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Diversity and Abundance of Pest Insects Associated with Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) in Bockle and Dang Localities (North-Cameroon)

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Abstract

Cowpea plants are damaged by insects in North-Cameroon. During ecological survey (2021 and 2022) in 44 plots of 4x3.5 m each, insects were captured on stems, leaves, flowers and pods, stored in vials containing 70° alcohol, identified in laboratory and the community structure was characterized. We captured 26,015 adults belonging to six orders, 13 families, 19 genera and 19 species. Coleoptera, Hemiptera and Hymenoptera were species-rich orders [five species each (26.3%)]. Hemiptera was mostly abundant (40.0%) followed by Coleoptera (27.6%), Hymenoptera (21.9%), Lepidoptera (0.9%). Heteroptera and Orthoptera were least abundant (0.8% respectively). We recorded five (26.3%) useful species [the West African predator species Cheilomenes sulphurea (Coleoptera: Coccinellidae), and four (21.1%) afrotropical Apidae species [Apis mellifera adamsonni, Amegilla calens, Amegilla sp. and Xylocopa olivacea]], seven (36.8%) phytophagous species [the indomalayan native Aulacophora indica (Coleoptera: Chrysomelidae), Nearctic native Danaus plexippus (Lepidoptera: Nymphalidae), Palaearctic native Dolerus sp. (Hymenoptera: Tenthredinidae), afrotropical native Hypolimnas misippus (Lepidoptera: Nymphalidae), afrotropical native Monolepta marginella (Coleoptera: Chrysomelidae), Palaearctic native Phyllotreta cruciferae (Coleoptera: Chrysomelidae) and the Eurasian native Tettigonia viridissima (Orthoptera: Tettigoniidae)]. We recorded seven (36.8%) sap-feeding species [the afrotropical native Anoplocnemis curvipes (Hemiptera: Coreidae), cosmopolitan Palaearctic native Aphis crassivora (Hemiptera: Aphididae), old world native Bothrogonia sp. (Hemiptera: Cicadellidae), subtropical native Dysdercus cingulata (Hemiptera: Pyrrhocoridae), western Palaearctic native Lagria hirta (Coleoptera: Tenebrionidae), North American native Poecilocapsus sp. (Hemiptera: Miridae) and the Palaearctic native Riptortus dentipes (Heteroptera: Alydidae)]. Giving up eight (42.1%) native species, 11 (57.9%) non-native species and 14 (73.7%) pest species [three natives species (15.8%) and 11 non-natives species (57.9%)]. The abundant species were M. marginella (39.9%), Poecilocapsus sp. (14.4%), Au. indica (11.4%), Ph. cruciferae (10.4%), C. sulphurea (4.6%), H. misippus (3.7%), L. hirta (3.4%), Ah. crassivora (2.4%) while 11 species (57.9%) were rare (<2.2% each). Insects associated with cowpea consisted mostly of non-native species and the situation calls for more research on the bio-ecology of recorded pests. Cowpea plants' insect assemblage mostly presented in 39 Moukhtar Mohammadou et al.: Diversity and Abundance of Pest Insects Associated with Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) in Bockle and Dang Localities (North-Cameroon) Bockle and Dang, a fairly significant regeneration force (Zipf and Zipf-Mandelbrot functioning models) and all conditions combine to soar. Due to the numerical and behavioural dominance of non-native insects, a significant number of resources are potentially exploitable. In due course, once the invaders would monopolize available resources and saturate the localities, they would not allow native species the niche opportunities to re-establish themselves. The consequences of loosing native species, which may well interact with the endemic flora and fauna, will be of extreme concern.
American Journal of Entomology
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Diversity and Abundance of Pest Insects Associated with
Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) in
Bockle and Dang Localities (North-Cameroon)
Moukhtar Mohammadou
1, 2, 3
, Boris Fouelifack-Nintidem
1
, Moïse Adamou
2, 3
, Taïmanga
4
,
Daniel Kossini
3
, Sedrick Junior Tsekane
1
, Babell Ngamaleu-Siewe
1
, Edith Laure Kenne
1
,
Abdel Kayoum Yomon
1
, Martin Kenne
1, *
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Cheilomenes sulphurea5!5"!.!"0I!0!2"!HApis mellifera adamsonni.
Amegilla calens. Amegilla  !" Xylocopa olivaceaJJ. 3 I ,!  H "!!,! !3
Aulacophora indica5!5,"!.&!!3Danaus plexippus/"!&,!"!.)!!!
!3 Dolerus  G,! =""!. !0! !3 Hypolimnas misippus /"! &,!"!.
!0!!3Monolepta marginella5!5,"!.)!!!!3Phyllotreta cruciferae5!
5,"! !"  @!! !3 Tettigonia viridissima *! ="!J B "" 3 I
!0"  H !0! !3 Anoplocnemis curvipes G! 5"!. ! )!!! !3
Aphis crassivora G! 2""!. " " !3 Bothrogonia  G! 5!""!. ! !3
Dysdercus cingulataG! ),"!.)!!!!3Lagria hirta5!="!.&
2!!3PoecilocapsusG!7"!!")!!!!3Riptortus dentipesG!2,""!J
83   #I !3  .  '(I !3  !" # I   H !3 
'I!"!3'(IJ=!"! M. marginella((I.Poecilocapsus##I.
2"!#I.Ph. cruciferae#I.C. sulphurea#I.H. misippusI./!#I.Ah. crassivora#I
'(I!KI!$!!"!",0!3!"
!!0!,0""5!!L !! ,"
( 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
1; !" 4!. ! 0!, 0! ! 0 +0 !" +07!" 0 " !" ! "
!4!!"!3!"!0!3.!0!0!
!,D!$".3!""F!3!!!"!!!.,
"!!3!3=90!3.
!,!"0!!"0!!.0D
Keywords:
Vigna unguiculata.)$.1"3,.&5!
1. Introduction
5! Vigna unguiculata / B!. # 6!!
6!!! !"  % 20! !" "!, !"
  " " @. /! 2! !"
%!2!H.J.03!"!0
 !3 !" " " 0   0 ! !"
3;HJ$!!!!
30.0.,."
!" 3! 2. 5 !" @ !" ! ; . .
!.!!.!."!"!
 "0 ! 3!  %%!!! 20!
H'J$%%!!!20! !
"   !"   !  "3
!!"!,.!"!!,0!
" ! 0 3! " !"  3!
"003!"!"
5"!0!!!3!!,.
3! 9 !3  !""  !
!3!!,H#J5!0"
!  0, ;  !3!! 0! 0 20!
00!!"
!"  ! "  D !"
"! 3 0 0!.  !; 0 3" 3!.
00  0 0F !"   0,.
D00!.!D.!
"3.!3.3!.!0
!!".0!!"!.!!;.!
! . !"9! ,!!, 
" "! !"  . ". !, 0
!! .   0,.   0 0"
!!!."!!"0!.
" ! "   9!,. !!
! ! !" !  H'(J 2 
!0!0!0"!"
"0!""!;!!.
!  !"   0! 0!. " 
,!0!!"9,!
 "  3! 9!, !" " ," 7!, !!
! ! !! !" "! !  !
!!  H. J  3! 0 2"! !"
G!"!0!! ! !H. J$!;
!" .   ; !  0! !3, 0
!  !!" , ! 0 !0
,!.  !" !;  H#J
5! ! ! ! ! 0 3! !!;  !
! 0. 3!" !!!" !F!
! ,! !" D,! !  %3!
!!   ! ! 
!"!"!!3!0.!!
! "   !   0  !3.  ! !
0"!"!!0!! 
!0.!3"!"0
!!"00"9"",
!.   !  !" 0"  ! 0
G!  ! !" !" !,  !  
!M!D!"!!!""!"
!"!!","HJB$
0"!!.!3! ""0"!;!
D  !!  !=H(J=
","!! !!"", !
 "! "! ! !00 "3, 0  !!;"
! H. J = !; 0 ," !" !" 
9!,0"!"!0!!""!.
!;!! 03!3!
5!=",!
0!.9!,0"!"!3,!00,"
&3!I0!!""
00 ! 0! !" !3  
!3!"33!HJ=3,"!"
3!!;"!"."!,
 , !  !3 !" !"9! !.
!"  !0 00  !. 3. 0
.  ! !"   D"   0
!0" ,  ! 0 ! ! H. #J =
!39!"!!30
,!!3!""0!!3
"0!! ! ! 0
3!!H'J&!"!,.!
! 0  !! " !  003
!!30.30!
6  . !, !! !""3 0 ! !3
 " 003     6
D! !0 !9 D! 0 3! !  
"003!!HJ=!
 0! ! !"  ! !3 
3, ""  5! H. J %"  
0!! !!"  ! ! !3 
""  3!  " -%2. $"!.
$"!.@,.7!!,!.%"!!"=!"!
"   !3 00 0   H. J
2 !  ", 3!"  5!. 
" "!! D   "3, 0 !!" 
&3. 0   0 ! 
!   3  ! 3! $ 
!3!!0 5!.3! !3
2!?!0@, #
  ! 0   0 !! " 
! ! ! !!3  , " 0 
!"!0!0HJ $!!!
0 8!! !" &!" &5!. !;
!"!3!.,!!",
,0!. "!".!" !"!!3 !
0!,   ),! "!! !  "" 
!, !" !! 4  "30" 0!
!" ! 0 !; !" !3,.   
0!   ,  !"
 0  !!"  ! ! =
0 ","0, !3 !
!!!!.;,"!!!
!"  09!, !"
9!, 0 !! ," = ; " !; 
  !!F   , 
!!"! 0""8!! !"
&!"&5!
2. Material and Methods
2.1. Study Sites
=",!""!"."!,
!.&5!=""
 !! 0 !" !"  !3!!, 0
3!80"!!;!
