{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T21:39:38Z","timestamp":1784929178416,"version":"3.55.0"},"reference-count":93,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2022,6,9]],"date-time":"2022-06-09T00:00:00Z","timestamp":1654732800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100007108","name":"George Washington University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100007108","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100006498","name":"Clemson University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006498","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IOS 0517240"],"award-info":[{"award-number":["IOS 0517240"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IOS 0817794"],"award-info":[{"award-number":["IOS 0817794"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100014548","name":"Society for Integrative and Comparative Biology","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100014548","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000522","name":"Company of Biologists","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000522","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2022,12,6]]},"abstract":"<jats:title>Synopsis<\/jats:title>\n               <jats:p>Many ecological factors influence animal movement, including properties of the media that they move on or through. Animals moving in terrestrial environments encounter conditions that can be challenging for generating propulsion and maintaining stability, such as inclines and deformable substrates that can cause slipping and sinking. In response, tetrapods tend to adopt a more crouched posture and lower their center of mass on inclines and increase the surface area of contact on deformable substrates, such as sand. Many amphibious fishes encounter the same challenges when moving on land, but how these finned animals modulate their locomotion with respect to different environmental conditions and how these modifications compare with those seen within tetrapods is relatively understudied. Mudskippers (Gobiidae: Oxudercinae) are a particularly noteworthy group of amphibious fishes in this context given that they navigate a wide range of environmental conditions, from flat mud to inclined mangrove trees. They use a unique form of terrestrial locomotion called \u201ccrutching,\u201d where their pectoral fins synchronously lift and vault the front half of the body forward before landing on their pelvic fins, while the lower half of the body and tail are kept straight. However, recent work has shown that mudskippers modify some aspects of their locomotion when crutching on deformable surfaces, particularly those at an incline. For example, on inclined dry sand, mudskippers bent their bodies laterally and curled and extended their tails to potentially act as a secondary propulsor and\/or anti-slip device. In order to gain a more comprehensive understanding of the functional diversity and context-dependency of mudskipper crutching, we compared their kinematics on different combinations of substrate types (solid, mud, and dry sand) and inclines (0\u00b0, 10\u00b0, and 20\u00b0). In addition to increasing lateral bending on deformable and inclined substrates, we found that mudskippers increased the relative contact time and contact area of their paired fins, while becoming more crouched, which are responses comparable to those seen in tetrapods and other amphibious fish. Mudskippers on these substrates also exhibited previously undocumented behaviors, such as extending and adpressing the distal portions of their pectoral fins more anteriorly, dorsoventrally bending their trunk, \u201cbelly-flopping\u201d on sand, and \u201cgripping\u201d the mud substrate with their pectoral fin rays. Our study highlights potential compensatory mechanisms shared among vertebrates in terrestrial environments while also illustrating that locomotor flexibility and even novelty can emerge when animals are challenged with environmental variation.