Sex Plant Reprod (1997) 10:293–299
O R I G I N A L PA P E R
&roles:Sandra L. Davis
Stamens are not essential as an attractant for pollinators in females
of cryptically dioecious Thalictrum pubescens Pursch. (Ranunculaceae)
&misc:Received: 4 February 1997 / Revision accepted: 20 May 1997
&p.1:Abstract It has been hypothesized that females of some
dioecious species maintain stamens that produce sterile
pollen as a means of attracting pollinators to promote
greater seed set. However, this hypothesis has rarely
been tested. This paper examines the role of pollinators
in the maintenance of sterile stamens in the cryptically
dioecious species, Thalictrum pubescens. Wind was
found to contribute to pollination; branches of female T.
pubescens enclosed in cheesecloth to exclude insects but
not wind were still able to set seed. Therefore, females
may not need the stamens for pollination. In 1994, in-
sects were found to discriminate against emasculated fe-
male flowers, but this discrimination did not lead to a
significant difference in either the amount of pollen re-
ceived or seed set. In 1995, emasculation was combined
with the addition of pollen to determine if emasculated
females were pollen limited. No significant effect of pol-
len addition or emasculation was found. Emasculated
branches in the pollen-addition treatment appeared to
have slightly lower seed set than non-emasculated
branches that received pollen, indicating that the slight
reduction in seed set was caused by emasculation rather
than pollen limitation. Since stamens do not appear to be
maintained as pollinator attractants, other explanations,
such as genetic constraints, must be investigated.
&kwd:Key words Thalictrum pubescens · Cryptic dioecy ·
Pollinator attractants · Wind pollination · Genetic
constraints&bdy:
Introduction
Plant species exhibit a variety of breeding systems rang-
ing from hermaphroditism, in which all individuals pro-
duce perfect flowers with both male and female function,
to dioecy, in which individuals produce either unisexual
S. L. Davis
Department of Biology, Indiana University, Bloomington,
IN 47401, USA
Fax +1–812–855–6705; e-mail: saldavis@ucs.indiana.edu&/fn-block:
© Springer-Verlag 1997
male flowers or unisexual female flowers. Between the
two extremes are several intermediates, such as gyn-
odioecy, subdioecy, polygamodioecy, etc., in which pop-
ulations may consist of a mixture of different types of in-
dividuals, and/or individuals may produce a mixture of
different types of flowers. Because 95% of all plant spe-
cies are hermaphroditic (Yamplosky and Yamplosky
1922), this breeding system has typically been consid-
ered the ancestral state to most others. The evolution of
dimorphic breeding systems (dioecy, in particular) from
hermaphroditism has been modelled extensively during
the past two decades (Charnov et al. 1976; Charlesworth
and Charlesworth 1978; Bawa 1980; Givnish 1980;
Bawa and Beach 1981; Beach, 1981; Lloyd 1982). One
conclusion that can be reached from these studies is that
the interaction between genetic effects and pollinator be-
havior can have a strong effect on the evolution of plant
breeding systems (Willson 1979; Bawa 1980). For exam-
ple, inbreeding depression is thought to selectively favor
the evolution of dioecy and other obligately outcrossing
breeding systems. However, Darwin (1877) pointed out
that cross-pollination, either through biotic or abiotic
factors, must be ensured before dioecy can evolve.
Achieving cross-pollination may be a particular problem
in insect-pollinated species in which pollen is an impor-
tant reward and/or in which nectar is absent, since fe-
males of most dioecious species lack stamens and pollen.
Consequently, pollinator discrimination against less re-
warding females may constrain the evolution of separate
sexes even when other selective factors are present (Bell
et al. 1984; Kay et al. 1984; Bierzychudek 1987). This
constraint can then affect the evolution of other traits
within the plant or the correlations between traits. For
example, Charlesworth (1993) recently suggested that
the alleviation or the removal of the constraint of pollina-
tor discrimination in those species pollinated by wind or
non-specialized insects may account for the correlation
between dioecy and these modes of pollination. This in
turn accounts for the correlation between dioecy and oth-
er traits associated with these modes of pollination, such
as small inconspicuous flowers.