8! 8)% =   ""  1;
(EN(O&.E'N#(O@.!"(!1;!"
4! E'M#MM&. EM##MM@. !" #  !
1;!"
"08!!&!"
4!  ! "  " 0 &!" 2"!!!
P 1 ! ! !"    8!
"" ! !3!! HJ !" "  
"!%!!!!F.! !"!"!
"! !0! H. (J = 3! ! 
!!,!%"!8!!
!!,!02*0!,!!"!
",!0&37!0  0 ,!
H#J=!!3!(E5!"!
!.# ,!= !3 ", 
6!,I.!"!3!!!,,I
H.(J69,3"!Cosmos sulphureus
5!3.(2!2!!, Helianthus annuus /.'
2!2!!, Tithonia diversifolia G,8!,.
2!2!!, Cajanus cajan /G.(
6!! 6!!!, Phaseolus vulgaris /. ' 6!!
6!!! !" Sesamum indicum / ' %!!
)"!!! = 0  " ! 
!!,"0Bombax costatum )!"C.
(#7!3!7!3!!, Borassus aethiopium 7!.
2! 2!!, Boswellia dalzielii G. (
%!"! 1!!, Commiphora africana 2 P
@.  %!"! 1!!, Hyparrhemia rufa
&%!0.(()!)!!.Lannea microcarpa @
!" > >!. ( %!"! 2!!"!!, Prosopis
africana 8!")=!.(6!!6!!!!"
Vittellaria paradoxa 568!.@!%!!!
)!! 0 Azadirachta indica %!"! 7!!.
Eucalyptus camaldulendis 7,! 7,!!, Cassia
occidentalis 6!! Caesalpiniaceae !" Hyptis
suaveolensis /!! /!!! ! 0" 5ultivated
areas were small plots of polycultures family farms
2.2. Sample Design
5!!  !F"  ,0
!" .  0 !"    ,
")!!!#@5''$&$"!"!"
!"!,!.!!3"5!
!"!,",0!H.#J!"!"
!9!3D!0!HCalotropis
procera 2 2.  8!! 2,!!.
Eucalyptus camaldulensis4.7,!7,!!
01;.!"Tithonia diversifoliaG28!,
2!2!!04!J83!!
0##0"0#D'!!"!!
"" 0"!" !"F"
; "  0 ! 2 ! 
!!!"00!!"0
0!) "!!
= !; 0 ! " 3!, 6; 
!"0$P248!!200!.!
"   ! #  ! ! !" ' 
 ! !" " # "!, !0   
!%D 0" !"!
"0=""
 ! ## " = ; !0 . 0
!   !!! 0  0 0. "
!!"!!"!"!6!!"
"!,!0!. !,!!
!"!5!!"
!" ( ! !"   !" ' .  !
!""0"!,0
 20! !!. !"
";.0!!0"."!!"0
!"!3=,!!"!""
!;0!0!
!!!!!.!3.0
!""5!0!#"!,!0!
!""!30")!"
""!,!;0!! "!,!"
 0"  . !. !3 !" 0. 
!"   0 !   0 0  0
!0,  !3! !6, 
 !"  !    !0 !F 
!!"!,2"00
3" 2#F!"30"";
!    !"  ;  !" 3!
! E ! 4  !3 ". 
0"    " = !0" " 
"!";"3,"!,!!,!
0!"10,!!!" !3!
"!"!"!!,
# 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
2.3. Identification of Insect Specimens
$  "0"    3  !
!0, !. ;, !"!" !! H##J 
/!!,02"+,.4!01!
%. 6!, 0 %. -3, 0 &!".
3""$""
"3!D,0"
!"!"!!
2.4. Data Analysis
4!!"!D!"34!!
!D0!"!00!!"!
,!")!!!" 0
3!! 0 " 2"!
"0!Q!"!"
= !  !"   %"  
3!!"!,!"9!3!!!"
$!""!!BD0
!"   7!B, 0 "" 
&!!!03!!"!!
>;!B!0%!%!0!
R!" ! ! !  4L
" 5! 0  09 ! " 
6L D! !" ! ! 0 3!
09!"66!G!0
%!S! 0!  !" !! ! 
!" " 0   0 !   
9! 10 " H#J P 9!
!""2&*C2"%!0
  !!  ""  )2%= '
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!"T%0""T0
   = !  = !
""   !"! !" !!
! 5! !"  !  %=: 
!" = " 0 "! , ! 0  !
3!!"1)!;"D$
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= 3!  3!! ! 3!!" 
%L " H#(J 1  ! !
3!!"  >"!L tau 00 = "!,
 !" ! ! ! 3!!" 
1!,5L "D !" ! 0"   !!
G!! 5 2!, !"  ?!!"L !,
"D=   ! "  -)872 !
H'J = !; !"!  ! "  ! 
!0%24!""0 00!%24!
! " ! !" , !!  )!L
! 00 !"  ! !F" ,
1!!>!  ! H'J B " 03 , "
!%24H'J031;;"
1%.8"87./!"/&.
+0"+!"+07!"+7"=
 0" " ! "  2$5 "
!F",?!"*!"HJ=!;!3!
0 P # 0!H'J "   !"   %24 1%
" ! !  !! D    !
!"!0 H'#J B"" !!0
87  /7    "" L 
""" 87 ""   !D !"! 0 
0!
!"7!L3!
=!!3"!,!0!"!
!;H''J+"!"+/"H'J.!"!
 !;"  "! " + "  !"  
!X!!!!F!.!"
Y!!"!,00!3!!,0
!!!0!H'.'J+7!!F""
Z!!"""7!9!"L!!9!
!!F",/!H'J!"7,H'(J.!
"!Z!"YB!!"Y0!!"
0"0"3"!!H.J
3. Results
3.1. Inventory of Insect Species
2 ! 0 .'   " .
  (I  1; !"'.# 
I4!="D".
0!. ( ! !" (  %3 0! 'I
H2""! /!.  G! /!. '.
2"! /!.  G,! /! '.
5,"! /!.  5! /!. '.
5!""! /!.  G! /!. '.
5"! /!  5! /!. '.
7"! G!.  G! /!. ' !"
="! /!.  5! /!. 'J
" "  !"     1; !"
4! 2,""! 2, [ %3. # G!
/!.  !  "" "   1;
5"! /!. ' G! /!. ' !"
="!>!.(*!/!(
"""1;),"!2,[
%3.#G!/!.'!""
"     &,!"! P!09. '
/"! /!. ' !" =""! /!.
 G,! /! '   ""
"4!2"""0!L"!!
=! .  "00 !   "! 3!
! 1; !" 4!!"  ,!
 !" .  !, !!,  0!
H1;    0! #.   T 
!DT#' !QT# Q''
"!3!7T((4!
0!.'. T. !DT''Q
2!?!0@, #
.7T'""!!
0!..T.!DT'.(Q#
7T#"3"!1;0!
'.'#"T'!DT.(
'Q7T'#"3"!4!
0!(.'T( 
!DT.''(Q("!T(
"3"! " "!!    0!. #'.#
.  T ' . !D T . 
.' Q #  "! T # "3"!
! ! 0 "! 3! 
>;!B!!!!,2&*C2GT
"0TT!!0!3!
!!!,2&*C26T((T
(JG3.!,!!"!0",.
3!!   ! 0  0 0! 
!!, 0!   66! G! 
G3.!,!!"!0",.!
3!!   ! 0  0 0! !
!!, 0! =!  )! !
" !  1; !"  ", ,!.  0!
  F"    "00
 33" 0!   0! H  0
03 0! 2""!. 5!""!. 5"!.
="! !" ="!.   0  0!
2"! !" 5"! !"   0  0!
5,"!!" 7"!J=!  !
".00!H000!
2""!. 2""!. 5!""! !" =""!.
  0  0! 5"!. &,!"! !"
="!.   0  0! 2,""! !"
),"! !"  # 0  0! 5"! !"
="!J=!$4!!"",,!.
0!   F" H  0  0!
2""!.2""!!"5,"!.0
0!2,""!.5"!!"=""!!"
00!&,!"!!"="!J=!
  !  "  .  0! 
H  0  0! 2,""!. 5"! !"
),"!.   0  0! 2"!.
5"! !" ="!. !"   0 
0!&,!"!!"=""!J=!1!"
!03!"""
,!!"",.0!,
"".!;"""!2""!
0((I0",5,"!#I
!" , 2"! # I =  0!  !,
""2,""!I5!""!I5"!
(I 5"! #I7"! 'I &,!"! (I
),"! #I ="! I =""!
'I!"="!I
4!!!!,!"=!"
!#I""",,!
!=Aphis crassivora
>. '# G! 2""!. Aulacophora indica
8. ( 5! 5,"!. Bothrogonia 
7!. ( G!! 5!""!. Cheilomenes
sulphurea *3. ( 5! 5"!. Lagria
hirta/!.' 5!="!.Monolepta
marginella B. ( 5! 5,"!.
Phyllotreta cruciferae 8F.  5!
5,"! !" Poecilocapsus  P. 
G! 7"! =  'I HAmegilla 
6. ( G,! 2"! !" Riptortus dentipes
6!.  G! 2,""!J ""  " 
1;  "   !! =  
'I HAmegilla calens / ). # G,!
2"!!"Dysdercus cingulata6!.'G!
),"!J"""!!
='IHDanaus plexippus/!.
' /"! &,!"!. Dolerus  )!F. 
G,! ="! !" Hypolimnas misippus
/!. # /"! &,!"!J  
""4!" !!.0
 !   !    6  I
HAnoplocnemis curvipes6!.G!5"!.
Apis mellifera adansonii /! # TApis mellifera
/!. ' G,! 2"!. Tettigonia
viridissima /!. ' *! ="! !"
Xylocopa (Kopthortosoma) olivacea 6!. 
G,! 2"!J   ""  1; "
!!=!
1!"   " "!!. M. marginella !  
"((I.0",Poecilocapsus##I.
Au. indica#I!"Ph. cruciferae#I'
(I HAh. crassivora#I. Am. calens'I.
Amegilla  I. An. curvipes (I. Ap. mellifera
adansoni I. Bothrogonia  I. C. sulphurea
#I. Dolerus  I. Da. plexippus 'I. Dy.
cingulata 'I. H. misippus I. L. hirta #I. R.
dentipesI.X. (Kopthortosoma) olivaceaI!"T.
viridissima #IJ  ! , " $
!'I"!0"!!=!Da.
plexippus"D3,1;",!
Ap. mellifera adansoni!""D3,"
  !!  4! 4   !!.
3!!0 !!0! 
03IHAm. calens6LD!T.
An. curvipes  T .Ap. mellifera adansoni T.
Bothrogonia T(!"Dy. cingulataTJ
$ !.   #I " 0!,
!01;HDa. plexippusT
(D(. Dolerus   T D#'. H. misippus  T
D.M. marginellaTD#.Ph. cruciferae
T D. Poecilocapsus   T D'. X.
(Kopthortosoma) olivaceaT'D(!"T. viridissima
 T DJ 6!, ! 0 D  
0!,4!HAmegillaTD.Ah.
crassivora  T D'. Au. indica  T (D. C.
sulphurea  T '#D. L. hirta  T D !" R.
dentipes  T DJ 4   !!.
!3!!!0!03
IHAmegilla6LD!T((.
BothrogoniaT.Dy. cingulataT##!"R.
# 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
dentipesTJ
Table 1. Occurrences and percentages of collected insects by family. Percentages are calculated from the total number of specimens.
Family
Year 2021 Year 2022 Both years (2021 and 2022)
A. Bockle
(%) B. Dang (%)
C. Global
(%) A. Bockle (%)
B. Dang (%) C. Global
(%) A. Bockle (%)
B. Dang
(%)
C. Global
(%)
$2,""!     # #'  #' 
$$2""! #   '# ( .#
 '# .#
((
$$$2"! (( ('  '  .'
( ((' '.''
#
$C5,"! .#''
.'# '.
' .( .''( #.#'#
#.'#( .#
#
.(
#
C5!""! (## #  # '' #(  ' #'
C$5"!  # '# ' # (# '( (# (
C$$5"!   #'  (# '#  ' '#
C$$$7"! .'
( .'(
(' #' #' .( .##''
#'
$S&,!"! '(' #( ( #  ' '( #( (
S),"!    ' ' ' ' ' #
S$="! # ( '#    ( '' 
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2021 Bockle: α’ (p)
Dang: α’ (p)
2021 Bockle: α’ (p) Dang: α’ (p) 2021 Bockle: α’ (p) Dang: α’ (p)
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#
:
$vs. S$$ 
##D

