Sex allocation of females and hermaphrodites in the gynodioecious Geranium sylvaticum

Seed production and patterns of sex allocation were studied in female and hermaphroditic plants in two gynodioecious populations of Geranium sylvaticum (Geraniaceae). Females produced more flower buds and seeds than hermaphrodites in one of the two study populations. The other female traits measured (pistil biomass, seed number per fruit, individual seed mass) did not differ between the gender morphs. The relative seed fitness of hermaphrodites differed between the study populations, with hermaphrodites gaining less of their fitness through female function in the population with a high frequency of females. However, the amount and size of pollen produced by hermaphrodites did not differ between populations. The number of flower buds was positively correlated with seed production in females, whereas in hermaphrodites a positive correlation between number of buds and seed production was found in only one of the two study populations. These results suggest that fitness gain through female function is labile in hermaphrodites of this species, and is probably affected by environmental factors such as the sex ratio of the population.     

Preferences of pollinators and herbivores in gynodioecious Geranium sylvaticum

BACKGROUND AND AIMS: For the maintenance of gynodioecy (i.e. the coexistence of female and hermaphroditic plants), females need to compensate for the lack of pollen production through higher seed production or better progeny quality compared to hermaphrodites. In Geranium sylvaticum, females produce more seeds per flower than hermaphrodites. This difference in seed production might be modified by biological interactions with pollinators and herbivores that may favour one sex and thus affect the maintenance of gynodioecy. METHODS: Sexual dimorphism in flower size and flowering phenology, and in attractiveness to pollinators, pre-dispersal seed predators and floral herbivores were examined in natural populations of G. sylvaticum. KEY RESULTS: Pollinators preferred hermaphrodites 25 % more often than females in two of the three study populations, and floral herbivores attacked hermaphrodites 15 % more often than females in two of the six study populations. These preferences might be explained by the larger flower size of hermaphrodites. In contrast, seed predators did not prefer either sex. CONCLUSIONS: The data suggest that pollinator preference does not benefit females, whereas the higher floral herbivory of hermaphrodites might enhance the maintenance of females in G. sylvaticum. Thus, although the data support the view that ecological factors may contribute to the maintenance of gynodioecy, they also suggest that these contributions may vary across populations and that they may function in opposite directions.     

银叶老鹳草(Geranium sylvaticum)种群中诱导保护抗性和病原菌相互作用的研究（英）

本文通过自然植物种群接种实验,人工接种实验和野外调查研究了（Geranium sylvaticum)对老鹳草单孢锈菌（Uromyces geranii)（长生活史）的抗病性在受柄锈菌（Puccinia)（短生活史）感病前后和受2种柄锈菌（P.leveillei或P.morthieri)感病后有无区别,以及在同一季节里感染同一寄主植物的长生活史单孢锈菌和短生活史柄锈菌间的相互作用,病原菌间的相互作用使寄主植物产生诱导保护抗性。结果表明,P.leveillei可诱导寄主植物产生短时间的保护抗性,而P.morthieri可能诱导寄主植物产生长时期的保护抗性,诱导保护抗性可能是影响自然植物种群中植病式样的重要因素之一。     

Sexes in gynodioecious Geranium sylvaticum do not differ in their isotopic signature or photosynthetic capacity

• In gynodioecious plants, females are expected to produce more or better seeds than hermaphrodites in order to be maintained within the same population. Even though rarely measured, higher seed production can be achieved through differences in physiology.
• In this work, we measured sexual dimorphism in several physiological traits in the gynodioecious plant Geranium sylvaticum. Photosynthetic rate, stomatal conductance, transpiration rate, WUE and isotopic signatures were measured in plants growing in two habitats differing in light availability.
• Females have been reported to produce more seeds than hermaphrodites. However, we did not observe any significant difference in seed output between the sexes in these experimental populations. Similarly, the sexes did not differ in any physiological trait measured. Seed production was strongly limited by light availability. Likewise, differences between plants growing in full light versus low light were detected in most physiological parameters measured.
• Our results show that the sexes in G. sylvaticum do not show any evidence of sexual dimorphism in physiology, which concurred with a lack of sexual differences in seed output.

