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.     

Active pollination favours sexual dimorphism in floral scent

Zoophilous flowers often transmit olfactory signals to attract pollinators. In plants with unisexual flowers, such signals are usually similar between the sexes because attraction of the same animal to both male and female flowers is essential for conspecific pollen transfer. Here, we present a remarkable example of sexual dimorphism in floral signal observed in reproductively highly specialized clades of the tribe Phyllantheae (Phyllanthaceae). These plants are pollinated by species-specific, seed-parasitic Epicephala moths (Gracillariidae) that actively collect pollen from male flowers and pollinate the female flowers in which they oviposit; by doing so, they ensure seeds for their offspring. We found that Epicephala-pollinated Phyllanthaceae plants consistently exhibit major qualitative differences in scent between male and female flowers, often involving compounds derived from different biosynthetic pathways. In a choice test, mated female Epicephala moths preferred the scent of male flowers over that of female flowers, suggesting that male floral scent elicits pollen-collecting behaviour. Epicephala pollination evolved multiple times in Phyllantheae, at least thrice accompanied by transition from sexual monomorphism to dimorphism in floral scent. This is the first example in which sexually dimorphic floral scent has evolved to signal an alternative reward provided by each sex, provoking the pollinator''s legitimate altruistic behaviour.     

Graft formation in Solanum pennellii (Solanaceae)

Three phases of cohesion were observable during the development of compatible autografts in Solanum pennellii. Phase I cohesion 1) lasted 4–5 d after grafting, 2) was characterized by an average increase in tensile strength of 4 g breaking weight (BW) mm^–2 graft area (GA) d^–1, and 3) correlated positively with cellular interdigitation at the graft interface. The fresh weight of the scion increased by approximately 5% d^–1 during the first 2 d after grafting. Phase II cohesion occurred 5–15 d after grafting, during which time 1) the tensile strength of the graft union increased by 14 g BW mm^–2 GA d^–1, 2) vascular differentiation across the graft interface was completed, and 3) the fresh weight of the scion increased by 9% d^–1. Phase III cohesion occurred subsequent to 15 d after grafting, during which time 1) the tensile strength of the graft union leveled off at a value similar to that of an ungrafted internode, and 2) the fresh weight of the scion increased by 14% d^–1. These results are discussed relative to mechanisms underlying the formation of compatible grafts.     

Sniffing out patterns of sexual dimorphism in floral scent

1.   A major transition in flowering plants has been the evolution of separate sexes (dioecy) from combined sexes (hermaphroditism). This transition is often, but not always, accompanied by the evolution of sexual dimorphism in attractive traits, and floral scent is no exception.
2.   In this review I aim to improve our understanding of variation in sexual dimorphism in floral scent characteristics by first explicating the relevant hypotheses, and then deriving explicit predictions for the pattern of floral scent from each.
3.   Next, I synthesize and qualitatively review published data on floral volatile emission rate and composition in 33 gender dimorphic species to identify emerging patterns, and evaluate these in light of predictions derived from the hypotheses.
4.   Although conclusions must be viewed as preliminary, a handful of strong patterns were revealed: (1) in the majority of the species studied males emit more volatiles per flower than females, (2) in over half of the species studied the sexes differed in at least one aspect of scent composition, (3) sexual dimorphism in scent composition was less common in species with rewardless females and/or brood site pollination than those that offered nectar and/or pollen, (4) a one-to-one correspondence between sex differences in overall scent composition and male organ-specific scent production is largely not found.
5.   This review has highlighted gaps in our understanding of the genesis of patterns of sexual dimorphism in floral scent, and makes clear that to move the field forward we need to shift our focus from pattern to process, and this will be best achieved by simultaneously testing alternative hypotheses at the same level of analysis.     

