5

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

István SZABÓ

Zoological Department of the Hungarian Natural History Museum, Budapest

• • •

Several papers d e a l i n g w i t h the n a t u r a l h i s t o r y of f l e a s record the common occurrence of several f l e a species on one host-spe­c i e s . These communications u s u a l l y r e f e r to a l l the f l e a spe­ci e s o c c u r r i n g on several i n d i v i d u a l s of the host species and only r a r e l y does a paper consider the f l e a p opulation found on a s i n g l e host specimen (or i n t h e i r nests, burrows, e t c . ) . I n my o p i n i o n i t would be more i n f o r m a t i v e i f the f l e a populations were evaluated on the basis o f s i n g l e i n d i v i d u a l s of the same host species, because the constant haunts of animals the loc a ­t i o n o f t h e i r n ests, the c o n d i t i o n s of t h e i r é.ssociutions w i t h other neighbouring species, and so on, d i f f e r even w i t h i n a small area of the h a b i t a t , and a l l these f a c t o r s i n f l u e n c e the common occurrence of f l e a species.

• . The common occurrence o f f l e a species aroused my i n t e r e s t some time ago. Examining.my m a t e r i a l c o l l e c t e d from 1958 t i l l the end o f 1967, I found t h a t although given host, usually c a r r i e s a s i n g l e f l e a species the common occurrence o f several f l e a spe­cies i s not rare.On one host species (or i n i t s nest, m a t e r i a l ) , I found one f l e a species i n 66.5 per cent o f cases, two I n 26.8 fo, three i n 5.8 and 4 i n 0.9 Since i n o n e - t h i r d o f the cases more than one f l e a species was found to occur on a host specimen, a study of the con d i t i o n s of coexistence neemed

worthwhile, with s p e c i a l regard to the following pointB of i n ­quiry: i s there any systematic r e l a t i o n s h i p between the species occurring together; are they i n any way interdependent on one another; do c e r t a i n species favour coexistence; are there spe­c i e s which seldom i f ever occur c o e x i s t with others; f i n a l l y , i s there any apparent reason, e c o l o g i c a l or otherwise, for the common occurrence of species or i s i t merely a spontaneous or chance phenomenon?

Materials and Methods

Since methods for c o l l e c t i o n of f l e a s or t h e i r hosts are w e l l known only a summary of methods used i n t h i s study w i l l be given.Mouse and shrew species were c o l l e c t e d by l i v e and l e t h a l t raps, dormice and hedgehogs by hand, moles and mole-rats by subterranean l e t h a l t r a p s , gophers e i t h e r by hand (flooding with water) or by shooting, bats by nets or by hand ( s i n g l e specimens), a l l other mammals ( s q u i r r e l s , foxes, badgers, w i l d ­c a t s , martens) by shooting. Immediately a f t e r capture, the host specimens were placed i n a sealed cotton sack and l a t e r i n chloroform v e s s e l s ; thus a l l f l e a s present on the host a t the moment of capture were securely c o l l e c t e d a f t e r narcotization. Great weight was l a i d on gathering the f l e a s separately by host specimen, except for cases when i n d i v i d u a l s of the same host species were caught i n traps on further than a few steps from one another, since i n a l l p r o b a b i l i t y t h e i r f l e a s occurred on members of the same family l i v i n g i n a common subterranean burrow.

The fundamental requirement for e s t a b l i s h i n g conditions of co­existence of f l e a species per i n d i v i d u a l host i s a recording system ( P i g . l ) which allows one to discover, a t any time, the numbers and species of f l e a s o r i g i n a t i n g from a given host specimen. The s e r i a l inventory number must a l s o be entered for the host i n question ( P i g . l ) since without t h i s , had the f l e a specimens been preserved e i t h e r i n alcohol or mounted on a

h* cm

ro

00 H

S e r i a l Number sorszám

Host Species gazdaállatfaj

CT CD O P H fcJ* CO p m H> 0» P o c_i. p.

ind.no. - db. Date of

C o l l e c t i o n gyűjtés i d e j e

L o c a l i t y lelőhely

Co l l e c t o r gyűjtő

Remarks megjegyzés

Table l - l . táblázat.

Research M a t e r i a l - Vizsgálati anyag

Host Species gazdafaj

M a t e r i a l I n v e s t i g a t e d vizsgált anyag

Nests, Burrows, and Holes

I n v e s t i g a t e d vizsgált fész­kek, kotorékok

és odúk Host Species gazdafaj

Total

összesen

Positive

pozitiv

S-S I I I I

Total

összesen

Positive

pozitiv

1 2 3 4 5 ! 6 7 Apodemus agrárius 109 30 79 27,5 Apodemus f l a v i c o l l i s 473 155 318 32,7 6 5 Apodemus s y l v a t i c u s 75 31 44 41,7 A r v i c o l a t e r r e s t . s c h . 5 - 5 0 1 -B a r b a s t e l l a b a rbast. 8 3 5 37,5 Canis familiáris 8 8 - 100,-C i t e l l u s c i t e l l u s 31 20 11 64,5 2 2 Clethrionomys g l a r e o l . 220 117 103 53,2 2 2 C r i c e t u s c r i c e t u s 6 - 6 0 1 -C r o c i d u r a leucodon 6 - 6 0 Crocidura suaveolens 1 - 1 0

1 • 2 3 4 5 6 7

Dryomis nite d u l a 1 1 - 100,- 3 1

Eptesicus serotinus 4 4 - 100,-

Erinaceus europaeus r . 18 7 11 38,8

F e l i s catus 3 3 - 100,-

F e l i s s y l v e s t r i s 1 1 - 100,-

G l i s g l i s 4 4 - 100,- 1 1

Lepus europaeus 3 1 2 33,3

Martes foina 3 3 - 100,-

Mêles mêles 11 11 - 100,-

Micromys minut.prat. 109 3 106 2,7 1 -Microtus a g r e s t i s 25 6 19 24,-

Microtus a r v a l i s 88 42 46 47,7 10 7

Microtus oeconom.méh. 33 9 24 27,3

Minlopterus s c h r e i b e r s i 154 2 152 1,3

Mus musculus s p i c i l e g u s 72 40 32 55,5 2 2

Muscardinus a v e l l a n a r . 11 7 4 63,6 11 5

Mustela n i v a l i s 2 1 1 50,-

Mustela putorius furo 2 2 - 100,-

Mustela putorius hung. 1 1 - 100,-

Myotis brandti 1 1 - 100,-

Myotis dasycneme 1 1 - 100,-

Myotis daubentoni 2 - 2 0 •

Myotis bechsteini 2 - 2 0

Myotis myotis 42 7 35 16,6

1 2 3 . 4 5 6 7

Myotis n a t t e r e r i 8 3 5 37,5 Myotis oxygnathus 206 47 159 22,8 Neomys anomalus m i l l e r i 14 8 6 57,1 Neomys fo d i e n a 7 5 2 71,4 Nyctalus l e i s l e r i 6 - 6 0 Nyctalus n o c t u l a 20 16 4 80,-Ondatra z i b e t h i c u s 1 1 - 100,-P i p i s t r e l l u s p i p i s t r . 27 21 6 77,8 Pitymya subterraneus 27 15 12 55,5 Plecotus a u r i t u s 2 - 2 0 Plecotus a u s t r i a c u s 14 11 3 78,5 Rattus norvegicus 2 - 2 0 Rhinolophus ferrumequ. 44 2 42 4,5 Rhinolophus h i p p o s i d . 2 - 2 0 Sciurus v u l g a r i s f u s e . 33 27 6 81,8 2 2 Sorex araneus 280 82 198 29,3 Sorex minutus 13 1 • 12 7,7 Spalax leucodon 18 16 2 88,9 Talpa europaea 49 28 21 57,1 Vulpes vulpes c r u c i g e r a 23 22 1 95,6

2,331 826

35,43 1°

1,505

64,57 fo

42 27

64,28 %

s l i d e (F i g. 2 ) , i t would "be imp o s s i b l e t o e s t a b l i s h l a t e r whether the f l e a s i n the c o l l e c t i o n come from the same host i n ­d i v i d u a l or n o t .

During the t e n year p e r i o d I had occasion t o examine 2331 spe­cimens and 42 nests or subterranean burrows o f 55 mammalian species. Fleas were found on 35.43 per cent of the hosts and i n 64.28 per cent o f t h e i r n e s t s ( T a b l e l ) . The Table shows the e x t e n t o f i n f e c t i o n o f the s e v e r a l host species.One hundred per c e n t , o r complete l a c k o f i n f e c t i o n by f l e a s occurred mainly i n those host species f o r which I was unable t o c o l l e c t suf­f i c i e n t m a t e r i a l . Presumably, a l a r g e number of specimens would not show t h i s h i g h degree or complete l a c k o f i n f e c t i o n . On mouse and shrew species and on moles and mole-rats t h e r e occur probably more f l e a s than i t was able t o e s t a b l i s h . This stems from the f a c t t h a t most o f these species were c o l l e c t e d by l e t h a l t r a p s and i t i s p o s s i b l e t h a t , d e s p i t e s e v e r a l examina­t i o n s o f t h e • t r a p s per day, a p a r t o f the f l e a p o p u l a t i o n had already l e f t the c o o l i n g hosts p r i o r t o c o l l e c t i o n . However, I estimate t h i s source o f e r r o r t o be not hi g h e r than 5-10 j6. A f t e r capture o f i n d i v i d u a l s o f other mammalian species, I was able t o prevent s c a t t e r i n g o f the f l e a s . S i m i l a r l y , t h e m a t e r i a l of n e s t s , burrows, holes were i n a l l cases placed immediately i n t o completely sealable bags and then t r a n s f e r r e d i n t o an ex­t r a c t o r ( F i g . 3) from which t h e p a r a s i t e s are unable to escape.

Results

Occurrences o f s i n g l e f l e a species

Before d e s c r i b i n g the coexistence o f d i f f e r e n t f l e a species on a s i n g l e h o st animal I propose t o l i s t t he cases o f s i n g l e oc­currences o f f l e a species as repeated reference w i l l l a t e r be made t o these. The number o f cases i s i n some i n s t a n c e s h i g h e r than t h a t o f the l i s t e d l o c a l i t i e s , because c o l l e c t i o n s were made a t s e v e r a l separate t i m e s .

