To present data is human, to communicate data is divine

Finding new and engaging ways to communicate science is of paramount importance. But, how many opportunities are there to practice the art of communication?

That’s how I began the lead-in piece for a series of student posts over a year ago (see piece here and the student posts can be found here).

Giving students the opportunity to hone their communication skills is a must. They need to be adept at engaging with all sorts of people who will cross their paths … from policy makers to scientists in the same field to an interested person when you’re in the field.

Clichés are normally clichés for a reason …. practice makes perfect (or at least a lot better). 

I’ve been lucky enough to expand my Science Communication course at the University of Alabama at Birmingham since I last taught it two years ago this fall (time does fly … more about that in a future post on the meetings I was supposed to cover <<insert chagrin here>>).

Students in the first round were able to write a blog post about a topic of their choice. Each student that submitted the blog post to Jeremy and myself got them published. Not only did they learn how to distill the primary literature, but they each got another line on their CV.

Over the last academic year, I have taught an Evolution course and the revamped Sci Comm, in which grad students in both courses had the opportunity to write a blog post again. I was impressed with the quality and excitement in the first round. I also wanted to try to provide other opportunities for students. As regular readers will know, I have found my time at TME to be incredibly rewarding.

Starting next Tuesday, each week a new blog post written by a student from my graduate Sci Comm course or from the graduate section of my Evolution course will go live.

There’ll be another series of student-written posts in the new year from my new Conservation Genetics course. I’m hoping this can be a series that will continue each time I teach a course with grad students at UAB.

The more I think about science communication … the more I wonder.

Illustration by Maki Naro

Is science communication a bit redundant? Should we not simply communicate? It’s probably a philosophical argument best saved for another day when a two-year review, a late piece for a society newsletter, and several manuscripts aren’t looming.

I hope you enjoy reading the posts over the next few weeks as much as I did working with the students to turn these into publishable pieces of science communication.

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Not my problem

Do American scientists know that doing research in America is a necessary step for many scientists from other parts of the world in order to get a permanent job in academia in their home country?

Once in the US, these researchers face many challenges outside of academia that can significantly affect their survival and well-being, and ultimately, their scientific output. These challenges include health care, visa issues, housing, taxes, the school system, and child care. In America, people can easily fall through the cracks. Many other countries have a safety net that protects you while working at an academic institution. In the US however, offices will not coordinate and fix problems without the affected individuals being involved.

Pssst! The following text is only for postdocs. (I also mean grad students and visiting scholars).

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Evolution 2018: assortative mating, combinatorial speciation and genome dynamics

The Evolution conference in Montpellier is over, and as the sun, wine and great science become a memory, here is my recap of some conference highlights following on from a great first day:

A sea of scientists waiting for a plenary lecture. Photo: Alex Twyford.

Sharon Strauss (University of California Davis) gave the ASN Presidential Address entitled “Diversity and coexistence in close relatives, and reflections on 150 years of the ASN”. In her talk, she discussed the coexistence of closely related plant species, and whether phylogenetic similarity predicts ecological similarity. Her work centres on herbaceous plant communities at the UC Bodega Marine Reserve in California and combines reciprocal transplant experiments with phylogenetic analysis. One of her main results was a curvilinear relationship in species performance and genetic divergence, i.e. that species perform best in sites of conspecific taxa that share similar ecological preferences, and in sites of distant relatives where there is less competition. She also showed rare species advantage, reproductive character displacement, and fine-scale exclusion within the plant community. The second part of her talk discussed the history of the American Society of Naturalist, a topic covered by the Molecular Ecologist earlier this year.

Tim Birkhead (University of Sheffield) gave the SSE Stephen Jay Gould Prize lecture, entitled “The most perfect thing: the inside (and outside) of a bird’s egg”. This was a fascinating general natural history talk full of wonderful facts about eggs. I had no idea that an hour before most birds lay an egg it rotates 180 degrees and is then laid point first, or that wrynecks are indeterminate egg layers and if you remove eggs as they are produced they will go on to lay 70 eggs. However, Tim’s talk did attract feedback on Twitter for using his lecture to vocally criticise other scientist’s research.

