Parasit. Hung. 6. 1973.
Seasonal Incidence of Larval Hydatigera taeniaef or mis Infection of Microtus arvalis in Hungary Gábor
NECHAY
Central Plant Protection and Quarantine Laboratory of the Ministry of Agriculture and Food, Laboratory of Rodents, Tanakajd
"Seasonal I n c i d e n c e of Larval Hydatigera taeniaeformis I n f e c t i o n o f M i c r o t u s a r v a l i s i n Hungary" - Hechay, G. - Paras i t . Hung. 6. 117-130. 1973. On the b a s i s o f p a r a s i t o l o g i c a l i n v e s t i g a t i o n o f 3097 f i e l d mice t h e f r e q u e n c y o f H y d a t i g e r a t a e n i a e f o r m i s i n f e c t i o n was found t o be 7.2 %. Seasonal s t u d i e s r e v e a l e d a peak o f i n f e c t i o n i n t h e w i n t e r and s p r i n g , w h i l e a d r o p i n i n f e c t i o n r a t e was n o t e d d u r i n g t h e summer. I n a n i m a l s living i n l u cerne f i e l d s , a maximal r a t e o f i n f e c t i o n i s reached a l r e a d y i n t h e autumn months, w h i l e i t o n l y t a k e s p l a c e d u r i n g n e x t s p r i n g i n those l i v i n g i n wheat f i e l d s . The f r e q u e n c y o f s t r o b i l o c e r c o s i s shows c o r r e l a t i o n w i t h t h e age d i s t r i b u t i o n o f t h e i n v e s t i g a t e d p o p u l a t i o n . Taking t h e t i m e , n e c e s s a r y f o r t h e development o f c y s t s and the i n f e c t i o n r a t e o f mice i n t o c o n s i d e r a t i o n , c e r t a i n demographical c o n c l u s i o n s can be r e a c h e d . Hydatigera
t a e n i a e f o r m i s Batsch,
mi s B a t s c h ) i s common
1786 ( s y n . : T a e n i a t a e n i a e f o r -
i n small r o d e n t s .
I n t h e r o d e n t s as i n -
t e r m e d i a t e h o s t s the l a r v a l f o r m o f t h i s " S t r o b i l o c e r c u s f a s c i o l a r i s " i s t o be
found i n the l i v e r t i s s u e w i t h i n f e c t i v e
vae i n c y s t s o f 8-10 mm d i a m e t e r . the p r i n c i p a l h o s t s .
Other
lar-
The house and w i l d c a t s a r e
predators
have n o t been
found as
p r i n c i p a l h o s t s i n Hungary. The p a r a s i t e i s endemic i n Hungary and has been KOTLÁN ( 1 9 6 1 ) , EDELÉNYI
r e p o r t e d by
and SZABÓ (1963ï t h e y r e p o r t
only the
a d u l t f o r m i n w i l d c a t s ) , NEMESÉRI and HOLLÓ ( 1 9 7 2 ) , KOTLÁN and KOBULEJ ( 1 9 7 2 ) , MURAI ( 1 9 7 2 ) , MURAI & TENORA ( 1 9 7 3 ) . KOTLÁN and KOBULEJ,
NEMESÉRI and HOLLÓ have r e p o r t e d i t as a common p a r a -
s i t e o f cats
demanding v e t e r i n a r y
a t t e n t i o n and a t t h e
t i m e r e p o r t e d t h e h o s t r o l e o f house mice
same
and r a t s . MURAI and 117
M 0 3
Pxg . 1 : L o c a l i t i e s o f t h e examined m a t e r i a l 1 . ábra: A vizsgált anyag gyűjtési h e l y e i
Table 1 Number o f i n f e s t e d common v o l e s c o l l e c t e d i n v a r i o u s c o m i t a t s i n Hungary i n 1971-72 Az ország különböző területein 1971-72-ben gyűjtött pockok fertőzőttsége
1972
1971 Locality
1 . Egyházasfalu 2. Kimle 3. Páli 4. Ujkér 5. Izmény ízmény 6. Decs 7. P u s z t a e g r e s 8. P u s z t a szabolcs 9. Velence 10. Kápolnásnyék 1 sztimer I12. ' ICsanád alberti 1 3 . Csorvás 14. Kamut 1 5 . Gyöngyös tarján 16. Sárvár Sárvár 17. Kenyéri Kenyéri 18. Nőtincs Nőtincs 19. Magyar— nándor