F i g . 3 - 3 . ábra.

E x t r a c t o r Féazekfuttató

Table 2 - 2 . táblázat.

Occurrence o f S i n g l e Flea Species B o l h a f a j o k egyenkénti előfordulása

Serial Number

sorszám

S iphonaptera Host - gazdaállat

L o c a l i t y lelőhely

Serial Number

sorszám -p CO CQ CO

Species - f a j i m cd o

-p CO 03 CO Species - f a j i

CO m cd o

L o c a l i t y lelőhely

1 2 3 4

1 Archaeopsylla 5 Erinaceus eur.roum.

5 Misina-tető Monor Orgovány Tompa

2 Cerat ." t r i b u l i s

1 C i t e l l u s c i t e l l u s

1 Tahitótfalu

3 Chaetops. globiceps

4 F e l i s s y l v e s t r i s

Vulpes vulpes 1

3

Diósjenő Kaposvár Szentmártonkáta P i l i s s z e n t k e r e s z t

4 Chaetops. r o t h s c h .

2 Martes f o i n a 2 Deszkáspuszta Mátraszentimre

5 C i t e l l o p h . m a r t i n o i

1 C i t e l l u s c i t e l l u s

1 Gyulafirátót

6 C i t e l l o p h . simplex

1 C i t e l l u s c i t e l l u s

1 Ha j dubago s

7 Ctenoceph. canis

2 Canis familiáris

lepus europaeus

1 1

Budapest Boly

8 Ctenoceph. f e l l s

4 Canis familiáris

F e l i s catus Mustela put or.

f u r o

1 2 1

Budapest Monor Budapest Gödöllő

1 2 3 4 9 Ctenoph.ag. 49 Apodemus 7 K i s b a l a t o n

"bosnic. agrárius Fityeház Zajk

Apodemus 18 Németbánya f l a v i c o l l . K i s b a l a t o n

Szabadegyháza Bakonybel Kiszépalmapuszta őriszentpéter Pálihálás

Apodemus 2 Pityeház s y l v a t i c u s Buesuta

Clethrionomys 11 Németbánya g l a r . K i s b a l a t o n g l a r .

Kaposmérő Szalafő Bucsuta

Micromys 2 Tatabánya m i n u t . p r a t . Németbánya

Mic r o t u s 2 K i s b a l a t o n a g r e s t i s

M i c r o t u s 2 Németbánya a r v a l i s

M i c r o t u s 1 K i s b a l a t o n oecon.méh.

Pitymys 4 Németbánya s u b t e r r a n . Iharkút

10 Ctenoph.ag. 7 Apodemus 6 Sarkadremete eurous f l a v i c o l l .

Mus musculus 1 Sarkadremete s p i c . T

I I Ctenoph.ag. 4 Apodemus 1 Tákos k l e i n s . agrárius

Apodemus 2 Táko s s y l v a t i c u s

M i c r o t u s 1 Lónya a r v a l i s

12 Ctenoph.ag. 7 Apodemus 3 Kisnána peusian. f l a v i c o l l . peusian.

Clethrionomys 4 Deszkáspuszta g l a r . K i snána

13 Ctenoph. 16 Apodemus 2 Hajdubagos a s s i m i l i a f l a v i c o l l . Tákos

Clethrionomys 2 Tahitótfalu g l a r . Deszkáspuszta

1 2 5 4 M i c r o t u s

a r v a l i s

Neomys fod i e n s

Sorex araneus Talpa

europaea

8

1 1 2

Orgovány K i s h a l a t o n Bugyi Németbánya Lippó Békéscsaba Nagykovácsi K i s b a l a t o n K i s b a l a t o n Sarkadremete Kisszépalmapuszta

14 Ctenoph. caucasicus

1 Spalax leucodon

1 Hajdubagos

15 Ctenoph. congener

3 Apodemus f l a v i c o l l .

Clethrionomys g l a r .

1 2

Németbánya Németbánya Bakonybél

16 Ctenoph. orientális

. 6 Apodemus f l a v i c o l l .

C i t e l l u s c i t e l l u s

Mustela n i v a l i s

1

4

K i snána Hollóháza Szabadszállás Fácánkert Bugac Ócsa

17 Ctenoph. s o l u t u s

14 Apodemus f l a v i c o l l .

Apodemus s y l v a t i c u s

Clethrionomys g l a r .

11

2 1

Gombáspuszta Nagysomlyóhegy Deszkáspuszta A l c s u t Pisznice-hegy Gézaháza Kisszépalmapuszta Kisnána Budakeszi L i t k e Kistolmács Deszkáspuszta

18 D o r a t o p s y l l a d.das.

10 Neomys f o d i e n s

Sorex araneus 1

9

Németbánya Németbánya Deszkáspuszta Szalafő

1 2 3 4 19 H y s t r i c h o p s .

t a l p . o r . 5 Apodemus

agrárius 1 K i s b a l a t o n

Apodemus f l a v i c o l l .

1 Cserfekvés Apodemus f l a v i c o l l .

M i c r o t u s 1 K i s b a l a t o n oecon.jméh.

Ondatra 1 Kapuvár z i h e t h i c u s

Talpa 1 Iharkút europaea

20 I s c h n o p s y l l u s 6 Myotis myotis 1 Pilismarót elong . Nyctalus

n o c t u l a 5 Rétszilas

Szikra-Tőserdő Lillafüred K i s b a l a t o n

21 I s c h n o p s y l l u s 15 Myotis myotis 2 Legény-barlang hexact. Myotis

oxygnathus

P i p i s t r e l l u s p i p i s t r .

6

1

Ördöglyuk-barlang Pálvölgyi-barlang S olymári-barlang Aba l i g e t i - b a r l a n g Legény-barlang Szekszárd

Plecotus 6 Tata a u s t r i a c u s Gyöngyös Budapest P i l i s Dorog Tompa

22 I s c h n o p s y l l u s i n t e r m .

17 Eptesicus s e r o t i n u s

M i n i o p t e r u s s c h r e i b .

Myotis dasycneme

Myotis myotis

Myotis oxygnathus

4

1

1

3

8

Pilismarót Budapest Alpár Németbánya A b a l i g e t i - b a r l a n g Szikra-Tőserdő Pilismarót Eger P i s z n i c e i - b a r l a n g Ördöglyuk-barlang Pálvölgyi-barlang Nagymaros A b a l i g e t i - b a r l a n g Leány-barlang Gyöngyös

1 2 3 4

23 I s c h n o p s y l l u s o c t a c t .

6 P i p i s t r e l l u s p i p i s t r .

6 Sopron Lillafüred Németbánya Bakonybél

24 Is c h n o p s y l l u s simpl.myst.

1 Myotis b r a n d t i

1 Baláta-tó

25 I s c h n o p s y l l u s s i m p l . s i m p l .

2 Myotis n a t t e r e r i

2 Leány-barlang Legény-barlang

26 L e p t o p s y l l a segnis

17 Apodemus f l a v i c o l l .

Mus muscuius s p i c i l .

2

15

Tákos Deszkáspuszta Orgovány Pel3Őgöd Sarkadremete K i sszépalmapuszta Budapest Tákos Pálihálás

27 Megabothris t u r b i d .

3 Apodemus f l a v i c o l l .

Clethrionomys g l a r .

M i c r o t u s a r v a l i s

1 1

1

őriszentpéter őriszentpéter Tákos

28 Megabothris w a l k e r i

2 M i c r o t u s a g r e s t i s

Sorex minutus

1

1 K i s b a l a t o n K i s b a l a t o n

29 Monopsyllus s c i u r .

26 Apodemus f l a v i c o l l .

Dryamis n i t e d u l a

G l i s g l i s

Mêles mêles Muscardlnus

a v e l l a n . M y o t i s

oxygnathus Sciurus v u l g .

f u s c o a t .

3

1

5

1 1

1

14

Gödöllő Iharkút Gödöllő Deszkáspuszta Pisznice-hegy Budakeszi Budapest Mecsek-hg Gödöllő Legény-barlang Diósjenő Pilisszántó Deszkáspuszta Budapest

1 2 3 4 Sciurus v u l g .

f u s c o a t . 14 Csákvár

Budakeszi Z i r c Németbánya Gödöllő Mecsek-hg.

30 No so pay U u s f a s c i a t .

9 Apodemus f l a v i c o l l .

Apodemus s y l v a t i c u s

M i c r o t u s a r v a l i s

Mua muaculua s p i c i l .

2 3 1 3

Pisznice-hegy Sarkadremete Tahitótfalu Sarkadremete Békéscsaba

Sarkadremete Békéscsaba

31 N y c t e r i d o p s . eusarca

3 Nyctalu s n o c t u r a

3 l e n g y e l Budapest

32 N y c t e r i d o p s . p e n t a c t .

7 B a r b a s t e l l a "barbast.

M y o t i s myotis M y o t i s

oxygnathus Plecotu s

a u s t r i a c u s

1 1 1 4

Vereshegyi-barlang A b a l i g e t i - b a r l a n g ördöglyuk-barlang Üllő Gödöllő Királyrét

33 Palaeops. k o h a u t i

3 Talpa europaea

3 K i s b a l a t o n Iharkút

34 Palaeops. s i m i l i s

10 Talpa europaea

10 Németbánya Szabadegyháza Deszkáspuszta Bakonybél Iharkút

35 Palaeops. . s o r . r o s i c .

18 Apodemus agrárius

Clethrionomys g l a r .

Micromys m i n u t . p r a t .

Neomys anomal . m i l l .