Luke Harmon (University of Idaho) gave the SSB Presidential Address on “Scaling the tree of life”. The talk addressed the broad biological issue of why some parts of the tree of life are so species rich. He also introduced the technical issue of why phylogenetic studies often find high speciation in young clades (Rosenblum et al. 2012). Luke suggested we are in the golden age of time-calibrated phylogenies, where we can now address these types of questions in replicate systems. I’ll save the details here as it’s hard to summarise briefly macroevolutionary pulses and hierarchical models, but it seems clear we need to take a critical view of how we measure speciation rates across trees.

Hopi Hoekstra (Harvard University) gave the SSE Presidential Address entitled “The genetic basis of Behavioural Ecology”. This brilliant talk described the genetic basis of parental behaviour in Peromyscus mice. She showed that differential expression of vasopressin underlies differences in nest-building behaviour (Bendesky et al. 2017), with genetic transformation of key genes changing this behaviour. This talk was one of the best examples of how merging ecological observations, population genomics, and functional genetics can improve our understanding of the evolution of key traits.

Highlights from presentations given in the parallel sessions include:

  • David Marques (University of Bern) used genomic sequencing to trace the origin of Lake Constance sticklebacks. He revealed this population has a complex mosaic genome, and proposed the new term combinatorial speciation to explain the growing number of cases where admixture is involved in the establishment of new taxa.
  • Axel Myer (Universität Konstanz) presented the genetic basis of large-lips in a lake radiation of cichlid fish. Large lips are advantageous as they aid crevice feeding. He showed that a single major QTL underlies this phenotype, and suggested experimental manipulation of lip size (using gelatine!) could be used to test the fitness benefit of this extraordinary phenotype.
  • Lesley Lancaster (University of Aberdeen) showed that the damselfly Ischnura elegans is expanding into colder climates beyond the range predicted by climate change.
  • David Baum (University of Wisconsin–Madison) described how species definitions are ‘artistic’ and there is no single correct species taxonomy. He also introduced exclusivity factors (which build on concordance factors) as a new approach to improve objective species delimitation.
  • Andrew Leitch (Queen Mary University of London) presented ecological evidence that nitrogen and phosphorus are limiting in the wild and that N + P rich environments have species with larger genomes (Guignard et al. 2016). He included a back-of-the-envelope calculation that polyploid genomes must lose around 367bp per generation to explain extant genome sizes, and asked what selective pressures explain this process of diploidization.
  • Greg Owens (The University of British Columbia) showed the importance of knowing the parents in studies of hybrid speciation. A long-term study of artificial hybrids intending to recreate the hybrid sunflower species Helianthus annuus subspecies texanus (with proposed parents H. annuus and H. debilis) revealed selection on particular haplotypes. But sequencing of natural populations revealed other species are more likely to be contributing to the hybrid species than H. debilis.

Common themes from the talks I saw were assortative mating, chromosomal structure and recombination, and balancing selection. I’m expecting these topics to continue to feature heavily as genomic data from more study systems become available.

The conference finished with a wonderful evening at Abbaye de Valmagne, a 13th century abbey surrounded by vineyards.

The stunning Abbaye de Valmagne, Montpellier. Photos: Alex Twyford.

Looking forward to the next Evolution meeting in Providence Rhode Island!

References

Bendesky A, Kwon Y-M, Lassance J-M, et al. (2017) The genetic basis of parental care evolution in monogamous mice. Nature 544, 434.

Guignard MS, Nichols RA, Knell RJ, et al. (2016) Genome size and ploidy influence angiosperm species’ biomass under nitrogen and phosphorus limitation. New phytologist 210, 1195-1206.

Rosenblum EB, Sarver BA, Brown JW, et al. (2012) Goldilocks meets Santa Rosalia: an ephemeral speciation model explains patterns of diversification across time scales. Evolutionary Biology 39, 255-261.

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ESA 2018 Recap

Something old, something new, something borrowed, something BLUE

…in which I shoe-horn a summary post of this giant meeting into a cutesy subtitle, but it mostly works.

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Evolution 2018 Day 1: From genomics in the wild, to new models of selection

It’s Evolution conference time! Evolution has long been my favourite fixture in the conference calendar, with its diverse mix of theoretical and empirical studies that span the full range of evolutionary biology. This year it’s the second Joint Congress on Evolutionary Biology, which brings together the European Society for Evolutionary Biology, the American Society of Naturalists, the Society for the Study of Evolution and the Society of Systematic Biologists all under one roof in the lovely city of Montpellier in the south of France.

