Abstract
In terrestrial legged locomotion, the distribution of mass can influence the gait characteristics. This can be due to a change in the magnitude or distribution of the load. The latter occurs in scorpions when they lift their large metasoma from a trailing position in ambulatory posture to the well-known arched forward position in the defensive posture. We measured how locomotion changes between these two postures by recording scorpions walking using high-speed video. We found that the metasoma in the fat-tailed scorpion (Androctonus australis) represents about a quarter of the total mass. Moving this mass anteriorly over the body changes the position of the center of mass forward 8.15 ± 1.86 mm. We found this increases the overall duty factor, and particularly that of the second leg pair, even when taking the reduced speed in defensive posture into account. In the five scorpions we recorded, also the ipsilateral phase of leg pairs 3 and 4 differed in defensive posture. We found that the trajectory the 4th foot describes during a single stride also differed significantly between postures, showing this to be a sensitive measure of changes in gait. The change from an ambulatory to a defensive posture places different demands on the gait of scorpions, possibly largely due to the forward displacement of the center of mass.
Similar content being viewed by others
References
Adams DC, Cerney MM (2007) Quantifying biomechanical motion using Procrustes motion analysis. J Biomech 40:437–444. https://doi.org/10.1016/j.jbiomech.2005.12.004
Adams DC, Otárola-Castillo E (2013) Geomorph: an R package for the collection and analysis of geometric morphometric shape data. Methods Ecol Evol 4:393–399. https://doi.org/10.1111/2041-210X.12035
Alexander RMN (2013) Principles of animal locomotion. Princeton University Press, Princeton
Alexander AJ, Ewer DW (1958) Temperature adaptive behaviour in the scorpion, Opisthophthalmus latimanus Koch. J Exp Biol 35:349–359
Autumn K, Han B (1989) Mimicry of scorpions by juvenile lizards, Teratoscincus roborowskii. Chin Herpetol Res 2:60–64
Bateman PW, Fleming PA (2009) To cut a long tail short: a review of lizard caudal autotomy studies carried out over the last 20 years. J Zool 277:1–14. https://doi.org/10.1111/j.1469-7998.2008.00484.x
Bowerman RF (1975a) The control of walking in the scorpion. II. Coordination modification as a consequence of appendage ablation. J Comp Physiol 100:197–209
Bowerman RF (1975b) The control of walking in scorpion. J Comp Physiol 100:183–196
Brady LF (1947) Invertebrate tracks from the Coconino Sandstone of Northern Arizona. J Paleontol 21:466–472
Brandão RA, Motta PC (2005) Circumstantial evidences for mimicry of scorpions by the neotropical gecko Coleodactylus brachystoma (Squamata, Gekkonidae) in the Cerrados of central Brazil. Phyllomedusa 4:139–145. https://doi.org/10.11606/issn.2316-9079.v4i2p139-145
Carlson BE, McGinley S, Rowe MP (2014) Meek males and fighting females: sexually-dimorphic antipredator behavior and locomotor performance is explained by morphology in bark scorpions (Centruroides vittatus). PLoS One 9:e97648. https://doi.org/10.1371/journal.pone.0097648
Chiari Y, van der Meijden A, Caccone A et al (2017) Self-righting potential and the evolution of shell shape in Galápagos tortoises. Sci Rep 7:15828. https://doi.org/10.1038/s41598-017-15787-7
Chippaux J-P, Goyffon M (2008) Epidemiology of scorpionism: a global appraisal. Acta Trop 107:71–79. https://doi.org/10.1016/j.actatropica.2008.05.021
Coelho P, Kaliontzopoulou A, Rasko M, van der Meijden A (2017) A ‘striking’ relationship: scorpion defensive behaviour and its relation to morphology and performance. Funct Ecol 31:1390–1404. https://doi.org/10.1111/1365-2435.12855
