The memory remains: application of historical DNA for scaling biodiversity loss

Few species worldwide have attracted as much attention in relation to conservation and sustainable management as Pacific salmon. Most populations have suffered significant reductions, many have disappeared, and even entire evolutionary significant units (ESUs) are believed to have been lost. Until now, no ‘smoking gun’ in terms of direct genetic evidence of the loss of a salmon ESU has been produced. In this issue of Molecular Ecology, Iwamoto et al. (2012) use microsatellite analysis of historical scale samples of Columbia River sockeye salmon (Oncorhynchus nerka) from 1924 ( Fig. 1 ) to ask the pertinent question: Do the historical samples contain salmon from extirpated populations or ESUs? They identified four genetic groups in the historical samples of which two were almost genetically identical to contemporary ESUs in the river, one showed genetic relationship with a third ESU, but one group was not related to any of the contemporary populations. In association with ecological data, the genetic results suggest that an early migrating Columbia River headwater sockeye salmon ESU has been extirpated. The study has significant importance for conservation and reestablishment of sockeye populations in the Columbia River, but also underpins the general significance of shifting baselines in conservation biology, and how to assess loss of genetic biodiversity. The results clearly illustrate the huge and versatile potential of using historical DNA in population and conservation genetics. Because of the extraordinarily plentiful historical samples and rapid advances in fish genomics, fishes are likely to spearhead future studies of temporal ecological and population genomics in non‐model organisms.

Figure 1 Open in figure viewer PowerPoint (a) Kokanee sampling site between Columbia and Windermere lakes on the upper Columbia River at Fairmont Hot Springs, British Columbia, Canada. (b) Bureau of Fisheries scale books that contained sockeye salmon (locally called ‘blueback’ salmon) scales collected from commercial fisheries during the 1920s in the lower Columbia River. (c) Kokanee on spawning beds in Kuskanax Creek, a tributary to Upper Arrow Lake, British Columbia. Photo credit Rick Gustafson and Jim Myers.     

Evolutionary, behavioural and molecular ecology must meet to achieve long-term conservation goals

Founder populations in reintroduction programmes can experience a genetic bottleneck simply because of their small size. The influence of reproductive skew brought on by polygynous or polyandrous mating systems in these populations can exacerbate already difficult conservation genetic problems, such as inbreeding depression and loss of adaptive potential. Without an understanding of reproductive skew in a target species, and the effect it can have on genetic diversity retained over generations, long‐term conservation goals will be compromised. In this issue of Molecular Ecology, Miller et al. (2009a) test how founder group size and variance in male reproductive success influence the maintenance of genetic diversity following reintroduction on a long‐term scale. They evaluated genetic diversity in two wild populations of the iconic New Zealand tuatara ( Fig. 1 ), which differ greatly in population size and genetic diversity, and compared this to genetic diversity in multiple founder populations sourced from both populations. Population viability analysis on the maintenance of genetic diversity over 400 years (10 generations) demonstrated that while the loss of heterozygosity was low when compared with both source populations (1–14%), the greater the male reproductive skew, the greater the predicted losses of genetic diversity. Importantly however, the loss of genetic diversity was ameliorated after population size exceeded 250 animals, regardless of the level of reproductive skew. This study demonstrates that highly informed conservation decisions could be made when you build on a solid foundation of demographic, natural history and behavioural ecology data. These data, when informed by modern population and genetic analysis, mean that fundamental applied conservation questions (how many animals should make up a founder population?) can be answered accurately and with an eye to the long‐term consequences of management decisions.

Figure 1 Open in figure viewer PowerPoint Large adult male tuatara attacking a smaller male. Photo by Jeanine Refsnider.     

Tales from scales: old DNA yields insights into contemporary evolutionary processes affecting fishes

