scholarly journals Sympatric ecological divergence with coevolution of niche preference

2020 ◽  
Vol 375 (1806) ◽  
pp. 20190749 ◽  
Author(s):  
Pavel Payne ◽  
Jitka Polechová
Keyword(s):  
Genetic Variation ◽  
Gene Flow ◽  
Reproductive Isolation ◽  
Parameter Space ◽  
Sympatric Speciation ◽  
Secondary Contact ◽  
Disruptive Selection ◽  
Theme Issue ◽  
Ecological Divergence ◽  
Trade Off

Reinforcement, the increase of assortative mating driven by selection against unfit hybrids, is conditional on pre-existing divergence. Yet, for ecological divergence to precede the evolution of assortment, strict symmetries between fitnesses in niches must hold, and/or there must be low gene flow between the nascent species. It has thus been argued that conditions favouring sympatric speciation are rarely met in nature. Indeed, we show that under disruptive selection, violating symmetries in niche sizes and increasing strength of the trade-off in selection between the niches quickly leads to loss of genetic variation, instead of evolution of specialists. The region of the parameter space where polymorphism is maintained further narrows with increasing number of loci encoding the diverging trait and the rate of recombination between them. Yet, evolvable assortment and pre-existing assortment both substantially broaden the parameter space within which polymorphism is maintained. Notably, pre-existing niche preference speeds up further increase of assortment, thus facilitating reinforcement in the later phases of speciation. We conclude that in order for sympatric ecological divergence to occur, niche preference must coevolve throughout the divergence process. Even if populations come into secondary contact, having diverged in isolation, niche preference substantially broadens the conditions for coexistence in sympatry and completion of the speciation process. This article is part of the theme issue ‘Towards the completion of speciation: the evolution of reproductive isolation beyond the first barriers'.

scholarly journals Sympatric ecological divergence with coevolution of niche preference

2019 ◽  
Author(s):  
Pavel Payne ◽  
Jitka Polechová
Keyword(s):  
Genetic Variation ◽  
Gene Flow ◽  
Assortative Mating ◽  
Parameter Space ◽  
Sympatric Speciation ◽  
Secondary Contact ◽  
Disruptive Selection ◽  
Ecological Divergence ◽  
Trade Off ◽  
Speciation Process

AbstractReinforcement, the increase of assortative mating driven by selection against unfit hybrids, is conditional on pre-existing divergence. Yet, for ecological divergence to precede the evolution of assortment, strict symmetries between fitnesses in niches must hold, and/or there must be low gene flow between the nascent species. It has thus been argued that conditions favouring sympatric speciation are rarely met in nature. Indeed, we show that under disruptive selection, violating symmetries in niche sizes and increasing strength of the trade-off in selection between the niches quickly leads to loss of genetic variation, instead of evolution of specialists. The region of the parameter space where polymorphism is maintained further narrows with increasing number of loci encoding the diverging trait and the rate of recombination between them. Yet, evolvable assortment and pre-existing assortment both substantially broaden the parameter space within which polymorphism is maintained. Notably, pre-existing niche preference speeds up further increase of assortment, thus facilitating reinforcement in the later phases of speciation. We conclude that in order for sympatric ecological divergence to occur, niche preference must co-evolve throughout the divergence process. Even if populations come into secondary contact, having diverged in isolation, niche preference substantially broadens the conditions for coexistence in sympatry and completion of the speciation process.

scholarly journals Speciation in the Cladophorales species flock of ancient Lake Baikal

2021 ◽  
Author(s):  
◽  
Sergio Diaz Martinez
Keyword(s):  
Population Genetics ◽  
Genetic Variation ◽  
Reproductive Isolation ◽  
Lake Baikal ◽  
Species Delimitation ◽  
Sympatric Speciation ◽  
Species Flock ◽  
Ancient Lake ◽  
Data Set ◽  
Dispersal Abilities

