• Evolution of ecological novelty
    Why does scale-eating rarely evolve?
  • Sympatric speciation in volcanic crater lakes
    Cast-netting for endemic cichlids in Lake Kotto, Cameroon
  • Measuring the adaptive landscape
    "When biologists think about the evolution of life, they think about climbing mountains." -Carl Zimmer
  • Developmental genetics
    Identifying novel craniofacial genes using in situ hybridization
  • Evolutionary diversification rates
    Morphogram illustrating trait diversification rates in San Salvador Island pupfishes
  • About the lab

    Welcome to the Martin fish speciation lab! We are evolutionary biologists broadly interested in the ecology, evolution, and genomics of adaptive radiation in fishes. We use field experiments, natural history, population genomics, behavioral ecology, functional morphology, quantitative and functional genetics, and phylogenetic methods to dissect this process at the mesoevolutionary scale: rapid radiations of three or more species.

    We are primarily developing two tropical field systems for studying the origins of adaptive radiation: 1) Caribbean microendemic radiations of trophic specialist pupfishes and 2) Cameroon crater lake cichlids, famous as putative examples of sympatric speciation.

    Common themes in our work include 1) the genetic, behavioral, and ecological origins of adaptive radiation in space and time, 2) major features of the fitness landscape driving this process, and 3) the evolution of novel phenotypes and ecological niches.

    If you are intrigued by the speciation process, drawn to tropical field systems, and you have a particular thing about fish, I strongly encourage you to contact me! I am currently accepting PhD students through Integrative Biology at the University of California, Berkeley. I especially encourage applications from BIPOC. Please see our lab's statement of support for the Black Lives Matter movement below and our code of conduct.

    The IB graduate program is described here (annual admissions deadline is December 3rd). For those considering a PhD, I would suggest reading the perspective of my undergraduate advisor here. I am the first Curator of Fishes at Cal and our lab is based inside the Museum of Vertebrate Zoology with extensive shared spaces for molecular genetics work, student offices, specimen preparation, data analysis, and the growing MVZ Ichthyology collection. We also maintain our fish colonies (over 1000 individuals in over 40 species) in four fish rooms plus roof-top experimental ponds.

    I am always open to sponsoring external and internal postdoctoral fellowship applicants, such as the Miller Research Fellowship, UC Mexus, or the Chancellor's fellowship. I am happy to discuss proposal ideas from prospective applicants.

    Martin Lab anti-racism statement

    The murders of Breonna Taylor, George Floyd, Ahmaud Arbery, Elijah McClain, Rayshard Brooks, Freddie Gray, Trayvon Martin, and so many more by our police state deeply saddens and enrages us. These ongoing killings show how deeply white supremacy has rooted itself in our country. We believe that Black Lives Matter and we stand in solidarity with our Black colleagues, students, and protesters. We are committed to anti-racism and increasing diversity and inclusion in our lab group, mentorship, research, teaching, and academic spaces overall. Although we are taking incremental steps toward becoming more anti-racist and continuing to educate ourselves through regular readings and discussion, we also recognize that white supremacy in STEM, particularly acute in our field of Evolution and Ecology and our department, is a major failing of our scientific community and is an ongoing crisis for science. Centuries of systemic racism in science and academia has left us with a largely homogeneous faculty of white men unequipped to take on our most pressing national and global crises in this century. We also recognize that our own research interests and understanding of nature have been skewed by this history of racism and the genocide of Indigenous communities and their traditional knowledge. The scientific contributions of BIPOC have been and continue to be ignored, stolen, and erased.

    We strongly believe that dismantling white supremacy is the only moral choice. This promotes innovation, accelerates discovery and scientific excellence by increasing equitable access to this profession that we love, and increases our ability to communicate science and serve as role models to the local, national, and international communities we serve. We are stronger together. We consider any form of support for white supremacy to be scientific misconduct.

    Lab Members

    Christopher Martin

    Principal Investigator

    Curriculum Vitae



    Recorded talks: Evolution 2014, 2016, 2017, 2019

    UC Berkeley 2015, 2018

    Emilie Richards

    Graduate Student 2016-

    I am broadly interested in how adaptation and speciation processes interact on a genomic level during adaptive radiations and how we harness the powerful resource of genomic data to address questions about evolutionary history. I am currently working on identifying the sources of genetic variation that fueled adaptive diversification in a radiation of trophic specialist pupfishes on a single island in the Bahamas. When I am not busy typing away at my computer, my hobbies include recreational scuba diving and watching nature documentaries with my cat. website email

    Chapter 1: Adaptive introgression into a microendmic radiation (PLOS Gen 2017)

