and ecological associations ing temporal scales extending from the that drive colonization. Mesozoic to Quaternary and Recent. For Hoberg and Adams (1992) discussed example, the Tetrabothriidae is an ap the issues surrounding host switching in parently archaic group, with origins in faunas typical of marine homeotherms. the Mesozoic. Ancestral hosts may be Colonization is a stochastic process represented by now extinct non-avian dependent on predictable guild associa archosaurs in marine systems, with tions over extended evolutionary time colonization of basal avian and later frames and may result in fauna! diversi mammalian hosts (Spasskii, 1993; fication within new host-groups, coadap Hoberg et al. , 1997b; Hoberg et al. tation, or parasite extinction. Broad 1999a, 1999b); such an hypothesis con diversification of a parasite group via trasts with previous concepts for the historical colonization indicates substan origin of tetrabothriids via colonization tial ecological similarity among defini of homeotherms by tetraphyllidean tive hosts of an assemblage. The dura cestodes of chondrichthians. Early colo tion, predictability and geographic extent nization of avian groups and diversifica of an ecological association within the tion of Te trabothrius may be indicated context of specific food-webs or trophic by patterns of putative basal cospecia linkages defines the potential fo r suc tion with marine bird assemblages (e.g., cessful colonization. It is also apparent Hoberg, 1996). Marine mammals were that not all associations, even those acquired independently as hosts fo r where host-specificity is particularly tetrabothriids in the middle to late Terti pronounced, involve deep temporal ary with colonization of cetaceans from roots. Examples of such assemblages seabirds and pinnipeds from odontocetes that have largely been structured by (Hoberg and Adams, 1992). colonization in already long established Campulids are also a relatively old vertebrate taxa include campulids and group with origins extending at a mini Anophryocephalus in pinnipeds and mum to the middle Miocene (� 10-15 Alcataenia in the Alcidae (Hoberg 1992; MY A), fo llowing radiation of odonto Hoberg et al. , 1997a). cetes including delphinids and phocoe nids (Fordyce and Barnes, 1994). Radia Definitive hosts as drivers of isolation tion of campulids occurred subsequent to and diversification host switching from marine fishes to An additional generality has been the cetaceans (see Fermindez et al. 1998a, observation of the impotiant role for 1998b) and secondarily among odonto definitive hosts as drivers in parasite cetes and pinnipeds. Diversification of speciation. In marine systems, parasite Orthosplanchnus involved both coevolu populations appear to be strongly influ tion among odobenids and colonization enced by the distributions of relatively 32 vagile definitive hosts that disseminate parasites could have proceeded inde infe ctive stages fo r an array of interme pendently from that of populations of diate hosts. Continuity fo r marine para intermediate hosts. site assemblages is reinforced though dependence on predictable foraging Phylogenetic context - building and zones by pinnipeds or seabirds, such as interpreting the tapestry systems of oceanic fronts or insular Phylogenetic studies of parasites and eddies, that represent fo ci fo r transmis hosts provide critical context for under sion. In insular and pelagic marine sys standing patterns in biodiversity, faunal tems, the putative role of intermediate structure and both contemporary and hosts is neutral with respect to both gene historical biogeography. flow and isolation (see Hoberg, 1992, The hierarchical order of the parasite 1995). Life history and transmission phylogenies reveals historical relation consequently are evolutionarily conser ships and constrains the range of possi vative and structured by guild dynamics ble explanations fo r faunal structure such that selection may be fo r continuity when examined within the context of in transmission rather than fo r associa host phylogeny. For example, in the case tions with a specific group of definitive of the relictual nature hosts. Orthosplanchnus of the assemblage is indicated by basal These predictions are consistent with associations of 4 species in Odobenus, low levels of cospeciation, where rapid but a derived placement fo r 0. antarcti shifts by parasites define the potential cus in Leptonychotes. In this assemblage fo r successful colonization that occurs the branching order of the parasite phy among ecologically similar hosts across logeny is largely congruent with its and within host-clades (Hoberg, 1986). hosts; relictual associations and parasite The common fa ctor in these associations extinction do not result in incongruence. is exploitation of a common prey re Overall patterns are consistent with basal source in communities occupying limited cospeciation and a now relictual distri biogeographic regions or zones. Faunas bution fo r species in Odobenus and structured by colonization are regionally Leptonychotes, but one of secondary limited and often depauperate (Hoberg, colonization among other phocines and 1986, 1992). marine mammals; the associations are Speciation is allopatric and is driven archaic, and antitropical in scale. by isolation of definitive hosts, whereas In contrast, the phylogenies of Ano ecological continuity and predictability phryocephalus and pinnipeds are highly are determined by tran smission dynam incongruent, consistent with an overall ics linked to intermediate hosts (Hoberg, history of colonization fo llowing initial 1995). Speciation at least fo r cestodes in diversification of species in ringed seals. pinnipeds and seabirds appears to be In these systems, diversification was driven by the limiting factor of the geo relatively recent and driven by well graphic ranges occupied by definitive defined patterns of environmental per hosts. Thus, isolation and speciation turbation in the Pleistocene; associations among diverse assemblages of marine are temporally circumscribed with re------ 33 spect to the origin and duration of pho ments of historical ecology and biogeog cids, and regionally delimited. raphy. Parasite and host phylogenies Deliamure (1955) observed that constitute the tapestry fo r elucidating the resolution of the history and biodiversity interaction of coevolution, colonization of helminth faunas in marine mammals and extinction on patterns of biodiver was dependent on accurate taxonomic sity, fa una! structure and ecological concepts, and an understanding of phy continuity across deep temporal and logenetic relationships for both parasite broad geographic scales in marine envi and host taxa. Nearly 50 years later, we ronments (Brooks and Hoberg, 2000). continue to have a paucity of phyloge netic studies for parasites in marine Acknowledgements homeotherms (Hoberg, 1997), and only We thank Kirill Galaktionov, Oleg recently has there been substantial reso Pugachev, and organizers of the sympo lution fo r relationships among the pri sium, "Ecological Parasitology on the mary host groups (Berta and Sumich, Turn of the Millennium" held 1-7 July 1999). 2000, at the Zoological Institute, St. Caveats aside, the power of integra Petersburg, Russia fo r their kind invita tion for phylogenetic, biogeographic, tion and the opportunity fo r us to discuss ecological, and geological history that our ideas about marine parasites. Devel leads to fo rmulation of synoptic hy opment of the concepts about cospecia potheses fo r fauna! development over tion and historical biogeography ad varying spatial and temporal scales is dressed in our paper benefitted from evident (e.g., Hoberg, 1997; Brooks and extensive discussion with Greg Klassen. Hoberg, 2000). Parasites are exquisite Additionally, Roberi L. Rausch gener phylogenetic and historical ecological ously shared his knowledge about the indicators revealing substantial informa systematics and biogeography of the tion about the marine biosphere (Hoberg, Diphyllobothriidae. 1996, 1997; Brooks and Hoberg, 2000). Phylogenetic hypotheses for parasites References are the fo undation for estimation of the Adams A, Rausch RL. A revrswn of the minimum age and duration of complex genus Orthosplanchnus Odhner, 1905 with ecological assemblages. In marine sys consideration of the genera Odhneriella tems, anisakine nematodes, cestodes, Skriabin, 1915 and Had wenius Price, 1932 digeneans and acanthocephalans, all with (Digenea: Campulidae). Can J Zool 1989; 67: complex life cycles, track broadly and 1268-78 predictably across trophic levels as Arduino P, Nascetti G, Cianchi R, P!Otz J, representatives of biological structure. Mattiucci S, D' Amelio S, Paggi L, Orecchia Patterns of association represent a con P, Bullini L. Isozyme variation and taxo tinuum from relatively recent to archaic. nomic rank of Contracaecum radiatum (v. Some associations are indicative of Linstow, 1907) from the Antarctic Ocean temporally deep structure of ecosystems (Nematoda, Ascaridoidea). Syst Parasitol and communities and serve to emphasize 1995; 30: 1-9 the imporiance of helminths in assess- 34 Arnason U, Bodin K, Gullberg A, Ledje C, Bullini L, Arduino P, Cianchi R, Nascetti G, Mouchaty S. A molecular view of pinniped D'Amelio S, Mattiucci S, Paggi L, Orecchia relationships with particular emphasis on the P. Genetic and ecological studies on nema true seals. J Molecular Evol 1995; 40: 78-85 tode endoparasites of the genera Contra caecum and Pseudoterranova in the Antarc Berta A, Demere TA. n. Callorhinus gilmorei tic and Arctic-Boreal regions. In: Proceed sp. (Carnivora: Otariidae) from the San ings 2nd Meeting, Antarctic Biology. Mar Diego Formation (Blancan) and its implica tello A, (ed). Padova, Italy: Universitare tions for otariid phylogeny. Trans San Diego Patavine, 1994; 131-46 Soc Nat Hist 1986; 111-26 Bullini L, Arduino P, Cianchi R, Nascetti G, Berta A, Sumich JL. Marine mammals: D'Amelio S, Mattiucci S, Paggi L, Orecchia Evolutionary biology. San Francisco: Aca P., Pltitz, J., Berland, B., Smith, J. W., Brat demic Press, 1999 tey, J. Genetic and ecological research on Berta A, Wyss AR. Pinniped phylogeny. In: anisakid endoparasites of fish and marine Berta A, and Demere TA, eds. Contributions mammals in the Antarctic and Arctic-Boreal in marine mammal paleontology honoring regions. In: Antarctic communities: species, Frank C. Whitmore Jr. Proc San Diego Soc structure and survival. Battaglia B, Valencia Nat Hist 1994; 29: 33-56 J, Walton DWH, eds. Cambridge, UK: Cam bridge University Press, 1997; 39-44 Bininda-Emonds ORP, Russell AP. A mor phological perspective on the phylogenetic Demere T. The family Odobenidae: A phy relationships of the extant phocid seals logenetic analysis of fossil and living taxa. (Mammalia: Carnivora: Phocidae). Bonner In: Berta A, and Demere TA, eds. Contribu Zoo! Monogr 1996; 1-256 tions in marine mammal paleontology honor ing Frank C. Whitmore Jr. Proc San Diego Bray R, Jones A, Hoberg EP. Observations Soc Nat Hist 1994; 29: 99-123 on the phylogeny of the cestode order Pseu dophyllidea Carus, 1863. Syst Parasitol 1999; Deliamure SL. Helminthofauna of marine 42: 13-20 mammals (ecology and phylogeny). Moskva: Izdatel'stvo Akad Nauk SSSR, 1955. (Eng Brooks DR. Historical ecology: A new ap lish Translation, Israel Program fo r Scientific proach to studying the evolution of ecologi Translations, Jerusalem, 1968) cal associations. Ann Missouri Bot Garden 1985; 72: 660-80. Fagerholm H, Gibson DI. A redescription of the pinniped parasite Brooks DR, Hoberg EP. Triage for the bio Contracaecum og (Nematoda: Ascaridoidea), with an sphere: the need and rationale for taxonomic morhini assessment of its antiboreal circumpolar inventories and phylogenetic studies of distribution. Zoo! Scripta 1987; !6: 19-24 parasites. Comparative Parasitology 2000; 67: 1-25. FernandezM, Aznar FJ, Latorre A, Raga JA. Molecular phylogeny of the families Cam Brooks DR, Mclennan DA. Phylogeny, pulidae and Nasitrematidae (Trematoda) ecology and behavior, a research program in based on mtDNA sequence comparison. Int J comparative biology. Chicago, USA: Univer Parasitol 1998a; 28: 767-75 sity of Chicago Press, 1991 Fernandez M, Littlewood DTJ, Laton-e A, Brooks DR, McLennan DA. Parascript Raga JA, Rollinson D. Phylogenetic relation parasites and the language of evolution. ships of the family Campulidae (Trematoda) Washington DC: Smithsonian Institution based on 18S rRNA sequences. Parasitology Press, 1993 1998b; 117: 383-91 35 Fordyce RE, Barnes LG. The evolutionary Hoberg EP, Adams AM. Phylogeny, histori history of whales and dolphins. Ann Rev cal biogeography, and ecology of Anophryo Earth Planet Sci 1994; 22: 419-55 cephalus spp. (Eucestoda: Tetrabothriidae) among pinnipeds of the Holarctic during the Friesen VL, Baker AJ, Piatt J. Phylogenetic late Tertiary and Pleistocene. Can J Zoo! relationships with the Alcidae (Charadriifor 1992; 70: 703-19 mes: Aves) inferred from total molecular evidence. Molec Bioi Evol 1996a; 13: 359- Hoberg EP, Brooks DR, Siegel-Causey D. 67 Host-parasite cospeciation: History, princi ples and prospects. In: Clayton D, and Moore Friesen VL, Montevecchi WA, Baker AJ, J, eds. Host-parasite evolution: general Barrett RT, Davidson WS. Population differ principles and avian models. Oxford, United entiation and evolution in the common guil Kingdom: Oxford University Press, 1997a; lemot Ur ia aalge. Molec Ecol 1996b; 5: 793- 212-35 805 Hoberg EP, Gardner SL, Campbell RA. Golvan YJ. Acanthocephales du genre Systematics of the Eucestoda: Advances Liihe 1904 parasites de mam Corynosoma toward a new phylogenetic paradigm, and miferes d' Alaska et de Midway. Ann Parasit observations on the early diversification of hum comp 1959; 34: 287-321 tapeworms and vertebrates. Syst Parasitol Hoberg EP. Evolution and historical bio 1999a; 42: 1-12 geography of a parasite-host assemblage: Hoberg EP, Jones A, Bray R. Phylogenetic spp. (Cyclophyllidea: Dilepidi Alcataenia analysis among the families of the Cyclophyl dae) in Alcidae (Charadriiformes). Can J lidea (Eucestoda) based on comparative Zoo! 1986; 64: 2576-89 morphology, with new hypotheses for eo Hoberg EP. Congruent and synchronic pat evolution in vertebrates. Syst Parasitol terns in biogeography and speciation among 1999b; 42: 51-73 seabirds, pinnipeds and cestodes. J Parasitol Hoberg EP Mariaux J, Justine J-L, Brooks 1992; 78: 601-15 DR, Weekes P. Phylogeny of the orders of Hoberg EP. Historical biogeography and the Eucestoda (Cercomeromorphae) based on modes of speciation across high latitude seas comparative morphology: Historical perspec of the Holarctic: Concepts for host-parasite tives and a new working hypothesis. J Parasi coevolution among Phocini (Phocidae) and tol 1997b; 83: 1128-47 Tetrabothriidae (Eucestoda). Can J Zoo! Hoberg EP, Mariaux J, Brooks DR. Phylog 1995; 73: 45-57 eny among the orders of the Eucestoda (Cer Hoberg EP. Fauna! diversity among avian comeromorphae) : Integrating morphology, parasite assemblages: the interaction of molecules and total evidence. In: Interrela history, ecology, and biogeography in marine tionships of the Platyhelminthes. Littlewood systems. Bull Scand Soc Parasitol, 1996; 65- DTJ, Bray R., eds. London, UK: Taylor and 89 Francis, 2000; In Press. Hoberg EP. Phylogeny and historical recon Iurakhno MV. Osobennosti evoliutsii struction: Host-parasite systems as keystones gel'mintov lastonogikh. In: Evoliutsiya in biogeography and ecology. In: Reaka Parazitov. Materialy Pervogo Vsesoyuznogo Kudla ML, Wilson DE, and Wilson EO, eds. Sympoziuma. Tol'yatti, Russia: Institut Biodiversity II: Understanding and protecting Ekologii Volzhskogo Basseina AN SSR our biological resources. Washington DC: 1991; 124-27 Joseph Henry Press, 1997; 243-61 36 Maddison WP, Maddison DR. MacClade: Paggi L, Nascetti G, Cianchi R, Orecchia P, Analysis of phylogeny and character evolu Mattiucci S, D'Amelio S, Berland B, Brattey tion. Version 3.05. Sunderland, Mass., USA: J, Smith JW, Bullini L. Genetic evidence for Sinauer Associates, 1992 three species within Pseudoterranova de czpwns (Nematoda: Ascaridida, Ascari Marincovich L, Jr, Gladenkov A Yu. Evi doidea) in the N01ih Atlantic and Norwegian dence for an early opening of the Bering and Barents Seas. Int J Parasitol 1991; 21: Strait. Nature 1999; 397: 149-5 1 195-212 Markowski S. The cestodes of seals fromthe Ray CE. Geography ofphocid evolution. Syst Antarctic. Bull British Mus (Nat Hist) Zoo! Zool 1976; 25: 391-406 1952a; 1: 123-50. Repenning CA, Ray CE, Grigorescu D. Markowski S. The cestodes of pinnipeds in Pinniped biogeography. In: Gray J, and the Arctic and other regions. J Helminthol Boucot AJ, eds. Historical biogeography, 1952b; 26: 171-214 plate tectonics, and the changing environ Mattiucci S, Paggi L, Nascetti G, Ishikura H, ment. Corvallis, USA: Oregon State Univer Kikuchi K, Sato N, Cianchi R, Bullini L. sity Press 1979; 357-69 Allozyme and morphological identification of Sher A. Traffic lights at the Beringian cross and Anisakis, Contracaecum Pseudoter roads. Nature 1999; 397: 103-4 ranova from Japanese waters (Nematoda, Ascaridoidea). Syst Parasitol 1998; 40: 81-92 Siegel-Causey D. Systematics and biogeog raphy of Notih Pacific shags, with a descrip Nascetti G, Cianchi R, Mattiucci S, tion of a new species. Occ Papers Mus Nat D'Amelio S, Orecchia P, Paggi L, Brattey J, Hist Univ Kansas 1991; 140: 1-17 Berland B, Smith JW, Bullini L. Three sibling species with Contracaecum oscula Spasskii AA. On the participation of marine turn (Nematoda, Ascaridida, Ascaridoidea), Mesozoic reptiles in the evolution of the from the Atlantic Arctic-Boreal region: suborder Tetrabothriata (Cestoda: Tetraphyl reproductive isolation and host preferences. lidea). Tezisy Dokladov. XI Conference of Int J Parasitol 1993; 23: 105-20 the Ukrainian Society of Parasitologists. Kiev, Ukraine, 1993; p. 154-155. (In Rus Orecchia P, Mattiucci S, D'Amelio S, Paggi sian. English translation, V. Tkach) L, Plotz J, Cianchi R, Nascetti, G, Arduino P, Bullini L. Two new members in the Contra Stanley H, Casey S, Carnahan JM, Goodman caecum osculatum complex (Nematoda, S, Harwood J, Wayne RK. Worldwide pat Ascaridoidea) from the Antarctic. Int J Para terns of mitochondrial DNA differentiation in sitol 1994; 24: 367-77 the harbor seal (Phoca vitulina). Mol Bioi Evol 1996; 13: 368-82 Paggi L, Bullini L. Molecular taxonomy in anisakines. Bull Scand Soc Parasitol 1994; 4: Swofford D. Phylogenetic analysis using 25-39. parsimony (PAUP). Version 3.1.1. Cham paign, USA: Illinois Natural History Survey, Paggi L, Mattiucci S, Gibson DI, Berland B, 1993 Nascetti G, Cianchi R, Bullini L. Pseudoter ranova decipiens species A and B (Nema Wojciechowska A, Zdzitowiecki K. Cestodes toda, Ascaridoidea): nomenclatural designa of Antarctic seals. Acta Parasitol Polon 1995; tion, morphological diagnostic characters and 40: 125-3 1 genetic markers. Syst Parasitol 2000; 45: Zdzitowiecki K. Acanthocephala of the 185-197 Antarctic. Pol Polar Res 1986; 7: 79-117 37 Appendix I. Phylogenetic analysis for 11. Egg. Length: 0= mean �100J.!m; 1= Orthosplanclmus spp. mean � 80 J.!m. Coding for this character was based on functional outgroup crite Phylogenetic analysis for Orthosplanchnus ria. was conducted with PAUP 3.0 using an 12. Vitellaria, Extent: 0= to near pharynx; exhaustive search (Swofford, 1993). Charac 1 = to acetabulum; 2= to near anterior ter-coding was based primarily on taxonomic edge of ovary. outgroup comparisons to Campula spp., 13. Vitellaria, Form: 0= dendritic in ante shown to be among the basal members of the rior; 1 = fo llicular. family Campulidae (Fermindez et al. , 1998a; 14. Cirrus, Spination: 0= absent; 1 = present. also Hoberg and Adams, unpublished data ). 15.Metraterm, Spination: 0= absent; 1= Additionally, a functional outgroup, the basal present. species of Orthosplanchnus , was used to 16. Habitat: 0= intestine; 1 = gall bladder. secondarily assess characters 11 and 20. A 17. Eye-Spots: 0= absent; 1 = present. single most parsimonious tree (MPT) result 18. Ovary, Position: 0= not overlapping ing from this analysis supported monophyly anterior testis; 1= overlapping anterior for the genus and was fully resolved (mini testis. mum length= 30 steps, 38 required; consis 19. Testes, Position: 0= tandem, not over tency index, Cl, excluding uninformative lapping; 1 = tandem, overlapping. characters= 0.778; homoplasy index, HI= 20. Body, L:W Ratio: 0= approximately 1:6- 0.222) 7; 1 = elongate, l: 14; 2= approximately 1:2-3. Coding for this character is Morphological characters for phylogenetic consistent with functional outgroup analysis of Ortlt osplanchnus spp. criteria. I. Cuticular Spination: 0= complete; 1 = Character Matrix fo r Phylogenetic Analysis confined to anterior. of Orthosplanchnus spp. 2. Body Length: 0= elongate, ;::: Omm; 1 1 = medium, � 4-7mm; minuscule, ::::: 2 mm. Characters 1-20 3. Oral Sucker, Diameter: 0= mean � Campula spp. * 00000 00000 00000 00000 500J.!m; 1 = mean ::::: 350 J.!m; 2= mean ::::: 0. arcticus 0 1001 11110 00111 10 1 00 250 J.!m. 0. fraterculus 0 1111 22110 11111 1 0000 4. Ventral Sucker, Diameter: 0= mean ;::: 0. rossicus 10001 11110 12111 1 00 11 500 J.!m; 1 = mean ;::: 350 J.!m; 2= mean ::::: 0. pygm aeus 12221 3 2121 1 0 111 0 1102 250j.!m. 0. antarcticus 01111 21110 11111 00001 5. OS:VS Ratio: 0= < 1:1; 1= 1:1; 2= 1:2. 0. oculatus 12221 32121 10111 01112 6. Pharynx, Diameter: 0= mean > 500 J.!m; 0. albamarinus 10002 10010 00000 00010 I= mean � 300 J.!m; 2= mean � 200 J.!m; 3= mean < 200 J.!m. *Outgroup 7. Ovary, Diameter: 0= mean > 450 J.!m; 1 =mean � 250 J.!m;2= mean < 200 J.!m. 8. Ovary, Form: 0= ellipsoidal to elongate; 1 = ovoid or subspherical. 9. Testes, Form: 0= dendritic, highly lo bate; 1 = elongate to ellipsoidal; 2= ovoid. 10. Testes, Position: 0= middle third of body; 1= third quarter of body. BullScand Soc Parasito/ 2000; 10 (2): 38 PARASITE APPLICATION IN CONCERVATION AND PEST CONTROL P.J. Hudson Institute of Biological Sciences, University of Stirling, Stirling Scotland, UK Parasitic infections have an impact on Direct and indirect means of applying the fitness of individuals and this can anthelmintic to red grouse (Lagopus have consequences for the population of lagopus) to control Tr ichostrongylus the host species. In natural populations tenuis will be examined. The control of of economic or conservation value then parasites in a reservoir host species and we may wish to reduce parasitic infec the timing of application in relation to tions to increase yield or maintain bio seasonal infection will also be examined. diversity. Alternatively we may wish to Parasites can also be used to control pest use parasites to control pest species. This species. The ways entomopathogenic paper will examine some of methods that nematodes can be used to control insect can be used and their efficacy in the pests will be considered. manipulation of parasites in wild animal populations. Bull Scand Soc Parasito/ 2000; 10 (2): 39-43 FRESHWATER FISH PARASITES AND HABITAT CHANGE C.R. Kennedy School of Biological Sciences, University of Exeter, Exeter, EX4 4PS ,UK Tel.: +44 1392 263757; Fax +44 1392 263700; e-mail: [email protected] The rate of habitat change appears to with such multiple causality it can be be exceptionally rapid in fr eshwaters. very difficult to distinguish the determi Many contemporary changes are anthro nant variables. Most studies relating fish pochore in origin, so all too often several parasite communities to habitat change factors change more or less simultane have fo cused on the local, anthropochore ously. These changes must and do influ changes and have been short term in ence the biota, including the parasite nature. However, to address the effects fa una. The effects on parasites may be of climatic induced habitat change it is direct on the fr ee living stages or on necessary to employ long term data sets ectoparasites, or indirect via effects on and these are sadly in short supply. fish or invertebrate hosts but the exis This present study employs long term tence of a causal relationship between data sets in order to investigate whether anthropochore habitat change and habitat changes caused by climatic change in metazoan parasite communi events can be detected and recognised as ties in freshwater fish is undeniable such in the presence of anthropochore (Kennedy, 1997a). changes and then correlated with These anthropochore changes are changes in helminth parasite communi essentially local in nature and generally ties in fi sh. The climatic variables se affect a single catchment or water body. lected are droughts and floods and tem They are, however, superimposed on a perature extremes, as these are generally background of regional changes which monitored on both local and regional are natural and climatic. These may be scales. If they do have an impact on discrete events such as floods and parasite communities we can predict droughts or a continual change such as a similar and simultaneous changes in rise in temperature. The two types of discrete localities within a region. This change may sometimes be linked as hypothesis will be tested by examining when a drought accentuates the effects the changes in parasite communities in of water abstraction, or floods negate the the European eel Anguilla anguilla in attempts at regulation, and when faced three rivers and a lake in South Devon 40 over periods varying from 14 to 30 tenerrima and A. clavula has become years. The data sets were originally very rare Eel specialists including Pro collected fo r other purposes and two of teocephalus macrocephalus and Dero them have been published (Kennedy, prists injlata have appeared in the com 1993, 1997b). munity since 1991, as have non The Environment Agency has identi specialists including Neoechinorhynchus fied 1976, 1984 and 1989 as drought rutili and Raphidascaris acus. The ap years in South Devon, in which mini pearance of D. injlata may relate to river mum flows were repotied from the River management, such that tidal influence Exe as 36.5, 103.4 and 104.9 respec may now be fe lt fmiher upstream, but tively in comparison to the normal of none of the other species appearances 165.4 (Mild). By contrast 1981, 1986 can be related to climatic or other par and 1988 were exceptionally wet years. ticular events. The crash in community The winters of 1978 and 1984/85 were species richness co-incided with the also exceptionally cold in South Devon, drought year of 1984, but there was no to the extent that in 1985 some lakes discernable effect of the drought of froze over for the first time in 22 years. 1989. These climatic variations are natural but The R. Culm is also a tributary of the severe enough to be rated as exceptional. R. Exe and joins the main river above In addition, the last decade has seen Exeter. It is 45km long, and until re some of the warmest years on record. cently had water quality close to class C The R. Clyst is a small (25km) tribu (fairly good) as a consequence of pollu tary of the R. Exe and joins the main tion from paper mills. The mills have river in the estuary. Its water quality is now closed and the river has become only class B (moderate), it does not grade B.Brown trout are present in the harbour brown trout, Salmo trutta, and it river and chub Leuciscus cephalus have is becoming enriched from farm drain been introduced into it. Samples have age. It has been subject to a multiplicity been taken over an 18 year period from of local changes including straightening, 1981 - 1998. In 1981 B. claviceps was deepening, dredging and bank stabiliza the only species present in the intestinal tion. Samples were taken over 20 years, community and this species is still pre from 1979 - 1998 (Kennedy, 1993). Up sent. In 1986 species richness increased until 1982 the intestinal helminth com to three with the detection of P. tener munity comprised three species, with rima and Acanthocephalus lucii. The Acanthocephalus clavula dominant over introduction of chub was coincidental Bothriocephalus claviceps and Para with the appearance of Pomphorhynchus quimperia tenerrima. The number of laevis in the river and by 1997 this spe species dropped to 2 in 1983 and to 0 in cies dominated the intestinal community 1984. By 1991 community richness had in eels. Species richness had increased to risen to 9 species and has subsequently 8 by 1998 with the appearance of eel fluctuated between 7 and 9 species. In specialists such as Acanthocephalus addition to enrichment, the identity of clavula and Sp initectus inermis but P. the dominant species has changed to P. laevis continues to dominate the corn- 41 munity. The community here has never drought of 1976. This enrichment also fa llen to zero richness or even declined, led to winter-kill of fish when the lake but has steadily increased in richness froze over during the witner of 1984/85 over the period and this may be related (Kennedy, 1996). These events have had to the recovery of the river from the a profound effe ct upon the fishery and effects of the mill pollution. There is no the parasites of fish in the lake. The indication that any of the droughts or sampling programme here extends over cold winters affected the community. 30 years, from 1971 to 2000. Because of The R. Otter catchment is adj acent to its isolation, parasite communities of fish that of the R. Exe. The river is 44 km tend to be rather poor. The lake is one of long and flows directly to the sea. It is the few localities from which the gill subject to borehole extraction and agri copepod Ergasilus gibbus has been cultural input, but the water quality is recorded from eels in Britain. In 1971 class A. The sampling programme has and 1975, the only intestinal species extended over 14 years, from 1985 - recorded from eels was Acanthocephalus 1996 (Kennedy, 1997b). In 1985 only B. lucii. By 1985, this had been replaced by claviceps was present, and it is possible A. clavula. By 1987 B.claviceps and P. that the low species richness might also tenerrima were also reported in eels, relate to the 1984 drought, but the com although A. clavula dominated the com munity had increased to 8 species by munity. Thereafter A. clavula, together 1987 and then declined to 1 in 1988 and with other species of acanthocephalans, 0 in 1989. This decline preceeded the gradually declined in the lake and even drought of 1990. Community richness tually, around 1990, became locally then varied between 4 and 8 from 1991 extinct. From 1995 - 1999 the commu to 1998. The only new species to appear nity comprised fo ur species with, P. after 1989 was the generalist Sp haero tenerrima dominating over B. claviceps, stoma bramae as all the other species P. macrocephalus and Pseudocapillaria had been reported in the river before this and E. gibbus is still present on the gills. Once again, none of these changes can time. Both P. laevis and S. inermis were first recorded in the 1987, but both in be related to local habitat changes or to creased in abundance from 1991 on climatic changes although both have had major effects upon the parasite commu wards until S. inermis dominated the community fr om 1996-1998. The ap nities of other species of fish in the lake. pearance of P. laevis is believed to be The expansion in range throughout due to stocking of the river with brown Europe of Anguillicola crassus and trout, but none of the other changes can Pseudodactylogyrus anguillae over the be clearly related to climatic or local last two decades has extended into habitat events. Devon and since 1995 both have been Slapton Ley is a small, isolated fo und in all fo ur localities. coastal lake in South Devon. The lake All four localities experienced the has suffe red from increasing eutrophica same regional climatic regrme of tion which switched into hyper droughts, floods and temperature eutrophication as a consequence of the changes. All fo ur, however, were also 42 subject to local anthropochore changes, ity in the early part of the study period. the nature and magnitude of which dif Changes in the proportions of these fered in each case. The R. Clyst was sub species occured throughout the study ject to management and flood relief period and A. clavula was often replaced changes, the R. Culm to pollution and by another, non-eel specialist acantho recovery therefrom, the R. Otter to bore cephalan, but the other species were hole abstraction and agricultural run off present throughout. and Slapton Ley to eutrophication. At 4. The identity of the dominant spe least two rivers were also subject to fish cies changed: P. tenerrima increased in stocking from another catchment. In all abundance in all localities and came to fo ur localities the helminth communities dominate 3 of them in the late 1990s. in eels showed changes over the period 5. In all four localities there was an of study. The problem lies in relating the parasite changes to the habitat changes, overall increase in component commu nity species richness over the period of and distinguishing regional and local study. This involved additions to the influences. common suite due to acquisition of the If regional, climatic factors are im specialist Pseudocapillaria in all four portant influences on helminth com localities and to the non-eel specialists munities then the changes in the com N rutili, R. . acus and S. bramae in the munities should be similar and synchro three rivers. In the rivers close to the nous. A number of such changes were in estuary and subject to tidal influences, fact observed : D. injlata appeared. In some cases, in 1. All fo ur communities were invaded troduced species such as P. laevis and S. by A. crassus and P. anguillae. This, inermis increased species richness. however was not related to local or There were also different and asynchro regional factors as such: both species nous changes between the communities: have been spreading throughout Britain over the last 15 years (Kennedy & 1. Eel parasite communities in the R. Fitch, 1990) and it is not surprising that Culm and R. Otter were enriched by they reached the far west so late. Dani species introduced with stocked fish: P. conema anguillae was also fo und in all laevis arriving with chub in the fo rmer fo ur localtities, but this is more likely to river and trout in the latter. These an reflect improved detection rather than a thropochore stockings were independent colonisation event. events. 2. Over the last 20 years there has 2. The rare eel specialist S. inermis been a decline in the prevalence and became common in the R. Otter and R. abundance of A. clavula in the region, Culm, but not in the R. Clyst. It was first such that this once common species has detected in the R. Otter in the same year become extinct in one locality and rare as P. laevis but this must be co in three others. incidental as eels are the only host for the species and cannot serve as a defini 3. A common suite of eel specialists tive host fo r P. laevis, S. inermis did not comprising B. claviceps, P. tenerrima appear in the R. Culm until 10 years and A. clavula was present in each local- 43 later, but the source of it in both rivers is ture state to a mature fo rm. Each phase unknown. in a life-cycle must be capable of carry 3. Community species richness de ing out its particular fu nction. Free liv clined to zero in the Rivers Clyst and ing larvae must be required to penetrate Otter but there was no such decline in a host, parasitic larvae must survive in the R. Culm which was adjacent to the the body of an intermediate host, adults Clyst. should be capable of reproduction. To ensure that these tasks may be completed 4. There was no synchrony in the successfully various adaptations (mor times of zero species richness. phological and behavioural) have been 5. Species richness increased steadily developed in the course of evolution. in the R. Culm, perhaps in response to These adaptations are specific and de recovery of the river from pollution and pend on the life-cycle phase and ecologi a corresponding increase in invetiebrate cal conditions of the environment. diversity. Therefore, it is impossible to discuss 6. The community also increased in evolution of life-cycles without consid richness in Slapton Ley over the period erating the ecological conditions that of eutrophication and through events influence these life-cycles. This is the such as winterkill that caused the disap essence of the ecological approach pearance of many parasite species fr om which we use in our studies. other species of fish (Kennedy, 1998). Whilst species of Dactylogyrus disap References peared, E. gibbus was unaffected. Kennedy CR. The dynamics of intestinal helminth communities in eels The overall conclusion must be that Anguilla an guilla in a small stream: long tenn changes in there is very little evidence that discrete, richness and structure. Parasitology 1993; regional, climatic events such as 107: 71-8 droughts have any lasting effect upon Kennedy CR. Freshwater fish parasites and helminth communities of eels. By con environmental quality: an overview and trast, these communities are strongly caution. Parassitologia 1997a; 39: 249-54 affected by local anthropochore events Kennedy CR. Long term and seasonal and especially introductions. is possi It changes in composition and richness of ble to detect correlations between cli intestinal helminth communities in eels mate improvement and increasing spe Anguilla anguilla of an isolated English cies richness in all fo ur localities: as river. Folia Parasitologica 1997b; 44: 267-73 with other suggestions on the effects of Kennedy CR. Aquatic birds as agents of global warming, however, the evidence parasite dispersal: a field test of the effec fo r causality is so far lacking. tiveness of helminth colonization strategies. Trematode life-cycles are amazingly Bull Scand Soc Parasit 1998; 8: 23-8 complex and diverse. They are character Kennedy CR, Fitch DJ. Colonization, larval ized by changes of generations (parthe survival and epidemiology of the nematode nogenetic and hermaphroditic) and ani Anguillicola crassus, parasitic in the eel mal hosts (intermediates and final). Each Anguilla anguilla, in Britain. J Fish Biol generation must develop fr om an imma- 1990; 36: 117-3 12 BullSc and Sac Parasito/ 2000; 10 (2): 44-48 THE EVOLUTIONARY ECOLOGY OF PARASITE-INDUCED CHANGES IN HOST PHENOTYPE R. Poulin Department of Zoology, University ofOtago, P.O. Box 56, Dunedin, New Zealand Phone: 64 3 479-7983; Fax: 64 3 479-7584, E-mail: [email protected] Since it was first convincingly illus nomena often become rapidly and widely trated by Holmes, Bethel (1972), the accepted by the scientific community. ability of many parasites to manipulate Following an initial enthusiasm fo r the the phenotype of their host in order to new paradigm, there is typically a de fac ilitate their own transmission has crease in interest fo llowed by a shift become a paradigm in the area of host toward another explanation. This pattern parasite evolutionary ecology. After two is so common that Kuhn ( 1996) pro decades during which examples had posed that it was the normal way in been accumulating, the fo cus of research which scientific progress and revolutions in the past ten years has shifted toward are achieved. The acceptance of host the evolution and adaptive nature of the phenotype manipulation as an adaptive phenomenon (Moore, Gotelli, 1990; parasite strategy was no diffe rent. Re Poulin, 1994, 1995; Km·is, 1997; Brown, cently, I used published estimates fr om 1999) and its physiological basis (Hurd, the past 30 years of the magnitude of 1990; Thompson, Kavaliers, 1994). parasite-induced changes in host behav More recent lines of research have high iour to test for a potential weakening of lighted different issues relating to host the manipulation paradigm (Poulin, manipulation by parasites, and this essay 2000). I used data on the effects of will explore some of them. I will first helminths on the behaviour of their host, draw attention to a rather disturbing and quantified the effects of parasites as historical trend in the quantification of the difference between the mean behav parasite-induced changes in host pheno iour of infected and control host groups, type, and then discuss the more general corrected for sample size and the pooled implications of altered host phenotypes standard deviation of individual behav for the evolution of hosts and parasites. ioural measures. I then tested the null New ideas that provide simple, ele hypothesis that the magnitude of the gant explanations for widespread phe- published effects of parasites has re mained the same through the past 30 45 