Testing the link between the latitudinal gradient in species richness and rates of molecular evolution

Numerous hypotheses have been proposed to explain latitudinal gradients in species richness, but all are subject to ongoing debate. Here we examine Rohde's (1978, 1992) hypothesis, which proposes that climatic conditions at low latitudes lead to elevated rates of speciation. This hypothesis predicts that rates of molecular evolution should increase towards lower latitudes, but this prediction has never been tested. We discuss potential links between rates of molecular evolution and latitudinal diversity gradients, and present the first test of latitudinal variation in rates of molecular evolution. Using 45 phylogenetically independent, latitudinally separated pairs of bird species and higher taxa, we compare rates of evolution of two mitochondrial genes and DNA-DNA hybridization distances. We find no support for an effect of latitude on rate of molecular evolution. This result casts doubt on the generality of a key component of Rohde's hypothesis linking climate and speciation.

Natural landscapes of the United States / by J. Francis Macbride, Curator of Peruvian Botany.

no.27(1950)

Water scarcity, social power and the production of an elite suburb: the political ecology of water in Matadepera, Catalonia

This article investigates the history of land and water transformations in Matadepera, a wealthy suburb of metropolitan Barcelona. Analysis is informed by theories of political ecology and methods of environmental history; although very relevant, these have received relatively little attention within ecological economics. Empirical material includes communications from the City Archives of Matadepera (1919-1979), 17 interviews with locals born between 1913 and 1958, and an exhaustive review of grey historical literature. Existing water histories of Barcelona and its outskirts portray a battle against natural water scarcity, hard won by heroic engineers and politicians acting for the good of the community. Our research in Matadepera tells a very different story. We reveal the production of a highly uneven landscape and waterscape through fierce political and power struggles. The evolution of Matadepera from a small rural village to an elite suburb was anything but spontaneous or peaceful. It was a socio-environmental project well intended by landowning elites and heavily fought by others. The struggle for the control of water went hand in hand with the land and political struggles that culminated – and were violently resolved - in the Spanish Civil War. The displacement of the economic and environmental costs of water use from few to many continues to this day and is constitutive of Matadepera’s uneven and unsustainable landscape. By unravelling the relations of power that are inscribed in the urbanization of nature (Swyngedouw, 2004), we question the perceived wisdoms of contemporary water policy debates, particularly the notion of a natural scarcity that merits a technical or economic response. We argue that the water question is fundamentally a political question of environmental justice; it is about negotiating alternative visions of the future and deciding whose visions will be produced.

Sex-dependent selection on an autosomal melanic female ornament promotes the evolution of sex ratio bias.

Sex-dependent selection often leads to spectacularly different phenotypes in males and females. In species in which sexual dimorphism is not complete, it is unclear which benefits females and males derive from displaying a trait that is typical of the other sex. In barn owls (Tyto alba), females exhibit on average larger black eumelanic spots than males but members of the two sexes display this trait in the same range of possible values. In a 12-year study, we show that selection exerted on spot size directly or on genetically correlated traits strongly favoured females with large spots and weakly favoured males with small spots. Intense directional selection on females caused an increase in spot diameter in the population over the study period. This increase is due to a change in the autosomal genes underlying the expression of eumelanic spots but not of sex-linked genes. Female-like males produced more daughters than sons, while male-like females produced more sons than daughters when mated to a small-spotted male. These sex ratio biases appear adaptive because sons of male-like females and daughters of female-like males had above-average survival. This demonstrates that selection exerted against individuals displaying a trait that is typical of the other sex promoted the evolution of specific life history strategies that enhance their fitness. This may explain why in many organisms sexual dimorphism is often not complete.

How life history and demography promote or inhibit the evolution of helping behaviours.

