Abstract: Equid evolution is characterized by a high diversity of extinct species and morphologies, whereas extant equids share a superficially similar, monodactyl morphology. This inferred musculoskeletal similarity of modern equid limbs remains unexplored, and it is often assumed that domestic horse limbs are representative for wild equids (e.g., zebras, onagers, etc.). Our aim was to quantitatively describe the muscle architecture and arrangement of all forelimb muscles in extant wild Equus species to test this assumption, and investigate any differences between the species. We hypothesized that there would be subtle variation linked to locomotion on the different substrates that these species encounter. Gross dissections were performed to record muscle attachment sites and to quantify architectural metrics: muscle-tendon unit (MTU) length, MTU mass, muscle mass, pennation angle, and fascicle length; physiological cross-sectional area (PCSA) of the muscles and tendon cross-sectional area (TCSA) of the distal ligaments were then calculated. Qualitative results confirm common origin and insertion sites of all muscles across all Equus species. When normalized for size, the forelimb muscles across equids generally exhibit comparable muscular architecture and force-generating capacities. However, we observed a trend for higher force-generating potential in the distal limb flexor muscles in two species of zebra naturally found in habitats with inclined or uneven substrates. Although limited by sample size, these results indicate that scaled data for modern wild equids are generally very comparable, which may enable much smoother translation of experimental data from domestic horses into digital simulations of wild equid locomotion, including for extinct equids.
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.
Overview
This study investigates the muscle structure of the forelimbs in wild horse species (Equus genus) to determine whether domestic horse limb data accurately represent wild equid muscular anatomy and function.
The research compares the muscle architecture of different wild equid species to identify potential adaptations linked to their natural environments and locomotion.
Introduction and Background
Equids (members of the horse family) have a rich evolutionary history with many extinct species that exhibited diverse limb morphologies.
Modern equids (like horses, zebras, and onagers) share a similar monodactyl (single-toed) limb structure, but it is not clear if this similarity extends to the underlying muscle architecture.
Typically, domestic horse limb anatomy is used as a reference for wild equids, yet this assumption has not been extensively tested.
The study aims to quantitatively assess the forelimb muscles of wild Equus species to clarify if domestic horse data are representative for the group.
Researchers hypothesize subtle differences in muscle characteristics among species that relate to their natural habitats and locomotion styles, especially since wild equids move on varied terrains.
Methods
Gross dissections were performed on forelimbs of multiple extant wild Equus species (examples include different types of zebras and onagers).
Muscle attachment sites were documented to verify common origins and insertions across species.
Architectural metrics measured included:
Muscle-tendon unit (MTU) length
MTU mass and muscle mass
Pennation angle (angle of muscle fibers relative to the tendon)
Fascicle length (length of bundles of muscle fibers)
Physiological cross-sectional area (PCSA) was calculated to estimate muscles’ force-generating capacity.
Tendon cross-sectional area (TCSA) of the distal ligaments was also measured.
All metrics were normalized to body size to allow fair comparison between different species.
Key Findings
Qualitative observations confirmed that all studied wild Equus species share similar muscle attachment points in their forelimbs, indicating a conserved musculoskeletal design.
When size differences were accounted for, the overall architectural traits of forelimb muscles were largely comparable across all species.
A noticeable trend was found in two zebra species that live on inclined or uneven terrains:
They exhibited higher force-generating potential, specifically in the distal limb flexor muscles (those responsible for bending the limb).
This suggests some level of muscular adaptation linked to the challenging substrate these zebras navigate regularly.
Despite some subtle differences, the general muscular architecture is fairly uniform among wild Equus species.
Implications and Significance
The study supports the common practice of using domestic horse limb data as a proxy for wild equids in biomechanical research and modeling.
This finding facilitates the application of domestic horse experimental data to digital simulations of wild equid locomotion, improving our understanding of how these animals move.
Insights gained may help reconstruct locomotor function in extinct equid species by extrapolation from modern analogs.
The observed minor muscular differences tied to habitat suggest that evolution fine-tunes muscle architecture to environmental demands but does so within a largely conserved framework.
Limitations and Future Directions
The study acknowledges a limited sample size, which may restrict the detection of subtle or species-specific differences.
Future research could include a larger number of specimens and species to better assess variability and confirm preliminary findings.
Additional studies could explore hindlimb muscle architecture and other anatomical features to provide a more comprehensive view.
Investigations incorporating locomotion kinematics and muscle function in different substrates would further clarify functional adaptations.
Cite This Article
APA
De Ridder T, Aerts P, MacLaren JA.
(2025).
Could “wild horses” drag you away? Quantifying muscular architecture in the forelimbs of extant, non-domestic equids (Perissodactyla: Equidae).
Anat Rec (Hoboken).
https://doi.org/10.1002/ar.70092
Department of Biology, Universiteit Antwerpen, Antwerp, Belgium.
