Rubenstein Lab

understanding why animal societies form and how organisms cope with environmental change

Twitter @DustRubenstein

integrating animal behavior  

 through genomics, neuroendocrinology, comparative methods, theory and more

spanning molecules to populations

from invertebrates to vertebrates

across levels of analysis, scales of biological organization and the globe

marine igaunas


house sparrows



snapping shrimps




pebble-mound mice

past systems

burying beetles

Egernia lizards

current systems

social evolution

 exploring the causes and consequences of sociality & evolutionary transitions in social organization

environmental uncertainty and animal societies. Work in cooperatively breeding starlings, weavers, pebble-mound mice and burying beetles examines how unpredictable climate variation influences social evolution. 

evolution of social diversity.  We are examining the evolution of social diversity in snapping shrimps, coral-dwelling gobies, and birds by quantifying life history variation and exploring the key evolutionary transitions among social states. 

genetic and neural mechanisms of social behavior.  We are studying the mechanistic bases of caste differentiation, social phenotypes, and social decision making in snapping shrimps, burying beetles, and Egernia lizards by examining role- and population-specific patterns of gene expression, signatures of genetic and epigenetic variation, and brain architecture. 

phenotypic consequences of social living.  We are studying how inter- and intra-sexual competition influence the evolution of social signaling (song and plumage) and patterns of sexual dimorphism in starlings, hummingbirds, and pebble-mound mice.

genotypic consequences of social living.  We are examining how social living influences the evolution of genome structure by studying the relationships among social organization, genome size, and transposable element accumulation in snapping shrimps using population genetic and demographic approaches. 

cooperation and ecological dominance.  We are exploring how cooperative behavior in birds, burying beetles, and snapping shrimps influences competitive ability against conspecifics, niche breadth and range expansion, ecological generalism vs. specialism, and ecological dominance.

environmental coping

determining the behavioral, physiological and molecular adaptations used to cope with environmental change

adaptation along environmental gradients.  We are examining stress physiology, immune function, color evolution, and patterns of genetic and epigenetic variation in different populations of starlings along temperature and precipitation gradients in the tropics. 

a framework for environmental coping.  Using game theory and comparative analyses we are exploring the evolution of risk averse strategies to cope with environmental uncertainty.  We are also developing a theoretical framework that predicts evolutionary responses to environmentally-driven fluctuating selection, and using it to explore the evolution of physiological coping mechanisms, as well as their genetic and epigenetic architectures.

epigenetics, plasticity and environmental variation.  We are examining how developmental conditions (both social and environmental) influence social phenotypes, stress physiology, and fitness later in life, and we are exploring DNA methylation as one potential mechanism underlying this relationship in starlings. Although we emphasize the stress axis, we also look globally at patterns of DNA methylation across the genome.

research articles


Siller, SJ and DR Rubenstein. 2019. A tissue comparison of DNA methylation in the glucocorticoid receptor (Nr3c1) gene promoter in the European starling. Integrative and Comparative Biology. In press.  PDF

Rubenstein DR, JA Ågren, L Carbone, NC Elde, HE Hoekstra, KM Kapheim, L Keller, CS Moreau, AL Toth, S Yeaman and HA Hofmann. 2019. Coevolution of genome architecture and social behavior. Trends in Ecology and Evolution. In press.  PDF

Cheng, Y-R, DR Rubenstein and S-F Shen. 2019. Nest predation predicts infanticide in a cooperatively breeding bird. Biology Letters 15:20193014.  PDF

Chak STC and DR Rubenstein. 2019. Social transitions in sponge-dwelling snapping shrimp. Current Opinion in Insect Science 34:33-39.  PDF

Lin, Y-H, S-F Chan, DR Rubenstein, M Liu and S-F Shen. 2019. Resolving the paradox of environmental quality and sociality: the ecological causes and consequences of cooperative breeding in two lineages of birds. The American Naturalist 194:207-216.  PDF

Rubenstein, DR. 2019. Animal society. In Encyclopedia of Animal Cognition and Behavior (Vonk, J and TK Schelford, eds.). Springer, New York, pp. 1-3.  PDF

Ellis, VA, EHR Sari, DR Rubenstein, RC Dickerson, S Bensch and RE Ricklefs. 2019. The global biogeography of avian haemosporidian parasites is characterized by local diversification and intercontinental dispersal. Parasitology 146:213-219.  PDF

Shen, S-F and DR Rubenstein. 2019. Environmental uncertainty and social behavior. In Encyclopedia of Animal Behavior, 2nd Edition (Choe, J, ed.). Elsevier, New York, Volume 4, pp. 807-815.  PDF


Guindre-Parker, S and DR Rubenstein. 2018. No short-term physiological costs of offspring care in a cooperatively breeding bird. Journal of Experimental Biology 221:jeb186569.  PDF

Guindre-Parker, S and DR Rubenstein. 2018. The oxidative costs of parental care in cooperative and pair-breeding African starlings. Oecologia 188:53-63.  PDF

Liu, M, DR Rubenstein, S-A Cheong and S-F Shen. 2018. Multitasking and the evolution of optimal clutch size in fluctuating environments. Ecology and Evolution 8:8803–8817.  PDF

Guindre-Parker, S and DR Rubenstein. 2018. Multiple fitness benefits of alloparental care in a fluctuating environment. Royal Society Open Science 5:172406.  PDF

Pikus, AE, S Guindre-Parker and DR Rubenstein. 2018. Testosterone, social status and parental care in a cooperatively breeding bird. Hormones and Behavior 97:85-93.  PDF


Gaynor, KM, JW Solomon, JE Duffy, L Jessell, S Siller and DR Rubenstein. 2017. Development of genome- and transcriptome-derived microsatellites in related species of snapping shrimps with highly duplicated genomes. Molecular Ecology Resources 17:e160-e173.  PDF

Brooks, KC, R Maia, JE Duffy, KM Hultgren and DR Rubenstein. 2017. Ecological generalism facilitates the evolution of sociality in snapping shrimps. Ecology Letters 20:1516-1525.  PDF

Dantzer, B and DR Rubenstein. 2017. Introduction to symposium: the developmental and proximate mechanisms causing individual variation in cooperative behavior. Integrative and Comparative Biology 57:560-565.  PDF

S-F Shen, ST Emlen, WD Koenig and DR Rubenstein. 2017. The ecology of cooperative breeding behaviour. Ecology Letters 20:708-720.  PDF

