Reproductive isolation between diploid and tetraploid individuals in mixed-cytotype populations of Lycium australe.

PREMISE: Whole-genome duplication is considered a major mechanism of sympatric speciation due to the creation of strong and instantaneous reproductive barriers. Although postzygotic reproductive isolation between diploids and polyploids is often expected, the extent of reproductive incompatibility must be empirically determined and compared to patterns of genetic isolation to fully characterize the reproductive dynamics between cytotypes. METHODS: We investigated reproductive compatibility between diploid and tetraploid Lycium australe in two mixed-cytotype populations using (1) controlled crossing experiments to evaluate fruit and seed production and (2) germination trials to test seed viability following homoploid and heteroploid crosses. We contrast these experiments with a single-nucleotide polymorphism (SNP) data set to measure genetic isolation between cytotypes and explore whether cytotype or population origin better explains patterns of genetic variation. Finally, we explore mating patterns using the observed germination rates of naturally produced seeds in each population. RESULTS: Although homoploid and heteroploid crosses resulted in similar fruit and seed production, reproductive isolation between co-occurring diploids and tetraploids was nearly complete, due to low seed viability following heteroploid crosses. Of 191,182 total SNPs, 21,679 were present in ≥90% of individuals and replicate runs using unlinked SNPs revealed strong clustering by cytotype and differentiation of tetraploids based on population origin. CONCLUSIONS: As often reported, diploid and tetraploid L. australe experience strong postzygotic isolation via hybrid seed inviability. Consistent with this result, cytotype explained a greater amount of variation in the SNP data set than population origin, despite some evidence of historical introgression.

Molecular signatures of long-distance oceanic dispersal and the colonization of Pacific islands in Lycium carolinianum.

PREMISE: Long-distance dispersal has been important in explaining the present distributions of many plant species. Despite being infrequent, such dispersal events have considerable evolutionary consequences, because bottlenecks during colonization can result in reduced genetic diversity. We examined the phylogeographic history of Lycium carolinianum, a widespread taxon that ranges from southeastern North America to several Pacific islands, with intraspecific diversity in sexual and mating systems. METHODS: We used Bayesian, likelihood, and coalescent approaches with nuclear and plastid sequence data and genome-wide single nucleotide polymorphisms to reconstruct the dispersal history of this species. We also compared patterns of genetic variation in mainland and island populations using single nucleotide polymorphisms and allelic diversity at the S-RNase mating system gene. RESULTS: Lycium carolinianum is monophyletic and dispersed once from the North American mainland, colonizing the Pacific islands ca. 40,100 years ago. This dispersal was accompanied by a loss of genetic diversity in SNPs and the S-RNase locus due to a colonization bottleneck and the loss of self-incompatibility. Additionally, we documented at least two independent transitions to gynodioecy: once following the colonization of the Hawaiian Islands and loss of self-incompatibility, and a second time associated with polyploidy in the Yucatán region of Mexico. CONCLUSIONS: Long-distance dispersal via fleshy, bird dispersed fruits best explains the unusually widespread distribution of L. carolinianum. The collapse of diversity at the S-RNase locus in island populations suggests that self-fertilization may have facilitated the subsequent colonization of Pacific islands following a single dispersal from mainland North America.

Colonization, Baker's law, and the evolution of gynodioecy in Hawaii: implications from a study of Lycium carolinianum.