Sorex araneus

1 1 1 4

11

K i s b a l a t o n K i s b a l a t o n Pákozd Németbánya K i s b a l a t o n Pákozd K i s b a l a t o n Pákozd

36 Peromyscops. f a l l a x

3 Clethrionomys g l a r .

1 Németbánya

1 2 3 4 Mi c r o t u s 1 Németbánya

a r v a l i s Pitymys 1 Németbánya

s u b t e r r a n . 37 Pul ex 15 Canis 5 Orgovány

i r r i t a n s familiáris Hont Sárcsikut Kapuvár Bugac

P e l i s catus 1 Csákvár Meie3 meles 3 Drégelypalánk

Lónya Mustela 1 Hajdúbagos

putor.hung. Hajdúbagos

Vulpes vulpes 5 Szigetmonostor c r u c i g . Szentendre

Haj dubagos Kapuvár Mihálygerge

38 Rhinolophops. , 2 Rhinolophus 2 Gyula u n i p. ferrume. A b a l i g e t i - b a r l a n g

I n T a b l e 2 c e r t a i n species occur i n only a few cases, but t h i s i s not an expression o f t h e i r r a r i t y since most o f them occur - as shown i n T a b l e 3 - more f r e q u e n t l y i n the company of oth e r species.On the b a s i s of m a t e r i a l c o l l e c t e d so f a r , the f o l l o w i n g f l e a species o f mammals may be considered r a r e : Chaetopsylla r o t h s c h i l d i Kohaut; I s c h n o p s y l l u s simplex simplex R o t h s c h i l d ; I . simplex mysticus Jordan; I . v a r i a b i l i s (Wagner); Megabothryis w a l k e r i ( R o t h s c h i l d ) ; P a l a e o p s y l l a ko­ h a u t i Dampf; Peromyscopsylla f a l l a x ( R o t h s c h i l d ) ; Rhadinopsylla isacantha isacantha ( R o t h s c h i l d ) ; R. pentacantha ( R o t h s c h i l d ) ; R. s t r o u h a l i Smit, h i t h e r t o found as a s i n g l e specimen only and i d e n t i f i e d c o n d i t i o n a l l y even by i t s d i s c o v e r e r ; Rhinolopho- p s y l l a u n i p e c t i n a t a u n i p e c t i n a t a (Taschenberg), and T a r s o p s y l l a octodecimdentata octodeclmdentata ( K o l e n a t i ) . As the host spe­c i e s o f these f l e a s are r a r e i n Hungary, t h e i r i n f r e q u e n t cap­t u r e i s q u i t e understandable.

Table 3 - 3 . táblázat.

Common Occurrence of Plea Species - Bolhafajok együttes előfordulása Serial

Numb

er

sorszám Common Occurrence

of F l e a Species Együttesen elő­

forduló bolhafajok Case eset

Host - gazdaállat Species - f a j

Case eset

L o c a l i t y lelőhely

1 2 3 4 5 6

1 Archaeopsylla e r i n . Pulex i r r i t a n s

1 Vulpes vulpes crue. 1 ócsa

2 Ceratoph.pullatus Ctenoph.ag•eurous Nosopsyllus f a s c .

1 Apodemus f l a v i c o l l . 1 Sarkadremete

3 Ceratoph.pullatus Monopsyllus s c i u r .

1 Muscardinus a v e l l . 1 Gödöllő

4 Cerato p h . t r i b u l i s Monopsyllus s c i u r .

1 Sciurus vulg.fuse. 1 Szeleste

5 Cera t o p h . t r i b u l i s Nosopsyllus f a s c .

1 C i t e l l u s c i t e l l u s 1 Tahitótfalu

6 Chaetops.globicepB Chaetops.trichosa Paracéras melis Pulex i r r i t a n s

1 Vulpes vulpes crue. 1 Deszkáspuszta

1 2 3 4 5 6

7 Chaetops.globiceps Pulex I r r i t a n s

6 Vulpes vulpes crue. . 6 Vokány Pécsudvar Inárcs Rákoscsaba Budakalász Drégelypalánk

8 Chaetops.globiceps Ctenoceph.canis Paraceras melis Pulex i r r i t a n s

1 Vulpes vulpes crue. 1 Mecsek-hg.

9 Chaetops,globiceps Monopsyllus s c i u r .

1 Sciurus v u l g . f u s e . 1 Németbánya

10 Chaetops.globiceps Monopsyllus s c i u r . Pulex i r r i t a n s

1 Vulpes vulpes crue. 1 Németbánya

11 Chaetops.globiceps Paraceras melis

1 Vulpes vulpes crue. 1 Deszkás-puszta

12 Chaetops.globiceps Paraceras melis Pulex i r r i t a n s

1 Meies meles 1 Ba jna

13 Chaetops.trichosa Paraceras melis Pulex i r r i t a n s

2 Meies meles o Sárcsikut L i t k e

14 Chaetops.trichosa Pulex i r r i t a n s

7 Canis familiáris Meies meles

1 Drégelypalánk Pusztavám S za ba d egy há za Egervár

M3

1 2 3 4 5 6

•

Vulpes vulpes crue. 3 Drégelypalánk Belvárgyula Kutberek

15 CItelloph.martinoi Ctenoph.orientális

3 C i t e l l u s c i t e l l u s 3 Hollóháza

16 Citelloph.simplex Ctenophth.assimllis

1 C i t e l l u s c i t e l l u s 1 Hajdubagos

Citelloph.simplex Ctenoph.caucasicus Ctenoph.orientális

1 Spalax leucodon 1 Ha jdubagos

18 Citelloph.simplex Ctenoph.orientális

3 C i t e l l u s c i t e l l u s 3 Ha jdubago-s Bátorliget

19 Ctenoceph.canis Ctenoceph.felis Pulex i r r i t a n s

1 Canis familiáris. 1 Biharugra

20 Ctenoph.ag.hosnic. Ctenoph.assimilis

4 Clethrionomys g l a r . Microtus oecon.méh.

3

1

B a l a t o n l e l l e Kisbalaton Kisbalaton

21 Ctenoph.ag.bosnic. Ctenoph.assimilis Megabothris walker!

» 1 Clethrionomys g l a r . 1 Kisbalaton

22 Ctenoph.ag.hosnic. Ctenoph.assimilis Palaeops.sor.ros.

1 Sorex araneus 1 Kisbalaton

1 2 3 4 5 6

23 Ctenoph.ag.bosnic. Ctenoph.congener

11 Apodemus f l a v i c o l l . Clethrionomys g l a r .

Neomys f o d i e n s

1 9

1

Zajk Sárcsikut Németbánya Kisszépalmapuszta Németbánya

24 Ctenoph.ag.bosnic. Ctenoph.congener Ctenoph.solutus

1 C l e t h r i o n o m y s . g l a r . 1 Sárcsikut

25 Ctenoph.ag.bosnic. Ctenoph.congener Doratops.d.dasycn.

2 Apodemus f l a v i c o l l . Clethrionomys g l a r .

1 1

Németbánya Németbánya

26 Ctenoph.ag.bosnic. Ctenoph.congener H y s t r i c h o p s . t a l p . o r .

1 Clethrionomys g l a r . 1 Gombáspuszta

27 Ctenoph.ag.hosnic. Ctenoph.congener Hy s t r i c h o p s . t a l p . o r . Peromyscops.fallax

1 Clethrionomys g l a r . ] Iharkút

28 Ctenoph.ag.hosnic. Ctenoph.congener Peromyscops.fallax

3 Clethrionomys g l a r . Pitymys subterraneus

2 1

Néme tbánya Németbány

29 Ctenoph.ag.hosnic. Ctenoph.congener Rh.vàtnops .isacantha

1 Clethrionomys g l a r . 1 Iharkút

30 Ctenoph,ag.boonlc Ctenoph.orientális

1 Apodemus f l a v i c o l l . 1 Mecsek-hg.

1 2 3 4 5 6

31 Ctenoph.ag.bosnic Ctenoph.solutus

7 Apodemus f l a v i c o l l .

Clethrionomys g l a r .

6

1

Sárcsikut Németbánya Pisznice-hegy Bakonynána Budakeszi Kaposmérő

32 Ctenoph.ag.bosnic. Doratops.a.dasycn.

2 Neomys a n o m a l . m i l l . Pitymys subterraneus

1 1

Németbánya Németbánya

33 Ctenoph.ag.hosnic H y s t r i c h o p s . t a l p . o r .

2 M i c r o t u s a r v a l i s Mus musculus s p i c .

1 1

Németbánya K i s b a l a t o n

34 Ctenoph.ag.bosnic Megabothris t u r b i d .

9 Apodemus f l a v i c o l l . Clethrionomys g l a r .

1 7

őriszentpéter őriszentpéter Zajk Szalafő K i s b a l a t o n M i c r o t u s a g r e s t i s 1

őriszentpéter őriszentpéter Zajk Szalafő K i s b a l a t o n

35 Ctenoph.ag.bosnic. Megabothris t u r b i d . Monopsyllus s c i u r .

1 Clethrionomys g l a r . 1 őriszentpéter

36 Ctenoph.ag.hosnic. Megabothris w a l k e r i

1 M i c r o t u s oecon.méh. 1 K i s b a l a t o n

37 Ctenoph.ag.hosnic. Megabothris w a l k e r i Palaeops.sor.ros.

1 Neomys f o d i e n s 1 K i s b a l a t o n

38 Ctenoph.ag.hosnic Monopsyllus s c i u r .

1 Apodemus f l a v i c o l l . 1 őriszentpéter

39 Ctenoph.ag.hosnic. Nosopsyllus f a s c . Palaeops.sor.ros. Peromyscops.fallax

1 Apodemus f l a v i c o l l . 1 Németbánya

1 2 3 4 5 6 40 Ctenoph.ag. "bosnic.