Chak STC, JE Duffy, KM Hultgren and DR Rubenstein. 2017. Evolutionary transitions towards eusociality in snapping shrimps. Nature Ecology & Evolution 1:0096.  PDF

Cornwallis, CK, CA Botero, DR Rubenstein, PA Downing, SA West and AS Griffin. 2017. Cooperation facilitates the colonization of harsh environments. Nature Ecology & Evolution 1:0057.  PDF

Rubenstein DR and P Abbot. 2017. The evolution of social evolution. In Comparative Social Evolution (Rubenstein DR and P Abbot, eds.). Cambridge University Press, Cambridge, pp. 1-18.  PDF  

Hultgren KM, JE Duffy and DR Rubenstein. 2017. Sociality in snapping shrimps. In Comparative Social Evolution (Rubenstein DR and P Abbot, eds.). Cambridge University Press, Cambridge, pp. 224-249.  PDF  

Rubenstein DR and P Abbot. 2017. Social synthesis: opportunities for comparative social evolution. In Comparative Social Evolution (Rubenstein DR and P Abbot, eds.). Cambridge University Press, Cambridge, pp. 427-452.  PDF  


Hofmeister, NR and DR Rubenstein. 2016. Environmental variability and the evolution of the glucocorticoid receptor (Nr3c1) in African starlings. Ecology Letters 19:1219-1227.  PDF

Hofmann, HA, SCP Renn and DR Rubenstein. 2016. Introduction to symposium: new frontiers in the integrative study of animal behavior: nothing in neuroscience makes sense except in the light of behavior. Integrative and Comparative Biology 56:1192-1196.  PDF

Rubalcaba JG, V Polo, R Maia, DR Rubenstein and JP Veiga. 2016. Sexual and natural selection in the evolution of extended phenotypes: the use of green nesting material in starlings. Journal of Evolutionary Biology 29:1585-1592.  PDF  

Jeffrey NW, KM Hultgren, TCS Chak, TR Gregory and DR Rubenstein. 2016. Patterns of genome size variation in snapping shrimp. Genome 59:393-402.  PDF  

Maia R, DR Rubenstein and MD Shawkey. 2016. Selection, constraint and the evolution of coloration in African starlings. Evolution 70:1064-1079.  PDF

Rubenstein DR, CA Botero and EA Lacey. 2016. Discrete but variable structure of animal societies leads to the false perception of a social continuum. Royal Society Open Science 3:160147.  PDF

Keen SC, CD Meliza, JA Piloswky and DR Rubenstein. 2016. Song in a social and sexual context: vocalizations signal identity and rank in both sexes of a cooperative breeder. Frontiers in Ecology and Evolution 4:46.  PDF

Rubenstein DR, HE Skolnik, A Berrio, F Champagne, S Phelps and J Solomon. 2016. Sex-specific fitness effects of unpredictable early life conditions are associated with DNA methylation in the avian glucocorticoid receptor. Molecular Ecology 25:1714-1728.  PDF

Rubenstein DR. 2016. Superb starlings: cooperation and conflict in an unpredictable environment. In Cooperative Breeding in Vertebrates: Studies of Ecology, Evolution, and Behavior (Koenig WD and JL Dickinson, eds.). Cambridge University Press, Cambridge, pp. 181-196.  PDF


Chak TCS, DR Rubenstein and JE Duffy. 2015. Social control of reproduction and breeding monopolization in the eusocial snapping shrimp Synalpheus elizabethae. The American Naturalist 186:660-668.  PDF

Pollack LJ and DR Rubenstein. 2015. The fitness consequences of kin-biased dispersal in a cooperatively breeding bird. Biology Letters 11:20150336.  PDF

Chak TCS, JE Duffy and DR Rubenstein. 2015. Reproductive skew drives patterns of sexual dimorphism in sponge-dwelling snapping shrimps. Proceedings of the Royal Society of London B 282:20150342.  PDF

Apakupakul K and DR Rubenstein. 2015. Bateman’s principle is reversed in a cooperatively breeding bird. Biology Letters 11:20150034.  PDF

Taborsky M, HA Hofmann, AK Beery, DT Blumstein, LD Hayes, EA Lacey, EP Martins, SM Phelps, NG Solomon and DR Rubenstein. 2015. Taxon matters: promoting integrative studies of social behavior. Trends in Neuroscience 38:189-191.  PDF

Weinman LR, J Solomon and DR Rubenstein. 2015. A comparison of single nucleotide polymorphism and microsatellite markers for analysis of parentage and kinship in a cooperatively breeding bird. Molecular Ecology Resources 15:502-511.  PDF

Botero CA, FJ Weissing, J Wright and DR Rubenstein. 2015. Evolutionary tipping points in the capacity to adapt to environmental change. Proceedings of the National Academy of Sciences USA 112:184-189.  PDF

Rubenstein DR and HA Hofmann. 2015. The integrative study of animal behavior. Current Opinion in Behavioral Sciences 6:v-viii.  PDF

Rubenstein DR and HA Hofmann. 2015. Proximate pathways underlying social behavior. Current Opinion in Behavioral Sciences 6:154-159.  PDF


Hofmann HA, AK Beery, DT Blumstein, ID Couzin, RL Earley, LD Hayes, PL Hurd, EA Lacey, SM Phelps, NG Solomon, M Taborsky, LJ Young and DR Rubenstein. 2014. An evolutionary framework for studying mechanisms of social behavior. Trends in Ecology and Evolution 29:581-589.  PDF

Sun S-J, DR Rubenstein, J-N Liu, M Liu, B-F Chen, S-F Chan, W Hwang, P-S Yang and S-F Shen. 2014. Climate-mediated cooperation promotes niche expansion in burying beetles. eLife 3:e02440.  PDF

Shen S-F, E Akçay and DR Rubenstein. 2014. Group size and social conflict in complex societies. The American Naturalist 183:301-310.  PDF


Keen SC, CD Meliza and DR Rubenstein. 2013. Flight calls signal group and individual identity but not kinship in a cooperatively breeding bird. Behavioral Ecology 24:1279-1285.  PDF

Meliza CD, SC Keen and DR Rubenstein. 2013. Pitch- and spectral-based dynamic time warping methods for comparing field recordings of harmonic avian vocalizations. Journal of the Acoustical Society of America 134:1407-1415.  PDF

