About Me

I am a Principal Scientist on the Nutrition Innovation team at Hill's Pet Nutrition with expertise in canine cognition and behavior. In this role, I lead scientific research to evaluate the efficacy of novel nutritional solutions aimed at improving the health and well-being of pets. My work blends citizen science approaches with on-site research at the Hill's Global Pet Nutrition Center, and includes developing new cognitive, behavioral, and biological endpoints to quantify pet health. I also cultivate and lead collaborative external partnerships that have the potential to advance these research aims.

Prior to my current role, I was a Postdoctoral Fellow in the Department of Psychology at Yale University, working with Dr. Laurie Santos and Dr. Julián Jara-Ettinger. I graduated from the University of North Carolina at Chapel Hill in 2016 with a B.S. in Psychology, and I received an M.A. (2018) and Ph.D. (2021) in Biological Anthropology from the University of Arizona under the direction of Dr. Evan MacLean. I have also previously worked as a Graduate Teaching Associate in both the School of Anthropology and the School of Mind, Brain and Behavior at the University of Arizona.

My research interests during my time in academia broadly centered on cognitive evolution, with a particular emphasis on social-cognitive abilities such as theory of mind, shared intentionality, and cooperative communication. For example, how do humans represent the mental states of others, and how do we use these representations to guide our goal-directed actions? Do other animals represent others' mental states in the same ways? Is our ability to form joint commitments unique, or do other animals also understand and share others' intentions? I investigated these and related questions by integrating non-invasive cognitive, behavioral, and biological research in humans, dogs, wolves, and free-ranging non-human primates.

My most recent academic research was funded by a Social, Behavioral and Economic Sciences Postdoctoral Research Fellowship from the National Science Foundation. This award supported my work jointly with the Comparative Cognition Lab and Computational Social Cognition Lab to build and evaluate agent-based computational models of non-human primate theory of mind.

Research Experience

Hill's Global Pet Nutrition Center, Topeka, KS (2023 – Present)
Computational Social Cognition Lab, New Haven, CT (2021 – 2023)
Comparative Cognition Lab, New Haven, CT (2021 – 2023)
Canine Cognition Center at Yale, New Haven, CT (2015; 2021 – 2023)
Cayo Santiago Biological Field Station, Punta Santiago, Puerto Rico (2016 – 2023)
Canine Companions for Independence, Santa Rosa, CA & Oceanside, CA (2016 – 2021)
Derby City Agility Association, Louisville, KY (2021)
Humane Society of Southern Arizona, Tucson, AZ (2018 – 2021)
Arizona Canine Cognition Center, Tucson, AZ (2016 – 2021)
Wildlife Science Center, Stacy, MN (2018)
Human Memory Lab, Chapel Hill, NC (2015 – 2016)
Lenovo User Experience Research Team, Morrisville, NC (2014 – 2016)
Somatosensory Research Lab, Chapel Hill, NC (2013 – 2014)

Peer-Reviewed Publications

Electronic versions of the articles listed below are provided to ensure timely dissemination of scholarly work. These files may not be reposted or shared without permission by the respective copyright holders and may be downloaded strictly for individual, non-commercial use.

18) Horschler, D.J.*, Berke, M.*, Santos, L.R., & Jara-Ettinger, J. (2023). Differences between human and non-human primate theory of mind: Evidence from computational modeling. [*Denotes equal contribution] [Preprint]

