Ulf Dieckmann

On the origin of species by sympatric speciation

Understanding speciation is a fundamental biological problem. It is believed that many species originated through allopatric divergence, where new species arise from geographically isolated populations of the same ancestral species. In contrast, the possibility of sympatric speciation (in which new species arise without geographical isolation) has often been dismissed, partly because of theoretical difficulties,. Most previous models analysing sympatric speciation concentrated on particular aspects of the problem while neglecting others. Here we present a model that integrates a novel combination of different features and show that sympatric speciation is a likely outcome of competition for resources. We use multilocus genetics to describe sexual reproduction in an individual-based model, and we consider the evolution of assortative mating (where individuals mate preferentially with like individuals) depending either on an ecological character affecting resource use or on a selectively neutral marker trait. In both cases, evolution of assortative mating often leads to reproductive isolation between ecologically diverging subpopulations. When assortative mating depends on a marker trait, and is therefore not directly linked to resource competition, speciation occurs when genetic drift breaks the linkage equilibrium between the marker and the ecological trait. Our theory conforms well with mounting empirical evidence for the sympatric origin of many species.

Hybridization and speciation

Hybridization has many and varied impacts on the process of speciation. Hybridization may slow or reverse differentiation by allowing gene flow and recombination. It may accelerate speciation via adaptive introgression or cause near‐instantaneous speciation by allopolyploidization. It may have multiple effects at different stages and in different spatial contexts within a single speciation event. We offer a perspective on the context and evolutionary significance of hybridization during speciation, highlighting issues of current interest and debate. In secondary contact zones, it is uncertain if barriers to gene flow will be strengthened or broken down due to recombination and gene flow. Theory and empirical evidence suggest the latter is more likely, except within and around strongly selected genomic regions. Hybridization may contribute to speciation through the formation of new hybrid taxa, whereas introgression of a few loci may promote adaptive divergence and so facilitate speciation. Gene regulatory networks, epigenetic effects and the evolution of selfish genetic material in the genome suggest that the Dobzhansky–Muller model of hybrid incompatibilities requires a broader interpretation. Finally, although the incidence of reinforcement remains uncertain, this and other interactions in areas of sympatry may have knock‐on effects on speciation both within and outside regions of hybridization.

The dynamical theory of coevolution: A derivation from stochastic ecological processes

In this paper we develop a dynamical theory of coevolution in ecological communities. The derivation explicitly accounts for the stochastic components of evolutionary change and is based on ecological processes at the level of the individual. We show that the coevolutionary dynamic can be envisaged as a directed random walk in the communitys trait space. A quantitative description of this stochastic process in terms of a master equation is derived. By determining the first jump moment of this process we abstract the dynamic of the mean evolutionary path. To first order the resulting equation coincides with a dynamic that has frequently been assumed in evolutionary game theory. Apart from recovering this canonical equation we systematically establish the underlying assumptions. We provide higher order corrections and show that these can give rise to new, unexpected evolutionary effects including shifting evolutionary isoclines and evolutionary slowing down of mean paths as they approach evolutionary equilibria. Extensions of the derivation to more general ecological settings are discussed. In particular we allow for multi-trait coevolution and analyze coevolution under nonequilibrium population dynamics.

Maturation trends indicative of rapid evolution preceded the collapse of northern cod

Northern cod, comprising populations of Atlantic cod (Gadus morhua) off southern Labrador and eastern Newfoundland, supported major fisheries for hundreds of years. But in the late 1980s and early 1990s, northern cod underwent one of the worst collapses in the history of fisheries. The Canadian government closed the directed fishing for northern cod in July 1992, but even after a decade-long offshore moratorium, population sizes remain historically low. Here we show that, up until the moratorium, the life history of northern cod continually shifted towards maturation at earlier ages and smaller sizes. Because confounding effects of mortality changes and growth-mediated phenotypic plasticity are accounted for in our analyses, this finding strongly suggests fisheries-induced evolution of maturation patterns in the direction predicted by theory. We propose that fisheries managers could use the method described here as a tool to provide warning signals about changes in life history before more overt evidence of population decline becomes manifest.

Speciation along environmental gradients

Traditional discussions of speciation are based on geographical patterns of species ranges. In allopatric speciation, long-term geographical isolation generates reproductively isolated and spatially segregated descendant species. In the absence of geographical barriers, diversification is hindered by gene flow. Yet a growing body of phylogenetic and experimental data suggests that closely related species often occur in sympatry or have adjacent ranges in regions over which environmental changes are gradual and do not prevent gene flow. Theory has identified a variety of evolutionary processes that can result in speciation under sympatric conditions, with some recent advances concentrating on the phenomenon of evolutionary branching. Here we establish a link between geographical patterns and ecological processes of speciation by studying evolutionary branching in spatially structured populations. We show that along an environmental gradient, evolutionary branching can occur much more easily than in non-spatial models. This facilitation is most pronounced for gradients of intermediate slope. Moreover, spatial evolutionary branching readily generates patterns of spatial segregation and abutment between the emerging species. Our results highlight the importance of local processes of adaptive divergence for geographical patterns of speciation, and caution against pitfalls of inferring past speciation processes from present biogeographical patterns.