Seddon N, CA Botero, JA Tobias, PO Dunn, H MacGregor, DR Rubenstein, A Uy,  JT Weir, LA Whittingham and RJ Safran. 2013. Sexual selection accelerates signal evolution during speciation in birds. Proceedings of the Royal Society of London B 280:20131065.  PDF

Maia R, DR Rubenstein and MD Shawkey. 2013. Key ornamental innovations facilitate diversification in an avian radiation. Proceedings of the National Academy of Sciences USA 110:10687-10692.  PDF

Mark MM and DR Rubenstein. 2013. Physiological costs and carry-over effects of avian interspecific brood parasitism influence reproductive tradeoffs. Hormones and Behavior 63:717-722.  PDF

Pilowsky JA and DR Rubenstein. 2013. Social context and the lack of sexual dimorphism in song in an avian cooperative breeder. Animal Behaviour 85:709-714.  PDF

Rubenstein DI and DR Rubenstein. 2013. Social behavior. In Encyclopedia of Biodiversity, 2nd Edition (Levin, SA, ed.). Elsevier, Volume 6, pp. 571-579.  PDF

Duffy JE, KS Macdonald, KM Hultgren, TCS Chak and DR Rubenstein. 2013. Decline and extinction of Caribbean eusocial shrimp. PLOS ONE 8:e54637.  PDF

Creel S, B Danzter, W Goymann and DR Rubenstein. 2013. The ecology of stress: effects of the social environment. Functional Ecology 27:66-80.  PDF


Rubenstein DR. 2012. Family feuds: social competition and sexual conflict in complex societies. Philosophical Transactions of the Royal Society B 367:2304-2313.  PDF

Rubenstein DR. 2012. Sexual and social competition: broadening perspectives by defining females roles. Philosophical Transactions of the Royal Society B 367:2248-2252.  PDF

Lovette IJ, BS Arbogast, RL Curry, RM Zink, CA Botero, JP Sullivan, AL Talaba, RB Harris, DR Rubenstein, RE Ricklefs and E Bermingham. 2012. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Molecular Phylogenetics and Evolution 63:219-229.  PDF

Botero CA and DR Rubenstein. 2012. Fluctuating environments, sexual selection and the evolution of flexible mate choice in birds. PLOS ONE 7:e32311.  PDF


Rubenstein DR. 2011. Spatiotemporal environmental variation, risk aversion and the evolution of cooperative breeding as a bet-hedging strategy. Proceedings of the National Academy of Sciences USA 108:10816-10822.  PDF

Jetz W* and DR Rubenstein*. 2011. Environmental uncertainty and the global biogeography of cooperative breeding in birds. Current Biology 21:72-78.  *contributed equally  PDF


Rubenstein DR and JA Kealey. 2010. Cooperation, conflict, and the evolution of complex animal societies. Nature Education Knowledge 1:47.  WEB

Blumstein DT, LA Ebensperger, LD Hayes, RA Vásquez, TH Ahern, JR Burger, AG Dolezal, A Dosmann, G González-Mariscal, BN Harris, EA Herrera, EA Lacey, J Mateo, L McGraw, D Olazabal, M Ramenofsky, DR Rubenstein, SA Sakhai, W Saltzman, C Sainz-Borgo, M Soto-Gamboa, ML Stewart, TW Wey, JC Wingfield and LJ Young. 2010. Towards an integrative understanding of social behavior: new models and new opportunities. Frontiers in Neuroscience 4:1-9.  PDF  


Rubenstein DR and IJ Lovette. 2009. Reproductive skew and selection on female ornamentation in social species. Nature 462: 786-789.  PDF   

Rubenstein DR and S-F Shen. 2009. Reproductive conflict and the costs of social status in cooperatively breeding vertebrates. The American Naturalist 173:650-661.  PDF   


Rubenstein DR and ME Hauber. 2008. Dynamic feedback between phenotype and physiology in sexually selected traits. Trends in Ecology and Evolution 23:655-658.  PDF

Rubenstein DR, AF Parlow, CR Hutch and LB Martin. 2008. Environmental and hormonal correlates of immune activity in a cooperatively breeding tropical bird. General and Comparative Endocrinology 159:10-15.  PDF

Vitousek MN, K Nelson, DR Rubenstein and M Wikelski. 2008. Are hotshots always hot? A longitudinal study of hormones, behavior, and reproductive success in male marine iguanas. General and Comparative Endocrinology 157:227-232.   PDF

Lovette IJ, BV McCleery, AL Talba and DR Rubenstein. 2008. A complete species-level molecular phylogeny for the “Eurasian” starlings (Sturnidae: Sturnus, Acridotheres, and allies): recent diversification in a highly social and dispersive avian group. Molecular Phylogenetics and Evolution 47:251-260.  PDF

Rubenstein DR, BV McCleery and JE Duffy. 2008. Microsatellite development suggests evidence of polyploidy in the social sponge-dwelling snapping shrimp Zuzalpheus brooksi. Molecular Ecology Resources 8:890-894.  PDF

Martin LB and DR Rubenstein. 2008. Stress hormones in tropical birds: patterns and future directions. Ornitologia Neotropical 19 (Suppl.):207-218.  PDF  


Rubenstein DR and IJ Lovette. 2007. Temporal environmental variability drives the evolution of cooperative breeding in birds. Current Biology 17:1414-1419.  PDF

Rubenstein DR. 2007. Territory quality drives intraspecific patterns in extrapair paternity. Behavioral Ecology 18:1058-1064.  PDF

Rubenstein DR. 2007. Female extrapair mate choice in a cooperative breeder: trading sex for help and increasing offspring heterozygosity. Proceedings of the Royal Society of London B 274:1895-1903.  PDF

Rubenstein DR. 2007. Temporal but not spatial environmental variation drives adaptive offspring sex allocation in a plural cooperative breeder. The American Naturalist 170:155-165.  PDF

Lovette IJ and DR Rubenstein. 2007. A comprehensive molecular phylogeny of the starlings (Aves: Sturnidae) and mockingbirds (Aves: Mimidae): congruent mtDNA and nuclear trees for a cosmopolitan avian radiation. Molecular Phylogenetics and Evolution 44:1031-1056.  PDF

Sachs JL and DR Rubenstein. 2007. The evolution of cooperative breeding; is there cheating? Behavioural Processes 76:13-137.  PDF

