On the Coexistence of Fleas (Siphonaptera) on Mammals in Hungary

Átírás

1 5 On the Coexistence of Fleas (Siphonaptera) on Mammals in Hungary István SZABÓ Zoological Department of the Hungarian Natural History Museum, Budapest Several papers dealing with the natural history of fleas record the common occurrence of several flea species on one host-species. These communications usually refer to all the flea species occurring on several individuals of the host species and only rarely does a paper consider the flea population found on a single host specimen (or in their nests, burrows, etc.). In my opinion i t would be more informative i f the flea populations were evaluated on the basis of single individuals of the same host species, because the constant haunts of animals the location of their nests, the conditions of their é.ssociutions with other neighbouring species, and so on, differ even within a small area of the habitat, and all these factors influence the common occurrence of flea species.. The common occurrence of flea species aroused my interest some time ago. Examining.my material collected from 958 t i l l the end of 967, I found that although given host, usually carries a single flea species the common occurrence of several flea species is not rare.on one host species (or in its nest, material), I found one flea species in 66.5 per cent of cases, two In 26.8 fo, three in 5.8 and 4 in 0.9 Since in one-third of the cases more than one flea species was found to occur on a host specimen, a study of the conditions of coexistence neemed

2 worthwhile, with special regard to the following pointb of inquiry: is there any systematic relationship between the species occurring together; are they in any way interdependent on one another; do certain species favour coexistence; are there species which seldom if ever occur coexist with others; finally, is there any apparent reason, ecological or otherwise, for the common occurrence of species or is it merely a spontaneous or chance phenomenon? Materials and Methods Since methods for collection of fleas or their hosts are well known only a summary of methods used in this study will be given.mouse and shrew species were collected by live and lethal traps, dormice and hedgehogs by hand, moles and mole-rats by subterranean lethal traps, gophers either by hand (flooding with water) or by shooting, bats by nets or by hand (single specimens), all other mammals (squirrels, foxes, badgers, wild cats, martens) by shooting. Immediately after capture, the host specimens were placed in a sealed cotton sack and later in chloroform vessels; thus all fleas present on the host at the moment of capture were securely collected after narcotization. Great weight was laid on gathering the fleas separately by host specimen, except for cases when individuals of the same host species were caught in traps on further than a few steps from one another, since in all probability their fleas occurred on members of the same family living in a common subterranean burrow. The fundamental requirement for establishing conditions of coexistence of flea species per individual host is a recording system ( P i g. l) which allows one to discover, at any time, the numbers and species of fleas originating from a given host specimen. The serial inventory number must also be entered for the host in question ( P i g. l) since without this, had the flea specimens been preserved either in alcohol or mounted on a

3 Serial Number sorszám Host Species gazdaállatfaj h* cm O P H fcj*co p m H> 0» P o c_i. p. CT CD ro ind.no. - db. Date of Collection gyűjtés ideje Locality lelőhely Collector gyűjtő 00 H Remarks megjegyzés

4 Table l - l. táblázat. Host Species gazdafaj Research Material - Vizsgálati anyag Material Investigated vizsgált anyag Nests, Burrows, and Holes Investigated vizsgált fészkek, kotorékok és odúk Total összesen Positive pozitiv S-S II II Total összesen Positive pozitiv ! 6 7 Apodemus agrárius ,5 Apodemus flavicollis ,7 6 5 Apodemus sylvaticus ,7 Arvicola terrest.sch Barbastella barbast ,5 Canis familiáris ,- Citellus citellus ,5 2 2 Clethrionomys glareol ,2 2 2 Cricetus cricetus Crocidura leucodon Crocidura suaveolens - 0

5 Dryomis nitedula - 00,- 3 Eptesicus serotinus ,- - Erinaceus europaeus r ,8 Felis catus ,- Glis glis ,- Felis sylvestris - 00,- Lepus europaeus ,3 Martes foina ,- - Mêles mêles - 00,- Micromys minut.prat ,7 - Microtus agrestis ,- Microtus arvalis ,7 0 7 Microtus oeconom.méh ,3 Minlopterus schreibersi ,3 Mus musculus spicilegus ,5 2 2 Muscardinus avellanar ,6 5 Mustela nivalis 2 50,- Mustela putorius furo ,- Mustela putorius hung. 00,- Myotis brandti 00,- - Myotis dasycneme - 00,- Myotis daubentoni Myotis bechsteini Myotis myotis ,6

6 Myotis nattereri ,5 Myotis oxygnathus ,8 Neomys anomalus milleri , Neomys fodiena ,4 Nyctalus leisleri Nyctalus noctula ,- Ondatra zibethicus - 00,- Pipistrellus pipistr ,8 Pitymya subterraneus ,5 Plecotus auritus Plecotus austriacus ,5 Rattus norvegicus Rhinolophus ferrumequ ,5 Rhinolophus hipposid Sciurus vulgaris fuse ,8 2 2 Sorex araneus ,3 Sorex minutus 3 2 7,7 Spalax leucodon ,9 Talpa europaea , Vulpes vulpes crucigera ,6 2,33 826, ,43 64,57 fo 64,28 %

7 slide (F i g. 2), i t would "be impossible to establish later whether the fleas in the collection come from the same host individual or not. During the ten year period I had occasion to examine 233 specimens and 42 nests or subterranean burrows of 55 mammalian species. Fleas were found on per cent of the hosts and in per cent of their nests ( T a b l e l ). The Table shows the extent of infection of the several host species.one hundred per cent,or complete lack of infection by fleas occurred mainly in those host species for which I was unable to collect sufficient material. Presumably, a large number of specimens would not show this high degree or complete lack of infection. On mouse and shrew species and on moles and mole-rats there occur probably more fleas than i t was able to establish. This stems from the fact that most of these species were collected by lethal traps and i t is possible that, despite several examinations of the traps per day, a part of the flea population had already left the cooling hosts prior to collection. However, I estimate this source of error to be not higher than 5-0 j6. After capture of individuals of other mammalian species, I was able to prevent scattering of the fleas. Similarly,the material of nests, burrows, holes were in a l l cases placed immediately into completely sealable bags and then transferred into an extractor ( F i g. 3) from which the parasites are unable to escape. Results Occurrences of single flea species Before describing the coexistence of different flea species on a single host animal I propose to l i s t the cases of single occurrences of flea species as repeated reference will later be made to these. The number of cases is in some instances higher than that of the listed localities, because collections were made at several separate times.

