Cytotypes of Andropogon gerardii Vitman (Poaceae): fertility and reproduction of aneuploids

In many North American prairies, populations of Andropogon gerardii Vitman (Poaceae) are composed of hexaploid and enneaploid cytotypes (2 n  = 60, 90), with intermediates occurring occasionally. Under controlled pollination, the two common cytotypes can be crossed, producing progeny with a range of chromosome numbers. In an investigation of fertility and compatibilities of intermediate cytotypes, individuals with chromosome numbers between 60 and 90 were crossed with each other, with the 2 n  = 60 and 90 cytotypes, and with South American Andropogon species having 60 chromosomes. Regardless of cytotype, all A. gerardii plants had some fertility and virtually all crosses produced seeds. Cytotype is only partially predictive of fertility. Inter-specific hybrids between A. gerardii and South American hexaploid species were vigorous but sterile. Gene flow in natural A. gerardii populations of mixed cytotype probably involves plants of all cytotypes.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 95–103.     

Intraclonal variation in RNA viruses: generation, maintenance and consequences

This paper explores the evolutionary implications of the enormous variability that characterizes populations of RNA viruses and retroviruses. It begins by examining the magnitude of genetic variation in both natural and experimental populations. In natural populations, differences arise even within individual infected patients, with the per-site nucleotide diversity at this level ranging from < 1% to 6%. In laboratory populations, two viruses sampled from the same clone differed by ∼0.7% in their fitness. Three different mechanisms that may be important in maintaining viral genetic variability were tested: (1) Fisher's fundamental theorem, to compare the observed rate of fitness change with the extent of fitness-related variation within the same experimental populations; (2) magnitude of genomic mutation rate, to assess whether it correlated with fitness-related variation, as predicted by the mutation-selection balance hypothesis; (3) frequency-dependent selection, which affords rare genotypes an advantage. The paper concludes with a discussion of two evolutionary consequences of variability: the fixation of deleterious mutations by drift in small populations and the role of clonal interference in large ones.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 17–26.     

Ageing, reproductive value, and the evolution of lifetime fighting behaviour

The study of sexual selection has recently been enriched by an explicit life history focus. This lifetime perspective has much to offer the field; however, most existing interdisciplinary exchanges have restricted themselves to the context of mate choice and associated phenomena. The present study explores the consequences of an explicit life history view upon the evolution of male–male contest competition. Key to this view is the fact that fighting typically has lifetime consequences, and thus the costs involved with contests are best considered in lifetime currencies such as residual reproductive value. The rate of contemporary contest cost accrual may vary among contestants due to differences between them in what they 'stand to lose', in terms of future reproductive opportunities. It is also suggested that it may be fruitful to partition key life history parameters into components that are either dependent or independent of future choice of mating strategy. Using a simple simulation, it is demonstrated that the optimum pattern of lifetime contest participation may vary depending upon lifetime variation in 'strategy-independent' components of reproductive value (such as the rate of mortality experienced in contexts unrelated to mating behaviour). In line with previous models of age-specific sexual advertisement, increasing lifetime aggression appears as the most favoured strategy; however, young age aggression is predicted if resource holding potential is expected to decline with age irrespective of prior contest participation. The model is discussed, along with key concepts surrounding the life history viewpoint, in light of prior life history/sexual selection models and the existing empirical data regarding lifetime fighting strategies in the wild.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 565–578.     

Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals

In theory, parthenogenetic lineages have low evolutionary potential because they inexorably accumulate deleterious mutations and do not generate much genotypic diversity. As a result, most parthenogenetic taxa occupy the terminal nodes of phylogenetic trees. The rate and mode of development of parthenogenesis are important factors to consider when assessing its costs and benefits since they determine both the level of genetic diversity and the ecological adaptability of the resulting lineages. The origin of parthenogenesis is polyphyletic in many taxa, suggesting that genetic systems maintaining sexuality are often labile. In addition, the loss of sex may be achieved in several ways, leading to parthenogenetic lineages with distinct genetic profiles. This could then influence not only the fate of such lineages in the long term, but also the outcome of competition with their sexual counterparts in the short term. In this paper, we review the possible evolutionary routes to parthenogenesis based on a survey of the phylogenetic relationships between sexual and parthenogenetic lineages in a broad range of animals. We also examine the different mechanisms by which parthenogenetic lineages could arise, and discuss the influence of these mechanisms on both the genetic properties and the ecological life styles of the resulting lineages.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 151–163.     