PREMISE: As Baker's law suggests, the successful colonization of oceanic islands is often associated with uniparental reproduction (self-fertility), but the high incidence of dimorphism (dioecy, gynodioecy) on islands complicates this idea. Lycium carolinianum is widespread, occurring on the North American mainland and the Hawaiian Islands. We examined Baker's ideas for mainland and island populations of L. carolinianum and examined inbreeding depression as a possible contributor to the evolution of gynodioecy on Maui. METHODS: Controlled crosses were conducted in two mainland populations and two populations in Hawaii. Treatments included self and cross pollination, unmanipulated controls, and autogamy/agamospermy. Alleles from the self-incompatibility S-RNase gene were isolated and compared between mainland and island populations. Given self-compatibility in Hawaii, we germinated seeds from self- and cross- treatments and estimated inbreeding depression using seven traits and a measure of cumulative fitness. RESULTS: Mainland populations of Lycium carolinianum are predominately self-incompatible with some polymorphism for self-fertility, whereas Hawaiian populations are self-compatible. Concordantly, S-RNase allelic diversity is reduced in Hawaii compared to the mainland. Hawaiian populations also exhibit significant inbreeding depression. CONCLUSIONS: Self-compatibility in Hawaii and individual variation in self-fertility in mainland populations suggests that a colonization filter promoting uniparental reproduction may be acting in this system. Comparison of S-RNase variation suggests a collapse of allelic diversity and heterozygosity at the S-RNase locus in Hawaii, which likely contributed to mate limitation upon arrival to the Pacific. Inbreeding depression coupled with autonomous self-fertilization may have led to the evolution of gynodioecy on Maui.

Floral size and shape evolution following the transition to gender dimorphism.

PREMISE OF THE STUDY: Floral morphology is expected to evolve following the transition from cosexuality to gender dimorphism in plants, as selection through male and female function becomes dissociated. Specifically, male-biased dimorphism in flower size can arise through selection for larger flowers through male function, selection for smaller flowers through female function, or both. The evolutionary pathway to floral dimorphism can be most effectively reconstructed in species with intraspecific variation in sexual system. We examined the evolution of flower size and shape in Lycium californicum, whose populations are either gender dimorphic with male and female plants, or cosexual with hermaphroditic plants. METHODS: Floral morphology was characterized in populations spanning the species' complete range. For a subset of the range where cosexual and dimorphic populations are in close proximity, we compared the size and shape of flowers from female and male plants in dimorphic populations to hermaphrodites in cosexual populations, accounting for variation associated with abiotic environmental conditions. KEY RESULTS: The magnitude of flower size dimorphism varied across dimorphic populations. After controlling for environmental variation across cosexual and dimorphic populations, flowers on males were larger than flowers on females and hermaphrodites, whereas flower size did not differ between females and hermaphrodites. Flower shape differences were associated with mating type, sexual system, and environmental variation. CONCLUSIONS: While abiotic environmental gradients shape both overall flower size and shape, male-biased flower size dimorphism in L. californicum appears to arise through selection for larger flowers in males but not smaller flowers in females.

Correlated polymorphism in cytotype and sexual system within a monophyletic species, Lycium californicum.

BACKGROUND AND AIMS: Polyploidy has important effects on reproductive systems in plants and has been implicated in the evolution of dimorphic sexual systems. In particular, higher ploidy is associated with gender dimorphism across Lycium species (Solanaceae) and across populations within the species Lycium californicum. Previous research on the association of cytotype and sexual system within L. californicum sampled a limited portion of the species range, and did not investigate evolutionary transitions between sexual systems. Lycium californicum occurs in arid regions on offshore islands and mainland regions in the south-western United States and Mexico, motivating a more comprehensive analysis of intraspecific variation in sexual system and cytotype across the full range of this species. METHODS: Sexual system (dimorphic vs. cosexual) was determined for 34 populations across the geographical range of L. californicum using field observations of pollen production, and was confirmed using morphological measurements and among-plant correlations of primary sexual traits. Ploidy was inferred using flow cytometry in 28 populations. DNA sequence data from four plastid and two nuclear regions were used to reconstruct relationships among populations and to map transitions in sexual system and ploidy. KEY RESULTS: Lycium californicum is monophyletic, ancestrally diploid and cosexual, and the association of gender dimorphism and polyploidy appears to have two evolutionary origins in this species. Compared with cosexual populations, dimorphic populations had bimodal anther size distributions, negative correlations between male and female floral traits, and larger coefficients of variation for primary sexual traits. Flow cytometry confirmed tetraploidy in dimorphic populations, whereas cosexual populations were diploid. CONCLUSIONS: Tetraploidy and gender dimorphism are perfectly correlated in L. californicum, and the distribution of tetraploid-dimorphic populations is restricted to populations in Arizona and the Baja California peninsula. The analysis suggests that tetraploidy and dimorphism likely established in Baja California and may have evolved multiple times.

