Mating system and fine-scale spatial genetic structure of Solanum lycocarpum St.Hil. (Solanaceae) in the Brazilian Cerrado

Restricted gene dispersion – resulting from both self-pollination and limited capability of pollen migration, as well as seed dispersion at short distances – has been considered the main reason for spatial genetic structuring in plant populations. This study evaluated the intrapopulation genetic structure and the mating system in four populations of Solanum lycocarpum, a woody bush occurring in Brazilian Cerrado vegetation. Two hundred and twenty-four individuals were genotyped through five nuclear SSR loci (30 alleles) and six cpSSR loci (82 haplotypes). The study evidenced that the species mates predominantly by outcrossings , that biparental inbreeding is not common, and that there are almost 10 trees participating as pollen donors per mother-tree. The populations were formed by many mother lineages, indicating efficient seed dispersion by the fauna and the occurrence of multiple foundation events. Spatial genetic structure was observed in three populations (average Sp=0.0184 ± 0.0030) and it resulted from both restricted seed dispersion and from vegetative reproduction. During the collection of seeds for ex situ conservation, seeds must be gathered from 150 to 200 mother trees, so that the effective size of 500 individuals is retained. The sampling must comprise the biggest possible number of populations in a wide area to enable the maintenance of the biggest possible haplotypic diversity.     

Floral biology,reciprocal herkogamy and breeding system in four Psychotria species (Rubiaceae) in Brazil

Distylous species have two floral morphs with reciprocal positions of sexual whorls, functioning to promote cross‐pollination. Additionally, most distylous species have an incompatibility system linked to the discrete variation of anthers and stigmas. Here we provide new data on heterostyly, reproductive biology and pollination in four Psychotria spp. from the Cerrado Region (Brazil). Psychotria deflexa, P. nitidula and P. trichophoroides fitted the distylous morphological syndrome, whereas P. prunifolia was monomorphic and self‐compatible. Reciprocity varied across species, with the upper whorl of anthers and stigmas being more reciprocal than the lower whorl. Psychotria nitidula has a heteromorphic incompatibility system, whereas P. deflexa and P. trichophoroides showed partial self‐compatibility. Psychotria prunifolia and P. trichophoroides were visited by large bees and exhibited lower reproductive efficacy than P. nitidula and P. deflexa, which are visited by wasps and small bees. Our results provide new evidence that the morphological distylous syndrome does not always co‐occur with heteromorphic incompatibility and additional mechanisms (e.g. disassortative pollen transfer) may operate to maintain the polymorphism and isoplethy of species with different incompatibility system.     

Genetic structure based on nuclear and chloroplast microsatellite loci of Solanum lycocarpum A. St. Hil. (Solanaceae) in Central Brazil

Solanum lycocarpum (Solanaceae) is a woody species found in the Brazilian Cerrado. The flowers are pollinated by Xylocopa spp bees, and seeds are dispersed by mammals with distinct home range sizes. As a consequence, relative contributions of pollen and seeds to overall gene flow can vary according to different spatial scales. We studied the genetic structure of four natural populations of S. lycocarpum separated by 19 to 128 km, including individuals located along dirt roads that interlink three of the populations. A total of 294 individuals were genotyped with five nuclear and six chloroplast microsatellite markers. Significant spatial genetic structure was found in the total set of individuals; the Sp statistic was 0.0086. Population differentiation based on the six chloroplast microsatellite markers (θ(pC) = 0.042) was small and similar to that based on the five nuclear microsatellite markers (θ(p) = 0.054). For this set of populations, pollen and seed flow did not differ significantly from one another (pollen-to-seed flow ratio = 1.22). Capability for long distance seed dispersion and colonization of anthropogenic sites contributes to the ability of S. lycocarpum to maintain genetic diversity. Seed dispersion along dirt roads may be critical in preserving S. lycocarpum genetic diversity in fragmented landscapes.     

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.     

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.     

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.     

