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.

What's so special about conversion disorder? A problem and a proposal for diagnostic classification.

Conversion disorder presents a problem for the revisions of DSM-IV and ICD-10, for reasons that are informative about the difficulties of psychiatric classification more generally. Giving up criteria based on psychological aetiology may be a painful sacrifice but it is still the right thing to do.

The number of planar central configurations for the 4-body problem is finite when 3 mass positions are fixed

In the n{body problem a central con guration is formed when the position vector of each particle with respect to the center of mass is a common scalar multiple of its acceleration vector. Lindstrom showed for n = 3 and for n > 4 that if n ? 1 masses are located at xed points in the plane, then there are only a nite number of ways to position the remaining nth mass in such a way that they de ne a central con guration. Lindstrom leaves open the case n = 4. In this paper we prove the case n = 4 using as variables the mutual distances between the particles.

Families of periodic orbits for the spatial isosceles 3-body problem

We study the families of periodic orbits of the spatial isosceles 3-body problem (for small enough values of the mass lying on the symmetry axis) coming via the analytic continuation method from periodic orbits of the circular Sitnikov problem. Using the first integral of the angular momentum, we reduce the dimension of the phase space of the problem by two units. Since periodic orbits of the reduced isosceles problem generate invariant two-dimensional tori of the nonreduced problem, the analytic continuation of periodic orbits of the (reduced) circular Sitnikov problem at this level becomes the continuation of invariant two-dimensional tori from the circular Sitnikov problem to the nonreduced isosceles problem, each one filled with periodic or quasi-periodic orbits. These tori are not KAM tori but just isotropic, since we are dealing with a three-degrees-of-freedom system. The continuation of periodic orbits is done in two different ways, the first going directly from the reduced circular Sitnikov problem to the reduced isosceles problem, and the second one using two steps: first we continue the periodic orbits from the reduced circular Sitnikov problem to the reduced elliptic Sitnikov problem, and then we continue those periodic orbits of the reduced elliptic Sitnikov problem to the reduced isosceles problem. The continuation in one or two steps produces different results. This work is merely analytic and uses the variational equations in order to apply Poincar´e’s continuation method.

Infinitely many periodic orbits for the rhomboidal five-body problem

We prove the existence of infinitely many symmetric periodic orbits for a regularized rhomboidal five-body problem with four small masses placed at the vertices of a rhombus centered in the fifth mass. The main tool for proving the existence of such periodic orbits is the analytic continuation method of Poincaré together with the symmetries of the problem. © 2006 American Institute of Physics.

El proyecto ‘Ecosportech’ como ejemplo de universidad emprendedora. Una experiencia a base del ‘Problem Based Learning’

El objetivo de este artículo es presentar el proyecto EcoSPORTech, cuya finalidad es la creación de una empresa social con jóvenes para la realización de actividades deportivas/ocio en el medio natural, integrando las nuevas tecnologías. Este proyecto supone una colaboración interdisciplinaria dentro de la Universidad de Vic, entre las facultades de Empresa y Comunicación (FEC), la de Ciencias de la Salud y el Bienestar (FCSB) y la de Educación (FE) e integra un equipo de profesionales procedentes de los ámbitos de la empresa, el marketing, el periodismo, el deporte y la terapia ocupacional. Estos profesores formarán al grupo de jóvenes con los que se creará la empresa y dirigirán la misma. Esta empresa (cooperativa) se integra en el vivero de empresas sociales que se está creando en la Universidad de Vic.

Double-Antiprism Central Configurations of the 3n-Body Problem

Abstract In this paper we study numerically a new type of central configurations of the 3n-body problem with equal masses which consist of three n-gons contained in three planes z = 0 and z = ±β = 0. The n-gon on z = 0 is scaled by a factor α and it is rotated by an angle of π/n with respect to the ones on z = ±β. In this kind of configurations, the masses on the planes z = 0 and z = β are at the vertices of an antiprism with bases of different size. The same occurs with the masses on z = 0 and z = −β. We call this kind of central configurations double-antiprism central configurations. We will show the existence of central configurations of this type.

On the existence of bi-pyramidal central configurations of the n + 2-body problem with an n-gon base

Abstract. In this paper we prove the existence of central con gurations of the n + 2{body problem where n equal masses are located at the vertices of a regular n{gon and the remaining 2 masses, which are not necessarily equal, are located on the straight line orthogonal to the plane containing the n{gon passing through its center. Here this kind of central con gurations is called bi{pyramidal central con gurations. In particular, we prove that if the masses mn+1 and mn+2 and their positions satisfy convenient relations, then the con guration is central. We give explicitly those relations.

Central configurations of nested regular polyhedra for the spatial 2n–body problem

We consider 2n masses located at the vertices of two nested regular polyhedra with the same number of vertices. Assuming that the masses in each polyhedron are equal, we prove that for each ratio of the masses of the inner and the outer polyhedron there exists a unique ratio of the length of the edges of the inner and the outer polyhedron such that the configuration is central.

Central configurations of three nested regular polyhedra for the spatial 3n–body problem

Three regular polyhedra are called nested if they have the same number of vertices n, the same center and the positions of the vertices of the inner polyhedron ri, the ones of the medium polyhedron Ri and the ones of the outer polyhedron Ri satisfy the relation Ri = ri and Ri = Rri for some scale factors R > > 1 and for all i = 1, . . . , n. We consider 3n masses located at the vertices of three nested regular polyhedra. We assume that the masses of the inner polyhedron are equal to m1, the masses of the medium one are equal to m2, and the masses of the outer one are equal to m3. We prove that if the ratios of the masses m2/m1 and m3/m1 and the scale factors and R satisfy two convenient relations, then this configuration is central for the 3n–body problem. Moreover there is some numerical evidence that, first, fixed two values of the ratios m2/m1 and m3/m1, the 3n–body problem has a unique central configuration of this type; and second that the number of nested regular polyhedra with the same number of vertices forming a central configuration for convenient masses and sizes is arbitrary.