CHAPTER 3: FINDING REFUGE
Hier fehlt eine Einführung
CHAPTER 3: FINDING REFUGE
Hier fehlt eine Einführung
DAD-010 (05)
The Ground Dweller
(Reductio alarum)
Description
Recurring heat waves and prolonged drought are turning the soil’s surface into an increasingly inhospitable habitat. Cooler layers of soil and sheltered cavities are becoming increasingly important. The Ground Dweller explores this hidden landscape with a body whose strength is focused on digging. Her wings have regressed to short, unevenly folded remnants; her front legs have broadened into serrated tools. They dislodge soil particles, widen crevices, and push her body through narrow passages. Within this evolutionary lineage, the subterranean lifestyle favors a powerful, compact body. The wasp gains access to spaces that were previously beyond her reach.
Scientific Context
The loss of wings may be accompanied by a shift in the allocation of physical resources: In stoneflies, smaller wings and reduced flight musculature have been linked to a larger ovary mass.¹ Studies of beetles associate the inability to fly with greater geographic differentiation and speciation.² In Hawaii, male crickets lost their ability to sing due to altered wing structures under the selective pressure of an acoustically oriented parasitoid.³ What becomes dispensable or dangerous may disappear—with consequences that extend beyond the individual function. The ground-dweller’s digging tools open up shelters, but its atrophied wings block escape routes. When water or pollutants penetrate the tunnels, the lifesaving habitat lies beyond what her body can still reach.
¹ G. A. McCulloch et al., ‘Dispersal-fecundity trade-offs in wild insect populations’, Journal of Evolutionary Biology (2025). https://pubmed.ncbi.nlm.nih.gov/39663214/
² ‘Genotyping-by-sequencing supports a genetic basis for wing reduction in an alpine New Zealand stonefly’, Scientific Reports (2018). https://doi.org/10.1038/s41598-018-34123-1
³ H. Ikeda et al., ‘Loss of flight promotes beetle diversification’, Nature Communications, 3 (2012), 648. https://doi.org/10.1038/ncomms1659
DAD-011 (10)
The Flood-Clingers
(Tarsi adhaesivi)
Description
The legs of the Flood-Clingers end in strong, curved claws, between which lie enlarged, soft adhesive pads. The different contact surfaces give the delicate legs a striking tool-like functionality: the claws can grip uneven surfaces, while the pads provide a firm hold on suitable smooth surfaces. Recurring flooding and sudden inflows of water constitute the environmental pressure driving this adaptation. On roots, rough bark, or exposed plant fibers, the wasp could maintain its position longer and reach a higher refuge. What matters most is what it clings to. Even a strong grip lasts only as long as the substrate itself holds.
Scientific Context
In ants, the claws and the movable adhesive pad between them work in tandem: Depending on the surface roughness, different parts of the foot are used.¹ Water can significantly disrupt this interaction. Studies of a tropical tree ant show that adhesive performance on wet surfaces depends on the properties of the substrate.² On the wet edges of certain pitcher plants, water films interrupt contact between the adhesive pads; whether the claws grip, in turn, depends on the surface structure.³ For flood-clinging ants, the potential advantage therefore lies primarily in mechanical grip on suitable rough structures. Their feet do not seal off either the body or the nest. A worker ant could withstand the water while the brood chambers beneath her fill with water. A gap opens up between the survival of the individual animal and the survival of its colony—a gap that even enlarged claws cannot bridge.