<\/jats:p>","DOI":"10.1093\/icb\/icac084","type":"journal-article","created":{"date-parts":[[2022,6,9]],"date-time":"2022-06-09T15:41:03Z","timestamp":1654789263000},"page":"1335-1356","source":"Crossref","is-referenced-by-count":15,"title":["Mudskippers Modulate their Locomotor Kinematics when Moving on Deformable and Inclined Substrates"],"prefix":"10.1093","volume":"62","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0703-7530","authenticated-orcid":false,"given":"Emily R","family":"Naylor","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, The George Washington University , Washington, D.C. 20052 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1856-5566","authenticated-orcid":false,"given":"Sandy M","family":"Kawano","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, The George Washington University , Washington, D.C. 20052 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2022,6,9]]},"reference":[{"key":"2023060902011238200_bib61","doi-asserted-by":"crossref","first-page":"20170213","DOI":"10.1098\/rsif.2017.0213","article-title":"The coefficient of determination R2 and intra-class correlation coefficient from generalized linear mixed-effects models revisited and expanded","volume":"14","author":"Nakagawa","year":"2017","journal-title":"J R Soc Interface"},{"key":"2023060902011238200_bib1","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1038\/nphys3568","article-title":"Robophysical study of jumping dynamics on granular media","volume":"12","author":"Aguilar","year":"2016","journal-title":"Nat Phys"},{"key":"2023060902011238200_bib2","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1111\/jzo.12934","article-title":"Vertical locomotion in the arboreal salamander Aneides vagrans","volume":"316","author":"Aretz","year":"2022","journal-title":"J Zool"},{"key":"2023060902011238200_bib3","volume-title":"MuMIn: Multi-Model Inference","author":"Barton","year":"2022"},{"key":"2023060902011238200_bib4","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting linear mixed-effects models using lme4","volume":"67","author":"Bates","year":"2015","journal-title":"J Stat Softw"},{"key":"2023060902011238200_bib5","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1093\/icb\/icaa018","article-title":"Mechanical ecology\u2014taking biomechanics to the field","volume":"60","author":"Bauer","year":"2020","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib44","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1242\/jeb.202.2.143","article-title":"Effects of incline and speed on the three-dimensional hindlimb kinematics of a generalized iguanian lizard (Dipsosaurus dorsalis)","volume":"202","author":"BC","year":"1999","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib6","doi-asserted-by":"crossref","first-page":"3706","DOI":"10.1242\/jeb.161109","article-title":"It's just sand between the toes: how particle size and shape variation affect running performance and kinematics in a generalist lizard","volume":"220","author":"Bergmann","year":"2017","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib7","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1093\/icb\/icu015","article-title":"The scaling of uphill and downhill locomotion in legged animals","volume":"54","author":"Birn-Jeffery","year":"2014","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib8","doi-asserted-by":"crossref","first-page":"20140456","DOI":"10.1098\/rsbl.2014.0456","article-title":"Geckos significantly alter foot orientation to facilitate adhesion during downhill locomotion","volume":"10","author":"Birn-Jeffery","year":"2014","journal-title":"Biol Lett"},{"key":"2023060902011238200_bib9","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1186\/s12983-016-0144-2","article-title":"Geckos decouple fore- and hind limb kinematics in response to changes in incline","volume":"13","author":"Birn-Jeffery","year":"2016","journal-title":"Front Zool"},{"key":"2023060902011238200_bib10","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1111\/j.1469-7998.2007.00315.x","article-title":"Ontogenetic change in novel functions: waterfall climbing in adult Hawaiian gobiid fishes","volume":"273","author":"Blob","year":"2007","journal-title":"J Zool"},{"key":"2023060902011238200_bib11","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.zool.2018.10.003","article-title":"A walking behavior generates functional overland movements in the tidepool sculpin, Oligocottus maculosus","volume":"131","author":"Bressman","year":"2018","journal-title":"Zoology"},{"key":"2023060902011238200_bib12","doi-asserted-by":"crossref","first-page":"obz026","DOI":"10.1093\/iob\/obz026","article-title":"Emersion and terrestrial locomotion of the Northern snakehead (Channa argus) on multiple substrates","volume":"1","author":"Bressman","year":"2019","journal-title":"Integr Org Biol"},{"key":"2023060902011238200_bib13","doi-asserted-by":"crossref","first-page":"608","DOI":"10.1643\/i2020084","article-title":"Reffling: a novel locomotor behavior used by Neotropical armored catfishes (Loricariidae) in terrestrial environments","volume":"109","author":"Bressman","year":"2021","journal-title":"Ichthyol Herpetol"},{"key":"2023060902011238200_bib14","first-page":"488","volume-title":"Model Selection and Multimodel Inference: a practical information-theoretic approach","author":"Burnham","year":"2002","edition":"2nd ed."