Ecological speciation! Or the lack thereof?This Perspective is based on the author’s J.C. Stevenson Memorial Lecture delivered at the Canadian Conference for Fisheries Research in Halifax, Nova Scotia, January 2008.

2009 ◽  
Vol 66 (8) ◽  
pp. 1383-1398 ◽  
Author(s):  
Andrew P. Hendry

Ecological speciation occurs when adaptation to different environments or resources causes the evolution of reproductive isolation. This process is now thought to be very important in the evolution of biological diversity. Indeed, support for ecological speciation is so often asserted in the literature that one can get the impression of ubiquity. Eager to ride on the coattails of this exciting work, my own research has investigated ecological speciation in guppies, sticklebacks, and Darwin’s finches. Much to my initial dismay, I failed to find simple and strong signatures of ecological speciation in the first two of these systems. Setting aside the possibility of personal incompetence, my apparent deviation from ubiquity might simply reflect an existing literature bias. This bias seems obvious in retrospect given that essentially all published studies of ecological speciation purport to be confirmatory, whereas many cases of divergent selection and adaptive divergence are associated with only weak to modest levels of reproductive isolation. In short, different populations can be arrayed along a continuum from panmixia to complete reproductive isolation. Variation along this continuum might profitably be used for studying factors, outlined herein, that can promote or constrain “progress” toward ecological speciation.

2013 ◽  
Vol 2013 ◽  
pp. 1-15 ◽  
Author(s):  
David Bierbach ◽  
Marina Penshorn ◽  
Sybille Hamfler ◽  
Denise B. Herbert ◽  
Jessica Appel ◽  
...  

Ecological speciation assumes reproductive isolation to be the product of ecologically based divergent selection. Beside natural selection, sexual selection via phenotype-assortative mating is thought to promote reproductive isolation. Using the neotropical fishPoecilia mexicanafrom a system that has been described to undergo incipient ecological speciation in adjacent, but ecologically divergent habitats characterized by the presence or absence of toxic H2S and darkness in cave habitats, we demonstrate a gradual change in male body colouration along the gradient of light/darkness, including a reduction of ornaments that are under both inter- and intrasexual selection in surface populations. In dichotomous choice tests using video-animated stimuli, we found surface females to prefer males from their own population over the cave phenotype. However, female cave fish, observed on site via infrared techniques, preferred to associate with surface males rather than size-matched cave males, likely reflecting the female preference for better-nourished (in this case: surface) males. Hence, divergent selection on body colouration indeed translates into phenotype-assortative mating in the surface ecotype, by selecting against potential migrant males. Female cave fish, by contrast, do not have a preference for the resident male phenotype, identifying natural selection against migrants imposed by the cave environment as the major driver of the observed reproductive isolation.


2012 ◽  
Vol 2012 ◽  
pp. 1-20 ◽  
Author(s):  
Anssi Karvonen ◽  
Ole Seehausen

Research on speciation and adaptive radiation has flourished during the past decades, yet factors underlying initiation of reproductive isolation often remain unknown. Parasites represent important selective agents and have received renewed attention in speciation research. We review the literature on parasite-mediated divergent selection in context of ecological speciation and present empirical evidence for three nonexclusive mechanisms by which parasites might facilitate speciation: reduced viability or fecundity of immigrants and hybrids, assortative mating as a pleiotropic by-product of host adaptation, and ecologically-based sexual selection. We emphasise the lack of research on speciation continuums, which is why no study has yet made a convincing case for parasite driven divergent evolution to initiate the emergence of reproductive isolation. We also point interest towards selection imposed by single versus multiple parasite species, conceptually linking this to strength and multifariousness of selection. Moreover, we discuss how parasites, by manipulating behaviour or impairing sensory abilities of hosts, may change the form of selection that underlies speciation. We conclude that future studies should consider host populations at variable stages of the speciation process, and explore recurrent patterns of parasitism and resistance that could pinpoint the role of parasites in imposing the divergent selection that initiates ecological speciation.


2012 ◽  
Vol 58 (1) ◽  
pp. 21-34 ◽  
Author(s):  
Megan V. Mcphee ◽  
David L. G. Noakes ◽  
Fred W. Allendorf

Abstract Morphologically divergent ecotypes arise in fish populations on postglacial time scales, and resource polymorphisms are often invoked to explain their origin. However, genetic recombination can constrain the ability of divergent selection to produce reproductive isolation in sympatry. Recombination breaks up favorable combinations of traits (“adaptive suites”) if individual traits are affected by different loci. Recombination also breaks up any association between traits under divergent selection and traits contributing to reproductive isolation. Thus, ecological speciation in the absence of preexisting barriers to gene flow is more likely when pleiotropy minimizes the number of loci involved. Here, we revisit research conducted by Carl Hubbs in the early 1900s on the effects of developmental rate on morphological traits in fishes. Hubbs’ work provides a mechanism to explain how sympatric divergence by trophic polymorphism can occur despite the challenges of recombination. We consider the implications of Hubbs’ observations for ecological speciation with gene flow in fishes, as well as rapid evolution in captive fish populations.


