Speciation in three-spined stickleback

Speciation, the division of one species into two, has provided evolutionary biologists with a rich ensemble of questions, conundrums and revelations for over a century, and yet our understanding of many of the factors affecting this complex, multidimensional process remains limited. In this thesis, I aimed to further our understanding of speciation using divergent populations of three-spined stickleback (Gasterosteus aculeatus) on the island of North Uist, Scottish Western Isles. Firstly, I explored the degree of morphological and genetic separation between three stickleback ecotypes, showing that both strongly reproductively isolated, and admixed populations exist in close proximity. I then attempted to identify the ecological and genetic origins of strongly isolated species-pairs, testing two competing explanations for their existence. I showed that a recent ‘double-invasion’ is unlikely, but found stark differences in the long-term genetic history between ecotypes, indicating that the evolution of species-pairs may be related to secondary contact between anciently diverged mitochondrial lineages. I then conducted mate choice trials to assess mating preferences between ecotypes, and to test for reinforcement in species-pairs. Consistent with the idea that speciation in this case is not driven purely by ecological factors, I found no evidence that reinforcement drives assortative mating in species-pairs. Rather, it appears that extant mating preferences have developed as a by-product of other adaptations. Finally, I took a brief interlude to document and investigate an exciting chance finding, internally developed embryos retained within the ovaries of a normally oviparous species, before concluding by summing up my findings, their relevance for scientific progress, and avenues opened up for further research

The Variable Stars and Blue Horizontal Branch of the Metal-Rich Globular Cluster NGC 6441

We present time-series VI photometry of the metal-rich ([Fe/H] = -0.53) globular cluster NGC 6441. Our color-magnitude diagram shows that the extended blue horizontal branch seen in Hubble Space Telescope data exists in the outermost reaches of the cluster. The red clump slopes nearly parallel to the reddening vector. A component of this slope is due to differential reddening, but part is intrinsic. The blue horizontal branch stars are more centrally concentrated than the red clump stars. We have discovered about 50 new variable stars near NGC 6441, among them eight or more RR Lyrae stars which are very probably cluster members. Comprehensive period searches over the range 0.2-1.0 days yielded unusually long periods (0.5-0.9 days) for the fundamental pulsators compared with field RR Lyrae of the same metallicity. Three similar long-period RR Lyrae are known in other metal-rich globulars. With over ten examples in hand, it seems that a distinct sub-class of RR Lyrae is emerging. The observed properties of the horizontal branch stars are in reasonable agreement with recent models which invoke deep mixing to enhance the atmospheric helium abundance, while they conflict with models which assume high initial helium abundance. The light curves of the c-type RR Lyrae seem to have unusually long rise times and sharp minima. Reproducing these light curves in stellar pulsation models may provide another means of constraining the physical variables responsible for the anomalous blue horizontal branch extension and sloped red clump observed in NGC 6441.Comment: 30 pages plus 6 EPS and 6 JPEG figures; uses AAS TeX. Accepted by the Astronomical Journal. Minor changes include computing He abundance, modifications to Figs 1 and 8, and expansion on idea that blue HB stars may be produced in binarie

