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Accepted Manuscript Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity Nicole Capdarest-Arest, Jorge P. Gonzalez, Tolga Turker PII: S0306-9877(14)00032-2 DOI: http://dx.doi.org/10.1016/j.mehy.2014.01.021 Reference: YMEHY 7484 To appear in: Medical Hypotheses Received Date: 23 August 2013 Accepted Date: 21 January 2014 Please cite this article as: N. Capdarest-Arest, J.P. Gonzalez, T. Turker, Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity, Medical Hypotheses (2014), doi: http://dx.doi.org/10.1016/j.mehy.2014.01.021 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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Accepted Manuscript

Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity

Nicole Capdarest-Arest, Jorge P. Gonzalez, Tolga Turker

PII: S0306-9877(14)00032-2DOI: http://dx.doi.org/10.1016/j.mehy.2014.01.021Reference: YMEHY 7484

To appear in: Medical Hypotheses

Received Date: 23 August 2013Accepted Date: 21 January 2014

Please cite this article as: N. Capdarest-Arest, J.P. Gonzalez, T. Turker, Hypotheses for Ongoing Evolution ofMuscles of the Upper Extremity, Medical Hypotheses (2014), doi: http://dx.doi.org/10.1016/j.mehy.2014.01.021

This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customerswe are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, andreview of the resulting proof before it is published in its final form. Please note that during the production processerrors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity 3

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1. NC; Nicole Capdarest-Arest, MA 5

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2. JPG; Jorge P Gonzalez 9

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3. TT; Tolga Turker, MD 13

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University of Arizona Health Sciences Center 18

The University of Arizona 19

1501 N. Campbell Ave., Room 4143C 20

Tucson, AZ 85724, USA 21

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*Correspondence to: 24

Tolga Turker, MD 25

Division of Reconstructive & Plastic Surgery 26

The University of Arizona Health Sciences Center 27

1501 N. Campbell Ave., Room 4143C 28

Tucson, AZ 85724, USA 29

Email: [email protected] 30

Telephone: 1-520-404-6850 31

Fax: 1-520-626-3330 32

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No support was received for this study. 35

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Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity

Abstract

There are organs and muscles in the human body that may be considered

rudimentary in that they have insignificant or undetermined function. Several such

muscles are found in the upper extremity. In this review, four muscles that appear

to be undergoing evolutionary changes are discussed: flexor digitorum superficialis

to the fifth finger, anconeus, palmaris longus, and anconeus epitrochlearis. The

present study synthesizes, advances and extends previously described work about

these muscles and extends the hypotheses and conclude that: (a) the flexor

digitorum superficialis to the fifth finger is currently under adaptive evolution, (b)

the anconeus has currently stabilized its evolution and is serving as a transient

stability augmenter during a short portion of the human lifespan, and (c) the entire

distal upper extremity is currently in the process of undergoing evolutionary

change. Understanding of these muscles and their evolutionary context is important

for understanding of impact on function, dysfunction, treatment and future

research.

Key Words: anatomy, anconeus, anconeus epitrochlearis, evolution, flexor

digitorum superficialis-V, forearm, morphology, palmaris longus.

*Abstract

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Hypotheses for Ongoing Evolution of Muscles of the Upper Extremity 38

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INTRODUCTION 40

As famously described by Charles Darwin and other scientists, there are organs and 41

muscles in the human body that are “rudimentary” in nature, namely they have uncertain 42

or no known current function(1). Additionally, natural selection also influences human 43

development and morphology by encouraging more economic and precise function 44

resulting in either agenesis or genesis of muscle. Accordingly, multiple reports of variations 45

in current human anatomy resulting from such morphology have been described. Studies 46

to date have focused on the physical description of individual muscle variations and 47

possible functional significance. However, four muscles in the upper extremity not only 48

show such variations, but, due to their demonstrated change in function, we hypothesize 49

that the entire distal upper extremity is currently in the process of evolutionary change. 50