17) ManyDogs Project, Espinosa, J., Stevens, J.R., Alberghina, D., Alway, H.E.E., Barela, J.D., Bogese, M., Bray, E.E., Buchsbaum, D., Byosiere, S.E., Byrne, M., Cavalli, C.M., Chaudoir, L.M., Collins-Pisano, C., DeBoer, H.J., Douglas, L.E.L.C., Dror, S., Dzik, M.V., Ferguson, B., Fisher, L., Fitzpatrick, L.C., Freeman, M.S., Frinton, S.N., Glover, M.K., Gnanadesikan, G.E., Goacher, J.E.P., Golanska, M., Alexandrina Guran, C.-N., Hare, E., Hare, B., Hickey, M., Horschler, D.J., Huber, L., Jim, H.-L., Johnston, A.M., Kaminksi, J., Kelly, D.M., Kuhlmeier, V.A., Lassiter, L., Lazarowski, L., Leighton-Birch, L., MacLean, E.L., Maliszewska, K., Marra, V., Montgomery, L.I., Murray, M.S., Nelson, E.K., Ostojic, L., Palermo, S.G., Parks Russell, A.E., Pelgrim, M.H., Pellowe, S.D., Reinholz, A., Rial, L.A., Richards, E.M., Ross, M.A., Rothkoff, L.G., Salomons, H., Sanger, J.K., Santos, L., Schirle, A.R., Shearer, S.J., Silver, Z.A., Silverman, J.M., Sommese, A., Srdoc, T., St. John-Mosse, H., Vega, A.C., Vekony, K., Völter, C.J., Walsh, C.J., Worth, Y.A., Zipperling, L.M.I., Zoledziewska, B., & Zylberfuden, S.G. (2023). ManyDogs 1: A multi-lab replication study of dogs’ pointing comprehension. Animal Behavior & Cognition, 10(3), 232-286. [PDF] [Data]

16) Byrne, M., Horschler, D.J., Schmitt, M., & Johnston, A.M. (2023). Pet dogs (Canis familiaris) re-engage humans after joint activity. Animal Cognition, 26, 1277-1282. [PDF]

15) ManyDogs Project, Alberghina, D., Bray, E.E., Buchsbaum, D., Byosiere, S.E., Espinosa, J., Gnanadesikan, G.E., Alexandrina Guran, C.-N., Hare, E., Horschler, D.J., Huber, L., Kuhlmeir, V.A., MacLean, E.L., Pelgrim, M.H., Perez, B., Ravid-Schurr, D., Rothkoff, L., Sexton, C., Silver, Z.A., & Stevens, J.R. (2023). The Many Dogs Project: A big team science approach to investigating canine behavior and cognition. Comparative Cognition & Behavior Reviews, 18, 059-077. [PDF]

14) Salomons, H., Smith, K.C.M., Callahan-Beckel, M., Callahan, M., Levy, K., Kennedy, B.S., Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., Gruen, M., Tan, J., White, P., vonHoldt, B.M., MacLean, E.L., & Hare, B. (2023). Response to Hansen Wheat et al.: Additional analysis further supports the early emergence of cooperative communication in dogs compared to wolves raised with more human exposure. Learning & Behavior, 51(2), 131-134. [PDF]

13) Mulligan, N.W., Susser, J.A., & Horschler, D.J. (2022). Action memory and metamemory. Journal of Experimental Psychology: Learning, Memory, and Cognition, 48(12), 1821–1832. [PDF]

12) Horschler, D.J., Bray, E.E., Gnanadesikan, G.E., Byrne, M., Levy, K.M., Kennedy, B.S., & MacLean, E.L. (2022). Dogs re-engage human partners when joint social play is interrupted: A behavioural signature of shared intentionality? Animal Behaviour, 183, 159-168. [PDF]

11) Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., Levy, K.M., Kennedy, B.S., Famula, T.R., & MacLean, E.L. (2021). Early-emerging and highly heritable sensitivity to human communication in dogs. Current Biology, 31(14), 3137-3144. [PDF] [Methods]
    ☨ Featured on the issue’s cover

10) Salomons, H., Smith, K.C.M., Callahan-Beckel, M., Callahan, M., Levy, K., Kennedy, B.S., Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., Gruen, M., Tan, J., White, P., vonHoldt, B.M., MacLean, E.L., & Hare, B. (2021). Cooperative communication with humans evolved to emerge early in domestic dogs. Current Biology, 31(14), 3132-3136. [PDF]

9) Horschler, D.J., Santos, L.R., & MacLean, E.L. (2021). How do non-human primates represent others’ awareness of where objects are hidden? Cognition, 212, 104658. [PDF]