8 Fig ábra. Extractor Féazekfuttató

9 Table 2-2. táblázat. Occurrence of Single Flea Species Bolhafajok egyenkénti előfordulása Serial Number sorszám Siphonaptera -p CO CQ CO Species - faj i m cd o Host - gazdaállat Species - faj -p CO 03 CO i CO m cd o Locality lelőhely Archaeopsylla 5 Erinaceus eur.roum. 5 Misina-tető Monor Orgovány Tompa 2 Cerat." tribulis Citellus citellus Tahitótfalu 3 Chaetops. globiceps 4 Felis sylvestris Vulpes vulpes 3 Diósjenő Kaposvár Szentmártonkáta Pilisszentkereszt 4 Chaetops. rothsch. 2 Martes foina 2 Deszkáspuszta Mátraszentimre 5 Citelloph. martinoi 6 Citelloph. simplex Citellus citellus Citellus citellus Gyulafirátót Ha j dubago s 7 Ctenoceph. canis 2 Canis familiáris lepus europaeus Budapest Boly 8 Ctenoceph. fells 4 Canis familiáris Felis catus Mustela furo put or. 2 Budapest Monor Budapest Gödöllő

10 Ctenoph.ag. 49 Apodemus 7 Kisbalaton "bosnic. agrárius Fityeház Zajk Apodemus 8 Németbánya flavicoll. Kisbalaton Szabadegyháza Bakonybel Kiszépalmapuszta őriszentpéter Pálihálás Apodemus 2 Pityeház sylvaticus Buesuta Clethrionomys Németbánya glar. Kisbalaton Kaposmérő Szalafő Bucsuta Micromys 2 Tatabánya minut.prat. Németbánya Microtus 2 Kisbalaton agrestis Microtus 2 Németbánya arvalis Microtus Kisbalaton oecon.méh. Pitymys 4 Németbánya subterran. Iharkút 0 Ctenoph.ag. 7 Apodemus 6 Sarkadremete eurous flavicoll. Mus musculus Sarkadremete spic. T I I Ctenoph.ag. 4 Apodemus Tákos kleins. agrárius Apodemus 2 Táko s sylvaticus Microtus Lónya arvalis 2 Ctenoph.ag. 7 Apodemus 3 Kisnána peusian. flavicoll. Clethrionomys 4 Deszkáspuszta glar. Ki snána 3 Ctenoph. 6 Apodemus 2 Hajdubagos assimilia flavicoll. Tákos Clethrionomys 2 Tahitótfalu glar. Deszkáspuszta

11 2 5 4 Microtus arvalis Neomys fodiens Sorex araneus Talpa europaea 8 2 Orgovány Kishalaton Bugyi Németbánya Lippó Békéscsaba Nagykovácsi Kisbalaton Kisbalaton Sarkadremete Kisszépalmapuszta 4 Ctenoph. caucasicus Spalax leucodon Hajdubagos 5 Ctenoph. congener 3 Apodemus flavicoll. Clethrionomys glar. 2 Németbánya Németbánya Bakonybél 6 Ctenoph. orientális. 6 Apodemus flavicoll. Citellus citellus Mustela nivalis 4 Ki snána Hollóháza Szabadszállás Fácánkert Bugac Ócsa 7 Ctenoph. solutus 4 Apodemus flavicoll. Apodemus sylvaticus Clethrionomys glar. 2 Gombáspuszta Nagysomlyóhegy Deszkáspuszta Alcsut Pisznice-hegy Gézaháza Kisszépalmapuszta Kisnána Budakeszi Litke Kistolmács Deszkáspuszta 8 Doratopsylla d.das. 0 Neomys fodiens Sorex araneus 9 Németbánya Németbánya Deszkáspuszta Szalafő

12 Hystrichops. talp.or. 5 Apodemus agrárius Kisbalaton Apodemus flavicoll. Microtus Cserfekvés Kisbalaton oecon.jméh. Ondatra Kapuvár zihethicus Talpa Iharkút europaea 20 Ischnopsyllus 6 Myotis myotis Pilismarót elong. Nyctalus 5 Rétszilas noctula Szikra-Tőserdő Lillafüred Kisbalaton 2 Ischnopsyllus 5 Myotis myotis 2 Legény-barlang hexact. Myotis 6 Ördöglyuk-barlang oxygnathus Pálvölgyi-barlang S olymári-barlang Pipistrellus pipistr. Plecotus austriacus Abalige Legény-barlang Szekszárd 6 Tata Gyöngyös Budapest Pilis Dorog Tompa ti-barlang 22 Ischnopsyllus interm. 7 Eptesicus serotinus Miniopterus schreib. Myotis dasycneme Myotis myotis Myotis oxygnathus Pilismarót Budapest Alpár Németbánya Abaligeti-barlang Szikra-Tőserdő Pilismarót Eger Pisznicei-barlang Ördöglyuk-barlang Pálvölgyi-barlang Nagymaros Abaligeti-barlang Leány-barlang Gyöngyös

13 Ischnopsyllus octact. 24 Ischnopsyllus simpl.myst. 25 Ischnopsyllus simpl.simpl. 26 Leptopsylla segnis 6 Pipistrellus pipistr. Myotis brandti 2 Myotis nattereri 7 Apodemus flavicoll. Mus muscuius spicil. 6 Sopron Lillafüred Németbánya Bakonybél Baláta-tó 2 Leány-barlang Legény-barlang 2 5 Tákos Deszkáspuszta Orgovány Pel3Őgöd Sarkadremete Ki sszépalmapuszta Budapest Tákos Pálihálás 27 Megabothris turbid. 3 Apodemus flavicoll. Clethrionomys glar. Microtus arvalis őriszentpéter őriszentpéter Tákos 28 Megabothris walkeri 2 Microtus agrestis Sorex minutus Kisbalaton Kisbalaton 29 Monopsyllus sciur. 26 Apodemus flavicoll. Dryamis nitedula Glis glis Mêles mêles Muscardlnus avellan. Myotis oxygnathus Sciurus vulg. fuscoat Gödöllő Iharkút Gödöllő Deszkáspuszta Pisznice-hegy Budakeszi Budapest Mecsek-hg Gödöllő Legény-barlang Diósjenő Pilisszántó Deszkáspuszta Budapest

14 2 3 4 Sciurus vulg. fuscoat. 4 Csákvár Budakeszi Zirc Németbánya Gödöllő Mecsek-hg. 30 No so pay Uu s fasciat. 9 Apodemus flavicoll. Apodemus sylvaticus Microtus arvalis Mua muaculua spicil Pisznice-hegy Sarkadremete Tahitótfalu Sarkadremete Békéscsaba Sarkadremete Békéscsaba 3 Nycteridops. eusarca 3 Nyctalus noctura 3 lengyel Budapest 32 Nycteridops. pentact. 7 Barbastella "barbast. Myotis myotis Myotis oxygnathus Plecotus austriacus 4 Vereshegyi-barlang Abaligeti-barlang ördöglyuk-barlang Üllő Gödöllő Királyrét 33 Palaeops. kohauti 34 Palaeops. similis 35 Palaeops.. sor.rosic. 36 Peromyscops. fallax 3 Talpa europaea 0 Talpa europaea 8 Apodemus agrárius Clethrionomys glar. Micromys minut.prat. Neomys anomal.mill. Sorex araneus 3 Clethrionomys glar. 3 Kisbalaton Iharkút 0 Németbánya Szabadegyháza Deszkáspuszta Bakonybél Iharkút 4 Kisbalaton Kisbalaton Pákozd Németbánya Kisbalaton Pákozd Kisbalaton Pákozd Németbánya

15 2 3 4 Microtus Németbánya arvalis Pitymys Németbánya subterran. 37 Pul ex 5 Canis 5 Orgovány irritans familiáris Hont Sárcsikut Kapuvár Bugac Pelis catus Csákvár Meie3 meles 3 Drégelypalánk Lónya Mustela Hajdúbagos putor.hung. Vulpes vulpes 5 Szigetmonostor crucig. Szentendre Haj dubagos Kapuvár Mihálygerge 38 Rhinolophops., 2 Rhinolophus 2 Gyula uni p. ferrume. Abaligeti-barlang In T a b l e 2 certain species occur in only a few cases, but this is not an expression of their rarity since most of them occur - as shown in T a b l e 3 - more frequently in the company of other species.on the basis of material collected so far, the following flea species of mammals may be considered rare : Chaetopsylla rothschildi Kohaut; Ischnopsyllus simplex simplex Rothschild; I. simplex mysticus Jordan; I. variabilis (Wagner); Megabothryis walkeri (Rothschild); Palaeopsylla kohauti Dampf; Peromyscopsylla fallax (Rothschild); Rhadinopsylla isacantha isacantha (Rothschild); R. pentacantha (Rothschild); R. strouhali Smit, hitherto found as a single specimen only and identified conditionally even by its discoverer; Rhinolophopsylla unipectinata unipectinata (Taschenberg), and Tarsopsylla octodecimdentata octodeclmdentata (Kolenati). As the host species of these fleas are rare in Hungary, their infrequent capture is quite understandable.