The conference kicks off with the ESEB Presidents’ Award delivered by Loeske Kruuk (Australian National University), with a talk entitled ‘Evolutionary dynamics and fitness in wild populations’. Studying quantitative genetics in the wild is challenging because many classical theoretical predictions don’t apply, and because robust inferences require long-term studies that genotype complete populations. Loeske discusses how her work generating a completely pedigree, along with large-scale phenotypic data, for the superb fairy-wren (Malurus cyaneus), has given insights into quantitative genetics in the wild. Interestingly she shows temporal covariance between body size and fitness, but this is because body size is related to other traits, and therefore there is no expectation of body size showing an evolutionary response. She also shows date of moult is heritable, and suggests this means that ‘the early bird gets the girl’. She finishes up by saying that there are less than 10 estimates of fitness for wild populations, and that there are some consistent effects between species (like cohort effects) but lots of variation. I’m really looking forward to seeing the paper where these comparisons of fitness are presented.

The superb fairy wren. Image JJ Harrison/Wikipedia.

Next up is the session ‘Consequences of hybridization: from swamping to speciation’, one of 78 thematic symposia in 13 parallel sessions. The highlight talk for me is Molly Schumer (Harvard University) discussing hybridisation in swordtail fishes. These remarkable fish all demonstrate over 10% hybrid ancestry in their genomes, suggesting a pervasive role of hybridisation in adaptation and speciation. She goes on to discuss how low coverage sequencing and local ancestry analyses reveal the minor parent ancestry being purged over time, as well as assortative mating related to hybrid ancestry. It’s a great demonstration of how fine-scale genomic analyses of independent geographic populations can reveal the repeatability of evolution. Other interesting talks in the session include Mario Vallejo-Marin (University of Stirling) discussing rapid evolution of hybrid monkeyflowers, and Amy Goldberg (Duke University) discussing global ancestry proportions inferred by mechanistic models.

The readily hybridising fish genus Xiphophorus. Image: Wikipedia.

After a lunch, I hop over to the session ‘Towards an integrated understanding of genomic and phenotypic divergence’. Thom Nelson (University of Montana) tell us how comparative genomic analyses of the monkeyflowers Mimulus lewisii and M. cardinalis show substantial genomic rearrangements, but with low divergence (as measured by Fst) and without elevated divergence in translocations and inversion. Konrad Lohse (who has the office next to me at the University of Edinburgh) gives a critical appraisal of verbal models of island of divergence, and discusses recent work to produce better genome scans for divergence. This talk seems particularly relevant given the widespread use of genome scans and the use of arbitrary cut-offs for outliers (something many of us have been guilty of at some point). Finally, David Field (IST Austria) gives an exciting overview of a hybrid zone between snapdragon (Antirrhinum) subspecies with contrasting flower colours. This study is an exceptional case of generating a pedigree of a plant population that incorporates all the complexities that occur in the wild (e.g. most Antirrhinum plants are annuals but 20% are perennial, and seeds can persist in the seedbank). These data reveal strong home-site advantage for the two subspecies, with strong clines for loci underlying flower colour in the hybrid zone.

The first day really shows the breadth of evolutionary research. Evolutionary biologist no longer appear to be getting dazzled by new technology (only one photo of a sequencer today) and genomic sequencing is now just a routine tool for investigating evolutionary questions. It’s also been nice to see such a well-organised conference, with clear cut-offs for talks to prevent people overrunning (using an interesting choice of music!), facilities for childcare, and minimal use of disposable materials.

Please let me your favourite moments of Evolution 2018 via Twitter (@alex_twyford) and I’ll include a selection in a conference round-up post once the conference dinner hangover eases.

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For flexible eDNA analysis, just capture whatever you want

This is a guest post by Taylor Wilcox and Katherine Zarn, whose article “Capture enrichment of aquatic environmental DNA: A first proof of concept” is online ahead of publication at Molecular Ecology Resources. Wilcox and Zarn wanted to elaborate on the usefulness of capture enrichment as an alternative to metabarcoding beyond what they could cover in that paper’s discussion, and this post is the result. — JBY

Environmental DNA sampling for multi-taxa species detection (i.e., the inference of species presence from genetic material in the environment) has been a hot topic lately. Some of the most exciting recent work has used high-throughput sequence (HTS) to simultaneously screen for the presence of large suites of taxa (Valentini et al. 2016), estimate relative species abundances (Ushio et al. 2018), and even make inferences about population structure (Sigsgaard et al. 2016). Most of these studies have relied on metabarcoding, which despite its obvious utility, has some real limitations. One fundamental limitation emerges from a reliance on shared primers for bulk amplification of mixed templates. This tends to generate skewed relative sequence abundances after enrichment and potential loss of species detection (Deiner et al. 2018, Piñol et al. 2018).

