Reproductive characters in a gynodioecious species, Silene italica (Caryophyllaceae), with attention to the gynomonoecious phenotype

Gynodioecious populations (i.e. populations with female and hermaphrodite individuals) often contain a third phenotype with an intermediate sex expression. In Silene italica , this phenotype is characterized by a mixture of pistillate and perfect flowers and is thus gynomonoecious. To characterize sexual functions of these gynomonoecious individuals and their potential influence in the maintenance of gynodioecy, reproductive characters of the three sexual phenotypes were compared over 2 years in several families of S. italica produced by crossing. We found that gynomonoecious individuals were intermediate for flower production and female fertility characters, although they did not always significantly differ from female individuals. Perfect flowers of gynomonoecious and hermaphrodite plants were similar in size and pollen production. Gynomonoecious individuals were thus intermediate in female and in male functions. Family effects were found for most of the characters. The female advantage (i.e. the fertility of females compared to the female fertility of pollen producing plants) was not dramatically different when gynomonoecious plants were taken into account.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 583–591.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

Pollen fates and the limits on male reproductive success in an orchid population

Male reproductive success in higher plants depends largely on the fates of pollen, but current methodologies have given only partial insights into this important aspect of plant mating. We present a detailed analysis of the proportions and absolute amounts of stained pollen involved in six key fates for the hawkmoth-pollinated African orchid Disa cooperi . Despite being packaged into pollinaria, high proportions (> 0.95) of the pollen removed from anthers were lost during transport by hawkmoths in both years. The proportion of pollen lost correlated positively with the number of pollinaria removed from a plant, so that pollen export did not vary with pollen removal. Most pollen was dispersed to neighbouring plants, with rare long-distance dispersal up to 65 m. Of the pollen that reached stigmas during both years, roughly equal amounts were involved in facilitated self-pollination vs. cross-pollination, but the relative proportions of these fates differed between years. Contrary to expectation, we found that self-pollination between flowers did not increase with the number of open flowers, even though moths probed significantly more flowers on larger plants. However, during both years the fraction of removed pollen exported to other plants declined significantly with increasing self-pollination on the source plant, indicating that once self-pollination occurred it reduced (discounted) subsequent pollen export opportunities. The packaging of pollen into pollinaria in orchids appears to increase overall transfer efficiency by at least an order of magnitude relative to plants with granular pollen. Nevertheless, considerable uncertainties remain in the male reproductive success of individual orchids. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 175–190.     

Diversity, flower visitation frequency and generalism of pollinators in temperate rain forests of Chiloé Island, Chile

Species richness and taxonomic composition of pollinator assemblages are documented for 26 plant species from temperate rain forests of northern Chiloé Island, southern Chile (42°30'S). We investigated the patterns of generalism and specialization among plants and animal pollinators by comparing the flower visit frequency by different pollen vectors during the spring and summer months of three consecutive years (2000–2002). Species studied exhibited a range of floral morphologies (radial vs. zygomorphic, open vs. tubular) and rewards (nectar and/or pollen). Overall, we recorded 172 pollinator species, with an average of 6.6 species of pollen vectors/plant species. Pollinators visited an average of 15.2 plant species/pollen vector. Pollinator assemblages were dominated by Coleoptera (75 species), Diptera (56 species) and Hymenoptera (30 species), but passerine birds and hummingbirds were also important. The most specialized plants were vines, including the bee-pollinated genus Luzuriaga (Philesiaceae) and two endemic species of hummingbird-pollinated Gesneriaceae. Hymenoptera contributed 41.2% of all visits, with the bumblebee Bombus dalhbomii accounting for 22.5% of these. Plants with unspecialized flower morphology supported a higher species richness of pollinators, but visiting rates did not differ from specialized flowers.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 399–416.     

Behavioural responses of wheat stem sawflies to wheat volatiles

1 Adult wheat stem sawflies Cephus cinctus, pests of cultivated cereals that also infests wild grasses, migrate into wheat fields where they oviposit in elongating, succulent stems.
2 Volatiles released by wheat plants at susceptible stages were analyzed to determine potential semiochemical compounds. Seven major compounds were identified and quantified.
3 A Y-tube bioassay was developed to evaluate upwind orientation of adult sawflies in response to an airstream that passed over elongated wheat plants. The bioassay was also conducted with synthetic volatile compounds. The compounds were tested using a range of concentrations spanning those identified in the airstream passing over wheat plants.
4 A significant number of adult females were attracted to wheat plants when given a choice of either purified air or the air passing over plants.
5 A significant number of female C. cinctus were attracted to ( Z )-3-hexenyl acetate, β-ocimene, and ( Z )-3-hexen-1-ol, but were repelled by 6-methyl-5-hepten-2-one. Females did not respond to ( E )-2-hexenal, or ( E )-2-hexenyl acetate. The behavioural responses were concentration dependent; the highest tested concentration of ( Z )-3-hexenyl acetate was repellent to females of this species.
6 Adult males did not discriminate between air passing over wheat plants and air from a purified airstream. Males did not respond to any tested synthetic compound at any concentration.
7 The present study demonstrates for the first time that adult females of wheat stem sawfly display innate behaviours in response to synthetic volatiles. These results provide a basis for the potential development of resistant wheat varieties and for the development of semiochemically-based pest management.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

