Episyrphus balteatus            !"  albomaculata (Dip.: Syrphidae) Functional response and mutual interference of Episyrphus balteatus and (Dip.: Syrphidae) fed on (Hom.: Aphididae)

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123  Episyrphus balteatus #/>>  # >> A>>)' B>>+C , <=>>%  ? >>% @>>. EF>+ (> CG &C F> B>, ! ± ',  )*2'- D2# , Scaeva albomaculata #>L >2- ,C>' /2>% (K>   2> (K J C+ ,  I, !! ± H #/> B+C , # ? % @. B. 2,# NO*' P QFRS "C # - ,Q  . (#

(Th)  Q>, ">'-  (a) CRQF> >L 5 > O% CW' B -# V' .  ' U, TC+ - #2,>Y' E. balteatus B>+C UC>  U, V 5 > > X  #> . (# #L ,Q ,C' B>  Q>, ">'- # 2 ,>Y'  /H HJ  /H H\ H  /H H!: V,>?' Z> %#% B> CRQF L B>>+C UC>>  U, 5 >> >> X #>> . >>2,# BE>>]' /H J  /H !HJ  / ! Z >> %#%   Q>, ">'- # 2,Y'  /H HJ!!  /H H!:H Z %#% B CRQF L #2,Y' S. albomaculata " > ' , ?' ^E% ' (, B _2Q+ & # . ' (, B /H !  /H !! Z %#% B E. balteatus #b> (P) UC> 5>  >  X `a#>% `Q 2 >)S  (a) CRQF L `Q 2)S   # <=% Z 2#d Y'  ,C , ?' S. albomaculata B+C , Bc  5 2 ' (+ ,C

∗ Corresponding author: [email protected]

! ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

B> /Hg \ V,>?' C fE BQ ,K HH `a#% - B 27e%  #b 5 2  UC 5  X . . ' (, <=>>%  ? >>% @>>a  Scaeva albomaculata  Episyrphus balteatus : 889 88 7 1 . . Myzus persicae

Abstract Functional response and mutual interference of two aphidophagous syrphid Episyrphus balteatus and Scaeva albomaculata fed on Myzus persicae were determined in laboratory conditions at 25 ± 2ºC, 55 ± 10 relative humidity and a photoperiod of 16L:8D h. Using logistic regression, a type II functional response was determined for both predators.

The Rogers model was used for estimating searching efficiency ( a) and handling time ( Th). For the first, second and third instars of E. balteatus , the estimated values of searching efficiency were 0.0359, 0.0420, and 0.0677 and the values of handling time were 2.257, 0.506 and 0.316, respectively. For the second and third instars of S. albomaculata , the estimated values of searching efficiency were 0.0590, and 0.0655 and the values of handling time were 0.575 and 0.252, respectively. According to the obtained results, no significant relationship was observed between log searching efficiency ( a) and log third instars density ( P) of E. balteatus , but this relationship was significant in S. albomaculata and the value of interference coefficient for third instars of this predator fed on 100 density of M. persicae was -0.274. Key words: Episyrphus balteatus , Scaeva albomaculata , Functional response, mutual interference, Myzus persicae. % - ,Q> B28 #  Y % (2#2'  `QF  52#%#j i '  52#% hC' #d0 V0 , V#>Q. 52f)2> > 6 52#% `$' - .(Dent, 1981)  ' CQ 2 # k  6 >?Sc' #> = B> , , , B>. ,#.   ? EG " , - ,Q B "C% ' 2  -> +  (>  #d0 V0 , . ( BQ2 @2f  " - ,Q lCm= , #W+   K D2#> , > B>+C n? 6#8 , EF+ (SC$ 5 o    pCQ2-8   #/ B . . (Jervis and Kid, 1996) (   V#Q. 6 52 6#QF qK )*2'-

! AB>C @ & ? 1*4 & => ( : <  *4 :;