:
D

D

:
$$$vs. S$$$ D
(
:
D

D
'
:
C$vs. S$ #D

: 
$vs. S$$$ 
D

:
D

'D
(
: $Cvs. C (D
#
: (D
#
: C$vs. S$$ 'D

: D
#

D

:
$$vs. $$$ (

'D

(D
#
: $Cvs. C$ D
#
: D
#
: C$vs. S$$$
('D
#
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
:
#D

D
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:
$$vs. $C D
#
:
D
#
:
$Cvs. C$$ D
#
: D
#
: C$$vs. C$$$
D
#
'D
(
:
D


D
'(
:
$$vs. C '
#D

:
'D
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D
#
: $Cvs. C$$$ (D
#
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: D
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: C$$vs. $S ((D
#
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
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D
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(D

:
$$vs. C$ 
'D

:
D

D

:
$Cvs. $S D
#
: D
#
: C$$vs. S 'D

: (
$$vs. C$$ 
D
#
:
D

#D
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$Cvs. S D
#
: D
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: C$$vs. S$ D
#
: D

D

:
$$vs. C$$$ D
#
D
#(
:
'D
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D
'
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$Cvs. S$ D
#
: #D
#
: C$$vs. S$$
(D
#'
: D

(D

:
$$vs. $S 
D

:
(D
#
#
D

: $Cvs. S$$ #D
#
: 'D
#
: C$$vs. S$$$
(D
#'
: D

#D

:
$$vs. S 
'D
(
:
D

'D
#
:
$Cvs. S$$$ 'D
#
: D
#
: C$$$vs. $S
(D
#
D
(
:
(D
#

D
(
:
$$vs. S$ #
D

:
D

#D

: Cvs. C$ 'D

:
'D
_
'D

:
C$$$vs. S D
#
D

:
('D
#
(
D

:
$$vs. S$$ 
(
((D
#
#
D

: Cvs. C$$ D

:
D

#D

:
C$$$vs. S$
D
#
(D

:
D

D

:
#' 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
2021 Bockle: α’ (p)
Dang: α’ (p)
2021 Bockle: α’ (p) Dang: α’ (p) 2021 Bockle: α’ (p) Dang: α’ (p)
)!!10""
C$$$vs. S$$
'D
#
(D
'
:
D
#

D
(
: $Svs. S$$ #D
(
:
' S3S$$$ 'D: D
#D(:
C$$$vs.
S$$$
D
#
'D
(
:
#D
#

D

: $Svs. S$$$ (D
#
'D

: #D

D

:
S$3S$$ D#: D#
D(:
$Svs. S 
'(D
(
:
D

'D

:
Svs. S$ (D
(
:
D

#D

:
S$3S$$$
D#: D#:
$Svs. S$ 
(
'D
#

D
(
: Svs. S$$ ##D

:
D

'D

:
S$$3S$$$
' D
D:
66G=66!G! ^0!3^L10"0!3 0!`^`^L:0!K^K^L
Table 2. Insect species composition in the study sites, number of foraging adults, and dominance status of recorded insect species.
Year 2021 Year 2022 Pooled two years
Order/Family
Species Bockle
(%)
Dang
(%) Total (%)
Bockle
(%)
Dang
(%) Total (%)
Bockle
(%) Dang (%)
Total (%)
5!
5,"!
Aulacophora indica8
(
a.$7.)
HJ (
(
.(
# #((
.
#(
.#
( 
.'

.(
#
Monolepta marginellaB
(
:.)
HJ '(
.#
(
.
'
.#(

'.'

.
#
.##
(#
.(##
'
.#
((
Phyllotreta cruciferae8F

a.)2.)
H#J '
#
.#
## 
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2"! Amegilla calens/)
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H
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Amegilla6.(
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H
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
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Apis mellifera adansoni
/!#
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HJ    '
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
Xylocopa olivacea6!

