Peer review of: Network-based diffusion analysis reveals context-specific dominance of dance (2020)

Reviewers' Comments: 
 
Reviewer #1: 
Remarks to the Author: 
This manuscript presents a very interesting experimental study on waggle dances of honey bees. The 
main focus lies in assessing the relative importance of alternative social information sources for (1) 
locating novel feeding sites and (2) re-activating known feeding sites. The results show that the 
discovery of novel feeding sites is mainly mediated by dances and re-activation of known feeding sites 
is mainly mediated by olfactory information, specifically by antennation. 
 
The conceptual framework and the statistical analyses are both based on communication networks. I 
think this technically a valid approach. However, the analyses and the manuscript as a whole could be 
simplified a lot by removing all or most parts related to networks. On the conceptual side I do not see 
any benefit in talking about “communication networks” instead of simply referring to different 
“information sources”. 
 
The statistical analyses were conducted by applying network-based diffusion analysis (NDBA). This 
application is technically correct. However, the data that has been collected in this study is so detailed 
that the involvement of networks becomes unnecessary. To understand this point it is useful to 
consider the original purpose of the NBDA. Essentially, NBDA had been developed to deal with 
incomplete information on social transmission opportunities. Usually this information is very difficult to 
collect and direct data on social transmission opportunities is not available. To deal with this problem 
NBDA essentially estimates the missing information. Specifically, NBDA combines information on social 
behavior (in form of networks) and individual trait acquisition to estimate individuals-specific changes 
in social transmission opportunities over time. In the current study it is not necessary anymore to 
estimate social transmission opportunities, because these opportunities are directly observed and 
recorded (e.g. the number of dances). The application of NBDA in this case means that information on 
social transmission opportunities is first put into a network, and then the same information is 
extracted again from the network in order to fit social transmission models. This whole process could 
be simplified by leaving out the network part and directly using recorded information on social 
transmission opportunities (number of dances etc.) for fitting alternative transmission models. As an 
example, the authors first recorded the number of dances for each demonstrator and then summed 
them up across demonstrators. The same could be achieved by ignoring the identity of the 
demonstrator from the beginning and just counting the total number of dances for each individual. 
 
 
Minor points 
 
Title: “in situ dominance” seems to contradict the results because the estimated influence of dancing 
on re-activating is weaker than the influence of antennation. Furthermore, I suggest adding 
information in the title that the study was performed on honey bees. 
 
L 106-107: I do not understand why it was surprising that olfactory information use did not predict 
arrival times at the novel feeder. It seems to me that the design of first experiment eliminates the 
possibility for using olfactory information in a meaningful way. Thus, this experiment seems to test 
whether information acquired through dances is used when olfactory information provides no 
information. It would be useful to explain this, or alternatively explain how it would be mechanistically 
possible that olfactory information can speed up arrival at the novel feeder in this experimental 
setting. 
 
Figure 1: The networks seem to be completely uninformative. I suggest removing them or otherwise 

explain what information the reader can extract from them. 
 
I wonder whether additional analyses would allow gaining insights on whether and how different 
information sources are combined. Currently all information sources are assumed to exert independent 
influences because it is assumed that all effects on the rate of arriving at a feeding site are additive. 
In principle it would be possible to fit models that include statistical interactions between different 
information sources, which could represent synergistic effects. 
 
 
 
Reviewer #2: 
Remarks to the Author: 
In “Network-based diffusion analysis reveals in situ dominance of dance communication networks”, 
Hasenjager et al. use the aforesaid mentioned technique to analyze honey bee forager interactions 
from a field trial to determine if waggle dances, antennation (olfactory info transfer), or trophallaxis 
(olfactory and reward info transfer) are best at predicting both recruitment, reactivation, or both. They 
find that recruitment to novel locations strongly is associated with waggle dance interactions and not 
the other two types of interactions. Reactivation, in contrast, seems to be more associated with 
antennation. 
 
I liked this paper very much because of its combination of techniques and, most importantly, because 
it provides support for the importance of the waggle dance in a field-realistic context, something that 
has been not well understood in the past. 
 
I really have no major concerns about the paper – I think the experiment and the analysis are both 
sound, and the discussion is thorough. My only medium concern is the scope and content may be less 
interesting to folks outside of honey bee foraging ecology or, at best, animal behavior in general. It’s 
hard for me to say that because I find the paper extremely interesting and exciting. 
 
I have a few smaller comments that I hope the authors will consider that might help with the general 
readability of the paper. Perhaps too these comments might help increase its appeal to a broader 
readership. 
 