     

Variable mycorrhizal benefits on the reproductive output of Geranium sylvaticum,with special emphasis on the intermediate phenotype

In several gynodioecious species, intermediate sex between female and hermaphrodite has been reported, but few studies have investigated fitness parameters of this intermediate phenotype. Here, we examined the interactions between plant sex and arbuscular mycorrhizal (AM) fungal species affecting the reproductive output of Geranium sylvaticum, a sexually polymorphic plant species with frequent intermediate sexes between females and hermaphrodites, using a common garden experiment. Flowering phenology, AM colonisation levels and several plant vegetative and reproductive parameters, including seed and pollen production, were measured. Differences among sexes were detected in flowering, fruit set, pollen production and floral size. The two AM species used in the present work had different effects on plant fitness parameters. One AM species increased female fitness through increasing seed number and seed mass, while the other species reduced seed mass in all sexes investigated. AM fungi did not affect intermediate and hermaphrodite pollen content in anthers. The three sexes in G. sylvaticum did not differ in their reproductive output in terms of total seed production, but hermaphrodites had potentially larger fathering ability than intermediates due to higher anther number. The ultimate female function – seed production – did not differ among the sexes, but one of the AM fungi used potentially decreased host plant fitness. In addition, in the intermediate sex, mycorrhizal symbiosis functioned similarly in females as in hermaphrodites.     

In search of better strategies of using activity traps to collect Dytiscidae and Noteridae (Coleoptera): a case study of a local fauna in central European Russia

Activity traps (ATs) are a widely used method of sampling water beetles (Coleoptera). This case study attempts to investigate the effect of mouth size and trapping effort on the performance of the ATs and reveal sex ratios and proportions of morphs in dimorphic females in populations of Dytiscidae and Noteridae in one particular area of Udomelsky District, Tver Oblast, Russia. ATs are more selective compared to hand netting but make it possible to collect a greater number of abundant species with relatively little expenditures of effort and time. The width of the trap mouth has no effect on efficiency. Long-time (48?h exposure) traps are not more efficient than short-time (24?h) ones, in terms of the relative intensity of collecting individuals and species. The applied number of ATs (up to 330 trap-days) was not sufficient to reveal the full diversity of diving beetles in the studied waterbodies.     

Factors determining survival of European eels in two unexploited sub‐populations

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Sex investment ratios in monogyne and polygyne populations of the fire ant Solenopsis invicta

Both monogyne (single queen per colony) and polygyne (multiple queens per colony) populations of the fire ant Solenopsis invicta are good subjects for tests of kin selection theory because their genetic and reproductive attributes are well-characterized, permitting quantitative predictions about the degree to which sex investment ratios should be female-biased if workers and not queens control reproductive allocation. In the study populations, an investment ratio of 3 females: 1 male is predicted (a proportional investment in females of 0.75) in the monogyne form, whereas a proportional investment in females between 0.637 and 0.740 is expected in the polygyne form. To test these predictions, colonies from a single population of each social form were collected and censused during three different seasons. Consistent with their alternative modes of colony founding, monogyne colonies invested more in reproduction (sexual production) and less in growth/maintenance (worker production) than did the polygyne colonies. Overall, the sex investment ratios were female-biased in both forms, although there was considerable seasonal variation. After adjusting for sex-specific energetic costs, the proportional investment in females was 0.607 in the monogyne population, a value in between those expected under complete control by either the queen or the workers. However, when combined with data from four other previously studied monogyne populations in the U.S.A., the mean investment ratio did not differ significantly from the value predicted if workers have exclusive control. In the polygyne population, the proportional investment in females of 0.616 was consistent with the level of female bias expected under partial to complete worker control, although the potential influence of two confounding factors — possible contact with monogyne colonies and the preponderance of sterile diploid males — weakens this conclusion somewhat. Taken as a whole, the sex investment ratios of monogyne and polygyne populations of S. invicta are consistent with at least partial worker control. Of several ultimate and proximate explanations that have been proposed to explain inter-colonial variation in the sex investment ratio, only the effect of the primary sex ratio (female-determined eggs: male-determined eggs) laid by the queen appears to account for the observed variation among monogyne colonies. In the polygyne population, there is limited support for the hypothesis that greater resource abundance favors investment in females.     