Dean J (1991) Effect of load on leg movement and step coordination of the stick insect Carausius morosus. J Exp Biol 159:449–471. https://doi.org/10.17705/1jais.00484
Eiseman C, Charney N, Carlson J (2010) Tracks and sign of insects and other invertebrates: a guide to North American Species. Stackpole Books, Mechanicsburg
Fernandez Rodriguez I, Brana F (2020) The movement dynamics of autotomized lizards and their tails reveal functional costs of caudal autotomy. Integr Zool 15:511–521. https://doi.org/10.1111/1749-4877.12443
Fet V, Fet E, Neff D, Graham M (2003) Metasoma of Orthochirus (Scorpiones: Buthidae): are scorpions evolving a new sensory organ? Rev Ibérica Aracnol 8:69–72
Fleming PA, Bateman PW (2007) Just drop it and run: the effect of limb autotomy on running distance and locomotion energetics of field crickets (Gryllus bimaculatus). J Exp Biol 210:1446–1454. https://doi.org/10.1242/jeb.02757
Full RJ (2011) Invertebrate locomotor systems. Comprehensive physiology. Wiley, Hoboken
Galbraith DA, Cloudsley-Thompson JL (1992) Ecophysiology of desert arthropods and reptiles. Springer Science and Business Media, Berlin
Geethabali BY, Rao KP (1973) A metasomatic neural photoreceptor in the scorpion. J Exp Biol 58:189–196
Gillis GB, Bonvini LA, Irschick DJ (2009) Losing stability: tail loss and jumping in the arboreal lizard Anolis carolinensis. J Exp Biol 212:604–609. https://doi.org/10.1242/jeb.024349
Gillis B, Kuo G, Irschick C-Y D (2013) The Impact of Tail Loss on Stability during Jumping in Green Anoles (Anolis carolinensis). Physiol Biochem Zool 86:680–689. https://doi.org/10.1086/673756
Görner M, Hirzinger G (2010) Analysis and evaluation of the stability of a biologically inspired, Leg loss tolerant gait for six- and eight-legged walking robots. In: 2010 IEEE international conference on robotics and automation. IEEE, pp 4728–4735
Hedrick TL (2008) Software techniques for two- and three-dimensional kinematic measurements of biological and biomimetic systems. Bioinspir Biomim 3:034001. https://doi.org/10.1088/1748-3182/3/3/034001
Hickman GC (1979) The mammalian tail: a review of functions. Mamm Rev 9:143–157. https://doi.org/10.1111/j.1365-2907.1979.tb00252.x
Holmes P, Full RJ, Koditschek D, Guckenheimer J (2006) The dynamics of legged locomotion: models, analyses, and challenges. SIAM Rev 48:207–304. https://doi.org/10.1137/S0036144504445133
Hsieh S-TT (2016) Tail loss and narrow surfaces decrease locomotor stability in the arboreal green anole lizard (Anolis carolinensis). J Exp Biol 219:364–373. https://doi.org/10.1242/jeb.124958
Lemelin P, Schmitt D (2004) Seasonal variation in body mass and locomotor kinetics of the fat-tailed dwarf lemur (Cheirogaleus medius). J Morphol 260:65–71. https://doi.org/10.1002/jmor.10214
Martin J, Avery RA (1998) Effects of tail loss on the movement patterns of the lizard, Psammodromus algirus. Funct Ecol 12:794–802. https://doi.org/10.1046/j.1365-2435.1998.00247.x
McGhee RB, Frank AA (1968) On the stability properties of quadruped creeping gaits. Math Biosci 3:331–351. https://doi.org/10.1016/0025-5564(68)90090-4
Merienne H, Latil G, Moretto P, Fourcassié V (2019) Walking kinematics in the polymorphic seed harvester ant Messor barbarus: influence of body size and load carriage. bioRxiv. https://doi.org/10.1101/614362
Mincer ST, Russo GA (2020) Substrate use drives the macroevolution of mammalian tail length diversity. Proc R Soc B Biol Sci 287:20192885. https://doi.org/10.1098/rspb.2019.2885
Moffett S, Doell GS (1980) Alteration of locomotor behavior in wolf spiders carrying normal and weighted egg cocoons. J Exp Zool 213:219–226. https://doi.org/10.1002/jez.1402130209
