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Proton
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They are trackers embedded into emails and images. Not from proton itself.
And it looks like part of that might be with how proton strips known trackers to remove some parameters.
Also worth noting that Proton opens any images or other elements server-side, so senders can only "track" Proton servers.
Right, you don't have to interact with an email to have trackers. Just opening it is enough, and trackers can be embedded in images or other elements.
I don't know much about Proton, but it may have features to prevent loading those elements.
I use Thunderbird, which optionally blocks those elements, and can work with Proton on some devices.