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Dolerus)!F
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Hypolimnas misippus
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HJ '(
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="! Tettigonia viridissima
/!'
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:
!0!!3
a
!3.
5*%
5!.
@-2
@!!!3.
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)
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)2
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!3.
&2
&2!.
&@
&!!3.
-
-0.
%-=
%!""
)!003I0!, 1;HDa. plexippusTD.Dolerus
2!?!0@, #
T(D#'.H. misippusTD.Ph. cruciferaeT
!" Poecilocapsus T #D##J6!, 
 ##I  , "  4! HAh.
crassivora6LD!T.An. curvipesT
(D. Ap. mellifera adansoni  T D#. Au.
indicaTD.C. sulphureaT(#D#.L. hirta
 T #D.M. marginella T . X. (Kopthortosoma)
olivaceaTD!"T. viridissima TDJ
2"   3! " "!! 0   ,!.
 1; !" 4!.  0! "00 !
"" 'IHAm. calens 6LD!
T!"L. hirtaT##J3
,""1;HPh. cruciferaeTD.
Poecilocapsus   T D'. X. (Kopthortosoma)
olivacea  T D. Da. plexippus  T 'D(.
DolerusTD.H. misippusT'D!"
T. viridissima  T D#J 6!,   'I
,"4!HAmegilla6LD!
TD.An. curvipesTD.Ap. mellifera
adansoniiTD#. Au. indicaTD(.Ah.
crassivora  T 'D. Bothrogonia   T . C.
sulphurea  T D. L. hirta  T D#. M.
marginellaT!"R. dentipesT#DJ)!
!!,0"0("
"  ,!    "  "3"
  0    "00  
   0!   
!!!,0!0!
!"003HAm. calens, Ap. mellifera adansoni,
Da. plexippus, Dy. cingulata!"T. viridissimaJ="
" 0 HAmegilla . An. curvipes
!"R. dentipesJ="!!"0
HAh. crassivora !" Dolerus J !"  0 
"0H/!!"GJ&
! 0 . 0!"00  ""
!Ap. mellifera adansoni Am.
calens, Da. plexippus!"Dy. cingulata=!2
2"      0 !
!  =!  Ap. mellifera adansoni !" Dy.
cingulata  "" D3, "   1;
!"4!Amegilla!"R. dentipes""
1; "   !! Da. plexippus !
"" D3,  1; "  !" 
!!Dolerus!"H. misippus""
4!"!!='IHAn.
curvipes. T. viridissima !" X. (Kopthortosoma) olivaceaJ
""1;"!!Am.
calens ! "" D3,  4! "  
!! 6!,   #I  ""
!, 1;!" 4!" ! ,!0
", Ap. crassivora. Au. indica, Bothrogonia . C.
sulphurea, L. hirta, M. marginella, Ph. cruciferae !"
Poecilocapsus  B 3  1;. ! ! 0 
#I"!!!"'
(I"!!$4!""!!
0 '   (I "   !! !" 
#(I"!!="00
   !  !  0! 6
D!T*!0
0!,2"!!"0I.
0" , 5,"!   'I. ,
/"!   'I !"   !
0! 2,""!. 2""!. 5!""!. 5"!.
5"!. 7"!. ),"!. ="!.
=""! !" ="!  !, ".
!,,'I=!1!
 !" ! ,! 3!! !  !!,
0!66!G!D!\]T#."0T
.T0!\]T'."0T.T0
!,!
*!0  0"!3. !3!!"
!0!".""'I
"!  C. sulphurea 5! 5"!
!3B20!.!"!!
 !D!, !! !". 0 I 0
20! !3 G,! 2"! ; !
!HAm. calens.Amegilla.Ap. mellifera adansoni
!"X. (Kopthortosoma) olivaceaJ.DI,!
  H "!!,! !3  Au. indica
5!5,"!.&!!3Da.
plexippus/"!&,!"!.!!!3
 H. misippus /"! &,!"!. 
20! !3  M. marginella 5!
5,"!.)!!!!3Ph. cruciferae
5!5,"!!" !! 0 @!!
!3 T. viridissima *! ="!J. 3
I! "; H !0!!3
 An. curvipes G! 5"!. 
! )!!! !3  Ah. crassivora. 
"" !3  Bothrogonia .  !
""  Dy. cingulata.   )!!!
!3  L. hirta 5! ="!. 
&! !3  Poecilocapsus  G!
7"! !"  )!!! !3  R. dentipes
G! 2,""!J = )!!! !3  !"
0DolerusG,!=""!
!""=Dolerus!3!!0"!.,
D!,."!3"0.
3"),
" 03 0   I 3 # 
I6LD!T(B3
! ! 0  20! !3  #I !"
0!3",!!!"D
'(I.  "00   0! 6LD!
T'
3.2. Insect Species Abundance
2!0.'!""",
%T(.#!"!Q!"!"
.(Q''"3"!"!3!7T#'!"
5"   "3""  . !" 0
1; (#I' .##' Q
7T!"!"'.#!"04!
(#I.(##!"(Q#(
# 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
7 T  !" 4   !!.  "
#."3"!1;#I.
Q(#7T ( !"!".'
!"04!'(I.#!"#
Q"3"!7T4!!.
"'.'#"3"!1;'(I#
.#(Q(7T'#!"!"
(.' !" 0 4!    #I (  '.'
!"'(Q("3"!7T#=!!"=
"00 "  0 ! "! ! 
0!>;!B!09!"!GT
"G T #T )!0"
 3!" 0!,    ! 
66!G!!,\]T.#."0T'.
T"!!\]T.."0T.T
"!!\]T.#."0T'.T"
!!\]T#.(."0T.T0"
 ,! "!! 1  ! . ! 0
!""3"!0!,"00"!H1;
 I versus 1;   'I 10L
!" "0!3^LT'.T#D

1;
versus 4!#I^LT.TD
(

1;versus 4!I^LT'D

.T
1; versus 4! ^L T'. T
D

1;versus 4!^LT.T
J)""!!0,!"!3!
,"1;(I!4!I
6L D!  \] T .##.  T  8!,
! 0   "" 3!, 0!,
0!\]T."0T.T((
&""D3,!=
!0!0!!3'(I0
!  HAmegilla . Am. calens, An. curvipes, Ap.
mellifera adansoni, C. sulphurea, H. misippus, M. marginella
!"X. (Kopthortosoma) olivaceaJ!!!,"00
0!0
#I 0  !  HAh. crassivora, Au. indica,
Bothrogonia , Da. plexippus, Dolerus , Dy. cingulata, L.
hirta, Ph. cruciferae, Poecilocapsus ., R. dentipes and T.
viridissimaJ%"T#."0T.T#
3.3. Insect Community Structure
=  0  "" 0    !
1;!"4!!!"3!"!
!. . ! 
3!=!
Table 3. Matrix of the species richness, diversity, evenness and dominance indexes for each study site.
Indices 2021 2022 Both years
Bockle Dang Global Bockle Dang Global Bockle Dang Global
2P"D
I #. .' . '.'# (.' '.# . '.# .'
% # '  '  (   (
!D
 #  #( ''  ## (# '
7 '#  #    # ' 
"T%         
5! # '  '  (   (
%!@00         
143,"D
GL  (( '  ' # ( ' #
GL
!D
T% (     (## ( ( (##
4 #  ' ( '  # ( 
&
T
GL
(  (  '  (  
&
T4  '    #   '
P!5!%       (  
G!T&
&
   (' '# '  ' 
5@3"D
?TGLGL
!D
#    '  '  (#
@4!"D
$
T
!D
 (    '( # ( ' ((
)!!0"3,"D%"
5! %!B!3"DGL %L"3,"D
1;34! T"0T.T#D
#
: T("0T.#T'D
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#
: T"0T#.'(TD
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13 T#"0T.'T'D
(
: T"0T.TD
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:
1,!1;34! T("0T'.#T#D

: T#"0T.'TD
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0!"00:0!"00!F!D!D!"!%3"77!!0"D
"!GL%!B!3"3,"DG!D!D%!B!3"3,"D4%L"3,"D&GL0"
"3,&TGL"""3,GGL"3,!?)L3"D$1)T1)!;"!"D
=!"1;%
T#7!!0"D7T'#!"T
!"!""4!
%T  7T" T=
"D3!0"D2
!! 0 "3,"D " !
2!?!0@, #
! 0! "00.   " ! 
"3,.!3,)L3!
,!"!"!,!01)!;
"D!!0"!3!=!
=!;!"! 0  " "!! " !
!3 !!!     
,0 "! 62 =! !
! "   !"   " 0
!"! %24 6 1 !" 5 = "3"!
!0!3" 0 ! !"
  ,! !!"  !! !!
 !  6 4 = ! ! 0 
!;!"!  ! 3"  ! ",  !"
"!,!6=33"1;"
!!"0!!!,!0.
     !"   
 ! "  4! "  6 1 6 !
!"!" ! 0 #.' "3"!.    4!
 !!"  "3 H@%T#.' T  Q J.
0" , 4!  H@%T#.' T ' Q  J. ,
1;H@%T#.'T'QJ!"!,,1;
H@%T#.'T #QJ5"5!
!! !.  !  "  !
!.! !D!I=!.!
!  !" "    1!" 
GM0!" """3,.0
,!"!!!0
"!!"3!0GM!3,
,=!!3.!!"!0
""!!,!0
Figure 1. Rank-frequency diagrams of the collected insects in Bockle and Dang during 2021 and 2022 showing species in order of numerical dominance and the
species rarefaction curves among the two study sites and the two collection years.
#( 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
Figure 2. Rank-frequency diagrams of relative insect species abundances collected from two study sites. For each site percentages were calculated on the total
number of individuals collected.
*  ! 0  GL 0 " "3,  
=!!3!"!;!"!6!".
0,!"!3!"003
I0!"4!
 ##I "   1; !"   
!, , !"! =,  "3""  
2!?!0@, '
 'I "" D3,  1; H
)!!! ,!   Dolerus 
G,! =""! ""  . 
!0! !3 !  X. (Kopthortosoma)
olivacea G,! 2"! ""   !" 
!!;Ah. crassivora G!
2""!""J.'I""
!1;!"4!!"D3,!1;
H!!!"!0!!3,!
H. misippus /"! &,!"! !"  
)!!!  !;  L. hirta 5!
="!J !" 0!, 03  I 
""!,  !"  ,! =
"!!,!!3,!Au.
indica 5! 5,"!,  !0! !3
"!C. sulphurea 5!5"!.
!0! !3 ,!  M. marginella
5! 5,"!.  )!! !3
,!  Ph. cruciferae 5!
5,"!!" &!!3 !; 
Poecilocapsus . G! 7"! P!  
3,=!6!"
2!0#!,!I"3""
!'I"D3,1
H!!0@!!!3,!
T. viridissima*!="!J!""
 ,! H &! !3 ,!  Da.
plexippus /"! &,!"!J = ! 
'I  " D3, ! 4!. !, 
 'I   H !0! !3 !
 Ap. mellifera adansoni G,! 2"! !"
!;L. hirta 5!="!J
!"   'I HH. misippus /"!
&,!"!J0"!3"!!0,!
B ""  !  HAh. crassivora G!
2""! !" An. curvipes G! 5"!J 
!1;!"",!!4!2!
 H ""  "; 
Bothrogonia  G! 5!""!J !
!,""" ,! ! !
 = !  H )!!!   !" "
 Dolerus  G,! =""!J !
""!4!!"!1;
= )!!! !; !  R. dentipes
G!2,""!!""!1;!"
" ,!4!=0!G,!
2"!HAmegilla!"X. (Kopthortosoma) olivaceaJ
""!"!!01;!""
!"4!6!,!HAu. indica
5! 5,"! !" Dy. cingulata G!
),"!  ""    1; !" 4!
3,1!"GL &
"D=! !"
%246!"."!
!" 3!"0'I
4!3I1;
%3 I ! "!  !3 
"0" $ . Dolerus  "!"  !!
"" 1; =  'I H !0!
!3"!C. sulphurea 5!5"!.
!!,! H. misippus /"!
&,!"!!" Poecilocapsus .J "!"
,!L !!  1; !" ! ""  4! 
 *  'I HPh. cruciferae 5!
5,"!J"!"",!!!
 1; !" ! ""  4!   6!, 
'IHM. marginella 5!5,"!
!" Au. indica 5! 5,"!J "!"
!!0!",!
1!"    . ! ! 0
!   !". ! "! 3 0
"!,!"!"1;!"
4! !! H1!,5 "D 1;  3 4!
15T'(#1;31;15T
4!34!15T1;34!
15T 1;34!15T
=!3,30"!,15T!"
1;!"4!!3,30
"!,15T#!"4!!"
1;=00!0"
"!DH1;,!vs.
4!,!15T1;!"4!vs.
1; !" 4!  15 T J =  !!,
!;F!!?!!"L!,
"D!"4!0"0.
1;0""=""
04!!"1;6
2"  0  %24   03 , ;
!""!0!0D9!,
""!!01;!"4!T((.T
D