¹ ‘Walking on smooth or rough ground: passive control of pretarsal attachment in ants’, Journal of Experimental Biology (2007). https://pubmed.ncbi.nlm.nih.gov/18060411/
² ‘Adhesion and running speed of a tropical arboreal ant (Cephalotes atratus) on wet substrates’, Royal Society Open Science (2018). https://pmc.ncbi.nlm.nih.gov/articles/PMC6281928/
³ ‘Insect aquaplaning: Nepenthes pitcher plants capture prey with the peristome, a fully wettable water-lubricated anisotropic surface’, Proceedings of the National Academy of Sciences (2004). https://pubmed.ncbi.nlm.nih.gov/15383667/
DAD-012 (11)
The Crevice Dweller
(CORPUS DEPRESSUM)
Description
Her body fills the last remaining spaces. Her head, thorax, and abdomen are distinctly flattened; her legs extend outward to the sides, and her wings lie close to her back. Where a curved body once met a boundary, a narrow crevice now suffices. The environmental pressure driving this adaptation stems from the loss of sheltered cavities due to sealed soils, airtight buildings, and the removal of old woodlands. The crevice dweller gains access to the remaining spaces in an increasingly sealed-off landscape: under loose bark, between building components, behind chipped masonry. Her form becomes a precise response to a world that leaves her with less and less space.
Scientific Context
Cockroaches can temporarily compress their bodies significantly and continue moving even in extremely confined spaces. This is made possible by the interaction of a deformable exoskeleton and altered leg positions.¹ Further experiments demonstrate how body contour, lateral rolling, and the positioning of the limbs facilitate passage between obstacles. ² ³ The permanent flattening of this crevice-dweller takes this idea further—until it reaches a problem that no amount of dexterous movement can solve: A crevice that offers a worker refuge is not yet a nest. Space must still remain for honeycombs, brood, and the queen. Thus, while the individual body might fit ever better into the remaining gaps, the colony’s shared living space disappears.
¹ K. Jayaram and R. J. Full, “Cockroaches traverse crevices, crawl rapidly in confined spaces, and inspire a soft, legged robot,” Proceedings of the National Academy of Sciences, 113, no. 8 (2016), pp. E950–E957. https://doi.org/10.1073/pnas.1514591113
² C. Li, A. O. Pullin, D. W. Haldane, H. K. Lam, R. S. Fearing, and R. J. Full, “Terradynamically streamlined shapes in animals and robots enhance traversability through densely cluttered terrain,” Bioinspiration & Biomimetics, 10, No. 4 (2015), Article 046003. https://doi.org/10.1088/1748-3190/10/4/046003
³ Y. Wang, R. Othayoth, and C. Li, “Cockroaches adjust body and appendages to traverse cluttered large obstacles,” Journal of Experimental Biology, 225, No. 10 (2022), Article jeb243605. https://doi.org/10.1242/jeb.243605
DAD-006
The Dust-Grooming
(Pedes pectinati)
Description
The dust-grooming carries a delicate cleaning tool on its front leg. An enlarged comb unfolds along its inner side: closely spaced teeth, sturdy bristles, and fine hairs form a tiered structure through which she pulls her antennae. The environmental pressure driving this evolution is constant exposure to stirred-up soil dust, soot, and abrasion. New particles are constantly depositing themselves on the sensitive sensory surfaces. In this evolved form of the German wasp, the unassuming cleaning device has developed into a striking bodily tool. With every stroke through the comb, she clears her sensory organs once again. The precision of this structure is astonishing: a single leg is responsible for maintaining the very senses the wasp uses to find food, fellow wasps, and her way through the world.
Scientific Context
In the ant Camponotus rufifemur, the bristles, comb, and brush on the front leg already function as a multi-stage particle filter: Different structures remove contaminants of varying sizes from the antennae.¹ Experiments on cockroaches show what happens without this grooming: Deposits cover the pores of the sensory hairs, and the antennae respond less sensitively to odors.² In houseflies, even brief exposure to airborne particles impaired their perception of food and mating odors.³ The dust-combing ant takes this existing cleaning principle to the level of a physical specialization. Its enlarged apparatus inherently limits the mobility of the front leg; the constant cleaning takes time that is otherwise needed for foraging and care. The body can still keep its senses clear. The environment, which repeatedly challenges it, remains.