16 Table 3-3. táblázat. Common Occurrence of Plea Species - Bolhafajok együttes előfordulása Serial Number sorszám Common Occurrence of Flea Species Együttesen előforduló bolhafajok Case eset Host - gazdaállat Species - faj Case eset Locality lelőhely Archaeopsylla erin. Pulex irritans 2 Ceratoph.pullatus Ctenoph.ag eurous Nosopsyllus fasc. 3 Ceratoph.pullatus Monopsyllus sciur. 4 Ceratoph.tribulis Monopsyllus sciur. 5 Ceratoph.tribulis Nosopsyllus fasc. 6 Chaetops.globicepB Chaetops.trichosa Paracéras melis Pulex irritans Vulpes vulpes crue. ócsa Apodemus flavicoll. Sarkadremete Muscardinus avell. Gödöllő Sciurus vulg.fuse. Szeleste Citellus citellus Tahitótfalu Vulpes vulpes crue. Deszkáspuszta

17 Chaetops.globiceps Pulex Irritans 8 Chaetops.globiceps Ctenoceph.canis Paraceras melis Pulex irritans 9 Chaetops,globiceps Monopsyllus sciur. 0 Chaetops.globiceps Monopsyllus sciur. Pulex irritans Chaetops.globiceps Paraceras melis 2 Chaetops.globiceps Paraceras melis Pulex irritans 3 Chaetops.trichosa Paraceras melis Pulex irritans 6 Vulpes vulpes crue.. 6 Vokány Pécsudvar Inárcs Rákoscsaba Budakalász Drégelypalánk Vulpes vulpes crue. Mecsek-hg. Sciurus vulg.fuse. Németbánya Vulpes vulpes crue. Németbánya Vulpes vulpes crue. Deszkás-puszta Meies meles Ba jna 2 Meies meles o Sárcsikut Litke 4 Chaetops.trichosa Pulex irritans 7 Canis familiáris Meies meles Drégelypalánk Pusztavám S za ba d egy há za Egervár

18 M CItelloph.martinoi Ctenoph.orientális 6 Citelloph.simplex Ctenophth.assimllis Citelloph.simplex Ctenoph.caucasicus Ctenoph.orientális 8 Citelloph.simplex Ctenoph.orientális 9 Ctenoceph.canis Ctenoceph.felis Pulex irritans Vulpes vulpes crue. 3 Drégelypalánk Belvárgyula Kutberek 3 Citellus citellus 3 Hollóháza Citellus citellus Hajdubagos Spalax leucodon Ha jdubagos 3 Citellus citellus 3 Ha jdubago - s Bátorliget Canis familiáris. Biharugra 20 Ctenoph.ag.hosnic. Ctenoph.assimilis 4 Clethrionomys glar. Microtus oecon.méh. 3 Balatonlelle Kisbalaton Kisbalaton 2 Ctenoph.ag.bosnic. Ctenoph.assimilis Megabothris walker! 22 Ctenoph.ag.hosnic. Ctenoph.assimilis Palaeops.sor.ros.» Clethrionomys glar. Kisbalaton Sorex araneus Kisbalaton

19 Ctenoph.ag.bosnic. Ctenoph.congener Apodemus flavicoll. Clethrionomys glar. Neomys fodiens 9 Zajk Sárcsikut Németbánya Kisszépalmapuszta Németbánya 24 Ctenoph.ag.bosnic. Ctenoph.congener Ctenoph.solutus Clethrionomys.glar. Sárcsikut 25 Ctenoph.ag.bosnic. Ctenoph.congener Doratops.d.dasycn. 2 Apodemus flavicoll. Clethrionomys glar. Németbánya Németbánya 26 Ctenoph.ag.bosnic. Ctenoph.congener Hystrichops.talp.or. 27 Ctenoph.ag.hosnic. Ctenoph.congener Hy strichops.talp.or. Peromyscops.fallax Clethrionomys glar. Gombáspuszta Clethrionomys glar. ] Iharkút 28 Ctenoph.ag.hosnic. Ctenoph.congener Peromyscops.fallax 3 Clethrionomys glar. Pitymys subterraneus 2 Néme tbánya Németbány 29 Ctenoph.ag.hosnic. Ctenoph.congener Rh.vàtnops.isacantha 30 Ctenoph,ag.boonlc Ctenoph.orientális Clethrionomys glar. Iharkút Apodemus flavicoll. Mecsek-hg.

20 Ctenoph.ag.bosnic Ctenoph.solutus 7 Apodemus flavicoll. Clethrionomys glar. 6 Sárcsikut Németbánya Pisznice-hegy Bakonynána Budakeszi Kaposmérő 32 Ctenoph.ag.bosnic. Doratops.a.dasycn. 2 Neomys anomal.mill. Pitymys subterraneus Németbánya Németbánya 33 Ctenoph.ag.hosnic Hystrichops.talp.or. 2 Microtus arvalis Mus musculus spic. Németbánya Kisbalaton 34 Ctenoph.ag.bosnic Megabothris turbid. 9 Apodemus flavicoll. Clethrionomys glar. Microtus agrestis 7 őriszentpéter őriszentpéter Zajk Szalafő Kisbalaton 35 Ctenoph.ag.bosnic. Megabothris turbid. Monopsyllus sciur. 36 Ctenoph.ag.hosnic. Megabothris walkeri 37 Ctenoph.ag.hosnic. Megabothris walkeri Palaeops.sor.ros. 38 Ctenoph.ag.hosnic Monopsyllus sciur. 39 Ctenoph.ag.hosnic. Nosopsyllus fasc. Palaeops.sor.ros. Peromyscops.fallax Clethrionomys glar. őriszentpéter Microtus oecon.méh. Kisbalaton Neomys fodiens Kisbalaton Apodemus flavicoll. őriszentpéter Apodemus flavicoll. Németbánya