years. Using only data from studies of However, the magnitude of the effects of the effects of helminths on the behaviour parasites on host phenotype are usually of their definitive host (in which a more subtle and much less spectacular change in host behaviour has no obvious than in the first published examples that effects on parasite fitness), this is indeed are still the most widely cited. what I fo und: the magnitude of published With this cautionary remark in mind, estimates of parasite-induced changes the time has come to examine other does not vary over time. However, using evolutionary aspects of the host manipu data fr om studies looking at the effects lation phenomenon. Beyond its fitness of parasitic helminths on the behaviour benefits for the manipulating parasite, of their intermediate host, the published the alteration in host phenotype can have estimates of parasite-induced changes in serious implications fo r the evolution of host behaviour decreased significantly the host itself, as well as that of other over time (Poulin, 2000). All these stud parasite species sharing it with the ma ies were explicit tests of the adaptive nipulating species. Looking first at the host manipulation hypothesis. They all host, it is clear that host manipulation by were on systems in which intermediate parasites can change the shape of the hosts must be ingested by the definitive frequency distributions of various con host fo r parasite transmission to occur, tinuous phenotypic variables within the and thus systems in which alterations in host population (Poulin, Thomas, 1999). intermediate host behaviour can affect Infection by a manipulating parasite can their susceptibility to predatory defini increase or decrease the mean value of a tive hosts. Another analysis on an inde phenotypic trait and increase its variance pendent data set, in which the parasite in the overall host population. This will mediated increase in predation of defini of course depend on the prevalence or tive hosts on intermediate hosts was abundance of the manipulating parasite. quantified instead of actual behavioural If the parasite is very common, the fr e changes, showed the same pattern: the quency distribution of host phenotypic published estimates of increases in sus traits may remain normal but be shifted ceptibility to predation (measured as the toward higher or lower values; if the increase in the probability of being eaten parasite is only moderately common, the resulting from infection) decreased distribution of host traits is likely to significantly over the past three decades become skewed (Poulin, Thomas, 1999). (Poulin, 2000). The importance of this effe ct is that What does this all mean? Well, a because of the parasite, the observed publication bias may be the main factor distribution of host traits in the popula responsible for the historical trend. tion no longer reflects the distribution Reviewers and editors may be more expected from host genotypes. This willing to publish weak results or excep parasite-mediated uncoupling between tions to the paradigm years after its first host genotype and phenotype can disrupt popular acceptance than at the begin selection acting on hosts. The strong ning. The existence of the host manipu background noise generated by parasites lation phenomenon is not in doubt. renders selection myopic, i.e. blind to the 46 genotype and only capable of seeing and site have received ve1y little attention to acting on the presently expressed pheno date (Poulin, Thomas, 1999). types. One possible consequence would Finally, what are the ecological and be a slowing down of evolutionary (ge evolutionary consequences of host ma netic) changes in specific host traits from nipulation by one parasite species fo r the generation to generation (Poulin, Tho other parasite species that utilise the mas, 1999). There has been no attempt to same host population? In the case of date at quantifying these s01ts of secon larval helminths exploiting the same dary evolutionary effects of parasite intermediate host population, there may manipulation of host phenotypes. be either shared interests or conflicts In more extreme cases, manipulating between the manipulating parasite and parasites cause such a large shift in host the other helminth species, depending on phenotype that they split the population whether they also share the same defini into two morphotypes, i.e. the frequency tive host. If we consider the intermediate distribution of certain host phenotypic host as a vehicle taking parasites toward traits becomes truly bimodal. This is true their definitive hosts, then the manipulat of ce1tain helminths that alter the colour ive parasite can be seen as the driver, ation or morphology of their intermedi since it influences the direction taken by ate hosts to the point that a human ob the vehicle, i.e. the definitive host most server can easily separate parasitized likely to eat the intermediate host individuals fr om healthy ones simply by (Lafferty, 1999). Other parasites with the looking at them (e.g., Hindsbo, 1972; same definitive host as the manipulator Plateaux 1972; LoBue, Bell, 1993). In can f0 1tuitously benefit fr om the other systems, manipulating parasites manipulation when they happen to share cause a shiftin the spatial distribution of an intermediate host with a manipulator, their hosts, such that parasitized indi or they can evolve the ability to seek viduals are segregated in space from manipulated hosts and 'hitch' a ride with healthy conspecifics. For example, a manipulating parasites (Thomas et al. , microphallid trematode causes its am 1998). If two manipulating species ex phipod intermediate hosts to move to ploit the same intermediate and defini ward the water surface whereas unpara tive host populations, they may evolve a sitized amphipods stay at the bottom of tendency to seek one another if their their lagoon habitats (Helluy, 1984 ). effects are additive, or one may lose its This marked spatial segregation causes manipulative ability and associate with parasitized amphipods to mate assorta the other one to obtain a cheap ride tively with one another, whereas unin toward the definitive host (Laffe rty et fe cted amphipods tend to mate only with al. , 2000). However, if a parasite species other uninfected conspecifics (Thomas et shares an intermediate host population al. , 1995). The evolutionary effects of with a manipulator, but if they have spatial segregation and reduced gene diffe rent definitive hosts, then there is a fl ow between parasitized and healthy potential fo r conflict between them. The individuals in a host population as a unlucky passenger may evolve ways to consequence of manipulation by a para- avoid the manipulator in order not to be 47 taken into a dead end (i.e. the wrong cused on some of the evolutionary con definitive host). If both parasite species sequences of host manipulation for are capable of manipulating intermediate organisms other than the manipulating hosts but influence diffe rent phenotypic parasite itself. Host manipulation weak traits, there may be a struggle for control ens the link between the genotype and of the host, with one species ending up the phenotype of the host, with possible in the driver seat and the other interfer effects on the speed of selection (Poulin, ing without success (see Lafferty et al., Thomas, 1999), and it modifies the host 2000). All these situations are simpli landscape in which other sympatric fied; in reality, the outcome of these parasites evolve (Lafferty et al. , 2000). sympatric associations would depend on In addition, manipulative parasites can the prevalence of the two parasites, on have indirect effects on other organisms, their life history and developmental not pmi of the host-parasite complex characteristics, etc. (see Thomas et al. , 1999). These kinds The point of all the above scenarios is of consequences of host manipulation, in that the evolution of host selection addition to further insight into the adap mechanisms and transmission strategies tive nature of manipulation, should be in parasites with complex life cycles is the fo cus of future studies in this area. not independent from the strategies used by other, sympatric parasite species References when host manipulation is involved. The Brown SP. Cooperation and conflict in host first prediction that can be made fr om manipulating parasites. Proc R Soc London B the observation that a manipulator shares 1999; 266: 1899-904 an intermediate host population with Helluy S. Relations h6tes-parasites du trema other species is that there should be tode Microphallus papillorobustus (Rankin, positive or negative associations be 1940). Ill. Facteurs impliques dans les modi tween parasite species among intermedi fications du comportement des Gammarus ate hosts, depending on what definitive h6tes intermediaires et tests de predation. hosts they use. Many recent studies (see Ann Parasitol Hum Comp 1984; 59: 41-56 Laffetiy et al. , 2000 for examples) have Hindsbo 0. Effects of Polymorphus (Acan indeed fo und such associations. The thocephala) on colour and behaviour of search for coevolved transmission Gammarus lacustris. Nature 1972; 238: 333 strategies among sympatric parasites Holmes JC, Bethel WM. Modification of should reveal a wide array of diffe rent intermediate host behaviour by parasites. In: tactics illustrating the broader conse Canning EU, Wright CA, eds. Behavioural quences of host manipulation. aspects of parasite transmission. London: Academic Press, 1972: 123-49 Despite the historical decrease in the magnitude of the published estimates of Hurd H. Physiological and behavioural its effect, the manipulation of host phe interactions between parasites and inveJie notype by parasites remains an impmiant brate hosts. Adv Parasitol 1990; 29: 271-3 18 phenomenon. It is widespread in natural Kuhn TS. The structure of scientific revolu systems, having evolved de novo in tions, 3'd edition. Chicago: University of many parasite taxa. This essay has fo - Chicago Press, 1996 48 Kuris AM. Host behavior modification: an Thomas F, Renaud F, Derothe JM, Lambert evolutionary perspective. In: Beckage NE, A, de Meetis T, Cezilly F. Assortative pairing ed. Parasites and pathogens: effects on host in Gammarus insensibilis (Amphipoda) hormones and behavior. New York: Chap infected by a trematode parasite. Oecologia man & Hall, 1997: 293-315 1995; 104: 259-64 Lafferty KD. The evolution of trophic trans Thomas F, Renaud F, Poulin R. Exploitation mission. Parasitol Today 1999; 15: 111-15 of manipulators: 'hitch-hiking' as a parasite transmission strategy. Anim Behav 1998 56: Lafferty KD, Thomas F, Poulin R. Evolution ; 199-206 of host phenotype manipulation by parasites and its consequences. In: Poulin R, Morand Thomas F, Poulin R, de MeetisT, Guegan I S, Skorping A, eds. Evolutionary biology of F, Renaud F. Parasites and ecosystem engi host-parasite relationships: theory meets neering: what roles could they play? Oikos reality. Amsterdam: Elsevier Science, 2000; 1999; 84: 167-71 (in press) Thompson SN, Kavaliers M. Physiological LoBue CP, Bell MA. Phenotypic manipula bases for parasite-induced alterations of host tion by the cestode parasite Schistocephalus behaviour. Parasitology 1994; 109: S 119- solidus of its intermediate host, Gasterosteus S138 aculeatus, the threespine stickleback. Am Nat 1993; 142: 725-35 Moore J, Gotelli NJ. A phylogenetic perspec tive on the evolution of altered host behav iours: a critical look at the manipulation hypothesis. In: Barnard CJ, Behnke JM, eds. Parasitism and host behaviour. London: Taylor & Francis, 1990: 193-233 Plateaux L. Sur les modifications produites chez une fourmi par la presence d'un parasite cestode. Ann Sci Nat Zoo! Bioi Anim 1972; 14: 203-20 Poulin R. The evolution of parasite manipula tion of host behaviour: a theoretical analysis. Parasitology 1994; 109: S 1 09-S 118 Poulin R. 'Adaptive' changes in the behaviour of parasitized animals: a critical review. Int J Parasitol 1995; 25: 1371-83 Poulin R. Manipulation of host behaviour by parasites: a weakening paradigm? Proc R Soc London B 2000; (in press) Poulin R, Thomas F. Phenotypic variability induced by parasites: extent and evolutionary implications. Parasitol Today 1999; 15: 28- 32 Bull Seand Soc Parasito/ 2000; 10 (2): 49-54 INFRACOMMUNITIES: STRUCTURE AND COMPOSITION O.N. Pugachev Zoological Institute Russian Academy of Sciences, Universitetskaja nab.l St. Petersburg, 199034, Russia Fax: +812 - 3282941, E-mail: [email protected], [email protected] Parasite communities from single host of 1-2 parasite species. Frequency individuals have one undoubted fe ature: distributions of parasite species are the overwhelming majority of parasite overdispersed (a2/M= 2.23) and can be infrapopulations cannot maintain their fitted by Gamma or Poisson distribu numbers by self-reproduction. These tions. The same distribution type was communities are rigorously defined spa fo und when infracommunities were tially. The existence of an infracommunity grouped according to fish ecology or is restricted to its individual host's life systematic position. This distribution is time. However, at this level parasites only typical fo r the frequency distribution of interact or do not interact, compete or do numbers of a single parasite species in not compete with each other, and either one host population. Parasite species fo rm or do not fo rm guilds. As a result of frequency distributions vary in differ these interactions isolationist or interac ent component commumt1es: from tive communities are fo rmed. typically overdispersed to asymmetri Holmes & Price (1986) suggested that cal. This reflects diffe rent life condi the host organism is analogous to an tions fo r different host populations. "island" which is inhabited by parasites. Thus, one of the impotiant parameters A number of studies of communities of of infracommunity - species number is intestinal helminths have clearly shown stochastic. that even component communities consist For all host individuals a low prob of few helminth species (5-10). But taking ability of events occurring and ap into account host numbers one can assume proximately equal probabilities of a a great number of empty or partly popu parasite being acquired (but not estab lated "islands" (Simberloff, 1990). Fre lishing) are the basis of overdispersed quency distributions of numbers of para or aggregated distributions. Research site species in 899 infracommunities into free-living commumttes often revealed the fact that most of them consist fo cuses on spatial segregation 50 ( overdispersion) and on changing patterns despite the fact that both parasite of spatial use as evidence fo r interspecific groups are nearly equally represented interactions, particularly competition in the region (44% and 56%). Thus, (Simberloff, 1990). Overdispersed species parasites with direct life cycles which distribution seems not to be a feature at are more dependent on the external the infracommunity level or even to have environment but with a greater prob no sense because of the impossibility of ability of infecting the host are less studying the same host individual several often dominant in infracommunities times. Neve1iheless it is difficult to fit than parasites with complex life cycles. another type of distribution fo r an associa This is one evidences of the interactive tion each member of which is overdis nature of infracommunities. On the persed. This phenomenon is based on a other hand spatial segregation very low probability fo r parasites to com ( overdispersion) is the result of interac plete their life cycle. If prevalence is the tion (ghost of competition in the past) probabilistic estimation fo r the event (to when all problems within the commu complete life cycle), the next probabilistic nity have been solved, whereas any estimation could be done fo r a cestode: P= disturbance of overdispersion is the 0,0005*0,05*0,8=0,00002, where the first sign of serious infracommunity number is prevalence of infection of a reorganization when competition plays planktonic copepod, the second - preva an important role. These periods can be lence of infection of the second interme shmi, fo r example during fish spawn diate host (fish) and the third number - ing, migration etc. prevalence of infe ction of the final host Simberloff (1990) stated that "host (predatory fish). Perhaps the probability individuals may more closely approxi of infecting a host increases fr om first mate replicates of one other, than is intermediate to final host. Hence the lands do". It could be supposed that a probability of occurring in the same host real island has a greater variety of individual comes to zero as we have to biotopes than a host individual, but host add another multiplier 1/N where N is the population genotype variety, diversity number of host specimens in the popula and the extent of host reaction to "para tion. This very low probability is compen site-colonists" are more diverse or at sated fo r by high fe cundity, reservoir least the same as on a real island. The parasitism etc. Such a prope1iy is typical main difference is that species coloniz of parasites with complicated life cycles ing a real island compete with each that are less dependent on the external other while the parasite habitat (host environment. It could be supposed that organism) reacts actively to such a parasites with direct life cycles which colonist's occurrence. The other diffe r have a greater probability of infecting the ence is that the number of "hosts host will be found more often and in islands" is much higher than the num higher numbers than parasites with com ber of islands in any archipelago. It is plex life cycles, which means that they well known that the number of species will be dominant species in infracommu inhabiting an island depends on the size nities. However, these species were domi of the island. In parasitology the analo- nant in only 25% of infracommunities 51 gous parameter could be host size. We use Cyprinidae show weak a significant fi sh length as the measure of host size. No correlation of S and N with fish length great correlation was fo und between fish which increases slightly when infra length and species number or fish length communities with S=O are excluded. and number of parasite individuals (Fig. Among Salmonidae S was correlated 1). None of the infracommunity parame with fish length in the case of graylings ters (dominance index, evenness, Bril (r=0,33; p=O,Ol) when infracommuni louin index) correlated with host size ties with S=O were included. A nega either when all infracommmunities were tive significant correlation was fo und included into analyses or when infracom between S, N and fish length in the munities without multicellular parasites case of round whitefish (Prosopium (S=O) were excluded. This means that the cylindraceum) (r=-0,4678,p=0,003; r=- host organism is not completely analogous 0,3488,p=0,032). Such a correlation to an island. The same fact was fo und was not fo und among other salmonid when infracommunities were grouped species (Salvelinus malma, Brachymys according to fish ecology. When the sys tax lenok, Coregonus sardinella). The tematic position of the host was included picture is much more diverse among in the analysis the situation changed. Cyprinidae. Species of the genus Phox Salmonidae s.l. show a weak significant inus in a combined sample did not correlation between S and In N with fish demonstrate significant correlations length which became less when infra between S, N and fish length, while in communities with S=O were excluded. the case of species of the related genus y=2,677+0,00 1 *x 0 14 0 0 000 0 0 0 00 0 0 0 0 0<00 0 00 0 0 00 0 00 0 CD 0 0 CDOO 0 aa:cCD 0 0 arnao:m cm alDCJDro -2 � -200 0 200 400 600 800 1000 1200 Fish length Fig. 1. Relationship between fish length and number of parasite species (r= 0,077). 52 Oreoleuciscus such correlations were ago" could correlations be revealed. observed (r=0,5058, p=0,0001; Host species of different sizes can r=0,3534,p=O,OOOI). Lake minnow (Phox harbour many parasite species, fo r inus perenurus) demonstrated a weak example minnow and pike (Pugachev, negative correlation between S and fish 1997). length only if infracommunities with S=O The numbers of parasites (N) in in were excluded (r=-0,2512, p=0,045). fracommunities varied greatly. 95% of River minnow (Ph. phoxinus) demon infracommunities had less than 1000 strated a weak positive correlation be parasites; 86% - less than 500; 76% - tween N and fish length only if infracom less than 200. Frequency distributions munities with S=O were excluded of parasite numbers were overdispersed (r=0,282,p=O, 16, r=0,24, p=0,02). The as were numbers of parasite species. lacustrine fo rm of the two species of west Numbers of species and numbers of mongolian minnows ( Oreoleuciscus po parasite individuals are connected by a tanini and 0. pewzowi) from Big Lake significant positive correlation, the V alley demonstrated significant correla value of which increased considera tions between S, N and fish length. The blyly when infracommunities with third species of the genus Oreoleuciscus more than 500 parasite individuals ( 0. humilis ), which has lacustrine and (14% of infracommunities) or more riverine fo rms, demonstrated a significant than 200 (24% of infracommunitis) negative correlation only between S and were excluded from the sample. This fish length (r=-0,3591 ,p=O,O 15). No sig correlation between S and N was re nificant correlations between these pa vealed in the case of fish ecological rameters were found among lacustrine groups, systematic groups (Salmonidae fo rms of this species while its riverine and Cyprinidae) and all fish species fo rm demonstrated a significant correla except burbot (Lata Iota). The value of tion between N and fish length only the significant correlation varied fr om (r=0,4529,p=O,O 12). 0.38 to 0.45 in the case of fish ecologi Thus, two main parameters - number cal groups and from 0.31 to 0.66 in the of species and number of parasite indi case of diffe rent fish species and was viduals - of the infracommunity show very 0.35 fo r Salmonidae and 0.45 fo r Cy diverse correlations with host size only prinidae. The numbers of parasite when we take into account host phyloge individuals m infracommunities netic relationships. The diversity of these reached "saturation level" on the aver correlations demonstrates the impossibil age when infracommunities consisted ity of making a direct analogy with free of 150 parasite individuals and 9-10 living animal ecology. If we would like to species (Fig. 2). This "saturation level" make such an analogy in the case of "host was revealed in all fish ecological and island" theory then every host species systematic groups. Thus, the number of would consist of "populations parasite individuals in the infracommu archipelagos which are situated in the nity being a stochastic and overdis same geographical zone". Only among persely distributed value shows the "islands" of one and the same "archipel- tendency to achieve saturation on the ��� ---�·------ 53 Median=-68,589+ 1 84,73 1 *log 1 O(x) 550 450 350 D 250 z J - 150 I Max 50 M in D 75% 25% -50 u-/ 0 2 3 4 6 7 8 9 10 11 12 13 14 15 16 � Median s Fig. 2. Relationship between number of species (S) and numbers of parasites (N) within infracommunities (regression line by average values, N<500) average. This is possible evidence fo r in a fish and are "interested in" the recognizing an infracommunity as interac host's death and its availability to a tive, nonisolationist and saturated, which predator. The others are not. 55% of is the result of interactions between host oversaturated infracommunities had organism and parasites as well as between dominant parasites of the first group. parasites. Only if we suggest that a host Thus, such parasite species exert an organism is an indifferent habitat can we important influence on the fate of an come to the conclusion that there are infracommunity. A rapid increase in "empty niches" and unlimited food re parasite numbers corresponds to their sources. Absence of parasites is not evi life strategy. These oversaturated infra dence of the existence of empty niches. communities have to have short life Niches can be "closed" by host reactions spans. In these cases parasites from the or even be non-existent. second group have little chance of There are infracommunities which completing their life cycles. consist of a large number of parasite An analysis of relationships be individuals and which could be defined as tween the main three parameters of "oversaturated" (Fig.2). These oversatu infracommunities was carried out. As rated infracommunities mask the satura Brillouin index and evenness are con tion. Infracommunities consist of parasite nected by fo rmula their correlation is species with diffe rent life strategies. Some not discussed. Dominance index was parasites cannot complete their life cycle significantly negatively correlated with 54 number of parasite species (S), evenness 2. The reasons for stochasticity are and Brillouin indices but not with number large numbers of hosts and diversity of of parasite individuals. These correlations interactions within different host were fo und for fishecological groups, and parasite systems. fo r most species of Salmonidae and Cy 3. Analysis of all infracommunities prinidae. Evenness index was positively has demonstrated saturation and even correlated with S but not with N. The oversaturation for parasite numbers; same relationship is characteristic fo r fu nctional dependence between parasite Brillouin index also. Thus, three main species number and number of indi parameters of infracommunities demon viduals; dependence of S and N on host strate the usual relationships. When the size when host phylogenetic relation number of species in the infracommunity ships were considered; clear correla increases, evenness and Brillouin index tions between main infracommunity increases but dominance decreases. This parameters. is further evidence of the interactive na ture of parasite infracommunities. The 4. Being stochastic by nature infra character of such interaction, presence or communities are sufficiently predict absence of competition between or within able and interactive. species, presence or absence of competi tion between or within guilds, needs fu r References ther investigations. Holmes JC, Price PW. Communities of Fundamental infracommunity features parasites. In: Anderson DJ, Kikkawa J, eds. are: existence is restricted to the lifetime Community Ecology: Patterns and Proc of an individual host; spatial distinctness; esses. Oxford: Blackwell ScientificPublica high rates of emigration and immigration; tions, 1986: 187-213 co-existence of species with diffe rent life Pugachev ON. Comparative analysis para strategies; habitat (host organism) reacts site communities of pike and minnow. In.: actively to the parasite; parasite in Alimov AF, ed. Ecological monitoring of frapopulations cannot maintain their parasites. St. Petersburg: Zoological Insti numbers by self-reproduction. These tute RAS, 1997; 83-5 (In Russian) fe atures clearly distinguish parasite infra Simberloff D. Free-living communities and communities from communities of free alimentary tract helminths: hypothesis and living animals. In this respect a parasite pattern analyses. In: Esch G, Bush A, Aho infracommunity is a "subcommunity" or J, eds. Parasite communities: Patterns and "hemicommunity". Processes. London: Chapman and Hall, 1990: 289-3 19 Summary !.Analysis of infracommunity structure fo r separate fish species did not reveal any clear tendencies fo r main infracommunity parameters. The infracommunity looks stochastic and unpredictable. �������·------ BullScand Sac Parasito/ 2000; 10 (2): 55-60 WHAT DETERMINES THE LONGEVITY OF MAMMALIAN NEMATODES? 1 A. Skorping and A.F. Read2 1Department of Zoology, University of Bergen, Allegaten 41, N-5007 Bergen, Norway, email: [email protected] 2Institute of Cell, Animal and Population Biology, University of Edinburgh, West Mains Road, Edinburgh, UK. The life of a nematode within its final same host and the same organ, live host can conveniently be divided into two their lives so diffe rently? Since all phases: the prepatency or developmental species are the result of past selection period, and the period the worm produces we must assume that both short- and eggs or larvae - usually termed the long-lived nematode species are opti patency period. As shown in Fig. 1, after mally adapted, but why is it optimal fo r entering the final host some nematode a hookworm to hang on to the gut species mature in less than a week while mucosa and keep producing eggs fo r others take more than a year to reach the years, but not for pinworms? Answer reproductive phase. Once they have ing such question fo r individual species started to reproduce, some species will is likely to require detailed information continue reproduction fo r years, while about their basic biology, but we may others burn out after a few days. Much of gain some general insights by searching this variation in longevity could be related fo r patterns in this diversity, and by to diffe rent habitats or different hosts, but using a theoretical framework devel even among nematode species sharing the oped by evolutionary biologists. same organ within a single host species we find a staggering range in longevity. In Life history evolution the human gut, fo r example, Enterobius The theory of life history evolution vermicularis has a life-span of about 60 is based on the simple idea that life days, while Necator americanus may live consists of compromises (Steams, fo r more than 4 years. 1992). Ideally, the optimal organism To any biologist interested in evolution would start reproduction immediately such variation begs the question: Why do after birth and produce an unlimited diffe rent species, even when infecting the number of offspring fo rever. The rea- 56 Cl) ..0 0 '+-0 0 z 0 500 1 000 1500 2000 2500 3000 3500 Patency 40 r---��--�--�----�--�--�----� 35 30 Cl) ..0 25 0 '+-0 0 z 0 0 50 1 00 150 200 250 300 350 400 450 500 Prepatency Fig. 1. Frequency distributions ofprepatency and patency times of mammalian nematodes. reason such "Darwinian demons" do not viduals therefore are less fecund than exist is because organisms are limited by larger ones. But reaching a ce1iain size energy and time. Spending more energy takes time. Higher fecundity must on reproduction leaves less fo r processes therefore be paid for by a longer devel related to the maintenance of life, such as opmental time. tissue repair. We would therefore expect The optimal way for an organism to long-lived organisms to have less avail mature and reproduce must depend on able energy fo r reproduction per time than its ecological situation, or, more pre shorter-lived ones. In poitkilothermic cisely, on the expected age- or stage organisms there is usually an association specific mortality. Many organisms, between size and fecundity. Small indi- 57 including parasites, have distinct devel controlled fo r (Trouve et al, 1998). opmental stages in different habitats. In Certainly, the large variation in fecun situations where juvenile mortality is low dity observed among diffe rent parasite an individual could increase its fitness by species indicates that egg-production is prolonging its developmental time, in traded off against some other traits creasing its body size and fe cundity. If a related to fitness. change to bigger reproductive stages leads to higher adult mortality (for instance by COMPARATIVE STUDIES IN increased predation or immune attack), we NEMATODES OF MAMMALS would also expect a shortening of adult life span. The opposite situation - high In nematodes the most important juvenile mortality - should select fo r a factor limiting the number of eggs a shotier developmental time. If this leads worm can produce appears to be its to lower adult mortality, fitness could be body size (Skorping et al, 1991). increased by prolonging the reproductive Among intestinal nematodes of mam period. Although this theory claims to be mals the typically impressive daily general, it has mainly been applied and output of hundreds of thousands of tested on relatively few taxa of fr ee-living eggs per female only occurs among the organisms. larger species, such as Ascaris spp. The slope of the relationship between pre Are trade-offs between life history traits patency and body size is significantly different in parasites? higher than 1 indicating that the fecun Is there anything about the parasitic dity gain per time increases the longer mode of life that shape their life histories the nematode stays in the prereproduc diffe rently fr om free-living animals? tive growth phase. Additionally, bigger Certainly many cestodes appear to come nematodes tend to have a longer repro close to Darwinian demons with their ductive period (Fig. 2). Spending a long enormous fe cundity combined with a time in the prepatency phase therefore reproductive lifespan that may equal that seems to give a nematode a twofold of its host. Parasites are repeatedly re advantage - it becomes more fe cund ferred to as extravagant reproducers with and will produce eggs for a longer time. a significant higher egg-output than their Why then, are not all nematodes big, free living relatives (e.g. Kennedy, 1975; fecund and long-lived? Calow, 1983; Whittington, 1998), and The cost of becoming big is a long Jennings & Calow (1975) suggested that developmental time (Fig. 2). This will parasites were more fecund because the increase generation time and also the host provides them with a stable environ risk of dying before ever reaching ment and virtually unlimited food re maturity. Apparently, this disadvantage sources. However, the only study that has is a strong constraint on how long a addressed this problem using many differ nematode can afford to keep growing - ent species, and controlling for common the majority of nematode species are ancestry, fo und no diffe rences in repro relatively small (Poulin, Morand, ductive capacity between parasitic and 1997). fr ee living flatworms when body size was 58 Skorping, 1995), suggesting growth a "' N rates might depend on the habitat fo r ·;;; . . >- "' development. Indeed, comparing nema 0 .0 "' tode species developing in the gastroin ro "'E u. testinal system with species having a juvenile phase in other tissues, the Prepatency time latter group was fo und to have a highly significant larger size and higher growth rate (Read, Skorping, 1995) . .. b . If we assume that within a habitat "' ·: � there is a fixed optimal growth rate that i$' c can be attained, it has been hypothe � n. sised that the average mortality rate during development is a major determi Prepatency time nant of how long a nematode can risk to stay in the developmental phase in order to increase growth (and fe cun dity). Gemmill et a! (1999) developed a simple model assuming the optimal time of development fo r an intestinal . nematode to be proportional to the . .. . inverse of juvenile mortality rate. The growth rate was estimated from the Female body size slope of the relationship between pre patency and body size, and juvenile mortality rate fr om experimental meas Fig. 2. Three important associations between ures of prepatent period fo r a range of mammalian nematode life history traits. species, assuming a constant death rate. Worms that take longer time to develop tend Although these assumptions are overly to be bigger (a), and have longer reproduc simple, the model fitted remarkably tive periods (b). Bigger worms also have higher fe cundity (c). (After Skorping, Read well with observed data. & Keymer 1991) THE ROLE OF HOSTS One way a nematode can reduce the cost of having a big size, is to increase the Several authors have fo und a posi rate of development. However, speeding tive relationship between host and up growth rate might increase the instan parasite body size (Harvey, K.eymer, taneous death rate if more resources are 1991; Morand et al, 1996; Sorci et al, allocated to growth rather than to proc 1997). Is this relationship mainly esses related to avoiding immune de caused by lower m01iality rates in fences. We have many reasons fo r assum bigger hosts, or because a larger host ing that immune defences differ both can provide more energy? Harvey, qualitatively and quantitatively in diffe r Keymer ( 1991 ), examining pinworms ent parts of the vertebrate body (Read, in primate hosts, found that host life ------ 59 expectancy explained more of the varia appear to be less susceptible to tion in parasite size than host body size, a anthelmintics, than in their adult, intes pattern which has been confirmed fo r a tinal phase. Continuous exposure to larger dataset on pinworms by Sorci et a! such stage-specific anthelmintics (1997). We have argued elsewhere that should therefore select for worms variations in host longevity is unlikely to spending more time in the larval phase, have much selective effect on maturation and less time in the reproductive stage. times of nematodes in mammals, because This could lead to an increase in worm most nematodes species live within much body size. Much of the pathology shorter time scales than their hosts (Read caused by this group of worms is asso et a!, 2000). To a pinworm, developing ciated with the tissue phase, so pro within a couple of weeks, it cannot mat longing this stage is likely to lead to ter much whether the host lives fo r 5 or more virulent worms. In order to pre 15 years. By looking at all nematode dict the long term consequences of species within mammals for which we chemotherapy, a better understanding have been able to find data (533 species), of parasite life-histmy evolution is host body size explains much more of the therefore crucial. variation in parasite body size (and there Parasite communities from single fo re prepatency times) than host life host individuals have one undoubted expectancy (Skorping, Read in prep.). fe ature: the overwhelming majority of Nematodes therefore do become larger parasite infrapopulations cannot main and have longer maturation times in tain their numbers by self-reproduction. bigger hosts, but this can not be ex These commumttes are rigorously plained by the fact that larger hosts have defined spatially. The existence of an lower motiality rates. infracommunity is restricted to its individual host's lifetime. However, at EVOLUTIONARY EFFECTS OF this level parasites only interact or do ANTHELMINTICS not interact, compete or do not compete Nematodes in mammals are among the with each other, and either fo rm or do most important targets of the drug indus not form guilds. As a result of these try. If an anthelmintic increases the juve interactions isolationist or interactive nile mmiality rate, we would expect selec communities are fo rmed. tion fo r fa ster, smaller, less fecund - and probably less virulent nematodes - accord References ing to the model developed by Gemmill et Calow P. Patternand paradox in parasite re a! (1999). Apparently, modern drug use production. Parasitology 1983; 86: 197-207 should therefore have positive evolution Gemmill A W, Skm-ping A, Read AF. Opti ary consequences, in the sense that they mal timing of firstreproduction in parasitic will select fo r parasites that produce less nematodes. J Evol Biol 1999; 12: 1148-56 disease. However, the selective effects of Harvey PH, Keymer AE. Comparing life modern chemotherapy are not always this histories using phylogenies. Phi! Trans simple (Skorping, Read, 1998). Tissue (Royal Society ofLondon) B 1991; 332: migrating nematodes in their larval phase 31-9 60 Jennings JB, Calow P. The relationship be tween high fe cundity and the evolution of entoparasitism. Oecologia 197 5; 21: 109-15 Kennedy CR. Ecological animal parasitology. Oxford London Edinburgh Melbourne: Black well, 1975 Morand S, Legendre P, Gardner SL, Hugot, JP. Body size evolution of oxyurid (Nematoda) parasites: the role of hosts. Oecologia 1996; 107: 274-82 Pou1in R, Morand S. Parasite body size distri butions: interpreting patterns of skewness. Int J Parasitol 1997; 27: 959-64 Read AF, Skorping A. The evolution of tissue migration by parasitic nematode larvae. Parasi tology 1995; 111: 359-71 Read AF, Gemmill A W, Skorping A. Evolu tion of nematode life histories: theory meets reality? In Poulin R, Morand S, Skorping A, eds. Evolutionary biology of host-parasite relationships: Theory meets reality. Amster dam: Elsevier, 2000: 237-46 Skorping A, Read AF, Keymer AE. Life his tory covariation in intestinal nematodes of mammals. Oikos 1991; 60: 365-72 Skorping A, Read AF. Drugs and parasites: global experiments in life history evolution. Ecology Letters 1998; 1: 10-2 Sorci G, Morand S, Hugot JP. Host-parasite coevolution: comparative evidence for covaria tion of life history traits in primates and oxyu rid parasites. Proc R Soc Lond. 1997; 264: 285-9 Steams SC. The evolution of life histories. Oxford: Oxford University Press 1992 Trouve S, Sasal P, Jourdane J, Renaud F, Morand S. The evolution of life-history traits in parasitic and free-livingpla tyhelminthes: a new perspective. Oecologia 1998; 115: 370-8 Whittington I. Reproduction and host-location among the parasitic platyhelminthes. Int J for Parasitol 1997; 27: 705-14 Bull Seand Sac Parasito/ 2000; 10 (2): 61-64 GETTING TO THE CORE OF THE PARASITE COMMUNITITES: SEPARATING REALITY FROM NOISE E. T. Valtonen, K. Pulkkinen and M. Julkunen Depariment of Biological and Environmental Science, University of JyvaskyHi., P. 0. Box 35, FIN-40351 Jyvaskyla, Finland, tel. +358 14 260 2329, fax. +358 14 260 2321, e-mail. [email protected] Introduction There has been a wide-ranging debate maintenance of the parasite population on the fa ctors determining species rich (Holmes et al., 1977), and thus their ness and predictability of parasite com role in the exchange patterns of para munities. Analyses of patterns and proc sites between hosts may be questioned. esses of parasite communities have often Applying equal sampling effo rt is been made from published lists of host important when comparing component parasite records without a thorough communities of hosts in different areas. knowledge of each of the systems in However, in the analysis of component cluded and the methods involved in col communities of sympatric hosts within lecting the data. Recently, the effects of one area, the real relative abundance of e.g. unequal sampling effort and the influ hosts, which affect the sampling effo rt, ence of host phylogeny on the analyses are an important factor in determining have been discussed (Gregory, Black the population densities of parasites burn, 1991; Guegan, Kennedy, 1996; and thus the exchange between hosts Poulin, 1995, 1997a,b). However, there (Leong, Holmes, 1981 ). In this work are other possible confounding factors we show how area specific ecological which may have an effect on the exchange elements and scattered occurrences of patterns of parasites between hosts and parasites may mask or obscure the real thus the structure and predictability of the patterns in exchange or sharing of component communities of parasites. parasites between 3 1 sympatric fish Factors that have not been previously species. addressed in this connection are the effect of ecological elements within each study area and the significance of the rare para Material, methods and study area site species having scattered occurrences. Our study system is located in the On the other hand, it is known that scat Bothnian Bay, the nmiheastern part of tered occurrences are not impmiant in the the Baltic Sea, the largest brackish 62 water area in the world. The Bothnian Bay Results and conclusions is a unique open system which comprises Altogether, 63 parasite species were components of both marine and freshwa fo und in the 3I fish species studied ter origin on all trophic levels side by fr om the Bothnian Bay (Valtonen, side. Most of the fi sh species occurring in Pulkkinen, Julkunen, Poulin, unpubl.). the area are of freshwater origin, but the The number of parasite species fo und fish hosts also receive parasites from the in one fish varied from the minimum of marine part of the Baltic Sea. This has 0 parasites in Ammotydes tobianus to occurred via the marine fish species that the maximum of 26 species in Lota Iota have migrated into the area and estab . The host range of one parasite species lished resident populations or via acciden varied fr om those found in only one tal visitors from the southern Baltic Sea. host species, to generalist species, that Our data comprises metazoan parasite were fo und from up to 25 fish species. records (excluding monogeneans) of 31 Majority of the parasite species had fish species, of which the common ones only I to 4 hosts. 