A two-pollinator model for the evolution of floral complexity

For a new, more complex floral form to become established in a population it must overcome the problem of frequency-dependent constancy to successfully attract pollinators. This may be achieved by complex floral forms offering absolute greater rewards than the simpler forms, or by complex flowers offering a higher probability of being rewarding because fewer pollinators are able to visit them. In this paper we examine the effect of three pollinator foraging strategies on the ratio of flights within and between floral morphs and hence on the probability of a new morph establishing in a population without offering a greater reward. We incorporate pollinator behaviour based around observations of two pollinator species systems into three models of competition for pollinators. In the first model the constancy of the pollinator of the new floral morph is a function only of the foraging strategy of the existing pollinator of the original floral morph. In the next model the constancy of the second pollinator is determined by the number of rewarding flowers of each floral morph left by the original pollinator and in the third model it is determined by the ratio of rewarding flowers of each morph left by the original pollinator. The results demonstrate that under conditions of intense competition for pollinators, new, more complex floral forms are indeed able to attract high levels of constant pollinators without offering intrinsically higher rewards. However, for this to occur constancy in one of the pollinators must be a function of the ratio of rewarding to non-rewarding flowers of both floral forms. One prediction from our results is that sympatric speciation of floral complexity based on a higher probability of reward is more likely to occur in flowers offering rewards of pollen rather than nectar. This is because the cost of visiting non-rewarding flowers is usually higher where the reward is pollen rather than nectar. We also predict that complex flowers occurring at low frequency, which offer rewards of nectar, may need intrinsically greater rewards if they are to successfully attract pollinators.     

FLORAL COLOR CHANGE IN LUPINUS ARGENTEUS (FABACEAE): WHY SHOULD PLANTS ADVERTISE THE LOCATION OF UNREWARDING FLOWERS TO POLLINATORS?

I examined the adaptive significance of two floral traits in the perennial herb, Lupinus argenteus: 1) the retention of corollas on “spent” flowers, i.e., flowers containing inviable pollen, unreceptive stigmas, and negligible pollinator rewards and 2) a change in corolla color of retained “spent” flowers, which is restricted to a spot on the banner petal. At anthesis, this spot is yellow, and approximately four days later, it changes to purple. After the change, purple flowers remain on plants an additional 5–7 days before corolla abscission occurs; purple flowers were avoided by pollinators, presumably because they contained less pollen (rewards) than yellow ones. I experimentally tested the hypothesis that purple flowers contribute to the floral display of the plant by removing varying numbers of spent flowers and assessing the effect on pollination visitation. Pollinators preferentially approached and foraged on plants with greater numbers of flowers per inflorescence; they did not discriminate between yellow (rewarding) and purple (nonrewarding) flowers at interplant distances greater than 0.4 meters but would preferentially forage on plants with more total flowers, even if these individuals contained fewer rewarding flowers. Thus, spent flowers increased the overall attractiveness of plants to pollinators. In theory, color change may benefit plants in two ways. First, by directing pollinators to rewarding flowers, the change may increase pollinator foraging efficiency, with the result that pollinators visit more flowers before leaving plants (pollinator-tenure mechanism). Second, by directing pollinators to receptive flowers, the color change may prevent incoming pollen from being wasted on unreceptive stigmas and may prevent collection of inviable pollen (pollination-efficiency mechanism). I tested the pollinator-tenure mechanism experimentally by removing pollen from yellow flowers, thereby reducing the reliability of the color-reward signal. Pollinators visited fewer total flowers on experimental plants than on controls, resulting in reduced seed production in one year.     

Are deception-pollinated species more variable than those offering a reward?

In most pollination systems, animals transfer pollen among plants of a given species. Pollinator visitations do not come without cost, so plants usually offer a reward. However, the flowers of some plant species, mostly orchids, lack rewards and deceive animals into visiting their flowers. Deceptive species are thought to have high levels of variation in traits associated with advertisement and pollinator attraction, which have been attributed to genetic drift, or disruptive selection due to pollinator behavior. Rewarding species are assumed to have less variation due to stabilizing selection. We compared variability in floral morphology and fragrance composition between deceptive and rewarding species. Because both suites of traits are often linked with floral advertisement and pollinator attraction, we expected variation to be greater in species with deceptive pollination systems than in those offering rewards. We obtained floral morphology metrics for 20 deceptive species and 41 rewarding species native or naturalized in Puerto Rico, Venezuela, and Ecuador. Floral fragrances were sampled from eight deceptive species and four rewarding species. We found that the amplitude of variation in floral morphology and fragrance composition covaries significantly. Comparison of coefficients of variation for morphology indicated that, overall, deceptive species show significantly higher variation than rewarding species, and this pattern was also found among just orchids or just nonorchids. There were no statistical differences in morphological variation between orchids and nonorchids within a functional pollination group. Fragrance variation, measured by Jaccard distance, tended to be greater for deceptive species than for rewarding species. Although overlap in measures of variation occurs between the two groups, the data support the hypothesis that populations of deception-pollinated species are more variable than rewarding species in traits associated with pollinator attraction.     

Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Sex allocation and investment into pre- and post-copulatory traits in simultaneous hermaphrodites: the role of polyandry and local sperm competition

Sex allocation theory predicts the optimal allocation to male and female reproduction in sexual organisms. In animals, most work on sex allocation has focused on species with separate sexes and our understanding of simultaneous hermaphrodites is patchier. Recent theory predicts that sex allocation in simultaneous hermaphrodites should strongly be affected by post-copulatory sexual selection, while the role of pre-copulatory sexual selection is much less clear. Here, we review sex allocation and sexual selection theory for simultaneous hermaphrodites, and identify several strong and potentially unwarranted assumptions. We then present a model that treats allocation to sexually selected traits as components of sex allocation and explore patterns of allocation when some of these assumptions are relaxed. For example, when investment into a male sexually selected trait leads to skews in sperm competition, causing local sperm competition, this is expected to lead to a reduced allocation to sperm production. We conclude that understanding the evolution of sex allocation in simultaneous hermaphrodites requires detailed knowledge of the different sexual selection processes and their relative importance. However, little is currently known quantitatively about sexual selection in simultaneous hermaphrodites, about what the underlying traits are, and about what drives and constrains their evolution. Future work should therefore aim at quantifying sexual selection and identifying the underlying traits along the pre- to post-copulatory axis.     

Association between floral traits and rewards in Erysimum mediohispanicum (Brassicaceae)

Background and Aims

Floral rewards may be associated with certain morphological floral traits and thus act as underlying factors promoting selection on these traits. This study investigates whether some traits that are under pollinator-mediated selection (flower number, stalk height, corolla diameter, corolla tube length and corolla tube width) in the Mediterranean herb E. mediohispanicum (Brassicaceae) are associated with rewards (pollen and nectar).

Methods

During 2005 the phenotypic traits and the visitation rate of the main pollinator functional groups were quantified in 720 plants belonging to eight populations in south-east Spain, and during 2006 the same phenotypic traits and the reward production were quantified in 400 additional plants from the same populations.

Key Results

A significant correlation was found between nectar production rate and corolla tube length, and between pollen production and corolla diameter. Visitation rates of large bees and butterflies were significantly higher in plants exhibiting larger flowers with longer corolla tubes.

Conclusions

The association between reward production and floral traits may be a factor underlying the pattern of visitation rate displayed by some pollinators.Key words: Erysimum, floral traits, nectar, pollen, pollinator visitation rate, reward     

Maternal investment and male reproductive success in angiosperms: parent-offspring conflict or sexual selection?

It is possible to interpret components of seed development in angiosperms from the perspective of parent-offspring conflict (a special case of kin selection) or sexual selection. Available parent-offspring conflict models predict the evolution of traits determining the outcome of competition among related individuals soliciting maternal resources. In such models, ‘selfishness’ may spread even if it reduces female fecundity and thus population mean fitness may decline. These models are limited, however, because most of them do not simultaneously consider selection among maternal genotypes varying in the tendency to respond to their offspring. Available sexual selection models, in contrast, do consider the joint evolution of polygenic male traits (influencing viability, mating success and fecundity) and female preferences (influencing the mating success of different male phenotypes). These models have shown that male traits may evolve that are non-optimal with respect to viability. Only one recent sexual selection model explicitly incorporates direct fecundity selection upon females; this model concludes that fecundity will be maximized at equilibrium. Hence population mean fitness may decline due to reduced male viability but not due to diminished female fecundity. Available sexual selection models, however, are limited because they do not consider the effects of interactions among relatives. The assumptions and qualitative results of the two types of models are compared and discussed in the context of seed development. Differential allocation of maternal resources among genetically distinct developing seeds may be viewed from the perspective of either. Because the results of the available models of parent-offspring conflict and sexual selection are not wholly consistent and because data confirming the genetic basis of maternal patterns of investment or differential male reproductive success are scant, it is not clear which set of conclusions is most appropriate to apply to plants. To achieve the generality towards which mathematical approaches aspire, new models concerning the evolution of traits influencing resource allocation in plants must incorporate the components of both parent-offspring conflict and sexual selection.     