<p>Understanding speciation is one of the great challenges in evolutionary biology as many of the processes involved in speciation, as well as the forces leading to morphological and genetic differentiation, are not fully understood. Three main modes of speciation have been described: allopatric, parapatric and sympatric. Sympatric speciation is the most enigmatic mode because in the absence of physical barriers, disruptive selection, assortative mating and hybridization play central roles in reproductive isolation. Although it is accepted that sympatric speciation is possible, only a few examples of this process exist to date. Another common method of speciation in plants and algae is via polyploidization. Recently, a promising system to study speciation in sympatry was discovered: the endemic Cladophorales species flock in ancient Lake Baikal, Russia. The flock consists of sixteen taxa grouped in four genera: Chaetocladiella, Chaetomorpha, Cladophora and Gemmiphora. In spite of their morphological diversity, recent molecular analyses have shown that this is a monophyletic group with low genetic variation and nested within the morphologically simple genus Rhizoclonium. Due to their high number of species, endemism and sympatric distribution, many interesting questions have arisen such as what processes are involved in speciation, and whether this group might be a novel example of sympatric speciation. In this study, we analysed the population genetics of the endemic Baikalian Cladophorales to infer the processes shaping the evolution of the group. First, a set of microsatellites was designed using high-throughput sequencing data. Second, species delimitation methods based on genetic clustering were performed. Third, the population genetics of three widely distributed species was analysed looking for evidence of panmixia, a common criteria to support sympatric speciation. A total of 11 microsatellites that mostly cross-amplify between most species were obtained. The genotyping revealed that most loci had more than two alleles per individual indicating polyploidy. As such, the analyses required a different approach which consisted in coding the genotypes as ‘allelic phenotypes’, allowing the use of individuals of different ploidy levels in the same data set. The species delimitation of 15 operative morphotaxa and 727 individuals supported reproductive isolation of five morphotaxa and two hypotheses of conspecificity. However, some morphotaxa showed unclear assignments revealing the need of further research to clarify their reproductive limits. Finally, the population genetics of Chaetomorpha moniliformis, Cladophora compacta and Cl. kursanovii revealed patterns of genetic variation and structure that suggest different reproductive strategies and dispersal abilities. This demonstrates that contrasting biological characteristics may arise in closely related lineages: Chaetomorpha moniliformis with dominant asexual reproduction and long dispersal abilities; Cladophora compacta with high genetic diversity, no population structure and likely to reproduce sexually; Cl. kursanovii with a structure congruent with geographic distribution and more restricted dispersal. The results suggest that polyploidy, rather than speciation with gene flow, is the force driving the reproductive isolation and evolution of this flock. Although many questions remain to be studied, this research provides the first insights into the diversification of this Cladophorales species flock and contributes to the understanding of speciation in freshwater algae.</p>

scholarly journals The limits to parapatric speciation 3: evolution of strong reproductive isolation in presence of gene flow despite limited ecological differentiation

2020 ◽  
Vol 375 (1806) ◽  
pp. 20190532 ◽  
Author(s):  
Alexandre Blanckaert ◽  
Claudia Bank ◽  
Joachim Hermisson
Keyword(s):  
Gene Flow ◽  
Reproductive Isolation ◽  
Local Adaptation ◽  
Theme Issue ◽  
Genetic Barrier ◽  
Incipient Species ◽  
Postzygotic Reproductive Isolation ◽  
Minimal Configuration ◽  
Parapatric Speciation ◽  
Two Populations

Gene flow tends to impede the accumulation of genetic divergence. Here, we determine the limits for the evolution of postzygotic reproductive isolation in a model of two populations that are connected by gene flow. We consider two selective mechanisms for the creation and maintenance of a genetic barrier: local adaptation leads to divergence among incipient species due to selection against migrants, and Dobzhansky–Muller incompatibilities (DMIs) reinforce the genetic barrier through selection against hybrids. In particular, we are interested in the maximum strength of the barrier under a limited amount of local adaptation, a challenge that many incipient species may initially face. We first confirm that with classical two-locus DMIs, the maximum amount of local adaptation is indeed a limit to the strength of a genetic barrier. However, with three or more loci and cryptic epistasis, this limit holds no longer. In particular, we identify a minimal configuration of three epistatically interacting mutations that is sufficient to confer strong reproductive isolation. This article is part of the theme issue ‘Towards the completion of speciation: the evolution of reproductive isolation beyond the first barriers’.

scholarly journals Host races in plant–feeding insects and their importance in sympatric speciation