31% of the parasite were collected during two years as species occurred at prevalence below monthly or bimonthly samples from the 5% and 30% with prevalence between catches of commercial fishermen and 5-10%, such that 61% of the host para additional samples were taken of the less site records had prevalence below I 0%. common species. The relative proportions The numbers of the host-parasite re fish species studied were a good reflection cords with a low prevalence (< 5%) of the actual fish species composition in increased with the increase in the num the area. Newly caught fish were studied ber of fish studied. The maximum for metazoan parasites (excluding mono prevalence of each parasite species in geneans) using standardized parasitologi one host was usually much greater than cal methods. Parasites that occurred in a the mean prevalence in all other in host species with prevalence below 5% fe cted hosts, indicating that most para were considered as a scattered occurrence site species had one important host and in that particular host. Scattered occur several where they occurred at lower rences may be significant in the mainte prevalence. This patternap plied both to nance of a parasite population, however, if host-specific parasites (1 to 4 hosts) they accumulate in the fo od web either as and parasites which had 5 to 25 hosts. larval stages from intermediate hosts to Altogether, 84% of the host-parasite definitive hosts or as adult stages from a records with low prevalence ( <5%) prey fi sh to a predatory fish (post-cyclic consisted of parasite species that can transmission). We based our evaluation not accumulate in the fo od web, and of the accumulation of these scattered which can thus be regarded as scattered occurrences to predatory fishes within the occurrences in hosts that are not impor fish community both on published data on tant fo r the maintenance of parasite the life-cycles of the parasites and their population. These scattered occur ability to undergo post-cyclic transmission rences fo rm most of the noise that may and partly on detailed dietary analysis of disturb the observation of the important the fish species studied (E.T.Valtonen, interactions between the component unpubl.). 63 communities of parasites in different the scattered occurrences and further hosts. Further on in this presentation we also the diplostomids were omitted examine how the exchange pattern of from the data, the numbers of other parasites between hosts is altered when hosts that the parasites were shared the scattered occurrences are omitted from with decreased considerably and it was the data. Diplostomid metacercariae in the revealed that the hosts with the highest eye of fishes comprised one third of all number of parasite species generally parasite specimens in the Bothnian Bay. also shared the highest number of Since these parasites occur in almost all parasites occurring at high prevalences hosts, they form a significant component (> 5%) with other hosts. When the of the parasite community that is shared sharing of the parasites was studied between the hosts but since they are un within and between the fish families specific to any host, they fo rm another that were presented with more than one component of the noise which prevents us species in the Bothnian Bay, it was seeing the structure of exchange of para seen that the families shared 1 0 to 17 sites between hosts. We also will show the parasites between other families, when effe ct of omitting"the diplostomid cover" all host-parasite records were taken into on the exchange pattern of parasites. account. An exception to this was the When all host-parasite records were taken Cyprinidae, which shared only 5 to 8 into account, the number of other hosts parasite species with other families with which each host shared any of its (Table 1 ) . When the scattered occur parasites was always high regardless of rences and diplostomids were omitted the number of parasites the host species from the data, the number of species harboured, indicating that even the fish shared with other families was de species with a low number of parasite creased in some cases to only one species shared generalist parasites with species and the highest number shared several other host species. However, when was found between Salmonidae and 0) 0) "' '0 4-<-5 � � 0 t:: ·�Vl 1-< - '0 · · '0 0) 0) a "' 0 .2::- · a 1-< � "' ] � � 0 ·c: '0 0) :;:! '0 E ·u E ·u , c::: E '§ 0.. 1-< tl :;:; ] ::� "' Table 1. The number of parasite species shared within a fish family or between the families (the first number gives the shared species when all parasite occurrences were included, in brackets are the numbers of species shared when host-parasite records with a prevalence < 5% were omitted and the third number is the number of species shared when both the occurrences with a prevalences < 5 % and diplostomids (Diplostomum spathaceum and Ty lodelphus clavata) were omitted) 64 Gadidae (7). However, the numbers of Poulin R. Phylogeny, ecology, and the parasite species shared within a family did richness of parasite communities in verte not decrease in most cases, they even brates. Ecol Monogr 1995; 65: 283-302 increased when the scattered occurrences Poulin R. Parasite faunas of fr eshwater fish: were omitted, showing that all families the relationship between richness and the harboured parasites specific to the group specificity of parasites. Int J Parasitol that occurred at high prevalences only 1997a; 27: 1091-8 within a fa mily. The fish species that Poulin R. Species richness of parasite shared the maximum number of parasite asseblages: Evolution and patterns. species between them were usually related Annual Reviews of Ecology and Sys species (e.g. among Coregonids, among tematics 1997b; 28: 341-58 Cyprinids) , non-related species that shared similar fo od habits (e.g. Esax lucius and Lata lata) or were a predator and a prey (L. lata and several small fish species). When the scattered occurrences and diplostomids were omitted, the core gonids still shared the maximum number of parasites within the family, but in Cyprinidae the prevalent parasites were shared with several other fish, indicating that they harboured mostly generalist parasites. References Gregory RD, Blackbum TM. Parasite preva lence and host sample size. Parasitology Today 1991; 7: 316-8 Guegan J-F, Kennedy CR. Maximum local helminth parasite community richness in British freshwater fish: a test of the coloniza tion time hypothesis. Parasitology 1993; 106: 91-l00 Holmes JC, Hobbs RP, Leong TS. Populations in perspective: community organization and regulation of parasite populations. In: Esch GW, ed. Regulations of Parasite Populations. Academic Press. New York 1977: 209-45 Leong TS, Holmes JC Communities of meta zoan parasites in open water fishes of Cold Lake, Alberta. J Fish Biol l981; 18: 693-7 13 BullScand Soc Parasito/ 2000; 10 (2): 65-94 SUBMITTED PAPERS- ORAL PRESENTATIONS SIGNIFICANCE OF THE AMOEBO gations of haemolymph cells around sporo CYTE-PRODUCING ORGAN OF BI cysts. Haemocytes from suJTounding tissues OMPHALARIA GLABRA TA SNAILS take part only in such local hemt fo rmation. (STRAINS SELECTED FOR SUS However, real capsules are not fo rmed and CEPTIBILITY/RESISTANCE) IN MS retain their ability to complete their CELLULAR RESPONSE TO ECH� migration to the heart of the snail (Ataev, NOSTOMA CA PR ONI MOTHER 1998). SPOROCYST INFECTION In addition, MS penetration activates APO situated between the pericardium and G.L. Ataevi1, A.A. Dobrovolski , A.V. 1 3 / A vanessiani and C. Coustau mantle epithelium. Therefore, rapid increase 'Department of Zoology, Russian State Peda in the number of haemolymph haemocytes gogical University, St. Petersburg, Russia is observed in snails of both strains after 2Department of Invertebrates, St. Petersburg State approximately one day of infection (Ataev, University, St. Petersburg, Russia Coustau, 1999). But MS encapsulation 3 Laboratoire de Biologie Anirnale, Centre de occurs in a resistant strain at 3-4 days post Biologie et d'Ecologie Tropicale et MeditetTa exposure (PE) only. The result of this is MS neenne, Universite de Perpignan, Perpignan death at 5-7 days PE, by which time APO Cedex, Franc reaches a full development. Haemocytes safely isolate disintegrated sporocyst re The dynamics of haemocyte encapsula mains and dead amoebocytes in the centre tion of Echinostorna caproni mother sporo of the capsule. The decrease in APO activity cysts (MS) in a resistant strain of Biom in resistant snails was noted at 10 days PE. phalaria glabrata snails was investigated After 15 days of infection there will be previously (Ataev, Coustau, 1999). The neither capsules nor agglutinations (haemo result of fmther research was the determina cyte accumulation not observed in MS tion of the role of the snail's amebocyte location). Snails remained viable throughout producing "organ" (APO) in this process the experiment. and the collection of information on the APO stays in active condition in suscep cellular response to sporocyst early devel opment. The first signs of a cellular response tible snails. However, haemocytes do not to Echinostoma caproni MS in susceptible encapsulate MS, and they become adherent and resistant strains of Biomphalaria in agglutinations. glabrata snails were observed several hours The observed variations of resistant after infection. This reaction is usually snails for single species are probably some manifested by insignificant infiltration of of the reasons for local differences of host small parenchymatous lacunas by haemo infection in natural populations. cytes. Sometimes one could see large aggre- 66 THE MAIN FEATURES AND macroparasite infestations correlated with FACTORS LIMITING METAZOAN fish age (body length), fe eding habits (con PARASITE COMMUNITIES IN FISH secutive change of trophic niches) and OF THE BALTIC SEA (LITHUANIAN migration. Macroparasite diversities and COASTAL ZONE) AND CURONIAN abundances within any particular fish LAGOON trophic group were fo und to be similar. The highest degree of parasite diversity and E. Bacevicius infestationwas fo und among benthophagous Fishery Research Laboratmy, Klaipeda,Lithuania fish. We have also observed several variations Parasite communities fo und in fresh and in parasite diversity, abundance and domi brackish water fish species are different in nating pattems of these (prevalence of their origin and structure as well as in their Cucullanelus minutus and Argulus fo liaceus patternsof seasonal and spatial variation. during the summer, glochidians of Anodont I have tried to summarize both literature eds in early spring, and Cucullanus hetero and my own data on brackish and freshwater chous in winter - spring). During summer fish and parasite systems. Fish biology, and autumn iri the sea coastal zone near the population structure, fe eding habits, distt·i estumy of the Curonian Lagoon several bution of fish belonging to different tt·ophic freshwater fish metazom1 parasites (Dige groups(nine each from Baltic andCuronian nean, Diplozoon sp., Ergasilus sieboldi, Ar Lagoon) as well as developmental cycles, gulus fo liaceus, Pradilepus scolecina, Pro population structure and biodiversity of teocephalus sp., Philometrasp., Camallanus parasites and their inte1mediate hosts, were lacustris, etc.) were fo und, but these could analyzed. not complete their entire life cycle in this environment and were counted as individu Both brackish and fresh waters were als eliminated frompar asitic systems. dominated by generalist (nonspecific) fish parasites with developmental cycles includ Neither enzooses nor noticeable pathol ing more than one intermediate stage. ogy to fish organs were observed. All Biodiversity of parasite species in fresh parasites fo und were of univoltic (in brack water was fo und to be higher than in the ish and fresh water) or bivoltic (in fresh brackish Baltic Sea (63/13). In brackish water) developmental types. waters the greatest number of species The main factors limitmg biodiversity of belonged to the fo llowing taxa: Acantho parasites and their intermediate hosts are cephala (5), Nematoda (3) and Cestoda (3), considered to be salinity, temperature, and while in fresh water they belonged to Tre bottom type. matoda (20), Monogenea (20), Cestoda (1 0), Nematoda (6), Acanthocephala (2) and Crustacea (2). Intennediate hosts of fish parasites in fresh water were Bivalvia (3), Gastropoda (5), Insecta (3), Crustacea (3) and Pisces (3), while in brackish water they were Crustacea (3), Gastropoda (2), Oli gochaeta (1) and Pisces (2). Fish with 67 THE COMMUNITY OF MYXO COMPARISON OF IMMUNE SPOREA FROM THE KIDNEY OF RESPONSE MECHANISMS IN THE ROACH RUTIL US RUTIL US RAINBOW TROUT AGAINST LACUSTRIS GYRODACTYLIDS AND CILIATES M.D. Badmaeva\ N.M. Pronin2 and S.V. Pron 1 K. Buchmann, C.V. Nielsen, M. Dalgaard, T. ina Lindenstr01lland A. Li 1Buryat State University,Ulan-Ude, Russia 2 Section of Fish Diseases, Department of Veteri Institute of General and Experimental Biology, nary Microbiology, Royal Veterinary and Agri SB RAS, Ulan-Ude, Russia cultural University, Frederiksberg, Denmark Objective: To investigate the commu Objective: Characterization and com nity of Myxosporea from the kidney of the parison of immune response mechanisms in roach Rutilus rutilus lacustris in the rainbow trout (Oncorhynchus mykiss) Chivirkuyski Bay of Lake Baikal in January fo llowing infections with monogeneans through August 1999. (Gyrodactylus derjavini) or ciliates (lch Materials and Methods: Counting of thyophthirius multifiliis). the myxosporean vegetative stages in the Materials and Methods: Various tissue of the kidney was done according to techniques fo r the study of fish immune Roitman (1997). mechanisms (cellular and humoral) were Results: The kidney of roach is infected applied for characterization of the responses with fo ur species of Myxosporea. The in the fish towards the ectoparasitic Gyro prevalence of infection by Myxidium rhodei dactylus derjavini and the epidermis invad was 98.48 %, Myxobolus ellipsoides 47.94 ing ciliate lchthyophthirius multifiliis. %, My xobolus mulleri 42.52 % and He nne Results: It was fo und that specific anti guya cutanea 9.52 %. These species formed body titres rose significantly fo llowing 16 combinations: single species infections - infection with Ic hthyophthirius multifiliis. In 4, two species - 6, three species - 5, fo ur contrast no binding of specific immu species - 1. noglobulins to the monogenean was de Conclusion: The absolutely predomi tected. The altemative complement cascade nant species in the community of Myxospo was in both cases fo und to affect the para rea from the kidney of Rutilus rutilus lacus sites significantly. Cellular reactions ( oxida tris is M rhodei. The correlation of one, tive burst reactions of macrophages) were two, three and fo ur species combinations also activated by both parasite species. In changes during the whole year and probably addition, putative mastcells (metachromatic depends on seasonal infection dynamics of thionin-positive cells) in the skin were seen some species. to degranulate as a reaction to infection with both ciliates and gyrodactylids. Mucous cell densities in the fish skin were affected differently by the two parasites but activity of lymphocytes may be involved in both reactions. The non-specific reactions may play a part in the host response and it was fo und that fish protected against gyrodac- 68 tylids showed a partial resistance to infec Results: Larvae from pasture and soil tions with lchthyophthirius multifiliis. were recorded on all deposition occasions. Conclusions: The immune response The highest recovery was from the mid third mechanisms involved in the acquired resis of the grazing season, both relative to egg tance of rainbow trout towards lchthyo count and in absolute numbers. Larvaewere phthirius multifiliis is relatively specific also found in soil. compared to the reactions against Gyrodac Conclusion: Faecal contamination tylus derjavini. However, both specific and throughout the grazing period contributes to non-specific elements play a role in the the overwintered larval population under response towards both species. Swedish conditions. However contamina tion early in the previous season is relatively more important in contributing to these PARASITIC GASTROENTERITIS OF larval numbers. Soil can also act as a signifi CATTLE IN SWEDEN: IMPOR cant reservoir. Because the period of snow TANCE OF THE ECOLOGY OF THE cover was limited and transient during the FREE-LIVING STAGES period of this study, the purported "buffer S.O. Dimander, J. Hi.iglundand P.J. Waller ing" effect fi"om extended snow cover, SWEPAR, SLU/SVA, Uppsala, Sweden resulting in enhanced survival of parasitic free living stages, could not be ascertained. Objective: Typically throughout Europe, Acknowledgement: This work was fi overwintered populations of free-living nancially supported by the Swedish Council stages of nematode parasites of cattle are for Forestry and Agricultural Resear·ch small. However, these are a very important (SJFR). source of infection to animals fo llowing turnout, whereas the fo llowing generations are responsible fo r production loss and HOST SPECIFICITY AND clinical disease. In Sweden, outbreaks of DISPERSAL STRATEGY OF clinical parasitism have been reported within GYRODACTYLIDS IN THE BLACK one month fo llowing turnout. What is the SEA COASTAL BIOCOENOSIS reason for this difference? This study was aimed at providing information on this E. Dmitrieva Department of Ecological Parasitology, important ecological question. Institute of Biology of the Southern Seas, Materials and Methods: A pasture plot Sevastopol, Ukraine study was conducted, whereby 12 artificial 400g dung pats derived from naturally Objective: Two parasite systems were parasitized young cattle, were deposited on studied: Gyrodactylus sp. 2 (G.s.2) with one seven occasions at three-weekly intervals host species (Blennius sphinx (B.s. )), at1d throughout the 1997 grazing season. Half Gyrodactylus sp. 1 ( G.s. 1) with eleven host the pats were protected from snow cover. species (excluding B.s. ). The fo llowing spring, pasture and soil Aims: 1 ). To show the role ofhost speci samples were taken fo r infective larval ficity of two gyrodactylid species and their recovery from a 28 cm circular area centred occurrence in the environment; 2). To deter- over the deposited dung pats. 69 mine the mechanisms and means of disper IS OSPORA (PROTOZOA, SPORO sal of gyrodactylidsin the biocoenosis. ZOA) INFECTION IN PASSERINE Results: The data were obtained asresult BIRDS OF VARIOUS FEEDING of laboratmy experiments. Individuals of HABITS G. s.2 were detached during several hours O.V. Dolnik after they were transferred to fish species Zoological Institute, Russian Academy of Sci other than B.s. Individuals of G.s.l survived, ences, St. Petersburg, Russia but did not reproduce, during 5-l 0 days after they were transferred to B.s and other fish Objective: Is ospora infects a new host species fi·om their natural host range. All via occasional swallowing of oocysts that detached specimens of G.s.2 were fo und at appear in bird's faeces. Thus, the diet of the the bottom and on the walls of the aquarium, bird can influence its chances to become in contrast to G. s.l which preferred macro infected by Is ospora. phytes (80 % - on macrophytes and 20 % - Materials and Methods: In total 1131 at the bottom). 72 % of G. s.2 specimens birds of 4 7 passerine species were trappedin transferred to uninfected hosts had function summer and autumn after 2 p.m. at the ing male reproductive systems and 28% had Baltic Sea coast (Curonian Spit) and the also empty uteri. Taking into consideration North Sea (Helgoland Island). Faecal sam that the worms belonging to this ontogenetic ples of every bird were examined for Is o stage account fo r only 5% of a gyrodactylid sp ora oocysts. The number of oocysts in population (Harris, 1985), their high propor every faecal sample was counted after tion among migrating gyrodactylids shows flotation centrifuging by standard method. that such specimens migratemore actively. Bird species were divided into fo ur groups, Conclusion: Gyrodactylid transmission according to their fe eding habits. is realized mainly by specimens with func Results:Both prevalence of infection (% tioning male systems, full fo rmation of of infected birds) and infection rate which may be a migration behavioural (C=number of oocysts per sample) increase trigger. Specimens of G. s.2 infect suitable together: birds catching insects in the air hosts due to genetic specificity and host (5%, C=3), collecting insects fi·om leafs and behaviour. Detached worms move on the sprouts (55%, C=l 47), including berries into substrate, which conforms to the behaviour diet (72%, C=730), and fe eding on the of hosts preferring to lie on parts of substrate ground (76%, C=3879). The difference without macrophytes. Specimens of G.s.l between the first and the last fe eding group can apparently use a wide range of transport is significant. hosts and macrophytes fo r transfer. Conclusions: The probability of trans fe rring oocysts is reduced if the bird catches insects in the air, but picking insects from leaves increases it. Berries attract many birds which get a chance to swallow oocysts in faeces that cover berries. Feeding on humid ground gives an oppmtunity fo r Is ospora infection because bird droppings 70 accumulate there and oocysts can survive Conclusion: It is obvious that the char there longer. acter of change of the community does not depend on the type of polluting substance. It PARASITE COMMUNITY STRUC is determined by the level of pollutant TURE AS AN INDEX OF ENVIRON influence on the resetvoir and the degree of MENTAL CONDITIONS biocoenosis degradation. G.N. Dorovskikh Syktyvkar State University, Syktyvkar, Russia P ARASITOLOGICAL MONI TORING IN EVACUATION ZONE NEAR CHERNOBYL NPP Objective: Parasites create a community by taking part in certain relationships, G. Efremova including quantitative. That the existence of Institute of Zoology, National Academy of a certain parasite community stmcture is Sciences, Minsk, Belarus detennined by the biomass of specific species was shown fo r Phoxinus phoxinus. Objective: Parasitological monitoring in Does this structure remain the same in the evacuation zone near Chemobyl Nuclear individual reservoirs and how does it react Power Plant was undettaken fo r the purpose to environmental conditions? of improvement of sanitaryconditions in the Materials and Methods: The present territory ofPolessky radio-ecological resetve work has been perfmmed on the basis of the and of the state of health of the staffof the parasite fauna of fish from the reservoirs of resetve. the nmth-eastem part of European Russia. Materials and Methods: The parasitic The combination and conditional biomass of arthropods of 697 birds and 367 bird nests species are taken as the characteristics of the were studied. community. Results: Positive dynamics of an in Results: In ecologically pure reservoirs, crease in the numbers of Ix odes ricinus L. and in parts of them, and in the areas which were recorded. The prevalence of tick are not heavily polluted, three groups of infestation of the birds was 1.5-2.0 times parasites have been identifiedin the parasite higher than on the control territmy. A large community of all host species researched. number of parasitic arthropods were fo und The logarithms of their conditional biomass in the nests (ticks fam Dermanyssidae, lie on line segments separated by gaps, and Laelaptidae, fleas, bugs). having different slope angles. As the reser Conclusion: The location of the popu voir is polluted and the biocoenosis de lated area near the evacuation zone in grades, species not typical of the community conjunction with the presence of the fo ci of stmcture appear (points of their conditional tick-borne encephalitis and tick-borne Lyme biomass do not lie on a line segment). Then borreliosis necessitate conducting regular the groups are divided into subgroups, and, annual epidemic controls on this territory. finally, in greatly polluted rese1voirs, the number of groups of parasites decreases from three to two. 71 THE EEL SWIMBLADDER NEMA increased prevalence (15%) and mean TODE ANGUILLICOLA CRASSUS intensity(8.6). (NEMATODA: DRACUNCULOIDEA) Conclusion: A. crassus is firmly estab INIRISH EELS lished in the Erne. Within this catchment it 2 has become more widely distributed by D.W. Evans1 and M.A. Matthews 1 utilizing the migratory behaviour of eels. Faculty of Agriculttrre and Food Science, Queen's University of Belfast, Belfast, Northem Moreover, with the commercial movement Ireland, UK of live eels from the Erne, furtherreports of this parasite within Ireland are anticipated. 2Northem Regional Fisheries Board, Ballyshan non, Ireland FISH ANISAKIDAE IN KHUZESTAN Objectives: The Erne lake system is the PROVINCE OF SOUTH WEST IRAN fo mth largest catchment in Ireland suppmt A. Farahnak ing an impottant commercial eel fishery. Department of Parasitology and Mycology, Anguillicola crassus was first documented School of Public Health, Tehran University of here in 1998 during investigations by the Medical Sciences, Tehran, Iran Erne Eel Enhancement Programme into eel population structures. Analysis has contin Objective: Fishes, including Barbus ued in order to monitor changes in preva spp., Cyprinus carpio, Liza abu, Aspius lence, intensity and parasite distribution. vorax, have a very important role in the Materials and Methods: In 1998 the economic condition of rural areas of swimbladders from 372 eels (328 yel Khuzestan province. These fishes are con low and 44 silver) from various lakes within sumed barbecued or roasted. These inade the Erne catchment were examined fo r quately cooked fishes could serve as a A. crassus . Total worm burdens were source of infection in these communities. recorded. This process was repeated in 1999 Material and Methods: For these rea analysing 658 eels ( 432 yellow and 226 sons 283 fishes were caught from fo ur silver). Lakes examined in 1998 were re lagoons (Hooral Azim, Shadegan, AI Hai, examined in 1999 in addition to several Seid naser) and transported live to a health lakes not surveyed before. research centre and their intestine, body Results: In 1998 prevalence in yellow cavity and muscle examined carefully. eels fo r the catchment was 7.3 % and mean Results: In 20.8% of fishes, infections intensity 4.3. In silver eels prevalence was with Contracaecum sp. (Anisakidae) and/or 4.5 % and mean intensity 2.5. The parasite Rhabdochona sp. (probably R denudata) was not distributed throughout the entire were fo und. These results suggested that catchment but was confmed to several human anisakiosis could be a medical localities in the northern half. In 1999 the problem in this area. range of the parasite had increased, moving fu rther upstream and downstream. Preva lence in yellow eels had risen to 10% and there was an increase in mean intensity to 6.7. Likewise, in silver eels there was an 72 THE ECOLOGY OF IXODIDAE In Novgorod Region before the drainage TICKS IN NOVGOROD FOREST of agricultural fields on vast territories there REGION UNDER THE CONDITION was a stubbing of shrubbety and low forest OF ANTHROPOGENIC LOADING which led to the decrease of biotopes fo r settingof I ricinus ticks and their. V.G. Fyodorova and Z.S. Lalaeva So, anthropogenic fa ctors are the main Novgorod State University, Novgorod Veliky, Russia ones in the change of enviromnent for the setting and activity of Ixodidae ticks. The problem of tick ecology relates to the studying of several different aspects COMPARATIVE ANALYSIS OF THE connected with anthropogenic loading of the MONOGENEAN FAUNAS OF THE environment. Firstly, at the end of the XX BLACK AND BALTIC SEAS centmy global changes in the biosphere, as a result of intensive economic activity, can be A Gaevskaya and E. Dmitrieva clearly seen, and this affects climate change. Department of Ecological Parasitology, Institute It is known (Pomerantsev,l935; ) that the of Biology of the Southem Seas, Ixodidae ticks are rather sensitive to changes Sevastopol, Ukraine in abiotic fa ctors (temperature, soil humidity and fo rest litter). Objective: The specific aim of this re search was to analyse the composition of Anthropogenic intetference in natural monogenean faunas fi·om two different seas complexes, especially industrial deforesta having similar hydrochemical conditions, tion, shmbbety, low fo rest stubbing, etc., namely the Black and Baltic Seas. improve natural landscapes. The change of forest biocoenosis into clarified agrocoeno Materials and Methods: Authors' own sis is the main fe ature in changing the and literature data on monogenean faunas ecological situation fo r Ixodidae ticks. fromthe Black and Baltic Seas were used. Accumulation of heavy metals in soil, Results: The monogenean fauna of the nitrogen through the use of mineral fettiliz Black Sea (38 species) is much richer than ers, electromagnetic radiation, heavy traffic that from the Baltic Sea (I 5). The latter and other factors influence the physiology, fa una is mainly represented by boreal the survival rate and the numbers of ticks. species and has no endemic species. The Black Sea monogenean fauna is represented During the last three years in Novgorod by Mediterranean invaders (30%), boreal Region, experimental investigations of tick Atlantic species (30%), endemic species (Ix odes persulcatus) populations were (20%) and includes small groups of cosmo carried out in fo rest areas treated with politan species and ponto-caspian relicts. mineral fertilizers in the village of Lesnaya, Only five from 15 Black Sea species which with simultaneous control in the identical occur in nmthem latitudes are also fo und in biotopes of the village of Golino. The the Baltic Sea: Nitzschia sturionis, Gyrodac experiments were also held in laboratories. tylus arcuatus, pungitii, jlesi and We considered that mineral fertilizers could G. G. G. harengi. The first-named species is a spe influence the development and the survival cific parasite of sturgeons and penetrates rate oflarvae of Ix odes persulcatus. into the seas with their hosts from river 73 systems. The fo llowing two species are between the local abundance of a species specific parasites of sticklebacks and appar and its regional distribution. ently occur over all the distribution of their Materials and Methods: 122 chubs hosts. Despite the wide distribution of the (Leuciscus cephalus) were caught by back fish host of G. jlesz; namely Platichthys pack electrofishing from March to October jlesus, this monogenean species was fo und 1998 in 13 water courses (northern Italy). only in the Baltic, White and Black Seas. G. Parasites were collected from gut, eyes and harengi parasitizes the herring in the Baltic, gills and identified to species level. The White and Black Seas and in the northern level of parasitic infestation was analysed part of the Pacific Ocean. According to for each species, using prevalence, intensity, Malmberg (1970), three subspecies of mean intensityand abundance. Clupea harengus are parasitized with three Results: 12 parasite species were identi diffe rent species of Gyrodactylus. G. jlesi fied: Plathyhelminthes: Dactylogyrus and G. harengi apparently have penetrated into the Black Sea during river floods in the vistulae, Paradiplozoon ergensi, Al anthropogenic period. The last 10 species locreadium isoporum, Bucephaluspol ymor phus, Diplostomum spathaceum, Ty lodel are typical marine monogeneans, the distri bution of which is restricted to a salinity of phys clavata, Proteocephalus torulosus; not less than 15%o. Nematoda: Rhabdochona denudata; Acan thocephala: Acanthocephalus anguillae and The present status of the Baltic Sea was Pomphorhynchus laevis; Crustacea: Lam fo rmed approximately four thousand years proglena pulehe !la; Mollusca: glochidia. ago, thus it is only half as old as the Black Sea. Its fauna can be enriched as was that of Mean parasite abundance, log trans the Black Sea, which continues to be ex fo rmed, and number of infected fish shown tended by species from the Mediterranean a significant correlation (Speatman rank Sea. correlation r-0.66, p P. Galli and G. Crosa Department of Landscape Environmental Sci ences, Universita degli Studi di Milano-Bicocca, Milano, Italy Objective: "Core and satellite" hypothe sis was tested on fish parasite populations according to the empirical relationship 74 DIVERSITY OF FISH PARASITES COMMUNITYECOLOGY OF THREE IN RELATION TO ENVIRON DIFFERENT MONOGENEAN PARA MENTAL STRESS SITE-HOST SYSTEMS 1 M. Gelnar, S. Sebelova, L. Dusek, J. Jarkovsk)l, M. Gelnar1, I. Matejusova1, A. Simkova AND S. 2 L. Machala, B. Koubkova and P. Jurajda Morand 1 Department of Zoology and Ecology, Masmyk Department of Zoology and Ecology, Masmyk University, Bmo, Czech Republic University, Bmo, Czech Republic 2Centre de Biologie et d'Ecologie Tropicale et Objective: A comparison of three differ Medite1raneenne, Universite de Perpignan, ent levels of parasite-host systems (e.g. Perpignan Cedex, France parasite individual, population and commu nity) has been perfonnedto assess the use of Objective: Three different monogenean parasites as indicators of environmental parasite-host systems were investigated to stress. analyse aggregation, coexistence and com petition among potentially competing Material and Methods: The current in parasites. fracommunity and component community approach using different biodiversity meas Material and Methods: Current meth ures (e.g. species abundance models, Q - ods of ecological and evolutionary parasi statistics) of various parasite assemblages tology (e.g. nestedness analysis, aggregation was used. model of coexistence, parsimony analysis) were used. Results: An apparent decrease in para site species richness and an increase in the Results: A nested pattem was recorded homogeneity of parasite distribution within in parasite assemblages of minnow, the host the population of the model host fish (Leu with a dominant number of specialist gyro ciscus cephalus) was found, corresponding dactylids. A non-nested pattern was ob to the levels of contamination of the river seJved in gyrodactylid assemblages of roach, bottom sediment and the bioaccumulation of the host with a dominant number of general selected pollutants in individual host fish. ists. Comparative analysis of dactylogyrid coexistence showed that parasite abundance Conclusion: This contribution repre is positively correlated with niche breadth. sents an attempt to compare and analyse, on A parsimony analysis of the evolution of the a precise current statistical basis, the rela parasite gill distribution indicates a change tionships among composition and structure in one parameter of the niche at each of parasite communities (infracommunity branching event. Inter- and intra-specific and component community levels) with competition between two monogeneans, respect to the level of bioaccumulation of Eudiplozoon nipponicum and Dactylogyrus pollutants by individual fish host and to the extensus, parasitic on the gills of carp ( Cy level of contamination of river bottom prinus carpio ), were investigeted. sediments. Conclusion: There is no doubt that monogenean parasites represent a perspec tive model fo r investigation of parasite aggregation, competition and coexistence. �--�----�------ 75 SPECIES LIFE CYCLE, DEVELOP- accurately. It can be distinguished as the MENTAL CYCLE AND LIFE way of LC realization in specific environ SCHEME IN PARASITOLOGY ments. Thus the life scheme is the ecological (DEFINITION AND CORRELATION aspect of the LC understood as the complex OF TERMS) of all life fo rms constituting it. This aspect is pa.tiicularly a subject of L.A. Ghitchenok parasitology. Biological Faculty, Moscow State University, Moscow, Russia A STUDY OF THE PARASITE FAUNA Life cycle (LC) is the term definedas the OF PHOXINUS PHOXINUS L. FROM system of adaptations ensuring continuous THE SMALL OF species existence in space and time and RIVERS NORTHERNDVINA BASIN applicable only to the species as a whole. The LC is based on the principle of "repro E. Golocova duction of similar not directly, but through a Syktyvkar State University, Syktyvkar, Russia more or less long chain of non-similarity". In other words, the LC is the sequence of Objective: The parasite fauna of Phox events which are necessary