294
There are several ways in which dioecious species
could circumvent the problem of insect discrimination
against less-rewarding females. Melampy and Hayworth
(1980) examined some possible solutions in several nec-
tarless plants, including wind pollination and shifts in
flowering phenology. Another solution to the problem of
pollinator discrimination is the production of stamens
containing non-viable pollen by females. The females ap-
pear hermaphroditic but are functionally unisexual. The
stamens may either provide the pollen as a reward for
pollinators (Kevan and Lack 1985; Anderson and Symon
1989) or may “trick” insects into visiting the less-reward-
ing sex (Schlessman et al. 1990). This type of dioecious
breeding system, in which one or both of the unisexual
morphs mimics perfect, hermaphroditic flowers by pro-
ducing non-functional organs of the opposite sex, is
termed cryptic dioecy (Mayer and Charlesworth 1991).
Although it is generally assumed that selection for in-
creased insect visitation maintains stamen and/or pollen
production as an attractant for pollinators (the pollinator-
attraction hypothesis), there are at least two other hy-
potheses that could account for this preservation (Ander-
son and Symon 1989; Mayer and Charlesworth 1991):
(1) there are strong, positive genetic correlations be-
tween anther and pollen production in males and their
production in females that constrain the evolution of sex-
ual dimorphism (the genetic-constraint hypothesis) and
(2) there is no selective benefit to their removal because
there is no “cost” to females for their production and
they are retained as vestiges of the ancestral state (the
ancestry hypothesis). Because the focus of studies on
cryptic dioecy until now has been on the pollinator-at-
traction hypothesis, researchers have primarily investi-
gated the pollination biology of such species (Kaplan
and Mulcahy 1971; Clay and Ellstrand 1981; Kevan and
Lack 1985; Kevan et al. 1985, 1990; Schlessman et al.
1990; Hoffman 1992) and ignored the role of the latter
two hypotheses. Since most cryptically dioecious species
are morphologically androdioecious (having male indi-
viduals that produce staminate flowers and female indi-
viduals that produce apparently perfect flowers) with
pollen as the main food reward for pollinators (Mayer
and Charlesworth 1991), the pollinator-attraction hy-
pothesis is an appealing explanation for the maintenance
of this breeding system. Even though the pollinator-at-
traction hypothesis may explain the presence of stamens
in the majority of cryptically dioecious species, it may
not provide the full story in all cases. For example, Sch-
lessman et al. (1990) found that females in Polyscias
pancheri present sterile pollen several days before the
stigmas become receptive. Hence, pollinator visits to
flowers displaying sterile pollen could not affect fertili-
zation at that time. The authors suggest that the pollina-
tors “learn” the location of the females during these vis-
its and, consequently, are more likely to visit them when
the stigmas do become receptive. However, the hypothe-
sis was not tested, and it is therefore not clear whether
sterile pollen production is retained due to this hypothe-
sis or to other factors.
In this study, I examined the role of insect pollinators
in the maintenance of cryptic dioecy (the pollinator-at-
traction hypothesis) in tall meadow rue, Thalictrum pu-
bescens Pursch. (also known as Thalictrum polygamum
Muhl., Park 1992). Females of this species produce full-
size stamens which contain sterile pollen that lack germi-
nation pores. Therefore, T. pubescens, like the majority of
cryptically dioecious species, is functionally dioecious
but appears androdioecious (Kaplan and Mulcahy 1971).
This is a particularly interesting species in which to study
the factors maintaining cryptic dioecy, since T. pubescens
is at least partially pollinated by wind (Kaplan and Mu-
lcahy 1971; Melampy and Hayworth 1980) and may not
need the sterile stamens for successful fertilization.
A primary assumption of the pollinator-attraction hy-
pothesis is that, in the absence of stamens, female repro-
ductive effort is limited by the amount of pollen being
delivered to the stigmas by insects. If this selective factor
is absent in T. pubescens, the pollinator-attraction hy-
pothesis cannot explain the maintenance of the non-func-
tional stamens. Secondly, if the stamens in T. pubescens
are maintained in the females through pollinator attrac-
tion, there must be a difference in pollinator response
that leads to a selective disadvantage toward females
with smaller or fewer (to zero) stamens compared to
those with larger displays, with results such as fewer pol-
linations and lower seed set.