2002 ◽  
Vol 357 (1420) ◽  
pp. 471-492 ◽  
Author(s):  
Michele Drès ◽  
James Mallet
Keyword(s):  
Linkage Disequilibrium ◽  
Gene Flow ◽  
Natural Selection ◽  
Reproductive Isolation ◽  
Sympatric Speciation ◽  
Accurate Information ◽  
Sympatric Populations ◽  
New Host ◽  
Host Races ◽  
Plant Feeding

The existence of a continuous array of sympatric biotypes—from polymorphisms, through ecological or host races with increasing reproductive isolation, to good species—can provide strong evidence for a continuous route to sympatric speciation via natural selection. Host races in plant–feeding insects, in particular, have often been used as evidence for the probability of sympatric speciation. Here, we provide verifiable criteria to distinguish host races from other biotypes: in brief, host races are genetically differentiated, sympatric populations of parasites that use different hosts and between which there is appreciable gene flow. We recognize host races as kinds of species that regularly exchange genes with other species at a rate of more than ca . 1% per generation, rather than as fundamentally distinct taxa. Host races provide a convenient, although admittedly somewhat arbitrary intermediate stage along the speciation continuum. They are a heuristic device to aid in evaluating the probability of speciation by natural selection, particularly in sympatry. Speciation is thereby envisaged as having two phases: (i) the evolution of host races from within polymorphic, panmictic populations; and (ii) further reduction of gene flow between host races until the diverging populations can become generally accepted as species. We apply this criterion to 21 putative host race systems. Of these, only three are unambiguously classified as host races, but a further eight are strong candidates that merely lack accurate information on rates of hybridization or gene flow. Thus, over one–half of the cases that we review are probably or certainly host races, under our definition. Our review of the data favours the idea of sympatric speciation via host shift for three major reasons: (i) the evolution of assortative mating as a pleiotropic by–product of adaptation to a new host seems likely, even in cases where mating occurs away from the host; (ii) stable genetic differences in half of the cases attest to the power of natural selection to maintain multilocus polymorphisms with substantial linkage disequilibrium, in spite of probable gene flow; and (iii) this linkage disequilibrium should permit additional host adaptation, leading to further reproductive isolation via pleiotropy, and also provides conditions suitable for adaptive evolution of mate choice (reinforcement) to cause still further reductions in gene flow. Current data are too sparse to rule out a cryptic discontinuity in the apparently stable sympatric route from host–associated polymorphism to host–associated species, but such a hiatus seems unlikely on present evidence. Finally, we discuss applications of an understanding of host races in conservation and in managing adaptation by pests to control strategies, including those involving biological control or transgenic parasite–resistant plants.

scholarly journals Speciation: more likely through a genetic or through a learned habitat preference?

2005 ◽  
Vol 272 (1571) ◽  
pp. 1455-1463 ◽  
Author(s):  
J.B Beltman ◽  
J.A.J Metz
Keyword(s):  
Reproductive Isolation ◽  
Habitat Preference ◽  
Sympatric Speciation ◽  
Genetic Mechanism ◽  
Dynamical Analysis ◽  
Disruptive Selection ◽  
Single Population ◽  
Preferred Habitat ◽  
Simultaneous Evolution ◽  
Evolution Proceeds

A problem in understanding sympatric speciation is establishing how reproductive isolation can arise when there is disruptive selection on an ecological trait. One of the solutions that has been proposed is that a habitat preference evolves, and that mates are chosen within the preferred habitat. We present a model where the habitat preference can evolve either by means of a genetic mechanism or by means of learning. Employing an adaptive-dynamical analysis, we show that evolution proceeds either to a single population of specialists with a genetic preference for their optimal habitat, or to a population of generalists without a habitat preference. The generalist population subsequently experiences disruptive selection. Learning promotes speciation because it increases the intensity of disruptive selection. An individual-based version of the model shows that, when loci are completely unlinked and learning confers little cost, the presence of disruptive selection most probably leads to speciation via the simultaneous evolution of a learned habitat preference. For high costs of learning, speciation is most likely to occur via the evolution of a genetic habitat preference. However, the latter only happens when the effect of mutations is large, or when there is linkage between genes coding for the different traits.

scholarly journals What role does natural selection play in speciation?

