Patterns of flowering and sex-ratio variation in the Mediterranean shrub Phillyrea angustifolia (Oleaceae): implications for the maintenance of males with hermaphrodites

The Mediterranean shrub Phillyrea angustifolia is one of the few plants that display androdioecy, a rare breeding system in which males co-occur with hermaphrodites. However, it has been difficult to account for male frequencies previously documented for this species, which have frequently been higher than 0.5. Here we present sex-ratio data from 13 sites in Spain and Portugal that throw further light on the maintenance of androdioecy in P. angustifolia . The inferred proportion of males in these populations was around 0.3, substantially lower that 0.5 and concurring with theoretical predictions for androdioecy. Differences in the proportion of hermaphrodites in flower between "stressed" and "nonstressed" sites suggest the existence of an important life-history trade-off between fruit production and the rate at which plants recover from recurrent disturbances such as fire. The reduced frequency of flowering by hermaphrodites relative to males may help to explain the maintenance of androdioecy in this long-lived, woody species.     

Breeding system and spatial variation in the pollination biology of the heterocarpic Thymelaea velutina (Thymelaeaceae)

Thymelaea velutina (Thymelaeaceae) is a unique dioecious and heterocarpic shrub, each female producing both dry and fleshy fruits. It is endemic to the Balearic Islands (western Mediterranean) and has suffered an important regression in recent decades, especially due to habitat loss. It lives in coastal areas, mainly in dunes, but a few populations occur up to 1300 m in the mountains of Mallorca. In the present study, we examined its breeding system and pollen vectors, determined whether seed production is pollen-limited, and quantified the relative importance of different insect visitors at coastal and mountain sites. Selective exclusions showed no differences between populations in the relative importance of insects and wind as pollen vectors. Pollen was limiting at both populations, though not consistently in time. Flowers of T. velutina were visited mostly by generalist insects, and species composition and abundance of flower visitors varied both in space and time. The flowering period is about a month later in the mountain than in the dune, and this results in a higher frequency of insect visits to plants in the mountain due to the higher insect abundance; insects were also more diverse in the mountain, although they visited a proportionally lower number of flowers than in the dune. Despite this spatio-temporal variation in the frequency of insect visits and in the importance of a particular insect group, the ambophilous system in this species (previously described as anemophilous) ensures a relatively high fruit set at the two different sites.     

Reproductive biology of Melanophryniscus montevidensis (Anura: Bufonidae) from Uruguay: reproductive effort,fecundity, sex ratio and sexual size dimorphism

We analyzed the reproductive traits of a population of Melanophryniscus montevidensis. The reproductive events were associated to rainfalls. Males called from dawn to late night, were more abundant than females, and adopted alternative tactics to find partners (active search). Females were larger than males, and had mature oocytes and lipid reserves after spawning; they showed no association between size and fecundity. As an explosive breeder, females are prepared all year long for reproduction. Several individuals revealed high site-fidelity for breeding. Knowledge about breeding pattern could help to propose suitable and efficient in situ conservation measures for this endangered species.     

Reproductive biology and pollination ecology of Triplaris gardneriana (Polygonaceae): a case of ambophily in the Brazilian Chaco

• Triplaris gardneriana (Polygonaceae) is a dioecious pioneer tree reported as insect‐pollinated, despite possessing traits related to anemophily. Here, we analyse the possible roles of insects and wind on the pollination of this species to establish whether the species is ambophilous.
• We carried out observations of floral biology, as well as on the frequency and behaviour of pollinators visiting flowers in a population of T. gardneriana in the Chaco vegetation of Brazil. We conducted experimental pollinations to determine the maternal fertility of female plants and whether they were pollen‐limited, and we also conducted aerobiological experiments to provide evidence of how environmental factors influence atmospheric pollen dispersal.
• The population comprised an area of approximately 152.000 m² and was composed of 603 female and 426 male plants (sex ratio = 0.59:0.41). We observed 48 species of insects visiting flowers of T. gardneriana, of which the bees Scaptotrigona depilis and Apis mellifera scutellata were the most effective pollinators. We recorded pollen grains dispersed by wind on 74% of the glass slides placed on females, located at different distances (1–10 m) from male plants.
• Airborne pollen concentration was negatively correlated with relative humidity and positively correlated with temperature. Our observations and experimental results provide the first evidence that T. gardneriana is an ambophilous species, with pollen dispersal resulting from both animal and wind pollination. This mixed pollination strategy may be adaptive in T. gardneriana providing reproductive assurance during colonisation of sites with different biotic and abiotic conditions.