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Transcriptome sequencing catches bats’ immune systems napping

A little brown bat (Myotis lucifugans) infected with the white-nose fungus. (Flickr: US Fish and Wildlife Service)

Populations of multiple North American bat species have been more than decimated by white-nose syndrome, a fungal disease that spreads within roosting colonies and becomes deadly during hibernation. A paper just released online early at Molecular Ecology adds support to a hypothesis that the reason for the fungus’s virulence is that hibernation puts bats’ immune systems to sleep — and waking up to fight the fungus costs more than they can afford.

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Just So Stories addendum: How the stickleback keeps getting its stickles

Model organisms have been essential tools for genetics research since the field was formed.  Kelle Freel discussed the characteristics that make for a good model organism in a previous TME post.  Briefly, traits like short generation time, lots of offspring, and easy-to-track developmental stages have been exploited to answer questions about molecular mechanisms behind transmission, cyto-, developmental, population and quantitative genetics and comparative genomics.  A lesser-known model organism is the three spined stickleback, Gasterosteus aculeatus (though it has been mentioned or discussed in several previous TME posts).

Why? Because this fish has the rare honor of being an evolutionary and ecological model organism.  Sticklebacks occur holarctically in marine, estuarine, and freshwater (freshwater) habitats in Europe, Asia, and North America.  During the last retreat of the glaciers in the Pleistocene, this historically marine species began to invade freshwater habitats many times on different timescales in different places. What piqued researchers’ interests about these fish, in the pre-genomic age, was the striking phenotypic variation in various newly-formed freshwater populations and how quickly these divergences happened.  Even though freshwater morphs can differ markedly from each other, even in the same lake, there are certain morphological changes that consistently happen in the evolution from marine/anadromous to freshwater forms, like the loss of body armor.

Fig 1 from Cresko et al 2004. A-C) are freshwater-derived representatives while D) is an anadromous specimen.

This in situ replication lends itself to the study of adaptive radiation and parallel evolution in the wild – a rare treat for evolutionary biologists.  Furthermore, scientists have been able to perform genetic crosses between different morphs and produce viable offspring.  By coupling laboratory crosses between divergent phenotypes with genomic information such as SNP locations across the genome (i.e. QTL mapping) previous studies (reviewed here) managed to pin down regions of the genome likely responsible for a wealth of phenotypic differences observed between environments like body size, feeding traits, and color.

Again, technology lurches forward and in 2010, the first high density SNP-based genome scan (45000 SNPs, 100 fish) of sticklebacks was undertaken.  Applying those observations of intrapopulation diversity and interpopulation divergence patterns to hypotheses of evolution and adaptation the researchers found: 1) the repeated parallel evolution observed in this system relies upon “freshwater haplotypes” that nevertheless persist in oceanic populations at low frequencies, then become frequent in newly minted freshwater populations (aka parallel hard sweeps), as opposed to new mutations cropping up over and over in replicated freshwater populations; 2) both balancing and divergent selection play a role in stickleback evolution; 3) many of the genomic regions previously identified as having a QTL related to divergent phenotypes overlap with regions they found to have high FST values, and that newly-detected regions of the genome also contain patterns of divergence between populations; 4) their results lend more evidence to the biogeographic hypothesis that a large, panmictic oceanic population has repeatedly given rise to divergent freshwater populations.

Fast forward to the July issue of Genetics and the newest installment of RAD-seq/SNP discovery/stickleback research. What novel insights are to be gained? First of all, the authors were able to take advantage of a quite recent colonization event of freshwater habitat via the 1964 Alaska earthquake, which was so powerful that three islands were seismically uplifted, trapping some sticklebacks in newly-formed freshwater ponds.