2010 ◽  
Vol 365 (1547) ◽  
pp. 1825-1840 ◽  
Author(s):  
N. H. Barton
Keyword(s):  
Gene Flow ◽  
Habitat Preference ◽  
Genetic Variance ◽  
Bimodal Distribution ◽  
Biological Species ◽  
Disruptive Selection ◽  
Linkage Equilibrium ◽  
Sexual Population ◽  
Trade Off ◽  
High Gene

If distinct biological species are to coexist in sympatry, they must be reproductively isolated and must exploit different limiting resources. A two-niche Levene model is analysed, in which habitat preference and survival depend on underlying additive traits. The population genetics of preference and viability are equivalent. However, there is a linear trade-off between the chances of settling in either niche, whereas viabilities may be constrained arbitrarily. With a convex trade-off, a sexual population evolves a single generalist genotype, whereas with a concave trade-off, disruptive selection favours maximal variance. A pure habitat preference evolves to global linkage equilibrium if mating occurs in a single pool, but remarkably, evolves to pairwise linkage equilibrium within niches if mating is within those niches—independent of the genetics. With a concave trade-off, the population shifts sharply between a unimodal distribution with high gene flow and a bimodal distribution with strong isolation, as the underlying genetic variance increases. However, these alternative states are only simultaneously stable for a narrow parameter range. A sharp threshold is only seen if survival in the ‘wrong’ niche is low; otherwise, strong isolation is impossible. Gene flow from divergent demes makes speciation much easier in parapatry than in sympatry.

scholarly journals Extensive introgression despite Haldane’s rule: insights from grasshopper hybrid zones

2021 ◽  
Author(s):  
Linda Hagberg ◽  
Enrique Celemin ◽  
Iker Irisarri ◽  
Oliver Hawlitschek ◽  
J L Bella ◽  
...  
Keyword(s):  
Gene Flow ◽  
Reproductive Isolation ◽  
Male Sterility ◽  
Secondary Contact ◽  
Hybrid Zones ◽  
Hybrid Male ◽  
Hybrid Male Sterility ◽  
Haldane’S Rule ◽  
Haldane's Rule ◽  
Species Formation

Although the process of species formation is notoriously idiosyncratic, the observation of pervasive patterns of reproductive isolation across species pairs suggests that generalities, or “rules”, underlie species formation in all animals. Haldane’s rule states that whenever a sex is absent, rare or sterile in a cross between two taxa, that sex is usually the heterogametic sex. Yet, understanding how Haldane’s rule first evolves and whether it is associated to genome wide barriers to gene flow remains a challenging task because this rule is usually studied in highly divergent taxa that no longer hybridize in nature. Here, we address these questions using the meadow grasshopper Pseudochorthippus parallelus where populations that readily hybridize in two natural hybrid zones show hybrid male sterility in laboratorial crosses. Using mitochondrial data, we infer that such populations have diverged some 100,000 years ago, surviving multiple glacial periods in isolated Pleistocenic refugia. Nuclear data shows that secondary contact has led to extensive introgression throughout the species range, including between populations showing hybrid male sterility. We find repeatable patterns of genomic differentiation across the two hybrid zones, yet such patterns are consistent with shared genomic constraints across taxa rather than their role in reproductive isolation. Together, our results suggest that Haldane’s rule can evolve relatively quickly within species, particularly when associated to strong demographic changes. At such early stages of species formation, hybrid male sterility still permits extensive gene flow, allowing future studies to identify genomic regions associated with reproductive barriers.

Evolutionary Branching in a Classical Model for Sympatric Speciation

2011 ◽  
Author(s):  
Michael Doebeli
Keyword(s):  
Reproductive Isolation ◽  
Adaptive Dynamics ◽  
Classical Model ◽  
Sympatric Speciation ◽  
Disruptive Selection ◽  
General Outcome ◽  
Wide Range ◽  
Genetic Mechanisms ◽  
Basic Concepts ◽  
Smith Model

This chapter begins by considering the Maynard Smith model. Much of this work concentrated on the genetic mechanisms for assortative mating and reproductive isolation, based on the assumption that the underlying niche ecology would generate disruptive selection. However, understanding the conditions under which disruptive selection arises in the first place is equally important, and indeed necessary for assessing whether diversification is a general outcome in the Maynard Smith model. The chapter then shows that disruptive selection and polymorphism are scenarios that occur generically, that is, for a wide range of parameters, in a classical and widely used speciation model. It also provides an introduction to some of the basic concepts of adaptive dynamics theory.