Materials and methods
Study organism
The genus Thalictrum shows a wide range of breeding systems
from hermaphroditism to complete dioecy, and species are polli-
nated by wind, insects, or both. Sex-determination in Thalictrum
spp. appears to be under nuclear control with males serving as the
heterogametic sex (reviewed in Meagher 1988). T. pubescens is a
summer-flowering perennial that grows in rich woods, low thick-
ets, swamps, wet meadows, and stream banks (Keener 1976). The
population used in this study is located in a small, unused, private-
ly owned field adjacent to Fort Hill State Historical Park near
Hillsboro, Ohio. Females of T. pubescens from the population near
Fort Hill produce pistillate flowers with an average of 11.7 pistils
with uniovulate ovaries and a wide range of zero to 15 stamens
(average=6.7), all containing inaperturate pollen. Males produce
only staminate flowers with an average of 34 stamens and contain
no vestigial pistils (Davis, unpublished data). T. pubescens lack
both petals and nectaries. Sepals are white, relatively small com-
pared to the stamens and pistils, and fall off soon after the flower
matures. The species is partially wind-pollinated (Boivin 1944;
Kaplan and Mulcahy 1971), with the primary insect visitors being
Syrphid flies. Because of these characteristics, Thalictrum species
provide an excellent system with which to explore the evolution of
cryptic dioecy and its maintenance.
Role of wind versus insect pollinators
An experiment to test whether wind can contribute to the pollina-
tion of females was conducted during the summer of 1995 by ex-
cluding either wind or wind and insects from female flowers. Two
branches were selected on each of 20 plants prior to flowering and
enclosed in either pollination bags (to exclude wind and insects)
or cheesecloth (to exclude insects only). An additional branch on
each plant was tagged as an open-pollinated control. At the end of

the season, the branches were collected and scored for fruit and
seed set. Seed set for all experiments was determined by counting
the number of mature seeds and aborted ovules produced and then
calculating the number of seeds per total ovules produced. If wind
is able to contribute to pollination, branches in the cloth-enclosed
group should have significantly higher seed set than the bagged
group. The seed set on the cloth-enclosed group is expected to be
an underestimate of the contribution of wind pollination, because
the cheesecloth undoubtedly prevents some pollen from reaching
the stigmas; therefore, the seed set of this group would not be ex-
pected to be as high as the open-pollinated controls.
Each pair of groups were compared with the Wilcoxon
matched pairs test. P values were adjusted to compensate for mul-
tiple tests by multiplying each by three, the number of tests per-
formed (all three possible pairwise tests were performed).
Emasculation experiment, 1994
Experimental manipulations were performed in the field to examine
the fitness consequences of the presence or absence of sterile sta-
mens in female flowers, independent of other correlated traits. Thir-
ty female plants were chosen at the beginning of the reproductive
season in July, 1994. Two similar branches on each female were
then chosen and randomly assigned either to the control or the treat-
ed group. All flowers on the treated branch were then “emasculat-
ed” by the removal of all sterile anthers. Choosing two branches on
the same plant allowed for the control of random factors, such as the
total number of flowers and location of the plant. Plants were fol-
lowed each day for the next week, and new flowers on the treated
branches were emasculated as they arose. Each branch was also ob-
served for a 45-min time period in the late morning, with the num-
ber and length of any insect visits recorded. Towards the end of the
week, one or two flowers of approximately the same age from each
branch were collected and preserved in 70% ETOH. Flowers were
taken back to Indiana University where several floral measurements
were then made using a micrometer under magnification of × 8.
Characters measured were stamen number, filament length, anther
length, pistil number, style length, ovary length, and sepal length
(except for anther and pistil number, measurements were made on
three structures per flower). Pollen deposition was also examined on
the flowers by mounting styles on slides and counting the number of
pollen grains under × 400 magnification. Styles were soaked in 8 N
NaOH for 24–48 h in order to soften the stigmatic tissue. The styles
were then soaked in distilled H2O for another 24–48 h. This re-
moved coloring from the stigmatic tissue while the pollen grains
were unaffected, making them easier to see. Because this procedure
most likely washed off some pollen grains, this is an underestimate
of the amount of pollen being received by means of both wind and
insects. At the end of the season, individual branches were collected
and scored for flower number, bud number, and seed set.
The data for both pollinator visits and pollen grains deposited
were highly skewed to the left, with many zeros. This skewness of
the data could not be improved by transformation, so differences
between the emasculated and control branches for pollinator visits
and the mean number of pollen grains deposited were tested using
the sign test. Seed set was transformed using the arcsine transfor-
mation (Sokal and Rohlf 1981), so that it approximated a normal
distribution, and then tested using a paired t-test.