Palaeops.sor.ros. 3 M i c r o t u s a g r e s t i s

Neomys a n o m a l . m i l l . Talpa europaea

1 1 1

K i s b a l a t o n Németbánya Kaposmérő

41 Ctenoph.ag.eurous H y s t r i c h o p s . t a l p . o r .

1 Apodemus f l a v i c o l l . 1 Sarkadremete

42 Ctenoph.ag.eurous H y s t r i c h o p s . t a l p . o r . Nosopsyllus f a s c .

1 Apodemus s y l v a t i c u s 1 Sarkadremete

43 Ctenoph.ag.eurous Nosopsyllus f a s c .

1 Apodemus f l a v i c o l l . 1 Sarkadremete

44 Ctenoph.ag.eurous Rhadinops.pentac.

1 Apodemus f l a v i c o l l . 1 Sarkadremete

45 Ctenoph.ag.kleins. C t e n o p h . a s s i m i l i s Megabothris t u r b i d .

1 Apodemus s y l v a t i c u s 1 Tákos

46 Ctenoph.ag.peusian. Ctenoph.congener *

2 Apodemus f l a v i c o l l . Clethrionomys g l a r .

1 1

Kisnána Kisnána

47 Ctenoph.ag.peusian. Ct e n o p h . o r i e n t a l ! s

1 Apodemus f l a v i c o l l . 1 Deszkáspuszta

48 Ctenoph.ag.peusian Ctenoph.solutus

5 Apodemus f l a v i c o l l . 5 Deszkáspuszta Kisnána

49 C te n o ph. ag. peus i a n . H y s t r i c h o p s . t a l p . o r .

. 1 Apodemus f l a v i c o l l . 1 Diósjenő

50 Ctenoph.ag.peus. X eurous C t e n o p h . a s s i m i l i s

1 Apodemus s y l v a t i c u s 1 Ócsa

1 2 3 4 5 6

51 Ctenc-ph.ag .peup. x eurous x k l e i n s . C t e n o p h . a s s i m i l i s

1 M i c r o t u s a r v a l i s 1 Ohat

52 Ctenoph.ag.ssp. Ctenoph.congener

1 Clethrionomys g l a r . 1 Deszkáspuszta

53 Ctenoph.ag.ssp. Ctenoph.orientális

1 Talpa europaea 1 Hajdubagos

54 Ctenoph.. as s i m i l i s Megabothris t u r b i d .

1 Clethrionomys g l a r . 1 K i s b a l a t o n

55 C t e n o p h . a s s i m i l i s Nosopsyllus f a s c .

5 Apodemus f l a v i c o l l . Apodemus s y l v a t i c u s M i c r o t u s a r v a l i s Talpa europaea

1 1 2

1

Bugac Tahitótfalu Pácánkert Békéscsaba Apaj

56 C t e n o p h . a s s i m i l i s Palaeops.sor.ros.

1 Neomys f o d i e n s 1 K i s b a l a t o n

57 Ctenoph.bisoctodent. Palaeops.kohauti P a l a e o p s . s i m i l i s

1 Talpa europaea 1 Deszkáspuszta

58 Ctenoph,bisoctodent. i P a l a e o p s . s i m i l i s

2 Talpa europaea 2 Deszkáspuszta

59 Ctenoph.caucasicus Ctehoph.congener Ctenoph,orientális

1 Spalax leucodon 1 Hajdubagos

60 Ctenoph.caucasius Ctenoph.orientális

1 Spalax: leucodon 4 Ha j dubago s

3 2 3 4 5 6 61 C t enoph. c ongener

Dorâtops.d.da syen. 1 Sorex araneus 1 Németbánya

62 Ot eno ph. c ongener Peromyscops.fallax

2 Clethrionomys g l a r . M i c r o t u s a r v a l i s

1 1

Németbánya Németbánya

63 Ctenoph. s o l u t u s R h a d i n o p s . s t r o u h a l i

1 Apodemus f l a v i c o l l . 1 Deszkáspuszta

64 Dorâtops.d.dasycn. H y s t r i c h o p s . t a l p . o r . Palaeops.sor.ros.

1 Neomys.anomal.mill. 1 Németbánya

65 Doratops.d.dasycn. Monopsyllus s c i u r .

1 Apodemus f l a v i c o l l . 1 Bakonynána

66 Doratops.d.dasycn. P a l a e o p s . s i m i l i s

2 Sorex araneus Talpa europaea

1 1

Deszkáspuszta Deszkáspuszta

67 Doratops.d.dasycn. Palaeops.sor.ros.

5 Sorex araneus 5 Németbánya K i s s zépalmapus z t a Iharkút

68 H y s t r i c h o p s . t a l p . o r . Megabothris w a l k e r i

1 M i c r o t u s oecon.méh. 1 K i s b a l a t o n

69 I s c h n o p s y l l . e l o n g . I s c h n o p s y l l . i n t e r m .

1 M yotis oxygnathus 1 Héviz

70 I s c h n o p s y l l . e l o n g . Nycteridops.eusarca

1 N yctalus n o c t u l a 1 Legény-barlang

71 I s c h n o p s y l l . h e x a c t . I s c h n o p s y l l . i n t e r m .

6 Epteslcuo s e r o t i n u s Myotis oxygnathus

1 5

Velence ördöclyuk-barlang A b a l i g e t i - b a r l a n g A g g t e l e k i - b a r l a n g

1 2 3 4 5 6

72 I s c h n o p s y l l . h e x a c t • I s c h n o p s y l l . i n t e r m . Nycteridops.pentact«

1 Myotis oxygnathus 1 A b a l i g e t i - b a r l a n g

73 I s c h n o p s y l l . h e x a c t . I s c h n o p s y l l . v a r i a h .

1 Myotis oxygnathus 1 ördöglyuk-barlang

74 I s c h n o p s y l l . h e x a c t . N y c t e r i d o p s . p e n t a c t .

3 B a r h a s t e l l a harhast. Plecotus a u s t r i a c u s

2

1

ördöglyuk-barlang Kec ske-barlang Budapest

75 Monopsyllus s c i u r . T a r s o p s y l l a octod.

2 Sciurus v u l g . f u s e . 2 Répáshuta Sopron

76 Paraceras m e l i s Pulex i r r i t a n s

3 Canis familiáris Meies meles

1 2

Németbánya Gödöllő Budakeszi

The case number o f the host species ( T a b l e 5) d i s p l a y s w i t h o u t need o f e x p l a n a t i o n the ex t e n t o f host s p e c i f i c i t y of the f l e a s pecies. This r e l a t i o n s h i p i s n o t so e v i d e n t from a study o f the l i s t of species found o c c u r r i n g t o g e t h e r .

Common occurrences o f s e v e r a l f l e a species

The coexistence o f f l e a species on the same host i n d i v i d u a l s and i n the nest m a t e r i a l s i s l i s t e d i n T a b l e 3.

Most o f the common occurrences were found i n one or two i n ­stances o n l y ; those noted i n t h r e e or more cases are worthy of s p e c i a l a t t e n t i o n . Besides the number of cases,the s i g n i f i c a n c e o f these common occurrences increases i f they are found i n more than one host species or i n g e o g r a p h i c a l l y remote l o c a l i t i e s . At present, the frequency o f cases seems t o be the most s i g n i ­f i c a n t ( T a b l e 4 ) .

Table 4 - 4 . táblázat. Frequency o f Common Occurrences of Flea Species According

to the Number of Species B o l h a f a j o k együttes előfordulásának gyakorisága a talált

f a j o k száma s z e r i n t

Number o f D i f f e r e n t Common Occurrences Comprising

2 Species J 3 Species | 4 Species Különféle együttes előfordulások száma

2 f a j 3 f a j 4 f a j

28 7 5 2 3 2 2 1 1

18 3 1

Frequency o f Common Occurrences

Együttes előfordu­lások gyakorisága

1 2 3 4 5 6 X 7 X

X X

9 11

F u r t h e r e v a l u a t i o n and i l l u s t r a t i o n are de r i v e d from two oth e r Tables. T a b l e 5 shows the frequency o f occurrences (com­bined cases o f s i n g l e and common occurrences) per host o f the f l e a s p e c i e s . T a b l e 6 d i s p l a y s the frequency o f common occurrences o f the i n d i v i d u a l f l e a species.

I n t he course o f my i n v e s t i g a t i o n s the f o l l o w i n g forms o f com­mon occurrences o f f l e a species were e s t a b l i s h e d .

Flea common occurrences on mouse species

I t was most f r e q u e n t l y on mouse species t h a t coexistence o f se­v e r a l f l e a species was found. This i s probably accounted f o r n e t only because the g r e a t m a j o r i t y o f c o l l e c t e d host i n d i v i ­d uals belonged t o t h i s group but also because o f the well-known f a c t t h a t most f l e a species can be found on members o f the f a m i l y Muridae i n Hungary. The m a j o r i t y of f l e a species l i v i n g on our mice belong t o the genus Ctenophthalmus, o f which Ct. a g y r t e s (C.a.bosnicus Wagner; C.a.eurous Jordan e t R o t h s c h i l d ; C a . k l e i n s c h m i d t i a n u s Peus; C a . peusianus Rosiczky) occur i n most p o p u l a t i o n s and i n r a t h e r w e l l d e f i n e d geographical areas, as was demonstrated i n an e a r l i e r paper (SMIT and SZABÓ, 1967). However, o t h e r species o f the genus a l s o occurred, v i z . C a s s i - m i l i s a s s i m i l i s (Taschenberg), Ccongener congener R o t h s c h i l d , and. C s o l u t u s s o l u t u s Jordan e t R o t h s c h i l d . Together w i t h those species, the Hungarian members o f the genera H y s t r i c h o p s y l l a , L e p t o p s y l l a , Monopsyllus, Nosopsyllus, P a l a e o p s y l l a , Peromysco- p s y l l a , and Rhadinopsylla occurred more r a r e l y . The two most f r e q u e n t p o p u l a t i o n s (on 9 and 11 occasions) a l s o occurred on mice ( s e r i a l numbers 23 and 34 ) . Both species are r a t h e r l o c a l , b eing found i n the Bakony Mts* and i n the neighbourhood o f the v i l l a g e őriszentpéter. Another i n t e r e s t i n g f a c t i s t h a t one i n ­stance o f both common occurrences was observed i n the environs of the v i l l a g e Zajk, Comitat Zala.

