Enhanced Forelimb Mobility in Insular Meles populations: Insights from Myological and Osteological Comparisons

Enhanced Forelimb Mobility in Insular Meles populations: Insights from Myological and Osteological Comparisons | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Enhanced Forelimb Mobility in Insular Meles populations: Insights from Myological and Osteological Comparisons Emma Dangerfield, Mao Shimoda, Sujoo Cho, Yuri Kimura This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7041610/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 02 May, 2026 Read the published version in Journal of Mammalian Evolution → Version 1 posted 11 You are reading this latest preprint version Abstract Island colonization can drive anatomical adaptations in mammals, yet few studies have examined how such changes occur in limb musculature and osteology. Here, we investigate the forelimb anatomy of Meles badgers to understand functional adaptations associated with insular environments. We conducted detailed myological dissections of the Japanese badger ( Meles anakuma ) and its continental relative ( Meles leucurus ), and compared osteological features with another continental-insular pair: Meles meles from mainland Europe and Meles canescens from Crete. Our results reveal several previously undescribed features, including the presence of m. tensor fasciae antebrachii in Meles , and a distal shift in the main belly of m. coracobrachialis in M. anakuma , both with considerable phylogenetic ramifications. While body size did not differ significantly between M. anakuma and M. leucurus , insular populations of both M. anakuma and M. canescens exhibit shared osteological traits, such as reduced projection of the humeral trochlea and enlargement of the cranial aspect of the ulnar head, which likely enhance forelimb mobility rather than maximize digging force. These results may indicate that insular Meles have evolved increased joint flexibility, potentially as an adaptation for locomoter versatility in more structurally complex or variable island habitats. Our findings establish Meles as a valuable model for studying evolutionary responses to island environments in subfossorial mesocarnivores. Comparative osteology forelimb myology Meles insular populations island adaptations Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Introduction The genus Meles , comprising of four extant species, Meles meles Linnaeus, 1758 (European badger), Meles leucurus Hodgson, 1847 (Asian badger), Meles anakuma Temminck, 1844 (Japanese badger), and the more recently redefined Meles canescens Blanford, 1875 (Caucasian badger), is widely distributed across Eurasia, from Europe to Eastern Asia and the Japanese Archipelago (Fig. 1 ) (Neal and Cheeseman 1996 ; Del Carro et al. 2010; Abramov and Puzachenko 2013 ). While the evolutionary history of these taxa have been documented through mitochondrial DNA and cranial morphology (Marmi et al. 2006 ; Koh et al. 2014 ; Lee et al. 2016 ; Law et al. 2018 ; Faggi et al. 2024 ), much less is known about how island colonization has shaped the functional morphology of their forelimbs, particularly in insular populations such as M. anakuma in Japan. Island environments are known to exert unique selective pressures on mammals, often resulting in shifts in body size. According to the ‘Island Rule’, large mammals tend to undergo dwarfism, while small mammals exhibit gigantism upon colonizing islands (Foster 1964 ; Meiri et al. 2004 ; Hayashi et al. 2023 ; Rozzi et al. 2023 ). However, this pattern is often less pronounced or predictable for medium-sized mesocarnivores (Meiri et al. 2004 ; Benítez-López et al. 2021 ). For instance, gigantism is evident in the island subspecies of Minorcan pine marten ( Martes martes minoricensis ) (López-Martín et al. 2006 ), in some insular otters (e.g. Megalenhydris barbaricina ) (Lyras et al. 2010 ) and the Arctic fox ( Vulpes lagopus ) (Nanova and Prôa 2017 ), while dwarfism was recorded for the island populations of the pygmy raccoon ( Procyon pygmaeus ) (Mcfaddon and Meiri 2012), the raccoon dog ( Nyctereutes procionoides ) (Kim et al. 2015 ), the leopard cat ( Prionailurus bengalensi ) (Sicuro and Oliveira 2015 ), and several fox species (e.g Urocyon littoralis ) (Moore and Collins 1995 ). Beyond body size, insular environments often influence behavior, ecology, and functional morphology. For example, field observations show the Channel Islands foxes have tendencies towards more arboreal activities and increased diurnal behavior likely as a response to ecological restraints (Wayne et al. 1991 ; Moore and Collins 1995 ; Coonan et al. 2010 ), while changes in cranial features in insular artic foxes reflect modified foraging strategies (Nanova and Prôa 2017 ). Similar ecological shifts have been found in Meles where island populations of M. meles and M. leucurus have shown altered diets and behaviors, likely driven by restricted resources (Oh 2007 ; Sleeman et al. 2009 ; Sleeman and Davenport 2016 ). These ecological differences are supported by recent morphological studies which report cranial and dental adaptations in insular Meles tax – including M. canescens from Crete and M. anakuma – such as increased interorbital distance, enhanced masticatory features and greater bite force (Savvidou et al. 2022 ). These traits are associated with shifts towards increased diurnal activity and more carnivorous diets. Nonetheless, these adaptations do not always coincide with significant changes in body size, suggesting that other morphological adjustments may be more crucial for ecological success on islands. Recent research emphasizes the value of studying medium-sized carnivores, including badgers, due to their high adaptability to rapid environmental changes (Marneweck et al. 2022 ). This make Meles a compelling case for investigating how island environments influence morphological adaptations. In particular, given the subfossorial lifestyle of badgers and their reliance on powerful forelimbs for digging and foraging (Moore 2011 ; Hildebrand and Goslow 2001 ), investigating forelimb myology and osteology are key to understanding functional responses to insular ecological pressures. Forelimb adaptations such as enhanced maneuverability or increased force production may compensate for the absence of major body size shifts, providing a functional strategy for unique ecological pressures in insular settings. Here, we provide the first detailed comparative myological analysis of the forelimbs in the insular M. anakuma and its continental counterpart M. leucurus , complemented by osteological comparisons with the European M. meles and an insular population of M. canescens from the Island of Crete. By integrating these anatomical observations, we aim to identify key musculoskeletal adaptations that have evolved in response to island-specific ecological demands. Materials and Methods Dissections We believe that no myological studies have been published on either M. anakuma , a badger species endemic to Japan, or on any M. leucurus individuals inhabiting South Korea. In this study, we dissected the forelimbs of four M. anakuma specimens and three M. leucurus specimens and analyzed their respective muscular structures (Table 1 ). All specimens were procured as roadkill victims and dissections were conducted at the NMNS Research center in Tsukuba for M. anakuma , and at both Seoul National University (SNU) and the Wildlife Rescue Center in Ulsan City for M. leucurus . For each badger specimen, the fur, organs and fatty tissues were carefully removed with scalpels and surgical scissors. Then the origin, insertion and morphological features of 28 forelimb intrinsic muscles were systematically identified and described (Table 2 ). Photographs were taken with a Canon EOS Kiss X9i (Canon Inc., Japan) to document the dissection process. Anatomical and osteological descriptions were based on terminology from Fishbeck and Sebastiani ( 2001 ), Evans et al. (2012), Ercoli et al. ( 2015 )d hmer et al. (2020). Individual muscles were traced over dissection photographs in the program Procreate version 5.2.8 (Savage Interactive Pty Ltd., Australia), then separated first by color gradients for each muscle group (shoulder, brachium, distal arm, and deep) and then by patterns for the individual muscle heads. Table 1 List of Meles specimens with locality, sex, and humerus length. Abbreviations: F , female; M , male; NA , not available. For museum abbreviations, see the main text Museum/ Institute Specimen ID ID No. for this study Meles species Locality Sex Humerus length (mm) NMNS POM-256 1 M. anakuma Fukushima, Japan NA 95.55 NMNS POM-259 2 M. anakuma Hiroshima, Japan M 94.56 NMNS POM-258 3 M. anakuma Kumamoto, Japan F 75.85 NMNS POM-260 4 M. anakuma Kagoshima, Japan NA 98.59 SNU NA 5 M. leucurus Seoul, South Korea F 86.09 SNU NA 6 M. leucurus Seoul, South Korea M 97.76 SNU NA 7 M. leucurus Ulsan, South Korea M 92.31 NHMC 80.5.63.19 - M. canescens Crete, Greece NA 84.57 NHMC 80.5.63.43 - M. canescens Crete, Greece NA 98.89 NHMC 80.5.63.46 - M. canescens Crete, Greece M 99.78 NHMC 80.5.63.6 - M. canescens Crete, Greece M 99.15 MCNG 19070501 - M. meles La Roca del Vallès, Spain M 114.51 MCNG 98042801-7507 - M. meles Lliçà d'Amunt, Spain M 116.15 MCNG 99031501 - M. meles Mieres, Spain F 109.07 MCNG 001022301 - M. meles Batet de la Serra, Spain F 108.26 Table 2 List of the origin, insertion and main locomotive function of each muscle analysed in this study. Anatomical and osteological descriptions are based on terminology from Fishbeck and Sebastiani ( 2001 ), Evans et al. (2012), Ercoli et al. ( 2015 )d hmer et al. (2020). No. Muscle name Abbr. Origin Insertion Main function Shoulder 1 Mm. supraspinatus - M. supraspinatus principle - M. supraspinatus intermediate - M. supraspinatus cranial SSp Supraspinous fossa, scapular spine Cranial aspect of the scapular spine Cranial aspect of supraspinous fossa Humeral greater tubercle Humeral greater tubercle Humeral great tubercle Shoulder extension, humeral protractor Shoulder extension, humeral protractor Shoulder extension, humeral protractor 2 M. infraspinatus ISp Infraspinous fossa, scapular spine Humeral greater tubercle Shoulder flexion, humeral rotation 3 M. subscapuaris SSc Subscapular fossa Humeral lesser tubercle Scapular adductor 4 M. teres major TMj Axillary border of scapula Medial margin of pectoral ridge Shoulder flexion, limb retraction 5 M. teres minor TMn Axillary border of scapula Humeral greater tubercle Shoulder flexion, humeral rotation 6 Mm. deltoideus - M. spinodeltoideus - M. acromiodeltoideus DSp DAc Scapular spine Acromion Deltoid crest of humerus Shoulder flexion and humeral abductor Brachium 7 M. coracobrachialis CBH Coracoid process of scapula Humeral diaphysis Shoulder stabilizer and humeral adductor 8 M. brachialis BCH Proximal humerus diaphysis Coronoid process of ulna Elbow flexion and forearm supinator 9 M. brachioradialis BRAD Proximal aspect of humeral epicondylar crest Distal aspect of radius Elbow flexion, forearm supinator 10 M. biceps brachii BB Tubercle of scapula Bicipital tuberosity of radius Shoulder extension, elbow flexion 11 Mm. triceps brachii - M. triceps brachii caput laterale - M. triceps brachii caput longum - M. triceps brachii caput mediale - M. triceps brachii caput accessorium - M. triceps brachii caput angulare TBLa TBLo TBMe TBAc TBAn Proximal surface of deltoid crest Mid-axillary border of scapula Mediocaudal humeral diaphysis Distal facet of humeral diaphysis Axillary edge of scapula Lateral aspect of ulnar olecranon Caudal aspect of ulnar olecranon Medial aspect of ulnar olecranon Medial aspect of ulnar olecranon Caudal aspect of ulnar olecranon Elbow extension Elbow extension, shoulder flexion Elbow extension Elbow extension Humeral retractor, elbow extension 12 M. tensor fasciae antebrachii TFA Caudal border of m. latissi dorsi Medial aspect of ulnar olecranon Elbow extension 13 M. anconeus ANC Distal epicondylar crest Lateral section of olecranon Elbow extension, forearm pronator Distal arm 14 15 16 17 18 19 Extensors M. extensor carpi radialis longus M. extensor carpi radialis brevis M. extensor digitorum communis M. extensor digitorum lateralis M. extensor carpi ulnaris - humeral head - ulnar head M. extensor digiti I and II ERL ERB EDC EDL EU ED Proximal aspect of humerus epicondylar crest Proximal aspect of humerus epicondylar crest Proximal aspect of humerus epicondylar crest Proximal aspect of humerus epicondylar crest Humeral epicondylar crest Ulnar and radial shaft Dorsal aspect of ulnar shaft Base of metacarpal II Base of metacarpal III Distal phalanges of digits II-V Distal phalanges of digits III-V Base of metacarpal V Distal phalanges of digits I-II Elbow flexion, wrist extension Elbow flexion, wrist and digit extension Elbow flexion, wrist and digit extension Elbow flexion, wrist and digit extension Elbow flexion, wrist extension Extensor of digits I-II 20 21 22 23 Flexors M. flexor digitorum profundus - humeral heads; profundus, mediale, and laterale - ulnar head - radial head M. flexor carpi radialis M. flexor carpi ulnaris M. flexor digitorum superficialis FDP FDPh p FDPh m FDPh l FDPu FDPr FCR FCU FDS Caudomedial aspect of the: humeral epicondyle, ulnar, and radial shafts Epicondyle of humerus Medial aspect of olecranon and epicondyle Superficial to distal FDP humeral heads Distal phalanges of digits II-V Base of metacarpal II and III Sesamoid proximal to metacarpal V Superficial to FDP Wrist and digital flexion Wrist flexion Wrist flexion Wrist flexion 24 M. palmaris longus PL Distal epicondyle of humerus Distal phalanges of digits II-V Wrist and digit flexion 25 M. abductor digiti I AD Lateral ulnar and radial shaft Metacarpal I Forearm abductor Deep 26 M. pronator teres PT Medial epicondyle Medial section of radius shaft Forearm pronator 27 M. supinator SU Lateral epicondyle Medial section of radius shaft Forearm supinator 28 M. pronator quadratus PQ Medial ulnar shaft Distal radial shaft Forearm pronator Comparative myology and osteology The anatomical structure of M. anakuma and M. leucurus were compared to the published myological descriptions of five mustelid species: Meles meles (Böhmer et al. 2020 ), two ictonychines ( Galictis cuja and Ictonyx striatus ) (Ercoli et al. 2015 ; Windle and Parsons 1897 ), and two guloines ( Martes martes and Martes foina ) (Böhmer et al. 2018 ; Yousefi et al. 2018 ; Böhmer et al. 2020 ), as well as two procynids: Potos flavus (Vélez-García et al. 2023 ) and Procyon lotor (Allen 1882 ; Feeney 1999 ). Melinae (to which Meles belongs), Ictonychinae, and Guloinae are all closely-related subfamilies within Mustelidae, whereas Procynidae is a separate family adjacent to Mustelidae but still within the same order, Carnivora (Law et al. 2018 ). Therefore, they share fundamental anatomical structures yet exhibit a range of forelimb functional specializations – including arboreal and generalist behaviors – which provide a valuable comparative framework for interpreting osteological data. For M. canescens , no published myological data was available and so, only osteological comparisons were conducted on specimens from the insular population on Crete. The 3D models for comparative specimens were generated using an EinScanSP 3D scanner (Shining 3D V3.1.0.1, China) and visually analysed in the program 3D Slicer V5.0.3 (Fedorov et al. 2012 ). Continental-insular pairwise comparisons and dispersion history of Meles In this study, we acknowledge that M. canescens has both continental and insular populations. However, for the continental-insular pairwise comparison analysis, we chose M. meles as the mainland counterpart to M. canescens individuals from the island population on Crete because M. meles is known from mDNA to be ancestral to M. canescens (Marmi et al. 2006 ), and since specimens of continental M. canescens were not accessible to us. This continental-insular comparison method has been previously successfully applied to M. canescens from Crete, revealing several shared cranial and dentition traits that reflect adaptations to island environments (Abramov and Puzachenko 2013 ; Savvidou et al. 2022 ). Here, we are working under the assumption that despite the shorter isolation period on Crete, M. canescens populations would still have undergone morphological transformation that distinguishes them from the mainland inhabitants. Following the evolutionary history of Meles summarized below, we grouped the genus into the following continental-insular pairs for comparative analysis: leucurus-anakuma and meles-canescens . Geographically, M. meles occupies much of Europe and the British Isles, while M. canescens ranges across southwestern Asia (Turkey, the Caucasus, Iran, etc.) and has established insular populations on the Mediterranean islands of Crete and Rhodes (Proulx et al. 2016 ). M. leucurus inhabits most of mainland China, Russia, and Korea, while M. anakuma is endemic to the Japanese archipelago, excluding Hokkaido and the Ryukyu Islands (Proulx et al. 2016 ). Palaeontological evidence suggests that Meles evolved from the genus Melodon (Neel and Cheeseman 1996), where the earliest known Meles fossil specimens found in Eurasia have been dated to the Late Pliocene (Madurell-Malapeira et al. 2011 ; Faggi et al. 2024 ). An initial divergence occurred between European ( M. meles and M. canescens ) and Asian ( M. leucurus and M. anakuma ) lineages during the Early Pleistocene, consistent in molecular and morphological studies (Marmi et al. 2006 ; Del Carro et al. 2010; Law et al. 2018 ; Abramov and Puzachenko 2013 ; Faggi et al. 2024 ). In the molecular study, the divergence between M. leucurus and M. anakuma is estimated at 1.09–0.21 Ma, whereas the split between M. meles and M. canescens is estimated to have occurred between 2.37 Ma and 450 ka (Marmi et al. 2006 ; Law et al. 2018 ). The dispersal of M. anakuma to Japan likely occurred via the “Korean land bridge” during the Middle Pleistocene, as supported by fossil occurrences (Shikama 1949 , 1962 ; Kawamura et al. 1989 ; Hasegawa 2012 ). Following this colonization, M. anakuma has remained genetically distinct from its continental relatives. The presence of badgers on Crete, represented by M. canescens , likely reflects a more recent, human-mediated introduction, with the oldest specimens dated only to approximately 3000 years ago (Masseti 1995 ). Meanwhile, M. meles populations in the British Isles became geographically isolated from continental Europe over 450 ka (Gupta et al. 2017 ), and small island populations, such as those on the Irish Peninsula, have arisen more recently (Sleemen et al. 2009). Institutional abbreviations: MCNG, Museu de Ciencies Natural de Granollers; NHMC, Natural History Museum of Crete; NMNS, National Museum of Natural Science in Tokyo; SNU, Seoul National University. Results The gross muscular topography of both Meles species was similar and shared the same general origin and insertion locations. Therefore, the muscular maps contain the complied results from all dissections undertaken for this study, with differences in muscle shape and structure between M. anakuma and M. leucurus described in detail in the following muscle descriptions. Shoulder muscles M. supraspinatus This muscle is oblong and located in the lateral aspect of the scapula covering the entire supraspinous fossa surface (Fig. 2a). It extends dorsally along the scapular spine and inserts via thick tendons and some fleshy fibers to the proximal region of the humerus greater tubercle (Fig. 3a and c, 6e and 7a). Three bellies were observed: an intermediate one, running along the dorsal edge of the scapular spine, a principle one, on the lateral fossa surface, and a cranial one, which runs along the cranial ridge of the scapula and projects slightly towards the medial aspect. M. infraspinatus This rectangular muscle is closely fused to the infraspinous fossa by fleshy fibers located on the lateral aspect of the scapula (Fig. 2a). It follows the caudal edge of the scapular spine, covers the internal surface of the metacromion and inserts via a thick tendinous band at the lateral aspect of the humeral greater tubercle, directly distal to m. supraspinatus (Fig. 3a, 6e and 7b). The caudal-most aspect of m. infraspinatus is distinguished from the rest by a notably thick tenacious band. M. teres major This muscle originates along the caudal axillary border of the scapula via fleshy fibers, partially superficially to m. infraspinatus, and extending across approximately 1/3 of the scapula length (Fig. 2b). It is a relatively flat, elongated and rectangular muscle that is caudally fused to m. latissimus dorsi and inserts at the medial aspect of the deltoid ridge on the humeral shaft via a short thick band of tendons. The insertion area is long and thin, roughly half the length of its origin point, and runs parallel and superficially to the m. triceps brachii caput mediale (Fig. 3b, 6e and 7c). M. teres minor A small fleshy muscle situated on the proximal lateral aspect of the scapular caudal edge below the glenoid fossa, directly inferior to m. infraspinatus (Fig. 3a and 6e). It runs parallel to that same muscle before inserting distally and slightly more caudally to m. infraspinatus below the humeral greater tubercle via a thickly banded tendon (Fig. 7b). M. deltoideus This muscle is located on the lateral aspects of the shoulder and is composed of two independent bellies: m. spinodeltoideus and m. acromiodeltoideus. M. spinodeltoideus originates along the cranial aspect of the scapular spine, caudal to m. supraspinatus and superficial to m. infraspinatus (Fig. 2). The origin of this belly varied between both species and individual specimens. Its fleshy fibers originated from approximately halfway and to the most distal aspect of the scapular spine. It is thin and elongated with a rounded caudal edge which fuses to the lateral distal aspect of m. infraspinatus via a wide thin band of fine tendons (Fig. 2a, 3a and c, 7a) This muscle continues proximally beneath m. acromiodeltoideus and, in several specimens, the two bellies fused together via fleshy fibers before inserting with fine tendons along the deltoid crest of humerus. In others, the two bellies remained independent with m. spinodeltoideus inserting caudally to m. acromiodeltoideus. M. acromiodeltoideus originates at the acromion on the proximal aspect of the scapular glenoid fossa (Fig. 2a, 3a and c, 7a). It is rhombus-like and connects with fine fascia along the proximal aspect of the humeral greater tubercle, distal to m. supraspinatus, and then inserts along the humeral deltoid ridge from the lateral to the cranial aspect. M. subscapuaris This rectangular muscle originates across the entire medial surface of the scapula on the subscapular fossa and inserts via short thick tendons along the caudal aspect of the humeral lesser tubercle (Fig. 2b). There are six fibrous groups originating from the proximal aspect of the scapula, which then fan out towards the distal scapular borders. Each fibrous group is weakly separated by fine tendonous fibers which are further imprinted on the surface and border of the subscapular fossa as bony ridges. The most caudally situated group is the largest and most elongated, stretching across the entire medial axillary edge of the scapula. The most cranially situated group inserts slightly superficially to the neighbouring group (Fig. 3b-c, and 7a). Brachium muscles M. coracobrachialis This muscle originates at the cranial aspect of the coracoid process on the scapula and extends down the medial aspect of the humeral diaphysis as a long, thin fleshy tendon before inserting on the medio-caudal region of the humeral shaft, dorsal to the medial epicondyle (Fig. 3a-c). For M. anakuma , m. coracobrachialis originates as a very long, thin tendon before broadening out approximately halfway down the humeral shaft and inserting as a long, triangular-like strip of fleshy fibers. M. leucurus , on the other hand, has a considerably shorter tendon at the origin, which expands outwards into an elongated strip of flat muscle at the medio-caudal aspect of the humeral lesser tubercle. It thins out once more just below m. teres major and inserts as a long thin tendon above the medial epicondyle (Figs. 4 and 7c-d). M. brachialis This muscle has an extensive origin, with thick fleshy fibers closely fused to the proximal aspect of the humerus diaphysis directly distal to m. triceps brachii laterale and up under the caudal-most aspect of the humeral head (Fig. 2a). This long, flat muscle continues down along the caudal humeral shaft and connects to m. brachioradialis via thick fleshy fibers (Fig. 5a). This then tampers out into a band of thin tendons and inserts at the ulnar tuberosity in the medial aspect (Fig. 3e and 7). M. brachioradialis The origin length of m. brachioradialis varied between both Meles species and specimens. This muscle originated from the proximal aspect of humeral lateral epicondylar crest, but in some specimens (e.g. M. anakuma : POM-259), it extends approximately halfway up the humeral diaphysis as a long fleshy band (Fig. 2, 5a and 6e). It is relatively flat at its origin and then extends along the forearm and fuses at the distal medial aspect of the radius (Fig. 7a). M. biceps brachii This muscle originates via a tight, compact band of tendons above the supraglenoid tubercle of the scapula and then rapidly expands out into a robust belly that extends down the medial aspect of the humerus (Fig. 2b, 5a and 6e). M. biceps brachii is held in place between the two humeral tubercle heads by its own retinaculum. It narrows again into a flattened tendon shortly before inserting on the bicipital tuberosity of the radius (Figs. 3 and 6e). M. anconeus This flat, triangular, entirely fleshy muscle fuses directly via fibrous tissue to the distal caudal surface of the lateral humeral epicondyle (Fig. 6e). It extends distally and inserts along the lateral aspect of the ulnar head and along the olecranon process. Fine, fleshy fibers also connect it to m. extensor carpi ulnaris (Fig. 7). M. triceps brachii This is the largest muscle in the brachium group and is situated in the lateral and