Salmon and trout populations are suffering declines in abundance and diversity over much of their range around the Atlantic and Pacific rims as a consequence of many factors. One method of dealing with the decline has been to produce them in hatcheries but the wisdom of this approach has been hotly debated (e.g. Hilborn & Winton 1993 ; Waples 1999 ; Brannon et al. 2004 ). One concern is that domesticated hatchery strains will interbreed with locally adapted wild fish; but how do we study the genetic effects if the introgression might have occurred in the past? Hansen (2002 ) used DNA isolated from archived scales from brown trout, Salmo trutta ( Fig. 1 ), to show that domesticated trout had, to varying degrees, genetically introgressed with wild, native trout in two Danish rivers. Extending that study, Hansen et al. (2009 ) have examined DNA from brown trout scales in six Danish rivers collected during historical (1927–1956) and contemporary (2000–2006) periods and from two hatchery source populations, to assess the effects of stocking nonlocal strains of hatchery trout and declining abundance on genetic diversity. Using 21 microsatellite loci, they revealed that genetic change occurred between the historic and contemporary time periods. Many populations appeared to have some low level of introgression from hatchery stocks and two populations apparently experienced high levels of introgression. Hansen et al. (2009 ) also showed that population structure persists in contemporary populations despite apparent admixture and migration among populations, providing evidence that the locally adapted populations have struggled against and, to some extent, resisted being overwhelmed by repeated introductions of and interbreeding with non‐native, hatchery‐produced conspecifics.

Figure 1 Open in figure viewer PowerPoint Photograph of a brown trout, Salmo trutta, one of many species of salmon and trout (Family Salmonidae) that are widely produced in hatcheries to enhance recreational and commercial fisheries. Photo by Peter Westley, Memorial University of Newfoundland, St John's NFLD, Canada.     

Heritability not missing—genetic basis of sexual weaponry uncovered

The chase to uncover the genetic underpinnings of quantitative traits of ecological and evolutionary importance has been on for a good while. However, the potential power of genome‐wide association studies (GWAS) as an approach to identify genes of interest in wild animal populations has remained untapped. Setting technical and economic explanations aside, the sobering lack of success in human GWAS might have fed this restraint. Namely, while GWAS have successfully identified genetic variants associated with hundreds of complex traits (e.g. Ku et al. 2010 ), these variants have generally captured only a low percentage of variance in traits known to be highly heritable—an observation came to be known as the ‘missing heritability’ ( Maher 2008 ; Aulchenko et al. 2009 ). Hence, if the vastly resourced human studies have been unsuccessful (but see: Yang et al. 2010 ), why should we expect that less resourced studies of wild animal populations would be able do better? In this issue of Molecular Ecology, Johnston et al. (2011) prove this line of thinking wrong. In an impressive and what may well be the most advanced gene mapping study ever performed in a wild population, they identify a single locus (RXFP2) responsible for explaining horn phenotype in feral domestic sheep from St Kilda ( Fig. 1 ). This same locus is also shown to account for up to 76% of additive genetic variance in horn size in male sheep: this contrasts sharply with most human GWAS where mapped loci explain only a modest proportion of genetic variation in a given trait.

Figure 1 Open in figure viewer PowerPoint The Soay sheep of the St Kilda archipelago are a primitive feral breed of domestic sheep. Pictured are a male with vestigial horns (=‘scurred’; left) and two normal‐horned males (centre and right). Photograph courtesy of Peter Korsten.     

Can we stop transgenes from taking a walk on the wild side?

Whether the potential costs associated with broad‐scale use of genetically modified organisms (GMOs) outweigh possible benefits is highly contentious, including within the scientific community. Even among those generally in favour of commercialization of GM crops, there is nonetheless broad recognition that transgene escape into the wild should be minimized. But is it possible to achieve containment of engineered genetic elements in the context of large scale agricultural production? In a previous study, Warwick et al. (2003) documented transgene escape via gene flow from herbicide resistant (HR) canola (Brassica napus) into neighbouring weedy B. rapa populations ( Fig. 1 ) in two agricultural fields in Quebec, Canada. In a follow‐up study in this issue of Molecular Ecology, Warwick et al. (2008) show that the transgene has persisted and spread within the weedy population in the absence of selection for herbicide resistance. Certainly a trait like herbicide resistance is expected to spread when selected through the use of the herbicide, despite potentially negative epistatic effects on fitness. However, Warwick et al.'s findings suggest that direct selection favouring the transgene is not required for its persistence. So is there any hope of preventing transgene escape into the wild?

