biology - Where does the energy come from in the Dune ecosystem?

In Frank Herbert's Dune books, the planet Arrakis has an interesting and well thought-out alien ecology. As summarised in this answer, the sandworms eat sandplankton (which consists, at least in part, of larval sandworms), while the sandplankton eat the spice, which is produced by side-effects of the sandworm life cycle.

However, one thing has always bothered me about this: where does the energy come from? On Earth, most ecosystems are powered ultimately by sunlight, while others are powered by chemotrophy. But either way, you need a source of energy or life can't survive.

There seems to be no substantial photosynthesis on Dune, since the planet is almost all either open sand or bare rock. (Some plants are mentioned, but they are all familiar Earth species and were presumably brought there by humans.) Perhaps the sand plankton, or even the sandtrouts or the sandworms themselves, are autotrophs --- but if so, where do they get their energy from? In short, where are the primary producers of the Dune ecosystem, and what do they eat?

To the best of my knowledge, this issue isn't mentioned in the original book, but Herbert wrote a lot of sequels, and it seems that having a plausible ecology is something he cared about, so I'm wondering if he addressed this point in any of his later works.

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