Matt,
I have no idea why anyone would cite that paper either for (or against) anything. It is one of the worst pieces of "research" I have ever seen published on tortoises. That said, plenty of people have accepted it without question:
For example, writing in “Biology of Turtles” (CRC Press, 2008), Pritchard states “current belief is that the condition derives from excessively dry conditions” and that “when hatchlings are raised on a substrate of wet sphagnum, shells will develop normally” (Biology of Turtles, pp. 70).
I love it. Wet sphagnum! Here?
We have just had the wettest winter for many years. It even rained a bit this morning, so, I popped out a couple of hours ago, right after my last post, to some tortoise habitat just to show you all what the ACTUAL relative humidity is, even among the vegetation used by Testudo graeca:

There you go.
I also have recorders running right down in the roots. The tortoises, including juveniles, are out and active. Basking and feeding. You can see that the air temperature is just below 21C. However, the shell temperature of the tortoises was as high as 37 C. Their core temperatures are in the 28-32 C range.
As a generalisation, the RH among the plant roots just below the surface where juveniles hide, tends to be around the 60% level.
This is the type of microclimate we're talking about (photographed a couple of weeks ago):

I have a lot of very reliable data on this. I can say with total confidence that the assertion made in that paper that "“many species of tortoises live most of their lives hidden either in caves (e.g. Testudo horsfieldii, Gopherus sp.) or on the ground surrounded by high humidity even during arid conditions” and that "areas with a humidity of nearly 100% for hiding should be provided to the tortoises at all times” is
simply wrong. These environments are not present in nature. There is an excellent, really well done study on Gopherus burrows. In the most detailed and comprehensive study yet carried out on Desert tortoise burrows in Nevada, Bulova (2002) established accurate environmental data for more than 70 individual burrows over an extended period and recorded burrow relative humidity levels ranging from 6.1 to 44.5%, compared to surface relative humidity levels of 4.1 to 32.2%.
Not one approached 100%. Over 15+ years I have carefully measured such microclimates all over the Mediterranean - and my finding are very consistent. The claim in that paper that there are areas of "high humidity" (with the inference this is around 100%) available to tortoises "even during arid conditions" is totally false.
Don't even get me started on the "cellular dehydration" rubbish!!!
Andy