..0!0!,9!,1;
T(.T'D
(
.#.1;T('.
TD
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.' ."!""!!
01;T(.T'#D
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.!"
3!!!T(.TD
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Figure 3. Hierarchical Cluster Analysis based on Jaccard’s index using the “Unweighted Pair Group Method with Arithmetic mean” (UPGMA) algorithm and
showing similarity in insect assemblages among two sampling sites and two colledction years (Cophenetic correlation: 0.821).
Table 4. Akaike Information Criteria (AIC) values for the adjusted theoretical models.
SAD model
AIC (BIC) values and the best fitted theoretical model
Year 2021 Year 2022 Both years
Bockle
S = 14;
n = 4,617
Dang
S = 15;
n = 6,251
Global
S = 16;
n = 10,838
Bockle
S = 15;
n = 5,584
Dang
S = 16;
n = 9,563
Global
S = 19;
n = 15,147
Bockle
S = 18;
n = 10,201
Dang
S = 18;
n = 15,814
Global
S = 19;
n = 26,015
1;%;1%  ## # ' ''( (#  (# (
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"3!  b T
'LTbT(%:T#(!!3
0!"!J$!+"0"
 %24 ""    4! =! # H
T '.'
"3"! !F! ! X T (.' "3"!
"3!##("!,00!3!
!,0 0! YT "
! 0!" !
T (.'

J = !  !
!"""!!0=!
#H
T."3"!XT'.#"3"!"3!
# (  Y T  "
T '.#

J
)" "!! 0  ,! "  !  4!
=!#H
T.(##"3"!!F!!XT
'.#"3"!"3! ' YT(
"
T '.#
(
J = +7 "  0" 
 ""    1; =! # H"3!
. X T #..
T . "3"!. % T # 
!D

T
=
!00!3!
c T  "! 0! d
T Z T # Y T (
"
T#.V#
(
!0!!"
0"0"3"!!YT((J
=!  ! "   ""!! 0
=!#H"3!#XT."3"!
T."3"!%TD

T
=
cT
2!?!0@, '
d
TZT'YT('"
T.V'
('
!0!!"0"0"3"!
!YT#'J6!,+7"0"
 !!""  1;=! #
H"3! '(#(. X T '.'# "3"!
T .#(
"3"!.%T'D

T
=
cTd
TZT
#YT"
T'.'#V#

!
0!! " 0  " 0 "3"! !
YTJ
3.4. Association and Correlation Between Species
*!0.#!0!
!",!.3!.D!3
!!  ! "!!   ( 
%LC!!!CPT'eB!
.(('#'K2!3 ! ! 
"   !    0
!!3!!
9!!,!=!'
Table 5. Kendall’s correlation tau τ between insect species recorded in 2,640 sample units.
Species 1 / Species 2 tau τ (p-value) Species 1 / Species 2 tau τ (p-value) Species 1 / Species 2 tau τ (p-value)
2Amegilla calens 4Aphis crassivora  $Da. plexipus
Amegilla  ('#D
#
:
H. misipus D

:
Dolerus  #'D
#
:
Ah crassivora #'(D
'
:
L. hirta D

:
H. misipus #''D

:
C. sulphurea '#D
(
:
Ph. cruciferae #D
((
:
M. marginella 'D

:
Da. plexipus (#((D
#
:
Poecilocapsus . #
Ph. cruciferae 'D
(
:
Dolerus . D

:
T. viridissima (D

:
X. olivacea #D

:
Dy. cingulatus 'D

: @Apis mellifera ?Dolerus 
H. misipus (D
('
:
C. sulphurea #(D

:
Dy. cingulatus D
(
:
L. hirta #D

:
Da. plexipus D
#
:
H. misipus #D

:
M. marginella #(D
'
:
Dy. cingulatus #D

:
L. hirta :
Poecilocapsus . (D

:
L. hirta (D

:
Ph. cruciferae ''D

:
B. Amegilla .
Poecilocapsus . D

:
R. dentipes :
Ap. mellifera #D
'
:
X. olivacea '(: >Dysdercus cingulatus
Ph. cruciferae #D

: 6Aulacophora indica
M. marginella ((##D
'
:
Poecilocapsus  'D
(
:
Bothrogonia . #(D
(
:
R. dentipes D

:
T. viridissima #(D

:
C. sulphurea #D

: /Hypolimnas misipus
X. olivacea #('D

:
Da. plexipus #'#D
(
:
M. marginella D
#
:
5Anoplocnemis curvipes
H. misipus #D

:
R. dentipes #'D
#
:
Bothrogonia . #D

:
X. olivacea D
'
:
T. viridissima #D
#
:
C. sulphurea D
#
: 8Bothrogonia  7Lagria hirta
H. misipus #D
(
:
C. sulphurea (''D

:
M. marginella #D

:
L. hirta D

:
Dolerus  ('(D

:
Ph. cruciferae D

:
M. marginella ((D
'
:
H. misipus #D

: &Monolepta marginella
R. dentipes (D

:
Ph. cruciferae '#D

:
T. viridissima 'D
#
:
T. viridissima D

:
R. dentipes #D

:
X. olivacea ((D

:
4Aphis crassivora
T. viridissima (D
'
: *Phyllotreta cruciferae
Ap. mellifera #D
(
:
X. olivacea 'D

:
Poecilocapsus . D

:
Au. indica ##D
'
: GC. sulphurea
R. dentipes D
#
:
Bothrogonia . D

:
Poecilocapsus  '
T. viridissima D

:
C. sulphurea 'D
(
:
R. dentipes ''#: )Poecilocapsus 
Da. plexipus (D
#
:
X. olivacea ('D

