Founder, Principal Investigator

Prof. Richard Connor, UMASS Dartmouth, USA

Connor_JTProf. Richard Connor 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., Prof. Connor 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 program in 1982 (with Rachel Smolker), is co-Director of the Dolphin Alliance Project (DAP) and President of an American Foundation, the Dolphin Alliance Project, Inc., that is currently 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, Dr. 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

Kruetzen_JTProf. Michael Krützen 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 and University of Zürich, Switzerland

Dr. Simon Allen is a Senior Research Associate at the University of Bristol and University of Zurich1 Profile pic. He completed an Australian tour of tertiary institutions, having gained his B.Sc. from Flinders University, his B.Sc. (Hons.) from the University of Queensland, his M.Sc. from Macquarie University and his Ph.D. from Murdoch University. His  research interests lie in studying the complex social and foraging behaviours of dolphins, and assessing the impacts of human activities (fisheries, tourism, coastal development and climate change) on marine fauna in general. Improved wildlife conservation and better management of the ways in which humans interact with wildlife is the end goal.

Dr. Allen has thus far published over 40 scientific articles and book chapters on dolphins, whales, reptiles and other fauna in journals and texts ranging from the taxon- and issue-specific (for example, Marine Mammal Science and Whale-Watching: Sustainable Tourism and Ecological Management) to the broad (Proceedings of the Royal Society BMolecular Ecology, Current Biology and Scientific Reports). He first ventured to Shark Bay, now one of his favourite places on the planet, in the year 2000 and has been returning ever since, including leading the 2005 Dolphin Alliance Project field season for Drs. Connor and Krützen, and running the inaugural 2007 Dolphin Innovation Project field season. 

Simon is now a PI of the Shark Bay Dolphin Research Alliance. He is a very keen field biologist, photographer and drone pilot, and holds a general interest in the behaviour, ecology and conservation of both marine and terrestrial wildlife. He would like to be head-hunted for a photo-journalism gig, or perhaps a fireman, when he eventually grows up. You can follow his occasional rants about politics, the environment or how much he loves dogs on Twitter: @SimonJAllen1

Principal Investigator

DSC_5390Dr. 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). 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. Whitney Friedman, University of California Santa Cruz

W Friedman_JT
Long-term behavioural observation in Shark Bay has revealed a nested structure of male alliances that is stable across the population, but often dynamic at the individual level. How is this complex social structure mediated through interactions among individuals?

Historically, such questions have been difficult to address as many interactions occur partially or completely underwater. For her dissertation research, Whitney employed a combination of aerial videography, underwater acoustic recordings, and boat-based observation to record fine-scale detail of interaction among 17 males within the Shark Bay population as they engage in first-, second-, and third-order associations.


Dr. Delphine Chabanne

Delphine’s research interests have primarily focused on understanding population ecology and genetics of coastal and estuarine dolphins. She is particularly interested in understanding the 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.



PhD Students

Livia Gerber, MSc

LG_JTPhD candidate, Anthropological Institute and Museum, University of Zurich, Switzerland

Male alliances in bottlenose dolphins are among the most complex social structures found outside humans and provide a rare example of male cooperation. However, little is known about their formation and how males choose their partners. Cooperative behaviours are often favoured among kin, as they allow the actors to benefit from inclusive fitness benefits. Whether kin selection explains alliance formation could not be answered reliably using classic genetic markers.

For her PhD project, Livia will apply a genomics approach, which allows us to investigate whether alliance formation is based on relatedness or other factors, such as familiarity.



Samuel Wittwer, MSc

Samuel Wittwer_JT

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

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?


Sonja Wild, MSc

PhD candidate, Leeds University, United Kingdom

Sonja Wild is a former M.S. student at the Anthropological Institute and Museum, University of Zurich, and now a Ph.D. student at Leeds University, U.K. Her interest lies in the emergence of animal culture and the process through which behavioural traits are passed on among individuals. Due to their cognitive abilities and highly social nature, bottlenose dolphins depict an ideal species to approach questions on how novel behaviours spread within wild animal populations.