Metamorphic duration: an under-studied variable in frog life histories

Over a narrow temperature range (25.5–28 °C), metamorphic duration in 14 taxonomically diverse frog species ranged from 2.0 to 7.3 days. For 12 species, duration was significantly related to tadpole body-size parameters, especially tail length. For the remainder, duration was longer than expected. Two patterns of tail regression were seen with either rapid or slow regression at the start. Most species lost about 30% of body wet weight during metamorphosis, but two were exceptional: X. laevis lost 58% and F. fitzgeraldi 13%. The results are discussed in the context of Wassersug & Sperry's (1977 ) hypothesis that selection acts to minimize metamorphic duration because metamorphosis is a highly vulnerable stage. Several other factors are outlined as possible determinants of metamorphic duration: developmental constraints, functional trade-offs, phylogeny, predator abundance, adaptive de-coupling, body condition and metabolic and water costs. There is evidence from the data for the operation of several of these factors, especially in species utilizing specialized habitats, and a plea is made for further study of metamorphic duration as an ecologically and evolutionarily important variable in amphibian life histories. Finally, the effects of different temperatures on metamorphic duration are presented for several species, and discussed in terms of local adaptation and reaction norms.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 261–272.     

A geometric morphometric assessment of the effects of environment and cladogenesis on the evolution of the turtle shell

In the largest group of extant turtles, the Testudinoidea, the acquisition of an aquatic or terrestrial way of life has occurred within two clades, allowing the study of homoplasy linked to environment (commonly named convergence). Here we appraise the respective importance of two sources of morphological variation: a major cladogenetic event and a major environmental shift (aquatic vs. terrestrial). The repeatability of the same evolutionary process (environmental change) allows an assessment of the weights of both natural selection and phylogenetic constraints on several morphological features of the shell. These sources of morphological variance on the complex shell structure were studied using geometric morphometrics. We depict the morphological variation of three parts of the turtle shell: epidermal carapace, bony carapace, and plastron. In the three structures, we found that both phylogeny and environment were significant sources of morphological variation, and geometric morphometrics allowed the pattern of morphological variation due to each effect to be assessed. The assessment of the homoplasy due to environment and of the pattern of morphological variability suggests that the carapace has undergone similar morphological changes between aquatic and terrestrial environments within the two clades. The radiation of the Testudinoidea is interpreted as an adaptive radiation.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 485–501.     

Gene flow and melanism in garter snakes revisited: a comparison of molecular markers and island vs. coalescent models

Within populations, the stochastic effect of genetic drift and deterministic effect of natural selection are potentially weakened or altered by gene flow among populations. The influence of gene flow on Lake Erie populations of the common garter snake has been of particular interest because of a discontinuous colour pattern polymorphism (striped vs. melanistic) that is a target of natural selection. We reassessed the relative contributions of gene flow and genetic drift using genetic data and population size estimates. We compared all combinations of two marker systems and two analytical approaches to the estimation of gene flow rates: allozymes (data previously published), microsatellite DNA (new data), the island model ( F _ST-based approach), and a coalescence-based approach. For the coalescence approach, mutation rates and sampling effects were also investigated. While the two markers produced similar results, gene flow based on F _ST was considerably higher (Nm > 4) than that from the coalescence-based method (Nm < 1). Estimates of gene flow are likely to be inflated by lack of migration-drift equilibrium and changing population size. Potentially low rates of gene flow (Nm < 1), small population size at some sites, and positive correlations of number of microsatellite DNA alleles and island size and between M , mean ratio of number of alleles to range in allele size, and island size suggest that in addition to selection, random genetic drift may influence colour pattern frequencies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 389–399.     

