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Principal investigator, Antónia Monteiro

Evolutionary Developmental Biology

Professor, 2020,  National University of Singapore and Yale-NUS-College, Associate Professor, 2013, National University of Singapore and Yale-NUS College   
Assistant Professor, 2006 Yale University
Assistant Professor, 2001-2006, University at Buffalo
Postdoctoral work, 1998-2001, Leiden University
Postdoctoral work, 1997-98, Harvard University
PhD, 1997, Edinburgh University

Contact information:

Department of Biological Sciences, National University of Singapore, 14 Science Drive 4
Singapore 117543
Yale-NUS-College, 6 College Avenue East,
Singapore 138614
tel: + 65 660 12108
Current lab members:
Postdoctoral fellow Emilie Dion described the chemical profile of wings of dry season and wet season B. anynana butterflies in a collaboration with Joanne Yew. She is now testing whether Bicyclus anynana females can learn to prefer novel sex pheromone blends.
Postdoctoral fellow Heidi Connahs is studying the role of Distal-less cis-regulatory elements using FAIRE-Seq and CRISPR experiments. She is also studying the role of decapendaplegic (dpp) in eyespot development in Nymphalid butterflies.
Postdoctoral fellow Yuji Matsuoka is investigating the role of multiple genes in color pattern development using CRISPR and introducing a PhiC31 landing pad in Bicyclus for future work with reporter constructs.
Postdoctoral fellow Anupama Prakash is performing single-cell sequencing on developing butterfly wings to assign transcripts to particular colored scales. She is also interested in the evolution of sexual dimorphism and in the evo-devo of structural colors on the wings of Junonia butterflies.
Postdoctoral fellow Ian Chan is working on the evolution of color on the wings of butterflies
Postdoctoral fellow Dantong Zhu is working on the comparative genomics of species with eyespots and species without eyespots

PhD student Tirtha Banerjee is interested in exploring the processes of early vein patterning and wing development in butterflies.

PhD student Jocelyn Wee is interested in exploring the gene expression differences between spots and eyespots to try and understand what makes these two wing patterns different.
PhD student Juan Olvido is interested in exploring the physiological correlates of white and brown eyeballs (yes, eyeballs, not eyespots) in two species of Orangutans from SE Asia.
PhD student Suriya Narayanan is comparing the set of genes expressed in eyespots to sets of genes expressed in other tissues to try and infer where eyespots came from. He is also testing the genetic basis for eyespot number variation with genetic mapping and CRISPR-Cas9.
PhD student Shen Tian is comparing the set of microRNAs expressed in the wings of dry and wet seasonal forms of Bicyclus anynana.
PhD student Gowri V. is working on larval odor learning in Bicyclus anynana
Master student Maniza Huq is testing why dry season males prefer to mate with females with dorsal and ventral eyespots intact. She is looking for correlations between female fecundity and female eyespot center size.
Honors student Yi Ting Ter is investigating the male coutship behavior of yellow mutant B. anynana butterflies comparatively to Wt male behavior.
Honors student Dean Tin Shuen Yew is working on sexual imprinting in Asian human males and females.
Laboratory technologist, Lu Wee Tan, is in charge of providing safety training to all lab members, as well as ordering consumables, equipment, etc.




