Kathleen L Prudic

Interests

Teaching

Biodiversity, Citizen Science, Conservation, Climate Change, Data Science, Ecology, Plant-Animal Interactions, R Programming, Wildlife

Research

Biodiversity, Butterflies, Citizen Science, Climate Change, Conservation, Ecology, Machine Learning, Plant-Animal Interactions, Pollinators, Research Computing, Restoration, Wildlife

Courses

Scholarly Contributions

Books

• Prudic, K. L. (2007). Warning and Deception: Chemical, Behavioral, and Phylogenetic Studies of Aposematic Coloration and Mimicry. The University of Arizona..
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  The study of aposematic coloration and mimicry has a long and distinguished history, and has stimulated scientific inquiry in areas as diverse as chemistry, evolution, ecology, and behavior. Yet, many questions regarding signal function and ecological dynamics remain unknown. This dissertation attempts to address some of these questions about how a visual warning signal functions and how the environment changes its efficacy. Using experimental methods, I evaluated the role of luminance contrast in aposematic signaling using milkweed bugs as model prey and Chinese mantids as model predators. The results of this laboratory experiment illustrated that luminance contrast with background can function as an aposematic signal between prey and predator. Predators learned to avoid unpalatable prey sooner and remembered to avoid unpalatable prey for longer when the prey had higher luminance contrast with the background. These results help define what makes a visual signal conspicuous and designate the importance of high luminance contrast in the efficacy of a warning color signal. Another important characteristic of warning coloration is a reason for the advertisement. This is either a physical or chemical defense that confers some protection against predation. In phytophagous insects this is often a chemical defense that is acquired from the insect’s host plant. By developing new analytical chemistry methods, I was able to identify and quantify the toxic compounds in both the host plant and the viceroy butterfly, a putative aposematic insect. These results provide a chemical mechanism for previous research that demonstrated that the viceroy was unpalatable to avian predators. Next, I was able to test the role of geographic variation in host plant and viceroy chemical defense and how