},{"key":"2023060902011238200_bib15","doi-asserted-by":"crossref","first-page":"1187","DOI":"10.1242\/jeb.012989","article-title":"Mechanics of limb bone loading during terrestrial locomotion in river cooter turtles (Pseudemys concinna)","volume":"211","author":"Butcher","year":"2008","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib16","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1152\/jn.1998.79.4.1687","article-title":"Forms of forward quadrupedal locomotion. II. a comparison of posture, hindlimb kinematics, and motor patterns for upslope and level walking","volume":"79","author":"Carlson-Kuhta","year":"1998","journal-title":"J Neurophysiol"},{"key":"2023060902011238200_bib17","doi-asserted-by":"crossref","first-page":"73","DOI":"10.4159\/harvard.9780674184404.c5","article-title":"Climbing","volume-title":"Functional Vertebrate Morphology","author":"Cartmill","year":"1985"},{"key":"2023060902011238200_bib18","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1898\/11-19.1","article-title":"Aquatic and terrestrial locomotion of the rock prickleback, Xiphister mucosus (Cottiformes: Zoarcoidei: Stichaeidae)","volume":"93","author":"Clardy","year":"2012","journal-title":"Northwest Nat"},{"key":"2023060902011238200_bib20","doi-asserted-by":"crossref","first-page":"jeb242906","DOI":"10.1242\/jeb.242906","article-title":"They like to move it (move it): walking kinematics of balitorid loaches of Thailand","volume":"225","author":"Crawford","year":"2022","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib19","volume-title":"These fish were made for walking: morphology and walking kinematics in balitorid loaches","author":"Crawford","year":"2021"},{"key":"2023060902011238200_bib21","doi-asserted-by":"crossref","first-page":"e13406","DOI":"10.1111\/apha.13406","article-title":"Evolutionary and cardio-respiratory physiology of air-breathing and amphibious fishes","volume":"228","author":"Damsgaard","year":"2020","journal-title":"Acta Physiologica"},{"key":"2023060902011238200_bib22","doi-asserted-by":"crossref","first-page":"jeb217562","DOI":"10.1242\/jeb.217562","article-title":"Asymmetrical gait kinematics of free-ranging callitrichine primates in response to changes in substrate diameter and orientation","volume":"223","author":"Dunham","year":"2020","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib23","doi-asserted-by":"crossref","first-page":"23711","DOI":"10.1038\/srep23711","article-title":"Tetrapod-like pelvic girdle in a walking cavefish","volume":"6","author":"Flammang","year":"2016","journal-title":"Sci Rep"},{"key":"2023060902011238200_bib24","doi-asserted-by":"crossref","first-page":"2288","DOI":"10.1242\/jeb.069856","article-title":"How forelimb and hindlimb function changes with incline and perch diameter in the green anole, Anolis carolinensis","volume":"215","author":"Foster","year":"2012","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib26","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1111\/j.1469-7998.2007.00395.x","article-title":"Locomotion of the Eurasian nuthatch on vertical and horizontal substrates","volume":"274","author":"Fujita","year":"2008","journal-title":"J Zool"},{"key":"2023060902011238200_bib27","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1111\/mms.12170","article-title":"Kinematics of terrestrial locomotion in harbor seals and gray seals: importance of spinal flexion by amphibious phocids","volume":"31","author":"Garrett","year":"2015","journal-title":"Mar Mamm Sci"},{"key":"2023060902011238200_bib28","first-page":"e1314298","article-title":"Neither bones nor feet: track morphological variation and \u201cpreservation quality\u201d","volume-title":"J Vertebr Paleontol","author":"Gatesy","year":"2017"},{"key":"2023060902011238200_bib30","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1093\/icb\/ict052","article-title":"Thrash, flip, or jump: the behavioral and functional continuum of terrestrial locomotion in teleost fishes","volume":"53","author":"Gibb","year":"2013","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib31","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1242\/jeb.201.7.949","article-title":"Environmental effects on undulatory locomotion in the American eel Anguilla rostrata: kinematics in water and on