K. Emily Knott

What do we need to know about speciation

Speciation has been a major focus of evolutionary biology research in recent years, with many important advances. However, some of the traditional organising principles of the subject area no longer provide a satisfactory framework, such as the classification of speciation mechanisms by geographical context into allopatric, parapatric and sympatry classes. Therefore, we have asked where speciation research should be directed in the coming years. Here, we present a distillation of questions about the mechanisms of speciation, the genetic basis of speciation and the relationship between speciation and diversity. Our list of topics is not exhaustive; rather we aim to promote discussion on research priorities and on the common themes that underlie disparate speciation processes.

Adaptation to a seasonally varying environment: a strong latitudinal cline in reproductive diapause combined with high gene flow in Drosophila montana

Adaptation to seasonal changes in the northern hemisphere includes an ability to predict the forthcoming cold season from gradual changes in environmental cues early enough to prepare for the harsh winter conditions. The magnitude and speed of changes in these cues vary between the latitudes, which induces strong selection pressures for local adaptation. We studied adaptation to seasonal changes in Drosophila montana, a northern maltfly, by defining the photoperiodic conditions leading to adult reproductive diapause along a latitudinal cline in Finland and by measuring genetic differentiation and the amount of gene flow between the sampling sites with microsatellites. Our data revealed a clear correlation between the latitude and the critical day length (CDL), in which half of the females of different cline populations enter photoperiodic reproductive diapause. There was no sign of limited gene flow between the cline populations, even though these populations showed isolation by distance. Our results show that local adaptation may occur even in the presence of high gene flow, when selection for locally adaptive life-history traits is strong. A wide range of variation in the CDLs of the fly strains within and between the cline populations may be partly due to gene flow and partly due to the opposing selection pressures for fly reproduction and overwinter survival. This variation in the timing of diapause will enhance populations’ survival over the years that differ in the severity of the winter and in the length of the warm period and may also help them respond to long-term changes in environmental conditions.

GENETIC VARIABILITY AND DRIFT LOAD IN POPULATIONS OF AN AQUATIC SNAIL

Abstract Population genetic theory predicts that in small populations, random genetic drift will fix and accumulate slightly deleterious mutations, resulting in reduced reproductive output. This genetic load due to random drift (i.e., drift load) can increase the extinction risk of small populations. We studied the relationship between genetic variability (indicator of past population size) and reproductive output in eight isolated, natural populations of the hermaphroditic snail Lymnaea stagnalis. In a common laboratory environment, snails from populations with the lowest genetic variability mature slower and have lower fecundity than snails from genetically more variable populations. This result suggests that past small population size has resulted in increased drift load, as predicted. The relationship between genetic variability and reproductive output is independent of the amount of nonrandom mating within populations. However, reproductive output and the current density of snails in the populations were not correlated. Instead, data from the natural populations suggest that trematode parasites may determine, at least in part, population densities of the snails.

THE EFFECTS OF MATING SYSTEM AND GENETIC VARIABILITY ON SUSCEPTIBILITY TO TREMATODE PARASITES IN A FRESHWATER SNAIL, LYMNAEA STAGNALIS

Abstract The amount and distribution of genetic variability in host populations can have significant effects on the outcome of host‐parasite interactions. We studied the effect of mating system and genetic variability on susceptibility of Lymnaea stagnalis snails to trematode parasites. Mating system of snails from eight populations differing in the amount of genetic variability was manipulated, and self‐ and cross‐fertilized offspring were exposed to naturally occurring trematode parasites in a controlled lake experiment. Susceptibility of snails varied between populations, but mating‐system treatment did not have a significant effect. Heterozygosity of snails was negatively correlated with the probability of trematode infection, however, suggesting that parasitic diseases may pose a serious threat to populations lacking genetic variability.