medial aspects of the forelimb with five bellies: m. triceps brachii caput laterale, m. triceps brachii caput mediale, m. triceps brachii caput accessorium, m. triceps brachii caput longum, and m. triceps brachii caput angulare (Fig. 2). All bellies insert on the proximal head of the ulnar olecranon process. M. triceps brachii caput laterale has a relatively long and narrow origin via thin tendons to the proximal lateral aspect of the deltoid ridge, immediately distal to m. teres minor (Fig. 2a). However, there are additional fleshy attachments along the cranial lateral aspect of the humeral head and some fine fibrous tissues connecting it superficially to the distal laterally situated extensor muscles, as well along the supracondyloid ridge border. This belly inserts on the proximal lateral facet of the olecranon tuber, directly above m. anconeus (Fig. 7). M. triceps brachii caput mediale is located in the medial aspect of the forelimb and partially fuses to m. triceps brachii caput lateral via fleshy fibers which extend towards the olecranon tuber insertion (Fig. 2b). More fleshy fibers fuse to the caudal aspect of the humeral head and the caudal proximal facet of m. subscapularis, while fine expanses of tendons also extend down along the medial aspect of the deltoid ridge, running parallel to the insertion of m. teres major (Fig. 7). M. triceps brachii caput accessorium is a short cylindrical muscle located in the medial aspect of the forearm (Fig. 2b). It is the smallest of the m. triceps brachii muscle group and is situated partially superficial to the insertion of m. coracobrachialis. It originates from the caudal-medial aspect of the humeral medial epicondyle and inserts on the proximal medial facet of the ulnar olecranon, directly distal to both m. triceps brachii caput longum and m. triceps brachii caput mediale (Fig. 4b, 6 and 7). M. triceps brachii caput longum is the largest and thickest belly of the m. triceps brachii group, located in both the medial and lateral aspects of the forelimb (Fig. 2). This muscle originates via abundant fleshy fibers at the mid-axillary border of the scapula, distal to m. triceps brachii caput laterale in the lateral view. It inserts on the most proximal caudal aspect of the ulnar olecranon (Fig. 3a, 4 and 6). M. triceps brachii caput angulare is a very flat, elongated muscle that is the most caudally located of the m. triceps brachii heads and is situated superficially to m. triceps brachii caput longum (Figs. 2 and 3a). It originates via fleshy fibers on the axillary-most border of the scapula in the medial view, directly superficial to m. teres major. It attaches via fine fibrous tendons on the caudal facet of the ulnar olecranon (Fig. 7b). M. tensor fasciae antebrachii This is a small, very flat muscle on the medial aspect of the forelimb and sits directly superficial to m. triceps brachii caput mediale and m. triceps brachii caput longum (Fig. 2b). It fuses directly to the caudal border of m. latissi dorsi via thin fascia for M. anakuma and fleshy fibers for M. leucurus (Fig. 5b) before inserting on the medial facet of the olecranon tuber immediately distal to m. triceps brachii caput longum. Distal arm muscles M. extensor The extensor group has six independent muscles: m. extensor carpi radialis longus, m. extensor carpi radialis brevis, m. extensor digitorum communis, m. extensor digitorum lateralis, m. extensor carpi ulnaris, and m. extensor pollicis, which are stacked vertically to each other along the humeral lateral epicondylar crest and lateral epicondyle (Fig. 2a). M. extensor carpi radialis longus and m. extensor carpi radialis brevis originates via fleshy fibers from the proximal aspect of the humeral lateral epicondyle (Fig. 2a). They are fused at their origin and split approximately one-third of the way towards the ulnar styloid, with m. extensor carpi radialis longus situated superficially to m. extensor carpi radialis brevis, and then become two tightly wound bands of flattened tendons. These muscles are held in place by their own retinaculum located at the distal anterior surface of the radius, with m. extensor carpi radialis longus inserting at the base of metacarpal II, and m. extensor carpi radialis brevis inserts at the base of metacarpal III (Fig. 7). M. extensor digitorum communis originates via fleshy fibers on the humeral lateral epicondylar crest, directly distal to m. extensor carpi radialis longus and m. extensor carpi radialis brevis (Fig. 2a). It extends distally towards the wrist before splitting into four groups of fine, elongated tendons that insert on the distal phalanges of digits II-V (Fig. 5a and 7). M. extensor digitorum lateralis originates via fleshy fibers on the distal aspect of humeral lateral epicondylar crest immediately distal to m. extensor digitorum communis (Fig. 2a). It is fused as a single entitity at its origin point however, it splits into two bellies – medial and lateral – approximately midway down the ulnar shaft. The medial belly further divides into two long flattened tendons that inserts in the distal phalanges of digits III and IV, while the lateral belly narrows into a single long thin tendon that inserts in the distal phalange of digit V (Fig. 5a and 7). M. extensor carpi ulnaris originates via fleshy fibers from the proximal aspect of the humeral lateral epicondyle directly distal to m. extensor digitorum lateralis (Fig. 2a). This muscle rapidly expands into a thick belly before narrowing again into a condensed cluster of tendons which insert beneath m. extensor digit I & II at the base of metacarpal V (Fig. 5a and 7). M. extensor digiti I & II is thick, fleshy and, unlike the other extensor muscles, is originates via fleshy fibers along the lateral ulnar shaft (Fig. 6e). It extends diagonally across the forearm towards the medial side, passing beneath all the extensor muscles excluding m. extensor carpi ulnaris, and then inserting as very fine tendons on the distal phalanges of digits I and II (Fig. 5a and 7). M. flexor The flexor group is located in the medial aspect of the humeral epicondyle and consists of four independent muscles: m. flexor digitorum profundus, m. flexor digitorum superficialis, m. flexor carpi radialis, and m. flexor carpi ulnaris (Fig. 2b). M. flexor digitorum profundus is the largest of the flexor muscles and is located in the deep caudal aspect of the distal forearm and consists of five heads: three with humeral origins – caput humerale laterale, caput humerale mediale, and caput humerale profundus, one originating on the ulna – caput ulnare, and one with origins on both the ulna and radius – caput radiale (Fig. 2b, 3 and 7). The humeral heads originate on the distal caudal-medial aspect of the humeral epicondyle however only caput laterale and caput profundus originate on the bone, as caput mediale fuses directly to caput laterale via fleshy fibers that extend midway along the ulna shaft towards the wrist. Caput profundus is considerably smaller, thinner and furtherly independent from the other two humeral heads, connected lengthwise only via thin fascia. Caput ulnare has a large heavy belly, originating via fleshy fibers from the proximal caudal aspect of the ulna shaft and extending two-thirds towards the wrist, while caput radiale is relatively flat and stretches between the lateral and cranial aspects of the radius and ulna shafts as fleshy fibers. M. flexor digitorum profundus heads fuse together (caput ulnare notably superficial to the rest) and are held in place by a thick retinaculum at the wrist before dividing into thin elongated tendons that insert into the distal phalanges I-V (Fig. 7). M. flexor digitorum superficialis is a fleshy relatively thin muscle situated directly superficial to m. flexor digitorum profundus (Fig. 6b). It does not appear to insert directly onto the caudal humeral medial epicondyle and instead its origin is fused to the distal surface of m. flexor digitorum profundus caput humerale laterale and mediale bellies. The insertion is also directly dorsal to m. flexor digitorum profundus, this time by thin fatty fascia superficial to m. flexor digitorum profundus caput ulnare, immediately between the diversion of the tendons towards digits I and V (Fig. 6b). M. flexor carpi radialis originates from the medial epicondyle of the humerus directly distal to m. pronator teres (Fig. 2b). It narrows into two thin long tendons that insert into the base of metacarpal II and III (Fig. 6b-d). It is also partially fused via thick fleshy fibers to m. pronator teres midway along the distal arm (Figs. 5 and 7). M. flexor carpi ulnaris is located on the medial side of the distal forearm, originating via fleshy fibers from the medial aspect of the ulnar olecranon tuber and medial humeral epicondyle as two independent bellies: caput ulnare and caput humerale, respectively (Fig. 2b). Caput ulnare is large and thick, while caput humerale is comparatively thin. These two heads fuse together partway along the ulnar shaft before inserting via a thin long tendon on the sesamoid most proximal to metacarpal V (Fig. 6a and 7). M. palmaris longus This muscle originates from the distal medial epicondyle of the humerus caudal to m. flexor carpi radialis (Fig. 2b and 6a-c). It rapidly expands out into a robust belly and then quickly thins out again into four flattened tendons which pass over the retinaculum that secures m. flexor digitorum profundus at the wrist and inserts into the distal phalanges of digits II-V (Fig. 7c-d). M. abductor digiti I It is a deep muscle that originates via fleshy fibers along the whole lateral surface of the ulna and radial shafts (Fig. 6e). It then narrows into a thin tendon that inserts into the base of metacarpal I (Fig. 7f-h). Deep muscles M. pronator teres It is a roughly semicircular muscle originating via fleshy fibers on the most proximal aspect of the humeral medial epicondyle, immediately cranial to the insertion of m. triceps brachii caput accessorium (Fig. 2b and 3d). It partially splits into two indistinct fleshy bellies which come together quickly to insert as a long insertion strip located cranially mid-way down the radial shaft (Fig. 5a, 6 and 7a). M. supinator This is a relatively flat, triangular muscle originating via thick fleshy fibers on the distal cranial aspect of the lateral humeral epicondyle, directly adjacent to m. extensor carpi ulnaris (Fig. 3d). It runs roughly parallel to m. pronator teres and then inserts on the dorsal-medial aspect of the radial shaft, approximately two-thirds of the way towards the distal styloid process (Fig. 6e). M. pronator quadratus This muscle is fused via thick fleshy fibers to the dorsal-medial distal surface of the ulna interosseous crest (Fig. 3d-e). It is rectangular, flat, and inserts on the caudal distal surface of the radius via fleshy fibers (Fig. 6d and 7b). Discussion In this study, we examined the forelimb myology of M. leucurus and M. anakuma and compared their muscular structure to published data of other closely related species. While the general topography of the forelimb muscular structure was similar, we found several key differences in the shape and form of the muscles which we will discuss in detail. Then, we analysed the forelimb osteology of Meles as continental-insular pairs ( leucurus-anakuma and meles-canescens , where here M. canescens refers to the insular communities inhabiting Crete) to comparatively investigate the functional adaptations of island Meles populations. Comparative muscular anatomy of M. anakuma and M. leucurus For the shoulder muscles, the three m. supraspinatus bellies described in Ercoli et al. ( 2015 ) for the lesser grison ( Galictis cuja ) are also evident in both M. anakuma and M. leucurus observed here (Fig. 2 ). However, both badgers have a comparatively more robust cranial belly and overall larger m. supraspinatus and m. infraspinatus surface areas than G. cuja (Fig. 2 ). The higher number of m. supraspinatus bellies has been linked to an increased range of motion in the shoulder joint, particularly during arm-extension (Ercoli et al. 2015 ). For m. teres major in both Meles species studied here, its origin is partially superficially fused to the distal caudal border of m. infraspinatus. This observation has also been reported for the kinkajou, Potos flavus (Vélez-García et al. 2023 ), and for the racoon, Procyon lotor (Feeney 1999 ; Allen 1882 ); however, it has not been described for any mustelid species although it appears to be present in several studies (e.g. G. cuja , Ercoli et al. 2015 ; Martes martes and Martes foina , Böhmer et al. 2020 ). The m. teres major origin itself is approximately one-fourth of the scapular length, which is considerably longer than in G. cuja (Ercoli et al. 2015 ). M. teres major is a powerful shoulder joint flexor and limb retractor used during the power stroke and so is particularly enlarged in specialised diggers (Moore 2011 ; Böhmer et al. 2020 ). In Meles , m. coracobrachialis is a small, thin muscle, yet its shape and nomenclature vary considerably among caniforms (Vélez-Garcia et al. 2023: table 3). Windle and Parsons ( 1897 ) described it as m. coracobrachialis medius in Meles , noting its more distal insertion on the humeral shaft compared to other species. In G. cuja , it has been termed m. coracobrachialis brevis, with a short insertion directly distal to the humeral lesser tubercle (Ercoli et al 2015 ). P. flavus possesses both brevis and longus heads, inserting on the proximal medial aspect of the humeral shaft and directly above the supracondylar foramen (Vélez-Garcia et al. 2023). The red panda, Ailurus fulgens (Fisher et al. 2009 )d martes (Yousefi et al. 2018 ) show only the more distal insertion. Böhmer et al. ( 2018 ) recorded a notably more distal insertion in M. foina and M. martes on the medial aspect of the olecranon. In this study, while origin and insertion were consistent, the position of the muscle belly varied between species. For M. leucurus , the belly lay directly beneath the humeral lesser tubercle, whereas in M. anakuma , it was situated distally along the humeral shaft (Fig. 4 ). M. anakuma also exhibited a second smaller belly that inserted directly distal to the humeral lesser tubercle (Fig. 4 b), resembling the dual insertion seen in P. flavus (Vélez-Garcia et al. 2023), although it was significantly reduced (Fig. 4 ). This distal insertion is most similar to A. fulgens in size and shape, while the dual arrangement mirrors that of arboreal taxa like P. flavus . Functionally, m. coracobrachialis is a weak shoulder stabilizer and humeral adductor, and its reduced appearance suggests a minimal impact on locomotion (Taylor 1974 ). However, its presence has been associated with arboreal activity (Salesa et al. 2008 ; Ercoli et al. 2015 ), and more distal insertions, as in P. flavus and A. fulgens , may enhance shoulder adduction strength and increase the animals’ climbing ability (Monroy-Cendales et al. 2020 ; Monroy-Cendales et al. 2023 ). Ercoli et al. ( 2015 ) considered the muscles’ presence a plesiomorphic trait within Carnivora. Our findings suggest interspecific variation in Meles , with M. anakuma retaining a dual insertion pattern similar to more arboreally-adapted taxa, potentially reflecting an ancestral or ecologically driven trait (Fig. 9 ) (Marmi et al. 2006 ; Law et al. 2018 ). In general, there are four m. triceps brachii heads: laterale, longum, mediale, and accessorium, which all insert on the proximal aspect of the ulnar olecranon however, there are frequent variations among caniforms. In all mustelids and mephitids (Windle and Parsons 1897 ; Moore et al. 2013 ; Ercoli et al. 2015 ), including the Meles species examined in this study, there is an additional head, m. triceps brachii caput angulare, which originates at the most distal caudal border of the scapula and inserts on the caudal aspect of the ulnar head (e.g. Figure 2 ). For P. flavus , m. triceps brachii caput angulare originates considerably more proximally along the scapular axillary border, likely making space for a larger m. teres major origin (Böhmer et al. 2020 ). Functioning in conjunction with m. triceps brachii to support elbow extension is the m. tensor fasciae antebrachii muscle, the presence of which has not been previously described for Meles . In both P. flavus (Vélez-García et al. 2023 )d lotor (Feeney 1999 ), m. tensor fasciae antebrachii had two heads (caudal and cranial), whereas for Meles , only the cranial belly was present and fused directly to the caudal border of m. latissi dorsi via thin fascia for M. anakuma , and fleshy fibers for M. leucurus (Fig. 2 b, 4 a and 5 b). For G. cuja (Ercoli et al. 2015 ) however, it was considered the modified intermediate belly of m. triceps brachii mediale. Our findings highlight the phylogenetic evolution of m. tensor fasciae antebrachii among caniforms (Fig. 9 ). In contrast to the four heads of m. flexor digitorum profundus described for M. meles (Böhmer et al. 2020 ) and typically present in canids and felids (Fishbeck and Sebastiani, 2001 ; Evans et al. 2012), five heads were consistently observed in our specimens, with an additional humeral head, m. flexor digitorum profundus humerale profundus (Fig. 5 c). In both M. anakuma and M. leucurus , m. flexor digitorum superficialis originates significantly more distally and exhibits a well-developed fleshy belly, differing from the shorter muscle belly observed in G. cuja (Ercoli et al. 2015 )d fulgens (Fisher et al. 2009 ). In this study, the muscle does not attach directly to the bone but instead fuses entirely with m. flexor digitorum profundus at the wrist bundle (Fig. 6 ), contrasting with the distinct distal insertion described for G. cuja and A. fulgens . Functionally, m. flexor digitorum superficialis supports wrist flexion however, given its reduced appearance in Meles and often misidentification or omission in anatomical descriptions, its comparative analysis across taxa remains challenging (Ercoli et al. 2015 ; Perdomo-Cárdenas et al. 2021 ). Continental-insular pairwise comparisons in Meles Comparative studies have successfully explored the close relationship between myology and osteology in describing locomotive habits (e.g. Van Valkenburgh 1987 ; Argot 2001 ; Fabre et al. 2013 ; Böhmer et al. 2019 ). In general, Meles badgers exhibit skeletal features consistent with a powerful ‘scratch-digger’ burrowing style. These include large muscle attachment sites at the shoulder and elbow joints, resulting in stout humeri and ulnae with thick shafts and robust proximal and distal features (Fig. 8 ) (Hildebrand and Goslow 2001 ; Moore et al. 2011; Samuels et al. 2013 ). The shoulder muscles – especially the rotator cuff muscle m. subscapularis and the abductor m. deltoideus – contribute to shoulder stabilization and humeral rotation (Fishbeck and Sebastiani 2001 ; Janis and Martín-Serra 2020 ), consistent with the enlarged humeral greater tubercles observed in all Meles specimens in this study (Fig. 8 ). Additionally, digging performance is enhanced by well-developed humeral medial and lateral epicondyles and an elongated ulnar olecranon, which are attachment sites for powerful distal flexor and extensor muscles (Fig. 8 ) (Argot 2001 ; Hildebrand and Goslow 2001 ; Moore et al. 2013 ; Samuels et al. 2013 ; Rose et al. 2014 ). In the leucurus-anakuma pair, the insular M. anakuma exhibits the cranially inclined olecranon typically seen in semifossorial and arboreal species (Fig. 8 h) (Fabre et al. 2013 ). In contrast, M. leucurus retains a straighter ulnar profile in the medial view with a caudally inclined olecranon process more common in terrestrial quadrupeds optimized for strong elbow extension (Fig. 8 g) (Henderson et al. 2017 ). Functionally, these changes alter the leverage and direction of muscle force across the elbow joint. In M. anakuma , the caudal head of the olecranon (insertion site for m. triceps brachii caput longum) is reduced, while the cranial head (for caput mediale) is enlarged (Fig. 8 h). This shift suggests decreased involvement of shoulder-extending caput longum, and increased reliance on caput mediale, which primarily supports late-phase elbow extension – often seen in species engaging in climbing or intricate foraging behaviours (Kholinne et al. 2018 ). Conversely, M. leucurus demonstrates stronger development of the medial epicondyle, particularly in attachment regions for m. pronator teres and other forearm flexors (Fig. 8 c), suggesting enhanced torque production during pronation-supination. This muscular development is likely associated with sweeping arm movements for overturning compact soil and possibly an adaptation to life in colder regions with harder substrates (Neal and Cheeseman, 1996 ) where badger setts are often dug deeper and exhibit more complex structures to withstand the lower temperatures (Choi et al. 2019 ). Additionally, the twisted orientation of the humeral lateral epicondyle relative to the medially tilted head further supports the presence of strain-induced remodeling from forceful digging behaviours (Fig. 8 c). M. anakuma also exhibits a slight reduction in the distal projection of the humeral trochlea, which may indicate an increase in manouverability in the elbow joint (Fig. 8 d). Therefore, the insular M. anakuma exhibits skeletal shifts that suggest increased arm flexibility, potentially indicating less intensive digging or more diverse substrate interaction, reflecting differences in island environments. In the meles-canescens pair, the insular population of M. canescens exhibited a decreased length of the humerus and ulna, and an increased robusticity of the deltoid ridge and epicondylar crest (Fig. 8 a–b, e-f). These changes suggest a functional shift toward more compact and powerful limb extension, potentially compensating for the reduced lever length via greater muscle mass at origin and insertion sites. The reduction in the humeral lesser tubercle – the insertion point for m. subscapularis, and the reduced distal projection of the humeral trochlea in the M. canescens specimens, may indicate increased mobility at the shoulder and elbow joints (Fig. 8 b) (Argot 2001 ; Salesa et al. 2020 ). This, combined with the enlarged cranial aspect of the olecranon and reduced projection in the humeral trochlea, implies enhanced arm-extension capabilities in the insular form, possibly reflecting greater reliance on limb flexibility (Fig. 8 a-b, e-f). These morphological patterns suggest that the Crete population of M. canescens has evolved toward a more versatile digging style, potentially as a response to differing substrate properties or ecological demands on the island. Functional responses shared in insular Meles Previous studies have reported reductions in the body size of insular Meles populations – including M. canescens inhabitants on Crete and M. anakuma in Japan – compared to their mainland counterparts, M. meles and M. leucurus respectively (Baryshnikov et al. 2002; Abramov and Puzachenko 2013 ; Proulx et al. 2016 ; Savvidou et al. 2022 ). Nonetheless, our study found no significant size difference (based on humerus length) between M. leucurus and M. anakuma (Table 1 ), although a reduction was evident between M. meles and the M. canescens populations of Crete (Fig. 9 ). This variation in body response highlights the complexity of insular adaptations in medium-carnivores and suggests that body size alone may not fully capture the extent of functional adaptation in island settings. Notably, we recognized significant morphological patterns in the forelimbs of the insular forms. Both island popuations of M. canescens and M. anakuma exhibited shared reductions in the distal and caudal projections of the humeral trochlea and enlargements of the cranial aspect of the olecranon tuber (Fig. 8 b, e, and 9 ), suggesting that increased elbow flexibility – possibly favoring greater forelimb mobility for handling variable or less resistant substrates, is a general trend among insular Meles species. These osteological changes complement ecological observations for both insular and mainland badger populations. Field-based ecology has been studied extensively for M. anakuma and M. meles , including annual hibernation periods (e.g. Kowalczyk et al. 2003 ; Tanaka 2006 ), social structure (e.g. Neel and Cheeseman 1996; Kaneko et al. 2014 ; Tanaka et al. 2022 ), and diet and distribution (e.g. Kaneko et al. 2006 ; Proulx et al. 2016 ), whereas ecological and behavioral data remain sparse for M. leucurus and M. canescens , making direct comparisons of the examined continental-insular pairs challenging. Despite these gaps, a few studies have observed dietary shifts and increased diurnal activity in M. meles populations on Rutland Island compared to their continental counterpart, interpretating these changes as insular traits (Sleeman et al. 2009 ; Sleeman and Davenport 2016 ). Furthermore, ecological studies on M. leucurus populations on Jeju Island recorded badgers as active from 3pm (Oh 2007 ), which is significantly earlier than what has been recorded for M. anakuma , with activity beginning from 6pm (Tanaka 2005). Possible increased diurnal activity was also extrapolated for other insular badgers ( M. canescens from Crete and M. anakuma ) based on cranial features such as greater interorbital distance (Savvidou et al. 2022 ). These patterns echo broader trends observed in other island-adapted carnivores – such as the Channel Islands foxes and Arctic fox populations which have shown both ecological and morphological shifts that align with increased arboreal activities and altered foraging strategies (Wayne et al. 1991 ; Moore and Collins 1995 ; Nanova and Prôa 2017 ). Collectively, our findings indicate that enhanced limb mobility may represent broader insular trends extending beyond terrestrial generalists to include subfossorial forms. We also note that direct behavioral observations are inherently limted by observer access, seasonal variation, and plastic ecological strategies especially for nocturnal, burrowing mammals like badgers. In contrast, skeletal features accumulate the selective pressures exerted over evolutionary time scales and can thus serve as good proxies for habitual locomotor