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Functional morphology and flight kinematics of Artibeus jamaicensis (Chiroptera, Phyllostomidae)

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Title:
Functional morphology and flight kinematics of Artibeus jamaicensis (Chiroptera, Phyllostomidae)
Creator:
Hermanson, John W
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Language:
English
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x, 160 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Bats ( jstor )
Electromyography ( jstor )
Hand bones ( jstor )
Head ( jstor )
Humerus ( jstor )
Kidnapping ( jstor )
Mammals ( jstor )
Muscles ( jstor )
Scapula ( jstor )
Tendons ( jstor )
Artibeus jamaicensis ( lcsh )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1983.
Bibliography:
Includes bibliographical references (leaves 153-159).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by John W. Hermanson.

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University of Florida
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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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FUNCTIONAL MORPHOLOGY AND FLIGHT KINEMATICS OF
Artibeus jamaicensis (CHIROPTERA, PHYLLOSTOMIDAE)










By


JOHN W. HERMANSON


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA


1983




































Copyright 1983

by

John W. Hermanson
















ACKNOWLEDGEMENTS


Financial support for this study was provided in part by a

Grant-in-Aid of Research from the Society of Sigma Xi; from the

Department of Zoology, University of Florida; and from the Department

of Natural Sciences, Florida State Museum. I received support in the

form of a Museum Assistantship in the Department of Natural Sciences,

Florida State Museum, and a Graduate Teaching Assistantship in the

Department of Physiological Sciences, College of Veterinary Medicine.

Sr. Tomas Blohm kindly allowed me to study on his ranch in

Venezuela in 1981. Dr. John Robinson invited me to travel with him to

Venezuela, and then took care of the necessary procedures to obtain

permission for me to collect and transport live bats in that country.

I also acknowledge the Department of Biology, University of New Mexico,

for allowing me free access to the facilities of the Museum of

Southwestern Biology.

I thank my committee members for advice and attention at a time

when none of them needed the additional burden of a dissertation to

review. My committee included Drs. John Anderson, Donald Dewsbury,

David Webb, Ronald Wolff, and Charles Woods. Pamela Johnson typed and

then retyped the manuscript. I appreciate her patience in this task.












I offer thanks to the four men in biology who have most influenced

my path. Dr. Scott Altenbach, of the University of New Mexico, opened

his laboratory and his home to me during our joint venture. We will

have many enjoyable moments to reflect on during the years to come.

Dr. David Klingener, of the University of Massachusetts, taught a

fabulous course in comparative anatomy and opened my eyes to a career

in biology. Dr. Ted Goslow, of Northern Arizona University, provided

me with high goals in both my personal and academic life. Finally, my

advisor Dr. Charles Woods has provided friendship and a wealth of

experiences in natural history and anatomy on which to build a career.

I am fortunate to have studied with these four gentlemen.

I thank my peers for those good moments that make graduate

education unique. These moments include late nights out on the silent

Arizona desert, evenings of light snow or Great-horned owl hootings in

the Green Mountains, or simply industrious evenings in the labs at the

Museum. In particular, I will miss those evenings out on the back

porch with the folks in our household at Newnan's Lake. All of these

times are irrevocably tied together in my mind.

Last but not least I wish to acknowledge my family. My mother and

father endured my endeavor and provided encouragement and support. My

aunt, Catherine Leonard, has provided inspiration and the gift of

laughter. My "other aunt," Ruth Cushman, has been a friend

throughout. Finally, I remember my companion of many treks, Japhy.













TABLE OF CONTENTS


Page

ACKNOWLEDGMENTS . .. iii

LIST OF FIGURES . ... vi

LIST OF TABLES . ... .viii

ABSTRACT . ix

INTRODUCTION . ... 1

MATERIALS AND METHODS . .. 10

FLIGHT AND AERODYNAMICS

Results . ... 15
Discussion . .. 24

OSTEOLOGY . .. 31

MYOLOGY

Trapezius Group . ... .53
Costo-spino-scapular Group ... 62
Latissimus Group . .. 71
Deltoid Group . 79
Supraspinous Group . 84
Triceps Group . 88
Pectoralis Group . .. 91
Flexor Group of Arm .. 103
Antebrachial Extensor Group .. 105
Antebrachial Flexor Group ... 123

ELECTROMYOGRAPHY OF FLYING BATS .. 133

CONCLUSIONS ... .. 149

LITERATURE CITED . 153

BIOGRAPHICAL SKETCH . 160














LIST OF FIGURES
Page

1. The wingbeat cycle of Artibeus jamaicensis. 7

2. Wing and body movements during slow forward flight in 17
Artibeus jamaicensis.

3. Wing and body movements during slow forward flight in 19
Artibeus jamaicensis.

4. Wing and body movements relative to still air during 21
slow forward flight in Artibeus jamaicensis.

5. High- and low-aspect ratio wingshapes. 28

6. Lateral view of the thoracic and axillary skeleton. 33

7. Cranial view of the right pectoral girdle. 35

8. Dorsal view of the scapula. 37

9. Medial view of the left humerus. 41

10. Articular surfaces of the humerus. 43

11. Dorsal view of the radius and ulna. 48

12. Dorsal and ventral views of the carpus. 51

13. Dorsal view of the shoulder and arm of Artibeus jamaicensis. 56

14. Activity patterns of shoulder and arm muscles in Artibeus 58
during slow flight.

15. Lateral view of serratus ventralis musculature. 66

16. Ventral view of the shoulder and arm of Artibeus 94
jamaicensis.

17. Electromyographic data for six regions in the pectoralis 99
muscle during slow flight.

18. Lateral view of the muscles of the elbow region. 108










Page

19. Dorsal view of the muscles of the carpal region. 110

20. Medial view of the muscles of the elbow region. 125

21. Ventral view of the muscles of the carpal region. 127

22. Numbers of coactive muscles during the wingbeat cycle. 145














LIST OF TABLES
Page

1. Aerodynamic parameters of the wing in two fruit bats, 22
Artibeus jamaicensis.

2. Electromyographic data for activity patterns of the 59
shoulder musculature in Artibeus jamaicensis during
slow flight.

3. Electromyographic data for activity patterns in six 100
regions of the pectoralis muscle of Artibeus
jamaicensis during slow flight.


viii














Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy


FUNCTIONAL MORPHOLOGY AND FLIGHT KINEMATICS OF
Artibeus jamaicensis (CHIROPTERA, PHYLLOSTOMIDAE)


By

John W. Hermanson

April 1983


Chairman: Dr. Charles A. Woods
Major Department: Zoology

The functional morphology of the pectoral girdle and arm of

Artibeus jamaicensis (Chiroptera, Phyllostomidae) is interpreted on

the basis of gross dissection and electromyographical analysis (EMG).

Electromyographic data obtained during flight for 15 muscles elucidate

several temporal patterns of activity associated with the wingbeat

cycle that are not similar to the patterns of flexor, extensor, and

bifunctional muscles observed in terrestrial mammals. Abductor

muscles exhibit intense activity associated with the early upstroke

phase of the wingbeat cycle and include clavotrapezius, acromiotrapezius,

latissimus dorsi, teres major, acromiodeltoideus, spinodeltoideus, and

triceps brachii (long and lateral heads). All abductors except for the

triceps brachii exhibit a secondary period of low-amplitude activity

associated with the early downstroke. Adductor muscles exhibit primary

activity immediately prior to and during the early downstroke phase.

The adductors include serratus ventralis thoracis, pectoralis, and












clavodeltoideus. Bifunctional muscles exhibit a single period of

activity through most of the wingbeat cycle, or two separate periods

of high-amplitude activity during a wingbeat cycle. The bifunctional

muscles include spinotrapezius, supraspinatus, infraspinatus, and

subscapularis. Dissection of all other muscles of the shoulder and

antebrachium form the basis of interpreting musculoskeletal movements

during flight in Artibeus. The major muscles of support in Artibeus

include serratus ventralis thoracis, pectoralis, and the trapezius

group. These muscles support the trunk between the wings during

flight or terrestrial locomotion. Propulsion during flight differs

from that observed during stepping in terrestrial mammals. During

the wingbeat, pectoralis provides the major component of thrust,

both by adducting and pronating the wing. Although latissimus dorsi

is a major propulsive muscle during stepping in terrestrial mammals,

its major function in Artibeus is to abduct the wing and reposition

the wing prior to the beginning of a downstroke.
















INTRODUCTION


Bat flight is energetically expensive (Thomas and Suthers, 1972;

Carpenter, 1975; Thomas, 1975, 1981) and requires extensive alterations

of the generalized mammalian morphology (Miller, 1907; Vaughan,

1970a). In order to understand how bats fly, one must both study their

anatomy as well as the subtle interactions between the nervous and

muscular systems of several bats must be studied. This study provides

information about the neuromuscular and osteological mechanisms

associated with flight in a frugivorous bat, Artibeus jamaicensis

(Microciroptera, Phyllostomidae). The objectives of the project are to

describe in detail the musculoskeletal system of Artibeus, to

illustrate the kinematics of the normal wingbeat cycle, and to obtain

electromyograms during flight for the major muscles of the shoulder

region and arm.

Three factors make Artibeus jamaicensis desirable for

morphological analysis. First, Artibeus is a ubiquitous neotropical

bat whose basic biology has been well documented. The available

information regarding the foraging and flight habits of this bat

permits deduction of the demands placed on the bat. Second, Artibeus











jamaicensis is easily maintained in captivity during the duration of an

experimental period (Rasweiler, 1977). Third, Artibeus is placed in

the family Phyllostomidae (Jones and Carter, 1976), a family that has

not been employed in previous EMG studies.

The diet of Artibeus includes fruits, flower products, and leaves

(Goodwin and Greenhall, 1961; Villa-R., 1967; Heithaus et al., 1975;

Gardner, 1977; Bonaccorso, 1979). Insectivory is insignificant in this

species even though Tuttle (1968) observed Artibeus jamaicensis in

Mexico feeding on blackflies within a cave. The bats usually transport

relatively heavy fruits from their source to adjacent feeding roosts

except when too large to carry efficiently (Morrison, 1978a;

Bonaccorso, 1979). Thus the wings of Artibeus need to be capable of

producing lift at low speeds without a great deal of maneuverability.

The flight style of Artibeus differs from that of Desmodus,

Antrozous, and Myotis, three bats previously studied with

electromyography. The vampire, Desmodus, is characterized by its

direct, swift flight (Altenbach, 1979). Antrozous and Myotis are both

insectivores but each employs slightly different foraging behaviors.

Antrozous is a long-eared bat and either gleans insects from foliage or

preys upon large, slow-moving insect species at low altitudes. Myotis

myotis tends to also be an opportunistic gleaner, searching for and

obtaining insects from the surface of foliage or the ground (Findley,

1972). Desmodus, Antrozous, and Myotis all forage for extended











periods of time during the night. Antrozous was reported to engage in

continuous foraging on the wing for one to four hours during early

evening and later for several hours before dawn (O'Shea and Vaughan,

1977). Myotis lucifugus similarly feeds for several hours during

early evening, and again for several shorter periods before dawn

(Anthony and Kunz, 1977). I expect behavioral studies will show Myotis

myotis has a similar time-energy budget. Turner (1975) reported that

Desmodus spends one to three hours foraging per night and found that

the bats spent a mean of one hour per night searching for their prey.

Artibeus, in contrast, tends to spend relatively little time in flight

during the night (Morrison, 1978a; 1978b). Artibeus also is not very

maneuverable and is easily captured in nets. There are two brief

flights between the day roost and the food source: one at dusk to the

food and one shortly before sunrise back to the day roost (Morrison,

1978a). Morrison found that day roosts were never very distant from a

source of ripe fruit. At night, Artibeus selects fruit at the source

tree and then carries the fruit to an adjacent feeding roost, often in

another tree or in a cave (Dalquest, 1953; Morrison, 1978a; Bonaccorso,

1979). After chewing the fruit and swallowing juice and edible parts,

the bat ejects the remaining fruit pulp and seeds before returning to

the source tree for another fruit. These feeding flights are repeated

many times during the night.