Moll K, Roces F, Federle W (2010) Foraging grass-cutting ants (Atta vollenweideri) maintain stability by balancing their loads with controlled head movements. J Comp Physiol A Neuroethol Sens Neural Behav Physiol 196:471–480. https://doi.org/10.1007/s00359-010-0535-3
Moll K, Federle W, Roces F (2012) The energetics of running stability: costs of transport in grass-cutting ants depend on fragment shape. J Exp Biol 215:161–168. https://doi.org/10.1242/jeb.063594
Moll K, Roces F, Federle W (2013) How load-carrying ants avoid falling over: mechanical stability during foraging in Atta vollenweideri grass-cutting ants. PLoS One 8:e52816. https://doi.org/10.1371/journal.pone.0052816
Newlands G, Cantrell AC (1985) A re-appraisal of the rock scorpions (Scorpionidae: Hadogenes). Koedoe 28:35–45. https://doi.org/10.4102/koedoe.v28i1.533
R Development Core Team R (2011) R: a language and environment for statistical computing. R Found Stat Comput 1:409
Rohlf FJ, Corti M (2000) Use of two-block partial least-squares to study covariation in shape. Syst Biol 49:740–753. https://doi.org/10.1080/106351500750049806
Rohlf FJ, Slice D (1990) Extensions of the Procrustes method for the optimal superimposition of landmarks. Syst Zool 39:40. https://doi.org/10.2307/2992207
Root TM, Bowerman RF (1978) Intra-appendage movements during walking in the scorpion Hadrurus arizonensis. Comp Biochem Physiol A 59:49–56. https://doi.org/10.1016/0300-9629(78)90305-5
Shultz BYJW (1992) Muscles firing patterns in two arachnids using different methods of propulsive leg extension. J. Exp. Biol. 162:313–329
Spagna JC, Peattie AM (2012) Terrestrial locomotion in arachnids. J Insect Physiol 58:599–606. https://doi.org/10.1016/j.jinsphys.2012.01.019
Spagna JC, Valdivia EA, Mohan V (2011) Gait characteristics of two fast-running spider species (Hololena adnexa and Hololena curta), including an aerial phase (Araneae: Agelenidae). J Arachnol 39:84–91. https://doi.org/10.1636/B10-45.1
Spenneberg D, McCullough K, Kirchner F (2004) Stability of walking in a multilegged robot suffering leg loss. In: IEEE international conference on robotics and automation, 2004. Proceedings. ICRA ’04, vol 3, 2004. IEEE, 2159–2164
Vogel S (2013) Comparative biomechanics: life’s physical world, 2nd edn. Princeton University Press, Princeton
Ward MJ, Ellsworth SA, Nystrom GS (2018) A global accounting of medically significant scorpions: epidemiology, major toxins, and comparative resources in harmless counterparts. Toxicon 151:137–155. https://doi.org/10.1016/j.toxicon.2018.07.007
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:e65788. https://doi.org/10.1371/journal.pone.0065788
Wolff JO, van der Meijden A, Herberstein ME (2017) Distinct spinning patterns gain differentiated loading tolerance of silk thread anchorages in spiders with different ecology. Proc R Soc B Biol Sci 284:20171124. https://doi.org/10.1098/rspb.2017.1124
Zollikofer CPE (1994) Stepping patterns in ants—influence of load. J Exp Biol 192:119–127
Acknowledgements
AvdM was financed through FCT-Fundação para a Ciência e a Tecnologia, I.P. under contract number DL57/2016/CP1440/CT0009. We thank Arendo Flipse for providing the scorpions.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Conflict of interest
The authors declare that they have no conflict of interests.
Additional information
Publisher’s Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Electronic Supplementary Material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Telheiro, A., Coelho, P. & van der Meijden, A. The effect of change in mass distribution due to defensive posture on gait in fat‐tailed scorpions. J Comp Physiol A 207, 117–125 (2021). https://doi.org/10.1007/s00359-021-01467-5
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00359-021-01467-5