Figure 1 Open in figure viewer PowerPoint Weedy Brassica rapa (orange flags) growing in a B. napus field. (Photo: MJ Simard)     

Beyond splitting hares and rabbiting on about major histocompatibility complex complexity

The genes of the major histocompatibility complex (MHC) have become the target of choice for studies wishing to examine adaptively important genetic diversity in natural populations. Within Molecular Ecology alone, there have been 71 papers on aspects of MHC evolution over the past few years, with an increasing year on year trend. This focus on the MHC is partly driven by the hypothesized links between MHC gene dynamics and ecologically interesting and relevant traits, such as mate choice and host–parasite interactions. However, an ability to pin down the evolutionary causes and ecological consequences of MHC variation in natural populations has proven challenging and has been hampered by the very issue that is attractive about MHC genes – their high levels of diversity. Linking high levels of MHC diversity to ecological factors in inherently complex natural populations requires a level of experimental design and analytical rigour that is extremely difficult to achieve owing to a plethora of potentially confounding and interacting variables. In this issue of Molecular Ecology, Smith et al. (2010) elegantly overcome the challenge of detecting complex interactions in complex systems by using an intricate analytical approach to demonstrate a role for MHC in the reproductive ability of a natural population of the European hare Lepus europaeus ( Fig. 1 ). Also in this issue, Oppelt et al. (2010) demonstrate a role for MHC variation in determining levels of hepatic coccidian infection in the European rabbit Oryctolagus cuniculus ( Fig. 2 ).

Figure 1 Open in figure viewer PowerPoint The European hare (Lepus europaeus).     

Revolution in food web analysis and trophic ecology: diet analysis by DNA and stable isotope analysis

Characterization of energy flow in ecosystems is one of the primary goals of ecology, and the analysis of trophic interactions and food web dynamics is key to quantifying energy flow. Predator‐prey interactions define the majority of trophic interactions and food web dynamics, and visual analysis of stomach, gut or fecal content composition is the technique traditionally used to quantify predator‐prey interactions. Unfortunately such techniques may be biased and inaccurate due to variation in digestion rates ( Sheppard & Hardwood 2005 ); however, those limitations can be largely overcome with new technology. In the last 20 years, the use of molecular genetic techniques in ecology has exploded ( King et al. 2008 ). The growing availability of molecular genetic methods and data has fostered the use of PCR‐based techniques to accurately distinguish and identify prey items in stomach, gut and fecal samples. In this month’s issue of Molecular Ecology Resources, Corse et al. (2010) describe and apply a new approach to quantifying predator‐prey relationships using an ecosystem‐level genetic characterization of available and consumed prey in European freshwater habitats ( Fig. 1a ). In this issue of Molecular Ecology, Hardy et al. (2010) marry the molecular genetic analysis of prey with a stable isotope (SI) analysis of trophic interactions in an Australian reservoir community ( Fig. 1b ). Both papers demonstrate novel and innovative approaches to an old problem – how do we effectively explore food webs and energy movement in ecosystems?

Figure 1 Open in figure viewer PowerPoint The aquatic habitats used for two studies of diet and trophic interactions that employed molecular genetic and stable isotope analyses. Panel a: Example of Rhone basin habitat (France) where fish diet was determined using PCR to classify prey to a series of ecological clades (photo by Emmanuel Corse). Panel b: A weir pool on the lower Murray River (Australia) where food web and prey use was evaluated using a combination of advanced molecular genetic and stable isotope analyses (photo credit: CSIRO).     

Cultivated and weedy rice interactions and the domestication process

Examining the targets of selection in crop species and their wild and weedy relatives sheds light on the evolutionary processes underlying differentiation of cultivars from progenitor lineages. On one hand, human‐mediated directional selection in crops favours traits associated with the streamlining of controllable and predictable monoculture practices alongside selection for desired trait values. On the other hand, natural selection in wild and especially weedy relatives presumably favours trait values that increase the probability of escaping eradication. Gene flow between crops and wild species may also counter human‐mediated selection, promoting the evolution and persistence of weedy forms. In this issue, two studies from a group of collaborators examine diversity and divergence patterns of genes underlying two traits associated with red rice (Oryza sp.), the conspecific relative of cultivated rice (Oryza sativa) that is a non‐native weed (see Fig. 1 ). In the first study by Gross et al. (2010) , genetic variation in the major gene underlying the hallmark red pigmentation characterizing most weedy rice (Rc) is found to have a pattern consistent with non‐reversion from U.S. cultivated rice (i.e. the cultivar did not ‘go feral’). This suggests that U.S. weedy rice is not an escaped lineage derived from U.S. cultivated rice populations; weedy rice likely differentiated prior to the selective sweep occurred in this gene within cultivated rice populations. Using the major seed shattering locus sh4 gene and the neighbouring genomic region, Thurber et al. (2010) track the molecular evolutionary history of the high shattering phenotype, a trait contributing dramatically to the success of crop selection in cultivated rice as well as the persistence and expansion of weedy red rice. In this study, the shared fixation of a sh4 mutation in both cultivated rice and weedy rice indicates that weedy rice arose subsequent to the strong selective sweep leading to significant reduction in seed shattering in cultivated rice.

