Shirey, V., N. Neupane, R. Guralnick, & L. Ries. (2024). Rising minimum temperatures contribute to 50 years of occupancy decline among cold‐adapted Arctic and boreal butterflies in North America. Global Change Biology, 30(2), e17205.

Larsen, E.A., M.W. Belitz, G.J. Di Cecco, J. Glassberg, A.H. Hurlbert, L. Ries and R. P. Guralnick. (In press) Overwintering strategy regulates phenological sensitivity and consequences for ecological services in a clade of temperate North American insects. Functional Ecology

Farr, M. T., Zylstra, E. R.,Ries, , & Zipkin, E. F. (2024). Overcoming data gaps using integrated models to estimate migratory species' dynamics during cryptic periods of the annual cycle. Methods in Ecology and Evolution

K. Wheeler, M. Dietze, D. LeBauer, J. Peters, A.D. Richardson, A. Ross, R.Q. Thomas, K. Zhu, U. Bhat, S. Munch, R.F. Buzbee, M. Chen, B. Goldstein, ... D. Pal, V. Shirey, Y. Song, ..., and L. Zachmann. (2024). ”Predicting spring phenology in deciduous broadleaf forests: NEON phenology forecasting community challenge.” Agricultural and Forest Meteorology 345.

Abarca, M., Parker, A.L., Larsen, E.A., Umbanhowar, J., Earl, C., Guralnick, R.P., Kingsolver, J., and Ries, L. (2024) How development and survival combine to determine the thermal sensitivity of insects. PLoS ONE 19(1): e0291393.

Supplementary info: List of species and location of populations included in analyses.: File Abarca et al 2024 supplementary S1 journal.pone_.0291393.s001.docx (323.41 KB)
Supplementary info: Model results from the “analytical” and “full” data sets: File Abarca et al 2024 supplementary info S2 journal.pone_.0291393.s002.docx (24.29 KB)
Supplementary info: Phylogenetic analyses: File Abarca et al 2024 PlosOne supplementary info S3 journal.pone_.0291393.s003.docx (871.63 KB)
Supplementary info: Models with phylogenetic correction: File Abarca et al 2024 PlosOne supplementary info S4 journal.pone_.0291393.s004.docx (1.34 MB)

Neupane, N., E.A. Larsen, and L.Ries (Online early). Ecological forecasts of insect range dynamics: a broad range of taxa include winners and losers under future climate. Invited paper at Current Opinion in Insect Science


V. Shirey and L. Ries. (2023). Population dynamics and drivers of the eastern monarch (Danaus plexippus) across its full annual cycle: a cross-scale synthesis of a model migratory species. Current Opinion in Insect Science 60.

Rippel, T. M., Minsavage-Davis, C. D., Shirey, V., & Wimp, G. M. (2023). Simple Machine Learning with Aerial Imagery Reveals Severe Loss of a Salt Marsh Foundation Species. Estuaries and Coasts, 46(4), 1110-1122.

Stevens, H. C., Smith, A. C., Buechley, E. R., Şekercioğlu, Ç. H., Shirey, V., Rosenberg, K. V., ... & Marra, P. P. (2023). Species-specific ecological traits, phylogeny, and geography underpin vulnerability to population declines for North American birds. Ornithological Applications, duad046.

Courter, JN, Z Liu, N Neupane, A Arab, J Siegrist (2023). Satellite-based environmental variables complement traditional variables in spatio-temporal models of purple martin migration. Wildlife Research

Shirey, V. and L. Ries (In press) Population dynamics and drivers of the eastern monarch (Danaus plexippus) across its full annual cycle: a cross-scale synthesis of a model migratory species. Invited paper at Current Opinion in Insect Science

Belitz, M. W., Larsen, E. A., Shirey, V., Li, D., & Guralnick, R. P. (2023). Phenological research based on natural history collections: Practical guidelines and a lepidopteran case study. Functional Ecology, 37(2), 234-247.

Kawahara, A. Y., Storer, C., Carvalho, A. P. S., Plotkin, D. M., Condamine, F. L., Braga, M. P., ... Larsen, E.A., .... Shirey, V., ... Ries, L., ... & Lohman, D. J. (2023). A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins. Nature Ecology & Evolution, 1-11.