The Geometry of Ecological Interactions: Simplifying Spatial Complexity

The field of spatial ecology has expanded dramatically in the past few years. This volume, written by world experts in the field, gives detailed coverage of the main areas of development in spatial ecological theory. Integrating a perspective from field ecology with novel methods for simplifying spatial complexity, it offers a didactical treatment with a gradual increase in mathematical sophistication. In addition, the volume features introductions to those fundamental phenomena in spatial ecology where emerging spatial patterns influence ecological outcomes qualitatively as well as quantitatively. An appreciation and understanding of such systematic departures from standard, nonspatial models is required if ecological theory is to move on in the 21st century. Written for graduate students and researchers in theoretical, evolutionary, and spatial ecology, applied mathematics, and spatial statistics, this book is a ground-breaking treatment of modern spatial ecological theory.

Evolutionary Branching and Sympatric Speciation Caused by Different Types of Ecological Interactions

Evolutionary branching occurs when frequency‐dependent selection splits a phenotypically monomorphic population into two distinct phenotypic clusters. A prerequisite for evolutionary branching is that directional selection drives the population toward a fitness minimum in phenotype space. This article demonstrates that selection regimes leading to evolutionary branching readily arise from a wide variety of different ecological interactions within and between species. We use classical ecological models for symmetric and asymmetric competition, for mutualism, and for predator‐prey interactions to describe evolving populations with continuously varying characters. For these models, we investigate the ecological and evolutionary conditions that allow for evolutionary branching and establish that branching is a generic and robust phenomenon. Evolutionary branching becomes a model for sympatric speciation when population genetics and mating mechanisms are incorporated into ecological models. In sexual populations with random mating, the continual production of intermediate phenotypes from two incipient branches prevents evolutionary branching. In contrast, when mating is assortative for the ecological characters under study, evolutionary branching is possible in sexual populations and can lead to speciation. Therefore, we also study the evolution of assortative mating as a quantitative character. We show that evolution under branching conditions selects for assortativeness and thus allows sexual populations to escape from fitness minima. We conclude that evolutionary branching offers a general basis for understanding adaptive speciation and radiation under a wide range of different ecological conditions.

Measuring probabilistic reaction norms for age and size at maturation.

Abstract We present a new probabilistic concept of reaction norms for age and size at maturation that is applicable when observations are carried out at discrete time intervals. This approach can also be used to estimate reaction norms for age and size at metamorphosis or at other ontogenetic transitions. Such estimations are critical for understanding phenotypic plasticity and life‐history changes in variable environments, assessing genetic changes in the presence of phenotypic plasticity, and calibrating size‐ and age‐structured population models. We show that previous approaches to this problem, based on regressing size against age at maturation, give results that are systematically biased when compared to the probabilistic reaction norms. The bias can be substantial and is likely to lead to qualitatively incorrect conclusions; it is caused by failing to account for the probabilistic nature of the maturation process. We explain why, instead, robust estimations of maturation reaction norms should be based on logistic regression or on other statistical models that treat the probability of maturing as a dependent variable. We demonstrate the utility of our approach with two examples. First, the analysis of data generated for a known reaction norm highlights some crucial limitations of previous approaches. Second, application to the northeast arctic cod (Gadus morhua) illustrates how our approach can be used to shed new light on existing real‐world data.

Adaptive changes in harvested populations: plasticity and evolution of age and size at maturation

We investigate harvest–induced adaptive changes in age and size at maturation by modelling both plastic variation and evolutionary trajectories. Harvesting mature individuals displaces the reaction norm for age and size at maturation toward older ages and larger sizes and rotates it clockwise, whereas harvesting immature individuals has the reverse qualitative effect. If both immature and mature individuals are harvested, the net effect has approximately the same trend as when harvesting immature individuals only. This stems from the sensitivity of the evolutionary response, which depends on the maturity state of harvested individuals, but also on the type of harvest mortality (negatively or positively density dependent, density independent) and the value of three life–history parameters (natural mortality, growth rate and the trade–off between growth and reproduction). Evolutionary changes in the maturation reaction norm have strong repercussions for the mean size and the density of harvested individuals that, in most cases, result in the reduction of biomass—a response that population dynamical models would overlook. These results highlight the importance of accounting for evolutionary trends in the long–term management of exploited living resources and give qualitative insights into how to minimize the detrimental consequences of harvest–induced evolutionary changes in maturation reaction norms.

POPULATION GROWTH IN SPACE AND TIME: SPATIAL LOGISTIC EQUATIONS

How great an effect does self-generated spatial structure have on logistic population growth? Results are described from an individual-based model (IBM) with spatially localized dispersal and competition, and from a deterministic approximation to the IBM describing the dynamics of the first and second spatial moments. The dynamical system incorporates a novel closure that gives a close approximation to the IBM in the presence of strong spatial structure. Population growth given by the spatial logistic model can differ greatly from that of the nonspatial logistic equation. Numerical simulations show that populations may grow more slowly or more rapidly than would be expected from the nonspatial model, and may reach their maximum rate of increase at densities other than half of the carrying capacity. Populations can achieve asymptotic densities substantially greater than or less than the carrying capacity of the nonspatial logistic model, and can even tend towards extinction. These properties of the spatial logistic model are caused by local dispersal and competition that affect spatial structure, which in turn affects population growth. Accounting for these local spatial processes brings the theory of single-species population growth a step closer to the growth of real spatially structured populations.