Thesis
These fish were made for walking: morphology and walking kinematics in balitorid loaches
Crawford, C.H.
Record Number:
5794
Year:
2021
University:
New Jersey Institute of Technology and Rutgers, The State University of New Jersey, Doctor of Philosophy
Abstract:
Terrestrial excursions have been observed in multiple lineages of marine and freshwater
fishes. These ventures into the terrestrial environment may be used when fish are
searching out new habitat during drought, escaping predation, laying eggs, or seeking
food sources. The physiological demands for life under water and on land are vastly
different and require different functional adaptations. Fish with terrestrial excursions
must be capable of dealing with the stresses of both aquatic and terrestrial environments
for varying periods of time. To deal with these stresses, amphibious fishes exhibit many
morphological and behavioral adaptations. These adaptations have led to a range of
locomotor strategies when traversing the terrestrial environment. Importantly, a broader
understanding of the ecomorphology and biomechanics of terrestrial excursions in living
fishes will aid in interpretation of both fossil fish and early tetrapod anatomy and
trackways.
The rheophilic hillstream loaches (Balitoridae) possess a pelvic morphology
which are attributed to adaptations for life in rapidly flowing water. The unique
connectivity of the pelvic plate to the vertebral column via a sacral rib, and the relative
size and shape of the sacral rib, fall within a spectrum of three discrete morphotypes.
These morphotypes, determined through skeletal morphology and compared
phylogenomically, are correlated with patterns observed in the pelvic muscle morphology of these fishes and are expected to provide a mechanical advantage for generating force
against the ground.
The skeletal connection via the sacral rib in balitorid loaches is hypothesized to
facilitate terrestrial locomotion observed in the family. Field and laboratory-collected
high-speed video is used to analyze terrestrial walking kinematics in seven balitorid
species representing both subfamilies and two of the three morphotypes. Contrary to the
hypothesis that robustness of the sacral rib would strongly influence walking
performance, there is not a large reduction in walking ability in the Morphotype 1
representative. Major differences in walking kinematics distinguish the two balitorid
subfamilies; with a generally greater walking performance in Balitorinae and reduced
capability in the Homalopteroidinae representatives. The connection between internal
anatomy and locomotion on land are explored with digitized video analysis, μCT scans,
and in the context of the phylogenetic history of this family of fishes.
The unexpected result of Homaloptera parclitella being capable of walking with
comparable performance to other balitorids prompted further exploration into the walking
mechanics in this species. To assess changes in terrestrial walking gait, walking
kinematics are explored at experimental inclines (0˚, 15˚, 30˚, and 45˚) along with
electromyography (EMG) during flat walking. H. parclitella is capable of walking at
inclines without much change to the distance traveled over time. Additionally, another
walking behavior is described; termed high walking, where the fish lifts its entire body
off the substrate and carries its mass on its fins.
Times Cited:
1
Relevent Species:
Related Records:
Kottelat, M. (1988)
Two species of cavefishes from northern Thailand in the genera Nemachilus and Homaloptera (Osteichthyes, Homalopteridae)
Kottelat, M. (1998)Homaloptera yuwonoi, a new species of hillstream loach from Borneo, with a new generic name for H. thamicola (Teleostei: Balitoridae)
Deharveng, L. and Bedos, A. (2001)Thailande
Borowsky, R.L. and Vidthayanon, C. (2001)Nucleotide diversity in populations of balitorid cave fishes from Thailand
Trajano, E., Mugue, N., Krejca, J., Vidthayanon, C., Smart, D. and Borowsky, R. (2002)Habitat, distribution, ecology and behaviour of cave balitorids from Thailand (Teleostei: Cypriniformes)
Prokofiev, A.M. (2010)Morphological classification of loaches (Nemacheilinae)
Kottelat, M. (2012)Conspectus cobitidum: An inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei)
Brancelj, A., Boonyanusith, C., Watiroyram, S. and Sanoamuang, L. (2013)The groundwater-dwelling fauna of Southeast Asia
Lujan, N.K. and Conway, K.W. (2015)Life in the fast lane: A review of rheophily in freshwater fishes
Sherwin, F. (2016)Wall-climbing cave fish: Evolutionary intermediate?
Zimmer, C. (2016)Researchers find fish that walks the way land Vertebrates do
Flammang, B.E., Suvarnaraksha, A., Markiewicz, J. and Soares, D. (2016)Tetrapod-like pelvic girdle in a walking cavefish
Anonymous (2018)New walking cavefish study explores origins of quadrapedal walking
Anonymous (2018)First steps. Scientists launch evolutionary study to explore the origins of fish that walk
Anonymous (2018)New walking cavefish study explores origins of quadrapedal walking
Ellis, M. (2018)The recorded fauna of the limestone caves of Thailand to April 2018
Willis, J., Burt de Perera, T., Newport, C., Poncelet, G., Sturr, C.J. and Thomas, A. (2019)The structure and function of the sucker systems of hill stream loaches
Liew, J.H., Chua, K.W.J., Arsenault, E.R, Thorp, J.H., Suvarnaraksha, A., Amirrudin, A. and Yeo, D.C.J. (2019)Quantifying terrestrial carbon in freshwater food webs using amino acid isotope analysis: Case study with an endemic cavefish
Zhongming, Z., Linong, L., Xiaona, Y., Wangqiang, Z. and Wei, L. (2020)Study of Asia's hillstream loaches reveals keys to fish family's land-walking abilities
Crawford, C.H., Randall, Z.S., Hart, P.B., Page, L.M., Chakrabarty, P., Suvarnaraksha, A. and Flammang, B.E. (2020)Skeletal and muscular pelvic morphology of hillstream loaches (Cypriniformes: Balitoridae)
Ellis, M. (2020)The caves of northern Thailand
Crawford, C.H. (2021)These fish were made for walking: morphology and walking kinematics in balitorid loaches
Crawford, C.H., Webber-Schultz, A., Hart, P.B., Randall, Z.S., Cerrato-Morales, C., Kellogg, A.B., Amplo, H.E., Suvarnaraksha, A., Page, L.M., Chakrabarty, P. and Flammang, B.E. (2022)They like to move it (move it): walking kinematics of balitorid loaches of Thailand
Flammang, B.E. (2022)Bioinspired design in research: Evolution as beta-testing