Abstract
Compliant pads based on a specific foam-like or fibrillar internal structure of material are a widespread type of adhesive structures made of cuticle. Such pads usually have a smooth surface at a macroscale and are supplemented by lipid-bearing secretions. Besides, in whip-spiders there is the special case of a soft pad with spatula-like microstructures directly connected with the surface. Soft pads usually occur on the pretarsus as so-called arolia, which serve dynamic attachment during locomotion and static attachment during resting. Arachnids have arolia of different shape, which influences their contact mechanics. There are cushion- or bubble-like pads, disc- or mushroom-like ones, and arolia with a stiff trunk and a broad soft lip (transverse-lip arolium). The latter type has evolved three times independently among arachnids, with a strikingly similar structure. The specific internal structure of soft adhesive pads allows for different detachment mechanisms, like invagination, folding and lateral peeling. In addition to the literature review, this chapter includes the first ultrastructural and functional original data on the arolia of pseudoscorpions, harvestman nymphs, and some mites.
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7.1 Electronic Supplementary Material
Video 7.1
Nymph of the scorpion Liocheles australasiae (Hemiscorpiidae) walking upside down on a glass coverslip. Playback with real speed (AVI 1353 kb)
Video 7.2
Detail of a leg tip (lateral view) of a L. australasiae nymph walking on a glass coverslip, showing the detachment of the arolium by invagination. Recorded with 50 fps and playback with 30 fps (AVI 1902 kb)
Video 7.3
Detail of a leg tip (lateral view) of a L. australasiae nymph walking on a glass coverslip, showing the detachment of the arolium by invagination. Recorded with 50 fps and playback with 15 fps (M2V 8573 kb)
Video 7.4
Detail of a leg tip (view from below) of a L. australasiae nymph walking on a glass coverslip, showing the detachment of the arolium by invagination. Recorded with 500 fps and playback with 30 fps (AVI 834 kb)
Video 7.5
Detail of two leg tips (view from below) of a L. australasiae nymph walking on a glass coverslip, showing the detachment of the arolium by invagination (transmission illumination and DIC optic). Note internal features like the retractor tendon and hemolymph cells. Recorded with 500 fps and playback with 30 fps (AVI 13655 kb)
Video 7.6
RICM-HSV of the arolium of a L. australasiae nymph, showing dry mode of adhesion (no traces left behind after detachment). Recorded with 1000 fps and playback with 30 fps (AVI 29525 kb)
Video 7.7
RICM-HSV of the arolium of a L. australasiae nymph, showing wet mode of adhesion (droplets of a fluid secretion left behind after detachment). Note that the secretion keeps on shrinking to droplets on the hydrophilic glass, indicating their lipid-like nature. Also note the dirt particle, which does not stick to the arolium after detachment, but remains on the glass slide. Hence, the high amount of fluids observed here, may be part of a rinsing mechanism of the arolium. Recorded with 1000 fps and playback with 30 fps (AVI 32975 kb)
Video 7.8
Tick walking upside down on a glass slide. The tick was trapped between a common microscopy slide and the table to reduce its mobility. Nonetheless it was capable to push itself out of the glass slide, showing an enormous strength of the animal and high friction of its arolia. Note the inflation and folding of the arolia during attachment and detachment. Playback with real speed (AVI 803 kb)
Video 7.9
Detail of a leg tip (view from above) of the whip-spider Charon cf. grayi (Charontidae) walking on a glass slide, showing the detachment of the arolium and movement of claws. Note the footprint left behind, indicating the deposition of fluid secretions. Recorded with 500 fps and playback with 30 fps (AVI 3463 kb)
Video 7.10
Detail of a leg tip (frontal view) of the whip-spider Charon cf. grayi (Charontidae) walking on a glass slide, showing the detachment of the arolium. Recorded with 3000 fps and playback with 30 fps (M2V 13515 kb)
Video 7.11
RICM-HSV of an arolium of the whip-spider Charinus cubensis (Charinidae), showing the contact behavior during detachment (dry mode of adhesion). Note that single spatulae jump independently out of contact, thus there is an uneven peel-off (stepwise crack propagation). Recorded with 6250 fps and playback with 15 fps (AVI 10102 kb)
Video 7.12
RICM-HSV of an arolium of the whip-spider C. cf. grayi, showing the contact behavior during detachment (wet mode of adhesion). Recorded with 3000 fps and playback with 15 fps (AVI 5697 kb)
Video 7.13
RICM-HSV of an arolium of the whip-spider C. cf. grayi, showing the contact behavior during detachment (partially wet mode of adhesion). Note that secretions occur only in single areas and do not spread over the pad. Recorded with 8000 fps and playback with 15 fps (AVI 10710 kb)
Video 7.14
RICM-HSV of an arolium of the whip-spider Sarax brachydactylus (Charinidae), showing the contact behavior during detachment (partially wet mode of adhesion). Note that a part of the secretions appears highly viscous, keeping the imprints of spatulae. Recorded with 8000 fps and playback with 25 fps (AVI 28453 kb)
Video 7.15
RICM-HSV of an arolium of the pseudoscorpion Chthonius sp. (Chthoniidae), showing detachment. Note the bi-lateral peel-off. Recorded with 5000 fps and playback with 30 fps (AVI 3525 kb)
Video 7.16
RICM-HSV of an arolium of the pseudoscorpion Neobisium sp. (Neobisidae), showing detachment. Note the bi-lateral peel-off. Recorded with 500 fps and playback with 30 fps (AVI 5441 kb)
Video 7.17
RICM-HSV of an arolium of the pseudoscorpion Neobisium sp., showing sliding of the arolium along the glass surface. Note that lipidoid fluids are deposited by the pointed tarsal setae. Recorded with 500 fps and playback with 30 fps (AVI 33222 kb)
Video 7.18
Tip of the walking leg of the predatory mite Anystis sp. (Anystidae), showing movements of the pretarsus. The foot exhibits both a pair of tenent setae and an arolium, and the high flexibility of the distal tarsus permits a stepwise contact of setae, arolium and claws. Recorded with 1000 fps and playback with 30 fps (AVI 35145 kb)
Video 7.19
RICM-HSV of the foot of Anystis sp. showing stepwise detachment of arolium and setae. Note the fluids left behind both by arolium and tenent setae. Recorded with 5400 fps and playback with 15 fps (AVI 3878 kb)
Video 7.20
RICM-HSV of the arolium of Anystis sp. showing its detachment. Note the bi-lateral peel-off and the lack of fluid remains. Recorded with 30000 fps and playback with 15 fps (AVI 406 kb)
Video 7.21
RICM-HSV of the foot of Anystis sp. showing detachment and attachment. Note that in this case only the tenent setae are brought in contact. Recorded with 500 fps and playback with 30 fps (AVI 10232 kb)
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Wolff, J.O., Gorb, S.N. (2016). Soft Adhesive Pads. In: Attachment Structures and Adhesive Secretions in Arachnids. Biologically-Inspired Systems, vol 7. Springer, Cham. https://doi.org/10.1007/978-3-319-45713-0_7
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