Welcome to the Ragland Lab at the University of Colorado, Denver. We are interested in the process of local adaptation, particularly in response to seasonal variability. Our goal is to understand the mechanisms and targets of natural selection, integrating studies from genomes, to physiology, to whole organism performance. You can read more about our research here. The lab is actively recruiting one PhD student (starting fall 2022). Please direct inquiries to Greg Ragland.

Lab News and Photos

10/31/2021 — Related to below, the lab is recruiting a new PhD student for Fall 2022.  I am happy to field informal inquiries.

10/312021 – Lots of change in lab personnel over the last few months. We are joined by postdoc Jimmy DeMayo, postdoc Shelly McCain, MS student Colin McSweeney, and undergraduate Eureca fellow Jenna Tomkinson. Will be updating the ‘people’ page soon.

4/8/2021 — The lab is now funded by a new NSF CAREER grant! We are extremely grateful for the funding, which will allow us to continue new studies on thermal plasticity of transcriptional dynamics, and more generally the evolution of plasticity in contrasting temperate and tropical environments. We will also be developing a new Course Based Undergraduate Research Experience based on the research specifically aimed and increasing participation and inclusion in STEM at CU Denver.

4/8/2021 — We collaborated on a new study led by Katherine Innskeep, Meredith Doellman, and Jeff Feder with a surprising result; magic traits that promote speciation via niche partitioning can also promote hybridization when the niche spaces begin to fill up! The paper is published in Molecular Ecology.

2/12/2021 — A paper led by former postdoc Jantina Toxopeus is now published in Comparative Biochemistry and Physiology A. The paper explores the consequences of not entering diapause prior to winter. It turns out that Rhagoletis flies can plastically depress metabolism quite a lot even when they are not in diapause, but the plastic response takes so long that they incur a significant metabolic cost, even at low temperatures. This is part of a special issue edited by Nick Teets and Scott Hayward: Combatting the cold: Comparative physiology of low temperature and related stressors in arthropods.

9/14/2020 — Our paper describing the role of general developmental processes in life history evolution has now been published in PNAS. Excited to see this finally out, it’s been many years in the making and has been a fun collaboration with former postdocs, students, and colleagues.

8/1/2020 — Joseph Tucker has joined our lab to complete a thesis and MS degree in Integrated Sciences. Joe will be working on thermal plasticity of transcriptional dynamics.

6/15/2020 — Dr. Jantina Toxopeus finished up in our lab this month and moved on to her new position as Assistant Professor in the Biology Department at St. Francis Xavier University. We look forward to seeing your new lab take shape!

6/2/2020 — Jantina won the Best Presentation Award for her talk in the Postdoctoral Association Seminar Series (2019-2020) at CU Denver/Anschutz.  Congrats!

4/15/2020 — Marianne Davenport successfully defended her Masters Thesis, and won the College of Liberal Arts and Sciences outstanding graduate student award! We look forward to continue working with Marianne in her current position with the US Forest Service.

2/14/2020 — Greg served as the opponent for Olle Lindestad’s doctoral dissertation defense at Stockholm University. The Swedes do dissertations (and after parties) right. Congratulations to Olle! Had a great visit with the Lep eco/evo group there, stood on hallowed ground for ecophysiologists in the Arrhenius lab, and found some window frosting that improbably appealed simultaneously to my love of bugs and surfing…


1/17/2020 — We welcomed in the new semester with a lab trip to the Butterfly Pavilion (with our friends in the Swallow lab)!

Back row: Lahari (M.S. student), Gabrielle (Swallow lab, research assistant), Greg (PI), Isaiah (B.S. student), Matin (B.S. student); Front row: Jantina (postdoc), Emily (Swallow lab, M.S. student), Lalitya (B.S. student), Manaal (B.S. student)

1/8/2020 — Greg Ragland and Jantina Toxopeus presented their recent research on Rhagoletis pomonella diapause at the Society for Integrative and Comparative Biology Annual Meeting in Austin TX this week. Jantina’s travel was funded by a Postdoctoral Association travel award from CU Denver.

12/16/2019 — We closed out the semester with a delicious potluck lunch (feat. so many cookies!) before everyone starts a well-deserved break!

10/17/2019 — Congratulations to Joseph Tucker, who has won a mini-UROP (Undegraduate Research Opportunity Program) grant to conduct some gene expression work with Drosophila melanogaster next semester!

9/28/2019 — Great lab hike today in Rocky Mountain National Park, with our friends in the Swallow lab!

Back row: Marianne (M.S. student), Lalitya (B.S. student); Middle row: Jantina (postdoc), Emily (Swallow lab, M.S. student), Isaiah (B.S. student), Matin (B.S. student), Manaal (B.S. student); Front row: Gabrielle (Swallow lab, research assistant), Graem (honorary entomologist for the day)

9/3/2019 — Had a good lab social at Atomic Cowboy (pizza AND trivia) to welcome new and returning lab members.