Four o'clock pollination biology: nectaries,nectar and flower visitors in Nyctaginaceae from southern South America

Floral nectary structure and nectar sugar composition were investigated in relation to other floral traits and flower visitors in contrasting species of Nyctaginaceae from southern South America, representing four tribes (Bougainvilleeae, Colignonieae, Nyctagineae, Pisoneae). Our comparative data will aid in the understanding of plant–pollinator interactions and in the development of hypotheses on the origin of floral and reproductive characters in this family. The nectaries are located on the inner side of the staminal tube. The nectariferous tissue is composed of an epidermis and three to ten layers of secretory parenchymal cells, supplied indirectly by the filament vascular bundles. Stomata appear to be associated with nectar secretion. For the first time in Nyctaginaceae, nectary ultrastructure is described in Boerhavia diffusa var. leiocarpa. Nectary parenchyma cells are densely cytoplasmic and contain numerous starch grains. Plasmodesmata connect the nectariferous cells. Flowers of Nyctaginaceae secrete a small volume of nectar of variable concentration (10–47%). Nectar is dominated by hexoses, but Mirabilis jalapa showed a balanced proportion of sucrose and hexoses. Hymenoptera are the most common visitors for most species; nocturnal Lepidoptera are the most common visitors for M. jalapa and Bougainvillea stipitata. We found relatively low variation in the nectary characteristics of Nyctaginaceae compared with broad variation in flower structure, shape, colour and nectar traits. © 2013 The Linnean Society of London     

Floral trait evolution associated with shifts between insect and wind pollination in the dioecious genus Leucadendron (Proteaceae)

Transitions between animal and wind pollination have occurred in many lineages and have been linked to various floral modifications, but these have seldom been assessed in a phylogenetic framework. In the dioecious genus Leucadendron (Proteaceae), transitions from insect to wind pollination have occurred at least four times. Using analyses that controlled for relatedness among Leucadendron species, we investigated how these transitions shaped the evolution of floral structural and signaling traits, including the degree of sexual dimorphism in these traits. Pollen grains of wind‐pollinated species were found to be smaller, more numerous, and dispersed more efficiently in wind than were those of insect‐pollinated species. Wind‐pollinated species also exhibited a reduction in spectral contrast between showy subtending leaves and background foliage, reduced volatile emissions, and a greater degree of sexual dimorphism in color and scent. Uniovulate flowers and inflorescence condensation are conserved ancestral features in Leucadendron and likely served as exaptations in shifts to wind pollination. These results offer insights into the key modifications of male and female floral traits involved in transitions between insect and wind pollination.     

广东地区外来种五爪金龙的传粉生物学研究（英文）

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006-2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花,花序为聚伞花序,花冠为紫色或蓝紫色喇叭状。开花时间在早晨4:30-5:20之间,下午17:40闭合,持续大约12h。每朵花平均分泌花蜜量为5.82±2.47μL(n=10),花蜜平均含糖量为41.03±6.52%,氨基酸含量为0.042mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18±264.58(n=10)。木蜂(Xylocopasp.)是其有效的传粉昆虫,蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触,有利于自花授粉。然而,繁育系统的研究表明五爪金龙是自交不亲和植物,其种子仅能通过异花授粉获得。     

Reproductive biology and pollination of Utricularia reniformis A.St.‐Hil. (Lentibulariaceae)

Utricularia reniformis is an endemic Brazilian carnivorous plant, most common in high‐altitude grasslands. Knowledge of the reproductive biology of U. reniformis is essential for planning conservation strategies, but it is currently poorly understood. Thus, we studied the floral morphology, floral biology, breeding system and pollination of this species. U. reniformis produces and stores nectar in the flower spur, a classic feature of bee‐pollinated flowers, and we recorded Xylocopa sp. and Bombus sp. as pollinators. Moreover, although it is self‐compatible it is an obligate animal‐pollinated species, as the sensitive stigma avoids self‐pollination. However, in natural conditions reproductive success is low due to the rarity of visits from pollinators. We suggest that the low reproductive success caused by a deficit of pollinators may affect gene flow, causing loss of genetic diversity in U. reniformis.     