Di Cecco, G.J., M.W. Belitz, R.J. Cooper, E.A. Larsen*, W.B. Lewis, L. Ries, R.P. Guralnick, and A.H. Hurlbert. (2023) Phenology in adult and larval Lepidoptera from structured and unstructured surveys across eastern North America. Frontiers in Biogeography 15(1).


Zylstra, E. R., Neupane, N., & Zipkin, E. F. (2022). Multi‐season climate projections forecast declines in migratory monarch butterflies. Global Change Biology, 28(21), 6135-6151.

Palacio, F. X., Callaghan, C. T., Cardoso, P., Hudgins, E. J., Jarzyna, M. A., Ottaviani, G., ..., Shirey, V. & Mammola, S. (2022). A protocol for reproducible functional diversity analyses. Ecography, 2022(11), e06287

Vaughn Shirey, Elise Larsen, Andra Doherty, Clifford A. Kim, Faisal T. Al-Sulaiman, Jomar D. Hinolan, Micael Gabriel A. Itliong, Mark Arcebal K. Naive, Minji Ku, Michael Belitz, Grace Jeschke, Vijay Barve, Gerardo Lamas, Akito Y. Kawahara, Robert Guralnick, Naomi E. Pierce, David J. Lohman, Leslie Ries. (2022) Lep Traits 1.0: Butterfly Traits 1.0: A globally comprehensive dataset of butterfly traits. Scientific Data 9:382

Larsen, E. A., Belitz, M. W., Guralnick, R. P., & Ries, L. 2022. Consistent Trait-Temperature Interactions Drive Butterfly Phenology in Both Incidental and Survey Data. Scientific Reports, 12(1), 1-10.

Neupane, N., M. Peruzzi, A. Arab, S.J. Mayor, J.C. Withey, L. Ries, and A.O. Finley. (2022). A novel model to accurately predict continental-scale timing of forest green-up. International Journal of Applied Earth Observation and Geoinformation. 108: 102747

M. Belitz, E. Larsen,V. Shirey, D. Li, and R. Guralnick. 2022. ”Phenological research based on natural history collections: practical guidelines and a Lepidopteran case study.” In Review.

V. Shirey, R. Khelifa, L.K. M’Gonigle, and M.L. Guzman. 2022. ”Utilizing occupancy-detection models with museum specimen data: promise and pitfalls.” Methods in Ecology and Evolution (Online early).

Neupane, N., E. Zipkin, S. Saunders, and L. Ries. 2022. Grappling with uncertainty in ecological projections for the migratory monarch butterfly using tailored climate models. Ecosphere


Neupane, N., Goldbloom-Helzner, A., & Arab, A. (2021). Spatio-temporal modeling for confirmed cases of lyme disease in Virginia. Ticks and Tick-borne Diseases, 12(6), 101822.

Belitz, M. W., Barve, V., Doby, J. R., Hantak, M. M., Larsen, E. A., Li, D., ... & Stucky, B. J. (2021). Climate drivers of adult insect activity are conditioned by life history traits. Ecology Letters, 24(12), 2687-2699.

Shirey, V., Carvalho, A. P. S., & Kawahara, A. Y. (2021). Early Evidence for Sexually Dimorphic, Ultraviolet Eyespots in Parnassius smintheus. The Journal of the Lepidopterists' Society, 75(3), 215-217.

Abarca, M., & Spahn, R. (2021). Direct and indirect effects of altered temperature regimes and phenological mismatches on insect populations. Current opinion in insect science, 47, 67-74.

Zylstra, Erin R., Leslie Ries, Naresh Neupane, Sarah P. Saunders, M. Isabel Ramírez, Eduardo Rendón-Salinas, Karen S. Oberhauser, Matthew T. Farr, and Elise F. Zipkin. "Changes in climate drive recent monarch butterfly dynamics." Nature Ecology & Evolution (2021): 1-12.

Report of research findings in news section of Nature Ecology and Evolution: PDF icon Bowler_2021_NEE_Review.pdf (932.48 KB)
Supplementary files: PDF icon Zylstra_etal_2021_ClimateDrivesMonarchPopulationDynamics_Supp.pdf (555.86 KB)

Larsen, E.A., V. Shirey. 2021. Method matters: pitfalls in analysing phenology from occurrence records. Ecology Letters 24:1287-1289

Degroot, D., Anchukaitis, K., Bauch, M. ..., Neupane, N., Newfield, T, et al. Towards a rigorous understanding of societal responses to climate change. Nature 591, 539–550 (2021).

