Hostname: page-component-745bb68f8f-grxwn Total loading time: 0 Render date: 2025-01-24T18:21:31.623Z Has data issue: false hasContentIssue false
  • English
  • Français

Requirements and limits of anatomy-based predictions of locomotion in terrestrial arthropods with emphasis on arachnids

Published online by Cambridge University Press:  09 May 2016

Tom Weihmann ,
Hanns Hagen Goetzke  and
Michael Günther
Tom Weihmann
Affiliation:
Department of Zoology, University of Cambridge, Downing Street, CB2 3EJ, United Kingdom 〈[email protected]〉; 〈[email protected]〉
Hanns Hagen Goetzke
Affiliation:
Department of Zoology, University of Cambridge, Downing Street, CB2 3EJ, United Kingdom 〈[email protected]〉; 〈[email protected]〉
Michael Günther
Affiliation:
Institute of Sport and Motion Science, University of Stuttgart, Allmandring 28, 70569 Stuttgart, Germany 〈[email protected]〉
Get access Rights & Permissions [Opens in a new window]

Abstract

Modern computer-aided techniques foster the availability and quality of 3D visualization and reconstruction of extinct and extant species. Moreover, animated sequences of locomotion and other movements find their way into motion pictures and documentary films, but also gain attraction in science. While movement analysis is well advanced in vertebrates, particularly in mammals and birds, analyses in arthropods, with their much higher variability regarding general anatomy and size, are still in their infancies and restricted to a few laboratory species. These restrictions and deficient understanding of terrestrial arthropod locomotion in general impedes sensible reconstruction of movements in those species that are not directly observable (e.g., extinct and cryptic species). Since shortcomings like over-simplified approaches to simulate arthropod locomotion became obvious recently, in this review we provide insight into physical, morphological, physiological, behavioral, and ecological constraints, which are essential for sensible reconstructions of terrestrial arthropod locomotion. Such concerted consideration along with sensible evaluations of stability and efficiency requirements can pave the way to realistic assessment of leg coordination and body dynamics.

Type
Articles
Information
Journal of Paleontology , Volume 89 , Issue 6 , November 2015 , pp. 980 - 990
Copyright
Copyright © 2016, The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Ache, J.M., and Matheson, T., 2012, Passive resting state and history of antagonist muscle activity shape active extensions in an insect limb: Journal of Neurophysiology, v. 107, p. 2756–2768.Google Scholar
Alexander, R.M., 1977, Mechanics and scaling of terrestrial locomotion, in Pedley, T.J., ed., Scale Effects in Animal Locomotion, New York, Academic Press, p. 93–110.Google Scholar
Alexander, R.M., and Bennet-Clark, H.C., 1977, Storage of elastic strain energy in muscle and other tissues: Nature, v. 265, p. 114–117.Google Scholar
Anderson, J.F., and Prestwich, K.N., 1975, The fluid pressure pumps of spiders (Chelicerata, Araneae): Zeitschrift für Morphologie der Tiere, v. 81, p. 257–277.Google Scholar
Arditti, J., Elliott, J., Kitching, I.J., and Wasserthal, L.T., 2012, ‘Good Heavens what insect can suck it’–Charles Darwin, Angraecum sesquipedale and Xanthopan morganii praedicta: Botanical Journal of the Linnean Society, v. 169, p. 403–432.Google Scholar
Barth, F.G., 2002, A Spider’s World Senses and Behavior, Berlin, Springer, 394 p.Google Scholar
Biancardi, C.M., Fabrica, C.G., Polero, P., Loss, J.F., and Minetti, A.E., 2011, Biomechanics of octopedal locomotion: kinematic and kinetic analysis of the spider Grammostola mollicoma: Journal of Experimental Biology, v. 214, p. 3433–3442.Google Scholar
Biewener, A., 1990, Biomechanics of mammalian terrestrial locomotion: Science, v. 250, p. 1097–1103.Google Scholar