Trends in the Globular Cluster Luminosity Function of Early-Type Galaxies

We present results from a study of the globular cluster luminosity function (GCLF) in a sample of 89 early-type galaxies observed as part of the ACS Virgo Cluster Survey. Using a Gaussian parametrization of the GCLF, we find a highly significant correlation between the GCLF dispersion, sigma, and the galaxy luminosity, M_B, in the sense that the GC systems in fainter galaxies have narrower luminosity functions. The GCLF dispersions in the Milky Way and M31 are fully consistent with this trend, implying that the correlation between sigma and galaxy luminosity is more fundamental than older suggestions that GCLF shape is a function of galaxy Hubble type. We show that the sigma - M_B relation results from a bonafide narrowing of the distribution of (logarithmic) cluster masses in fainter galaxies. We further show that this behavior is mirrored by a steepening of the GC mass function for relatively high masses, M >~ 3 x 10^5 M_sun, a mass regime in which the shape of the GCLF is not strongly affected by dynamical evolution over a Hubble time. We argue that this trend arises from variations in initial conditions and requires explanation by theories of cluster formation. Finally, we confirm that in bright galaxies, the GCLF "turns over" at the canonical mass scale of M_TO ~ 2 x 10^5 M_sun. However, we find that M_TO scatters to lower values (~1-2 x 10^5 M_sun) in galaxies fainter than M_B >~ -18.5, an important consideration if the GCLF is to be used as a distance indicator for dwarf ellipticals.Comment: 4 pages, 3 figures. Accepted for publication in ApJ Letters. Also available at http://www.cadc.hia.nrc.gc.ca/community/ACSVCS/publications.htm

The PCP genes Celsr1 and Vangl2 are required for normal lung branching morphogenesis

The lungs are generated by branching morphogenesis as a result of reciprocal signalling interactions between the epithelium and mesenchyme during development. Mutations that disrupt formation of either the correct number or shape of epithelial branches affect lung function. This, in turn, can lead to congenital abnormalities such as cystadenomatoid malformations, pulmonary hypertension or lung hypoplasia. Defects in lung architecture are also associated with adult lung disease, particularly in cases of idiopathic lung fibrosis. Identifying the signalling pathways which drive epithelial tube formation will likely shed light on both congenital and adult lung disease. Here we show that mutations in the planar cell polarity (PCP) genes Celsr1 and Vangl2 lead to disrupted lung development and defects in lung architecture. Lungs from Celsr1Crsh and Vangl2Lp mouse mutants are small and misshapen with fewer branches, and by late gestation exhibit thickened interstitial mesenchyme and defective saccular formation. We observe a recapitulation of these branching defects following inhibition of Rho kinase, an important downstream effector of the PCP signalling pathway. Moreover, epithelial integrity is disrupted, cytoskeletal remodelling perturbed and mutant endoderm does not branch normally in response to the chemoattractant FGF10. We further show that Celsr1 and Vangl2 proteins are present in restricted spatial domains within lung epithelium. Our data show that the PCP genes Celsr1 and Vangl2 are required for foetal lung development thereby revealing a novel signalling pathway critical for this process that will enhance our understanding of congenital and adult lung diseases and may in future lead to novel therapeutic strategies

Courtship behavior, nesting microhabitat, and assortative mating in sympatric stickleback species pairs

© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. The maintenance of reproductive isolation in the face of gene flow is a particularly contentious topic, but differences in reproductive behavior may provide the key to explaining this phenomenon. However, we do not yet fully understand how behavior contributes to maintaining species boundaries. How important are behavioral differences during reproduction? To what extent does assortative mating maintain reproductive isolation in recently diverged populations and how important are “magic traits”? Assortative mating can arise as a by-product of accumulated differences between divergent populations as well as an adaptive response to contact between those populations, but this is often overlooked. Here we address these questions using recently described species pairs of three-spined stickleback (Gasterosteus aculeatus), from two separate locations and a phenotypically intermediate allopatric population on the island of North Uist, Scottish Western Isles. We identified stark differences in the preferred nesting substrate and courtship behavior of species pair males. We showed that all males selectively court females of their own ecotype and all females prefer males of the same ecotype, regardless of whether they are from species pairs or allopatric populations. We also showed that mate choice does not appear to be driven by body size differences (a potential “magic trait”). By explicitly comparing the strength of these mating preferences between species pairs and single-ecotype locations, we were able to show that present levels of assortative mating due to direct mate choice are likely a by-product of other adaptations between ecotypes, and not subject to obvious selection in species pairs. Our results suggest that ecological divergence in mating characteristics, particularly nesting microhabitat may be more important than direct mate choice in maintaining reproductive isolation in stickleback species pairs

