Impact of prairie fragment size on proportion of females and reproductive success of Lobelia spicata Lam., a gynodioecious species

• Due to ongoing human impacts, plant species increasingly occur in landscapes that are highly fragmented, with remaining natural habitats occupying small areas, resulting in populations that are smaller and more isolated than in previous time periods. This changed metapopulation structure is expected to have negative impacts on seed production. For example, the proportion of female plants within gynodioecious populations may be more volatile due to genetic drift in small populations associated with small habitat fragments, with concomitant impacts on seed production. My aims were to determine: (i) if variation in proportion of females is larger in smaller fragments; and (ii) if such changes in female frequency in small fragments result in reduced seed production.
• Thirty‐two populations of Lobelia spicata Lam., a gynodioecious species, were surveyed in 2000, 2001 and 2009 in the tallgrass prairie region of Midwestern North America (Illinois and Indiana, USA). Data were collected for: proportion of female plants, total number of flowering plants (measure of population size), seed set per plant and prairie fragment size (another measure of population size).
• The proportion of females is more variable in smaller prairie fragments. Seed number per fruit decreases as the proportion of females increases in a population, but only significantly for female plants. The number of flowering plants is positively associated with fruit production for both genders. Populations within larger prairie fragments have higher seed production.
• The reproductive consequences of habitat fragmentation depend on the plant breeding system. While both sexes were negatively impacted, females were more adversely affected.

     

Inbreeding effects on progeny sex ratio and gender variation in the gynodioecious Silene vulgaris (Caryophyllaceae)

In gynodioecious species, sex expression is generally determined through cytoplasmic male sterility genes interacting with nuclear restorers of the male function. With dominant restorers, there may be an excess of females in the progeny of self-fertilized compared with cross-fertilized hermaphrodites. Moreover, the effect of inbreeding on late stages of the life cycle remains poorly explored. Here, we used hermaphrodites of the gynodioecious Silene vulgaris originating from three populations located in different valleys in the Alps to investigate the effects of two generations of self- and cross-fertilization on sex ratio and gender variation. We detected an increase in females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for female and male fertility. Male fertility correlated positively with sex ratio differences between outbred and inbred progeny, suggesting that dominant restorers are likely to influence male fertility qualitatively and quantitatively in S. vulgaris. We argue that the excess of females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for gamete production may contribute to the maintenance of females in gynodioecious populations of S. vulgaris because purging of the genetic load is less likely to occur.     

Four species from western Mexico new to Lobelia (Campanulaceae: Lobelioideae)

Lobelia knoblochii, L. mcvaughii, andL. villaregalis are described and referred, withHeterotoma macrocentron Benth., toLobelia “grex Eriniformes” Wimmer. All are narrow endemics occurring on volcanic soils in the Sierra Madre Occidental of Mexico.Lobelia macrocentron, L. mcvaughii, andL. villaregalis are known only from the type localities.     

A cost of restoration of male fertility in a gynodioecious species,Lobelia siphilitica

Abstract.— Models allowing the coexistence of females and hermaphrodites in gynodioecious populations assume a simple genetic system of sex determination, a seed fitness advantage of females (compensation), and a negative pleiotropic effect of nuclear sex-determining genes on fitness (cost of restoration). In Lobelia siphilitica , sex is determined by both mitochondrial genes causing cytoplasmic male sterility (CMS) and nuclear genes that restore fertility when present with specific CMS haplotypes (nuclear restorers). I tested for a cost of restoration in L. siphilitica by measuring restored hermaphrodites for five fitness components and estimating the number of nuclear restorers by crosses with females carrying CMS1 and CMS2. A cost of restoration appears as a significant negative coefficient (B) in the regression model explaining fitness. I found that hermaphrodites carrying more nuclear restorer genes for CMS2 (or restorer genes of greater effect) have lower pollen viability (B =– 1.08, P = 0.001). This pollen viability cost of restoration in L. siphilitica supports the theoretical prediction that negative pleiotropic effects of restorers will exist in populations of gynodioecious species containing females. The existence of such a cost supports the view that gynodioecy can be a stable breeding system in nature.     