« Mountains

Table 5 - 5. táblázat. Frequency o f Flea Species Found on Host Species -

Gazdaállatfajokon talált b o l h a f a j o k 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 H •H 1-1 CD ri •H CD • cd p ri Si ft CD U CD fl CQ ri 03 •H ri ri •H M •H H o ri H 03 ft P ri CD CD

i ra •H ri fl ri Ü ri -P •H ra CQ H ri o fl •H H 03 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 CD ri M ft-H P m ri -p ri o cd tUDP ri Ü H rH ri 03 pi ci O a ri cd ri rH ri •H cd CQ o P o p P cö ri •H fl fl o ri çrj H o ri H CD • • CO a O a CD ri p >i -p ri cd CD ri ft ft •H CQ P CQ H ri p ri •H • 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 ri CQ ri fl £3 ri (U • cd ra CD p CQ O H cö 3 H O ri S ra o H . . . • . • CQ m ri ri •H H cd ra H 0) ri o •H •H cd cd CQ CQ CQ CQ ri CD o •H O 0 CQ P •H ft 0Q ft •

W) U p . • • Í5JJ bo bO bû tiD CQ •H ci o ri O • r i P- 03 fi •H o ra ra i> H -H •H ri ri CD ft^ CQ ra ft-H • • • O H

CQ r-j •H fl A fl A cd CÖ ri cd cd cd fl o o o ra • o • . • . • « H ri ri H H • • • • • ra o ri ra ra CQ fl H

ft ri u CQ CQ ra ft ft ft ft • • * . . . • . ra .ri CQ CQ ra CQ m ra ra >>fl fl H H ra ra ra ri o ri ft ft ft ft >> O ft-P ft ft ft o o CD CD - r i fl fl fl fl fl A A fl A A ft o ft ft ft ft ft ft ft CQ P P >j >•> H •H ft ft ft ri CQ iH o o O o ra <D • • O O o H rH O O ft ft ft ft ft ft fti ft 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 H 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 s i ri ri ri ri ri o fl fl ft ft

CD CD CD CD 03 O a 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 -P cd cd o o P P ri ri ri ri O 03 rH ri m ü u U 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 O O rH H H ri ri H ri ci •H ri U CD CD S fii fi •H •H P -P -P P P P P p -P P P P P -P P o >Î CQ CQ m ra ra ra ra 03 03 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 1

3 3 23 5 \

Apodemus f l a v i c o l l i s 30 C 3

H 1 3 4 3 23 2 3 2 2 5 6 1 4 4 4

Apodemus s y l v a t i c u s 2 1 3 4 3 2 5 3 B a r b a s t e l l a b a r b a s t . 2 3

Canis familiáris 1 2 2 4 8 C i t e l l u s c i t e l l u s 2 4 5 4 10 C l e t h r i o n o m y s g l a r . 40 5 i 6 21 3 1 2 9 4 5 4 Dryomis n i t e d u l a

5 E p t e s i c u s s e r o t i n u s 5

< 5 Erinaceus europaeus 5 F e l i s c a t u s 2 4 F e l i s s i l v e s t r i s 4 G l i s g l i s 5 Lepus europaeus Martes f o i n a 2 Meles meles 4 5 5 Micromys m i n u t . p r a t . 2 1 M i c r o t u s a g r e s t i s 4 M i c r o t u s a r v a l i s 3 4 3 1 M i c r o t u s oecon.méhelyi 4 2 2 M i n i o p t e r u s s c h r e i b . Mus musculus s p i c i l . j { \ 3 Muscardinus a v e l l a n . \ 2 Mustela n i v a l i s Mustela p u t o r . f u r o 1 Mustela putor.hung. •

M y o t i s b r a n d t i i M y o t i s dasycneme \

M y o t i s m y o t i s 4 2 3 M y o t i s n a t t e r e r i 2 M y o t i s oxygnathus 4 3 45 4 2 Neomys anomalus m i l l . 2 3 1 6 Neomys f o d i e n s 2 1 2 Myctalus n o c t u l a 6 4 Ondatra z i b e t h i c u s P i p i s t r e l l u s p i p i s t r . Pitymys subterraneus 6 < 4 2 P l e c o t u s a u s t r i a c u s 7 5 Rhinolophus ferrume. 2 S c i u r u s v u l g a r i s f u s e . \ 4 3 Sorex araneus 1 16 \ 47 Sorex minutus Spalax leucodon 1 7 6 Talpa europaea i i 3 3 1 1 4 14 4 Vulpes v u l p e s crue. \ 43 4 1 1 3 48

Table 6 - 6 . táblázat. Frequency of Common Occurrences of F l e a Species -Bolhafajok együttes előfordulásának gyakorisága

0 g

• CQ . p ri CQ CÛ • •H p ri P •H PI CQ •H ri . ri X P! CQ CQ ci o Cû ri • •H P ri 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 ri CÖ

u CD CQ ri 0 •H ri ri •H 0 fl •H •d o ri H 0 ftP 0 0 •H fl ri o ci p •H CQ CQ H ri o fl O 0 Ü O • H H CQ CQ ri o •H CQ fl . H o •H 0 ri CÛ ri H CÖ p a fl ri H •H CQ o o P •H O CÛ H cd CÖ ri P

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 od

H

p •H fl O ri 0 ri H O ri r—t 0 CQ a o ri 0 ri p >> P ri  0 •H p CQ fl ri P ri •-H • H fl ri ri ri ri O od H

-p H O •H cd •H a 0 fl 0 fl fl A PH CQ •H o o CD ri CQ • o X p ri CQ ri fl a ri 0 • ri CÛ 0 P

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 \>> H fl tu) P . . . bO EVD bQ CQ •H cd o ri O -d fl 0 fl •H > H •H ri ri 0 fl fl CQ CQ fl •H • • • H CO H •H . A fl A fl CÖ ci ri cd ri cd ri ri fl o o o CQ • O « . • » ri ri H H • . • • • CÛ O ri CQ CQ CQ H fl P ri CQ CQ Pi fl P H P H

cd • • . . • CQ fl cû Cû CQ CQ ri fl H H fl TT) CQ CQ CQ cö o ri fl P H 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 P H P H fl p P •H •H fl fl fl ri CÛ •H O O O Cû CD • • O O H H O O P H fl fl fl fl PH P H fl fl fl fl fl fl O •H O O O O o o CQ CQ ri ri O O O 0 b

X 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 P H 0 0 0 0 0 O a X H •H •H o fl CÖ p 0 0 0 0 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 ri ri cd ri c 0 fl T J fl CÛ

o ri u cd ri P P 0 0 0 0 0 0 0 0 0 0 0 0 a> o 0 ri CQ o o o o ri CQ o o fl H H ri ri H ri CÖ ri ÍH 0 CD fl fl •H •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 0 ri fl fl fl CÖ

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Chaetops.globiceps Chaetops t r i c h o s a

\ i 2 4 3 Chaetops.globiceps Chaetops t r i c h o s a 1 3 <0 Ci t e l l o p h . m a r t i n o i 3 Citelloph.simplex \ i « Ctenoceph.canis 1 i 2 Ctenoceph.felis Ctenoph.ag.bosnicus 5 20 0 4 V 10 3 2 i 6 5 Ctenoph.ag.eurous 1 2 3 i Ctenoph.ag.kleins. \ Ctenoph.ag.peusian. 2 1 4 1 Ctenoph.ag.p.x e. Ctenoph.ag.p.x k.x e. Ctenoph.ag. ssp.' i 1 Ctenoph.assimilis 5 \ i 2 5 2 Ctenoph.bisoctodent. 3 Ctenoph.caucasicus \ 6 Ctenoph.congener 20 2 \ i 3 2 6 i Ctenoph.orientális 3 4 \ \ 6 \ Ctenoph.solutus 8 4 \ Doratops.dasycnema 4 3 \ 2 6 Hy s t r i c h o p s . t a l p . o r . 4 2 l i i <

Ischnops.elongatus \

Ischnops.hexactenus 7 \ 3 Ischnops.intermed. 7 1 I s c h n o p s . v a r i a b i l i s Megabothris turbid. <Û i 2 Megabothris walkeri 3 \ Monopsyllus s c i u r . \ \ 2 2 i i h 3 Nosopsyllus f a s c i a t . i 1 \ 3 5 \ i Nycterid.eusarca Nycterid.pentactena 3 \ Palaeops.kohauti 1* \ P a l a e o p s . s i m i l i s 3 l \ Palaeops.sor.ros. 6 2 6 h \ Paraceras melis 4 3 8 Peromyscops.fallax 5 6 i \ i Pulex i r r i t a n s \ 9 2 1 8 Rhadinops.isacantha \ Rhadinops.pentacant. Rhadinops.strouhali T a r s o p s y l l a octod. 3

Plea common occurrences on shrew species

The most f r e q u e n t f l e a o c c u r r i n g on shrew species i n Hungary i s Pa l a e o p s y l l a s o r i c i s r o s i c z k y i Smit, f o l l o w e d hy D o r a t o p s y l l a das.dasyenema ( R o t h s c h i l d ) . E i t h e r one o f those species,or both appear i n most p o p u l a t i o n s , f r e q u e n t l y a s s o c i a t e d w i t h Ctenoph- thalmus a g y r t e s bosnicus Wagner, or more r a r e l y w i t h C t . a s s i m i ­ l i s a s s i m i l i s (Taschenberg), Ct. congener congener R o t h s c h i l d , H y s t r i c h o p s y l l a t a l p a e orientális Smit, Megabothris w a l k e r i ( R o t h s c h i l d ) , and P a l a e o p s y l l a s i m i l i s s i m i l i s Dampf.

Plea common occurrences on the f o x and the badger

A s p e c i f i c a l f o x f l e a i s Chaetopsylla globiceps (Taschenberg), f l e a s s p e c i f i c f o r the badger are Chaetopsylla t r i c h o s a t r i c h o -sa Kohaut and Paraceras m e l i s melis ( W a l k e r ) . Besides those t h r e e species, Pulex i r r i t a n s Linné i s als o f r e q u e n t on both host species. Since the f o x and the badger f r e q u e n t l y a l t e r n a t e t h e i r l a i r s t h e i r f l e a s can i n t e r m i n g l e e a s i l y , a l t h o u g h the s p e c i f i c f l e a s mentioned above u s u a l l y predominate on t h e i r own host species. I n the e i g h t cases of common occurrence found on the badger, only the f o u r f l e a species l i s t e d above occurred i n v a r i o u s combinations whereas i n the f o u r t e e n p o p u l a t i o n s of f l e a s found on the f o x , the species Monopsyllus s c i u r o r u a (Schrank) and Archaeopsylla e r i n a c e i e r i n a c e i (Bouché) a l s o ap­peared i n one case each. I should l i k e t o remark t h a t the f l e a species of the f o x and the badger are als o f r e q u e n t l y found on breeds o f dog used i n hunting badger-dog, f o x - t e r r i e r , and t h e i r mongrels.