and sufficient inus phoxinus L. from small rivers of the fo r species preservation. Thus realization of Northern Dvina basin and seasonal dynam the LC is the form of species existence, and ics of infection were investigated. the notion of the LC proper is basic and Materials and Methods: Two hundred universal, at least fo r all eukaryotes includ and fifty-seven fish specimens were caught ing parasitic species. from the small rivers of Vychegda basin in Life cycles of exact species could be ex the vicinity of Syktyvkar in 1996-1999. The amined through two aspects. examination of material was carried out by The morphogenetic aspect reflects the the method of total parasitological dissec realization of a genetically determined tion. consecutive developmental programme Results: Twenty-two parasite species peculiar to this species; that is morphogene belonging to six classes were recovered sis as such, independently from the exact from P.phoxinus during the research period. environment of its realization. The fo llowing species were found fo r the The term developmental cycle corre first time in P. phoxinus fl-om the Northern sponds to the essence of this aspect (the Dvina River basin: Apiosoma piscicolum morphogenetic aspect of the LC), recog ssp. perci, lvfyxidium rhodei, My xobolus nized as the ontogenetic complex of all muelleri, M musculi, M albovae, Dac generations that compose the species LC. tylogyrus borealis, Pellucidhaptor merus, Gyrodactylus magnifzcus, G. minimus, G. This aspect is not a subject of parasitol laevis. It was shown that the minnow para ogy asan ecological discipline. site fauna from the River Chovju (20 spe The ecological aspect reflects the adap cies) and from the River Kyliymju (18 tion of the LC to the exact environment. The species) differ only in their rarely occurring notion of the species life scheme (Beklem species. The parasite fa una of P. phoxinus is ishev, 1957) corresponds to this aspect most considerably impoverished in the River 76 Dyrnos (eight species). Seasonal changes in is only observed as an exception in shmt the infection rate in the River Chovju and lived species, which generally do not change River Kyltymj u showed a general regularity: growth rate when parasitized - Basommato the majorityof parasites increase in numbers phora (Pulmonata). The only trend mani during May through June, and common fested in infected long-lived species (6-27 fe atures are determined by a number of yr) - Littorinacea and Cerithiacea (Proso fa ctors: hydrological, hydrobiological and branchia) - is a decrease in growth rate. The anthropogenic. morpho-functional distinctions in the sys Conclusion: The available data allow us tems fmmed by n·ematodes and molluscs to conclude that the River Chovju and River from different taxa are expressed in the Kyltymj u are subject to anthropogenic pattern of parthenitae localization in the host eutrophication, and the River Dyrnos is body and in the degree of digestive gland characterized by an extremely high rate of and/or gonad destruction. pollution,. Conclusions: Snail species from differ ent taxa are not even in terms of the living space «granted» to the parasite without risk EXPLOITATION OF GASTROPODS of host viability reduction. Hypothetically, BY TREMATODES: HOST MOR an «equilibrium» exploitation sn·ategy is PHOLOGY AND PARASITE favoured in long-lived hosts which may be STRATEGY viewed as a stable environment with low A. M. Gorbushin variation in habitat quality. Trematodes Department of Cytology and Histology, St. parasiting shorter lived hosts may use an Petersburg State University, St. Petersburg, «oppmtunistic» strategy which maximizes Russia the exploiting capability in environments that change frequently on a relatively small Objective: Morpho-functional patterns of temporal scale. pathogenesis in various mollusc - trematode combinations are very different. To evaluate the level of antagonism between the host PARASITIC SYSTEMS OF TREMA and parasite it is reasonable to use a summa TODES AND THE MECHANISMS OF rized parameter of the general physiological THEIRREGULATION state of the infected individual. AI. Granovitch Materials and Methods: Analysis of host Department of Invertebrate Zoology, growth rate fo llowing infection and patterns St.Petersbmg State University, St. Petersbmg, of histopathological modification was used Russia in this study. A comparative analysis was conducted of original and literature data on A parasitic system is a complex of popu 40 snail - trematode combinations. lation systems of hosts, united by the popu Results: Growth response to infection lation system of the parasite on the basis of depends on the life history of the mollusc. stable parasite-host community relation Growth acceleration (gigantism) is common ships. The structures of parasitic systems are fo r snails with intermediate longevity (3-4 highly variable. However there are two yr) - Rissoacea (Prosobranchia). Gigantism general aspects of their organization: a) 77 depending on the specific life cycle peculi POPULATION DYNAMICS OF arities of the parasite; b) depending on the ARGULUS COREGONI THORELL environmental subdivision of the parasite. (CRUSTACEA: BRANCHIURA) Correspondingly, we can separate meta- and para- aspects of the structure of a parasitic T. Hakalathi and E.T. Valtonen system. Department of Environmental and Biological Science, University of Jyvaskyla, Jyvaskyla, Mechanisms which lead to the stability Finland of parasitic systems of trematodes were considered. The investigation was cruTied Argulus coregoni is a free-swimming, out in two regions of the White Sea (Kan obligate ectoparasite of fishes. After copula dalaksha Bay, Russia) and one of the North tion they leave their fish hosts in order to Sea (Skagerrak, Sweden). Populations of deposit eggs on hard substrata. Ectoparasites marine littoral molluscs and their trematode from boreal regions face a severe and long parasites were examined. winter when their populations are sup The specific regulatory mechanisms in pressed. It has been suggested that Argulus the parasitic systems may be considered on eggs can rest throughout winter and hatch two levels at least (of thesystems of differ next summer having 2-3 generations in a ent «scale»). year. 1. The level of host and parasite popula Objectives: To describe A. coregoni tions. The population of separate mollusc population rhythms in a fish farm during species interacts with the hemipopulation summer: number of generations, hatching group of trematodes (i.e. individuals of one rhythms, hatching duration and the influence stage of their life cycle). Severe prevalence of dtyingand temperature on hatching. may deform various ecological, behavioural Materials and Methods: The size and and demographic characteristics of the sex structure of the A. coregoni population mollusc population. However, the persis was monitored weekly in a fish farm in tence of the host population under condi Central Finland during the open water tions of heavy infection suggests the pres period in 1999. Overwintering eggs were ence of mechanisms that compensate fo r the collected in September and kept at a steady influence of the parasites. temperature in the laboratory to study their 2. The level of the parasitic system. The hatching dyna111ics afterdiff erent treatments. distribution of the trematode hemipopulation Results: Results indicate that A. core over several host species (paraxenia) ap goni overwinter as eggs and there might be pears to be ru1 important possibility fo r only one generation in a year. However the regulation of the parasitic system (paraxenic hatching of eggs is very prolonged. Drying regulation). Other essential mechanisms at in all treatment combinations and freezing at this level are metaxenic regulations of the -18°C fo r 12 days killed all the A. coregoni parasitic system. That is the possibility to eggs. After other treatments ( + 4 °C, + 1 0°C change the «flow» of parasites through the and normal winter temperature with fluctua successive (metaxenic) hosts, which are tions) juveniles were still able to hatch. The used by successive stages of the life cycle of stones with A. coregoni eggs produced the parasite. 78 juveniles continuously from November to Results: All herds, except one, were in March. fected with D. viviparus at some time during Conclusion: After the ice cover period, 1999. The overall seroprevalence was 8.3% ove1wintered eggs will hatch synchronously. (101/1212). Elevated levels of antibodies First A. coregoni females mature and lay were identified in 4.6% (28/604) and 12.0% eggs in late June in Finland. During the (73/608) of the serum samples at turnoutand summer newly hatched juveniles appear housing, respectively. On the infected farms, fi·om overwintered eggs continuously as seroprevalences varied between 0% - 15.7% new females mature and lay eggs. We at turnoutand 5.2% - 24.6% at housing. The suggest that eggs, laid at any time in the seroprevalence in calves < 1 year and in summer, will hatch only afterwinter in the older cattle >2 years was 20.3% and 6.0%, fo llowing summer. This is why we suggest respectively. No clinical signs of disease the occurrence of only one generation with a were observed. long annual recruitment time. This however Conclusion: Although the highest sera needs experimental verification. prevalence was noted at housing, some animals were also infected at turnout. Despite that there was a significantly greater A SEROEPIDEMIOLOGICAL SUR propmtion of calves than older cattle in VEY OF DICTYOCA UL US VIVIPA RUS, fected, and seropositive animals were fo und LUNGWORM INFECTION OF in all age groups. These results suggest that CATTLE IN SWEDEN arrested larvae and older cattle as a means of J. Hoglund, S.O. Dimander and P.J. Wallere survival over-winter may play impmtant SWEPAR, SLU/SVA, Uppsala, Sweden roles in the epidemiology of lungworm infection in Sweden. Objective: To investigate the seropreva Acknowledgement: This work was fi lence of Dictyocaulus viviparus infection in nancially supported by the Swedish Farmers Swedish cattle, with particular emphasis on Foundation for Agricultural Research (SLF) the role of adult cattle as silent carriers. and Swedish Meats. Material and Methods: This survey was carried out in 1999 in 10 herds which ECOLOGICAL CHARACTERISTICS had a history of dictyocaulosis in 1998. OF FISH PARASITE FAUNAS IN Accordingly, some animals on each of the NORTH-EASTERN AZERBAIJAN farms were treated either with ivermectin, morantel or febantel. Neve1theless, all cattle RIVERS that had been on pasture during 1998 were S.R. Ibrahimov and B.S. Agayeva sampled at the beginning (turnout) and end Institute of Zoology, Azerbaijan Academy of (housing) of the grazing period in 1999. Sciences, Balm, Azerbaijan From each fa rm, blood samples were col Objective: To defme the ecological lected from 23 - 114 animals, serum was characteristics of the fish parasite faunas in obtained and specific levels of serum anti north-easternAzerbai jan rivers. bodies to Dictyocaulus viviparus were Material and Methods: In 1994-98, 657 measured with a commercially available fishes belonging to 24 species were exam ELlSA. ined by total parasitological dissection 79 method in north-eastern Azerbaijan rivers: method consist of a dominant peak in Samur, Gusarchay, Gudialchay, Deve activity in spring and a minor peak in au chichay and Gilgilchay. tumn. Flagging, however, does not reveal Results: 121 fish parasite species were any detailson whether the seasonal variation fo und in the north-easternAzerbaij an rivers. in activity originates from variation in population size or activityof the tick popula Conclusion: Only four species of ich tion. It is thus unknown whether the differ thyoparasites are typically rheophilous, ences observed by flagging vegetation are a while the other 117 species are limnophi measure of population size or activity of the lous. The majority ofthe recovered parasites individual ticks. The purpose of this study are typical freshwater inhabitants, some are was to obtain information on how well the emyhaline, and 10 species are marine. All peaks in activityrelate to population size. marine parasites were noted in marinefish and migratmy fish only. The typically Material and Methods: Ticks were col marine Corynosoma caspicum infects all lected by flagging vegetation for three species of marine and migratory fishes, and consecutive days on eight occasions. it can be a natural indicator, petmitting Results: The first day's flagging showed differentiation of typically fi·eshwater and a bimodal pattern, with a peak in abundance migratory populations of fishes. of nymphs in June and September which The predatmy fishes accumulate 13 resembles traditional flagging results. helminth species. The plankton-eating fishes Results from the third day's flagging did not were infected by six helminth species, latvae show a clear bimodal pattem. The hostseek of which live in planktonic invertebrates. ing period as defmed by the ratio between The benthophagous fishes have 12 helminth first and third day samples appem·ed almost species connected with the benthos, but they exclusively to be responsible for the bimodal also have 12 helminth species com1ected pattern of activity in first day samples. with the plankton because some copepods Conclusion: The variation in hostseek inhabit both the water column and bottom ing periods may have profound influence on areas. the observed abundance and seasonality of ticks. Consecutive flagging appears to be a convenient method of obtaining infotmation CONSECUTIVE FLAGGING on the hostseeking period and ascending SAMPLES - A SIMPLE METHOD TO numbers of ticks. The latter may possibly be OBTAIN DETAILED INFORMATION closely related to population size. ON THE HOSTSEEKING ACTIVITY OF IXODES RICINUS P.M. Jensen Zoology Section, Department for Ecology, Royal Veterinary and Agricultural University, Frederiksberg, Denmark Objective: Observations on tick activity are mostly based on flagging satnples. The typical patterns of tick activity by this 80 VARIABILITY OF THE PARASITE has an effect on parasite fauna. As fish FAUNA OF OCEANIC FISH SPECIES mature, most of them begin spawning (Irish DURING DIFFERENT PERIODS OF A shelf), feeding and wintering (the Norwe MIGRATION CYCLE (BY THE gian Sea) migrations. However, there are EXAMPLE OF BLUE WIDTING OF also migrants with delayed gonadal devel THENORTH WEST ATLANTIC) opment, as well as specimens missing spawning. In the north of the distribution AB. Karasev area pseudopopulations of never-spawning Polar Research Institute of Marine Fisheries and fish are formed. In different periods of the Oceanography, Munnansk, Russia migration cycle fish stayeither in the centres Objective: The parasite fa una of blue of parasite infestation, or spatially fa r from whiting Micromesistius poutassou (Risso) them. was studied in different intrapopulation host Conclusion: On the basis of parasi groups. tological data using indicator parasites, the Materials and Methods: Parasitological relationships between intrapopulation and ichthyological samples from blue groupings of blue whiting were mapped. wh iting of the Hebrido-Norwegian popula GENETIC HETEROGENEITY OF tion taken in the area from the Irish shelf to TREMATODE SPECIES REFLECTS Spitsbergen during 10 years were analysed. THE STRUCTURE OF THEIR 1210 fishes were examined by the total NATURAL POPULATIONS parasitological dissection method. Incom K.V. Khalturin1, N.A. Mikhailova2 and AI. plete dissection of 15 000 specimens was 3 pe1fonned. The material encompassed Granovitch 1Zoological Institute, University of Kiel, Kiel, different intrapopulation groupings of blue Ge1many whiting at the ages of 1-16 years. 2Institute of Cytology, Russian Academy of Results: In the distribution area of blue Sciences , St. Petersburg, Russia 3 whiting 37 parasite species of 10 taxawere Department of Invertebrate Zoology, St. Peters recorded. Ecological analysis revealed the burg State University,St. Petersburg, Russia characteristic fe atures of tropho-parasitic Objective: We studied two sibling tre relations of the host, typical fe atures of age matode species Microphallus piriformes and and area-related variability of infestation on pygmaeus belonging to the Microphalli the spawning, fe eding and wintering M dae family. About 400 individual sporocysts grounds of young and mature fish. Indicator (containing metacercariae) infesting Lit parasites - microsporidians Pleistophora sp. farina sa:xatilis and L. littorea periwinkles and trematodes Bucephaloides gracilescens were used fo r the analysis. All individuals etc. - were used to trace the formation of were collected in the Chupa Bay of the peculiar fe atures of the parasite fauna in the White Sea and the largest distance between intrapopulation groupings. The distribution the analyzed localities was 20 of immature fish is related to the dispersal of km. larvae and fry by currents. In different parts Methods: We used Chelex 100 method of the distribution area (the North, Norwe to extract total DNA from individual trema gian and Barents Seas, Iceland) fish live in todes. This extraction has earlier been different environmental conditions, which proven to be useful fo r very small sample 81 sizes and we successfully scored individual THE EFFECT OF HOST DENSITY ON sporocysts of both species. TI1e random ECTOPARASITEDISTRIBUTION amplified polymorphic DNA (RAPD) 1 2 technique was applied for the studies of B. Krasnov and I. Khokhlova 1Ramon Science Center, Jacob Blaustein Institute genetic heterogeneity. for Desert Research, Ben-Gurion University of Results: Amplification of the total DNA, the Negev, Mizpe Ramon, Israel extracted from the whole sporocysts, never 2Wyler Department of D1yland Agriculture, Ben showed any differences in RAPD patterns Gurion University of the Negev, Beer-Sheva, between the parasites derived from one Israel infested snail (local parasite hemipopula tion). That allowed us to compare different Objective: We studied the pattern of parasite populations, referring to the RAPD parasitism of the fleas Xe nopsylla dipodilli pattern of one sporocyst from a snail as (mainly spring and summer flea) and representative of one local hemipopulation. Nosopsyllus iranus theodori (winter flea) on a small rodent, Gerbillus dasyurus, in The data showed considerable genetic different habitats of the Negev Desert, Israel. differences between the subpopulations of We predicted that the intensity of flea M piriformes infesting different paraxenic infestation would increase with the increase intermediate hosts -L. saxatilis and L. obtu of the host density curvilinearly to a plateau sata - which were statistically significant fo r that will be attained at the level of the host two out of three localities studied. No such densitythat corresponds to the abundance of heterogeneity within populations was recor resident individuals (those having their own ded fo r M pygmaeus. Moreover, both M bunows). We predicted also that the preva piriformes and M pygmaeus are character lence of flea infestation plotted against host ized by the genetic differentiation on the densitywould be hump-shaped. microgeographic scale. Accordingto the fre quencies of the RAPD-loci, parasites from Material and Methods: We captured the sheltered locality differ significantly rodents on 20 one-hectare plots and col fi·om the parasites of the other two localities lected fleas fi·om them. In total, 657 gerbils exposed to the open sea. For both species were captured and 1265 fleas were col the degree of genetic similarity between the lected. populations correlates positively with the Results: The distributions of both fleas distance between the localities. on G. dasyurus confotm to the negative Conclusions: Genetic population struc binomial distribution. The intensity of ture of microphallids is dependent on the infestation by both fleas increased with the population structures of their intermediate increased gerbil density to a plateau. How and defmitive hosts and the geographical ever, the threshold host density for this structure of the area. Because of low motil plateau coincided with the number of resi dent individuals fo r X dipodilli and with ity of snails, we believe that the distribution ' migration and species composition of the overall host density fo r N i. theodori. definitive hosts play the key role in the Prevalence of infestation by X dipodilli genetic structuring of M pygmaeus and M plotted against host density was hump piriformes hemipopulations. shaped, whereas that for N i. theodori 82 increased linearly with increase in host Europe and North America. The disappem· density. ance of heavily infected parr (age < 3+) Conclusion: The habitat patch of a flea during the winter in the upper part of the should be considered not only as a pmiicular river indicates parasite dependent mortality. host, but rather as a particular host with a Possible spreading mechanisms of the particulm· burrow or nest in a pmiicular parasite include attraction of the fm al hosts habitat. The flea abundance parameters are (seagulls) by dead fish in the effluent water influenced by the density of the resident fromthe turbines. hosts rather than by the overall host density. INTERRELATIONSHIPS OF CRABS, MORTALITY OF ATLANTIC LEECHES, FISH AND TRYPA SALMON PARR DUE TO IDGH NOSOMES IN COASTAL WATERS INTENSITIES OF EYEFLUKES OF NORTH NORWAY K. Mackenzie1, W. Hemmingsen2 and P.A. T. Larsen and F.R. Lund 3 Finnmark College, Alta,N01way Jansen 1Depmiment of Zoology, Universityof Aberdeen, Objective: The density of Atlantic Aberdeen, Scotland, UK 2Institute of Biology, Depmiment of Ecology and salmon parr in the upper pati of the Alta Zoology, Universityof Troms0, Troms0, Norway river has declined 90 % since the regulation 3Institute of Zoology, Faculty of Cherrtistry and of the river fo r hydroelectric power produc Biology, NTNU, Trondheim, N01way tion in 1987. No changes in density were detected in the lower part. Preliminmy The red king crab, Paralithodes studies have demonstrated high intensities of camstchaticus, was introduced to the Bm· eyeflukes in the Atlantic salmon patr of the ents Sea fi·om its native North Pacific by river. The aim of the present study was to Russian scientists in the 1960s and 1970s. A investigate whether the decline in the den thriving population now established in the sity of Atlantic salmon patT in the upper pati southern Bm·ents Sea is steadily spreading of the river are due to high intensities of westwm·ds across the coast of Finnmark. eyeflukes, and whether the high intensities The arctic leech, Jo hannsonia arctica, lays of eyeflukes are connected with the regula its eggs on the cm·apace of the crabs and is a tion of the river. vector for the haematozoan T!ypanosoma Materials and Methods: Atlantic murmanensis, which it transmits to cod m1d salmon parr were collected by electrofish other fish while fe eding on the blood of the ing. Metacercm·ia of Ty lodelphys clavata fish. and Diplostomum sp. (eyeflukes) were In October 1999 we investigated the oc registered in each year class in autumn and currence of trypm1osomes in cod caught at spnng. 18 stations along the coast ofFinnmark. We Results: The intensity of eyeflukes in fo und two fo ci of infection: one in Va Atlantic salmon patT of the Alta river was rangerfjord in easternFi nnmark where there more that ten timeshigher than that recorded is a dense population of king crabs, and one in neighbouring rivers and what is reported in western Finnmark where there are no king from Atlantic salmon patr in other rivers in crabs. Station prevalences varied from 0 to 83 96%, with both extremes occurring in main parts: ectosoma, located in the external western Finnmark. The heaviest infection environment (environment of the first order) was found in an area where native spider and endosoma, located inside the organism crabs Lithodes maja are known to be par of the host (environment of the second ticularly abundant. We discuss the interrela order). tionships of crabs, leeches, fish and trypano 2. Functional subdivision of the body of somes in the study area, including the the parasite into two parts: endosoma, which possibilitythat we may be dealing with two performsthe trophicfunction, and ectosoma, species of trypanosome with different leech with genital organs performing the repro vectors. ductive function 3. Concurrent existence in the environ NOTES ON MESOPARASITISM ments of the first and the second order, each of the environments influencing the parasite A. Marchenkov immediately. Department of Parasitic Wonns, Zoological Institute, Russian Academy of Sciences, St. Up to now the typical mesoparasites Petersburg, Russia have been only found among the parasitic copepods. In parasitology it is accepted that para sites can be subdivided into two categories traditionally based on their spatial relations PATTERNS OF SIMILARITY IN THE with the host organism. Ectoparasites, i.e. PARASITE FAUNAS OF LARUS organisms which live «on external covers, GULLS skin, gills», are placed in the first category. J. Mellen Endoparasites, i.e. organisms living «in Biology Department, Bemidji State internal cavities, tissues and cells of the University, Bemidji, Minnesota, USA host» are placed in the second categmy (Dogiel, 1941). Objective: The repotted parasite fauna Feizullaev (1971) offered a third cate of Larus gulls was reviewed for complete gory - mesoparasites - where he places «the ness, similarity, and evidence of geographic parasites inhabiting localities which are patterns. connected with cavities opening to the Materials and Methods: Analysis was external environment>>. According to the conducted on reported parasites for 21 above author such places are oral, nasal, eye species of Larus gulls with two or more cavity, cloaca, etc. reports recorded in the Index Catalog of Sincesome parasitic copepods living in a Medical and Veterinary Zoology (USDA), certain kind of spatial relation with the host and Helminthological Abstracts. The influ can be referred neither to ectoparasites nor ence of the size of host breeding range to endoparasites, the author offers the (drawn on Fuller-projections) and number of fo llowing three criteria fo r the defmition of studies reported on parasite species richness mesoparasitism. of the compound fauna of 21 gull species was evaluated by pattial correlation analysis. 1. Morphological and anatomic subdivi Similarity analysis was conducted using sion of the body of the parasite into two 84 Simpson's and Jaccard's coefficients and HOW DOES ARGULUS FOLIA CEUS L. UMPGA clustering. (CRUSTACEA: BRANCHIURA) Results: Host breeding range was not MATCH WITH ITS HOSTS AT THE significantly correlated with the species BEHAVIOURAL AND ECOLOGICAL riclmess of parasite fauna. However, the LEVELS? number of published studies was statistically V. Mikheev1, A. Pastern� and E.T. Valtonen3 significant. Simpson's coefficient generated 1Laboratory of Behaviour of Lower Vertebrates, similarity patterns of parasite faunae that Institute of Ecology and Evolution, RAS, Mos were highly consistent with tl1e bio cow, Russia geographic realms associated with the host 2Laboratory of Plankton Ecology, Institute of breeding range. Oceanology,RAS, Moscow, Russia 3Department of Environmental and Biological Conclusions: La The parasite fauna of Science, University of Jyvaskyla, rus gulls remains poorly known fo r many Jyvaskyla,Finland species, particularly in the southern hemi sphere. New records represent a large Argulus fo liaceus, an obligatory fish ec proportion of rep01ts. The highest affinities toparasite, has to match with its relatively (> 80 percent) between parasite faunae were rare, poorly predictable, and evasive hosts among two pairs of gulls with overlapping temporally and spatially. Parasites need to or neighboring breeding ranges in northern adjust recruitment to the period of high fish Europe (L . canus-L. fuscus and L. argen availability. Free swimming individuals tatus-L. hyp erboreus) . Other high similari have to find the hosts as quickly as possible. ties in parasite faunae were fo und between Objectives: Are a synchronized recruit host pairs composed of one large and one ment and regular fluctuations of the popula small geographic distribution, a condition to tion number typical of A. fo liaceus? What which the Simpson's coefficient is sensitive. behavioural traits ensure efficient host Results of preliminary field work in 1999 searching inA.joliaceus? suggest the high similarity between L. argentatus and L. canus near the White Sea Materials and Methods: Size and sex Biological Station (Chupa, Karelia) is not an structure of A. jo liaceus populations were artifact of analysis. monitored in a lake and fish fmm in Central Finland. Hatching dynamics were observed in the laboratory. Host searching activity patterns together with external and internal controlling fa ctors were studied using specially designed experimental aquariums. Results: Synchronized recruitment con trolled abiotically (temperature and, perhaps, illumination) was observed only in late spring. Later, asynchronous egg laying and, especially, hatching extended recruitment with no clear seasonal peaks of larvae. A distinct midsummer peak of larvae was observed only in a densely farmed popula- 85 tion of Onchorhynchus mykiss. Light during the study: adult males, adult females mediated switching from a «hover-and and immature chaffrnches. wait» (day) to «cruising» (night) host search Results: The dynamics of M micro ing strategy makes parasites efficient around phylla micropopulations on all chaffinch the clock. A. fo liaceus swimming activity is groups were similar and gradually increased controlled both internally (hunger state) and during the spring migration (April - May). externally (visual, chemical, and hydrome On the adult male chaffrnchs, the mite chanical stimuli related to fish). number continued to increase during the Conclusions: A prolonged period of re nesting period (June) and began to decrease cruitment in A. jo liaceus plays an adaptive significantly in July, because of plumage role in unpredictable ambient conditions, postrnating moult; the mite number slowly extending a "net of infectivity". A number of decreased at the end of summer and tl1e individual behavioural adaptations facilitate autumn migration (September - October). finding of rare and evasive hosts. Intensity On the adult fe male chaffrnchs, the mite and prevalence of infection depend on number decreased abruptly during the interplay between the behaviour of parasites nesting period, when the fe males sit together and their fish hosts. with nestlings. About 60% of mites mi grated from the adultfe male chaffrnches on to young birds. The mite micropopulation SEASONAL DYNAMICS OF THE on the nestling ready to leave the nest was FEATHER MITE MONOJOUBERTIA represented mainly by males, females (about MICR OPHYLLA (ASTIGAMATA: 25% each) and tritonymphs (38%). The mite ANALGOIDEA: PROCTOPHYLLO numbers on the female chaffinches contin DIDAE) ON THE CHAFFINCH ued to decrease in July under the in:fluece of FR INGILLA COELEBS plumage moult and slightly increased up to S.V. Mironov the autumn migration only. On young Zoological Institute, Russian Academy of Sci chaffmches, mite numbers quickly increased ences, St. Petersburg, Russia up to the begirming of the autumn migration due to the intense reproduction of mites. The Objective: Seasonal dynamics of the decrease of the micropopulation on the structure of Mo nojoubertia microphylla young birds began only in the end of autumn (Robin, 1877) micropopulations on different migration. sex and age groups of chaffinches Fringilla Conclusion: The dynamics of M micro coelebs (Passeriformes: Fringillidae) have phylla micropopulations during the nesting been studied during the nesting and migra period and autumn migration are considera tion periods in the North-West of Russia (in bly different on adult males, females and 1981-1982 and 1999). immature chaffrnches. These differences are Materials and Methods: The data on detetmined by the different roles of adult structure of mite micropopulations were chaffrnch males and females in the process obtained monthly from April to October, of infesting the nestlings with mites. The and 353 live individuals of chaffrncheswere mite micropopulation migrating from the examined. Three bird groups were deter adult fe male chaffrnchs onto the nestlings is mined within the chaffinch population represented mainly by older instars ready for 86 reproduction, fe males, males and trito than CIV s, and prevalence in the adults was nymphs. In the migration periods and 2.4 times higher than in the CVs, which probably the winter, the main part of the could result from a higher fe eding activityof mite population is represented by females, older stages. Similar results were obtained while in sununer the preimaginal instars on the wild population of C. strenuus in predominated. fe cted with Diphyllobothrium spp. Infection with both parasites impaired the escape ability of copepods, and influenced the FEEDING RELATIONSHIPS OF number of takeoffs to swim and time spent INTERMEDIATE HOSTS AFFECT swimming. In experiments, the infected TRANSMISSION SUCCESS OF copepods stayed near the swface while the LARVAL CESTODES uninfected remained close to the bottom. A 1 3 higher percentage of infected copepods was A Pastemak: , K. Pulkkinen , V. Mikheev and E.T. Valtonen3 consumed by fish in the laboratory. 1 Laboratory of Plankton Ecology, Institute of Conclusions: The success of parasite Oceanology, RAS, Moscow, Russia 2 transmission depends not only on Laboratory of Behaviour of Lower Vertebrates, host/parasite concentrations, but also on Institute of Ecology and Evolution, RAS, Mos availabilityof alternative prey. Prevalence is cow, Russia 3Department of Enviromnental and Biological higher in older copepodids due to their Science, Universityof Jyvaskyla, increased fe eding activity. Alterations in Jyvaskyla, Finland escape ability and motility influencing conspicuousness of infected copepods Copepods often serve as first intermedi increased the risk of being eaten by fish. ate hosts for tapeworms. Parasite larvae are eaten by copepods which, in turn, are con sumed by fish. The entire life cycle of KARYOTYPICAL DIFFERENTI tapeworms is determined by the fe eding ATION OF THE SPECIES OF relationships of the hosts. EUBOTHRIUM (CESTODA: PSEUDO PHYLLIDEA) Objectives: Does the presence of alter native fo od influence the acquisition of R. PetkeviciUte coracidia by copepods? Is acquisition of Institute of Ecology, Vilnius, Lithuania coracidia cmmected with the developmental stages of copepods? How do procercoids Objective: The identification of the spe alter the vulnerabilityof copepods to fish? cies of Eubothrium has always been a complex problem. The problem is pruticu Material and Methods: Experimental larly acute in respect of E. crassum and E. and field data on Cyclops strenuus - Triaeno salvelini. There ru·e a limited number of phorus crassus and strenuus - Diphyllo C. morphological characters available, several bothrium spp. systems. of these are variable and hence unreliable. Results: Alternative fo od reduced acqui Cytogenetic information can reveal differ sition of coracidia by copepods. In experi ences and similarities that may not be ments, the CVs of C. strenuus were infected obvious at the morphological level. with T crassus four times more frequently 87 Matedals and Methods: Karyotypes of CHANGING RISK OF TICK E. salvelini from Salvelinus leucomaenis BORNE ENCEPHALITIS IN and S. malma, E. crassum from Salmo salar SCANDINAVIA WITH PREDICTED and S. trutta trutta and Eubothrium sp. from CLIMATE CHANGE Clupea harengus membras were studied using conventional Giemsa staining and S.E. Randolphand D.J. Rogers Department of Zoology, University of Oxford, comparative karyometric analysis. Oxford, UK Results: The karyotypes, reported here fo r the first time, consist of eight chromo Objective: To understand the detenni some pairs (2n=l6). In the kruyotype of E. nants of the current distribution of tick salvelini chromosome pairs 1 and 2 are borne encephalitis virus (TBEv) in order to metacentric, pairs 3 and 4 are subtelocentric, predict any future changes in response to pair 5 is subtelo to submetacentric, pairs 6 climate change. and 7 are submetacentric and pair 8 is Methods: Complementary statistical and acrocentric. The two first pairs of homo biological approaches were used to relate logues are mru·kedly larger than the remain the observed spatial patterns of TBEv fo ci ing elements. The kruyotype of E. crassum and the temporal dynrunics of the transmis shows fa irly close affmity with that of E. sion process to satellite-derived climatic salvelini. The only statistically significant factors. The results were applied to future interspecific difference in relative lengths climate scenarios, based on Global Circula was revealed between chromosomes pair 5. tion Models, to predict future distributions. The best cytogenetic marker for species discrimination is the last 8tl1 pair of chromo Results: Enzootic cycles of TBEv were somes, which is metacentric in the karyo shown to depend on specific seasonal temperature profiles which determine a type of E. crassum. Comparative study revealed no significant interspecific differ specific pattern of the seasonal dynrunics of ences between kruyotypes of Eubothrium tick populations necessruy fo r TBEv trans mission. Predicted warmer winters and drier sp. and E. crassum. summers in the future apperu· to disrupt this Conclusions: The obvious similarity in system, leading to a northward shift, but kruyotype structure do not exclude the overall decrease in the geographical range of possibility to discriminate between E. TBEv. salvelini and E. crassum by karyotypic characters. Karyological observations Conclusions: When all the biological provide strong evidence to assign multi-factorial variables are taken into account, the threat of global warming does Eubothrium sp. from C. harengus membras not necessarily presage a mru·kedly worsen to E. crassum. ing situation for all vector-borne disease systems. 88 NEMATODE FAUNA OF SHEEP Conclusion: Trichostrongylus had the WITH EMPHASIS ON ZOONOSES IN lowest prevalence and intensity. The zoono THE WEST OF IRAN tic nematodes had low prevalences and intensities in the region. S.M. Sadjjadi Department of Parasitology, School of Medicine, Shiraz University of Medical Sciences, Shiraz, ELECTROPHORETYPES, GENO Iran TYPES AND BORRELIA AGENTS OF IXODES PERSULCATUS TICKB Objective: Determination of gastrointes tinal nematode fa una of sheep, especially A.V. Semenov1•2, A. N. Alekseel AND H.V. those which cause disease in man and are Dubinina2 1 zoonotic. Municipal Center ofViriologicalResearch, PCR Department,St. Petersburg, Russia Materials and Methods: The contents 2Zoological Institute, Russian Academy of of the abomasum, small intestine, colon and Sciences, St. Petersburg, Russia caecum of 160 slaughtered sheep were collected separately. The contents of each Objective: Ixodes persulcatus ticks col part of the gastrointestinal tract were filtered lected in the vicinity of St. Petersburg were through meshes placed in a column and tested for MDH (malatdehydrohenase) loci deposits were collected and fixed in 10% heterogeneity, Borrelia burgdoiferi species fo rmol saline solution. All nematodes were and etiological agents of echrlichiosis (HGE detected, counted and identified using - human granulocytarian echrlichiosis and azocarmine lactophenol. HME-human monocytarian echrlichiosis) . Results: More than 90% of the sheep Material and Methods: Isoenzyme were infected by at least one genus or analysis was used fo r MDH electrophore species of worm. A total of 24 genera or typing and genotyping. All ticks were species of nematodes were identified as examined using darkfield microscopy (DF) fo llows: From abomasum: Ma rshallagia to detect live spirochaetes. DF positive marshalli, Ostertagia occidentalis, 0. specimens were used fo r Borrelia and circumcincta, Ostertagia. sp., Parabronema Ehrlichia species determination. Borrelia skrjabini, Haemonchus contortus, Nemato species (B. aft elii, B. garinii, B. burgdoiferi dirus abnormalis, N oiriatianus, N archari. sensu stricto) and HGE & HMEagents were From small intestine: Nematodirus abnor detected using genus- and species-specific malis 58 %, N oiriatianus 21 %, N helve primers by PCR analysis. tianus 5 %, N spathiger 1%, N fzlicollis Results: Six detected MDH- 2%, battus 1%, archari 1%, sp . 