The first record of gynodioecy in a species of Gnidia (Thymelaeaceae) from South Africa

Sexual polymorphism was studied in the shrub Gnidia wikstroemiana (Thunb.) Meisn. from the semiarid Nama Karoo Biome, South Africa. The populations comprised plants bearing either female flowers, or hermaphrodite flowers with variable female function. In two populations, female plants accounted for 36–37% of the flowering plants. Female flowers were smaller and their stamens were reduced to staminodes, but their styles were significantly longer than those of hermaphrodite flowers. Energy investment in flowers and fruits for females and hermaphrodites was measured using bomb calorimetry. Females produce a greater number of less costly flowers than hermaphrodites, and invest less energy per unit in production of flowers and inflorescences. In contrast, females invest more energy per unit in production of fruits and infructescences than hermaphrodites. Females overall invest 7.3% more energy in reproduction than hermaphrodites. Female flowers were obligate out-crossers (xenogamous), with 35% of nonmanipulated, open-pollinated flowers setting fruit, comparable with fruit set among selfed hermaphrodite flowers. The breeding strategy of G. wikstroemiana most closely resembles gynodioecy. This is the first report of sexual dimorphism in Gnidia L. and sub-Saharan Thymelaeaceae.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 219–233.     

Volatile chemical release by bethylid wasps: identity, phylogeny, anatomy and behaviour

The structures of volatile chemicals released by parasitic wasps in the family Bethylidae are shown to correspond to the subfamily to which the species belong. Species in the Epyrinae release skatole (3-methylindole) and species in the Bethylinae release a spiroacetal (2-methyl-1,7-dioxaspiro [5.5]undecane): these compounds are chemically very different. The enantiomeric composition of the spiroacetal differs between congeneric species. Chemical release is a discrete event under the active control of both male and female wasps. Structural differences between the mandibular glands and intramandibular glands suggest the mandibular glands to be the source of released volatiles. Real-time mass spectrometry shows that the spiroacetal is released by Goniozus nephantidis females during dyadic resource contests, with release more common during more aggressive interactions. Chemical tagging with deuterium further shows that the volatile is released by the loser of an agonistic interaction and not the winner. The function of spiroacetal and skatole release by bethylids is discussed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 837–852.     

Factors affecting pollination ecology of Quercus anemophilous species in north-west Spain

Pollination ecology of Quercus is influenced by meteorological, biotic and genetic factors. This study was undertaken to ascertain the effect induced by these factors on pollen production, release and dispersion. Aerobiological data have been used in recent years as phenological information, because the presence of pollen in the air is the result of flowering across a wide area. The onset of the Quercus pollen season and the atmospheric pollen concentrations during the pollination period in two localities of north-west Spain (Ourense and Santiago) were determined from 1993 to 2001. There were important variations in total annual pollen as a result of meteorological conditions, lenticular galls produced by Neuropterus on catkins and biennial genetic rhythms of pollen production. In order to determine the beginning of flowering, a thermal time model has been used. Chill requirements were around 800 chilling hours (CH) and heat requirements were 953 growth degree days (GDD in °C) in Santiago and 586 GDD in Ourense. Pollen in the air show positive correlation (99% significance) with daily thermal oscillation, maximum and minimum temperatures, and hours of sunshine. Regression analysis with previous days' pollen concentrations explained the high percentage of pollen concentration variability, as meteorological variables do not, on their own, explain pollen production and release.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 283–297.     

Bauhinia larsenii, a fossil legume from Guangxi, China

Fossil leaves and a branch with a leaf and a pod attached have been discovered from Ningming County, Guangxi Autonomous Region, China, and are described as Bauhinia larsenii sp. nov. (Leguminosae: Cercideae). The strata from which the Bauhinia fossils were collected, namely the Ningming Formation, were dated as Late Eocene–Oligocene based on a combination of pollen, fauna and flora. The affinity of the new species to Bauhinia section Micralvesia subsection Viridescentes species is also discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 437–440.     