    Chapter 2: Introgression in Barombi Mbo cichlids (Evo Letters 2018)

    Chapter 3: Searching for sympatric speciation in the genomic era (BioEssays 2019)

    Chapter 4: A vertebrate adaptive radiation is assembled from an ancient and disjunct spatiotemporal landscape

    Michelle St. John

    Graduate Student 2017-

    I am interested in investigating the speciation process through the lens of fish behavior. This involves studying behavioral differences between diverging species, exploring interactions between species, and ultimately determining how these differences and interactions affect the evolution of reproductive isolation. When I am not observing fish, I enjoy cooking with my husband and hanging out with my two cats (Tulip and Alfalfa). website email

    Chapter 1: The aggressive behavioral origins of scale-eating (BEH 2019)

    Chapter 2: Adaptive scale-eating feeding kinematics (JEB 2020)

    Chapter 3: The novel behavior of oral-shelling (JFB 2020)


    David Tian

    Graduate Student 2019-

    By combining population genetics, functional genetics, and fitness experiments, I aim to gain a comprehensive and mechanistic understanding of adaptation. Specifically, I am interested in how genetic variation underlying adaptive traits is shaped across time and space in populations and how these loci interact with each other at a molecular level to produce phenotypes. To this end, I am focused on investigating how and why a rare scale-eating trophic specialist evolved its novel and extreme craniofacial morphology. website email


    Austin Patton

    Postdoctoral Scholar 2020-present

    I am broadly interested in hybridization and diversification, with a particular focus on adaptive radiation. My research bridges the macroevolutionary and microevolutionary scales through the use of comparative phylogenetic and genomic approaches. My past and current work includes macroevolutionary studies of salamanders and Anolis lizards, as well as more microevolutionary studies of Tasmanian devils and their transmissible cancer. While in the Martin lab, I will be working to understand the underlying genomic basis of fitness differences in San Salvador Island pupfishes. My dog and partner in crime, Bailey, is probably my biggest fan, and for that I am grateful. website email


    Maria Fernanda (Feña) Palominos

    Postdoctoral Scholar 2021-present

    As a developmental neurobiologist, I am broadly interested in the genetic basis of vertebrate sensory systems evolution. In the Martin lab, I will use my expertise in developmental neurogenetics and microscopy to understand how the genetic variation observed in San Salvador's Cyprinodon pupfishes creates distinctive craniofacial morphologies. When not in the lab, I can probably be found writing, reading, biking, walking, or just looking into nature. email

    Group Photos

    LAB MEETING (2020)

    Fish collecting trip to Albany Marine Park, CA (2019)

    FISH FACES (2019)

    How to transport 1,000+ fishes +1 pug across the country...

    Loading up the live fishes!

    Darwin Day at the NC Museum of Natural Sciences (2019)

    Pupfishing on San Salvador Island, Bahamas (2018)

    Collecting pupfish at Fort Fisher, NC (2017)

    March for Science (2017) (guess which 2 are lab members?!)

    Alumni: Grad students and Postdocs


    Jacquelyn Galvez

    Graduate Student 2019-2021

    Currently a PhD candidate in Jack Tseng's FAVE lab. email


    Joseph Heras

    Postdoctoral Scholar 2020

    Microbiome diversity and function in Caribbean pupfishes (in prep). Currently (August, 2020) an Assistant Professor in the Biology Department at CSU San Bernardino. website email

    Jelmer Poelstra

    Postdoctoral scholar 2016-2017

    Demographic history of diversification in Cameroon crater lake cichlids (Mol Ecol 2018). Following a second postdoc in Anne Yoder's lab at Duke University, Jelmer is now a bioinformaticist at the Molecular and Cellular Imaging Center, Ohio State University.

    Alumni: Lab managers and undergrads

    Nicole Suren

    Lab Manager 2019-2020

    Kristi Dixon

    Lab Manager 2018-2019

    Currently lab manager for the Peifer lab.

    Maria Solano

    Lab Technician 2018-2019

    Delaney O'Connell

    Undergraduate 2019-2020

    Logan Buie

    Lab Manager 2017-2018

    Currently staff at the North Carolina Museum of Natural Sciences

    Jackson Waite-Himmelright

    Barombi Mbo, Cameroon field technician 2016

    Currently a prospective graduate student

    Stephanie Jeselson

    Undergraduate 2016-2017

    Clare Bocklage

    Undergraduate 2015-2017

    Katelyn Gould

    Lab Manager 2015-2017

    Currently a grad student in John Bruno's lab at UNC Chapel Hill


    Google scholar profile

    *Graduate students, †Postdoctoral scholars, •Undergraduate students

    Richards EJ*, Martin CH. 2021. We get by with a little help from our friends: Diversity begets diversity through shared adaptive genetic variation. preprint