Journals/Publications

• Crimmins, T. M., Posthumus, E. E., Schaffer, S. N., & Prudic, K. L. (2020). COVID-19 impacts on participation in large scale community science projects in the United States.. Biological Conservation.
• Prudic, K. L. (2021). COVID-19 impacts on participation in large scale biodiversity-themed community science projects in the United States. Biological Conservation.
• Prudic, K. L. (2021). Climate change and local host availability drive the northern range boundary in the rapid expansion of a specialist insect herbivore, Papilio cresphontes. Frontiers in Ecology and Evolution.
• Prudic, K. L. (2021). Fewer butterflies seen by community scientists across the warming and drying landscapes of the American West. Science.
• Prudic, K. L., Schaffer, S. N., Posthumus, E. E., & Crimmins, T. M. (2021). COVID-19 impacts on participation in large scale community science projects in the United States.. Biological Conservation, 256, 109017. doi:doi.org/10.1016/j.biocon.2021.109017
• Prudic, K. L., Timmermann, B. N., Papaj, D. R., Ritland, D. B., & Oliver, J. C. (2019). Mimicry in viceroy butterflies is dependent on abundance of the model queen butterfly. Communications Biology, 2, 68.
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  Mimics should not exist without their models, yet often they do. In the system involving queen and viceroy butterflies, the viceroy is both mimic and co-model depending on the local abundance of the model, the queen. Here, we integrate population surveys, chemical analyses, and predator behavior assays to demonstrate how mimics may persist in locations with low-model abundance. As the queen becomes less locally abundant, the viceroy becomes more chemically defended and unpalatable to predators. However, the observed changes in viceroy chemical defense and palatability are not attributable to differing host plant chemical defense profiles. Our results suggest that mimetic viceroy populations are maintained at localities of low-model abundance through an increase in their toxicity. Sharing the burden of predator education in some places but not others may also lower the fitness cost of warning signals thereby supporting the origin and maintenance of aposematism.
• Prudic, K. L., Wilson, J. K., Toshack, M. C., Gerst, K. L., Rosemartin, A., Crimmins, T. M., & Oliver, J. C. (2019). Creating the Urban Farmer's Almanac with Citizen Science Data. Insects, 10(9).
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  Agriculture has long been a part of the urban landscape, from gardens to small scale farms. In recent decades, interest in producing food in cities has grown dramatically, with an estimated 30% of the global urban population engaged in some form of food production. Identifying and managing the insect biodiversity found on city farms is a complex task often requiring years of study and specialization, especially in urban landscapes which have a complicated tapestry of fragmentation, diversity, pollution, and introduced species. Supporting urban growers with relevant data informs insect management decision-making for both growers and their neighbors, yet this information can be difficult to come by. In this study, we introduced several web-based citizen science programs that can connect growers with useful data products and people to help with the who, what, where, and when of urban insects. Combining the power of citizen science volunteers with the efforts of urban farmers can result in a clearer picture of the diversity and ecosystem services in play, limited insecticide use, and enhanced non-chemical alternatives. Connecting urban farming practices with citizen science programs also demonstrates the ecosystem value of urban agriculture and engages more citizens with the topics of food production, security, and justice in their communities.
• Tyler, F., Larrivée, M., Prudic, K. L., & Ryan Norris, D. (2019). Estimating the annual distribution of monarch butterflies in Canada over 16 years using citizen science data. FACETS, 4(1), 238-253.
• Bhardwaj, S., Prudic, K. L., Bear, A., Dasgupta, M., Wasik, B. R., Tong, X., Cheong, W. F., Wenk, M. R., & Monteiro, A. (2018). Sex Differences in 20-Hydroxyecdysone Hormone Levels Control Sexual Dimorphism in Bicyclus anynana Wing Patterns. MOLECULAR BIOLOGY AND EVOLUTION, 35(2), 465-472.
• Dornelas, M., Antao, L. H., Moyes, F., Bates, A. E., Magurran, A. E., Adam, D., Akhmetzhanova, A. A., Appeltans, W., Arcos, J. M., Arnold, H., Ayyappan, N., Badihi, G., Baird, A. H., Barbosa, M., Barreto, T. E., Baessler, C., Bellgrove, A., Belmaker, J., Benedetti-Cecchi, L., , Bett, B. J., et al. (2018). BioTIME: A database of biodiversity time series for the Anthropocene. GLOBAL ECOLOGY AND BIOGEOGRAPHY, 27(7), 760-786.
• Long, E. C., Brown, B. V., Oliver, J. C., & Prudic, K. L. (2018). Comparisons of Citizen Science Data-Gathering Approaches to Evaluate Urban Butterfly Diversity. Insects, 9(4), 186. doi:10.3390/insects9040186
• Oliver, J. C., Prudic, K. L., & Clement, W. L. (2018). Exploring how climate will impact plant-insect distributions and interactions using open data and informatics. Teaching Issues and Experiments in Ecology, 14.
• Prudic, K. L., Oliver, J. C., Brown, B. V., & Long, E. C. (2018). Comparisons of Citizen Science Data-Gathering Approaches to Evaluate Urban Butterfly Diversity. Insects, 9(4).
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  By 2030, ten percent of earth's landmass will be occupied by cities. Urban environments can be home to many plants and animals, but surveying and estimating biodiversity in these spaces is complicated by a heterogeneous built environment where access and landscaping are highly variable due to human activity. Citizen science approaches may be the best way to assess urban biodiversity, but little is known about their relative effectiveness and efficiency. Here, we compare three techniques for acquiring data on butterfly (Lepidoptera: Rhopalocera) species richness: trained volunteer Pollard walks, Malaise trapping with expert identification, and crowd-sourced iNaturalist observations. A total of 30 butterfly species were observed; 27 (90%) were recorded by Pollard walk observers, 18 (60%) were found in Malaise traps, and 22 (73%) were reported by iNaturalist observers. Pollard walks reported the highest butterfly species richness, followed by iNaturalist and then Malaise traps during the four-month time period. Pollard walks also had significantly higher species diversity than Malaise traps.
• Bear, A., Prudic, K. L., & Monteiro, A. (2017). Steroid hormone signaling during development has a latent effect on adult male sexual behavior in the butterfly Bicyclus anynana. PLOS ONE, 12(3).
• Bear, A., Prudic, K. L., & Monteiro, A. (2017). Steroid hormone signaling during development has a latent effect on adult male sexual behavior in the butterfly Bicyclus anynana. PloS one, 12(3), e0174403.
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  It is well established that steroid hormones regulate sexual behavior in vertebrates via organizational and activational effects. However, whether the organizational/activational paradigm applies more broadly to the sexual behavior of other animals such as insects is not well established. Here we describe the hormonal regulation of a sexual behavior in the seasonally polyphenic butterfly Bicyclus anynana is consistent with the characteristics of an organizational effect. By measuring hormone titer levels, quantifying hormone receptor gene expression in the brain, and performing hormone manipulations, we demonstrate steroid hormone signaling early in pupal development has a latent effect on adult male sexual behavior in B. anynana. These findings suggest the organizational/activational paradigm may be more highly conserved across animal taxa than previously thought.