In natural populations, dispersal tends to be limited so that individuals are in local competition with their neighbours. As a consequence, most behaviours tend to have a social component, e.g. they can be selfish, spiteful, cooperative or altruistic as usually considered in social evolutionary theory. How social behaviours translate into fitness costs and benefits depends considerably on life-history features, as well as on local demographic and ecological conditions. Over the last four decades, evolutionists have been able to explore many of the consequences of these factors for the evolution of social behaviours. In this paper, we first recall the main theoretical concepts required to understand social evolution. We then discuss how life history, demography and ecology promote or inhibit the evolution of helping behaviours, but the arguments developed for helping can be extended to essentially any social trait. The analysis suggests that, on a theoretical level, it is possible to contrast three critical benefit-to-cost ratios beyond which costly helping is selected for (three quantitative rules for the evolution of altruism). But comparison between theoretical results and empirical data has always been difficult in the literature, partly because of the perennial question of the scale at which relatedness should be measured under localized dispersal. We then provide three answers to this question.

Evolutionary ecology of learning: insights from fruit flies

Ecologically and evolutionarily oriented research on learning has traditionally been carried out on vertebrates and bees. While less sophisticated than those animals, fruit flies (Drosophila) are capable of several forms of learning, and have an advantage of a short generation time, which makes them an ideal system for experimental evolution studies. This review summarizes the insights into evolutionary questions about learning gained in the last decade from evolutionary experiments on Drosophila. These experiments demonstrate that Drosophila have the genetic potential to evolve substantially improved learning performance in ecologically relevant learning tasks. In at least one set of selected populations the improved learning generalized to another task than that used to impose selection, involving a different behavior, different stimuli, and a different sensory channel for the aversive reinforcement. This improvement in learning ability was associated with reduction in other fitness-related traits, such as larval competitive ability and lifespan, pointing out to evolutionary trade-offs of improved learning. These trade-offs were confirmed by other evolutionary experiments where reduction in learning performance was observed as a correlated response to selection for tolerance to larval nutritional stress or for delayed aging. Such trade-offs could be one reason why fruit flies have not fully used up their evolutionary potential for learning ability. Finally, another evolutionary experiment with Drosophila provided the first direct evidence for the long-standing ideas that learning can under some circumstances accelerate and in other slow down genetically-based evolutionary change. These results demonstrate the usefulness of fruit flies as a model system to address evolutionary questions about learning.

Experimental evolution.

Experimental evolution is the study of evolutionary processes occurring in experimental populations in response to conditions imposed by the experimenter. This research approach is increasingly used to study adaptation, estimate evolutionary parameters, and test diverse evolutionary hypotheses. Long applied in vaccine development, experimental evolution also finds new applications in biotechnology. Recent technological developments provide a path towards detailed understanding of the genomic and molecular basis of experimental evolutionary change, while new findings raise new questions that can be addressed with this approach. However, experimental evolution has important limitations, and the interpretation of results is subject to caveats resulting from small population sizes, limited timescales, the simplified nature of laboratory environments, and, in some cases, the potential to misinterpret the selective forces and other processes at work.

Genes as leaders and followers in evolution.

A major question for the study of phenotypic evolution is whether intra- and interspecific diversity originates directly from genetic variation, or instead, as plastic responses to environmental influences initially, followed later by genetic change. In species with discrete alternative phenotypes, evolutionary sequences can be inferred from transitions between environmental and genetic phenotype control, and from losses of phenotypic alternatives. From the available evidence, sequences appear equally probable to start with genetic polymorphism as with polyphenism, with a possible dominance of one or the other for specific trait types. We argue in this review that to evaluate the prevalence of each route, an investigation of both genetic and environmental cues for phenotype determination in several related rather than in isolated species is required.