Biewener, A., 2003, Animal Locomotion: Oxford University Press, 294 p.Google Scholar
Blickhan, R., 1989, The spring-mass model for running and hopping: Journal of Biomechanics, v. 22, p. 1217–1227.Google Scholar
Blickhan, R., and Full, R.J., 1987, Locomotion energetics of the ghost crab: II Mechanics of the center of mass: Journal of Experimental Biology, v. 130, p. 155–174.Google Scholar
Blickhan, R., and Full, R.J., 1993, Similarity in multilegged locomotion: bouncing like a monopode: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 173, p. 509–517.Google Scholar
Blickhan, R., Seyfarth, A., Geyer, H., Grimmer, S., Wagner, H., and Günther, M., 2007, Intelligence by mechanics: Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Science, v. 365, p. 199–220.Google Scholar
Bowerman, R.F., 1975, The control of walking in the scorpion: Journal of Comparative Physiology, v. 100, p. 197–209.Google Scholar
Brüssel, A., 1987, Belastungen und Dehnungen im Spinnenskelett unter natürlichen Verhaltensbedingungen [Dissertation]: Frankfurt am Main, J.W.Goethe Universität, 178 p.Google Scholar
Bullard, B., Garcia, T., Benes, V., Leake, M.C., Linke, W.A., and Oberhauser, A.F., 2006, The molecular elasticity of the insect flight muscle proteins projectin and kettin: Proceedings of the National Academy of Sciences, v. 103, p. 4451–4456.Google Scholar
Burden, S.A., 2014, A Hybrid Dynamical Systems Theory for Legged Locomotion [Ph.D. dissertation]: University of California, Berkeley, 123 p.Google Scholar
Cavagna, G.A., Saibene, F.P., and Margaria, R., 1964, Mechanical work in running: Journal of Applied Physiology, v. 19, p. 249–256.Google Scholar
Clarke, J., 1986, The comparative functional morphology of the leg joints and muscles of five spiders: Bulletin of the British Arachnological Society, v. 7, p. 37–47.Google Scholar
Cruse, H., 1976, The function of the legs in the free walking stick insect, Carausius morosus: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 112, p. 235–262.Google Scholar
Cruse, H., Dürr, V., and Schmitz, J., 2007, Insect walking is based on a decentralized architecture revealing a simple and robust controller: Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Science, v. 365, p. 221–250.Google Scholar
De Groote, F., Jonkers, I., and Duysens, J., 2014, Task constraints and minimization of muscle effort result in a small number of muscle synergies during gait: Frontiers in Computational Neuroscience, v. 8, 11 p.CrossRefGoogle Scholar
Dickinson, M.H., Farley, C.T., Full, R.J., Koehl, M.A.R., Kram, R., and Lehman, S., 2000, How animals move: An integrative view: Science, v. 288, p. 100–106.Google Scholar
Dudek, D.M., and Full, R.J., 2006, Passive mechanical properties of legs from running insects: Journal of Experimental Biology, v. 209, p. 1502–1515.Google Scholar
Dudek, D.M., and Full, R.M., 2007, An isolated insect leg’s passive recovery from dorso-ventral perturbations: Journal of Experimental Biology, v. 210, p. 3209–3217.Google Scholar
Dunlop, J.A., and Brauckmann, C., 2006, A new trigonotarbid arachnid from the Coal Measures of Hagen-Vorhalle, Germany: Fossil Record, v. 9, p. 130–136.Google Scholar
Ehlers, M., 1939, Untersuchungen über Formen aktiver Lokomotion bei Spinnen: Zoologische Jahrbücher: Abteilung für Systematik, Oekologie und Geographie der Tiere, v. 72, p. 373–499.Google Scholar
Endlein, T., and Federle, W., 2008, Walking on smooth or rough ground: passive control of pretarsal attachment in ants: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 194, p. 49–60.Google Scholar
Falkingham, P., 2014, Interpreting ecology and behavior from the vertebrate fossil track record: Journal of Zoology, v. 292, p. 222–228.Google Scholar