1. Title – the title does little to convey what’s going on to anyone unfamiliar with NBDA. In fact, 
having both the NBDA and the “in situ” made me think this was just a modelling paper without a field 
experiment. We don’t even know the organism! The word “network” is used twice, which always feels 
clunky to me. Consider a new title that is more informative, doesn’t overly rely on the analysis method 
at the expense of other, important info (organism!), and that conveys the fundamental take-home of 
the paper – that there are contexts in which the dance communication is dominant. 
 
2. Ln 19 – for those of us that work with the waggle dance, the word “direction” has a very specific 
meaning: the dance communication encodes a distance and a direction to a resource. So when we 
want to use one word to describe what the dance provides, we usually say “spatial” or “location” (and 
indeed, the authors also use “spatial”, see Ln 272). Consider replacing “directions” with “location 
information” 
 
3. Ln 25 – remove comma before “because” 
 
4. Ln 38 – very good explanation of NBDA – understandable to someone who hasn’t yet heard of it 
 
5. Ln 42 – “trait” – is it more of a behavioral event? “Behavioral trait” generally means something else 

to behavioral ecologists. 
 
6. Table 1 – For trials 3 and 4, you have very few recruits to the full feeder. Do you have any thoughts 
why? If so, can you discuss them in the discussion? 
 
I suspect that, once you get to mid-September, there actually are other foraging resources available, 
and this might make the bees less likely to be recruited to another site. You talk in your discussion 
how you train during autumn b/c there’s a forage dearth, but your citations are from Seeley and von 
Frisch, who definitely say that training is easier when nectar is in short supply, but say nothing about 
the seasonality of forage availability for southern England. 
 
In particular, ivy is a very important autumn nectar flow for your part of the world, as been shown 
with research from Sussex. Please see (1) Garbuzov & Ratnieks (2013) Insect Conservation and 
Diversity (2) Couvillon et al (2014) Plos One for better references re: forage availability in autumn. 
 
7. Fig 1: I found most of figure 1 to be very helpful, and indeed I referred to it many times to make 
sure I understood the experimental design. The only notable exception was 1e – there’s nothing to be 
gained here at all. What am I looking at? I know what a general network looks like – this adds nothing 
new. Does the dance versus trophallaxis versus antennation network look different? Is there a better 
way to represent what you are trying to show? 
 
Consider replacing 1e with something more meaningful to the reader – a picture of a waggle dance 
versus antennation versus trophallaxis. Someone not directly working with the waggle dance might 
find that much more useful than an unnamed / unlabeled network. 
 
8. Ln 113 – you throw in “asocial” learning, but a reader might not know what you mean. Can you 
define social versus asocial (including examples both from the paper and from other model systems)? 
 
9. It struck me that one good example of how the odor issue dominated people’s thinking of the 
waggle dance is the dance controversy (i.e., Adrian Wenner). For fun, read Munz (2005) Journal of 
History of Biology. And you can maybe mention a line in your discussion how, in some ways, your 
results really put the final nail in the coffin of the idea that odor alone is sufficient and necessary to 
recruit new foragers to a novel site (Ln 130 it is mentioned, but perhaps you can say something in Ln 
303). I think the “dance controversy” underscores how contentious the conversation was about the 
benefit / use of the dance. 
 
10. Ln 131 – I thought it was interesting that the number of waggle runs followed predicted 
recruitment order. We know from Seeley that a recruit has a greater chance of following a novel site if 
the recruit follows more waggle runs. So your results suggest a similar dose-dependent effect of the 
dance? That strengthens too your main argument. Can that been seen in figure form? Would be nice 
to have some data graphs - 
 
11. Ln 137 – “differing degrees” – how so? Can you explain? 
 
12. Ln 146 – so the more persistent bees (see Al Toufalia (2013) Ethology, it’s the ones more 
determinedly visiting the EMPTY feeder) are less likely to be recruited to new site? That seems very 
interesting to me. 
 
13. Ln 352 – Again, consider referencing something specific to your area (see comment 6). 
 
14. Ln 376 – clearly your experimental procedure worked, but just a note of interest – 50 uL / L is a 

strong scent! We usually used 10 uL / L – very strong scent can inhibit dancing. 
 
15. Lastly – is there a way to broaden the readership of the paper so it is interesting to people outside 
of bee foraging ecology and communication / self organization? Persistence of presumably costly 
behavior when the adaptive benefit has been unclear? 
 
Well done on a very interesting read! 
 