Developmental and Anatomical Constraints on Vertebrate Genome Evolution

Pendant ma thèse de doctorat, j'ai utilisé des espèces modèles, comme la souris et le poisson-zèbre, pour étudier les facteurs qui affectent l'évolution des gènes et leur expression. Plus précisément, j'ai montré que l'anatomie et le développement sont des facteurs clés à prendre en compte, car ils influencent la vitesse d'évolution de la séquence des gènes, l'impact sur eux de mutations (i.e. la délétion du gène est-elle létale ?), et leur tendance à se dupliquer. Où et quand il est exprimé impose à un gène certaines contraintes ou au contraire lui donne des opportunités d'évoluer. J'ai pu comparer ces tendances aux modèles classiques d'évolution de la morphologie, que l'on pensait auparavant refléter directement les contraintes s'appliquant sur le génome. Nous avons montré que les contraintes entre ces deux niveaux d'organisation ne peuvent pas être transférées simplement : il n'y a pas de lien direct entre la conservation du génotype et celle de phénotypes comme la morphologie. Ce travail a été possible grâce au développement d'outils bioinformatiques. Notamment, j'ai travaillé sur le développement de la base de données Bgee, qui a pour but de comparer l'expression des gènes entre différentes espèces de manière automatique et à large échelle. Cela implique une formalisation de l'anatomie, du développement et de concepts liés à l'homologie grâce à l'utilisation d'ontologies. Une intégration cohérente de données d'expression hétérogènes (puces à ADN, marqueurs de séquence exprimée, hybridations in situ) a aussi été nécessaire. Cette base de données est mise à jour régulièrement et disponible librement. Elle devrait contribuer à étendre les possibilités de comparaison de l'expression des gènes entre espèces pour des études d'évo-devo (évolution du développement) et de génomique. During my PhD, I used model species of vertebrates, such as mouse and zebrafish, to study factors affecting the evolution of genes and their expression. More precisely I have shown that anatomy and development are key factors to take into account, influencing the rate of gene sequence evolution, the impact of mutations (i.e. is the deletion of a gene lethal?), and the propensity of a gene to duplicate. Where and when genes are expressed imposes constraints, or on the contrary leaves them some opportunity to evolve. We analyzed these patterns in relation to classical models of morphological evolution in vertebrates, which were previously thought to directly reflect constraints on the genomes. We showed that the patterns of evolution at these two levels of organization do not translate smoothly: there is no direct link between the conservation of genotype and phenotypes such as morphology. This work was made possible by the development of bioinformatics tools. Notably, I worked on the development of the database Bgee, which aims at comparing gene expression between different species in an automated and large-scale way. This involves the formalization of anatomy, development, and concepts related to homology, through the use of ontologies. A coherent integration of heterogeneous expression data (microarray, expressed sequence tags, in situ hybridizations) is also required. This database is regularly updated and freely available. It should contribute to extend the possibilities for comparison of gene expression between species in evo-devo and genomics studies.

Variation and evolution of herkogamy in Exochaenium (Gentianaceae): implications for the evolution of distyly.

Backgrounds and Aims The spatial separation of stigmas and anthers (herkogamy) in flowering plants functions to reduce self-pollination and avoid interference between pollen dispersal and receipt. Little is known about the evolutionary relationships among the three main forms of herkogamy - approach, reverse and reciprocal herkogamy (distyly) - or about transitions to and from a non-herkogamous condition. This problem was examined in Exochaenium (Gentianaceae), a genus of African herbs that exhibits considerable variation in floral morphology, including the three forms of herkogamy. Methods Using maximum parsimony and maximum likelihood methods, the evolutionary history of herkogamic and non-herkogamic conditions was reconstructed from a molecular phylogeny of 15 species of Exochaenium and four outgroup taxa, based on three chloroplast regions, the nuclear ribosomal internal transcribed spacer (ITS1 and 2) and the 5·8S gene. Ancestral character states were determined and the reconstructions were used to evaluate competing models for the origin of reciprocal herkogamy. Key results Reciprocal herkogamy originated once in Exochaenium from an ancestor with approach herkogamy. Reverse herkogamy and the non-herkogamic condition homostyly were derived from heterostyly. Distylous species possessed pendent, slightly zygomorphic flowers, and the single transition to reverse herkogamy was associated with the hawkmoth pollination syndrome. Reductions in flower size characterized three of four independent transitions from reciprocal herkogamy to homostyly. Conclusions The results support Lloyd and Webb's model in which distyly originated from an ancestor with approach herkogamy. They also demonstrate the lability of sex organ deployment and implicate pollinators, or their absence, as playing an important role in driving transitions among herkogamic and non-herkogamic conditions.