Understanding reversals in the relative strength of sexual selection on males and females: a role for sperm competition?

Sperm competition affects sexual selection intensity on males, but models suggest it cannot affect the relative intensity of sexual selection on males compared to females. However, if sperm competition depresses the payoff for male multiple mating, it could affect the relative intensity of sexual selection and even cause sexual selection to be more intense on females than males (reversal of typical pattern). To evaluate how sperm competition, energy availability, and parental investment affect the intensity of sexual selection on each sex, I constructed a simulation model using the relationship between fecundity and number of mates to estimate sexual selection gradients. Unlike earlier models, I include a trade-off between paternal investment and sperm competition ability. The amount of energy available for reproduction affects the sexual selection gradient for each sex. Reversals in the sex experiencing stronger sexual selection do occur when additional paternal investment reduces a male's ability to compete for fertilizations within females. The shape of the distribution of mates for each sex (determined by mate competition) is also important. Output from the model is qualitatively similar to empirical data from insects with paternal investment. This model challenges previous thinking about the role of sperm competition in sex-role reversal.     

The relative importance of intra- and intersexual selection on human male sexually dimorphic traits

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N?=?164), we sought to provide further evidence on the effects of men's physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective ratings of male sexually dimorphic traits purportedly under sexual selection (height, vocal and facial masculinity, upper body size from 3D scans, physical strength, and baseline testosterone) and observer perceptions of physical dominance and sexual attractiveness based on self-presentation video recordings were assessed and associated with mating success (sociosexual behaviour and number of potential conceptions) in a partly longitudinal design. Results from structural equation models and selection analyses revealed that physical dominance, but not sexual attractiveness, predicted mating success. Physical dominance mediated associations of upper body size, physical strength, as well as vocal and facial physical dominance and attractiveness with mating success. These findings thus suggest a greater importance of intrasexual competition than female choice in human male sexual selection.     

Pollinators and seed predators generate conflicting selection on Dalechampia blossoms

Plant–pollinator relationships are often mediated by floral traits that advertise the presence or amount of rewards. However, herbivores may also use these traits to find their hosts. In Dalechampia scandens, we tested whether floral advertisements that attract pollinators were also used by seed predators, and whether this generated conflicting selection pressures. We studied the influence of natural variation in the size of showy bracts, amount of reward, and two shape traits on pollinator visitation, pollen arrival on stigmas, seed production and seed predation. We then built a multivariate fitness function for these traits to estimate selection generated by pollinators and seed predators. Blossoms with larger bracts were visited by bees more frequently and received more pollen on their stigmas. Seed predators laid more eggs on blossoms with larger bracts and also on blossoms later producing more seeds. Consequently, selection for larger bract size exerted by pollinators was counteracted by the selection exerted by seed predators. As a result, net selection on bract size tended to be stabilizing. Additionally, we found selection on traits that increased the rate of self‐pollination (assuming uniform seed quality). These results illustrate the importance of both mutualists and antagonists in floral evolution, as well as the value of taking an integrative approach to assessing selection on floral traits.     

Optimal pollinator attraction strategies in Trollius ranunculoides Hemsl. (Ranunculaceae) at different altitudes: increased floral display or promotion of nectar output?

For alpine plant species, patterns of resource allocation to functional floral traits for pollinator attraction can be highly significant in adaptation to low pollinator abundance and consequent pollen limitation. Increased pollination can be achieved either through a larger floral display or production of more pollen rewards. In this study, variation in resource allocation to different components for pollinator attraction was studied along an altitudinal gradient in Trollius ranunculoides, an obligate self‐incompatible out‐crosser of the Qinghai–Tibet Plateau. We compared resource allocation to conspicuous yellow sepals (which mainly provide visual attraction) and degenerate petals (which provide the major nectar reward) between populations at four altitudes. Furthermore, we investigated the contribution of sepals and petals to pollinator attraction and female reproductive success in an experiment with sepal or petal removal at sites at different altitudes. At the level of single flowers, resource allocation increased to sepals but decreased to petals with increasing altitude. Consistent with these results, sepals contributed much more to visitation rate and seed set than petals, as confirmed in the sepal or petal removal experiment. Sepals and petals contributed to female reproductive success by ensuring visitation rate rather than visitation duration. To alleviate increasing pollen limitation with increasing altitude, resource allocation patterns of T. ranunculoides altered to favour development of sepals rather than petals. This strategy may improve pollination and reproductive success through visual attraction (sepal) rather than nectar reward (petal) over a gradient of decreasing pollinator abundance.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