2019 ◽  
Vol 59 (4) ◽  
pp. 900-911 ◽  
Author(s):  
M Tobler ◽  
N Barts ◽  
R Greenway

Abstract Mitochondria have been known to be involved in speciation through the generation of Dobzhansky–Muller incompatibilities, where functionally neutral co-evolution between mitochondrial and nuclear genomes can cause dysfunction when alleles are recombined in hybrids. We propose that adaptive mitochondrial divergence between populations can not only produce intrinsic (Dobzhansky–Muller) incompatibilities, but could also contribute to reproductive isolation through natural and sexual selection against migrants, post-mating prezygotic isolation, as well as by causing extrinsic reductions in hybrid fitness. We describe how these reproductive isolating barriers can potentially arise through adaptive divergence of mitochondrial function in the absence of mito-nuclear coevolution, a departure from more established views. While a role for mitochondria in the speciation process appears promising, we also highlight critical gaps of knowledge: (1) many systems with a potential for mitochondrially-mediated reproductive isolation lack crucial evidence directly linking reproductive isolation and mitochondrial function; (2) it often remains to be seen if mitochondrial barriers are a driver or a consequence of reproductive isolation; (3) the presence of substantial gene flow in the presence of mito-nuclear incompatibilities raises questions whether such incompatibilities are strong enough to drive speciation to completion; and (4) it remains to be tested how mitochondrial effects on reproductive isolation compare when multiple mechanisms of reproductive isolation coincide. We hope this perspective and the proposed research plans help to inform future studies of mitochondrial adaptation in a manner that links genotypic changes to phenotypic adaptations, fitness, and reproductive isolation in natural systems, helping to clarify the importance of mitochondria in the formation and maintenance of biological diversity.


2012 ◽  
Vol 106 (1) ◽  
pp. 1-17 ◽  
Author(s):  
RÜDIGER RIESCH ◽  
LANCE G. BARRETT-LENNARD ◽  
GRAEME M. ELLIS ◽  
JOHN K. B. FORD ◽  
VOLKER B. DEECKE

2011 ◽  
Vol 2011 ◽  
pp. 1-15 ◽  
Author(s):  
Aneil F. Agrawal ◽  
Jeffrey L. Feder ◽  
Patrik Nosil

The evolution of intrinsic postmating isolation has received much attention, both historically and in recent studies of speciation genes. Intrinsic isolation often stems from between-locus genetic incompatibilities, where alleles that function well within species are incompatible with one another when brought together in the genome of a hybrid. It can be difficult for such incompatibilities to originate when populations diverge with gene flow, because deleterious genotypic combinations will be created and then purged by selection. However, it has been argued that if genes underlying incompatibilities are themselves subject to divergent selection, then they might overcome gene flow to diverge between populations, resulting in the origin of incompatibilities. Nonetheless, there has been little explicit mathematical exploration of such scenarios for the origin of intrinsic incompatibilities during ecological speciation with gene flow. Here we explore theoretical models for the origin of intrinsic isolation where genes subject to divergent natural selection also affect intrinsic isolation, either directly or via linkage disequilibrium with other loci. Such genes indeed overcome gene flow, diverge between populations, and thus result in the evolution of intrinsic isolation. We also examine barriers to neutral gene flow. Surprisingly, we find that intrinsic isolation sometimes weakens this barrier, by impeding differentiation via ecologically based divergent selection.


2013 ◽  
Vol 9 (5) ◽  
pp. 20130327 ◽  
Author(s):  
Matthew Schrader ◽  
Rebecca C. Fuller ◽  
Joseph Travis

Crosses between populations or species often display an asymmetry in the fitness of reciprocal F 1 hybrids. This pattern, referred to as isolation asymmetry or Darwin's Corollary to Haldane's Rule, has been observed in taxa from plants to vertebrates, yet we still know little about which factors determine its magnitude and direction. Here, we show that differences in offspring size predict the direction of isolation asymmetry observed in crosses between populations of a placental fish, Heterandria formosa . In crosses between populations with differences in offspring size, high rates of hybrid inviability occur only when the mother is from a population characterized by small offspring. Crosses between populations that display similarly sized offspring, whether large or small, do not result in high levels of hybrid inviability in either direction. We suggest this asymmetric pattern of reproductive isolation is due to a disruption of parent–offspring coadaptation that emerges from selection for differently sized offspring in different populations.