• Bhardwaj, S., Prudic, K. L., Bear, A., Gupta, M. D., Wasik, B. R., Tong, X., Cheong, W. F., Wenk, M. R., & Monteiro, A. (2017). Sex differences in 20-hydroxyecdysone hormone levels control sexual dimorphism in Bicyclus anynana butterfly wing patterns. Molecular Biology and Evolution.
• Prudic, K. L., McFarland, K. P., Oliver, J. C., Hutchinson, R. A., Long, E. C., Kerr, J. T., & Larrivée, M. (2017). eButterfly: Leveraging Massive Online Citizen Science for Butterfly Consevation. Insects, 8(2).
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  Data collection, storage, analysis, visualization, and dissemination are changing rapidly due to advances in new technologies driven by computer science and universal access to the internet. These technologies and web connections place human observers front and center in citizen science-driven research and are critical in generating new discoveries and innovation in such fields as astronomy, biodiversity, and meteorology. Research projects utilizing a citizen science approach address scientific problems at regional, continental, and even global scales otherwise impossible for a single lab or even a small collection of academic researchers. Here we describe eButterfly an integrative checklist-based butterfly monitoring and database web-platform that leverages the skills and knowledge of recreational butterfly enthusiasts to create a globally accessible unified database of butterfly observations across North America. Citizen scientists, conservationists, policy makers, and scientists are using eButterfly data to better understand the biological patterns of butterfly species diversity and how environmental conditions shape these patterns in space and time. eButterfly in collaboration with thousands of butterfly enthusiasts has created a near real-time butterfly data resource producing tens of thousands of observations per year open to all to share and explore.
• Monteiro, A., Tong, X., Bear, A., Liew, S. F., Bhardwaj, S., Wasik, B. R., Dinwiddie, A., Bastianelli, C., Cheong, W. F., Wenk, M. R., Cao, H., & Prudic, K. L. (2015). Differential Expression of Ecdysone Receptor Leads to Variation in Phenotypic Plasticity across Serial Homologs. PLOS GENETICS, 11(9).
• Monteiro, A., Tong, X., Bear, A., Liew, S. F., Bhardwaj, S., Wasik, B. R., Dinwiddie, A., Bastianelli, C., Cheong, W. F., Wenk, M. R., Cao, H., & Prudic, K. L. (2015). Differential Expression of Ecdysone Receptor Leads to Variation in Phenotypic Plasticity across Serial Homologs. PLoS genetics, 11(9), e1005529.
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  Bodies are often made of repeated units, or serial homologs, that develop using the same core gene regulatory network. Local inputs and modifications to this network allow serial homologs to evolve different morphologies, but currently we do not understand which modifications allow these repeated traits to evolve different levels of phenotypic plasticity. Here we describe variation in phenotypic plasticity across serial homologous eyespots of the butterfly Bicyclus anynana, hypothesized to be under selection for similar or different functions in the wet and dry seasonal forms. Specifically, we document the presence of eyespot size and scale brightness plasticity in hindwing eyespots hypothesized to vary in function across seasons, and reduced size plasticity and absence of brightness plasticity in forewing eyespots hypothesized to have the same function across seasons. By exploring the molecular and physiological causes of this variation in plasticity across fore and hindwing serial homologs we discover that: 1) temperature experienced during the wandering stages of larval development alters titers of an ecdysteroid hormone, 20-hydroxyecdysone (20E), in the hemolymph of wet and dry seasonal forms at that stage; 2) the 20E receptor (EcR) is differentially expressed in the forewing and hindwing eyespot centers of both seasonal forms during this critical developmental stage; and 3) manipulations of EcR signaling disproportionately affected hindwing eyespots relative to forewing eyespots. We propose that differential EcR expression across forewing and hindwing eyespots at a critical stage of development explains the variation in levels of phenotypic plasticity across these serial homologues. This finding provides a novel signaling pathway, 20E, and a novel molecular candidate, EcR, for the regulation of levels of phenotypic plasticity across body parts or serial homologs.
• Prudic, K. L., Stoehr, A. M., Wasik, B. R., & Monteiro, A. (2015). Eyespots deflect predator attack increasing fitness and promoting the evolution of phenotypic plasticity. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES, 282(1798).
• Prudic, K. L., Stoehr, A. M., Wasik, B. R., & Monteiro, A. (2015). Eyespots deflect predator attack increasing fitness and promoting the evolution of phenotypic plasticity. Proceedings. Biological sciences, 282(1798), 20141531.
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  Some eyespots are thought to deflect attack away from the vulnerable body, yet there is limited empirical evidence for this function and its adaptive advantage. Here, we demonstrate the conspicuous ventral hindwing eyespots found on Bicyclus anynana butterflies protect against invertebrate predators, specifically praying mantids. Wet season (WS) butterflies with larger, brighter eyespots were easier for mantids to detect, but more difficult to capture compared to dry season (DS) butterflies with small, dull eyespots. Mantids attacked the wing eyespots of WS butterflies more frequently resulting in greater butterfly survival and reproductive success. With a reciprocal eyespot transplant, we demonstrated the fitness benefits of eyespots were independent of butterfly behaviour. Regardless of whether the butterfly was WS or DS, large marginal eyespots pasted on the hindwings increased butterfly survival and successful oviposition during predation encounters. In previous studies, DS B. anynana experienced delayed detection by vertebrate predators, but both forms suffered low survival once detected. Our results suggest predator abundance, identity and phenology may all be important selective forces for B. anynana. Thus, reciprocal selection between invertebrate and vertebrate predators across seasons may contribute to the evolution of the B. anynana polyphenism.
• Oliver, J. C., Ramos, D., Prudic, K. L., & Monteiro, A. (2013). Temporal gene expression variation associated with eyespot size plasticity in Bicyclus anynana. PloS one, 8(6), e65830.
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  Seasonal polyphenism demonstrates an organism's ability to respond to predictable environmental variation with alternative phenotypes, each presumably better suited to its respective environment. However, the molecular mechanisms linking environmental variation to alternative phenotypes via shifts in development remain relatively unknown. Here we investigate temporal gene expression variation in the seasonally polyphenic butterfly Bicyclus anynana. This species shows drastic changes in eyespot size depending on the temperature experienced during larval development. The wet season form (larvae reared over 24°C) has large ventral wing eyespots while the dry season form (larvae reared under 19°C) has much smaller eyespots. We compared the expression of three proteins, Notch, Engrailed, and Distal-less, in the future eyespot centers of the two forms to determine if eyespot size variation is associated with heterochronic shifts in the onset of their expression. For two of these proteins, Notch and Engrailed, expression in eyespot centers occurred earlier in dry season than in wet season larvae, while Distal-less showed no temporal difference between the two forms. These results suggest that differences between dry and wet season adult wings could be due to a delay in the onset of expression of these eyespot-associated genes. Early in eyespot development, Notch and Engrailed may be functioning as repressors rather than activators of the eyespot gene network. Alternatively, temporal variation in the onset of early expressed genes between forms may have no functional consequences to eyespot size regulation and may indicate the presence of an 'hourglass' model of development in butterfly eyespots.
