Fundamental Science and Engineering Questions in Planetary Cave Exploration.

Nearly half a century ago, two papers postulated the likelihood of lunar lava tube caves using mathematical models. Today, armed with an array of orbiting and fly-by satellites and survey instrumentation, we have now acquired cave data across our solar system-including the identification of potential cave entrances on the Moon, Mars, and at least nine other planetary bodies. These discoveries gave rise to the study of planetary caves. To help advance this field, we leveraged the expertise of an interdisciplinary group to identify a strategy to explore caves beyond Earth. Focusing primarily on astrobiology, the cave environment, geology, robotics, instrumentation, and human exploration, our goal was to produce a framework to guide this subdiscipline through at least the next decade. To do this, we first assembled a list of 198 science and engineering questions. Then, through a series of social surveys, 114 scientists and engineers winnowed down the list to the top 53 highest priority questions. This exercise resulted in identifying emerging and crucial research areas that require robust development to ultimately support a robotic mission to a planetary cave-principally the Moon and/or Mars. With the necessary financial investment and institutional support, the research and technological development required to achieve these necessary advancements over the next decade are attainable. Subsequently, we will be positioned to robotically examine lunar caves and search for evidence of life within Martian caves; in turn, this will set the stage for human exploration and potential habitation of both the lunar and Martian subsurface.

Using near-surface temperature data to vicariously calibrate high-resolution thermal infrared imagery and estimate physical surface properties.

Thermal response of the surface to solar insolation is a function of the topography and the thermal physical characteristics of the landscape, which include bulk density, heat capacity, thermal conductivity and surface albedo and emissivity. Thermal imaging is routinely used to constrain thermal physical properties by characterizing or modeling changes in the diurnal temperature profiles. Images need to be acquired throughout the diurnal cycle - typically this is done twice during a diurnal cycle, but we suggest multiple times. Comparison of images acquired over 24 hours requires that either the data be calibrated to surface temperature, or the response of the thermal camera is linear and stable over the image acquisition period. Depending on the type and age of the thermal instrument, imagery may be self-calibrated in radiance, corrected for atmospheric effects, and pixels converted to surface temperature. We used an experimental instrumentation where the calibration should be stable, but calibration coefficients are unknown. Cases may occur where one wishes to validate the camera's calibration. We present a method to validate and calibrate the instrument and characterize the thermal physical properties for areas of interest. Finally, in situ high-temporal-resolution oblique thermal imaging can be invaluable in preparation for conducting overflight missions. We present the following:•The use of oblique thermal high temporal resolution thermal imaging over diurnal or multiday periods for the characterization of landscapes has not been widespread but poses great potential.•A method of collecting and analyzing thermal data that can be used to either determine or validate thermal camera calibration coefficients.•An approach to characterize thermophysical properties of the landscape using oblique temporally high-resolution thermal imaging, combined with in situ ground measurements.

Towards evidence-based conservation of subterranean ecosystems.

Subterranean ecosystems are among the most widespread environments on Earth, yet we still have poor knowledge of their biodiversity. To raise awareness of subterranean ecosystems, the essential services they provide, and their unique conservation challenges, 2021 and 2022 were designated International Years of Caves and Karst. As these ecosystems have traditionally been overlooked in global conservation agendas and multilateral agreements, a quantitative assessment of solution-based approaches to safeguard subterranean biota and associated habitats is timely. This assessment allows researchers and practitioners to understand the progress made and research needs in subterranean ecology and management. We conducted a systematic review of peer-reviewed and grey literature focused on subterranean ecosystems globally (terrestrial, freshwater, and saltwater systems), to quantify the available evidence-base for the effectiveness of conservation interventions. We selected 708 publications from the years 1964 to 2021 that discussed, recommended, or implemented 1,954 conservation interventions in subterranean ecosystems. We noted a steep increase in the number of studies from the 2000s while, surprisingly, the proportion of studies quantifying the impact of conservation interventions has steadily and significantly decreased in recent years. The effectiveness of 31% of conservation interventions has been tested statistically. We further highlight that 64% of the reported research occurred in the Palearctic and Nearctic biogeographic regions. Assessments of the effectiveness of conservation interventions were heavily biased towards indirect measures (monitoring and risk assessment), a limited sample of organisms (mostly arthropods and bats), and more accessible systems (terrestrial caves). Our results indicate that most conservation science in the field of subterranean biology does not apply a rigorous quantitative approach, resulting in sparse evidence for the effectiveness of interventions. This raises the important question of how to make conservation efforts more feasible to implement, cost-effective, and long-lasting. Although there is no single remedy, we propose a suite of potential solutions to focus our efforts better towards increasing statistical testing and stress the importance of standardising study reporting to facilitate meta-analytical exercises. We also provide a database summarising the available literature, which will help to build quantitative knowledge about interventions likely to yield the greatest impacts depending upon the subterranean species and habitats of interest. We view this as a starting point to shift away from the widespread tendency of recommending conservation interventions based on anecdotal and expert-based information rather than scientific evidence, without quantitatively testing their effectiveness.