8) Arre, A.M.* & Horschler, D.J.* (2021). Swimming and diving as social play in juvenile rhesus macaques (Macaca mulatta). Behaviour, 158(6), 529-546. [*Denotes equal contribution] [PDF]

7) Bray, E.E., Gruen, M.E., Gnanadesikan, G.E., Horschler, D.J., Levy, K.M., Kennedy, B.S., Hare, B.A., & MacLean, E.L. (2020). Dog cognitive development: A longitudinal study across the first two years of life. Animal Cognition, 24(2), 311-328. [PDF] [Methods]

6) Horschler, D.J., MacLean, E.L., & Santos, L.R. (2020). Advancing gaze-based research on primate theory of mind. Trends in Cognitive Sciences, 24(10), 778-779. [PDF]

5) Bray, E.E., Gruen, M.E., Gnanadesikan, G.E., Horschler, D.J., Levy, K.M., Kennedy, B.S., Hare, B.A., & MacLean, E.L. (2020). Cognitive characteristics of 8-to-10-week-old assistance dog puppies. Animal Behaviour, 66, 193-206. [PDF] [Methods]

4) Horschler, D.J., MacLean, E.L., & Santos, L.R. (2020). Do non-human primates really represent others’ beliefs? Trends in Cognitive Sciences, 24(8), 594-605. [PDF]

3) Horschler, D.J. & MacLean, E.L. (2019). Leveraging brain-body scaling relationships for comparative studies. Animal Cognition, 22(6), 1197-1202. [PDF]

2) Horschler, D.J., Santos, L.R., & MacLean, E.L. (2019). Do non-human primates really represent others' ignorance? A test of the awareness relations hypothesis. Cognition, 190, 72-80. [PDF]

1) Horschler, D.J., Hare, B., Call, J., Kaminski, J., Miklósi, Á., & MacLean, E.L. (2019). Absolute brain size predicts dog breed differences in executive function. Animal Cognition, 22(2), 187-198. [PDF]
    ☨ Selected as the issue’s highlight article

Manuscripts

Silver, Z.A., Horschler, D.J., Richards, E.M., Siegel, L.A., Furlong, E.E., & Santos, L.R. (Under review). The evolutionary origins of social evaluation: Do dogs (Canis lupus familiaris) evaluate others based on competence?

Gnanadesikan, G.E., Bray, E.E., Levy, K.M., Horschler, D.J., Hargrave, S., Douglas, L.E.L.C., Kennedy, B.S., Watowich, M.M., Synder-Mackler, N., & MacLean, E.L. (Under review). Characterizing the heritability of cognitive and behavioral traits across development in domestic dogs.

Horschler, D.J., Silver, Z.A., Leimgruber, K.L., Sánchez, C.A., Widness, J. & Santos, L.R. (In prep). Capuchin monkeys (Sapajus apella) engage in generalized reciprocity.

Bray, E.E., Douglas, L.E.L.C., Levy, K.M., Gnanadesikan, G.E., Horschler, D.J., Kennedy, B.S., & MacLean, E.L. (In prep). Effects of maternal style and early rearing environment on puppy behavior and cognition.

Silver, Z.A., Thomas, A., Derks, N., Horschler, D.J., Richards, E., Furlong, E., & Santos, L.R. (In prep). Do dogs prefer the winners of zero-sum conflicts?

Conference Presentations

29) Silver, Z.A., Horschler, D.J., Richards, E.M., Siegel, L.A., Furlong, E.E., & Santos, L.R. (2022). Evaluations of prosociality and competence in domestic dogs (Canis lupus familiaris). Talk presented at the Canine Science Conference, Hamilton, NY.

28) Bray, E.E., Douglas, L.E.L.C., Levy, K.M., Gnanadesikan, G.E., Horschler, D.J., Kennedy, B.S., & MacLean, E.L. (2022). Effects of early rearing environment on working dog puppy behavior and outcome. Talk presented at the Canine Science Conference, Hamilton, NY.