• Noge, K., Prudic, K. L., & Becerra, J. X. (2012). Defensive roles of (E)-2-alkenals and related compounds in heteroptera. Journal of chemical ecology, 38(8), 1050-6.
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  We examined whether shared volatiles found in various heteropteran species and developmental stages function to repel predators. The nymphal dorsal abdominal gland secretions of Riptortus pedestris (Heteroptera: Alydidae) and Thasus acutangulus (Heteroptera: Coreidae), and the metathoracic scent gland secretion of Euschistus biformis (Heteroptera: Pentatomidae) adults were identified by gas chromatography/mass spectrometry (GC/MS). (E)-2-Hexenal, 4-oxo-(E)-2-hexenal (4-OHE), and (E)-2-octenal were found in all three species and deemed likely candidates for repelling predators. In addition to (E)-2-alkenals, the adult E. biformis secreted (E)-2-hexenyl acetate, (E)-2-octenyl acetate, and four hydrocarbons. We evaluated the potential predator repellent properties of these compounds and compound blends against a generalist, cosmopolitan insect predator, the Chinese praying mantid (Mantodea: Mantidae: Tenodera aridifolia sinensis). Mantids that experienced (E)-2-hexenal, (E)-2-octenal, and (E)-2-octenyl acetate moved away from the site of interaction, while 4-OHE and (E)-2-hexenyl acetate did not affect mantid behavior. The compound blends did not have additive or synergistic repellency effects on predator behavior. Compound repellency was not related to compound volatility. Instead, the repellent effect is likely related to predator olfaction, and the affinity of each compound to receptors on the antennae. Our results also suggest the repellents might intensify the visual defensive signals of aposematism (T. acutangulus nymphs) and mimicry (R. pedestris nymphs) in heteropteran bugs.
• Prudic, K. L., Jeon, C., Cao, H., & Monteiro, A. (2011). Developmental plasticity in sexual roles of butterfly species drives mutual sexual ornamentation. Science (New York, N.Y.), 331(6013), 73-5.
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  Current explanations for why sexual ornaments are found in both sexes include genetic correlation, same sex competition, and mutual mate choice. In this study, we report developmental plasticity in mating behavior as induced by temperature during development in the butterfly Bicyclus anynana. Males and females reciprocally change their sexual roles depending on their larval rearing temperatures. This switch is correlated with a change in mating benefits to females and costs to males. The discrete seasonal environments, wet season and dry season, are known to produce the two developmental forms and as a consequence impose alternating, symmetrical patterns of sexual selection, one season on male ornaments, the following season on female ornaments. Thus, reciprocal selection through time may result in mutual sexual ornamentation.
• Oliver, J. C., & Prudic, K. L. (2010). Are mimics monophyletic? The necessity of phylogenetic hypothesis tests in character evolution. BMC evolutionary biology, 10, 239.
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  The processes governing the origin and maintenance of mimetic phenotypes can only be understood in a phylogenetic framework. Phylogenetic estimates of evolutionary relationships can provide a context for analyses of character evolution; however, when phylogenetic estimates conflict, rigorous analyses of alternative evolutionary histories are necessary to determine the likelihood of a specific history giving rise to the observed pattern of diversity. The polyphenic butterfly Limenitis arthemis provides a case in point. This species is comprised of three lineages, two of which are mimetic and one of which is non-mimetic. Conflicting estimates of the relationships among these three lineages requires direct evaluation of the alternative hypotheses of mimicry evolution.
• Prudic, K. L. (2009). Predation on Animals. Encyclopedia of Life Sciences (eLS). doi:10.1002/9780470015902.A0003284
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  The word predator often invokes a vision of a fierce and cruel animal the world would be better without. However, predation is a major ecological process controlling both the structure and function of communities. Predation affects the distribution and abundance of species, the strength and direction of energy flow within a system and the diversity and composition of communities. Predators play an essential role in evolution. Traits that decrease the likelihood of being predated and traits that increase the efficacy of the predating are under strong selection. This process has resulted in a vast array of prey defences and predator counter-defences. Also, according to recent studies of the fossil record, predation has played a central role in determining the history of life on earth. Thus, predators are not to be viewed as cold-blooded killers; instead, predation is a critical process for maintaining species diversity in both ecological and evolutionary time. Key Concepts Predators are animals which actively catch and consume other animals. Predators may be limited in the type of prey they consume; these limitations may be extrinsic, due to factors such as prey abundance, or they may be intrinsic, where the predators’ physiological, reproductive or ecological requirements result in specialization on certain types of prey. Predation is a crucial ecological force because it moves energy through the system. Mathematical models and empirical research illustrate how predators and their prey coexist and should cycle through time. Prey species evolve defences that reduce their vulnerability to predators whereas predators evolve adaptations to counter the defences of the prey. Predator–prey interactions structured ancient, fossil ecosystems and played a role in determining the history of life on earth. Keywords: population dynamics; community dynamics; prey defence; predator counter defence; predator–prey interactions; coevolution
• Prudic, K. L., & Oliver, J. C. (2008). Once a Batesian mimic, not always a Batesian mimic: mimic reverts back to ancestral phenotype when the model is absent. Proceedings. Biological sciences, 275(1639), 1125-32.