:
X. olivacea ((D

:
Dy. cingulatus ((D
(
:
0!!:0!!&120"!."!!00!;"
!
= 0 !0! !3 ! Amegilla calens
G,!2"!!!3,!"3
 =! '2 H ! )!!! !3
!; Aphis crassivora G! 2""!. 
0 !0! !3 "! Cheilomenes sulphurea
5! 5"!.  )!!! !3
,! Dolerus  G,! =""!.
 ! "" !;  Dysdercus
cingulatus G! ),"!.  
)!!! !3 !; Lagria hirta 5!
="!.  !0! !3 ,! 
Monolepta. marginella 5! 5,"! !" 
! ! 0 @!! !3 ,! Tettigonia
viridissima*!="!J!"!3,
!"    =! '2 H 0
!0! !3 "! Amegilla  G,!
2"!. &!!3 ,! Danaus plexipus
/"! &,!"! !"  !! ""
,! Hypolimnas misipus /"!
&,!"!JAmegilla . !!3, !" 
 =! '1 H)!!! ,!
Phyllotreta cruciferae 5! 5,"!, 
' 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
& 2! !3 !; Poecilocapsus 
G!7"!!"T. viridissimaJ!"!3,
!"     =! '1 H 0
G,! 2"! !0! ! Apis mellifera
adansonni !" Xylocopa olivaceaJ Anoplocnemis curvipes
!!3,!"=!'5H"
"  !; Bothrogonia . G!
5!""!!" H. misipusJ!" 3,!" 
03  =! '5 HBothrogonia ., L. hirta, M.
marginella, R. dentipes !"T. viridissimaJAphis crassivora
!!3,!"=!'4HAp.
mellifera adansonni, Da. plexipus !"H. misipusJ!"!
3, !"  3  =! '4 H
"!!,! !3 ,! Auacophora indica
!5,"!. Bothrogonia . C. sulphurea,
Dy. cingulatus, L. hirta, Ph. cruciferae !"T. viridissimaJAp.
mellifera adansonni ! !3, !"  0
=!'@HC. sulphurea, Dy. cingulatus, L. hirta !"
Poecilocapsus .J !" !3,!" 
=!'@HDa. plexipus !"X. olivaceaJAu. indica
!!3,!"=!'6HDa.
plexipus, H. misipus !" X. olivaceaJ !"  ! 3,
!!"=!'6HBothrogonia . !"C.
sulphureaJ Bothrogonia  ! !3, !" 
=!'8HH. misipus !"X. olivaceaJ!"!
3, !"  03  =! '8 HC.
sulphurea,  )!!! !3 ,! Dolerus 
G,! =""!. Ph. cruciferae, R. dentipes
!" T. viridissimaJ C. sulphurea ! !3, !"
 X. olivacea !" 3, !"  R. dentipes
=!'GDa. Plexipus !!3,!"
 =! '$ HDolerus . M. marginella !" Ph.
cruciferaeJ!"!3,!"
=! '$ HH. misipus !" X. olivaceaJ Dolerus  !
!3, !"  H. misipus !" 3,
!"  0  =! '? HDy. cingulatus, L.
hirta, Ph. cruciferae !"R. dentipesJDy. cingulatus !,
3, !"  M. marginella !" R. dentipes
=!'> H. misipus !, !3,!"
=!'/HM. marginella, R. dentipes !"
T. viridissimaJL. hirta !,3,!"M.
marginella !" Ph. cruciferae =! '7  M.
marginella ! !3, !"  X. olivacea !"
3, !"  T. viridissima =! '& Ph.
cruciferae ! !3, !"  Poecilocapsus .
!"3,!" R. dentipes !" T. viridissima
=! '* 6!, Poecilocapsus  ! !3,
!"  X. olivacea =! ') =  
!"0!!"!!
00"!!
4. Discussion
4.1. Species Richness, Diversity, Abundance and Dominance
*"3!"0(.(!.
 0! !" D " !!"  ! !
1!"   0" . G! " 
! #I 0   !". 0" , 5!
I. G,! (I. /"! (I.
G!I!"!,*!"I0
" =!09,"!
  !; !"  3!  "
5!H. #(J =!, 0    !
, ! !3  ! !0  " !
!!.F!00"
0   !" 0!.   ! 0
!! ! ! , !!.  ! !!! 0
"3"!!!"P!"
!3  "3" ! ! 9 0 !! !"
"  0 " , ""!" 0! 
5! H. #. (. (J Dolerus  G,!
=""! !  !  0  !" "
),!  H5! 5,"! #I.
G,! =""! 'I. /"!
&,!"! (I !" *! ="! IJ
!" !0"  H5! ="! I.
G!2,""!I.G!2""!((I.
5!""! I. 5"! #I. 7"! 'I. !"
),"!#IJ.!3,"I0
 =   !,   " 
!0"$"!HJ.@,H(J.&!H(J.
  8! %!3!! 0 5! !"  %"!
%!!2!+H(J!"!;!"
!  5! H. (J.   " G!
2""!!!"!$!01;!"
4! &5!.    0 
!!" !. ! 9  3!
!",3!!!"20(
  !!"  ! !  " ,
+!!!H(#JB"" HAphis craccivora
> G! 2""!. Anoplocnemis curvipes
6!  G! 5"! !" Riptortus
dentipes6!G!2,""!J1;
!" 4! ! & 5! $ $"!.
3!0!0!!!!3!
  ! " "" "3, !" 3, 3
!. 3, !"  "!  
/"! !" 5!M " H. ('J G3
!"!!.3!!0!
!"!"!!0!!!G,!
!"*!M "7!.  3! 
."!!0"G!
0!",$%P! !!$"!.!
" !    D 0 ! !
!F"  ' " G!. =,!!.
5!.4!!"/"!#0!.
 0 0!! " ! 0 0! .
!".0.!;.!0..!".
""!" ,!"!!H(J5!"
3!.0D!%"!.
 " "!! !" ! 0". "3""
  ,! . 3  ! ",
0 !" 0.   !"  
2!?!0@, '#
H#J %!,  1!!".   ! "!!
! ! ! "3""  ' ,! .
00!"0!"
H(J = !3!!, 0 "    
0! 0 !  "  ! !!
0!3!   !! 0   = "!!
!",,! 5!5,"!.
G,!. /"! !" *! !" !0"
5! ="! !" G!  ! 
!3 !" "  !  ! 0 I = 
D!" ,  0! ! !  0 !.  !
! 3! " !! !
0 !" !!""!
0  ,!  !" 3! !3
 " !;  !"  !" "
=!0"!01;8!!
!" 4! &!"  3!" !  
0$!;!"!!
$3,5!.!!!!!!3.0!"
"!!!;"!.""
" " HJ = "00 3"  " 
D!",  09,0!3,!  !
". 0 ",,"!"
0!.  !!;   0" ,  3!,
"!,!"!!0!
*", 0  3!!!0!3!"
!3!!0!
! 3!"  1; !" 4! !
&5! !,   ! ,
"!9!3D!0!"!=
3!" !  " ! !3,  
 !" "3, !"!.  3 0 
3    ! 0 !3 
 %!  ! "  "" !
!F"3H'.(.((JP
!!"!"0!"!!
 ! Piper nigrum /   !, 0
) !5!. ! !" ! ! 
1!5!H. (.J ="3,0
!!"!"!!3
#I0!!"'I
0  ! " .    !;
D!0 = D!0  0" !"
  ! , !3" , !3  
!3   '(I 0    !"
'I 0 ! !"!%!"
!!"!!0"1!5!
H.(J=!"!303!3!3
""!;HJ=
"3,""0!3000
!! 0 !3
HAnoplocnemis curvipes 6!  G!
5"!. Monolepta marginella B ( 5!
5,"!. Hypolimnas misippus /! #
/"! &,!"!J !"   !3 
HAphis crassivora > '# G! 2""!.
Aulacophora indica 8 ( 5!
5,"!.Bothrogonia7!(G!
5!""!. Danaus plexippus /! '
/"! &,!"!. Dolerus  )!F 
G,! =""!. Dysdercus cingulata
6! ' G! ),"!. Lagria hirta
/! ' 5! ="!. Phyllotreta
cruciferae 8F  5! 5,"!.
Poecilocapsus  P  G! 7"!.
Riptortus dentipes6!G!2,""!.
Tettigonia viridissima /! ' *!
="!J = "" !3  ! 09,
"!0"  3!!An. curvipes
G! 5"!   0     0
!"!"!"5!!;
,!0"0"!""!
6D!5f"M$3A!HJ!3
! An. curvipes !    0  !.
0 (I 0  ! 3," " Anacardium
occidentalis %!"! 2!!"!!. Mangifera indica
2!!"!!. Cocos nucifera 2! 2!!.
Elaeis guineensis 2!!. Ipomoea batatas %!!
533!!.Cucumus sativusC!5!!.
Cucurbita maxima 5!!. Langenaria vulgaris
5!!. Hevea brasiliensis 7!!
@!!. Manihot esculenta 7!!
@!!. Arachis hypogea 6!! 6!!!.
Glycine max6!!!.Phaseolus lunatus 6!!!.Vigna
unguiculata 6!!!. Persea africana /!!
/!!!.Abelmoschus esculentus 7!3!7!3!!.
Gossypium barbadense 7!3!!. Musa paradisiaca
+! 7!!. Psidium guajava 7,!
7,!!. Passiflora edulis C! )!0!!.
Oryza glabberima 5,!)!!.Zea mays)!!.
Coffea arabica P! P!!. Citrus limosus
%!"! P!!. C. maxima P!!. C. sinensis
P!!.Lycopersicon esculentum%!!%!!!.
Solanum nodiflorum%!!!.S. melongena%!!!
!"Theobroma cacao7!3!%!!73.
!""!!5itrus!"&!HJ
=!0M. marginella5!5,"!.
;!!;!3!"0 !" 
!"!3!""!!"3
0 !, !"   ! 0" " !3!" 
!"HJMonolepta53!.!
08!!  ! ""
H#J !"  0   ! ""  !
H'JH. misippus/"!&,!"!  !
; 20! , !"  ,!"
0,. ,!.  ! !! "
5!! 0"  ! 3!, 0 ! 0! "
533!!. 7!3!! !" )!!! !" 
!,0",!)!!!Portulaca oleracea!"
Po. quadrifida HJ&!3  "!! 3!"
!  ,   !3 !  !  !! 0
"=!0!3H.#.
. .. . J! "" 1; !"
4!
'' 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
%!0"  3 0 ! 3 ! ,
,!!"!3"3"!3!,
"  3!  =!0 0 !0"
00!3!"!
!,;0""!"!!0!!
.!!!0!!0"
!"5!H.J1!"
  !0 !3, 0 !3   
!0".,"!,!!!3,
!8!!!"&!"&5!
*"! 30 !3
.    0  !3  !,
"" ! !   !, "3 
3!" !!  , !3  "" = 
!0!3"0
!0!,!!!.0!
!3 0 0 ""  B"". ,
" ! , " 0   1
!!!!3!!
"  !, !" 00.  ! 3!
.!00!"!!!F!"!"
"30!!!!0 "
H#J 2 ! ! " !  1; !" "!
!0,!!,!"!;!"9!
!"!!"!"3
!"!
4.2. Community Structure and Functioning Model
=!!0,!L"
"!!  1; ! 0, 0" ,  !
"//7!M "87.
7!L3!!!!
3!     T ( 4!  !"  3!
"   0"  ! 
"/&)L")L3!
!! ! 3!L T( !"L T (
3,=870"!
! !   !"! 0  
!0"!!!"!;5!,
/&)""!
!0!"!!"0!;0
  !"  0 ! ,  
! , 0   "! !"!.  87
7! " "  ! ,   !
""    !, "! 87 
!"  "0%24 03!
.0D!""!!!
!D0F!"!!!!D!!"!
),6!H(J.!!0 "
 ! !" 0  % 2 HJ. !"0,
   7!,  0 5 HJ. 
5!!"! !" G! ! !" 3 !"
!"g! !  !!" !!"  !
6!" HJ.  !! 0 !  "00
,03!!F05!HJ.
!! 0  !!"  ! ! 
1! 5! H(J 83 !  !
!! 0      0 0  !
0! !"   0  00,   
!!"   /!  /! " H#J.
"0!!F!"0
0 !" "" 3    
0D!0!3!!
=4!.,!L"
"!!  4! !"  3! " !! 0 
 !"  ,!   0" ,  +0 + "
    1;   !"  !" 
" !! 0      0" 
+07!" +7 " +0M !. 3, "
D3,.09,!"!!
,!!F!"!"= 
"!,!","!!F%240
H'.(.J6+"09,0
!3,!!;!""
"!"0!!!0!"
"!9 3! " =!
9   !! 3, !, !"  !"!.
 ! !F"   !! !.   0
!3"0"3.!"!
+!"+7"!09, 0" 0
!!3.  33",
;" " !!
 0  ! 0 0! !"  
!! ! H'. '. J = 
!! !!"  ! !  1; !"
4!!.0!0!!0!
D 0! ; "3"   !" !
+0 !" +07!" " "  ! 0
33"3! !!!"
"!0!,0!!0!"
""!3
5. Conclusion
=0",8!!!"&!"!
""3,0!!"!
! !" !!F  ,  5"
0 " D ". 0!.(
! !" (  )" "!! "  
."3,!"!"!,!0
1!"     0  " "!!.
5!. G! !" G,!  
" H03  ! IJ  /"! !
" , ,   'I. G! !"
*! !,",, 
'I !1!"!"!.G!!
, !"! #I 0" , 5! I.
G,!(I./"! (IG!!"
*!  ! !"! I 3, =
3!"3, 0 ! !!"
3!!"!!3
#I 0  !   !" 'I 0  !
 !"!.    !; D! 0
 = D! 0  0" !"   !
2!?!0@, '
,!3",!3!3
 '(I 0    !" 'I 0  !
!"! $ 1; !" 4! !. ! 
""   !!"  ! !. 
20! !3   "" HAnoplocnemis
curvipes G! 5"!. Hypolimnas misippus
/"! &,!"! !" Monolepta marginella
5! 5,"!J 4 ! !.
!!"!!"0",,!3
  = ,  0  !!"
 ! !  1; !" 4! . " ! 0!,
0! ! 0 !"  "" !
3 $  ! 0  ,. ! 
"   ! 4   ! !"
!3! "! 0 !3 . ! 0!
0!!,D!$".
  3!" " , F !3!!
 !" !!  !. , "  !
!3      !
3=90!3.
 !,  !   "  0!.   0
D  =   0  !
 ! 0  !! ,!!,  3 
"
Acknowledgements
= ! !;"  5!! 7, 0
G @"! 0 3" 0"   !
!=, !; "0 /!!,0
2"+,0-3,0&!"0!!
!"0!!"!!!
Appendix
Table A1. Pairwise comparisons of the pooled species occurrences from the two study sites and two years collections using Bonferroni’s procedure and the
Fisher’s exact p-values.
α' (p-value) α' (p-value)
α' (p-value) α' (p-value) α' (p-value) α' (p-value)
!vs.  D
#
:
vs. 0 #D
#
:
vs. ;
D
#
: "vs. 9
#D
#
(D