Marked variation in parasite resistance between two wild populations of the Trinidadian guppy, Poecilia reticulata (Pisces: Poeciliidae)

This is the first study to demonstrate significant differences between two natural Trinidadian guppy populations in susceptibility to a pathogenic monogenean parasite, Gyrodactylus turnbulli . Following experimental infection with an isogenic laboratory culture of G. turnbulli , fish from the Upper Aripo (UA) lost parasites more slowly and carried up to three times as many parasites at peak infection than did those from the Lower Aripo (LA). The UA population appeared to be more susceptible than the LA fish, even though fish of both populations were naïve to this particular laboratory strain of G. turnbulli and had not encountered any gyrodactylid infection for at least 65 days. The parasite infection reduced the feeding response and feeding activity of UA and LA fish to a similar extent. Our findings suggest that this ectoparasite may have an important impact on the evolutionary biology of guppies (and possibly of other teleosts), particularly as the prevalence of Gyrodactylus infection in natural populations may be as high as 50%. We discuss the role of parasite infections on natural and sexual selection, the good genes model and the implications for immunogenetics in small genetically isolated host populations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 645–651.     

Transitions in cuticular composition across a hybrid zone: historical accident or environmental adaptation?

Two subspecies of the grasshopper Chorthippus parallelus form a hybrid zone in the Pyrenees. Transitions across the zone, including changes in mating signals and reproductive isolation have been intensively studied. Cuticular pheromones have been identified as likely mate recognition signals. Since the major role of the cuticle is in waterproofing, environmental adaptation of cuticular composition has the interesting potential to generate assortative mating as an incidental by-product. We describe the pattern of variation in cuticular hydrocarbon blend in four transects through the hybrid zone. We find no evidence for a previously observed displaced cline in one blend component. There were differences between subspecies but these varied among transects and were small compared with variation between transects. We examined environmental variation within one transect and found a correlation between vegetation and cuticular composition, suggesting that environment influences the constitution of the cuticle, and hence natural selection may interact with mating signals in this species.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 193−201.     

Direct and correlated responses to selection for longevity in Drosophila buzzatii

The possible associations between longevity, early fecundity, and stress-resistance traits were explored using artificial selection on longevity in a laboratory population of Drosophila buzzatii . Three replicated lines were selected for increased lifespan (L lines) and compared with the respective unselected controls (C lines) after the 14th generation of selection. Mean longevity exhibited a significant response to selection. The baseline mortality tended to decrease in the L lines and a negative correlated response to longevity selection was found for early fecundity. Egg-to-adult developmental time increased in L lines. Longevity selection increased stress resistance for both high and low temperatures, as measured by heat knockdown resistance and chill-coma recovery. Starvation resistance also tended to be higher in L than in C lines. The results obtained are consistent with the hypothesis of trade-offs between longevity and early fecundity, and also suggest a trade-off association between adult longevity and developmental time. Correlated selection responses were generally consistent with correlations among the traits previously inferred from altitudinal clines for longevity and stress-resistance phenotypes.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 738–748.     

Arbuscular mycorrhizal fungi: genetics of multigenomic, clonal networks and its ecological consequences

Arbuscular mycorrhizal fungi are thought to have remained asexual for 400 million years, although recent studies have suggested that considerable genetic and phenotypic variation could potentially exist in populations. A brief discussion of these multigenomic organisms is presented.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 59–60.     

Food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology

Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 333–343.     

Plant clonality, mutation, diplontic selection and mutational meltdown

Apomixis is a very common characteristic in vascular plants. It occurs in two general forms: either subversion of the sexual system (agamospermous seeds or apogamous sporophytes in non-seed plants) or vegetative reproduction. In this communication, only the mutational consequences of vegetative reproduction are considered. Vegetative reproduction involves the replication of apical meristems, especially shoot apical meristems. Three general types of shoot apical meristems occur in the vascular plants: single tetrahedral apical initial, unstratified with impermanent initials and stratified with impermanent initials. Each meristem type has different consequences with regard to mutation, diplontic selection and the possibility of mutational meltdown.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 61–67.     