     

Multiple mating and its effect on the reproductive success of female Epiphyas postvittana (Lepidoptera: Tortricidae)

Abstract.

• 1 Multiple mating and its effect on reproductive performance of female Epiphyas postvittana (Walker) moths were studied under controlled conditions.
• 2 The age at which the moths mated for the first time ranged from the first to the tenth day after emergence, but 71% of first matings were during the first 3 days.
• 3 The majority (63%) of females had one or two spermatophores in the bursa copulatrix. Some (24%) were found with three to five spermatophores, whereas no successful mating occurred among 13% of individuals. The number of matings was partly dependent on the number of mates available to the female. Between the range of sex ratios of one male to one female and four males to one female maximal mating success occurred at the ratio of three males to one female.
• 4 Virgin females were capable of egg-laying, but mating stimulated and accelerated oviposition. Mated individuals laid twice as many eggs as unmated ones.
• 5 The level of copulatory activity did not influence the longevity of females irrespective of the number of males available to them.
• 6 Sex ratios with greater than one male to a female improved the reproductive success by marginally increasing fecundity and fertility.
• 7 It is concluded that multiple mating would enhance population growth, and is of particular benefit to populations with a preponderance of females, as is known to occur naturally in this species.

     

Pollination biology and reproductive success in four Brazilian species of Gomesa (Orchidaceae: Oncidiinae): Specific pollinators,but high pollen loss and low fruit set

Four species of Gomesa (Gomesa flexuosa, Gomesa ranifera, Gomesa cornigera, and Gomesa riograndensis), a horticulturally important, albeit scarcely studied orchid genus, were investigated. Pollination biology was studied through fieldwork in Rio Grande do Sul in Southern Brazil during two flowering seasons. Breeding systems were studied through controlled pollinations in cultivated plants. All studied species secrete floral oils through elaiophores and are exclusively pollinated by native, oil-collecting bees of the Centridini or Tapinotaspidini tribes. Pollinarium reconfiguration after withdrawal, a feature likely promoting cross-pollination, was recorded in G. cornigera and G. riograndensis. All studied species are pollinator-dependent and either predominantly or fully self-incompatible, that is, unable to set fruit following self-pollination. The used indexes of pollination efficiency consistently scored values below 1, indicating that in all studied species less than one flower is pollinated per pollinarium removed. Accordingly, pollen loss was high, ranging from 40% to 75%. Furthermore, inflorescences in natural populations displayed low fruit sets (below 13%). Gomesa flexuosa presented the highest values of pollinarium removal (36.24%) and deposition (15.34%). Significantly, this is the only studied species bearing a tabula infraestigmatica, a column swelling that bees hold to stabilize themselves while gathering the floral oils. Since pollinators were observed in all studied species spending considerable time (up to 3 min 46 s) and visiting several (up to 19) flowers, we propose that the observed low fruit set is the result of a combination of self-incompatibility, pollinator-mediated self-pollination, and pollen loss.     

Population structure and reproductive period of whitebelly prawn Nematopalaemon schmitti (Holthuis 1950) (Decapoda: Caridea: Palaemonidae) on the southeastern coast of Brazil