and foraging behaviour. By linking skeletal morphology to inferred changes in habitat use, foraging strategy and locomotor behavior, our study reinforces the importance of integrating ecological and anatomical approaches to better understand how medium-sized mesocarnivores like Meles adapt to island ecosystems. Conclusions This study provides the first comparative myological analysis of the forelimbs in M. anakuma and M. leucurus , revealing both conserved and novel anatomical traits. Notably, the m. coracobrachialis muscle, which has been described as a plesiomorphic trait in Mustelidae, has a more distal main belly and second shorter head for M. anakuma compared to M. leucurus , which may represent taxon-specific variations not previously documented in other mustelid species. The presence of m. tensor fasciae antebrachii is observed in Meles for the first time and highlights the phylogenetic significance of this muscle among caniforms. Osteological comparisons with other continental-insular Meles pairs show shared morphological adaptations—such as increased forelimb flexibility—in insular species. Contrary to expectations based on prior studies of insular dwarfism, M. anakuma and M. leucurus did not differ significantly in body size, suggesting that other undetermined factors had a more promeninet influence on forelimb development in M. anakuma . These findings underscore the complex interplay between phylogeny, ecology, and biomechanics in shaping the musculoskeletal evolution following dispersal events. Future research should prioritize integrating myological, morphological and ecological data – particularly with specimens from underrepresented regions such as Greece, South Korea, and mainland China – to further refine our understanding of functional and evolutionary patterns in Meles and other mustelids. Detailed investigations into the finer-scale factors which influence intraspecific variation, such as sexual dimorphism and ontogenetic growth, would also greatly enhance our understanding of the ecological and behavioral roles favored by different Meles species. Declarations Acknowledgements We would like to express our gratitude to Chang-Yong Choi (Seoul National University) and Hee-Jong Kim (Wildlife Rescue Center, Ulsan) for providing samples and allowing us to perform dissections in their laboratories. We are especially grateful to Shin-ichiro Kawada (National Museum of Nature and Science, Japan), Petros Lymberakis and Panagiotis Georgiakakis (Museum of Crete, Greece), Toni Arrizabalaga and Anna Surroca (Museu de Ciencies Natural de Granollers, Spain), Roberto Portela Miguez and Phaedra Kokkini (National Museum of Natural Science in London), Géraldine Veron and Lucile Armand (National Museum of Natural Science in Paris), Aleksandra Panyutina (Tel Aviv University), Yuusuke Goto (Ibaraki Nature Museum), Eri Akasaki and Hiroshi Tanaka (Yamaguchi Prefectural Museum), Keiichi Takahashi (Lake Biwa Museum), Nozomi Nakanishi (Kita-Kyushu Museum of Natural and Human History), Fumio Takahashi (Mine City Museum of History and Folklore), Hashimoto Tatsuya and Mitsuharu Matsumoto (Kagoshima University), Yohoko Okumura (Kuzu Fossil Museum), Jun Nemoto (Tohoku University Museum), Taruno Hiroyuki (Osaka Museum of Natural History), and Satoshi Suzuki (Kanagawa Prefectural Museum of Natural History) for granting us access to their extant badger collections. We are also thankful to Haruto Sugeno, Shingo Nishimura and Haemin Seo for supplying us with the M. anakuma and M. leucurus roadkill specimens, and Mao Shimoda, Kyunghae Min and Tri Sayektiningsih for assisting with the dissections. Finally, we would further like to thank Kohei Tanaka (University of Tsukuba), Yoshikazu Hasegawa (Iida City Museum, Nagano), Yayoi Kaneko (Tokyo University of Agriculture and Technology), and Qigao Jiangzuo and Peiran Li (Institute of Vertebrate Paleontology and Paleoanthropology, China) for their constructive comments on the early draft. Author contributions All authors have read and agreed to the definitive version of the manuscript. Funding This work was partially funded by the Japanese Government Monbukagakusho Scholarship (grant number 210348). Data availability statement The datasets generated and analyzed during the current study are included in this published article. Declarations Competing interests The authors have no competing interests to declare that are relevant to the content of this article. References Abramov AV, Puzachenko AY (2013) The taxonomic status of badgers (Mammalia, Mustelidae) from Southwest Asia based on cranial morphometrics, with the redescription of Meles canescens . Zootaxa 3681(1):44–58. https://doi.org/10.11646/zootaxa.3681.1.2 Allen H (1882) The muscles of the limbs of the raccoon ( Procyon lotor ). Proc Acad Nat Sci Philadelphia 115–144. Argot C (2001) Functional-adaptive anatomy of the forelimb in the Didelphidae, and the paleobiology of the Paleocene marsupials Mayulestes ferox and Pucadelphys andinus . J Morphol 247(1):51–79. https://doi.org/10.1002/1097-4687(200101)247:13.0.CO;2-%23 Baryshnikov GF, Puzachenko AY, Abramov AV (2003) New analysis of variability of cheek teeth in Eurasian badgers (Carnivora, Mustelidae, Meles ). Russian Journal of Theriology 1(2):133–149. https://doi.org/10.15298/rusjtheriol.01.2.07 Benítez-López A, Santini L, Gallego-Zamorano J, Milá B, Walkden P, Huijbregts MA, Tobias JA (2021) The island rule explains consistent patterns of body size evolution in terrestrial vertebrates. Nat Ecol Evol 5(6):768–786. https://doi.org/10.1038/s41559-021-01426-y Böhmer C, Fabre AC, Herbin M, Peigné S, Herrel A (2018) Anatomical basis of differences in locomotor behavior in martens: A comparison of the forelimb musculature between two sympatric species of Martes . Anat Rec 301(3):449–472. https://doi.org/10.1002/ar.23742 Böhmer C, Fabre AC, Taverne M, Herbin M, Peigné S, Herrel A (2019) Functional relationship between myology and ecology in carnivores: do forelimb muscles reflect adaptations to prehension? Biol J Linn Soc 127(3):661–680. https://doi.org/10.1093/biolinnean/blz036 Böhmer C, Theil JC, Fabre AC, Herrel A (2020) Atlas of Terrestrial Mammal Limbs. CRC Press Choi TY, Lee SK, Woo DG (2019) Study on spatial ecology of Asian Badger ( Meles leucurus ) by radio telemetry and camera trapping: focusing on home-range and hibernation sett use pattern. J Korean Cadastre Inform Assoc 21:151–163. (in Korean) Coonan TJ, Schwemm CA, Garcelon DK (2010) Decline and Recovery of the Island Fox: A Case Study for Population Recovery. Cambridge University Press, England. Del Cerro I, Marmi J, Ferrando A, Chashchin P, Taberlet P, Bosch M (2010) Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora). Zool Scr 39(5):415–425. https://doi.org/10.1111/j.1463-6409.2010.00436.x Ercoli MD, Álvarez A, Stefanini MI, Busker F, Morales MM (2015) Muscular anatomy of the forelimbs of the lesser grison ( Galictis cuja ), and a functional and phylogenetic overview of Mustelidae and other Caniformia. J Mammal Evol 22:57–91. https://doi.org/10.1007/s10914-014-9257-6 Evans HE, De Lahunta A (2012) Miller's Anatomy of the Dog. Elsevier health sciences Fabre AC, Cornette R, Slater G, Argot C, Peigné S, Goswami A, Pouydebat E (2013) Getting a grip on the evolution of grasping in musteloid carnivorans: a three-dimensional analysis of forelimb shape. J Evol Biol 26(7):1521–1535. https://doi.org/10.1111/jeb.12161 Faggi A, Bartolini-Lucenti S, Madurell-Malapeira J, Abramov AV, Puzachenko AY, Jiangzuo Q, Peiran L, Rook L (2024) Quaternary Eurasian badgers: Intraspecific variability and species validity. J Mammal Evol 31(1):3. https://doi.org/10.1007/s10914-023-09696-y Fedorov A, Beichel R, Kalpathy-Cramer J, Finet J, Fillion-Robin J-C, Pujol S, Bauer C, Jennings D, Fennessy F, Sonka M, Buatti J, Aylward SR, Miller JV, Pieper S, Kikinis R (2012) 3D Slicer as an Image Computing Platform for the Quantitative Imaging Network. JMRI 30(9):1323–1341. https://doi.org/10.1016/j.mri.2012.05.001 Feeney S (1999) Comparative osteology, myology, and locomotor specializations of the fore and hind limbs of the North American foxes Vulpes vulpes and Urocyon cinereoargenteus . Dissertation, University of Massachusetts Amherst Fishbeck DW, Sebastiani A (2001) Comparative Anatomy: Manual of Vertebrate Dissection. Morton Publishing Company, Colarado Fisher RE, Adrian B, Barton M, Holmgren J, Tang SY (2009) The phylogeny of the red panda ( Ailurus fulgens ): evidence from the forelimb. J Anat 215(6):611–635. https://doi.org/10.1111/j.1469-7580.2009.01156.x Foster JB (1964) Evolution of mammals on islands. Nat 202(4929):234–235. https://doi.org/10.1038/202234a0 Giannico GR, Nagorsen DW (1989) Geographic and sexual variation in the skull of Pacific coast marten ( Martes americana ). Can J Zool 67(6):1386–1393. https://doi.org/10.1139/z89-197 Goltsman M, Kruchenkova EP, Sergeev S, Volodin I, Macdonald DW (2005) ‘Island syndrome’in a population of Arctic foxes ( Alopex lagopus ) from Mednyi Island. J Zool 267(4):405–418. https://doi.org/10.1017/S0952836905007557 Gupta S, Collier JS, Garcia-Moreno D, Oggioni F, Trentesaux A, Vanneste K et al (2017) Two-stage opening of the Dover Strait and the origin of island Britain. Nat Commun 8(1):15101. https://doi.org/10.1038/ncomms15101 Hasegawa Y (2012) Origin of modern mammals in Japan. Mammal Sci 52(2):233–247. https://doi.org/10.11238/mammalianscience.52.233 . (in Japanese) Hayashi S, Kubo MO, Sánchez-Villagra MR, Taruno H, Izawa M, Shiroma T, Nakano T, Fujita M (2023) Variation and process of life history evolution in insular dwarfism as revealed by a natural experiment. Front Earth Sci 11:1095903. https://doi.org/10.3389/feart.2023.1095903 Henderson K, Pantinople J, McCabe K, Richards HL, Milne N (2017) Forelimb bone curvature in terrestrial and arboreal mammals. PeerJ 5:e3229. https://doi.org/10.7717/peerj.3229 Hildebrand M, Goslow GE (2001) Digging, and crawling without appendages. In: Hildebrand M, Goslow GE, Hildebrand V (eds) Analysis of Vertebrate Structure. John Wiley and Sons Inc, New Jersey, pp 455–474 Janis CM, Martín-Serra A (2020) Postcranial elements of small mammals as indicators of locomotion and habitat. PeerJ 8:e9634. https://doi.org/10.7717/peerj.9634 Kaneko Y, Kanda E, Tashima S, Masuda R, Newman C, Macdonald DW (2014) The socio-spatial dynamics of the Japanese badger ( Meles anakuma ). J Mammal 95(2):290–300. https://doi.org/10.1644/12-MAMM-A-158 Kaneko Y, Maruyama N, Macdonald DW (2006) Food habits and habitat selection of suburban badgers ( Meles meles ) in Japan. J Zool 270(1):78–89. https://doi.org/10.1111/j.1469-7998.2006.00063.x Kawamura Y, Kamei T, Taruno H (1989) Middle and Late Pleistocene mammalian faunas in Japan. Quat Res 28(4):317–326. https://doi.org/10.4116/jaqua.28.317 Kholinne E, Zulkarnain RF, Sun YC, Lim S, Chun JM, Jeon IH (2018) The different role of each head of the triceps brachii muscle in elbow extension. Acta orthop traumatol turc 52(3):201–205. https://doi.org/10.1016/j.aott.2018.02.005 Kim SI, Oshida T, Lee H, Min MS, Kimura J (2015) Evolutionary and biogeographical implications of variation in skull morphology of raccoon dogs ( Nyctereutes procyonoides , Mammalia: Carnivora). Biol J Linn Soc 116(4):856–872. https://doi.org/10.1111/bij.12629 Koh HS, Kryukov A, Oh JG, Bayarkhagva D, Yang BG, Ahn NH, Bazarsad D (2014) Two sympatric phylogroups of the Asian badger Meles leucurus (Carnivora: Mammalia) identified by mitochondrial DNA cytochrome b gene sequences. Russ J Theriol 13(1):1–8. https://doi.org/10.15298/rusjtheriol.13.1.01 Kowalczyk R, Jȩdrzejewska B, Zalewski A (2003) Annual and circadian activity patterns of badgers ( Meles meles ) in Białowieża Primeval Forest (eastern Poland) compared with other Palaearctic populations. J Biogeogr 30(3):463–472. https://doi.org/10.1046/j.1365-2699.2003.00804.x Law CJ, Slater GJ, Mehta RS (2018) Lineage diversity and size disparity in Musteloidea: testing patterns of adaptive radiation using molecular and fossil-based methods. Syst Biol 67(1):127–144. https://doi.org/10.1093/sysbio/syx047 Lee MY, Lee SM, Song EG, An JH, Voloshina I, Chong JR, Johnson WE, Min M, Lee H (2016) Phylogenetic relationships and genetic diversity of badgers from the Korean Peninsula: Implications for the taxonomic status of the Korean badger. Biochem Syst Ecol 69:18–26. https://doi.org/10.1016/j.bse.2016.07.015 López-Martín JM, Ruiz-Olmo J, Padró I (2006) Comparison of skull measurements and sexual dimorphism between the Minorcan pine marten ( Martes martes minoricensis ) and the Iberian pine marten ( M. m. martes ): A case of insularity. Mamm Biol 71(1):13–24. https://doi.org/10.1016/j.mambio.2005.08.006 Lyras GA, van der Geer AA, Rook L (2010) Body size of insular carnivores: evidence from the fossil record. J Biogeogr 37(6):1007–1021. https://doi.org/10.1111/j.1365-2699.2010.02312.x Madurell-Malapeira J, Martínez-Navarro B, Ros-Montoya S, Espigares MP, Toro I, Palmqvist P (2011) The earliest European badger ( Meles meles ), from the late villafranchian site of Fuente Nueva 3 (Orce, granada, se iberian peninsula). Comptes Rendus Palevol 10(8):609–615. https://doi.org/10.1016/j.crpv.2011.06.001 Marmi J, Lopez-Giraldez F, Macdonald DW, Calafell F, Zholnerovskaya E, Domingo‐Roura X (2006) Mitochondrial DNA reveals a strong phylogeographic structure in the badger across Eurasia. Mol Ecol 15(4):1007–1020. https://doi.org/10.1111/j.1365-294X.2006.02747.x Marneweck CJ, Allen BL, Butler AR, Do Linh San E, Harris SN, Jensen AJ et al (2022) Middle-out ecology: small carnivores as sentinels of global change. Mamm Rev 52(4):471–479. https://doi.org/10.1111/mam.12300 Masseti M (1995) Quaternary biogeography of the Mustelidae family on the Mediterranean islands. HYSTRIX 7:17–34. https://doi.org/10.4404/hystrix-7.1-2-4048 Meiri S Dayan T, Simberloff D (2004) Body size of insular carnivores: little support for the island rule. The Am Nat 163(3):469–479. https://doi.org/10.1086/382229 Monroy-Cendales MJ, Vélez‐García JF, Castañeda‐Herrera FE (2020) Gross anatomy of the shoulder and arm intrinsic muscles in the white‐footed tamarin ( Saguinus leucopus –Günther, 1876): Inter- and intraspecific anatomical variations. J Med Primatol 49(3):123–135. https://doi.org/10.1111/jmp.12465 Monroy-Cendales MJ, Vélez‐García JF, Castañeda‐Serrano RD, Miglino MA (2023) Gross anatomical description of the extrinsic and intrinsic scapular and brachial muscles of Sapajus apella (Linnaeus, 1758). J Med Primatol 52(1):3–16. https://doi.org/10.1111/jmp.12619 Moore A (2011) Functional specialization in the intrinsic forelimb musculature of the American badger (Taxidea taxus) . Dissertation, Youngstown State University Moore AL, Budny JE, Russell AP, Butcher MT (2013) Architectural specialization of the intrinsic thoracic limb musculature of the American badger ( Taxidea taxus ). J Morphol 274(1):35–48. https://doi.org/10.1002/jmor.20074 Moore CM, Collins PW (1995) Urocyon littoralis . Mamm Spec 489:1–7. https://doi.org/10.2307/3504160 Nanova O, Prôa M (2017) Cranial features of mainland and Commander Islands (Russia) Arctic foxes ( Vulpes lagopus ) reflect their diverging foraging strategies. Polar Res 36:7 https://doi.org/10.1080/17518369.2017.1310976 Neal EG, Cheeseman CL (1996) Badgers. Poyster Natural History, London Oh JG (2007) Behavioral Characteristics of Jeju Badgers. Hallasan Research Institute Research Report:37–49 Perdomo-Cárdenas V, Patiño‐Holguín C, Vélez‐García JF (2021) Evolutionary and terminological analysis of the flexor digitorum superficialis, interflexorii and palmaris longus muscles in kinkajou ( Potos flavus ) and crab‐eating racoon ( Procyon cancrivorus ). Anat Histol Embryol 50(3):520–533. https://doi.org/10.1111/ahe.12656 Proulx G, Abramov AV, Adams I, Jennings AP, Khorozyan I, Rosalino LM et al (2016) World distribution and status of badgers–A review. In: Proulx G, Do Linh San, E (eds) Badgers: Systematics, Biology, Conservation and Research Techniques. Alpha Wildlife Publications, Canada, pp 31–116 Rose J, Moore A, Russell A, Butcher M (2014) Functional osteology of the forelimb digging apparatus of badgers. J Mammal 95(3):543–558. https://doi.org/10.1644/13-MAMM-A-174 Rozzi R, Lomolino MV, van der Geer AA, Silvestro D, Lyons SK, Bover P et al (2023) Dwarfism and gigantism drive human-mediated extinctions on islands. Science 379(6636):1054–1059. https://doi.org/10.1126/science.add8606 Salesa MJ, Anton M, Peigne S, Morales J (2008) Functional anatomy and biomechanics of the postcranial skeleton of Simocyon batalleri (Viret, 1929)(Carnivora, Ailuridae) from the Late Miocene of Spain. Zool J Linn Soc 152(3):593–621. https://doi.org/10.1111/j.1096-3642.2007.00370.x Salesa MJ, Siliceo G, Antón M, Fabre AC, Pastor JF (2020) Functional inferences on the long bones of Ischyrictis zibethoides (Blainville, 1841)(Carnivora, Mustelidae) from the middle Miocene locality of Sansan (Gers, France). Geodiversitas 42(1):1–16. https://doi.org/10.5252/geodiversitas2020v42a1 Samuels JX, Meachen JA, Sakai SA (2013) Postcranial morphology and the locomotor habits of living and extinct carnivorans. J Morphol 274(2):121–146. https://doi.org/10.1002/jmor.20077 Savvidou A, Youlatos D, Spassov N, Tamvakis A, Kostopoulos DS (2022) Ecomorphology of the Early Pleistocene Badger Meles dimitrius from Greece. J Mammal Evol 29(3):585–607. https://doi.org/10.1007/s10914-022-09609-5 Shikama T (1949) The Kuzuu Ossuaries, Geological and Palaeontological Studies of the Limestone Fissure Deposits in Kuzuu, Totigi Prefecture. Sci Rep Tohoku Uni II Ser 23: 1–209. Shikama T (1962) Quaternary Land Connections of Japanese Islands with Continent from the Viewpoints of Palaeomammalogy. Quat Res 2(4–5):146–153. https://doi.org/10.4116/jaqua.2.146 Sicuro FL, Oliveira LFB (2015) Variations in leopard cat ( Prionailurus bengalensis ) skull morphology and body size: sexual and geographic influences. PeerJ 3:e1309. https://doi.org/10.7717/peerj.1309 Sleeman DP, Davenport J (2016) Evidence of poor diet quality in the small-bodied Eurasian Badgers ( Meles meles ) on Rutland Island, Co. Donegal. Ir Nat J 35(1):22–26. Sleeman DP, Davenport J, Cussen RE, Hammond RF (2009) The small-bodied badgers ( Meles meles (L.)) of Rutland Island, Co. Donegal. Ir Nat J 30(1):1–6. Tanaka H (2006) Winter hibernation and body temperature fluctuation in the Japanese badger, Meles meles anakuma . Zool Sci 23(11):991–997. https://doi.org/10.2108/zsj.23.991 Tanaka H, Fukuda Y, Yuki E, Ota Y, Hosoi E, Kojima W (2022) Cooperative den maintenance between male Japanese badgers that are delayed dispersers and their mothers. J Ethol 40(1):3–11. https://doi.org/10.1007/s10164-021-00718-x Taylor ME (1974) The functional anatomy of the forelimb of some African Viverridae (Carnivora). J Morphol 143(3):307–335. https://doi.org/10.1002/jmor.1051430305 Van Valkenburgh B (1987) Skeletal indicators of locomotor behavior in living and extinct carnivores. J Vertebr Paleontol 7(2):162–182. https://doi.org/10.1080/02724634.1987.10011651 Vélez-García JF, Blanco DAC, Gómez GM, Cañas SSM (2023) Descriptive study of the intrinsic muscles of the shoulder and brachium in kinkajou ( Potos flavus ) and an evolutionary analysis within the suborder Caniformia. Vertebr Zool 73:957–980. https://doi.org/10.3897/vz.73.e102645 Vélez García JF, de Carvalho Barros RA, Miglino MA (2025) Is the articularis humeri muscle homologous to the coracobrachialis muscle in carnivorans? An evolutionary and terminological answer based on the shoulder myology of the coati ( Nasua nasua , Carnivora, Procyonidae). Anat, Histol, Embryol 54(2):e70034. https://doi.org/10.1111/ahe.70034 Wayne RK, George SB, Gilbert D, Collins PW, Kovach SD, Girman D, Lehman N (1991) A morphologic and genetic study of the island fox, Urocyon littoralis . Evolution 45(8):1849–1868. https://doi.org/10.1111/j.1558-5646.1991.tb02692.x Windle BCA, Parsons FG (1897) On the Myology of the Terrestrial Carnivora. Part I: muscles of the head, neck, and fore-limb. Proc Zool Soc Lond 65:370–409. https://doi.org/10.1111/j.1469-7998.1897.tb00023.x Yousefi MH, Rasouli B, Ghodrati S, Adibi MA, Taherdoost M, Omidbakhsh S, Behnam G (2018) Anatomical study of extrinsic and some intrinsic muscles of the thoracic limb in Iranian pine marten ( Martes martes ): a case report. Iran J Vet Med 12(3):273–282. https://doi.org/10.22059/ijvm.2018.252150.1004876 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 02 May, 2026 Read the published version in Journal of Mammalian Evolution → Version 1 posted Editorial decision: Revision requested 04 Aug, 2025 Reviews received at journal 01 Aug, 2025 Reviews received at journal 30 Jul, 2025 Reviewers agreed at journal 10 Jul, 2025 Reviews received at journal 10 Jul, 2025 Reviewers agreed at journal 10 Jul, 2025 Reviewers agreed at journal 09 Jul, 2025 Reviewers invited by journal 09 Jul, 2025 Editor assigned by journal 04 Jul, 2025 Submission checks completed at journal 04 Jul, 2025 First submitted to journal 03 Jul, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7041610","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":483724800,"identity":"de58f704-1fac-4f9e-8e45-3fbc1d851dfb","order_by":0,"name":"Emma Dangerfield","email":"","orcid":"","institution":"University of Tsukuba","correspondingAuthor":false,"prefix":"","firstName":"Emma","middleName":"","lastName":"Dangerfield","suffix":""},{"id":483724802,"identity":"da4a641d-cf4c-404a-97df-9e9a8c8e340f","order_by":1,"name":"Mao Shimoda","email":"","orcid":"","institution":"University of Tsukuba","correspondingAuthor":false,"prefix":"","firstName":"Mao","middleName":"","lastName":"Shimoda","suffix":""},{"id":483724803,"identity":"0d4343be-843d-497f-8b2d-f638de81b2f5","order_by":2,"name":"Sujoo Cho","email":"","orcid":"","institution":"Seoul National University","correspondingAuthor":false,"prefix":"","firstName":"Sujoo","middleName":"","lastName":"Cho","suffix":""},{"id":483724804,"identity":"cd15f8a9-713f-4380-bdb9-9cd408b76749","order_by":3,"name":"Yuri Kimura","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABDElEQVRIiWNgGAWjYDACHsYGFH4CP5gsIEWLJIifYIBPCxo/weAAiMKjxeDM4dbNvHvsGMzb2x8++LnHLs/4/OrEDw8MGOT5xQ5g13K2se02z7NkBpkzZ4wNe54lF5vdeLtZAugww5mzE7BrOc8I1HKAmUFCIodNgufAgcRtN85uAGlJMLiNV0s9g4T88+c//wC1bJ5xdvMPvFrADjtwGGgLgxkzyJYN/L3b8NoieeZg2805B47zSPDkGEvLHEhOnHGDd5tFgoEETr/wnUl/duPNgWo5CfbjDz++OWCX2N9/dvPNHxU28vzS2LUoHIDQSNEjAVYpgVU5CMg3YAjxH8CpehSMglEwCkYmAAAQvWdPRW933wAAAABJRU5ErkJggg==","orcid":"","institution":"National Museum of National Science in Tokyo","correspondingAuthor":true,"prefix":"","firstName":"Yuri","middleName":"","lastName":"Kimura","suffix":""}],"badges":[],"createdAt":"2025-07-03 22:53:03","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7041610/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7041610/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10914-025-09796-x","type":"published","date":"2026-05-02T15:58:38+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":86802293,"identity":"93d568f4-dd75-4554-aca2-eae0bee128d1","added_by":"auto","created_at":"2025-07-15 17:18:44","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4891250,"visible":true,"origin":"","legend":"\u003cp\u003eMap of Meles species distribution. Population boundaries are distinguished by colors, M. meles in red, M. canescens in orange, M. leucurus in blue, and M. anakuma in green. M. leucurus and M. anakuma specimens dissected in this study are numbered 1-7. Locations of museum collections of extant M. meles and M. canescens specimens used in this study are indicated by red and orange stars, respectively. Figure modified from Faggi et al. (2024) with the inclusion of Crete Island and Rhodes Island M. canescens populations and Jeju Island M. leucurus inhabitants.