>? EG " , 52#% `$' -  #   A)'   r-*. # W+ C= BQ  #/ 27>k `>2s .(Francis et al., 2001) + '   B p,  BSF/2  (*. ,  52 > X > BQ=> > B>+C #>u. ,  ,C> TC >Q' tS + <0#' ,  ,C+= 52 ,# x0 ,  $' @Y+  BQ v2 B  wQO'  # - B27e%  B.  ' #/ .(Peck, 1988) . ' 2 -*.  `QF C. P2sCSC V,?% B>a B>a > >' Syrphidae ,C+>=  B+C - /2 Episyrphus balteatus De Geer A)' >> >>' >> BQ>> P>>2sCSC V#>>Q. <>>'CK 52#>>% `>>$' B>>  - " #/>> >> X , B>+C 5>2 . (Ankersmit et al , 1986; Sadeghi and Gilbert, 2000; Alhmedi et al., 2008 ) BQ> ( ?  B. 2,# 6f ZSk B+C "CK B 5 y "C2 (S2 ,  UR+ ?Sc' #> z>K . (Youzhen and Xueyan, 2000) 2 + ' V#Q.  Myzus persicae (Sulzer) C fE > >' {#>c' f + "QF)+ C*. -*. d , ZSk B+C "CK B |C B+C 52 > B>+C - #)2, /2 Scaeva albomaculata (Maquart) A)' .(Dean, 1982; Gilbert, 1986) (>> >> 6f>> w>>QO'  C*>>. - B>>. >> >>' ,C+>>= 5>>2 # />> B>  -  #   A)' - B+C ( Radjabi (1989) f + "#2 , . (Soos and Papp, 1988) > BQ P2sCSC V#Q. <'CK - ) B. ( ,#. #?'  Scaeva A - B+C P2 BQ>> P>>2sCSC V#>>Q. <>>'CK - >>/2 "C>>K B>> B>>+C 5>>2 . >> >>' C>> ' ">>Q=, (>   > 6f> f> + ,#.#$> U, k , Brachycaudus amygdalinus (Schouteden) . . (Nourbakhsh et al ., 2008) >KzG > " 2. "f ' - 2 Q   V#Q. , ? EG " , - ,Q - >L >> "  7>> B2'#>> B>> (EF>>+ B>>0#' 5>>2 - A>>8  , (>>, B>> >>. > #Q'>8 ># h>2#G - >? EG " >, >2- VC ?' CG B . (Waage, 1990) ,C ,#>>  >>' C>>I ">> f ' >>2 B>> ?G >> ^>>E% , >> " #)>>. >>Q  QF>>2- B> "C>% >' B>?Sc' < L Q  v2 5 2#Q $' B  - . (Donelly and Phillips, 2001)    #b> `a#% # e% # j%  ( ? % @. ) / "f ' # B ?G `.#% # e% # j% > U>R+ B?Sc' , .(Jervis and Kidd, 1996) ,C +  (<=%  ) " #CRQF L

!: ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

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J ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

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N a = N t [1 − exp (a(Th N a − T ))]

" f ' B S `a#% = N t  ,C=   " f ' ,?% = N a

 Q, "'- = Th ? EG `Q2)S B28 = exp CRQF L = a @2'- "'- ' = T

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J ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