21 Ctenoph.ag. "bosnic. Palaeops.sor.ros. 3 Microtus agrestis Neomys anomal.mill. Talpa europaea Kisbalaton Németbánya Kaposmérő 4 Ctenoph.ag.eurous Hystrichops.talp.or. 42 Ctenoph.ag.eurous Hystrichops.talp.or. Nosopsyllus fasc. 43 Ctenoph.ag.eurous Nosopsyllus fasc. 44 Ctenoph.ag.eurous Rhadinops.pentac. 45 Ctenoph.ag.kleins. Ctenoph.assimilis Megabothris turbid. Apodemus flavicoll. Sarkadremete Apodemus sylvaticus Sarkadremete Apodemus flavicoll. Sarkadremete Apodemus flavicoll. Sarkadremete Apodemus sylvaticus Tákos 46 Ctenoph.ag.peusian. Ctenoph.congener * 2 Apodemus flavicoll. Clethrionomys glar. Kisnána Kisnána 47 Ctenoph.ag.peusian. Ctenoph.oriental!s 48 Ctenoph.ag.peusian Ctenoph.solutus 49 C te n o ph. ag. peus ian. Hystrichops.talp.or. 50 Ctenoph.ag.peus. X eurous C tenoph.assimilis Apodemus flavicoll. Deszkáspuszta 5 Apodemus flavicoll. 5 Deszkáspuszta Kisnána. Apodemus flavicoll. Diósjenő Apodemus sylvaticus Ócsa

22 Ctenc-ph.ag.peup. Microtus arvalis Ohat x eurous x kleins. Ctenoph.assimilis 52 Ctenoph.ag.ssp. Ctenoph.congener 53 Ctenoph.ag.ssp. Ctenoph.orientális 54 Ctenoph.. as similis Megabothris turbid. 55 Ctenoph.assimilis Nosopsyllus fasc. 56 Ctenoph.assimilis Palaeops.sor.ros. 57 Ctenoph.bisoctodent. Palaeops.kohauti Palaeops.similis 58 Ctenoph,bisoctodent. i Palaeops.similis 59 Ctenoph.caucasicus Ctehoph.congener Ctenoph,orientális 60 Ctenoph.caucasius Ctenoph.orientális Clethrionomys glar. Deszkáspuszta Talpa europaea Hajdubagos Clethrionomys glar. Kisbalaton 5 Apodemus flavicoll. Bugac Apodemus sylvaticus Tahitótfalu Microtus arvalis 2 Pácánkert Békéscsaba Talpa europaea Apaj Neomys fodiens Kisbalaton Talpa europaea Deszkáspuszta 2 Talpa europaea 2 Deszkáspuszta Spalax leucodon Hajdubagos Spalax: leucodon 4 Ha j dubago s

23 C t enoph. c ongener Dorâtops.d.da syen. Sorex araneus Németbánya 62 Ot eno ph. c ongener Peromyscops.fallax 2 Clethrionomys glar. Microtus arvalis Németbánya Németbánya 63 Ctenoph. solutus Rhadinops.strouhali 64 Dorâtops.d.dasycn. Hystrichops.talp.or. Palaeops.sor.ros. 65 Doratops.d.dasycn. Monopsyllus sciur. Apodemus flavicoll. Deszkáspuszta Neomys.anomal.mill. Németbánya Apodemus flavicoll. Bakonynána 66 Doratops.d.dasycn. Palaeops.similis 2 Sorex araneus Talpa europaea Deszkáspuszta Deszkáspuszta 67 Doratops.d.dasycn. Palaeops.sor.ros. 68 Hystrichops.talp.or. Megabothris walkeri 69 Ischnopsyll.elong. Ischnopsyll.interm. 70 Ischnopsyll.elong. Nycteridops.eusarca 5 Sorex araneus 5 Németbánya Ki s s zépalmapus z ta Iharkút Microtus oecon.méh. Kisbalaton Myotis oxygnathus Héviz Nyctalus noctula Legény-barlang 7 Ischnopsyll.hexact. Ischnopsyll.interm. 6 Epteslcuo serotinus Myotis oxygnathus 5 Velence ördöclyuk-barlang Abaligeti-barlang Aggteleki-barlang

24 Ischnopsyll.hexact Ischnopsyll.interm. Nycteridops.pentact«73 Ischnopsyll.hexact. Ischnopsyll.variah. Myotis oxygnathus Abaligeti-barlang Myotis oxygnathus ördöglyuk-barlang 74 Ischnopsyll.hexact. Nycteridops.pentact. 3 Barhastella harhast. Plecotus austriacus 2 ördöglyuk-barlang Kec ske-barlang Budapest 75 Monopsyllus sciur. Tarsopsylla octod. 2 Sciurus vulg.fuse. 2 Répáshuta Sopron 76 Paraceras melis Pulex irritans 3 Canis familiáris Meies meles 2 Németbánya Gödöllő Budakeszi

25 The case number of the host species ( T a b l e 5) displays without need of explanation the extent of host specificity of the flea species. This relationship is not so evident from a study of the l i s t of species found occurring together. Common occurrences of several flea species The coexistence of flea species on the same host individuals and in the nest materials is listed in T a b l e 3. Most of the common occurrences were found in one or two instances only; those noted in three or more cases are worthy of special attention. Besides the number of cases,the significance of these common occurrences increases i f they are found in more than one host species or in geographically remote localities. At present, the frequency of cases seems to be the most significant ( T a b l e 4). Table 4-4. táblázat. Frequency of Common Occurrences of Flea Species to the Number of Species According Bolhafajok együttes előfordulásának gyakorisága a talált fajok száma szerint Number of Different Common Occurrences Comprising 2 Species J 3 Species 4 Species Különféle együttes előfordulások száma 2 faj 3 faj 4 faj Frequency of Common Occurrences Együttes előfordulások gyakorisága X 7 X 9 X X

26 Further evaluation and illustration are derived from two other Tables. T a b l e 5 shows the frequency of occurrences (combined cases of single and common occurrences) per host of the flea species. T a b l e 6 displays the frequency of common occurrences of the individual flea species. In the course of my investigations the following forms of common occurrences of flea species were established. Flea common occurrences on mouse species It was most frequently on mouse species that coexistence of several flea species was found. This is probably accounted for net only because the great majority of collected host individuals belonged to this group but also because of the well-known fact that most flea species can be found on members of the family Muridae in Hungary. The majority of flea species living on our mice belong to the genus Ctenophthalmus, of which Ct. agyrtes (C.a.bosnicus Wagner; C.a.eurous Jordan et Rothschild; Ca. kleinschmidtianus Peus; Ca. peusianus Rosiczky) occur in most populations and in rather well defined geographical areas, as was demonstrated in an earlier paper (SMIT and SZABÓ, 967). However, other species of the genus also occurred, viz. Cassimilis assimilis (Taschenberg), Ccongener congener Rothschild, and. Csolutus solutus Jordan et Rothschild. Together with those species, the Hungarian members of the genera Hystrichopsylla, Leptopsylla, Monopsyllus, Nosopsyllus, Palaeopsylla, Peromyscopsylla, and Rhadinopsylla occurred more rarely. The two most frequent populations (on 9 and occasions) also occurred on mice (serial numbers 23 and 34). Both species are rather local, being found in the Bakony Mts* and in the neighbourhood of the village őriszentpéter. Another interesting fact is that one instance of both common occurrences was observed in the environs of the village Zajk, Comitat Zala. «Mountains