1 %, N N N electrophoretypes were referred to three Capillaria sp. 2%, Tr ichostrongylus colu alleles of MDH gene and marked as geno briformis 1% and vitrinus 1%. From T types 1.1 (65% of total observations), 1.2 colon and caecum: Skrjabinema avis, (5%), 1.3 (19%), 2.2 (2%), 2.3 (3%), 3.3 Tr ichuris avis, skrjabini, !ani, ovina, T T T (6%) respectively. 56% ofDF positive ticks T disco/or and Trichuris sp. More than 10 were Borrelia-positive (13.4% of them were 000 nematodes were fo und in the small multiinfected with two or three types of intestine of which Nematodirus abnormalis Borrelia simultaneously). The most fre- at 7078 had the highest intensity. 89 quently detected Borrelia agent was B. at the larval phase because the utilization of aft elii (45% of all cases, B. garinii - 21%, the embtyonic yolk takes place already in B. burgdoiferi s. s. - 3%). HGE and HME the larva. This, however, does not occur agents were fo und in 11.8% of samples because it is necessary fo r a small larva to tested (9.4% - HME, 2.4% - HGE). attain, during its parasitic feeding, the large sizes of deutonymph and adult mites, which Conclusion: The 1 persulcatus popula tion in the vicinity of St. Petersburg was reproduce a rather conservative morphologi fo und to be heavily infected with Borrelia cal prototype with apparent secondary and HME agents. The degree of Borrelia simplification. The rapid ingestion of large infection between MDH-genotypes was masses of fo od by the parasitic larva has an different. This is the first case of HGE agent inevitable result in non-feeding and signifi fm ding in the St. Petersburg region. cant regression of the protonymphal instar. The tritonymphal instar undergoes a quite Acknowledgements: The research de synchronous morphological reduction that scribed in this publication was made possi obviously indicates the concordant evolution ble by Grant 9600864 Danish Research of the whole ontogenesis in this group of Council. trombidiformmites. MORPHOPHYSIOLOGICAL AND MORPHOLOGICAL AND KARYO ECOLOGICAL ANALYSIS OF LOGICAL STUDIES OF ECHINO ONTOGENESIS IN TERRESTRIAL CHASMUS SP. CERCARIAE (TREMA P ARASITENGONA (ACARIFORMES) TODA, ECHINOCHASMIDAE) By AB. Shatrov G. Staneviciilte,V. Kiseline and R. Petkeviciilte Zoological Institute, Russian Academy of Sci Institute ofEcology, Vilnius, Lithuania ences, St. Petersburg, Russia Objective: To present the results of mor Objective: A combined analysis of on phological and karyological investigations of togenesis in parasitengona mites that is Echinochasmus sp. Cytogenetic irtformation characterized by the phenomenon of «alter can reveal differences and similarities that nating calyptostasy» was carried out from may not be obvious at the morphological the standpoint of their ecological and mor level. phological specialization. Materials and Methods: Macrocercous Materials and Methods: Field and cul (zygocercous) cercariae were emitted by ture observations as well as light and elec naturally infected hydrobiid prosobranch tron microscopy of different organ systems snails Lithoglyphus naticoides. Molluscs during life cycle were used for the purpose were collected in the River Nemunas near of this study. Kaunas (Lithuania) during the years 1996- Results and conclusion: The enrich 1998. Cercariae were observed alive in egg ment of eggs in yolk and the progressive albumin in order to reveal the morphology, embryonization of the prelarva instar repre especially the excretory system. The mitotic sent an evolutionary tendency fo r acceler chromosomes of somatic cells of parthenites ated development that theoretically could lead to a complete loss of feeding capability 90 were analyzed using air-drying technique THE ECOLOGICAL VALUE OF and Giemsa staining. ACANTHOCEPHALANS FOR Results: Macrocercous (zygocercous) DETECTING TRAFFIC RELATED cercariae of Echinochasmus sp. have an oval POLLUTION body and vety large tail (reddish and 3-4 B. Sures longer than body). Flame cell formula: 2 Zoologisches lnstitut I - Okologie/Parasitologie, [(1+1+1+1) + (1+1+1+1)] = 16. The ducts Universitat Karlsruhe, Karlsruhe, Germany of excretory systems contain 8- 1 0 large composite and 2-3 small granules. The A major source of aquatic metal con caudal excretory duct is bifurcatedat its end, tamination is road runoffentering lakes and opening in the fr rst third of tail. rivers. This runoff contains a complex The mitotic complement of Echino mixture of potential toxicants. High sedi chasmus sp. consisted from 20 chromo ment and water concentrations of heavy somes, ranging in size from 2.11 to 7.64 f.tm. metals like zinc (Zn), cadmium (Cd), chro The ftrstpair of submetacentric-metacentric mium (Cr) and lead (Pb) m·e commonly homologues was larger than the remaining fo und near highways as these toxic metals elements, which decrease in size fairly are constituents of fuel, brake linings and gradually and represent acrocentrics (2"d vehicle tyres. In recent years another traffic pair), submetacentrics (6tl1 pair) and subtelo related metal emission could be observed: centics (all the rest) units of genome. fo llowing the introduction of automobile catalysts in the middle of the 80s there is an Conclusions: Cytogenetic data about increasing emission of the platinum-group this genus is poor and controversial. Accord metals (PGM) platinum (Pt), palladium (Pd) ing to existing data, Echinochasmus beleo and rhodium (Rh). Still, it remains unclear if cephalus has diploid chromosome number these metals are bioavailable for aquatic 2n= 14 and morphology of chromosomes are animals at all, and to what extent they vety similar to the chromosome set of become accumulated by the aquatic bio Sp haeridiotremaglo bulus. The katyotype of sphere. Echinochasmus sp. displays a pattern typical Therefore fteld studies as well as ex of most genera of Echinostomatidae - Echinopharyphium, Mo liniella, Hyp oder perimental investigations were conducted on aeum. According to the morphological data the accumulation of the above mentioned on Echinochasmus cercm·iae, this genus metals in different ftsh species, their acan seems to belong to a valid family Echino thocephalans and the established free living chasmidae, which appem·s more closely bioindicator, Dreissena polymorpha. To related to Psilostomidae than to Echi analyse metal levels in the ftsh tissues ' the nostomatidae. parasites and the mussels, various modern a11alytical techniques like electrothermal atomic absorption spectrometry (Et-AAS), inductively coupled mass spectrometry (ICP-MS) and total-reflection X-ray fluores cence analysis (TXRF) were applied. The results demonstrate that the zebra mussels and the parasites are suitable to 91 detect differences in contamination derived rats reached higher levels than in the unin fi·om mototway tunoff. But the bioconcen fected exposed rats. Additionally, the adre tr·ation of metals by the acanthocephalans nal tissue of only the exposed and infected was several times greater than that by zebra rats showed hyperplasia, indicating that the mussels. Acanthocephalans are therefore infection acts as a synergistic stressor in more sensitive indicators fo r detecting low addition to cadmium. While the acantho concentrations of dissolved heavy metals in cephalan infection leads to increases in the aquatic ecosystems than zebra mussels. The catecholamine levels, the levels of adrena experimental studies revealed fo r the first line and noradrenaline, as determined by time an uptake and therefore the bioavail high performance liquid chromatography ability of traffic related Pd by the exposed (HPLC), decreased after heavy metal expo aquatic organisms. sure. Accordingly, the infection was antago nistic to Cd-exposure. The mean cadmium concentration in M moniliformis determined FIRST STUDIES ON THE STRESS by electrothermal atomic absorption spec RESPONSE OF MAMMALS FOL trometry (ET-AAS) was 5,8 11g g-1 wet LOWING HELl\flNTH INFECTIONS weight, which was 20, 23 and 119 times ANDHEAVY METAL EXPOSURE higher than in host kidney, liver and intes B. Sures\ G. Scheef, W. Kloas3 AND H. tine, respectively. While female wonns Taraschewski 1 accumulated higher amounts of cadmium 1Zoologisches Institut I- Okologie/Parasitologie, than males, no tendency emerged between Universitat Karlsruhe, Karlsruhe, Germany the cadmium concentration and the weight 2Paul-Ehrlich-Institut, Bundesarnt fi.ir Sera und of individual acanthocephalans. Impfstoffe, Gentechnische Impfstoffe, Langen, Thus, our results show that parasites Germany 3Department of Inland Fisheries, Institute of have to be considered in (eco-) toxicological Freshwater Ecology and Inland Fisheries, Berlin, studies as they are able to accumulate high Germany amounts of heavy metals taken up by the host organism. On the other hand they also The impact of an infection with the acan affect the health of their hosts as demon thocephalan Mo niliformis moniliformis and strated by the exacerbation of cadmium a simultaneous Cd-exposure on the stress induced stress due to the acanthocephalan hormone levels of rats was studied. Afterthe infection. application of cadmium, cortisol levels in rats, as quantified by radioimmunoassay (RIA), significantly increased. Although the animals were heavily str·essed by the metal, cortisol levels at the time of killing did not significantly differ fi·om those in control rats, indicating that their adrenal systems were exhausted. Experimental infections with the acanthocephalan M moniliformis enhanced the impact of cadmium on the host animal, as cortisol concentr·ations in infected 92 ESTONIAN OESOPHA GOSTOMUM dominance which showed unstable egg SPP. ISOLATES INPIGS excretion. H. Talvik1, C.M. Christensen2 and T. Jarvis1 Conclusion: The present study describes 1Estonian AgriculturalUniversity, Tartu, Estonia for the first time 0. quadrispinulatum in 2Danish Centre for Experimental Parasitology, Estonian pigs. We suggest that this species The Royal Veterinary and Agricultural Univer may occasionally give rise to prolonged sity, Copenhagen, Denmark prepatent periods. Objective: Until recently, only the pres ence of Oesophagostomum dentatum had SEXUAL ORNAMENTS SIGNAL been documented in Estonia and no observa PARASITE RESISTANCE IN A MALE tions had been carried out to determine the FISH length of the prepatent period of the porcine J. Taskinen, R. Kortet ANDT. Sinisalo nodular worm infection. Department of Environmental and Biological Materials and Methods: Therefore, Science, University of Jyvaskyla, Jyvaskyla, eight Landrace/Estonian Big White Pig Finland crossbred piglets of approx. 15 kg body weight were inoculated with 2000 infective Objective: The «Hamilton and Zuk hy larvae of fo ur Oesophagostomum spp. pothesis» proposed that male sexual orna isolates collected from different Estonian ments could signal parasite resistance. We pig farms. Daily faecal samples were exam tested the hypothesis by applying a novel ined from day 15 p.i. and pigs were slaugh method-separation of dead and live para tered between days 44-60 p.i. Worms were sites. recovered by the agar-gel migration tech Materials and Methods: We used the nique and 100 worms from each pig were leaking fish Rutilus rutilus (males of which identified. develop breeding tubercles as sexual orna Results: The pigs commenced excretion ments), and the fo llowing measures of host of Oesophagostomum spp. eggs between resistance: proportion of dead parasites, days 18 to 43 p.i. The length ofthe prepatent parasite load (intensity/prevalence) and period was fo und to depend on both isolate immune function (spleen size). and pig. Two pigs showed single eggs on Results: As anticipated by the hypothe day 24 p.i., whereafter all samples were sis, there was a significant positive relation negative again until days 32 and 43 which ship between male ornamentation and host were therefore suggested to be the «true» resistance (proportion of dead parasites) prepatent period. All Estonian isolates against the gill parasite Rhipidocotyle contained 0. quadrispinulatum that pre campanula (Trematoda) in all of the three dominated in the two isolates which had the populations where the parasite occuned. latest average onset of egg excretion. The Correspondingly, numbers of live R cam two isolates dominated by 0. dentatum panula were significantly negatively associ showed markedly shorter prepatent periods ated with ornamentation in two populations. and fa irly constant egg excretion, compared However, we fo und no relationship between to isolates with 0. quadrispinulatum pre- ornamentation and the proportion of dead liver parasites Raphidascaris acus (Nema- 93 toda), load of R campanula or R acus, load Conclusions: The juvenile Crossbills of the gill parasite Myx obolus sp. (Protozoa), born during a cold season can be infected load of the liver parasite M miilleri, load of only after the breeding period (outside of the gill parasite Paradoplozoon homoion nest). The high haematozoa prevalence in (Monogenea), relative spleen size, or so the juvenile Crossbills shows that the period matic condition of fish. while small passerines are in the nest does Conclusions: Breeding tubercles on the not play a decisive role in their infection. body of male roach may signal parasite This contributes to the discussion on therole resistance in the present system against R of the nesting period in the infection of birds campanula, which also happened to be the with blood parasites. Unusual life histories locally most prevalent and abundant para may protect birds from infection with some site. In addition, we propose the proportion parasites either completely (Cuckoo, Swift) of dead parasites to be a useful measure of or at least during first months of their life resistance since it may provide a specific, (Crossbill) when individuals are especially direct, long-term measure of host immu vulnerable to parasitic infections. Conceiva nological response. bly, this advantage contributed to the evolu tion of unusual life histories ofbirds, such as the brood parasitism of Cuckoo, winter PARASITES IN HOSTS WITH breeding in Cross bill, nearly all-day flight in SPECIALLIFE IDSTORlES Swift.The parasites in hosts withspecial life histories provide opportunities to answer G. Valkilinasand T. Iezova some intricate questions in ecological Institute of Ecology,Vilnius , Lithuania parasitology, using relatively simple and inexpensive methods. Objective: To analyse the haemosporid ian parasites (Protista: Haemosporida) in birds with special life histories. ENVIRONMENTAL IMPACT AS Material and Methods: A total of 107 SESSMENT OF PARASITE CON specimens of Cuckoo (this bird species is TROL MEASURESIN CATTLE characterised by brood parasitism), 307 P.J. Waller1, S.O. Dimander\ G.W. Yeates2 and J. specimens of juvenile Crossbill (the birds Hoglund1 breed during the cold months of the year) 1SWEPAR, SLU/SVA, Uppsala, Sweden and 57 specimens of Swift (the birds spend 2Landcare Research, Palmerston North, New up to 20 hours per day in the sky and even Zealand more when they do not breed) were exam ined in the Western Baltics using the blood Objective: Environmental impact as smear technique. sessments are important issues with regard Results: The haemosporidians were not to registration of control agents (principally recorded in Cuckoo and Swift,but they were chemicals and drugs) fo r widespread agri common in juvenile Crossbills. Haemopro cultural use. This study was undertaken to teus tartakovskyi (7.2%) and Leucocytozoon determine the effect of long-term admini fringillinarum (14.3%) were the dominant stration of ivermectin (Ivomec® controlled species of parasite. release device), and the nematophagous fungus Duddingtonia jlagrans, compared 94 with untreated control cattle, on the range of generally beneficial soil nematode popula tions fo und in pasture soils. Materials and Methods: A plot study and a paddock field trial with the various parasite control options in cattle were monitored during two grazing seasons (1998, 1999). Soil samples were taken to a depth of about 3 cm using soil corers. Frequency of sampling and number of samples varied according to the design of the two investigations. Soil nematodes were recovered by standard procedures, counted, differentiated into various taxa and then aggregated into "functional groups" (i.e. plant feeders, fungivores, bacteriovores, camivores, omnivores, plant-associated etc ). This study will be continued during the grazingyear 2000. Results: In analysis of initial data, no significant differences in either total abun dance, or of functional groups, emerged. However, in early samples there appeared to be a greater proportion of bacterial-feeding and predacious nematodes in soil from the bolus treated paddock, whereas the propor tion of plant-feeding nematodes was lower. Although the same general trend is reflected in the plot study, on some sample dates there was an apparent decrease in the fungivorous nematodes Aphelenchus, Aphelenchoides. Conclusion: Results suggest that differ ences may emerge in soil microfaunal populations in relation to parasitological treatments applied to grazing livestock. These may affect decomposition and nutri ent cycling processes. Acknowledgement: This work was fi nancially supported by the Swedish Council for Forestry and Agricultural Research (SJFR). Bull Se and Soc Parasito/ 2000; 10 (2): 95-138 SUBMITTED PAPERS- POSTER PRESENTATIONS SPOROZOANS (SPOROZOA, API not only seasonal, but also age character COMPLEXA) - BLOOD PARASITES istics of haemoparasite infection of hosts OF TUNDRA WATER-SWAMP in a discrete population. BIRDS, SPENDING WINTER IN AZERBAIJAN COMPARATIVE ANALYSIS OF THE HELMINTHOCOENOSES OF NA M. Aliyev, G. Gaibova, M. Musayev and E. TIVE RIP ARIAN MUSTELIDS (L U Sultanov TR A LUTR A, MUSTELA LUTR EOLA) Laboratory of Protistology and Laboratory of IN RELATION TO WIDTH OF FOOD Ornithology, Institute of Zoology, Academy of SPECTRA Sciences of Azerbaijan, Baku, Azerbaijan E. Anisimova and V. Sidorovich Objective: Haemoparasites of birds of The Vertebrate Predation Research Group, passage (seagulls, swans, diving ducks and Institute of Zoology, Minsk, Belarus geese) wintering in the Island of Kur-Cosa (Kizil-Agadj Bay) of theCaspian Sea were Objective: Helminthocoenoses of the investigated. otter (Lutra lutra) and European mink Materials and Methods: Approxi (Mustela lutreola) were compared. mately 100 birds were examined. Blood of Materials and Methods: In Belarus, 49 the birds caught with traps and nets was otters and 41 European mink were exam taken by piercing the vein or cutting the ined for helminth parasites. At the same claw (in small species). Thin smears of time the width of fo od spectra of these blood were taken and stained azure-eosin by predators was evaluated.and results: The Giemsa-Romanowsky. The slides were dissected otters were infested only by five examined using a light microscope. helminth species. The helminthocoenosis of Results: Blood sporozoans were dis the European mink population was charac covered in one species of the family Anati terized by higher diversity (17 helminth dae and in one species of the family Lari species). Analysis of 1474 seats of the dae. European minks revealed 49 prey species. Conclusion: At the present stage of Food niche breadth calculated fo r eight prey research it is too early to make conclu categories comprised 3.36. By analysing sions about the poor fauna of haemococ 802 otter spraints, 29 prey species (includ cidians of birds. Investigation of blood of ing 19 species of fish) were fo und. Food birds of passage at the end of the winter niche breadth fo r eight prey categories was is not enough to understand the circula significantly lower than that of the Euro tion of parasites. It is necessary to study pean minks (2.62 versus 3.36). 96 Conclusion: Width of ecological niche genoides medians and Me torchis inter is one of the main factors influencing the memius - were conserved. Analysis of the fo rmation of helminthocoenosis in mustelid age structure of the mollusc population and populations. of the seasonal dynamics of their invasion allows us to conclude that life cycle stability in many trematode species is provided by THE SIGNIFICANCE OF MOLLUSCS intramolluscan stages of development. IN CONSERVATION OF TREMA TODE LIFE CYCLES The above conclusion is well supported by examples from parasites of Anatidae - N imbricatus, P. tuberculata, S globules G.L. Ataev\ E.V. Kozminslifand AA. Dobro and H volgensis. At least 30% of molluscs volski / containing parthenites of these species sur 1 Departrnent of Zoology, Russian State Peda vive the winter period. It is these that be gogical University,St. Petersburg, Russia come a source of infection fo r birds the 2Departrnent of Invertebrates, Saint Petersburg fo llowing year. State University, St. Petersburg, Russia The data obtained demonstrate that un der temperate climatic conditions molluscs The role of molluscs is generally recog can play an important role in supporting nized not only in supporting trematode life natural hotbeds of infection. Earlier a simi cycles as intermediate hosts, but also in lar conclusion was reached for more north dispersion of the parasites in the environ erly latitudes (Galaktionov, 1993). Probably ment. However, the question of the signifi such a conclusion is especially true fo r cance of molluscs in long-term conservation models where migrating species play the of hotbeds of trematode invasion remains role of the defmitive host. insufficiently studied. Moreover, the leading role in this is often attributed to definitive hosts. PARASITE SYSTEMS WITH ACAN THOCEPHALANS OF THE GENUS During 1990-1998 monitoring of the FIL/COLLIS LUHE, 1911 (ACAN parasite fa una of molluscs Bithynia tentacu THOCEPHALA) AND THEIR GEO lata was conducted in a pond in the St. Pe GRAPIDCAL MODIFICATIONS tersburg area. About 6000 snails were col lected and investigated. The overall preva lence of trematode infection was approxi G .I. Atrashkevich mately 30%. During the investigation de Institute of Biological Problems of the North formation of the age structure of the FEB RAS, Magadan, Russia Bithynia population took place, induced by the elevated mortality of molluscs born that Objective: Long-term studies of bird year and by a decrease in their fertility in acanthocephalans were conducted to show 1992 through 1993. However, despite the the main fe atures of parasitic helminth sys almost complete renovation of the mollusc tems of migratorybirds. population, all trematode species - Noto cotylus imbricatus, Psilotrema tuberculata, Materials and Methods: Original mate Sp haeriodiotrema globulus, Holostephanus rials from the Asiatic Subarctic were col lected during twenty years study using the volgensis, Cercaria helvetica XII, Pleura------�- 97 original methods of Kontrimavichus & IITSTOLOGICAL AND IDSTO Atrashkevich (1982). CHEMICAL CHANGES IN TOXOCARA Results: The genus Filicollis includes CANIS TISSUES UNDERTHE ACTION OF LEVAMISOLE two species -F.anatis (Schrank, 1788) and F. trophimenlwi Atrashkevich, 1982. The 1 dabbling duck A. acuta (Anatidae, Anseri R. AukStikalniene\ 0. Kublickiene and A 2 fo nnes) is absolutely dominant as a defmi Vysniauskas 1 tive host (90%) in northern populations of Faculty of Natural Sciences, Vilnius University, transpalaearctic F. anatis, but in some Vilnius, Lithuania 2 southern regions (Ukraine, Caucasus, S. Veterinary Institute, Kaisiadorys, Lithuania Ural) the main defmitive host is Fulica atra (Rallidae) , whose distribution range hardly reaches the subArctic. Only Asellus aquati Changes in the micromorphology and of cus (Isopoda) is known to be the intermedi glycogen deposits in the tissues of T canis ate host fo r F. anatis in Europe, whereas in under the action of 7 mglkg levamisole Asia five other endemic species of the ge were studied in vivo. Puppies naturally in nus Asellus have been identified as interme fe cted with T canis were used and exami diate hosts (Atrashkevich, 1997). nations were carried out on the worms eliminated from the host intestine 4, 10, 15, F. trophimenlwi is endemic to North 25 and 46 hr afterthe levamisole treatment. East Asia. Among 20 defmitive host species Semithin sections of the worms were only a few diving ducks are obligatory stained standardly with haematoxylin - hosts, with Clangula hyemalis dominant (up eosin for micromorphological investigation to 70% prevalence). Only two species of and fo r PAS reaction fo r glycogen deposit intermediate hosts are known. Parasitic changes in tissues of the worms. Amylase systems of both acanthocepalan species in was used to control PAS reaction. sympatric areas include the same intermedi ate host - one of the endemic Asellus spe The results show that levamisole causes cies, but different defmitive hosts (F. anatis serious alterations in the structures of cuti in A. acuta and F. trophimenlwi in C. hye cle, intestinal cells, hypodermis and muscle malis). cells of T canis and influences their carbo hydrate metabolism. The changes seen in T Conclusion: The spatial structures of canis afterthe treatment with levamisole in parasite systems with subarctic trophi F. the first few hours were slight. At 4 hr after menkoi are remarkable for the homogeneity treatment noticeable alterations were seen of their intermediate hosts. The transpalae only in cuticle. Small granules and vacuoles arctic anatis consists of heterogeneous F. appeared in other tissues only at 25 hr post parasite systems that differ in that their in treatment. Weak swelling of all tissues was termediate and obligatory defmitive host observed after treatment with levamisole. species are multifunctionalthrough changes The fine structure of nerve cords showed no in the main defmitive hosts. Thus, the noticeable changes. widely distributed parasite species of migra tory birds can fo rm relatively isolated PAS reaction was used to examine gly populations, even in northern regions, with cogen deposits in T canis tissues. Under the little or no mixing between them. action of levamisole the decrease in glyco gen was seen only in the intestinal cells at 4- 98 25 lu·. At the end of the experiment there HAEMOSPORIDIAN PARASITES was positive PAS reaction in all tissues. OF BIRDS FROM BELARUS It is concluded from these obvious and quick changes of cuticle after treatment with A. Babushnikova levamisole that this drug penetrates the T. Institute of Zoology, National Academy of cants body surfa ce. Sciences, Minsk, Belarus Objective: Based on the results of pre ECOLOGICAL ANALYSIS OF FISH liminary original investigations and litera PARASITE FAUNA IN KALININ ture data, to collate the available informa GRAD REGION RESERVOIRS tion on the distribution of haemosporidian parasites (Protista: Haemosporida) of birds E. Avdeeva and E. Evdokimova in Belarus. Kaliningrad State Technical University, Materials and Methods: A total of 21 Kaliningrad, Russia specimens of birds belonging to 10 species of the order Passeriformes were investigated Objective: During 1990 through 1999 a fo r haematozoa using the blood smear tech complex of investigations was carried out nique. The birds were sampled from the with the aim of determining the fish parasite State National Park Belovezhskaya Puscha fauna inKaliningrad Region reservoirs. in June 1999. Materials and Methods: eight species Results: Parasites belonging to three of fish in Rivers Pregel and Prokhladnaya, genera of haemosporidians were fo und: seven species of fish of Kaliningrad Bay Haemoproteus, Leucocytozoon and Plas and five species of fish in Lake Chistoye modium. We recorded haemoproteids in were investigated with the method of total tlrree species of birds. The highest intensity parasitological dissection. (3-4parasites per one field of the oil inrrner sion objective) was found in Luscinia lus Results: Parasites with a direct life cycle cinia. According to the literature, Haemo dominated in 17 fish species. Parasites with proteus sp. has been recorded in 22 species a complex life cycle were represented by 13 ofbirds in species. In most cases trematodes infected fish as metacercariae. Infections by other Belarus so fa r. We fo und Leucocytozoon types of parasites were insignificant. sp. in Turdus merula. The common passer ine hosts of leucocytozoids in Belarus are Conclusion: Deterioration of the eco crows (Corvidae). Nine species of leucocy system in a reservoir resulted in a decrease in the number of smooth myxosporean tozoids have been repotted in Belarus so fa r. In merula, we fo und Plasmodium sp in spores, and in variations in the size of their T. mixed infections with Leucocytozoon sp. valves and polar capsules. Sexually mature Only one species of Plasmodium has been trematodes were rarely represented in pol recorded in birds in Belarus. luted reservoirs. Few parasite species de velop through Copepoda. Apparently the Conclusion: Haemosporidian parasites influence of sewage water in reservoirs are common in birds in Belarus. All genera results in depression of tl1e development of of these parasites known to parasitize Palae planktonic Copepoda. arctic birds have been recorded. However, 99 the fauna and its distribution pattern remain The developmental cycle of P. laevis insufficiently studied. does not include a stage of encapsulated cystacanth. The bodies of acanthocephalans deformed and extruded from the intestine POMPHORHYNCHUS LAEVIS m·e not involved in the developmental cycle (M(JLLER, 1776) (ACANTHOCE of the parasite. PHALA) REMAJNS IN MESENTE RIUM TISSUE OF FLOUNDER P. laevis remains can be classified into (PLATICHTH YS FLESUS L): MOR three typesaccording to their origin: PHOLOGY, LOCALIZATION AND a) residuals of early stage or adult indi ORIGIN viduals that migrated but settled in unfa vourable environmental conditions (20%), E. Bacevicius b) residuals of migrating unsuccessful Fishery Research Laboratory, Klaipeda, cystacanths which had returned and adhered Lithuania by suction to the external wall of the intes tine (11 %), More or less decomposed remains of c) residuals of dead adults extruded by Pomphorhynchus laevis were fo und in the the immune response from the intestinal body cavities of flounder. There is no com wall (69%). mon opinion about the origin, structure, and Flounders infected with encapsulated role of those remains in the life cycle of P. worms do not fillthe role of transport hosts. laevis. Two hundred P. laevis remains from flounder (n=850, 1=5-35 cm) intestinal tracts were studied. All the remains could be clas GAMMARIDS OF LAKE BAIKAL sified as fo llows: those having only a pro BASIN AS INTERMEDIATE HOSTS boscis sucker - 37 %, with proboscis bulb OF HELMINTHS and neck - 53 %, and those mummified by salinization- 10 %. Deformed, shrunk bod D.R Baldanova, U.A. Kritskaya and N.M. ies («mummies») were fo und on liver mes Pronin entery. Depending on the location in the Institute of General and Experimental host body cavity proboscides were either Biology, Ulan-Ude, Russia connected to the external side of the intesti nal wall by a fibroblast thread or, in those Objective: Helminthological research of attached to a liver mesentery or in the peri gammarids in Baikal was conducted. toneal cavity (70 %), without a thread. Re Gammarids are one of the most diverse mains attached via the connecting tissue are groups of animals in Baikal (270 species mostly aggregated in the middle patt of the and 80 subspecies). Two hundred and intestinal tract or on either side of a rectal twenty-nine gammarid species are endemic sphincter (68 %). The length of fibroblast to Baikal. thread (mm) was positively correlated with the distance from the rectal (r=0.95) or in Materials and Methods: Helmintho testinal (r=0.72) wall. The longer the thread, logical dissections of 44121 specimens the older were the P. laevis remains. belonging to 209 species was carried out. Literature data (Bekman, 1954; Zaika, 1965; Shabaev, 1966; Pronin et al., 1986; 100 Baldanova, Pronin, 1992; Baldanova, 1998) COMPARATIVE ANALYSIS OF THE and results of the authors' own research of PARASITE FAUNAS OF BROWN gammarids are summarised. TROUT PARR (SALMO TR UTTA L.) Results: Gammarids are infected with FROM SOME WATER BODIES OF 14 helminth species. The hosts of Cyatho THEP AANAJARVI NATIONAL PARK cephalus truncatus are Gammarus lacustris, ANDTHE RIVER OULANKA Gmellinoides fa sciatus, Pallasea cancellus, P. cancelloides, Poecilogammarus pictus Y.Y. Barskaya, B.S. Shulman and E.P. and Eulimnogammarus fuscus. A cestode of leshkov fam. Hymenolepididae has been reported in Institute of Biology, Karelian Research Gmellinoides fasciatus. The trematode Center, RAS, Petrozavodsk, Karelia, Russia Crepidostomum fa rion is has been fo und in Micruropus posolskii and Gmellinoides Objective: The purpose of the present ja sciatus, while Crepidostomum metoecus study was to reveal the diversity of the para was fo und in Gammarus lacustris and site fauna of brown trout parr (Salmo trutta Gammarus kozhowi. Allocreadium sp. has L.) of the Oulanka - Paanajarvi - Olanga been recorded in Acanthogammarus victo lake-river system. rii. The nematode Cystidicola fa rionis is a Materials and Methods: Brown trout parasite of Gammarus lacustris and Gam parr aged 2+ from the Paanajarvi National marus kozhowi. Te trameres fissispina and Park (Mutkajoki River, Siltajoki River, Streptocara crassicauda have been reported Lohioya River) and the River Oulanka in Gammarus lacustris. The acanthoce (Finland) were studied. The research was palans Polymorphus magnus and P. minu carried out in 1998-1999. Fifteen specimens tus have been recorded in Gammarus lacus from each locality were dissected. The tris and Gmellinoides fasciatus. Hosts of study was catried out according to standard Echinorhynchus borealis are G. fa sciatus, methods (Bikhovskaya-Pavlovskaya, 1985). Eulimnogammarus cyanoides and Pallasea Results: In the Mutkajoki River five cancelloides. Hosts of E. salmonis are G. species of parasites were fo und: Myxospo fasciatus, E. cyanoides and Micruropus rea - 1, Ciliophora - 1, Protozoa incertae posolskii. The host of E. truttae is M pos sedis - 1, Cestoda - 1, Acanthocephala - 1. solskii. The parr in this river were mostly infected Conclusion: Palaearctic helminth spe by Dermocystidium sp. (prevalence - 67%, cies use endemic species of gammarids as intensity - 40). The parasite fa una of brown intermediate hosts inBaikal Lake. trout parr from the Siltajoki River consisted The research was supported by the Rus of eight species: Myxosporea - 1, Cilio sian Foundation for Basic Research (NN phora - 3, Trematoda - 2, Acanthocephala - 97-04-96212 and 00-04-49598). 1, Copepoda - 1. Crepidostomum ja r ion is was the most widespread parasite. The spe cies composition of brown trout parr in the Lohioya River consisted of nine species: Myxosporea - 4, Ciliophora - 3, Monogenea - 1, Trematoda - 1. The prevalence and intensity of infection by all parasites were low. In the Oulanka River 12 species of 101 parasites were fo und: Ciliophora - 3, Nema ter depends directly on the species of toda - 3, Trematoda - 5, Acanthocephala - 1. obligatory intermediate host of a cestode The acanthocephalan Neoechinorhynchus and its trophic relationships. An example of crassus was recorded in Fennoscandia fo r fo ur species of Hy menolep is (s.l.) parasitiz the first time. The prevalence and intensity ing the gulls of the subarctic Region and of infection by Ciliophora (Apiosoma sp. using the same cmstaceans (Branchiopoda) 78/22, Capriniana piscium 83/39) were as intermediate hosts shows distinct specific quite high. differences in the structure of their eggs Conclusion: Species composition of which determine their flotation qualities. parasites of brown trout parr from therivers This character, together with the determina studied was fo und to consist of 23 species. tion of differences in the morphology of Analysis of the parasite fauna shows that copulative organs, enabled us to differenti the species composition in some of the riv ate between species, while the participation ers displayed low species diversity. Para of cmstaceans as intermediate hosts (2) sitological data indicate that the Oulanka allowed to distinguish this group of cestodes River ecosystem is exposed to considerable from the composition of the genus Ward ium anthropogenic impact whereas the Paana (= Hymenolep is s.l.). The taxonomic rank of jarvi NP water bodies remain undisturbed. hosts (2) and localization of parasites (3) played an important role in the description of new Cestoda species parasitizing the IMPORTANCE OF ECOLOGICALLY diving ducks and shorebirds of the Subarc DETE�D CEUUlACTERS � tic region. A special place in solving the THETAXONOMY OF CESTODES problems related to our revision of cestodes of the subfamily Aploparaksinae is given by (4). Allocation of the metacestodes of S. Bondarenk and V. Kontrimavichus closely related species of cestodes to differ Institute of Ecology, Vilnius, Lithuania ent types of cysticercoid was useful in most cases in solving disputed questions of Sometimes, in order to differentiate be cestode identification. tween closely related species of cestodes mainly parasitizing the same or similar species of defmitive hosts, investigation of PARASITE COMMUNITIES OF the morphological characters of the para SMALL RODENTS � THE PAANA sites is not sufficient. The problem can be JARVI AND OULANKA NATIONAL solved by analyzing ecological data ac PARKS quired during the investigation of life cycles S.V. Bugmyrin, E.P. Ieshko, V.S. Anikanova of the parasites. The most important data and L.A. Bespyatova are: 1) the stmcture of outer membranes of Institute of Biology, Karelian Research live eggs determining the mode of infection Centre, RAS, Petrozavodsk, Karelia, Russia of their intermediate hosts; 2) the taxonomic rank of hosts ( defmitive and intermediate); Objective: The diversity of helminth 3) localization of parasites in the defmitive and ectoparasite communities of rodents in and intermediate hosts; 4) the structure of undisturbed ecosystems (Paanajarvi Na cysticercoids. The structure of the outer tional Park, Russia and Oulanka National membrane of eggs (1) as a specific charac- Park, Finland) was studied. 102 Materials and Methods: The material in the sample predete1mines the highest H was collected early in July 1998 (i), early in value. October 1999 (ii) in the south-eastem prut of the PNP and in late September 1999 (iii) ru·ound Oulru1ka Biological Station. Hosts SPECIAL FEATURES OF THE FOR were captured by snap-traps arranged in a MATION OF RODENT HELMIN line. 63 small rodents belonging to three THOCOENOSES IN URBAN LAND species were examined fo r parasites during SCAPES OF BELARUS the period of investigation. The diversity (H) and evenness (E) of parasite communi E. Bychkova ties was measured using the Shrumon index. Institute of Zoology, National Academy of Results: (i) Fifteen individuals of the Science, Minsk, Belarus family Cricetidae were examined, repre senting the species Microtus oeconomus (3) The objective of this investigation was and Clethrionomys glareolus (12) of two to study specific features of the fo rmation of age groups: overwintered adults and juve helminthocoenoses under urban pressure. niles. Their pru·asite fauna was represented Materials and Methods: In 1996-99, by 15 species, belonging to fo ur taxonomic 1520 rodents from Minsk belonging to the groups: Cestoda (4), Nematoda (3), Ixodi fo llowing eight species were dissected and dae (1) and Gamasina (7) (also Siphonap examined for helminth parasites: Apodemus tera and Anoplura). Lice (55%, 4.9), fleas jlavicollis, Melchior, 1834 - 100 specimens; (53%, 2.8), Hirstionyssus isabellinus (30%, Apodemus sylvaticus, Linnaeus, 1758 - 4.3) and the nematode Heligmosomum mix eight specimens; Apodemus agrarius Pallas, tum (52%, 1.1) domi\nated the community. 1771 - 120 specimens; Clethrionomys The measured diversity index and evenness glareolus Schreber, 1780 - 27 specimens; were 2.34 and 0.8, respectively. Microtus oeconomus Pallas, 1778 - one (ii) The host material consisted of 17 specimen; Microtus arvalis, Pallas, 1778 - bank voles oftwo age groups:ju veniles and 72 specimens; Mus musculus Linnaeus, subadults. Nine parasite species were fo und: 1758 - 1087 specimens; and Rattus rattus Nematoda (4), Cestoda (2) and Gamasina Linnaeus, 1758- 105 specimens. (3). The dominant nematodes were Syp ha Results: The community of rodents in cia petrusewiczi (30%, 7.1) and H mixtum Minsk consists of synanthropic species 78.4 (88%, 2.5). H=1.73, E=0.72. %, fo rest species 8.9% and field rodents (iii) Thirty-one bank voles and two grey 12. 