Dormancy and the influence of photoperiod and temperature on sexual maturity in Nicrophorus nepalensis (Coleoptera: Silphidae)

Abstract Carrion beetles (Nicrophorus spp.) use small vertebrate carcasses for food and reproduction. Their ecology and behaviors are highly affected by the availability of carcasses and the surrounding environmental conditions. Our results revealed that in subtropical Fushan, northern Taiwan, N. nepalensis was mainly active in spring (February to May), and could also be found in autumn (October and November); but there was no capture record in summer (June to September) and winter (December and January). A laboratory temperature tolerance study indicated that N. nepalensis adults become inactive at temperatures above 26°C, and had the highest mortality when the temperature was raised from 27°C to 28°C. Furthermore, N. nepalensis became sexually mature at 20°C, depending on the photoperiod: the longer the day, the lower the percentage of sexually mature 2‐week‐old females after emergence. In another experiment, N. nepalensis virgins were paired under three possible conditions at Fushan. At 15°C and 20°C, if carcasses were presented to the pairs within 3 days after emergence, all laid eggs in the second week after emergence. If carcasses were presented 1 week after emergence, most began to reproduce at 20°C with 12.5 h of daylight. However, at 15°C with 11 h of daylight, the carrion beetles hibernated first, and reproduced in the ninth week after emergence. At 25°C with 14 h of daylight, carrion beetles did not bury the mouse carcasses, the females did not lay eggs, and the adult lifespan was only one‐third of that at 20°C. This study revealed that both photoperiod and temperature influence the time needed to reach the sexual maturity of N. nepalensis; and also implied that the narrow temperature tolerance range and dormancy behavior of carrion beetles are highly regulated by those environmental factors.     

Breeding system and thrips (Thysanoptera) pollination in the endangered tree Ocotea porosa (Lauraceae): implications for conservation

Ocotea porosa has been extensively exploited over the past few decades because of the quality of its wood. Today, populations are reduced and the species is now included in the Red List of threatened species by the International Union for Nature Conservation. For conservation and management purposes, it is extremely important that we understand its reproductive ecology. Floral morphology was described based on field and scanning electron microscopic examination. The reproductive system was determined through experimentally controlled pollination along with observations of pollen tube growth. Pollinators were identified through field observations. Flowers of O. porosa are small, shallow, inconspicuous, asynchronous and grouped in inflorescences. This species presented self‐compatibility, but did not reproduce through apomixy and spontaneous self‐pollination was very rare (5%). Despite being monoclinal, flowers were protogynic, and the gynoecium was receptive after the first day of anthesis when the anthers were closed. Spontaneous self‐pollination was avoided by the extrorse position of the anthers of the internal stamens. Frankliniella gardeniae (Thysanoptera) was the only pollinating species observed and, after visiting several asynchronous flowers in the same inflorescence, favors geitonogamy. Together the small distance supposedly achieved by thrips in flight and the small population density of O. porosa can reduce the chances of cross‐pollination in this species. If so, conservation measures must include preservation of the current population and possibly planting to increase population density. This would reduce the distance between individuals and increase genetic variability. Thrips as pollinators must be included in conservation planning for O. porosa.     

Random amplified polymorphic DNA variation in the eggplant,Solanum melongena L. (Solanaceae)

RAPD analysis was carried out on 52 accessions of Solanum melongena (eggplant) and related weedy forms known as insanum. Twenty-two primers amplified 130 fragments. Solanum melongena exhibited 117 of the fragments, all of which were also present in insanum. Insanum displayed an additional 13 fragments not found in S. melongena. Overall, the insanum accessions were more diverse than those of S. melongena. The calculated similarity between them was 0.947. The RAPD results were closely concordant with the results of an electrophoretic isozyme survey performed on the same accessions. The concordance of the results shows that even though S. melongena and insanum are highly diverse morphologically, it is no longer appropriate to distinguish them taxonomically.     

