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.     

Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

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.     

Fruit-Set Induced Changes in the Sex of Flowers in Caesalpinia calycina (Leguminosae)

Abstract: The proportions of hermaphrodite to staminate flowers in inflorescences of the andromonoecious species Caesalpinb calycina vary. Analysis of fruit position along the flowering rhachis, and also of the sex of floral buds in inflorescences with fruits set in different positions, indicate that fruiting success influences the sex of flowers in distal positions along the rhachis. Other reports of andromonoecy in caesalpinoid species are examined with reference to floral sex lability and the influence of fruit set.     

FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

Thrips pollination of androdioecious Castilla elastica (Moraceae) in a seasonal tropical forest

Androdioecy is a rare sexual system in nature, as predicted theoretically. Among the androecious species reported so far, Castilla elastica (Moraceae) is unique in that flowers are unisexual and staminate and pistillate flowers on cosexual plants are produced on different inflorescences. In addition, inflorescence structure of staminate inflorescences on males and staminate and pistillate inflorescences on cosexes is markedly different. Staminate inflorescences on males are bivalvate, while staminate inflorescences on cosexes are "fig-like" and urceolate. Pistillate inflorescences are discoidal. The difference may reflect different roles and requirements of the three inflorescences in pollination and protection from herbivores. This study reports thrips pollination of C. elastica, demonstrated by a pollinator introduction experiment. Thrips pollination of the species may be an example of mutualism originating from plant-herbivore interactions. In the Moraceae, shifts from simple herbivores on flowers to pollinators might have occurred many times, evolving into diverse pollination systems including the fig-fig wasp mutualism. The family, of which little is known about pollination systems, provides interesting and unique opportunities to study evolution of pollination systems and roles of nonpollinating associates of inflorescences.     

Contribution to the flower structure of Sararanga (Pandanaceae)

In Sararanga , the fruit is a berry as in Freycinetia. The testa comprises a lignifled outer integument with several cell layers, and an unlignified inner integument with two cell layers. Abortive fruits are frequent; they correspond to normal fruits that do not have carpels and sometimes have a lateral process that suggests an abortive carpel. The staminate flowers have a pistillode as in Freycinetia. The anther walls have 1–3 cell layers with endothecial thickenings, one layer in the distal part, 2–3 layers in the proximal part, as in Pandanus. Thus, within the family Pandanaceae, Sararanga has an intermediate position between Pandanus and Freycinetia. Generally speaking, there is a gradient in the vascularization of the bracts on the inflorescences: upper bracts are unvascularized, lower bracts vascularized. Anatomy suggests that the cupules are a perianth.     

New taxa and combinations in Protium Burm. f. Studies in neotropical Burseraceae VI

Revision ofProtium in eastern Amazonia and the Guianas yielded one new section, three new species, three new subspecies, and five new combinations in the genus. SectionSarcoprotium Daly is distinguished principally by the cucullate petals with clawed bases and usually with a dorsal thickening produced vertically as an appendage; staminate inflorescences with a few 3-merous flowers; thecae of the anthers completely separated by the connective and the dehiscence latero-introrse; and fruit ellipsoid when only one pyrene develops.     

Morphological and cytological development and starch accumulation in hermaphrodite and staminate flowers of olive (Olea europaea L.)

In olive (Olea europaea L.), the formation of functionally staminate flowers rather than fully functional hermaphrodites is one of the major factors limiting fruit set, as flowers with aborted pistils are incapable of producing fruit. Studies conducted on various angiosperm species have shown a correlation between flower abortion and starch content. Thus, it is important to know if starch content plays a role in regulating pistil development in olive and if so, what mechanism regulates starch distribution. Cyto-histological observations of staminate and hermaphrodite olive flowers show that pistil development in staminate flowers is interrupted after the differentiation of the megaspore mother cell. At that stage, starch grains were only detected in the ovary, style and stigma of the hermaphrodite flowers. No starch was observed in the pistils of the staminate flowers. This finding suggests a tight correlation between starch content and pistil development. The secondary origin of starch within the flower is indicated by low chlorophyll content in the gynoecium, undetectable Rubisco activity in the pistils of these two kinds of flowers and by the ultrastructure of the plastids observed by transmission electron microscope analysis. The plastids have few thylakoid membranes and grana and in the staminate flowers appeared very similar to proplastids. Considering differences in starch content between staminate and hermaphrodite flowers and the secondary origin of the starch, differences in pistil development in the staminate and hermaphrodite flowers could be related to differences in the sink strength of these two types of flowers.     