Evolution of life history traits in a natural population of the shrew Crocidura russula

Abstract Life history traits encompass all the decisions concerning fitness an individual is faced with during his life. The study of these traits is crucial to understand the factors shaping the biology of living organisms. Up until now, most of the information on the evolution of life history traits comes from laboratory studies. While these studies are interesting to test the effect of specific parameters, their conclusions are difficult to extrapolate to natural populations. Investigating the evolution of life history traits in natural populations is of great interest. This may be tricky because it requires information on reproduction, survival and morphology of individuals. Mark-recapture methods allow most of this information to be obtained. However, when direct observations of a species are not possible due to its ecology, indirect methods must be used to infer lifetime reproductive success. In this case, molecular markers are particularly helpful in assessing the genetic relationships between individuals and allow the construction of a pedigree. This thesis focuses on a natural population of a small insectivorous mammal, the greater white-toothed shrew, Crocidura russula. Because of its hidden lifestyle, the two complementary techniques mentioned above were combined to gather information on this population. The data were used to explore diverse aspects of evolutionary biology. We demonstrated that the high genetic variance displayed by the species was not maintained by its mating system because this shrew was less monogamous than previously thought. The large genetic diversity was most likely promoted by gene flow from the neighborhood. Dispersal was thus a central topic in this thesis. We showed that dispersal was not driven by inbreeding avoidance. In addition, we did not find any inbreeding depression in the population. Dispersal was promoted by a high number of vacant territories in the population for both sexes, meaning that territory acquisition played an important role in driving dispersal. Moreover, dispersal propensity was shown to have a genetic basis and, once achieved, to have no effect on individual fitness. Body mass was found to be a life history trait strongly influenced by sexual and viability selection in both sexes. Larger individuals had higher access to reproduction through territory acquisition and defense than lighter ones. By contrast, intermediate size individuals were favored by viability selection presumably because of ecological constraints and metabolic costs. Finally, we demonstrated that the majority of the life history traits in our shrew population has the potential to evolve because they maintained substantial amounts of additive genetic variance. Nonetheless, life history traits had no significant heritability due to their high level of nonadditive or environmental variance. Résumé Les traits d'histoire de vie comprennent toutes les décisions auxquelles un individu est confronté au cours de sa vie et qui concernent sa valeur adaptative. L'étude de ces traits est cruciale pour comprendre les facteurs qui façonnent la biologie des êtres vivants. Jusqu'à ce jour, la majorité des informations sur l'évolution des traits d'histoire de vie provient d'études réalisées en laboratoire. Alors que ces études sont intéressantes pour tester l'effet de paramètres spécifiques, leurs conclusions sont difficilement extrapolables aux populations naturelles. Il est particulièrement intéressant d'étudier l'évolution des traits d'histoire de vie dans des populations naturelles. Toutefois, ces études peuvent se révéler difficiles parce qu'elles requièrent des informations sur la reproduction, la survie et la morphologie des individus. Des méthodes de marquage-recapture permettent d'obtenir ces informations. Cependant, lorsque l'écologie de l'espèce rend les obervations directes impossibles, des méthodes indirectes doivent être utilisées pour obtenir le succès reproducteur des individus. Dans ce cas, les marqueurs moléculaires sont particulièrement utiles pour évaluer les relations génétiques entre individus et permettre la construction d'un pedigree. Cette thèse porte sur une population naturelle d'un petit mammifère insectivore, la musaraigne musette, Crocidura russula. Parce que cette espèce présente un mode de vie souterrain, les deux techniques complémentaires mentionnées ci-dessus ont été combinées pour acquérir les informations nécessaires. Les données ont été utilisées pour explorer divers aspects de biologie evolutive. Nous avons montré que la grande quantité de variance génétique trouvée chez cette espèce n'est pas maintenue par son système d'appariement. Celle-ci s'est en effet avérée être moins monogame que ce qui était admis jusqu'ici. Sa grande diversité génétique est plutôt entretenue par le flux de gènes provenant du voisinage. La dispersion a donc été un sujet phare dans cette thèse. Nous avons montré qu'elle n'est pas provoquée par un évitement de la consanguinité et nous n'avons pas trouvé de dépression de consanguité dans notre population. L'acquisition d'un territoire joue par contre un rôle important dans la dispersion. En outre, la dispersion possède une base génétique chez cette espèce. De plus, une fois qu'ils ont dispersé, les individus n'ont pas une valeur adaptative differente d'individus philopatriques. Le poids s'est avéré être un trait d'histoire de vie fortement influencé par la sélection sexuelle et de viabilité chez les deux sexes. Les gros individus ont accès à la reproduction parce qu'ils acquièrent et défendent un territoire plus facilement que les plus légers. Au contraire, les individus de taille intermédiaire sont favorisés par la sélection de viabilité, certainement à cause de contraintes écologiques et de coûts métaboliques. Finalement, nous avons montré que la majorité des traits d'histoire de vie dans notre population a le potentiel d'évoluer parce qu'elle maintient des quantités considérables de variance génétique additive. Néanmoins, l'héritabilité de ces traits d'histoire de vie n'est pas significative à cause de la grande quantité de variance non-additive ou environmentale associée à ces traits.