Former lab members




Master student Zohara Rafi conducted field tests of predation on Bicyclus butterfies to test the role of UV-patterns and eyespot number on attack risk in the wild.
Honors student Ling Sheng Loh investigated the role of Armadillo in eyespot and spot development using CRISPR-Cas9.
Honors student Sharran Rajendran investigated the role of a novel Distal-less CRE in wing and eyespot development using CRISPR-Cas9.
Visiting student, Lin Naing, from Monash University, tested how variation in eyespot number in B. anynana affects predation risk by praying mantids in mesocosmos experiments.
PhD student Shivam Bhardwaj compared mechanisms of eyespot plasticity across nymhalid butterflies. He explored the developmental/molecular origin of eyespot size plasticity.
Visiting undergraduate student, Gowri V, worked on the transmission of novel learned food odors to the next generation in Bicyclus anynana larvae.
Visiting undergraduate student, Athmaja, worked on the transmission of novel learned food odors to the next generation in Bicyclus anynana larvae.
Postdoctoral fellow Eunice Tan investigated the set of genes that are differentially expressed in male pupal brains of wet and dry season forms, as well as hormone-manipulated brains, and identified candidates that may cause the different sexual behaviors of these males.
Master's student (and former Honors student) Li Xian Pui showed that Bicyclus anynana females that learn to prefer new pheromone blends pass on that preference to their female offpring. She also explored whether epigenetic inheritance is involved in this process.
Honors student Jeremy Pang tested the role of a sex pheromone odor receptor in sexual behavior using CRISPR-Cas9.
Honors student Khai Ann Hiew tested whether the false 3D head of the Common Red Flash butterfly, Rapala iarbus iarbus, may give it a protective advantage in interactions with invertebrate mantid predators
Postdoctoral fellow Kathy Su was interested in the role of doublesex in promoting the development of a novel male-specific sexual organ in the abdomen of sepsid flies. Kathy had her own independent funding (She was a Lee Kuan Yew fellow).
Honors student Yue Qian Tan, tested the effect of "haze" on butterfly growth rate and survival.
Undergraduate student Ling Sheng Loh investigated the temporal and spatial expression of wingless genes on developing butterfly wings.
Honors student Xiang Hui Low was interested in undertsanding the process that restricts mimicry of distatesful models to females of Papilio polytes butterflies. He tested whether P. polytes females differed in their mate preferences towards artificially colored mimetic males and typical non-mimetic males.
Honors student Brent Tan tested the role of the banding pattern and the head-to-tail mimicry in the Banded Swallowtail butterfly (Papilio demolion demolion) with paper models in the wild.
Research Assistant Jonathan Peh Jun Jie tested the role of three candiate genes in the regulation of eyespot number variation (between Spotty mutants and Wildtype individuals) using CRISPR-Cas9.
PhD student Nesibe Ozsu identified members of the eyespot gene regulatory network during early pupation using RNA-seq. She also tested the role of wingless in eyespot development using transgenenic lines.
Undergraduate student Tricia Loo investigated the process of eyespot center differentiation using computer modeling.
Visiting undergraduate student Sarah Monroe tested the role of the gene Spalt in eyespot and wing development in Bicyclus anynana butterflies.
Postdoctoral fellow Arjen van 't Hof mapped the mutation, Spotty, that introduces two extra eyespots on the wings of B. anynana butterflies.
Masters student Jelle van Creij, visiting from The Netherkands, tested the role of Distal-less in eyespot development using CRISPR-Cas9
Honors student Swit Yee Ng tested how dry season males invest in reproduction when mated with females with manipulated sexual ornaments.
Honors student Jocelyn Wee tested whether different colors on the wings of the Painted Jezebel (orange, yellow, white and black) function as aposematic colors using manipulated paper models in field experiments.
Honors student Kenneth Neo tested whether learned visual mate preferences lead to different levels of opsin expression in B. anynana females.
Honors student Qian Yi Chan tested the role of knocking-down the signaling ligand wingless on eyespot size in Bicyclus anynana butterflies.
Honors student Shearel Sim tested the function of candidate cis-regulatory elements for eyespot genes using CRISPR-Cas9
Undergraduate student, Soo Yee Ng, tested the role of the conspicuous wing flaps in the Red Flash butterfly using mantid predators and paper models of the butterflies.
Undergraduate student, Yue Qian Tan, tested the predation risk of mimetic and non-mimetic wing patterns in Papilio polytes butterflies using paper models in the field.
Postdoctoral fellow Luqman Aslam mapped the genomic region responsible for a eyespot number variant of Bicyclus anynana, Spotty, with four eyespots on the forewing instead of the usual two eyespots, using RAD-Tags.