7%. Host ecology, dynamics of in sided voles were examined. Six parasite trapopulation and interspecific relationships species were recorded: Nematoda (2), had great influences on the formation of Cestoda (1) and Gamasina (3). The most helminthocoenoses. The low population abundant were S petrusewiczi (18%, 4.6), density of forest rodents is the reason for H mixtum (97%, 5.2) atld lice (55%, 5.6). low helminth infestation in the city. The H= 1.54, E=0.74. specific habitats of rodents in urban territo Conclusion: The species diversity in a ries limit contact between different ecologi parasite community depends on the age cal groups of rodents. Rare contacts caused structure of the host population. Thus, in our decreased richness of the helminthofauna in case, the presence of overwintered animals different ecological groups of rodents. 103 Conclusion: The specific feature of the mia, anthrax, paraphylariosis and other helminthofauna of different species of ro diseases. dents in urban territ01y is the dominance of Conclusion: The study of horsefly spe helminthsfrom synant hropic rodent groups. cies composition and abundance in various ecosystems of the Lower Volga Region ROLE OF HORSEFLIES (DIPTERA, allowed us to determine the location where TABANIDAE) IN THE ECOLOGICAL tabanids had a severe influence on the pro BALANCE OF BlOTA ductivity and resistance of domestic ani mals. The presence of natural fo ci of infec tions, the agents of which can be transmitted P.A. Chirov and A.M. Peterson by horseflies, necessitates protecting ani Saratov State University, Saratov, Russia mals against their attack. Objective: The nature of horseflies lo calisation beingheterogene ous, their species HOUSE DUST MITES IN FOUR VIL composition, abundance and probable role NIUS PRE-SCHOOLS in agent transmission in different ecosys tems of the Lower Volga Region were stud ied. A. Dauartiene and A. Jurkuvenaite Department of Zoology, Vilnius University, Materials and Methods: Tabanids Vilnius, Lithuania were collected from horses in various eco systems in the territory of Kalmykia, Astra Objectives: To describe house dust mite khan, Volgograd and Saratov regions. The species, their prevalence and seasonal varia total number of horseflies collected was tion in Vilnius pre-schools. 25000. Materials and Methods: The study was Results: Forty-eight species of horse carried out during one year - March 1999- flies were found, 13 of which were new for February 2000. From the total list ofVilnius the region studied. Horsefly females were pre-schools (total number 117) fo ur were able to have over five gonotrophic cycles randomly selected: two where children stay during a season. This determined their high for the whole week and two of 12 working fe cundity and abundance in their habitats hours per day. Dust was collected in March, with stable conditions fo r their develop June, October and January, taking 14 sam ment. The duration of horsefly activity was ples at each institution per day (from mat 100-150 days. Mass flight lasted from 1 to tresses, stuffed toys, and carpets) and later 1.5 months in different landscapes. Particu examined using the Acarological method. larly high abundance (up to 210 individuals The total number of samples examined was per collection), which have negative effects 224. on the horses pastures, was noted in the north-western region of the Caspian Sea. Results: Twelve mite species were The weights of wild and domestic animals fo und in all samples examined. Dermato subject to mass attack by tabanids declined phagoides pteronyssinus (Trouessart) and D. and the animals' resistance to transmitted far inae Hughes (Pyroglyphidae) were present diseases decreased. Decrease in resistance in samples at 2-2000 and 2-200 individu was conducive to transmission of tularae- als/gram respectively. Mites were present in all room sites examined. The numbers of 104 mites detected in one gram of dust varied in relationship between eggs and uterine wall, samples taken from different floors and at the so called placental-like interaction, was different seasons. In the samples gathered established. The maturing eggs of N mo from the ground floor mites were present gurndae are attached to the uterine wall. 2.95 times more frequentlythan on the first The cytoplasmic outgrowths and lamellae floor. For the first time in Vilnius stuffed of uterine epithelium are believed to inter toys were examined and the numbers of lace intimately with the shell and an outer mites fo und was surprisingly high. Accord envelope surrounding the eggs. ing to our study the number of mites was Conclusion: The process of uterus de highest in October and lowest in March. velopment in N mogurndae is similar to that in a group of lowest cestodes of the DIFFERENTIATION AND PLACEN order Caryophyllidea and is one of the gen TAL-LIKE ORGANIZATION OF THE eral ways to form both the uterus and ducts UTERUS IN NIPPOTAENIA MO of the reproductive system. GURNDAE (CESTODA) Acknowledgments: The study was sup ported by the Russian Foundation fo r Basic 1 2 Research (grantN 99-04-48465). V. G. Davydov and J.V. Komeva 1Zoological Institute, Russian Academy of Sci ences, St. Petersburg. Russia ENDOPARASITES OF JUVENILE 2Institute fo r Biology of Inland Waters, Russian (AGE 0+, 1+) COD, GAD US MORHUA, Academy of Sciences, Borok, Russia IN ICELANDIC WATERS - PRELIMI NARY RESULTS Objective: Investigations were carried out on Nippotaenia mogumdae Yamaguti et M. Eydal, A. Kristmundsson, S.H. Bambir, R. Miyata, 1940 (Cestoda: Nippotaeniidea). Magnt.lsd6ttir and S. Helgason Materials and Methods: Adult worms Institute for Experimental Pathology, were removed from the intestine of the fish University of Iceland, Keldur, Reykjavik, Perccottus glenii, collected fi·om shallow Iceland lakes near Vladivostok, Russia. Materials were fixed in 4% glutaraldehyde in cacody Objective: To investigate when late buffer, post-fixed in Os04 (for 1 h), endoparasites start to infect wild cod (age dehydrated and processed fo r TEM exami 0+ and 1 +) in Icelandic waters and to ana nation. lyse the progress of infection. Results: The process of uterine differen Materials and Methods: Approxi tiation can be divided into three main mately 1,000 fish caught in 1998-1999, at 2, stages. In the first stage, aggregation of 4, 6, 10 and 18 months of age, have been undifferentiated cells leads to the syncytium examined so far. Two year classes, 1998 formation. In the second stage, as a result of and 1999, were studied. All major organs active autophagic processes, the uterine were screened fresh, frozen or fotmalin cavity is fo rmed. Then the uterus is sepa fixed (conventional histology) and blood rated from the parenchyma by the interstitial samples were taken. basal plate and is surrounded by bands of Results: The fo llowing parasites were circular muscle. For cestodes the unique fo und: Protozoa - coccidian oocysts and 105 other coccidian stages, Lama sp.; Myxospo HOW NEMATODES COMMUNI rea; Digenea - Brachyphallus crenatus, CATE, - WITH A NOTE ON PHAS Derogenes varicus, Lepidapedon elonga MIDULTRASTRUCTURE tum, Podocotyle atomon, Prosorhynchoides gracilescens (metacercariae) Stephanosto 1 2 H.-P. Fagerholm , M. Brunaska , A. 4 4 mum sp. (metacercariae); Cestoda - Pseu Roepstorpf, G. Jungersen and L. Eriksen dophyllidea and other plerocercoids (lar 1 Departrnent of Biology, Institute of Parasitol vae); Nematoda - Anisakis simplex, Cucul ogy, Abo AkademiUniversity, Abo, Finland lanus cirratus, Hy sterothylacium aduncum, 2 Pseudoterranova decipiens; Acantho Institute of Parasitology, Slovak Academy of Sciences, Kosice, Slovak Republik cephala - Corynosoma sp. (larvae), Echi 3 norhynchus gadi. No blood parasites were Royal Veterinary Agricultural University, CEP, fo und. Endoparasites (i.e. metacercariae) Denmark 4 were first detected in a two months old Royal Veterinary Agricultural University, juvenile cod. At least six species of parasites Clinical Studies, InternalMedicine, Denmark have been found in four months old cod, 11 in six months old, 11 in 10 months old and In ascaridoidean nematodes, only a few 17 species in 18 months old cod. Prevalence somatic sense organs are present: Al. Lat and intensity of each species generally in eral amphids (n=2) in the head region, B/. creased with age. Cephalic sense organs (4), Cl. Labial sense Some species reached a prevalence of organs (12), D/. Cervical deirids (2), E/. 100% (D. varicus, P. atomon, A. simplex). Asymmetrically oriented midbody centrids Among the helminths, digenean species (Fagerholm et al.: J. Parasitol., 1999, 85: 41- reached the highest intensities, with num 47; probably homologous to the not well bers up to 270 per fish(L. elongatwn). defmed structures often called post-deirids), Some parasites/pathogens still remain to F/. Lateral phasmids on the tail (2). Caudal be identified. Other diseases/pathogens sense organs of the male (21- ea. 200). Re detected include: Pseudobranch tumor, sults to date on the structure and function of epitheliocystis (bacterium) and Ichthyo somatic sense organs deal primarily with phonus hoferi (fungus). the single pair of lateral cephalic amphids (see Ashton & Shad: Vet. Parasitol. 84, Conclusions: Diversity of parasites in (1999) 297-3 16). With the sequencing of creases with age, as does prevalence and the total C. elegance genome in 1998 a intensity of infection, although there are perception of the number of genes involved differences between year classes and locali in the function of sensory dendrites has ties. It seems that metacercariae of been obtained. The number of functional Stephanostomum sp. (found mainly in the genes involved in chemoreception amounts eyes) have not been reported earlier in this to no less than some 500. These can be host. divided into fo ur basic receptor families This study is supported by the Icelandic (Troemel (1999), BioEssay 21: 1011-1020). Research Council and The Icelandic Repub The function of the 11 pairs of chemosen lic Fund. sory neurons fo und in the amphid is now known while that of the neurons in other sensory organs largely remain to be investi gated. This applies also to basic structure. 106 The phasmid ultrastructure has, so far, only INTERCOMMUNICATIONS BE rarely been studied. This is the case al TWEEN ZONE OF MITEBITE (GE though the presence or absence of phasmids NUS IXODES) LOCALIZATION AND is a fe ature traditionally used for dividing THE HUMAN BIOENERGETICAL nematodes into basic clades. In the present SYSTEM study serial ultrathin sections of the phas mid of Hy sterothylacium auctum (Rud. V. Fyodorova and V. Kosenco 1819) (Ascaridoidea: Raphidascarididae) from the gut of Zoarces viviparus L. were Novgorod State University, Novgorod studied under TEM. The results give sup Veliky, Russia port to the general view of a single dendrite being present in phasmids. Each phasmid The problem of the increase in the num was laterally situated and the dendrite runs ber of mitebites (Ixodes) among local in anteriorly (The nucleus apparently being habitants is a vital question toward the end of the 20tl1century. situated at the level of the cloaca, as in C. elegans). Close to the papilla the dendrite of This is the result of social problems in the left-hand phasmid was provided with modem Russia because townspeople, owing numerous blunt and rather short fm gerlike to the state of national economics, must protuberances extending into the sheath cell. work on plots of cultivated land, so contact These are apparently homologous with the between the mban population and the natu abundant rather thin radiating extensions ral environment is becoming greater. The that we have observed in Ascaris suum. number of persons who had mitebites in When phasmids are present in nematodes a creased fo urfold during the last 10 years consetvative basic ultrastructure of these (1987-1997) in the territory of Novgorod would give additional support fo r Se Region alone (Bolshacova et al., 1997). cementea forming one basic clade in the The obj ect of our research is the study of Nematoda. However, only by research on intercommunication between zones of fu rther representative entities can this be mitebites (Ixodes) and the human bioener verified. It is wmth noting that the orienta getic system. tion of the caudal male sense organs is also governed by specific genes, as recently Methods: Measurements were carried shown in studies by Emmons and Pitch. It is out with the help of device ACEC (analyser evident that nematode sense organs are of condition of power canals) with appro much more complicated both structurally priate programme-apparatus complex. The and functionally than previously imagined basic working principle of the device is a and that communication between individual diagnosticmethod, based on the registration wotms and the perception of the environ of change in the impedance of the tissues as ment are well evolved. a result of the vital activity of the organism. Device ACEC allows us to find biologically active points (BAP), to measure power parameters (module of full electrical resis tance) of these points and with the help of the programme-apparatus module to proc ess the results of the changes. 107 Results: During the research 13 zones and stained with carbol-fuchsin by the were identified where bites of the ticks were Ziehl-Nielsen technique. The faeces were fo und. The greatest number of large bites of also preserved in 2.5% solution of potas these ticks takes place on the ear, neck, sium bichromate. The preserved faeces armpit and on the head. This is explained by were centrifuged with the flotation liquid the fact that except for these areas the whole according to the standard technique. All body is covered by clothes, so it is easier fo r preparations were examined with a light ticks to bite the above mentioned places. microscope. The human body consists of bioenergetic Results: Oocysts of coccidia were de points which are very visible on the outside tected in two swan species and in one gull surface of the human body. species (Larus argentatus). When 53 tick bitten victims with bio energetic points were compared it was Conclusion: In late winter the fauna of noted that the sites of bites were the same intestinal coccidia in the water and marsh fo r these two species of ticks 1 persulcatus, fowl of the tundra is rather poor. Examina and 1 ricinus. tion of these birds at the time of their arrival These experiments showed that the fe fo r overwintering is desirable. males always sucked blood from the region of medium resistance. Hailer's organ helps CESTODES OF THE EIDER DUCK these species to fmd places of high biologi POPULATION INHABITING SPITS cal activity on the body (BAP). BERGEN AND FRANZ JOSEF LAND (PRELIMINARY COMMUNICA COCCIDIA (SPOROZOA, APICOM TION) PLEXA) OF WATER AND MARSH FOWL OF THE TUNDRA OVER A.K. Galkin\ K.V. Galaktionov1 and S.F. WINTERING INAZERBAIJAN Marasae� 1Zoological Institute, Russian Academy of Sci G. Gaibova, M. Aliyev, M. Musayev and E. ences, St. Petersburg, Russia Sultanov 2Murmansk Marine Biological Institute, Kola Laboratory of Protistology and Laboratory of Scientific Center, Russian Academy of Sciences, Ornithology, Institute of Zoology, Academy of Murmansk,Russia Sciences of Azerbaijan, Baku, Azerbaijan Objective: The cestode fauna of eider Objective: Some water and marsh birds duck, Somateria mollissima, is reported of the tundra overwintering on Kur-Kosa fo r the first time from the northern bor Island in the Caspian Sea (Kyzylagaj Bay) der of the species distribution, Spitsber in the winter-spring period were studied to gen and Franz JosefLand. fm d out if they carry oocysts of coccidia (Eucoccidia, Coccidia, Apicomplexa). Materials and Methods: In August 1991 three birds were collected in Spits Materials and Methods: Fresh faeces bergen (Bellsund) and fo ur on Franz of passing gulls, swans, pochards, and geese JosefLand (FJL). In August - September caught by special nets were collected. Fine 1992 10 more birds were captured on smears of faeces on slides were prepared FJL: six in the vicinity of Hooker Island, 108 as. in 1991, and fo ur on Apollonov Island PARASITE INVASION AS A RISK ' (81 20 · N), much farther to the north. FACTOR IN THE DEVELOPMENT Results: All cestodes collected belong OF INFLAMMATORY DISEASES OF to the genus Microsomacanthus. Prevalence THEPELVIS ORGANS 1 1 of cestodes in eiders eve1ywhere was 100% T.A. Gasanova , AV. Mikhailov and P.A. and the parasite load was very high (up to Chirol 250.000 specimens per bird). In Spitsbergen 1Perinatal Centre, the Town of Engels, Saratov and in the southern parts of FJL M micro State Medical Un iversity, Saratov, Russia soma dominated, while the subdominant 2Saratov State Un iversity, Saratov, Russia species was M jagerskioldi. These species Objective: The aim of the study is to es are distinguished from each other mainly by timate the parasite (Giardia Iamblia, pin the form of the strobila, slender and long in wmms, Toxocara) infection among female M microsoma and shmt but broad in M patients with chronic inflammato1y diseases jagerskioldi. The third related species, M of pelvis organs (pelvis inflammation dis diorchis, only once noted in Spitsbergen, ease - PID). was more widespread in the vicinity of Hooker Island. At the northern extremity of Material and Methods: Immuno FJL all the eider ducks examined, juve enzymatic analyses [Test sets of Toxocara niles and adult, were infected by M duc strip, LambliaAT-strip of Vector (Best pro ti/is. The latter is a common parasite of gulls duction)] were used to investigate the level and has never been recorded before in eider of Toxocara and G. Iamblia invasion. Fae ducks. ces smears and solutions were also analysed to detect Giardia cysts. Pinwmm eggs were Conclusion: In comparison with Su detected in the morning urine centrifugate barctic regions such as Iceland (Baer, 1962) and also by the study of perianal fo lds im and Murman Coast (Belopolskaya, 1952; prints on adhesive tape. Total number of Galkin, 1997), the cestode fa una of eider patients - 144 females. Age of patients duck in Spitsbergen and FJL is scanty in ranged from 2 to 36. species composition but not in prevalence and intensity of infection. Further investiga Results: The fr equency of PID cases is tions may confi1m the lack of dilepidids shown in Table 1 (next page). (Lateriporus), fm 1briariins and M hetero The high level of parasite invasion spinus there. V erifled systematic data fo r common in children at ages 2-5 might be a comparative ecological studies is of great result of insufficienthy giene. value. Correct analysis of eider ducks' para Conclusion: Parasite invasion of PID site fauna is impossible without the detailed patients increases the inflammatory proc description of microsoma (Creplin, M esses, lowers the effectiveness of antibacte 1829) - the type species of the genus - rial therapy, prolongs the recovery period based on old collections (Fuhm1arm, 1913) and is a main risk factor for development of and newly obtainedmat erial. chronic PID promoting decompensation of The work was supported by INTAS genitourinary tract mucous membrane dis (project N 97-10224). bacteriosis and activation of persistent in fe ctions. All PID patients had to be exam ined for giardiasis, enterobiasis and toxocarosis for the female PID prophylaxis. 109 Parasites Ratio of patients (%) Girls of2-5 yr Girls of9-14 yr Women with with chronic vul- with acute salpin- infeiiilitym vovagmits goophoritis marriage (n = 27) (n = 34) (n = 83) Giardia 25.9 11.7 30.1 Pmworms 40.7 14.7 15.7 Toxocara 40.7 20.6 40.9 Table 1. Frequency ofPIDcases EUDIPL OZOON NIPPONICUM (DI- describe tegumentary structures durmg the PLOZOIDAE, MONOGENEA) parasite's ontogenetic development. COMPARATIVE ANALYSIS OF MI Results: Considerable seasonal changes CROECOLOGY AND HABITAT were observed m population dynamics, SPECIFICITY IN RELATION TO population age structure, development of PARASITE ONTOGENETIC DEVEL sexual organs and tegumentaJy structures. OPMENT Conclusion: There is no doubt that E. nipponicum seems to be a very useful M. Gelnar, I. Hodova, B. Koubkova, A. model parasite species for microecological S imkova ANDT.H. Zurawski studies of parasite spatial distribution m Department of Zoology and Ecology, Masmyk cludmg analysis of niche segregation, mtra University, Bmo, Czech Republic and mterspecific competition and aggrega tion. There is also need fo r a detailed TEM Objective: Study of Eudiplozoon nip study to clarifY the structure and function of ponicum (Goto, 1891) (Monogenea, Diplo some tegumentaJy papillae and their role m zoidae), parasitic on the gills of carp ( Cy parasite microhabitat segregation. prinus carpio ), a parasite of Far East origin which was introduced to Europe. Materials and Methods: Durmg the period fi·om 1988 to 1998, a long term study of the morphology, biology and ecology of E. nipponicum was performed under the conditions of a fish farm pond system. Scannmg electron microscope techniques were used to study the parasite surface to 110 TAXONOMY AND SYSTEMATICS SOME CHARACTERISTICS OF THE OF THE MONOGENEAN FAMILY DISTRIBUTION OF TE TR AONCHUS DIPLOZOIDAE IN THE LIGHT OF MONENTERON (MONOGENEA) AND RECENT KNOWLEDGE ERGA SIL US SIEBOLDI (CRUSTA CEA) ON GILLS OF PIKE ESOX LUCIUS M. Gelnar, B. Koubkova, I. Hricova, M. Dlouha and M. Klfmova Department of Zoology and Ecology, Masaryk P .I. Gerasev University, Bmo, Czech Republic Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia Objective: Members of the Diplozoidae are unique among platyhelminths in that Objective: Study of gill parasites of two adult worms are fused in permanent pike. copulation, each individual being unable to Matelials and Methods: Pike (N=22) survive alone. Until now there have been no were collected in Pskov Region in May data on the morphometric variability of 1995 and 1996. All parasites, plus areas of structures of taxonomic importance. Be gills, hemibranchs and sectors of gills were cause of this species identification is very counted as percentages. For comparisons of difficult. percentages the Fisher criterion was applied. Matelials and Methods: Phase Con Results: Significant positive correlations trast microscopy, Nomarski DIC, special were fo und between monogeneans and area staining techniques and digital image analy of 1) gills (r = +0.91), 2) hemibranchs (r = sis have been used fo r investigation of struc +0.86) and 3) sum of areas of two hemi tures of taxonomic importance. branchs of gill pairs adj acent to each other Results: Out of nine tested methods fo r (r=+0.92). Percentage distribution of crusta staining of attachment apparatus sclerites of ceans had 1) negative correlations with the individuals of model parasite species area of gills (r = -0.34), 2) hemibranchs (r = (Paradiplozoon homoion, P. bliccae, Eudip -0.19) and 3) a positive correlation (r = lozoon nipponicum and Diplozoon para +0.58) with the percentage of total area of doxum) , three methods are suggested for two hemibranchs of adj acent pairs of gills. use: Gomori trichrome, Lignin Pink and Preference for I-II pairs of gills and middle modified peracetic acid alcian blue for SS and angular sectors of gills was revealed in groups. monogeneans and preference fo r IIl-IV pair of gills and the anterior and posterior sectors Conclusion: Due to the above of gills was revealed in crustaceans. Signifi mentioned staining techniques, a com cant negative cOITelations between wonns parative analysis of the morphometric vari and crustaceans in the middle (r = -0.96) ability of structures of taxonomic impor and posterior sectors of gills (r = -0.91 ), and tance were petfonned to evaluate current a high positive conelation (r = +0.56) be criteria used in species identification. Taxo tween both parasites in the sum of areas of nomic revision of this parasite group is two hemibranchs of adjoining pairs of gills recommended. were shown. 111 Conclusion: Our data suggest antago Fish species of the same genus had nistic relationships between these parasites lower similarity indices of monogenean and indicate a direct relationship of occur populations (MP) (18.3%; 5.4%) than those rence of these gill parasites and the flow of from different genera within one family - respiratory cunents passing through slits 32%; and from different families - 23%. between the gill pairs. MP from fishes of the same ecological group had relatively high similarity indices (33%; 73%). The lowest index (7.6%) was COMPARATIVE ANALYSIS OF recorded for MP from the most ecologically MONOGENEAN FAUNAS AND different groups of fish. POPULATIONS FROM SEVERAL BELONIFORM FISHES Conclusion: The similarity of MF de pends mainly on the degree of affmity(sys tematic) between the hosts. In contrast, the L.A. Ghitchenok similarity of MP does not show any clear Biological Faculty, Moscow State dependence on the affinities between the University, Moscow, Russia hosts, but depends mainly on their ecologi cal similarity. Objective: A comparative analysis was canied out of both fauna and populations of monogeneans from beloniform fish belong THE LABORATORY RAT AS DE ing to different ecological groups (the oce FINITIVE AND INTERMEDIATE anic group proper, the pseudo-oceanic and HOST FOR SA RCOCYSTIS RODEN neritic). The fauna of a particular unit of the TIFELIS (PROTISTA: COCCIDIA) environment is a list of the animal species FROM THEBLACK RAT occupying that unit, while the population is a real community of animals, which de J. Grikieniene andL. Kutkiene pends also on the numbers of single-species Institute of Ecology, Vilnius, Lithuania populations (Beklemishev, 1970). Materials and Methods: Monogeneans Objective: As we have shown recently, from 1 520 specimens of fishes belonging S. rodentifelis can circulate among Wistar to 6 species of Beloniformes. A compara Norway laboratory rats (Rattus norvegicus) tive analysis was carried out using quantita by means of carmibalism. In such a life tive measurements of similarity. cycle both sexual (oocysts and sporocysts) Results: Fifteen species of mono and asexual (sarcocysts) stages of the para geneans were recorded. Relatively high site were detected in the same host species. similarity indices of monogenean faunas This study aimed to investigate the possibil (MF) were fo und in fish species of the same ity of transmission of S. rodentifelis from genus (0.56; 0.57), while the lowest was black rats (R rattus) to Norway rats without fo und in fishes of different families (0.20). the participation of a carnivorous defmitive Fishes of the same ecological group had host (cat). lower similarity indices MF (0.30; 0.49) Materials and Methods: Three cap than the most ecologically different groups tured black rats which had been infected of fishes(0.33). with sarcocysts, morphologically in distinguishable from S. rodentifelis, served 112 as the initial source of material. Laboratmy two sites near Vienna, namely Stockerau rats were infected by fe eding them with and Lobau, between 1960 and 1962. A muscle from black rats. Microscopy of comparison of the helminth species stained muscle sections and unstained composition and the intensities are given squashes of sarcocysts and of small intestine and the dependence of the helminth samples were used, as well as various flota fauna on the sex and age of the host was tion and concentration methods fo r also investigated. investigation of other coccidia. Materials and Methods: One hun Results: Sarcocysts were detected in dred and fifty-five specimens of the muscles of laboratol}' rats after eating mus common shrew, Sorex araneus L. , 115 cles of black rats containing sarcocysts with obtained from Stockerau and 40 from mature cyst merozoites. In another experi Lobau, were dissected. Body cavity, the ment, thin-walled sporulated oocysts and organs (heart, liver, lung, spleen, kid sporocysts were found in the small intes neys, pancreas and urinary bladder) and tines of rats on the 9th and 30th days after the alimentary tract (oesophagus, stom having fe d them with infected muscle. ach and the intestine) were examined. These intestines were infective fo r healthy Results: So far, four species of cesto rats. The laboratol}' rat served both as de des (Molluscotaenia crassiscolex, Hy finitive and intermediate host fo r S. roden menolepis diaphana, Neoskrjabinolepis tifelis. schaldybini, Hy menolep is singularis), eight species of nematodes ( Calodium Conclusion: S. rodentifelis can spread between individuals of two different rat cholidicola, Crenosoma skrjabini, Euco species by predation. The Norway rat acts leus oesophagicola, Liniscus incrassatus, as a predatmy defmitive host in the para Longistriata sp. l, Longistriata sp.2, site's life cycle while the intermediate host Parastrongyloides winchesi, Porro larvae) and two species of is the black rat. Definitive hosts fo r S. ro caecum sp . de ntifelis (cat and Notway rat) belong to trematodes (Brachylaemus fu lvus, Dicro two different orders. coelium soricis) have been fo und. In total, 92.9 % (144 out of 155 shrews) carried at least one species of helminth. THE HELMINTH FAUNA OF THE 78.7 % (122/155) had cestodes, 71.6 % COMMON SHREW, SOREX ARA (11 1/155) nematodes and 54.8% NE US L., FROM TWO SITES NEAR (85/155) trematodes. The maximum in VIENNA,AUSTRIA tensity of cestodes was 1419, of nema todes 85 and oftrematodes 16 per shrew. In general adult shrews were more heav C. Hoerweg ily infested than juveniles. Department of Systematic Zoology, Institute of Zoology, University of Vienna, Vienna, Conclusion: The high prevalence of Austria helminths in Sorex araneus L., as fo und in the neighbouring countries of Austria, Objective: The fauna and infection was confirmed fo r the first time for the levels of intestinal helminths of the environs of Vienna. Nevertheless, there common shrew, Sorex araneus L., were is a need fo r further investigations, espe studied. The shrews were collected at cially with recent collections. 113 ARE POPULATION DYNAMICS OF dergone a steady increase in density during �LOW PTARNUGAN AFFECTED the last five years, and our continued study BY PARASITES? will allow us to obtain detailed, long-tenn quantitative data on the changes in the para 1 2 1 site community with changing host density. P. Holmstad , P. Hudson and A. Skorping 1 More specifically we will ask the fo llowing Department of Zoology, Un iversity of Bergen, questions in our study: Bergen, Norway Do parasite burdens increase prior to a 2 Un it of Wildlife Ep idemiology, Department of Biological and Molecular Sciences, Un iversity of host population decline? Sterling, Stirling, UK Which are the dominant parasites asso ciated with the decline? Most field investigations of host-parasite Is there a relationship between breeding population dynamics have been restricted to production and parasite intensity? the study of a single parasite species, and Is there an association between estimates often conducted over short time intervals of host body condition and parasite inten relative to the life expectancy of the host. sity? Willow ptarmigan (Lagopus lagopus L.), Is there a positive association between like most other vertebrate populations, carry parasite species? a whole community of parasites and the transmission of each is likely to be affected Is this relationship linear or non-linear both by host densities and climate. indicating synergism between parasite spe cies? In order to get a long-tetm data set in which data fo r both host and parasite densi ties could be obtained, we have collected ECOLOGICAL ANALYSIS OF THE autumn samples of willow ptarmigan since FISH MYXOSPOREAN FAUNA OF 1992. All hosts have been shot from a single AZERBAIJAN WATER BODIES IN coastal locality in Kattfjord (69°40' N, THE LIGHT OF NEW DATA ABOUT 18°15' E), at the island Kvaleya in Troms THE BIOLOGY OF THESE PARA county, Norway. This sampling program SITES will continue at least until 2002. Each ptar migan is thoroughly searched for protozo ans and helminths in the intestinal system, Sh.R. Ibrahimov and R.K. Kerimova blood and other tissues. Institute of Zoology, Azerbaijan Academy of Sciences, Baku, Azerbaijan So far we have fo und fo ur species of nematodes - Ascaris compar, Capillaria Objective: The analyses of fish infec caudiriflata, Sp lendidofilaria papillocerca tion with myxosporeans in relation to the and Tr ichostrongylus tenuis; and two spe location of spores in a water body. cies of cestodes - Hy menolepis microps and Paroniella urogalli. Oocysts of Eimeria sp. Material and Methods: In 1972-99, 4167 have been fo und in the caecal contents. fishes belonging to 63 species and subspe Leucocytozoon lovati, Trypanosoma sp. and cies were examined in Azerbaijan water microfilariae have been fo und in blood bodies. The myxosporean collection of the smears. The ptarmigan population has un- 114 Institute of Zoology of Azerbaijan Acad Materials and Methods: Twenty-two emy of Sciences was also studied. birds of five species, Somateria mollissima, Results: Sixty-seven species were fo und Rissa tridactyla, Calidris maritima, Larus in Azerbaijan water bodies. argentatus, L. marinus, and about 8 000 specimens of the intertidal molluscs Lit Conclusion: The myxosporean fauna of farina saxatilis and Mytilus edulis, were benthic fishes is richer ( 60 species) than that examined. of pelagic fishes (24) because oligochaetes in which the actinosporean phase grows Results: Himasthla larina was fo und in usually inhabit the bottom. The myxo the hetTing gull, while Himasthla leptosoma sporean fauna of lowland water bodies is infected the population of the purple sand richer (60) than that of pre-mountain (42) piper. The kittiwakes and eiders were free and especially mountain ( 18) water bodies of infection by echinostomatid trematodes. because the water current carries the myxo The molluscs infected with larval echi sporean away from mountain and pre nostomatids were fo und only in the mountain areas to the lowland. Yamyshnaya and Khlebnaya inlets in the samples collected on the sand and mud The myxosporean fa una depends on the intertidal flats. The infection levels of dimensions, geographical location and eco whelks were rather low (not higher than logical conditions of every water basin. All 3%), while the blue mussels were heavily Azerbaijan rivers flow down into the Cas infected (80-100 %). pian Sea (53 species) and bring myxo sporean spores. Kura ( 49) is the largest river Conclusion: Representatives of only of this region. Araz (29), nmth-eastem one genus of the family Echinostomatidae, Azerbaijan rivers (17) and Lenkoran rivers Himasthla, were recorded in the seabirds of (15) are smaller and have very fast currents. the Barents Sea. Their major definitive Apsheron Peninsula (12) is characterized by hosts here are purple sandpipers and sea semi-desett climate. gulls. The fo rmation of infection fo cuses occurs strictly in cettain areas, which are enclosed or well-sheltered embayments and ON THE FORMATION OF INFEC basins of a fjord type witl1 vast sandy or TION FOCUSES OF ECIDNO muddy intettidal flats. The successful estab STOMOSIS IN SEABIRDS IN THE lishment of the infection fo cuses is facili BARENTS SEA tated by the fact that infection of the second intetmediate host, blue mussel, is very high (up to 100 %). This is due to the existence D. Ishkulov of the endogenic reproductive stage in the Murmansk Marine Biological Institute, Kola first intetmediate host L. saxatilis and the Scientific Centre, Russian Academy of Sciences, continuous emission of cercariae into the Murmansk, Russia environment. The work was supported by INTAS Objective: The echinostomatid trema (project N 97-1 0224). tode fauna was studied in the seabird and intertidal mollusc populations of the Barents Sea (East Murman coast) in 1995-1997. 115 THE TRICHJNELLOSIS SITUATION EFFICACY OF SOME INTEGRATED INESTONIA AND NON-CHEMOTHERAPEUTIC CONTROL METHODS AGAINST 1 2 LAMB MONIEZIOSIS INESTONIA T. Jarvis\ I. Miller and E. Pozio 1 Department of Parasitology, Estonian Agricul tural University, Tartu,Estonia A Kaarma and E. Magi 2 Laboratmy of Parasitology, Instituto Superiore Institute of Infectious Diseases, Estonian di Sanita, Rome, Italy Agricultural University, Tartu, Estonia Objective: Lamb monieziosis one of the Objective: To evaluate the present epi frequently registered diseases in Estonia. demiological situation of trichinellosis. According to our investigations, approxi Materials and Methods: From 1992 to mately 9-14% of young sheep were in 1999 muscle samples from 1987 sylvatic, fected. To work out control measures, dif domestic and synanthropic animals were fe rent pasture regimes were combined with collected. Using the method of artificial prophylactic treatments. The main purposes digestion the prevalence and intensity of of our trials were to reduce the use of che trichinellosis were determined. Species of motherapeutic preparations and to avoid Tr ichinella were identified by PCR (RAPD) contamination of the environment. analysis. Materials and Methods: The major Results: Cases of human trichinellosis invasion sources of sheep monieziosis in have been registered almost every year. Estonia were examined. Trials were con Trichinella infection was established in all ducted on sheep flocks of several Estonian animal species (14) investigated. The preva large-scale farms where lambs were grazed lence of infection was highest in wolves on cultivated pastures. Investigations on the (79.4%), raccoon dogs (50.0%), lynxes effectiveness of applied measures were (47.4%) and red foxes (42.1%). Intensity of estimated by identifYing the eggs of invasion was highest in raccoon dogs and Mo niezia on the basis of faecal flotation red foxes (up to 213 larvae/g). 1.0% of wild procedures. boars and 29.4% of bears were infected. Results: In three trials, Mo niezia-free Only two fo ci of trichinellosis in domestic lambs were allowed to graze alone on pas pigs were registered, T britovi and T sp i tures contaminated with parasites of the ralis being identified. Most of the wild ani previous year. After the observation period mal species were infected with T nativa 16%, 24% and 55% respectively were in and T britovi. In farm fur-animals T nativa fe cted. The mean prevalences of infection and T sp iralis, and in brown rats T sp iralis, of weaned lambs, grazed with ewes on were fo und. contaminated pastures, were 8%, 18% and Conclusion: Trichinella spp. are widely 24% respectively. Lambs weaned during distributed in the wildlife fauna in Estonia. the indoor holding period and grazed sepa Domestic animals and human beings are rately on contamination-free pastures were endangered by trichinellosis. not infected with Moniezia eggs. With sepa rate grazing on contaminated pasture and a single treatment after20 days of the grazing period, 3-8% of lambs were still infe cted 116 in October. If the lambs were treated twice, pupation and emergence, numbers of dead after 20 and 45 days, 1.5-2.2% were still individuals, wing length and dry weight of infected in autumn. Clinical symptoms of the adult mosquitoes, were recorded. the disease were not observed. Results: Under abnormal light condi Conclusions: Pastures contaminated tions, the rates of development and mortal with parasites of the previous year are the ity increased and the ratio of wing length main sources of infection for lambs. Infec and dry weight changed. Response to con tion levels in lambs grazed with ewes on stant light conditions of some values (dura contaminated pastures were lower than in tion of larval development, m01tality rate) lambs grazed separately on comparable was less pronounced than to constant dark pastures. Two treatments of lambs after 20 ness. and 45 days of the grazing period on con Conclusion: The results obtained agreed taminated pastures eliminated the spread of with the data on the influence of some un monieziosis in sheep flocks. favourable factors (high temperature, high salinity, overcrowding, toxicants) on Culi SEVERAL ASPECTS OF THE IN cidae. Thus, the hypothesis concerning the FLUENCE OF ABNORMAL LIGHT existence of a non-specific response to ab CONDffiONS ON THE PREIMAGI normal conditions in blood-sucking mosqui NAL STAGES OF BLOOD-SUCKING toes has been confirmed. More stable values MOSQUITOES of Aedes communisunder constant light, in comparison with constant darkness, may be caused by development of preimaginal S. Karpova stages in the nearly constant natural light Laboratory of Parasitology, Institute of conditions. Biology, Petrozavodsk, Russia Objective: The influence of abnormal THE HELMINTHS OF REPTILES light conditions (constant darkness, constant FROM SOUTH URALS light) on the preimaginal stages of Aedes communis, one of the abundant species of V. Khabibullin blood-sucking mosquitoes in Karelia, was Department of Biology, Bashkir State studied to test the hypothesis concerning the University, Ufa, Russia existence of a non-specific response to the unfavourable conditions in Culicidae. The Objectives: To investigate the helmin rates of larval and pupal development and thofauna of eight species of reptiles (out of mortality at preimaginal stages, and the 13 species inhabiting South Ural). morphometric fe atures of imago were in vestigated. Materials and Methods: Reptiles were collected from the territory of Bashkiria Materials and Methods: Fourth-instar (South Ural, Russia) during 1997-1999 field larvae collected fi·om natural reservoirs seasons. One hundred and fo urteen speci were reared under different light condi mens of reptiles belonging to eight species tions: 1) natural photoperiod (control; (including one turtle, three lizards and fo ur light:dark =17:7 h.), 2) constant darkness, snakes) were examined by total helmintho- 3) constant light. Onset and duration of 117 logical dissection. All helminths found were neans Pseudodactylogyrus anguillae and P. counted, sorted and collected. bini from the gills and the blood flagellate Results: We fo und 16 species of reptile Trypanosoma granulosum were found fo r hosted helminths: eight species of trema the first time in reservoirs of Latvia. The todes and eight species of nematodes; five most widespread parasite of the eels was the species were present as larval stages. Anguis nematode A. crassus, which was found for fragilis and Natrix natrix were 100% in the first time by us in 1994. Nematode in fe sted (3/3 and 15/15 respectively). The fection has increased gradually during the most common parasites were Te lorchis fo llowing years and reached a maximum in 1998 when all fishes were infected with a assula (Dujardin, 1845) from N natrix and Neoxysomatium brevicaudatum (Zeder, maximum intensity of 126 nematodes. In I 800) fromA. jragilis. Usmas Lake a mass disease of eels oc curred, with mortalities recorded. Conclusion: The intensity of infec tion varied considerably from one to 403 A preliminary study of the A. crassus parasite individuals per host (74.0 on life cycle shows that paratenic hosts of this average); the mean prevalence of inva parasite in Latvia are mainly ruffand rarely sion was 27.2%. The results obtained are perch. the first contribution to the investigation Both species of Pseudodactylogyrus, as of the helminthofauna of reptiles in well as Acanthocephalus lucii, were fo und South Urals. very frequently. Although the maximum intensity of infection by monogeneans was high, pathogenic changes in the gills were PARASITES OF THE EEL IN LAT not observed. This was also the case with VIA Bothriocephalus claviceps, Raphidascaris acus and Camallanus lacustris. 