马先蒿属植物花冠分化与繁殖适应的研究进展

结合已有的研究报道和作者近年来的工作,对马先蒿属（Pedicularis）植物的花冠多样化成因与繁殖适应特性进行了总结和探讨。通过对该属4种进化花冠型的花器官发生和分化的研究发现,花部各器官在发生和发育初期基本一致,后期上唇形态的分化是导致成熟花形态结构产生较大差异的重要阶段。孢粉学研究认为,花冠类型与花粉萌发孔类型之间具有显著相关性;萌发沟的演化可能与繁殖适应有一定的关系。分子系统学研究表明,多样化的花冠类型在不同的谱系内经过若干次的独立进化而表现出了高度的平行演化（parallelism）。传粉生物学研究证实,该属植物花冠多样化与其主要传粉者熊蜂属（Bombus）昆虫的传粉行为存在较为密切的关系。具有相同（似）花冠类型的马先蒿可能被同种或不同种的熊蜂以相同的方式访问,但在花粉落置位置上存在显著差异,这可能有助于同域分布重叠的物种间在生殖上的机械隔离,而花冠的分化在一定程度上促进了新的物种形成。     

Histomorphological and endocrine assessment of female Arabian carpetshark,Chiloscyllium arabicum (Elasmobranchii: Hemiscylliidae) from the Persian Gulf during annual reproductive cycle

The present study aimed at assessing the annual reproductive cycle of female Arabian carpetshark, Chiloscyllium arabicum from the Persian Gulf by a macroscopic and microscopic evaluation of the reproductive tract. The annual cycle of gonadal steroids [17β-estradiol (E2), progesterone (P4) and testosterone (T)] was also assessed in this shark. In total, 130 female C. arabicum were collected from the Bahrakan Creek (located northwest of the Persian Gulf) between January 2018 and March 2019. Females were oviparous with an external-type ovary and only one functional ovary. Five sexual maturity stages were recognized based on macroscopic and microscopic evaluation: Immature I (August–October), Immature II (November–January), Mature (February–March), Pregnant (April–May) and Spent (June–July). The structural changes in the oviducts, oviducal glands and uterus throughout the annual reproductive cycle were consistent with their roles in the egg movement, the egg capsule production and sperm storage. The plasma levels of the gonadal steroids were associated with morphological changes in the reproductive tract. E2 showed two detectable peaks during March (close to ovulation) and June (just before mating). P4 and T displayed a peak just before ovulation.     

Density affects gametophyte growth and sexual expression of Osmunda cinnamomea (Osmundaceae: Pteridophyta)

BACKGROUND AND AIMS: To understand how gametophyte densities affect the sexual expression and sizes of Osmunda cinnamomea and to provide information on the density of growth needed to favour successful reproduction, fresh spores were sown at various densities and subsequent gametophyte growth was studied. METHODS: Spores were sown and cultured in the laboratory. Subsequent gameophytes at different population densities were sampled and their sexual expression and sizes were recorded. KEY RESULTS: One-year-old multispore cultures of the fern O. cinnamomea demonstrated that population density affected gametophyte growth and sexual expression. As density increased, gametophytes became significantly smaller and more slender. Female and asexual gametophytes dominated in populations of low and high densities, respectively. At intermediate population densities, hermaphroditic and male gametophytes were dominant. Female gametophytes were larger than gametophytes of all other types. Hermaphroditic gametophytes were larger than male gametophytes, which were larger than asexual gametophytes. Large gametophytes were wide-cordate, whereas smaller ones tended to be narrow-spathulate. CONCLUSIONS: Gametophyte size of O. cinnamomea is negatively related to the population density, which significantly affects gametophytes' sexual expression. The presence of unisexual and bisexual gametophytes at intermediate densities indicates that both intergametophytic and intragametophytic selfing may occur.     

广东地区外来种五爪金龙的传粉生物学研究

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006–2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花, 花序为聚伞花序, 花冠为紫色或蓝紫色喇叭状。开花时间在早晨4: 30–5: 20之间, 下午17: 40闭合, 持续大约12 h。每朵花平均分泌花蜜量为5.82 ± 2.47 μL (n = 10), 花蜜平均含糖量为41.03 ± 6.52%, 氨基酸含量为0.042 mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18 ± 264.58 (n = 10)。木蜂(Xylocopa sp.)是其有效的传粉昆虫, 蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触, 有利于自花授粉。然而, 繁育系统的研究表明五爪金龙是自交不亲和植物, 其种子仅能通过异花授粉获得。     

Spatio-temporal variation of natural and sexual selection in breeding populations of the coreid bug,Colpula lativentris (Heteroptera: Coreidae)

