Founder, Principal Investigator

Prof. Richard Connor, UMASS Dartmouth, USA

Richard is a Professor of Biology at the University of Massachusetts at Dartmouth. He obtained his Bachelor’s degree from the University of California at Santa Cruz, his Ph.D. from the University of Michigan. After obtaining his Ph.D., he had post-doctoral appointments at the Woods Hole Oceanographic Institution and Harvard University. He was a fellow at the Michigan Society of Fellows and at the Center for Advanced Study in the Behavioral Sciences at Stanford University.

Prof. Connor co-founded the Shark Bay dolphin research project in 1982 (with Rachel Smolker), is a Principal Investigator of the Dolphin Alliance Project (DAP) and President of an American Foundation, the Dolphin Alliance Project, Inc., that is seeking contributions to continue funding DAP research.

The main focus of Prof. Connor’s work in DAP has been dolphin behaviour and the males’ alliance relationships. He examines the alliance relationships from a broad, comparative perspective and has developed new theory on alliance formation and cooperation in general.

Prof. Connor has published over 75 scientific articles and book chapters on dolphins and his other major interest, the evolution of cooperation and mutualism. His research has been published in top scientific journals (e.g. Nature, Proceedings of the National Academy of Sciences, Proceedings of the Royal Society), featured in television documentaries, including the National Geographic Society’s Dolphins the Wild Side and NOVA’s The Private Lives of Dolphins, and reported in major news outlets (e.g. The New York Times, Science Online).

The Dolphin Alliance Project’s research has produced important discoveries with profound implications for understanding the evolution of large brains and intelligence, including that in humans. Accordingly, Prof. Connor has been invited to speak to academic audiences in a range of disciplines, including biology, anthropology, psychology and even political science, and at such prestigious institutions as Harvard, Yale, Oxford, the École Normale Supérieure in Paris and the Royal Society of London. He is available for public speaking engagements.

Principal Investigator

Prof. Michael Krützen, University of Zurich, Switzerland

Michael is a Professor of Evolutionary Anthropology and Genomics at the University of Zurich. His broad interests are in the social evolution of both primates and cetaceans. Using modern DNA techniques, his work focuses on relationships among cooperating individuals, measuring the number of paternities, and trying to link genetic population structure with social correlates. Prof. Krützen’s group at the University of Zurich also work on the demographic reconstruction of animal populations and delineation of conservation units.

Prof. Krützen has also been fascinated by the study of how knowledge is transferred in primates and cetaceans. His work on social transmission of tool use in dolphins in Shark Bay and orang-utans in Borneo and Sumatra has helped further our understanding of how culture has evolved in humans.

Recently, Prof. Krützen’s group has employed population genomics approaches to identify the genetic signatures of adaptive evolution in orang-utans and dolphins. Using landscape genomics approaches, this work aims to disentangle adaptive evolution from non-adaptive processes, such as genetic drift, by taking into account demographic, stochastic and environmental processes.

Principal Investigator

Dr. Simon Allen, University of Bristol, UK (University of Zürich, University of Western Australia)

Simon is a Senior Research Associate at the Universities of Bristol and Zurich, and an Adjunct Research Fellow at the University of Western Australia. His research interests include the complex social and foraging behaviours of dolphins, and assessing the impacts of fisheries, tourism and climate breakdown on marine fauna. Improved wildlife conservation and better management of the ways in which humans interact with wildlife are the end goals.

Dr. Allen has published over 50 scientific articles (as well as a few book chapters and species accounts) on dolphins, whales, reptiles and other fauna. He first ventured to Shark Bay in the year 2000, was left in awe, and has been returning ever since.

Simon is a PI at Shark Bay Dolphin Research, a keen field biologist, photographer and drone pilot, and holds general interests in behavioural ecology and conservation. You can follow his occasional rants about politics, the environment or how much he loves dogs on Twitter: @SimonJAllen1

Principal Investigator

Dr. Stephanie King, University of Bristol, UK

Dr. Stephanie King is a Senior Lecturer at the University of Bristol and a Branco Weiss Fellow (The Branco Weiss Fellowship – Society in Science). She is also an Adjunct Research Fellow at the University of Western Australia. Her research interests lie in understanding the role vocal communication plays in mediating complex social behaviours, such as cooperation, in animal systems. The nested structure of male alliance formation found in the Shark Bay dolphin population provides a unique opportunity to understand the interplay between vocal communication and cooperative strategies. Her group uses techniques such as hydrophone arrays, overhead video and sound playback experiments to explore the role of communication in the formation and maintenance of male alliances, and the communicative strategies these males employ when making decisions of when and with whom to cooperate.

