Passive water collection with the integument: mechanisms and their biomimetic potential.

Several mechanisms of water acquisition have evolved in animals living in arid habitats to cope with limited water supply. They enable access to water sources such as rain, dew, thermally facilitated condensation on the skin, fog, or moisture from a damp substrate. This Review describes how a significant number of animals - in excess of 39 species from 24 genera - have acquired the ability to passively collect water with their integument. This ability results from chemical and structural properties of the integument, which, in each species, facilitate one or more of six basic mechanisms: increased surface wettability, increased spreading area, transport of water over relatively large distances, accumulation and storage of collected water, condensation, and utilization of gravity. Details are described for each basic mechanism. The potential for bio-inspired improvement of technical applications has been demonstrated in many cases, in particular for several wetting phenomena, fog collection and passive, directional transport of liquids. Also considered here are potential applications in the fields of water supply, lubrication, heat exchangers, microfluidics and hygiene products. These present opportunities for innovations, not only in product functionality, but also for fabrication processes, where resources and environmental impact can be reduced.

Cutaneous water collection by a moisture-harvesting lizard, the thorny devil (Moloch horridus).

Moisture-harvesting lizards, such as the Australian thorny devil, Moloch horridus, have the remarkable ability to inhabit arid regions. Special skin structures, comprising a micro-structured surface with capillary channels in between imbricate overlapping scales, enable the lizard to collect water by capillarity and transport it to the mouth for ingestion. The ecological role of this mechanism is the acquisition of water from various possible sources such as rainfall, puddles, dew, condensation on the skin, or absorption from moist sand, and we evaluate here the potential of these various sources for water uptake by M. horridus The water volume required to fill the skin capillary system is 3.19% of body mass. Thorny devils standing in water can fill their capillary system and then drink from this water, at approximately 0.7 µl per jaw movement. Thorny devils standing on nearly saturated moist sand could only fill the capillary channels to 59% of their capacity, and did not drink. However, placing moist sand on skin replicas showed that the capillary channels could be filled from moist sand when assisted by gravity, suggesting that their field behaviour of shovelling moist sand onto the dorsal skin might fill the capillary channels and enable drinking. Condensation facilitated by thermal disequilibrium between a cool thorny devil and warm moist air provided skin capillary filling to approximately 0.22% of body weight, which was insufficient for drinking. Our results suggest that rain and moist sand seem to be ecologically likely water sources for M. horridus on a regular basis.

Phylogenetic and morphological influence on habitat choice in moisture-harvesting horned lizards (Phrynosoma spp.).

In previous studies, the superhydrophilic skin of moisture-harvesting lizards has been linked to the morphological traits of the lizards' integument, that is, the occurrence of honeycomb-shaped microstructures. Interestingly, these structures can also cover the skin of lizards inhabiting wet habitats. We therefore tested the influence of the microstructures' main features on the habitat choice and wettability in the genus Phrynosoma. The genus Phrynosoma comprises moisture-harvesting species as well as nonspecialists. Lizards of this genus inhabit large areas of North America with diverse climatic conditions. Remarkably, the differences in the manifestation of microstructures are just as versatile as their surroundings. The phylogeny of the lizards as well as the depth of their ventral microstructures, though independent of each other, correlated with the precipitation in their respective habitat. All other morphological traits, as well as the skin's wettability itself, could not predict the habitat of Phrynosoma species. Hence, it is unlikely that the microstructure influences the wettability, at least directly. Hence, we presume an indirect influence for the following reasons: (a) As the ventral side cannot get wet by rain, but the belly could easily interact with a wet surface, the microstructure might facilitate water absorption from wet soil following precipitation. (b) We found the number of dorsal microstructures to be linked to the occurrence of silt in the habitat. In our study, we observed scales being heavily contaminated, most likely with a mixture of dead skin (after shedding) and silt. As many lizards burrow themselves or even shovel sand onto their backs, deploying the substrate might be a mechanism to increase the skin's wettability.