The population biology of Nematopalaemon schmitti in the Ubatuba region on the northern coast of the state of São Paulo was characterized through analyses of the length-frequency distribution, sex ratio, reproductive period, and juvenile recruitment. The specimens were caught monthly from January 1998 to December 1999, from a shrimp boat equipped with double-rig trawl nets. A total of 1073 specimens were analyzed, and the sex and carapace length (0.1?mm) of each individual were noted. The analyzed specimens consisted of 152 juveniles, 437 adult males, 296 adult females, and 188 ovigerous females (OFs). The monthly distribution of N. schmitti by size classes revealed a stable population structure, with both juveniles and adults continuously present. This population showed a unimodal distribution; however, bimodality was identified in June 1998 and 1999 for males, and June 1998 and July 1998 and 1999 for females, probably related to recruitment pulses of juveniles. Sexual dimorphism was indicated by the presence of females in the larger size classes, probably because of their growth rate. The proportion between males and females differed from 1?:?1 in certain size classes and seasons of the year (χ ²,?p?<?0.05); in some situations, the females were more abundant than the males, or vice versa. No significant relationship was detected between the seasonal mean temperatures of the bottom water and the percentages of OFs and young, or for the relationship between these two groups (Spearman, p?>?0.05). The continuous presence of OFs and young in the population suggests a pattern of continuous reproduction for N. schmitti in the Ubatuba region.     

Flower and fruit structure of the endangered species Petagnaea gussonei (Sprengel) Rauschert (Saniculoideae,Apiaceae) and implications for its reproductive biology

Petagnaea gussonei (Apiaceae) is an endangered species endemic to the Nebrodi mountains (north-eastern Sicily). Although an increasing number of studies have been performed on this species, its reproductive biology remains poorly understood. The aim of this study was to investigate in detail the structure of the flower and the fruit of Petagnaea, and the possible implications for its breeding system and seed dispersal mechanism. Results from fieldwork, light microscopy and scanning electron microscopy suggest (1) the presence of protandrous hermaphrodite flowers; (2) geitonogamy, autogamy, and allogamy as breeding system mechanisms of P. gussonei, even if asexual reproduction is preferred by the plant; and (3) epizoochory and hydrochory as possible modalities of fruit dispersal.     

Reproductive ecology of the endangered Alpine species Eryngium alpinum L. (Apiaceae): phenology, gene dispersal and reproductive success

BACKGROUND AND AIMS: Eryngium alpinum (Apiaceae) is an endangered perennial, characteristic of the Alpine flora. Because the breeding system influences both demographic (reproductive success) and genetic (inbreeding depression, evolutionary potential) parameters that are crucial for population maintenance, the reproductive ecology of E. alpinum was investigated. Specifically, the aims of the study were (1) to determine the factors (resources and/or pollen) limiting plant fitness; and (2) to assess the potential for gene flow within a plant, within a patch of plants, and across a whole valley where the species is abundant. METHODS: Field experiments were performed at two sites in the Fournel valley, France, over three consecutive years. Studies included a phenological survey, observations of pollinators (visitation rates and flight distances), dispersal of a fluorescent powder used as a pollen analogue, the use of seed traps, determination of the pollen/ovule ratio, and an experiment to test whether seed production is limited by pollen and/or by resources. KEY RESULTS: E. alpinum is pollinated by generalist pollinators, visitation rates are very high and seed set is resource- rather than pollen-limited. The short flights of honeybees indicate a high potential for geitonogamy, and low pollen and seed dispersals suggest strong genetic structure over short distances. These results are interpreted in the light of previous molecular markers studies, which, in contrast, showed complete outcrossing and high genetic homogeneity. CONCLUSION:S. The study highlights the usefulness of adopting several complementary approaches to understanding the dynamic processes at work in natural populations, and the conservation implications for E. alpinum are emphasized. Although the studied populations do not seem threatened in the near future, long-term monitoring appears necessary to assess the impact of habitat fragmentation. Moreover, this study provides useful baseline data for future investigations in smaller and more isolated populations.     

Marking of host fruit by maleRhagoletis boycei Cresson flies (Diptera: Tephritidae) and its effect on egg-laying

MaleRhagoletis boycei flies (Cresson) have been shown previously to guard egg-laying punctures on host walnut (Juglans major) fruit. During their tenure,R. boycei males were observed to dip the posterior portion of their abdomen repeatedly toward the fruit surface. Closeup video analysis revealed that, during this dipping behavior, a clear viscous substance emanating from the cercus was deposited directly on the fruit. Laboratory assays indicated that the rate of abdomen-dipping was significantly higher on fruit bearing simulated egg-laying punctures than on control fruit and significantly higher in the area of the puncture than elsewhere on the fruit. In a final experiment, the occurrence of abdomen-dipping in the vicinity of a puncture was manipulated independently on separate punctures on the same fruit. On such fruit and in the absence of males, previously mated, reproductively mature females attempted oviposition significantly more often in or near a puncture exposed to males than in or near an unexposed control puncture. Possible functions of the putative male-marking behavior from the perspectives of both male and female are discussed.     