Artibeus jamaicensis transports fruit on average 270 m from the

parental trees to feeding roosts (Janzen et al., 1976).











Morrison (1978a) observed that the distance between several Ficus

insipida and Ficus yaponensis, two species of figs important as food

trees, and the adjacent feeding roosts ranged from 25-400 m (mean 175

m). It is not clear why these bats carry fruits away from the parent

tree. Janzen et al. (1976) speculated that branch morphology and odor

in Andira inermis evolved to insure dispersal of the seeds away from

the parent tree. Seed mortality resulting from fungal infections and

weevil predation was most severe beneath the parent trees but declined

with distance away from the tree. They were unable to demonstrate a

clear cause and effect relationship, but did speculate that bats were

intolerant of the odor associated with many ripe Andira fruit. As an

alternative hypothesis, several authors suggested that Artibeus carried

fruit to a distinct feeding roost to decrease predation (Fenton and

Fleming, 1976; Morrison, 1978a; August, 1979). Artibeus avoid visually

orienting predators such as owls and opossums, by feeding only on dark

nights or during dark periods of the night (Morrison, 1978b). Specific

cases of predation by Didelphis marsupialis upon Artibeus were reported

in Venezuela (August, 1979). In any case, Artibeus make a number of

short flights during the night to and from a ripe fruit tree.

The neuromuscular control of flight can be correlated with similar

mechanisms in operation in the control of terrestrial locomotion.

Analysis of terrestrial involved resolution of the locomotor activity

of each limb into successive strides or step cycles, and quantification












of the interaction of these events between the limbs (gait analysis)

(Wetzel and Stuart, 1976). Philippson (1905) described four stages in

the step cycle of dogs, beginning with foot liftoff: F or flexion

phase is when most joint angles in the limbs decrease; El is the final

portion of the "swing" or non-support phase of the stride; E2 is the

first portion of the "stance" or support phase and includes some

passive joint flexion as a result of the limbs yielding under the

animal's weight; E3 is the final portion of the "stance" phase during

which all joints are actively being extended and maximum thrust is

being generated by the limb. Neurophysiologists studying vertebrate

locomotion have long focused on events surrounding the two reversal

points in the stride. For example, Sherrington (1910) made the

observation that a graded alternation of activities occurred in the

flexor and extensor muscles and that this alternation could be invoked

in a model of reflexive control of the step cycle (cf. Wetzel and

Stuart, 1976).

As an analogy, Hermanson and Altenbach (1981) expressed the

chiropteran wingbeat cycle as being a quantifiable and repeated unit of

normal bat flight. Individual wingbeats exhibit little variation among

conspecific individuals performing the same locomotory tasks under

similar conditions. The chiropteran downstroke, or adductor phase, is

similar in basic form and timing to the E2-E3 phases of the Philippson

step cycle and produces components of both lift and thrust (Norberg,

1976). The chiropteran upstroke, or abductor phase, parallels the F-El
































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phases of overground locomotion in its function as a recovery stroke.

The last portion of the upstroke often includes a thrust-producing

"flick phase" (Norberg, 1970, 1976). Except for the flick phase, it is

not practical to subdivide the upstroke or downstroke because of the

smooth arc followed by the wing during each phase (Figure 1).

Engberg and Lundberg (1969) demonstrated that cat limb muscles

exhibit characteristic activity profiles during overground locomotion:

extensor or flexor muscles produce a cumulative effect on the skeletal

system through overlap or coactivation, and muscles that span two

joints produce a complex function including both flexion and

extension. Engberg and Lundberg reported extensor activity prior to

foot touchdown during normal stepping. This important observation

indicated that muscle activity could be preprogrammed at the spinal or

supraspinal level, taking precedence over reflexive control of the step

cycle (Wetzel and Stuart, 1976). Subsequent studies demonstrated that

the EMG profiles for homologous mammals are conservative among mammals,

with subtle differences in timing related to differing functional

demands being placed upon the musculoskeletal system in various species

(English, 1978a, 1978b; Gambaryan, 1974; Rasmussen et al., 1978;

Tokuriki, 1973a, 1973b).

Hermanson and Altenbach (1981, 1983) assembled EMG data for muscle

activity during flight in an insectivorous bat, Antrozous pallidus

(Vespertilionidae). Of 17 muscles studied, 11 muscles demonstrated

patterns of either adductor or abductor activity. Only six muscles











exhibited a complex bifunctional pattern of activity. The adductor

muscles demonstrated an onset of activity on average 15-25 msec prior

to initiation of the downstroke. This pattern is similar to the El

extensor coactivation observed prior to foot touchdown in cats (Engberg

and Lundberg, 1969) and suggests that similar neural control mechanisms

are involved in regulating both the terrestrial step cycle and the

aerial wingbeat cycle. Despite gross anatomical modification during

the evolution of the chiropteral musculoskeletal system, the patterns

of limb movement and muscle activity remained relatively unchanged from

the basic mammalian condition.

This study presents the descriptive anatomy of the locomotor

system of A. jamaicensis, including the shoulder girdle, arm, and

forelimb portions of the wing. Descriptions are accompanied by

functional hypotheses for each muscle following a format employed in a

number of studies on the mammalian musculoskeletal system (Howell,

1926; Rinker, 1954; Klingener, 1964; Woods, 1972). Additionally, EMG

data are presented for 15 of the 17 muscles that were previously

analyzed in Antrozous pallidus (Hermanson and Altenbach, 1983). These

data are compared with simialr EMG observations upon terrestrial

mammals.
















MATERIALS AND METHODS


Ten skeletons and ten alcoholic specimens of Artibeus jamaicensis

were used for descriptive study. Dissection specimens were preserved

in a solution of 10 percent solution of formalin for 10-14 days and

were later stored in 70 percent ethanol or 40 percent isopropanol.

Complete dissections performed on the shoulder, arm, and antebrachial

regions form the basis of descriptions and illustrations presented in

this study and for the design of electromyography experiments.

Terminology used in my study conforms to recommendations by the

International Commission on Veterinary Anatomical Nomenclature (Nomina

Anatomica Veterinaria, 1975). Although this procedure initially

appears to pose difficulty in making comparisons with previous studies

on chiropteran morphology, relatively few new terms are introduced.

Strict adherence to the terminology of the Nomina Anatomica Veterinaria

(NAV) will provide a consistent vocabulary for effective communication

about the subject and will facilitate communication with researchers

who have not mastered the specialized jargon of bat anatomy.

Several terms are used throughout this text to indicate direction

along the body axis or direction of movement at a joint. The terms











cranial and caudal have replaced anterior and posterior, respectively,

in all cases where the latter terms have been used in the chiropteran

literature. Cranial refers to the cranial end of the animal, or

indicates that a structure passes from a point on the body along a line

towards the transverse plane of the head. Caudal indicates the

opposite direction to cranial: to or towards a transverse plane

containing the caudal vertebrae. Medial and lateral are used to

indicate position or movement towards or away from the medial plane of

the body. Flexion and extension refer to a decrease or increase,

respectively, in the angle between two bones. Adduction is used to

denote movements of the wings, humerus, or clavicle towards the medial

plane on the ventral surface of the body. Abduction of the wings,

humerus, or clavicle is a movement away from the median plane of the

ventral surface of the body. Rotation of a bone about its long axis is

described as supination, or outward rotation of the top or lateral

surface of the bone, and pronation, an inward rotation of the top or

lateral surface of the bone. To facilitate comparison with terrestrial

mammals, the arm and antebrachium are described in the position of

maximal adduction. Thus these bones have cranial, caudal, lateral, and

medial aspects. The carpus and digits are described as if they are

held horizontally at the side of the body. These elements have

cranial, caudal, dorsal, and ventral palmarr) aspects. The apparent

inconsistency in terms and orientation between the anatomical position

of the arm and carpus is in fact the traditional and accepted format

for descriptive purposes.











Skeletons were measured with dial calipers in order to describe

shape and size of individual bones. This information is useful in

estimating position or attachments of soft tissues with respect to the

skeleton.

Muscle origins and insertions were described relative to named

structures or surfaces of individual bones: the extent of muscle

attachment to long bones is expressed as a percentage or proportion of

the total length of the bones. Fiber orientation was expressed with

respect to the long axis of a muscle, along a line fit by eye between

the lines of attachment.

Synchronized film and EMG records were obtained with a system

initially described in Altenbach (1972, 1979) and modified by Hermanson

and Altenbach (1981). A Hycam high-speed cine camera (Red Lakes

Laboratories, Inc.) photographed flight sequences at film speeds of

300-400 frames per second. Myopotentials were recorded from a bipolar

fine copper wire electrode implanted in the muscle belly. Implantation

was achieved by a simple microsurgical procedure. An incision was

first made in the dorsal integument superficial to the lumbar region

and a silicone plug assembly was sutured to the muscle and ligaments of

the paravertebral region. Electrodes were passed subcutaneously to a

second incision at the level of the specific muscle being studied. A

shielded cable surrounding the electrode assembly served as a tissue

ground. With the aid of a microscope, the electrode tips were

inspected to insure than less than











0.5 mm of the tip was bared. The tips were then placed in the muscle

belly with a 27 guage hypodermic needle, and both incisions were

sutured shut with 6-0 Ethicon nylon suture. Methoxyfluorane was used

as a short term anesthetic agent with excellent results. Data were

recorded shortly after the bat exhibited normal locomotor behavior and

within three hours of surgery. Extended delays between the time of

implantation and recording sessions resulted in less than satisfactory

results, apparently as a result of toxic reactions between the copper

electrode material and adjacent tissues. Inspection of one

implantation that had been in place for 24 hours showed significant

tissue necrosis in the muscle for a 2 mm diameter around the electrode

tip. Only one muscle was studied per experiment. Each implant was

visually inspected at the conclusion of each recording session to

verify that the electrode remained in place. Each animal was allowed

to recover for at least one week before being used in further

experiments.

Normal flight was studied with single frame analysis of films of

normal horizontal flight sequences. Single film frame images were

projected with an L and W motion analyzer (Model 224A) and were traced

to ascertain wingtip motion and joint position in the wing. Vertical

displacement of the wingtips was used to determine the turnover points

of adduction and abduction. The beginning of a downstroke was treated

as the beginning of a wingbeat cycle and was assigned a numerical value

of 0.000 (Figure 1, frame B). A complete wing cycle included values

between 0.000 and 1.00, with 1.00 representing the final frame of an











upstroke (Figure 1, frame M). The transition between downstroke and

upstroke (Figure 1, frame H) and periods of muscle activity are

expressed as a mean percentage of a complete cycle.

Estimation of aerodynamic parameters was accomplished by measuring

the dimensions of the wing and body of two fresh specimens of Artibeus

jamaicensis. The mass of each bat was obtained to the nearest 0.5 g.

Wing area was estimated by drawing the outline of the body and fully

outstretched wing of each bat on paper. The cut-out outline of the

airfoil was then weighed to the nearest 0.0005 g and then compared to

the mass of a .100 m2 square of the same paper to calculate actual

wing area. Wingspan was measured along a line between the two

wingtips. Aspect ratio is the ratio of length to width of the

airfoil. The equation for aspect ratio is b2/A (Norberg, 1981).

Wingloading is the ratio of body weight to wing area, W/A (Norberg,

1981).
















FLIGHT AND AERODYNAMICS


Results

The temporal relationship between the downstroke and upstroke is

critical in the interpretation of EMG data. In part, this relationship

reflects whether or not the activity of a given muscle is coincident

with or occurs before or after a particular event, such as the

beginning of a downstroke. During all filming sessions, flight was

observed to detect any grossly abnormal flight behaviors. All flight

sequences exhibiting turning behavior or altitude loss were eliminated

from consideration. The wingbeat cycle of Artibeus appeared to be

conservative during slow horizontal flight, based upon unaided visual

observation as well as initial study of the film records. However,

analysis of the film revealed slight differences in the flight of

restrained and unrestrained animals.