27) Byrne, M., Horschler, D.J., Schmitt, M., & Johnston, A.M. (2022). Re-engagement after disrupted play; evidence for shared intentionality in pet dogs? Poster presented at the Canine Science Conference, Hamilton, NY.

26) Horschler, D.J., Berke, M., Santos, L.R., & Jara-Ettinger, J. (2022). Multiple representational theories explain non-human primate perspective-taking: Evidence from computational modeling. Poster presented at the Bridging the Technological Gap Workshop, Göttingen, DE.

25) Horschler, D.J., Berke, M., Santos, L.R., & Jara-Ettinger, J. (2022). Multiple representational theories explain non-human primate perspective-taking: Evidence from computational modeling. Poster presented at the 44th Annual Conference of Cognitive Science Society, Toronto, CA.

24) Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., Levy, K.M., Kennedy, B.S., Famula, T.R., & MacLean, E.M. (2021). Development and heritability of cognitive traits in Canine Companions dogs. Talk presented at the International Working Dog Conference (Virtual meeting).

23) Horschler, D.J., MacLean, E.L., & Santos, L.R. (2021). What do non-human primates represent about others’ knowledge of hidden objects? Talk presented at the Annual Conference of the European Society for Philosophy and Psychology (Virtual meeting).

22) Horschler, D.J., Bray, E.E., & MacLean, E.L. (2021). Interpreting dogs' re-engagement behaviors after disrupted joint activities. Talk presented at the Yale University East Coast Canine Cognition Workshop (Virtual meeting).

21) Horschler, D.J., Bray, E.E., & MacLean, E.L. (2021). Do dogs form joint intentions with humans in social play? Talk presented at the Society for Personality and Social Psychology’s Origins of the Social Mind Preconference (Virtual meeting).

20) Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., & MacLean, E.L. (2021). Heritable variation in dog social cognition. Thread presented at the Animal Behavior Society and Association for the Study of Animal Behaviour Twitter Conference (Virtual meeting).

19) Gnanadesikan, G.E., Espinsoa, J.H., & ManyDogs. (2021). ManyDogs 1: An International Collaborative Approach to Pointing Comprehension in Domestic Dogs. Thread presented at the Animal Behavior Society and Association for the Study of Animal Behaviour Twitter Conference (Virtual meeting).

18) Horschler, D.J., Bray, E.E., & MacLean, E.L. (2020). Do dogs re-engage humans when joint experiences end? Talk presented at the Yale University East Coast Canine Cognition Workshop, New Haven, CT.

17) Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., & MacLean, E.L. (2020). Early development and longitudinal stability of cognitive traits in working dogs. Talk presented at the Yale University East Coast Canine Cognition Workshop, New Haven, CT.

16) MacLean, E.L., Bray, E.E., Gnanadesikan, G.E., & Horschler, D.J. (2020). Associations between individual differences in cognition and training outcomes in assistance dogs. Talk presented at the Yale University East Coast Canine Cognition Workshop, New Haven, CT.

15) Horschler, D.J., Santos, L.R., & MacLean, E.L. (2019). Awareness relations underpin knowledge representation in rhesus monkeys. Talk presented at the UA-ASU Cognitive Science Conclave, Tucson, AZ.

14) Horschler, D.J., Santos, L.R., & MacLean, E.L. (2019). How do non-human primates represent what others know? Talk presented at the 7th Annual Conference of the Southwestern Association of Biological Anthropologists, Tempe, AZ.
    ☨ Awarded prize for best student podium presentation

13) MacLean, E.L., Bray, E.E., Gnanadesikan, G.E., & Horschler, D.J. (2019). Ontogeny and heritability of cognitive and temperamental traits in an assistance dog population. Talk presented at the International Canine Science Conference, Tempe, AZ.

12) Horschler, D.J., Santos, L.R., & MacLean, E.L. (2019). Do non-human primates really represent others' ignorance? Talk presented at the 26th International Conference on Comparative Cognition, Melbourne Beach, FL.
    ☨ Selected as a finalist for the Ron Weisman Outstanding Student Presentation Award

11) Bray, E.E., Gnanadesikan, G.E., Horschler, D.J., & MacLean, E.L. (2019). Early emerging cognition in 9-week-old puppies. Talk presented at the 26th International Conference on Comparative Cognition, Melbourne Beach, FL.