Plea common occurrences on the mole-rat and the gopher

Owing t o the much decreased numbers o f Spalax leucodon Kordmann i n Hungary, the above two host species occur t o g e t h e r i n only a few l o c a l i t i e s . However, I t h i n k t h a t the common occurrences o f

t h e i r f l e a a should he discussed t o g e t h e r , since i n an i n v e s t i ­g a t i o n of a l o c a l i t y (Hajdubagos), where the two host species occur t o g e t h e r the f l e a s o f the gopher can f r e q u e n t l y he found on the mole-rat a l s o , e i t h e r alone or t o g e t h e r w i t h o t h e r f l e a species. Ctenophthalmus caucasicus (Taschenberg) i s c l e a r l y host s p e c i f i c to the m o l e - r a t ; i t occurred i n a l l observed cases t o g e t h e r w i t h the f l e a species o f the gopher, and i n one case w i t h Ctenophthalmus congener congener R o t h s c h i l d . The f l e a s of t h e gopher are C i t e l l o p h i l u s m a r t i o n i (Wagner e t I o f f ) , C. simplex (Wagner), and Ctenophthalmus orientális (Wagner). I n the e i g h t cases o f common occurrences found on the gopher one or o t h e r o f these species have, w i t h one e x c e p t i o n , always been present.Besides the f o r e g o i n g species 3 o t h e r f l e a s were found, each on one occasion. They were Ceratophyllus t r i b u l i s (Jordan) presumably t r a n s f e r r e d t o the gopher from a b i r d n e s t i n g on the ground. Ctenophthalmus a s s i m i l i s a s s i m i l i s (Taschenberg), and Nosopsyllus f a s c i a t u s (Bosc).Two v a r i e t i e s of common occurrence ( s e r i a l numbers 17 and 18) were found i n three cases each. I t deserves s p e c i a l mention t h a t whereas the f l e a s o f the gopher very f r e q u e n t l y occur on the m o l e - r a t , the c h a r a c t e r i s t i c f l e a o f the mole-rat has never been observed on a gopher occupying the same l o c a l i t y and h a b i t a t .

Plea common occurrences on the mole

I found s i x k i n d s o f f l e a p o p u l a t i o n s on the mole w i t h the host s p e c i f i c P a l a e o p s y l l a s i m i l i s s i m i l i 3 Dampf appearing i n most of them. I n t h r e e cases Ctenophthalmus b i s o c t o d e n t a t u s b i s o c t o - dentatus K o l e n a t i was a l s o observed - p r e v i o u s l y c o l l e c t e d only from the mole i n Hungary - so t h a t i t can be regarded w i t h some j u s t i f i c a t i o n aa host s p e c i f i c . Besides the two species men­t i o n e d above, Ctenophthalmus a g y r t e s bosnicus Wagner, C t . a s s i ­ m i l i s a s s i m i l i s (Taschenberg), Ct.orientális (Wagner), Dorato- p s y l l a dasyonema dasycnema ( R o t h s c h i l d ) , Nosopsyllus f a s c i a t u s (Bosc), P a l a e o p s y l l a k o h a u t i Dampf, a n d , P . s o r i c i s r o s i c z k y i Smit were a l s o c o l l e c t e d i n one case each. U n f o r t u n a t e l y , I have not

y e t had occasion to c o l l e c t the m a t e r i a l of a mole burrow, though the r i c h f l e a fauna o f i t s nest has a t t r a c t e d the a t t e n ­t i o n o f s e v e r a l research workers (OUDEMANS, 1913; WAGNER, 1936). Recently ROSICKY (1937), SMIT (1962), and JTJRIK (1968) from an i n v e s t i g a t i o n o f considerable m a t e r i a l e s t a b l i s h e d the presence of many f l e a species,both on the animal and i n i t s nest.Regret-a b l y , none of these a u t h o r s , w i t h the e x c e p t i o n o f ROSICKY have publ i s h e d i n f o r m a t i o n on the f l e a p o p u l a t i o n s o f the se­v e r a l hosts and t h e i r n e s t s but have merely discussed the oc­currence of the f l e a species observed.

Plea common occurrences on the bat

I t might be assumed t h a t s e v e r a l types of f l e a p o p u l a t i o n would occur on bats because c e r t a i n bat species aggregate i n masses i n b o t h t h e i r w i n t e r and summer q u a r t e r s . T r u e , i n most c o l o n i e s only members of a given species are c l o s e l y a s s o c i a t e d w i t h each o t h e r but there would s t i l l be more p o s s i b i l i t i e s f o r t r a n s m i s s i o n o f d i v e r s f l e a species than i n the case o f o t h e r mammalian groups. However, I found only s i x k i n d s of common oc­currences o f f l e a s , among which only two occurred i n t h r e e and s i x cases, r e s p e c t i v e l y ( s e r i a l numbers 71 and 7 4 ) , w h i l e the f o u r o t h e r compositions were observed i n only one case each. The most f r e q u e n t species of the cases of coexistence i s I s c h ­ n o p s y l l u s hexactenus ( K o l e n a t i ) , which., occurred i n f o u r v a r i a ­t i o n s i n eleven cases. The occurrence o f I . i n t e r m e d i u s (Roth­s c h i l d ) w i t h other species i s f a i r l y common having been found i n t h r e e v a r i a t i o n s on e i g h t occasions.

I n a d d i t i o n there are o t h e r , somewhat o c c a s i o n a l , comron occur­rences o f smaller s p e c i f i c numbers and r a t h e r r a r e appearence. Thus, besides Monopsyllus sciurorum sciurorum (Schrank), the s p e c i f i c f l e a o f the s q u i r r e l ( S c i u r u s v u l g a r i s f u s c o a t e r Altum) and i t s n e s t , one may o c c a s i o n a l l y f i n d i n i t s company Tarso- p s y l l a octodecimdentata octodecimdentata ( K o l e n a t i ) , a species r a t h e r r a r e i n Hungary. The v a r i o u s dormouse species show a de-

f i n i t e preference f o r a r t i f i c i a l n e s t i n g boxes,where the s q u i r ­r e l f l e a s (M.sciurorum) l i v i n g on the dormice may be found t o ­g e t h e r w i t h the f l e a s o f a v i a n species which had p r e v i o u s l y nested t h e r e . This a s s o c i a t i o n w i l l n o t be considered as i t i s the r e s u l t o f human i n t e r f e r e n c e and not n a t u r a l . Nor do I wish t o discuss the v a r i o u s common occurrences o f f l e a species t o be found on our domestic animals.

Discussion

From the f o r e g o i n g data o f observed c o e x i s t i n g f l e a species, an attempt w i l l be made t o answer the problems r a i s e d i n the i n t r o d u c t i o n .

1 . The f o l l o w i n g systematic r e l a t i o n s h i p can be e s t a b l i s h e d between f l e a species o c c u r r i n g t o g e t h e r on a give n host or i n i t s n e s t s : the members o f the p o p u l a t i o n s belong t o

v a r i o u s f a m i l i e s and genera i n . . . . 42.1 fo o f cases; the same f a m i l y i n 18.4 ̂ of cases; the same f a m i l y and genus i n 39.5 i° o f cases.

Taking i n t o c o n s i d e r a t i o n t h a t h a l f the species of Hungarian Siphonaptera belongs t o the f a m i l y H y s t r i c h o p s y l l i d a e and n e a r l y o n e - t h i r d t o the genus Ctenophthalmus - whose members are the most commonly o c c u r r i n g f l e a s - one can s a f e l y conclude t h a t the apparent connections between species o c c u r r i n g t o g e t h e r stem more from the composition o f the fauna than from any i n ­c l i n a t i o n o f c l o s e l y r e l a t e d species t o form c e r t a i n types of a s s o c i a t i o n . There i s as y e t no j u s t i f i c a t i o n f o r r e l a t i n g the s p e c i f i c composition o f mixed p o p u l a t i o n s t o any nearer i n t e r ­r e l a t i o n s h i p .

2. I sought i n v a i n t o e x p l a i n the i n c l i n a t i o n of the species t o l i v e t o g e t h e r . Nor i s there any reason t o suppose an i n t e r ­dependence o f the' species: they d e r i v e no advantage or d i s a d -

vantage from the presence or absence o f o t h e r f l e a species. The food requirements o f the l a r v a e and f u l l y developed f l e a s h a r d l y d i f f e r . And i t would be unreasonable t o suppose t h a t some s m a l l e r - s i z e d f l e a would have l e s s food because of the presence o f the l a r g e - s i z e d H y s t r i c h o p s y l l a t a l p a e orientális. There i s no i n f o r m a t i o n to suggest t h a t any one o f the species could i n t e r f e r e w i t h the f e e d i n g or r e p r o d u c t i o n o f o t h e r s . (One might presume disadvantages a r i s i n g from the presence of c e r t a i n species t o c o e x i s t i n g species i n only one case: v i z , i f some fav o u r a b l e circumstances enabled one species t o o v e r p r o l i -f e r a t e i n a h a b i t a t / n e s t , burrow, l i t t e r / , though even then the members o f the m i n o r i t y species might o b t a i n l e s s food only i n p r o p o r t i o n t o the number o f i n d i v i d u a l s o f the species.)