Intraspecific variation in gender strategies in Lycium (Solanaceae): associations with ploidy and changes in floral form following the evolution of gender dimorphism.

UNLABELLED: • PREMISE OF THE STUDY: An association between polyploidy and gender dimorphism has been noted in several plant lineages. Whereas the majority of Lycium species are diploid and have hermaphroditic flowers in cosexual populations, gender dimorphism (gynodioecy, dioecy) has been shown to be uniformly associated with polyploidy in previous studies. Preliminary field observations suggested that some populations of Lycium carolinianum were dimorphic, providing a test of this association.• METHODS: We assessed sexual systems and cytotype variation (to infer ploidy) across 17 populations of L. carolinianum. Comparison of flowers in cosexual and dimorphic populations were used to infer changes in reproductive morphology associated with the evolution of gynodioecy.• KEY RESULTS: The majority of populations were cosexual in gender expression, but dimorphism was present in the Yucatán and in some populations in Hawaii. Populations varied in ploidy and were either diploid or tetraploid. Floral sexual dimorphism was present in all gynodioecious populations, though the magnitude differed and was cryptic in some cases. Our results are consistent with the hypothesis that following the evolution of gynodioecy, flowers on hermaphrodites increased in size.• CONCLUSIONS: Dimorphic sexual systems have likely evolved convergently in L. carolinianum. In contrast to previous studies, dimorphism is not perfectly associated with polyploidy. Although our sample from the Yucatán was both tetraploid and dimorphic, all populations in Hawaii were diploid regardless of sexual system. Ongoing phylogeographic and mating system studies will contribute to our understanding of reproductive evolution in this widespread, polymorphic species.

Developmental plasticity, genetic assimilation, and the evolutionary diversification of sexual expression in Solanum.

PREMISE OF THE STUDY: For over a century, it has been hypothesized that selection can convert an environmentally induced phenotype (i.e., plasticity) into a fixed (constitutively produced) phenotype, a process known as genetic assimilation. While evidence of assimilation is accumulating, the role of plasticity generally and assimilation specifically in evolutionary diversification has rarely been examined from a comparative phylogenetic perspective. • METHODS: We combined experimental analyses of plasticity with ancestral state reconstructions to examine the evolutionary dynamics of sexual expression in two well-characterized sections (Acanthophora and Lasiocarpa) in Solanum subgenus Leptostemonum. We examined sexual expression phenotypes and the proportion of staminate flowers produced under contrasting resource conditions in 10 species and combined these data with previous studies. • KEY RESULTS: Staminate flower production was phenotypically plastic for nine of 14 species and unaffected by treatment in five species. Two of the nonplastic species bore few staminate flowers, and three constitutively produced large numbers of staminate flowers. For individuals and species producing staminate flowers, these flowers occurred in a distinctive architectural pattern that was qualitatively the same in both plastic and nonplastic species. Parsimony and Bayesian reconstructions demonstrate that plasticity is ancestral among the species studied. • CONCLUSIONS: Plasticity has been lost independently in sections Acanthophora and Lasiocarpa, and the consequence of its loss results in evolutionary diversification of sexual expression. In section Acanthophora, loss of plasticity represents a reversion to production of predominantly hermaphroditic flowers. In contrast, the fixed production of staminate flowers in Lasiocarpa has the hallmarks of evolution via genetic assimilation.

Out of America to Africa or Asia: inference of dispersal histories using nuclear and plastid DNA and the S-RNase self-incompatibility locus.