Secondly, by training a general Hidden Markov model to segment the genome into regions of high and low divergence using genomic data from regions of known high and low FST values from other studies, and defining and classifying haplotypes (~100bp RAD loci, instead of individual SNPs) into different categories and analyzing the pattern between regions of high and low divergence across freshwater and marine populations, they were able to gain insight into mechanisms of evolution absent in other studies.

Fig 2 from Bassham et al 2018. I-XXI are linkage groups of the stickleback genome.  Each concentric circle, numbered outward, is a pairwise comparison of FST’ values.  “OC” indicates a marine population.  The complete lack of high FST’ (red; a measure of haplotype diversity) in ring 1 illustrates the low haplotype divergence between two oceanic populations.  Rings 2-5 are comparisons between a newly-formed freshwater population and 3 pooled marine populations. Note that regions with high FST’ values overlap across comparisons. Rings 4 and 5 represent the youngest of the populations and so have less marine-freshwater divergence.

For example, marine sticklebacks have lower absolute haplotype diversity across divergent regions as opposed to non-divergent regions, which underscores the possibility of strong selection in both regimes, not just freshwater. Furthermore, most haplotypes in majority in freshwater occur in marine fish as well, but are rare. Therefore, greater purifying selection in marine sticklebacks across divergent genomic regions coupled with the fact that haplotypes that are more successful in freshwater habitats persist in marine fish at low frequencies could contribute to the magnitude of the divergence the authors observe as well as the speed with which freshwater morphotypes arise from oceanic stickleback when they colonize a new freshwater habitat. In fact, both population types could be acting as reservoirs of genetic diversity.

Another striking finding was the percentage of the genome deemed to be divergent, nearly 25%, which is a 30-fold increase from previous estimations. However, the authors argue deeper population genomic sampling will detect more changes in allele frequencies across loci as compared to previous methods.

So to reiterate: The remarkable thing about this system is that A) there is a powerful signal of divergent selection, more powerful than previously assumed, even in the presence of gene flow and B) this parallel evolution phenomenon has happened over and over, in the same regions in the genome, in a time period as short as 50 years, though the evolutionary history is much deeper.  Truly, the stickleback is a fascinating and powerful evolutionary and ecological model organism.

References

Bassham, S., Catchen, J., Lescak, E., von Hippel, F. A., Cresko, W. A. (2018) Repeated Selection of Alternatively Adapted Haplotypes Creates Sweeping Genomic Remodeling in Stickleback. GENETICS 209 (3) 921-939

Cresko, W. A., Amores, A., Wilson, C., Murphy, J., Currey, M., Phillips, P., Bell, M. A., Kimmel, C. B., Postlethwait, J. H. (2004) Parallel genetic basis for repeated evolution of armor loss in Alaskan threespine stickleback populations. Proceedings of the National Academy of Sciences, 101 (16) 6050-6055

Hohenlohe PA, Bassham S, Etter PD, Stiffler N, Johnson EA, Cresko WA (2010) Population Genomics of Parallel Adaptation in Threespine Stickleback using Sequenced RAD Tags. PLoS Genet 6(2)

Marques DA, Lucek K, Meier JI, Mwaiko S, Wagner CE, Excoffier L, Seehausen, O. (2016) Genomics of Rapid Incipient Speciation in Sympatric Threespine Stickleback. PLoS Genet 12(2)

Peichel, C.L. and Marques, D. A. Marques (2017) The genetic and molecular architecture of phenotypic diversity in sticklebacks. Philosophical Transactions of the Royal Society B-Biological Sciences 372 (1713)

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Robin Waples awarded the 2018 Molecular Ecology Prize

The 2018 Molecular Ecology prize has been awarded to Robin Waples for his work on conservation biology and management, particularly as the leading expert on approaches for using molecular markers to estimate and understand effective population size in natural populations, including subdivided and continuously distributed populations, and use of time series analyses. His studies of populations with overlapping generations have illuminated the evolution of life-history changes in species that are harvested by humans, and made important contributions to understanding fisheries populations. By adapting population genetic models to real-life situations, including structured populations with gene flow, and developing statistically rigorous analyses, his contributions have significantly advanced both conservation and evolutionary ecology.