In addition, a relationship between seed set and mean anther
number was tested for by a Pearson-product correlation using data
from the control branches only. The relationship between anther
number and seed set within the control branches should also re-
flect this same relationship in the rest of the plant and within other
plants in the population, since only one branch out of many was
emasculated on these plants.
Emasculation and pollen augmentation experiment, 1995
In order to separate the physical side effects of the emasculation
treatment and the possible reduction of pollination due to the treat-
ment, I performed hand-pollinations of emasculated females and
295
compared them to open-pollinated emasculated females. At the
beginning of the flowering season in the summer of 1995, four
branches on each of 30 plants growing at the Hillsboro site were
selected and each branch was randomly assigned to one of the four
possible combinations of two treatment/control groups: emasculat-
ed/non-emasculated and pollen addition/natural pollination (e.g.,
all the flowers on one branch were emasculated and hand pollinat-
ed, or non-emasculated and pollinated, etc.). Each of the 30 plants
received all four combinations of treatments in order to control for
inter-plant differences that might have arisen due to microsite vari-
ation, flower-number differences, etc. If the emasculation experi-
ment causes reduced seed set due to damage effects rather than re-
duced pollination, then the emasculated branches will have lower
seed set with both natural and hand pollination. On the other hand,
if emasculated branches have lower seed set due to reduced polli-
nation by insects, the addition of pollen by hand should increase
the seed set of emasculated branches. In addition, if pollen aug-
mentation increases the seed set of non-emasculated branches, this
would also indicate pollen limitation and support the hypothesis of
competition among females for pollinators.
Data were analyzed using a mixed-model three-way ANOVA.
Independent factors were the individual plant (random), the addi-
tion of pollen (fixed), and emasculation (fixed), with the denomi-
nator mean squares calculated according to Zar (1984). Because
there were no degrees of freedom left in the residual term of this
model, when the denominator mean square called for was the
mean square error, the mean square for the three-way interaction
was substituted. A significance test on the three-way interaction
could not be performed due to the lack of an appropriate denomi-
nator mean square term. The dependent factor, seed set, was arc-
sine-transformed to improve normality (Sokal and Rohlf 1981).
Levene’s test for homogeneity of variances showed no significant
differences between treatment groups.
Results
Role of wind versus insect pollination
When both wind and insects were excluded from flow-
ers, seed set was near zero (Fig. 1) indicating that out-
side pollen is needed for fertilization. Although some
plants that did mature a few seeds (maximum seed set
was 0.071 seeds/pistil for this treatment), this was likely
Fig. 1 Box plot showing the median (dark line), interquarile range
(box) and outliers (circles) for seed set of branches of Thalictrum
pubescens when various pollination modes are excluded. Different
letters represent significant differences at the P=0.05 level&ig.c:/f
296
Fig. 2A–C Effects of emasculation treatment on branches of fe-
male T. pubescens. A Number of pollinators visiting. B Amount of
pollen deposited on stigmas. C Seed set. Error bars represent one
standard error of the means&ig.c:/f
to have been caused by contamination (e.g., small holes
were found in the pollination bags; the seal around the
bag at the bottom of the branch may not have been tight;
or, because the flowers do not have buds tightly enclosed
within petals, they may have received some pollen prior
to bagging).
When only pollen delivered by wind was allowed to
reach the flowers (the cheesecloth treatment), mean seed
set increased significantly, rising from 2% to 15%
(P=0.02). This indicates that wind can contribute to pol-
lination in T. pubescens and, in one case, seed set was
near 80% (Fig. 1). When both wind and insects were al-
lowed to reach flowers (the control group), mean seed
set was significantly greater than in the wind-only
group, increasing from 15% to 46% (P=0.002). Two dis-
tinct, non-mutually exclusive factors may contribute to
this difference: (1) insects provided some additional pol-
Fig. 3 Relationship between anther number and seed set of fe-
male T. pubescens&ig.c:/f
linations and (2) the cheesecloth blocked some wind-
borne pollen from reaching the stigmas in the wind-only
treatment.
Emasculation experiment, 1994
Emasculated branches received significantly fewer insect
visits than the controls (x ± SE=0.533 ± 0.157 vs 0.967 ±
0.176, P=0.049, Fig. 2A). Nevertheless, because T. pu-
bescens is also capable of being wind pollinated, fewer
insect visits may not necessarily lead to a fitness disad-
vantage for the emasculated branches. For this reason,
both the amount of pollen being delivered to stigmas and
seed set were determined for the emasculated and control
branches.