Fayers, S.R., and Trewin, N.H., 2003, A review of the palaeoenvironments and biota of the Windyfield chert: Transactions of the Royal Society of Edinburgh: Earth Sciences, v. 94, p. 325–339.Google Scholar
Ferdinand, W., 1981, Die Lokomotion der Krabbenspinnen (Araneae, Thomisidae) und das Wilsonsche Modell der metachronen Koordination: Zoologische Jahrbucher Abteilung fuer Allgemeine Zoologie und Physiologie der Tiere, v. 85, p. 46–65.Google Scholar
Fischer, M.S., and Blickhan, R., 2006, The tri-segmented limbs of therian mammals: kinematics, dynamics, and self-stabilization—a review: Journal of Experimental Zoology Part A: Ecological Genetics and Physiology, v. 305, p. 935–952.Google Scholar
Fröhlich, A., 1978, Verhaltensphysiologische Untersuchungen zur Lokomotion der Spinnen Agelena labyrinthica Cl. und Sitticus pubescens F. [Inaugural-Dissertation]: Berlin, FU – Berlin, 129 p.Google Scholar
Fröschl, M., Handschuh, S., Erlach, R., Schwaha, T., Goldammer, H., Fragner, R., and Walzl, M.G., 2014, Computer-generated images of microscopic soil organisms for documentary films: Soil Organisms, v. 86, p. 95–102.Google Scholar
Full, R.J., and Koditschek, D.E., 1999, Templates and anchors: neuromechanical hypotheses of legged locomotion on land: Journal of Experimental Biology, v. 202, p. 3325–3332.Google Scholar
Full, R.J., and Koehl, M.A.R., 1992, Drag and lift on running insects: Journal of Experimental Biology, v. 176, p. 89–101.Google Scholar
Full, R.J., Blickhan, R., and Ting, L.H., 1991, Leg design in hexapedal runners: Journal of Experimental Biology, v. 158, p. 369–390.Google Scholar
Fusi, L., Brunello, E., Reconditi, M., Piazzesi, G., and Lombardi, V., 2014, The non-linear elasticity of the muscle sarcomere and the compliance of myosin motors: Journal of Physiology, v. 592, p. 1109–1118.Google Scholar
Garwood, R., and Dunlop, J., 2014, The walking dead: Blender as a tool for paleontologists with a case study on extinct arachnids: Journal of Paleontology, v. 88, p. 735–746.Google Scholar
Geyer, H., Seyfarth, A., and Blickhan, R., 2003, Positive force feedback in bouncing gaits?: Proceedings of the Royal Society B: Biological Sciences, v. 270, p. 2173–2183.Google Scholar
Geyer, H., Seyfarth, A., and Blickhan, R., 2005, Spring-mass running: simple approximate solution and application to gait stability: Journal of Theoretical Biology, v. 232, p. 315–328.Google Scholar
Goyens, J., Dirckx, J., Aerts, P., and Davidowitz, G., 2015, Costly sexual dimorphism in Cyclommatus metallifer stag beetles: Functional Ecology, v. 29, p. 35–43.Google Scholar
Gradstein, F., and Ogg, J., 2004, Geologic time scale 2004–why, how, and where next!: Lethaia, v. 37, p. 175–181.Google Scholar
Grimmer, S., Ernst, M., Günther, M., and Blickhan, R., 2008, Running on uneven ground: leg adjustment to vertical steps and self-stability: Journal of Experimental Biology, v. 211, p. 2989–3000.Google Scholar
Günther, M., and Weihmann, T., 2011, The load distribution among three legs on the wall: model predictions for cockroaches: Archive of Applied Mechanics, v. 81, p. 1269–1287.Google Scholar
Günther, M., and Weihmann, T., 2012, Climbing in hexapods: a plain model for heavy slopes: Journal of Theoretical Biology, v. 293, p. 82–86.Google Scholar
Günther, M., Keppler, V., Seyfarth, A., and Blickhan, R., 2004, Human leg design: optimal axial alignment under constraints: Journal of Mathematical Biology, v. 48, p. 623–646.Google Scholar
Guo, S., Chang, J., Yang, X., Wang, W., and Zhang, J., 2014, Locomotion skills for insects with sample-based controller: Computer Graphics Forum, v. 33, p. 31–40.Google Scholar
Haeufle, D.F.B., Günther, M., Wunner, G., and Schmitt, S., 2014, Quantifying control effort of biological and technical movements: An information-entropy-based approach: Physical Review E, 89:1:012716, http://dx.doi.org/10.1103/PhysRevE.89.012716.Google Scholar