    Patton AH†, Richards EJ*, Gould KJ, Buie LK, Martin CH. 2021. The first genotypic fitness network in a vertebrate reveals that hybridization increases access to novel fitness peaks. preprint

    St. John ME*, Richards EJ*, Dunker JC•, Romero S•, Martin CH. 2021. Parallel genetic changes underlie integrated craniofacial traits in an adaptive radiation of trophic specialist pupfishes. preprint

    Heras J†, Martin CH. 2021. Nonadaptive radiation of the gut microbiome in an adaptive radiation of Cyprinodon pupfishes with minor shifts for scale-eating. preprint

    Richards EJ*, McGirr JA*, Jeremy Wang J, St John ME*, Poelstra JW†, Solano MJ•, O'Connell DC•, Turner BJ, Martin CH. 2021. A vertebrate adaptive radiation is assembled from an ancient and disjunct spatiotemporal landscape. Proceedings of the National Academy of Sciences pdf supplement

    McGirr JA*, Martin CH. 2021. Few fixed variants between trophic specialist pupfish species reveal candidate cis-regulatory alleles underlying rapid craniofacial divergence. Molecular Biology and Evolution 38:405-423. pdf

    Higham TE, Ferry LA, Schmitz L, Irschick DJ, Starko S, Anderson PSL, Bergmann PJ, Jamniczky HA, Monteiro LR, Navon D, Messier J, Carrington E, Farina SC, Feilich KL, Hernandez LP, Johnson MA, Kawano SM, Law CJ, Longo SJ, Martin CH, Martone PT, Rico-Guevara A, Santana SE, Niklas KJ. 2021. Linking ecomechanical models and functional traits to understand phenotypic diversity. Trends in Ecology and Evolution. In press. pdf

    Perevolotsky T, Martin CH, Rivlin A, Holzman R. 2020. Work that body: fin and body movements determine herbivore feeding behavior in a natural reef environment. Proceedings of the Royal Society B 287:20201903. pdf

    Olsson KH, Martin CH, Holzman R. 2020. Hydrodynamic simulations of the performance landscape for suction-feeding fishes reveal multiple peaks for different prey types. Integrative and Comparative Biology pdf

    Martin CH, Gould K. 2020. Surprising spatiotemporal stability of a multi-peak fitness landscape revealed by field experiments measuring hybrid fitness. Evolution Letters 4:530-544. pdf

    McGirr JA*, Martin CH. 2020. Ecological divergence in sympatry causes gene misexpression in hybrids. Molecular Ecology pdf

    St. John ME*, Dixon KE•, Martin CH. 2020. Oral shelling within an adaptive radiation of pupfishes: Testing the adaptive function of a novel nasal protrusion and behavioral preference. Journal of Fish Biology pdf

    St. John ME*, Holzman R, Martin CH. 2020. Rapid adaptive evolution of scale-eating kinematics to a novel ecological niche. pdf

    Gillespie RG, Bennett GM, De Meester L, Feder JL, Fleischer RC, Harmon LJ, Hendry AP, Knope ML, Mallet J, Martin CH, Parent CE, Patton AH†, Pfennig KS, Rubinoff D, Schluter D, Seehausen O, Shaw KL, Stacy E, Stervander M, Stroud JT, Wagner CE, Wogan GO. 2020. Comparing adaptive radiations across space, time, and taxa. Journal of Heredity. 111:1-20. pdf

    Martin CH, Richards EJ*. 2019. The paradox behind the pattern of rapid adaptive radiation: How can the speciation process sustain itself through an early burst? Annual Review of Ecology, Evolution, and Systematics 50:569-593. pdf

    Martin CH, McGirr JA*, Richards EJ*, St. John ME*. 2019. How to investigate the origins of novelty: insights gained from genetic, behavioral, and fitness perspectives. Integrative Organismal Biology pdf

    McGirr JA*, Martin CH. 2019. Hybrid gene misregulation in multiple developing tissues within a recent adaptive radiation of Cyprinodon pupfishes. PLOS One 14:e0218899. pdf

    Richards EJ*, Servedio MR, Martin CH. 2019. Searching for sympatric speciation in the genomic era. BioEssays 41:1900047. pdf

    St. John ME*, McGirr JA*, Martin CH. 2019. The behavioral origins of novelty: did increased aggression lead to scale-eating in pupfishes? Behavioral Ecology 30:557-569. pdf