Shirey, V., M. Belitz, V. Barve, R. Guralnick. 2021. "A complete inventory of North American butterfly occurrence data: narrowing data gaps, but increasing bias." Ecography 44(4), 537-547.

Mushegian, A., N. Neupane, Z. Batz, M. Mogi, N. Truno, T. Toma, I. Miyagi, L. Ries and P. Armbruster (2021). Letter: Ecological mechanism of climate-mediated selection in a rapidly evolving invasive species. Ecology Letters 24:698-707. doi: 10.1111/ele.13686


Bried, J. L. Ries, B. Smith, M. Patten, J. Abbott, J. Ball-Damerow, R. Cannings, A. Cordero-Rivera, A. Córdoba-Aguilar, P. De Marco Jr., K. Dijkstra, A. Dolný, R. van Grunsven, D. Halstead, F. Harabiš, C. Hassall, M. Jeanmougin, C. Jones, L. Juen, V. Kalkman, G. Kietzka, C. Searles Mazzacano, A. Orr, M. Perron, M. Rocha-Ortega, G. Sahlén, M. Samways, A. Siepielski, J. Simaika, F. Suhling, L. Underhill, and E. White. In press. Towards global volunteer monitoring of dragonfly abundance. Bioscience

Belitz, M.W., E.A.Larsen*, L.Ries and R.P. Guralnick. 2020. The accuracy of phenology estimators for use with sparsely sampled presence-only observations. Methods in Ecology and Evolution 11:1273–1285

Mainali*, K., T. Hefley, L. Ries, and W.F. Fagan. 2020. Matching expert range maps with species distribution model predictions. Conservation Biology 24:1292-1304.

Campbell, D., A.Thessen, and L. Ries. 2020. A novel curation system to facilitate data integration across regional citizen science survey programs. PeerJ 8:e9219

Abarca, M., Lill, J. T., & Weiss, M. R. (2020). Host Plant and Thermal Stress Induce Supernumerary Instars in Caterpillars. Environmental entomology, 49(1), 123-131.


Abarca, M. (2019). Herbivore seasonality responds to conflicting cues: Untangling the effects of host, temperature, and photoperiod. PLoS ONE, 14(9).

Wepprich T, Adrion JR, Ries L, Wiedmann J, Haddad NM (2019) Butterfly abundance declines over 20 years of systematic monitoring in Ohio, USA. PLoS ONE 14(7): e0216270.

Supplemental Appendix: File Wepprich et al 2019 PLoS ONE Supplemental.docx (308.98 KB)

Ries, L., E. Zipkin, R. Guralnick. 2019. Tracking trends in monarch abundance over the 20th century is currently impossible using museum records. Proceedings of the National Academy of Sciences Jul 2019, 116 (28) 13745-13748; DOI: 10.1073/pnas.1904807116

Abarca, M., E.A. Larsen, and L. Ries. (2019). Heatwaves and Novel Host Consumption Increase Overwinter Mortality of an Imperiled Wetland Butterfly. Front. Ecol. Evol. 7: 193. doi: 10.3389/fevo.

Saunders, S.P., L. Ries, N. Neupane, M. I. Ramirez, E.Garcia-Serrano, E. Rendon-Salinas, E.F. Zipkin. 2019. Multi-scale seasonal factors drive the size of winter monarch colonies. Proceedings of the National Academy of Sciences. 201805114

Wimp, G., L. Ries, D. Lewis, and S. Murphy. In Press. Habitat edge responses of generalist predators are predicted by prey and structural resources. Ecology


Ries, L., N. Neupane, K. A. Baum, E. F. Zipkin. (2018). Flying through hurricane central: impacts of hurricanes on migrants with a focus on monarch butterflies. Animal Migration 5(1): 94-103..

Abarca, M., E. Larsen, J.T. Lill, M. Weiss, E. Lind, L. Ries. 2018. Inclusion of host quality data improves predictions of herbivore phenology. Entomologia Experimentalis et Applicata 166 (8), 648-660

Fletcher Jr, R. J., Didham, R. K., Banks-Leite, C., Barlow, J., Ewers, R. M., Rosindell, J., ... Ries, L.,...& Melo, F. P. (2018). Is habitat fragmentation good for biodiversity?. Biological Conservation, 226, 9-15.