Much of Dr. King’s earlier work has involved logistically challenging field experiments with bottlenose dolphins. She obtained the first evidence of a non-human mammal using learned signals as labels for individuals (published in Proceedings of the National Academy of Sciences), and that the formation of individual vocal labels takes precedence over vocal convergence in animal societies where biological markets are prevalent (published in Current Biology).

Dr. King’s research has received worldwide media coverage and has been the focus of TV documentaries (BBC’s Britain’s Secret Seas and Winterwatch), radio broadcasts (BBC, ABC and NPR) and has featured in many high-profile science magazine articles (e.g. New Scientist, Scientific American, and National Geographic, amongst many others).

Twitter: @_StephanieLKing

Research Associates

Dr. Delphine Chabanne, University of Zurich, Switzerland

Delphine’s research interests have primarily focused on understanding population ecology and genetics of coastal and estuarine dolphins. She is particularly interested in understanding spatial and temporal variation in population genetic structure. She completed her PhD thesis in July 2017, in which she assessed the demographic, social and genetic structure of Indo-Pacific bottlenose dolphins inhabiting coastal and estuarine waters of Perth, Western Australia. She has assisted on a number of projects involving coastal dolphin species around the vast Western Australian coastline.



Dr. Livia Gerber, University of Zurich, Switzerland

Livia completed her PhD on the ontogeny of alliance formation at the University of Zurich.

Using a genomics approach, she investigated whether alliance formation is based on relatedness or other factors, such as familiarity (see the publication page under 2021 and 2020). Livia will soon be taking on a post-doctoral fellowship through the University of New South Wales.

Dr. Katharina Peters, University of Zurich, Switzerland

Reproductive success is considered the hallmark of evolution, as transferring one’s genes into the next generation is the key determinant of evolutionary fitness. This project synthesizes unparalleled genomic and behavioural data of a long-lived mammal into a unique data set spanning 35 years, to compare lifetime reproductive success of males and females. There will be a special emphasis on male cooperation, as male bottlenose dolphins (Tursiops aduncus) in Shark Bay, Western Australia, form alliances on three different levels for the purpose of coercing females in reproductive condition. The complexity of such alliances is unparalleled outside humans. As yet, however, relatively little is known about how male cooperation translates into reproductive success and if, and to what extent, important parameters of life-time reproductive success vary between males and females.


Recent advances in statistical methodology and genomic analyses of wild animal populations allow us to integrate molecular age determination via epigenetic DNA methylation patterns with long-term behavioural and genetic data. For the first time, I will be able to directly compare lifetime reproductive success between males and females in this remarkable system. Moreover, I will finally be able to undertake a long overdue analysis that will investigate to which extend sociality and age are linked to male reproductive success, thereby investigating important proximate mechanisms explaining the evolution and maintenance of male alliance formation. Findings of this work improve our understanding of the evolution of life histories in one of the most complex cooperative behaviours outside our own species.

PhD Candidates

Samuel Wittwer, MSc
PhD candidate, Anthropological Institute and Museum, University of Zurich, Switzerland

Samuel Wittwer_JT
The nested alliance structure, where males work together to obtain and defend females, among bottlenose dolphins in Shark Bay is one of the most complex social structures observed outside of humans. This pattern of nested alliances is especially fascinating because the resource for which males cooperate (succesfully fathering offspring) is indivisible between the cooperating males. A male’s cooperative alliance partners are thus also his competitors. Apart from their fascinating social complexity, bottlenose dolphins are also an ideal system to study genomic patterns of local adaptation due to their global distribution and highly heterogeneous habitat. In his PhD project, Sam will take advantage of recent advances in Next Generation Sequencing technologies to look at genome scale questions in different populations. Does relatedness play a role in alliance formation? Are certain regions of the genome under divergent selection between populations? Are visible signatures of selection within the genome associated with alliance formation or other behavioural patterns?


Kathryn Holmes, MRes
PhD candidate, School of Biological Sciences, University of Western Australia, Australia

Although much has been learned about the alliances between sexually mature males in the Shark Bay population, we do not know how social development during the juvenile period (between the ages of weaning and early adulthood) shapes the formation of these alliances and affects the future social roles and fitness of individual males. For her PhD project, Katy will use acoustic, behavioural and genetic data to investigate the ontogeny of alliance formation, focusing on the development of vocal and physical behaviours that mediate social bonds, or become instrumental in adult consortships. She will also examine juvenile male social networks and specific dyadic associations, and further investigate the factors that contribute to alliance partner choice and male fitness.  