Figure 1 Open in figure viewer PowerPoint A weedy, brown hulled red rice individual with long awns surrounded by a field of cultivated rice (photo by A. Lawton‐Rauh).     

Selection and sticklebacks

Over the last decade, there has been increasing circumstantial evidence for the action of natural selection in the genome, arising largely from molecular genetic surveys of large numbers of markers. In nonmodel organisms without densely mapped markers, a frequently used method is to identify loci that have unusually high or low levels of genetic differentiation, or low genetic diversity relative to other populations. The paper by Mäkinen et al. (2008a) in this issue of Molecular Ecology reports the results of a survey of microsatellite allele frequencies at more than 100 loci in seven populations of the three‐spined stickleback (Gasterosteus aculeatus). They show that a microsatellite locus and two indel markers located within the intron of the Eda gene, known to control the number of lateral plates in the stickleback ( Fig. 1 ), tend to be much more highly genetically differentiated than other loci, a finding that is consistent with the action of local selection. They identify a further two independent candidates for local selection, and, most intriguingly, they further suggest that up to 15% of their loci may provide evidence of balancing selection.

Figure 1 Open in figure viewer PowerPoint Three‐spined stickleback (Gasterosteus aculeatus).     

Fungal defences against animal antagonists – lectins &; more

Unlike in the laboratory, in nature fungi are exposed to antagonists including competitors, pathogens, parasites and predators. In this issue of Molecular Ecology, an exciting paper by Bleuler‐Martínez et al. (2011) has unearthed on one of the processes used by fungi to protect themselves against animal antagonists. The authors from Markus Künzler′s group at the ETH Zürich, Switzerland present direct evidence that filamentous fungi possess an inducible resistance mechanism against predators and parasites. This is based on cytoplasmic lectins, which specifically bind to glycans of these predators and parasites, and thus provide toxicity against them. These lectins are expressed at high levels in fruiting bodies and sclerotia of these fungi ( Fig. 1 ). While there have been previous suggestions and efforts to implicate mycotoxins such as sterigmatocystin into fungal defence mechanisms and as an evolutionary force (see Kempken & Rohlfs 2010 ), the data presented by Markus Künzler′s group highlight the ecological relevance of lectins in defending fungi from parasites and fungivorous animals. As such this paper provides important ecological clues and suggests that secondary metabolites are not the sole player in fungal–animal competition. It rather appears that fungi have evolved several lines of defence against antagonistic organisms.

Figure 1 Open in figure viewer PowerPoint Fungi protect their fruiting bodies by the use of lectin (picture of Aleuria aurantia provided by Taylor F. Lockwood).     

Modelling evolutionary processes in small populations: not as ideal as you think

Evolutionary processes are routinely modelled using ‘ideal’ Wright–Fisher populations of constant size N in which each individual has an equal expectation of reproductive success. In a hypothetical ideal population, variance in reproductive success (V_k) is binomial and effective population size (N_e) = N. However, in any actual implementation of the Wright–Fisher model (e.g., in a computer), V_k is a random variable and its realized value in any given replicate generation () only rarely equals the binomial variance. Realized effective size () thus also varies randomly in modelled ideal populations, and the consequences of this have not been adequately explored in the literature. Analytical and numerical results show that random variation in  and  can seriously distort analyses that evaluate precision or otherwise depend on the assumption that  is constant. We derive analytical expressions for Var(V_k) [4(2N – 1)(N – 1)/N³] and Var(N_e) [N(N – 1)/(2N – 1) ≈ N/2] in modelled ideal populations and show that, for a genetic metric G = f(N_e), Var(?) has two components: Var_Gene (due to variance across replicate samples of genes, given a specific ) and Var_Demo (due to variance in ). Var(?) is higher than it would be with constant N_e = N, as implicitly assumed by many standard models. We illustrate this with empirical examples based on F (standardized variance of allele frequency) and r² (a measure of linkage disequilibrium). Results demonstrate that in computer models that track multilocus genotypes, methods of replication and data analysis can strongly affect consequences of variation in . These effects are more important when sampling error is small (large numbers of individuals, loci and alleles) and with relatively small populations (frequently modelled by those interested in conservation).     