Karyotypic studies in Solanum section Lasiocarpa (Solanaceae)

Mitotic chromosomes of 13 species included in Solanum sect. Lasiocarpa were studied. All species have 2n = 24. The chromosome numbers of S. stagnale, S. felinum, and S. repandum are reported for the first time. Statistical analyses of chromosomes, genome length, and centromere position yielded estimates of karyotype composition and asymmetry. A generalized karyotype of the section shows that most of the chromosomes are metacentric (73%) or submetacentric (25.6%). Two pairs of subtelocentric chromosomes are present only in S. sessiliflorum. Satellites are quite common, characterizing chromosomes for ten species and are attached to the short arms of either m or sm chromosomes. They could be seen in 10% to 70% of the cells. Only S. pseudolulo bears two satellited chromosomes, one of them attached to a long chromosome arm. Although the section is chromosomally quite homogeneous, species can be distinguished by karyotype formula, presence of satellites in a particular chromosome pair, and total chromosome length. There are no indications of major chromosomal rearrangements within the section. Using cytological data exclusively, cluster analysis indicates S. sessiliflorum is isolated in the section. Solanum candidum and S. vestissimum are somewhat isolated as well and have unique karyological traits. Solanum pectinatum has a peculiar karyotype, but in the phenogram it is not particularly separated. Karyotype features suggest that morphological differentiation was not always followed by chromosomal divergence. The origin of the domesticated S. quitoense is no further elucidated by our data.     

Wind pollination, sexual dimorphism, and changes in floral traits of Schiedea (Caryophyllaceae)

Both changes in sex allocation and pollination mode may promote the separation of sexes in plant populations. Simultaneous evolution of wind pollination and dimorphism has occurred in Schiedea, where species with different female frequencies provide an opportunity to observe the effect of wind pollination on sex allocation and floral morphology. Differences among species in the ratio of anther to ovary volume were not the result of sex allocation trade-offs, but instead resulted from production of vestigial stamens in females; there were no changes in ovary volume in males and hermaphrodites (MH) of dimorphic species. Relative to hermaphroditic species, dimorphic species had more condensed inflorescences, a pattern often associated with wind pollination. Within dimorphic species, MH had longer filament lengths than females, and females had longer stigmas than MHs. These traits are characteristic of wind pollination, but there was no relationship between the degree of sexual dimorphism and female frequency. Ovary volume and ovule number and size had positive phenotypic correlations between females and MH of dimorphic species, making sex specialization more difficult. In dimorphic Schiedea species, selection for wind pollination may have a greater effect on floral traits than trade-offs in allocation between male and female function.     

A revision of Solanum section Brevantherum (Solanaceae)

The present revision of Solatium sect. Brevantherum Seithe recognizes 27 species in North and South America, two of which have become naturalized in parts of the Old World. Three species,S. lltisii, S. Goodspeedii, and S. appressum, are described as new. Comparative morphology, geographical distribution, and ecological data provide evidence for species recognition. Some of the most diagnostic features are those associated with the buds, leaves, and the hairs, which range from simple to porrect-stellate and more complex types, described herein as multangulate, dendritic-multangulate, echinoid, and dendriticechinoid.     

Seed coat structure and development in Solanum L. section Solanum (Solanaceae)

Seed coat structure in Solanum L. section Solanum (Solanaceae). Seed coat structure in Solanum L. section Solanum was examined using SEM. The seed coat patterns observed were fairly uniform throughout the section, and thus of little diagnostic use. Moreover, all species are characterized by their seeds being covered by hairlike structures, which are bands of lignified thickening in the lateral (and basal) walls of the outer epidermal cells of the testa, and which only become obvious when the seeds are wetted. The development of the seed coat and its characteristic thickening was followed from the unfertilized ovule to the mature seed in vouchered material of Solanum nigrum L. since this species is the generic type, using sections of freshly fixed material.     

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 na?ve 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.     