Rubenstein DR. 2007. Stress hormones and sociality: integrating social and environmental stressors. Proceedings of the Royal Society of London B 274:967-975.  PDF

Vitousek MN, DR Rubenstein and M Wikelski. 2007. The evolution of foraging behavior in the Galápagos marine iguana: natural and sexual selection on body size drives ecological, morphological, and behavioral specialization. In Foraging Behavior in Lizards (Reilly SM, DB Miles and LD McBrayer, eds.). Cambridge University Press, Cambridge, pp. 491-507.  PDF

prior to 2007

Rubenstein DR, DI Rubenstein, PW Sherman and TA Gavin. 2006. Pleistocene Park: does re-wilding North America represent sound conservation for the 21st century? Biological Conservation 132:232-238.  PDF

Lovette IJ, DR Rubenstein and WN Watetu. 2006. Provisioning of fledgling conspecifics by males of the brood-parasitic cuckoos Chrysococcyx klaas and C. caprius. The Wilson Journal of Ornithology 118:99-101.  PDF

Rubenstein DR. 2005. Isolation and characterization of polymorphic microsatellite loci in the plural cooperatively breeding superb starling, Lamprotornis superbus. Molecular Ecology Notes 5:739-744.  PDF

Rubenstein DR and M Wikelski. 2005. Steroid hormones and aggression in female Galápagos marine iguanas. Hormones and Behavior 48:329-341.  PDF

McRae SB, ST Emlen, DR Rubenstein and SM Bogdanowicz. 2005. Polymorphic microsatellite loci in a plural breeder, the grey-capped social weaver (Pseudonigrita arnaudi), isolated with an improved enrichment protocol using fragment size-selection. Molecular Ecology Notes 5:16-20.  PDF

Royle JA and DR Rubenstein. 2004. The role of species abundance in determining breeding origins of migratory birds with stable isotopes. Ecological Applications 14:1780-1788.  PDF

Rubenstein DR and KA Hobson. 2004. From birds to butterflies: animal movement patterns and stable isotopes. Trends in Ecology and Evolution 19:256-263.  PDF

Rubenstein DR and M Wikelski. 2003. Seasonal changes in food quality: a proximate cue for reproductive timing in marine iguanas. Ecology 84:3013-3023.  PDF

Rubenstein DR, CP Chamberlain, RT Holmes, MP Ayres, JR Waldbauer, GR Graves and NC Tuross. 2002. Linking breeding and wintering ranges of a migratory songbird using stable isotopes. Science 295:1062-1065.  PDF

Rittschof D, J Sarrica and DR Rubenstein. 1995. Shell dynamics and microhabitat selection by striped legged hermit crabs, Clibanarius vittatus (Bosc). Journal of Experimental Marine Biology and Ecology 192:157-172.  PDF

other articles

Rubenstein DR and DI Rubenstein. 2016. From Pleistocene to trophic rewilding: a wolf in sheep’s clothing. Proceedings of the National Academy of Sciences USA 113:E1.  PDF

Rubenstein DR, H Hofmann, E Akçay, S Alonzo, E Archie, A Beery, R Calisi-Rodríguez, K Carleton, B Chow, J Dubnau, C Grozinger, E Ketterson, A Leifer, T Linksvayer, M MacManes, L Martin, K McGraw, L McGraw, T Mendelson, L O'Connell, A Ophir, L Remage-Healey, S Renn, T Roth, J Tung and S Woolley. 2014. New frontiers for the integrative study of animal behavior. National Science Foundation White Paper.  WEB

Rubenstein DR. 2012. The Flexible Phenotype: A Body-Centered Integration of Ecology, Physiology, and Behaviour (Book Review). The Quarterly Review of Biology 87:264.  PDF

Rubenstein DR and JE Duffy. 2012. Scientists at work: notes from the field in Belize. The New York Times July 18-27, 2012.  WEB

Rubenstein DR. 2011. From the big city to the bush. Mpala Memos July:7.  PDF

Rubenstein DR. 2010. Evolutionary Behavioral Ecology (Book Review). The Quarterly Review of Biology 85:504.  PDF

Rubenstein DR. 2010. Scientists at work: notes from the field in Kenya. The New York Times July 12-28, 2010.  WEB

Rubenstein DR. 2009. The secret lives of starlings. Natural History 118:28-33.  PDF

Rubenstein DR. 2009. Why I do science: the freedom to explore. SEED 21:34.  PDF

Rubenstein DR, PW Sherman, DI Rubenstein and TM Caro. 2007. Rewilding rebuttal. Scientific American October:12.  PDF

Rubenstein DR. 2006. Searching for starlings. Travel News April:58.  PDF

Rubenstein DR. 2006. Chasing starlings, chased by a lion. Living Bird 26:26-32.  PDF

Rubenstein DR. 2005. The uncommon lifestyle of the superb starling. BirdScope 19:20.  PDF

Rubenstein DR. 2001. The places you can go. Dartmouth Alumni Magazine May/June:24-25.  PDF


Rubenstein DR and J Alcock. 2018. Animal Behavior, 11th Edition. Oxford University Press, New York.  WEB

Rubenstein DR and P Abbot. 2017. Comparative Social Evolution. Cambridge University Press, Cambridge.  WEB

edited volumes

Rubenstein DR and HA Hofmann. 2015. New frontiers for the integrative study of animal behavior. Current Opinion in Behavioral Sciences 6:1-182.  WEB

Rubenstein DR, RO Prum and M Levandowsky. 2012. Sexual selection, social conflict and the female perspective. Philosophical Transactions of the Royal Society B 367:2248-2375.  WEB

principle investigator

CV          BIO

Dustin R. Rubenstein 

Dustin is an Associate Professor of Ecology, Evolution and Environmental Biology at Columbia University. His research takes an integrative approach to understand why complex animal societies form and how organisms cope with environmental change through studies that combine behavior, ecology, and evolution with those of the underlying molecular and neuroendocrine mechanisms. He has studied a variety of animals, including reptiles, birds, insects, crustaceans, and mammals throughout Africa, Asia, Australia, Central and South America, and the Caribbean.


Shana Caro ’17 - ’20 

Simons Foundation Fellow

Shana is interested in the evolution of social behavior – particularly in communication and parental care. During her PhD and as a postdoc, she investigated how ecology and life history traits affect parent-offspring communication both across and within bird species. At Columbia, she is integrating physiological, genetic, and behavioral data to explore how environmental and social factors shift how starlings communicate, and illuminate the mechanisms underlying these shifts.