2019 ◽  
Author(s):  
Joseph A. McGirr ◽  
Christopher H. Martin

AbstractEcological speciation occurs when reproductive isolation evolves as a byproduct of adaptive divergence between populations. However, it is unknown whether divergent ecological selection on gene regulation can directly cause reproductive isolation. Selection favoring regulatory divergence between species could result in gene misregulation in F1 hybrids and ultimately lower hybrid fitness. We combined 58 resequenced genomes with 124 transcriptomes to test this hypothesis in a young, sympatric radiation of Cyprinodon pupfishes endemic to San Salvador Island, Bahamas, which consists of a dietary generalist and two novel trophic specialists – a molluscivore and a scale-eater. We found more differential gene expression between closely related sympatric specialists than between allopatric generalist populations separated by 1000 km. Intriguingly, 9.6% of genes that were differentially expressed between sympatric species were also misregulated in their F1 hybrids. Consistent with divergent ecological selection causing misregulation, a subset of these genes were in highly differentiated genomic regions and enriched for functions important for trophic specialization, including head, muscle, and brain development. These regions also included genes that showed evidence of hard selective sweeps and were significantly associated with oral jaw length – the most rapidly diversifying skeletal trait in this radiation. Our results indicate that divergent ecological selection in sympatry can cause hybrid gene misregulation which may act as a primary reproductive barrier between nascent species.SignificanceIt is unknown whether the same genes that regulate ecological traits can simultaneously contribute to reproductive barriers between species. We measured gene expression in two trophic specialist species of Cyprinodon pupfishes that rapidly diverged from a generalist ancestor. We found genes differentially expressed between species that also showed extreme expression levels in their hybrid offspring. Many of these genes showed signs of selection and have putative effects on the development of traits that are important for ecological specialization. This suggests that genetic variants contributing to adaptive trait divergence between parental species negatively interact to cause hybrid gene misregulation, potentially producing unfit hybrids. Such loci may be important barriers to gene flow during the early stages of speciation, even in sympatry.


Author(s):  
Zachariah Gompert

Biological diversity results from speciation, which generally involves the splitting of an ancestral species into descendant species due to adaptation to different niches or the evolution of reproductive incompatibilities (Coyne and Orr 2004). The diverse flora and fauna of the world, including the native inhabitants of the Greater Yellowstone Area (GYA), exist as a result of the speciation process. The central role speciation plays in generating biological diversity imbues importance to our understanding of this process. The general importance of a thorough understanding of speciation is amplified because of the current high rates of extinction on the planet. This is because a long term solution to the present extinction crisis will require maintaining the processes that create species (speciation) not simply preventing extinction. However, many central questions regarding speciation remain to be answered. One fundamental question in speciation research is whether diverging species are isolated (i.e., prevented from interbreeding) due to differences in one, a few, or many characters and whether each of these character differences results from different alleles at a few or many genes. For example, speciation and reproductive isolation might involve divergence along multiple phenotypic axes, such as mate preference, habitat use or preference, and phenology (the timing of life-cycle events). Alternatively, isolation could result from differentiation of a single character. I propose to address this question by assessing patterns of variation for a suite of characters across a hybrid zone between two butterfly species. This is possible because patterns of character variation across hybrids zones allow for inferences about reproductive isolation (Barton and Hewitt 1985).


Author(s):  
Zachariah Gompert

Biological diversity results from speciation, which generally involves the splitting of an ancestral species into descendant species due to adaptation to different niches or the evolution of reproductive incompatibilities (Coyne and Orr 2004). The diverse flora and fauna of the world, including the native inhabitants of the Greater Yellowstone Area (GYA), exist as a result of the speciation process. The central role speciation plays in generating biological diversity imbues importance to our understanding of this process. The general importance of a thorough understanding of speciation is amplified because of the current high rates of extinction on the planet. This is because a long term solution to the present extinction crisis will require maintaining the processes that create species (speciation) not simply preventing extinction. However, many central questions regarding speciation remain to be answered. One fundamental question in speciation research is whether diverging species are isolated (i.e., prevented from interbreeding) due to differences in one, a few, or many characters and whether each of these character differences results from different alleles at a few or many genes. For example, speciation and reproductive isolation might involve divergence along multiple phenotypic axes, such as mate preference, habitat use or preference, and phenology (the timing of life-cycle events). Alternatively, isolation could result from differentiation of a single character. I propose to address this question by assessing patterns of variation for a suite of characters across a hybrid zone between two butterfly species. This is possible because patterns of character variation across hybrids zones allow for inferences about reproductive isolation (Barton and Hewitt 1985).


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