While karst is not biogenic in the same sense as, say, coral reefs or peat bogs, and carbonate dissolution can occur abiotically, formation of karst landscapes would not occur in the absence of the biosphere. Seven levels of biogeomorphic biotic-abiotic interactions are identified, from indirect impacts to landforms as extended phenotypes. Karst is generally near the biogenic end of that spectrum, featuring reciprocal interactions and mutual adjustments between biota and landforms and interrelated geomorphological and ecological processes. Karst biogeomorphology may also involve niche construction. In many cases biogeomorphic ecosystem engineering in karst is contingent, in the sense that the engineer organisms may have no, or different, biogeomorphic impacts in non-karst environments. Several examples of contingent ecosystem engineering in karst are given, including biogeomorphic effects of chinkapin oak. Abiotic geomorphic features exist on Earth, but consideration of landform types lying between the biotic-abiotic extremes would likely yield broadly similar conclusions to those about karst. However, it is also clear that we know very little about niche construction and coevolution in karst biogeomorphology, and whether karst or any specific karst features can be considered an extended (composite) phenotype is still an open question. Thus far, most work on biogeomorphology and ecosystem engineering has focused on what might be called obligate engineers—organisms whose engineering effects are at least inevitable, if not necessary to their survival. However, in some cases contingent ecosystem engineers have substantial geomorphic impacts.
Proliferation of Isoëtalean Lycophytes During the Permo-Triassic Biotic Crises: A Proxy for the State of the Terrestrial Biosphere