Renata Mazzei ’19 - ’21 

Swiss National Science Foundation Early Postdoc.Mobility Fellow

During her PhD, Renata studied social behavior in gobies from both a proximate and and ultimate perspective. At Columbia, she will take a comparative and empirical approach to continue to study social evolution in gobies, combining field work throughout the Caribbean to identify key life history traits of goby species with phylogenetic comparative work of this group of fishes.

graduate students

Jay Falk ’13 -  PhD

NSF Graduate Research Fellow 

Jay is a PhD student in the Department of Neurobiology and Behavior at Cornell University co-advised by Mike Webster. For his undergraduate work, he studied reproductive isolation and mating behavior in Tribolium beetles. He is interested in sexual selection and trait evolution in a variety of species including insects, crustaceans, and birds. For his PhD, Jay is studying the evolution and maintenance of female plumage polymorphism in hummingbirds.

Yi-Ru Cheng ’14 -  PhD

Yiru studies the evolution of avian life history strategies and social behavior. For her MS, she studied how growth strategies could have evolved under different levels of predation pressure in North America passerine birds. For her PhD, she is examining social network dynamics in grey-capped social weavers living the East African savanna ecosystem. Her work emphasizes the formation and maintenance of social groups, as well as how environmental variation influences the stability and robustness of social networks across year.

Stefanie Siller ’16 -  PhD

NSF Graduate Research Fellow 

Stefanie is examining how early life conditions influence fitness later in life. After graduating college, she spent a year in Kenya working at the Mpala Research Centre doing conservation outreach with local schools and studying extrapair paternity in birds. For her PhD, she is examining how environmental conditions during early life influence DNA methylation in wild and captive house sparrows.

Shailee Shah ’16 -  PhD

Shailee is interested in social dominance, communication, and sexual selection in birds. As an undergraduate, she studied alarm calling in herring gulls. After graduation she worked on a project on Himalayan birds in her native India, and worked at the Cornell Lab of Ornithology doing multimedia and citizen science. For her PhD, she is using field experiments, long-term data, and population genetics to examine dispersal and the direct benefits that superb starlings receive while living in complex social groups with surprisingly low kin structure.

Alexis Earl ’19 -  PhD

Alexis is interested in the evolution of complex social behavior, communication, and cognition. For her MS she studied whether female ornamentation in Indian peafowl signals social status, and whether hormonal mechanisms link variation in ornamentation with status. For her PhD she will use long-term behavioral and epigenetic data in super starlings to investigate how social and ecological environments influence individual level trajectories of roles within complex social groups, and subsequent impacts on group stability and structure.

Catherine Yung-Yi Lan ’21  MA

Catherine is interested in the evolution of behavioral traits. After graduating college, she spent a year at Academia Sinica in Taiwan doing research on the effects of photoperiodism on burial behavior in hybrid beetles, as well as analyzing gene sequences for wild-caught beetles. For her MA, she will continue studying social behavior in burying beetles.

Jerry Shuzhe Guan ’21  MA

coming soon

undergraduate students

Arden Berlinger ’20 CC  

Science Research Fellow

Arden is a Science Research Fellow studying kin selection and helping behavior in superb starlings.

Faith Ajayi ’21 CC 

Rabi Scholar

Faith is a Rabi Scholar studying DNA methylation and gene expression in superb starlings.

high school students

research associates

Zhaleh Mahootian ’20 

Bronx High School of Science 

As part of the Biological and Physical Research Mentorship Program at her high school, Zhaleh is examining fecundity variation among species and within colonies of eusocial snapping shrimps.

Joseph Solomon

Joe received his MA from Hunter College before joining the lab. He manages the lab in New York, overseeing students, helping with bioinformatics and data analysis, and all aspects of lab work.

Wilson Nderitu

Wilson has worked on the African starling project since 2001. He left in 2006 to complete his Diploma in Wildlife Management at the Kenya Wildlife Service Training Institute.

Elkana Korir

Korir has worked on the African starling project since 2019.  Previously he worked on weavers and guinea fowl projects at the Mpala Research Centre.

former lab members

Solomon Chak ’16 - ’19 

Life Sciences Research Fellow 

Postdoctoral Scholar, NJIT 

Assistant Professor, SUNY Old Westbury 

(starting Fall 2020)

Rafael Maia ’15 - ’18 

Simons Foundation Fellow

Senior Data Scientist, Instacart

Katherine Brooks ’14 - ’17 

Columbia Frontiers of Science Fellow

Collection Analysis Librarian

Columbia University

Stephen Harris ’15 - ’16 

Columbia Frontiers of Science Fellow

Assistant Professor, SUNY Purchase

Melissa Mark ’09 - ’12 

NSF Postdoctoral Fellow

Director, Doris Duke Conservation Scholars Prg

Northern Arizona University

Sarah Guindre-Parker ’17 PhD

NSERC Graduate Fellow

Assistant Professor, Kennesaw State University

Eleanor Diamant ’17 MA

PhD student, UCLA

Yuki Haba ’17 MA

PhD student, Princeton University

Alyxandra Pikus ’17 MA

Associate Director of Program Development

Oxbridge Academic Programs

Natalie Hofmeister ’15 MA

PhD student, Cornell University

Rebecca Kelley ’13 MA

PhD, New Mexico State University

Data Scientist, Amazon

James Kealey ’12 MA

Science teacher

Bret Harte Middle School, Oakland

Kathleen Apakupakul ’12 MA

Research Associate

Institute for Conservation Medicine

Saint Louis Zoo

Sara Keen ’11 MA

PhD student, Cornell University

Laura Booth ’15 CC

Park Ranger, Muir Woods National Monument

Hannah Skolnik ’15 CC

DVM student, UC Davis

Lucia Weinman ’14 CC

PhD student, Rutgers University

Lea Pollack ’12 CC

PhD student, UC Davis

Julia Pilowsky ’12 CC

MA, Tufts University

PhD student, University of Copenhagen

Caitlin Dean ’12 CC

JD, University of Michigan

Associate, Fish & Richardson

Jeremy Law ’11 CC

MA, Columbia University

Manager, New York Tree Trust

undergraduate interns

Sarah Marazzi-Sassoon BC ’22

Ona Jae Lee SEAS ’21

Shoshana Sernik GS ’20

Sophia Kislik BC ’20

Christian Eggers CC ’20

Jake Arlow BC ’19

Joseph Knee CC ’19

Kaiulani Sakaguchi CC ’19

Francesca Garofalo CC ’18

Tatini Mal-Sarkar CC ’17

Michael Spiotta GS ‘17

Karen Bao CC ’16

Fayme Cai CC ’16

Catherine Chen CC ’15

Brahadheeshwar Sundararaju CC ’15

Elora Lopez CC ’15

Nathen Huang CC ’14

Kerstin Nolan CC ’14

Sonalee Rau CC ’14

Madeline Cohen CC ’13

Heather D’Angelo GS ’12

Ben Eckersley CC ’13

Nathan Bailey CC’ 12

Jordan Hollarsmith CC ’12

Linnet Jessell (University of Kings College) 