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  Batesian mimics gain protection from predation through the evolution of physical similarities to a model species that possesses anti-predator defences. This protection should not be effective in the absence of the model since the predator does not identify the mimic as potentially dangerous and both the model and the mimic are highly conspicuous. Thus, Batesian mimics should probably encounter strong predation pressure outside the geographical range of the model species. There are several documented examples of Batesian mimics occurring in locations without their models, but the evolutionary responses remain largely unidentified. A mimetic species has four alternative evolutionary responses to the loss of model presence. If predation is weak, it could maintain its mimetic signal. If predation is intense, it is widely presumed the mimic will go extinct. However, the mimic could also evolve a new colour pattern to mimic another model species or it could revert back to its ancestral, less conspicuous phenotype. We used molecular phylogenetic approaches to reconstruct and test the evolution of mimicry in the North American admiral butterflies (Limenitis: Nymphalidae). We confirmed that the more cryptic white-banded form is the ancestral phenotype of North American admiral butterflies. However, one species, Limenitis arthemis, evolved the black pipevine swallowtail mimetic form but later reverted to the white-banded more cryptic ancestral form. This character reversion is strongly correlated with the geographical absence of the model species and its host plant, but not the host plant distribution of L. arthemis. Our results support the prediction that a Batesian mimic does not persist in locations without its model, but it does not go extinct either. The mimic can revert back to its ancestral, less conspicuous form and persist.
• Prudic, K. L., Noge, K., & Becerra, J. X. (2008). Adults and nymphs do not smell the same: the different defensive compounds of the giant mesquite bug (Thasus neocalifornicus: Coreidae). Journal of chemical ecology, 34(6), 734-41.
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  Heteropteran insects often protect themselves from predators with noxious or toxic compounds, especially when these insects occur in aggregations. The predators of heteropteran insects change from small insect predators to large avian predators over time. Thus, a chemical that is deterrent to one type of predator at one point in time may not be deterrent to another type of predator at another point in time. Additionally, these predator deterrent compounds may be used for other functions such as alarm signaling to other conspecifics. Defensive secretion compounds from the adult and the nymph giant mesquite bug (Thasus neocalifornicus: Coreidae) were isolated and identified by gas chromatography-mass spectrometry and NMR. The predominant compounds isolated from the nymph mesquite bugs during a simulated predator encounter were (E)-2-hexenal and 4-oxo-(E)-2-hexenal. In adults, the major compounds released during a simulated predator encounter were hexyl acetate, hexanal, and hexanol. Results from predator bioassays suggest the nymph compounds are more effective at deterring an insect predator than the adult compounds. By using behavioral bioassays, we determined the role of each individual compound in signaling to other mesquite bugs. The presence of the nymph secretion near a usually compact nymph aggregation caused nymph mesquite bugs to disperse but did not affect adults. Conversely, the presence of the adult secretion caused the usually loose adult aggregation to disperse, but it did not affect nymph aggregation. The compounds that elicited nymph behavioral responses were (E)-2-hexenal and 4-oxo-(E)-2-hexenal, while those that elicited adult behavioral responses were hexyl acetate and hexanal. The differences between the chemical composition of nymph and adult defensive secretions and alarm behavior are possibly due to differences in predator guilds.
• Warren, A. D., Prudic, K. L., & Llorente-bousquets, J. (2008). Molecular and morphological evidence reveals three species within the California sister butterfly, Adelpha bredowii (Lepidoptera: Nymphalidae: Limenitidinae). Zootaxa, 1819(1), 1-24. doi:10.11646/zootaxa.1819.1.1
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  In recent decades the butterfly Adelpha bredowii (Lepidoptera: Nymphalidae), known in the USA as the California Sister, has been considered to be a polytypic array of taxa, composed of two, three or four subspecies. The most recent review of the genus (Willmott 2003a) recognized three: A. b. bredowii, A. b. eulalia (= guatemalensis) and A. b. californica. We used mitochondrial (COII) and nuclear (Tpi) DNA sequence data, coupled with a reevaluation of traditional morphological characters, to determine the phylogenetic relationships between members of the A. bredowii complex, and to elucidate their taxonomic status. Phylogenetic analysis of molecular data corroborated the monophyly of each of the three subspecies (sensu Willmott), with the following topology: (bredowii + (eulalia + californica)). Average levels of COII variation among these taxa were much greater than the average variation within each taxon (3.4% vs. 0.2%, respectively). There were no shared alleles among these taxa, even from localities where two lineages (bredowii and eulalia) were collected in exact sympatry and synchrony. The degree of genetic divergence, reciprocal monophyly, and absence of shared alleles between taxa, coupled with unique morphological and distributional attributes of each, strongly suggest that A. bredowii, A. eulalia and A. californica are all species-level taxa, as they are treated herein (rev. stat.). Adelpha bredowii is endemic to western, central and southern Mexico: the syntype female (herein designated Lectotype) most likely originated near Oaxaca City, Oaxaca. Adelpha eulalia occurs from Guatemala, through much of Mexico, to Arizona, New Mexico and Texas, United States (as permanent residents; further north as vagrants), and occurs in exact sympatry with A. bredowii at least in Oaxaca, Mexico, but potentially in much of western and southwestern Mexico. The syntype female of eulalia (herein designated Lectotype) most likely originated from south-central Mexico. Adelpha californica is allopatric with respect to the other two species, occurring in northern Baja California, much of California, far western Nevada, Oregon and Washington. Our results highlight the continuous need for systematic scrutiny of familiar taxa, and raise the prospect that the genus Adelpha may be comprised of many more hidden species.
• Frentiu, F. D., Bernard, G. D., Cuevas, C. I., Sison-Mangus, M. P., Prudic, K. L., & Briscoe, A. D. (2007). Adaptive evolution of color vision as seen through the eyes of butterflies. Proceedings of the National Academy of Sciences of the United States of America, 104 Suppl 1, 8634-40.
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  Butterflies and primates are interesting for comparative color vision studies, because both have evolved middle- (M) and long-wavelength- (L) sensitive photopigments with overlapping absorbance spectrum maxima (lambda(max) values). Although positive selection is important for the maintenance of spectral variation within the primate pigments, it remains an open question whether it contributes similarly to the diversification of butterfly pigments. To examine this issue, we performed epimicrospectrophotometry on the eyes of five Limenitis butterfly species and found a 31-nm range of variation in the lambda(max) values of the L-sensitive photopigments (514-545 nm). We cloned partial Limenitis L opsin gene sequences and found a significant excess of replacement substitutions relative to polymorphisms among species. Mapping of these L photopigment lambda(max) values onto a phylogeny revealed two instances within Lepidoptera of convergently evolved L photopigment lineages whose lambda(max) values were blue-shifted. A codon-based maximum-likelihood analysis indicated that, associated with the two blue spectral shifts, four amino acid sites (Ile17Met, Ala64Ser, Asn70Ser, and Ser137Ala) have evolved substitutions in parallel and exhibit significant d(N)/d(S) >1. Homology modeling of the full-length Limenitis arthemis astyanax L opsin placed all four substitutions within the chromophore-binding pocket. Strikingly, the Ser137Ala substitution is in the same position as a site that in primates is responsible for a 5- to 7-nm blue spectral shift. Our data show that some of the same amino acid sites are under positive selection in the photopigments of both butterflies and primates, spanning an evolutionary distance >500 million years.
