Caves and other subterranean habitats fulfill the requirements of experimental model systems to address general questions in ecology and evolution. Yet, the harsh working conditions of these environments and the uniqueness of the subterranean organisms have challenged most attempts to pursuit standardized research. Two main obstacles have synergistically hampered previous attempts. First, there is ahabitat impedimentrelated to the objective difficulties of exploring subterranean habitats and our inability to access the network of fissures that represents the elective habitat for the so‐called “cave species.” Second, there is abiological impedimentillustrated by the rarity of most subterranean species and their low physiological tolerance, often limiting sample size and complicating laboratory experiments. We explore the advantages and disadvantages of four general experimental setups (in situ,quasi in situ,ex situ, andin silico) in the light of habitat and biological impediments. We also discuss the potential of indirect approaches to research. Furthermore, using bibliometric data, we provide a quantitative overview of the model organisms that scientists have exploited in the study of subterranean life. Our over‐arching goal is to promote caves as model systems where one can perform standardized scientific research. This is important not only to achieve an in‐depth understanding of the functioning of subterranean ecosystems but also to fully exploit their long‐discussed potential in addressing general scientific questions with implications beyond the boundaries of this discipline.
Orb-web spiders and their webs constitute an ideal model system in which to study behavioural flexibility and spatial cognition in invertebrates due to the easily quantifiable nature of the orb web. A large number of studies demonstrate how spiders are able to modify the geometry of their webs in response to a range of different conditions including the ability to adapt their webs to spatial constraints. However, the mechanisms behind this impressive web-building flexibility in these cognitively limited animals remain poorly explored. One possible mechanism though may be spatial learning during the spiders’ exploration of their immediate surroundings. This review discusses the importance of exploration behaviour, the reliance on simple behavioural rules, and the use of already laid threads as guidelines for web-building in orb-web spiders. The focus is on the spiders’ ability to detect and adapt their webs to space limitations and other spatial disruptions. I will also review the few published studies on how spatial information is gathered during the exploration phase and discuss the possibility of the use of ‘cognitive map’-like processes in spiders. Finally, the review provides suggestions for designing experimental studies to shed light on whether spiders gather metric information during the site exploration (cognitive map hypothesis) or rely on more simple binary information in combination with previously laid threads to build their webs (stigmergy hypothesis).
Studies on spiders in their natural habitats are necessary for determining the full range of plasticity in their web-building behaviour. Plasticity in web design is hypothesised to be important for spiders building in habitats where environmental conditions cause considerable web damage. Here we compared web characteristics of the orb spider Metellina mengei (Araneae, Tetragnathidae) in two different forest habitats differing in their wind exposure. We found a notable lack of differences in web geometry, orientation and inclination between webs built along an exposed forest edge and those built inside the forest, despite marked differences in wind speed. This suggests that M. mengei did not exhibit web-building plasticity in response to wind in the field, contrasting with the findings of laboratory studies on other species of orb spiders. Instead, differences in prey capture and wind damage trade-offs between habitats may provide an explanation for our results, indicating that different species employ different strategies to cope with environmental constraints.
Interest for subterranean biology has risen sharply in recent years due to the simplicity of the cave environment. However, most studies have focussed on morphology with few studies looking at behaviour. The cave orb spiders show some unique behavioural adaptations compared to other orb spiders, including rudimentary orb webs, off-web foraging and a complex life cycle with a surface phase. Here, we compare these behavioural adaptations in the European Meta menardi and Meta bourneti to similar behaviours in surface-dwelling orb spiders. We find that current data suggest (1) an extreme reduction in the number of frame threads, (2) evidence of capturing non-flying prey, but not necessarily evidence for off-web foraging and (3) dispersal through a surface-dwelling life stage, but with data lacking on the role of ballooning and their return to caves. We conclude that Meta spiders have potential as model organisms for studies on behavioural adaptations and flexibility.