During unrestrained flight in two bats, the downstroke encompassed

55.1 percent of the wingbeat. In contrast, bats that carried an

electrode and plug assembly exhibited a mean downstroke of 49.44

percent. During unrestrained locomotion, the bats averaged 10.08

wingbeats per second. Records analyzed from four EMG experiments

(clavotrapezius, spinotrapezius, clavodeltoideus, and infraspinatus)

demonstrated a mean wingbeat frequency of 10.45 wingbeats per second.


































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Figure 4.--Wing and body movements plotted relative to still
air during slow forward flight in Artibeus jamaicensis. The
bat flew from left to right at a velocity of 2.15 m/sec.
(A) The movements of the wingtip are plotted relative to the
body. (B) The movement of the wrist is plotted relative to
the body.













































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Table 1. Aerodynamic parameters of the wing in two fruit bats,
Artibeus jamaicensis. All measurements are recorded in
meter-kilogram-second units.


mass

weight

wing span (b)

wing area (A)

aspect ratio (b2/A)

wing loading (W/A)


0.04 kg

0.40 N

0.39 m

0.02 m2

6.79

16.74 N/m2


0.03 kg

0.32 N

0.32 m

0.02 m2

6.55

15.47 N/m2











Analysis of one sequence of unrestrained flight is provided to

illustrate airspeed and movement of several parts of the body and wing

(Figures 2-4). Plots of head movement exhibited forward velocity

ranging from 2.19-2.85 m/sec. At the beginning of the downstroke, the

wingtip was positioned above and caudal to the center of the body. The

wing was adducted, but moved with an initial caudal sweep of the

wingtip relative to the center of the body. After approximately 15

msec, the adduction of the wingtip exhibited an increasing cranial

component of movement. The wrist moved in synchrony with the wingtip

but travelled through a smaller excursion. The wingtip continued

moving ventrally and cranially through 62.5 percent of the wingbeat

(Figure 2, Symbols 2-12). The wingtip and wrist synchronously showed

abduction at the beginning of the upstroke. There was a cranial

movement of both elements for approximately 5 msec. Subsequently,

abduction of the wingtip and the wrist included a caudal component of

movement for about 26 msec until the beginning of the flick phase.

During the flick phase (Figure 2, Symbols 18-21) the outer portion of

the chiropatagium was rapidly pronated. Cranial movement of the

wingtip occurred during the flick phase, which lasted approximately 5

msec, or until the beginning of the downstroke. During the entire

upstroke, movement of the wrist was in a dorsal and caudal direction.

Thus, during the wingbeat cycle the wrist followed an elongated

elliptical path. The path of the wingtip was best characterized as a

tight and complex figure eight when viewed laterally.











Several measures are used in the description and comparison of the

aerodynamic properties of Artibeus jamaicensis. Based on two specimens

obtained in Haiti, the following means were obtained for these

parameters: mean wing area, inclusive of the body, 0.02 m2;

wingspan, 0.38 m; aspect ratio, 6.67; wingloading, 16.11 N/m2.



Discussion

The wing and flight characteristics of Artibeus jamaicensis are

generalized in comparison to many other bats. Analysis of the flight

kinematics and wingshape demonstrates that Artibeus is capable of slow

flight in the forest canopy but not the faster, more maneuverable

flight associated with the pursuit of insects among dense vegetation or

in open airspace.

The data obtained during EMG experiments on Artibeus represent a

departure from the mean downstroke times observed during the

unrestrained flight sequences. These EMG flight data are comparable to

downstroke phase relationships reported during EMG studies in Antrozous

where a range of 39.7 to 52.3 percent was reported for the downstroke

period (Hermanson and Altenbach, 1983). The downstroke of flying

animals lasts longer than the accompanying upstroke (Aymar, 1935;

Brown, 1963; Norberg, 1976; Brandon, 1979). Flying birds or bats can

be seen to oscillate in a vertical plane during flight, losing a slight

amount of altitude during each upstroke phase, and regaining altitude

during the subsequent downstroke (Eisentraut, 1936). The upstroke does











not provide sufficient lift to counteract the effect of gravity except

in certain species that employ a hovering mode of flight (Norberg,

1975).

The flight speeds observed in the present study are similar to

those observed in Plecotus auritus by Norberg (1976). In her study,

the bats were filmed while flying unrestrained within a room and flight

speeds ranged between 2 a 3 m/sec. Higher velocities were exhibited by

Noctilio albiventris (3.51-10.38 m/sec) and Tadarida brasiliensis

(4.33-9.40 m/sec) while flying in a wind tunnel (Brandon, 1979).

Leptonycteris sanborni, a facultative hovering bat, flew between 2.0

and 6.0 m/sec in a wind tunnel study. The higher speeds reported in

the wind tunnel studies were not duplicated in the present study.

Predictable flight, suitable for EMG analysis, was facilitated by

filming the bats shortly after takeoff and before they reached maximum

velocities. Also, it is not known if Artibeus is capable of flying up

to 4.47 m/sec as was generalized for many bats (Hayward and Davis,

1964).

The path of the wingtip during the wingbeat cycle changes with

increasing airspeed. During slow flight in Plecotus auritus, the

wingtip path was directed upwards and caudally relative to still air

during the upstroke (Norberg, 1976). At higher velocities (2-2.5

m/sec), the upstroke of Plecotus was directed almost vertically, and at

higher velocities (greater than 3.0 m/sec) exhibited a cranial and

dorsal movement relative to still air. The same tendency was present

in Artibeus flying between 2-2.85 m/sec. The wing was directed











caudally during the upstroke at low speeds (Figure 2) and more

vertically during the upstroke at higher velocities (Figure 3). If

wing movements are different at several airspeeds, these relationships

must be considered when evaluating EMG recordings. The difference in

wing movements may be correlated with differences in muscular activity.

Flying animals have been classified according to several

descriptors of wingshape including the aspect ratio (Saville, 1962).

Vaughan (1959) discussed the relationship between aspect ratio and

foraging habits. Struhsaker (1961) was the first author to describe

the aspect ratios of a diversity of chiropteran wings. Aspect ratios

were also provided for bats by Farney and Fleharty (1969), Findley et

al. (1972), Lawlor (1973), Smith and Starrett (1979), and Norberg

(1981). My calculation of an aspect ratio for Artibeus jamaicensis,

6.67, is slightly greater than but comparable to the value of 6.36

obtained by Norberg (1981). In general, bats with low aspect ratios

are common in heavily forested habitat, whereas high aspect ratio bats

are commonly associated with flight in open, uncluttered airspace

(Findley et al., 1972). For example, Vaughan (1959) compared the

high-altitude flight of Eumops perotis with that of swifts. These bats

fed high over the forest canopy at high airspeeds. Vaughan reported

that the aspect ratio of Eumops was 11.9, the highest value among the

three species that he studied. Intermediate aspect ratio wings are not

well correlated with any particular behavioral repertoire or foraging

environment (Norberg, 1981). Wing shape in Artibeus probably evolved


































Figure 5.--High- and low-aspect ratio wingshapes. Tadarida
brasiliensis (Molossidae) exemplifies the long and narrow
outline of a high aspect ratio wing. In contrast, the wing
of Artibeus jamaicensis exhibits a lower aspect ratio,
characterized by a broad and stubby appearance.













Tadarida









Artibeus











in response to the needs of the bat in carrying large food items and

transporting infants. Artibeus females have been observed carrying

young weighing 10.0 g (personal observation). Low aspect ratio wings

promote increased lift at lower flight speeds (Findley et al., 1972),

permitting more careful scrutiny of potential food or of potential

feeding sites. Maneuverability does not appear to be characteristic of

the flight pattern of Artibeus in comparison with some more agile

bats. This was supported by repeated netting success while trying to

collect bats in mist nets (personal observation). In contrast, other

phyllostomids such as Micronycteris have low aspect ratio wings (Smith

and Starrett, 1979) and were frequently observed eluding my nets during

field work in Venezuela.

Wing loading is an estimate of the relationship between the weight

of a bat and the lift-producing potential of a wing. To fly slowly,

which would be adaptive to an Artibeus investigating a fig tree as a

potential food source, two strategies might be employed: the species

might exhibit low body weight or a greater airfoil surface area

(Findley et al., 1972). My data for wing loading in Artibeus were

slightly less than the 16.65 N/m2 reported by Norberg and greater

than the 15.94 N/m2 reported by Smith and Starrett (1979). Norberg

found that the wingloading of frugivorous microbats (this included most

phyllostomid fruit eaters) was intermediate between the insectivorous

Molossidae and Vespertilionidae. The molossids of equivalent weight

have high aspect ratios and must maintain high airspeeds to avoid a











stall (Vaughan, 1966; Norberg, 1981). In contrast, the vespertilionids

generally have a large wing area for a given body size (Norberg, 1981)

and fly at relatively lower airspeeds than similar-sized molossids

(Hayward and Davis, 1964; Brandon, 1979). Thus, the wing of Artibeus

represents a compromise between the two strategies suggested by Findley

et al. (1972). The bats have large body weights but have not

substantially increased wing surface area like vespertilionids have.

This combination leads to reduced maneuverability, different from the

butterfly-like flight observed in some vespertilionids. Also, I have

never observed Artibeus hovering. The bats tend to fly directly along

aerial pathways to and from a feeding tree, and fly past a tree several

times when investigating or searching for food.

In conclusion, the wing of Artibeus represents a generalized

condition relative to certain other bats. I suggest that this reflects

the heterogeneous nature of the airspace through which the bat must

travel every night. A high aspect ratio wing is not suited to travel

through the cluttered environment of most neotropical forests. Also,

at slow airspeeds, the larger lift production that accrues from a low

aspect ratio wing permits the bat to forage upon and to transport large

energy-rich foods. A secondary consequence of the large body mass and

low aspect ratio of Artibeus may relate to post-natal development and

maternal care of the young bats.
















OSTEOLOGY


Results



Clavicle

The clavicle in five specimens had a mean length of 16.10 mm and a

mean diameter at the midshaft of 1.15 mm. The bone articulates

proximally with the manubrium by a synovial joint. Distally, the

clavicle articulates with the scapula both by a synovial joint at the

base of the coracoid process and by a ligamentous connection with the

tip of the acromian process. Viewed cranially, the bone is straight,

with slight lateral curvature near its distal end. Viewed laterally,

the bone has a slight caudal curvature at the distal end. The distal

one-third of the cranial border of the bone is expanded for insertion

of part of the subclavius muscle. In lateral view the distal end of

the bone lies cranial relative to the proximal end.

Scapula

The scapula is a thin, relatively flat bone positioned over the

dorsal surface of the cranial part of the ribcage. It articulates

proximally with the clavicle as described above. The scapula




























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_


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Figure 7.--Cranial view of the right pectoral girdle.
UF 16265.

















acromion process












4
\
\
\










glenoid fossa

coracoid
coracoid process


craniomedial flange


) -.- caudal angle


.-- subscapular fossa





----------- clavicle


manubrium ---


5 MM















































C-s


r-4










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articulates with the humerus at the glenoid fossa, which forms the

shoulder joint. The dorsal surface of the scapula is divided into two

major subdivisions by the scapular spine. Cranially, the supraspinous

fossa forms a small flat shelf immediately cranial to the scapular

spine. Caudal to the scapular spine is the infraspinous fossa which

is, in cranial to caudal sequence, divided into three subdivisions:

craniomedial facet, intermediate facet, and caudolateral facet. The

ventral surface of the scapula is the subscapular fossa, composed of

two concave facets. The cranial subscapular facet corresponds with the

contours of the combined supraspinous fossa and craniomedial facet of

the dorsum. The caudal facet of the subscapular fossa corresponds with

the contours of the combined intermediate and caudolateral facets of

the dorsum. There are two borders on the scapula. Medially, the

vertebral border lies in a sagittal plane extending from the caudal

angle to the scapular spine. Cranial to the scapular spine, the

vertebral border curves laterally and becomes continuous with the

craniomedial flange of the scapula. On the lateral aspect of the

scapula, the axillary border is convex, coursing cranially from the

caudal angle to the glenoid fossa. The infraglenoid tubercle is a

small projection from the axillary border 1-5 mm caudal to the glenoid

fossa. The supraglenoid fossa is slightly concave and lies between the

glenoid fossa and the dorsal base of the coracoid process. This

surface abuts with the greater tubercle of the humerus when the wing is

fully abducted.











