The evolution of social discounting in hierarchically clustered populations.

The expression of a social behaviour may affect the fitness of actors and recipients living in the present and in the future of the population. When there is a risk that a future reward will not be experienced in such a context, the value of that reward should be discounted; but by how much? Here, we evaluate social discount rates for delayed fitness rewards to group of recipients living at different positions in both space and time than the actor in a hierarchically clustered population. This is a population where individuals are grouped into families, families into villages, villages into clans, and so on, possibly ad infinitum. The group-wide fitness effects are assumed to either increase or decrease the fecundity or the survival of recipients and can be arbitrarily extended in space and time. We find that actions changing the survival of individuals living in the future are generally more strongly discounted than fecundity-changing actions for all future times and that the value of future rewards increases as individuals live longer. We also find that delayed fitness effects may not only be discounted by a constant factor per unit delay (exponential discounting), but that, as soon as there is localized dispersal in a population, discounting per unit delay is likely to fall rapidly for small delays and then slowly for longer delays (hyperbolic discounting). As dispersal tends to be localized in natural populations, our results suggest that evolution is likely to favour individuals that express present-biased behaviours and that may be time-inconsistent with respect to their group-wide effects.

The evolution of unicoloniality in ant societies : recognition, dispersal and population genetic structure

Introduction Societies of ants, bees, wasps and termites dominate many terrestrial ecosystems (Wilson 1971). Their evolutionary and ecological success is based upon the regulation of internal conflicts (e.g. Ratnieks et al. 2006), control of diseases (e.g. Schmid-Hempel 1998) and individual skills and collective intelligence in resource acquisition, nest building and defence (e.g. Camazine 2001). Individuals in social species can pass on their genes not only directly trough their own offspring, but also indirectly by favouring the reproduction of relatives. The inclusive fitness theory of Hamilton (1963; 1964) provides a powerful explanation for the evolution of reproductive altruism and cooperation in groups with related individuals. The same theory also led to the realization that insect societies are subject to internal conflicts over reproduction. Relatedness of less-than-one is not sufficient to eliminate all incentive for individual selfishness. This would indeed require a relatedness of one, as found among cells of an organism (Hardin 1968; Keller 1999). The challenge for evolutionary biology is to understand how groups can prevent or reduce the selfish exploitation of resources by group members, and how societies with low relatedness are maintained. In social insects the evolutionary shift from single- to multiple queens colonies modified the relatedness structure, the dispersal, and the mode of colony founding (e.g. (Crozier & Pamilo 1996). In ants, the most common, and presumably ancestral mode of reproduction is the emission of winged males and females, which found a new colony independently after mating and dispersal flights (Hölldobler & Wilson 1990). The alternative reproductive tactic for ant queens in multiple-queen colonies (polygyne) is to seek to be re-accepted in their natal colonies, where they may remain as additional reproductives or subsequently disperse on foot with part of the colony (budding) (Bourke & Franks 1995; Crozier & Pamilo 1996; Hölldobler & Wilson 1990). Such ant colonies can contain up to several hundred reproductive queens with an even more numerous workforce (Cherix 1980; Cherix 1983). As a consequence in polygynous ants the relatedness among nestmates is very low, and workers raise brood of queens to which they are only distantly related (Crozier & Pamilo 1996; Queller & Strassmann 1998). Therefore workers could increase their inclusive fitness by preferentially caring for their closest relatives and discriminate against less related or foreign individuals (Keller 1997; Queller & Strassmann 2002; Tarpy et al. 2004). However, the bulk of the evidence suggests that social insects do not behave nepotistically, probably because of the costs entailed by decreased colony efficiency or discrimination errors (Keller 1997). Recently, the consensus that nepotistic behaviour does not occur in insect colonies was challenged by a study in the ant Formica fusca (Hannonen & Sundström 2003b) showing that the reproductive share of queens more closely related to workers increases during brood development. However, this pattern can be explained either by nepotism with workers preferentially rearing the brood of more closely related queens or intrinsic differences in the viability of eggs