Postdoctoral fellow Katy Prudic tested why males and females both have "ornaments", i.e., eyespots, on the hidden surfaces on their wings. She showed that both sexes display these eyespots to the opposite sex during courtship. This behavior is phenotypically plastic, however, and wired during larval/pupal development. Katy also studied the role of insect predators in maintaining the phenotypically plastic ventral wing patterns in Bicyclus anynana and examined the physiological mechanisms underlying dorsal eyespot size and brightness plasticity.
Postdoctoral fellow Mainak Das Gupta identified a series of putative cis-regulatory elements of eyespot genes expressed during the larval and pupal stages of development using FAIRE-Seq.
Honors student Zhe Ching Ngan showed that mantid predators have a different attack behavior toward butterflies with different eyespots on the forewing ventral surface.
Visting undergraduate student Sam Chan documented the loss of green fluorescence in the eyes of a transgenic line of Bicyclus anynana butterflies with adult age.
Honors student Melissa Teo Hui Jing worked on the molecular basis of sexual dimorphism in butterflies.
Honors student Sebastian Ho worked on the function of eyespots in the field using natural predators. He tested how eyespot size and eyespot number alters eyespot function from intimidating to attracting predators.
Undergraduate student Jonathan Peh Jun Jie tested whether learned preferences can be passed on to the next generation.
Postdoctoral fellow Bethany Wasik worked on an artificial selection experiment to produce violet structural colors on the wings of B. anynana. She also worked on the positional cloning of an eyespot network "switch" locus, Spotty, using RAD-tag markers.
Undergraduate student Stacey Yuen tesed the function of a very interesting wing pattern in a local moth species using manipulations in paper models and natural predators.
Graduate student Ashley Bear explored the (neuro)physiological and genetic basis of courtship reversals induced by environmental temperature in male and female Bicyclus anynana. She previously worked on the physiological/behavioral traits of a dark larval color mutation also in Bicyclus .
April Dinwiddie Graduate student April Dinwiddie worked on the development of structural colors in butterfly wings. She is also interested in exploring the evolution and development of wing scales across insects.
Undergraduate student Chris Tokita performed a survey of eyespot sexual dimorphism across 450 species of Nymphalid butterflies and discovered that females, on average, have more eyespots than males, and that "hidden" dorsal and forewing surfaces are disproportionately more dimorphic than "exposed" ventral hindwing surfaces.
Graduate student Erica Westerman worked on the role of learning in shaping female preference for novel male wing patterns. She showed that females don't have fixed preferences for males with particular wing patterns and can learn to prefer males with novel patterns. She also showed that smell is an important mediator of visual learning.
Postdoctoral fellow Xiaoling Tong tested the function of the hox gene Ultrabithorax in regulating eyespot patterns in Bicyclus anynana. She used transgenic tools to down-regulate Ubx at different times during development. Xiaoling also described the expression of Ecdysone Receptor (EcR) on the wings of Bicyclus anynana.
Postdoctoral fellow Jeffrey Oliver mapped the evolution of eyespots on the phylogeny of nymphalid butterflies. He also did comparative gene expression work in a variety of nymphalid species to understand the evolution of the eyespot developmental gene network. He discovered that eyespots evolved once, at the base of the nymphalid clade, and that many of the genes associated with early eyespot development were probably co-opted as a cluster, from a preexisting network.
Graduate student Karin van der Burg completed her MS degree in the lab. She examined how the expression of Wingless and Decapentaplegic were alterered by over-expression of Ultrabithorax in early pupal wings.
Andrew Everett Undergraduate student, Andrew Everett described eye sexual dimorphism and morphological plasticity in Bicyclus anynana across a temperature gradient.
Joseph Walker Visiting undergraduate student, Joseph Walker, from Purdue University worked on black spot formation in the Cabbage White Butterfly, using classic cell ablation techniques in live pupal wings.
Postdoctoral fellow Andrew Stoehr developed functional genetic tools for the Pieris rapae in order to understand the developmental basis of phenotypic plasticity in this species. He also performed comparative gene expression work across the Pieridae.
Visiting Ms student, Carole Bastianelli, from Lyon, France worked on describing the expression dynamics of the Ecdysone Receptor in Bicyclus wings of the wet and dry seasonal forms. This work will help us understand how the endocryne system evolved to regulate a classic seasonal polyphenism in a butterfly wing pattern.
Undergraduate student Anna Lindemann worked on the role of the hedgehog signaling pathway in differentiating eyespot centers.
Paleontology graduate student Joanna Wolfe is interested in understanding the evolution of the short anterior pair of legs of nymphalid butterflies (the brushfoots), both at the molecular level and through the fossil record.
Paul Shamble photographed most of the nymphalid butterflies in the collection at the Peabody Museum (~1500 species). This will allow us to answer many questions about the evolution of wing patterns in this group using the comparative method.
Postdoctoral fellow Ondrej Podlaha tested the function of the hox gene Ultrabithorax in regulating eyespot patterns in Bicyclus anynana. He used transgenic tools to over-express Ubx at different times during development and characterized both Ubx mRNA expression and the adult wing pattern phenotype. He is interested in molecular evolution and in testing ideas of gene network co-option at the empirical level.
Rotating graduate student Andrea Hodgins-Davis worked on the role of prior experience in female mate choice.
Undergraduate student Robert Tunney worked on the developmental evolution of eyespots in local CT nymphalid butterfly species.
Undergraduate student, Ariel Simons, from the University of Rochester, worked out the genetics of a new spontaneous mutation in our stock that produces a dark larval cuticle. She visited us during the summer of 2008.
Undergraduate student Lee Driftmier tested the role of the sexually dimorphic black spots on the wings of Pieris rapae in the context of female choice.
Graduate student Diane Ramos worked on the development of a laser-mediated heat-shocking mechanism that enables the spatial control of transgene expression on the pupal wing. She cloned several copies of the Engrailed gene and tested their localization using in-situs. She tested the role of over-expressing the transcription factor Spalt in vein and eyespot development.
Visiting scientist, Bin Chen, cloned a series of transcription factors from B. anynana in order to test their function in wing and color pattern development. He also developed new piggyBac vectors for both ectopic expression as well as gene knockdown via transgenic RNAi.
Graduate student Kendra Roberston tested whether color pattern differences on the wings of males have an effect on female preference.