Aerts, Peter
Department of Biology, Universiteit Antwerpen, Antwerp, Belgium.
Movement and Sports Sciences, University of Ghent, Ghent, Belgium.
MacLaren, Jamie A
Department of Biology, Universiteit Antwerpen, Antwerp, Belgium.
Evolution & Diversity Dynamics Lab, UR Geology, Université de Liège, Liège, Belgium.
Palaeobiosphere Evolution Lab, Institute of Natural Sciences, Brussels, Belgium.
Grant Funding
Fonds Wetenschappelijk Onderzoek
University Research Fund (BOF-UA) of the University of Antwerp
References
This article includes 89 references
Andrew SM, Totland Ø, Moe SR. Spatial variation in plant species richness and diversity along human disturbance and environmental gradients in a tropical wetland. Wetlands Ecology and Management 23(3), 395–404.
Bates K, Maidment SCR, Schachner ER, Barrett PM. Comments and corrections on 3D modeling studies of locomotor muscle moment arms in archosaurs. PeerJ 3, e1272.
Bishop PJ, Michel KB, Falisse A, Cuff AR, Allen VR, de Groote F, Hutchinson JR. Computational modelling of muscle fibre operating ranges in the hindlimb of a small ground bird (Eudromia elegans), with implications for modelling locomotion in extinct species. PLoS Computational Biology 17(4), e1008843.
Böhmer C, Theil J, Fabre A, Herrel A. Atlas of Terrestrial Mammal Limbs. .
Brassey CA, Maidment SCR, Barrett PM. Muscle moment arm analyses applied to vertebrate paleontology: A case study using stegosaurus stenops marsh, 1887. Journal of Vertebrate Paleontology 37(5), e1361432.
Carnicer D, Coudry V, Denoix J. Ultrasonographic examination of the palmar aspect of the pastern of the horse: Sesamoidean ligaments. Equine Veterinary Education 25, 256–263.
Charles JP, Cappellari O, Hutchinson JR. A dynamic simulation of musculoskeletal function in the mouse hindlimb during trotting locomotion. Frontiers in Bioengineering and Biotechnology 6, 61.
Cuff AR, Demuth OE, Michel K, Otero A, Pintore R, Polet DT, Wiseman ALA, Hutchinson JR. Walking—And running and jumping—With dinosaurs and their cousins, viewed through the lens of evolutionary biomechanics. Integrative and Comparative Biology 62(5), 1281–1305.
Demuth OE, Herbst E, Polet DT, Wiseman ALA, Hutchinson JR. Modern three‐dimensional digital methods for studying locomotor biomechanics in tetrapods. Journal of Experimental Biology 226, jeb245132.
Demuth OE, Wiseman ALA, van Beesel J, Mallison H, Hutchinson JR. Three‐dimensional polygonal muscle modelling and line of action estimation in living and extinct taxa. Scientific Reports 12(1), 3358.
Der Sarkissian C, Vilstrup JT, Schubert M, Seguin‐Orlando A, Eme D, Weinstock J, Alberdi MT, Martin F, Lopez PM, Prado JL, Prieto A, Douady CJ, Stafford TW, Willerslev E, Orlando L. Mitochondrial genomes reveal the extinct Hippidion as an outgroup to all living equids. Biology Letters 11(3), 20141058.
Eisenmann V, Beckouche S. Identification and Discrimination of Metapodials from Pleistocene and Modern Equus, Wild and Domesstic. In R. H. Meadow & H.‐P. Uerpmann (Eds.), Equids in the ancient world (pp. 67–163).
Eisenmann V, Karchoud A. Analyses multidimensionnelles des metapodes d'Equus. Bulletin du Museum National d'Histoire Naturelle 4, 75–103.
Etienne C, Houssaye A, Hutchinson JR. Limb myology and muscle architecture of the Indian rhinoceros Rhinoceros unicornis and the white rhinoceros Ceratotherium simum (Mammalia: Rhinocerotidae). PeerJ 9, e11314.
Feh C, Munkhtuya B, Enkhbold S, Sukhbaatar T. Ecology and social structure of the Gobi khulan Equus hemionus subsp. in the Gobi B National Park, Mongolia. Biological Conservation 101(1), 51–61.
Gaunitz C, Fages A, Hanghøj K, Albrechtsen A, Khan N, Schubert M, Seguin‐Orlando A, Owens IJ, Felkel S, Bignon‐Lau O, de Barros Damgaard P, Mittnik A, Mohaseb AF, Davoudi H, Alquraishi S, Alfarhan AH, Al‐Rasheid KAS, Crubézy E, Benecke N, Orlando L. Ancient genomes revisit the ancestry of domestic and Przewalski's horses. Science 360(6384), 111–114.