laid by queens. In the first chapter, we designed an experiment to disentangle nepotism and differences in brood viability. We tested if workers prefer to rear their kin when given the choice between highly related and unrelated brood in the ant F. exsecta. We also looked for differences in egg viability among queens and simulated if such differences in egg viability may mistakenly lead to the conclusion that workers behave nepotistically. The acceptance of queens in polygnous ants raises the question whether the varying degree of relatedness affects their share in reproduction. In such colonies workers should favour nestmate queens over foreign queens. Numerous studies have investigated reproductive skew and partitioning of reproduction among queens (Bourke et al. 1997; Fournier et al. 2004; Fournier & Keller 2001; Hammond et al. 2006; Hannonen & Sundström 2003a; Heinze et al. 2001; Kümmerli & Keller 2007; Langer et al. 2004; Pamilo & Seppä 1994; Ross 1988; Ross 1993; Rüppell et al. 2002), yet almost no information is available on whether differences among queens in their relatedness to other colony members affects their share in reproduction. Such data are necessary to compare the relative reproductive success of dispersing and non-dispersing individuals. Moreover, information on whether there is a difference in reproductive success between resident and dispersing queens is also important for our understanding of the genetic structure of ant colonies and the dynamics of within group conflicts. In chapter two, we created single-queen colonies and then introduced a foreign queens originating from another colony kept under similar conditions in order to estimate the rate of queen acceptance into foreign established colonies, and to quantify the reproductive share of resident and introduced queens. An increasing number of studies have investigated the discrimination ability between ant workers (e.g. Holzer et al. 2006; Pedersen et al. 2006), but few have addressed the recognition and discrimination behaviour of workers towards reproductive individuals entering colonies (Bennett 1988; Brown et al. 2003; Evans 1996; Fortelius et al. 1993; Kikuchi et al. 2007; Rosengren & Pamilo 1986; Stuart et al. 1993; Sundström 1997; Vásquez & Silverman in press). These studies are important, because accepting new queens will generally have a large impact on colony kin structure and inclusive fitness of workers (Heinze & Keller 2000). In chapter three, we examined whether resident workers reject young foreign queens that enter into their nest. We introduced mated queens into their natal nest, a foreign-female producing nest, or a foreign male-producing nest and measured their survival. In addition, we also introduced young virgin and mated queens into their natal nest to examine whether the mating status of the queens influences their survival and acceptance by workers. On top of polgyny, some ant species have evolved an extraordinary social organization called 'unicoloniality' (Hölldobler & Wilson 1977; Pedersen et al. 2006). In unicolonial ants, intercolony borders are absent and workers and queens mix among the physically separated nests, such that nests form one large supercolony. Super-colonies can become very large, so that direct cooperative interactions are impossible between individuals of distant nests. Unicoloniality is an evolutionary paradox and a potential problem for kin selection theory because the mixing of queens and workers between nests leads to extremely low relatedness among nestmates (Bourke & Franks 1995; Crozier & Pamilo 1996; Keller 1995). A better understanding of the evolution and maintenance of unicoloniality requests detailed information on the discrimination behavior, dispersal, population structure, and the scale of competition. Cryptic genetic population structure may provide important information on the relevant scale to be considered when measuring relatedness and the role of kin selection. Theoretical studies have shown that relatedness should be measured at the level of the `economic neighborhood', which is the scale at which intraspecific competition generally takes place (Griffin & West 2002; Kelly 1994; Queller 1994; Taylor 1992). In chapter four, we conducted alarge-scale study to determine whether the unicolonial ant Formica paralugubris forms populations that are organised in discrete supercolonies or whether there is a continuous gradation in the level of aggression that may correlate with genetic isolation by distance and/or spatial distance between nests. In chapter five, we investigated the fine-scale population structure in three populations of F. paralugubris. We have developed mitochondria) markers, which together with the nuclear markers allowed us to detect cryptic genetic clusters of nests, to obtain more precise information on the genetic differentiation within populations, and to separate male and female gene flow. These new data provide important information on the scale to be considered when measuring relatedness in native unicolonial populations.