Pernille Meyer Sørensen, MSc
PhD candidate, School of Biological Sciences, University of Bristol, United Kingdom

The alliance structure of male bottlenose dolphins in Shark Bay varies as a function of the physical structure of the habitat. Whereas trios are most often seen in the northern, open water habitat, males usually form pairs in the southern habitat, characterized by shallow flats bisected by deeper channels. With sound being the primary sensory modality for all cetaceans, including bottlenose dolphins, Pernille will for her PhD explore how the acoustic habitat may  influence on alliance structure and mating success. To investigate this Pernille will collect underwater noise and sound propagation measurements, along with bottlenose dolphin source level estimations, to understand spatio-temporal variations in detection range. These estimates will be combined with long-term data on alliance behaviour to test how the acoustic characteristics of the habitat may affect conspecific encounter rates, consortship rates and acoustic behaviour within alliances.


Manuela Bizzozzero, MSc
PhD candidate, Anthropological Institute and Museum, University of Zurich, Switzerland

 The social structure and space use of dolphins differ with varying food availability and predation risk. In Shark Bay, the seascape represents a mosaic of different habitat types encompassing seagrass beds and sandy patches in shallower waters and deep sandy channels. Male bottlenose dolphins display a complex social structure that varies with changing habitats. Furthermore, the dolphins are known for their habitat dependent, versatile foraging techniques, some of which include tool use (with sponges and shells). For her PhD project, Manuela will use remote sensing satellite data to characterize different habitat types, and environmental DNA to investigate how food availability changes across the seascape. This will allow her to investigate how differences in food availability affect the social structure and habitat use of bottlenose dolphins.




Svenja Marfurt
PhD candidate, Anthropological Institute and Museum, University of Zurich, Switzerland

For her Master’s thesis Svenja investigated association patterns and community structure of female Indo-Pacific Bottlenose Dolphins (T. aduncus) in western Shark Bay (WA) using behavioral and genetic determinants.





MSc Students


Rufus Sowerbutts
M.Sc. candidate, University of Bristol, UK

Rufus graduated from University of Bristol with a BSc in Biology. For his Master’s thesis, Rufus will be exploring the social and ecological drivers of signature whistle complexity.



Emma Chereskin
M.Sc. candidate, University of Bristol, UK 

Emma graduated from the University of Vermont with a B.A. in Zoology. For her Master’s thesis, Emma will be examining the role of keystone individuals in information distribution in male bottlenose dolphin alliances.






Natalie Kelpacova
M.Sc. candidate, University of Bristol, UK 

Natalie graduated from the University of Bristol with a BSc in Biology. For her Master’s thesis, Natalie will be exploring the long term effects of extreme climatic events on the behavioural activity of Shark Bay’s iconic dolphin population.





External Collaborators

Dr. Sonja Wild

Postdoctoral Researcher, University of Konstanz & Max Planck Institute of Animal Behavior, Germany

Dr Sonja Wild is a postdoctoral researcher at the University of Konstanz & Max Planck Institute of Animal Behavior in Radolfzell, Germany, studying conditions that trigger switches from established to alternative behaviour in a social context using small passerine birds as a model system. Her main research interests lie in the mechanisms underlying the spread of behaviour in wild animal populations and how animals respond behaviourally to changes in the (social and physical) environment.

For her MSc degree at the University of Zurich (Switzerland) and her PhD at the University of Leeds (UK), Sonja spent 7 field seasons in Shark Bay to investigate the spread of foraging behaviour in the dolphin population by integrating behavioural, environmental and genetic data. She also investigated the impacts of a marine heatwave on the population’s demography.

Twitter: @wild_sonja


Rachel Smolker, PhD (co-founder of Shark Bay dolphin research with Richard Connor in the early 1980s)
Acoustic communication of bottlenose dolphins
PhD thesis, University of Michigan, Ann Arbor, USA


Amy Samuels, PhD (great field biologist and dearly missed mentor, colleague and friend) AmySamuelss_84448
Woods Hole Oceanographic Institute
Memories: Amy Samuels




Coco Ackermann, MSc
Contrasting vertical skill transmission patterns of a tool use behaviour in two groups of wild bottlenose dolphins (Tursiops sp.), as revealed by molecular genetic analyses
MSc thesis, University of Zurich, Switzerland


Kathrin Bacher, MSc
The evolutionary and ecological mechanisms underlying the rise of material culture in bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia

MSc thesis, University of Zurich, Switzerland


Lynne Barre, MSc
Dedicated field biologist in the 1990s and coauthor of several important papers with Prof. Richard Connor.