These four muscles are: flexor digitorum superficialis to the fifth finger (FDS-V), anconeus, 51

palmaris longus, and anconeus epitrochlearis (AE). Two of these muscles already appear to 52

be in a rudimentary stage, one appears to be undergoing adaptive evolution, and one is 53

stabilized and acting as a transient stability augmenter. The present study synthesizes, 54

advances and extends previously described work about these muscles and extend the 55

hypotheses that: (a) the FDS-V is currently under adaptive evolution, (b) the anconeus has 56

currently stabilized its evolution and is serving as a transient stability augmenter during a 57

short portion of the human lifespan, and (c) the entire distal upper extremity is currently in 58

the process of undergoing evolutionary change. 59

60

3

HYPOTHESES 61

Quadrupedal or arboreal locomotion to habitual upright bipedal locomotion has caused 62

changes to the necessary musculature of the upper extremity. Muscles that, in the past, 63

may have been useful for quadrupedal or arboreal locomotion and fewer fine motor tasks 64

may be no longer as useful in the modern environment. We hypothesize that evolution is 65

currently underway in the human forearm, as demonstrated by four muscles that are in the 66

midst of evolutionary change or have recently evolved to their current presentation. These 67

four muscles are: (a) FDS-V, (b) anconeus, (c) palmaris longus, and (d) AE. From our review 68

of the FDS-V, we hypothesize that the incidence of FDS-V in humans will continue to 69

increase in order to perform fine tasks with precise range of motion. Regarding the 70

anconeus muscle, which is always present in humans and is currently thought to be an 71

extension of the triceps muscle with unknown current function, we hypothesize that this 72

muscle has a primary function in infants as a stabilizer during the relatively brief period of 73

human development where infants crawl and that it has currently stabilized evolutionarily. 74

As humans grow, learn to walk and the elbow joint finishes developing in later childhood, 75

we postulate that the anconeus takes on a more accessory-type role, which explains why 76

the literature to date does not elucidate function of this muscle. Finally, we hypothesize 77

that the entire distal upper extremity is currently in the process of undergoing 78

evolutionary changes, as evidenced by the adaptive changes of FDS-V, the stabilization of 79

the anconeus, and other changes occurring in the palmaris longus and the AE. 80

81

4

DISCUSSION 82

Flexor Digitorum Superficialis-V 83

The FDS-V is another muscle that has variable absence in humans. This muscle starts from 84

the forearm and it inserts onto the radial and ulnar aspects of the proximal half of the 85

middle phalanx of the fifth finger when it is present and is enervated by the median 86

nerve(2). This muscle has been shown to be absent in 2% of the Japanese population(3), 87

and others have reported its absence to be 6% (bilateral) and 6.8% (unilateral)(4). 88

Contradictory hypotheses for the development of the FDS muscle belly(3) have been 89

proposed, single origin (either antebrachial or palmar) and dual origin (both antebrachial 90

and palmar). Kobayashi et al. referred to Yamada’s supposition that the FDS muscle 91

originates first in the palm and then migrates to the forearm(3) due to the results of their 92

study as well as another reporting “brevis-type” variations of the FDS to the finger in 93

question. It does not seem that “brevis-type” variations would be possible with a single 94

origin of the FDS-V. However, the FDS muscle motor nerve is distributed in the forearm 95

level and there is no recurrent branch that may have originated in the palm and goes back 96

to the forearm for FDS innervation described yet. It is possible that the muscle portion of 97

FDS comes from the forearm and the tendon portions originate from the hand because of 98

previously described anomalies and the motor branch of the muscle. Furthermore, 99

Shrewsbury and Kuczynski(5) noted that the FDS-V was absent in about 20% of the studied 100

population. Interestingly, even if the tendon of FDS-V is absent, the distal components of 101

the tendon are present(5). We have similarly noted an instance of a patient without an FDS 102

but possessing intact distal components. Such instances are important for clinicians to be 103

aware of in the event they encounter distal tendinous portions but not the FDS-V tendon 104