Brynn McCleery (Cornell University)

Suraj Nagaraj (UC Berkeley)

post-baccalaureate interns

Tyler Davis (Cornell University)

Kaitlyn Gaynor (Columbia University)

Rebecca Harris (Cornell University)

Sarah Khalil (Cornell University)

Brynn McCleery (Cornell University)

Lea Pollack (Columbia University)

Stefanie Siller (Princeton University)

Gabrielle Welsh (University of Maryland)

high school interns

Zhaleh Mahootian (Bronx High School of Science ’20)

Hank Marriott (Ethical Culture Fieldston School ’20)

Livia Marchese (Briarcliff High School ’19)

Regina Hashim (home school student ’19)

Samuel Levy (Abram Joshua Heschel School ’18)

Rebecca Marcus (Mamaroneck High School ’18)

Myron Huang (Bronx High School of Science ’18)

Katherine Grygiercyzk (Valley Stream South High School ’17)

Gillian Carling (The Bronx High School of Science ’13)

visiting graduate students and postdocs

Rebecca Calisi (University of California, Berkeley)

Rafael Maia (University of Akron)

Daniel Meliza (University of Chicago)

Juan Rubalcaba (King Juan Carlos University)

visiting faculty

Tim Greives (North Dakota State University)

behind the scenes looks at our work on avian and shrimp social evolution

edited volume in Current Opinion in Behavioral Sciences and NSF White Paper on integrative animal behavior

Program in Tropical Biology and Sustainability featured in:

                            Columbia College Annual Report                                                            Columbia College Today

                            a movie about the Columbia Global Centers                                    a movie about Columbia College

Rubenstein Lab featured in Columbia Magazine and Columbia College Today

field expeditions to Kenya and Belize featured in The New York Times


Solomon Chak accepted a position as an Assistant Professor at SUNY Old Westbury. Before he begins in Fall 2020, he will continue to study snapping shrimps as a postdoc for one more year at NJIT.  (6/19)

Shailee Shah received a Graduate Student Travel Award to attend this year's Animal Behavior Society meetings.  (6/19)

Shailee Shah was awarded a Sigma Xi Grant-in-Aid of Research.  (6/19)

Renata Mazzei received an Early Postdoc.Mobility Fellow funded by the Swiss National Science Foundation to study social evolution in gobies in the Rubenstein Lab.  (6/19)

Shailee Shah received a Van Tyne Research Award from the American Ornithological Society.  (4/19)

Stefanie Siller was named a Lead Teaching Fellow by Columbia’s Center for Teaching and Learning for 2019-2020.  (4/19)

Shailee Shah was awarded a Chapman Grant from the American Museum of Natural History.  (3/19)

Shailee Shah’s research was featured in a film by the Cornell Lab of Ornithology: A Day in the Life of a Field Researcher.  (3/19)

Shailee Shah received a travel award from the American Ornithological Society to present at the 2019 meetings.  (3/19)

Stefanie Siller and Shailee Shah were awarded an Education Outreach Grant from the Animal Behavior Society and an Outreach Grant from the Society for the Study of Evolution to write and illustrate a children’s book on superb starlings for the Northern Kenya Conservation Clubs.  (2/19) 

Rafael Maia accepted a position as a Senior Data Scientist at Instacart.  (1/19)


Shana Caro won the Thomas Henry Huxley Award & Marsh Prize from the London Zoological Society for the best thesis in the UK.  (4/18)

Stefanie Siller received a Van Tyne Research Award from the American Ornithological Society.  (4/18)

Jay Falk won an American Society of Naturalists Student Research Award.(4/18)

High school student Myron Huang won first prize in Animal Sciences at the New York City Science and Engineering Fair.  (3/18)

Jay Falk was awarded a Smithsonian Predoctoral Fellowship.(2/18)

Becca Marcus was named one of the top 300 high school scholars nationally in the Regeneron Science Talent Search.  (1/18)


Shailee Shah received a Travel Grant to present her work at the 2017 American Ornithology Society Meetings.  (8/17)

Katherine Brooks accepted a job as a Collection Analysis Librarian at Columbia University.  (6/17)

High school student Becca Marcus won an Acorda Scientific Excellence Award.  (6/17)

Congratulations to all of the new Rubenstein Lab graduates: Dr. Sarah Guindre-Parker, PhD and newly minted MAs Eleanor Diamant, Yuki Haba, and Alyxandra Pikus.  (5/17)

Stephen Harris accepted an Assistant Professor position at SUNY Purchase.  (5/17)

The Rubenstein Lab was featured in Columbia College Today.  (4/17)

Sarah Guindre-Parker was awarded a graduate student travel grant to present at the Animal Behavior Society Meetings.  (4/17)

High school student Becca Marcus won 4th place for her presentation at the Westchester Science and Engineering Fair.  (3/17)

Behind the Paper: a behind the scenes look at our latest work on shrimp social evolution.  (3/17)

Rubenstein Lab members moving on to top graduate programs: Eleanor Diamant MA ’17 (UCLA, PhD), Yuki Haba MA ’17 (Princeton University, PhD), Lucia Weinman CC ’14 (Rutgers University, PhD), Hannah Skolnik CC ’15 (UC Davis, DVM), and Maddy Cohen CC ’13 (University of Miami, MD).  (3/17)

Sarah Guindre-Parker was awarded a NSERC Postdoctoral Fellowship to do research on squirrels at the University of Guelph.  (3/17)

Behind the Paper: a behind the scenes look at our latest work on avian social evolution.  (2/17)


Yuki Haba was awarded a Sigma Xi Grant-in-Aid of Research.  (12/16)