Environmental stressors affect sex ratios in sexually dimorphic plant sexual systems

• Revealing the environmental pressures determining the frequency of females amongst populations of sexually dimorphic plants is a key research question. Analyses of sex ratio variation have been mainly done in dioecious plants, which misses key plant sexual systems that might represent intermediate stages in the evolution of dioecy from hermaphroditism.
• We investigated female frequency across populations of sexually dimorphic plant species in relation to environmental stressors (temperature, precipitation), totaling 342 species, 2011 populations, representing 40 orders and three different sexual systems (dioecy, gynodioecy and subdioecy). We also included the biome where the population was located to test how female frequency may vary more broadly with climate conditions.
• After correcting for phylogeny, our results for gynodioecious systems showed a positive relationship between female frequency and increased environmental stress, with the main effects being temperature‐related. Subdioecious systems also showed strong positive relationships with temperature, and positive and negative relationships related to precipitation, while no significant effects on sex ratio in dioecious plants were detected.
• Combined, we show that female frequencies in an intermediate sexual system on the pathway from hermaphroditism to dioecy respond strongly to environmental stressors and have different selective agents driving female frequency.

     

Sex investment ratios in eusocial Hymenoptera support inclusive fitness theory

Inclusive fitness theory predicts that sex investment ratios in eusocial Hymenoptera are a function of the relatedness asymmetry (relative relatedness to females and males) of the individuals controlling sex allocation. In monogynous ants (with one queen per colony), assuming worker control, the theory therefore predicts female‐biased sex investment ratios, as found in natural populations. Recently, E.O. Wilson and M.A. Nowak criticized this explanation and presented an alternative hypothesis. The Wilson–Nowak sex ratio hypothesis proposes that, in monogynous ants, there is selection for a 1 : 1 numerical sex ratio to avoid males remaining unmated, which, given queens exceed males in size, results in a female‐biased sex investment ratio. The hypothesis also asserts that, contrary to inclusive fitness theory, queens not workers control sex allocation and queen–worker conflict over sex allocation is absent. Here, I argue that the Wilson–Nowak sex ratio hypothesis is flawed because it contradicts Fisher's sex ratio theory, which shows that selection on sex ratio does not maximize the number of mated offspring and that the sex ratio proposed by the hypothesis is not an equilibrium for the queen. In addition, the hypothesis is not supported by empirical evidence, as it fails to explain ‘split’ (bimodal) sex ratios or data showing queen and worker control and ongoing queen–worker conflict. By contrast, these phenomena match predictions of inclusive fitness theory. Hence, the Wilson–Nowak sex ratio hypothesis fails both as an alternative hypothesis for sex investment ratios in eusocial Hymenoptera and as a critique of inclusive fitness theory.     

Sex ratios observed in 80 species of parrots

A number of potential evolutionary and physiological factors may be involved in avian sex ratio bias so that under certain conditions a sex ratio bias may favour males or females within a population. In addition different factors may be important in manipulating sex ratio bias through the different life stages. In this study sex ratio bias was examined in a total of 16 570 captive parrots, representing 80 species, many of which are endangered in the wild, using database records originating form commercial laboratories that offer genetic sexing. Within the species examined 72% showed a male bias this was significant in three species, when adjusted for multiple comparisons. This preliminary study is limited due to lack of data on the age of the individuals sampled. However, the large dataset do suggest that this phenomenon should be further considered by investigators working at a species level where such data can be collected.     