Effects of BRCA1 and BRCA2 mutations on female fertility

Women with BRCA1/2 mutations have a significantly higher lifetime risk of developing breast or ovarian cancer. We suggest that female mutation carriers may have improved fitness owing to enhanced fertility relative to non-carriers. Here we show that women who are carriers of BRCA1/2 mutations living in natural fertility conditions have excess fertility as well as excess post-reproductive mortality in relation to controls. Individuals who tested positive for BRCA1/2 mutations who linked into multi-generational pedigrees within the Utah Population Database were used to identify putative obligate carriers. We find that women born before 1930 who are mutation carriers have significantly more children than controls and have excess post-reproductive mortality risks. They also have shorter birth intervals and end child-bearing later than controls. For contemporary women tested directly for BRCA1/2 mutations, an era when modern contraceptives are available, differences in fertility and mortality persist but are attenuated. Our findings suggest the need to re-examine the wider role played by BRCA1/2 mutations. Elevated fertility of female mutation carriers indicates that they are more fecund despite their elevated post-reproductive mortality risks.     

Sexual size dimorphism and sex ratios in dragonflies (Odonata)

Sexual size dimorphism and biased sex ratios are common in animals. Rensch's rule states that sexual size dimorphism (SSD) would increase with body size in taxa where males are larger than females and decrease with body size in taxa where females are larger. We tested this trend in dragonflies (Odonata) by analysing body size of 21 species and found support for Rensch's rule. The increase in SSD with increasing size among species can be explained by sexual selection favouring large males. We also estimated the slope of the relationship between sex ratio and size ratio in populations of the 21 species. A negative slope would suggest that the larger sex suffers from high mortality in the larval stage, consistent with riskier foraging. The slope of this relationship was negative, but after correcting for phylogentic non-independence with independent contrasts the relationship was no longer statistically significant, perhaps because of phylogenic inertia or low sample size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 507–513.     

A new species of Valantia (Rubiaceae) from Spain

A new species of Valantia , V. lainzii Devesa & Ortega-Olivencia, endemic to the coastal zone of Granada (southern Spain) is described. The species recalls V. muralis L. in its general aspect, and V. deltoidea Brullo in the type of fructiferous body.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 331–335.     

DOES INCREASED MORTALITY FAVOR THE EVOLUTION OF MORE RAPID SENESCENCE?

It is widely believed (following the 1957 hypothesis of G. C. Williams) that greater rates of “extrinsic” (age- and condition-independent) mortality favor more rapid senescence. However, a recent analysis of mammalian life tables failed to find a significant correlation between minimum adult mortality rate and the rate of senescence. This article presents a simple theoretical analysis of how extrinsic mortality should affect the rate of senescence (i.e., the rate at which probability of mortality increases with age) under different evolutionary and population dynamical assumptions. If population dynamics are density independent, extrinsic mortality should not alter the senescence rate favored by natural selection. If population growth is density dependent and populations are stable, the effect of extrinsic mortality depends on the age specificity of the density dependence and on whether survival or reproduction (or both) are functions of density. It is possible that higher extrinsic mortality will increase the rate of senescence at all ages, decrease the rate at all ages, or increase it at some ages while decreasing it at others. Williams's hypothesis is most likely to be supported when density dependence acts primarily on fertility and does not differentially decrease the fertilities of older individuals. Patterns contrary to Williams's prediction are possible when density dependence acts primarily on the survival or fertility of later ages or when most variation in mortality rates is due to variation in nonextrinsic mortality.     

The importance of phylogeny and ecology in microgeographical variation in the morphology of four Canarian species of Aeonium (Crassulaceae)

The relative importance of natural selection in the diversification of organisms can be assessed indirectly using matrix correspondence. The present study determines the environmental and genetic correlates of microgeographical variation in the growth form, leaf form and flower morphology in populations of four Aeonium species from section Leuconium using partial regression methods. The phylogeny of the four species and the other 12 species in the section was deduced from amplified fragment length polymorphism (AFLP). Pubescence of floral organs and flower size correlate with the phylogeny while traits related to growth form, leaf form, flower construction and inflorescence size correlate with ecological factors. The variation in the latter four traits may therefore reflect selection by current ecological conditions while variation in pubescence and flower size may reflect historical events like neutral mutations, founder events and drift. Additionally, the morphological analyses revealed a large amount of variation in all traits within populations. This suggests a possible influence of microhabitat on the variation in morphology of Aeonium in the Canary Islands.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 76 , 521–533.