Opening Pandora's box: comparative studies of genetic mating systems reveal reproductive complexity

Genetic analyses of realized reproductive success have fundamentally changed our understanding of mating behaviour in natural systems. While behavioural ecologists have long been interested in what factors influence mating behaviour, early studies were limited to direct observations of matings and thus provided an incomplete picture of reproductive activity. Genetic assessments of parentage have revolutionized the study of reproductive behaviour, revealing that many individuals engage in extra‐pair copulations ( Griffith et al. 2002 ) and that social mating partners frequently invest substantial resources into raising offspring that are unrelated to one or both of them ( Avise et al. 2002 ). While these findings have changed the way we think about reproductive behaviour, most investigations of genetic parentage have been restricted to single populations at a single point in time, obscuring spatial and/or temporal variation in mating behaviour and limiting our ability to determine how environmental changes can lead to shifts in mating strategies. In this issue of Molecular Ecology, Mobley & Jones (2009) compare genetic mating behaviour across five populations of Syngnathus floridae ( Fig. 1 ), a widespread species of pipefish distributed along the Gulf‐ and Atlantic Coasts of North America. The authors document how genetic mating behaviour varies across space in S. floridae and identify correlations between reproductive variation and particular ecological characteristics. Mobley & Jones’ paper is one of an increasing number of studies which address the question of how ecological variables influence mating behaviour, and highlights how our understanding of mating system variation and evolution is likely to expand through the wider application of high‐throughput parentage assessment in a comparative context.

Figure 1 Open in figure viewer PowerPoint A pregnant male dusky pipefish (Syngnathus floridae) in its natural habitat. Photo credit: Joe O’Hop.     

Incipient ring speciation revealed by a migratory divide

Ever since Ernst Mayr (1942) called ring species the ‘perfect demonstration of speciation’, they have attracted much interest from researchers examining how two species evolve from one. In a ring species, two sympatric and reproductively isolated forms are connected by a long chain of intermediate populations that encircle a geographic barrier. Ring species have the potential to demonstrate that speciation can occur without complete geographic isolation, in contrast to the classic model of allopatric speciation. They also allow researchers to examine the causes of reproductive isolation in the contact zone and to use spatial variation to infer the steps by which speciation occurs. According to the classical definition, a ring species must have (i) gradual variation through a chain of populations connecting two divergent and sympatric forms, and (ii) complete or nearly complete reproductive isolation between the terminal forms. But evolutionary biologists now recognize that the process of speciation might often occur with some periods of geographic contact and hybridization between diverging forms; during these phases, even partial reproductive isolation can limit gene flow and permit further divergence to occur. In this issue Bensch et al. (2009) make an exciting and important contribution by extending the ring species concept to a case in which the divergence is much younger and not yet advanced to full reproductive isolation. Their study of geographic variation in willow warblers (Phylloscopus trochilus; Fig. 1 ) provides a beautiful example of gradual variation through a ring of populations connecting two forms that are partially reproductively isolated where they meet, possibly due to divergent migratory behaviours of the terminal forms.

Figure 1 Open in figure viewer PowerPoint A male willow warbler resembling the southeastern‐migrating form (Phylloscopus trochilus acredula), on its breeding territory in central Sweden. (Photo: Anders Hedenström).     

Connecting genetic variation to phenotypic clines

From early allozyme work to recent genome‐wide scans, many studies have reported associations between molecular markers and latitude. These geographic patterns are tantalizing because they hint at the possibility of identifying specific mutations responsible for climatic adaptation. Unfortunately, few studies have done so because these exciting first glances often prove extremely challenging to follow up. Many difficulties can hinder connecting genetic and phenotypic variation in this context, and without such links, distinguishing the action of spatially varying selection from the other evolutionary processes capable of generating these patterns can be quite thorny. Nevertheless, two papers in this issue report excellent progress in overcoming these obstacles and provide persuasive evidence supporting the involvement of specific natural variants in clinal adaptation of Drosophila melanogaster populations ( Fig. 1 ). In the first paper, Paaby et al. (2010) describe replicated allele frequency clines for a coding polymorphism in the Insulin‐like Receptor (InR) gene on two continents, findings that strongly point to selection acting at this locus and that likely reflect life history adaptation. McKechnie et al. (2010) report compelling functional evidence that cis‐regulatory variation in the Dca (drosophila cold acclimation) gene contributes to an adaptive cline in wing size. Notably, these papers employ largely alternative and complementary approaches, and together they exemplify how diverse strategies may be interwoven to draw convincing connections between genotype, phenotype, and evolutionary process.