Note:
Spring
Autumn
L
C
-
-
18/2 4/0 13/0 3/0 7/3 3/0 25/4
-
C
-
-
248/10 57/15 105/13 40/9 26/3
12/0 4/0 6/0
-
L
-
2/0
i/o
-
-
-
-
2/1
—
-
-
-
16/5
7/0 -
5/0 5/0
-•
4/1
-
-
-
26/0 9/0 1/1 3/0 305/14 47/3
-
50/4
22/7 18/3 26/6 14/3 -
L = a l f a l f a or clover Figures'before
-
-
-
_
11/0
-
-
_-
97/4
_
-
— — —
-
_
C
156/2
-
-
4/1
-
L
-
15/2
-
C
2/0
-
-
Autumn
2/0
-. -
16/3 27/1
-
L
Spring
-
-
i/o
— —
-
5/0
-
5/0
-
-
—
-
-
--
-
9/2 2/0
-
1/1 i/o
-
--
26/0
-
-
-
-
i/o 2/1 3/0
7/0
-
i/o
-
5/0
—
-
-
i/o
-
C = autumn wheat or b a r l e y
place-names r e f e r t o F i g . 1 .
A gyűjtőhelyek száma megegyezik a térkép számaival.
119
TENORA have shown t h e o c c u r r e n c e o f l a r v a e i n numerous s m a l l mammalian s p e c i e s , namely i n : 5 % o f Apodemus f l a v i c o l l i s , 2 % of A. s y l v a t i c u s , 5 % o f M i c r o t u s a r v a l i s , 0 % o f M. a g r e s t i s , 6 % o f Mus musculus s p i c i l e g u s , 1 % o f a l b i n o l a b o r a t o r y breeds Of Mus musculus and R a t t u s r a t t u s , 2 % o f C l e t h r i o n o m y s g l a r e o l u s and 1 % o f P i t y m y s s u b t e r r a n e u s . Since 1969, t h e C e n t r . P l a n t P r o t , and Quarantine Lab., Rodent L a b o r a t o r y M i n . o f Ag. Fd., has been c o n t i n u i n g p o p u l a t i o n - d y namic i n v e s t i g a t i o n s on s m a l l r o d e n t s i n a g r i c u l t u r a l f i e l d s . D u r i n g t h i s work we have a l s o s t u d i e d and e s t i m a t e d l a r v a l i n f e c t i o n s ( s t r o b i l o c e r c o s i s ) o f Hydatigera taeniaeformis i n the common v o l e , h y p o t h e s i z i n g t h a t i t has an e f f e c t on t h e v o l e population. Material and Methods
H a l f t h e examined m a t e r i a l
representing
one population"*" came
f r o m permanent c o l l e c t i o n s near t h e v i l l a g e Bük (West Com. Vas,
see P i g .
l ) i n 1969-72.
(Trapping i n
Hungary, lines
and
2 s n a p - t r a p s taken i n areas o f 1000 m i n p e r e n n i a l c u l t u r e s a l f a l f a , c l o v e r , i t a l i a n meadow-grass o r a m i x t u r e o f these t w o l a t t e r and i n c o r n - autumn wheat and b a r l e y . ) The o t h e r h a l f o f o u r m a t e r i a l came f r o m c o l l e c t i o n s made i n o t h e r p a r t s o f the c o m i t a t s (Békés, Csongrád, Győr, Fejér, Heves, Nógrád, T o l na and Vas, see F i g . 1.) i n two r e g i o n s o f each c o m i t a t and t w i c e i n the same l o c a l i t y f o r t h r e e days c o n s e c u t i v e l y i n l e guminous and c o r n c u l t u r e s d u r i n g s p r i n g ( M a r c h - A p r i l ) and autumn (October-December) i n 1971-72. I n these areas t h e method of c o l l e c t i o n was always l i n e - t r a p p i n g ( i n d e p e n d e n t o f abundance and h o l e s i t e s , w i t h 100 s n a p - t r a p s / m e t e r ) . On t h e b a s i s o f t h e home range o f t h e common v o l e s and t h e i r m i g r a t i o n f r o m one c u l t u r e t o a n o t h e r i n c r e a s e d p a r t i c u l a r l y owing t o human a c t i v i t y ( e . g . r o t a t i o n of crops, h a r v e s t i n g , p l o u g h i n g and h a r v e s t i n g o f p e r e n n i a l c u l t u r e s , p e s t c o n t r o l e t c . ) and r e s u l t i n g i n sudden changes o f t h e e n v i r o n m e n t , the specimens l i v i n g i n open areas ( a t t h e b o r d e r t e r r i t o r i e s of one a g r i c u l t u r a l c o o p e r a t i v e ) have t o be c o n s i d e r e d as one population. 120