3. A comparison of T a b l e s 5 and 6 f u l l y i d e n t i f i e s those species which seldom or never a s s o c i a t e w i t h o t h e r spe­c i e s . The f o l l o w i n g were never found i n s p e c i f i c a s s o c i a t i o n s : Chaetopsylla r o t h s c h i l d i , I s c h n o p s y l l u s octactenus, I . simplex m y s t i c u s , I . s. simplex, L e p t o p s y l l a segnis, R h i n o l o p h o p s y l l a u. u n i p e c t i n a t a . .The species Archaeopsylla e. e r i n a c e i , I s c h n o ­ p s y l l u s v a r i a b i l i s , N y c t e r i d o p s y l l a eusarca, Rhadinopsylla pen- t a c a n t h a , and R h . s t r o u h a l i occurred t o g e t h e r w i t h o t h e r species i n one case onl y . The m a j o r i t y o f these species, having noTten­dency t o appear i n the company of o t h e r s , have been c o l l e c t e d a few times o n l y ; most of them are probably t r u l y r a r e , but there may be some which appear t o be r a r e only because t h e i r hosts a l s o occur i n f r e q u e n t l y i n Hungary, or because o f some aspect of c o l l e c t i o n . I t should be emphasized again t h a t the number o f r a r e f l e a species i s c o n s i d e r a b l y l e s s than we t h i n k and we are r a t h e r i n c l i n e d t o regard those which are seldom captured as r a r e whereas the cause o f t h i s phenomenon a c t u a l l y l i e s i n f a u l t s o f the c o l l e c t i n g apparatus or techniques. I f the spe­c i e s mentioned above are captured s e v e r a l times i n f u t u r e c o l ­l e c t i o n s , they might a l s o be found t o c o e x i s t w i t h o t h e r spe­c i e s . However, i n the case o f species which are f r e q u e n t and s t i l l f a i l t o occur i n the company of o t h e r s , the p o s s i b i l i t y remains t h a t , f o r some unknown reason they have no „inclination"

t o p a r t i c i p a t e i n s p e c i f i c a s s o c i a t i o n s . Such are I . o c t a c t e n u s , h.segnis, A . e r i n a c e i , and N.eusarca. I have c o l l e c t e d the hosts (Erinaceus europaeus roumanicus Barr.-Ham., and Mus musculus s p i c i l e g u s Perényi) o f two o f these species ( A . e r i n a c e i and L. segnis) i n moderate numbers, but I have never found them t o occur t o g e t h e r w i t h o t h e r f l e a species, even though b o t h hosts have ample o p p o r t u n i t y , b o t h i n t h e i r nests and sphere o f a c t i ­v i t y , t o acq u i r e a number o f o t h e r p a r a s i t e s . One might t h e r e ­f o r e i n f e r w i t h some j u s t i f i c a t i o n t h a t these two f l e a species are f o r some reason l e s s i n c l i n e d t o c o e x i s t w i t h o t h e r s or t h a t they shun the presence o f the l a t t e r , o r , indeed, v i c e versa. One should a l s o r e c a l l t h a t Ct.caucasicus, the f l e a o f the m o l e - r a t , has never been found on gophers though a l l f l e a species o f t h a t l a t t e r animal f r e q u e n t l y occur on the m o l e - r a t . The problem of host s p e c i f i c i t y i n f l e a s , perhaps depending t o some e x t e n t on the type o f food ( b l o o d ) may be c l a r i f i e d by l a ­b o r a t o r y experiments.

4. The l a 3 t and most d i f f i c u l t q u e s t i o n i n v o l v e s the a c t u a l e c o l o g i c a l , or some o t h e r , cause of the coexistence of f l e a species on a h o s t , o r the p o s s i b i l i t y of i t s chance (spontaneous) n a t u r e . The s p e c i f i c f l e a s o f the hosts are known, and we a l s o have i n f o r m a t i o n on animals which could serve as hosts f o r the gi v e n f l e a species. Those f l e a s considered most s p e c i f i c appear i n c i d e n t a l l y though seldom, on o t h e r than t h e i r „true" h o s t s , b u t these are merely o c c a s i o n a l phenomena. I f , however, the host changes i t s haunt (mainly i t s s i t e of r e p r o d u c t i o n ) f o r any l e n g t h of time,and i f t h i s be the h a b i t a t o f hosts support­i n g o t h e r f l e a species, then those appearing l a t e r on the im­migrant host can h a r d l y be d e f i n e d as merely o c c a s i o n a l para­s i t e s . (The number o f i n d i v i d u a l s o f a f l e a species found on hosts and i n t h e i r n e s t s i n d i c a t e s the oc c a s i o n a l or s p e c i f i c n a t u r e o f the f l e a species.) I t was many times my experience t h a t the f l e a fauna o f a host found i n an environment d i f f e r e n t from i t s usual h a b i t a t ( n e s t i n g s i t e ) i s n o t composed of the species t o be expected.

I should l i k e t o put forward a most c h a r a c t e r i s t i c example (not contained i n the Tables, because the case was s t u d i e d a f t e r the p e r i o d under i n v e s t i g a t i o n ) . I n October, 1968, i n the southern p a r t of the country (Madaras) an ermine (Mustela erminea a e s t i v a K e r r ) made i t s subterranean q u a r t e r s i n a f i e l d much i n f e s t e d by gophers. We succeeded i n s h o o t i n g the a n i m a l . Fourteen of the f i f t e e n f l e a specimens found on i t proved t o belong to the s p e c i f i c f l e a s of the gopher ( C i t e l l o p h i l u s m a r t i n o i and Cteno­phthalmus orientális). This comparatively l a r g e number of i n d i ­v i d u a l s a l s o i n d i c a t e s t h a t the f l e a species mentioned do not a t t a c h themselves so much t o t h e i r h o s t s as t o the h a b i t a t of the n e s t i n g s i t e .

DUDICH (1939) has already remarked t h a t one u s u a l l y encounters d i f f i c u l t i e s when a t t e m p t i n g t o e s t a b l i s h the b i o t o p e s o f spe­c i e s . This statement i s e s p e c i a l l y t r u e of p a r a s i t e s and thus of f l e a s . What then, are the biotopes of f l e a s ? Environment i n the s t r i c t e r or wider sense of the term? There are species which occur on animals l i v i n g i n pastures and a g r i c u l t u r a l areas o f the p l a i n s , and ethers on f o r e s t mammals. Some are more f r e q u e n t i n a r i d , o t hers i n damper h a b i t a t s . I f a i l e d to f i n d any.demonstrable connection between the p l a n t cover, the f o r e s t stand,and the occurrence o f f l e a species; - i f the hosts are p r e s e n t , and t h e i r burrows and n e s t s s u i t a b l e f o r i n v a s i o n by and r e p r o d u c t i o n o f f l e a s , we might expect the appearance of c e r t a i n species. Hence the biotope o f f l e a species i s not the w i d e r but the more r e s t r i c t e d h a b i t a t ; the s m a l l area where they reproduce, l a y t h e i r eggs, the l a r v a e f i n d t h e i r food, and the pupae t r a n s f o r m i n t o imagos. I n the search f o r the b i o t o p e of the f l e a s we have thus a r r i v e d a t the l a i r s of t h e i r h o s t s , be they subterranean burrows, h o l e s , or nests c o n s t r u c t e d on the ground, on clumps of grass, shrubs, or t r e e s . (Tne b i z a r r e idea t h a t the biotope o f the f l e a species i s the host i t s e l f should be r e j e c t e d a t once; t h i s assumption may be j u s t i f i e d only i n some cases of endoparasitism.) The b i o t o p e s mentioned above can be d e l i n e a t e d r a t h e r s a t i s f a c t o r i l y and a l s o assigned t o the b i o c o e n o l o g i c a l h o r i z o n s o f the temperate regions es-

ta"blished by BALOGH (1953). The communication of the s e v e r a l species u s u a l l y occurs between n e i g h b o u r i n g h o r i z o n s o n l y . Rare exceptions n a t u r a l l y occur, e.g. a s q u i r r e l f l e a (M.s.sciuro­ rum) found on one occasion on a f o x ; i t might have a r r i v e d on i t s unusual host e i t h e r by having f a l l e n from the nest or i n the course o f the f o o d - c h a i n , or e v e n t u a l l y from the ground, as, f o r i n s t a n c e , was the s q u i r r e l f l e a c o l l e c t e d by s o i l sur­face e x t r a c t i o n methods.

On the basis o f my experiences gained i n the course o f c o l l e c ­t i o n s and i n v e s t i g a t i o n s , I venture t o s t a t e t h a t a g r e a t majo­ r i t y o f f l e a species appear t o be more bound t o c e r t a i n types o f n e s t s and t h e i r m i c r o c l i m a t e than to the hosts themselves. I s h a l l t r y t o prove t h i s a s s e r t i o n by some examples. The f o x and the badger i n h a b i t the same bio t o p e where i n most of our mouse species a l s o occur, but I have never found a f l e a species para­s i t i z i n g mice on e i t h e r the f o x or the badger. S i m i l a r l y , the c h a r a c t e r i s t i c f l e a s o f these l a t t e r two hosts have never oc­c u r r e d on mice,although t h e r e are innumerable o p p o r t u n i t i e s f o r the exchange or t r a n s m i s s i o n o f f l e a s l i v i n g on these h o s t s . That i s n o t to say, however, t h a t one or two mouse f l e a s w i l l n o t be found on f o x or badger, or v i c e v e r s a , though t h i s can only be an o c c a s i o n a l phenomenon. (Even i n the l i t e r a t u r e t h e r e are only a few records o f such an occurrence.) I n my o p i n i o n , l a c k o f such i n t e r c h a n g e o f f l e a species i s not due simply t o d i f f e r e n c e s i n the h o s t s , but t o the f a c t t h a t nests o f mice have a completely d i f f e r e n t m i c r o c l i m a t e from those of the deep subterranean burrows o f the f o x and the badger. I n c i d e n t a l l y , PETJS (1953) has a l r e a d y p o i n t e d out the importance of the c o r ­r e l a t i o n between the occurrence o f c e r t a i n f l e a species and the environmental f a c t o r s , so my experiences would seem to c o r r o ­borate h i s statements.