27 Table 5-5. táblázat. Frequency of Flea Species Found on Host Species - Gazdaállatfajokon talált bolhafajok előfordulási esetszámai H o H.. *J. ra p U CQ -p. ft ra H -p ri ci -p H ri. CQ H f-+ ri. ri ri CQ ra Sa o CQ CQ ri CQ o H CQ rp ri. ci ri fl ri H o H ft-h CÖ o M o ra ra cd CQ CD ri H ri. ri ri H - CD ri cd p ri Si ft U CD fl CQ ri ri ri o ri ft P ri CD CD ra ri i fl ri ri H ra ri o fl 03 H H O ü O H H ra CQ ri o H CD H O H (D cd ra ri rh cd CD a P fl 8)H H H CQ o O P H O CQ H ri ri ri ri O CQ CQ H CQ fl P ft H H ra ri ri M ft-h P m ri ri o cd tudp ri ri 03 pi ci O a ri cd ri ri cd CQ o P o p P cö ri H fl fl o ri çrj o ri CO a O a CD ri CD ri cd ri ft ft CQ H ri ri H <H ri ri ri ri Ü H P H o -P H ci H CÖ CD fl CD fl ft ft ft CQ -H o o CD ri CQ. o X p P S a CQ ri ri fl 3 ri (U cd ra CD p CQ O H cö 3 H O ri ra o H.... CQ m ri ri H H cd ra ri H H cd cd CQ CQ CQ CQ ri o H O 0 CQ P H ft ft W) U p. Í5JJ bo bo bû tid CQ H ci ri O r i P- 03 fi H o ra ra ri ri ft^ CQ ra ft-h O H CQ r-j H fl fl cd CÖ ri cd cd cd fl o o o ra o.. «H ri ri ra ri ra ra fl ft ri H u CQ CQ ra ft ft ft ft *.... ra.ri CQ CQ ra m ra ra >>fl fl H ra ra ra ri o ri ft ft ft ft O ft-p ft ft o ft o CD CD -ri fl fl fl fl fl fl A A ft ft o ft ft ft ft ft ft >j > > H H ft ft ft ri CQ ih o o O o <D ra O O o H rh O O ft ft ft ft ft fti ft ft ft ft o H O O o O o o o ft o o ra ra ri ri o o o CD b ^ ri ri ri ft CÖ -P P -P -P -P rh H o o o o o o o o o o o o o O o p ri ri ri ri ri ri o fl fl ft ft s i X H H H o o fl CÖ CÖ CD CD CD CD CD ri ri ri ri ri ri ri ri ri ri ri ri ri ri ri cd P fl fl A A fi fl -Pcd cd o o P P ri ri ri ri O03 rh ri ü u U m CÖ CÖ cd P P CD CD CD CD CD CD CD CD CD CD CD CD CD CD ri ra o o o o o o o ft tto QC ri ra rh H H ri ri H ri ci H ri U CD CD S fii fi -P P o >Î CQ CQ m ra ra ra ra CD o o l>i ï>> a ri ri ri 03 pi fl fl fl fl ri 4 o O o o o O O O O O o O O O o O o O o Ü O O H H M M H H M H: S S S ri; K PH PH PH PH PH ft cri fl fl fl EH Apodemus agrárius 7 4 \ Apodemus flavicollis 30 C H Apodemus sylvaticus Barbastella barbast. 2 3 Canis familiáris Citellus citellus Clethrionomys glar i Dryomis nitedula Eptesicus serotinus < 5 Erinaceus europaeus 5 Felis catus 2 4 Felis silvestris 4 Glis glis 5 Lepus europaeus Martes foina 2 Meles meles Micromys minut.prat. 2 Microtus agrestis 4 Microtus arvalis Microtus oecon.méhelyi Miniopterus schreib. j { \ 3 Muscardinus avellan. \ 2 Mustela nivalis Mus musculus spicil. Mustela putor.furo Mustela putor.hung. Myotis brandti i Myotis dasycneme \ Myotis myotis Myotis nattereri 2 Myotis oxygnathus Neomys anomalus mill Neomys fodiens 2 2 Myctalus noctula 6 4 Ondatra zibethicus Pipistrellus pipistr. Pitymys subterraneus 6 < 4 2 Plecotus austriacus 7 5 Rhinolophus ferrume. 2 Sciurus vulgaris fuse. \ 4 3 Sorex Sorex araneus minutus 6 \ 47 Spalax leucodon 7 6 Talpa europaea i i Vulpes vulpes crue. \

28 Table 6-6. táblázat. Frequency of Common Occurrences of Flea Species - Bolhafajok együttes előfordulásának gyakorisága 0 g CQ p CQ CÛ. ri H p P ri H PI CQ H ri. ri X P! CQ CQ ci o Cû ri H P ri ri ri X ri H H PH P O Ü CQ CQ ri. CQ 0 ri H ä ri rp H H 0 ri H 0 cd p u ri CÖ CD CQ ri 0 H ri ri H 0 H d o ri H 0 ftp 0 0 H fl ri o ci p H CQ CQ H ri o fl O Ü O H CQ CQ 0 H ri o fl H. CQ H o H 0 ri ri a fl ri o o P H O CÛ H cd CÖ ri CO CQ H fl -P ri< H H CQ ri 0 ri X X fl H p CÛ ri -p ri o ri ÏÏO o ri P ri cd o cd ri ri ri H ri H ri CQ o p o Ü p H fl O ri 0 ri ri 0 CQ a o ri ri ri  0 H p CQ fl ri P ri od H fl ri ri ri ri -p H O H cd H a 0 fl 0 fl fl PH CQ H CD o o ri o X p CQ ri ri fl a ri ri H cd p rh ri a.. CQ o P,.. CQ CQ ri ri H H ri CQ rh 0 ri cd CQ CQ CQ CQ ri 0 o H o a CQ p H P CQ ri \>> fl tu). P.. bo EVD bq CQ H cd o ri -d fl fl > H H ri ri CQ CQ fl fl fl H CO H H. A fl A fl CÖ ci ri ri cd ri ri fl o o o CQ «. O» ri ri. CÛ O ri H fl P ri CQ CQ cd Pi fl PH PH.. CQ fl cû Cû CQ CQ ri fl H H fl TT) CQ CQ CQ cö o ri fl PH P o P H P fl ft o O 0 0 fl fl fl fl fl fl fl fl fl fl fl fl fl fl Ü fl PH PH fl p P H H fl fl fl ri CÛ H O O O Cû CD O O H O O PH fl fl fl fl PH PH fl fl fl fl fl fl O H O O O O o o CQ CQ ri ri O O O 0 b ri ri ri fl CÖ P p p P H H O O O o O o o O O o O o o o O P ri ri ri ri ri fl fl P PH O a X H H H o fl CÖ p ri ri ri ri ri ri ri ri ri ri ri ri ri ri ri ri p fl fl fl fl ri ri O O P P 0 ri ri cd ri c fl TJ fl CÛ o ri u cd ri P P a> o 0 ri CQ o o o o ri CQ o o fl H H ri ri H ri CÖ ÍH 0 CD fl fl ri H p -P p p p p p p p p P p p p p o!>> CQ CD CQ CQ 0 0 o O >> >> cd cd ri ri ri fl fl fl < CÖ o o o O O o o o o o o o o o o O o o o o fl fl M H H H S S S fl fl fl fl fl fl W fl fl fl fl fl Archaeopsylla erin. ( C erat.pullatus C erat.tribuli s < Chaetops.globiceps \ i Chaetops trichosa 3 <0 Citelloph.martinoi 3 Citelloph.simplex \ i «Ctenoceph.canis i 2 Ctenoceph.felis Ctenoph.ag.bosnicus V i 6 5 Ctenoph.ag.eurous 2 3 i Ctenoph.ag.kleins. \ Ctenoph.ag.peusian. 2 4 Ctenoph.ag.p.x e. Ctenoph.ag.p.x k.x e. Ctenoph.ag. ssp.' i Ctenoph.assimilis 5 \ i 2 Ctenoph.bisoctodent Ctenoph.caucasicus \ 6 Ctenoph.congener 20 2 \ i i Ctenoph.orientális 3 4 \ \ 6 \ Ctenoph.solutus 8 4 \ Doratops.dasycnema 4 3 \ 2 6 Hy strichops.talp.or. 4 2 l i i < Ischnops.elongatus \ Ischnops.hexactenus 7 \ 3 Ischnops.intermed. 7 Ischnops.variabilis Megabothris turbid. <Û i 2 Megabothris walkeri 3 \ Monopsyllus sciur. \ \ 2 2 i i h 3 Nosopsyllus fasciat. i \ 3 5 \ i Nycterid.eusarca Nycterid.pentactena 3 \ Palaeops.kohauti * \ Palaeops.similis 3 l \ Palaeops.sor.ros h \ Paraceras melis Peromyscops.fallax 5 6 i \ i Pulex irritans \ Rhadinops.isacantha Rhadinops.pentacant. \ Rhadinops.strouhali Tarsopsylla octod. 3