: 0vs. ;
D
#
#D
'
: vs.  #D
#
:
!vs.  D
#
:
vs.  #D
#
:
vs.
D
#
: "vs.
D

D

: 0vs.  '(D
#
D

: vs. (D

D

:
!vs. " D
#
:
vs.  #D
#
:
vs.
D
#
: "vs.
#D
#
D
#'
: 0vs.
D
#
(D

: vs. ;
(

!vs.  D
#
:
vs.  D

D

: vs.
D
#
: vs. 0
'D

D

: 0vs.
D
#
D
'
: vs.  D

'D
(
:
!vs. 0 D
#
:
vs. #D
#
:
vs.
(D
#
: vs.
(D
#
D

: 0vs.
((D
#
D

: vs.
D

D

:
!vs.  D
#
:
vs. ; #D
#
:
vs.
(D
#
: vs.
D
#
D
(
: 0vs.
'
 vs.


!vs.  D
#
:
vs.  'D
#
:
vs. 9
(D
#
: vs.
#D
#
: 0vs. 9
'D
#
D

: vs.
(D

D

:
!vs.  D
#
:
vs.  'D
#
:
vs.
#D
#
##D

: vs.
'D
#
D
'
: 0vs.  D

'D

: vs.
('D
#
#D

:
!vs. D
#
:
vs.  'D
#
:
vs.
(D
#
: vs. ;
D
#
D

: 0vs.
D
#
'D
(
: vs. 9
D

D

:
!vs. ; D
#
:
vs.  'D
#
:
"vs.
#D

D

: vs.
'D
#
#(D

: vs.
D

D

: vs.
D
#
D

:
!vs.  D
#
:
vs.  D
#
:
"vs. 0
#D
#
D
#
: vs.
''D
#
'D

: vs.
#D
#
: vs.
#D

D

:
!vs.
D
#
:
vs. 9 D
#
:
"vs.
D
#
#'D

: vs.
D
#
(D

: vs.
D
#
D
(
: vs. #D
#
:
!vs.  D
#
:
vs.  #D
#
D
'
:
"vs.
'D
#
D

: vs.
(D
#
D

: vs. ;
D

D

: vs. ; #D
#
:
!vs.  D
#
:
vs.  D
#
:
"vs.
#D
#
: vs.
D

D
(
: vs.
D
#
D
'
: vs.  #D
#
:
!vs.  D
#
:
vs. " ''D
#
(D
#
:
"vs.
#D
#
D
#
: vs. 9
'D
#
(D

: vs.
'D
#
D
#
: vs.
#D
#
:
!vs. 9 D
#
:
vs.  #D
#
D
'
:
"vs. ;
'D
#
D

: vs.
'
vs.
D

#D

: vs.  #D
#
:
!vs.  D
#
:
vs. 0 D
#
:
"vs.
#'D
#
D

: vs.
'D
#
#D

: vs.
D

#D

: vs.  #D
#
:
!vs.  D
#
:
vs.  D
#
:
"vs.

#D
#
(D

: 0vs.
D

D

: vs.
D

D

: vs.  #D
#
:
vs.  D

D
#
: vs.  D
#
:
"vs.
'D
#
'D

: 0vs.
(D
#
(D
##
: vs. 9
D
#
(D
##
: vs. 9 ##D
#
:
vs. " #(D
#
'#D

:
vs.  D
#
D
#
: "vs.
'D
#
D
(
: 0vs.
#D
#
: vs.
D
#
'D
##
: vs.  ##D
#
:
vs.  #'D
#
D

:
vs. D
#
:
"vs.
#D
#
D
'
: 0vs.
D
#
#D

: vs.
'D
#
D
''
: vs.  #'D
#
:
' 7;!7!!"et al43,!"2"!0)$2!"Vigna unguiculata/B!.#
6!!6!!!1;!"4!/!&5!
Table A1. Continued.
α' (p-value) α' (p-value) α' (p-value) α' (p-value) α' (p-value) α' (p-value)
vs. ; D

D
#
: vs.
#(
;vs.
D

'D

: vs.
D

D
'
: vs. 9
D

D

: vs.
(D
#
#D

:
vs.  D

D
'
: ;vs.
D

D
#
: vs.
#D

D
'
: vs.
D

D
'
: vs.
'D
#
'D

: vs.
D
#
'D

:
vs.  '
(#
;vs.