Features related to anther opening in Solanum species (Solanaceae)

The mode of anther opening and the morphological and histological variability of the stomium are described in 30 Solanum species. Poricidal, poricidal‐longitudinally dehiscing and longitudinally dehiscing anthers are observed. In the three types, the stomium may be diverse with regard to shape and histological characteristics before opening, but is always composed of small epidermal cells as the sole anther wall layer; the stomial cells may be differentiated only in part of the anther length. Particular crescent‐shaped structures in the epidermis, called ‘ridges’, are observed to line the stomium in most species. These ridges may be related to the stomium opening, working together with the cells with thickened walls of the anther. Cells with thickened walls are developed in the endothecium, middle layers and/or connective tissue at the apical end of the anther, surrounding the pore; only in the longitudinally dehiscing anthers of S. nitidum does an endothecium with thickened cell walls develop along its entire length. At least two histological features (the differentiation of small stomial epidermal cells as a unique layer, and the distribution of cells with thickened walls) seem to constrain the form of the open stomium. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 344–354.     

Dichogamy and sexual dimorphism in floral traits in the andromonoecious Euphorbia boetica

BACKGROUND AND AIMS: Euphorbia boetica (Euphorbiaceae) is a functional andromonoecious species that shows both intra- and interfloral dichogamy, hermaphrodite cyathia being protogynous. Sexual dimorphism of the cyathia of E. boetica is examined according to their gender and arrangement on the inflorescence. METHODS: Data were obtained from two natural populations, where the distribution of male and hermaphrodite cyathia in the inflorescence was recorded. The size, pollen production and viability, and nectar secretion were measured in both types of cyathia. KEY RESULTS: Most cyathia were male at the first levels of the inflorescence, then hermaphrodite cyathia predominated at the successive levels, although at the last levels the proportion of male cyathia increased. Male cyathia at basal positions lack ovaries, whereas those at distal positions showed vestigial ovaries. The size of the cyathia varied significantly depending on the level of the inflorescence where they were produced: those of the last levels were usually smaller. The hermaphrodites were significant bigger than males; however, these differences were due to the differential distribution of each cyathium type in the inflorescence. Male cyathia produced significantly more pollen and nectar than hermaphrodites. CONCLUSIONS: In Euphorbia boetica, basal male cyathia could be explained by the presence of protogyny, and apical male cyathia seem to respond to a preemption of resources. A true dimorphism affecting primary sexual characters and related to gender function appears at lower levels of the inflorescence, whereas an apparent size dimorphism due to positional effects occurs at upper positions. Longevity and distribution of cyathia, and their pattern of nectar production, could improve both male and female fitness.     

Observations on stylar heteromorphism in Solanum torvum Sw. (Solanaceae)

In several species of Solanum L. two forms of flower are noticeable in each inflorescence. In one form, the style is long and distinctly exserted; in the other, it is short and included within the conically connivent anthers, so that it is not visible from outside in an open flower. To this kind of difference in style length the term stylar heteromorphism has been applied here, just to distinguish it from heterostyly in the traditional sense. The phenomenon does not seem to have received much attention from botanists so far, and consequently rather little is known about its biological significance and possible evolutionary importance. The present report consists mainly of some observations made on stylar heteromorphism in one of these species, Solanum torvum , a shrubby weed widely distributed in the humid tropics.     

Correlated evolution of self-incompatibility and clonal reproduction in Solanum (Solanaceae)

It has been suggested that clonality provides reproductive assurance in cross-fertilizing species subject to pollen limitation, relieving one of the main selective pressures favoring the evolution of self-fertilization. According to this hypothesis, cross-fertilizing species subject to pollen limitation should often be clonal. Here, we investigated the association between clonality and a genetic mechanism enforcing outcrossing, self-incompatibility, in Solanum (Solanaceae). We collected self-incompatibility and clonality information on 87 species, and looked for an association between these two traits. To account for the contribution of shared evolutionary history to this association, we incorporated phylogenetic information from chloroplast (NADH dehydrogenase subunit F) sequence data. We found that self-incompatibility is strongly associated with clonal reproduction: all self-incompatible species reproduce clonally, while the absence of clonality is widespread among self-compatible taxa. The observed correlation persists after taking into account shared phylogenetic history, assumptions about the evolutionary history of self-incompatibility, uncertainty associated with phylogeny estimation, and associations with life history (annual/perennial). Our results are consistent with the hypothesis that clonality provides reproductive assurance, and suggest that the consequences of clonal growth in the evolution of plant reproductive strategies may be more significant than previously thought.     

