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Insect Choice and Floral Size Dimorphism: Sexual Selection or Natural Selection?

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Abstract

In considerations of sexual floral size dimorphism, there is a conflict between sexual selection theory, which predicts that larger floral displays attract more pollinators, and optimality theory—particularly the ideal free distribution—which predict that pollinators' visits should match nutritional rewards. As an alternate explanation of this dimorphism, Müller reported that pollinators tend to visit larger male flowers before visiting smaller female flowers, thereby promoting effective pollination. To investigate optimality predictions, I offered pollinators a choice between smaller, less numerous, but more rewarding flowers; and larger, more numerous, but less rewarding flowers. Foragers initially favored the larger and more numerous flowers, but rapidly shifted preferences to conform with the predictions of the ideal free distribution. To test Müller's hypothesis, I offered pollinators choices between larger and smaller corollas of equal caloric reward. Results showed that although pollinators tended to visit larger corollas first, they did not visit them more often. These experiments highlight the need for further investigation into the tradeoff between natural and sexual selection, and their respective influences in pollination ecology.
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ABSTRACT%6*
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$?*('%"&
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*DA,44/*,44,*,44+GA$
'EF,4.4=I,451=,45+=C
",454G'#%"@'
(#(AB7,45/*#=B,45,=
B,451GF
'%*7
A ,44+G"
7'*$88$
%688#?'('" #%
'%?'%
 ADC,45-=J(",45+=B,450=
$,455=#,454= ,44,=#"
,44,=C,44,=",44,*,44-=EC(,441=
,443=B(D(,440=",44.G"D
7CADC
,4.+G"*#%'>
'%(67A
B,451=',450=)(",454=D
",44+=,440=B(D(,440*,44.G"
($(6
'$(#%'%9
?"BC*%
'$$#($*#(#$
"67*7$(
%$$%%"
%'%9?*
'%$$(A@&G<'$
%%*""*C%$$
#($AE(C"
,4..=E(C,4.5=J#',441G*%"
''7%6A'%G*
#(#(%
67%@&"
##(%$@&"
E'''('$$7
AD#"-//-G*'7'
*7#'
*$'>(<*6
%AB,4+0=,433=,435=&E7$
,45/=,4.0=#",44,=:,441G"*
(7%88F<67*#
(<88<7*'%
6A:$",404=,435=E(C",4..=
,4.3*,451=C#,451=#",444G"
(*7'$$$#
%6''7%
(67A7,555G"&(*
'$%(*79@&
H&7677$$*
$*7%#'#(*7
%#KHA7,504*,/-G"
&7'(
6"&?7#8
'' **#67"* 7(
''%7*"&
#%7'$
%$67A$'EF,4.4=
",450=EC,45.G*'7%
767#7$A
,44/G"*%67
<7*'*AJ
:(,4.,=B7,45/*#=B",45+=L$",453= 
,44,G"&7(%
?*%
*#%'$"
:;A,5.1G<#(*H>(*H
$%('$#%'$
"$$6<'
<'%'$*8$
67?**8$67%*
"
&'%##('"
*$ #$'" 
%*%$$ ?
'#%( A"-//,G"*(#
$ $(*'(*
$?7"
*'9$
((morning (Young and Stanton 1990; Delph and Lively 1992), and I
have unpublished experiments that support this. Such a decay suggests an intriguing third
hypothesis, that pollinators are choosing flowers in sequence only until they have acquired
sufficient residual pollen load to insure continued fertilization. If so, this points to a more
complicated situation, in that pollinators may be selected to maximize food crops for future
generations.
7#F:;9>(*$
%%7%$M($
*?(*H(#$%
HA7,555G")(D##A,4..G'7
:;9(*%79"79
'#*7'*$#
>("
79H#HC*
'$*% $
A"@G"
$%''#7*
$(C("%(C$>##7
$$'*7$
(*'('$7'
#'9?"
&''%%
6*#7
("
%7'$$#"&?