29 Plea common occurrences on shrew species The most frequent flea occurring on shrew species in Hungary is Palaeopsylla soricis rosiczkyi Smit, followed hy Doratopsylla das.dasyenema (Rothschild). Either one of those species,or both appear in most populations, frequently associated with Ctenophthalmus agyrtes bosnicus Wagner, or more rarely with Ct.assimil i s assimilis (Taschenberg), Ct. congener congener Rothschild, Hystrichopsylla talpae orientális Smit, Megabothris walkeri (Rothschild), and Palaeopsylla similis similis Dampf. Plea common occurrences on the fox and the badger A specifical fox flea is Chaetopsylla globiceps (Taschenberg), fleas specific for the badger are Chaetopsylla trichosa trichosa Kohaut and Paraceras melis melis (Walker). Besides those three species, Pulex irritans Linné is also frequent on both host species. Since the fox and the badger frequently alternate their lairs their fleas can intermingle easily, although the specific fleas mentioned above usually predominate on their own host species. In the eight cases of common occurrence found on the badger, only the four flea species listed above occurred in various combinations whereas in the fourteen populations of fleas found on the fox, the species Monopsyllus sciurorua (Schrank) and Archaeopsylla erinacei erinacei (Bouché) also appeared in one case each. I should like to remark that the flea species of the fox and the badger are also frequently found on breeds of dog used in hunting badger-dog, fox-terrier, and their mongrels. Plea common occurrences on the mole-rat and the gopher Owing to the much decreased numbers of Spalax leucodon Kordmann in Hungary, the above two host species occur together in only a few localities. However, I think that the common occurrences of

30 their fleaa should he discussed together, since in an investigation of a locality (Hajdubagos), where the two host species occur together the fleas of the gopher can frequently he found on the mole-rat also, either alone or together with other flea species. Ctenophthalmus caucasicus (Taschenberg) is clearly host specific to the mole-rat; i t occurred in a l l observed cases together with the flea species of the gopher, and in one case with Ctenophthalmus congener congener Rothschild. The fleas of the gopher are Citellophilus martioni (Wagner et Ioff), C. simplex (Wagner), and Ctenophthalmus orientális (Wagner). In the eight cases of common occurrences found on the gopher one or other of these species have, with one exception, always been present.besides the foregoing species 3 other fleas were found, each on one occasion. They were Ceratophyllus tribulis (Jordan) presumably transferred to the gopher from a bird nesting on the ground. Ctenophthalmus assimilis assimilis (Taschenberg), and Nosopsyllus fasciatus (Bosc).Two varieties of common occurrence (serial numbers 7 and 8) were found in three cases each. I t deserves special mention that whereas the fleas of the gopher very frequently occur on the mole-rat, the characteristic flea of the mole-rat has never been observed on a gopher occupying the same locality and habitat. Plea common occurrences on the mole I found six kinds of flea populations on the mole with the host specific Palaeopsylla similis simili3 Dampf appearing in most of them. In three cases Ctenophthalmus bisoctodentatus bisoctodentatus Kolenati was also observed - previously collected only from the mole in Hungary - so that i t can be regarded with some justification aa host specific. Besides the two species mentioned above, Ctenophthalmus agyrte s bosnicus Wagner, Ct.assimilis assimilis (Taschenberg), Ct.orientális (Wagner), Doratopsylla dasyonema dasycnema (Rothschild), Nosopsyllus fasciatus (Bosc), Palaeopsylla kohauti Dampf, and,p.soricis rosiczkyi Smit were also collected in one case each. Unfortunately, I have not

31 yet had occasion to collect the material of a mole burrow, though the rich flea fauna of its nest has attracted the atten tion of several research workers (OUDEMANS, 93; WAGNER, 936). Recently ROSICKY (937), SMIT (962), and JTJRIK (968) from an investigation of considerable material established the presence of many flea species,both on the animal and in its nest.regretably, none of these authors, with the exception of ROSICKY have published information on the flea populations of the several hosts and their nests but have merely discussed the occurrence of the flea species observed. Plea common occurrences on the bat I t might be assumed that several types of flea population would occur on bats because certain bat species aggregate in masses in both their winter and summer quarters. True,in most colonies only members of a given species are closely associated with each other but there would s t i l l be more possibilities for transmission of divers flea species than in the case of other mammalian groups. However, I found only six kinds of common occurrences of fleas, among which only two occurred in three and six cases, respectively (serial numbers 7 and 74), while the four other compositions were observed in only one case each. The most frequent species of the cases of coexistence is Ischnopsyllus hexactenus (Kolenati), which., occurred in four variations in eleven cases. The occurrence of I.intermedius (Rothschild) with other species is fairly common having been found in three variations on eight occasions. In addition there are other, somewhat occasional, comron occurrences of smaller specific numbers and rather rare appearence. Thus, besides Monopsyllus sciurorum sciurorum (Schrank), the specific flea of the squirrel (Sciurus vulgaris fuscoater Altum) and its nest, one may occasionally find in its company Tarsopsylla octodecimdentata octodecimdentata (Kolenati), a species rather rare in Hungary. The various dormouse species show a de-

32 finite preference for a r t i f i c i a l nesting boxes,where the squirrel fleas (M.sciurorum) living on the dormice may be found to gether with the fleas of avian species which had previously nested there. This association will not be considered as i t is the result of human interference and not natural. Nor do I wish to discuss the various common occurrences of flea species to be found on our domestic animals. Discussion From the foregoing data of observed coexisting flea species, an attempt will be made to answer the problems raised in the introduction.. The following systematic relationship can be established between flea species occurring together on a given host or in its nests: the members of the populations belong to various families and genera in fo of cases; the same family in 8.4 ^ of cases; the same family and genus in 39.5 i of cases. Taking into consideration that half the species of Hungarian Siphonaptera belongs to the family Hystrichopsyllidae and nearly one-third to the genus Ctenophthalmus - whose members are the most commonly occurring fleas - one can safely conclude that the apparent connections between species occurring together stem more from the composition of the fauna than from any inclination of closely related species to form certain types of association. There is as yet no justification for relating the specific composition of mixed populations to any nearer interrelationship. 2. I sought in vain to explain the inclination of the species to live together. Nor is there any reason to suppose an interdependence of the' species: they derive no advantage or disad-