A Revision of Lobelia Subgen. Tupa (Campanulaceae) in China

Based on specimens as many as available, a quantitative analysis of character variability and correlation in Lobelia Subgen. Tupa was carried out with the results shown by pictorialized scatter diagram and histogram. Pollen grains and seed coat were examined under SEM. The classitication of the subgenus in China is revised. Twelve specipes are recognized; a new species, L. foliiformis, and a new subspecies, L. colorata subsp. guizhouensis, are described; L. taliensis is reduced to a subspecies or L. colorata, which consists of three subspecies; L. kwangsiensis is treated as a variety of L. davidii in accordance with the treat-ment made by Lian Yong-shan in Fl. Reip. Pop. Sin.73 (2)     

Two new taxa of Lobelia (Campanulaceae) from the Chihuahuan desert region of Mexico

Two taxa of Lobelia from north–central Mexico are described as new, both being in sect. Lobelia subsect. Leiospermae E. Wimmer, viz. L. henricksonii (in "grex" Rosulatae E. Wimmer) and L. anatina E. Wimmer var. riskindii (in "grex" Sub–spicatae E. Wimmer). These two taxa are confined to the Mexican state of Coahuila in areally restricted mesic habitats above 1400 m alt.     

Mating system, sex ratio, and persistence of females in the gynodioecious shrub Daphne laureola L. (Thymelaeaceae)

Although in gynodioecious populations male steriles require a fecundity advantage to compensate for their gametic disadvantage, southern Spanish populations of the long-lived shrub Daphne laureola do not show any fecundity advantage over hermaphrodites in terms of seed production and early seedling establishment. By using allozyme markers, we assess the mating system of this species in five populations differing in sex ratio, and infer levels of inbreeding depression over the whole life cycle by comparing the inbreeding coefficients at the seed and adult plant stages. Extremely low outcrossing rates (0.001相似文献   

Delayed pollination,stigma length,sex expression,and progeny sex ratio in spinach,Spinacea oleracea (Chenopodiaceae)

Delayed outcrossed pollination of female spinach plants (Spinacea oleracea L.) resulted in increased stigma length and a male-biased progeny sex ratio. One group of females was outcrossed 10–14 d after anthesis, a second group was never outcrossed, and a third group, the control, consisted of females that were outcrossed as soon as stigmas appeared. Stigma length was significantly greater for plants in the delayed and never outcrossed groups compared to the control. Furthermore, stigmas of virgin flowers grew until they were either pollinated or the plants produced anthers. Plants that were never outcrossed produced their own anthers and self pollinated. The resulting progeny were all female. The sons of the delayed outcrossed group produced more stamens, on average, than sons of the control group. The observed male-biased sex ratio among the progeny of delayed outcrossed plants could be due to gametic selection. To test for this, plants were held virgin until their stigmas reached a length of at least 3 mm. These stigmas were then pollinated either distally or proximally. No significant difference was found between the progeny sex ratios of these two treatments. However, both sex ratios were more male biased than progeny of plants pollinated the day of anthesis (control group of the first experiment). We conclude that maternal factors, rather than gametophytic selection, may be responsible for the male-biased sex ratio observed in the first experiment.     

Lobelia vivaldii (Campanulaceae: Lobelioideae), a remarkable new species of sect. Tylomium from Isla de Mona,Puerto rico

A new species ofLobelia (Campanulaceae: Lobelioideae) is described from Isla de Mona, a Caribbean island not previously known to harbor the genus. The small woody rosette-plants were collected in 1988 and 1992 at the northeastern corner of the island, where they grew from crevices in the vertical limestone cliffs, approximately 60 m above the sea. The new species,Lobelia vivaldii, belongs to sect.Tylomium, a group of approximately 36 suffruticose and woody species endemic to the Caribbean basin. Within that section, it is most similar to and apparently derived fromL. cirsiifolia, a less woody rain forest species found at higher elevations on several islands in the Lesser Antilles.     

Female compensation through the quantity and quality of progeny in a gynodioecious plant, Geranium maculatum (Geraniaceae)

One of the major evolutionary trends in flowering plants is the evolution of unisexual flowers (male or female) from perfect flowers. This transition has occurred repeatedly in many taxa and has generated a wonderful array of variation in sexual expression among species. Theoretical studies have proposed a number of mechanisms to explain how this level of variation could be maintained in natural systems. One possible mechanism is the female compensation hypothesis, which predicts that female mutants require an increase in their seed fitness in order to invade a hermaphroditic system. Using Geranium maculatum, I tested this hypothesis and showed that female mothers produced more and larger seeds than hermaphroditic mothers even though they were indistinguishable in their vegetative traits and the flower production. Seeds from females were also more likely to germinate and produced seedlings with larger above- and belowground biomass. These seedlings were more likely to flower than those from hermaphrodites in at least one of the two populations studied. Combined, these results indicated that females in G. maculatum did compensate for their loss of male function by producing more and better seeds than hermaphrodites. This provides a mechanism for the maintenance of female plants in this species.     