%$%@&"&
%$%:;9>
("
:&@
!(%,443*%7'%86($
###AB#"G776%
,0A#()"G"&7%7
$*(%$8# C"&
7)%(*)(0/%
%'(%)"&#7$$7
(,-$*$$(87"
&67%'(5
"&(7$
'< "  +/N%$
(0*7C(
*#>$"
&%(#$$#7
#%?'"77
$/"0($7#F0/0/
7#('"D677*(7
69CA""*78#$#7%
$G?77%OA%*"*D#*
"" "G*%9?"
,
677'%%%7
H$HA""*#'%$G*%
7"
?'*,-677%
7*%(+/N%=
7*,5677%%"
7*677$
7/"/,%?##'"&,-
67*7'*7#$$%,0
#'*7,5$677
#$$%'(,0#'*$
67'%%"
'*5677*
$+/N%*7,-%8677
7"7%
''*5677
#$$%,0#'*7,-$
677#$$%'(*
$67'8%
"
#F67#$%*
67$#'7("
###'67#(%$(#'%
$%,0#'"
B%('#%
$788*$*(*$%
*"88###''7($(*'
%(("*%'$###*
%$'*(%
(" %'(*#
$'%7(7'#*7
$%##%#C$("
&%%?%%
7$H?H%#"
'%77$$*'7
%"@#'7#$(
$*$,/#'*7C#
3"0"
&#%'$7#*
#'7$(%"2
$7$%'%
#'"
-
?*6767
###7F%67*77
67"%67
%8*#
$%$#(-/N*+/N*3/N"
E*7'*'''67
%*#(676
A,433=!,4.,=E(C,4.5*,45-=D$
,4.4=#",45,=&",45-*,454=(:
,45+=B,450G"B677<(
$*7#%>('67
6"*7
%"67769C?7
77%*$*7
?'"&%7$
#%767#'"&$
'6%#(67*7('%
67'(1/"$67#
(7?"'(,0/"/,%
0/0/77F#%
67"&67<##(
"
#'*9'7
#F*7677'?"
)&
,
&%#AG%$7#
'$7*#($$(*
>'#7AJ#'
,441G"%###'$7$7
*#%'>
AE,45,=)'(,44-=$,440G"?
'*'7#'
HHA""*#7$G$*##(
%7$
 ,- xPPPP 
,-xQ,5A/"0xGR,//NS0.N*
7x%F$*/"0
%$%HFH
A$*7$G$"
(*#%'$
'7###(
 5xPPPPPP
5xQ,-A/"11xGR,//NS3.N"
$",-7$%'$*
77"E7$
#(-$%?"@>7
$?(<%AT/"/0G"
-
E9#%%$7

absolute preference = total visits to larger !ower ÷ total visits to both
!owers;
>%7
sequence preference = &rst visits to larger !ower ÷ &rst visits to both
!owers.
B$$"1"
#%*>%7(
$?($#%AT/"/0G#+/N
3/N%*8$?-/N"
&%%77-7
$A0/NG"
@
#*%
$%"
?*#(%$*
'#(7'67"D
%<$7*7'*>C(
F'('$
%("&?#%'7$
$#(7%*7
%%$*"
7'%%
$ "&*<>
$% "B%
 *77'#'%-,N
$*7$%?*
#"B%67#*#
7''%+/N$*
$%*#?"
&%$("%
($%(8'*
C%%7=(
 (''%7"
<$7$%*#<$6
#<
M" $677
%7'67"%78$
$*'(67$$
%$*(
=(7C$#?
*867A%,45+G"&(
$#%("
####('$#(
*7%#"&
##$'#>(%%*%7
$%%*%$$(
*777%$
'"
*(#(
(#7#%
"7>%*7'*'(
*7#%(
$#%A0/NG"&%
>("
$%:;9(*
%A$77G*
6767$*'%
A (,454G"<
%'*#>
($"(7#$
$#%*#
'*$*($*67
$(67#"&7
$##(7'
67#%67*%$7
%*8$"&7##
* A:(C
U%-//1G"
&#7
*'7A,5.,G'
%"7A #,443G'$$
%"&
*7%$%
(#77"
C7$$'JF"B7 "#
%'#$$"
)& &&
#*!""A,443G")# $(%
" B46*-1810"
L$*!"*>'*&"&* "A,453G"E#(*6
#)"@$70*
11-8115"
*"!"(*""A,454G"(
("'43*-/+8-,4"
*"A,44,G"( 
A G"""14*,538,4,"
*"!"A,44+G"*
($#('!" " ""144*,8,+4"
*&")"*:")"A,44,G"'
(%($"
A:'G"$(72*4418,//1
BC*""A,4+5G"$($(
%67$"E")E")""5*,138,14"
B7*J""A,45/G":(%#(%67
%!A"GD
AG"'34*+3.8+.+"
B7*J""A,45/#G"'%(67$" "'""
("11*,0814"
B* !" " A,45,G" E %$$   '%(" "
"118*0.-80.."