land","volume":"201","author":"Gillis","year":"1998","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib32","doi-asserted-by":"crossref","first-page":"208001","DOI":"10.1103\/PhysRevLett.108.208001","article-title":"Entangled granular media","volume":"108","author":"Gravish","year":"2012","journal-title":"Phys Rev Lett"},{"key":"2023060902011238200_bib33","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1111\/j.1469-7998.1960.tb05921.x","article-title":"On the locomotion of the mud-skipper Periophthalmus koelreuteri (pallas): (gobiidae)","volume":"134","author":"Harris","year":"1960","journal-title":"Proc Zool Soc Lond"},{"key":"2023060902011238200_bib34","doi-asserted-by":"crossref","first-page":"e4794","DOI":"10.7717\/peerj.4794","article-title":"A brief introduction to mixed effects modelling and multi-model inference in ecology","volume":"6","author":"Harrison","year":"2018","journal-title":"PeerJ"},{"key":"2023060902011238200_bib35","doi-asserted-by":"crossref","first-page":"034001","DOI":"10.1088\/1748-3182\/3\/3\/034001","article-title":"Software techniques for two- and three-dimensional kinematic measurements of biological and biomimetic systems","volume":"3","author":"Hedrick","year":"2008","journal-title":"Bioinspir Biomim"},{"key":"2023060902011238200_bib36","doi-asserted-by":"crossref","first-page":"e0153446","DOI":"10.1371\/journal.pone.0153446","article-title":"Flapping before flight: high resolution, three-dimensional skeletal kinematics of wings and legs during avian development","volume":"11","author":"Heers","year":"2016","journal-title":"PLoS One"},{"key":"2023060902011238200_bib37","doi-asserted-by":"crossref","first-page":"3599","DOI":"10.1242\/jeb.090027","article-title":"The effect of substrate diameter and incline on locomotion in an arboreal frog","volume":"216","author":"Herrel","year":"2013","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib38","doi-asserted-by":"crossref","first-page":"860","DOI":"10.1016\/j.tree.2021.05.009","article-title":"Linking ecomechanical models and functional traits to understand phenotypic diversity","volume":"36","author":"Higham","year":"2021","journal-title":"Trends Ecol Evol"},{"key":"2023060902011238200_bib39","doi-asserted-by":"crossref","first-page":"38","DOI":"10.4159\/harvard.9780674184404.c3","article-title":"Walking and running","volume-title":"Functional Vertebrate Morphology","author":"Hildebrand","year":"1985"},{"key":"2023060902011238200_bib41","doi-asserted-by":"crossref","first-page":"e11197","DOI":"10.1371\/journal.pone.0011197","article-title":"A locomotor innovation enables water-land transition in a marine fish","volume":"5","author":"Hsieh","year":"2010","journal-title":"PLoS One"},{"key":"2023060902011238200_bib40","doi-asserted-by":"crossref","first-page":"4363","DOI":"10.1242\/jeb.00679","article-title":"Three-dimensional hindlimb kinematics of water running in the plumed basilisk lizard (Basiliscus plumifrons)","volume":"206","author":"Hsieh","year":"2003","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib42","doi-asserted-by":"crossref","DOI":"10.1201\/9781315119861","volume-title":"Fishes out of water: Biology and ecology of mudskippers","author":"Jaafar","year":"2017"},{"key":"2023060902011238200_bib43","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1103\/RevModPhys.68.1259","article-title":"Granular solids, liquids, and gases","volume":"68","author":"Jaeger","year":"1996","journal-title":"Rev Mod Phys"},{"key":"2023060902011238200_bib45","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1093\/icb\/ict034","article-title":"Atmospheric oxygen levels affect mudskipper terrestrial performance: implications for early tetrapods","volume":"53","author":"Jew","year":"2013","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib46","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1111\/jzo.12473","article-title":"Arboreality in acacia rats (Thallomys paedulcus; Rodentia, Muridae): gaits and gait metrics","volume":"303","author":"Karantanis","year":"2017","journal-title":"J Zool"},{"key":"2023060902011238200_bib47","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1093\/icb\/ict051","article-title":"Propulsive forces of mudskipper fins and salamander limbs during terrestrial locomotion: implications for the invasion of land","volume":"53","author":"Kawano","year":"2013","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib48","doi-asserted-by":"crossref","first-page":"jeb.242795","DOI":"10.1242\/jeb.242795","article-title":"Terrestrial force production by the limbs of a semi-aquatic