"> ' , . >2 >' , Z+R' CI B (2$+ , BI0 ]'  BQ2 @ . _2% B  5 >, UC> TC>+ ? >% @a , DY ( Holling, 1959 )  {#c' ? %   @a TC+ #>Q$ B>R Q+ ,  ,#>. <> K `a#% B BQF  CI B B ?G `a#% -  ,]' , ? EG - #>)2, >/2 S. pyrastri  > U>R+ B>?Sc' , . >2 + V#>Qa  " ( ?  +C% ' TC>+ ? % @. f + Z  fE BQ wQO'  `.#% - B27e% ,  #  #/  B+C >  X ? >% @. 5 y , #)2, * v8 , . (Fan and Zheng, 1990) ' (, B U, >' (>, B> U, TC>+ - f> + C> fE> BQ> w>QO' > `.#>% B> (EF+ E. balteatus . ,, (>>Y c' #>d0 h>> Y]% - >' (>> _2>Q+ > B>>a (Youzhen and Xueyan, 2000) TC>+ - f> + A. pisum  A. craccivora > BQ> B (EF+ E. balteatus  X ? % @. . . (Susetya Putra and Yasuda ,2008) ( 2 ,#  ,# U, E. balteatus B>+C UC % V 5   X  Q, "'-  CRQF L  #Q'8 BF >2 Y' . (  ,, "*+ V , S. albomaculata B+C UC  U, 5   X  > X , CRQF> >L 52#Q *> >y # Ba ,, "*+ |C   #Q'8 " G ,0 " > ' >  *>' >% >$% >'  ' UC 5 B ^C #' E. balteatus B+C #/ z>K . ,C , ?'  ' “]S - #b 5 2  UC  V 5    X  CRQF L - f > + |C> B+C  X wQO' 5   Q, "'- # 2 ,Y' ,   *' zQ= 5 2  # , UC> 5> > X f> + S. albomaculata B>+C , .( V> ) ,C , ?' ' #W+ #>W+ - >  *>' >zQ= ' +,C  #Q* CRQF L , U, 5  BF2Y'   > X  Q, "'- # 2 ,Y' ,   *'   % ,C 5   2 . ,CE+ , ?' '   > X , #Q'>8 5 >2  Y'  ,C , ?' ' “]S - #b 5 2  UC  U, 5  B>?Sc' ,C' #/ B+C , BF2Y' , .( V ) ,C U, 5   X - #Q a UC 5  CRQF> >L   #Q'8 # 2 ,Y' , %zQ= y # B. 2,# NO*' # 52 , >zQ= $% '   *' B?Sc' ,C' B+C , UC  U, 5   X  Q, "'-   >' #>W+ - B>?Sc' ,C' B+C , UC 5   X  Q, "'- # 2 ,Y' ,   *' .( .( V ) ,C , ?'

J\ AB>C @ & ? 1*4 & => ( : <  *4 :;

#/  A)' ? % @. @2'- , P QFRS "C # - ' (  #Q'8 ST R ( C fE BQ wQO'  `a#% B (EF+ Scaeva albomaculata  Episyrphus balteatus Table 1. Results of logistic regression analysis of the proportion of Myzus persicae consumed by predator flies of Episyrphus balteatus and Scaeva albomaculata

Predator species Instars Parameter Estimated value SE P-value

E. balteatus First Intercept 1.6052 0.6497 0.0135

Linear (N 0) -0.2168 0.0940 0.0211

Quadratit (N 02 ) 0.00562 0.00346 0.1041

Cubic (N 03 ) -0.00005 0.000034 0.1555 Second Intercept 1.2746 0.3763 0.0007

Linear (N 0) -0.0429 0.0278 0.1228

Quadratit (N 02 ) 0.00051 0.000518 0.3254

Cubic (N 03 ) -2.000048 2.000539 0.3277 Third Intercept 2.6088 0.1815 0.0001

Linear (N 0) -0.0277 0.00320 0.0001

Quadratit (N 02 ) 0.000065 0.000015 0.0001

Cubic (N 03 ) -5.00000059 1.00000808 0.0020 S. albomaculata Second Intercept 2.1070 0.2766 0.0001