¹ A. Hackmann et al., ‘Functional morphology and efficiency of the antenna cleaner in Camponotus rufifemur ants’, Royal Society Open Science, 2 (2015), 150129. https://doi.org/10.1098/rsos.150129
² K. Böröczky et al., ‘Insects groom their antennae to enhance olfactory acuity’, Proceedings of the National Academy of Sciences (2013). https://pmc.ncbi.nlm.nih.gov/articles/PMC3587213/
³ ‘Short-term particulate matter contamination severely compromises insect antennal olfactory perception’, Nature Communications, 14 (2023), 4112. https://doi.org/10.1038/s41467-023-39469-3
DAD-008
The Dew Harvesters
(Setae aquiferae)
Description
An irregular fringe of elongated bristles rises above the head, thorax, and abdomen of the dew harvesters. Individual water droplets hang between these fine structures, focusing the light and causing the dark body surface to glow in places. Prolonged dry spells and the disappearance of reliable water sources constitute the environmental pressures faced by this species. Their bristles serve as collection surfaces during the brief humid hours of the day: when the surface cools below the dew point, water can condense. Larger droplets merge between adjacent bristles; accessible accumulations are scraped off with the legs and guided to the mouthparts. The body taps into a fleeting water source, drop by drop.
Scientific Context
Certain beetles in the Namib Desert, located on the southwestern coast of Africa, obtain drinking water from fog. They orient their bodies toward the moist wind and assume a posture that allows the collected droplets to flow toward their mouths.¹ Comparative experiments show how crucial the conveyance of water is, in addition to its collection; flow simulations further illustrate the influence of body posture on yield.² ³ Fog collection captures droplets already in the air, whereas dew only forms on a sufficiently cooled surface. Both methods tie access to water to a close interplay between the body and the weather. The dew-combing beetle visibly bears this dependence on its back. Its bristles can hold water, but they cannot create the conditions for its formation. If the nights remain too dry or the body does not cool sufficiently, the collection apparatus remains empty.
¹ T. Nørgaard und M. Dacke, ‘Fog-basking behaviour and water collection efficiency in Namib Desert Darkling beetles’, Frontiers in Zoology, 7 (2010), 23. https://doi.org/10.1186/1742-9994-7-23
² T. Nørgaard, M. Ebner und M. Dacke, ‘Animal or Plant: Which Is the Better Fog Water Collector?’, PLOS ONE, 7 (2012), e34603. https://doi.org/10.1371/journal.pone.0034603
³ U. Chakrabarti et al., ‘Importance of Body Stance in Fog Droplet Collection by the Namib Desert Beetle’, Biomimetics, 4 (2019), 59. https://doi.org/10.3390/biomimetics4030059
DAD-009
The Stilt-Walker
(Pedes elongati)
Description
The legs of the Stilt-Walker extend far beyond the familiar outline of its body. Elongated limbs form narrow, jointed arches; the small claws remain intact at their ends. When the body is at rest, these struts lie side by side, folded unevenly and stretched out. Their potential benefit becomes apparent when walking: a carefully coordinated leg posture lifts the thorax and abdomen further off the ground. Highly heated, unshaded ground surfaces provide the environmental pressure that shapes this form. An additional gap is created between the body and the surface, which can already provide access to a different thermal environment near the ground. The wasp carries the possibility of a small refuge within the length of its limbs.
Scientific Context
Heat-loving desert ants of the genera Cataglyphis and Ocymyrmex have particularly long legs that can lift their bodies into cooler layers of air above the hot ground.¹ Studies of their locomotion also show how closely leg length, stride sequence, and ground contact interact; particularly high speeds can be achieved even with comparatively shorter legs and higher stride frequencies.² ³ In the case of the “stilt-walker,” this elongation also alters the relationship to its own nest. When passing through narrow passages, turning between the honeycombs, and working on the cell edges, the legs must be tucked in and repositioned. If larger distances between honeycombs and wider entrances become necessary, the colony’s space requirements increase. If the nest architecture remains unchanged, certain construction and maintenance tasks could increasingly fall to the shorter-legged workers. The long-legged worker gains distance from the hot ground but at the same time risks losing access to parts of her own nest.