The plant genus Lycium (Solanaceae) originated in the Americas and includes approximately 85 species that are distributed worldwide. The vast majority of Old World species occur in southern Africa and eastern Asia. In this study, we examine biogeographic relationships among Old World species using a phylogenetic approach coupled with molecular evolutionary analyses of the S-RNase self-incompatibility gene. The phylogeny inferred from nuclear granule-bound starch synthase I (GBSSI), nuclear conserved ortholog set II (COSII) marker C2_At1g24360, and plastid spacer data (trnH-pbsA, trnD(GUC)-trnT(GGU), rpl32-trnL(UAG), and ndhF-rpl32) includes a clade of eastern Asian Lycium nested within the African species, suggesting initial dispersal from the Americas to Africa, with subsequent dispersal to eastern Asia. Molecular dating estimates suggest that these dispersal events occurred relatively recently, with dispersal from the Americas to Africa approximately 3.64 Ma (95% highest posterior density [HPD]: 1.58-6.27), followed by subsequent dispersal to eastern Asia approximately 1.21 Ma (95% HPD: 0.32-2.42). In accordance, the S-RNase genealogy shows that S-RNases isolated from Old World species are restricted to four lineages, a subset of the 14 lineages including S-RNases isolated from New World Lycium species, supporting a bottleneck of S-RNase alleles concomitant with a single dispersal event from the Americas to the Old World. Furthermore, the S-RNase genealogy is also consistent with dispersal of Lycium from Africa to Asia, as eastern Asian alleles are restricted to a subset of the lineages that also include African alleles. Such a multilocus approach, including complementary data from GBSSI, COSII, plastid spacer regions, and S-RNase, is powerful for understanding dispersal histories of closely related species.

Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation.

Unambiguous examples of ecological causation of sexual dimorphism are rare, and the best evidence involves sexual differences in trophic morphology. We show that moderate female-biased sexual dimorphism in bill curvature is the ancestral condition in hermit hummingbirds (Phaethornithinae), and that it is greatly amplified in species such as Glaucis hirsutus and Phaethornis guy, where bills of females are 60 per cent more curved than bills of males. In contrast, bill curvature dimorphism is lost or reduced in a lineage of short-billed hermit species and in specialist Eutoxeres sicklebill hermits. In the hermits, males tend to be larger than females in the majority of species, although size dimorphism is typically small. Consistent with earlier studies of hummingbird feeding performance, both raw regressions of traits and phylogenetic independent contrasts supported the prediction that dimorphism in bill curvature of hermits is associated with longer bills. Some evidence indicates that differences between sexes of hermit hummingbirds are associated with differences in the use of food plants. We suggest that some hermit hummingbirds provide model organisms for studies of ecological causation of sexual dimorphism because their sexual dimorphism in bill curvature provides a diagnostic clue for the food plants that need to be monitored for studies of sexual differences in resource use.

The utility of nuclear conserved ortholog set II (COSII) genomic regions for species-level phylogenetic inference in Lycium (Solanaceae).

The identification of genomic regions with sufficient variation to elucidate fine-scale relationships among closely related species is a major goal of phylogenetic systematics. However, the accumulation of such multi-locus data sets brings its own challenges, given that gene trees do not necessarily represent the true species tree. Using genomic tools developed for Solanum (Solanaceae), we have evaluated the utility of nuclear conserved ortholog set II (COSII) regions for phylogenetic inference in tribe Lycieae (Solanaceae). Five COSII regions, with intronic contents ranging from 68% to 91%, were sequenced in 10 species. Their phylogenetic utility was assessed and compared with data from more commonly used nuclear (GBSSI, nrITS) and cpDNA spacer data. We compared the effectiveness of a traditional total evidence concatenation approach versus the recently developed Bayesian estimation of species trees (BEST) method to infer species trees given multiple independent gene trees. All of the sampled COSII regions had high numbers of parsimony-informative (PI) characters, and two of the COSII regions had more PI characters than the GBSSI, ITS, and cpDNA spacer data sets combined. COSII markers are a promising new tool for phylogenetic inference in Solanaceae, and should be explored in related groups. Both the concatenation and BEST approaches yielded similar topologies; however, when multiple individuals with polyphyletic alleles were included, BEST was clearly the more robust approach for inferring species trees in the presence of gene tree incongruence.

Phylogenetic utility of nuclear nitrate reductase: a multi-locus comparison of nuclear and chloroplast sequence data for inference of relationships among American Lycieae (Solanaceae).

There has been considerable interest and research into the benefits of multiple low-copy nuclear regions for phylogenetic studies at low taxonomic levels. In this study, the phylogenetic utility of DNA sequence data from two low-copy nuclear genes, nitrate reductase (NIA) and granule-bound starch synthase I (GBSSI), was compared with data from nuclear ribosomal ITS and the cpDNA spacers trnT-trnF and trnD-trnT across 33 closely related taxa in tribe Lycieae (Solanaceae). The NIA data (introns 1-2) had the most parsimony-informative characters, with over twice the number provided by GBSSI, but NIA also had greater homoplasy. Although gene trees were generally concordant across the four regions, there were some notable areas of incongruence, suggesting both incomplete lineage sorting as well as possible reticulate origins.