5

itself, and may also support the theory that this muscle may have a dual origin and is 105

formed in the hand first(5). It seems that, from research done so far, the dual-origin 106

hypothesis may make more sense. 107

108

Comparing humans to other species, limbed amphibians and reptiles are similar to 109

mammals in that they have flexor digitorum longus muscles. However, mammals have 110

developed two layers in order to flex the digits: the flexor digitorum profundus and the 111

FDS(6). The short hand muscles seen in reptiles and amphibians may have evolved 112

proximally in mammals since there is no relation between palmaris longus and the FDS in 113

reptiles and amphibians since the FDS location in the hand is replaced by a set of short, 114

superficial finger flexors(7). In this case, Kobayashi et al. may be correct in restating 115

Yamada’s assertion that the FDS muscle originates first in the palm and then migrates to 116

the forearm, however the nerves still emanate proximally(3). Intuitively, it makes sense for 117

more primitive species to retain muscles in the hands because they do not use their hands 118

for precise tool handling. Having less bulky muscles in the hand would be an advantageous 119

evolutionary development for improved tool handling capabilities. In support of this 120

theory, one may note that the fully oppositional motion of the human thumb is made 121

possible by the distinct nature of the double saddle-shaped design of the first 122

carpometacarpal joint. The first carpometacarpal joint is comprised of two articulating 123

saddle-shaped bones facing one another, with the saddles meeting to form an “X”. Because 124

of this unique configuration, the thenar area has two fewer muscles than would be present 125

if the joint was otherwise shaped and thus keeps the hand less bulky(8). Some might argue 126

that since the FDS-V is likely a less frequently used FDS muscle (when compared to FDS to 127

6

the index finger, for example), it is less economically important to humans and we may 128

continue to see more agenesis of this muscle; however, for the reasons outlined below, we 129

hypothesize that, conversely, we will see increasing genesis of the FDS-V. 130

131

Other research has also confirmed the variability of the FDS-V in humans. In a study on 70 132

cadaveric hands, it was found that 13% of the hands had anatomical variations for the FDS-133

V(9). Additionally, the variations noted were mostly irregular themselves, such as unusual 134

variants of the FDS decussation and even complete absence of FDS muscles(9). Regarding 135

the impact of cases in which the FDS-V is completely absent, research has indicated that the 136

relatively common absence of this muscle can impact grip strength(10). Out of 171 137

subjects, it was found that the FDS-V was absent in 18.6% of females and 15.3% of males 138

and in those subjects, grip strength was significantly lower than in subject groups with 139

independent or common (attached to FDS of fourth finger) function of FDS-V(10). Another 140

study, in contrast, found that there was no significant difference seen in grip strength 141

between subjects who had an FDS-V and those who did not(11). Despite the wide 142

variability of the presence of and the conflicting information as to the functional impact of 143

the FDS-V, current clinical examination techniques are inadequate to discriminate among 144

the possible variations or absence of FDS(12). There are also several muscle tendon 145

variations that have been described in the flexor compartment of the forearm and, 146

interestingly, most variations were related to the fifth finger(12). It is possible that the 147

hand is evolving to either have decreased incidence of FDS-V or, more likely, it is possible 148

that the incidence of FDS-V in humans will continue to increase in order to perform fine 149

tasks (e.g., playing the piano or typewriting) or with precise range of motion, as humans 150

7

often do in the modern environment. In this case, muscle tendon variations may be 151

considered atavistic, in that they appear to represent a more primitive evolutionary 152

presentation. 153

Anconeus 154

The anconeus is a muscle that originates from the lateral epicondyle of humerus and makes 155

an insertion at the posterior olecranon process of the ulna. The radial nerve acts as the 156

motor branch for this muscle(2). The anconeus is a muscle common to many species, is 157

present in all mammals(6), and presents with similar anatomical location and possible 158

proposed action in chimpanzees and Rhesus macaques. In other primates, the anconeus 159

sometimes appears as a distinct muscle and in others appears as ill defined or a 160

continuation of the triceps. Although the function of anconeus is unclear, it is linked with 161

the extension of the forearm at the elbow in all non-human primates. 162

163

There is not clear consensus about function of the anconeus in humans and multiple 164

studies have been conducted with varying conclusions. Gleason et al.’s EMG study supports 165