Clockwise, starting at bottom left: Greg (PI), Joseph (B.S. student), Lalitya (B.S. student), Jantina (postdoc), Isaiah (B.S. student), Ahn (honorary trivia team member for the day), Lahari (M.S. student), Matin (B.S. student), Marianne (M.S. student)

8/14/2019 — Welcome to our new Master’s student Lahari Gadey, who is excited to work on all sorts of fly research.

7/29/2019 — Greg Ragland and Jantina Toxopeus are presenting their recent research on Drosophila melanogaster this week at the International Symposium on the Environmental Physiology of Ectotherms and Plants – in Buenos Aires, Argentina! Congratulations to Jantina for receiving a Sable Systems Young Scientist Award to support her travel to the meeting.

7/1/2019 — Congratulations to our postdoc Jantina Toxopeus, who has accepted a tenure track position beginning next summer at St. Francis Xavier University in her homeland of Canada!

6/21/2019 – 6/25/2019 — Greg Ragland and Mac Calvert each presented their research on the predictability of evolution at the Evolution meeting in Providence, RI. You can see a video of Greg’s talk here.

6/17/2019 — Welcome to our newest lab member, Isaiah Sower, who joins our lab as an undergraduate volunteer helping Marianne with her work on beetle pests.

6/3/2019 — Welcome to our newest lab member, Joseph Tucker, who joins our lab as an undergraduate volunteer helping Jantina with some RNAi work in Drosophila melanogaster.

5/13/2019 — Jantina Toxopeus is off to the Canadian Society of Zoologists (CSZ) annual meeting this week to present some of our latest work on Rhagoletis pomonella overwintering biology. Her travel is partially funded by a CSZ travel award for postdoctoral fellows!

5/11/2019 — We had an excellent get-together (featuring homemade tacos) to celebrate the many Ragland lab successes this semester, but neglected to collect any photographic evidence!

4/30/2019 — Welcome back to Marianne Davenport, who spent last week in Alaska sharing her work on spruce beetle diapause at the Western Forest Insect Work Conference in Alaska.

4/26/2019 — Congratulations to Mac Calvert who, along with defending his thesis this week, has won the Integrative Biology department’s award for Outstanding Graduate Student.

4/26/2019 — Excellent work by all the Ragland lab students that presented their research projects today at CU Denver’s RaCAS (Research and Creative Activities Symposium). Manaal Dawaldi won 2nd place in the poster competition!

Lalityla Andaloori presenting her UROP project on fly metabolism
Manaal Dawaldi presenting the work he’s been doing with Marianne Davenport on beetle diapause
Erina Rader presenting her UROP and EUReCA!-funded work on apple maggot fly genetics

4/24/2019 — M.S. student Mac Calvert defended his thesis today, and passed with flying colors. He will be starting a Ph.D. in the fall at the University of Pittsburgh. Congratulations, Mac!

Greg (left) and Mac (right) with big smiles post-thesis defense

4/13/2019 — Had a visit from Brent Sinclair this week – he gave a great seminar on the role of ‘omics in physiology, and had time for some skiing too.

Brent Sinclair (left) and Greg (right) at the “faculty club”

4/8/2019 — Congratulations to Matin Sanaei, who has been awarded UROP (undergraduate research opportunity program) funding to do an independent research project in the Ragland lab during his sophomore year!

3/31/2019 — The lab got out to enjoy the spring weather this weekend on a hike through Reynolds Park.

At the top of the Reynolds Park hike, from left to right: Manaal (B.S. student), Lalitya (B.S. student), Dana (honorary lab member for the day), Mac (M.S. student), Greg (chief form signer), Marianne (M.S. student), Jantina (post-doc)

1/22/2019 — Our article on the genomics of host race formation in Rhagoletis pomonella led by Meredith Doellman in Jeff Feder’s lab is now published in Ecology and Evolution.

12/10/2018 — Marianne Davenport passed her M.S. proposal with flying colors!

11/6/2018 — Phil Freda successfully completed his Ph.D. defense today. Well done, Phil!

9/21/2018 — Jantina Toxopeus has defended her Ph.D. dissertation with Brent Sinclair, and will be joining the lab as a postdoc in a few weeks. Welcome, Jantina!

9/21/2018 — Our paper on sixtoothed spruce bark beetle phylogeography led by Martin Schebeck in Christian Stauffer’s lab is now published.

8/28/2018 — Erina Rader has been awarded a EUReCA! fellowship to support a new project on rose fly population structure.

8/23/2018 — Mac Calvert presented his M.S. research on genetic modularity of seasonal adaptation in an invited symposium at the Joint Congress on Evolutionary Biology in Montpellier, France.

5/18/2018 — Our paper detailing how geographic, genetic variation contributes to local adaptation is now published in Genes.