Predominance of outcrossing in Lymnaea stagnalis despite low apparent fitness costs of self-fertilization

We have quantified the natural mating system in eight populations of the simultaneously hermaphroditic aquatic snail Lymnaea stagnalis, and studied the ecological and genetic forces that may be directing mating system evolution in this species. We investigated whether the natural mating system can be explained by the availability of mates, by the differential survival of self‐ and cross‐fertilized snails in nature, and by the effects of mating system on parental fecundity and early survival. The natural mating system of L. stagnalis was found to be predominantly cross‐fertilizing. Density of snails in the populations had no relationship with the mating system, suggesting that outcrossing rates are not limited by mate availability at the population densities observed. Contrary to expectations for outcrossing species, we detected no evidence for inbreeding depression in survival in nature with inferential population genetic methods. Further, experimental manipulations of mating system in the laboratory revealed that self‐fertilization had no effect on parental fecundity, and only minor effects on offspring survival. Predominance of cross‐fertilization despite low apparent fitness costs of self‐fertilization is at odds with the paradigm that high self‐fertilization depression is necessary for maintenance of cross‐fertilization in self‐compatible hermaphrodites.

Identification of Asteroid Genera With Species Capable of Larval Cloning

Asexual reproduction in larvae, larval cloning, is a recently recognized component of the complex life histories of asteroids. We compare DNA sequences of mitochondrial tRNA genes (Ala, Leu, Asn, Pro, and Gln) from larvae in the process of cloning collected in the field with sequences from adults of known species in order to identify asteroid taxa capable of cloning. Neighbor-joining analysis identified four distinct groups of larvae, each having no, or very little, sequence divergence (p distances ranging from 0.00000 to 0.02589); thus, we conclude that each larval group most likely represents a single species. These field-collected larvae cannot be identified to species with certainty, but the close assemblage of known taxa with the four larval groups indicates generic or familial identity. We can assign two of the larval groups discerned here to the genera Luidia and Oreaster and another two to the family Ophidiasteridae. This study is the first to identify field-collected cloning asteroid larvae, and provides evidence that larval cloning is phylogenetically widespread within the Asteroidea. Additionally, we note that cloning occurs regularly and in multiple ways within species that are capable of cloning, emphasizing the need for further investigation of the role of larval cloning in the ecology and evolution of asteroids.

Introduction to Symposium: Poecilogony—A Window on Larval Evolutionary Transitions in Marine Invertebrates

Poecilogony is the intraspecific variation in developmental mode that has been described in some marine invertebrates. Poecilogonous species produce different larval forms (e.g., free-swimming planktotrophic larvae as well as brooded lecithotrophic or adelphophagic larvae). Poecilogony can be a controversial topic, since it is difficult to identify and characterize the phenomenon with certainty. It has been challenging to determine whether poecilogony represents developmental polymorphism with a genetic basis or developmental polyphenism reflecting plastic responses to environmental cues. Other outstanding questions include whether common mechanisms underlie the developmental variation we observe in poecilogonous species, and whether poecilogony is maintained in different taxa through similar mechanisms or selective pressures. Poecilogonous species provide a unique opportunity to elucidate the cellular, developmental, and genetic mechanisms underlying evolutionary transitions in developmental mode, as well as to help clarify the selective pressures and possible ecological circumstances that might be involved. Here, we describe an integrative approach to the study of poecilogony and its role in larval evolutionary transitions highlighted during a symposium held at the 2012 annual meeting of the Society for Integrative and Comparative Biology.

Polymorphism in developmental mode and its effect on population genetic structure of a spionid polychaete, Pygospio elegans.

Population genetic structure of sedentary marine species is expected to be shaped mainly by the dispersal ability of their larvae. Long-lived planktonic larvae can connect populations through migration and gene flow, whereas species with nondispersive benthic or direct-developing larvae are expected to have genetically differentiated populations. Poecilogonous species producing different larval types are ideal when studying the effect of developmental mode on population genetic structure and connectivity. In the spionid polychaete Pygospio elegans, different larval types have been observed between, and sometimes also within, populations. We used microsatellite markers to study population structure of European P. elegans from the Baltic Sea (BS) and North Sea (NS). We found that populations with planktonic larvae had higher genetic diversity than did populations with benthic larvae. However, this pattern may not be related to developmental mode, since in P. elegans, developmental mode may be associated with geography. Benthic larvae were more commonly seen in the brackish BS and planktonic larvae were predominant in the NS, although both larval types also are found from both areas. Significant isolation-by-distance (IBD) was found overall and within regions. Most of the pair-wise F(ST) comparisons among populations were significant, although some geographically close populations with planktonic larvae were found to be genetically similar. However, these results, together with the pattern of IBD, autocorrelation within populations, as well as high estimated local recruitment, suggest that dispersal is limited in populations with planktonic larvae as well as in those with benthic larvae. The decrease in salinity between the NS and BS causes a barrier to gene flow in many marine species. In P. elegans, low, but significant, differentiation was detected between the NS and BS (3.34% in AMOVA), but no clear transition zone was observed, indicating that larvae are not hampered by the change in salinity.