Duchenne’s original proposal for anconeus function, namely, that the anconeus abducts the 166

ulna during pronation of the forearm. They observed electric silence in the muscle in 167

flexion and extension of the elbow however they were able to show electric activity while 168

the forearm pronates the axis of the second digit(13). Another EMG study performed on ten 169

volunteers showed some muscle activity during both pronation and supination, but the 170

researchers could not conclude the function of the anconeus(14). They postulated, 171

however, that the muscle most likely acts as lateral stabilizer of the elbow joint(14). 172

Another study suggested that the anconeus muscle is one of the elbow extensors along with 173

8

the triceps and flexor carpi ulnaris muscles, however the role of the anconeus in elbow 174

stabilization was not mentioned(15). The anconeus was also thought to be an important 175

dynamic elbow stabilizer among triceps and brachialis according to O’Driscoll et al(16), 176

and Molinier et al.’s(17) anatomy study shows close relation of the anconeus to the lateral 177

head of the triceps muscle and they considered the two muscles to act synergistically. 178

Additionally, they mentioned that the anconeus provides lateral stability for the extended 179

elbow joint due to the close relationships between (i) the triceps and the anconeus and (ii) 180

the joint capsule and the anconeus(17). 181

182

This proposed theory for the function of the anconeus muscle as a stabilizer for the elbow 183

may be explained with mammalian humeroulnar joint evolutional theory explained by 184

Jenkins(18). According to Jenkins, the pelycosaur humeroulnar joint, which is a less 185

constrained joint than the human humeroulnar joint, has a torsional stress due to humeral 186

rotation and the joint may disarticulate under torque. In order to prevent dislocation, the 187

ulna rotates in conjunction with the humerus. As mammals evolved in the Jurassic period, 188

the joint becomes more constrained and the forearm remains parallel to the humerus, 189

which does a complex motion of rotation and adduction. Evolution continued so the joint 190

becomes even more constrained and deep with a well-defined trochlea in nonhuman 191

primates(18). 192

193

If the humeroulnar joint follows a common frame evolutionarily, it may be that in the 194

earlier stages of tetrapod evolution, the joint was shallower requiring more active dynamic 195

stabilization and, therefore, the anconeus muscle might have played a more primary role in 196

9

earlier stages of humeroulnar joint evolution. In this way, the anconeus may have kept the 197

forearm tracking properly while the humerus underwent the complex rotational motion 198

with the tetrapodal gait. If this theory is correct, humans may not need this muscle 199

anymore since the joint itself is anatomically stable. We postulate that it could be that the 200

anconeus plays a more important role as a stabilizer in infancy during the relatively brief 201

period of human development where infants crawl. As humans grow, learn to walk and the 202

elbow joint finishes developing in later childhood, perhaps the anconeus takes on this more 203

accessory-type role. The current literature does not have a comprehensive discussion of 204

humeroulnar joint biomechanics during crawling and this may be an area meriting further 205

research. It may be that the humeroulnar joint may be a relatively young joint 206

evolutionarily. Due to the relative recent development of this joint and the transition to 207

bipedalism, perhaps the anconeus will continue to adapt into a more rudimentary stage, 208

unless there are stresses placed on it that stabilize the joint during a crucial period of 209

human growth. In order to demonstrate whether or not the anconeus actually has any 210

current function in modern humans, it may be informative to research and measure 211

function of this muscle in crawling infants. 212

213

Evolution of the Distal Upper Extremity 214

Given the adaptive changes of the FDS-V and the current stabilization of the anconeus as 215

hypothesized above, we also postulate that, due to these and additional changes occurring 216

in the palmaris longus and the AE, the forearm as a whole is currently undergoing adaptive 217

evolutionary change. The palmaris longus is a muscle that mostly appears in the human 218

populations, varying among ethnic groups. This muscle is reported as one of the most 219