A c e r t a i n k i n d of c o r r e l a t i o n i n the s y s t e m a t i c a l r e l a t i o n s h i p o f the hosts and t h e i r f l e a s (and a l s o t h e i r o t h e r p a r a s i t e s ) appears t o be f r e q u e n t l y demonstrable.This statement a p p a r e n t l y c o n t r a d i c t s my p r e v i o u s a s s e r t i o n , because i t might be supposed

t h a t the f l e a s o f the hosts mentioned above do n o t mingle on account o f the d i f f e r e n t s y s t e m a t i c a l c a t e g o r i e s of the h o s t species. But then why do the f l e a s o f the badger occur on the f o x , and v i c e versa? - the two host species are assigned t o two d i f f e r e n t mammalian f a m i l i e s . The mice stand a t l e a s t as f a r removed from the shrews as from the f o x and the badger, but the f l e a s c h a r a c t e r i z i n g the one s t i l l occur on the o t h e r . The ob­j e c t i o n t h a t the f l e a species of the mice and the shrews s t i l l d i f f e r may t h e r e f o r e be a p p a r e n t l y j u s t i f i e d - these hosts l i v e i n a common h a b i t a t and i n subterranean burrows of n o t too d i f ­f e r i n g t y p e s . I n the course of making my c o l l e c t i o n s , I have i n f a c t found mouse and shrew species w i t h i n the same h a b i t a t i n many places but the shrews i n v a r i a b l y occurred i n the v i c i n i t y of the m o i s t e r areas ( s m a l l e r bodies o f s t a n d i n g waters, shores of b r o o k s ) , so the m i c r o c l i m a t e o f t h e i r nests a l s o i s probably d i f f e r e n t from t h a t o f mice.

Evidence s u p p o r t i n g my t h e s i s can be found i n the mouse f a m i l y . The f l e a G t . a . a s s i m i l i s i s more or l e s s s p e c i f i c t o the f i e l d v o l e ( M i c r o t u s a . a r v a l i s P a l l a s ) . I t i s known t h a t the f i e l d v o l e i n h a b i t s r a t h e r dry meadows, pa s t u r e s , . and a g r i c u l t u r a l f i e l d s , and so the environment of i t s nests i s d r i e r than t h a t of mice (Apodemus, Clethrionomys, Pitymys, Micromys). Now I have c o l l e c t e d Ct«assimilis r a t h e r f r e q u e n t l y on mouse species i f t h e i r h a b i t a t s were adjacent t o dry areas, and even i n cases when I found no f i e l d v o l e s i n the v i c i n i t y . I t might be assumed i n t h i s case - and w i t h j u s t i f i c a t i o n - t h a t C t . a s s i m i l i s i s bound n o t so much t o M . a r v a l i s as t o the m i c r o c l i m a t e of i t s nests and t h e i r s t r i c t e r environment. Though I have discussed here only occurrences jof f l e a s p a r a s i t i z i n g mammals, I should l i k e t o s u b s t a n t i a t e my theory by an example drawn from a v i a n f l e a s . A g r e a t number o f h o s t s , n o t too f a r removed from one another s y s t e m a t i c a l l y , o f a v i a n f l e a species are known i n Hun­gary. There i s one e x c e p t i o n , namely C e r a t o p h y l l u s s t y x s t y x R o t h s c h i l d , demonstrated only from the sandmartin ( R i p a r i a r i -p a r i a L . ) ; nor d i d I f i n d any o t h e r f l e a species on t h i s h o s t . Even PEUS's (1968) abundant research m a t e r i a l f a i l e d t o produce

other f l e a s from t h i s host, and C.styx was found i n only a single case on the stonechat (Oenanthe oenanthe L . ) , which oc­c a s i o n a l l y nests i n s i t e s s i m i l a r to the subterranean nesting c a v i t i e s of the sand-martin. Thus we have the s p e c i f i c f l e a of a nest type which d i f f e r s fundamentally from the nests of other species. I t would be f u r t h e r proof of my assumption i f C.s.styx were found i n the s i m i l a r l y constructed nest of the bee-eater (Merops a p i a s t e r L . ) . These examples s u f f i c e to prove the merit of t h i s type of i n v e s t i g a t i o n s and to draw attention to my hy­pothesis .

F i n a l l y , I should l i k e to add some remarks concerning my work and future research. Though numerical data r e f e r r i n g to i n d i v i ­duals of coexisting species are a v a i l a b l e , I have discussed only cases of s p e c i f i c a s s o c i a t i o n s . I n l a t e r communications I intend to consider, whose studying f l e a populations of a given host species, the numerical r a t i o s and sex d i s t r i b u t i o n sex of the p a r a s i t i c species. A more in t e n s i v e c o l l e c t i n g of nests i s also indicated, as w e l l as the evaluation of the occurrence of f l e a s by seasons (YYSOTSKAJA, 1967), the comparison of f l e a species found on the hosts and t h e i r nests (JURIK, 1968), and demonstration of c o r r e l a t i o n s between the development of f l e a s found on the hosts and t h e i r n ests (ROTHSCHILD, 1967). One or more of these aspects have already been examined- by the above-c i t e d authors and by others, without, however, a l l aspects being considered i n the course of a given i n v e s t i g a t i o n . Many ques­tions might be answered i f i t could be determined by labo­ratory i n v e s t i g a t i o n s how f a r the f l e a species found i n the samples show food preferences. Also, as already stated, I attach great importance to moisture, humidity, and temperature condi­tions a f f e c t i n g the s e v e r a l biotopes.The i n v e s t i g a t i o n of these l a t t e r seems to present most d i f f i c u l t i e s , since a f t e r the exposure of the nests the o r i g i n a l microclimatic conditions can no longer e x i s t . I f a t e c h n i c a l solution to t h i s problem were assured, we would be much nearer a demonstration of the nest-s p e c i f i c i t y of f l e a species.

The present paper discusses the most frequent v a r i a t i o n s of co­existence or common occurrences o f f l e a species o c c u r r i n g on mammals i n Hungary. The coexistence of f l e a s on mouse and shrew species, on the fox and the badger, on the mole-rat and the gopher, as w e l l as the mole and the bat species, appeared i n (comparatively) most d e f i n i t e and f r e q u e n t l y repeated composi­t i o n s . No systematic r e l a t i o n s h i p could be demonstrated between the f l e a species c o e x i s t i n g on the h o s t s . I t could not be proven t h a t any one of the f l e a species of the var i o u s a s s o c i a t i o n s was e s p e c i a l l y i n c l i n e d t o occur i n s p e c i f i c compositions. Some species occur very seldom or not a t a l l i n the company of other f l e a species. The most probable c o n d i t i o n s a f f e c t i n g the evolu­t i o n of coexistence, namely the environment of the nests, t h e i r p o s t i t i o n as to h o r i z o n , t h e i r s t r u c t u r e and microc l i m a t e , imply t h a t i f these f a c t o r s are favourable f o r the occurrence of cer­t a i n f l e a species present i n the area,then the p o s s i b i l i t i e s of forming s p e c i f i c a s s o c i a t i o n s are a v a i l a b l e f o r these species.

SZABÓ, I . : A magyarországi emlősök bolháinak (Siphonaptera) együttélési viszonyai

A szerző 1958-tól 1967 végéig 55 emlősfaj 2331 példányát és 42 fészkét vizsgálta meg bolhászati szempontból, melyeknek 35,4, i l l e t v e 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 j e l e n t k e z t e k . A gazdaállatokon együttesen előfordult bol h a f a j o k között nem sikerült származás­t a n i összefüggést találni; nem látszik bizonyítottnak, hogy az együttesen előfordult f a j o k közül valamelyik különösképpen h a j ­lamos lenne a társulásra; a magyar fauna b o l h a f a j a i között akad

néhány, melyek csak i g e n ritkán, vagy egyáltalán nem f o r d u l n a k elő társulásban. Végül megállapítja, hogy ha a fészkek környe­ze t e , s z i n t b e l i elhelyezkedése, struktúrája és mikroklímája kedvező a területen előforduló bolhák bizonyos f a j a i n a k , akkor e f a j o k részére fennállanak az együttes előfordulás lehetősé­g e i . E d d i g i gyűjtései és anyagának kiértékelése során s z e r z e t t t a p a s z t a l a t o k alapján a szerző ugy véli, hogy a b o l h a f a j o k jó-része inkább ragaszkodik bizonyos fészektipusokhoz i l l e t v e azok-mikrokiimájához, mint magához a gazdaállathoz.

R e f e r e n c e s

BALOGH,J.: A zoocönológia a l a p j a i . - Akadémiai Kiadó, Budapest, pp. 248. 1953.

DUDICH,E.: „Élettér", élőhely, életközösség. - Term. Tud. Közi. Pótf. 71. 49-64. 1939.

JURIK,M.: Pleas o f the mole Talpa europaea L. i n CzehoSlovakia ( A p h a n i p t e r a ) . - Acta Ent. Bohemoslov. 6_5_. 67-75. 1968,

OUDEMANS ,A. : Suctorologisch.es aus Maulwurfnestern. - T i j d s c h r . v. Entom. 5j6. 238-280. 1913-

PEUS,F.: Flöhe. - Akademische Verlagsges.Geest.u.Poriing K.-G., .L e i p z i g , pp. 43. 1953.

PEUS,F.: Zur Kenntnis der Flöhe Deutschlands I I . F a u n i s t i k und Ökologie der Vogelflöhe ( I n s e c t a , Siphonaptera). - Zool. Jb. Syst. 95. 571-633. 1968.

ROSICKY,B.: Aphaniptera zimnich h n i z d k r t k a obecného (Talpa eu­ropaea L.) V ruznych b i o t o p e c h . - Öeskoslov. P a r a s i t . 4. 275-290. 1957.

ROTHSCHILD,M.: The Rabbit Flea and Hormones. - Penguin Sci.Surv. B i o l . 189-199: 1967.

SHIT,F.: Siphonaptera c o l l e c t e d from moles and t h e i r nests a t w i l p , Netherlands, by Jhr.W.C. van Heurn. - Tijdschr.v.Entom. 105. 29-44. 1962.

SMIT,P. - SZABÓ,I.: The d i s t r i b u t i o n of Subspecies of Ctenoph­thalmus agyrtes i n Hungary (Siphonaptera:Hystrichopsyllidae). - Ann. Hist.-Nat. Mus. Nat. Hung. 59. 345-351. 1967-

VYSOTSKAJA,S.: Biocenotitcheskie otnoseni a mezdu ektoparasitami grüsunov i obtateliami i h gnesd. - P a r a s i t . Sbornik. 2_3. I 9 ­60. 1967.

WAGNER,J.: Über die Aphanipterenfauna der Maulwurfnester. - Eo-nowia 15. 97-101. 1936.

Received: 24.4.1969. I - SZABÓ Zoological Department of the Hun­garian Natural History Museum, Budapest, V I I I . Baross u. 13.