Shana Caro received a Simons Foundation Junior Society Fellowship to join the lab to study parental care in starlings.  (12/16)

Katherine Brooks won a poster award at the 2016 Columbia Postdoc Symposium.  (10/16)

Eleanor Diamant was awarded a research grant from the Wilson Ornithological Society.  (6/16)

Postbac students Stefanie Siller and Sarah Khalil both received NSF Graduate Research Fellowships, and MA student Eleanor Diamant received an honorable mention. Former undergraduate student Lea Pollack CC ’12 also received the award.  (3/16)

Yuki Haba received a travel grant from the Columbia Earth Institute to study burying beetles in Japan this summer.  (3/16)

Solomon Chak received a Fellowship from the Life Sciences Research Foundation to do a postdoc at Columbia.  (3/16)

Sarah Guindre-Parker was awarded a travel grant from the International Society for Behavioral Ecology to attend the 2016 annual meetings in Exeter, UK.  (4/16)

In an article about study abroad programs at Columbia, the Columbia Daily Spectator highlighted the Program in Tropical Biology and Sustainability in Kenya.  (1/16)

Sarah Guindre-Parker was awarded the Lynn Riddiford Best Student Poster Award from the Division of Comparative Endocrinology at the Society for Integrative and Comparative Biology annual meetings.  (1/16)


Sarah Guindre-Parker was awarded a Sigma Xi Grant-in-Aid of Research to complete a collaborative project on avian malaria.  (12/15)

Jay Falk was awarded a Smithsonian Institution Short-term Fellowship and designated an Ernst Mayer Fellow.(11/15)

Rubenstein Lab research on superb starlings and grey-capped social weavers was highlighted in the new PBS documentary Animal Homes.  (4/15)

Jay Falk was awarded a Sigma Xi Grant-in-Aid of Research.  (4/15)

Sarah Guindre-Parker received an NSF Doctoral Dissertation Improvement Grant (DDIG) for her work on superb starlings.  (3/15)

Rubenstein Lab members moving on to top PhD programs: Sara Keen MA ’11 (Cornell), Lea Pollack CC ’12 (UC Davis), and Natalie Hofmeister MA ’15 (Cornell).  (3/15)

Stephen Harris received a a Columbia Frontiers of Science Fellowship and will be joining the lab in July.  (3/15)

Rafael Maia received a Simons Foundation Junior Society Fellowship and will be joining the lab in July.  (1/15)


Dustin Rubenstein was interviewed on ESPN Outside the Lines about co-evolution and the defensive shift in baseball.  (6/14)

Dustin Rubenstein was awarded a grant from the National Geographic Society to return to northern Kenya to study starling stress physiology.  (5/14)

Dustin Rubenstein was awarded a grant from the President’s Office at Columbia University to expand the Kenyan study abroad program.  (5/14)

Dustin Rubenstein and Hans Hofmann were awarded an NSF grant to organize a workshop entitled New Frontiers for the Integrative Study of Animal Behavior at the New York Genome Center in August 2014.  (5/14)

Sarah Guindre-Parker and Natalie Hofmeister both received research grants from the American Ornithologists’ Union.  (4/14)

Katherine Brooks was awarded a Columbia Frontiers of Science Fellowship to do postdoctoral work in the lab.  (4/14)

Dustin Rubenstein was promoted to Associate Professor after a successful 5th Year review.  (4/14) 

Lab alumnus Maddy Cohen CC ’13 was featured in 2012-2013 Columbia College Annual Report about her global study abroad experiences while at Columbia.  (2/14)


Lab alumni Julia Pilowsky CC ’12 and Maddy Cohen CC ’13 were featured in Columbia College Today about their study abroad and research experiences.  (12/13)

Sarah Guindre-Parker was awarded an Animal Behavior Society Student Research Grant.  (5/13)

Gillian Carling, a senior at The Bronx High School of Science, was admitted to Columbia’s class of 2017 and will be starting in the fall.  (4/13) 

Hannah Skolnik was admitted to the 2013 Amgen Scholar’s Program as 1 of 250 students from a national pool of more than 6,200 applications.  (4/13)

Kaitlyn Gaynor and Julia Pilowsky won NSF Graduate Research Fellowships to fund their PhD work at UC Berkeley and Tufts, respectively.  (4/13)

Rebecca Kelley received an NSF Graduate Research Fellowship honorable mention.  (4/13)

High school student Gillian Carling was named a Finalist at the New York City Science and Engineering Fair for her work on snapping shrimp.  (3/13)

Julia Pilowsky’s undergraduate senior was published in the April edition of Animal Behaviour, where it is featured in the In Focus section.  (3/13)


Dustin Rubenstein’s research trip with Emmett Duffy to Belize was featured in The New York Times’ Scientists at Work Blog.  (7/12)

Dustin Rubenstein co-edited a recent issue of the Philosophical Transactions of the Royal Society B.  (7/12)

Lea Pollack was awarded Departmental Honors, the only EB major to receive the award.  (5/12)

Rebecca Kelley was awarded an Animal Behavior Society Student Research Grant.  (4/12)

Rebecca Kelley received an honorable mention for her NSF Graduate Research Fellowship Application.  (4/12)

Sarah Guindre-Parker was awarded a Canadian NSERC Fellowship to fund her PhD work.  (4/12)

Julia Pilowsky received a Charles Turner Travel Award from the Animal Behavior Society to present at the Annual Meetings this summer.  (3/12)


Rebecca Kelley was awarded a Sigma Xi Grant-in-Aid of Research.  (12/11)

James Kealey’s work on snapping shrimp was profiled in the STRI News.  (12/11)

Julia Pilowsky’s senior thesis research on song in superb starlings was featured in the Columbia Spectator.  (9/11)

Sara Keen was awarded a Conference Travel Award by the American Ornithologists’ Union to present her thesis work at the Annual Meetings.  (5/11)

James Kealey was awarded a Smithsonian Institution Short-term Fellowship and the STRI Ernst Mayer Fellowship.  (4/11)

James Kealey was awarded a Columbia Institute of Latin American Studies research grant.  (4/11) 

Rubenstein Lab was featured in the Spring 2011 issue of the Columbia Magazine.  (4/11)

Sara Keen received her second honorable mention for her NSF Graduate Research Fellowship Application.  (4/11)

Dustin Rubenstein co-organized a one-day symposium at Columbia on “Sexual selection, social conflict and the female perspective”.  (4/11)