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.

Two new subterranean-adapted pseudoscorpions (Pseudoscorpiones: Neobisiidae: Parobisium) from Beijing, China.

Two new troglomorphic pseudoscorpion species, Parobisium magangensis sp. n. and P. yuantongi sp. n., belonging to the family Neobisiidae, are described based on specimens collected in karst caves from Beijing, China. These are the first troglomorphic pseudoscorpions discovered from caves in northern China. Detailed diagnosis, descriptions, and illustrations are provided. We also offer future research and management recommendations for these two new pseudoscorpion species.

A new millipede, Austrotyla awishoshola n. sp., (Diplopoda, Chordeumatida, Conotylidae) from New Mexico, USA, and the importance of cave moss gardens as refugial habitats.

Austrotyla awishoshola n. sp. is described from the moss gardens of one lava tube cave in El Malpais National Monument, Cibola Co., New Mexico. Most chordeumatidans require mesic conditions, and these environments are limited to moss gardens in several cave entrances and beneath cave skylights in El Malpais. Presently, this species is known from the moss gardens of a single of cave in the monument. We suggest A. awishoshola may be a climatic relict, having become restricted to the cave environment following the end of the Pleistocene. We discuss the importance of cave moss gardens as refugial and relictual habitats. Recommendations are provided to aid in the conservation and management of A. awishoshola and these habitats.

The terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island), with descriptions of two new species.

Nine species of terrestrial isopods are reported for the Polynesian island of Rapa Nui (Easter Island) based upon museum materials and recent collections from field sampling. Most of these animals are non-native species, but two are new to science: Styloniscusmanuvaka sp. n. and Hawaiiosciarapui sp. n. Of these, the former is believed to be a Polynesian endemic as it has been recorded from Rapa Iti, Austral Islands, while the latter is identified as a Rapa Nui island endemic. Both of these new species are considered 'disturbance relicts' and appear restricted to the cave environment on Rapa Nui. A short key to all the oniscidean species presently recorded from Rapa Nui is provided. We also offered conservation and management recommendations for the two new isopod species.

Collembola of Rapa Nui (Easter Island) with descriptions of five endemic cave-restricted species.

Eight species of Collembola are reported from recent collections made in caves on the Polynesian island of Rapa Nui (Easter Island). Five of these species are new to science and apparently endemic to the island: Coecobrya aitorererere n. sp., Cyphoderus manuneru n. sp., Entomobrya manuhoko n. sp., Pseudosinella hahoteana n. sp. and Seira manukio n. sp. The Hawaiian species Lepidocyrtus olena Christiansen & Bellinger and the cosmopolitan species Folsomia candida Willem also were collected from one or more caves. Coecobrya kennethi Jordana & Baquero, recently described from Rapa Nui and identified as endemic, was collected in sympatric association with C. aitorererere n.sp. With the exception of F. candida, all species are endemic to Rapa Nui or greater Polynesia and appear to be restricted to the cave environment on Rapa Nui. A key is provided to separate Collembola species reported from Rapa Nui. We provide recommendations to aid in the conservation and management of these new Collembola, as well as the other presumed cave-restricted arthropods.

Genus Cyptophania Banks (Psocodea: 'Psocoptera': Lepidopsocidae): unique features, augmented description of the generotype, and descriptions of three new species.

We define the genus Cyptophania with characters that clearly separate it from other genera of the Family Lepidopsocidae in which wing reduction has occurred. We redescribe the generotype, C. hirsuta Banks (Hawaii, presumably introduced), and describe three new species, C. australica n.sp. (Queensland, Australia), C. costalis n.sp. (Gulf of Mexico and Caribbean), and C. pakaratii n.sp. (Rapa Nui = Easter Island, probably endemic). The latter species is described from both sexes and presents the first males known for the genus. One female of this species presents a large spermatophore protruding from the genital chamber, thus indicating the mode of sperm transfer in sexual members of this genus. A key to the known species is included. All of the species of Cyptophania are highly neotenic, but differences in the level of neoteny are noted among the species studied. We question the synonymy of the genus Ptenocorium Enderlein with Cyptophania on the basis of several characters illustrated in the original description of Ptenocorium. We note similarities of Cyptophania to the entirely macropterous genus Lepidopsocus Enderlein and suggest a possible close relationship between the two genera.