10) MacLean, E.L., Bray, E.E., Gnanadesikan, G.E., & Horschler, D.J. (2019). Heritability of cognitive traits in a pedigreed dog population. Talk presented at the 26th International Conference on Comparative Cognition, Melbourne Beach, FL.

9) Gnanadesikan, G.E., Horschler, D.J., & MacLean, E.L. (2019). Social cues and hormonal profiles over development in wolf puppies. Poster presented at the GPSC Student Showcase, Tucson, AZ.

8) Horschler, D.J. & MacLean, E.L. (2018). Absolute brain size predicts dog breed differences in executive function and social cognition. Poster presented at the GPSC Student Showcase, Tucson, AZ.

7) Horschler, D.J. & MacLean, E.L. (2018). Dogs as a model for the cognitive implications of neuroanatomical variation. Talk presented at the Yale University East Coast Canine Cognition Workshop, New Haven, CT.

6) MacLean, E.L. & Horschler, D.J. (2018). Breed differences in dog cognition are predicted by absolute brain volume. Talk presented at the 126th Annual Convention of the American Psychological Association, San Francisco, CA.

5) Horschler, D.J. & MacLean, E.L. (2017). Absolute brain size predicts dog breed differences in executive function and social cognition. Poster presented at the UA-ASU Cognitive Science Conclave, Tucson, AZ.

4) Horschler, D.J. & MacLean, E.L. (2017). Absolute brain size predicts dog breed differences in working memory and social cognition. Poster presented at the North American Canine Science Forum, Tempe, AZ.

3) Horschler, D.J., Susser, J.A., & Mulligan, N.W. (2016). Metamemory for performed and observed actions: Within-subjects. Poster presented at the UNC Psychology and Neuroscience Honors Poster Symposium, Chapel Hill, NC.

2) Horschler, D.J., Susser, J.A., & Mulligan, N.W. (2016). Metamemory for performed and observed actions: Within-subjects. Poster presented at the UNC Celebration of Undergraduate Research, Chapel Hill, NC.

1) Horschler, D.J. (2014). Insights into user experience at Lenovo. Poster presented at the Karen M. Gil Internship Program Poster Symposium, Chapel Hill, NC.

Invited Talks

Collège de France, Decoupling and Mentalizing Workshop (2022)
University of Arizona, Spiritual and Scientific Approaches to Consciousness (2021)
Yale University, Philosophy of Language (2020)
Boston College, Canine Cognition Center and Social Learning Laboratory (2020)
University of Arizona, AGUA Student Lecture Series (2019)
University of Arizona, Grad Slam Competition (2019)
Yale University, Comparative Cognition Laboratory (2019)
University of North Carolina at Chapel Hill, The Senses of Animals (2015)

Selected Media Coverage


Commentary on Related Research

Teaching Experience

Fundamentals of Neuroscience and Cognitive Science (Spring 2020)
University of Arizona | Department of Neuroscience | School of Mind, Brain & Behavior
Graduate Teaching Associate | ~160 students

Animal Minds (Fall 2019)
University of Arizona | School of Anthropology | College of Social and Behavioral Sciences
Graduate Teaching Associate | Two sections (~30 students each)

Dog Thought (Spring 2019)
University of Arizona | School of Anthropology | College of Social and Behavioral Sciences
Graduate Teaching Associate | ~150 students

Research Methods in Cognitive Science (Fall 2018)
University of Arizona | Department of Neuroscience | School of Mind, Brain & Behavior
Graduate Teaching Assistant | ~110 students

Human Variation in the Modern World (Fall 2017 & Spring 2018)
University of Arizona | School of Anthropology | College of Social and Behavioral Sciences
Graduate Teaching Assistant | Two sections per semester (~30 students each)

Contact Information

Email Address
daniel_horschler [at] hillspet.com