    Martin CH, Richards EJ*. 2019. The paradox behind the pattern of rapid adaptive radiation: how can the speciation process sustain itself through an early burst? Annual Review of Ecology, Evolution, and Systematics pdf

    Davis AL•, Babb MH•, Lowe MC•, Yeh AT•, Lee BT•, Martin CH. 2019. Testing Darwin’s hypothesis about the wonderful Venus flytrap: marginal spikes form a “horrid prison” for moderate-sized insect prey. American Naturalist 93:309-317. pdf Media: Atlas Obscura Phys.org ScienceDaily

    McGirr JA*, Martin CH. 2018. Parallel evolution of gene expression between trophic specialists despite divergent genotypes and morphologies. Evolution Letters 2:62-75. pdf

    Richards EJ*, Poelstra JW†, Martin CH. 2018. Don’t throw out the sympatric species with the crater lake water: fine-scale investigation of introgression provides weak support for functional role of secondary gene flow. Evolution Letters 2:524-540. pdf

    Poelstra JW†, Richards EJ*, Martin CH. 2018. Speciation in sympatry with ongoing secondary gene flow and a potential olfactory trigger in a radiation of Cameroon cichlids. Molecular Ecology 27:4270–4288. pdf

    Martin CH, Turner BJ. 2018. Long-distance dispersal over land by fishes: extremely rare ecological events become probable over millennial timescales. Proceedings of the Royal Society B 285: 20172436. pdf

    Zeng Y•, Martin CH. 2017. Oxford Nanopore sequencing in a research-based undergraduate course. preprint

    Keren T, Kiflawi M, Martin CH, China V, Mann O, Holzman R. 2017. A complex performance landscape for suction-feeding reveals constraints and adaptations in a population of reef damselfish. preprint

    Hernandez LP, Adriaens D, Martin CH, Wainwright PC, Masschaele B, Derick M. 2017. Building trophic specializations that result in substantial niche partitioning within a young adaptive radiation. Journal of Anatomy. 232:173-185. pdf

    Richards EJ*, Martin CH. 2017. Adaptive introgression from distant Caribbean islands contributed to the diversification of a microendemic radiation of trophic specialist pupfishes. PLOS Genetics 13:e1006919. pdf Media: ScienceDaily

    Stager JC, Alton K, Martin CH, King DT, Livingstone DT. 2017. On the age and origin of Lake Ejagham and its endemic fishes. 2017. Quaternary Research. 89:21-32. pdf

    Martin CH, Höhna S. 2017. New evidence for the recent divergence of Devil's Hole pupfish and the plausibility of elevated mutation rates in endangered taxa. Molecular Ecology. 27:831-83. pdf

    Martin CH, Höhna S, Crawford JE, Turner BJ, Richards EJ*, Simons LH. 2017. The complex effects of demographic history on the estimation of substitution rate: concatenated gene analysis results in no more than twofold overestimation. Proceedings of the Royal Society B 284:20170537. pdf

    Martin CH, Erickson PA, Miller CT. 2017. The genetic architecture of novel trophic specialists: higher effect sizes are associated with exceptional oral jaw diversification in a pupfish adaptive radiation. Molecular Ecology 26:624-638. pdf supplement

    McGirr JA*, Martin CH. 2016. Novel candidate genes underlying extreme trophic specialization in Caribbean pupfishes. Molecular Biology and Evolution 34:873-888. pdf supplement Media: Phys.org ScienceDaily

    Martin CH. 2016. The cryptic origins of evolutionary novelty: 1,000-fold-faster trophic diversification rates without increased ecological opportunity or hybrid swarm. Evolution 70:2504-2519. pdf supplement

    Higham TE, Rogers SM, Langerhans RB, Jamniczky HA, Lauder GV, Stewart WJ, Martin CH, Reznick DN. 2016. Speciation through the lens of biomechanics: locomotion, prey capture, and reproductive isolation. Proceedings of the Royal Society B 283:1294-1304. pdf

    Martin CH. 2016. Context-dependence in complex adaptive landscapes: frequency and trait-dependent selection surfaces within an adaptive radiation of Caribbean pupfishes. Evolution 69:1406-1422. pdf supplement

    Martin CH, Crawford JE, Turner BJ, Simons LH. 2016. Diabolical survival in Death Valley: recent pupfish colonization, gene flow, and genetic assimilation in the smallest species range on earth. Proceedings of the Royal Society B 283:23-34. pdf supplement Media: Nature Science Science Magazine BBC Discovery News New Scientist Quirks and Quarks CBC radio interview