33 vantage from the presence or absence of other flea species. The food requirements of the larvae and fully developed fleas hardly differ. And i t would be unreasonable to suppose that some smaller-sized flea would have less food because of the presence of the large-sized Hystrichopsylla talpae orientális. There is no information to suggest that any one of the species could interfere with the feeding or reproduction of others. (One might presume disadvantages arising from the presence of certain species to coexisting species in only one case: viz, i f some favourable circumstances enabled one species to overproliferate in a habitat /nest, burrow, l i t t e r /, though even then the members of the minority species might obtain less food only in proportion to the number of individuals of the species.) 3. A comparison of T a b l e s 5 and 6 fully identifies those species which seldom or never associate with other species. The following were never found in specific associations: Chaetopsylla rothschildi, Ischnopsyllus octactenus, I. simplex mysticus, I. s. simplex, Leptopsylla segnis, Rhinolophopsylla u. unipectinata..the species Archaeopsylla e. erinacei,ischnopsyllus variabilis, Nycteridopsylla eusarca, Rhadinopsylla pentacantha, and Rh.strouhali occurred together with other species in one case only. The majority of these species, having nottendency to appear in the company of others, have been collected a few times only; most of them are probably truly rare, but there may be some which appear to be rare only because their hosts also occur infrequently in Hungary, or because of some aspect of collection. I t should be emphasized again that the number of rare flea species is considerably less than we think and we are rather inclined to regard those which are seldom captured as rare whereas the cause of this phenomenon actually lies in faults of the collecting apparatus or techniques. I f the species mentioned above are captured several times in future collections, they might also be found to coexist with other species. However, in the case of species which are frequent and s t i l l f a i l to occur in the company of others, the possibility remains that,for some unknown reason they have no inclination"

34 to participate in specific associations. Such are I.octactenus, h.segnis, A.erinacei, and N.eusarca. I have collected the hosts (Erinaceus europaeus roumanicus Barr.-Ham., and Mus musculus spicilegus Perényi) of two of these species (A.erinacei and L. segnis) in moderate numbers, but I have never found them to occur together with other flea species, even though both hosts have ample opportunity, both in their nests and sphere of activity, to acquire a number of other parasites. One might therefore infer with some justification that these two flea species are for some reason less inclined to coexist with others or that they shun the presence of the latter, or, indeed, vice versa. One should also recall that Ct.caucasicus, the flea of the mole-rat, has never been found on gophers though a l l flea species of that latter animal frequently occur on the mole-rat. The problem of host specificity in fleas, perhaps depending to some extent on the type of food (blood) may be clarified by laboratory experiments. 4. The la3t and most difficult question involves the actual ecological, or some other, cause of the coexistence of flea species on a host,or the possibility of its chance (spontaneous) nature. The specific fleas of the hosts are known, and we also have information on animals which could serve as hosts for the given flea species. Those fleas considered most specific appear incidentally though seldom, on other than their true" hosts, but these are merely occasional phenomena. I f, however, the host changes its haunt (mainly its site of reproduction) for any length of time,and i f this be the habitat of hosts supporting other flea species, then those appearing later on the immigrant host can hardly be defined as merely occasional parasites. (The number of individuals of a flea species found on hosts and in their nests indicates the occasional or specific nature of the flea species.) I t was many times my experience that the flea fauna of a host found in an environment different from its usual habitat (nesting site) is not composed of the species to be expected.

35 I should like to put forward a most characteristic example (not contained in the Tables, because the case was studied after the period under investigation). In October, 968, in the southern part of the country (Madaras) an ermine (Mustela erminea aestiva Kerr) made its subterranean quarters in a field much infested by gophers. We succeeded in shooting the animal. Fourteen of the fifteen flea specimens found on i t proved to belong to the specific fleas of the gopher (Citellophilus martinoi and Ctenophthalmus orientális). This comparatively large number of individuals also indicates that the flea species mentioned do not attach themselves so much to their hosts as to the habitat of the nesting site. DUDICH (939) has already remarked that one usually encounters difficulties when attempting to establish the biotopes of species. This statement is especially true of parasites and thus of fleas. What then, are the biotopes of fleas? Environment in the stricter or wider sense of the term? There are species which occur on animals living in pastures and agricultural areas of the plains, and ethers on forest mammals. Some are more frequent in arid, others in damper habitats. I failed to find any.demonstrable connection between the plant cover, the forest stand,and the occurrence of flea species; - i f the hosts are present, and their burrows and nests suitable for invasion by and reproduction of fleas, we might expect the appearance of certain species. Hence the biotope of flea species is not the wider but the more restricted habitat; the small area where they reproduce, lay their eggs, the larvae find their food, and the pupae transform into imagos. In the search for the biotope of the fleas we have thus arrived at the lairs of their hosts, be they subterranean burrows, holes, or nests constructed on the ground, on clumps of grass, shrubs, or trees. (Tne bizarre idea that the biotope of the flea species is the host itself should be rejected at once; this assumption may be justified only in some cases of endoparasitism.) The biotopes mentioned above can be delineated rather satisfactorily and also assigned to the biocoenological horizons of the temperate regions es-

36 ta"blished by BALOGH (953). The communication of the several species usually occurs between neighbouring horizons only. Rare exceptions naturally occur, e.g. a squirrel flea (M.s.sciurorum) found on one occasion on a fox; i t might have arrived on its unusual host either by having fallen from the nest or in the course of the food-chain, or eventually from the ground, as, for instance, was the squirrel flea collected by soil surface extraction methods. On the basis of my experiences gained in the course of collections and investigations, I venture to state that a great majority of flea species appear to be more bound to certain types of nests and their microclimate than to the hosts themselves. I shall try to prove this assertion by some examples. The fox and the badger inhabit the same biotope where in most of our mouse species also occur, but I have never found a flea species parasitizing mice on either the fox or the badger. Similarly, the characteristic fleas of these latter two hosts have never occurred on mice,although there are innumerable opportunities for the exchange or transmission of fleas living on these hosts. That is not to say, however, that one or two mouse fleas will not be found on fox or badger, or vice versa, though this can only be an occasional phenomenon. (Even in the literature there are only a few records of such an occurrence.) In my opinion, lack of such interchange of flea species is not due simply to differences in the hosts, but to the fact that nests of mice have a completely different microclimate from those of the deep subterranean burrows of the fox and the badger. Incidentally, PETJS (953) has already pointed out the importance of the correlation between the occurrence of certain flea species and the environmental factors, so my experiences would seem to corroborate his statements. A certain kind of correlation in the systematical relationship of the hosts and their fleas (and also their other parasites) appears to be frequently demonstrable.this statement apparently contradicts my previous assertion, because i t might be supposed