B*"A,450G"@%%67"E""")"B224*--18
-30"
B*"*)%#'*)"**)"8 "D(*"A,45+G"E
%%%67%#"
@$65*-5.8-4+"
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'49*54844"
B(*"B"D(*"A,44.G"&(
'I" ""149*0408044"
B*""A,4+0G"&6%'(#$%
%'(#'(" %
#7'(
#"!""B"21*08,-"
#*""A,454G"6%%
(" "!"B"76*.1/8.15"
#*""*D*":"*E*:"2"*)(*" ":*"!"
A,44,G"%(67
7$$$"'45*,+058,+3."
*"!"DC*!")"A,45-G"$'
$?%6(%%C7"$(63*
,50.8,5./"
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E7*:"A,444G"67'67
%(8?'" "B"83*.1853"
*!"J"*"A,443G"<%67#
'7*"@$
104*+/480,3"
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D(8%$$#77C7
6@$51*.4851"
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15*10481.5"
7*"A,504G"@@$%#(:%"
!":(*)"
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:(*)"
7*"A,555G"&<%7E%
"'(%$E*$"
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@C72*,3,8,.-"
C*2":"A,44,G"&<%6('
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"5*1./815+"
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,+54"
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,5+8,4-"
*" ",405"&&(%"'*7
UC"
$*""*""A,455G"'
6%$#A@G" "!"B"
75*,1.,8,1.."
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67$"!*"")*"!"A"G*
#C%EB$("?W 
*7UC*"13/81.-"
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""5*,348,.5"
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%$#67" ""100*5084."
*J"A,440G""E"" "
" 92*,-+.8,-0/"
*)""**:"B"A,44/G" '%($
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"""4*00480.-"
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'6$(
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-4,"
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%$$<(%)(("@$61*
-+48-01"
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,./"
*B"A,4.3G"&%$$%'###"
":"46*,/08,-5"
*B"A,451G"%$$$"!*"")*"
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*7UC*",5.8-,+"
(*!""A,415G"79(%
##(*$%" ""72*+,38+11"
!*""A,4.,G"$#$%
"171*-/18-/0"
!*""A,4..G" #(" ""
111*13081.,"
J*":":(*")"A,4.,G":%6
(&"'25*3048335"
J(*I"@""*)C* "!"*B#*""'*""A,45+G"
<#76$(*
'"7E("98*
0,080-4"
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'"43*,..8-,3"
)(*""*D*":"*E*:"2"* '*!":* ""
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%%" ""134*+/48+11"
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:$*""* *":"*J*""!"*B*""!"A,404G"
B'$67"!"""52*,//-8,//+"
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#'?"'47*-08
10"
:(C*&"U%*:"A-//1G"<'
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$(70*.,8.3"
E*!":",45,"B##'
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EC(*""A,441G"'%677<'
#(HH67$(74*--008--3/"
EC*"B"A,45.G"$67*%*" "
'""("18*+/48+1/"
E(C*""A,4.5G"@%$$'%###
#76"&"E"B"13*.-845"
E(C*""A,45-G"$$###%%
6"!"%V"60*+,.8+-5"
E(C*""*E*""'*")"A,4..G"@%$$
''7%("I"'"B"52*,1.8,0+"
I*""A,451G""
305*./38./."