    Martin CH, Cutler JS, Friel JP, Dening Touokong C, Coop G, Wainwright PC. 2015. Complex histories of repeated gene flow in Cameroon crater lake cichlids cast doubt on one of the clearest examples of sympatric speciation. Evolution 69:1406-1422. pdf Jerry Coyne's jubilant blog posts: 1 2

    Musilová Z, Indermaur A, Nyom ARB, Tropek R, Martin CH, Schliewen UK. 2014. Persistence of Stomatepia mongo, an endemic cichlid fish of the Barombi Mbo Crater Lake, Southwestern Cameroon, with notes on its life history and behavior. Copeia 2014:556-560. pdf

    Martin CH, Feinstein LC. 2014. Novel trophic niches drive variable progress toward ecological speciation within an adaptive radiation of pupfishes. Molecular Ecology. 23: 1846-1862. pdf

    Martin CH, Wainwright PC. 2013. A remarkable species flock of Cyprinodon pupfishes endemic to San Salvador Island, Bahamas. Bulletin of the Peabody Museum of Natural History. 54:231-240. pdf  Media: UC Davis blog, Seriously Fish.

    Schmitz L, Motani R, Oufiero CE,Martin CH, McGee MD, Wainwright PC. 2013. Potential enhanced ability of giant squid to detect sperm whales is an exaptation tied to their large body size. BMC Evolutionary Biology 13:226. pdf

    Friedman M, Keck BP, Dornburg A, Eytan RI, Martin CH, Hulsey CD, Wainwright PC, Near TJ. 2013. Molecular and fossil evidence place the origin of cichlid fishes long after Gondwanan rifting. Proceedings of the Royal Society B 280:1770. pdf

    Martin CH, Wainwright PC. 2013. On the measurement of ecological novelty: scale-eating pupfish are separated by 168 million years from other scale-eating fishes. PLOS One 8:e71164. pdf

    Martin CH. 2013. Strong assortative mating by diet, color, size, and morphology but limited progress toward sympatric speciation in a classic example: Cameroon crater lake cichlids. Evolution 67:2114-2123. pdf

    Martin CH, Wainwright PC. 2013. Multiple fitness peaks on the adaptive landscape drive adaptive radiation in the wild. Science 339:208-211. pdf supplement press release Media: Carl Zimmer's blog.  Seriously Fish. Der Spiegel. Nothing in Biology Makes Sense.WhyFiles. Davis Enterprise.

    Schmitz L, Motani R, Oufiero CE, Martin CH, McGee MD, Gamarra AR, Lee JJ, Wainwright PC. 2013. Allometry indicates giant eyes of giant squid are not exceptional. BMC Evolutionary Biology 13:45. pdf

    Martin CH. 2012. Weak disruptive selection and incomplete phenotypic divergence in two classic examples of sympatric speciation: Cameroon crater lake cichlids. American Naturalist 180:E90-109. pdf supplement press release 

    Martin CH, Wainwright PC. 2011. Trophic novelty is linked to exceptional rates of morphological diversification in two adaptive radiations of Cyprinodon pupfishes. Evolution 65:2197-2212. pdf.supplement. press release. F1000. Media: MSNBC. UC Davis Aggie. Practical Fishkeeping. KillieNutz. LiveScience. anti-evolution press: Lutheran. spoof attack.

    Martin CH. 2010. Unexploited females and unreliable signals of male quality in a Malawi cichlid bower polymorphism. Behavioral Ecology 21:1195-1202. pdf

    Martin CH, Genner MJ. 2009. A role for male bower size as an intrasexual signal in a Lake Malawi cichlid fish. Behaviour 146:963-978. pdf

    Martin CH, Genner MJ. 2009. High niche overlap between two successfully coexisting pairs of Lake Malawi cichlids.  Canadian Journal of Fisheries and Aquatic Sciences. 66:579-588. pdf supplement

    Martin CH, Johnsen S. 2007. A field test of the Hamilton-Zuk hypothesis in the guppy Poecilia reticulata. Behavioral Ecology and Sociobiology 61:1897-1909. pdf

    Yang AS, Martin CH, Nijhout HF. 2004. Geographic variation of caste structure among ant populations. Current Biology 14: 514-519. pdf.


    Red Sea 2019

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    Bahamas 2017

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    Cameroon 2016

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    Bahamas 2015

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    Bahamas 2014

    Bahamas 2013

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    Death Valley 2012

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    Bahamas 2011

    Dominican Republic 2011

    Limia perugiae collected from Lago Enriquillo.

    Cameroon 2010

    Sarotherodon knauerae, previously S.

    Mexico 2009

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    Bahamas 2008

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    Malawi 2006-2007

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    Madagascar 2005

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    Peru 2004

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    Trinidad 2004

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    Costa Rica 2003

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