Fundamental research questions in subterranean biology.

Five decades ago, a landmark paper in Science titled The Cave Environment heralded caves as ideal natural experimental laboratories in which to develop and address general questions in geology, ecology, biogeography, and evolutionary biology. Although the 'caves as laboratory' paradigm has since been advocated by subterranean biologists, there are few examples of studies that successfully translated their results into general principles. The contemporary era of big data, modelling tools, and revolutionary advances in genetics and (meta)genomics provides an opportunity to revisit unresolved questions and challenges, as well as examine promising new avenues of research in subterranean biology. Accordingly, we have developed a roadmap to guide future research endeavours in subterranean biology by adapting a well-established methodology of 'horizon scanning' to identify the highest priority research questions across six subject areas. Based on the expert opinion of 30 scientists from around the globe with complementary expertise and of different academic ages, we assembled an initial list of 258 fundamental questions concentrating on macroecology and microbial ecology, adaptation, evolution, and conservation. Subsequently, through online surveys, 130 subterranean biologists with various backgrounds assisted us in reducing our list to 50 top-priority questions. These research questions are broad in scope and ready to be addressed in the next decade. We believe this exercise will stimulate research towards a deeper understanding of subterranean biology and foster hypothesis-driven studies likely to resonate broadly from the traditional boundaries of this field.

Shifting stream planform state decreases stream productivity yet increases riparian animal production.

In the Colorado Front Range (USA), disturbance history dictates stream planform. Undisturbed, old-growth streams have multiple channels and large amounts of wood and depositional habitat. Disturbed streams (wildfires and logging < 200 years ago) are single-channeled with mostly erosional habitat. We tested how these opposing stream states influenced organic matter, benthic macroinvertebrate secondary production, emerging aquatic insect flux, and riparian spider biomass. Organic matter and macroinvertebrate production did not differ among sites per unit area (m-2), but values were 2 ×-21 × higher in undisturbed reaches per unit of stream valley (m-1 valley) because total stream area was higher in undisturbed reaches. Insect emergence was similar among streams at the per unit area and per unit of stream valley. However, rescaling insect emergence to per meter of stream bank showed that the emerging insect biomass reaching the stream bank was lower in undisturbed sites because multi-channel reaches had 3 × more stream bank than single-channel reaches. Riparian spider biomass followed the same pattern as emerging aquatic insects, and we attribute this to bottom-up limitation caused by the multi-channeled undisturbed sites diluting prey quantity (emerging insects) reaching the stream bank (riparian spider habitat). These results show that historic landscape disturbances continue to influence stream and riparian communities in the Colorado Front Range. However, these legacy effects are only weakly influencing habitat-specific function and instead are primarily influencing stream-riparian community productivity by dictating both stream planform (total stream area, total stream bank length) and the proportional distribution of specific habitat types (pools vs riffles).

Quantitative food web analysis supports the energy-limitation hypothesis in cave stream ecosystems.

Energy limitation has long been the primary assumption underlying conceptual models of evolutionary and ecological processes in cave ecosystems. However, the prediction that cave communities are actually energy-limited in the sense that constituent populations are consuming all or most of their resource supply is untested. We assessed the energy-limitation hypothesis in three cave streams in northeastern Alabama (USA) by combining measurements of animal production, demand, and resource supplies (detritus, primarily decomposing wood particles). Comparisons of animal consumption and detritus supply rates in each cave showed that all, or nearly all, available detritus was required to support macroinvertebrate production. Furthermore, only a small amount of macroinvertebrate prey production remained to support other predatory taxa (i.e., cave fish and salamanders) after accounting for crayfish consumption. Placing the energy demands of a cave community within the context of resource supply rates provided quantitative support for the energy-limitation hypothesis, confirming the mechanism (limited energy surpluses) that likely influences the evolutionary processes and population dynamics that shape cave communities. Detritus-based surface ecosystems often have large detrital surpluses. Thus, cave ecosystems, which show minimal surpluses, occupy the extreme oligotrophic end of the spectrum of detritus-based food webs.