The effect of different exogenous kisspeptins on sex hormones and reproductive indices of the goldfish (Carassius auratus) broodstock

Despite several studies on fish hormone therapy, finding new candidates may provide more reproductive efficiency in artificial propagation. Kisspeptins, being upstream of the hypothalamic–pituitary-gonadal axis, appear to play a key role in the reproduction process. In the present study, the effect of different variants of kisspeptide, including goldfish (Carassius auratus) kiss1 kisspeptin (Kiss1), human kisspeptin (Hkiss), and their combination (Kiss1 + H), on the reproductive indices of goldfish broodstock in comparison to Ovaprim (a typical synthetic Gnrh hormone) was investigated. Peptides (Kiss1 and Hkiss) were synthesized using a solid-phase synthesis approach. Kiss1 and Hkiss were injected at a dose of 100 μg kg⁻¹ body weight, blood samples were taken 6 h after injection and sex hormones (E2, Dhp, and 11-Kt), gonadotropins (Lh and Fsh), cortisol and reproductive indices (fecundity, fertilization and hatching percentage) were measured. The results showed a significant increase of plasma sex hormones and gonadotropins in fish treated with kisspeptins. In addition, the cortisol and lipoprotein lipase in Kiss1, Hkiss and Kiss1 + H were remarkably increased compared to Ovaprim. In conclusion, kisspeptins could be a more suitable candidate than Ovaprim for accelerating and synchronizing oocyte maturation in the fisheries industry.     

The effect of ejaculate mass on female reproductive output in the european swallowtail butterfly,Papilio machaon (L.) (Lepidoptera: Papilionidae)

Previously, we showed that virgin males of Papilio machaondeliver ejaculates that are twice as big as any ejaculates they transfer at later matings. Here, we investigate the consequences of these two size classes of ejaculates on female reproductive output and demonstrate that females that received one small ejaculate laid as many eggs, fertilized the same proportion of eggs, and lived as long as those that had received one big ejaculate. However, females that received big ejaculates laid heavier eggs, but only between the twelfth and the fifteenth days of egg-laying. We conclude that male-derived nutrients appear to have a limited effect on female reproductive output in P. machaonand that the large size of ejaculates delivered by male butterflies are determined primarily by selection on males to produce longer refractory periods in females.     

Evidence of gynodioecy and sex ratio variation in Wurmbea biglandulosa (Colchicaceae)

 A valuable approach to understanding the evolution of gender dimorphism involves studies of single species that exhibit intraspecific variation in sexual systems. Here we survey sex ratios in 35 populations of Wurmbea biglandulosa, previously described as hermaphroditic. We found pronounced intraspecific variation in sexual systems; populations in the northeastern part of the species' range were hermaphroditic, whereas other populations were gynodioecious and contained 2–44% females. Populations with lower annual rainfall were more likely to be gynodioecious, supporting the view that gender dimorphism evolves more frequently in harsher environments. In gynodioecious populations, however, female frequency was not related to either annual rainfall or habitat, indicating that other factors are important in determining sex ratio variation. Females had smaller flowers and shorter stems than did hermaphrodites, potentially providing a basis for resource compensation. A female fecundity advantage may contribute to the maintenance of females in populations because females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites. Received March 2, 2001 Accepted February 25, 2002     

Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits

The density of great tit Parus major L. and blue tit Parus caeruleus L. was artificially increased by placing nest-box colonies for these species in the vicinity of the nests of breeding tawny owls during 1993–1997. Bird prey composition in the owl nests, the proportion of parents disappearing from the breeding tit populations and the reproductive performance of the widowed parents were analysed. The frequency of predation on tits by tawny owls was greater in areas where tit density had been artificially increased. Owls preyed more on tits during the feeding period of owlets than during the incubation period and more in years when snow covered the ground during the incubation period than when it did not. Mortality due to predation was male biased and more females lost their mates in populations breeding near tawny owl nests. Reproductive performance of the widowed parents was lower and their body weights were lighter at the end of the nestling period than those found in birds rearing youngs with their mates. Predation by owls increased the between-year turnover in the breeding tit population: widowed parents did not return to the nesting site for the next breeding season.     