P i g . 2:
I n f e s t a t i o n by H y d a t i g e r a t a e n i a e f o r m i s l a r v a e o f d i f f e r e n t common v o l e s a c c o r d i n g t o sex and age
2. ábra: Pockok H. t a e n i a e f o r m i s fertőzöttsége nem és k o r sze rint
= number o f observed animals l = 1458 = 1454
f J
t
number o f a n i m a l s infected with developing cysts n = 60 n | = 49 0
number o f a n i m a l s infected with f u l l y developed c y s t s n = 59 nï = 52 ç
Age groups= maximum 1 month, 1 - 2 months, 2- 4 months, 4-6 months, o l d e r t h a n 6 months 121
The
investigations
were c a r r i e d o u t
frozen animal bodies. the d e v e l o p i n g
cysts
To e s t i m a t e
on f r e s h o r
occasionally
the i n f e c t i o n s
i n the
liver
( p o i n t - l i k e on t h e s u r f a c e , embedded
small
c y s t i n t i s s u e s ) o r c y s t s c o n t a i n i n g d e v e l o p e d l a r v a e were con s i d e r e d . The l a r v a e were i d e n t i f i e d on the b a s i s o f samples sent t o Dr. É . MURAI ( H u n g a r i a n N a t . The
H i s t . Museum, B u d a p e s t ) .
age o f the common v o l e s was a l s o c o n s i d e r e d
i n the
investi
g a t i o n . The body l e n g t h o f specimens was t a k e n as an I n d i c a t i o n of age.
The animals
below 75 mm ( l ) , (4),
were d i v i d e d i n t o f i v e g r o u p s
7 5 . 1 - 8 5 mm ( 2 ) ,
and above 105 mm ( 5 ) (see
vestigations of SYKORA, 1959)
t h e ages
phasize
the p o s s i b i l i t y
t h e r e f o r e the
95-1-105 mm
2 ) . Supported by t h e i n
(BASENLNA, 1 9 5 3 ;
o f t h e s e g r o u p s were
SEBEK, 1959;
considered
about 1-2 m o n t h s , ( 3 )
months, ( 4 ) about 4-6 months and s i d e r i n g the w e l l - k n o w n
8 5 . 1 - 9 5 mm ( 3 ) , Fig.
several authors
( l ) younger t h a n 1 month ( 2 ) ,
as f o l l o w s :
( 5 ) o l d e r than
t o be
about 2-4
6 months. Con
problems o f a g e i n g I w o u l d l i k e t o em o f overlapping
between
t h e groups,
f r e q u e n c y o f s t r o b i l o c e r c i examined i n
each age
group and t h e t r u e age o f the a n i m a l s i s o n l y a p p r o x i m a t e . From a synoecological
standpoint
can be t r u l y e s t i m a t e d ,
the
i n f e s t a t i o n o f g r o u p s 1 and 5
because specimens o f t h e s e g r o u p s a r e
s u r e l y younger t h a n 1 month o r o l d e r t h a n 6 months.
Results and Conclusions
l)
S t r o b i l o c e r c o s i s was f o u n d g e n e r a l l y i n common v o l e
tions from Microtus
varied agricultural cultures;
a r v a l i s specimens were i n f e s t e d
popula
222 ( 7 . 2 %) o f 3097 (Tables
1 and 2 ) . The
degree o f i n f e s t a t i o n i n t h e more abundant p o p u l a t i o n was g r e a t e r t h a n i n the
smaller populations.