10

variable muscles in the human body(19); its absence is reported between 3% and 220

63.9%(20). This muscle originates at the medial epicondyle of the humerus and inserts to 221

the palmar aponeurosis. The median nerve acts as the motor branch for the palmaris 222

longus(2). The function of the palmaris longus is uncertain in humans, but it is generally 223

considered to be a contributor for flexing the hand at the wrist and tensing the palmar 224

aponeurosis. Additionally, it is a well-known common option for use in tendon grafts. 225

Most forearm muscles have a large and long muscle belly filling most of the length of the 226

forearm and then, distally, the muscle becomes tendon 3-4 cm proximal of the wrist joint. 227

The palmaris longus when present, however, has a long tendon and a short muscle belly 228

[Figure 1], although variations have been found in humans wherein the muscle belly is 229

more or less formed, attachment sites are varied, or accessory tendinous slips are present. 230

Additionally, absence or presence of palmaris longus does not influence flexion of the 231

wrist(21). These anatomical properties make the palmaris longus a popular choice for 232

reconstructive purposes. 233

234

So if the palmaris longus does not have any significant function since humans can perform 235

the same tasks regardless of whether they have palmaris longus, why do most humans 236

have this muscle? To explore the answer this question we looked at human evolution and 237

function of this muscle in different species. In orangutans, the only strictly arboreal ape, the 238

palmaris longus is always present(22). Having a strong palmaris longus muscle might 239

therefore be more important in species such as orangutans that perform most daily 240

activities in trees or use the forelimbs for ambulation, providing improved ability to grip 241

and move in trees or with the forelimbs, and may be evolutionarily less important in other 242

11

primates or in humans who are mostly terrestrial. The lower incidence of palmaris longus 243

in other primates, who are more or completely terrestrial, may support this. 244

245

In other species, the palmaris longus is also variably found. Abdala and Diogo(6) performed 246

a comprehensive comparison of anatomy among several species. They showed that 247

salamanders, crocodiles, chickens and frogs do not have palmaris longus, however, 248

semiaquatic turtles and rats have palmaris longus and some lizards also have it(6). 249

According to the framework described by Abdala and Diogo, it makes sense for mammals 250

have the muscle, but why? We postulate that since evolution wants to economize energetic 251

output, it is possible that in the past, it may have been advantageous to have palmaris 252

longus, but this may no longer be the case. However, according to current anatomical 253

studies and the frequency of occurrence of the palmaris longus in humans and other 254

species, it is likely that the palmaris longus is under regression and becoming a 255

rudimentary muscle. This might be evidenced by the increase in size of the muscle bellies 256

of the other muscles of the forearm and the relatively much smaller size of the palmaris 257

longus. Additionally, although palmaris longus may vary among humans as described 258

above, in general the smaller muscle belly and incidence of agenesis in humans is in 259

contrast to the higher incidence and larger muscle belly in lower primates, mammals and 260

other species. One can make the assumption that, if human evolution continues along 261

similar lines wherein the muscle belly continues to phylogenetically reduce, it is expected 262

that this muscle will eventually not be found in humans(23). 263

264

12

The AE, also known as epitrochleo-anconeus or anconeus sextus, is a muscle found in a 265

variety of species. In humans it originates from the inferior surface of the medial 266

epicondyle, crosses the ulnar nerve, and inserts on the olecranon. It is innervated from the 267

ulnar nerve(24). The muscle function varies in different species, and the function in 268

humans is unclear. In humans the AE is sometimes referred to as an independent muscle 269

and sometimes as an accessory muscle or factor of the triceps brachii, whereas in other 270

mammals it is always an independent structure. It is suggested that the muscle (i) serves to 271

keep the ulnar nerve in position and guard the vessels that accompany it from 272

pressure(24), and (ii) acts to assist the triceps brachii and the ligamentum cubiti mediale to 273

support the median aspect of the elbow joint(25). This muscle presents with similar 274

undifferentiated anatomic structure in gorillas, orangutans, and most other primates(26). 275