2 M. Kirj usina 1 and K. Vismanis 1 Other parasite species infect eels seldom National Veterinary Laboratory, Riga, Latvia and even rarely. The exception is Anodonta 2 Biology Faculty, Latvian University, Riga, sp., maximum numbers of which were ob Latvia served in the early spring. It occurred fre quently at high intensifies, but caused no Objective: Eels from the Venta River, disease. Usmas Lake and coastal waters of the Riga Conclusion: The parasite fauna of the Gulf were investigated. eel in reservoirs of Latvia is poorer than Materials and Methods: Since 1994, repotted in the literature by other authors 63 eels were studied by the method of total (Shulman, 1949; K.0ie, 1988; Kennedy et parasitological examination. Thirty-four al. , 1992). It is necessary to study other fishes were investigated completely, while reservoirs in the country to make a final 29 were examined solely fo r infection by conclusion. the nematode Anguillicola crassus. To study the biology of this nematode we also examined 4 7 perch and 18 ruff. Results and discussion: A total of 17 parasite species were found. The monoge- 118 ON THE ISOPOD FISH PARASITE parasites of the family Cymothoidae are FAUNA OF THE ATLANTIC COAST characteristic mainly of the tropical regions OF NORTH-WESTERN EUROPE of the ocean. Most probably this fact is connected with the level of tension witl1in trophic links in different geographic zones A.F. Kononenk:o of the world's oceans. Depmtment of Invertebrate Zoology, Moscow State University, Moscow, Russia ISOPODA PARASITES OF NORTH Objective: The present study focuses on SEA FISHES the analyses of the Isopoda fish parasite fa una of the Atlantic coast of north-western A.F. Kononenko Europe, which are represented by faculta tive, tempormy and permanent parasites. Department of Invertebrate Zoology, Mos cow State University, Moscow, Russia Matelials and Methods: From the au thor's own records and from literature data, about 195 species of Isopoda fish parasites Objectives: The present study fo cuses are known for the Atlantic Ocean basin. on the analysis of the Isopoda fish parasite Twenty-six species have been recorded in fauna of the North Sea which is represented the North Sea, including three facultative, by facultative, tempormy and permanent 17 tempormy and 6 permanent parasite parasites. species. Fifteen parasitic isopods are known Matelials and Methods: From the au fo r the Irish Sea, including four fa cultative thor's own records and from literature data, and 11 tempormy species. Representatives 26 species of parasitic isopods infecting 29 of the fam ily Cymothoidae - petmanent fish species have been recorded from the parasites of fishes- have not been recorded North Sea. Facultative and tempormy fo rms in this region. Only one species was re are dominant (80%). The family Aegidae is corded in the Baltic Sea - Aega psora L., most numerous both in number of species 1759, which is widespread throughout the and in number of infected hosts. Four eco North Atlantic. Aega psora is a tempormy logical groups of fishes infected with para parasite of fish known also from the North sitic isopods are recognized fo r the North and Irish Seas. Sea. Fishes of the coastal shelf group are Results: Thus, the parasitic isopod fau dominant (15), and the largest number of nas can be compm·ed only for the North and parasitic isopods is also known from iliese Irish seas. There are 11 common parasitic (14). 45.8% of them are representatives of isopod species in tl1ese seas (Czekanovski the family Aegidae - the characteristic ele Sorensen index of similarity is equal to ment of the isopod fauna of bottom and 0.54). Of these, two species are facultative near-bottom fishes. Five coastal pelagic fish and nine species are tempormy. There are species have four isopod species. Seven no common species belonging to the family isopod species, common in the shelf waters Cymothoidae. of the north-eastern Atlantic, have been recorded on the six fish species of the Conclusion: Isopods representing facul bathypelagic group. One isopod species has tative and temporary parasites are the usual been recorded on three species of brackish fonns fo und in boreal waters. Permanent water and euryhaline fishes. 119 Results: Most of the 26 isopod species Results: There are critical values of PI recorded from the North Sea belong to the above which the host population dies out. families Aegidae and Cirolanidae. Wide These values of PI are proposed to be used spread in northern waters, they are distrib as the quantity estimation of permissible uted far to the south, having been recorded PrP. For its designation the term «parasite even near the coast of Africa. Only five of capacity of the host population» (PCP) is the most eurybiotic species of the family used. The mathematical approach to esti Cymothoidae have reached this sea, which mate PCP is developed and its strict defrni is the northern boundary of their distribu tion is given: tion. PCP is the heaviest possible PI, at Conclusion: The isopod fauna of the which, with the generation number T ap North Sea is heterogeneous, including ele proaching infinity, there exists at least one ments of both boreal and Mediterranean initial population size nJor which the prob Lusitanian faunas. ability of size decrease through T genera THE PARASITE CAPACITY OF THE tions is less than the probability of its in HOST POPULATION crease. Conclusion: Use of empirical values of PI as an estimation of PrP is insufficient. E.V. Kozminsky The estimate of PrP should be carried out in Department of lnvertebrate Zoology, Faculty the light of the changing extinction prob of Biology and Soil Sciences, St. Petersburg ability of the host population under the in State University, St. Petersburg, Russia fluence of parasites. Objective: The problem of parasitic pressure on the host population (PrP) is ECTOPARASITES OF JUVENILE considered. (AGE 0+, 1+) COD, GADUS MORHUA, Materials and Methods: The approach IN ICELANDIC WATERS - PRE is based on the concept of changing extinc LIMINARYRESULTS tion probability of a host population under the influence of parasites. A. Kristmundsson, M. Eydal, S.H. Bambir, R. The change of host population size is Magntisd6ttir and S. Helgason considered as Markov process. The prob Institute fo r Experimental Pathology, abilities of all changes of population size for University of Iceland, Keldur, Reykjavik, a generation are described by a matrix of Iceland transition (n) with dimensions N,11axXN,11ax (maximum population size). The probabili Objective: To investigate when ecto ties of all possible size changes for T gen parasites start to infect juvenile wild cod erations can be calculated as nr. (age 0+, 1 +) in Icelandic waters and to ana lyse the progress of infection. Only available parameters are used in Materials and Methods: In 1998-2000 the construction of matrix n - sex ratio, fe cundity, mortality, prevalence of infection approximately 720 fish at four, six, 10 and (PI). It is assumed that these parameters are 18 months of age were examined for ecto constant and that the parasitic castration of parasites. In addition, 30 cod at 22 months the host takes place. 120 of age were screened fo r Lernaeocera PARASITISM AS A BIOINDICATOR branchia/is. OF ECOLOGICAL AND SEASONAL Results: Tr ichodina sp. was fo und on CHARACTERISTICS OF SEABlliD six and 18 months old fish with a preva POPULATIONS (BY THE EXAMPLE lence of <7% (intensity 1-29). Gyrodactylus OF THE MURRES FROM SEVEN sp. was fo und on one six months old fish ISLANDSARCHIPELAGO) with a prevalence of <1% (1 1 ). Caligus sp: larval stages were fo und on all age groups. V .V. Kuklin At fo ur months of age no Caligus larvae Mwmansk Marine Biological Institute, Kola were detected in 1998, but in 1999 the Scientific Centre, Russian Academy of Sciences, prevalence at the same age was 20% (1-3); Mwmansk, Russia > 98% (1-5 1) at six months; 53% (1-8) at 10 months and 36% (1-2) at 18 months. The Objective: In 1993 and 1999, a parasi prevalence of Clavella adunca was <12% tological survey of two murre populations, (1) at six months, 22% (1-3) at 1 0 months the cmmnon guillemot, Ur ia aalge, and and 46% (1-6) at 18 months. Lernaeocera Brunnich's guillemot, U. lomvia, of Seven branchia/is was only fo und on the 18 and Islands Archipelago was carried out to 22 months age groups with a prevalence of study the main features of their ecology and 10% (1) and 37% (1-2) respectively. seasonal dynamics using the methods of Conclusions: The diversity of ectopara helminthological testing. sites increases with age. Trichodina sp. and Matelials and Methods: Parasitologi Gyrodactylus sp. were occasionally fo und, cal examination of digestive tracts and their prevalence possibly being underesti stomach contents of 20 specimens of U. mated because these parasites are easily lost aalge and 21 specimens of U. lomvia was during sampling. The prevalence of L. conducted. Helminths were preserved with branchia/is and C. adunca increases with either 70 % ethanol or 4 % neutralised fo r the age of the host. The prevalence and maldehyde. intensity of infection with Caligus sp. larvae reaches a peak at six months of age, fo l Results: 20 % of U. aalge were infected lowed by a marked decrease. The study is with cestodes with a maximum intensity of supported by the Icelandic Research Coun infection of nine worms per specimen, 10 % cil and the Icelandic Republic Fund. with nematodes (about one worm per bird). The population of U. lomvia was much more heavily infected with parasites. 47.6 % of birds were infected with cestodes (55 worms per specimen), and 52.4 % with nematodes (47 worms per specimen). It is noteworthy that in U. aalge, mostly para sites with garnmarids and ammodytids as intermediate hosts were fo und, whereas in U. lomvia the intennediate hosts were euphausiids and capelin. Prevalences and abundances of infection of birds were higher in 1999 than in 1993. 121 Conclusion: The basic diet of U aalge todes. Increasing intensity of infection by in the study area consisted largely of Am cestodes was accompanied by an increase in modytes spp., whereas the capelin repre glucose concentration. The highest content sents the main fo od item fo r U lomvia. of y-globulins was observed in birds in Feeding grounds of the former species are fected with larval cestodes and a somewhat located much closer to the shoreline. The lower content in birds infected with adult infection of birds with helminths increases cestodes. The lowest concentrations of glu in the years characterised by a peak abun cose and y-globulins were fo und in a gull dance of birds, and vice versa. freeof these worms. The work was supported by INTAS Conclusion: The cestodes are the most (project N 97-1 0224). pathogenic group of helminths infecting the seagull populations on East Murman. The highest stress on the immune system of ASSESSMENT OF THE PATHO infected birds was inflicted by the larval GENICITY OF SEABIRD BEL cestodes. MINTHS USING BIOCHEMICAL TESTING The work was supported by INTAS (project N 97-1 0224). M.M. Kuklina and V .V. Kuklin Murmansk Marine Biological Institute, Kola COPROPHAGY AND TRANSPLA Scientific Centre, Russian Academy of CENTAL TRANSMISSION AS POS Sciences, Murmansk, Russia SIDLE WAYS OF DISTRIBUTION OF SOME SARCOSPORIDIAN (PROTIS TA: COCCIDIA) SPECIES IN RO Objective: The pathogenicity of differ DENTS ent groups of seabird helminths was exam ined in 1999 using the methods of bio chemical testing. The study was carried out L. Kutkiene and J. Grikieniene in the Dalnezelenetskaya Inlet (East Mur Institute of Ecology, Vilnius, Lithuania man) in two !arid populations, the black backed gull,Larus marinus, and the herring Objective: Our knowledge of the modes gull, Larus argentatus. of transmission of Protozoa of the genus Materials and Methods: Nine speci Sarcocystis is insufficient. Such studies are mens of L. marinus and three specimens of of prime importance fo r species with a can L. argentatus were studied. Samples of nibalistic mode of transmission. The aim of plasma were extracted from bird blood, in the present work was to clarifY the ecologi which the content of glucose and y cal significance of coprophagy and trans globulins was measured. The birds were placental transmission of Sarcocystis roden also examined fo r parasites, then the rela tifelis. tionships between the parasitological and Materials and Methods: Three ex biochemical characteristics were studied. periments were carried out on 340 Wistar Results: There were no correlations be Norway laboratory rats. Sarcocysts in mus tween the biochemical parameters and lev cle sections were determined microscopi els of infection by nematodes and trema- cally. To detect oocysts or sporocysts, fae ces were examined by differentmet hods. 122 Results: It was determined that the in like particles inside them were noted on the testines of rats infected with sarcocysts of skin of perch (Perca jluviatilis) in three of S rodentifelis became infective for intact fo ur lakes studied in central Finland. They rats during the period fi·om the 4th-8th to were found both on scales and on scaleless 54th-70th day (duration of the experiment). regions. The surrounding tissue was not Intact rats became infected on the 6th to affected, the border between the nodules 80th day after they were fe d on faeces of the and the skin being clearly structured. The rats infected with sarcocysts. Sporocysts occurrence of lymphocystis was greatly dependent on the season of the year. High and oocysts of S rodentifelis were fo und in the faeces of rats infected with sarcocysts. est prevalences were fo und in two subareas This is the firstrecord of the fm ding of Sar of the largest and deepest oligotrophic lake cocystis oocysts and sporocysts in the faeces of the study in March and April ( 60%), of rodents. while nodules were practically absent dur ing the summer and autumn months. The Sarcocysts were not detected in 83 rat seasonality of the occurrence oflymph ocys lings born from eight females fe d on the tis is suggested to be due to spawning stress intestines with oocysts and sporocysts of S of the host in spring and a temperature rodentifelis at different days during preg activated immune response of the host in nancy. summer, causing the disappearance of the Conclusions: The faecal-oral way is nodules. In addition to the seasonal study in important in distribution of S rodentifelis, the largest lake, perch were also monitored whereas the transplacental route of trans from all of the fo ur lakes in May during two mission is ecologically insignificant. subsequent years (1997, 1998). The highest The studies were subsidised by the prevalences were fo und in the oligot:rophic Lithuanian State Research and Higher Edu lake in both years (39%, 34% respectively) cation Fund (1999, Nr. 353). and in the most isolated and smallest fo rest lake (3 1 %, 12% respectively) while in the other forest lake the prevalence was only A NEW DISEASE IN PERCH (PERCA 10% in both years. In the fo urth lake influ FL UVIATIL IS) : OCCURRENCE OF enced by humic substances, lymphocystis LYMPHOCYSTIS IN FINNISH was absent. A study of the localisation of LAKES lymphocystis in five sites in the large oligotrophic lake in May 1997 and 1998 KM. Leskisenoja1, E.T. Valtonen1 and R.W. revealed that lymphocystis occurred abun Hoffinann2 dantly in all areas of the lake (in 1997 1Department of Biological and Environmental prevalence varied between 33-61% and in Science, University of Jyvaskyla, Jyvaskyla, 1998 between 20-45%). Lymphocystis has Finland not been reported earlier on perch. How common it is, and what fa ctors induce its 2Institut flir Zoologie und Hydrobiologie, Tier arztliche Fakultat der Universitat MUnchen, appearance need further studies. Munich, Ge1many Small, white lymphocystis nodules (< 1mm) of hard consistency and with vims- 123 NEW DATA ON THE EPIDEMI with the appearance of the Chaffinch off OLOGY OF BIRD HAEMOPRO spring. C. impunctatus is abundant on the TEIDS ON THECURONIAN SPIT study site and is highly susceptible to infec tion with haemoproteids. The most active G. Liutkevicius transmission of haemoproteids occurs in May-June, and C. impunctatus is the most Institute of Ecology, Vilnius, Lithuania likely vector of haemoproteids on the Curonian Spit. Objective: To detennine possible vec tors of haemoproteids (Haemosporida: Haemoproteidae) on the Curonian Spit and EFFECTS OF DIFFERENT NATURAL to investigate the peculiarities of the vectors PRODUCTS AGAINST SARCOPTIC and the distribution of parasites there. MANGE MITESIN SWINE Materials and Methods: The material was collected in May-August 1997- 1999. E. Magi and A. Kaarma To investigate the fauna of vectors, 295 Institute of Infectious Diseases, Estonian specimens of biting midges (genus Culi Agricultural University, Tartu, Estonia coides) and 106 specimens of hippoboscid flies (genus Ornithomyia) were collected. Objective: Pig mange caused by the To detennine spontaneous infection of vec parasitic arthropod mite Sarcoptes scabiei tors with haemoproteids, the salivruy glands var. suis is a widespread contagious skin of 201 specimens of Culicoides sp. and 106 disease and is recognized as a major con specimens of Ornithomyia sp. were investi tributor to losses in swine productivity. As gated. Twenty five specimens of C. impunc several plants with insect reproductive in tatus were also experimentally infected with hibitors and repellents have long been used Haemoproteus fringillae. Spontaneous infec fo r medical purposes, the possibility of the tion of the Chaffmch (Fringilla coelebs) with use of natural plant extracts against swine haemoproteids was monitored on the study mange mites was investigated. site. Materials and Methods: Laboratoty Results: C. impunctatus was the most tests on the viability of swine sarcoptic abundant species of biting midge (94.6% of mange mites were carried out. Our investi the collected specimens), and 0. avicularia gations were made on the effect of ethereal was the most frequently fo und species oils of tea tree (Melaleuca altemifolia), among hippoboscid flies (47.2%). Culi black pepper (Piperni ghrum), sweet orange coides sp. and Ornithomyia sp. were not (Citrus sinensis), pennyroyal (Mentha spontaneously infe cted with haemoproteids. pylegium), citronella (Cymbopogon nar Sporozoites were fo und in 44.0% of C. dus), eucalyptus (E ucalyptus globulus) and impunctatus, which were experimentally juniper (Juniperus communis) and on ex infected with H fringillae. Over 45% of tractof garlic (Allium sativum). adult and 15% of juvenile Chaffmches were spontaneously infected with haemoproteids. Results: According to the results of our laboratory trials, all the tested plant products Conclusions: The Chaffinch is a com proved to be effective against adult swine mon host of haemoproteids. The highest mange mites: in their 0.5-2% water emul seasonal activity of Culicoides sp. coincides sions the lethality in vitro was registered up 124 to 95-100% in 24 hours. Our data show no cilia were found in the vitelline reservoir. that the most effective preparation against The oviduct wall was elevated in the form mange mites was tea tree oil - even after of lamellae, which were plicated along their only one hour a 1 00% death rate of para length. The ootypewas characterized by the sites was recorded. After three hours com presence of lamellae, cilia and unicellular parisons of means were used to compare the Mehlis' glands. It was shown that the fo r results of different variants. mation of egg shells was implemented by Conclusion: Our results of trials carried the deposition of vitelline globules in their out with various plant extracts led to the surface in the ootype-uterine duct. The re conclusion that on the basis of such studies gional differentiations of the uterus wall it would be possible to develop new bio were recorded. The proximal region of the logical parasite control methods, and that uterus was covered by microlamellae and natural plant products might be used against the distal one by microtriches. Filamentous animal external parasites as alternatives to microtriches were observed on the apical neurotoxic insecticides. surface of thevagina. Conclusion: Structural and functional differences and the origin of different patis STUDIES ON THE FEMALE RE of the fe male reproductive appmatus for PRODUCTIVE SYSTEM OF DIPHYL vmious groups of cestodes are conditioned LOBOTHR IUM LATUM by the biology of each species. Acknowledgments: The study was sup L.G. Poddubnaya ported by the Russian Foundation fo r Basic Institute for Biology ofinland Waters RAS, Research (N 99-04-48465). Borok, Yaroslavl region, Russia Objective: The study was carried out on INFLUENCE OF ENVIRONMENTAL Diphyllobothrium latum (L.) (Cestoda: FACTORS ON SURVIVAL OF TRE Pseudophyllidea). MATODE CERCARIAE FROM LIT TORAL PROSOBRANCHS OF THE Materials and Methods: Adult worms WHITE AND THE BARENTS SEAS: of latum were obtained from an experi D. ANEXPERIMENTAL STUDY mentally infected cat. Plerocercoids of these cestodes were removed from the body cav ity of a pike (Esox lucius) collected from V.V. Prokofiev Rybinsk reservoir. Materials were fixed in Pskov Pedagogical Institute, Pskov, Russia 4% glutaraldehyde in cacodylate buffer, then in Os04, dehydrated fo r TEM. Objective: To study the influence of dif Results: The epithelium of the fe male fe rent water temperatures and salinities on reproductive system ducts consists of a the survival of free-living larvae (cercariae) nucleate syncytial layer. Structural differ of marine trematodes. ences in apical surface of the ducts, the Material and Methods: The average lon number of nuclei and organoids in syncytial gevities (LT50) of cercariae of seven trema layer as well as the number of underlying tode species were studied experimentally at ° ° ° muscles were revealed. Numerous cilia three temperatures (5 C, 10 C, 20 C) and were fo und on the vitelloduct surface, but fo ur salinities (8, 16, 24 and 32%0). Cer- 125 cariae used in experiments were released in PLATELET ACTIVATION MARK the laboratory from naturally infected inter ERS INPARASITIC DISEASES tidal molluscs Hy drobia ulvae and Littorina sa:xatilis. H ulvae infected by Cryptocotyle D. Prokopowicz1, J. Matowicka-Karni and A. concavum (Heterophyidae), Levinseniella Panasiuk1 brachysoma, Maritrema subdolum (Micro 1Department of Infectious Diseases, Medical phallidae), Himasthla elongata (Echi Academy ofBialystok,Poland nostomatidae ), and L. sa:xatilis by Renicola 2Department of Clinical Diagnostics, Medical roscovita (Renicolidae), were collected in the Academy ofBialystok, Poland White Sea, and L. sa:xatilis infected by Po docotyle atomon (Opecoelidae) and Reni cola thaidus (Renicolidae) in the Barents Objective: Blood platelets actively par Sea. ticipate in the immune mechanisms of the Results: The results of the experiments organism (e.g. antiparasitic immunity). showed that temperature resistance and Platelet stimulation may be caused by con salinity tolerance of cercariae from the tact with a parasite, the increase of IgE anti White Sea snails were higher than of those body concentration, and the presence of fr om the Barents Sea. Significant decreases complement and lymphokines. In our stud in the LTso of the «White Sea cercariae» ies we have tried to answer the question of was recorded in water of 8%0salinity and of whether a parasitic infection can cause the the «Barents Sea cercariae» even at 16%0. platelet activation. The rate of LTso decrease with increase in Materials and Methods: Thirty-five water temperature was also lower in the patients (aged 19 - 58) treated in the De fo rmer cercariae than in the latter. partment of Infectious Diseases underwent Conclusion: Cercariae shed from the the examinations. Twenty-one patients were White Sea molluscs are able to survive infected with Giardia intestinalis (Gi) and under a relatively wider range of environ 14 patients with Echinococcus granulosus mental factors (temperature, salinity) than (Eg). Blood samples were taken twice: larvae from the Barents Sea periwinkles. before treatment (AI) and after antiparasitic This might be determined by the fact that therapy (A2). The control group consisted the seasonal and daily fluctuations of water of 33 healthy individuals, aged 19- 45. The temperature and salinity in the intertidal venal blood was transferred to Vacutainer zone of the White Sea are higher than of the testing tubes containing CTAD (citrate, Barents Sea. theophylline, adenosine, and dipiridamol). �-thromboglobulin and PF4 concentrations The work was supported by INTAS were determined with the use of AS (project N 97-10224) and the Russian SERACHROM kit (Boehringer Mannheim) Foundation fo r Basic Research (N 98-09- by the immunoenzymaticmet hod. 49706). Results: Blood platelet secretory activ ity was determined by the release of two proteins (�-thromboglobulin and PF4) out of a-granules. The concentration of these proteins is shown in Table 1 (next page). 126 Parameters Examined group, Control group, (K) p* n=35 n=33 AI A2 Platelet factor 4 20.3±9.4 6.0±3.0 2.27±0.88 Al:K p<0.05* (IU/ml) A2:K p<0.05* Al:A2 p<0.05* �-thromboglobulin 33.6±5.8 7.5±2.8 4.73±2.1 Al:K p<0.05* (IU/ml) A2:K p<0.05* Al:A2 p Table 1. Blood protein concentrations Conclusion: Despite the lack of platelet included in the tests were sampled from direct contact with a parasite, the increase in both in-vitro cultures and fi·om infected concentrations of �-thromboglobulin and rainbow trout ( Oncorhynchus mykiss). 1 PF4 (especially before antiparasitic treat multifzliis was maintained in Blue fin (BF) ment) points to the increased platelet activa cell lines at l5°C using minimal essential tion. medium (MEM). The selected PCR primers A PCR - BASED METHOD FOR THE D-1 (5- ' CTAA TTGTTG GGCTAA T ACA TG-3 ' ), DETECTION OF TE TRAHYMENA D-2 (5'- TCAATGCCGTAGAGAGAAGAATC-3 '), CORLISS/ CONTAMINATION OF and ICHTHYOPHTHIRIUS MULTIFILIIS D-3 (5'-GACGAGTCGITATGAGTCTG-3') ININ VITRO CULTURE were obtained from GIBCO BRL Life technologies (Base!, Switzerland). 1 1 2 D. Pugovkin , T. Wahli and R. Felleisen Results: The PCR test used was fo und 1National Fish Disease Laboratmy, Institute for to be sensitive even for single cell analysis, Animal Pathology, University of Bem, Bem, and an effective discriminator of the two Switzerland species during all life-cycle stages. 2 Institute of Parasitology, University of Bem, Conclusion: This test was fo und to be a Bem, Switzerland rapid and accurate tool fo r the detection and differentiation of the two related para Objective: Development of a PCR sites 1 multifzliis and T corlissi. This pro based method to test in-vitro Ichthyophthir cedure will now facilitate the in-vitro cul ius multifzliis cultures fo r Tetrahymena ture of 1 multifzliis by overcoming the corlissi contamination. initial difficulties encountered in determin ing the contamination status of cell cul Materials and Methods: Primers de tures. rived from the 18S-rRNA gene were tested for specificity and sensitivity fo r detection of 1 multifzliis and T corlissi. Parasites 127 DEPENDENCE OF TELEOSTEI consumed by Antarctic fishes in the south HELMINTH FAUNA FROM ATLAN em Antarctic areas (South Shetlands and TIC ANTARCTIC WATERS ON Antarctic Peninsula). THEffiFEEDING PECULIA RITIES The helminth fauna of Antarctic fishes depends on the features of the Antarctic G. Rodjuk trophic network. Atlantic Scientific Research Institute of Marine Fisheries and Oceanography (AtlantNIRO), THE INFLUENCE OF MAGNETIZED Kaliningrad,Ru ssia WATER ON THE LARVAE OF CULEX PIP/ENS Teleost fishes from South Georgia, the South Shetlands and Antarctic Peninsula 0. Rogachyova waters were examined to determine the Novgorod State University, Novgorod dependence of their helminth fauna (species Veliky, Russia with complex life cycles) on the host's fe ed ing habits. The helminths were collected from fresh and frozen fish specimens in During recent years there has been con 1972-1989. siderable interest in studies of the influence of magnetism on living organisms. Studies In total 1036 specimens belonging to 38 of the influence of magnetic fields on vari fish species (12 families) were investigated. ous biological subjects are increasingly Forty helminth species (11 acanthocepha needed. However, published experimental lans, 7 cestodes, 14 trematodes, 8 nema data are insufficient and do not always re todes) were found (32 species at South flect modem views upon the biological Georgia, 33 at South Shetlands and 19 at effect of a magnetic field. By studying an Antarctic Peninsula). The helminth species organism on the quantum level scientists composition indicates that Antarctic fishes point to the fact that every chemical and fe ed intensively on krill and benthic inver biological reaction is closely bonded to tebrates, such as mysids and amphipods. magnetic energy (Holodov, 1996). It is The highest similarity of fish helminth knownthat the mechanism of the effect of a fauna was observed between South Georgia magneticfield on a living organism is based and South Shetlands (Sorensen on significant changes in the energy of Czekanowski index - 81.8%). Despite the chemical bonds during biological processes. fact that both areas were similar inthe num These researches testifYthat an organism is ber of helminth species, they differed in the a multilevel system which is characterized number of helminth species developing via by selected magneticphen omena. krill as well as in quantitative aspects. The Under laboratory conditions a number of prevalence and intensity of infection by experiments were conducted by us on the helminth species developing via krill in influence of magnetic fields on living creased in the more southernAntarctic areas organisms from unicellular organisms to (South Shetlands and Antarctic Peninsula) larvae of the dipteran Culex pipiens. as compared with South Georgia. The dif ferences in helminth faunas were a result of Methods: In the experiments larvae of an increase in the proportion of euphausiids II-IV chrysalis stages of Culex pipiens, 128 Anopheles messeae and Chaoborinae were techniques (de V os, Dick, 1989). All other used. techniques were as described by 1. Water fr om a natural source was Kuznedelov et al. (1996). The sequences' poured into two vessels and the comparison analysis was made with data second vessel was magnetized using registered in EMBL under the accession a device called a 'magnometer'. numbers Limnadia lenticularis - L81934 2. The same conditions were maintained Cancrincola plumipes - L81938, Calanu; in both vessels: temperature regime, pacificus - L81939, Eucyclops serrulatus - lighting, pH, and intervals of L8 1940, Bemtia purpurea - 26511 , Am observations. blytelus curtus - AF0 12482, Raphetis sp. - The experiments showed that Culicidae AF012485, Mecyclothorax vu/cans larvae are sensitive to a magnetic field. In AFOI2482, Tropopterus sp. - AF012483, magnetized water they were the first to Clambus arnetti - AF012526, Pamborus become excited and move to the upper sur guerinii - AFOI2508, Me codemafulgidum fa ce of the water, which is probably con AFOI2501. Comparative analysis was done nected with changes in the viscosity of the by calculation of evolutionary distances magnetized water. The most important (number of differences) between the se result of the experiment was the speeding quences according to the method of Jukes & Cantor (1969). Evolutionary distance matrix up of metamorphoses of C. pipiens and Anopheles messeae larvae. Therefore, mag obtained in pairwise comparison was used netized water influences the behaviour of to construct dendrograms according to the Culicidae larvae and the process of meta method of Saitou & Nei (1987), showing morphosis. the degree of affmity between sequences compared. TREECON program package (Van de Peer, De Wachter, 1993) was used PRELIMINARY DATA ON NUCLEO fo r comparative analysis. TIDE SEQUENCES OF 18S rDNA OF Results: Dete1mination of nucleotide TWO SPECIES OF CRUSTACEAN sequences in amplified fragments revealed PARASITESFROM LAKEBAIKAL 537 aligned nucleotide positions. Conclusion: The phylogenetic analysis 0. Rusinek1, K. Kuznedelov1 andE. Rusine!(l showed that molecular data of S. thymalli 1Limnological Institute, Irkutsk, Russia and B. briani have I 00 % statistical support. 2IrkutskState University, Irkutsk, Russia These data correlate well with the phyloge netic conception of Kabata (1979), suggest ing that species of the genera Basanisthes Objective: In 1998-99, material from and Salmincola are closely related and that Salmincola thymalli (from Thymallus arcti the first-named genus originated fi·om the cus) and Basanisthes briani (from second. Brachymystax lenok) were sampled from rivers of Barguzinski Nature Reserve to investigate their molecular structure. Materials and Methods: Total DNA was extracted from 96%-ethanol-fixed crus taceans according to previously described 129 COMPARISON OF THE HELMINTH SOME RESULTS OF INVESTI FAUNAS OF BUFO VIRIDIS GATION OF "BLACK SPOT" DIS LAURENTI, 1768 AND B. REGULARIS EASE OF BREAM (ABRAMIS REUSS, 1828 COLLECTED INEGYPT BRAMA) FROM THE CURONIAN LAGOON (SOUTH-EAST PART OF D. Schneider1 and H. Sattmarm2 THEBALTIC SEA) 1Department of Systematic Zoology, Institute of Zoology, UniversityVienna, Vienna, Austria 0. Shugalter 23'd Department of Zoology, Naturhistorisches Atlantic ScientificResearch Institute of Museum Wien, Vienna, Austria Marine Fisheries and Oceanography (AtlantNIRO), Kaliningrad, Russia Objective: Toads of the species Bufo "Black spot" disease of bream fromthe regularis, B. viridis and several frogs from Curonian Lagoon was studied during 1998- Egypt were examined to determine differ 99. A total of3919 specimens (TL=2.8-54.0 ences and conformities in their helmintho cm) were examined. The disease occurred fa unas. In addition fo od analyses were on average in 23 % of all the fishes exam made to investigate an ecological fa ctor ined. The numbers of fishes with black influencing the composition of the helminth spots in samples varied from 4.9 % up to 46 fa unas. %, and the number of spots on a single Materials and Methods: In 1982, 106 specimen variedfrom 1 to 268. anurans from different locations in Egypt Two types of black spot were found. were collected. From dissections of these The first type did not contain cysts with 106 host specimens, Nematoda, Trematoda, metacercariae and occurred within the Cestoda and Acanthocephala were ex whole period of observation. The spots of tracted. The parasites were preserved, the second type, containing cysts with stained and cleared according to standard metacercariae of Posthodiplostomum cuti methods. cola, were only fo und from April through Results: The mean prevalence of infec November. The metacercariae had a large tion was 86.8%. The great majority of ex size range (0.7 - 2.1 mm). Features of their tracted helminths were nematodes. Ecologi allometricgrowth were studied. cal and geographical aspects will be dis The relationship between fish length and cussed. prevalence of black spots was established: prevalence of black spots increased from 0.5 to 46% with increasing fish length from 2.8 to 54.0 cm. The seasonal dynamics of black spot disease of bream were also studied. Only black spots of the first type were fo und from November through April. Living metacer cariae were found on bream from April through October. 130 WHY DO BROWN RATS RAITUS Conclusions: All parasites fo und in the NOR VEGICUS LIVING IN THE survey are known parasites of brown rats, SEWER SYSTEM OF THE OLD although all were rep01ied for the first time TOWN OF REYKJAviK HAVE SO in rats in Iceland in the present study. A FEW PARASITES? feature which all the species fo und have in common is that they have direct life-cycles. K. Skimisson and E. J6hannisdottir A provisional list of brown rat parasites, Institute fo r Experimental Pathology, compiled from surveys carried out in vari University of Iceland, Keldur, Reykjavik, ous countries of the world, includes at least Iceland 20 protozoans, I 0 tJ·ematodes, 7 cestodes, 19 nematodes and 24 ectoparasites - alto Introduction: During the last decade of gether 80 species. Only 11% of these para the 20tl1 century effective pest control meas sites were reported in the present study. ures fm ally exterminated brown rats (Rattus Various factors were considered to ex norvegicus) eve1ywhere in Reykjavik ex plain why so few parasites occur among the cept in the old town where isolated popula sewer rats in Reykjavik. Firstly, potential tions still survive in association with the intermediate hosts like gastropods, crusta sewer system. The nests are usually located ceans and insects are absent in the closed, in recesses beneath buildings where pipes of uniform sewage environment. Therefore, the sewer system are open or damaged. brown rat parasites having complicated life Black rats (R rattus), however, have not cycles have no possibility of being main occurred in Reykjavik fo r decades. tained in the population. Secondly, in past Objective: To study the parasite fa una decades intensive control measures have of the brown rat in the sewer system. been carried out during the summer months whereby baits containing rodenticides have Materials and Methods: A total of 24 been systematically put into manholes of subadult and adult brown rats, trapped the sewer system. As a result the brown rat within houses in the old town of Reykjavik population has temporarily been conti'OIIed where damaged sewers or open water traps every year, and even eradicated in some had enabled rats to enter the buildings, were parts of the sewer system. As a conse examined fo r intestinal endoparasites and quence, rare parasites could also have been ectoparasites. driven to extinction. Thirdly, due to the Results: Nine parasite species were effective pest control measures no rats occur fo und: The protozoans Eimeria ja lciformis, anymore in Reykjavik outside of the closed E. nieschulzi and Cryptosporidium parvum; sewer system. Therefore, import of «new» the cestode Hy menolepis nana; the nema parasites from other populations into this todes Trichuris muris, Syp hacia sp. and closed environment seems veryunlikely. Heterakis sp umosa; the mite Radfordia af finis and nit of a louse, probably Polyp lax sp inulosa. 