¹ S. Sommer und R. Wehner, ‘Leg allometry in ants: Extreme long-leggedness in thermophilic species’, Arthropod Structure & Development (2012). https://pubmed.ncbi.nlm.nih.gov/21992805/
² ‘Walking and running in the desert ant Cataglyphis fortis’, Journal of Comparative Physiology A (2015). https://doi.org/10.1007/s00359-015-0999-2
³ ‘High-speed locomotion in the Saharan silver ant, Cataglyphis bombycina’, Journal of Experimental Biology, 222 (2019), jeb198705. https://pubmed.ncbi.nlm.nih.gov/31619540/
DAD-012
The Breaker
(Mandibulae robustae)
Description
The head becomes the center of power. Two short, massive mandibles with widened bases and powerful inner teeth define its form. Behind them, the enlarged head capsule creates space for stronger closing muscles. The environmental pressure driving this design stems from the scarcity of easily removable wood fibers: In a tidied-up, sealed landscape, weathering wood surfaces disappear, while harder materials remain. The breaker taps into this stubborn residue. Her mouthparts grip, notch, and extract fibers from surfaces where other workers fail. Nest-building begins with a forceful intervention into the material; resistance is transformed back into a habitable space.
Scientific Context
The mouthparts of Vespula germanica already combine material collection, prey processing, and brood care.¹ Studies of other invertebrates demonstrate just how effective such tools can become: structures rich in zinc and manganese enable hard, wear-resistant cutting edges; in addition to size, the texture of their contact surfaces is also crucial. ² In leaf-cutting ants, wear on the mandibles even alters the division of labor: ants with worn cutting edges cut more slowly and more frequently take on transport duties. ³ For the nest builders, this spells a precarious future. A colony whose building material can be harvested by only a few specialized workers depends on the condition of their tools. With each cutting edge that becomes blunter, the ability to expand the nest or repair damage could diminish.
¹ B. Baranek, K. Kuba, J. Bauder, and H. Krenn, “Mouthpart dimorphism in male and female wasps of Vespula vulgaris and Vespula germanica (Vespidae, Hymenoptera),” Deutsche Entomologische Zeitschrift, 65, No. 1 (2018), pp. 65–74. https://doi.org/10.3897/dez.65.23593
² R. M. S. Schofield et al., “The homogeneous alternative to biomineralization: Zn- and Mn-rich materials enable sharp organismal ‘tools’ that reduce force requirements,” Scientific Reports, 11 (2021), Article 17481. https://doi.org/10.1038/s41598-021-91795-y
³ R. M. S. Schofield, K. D. Emmett, J. C. Niedbala, and M. H. Nesson, “Leaf-cutter ants with worn mandibles cut half as fast, spend twice the energy, and tend to carry instead of cut,” Behavioral Ecology and Sociobiology, 65 (2011), pp. 969–982. https://doi.org/10.1007/s00265-010-1098-6
DAD-013
The Long-Tongued Wasp
(Partes oris elongatae)
Description
Below the head, a slender, widely extending mouthpart unfolds. The elongated portions of the lower lip and maxillae together form a movable apparatus for taking in liquids; the lateral mandibles remain intact. The environmental pressure driving this design is a changing foraging landscape: Recurring dry spells and intensive land management are causing a decline in easily accessible flower resources. Where deep-calyx plants remain as reliable sources of nectar, an extended reach provides an advantage. The Long-Reaching Forager reaches food behind narrow flower openings without having to force its head inside. Her body opens up a new spatial dimension of foraging: depth becomes a storage space.
Scientific Context
The German wasp already possesses an apparatus for fluid intake formed by several mouthparts, whose bristles and internal structures filter food particles.¹ South African Prosoeca flies demonstrate just how far this extension can go: their extremely long mouthparts allow them to access nectar in deep flower tubes. ² However, the relationship between body structure and food availability can also be reversed. A study spanning four decades documented shorter tongues in two alpine bumblebee species; the researchers linked this development to the decline in flower availability during warmer summers and a more diverse foraging behavior. ³ For long-range feeders, the danger therefore lies in the permanence of their specialization. If the plants whose nectar they can now reach disappear, their elaborately elongated anatomy remains. A body may still be tailored to a food source that has long since ceased to exist.