• Papaj, D. R., Skemp, A. K., Prudic, K. L., & Papaj, D. R. (2007). Aposematic coloration, luminance contrast, and the benefits of conspicuousness. Behavioral Ecology, 18(1), 41-46. doi:10.1093/beheco/arl046
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  Many organisms use warning, or aposematic, coloration to signal their unprofitability to potential predators. Aposematically colored prey are highly visually conspicuous. There is considerable empirical support that conspicuousness promotes the effectiveness of the aposematic signal. From these experiments, it is well documented that conspicuous, unprofitable prey are detected sooner and aversion learned faster by the predator as compared with cryptic, unprofitable prey. Predators also retain memory of the aversion longer when prey is conspicuous. The present study focused on the elements of conspicuousness that confer these benefits of aposematic coloration. Drawing on current understanding of animal vision, we distinguish 2 features of warning coloration: high chromatic contrast and high brightness, or luminance, contrast. Previous investigations on aposematic signal efficacy have focused mainly on the role of high chromatic contrast between prey and background, whereas little research has investigated the role of high luminance contrast. Using the Chinese mantid as a model predator and gray-painted milkweed bugs as model prey, we found that increased prey luminance contrast increased detection of prey, facilitated predator aversion learning, and increased predator memory retention of the aversive response. Our results suggest that the luminance contrast component of aposematic coloration can be an effective warning signal between the prey and predator. Thus, warning coloration can even evolve as an effective signal to color blind predators. Copyright 2007.
• Prudic, K. L., Khera, S., Sólyom, A., & Timmermann, B. N. (2007). Isolation, identification, and quantification of potential defensive compounds in the viceroy butterfly and its larval host-plant, Carolina willow. Journal of chemical ecology, 33(6), 1149-59.
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  The viceroy-monarch and viceroy-queen butterfly associations are classic examples of mimicry. These relationships were originally classified as Batesian, or parasitic, but were later reclassified as Müllerian, or mutalistic, based on predator bioassays. The Müllerian reclassification implies that viceroy is unpalatable because it too is chemically defended like the queen and the monarch. However, unlike the queen and the monarch, the viceroy defensive chemistry has remained uncharacterized. We demonstrate that the viceroy butterfly (Limenitis archippus, Nymphalidae) not only sequesters nonvolatile defensive compounds from its larval host-plant, the Carolina willow (Salix caroliniana, Salicaceae), but also secretes volatile defensive compounds when disturbed. We developed liquid chromatography-mass spectrometry-mass spectrometry methods to identify a set of phenolic glycosides shared between the adult viceroy butterfly and the Carolina willow, and solid phase microextraction and gas chromatography-mass spectrometry methods to identify volatile phenolic compounds released from stressed viceroy butterflies. In both approaches, all structures were characterized based on their mass spectral fragmentation patterns and confirmed with authentic standards. The phenolics we found are known to deter predator attack in other prey systems, including other willow-feeding insect species. Because these compounds have a generalized defensive function at the concentrations we described, our results are consistent with the Müllerian reclassification put forth by other researchers based on bioassay results. It seems that the viceroy butterfly possesses chemical defenses different from its monarch and queen butterfly counterparts (phenolic glycosides vs. cardiac glycosides, respectively), an unusual phenomenon in mimicry warranting future study.
• Prudic, K. L., Oliver, J. C., & Sperling, F. A. (2007). The signal environment is more important than diet or chemical specialization in the evolution of warning coloration. Proceedings of the National Academy of Sciences of the United States of America, 104(49), 19381-6.
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  Aposematic coloration, or warning coloration, is a visual signal that acts to minimize contact between predator and unprofitable prey. The conditions favoring the evolution of aposematic coloration remain largely unidentified. Recent work suggests that diet specialization and resultant toxicity may play a role in facilitating the evolution and persistence of warning coloration. Using a phylogenetic approach, we investigated the evolution of larval warning coloration in the genus Papilio (Lepidoptera: Papilionidae). Our results indicate that there are at least four independent origins of aposematic larval coloration within Papilio. Controlling for phylogenetic relatedness among Papilio taxa, we found no evidence supporting the hypothesis that either diet specialization or chemical specialization facilitated the origin of aposematic larvae. However, there was a significant relationship between the signal environment and the evolution of aposematic larvae. Specifically, Papilio lineages feeding on herbaceous or narrow-leaved plants, regardless of the plants' taxonomic affiliation, were more likely to evolve aposematic larvae than were lineages feeding only on trees/shrubs or broad-leaved plants. These results demonstrate that factors other than diet specialization, such as the signal environment of predator-prey interactions, may play a large role in the initial evolution and persistence of aposematic coloration.
• Prudic, K. L., Oliver, J. C., & Collinge, S. K. (2006). BOULDER COUNTY OPEN SPACE BUTTERFLY DIVERSITY AND ABUNDANCE. Ecology, 87(4), 1066-1066. doi:10.1890/0012-9658(2006)87[1066:bcosbd]2.0.co;2
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  Abstract. The purpose of this data set was to investigate the effects of habitat characteristics and urbanization on grassland butterfly diversity. We sampled butterflies at 66 sites in Boulder County Open Space in the years 1999 and 2000. We tested for effects of habitat characteristics (grassland type and quality) and landscape context (percentage surrounding urbanization) on butterfly diversity and abundance. Although both habitat characteristics did affect butterfly diversity, landscape context did not. The data contain butterfly species diversity and individual species' abundance for the five periods sampled: July 1999, August 1999, June 2000, July 2000, and August 2000. Our samples included 58 species from five butterfly families: Hesperiidae (17 species), Papilionidae (4 species), Pieridae (7 species), Lycaenidae (11 species), and Nymphalidae (19 species). These data include endangered butterfly species, as well as surveys from relictual tallgrass prairie. There is some limitation to these data, as...
• Prudic, K. L., Oliver, J. C., & Bowers, M. D. (2005). Soil nutrient effects on oviposition preference, larval performance, and chemical defense of a specialist insect herbivore. Oecologia, 143(4), 578-87.