Lars Bejder, PhD   
Linking short and long-term effects of nature-based tourism on cetaceans
PhD thesis, Dalhousie University, Canada


Manuela Bizzozzero, MSc
Alliance formation and sponge tool use in male bottlenose dolphins (Tursiops sp.)
MSc thesis, University of Zurich, Switzerland


Riccardo Cicciarella, MSc
Long-term movements of Shark Bay bottlenose dolphins
MSc thesis, UMASS Dartmouth, USA


Whitney Friedman, PhD
Social and Cognitive Complexity in Wild Bottlenose Dolphins (Tursiops sp.)
PhD thesis, University of California, San Diego, USA


Livia Gerber, PhD
The Influence of Social and Genetic Relationships on Cooperation in Male Bottlenose Dolphins
PhD thesis, University of Zurich, Switzerland


Maja Greminger, PhD
The quest for the Y ─ Development and application of male-specific markers in orangutans (Pongo spp.) and bottlenose dolphins (Tursiops spp.)

MSc thesis, University of Zurich, Switzerland


Mike Heithaus, PhD
PhD thesis, Simon Fraser University, Vancouver, Canada


Kay Horlacher, Msc
Reproductive success in male bottlenose dolphin alliances in Shark Bay, Western Australia
MSc thesis, University of Zurch, Switzerland


Stephanie Kalberer, MSc
Relatedness patterns
in male bottlenose dolphin alliances in Shark Bay, Western Australia

MSc thesis, University of Zurch, Switzerland


Anna Kopps, PhD
Ecological, social and genetic forces shaping behavioural variation in bottlenose dolphins
Ph.D. thesis, The University of New South Wales, Sydney, Australia
Who is the one? Paternity assessment in bottlenose dolphins (Tursiops sp.) – a species with multi-level alliances

MSc thesis, University of Zurich, Switzerland


Sina Kreicker, MSc
Culturally transmitted tool use in bottlenose dolphins (Tursiops sp.) – utilization of an unexploited niche?

MSc thesis, University of Zurich, Switzerland


Svenja Marfurt, MSc
Using social network analysis to investigate association patterns and community structure of female Indo-Pacific bottlenose dolphins (Tursiops aduncus)
MSc thesis, University of Zurich, Switzerland

Oliver Manlik, PhD
Fitness and major histocompatibility complex diversity of two bottlenose dolphin populations
PhD thesis, University of New South Wales, Australia


Séverine Miszak, MSc
Population Genetics and Phylogenetic Analyses of Australian Bottlenose Dolphins (Tursiops spp.) using reduced representation libraries
MSc thesis, University of Zurich, Switzerland


Bronte Moore, MSc
Vocal synchrony in allied male Indo-Pacific bottlenose dolphins (Tursiops aduncus)
MSc thesis, University of Western Australia, Australia


Alexander Nater, PhD
New microsatellite markers in orang-utans (Pongo spp.) and bottlenose dolphins (Tursiops spp.)

MSc thesis, University of Zurich, Switzerland


Krista Nicholson, MSc
Abundance, survival and temporary emigration of bottlenose dolphins (Tursiops sp.) in the Western gulf of Shark Bay, Western Australia

MSc thesis, Murdoch University, Perth, Australia


Orla O’Brien, MSc
Alliance-specific habitat selection by male dolphins in Shark Bay, Western Australia
MSc thesis, UMASS Dartmouth, USA


Haley Pedersen, MSc
Population level assessment of BaitSTR performance – a test case in deep-pedigree reconstruction of Indo-Pacific bottlenose dolphins using genomic microsatellites
MSc thesis, University of Zurich, Switzerland


Srđan Randić, MSc 
Spatial analysis of distribution and home ranges of male bottlenose dolphins in Shark Bay, Australia.
MSc thesis, UMASS Dartmouth, USA


Andrew Richards, PhD
Life history and behavior of female dolphins (Tursiops sp.) in Shark Bay, Western Australia
PhD thesis, University of Michigan, USA


Julian Tyne, PhD
Does sponge distribution lead to sponging behaviour by bottlenose dolphins in Shark Bay?

BSc(Hons) thesis, Murdoch University, Perth, Australia


Sonja Wild, PhD
Social transmission of foraging behaviour in bottlenose dolphins and its interplay with climate change
PhD thesis, University of Leeds, UK