Hill, D.A., 2006, Targeted jumps by salticid spiders (Araneae, Salticidae, Phidippus). Peckhamia Version 9, p. 1–28, http://www.peckhamia.com/epublications/Hill%202006%20Targeted%20jumps%20by%20salticid%20spiders%20V9%20EB%20PDF.pdf.Google Scholar
Hooper, S.L., Guschlbauer, C., Blumel, M., Rosenbaum, P., Gruhn, M., Akay, T., and Buschges, A., 2009, Neural control of unloaded leg posture and of leg swing in stick insect, cockroach, and mouse differs from that in larger animals: Journal of Neuroscience, v. 29, p. 4109–4119.Google Scholar
Hughes, G.M., 1952, The co-ordination of insect movements. I: The walking movements of insects: Journal of Experimental Biology, v. 29, p. 267–284.Google Scholar
Ijspeert, A.J., 2008, Central pattern generators for locomotion control in animals and robots: a review: Neural Networks, v. 21, p. 642–653.Google Scholar
Jacobi-Kleemann, M., 1953, Über die Locomotion der Kreuzspinne Aranea diadema beim Netzbau (nach Filmanalysen): Journal of Comparative Physiology, v. 34, P. 606–654.Google Scholar
Lewis, J.G.E., 1981, The Biology of Centipedes, Cambridge, UK, Cambridge University Press, 476 p.Google Scholar
Lipfert, S.W., Günther, M., Renjewski, D., and Seyfarth, A., 2014, Impulsive ankle push-off powers leg swing in human walking: Journal of Experimental Biology, v. 217, p. 1218–1228.Google Scholar
Maier, L., Root, T.M., and Seyfarth, E.A., 1987, Heterogeneity of spider leg muscle: Histochemistry and electrophysiology of identified fibers in the claw levator: Journal of Comparative Physiology B: Biochemical, Systems, and Environmental Physiology, v. 157, p. 285–294.Google Scholar
Manton, S.M., 1950, The evolution of arthropodan locomotory mechanisms.—Part I. The locomotion of Peripatus: Zoological Journal of the Linnean Society, v. 41, p. 529–570.Google Scholar
Manton, S.M., 1952, The evolution of arthropodan locomotory mechanisms.—Part 2. General introduction to the locomotory mechanisms of the arthropoda: Zoological Journal of the Linnean Society, v. 42, p. 93–117.Google Scholar
Manton, S.M., 1977, The Arthropoda: Habits, Functional Morphology, and Evolution, Oxford, Clarendon Press, 527 p.Google Scholar
Melchers, M., 1967, Der Beutefang von Cupiennius salei Keyserling (Ctenidae): Zeitschrift für Morphologie und Ökologie der Tiere, v. 58, p. 321–346.Google Scholar
Mendes, C.S., Bartos, I., Akay, T., Márka, S., and Mann, R.S., 2013, Quantification of gait parameters in freely walking wild type and sensory deprived Drosophila melanogaster: eLife, 2013:2:e00231, doi: http://dx.doi.org/10.7554/eLife.00231.001.Google Scholar
Minter, N.J., Mángano, M.G., and Caron, J.-B., 2012, Skimming the surface with Burgess Shale arthropod locomotion: Proceedings of the Royal Society B: Biological Sciences, v. 279, p. 1613–1620.Google Scholar
Mochon, S., and McMahon, T.A., 1980, Ballistic walking: Journal of Biomechnics, v. 13, p. 49–57.Google Scholar
Moffett, S., and Doell, G. S., 1980, Alteration of locomotor behavior in wolf spiders carrying normal and weighted egg cocoons: Journal of Experimental Zoology, v. 213, p. 219–226.Google Scholar
Moll, K., Roces, F., and Federle, W., 2013, How load-carrying ants avoid falling over: mechanical stability during foraging in Atta vollenweideri grass-cutting ants: PLoS One, 8:1:e52816, doi:10.1371/journal.pone.0052816.Google Scholar
Parry, D.A., and Brown, R.H.J., 1959a, The hydraulic mechanism of the spider leg: Journal of Experimental Biology, v. 36, p. 423–433.Google Scholar
Parry, D.A., and Brown, R.H.J., 1959b, The jumping mechanism of salticid spiders: Journal of Experimental Biology, v. 36, p. 654–664.Google Scholar
Patek, S., Dudek., D., and Rosario, M., 2011, From bouncy legs to poisoned arrows: elastic movements in invertebrates: Journal of Experimental Biology, v. 214, p. 1973–1980.Google Scholar