¹ K. Kuba et al., “Filtration of Nutritional Fluids in the German Wasp Vespula germanica (Vespidae, Hymenoptera),” Insects, 13, No. 2 (2022), Article 185. https://doi.org/10.3390/insects13020185
² F. Karolyi, N. U. Szucsich, J. F. Colville, and H. W. Krenn, “Adaptations for nectar-feeding in the mouthparts of long-proboscid flies (Nemestrinidae: Prosoeca),” Biological Journal of the Linnean Society, 107, No. 2 (2012), pp. 414–424. https://doi.org/10.1111/j.1095-8312.2012.01945.x
³ N. E. Miller-Struttmann et al., “Functional mismatch in a bumblebee pollination mutualism under climate change,” Science, 349, No. 6255 (2015), pp. 1541–1544. https://doi.org/10.1126/science.aab0868
DAD-014
Vibration Reader
(SENSILLA TARSALIA AUCTA)
Description
The legs become instruments of perception that function without light. Widened foot segments establish contact with the ground; enlarged mechanosensory organs are embedded in the reinforced rails. The Vibration Reader pauses, spreads its legs across the surface, and detects the faintest vibrations traveling through wood, plant stems, or nest walls. The environmental pressure in this design is sustained daytime heat, which shifts activity into twilight and into sheltered, dark cavities. Where visual orientation reaches its limits, the movement of the material takes on greater significance. A hidden rustling, a touch on the nest shell, the footsteps of another worker: the ground becomes a conduit for messages. The world extends beyond what is visible.
Scientific Context
Some parasitoid wasps locate hidden hosts using substrate vibrations they generate themselves. Subgenual organs inside the tibiae are involved in this perception; conspicuous external hair fields alone do not explain this ability.¹ Experiments with Pimpla turionellae also show that ambient temperature influences the activity and success of this mechanosensory search. ² However, sensitive perception remains dependent on distinguishable signals: In the green stink bug Nezara viridula, experimentally generated interference vibrations impaired the detection and localization of mating signals—even outside their actual frequency range.³ For the vibration-detecting insect, any additional contact with the environment could thus also introduce a new source of interference. Traffic, machinery, and construction work would intrude upon her perception precisely where she seeks shelter. The nest would remain dark, but it would no longer be a place of peace.
¹ G. R. Broad and D. L. J. Quicke, “The adaptive significance of host location by vibrational sounding in parasitoid wasps,” Proceedings of the Royal Society B: Biological Sciences, 267, No. 1460 (2000), pp. 2403–2409. https://doi.org/10.1098/rspb.2000.1298
² J. Samietz, S. Kroder, D. Schneider, and S. Dorn, “Ambient temperature affects mechanosensory host location in a parasitic wasp,” Journal of Comparative Physiology A, 192 (2006), pp. 151–157. https://doi.org/10.1007/s00359-005-0057-6
³ R. Janža, N. Stritih-Peljhan, A. Škorjanc, J. Polajnar, and M. Virant-Doberlet, “Vibrational noise disrupts Nezara viridula communication, irrespective of spectral overlap,” Communications Biology, 7 (2024), Article 1533. https://doi.org/10.1038/s42003-024-07185-3
DAD-015
The Rasping One
(Apparatus stridulatorius)
Description
A finely ribbed surface forms at the junction of two abdominal segments. A hardened edge of the adjacent segment strokes across it as soon as the abdomen moves: the body becomes an instrument. Short, rhythmic impulses are transmitted through the outstretched legs into the nest structure. The environmental pressure driving this design is the increasing daytime heat, which prolongs stays in the sheltered nest and intensifies the shared use of limited retreat spaces. Amid densely packed bodies and winding honeycombs, the “Rasping One” supplements chemical and tactile communication with mechanical signals. She can attract attention without directly touching another worker. The paper walls become a medium of transmission; a small movement generates a message that extends beyond her own body.