THE EVOLUTION OF POLLINATOR–PLANT INTERACTION TYPES IN THE ARACEAE

Most plant–pollinator interactions are mutualistic, involving rewards provided by flowers or inflorescences to pollinators. Antagonistic plant–pollinator interactions, in which flowers offer no rewards, are rare and concentrated in a few families including Araceae. In the latter, they involve trapping of pollinators, which are released loaded with pollen but unrewarded. To understand the evolution of such systems, we compiled data on the pollinators and types of interactions, and coded 21 characters, including interaction type, pollinator order, and 19 floral traits. A phylogenetic framework comes from a matrix of plastid and new nuclear DNA sequences for 135 species from 119 genera (5342 nucleotides). The ancestral pollination interaction in Araceae was reconstructed as probably rewarding albeit with low confidence because information is available for only 56 of the 120–130 genera. Bayesian stochastic trait mapping showed that spadix zonation, presence of an appendix, and flower sexuality were correlated with pollination interaction type. In the Araceae, having unisexual flowers appears to have provided the morphological precondition for the evolution of traps. Compared with the frequency of shifts between deceptive and rewarding pollination systems in orchids, our results indicate less lability in the Araceae, probably because of morphologically and sexually more specialized inflorescences.     

Investment in defense and cost of predator-induced defense along a resource gradient

An organism’s investment in different traits to reduce predation is determined by the fitness benefit of the defense relative to the fitness costs associated with the allocation of time and resources to the defense. Inherent tradeoffs in time and resource allocation should result in differential investment in defense along a resource gradient, but competing models predict different patterns of investment. There are currently insufficient empirical data on changes in investment in defensive traits or their costs along resource gradients to differentiate between the competing allocation models. In this study, I exposed tadpoles to caged predators along a resource gradient in order to estimate investment in defense and costs of defense by assessing predator-induced plasticity. Induced defenses included increased tail depth, reduced feeding, and reduced swimming activity; costs associated with these defenses were reduced developmental rate, reduced growth, and reduced survival. At low resource availability, these costs predominately resulted in reduced survival, while at high resource availability the costs yielded a reduced developmental rate. Defensive traits responded strongly to predation risk, but did not respond to resource availability (with the exception of feeding activity), whereas traits construed as costs of defenses showed the opposite pattern. Therefore, defensive traits were highly sensitive to predation risk, while traits construed as costs of defense were highly sensitive to resource allocation tradeoffs. This difference in sensitivity between the two groups of traits may explain why the correlation between the expression of defensive traits and the expression of the associated defense costs was weak. Furthermore, my results indicate that genetic linkages and mechanistic integration of multiple defensive traits and their associated costs may constrain time and resource allocation in ways that are not addressed in existing models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Does competition with wind‐pollinated species alter Echium plantagineum's attractiveness to a common pollinator Bombus terrestris?

1. In insect‐pollinated plants, pollinator attraction is influenced by flowers (e.g. number, size) and their associated rewards (e.g. pollen, nectar). These traits can depend on plant interactions. Indeed, below‐ground competition between plants can lead to a decrease in flower or reward production in insect‐pollinated species. 2. Wind‐pollinated plants, in particular, which are almost never studied in plant–pollinator networks, can alter insect‐pollinated plants' attractiveness through competition for nutrients. The response of pollinators to such changes has never been investigated. 3. A pot experiment was carried out in which an insect‐pollinated species, Echium plantagineum, was grown in binary mixture with three wind‐pollinated species selected to exert a panel of competitive interactions. Below‐ground competition was controlled using dividers limiting interspecific root competition. Floral traits of E. plantagineum (i.e. flower production, floral display size, flower size and nectar production) were measured. For each species mixture, the visits (i.e. first visit, number of visits, 10‐min sequences) of Bombus terrestris individuals released in a flight cage containing two pots were followed, one with and one without below‐ground competition. 4. Below‐ground competition significantly affected nectar's sucrose concentration but did not influence flower and nectar production. Likewise, pollinator visits were not influenced by below‐ground competition. Competitor identity significantly influenced flower and reward production of E. plantagineum, with a decrease in the presence of the most competitive wind‐pollinated species. A tendency for faster flower visitation events was also detected in the presence of the least competitive competitor. This study raises new questions regarding the influence of wind‐pollinated plants on plant–pollinator interactions.     

