
Publications






2025
The Developmental Origin of Novel Complex Morphological Traits in Lepidoptera
Monteiro, A., Murugesan, S. N., Prakash, A., & Papa, R.​
We explore the developmental and genetic mechanisms underlying the origin of novel complex morphological traits in Lepidoptera, highlighting how evolutionary changes shape wing patterns and structures.
Validity evidence for the translation and cultural adaptation of the measure of acceptance of the theory of evolution in Español (MATE-E)
Pérez-Vega, Á. E., Papa, R., & Borrero, M.
This paper validates the Spanish adaptation of the Measure of Acceptance of the Theory of Evolution (MATE-E), ensuring its cultural relevance and reliability for assessing evolution acceptance in Spanish-speaking populations.
2024
Transforming in-service high school biology teachers’ acceptance of evolution through professional development
Pérez-Vega, Á. E., Papa, R., & Borrero, M.
This paper examines how professional development programs can enhance in-service high school biology teachers' acceptance and understanding of evolution, focusing on initiatives led by the Papa Lab in Puerto Rico
A long noncoding RNA at the cortex locus controls adaptive coloration in butterflies
Livraghi, L., Hanly, J. J., Evans, E., et al.
We identified a long noncoding RNA (lncRNA) named 'ivory' within the cortex locus that regulates adaptive coloration in butterflies by orchestrating transitions between dark and light wing patterns during development.
Many functionally connected loci foster adaptive diversification along a neotropical hybrid zone
Lewis, J. J., Van Belleghem, S. M., Papa, R., Danko, C. G., & Reed, R. D.
We examined the genetic architecture underlying adaptive diversification in Heliconius butterflies, revealing that multiple functionally connected loci, regulated by the master gene Optix, collectively maintain distinct red color pattern phenotypes in natural populations
Recolonization of secondary forests by a locally extinct Caribbean anole through the lens of range expansion theory
Acevedo, M. A., Fankhauser, C., Gonzalez, L., Quigg, M., Gonzalez, B., & Papa, R.
We examined the recolonization of secondary forests in Puerto Rico by the shade-specialist anole (Anolis gundlachi), applying range expansion theory to understand the ecological processes involved.
2023
Evolutionary dynamics of genome size and content during the adaptive radiation of Heliconiini butterflies
Edelman, N. B., Frandsen, P. B., Miyagi, M. T., Clavijo, B., Mallet, J., & Papa, R.
We investigated the evolutionary dynamics of genome size and content during the adaptive radiation of Heliconiini butterflies, highlighting the role of hybridization and introgression in their diversification.
SARS-CoV-2 Omicron Replacement of Delta as Predominant Variant, Puerto Rico
Reich, N. G., McGibbon, A. R., Miller, B. W. R., et al.
We analyzed the genomic evolution of SARS-CoV-2 in Puerto Rico from October 2021 to May 2022, demonstrating that the Omicron BA.1 variant emerged and supplanted the Delta variant as the predominant strain by December 2021.
High level of novelty under the hood of convergent evolution
Van Belleghem, S. M., Ruggieri, A. A., Concha, C., et al
We investigated the mechanisms of convergent evolution, highlighting how different evolutionary pathways can lead to the development of similar traits, thereby providing insights into the processes that drive novelty in evolutionary biology.
Recolonization of secondary forests by locally extinct fauna through the lens of range expansion:
Four open questions
Acevedo, M. A., Fankhauser, C., & Papa, R.
We explored the recolonization of secondary forests by locally extinct fauna through the lens of range expansion, posing four open questions to guide future research in this area.
2022
Widespread Gene Expression Divergence in Butterfly Sensory Tissues Plays a Fundamental Role During Reproductive Isolation and Speciation
Wu, N., Evans, E., van Schooten, B., et al.
We investigated the role of gene expression divergence in butterfly sensory tissues—antennae, legs, and mouthparts—in reproductive isolation and speciation among Heliconius butterflies.
A butterfly pan-genome reveals that a large amount of structural variation underlies the evolution of chromatin accessibility
Ruggieri, A. A., Livraghi, L., Lewis, J. J., et al.
We examined the butterfly pan-genome, revealing that substantial structural variation contributes to the evolution of chromatin accessibility
2021
Balanced polymorphisms and their divergence in a Heliconius butterfly
Ogilvie, J. G., Van Belleghem, S. M., Range, R. R., et al.