The co-evolution of culturally inherited altruistic helping and cultural transmission under random group formation.

Limited migration results in kin selective pressure on helping behaviors under a wide range of ecological, demographic and life-history situations. However, such genetically determined altruistic helping can evolve only when migration is not too strong and group size is not too large. Cultural inheritance of helping behaviors may allow altruistic helping to evolve in groups of larger size because cultural transmission has the potential to markedly decrease the variance within groups and augment the variance between groups. Here, we study the co-evolution of culturally inherited altruistic helping behaviors and two alternative cultural transmission rules for such behaviors. We find that conformist transmission, where individuals within groups tend to copy prevalent cultural variants (e.g., beliefs or values), has a strong adverse effect on the evolution of culturally inherited helping traits. This finding is at variance with the commonly held view that conformist transmission is a crucial factor favoring the evolution of altruistic helping in humans. By contrast, we find that under one-to-many transmission, where individuals within groups tend to copy a "leader" (or teacher), altruistic helping can evolve in groups of any size, although the cultural transmission rule itself hitchhikes rather weakly with a selected helping trait. Our results suggest that culturally determined helping behaviors are more likely to be driven by "leaders" than by popularity, but the emergence and stability of the cultural transmission rules themselves should be driven by some extrinsic factors.

The evolution, maintenance and adaptive function of genetic colour polymorphism in birds.

The hypothesis that ornaments can honestly signal quality only if their expression is condition-dependent has dominated the study of the evolution and function of colour traits. Much less interest has been devoted to the adaptive function of colour traits for which the expression is not, or is to a low extent, sensitive to body condition and the environment in which individuals live. The aim of the present paper is to review the current theoretical and empirical knowledge of the evolution, maintenance and adaptive function of colour plumage traits for which the expression is mainly under genetic control. The finding that in many bird species the inheritance of colour morphs follows the laws of Mendel indicates that genetic colour polymorphism is frequent. Polymorphism may have evolved or be maintained because each colour morph facilitates the exploitation of alternative ecological niches as suggested by the observation that individuals are not randomly distributed among habitats with respect to coloration. Consistent with the hypothesis that different colour morphs are linked to alternative strategies is the finding that in a majority of species polymorphism is associated with reproductive parameters, and behavioural, life-history and physiological traits. Experimental studies showed that such covariations can have a genetic basis. These observations suggest that colour polymorphism has an adaptive function. Aviary and field experiments demonstrated that colour polymorphism is used as a criterion in mate-choice decisions and dominance interactions confirming the claim that conspecifics assess each other's colour morphs. The factors favouring the evolution and maintenance of genetic variation in coloration are reviewed, but empirical data are virtually lacking to assess their importance. Although current theory predicts that only condition-dependent traits can signal quality, the present review shows that genetically inherited morphs can reveal the same qualities. The study of genetic colour polymorphism will provide important and original insights on the adaptive function of conspicuous traits.

Inbreeding load, bet hedging, and the evolution of sex-biased dispersal

Inbreeding load affects not only the average fecundity of philopatric individuals but also its variance. From bet-hedging theory, this should add further dispersal pressures to those stemming from the mere avoidance of inbreeding. Pressures on both sexes are identical under monogamy or promiscuity. Under polygyny, by contrast, the variance in reproductive output decreases with dispersal rate in females but increases in males, which should induce a female-biased dispersal. To test this prediction, we performed individual-based simulations. From our results, a female-biased dispersal indeed emerges as both polygyny and inbreeding load increase. We conclude that sex-biased dispersal may be selected for as a bet-hedging strategy.