Semmens, J.E. Diffendorfer, K. Bagstad, R. Wiederholt, K. Oberhauser, L. Ries, B. Semmens and J. Goldstein J. Loomis, W.E. Thogmartin, B.J. Mattsson, L.Lopez-Hoffman. Online early. Quantifying ecosystem service flows at multiple scales across the range of a long-distance migratory species. Ecosystem Services ong-distance migratory species. Ecosystem services, 31:255-264.

Saunders, S., L. Ries, K. Oberhauser, W. Thogmartin, E. Zipkin. 2018. Local and cross-seasonal effects of climate and land-use on migratory monarch butterflies. Ecography. 41 (2), 278-290


Ries, L., S.M. Murphy, G.M. Wimp, R.J. Fletcher. 2017. Closing persistent gaps in knowledge about edge ecology. Invited review in Current Landscape Ecology Reports 2:30-41.

Oberhauser, K., R. Wiederholt, J. E. Diffendorfer, D. Semmens, L. Ries, W. E. Thogmartin, L.A. Lopez‐Hoffman, B. Semmens. 2017. A trans‐national monarch butterfly population model and implications for regional conservation priorities. Ecological Entomology, 42(1), 51-60.


Schmucki R., Pe’er G., Roy D.B., Stefanescu C., van Swaay C.A.M., Oliver T.H., Kuussaari M., van Strien A.J., Ries L., Settele J., Musche M., Carnicer J., Schweiger O., Brereton T., Harpke A., Heliola J., Kuhn E., and R. Julliard. 2016. Regionally informed abundance index for supporting integrative analyses across butterfly monitoring schemes. Journal of Applied Ecology. 53:501-510.

Murphy, S.M., A.H. Battocletti, R.M. Tinghitella, G.M. Wimp, and L. Ries. 2016. Complex community and evolutionary responses to habitat fragmentation and habitat edges: what can we learn from insect science? Invited paper in Current Opinion in Insect Science 14:61-65.

Saunders, S.P., L. Ries, K.S. Oberhauser, and E.F. Zipkin. 2016. Evaluating confidence in population-level predictions from climate impacts: Summer abundances of the monarch butterfly. Global Ecology and Biogeography 25:1000-1012.

Thorson, J.T., J.N. Ianelli, E.A. Larsen, L. Ries, M.D. Scheuerell, C. Szuwalski, and E.F. Zipkin. 2016. Joint dynamic species distribution models: a tool for community ordination and spatio‐temporal monitoring. Global Ecology and Biogeography.


Oberhauser, K., L. Ries, S. Altizer, R. Betalden, J. Ekstrum, M. Garland, E. Howard, S. Jepsen, J. Lovett, M. Monroe, G. Morris, E. Rendon, R. RuBino, A. Ryan, C. Taylor, R. Trevino, F. Villablance, D. Walton. 2015. Citizen scientists and an iconic insect: 70 years and counting in (K. Oberhauser, ed.) Monarchs in a changing world: Biology and conservation of an iconic butterfly. Cornell University Press. Ithaca, NY.

Ries, L., K. Oberhauser, D. Taron, E. Rendon-Salinas. 2015. Connecting eastern monarch population dynamics across their migratory cycle. Invited book chapter in (K. Oberhauser, ed.) Monarchs in a changing world: Biology and conservation of an iconic butterfly. Cornell University Press. Ithaca, NY.

Taron, D. and L. Ries. 2015. Butterfly monitoring for Conservation. Invited book chapter in (J. Daniels, editor) Butterfly Conservation in North America. Springer Press.

Ries, L., E. Rendon and D. Taron. 2015. The disconnect between summer and winter monarch trends for the Eastern migratory population: possible links to differing drivers. Invited paper in Annals of the Entomological Society of America (Online early).

Cayton, H, N. Haddad, K. Gross, S. E. Diamond and L. Ries. 2015. Do growing degree days predict phenology across butterfly species? Ecology 96: 1473–1479

Ries, L. and K. Oberhauser. 2015. A Citizen-Army for Science: Quantifying the Contributions of Citizen Scientists to our Understanding of Monarch Butterfly Biology. Bioscience 65:419-430.


Diamond, S.E., H. Lessig*, T. Wepprich, C.N. Jenkins, R.R. Dunn, N.M. Haddad, and L. Ries. 2014. Unexpected phenological responses of butterflies to the interaction of urbanization and geographic temperature. Ecology 95:2613-2621.