D

D
#
: vs.
D

D
'
: vs.
D
#
D

: vs.
'D

D

: 9vs.
'D
#
D

:
vs.  D

D
#
: ;vs.
'
vs.
(D
#
D
#
: vs. 9

vs.
D

'D

: 9vs.


vs.  D

(D

: ;vs.
D

D
'
: vs.
D
#
D
(
: vs.
'D
#
(D
'
: vs. 9
D

D

: vs.  'D
#
D

:
vs.  D
#
D
'(
: ;vs.
D
#
'D

: vs. 9
#D

'(D
#
: vs.
'
vs.
#D
#
D

:
vs. 9 
' ;vs. 9
D

D
'
: vs.
#(D
#
D
(
: vs.
D

D
'
: vs.
D

D
'
:
vs.  '#D
#
D
''
:
;vs.
'D
#
D

: vs.

vs.
(D
#
D

: vs. 9
D
#
#D
#
:
!7!!!5!5,"!2!!"!5!5,"!),!0!5!5,"!"
5!5!5"!/!!!5!="!02!3!G!2""!1!
G!5!""!  2 3G! 5"! )!  G! 7"!  4," !!G!
),"!;P"G!2,""!20!!"!G,!2"!2!!G,!2"!
2!  G,! 2"!  S,! 3!! G,! 2"!  4  G,! =""! 9 4!! D
/"!&,!"!G,!/"!&,!"!=!3"!*!="!
References
HJ @".G *[ $. 5 * 203!
! " ! ! " 0!  @,
%!.&!2P)&?21%.'
HJ 1!"!. 6 2. 40. 7 [ 40. 4 # 5!.
)('# $ 7 %. $ % 1 [ 7 4! "
1!"  8 1! 0 8! /. %
$"!4*$(h'
HJ 2. 1 > [ 2!,. 7 =   2 3 0 
!  0 ! C! !! / B! 0
! !" !! " ? 2 6" P.  
"  !0
H#J *!".?*.2L.8.*;.7[7!L.2B
230 0!!30"
!",@!20!6"%&.#
"0
H'J G. / [ %. G  )" ! !"
" !! 0 ! 3 0 "
!%%!!!20!22%.
'(4*$ !!
HJ 1!!.>.@!.)&.&"[*;,.@6!
200"3,0%!"!81
4. 8  +. *! P! %!. @! 43
5,".(
HJ &!=!!".&.>;&;.41[4 
&!!. @  @3!! " 0! " 9
3!D!!!3!!""
!!!!5!!"1!!L*
5!20%.(
HJ 7. > 5. *. % [ /. % G ( = =!
@00, 0 %!" $ )! )"  %!!
%"3.5!2.('(
H(J 8!. A. B;. B. 7!". G [ -!. @ 
)"5!.6!)0"!!"6!
%,05!%@!P!%9!.
4*$#'(#3
HJ 20"!./5 2. %F.4. B0.8[B,"!.>
1!  0 Macrophomina phaseolina  !
Vigna unguiculata " ", " , !!
!! $ ? 1 5%.   '' 4*$
"D"## 3'
HJ 7!. $. P,. % [ 7!. 1 >  5!!3
1, 0 = 5" )"!  5! 2"
Aphis craccivora $"! ? @.  4*$
'('$?@'
HJ =!!! [ = 6. 6&  )!
00, 0 Apis mellifera /! ' G,!
2"!  Mimosa pudica /! ' 6!!!
0!A!!4!!5!?@+
%".'
HJ 4!. 1 &. ?!;!. / @. 2!. 1 2 [
2;7!. 5  )!  5! Vigna
unguiculata $! 0 $  @!
1"3, $.   '#
"('#
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C!! !.7.%! P!. 2C7.7!!. 7 [%!,
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>J !" 5" HMenochilus sexmaculatus 6!J
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''" !
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)".1P[ 7";. / /  = "
.Maruca vitrata6!/"!5!"!.!
!  0 ! ! 3!F B
20! $ ? = $ % (
"#((#
2!?!0@, '
HJ +!. 8 C. 7!F. % [ = 6. 6& 
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G,! 2"!  Psorospermum febrifugum
G,!!!"!&!".5!?@
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Camponotus compressus 0" ", !" !3! 0!
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!%!!/!"!
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... Pest insects (case of Hemipteran [6]) and phytophagous insects (case of Coleoptera and Lepidoptera larvae) damage organs of wild or cultivated plants. Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) is a largely cultivated plant species in West and Central Africa [6][7][8][9]. Several studies on the floricultural entomofauna highlighted the negative effect of pest insects [2,3,[8][9][10][11][12][13][14][15]. Farmers frequently use approved synthetic insecticides [16] or unapproved ones in order to control harmful insects. ...
... Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) is a largely cultivated plant species in West and Central Africa [6][7][8][9]. Several studies on the floricultural entomofauna highlighted the negative effect of pest insects [2,3,[8][9][10][11][12][13][14][15]. Farmers frequently use approved synthetic insecticides [16] or unapproved ones in order to control harmful insects. ...
... Farmers frequently use approved synthetic insecticides [16] or unapproved ones in order to control harmful insects. In Adamaoua Region (Cameroon), the synthetic insecticide Parastar (mixture of 20 g.l -1 imidacloprid and 20 g.l -1 lamda-cyhalothrin, l p.c..ha -1 ) is frequently used [8,9,17]. The abusive use of synthetic pesticides leads very often to harmful effects on the health of humans, the environment and on useful insects [18][19][20][21]. ...
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Synthetic pesticides present risks of pollution of the environment, humans and livestock and the alternative proposed today is to use botanical extracts in the fields against crop pests. But in North Cameroon, little information exists concerning the effect of these extracts on useful pollinating insects in general and no information exists in particular on foragers of the genus Amegilla Friese, 1897 (Apidae: Apinae: Anthophorini). The frequency and foraging activities of Amegilla , on newly blooming flowers of Vigna unguiculata (L.) Walp., 1843 (Fabales: Fabaceae) were recorded during five consecutive days in 2021 and 2022 planting campaigns. Plants were divided into untreated plots and plots treated using the synthetic insecticide Parastar (l p.c..ha-1) or 10%, 20% and 30% aqueous leaf extracts of Calotropis procera (Aiton) Aiton, 1811 (Gentianales: Apocynaceae), Eucalyptus camaldulensis Dehnh., 1832 (Myrtales: Myrtaceae) and Tithonia diversifolia (Hemsley) Gray, 1883 (Asterales: Asteraceae) respectively. Among 8,987 insects collected (48.9% in 2021), Amegila calens Le Peletier. 1841 with stockier foragers (2021 campaign: 2.2% of the total collection, entomophily F A. calens =4.5%; 2022 campaign: 0.7%, F A. calens =1.3%; pooled campaigns: 2.9%, F A. calens =2.9%) and Amegilla sp. with slender foragers (2021: 3.8%, F Amegilla sp.=7.7%; 2022: no data) were recorded. Foragers started activity from 6 a.m. and stopped foraging before noon, with a peak of activity in 8 to 9 a.m. time slot for A. calens and 10 to 11 a.m. time slot for Amegilla sp.. During the five consecutive days from the first blooming day of the flowers, 598 visits (89.8% in 2021 and 10.2% in 2022) were recorded with a peak of visits during the 3rd day and then declined until it stopped during the 5th day. Treatments including the synthetic insecticide (which was the most repellent to the wild bees), did not significantly reduce the frequency of visits. But 20% aqueous extract of Ca. procera showed a significant increased of the mean duration of visits of the bees, compare to the results recorded in Parastar-treated plots. Therefore, the tested extracts, especially 20% aqueous leaves extract of Ca. procera may be recommended to control field insect pests and for preservation of foraging activities of Amegilla genus.
... It is for example the case of aqueous leave extracts of the fake Kinkeliba Cassia occidentalis L. (=Senna occidentalis (L.) Link, 1829) (Fabales: Fabaceae), the apple tree of Sodom Calotropis procera (Aiton) W. T. Aiton, 1811 (Gentianales: Apocynaceae), the red gum tree Eucalyptus camaldulensis Dehnh., 1832 (Myrtales: Myrtaceae), the Chan grass Hyptis suaveolens (L.) Poit., 1806 (Lamiales: Lamiaceae) and the Mexican sunflower Tithonia diversifolia (Hemsl.) A. Gray, 1883 (Asterales: Asteraceae) [11,14]. Floricultural entomofauna of V. unguiculata is well documented in cowpea producing countries in general and more particularly in Benin [34], in Ghana [35], in Nigeria [36], and in Cameroon [19,23,37]. These authors have reported the pollinating abilities of several insects including Ap. mellifera, Halictus sp. ...
... But our main objective was just to identify the main pollinating insect of the locality and to evaluate the effect of botanical pesticides on their behavior and not to study the entomofauna associated with the stages of plant development. [20,37]. In Benin, the most frequent insect on cowpea flowers was reported as X. olivacea [34] while in Nigeria, Ap. mellifera and X. olivacea predominated on V. unguiculata flowers [36] and in Ghana, Ap. mellifera and Halictus sp. ...
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