Mechanisms and evolution of deceptive pollination in orchids

The orchid family is renowned for its enormous diversity of pollination mechanisms and unusually high occurrence of non-rewarding flowers compared to other plant families. The mechanisms of deception in orchids include generalized food deception, food-deceptive floral mimicry, brood-site imitation, shelter imitation, pseudoantagonism, rendezvous attraction and sexual deception. Generalized food deception is the most common mechanism (reported in 38 genera) followed by sexual deception (18 genera). Floral deception in orchids has been intensively studied since Darwin, but the evolution of non-rewarding flowers still presents a major puzzle for evolutionary biology. The two principal hypotheses as to how deception could increase fitness in plants are (i) reallocation of resources associated with reward production to flowering and seed production, and (ii) higher levels of cross-pollination due to pollinators visiting fewer flowers on non-rewarding plants, resulting in more outcrossed progeny and more efficient pollen export. Biologists have also tried to explain why deception is overrepresented in the orchid family. These explanations include: (i) efficient removal and deposition of pollinaria from orchid flowers in a single pollinator visit, thus obviating the need for rewards to entice multiple visits from pollinators; (ii) efficient transport of orchid pollen, thus requiring less reward-induced pollinator constancy; (iii) low-density populations in many orchids, thus limiting the learning of associations of floral phenotypes and rewards by pollinators; (iv) packaging of pollen in pollinaria with limited carry-over from flower to flower, thus increasing the risks of geitonogamous self-pollination when pollinators visit many flowers on rewarding plants. All of these general and orchid-specific hypotheses are difficult to reconcile with the well-established pattern for rewardlessness to result in low pollinator visitation rates and consequently low levels of fruit production. Arguments that deception evolves because rewards are costly are particularly problematic in that small amounts of nectar are unlikely to have a significant effect on the energy budget of orchids, and because reproduction in orchids is often severely pollen-, rather than resource-limited. Several recent experimental studies have shown that deception promotes cross-pollination, but it remains unknown whether actual outcrossing rates are generally higher in deceptive orchids. Our review of the literature shows that there is currently no evidence that deceptive orchids carry higher levels of genetic load (an indirect measure of outcrossing rate) than their rewarding counterparts. Cross-pollination does, however, result in dramatic increases in seed quality in almost all orchids and has the potential to increase pollen export (by reducing pollen discounting). We suggest that floral deception is particularly beneficial, because of its promotion of outcrossing, when pollinators are abundant, but that when pollinators are consistently rare, selection may favour a nectar reward or a shift to autopollination. Given that nectar-rewardlessness is likely to have been the ancestral condition in orchids and yet is evolutionarily labile, more attention will need to be given to explanations as to why deception constitutes an 'evolutionarily stable strategy'.     

A Tale of Two Morphs: Modeling Pollen Transfer,Magic Traits,and Reproductive Isolation in Parapatry

The evolution of the flower is commonly thought to have spurred angiosperm diversification. Similarly, particular floral traits might have promoted diversification within specific angiosperm clades. We hypothesize that traits promoting the precise positional transfer of pollen between flowers might promote diversification. In particular, precise pollen transfer might produce partial reproductive isolation that facilitates adaptive divergence between parapatric populations differing in their reproductive-organ positions. We investigate this hypothesis with an individual-based model of pollen transfer dynamics associated with heterostyly, a floral syndrome that depends on precise pollen transfer. Our model shows that precise pollen transfer can cause sexual selection leading to divergence in reproductive-organ positions between populations served by different pollinators, pleiotropically causing an increase in reproductive isolation through a “magic trait” mechanism. Furthermore, this increased reproductive isolation facilitates adaptive divergence between the populations in an unlinked, ecologically selected trait. In a different pollination scenario, however, precise pollen transfer causes a decrease in adaptive divergence by promoting asymmetric gene flow. Our results highlight the idea that magic traits are not “magic” in isolation; in particular, the effect size of magic traits in speciation depends on the external environment, and also on other traits that modify the strength of the magic trait''s influence on non-random mating. Overall, we show that the evolutionary consequences of pollen transfer dynamics can depend strongly on the available pollinator fauna and on the morphological fit between flowers and pollinators. Furthermore, our results illustrate the potential importance of even weak reproductive isolating barriers in facilitating adaptive divergence.     