( I tis
shown i n Table 1
t h a t t h e abundance was d i r e c t l y p r o p o r t i o n a l t o t h e number o f examined s p e c i m e n s . I n Bük f i e l d
t h e i n v e s t i g a t i o n s were c a r r i e d
out d u r i n g one p o p u l a t i o n c y c l e . There would have
been a
peak
year i n 1 9 7 1 , b u t t h e p o p u l a t i o n was decreased by c o n t r o l s c a r r i e d out r e p e a t e d l y 122
i n s p r i n g and autumn. T h e r e f o r e
the
popula-
Table 2 Infestation i n 1969-72
i n t h e Bük p o p u l a t i o n a c c o r d i n g t o t h e seasons and c u l t u r e s A b ü k i p o p u l á c i ó f e r t ő z ö t t s é g e 1969-72-ben é v s z a k o k é s k u l túrák szerint
c o r n
leguminous D a t
1969.
1970.
1971.
1972 .
e
n
inf .
n
97 34
2 1
2.1 2.9
1 4 47
4.7 1.4 9.4
11
inf.
%
2
2.4
—
4 49 120
1 0 4
25.0
1.7 11.3
3.3
-
21 279 499
4 1 16
26.6 1.2 8.4
60 16 68
7 2
11.6 12.5 4.4
75 94 258
19
3
14.6 3.2 7.3
1 1
4.3 6.2
88 60 19
6 2 0
6.8 3,3
111 76 19
7 3 0
6.3 3.9
568
27
4.7
1563
98
6.3
0
17 230 379
0 4 45
II-V VI-VIII IX-XII
15 78 190
II-V VI-VIII IX-XI I
16
—
961
t o t a l
%
—
II-V VI-VIII IX-XII
Összesen
inf.
—
13
23
n
84
VI-VIII IX-XI
"
Total -
%
72
3
—
—
"
7.3
t i o n was moat abundant i n autumn 1970, a t a minimum i n 1969 and small i n 1972.) 2)
The d i f f e r e n c e s a c c o r d i n g t o sex and age a r e shown i n F i g .
2. ( I t i s based on 2912 - o r 220 i n f e s t e d of t h e t o t a l
specimens examined
number o f 3097 specimens c o l l e c t e d . )
f i n d a r e l a t i o n s h i p between age
We d i d n o t
o f specimens and degree o f i n
f e s t a t i o n w i t h s t r o b i l o c e r c o s i s i n mobile v o l e s . We have a s c e r t a i n e d t h a t t h e c y s t s have no d i s c e r n b l e e f f e c t on the l i f e
of the hosts.
The d i f f e r e n c e between sexes was n e g l i
g i b l e , b u t t h e degree o f i n f e s t a t i o n i n t h e v a r i e d specimens o f different the
sexes and
ages was o f i n t e r e s t
i n some r e s p e c t s .
On
b a s i s o f t h e presence o f f u l l y d e v e l o p e d c y s t s among t h e
w e a n l i n g v o l e s (2 cases, see m a l e s ) and r e g a r d i n g s e v e r a l expe riments
(MLODZIMOWSKA, 1931;
1962) showing t h a t more s i t e t o develop
JARRETT, I 9 6 0 ;
OLIVIER,
t h a n 30 days a r e needed f o r t h i s p a r a
i n a l b i n o mice,
i n f e s t a t i o n w o u l d have
DOW &
i t can be concluded
occurred during the e a r l y
that the
s u c k l i n g age
by p a r a s i t e ova b r o u g h t i n t o t h e n e s t .The i n v e r s i o n between t h e number o f d e v e l o p i n g and
f u l l y d e v e l o p e d c y s t s i n age groups 2
and 3 o f f e m a l e s p r o b a b l y i n d i c a t e s a more equal development o f f e m a l e s . But t h e males can be supposed t o be more s e n s i t i v e t o an oncosphera i n v a s i o n resulting
i n t h e e a r l y phase
o f the i n f e s t a t i o n ,
i n a g r e a t e r degree o f m o r t a l i t y
and a s m a l l e r number
of m a n i f e s t e d s t r o b i l o c e r c o s i s . (The home range - area o f a c t i v i t y - i s much g r e a t e r i n males - even o f j u v e n i l age - t h a n i n females (REICHSTEIN,
1951),
posure t o t h e i n f e s t a t i o n .