Variation in seed traits of Lobelia inflata (Campanulaceae): sources and fitness consequences

Seed germination constitutes an important event in the life cycle of plants. Two related seed traits affect fitness: seed size and the timing of seed germination. In three sets of experiments, we (1) partition the sources of seed-size variance in Lobelia inflata into components attributable to fruit size, relative fruit position, and parental identity; (2) examine the influence of pregermination conditions and seed size on time to germination; and (3) assess the fitness consequences of seed size and germination timing under seminatural, harsh conditions. Seed-size variance is attributable to both parental identity and fruit position within an individual. Distal fruits produce larger but fewer seeds. No significant correlation exists between fruit size and seed size, but a trade-off is found between the number and size of seeds contained in a fruit after correcting for fruit size. The timing of germination is influenced by seed size, light conditions before winter, and winter duration. Germination timing influences survival, and despite small seed size in this species (2 × 10 g/seed), seed size has a persistent and significant association with both final plant size and the probability of survival to autumn.     

Female resistance behaviour and progeny sex ratio in two Bradysia species (Diptera: Sciaridae) with paternal genome elimination

The relationship between female mating preferences and sex allocation has received considerable theoretical and empirical support. Typically, choosier females adjust their progeny sex ratio towards sons, who inherit the attractive traits of their father. However, in species with paternal genome elimination, where male sperm do not contain the paternal genome, predictions for the direction of progeny sex ratio biases and their relationship with female choosiness are atypical. Paternal genome elimination also creates a potential for male–female conflict over sex allocation, and any influence of female mate choice on sex ratio outcomes have interesting implications for sexually antagonistic coevolution. Within the Sciaridae (Diptera) are species that produce single‐sex progeny (monogenic species) and others in which progeny comprise both sexes (digenic species). Paternal genome elimination occurs in both species. We explore female mate resistance behaviour in a monogenic and digenic species of mushroom gnat from the genus Bradysia. Our experiments confirmed our theoretical predictions, revealing that in the monogenic and digenic species, females producing female‐biased progeny were more likely to have resisted at least one mating attempt.     

Microsatellite evidence for obligate autogamy,but abundant genetic variation in the herbaceous monocarp Lobelia inflata (Campanulaceae)

Although high levels of self‐fertilization (>85%) are not uncommon in nature, organisms reproducing entirely through selfing are extremely rare. Predominant selfers are expected to have low genetic diversity because genetic variation is distributed among rather than within lineages and is readily lost through genetic drift. We examined genetic diversity at 22 microsatellite loci in 105 individuals from a population of the semelparous herb Lobelia inflata L. and found (i) no evidence of heterozygosity through outcrossing, yet (ii) high rates of genetic polymorphism (2–4 alleles per locus). Furthermore, this genetic variation among lineages was associated with phenotypic traits (e.g. flower colour, size at first flower). Coupled with previous work characterizing the fitness consequences of reproductive timing, our results suggest that temporal genotype‐by‐environment interaction may maintain genetic variation and, because genetic variation occurs only among lineages, this simple system offers a unique opportunity for future tests of this mechanism.     

Effects of inbred/outbred crosses on progeny sex ratio in Silene latifolia (Caryophyllaceae)

Sex ratio polymorphism has been extensively studied in Silene latifolia, but it is neither known whether inbreeding (which is likely to occur under field conditions) affects it, nor which of the proposed mechanisms (Y degeneration, X-linked drive) is more important. Both mechanisms predict reduced pollen performance. In this study, females were crossed with pollen from related and unrelated males in single-donor and two-donor crosses, and the sex ratio of offspring (n = 866, 60 crosses), sons'in vitro pollen germination and sex ratios in parental families were scored. Flowers receiving only unrelated pollen produced a significant excess of sons. Sex ratios were not significantly correlated between generations. Sons'in vitro pollen germination was significantly negatively correlated with the 'sex-ratio phenotype' of maternal grandfathers, but not of fathers. This generation leap may be consistent with X-linked determinants because Y-linked determinants alone cannot explain it (grandfathers, fathers and sons share the same Y chromosome). Further work is required, but inbreeding and limited dispersal may lead to local accumulation of biasing factors, a process potentially countered by conditional shifts to produce more sons in pure outbred crosses.     

Variation in sex ratio, morph-specific reproductive ecology and an experimental test of frequency-dependence in the gynodioecious Kallstroemia grandiflora (Zygophyllaceae)

An enduring puzzle in gynodioecious species is the great variation in female frequency seen among populations. We quantified sex ratio in 44 populations of gynodioecious Kallstroemia grandiflora. Then, we measured pollinator visitation, pollen deposition, autonomous selfing rate and pollen limitation of females. Finally, using experimental populations, we tested whether female fitness responds to the frequency of female plants. We found broad variability in sex ratio among populations (0-44% female). Hermaphrodite flowers received more pollinator visits and pollen grains than females, and bagged hermaphrodite flowers produced fruits. However, we found no evidence of pollen limitation in females. In experimental populations, female plants showed no evidence of frequency-dependent pollinator visitation, fruit set, seed set or total seed mass. These results do not support frequency-dependent variation in fitness as a major mechanism affecting female frequencies in K. grandiflora. Within the context of this study, pollinators are abundant and pollinator movement appears to operate at a large enough scale to overcome the potential reproductive disadvantages of producing solely female flowers.     