\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/8f597e95495850bcbdd9e254.png"},{"id":86802961,"identity":"f517f44b-b3da-4108-b71a-8a27ace79ba8","added_by":"auto","created_at":"2025-07-15 17:26:46","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":18649624,"visible":true,"origin":"","legend":"\u003cp\u003eSuperficial musculature of \u003cem\u003eMeles anakuma \u003c/em\u003ein lateral\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e),\u003cstrong\u003e \u003c/strong\u003eand medial\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003eb\u003c/strong\u003e)\u003cstrong\u003e \u003c/strong\u003eviews. Muscles are grouped by color and denoted by gradience. Abbreviations: \u003cstrong\u003eBB\u003c/strong\u003e, m. biceps brachii; \u003cstrong\u003eBCH\u003c/strong\u003e, m. brachialis; \u003cstrong\u003eBRAD\u003c/strong\u003e, m. brachioradialis;\u003cstrong\u003eDAc\u003c/strong\u003e, m. acromiodeltoideus;\u003cstrong\u003e DSp\u003c/strong\u003e, m. spinodeltoideus;\u003cstrong\u003e EDC\u003c/strong\u003e, m. extensor digitorum communis;\u003cstrong\u003e EDL\u003c/strong\u003e, m. extensor digitorum lateralis;\u003cstrong\u003e ERB\u003c/strong\u003e, m. extensor carpi radialis brevis;\u003cstrong\u003e ERL\u003c/strong\u003e, m. extensor carpi radialis longus;\u003cstrong\u003e EU\u003c/strong\u003e, m. extensor carpi ulnaris;\u003cstrong\u003e FCR\u003c/strong\u003e, m. flexor carpi radialis;\u003cstrong\u003e FCU\u003c/strong\u003e, m. flexor carpi ulnaris;\u003cstrong\u003e FDP\u003c/strong\u003e, m. flexor digitorum profundus;\u003cstrong\u003e ISp\u003c/strong\u003e, m. infraspinatus;\u003cstrong\u003e PL\u003c/strong\u003e, m. palmaris longus;\u003cstrong\u003e PT\u003c/strong\u003e, m. pronator teres;\u003cstrong\u003e SSc\u003c/strong\u003e, m. subscapuaris;\u003cstrong\u003e SSp\u003c/strong\u003e, m. supraspinatus;\u003cstrong\u003eTBAc\u003c/strong\u003e, m. triceps brachii caput accessorium;\u003cstrong\u003e TBAn\u003c/strong\u003e, m. triceps brachii caput angulare;\u003cstrong\u003e TBLa\u003c/strong\u003e, m. triceps brachii caput laterale;\u003cstrong\u003e TBLo\u003c/strong\u003e, m. triceps brachii caput longum;\u003cstrong\u003e TBMe\u003c/strong\u003e, m. triceps brachii caput mediale;\u003cstrong\u003eTMj\u003c/strong\u003e, m. teres major; \u003cstrong\u003eTFA\u003c/strong\u003e, m. tensor fasciae antebrachii. Scale bar equals 5 cm\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/21629d08ec6f586a2e6967dd.png"},{"id":86802295,"identity":"58ece024-240f-4eac-a629-963c6011171d","added_by":"auto","created_at":"2025-07-15 17:18:44","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":9491155,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eMeles anakuma\u003c/em\u003e muscle maps in the lateral (\u003cstrong\u003ea\u003c/strong\u003e), medial (\u003cstrong\u003eb\u003c/strong\u003e), and ventral view (\u003cstrong\u003ec\u003c/strong\u003e) of the scapula; posterior (\u003cstrong\u003ed\u003c/strong\u003e), and anterior (\u003cstrong\u003ee\u003c/strong\u003e) views of the radius. Abbreviations: \u003cstrong\u003eBB\u003c/strong\u003e, m. biceps brachii; \u003cstrong\u003eBCH\u003c/strong\u003e, m. brachialis; \u003cstrong\u003eCBH\u003c/strong\u003e, m. coracobrachialis; \u003cstrong\u003eDAc\u003c/strong\u003e, m. acromiodeltoideus; \u003cstrong\u003eDSp\u003c/strong\u003e, m. spinodeltoideus; \u003cstrong\u003eFDPr\u003c/strong\u003e, m. flexor digitorum profundus radial head;\u003cstrong\u003e ISp\u003c/strong\u003e, m. infraspinatus; \u003cstrong\u003ePT\u003c/strong\u003e, m. pronator teres; \u003cstrong\u003ePQ\u003c/strong\u003e, m. pronator quadratus;\u003cstrong\u003e SSc\u003c/strong\u003e, m. subscapuaris; \u003cstrong\u003eSSp\u003c/strong\u003e, m. supraspinatus; \u003cstrong\u003eSU\u003c/strong\u003e, m. supinator; \u003cstrong\u003eTBAn\u003c/strong\u003e, m. triceps brachii caput angulare;\u003cstrong\u003e TBLo\u003c/strong\u003e, m. triceps brachii caput longum; \u003cstrong\u003eTMj\u003c/strong\u003e, m. teres major; \u003cstrong\u003eTMn\u003c/strong\u003e, m. teres minor\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/48e19e7a2a577f4bce934c5a.png"},{"id":86802297,"identity":"e965f152-b97a-4ed8-bdd0-fdea13eac8b2","added_by":"auto","created_at":"2025-07-15 17:18:44","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":15222190,"visible":true,"origin":"","legend":"\u003cp\u003eIntrinsic muscular structure of m. coracobrachialis\u003cem\u003e \u003c/em\u003ein\u003cem\u003e Meles leucurus\u003c/em\u003e (\u003cstrong\u003ea\u003c/strong\u003e), and\u003cem\u003e M. anakuma \u003c/em\u003e(\u003cstrong\u003eb\u003c/strong\u003e)\u003cstrong\u003e \u003c/strong\u003ein the medial\u003cstrong\u003e \u003c/strong\u003eview. Muscle belly is indicated by coloured regions. Abbreviations: \u003cstrong\u003eCBH\u003c/strong\u003e, m. coracobrachialis; \u003cstrong\u003eTBAc\u003c/strong\u003e, m. triceps brachii caput accessorium;\u003cstrong\u003e TBLo\u003c/strong\u003e, m. triceps brachii caput longum; \u003cstrong\u003eTBMe\u003c/strong\u003e, m. triceps brachii caput mediale. Scale bar equals 5 cm\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/a447859f40ea9ff04f621d51.png"},{"id":86802298,"identity":"13fff1b1-2b24-4be7-b648-7335e6169412","added_by":"auto","created_at":"2025-07-15 17:18:45","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":14684552,"visible":true,"origin":"","legend":"\u003cp\u003eMuscle map of the distal arm in the anterior view for \u003cem\u003eMeles anakuma \u003c/em\u003e(\u003cstrong\u003ea\u003c/strong\u003e); m. tensor fasciae antebrachii in the medial view for \u003cem\u003eM. leucurus \u003c/em\u003e(\u003cstrong\u003eb\u003c/strong\u003e). Abbreviations: \u003cstrong\u003eAD\u003c/strong\u003e, m. abductor digiti I; \u003cstrong\u003eBB\u003c/strong\u003e, m. biceps brachii; \u003cstrong\u003eBCH\u003c/strong\u003e, m. brachialis; \u003cstrong\u003eBRAD\u003c/strong\u003e, m. brachioradialis; \u003cstrong\u003eED\u003c/strong\u003e, m. extensor digiti I and II; \u003cstrong\u003eEDC\u003c/strong\u003e, m. extensor digitorum communis; \u003cstrong\u003eEDL\u003c/strong\u003e, m. extensor digitorum lateralis; \u003cstrong\u003eERB\u003c/strong\u003e, m. extensor carpi radialis brevis; \u003cstrong\u003eERL\u003c/strong\u003e, m. extensor carpi radialis longus; \u003cstrong\u003eEU\u003c/strong\u003e, m. extensor carpi ulnaris; \u003cstrong\u003eFCR\u003c/strong\u003e, m. flexor carpi radialis; \u003cstrong\u003eFDP\u003c/strong\u003e, m. flexor digitorum profundus; \u003cstrong\u003ePT\u003c/strong\u003e,\u003cem\u003e \u003c/em\u003em. pronator teres; \u003cstrong\u003eTBLo\u003c/strong\u003e, m. triceps brachii caput longum; \u003cstrong\u003eTBMe\u003c/strong\u003e, m. triceps brachii caput mediale; \u003cstrong\u003eTFA\u003c/strong\u003e, m. tensor fasciae antebrachii. Scale bar equals 2.5 cm\u003c/p\u003e","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/4924f8daf3d434e681336d51.png"},{"id":86802302,"identity":"06375262-514c-489b-8e0d-b5c3856f789e","added_by":"auto","created_at":"2025-07-15 17:18:45","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":24998137,"visible":true,"origin":"","legend":"\u003cp\u003eDeep musculature (depth direction indicated with an arrow) of m. palmaris longus\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e), m. flexor digitorum superficialis (\u003cstrong\u003eb\u003c/strong\u003e),\u003cstrong\u003e \u003c/strong\u003ethe five heads of m. flexor digitorum profundus\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003ec\u003c/strong\u003e),\u003cstrong\u003e \u003c/strong\u003em. pronator quadratus and m. flexor carpi radialis\u003cem\u003e \u003c/em\u003e(\u003cstrong\u003ed\u003c/strong\u003e); the deep distal arm muscles of \u003cem\u003eMeles anakuma\u003c/em\u003e in the lateral view (\u003cstrong\u003ee\u003c/strong\u003e). Abbreviations: \u003cstrong\u003eAD\u003c/strong\u003e, m. abductor digiti I; \u003cstrong\u003eANC\u003c/strong\u003e, m. anconeus; \u003cstrong\u003eBB\u003c/strong\u003e, m. biceps brachii;\u003cstrong\u003e BCH\u003c/strong\u003e, m. brachialis; \u003cstrong\u003eBRAD\u003c/strong\u003e, m. brachioradialis;\u003cstrong\u003e ED\u003c/strong\u003e, m. extensor digiti I and II;\u003cstrong\u003e FCR\u003c/strong\u003e, m. flexor carpi radialis;\u003cstrong\u003e FCU\u003c/strong\u003e, m. flexor carpi ulnaris;\u003cstrong\u003e FDP\u003c/strong\u003e, m. flexor digitorum profundus; \u003cstrong\u003eFDS\u003c/strong\u003e, m. flexor digitorum superficialis;\u003cstrong\u003e ISp\u003c/strong\u003e, m. infraspinatus;\u003cstrong\u003e PL\u003c/strong\u003e, m. palmaris longus; \u003cstrong\u003ePT\u003c/strong\u003e,\u003cem\u003e \u003c/em\u003em. pronator teres\u003cem\u003e; \u003c/em\u003e\u003cstrong\u003ePQ\u003c/strong\u003e, m. pronator quadratus; \u003cstrong\u003eSSp\u003c/strong\u003e, m. supraspinatus; \u003cstrong\u003eSU\u003c/strong\u003e, m. spinator; \u003cstrong\u003eTBAc\u003c/strong\u003e, m. triceps brachii caput accessorium;\u003cem\u003e \u003c/em\u003e\u003cstrong\u003eTMj\u003c/strong\u003e, m. teres major; \u003cstrong\u003eTMn\u003c/strong\u003e, m. teres minor. M. flexor digitorum profundus heads are expressed as: h\u003csub\u003ep\u003c/sub\u003e, humeral profundus; h\u003csub\u003em\u003c/sub\u003e, humeral medial; h\u003csub\u003el\u003c/sub\u003e, humeral laterale; u, ulnare; r, radial. Broken lines indicate location of a muscle hidden from view. Scale bar equals 2.5 cm\u003c/p\u003e","description":"","filename":"Fig6.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/5e7341bc231300fd02fb0c13.png"},{"id":86802306,"identity":"c4677dc1-68bc-4dc7-82ae-9da0322c11c3","added_by":"auto","created_at":"2025-07-15 17:18:45","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":19696675,"visible":true,"origin":"","legend":"\u003cp\u003eMuscular maps on the humerus (\u003cstrong\u003ea-e\u003c/strong\u003e) and ulna (\u003cstrong\u003ef-j\u003c/strong\u003e) in \u003cem\u003eMeles anakuma \u003c/em\u003ein the cranial, lateral, medial, dorsal, and caudal views. Muscles are grouped by color gradients, and individual bellies are further distinguished by patterns. Abbreviations: \u003cstrong\u003eAD\u003c/strong\u003e, m. abductor digiti I; \u003cstrong\u003eANC\u003c/strong\u003e, m. anconeus; \u003cstrong\u003eBCH\u003c/strong\u003e, m. brachialis; \u003cstrong\u003eBRAD\u003c/strong\u003e, m. brachioradialis;\u003cstrong\u003e CBH\u003c/strong\u003e, m. coracobrachialis;\u003cstrong\u003eDAc\u003c/strong\u003e, m. acromiodeltoideus;\u003cstrong\u003e DSp\u003c/strong\u003e, m. spinodeltoideus;\u003cstrong\u003e ED\u003c/strong\u003e, m. extensor digiti I and II; \u003cstrong\u003eEDC\u003c/strong\u003e, m. extensor digitorum communis; \u003cstrong\u003eEDL\u003c/strong\u003e, m. extensor digitorum lateralis;\u003cstrong\u003e ERB\u003c/strong\u003e, m. extensor carpi radialis brevis;\u003cstrong\u003eERL\u003c/strong\u003e, m. extensor carpi radialis longus; \u003cstrong\u003eEU\u003c/strong\u003e, m. extensor carpi ulnaris;\u003cstrong\u003e FCR\u003c/strong\u003e, m. flexor carpi radialis; \u003cstrong\u003eFCU\u003c/strong\u003e, m. flexor carpi ulnaris;\u003cstrong\u003e ISp\u003c/strong\u003e, m. infraspinatus;\u003cstrong\u003e PL\u003c/strong\u003e, m. palmaris longus; \u003cstrong\u003ePT\u003c/strong\u003e,\u003cem\u003e \u003c/em\u003em. pronator teres; \u003cstrong\u003ePQ\u003c/strong\u003e, m. pronator quadratus; \u003cstrong\u003eSSc\u003c/strong\u003e, m. subscapuaris; \u003cstrong\u003eSSp\u003c/strong\u003e, m. supraspinatus; \u003cstrong\u003eTBAc\u003c/strong\u003e, m. triceps brachii caput accessorium; \u003cstrong\u003eTBAn\u003c/strong\u003e, m. triceps brachii caput angulare; \u003cstrong\u003eTBLa\u003c/strong\u003e, m. triceps brachii caput laterale; \u003cstrong\u003eTBLo\u003c/strong\u003e, m. triceps brachii caput longum; \u003cstrong\u003eTBMe\u003c/strong\u003e, m. triceps brachii caput mediale; \u003cstrong\u003eTMj\u003c/strong\u003e, m. teres major; \u003cstrong\u003eTMn\u003c/strong\u003e, m. teres minor. M. flexor digitorum profundus heads are expressed as: FDPh, humeral; FDPu, ulnare, and FDPr, radial. (+) indicates muscle fusion prior to origin or insertion point\u003c/p\u003e","description":"","filename":"Fig7.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/e4e7fc833bb628fd804d1ddc.png"},{"id":86802308,"identity":"c093c249-1a56-4c89-ba02-a61b52e3465f","added_by":"auto","created_at":"2025-07-15 17:18:45","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":10495245,"visible":true,"origin":"","legend":"\u003cp\u003ePairwise comparisons of the humerus\u003cstrong\u003e \u003c/strong\u003ein the caudal, medial and distal views\u003cstrong\u003e, \u003c/strong\u003eand ulna in the medial and cranial views,\u003cstrong\u003e \u003c/strong\u003efor each extant \u003cem\u003eMeles\u003c/em\u003e species. \u003cstrong\u003ea\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003eand \u003cstrong\u003ee\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eM. meles \u003c/em\u003efrom Spain MCNG-19070501; \u003cstrong\u003eb\u003c/strong\u003e. and \u003cstrong\u003ef\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eM. canescens \u003c/em\u003efrom Crete NMC-80.5.63.6; \u003cstrong\u003ec\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003eand \u003cstrong\u003eg\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eM. leucurus \u003c/em\u003efrom South Korea #7; \u003cstrong\u003ed\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003eand \u003cstrong\u003eh\u003c/strong\u003e.\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eM. anakuma \u003c/em\u003efrom Japan POM-256. Arrows indicate dispersion direction. Key osteological features are colored red for \u003cem\u003emeles-canescens\u003c/em\u003e, blue for \u003cem\u003eleucurus-anakuma\u003c/em\u003e, and purple for shared features present in both groups. The scale equals 5 cm\u003c/p\u003e","description":"","filename":"Fig8.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/37b5dc9880eb967d8924f5da.png"},{"id":86802312,"identity":"fd580569-3991-474d-984f-7a3c74848161","added_by":"auto","created_at":"2025-07-15 17:18:45","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":2487375,"visible":true,"origin":"","legend":"\u003cp\u003eA simplified phylogenetic overview of the major osteological and myological changes that occur between continental species \u003cem\u003eMeles meles\u003c/em\u003eand \u003cem\u003eM. leucurus\u003c/em\u003e, and insular populations of \u003cem\u003eM. canescens\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e. (*) distinguishes the \u003cem\u003eM. canescens\u003c/em\u003e population used in this study, and insular species are indicated with an island symbol. Shared features are highlighted in bold, and badger silhouettes signify changes in body size. Arrows indicate the direction of dispersion from continental to insular populations. Nodes in bold indicate dispersion events of \u003cem\u003eMeles\u003c/em\u003e while broken gray lines show major myological changes that occurred prior to and preceding \u003cem\u003eMeles \u003c/em\u003ewith (?) to indicate dubious trends. Phylogenetic relationships are based on mtDNA analyses detailed in Marmi et al. (2006) and myological trends are from Ercoli et al. (2015)\u003c/p\u003e","description":"","filename":"Fig9.png","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/f4d51bdc706f94c674dd3517.png"},{"id":108498100,"identity":"70e4f956-7084-47ac-b3eb-64dca2a8561d","added_by":"auto","created_at":"2026-05-05 10:14:06","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":120919860,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7041610/v1/ca4fa2a0-bf97-4a50-87a5-54ba775f24ac.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Enhanced Forelimb Mobility in Insular Meles populations: Insights from Myological and Osteological Comparisons","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe genus \u003cem\u003eMeles\u003c/em\u003e, comprising of four extant species, \u003cem\u003eMeles meles\u003c/em\u003e Linnaeus, 1758 (European badger), \u003cem\u003eMeles leucurus\u003c/em\u003e Hodgson, 1847 (Asian badger), \u003cem\u003eMeles anakuma\u003c/em\u003e Temminck, 1844 (Japanese badger), and the more recently redefined \u003cem\u003eMeles canescens\u003c/em\u003e Blanford, 1875 (Caucasian badger), is widely distributed across Eurasia, from Europe to Eastern Asia and the Japanese Archipelago (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) (Neal and Cheeseman \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Del Carro et al. 2010; Abramov and Puzachenko \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). While the evolutionary history of these taxa have been documented through mitochondrial DNA and cranial morphology (Marmi et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Koh et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Lee et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Law et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Faggi et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), much less is known about how island colonization has shaped the functional morphology of their forelimbs, particularly in insular populations such as \u003cem\u003eM. anakuma\u003c/em\u003e in Japan.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIsland environments are known to exert unique selective pressures on mammals, often resulting in shifts in body size. According to the \u0026lsquo;Island Rule\u0026rsquo;, large mammals tend to undergo dwarfism, while small mammals exhibit gigantism upon colonizing islands (Foster \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1964\u003c/span\u003e; Meiri et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Hayashi et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Rozzi et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, this pattern is often less pronounced or predictable for medium-sized mesocarnivores (Meiri et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ben\u0026iacute;tez-L\u0026oacute;pez et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). For instance, gigantism is evident in the island subspecies of Minorcan pine marten (\u003cem\u003eMartes martes minoricensis\u003c/em\u003e) (L\u0026oacute;pez-Mart\u0026iacute;n et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), in some insular otters (e.g. \u003cem\u003eMegalenhydris barbaricina\u003c/em\u003e) (Lyras et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and the Arctic fox (\u003cem\u003eVulpes lagopus\u003c/em\u003e) (Nanova and Pr\u0026ocirc;a \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), while dwarfism was recorded for the island populations of the pygmy raccoon (\u003cem\u003eProcyon pygmaeus\u003c/em\u003e) (Mcfaddon and Meiri 2012), the raccoon dog (\u003cem\u003eNyctereutes procionoides\u003c/em\u003e) (Kim et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), the leopard cat (\u003cem\u003ePrionailurus bengalensi\u003c/em\u003e) (Sicuro and Oliveira \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), and several fox species (e.g \u003cem\u003eUrocyon littoralis\u003c/em\u003e) (Moore and Collins \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Beyond body size, insular environments often influence behavior, ecology, and functional morphology. For example, field observations show the Channel Islands foxes have tendencies towards more arboreal activities and increased diurnal behavior likely as a response to ecological restraints (Wayne et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Moore and Collins \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Coonan et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), while changes in cranial features in insular artic foxes reflect modified foraging strategies (Nanova and Pr\u0026ocirc;a \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Similar ecological shifts have been found in \u003cem\u003eMeles\u003c/em\u003e where island populations of \u003cem\u003eM. meles\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e have shown altered diets and behaviors, likely driven by restricted resources (Oh \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Sleeman et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sleeman and Davenport \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). These ecological differences are supported by recent morphological studies which report cranial and dental adaptations in insular \u003cem\u003eMeles\u003c/em\u003e tax \u0026ndash; including \u003cem\u003eM. canescens\u003c/em\u003e from Crete and \u003cem\u003eM. anakuma\u003c/em\u003e \u0026ndash; such as increased interorbital distance, enhanced masticatory features and greater bite force (Savvidou et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These traits are associated with shifts towards increased diurnal activity and more carnivorous diets. Nonetheless, these adaptations do not always coincide with significant changes in body size, suggesting that other morphological adjustments may be more crucial for ecological success on islands.\u003c/p\u003e\u003cp\u003eRecent research emphasizes the value of studying medium-sized carnivores, including badgers, due to their high adaptability to rapid environmental changes (Marneweck et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This make \u003cem\u003eMeles\u003c/em\u003e a compelling case for investigating how island environments influence morphological adaptations. In particular, given the subfossorial lifestyle of badgers and their reliance on powerful forelimbs for digging and foraging (Moore \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Hildebrand and Goslow \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), investigating forelimb myology and osteology are key to understanding functional responses to insular ecological pressures. Forelimb adaptations such as enhanced maneuverability or increased force production may compensate for the absence of major body size shifts, providing a functional strategy for unique ecological pressures in insular settings.\u003c/p\u003e\u003cp\u003eHere, we provide the first detailed comparative myological analysis of the forelimbs in the insular \u003cem\u003eM. anakuma\u003c/em\u003e and its continental counterpart \u003cem\u003eM. leucurus\u003c/em\u003e, complemented by osteological comparisons with the European \u003cem\u003eM. meles\u003c/em\u003e and an insular population of \u003cem\u003eM. canescens\u003c/em\u003e from the Island of Crete. By integrating these anatomical observations, we aim to identify key musculoskeletal adaptations that have evolved in response to island-specific ecological demands.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eDissections\u003c/span\u003e\u003c/p\u003e\u003cp\u003eWe believe that no myological studies have been published on either \u003cem\u003eM. anakuma\u003c/em\u003e, a badger species endemic to Japan, or on any \u003cem\u003eM. leucurus\u003c/em\u003e individuals inhabiting South Korea. In this study, we dissected the forelimbs of four \u003cem\u003eM. anakuma\u003c/em\u003e specimens and three \u003cem\u003eM. leucurus\u003c/em\u003e specimens and analyzed their respective muscular structures (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All specimens were procured as roadkill victims and dissections were conducted at the NMNS Research center in Tsukuba for \u003cem\u003eM. anakuma\u003c/em\u003e, and at both Seoul National University (SNU) and the Wildlife Rescue Center in Ulsan City for \u003cem\u003eM. leucurus\u003c/em\u003e. For each badger specimen, the fur, organs and fatty tissues were carefully removed with scalpels and surgical scissors. Then the origin, insertion and morphological features of 28 forelimb intrinsic muscles were systematically identified and described (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Photographs were taken with a Canon EOS Kiss X9i (Canon Inc., Japan) to document the dissection process. Anatomical and osteological descriptions were based on terminology from Fishbeck and Sebastiani (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), Evans et al. (2012), Ercoli et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e)d hmer et al. (2020). Individual muscles were traced over dissection photographs in the program Procreate version 5.2.8 (Savage Interactive Pty Ltd., Australia), then separated first by color gradients for each muscle group (shoulder, brachium, distal arm, and deep) and then by patterns for the individual muscle heads.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eList of \u003cem\u003eMeles\u003c/em\u003e specimens with locality, sex, and humerus length. Abbreviations: \u003cb\u003eF\u003c/b\u003e, female; \u003cb\u003eM\u003c/b\u003e, male; \u003cb\u003eNA\u003c/b\u003e, not available. For museum abbreviations, see the main text\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMuseum/\u003c/p\u003e\u003cp\u003eInstitute\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSpecimen ID\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eID No. for this study\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eMeles\u003c/em\u003e\u003c/p\u003e\u003cp\u003especies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLocality\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eSex\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eHumerus length (mm)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNMNS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePOM-256\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. anakuma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eFukushima, Japan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e95.55\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNMNS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePOM-259\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. anakuma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHiroshima, Japan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e94.56\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNMNS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePOM-258\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. anakuma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eKumamoto, Japan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e75.85\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNMNS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePOM-260\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. anakuma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eKagoshima, Japan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e98.59\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSNU\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. leucurus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSeoul, South Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e86.09\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSNU\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. leucurus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSeoul, South Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e97.76\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSNU\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. leucurus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eUlsan, South Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e92.31\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNHMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80.5.63.19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. canescens\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCrete, Greece\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e84.57\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNHMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80.5.63.43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. canescens\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCrete, Greece\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e98.89\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNHMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80.5.63.46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. canescens\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCrete, Greece\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e99.78\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNHMC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80.5.63.