Kendra also mapped the evolution of wing patterns of 54 of the 80 members of the genus Bicyclus on the phylogeny of the genus.

Jessica Decker was a visiting student from the Concordia College Moorhead in Minnesota who worked in the lab during the Summer 2007 on sexual selection in Pieris butterflies.
Lisa Fazzino was a visiting high-school student from Wallinford, CT, who helped us gather functional data for some transgenic lines of Bicyclus. She and Jessica also worked together on developmental perturbations of Pieris wing patterns.
Undergraduate student Mark Fisher tested the role of behavioral plasticity in female choice in Bicyclus anynana
Graduate student, Gary Glaser, cloned the wingless gene from B. anynana and determined that this gene is expressed in the eyespot centers in developing pupal wings using antibody stainings.
Graduate student, Firdous Kamal, from the Department of Electrical Engineering, developed a new laser system in order to heat-shock the pupal wing epidermis of Bicyclus anynana without killing the cells.
Graduate student, Katie Costanzo, worked on the relative role that male phenoromes and visual signals play in female choice in Bicyclus anynana.
Graduate student, Kyle Golden, worked on developing a "fast" method to introduce genes into wing epidermal cells, using in-vivo electroporation.
Min Undergraduate student, Min Guo, worked on testing the role of Distal-less in eyespot development.
Undergraduate student, Andrew Goldman, continued working on the color learning ability of young virgin female butterflies, and how early experience with particular color patterns influences subsequent mate choice.
Undergraduate student, Taid Rahimi, worked on the color learning ability of young virgin female butterflies, and how early experience with particular color patterns influences subsequent mate choice.
Undergraduate student, Veena Raju, helped develop a "fast" method to introduce genes into wing epidermal cells, using in-vivo electroporation.
Graduate student, Sara Kremmer, worked on an educational-research project. She helped develop a new course syllabus for the Evolutionary Genetics course that integrates quiz questions, homework and class exercises to improve student learning.
Undergraduate student, Sam Arbesman, worked on a web based animation (Ancient Wings) that reconstructs the putative ancestral wing patterns of 54 of the 80 species of Bicyclus butterflies, and morphs these patterns across the phylogenetic tree of Bicyclus.
Undergraduate student, Steven Stockslager, worked on the effect of wing damage on the activation of certain transcription factors around the site of damage. Damage ultimately results in the production of an ectopic eyespot on the wings.
Undergraduate honors student, Nathan Markwarth, worked on developing a "fast" method to introduce genes into wing epidermal cells, using in-vivo electroporation.
Undergraduate student, Laura Blodgett, visiting from Rochester University, worked with paper models of Bicyclus butterflies to determine to what extent males detect conspecifc females based on visual cues.
Undergraduate student, Jarod Masci, developed a new protocol to dechorionate eggs and visualize the early embryos of B. anynana. He co-authored a paper, as first author, in the journal Zygote.
Undergraduate student, Sashti Balasundaram, together with rotating graduate student, Nathan Kirk, developed a protocol for artificially inseminating B. anynana butterflies, using whole spermatophore transfers into virgin females.
Undergraduate student, Laura Falkowski, tested whether the presence of extra eyespots on the dorsal surface of the wings of B. anynana males is more attractive to females relative to the wild type pattern.
Undergraduate student, Lauren Scott, tested the effects of two mutations, Missing and Spotty, when in homozygote and heterozygote condition, and when present simultaneously in the same genome. She found that each mutation is codominant relative to wildtype and influences both the fore and hindwing pattern of Bicyclus anynana. These mutations also display an epistatic effect when present together in the genome.
Jeff_Marcus Postdoctoral Fellow Jeffrey Marcus developed the first line of transgenic butterflies.