Reproductive biology of Boswellia serrata, the source of salai guggul, an important gum-resin

Detailed studies were carried out on the phenology, floral biology, pollination ecology and breeding system of Boswellia serrata Roxb. (Burseraceae) the source of 'salai guggul'. The trees remain leafless during the entire period of flowering and fruiting. The inflorescence is a terminal raceme and produces up to 90 bisexual, actinomorphic flowers. On average a flower produces 10 044 ± 1259 starch-filled pollen grains. About 85% of the fresh pollen grains are viable; the pollen to ovule ratio is 3348 : 1. The stigma is of the wet papillate type. The style is hollow with three flattened stylar canals filled with a secretion product. The stylar canals are bordered by a layer of glandular canal cells. The inner tangential wall of the canal cells shows cellulose thickenings. The ovary is trilocular and bears three ovules, one in each locule. Flowers offer nectar and pollen as rewards to floral visitors. The giant Asian honey bee ( Apis dorsata ) and A. cerana var. indica (Indian honey bee) are the effective pollinators. The species is self-incompatible and the selfed pollen tubes are inhibited soon after their entry into the stigma. Self-pollen tubes develop a characteristic 'isthmus' as a result of enlargement of the tube soon after emergence through the narrow germ pore. Cross-pollinated flowers allowed normal pollen germination and pollen tube growth, and resulted in fruit- and seed-set. Under open pollination fruit-set was only about 10%. Although manual cross-pollinations increased fruit set, it was only up to about 20%. Low fruit set appears to be the result of inadequate cross-pollination and other constraints, presumably limitation of available nutrients.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 73–82.     

Signal components, acoustic preference functions and sexual selection in a cricket

In many sexually reproducing organisms, females choose mates based on multiple male traits. This study examined how two temporal components of the male mating call – chirp rate and chirp duration – affect female mating preference in five populations of a widely distributed North American cricket, Allonemobius socius (Orthoptera, Gryllidae). Chirp rate and chirp duration of the A. socius mating call were varied independently, and the responses of virgin females to these experimentally manipulated calls were repeatedly measured using a sequential single-stimulus design. Significant among- and within-population variation in chirp-duration preferences of females were found. Contrary to many previous studies, call chirp rate had no effect on female phonotaxis. Also there was no evidence of an interaction between chirp rate and chirp duration on female response to male mating calls. Moreover, female responsiveness to average and above-average chirp duration appeared to decline with female (adult) age. Overall, these results suggest evolved differences among populations in chirp-duration preferences, and that selection can act within populations on female chirp-duration preference.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 461–472.     

Development and ultrastructure of the megagametophyte in Passiflora caerulea L. (Passifloraceae)

Megasporogenesis and megagametogenesis of Passiflora caerulea L. were studied using light and transmission electron microscopy. The archesporial tissue is generally formed by one cell. The megaspore mother cell gives rise to a linear tetrad of megaspores. The chalazal megaspore is the functional one, and originates a Polygonum -type female gametophyte. The antipodals are ephemeral. Abundant starch is found in the nucellar cells, specially the ones adjacent to the megagametophyte. The two synergids show ultrastructural differences, involving the filiform apparatus, the nucleolus and the endoplasmic reticulum; these differences suggest a functional differentiation, probably related to the reception of the pollen tube. This is the first report in angiosperms of substantial morphological differences between the two synergids.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 73–81.     

Phyllanthus section Choretropsis (Euphorbiaceae) in Brazil

Section Choretropsis genus Phyllanthus (Euphorbiaceae) is formed by plants whose branches are modified into phylloclades. Despite the suitable characteristics of phylloclades within the Phyllanthus genus, the systematics of these species are poorly understood. Morphological data are presented here to allow future revision of this taxonomic group. The section is represented by nine species distributed in South America, eight of them endemic to Brazil. Our taxonomic studies distinguish the species and propose the synonymization of nine varieties of P. klotzschianus and two varieties of P. flagelliformis based on vegetative and reproductive characters. The diversity of branching and the morphology of phylloclades are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 131–164.     

Different rewards in female and male flowers can explain the evolution of sexual dimorphism in plants