Floral biology,breeding system and pollination ecology of Justicia betonica L. (Acanthaceae): An assessment of its low reproductive success in West Bengal,India

Besides its importance as an ornamental plant, Justicia betonica L. is also used as a medicinal plant for the treatment of several human disorders. However, the population size and abundance of the plant species are very low in Indian states, including West Bengal. The breeding system and pollination ecology of J. betonica are still unclear. Therefore, some reproductive aspects were investigated in order to provide important information for the sustainability of the species. The flowering duration spanned from December to May, with its peak in February–March. Flower opening time was much earlier (5.00–8.00 a.m.) on a hot day (in April–May) than on a cold day (10.00 a.m.–12.00 p.m. in December–January). The time of anther dehiscence coincided with the flower opening time; however, the stigma became receptive later. The plant species is self-compatible but dependent on pollinators for fruit and seed sets. Members of different insect groups like ants, bees, butterflies, flies, moths and wasps visit the flowers. Among them, Allorhynchium metallicum and Halictus acrocephalus are the primary pollinators (based on the “relative pollinator service”). The plant species showed a significant pollination deficit (coefficient of pollination deficit [D] = 0.32) in the open condition. Larvae of the florivorous moth Gatesclarkeana erotias significantly reduced reproductive success by eating their floral parts, ovules and immature seeds. In conclusion, combining the pollination deficit and the feeding activity of phytophagous insects (moth larvae) may reduce the reproductive fitness of J. betonica in the study regions.     

Breeding systems and phylogeny in Poa,with special attention to Northeast Asia: The problem of Poa shumushuensis and sect. Nivicolae (Poaceae)

Poa sect. Poa subsect. Nivicolae (Prob.) Tzvelev was circumscribed to include four species of the Soviet Union: Poa shumushuensis, Poa caucasica, Poa irkutica, and Poa veresczaginii. Bayesian phylogenetic analyses of plastid and nuclear‐ribosomal DNA revealed that it is polyphyletic, none of these species are closely related. Poa shumushuensis, type of sect. Nivicolae, or its ancestor, likely displayed the nrDNA genotype characteristic of the higher polyploid P. sect. Malacanthae. Genotype codes are designated for each species: Hx, C c, Php, and Shp. Poa sect. Nivicolae s.s. is restricted to P. shumushuensis; P. sect. Irkuticae is restricted to P. irkutica; P. caucasica is moved to P. subg. & sect. Caucasicae nov.; and P. sect. Dschungaricae is resurrected for P. veresczaginii and two other species. Although diclinous breeding systems are known in many western hemisphere species of Poa, dicliny is infrequent and little studied in Asian Poa. Poa shumushuensis is judged to be either sequentially gynomonoecious or gynodioecious. A ratio of 2 perfect‐, to 2 mixed‐, to 1 pistillate‐flowered inflorescences from different plants in P. shumushuensis is suggestive of a recessive allele for stamen suppression, and this is associated with subtle sexual‐dimorphism. Poa irkutica is diclinous with a breeding system between simple gynomonoecy and sequential gynomonoecy; P. caucasica is perfectly flowered; and P. veresczaginii has infrequent abortive anthers, indicative of limited dicliny, or sterility for other reasons possibly related to its reticulate origin. In total, 23 Asian species are here reported to be diclinous and their breeding systems are characterized. A lectotype is designated for P. fauriei.     

Comparative biology,predation capacity and effect of an artificial diet on reproductive parameters of green lacewing Mallada boninensis (Neuroptera: Chrysopidae)

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