AFLP markers support separation of Solanum nodiflorum from Solanum americanum sensu stricto (Solanaceae)

This study was aimed at examining the relationships between the African material of Solanum americanum (also designated as S. nodiflorum), accessions of this taxon from other geographical areas, and American S. americanum using AFLP markers. 96 individuals representing 39 accessions of S. americanum sensu lato and related diploid species from the widest possible geographical range, and one accession of S. dulcamara (as outgroup) were used. The AFLP results suggested that American S. americanum differs from S. nodiflorum and that the material investigated in this study can be assigned to three different species: S. americanum sensu stricto, S. nodiflorum and a Solanum species from Brazil. These species can be differentiated based on a combination of floral and fruit characteristics.     

Allozyme variation in the eggplant,Solanum melongena L. (Solanaceae)

Enzyme electrophoretic studies were made in cultivated Solanum melongena L. (eggplant) and similar wild and weedy forms, several of which have been thought to be different species/taxa. Twenty-nine accessions of S. melongena, 33 accessions of weedy forms (referred to as insanum) and 2 accessions of wild forms (referred to as incanum) were surveyed for 29 isozyme loci. In S. melongena, 22 of the 29 loci were monomorphic, and nearly all of its genes were either also monomorphic or in similar frequencies in insanum and incanum. The results demonstrate that the three taxa have a very close genetic relationship. The high genetic identities between them (0.913–0.967) suggests that they are conspecific even though they include extensive morphological diversity.     

Functional gametophytic self-incompatibility in a peripheral population of Solanum peruvianum (Solanaceae)

The transition from self-incompatibility to self-compatibility is a common transition in angiosperms often reported in populations at the edge of species range limits. Geographically distinct populations of wild tomato species (Solanum section Lycopersicon (Solanaceae)) have been described as polymorphic for mating system with both self-incompatible and self-compatible populations. Using controlled pollinations and sequencing of the S-RNase mating system gene, we test the compatibility status of a population of S. peruvianum located near its southern range limit. Pollinations among plants of known genotypes revealed strong self-incompatibility; fruit set following compatible pollinations was significantly higher than following incompatible pollinations for all tested individuals. Sequencing of the S-RNase gene in parents and progeny arrays was also as predicted under self-incompatibility. Molecular variation at the S-RNase locus revealed a diverse set of alleles, and heterozygosity in over 500 genotyped individuals. We used controlled crosses to test the specificity of sequences recovered in this study; in all cases, results were consistent with a unique allelic specificity for each tested sequence, including two alleles sharing 92% amino-acid similarity. Site-specific patterns of selection at the S-RNase gene indicate positive selection in regions of the gene associated with allelic specificity determination and purifying selection in previously characterized conserved regions. Further, there is broad convergence between the present and previous studies in specific amino-acid positions inferred to be evolving under positive selection.     

New species of Solanum section Geminata (Solanaceae) from South America

Fourteen new species ofSolanum sectionGeminata are described and illustrated. Eight of these,S. erosomarginatum, S. foetens, S. lasiocladum, S. lucens, S. ombrophilum, S. tanysepalum, S. tenuiflagellatum, andS. turgidum are from Venezuela.Solanum cucullatum andS. bellum are from Ecuador,S. abitaguense from Ecuador and northeastern Peru,S. amnicola andS. yanamonense from eastern Peru, andS. mapiricum from eastern Bolivia. Relationships of each of the species to others in its geographical area are discussed. Chromosome numbers are reported for ten of these new species.