(*:"!"* ""A,44/G"&(#%%
X$%$*E(YA
%(G"'44*1/081,+"
*!"*&*!"C*)"A-//,G"2%$$
###67<6$
#'"E"" ""98*1545814/1
*:")"*U$*"!"**""$$*!":"A,44,G"
>%6'%%
%7*')"
'45*-358-5/"
*:")"* *&"8)"*7(*)""*U$*"!"A,44-G"
$?%"D(*"
A"G*$('%E"W*
7UC*"3-84/"
* ""B*""A,451G":*%*
"*)"A"G*EB$("
E*7UC*",,/8,+4"
*"*)""A,45+G"@%?
%%"$(65*,/418,,/+"
&*!""E7$*""A,45/G"E('*
7*#'7%'
"@$46*358.+"
&*!""*:*D"E"*E7$*""A,45-G"B'%
###% 2"A G"@$
54*1-38113"
&*!""*:J*:" "*:"B"A,454G"&
% 
%'"$(70*,/3,8,/35"
D$*"*B"A,4.4G"&%$$'%
###'6("!"
"E("134*,,18,,."
D#* "!"**"*D":"A-//-G"(#
$677$%$$#" B'"
63*,83
D*:""A,4.4G"" ""113*...8.4/"
D*:""A,44/G""&5*-,/8
-,+"
D*:""A,44,G"*
($("'""5*3485."
D*:""A,44+G"'''7"
""144*,1814"
D*:""C*B"!"A,4.+G" '$%6
( ()" ":""92*+.80."
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... That is, Bell's hypothesis in insufficient to explain the small reward offered by larger male flowers, but it can be explained by the sequence hypothesis. However, few studies on the sequence hypothesis have considered both advertisement traits and reward traits simultaneously 5,21 , which makes it difficult to evaluate which of the two hypotheses is more likely to explain sexual dimorphism in floral traits. ...
... In this experiment, we could not control the floral scent, but the stronger scent of male flowers compared with female flowers 24 suggests that floral scent might serve as an advertisement trait for pollinators in open air conditions. Prior experimental research has demonstrated partial support for the sequence hypothesis in another plant species, Hibiscus syriacus L., in which pollinators visit larger flowers first in the field 21 . ...
Article
Full-text available
Sexual dimorphism is a pervasive form of variation within species. Understanding how and why sexual dimorphism evolves would contribute to elucidating the mechanisms underlying the diversification of traits. In flowering plants, pollinators are considered a driver of sexual dimorphism when they affect female and male plant fitness in distinct ways. Here, we found that flowers appear to manipulate the behavior of pollinators using sexually dimorphic traits in the dioecious tree Eurya japonica. In this plant, female flowers present a higher-quality reward for pollinators, whereas male flowers have a more conspicuous appearance. Plants benefit by inducing pollinators to carry pollen from male to female flowers, and their sexual dimorphism might thus facilitate pollen movement through pollinator behavior. In two-choice experiments, pollinators frequently moved from male to female flowers, whereas computer simulation suggested that sexually dimorphic traits would evolve if pollinators changed behavior depending on the traits of the flowers they had just visited. These results suggest that the floral traits affecting the visiting order of pollinators have evolved in plants. Using E. japonica, we theoretically show that the induction of sequential behavior in pollinators might be crucial to the evolution of sexual dimorphism in flowers, and our experiments support these findings.
... This may be caused by the differences in the composition of floral scent or the by the fact that male flowers may release more scent compounds. In this experiment, honeybees and bumblebees were the main pollinators; they have great learning abilities and might distinguish floral scent patterns and might transfer these to other corresponding patterns [18,52]. ...
Article
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Eurya plants are usually dioecious or subdioecious with small fragrant flowers. Here, we investigate the floral scent components of the subdioecious species Eurya loquaiana Dunn and how floral scent affects pollinators. Headspace solid-phase microextraction–gas chromatography–mass spectrometry (HS-SPME-GC-MS) was used to compare the floral scents of male, female, and hermaphrodite flowers. We also test whether differences in floral scent affect the foraging behaviors of pollinators and describe the flower morphological traits of the three sexes. Twenty-eight floral scent compounds were tentatively identified, and four monoterpenoids were tentatively identified as the most abundant compounds: linalool oxide (pyranoid), linalool, lilac aldehyde, and linalool oxide (furanoid). There were floral scent differences among the sex types, and male flowers were more attractive to pollinators in the wild, even when visual factors were excluded, indicating that pollinators likely distinguish sexual differences by floral scent. In the competition for pollinators, the advantage that male flowers have over female and hermaphrodite flowers can likely be accounted for the differences in floral scent and display size.