Figure 1 Open in figure viewer PowerPoint Drosophila melanogaster mating in the field. Credit: Annalise Paaby.     

Determining parentage and relatedness from genetic markers sheds light on patterns of marine larval dispersal

Population connectivity, the extent to which geographically separated subpopulations exchange individuals and are demographically linked, is important to the scientific management of marine living resources. In theory, the design of a marine protected area, for example, depends on an explicit understanding of how dispersal of planktonic larvae affects metapopulation structure and dynamics ( Botsford et al. 2001 ). In practice, for most marine metazoans with planktonic larvae, the mean and variance of the distances that larvae disperse are unobservable quantities, owing to the small sizes of larvae and the very large volumes through which they are distributed. Simulation of dispersal kernels with models that incorporate oceanography and limited aspects of larval biology and behaviour, coupled with field studies of larval distribution, abundance, and settlement, have provided the best available approaches to date for understanding connectivity of marine populations ( Cowen et al. 2006 ). On the other hand, marine population connectivity has often been judged by spatial variation in the frequencies of alleles and genotypes, although the inherent limitations of this indirect approach to measuring larval dispersal have often been overlooked ( Hedgecock et al. 2007 ). More recently, researchers have turned to genetic methods and highly polymorphic markers that can provide direct evidence of population connectivity in the form of parentage or relatedness of recruits (e.g. Jones et al. 2005 ). In this issue, Christie et al. (2010) provide a particularly elegant example, in which both indirect and novel direct genetic methods are used to determine the major ecological processes shaping dispersal patterns of larval bicolour damselfish Stegastes partitus, a common and widespread reef fish species in the Caribbean Basin ( Fig. 1 ).

Figure 1 Open in figure viewer PowerPoint The bicolour damselfish Stegastes partitus shows substantial self‐recruitment of juveniles to their natal coral reef habitat. Below, a male guarding an artificial nest made from PVC pipe; differential reproductive success of parents or differential survival of egg clutches or the larvae that hatch from them may account for signals of sweepstakes reproductive success in this species (photo credits: top, Bill Harward; bottom, Darren Johnson).     

Testing evolutionary hypotheses for phenotypic divergence using landscape genetics

Understanding the evolutionary causes of phenotypic variation among populations has long been a central theme in evolutionary biology. Several factors can influence phenotypic divergence, including geographic isolation, genetic drift, divergent natural or sexual selection, and phenotypic plasticity. But the relative importance of these factors in generating phenotypic divergence in nature is still a tantalizing and unresolved problem in evolutionary biology. The origin and maintenance of phenotypic divergence is also at the root of many ongoing debates in evolutionary biology, such as the extent to which gene flow constrains adaptive divergence ( Garant et al. 2007 ) and the relative importance of genetic drift, natural selection, and sexual selection in initiating reproductive isolation and speciation ( Coyne & Orr 2004 ). In this issue, Wang & Summers (2010) test the causes of one of the most fantastic examples of phenotypic divergence in nature: colour pattern divergence among populations of the strawberry poison frog (Dendrobates pumilio) in Panama and Costa Rica ( Fig. 1 ). This study provides a beautiful example of the use of the emerging field of landscape genetics to differentiate among hypotheses for phenotypic divergence. Using landscape genetic analyses, Wang & Summers were able to reject the hypotheses that colour pattern divergence is due to isolation‐by‐distance (IBD) or landscape resistance. Instead, the hypothesis left standing is that colour divergence is due to divergent selection, in turn driving reproductive isolation among populations with different colour morphs. More generally, this study provides a wonderful example of how the emerging field of landscape genetics, which has primarily been applied to questions in conservation and ecology, now plays an essential role in evolutionary research.

Figure 1 Open in figure viewer PowerPoint Divergent colour morphs observed among populations of the strawberry poison frog, Dendrobates pumilio. Frogs are from San Cristobal (upper left), Cerro Brujo (upper right), Bastimentos (lower right), and Agua (lower left).     