A lower pollinator visitation rate did not lead to a sig-
nificant difference in the number of pollen grains re-
ceived per stigma (2.172 ± 1.166 vs 2.928 ± 0.760,
P=0.06, Fig. 2B) or in seed set (0.591 ± vs 0.669,
P=0.08, Fig. 2C), although there does appear to be a
trend for the emasculated flowers to have lower values
for each of these variables.
There was no significant correlation between anther
number and seed set (Fig. 3). This indicates that if in-
sects do discriminate against plants on the basis of anther
number, it does not lead to a difference in seed set.
Emasculation and pollen augmentation experiment, 1995
By separating the effects of pollen limitation and emas-
culation, this experiment was designed to more conclu-
sively distinguish between the two possible causes of re-
duced seed set in the 1994 emasculation treatment.
Neither treatment had an effect on seed set (Table 1).
The only significant difference found was due to a plant
effect (Table 1). When no pollen was added (first two
bars in Fig. 4), seed set was almost identical in the emas-
culated and non-emasculated branches (0.5139 ± 0.0397
vs 0.5181 ± 0.0415). A larger difference (although still
 297
Table 1 ANOVA table analyz-
ing effects of the individual Source of variation Sum of df Mean F P
(random), the addition of pol- squares square
len (fixed) and the removal of
anthers (fixed) on seed set of Main effects
female Thalictrum pubescens. Plant 5.490 25 0.220 13.75 <0.001
Seed set (seeds per ovule) was Pollen 0.010 1 0.010 0.476 0.506
arcsine transformed to improve Emasculation 0.003 1 0.003 0.136 0.697
normality&/tbl.c:& Interactions
plant+pollen 0.525 25 0.021 1.313 0.263
plant+emasculation 0.558 25 0.022 1.375 0.217
pollen+emasculation 0.001 1 0.001 0.063 0.786
plant+pollen+emasculation 0.407 25 0.016
Explained 6.994 103 0.068
Residual 0.000 0 0.000

Total 6.994 103 0.068

&/tbl.:

be a result of a compromise between all of these factors.

Fig. 4 Effects of emasculation (emas) and pollen addition (+ pol-
len) on seed set of female T. pubescens (mean ± 1 SE). There are
no statistical differences among the groups&ig.c:/f
not significant) between emasculated and non-emasculat-
ed groups occurred when pollen was added (last two
bars, Fig. 4, 0.5252 ± 0.0413 vs 0.5525 ± 0.0448).
Hence, emasculated branches had slightly lower seed set
even when ample pollen should have been available.
This suggests that the slight trend found in 1994 for low-
er seed set among the emasculated branches may have
been caused by damage during emasculation.
Discussion
Because many plant species mate indirectly through pol-
linators, insects are a strong selective force in the evolu-
tion of breeding systems and floral morphology. This
does not mean, however, that every floral trait displayed
by a plant can be explained as an adaptation for pollina-
tor attraction. Plants may be under numerous opposing
selective pressures or constraints, including developmen-
tal, genetic or ecological. Hence, floral morphology may
In order for the pollinator-attraction hypothesis to ex-
plain the maintenance of stamens in females, two criteria
have to be met. First, insects must show discrimination
against females that have fewer or smaller (or no) sta-
mens. Second, this discrimination must also lead to a fit-
ness disadvantage, such as lower seed set, in order to be
a selective force.
The first of these criteria appears to be met in T. pu-
bescens. Insects visited emasculated branches less often
than the control branches in 1994 (Fig. 2A) and there
was a trend for emasculated flowers to receive less pol-
len per stigma (Fig. 2B). Nonetheless, this trend may
have little biological relevance. The amount of pollen re-
ceived by the emasculated flowers (2–3 pollen
grains/stigma) may be sufficient for full seed set since
each stigma services only one ovule. In addition, this
was an underestimate of the amount of pollen being re-
ceived, since the preparation procedure may have re-
moved some pollen. Therefore, lower visitation will not
necessarily lead to pollen limitation.
The results from both of the emasculation experi-
ments presented in this paper fail to support the second
criterion for the pollinator-attraction hypothesis. In nei-
ther year was there a significant effect of emasculation
on seed set, although there was such a trend in 1994. The
1995 data support the possibility that the trend for emas-
culated branches having lower seed set in 1994 could
have been due to unforseen treatment effects; the physi-
cal removal of anthers apparently caused damage to the
flowers, lowering seed set. When pollen was supple-
mented by hand, so that there was no pollen limitation,
the emasculated group still had slightly lower seed set
(last two bars of Fig. 4) than the non-emasculated group.