细枝柃性别表达特性及资源分配

已有的资料将柃木属(Eurya)描述为严格的雌雄异株植物, 性别变异现象极为少见。目前仅在柃木(E. japonica)和钝叶柃(E. obtusifolia)等少数种类中报道过两性花的存在。近几年笔者发现细枝柃(E. loquaiana)存在性别变异现象, 性别变异株上具有不同性别类型的花。该文从单花和植株水平分析了细枝柃的性别表达特性, 并对不同类型花的花部构件生物量分配进行比较分析。结果表明, 细枝柃具有6种类型的花, 从单花水平上看, 细枝柃性别有雌性、雄性及两性3种类型; 细枝柃性别在植株水平上体现较为复杂, 有雌株, 雄株, 雌花和两性花同株, 雄花和两性花同株, 雌雄异花同株及雌花、雄花、两性花同株6种类型; 在细枝柃花部构件生物量分配中, 雄花(包括雄株花和变异株雄花)花部构件生物量分配中雄蕊生物量的分配低于雌花(包括雌株花和变异株雌花)中雌蕊生物量的分配; 两性花中, 雄蕊生物量分配低于雌蕊, 这是其优化资源分配的手段, 进而获取最大适合度收益。     

Neither insects nor wind: ambophily in dioecious Chamaedorea palms (Arecaceae)

Pollination of Neotropical dioecious trees is commonly related to generalist insects. Similar data for non‐tree species with separated genders are inconclusive. Recent studies on pollination of dioecious Chamaedorea palms (Arecaceae) suggest that species are either insect‐ or wind‐pollinated. However, the wide variety of inflorescence and floral attributes within the genus suggests mixed pollination mode involving entomophily and anemophily. To evaluate this hypothesis, we studied the pollination of Chamaedorea costaricana, C. macrospadix, C. pinnatifrons and C. tepejilote in two montane forests in Costa Rica. A complementary morphological analysis of floral traits was carried out to distinguish species groups within the genus according to their most probable pollination mechanism. We conducted pollinator exclusion experiments, field observations on visitors to pistillate and staminate inflorescences, and trapped airborne pollen. A cluster analysis using 18 floral traits selected for their association with wind and insect pollination syndromes was carried out using 52 Chamaedorea species. Exclusion experiments showed that both wind and insects, mostly thrips (Thysanoptera), pollinated the studied species. Thrips used staminate inflorescences as brood sites and pollinated pistillate flowers by deception. Insects caught on pistillate inflorescences transported pollen, while traps proved that pollen is wind‐borne. Our empirical findings clearly suggest that pollination of dioecious Chamaedorea palms is likely to involve both insects and wind. A cluster analysis showed that the majority of studied species have a combination of floral traits that allow for both pollination modes. Our pollination experiments and morphological analysis both suggest that while some species may be completely entomophilous or anemophilous, ambophily might be a common condition within Chamaedorea. Our results propose a higher diversity of pollination mechanisms of Neotropical dioecious species than previously suggested.     

Thrips pollination of the dioecious ant plant Macaranga hullettii (Euphorbiaceae) in Southeast Asia

Discussion about thrips (Thysanoptera) as main pollinators has been controversial in the past because thrips do not fit the preconception of an effective pollinator. In this study, we present evidence for thrips pollination in the dioecious pioneer tree genus Macaranga (Euphorbiaceae). Macaranga hullettii is pollinated predominantly by one thrips species, Neoheegeria sp. (Phlaeothripidae, Thysanoptera). As a reward for pollinators, the protective floral bracteoles function as breeding sites for thrips and trichomal nectaries on the adaxial surface of the floral bracteoles provide alimentation. Flowering phenology of both staminate and pistillate trees was highly synchronized within 3-4 wk periods. In contrast to pistillate trees, staminate trees start to breed the thrips inside the developing inflorescences ～2 wk before anthesis. Breeding of Neoheegeria sp. in the laboratory indicates that the thrips development is completed within ～17 d. Thus, staminate trees offer breeding sites for one thrips generation until the onset of pollen presentation. Intraspecific pollen transfer by thrips was proved by pollen loads of thrips taken from receptive pistillate inflorescences of M. hullettii. Bagging experiments of different mesh sizes showed that seed set reached almost the level of open-pollinated flowers when exclusively tiny insects like thrips were able to enter the net bags, but no apomictic seed set occurred when no insect access was given to the flowers.     

中国蜘蛛抱蛋属植物一新记录种——垂花蜘蛛抱蛋

报道了产自云南屏边的中国蜘蛛抱蛋属一新记录种——垂花蜘蛛抱蛋（Aspidistra nutans Aver.&Tillich）。该种是2017年发表的新种,采集于越南西北部地区,原文缺少果实描述,现补充描述该种果的形态特征。本种与辐花蜘蛛抱蛋（A.subrotata Y.Wan&C.C.Huang）在形态上相近,但其叶片中脉和侧脉明显突出,花梗近先端向下弯曲,长0.8~4.5 cm,花下垂,常贴生于地面,花被紫褐色,常平展,柱头纯白色（后者叶脉凹陷,花梗直立,长约5 mm,花斜向上开,花被暗紫色,内面具有小乳突,柱头白色带紫色斑点）而显著有别于后者。     