New Triassic beetles (Coleoptera) from northern European Russia

Beetle remains from the Triassic Khey-Yaga locality, Nenets National District, Korotaikha Basin, Nyadeita Formation, Olenekian-Anisian are described. Only isolated elytra have been found. Tetracoleus tshalyshevi gen. et sp. nov. and T. minimus sp. nov. are described and assigned to Tricoleidae, and six new species are described in formal taxa. Tetracoleus is close to Permian tricoleids just found in the Newcastle Group of Australia and the Vyatkian Aristovo locality of northern European Russia. The composition of elytra in the locality is closer to that of Lower Triassic localities than other Middle Triassic localities and characterizes the beginning of a recovery after the Permian-Triassic ecological crisis. Longxianocupes Hong in Liu, Liu et Hong, 1985 is shown to be a junior synonym of Sogdelytron Ponomarenko, 1969. The species Palademosyne ovum Ponomarenko, 2004, P. elongatum Ponomarenko, 2004, and P. latum Ponomarenko, 2004 are transferred to the genus Pseudochrysomelites Handlirsch, 1906.     

Sex ratio variation in female-biased populations of Notostracans

Females from female-biased populations of the notostracan Triops newberryi produce viable eggs when reared in isolation. Clutches produced under such conditions exhibit sex-ratio polymorphism. One category of females (monogenics) produces only female offspring; the second category (amphigenics) produces clutches with a sex ratio of 3 females: 1 male. The relative proportions of the two categories of females varies significantly between populations and is correlated with population sex ratio. This correlation, and the pattern of offspring distribution in amphigenics, suggests that sex is determined by an autosomal Mendelian gene locus for which the male-determining allele is recessive. Limited pedigree analysis of lineages under selfing support the genetic model.     

Late Permian Discordichthyiformes (Osteichthyes) from European Russia

New Late Permian fishes combining features characteristic of several classes are analyzed and redescribed. They were assigned to the family Discordichthyidae of the order Discordichthyiformes comprising three genera and four species: Mutovinia stella Minich, Geryonichthys longus A. Minich, G. burchardi A. Minich, and Discordichthys spinifer A. Minich (Minikh, 1998). A new species, Mutovinia sennikovi sp. nov., is described. Due to rapid evolutionary replacement, discordichthyids are rather useful stratigraphically. The photographs of the skeleton and membrane bones of these fishes are published for the first time.     

Sex ratios in natural populations ofAphelinus mali (Hym.: Aphelinidae) in relation to host size and host density

In an apple orchard at Armidale, the Northern Tablelands of NSW, population sex ratios ofAphelinus mali (Haldeman), an endoparasitoid of the woolly apple aphid,Eriosoma lanigerum (Hausmann) varied from 0.51 (proportion of males) at low host densities to female-biased at high host densities (proportion of males ranged from 0.35–0.39). This shift in sex ratio seems to be caused by the differences in allocation of sons and daughters to hosts of different sizes. In the fieldA. mali parasitizes all life stages (four nymphal instars and adult) of the woolly aphid upon encountering. According to Hughes'(1979) optimal diet model, such general host acceptance seems to be the best strategy. However, it allows the host nymphs or adults to continue to develop or reproduce until about to mummify (pupate). No mortality was observed when first or second-instar hosts were parasitized in the laboratory. Field collected small mummified hosts yielded male-biased sex ratios whereas large mummified hosts produced mainly females. In the laboratory, progeny from smaller hosts (first to third-instars) produced sex ratios which were not significantly different from 0.5 whereas progeny from larger hosts (third and fourth-instars) produced female-biased sex ratio. During winter (June–August) and early spring (September–October) when the host populations in the orchard were predominantly nymphs, the parasitoid tended to allocate equal resources to male and female offspring. In contrast, at peak population densities in summer and autumn (December–May) when larger hosts were available, the sex ratios were female-biased. The host size ofE. lanigerum andA. mali is, therefore, an important component in the dynamics of host-parasitoid interactions.     