Spatio-temporal variations of lifetime reproductive succes (LRS) of both male and female individuals of a coreid bugColpula lativentris were measured and analyzed using the multiple regression method of Arnold and Wade (1984a, b). The standardized variance of LRS was larger in males than that in females as males often to secure mates for a long period whereas females could easily find mates and oviposit simply dependent on ovarial maturation. LRS was partitioned into 4 consecutive fitness components: (1) reproductive lifespan, (2) copulating efficiency, (3) guarding efficiency (for males) or oviposition efficiency (for females), and (4) number of eggs per clutch. In males copulating efficiency was the largest determining factor of LRS, whereas in females reproductive lifespan was the most important factor. Such tendencies were stable on both a yearly and local basis. Patterns of relative contribution of natural selection (reproductive lifespan and number of eggs per clutch) and sexual selection (copulating efficiency and guarding or oviposition efficiency) to LRS were clearly different between males and females. This sexual difference is, at least to some extent, thought to be brought about by sexual selection among males for mating opportunity, though no physical fight was observed among males. Directional selection on body length was found only in relation to the clutch size of females because large females tended to lay larger clutches. No significant directional selection was found in other fitness components.     

How do sexual expression,reproductive phenology and reproductive success relate to sexual systems in Fissidens Hedw. (Fissidentaceae)? A case study comparing two different sexual systems in mosses

• Reproductive performance is known to differ between co‐sexual and non‐co‐sexual species. Thus, our aim was to determine whether: (i) the distance between sex structures is negatively associated with sex expression; (ii) male gametangia take longer to mature in rhizautoicous species than in gonioautoicous species; and (iii) the gonioautoicous sexual system has greater reproductive success (i.e. percentage of ramets with sporophyte) than the rhizautoicous sexual system.
• One population each of the mosses Fissidens scariosus and F. submarginatus, rhizautoicous and gonioautoicous, respectively, in a remnant of Atlantic Forest in Brazil were sampled monthly from September 2016 until August 2017. The number and phenophases of antheridia, archegonia and sporophytes were analysed for each species. Sexual expression and reproductive success were calculated, and reproductive phenology compared across environmental variables.
• As expected, sexual expression was significantly higher for the gonioautoicous species, which produced antheridia throughout the year and archegonia over many months, while gametangia production by the rhizautoicous species occurred only during the rainy season. Mean number of gametangia per perigonium and perichaetium were slightly higher for the rhizautoicous species (6.84 antheridia; 11.38 archegonia) than for the gonioautoicous species (4.39; 7.62). Gametangia and sporophyte production in the rhizautoicous species were markedly seasonal compared to that of the gonioautoicous species, although reproductive success did not differ.
• Therefore, we infer that the rhizautoicy (i.e. a functional dioicy) promotes lower expression of gametangia compared to gonioautoicy but is more efficient and so obtains the same reproductive success.

     

Morphometric analysis of floral variation in the Pyrola picta species complex (Ericaceae): interpretation and implications for ecological and phylogenetic differentiation

The Pyrola picta species complex of western North America comprises four species (P. picta, P. dentata, P. aphylla and P. crypta) that grow sympatrically in some parts of their collective ranges, have remarkably similar flowers and share pollinators. These species do not exhibit the genetic signatures typical of random or heterospecific mating, but instead show genetic divergence patterns indicating that they maintain surprising levels of reproductive isolation. To better understand how species boundaries are maintained, the current study uses statistical ordination analyses to determine whether species isolation across shared geographical ranges might be achieved through subtle differences in floral characters among species. The possible contribution of differences in flowering phenology (e.g. temporal reproductive isolation) to reproductive isolation was also evaluated for the small subset of populations in which two or more species occur in direct sympatry. Among species in the P. picta complex, there are both phylogenetic and geographical trends in some floral characteristics, whereas other characters do not covary with either geography or species identity. In several sympatric populations, differences in flowering phenology among species suggest that timing plays a major role in non‐random (i.e. mainly conspecific) mating. The conclusions of this study are that reproductive isolation in the P. picta species complex is reinforced by differences in the timing of floral maturation and the morphologies of androecium and floral display characters. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00 , 000–000.