131 HOST SPECIFICITY OF GAMASID the southern part of the studied region (e.g. MITES (GAMASINA: SPINTURNI Barbastella) are infected by a smaller num CIDAE, MACRONYSSIDAEJ FROM ber of gamasid species. Bats inhabiting the BATS (CHIROPTERAJ northern part of the region are infected by a considerably larger number of gamasid M.K. Stanyukovich species. In mixed colonies of bats the diver sity of parasites is richer than in monospe Zoological Institute, Russian Academy of cies colonies. There appears to be a weak Sciences, St Petersburg, Russia correlation between the numbers of spin turnicids (permanent parasites) and macro Objective: Analysis of host specificity nyssids (some are permanent [Macronys of gamasids (Gamasina: Spinturnicidae, sus], some are temporary parasites [Steato Macronyssidae ), parasitizing bats (Chirop nyssus, Ornithonyssus ]). The species com tera) in Russia and adjacent countries. position of gamasids on bats of the same Materials and Methods: About 16 000 species may be different in different popula gamasids of 42 species (own material and tions. This depends on certain characteristics collection of the Zoological Institute, Rus of host populations or colonies, i.e. geo sian Academy of Sciences) were collected graphical position, other species of bats in a fr om more then 2000 bats belonging to 34 mixed colony, season, migrationroutes, etc. species. Results: Monoxenic species: Sp inturnix IS THE MONOGENEAN ANCYRO acuminatus (on Ny ctalus noctula), S.psi (on CEPHALUS PA RADOXUS CAPABLE Miniopterus schreibersii), S. emarginatus OF REGULATING THE POPU (on M emarginatus), Eyndhovenia euryalis LATION STRUCTURE OF 0+ CLASS euryalis (on Rhinolophus euryale ), Ma cro OF STIZOSTEDION LUCIOPERCA ? nyssus barbastellinus (on Barbastella bar bastella), etc. Oligoxenic species: S. myoti 1 (on My otis), Paraperiglishrus rhinolophinus V.K. Starovoitov and P.I. GeraseY 1 (on Rhinolophus), etc. Polyxenic species: M Atlantic Research Institute of Fisheries and cyclaspis, Steatonyssus sp inosus, Orni Oceanography, Kaliningrad,Russia thonyssuspipi strelli, etc. 2Zoological Institute, Russian Academy of Sci Conclusion: Very close relationships ences, St. Petersburg, Russia between gamasids and their hosts - bats - have led to the appearance of monoxenic Objective: In the Kurish Bay (Curonian and oligoxenic species. The number of Lagoon) of the Baltic Sea, mature fo rms of polyxenic species is considerably smaller. the typical gill monogenean Ancyrocepha All polyxenic species of gamasids are from lus paradoxus were found parasitizing the family Macronyssidae. The latter parasi zander on the isthmus (lower surface of the tize bats of different species in one colony. head between opercula) in mature (4-14 +; All mites of Spinturnicidae are monoxenic 5%) , but mostly in young (1-3+; 54%) fish. or oligoxenic. The fauna of parasitic gama Materials and Methods: Fishes of 0+ sids is richest on l'vfyotis mystacinus (17 class (N=454) were collected in 1997 from species), M daubentoni (15), M brandti May through October. The sample of hosts and Plecotus auritus (14). Bats inhabiting was split into 10 mm size classes. 132 Results: In the total sample, size group 4300 cyprinid specimens were caught: II (3 1-40 mm) had 2.4% prevalence and the Abramis brama (L.) (2953), Rutilus rutilus lowest intensities of infection (abun (L.) (1022), Carassius carassius (L.) (151), dance=0.45), with the propmtion of worms Pelecus cultratus (L.) (164), Vimba vimba infecting the gills being vety high ( 60% ). (L.) (8). The frequency of occurrence of Group V (61-70 mm; A= 10.3) had 36% external tumours, their types and distribu prevalence, with the propmtion of worms tion on the fish body, fish size, sex and age, being minimal on the gills (19.3%) and and internaltumours were all recorded. maximal on the isthmus (more than 80%). Results: Prevalence of tumours was Prevalences in groups VI and VII (71-90 19.9% in bream, 9.0% in roach, 6.0% in mm) decreased (12% and 4%), intensities of crucian carp, and only 0.6% in P. cultratus. infection increased (A=23; 26.5), and the Seven morphological types of tumours were proportion of worms on the gills was 20%. recognized: type I - epithelium-like non Conclusion: It can be concluded that pigmented, II - epithelium-like pigmented, survival of the host and its parasite depends m - papilloma-like non-pigmented, N - on a decrease in the proportion of worms papilloma-like pigmented, V -lipoma-like, parasitizing the gills and on the possibility VI - carcinoma-like, VII - tumours of in of monogeneans locating on the isthmus. ternal organs. Bream had all types of tu Fishes of the 0+ age class with wonns mours, with type I predominant (68.3%). In mostly on gills may suffe r fi"om anaemia, roach, types I (62.5%) and II (37.5%) pre delay in growth, etc., and then may be dominated. In crucian carp, only types N eliminated by cannibalism. Monogeneans in (1 00%) and m (11.1%) were fo und. the parasite/0+ host relationship apparently Analysis of infections in bream and act as a factor in regulating the population roach by tumour occurrence in different age of middle size groups. classes showed 50.0% type I tumour occur rence in class 1-3; 22.7% in 4-6, 18.2 % in TUMOUR-LIKE INFECTIONS OF 7-9; 4.5% in 10-12; 2.3% in 13-15, 2.3% in CYPRINID FISH (CYPRINIDAE) 16-18. FROM CURONIAN LAGOON (THE Conclusion: Infections of different BALTIC SEA) types of tumours in bream and roach are similar. Crucian carp differs in the nature R.V. Sudarev and V.K. Starovoitov and degree of infection, apparently related to the degree of relationship between spe Atlantic Research Institute of Fisheries and cies, different infection nature and host Oceanography (AtlantNIRO), Kaliningrad, Russia population density. All three species are benthophagous; P. cultratus is planktono phagous, tumours in it being rare. Objective: In the south-eastern Curonian Lagoon five cyprinid species were examined for tumours from September 1998 through May 1999. Materials and Methods: Fishes were caught by nets (mesh 40 and 70 mm) at 800-1500 m from the shoreline. A total of 133 DISTRIBUTION OF MOSQUITOES DYNAMICS OF ABUNDANCE OF (DIPTERA: CULICIDAE) IN DIF HYS TER OTHYLACIUM ADUNCUM FERENT HABITATS OF URBAN (NEMATODA), A PARASITE OF TERRITORY SPRAT IN THEBLACK SEA N. Tereshkina and M. Trukhan L. Tkachuk Institute of Zoology, National Academy of Department of Ecological Parasitology, Institute Sciences, Minsk, Belarus of Biology of the Southern Seas, Sevastopol, Crimea, Ukraine Objective: To study the influence of ur ban pressure upon the fauna and larval Objective: Long-term studies of infec stages of mosquitoes. tion of sprat, Sp rattus sprattus phalericus Materials and Methods: In 1996-99 with larvae of the nematode Hy sterothy larval stages of mosquitoes were fo und in lacium aduncum have shown increasing different parts of Minsk territory, such as intensity of infection from single specimens suburban areas, park zones, green areas of in 5-6 cm fish to 317 in 12 cm fish. housing estates and the centre of the city. Material and Methods: Sampling was Larvae and imago captures were carried out carried out from vessels along the Crimean using entomological and water nets. coast from 1976 through 1999. Fresh and Results: Of 39 species of mosquitoes frozen fish of 5-12 cm length were studied. occurring in Belarus, 25 have been recorded Standard parasitological methods were in Minsk. Fourteen species which were not used. Tests of statistical significance on fo und in the city belong to Holarctic and mean differences were conducted. Palearctic fauna and are not common. Ur Results: The nematode H aduncum ban pressure caused a change in the compo uses sprat as a transport host. Adult forms of sition of dominant species. C. pipiens, A. this parasite were fo und in the stomach and dorsalis, A. jlavescens and A. excrucians intestine of about 20 Black Sea fish species. were dominant in urban territory, while A. In sprat (2-12 mm), larvae were fo und on communis, A. cantans and A. excrucians the pyloric caeca and in the abdominal cav were the most numerous in Belarus. Eight ity. Intensity of infection increases with fish water body types ofnatural origin and fo ur length. Specimens of the 5-6 cm size group types of man-made larval habitats have were either not infected or infected with a been recorded in the Minsk region. The single worm. In the largest sprats (8.1-9.0 highest densities of larvae were in tempo cm) parasite abundance was greater than rary water bodies in park zones and in twice that of fish in the 7.1-8.0 cm group. basement of houses. Intensity of infection in similar-sized fish is Conclusion: Urban pressure caused a always similar. In the first and second decrease in the number of species to 64%, length-groups, parasites were fo und on the influenced the composition of dominant pyloric caeca, in the third length-group on species, decreased the number of species the pyloric caeca and in the abdominal cav and changed the types of larval habitats. ity. From 988 sprats 800 were examined immediately after capture, in the region of the fishery. Another 188 sprats were dis- 134 sected after freezing or pickling. In the first material was sampled yearly from May group the parasite distribution was normal, through September in Karantin Bay isolated whereas in the second almost all the inner from the sea at a site of unftltrated urban organs were infected. The greatest number wastes discharge, and in the esturuy of of worms was observed in those organs and Omega Bay with good water flow and low tissues with the greatest fat content. No pollution. Standard parasitological methods significant differences in mean lengths of were used in the work. parasites were fo und. Results: Shrimps were infected with Conclusion: Sprat habitats distant from parasites in both regions. Actinomyces sp. the shore in the Black Sea facilitated preser inhabits the cuticular shell of all the vation of established host-parasite relation shrimps' organs, but most often the cara ships. Coastal marine communities suffer pace, epimeral plates and uropods. The most from pollution. Parasite abundance infection level of cuticular shell in Karantin increased with fish length. Changes in the Bay exceeded that from Omega Bay three condition of the fish before examination fo ld. T lenticularis pru·asitizes the thickness (frozen or salted) caused an increase of of the shrimp's cuticle. It was abundant in parasite activity, which was expressed by rostrum cuticula, cru·apace, genital segment, changes in distribution. and uropods. Infusoria infection in Kru·antin Bay was two times higher than in Omega Bay. Trematode metacercru·iae occur more THE INFLUENCE OF AN ENVIRON often in the anterior muscles of of shrimps, MENTAL FACTOR (POLLUTION) less often in the muscles of posterior seg ON ABUNDANCE OF THE DOMI ments and swimming legs. Its intensity of NANT PARASITE SPECIES OF infection in Karantin Bay exceeded that in SHRIMP PALAEMON ELEGANS Omega Bay 6-1 6-fold, and its prevalence 2- RATHKE IN THESEA BLACK 6-fold. Intensity of infection of all parasite species in similar-sized shrimps (50-60 L. Tkachuk mm), according to the long-term data, Department of Ecological Parasitology, shows a considerable difference between Institute of Biology ofthe Southern Seas, the two regions. Isopods occwTed in 11.3% Sevastopol, Crimea, Ukraine of shrimps in 1970 and 1.6% of slu·imps in 1975, but from 1997 up to the present time Objective: Monitoring of the parasite these parasites have not been fo und. fauna (1995-1999) of shrimp Palaemon elegans Rathke, 183 7 in two differently Conclusion: The disappearance of B. polluted bays near Sevastopol showed sharp squillarum led to a decrease of diversity of increases in the numbers of three parasite P. elegans parasites. A sharp increase in the species: the fungus Actinomyces sp., the number of dominant parasite species took infusorian Terebrospira lenticularis and the place because they (fungi, infusoria) or their trematode Helicometra fasciata, and the hosts, taking part in the trematode life cycle, disappearance of the previously common are chru·acterized by adaptation to ru1 abun isopod species Bopyrus squillarum. dance of organic renmants and low oxygen content in the water. But it should be noted Material and Methods: About 3000 that the extent of organic pollution deter slu·imps 27-65 mm in length, caught at mines this situation. depths of 0.5-1.0 m, were examined. The 135 TENDENCY FOR A POLYCYCLIC albula. It may be hypothesized that the SEASONAL DEVELOPMENT OF changes in the fe eding activity of the defini PR OTEOCEPHAL US CERNUAE IN tive host depending on the type of water THE RYBINSKRESERVOIR body affects the biologyof the cestodes. AV. Tyutin REGULARITIES IN THE DISTRI Institute for Biology ofinland Waters RAS, BUTION OF LETIIAL HAEMOSPO Borok, Yaroslavl region, Russia RIDIAN PARASITES OF POULTRY Objective: To evaluate the level of in G. ValkiiinasAND T. Iezhova fe ction of ruff ( Gymnocephalus cernua) by the cestode Proteocephalus cernuae Institute ofEcology, Vilnius, Lithuania (Gmelin, 1790) and to study its seasonal maturation in the Rybinsk Reservoir, one of Objective: To specifythe regularities of the largest reservoirs in the world. distribution of haemosporidians (Protista: Haemosporida) which cause lethal diseases Materials and Methods: In 1992-1994 in poultry. a total of 716 specimens of fish, mainly from the Volga part of the reservoir, were Material and Methods: The original data captured and examined. of a long-term period of investigation by the authors using traditional methods as well as Results: Seasonal variation in preva analyses of collections and literature data. lence of the tapeworm was observed (win ter: 51.2±2.9, spring: 60.5±3 .9, summer: Results: So far, 210 species of 48.6±3.7, autumn: 37.7±5.5%). Unlike P. Haemosporida are known to parasitize percae, the proportions of adult P. cernuae birds. Only 14 of them are of practical sig in the total worm population appeared to be nificance, among them one species of Hae greatest from May to September (maxi moproteus, eight of Plasmodium, and five mums in June and August). A certain num of Leucocytozoon. Approximately 80% of ber of these maturating cestodes were found the practically important species parasitize even in winter: 6.0±1.0% (spring: 19.3±2.2, galliform birds. H mansoni (=H melea summer: 62.9±4.1, autumn: 34.1±7.2%). gridis) and L. smithi cause lethal diseases in The ratio of variance to abundance of adult turkeys in the south east (SE) of the USA, worms was always about 1, therefo re the L. simondi in ducks and geese in the North distribution of this group of helminths was em Holarctic, L. caulleryi in hens in SE uniform. A two-month delay was observed Asia, L. struthionis in ostriches in S. Africa, between peaks (May and October) of P. P. gallinaceum in hens in the Oriental re cernuae adults in comparison with overall gion, and P. juxtanucleare in hens in the prevalence and abundance. tropics worldwide except for Australia. Lethal haemosporidioses are totally absent Conclusion: There are at least two para in Australia and rare in S. America, but they site generations with maturation periods in may be introduced. winter-spring and summer-autumn. Similar trends in the polycyclic seasonal develop Conclusions: Haemosporidian parasites ment were previously observed by many of poultryare clearly spotty in their distribu authors fo r P. exiguus from Coregonus tion, and the majority oftheir species parasi tize galliform birds which are the main 136 source fo r introduction of the diseases into also important in detetmining the preva new territories. Leucocytozoonosis is the lence and intensityof infection. main practically important haemosporidio Conclusions: In general fishes of the sis. Volga and the Gulf of Finland are highly infected with P. ovatus. The northern border DISTRIBUTION OF PARACOENO of this parasite's distribution coincides with GONIMUS OVATUS KATSURADA, the distribution of its first intermediate hosts 1914 INFISHES OF RUSSIA - the molluscs Viviparus viviparus and V contectus. But the finding of P. ovatus in the rivers Amur and Ittysh do not coincide 1 1 1 V. Voronin , R. Kudenzova , Y. Strelkov , N. with the area of these molluscs. There is an Chemyshova\ E. Golubeva1, A. Zhokhov, M. opinion that P. ovatus is a complex of sev Safronov1 and V. Fetisov1 eral species. 1State Research Institute fo r Lake and River Fisheries, St. Petersburg, Russia 2Institute for Biology of Inland Waters RAS, OCCURRENCE OF MICROSPO Borok, Russia RIDIA AND SCHIS TOCEPHAL US SPP. IN STICKLEBACKS OF THE GULF OF FINLAND NEAR ST. PE Objective: Paracoenogonimus ovatus TERSBURG: RESULTS OF LONG Katsurada, 1914 is one of the common TERM OBSERVATIONS parasites of many species of freshwater fishes in Europe and in Asia. Definitive hosts of P. ovatus are different fish-eating V.N. Voronin and M.E. Safronov birds and carnivores (experimental data). State Research Institute for Lake and River Fisheries, St. Petersburg, Russia Materials and Results: During 1999, 49 1 muscle samples of 18 fish species from the Gulf of Finland and the Volga River as Objective: The three-spined stickleback well as literature data were analyzed to Gasterosteus aculeatus L. and nine-spined study the ecological aspects and geographical stickleback Pungitius pungitius (L.) are distribution of P. ovatus in Russia. Among commercial fishes of the Gulf of Finland. cyprinids the highest prevalences and intensi Catches of these fish near St. Petersburg ties of infection were observed in roach during 1972-1975 were approximately 3000 (Rutilus rutilus), bream (Abramis brama), tons per year but only 300 tons per year silver bream (Blicca bj oerkna) and ide (Leu during 1998-1999. Two new microsporid ciscus idus). The lowest levels of infection ian species, Glugea gasterostei Voronin, were observed in pelagic fishes: bleak (Al 1974 and Thelohania baueri Voronin, 1974 burnus alburnus) and ziege (Pelecus cultra were detected in sticklebacks of the Gulf of tus). Among percids the occurrence of P. Finland. ovatus was low in contrast to cyprinids and Materials and Results: Of 1659 G. minimal infestation was observed for perch aculeatus specimens examined during (Percajluviatilis). The predatory fishes pike 1972-1975 36 (2.2%) were infected by (Esox lucius) and asp (Aspius aspius) were Glugea gasterostei and 54 of 1084 (5.0%) also infected. Ecological conditions in the fe male specimens were infected by Thelo water reservoir or its different localities are hania baueri. At the same time, 135 of 1659 137 G. aculeatus (8.1%) and 81 of 924 P. fo rest in Silute districtand from fo rest and a ptmgitius (8.8%) were infected by plerocer river pond in Sakiai district. The specimens coids of Schistocephalus spp. During 1998- were examined by the method of total 1999 (about 25 years later) sticklebacks parasitological autopsy. from the same localities were reinvesti Results: Six species of cestodes and gated. G. gasterostei was fo und in 2.8% of three species of trematodes were found in Gasterosteus aculeatus and T baueri in Sorex minutus, fo ur species of cestodes 3.0% of females of 425 fish specimens. were found in Sorex araneus, two species of Schistocephalus solidus was found in trematodes were found in Neomys fo diens, 25.5% G. aculeatus (60 of 235) during and one species of cestode in Talpa eu autumn 1998 and in 40.0% specimens (76 ropaea. The prevalence of infection was of 190) during autumn 1999. 86.96% in Soricidae and 7.69% in Talpidae Conclusions: Our long-term observa Conclusion: All of the platyhelminths tions demonstrated the different models of tended to occupy a specific site in the host parasite-host relations fo r Microsporidia and gastrointestinal tract. The cestodes plerocercoids of Schistocephalus solidus. In Neoscrjabinolep is schaldybini and Mo llus general the permanent low level of Micro cotaenia crasiscolex were the dominant sporidia infection of sticklebacks is con playthelminth species. Trematode species firmed. On the other hand, the significant were recorded from Neomys fo diens and increase of S. solidus infection of three Sorex minutus only in Sakiai district il1 wet spilled stickleback during recent years is places where molluscs are abundant. unusual. The reasons may be changes in the ecological situation as a result of dam con struction or the effect of low fishing effort ANTHROPOGENIC PRESSURE IN on the host population. FLUENCES ON IXODES TICK POPULATIONS THE PLATYHELMINTHS OF IN 2 SECTIVORES INLITHUANIA S.D. Zharkov\ P.M. Jensen\ H.V. Dubinina and A.N. AlekseeY 1Royal Veterinary and Agricultural University, R. Zqsityte Copenhagen, Denmark Institute ofEcology, Vilnius, Lithuania 2Zoological Institute, Russian Academy of Sci ences, St. Petersburg, Russia Objective: Plathelminthes of Lithua nia's insectivores: Sorex araneus, Sorex minutus, Ne omys fo diens, Talpa europaea, Objective: About 3000 Ix odes ticks previously collected and fixed in 70% etha Erinaceus concolor, were studied for the nol, and 3000 examined as live specimens, first time. were investigated for detection of exoskele Materials and Methods: Twenty three ton abnormalities. Ticks were collected specimens of shrews (Soricidae), thirteen from different parts of their area within a specimens of moles (Talpidae) and one nineyears period ( 1990-1 999) by flagging. specimen of hedgehog (Erinaceidae) were Materials and Methods: Abnormalities collected from June to October in 1999 were detected using a stereomicroscope. from a town park in Vilnius, the edge of a 138 Dark field microscopy, IFA and PCR meth persulcatus tick populations greatly de ods were used to detect infections. Every pends on anthropogenic pressure. Anoma specimen was investigated to determine the lous ticks differed from normal ones in type of infection by different pathogens many parameters and were more suscepti (e.g. Borrelia, Ehrlichia and tick-borne ble to infection. Exoskeleton abnormality encephalitis virus). Types of abnormalities research might be one of the reliable ways were described previously (Alekseev & of biomonitoring. Further study of abnor Dubinina, 1996). mality-causing agents is needed. Results: The minimal level of abnor Acknowledgements: The research de malities ( 10%) was fo und in protected areas scribed in this publication was made possi (Zhiguli Preserve, Russia and Kingelund ble by Grant 9600864 Danish Research Forest, Denmark). The maximum (50%) Council. was observed in the Cherepovets area (Vo logda Region, Russia). The activity of anomalous ticks differed from that of nor References mal ones, and their infection rate was higher Alekseev AN, Dubinina HV. Some than that of normal ticks. aspects of mite (Oppiidae) and tick (Ixodidae) pathology as a result of Conclusion: The amount of exoskeleton antropogenic pressure. Acarology IX. abnonnalities in Ix odes ricinus and Ix odes Columbus, Ohio 1996; 1: 117-20 NEWS Minutes fr om the SSP Board telephone meeting 28. November, 2000: Present: Tellervo Valtonen, Jorun Tharaldsen, Ingela Krantz, Karl Skirnisson, Mats Wahlgren and Maria Vang Johansen Repmier: Maria Agenda fo r the meeting: 1) Report from the meeting in St Petersburg (Te llervo) 2) Next SSP meeting including connected nematode meeting, The history of SSP presentation by Jorn Andreassen and young scientist support (Ma ts) . 3) Financial status of SSP and new members. Each local treasure should report. 4) New president fo r SSP to be selected in Stockholm in 2001 (Te llervo). 5) Report from the General Assembly of the European Federation of Parasitologists (Karl). 6) Information on new initiatives on parasitological courses in Denmark (Ma ria) 7) Other issues. 139 Ad 1) Tellervo informed that the joint meeting of The Russian Soc. for Parasitology and SSP in St Petersburg in July 2000, had been very successful, with many interest ing speeches and discussions and very well organised. A total of 121 people from 25 countries patiicipated. Jorun informed that the next issue ofthe Bulletin containing the abstracts (150 pages) will be fi nished shortly and 2/3 of the publication expenses will be covered by extra mural fu nding. Ad 2) Planning of the 201h SSP meeting to be held in Stockholm in 2001 is going very well. The meeting will be held at the Karolinska Institute in Stockholm. As the SSP meeting is planned in connection with a special meeting on 'free-living and parasitic nematodes' (organised by Hans P. Fagerholm) as well as a course on 'antigen varia tion', many speakers can contribute without payment. Hans Peter has now got funding fo r his meeting. The 2nd announcement will come out soon. It was agreed to ask J0rn Andreassen (Denmark) to give an anniversary-talk about the 30 years of SSP at the meeting in Stockholm, although the actual anniversmy is in 2002. Karl told that he had poster presented the 1st announcement of the SSP meeting 2001 at the EMOP meeting in Poland this summer. It was decided that the meeting should be announced in Parasi tology Today, Journal of Helminthology, SSP Bulletin and also communicated to na tional parasitological and tropical medicine societies (Norwegian Vet. Ass., Swedish Trop.Soc. etc.). Mats will take action on the international announcements. Mats also informed about a very informal meeting among Swedish parasitologists last week. Eighty persons participated, but no abstracts were provided. Mats will give Jorun a sh01i report from the meeting fo r the Bulletin. Ad 3) In order to get an updated list of our members Tor and Maria should communi cate as soon as possible. Before acceptance of new members the candidate should be presented to the board and their names published in the Bulletin. The financial status IS: Norway: 15,000 Norwegian Kr. Denmark: 9,290 Danish Kr. (26 persons paid in 2000) Iceland: 2,900 Norwegian Kr. (6 persons paid in 2000) Sweden: 2,000 Swedish Kr. Finland: 4,300 Fmk (26 persons paid in 2000) Ingela reported that there had been some problems with the Swedish giro account, but she would look in to this. It was agreed upon that the SSP meetings have to be self-supporting and that SSP in connection with meetings only provide money for travel and attendance stipends fo r young scientists. The membership fe e is 25 EURO fo r ordinary members and 1 0 EURO fo r students. Ad 4) The board discussed possible new candidates to replace the president and the treasurer of SSP from autumn 2001. 140 Ad 5) Karl reported fr om the board meeting of the European Federation of Parasitologist (EFP) in Poznan in Poland in September this year where he represended SSP. 19 societies of the 29 member societies were present at the meeting. The new board of EFP were elected as fo llows: President: R. Houin fr om Creteil in France, (Echinococcus) Secretary: Y. Gi.iri.iz fr om Izmir in Turkey, (Echinococcus) Treasurer: J. Sluiters fr om Rotterdam in the Netherlands (Giardia) Vice-president: H. Wedrychowicz from Warshaw in Poland (DNA Waccination) Additional board members are: J. Corba from the Slovak Academy of Sciencss in Kosice (Echinococcus) Andrey Alekseev from Saint Petersburg ( ectoparasites) Tor Bakke from Oslo. The fo urth candidate, Kolarova from the Czech Republik parasitological Society only got 7 votes and was not elected. Auditor is C. Holland from Dublin in Ireland. Finally it was accepted by all representatives that the next EMOP will be organized during the third week of July in 2004 in Valencia in Spain. No other candidates asked to organize this meeting but the British Society had also shown some interest in some preliminary stages. Ad 6) Maria informed that a 2-year Master programme in parasitology is being planed at the Royal Veterinary and Agricultural University (KVL) in Copenhagen. The course will be in English and be announced internationally. Additionally, a 3-week summer course in parasitic zoonoses will be announced in the beginning of next year. The summer course has been sponsored by NOVA. Maria will give more details as soon as possible to Jorun fo r the Bulletin. Jorun asked fo r written information for the Bulletin about the 2 new research professorships in parasitology at KVL as well as information about the new leader of the Danish Centre fo r Experimental Parasitology, Darwin Murrell. Maria answered that she had asked the relevant people to write about it. She will remind them again. Ad 7) Tellervo mentioned that a one-week course in 'advances in parasite ecology' will be offered at the International Summer School at University of Jyvaskyla, Finland next autumn. Mats informed about 2 new books he is involved in that are coming out soon. One is about 'parasites of the Nordic areas' and the other is a 'Method book in malaria re search'. Jorun informed that parasitic zoonoses are currently receiving a lot of attention in Norway and a recent meeting on the issue attracted 120 participants. Tellervo told that the Baltic Society for Parasitology had asked SSP to make a joint meeting in 2003 in Vilnius. It was then discussed whether our regular meetings (which should be in Norway in 2003) should be held in a Nordic country or whether it could be a part of a joint meeting outside the Nordic countries. The board all agreed that 1 41 there would be no problem in holding the meeting in Vilnius, but it was an issue to be decided at the General Assembly. Tellervo will write to the Baltic society with a copy to Maria. 29.1 1.00 Maria Vang Johansen Book information Methods in Malaria Research is a collection of a variety of protocols in malaria research. The book contains 73 pages of protocols and was edited by Martha Schlich therle and Mats Wahlgren (Karolinska Institutet, Stockholm Sweden), Hedvig Perlmann (Stockholm University, Sweden) and Artur Scherf(Institut Pasteur, Paris, France). Protocols were contributed as a collaborative effo ti from labs of the editors as well as the MR4 at ATCC (Manassas; Virginia, USA) and KEMRI/CGMRC I Well come Trust Research Unit (Kilif'i,Kenya). The book aims to be an all-round collection of practical protocols that students and researchers of malaria can use as they conduct their research. Some of the high-lights include methods in: the handling of parasites (such as culturing, freezing, thawing, staining, purification, micromanipulation, fixation, sampling of blood in the field), immunochemistry (surface iodination and metabolic labeling), serology (immunofluo rescence, ELISA, agglutination assay), cellular methods (parasite antigen preparations, T-cell proliferation assay, cytokine RT-PCR), molecular biology (gDNA preparation, RNA- and Northern preparations, cDNA library construction, single cell RT-PCR, PCR species identification, pulsed-field gel electrophoresis, recombinant protein ex pression, transfection protocols). To make future editions of this book of protocols even more complete, we hope that our readers will want to contribute their own well working methods. A criterion we have set to include a protocol, is that it be well-tried by the lab or person contributing it. We aim to maintain this idea in the future and thus wish fo r the book to be a living document of practical malaria methodology. Methods in Malaria Research is now available as an internet pdf-document on the fo llowing address: http://www.malaria.mr4.org/mr4pages/MR4 Protocols.html Parasitological activities in Sweden The second Swedish Parasitology Meeting (SPAM) was held at the Swedish Insti tute of Infectious Disease Control in Stockholm on the 17th of November. The initia tive to meet among parasitologists in Sweden had come from the Veterinary parasi tologists in Uppsala and the first meeting was held 1999 in Uppsala. This year was an 142 expanded version of SP AM where we had talks, among others, about the infectious biology of giardia, malaria, scabies, toxoplasma, and the genome sequencing project of Trypanosoma cruzi which is taking place at Uppsala Unviversity. Mats Wahlgren Parasitological activities in Norway A course in Serious, Contagious Diseases in Animals was held in Oslo, September 3-8, 2000, jointly organized by the Norwegian School of Veterinary Science, the Na tional Veterinary Institute, VESO, the Faculty of Veterinary Science of the University of Pretoria and the Onderstepoort Veterinary Institute of South Africa. There were participants from Denmark, Sweden, Italy/Ethiopia and Norway. This course also included a session on zoonotic parasitic diseases, with a lecture by T. Krecek (An African perspective), and J. Tharaldsen (Nordic Perspective). The Norwegian Zoonosis Centre organized a one-day Seminar on parasitic Zoonoses at the National Veterinary Institute, Oslo, November 8, 2000. The seminar was very well attended, with more than 120 participants. Jorun Tharaldsen 143 GUIDELINES FOR CONTRIBUTORS All contributions should be submitted as word-processed manuscripts on floppy disk, accompanied by two exactly matching print-outs of good reading-quality. The preferred storage medium is a 3\12inch disk in MS-DOS or Windows compatible fo rmat. The text should be written in Word or WordPerfect or other word processing programs converti ble to these. With a Macintosh computer, save the file in the MS-DOS compatible option. Please indicate the word processor (and version) used to generate the file, the type of computer, the operating system, and the fo rmatted capacity of the diskette. The atiicles/communications should normally not exceed 4 printed pages, including tables, figures, and references, and may contain a maximum of 2000 words if there are no figures or tables. The first page should show the title of the article, and the name(s) of the author(s). The authors' addresses should be given, and the complete correspondence address with telephone and telefax number (if available). The text should fo llow, with out subheadings, but a sh01t summary, maximum 100 words, may be included. The text should be typed unjustified (unaligned right margins), without hyphenation (except fo r compound words), and at 1 \12 line spacing. Do not typepage numbers. Label the hard copies by hand at the bottom of the page. Please ensure that the digit 1 and the letter '1' have been used properly, likewise with the digit 0 and the letter '0'. Do not use decorative fo rmatting, such as boldface and centred headings, or underlining of titles or subheads. Authors are obliged to fo llow the rules governing biological nomenclatures, as laid down in e.g. the In ternational Code of Zoological Nomenclature. Disease names should fo llow the principles of Standardized Nomenclature of Parasitic Diseases (SNOPAD). Figure legends must be included on the diskette. If possible, computer made figures should also be included, but good print-outs should be provided with the manuscript. Otherwise the figures will be handled conventionally. They should be marked on the back with the title of the article and name of the (first)auth or. Line drawings should be provided as good quality hard copies suitable fo r reproduc tion as submitted. 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 are more than two authors, only the first name plus et al. is given (Lund-Larsen et a!, 1977). The reference list should be in alphabetical order, and fo llow the style set forth in Un iform Requirements to Manuscripts Submitted to Biomedical Jo urnals, Br Med J 1988; 296: 401-5. References to journals should contain names and initials of the au thors, article title, the abbreviated name of the journal, year of publication, volume, and first and last page numbers of the paper. Journals should be abbreviated according to the 144 "List ofjo urnals indexed in Index Medicus". Authors without access to this list may type the fu ll name of the journal, and the Editor will take care of the abbreviations. If there are more than six authors, list only the first three and add 'et al'. Personal communica tions and unpublished data should not be used as references, but may be inseiied in the text (within parenthesis marks). Examples of correct forms of references are given below: Standardjournal article: Anonymous. Some fa cts on small animal practice. Vet Rec 1987; 120: 73 Horsberg TE, Berge GN, Hoy T et al. Diklorvos som avlusningsmiddel fo r fisk: klinisk utpmving og toksisitetstesting. Nor Vet Tidsskr 1987; 99: 611-5 Lund-Larsen TR, Sundby A, Kruse V, Velle W. Relation between growth rate, serum somatomedin and plasma testosterone in young bulls. J Anim Sci 1977; 44: 189-94 Books and other monographs: Austin B, Austin DA. Bacterial fish pathogens: disease in fa rmed and wild fish. Chi chester: Ellis Horwood, 1987 McFerran JB, McNulty MS, eds. Acute virus infections of poultry: a seminar in the CEC programme, Brussels 1985. Dordrecht: Martinus Nijhoff, 1986. (Current topics in vet erinary medicine and animal science 3 7) Sosialdepartementet. Tsjernobyl-ulykken: Rappmi fr a Helsedirektoratets radgivende faggruppe. Oslo: Universitetsforlaget, 1987 (Norges offentlige utredninger NOU 1987: 1) Thornhill JA. Renal endocrinology. In: Drazner FH, ed. Small animal endocrinology. New York: Churchill Livingstone, 1987: 315-39 The manuscript (diskette and paper copies) should be sent to one of the editors in 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 accep tance and expected time of publication. REPRINTSWILL NOT BE A VAILABLE. In the interest of speed, no proofs will be sent to authors. It is therefore of vital importance that the manuscripts are carefully checked before submission. BULLETIN OF THE SCANDINAVIAN SOCIETY FOR PARASITOLGY Editor: Jon1n Tharaldsen, National Veterinary Institute, P.O. Box 8156 Dep, N-0033 Oslo, NORWAY. Telephone: 147 22%4617 Fax: +47 22600981 e-mail [email protected] Editorial board: Denmark: E. Tellervo Valtonen, Sweden: Flemming Frandscn, Royal University of Jyviiskylii, J ohan Hoglund Vet. and Agric. Univ., Sect. Dept. ofBiology, P.O. Box National V et. Inst./ Swedish for Zool., Inst. for Ecol. and 35, FIN-40351 Jyviiskylii Univ. Agric. Scient., Dept. Molec. Biology, BU!owsvej (Tcl: +358 14 602329, Fax: ofParasitol., P.O. Box 13, DK-1870 Fredrikslwrg ( ' +358 14 602321) 7073, S-750 07 Uppsala, (Tel: +45 3528277), Fax: 1 tJ ') e-mail: etvalto(cDt11kki.jyu.fi (Tel: +46 18674156, Fax: 35282774) +46 18309162) e-mail: e-mail: ccol(a)kvl.dk Iceland: J [email protected] Siguri)urRi chter, Maria Vang .Johanst.�n, Danish l Jnivcrsity of Iceland, Inst. Lars-Ake Nilsson, Bilharziasis I �ab., .la:gcrsl)()rg for Exp. Pathol. Keldur, University of Goteborg, Alle 1 D, P.O. Box l\5110, lS-112 Inst. of Med. Microbial. & DK-2920 Charlottenlund Reykjavik (Tcl: +354 Immunol., Guldhedsgatan (Tel: +45 3%26!(lX, h1 x: I tJ (i 5674700, Fax: 1354 5673- 10, S-413 46 Goteborg 39626121) c·mail: <)'J!)) (Tel: 46 31 6047 17, Fax MVJ@Bilharziasis. l )K c·lllail: shr(olrhi.hi.is +46 31604688) Eskild Petersen, Statcns Karl Skimisson, University Jan Thulin Semminstitut, I .ab. of of Iceland, Inst. fo r Exp. National Board of Parasitology, DK ·2300 Pathol., Kcldur, IS-112 Fisheries, Inst of Marine Copenhagen S (Tcl: 14'\ Reykjavik Research, P.O. Box 4, S- 32683223, Fax: 1 45 (Tcl i 354 5674700, Fax: 453 21 Lysekil (Tel: +46 32683033) e-mail: cp(alssi.dk i 3S4 5673979) 52314180, Fax: +46 e·mai I: [email protected] 523 13977) e-mail: Finland: [email protected] Margaretha Gustafsson Norway: Abo Akademi, Dept. of Bioi., Tor A Bakkc, Zoological Editor of Baltic News: BIOCITY, Artillerigt. 6, FIN· Museum, University of Christian M. Kapel, 20520 Abo (Tel : + 35X Oslo, Sarsgt. 1, N-0562 Royal V et. and Agric·. 212654603, Fax: +35X Oslo (Tel: +47 22851 678, Univ., Danish Center. for 212654748) e-mail: Fax: +47 2285183 7) e-mai1: Exp. Parasitol. BUlowsvej [email protected] [email protected] 13, DK-1870 Fredriksberg C (Tel: +45 35282778, Fax: Hannu Kyronseppii,Auroran Svein G. Gundersen, +45 35282774), Sairaala, Nordenskioldsgt. 20, Ullevaal Hospital, Dept. of. e-mail: [email protected] FIN-00250 Helsinki Inf. Diseases, N-0407 Oslo (Tel: +358 9 471 5983, Fax: (Tel: +47 22119119, +358 9 471 5900) Fax: +47 22119125) BULLETIN OF THE SCANDINAVIAN SOCIETY FOR PARASITOLOGY VOL. lO No. 2 CONTENTS December 2000 Proceedings of the symposium on Ecological Parasitology on the turn of Millenium, St. Petersburg, Russia, 1-7 July, 2000 Kirill V. Galaktionov, Oleg Pugachev, Ken Ma cKenzie and Karl Skirnisson (E ditors) ...... 1 SOCIETY NEWS Minutes fr om Board meeting 28 Nov. 2000 ...... 138 New book: Methods in Malaria Research ...... 141 Parasitological activities in Sweden ...... 141 Parasitological activities in Norway ...... 142 Guidelines for contributors ...... 143