meaning t h e o r e t i c a l l y a g r e a t e r ex A c c o r d i n g t o DOW and
male mice were more d i s p o s e d t o t h e i n f e s t a t i o n ,
JARRETT the
OLIVIER
expe
r i m e n t e d o n l y on m a l e s . A c c o r d i n g t o MLODZIATTOWSKA, t h e j u v e n i l mice were much more s e r i o u s l y a f f e c t e d symptoms o f
by t h e i n f e s t a t i o n . The
i l l n e s s and g e n e r a l l y t h e d e a t h s
f o l l o w i n g a few
days a f t e r i n f e s t a t i o n were a l s o e x p e r i e n c e d b y t h e s e a u t h o r s . ) 3)
The above c o n c l u s i o n i s
the
i n t e n s i t y o f i n f e s t a t i o n . A c c o r d i n g t o t h e numbers o f deve
loping cysts,
c o n f i r m e d by t h e
29 males were i n f e s t e d
i n v e s t i g a t i o n of
by one and
20 o t h e r s by
F i g . 3:
Seasonal i n c i d e n c e o f s t r o b i l o c e r c o s i s i n t h e Bük po p u l a t i o n A s t r o b i l o c e r c o s i s szezonális előfordulása a Bük-i populációban
Spring
n = number o f examined specimens vizsgált állatok száma leguminous pillangós kultúrák corn gabona
Summer
Autumn
I n f e s t a t i o n of voles i n leguminous s t a n d s pil langósokban élő populá ció fertőzöttsége I n f e s t a t i o n of voles i n corn f i e l d s gabonában élő populáció fertőzöttsége
125
more t h a n one c y s t s . These v a l u e s i n females were 41 and 19 r e s p e c t i v e l y . C o n s i d e r i n g t h e f u l l y developed l a r v a e , 43 males and 50 females were i n f e s t e d by one and 9 males and 9 females by more t h a n one l a r v a e . We f o u n d s e v e r a l t i m e s some l a c t a t i n g and a t t h e same t i m e g r a v i d f e m a l e s w i t h an i n t e n s i v e i n f e s t a t i o n . The g r e a t e s t number o f l a r v a e f o u n d was 6. 4 ) There were i m p o r t a n t d i f f e r e n c e s i n t h e o c c u r r e n c e o f s t r o bilocercosis a c c o r d i n g t o t h e seasons (Table 2 and P i g . 3 ) . The i n f e s t a t i o n was a t a minimum d u r i n g summer o r t h e p e r i o d o f v e g e t a t i o n , and a t a maximum i n w i n t e r p o p u l a t i o n s . T h i s i s p r o b a b l y connected w i t h t h e v o l e - k i l l i n g a c t i v i t y o f p r e d a t o r s o r some o t h e r f a c t o r ( d u r i n g t h e i n f e c t i v e phase o f oncosphera, resistance o f voles). Aware o f t h e r e p r o d u c t i v e c y c l e i n t h e v o l e p o p u l a t i o n , the specimens i n f e s t e d by s t r o b i l o c e r c o s i can be c o n s i d e r e d i n d i c a t i v e o f t h e whole p o p u l a t i o n and t h e i r disappearance f r o m t h e p o p u l a t i o n u n t i l summer t i m e i n d i c a t e s a r e l a t i v e l y q u i c k mor t a l i t y o f o v e r w i n t e r e d animals n e x t s p r i n g . These d a t a c o n f i r m the r e s u l t s of t h e seasonal changes o f t h e s t r u c t u r e i n M i c r o t u s a r v a l i s p o p u l a t i o n s shown by MARTINET ( 1 9 6 7 ) , MARTINET and SPITZ ( 1 9 7 1 ) . ( I n o u r e x p e r i m e n t a l area we f o u n d the sexually i n a c t i v e - non b r e a d i n g - season i n 1968/69 f r o m t h e m i d d l e of Oct. t o t h e middle o f A p r i l , i n 1969/70 f r o m Dec. t o t h e end o f March, i n 1970/71 f r o m Oct. t o t h e m i d d l e o f P e b r . and i n 1 9 7 1 / 72 f r o m the end o f Sept. t o t h e m i d d l e o f March.) I n autumn, t h e s m a l l i n f e s t a t i o n s o f c o r n c u l t u r e s ( P i g . 3) are p r o b a b l y due t o t h e f a c t t h a t these a r e ploughed and sown i n October-November .The m a j o r i t y o f t h e s e p o p u l a t i o n s c o n s i s t e d of recent immigrants from other t e r r i t o r i e s . The m i g r a t i o n i s p r o b a b l y an e x p l a n a t i o n o f t h e f a c t t h a t strobilocercosis i s more f r e q u e n t even i n autumn i n these areas t h a n d u r i n g t h e summer. 5) Considering the i n f e c t i o u s sources m e n t i o n e d , o n l y t h e Bük v o l e p o p u l a t i o n i s r e a l l y known . The area r e p r e s e n t e d by c o l 126