Welfare regimes and change in the employment structure: Britain, Denmark and Germany since 1990

Welfare states are often reduced to their role as providers of social protection and redistribution. In 1990, Esping-Andersen argued that they also affect employment creation and the class structure. We analyse the stratification outcomes for three welfare regimes - Britain, Germany and Denmark - over the 1990s and 2000s. Based on individual-level surveys, we observe a disproportionate increase among professionals and managers, and a decline among production workers and clerks. The result is clear-cut occupational upgrading in Denmark and Germany. In Britain, high and low-end service jobs expanded, resulting in a polarized version of upgrading. Growth in low-end service jobs - and thus polarization - is no precondition for full employment. Both Britain and Denmark halved their low-educated unemployment rate between 1995 and 2008. Yet low-end service jobs expanded only in Britain, not in Denmark. The cause is the evolution of labour supply: rising educational attainment means that fewer low-educated workers look for low-skilled jobs.

Geochemical evolution of active porphyry copper tailings impoundments : from alkaline deposition towards acidification

J. Smuda: Geochemical evolution of active porphyry copper tailings impoundments Thesis abstract Mine waste is the largest volume of materials handled in the world. The oxidation of sulfidic mine waste may result in the release of acid mine drainage (AMD) rich in heavy metals and arsenic to the environment, one of the major problems the mining industry is facing today. To control and reduce this environmental impact, it is crucial to identify the main geochemical and hydrological processes influencing contaminant liberation, transport, and retention. This thesis presents the results of a geochemical, mineralogical and stable isotope study (δ2H, δ18O, δ34S) from two active porphyry copper tailings impoundments in Mediterranean (Carén tailings impoundment, El Teniente mine, Central Chile) and hyper-arid climate (Talabre tailings impoundment, Chuquicamata, Northern Chile) from the deposition in alkaline environment (pH 10.5) towards acidification after several years of exposure. The major hydrological results were the identification of vertical contaminant and water transport in the uppermost, not water-saturated zone, triggered by capillary rise due to evaporation, and infiltration downwards due to new tailings deposition, and of horizontal transport in the groundwater zone. At the surface of the sedimented tailings, evaporation of pore water led to the precipitation of Na-Ca-Mg sulfates (e.g., gypsum, tenorite), in hyper-arid climate also halite. At the Carén tailings impoundment, renewed deposition in a 4-week interval inhibited a pH decrease below neutral values and the formation of an efflorescent salt crust. At the Talabre tailings impoundment, deposition breaks of several years resulted in the formation of acidic oxidation zones in the timeframe of less than 4 years. This process enabled the transport of liberated Cu, Zn, and Fe via capillary rise to the surface, where these metals precipitated as heavy-metal sulfates (e.g., devilline, krohnkite) and chlorides (eriochalcite, atacamite). Renewed depositing may dissolve efflorescent salts and transport liberated elements towards the groundwater zone. This zone was found to be highly dynamic due to infiltration and mixing with water from different sources, like groundwater, catchment water, and infiltration from superficial waters. There, Cu was found to be partially mobile due to complexation with Cl (in Cl-rich groundwater, Talabre) and dissolved organic matter (in zones with infiltration of catchment water rich in dissolved organic matter, Carén). A laboratory study on the isotopic fractionation of sulfur and oxygen of sulfate in different minerals groups (water-soluble sulfates, low- and high-crystalline Fe(III) oxyhydroxides) contributed to the use of stable isotopes as tracer of geochemical and transport processes for environmental studies. The results highlight that a detailed geochemical, stable isotope and mineralogical study permits the identification of contamination processes and pathways already during the deposition of mine tailings. This knowledge allows the early planning of adequate actions to reduce and control the environmental impact during tailings deposition and after the closing of the impoundment. J. Smuda: Geochemical evolution of active porphyry copper tailings impoundments Résumé de these Les déchets miniers constituent les plus grands volumes de matériel gérés dans le monde. L'oxydation des déchets miniers sulfuriques peut conduire à la libération de drainages miniers acides (DMA) riches en métaux et arsenic dans l'environnement, ce qui