Scientific Context
In leafcutter ants, substrate vibrations generated by stridulation serve to recruit nestmates in the immediate vicinity.¹ In Myrmica ants, auditory signals can also convey social status: the sounds produced by queens trigger special attention from workers—an effect that certain butterfly larvae and pupae exploit by mimicking these sounds. ² At the same time, experiments with Nezara viridula show how disruptive vibrations can impair the detection and localization of social signals.³ For leaf-cutting ants, this creates a delicate balance between communication and interference. What allows a single worker to be heard could cause signals to overlap when many ants are transmitting simultaneously. Traffic and machinery would introduce additional vibrations. The colony could generate more and more messages yet be able to reliably distinguish fewer and fewer of them.
¹ F. Roces and B. Hölldobler, “Use of stridulation in foraging leaf-cutting ants: Mechanical support during cutting or short-range recruitment signal?”, Behavioral Ecology and Sociobiology, 39, No. 5 (1996), pp. 293–299. https://doi.org/10.1007/s002650050292
² F. Barbero, J. A. Thomas, S. Bonelli, E. Balletto, and K. Schönrogge, “Queen ants make distinctive sounds that are mimicked by a butterfly social parasite,” Science, 323, No. 5915 (2009), pp. 782–785. https://doi.org/10.1126/science.1163583
³ R. Janža, N. Stritih-Peljhan, A. Škorjanc, J. Polajnar, and M. Virant-Doberlet, “Vibrational noise disrupts Nezara viridula communication, irrespective of spectral overlap,” Communications Biology, 7 (2024), Article 1533. https://doi.org/10.1038/s42003-024-07185-3
About the Project
In the summer of 2024, a Nosferatu spider crawled into my Berlin apartment through an open window. To me, it was terrifyingly large and incredibly fast. The fact that it can bite didn’t make the situation any easier. I couldn’t find a pest control technician willing to remove it, so I spent several days with it in my apartment. It was a nightmare. At the same time, I began to read: about its origins, its behavior, what it’s capable of, and what there actually is to fear from it. I wanted to understand what I was involuntarily sharing my living space with.
Zoropsis spinimana is native to the Mediterranean region, including North Africa, and has expanded its range into Central Europe. Since it was first recorded in Germany in 2005, it has been documented in numerous other locations, including Berlin.¹ ² Human transport, sheltered habitats within buildings, and possibly more favorable climatic conditions are among the proposed explanations for this development.
My research on this one animal led me to explore changes that have long been taking place. Range boundaries are shifting; studies of European bird and butterfly communities show a trend toward thermophilic species. The seasonal activity patterns of butterflies are also changing as temperatures rise.³ ⁴ What I had previously perceived primarily as a prediction became a question directed at my immediate surroundings: What is already changing, even as I still regard it as a single, unusual encounter?
This question led me from a spider to insects, from observable changes to possible futures, and finally to my collection of dead wasps. Directory of Anticipated Deviations gives form to this exploration. Using AI, the microscopic photographs are transformed into bodies that reveal the possibilities and limitations of survival under changing conditions. I am also preoccupied with my own reaction: how quickly the unfamiliar triggers fear, how little I know about the animals around me, and how my perspective shifts as soon as I look more closely.
Sources
¹ A. Hänggi and A. Bolzern, ‘Zoropsis spinimana (Araneae: Zoropsidae) neu für Deutschland’, Arachnologische Mitteilungen, 32 (2006). https://doi.org/10.5431/aramit3201
² ‘Updated distribution of Zoropsis spinimana (Dufour, 1820; Araneae: Zoropsidae) in Germany and novel insights into its ecology based on a citizen science survey’, Frontiers in Arachnid Science (2024). https://doi.org/10.3389/frchs.2024.1383339
³ V. Devictor et al., ‘Differences in the climatic debts of birds and butterflies at a continental scale’, Nature Climate Change, 2 (2012), S. 121–124. https://doi.org/10.1038/nclimate1347
⁴ J. C. Habel et al., ‘Active around the year: Butterflies and moths adapt their life cycles to a warming world’, Global Change Biology, 30, e17103. https://doi.org/10.1111/gcb.17103