Adsorption and movement of water by skin of the Australian thorny devil (Agamidae: Moloch horridus).

Moisture-harvesting lizards, such as the Australian thorny devil Moloch horridus, have remarkable adaptations for inhabiting arid regions. Their microstructured skin surface, with channels in between overlapping scales, enables them to collect water by capillarity and passively transport it to the mouth for ingestion. We characterized this capillary water transport for live thorny devils using high-speed video analyses. Comparison with preserved specimens showed that live lizards are required for detailed studies of skin water transport. For thorny devils, there was no directionality in cutaneous water transport (unlike Phrynosoma) as 7 µl water droplets applied to the skin were transported radially over more than 9.2 mm. We calculated the total capillary volume as 5.76 µl cm-2 (dorsal) and 4.45 µl cm-2 (ventral), which is reduced to 50% filling by the time transportation ceases. Using micro-computed tomography and scanning electron microscopy of shed skin to investigate capillary morphology, we found that the channels are hierarchically structured as a large channel between the scales that is sub-divided by protrusions into smaller sub-capillaries. The large channel quickly absorbs water whereas the sub-capillary structure extends the transport distance by about 39% and potentially reduces the water volume required for drinking. An adapted dynamics function, which closely reflects the channel morphology, includes that ecological role.

Directional, passive liquid transport: the Texas horned lizard as a model for a biomimetic 'liquid diode'.

Moisture-harvesting lizards such as the Texas horned lizard (Iguanidae: Phrynosoma cornutum) live in arid regions. Special skin adaptations enable them to access water sources such as moist sand and dew: their skin is capable of collecting and transporting water directionally by means of a capillary system between the scales. This fluid transport is passive, i.e. requires no external energy, and directs water preferentially towards the lizard's snout. We show that this phenomenon is based on geometric principles, namely on a periodic pattern of interconnected half-open capillary channels that narrow and widen. Following a biomimetic approach, we used these principles to develop a technical prototype design. Building upon the Young-Laplace equation, we derived a theoretical model for the local behaviour of the liquid in such capillaries. We present a global model for the penetration velocity validated by experimental data. Artificial surfaces designed in accordance with this model prevent liquid flow in one direction while sustaining it in the other. Such passive directional liquid transport could lead to process improvements and reduction of resources in many technical applications.

Moisture harvesting and water transport through specialized micro-structures on the integument of lizards.

Several lizard species that live in arid areas have developed special abilities to collect water with their bodies' surfaces and to ingest the so collected moisture. This is called rain- or moisture-harvesting. The water can originate from air humidity, fog, dew, rain or even from humid soil. The integument (i.e., the skin plus skin derivatives such as scales) has developed features so that the water spreads and is soaked into a capillary system in between the reptiles' scales. Within this capillary system the water is transported to the mouth where it is ingested. We have investigated three different lizard species which have developed the ability for moisture harvesting independently, viz. the Australian thorny devil (Moloch horridus), the Arabian toadhead agama (Phrynocephalus arabicus) and the Texas horned lizard (Phrynosoma cornutum). All three lizards have a honeycomb like micro ornamentation on the outer surface of the scales and a complex capillary system in between the scales. By investigation of individual scales and by producing and characterising polymer replicas of the reptiles' integuments, we found that the honeycomb like structures render the surface superhydrophilic, most likely by holding a water film physically stable. Furthermore, the condensation of air humidity is improved on this surface by about 100% in comparison to unstructured surfaces. This allows the animals to collect moisture with their entire body surface. The collected water is transported into the capillary system. For Phrynosoma cornutum we found the interesting effect that, in contrast to the other two investigated species, the water flow in the capillary system is not uniform but directed to the mouth. Taken together we found that the micro ornamentation yields a superhydrophilic surface, and the semi-tubular capillaries allow for an efficient passive - and for Phrynosoma directed - transport of water.