salamander provides insight into the evolution of terrestrial locomotor mechanics","volume":"225","author":"Kawano","year":"2022","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib50","doi-asserted-by":"crossref","first-page":"4154","DOI":"10.1242\/jeb.02493","article-title":"Locomotor kinetics and kinematics on inclines and declines in the gray short-tailed opossum Monodelphis domestica","volume":"209","author":"Lammers","year":"2006","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib51","doi-asserted-by":"crossref","first-page":"215824401882038","DOI":"10.1177\/2158244018820380","article-title":"Model misspecification and assumption violations with the linear mixed model: a meta-analysis","volume":"8","author":"LeBeau","year":"2018","journal-title":"SAGE Open"},{"key":"2023060902011238200_bib52","doi-asserted-by":"crossref","first-page":"2071","DOI":"10.1242\/jeb.201.13.2071","article-title":"Mechanics and energetics of human locomotion on sand","volume":"201","author":"Lejeune","year":"1998","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib53","article-title":"Emmeans: estimated marginal means, aka least-squares means","author":"Lenth","year":"2022"},{"key":"2023060902011238200_bib54","first-page":"3293","article-title":"Multi-functional foot use during running in the zebra-tailed lizard (Callisaurus draconoides)","volume":"215","author":"Li","year":"2012","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib55","doi-asserted-by":"crossref","first-page":"1408","DOI":"10.1126\/science.1229163","article-title":"A terradynamics of legged locomotion on granular media","volume":"339","author":"Li","year":"2013","journal-title":"Science"},{"key":"2023060902011238200_bib56","doi-asserted-by":"crossref","first-page":"jeb242395","DOI":"10.1242\/jeb.242395","article-title":"Patterns and processes in amphibious fish: biomechanics and neural control of fish terrestrial locomotion","volume":"225","author":"Lutek","year":"2022","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib57","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1126\/science.aaf0984","article-title":"Tail use improves performance on soft substrates in models of early vertebrate land locomotors","volume":"353","author":"McInroe","year":"2016","journal-title":"Science"},{"key":"2023060902011238200_bib58","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1111\/jzo.12794","article-title":"Terrestrial locomotion in elongate fishes: exploring the roles of morphology and substrate in facilitating locomotion","volume":"315","author":"Mehta","year":"2020","journal-title":"J Zool"},{"key":"2023060902011238200_bib59","doi-asserted-by":"crossref","DOI":"10.3853\/j.0812-7387.11.1989.93","volume-title":"A taxonomic revision and cladistic analysis of the oxudercine gobies (Gobiidae: Oxudercinae)","author":"Murdy","year":"1989"},{"key":"2023060902011238200_bib60","first-page":"99","article-title":"Systematics of Oxudercinae","volume-title":"The biology of gobies","author":"Murdy","year":"2011"},{"key":"2023060902011238200_bib62","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1111\/j.2041-210x.2012.00261.x","article-title":"A general and simple method for obtaining R2 from generalized linear mixed-effects models","volume":"4","author":"Nakagawa","year":"2013","journal-title":"Methods Ecol Evol"},{"key":"2023060902011238200_bib63","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1111\/1365-2435.13969","article-title":"High-speed terrestrial substrate transitions: how a fleeing cursorial day gecko copes with compliance changes that are experienced in nature","volume":"36","author":"Naylor","year":"2022","journal-title":"Funct Ecol"},{"key":"2023060902011238200_bib64","doi-asserted-by":"crossref","first-page":"jeb.211706","DOI":"10.1242\/jeb.211706","article-title":"Cling performance and surface area of attachment in plethodontid salamanders","author":"O'Donnell","year":"2020","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib65","doi-asserted-by":"crossref","first-page":"1747","DOI":"10.1111\/evo.12971","article-title":"Repeated evolution of amphibious behavior in fish and its implications for the colonization of novel environments","volume":"70","author":"Ord","year":"2016","journal-title":"Evolution"},{"key":"2023060902011238200_bib66","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1007\/978-3-319-63537-8_55","article-title":"Geometric mechanics applied to tetrapod locomotion on granular media","volume-title":"Biomimetic and Biohybrid