... The abundance of male flowers (or equivalents) translates directly into an abundance of produced pollen. In addition, in animal-pollinated plants, the flower abundance attracts pollinators and increases pollen transfer(Glaettli & Barrett, 2008).Also, floral characters (floral display, floral colour, nectar and scent) may be important for male success because they influence visitation frequency and foraging time of pollinators, and indirectly pollen dispersal between plants(Abraham, 2005;Huang et al., 2006;Yan et al., 2016). However, flower traits are often difficult to quantify, especially if there are many candidate parents within a study plot or if a study species is characterized by large dimensions, as many trees. ...
Article
Individual differences in male reproductive success drive genetic drift and natural selection, altering genetic variation and phenotypic trait distributions in future generations. Therefore, identifying the determinants of reproductive success is important for understanding the ecology and evolution of plants. Here, based on the spatially‐explicit mating model (the neighborhood model), we develop a hierarchical probability model that links co‐dominant genotypes of offspring and candidate parents with phenotypic determinants of male reproductive success. The model accounts for pollen dispersal, genotyping errors as well as individual variation in selfing, pollen immigration, and differentiation of immigrant pollen pools. Unlike the classic neighborhood model approach, our approach is specially designed to account for excessive variation (overdispersion) in male fecundity. We implemented a Bayesian estimation method (the Windows computer program available at https://www.ukw.edu.pl/pracownicy/plik/igor_chybicki/1806/) that, among others, allows for selecting phenotypic variables important for male fecundity and assessing the fraction of variance in fecundity (R2) explained by selected variables. Simulations showed that our method outperforms both the classic neighborhood model and the two‐step approach, where fecundities and the effects of phenotypic variables are estimated separately. The analysis of two data examples showed that in wind‐pollinated trees, male fecundity depends on both the amount of produced pollen and the ability to pollen spread. However, despite that the tree size was positively correlated with male fecundity, it explained only a fraction of the total variance in fecundity, indicating the presence of additional factors. Finally, case studies highlighted the importance of accounting for pollen dispersal in the estimation of fecundity determinants.
... Other studies have also found that flower size affects the behaviour of pollinators. Abraham found that naïve bumblebees (Bombus spp.) visited all of the flowers of 15 Althea (Hibiscus syriacus L.) plants in equal numbers but visited the larger flowers first [32]. It was suggested that this was an optimal foraging strategy, as larger flowers were likely to produce more nectar. ...
Article
Full-text available
Cytoplasmically male-sterile (CMS) carrot cultivars suffer from low pollination rates. In this study, insect visitation varied more than eightfold between 17 CMS carrot cultivars in a field-based cultivar evaluation trial. The visitation rates of honey bees, nectar scarabs, muscoid flies, and wasps each significantly differed among these cultivars. No significant difference in visitation rates was observed among cultivars of different CMS type (brown-anther or petaloid) or flower colour, but cultivars of Berlicumer root type had significantly higher insect visitation rates than Nantes. Six cultivars were further compared in regard to selected umbel traits: as umbel diameter increased, so did the visitation of soldier beetles, while that of honey bees decreased. Finally, nectar of these six cultivars was analysed for sugar content, which revealed monosaccharides to be the most common sugars in all. There was high variation in the levels of sugars from individual umbellets but no significant difference in nectar sugar composition among cultivars, suggesting that nectar sugar composition is of minor importance regarding pollinator attraction to hybrid CMS carrot umbels.
... The associative learning of reward availability with floral size enables experienced bees to choose a rewarding floral size when initially preferred flowers become unrewarding (Blarer et al. 2002). Furthermore, my data show that bees' initial size preferences quickly disappear when reward size is not different among flower sizes, which corroborates the findings of Abraham (2005). With reward-based foraging behaviour, experienced bees show no preferences for flower sizes that have the same amount of reward. ...