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. canescens\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCrete, Greece\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e99.15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMCNG\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e19070501\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. meles\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLa Roca del Vall\u0026egrave;s, Spain\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e114.51\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMCNG\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e98042801-7507\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. meles\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLli\u0026ccedil;\u0026agrave; d'Amunt, Spain\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e116.15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMCNG\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e99031501\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. meles\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMieres, Spain\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e109.07\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMCNG\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e001022301\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. meles\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBatet de la Serra, Spain\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e108.26\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eList of the origin, insertion and main locomotive function of each muscle analysed in this study. Anatomical and osteological descriptions are based on terminology from Fishbeck and Sebastiani (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), Evans et al. (2012), Ercoli et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e)d hmer et al. (2020).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNo.\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMuscle name\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAbbr.\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOrigin\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eInsertion\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eMain function\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eShoulder\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eMm. supraspinatus\u003c/b\u003e\u003c/p\u003e\u003cp\u003e- M. supraspinatus principle\u003c/p\u003e\u003cp\u003e- M. supraspinatus intermediate\u003c/p\u003e\u003cp\u003e- M. supraspinatus cranial\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSSp\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSupraspinous fossa, scapular spine\u003c/p\u003e\u003cp\u003eCranial aspect of the scapular spine\u003c/p\u003e\u003cp\u003eCranial aspect of supraspinous fossa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHumeral greater tubercle\u003c/p\u003e\u003cp\u003eHumeral greater tubercle\u003c/p\u003e\u003cp\u003eHumeral great tubercle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder extension, humeral protractor\u003c/p\u003e\u003cp\u003eShoulder extension, humeral protractor\u003c/p\u003e\u003cp\u003eShoulder extension, humeral protractor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e2\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. infraspinatus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eISp\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eInfraspinous fossa, scapular spine\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHumeral greater tubercle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder flexion, humeral rotation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e3\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. subscapuaris\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSSc\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSubscapular fossa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHumeral lesser tubercle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eScapular adductor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e4\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. teres major\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTMj\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAxillary border of scapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMedial margin of pectoral ridge\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder flexion, limb retraction\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e5\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. teres minor\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTMn\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAxillary border of scapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHumeral greater tubercle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder flexion, humeral rotation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e6\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eMm. deltoideus\u003c/b\u003e\u003c/p\u003e\u003cp\u003e- M. spinodeltoideus\u003c/p\u003e\u003cp\u003e- M. acromiodeltoideus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDSp\u003c/p\u003e\u003cp\u003eDAc\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eScapular spine\u003c/p\u003e\u003cp\u003eAcromion\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDeltoid crest of humerus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder flexion and humeral abductor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eBrachium\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e7\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. coracobrachialis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCBH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCoracoid process of scapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHumeral diaphysis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder stabilizer and humeral adductor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e8\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. brachialis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBCH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProximal humerus diaphysis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCoronoid process of ulna\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow flexion and forearm supinator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e9\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. brachioradialis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBRAD\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProximal aspect of humeral epicondylar crest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDistal aspect of radius\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow flexion, forearm supinator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e10\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. biceps brachii\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eTubercle of scapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBicipital tuberosity of radius\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eShoulder extension, elbow flexion\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e11\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eMm. triceps brachii\u003c/b\u003e\u003c/p\u003e\u003cp\u003e- M. triceps brachii caput laterale\u003c/p\u003e\u003cp\u003e- M. triceps brachii caput longum\u003c/p\u003e\u003cp\u003e- M. triceps brachii caput mediale\u003c/p\u003e\u003cp\u003e- M. triceps brachii caput accessorium\u003c/p\u003e\u003cp\u003e- M. triceps brachii caput angulare\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTBLa\u003c/p\u003e\u003cp\u003eTBLo\u003c/p\u003e\u003cp\u003eTBMe\u003c/p\u003e\u003cp\u003eTBAc\u003c/p\u003e\u003cp\u003eTBAn\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProximal surface of deltoid crest\u003c/p\u003e\u003cp\u003eMid-axillary border of scapula\u003c/p\u003e\u003cp\u003eMediocaudal humeral diaphysis\u003c/p\u003e\u003cp\u003eDistal facet of humeral diaphysis\u003c/p\u003e\u003cp\u003eAxillary edge of scapula\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLateral aspect of ulnar olecranon\u003c/p\u003e\u003cp\u003eCaudal aspect of ulnar olecranon\u003c/p\u003e\u003cp\u003eMedial aspect of ulnar olecranon\u003c/p\u003e\u003cp\u003eMedial aspect of ulnar olecranon\u003c/p\u003e\u003cp\u003eCaudal aspect of ulnar olecranon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow extension\u003c/p\u003e\u003cp\u003eElbow extension, shoulder flexion\u003c/p\u003e\u003cp\u003eElbow extension\u003c/p\u003e\u003cp\u003eElbow extension\u003c/p\u003e\u003cp\u003eHumeral retractor, elbow extension\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e12\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. tensor fasciae antebrachii\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTFA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCaudal border of \u003cem\u003em. latissi dorsi\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMedial aspect of ulnar olecranon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow extension\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e13\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. anconeus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eANC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eDistal epicondylar crest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eLateral section of olecranon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow extension, forearm pronator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eDistal arm\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e14\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e15\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e16\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e17\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e18\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e19\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eExtensors\u003c/b\u003e\u003c/p\u003e\u003cp\u003eM. extensor carpi radialis longus\u003c/p\u003e\u003cp\u003eM. extensor carpi radialis brevis\u003c/p\u003e\u003cp\u003eM. extensor digitorum communis\u003c/p\u003e\u003cp\u003eM. extensor digitorum lateralis\u003c/p\u003e\u003cp\u003eM. extensor carpi ulnaris\u003c/p\u003e\u003cp\u003e- humeral head\u003c/p\u003e\u003cp\u003e- ulnar head\u003c/p\u003e\u003cp\u003eM. extensor digiti I and II\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eERL\u003c/p\u003e\u003cp\u003eERB\u003c/p\u003e\u003cp\u003eEDC\u003c/p\u003e\u003cp\u003eEDL\u003c/p\u003e\u003cp\u003eEU\u003c/p\u003e\u003cp\u003eED\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProximal aspect of humerus epicondylar crest\u003c/p\u003e\u003cp\u003eProximal aspect of humerus epicondylar crest\u003c/p\u003e\u003cp\u003eProximal aspect of humerus epicondylar crest\u003c/p\u003e\u003cp\u003eProximal aspect of humerus epicondylar crest\u003c/p\u003e\u003cp\u003eHumeral epicondylar crest\u003c/p\u003e\u003cp\u003eUlnar and radial shaft\u003c/p\u003e\u003cp\u003eDorsal aspect of ulnar shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBase of metacarpal II\u003c/p\u003e\u003cp\u003eBase of metacarpal III\u003c/p\u003e\u003cp\u003eDistal phalanges of digits II-V\u003c/p\u003e\u003cp\u003eDistal phalanges of digits III-V\u003c/p\u003e\u003cp\u003eBase of metacarpal V\u003c/p\u003e\u003cp\u003eDistal phalanges of digits I-II\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eElbow flexion, wrist extension\u003c/p\u003e\u003cp\u003eElbow flexion, wrist and digit extension\u003c/p\u003e\u003cp\u003eElbow flexion, wrist and digit extension\u003c/p\u003e\u003cp\u003eElbow flexion, wrist and digit extension\u003c/p\u003e\u003cp\u003eElbow flexion, wrist extension\u003c/p\u003e\u003cp\u003eExtensor of digits I-II\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e20\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e21\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e22\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e23\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eFlexors\u003c/b\u003e\u003c/p\u003e\u003cp\u003eM. flexor digitorum profundus\u003c/p\u003e\u003cp\u003e- humeral heads; profundus, mediale, and laterale\u003c/p\u003e\u003cp\u003e- ulnar head\u003c/p\u003e\u003cp\u003e- radial head\u003c/p\u003e\u003cp\u003eM. flexor carpi radialis\u003c/p\u003e\u003cp\u003eM. flexor carpi ulnaris\u003c/p\u003e\u003cp\u003eM. flexor digitorum superficialis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eFDP\u003c/p\u003e\u003cp\u003eFDPh\u003csub\u003ep\u003c/sub\u003e\u003c/p\u003e\u003cp\u003eFDPh\u003csub\u003em\u003c/sub\u003e\u003c/p\u003e\u003cp\u003eFDPh\u003csub\u003el\u003c/sub\u003e\u003c/p\u003e\u003cp\u003eFDPu\u003c/p\u003e\u003cp\u003eFDPr\u003c/p\u003e\u003cp\u003eFCR\u003c/p\u003e\u003cp\u003eFCU\u003c/p\u003e\u003cp\u003eFDS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCaudomedial aspect of the:\u003c/p\u003e\u003cp\u003ehumeral epicondyle,\u003c/p\u003e\u003cp\u003eulnar,\u003c/p\u003e\u003cp\u003eand radial shafts\u003c/p\u003e\u003cp\u003eEpicondyle of humerus\u003c/p\u003e\u003cp\u003eMedial aspect of olecranon and epicondyle\u003c/p\u003e\u003cp\u003eSuperficial to distal FDP humeral heads\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDistal phalanges of digits II-V\u003c/p\u003e\u003cp\u003eBase of metacarpal II and III\u003c/p\u003e\u003cp\u003eSesamoid proximal to metacarpal V\u003c/p\u003e\u003cp\u003eSuperficial to FDP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eWrist and digital flexion\u003c/p\u003e\u003cp\u003eWrist flexion\u003c/p\u003e\u003cp\u003eWrist flexion\u003c/p\u003e\u003cp\u003eWrist flexion\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e24\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. palmaris longus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eDistal epicondyle of humerus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDistal phalanges of digits II-V\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eWrist and digit flexion\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e25\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. abductor digiti I\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAD\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLateral ulnar and radial shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMetacarpal I\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eForearm abductor\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003eDeep\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e26\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. pronator teres\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePT\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMedial epicondyle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMedial section of radius shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eForearm pronator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e27\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. supinator\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSU\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLateral epicondyle\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMedial section of radius shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eForearm supinator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e28\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eM. pronator quadratus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePQ\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMedial ulnar shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDistal radial shaft\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eForearm pronator\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eComparative myology and osteology\u003c/span\u003e\u003c/p\u003e\u003cp\u003eThe anatomical structure of \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e were compared to the published myological descriptions of five mustelid species: \u003cem\u003eMeles meles\u003c/em\u003e (B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), two ictonychines (\u003cem\u003eGalictis cuja\u003c/em\u003e and \u003cem\u003eIctonyx striatus\u003c/em\u003e) (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Windle and Parsons \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e1897\u003c/span\u003e), and two guloines (\u003cem\u003eMartes martes\u003c/em\u003e and \u003cem\u003eMartes foina\u003c/em\u003e) (B\u0026ouml;hmer et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yousefi et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), as well as two procynids: \u003cem\u003ePotos flavus\u003c/em\u003e (V\u0026eacute;lez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and \u003cem\u003eProcyon lotor\u003c/em\u003e (Allen \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1882\u003c/span\u003e; Feeney \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). Melinae (to which \u003cem\u003eMeles\u003c/em\u003e belongs), Ictonychinae, and Guloinae are all closely-related subfamilies within Mustelidae, whereas Procynidae is a separate family adjacent to Mustelidae but still within the same order, Carnivora (Law et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Therefore, they share fundamental anatomical structures yet exhibit a range of forelimb functional specializations \u0026ndash; including arboreal and generalist behaviors \u0026ndash; which provide a valuable comparative framework for interpreting osteological data. For \u003cem\u003eM. canescens\u003c/em\u003e, no published myological data was available and so, only osteological comparisons were conducted on specimens from the insular population on Crete. The 3D models for comparative specimens were generated using an EinScanSP 3D scanner (Shining 3D V3.1.0.1, China) and visually analysed in the program 3D Slicer V5.0.3 (Fedorov et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eContinental-insular pairwise comparisons and dispersion history of\u003c/span\u003e \u003cspan type=\"ItalicUnderline\" class=\"ItalicUnderline\" name=\"Emphasis\"\u003eMeles\u003c/span\u003e\u003c/p\u003e\u003cp\u003eIn this study, we acknowledge that \u003cem\u003eM. canescens\u003c/em\u003e has both continental and insular populations. However, for the continental-insular pairwise comparison analysis, we chose \u003cem\u003eM. meles\u003c/em\u003e as the mainland counterpart to \u003cem\u003eM. canescens\u003c/em\u003e individuals from the island population on Crete because \u003cem\u003eM. meles\u003c/em\u003e is known from mDNA to be ancestral to \u003cem\u003eM. canescens\u003c/em\u003e (Marmi et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), and since specimens of continental \u003cem\u003eM. canescens\u003c/em\u003e were not accessible to us. This continental-insular comparison method has been previously successfully applied to \u003cem\u003eM. canescens\u003c/em\u003e from Crete, revealing several shared cranial and dentition traits that reflect adaptations to island environments (Abramov and Puzachenko \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Savvidou et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Here, we are working under the assumption that despite the shorter isolation period on Crete, \u003cem\u003eM. canescens\u003c/em\u003e populations would still have undergone morphological transformation that distinguishes them from the mainland inhabitants. Following the evolutionary history of \u003cem\u003eMeles\u003c/em\u003e summarized below, we grouped the genus into the following continental-insular pairs for comparative analysis: \u003cem\u003eleucurus-anakuma\u003c/em\u003e and \u003cem\u003emeles-canescens\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eGeographically, \u003cem\u003eM. meles\u003c/em\u003e occupies much of Europe and the British Isles, while \u003cem\u003eM. canescens\u003c/em\u003e ranges across southwestern Asia (Turkey, the Caucasus, Iran, etc.) and has established insular populations on the Mediterranean islands of Crete and Rhodes (Proulx et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). \u003cem\u003eM. leucurus\u003c/em\u003e inhabits most of mainland China, Russia, and Korea, while \u003cem\u003eM. anakuma\u003c/em\u003e is endemic to the Japanese archipelago, excluding Hokkaido and the Ryukyu Islands (Proulx et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Palaeontological evidence suggests that \u003cem\u003eMeles\u003c/em\u003e evolved from the genus \u003cem\u003eMelodon\u003c/em\u003e (Neel and Cheeseman 1996), where the earliest known \u003cem\u003eMeles\u003c/em\u003e fossil specimens found in Eurasia have been dated to the Late Pliocene (Madurell-Malapeira et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Faggi et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). An initial divergence occurred between European (\u003cem\u003eM. meles\u003c/em\u003e and \u003cem\u003eM. canescens\u003c/em\u003e) and Asian (\u003cem\u003eM. leucurus\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e) lineages during the Early Pleistocene, consistent in molecular and morphological studies (Marmi et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Del Carro et al. 2010; Law et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Abramov and Puzachenko \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Faggi et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the molecular study, the divergence between \u003cem\u003eM. leucurus\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e is estimated at 1.09\u0026ndash;0.21 Ma, whereas the split between \u003cem\u003eM. meles\u003c/em\u003e and \u003cem\u003eM. canescens\u003c/em\u003e is estimated to have occurred between 2.37 Ma and 450 ka (Marmi et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Law et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The dispersal of \u003cem\u003eM. anakuma\u003c/em\u003e to Japan likely occurred via the \u0026ldquo;Korean land bridge\u0026rdquo; during the Middle Pleistocene, as supported by fossil occurrences (Shikama \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1949\u003c/span\u003e, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1962\u003c/span\u003e; Kawamura et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Hasegawa \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Following this colonization, \u003cem\u003eM. anakuma\u003c/em\u003e has remained genetically distinct from its continental relatives. The presence of badgers on Crete, represented by \u003cem\u003eM. canescens\u003c/em\u003e, likely reflects a more recent, human-mediated introduction, with the oldest specimens dated only to approximately 3000 years ago (Masseti \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Meanwhile, \u003cem\u003eM. meles\u003c/em\u003e populations in the British Isles became geographically isolated from continental Europe over 450 ka (Gupta et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and small island populations, such as those on the Irish Peninsula, have arisen more recently (Sleemen et al. 2009).\u003c/p\u003e\u003cp\u003eInstitutional abbreviations: MCNG, Museu de Ciencies Natural de Granollers; NHMC, Natural History Museum of Crete; NMNS, National Museum of Natural Science in Tokyo; SNU, Seoul National University.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eThe gross muscular topography of both \u003cem\u003eMeles\u003c/em\u003e species was similar and shared the same general origin and insertion locations. Therefore, the muscular maps contain the complied results from all dissections undertaken for this study, with differences in muscle shape and structure between \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e described in detail in the following muscle descriptions.\u003c/p\u003e\n\u003cp\u003eShoulder muscles\u003c/p\u003e\n\u003cp\u003eM. supraspinatus\u003c/p\u003e\n\u003cp\u003eThis muscle is oblong and located in the lateral aspect of the scapula covering the entire supraspinous fossa surface (Fig.\u0026nbsp;2a). It extends dorsally along the scapular spine and inserts via thick tendons and some fleshy fibers to the proximal region of the humerus greater tubercle (Fig.\u0026nbsp;3a and c, 6e and 7a). Three bellies were observed: an intermediate one, running along the dorsal edge of the scapular spine, a principle one, on the lateral fossa surface, and a cranial one, which runs along the cranial ridge of the scapula and projects slightly towards the medial aspect.\u003c/p\u003e\n\u003cp\u003eM. infraspinatus\u003c/p\u003e\n\u003cp\u003eThis rectangular muscle is closely fused to the infraspinous fossa by fleshy fibers located on the lateral aspect of the scapula (Fig.\u0026nbsp;2a). It follows the caudal edge of the scapular spine, covers the internal surface of the metacromion and inserts via a thick tendinous band at the lateral aspect of the humeral greater tubercle, directly distal to m. supraspinatus (Fig.\u0026nbsp;3a, 6e and 7b). The caudal-most aspect of m. infraspinatus is distinguished from the rest by a notably thick tenacious band.\u003c/p\u003e\n\u003cp\u003eM. teres major\u003c/p\u003e\n\u003cp\u003eThis muscle originates along the caudal axillary border of the scapula via fleshy fibers, partially superficially to m. infraspinatus, and extending across approximately 1/3 of the scapula length (Fig.\u0026nbsp;2b). It is a relatively flat, elongated and rectangular muscle that is caudally fused to m. latissimus dorsi and inserts at the medial aspect of the deltoid ridge on the humeral shaft via a short thick band of tendons. The insertion area is long and thin, roughly half the length of its origin point, and runs parallel and superficially to the m. triceps brachii caput mediale (Fig.\u0026nbsp;3b, 6e and 7c).\u003c/p\u003e\n\u003cp\u003eM. teres minor\u003c/p\u003e\n\u003cp\u003eA small fleshy muscle situated on the proximal lateral aspect of the scapular caudal edge below the glenoid fossa, directly inferior to m. infraspinatus (Fig.\u0026nbsp;3a and 6e). It runs parallel to that same muscle before inserting distally and slightly more caudally to m. infraspinatus below the humeral greater tubercle via a thickly banded tendon (Fig.\u0026nbsp;7b).