G’day Marmamers,

We are most pleased to bring to your attention two papers on dolphin social 
complexity in the forthcoming issue of Current Biology, both stemming from our 
long-term research programme in Shark Bay, Western Australia 
(www.sharkbaydolphins.org<http://www.sharkbaydolphins.org>).

In the first paper, led by University of Bristol MSc graduate Emma Chereskin, 
we show that vocal exchanges can function as a replacement of physical bonding 
in dolphin alliances. This is the first evidence for Robin Dunbar’s social 
bonding hypothesis and, interestingly, comes from outside of the primate 
lineage.

Paper 1: Chereskin E, Connor RC, Friedman WR, Jensen FH, Allen SJ, Sørensen PM, 
Krützen M, King SL (2022). Allied male dolphins use vocal exchanges to 
‘bond-at-a-distance’. Current Biology https://doi.org/10.1016/j.cub.2022.02.019

Summary: Vocal interactions are intrinsic features of social groups and can 
play a pivotal role in social bonding. Dunbar’s social bonding hypothesis 
posits that vocal exchanges evolved to “groom at a distance” when social groups 
became too large or complex for individuals to devote time to physical bonding 
activities. Tests of this hypothesis in non-human primates, however, suggest 
that vocal exchanges occur between more strongly bonded individuals that engage 
in higher grooming rates and thus do not provide evidence for replacement of 
physical bonding. Here, we combine data on social bond strength, whistle 
exchange frequency, and affiliative contact behavior rates to test this 
hypothesis in wild male Indo-Pacific bottlenose dolphins, who form multi-level 
alliances that cooperate over access to females. We show that, although whistle 
exchanges are more likely to occur within the core alliance, they occur more 
frequently between those males that share weaker social bonds, i.e., between 
core allies that spend less time together, while the opposite occurs for 
affiliative physical contact behavior. This suggests that vocal exchanges 
function as a low-cost mechanism for male dolphins that spend less time in 
close proximity and engage in fewer affiliative contact behaviors to reinforce 
and maintain their valuable alliance relationships. Our findings provide new 
evidence outside of the primate lineage that vocal exchanges serve a bonding 
function and reveal that, as the social bonding hypothesis originally 
suggested, vocal exchanges can function as a replacement of physical bonding 
activities for individuals to maintain their important social relationships.

In the second paper, led by University of Zürich PhD graduate Livia Gerber, we 
show that ‘popular’ allied male dolphins (those that are well-integrated and 
have homogenous social bonds with their allies) enjoy higher reproductive 
success.

Paper 2: Gerber L, Connor RC, Allen SJ, Horlacher K, King SL, Sherwin WB, 
Willems E, Wittwer S, Krützen M (2022). Social integration influences fitness 
in allied male dolphins. Current Biology. 
https://doi.org/10.1016/j.cub.2022.03.027

Summary: Understanding determinants of differential reproductive success is at 
the core of evolutionary biology because of its connection to fitness. Early 
work has linked variation in reproductive success to differences in age, rank, 
or size, as well as habitat characteristics. More recently, studies in 
group-living taxa have revealed that social relationships also have measurable 
effects on fitness. The influence of social bonds on fitness is particularly 
interesting in males who compete over reproductive opportunities. In Shark Bay, 
Western Australia, groups of 4-14 unrelated male bottlenose dolphins cooperate 
in second-order alliances to compete with rival alliances over access to 
females. Nested within second-order alliances, pairs or trios of males, which 
can vary in composition, form first-order alliances to herd estrus females. 
Using 30 years of behavioral data, we examined how individual social factors, 
such as first-order alliance stability, social connectivity, and variation in 
social bond strength within second-order alliances, affect male fitness. 
Analyzing the reproductive careers of 85 males belonging to 10 second-order 
alliances, we found that the number of paternities a male achieved was 
positively correlated with his cumulative social bond strength but negatively 
correlated with his variation in bond strength. Thus, well-integrated males 
with more homogeneous social bonds to second-order allies obtained most 
paternities. Our findings provide novel insights into the fitness benefits of 
polyadic cooperation among unrelated males and highlight the adaptive value of 
social bonds in this context.

If you’d like a PDF or have any queries, please don’t hesitate to reach out to 
the primary authors 
(emma.cheres...@bristol.ac.uk<mailto:emma.cheres...@bristol.ac.uk> and 
livia.ger...@uzh.ch<mailto:livia.ger...@uzh.ch>).

All the best,



Simon

~~~~~~~~~~~~~
Dr Simon J Allen
Senior Lecturer
School of Biological Sciences
University of Bristol

Mob: +44 (0) 77047 53101 [UK] / +61 (0) 416 083 653 [AU]
Email: simon.al...@bristol.ac.uk<mailto:simon.al...@bristol.ac.uk> / 
simon.al...@uwa.edu.au<mailto:simon.al...@uwa.edu.au>
Web: http://www.sharkbaydolphins.org
Twitter: @SimonJAllen1

 [Chart, scatter chart  Description automatically generated]

Recent papers: Estimating sustainable limits to human-caused wildlife mortality 
https://conbio.onlinelibrary.wiley.com/doi/abs/10.1111/cobi.13897
Cooperation-based concept formation in bottlenose dolphins 
https://www.nature.com/articles/s41467-021-22668-1
Non-vertical transmission of a dolphin foraging innovation 
https://www.cell.com/current-biology/fulltext/S0960-9822(20)30756-9
Declines in dolphin survival and reproduction following a heatwave 
https://www.cell.com/action/showPdf?pii=S0960-9822%2819%2930217-9

“Believe me, my young friend, there is nothing–absolutely nothing–half so much 
worth doing as simply messing about in boats.” (Kenneth Grahame)
“I must say here, in passing, that those captains who have scientists… aboard 
their ships, must take with them a good supply of patience. I admit that 
although I have no lack of it, the scientists have frequently driven me to the 
end of my tether...” (Nicolas Baudin)



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