Piazzesi, G., Reconditi, M., Linari, M., Lucii, L., Bianco, P., Brunello, E., Decostre, V., Stewart, A., Gore, D.B., and Irving, T.C., 2007, Skeletal muscle performance determined by modulation of number of myosin motors rather than motor force or stroke size: Cell, v. 131, p. 784–795.Google Scholar
Pohl, H., Wipfler, B., Grimaldi, D., Beckmann, F., and Beutel, R.G., 2010, Reconstructing the anatomy of the 42-million-year-old fossil—Mengea tertiaria (Insecta, Strepsiptera): Naturwissenschaften, v. 97, p. 855–859.Google Scholar
Prange, H.D., 1977, The Scaling and Mechanics of Arthropod Exoskeletons, in Pedley, T.J., ed., Scale Effects in Animal Locomotion, London, Academic Press, p. 169–181.Google Scholar
Reinhardt, L., Weihmann, T., and Blickhan, R., 2009, Dynamics and kinematics of ant locomotion: do wood ants climb on level surfaces?: Journal of Experimental Biology, v. 212, p. 2426–2435.Google Scholar
Ringrose, R.P., 1997, Self-stabilizing Running, Boston, Massachusetts Institute of Technology, 131 p.Google Scholar
Rode, C., Siebert, T., Herzog, W., and Blickhan, R., 2009, The effects of parallel and series elastic components on the active cat soleus force-length relationship: Journal of Mechanics in Medicine and Biology, v. 9, p. 105–122.Google Scholar
Rosenfeld, E.V., and Günther, M., 2014, An enhanced model of cross-bridge operation with internal elasticity: European Biophysics Journal, v. 43, p. 131–141.Google Scholar
Rovner, J.S., 1980, Morphological and ethological adaptations for prey capture in wolf spiders (Aranae, Lycosidae): Journal of Arachnology, v. 8, p. 201–215.Google Scholar
Ruhland, M., and Rathmayer, W., 1978, Die Beinmuskulatur und ihre Innervation bei der Vogelspinne Dugesiella hentzi (Ch.) (Araneae, Aviculariidae): Zoomorphologie, v. 89, p. 33–46.Google Scholar
Ruina, A., Bertram, J.E., and Srinivasan, M., 2005, A collisional model of the energetic cost of support work qualitatively explains leg sequencing in walking and galloping, pseudo-elastic leg behavior in running and the walk-to-run transition: Journal of Theoretical Biology, v. 237, p. 170–192.Google Scholar
Schmitt, J., and Bonnono, S., 2009, Dynamics and stability of lateral plane locomotion on inclines: Journal of Theoretical Biology, v. 261, p. 598–609.CrossRefGoogle ScholarPubMed
Schmitt, J., and Holmes, P., 2000, Mechanical models for insect locomotion: dynamics and stability in the horizontal plane-II. Application: Biological Cybernetics, v. 83, p. 517–527.Google Scholar
Sens, J., 1996, Funktionelle Anatomie und Biomechanik der Laufbeine einer Vogelspinne [Dissertation]:, Saarbrücken, Universität des Saarlandes, Saarbrücken, 344 p.Google Scholar
Sensenig, A.T., and Shultz, J.W., 2003, Mechanics of cuticular elastic energy storage in leg joints lacking extensor muscles in arachnids: Journal of Experimental Biology, v. 206, p. 771–784.Google Scholar
Sensenig, A.T., and Shultz, J.W., 2006, Mechanical energy oscillations during locomotion in the harvestman Leiobunum vittatum (Opiliones): Journal of Arachnology, v. 34, p. 627–633.Google Scholar
Seyfarth, A., Günther, M., and Blickhan, R., 2001, Stable operation of an elastic three-segment leg: Biological Cybernetics, v. 84, p. 365–382.Google Scholar
Seyfarth, A., Geyer, H., Günther, M., and Blickhan, R., 2002, A movement criterion for running: Journal of Biomechanics, v. 35, p. 649–655.Google Scholar
Shultz, J.W., 1987, Walking and surface film locomotion in terrestrial and semiaquatic spiders: Journal of Experimental Biology, v. 128, p. 427–444.Google Scholar
Shultz, J.W., 1989, Morphology of locomotor appendages in Arachnida: evolutionary trends and phylogenetic implications: Zoological Journal of the Linnean Society, v. 97, p. 1–56.Google Scholar