\u003c/p\u003e\n\u003cp\u003eM. deltoideus\u003c/p\u003e\n\u003cp\u003eThis muscle is located on the lateral aspects of the shoulder and is composed of two independent bellies: m. spinodeltoideus and m. acromiodeltoideus.\u003c/p\u003e\n\u003cp\u003eM. spinodeltoideus originates along the cranial aspect of the scapular spine, caudal to m. supraspinatus and superficial to m. infraspinatus (Fig.\u0026nbsp;2). The origin of this belly varied between both species and individual specimens. Its fleshy fibers originated from approximately halfway and to the most distal aspect of the scapular spine. It is thin and elongated with a rounded caudal edge which fuses to the lateral distal aspect of m. infraspinatus via a wide thin band of fine tendons (Fig.\u0026nbsp;2a, 3a and c, 7a) This muscle continues proximally beneath m. acromiodeltoideus and, in several specimens, the two bellies fused together via fleshy fibers before inserting with fine tendons along the deltoid crest of humerus. In others, the two bellies remained independent with m. spinodeltoideus inserting caudally to m. acromiodeltoideus.\u003c/p\u003e\n\u003cp\u003eM. acromiodeltoideus originates at the acromion on the proximal aspect of the scapular glenoid fossa (Fig.\u0026nbsp;2a, 3a and c, 7a). It is rhombus-like and connects with fine fascia along the proximal aspect of the humeral greater tubercle, distal to m. supraspinatus, and then inserts along the humeral deltoid ridge from the lateral to the cranial aspect.\u003c/p\u003e\n\u003cp\u003eM. subscapuaris\u003c/p\u003e\n\u003cp\u003eThis rectangular muscle originates across the entire medial surface of the scapula on the subscapular fossa and inserts via short thick tendons along the caudal aspect of the humeral lesser tubercle (Fig.\u0026nbsp;2b). There are six fibrous groups originating from the proximal aspect of the scapula, which then fan out towards the distal scapular borders. Each fibrous group is weakly separated by fine tendonous fibers which are further imprinted on the surface and border of the subscapular fossa as bony ridges. The most caudally situated group is the largest and most elongated, stretching across the entire medial axillary edge of the scapula. The most cranially situated group inserts slightly superficially to the neighbouring group (Fig.\u0026nbsp;3b-c, and 7a).\u003c/p\u003e\n\u003cp\u003eBrachium muscles\u003c/p\u003e\n\u003cp\u003eM. coracobrachialis\u003c/p\u003e\n\u003cp\u003eThis muscle originates at the cranial aspect of the coracoid process on the scapula and extends down the medial aspect of the humeral diaphysis as a long, thin fleshy tendon before inserting on the medio-caudal region of the humeral shaft, dorsal to the medial epicondyle (Fig.\u0026nbsp;3a-c). For \u003cem\u003eM. anakuma\u003c/em\u003e, m. coracobrachialis originates as a very long, thin tendon before broadening out approximately halfway down the humeral shaft and inserting as a long, triangular-like strip of fleshy fibers. \u003cem\u003eM. leucurus\u003c/em\u003e, on the other hand, has a considerably shorter tendon at the origin, which expands outwards into an elongated strip of flat muscle at the medio-caudal aspect of the humeral lesser tubercle. It thins out once more just below m. teres major and inserts as a long thin tendon above the medial epicondyle (Figs.\u0026nbsp;4 and 7c-d).\u003c/p\u003e\n\u003cp\u003eM. brachialis\u003c/p\u003e\n\u003cp\u003eThis muscle has an extensive origin, with thick fleshy fibers closely fused to the proximal aspect of the humerus diaphysis directly distal to m. triceps brachii laterale and up under the caudal-most aspect of the humeral head (Fig.\u0026nbsp;2a). This long, flat muscle continues down along the caudal humeral shaft and connects to m. brachioradialis via thick fleshy fibers (Fig.\u0026nbsp;5a). This then tampers out into a band of thin tendons and inserts at the ulnar tuberosity in the medial aspect (Fig.\u0026nbsp;3e and 7).\u003c/p\u003e\n\u003cp\u003eM. brachioradialis\u003c/p\u003e\n\u003cp\u003eThe origin length of m. brachioradialis varied between both \u003cem\u003eMeles\u003c/em\u003e species and specimens. This muscle originated from the proximal aspect of humeral lateral epicondylar crest, but in some specimens (e.g. \u003cem\u003eM. anakuma\u003c/em\u003e: POM-259), it extends approximately halfway up the humeral diaphysis as a long fleshy band (Fig.\u0026nbsp;2, 5a and 6e). It is relatively flat at its origin and then extends along the forearm and fuses at the distal medial aspect of the radius (Fig.\u0026nbsp;7a).\u003c/p\u003e\n\u003cp\u003eM. biceps brachii\u003c/p\u003e\n\u003cp\u003eThis muscle originates via a tight, compact band of tendons above the supraglenoid tubercle of the scapula and then rapidly expands out into a robust belly that extends down the medial aspect of the humerus (Fig.\u0026nbsp;2b, 5a and 6e). M. biceps brachii is held in place between the two humeral tubercle heads by its own retinaculum. It narrows again into a flattened tendon shortly before inserting on the bicipital tuberosity of the radius (Figs.\u0026nbsp;3 and 6e).\u003c/p\u003e\n\u003cp\u003eM. anconeus\u003c/p\u003e\n\u003cp\u003eThis flat, triangular, entirely fleshy muscle fuses directly via fibrous tissue to the distal caudal surface of the lateral humeral epicondyle (Fig.\u0026nbsp;6e). It extends distally and inserts along the lateral aspect of the ulnar head and along the olecranon process. Fine, fleshy fibers also connect it to m. extensor carpi ulnaris (Fig.\u0026nbsp;7).\u003c/p\u003e\n\u003cp\u003eM. triceps brachii\u003c/p\u003e\n\u003cp\u003eThis is the largest muscle in the brachium group and is situated in the lateral and medial aspects of the forelimb with five bellies: m. triceps brachii caput laterale, m. triceps brachii caput mediale, m. triceps brachii caput accessorium, m. triceps brachii caput longum, and m. triceps brachii caput angulare (Fig.\u0026nbsp;2). All bellies insert on the proximal head of the ulnar olecranon process.\u003c/p\u003e\n\u003cp\u003eM. triceps brachii caput laterale has a relatively long and narrow origin via thin tendons to the proximal lateral aspect of the deltoid ridge, immediately distal to m. teres minor (Fig.\u0026nbsp;2a). However, there are additional fleshy attachments along the cranial lateral aspect of the humeral head and some fine fibrous tissues connecting it superficially to the distal laterally situated extensor muscles, as well along the supracondyloid ridge border. This belly inserts on the proximal lateral facet of the olecranon tuber, directly above m. anconeus (Fig.\u0026nbsp;7).\u003c/p\u003e\n\u003cp\u003eM. triceps brachii caput mediale is located in the medial aspect of the forelimb and partially fuses to m. triceps brachii caput lateral via fleshy fibers which extend towards the olecranon tuber insertion (Fig.\u0026nbsp;2b). More fleshy fibers fuse to the caudal aspect of the humeral head and the caudal proximal facet of m. subscapularis, while fine expanses of tendons also extend down along the medial aspect of the deltoid ridge, running parallel to the insertion of m. teres major (Fig.\u0026nbsp;7).\u003c/p\u003e\n\u003cp\u003eM. triceps brachii caput accessorium is a short cylindrical muscle located in the medial aspect of the forearm (Fig.\u0026nbsp;2b). It is the smallest of the m. triceps brachii muscle group and is situated partially superficial to the insertion of m. coracobrachialis. It originates from the caudal-medial aspect of the humeral medial epicondyle and inserts on the proximal medial facet of the ulnar olecranon, directly distal to both m. triceps brachii caput longum and m. triceps brachii caput mediale (Fig.\u0026nbsp;4b, 6 and 7).\u003c/p\u003e\n\u003cp\u003eM. triceps brachii caput longum is the largest and thickest belly of the m. triceps brachii group, located in both the medial and lateral aspects of the forelimb (Fig.\u0026nbsp;2). This muscle originates via abundant fleshy fibers at the mid-axillary border of the scapula, distal to m. triceps brachii caput laterale in the lateral view. It inserts on the most proximal caudal aspect of the ulnar olecranon (Fig.\u0026nbsp;3a, 4 and 6).\u003c/p\u003e\n\u003cp\u003eM. triceps brachii caput angulare is a very flat, elongated muscle that is the most caudally located of the m. triceps brachii heads and is situated superficially to m. triceps brachii caput longum (Figs.\u0026nbsp;2 and 3a). It originates via fleshy fibers on the axillary-most border of the scapula in the medial view, directly superficial to m. teres major. It attaches via fine fibrous tendons on the caudal facet of the ulnar olecranon (Fig.\u0026nbsp;7b).\u003c/p\u003e\n\u003cp\u003eM. tensor fasciae antebrachii\u003c/p\u003e\n\u003cp\u003eThis is a small, very flat muscle on the medial aspect of the forelimb and sits directly superficial to m. triceps brachii caput mediale and m. triceps brachii caput longum (Fig.\u0026nbsp;2b). It fuses directly to the caudal border of m. latissi dorsi via thin fascia for \u003cem\u003eM. anakuma\u003c/em\u003e and fleshy fibers for \u003cem\u003eM. leucurus\u003c/em\u003e (Fig.\u0026nbsp;5b) before inserting on the medial facet of the olecranon tuber immediately distal to m. triceps brachii caput longum.\u003c/p\u003e\n\u003cp\u003eDistal arm muscles\u003c/p\u003e\n\u003cp\u003eM. extensor\u003c/p\u003e\n\u003cp\u003eThe extensor group has six independent muscles: m. extensor carpi radialis longus, m. extensor carpi radialis brevis, m. extensor digitorum communis, m. extensor digitorum lateralis, m. extensor carpi ulnaris, and m. extensor pollicis, which are stacked vertically to each other along the humeral lateral epicondylar crest and lateral epicondyle (Fig.\u0026nbsp;2a).\u003c/p\u003e\n\u003cp\u003eM. extensor carpi radialis longus and m. extensor carpi radialis brevis originates via fleshy fibers from the proximal aspect of the humeral lateral epicondyle (Fig.\u0026nbsp;2a). They are fused at their origin and split approximately one-third of the way towards the ulnar styloid, with m. extensor carpi radialis longus situated superficially to m. extensor carpi radialis brevis, and then become two tightly wound bands of flattened tendons. These muscles are held in place by their own retinaculum located at the distal anterior surface of the radius, with m. extensor carpi radialis longus inserting at the base of metacarpal II, and m. extensor carpi radialis brevis inserts at the base of metacarpal III (Fig.\u0026nbsp;7).\u003c/p\u003e\n\u003cp\u003eM. extensor digitorum communis originates via fleshy fibers on the humeral lateral epicondylar crest, directly distal to m. extensor carpi radialis longus and m. extensor carpi radialis brevis (Fig.\u0026nbsp;2a). It extends distally towards the wrist before splitting into four groups of fine, elongated tendons that insert on the distal phalanges of digits II-V (Fig.\u0026nbsp;5a and 7).\u003c/p\u003e\n\u003cp\u003eM. extensor digitorum lateralis originates via fleshy fibers on the distal aspect of humeral lateral epicondylar crest immediately distal to m. extensor digitorum communis (Fig.\u0026nbsp;2a). It is fused as a single entitity at its origin point however, it splits into two bellies \u0026ndash; medial and lateral \u0026ndash; approximately midway down the ulnar shaft. The medial belly further divides into two long flattened tendons that inserts in the distal phalanges of digits III and IV, while the lateral belly narrows into a single long thin tendon that inserts in the distal phalange of digit V (Fig.\u0026nbsp;5a and 7).\u003c/p\u003e\n\u003cp\u003eM. extensor carpi ulnaris originates via fleshy fibers from the proximal aspect of the humeral lateral epicondyle directly distal to m. extensor digitorum lateralis (Fig.\u0026nbsp;2a). This muscle rapidly expands into a thick belly before narrowing again into a condensed cluster of tendons which insert beneath m. extensor digit I \u0026amp; II at the base of metacarpal V (Fig.\u0026nbsp;5a and 7).\u003c/p\u003e\n\u003cp\u003eM. extensor digiti I \u0026amp; II is thick, fleshy and, unlike the other extensor muscles, is originates via fleshy fibers along the lateral ulnar shaft (Fig.\u0026nbsp;6e). It extends diagonally across the forearm towards the medial side, passing beneath all the extensor muscles excluding m. extensor carpi ulnaris, and then inserting as very fine tendons on the distal phalanges of digits I and II (Fig.\u0026nbsp;5a and 7).\u003c/p\u003e\n\u003cp\u003eM. flexor\u003c/p\u003e\n\u003cp\u003eThe flexor group is located in the medial aspect of the humeral epicondyle and consists of four independent muscles: m. flexor digitorum profundus, m. flexor digitorum superficialis, m. flexor carpi radialis, and m. flexor carpi ulnaris (Fig.\u0026nbsp;2b).\u003c/p\u003e\n\u003cp\u003eM. flexor digitorum profundus is the largest of the flexor muscles and is located in the deep caudal aspect of the distal forearm and consists of five heads: three with humeral origins \u0026ndash; caput humerale laterale, caput humerale mediale, and caput humerale profundus, one originating on the ulna \u0026ndash; caput ulnare, and one with origins on both the ulna and radius \u0026ndash; caput radiale (Fig.\u0026nbsp;2b, 3 and 7). The humeral heads originate on the distal caudal-medial aspect of the humeral epicondyle however only caput laterale and caput profundus originate on the bone, as caput mediale fuses directly to caput laterale via fleshy fibers that extend midway along the ulna shaft towards the wrist. Caput profundus is considerably smaller, thinner and furtherly independent from the other two humeral heads, connected lengthwise only via thin fascia. Caput ulnare has a large heavy belly, originating via fleshy fibers from the proximal caudal aspect of the ulna shaft and extending two-thirds towards the wrist, while caput radiale is relatively flat and stretches between the lateral and cranial aspects of the radius and ulna shafts as fleshy fibers. M. flexor digitorum profundus heads fuse together (caput ulnare notably superficial to the rest) and are held in place by a thick retinaculum at the wrist before dividing into thin elongated tendons that insert into the distal phalanges I-V (Fig.\u0026nbsp;7).\u003c/p\u003e\n\u003cp\u003eM. flexor digitorum superficialis is a fleshy relatively thin muscle situated directly superficial to m. flexor digitorum profundus (Fig.\u0026nbsp;6b). It does not appear to insert directly onto the caudal humeral medial epicondyle and instead its origin is fused to the distal surface of m. flexor digitorum profundus caput humerale laterale and mediale bellies. The insertion is also directly dorsal to m. flexor digitorum profundus, this time by thin fatty fascia superficial to m. flexor digitorum profundus caput ulnare, immediately between the diversion of the tendons towards digits I and V (Fig.\u0026nbsp;6b).\u003c/p\u003e\n\u003cp\u003eM. flexor carpi radialis originates from the medial epicondyle of the humerus directly distal to m. pronator teres (Fig.\u0026nbsp;2b). It narrows into two thin long tendons that insert into the base of metacarpal II and III (Fig.\u0026nbsp;6b-d). It is also partially fused via thick fleshy fibers to m. pronator teres midway along the distal arm (Figs.\u0026nbsp;5 and 7).\u003c/p\u003e\n\u003cp\u003eM. flexor carpi ulnaris is located on the medial side of the distal forearm, originating via fleshy fibers from the medial aspect of the ulnar olecranon tuber and medial humeral epicondyle as two independent bellies: caput ulnare and caput humerale, respectively (Fig.\u0026nbsp;2b). Caput ulnare is large and thick, while caput humerale is comparatively thin. These two heads fuse together partway along the ulnar shaft before inserting via a thin long tendon on the sesamoid most proximal to metacarpal V (Fig.\u0026nbsp;6a and 7).\u003c/p\u003e\n\u003cp\u003eM. palmaris longus\u003c/p\u003e\n\u003cp\u003eThis muscle originates from the distal medial epicondyle of the humerus caudal to m. flexor carpi radialis (Fig.\u0026nbsp;2b and 6a-c). It rapidly expands out into a robust belly and then quickly thins out again into four flattened tendons which pass over the retinaculum that secures m. flexor digitorum profundus at the wrist and inserts into the distal phalanges of digits II-V (Fig.\u0026nbsp;7c-d).\u003c/p\u003e\n\u003cp\u003eM. abductor digiti I\u003c/p\u003e\n\u003cp\u003eIt is a deep muscle that originates via fleshy fibers along the whole lateral surface of the ulna and radial shafts (Fig.\u0026nbsp;6e). It then narrows into a thin tendon that inserts into the base of metacarpal I (Fig.\u0026nbsp;7f-h).\u003c/p\u003e\n\u003cp\u003eDeep muscles\u003c/p\u003e\n\u003cp\u003eM. pronator teres\u003c/p\u003e\n\u003cp\u003eIt is a roughly semicircular muscle originating via fleshy fibers on the most proximal aspect of the humeral medial epicondyle, immediately cranial to the insertion of m. triceps brachii caput accessorium (Fig.\u0026nbsp;2b and 3d). It partially splits into two indistinct fleshy bellies which come together quickly to insert as a long insertion strip located cranially mid-way down the radial shaft (Fig.\u0026nbsp;5a, 6 and 7a).\u003c/p\u003e\n\u003cp\u003eM. supinator\u003c/p\u003e\n\u003cp\u003eThis is a relatively flat, triangular muscle originating via thick fleshy fibers on the distal cranial aspect of the lateral humeral epicondyle, directly adjacent to m. extensor carpi ulnaris (Fig.\u0026nbsp;3d). It runs roughly parallel to m. pronator teres and then inserts on the dorsal-medial aspect of the radial shaft, approximately two-thirds of the way towards the distal styloid process (Fig.\u0026nbsp;6e).\u003c/p\u003e\n\u003cp\u003eM. pronator quadratus\u003c/p\u003e\n\u003cp\u003eThis muscle is fused via thick fleshy fibers to the dorsal-medial distal surface of the ulna interosseous crest (Fig. 3d-e). It is rectangular, flat, and inserts on the caudal distal surface of the radius via fleshy fibers (Fig. 6d and 7b).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we examined the forelimb myology of \u003cem\u003eM. leucurus\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e and compared their muscular structure to published data of other closely related species. While the general topography of the forelimb muscular structure was similar, we found several key differences in the shape and form of the muscles which we will discuss in detail. Then, we analysed the forelimb osteology of \u003cem\u003eMeles\u003c/em\u003e as continental-insular pairs (\u003cem\u003eleucurus-anakuma\u003c/em\u003e and \u003cem\u003emeles-canescens\u003c/em\u003e, where here \u003cem\u003eM. canescens\u003c/em\u003e refers to the insular communities inhabiting Crete) to comparatively investigate the functional adaptations of island \u003cem\u003eMeles\u003c/em\u003e populations.\u003c/p\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eComparative muscular anatomy of\u003c/span\u003e \u003cspan type=\"ItalicUnderline\" class=\"ItalicUnderline\" name=\"Emphasis\"\u003eM. anakuma\u003c/span\u003e \u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eand\u003c/span\u003e \u003cspan type=\"ItalicUnderline\" class=\"ItalicUnderline\" name=\"Emphasis\"\u003eM. leucurus\u003c/span\u003e\u003c/p\u003e\u003cp\u003eFor the shoulder muscles, the three m. supraspinatus bellies described in Ercoli et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) for the lesser grison (\u003cem\u003eGalictis cuja\u003c/em\u003e) are also evident in both \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e observed here (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). However, both badgers have a comparatively more robust cranial belly and overall larger m. supraspinatus and m. infraspinatus surface areas than \u003cem\u003eG. cuja\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The higher number of m. supraspinatus bellies has been linked to an increased range of motion in the shoulder joint, particularly during arm-extension (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). For m. teres major in both \u003cem\u003eMeles\u003c/em\u003e species studied here, its origin is partially superficially fused to the distal caudal border of m. infraspinatus. This observation has also been reported for the kinkajou, \u003cem\u003ePotos flavus\u003c/em\u003e (V\u0026eacute;lez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and for the racoon, \u003cem\u003eProcyon lotor\u003c/em\u003e (Feeney \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Allen \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1882\u003c/span\u003e); however, it has not been described for any mustelid species although it appears to be present in several studies (e.g. \u003cem\u003eG. cuja\u003c/em\u003e, Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; \u003cem\u003eMartes martes\u003c/em\u003e and \u003cem\u003eMartes foina\u003c/em\u003e, B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The m. teres major origin itself is approximately one-fourth of the scapular length, which is considerably longer than in \u003cem\u003eG. cuja\u003c/em\u003e (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). M. teres major is a powerful shoulder joint flexor and limb retractor used during the power stroke and so is particularly enlarged in specialised diggers (Moore \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn \u003cem\u003eMeles\u003c/em\u003e, m. coracobrachialis is a small, thin muscle, yet its shape and nomenclature vary considerably among caniforms (V\u0026eacute;lez-Garcia et al. 2023: table 3). Windle and Parsons (\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e1897\u003c/span\u003e) described it as m. coracobrachialis medius in \u003cem\u003eMeles\u003c/em\u003e, noting its more distal insertion on the humeral shaft compared to other species. In \u003cem\u003eG. cuja\u003c/em\u003e, it has been termed m. coracobrachialis brevis, with a short insertion directly distal to the humeral lesser tubercle (Ercoli et al \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). \u003cem\u003eP. flavus\u003c/em\u003e possesses both brevis and longus heads, inserting on the proximal medial aspect of the humeral shaft and directly above the supracondylar foramen (V\u0026eacute;lez-Garcia et al. 2023). The red panda, \u003cem\u003eAilurus fulgens\u003c/em\u003e (Fisher et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e)d \u003cem\u003emartes\u003c/em\u003e (Yousefi et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) show only the more distal insertion. B\u0026ouml;hmer et al. (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) recorded a notably more distal insertion in \u003cem\u003eM. foina\u003c/em\u003e and \u003cem\u003eM. martes\u003c/em\u003e on the medial aspect of the olecranon. In this study, while origin and insertion were consistent, the position of the muscle belly varied between species. For \u003cem\u003eM. leucurus\u003c/em\u003e, the belly lay directly beneath the humeral lesser tubercle, whereas in \u003cem\u003eM. anakuma\u003c/em\u003e, it was situated distally along the humeral shaft (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003e). \u003cem\u003eM. anakuma\u003c/em\u003e also exhibited a second smaller belly that inserted directly distal to the humeral lesser tubercle (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003eb), resembling the dual insertion seen in \u003cem\u003eP. flavus\u003c/em\u003e (V\u0026eacute;lez-Garcia et al. 2023), although it was significantly reduced (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003e). This distal insertion is most similar to \u003cem\u003eA. fulgens\u003c/em\u003e in size and shape, while the dual arrangement mirrors that of arboreal taxa like \u003cem\u003eP. flavus\u003c/em\u003e. Functionally, m. coracobrachialis is a weak shoulder stabilizer and humeral adductor, and its reduced appearance suggests a minimal impact on locomotion (Taylor \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1974\u003c/span\u003e). However, its presence has been associated with arboreal activity (Salesa et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), and more distal insertions, as in \u003cem\u003eP. flavus\u003c/em\u003e and \u003cem\u003eA. fulgens\u003c/em\u003e, may enhance shoulder adduction strength and increase the animals\u0026rsquo; climbing ability (Monroy-Cendales et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Monroy-Cendales et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Ercoli et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) considered the muscles\u0026rsquo; presence a plesiomorphic trait within Carnivora. Our findings suggest interspecific variation in \u003cem\u003eMeles\u003c/em\u003e, with \u003cem\u003eM. anakuma\u003c/em\u003e retaining a dual insertion pattern similar to more arboreally-adapted taxa, potentially reflecting an ancestral or ecologically driven trait (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e9\u003c/span\u003e) (Marmi et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Law et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIn general, there are four m. triceps brachii heads: laterale, longum, mediale, and accessorium, which all insert on the proximal aspect of the ulnar olecranon however, there are frequent variations among caniforms. In all mustelids and mephitids (Windle and Parsons \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e1897\u003c/span\u003e; Moore et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), including the \u003cem\u003eMeles\u003c/em\u003e species examined in this study, there is an additional head, m. triceps brachii caput angulare, which originates at the most distal caudal border of the scapula and inserts on the caudal aspect of the ulnar head (e.g. Figure\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). For \u003cem\u003eP. flavus\u003c/em\u003e, m. triceps brachii caput angulare originates considerably more proximally along the scapular axillary border, likely making space for a larger m. teres major origin (B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Functioning in conjunction with m. triceps brachii to support elbow extension is the m. tensor fasciae antebrachii muscle, the presence of which has not been previously described for \u003cem\u003eMeles\u003c/em\u003e. In both \u003cem\u003eP. flavus\u003c/em\u003e (V\u0026eacute;lez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2023\u003c/span\u003e)d \u003cem\u003elotor\u003c/em\u003e (Feeney \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), m. tensor fasciae antebrachii had two heads (caudal and cranial), whereas for \u003cem\u003eMeles\u003c/em\u003e, only the cranial belly was present and fused directly to the caudal border of m. latissi dorsi via thin fascia for \u003cem\u003eM. anakuma\u003c/em\u003e, and fleshy fibers for \u003cem\u003eM. leucurus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb, \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ea and \u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003eb). For \u003cem\u003eG. cuja\u003c/em\u003e (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) however, it was considered the modified intermediate belly of m. triceps brachii mediale. Our findings highlight the phylogenetic evolution of m. tensor fasciae antebrachii among caniforms (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e9\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn contrast to the four heads of m. flexor digitorum profundus described for \u003cem\u003eM. meles\u003c/em\u003e (B\u0026ouml;hmer et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and typically present in canids and felids (Fishbeck and Sebastiani, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Evans et al. 2012), five heads were consistently observed in our specimens, with an additional humeral head, m. flexor digitorum profundus humerale profundus (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e5\u003c/span\u003ec). In both \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e, m. flexor digitorum superficialis originates significantly more distally and exhibits a well-developed fleshy belly, differing from the shorter muscle belly observed in \u003cem\u003eG. cuja\u003c/em\u003e (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e)d \u003cem\u003efulgens\u003c/em\u003e (Fisher et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In this study, the muscle does not attach directly to the bone but instead fuses entirely with m. flexor digitorum profundus at the wrist bundle (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e6\u003c/span\u003e), contrasting with the distinct distal insertion described for \u003cem\u003eG. cuja\u003c/em\u003e and \u003cem\u003eA. fulgens\u003c/em\u003e. Functionally, m. flexor digitorum superficialis supports wrist flexion however, given its reduced appearance in \u003cem\u003eMeles\u003c/em\u003e and often misidentification or omission in anatomical descriptions, its comparative analysis across taxa remains challenging (Ercoli et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Perdomo-C\u0026aacute;rdenas et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eContinental-insular pairwise comparisons in\u003c/span\u003e \u003cspan type=\"ItalicUnderline\" class=\"ItalicUnderline\" name=\"Emphasis\"\u003eMeles\u003c/span\u003e\u003c/p\u003e\u003cp\u003eComparative studies have successfully explored the close relationship between myology and osteology in describing locomotive habits (e.g. Van Valkenburgh \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Argot \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Fabre et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; B\u0026ouml;hmer et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In general, \u003cem\u003eMeles\u003c/em\u003e badgers exhibit skeletal features consistent with a powerful \u0026lsquo;scratch-digger\u0026rsquo; burrowing style. These include large muscle attachment sites at the shoulder and elbow joints, resulting in stout humeri and ulnae with thick shafts and robust proximal and distal features (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003e) (Hildebrand and Goslow \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Moore et al. 2011; Samuels et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The shoulder muscles \u0026ndash; especially the rotator cuff muscle m. subscapularis and the abductor m. deltoideus \u0026ndash; contribute to shoulder stabilization and humeral rotation (Fishbeck and Sebastiani \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Janis and Mart\u0026iacute;n-Serra \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), consistent with the enlarged humeral greater tubercles observed in all \u003cem\u003eMeles\u003c/em\u003e specimens in this study (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003e). Additionally, digging performance is enhanced by well-developed humeral medial and lateral epicondyles and an elongated ulnar olecranon, which are attachment sites for powerful distal flexor and extensor muscles (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003e) (Argot \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Hildebrand and Goslow \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Moore et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Samuels et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Rose et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIn the \u003cem\u003eleucurus-anakuma\u003c/em\u003e pair, the insular \u003cem\u003eM. anakuma\u003c/em\u003e exhibits the cranially inclined olecranon typically seen in semifossorial and arboreal species (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003eh) (Fabre et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). In contrast, \u003cem\u003eM. leucurus\u003c/em\u003e retains a straighter ulnar profile in the medial view with a caudally inclined olecranon process more common in terrestrial quadrupeds optimized for strong elbow extension (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003eg) (Henderson et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Functionally, these changes alter the leverage and direction of muscle force across the elbow joint. In \u003cem\u003eM. anakuma\u003c/em\u003e, the caudal head of the olecranon (insertion site for m. triceps brachii caput longum) is reduced, while the cranial head (for caput mediale) is enlarged (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003eh). This shift suggests decreased involvement of shoulder-extending caput longum, and increased reliance on caput mediale, which primarily supports late-phase elbow extension \u0026ndash; often seen in species engaging in climbing or intricate foraging behaviours (Kholinne et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Conversely, \u003cem\u003eM. leucurus\u003c/em\u003e demonstrates stronger development of the medial epicondyle, particularly in attachment regions for m. pronator teres and other forearm flexors (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003ec), suggesting enhanced torque production during pronation-supination. This muscular development is likely associated with sweeping arm movements for overturning compact soil and possibly an adaptation to life in colder regions with harder substrates (Neal and Cheeseman, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) where badger setts are often dug deeper and exhibit more complex structures to withstand the lower temperatures (Choi et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Additionally, the twisted orientation of the humeral lateral epicondyle relative to the medially tilted head further supports the presence of strain-induced remodeling from forceful digging behaviours (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003ec). \u003cem\u003eM. anakuma\u003c/em\u003e also exhibits a slight reduction in the distal projection of the humeral trochlea, which may indicate an increase in manouverability in the elbow joint (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003ed). Therefore, the insular \u003cem\u003eM. anakuma\u003c/em\u003e exhibits skeletal shifts that suggest increased arm flexibility, potentially indicating less intensive digging or more diverse substrate interaction, reflecting differences in island environments.\u003c/p\u003e\u003cp\u003eIn the \u003cem\u003emeles-canescens\u003c/em\u003e pair, the insular population of \u003cem\u003eM. canescens\u003c/em\u003e exhibited a decreased length of the humerus and ulna, and an increased robusticity of the deltoid ridge and epicondylar crest (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003ea\u0026ndash;b, e-f). These changes suggest a functional shift toward more compact and powerful limb extension, potentially compensating for the reduced lever length via greater muscle mass at origin and insertion sites. The reduction in the humeral lesser tubercle \u0026ndash; the insertion point for m. subscapularis, and the reduced distal projection of the humeral trochlea in the \u003cem\u003eM. canescens\u003c/em\u003e specimens, may indicate increased mobility at the shoulder and elbow joints (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003eb) (Argot \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Salesa et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This, combined with the enlarged cranial aspect of the olecranon and reduced projection in the humeral trochlea, implies enhanced arm-extension capabilities in the insular form, possibly reflecting greater reliance on limb flexibility (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003ea-b, e-f). These morphological patterns suggest that the Crete population of \u003cem\u003eM. canescens\u003c/em\u003e has evolved toward a more versatile digging style, potentially as a response to differing substrate properties or ecological demands on the island.\u003c/p\u003e\u003cp\u003eFunctional responses shared in insular \u003cem\u003eMeles\u003c/em\u003e\u003c/p\u003e\u003cp\u003ePrevious studies have reported reductions in the body size of insular \u003cem\u003eMeles\u003c/em\u003e populations \u0026ndash; including \u003cem\u003eM. canescens\u003c/em\u003e inhabitants on Crete and \u003cem\u003eM. anakuma\u003c/em\u003e in Japan \u0026ndash; compared to their mainland counterparts, \u003cem\u003eM. meles\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e respectively (Baryshnikov et al. 2002; Abramov and Puzachenko \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Proulx et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Savvidou et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Nonetheless, our study found no significant size difference (based on humerus length) between \u003cem\u003eM. leucurus\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), although a reduction was evident between \u003cem\u003eM. meles\u003c/em\u003e and the \u003cem\u003eM. canescens\u003c/em\u003e populations of Crete (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e9\u003c/span\u003e). This variation in body response highlights the complexity of insular adaptations in medium-carnivores and suggests that body size alone may not fully capture the extent of functional adaptation in island settings. Notably, we recognized significant morphological patterns in the forelimbs of the insular forms. Both island popuations of \u003cem\u003eM. canescens\u003c/em\u003e and \u003cem\u003eM. anakuma\u003c/em\u003e exhibited shared reductions in the distal and caudal projections of the humeral trochlea and enlargements of the cranial aspect of the olecranon tuber (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e8\u003c/span\u003eb, e, and \u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e9\u003c/span\u003e), suggesting that increased elbow flexibility \u0026ndash; possibly favoring greater forelimb mobility for handling variable or less resistant substrates, is a general trend among insular \u003cem\u003eMeles\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003eThese osteological changes complement ecological observations for both insular and mainland badger populations. Field-based ecology has been studied extensively for \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. meles\u003c/em\u003e, including annual hibernation periods (e.g. Kowalczyk et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Tanaka \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), social structure (e.g. Neel and Cheeseman 1996; Kaneko et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Tanaka et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and diet and distribution (e.g. Kaneko et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Proulx et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), whereas ecological and behavioral data remain sparse for \u003cem\u003eM. leucurus\u003c/em\u003e and \u003cem\u003eM. canescens\u003c/em\u003e, making direct comparisons of the examined continental-insular pairs challenging. Despite these gaps, a few studies have observed dietary shifts and increased diurnal activity in \u003cem\u003eM. meles\u003c/em\u003e populations on Rutland Island compared to their continental counterpart, interpretating these changes as insular traits (Sleeman et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sleeman and Davenport \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Furthermore, ecological studies on \u003cem\u003eM. leucurus\u003c/em\u003e populations on Jeju Island recorded badgers as active from 3pm (Oh \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), which is significantly earlier than what has been recorded for \u003cem\u003eM. anakuma\u003c/em\u003e, with activity beginning from 6pm (Tanaka 2005). Possible increased diurnal activity was also extrapolated for other insular badgers (\u003cem\u003eM. canescens\u003c/em\u003e from Crete and \u003cem\u003eM. anakuma\u003c/em\u003e) based on cranial features such as greater interorbital distance (Savvidou et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These patterns echo broader trends observed in other island-adapted carnivores \u0026ndash; such as the Channel Islands foxes and Arctic fox populations which have shown both ecological and morphological shifts that align with increased arboreal activities and altered foraging strategies (Wayne et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Moore and Collins \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Nanova and Pr\u0026ocirc;a \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Collectively, our findings indicate that enhanced limb mobility may represent broader insular trends extending beyond terrestrial generalists to include subfossorial forms. We also note that direct behavioral observations are inherently limted by observer access, seasonal variation, and plastic ecological strategies especially for nocturnal, burrowing mammals like badgers. In contrast, skeletal features accumulate the selective pressures exerted over evolutionary time scales and can thus serve as good proxies for habitual locomotor and foraging behaviour. By linking skeletal morphology to inferred changes in habitat use, foraging strategy and locomotor behavior, our study reinforces the importance of integrating ecological and anatomical approaches to better understand how medium-sized mesocarnivores like \u003cem\u003eMeles\u003c/em\u003e adapt to island ecosystems.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThis study provides the first comparative myological analysis of the forelimbs in \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e, revealing both conserved and novel anatomical traits. Notably, the m. coracobrachialis muscle, which has been described as a plesiomorphic trait in Mustelidae, has a more distal main belly and second shorter head for \u003cem\u003eM. anakuma\u003c/em\u003e compared to \u003cem\u003eM. leucurus\u003c/em\u003e, which may represent taxon-specific variations not previously documented in other mustelid species. The presence of m. tensor fasciae antebrachii is observed in \u003cem\u003eMeles\u003c/em\u003e for the first time and highlights the phylogenetic significance of this muscle among caniforms. Osteological comparisons with other continental-insular \u003cem\u003eMeles\u003c/em\u003e pairs show shared morphological adaptations\u0026mdash;such as increased forelimb flexibility\u0026mdash;in insular species. Contrary to expectations based on prior studies of insular dwarfism, \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e did not differ significantly in body size, suggesting that other undetermined factors had a more promeninet influence on forelimb development in \u003cem\u003eM. anakuma\u003c/em\u003e. These findings underscore the complex interplay between phylogeny, ecology, and biomechanics in shaping the musculoskeletal evolution following dispersal events. Future research should prioritize integrating myological, morphological and ecological data \u0026ndash; particularly with specimens from underrepresented regions such as Greece, South Korea, and mainland China \u0026ndash; to further refine our understanding of functional and evolutionary patterns in \u003cem\u003eMeles\u003c/em\u003e and other mustelids. Detailed investigations into the finer-scale factors which influence intraspecific variation, such as sexual dimorphism and ontogenetic growth, would also greatly enhance our understanding of the ecological and behavioral roles favored by different \u003cem\u003eMeles\u003c/em\u003e species.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAcknowledgements\u003c/p\u003e\n\u003cp\u003eWe would like to express our gratitude to Chang-Yong Choi (Seoul National University) and Hee-Jong Kim (Wildlife Rescue Center, Ulsan) for providing samples and allowing us to perform dissections in their laboratories. We are especially grateful to Shin-ichiro Kawada (National Museum of Nature and Science, Japan), Petros Lymberakis and Panagiotis Georgiakakis (Museum of Crete, Greece), Toni Arrizabalaga and Anna Surroca (Museu de Ciencies Natural de Granollers, Spain),\u0026nbsp;Roberto Portela Miguez and Phaedra Kokkini (National Museum of Natural Science in London),\u0026nbsp;Géraldine Veron and Lucile Armand (National Museum of Natural Science in Paris), Aleksandra Panyutina (Tel Aviv University), Yuusuke Goto (Ibaraki Nature Museum), Eri Akasaki and\u0026nbsp;Hiroshi Tanaka (Yamaguchi Prefectural Museum), Keiichi Takahashi (Lake Biwa Museum), Nozomi Nakanishi (Kita-Kyushu Museum of Natural and Human History), Fumio Takahashi (Mine City Museum of History and Folklore), Hashimoto Tatsuya and Mitsuharu Matsumoto (Kagoshima University), Yohoko Okumura (Kuzu Fossil Museum), Jun Nemoto (Tohoku University Museum), Taruno Hiroyuki (Osaka Museum of Natural History), and Satoshi Suzuki (Kanagawa Prefectural Museum of Natural History) for granting us access to their extant badger collections. We are also thankful to Haruto Sugeno, Shingo Nishimura and Haemin Seo for supplying us with the \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e roadkill specimens, and Mao Shimoda, Kyunghae Min and Tri Sayektiningsih for assisting with the dissections. Finally, we would further like to thank Kohei Tanaka (University of Tsukuba), Yoshikazu Hasegawa (Iida City Museum, Nagano), Yayoi Kaneko (Tokyo University of Agriculture and Technology), and Qigao Jiangzuo and Peiran Li (Institute of Vertebrate Paleontology and Paleoanthropology, China) for their constructive comments on the early draft.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAuthor contributions\u003c/p\u003e\n\u003cp\u003eAll authors have read and agreed to the definitive version of the manuscript.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFunding\u003c/p\u003e\n\u003cp\u003eThis work was partially funded by the Japanese Government Monbukagakusho Scholarship (grant number 210348).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eData availability statement\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are included in this published article.\u003c/p\u003e\n\u003cp\u003eDeclarations\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e The authors have no competing interests to declare that are relevant to the content of this article.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbramov AV, Puzachenko AY (2013) The taxonomic status of badgers (Mammalia, Mustelidae) from Southwest Asia based on cranial morphometrics, with the redescription of \u003cem\u003eMeles canescens\u003c/em\u003e. Zootaxa 3681(1):44\u0026ndash;58. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.11646/zootaxa.3681.1.2\u003c/span\u003e\u003cspan address=\"10.11646/zootaxa.3681.1.2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eAllen H (1882) The muscles of the limbs of the raccoon (\u003cem\u003eProcyon lotor\u003c/em\u003e). Proc Acad Nat Sci Philadelphia 115\u0026ndash;144.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eArgot C (2001) Functional-adaptive anatomy of the forelimb in the Didelphidae, and the paleobiology of the Paleocene marsupials \u003cem\u003eMayulestes ferox\u003c/em\u003e and \u003cem\u003ePucadelphys andinus\u003c/em\u003e. J Morphol 247(1):51\u0026ndash;79. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/1097-4687(200101)247:1\u0026lt;51::AID-JMOR1003\u0026gt;3.0.CO;2-%23\u003c/span\u003e\u003cspan address=\"10.1002/1097-4687(200101)247:1%3C51::AID-JMOR1003%3E3.0.CO;2-%23\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBaryshnikov GF, Puzachenko AY, Abramov AV (2003) New analysis of variability of cheek teeth in Eurasian badgers (Carnivora, Mustelidae, \u003cem\u003eMeles\u003c/em\u003e). Russian Journal of Theriology 1(2):133\u0026ndash;149. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.15298/rusjtheriol.01.2.07\u003c/span\u003e\u003cspan address=\"10.15298/rusjtheriol.01.2.07\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBen\u0026iacute;tez-L\u0026oacute;pez A, Santini L, Gallego-Zamorano J, Mil\u0026aacute; B, Walkden P, Huijbregts MA, Tobias JA (2021) The island rule explains consistent patterns of body size evolution in terrestrial vertebrates. Nat Ecol Evol 5(6):768\u0026ndash;786. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/s41559-021-01426-y\u003c/span\u003e\u003cspan address=\"10.1038/s41559-021-01426-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eB\u0026ouml;hmer C, Fabre AC, Herbin M, Peign\u0026eacute; S, Herrel A (2018) Anatomical basis of differences in locomotor behavior in martens: A comparison of the forelimb musculature between two sympatric species of \u003cem\u003eMartes\u003c/em\u003e. Anat Rec 301(3):449\u0026ndash;472. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ar.23742\u003c/span\u003e\u003cspan address=\"10.1002/ar.23742\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eB\u0026ouml;hmer C, Fabre AC, Taverne M, Herbin M, Peign\u0026eacute; S, Herrel A (2019) Functional relationship between myology and ecology in carnivores: do forelimb muscles reflect adaptations to prehension? Biol J Linn Soc 127(3):661\u0026ndash;680. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/biolinnean/blz036\u003c/span\u003e\u003cspan address=\"10.1093/biolinnean/blz036\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eB\u0026ouml;hmer C, Theil JC, Fabre AC, Herrel A (2020) Atlas of Terrestrial Mammal Limbs. CRC Press\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eChoi TY, Lee SK, Woo DG (2019) Study on spatial ecology of Asian Badger (\u003cem\u003eMeles leucurus\u003c/em\u003e) by radio telemetry and camera trapping: focusing on home-range and hibernation sett use pattern. J Korean Cadastre Inform Assoc 21:151\u0026ndash;163. (in Korean)\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCoonan TJ, Schwemm CA, Garcelon DK (2010) Decline and Recovery of the Island Fox: A Case Study for Population Recovery. Cambridge University Press, England.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDel Cerro I, Marmi J, Ferrando A, Chashchin P, Taberlet P, Bosch M (2010) Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora). Zool Scr 39(5):415\u0026ndash;425. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1463-6409.2010.00436.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1463-6409.2010.00436.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eErcoli MD, \u0026Aacute;lvarez A, Stefanini MI, Busker F, Morales MM (2015) Muscular anatomy of the forelimbs of the lesser grison (\u003cem\u003eGalictis cuja\u003c/em\u003e), and a functional and phylogenetic overview of Mustelidae and other Caniformia. J Mammal Evol 22:57\u0026ndash;91. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10914-014-9257-6\u003c/span\u003e\u003cspan address=\"10.1007/s10914-014-9257-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eEvans HE, De Lahunta A (2012) Miller's Anatomy of the Dog. Elsevier health sciences\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFabre AC, Cornette R, Slater G, Argot C, Peign\u0026eacute; S, Goswami A, Pouydebat E (2013) Getting a grip on the evolution of grasping in musteloid carnivorans: a three-dimensional analysis of forelimb shape. J Evol Biol 26(7):1521\u0026ndash;1535. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/jeb.12161\u003c/span\u003e\u003cspan address=\"10.1111/jeb.12161\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFaggi A, Bartolini-Lucenti S, Madurell-Malapeira J, Abramov AV, Puzachenko AY, Jiangzuo Q, Peiran L, Rook L (2024) Quaternary Eurasian badgers: Intraspecific variability and species validity. J Mammal Evol 31(1):3. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10914-023-09696-y\u003c/span\u003e\u003cspan address=\"10.1007/s10914-023-09696-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFedorov A, Beichel R, Kalpathy-Cramer J, Finet J, Fillion-Robin J-C, Pujol S, Bauer C, Jennings D, Fennessy F, Sonka M, Buatti J, Aylward SR, Miller JV, Pieper S, Kikinis R (2012) 3D Slicer as an Image Computing Platform for the Quantitative Imaging Network. JMRI 30(9):1323\u0026ndash;1341. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mri.2012.05.001\u003c/span\u003e\u003cspan address=\"10.1016/j.mri.2012.05.