We examined the selective pressures maintaining polymorphic mimicry in Heliconius doris, highlighting the complexity of positive frequency-dependent selection and its impact on interspecific variation of mimetic warning coloration.
Novel Doublesex Duplication Associated with Sexually Dimorphic Development of Dogface Butterfly Wings
Rodriguez-Caro, F., Fenner, J., Bhardwaj, S., et al.
We investigated a novel duplication of the doublesex (dsx) gene in the Southern Dogface butterfly (Zerene cesonia), revealing its role in sexually dimorphic wing coloration through sex-specific expression patterns.
Selection and isolation define a heterogeneous divergence landscape between hybridizing Heliconius butterflies
Van Belleghem, S. M., Cole, J. M., Montejo-Kovacevich, G., et al.
We investigated the divergence between hybridizing Heliconius butterflies, highlighting how selection and isolation shape their genomic landscapes.
Heliconius butterflies: A window into the evolution and development of diversity.
Van Belleghem, S. M., Lewis, J. J., Papa, R., et al.
This review explores the evolution and development of diversity in Heliconius butterflies, emphasizing their role as models for studying phenotypic variation and speciation mechanisms.
Cortex cis-regulatory switches establish scale colour identity and pattern diversity in Heliconius
Livraghi, L., Ruggieri, A. A., Lewis, J. J., et al.
We identified how cortex cis-regulatory switches control scale color identity and contribute to the pattern diversity observed in Heliconius butterflies.
2020
Many functionally connected loci foster adaptive diversification along a neotropical hybrid zone
Lewis, J. J., Van Belleghem, S. M., Papa, R., Danko, C. G., & Reed, R. D.
We examined how multiple functionally connected loci contribute to adaptive diversification in Heliconius butterflies along a neotropical hybrid zone.
Visual mate preference evolution during butterfly speciation is linked to neural processing genes
Rossi, M., Hausmann, A. E., Thurman, T. J., et al.
We identified five genes linked to divergent visual preferences in Heliconius butterflies, suggesting that changes in visual processing contribute to speciation through modified mate selection behaviors.
Perfect mimicry between Heliconius butterflies is constrained by genetics and development
Van Belleghem, S. M., Alicea Roman, P. A., Papa, R., Jiggins, C. D., & McMillan, W. O.
We explored how genetic and developmental constraints limit the perfect mimicry between Heliconius butterfly species, despite strong selective pressures favoring convergent wing patterns.
Divergence of chemosensing during the early stages of speciation
Papa, R., Jiggins, C. D., McMillan, W. O., & Counterman, B. A.
We investigated the divergence of chemosensory genes between Heliconius melpomene and Heliconius cydno butterflies, suggesting that chemosensory differences play a role in early speciation stages.
Wnt Genes in Wing Pattern Development of Coliadinae Butterflies
Fenner, J., Benson, C., Rodriguez-Caro, L., et al.
We examined the role of Wnt genes in wing pattern development of Coliadinae butterflies, revealing conserved gene content but divergent expression patterns compared to nymphalid butterflies.
Selection and gene flow define polygenic barriers between incipient butterfly species
Van Belleghem, S. M., Cole, J. M., Montejo-Kovacevich, G., et al.
We examined how selection and gene flow establish polygenic barriers between incipient Heliconius butterfly species, utilizing demographic modeling and simulations to infer that periods of isolation and selection have led to genome-wide correlation patterns affecting recombination rates and divergence.
Multiple Loci Control Eyespot Number Variation on the Hindwings of Bicyclus anynana Butterflies
Rivera-Colón, A. G., Westerman, E. L., Van Belleghem, S. M., et al.
We identified multiple genetic loci influencing eyespot number variation on the hindwings of Bicyclus anynana butterflies, highlighting a polygenic architecture with moderate heritability.
​Next generation sequencing panel in undifferentiated autoinflammatory diseases identifies patients with colchicine-responder recurrent fevers
Papa, R., Rusmini, M., Volpi, S., et al.
We evaluated a next-generation sequencing (NGS) panel comprising 41 genes in patients with undifferentiated systemic autoinflammatory diseases (uSAID), identifying a subgroup of patients with recurrent fevers responsive to colchicine treatment.