scholarly journals Variability of Anthoxanthum species in Poland in relation to geographical-historical and environmental conditions: isozyme variation

2013 ◽  
Vol 30 (1) ◽  
pp. 63-93 ◽  
Author(s):  
Maria Drapikowska
Keyword(s):  
Genetic Variation ◽  
Gene Flow ◽  
Reproductive Isolation ◽  
Environmental Conditions ◽  
Habitat Type ◽  
Environmental Variation ◽  
Small Extent ◽  
Isozyme Variation ◽  
Isozyme Markers ◽  
Isozyme Variability

Abstract Variation of 9 isozyme systems was studied in Polish populations of 3 species of the genus Anthoxanthum: the native A. odoratum s. str. L. and A. alpinum Á. Löve & D. Löve, as well as the alien A. aristatum Boiss. Results of this study show that A. odoratum is characterized by a high isozyme variability of lowland populations, weakly correlated with habitat type, and partial genetic distinctness of montane populations. Moreover, 5 isozyme markers have been identified (Pgi-2, Dia-2, Mdh, Idh, Pgm) for the allopolyploid A. odoratum. Populations of A. aristatum are highly polymorphic (P = 98%). The observed isozyme differentiation of its populations (FST = 0.087) is low and gene flow between them (Nm = 5.314) is high. The genetic variation reflects environmental variation only to a small extent and is not significantly related to the phase of chorological expansion of this species. Altitudinal vicariants, A. alpinum and A. odoratum, are characterized by morphological and isozymatic distinctness, indicating their reproductive isolation. In populations of A. alpinum, polymorphism is high (P = 76.92%), differentiation among populations is moderate (FST = 0.198), and gene flow between populations along the altitudinal transect (Nm = 1.709) is relatively low

scholarly journals Asymmetric Reproductive Barriers and Gene Flow Promote the Rise of a Stable Hybrid Zone in the Mediterranean High Mountain

2021 ◽  
Vol 12 ◽  
Author(s):  
Mohamed Abdelaziz ◽  
A. Jesús Muñoz-Pajares ◽  
Modesto Berbel ◽  
Ana García-Muñoz ◽  
José M. Gómez ◽  
...  
Keyword(s):  
Gene Flow ◽  
Reproductive Isolation ◽  
Sierra Nevada ◽  
Hybrid Zone ◽  
High Elevation ◽  
Secondary Contact ◽  
High Mountain ◽  
Hybrid Zones ◽  
Reproductive Barriers ◽  
Sierra Nevada Mountains

Hybrid zones have the potential to shed light on evolutionary processes driving adaptation and speciation. Secondary contact hybrid zones are particularly powerful natural systems for studying the interaction between divergent genomes to understand the mode and rate at which reproductive isolation accumulates during speciation. We have studied a total of 720 plants belonging to five populations from two Erysimum (Brassicaceae) species presenting a contact zone in the Sierra Nevada mountains (SE Spain). The plants were phenotyped in 2007 and 2017, and most of them were genotyped the first year using 10 microsatellite markers. Plants coming from natural populations were grown in a common garden to evaluate the reproductive barriers between both species by means of controlled crosses. All the plants used for the field and greenhouse study were characterized by measuring traits related to plant size and flower size. We estimated the genetic molecular variances, the genetic differentiation, and the genetic structure by means of the F-statistic and Bayesian inference. We also estimated the amount of recent gene flow between populations. We found a narrow unimodal hybrid zone where the hybrid genotypes appear to have been maintained by significant levels of a unidirectional gene flow coming from parental populations and from weak reproductive isolation between them. Hybrid plants exhibited intermediate or vigorous phenotypes depending on the analyzed trait. The phenotypic differences between the hybrid and the parental plants were highly coherent between the field and controlled cross experiments and through time. The highly coherent results obtained by combining field, experimental, and genetic data demonstrate the existence of a stable and narrow unimodal hybrid zone between Erysimum mediohispanicum and Erysimum nevadense at the high elevation of the Sierra Nevada mountains.

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