Several processes are evident on the scapula of Artibeus. The

cartilaginous expansion is triangular and projects from the caudal

angle and curves ventrally. The acromion process projects cranially

and dorsally from the scaular spine, and curves cranially. A 2 mm long

flared part of the acromion process lies dorsal to the clavoscapular

articulation. The coracoid process extends cranially and ventrally

from the supraglenoid region of the scapula and curves laterally to

terminate 2 mm ventral to the glenoid fossa.

Humerus

The humerus averaged 31.90 mm in length (measured between the

articular surfaces) and 1.98 mm in diameter (along the craniocaudal

axis) at the midshaft. In cranial view the distal end of the shaft

curves slightly lateral. The shaft is unmarked. Both ends of the

humerus are modified to form articular surfaces and to accommodate

muscle attachments.

The shoulder joint is formed in part by the head of the humerus.

The head is a convex elliptical surface that articulates proximally

with the glenoid fossa. The greater tubercle lies lateral to the head

and projects 1 mm proximally. The greater tubercle provides attachment

for the suprascapular muscles and abuts with the supraglenoid fossa of

the scapular when the humerus is fully abducted. This lock occurs when

the humerus is abducted about 32 degrees dorsal to the horizontal plane

of the scapula. The lesser tuberosity also extends slightly proximal

to and lies on the medial aspect of the head. Several ridges extend


































Figure 9.--Medial view of the left humerus. UF 16265.



















greater tubercle


lesser tubercle

head ---------


lateral ridge ------'






















medial epicondyle -----------


apex of pectoral crest


pectoral crest









shaft


5 MM


































Figure 10.--Articular surfaces of the humerus. (A) Caudal
aspect of the proximal left humerus. (B) Cranial aspect of
the distal left humerus. UF 16265.


















greater tubercle


lateral ridge






A


I
I
I
I


medial process


,- lesser tubercle




.-- ------- medial ridge




















Sepicondylar ridge


lateral epicondyle




..

capitulum

troch
trochlea


head











distally from the lesser and greater tuberosities. From the lesser

tuberosity, the medial ridge extends 6 mm along the proximal medial

border of the humerus and serves as the point of attachment of the

teres major and latissimus dorsi muscles. A second indistinct ridge

crosses the proximal surface of the humerus and courses along the base

of the ventral surface of the pectoral ridge. This ridge is the site

of insertion of the pectoralis abdominalis. The lateral ridge is a

small indistinct ridge on the lateral surface of the humerus courses 7

mm distal to the greater tuberosity. This forms the dorsolateral

border of the origin of triceps brachii, lateral head. The most

prominent ridge from the greater tuberosity is continuous distally with

the pectoral ridge, a large flat projection on the lateral aspect of

the humerus. The pectoral ridge extends 8 mm along the lateral edge of

the humerus. The widest portion of the pectoral ridge, 2-4 mm distal

to the head, is termed the apex. The pectoral ridge provides insertion

sites for the pectoralis muscles medially, and the deltoid muscles

laterally.

At the elbow the radius articulates with the cranial surface of

the humerus. The ulna articulates along the distal and caudal surface

of the humerus. The capitulum and trochlea are fused cranially to

present a synovial surface for the humeroradial articulation. The

trochlea extends around the distal and caudal surface of the humerus to

provide an articular surface for the ulna. Medial to the trochlea and

composing one-third of the distal end of the bones is a medial











process. This region provides an attachment surface for several flexor

muscles of the antebrachium. On the lateral end of the capitulum is

the spool-shaped lateral epicondyle. Extending from the lateral

epicondyle proximally onto the shaft of the humerus is the short

epicondyle ridge. The lateral epicondyle and epicondylar ridge provide

an area of attachment for several antebrachial extensor muscles. The

elbow articulation is not a pure hinge joint. Slight rotation of the

radius relative to the capitulum is evident in fresh specimens of

Artibeus.

Radius

The radius is slightly curved along its length. In five

specimens, the mean length is 53 mm. At the midshaft, it is thickest

in the dorsoventral axis (1.8 mm) to resist bending forces associated

with the wingbeat cycle. The proximal articular region is thickened to

provide a surface for articulation with the humerus and to provide a

strong site for attachment of the biceps brachii. The distal articular

region is wide and flat. This provides a structural mechanism for

separation of the tendons of insertion of the antebrachial muscles and

increases the mechanical advantage of several of these muscles as they

cross the carpus.

The proximal articular surfaces is oriented cranially and includes

a large concave fossa for articulation with the capitulum. Laterally a

smaller elliptical concavity articulates with the trochlea. A ridge on

the caudal aspect of the radius courses 5 mm distal from the elbow











joint. A ridge on the medial aspect of the proximal radius courses 3

mm distal to the elbow joint and provides the site for insertion of the

biceps brachii.

The styloid process and pseudostyloid process are small bumps on

the distal articular end of the radius but are not greatly developed in

bats. The pseudostyloid process is on the medial side of the distal

articular surface, analogous in position to the styloid process of the

ulna in other mammals.

The axis of the distal articular surface is laterally rotated

about 20 degrees from the orientation of the proximal articular

surface. The distal articular surface presents a concave surface for

articulation with the proximal carpal bones. When the wing is

laterally outstretched, the radiocarpal joint permits flexion and

extension in a mediolateral plane and no movement in the dorsoventral

plane. This restriction braces the joint against displacement caused

by the force of the airstream.

Ulna

The ulna is reduced in bats to a length approximately one-half the

length of the adjacent radius. The proximal articular surface

comprises the cranial surface of the olecranon process and articulates

with the caudal extension of the trochlea of the humerus. An

interosseous membrane is found between the radius and ulna. The ulna

tapers towards its distal articulation with the radius and presents no


processes.



































Figure 11.--Dorsal view of the radius and ulna.
























radius
\


Sulna


olcranon


process


10 mm











Manus

The diminutive size of and the tightly bound joint cavities

between the bones of the carpus make it difficult to adequately

describe the bones of this region. A perspective on the carpal anatomy

can be obtained by studying Figure 12. The carpus of Artibeus includes

eight bones as is typical of many mammals.

The proximal row of carpal bones includes the scaphoid, lunar, and

triquetrum. The largest carpal bone, the lunar, articulates proximally

with the radius and distally with the trapezoid, trapezium, and

capitate. On the lateral aspect of the distal articular surface of the

radius, the triquetrum (-cuneiform of Vaughan, 1959) is bound to the

radius and forms the caudoventral border of the canal traversed by the

antebrachial extensor muscles. The scaphoid is a small bone located

craniodorsally in the carpus along the cranial surface of the lunar.

The scaphoid, unlike the lunar and triquetrum, is not bound to the

distal surface of the radius.

The distal row of carpal elements includes the trapezium,

trapezoid, capitate, hamate, and pisiform. The trapezium articulates

with the proximal bases of metacarpals I and II, and with the

craniodistal surface of the lunar. The trapezoid is compressed

craniocaudally and articulates with the proximal base of metacarpal

II. The capitate (=magnum of Vaughan, 1959) is a large bone that has

an hourglass shape when viewed dorsally. It articulates with the

proximal bases of metacarpals II and III. Lateral to the capitate, the




































Figure 12.--Dorsal and ventral views of the carpus.












styloid process
lunar

triquetrum

radius









hamat














m


trape


mc II





mc III


scaphoid


capitate


mc I


mc II


e


mc III


mc IV


mc V


trapezium


Ic I


mc IV


', 'triquetrum
'pisiform

hamate
capitate


mc V


5 mm











hamate uniformom of Vaughan, 1959) articulates distally with the bases

of metacarpals IV and V, and proximally with the lunar and triquetrum.

The pisiform is a small rod-shaped bone on the palmar surface of the

carpus and is firmly bound to the ventral surface of the capitate and

hamate.

The distal part of the wing is supported by digits II through V.

The metacarpals diverge within the wing membrane and are the largest

elements in each digit. Movement at the carpometacarpal joints of

digits II through V is restricted to flexion and extension. Flexion of

the carpometacarpal joints brings the digits parallel to and into

juxtaposition with the radius. The metacarpophalangeal and

interphalangeal joints permit only palmar flexion or dorsiflexion.

These movements adjust the camber of the wing during the wingbeat

cycle. Digit II has one phalanx. Digits III through V have three

phalanges. Digit I, the pollex, is unique because of the range of

motion possible at the carpometacarpal joint. Besides flexion and

extension, the pollical carpometacarpal joint permits the extensive

rotation necessary for use of the pollex in several functions: the

pollex is a prehensile organ used during landing maneuvers or while

manipulating food. The second phalanx of the pollex bears a claw. The

proximal and second phalanges of the pollex are both free of the

leading edge of the dactylopatagium minus and the propatagium.
















MYOLOGY


Trapezius Group

Clavotrapezius and Acromiotrapezius

Form. The clavotrapezius originates by a fibrous attachment on

the dorsal surface of the seventh cervical vertebrae. Fibers of

acromiotrapezius arise from the dorsal surfaces of thoracic vertebrae

one through six. There is no apparent separation between adjacent

fibers of the clavotrapezius and acromiotrapezius. Insertion of the

clavotrapezius is by fibers on the distal 2.5 mm of the craniomedial

surface of the clavicle. The acromiotrapezius inserts separately from

the clavotrapezius: insertion is by fibers convergent upon the

anterior and dorsal surface of the acromion process and on a muscular

raphe caudal to the acromion process. Innervation of both muscles is

by cranial nerve XI (spinal accessory nerve).

Comparative aspects. The clavotrapezius and acromiotrapezius

muscles have been treated as two separate muscles in the Phyllostomidae

(Macalister, 1872; Walton, 1967), and also in the specialized vampire

bats (Altenbach, 1979). In contrast, the condition observed in many

other bats, including phyllostomatids, demonstrated fusion of these two

bellies (Strickler, 1978; Vaughan, 1959, 1970a). In the three species

he studied, Vaughan (1959) noted the fusion of the two bellies only in











Macrotus (Phyllostomidae). Macalister (1872; p. 138) most likely made

reference to the clavotrapezius of Artibeus sp. when he described "a

semi-detached upper slip passed from the two lowermost cervical spines

to the outer fifth of the clavicle." The two muscles are

distinguishable in Artibeus only by a fascial division coursing from

the C7-T1 articulation to the distal end of the clavicle. Despite

their separate insertion, I consider the two muscles to be fused and to

share a similar function. The clavotrapezius and acromiotrapezius form

a flat sheet of parallel fibers, thickest at the cranial end

(clavotrapezius portion).

Functional aspects. The clavotrapezius and acromiotrapezius

exhibit a similar pattern of EMG activity during the wingbeat cycle.

Clavotrapezius has a major period of EMG activity during the transition

between the late downstroke and early upstroke. A second period of

activity occurs during the middle to late upstroke phase, but the

myopotentials have less than one-half of the amplitude observed during

the earlier EMG burst. Acromiotrapezius also has two periods of EMG

activity, but both exhibit similar amplitude and frequency. The first

period of activity in acromiotrapezius precedes clavotrapezius activity

by approximately 10 m/sec. Both muscles are anatomically positioned to

effect the upstroke by drawing the scapula medially (Vaughan, 1959;

Strickler, 1978; Altenbach, 1979). The primary activity of each muscle

contributes to this upstroke function. The secondary burst of activity

possibly stabilizes the scapula in response to the adductor activity of





























U)
a4-1


u1
4 -4


0 01
m










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0) 414 M
00 m 0
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r-1 (
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0 0


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(DO0 0 0 0
0) 0 'O 1 OC


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C- O I C
ca"0 *H *c

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, co w > H w
o c 0 c 0 r- 4 J.
0 4-1 U 0
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3 u 1o Q) c au

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w -0 0 U 0 0
0 *0 ol c
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t 0 > *H i
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1-1 0B 0i -i!< J






58












-Y

0o

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1o 0


SCO


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S, | 0 C

























C M 0 .I 7 c
a V- co C -

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< S0 V5















Table 2.--Electromyographic data for activity patterns of the shoulder
musculature in Artibeus jamaicensis during slow flight. N= number of
wingbeats analyzed per muscle. Mean duration of the downstroke is
expressed as a percentage of the wingbeat cycle duration. Mean times
of muscle activity onset and termination are recorded in relation to
the wingbeat cycle from 0.00 to 1.00, and one standard deviation is
recorded.