5/8/18 — Lalitya Andaloori and Erina Rader have both won CU Denver UROP (undergraduate research opportunity program) awards to support their research projects. Congrats!


8/2/2017 — Start of a spruce beetle gallery. This happens when the female bores into the tree and then she will lay eggs on either side of the gallery. Photo taken by Marianne at Guanella Pass, Colorado.


6/7/2017 — Photo from Marianne of fir engraver bark beetle gallery under the bark. This was taken near Ouray, Colorado.

6/1/2017 — Marianne Davenport has taken a summer position with the forest service to survey bark beetle infestations in Colorado, and will join the lab to pursue a M.S. in the fall. She will focus on seasonality and population dynamics of forest pest insects.


5/24/2017 — Enjoying the food truck-a-palooza at Civic Center Eats

5/13/2017 — Our new Molecular Ecology paper studying genetic modularity during rapid adaptations has two important implications. First, it suggests that genetic independence among phenotypes may facilitate the process of speciation with gene flow. Second, it suggests that multiple diapause components, such as initiation and termination, may evolve independently.


4/22/2017 — March for Science, CO State Capital. A great turnout from CU Denver and in general, with a very positive vibe.

4/17/2017 — Our paper examining diapause transcriptomics across insect species is now available on early view, to be published in a special issue of Physiological Entomology associated with the ICE 2016 symposium on Photoperiodic induction of diapause and seasonal morphs. The results support multiple evolutionary origins of insect diapause, but also reveal a strong signal of convergence on similar patterns of gene expression during diapause across Insecta.

3/31/2017 — The Entomological Society of America meetings will be in Denver for 2017, and we are hosting a symposium, ‘Genomics of Adaptation: Linking the Next Generation of Genome-Wide Analysis to Understand and Manage Complex Traits’, co-organized by Greg Ragland, Glen Hood, Scott Egan, Dan Hahn, and Meredith Doellman. We’ve got a fantastic group of speakers, schedule should be available in the summer.

1/15/2017 — The symposium ‘Evolutionary Impacts of Seasonality‘ at SICB 2017, co-organized by Caroline Williams and Greg Ragland, was great fun and a fantastic opportunity to hear about and discuss the latest and greatest research with the community. Stay tuned for a companion special issue in Integrative and Comparative Biology…

12/15/16 — McCall Calvert (Mac) joins the lab to pursue an MS. He will be working on genomic differentiation and speciation in flies and parasitoid wasps


9/18/16 — Field site for Rose Hip fly (R. basiola) collections, near Mt. Evans, CO.


9/7/2016 — Easy field day in the park. Haw collections, Cheeseman Park.


7/20/16 — The Ragland lab will be re-locating to the University of Colorado, Denver, Department of Integrative Biology, starting 15 August 2016. Dr. Eddy Dowle will be moving to Denver as well, while Phil Freda will remain at KSU to complete his PhD. Photo: lab farewell BBQ, Manhattan, KS.

7/1/16 — Our NSF Dimensions of Biodiversity proposal with Jeff Feder, Dan Hahn, and Tom Powell has been recommended for funding! This project will examine how seasonal adaptations to changing environments affect co-evolution with parasitoid communities, following up on Glen Hood’s and Adrew Forbe’s excellent work on cascading speciation.

7/1/16 — Our Journal of Experimental Biology paper using RNAseq to test for adaptive divergence in regulation of seasonal timing during winter is now available online. We also found an interesting pattern consistent with active suppression of growth/development during exposure to elevated temperatures.

8/20/15 — Greg presented a comparative transcriptomic study of general stress responses at the ISEPEP meeting in Aarhus, Denmark. Great to catch up with the comparative physiology crowd, very fun meeting with a good mix of sub-organismal physiology, physiological ecology, and ecological genomics.

8/5/15 — Christian Stauffer arrived in Manhattan to start his Fulbright Fellowship. Excited to have him at KSU. Should be a productive semester for our Ips typograpus diapause project.

8/1/15 — Full NSF proposal on acclimation in eastern Newts submitted in collaboration with Nancy Berner at the University of the South and Berea College. Fingers crossed!

6/19/15 — Eddy, Phil, Adam, and Greg presented research at the Arthropod Genomics Symposium in Manhattan, Ks. Nice having a great international crowd just steps from your own front door.

6/15/15 — Our Ecology Letters paper on natural selection and genomic divergence in Rhagoletis flies has now been published (open access). Seems to be getting some good press coverage!

5/14/15 — Our study of transcriptomic responses to host shifts has been published in Molecular Ecology. Cool result suggesting that early in a host shift selection may act strongly against non-adaptive plasticity.

2/15/15 — Phil wins the Reginald Painter memorial scholarship through KSU entomology. Well done, Phil!

5/8/2014 — Christian Stauffer’s Austrian FWF proposal to study the genetics of obligate vs. facultative diapause in spruce bark beetles has been funded! We are collaborators on this work, and very excited to get the project up and running.