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They joy of genome sequencing: when genomics meets natural history

When I have a massive pile of papers that I need to read, I can’t help but look at the ones with interesting natural history first. There’s something exceptionally satisfying about using modern tools to dig deeper into the features that make each species so interesting. Many molecular ecologists, myself included, started a career in biology because of a love of natural history, and I think it’s great when this passion can be captured in modern research. One area where an understanding of an organism’s natural history is perhaps surprisingly important is in whole genome sequencing. While it’s becoming increasingly common to sequence, assemble and annotate genomes (though I’d still argue it’s challenging to do it well), many papers do more than just generate a genomic resource, and relate genomic variants to unique properties of a species. Even better is when these interesting features of a species can be related to other experimental work, or sequencing additional natural populations, to gain deeper insights into organismal biology.

The pitcher plant Cephalotus follicularis (picture H. Zell for Wikipedia)

One example is the paper that describes the genome of the carnivorous pitcher plant Cephalotus (Fukushima et al. 2017). Carnivorous plants have evolved on multiple occasions and use a sophisticated range of traps to attract, catch and digest prey in nutrient-poor environments. The highlight of the paper is that the authors manage to induce the switch between pitcher trap and flat leaf by changing ambient temperature, and then use transcriptomic comparisons to pinpoint differentially expressed genes involved in leaf development. This neat trick reveals a number of candidate genes involved in pitcher formation, including AS2, YAB5, and WOX1 orthologues. They also looked at the digestive proteins in Cephalotus and other taxa representing independent origins of carnivory, and find shared proteins that are repeatedly co-opted in the evolution of this novel phenotype. I came away from reading the paper being even more intrigued about how these amazing plants have evolved, and thinking that there’s so much we can now address with careful genomic and transcriptomic analyses.

And who can forget the passenger pigeon genome papers? The passenger pigeon (Ectopistes migratorius) was at one point perhaps the most abundant bird species on earth, with a census population size in the billions, before its unprecedented extinction in the 19th century due to over-hunting. Two papers, one lead by Chih-Ming Hung published in PNAS, and one by Gemma Murray published in Nature, produced genome data from museum specimens and from related extant pigeon species. Both studies showed surprisingly low genomic diversity indicative of a low effective population size. They differ in what they think caused this low diversity, with Hung et al. (2014) pointing to population fluctuations, and Murray et al. (2017) inferring pervasive natural selection. I haven’t looked at the details to decide which is more likely to be correct, but either way, this is a remarkable example of low diversity in a high abundance species, and a great use of museum sequencing.

Genomes come in all sizes and levels of complexity, and I think there’s something to learn from sequencing them all. From tiny desiccation-tolerant bdelloid rotifers (Nowell et al. 2018) and wonderful water bears (Koutsovoulos et al. 2016), to enigmatic Gnetum (one to look up, plant fans, see Wan et al. 2018) and giant lilly genomes (Kelly et al. 2015), each with their own fascinating biology. The next time whole genome sequencing fatigue sets in as you see the boiler plate title “the genome of XXX reveals YYY”, remember that genome sequencing is a superb tool to make amazing discoveries about the natural world.

References

Fukushima K, Fang X, Alvarez-Ponce D, et al. (2017) Genome of the pitcher plant Cephalotus reveals genetic changes associated with carnivory. Nature ecology & evolution 1, 0059.

Hung C-M, Shaner P-JL, Zink RM, et al. (2014) Drastic population fluctuations explain the rapid extinction of the passenger pigeon. Proceedings of the National Academy of Sciences 111, 10636-10641.

Kelly LJ, Renny‐Byfield S, Pellicer J, et al. (2015) Analysis of the giant genomes of Fritillaria (Liliaceae) indicates that a lack of DNA removal characterizes extreme expansions in genome size. New Phytologist 208, 596-607.

Koutsovoulos G, Kumar S, Laetsch DR, et al. (2016) No evidence for extensive horizontal gene transfer in the genome of the tardigrade Hypsibius dujardini. Proceedings of the National Academy of Sciences, 201600338.

Murray GG, Soares AE, Novak BJ, et al. (2017) Natural selection shaped the rise and fall of passenger pigeon genomic diversity. Science 358, 951-954.

Nowell RW, Almeida P, Wilson CG, et al. (2018) Comparative genomics of bdelloid rotifers: Insights from desiccating and nondesiccating species. PLoS biology 16, e2004830.

Wan T, Liu Z-M, Li L-F, et al. (2018) A genome for gnetophytes and early evolution of seed plants. Nature Plants 4, 82.

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