Thesis
Full-text available
The flower morphology and reward availability of animal pollinated plants are intrinsically related to the foraging behaviour and preferences of their pollinators. However, it is often difficult to test how pollinator preferences may have helped to shape floral morphology because the morphology of many animal pollinated flowers is an adaptive compromise to optimise both male and female function. This may be overcome by studying the foraging decisions of pollinators in relation to flower morphology of species with unisexual flowers. The inherent difficulty of studying diclinous species is that in nearly all of these species the flowers of different sexes do not only differ in morphology, but also in reward type: male flowers offer pollen and possibly nectar, while female flowers offer nectar only. Solanum is an ideal genus to investigate evolutionary links between pollinators and flower morphology for two reasons. First, it demonstrates a large variation in sex system with hermaphrodite, andromonoecious and dioecious species. The diclinous species of Solanum have evolved sexual dimorphisms involving floral size and the number of flowers per inflorescence. This variation allows the evaluation of floral morphology in a phylogenetically informed way. Second, pollen is the only reward, and is present in apparently equal amounts in both male and hermaphrodite/female flowers. This allows the investigation of sexual floral morphology in the absence of differences in reward type and amount. The genus Solanum is further suitable for such investigations because it relies for pollination on a relatively small number of buzz-pollinating bee species. The main objective of this study was to examine how sexual dimorphisms in floral display and reward availability influence bee foraging behaviour, as this could lead to an understanding of the evolution of floral traits in association with changes in sex systems in the Australian members of Solanum subgenus Leptostemonum. To investigate this, buzz pollinating bees were tested for their responses to dimorphisms in three floral traits: corolla size, flower number and style length. Although Amegilla murrayensis had an initial preference for larger flower size, this preference quickly disappeared in the absence of differences in rewards among flowers. Clusters of flowers were more attractive than solitary flowers, even when the clustered flowers were smaller in size. In another experiment, Amegilla chlorocyanea showed no differences in the number of buzzes and time spent on each buzz between two floral morphs of andromonoecious Solanum elaeagnifolium that differed in their style length. Furthermore, foraging decisions by individual bees were analysed in relation to variation in pollen availability. Amegilla chlorocyanea showed no difference in the total number of times they buzzed pollen-full and pollen-empty flowers before they left the patch. However significant differences observed between first and second visits to flowers indicated that bees could perceive recent visitation by a bee and adjust their visitation behaviour. Since an overall lack of support was found for the evolution of floral sexual dimorphism as a direct response to bee foraging preferences, a phylogenetic analysis was performed to investigate other possible explanatory models for the evolution of floral dimorphism in the diclinous species of Solanum. First, molecular phylogeny was inferred based on three gene region sequences of 71 Australian members of Solanum subgenus Leptostemonum. The analysis showed that the evolution of andromonoecy from hermaphroditism is most likely preceded by the evolution of large fruit, and thus the selection for large fruit size is the main driving force for the evolution and maintenance of andromonoecy in this group.
... The rates of pollinator visitations may vary with different floral features, e.g. flower color ( Staton 1987), size ( Conner & Rush 1996, Abraham 2005), nectar production ( Mitchell 1994) and display morph ( de Jong & Klinkhamer 1994, Thompson 2001) etc. Insect visitation may also be sconsiderably influenced by the spatial and temporal arrangement of floral traits ( Campbell et al. 1997, Mitchell et al. 2004), and this interaction is likely to be particularly important since different pollinators often show marked spatialtemporal variation in abundance ( Herrera 1988) and pollination effectiveness ( Wilson & Thomson 1991). ...
Article
Seed production of alfalfa (Medicago sativa L.) is important in determining the effective distribution of new cultivars to farmers. However, little is known about the biodiversity and their community function of native wild pollinators of alfalfa in agronomic systems. We investigated the biodiversity of insects which visited alfalfa flowers and their temporal foraging characters in Hexi Corridor, China. A high biodiversity of insect visitors was discovered, 20 insect taxa in all, including 13 species of Hymenoptera, 3 species of Coleóptera, 3 species of Lepidoptera and 1 species of Diptera. Three native bee species, Andrena squamata, Anthophora melanognatha and Megachile abluita, were validated as the principal pollinators. They showed significant variations in tripping mode and their diurnal distribution patterns. Our results indicated that the native wild bees are diverse and they complement each other. This means they have developed a more complex system for the pollination of alfalfa than has been previously found out.