Diffendorfer, J.E., J.B. Loomis, L. Ries, K. Oberhauser, L. Lopez-Hoffman, D. Semens, 79B. Semmens, B. Butterfield, K. Bagstad, J. Goldstein, R. Wiederholt, B. Mattsson, and W.E. Thogmartin. 2014. National valuation of monarch butterflies suggests incentive-based conservation strategies could be effective. Conservation Letters 7:253-262.


Butler, L.K., L. Ries, I.A. Bisson, T.J. Hayden, M.M. Wikelski, L.M. Romero. 2013. Opposite but analogous effects of road density on songbirds with contrasting habitat preferences. Animal Conservation 16:77-85


Fagan, W. F. and L. Ries. 2012. Edge effects. Berkshire Encyclopedia of Sustainability, Vol. 5: Ecosystem Management and Sustainability. Pp. 118-124.

Zipkin, E. F., L. Ries, R. Reeves, J. Regetz and K. S. Oberhauser. 2012. Tracking climate impacts on the migratory monarch butterfly. Global Change Biology 18:3039-3049

Full article: PDF icon Zipkin et al 2012.pdf (298.77 KB)
Related article: PDF icon Ries et al 2004 AnnReview.pdf (436.36 KB)

Soykan, C.U., L.A. Brand, L. Ries, J.C. Stromberg, C.Hass, D.A.Simmons, Jr., W.J.D. Patterson, J.L. Sabo. 2012. Multitaxonomic diversity patterns along a desert riparian–upland gradient. PLoS ONE 7(1):e28235


Wimp, G.M., S.M. Murphy, D. Lewis, L. Ries. 2011. Do edge responses cascade up or down a multi-trophic food web? Ecology Letters 14:863-870

Koenig, W. D., L. Ries, V.B.K. Olsen*, and A. M. Liebhold. 2011. Avian predators are less abundant during periodical cicada emergences, but why? Ecology 92:784-790.


Ries, L. and T. D. Sisk. 2010. What is an 'edge species'? The implications of sensitivity to habitat edges. Oikos 119:1636-1642.


Hannon*, L., L. Ries and K. S. Williams. 2009. Invertebrates of the San Pedro River. Invited book chapter in (J. Stromberg & B. Tellman, eds.) Ecology and Conservation of the San Pedro River. Island Press.


Ries, L. & S. P. Mullen. 2008. A rare model limits the distribution of its more common mimic: a twist on frequency-dependent Batesian mimicry. Evolution 62:1798-1803.

Calabrese*, J. M., L. Ries, S. F. Matter, D. M. Debinski, J. N. Auckland, J. Roland and W. F. Fagan. 2008. Reproductive asynchrony in natural butterfly populations and its consequences for female matelessness. Journal of Animal Ecology 77:746-756.

Ries, L. and T. D. Sisk. 2008. Butterfly responses to habitat edges are predicted by a simple model in a complex landscape. Oecologia 156:75-86.

Supplementary material: Study site information: Microsoft Office document icon 442_2008_976_MOESM1_ESM.doc (132 KB)
Supplementary material: Habitat preference analysis: Microsoft Office document icon 442_2008_976_MOESM2_ESM.doc (244 KB)


Fletcher Jr., R. J., L. Ries, J. Battin and A. D. Chalfoun. 2007. The role of habitat area and edge in fragmented landscapes: definitively distinct or inevitably intertwined? Canadian Journal of Zoology 85: 1017-1030.


Ries, L., R. J. Fletcher, J. Battin, and T. D. Sisk. 2004. Ecological responses to habitat edges: mechanisms, models and variability explained. Annual Review of Ecology, Evolution and Systematics 35:491-522.

Ries, L. and T. D. Sisk. 2004. A predictive model of edge effects. Ecology 85:2917-2926.


Ries, L. and W. F. Fagan. 2003. Habitat edges as a potential ecological trap for an insect predator. Ecological Entomology 28:567-572.


Ries, L., D. M. Debinski, and M. L. Wieland*. 2001. Conservation value of roadside prairie restoration to butterfly communities. Conservation Biology 15: 401-411.

Ries, L. and D. M. Debinski. 2001. Butterfly responses to habitat edges in the highly fragmented prairies of central Iowa. Journal of Animal Ecology 70: 840-852.