About

Daniel Papaj is a Professor in the Department of Ecology & Evolutionary Biology at the University of Arizona. His research focuses on learning and cognition in bees, tephritid fruit flies, and butterflies, as well as plant-insect interactions. He is involved in exploring the behavioral and ecological aspects of these insects, contributing to the understanding of their interactions with their environment and other species.

Research topics

• Biology
• Psychology
• Neuroscience
• Ecology
• Communication
• Computer Science
• Cognitive psychology
• Genetics
• Audiology
• Zoology
• Evolutionary biology

Selected publications

• Learning

  Elsevier eBooks · 2026-01-01

  book-chapter1st authorCorresponding
  PublisherDOI

• Why some flowers have different forms of anthers &amp; handedness: a commentary on ‘Functions of heteranthery and enantiostyly for wing pollination by pollen-collecting bees in <i>Dilatris ixioides</i> (Haemodoraceae)’

  Annals of Botany · 2025-11-15

  articleOpen accessSenior author
  PublisherOA PDFDOI

• The role of similarity of stimuli and responses in learning by nectar-foraging bumble bees: a test of Osgood’s model

  Animal Behaviour · 2024-12-14

  articleSenior author
  PublisherDOI

• The Role of Similarity of Stimuli and Responses in Learning by Nectar-Foraging Bumble Bees: A Test of Osgood’s Model

  bioRxiv (Cold Spring Harbor Laboratory) · 2024-04-28 · 3 citations

  preprintOpen accessSenior author

  Abstract Learning stimulus – response associations helps animals to adjust to changing environments. Sequentially learned associations may interact with each other, either reinforcing memory, a process referred to as ‘transfer’, or hindering memory, a process referred to as ‘interference.’ According to Osgood’s (1949) model, close similarity between new and previously learned stimuli can enhance the transfer of memory through a process of stimulus generalization. In contrast, the model proposes that if responses are different from those previously learned, generalizing stimuli may lead to confusion, resulting in the interference of memory. Except for some work in humans, the interaction between stimulus similarity and response similarity is poorly documented. Here, we tested Osgood’s model using bumble bees ( Bombus impatiens ) foraging for sucrose on artificial flowers with varied colours (= stimuli) that required either legitimate visitation or nectar robbing (= responses). Bees were first allowed to forage on one type of flower, were then switched to another, and finally were returned to the initial flower type. We measured learning performance via flower handling time and the number of failed visits. Consistent with Osgood’s model, bees made more failed visits when they switched between similarly coloured flowers requiring different foraging techniques but made fewer failed visits when switching between similarly coloured flowers with the same technique. Regardless of similarities in stimuli or responses, however, experienced bees were faster in handling flowers than were naïve bees. Results taken together thus provided mixed support for Osgood’s model. Possible explanations for the mixed results are discussed.

  PublisherOA PDFDOI

• Hung out to dry: diminished flowers offer less to pollinators and us

  New Phytologist · 2024-08-08

  letterOpen accessSenior author

  This article is a Commentary on Barman et al . (2024), 244 : 1013–1023.

  PublisherOA PDFDOI

• Floral cues and flower-handling tactics affect switching decisions by nectar-foraging bumble bees

  Animal Behaviour · 2024-08-03

  articleSenior author
  PublisherDOI

• Seasonal Alternation of Putative Camouflage Wing Morphs of the American Snout Butterfly (<i>Libytheana carinenta</i>)

  The American Naturalist · 2024-05-07 · 2 citations

  article1st authorCorresponding

  Intraspecific variation in camouflage is common in animals. Sexual dimorphism in camouflage is less common and, where observed, attributed to trade-offs between natural selection for predator avoidance and sexual selection for conspicuous mating signals. Here we report on variation in putatively cryptic ventral hindwing patterns in the American snout butterfly, Libytheana carinenta. We use field surveys and crowdsourced data to characterize three morphs. One is found in both sexes, one is male specific, and one is female specific. The sex-specific morphs constitute a sexually dimorphic set whose frequencies change together in time. Field surveys indicate that butterflies in southern Arizona transition from midsummer dominance of the sexually monomorphic pattern to early-fall dominance of the sexually dimorphic set. Crowdsourced data indicate that the sexually dimorphic set dominates in early spring, transitioning later into a mixture of morphs dominated by the monomorphic pattern, with the dimorphic set rising in frequency again in late fall. We discuss this unique pattern of camouflage variation with respect to contemporary theory on animal coloration.

  PublisherDOI

• Floral Cues and Flower Handling Tactics Affect Switching Decisions by Nectar-Foraging Bumble Bees

  bioRxiv (Cold Spring Harbor Laboratory) · 2024-05-02

  preprintOpen access

  Abstract Nectar foraging bees change their use of floral resources as plant species appear in the environment and disappear over their lifetimes. The new flowers used may involve different cues and different nectar extraction tactics. Although bumble bees can adapt to changes in floral cues and required tactics, little is known about whether bees prioritize switching tactics or floral cues when deciding which plant species to switch to. In a laboratory assay, we forced Bombus impatiens (common eastern bumble bee) workers either to switch the handling tactic they were using or to continue using the tactic but to switch the colour of artificial flowers foraged on. We examined whether bees’ tendency to change their tactics was influenced by how similar in colour novel flowers were to familiar ones. We conducted a 2 × 2 factorial experiment using artificial flowers, manipulating the handling tactic that bees were initially trained (legitimate visitation or nectar robbing) and the similarity between novel and trained colours (similar or distinct). We found that under most conditions bees preferred to switch flower colours and retain handling tactics. However, when given experience with legitimate visitation and when novel flowers were markedly different in colour from those they had experienced previously, bees tended to switch tactic while continuing to forage on flowers of the same colour. These findings suggest that the similarity in colour of a new floral resource to the currently exploited resource, along with the flower handling tactic employed by bees, jointly plays an important role in decision-making by foraging bumble bees.

  PublisherOA PDFDOI

• The relationship between preference and switching in flower foraging by bees

  Behavioral Ecology and Sociobiology · 2024-03-01 · 5 citations

  article1st authorCorresponding
  PublisherDOI

• The interplay of experience and pre-existing bias in nectar-robbing behavior by the common eastern bumble bee

  Behavioral Ecology and Sociobiology · 2023-03-01 · 5 citations

  articleSenior author
  PublisherDOI

Recent grants

• Flowers as complex signals: The role of signal uncertainty

  NSF · $420k · 2009–2014

Frequent coauthors

• Henar Alonso-Pimentel

  University of Arizona

  21 shared

• Ronald J. Prokopy

  20 shared

• Anne S. Leonard

  University of Nevada, Reno

  19 shared

• Avery L. Russell

  Missouri State University

  18 shared

• César R. Nufio

  Howard Hughes Medical Institute

  15 shared

• Anna Dornhaus

  University of Arizona

  13 shared

• L.E.M. Vet

  11 shared

• Goggy Davidowitz

  University of Arizona

  9 shared

Labs

• Daniel Papaj LaboratoryPI