Downstroke Onset Termination
Muscle N Duration Time (S.D.) Time (S.D.)


ABDUCTORS
Clavotrapezius


Acromiotrapezius


Latissimus dorsi


Teres major

Acromiodeltoideus
cranial


caudal


Spinodeltoideus


Triceps brachii
lateral


12 0.515


9 0.477


13 0.428


3 0.412


10 0.536


10 0.532


11 0.481



3 0.594


0.332 (.099)
0.894 (.030)

0.236 (.033)
0.794 (.080)

0.329 (.028)
0.869 (.035)

0.259 (.022)
0.922 (.046)

0.355 (.056)
0.799 (.041)

0.315 (.053)
0.894 (.051)

0.309 (.030)
0.001 (.115)


0.564 (.017)


15 0.513 0.429 (.069)


0.631 (.045)
0.092 (.047)

0.572 (.104)
0.903 (.104)

0.644 (.025)
0.098 (.025)

0.519 (.016)
0.077 (.045)

0.628 (.040)
0.081 (.055)

0.556 (.062)
0.103 (.065)

0.608 (.030)
0.114 (.080)


0.741 (.053)

0.714 (.074)


long














Table 2.--continued


Downstroke Onset Termination
Muscle N Duration Time (S.D.) Time (S.D.)


ADDUCTORS
Clavodeltoideus 19 0.544 0.853 (.059) 0.135 (.027)
0.481 (.038) 0.617 (.039)

Serratus ventralis 11 0.484 0.823 (.036) 0.091 (.058)
0.515 (.058) 0.599 (.109)

Pectoralis 7 0.526 0.781 (.027) 0.094 (.086)

BIFUNCTIONAL
Spinotrapezius 7 0.452 0.254 (.031) 0.672 (.032)
0.784 (.020) 0.123 (.047)

Supraspinatus 6 0.518 0.706 (.263) 0.142 (.084)

Infraspinatus 7 0.449 0.346 (.039) 0.607 (.046)
0.942 (.059) 0.120 (.039)

Subscapularis 10 0.450 0.420 (.060) 0.174 (.033)













the early downstroke, or causes active lengthening of the pectoralis

fibers during their early period of activity. Similar passive

lengthening, or "eccentric contractions," increased force production of

several locomotory muscles in other animals (Cavagna et al., 1977;

Heglund et al., 1979; Goslow et al., 1981). The biphasic activity

observed in Artibeus contrasts with the single EMG burst observed in

Antrozous (Hermanson and Altenbach, 1983). The heavier wingloading of

Artibeus may necessitate the extra muscular activity. In conclusion,

both clavotrapezius and acromiotrapezius show abductor activity, but at

slightly different times in the wingbeat cycle. The secondary activity

in each muscle stabilizes the scapula against the force of the

adductors.

Spinotrapezius

Form. The spinotrapezius originated by fibers on the dorsal

midline over lumbar vertebrae two through three. It inserts by an

aponeurosis on the vertebral border of the scapula, extending from the

caudal angle to the point 2 mm anterior to the scapular spine. The

spinotrapezius is a thin, parallel-fibered muscle. Based upon its

anatomy, the muscle tips the vertebral border of the scapula ventrally,

and retracts the scapula (Strickler, 1978). Innervation is by the

spinal accessory nerve.

Functional aspects. Spinotrapezius exhibits biphasic EMG

activity. Both periods of activity have approximately equal amplitude

and frequency characteristics. One period commences at 0.254 and











terminates at 0.672: This contributes to the overall abduction of the

wing along with the activity of clavotrapezius and acromiotrapezius. A

second period of activity commences at 0.784 and terminates at 0.123 of

the following wingbeat. Spinotrapezius is quiescent only during about

24 percent of the wingbeat, an observation similar to EMG data from

Desmodus (Altenbach, 1979), but in contrast to the shorter, single

activity period observed in Antrozous (Hermanson and Altenbach, 1983).

The extent of EMG activity throughout the wingbeat suggests a multiple

function for spinotrapezius as proposed by Vaughan (1959):

stabilization of the scapula and abduction of the wing and scapula are

likely functions.



Costo-spino-scapular Group

Serratus Ventralis Cervicis

The serratus ventralis cervicis was previously called the elevator

scapulae in the chiropteran anatomy literature.

Form. The serratus ventralis cervicis originates from fibrous

attachments along the dorsolateral aspect of the transverse processes

of cervical vertebrae three through six. The muscle has a fleshy

insertion on the ventral surface of the craniomedial flange of the

scapula, and along the vertebral border of the scapula craniad of the

scapular spine. The muscle is composed of two parallel-fibered slips:

the cranial slip originated from cervical vertebrae three and four, and

the caudal slip from five and six. Innervation is by a branch of the

dorsal scapular nerve, and by cervical nerve five.












Comparative aspects. In many mammals, the serratus ventralis is

composed of a continuous sheet of muscle, including fibers that

originate from the cervical vertebrae serratuss ventralis cervicis) and

from the lateral surface of the ribcage serratuss ventralis thoracis).

The terminology and comparative anatomy are discussed with the serratus

ventralis thoracis.

Strickler (1978) noted that although the cranial origin normally

extends to cervical vertebra five to seven the origin of serratus

ventralis cervicis varies in bats. Phyllostomoids were characterized

by a relatively large insertion, usually extending along the vertebral

border of the scapula caudal to the scapular spine. Walton (1967)

observed a similar insertion in Artibeus lituratus extending along the

"coracoid border" and vertebral border of the scapula to a point

immediately posterior to the spine.

Walton (1967) described the serratus ventralis cervicis of A.

literatus to be composed of three large slips. Macalister observed

only two slips in Artibeus sp. (Macalister, 1872). I concur with

Macalister.

Functional aspects. Strickler (1978) concluded that a correlation

existed between degree of humeroscapular "locking" observed in some

bats and the size of the serratus ventralis cervicis muscle. He

observed that this muscle was relatively enlarged while the rhomboideus

was relatively small in vespertilionids and phyllostomatids. In

contrast, the serratus ventralis cervicis was small and the











rhomboideus large in emballonurids and pteropodids. These two groups

possess poorly developed humeroscapular locking mechanisms (Vaughan,

1970a, 1970b; Strickler, 1978). Strickler suggested that the major

role of serratus ventralis cervicis is to stabilize the medial border

of the scapula against rotation about its longitudinal axis. No EMG

data are available to test this hypothesis.

Serratus Ventralis Thoracis

Macalister (1872) observed and described the serratus magnus and

divided the muscle into superior and inferior portions. The same

muscle was described by Vaughan (1959) under the name serratus

anterior. He also divided the serratus anterior into two parts, the

anterior and posterior divisions. Subsequent authors followed his

precedent for bats (Walton, 1967; Norberg, 1970; Strickler, 1978;

Altenbach, 1979; Hermanson and Altenbach, 1981, 1983).

Serratus Ventralis Thoracis, Cranial Division

Form. The origin is a fibrous attachment on the dorsolateral

surface of the first rib, first intercoastal space, and cranial surface

of the second rib, medial to the origin of subclavius. The muscle has

a broad fleshy insertion along the ventral surface of the craniomedial

flange of the scapula, lateral to the insertion of serratus ventralis

cervicis. At the origin, the belly is small and rounded. Distally the

belly becomes wider and flatter, being thickest along the medial

border. The muscle is innervated by a branch of the dorsal scapular

nerve.



































Figure 15.--Lateral view of serratus ventralis musculature.
In the upper figure, subclavius is intact. In the lower
figure, subclavius is removed to reveal the position of the
deeper serratus ventralis cervicis.



















serratus ventralis


scapula


coracoid process


serratus


serratus ventralis thoracis,

...---- cranial div.
\" .... caudal div.







i:-



















serratus ventralis thoracis,
.... cranial div.
S caudal div.
.~----~


subclavius











Comparative aspects. The cranial division of serratus ventralis

thoracis is large in hovering bats and in slow-flying bats (Strickler,

1978).

Functional aspects. Vaughan (1959) proposed that the cranial

division functioned as a wing abductor by pulling ventrally on the

craniomedial edge of the scapula. The cranial division is large in

hovering bats, the glossophagines, and slow-flying bats, the

rhinolophids (Strickler, 1978). In both groups, the wingbeat amplitude

is large and may include extensive tipping of the lateral border of the

scapula ventrad during the downstroke. The cranial division is

situated to initiate abduction of the wing by arresting outward

rotation and by initiating inward rotation of the scapula. No EMG data

are available to test this hypothesis.

Serratus Ventralis Thoracis, Caudal Division

Form. The muscle originates by fibers along the dorsolateral

aspect of ribs two through 11, adjacent and dorsal to each

costochondral junction. A small, separate slip originates from the

caudolateral aspect of the first costal cartilage, caudal to the origin

of subclavius. The fibers of this small slip course caudally to insert

on the middle one-third of the axillary border of the scapula. The

fibers of the larger portion of the caudal division insert by fleshy

attachment along the caudal three-quarters to seven-eighths of the

axillary border and on the caudal angle of the scapula. The caudal

fibers inserting upon the caudal angle constitute the thickest portion











of the muscle. Fibers are arranged in parallel. The fibers

interdigitate at their origin with the scaleneus muscles cranially and

the external abdominal oblique muscles caudally. Innervation of the

caudal division is by the long thoracic nerve, coursing along the

lateral surface of the muscle.

Comparative aspects. Strickler (1978) found that the caudal

division is relatively large in most phyllostomids and small in the

Pteropidae and most of the Verpertilionidae. He felt that the function

and size of the muscle vary with respect to the development of the

humeroscapular locking mechanism.

Functional aspects. The shoulder locking mechanism described by

Vaughan (1959) provides a mechanical link between the action of the

caudal division upon the scapula and the resultant adduction of the

wing. Thus, contraction of the serratus ventralis thoracis was

hypothesized to power the downstroke. Electromyograms obtained from

Artibeus indicate that the posterior division has a major period of

activity that commenced before onset of the downstroke and ended during

the early downstroke. A second burst of activity is of shorter

duration and occurs during the early upstroke phase. Hermanson and

Altenbach (1981, 1983) reported a single period of activity for this

muscle in Antrozous during the transition between the upstroke and

downstroke. In both Antrozous and Artibeus, the pectoralis muscles are

active prior to the major burst of the caudal division of serratus

ventralis thoracis. The caudal division and pectoralis are coactive as

adductors throughout the early downstroke.











The second period of EMG activity occurs after the upstroke has

begun. This burst provides a force antagonistic to activity in

clavodeltoideus and latissimus dorsi. Clavodeltoideus activity tends

to protract the humerus, and thus advances the scapula craniad relative

to the thorax. Latissimus dorsi exhibits its most pronounced activity

as an abductor and pronator of the wing, causing retraction of the

scapula and arm.

English (1978a, 1978b) presented data for the serratus ventralis

cervicis and serratus ventralis thoracis of cats indicating biphasic

activity patterns. He concluded that serratus ventralis thoracis is

important in transmitting the weight of the body to the forelimbs

during the stance phase, an observation suggested by earlier anatomists

working only with cadaverous material (Davis, 1949; Gray, 1968). Of

greater interest was English's observation that coactivation of

cervical and thoracic portions of serratus ventralis during the swing

phase effected shoulder extension and repositioning of the limb for the

following step. Jenkins and Weijs (1979) found no significant

difference between EMG activity of different slips of serratus

ventralis thoracis in Didelphis. During walking, serratus ventralis of

Didelphis was active during the propulsive phase. These authors

commented upon a secondary period of activity during the swing phase in

Didelphis, but did not discuss its role. A secondary phase of activity

was not recorded in Antrozous (Hermanson and Altenbach, 1983).