A tale of two continents: Baker's rule and the maintenance of self-incompatibility in Lycium (Solanaceae).

Over 50 years ago, Baker (1955, 1967) suggested that self-compatible species were more likely than self-incompatible species to establish new populations on oceanic islands. His logic was straightforward and rested on the assumption that colonization was infrequent; thus, mate limitation favored the establishment of self-fertilizing individuals. In support of Baker's rule, many authors have documented high frequencies of self-compatibility on islands, and recent work has solidified the generality of Baker's ideas. The genus Lycium (Solanaceae) has ca. 80 species distributed worldwide, and phylogenetic studies suggest that Lycium originated in South America and dispersed to the Old World a single time. Previous analyses of the S-RNase gene, which controls the stylar component of self-incompatibility, have shown that gametophytically controlled self-incompatibility is ancestral within the genus, making Lycium a good model for investigating Baker's assertions concerning reproductive assurance following oceanic dispersal. Lycium is also useful for investigations of reproductive evolution, given that species vary both in sexual expression and the presence of self-incompatibility. A model for the evolution of gender dimorphism suggests that polyploidy breaks down self-incompatibility, leading to the evolution of gender dimorphism, which arises as an alternative outcrossing mechanism. There is a perfect association of dimorphic gender expression, polyploidy, and self-compatibility (vs. cosexuality, diploidy, and self-incompatibility) among North American Lycium. Although the association between ploidy level and gender expression also holds for African Lycium, to date no studies of mating systems have been initiated in Old World species. Here, using controlled pollinations, we document strong self-incompatibility in two cosexual, diploid species of African Lycium. Further, we sequence the S-RNase gene in 15 individuals from five cosexual, diploid species of African Lycium and recover 24 putative alleles. Genealogical analyses indicate reduced trans-generic diversity of S-RNases in the Old World compared to the New World. We suggest that genetic diversity at this locus was reduced as a result of a founder event, but, despite the bottleneck, self-incompatibility was maintained in the Old World. Maximum-likelihood analyses of codon substitution patterns indicate that positive Darwinian selection has been relatively strong in the Old World, suggesting the rediversification of S-RNases following a bottleneck. The present data thus provide a dramatic exception to Baker's rule, in addition to supporting a key assumption of the Miller and Venable (2000) model, namely that self-incompatibility is associated with diploidy and cosexuality.

Correlated evolution of fruit size and sexual expression in andromonoecious Solanum sections Acanthophora and Lasiocarpa (Solanaceae).

Andromonoecy is hypothesized to evolve as a mechanism enabling plants to independently allocate resources to female and male function. If staminate flower production is a mechanism to regulate allocation to female function (i.e., fruit production), then large-fruited species should be more strongly andromonoecious than smaller-fruited taxa because more resources are required to mature large fruit. We combined phylogenetically independent contrast analyses with extensive phenotypic characterization under common greenhouse conditions to examine the predicted relationship between fruit mass and the strength of andromonoecy among 13 species in Solanum sections Acanthophora and Lasiocarpa. The strength of andromonoecy, defined as the proportion of staminate flowers produced within inflorescences, was significantly and positively associated with fruit mass in both naïve and phylogenetically independent analyses. Our results are consistent with the hypothesis that andromonoecy functions as a mechanism to regulate allocation to female function and suggest that the strength of andromonoecy is also associated with resource limitation. In general, we find that strong andromonoecy appears to arise via reductions in hermaphroditic flower number. However, increases in staminate flowers have also contributed to transitions to strong andromonoecy in certain species. Finally, our analyses identified a suite of correlated characters (flower size, ovary width, fruit mass) that are associated with changes in the sexual expression of andromonoecy.

Association of ploidy and sexual system in Lycium californicum (Solanaceae).