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  This study examined the effects of increased leaf nitrogen in natural host-plants (Plantago spp.) on female oviposition preference, larval performance, and larval chemical defense of the butterfly Junonia coenia. Increased availability of soil nutrients caused the host-plant's foliar nitrogen to increase and its chemical defense to decrease. Larval performance did not correlate with increases in foliar nitrogen. Larval growth rate and survival were equivalent across host-plant treatments. However, larvae raised on fertilized host-plants showed concomitant decreases in chemical defense as compared to larvae reared on unfertilized host-plants. Since most butterfly larvae cannot move long distances during their first few instars and are forced to feed upon the plant on which they hatched, J. coenia larval chemical defense is determined, in large part, by female oviposition choice. Female butterflies preferred host-plants with high nitrogen over host-plants with low nitrogen; however, this preference was also mediated by plant chemical defense. Female butterflies preferred more chemically defended host-plants when foliar nitrogen was equivalent between host-plants. J. coenia larvae experience intense predation in the field, especially when larvae are not chemically well defended. Any qualitative or quantitative variation in plant allelochemical defense has fitness consequences on these larvae. Thus, these results indicate that females may be making sub-optimal oviposition decisions under a nutrient-enriched regime, when predators are present. Given the recent increase in fertilizer application and nitrogen deposition on the terrestrial landscape, these interactions between female preference, larval performance, and larval chemical defense may result in long-term changes in population dynamics and persistence of specialist insects.
• Prudic, K. L., Oliver, J. C., & Collinge, S. K. (2003). Effects of Local Habitat Characteristics and Landscape Context on Grassland Butterfly Diversity. Conservation Biology, 17(1), 178-187. doi:10.1046/j.1523-1739.2003.01315.x
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  Recent ecological studies suggest that the landscape context of native habitat remnants may signif- icantly influence plant and animal abundance and distribution within those remnants. Other research has revealed a weak link between landscape context and native community composition. To understand the rela- tive importance of local and regional habitat characteristics for grassland butterflies, we assessed butterfly community diversity in four types of grassland habitats surrounded by varying amounts of urban develop- ment near Boulder, Colorado (U.S.A.). We recorded butterfly species abundance and composition in 66 grass- land study plots on five sampling dates in 1999 and 2000. Grasslands were of four types: native shortgrass, native mixed grass, native tallgrass, and planted hayfields. Grasslands also varied in quality, determined by the abundance of native versus exotic plant species. We observed highly significant effects of grassland type on butterfly species richness and composition. For example, tallgrass plots supported significantly higher but- terfly species richness than shortgrass plots ( p � 0.01). Habitat quality also affected butterfly species richness and composition. Low-quality plots generally supported fewer species than moderate- or high-quality plots ( p � 0.05). Landscape context—the percentage of urbanization in the surrounding landscape—did not sig- nificantly predict butterfly species richness or composition. Our observations suggest that for the grassland butterfly communities in our study, (1) grassland type was the primary determinant of species richness and composition, (2) habitat quality secondarily affected butterfly community diversity, and (3) landscape con- text did not significantly predict butterfly species composition. Our findings emphasize the importance of maintaining high-quality grassland habitat to protect native butterfly diversity.
• Voss, S. R., Shaffer, H. B., Prudic, K. L., & Oliver, J. C. (2003). Candidate gene analysis of metamorphic timing in ambystomatid salamanders.. Molecular ecology, 12(5), 1217-23. doi:10.1046/j.1365-294x.2003.01806.x
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  Although much is known about the ecological significance of metamorphosis and metamorphic timing, few studies have examined the underlying genetic architecture of these traits, and no study has attempted to associate phenotypic variation to molecular variation in specific genes. Here we report on a candidate gene approach (CGA) to test specific loci for a statistical contribution to variation in metamorphic timing. Three segregating populations (SP1, SP2 and SP3) were constructed utilizing three species of paedomorphic Mexican ambystomatid salamander, including the axolotl, Ambystoma mexicanum. We used these replicated species to test the hypothesis that inheritance of alternate genotypes at two thyroid hormone receptor loci (TRalpha, TRbeta) affects metamorphic timing in ambystomatid salamanders. A significant TRalpha*SP effect indicated that variation in metamorphic timing may be influenced by TRalpha genotype, however, the effect was not a simple one, as both the magnitude and direction of the phenotypic effect depended upon the genetic background. These are the first data to implicate a specific gene in contributing to variation in metamorphic timing. In general, candidate gene approaches can be extended to any number of loci and to any organism where simple genetic crosses can be performed to create segregating populations. The approach is thus of particular value in ecological studies where target genes have been identified but the study organism is not one of the few well-characterized model systems that dominate genetic research.
• Shapiro, A. M., Prudic, K. L., & Clayton, N. S. (2002). Evaluating a putative mimetic relationship between two butterflies, Adelpha bredowii and Limenitis lorquini. Ecological Entomology, 27(1), 68-75. doi:10.1046/j.0307-6946.2001.00384.x
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  Abstract 1. A laboratory bioassay experiment was performed in order to investigate a long-standing putative mimetic relationship between two butterflies, the Lorquin's admiral Limenitis lorquini and the California sister Adelpha bredowii (Lepidoptera, Nymphalidae). These butterfly species are commonly sympatric in their broad distribution on the west coast of the United States. The wing colour pattern of L. lorquini differs from the patterns exhibited by its North American congeners, resembling A. bredowii instead. 2. The feeding responses of California scrub jays Aphelocoma californica (Passeriformes, Corvidae) to these prey types and a known palatable control, the buckeye butterfly Junonia coenia (Lepidoptera, Nymphalidae), were examined quantitatively. The birds usually demonstrated long handling times of A. bredowii before consumption, while L. lorquini was manipulated and consumed at the same rate as the control butterfly. The birds also exhibited feather ruffling and bill wiping after consuming A. bredowii while they did not exhibit such behaviours after consuming L. lorquini and J. coenia. 3. The birds did not discriminate between the dorsal colour patterns of the two species in a choice experiment. 4. Because A. bredowii was moderately unpalatable to the captive scrub jays and the birds were not discriminating in their choice, this suggests a Batesian mimetic relationship between L. lorquini and A. bredowii.