Repositioning of the limb for the downstroke may be effected in











Antrozous during the upstroke by the force of the airstream on the wing

(Vaughan, 1959; Norberg, 1976) and by the action of the dorsal shoulder

musculature (Hermanson and Altenbach, 1983). The greater wingloading

in Artibeus may require an additional burst of muscle activity.

Rhomboideus

Form. The rhomboideus originates by fibers on the dorsal surface

of the transverse process of cervical vertebrae seven, lateral to the

cranial fibers of clavotrapezius, through thoracic vertebra six,

lateral to the proximal fibers of acromiotrapezius. The muscle fibers

course obliquely, caudodorsally through the interscapular region to

insert on the entire vertebral border of the scapula caudal to the

scapular spine. Innervation is by the dorsal scapular nerve.

Comparative aspects. The rhomboideus of the Phyllostomidae was

characterized by Strickler (1978) as being relatively small. In

contrast, both Macalister (1872) and Walton (1967) described the muscle

as robust in the Phyllostomidae. I concur with Strickler's

interpretation in part because of the muscle mass data presented in his

arguments, whereas the other authors relied upon subjective

interpretations. The romboideus is a thin, flat muscle in Artibeus.

Strickler (1978) hypothesized that either rhomboideus or elevator

scapulae are well developed in bats depending on locomotor

requirements. In the Molossidae, he observed that the rhomboideus was

relatively large and adapted to rotating the caudal angle of the












scapula medially relative to an axis perpendicular to the dorsal

surface of the scapula. He speculated that this action is important in

terrestrial movements and not in the scapular movements associated with

the phyllostomid shoulder during flight.

Functional aspects. Based upon the attachments and orientation, I

deduce that rhomboideus pulls the caudal angle of the scapula

ventromedially. There are no EMG data with which to elaborate upon

this hypothesis.



Latissimus-subscapular Group

Latissimus dorsi

Form. The muscle originates from the superficial fascia overlying

thoracic vertebra eleven through lumbar vertebra four. At the origin,

the belly is thicker over thoracic vertebrae and grades into a thin

aponeurosis caudally over the lumbar region. Latissimus dorsi inserts

by a thin, 1 mm wide tendon 3-4 mm distal to the shoulder joint on the

medial ridge of the humerus. The tendon lies ventral to and on the

proximal edge of the fibrous insertion of teres major. Fibers in the

latissimus dorsi are arranged in parallel at the origin, but converge

distally on the tendon of insertion. Innervation is received from the

thoracodorsal nerve.

Comparative aspects. Strickler (1978) did not observe much

variation in the latissimus dorsi attachments of bats, except in the

Rhinolophidae. In several phyllostomid species studied, the origin











of latissimus dorsi ranged between thoracic vertebra 10 and lumbar

vertebra 5, but no correlation was established between the attachments

and taxonomy or locomotor modes. Strickler's muscle mass data

indicated that the muscle was largest in slow fliers, and average-sized

in hovering and moderately fast frugivorous bats.

Functional aspects. The action of latissimus dorsi for Artibeus

and for bats in general is confusing because of conflicting evidence

from several sources. Latissimus dorsi was described by Vaughan (1959)

as a pronator and flexor of the humerus, and also as an important

retractor during terrestrial locomotion (1970b). Strickler (1978)

added that the muscle might also serve in abduction of the wing during

upstroke movements. For Desmodus, Altenbach (1978) obtained EMG data

suggestive of a major role in humerus pronation and adduction, but not

in abduction. In Antrozous latissimus dorsi functions primarily as a

pronator during the downstroke (Hermanson and Altenbach, 1983).

The EMG data for latissimus dorsi in Artibeus are difficult to

interpret in comparison with the data obtained from other studies. In

17 wingbeats, latissimus dorsi exhibits a biphasic activity pattern,

with highest intensity EMG myopotentials observed during the late

downstroke to early upstroke transition period (onset mean was 0.329,

average onset of downstroke movements was 0.428). A second period of

activity, of shorter duration and with lower amplitude mypotentials,

occurs during the late upstroke and early downstroke. These patterns

are consistent in all wingbeats. These data contrast with the EMG











pattern reported for Antrozous (Hermanson and Altenbach, 1983).

Although the greatest activity for latissimus dorsi in Artibeus was

observed during the late downstroke, the muscle was quiescent in

Antrozous during the same period.

English (1978a) observed variability in the EMG patterns of

latissimus dorsi in cats. Although the muscle was characterized by

English as monophasic and active predominantly during the propulsive

phase (EI-E2), he observed inconsistent bursts of activity during the

swing phase (F). These inconsistent bursts are similar in timing to

the late downstroke activity observed in Artibeus. Jenkins and Weijs

(1979) reported monophasic activity in the posterior part of latissimus

dorsi of walking Didelphis, however, they noted a consistent biphasic

pattern for recordings made with electrodes located in the anterior

fibers of the muscle. Placement of electrodes in intermediate

positions resulted in "hybrid" EMG patterns: this region of latissimus

dorsi was monophasic during some step cycles and biphasic during

others. Tokuriki (1973a, 1973b, 1974) observed two distinct phases of

activity in the latissimus dorsi of dogs at all gaits. One phase

occurred with other flexor muscles during the F phase, contributing to

lift the limb off the ground. A second and evidently more intense

period of activity occurred throughout the propulsive phase. In

terrestrial mammals, the activity of latissimus dorsi appears to be

biphasic, although the most intense activity occurs during a different

period of the locomotory cycle than observed in Artibeus.











Based on my EMG data and on comparative data for other mammals, I

conclude that latissimus dorsi in both Artibeus and Antrozous

stabilizes the shoulder and pronates the wing throughout the

downstroke. Primary EMG activity occurs during the transition from

downstroke to upstroke. Assuming a contraction time of 25 msec (Burke,

1978; Hermanson and Altenbach, 1981), the action of this muscle burst

pronates the wing during the first third of the upstroke. Analysis of

our film records indicates that the wing is continuously supinated

during the first two-thirds of the upstroke (Norberg, 1976; Altenbach,

1978), an action effected by the dorsal abductor muscles and by the

force of the airstream (Vaughan, 1959). The large burst of activity in

latissimus dorsi may therefore be necessary to counteract the

supinators and stabilize the shoulder during the early upstroke. The

temporal activity pattern of latissimus dorsi indicates that the muscle

is primarily a wing abductor.

Teres Major

Form. The teres major originates on the lateral and dorsal aspect

of the caudal angle of the scapula, and along the caudal 3 mm of the

axillary border of the scapula, by fibers. The origin does not extend

to the cartilaginous expansion at the caudal angle of the scapula. The

muscle inserts along the medial ridge of the humerus, 3-6 mm distal to

the shoulder joint. The insertion is primarily a fibrous attachment,

adjacent to but not conjoined with the tendon of insertion of the

latissimus dorsi. All myofibers are arranged in parallel. Innervation

is provided by several branches of the subscapular nerve.











Comparative aspects. Vaughan (1959) observed that the teres

major in Macrotus (Phyllostomidae) is larger relative to the latissimus

dorsi, than in Myotis (Vespertilionidae) and Eumops (Molossidae).

Strickler (1978) found that both the teres major and latissimus dorsi

exhibit positive allometry with respect to body mass. Triceps brachii

(long head) and biceps brachii (short head) were the only other muscles

in which Strickler observed positive allometry. He hypothesized that

whereas musculoskeletal specializations or lift provided by the

airstream permitted isometry to prevail in the primary adductors, and

abductors, respectively, the muscles associated with powering the flick

phase of the upstroke cannot be assisted by outside forces. Therefore,

a greater degree of positive allometry is exhibited in these four

muscles than in other muscles. Strickler did not include Artibeus in

his sample but did study other phyllostomids, including Carollia and

Phyllostomus, representative of locomotor styles broadly comparable to

Artibeus.

Functional aspects. Electromyographic data exhibit two phases of

activity. High-amplitude activity occurs during the late downstroke to

upstroke transition. A second burst of one-half amplitude activity

occurs during the late upstroke to downstroke transition. The

association of muscle activity with turnover periods of the wingbeat

cycle indicates these muscles play an important role in stabilizing the

shoulder joint. Previous investigators used dissection studies and

determined the role of teres major is responsible for pronation of the

wing and a flexion of the shoulder joint (Vaughan, 1959; Norberg,











1970, 1972; Strickler, 1978; Altenbach, 1979). Hermanson and Altenbach

(1983) reported EMG data for four wingbeats in Antrozous but made no

specific conclusions beyond the classification of teres major as a

bifunctional muscle. In Artibeus, the biphasic activity pattern is

observed but is significantly different from that observed in

Antrozous. The primary activity of teres major in Antrozous, during

the final stages of the downstroke, probably pronates the wing

synergistically with latissimus dorsi. Together, the two muscles

counteract the onset of supination effected by the abductor muscles and

airstream, and provide a smooth transition of wing movements between

the downstroke and upstroke.

Subscapularis

Form. The subscapularis fills the entire subscapular fossa. The

muscle originates along the periphery and on the elevated ridges of the

subscapular fossa, and from the ventral surface of the cartilaginous

expansion of the caudal angle of the scapula. The fibers exhibit a

multipinnate arrangement, with two main tendons converging to the

insertion on the lesser tuberosity of the humerus. The insertion is

composed of a broad tendinous attachment, and by fibers coursing

lateral of the tendon. Innervation is received from the two

subscapular nerves, derived from the dorsal division of the brachial

plexus.

Comparative aspects. The attachments of the subscapularis are

conservative in all bats (Strickler, 1978). The large size of the











subscapularis relative to body mass was noted by Macalister (1872) as

being larger than observed in other mammals. Vaughan (1970a)

elaborated upon this point and provided data for the mass of

subscapularis relative to the total mass of the primary downstroke

muscles. This value ranged from 16.1 to 19.4 percent. Overall, the

relative weights of these primary downstroke muscles are consistent in

all bats. The data from Strickler (1978) are in agreement with Vaughan

except for in the Pteropodidae and for the phyllostomid genus

Desmodus. Strickler commented that pteropodids were unique among bats

in having small subscapularis muscles and small ventral muscle masses.

It was not established whether the small muscle masses represent a

primitive condition for chiropterans, or result from functional

constraints. In contrast, the large subscapularis muscle of Desmodus

appeared to be related to the terrestrial habits of the vampire bat,

for which there are increased demands for shoulder extension and wing

adduction (Strickler, 1978).

Functional aspects. Electromyography of the subscapularis in

free-flying Artibeus indicates that the muscle remains active through

approximately 75 percent of the wingbeat cycle. High-amplitude EMG

activity is observed through approximately 26 percent of the wingbeat

cycle and coincides with the second one-half of the upstroke and first

one-tenth of the subsequent downstroke. This activity precedes

myopotentials recorded from the other two downstroke muscles, serratus

ventralis thoracis and pectoralis. Subscapularis was discussed as a











component of the rotator cuff complex of primates and other mammals

(Tuttle and Basmajian, 1978) and is important in stabilizing the

shoulder joint. This stabilization includes the role of holding the

articular surface of the humerus within the glenoid fossa. Hermanson

and Altenbach (1981, 1983) concluded that subscapularis was primarily

responsible for fine control and stability of the shoulder joint in

Antrozous. They also observed that the expression of continuous versus

episodic activity in subscapularis was facultative and therefore

dependent upon the nature of the flight maneuver being attempted at any

given time. Kovtun and Moroz (1974) also used electromyography and

observed that subscapularis preceded pectoralis activity and ceased

activity 6-10 msec after mypotentials terminated in the "chest muscles"

of Myotis myotis. It was not clear whether their data were pooled from

several wingbeats or represented only one wingbeat.

The timing of subscapularis activity does not support Vaughan's

hypothesis (1959) that subscapularis is the third muscle among the

primary downstroke muscles to commence activity during a wingbeat.