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFeeney S (1999) Comparative osteology, myology, and locomotor specializations of the fore and hind limbs of the North American foxes \u003cem\u003eVulpes vulpes\u003c/em\u003e and \u003cem\u003eUrocyon cinereoargenteus\u003c/em\u003e. Dissertation, University of Massachusetts Amherst\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFishbeck DW, Sebastiani A (2001) Comparative Anatomy: Manual of Vertebrate Dissection. Morton Publishing Company, Colarado\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFisher RE, Adrian B, Barton M, Holmgren J, Tang SY (2009) The phylogeny of the red panda (\u003cem\u003eAilurus fulgens\u003c/em\u003e): evidence from the forelimb. J Anat 215(6):611\u0026ndash;635. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-7580.2009.01156.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-7580.2009.01156.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFoster JB (1964) Evolution of mammals on islands. Nat 202(4929):234\u0026ndash;235. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/202234a0\u003c/span\u003e\u003cspan address=\"10.1038/202234a0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGiannico GR, Nagorsen DW (1989) Geographic and sexual variation in the skull of Pacific coast marten (\u003cem\u003eMartes americana\u003c/em\u003e). Can J Zool 67(6):1386\u0026ndash;1393. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1139/z89-197\u003c/span\u003e\u003cspan address=\"10.1139/z89-197\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGoltsman M, Kruchenkova EP, Sergeev S, Volodin I, Macdonald DW (2005) \u0026lsquo;Island syndrome\u0026rsquo;in a population of Arctic foxes (\u003cem\u003eAlopex lagopus\u003c/em\u003e) from Mednyi Island. J Zool 267(4):405\u0026ndash;418. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1017/S0952836905007557\u003c/span\u003e\u003cspan address=\"10.1017/S0952836905007557\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGupta S, Collier JS, Garcia-Moreno D, Oggioni F, Trentesaux A, Vanneste K et al (2017) Two-stage opening of the Dover Strait and the origin of island Britain. Nat Commun 8(1):15101. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/ncomms15101\u003c/span\u003e\u003cspan address=\"10.1038/ncomms15101\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHasegawa Y (2012) Origin of modern mammals in Japan. Mammal Sci 52(2):233\u0026ndash;247. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.11238/mammalianscience.52.233\u003c/span\u003e\u003cspan address=\"10.11238/mammalianscience.52.233\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. (in Japanese)\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHayashi S, Kubo MO, S\u0026aacute;nchez-Villagra MR, Taruno H, Izawa M, Shiroma T, Nakano T, Fujita M (2023) Variation and process of life history evolution in insular dwarfism as revealed by a natural experiment. Front Earth Sci 11:1095903. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/feart.2023.1095903\u003c/span\u003e\u003cspan address=\"10.3389/feart.2023.1095903\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHenderson K, Pantinople J, McCabe K, Richards HL, Milne N (2017) Forelimb bone curvature in terrestrial and arboreal mammals. PeerJ 5:e3229. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7717/peerj.3229\u003c/span\u003e\u003cspan address=\"10.7717/peerj.3229\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHildebrand M, Goslow GE (2001) Digging, and crawling without appendages. In: Hildebrand M, Goslow GE, Hildebrand V (eds) Analysis of Vertebrate Structure. John Wiley and Sons Inc, New Jersey, pp 455\u0026ndash;474\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eJanis CM, Mart\u0026iacute;n-Serra A (2020) Postcranial elements of small mammals as indicators of locomotion and habitat. PeerJ 8:e9634. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7717/peerj.9634\u003c/span\u003e\u003cspan address=\"10.7717/peerj.9634\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKaneko Y, Kanda E, Tashima S, Masuda R, Newman C, Macdonald DW (2014) The socio-spatial dynamics of the Japanese badger (\u003cem\u003eMeles anakuma\u003c/em\u003e). J Mammal 95(2):290\u0026ndash;300. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/12-MAMM-A-158\u003c/span\u003e\u003cspan address=\"10.1644/12-MAMM-A-158\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKaneko Y, Maruyama N, Macdonald DW (2006) Food habits and habitat selection of suburban badgers (\u003cem\u003eMeles meles\u003c/em\u003e) in Japan. J Zool 270(1):78\u0026ndash;89. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-7998.2006.00063.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-7998.2006.00063.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKawamura Y, Kamei T, Taruno H (1989) Middle and Late Pleistocene mammalian faunas in Japan. Quat Res 28(4):317\u0026ndash;326. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4116/jaqua.28.317\u003c/span\u003e\u003cspan address=\"10.4116/jaqua.28.317\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKholinne E, Zulkarnain RF, Sun YC, Lim S, Chun JM, Jeon IH (2018) The different role of each head of the triceps brachii muscle in elbow extension. Acta orthop traumatol turc 52(3):201\u0026ndash;205. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.aott.2018.02.005\u003c/span\u003e\u003cspan address=\"10.1016/j.aott.2018.02.005\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKim SI, Oshida T, Lee H, Min MS, Kimura J (2015) Evolutionary and biogeographical implications of variation in skull morphology of raccoon dogs (\u003cem\u003eNyctereutes procyonoides\u003c/em\u003e, Mammalia: Carnivora). Biol J Linn Soc 116(4):856\u0026ndash;872. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/bij.12629\u003c/span\u003e\u003cspan address=\"10.1111/bij.12629\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKoh HS, Kryukov A, Oh JG, Bayarkhagva D, Yang BG, Ahn NH, Bazarsad D (2014) Two sympatric phylogroups of the Asian badger \u003cem\u003eMeles leucurus\u003c/em\u003e (Carnivora: Mammalia) identified by mitochondrial DNA cytochrome b gene sequences. Russ J Theriol 13(1):1\u0026ndash;8. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.15298/rusjtheriol.13.1.01\u003c/span\u003e\u003cspan address=\"10.15298/rusjtheriol.13.1.01\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKowalczyk R, Jȩdrzejewska B, Zalewski A (2003) Annual and circadian activity patterns of badgers (\u003cem\u003eMeles meles\u003c/em\u003e) in Białowieża Primeval Forest (eastern Poland) compared with other Palaearctic populations. J Biogeogr 30(3):463\u0026ndash;472. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1046/j.1365-2699.2003.00804.x\u003c/span\u003e\u003cspan address=\"10.1046/j.1365-2699.2003.00804.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLaw CJ, Slater GJ, Mehta RS (2018) Lineage diversity and size disparity in Musteloidea: testing patterns of adaptive radiation using molecular and fossil-based methods. Syst Biol 67(1):127\u0026ndash;144. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/sysbio/syx047\u003c/span\u003e\u003cspan address=\"10.1093/sysbio/syx047\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLee MY, Lee SM, Song EG, An JH, Voloshina I, Chong JR, Johnson WE, Min M, Lee H (2016) Phylogenetic relationships and genetic diversity of badgers from the Korean Peninsula: Implications for the taxonomic status of the Korean badger. Biochem Syst Ecol 69:18\u0026ndash;26. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.bse.2016.07.015\u003c/span\u003e\u003cspan address=\"10.1016/j.bse.2016.07.015\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eL\u0026oacute;pez-Mart\u0026iacute;n JM, Ruiz-Olmo J, Padr\u0026oacute; I (2006) Comparison of skull measurements and sexual dimorphism between the Minorcan pine marten (\u003cem\u003eMartes martes minoricensis\u003c/em\u003e) and the Iberian pine marten (\u003cem\u003eM. m. martes\u003c/em\u003e): A case of insularity. Mamm Biol 71(1):13\u0026ndash;24. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambio.2005.08.006\u003c/span\u003e\u003cspan address=\"10.1016/j.mambio.2005.08.006\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLyras GA, van der Geer AA, Rook L (2010) Body size of insular carnivores: evidence from the fossil record. J Biogeogr 37(6):1007\u0026ndash;1021. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2699.2010.02312.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2699.2010.02312.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMadurell-Malapeira J, Mart\u0026iacute;nez-Navarro B, Ros-Montoya S, Espigares MP, Toro I, Palmqvist P (2011) The earliest European badger (\u003cem\u003eMeles meles\u003c/em\u003e), from the late villafranchian site of Fuente Nueva 3 (Orce, granada, se iberian peninsula). Comptes Rendus Palevol 10(8):609\u0026ndash;615. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.crpv.2011.06.001\u003c/span\u003e\u003cspan address=\"10.1016/j.crpv.2011.06.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMarmi J, Lopez-Giraldez F, Macdonald DW, Calafell F, Zholnerovskaya E, Domingo‐Roura X (2006) Mitochondrial DNA reveals a strong phylogeographic structure in the badger across Eurasia. Mol Ecol 15(4):1007\u0026ndash;1020. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-294X.2006.02747.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-294X.2006.02747.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMarneweck CJ, Allen BL, Butler AR, Do Linh San E, Harris SN, Jensen AJ et al (2022) Middle-out ecology: small carnivores as sentinels of global change. Mamm Rev 52(4):471\u0026ndash;479. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/mam.12300\u003c/span\u003e\u003cspan address=\"10.1111/mam.12300\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMasseti M (1995) Quaternary biogeography of the Mustelidae family on the Mediterranean islands. HYSTRIX 7:17\u0026ndash;34. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4404/hystrix-7.1-2-4048\u003c/span\u003e\u003cspan address=\"10.4404/hystrix-7.1-2-4048\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMeiri S Dayan T, Simberloff D (2004) Body size of insular carnivores: little support for the island rule. The Am Nat 163(3):469\u0026ndash;479. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1086/382229\u003c/span\u003e\u003cspan address=\"10.1086/382229\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMonroy-Cendales MJ, V\u0026eacute;lez‐Garc\u0026iacute;a JF, Casta\u0026ntilde;eda‐Herrera FE (2020) Gross anatomy of the shoulder and arm intrinsic muscles in the white‐footed tamarin (\u003cem\u003eSaguinus leucopus\u003c/em\u003e\u0026ndash;G\u0026uuml;nther, 1876): Inter- and intraspecific anatomical variations. J Med Primatol 49(3):123\u0026ndash;135. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/jmp.12465\u003c/span\u003e\u003cspan address=\"10.1111/jmp.12465\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMonroy-Cendales MJ, V\u0026eacute;lez‐Garc\u0026iacute;a JF, Casta\u0026ntilde;eda‐Serrano RD, Miglino MA (2023) Gross anatomical description of the extrinsic and intrinsic scapular and brachial muscles of \u003cem\u003eSapajus apella\u003c/em\u003e (Linnaeus, 1758). J Med Primatol 52(1):3\u0026ndash;16. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/jmp.12619\u003c/span\u003e\u003cspan address=\"10.1111/jmp.12619\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMoore A (2011) Functional specialization in the intrinsic forelimb musculature of the American badger \u003cem\u003e(Taxidea taxus)\u003c/em\u003e. Dissertation, Youngstown State University\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMoore AL, Budny JE, Russell AP, Butcher MT (2013) Architectural specialization of the intrinsic thoracic limb musculature of the American badger (\u003cem\u003eTaxidea taxus\u003c/em\u003e). J Morphol 274(1):35\u0026ndash;48. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/jmor.20074\u003c/span\u003e\u003cspan address=\"10.1002/jmor.20074\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMoore CM, Collins PW (1995) \u003cem\u003eUrocyon littoralis\u003c/em\u003e. Mamm Spec 489:1\u0026ndash;7. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/3504160\u003c/span\u003e\u003cspan address=\"10.2307/3504160\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNanova O, Pr\u0026ocirc;a M (2017) Cranial features of mainland and Commander Islands (Russia) Arctic foxes (\u003cem\u003eVulpes lagopus\u003c/em\u003e) reflect their diverging foraging strategies. Polar Res 36:7 \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/17518369.2017.1310976\u003c/span\u003e\u003cspan address=\"10.1080/17518369.2017.1310976\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNeal EG, Cheeseman CL (1996) Badgers. Poyster Natural History, London\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eOh JG (2007) Behavioral Characteristics of Jeju Badgers. Hallasan Research Institute Research Report:37\u0026ndash;49\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePerdomo-C\u0026aacute;rdenas V, Pati\u0026ntilde;o‐Holgu\u0026iacute;n C, V\u0026eacute;lez‐Garc\u0026iacute;a JF (2021) Evolutionary and terminological analysis of the flexor digitorum superficialis, interflexorii and palmaris longus muscles in kinkajou (\u003cem\u003ePotos flavus\u003c/em\u003e) and crab‐eating racoon (\u003cem\u003eProcyon cancrivorus\u003c/em\u003e). Anat Histol Embryol 50(3):520\u0026ndash;533. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/ahe.12656\u003c/span\u003e\u003cspan address=\"10.1111/ahe.12656\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eProulx G, Abramov AV, Adams I, Jennings AP, Khorozyan I, Rosalino LM et al (2016) World distribution and status of badgers\u0026ndash;A review. In: Proulx G, Do Linh San, E (eds) Badgers: Systematics, Biology, Conservation and Research Techniques. Alpha Wildlife Publications, Canada, pp 31\u0026ndash;116\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRose J, Moore A, Russell A, Butcher M (2014) Functional osteology of the forelimb digging apparatus of badgers. J Mammal 95(3):543\u0026ndash;558. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/13-MAMM-A-174\u003c/span\u003e\u003cspan address=\"10.1644/13-MAMM-A-174\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRozzi R, Lomolino MV, van der Geer AA, Silvestro D, Lyons SK, Bover P et al (2023) Dwarfism and gigantism drive human-mediated extinctions on islands. Science 379(6636):1054\u0026ndash;1059. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1126/science.add8606\u003c/span\u003e\u003cspan address=\"10.1126/science.add8606\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSalesa MJ, Anton M, Peigne S, Morales J (2008) Functional anatomy and biomechanics of the postcranial skeleton of \u003cem\u003eSimocyon batalleri\u003c/em\u003e (Viret, 1929)(Carnivora, Ailuridae) from the Late Miocene of Spain. Zool J Linn Soc 152(3):593\u0026ndash;621. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1096-3642.2007.00370.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1096-3642.2007.00370.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSalesa MJ, Siliceo G, Ant\u0026oacute;n M, Fabre AC, Pastor JF (2020) Functional inferences on the long bones of \u003cem\u003eIschyrictis zibethoides\u003c/em\u003e (Blainville, 1841)(Carnivora, Mustelidae) from the middle Miocene locality of Sansan (Gers, France). Geodiversitas 42(1):1\u0026ndash;16. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.5252/geodiversitas2020v42a1\u003c/span\u003e\u003cspan address=\"10.5252/geodiversitas2020v42a1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSamuels JX, Meachen JA, Sakai SA (2013) Postcranial morphology and the locomotor habits of living and extinct carnivorans. J Morphol 274(2):121\u0026ndash;146. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/jmor.20077\u003c/span\u003e\u003cspan address=\"10.1002/jmor.20077\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSavvidou A, Youlatos D, Spassov N, Tamvakis A, Kostopoulos DS (2022) Ecomorphology of the Early Pleistocene Badger \u003cem\u003eMeles dimitrius\u003c/em\u003e from Greece. J Mammal Evol 29(3):585\u0026ndash;607. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10914-022-09609-5\u003c/span\u003e\u003cspan address=\"10.1007/s10914-022-09609-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eShikama T (1949) The Kuzuu Ossuaries, Geological and Palaeontological Studies of the Limestone Fissure Deposits in Kuzuu, Totigi Prefecture. Sci Rep Tohoku Uni II Ser 23: 1\u0026ndash;209.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eShikama T (1962) Quaternary Land Connections of Japanese Islands with Continent from the Viewpoints of Palaeomammalogy. Quat Res 2(4\u0026ndash;5):146\u0026ndash;153. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4116/jaqua.2.146\u003c/span\u003e\u003cspan address=\"10.4116/jaqua.2.146\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSicuro FL, Oliveira LFB (2015) Variations in leopard cat (\u003cem\u003ePrionailurus bengalensis\u003c/em\u003e) skull morphology and body size: sexual and geographic influences. PeerJ 3:e1309. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7717/peerj.1309\u003c/span\u003e\u003cspan address=\"10.7717/peerj.1309\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSleeman DP, Davenport J (2016) Evidence of poor diet quality in the small-bodied Eurasian Badgers (\u003cem\u003eMeles meles\u003c/em\u003e) on Rutland Island, Co. Donegal. Ir Nat J 35(1):22\u0026ndash;26.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSleeman DP, Davenport J, Cussen RE, Hammond RF (2009) The small-bodied badgers (\u003cem\u003eMeles meles\u003c/em\u003e (L.)) of Rutland Island, Co. Donegal. Ir Nat J 30(1):1\u0026ndash;6.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eTanaka H (2006) Winter hibernation and body temperature fluctuation in the Japanese badger, \u003cem\u003eMeles meles anakuma\u003c/em\u003e. Zool Sci 23(11):991\u0026ndash;997. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2108/zsj.23.991\u003c/span\u003e\u003cspan address=\"10.2108/zsj.23.991\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eTanaka H, Fukuda Y, Yuki E, Ota Y, Hosoi E, Kojima W (2022) Cooperative den maintenance between male Japanese badgers that are delayed dispersers and their mothers. J Ethol 40(1):3\u0026ndash;11. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10164-021-00718-x\u003c/span\u003e\u003cspan address=\"10.1007/s10164-021-00718-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eTaylor ME (1974) The functional anatomy of the forelimb of some African Viverridae (Carnivora). J Morphol 143(3):307\u0026ndash;335. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/jmor.1051430305\u003c/span\u003e\u003cspan address=\"10.1002/jmor.1051430305\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eVan Valkenburgh B (1987) Skeletal indicators of locomotor behavior in living and extinct carnivores. J Vertebr Paleontol 7(2):162\u0026ndash;182. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/02724634.1987.10011651\u003c/span\u003e\u003cspan address=\"10.1080/02724634.1987.10011651\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eV\u0026eacute;lez-Garc\u0026iacute;a JF, Blanco DAC, G\u0026oacute;mez GM, Ca\u0026ntilde;as SSM (2023) Descriptive study of the intrinsic muscles of the shoulder and brachium in kinkajou (\u003cem\u003ePotos flavus\u003c/em\u003e) and an evolutionary analysis within the suborder Caniformia. Vertebr Zool 73:957\u0026ndash;980. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3897/vz.73.e102645\u003c/span\u003e\u003cspan address=\"10.3897/vz.73.e102645\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eV\u0026eacute;lez Garc\u0026iacute;a JF, de Carvalho Barros RA, Miglino MA (2025) Is the articularis humeri muscle homologous to the coracobrachialis muscle in carnivorans? An evolutionary and terminological answer based on the shoulder myology of the coati (\u003cem\u003eNasua nasua\u003c/em\u003e, Carnivora, Procyonidae). Anat, Histol, Embryol 54(2):e70034. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/ahe.70034\u003c/span\u003e\u003cspan address=\"10.1111/ahe.70034\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eWayne RK, George SB, Gilbert D, Collins PW, Kovach SD, Girman D, Lehman N (1991) A morphologic and genetic study of the island fox, \u003cem\u003eUrocyon littoralis\u003c/em\u003e. Evolution 45(8):1849\u0026ndash;1868. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1558-5646.1991.tb02692.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1558-5646.1991.tb02692.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eWindle BCA, Parsons FG (1897) On the Myology of the Terrestrial Carnivora. Part I: muscles of the head, neck, and fore-limb. Proc Zool Soc Lond 65:370\u0026ndash;409. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-7998.1897.tb00023.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-7998.1897.tb00023.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eYousefi MH, Rasouli B, Ghodrati S, Adibi MA, Taherdoost M, Omidbakhsh S, Behnam G (2018) Anatomical study of extrinsic and some intrinsic muscles of the thoracic limb in Iranian pine marten (\u003cem\u003eMartes martes\u003c/em\u003e): a case report. Iran J Vet Med 12(3):273\u0026ndash;282. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.22059/ijvm.2018.252150.1004876\u003c/span\u003e\u003cspan address=\"10.22059/ijvm.2018.252150.1004876\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-mammalian-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jomm","sideBox":"Learn more about [Journal of Mammalian Evolution](http://link.springer.com/journal/10914)","snPcode":"10914","submissionUrl":"https://submission.nature.com/new-submission/10914/3","title":"Journal of Mammalian Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Comparative osteology, forelimb myology, Meles, insular populations, island adaptations","lastPublishedDoi":"10.21203/rs.3.rs-7041610/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7041610/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eIsland colonization can drive anatomical adaptations in mammals, yet few studies have examined how such changes occur in limb musculature and osteology. Here, we investigate the forelimb anatomy of \u003cem\u003eMeles\u003c/em\u003e badgers to understand functional adaptations associated with insular environments. We conducted detailed myological dissections of the Japanese badger (\u003cem\u003eMeles anakuma\u003c/em\u003e) and its continental relative (\u003cem\u003eMeles leucurus\u003c/em\u003e), and compared osteological features with another continental-insular pair: \u003cem\u003eMeles meles\u003c/em\u003e from mainland Europe and \u003cem\u003eMeles canescens\u003c/em\u003e from Crete. Our results reveal several previously undescribed features, including the presence of m. tensor fasciae antebrachii in \u003cem\u003eMeles\u003c/em\u003e, and a distal shift in the main belly of m. coracobrachialis in \u003cem\u003eM. anakuma\u003c/em\u003e, both with considerable phylogenetic ramifications. While body size did not differ significantly between \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. leucurus\u003c/em\u003e, insular populations of both \u003cem\u003eM. anakuma\u003c/em\u003e and \u003cem\u003eM. canescens\u003c/em\u003e exhibit shared osteological traits, such as reduced projection of the humeral trochlea and enlargement of the cranial aspect of the ulnar head, which likely enhance forelimb mobility rather than maximize digging force. These results may indicate that insular \u003cem\u003eMeles\u003c/em\u003e have evolved increased joint flexibility, potentially as an adaptation for locomoter versatility in more structurally complex or variable island habitats. Our findings establish \u003cem\u003eMeles\u003c/em\u003e as a valuable model for studying evolutionary responses to island environments in subfossorial mesocarnivores.\u003c/p\u003e","manuscriptTitle":"Enhanced Forelimb Mobility in Insular Meles populations: Insights from Myological and Osteological Comparisons","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-15 17:18:39","doi":"10.21203/rs.3.rs-7041610/v1","editorialEvents":[{"type":"communityComments","content":1},{"type":"decision","content":"Revision requested","date":"2025-08-05T00:44:27+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-02T03:33:26+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-30T16:48:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"271651386692123044207922989066507528061","date":"2025-07-10T22:29:42+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-10T18:15:22+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"101301014113185659305528845654298339266","date":"2025-07-10T18:06:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"295482259172025838118697721278855897732","date":"2025-07-09T20:08:04+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-09T19:51:39+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-07-04T04:54:37+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-07-04T04:53:50+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Mammalian Evolution","date":"2025-07-03T22:37:22+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-mammalian-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jomm","sideBox":"Learn more about [Journal of Mammalian Evolution](http://link.springer.com/journal/10914)","snPcode":"10914","submissionUrl":"https://submission.nature.com/new-submission/10914/3","title":"Journal of Mammalian Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"2475a00e-4acb-438e-a717-c59d60ef0ae1","owner":[],"postedDate":"July 15th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-05-05T10:02:12+00:00","versionOfRecord":{"articleIdentity":"rs-7041610","link":"https://doi.org/10.1007/s10914-025-09796-x","journal":{"identity":"journal-of-mammalian-evolution","isVorOnly":false,"title":"Journal of Mammalian Evolution"},"publishedOn":"2026-05-02 15:58:38","publishedOnDateReadable":"May 2nd, 2026"},"versionCreatedAt":"2025-07-15 17:18:39","video":"","vorDoi":"10.1007/s10914-025-09796-x","vorDoiUrl":"https://doi.org/10.1007/s10914-025-09796-x","workflowStages":[]},"version":"v1","identity":"rs-7041610","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7041610","identity":"rs-7041610","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}