臭椿花器官分化的初步研究

利用扫描电镜(SEM)和光镜(LM)对臭椿花序及花器官的分化和发育进行了初步研究,表明:1)臭椿花器官分化于当年的4月初,为圆锥花序;2)分化顺序为花萼原基、花冠原基、雄蕊原基和雌蕊原基。5个萼片原基的发生不同步,并且呈螺旋状发生;5个花瓣原基几乎同步发生且其生长要比雄蕊原基缓慢;雄蕊10枚,两轮排列,每轮5个原基的分化基本是同步的;雌蕊5,其分化速度较快;3)在两性花植株中,5个心皮顶端粘合形成柱头和花柱,而在雄株中,5个心皮退化,只有雄蕊原基分化出花药和花丝。本研究着重观察了臭椿中雄花及两性花发育的过程中两性花向单性花的转变。结果表明,臭椿两性花及单性花的形成在花器官的各原基上是一致的(尽管时间上有差异),雌雄蕊原基同时出现在每一个花器官分化过程中,但是,可育性结构部分的形成取决于其原基是否分化成所应有的结构:雄蕊原基分化形成花药与花丝,雌蕊原基分化形成花柱、柱头和子房。臭椿单性花的形成是由于两性花中雌蕊原基的退化所造成,其机理有待于进一步研究。     

Observations of inflorescence patterns and flower sex in a population of Sagittaria papillosa Buch. (Alismataceae)

Sagittaria papillosa Buch. is monoecious with unisexual flowers, pistillate below, staminate above, typically with an unbranched scape. A large population with unusual numbers of staminate and bisexual flowers on the lowest whorl of the inflorescence and many particles was quantitatively evaluated. First-formed inflorescences had more staminate and bisexual flowers than those produced later. Branched scapes were predominantly found to be the second inflorescence produced by a given plant. Genetic crosses between flowers on recemes and panicles produced no branched inflorescences. When grown under greenhouse conditions all tested plants had racemes with pistillate flowers in the lower whorls and staminate ones above. Data from soil parameters, daylengths and air temperatures are compared to reported information on modification of flower sexuality by these factors.     

Floral anatomy in Dypsis (Arecaceae–Areceae): a case of complex synorganization and stamen reduction

Male (staminate) flowers of Dypsis possess either six or three stamens and/or staminodes, in contrast to most other palms, in which the basic stamen number is six (although polyandry is also common in palms). Significant variation among the tristaminate forms, both in stamen morphology and stamen position with respect to the perianth parts, indicates that stamen reduction from six to three has occurred more than once, and possibly several times within Dypsis. A few species include teratological forms with zygomorphic flowers; for example, Dypsis lantzeana normally possesses three antesepalous stamens, but in some specimens only the median outer (abaxial) stamen is expressed, perhaps indicating incipient zygomorphy correlated with complex synorganization. Inclusion of earlier historic genera such as Neophloga and Chrysalidocarpus within a broadly circumscribed Dypsis appears to be justified, although the informal taxonomic groupings within Dypsis require review, in particular the taxonomic significance of the different types of anther morphology. The discovery here that adnation of staminodes to the pistillode (complex synorganization) occurs in species other than D. mirabilis opens further questions about the taxonomic utility of this character in Dypsis, in which stamen/staminode development and adnation are apparently unusually labile. Such androecial–gynoecial adnation is otherwise rare in palms, as also in other monocots, in which probably the best‐known example occurs in orchids. Septal nectaries are present in some, but not all, staminate flowers in species of Dypsis. Dypsis bejofo is exceptional in that in staminate flowers the pistillode is distally bulbous and bears three prominent modified supralocular septal nectaries. Female (pistillate) flowers in Dypsis are syncarpous, normally pseudomonomerous (as in many other Arecoideae), and possess septal nectaries that effectively delimit the carpel margins and indicate insect pollination. There is a central solid transmitting tissue that extends from the placenta to three stylar canals. The stylar canals of the two sterile carpels are apparently functional, in addition to that of the fertile carpel. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 115?133.     

A new species of Myrcia (Myrteae, Myrtaceae) from the Atlantic Forest of southern Bahia, Brazil

Myrcia thomasii, a new species from the Atlantic Forest of southern Bahia, is described and illustrated. The new species is similar to M. micropetala, but differs in having the inflorescence and basal bracts puberulent, lanceolate bracts at the base of flowers, longer calyx lobes and petals, and a 3-locular ovary.     

An ecologically significant new species of <Emphasis Type="Italic">Dacryodes</Emphasis> from the northern Andes. Studies in neotropical Burseraceae XV

Dacryodes uruts-kunchae is described, illustrated, and contrasted with D. steyermarkii. It is from white-sand plateaus and slopes of the sub-Andean cordilleras east of the Eastern Cordillera of the Andes in Ecuador, and from eastern foothills of the Andes in Peru. The new species is unusual for a number of reasons: (1) relatively few Burseraceae are found in tropical pre-montane and montane habitats; (2) it is dominant where it occurs, a rare phenomenon for Neotropical Dacryodes; (3) its staminate flowers show three stages in the transition from disk+pistillode to “ovariodisk”; and (4) it has relatively small, densely glandular-punctate, sclerophyllous leaflets.