Scott -- The Mullard design (which you are using) has been as roundly praised for its performance capabilities as it has been criticized for its unbearably excessive gain.
To put it into perspective, the same Mullard circuit you are using is the basis for many of the Eico amplifiers -- except that Eico wisely used a 6SN7 in place of the 12AX7 for inverter service that not only provided a lower impedance (i.e. better) drive for the output stage, but also cut a big chunk of the gain out of that circuit. But still -- even with that modification, the Eico amplifiers still require only .5 volt input to generate full power output. This is still too sensitive for my tastes, causing problems with noisy interconnects and amplifying the quiescent noise from the preamp. As a result, Eico still had to provide input level controls to contain the still excessive gain from the design.
In your case, the gain issue is a bit of a big elephant in the middle of the room. Even with the FB issues dealt with, you're still left will all that gain. To that point, in my opinion, using large attenuation networks ahead of the input stage is poor design. It's a huge invitation to reducing HF performance and loss of detail. More directly, why produce all that gain, only to have to compensate for it (i.e attenuate it) somewhere else?
If it is the problem for you I think it is, it should really be dealt with first -- and properly -- before moving forward. If it is to be dealt with through basic circuit design, then just like using the actual OPT when developing the feedback/stability circuits, the actual amplifying block should be used as well. Otherwise, just like before, you've got to do it all over again when the actual circuit is used.
I would strongly consider converting your 12AX7 inverter tube to a 6FQ7/6CG7 (basically a miniature version of the 6SN7). This tube uses the same size socket, and can be converted with simple rewiring. It will require new components associated with that tube, but it will also be a big step in the right direction regarding this problem. Then, if the gain is still excessive for you, we can reconfigure the input stage to operate in triode mode. This -- along with the inverter change -- will deal a serious blow to all that unnecessary gain, and almost certainly eliminate the need for any input attenuation circuits -- which you would not want to use with a triode type input tube anyway.
Regarding the function of the step network components, both components work together in harmony -- in conjunction with the circuit impedance present -- to control not only the point at which the HF begins to roll off, but also the amount of overall attenuation the network provides at those frequencies. Therefore, a change in one component within the network requires a change in the other component to achieve both goals being targeted. For example, if a given network provides the correct roll off frequency, but not enough attenuation, then reducing the R component will provide more attenuation, but also change the "corner" frequency of the network. So, the C component must then be adjusted to compensate, etc., -- and so it goes.
As I said, this is quite an ambitious project, but one if you see through, you will also learn a lot from.
Dave