Siebert, T., Weihmann, T., Rode, C., and Blickhan, R., 2010, Cupiennius salei: biomechanical properties of the tibia-metatarsus joint and its flexing muscles: Journal of Comparative Physiology B: Biochemical, Systems, and Environmental Physiology, v. 180, p. 199–209.Google Scholar
Spagna, J.C., Goldman, D.I., Lin, P.C., Koditschek, D.E., and Full, R.J., 2007, Distributed mechanical feedback in arthropods and robots simplifies control of rapid running on challenging terrain: Bioinspiration & Biomimetics, v. 2, p. 9–18.Google Scholar
Spagna, J.C., Valdivia, E.A., and Mohan, V., 2011, Gait characteristics of two fast-running spider species (Hololena adnexa and Hololena curta), including an aerial phase (Araneae: Agelenidae): Journal of Arachnology, v. 39, p. 84–91.Google Scholar
Sponberg, S., and Full, R.J., 2008, Neuromechanical response of musculo-skeletal structures in cockroaches during rapid running on rough terrain: Journal of Experimental Biology, v. 211, p. 433–446.Google Scholar
Stewert, D.M., and Martin, A.W., 1974, Blood pressure in the tarantula Dugesiella hentzi: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 88, p. 141–172.Google Scholar
Sutton, M.D., 2008, Tomographic techniques for the study of exceptionally preserved fossils: Proceedings of the Royal Society B: Biological Sciences, v. 275, p. 1587–1593.Google Scholar
Ting, L.H., Blickhan, R., and Full, R.J., 1994, Dynamic and static stability in hexapedal runners: Journal of Experimental Biology, v. 197, p. 251–269.Google Scholar
Wahl, V., Pfeffer, S.E., and Wittlinger, M., 2015, Walking and running in the desert ant Cataglyphis fortis: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 201, p. 645–656.Google Scholar
Ward, T.M., and Humphreys, W.F., 1981, Locomotion in burrowing and vagrant wolf spiders (Lycosidae): Journal of Experimental Biology, v. 92, p. 305–321.Google Scholar
Weihmann, T., 2007, Biomechanische Analyse der ebenen Lokomotion von Ancylometes bogotensis (Keyserling, 1877) (Chelicerata, Arachnida, Lycosoidea) [doctoral thesis]:, Jena, Friedrich-Schiller-Universität, 99 p. urn:nbn:de:gbv:27-20080418-114748-4.Google Scholar
Weihmann, T., 2013, Crawling at High Speeds: Steady Level Locomotion in the Spider Cupiennius salei—Global Kinematics and Implications for Centre of Mass Dynamics: PLoS One, 8:6:e65788, doi:10.1371/journal.pone.0065788.Google Scholar
Weihmann, T., Günther, M., and Blickhan, R., 2012, Hydraulic leg extension is not necessarily the main drive in large spiders: Journal of Experimental Biology, v. 215, p. 578–583.Google Scholar
Weihmann, T., Karner, M., Full, R.J., and Blickhan, R., 2010, Jumping kinematics in the wandering spider Cupiennius salei: Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, v. 196, p. 421–438.Google Scholar
Weihmann, T., Siebert, T., and Blickhan, R., 2009, Muscle properties of the tibia-metatarsus joint flexors in the labidognath spider Cupiennius salei: Comparative Biochemistry and Physiology - Part A: Molecular & Integrative Physiology, v. 153, p. 124–125.Google Scholar
Wendler, G., 1964, Laufen und Stehen der Stabheuschrecke Carausius morosus: Sinnesborstenfelder in den Beingelenken als Glieder von Regelkreisen: Zeitschrift für vergleichende Physiologie, v. 48, p. 198–250.Google Scholar
Wilson, D.M., 1967, Stepping patterns in tarantula spiders: Journal of Experimental Biology, v. 47, p. 133–151.Google Scholar
Wosnitza, A., Bockemühl, T., Dübbert, M., Scholz, H., and Büschges, A., 2013, Inter-leg coordination in the control of walking speed in Drosophila: Journal of Experimental Biology, v. 216, p. 480–491.Google Scholar
Wu, G.C., Wright, J.C., Whitaker, D.L., and Ahn, A.N., 2010, Kinematic evidence for superfast locomotory muscle in two species of teneriffiid mites: Journal of Experimental Biology, v. 213, p. 2551–2556.Google Scholar
Zollikofer, C., 1994, Stepping patterns in ants—influence of load: Journal of Experimental Biology, v. 192, p. 119–127.Google Scholar