Temporal genetic structure in a poecilogonous polychaete: the interplay of developmental mode and environmental stochasticity.

BackgroundTemporal variation in the genetic structure of populations can be caused by multiple factors, including natural selection, stochastic environmental variation, migration, or genetic drift. In benthic marine species, the developmental mode of larvae may indicate a possibility for temporal genetic variation: species with dispersive planktonic larvae are expected to be more likely to show temporal genetic variation than species with benthic or brooded non-dispersive larvae, due to differences in larval mortality and dispersal ability. We examined temporal genetic structure in populations of Pygospio elegans, a poecilogonous polychaete with within-species variation in developmental mode. P. elegans produces either planktonic, benthic, or intermediate larvae, varying both among and within populations, providing a within-species test of the generality of a relationship between temporal genetic variation and larval developmental mode.ResultsIn contrast to our expectations, our microsatellite analyses of P. elegans revealed temporal genetic stability in the UK population with planktonic larvae, whereas there was variation indicative of drift in temporal samples of the populations from the Baltic Sea, which have predominantly benthic and intermediate larvae. We also detected temporal variation in relatedness within these populations. A large temporal shift in genetic structure was detected in a population from the Netherlands, having multiple developmental modes. This shift could have been caused by local extiction due to extreme environmental conditions and (re)colonization by planktonic larvae from neighboring populations.ConclusionsIn our study of P. elegans, temporal genetic variation appears to be due to not only larval developmental mode, but also the stochastic environment of adults. Large temporal genetic shifts may be more likely in marine intertidal habitats (e.g. North Sea and Wadden Sea) which are more prone to environmental stochasticity than the sub-tidal Baltic habitats. Sub-tidal and/or brackish (less saline) habitats may support smaller P. elegans populations and these may be more susceptible to the effects of random genetic drift. Moreover, higher frequencies of asexual reproduction and the benthic larval developmental mode in these populations leads to higher relatedness and contributes to drift. Our results indicate that a general relationship between larval developmental mode and temporal genetic variation may not exist.

Low Levels of Mitochondrial DNA and Symbiont Diversity in the Worldwide Agricultural Pest, the Greenhouse Whitefly Trialeurodes vaporariorum (Hemiptera: Aleyrodidae)

Trialeurodes vaporariorum, the greenhouse whitefly, is a cosmopolitan agricultural pest. Little is known about the genetic diversity of T. vaporariorum and the bacterial symbionts associated with this species. Here, we undertook a large phylogeographic study by investigating both the mitochondrial (mt) diversity and the infection status of 38 T. vaporariorum collections from 18 countries around the world. Genetic diversity of T. vaporariorum was studied by analyzing sequence data from the mt cytochrome oxidase I, cytochrome b, and NADH dehydrogenase subunit 5 genes. Maximum-likelihood (ML) phylogeny reconstruction delineated 2 clades characterized by limited sequence divergence: one clade comprised samples only from the Northern hemisphere whereas the other comprised samples from a broader geographical range. The presence of secondary symbionts was determined by PCR using primers specific for Hamiltonella, Rickettsia, Arsenophonus, Cardinium, Wolbachia, and Fritschea. Most individuals examined harbored at least one secondary endosymbiont, and Arsenophonus was detected in almost all male and female individuals. Wolbachia was present at a much lower frequency, and Cardinium was detected in only a few individuals from Greece. Rickettsia, Hamiltonella, and Fritschea were not found. Additionally, we set out to further analyze Arsenophonus diversity by multilocus sequence typing analysis; however, the Arsenophonus sequences did not exhibit any polymorphism. Our results revealed remarkably low diversity in both mtDNA and symbionts in this worldwide agricultural pest, contrasting sharply with that of the ecologically similar Bemisia tabaci.