Linear (N 0) -0.0430 0.00998 0.0001

Quadratit (N 02 ) 0.000186 0.000093 0.0452

Cubic (N 03 ) -3.00000006 2.0000281 0.1802 Third Intercept 2.7189 0.2240 0.0001

Linear (N 0) -0.0239 0.00387 0.0001

Quadratit (N 02 ) 0.000048 0.000018 0.0068

Cubic (N 03 ) -3.00000053 2.00000146 0.1002

J! ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

D>C% ( > ,C>=   BQ I,  ,?% )  #b "f ' #ua0 B ^C #' #2,Y' @>a   ]' ` #% ,  2,# ,# ?  % @a TC+ 5 ?%  # ,Q ,C' V' .(   >   V> ) (># #>L ,Q> ,C>' f> +  #b I,  ? % > X , #/ "f ' #u.0 B.  , ' "*+   V ,  Bp KzG , f> + #Q'8 52 #2,Y' 52 #Q .   ' UC 5  X B ^C #' #/ B+C , # . ' (, B U,  V 5   X , Z %#% B S. albomaculata  E. balteatus  B+C , UC>  U, 5 > > X , #Q'>8 52 ' (, B #2,Y' BF2Y' , 52 # zK B>+C U, 5> X # /> "f> ' #u.0 B. 2,# NO*' B?Sc' ,C' #/ B+C 5>2 #2,Y' S. albomaculata B+C , S  ' S. albomaculata B+C - #Q* E. balteatus . . ,C #Q* UC 5 X , #Q'8

Episyrphus balteatus #/  A)' (±SE) ? % @a  #Q'8 #2,Y' UT R ( C fE BQ wQO'  `a#% B (EF+ Scaeva albomaculata  Table 2. Estimates of functional response parameters (±SE) of predator flies of Episyrphus balteatus and Scaeva albomaculata to proportion of Myzus persicae

2 Predator species Instars (a) (Th) (T/T h) (r )

0.0359±0.00718 2.257±0.2192 E. balteatus First 10.63 99.07 (0.0212-0.0506) (1.8077-2.7059) 0.0420±0.00555 0.506±0.0472 Second 47.43 98.00 (0.0307-0.0533) (0.4101-0.6018) 0.0677±0.00812 0.316±0.0143 Third 75.71 94.23 (0.0515-0.0839) (0.2876-0.3449) 0.0590±0.00786 0.575±0.0312 S. albomaculata Second 41.74 0.87 (0.0431-0.0749) (0.5118-0.6381) 0.0655±0.00902 0.252±0.0144 Third 92.24 0.87 (0.0473-0.0836) (0.2233-0.2814)

JJ AB>C @ & ? 1*4 & => ( : <  *4 :;

: <*2 K (  ) K LM( :*N  *H IJ  F9 GV  E. balteatus B+C UC 5  X `.#% `Q2)S " '  , ?' ^E% B. ,, "*+ _2Q+ ' “]S - C fE BQ ,K ‘‘ `.#% < Y' ,  " B+# CRQF L `Q2)S L 5 )+ ' B. ( " #)+*+ , ?' ^E% ,C UK . (P > 0.05) (+ ,C V )  2 + @ .  " `.#% @2f  #/ 52 UC 5  X B+# CRQF Ž.(

#/  A)' #/ <.  B+# B+# CRQF L (  ?' c= ±) 5 )+ ' WT R ( C fE BQ ,K ‘‘ `.#% < Y' , Scaeva albomaculata  Episyrphus balteatus Table 3. Mean (±SE) per capita searching efficiency, per capita and total predation rate of Episyrphus balteatus and Scaeva albomaculata to 100 Myzus persicae

Predator Total Per capita Per capita searching Predator species density predation rate predation rate efficiency

* E. balteatus 1 59.00±3.75b 59.00±3.75a 0.91±0.09a 2 79.00±5.94a 39.50±2.97b 0.97±0.23a 3 85.00±3.80a 28.33±1.27c 0.68±0.08ab 4 88.50±4.03a 22.13±1.01cd 0.83±0.11ab 5 90.50±3.00a 18.10±0.59d 0.40±0.07b S. albomaculata 1 74.00±4.70c 74.00±4.70a 1.47±0.25a 2 91.17±2.96b 45.58±1.48b 1.44±0.24a 3 97.00±1.15ab 32.33±0.38c 1.19±0.12a 4 98.50±0.56ab 24.62±.014d 1.14±0.00a 5 99.00±0.51a 19.80±0.10d 0.91±0.00a % ‹ V Q0 ~c , , ?' zQ= ,C UK  , "*+ ( B+C # # ) "CQ # , B *' #0   5 )+ ' * .(Duncan )  ' *Means followed by the same letter in a column (for each species) are not significantly different at 5% probability level (Duncan).

J ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

( ( ) C fE BQ  ,C=  B ?G I,  (•y ) ? % @.  ]' ST 2 Episyrphus balteatus A)' ( 5 p8 ) UC  ( D ) U, , ( X ) V 5   X DC% Fig. 1. Functional response (left) and percentage of prey eaten of Myzus persicae (right) by first (upside), second (navel) and third (bottom) instars of Episyrphus balteatus

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( ( ) C fE BQ  ,C=  B ?G I, ( •y ) ? % @.  ]' ?T E  Scaeva albomaculata A)' ( 5 28 ) UC ( X ) U, 5   X DC% Fig. 2. Functional response (left) and percentage of prey eaten of Myzus persicae (right) by second (upside) and third (bottom) instars of Scaeva albomaculata

 #/> 5>2 UC> 5>  X `.#% `Q2)S " ' ^E% S. albomaculata B+C , ,C> , >?' C fE BQ ,K ‘‘ `.#% < Y' ,  " B+# CRQF L `Q2)S . >' (>, B> Log a = 0.200 – 0.274 Log P C>I B "C # D= BS,?'  (P < 0.05) "C ># D>= Z  ",C ' B BC%  . ,C ‘/‘  # # f + (r2) )QFE Z2#d "f ' #> - B> B+#> CRQF> L 5 )+ ' X `.#% @2f  B. (# BR Q+ "C% ' `.#>% B> BQF>  <>' K P>2 "C>K B> <=>% ,C #)+*+ #' 52   2 ' @ . X > BF>2Y' , S. albomaculata B+C UC 5  X , <=% ,C .( Ž V )  '

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@2f> - B> B+#> CRQF> L 5 )+ ' #Q* @ . (K B —X Q0 E. balteatus  X #Qf Bu < S, B +C% ' f + #' 52 B.  ' B+C 52 #/  X `.#% TC>dC' B>. >+YQ?' 5 YY]' =# .  E. balteatus B+C  BF2Y' , B+C 52 UC 5 2#]>I D2#> ,  ,, >EQK ,>]' > D ]'  )*2'- D2# , DY <=% L 5 )+ ' # , . ( ˜+  F #b B+C } 2 ,# 5 <=% ,C V Q0  BQ> , ,C> 2#]>I  )*>2'-  . _2Q+ 5 ,2- zQ= B+# CRQF  B>?Sc' V>0 # B . (Jones and Hassell, 1988) ( ,C #Q . ? EG D2# , <=% "f '  > B>+O >+' ,>]' > D >]' , >? EG " , -  CW' B <=% 28 . . ( ,C=# #Q* (  - -* a XCm]'  E+ B>+C , #> , #/>  X `.#% @2f B. ,, "*+  @2'- -  ) ,,#> >' B+#  <. #/ "f ' @ .  @2f ZE Z %#% B B?Sc' ,C' , Z> %#% B> #/> <>. "f> ' 52#>Q .  52#Q* #/ A)' B+C , # , .(  B+#> "f> ' 52#Q*> 52 # zK . ' (, B UC 5  X ,K   !  `.#% >' (>, B> UC> 5> X ,>K  `.#% , #/ A)' B+C , # , #/ . . ∗ ( V )

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ALHMEDI, A., E. HAUBRUGE and F. FRANCIS, 2008. Role of prey-host plant associations on Harmonia axyridis and Episyrphus balteatus reproduction and predatory efficiency. Entomologia Experimentalis et Applicata, 128: 49-56.

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5 >' &>$'  ES>G #e>I >K #>Q., ">   ,-# &$' C8 ]Q ƒCY?2 #Q., : '<K*K KYK ∗ QF>>8 |>>I &>>' (>> #% )*>>+, -*>>. /*>>+, - *>>. >> #*>>0 #>> " %>>I ."#2 "#$%  \! gJ

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Address of the authors: Eng. F. JALILIAN, Dr. Y. FATHIPOUR, Dr. A. A. TALEBI and Eng. A. SEDARATIAN, Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, P. O. Box 14115-336, Tehran, Iran.

 ... Scaeva albomaculata Episyrphus balteatus       : '*24 '(

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