Quantifying the strength and form of sexual selection on men's traits

Although recent research has increasingly focused on human sexual selection, fundamental questions remain concerning the relative influence of individual traits on success in competition for mates and the mechanisms, form, and direction of these sexual selective pressures. Here, we explore sexual selection on men's traits by ascertaining men's dominance and attractiveness from male and female acquaintances. On a large American university campus, 63 men from two social fraternities provided anthropometric measurements, facial photographs, voice recordings, and reported mating success (number of sexual partners). These men also assessed each other's dominance, and 72 women from two socially affiliated sororities assessed the men's attractiveness. We measured facial masculinity from inter-landmark distances and vocal masculinity from acoustic parameters. We additionally obtained facial and vocal attractiveness and dominance ratings from unfamiliar observers. Results indicate that dominance and the traits associated with it predict men's mating success, but attractiveness and the traits associated with it do not. These findings point to the salience of contest competition on men's mating success in this population.     

ECOLOGICAL COSTS OF PLANT RESISTANCE TO HERBIVORES IN THE CURRENCY OF POLLINATION

In this paper, we examine how ecological costs of resistance might be manifested through plant relationships with pollinators. If defensive compounds are incorporated into floral structures or if they are sufficiently costly that fewer rewards are offered to pollinators, pollinators may discriminate against more defended plants. Here we consider whether directional selection for increased resistance to herbivores could be constrained by opposing selection through pollinator discrimination against more defended plants. We used artificial selection to create two populations of Brassica rapa plants that had high and low myrosinase concentrations and, consequently, high and low resistance to flea beetle herbivores. We measured changes in floral characters of plants in both damaged and undamaged states from these populations with different resistances to flea beetle attack. We also measured pollinator visitation to plants, including numbers of pollinators and measures of visit quality (numbers of flowers visited and time spent per flower). Damage from herbivores resulted in reduced petal size, as did selection for high resistance to herbivores later in the plant lifetime. In addition, floral display (number of open flowers) was also altered by an interaction between these two effects. Changes in floral traits translated into overall greater use of low-resistance, undamaged plants based on total amount of time pollinators spent foraging on plants. Total numbers of pollinators attracted to plants did not differ among treatments; however, pollinators spent significantly more time per flower on plants from the low-resistance population and tended to visit more flowers on these plants as well. Previous work by other investigators on the same pollinator taxa has shown that longer visit times are associated with greater male and female plant fitness. Because initial numbers of pollinators did not differ between selection regimes, palatability and/or amount of rewards offered by high- and low-resistance populations are likely to be responsible for these patterns. During periods of pollinator limitation, less defended plants may have a selective advantage and pollinator preferences may mediate directional selection imposed by herbivores. In addition, if pollinator preferences limit seed set in highly defended plants, then lower seed set previously attributed to allocation costs of defense may also reflect greater pollinator limitation in these plants relative to less defended plants.     

The role of maternal and paternal effects in the evolution of parental quality by sexual selection

Genetic models of maternal effects and models of mate choice have focused on the evolutionary effects of variation in parental quality. There have been, however, few attempts to combine these into a single model for the evolution of sexually selected traits. We present a quantitative genetic model that considers how male and female parental quality (together or separately) affect the expression of a sexually selected offspring trait. We allow female choice of males based on this parentally affected trait and examine the evolution of mate choice, parental quality and the indicator trait. Our model reveals a number of consequences of maternal and paternal effects. (1) The force of sexual selection owing to adaptive mate choice can displace parental quality from its natural selection optimum. (2) The force of sexual selection can displace female parental quality from its natural selection optimum even when nonadaptive mate choice occurs (e.g. runaway sexual selection), because females of higher parental quality produce more attractive sons and these sons counterbalance the loss in fitness owing to over-investment in each offspring. (3) Maternal and paternal effects can provide a source of genetic variation for offspring traits, allowing evolution by sexual selection even when those traits do not show direct genetic variation (i.e. are not heritable). (4) The correlation between paternal investment and the offspring trait influenced by the parental effects can result in adaptive mate choice and lead to the elaboration of both female preference and the male sexually selected trait. When parental effects exist, sexual selection can drive the evolution of parental quality when investment increases the attractiveness of offspring, leading to the elaboration of indicator traits and higher than expected levels of parental investment.