Dustin Rubenstein and colleagues (Jennifer Fewell and Jim Hunt) were awarded a grant from NESCent to organize a working group entitled Large-scale demographic, network and behavioral trait analyses of sociality from 2011 - 2013.  (2/11)  


Dustin Rubenstein and colleagues (Eileen Lacey, Steven Phelps, and Nancy Solomon) were awarded a grant from NESCent to organize a working group entitled Integrative models of vertebrate sociality: evolution, mechanism and emergent properties from 2011 - 2012.  (10/10)           

Dustin Rubenstein was awarded the 2011 Young Investigator Award by the Animal Behavior Society.  (8/10)

Rubenstein Lab is now on Facebook.  (8/10)

Dustin Rubenstein’s research trip to northern Kenya was featured in The New York Times’ Scientists at Work Blog.  (7/10)

Godffrey Manyaas is the proud father of a beautiful new baby girl.  (4/10)

Sara Keen received an honorable mention for her NSF Graduate Research Fellowship Application.  (4/10)

Dustin Rubenstein was awarded the 2010 The Ned K. Johnson Young Investigator Award by the American Ornithologists‘ Union.  (2/10)

James Kealey was awarded a Sigma Xi Grant-in-Aid of research.  (1/10) 


Sara Keen and James Kealey were both awarded research travels grants from Columbia’s Earth Institute.  (12/09)

Dustin Rubenstein was profiled in Nature and interviewed on the Nature Podcast.  (12/09)

Wilson Nderitu was profiled in Mpala Memos.  (10/09)

Dustin Rubenstein’s 2004 TREE paper was named as a “Fast Moving Front” by Thomson Reuters.  (9/09)

Rubenstein Lab has finally moved to Columbia University and was profiled in The Record.  (8/09)

Melissa Mark was awarded a 3-yr NSF Minority Postdoctoral Research Fellowship to work in the Rubenstein Lab starting in Jan 2010.  (5/09)


Tyler Davis was awarded an Explorers’ Club Grant for his research on sexual conflict in superb starlings.  (6/08)

Wilson Nderitu graduated with his Diploma in Wildlife Management from the Kenya Wildlife Service Training Institute.  (6/08)

Dustin Rubenstein will start as an Assistant Professor of Ecology, Evolution and Environmental Biology at Columbia University in Fall 2009.  (3/08)

Rebecca Calisi was awarded a Grant-in-Aid of Research from the Society for Integrative and Comparative Biology to study starlings in Kenya.  (1/08) 

Rebecca Calisi won the award for Best Student Poster from the Society of Integrative and Comparative Biology’s Division of Neuroscience.  (1/08)

Columbia is a great place to begin or to continue your academic career. In addition to the diverse community in the Department of Ecology, Evolution and Environmental Biology and its partner institutions, the Rubenstein Lab has strong links to colleagues in the Department of Biological Sciences, Department of Psychology, Program in Neurosciences and Behavior, and the Zuckerman Mind, Brain, Behavior Institute. We also interact regularly with colleagues from CUNY, Fordham, Rockefeller, NJIT, NYU, and the other many academic institutions in NYC. Members of the lab are typically are affiliated with the American Museum of Natural History, where we do much of our molecular work. We also work closely with the New York Genome Center and a variety of core labs at the Columbia University Medical Center and other local hospitals. While Columbia may be relatively small in size, NYC is not. Whether it is access to a lab, a collaborator, or a colleague, most of what you could ever need is only a short subway ride away. And with three international airports, it’s equally easy to do field work all over the world.

prospective PhD students

I encourage you to think big! What are the most interesting problems in behavioral and evolutionary ecology right now and how are you going to solve them? I am particularly interested in applicants looking to integrate studies of behavior, evolution, ecology, endocrinology, neuroscience, genomics, or epigenetics. Your goal as a graduate student should be to not only master your topic of study and become an expert in your discipline and study system, but also to push the field and further develop a body of evolutionary or ecological theory. My job is to help you succeed in doing this. My approach is one of guided independence, but I will work with you to develop questions and formulate hypotheses. I will help you become a better writer and thinker. Although I want students to do fieldwork, I will also require that you learn a variety of laboratory techniques so that you become trained as an integrative biologist that can think and work across disciplines. I believe that integrative research is the future of animal behavior. Recognizing this now and being trained this way during your PhD will make you more marketable for postdocs and jobs in the future.

Prospective students should have extensive field and/or lab experience, including having conducted their own independent research. Interested applicants should demonstrate creativity, perseverance, and a passion of science. Students admitted to the PhD program in Environmental Biology will be offered up to five years of support, including a generous stipend, subsidized housing near campus, and modest research funds. 

Interested applicants should email their CV, a summary of previous research experience, and a description of the project that they would like to do at Columbia.

prospective MA students

Masters students are generally accepted into the E3B program and then find an advisor during their first year at Columbia. However, if you are interested in working with me, feel free to get in touch.

prospective undergraduate students

There are many opportunities for undergrads to gain experience with molecular techniques, hormone assays, immune assays, bioinformatics, comparative methods, video analysis, or field work. Students work initially with an older member of the lab, eventually developing their own project for a senior thesis. Ultimately, I hope that each undergraduate will publish their thesis as a first-authored paper in a peer-reviewed journal. 

prospective postdocs

Because my interests are broad and varied, I am open to postdocs wanting to work on a variety of systems and question. If you might be interested in working in my lab, please email me and we can discuss ideas and opportunities. I currently do not have funding to support postdocs, but I am happy to work together on fellowship applications to fund training at Columbia. The best current opportunities for postdoctoral support are fellowships from the Simons Foundation, Fulbright, Marie Curie, NSF, NERC, NSERC, Life Sciences Research Foundation, The Human Frontiers Program, etc.

     Columbia University, Department of Ecology, Evolution and Environmental Biology, 10th Floor Schermerhorn Extension, MC5557, 1200 Amsterdam Avenue, New York, NY 10027

     Office: 90 Morningside Drive, Basement #3 • Lab: 851-854 Schermerhorn Extension

     Tel: 212-854-4881 • Fax: 212-854-8188 • Lab Tel: 212-854-5330 • Email: dr2497[at] • Twitter: @DustRubenstein

Last updated 11 August 2019

© 2002-2019   Dustin R. Rubenstein

Columbia University in the City of New York