During her M.S. studies, Sonja focused on a foraging strategy called “shelling”. By combining behavioural and ecological data with molecular techniques, she was able to assess the relative importance of social learning, ecological variation and relatedness in the spread of the shelling behaviour.



Kathryn Holmes, MRes

PhD student, 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 Student, 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.


MSc Students

Svenja Marfurt

M.Sc. Student, Department of Anthropology, University of Zurich, Switzerland

The high complexity of social structures found in bottlenose dolphins (Tursiops sp.) are comparable to those found in primates, including humans. Although the general social pattern of a dolphin population is characterized by a fission and fusion system, preferred associations between individuals can lead to community structure within populations. While male bottlenose dolphins in Shark Bay engage in long-term associations and form highly nested alliances, association patterns in female bottlenose dolphins are not yet well understood.

In her master thesis, Svenja will investigate association patterns and community structure of female Indo-Pacific Bottlenose Dolphins (T. aduncus) in Western Shark Bay (WA) using behavioral and genetic determinants. She specifically aims to examine the role of maternal and biparental kinship as well as similarity of reproductive state and foraging technique in female association patterns and community structure.


Séverine Miszak

M.Sc. Student, Department of Anthropology, University of Zurich, Switzerland

Séverine completed her undergraduate in Biology at the ETH Zurich. With a special interest in marine biology, she switched to the Evolutionary Genetics group at the University of Zurich for her Master’s. In her Master’s thesis she is focusing on reconstructing the demographic history of australian Tursiops populations in South Western Australia.


Haley Pedersen

M.Sc. Student, Department of Anthropology, University of Zurich, Switzerland

Haley’s Master’s thesis will focus on implementing recent advancements in genome-scale STR marker development to assemble pedigrees for the Shark Bay dolphin populations.  

Prior to joining Prof. Krützen’s group, Haley completed her undergraduate degree at the University of Alberta, specializing in Animal Behaviour. She has also worked in the fields of cancer biology, and wildlife conservation.


Riccardo Ciccarella

M.Sc. Student, UMASS Dartmouth, USA

Riccardo graduated from Humboldt State University, CA, with a B.S. in Marine Biology. After studying pink and coho salmon in Alaska during his undergraduate career, he became interested in animal behavior and social interactions.  He will be working on the behavior of allied males.


External Collaborators

Dr. Will Hoppitt, Leeds University, United Kingdom

dr_william_hoppitt.Maincontent.0007.ImageGal..134.Image.~etc~medialib~fst_2009~staff_2014~~~Par~~0048~~ImageDr. Hoppitt’s main research interests concern the development of statistical methods to study social transmission of behavioural traits in groups of animals, and the application of these methods to data from groups of wild and captive animals. Will obtained a first degree in Biological Sciences in 1999 and a Masters degree in Integrative Biosciences in 2000, both from Oxford University. He then moved to Cambridge University to do a Ph.D. with Kevin Laland on Social Processes Influencing Learning: Combining Theoretical and Empirical Approaches.

After a brief spell as a safari guide in Botswana, Will continued his research on social learning in non-human animals, again in Kevin Laland’s research group, now at St. Andrews University in Scotland. It was here that Will started his work on methods for studying social learning in a field setting, most notably extending the novel technique, network based diffusion analysis (NBDA). He worked with a wide range of researchers to apply NBDA to a number of different species, showing strong evidence of social transmission of a novel foraging technique in humpback whales, and of novel tool use behaviour in wild chimpanzees. Will spent a period of three years as a Senior Lecturer in Zoology at Anglia Ruskin University (UK, 2012-2015), before moving to his current position as a Lecturer in Zoology at Leeds University.



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 (legendary 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


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


Oliver Manlik, PhD

Fitness and major histocompatibility complex diversity of two bottlenose dolphin populations

PhD thesis, University of New South Wales, Australia


Bronte Moore

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


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