Harbers H, Zanolli C, Cazenave M, Theil J‐C, Ortiz K, Blanc B, Locatelli Y, Schafberg R, Lecompte F, Baly I, Laurens F, Callou C, Herrel A, Puymerail L, Cucchi T. Investigating the impact of captivity and domestication on limb bone cortical morphology: An experimental approach using a wild boar model. Scientific Reports 10, 19070.
Hermanson JW, MacFadden BJ. Evolutionary and functional morphology of the shoulder region and stay‐apparatus in fossil and extant horses (Equidae). Journal of Vertebrate Paleontology 12(3), 377–386.
Herrel A, Locatelli Y, Ortiz K, Theil J, Cornette R, Cucchi T. Cranial muscle architecture in wild boar: Does captivity drive ontogenetic trajectories?. Journal of Morphology 285(2), e21676.
Hutchinson JR, Anderson FC, Blemker SS, Delp SL. Analysis of hindlimb muscle moment arms in Tyrannosaurus rex using a three‐dimensional musculoskeletal computer model: Implications for stance, gait, and speed. Paleobiology 31(4), 676.
Jónsson H, Schubert M, Seguin‐Orlando A, Ginolhac A, Petersen L, Fumagalli M, Albrechtsen A, Petersen B, Korneliussen TS, Vilstrup JT, Lear T, Myka JL, Lundquist J, Miller DC, Alfarhan AH, Alquraishi SA, Al‐Rasheid KAS, Stagegaard J, Strauss G, Orlando L. Speciation with gene flow in equids despite extensive chromosomal plasticity. Proceedings of the National Academy of Sciences of the United States of America 111(52), 18655–18660.
Kaczensky P, Kuehn R, Lhagvasuren B, Pietsch S, Yang W, Walzer C. Connectivity of the Asiatic wild ass population in the Mongolian Gobi. Biological Conservation 144(2), 920–929.
MacLaren JA, Corssmit E, MacMillan M, Rojas‐Jimenez J. A quantitative analysis of the manus musculature in tapirs (Perissodactyla: Tapiridae). Journal of Morphology 286(5), e70051.
Mendez J, Keys A. Density and composition of mammalian muscle. Metabolism 9, 184–188.
Michilsens F, Vereecke EE, D'août K, Aerts P. Functional anatomy of the gibbon forelimb: Adaptations to a brachiating lifestyle. Journal of Anatomy 215(3), 335–354.
Moehlman PD. Status survey and conservation action plan equids: Zebras, asses and horses. In P. D. Moehlman (Ed.).
Neaux D, Blanc B, Ortiz K, Locatelli Y, Schafberg R, Herrel A, Debat V, Cucchi T. Constraints associated with captivity alter craniomandibular integration in wild boar. Journal of Anatomy 239(2), 489–497.
Oakenfull EA, Ryder OA. Genetics of equid species and subspecies. In Equids: Zebras, asses and horses. Status survey and conservation action plan (pp. 108–112).
Otero A, Allen V, Pol D, Hutchinson JR. Forelimb muscle and joint actions in Archosauria: Insights from Crocodylus johnstoni (Pseudosuchia) and Mussaurus patagonicus (Sauropodomorpha). PeerJ 2017(11), e3976.
Pelletier M, Niinimäki S, Salmi AK. Influence of captivity and selection on limb long bone cross‐sectional morphology of reindeer. Journal of Morphology 282(10), 1533–1556.
Richards HL, Adams JW, Evans AR. Hanging on and digging deep: Comparative forelimb myology of the koala (Phascolarctos cinereus) and common wombat (Vombatus ursinus). Zoological Journal of the Linnean Society 199, 60–82.
Schulz E, Kaiser TM. Historical distribution, habitat requirements and feeding ecology of the genus Equus (Perissodactyla). Mammal Review 43(2), 111–123.
Shanungu GK, Kaumba CH, Beilfuss R. Current population status and distribution of large herbivores and floodplain birds of the Kafue flats wetlands, Zambia: Results of the 2015 Wet Season Aerial Survey. .
Tesfai RT, Owen‐Smith N, Parrini F, Moehlman PD. Viability of the critically endangered African wild ass (Equus africanus) population on Messir Plateau (Eritrea). Journal of Mammalogy 100(1), 185–191.
Tidière M, Duncan P, Lemaître J‐F, Gaillard J‐M, Lackey LB, Müller DWH, Clauss M. Do equids live longer than grazing bovids?. Journal of Mammalian Evolution 27(4), 809–816.
Wareing K, Tickle PG, Stokkan KA, Codd JR, Sellers WI. The musculoskeletal anatomy of the reindeer (Rangifer tarandus): Fore‐ and hindlimb. Polar Biology 34(10), 1571–1578.
Witmer LM. The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils. In J. Thomason (Ed.), Functional morphology in vertebrate paleontology (pp. 19–33).