In North American Lycium (Solanaceae), the evolution of gender dimorphism has been proposed as a means of restoring outcrossing after polyploidization causes the loss of self-incompatibility. Previous studies of this process in Lycium focused on comparisons between species that differ in ploidy. We examined intraspecific variation in floral morphology and DNA content in populations of L. californicum to determine correlations between sexual system and cytotype. We also used nuclear ITS and GBSSI sequence data to determine whether diploid and polyploid forms represent the same phylogenetic species, and the phylogeographic relationships among populations and ploidy levels. Within populations, no variation in ploidy was found, although among populations there was a perfect correspondence between sexual system and cytotype. Diploid populations were all hermaphroditic, whereas tetraploid populations were all gender dimorphic. There was no clear geographic pattern to the occurrence of diploid and tetraploid forms. Phylogenetic analysis confirms that L. californicum, regardless of ploidy, forms a monophyletic group within the genus Lycium. Sequences from diploid and polyploid individuals did not form reciprocally monophyletic clades, indicating either multiple gains of polyploidy, ongoing gene flow between cytotypes, or lack of lineage sorting since the evolution of polyploidy. The correspondence between ploidy and sex expression is consistent with the hypothesis that polyploidization triggers the evolution of gender dimorphism in this and other Lycium species.

Relationships within tribe Lycieae (Solanaceae): paraphyly of Lycium and multiple origins of gender dimorphism.

We infer phylogenetic relationships among Lycium, Grabowskia, and the monotypic Phrodus microphyllus, using DNA sequence data from the nuclear granule-bound starch synthase gene (GBSSI, waxy) and the chloroplast region trnT-trnF. This is the first comprehensive molecular phylogenetic study of tribe Lycieae (Solanaceae). In addition to providing an understanding of evolutionary relationships, we use the phylogenetic hypotheses to frame our studies of breeding system transitions, floral and fruit evolution, and biogeographical patterns within Lycieae. Whereas Lycium is distributed worldwide, Phrodus and the majority of Grabowskia species are restricted to South America. Tribe Lycieae is strongly supported as monophyletic, but Lycium likely includes both Grabowskia and Phrodus. Results also suggest a single dispersal event from the Americas to the Old World, and frequent dispersal between North and South America. The diversity of fruit types in Lycieae is discussed in light of dispersal patterns and recent work on fruit evolution across Solanaceae. Dimorphic gender expression has been studied previously within Lycium, and results indicate that transitions in sexual expression are convergent, occurring multiple times in North America (a revised estimate from previous studies) and southern Africa.

Architectural effects mimic floral sexual dimorphism in Solanum (Solanaceae).

Factors underlying apparent floral sexual dimorphism were examined in six species of andromonoecious Solanum section Lasiocarpa (Solanaceae). Both multivariate and univariate analyses show that hermaphroditic flowers are significantly larger than staminate flowers for all features measured. Thus, flowers could be characterized as sexually size dimorphic. However, when size variation due to flower position (architecture) is controlled experimentally, differences between the floral genders for the nongynoecial characters disappear; there is no difference in corolla or androecium size. Staminate flowers appear to be generally smaller than hermaphroditic flowers, not because of any difference related to primary sexual function, but because they tend to occur in the distal regions of each inflorescence. In contrast, significant differences between hermaphroditic and staminate flowers for primary female traits (ovary, style, and stigma) remain after controlling for position: the two floral types are truly dimorphic for these characters. We show that consideration of architectural effects can direct and refine hypotheses concerning the evolution of andromonoecy. More generally, if architectural effects on flower size are common among taxa with unisexual flowers, then these effects may contribute to the common perception of size dimorphism in taxa with unisexual flowers.

Floral morphometrics and the evolution of sexual dimorphism in Lycium (Solanaceae).