Tobacco mosaic virus, not just a single component virus anymore

Taxonomy: Tobacco mosaic virus (TMV) is the type species of the Tobamovirus genus and a member of the alphavirus-like supergroup. Historically, many tobamoviruses are incorrectly called strains of TMV, although they can differ considerably in sequence similarities and host range from each other and from TMV. Physical properties: TMV virions are 300 × 18 nm rods with a central hollow cavity ( Fig. 1 ) and are composed of 95% capsid protein (CP), and 5% RNA. Each CP subunit interacts with 3-nts in a helical arrangement around the RNA. Virions are stable for decades; infectivity in sap survives heating to 90 °C.

Figure 1 Open in figure viewer PowerPoint Electron micrograph of TMV virions stained with uranyl acetate. Courtesy of Dr J.N. Culver, University of Maryland Biotechnology Institute.     

The nearness of you: the effect of population structure on siring success in a gynodioecious species

Theoretically, both balancing selection and genetic drift can contribute to the maintenance of gender polymorphism within and/or among populations. However, if strong differences exist among genotypes in the quantity of viable gametes they produce, then it is expected that these differences will play an important role in determining the relative frequency of the genotypes and contribute to whether or not such polymorphism is maintained. In this issue, De Cauwer et al. (2010) describe an investigation of gynodioecious wild sea beet, which in addition to containing females, contain two types of hermaphrodites: restored hermaphrodites carrying a cytoplasm that causes pollen sterility and a nuclear gene that restores pollen fertility, and hermaphrodites without the sterilizing cytoplasm. The results show that restored hermaphrodites, who have relatively low pollen viability, achieve disproportionately high siring success simply because of where they are located in a patchy population ( Fig. 1 ). Notably, these individuals tend to be close to females because of the genetics of sex determination. These results indicate that population structure caused by drift processes can have an unexpectedly large effect on the fitness of these low quality hermaphrodites, thereby contributing in the short term to the maintenance of gynodioecy in this population. While these results indicate that population structure caused by drift processes can have a large effect on the relative fitness of genetic variants, whether these effects promote or discourage the maintenance of polymorphism in the long term is still up for debate.

Figure 1 Open in figure viewer PowerPoint A stretch of beach along which wild sea beet can be seen to be growing among the rocks above the splash zone. This linear arrangement enhances the potential for mating success to depend on proximity to other plants (Photo: J.‐F. Arnaud).     

The flickering genes of the last mammoths

Woolly mammoths, Mammuthus primigenius, are arguably the most iconic of the extinct Pleistocene megafauna, and an abundance of large permafrost‐embedded bone and ivory material ( Fig. 1 ) means they were also among the first to yield credible DNA sequences ( Hagelberg et al. 1994 ; Hoss et al. 1994 ). Despite mammoth remains being numerous throughout northern Eurasia and North America, both the earliest and most recent fossils are found in northeast Siberia, with the last known population being confined to Wrangel Island in the Arctic Ocean from around 10,000 years ago until their extinction around 4,000 years ago. The extent to which these Holocene mammoths were descended from the Pleistocene populations of Wrangel Island and the demographic nature of their terminal decline have, until now, remained something of a mystery. In this issue of Molecular Ecology, Nyström et al. (2012) report the first use of autosomal variation to track the decline of the last mammoths and, in doing so, take a significant step towards resolving these questions. The authors genotyped four microsatellite loci in 59 Pleistocene and Holocene mammoths from Wrangel Island and Chukotka in mainland northeastern Siberia and showed that while the Pleistocene‐to‐Holocene transition is associated with a significant reduction in genetic diversity, subsequent levels of variation remain constant until extinction. Such a pattern is somewhat surprising as it indicates that while the last mammoths were confined to only a few Arctic islands, their final extinction on Wrangel Island was not a gradual process resulting from loss of genetic diversity/inbreeding. Instead, it seems they maintained a viable effective population size of around 500 until near their presumably rapid extinction. While the ultimate agent of mammoth extinction remains unknown, the work of Nyström et al. (2012) . suggests that we should be looking for something sudden, like a rapid change in climate/ecology or perhaps the arrival of humans.

Figure 1 Open in figure viewer PowerPoint Sergey Vartanyan during field collection, holding a woolly mammoth tusk found along a river embankment in northeast Siberia (photographer: Diana Solovyeva).