est l'un des principaux problèmes de l'industrie minière aujourd'hui. Pour contrôler et réduire ces impacts sur l'environnement, il est crucial d'identifier les principaux processus géochimiques et hydrologiques influençant la libération, le transport et la rétention des contaminants. Cette thèse présente les résultats d'une étude géochimique, minéralogique et des isotopes stables (δ2H, δ18O, δ34S) sur des déchets miniers de 2 sites de dépôt actifs en climat méditerranéen (Dépôt de déchets de Carén, mine de El Teniente, Centre du Chili) et en climat hyper-aride (Dépôt de déchets de Talabre, mine de Chuquicamata, Nord du Chili). L'objectif était d'étudier l'évolution des déchets de la déposition en milieu alcalin (pH = 10.5) vers l'acidification après plusieurs années d'exposition. Le principal résultat hydrologique a été l'identification de 2 types de transport : un transport vertical de l'eau et des contaminants dans la zone non saturée en surface, induit par la montée capillaire due à l'évaporation et par l'infiltration subséquente de la déposition de sédiments frais ; et un transport horizontal dans la zone des eaux souterraines. À la surface des déchets, l'évaporation de l'eau interstitielle conduit à la précipitation de sulfates de Na-Ca-Mg (ex. gypse, ténorite) et halite en climat hyper-aride. Dans le site de Carén, une nouvelle déposition de déchets frais à 4 semaines intervalle a empêché la baise du pH en deçà des valeurs neutres et la formation d'une croûte de sels efflorescentes en surface. Dans le site de Talabre, les fentes de dessiccation des dépôts ont entraîné la formation d'une zone d'oxydation à pH acide en moins de 4 ans. Ce processus a permis la libération et le transport par capillarité de Cu, Zn, Fe vers la surface, où ces éléments précipitent sous forme de sulfates de métaux lourds (ex., dévilline, krohnkite) de chlorures (ex. ériochalcite, atacamite). Une nouvelle déposition de sédiments frais pourrait dissoudre ces sels et les transporter vers la zone des eaux souterraines. Cette dernière zone était très dynamique en raison du mélange d'eaux provenant de différentes sources, comme les eaux souterraines, l'eau de captage et l'infiltration des eaux superficielles. Egalement dans cette zone, le cuivre était partiellement mobile à cause de la formation de complexe avec le chlore (dans les zone riche en Cl, Talabre) et avec la matière organique dissoute (dans les zones où s'infiltre l'eau de captage riche en matière organique, Carén). Une étude en laboratoire sur le fractionnement des isotopes stables de sulfure et d'oxygène des sulfates dans différents groupes de minéraux (sulfates hydrosolubles, sulfures de oxy-hydroxyde de Fe(III) faiblement ou fortement cristallins) a permis d'apporter une contribution à leur utilisation comme traceurs dans l'étude des processus géochimiques et de transport lors d'études environnementales. Les résultats montrent qu'une étude détaillée de la géochimie, des isotopes stables et de la minéralogie permet d'identifier les processus et les voies de contamination déjà pendant la période de dépôt des déchets miniers. Cette connaissance permet de planifier, dès le début de l'exploitation, des mesures adéquates pour réduire et contrôler l'impact sur l'environnement pendant la période de dépôts de déchets miniers et après la fermeture du site.

Is our heart a well-designed pump? The heart along animal evolution.

A carrier system for gases and nutrients became mandatory when primitive animals grew larger and developed different organs. The first circulatory systems are peristaltic tubes pushing slowly the haemolymph into an open vascular tree without capillaries (worms). Arthropods developed contractile bulges on the abdominal aorta assisted by accessory hearts for wings or legs and by abdominal respiratory motions. Two-chamber heart (atrium and ventricle) appeared among mollusks. Vertebrates have a multi-chamber heart and a closed circulation with capillaries. Their heart has two chambers in fishes, three chambers (two atria and one ventricle) in amphibians and reptiles, and four chambers in birds and mammals. The ventricle of reptiles is partially divided in two cavities by an interventricular septum, leaving only a communication of variable size leading to a variable shunt. Blood pressure increases progressively from 15 mmHg (worms) to 170/70 mmHg (birds) according to the increase in metabolic rate. When systemic pressure exceeds 50 mmHg, a lower pressure system appears for the circulation through gills or lungs in order to improve gas exchange. A four-chamber heart allows a complete separation of systemic and pulmonary circuits. This review describes the circulatory pumping systems used in the different classes of animals, their advantages and failures, and the way they have been modified with evolution.