However, the high-amplitude activity fit well with the temporal pattern

observed in downstroke muscles: myopotentials commence approximately

25 msec before wing adduction begins and terminates during the early

downstroke. Low-amplitude myopotentials during the early downstroke

and early upstroke indicate stabilization activity at the shoulder

joint. Low-amplitude activity at the transition between downstroke and

upstroke, and during the early upstroke stabilizes the shoulder against











forces produced by the abductor muscles during this period. Because of

these two periods of activity and the roles that the muscle carries

out, adduction and stabilization, the subscapularis is classified as a

bifunctional muscle and serves primarily to stabilize the shoulder

joint.



Deltoid Group

The deltoideus of bats has long been recognized as a tripartite

muscle composed of three parts: clavodeltoideus, acromiodeltoideus,

and spinodeltoideus (Macalister 1872). Because of functional

specializations associated with each of these parts, I will describe

and refer to each part specifically. All parts are innervated by the

axillary nerve.

Clavodeltoideus

Form. The clavodeltoideus originates by fibers along the

expanded, distal three-quarters of the clavicle. The muscle fibers

course dorsally, proximal to the apex of the pectoral ridge of the

humerus, and insert near the proximal insertion of acromiodeltoideus.

Fibers are arranged in parallel.

Comparative aspects. Vaughan (1959) pointed out that the

clavodeltoideus is separated from adjacent fibers of the pectoralis

only with great difficulty. Visualization of the axillary nerve is

necessary to correctly identify muscle bundles belonging to the











deltoideus group. Strickler (1978) illustrated some of the variation

in extent of origin that is possible within the order Chiroptera and

thus the need to undertake careful dissection of this region.

Functional aspects. The functional analysis of clavodeltoideus is

discussed along with the spinodeltoideus.

Acromiodltoideus

Form. The acromiodeltoideus has a fibrous origin along the entire

tip of the acromion process and proximally, along the base of the

acromion process as it intergrades with the scapular spine. Insertion

is by fibrous attachment along the lateral ridge of the humerus, from 1

mm distal to the greater tubercle along the proximal one-third of the

humerus. The myofiber architecture is parallel with some fibers

diverging distally along the insertion. A thin fascial plane bisects

the muscle into anterior and posterior portions and extends distally

from the middle of the acromion process.

Comparative aspects. The primitive condition for the origin of

the chiropteran acromiodeltoideus is restricted to the acromion process

(Strickler, 1978). In advanced families, the origin of the muscles has

extended medially along the scapular spine, a position that places

fibers farther from the shoulder joint and that increases the

effectiveness of acromiodeltoideus as an abductor and flexor of the

shoulder (Strickler, 1978). In Antrozous, fibers of acromiodeltoideus

and acromiotrapezius attach to each other along the dorsal scapular











ligament (Hermanson, 1978). The situation in Artibeus, in contrast, is

for both acromiodeltoideus and acromiotrapezius to insert upon a solid

mass of bone, adjacent to the acromion process.

Functional aspects. The action of the acromiodeltoideus is

discussed along with the spinodeltoideus.

Spinodeltoideus

Form. The spinodeltoideus originates by fibers from the dorsal

scapular ligament (sensu Hermanson and Altenbach, 1983), from the

vertebral border of the scapula caudal to the scapular spine, and over

the cartilaginous plate at the caudal angle of the scpaula. Insertion

is by an aponeurosis on the medial aspect of the dorsal ridge of the

humerus, deep to the insertion of the acromiodeltoideus and immediately

distal to the insertion of teres minor. The muscle is parallel-fibered

and forms a broad but thin sheet over the caudal scapular region.

Vaughan (1959) commented that spinodeltoideus was composed of two parts

in Macrotus.

Functional aspects. The deltoideus muscle of man is a single

muscle mass that was divided by anatomists into an anterior and

posterior portion. Each division can be correlated with differential

EMG activity while performing several actions (Inman et al., 1944).

Similarly, Hermanson and Altenbach (1983) presented functional

interpretations for all three heads in Antrozous and discussed these

data with respect to published data for several terrestrial mammals.

The acromiodeltoideus and spinodeltoideus EMG patterns for Antrozous











were in general agreement with the hypotheses of earlier descriptive

studies (Vaughan, 1959, Strickler, 1978): the muscles functioned as

abductors of the wing. The clavodeltoideus, however, was active almost

in synchrony with the primary downstroke muscles. This observation,

and dissection studies (Norberg, 1972; Strickler, 1978), led Hermanson

and Altenbach to conclude that the muscle was an important humeral

protractor and also a humeral adductor.

During the propulsive phase in Antrozous, a lack of muscle

activity in acromiodeltoideus and spinodeltoideus contrasted with

observations of EMG patterns in the deltoideus of terrestrial mammals.

Deltoideus activity occurred during the stance phase in stepping cats

at several speeds of locomotion (English, 1978a). Biphasic activity

occurred in the deltoideus muscle in Didelphis: the largest amplitude

activity occurred during the swing phase, and a less intense period of

activity during the stance phase (Jenkins and Weijs, 1979). When the

acromioclavicular and spinal portions of the Didelphis deltoideus were

monitored simultaneously, the spinal portion exhibited intense activity

during the first one-third of the swing phase, while the

acromioclavicular portion was active during the final two-thirds of the

swing phase (Jenkins and Weijs, 1979). Recordings from the deltoideus

of Canis indicated continuous myopotentials throughout the step cycle

of walking, trotting, and galloping dogs (Tokuriki, 1973a, 1973b,

1974).











Data for the deltoideus of Artibeus provide contrasts and

similarities with the Antrozous observations. For example, the primary

(highest amplitude) EMG activity in the clavodeltoideus was evident

during the downstroke portion of the wingbeat. Myopotentials commenced

almost in synchrony with the pectoralis, contributing to wing adduction

but also to the anterior movement of the wing. This wing protraction

is an important aspect of the "Rudderflug" or rowing flight described

by Eisentraut (1936) and Norberg (1976). Low-amplitude activity of

brief duration occurs during the transition between downstroke and

upstroke and serves to stabilize the scapula. In contrast,

highest-amplitude activity in both the acromiodeltoideus and

spinodeltoideus occurred during the late downstroke and early

upstroke. These data therefore suggest an important role for

acromiodeltoideus and spinodeltoideus in wing abduction and agree with

observations in Antrozous (Hermanson and Altenbach, 1983). Both

acromiodeltoideus and spinodeltoideus were biphasically active. The

posterior part of acromiodeltoideus showed a low-amplitude phase of

activity roughly coincident with the primary burst of clavodeltoideus,

although of shorter duration and with less amplitude than in the latter

muscle. Spinodeltoideus exhibited low-amplitude activity during the

first one-third of the downstroke. These secondary EMG bursts in the

acromiodeltoideus and spinodeltoideus may stabilize the shoulder

against the powerful actions of the pectoralis.












Suprascapular Group

Supraspinatus

Form. Supraspinatus originates by fibers along the cranial and

medial perimeter of the supraspinous fossa, deep to the belly of

acromiotrapezius. The muscle fibers are arranged in parallel. This

muscle has a fibrous insertion on the tip of the greater tubercle of

the humerus. Innervaton is by the suprascapular nerve.

Comparative aspects. The supraspinatus exhibits little variation

in its attachments for all bat species (Vaughan, 1959, 1970a;

Strickler, 1978). The regression of supraspinatus mass against body

mass suggests that the size of this muscle is average to relatively

large size in slow-flying bats, and smaller in faster-flying species

(Strickler, 1978).

Functional aspects. The mammalian supraspinatus is considered one

of the rotator cuff muscles of the shoulder: its primary role is to

stabilize the glenohumeral articulation in concert with the

infraspinatus, subscapularis, and teres minor (Hollinshead, 1974;

Tuttle and Basmajian, 1978). For bats, Vaughan (1959) described the

general action of the supraspinatus to be shoulder extension and

humerus supination, a description accepted by subsequent authors (cf.

Strickler, 1978). Vaughan (1959) suspected that supraspinatus was most

effective as a shoulder extensor in the phyllostomid Macrotus, as

compared to Eumops and Myotis. The shoulder joint of Macrotus allows











more freedom of movement than is observed in Eumops and Myotis. Both

of the latter species had bony locking mechanisms that restricted

movement of the humerus in both the dorsoventral and craniocaudal

directions. Thus, the humerus of Macrotus could be extended cranial

to a transverse plane through the shoulder, while the humerus of Eumops

could only be extended forward to this plane.

Electromyographic data for the supraspinatus is available for only

six wingbeats, primarily because the small size and deep position of

the muscle make the surgical implantation a traumatic procedure for the

animal. The muscle exhibited a single, extended period of activity

commencing during the early upstroke, and continuing through the early

downstroke. During two of the wingbeats, supraspinatus exhibited two

periods of normal-amplitude EMG activity with a brief period of

quiescence interposed.

Biphasic EMG activity was observed in supraspinatus of Antrozous

(Hermanson and Altenbach, 1983). Biphasic activity was also observed

during the step cycle of Didelphis (Jenkins and Weijs, 1978), but not

during stepping in cats (English, 1978a) or dogs (Tokuriki, 1973a,

1973b, 1974). In several primate species, Tuttle and Basmajian (1978)

reported prominent activity of the supraspinatus during humeral

elevation and cautioned that the function of this muscle may be one of

acceleration of the humerus rather than as a simple stabilizer (i.e.,

rotator cuff). The variability observed in all of these studies may be

attributed to small sample sizes, at least for the two bat species











studied, and to an inconsistent pattern of locomotor movements studied

in the primate examples. In the latter study, movements of the animal

were not restricted to forward progression, but instead included

forward, lateral, and vertical movements. Support for a role in wing

supination is provided by EMG data indicating activity during the early

upstroke when the wing is supinated and abducted.

Infraspinatus

Form. The infraspinatus originates from the periphery of the

infraspinous fossa of the scapula: cranially from the spine of the

scapula; medially from the dorsum of the vertebral border of the

scapula; and laterally from the raised ridge of the infraspinous

fossa. All fibers insert upon a central tendon that courses through

the middle of the muscle. This tendon forms the sole insertion of the

infraspinatus upon the lateral ridge of the humerus. Fibers at the

medial end (nearest the vertebral border) are the longest in the muscle

and insert upon the central tendon at an acute angle. Fibers that

insert more distally on the central tendon are shorter and tend to

intersect the tendon at a larger angle than the medial fibers.

Innervation is by the suprascapular nerve, after that nerve provides a

branch to the supraspinatus and courses laterally under the tip of the

acromion process.

Comparative aspects. Strickler (1978) noted little variation in

the attachments of the infraspinatus in all bat species that he

studied. In contrast to the regression of muscle mass on body mass











obtained for supraspinatus, Strickler observed that the muscle was

larger in fast-flying species, but smaller in fast-flying frugivores.

No functional explanation was provided. Within families, there was no

consistent ratio observed between the mass of infraspinatus and

supraspinatus.

Functional aspects. Two phases of activity were evident in the

infraspinatus of Artibeus. The first period and the one exhibiting the

highest amplitude myopotentials commenced immediately prior to the

onset of downstroke movements, and continued during the first third of

the downstroke. A second, less intense period of activity was observed

in six wingbeats during the late downstroke and early upstroke. These

data were similar to observations made on the infraspinatus of

Antrozous (Hermanson and Altenbach, 1983). Although two distinct

phases were also observed in walking Didelphis (Jenkins and Weijs,

1979), only one period of activity was noted in cats (English, 1978)

and dogs (Tokuriki, 1973a, 1973b, 1974) at all gaits.