37 that the fleas of the hosts mentioned above do not mingle on account of the different systematical categories of the host species. But then why do the fleas of the badger occur on the fox, and vice versa? - the two host species are assigned to two different mammalian families. The mice stand at least as far removed from the shrews as from the fox and the badger, but the fleas characterizing the one s t i l l occur on the other. The objection that the flea species of the mice and the shrews s t i l l differ may therefore be apparently justified - these hosts live in a common habitat and in subterranean burrows of not too differing types. In the course of making my collections, I have in fact found mouse and shrew species within the same habitat in many places but the shrews invariably occurred in the vicinity of the moister areas (smaller bodies of standing waters, shores of brooks), so the microclimate of their nests also is probably different from that of mice. Evidence supporting my thesis can be found in the mouse family. The flea Gt.a.assimilis is more or less specific to the field vole (Microtus a.arvalis Pallas). I t is known that the field vole inhabits rather dry meadows, pastures,. and agricultural fields, and so the environment of its nests is drier than that of mice (Apodemus, Clethrionomys, Pitymys, Micromys). Now I have collected Ct«assimilis rather frequently on mouse species i f their habitats were adjacent to dry areas, and even in cases when I found no fieldvoles in the vicinity. I t might be assumed in this case - and with justification - that Ct.assimilis is bound not so much to M.arvalis as to the microclimate of its nests and their stricter environment. Though I have discussed here only occurrences jof fleas parasitizing mammals, I should like to substantiate my theory by an example drawn from avian fleas. A great number of hosts, not too far removed from one another systematically, of avian flea species are known in Hungary. There is one exception, namely Ceratophyllus styx styx Rothschild, demonstrated only from the sandmartin (Riparia r i - paria L.); nor did I find any other flea species on this host. Even PEUS's (968) abundant research material failed to produce

38 other fleas from this host, and C.styx was found in only a single case on the stonechat (Oenanthe oenanthe L.), which occasionally nests in sites similar to the subterranean nesting cavities of the sand-martin. Thus we have the specific flea of a nest type which differs fundamentally from the nests of other species. It would be further proof of my assumption if C.s.styx were found in the similarly constructed nest of the bee-eater (Merops apiaster L.). These examples suffice to prove the merit of this type of investigations and to draw attention to my hypothesis. Finally, I should like to add some remarks concerning my work and future research. Though numerical data referring to individuals of coexisting species are available, I have discussed only cases of specific associations. In later communications I intend to consider, whose studying flea populations of a given host species, the numerical ratios and sex distribution sex of the parasitic species. A more intensive collecting of nests is also indicated, as well as the evaluation of the occurrence of fleas by seasons (YYSOTSKAJA, 967), the comparison of flea species found on the hosts and their nests (JURIK, 968), and demonstration of correlations between the development of fleas found on the hosts and their nests (ROTHSCHILD, 967). One or more of these aspects have already been examined- by the abovecited authors and by others, without, however, all aspects being considered in the course of a given investigation. Many questions might be answered if it could be determined by laboratory investigations how far the flea species found in the samples show food preferences. Also, as already stated, I attach great importance to moisture, humidity, and temperature conditions affecting the several biotopes.the investigation of these latter seems to present most difficulties, since after the exposure of the nests the original microclimatic conditions can no longer exist. If a technical solution to this problem were assured, we would be much nearer a demonstration of the nestspecificity of flea species.

39 The present paper discusses the most frequent variations of coexistence or common occurrences of flea species occurring on mammals in Hungary. The coexistence of fleas on mouse and shrew species, on the fox and the badger, on the mole-rat and the gopher, as well as the mole and the bat species, appeared in (comparatively) most definite and frequently repeated compositions. No systematic relationship could be demonstrated between the flea species coexisting on the hosts.it could not be proven that any one of the flea species of the various associations was especially inclined to occur in specific compositions. Some species occur very seldom or not at all in the company of other flea species. The most probable conditions affecting the evolution of coexistence, namely the environment of the nests, their postition as to horizon, their structure and microclimate, imply that i f these factors are favourable for the occurrence of certain flea species present in the area,then the possibilities of forming specific associations are available for these species. SZABÓ, I.: A magyarországi emlősök bolháinak (Siphonaptera) együttélési viszonyai A szerző 958-tól 967 végéig 55 emlősfaj 233 példányát és 42 fészkét vizsgálta meg bolhászati szempontból, melyeknek 35,4, illetve 64,3 $-át találta bolhával fertőzöttnek. Sikerült megállapítania az egérfélék-, cickányfélék-, róka és borz-, földikutya és ürge-, vakond- és denevérfélék bolháinak együttes előfordulásait, melyek a vizsgálati anyagban a leghatározottabban és gyakran ismétlődő formában jelentkeztek. A gazdaállatokon együttesen előfordult bolhafajok között nem sikerült származástani összefüggést találni; nem látszik bizonyítottnak, hogy az együttesen előfordult fajok közül valamelyik különösképpen hajlamos lenne a társulásra; a magyar fauna bolhafajai között akad

40 néhány, melyek csak igen ritkán, vagy egyáltalán nem fordulnak elő társulásban. Végül megállapítja, hogy ha a fészkek környezete, szintbeli elhelyezkedése, struktúrája és mikroklímája kedvező a területen előforduló bolhák bizonyos fajainak, akkor e fajok részére fennállanak az együttes előfordulás lehetőségei. Eddigi gyűjtései és anyagának kiértékelése során szerzett tapasztalatok alapján a szerző ugy véli, hogy a bolhafajok jórésze inkább ragaszkodik bizonyos fészektipusokhoz illetve azokmikrokiimájához, mint magához a gazdaállathoz. R e f e r e n c e s BALOGH,J.: A zoocönológia pp alapjai. - Akadémiai Kiadó, Budapest, DUDICH,E.: Élettér", élőhely, életközösség. - Term. Tud. Közi. Pótf JURIK,M.: Pleas of the mole Talpa europaea L. in CzehoSlovakia (Aphaniptera). - Acta Ent. Bohemoslov. 6_5_ , OUDEMANS,A. : Suctorologisch.es aus Maulwurfnestern. - v. Entom. 5j Tijdschr. PEUS,F.: Flöhe. - Akademische Verlagsges.Geest.u.Poriing K.-G.,.Leipzig, pp PEUS,F.: Zur Kenntnis der Flöhe Deutschlands I I. Faunistik und Ökologie der Vogelflöhe (Insecta, Siphonaptera). - Zool. Jb. Syst ROSICKY,B.: A p h a n i p t e r a z i m n i c h h n i z d k r t k a obecného (T a l p a europaea L.) V ruznych biotopech. - Öeskoslov. Parasit ROTHSCHILD,M.: The Rabbit Flea and Hormones. - Penguin Sci.Surv. Biol : 967. SHIT,F.: Siphonaptera collected from moles and their nests at wilp, Netherlands, by Jhr.W.C. van Heurn. - Tijdschr.v.Entom

41 SMIT,P. - SZABÓ,I.: The distribution of Subspecies of Ctenophthalmus agyrtes in Hungary (Siphonaptera:Hystrichopsyllidae). - Ann. Hist.-Nat. Mus. Nat. Hung VYSOTSKAJA,S.: Biocenotitcheskie otnoseni a mezdu ektoparasitami grüsunov i obtateliami ih gnesd. - Parasit. Sbornik. 2_3. I WAGNER,J.: Über die Aphanipterenfauna der Maulwurfnester. - Eonowia Received: I- SZABÓ Zoological Department of the Hungarian Natural History Museum, Budapest, VIII. Baross u. 3.