A new classification of the Turkish terranes and sutures and its implication for the paleotectonic history of the region

The Turkish part of the Tethyan realm is represented by a series of terranes juxtaposed through Alpine convergent movements and separated by complex suture zones. Different terranes can be defined and characterized by their dominant geological background. The Pontides domain represents a segment of the former active margin of Eurasia, where back-arc basins opened in the Triassic and separated the Sakarya terrane from neighbouring regions. Sakarya was re-accreted to Laurasia through the Balkanic mid-Cretaceous orogenic event that also affected the Rhodope and Strandja zones. The whole region from the Balkans to the Caucasus was then affected by a reversal of subduction and creation of a Late Cretaceous arc before collision with the Anatolian domain in the Eocene. If the Anatolian terrane underwent an evolution similar to Sakarya during the Late Paleozoic and Early Triassic times, both terranes had a diverging history during and after the Eo-Cimmerian collision. North of Sakarya, the Küre back-arc was closed during the Jurassic, whereas north of the Anatolian domain, the back-arc type oceans did not close before the Late Cretaceous. During the Cretaceous, both domains were affected by ophiolite obduction, but in very different ways: north directed diachronous Middle to Late Cretaceous mélange obduction on the Jurassic Sakarya passive margin; Senonian synchronous southward obduction on the Triassic passive margin of Anatolia. From this, it appears that the Izmir-Ankara suture, currently separating both terranes, is composite, and that the passive margin of Sakarya is not the conjugate margin of Anatolia. To the south, the Cimmerian Taurus domain together with the Beydağları domain (part of the larger Greater Apulian terrane), were detached from north Gondwana in the Permian during the opening of the Neotethys (East-Mediterranean basin). The drifting Cimmerian blocks entered into a soft collision with the Anatolian and related terranes in the Eo-Cimmerian orogenic phase (Late Triassic), thus suturing the Paleotethys. At that time, the Taurus plate developed foreland-type basins, filled with flysch-molasse deposits that locally overstepped the lower plate Taurus terrane and were deposited in the opening Neotethys to the south. These olistostromal deposits are characterized by pelagic Carboniferous and Permian material from the Paleotethys suture zone found in the Mersin mélange. The latter, as well as the Antalya and Mamonia domains are represented by a series of exotic units now found south of the main Taurus range. Part of the Mersin exotic material was clearly derived from the former north Anatolian passive margin (Huğlu-type series) and re-displaced during the Paleogene. This led us to propose a plate tectonic model where the Anatolian ophiolitic front is linked up with the Samail/Baër-Bassit obduction front found along the Arabian margin. The obduction front was indented by the Anatolian promontory whose eastern end was partially subducted. Continued slab roll-back of the Neotethys allowed Anatolian exotics to continue their course southwestward until their emplacement along the Taurus southern margin (Mersin) and up to the Beydağları promontory (Antaya-Mamonia) in the latest Cretaceous-Paleocene. The supra-subduction ocean opening at the back of the obduction front (Troodos-type Ocean) was finally closed by Eocene north-south shortening between Africa and Eurasia. This brought close to each other Cretaceous ophiolites derived from the north of Anatolia and those obducted on the Arabian promontory. The latter were sealed by a Maastrichtian platform, and locally never affected by Alpine tectonism, whereas those located on the eastern Anatolian plate are strongly deformed and metamorphosed, and affected by Eocene arc magmatism. These observations help to reconstruct the larger frame of the central Tethyan realm geodynamic evolution.