... Pollen limitation combined with pollinator preference exerts selection pressures on flowering traits. Many studies have found significant relationships between attraction of pollinators and floral display, both in field studies (Willson 1979;Ohashi & Yahara 1998) and in manipulation experiments (Andersson 1996;Abraham 2005) for a wide range of plant families (e.g. Asteraceae, Malvaceae). ...
Article
The plant life cycle is often affected by animal–plant interactions. In insect-pollinated plants, interaction with pollinators is very important. When pollen transfer due to a lower abundance of pollinators limits seed production, selection pressures on plant traits related to plant attraction to pollinators might occur, e.g. on flowering phenology, height or number of flowerheads. Landscape changes (e.g. habitat fragmentation or changed habitat conditions) may cause plant–pollinator systems to lose balance and consequently affect population dynamics of many plant species. We studied the relationship between measured plant traits, environmental variables and pollinator preferences in Scorzonera hispanica (Asteraceae), a rare perennial, allogamous herb of open grasslands. We estimated the pollen limitation by comparing seed set of supplemental-pollinated plants with that of open-pollinated ones. Pollinators selected plants based on position within the locality (isolated plants close to trees) rather than on their traits. In spite of a high proportion of undeveloped seeds on the plants, we demonstrated that they are not pollen limited. Instead, seed set and weight of seeds was correlated with plant size traits (height and flowerhead number), with larger plants producing more and larger seeds. This suggests that the studied plants are likely resource limited. Overall, the results suggest that pollinators are not a selection factor in this system, in contrast to studies on various plant species, including self-compatible species of the Asteraceae. The lack of any effect of pollinators in the system may be caused by a strong negative effect of ungulate herbivores, which could play a decisive role in functioning of the system.
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In many plant species, larger flowers offer larger or higher quality rewards to visitors, and flower visitors preferentially visit these larger, more rewarding flowers. Limited evidence suggests that plants in which flower size provides more reliable information about reward value have higher reproductive success than plants in which flower size offers less reliable information. We tested whether bumblebees could learn to respond to flower size when it was an informative reward cue while simultaneously learning not to respond to it when it was not informative. We also tested whether bees would develop a preference for a flower type that provided reward cues over one that did not. Bees were allowed to forage on an artificial array containing 2 flower types, each with a unique color and scent. In the informative flower type, large flowers contained sucrose rewards, whereas small flowers contained quinine solution (a deterrent). In the uninformative flower type, both flower sizes were equally likely to contain sucrose or quinine. Bees learned to prefer large flowers in the informative flower type and to be indifferent to flower size in the uninformative type. However, contrary to expectations, bees did not develop a preference for the informative type. These results suggest that although bees may benefit from floral cues that give information about rewards, plants offering these cues may not receive more flower visits than plants that do not.
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We studied the relationship between inflorescence size and male fitness in the andromonoecious lily Zigadenus paniculatus, using experimentally manipulated inflorescences to eliminate possible correlations between flower number, resource availability, and other floral traits. Allozyme markers were used to determine the siring success of large versus small plants in 14 arrays of plants, each array containing five large and five small plants. The inflorescence size of small plants was held constant both within and among arrays; the size of large plants was held constant within an array but was varied among arrays. Large plants sired more than half the seeds in 12 of the 14 arrays, and significantly more than half in six of these 12. However, in eight of the arrays, large plants sired significantly fewer seeds than expected on the basis of their size advantage. Furthermore, there was no significant relationship between relative size and relative siring success in comparisons among arrays. A maximum-likelihood model estimated that 28% of seeds were sired by imported pollen, with 95% confidence limits of 13% and 50%. Within these limits, high import rates tended to mask the relative success of large plants in several arrays. These results suggest that the evolution of inflorescence size in Z. paniculatus is at least partly driven by selection for increased male success, assuming genetic variation for flower number. However, the data also support a growing body of evidence that estimates of male fitness in plants can be highly variable. We discuss the sources of this variability and the possible effects of inflorescence design on the relationship between inflorescence size and fitness.