Latitudinal patterns in plant defense: evolution of cardenolides, their toxicity and induction following herbivory.

Attempts over the past 50 years to explain variation in the abundance, distribution and diversity of plant secondary compounds gave rise to theories of plant defense. Remarkably, few phylogenetically robust tests of these long-standing theories have been conducted. Using >50 species of milkweed (Asclepias spp.), we show that variation among plant species in the induction of toxic cardenolides is explained by latitude, with higher inducibility evolving more frequently at lower latitudes. We also found that: (1) the production of cardenolides showed positive-correlated evolution with the diversity of cardenolides, (2) greater cardenolide investment by a species is accompanied by an increase in an estimate of toxicity (measured as chemical polarity) and (3) instead of trading off, constitutive and induced cardenolides were positively correlated. Analyses of root and shoot cardenolides showed concordant patterns. Thus, milkweed species from lower latitudes are better defended with higher inducibility, greater diversity and added toxicity of cardenolides.

Social evolution and defences against pathogens in ants

Pathogens represent a threat to all organisms, which generates a coevolutionary arms race. Social insects provide an interesting system to study host-pathogen interactions, because their defences depend on both the individual and collective responses, and involve genetic, physiological, behavioral and organizational mechanisms. In this thesis, I studied the evolutionary ecology of the resistance of ant queens and workers to natural fungal pathogens. Mechanisms that increase within-colony genetic diversity, like polyandry and polygyny, decrease relatedness among colony mates, which reduces the strength of selection for the evolution and maintenance of altruistic behavior. A leading hypothesis posits that intracolonial genetic diversity is adaptive because it reduces the risk of pathogen transmission. In chapter 1, I examine individual resistance in ant workers of Formica selysi, a species that shows natural variation in colony queen number. I discuss how this variation might be beneficial to resist natural fungal pathogens in groups. Overall my results indicate that there is genetic variation for fungal resistance in workers, a requirement for the 'genetic diversity for pathogen resistance' hypothesis. However I was not able to detect direct evidence that group diversity improves the survival of focal ants or reduces pathogen transmission. Thus, although the coexistence of multiple queens increases the within-colony variance in worker resistance, it remains unclear whether it protects ant colonies from pathogens and whether it is comparable to polyandry in other social insects. Traditionally, it was thought that the immune system of invertebrates lacked memory and specificity. In chapter 2, I investigate individual immunity in ant queens and show that they may be able to adjust their pathogen defences in response to their current environment by means of immune priming, which bears similarities with the adaptive immunity of vertebrates. However, my results indicate that the expression of immune priming in ant queens may be influenced by factors like mating status, mating conditions or host species. In addition, I showed that mating increases pathogen resistance in çhe two ant species that I studied (F. selysi and Lasius niger). This raises the question of how ant queens invest heavily in both maintenance and reproduction, which I discuss in the context of the evolution of social organization. In chapter 3,1 investigate if transgenerational priming against a fungal pathogen protects the queen progeny. I failed to detect this effect, and discuss why the detection of transgenerational immune priming in ants is a difficult task. Overall, this thesis illustrates some of the individual and collective mechanisms that likely played a role in allowing ants to become one of the most diverse and ecologically successful groups of organisms. -- Les pathogènes représentent une menace pour tous les organismes, ce qui a engendré l'évolution d'une course aux armements. Les insectes sociaux sont un système intéressant permettant d'étudier