Insects use floral signals to find rewards in flowers, transferring pollen in the process. In unisexual plants, the general view is that staminate (male) and pistillate (female) flowers obtain conspecific pollen transfers by advertising their rewards with similar floral signals. For female plants lacking food rewards, this can lead to floral mimicry and pollination by deceit. In this study, we challenge this view by presenting evidence for different rewards offered by flowers on females and males, as a mechanism promoting sexual dimorphism in Leucadendron xanthoconus (Proteaceae), a clearly sexually dimorphic shrub. The tiny beetle pollinators Pria cinerascens (Nitidulidae) depend entirely on the plants they pollinate for survival and reproduction. Male flowers provide mating and egglaying sites, and food for adults and larvae. Female flowers lack nectar and function to shelter pollinators from rain. Their flower heads have cup‐shaped display leaves, and are more closed than are those in males. On rainy days, flowers on females received 30% more visits than did flowers on males, and 90% more than they did on sunny days. When we removed display leaves in females, intact flower heads received 14 times more P. cinerascens visits than did manipulated flower heads, indicating that the cup shape attracts the beetles. In both sexes, having many flowers increased the probability of visits and the number of P. cinerascens visiting a plant. In males, the number of larvae was positively correlated with floral‐display size, while in females, seed set (pollen transfers) showed no relationship with floral‐display size. Ninety‐five per cent of the ovules received pollen and 52% matured into seeds. We explain the sexual dimorphism in L. xanthoconus as a result of an intimate partnership with P. cinerascens pollinators, in conjunction with a rainy climate. Pollinators favour large male floral displays, because they offer a reliable food source for adults and larvae. Frequent rains drive the P. cinerascens to leave males in search of the protection offered by females. Because females offer shelter, an essential resource that is not offered by male plants, they receive sufficient pollen independent of their floral‐display size. This pollination system promotes the evolution of sexually dimorphic floral signals, guiding pollinators to different rewards in male and female flowers. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 97–109.     

Hesperis turkmendaghensis (sect. Hesperis) (Cruciferae/Brassicaceae), a new species from the Central Anatolia region, Turkey

Hesperis turkmendaghensis A.Duran & A.Ocak sp. nov. (Cruciferae) is described and illustrated from Anatolia, Turkey. The species grows under mixed forest, open forest and shady slopes in Türkmen Dağı (B3 Eskişehir) in Central Anatolia. It is closely related to H. matronalis L. ssp. matronalis , an endemic confined to Central Anatolia. Diagnostic morphological characters are discussed. Notes are also presented on its ecology, biogeography and conservation status. In addition, the pollen characteristics and seed coat surface of H. turkmendaghensis and H. matronalis are examined by SEM.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 239–247.     

Reproductive trait divergence and hybrid fertility patterns between chromosomal races of the house mouse in Tunisia: analysis of wild and laboratory-bred males and females

The divergence in reproductive features and hybrid fertility patterns between two chromosomal races (2 n  = 40, 40St, and 2 n  = 22, 22Rb) of the house mouse in Tunisia were re-assessed on a larger sample of wild and laboratory-bred individuals than studied hitherto. Results showed that litter sizes were significantly smaller in 40St than in 22Rb mice, contrary to previous analyses. This suggests that variation in litter size between the two chromosomal races is more likely related to selective and/or environmental factors acting locally than to interracial reproductive trait divergence. However, the significantly reduced litter size of F₁ hybrids compared with parental individuals was confirmed, and further highlighted a sex difference in hybrid infertility, as F₁ females produced fewer litters and of smaller size than males. Histological analyses of F₁ and backcrosses showed a breakdown of spermatogenesis in males and a significantly reduced primordial follicle pool in females. The degree of gametogenic dysfunction was not related to the level of chromosomal heterozygosity per se , but a significant effect of two Rb fusions on follicle number was observed in hybrid females. These results suggest that genetic incompatibilities contribute to primary gametogenic dysfunction in hybrids between the chromosomal races in Tunisia.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 407–416.     

The cytological origin of the polyads and their significance in the reproductive biology of Mimosa bimucronata

The mechanism of polyad formation in Mimosa bimucronata (DC.) Kuntze is described and its significance in the reproductive biology is discussed. In this species c . 500 complexes of two coupled pollen mother cells (PMCs) per pollen sac are formed after the last mitotic division of the sporogenous tissue. All the pollen grains of each complex remain attached and form a bitetrad. This mechanism leads to the formation of many polyads per pollen sac. On this basis, we propose that in Mimosa the number of pollen grains per polyad (eight or 16) is determined by the number of cells that remain attached after the last mitotic divisions of the sporogenous tissue. Concerning the biological significance of the polyads in M. bimucronata , we find that the number of pollen grains per polyad is correlated with the number of ovules per ovary, and that the internal dimensions of the pit-shaped stigma are sufficient just for one polyad to be localized in it. Based on the number of polyads per flower this species has a high pollen/ovule ratio characteristic of xenogamous plants, in opposition to that expected for plants with composite pollen dispersal units. In contrast, it has low pollination efficiency, low fruit set, a 1/1 relationship between the pollen grains of a polyad and the number of ovules per ovary, and high seed set per fruit. These facts support the hypothesis that the occurrence of polyads in M. bimucronata would be an adaptation helping to ensure high seed set following a single cross pollination event.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 343–349.