Their earlier work involved studying rat and bat navigation in unnatural environments - a "2D" space which was really a 3D space constrained to be along a single level. This forced or even allowed the rat brains to dumb things down to a situation where they had no need or opportunity to consider things outside that unnaturally limited space and so their brains naturally optimized for that simplistic situation.
You have dumbed down their environment to the point where you have established a minimal ability that they need to master in order to function and navigate that environment.
Then when you discover that their memory encodes navigational information from this dumbed-down, unnatural environment in a regularized grid, this should be no surprise. They are minimizing the energy required to thrive in that environment.
Once you allow them to navigate in a more natural environment you should expect to see this regular grid disappear since their options for reaching a destination are no longer constrained to a single path along most of the route to their destination. Their brains will have to incorporate many more clues to guide their route selection and those clues could be time-varying as in the case where they are tracking the source of an odor as they search for food so they will need to monitor air flow and direction, odor intensity, potential obstacles between themselves and the source, alternate routes which present themselves as they move through space, threat detection from predators or bait traps, etc. In short, there so many variables that will need to be evaluated once you remove the flat plane constraints that led to their grid discovery that it should be no surprise to discover that a more complex environment uses different optimizations, some of which are encoded in a regularized notation and others, probably situational events on the path-picking decision tree, are more random.