Kind regards, 
Dr. Margaret Couvillon 
 
 
 
Reviewer #3: 
Remarks to the Author: 
Review: 
 
This paper describes an elegant experiment that uses social network analysis to disentangle the 
relative contributions of waggle dances, olfactory cues and nectar donation to 1) the discovery of 
novel food sites, and 2) rediscovery/return to previously rewarding sites. To do so, the authors apply a 
modified form of network-based diffusion analysis (NBDA), comparing the fit between the order of 
arrival of bees to the food sites with social networks built from observations of interacting bees at the 
hive. 
 
NBDA is a well-established modelling approach to investigating the transmission of information. It is 
more usually used with a single information network, and I haven’t previously seen it used to compare 
alternative information networks in quite this way (Farine et al. 2015 possibly excepted). However, it 
seems to be a logical extension of the approach and I can see lots of potential future applications 
following a similar methodology to disentangle alternative possible information pathways. 
 
In general, I think the paper is well written, of broad interest, and well suited to publication in Nature 
Communications. I think the first experiment (discovery of new feeder) is excellent with a very clear 
result, and I only have minor comments requiring clarification. The second experiment (reactivation) 
is a little harder to understand, and I would appreciate more information/clarity on the experimental 
design and analysis. 
 
Experiment 1: 
 
1) Could the authors explain why they had an experimental design that involved the paired “empty” / 
“full” feeders, rather than simply one novel food site that some proportion of individuals had been 
trained on? At L50 they state it is “context mimicking natural depletion of one patch…followed by 
discovery of another”, but it is not clear to me why it this empty site is necessary at all to help answer 
the central question, that is, what information individuals use to discover new food sources. 
 
2) Why were there so few recruits to the full feeder for the third and fourth trials? These two trials 
appear to much fewer bees discovering the feeder (e.g. 4 in the third trial vs. 34 in the second). Does 
this have any effect on the analysis? 
 
3) Table 1 - I was scratching my head over this for a while to figure it out. I’m not convinced this table 
is needed in the main text at all, but if the authors want it, can it be simplified? For example, is it 
really necessary to know the mean air temperatures, given there was likely a lot of variation over the 

day? 
 
4) Did the bees arrive to the new food sites in groups, or follow each to food sources outside the hive? 
Were these possible “out of hive information transfers” controlled for in any way in the NBDA model? 
 
Figure 1 – these are pretty uninformative networks. I would suggest that instead of showing 4 
‘dancing’ networks, you instead show 3 networks for one site as an example: 1 x waggle dance, 1 x 
trophallaxis, 1x antennal (you could include all the networks for the other three sites in the 
supplementary). That would allow the reader to compare and contrast the networks derived these 
different interactions. 
 
Experiment 2: 
 
I find this experiment a little confusing – but if I can attempt to summarise, previous to the trial, the 
two feeders are equally profitable and provide identical scent cues. During the trial, only one is baited, 
but all decisions to return to either feeder are recorded as reactivation events. 
 
1) If reactivation to either feeder was included in the analysis, what was the thinking behind only 
filling one feeder? Why not just record individuals returning to 1 previously rewarding feeder, or even 
2 previously rewarding feeders? 
 
2) The authors state 173 that some individuals were already present and waiting at the familiar 
feeders, having presumably “reactivated themselves” based off their memory, and without any extra 
social information. This process presumably continued on throughout the trial. Can the authors make 
it clearer how they accounted for this personal information use operating parallel to the social 
information? I realise that they included a model that only had asocial effects (L 187), but it doesn’t 
seem to be me to be a sufficient control to assume it is either one or the other – rather having 
extensive personal information use would presumably be making their results much “messier” and 
more difficult to interpret. I confess I’m finding it difficult to see how the model could accurately 
estimate all these information sources (including bees dancing for natural sources as well!), with 
multiple potentially information users and information providers. More justification/information would 
be appreciated. 
 
3) The hypothesis stated on line 178-180 is that olfactory and motivation information provided by 
return successful foragers would be relevant for reactivation to either feeder. Presumably it also 
follows that waggle dances would only be relevant for the full feeder. 
 
Is it therefore the first prediction that kinds of different information use should lead different numbers 
of forages presenting at the full and empty feeder? After I read this paragraph I expected the first 
presented result to the proportion/numbers of individuals returning to each site, with a bias to the full 
feeder indicating a role for the spatial information provided by the waggle dance. While in L228 the 
authors include the feeder status in the NBDA model, it seems to me that the raw data is also 
informative. Can you provide this information? 
 
4) L242 – why were individuals trained to the empty feeder slower to reactivate? 
 
Signed, 
Lucy Aplin