Systems","author":"Ozkan\u00a0Aydin","year":"2017"},{"key":"2023060902011238200_bib67","doi-asserted-by":"crossref","first-page":"2279","DOI":"10.1242\/jeb.029041","article-title":"Mudskipper pectoral fin kinematics in aquatic and terrestrial environments","volume":"212","author":"Pace","year":"2009","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib68","doi-asserted-by":"crossref","first-page":"530","DOI":"10.1242\/jeb.047902","article-title":"Locomotor behavior across an environmental transition in the ropefish, Erpetoichthys calabaricus","volume":"214","author":"Pace","year":"2011","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib69","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1111\/jfb.12318","article-title":"Sustained periodic terrestrial locomotion in air-breathing fishes","volume":"84","author":"Pace","year":"2014","journal-title":"J Fish Biol"},{"key":"2023060902011238200_bib70","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1002\/jez.2594","article-title":"Kinematic comparisons between mudskipper fins and salamander limbs during terrestrial locomotion","volume":"337","author":"Quigley","year":"2022","journal-title":"J Exp Zool A Ecol Integr Physiol"},{"key":"2023060902011238200_bib71","volume-title":"R: A language and environment for statistical computing Vienna","author":"R Core Team","year":"2020"},{"key":"2023060902011238200_bib72","volume-title":"RStudio: Integrated Development for R RStudio, PBC","author":"R Studio Team","year":"2021"},{"key":"2023060902011238200_bib73","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1093\/icb\/icaa016","article-title":"Terrestrial locomotion in American eels (Anguilla rostrata): how substrate and incline affect movement patterns","volume":"60","author":"Redmann","year":"2020","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib74","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1111\/j.1365-2435.2011.01943.x","article-title":"The effect of natural substrates on jump height in click-beetles","volume":"26","author":"Ribak","year":"2012","journal-title":"Funct Ecol"},{"key":"2023060902011238200_bib75","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1111\/j.1467-2979.2005.00193.x","article-title":"Adaptations of amphibious fish for surviving life out of water","volume":"6","author":"Sayer","year":"2005","journal-title":"Fish Fish"},{"key":"2023060902011238200_bib76","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat Methods"},{"key":"2023060902011238200_bib77","doi-asserted-by":"crossref","first-page":"2544","DOI":"10.1242\/jeb.051086","article-title":"The kinematic consequences of locomotion on sloped arboreal substrates in a generalized (Rattus norvegicus) and a specialized (Sciurus vulgaris) rodent","volume":"214","author":"Schmidt","year":"2011","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib95_1655760866177","first-page":"icac047","article-title":"The spring-mass model and other reductionist models of bipedal locomotion on inclines","author":"Selvitella","year":"2022","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib78","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1002\/jez.2020","article-title":"Effects of substrate size and orientation on quadrupedal gait kinematics in Mouse Lemurs (Microcebus murinus)","volume":"325","author":"Shapiro","year":"2016","journal-title":"J Exp Zool A Ecol Genet Physiol"},{"key":"2023060902011238200_bib79","doi-asserted-by":"crossref","first-page":"046009","DOI":"10.1088\/1478-3975\/12\/4\/046009","article-title":"Controlled preparation of wet granular media reveals limits to lizard burial ability","volume":"12","author":"Sharpe","year":"2015","journal-title":"Phys Biol"},{"key":"2023060902011238200_bib80","first-page":"R1","article-title":"A guerilla guide to common problems in \u2018neurostatistics\u2019: essential statistical topics in neuroscience","volume":"16","author":"Smith","year":"2017","journal-title":"J Undergrad Neurosci Educ"},{"key":"2023060902011238200_bib81","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.zool.2016.05.001","article-title":"Locomotor flexibility of Polypterus senegalus across various aquatic and terrestrial substrates","volume":"119","author":"Standen","year":"2016","journal-title":"Zoology"},{"key":"2023060902011238200_bib82","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1038\/nature13708","article-title":"Developmental plasticity and the origin of tetrapods","volume":"513","author":"Standen","year":"2014","journal-title":"Nature"},{"key":"2023060902011238200_bib83","doi-asserted-by":"crossref","first-page":"18","DOI":"10.2307\/1440166","article-title":"Observations