Sex ratio variation in gynodioecious Lobelia siphilitica: effects of population size and geographic location

Variation in population sex ratio can be influenced by natural selection on alternate sex phenotypes as well as nonselective mechanisms, such as genetic drift and founder effects. If natural selection contributes to variation in population sex ratio, then sex ratio should covary with resource availability or herbivory. With nonselective mechanisms, sex ratio should covary with population size. We estimated sex ratio, resource availability, herbivory and size of 53 populations of gynodioecious Lobelia siphilitica. Females were more common in populations with higher annual temperatures, lower soil moisture and lower predation on female fruits, consistent with sex-specific selection. Females were also more common in small populations, consistent with drift, inbreeding or founder effects. However, small populations occurred in areas with higher temperatures than large populations, suggesting that female frequencies in small populations could be caused by sex-specific selection. Both selective and nonselective mechanisms likely affect sex ratio variation in this gynodioecious species.     

Sex reversal and primary sex ratios in the common frog (Rana temporaria)

Sex reversal has been suggested to have profound implications for the evolution of sex chromosomes and population dynamics in ectotherms. Occasional sex reversal of genetic males has been hypothesized to prevent the evolutionary decay of nonrecombining Y chromosomes caused by the accumulation of deleterious mutations. At the same time, sex reversals can have a negative effect on population growth rate. Here, we studied phenotypic and genotypic sex in the common frog (Rana temporaria) in a subarctic environment, where strongly female‐biased sex ratios have raised the possibility of frequent sex reversals. We developed two novel sex‐linked microsatellite markers for the species and used them with a third, existing marker and a Bayesian modelling approach to study the occurrence of sex reversal and to determine primary sex ratios in egg clutches. Our results show that a significant proportion (0.09, 95% credible interval: 0.04–0.18) of adults that were genetically female expressed the male phenotype, but there was no evidence of sex reversal of genetic males that is required for counteracting the degeneration of Y chromosome. The primary sex ratios were mostly equal, but three clutches consisted only of genetic females and three others had a significant female bias. Reproduction of the sex‐reversed genetic females appears to create all‐female clutches potentially skewing the population level adult sex‐ratio consistent with field observations. However, based on a simulation model, such a bias is expected to be small and transient and thus does not fully explain the observed female‐bias in the field.     

New Tatarian representatives of the subfamily Chaulioditinae (Insecta: Grylloblattida: Chaulioditidae) from European Russia

New Late Permian grylloblattids of the subfamily Chaulioditinae (Chaulioditidae) are described: Chauliodites afonini sp. nov. (Sokovka locality, Vladimir Region; Vyatkian Stage), Ch. gomankovi sp. nov. (Novo-Aleksandrovka locality, Orenburg Region; Severodvinian Stage), and Ch. ponomarenkoi sp. nov. (Isady locality, Vologda Region, upper Severodvinian Stage). Stratigraphic distribution of the subfamily is discussed.     

Adult sex ratios in wild bird populations

Offspring sex ratios in wild bird populations, and the extent to which they vary from the equality expected by random genotypic sex determination, have received much recent attention. Adult sex ratios (ASRs) in wild birds, on the other hand, remain very poorly described, and many of the questions about them posed by Ernst Mayr in 1939 remain unanswered. This review assesses population-level sex ratio patterns in wild bird populations, with an emphasis on the ASR. A quantitative assessment of over 200 published estimates of ASR, covering species from a wide range of taxa, regions and habitats, supported Mayr's assertion that skewed ASRs are common in wild bird populations. On average, males outnumbered females by around 33%, and 65% of published estimates differed significantly from equality. In contrast, population-level estimates of offspring sex ratio in birds did not generally differ from equality, and mean ASR across a range of wild mammal species was strongly female-skewed. ASR distortion in birds was significantly more severe in populations of globally threatened species than in non-threatened species, a previously undescribed pattern that has profound implications for their monitoring and conservation. Higher female mortality, rather than skewed offspring sex ratio, is the main driver of male-skewed ASRs in birds, and the causes and implications of this are reviewed. While estimates of ASR in wild bird populations may be subject to a number of biases, which are discussed, there is currently no quantitative evidence that an ASR of one male to one female represents the norm in birds. A better understanding and reporting of ASRs in wild bird populations could contribute greatly to our understanding of population processes and could contribute much to theoretical and applied research and conservation.