Best Practices for Archival Term Positions

This document presents best practices for administrators, hiring managers, and supervisors to put into action when planning for and employing archival term positions. At the foundation of this document is the recognition that temporary labor is detrimental to employees, as well as to sustainable and holistic collection stewardship. This document is on a 5 year review cycle; it will be reviewed for potential updates in 2027

The Hubble Space Telescope Extragalactic Distance Scale Key Project XXIII. The Discovery of Cepheids In NGC 3319

The distance to NGC 3319 has been determined from Cepheid variable stars as part of the Hubble Space Telescope Key Project on the Extragalactic Distance Scale. Thirteen and four epochs of observations, using filters F555W (V) and F814W (I) respectively, were made with the Wide Field Planetary Camera 2. Thirty-three Cepheid variables between periods of 8 and 47 days were discovered. Adopting a Large Magellanic Cloud distance modulus of 18.50 +- 0.10 mag and extinction of E(V-I)=0.13 mag, a true reddening-corrected distance modulus (based on an analysis employing the ALLFRAME software package) of 30.78 +- 0.14 (random) +- 0.10 (systematic) mag and the extinction of E(V-I) = 0.06 mag were determined for NGC 3319. This galaxy is the last galaxy observed for the HST H0 Key Project.Comment: 22 pages. A gzipped tar file containing 16 figures can be obtained from http://www.ipac.caltech.edu/H0kp/n3319/n3319.htm

Reproductive isolation in a three‐way contact zone

Contact zones between divergent forms within a species provide insight into the role of gene flow in adaptation and speciation. Previous work has focused on contact zones involving only two divergent forms, but in nature, many more than two populations may overlap simultaneously and experience gene flow. Patterns of introgression in wild populations are, therefore, likely much more complicated than is often assumed. We begin to address this gap in current knowledge by investigating patterns of divergence and introgression across a complex natural contact zone. We use phenotypic and genomic data to confirm the existence of a three-way contact zone among divergent freshwater resident, saltwater resident and saltwater migratory three-spined stickleback (Gasterosteus aculeatus) on the island of North Uist, Scottish Western Isles. We find evidence for hybridization, mostly between saltwater resident and saltwater migratory forms. Despite hybridization, genomic analyses reveal pairwise islands of divergence between all forms that are maintained across the contact zone. Genomic cline analyses also provide evidence for selection and/or hybrid incompatibilities in divergent regions. Divergent genomic regions occur across multiple chromosomes and involve many known adaptive loci and several chromosomal inversions. We also identify distinct immune gene expression profiles between forms, but no evidence for transgressive expression in hybrids. Our results suggest that reproductive isolation is maintained in this three-way contact zone, despite some hybridization, and that reduced recombination in chromosomal inversions may play an important role in maintaining this isolation

On the origins of phenotypic parallelism in benthic and limnetic stickleback

Rapid evolution of similar phenotypes in similar environments, giving rise to in situ parallel adaptation, is an important hallmark of ecological speciation. However, what appears to be in situ adaptation can also arise by dispersal of divergent lineages from elsewhere. We test whether two contrasting phenotypes repeatedly evolved in parallel, or have a single origin, in an archetypal example of ecological adaptive radiation: benthic–limnetic three-spined stickleback (Gasterosteus aculeatus) across species pair and solitary lakes in British Columbia. We identify two genomic clusters across freshwater populations, which differ in benthic–limnetic divergent phenotypic traits and separate benthic from limnetic individuals in species pair lakes. Phylogenetic reconstruction and niche evolution modeling both suggest a single evolutionary origin for each of these clusters. We detected strong phylogenetic signal in benthic–limnetic divergent traits, suggesting that they are ancestrally retained. Accounting for ancestral state retention, we identify local adaptation of body armor due to the presence of an intraguild predator, the sculpin (Cottus asper), and environmental effects of lake depth and pH on body size. Taken together, our results imply a predominant role for retention of ancestral characteristics in driving trait distribution, with further selection imposed on some traits by environmental factors