on the natural history of the mud-skipper, Periophthalmus sobrinus","volume":"1961","author":"Stebbins","year":"1961","journal-title":"Copeia"},{"key":"2023060902011238200_bib84","article-title":"Do we all walk the walk? A comparison of walking behaviors across tetrapods","author":"Struble","year":"","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib86","doi-asserted-by":"crossref","first-page":"4037","DOI":"10.1242\/jeb.01237","article-title":"Kinematics of aquatic and terrestrial escape responses in mudskippers","volume":"207","author":"Swanson","year":"2004","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib87","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1152\/physiol.00012.2021","article-title":"More than breathing air: evolutionary drivers and physiological implications of an amphibious lifestyle in fishes","volume":"36","author":"Turko","year":"2021","journal-title":"Physiology"},{"key":"2023060902011238200_bib88","first-page":"icac009","article-title":"What is stance phase on deformable substrates?","author":"Turner","year":"2022","journal-title":"Integr Comp Biol"},{"key":"2023060902011238200_bib89","doi-asserted-by":"crossref","first-page":"obaa029","DOI":"10.1093\/iob\/obaa029","article-title":"Tiger salamanders (Ambystoma tigrinum) increase foot contact surface area on challenging substrates during terrestrial locomotion","volume":"2","author":"Vega","year":"2020","journal-title":"Integr Org Biol"},{"key":"2023060902011238200_bib90","doi-asserted-by":"crossref","first-page":"3523","DOI":"10.1242\/jeb.007187","article-title":"Stereotypy, flexibility and coordination: key concepts in behavioral functional morphology","volume":"211","author":"Wainwright","year":"2008","journal-title":"J Exp Biol"},{"key":"2023060902011238200_bib91","doi-asserted-by":"crossref","DOI":"10.1109\/ROBIO.2013.6739610","article-title":"Experimental study on morphology and kinematics of mudskipper in amphibious environments","volume-title":"Presented at the 2013 IEEE International Conference on Robotics and Biomimetics (ROBIO), Shenzhen, China","author":"Wang","year":"2013"},{"key":"2023060902011238200_bib92","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1016\/j.zool.2016.06.007","article-title":"The significance of pelvic fin flexibility for tree climbing fish","volume":"119","author":"Wicaksono","year":"2016","journal-title":"Zoology"},{"key":"2023060902011238200_bib93","doi-asserted-by":"crossref","first-page":"2121","DOI":"10.1017\/S0025315417001722","article-title":"A mechanical piston action may assist pelvic\u2013pectoral fin antagonism in tree-climbing fish","volume":"98","author":"Wicaksono","year":"2018","journal-title":"J Mar Biol Assoc UK"},{"key":"2023060902011238200_bib94","doi-asserted-by":"crossref","first-page":"2472","DOI":"10.1111\/1365-2435.13919","article-title":"Clinging performance on natural substrates predicts habitat use in anoles and geckos","volume":"35","author":"Wright","year":"2021","journal-title":"Funct Ecol"},{"key":"2023060902011238200_bib95","first-page":"345","article-title":"Morphological adaptations to special environments of gobies","volume-title":"The biology of gobies","author":"Zander","year":"2011"},{"key":"2023060902011238200_bib96","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1177\/0278364913481690","article-title":"Ground fluidization promotes rapid running of a lightweight robot","volume":"32","author":"Zhang","year":"2013","journal-title":"Int J Rob Res"}],"container-title":["Integrative And Comparative Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/icb\/advance-article-pdf\/doi\/10.1093\/icb\/icac084\/44329145\/icac084.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/icb\/article-pdf\/62\/5\/1335\/47612544\/icac084.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/icb\/article-pdf\/62\/5\/1335\/47612544\/icac084.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,6,9]],"date-time":"2023-06-09T02:01:50Z","timestamp":1686276110000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/icb\/article\/62\/5\/1335\/6604839"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,6,9]]},"references-count":93,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2022,6,9]]},"published-print":{"date-parts":[[2022,12,6]]}},"URL":"https:\/\/doi.org\/10.1093\/icb\/icac084","relation":{},"ISSN":["1540-7063","1557-7023"],"issn-type":[{"value":"1540-7063","type":"print"},{"value":"1557-7023","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2022,11]]},"published":{"date-parts":[[2022,6,9]]}}}