276

According to Dellon, the AE exists in only 11% of the human population, and the Osborne 277

ligament is taking its place due to the increasing amount of work humans perform with a 278

flexed elbow position. The Osborne ligament would provide more stability for the elbow 279

when performing tasks in flexion(24). Husarik et al. performed an MRI study on sixty 280

patients with asymptomatic elbows and found that 23% had an AE muscle(27). In Galton’s 281

study published in 1874(25), he performed dissections in Edentata and stated that this 282

muscle was always present and well developed. He also declared that this muscle is not 283

always seen in bats and occurs rarely in hoofed animals, but mentioned that the AE is seen 284

very often in other mammals. In addition, he stated that it seen less frequently among the 285

lower-order primates, is not seen in anthropoid apes, and seen only occasionally in humans 286

as an anomaly(25). When a comparison of the AE’s presence among the different species 287

13

with an emphasis on the evolutionary track of the muscle is taken, there appears to be a 288

reduction in the frequency of the occurrence of the muscle. There is a higher presence in 289

the lower-order monkeys and lemurs, although it is not universally present among 290

these(25). In any case, it seems to become lost among the anthropoid apes, and occurs 291

again infrequently in humans, though called a variation(27) or anomaly(25). 292

293

There is evidence that the AE is becoming or may be already considered rudimentary as its 294

structure is retained from an earlier and more primitive condition of existence while in 295

other mammals it is a necessary and functional mechanical appendage of the elbow 296

joint(24). As over 70% of the human population does not even have this muscle(27), one 297

can see that the muscle must not be necessary in humans and may be indicative of a late 298

evolutionary process. Indeed, presence of an AE may not only be unnecessary, but may 299

cause problems such as ulnar neuropathy, cubital tunnel syndrome and elbow pain(28, 29). 300

It is therefore important for clinicians to be aware of the potential presence of the AE 301

[Figure 2], as it may potentially be a contributing factor to patients presenting with elbow 302

pain or neuropathy. 303

304

In thinking about evolution, we tend to think of large branching moments in the 305

evolutionary tree that occurred hundreds of thousands or millions of years ago – Homo 306

heidelbergensis to Homo sapiens or Australopithecus anamensis and Australopithecus 307

afarensis. In between those large branching “moments,” however, evolution also occurs on 308

a more individual rather than a species level. Over time, these individual changes take hold 309

if they are beneficial to the endurance of the species, playing a part in adaptation, natural 310

14

selection and overall survival of the species. In this study we have examined four muscles 311

of the distal upper extremity that appear to be currently undergoing evolutionary 312

processes. It will be illuminative for researchers to continue to monitor the changes that 313

these muscles are undergoing over time to see whether they continue to change or 314

completely disappear in humans. Regarding current presentations of the FDS-V, it will also 315

be of interest, especially to the fields of medicine and anatomy, to determine the origin 316

(single origin vs. dual origin) and evolution of this muscle. Finally, with regard to the 317

anconeus, the role of the anconeus as a potential stabilizer of the humeroulnar joint during 318

infancy may be examined. Darwin wrote at length about various end-effects of natural 319

selection on evolution, and we believe as discussed herein using the examples of these four 320

muscles, that evolution is an ongoing process across a long-term continuum. 321

322

All authors declare that there is no conflict of interest. 323

15

References 324

[1] C. Darwin, The descent of man, and selection in relation to sex. , Princeton University Press, 325 Princeton, N.J. (1981; 1871). 326

[2] S. Standring, H. Ellis, J.C. Healy, et al., Gray's anatomy: the anatomical basis of clinical practice, 327 American Journal of Neuroradiology 26 (2005), p. 2703. 328

[3] N. Kobayashi, S. Saito, H. Wakisaka and S. Matsuda, Anomalous flexor of the little finger, Clin 329 Anat 16 (2003), pp. 40-43. 330