Branches were chosen as the experimental unit in the
design of these experiments rather than entire plants for
several reasons. Insects are more likely to be attracted to
other aspects of the floral display from a distance, such
as flower number, inflorescence size, etc., which are
controlled for by comparing branches on the same plant.
In this experiment, insects did discriminate against
298
emasculated branches by visiting them significantly less
often then their non-emasculated counterparts. However,
even with this discrimination, any effect on seed set ap-
pears to be due to damage by the emasculation proce-
dure and not due to fewer insect visits. There was also
no correlation between anther number and seed set, so if
there is insect discrimination at the whole plant level, it
does not lead to a difference in seed set (Fig. 3). There-
fore, the fact that branches were chosen rather than en-
tire plants is unlikely to change the conclusions of this
experiment.
Besides these results, there are several lines of evidence
to suggest that T. pubescens is not pollen limited. In an
initial study by Kaplan and Mulcahy (1971), females en-
closed in bridal veil, to exclude pollination by insects but
not wind, were still able to set seed at slightly less than
half the rate of unenclosed control females. The results of
this paper support their findings that wind contributes sub-
stantially to pollination (Fig. 1). Furthermore, this is an
underestimate of the contribution of wind pollination giv-
en that the cheesecloth undoubted blocked some pollen
transfer by wind. Since wind contributes to pollination,
pollen delivered by insects may be superfluous with re-
spect to seed set. The additional pollen could, however,
contribute to pollen tube competition and perhaps increase
the vigor of the progeny produced (Mulcahy and Mulcahy
1975; Janzen 1977; Winsor et al. 1987).
Second, the fact that there was no evidence of pollen
limitation in 1995 is not due to the fact that pollen was
especially abundant that year. The level of seed set in
1995 was lower than in 1994, even though the 1994
plants received no supplemental pollen. If the lack of a
pollen effect was due to pollen being particularly plenti-
ful in 1995, the level of seed set should have been higher
for all groups in 1995.
Another possibility is that the anthers help to “cap-
ture” pollen being carried by wind or to direct pollen to
the stigmas. The 1995 experiment also rules out this hy-
pothesis. If emasculation causes reduced seed set due to
this hypothesis, rather than damage, the addition of pol-
len would have raised the seed set of the emasculated
branches, but this did not happen (Fig. 4).
Collectively, this evidence argues that the pollinator-at-
traction hypothesis may have only a secondary role in the
maintenance of stamens in females of T. pubescens. The
results from both years indicate that emasculated flowers
do not have lower seed set than the non-emasculated con-
trols. Consequently, in order to understand why these or-
gans are present, attention must now be focused on the two
other hypotheses that may account for their maintenance.
Preliminary evidence also argues against the idea that
the stamens are simply vestiges of the ancestral state (the
ancestry hypothesis); there is a negative phenotypic cor-
relation between stamen number and pistil number with-
in females (Davis, unpublished data). In other words, for
each stamen produced, fewer pistils are produced per
flower. This, then, directly affects fitness, as each pistil
makes one seed. If this phenotypic correlation has a ge-
netic basis, this would constitute a substantial cost to the
production of stamens and would argue against the an-
cestry hypothesis.
The third possibility is that the stamens are main-
tained by genetic constraints. When dioecy first arises,
sexual dimorphism should be low and, presumably, ho-
mologous characters in the two sexes should be under
the control of the same or overlapping sets of genes in
each sex (Meagher 1992). Under these conditions, selec-
tion for reduction of the nonfunctional organs in one sex
may be hindered by strong positive genetic correlations
with the homologous organs in the opposite sex; i.e., se-
lection for resource reallocation and reduction of anthers
in females would be countered by selection favoring an-
thers in males. Such genetic correlations between the
sexes would limit the independent evolution of these
traits in males and females while sustaining genetic vari-
ation. This hypothesis is currently being investigated.
&p.2:Acknowledgements The author wishes to thank L. Delph, D. Du-
dle, J. Gehring, D. Marr, D. McClellan, and L. Rieseberg for help-
ful comments on an earlier version of the manuscript; J. Price for
insect identification; and H. Lindman for statistical advice. This
research was supported by the Indiana Academy of Sciences.
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