Effect of superparasitism on the development,sex ratio and progeny ofBracon greeni Ashmead

Résumé Bracon greeni Ashmead est un ectoparasite larvaire de la noctuelleEublemma amabilis Moore, prédatrice sur laquiers en Inde. L'effet de superparasitisme sur le développement, le sex ratio et la descendance du parasite ont été étudiés en détail. Le pourcentage de développement du parasite décro?t quand le nombre de parasites développés par h?te augmente. La période de développement, de l'œuf à l'éclosion de l'adulte, est de 10, 85 jours et ce délai est un peu plus court quand l'h?te est fortement superparasité. La taille de la femelle adulte décro?t avec l'augmentation du nombre de parasites développés par h?te. Il y a prédominance du sexe femelle chez le parasite. Le superparasitisme ne para?t avoir que peu d'action sur le taux sexuel. Quand plus de trois parasites se partagent un seul h?te, la longévité de l'adulte et la fécondité du parasite sont réduits. De tels individus sont mauvais voiliers et inactifs dans leur comportement.      

Testing why the sex of the maternal parent affects seedling survival in a gynodioecious species

In gynodioecious plants, seed offspring from hermaphrodites often perform less well than those from females. This lower performance sometimes can be attributed to inbreeding by hermaphrodites or to relatively greater provisioning of individual seeds by females. However, these hypotheses are not explanatory when only outcrossing occurs and when individual seeds of the two morphs are equally well provisioned. Three hypotheses may explain the lower fitness of seed offspring from hermaphrodites in such cases. The morphology hypothesis states that the opportunity for gametophytic selection is lower within flowers of hermaphrodites compared to flowers on females, because the perfect flowers of hermaphrodites are relatively short-styled. The cytotype hypothesis states that the performance difference is directly caused by an individual's cytotype, whose frequency in the population may differ for the two sex morphs. The pleiotropy hypothesis states that negative pleiotropic effects of nuclear restorer alleles or alleles hitchhiking with them are expressed more often by offspring from hermaphrodites. We performed two experiments using the gynodioecious plant Silene acaulis to contrast these hypotheses. In our first experiment we contrasted the morphology and pleiotropy hypotheses by performing controlled pollinations and subsequently planting seeds in both the greenhouse and field. Hermaphrodites of S. acaulis can produce both pistillate and perfect flowers, which allowed us to determine whether flower morphology affects offspring survivorship independent of the sex of the maternal parent. We found that neither seed mass nor germination differed between seeds from females and hermaphrodites. Offspring from pistillate flowers on hermaphrodites did not differ significantly in their survival compared to offspring from perfect flowers on hermaphrodites, but had lower survivorship compared to offspring from pistillate flowers on females, refuting the morphology hypothesis. In a second experiment, we compared offspring survival of full-sibling pairs of females and hermaphrodites (who shared the same cytoplasm) to contrast the cytotype and pleiotropy hypotheses. We found that seed offspring from females and hermaphrodites that shared the same cytoplasm differed in their survival, which is counter to the prediction of the cytotype hypothesis. In both experiments, the sex of the maternal parent significantly affected offspring survival, with seed offspring from hermaphrodites surviving less well than those from females. These results support the pleiotropy hypothesis. We conclude by discussing alternative ways of thinking about negative pleiotropic effects of nuclear restorers or "the cost of restoration."     

Host-size-dependent feeding behaviour and progeny sex ratio of Anisopteromalus calandrae (Hym., Pteromalidae)

Host-size related feeding and oviposition behaviour, and allocation of progeny sex by Anisopteromalus calandrae (Howard) were tested on Sitophilus oryzae L. The parasitoid showed a host-size-dependent partition of feeding and oviposition behaviour, preferring small hosts for feeding, but large hosts for oviposition. Neither the mutual interference nor the host density showed any effect on the behaviour of the parasitoid. Allocation of progeny sex by the female parasitoid appeared to be based more likely on absolute than on relative host size encountered. A model for the progeny sex ratio was constructed based on: (1) ovipositional preference of the parasitoid on large hosts; (2) feeding preferentially on small hosts; and (3) host-size-related regulation of progeny sex ratio. The progeny sex ratio of the parasitoid predicted by the model was in close agreement with the observed value.