[4] W.A. Townley, M.C. Swan and R.L. Dunn, Congenital absence of flexor digitorum superficialis: 331 implications for assessment of little finger lacerations, The Journal of hand surgery, European 332 volume 35 (2010), pp. 417-418. 333

[5] M.M. Shrewsbury and K. Kuczynski, Flexor digitorum superficialis tendon in the fingers of the 334 human hand, The Hand 6 (1974), pp. 121-133. 335

[6] V. Abdala and R. Diogo, Comparative anatomy, homologies and evolution of the pectoral and 336 forelimb musculature of tetrapods with special attention to extant limbed amphibians and 337 reptiles, J Anat 217 (2010), pp. 536-573. 338

[7] R.W. Haines, The flexor muscles of the forearm and hand in lizards and mammals, J Anat 84 339 (1950), pp. 13-29. 340

[8] P.W. Brand and A. Hollister, Clinical mechanics of the hand, Mosby Year Book Burlington, MA 341 (1993). 342

[9] M.H. Gonzalez, J. Whittum, M. Kogan and N. Weinzweig, Variations of the flexor digitorum 343 superficialis tendon of the little finger, Journal of hand surgery (Edinburgh, Scotland) 22 (1997), 344 pp. 277-280. 345

[10] P. Bowman, L. Johnson, A. Chiapetta, A. Mitchell and E. Belusko, The clinical impact of the 346 presence or absence of the fifth finger flexor digitorum superficialis on grip strength, J Hand 347 Ther 16 (2003), pp. 245-248. 348

[11] M.E. Puhaindran, S.J. Sebastin, A.Y.T. Lim, W.X. Xu and Y.M. Chen, Absence of flexor digitorum 349 superficialis tendon in the little finger is not associated with decreased grip strength, Journal of 350 Hand Surgery (European Volume) 33 (2008), pp. 205-207. 351

[12] J.S. Tan, L. Oh and D.S. Louis, Variations of the flexor digitorum superficialis as determined by an 352 expanded clinical examination, The Journal of hand surgery 34 (2009), pp. 900-906. 353

[13] T.F. Gleason, W.M. Goldstein and R.D. Ray, The function of the anconeus muscle, Clinical 354 orthopaedics and related research 192 (1985), pp. 147-148. 355

[14] J.V. Basmajian and W.R. Griffin, Jr., Function of anconeus muscle. An electromyographic study, 356 The Journal of bone and joint surgery. American volume 54 (1972), pp. 1712-1714. 357

[15] K.N. An, F.C. Hui, B.F. Morrey, R.L. Linscheid and E.Y. Chao, Muscles across the elbow joint: a 358 biomechanical analysis, Journal of biomechanics 14 (1981), pp. 659-669. 359

[16] S.W. O Driscoll, J.B. Jupiter, G.J. King, R.N. Hotchkiss and B.F. Morrey, The unstable elbow, 360 Instructional course lectures-american academy of orthopaedic surgeons 50 (2001), pp. 89-104. 361

[17] F. Molinier, J.-M. Laffosse, O. Bouali, J.-L. Tricoire and J. Moscovici, The anconeus, an active 362 lateral ligament of the elbow: new anatomical arguments, Surgical and radiologic anatomy 33 363 (2011), pp. 617-621. 364

[18] F.A. Jenkins, The functional anatomy and evolution of the mammalian humero‐ulnar 365 articulation, American Journal of Anatomy 137 (1973), pp. 281-297. 366

[19] M.D. Cassell and R.A. Bergman, Palmaris longus muscle substituting for the ring finger slip of 367 flexor digitorum superficialis, Anatomischer Anzeiger 171 (1990), pp. 201-204. 368

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394

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Figure Legend 395

396

Figure 1: Palmaris longus tendon harvested during reconstructive surgery of an 8-year-old 397

female patient. Please note the short muscle belly and long tendinous portion of the graft. 398

Figure 2: Intraoperative image of cubital tunnel release showing the anconeus 399

epitrochlearis. Please note a small anconeus epitrochlearis (black arrow) muscle and its 400

relation to the ulnar nerve (asterisk). 401