l e c t i o n s i s about 3 km,as t h e crow f l i e s , f r o m i n h a b i t e d a r e a s . S t r a y c a t s have n o t been seen on our s a m p l i n g s i t e . During t h e autumn and w i n t e r seasons o f t h e f o u r year i n v e s t i g a t i o n s we have s e v e r a l t i m e s e x p e r i e n c e d , t h a t weasels p r e d a t e d t h e com mon v o l e s . I n 1971,the t r a p p e d v o l e s were a l s o e a t e n b y a f i e l d p o l e c a t ( i t s h o l e was t r e a t e d by g a s s i n g w i t h PH^ and t h e body dug o u t ) . On the b a s i s o f t h e s e o b s e r v a t i o n s , H y d a t i g e r a t a e n i a e f o r m i s can be supposed t o be a common p a r a s i t e i n o t h e r p r e d a t o r s p e c i e s . T h i s i s c o n f i r m e d by s e v e r a l p u b l i c a t i o n s : NEMESÉRI & HOLLÓ ( 1 9 6 I ) mentioned s t r o b i l o c e r c o s i s t o occur i n r a t s and m i c e - e a t i n g dogs; HINAYDI (1971) i d e n t i f i e d t h i s spe c i e s i n f o x e s i n A u s t r i a ; PALOTÁS (1969) s t a t e d , t h a t t h e house cat was preceded by f o x , f i e l d p o l e c a t and weasel f r o m t h e p o i n t of v i e w o f v o l e - e a t i n g .
Acnowledgement
I w i s h t o express my thanks f o r i d e n t i f i c a t i o n work and p r o f e s s i o n a l a i d t o Dr. É. MURAI-, and t o my coworkers M. VERHÁS and Miss Zs.VITÁLYOS f o r p r o v i d i n g a s s i s t a n c e i n c o l l e c t i o n s and e l a b o r a t i n g t h e d a t a , and t o my w i f e E. NIKODÉMUSZ, f o r t h e t r a n s l a t i o n , and t o M. REEVE f o r s u p e r v i s i n g t h e t r a n s l a t i o n .
JNECHAY, G . : A mezei pocok Hydatigera
taeniaeformu
lárvákkal
való
fertőzöttségének évszakos változása Magyarországon
Az ország különböző részeiből származó 3097 mezei pocok 7,2%-át találta fertőzőttnek.A fertőzöttség gyakorisága a gazdák egyed sűrűségének növekedésével e m e l k e d i k . Szezonális ingadozás i s észlelhető, l e g k i s e b b a fertőzöttség nyáron, a vegetációs idő szakban. Szerző feltételezi, hogy a hímek gyakrabban fertőződ nek, de a p a r a z i t a kártétele iránti f o k o z o t t érzékenységük m i a t t - különösen az invázió szakaszában - nagyobb a mortalitásuk i s . Ezért a fertőzött nőstények száma v a l a m i v e l nagyobb, m i n t a fertőzött hímeké. A k i f e j l e t t cysták jelentős károsodást való színűleg nem okoznak. Populáció-dinamikai szempontból a fertő127
zött gazdák "jelölt" egyedeknek tekinthetők, amelyek segítségé v e l demográfiai következtetések vonhatók l e , p l . a szaporodási időszak kezdetén az áttelelt egyedek éliminaiódásának időtarta mát illetően. Megfigyelések és i r o d a l m i a d a t o k alapján feltéte lezhető, hogy a H y d a t i g e r a t a e n i a e f o r m i s a macskán kívül más ragadozó f a j o k n a k i s parazitája l e h e t .
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SYKORA, I . :
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NECHAY, G. MÉM KNKL Rágcsáló-Biológiai Laboratóriuma 9762 T a n a k a j d , Hungary