Proceedings Publications

• Prudic, K. L., Larrive, M., & Mcfarland, K. P. (2016). Symposium: Keeping Science in Citizen Science. In International Congress of Entomology.

Presentations

• Prudic, K. L. (2022). Big data drives big challenges in biodiversity monitoring. Lewis and Clark Departmental Seminar.
• Prudic, K. L. (2022, Dec 2022). Community science, climate change, and monitoring. Research Interests in Semiarid Ecosystems. Tucson, AZ.
• Prudic, K. L., & Crimmins, T. M. (2022). How you and your phone can save animals from extinction. TenWest. Tucson, AZ.
• Prudic, K. L., & Crimmins, T. M. (2022). How you and your phone can save animals from extinction. Wonder House, SXSW.
• Prudic, K. L. (2020, Summer). Harnessing big citizen science data: Climate change and local host availability drive the northern range boundary in the rapid northward expansion of the eastern giant swallowtail butterfly. Ecology Society of America.
• Prudic, K. L. (2020, Summer). Improving citizen science data collection for biodiversity conservation through targeted sampling. Ecological Society of America.
• Prudic, K. L. (2020, Fall). Data Science and Conservation Takes Flight in Butterflies. Florida Museum of Natural History Seminar.
• Giebink, N., Wilson, J. K., & Prudic, K. L. (2019, November). Unlocking big data for pollinator conservation. University of Arizona - EEB Department.
• Prudic, K. L. (2019, April). Data Science and Conservation. Women in Data Science - University of Arizona.
• Prudic, K. L. (2019, November). Citizen Science and Conservation Take Flight in Butterflies. Entomological Society of America.
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  Invited speaker to ESA 2019 program symposium: Marketing Entomology in the 21st Century: Delivering Knowledge, Changing Attitudes, and Encouraging Action organized by David Onstad and Keri Carstens
• Prudic, K. L. (2019, October). Data Science Conservation Takes Flight in Butterflies. Iowa State University - Entomology Department.
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  Invited departmental seminar
• Prudic, K. L. (2019, September). Citizen Science and Environmental Education, A Match Made in Heaven. Arizona Association for Environmental Education.
• Prudic, K. L. (2019, September). Citizen Science for Health Care: Lessons from Butterflies. University of Arizona Grand Rounds - Department of OB/GYN.
• Prudic, K. L. (2019, September). How butterflies make a living and how we can help them in a changing world. Boise State University - Biology Department.
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  Invited seminar sponsored by graduate students
• Bronstein, J., Davidowitz, G., Prudic, K. L., & Busby, K. (2018, June). Forecasting the developmental niche of Xylocopa californica in a changing thermal world. Entomological Society of America, Pacific Branch. Reno, NV: ESA.
• Prudic, K. L. (2017, August). Data Science and Biodiversity - Where do we go from here?. The Lepidopterists' Society. Tucson, Arizona: The Lepidopterists' Society.
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  Keynote address
• Prudic, K. L. (2017, January). Ecophysiology: tales from the field. Departmental SeminarUniversity of Illinois, Urbana-Champagne.
• Prudic, K. L., Kent, M. P., Rebecca, H., Oliver, J. C., Jeremy, K., Maxim, L., & Elizabeth, L. C. (2017, November). Real time massive online citizen science biodiversity programs: Lessons from butterflies. Entomological Society of America. Denver, CO.
• Prudic, K. L. (2016, November). Citizen Science, massive data, and new directions. Entomological Society of America. Orlando, FL.
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  Organized a symposium with 8 speakers all involved in insect citizen science programs. Also gave a talk in symposium

Poster Presentations

• Prudic, K. L., Melkonoff, N., & Zylstra, E. (2022). Butterflies, hostplants and their distributional responses to climate change. Ecology Society of America.
• Prudic, K. L., Melkonoff, N., & Zylstra, E. (2022). Butterflies, hostplants and their distributional responses to climate change. Society for Integrative and Comparative Biology.
• Giebink, N., Wilson, J. K., & Prudic, K. L. (2019, November). Does targeted sampling in remote areas solve the sampling bias challenge in citizen science data?. Society for Ecological Restoration - Southwest.
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  Authors: Noah Giebink (50%), Keaton Wilson (25%) , Katy Prudic (25%)

Other Teaching Materials

• Clement, W. L., Prudic, K. L., & Oliver, J. C. (2019. Exploring how climate will impact plant-insect distributions and interactions using open data and informatics. QUBES Educational Resources.
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  This teaching module provides an entry point for students to learn about data science, open data repositories (e.g., citizen science data), and species distribution modeling to study the effects of climate change on butterfly-host plant interactions.