Plants of Lycium californicum, L. exsertum, and L. fremontii produce flowers that are either male-sterile (female) or hermaphroditic, and populations are morphologically gynodioecious. As is commonly found in gynodioecious species, flowers on female plants are smaller than those on hermaphrodites for a number of floral traits. Floral size dimorphism has often been hypothesized to be the result of either a reduction in female flower size that allows reallocation to greater fruit and seed production, or an increase in hermaphroditic flower size due to the increased importance of pollinator attraction and pollen export for hermaphroditic flowers. We provide a test of these two alternatives by measuring 11 floral characters in eight species of Lycium and using a phylogeny to reconstruct the floral size shifts associated with the evolution of gender dimorphism. Our analyses suggest that female flowers are reduced in size relative to the ancestral condition, whereas flowers on hermaphrodites have changed only slightly in size. Female and hermaphroditic flowers have also diverged both from one another and from ancestral cosexual species in several shape characteristics. We expected sexual dimorphism to be similar among the three dimorphic taxa, as gender dimorphism evolved only a single time in the ancestor of the American dimorphic lineage. While the floral sexual dimorphism is broadly similar among the three dimorphic species, there are some species-specific differences. For example, L. exsertum has the greatest floral size dimorphism, whereas L. fremontii had the greatest size-independent dimorphism in pistil characters. To determine the degree to which phylogenetic uncertainty affected reconstruction of ancestral character states, we performed a sensitivity analysis by reconstructing ancestral character states on alternative topologies. We argue that investigations such as this one, that examine floral evolution from an explicitly phylogenetic perspective, provide new insights into the study of the evolution of floral sexual dimorphism.

Diversification of andromonoecy in Solanum section Lasiocarpa (Solanaceae): the roles of phenotypic plasticity and architecture.

Quantitative analyses of sexual expression show extensive interspecific variation in the strength of andromonoecy (proportions of hermaphroditic and staminate flowers) among Solanum species in the monophyletic section Lasiocarpa. The roles of phenotypic plasticity and inter- and intra-inflorescence architecture in the diversification of andromonoecy within this small clade were analyzed. Four species that represent a range of expression of andromonoecy were examined. Staminate flowers produced within inflorescences ranged from 3% (S. candidum) to 7% (S. ferox) in weakly andromonoecious species and from 39% (S. pseudolulo) to 60% (S. quitoense) in more strongly andromonoecious species. Manipulation of fruit set on clonal replicates of multiple genotypes demonstrated variation among species for phenotypic plasticity. The strongly andromonoecious species, S. pseudolulo and S. quitoense, were not plastic and produced a large proportion of staminate flowers regardless of fruiting treatment, whereas S. candidum and S. ferox were phenotypically plastic and produced significantly more staminate flowers in the presence of developing fruit. Staminate flower production of all four species varied both within and among inflorescences. A greater proportion of staminate flowers were produced in distal (later produced) inflorescences. Within inflorescences, hermaphroditic flowers occurred in basal positions, whereas staminate flowers, when produced, occurred more distally. This pattern of staminate flower production is qualitatively the same in all species investigated; however, quantitative variation in the transition from hermaphroditic to staminate flower production within and among inflorescences is associated with variation in the strength of andromonoecy. At least three factors have contributed to the diversification of andromonoecy in section Lasiocarpa including the presence or absence of phenotypic plasticity in response to fruit set, quantitative variation in intra- and inter-inflorescence architectural effects, and total flower production.

The transition to gender dimorphism on an evolutionary background of self-incompatibility: an example from Lycium (Solanaceae).

Populations of three North American species of Lycium (Solanaceae) are morphologically gynodioecious and consist of male-sterile (i.e., female) and hermaphroditic plants. Marked individuals were consistent in sexual expression across years and male sterility was present throughout much of the species' ranges. Controlled pollinations reveal that L. californicum, L. exsertum, and L. fremontii are functionally dioecious. Fruit production in females ranged from 36 to 63%, whereas hermaphrodites functioned essentially as males. Though hermaphrodites were mostly male, investigation of pollen tube growth reveals that hermaphrodites of all dimorphic species were self-compatible. Self-fertilization and consequent inbreeding depression are commonly invoked as important selective forces promoting the invasion of male-sterile mutants into cosexual populations. A corollary prediction of these models is that gender dimorphism evolves from self-compatible ancestors. However, fruit production, seed production, and pollen tube number following outcross pollination were significantly higher than following self-pollination for three diploid, cosexual species that are closely related to the dimorphic species. The data presented here on incompatibility systems are consistent with the hypothesis that polyploidy disrupted the self-incompatibility system in the gynodioecious species leading to the evolution of gender dimorphism.