les interactions hôtes-pathogènes, car leurs défenses dépendent de réponses aussi bien individuelles que collectives, et impliquent des mécanismes génétiques, physiologiques, comportementaux et organisationnels. Dans cette thèse, j'ai étudié l'écologie évolutive de la résistance des reines et des ouvrières de fourmis exposées à des champignons pathogènes. Les facteurs augmentant la diversité génétique à l'intérieur de la colonie, comme la polyandrie et la polygynie, diminuent la parenté, ce qui réduit la pression de sélection pour l'évolution et la maintenance des comportements altruistes. Une hypothèse dominante stipule que la diversité génétique à l'intérieur de la colonie est adaptative car elle réduit le risque de transmission des pathogènes. Dans le chapitre 1, nous examinons la résistance individuelle à des pathogènes fongiques chez les ouvrières de Formica selysi, une espèce présentant une variation naturelle dans le nombre de reines par colonie. Nous discutons aussi de la possibilité que ces variations individuelles augmentent la capacité du groupe à résister à des champignons pathogènes. Dans l'ensemble, nos résultats indiquent une variation génétique dans la résistance aux champignons chez les ouvrières, un prérequis à l'hypothèse que la diversité génétique du groupe augmente la résistance aux pathogènes. Cependant, nous n'avons pas pu détecter une preuve directe que la diversité du groupe augmente la survie de fourmis focales ou réduise la transmission des pathogènes. Ainsi, bien que la coexistence de plusieurs reines augmente la variance dans la résistance des ouvrières à l'intérieur de la colonie, la question de savoir si cela protège les colonies de fourmis contre les pathogènes et si cela est comparable à la polyandrie chez d'autres insectes sociaux reste ouverte. Traditionnellement, il était admis que le système immunitaire des invertébrés ne possédait pas de mémoire et était non-spécifique. Dans le chapitre 2, nous avons étudié l'immunité individuelle chez des reines de fourmis. Nous avons montré que les reines pourraient être capables d'ajuster leurs défenses contre les pathogènes en réponse à leur environnement, grâce à une pré-activation du système immunitaire (« immune priming ») ressemblant à l'immunité adaptative des vertébrés. Cependant, nos résultats indiquent que cette pré-activation du système immunitaire chez les reines dépend du fait d'être accouplée ou non, des conditions d'accouplement, ou de l'espèce. De plus, nous avons montré que l'accouplement augmente la résistance aux pathogènes chez les deux espèces que nous avons étudié (F. selysi et Lasius niger). Ceci pose la question de la capacité des reines à investir fortement aussi bien dans la maintenance que dans la reproduction, ce que nous discutons dans le contexte de l'évolution de l'organisation sociale. Dans le chapitre 3, nous étudions si la pré-activation trans-générationelle du système immunitaire [« trans-generational immune priming ») protège la progéniture de la reine contre un champignon pathogène. Nous n'avons par réussi à détecter cet effet, et discutons des raisons pour lesquelles la détection de la pré-activation trans-générationelle du système immunitaire chez les fourmis est une tâche difficile. Dans l'ensemble, cette thèse illustre quelques-uns des mécanismes individuels et collectifs qui ont probablement contribué à la diversité et à l'important succès écologique des fourmis.

Assessing rapid evolution in a changing environment.

Climate change poses a serious threat to species persistence. Effective modelling of evolutionary responses to rapid climate change is therefore essential. In this review we examine recent advances in phylogenetic comparative methods, techniques normally used to study adaptation over long periods, which allow them to be applied to the study of adaptation over shorter time scales. This increased applicability is largely due to the emergence of more flexible models of character evolution and the parallel development of molecular technologies that can be used to assess adaptive variation at loci scattered across the genome. The merging of phylogenetic and population genetic approaches to the study of adaptation has significant potential to advance our understanding of rapid responses to environmental change.