On the basis of anatomical position the infraspinatus is a

shoulder flexor, and a humerus abductor and supinator (Hermanson and

Altenbach, 1983; Strickler, 1978; Altenbach, 1979). The biphasic

activity pattern suggests an underlying stabilizing role at the

shoulder; the secondary EMG activity observed during the early

downstroke counteracts the action of the pectoralis and protects the

glenohumeral joint in association with other intrinsic muscles of the

shoulder. Because the wing of a bat is not capable of being











"feathered" during the upstroke like that of a bird, the role of the

abductors is accentuated in bats (Vaughan, 1959). Thus, the

infraspinatus exhibits strong activity during the non-propulsive

upstroke, and acts in concert with the more superficial spinodeltoideus

to abduct the wing.



Triceps Group

The triceps brachii is composed of three heads in most mammals.

Each head is innervated by branches of the radial nerve. Because of

the different origins of these heads, each is treated separately.

Triceps Brachii, Medial Head

Form. The medial head has a fleshy origin along the caudal

surface of the humerus, approximately 10-13 mm distal to the greater

tubercle. Muscle fibers pass medial to the radial nerve. Fibers

insert distally in a unipinnate fashion on the main triceps brachii

tendon. This is the smallest of the three heads of triceps brachii.

Comparative aspects. In most phyllostomids, the medial head

originates on the middle one-third of the ventral surface of the

humerus (Strickler, 1978). The attachments are conservative in most

chiropterans.

Functional aspects. The medial head of triceps brachii spans one

joint, the elbow. Vaughan (1959) suggested that the medial head

stabilizes the sesamoid bones found at the insertion of the triceps

brachii tendons in many bats. No EMG data are available for any bats

to corroborate or refute the hypothesis.












Triceps Brachii, Lateral Head

Form. The lateral head originates by fibers on the proximal

one-quarter of the humerus, primarily on the caudal surface adjacent to

the greater tubercle, surgical neck, and lesser tubercle. All fibers

pass lateral to the radial nerve. Central fibers insert directly upon

a central tendon of insertion. Peripheral fibers insert on either side

of this tendon at an angle of less than 10 degrees.

Functional aspects. The lateral head of triceps brachii crosses

one joint, the elbow joint, and is capable to effecting extension of

that joint. EMG data is discussed along with the long head of triceps

brachii.

Triceps Brachii, Long Head

Form. The long head arises from the axillary border of the

scapula, adjacent to the infraglenoid tubercle. The muscle is

pinnate. Fibers course distally to insert upon a central tendon in

common with the other two heads of triceps brachii. The main

insertional tendon contains a small sesamoid caudal to the elbow

joint. A synovial cavity is present deep to the tendon, permitting

movement of the tendon relative to the joint in this area. The

insertional tendon of the triceps brachii attaches to the caudal

surface of the olecranon region of the ulna.

Comparative aspects. The anatomy of the triceps brachii does not

depart significantly from the general pattern observed in other bats

(cf. Strickler, 1978): the long head spans the shoulder and elbow

joints, the lateral head spans only the elbow joint.












Functional aspects. In terrestrial mammals, the triceps brachii

function as anti-gravity muscles principally by maintaining elbow

extension during standing or locomotion (Armstrong et al., 1982;

English, 1978b; Jenkins and Weijs, 1979). EMG data for long and

lateral head of triceps brachii in Antrozous did not clearly support an

elbow extensor hypothesis (Hermanson and Altenbach, 1983). In A.

jamaicensis, muscle activity in the long head commenced prior to

initiation of the upstroke, and continued throughout the first one-half

of the upstroke. The lateral head also commenced activity prior to the

downstroke, but relative to the long head, EMG activity began later and

was of shorter duration. These temporal patterns are similar to

observations of triceps brachii activity in Antrozous (Hermanson and

Altenbach, 1983). In both bat species, the long and lateral heads of

triceps brachii exhibited activity patterns assoicated with wing

abduction. It is not clear how either portion of triceps brachii

effects elbow extension, a movement associated with the downstroke

phase of the wingbeat cycle (Eisentraut, 1936; Norberg, 1976; Hermanson

and Altenbach, 1983). Elbow extension automatically contributes to

spreading the distal wing during the downstroke in many bat species

(Vaughan, 1966; Vaughan and Bateman, 1970). In Artibeus, the amount of

time between EMG onset and downstroke initiation was approximately 55

msec. Armstrong et al. (1977) and Hermanson and Foehring (1982)

demonstrated the existence of fast type histochemical profiles for

triceps brachii in Myotis and Tadarida, respectively. In several

mammalian muscles, fast type profiles are correlated with contraction




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PAGE 173

81,9(56,7< 2) )/25,'$


FUNCTIONAL MORPHOLOGY AND FLIGHT KINEMATICS OF
Artibeus jamaicensis (CHIROPTERA, PHYLLOSTOMIDAE)
By
JOHN W. HERMANSON
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA

Copyright 1983
by
John W. Hermanson

ACKNOWLEDGEMENTS
Financial support for this study was provided in part by a
Grant-in-Aid of Research from the Society of Sigma Xi; from the
Department of Zoology, University of Florida; and from the Department
of Natural Sciences, Florida State Museum. I received support in the
form of a Museum Assistantship in the Department of Natural Sciences,
Florida State Museum, and a Graduate Teaching Assistantship in the
Department of Physiological Sciences, College of Veterinary Medicine.
Sr. Tomas Blohm kindly allowed me to study on his ranch in
Venezuela in 1981. Dr. John Robinson invited me to travel with him to
Venezuela, and then took care of the necessary procedures to obtain
permission for me to collect and transport live bats in that country.
I also acknowledge the Department of Biology, University of New Mexico,
for allowing me free access to the facilities of the Museum of
Southwestern Biology.
I thank my committee members for advice and attention at a time
when none of them needed the additional burden of a dissertation to
review. My committee included Drs. John Anderson, Donald Dewsbury,
David Webb, Ronald Wolff, and Charles Woods. Pamela Johnson typed and
then retyped the manuscript. I appreciate her patience in this task.
iii

I offer thanks to the four men in biology who have most influenced
my path. Dr. Scott Altenbach, of the University of New Mexico, opened
his laboratory and his home to me during our joint venture. We will
have many enjoyable moments to reflect on during the years to come.
Dr. David Klingener, of the University of Massachusetts, taught a
fabulous course in comparative anatomy and opened my eyes to a career
in biology. Dr. Ted Goslow, of Northern Arizona University, provided
me with high goals in both my personal and academic life. Finally, my
advisor Dr. Charles Woods has provided friendship and a wealth of
experiences in natural history and anatomy on which to build a career.
I am fortunate to have studied with these four gentlemen.
I thank my peers for those good moments that make graduate
education unique. These moments include late nights out on the silent
Arizona desert, evenings of light snow or Great-horned owl hootings in
the Green Mountains, or simply industrious evenings in the labs at the
Museum. In particular, I will miss those evenings out on the back
porch with the folks in our household at Newnan's Lake. All of these
times are irrevocably tied together in my mind.
Last but not least I wish to acknowledge my family. My mother and
father endured my endeavor and provided encouragement and support. My
aunt, Catherine Leonard, has provided inspiration and the gift of
laughter. My "other aunt," Ruth Cushman, has been a friend
throughout. Finally, I remember my companion of many treks, Japhy.
iv

TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS iii
LIST OF FIGURES vi
LIST OF TABLES viii
ABSTRACT ix
INTRODUCTION 1
MATERIALS AND METHODS 10
FLIGHT AND AERODYNAMICS
Results 15
Discussion 24
OSTEOLOGY 31
MYOLOGY
Trapezius Group 53
Costo-spino-scapular Group 62
Latissimus Group 71
Deltoid Group 79
Supraspinous Group 84
Triceps Group 88
Pectoralis Group
Flexor Group of Arm 103
Antebrachial Extensor Group 103
Antebrachial Flexor Group 123
ELECTROMYOGRAPHY OF FLYING BATS 133
CONCLUSIONS 149
LITERATURE CITED 153
BIOGRAPHICAL SKETCH 160
v

LIST OF FIGURES
Page
1. The wingbeat cycle of Artibeus jamaicensis. 7
2. Wing and body movements during slow forward flight in 17
Artibeus jamaicensis.
3. Wing and body movements during slow forward flight in 19
Artibeus jamaicensis.
4. Wing and body movements relative to still air during 21
slow forward flight in Artibeus jamaicensis.
5. High- and low-aspect ratio wingshapes. 28
6. Lateral view of the thoracic and axillary skeleton. 33
7. Cranial view of the right pectoral girdle. 35
8. Dorsal view of the scapula. 37
9. Medial view of the left humerus. 41
10. Articular surfaces of the humerus. 43
11. Dorsal view of the radius and ulna. 48
12. Dorsal and ventral views of the carpus. 51
13. Dorsal view of the shoulder and arm of Artibeus jamaicensis. 56
14. Activity patterns of shoulder and arm muscles in Artibeus 58
during slow flight.
15. Lateral view of serratus ventralis musculature. 66
16. Ventral view of the shoulder and arm of Artibeus 94
jamaicensis.
17. Electromyographic data for six regions in the pectoralis 99
muscle during slow flight.
18. Lateral view of the muscles of the elbow region. 108
vi

Page
19. Dorsal view of the muscles of the carpal region. 110
20. Medial view of the muscles of the elbow region. 125
21. Ventral view of the muscles of the carpal region. 127
22. Numbers of coactive muscles during the wingbeat cycle. 145
vii

LIST OF TABLES
Page
1. Aerodynamic parameters of the wing in two fruit bats, 22
Artibeus jamaicensis.
2. Electromyographic data for activity patterns of the 59
shoulder musculature in Artibeus jamaicensis during
slow flight.
3. Electromyographic data for activity patterns in six 100
regions of the pectoralis muscle of Artibeus
jamaicensis during slow flight.
viii

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
FUNCTIONAL MORPHOLOGY AND FLIGHT KINEMATICS OF
Artibeus jamaicensis (CHIROPTERA, PHYLLOSTOMIDAE)
By
John W. Hermanson
April 1983
Chairman: Dr. Charles A. Woods
Major Department: Zoology
The functional morphology of the pectoral girdle and arm of
Artibeus jamaicensis (Chiroptera, Phyllostomidae) is interpreted on
the basis of gross dissection and electromyographical analysis (EMG).
Electromyographic data obtained during flight for 15 muscles elucidate
several temporal patterns of activity associated with the wingbeat
cycle that are not similar to the patterns of flexor, extensor, and
bifunctional muscles observed in terrestrial mammals. Abductor
muscles exhibit intense activity associated with the early upstroke
phase of the wingbeat cycle and include clavotrapezius, acromiotrapezius,
latissimus dorsi, teres major, acromiodeltoideus, spinodeltoideus, and
triceps brachii (long and lateral heads). All abductors except for the
triceps brachii exhibit a secondary period of low-amplitude activity
associated with the early downstroke. Adductor muscles exhibit primary
activity immediately prior to and during the early downstroke phase.
The adductors include serratus ventralis thoracis, pectoralis, and
ix

clavodeltoideus. Bifunctional muscles exhibit a single period of
activity through most of the wingbeat cycle, or two separate periods
of high-amplitude activity during a wingbeat cycle. The bifunctional
muscles include spinotrapezius, supraspinatus, infraspinatus, and
subscapularis. Dissection of all other muscles of the shoulder and
antebrachium form the basis of interpreting musculoskeletal movements
during flight in Artibeus. The major muscles of support in Artibeus
include serratus ventralis thoracis, pectoralis, and the trapezius
group. These muscles support the trunk between the wings during
flight or terrestrial locomotion. Propulsion during flight differs
from that observed during stepping in terrestrial mammals. During
the wingbeat, pectoralis provides the major component of thrust,
both by adducting and pronating the wing. Although latissimus dorsi
is a major propulsive muscle during stepping in terrestrial mammals,
its major function in Artibeus is to abduct the wing and reposition
the wing prior to the beginning of a downstroke.
x

INTRODUCTION
Bat flight is energetically expensive (Thomas and Suthers, 1972;
Carpenter, 1975; Thomas, 1975, 1981) and requires extensive alterations
of the generalized mammalian morphology (Miller, 1907; Vaughan,
1970a). In order to understand how bats fly, one must both study their
anatomy as well as the subtle interactions between the nervous and
muscular systems of several bats must be studied. This study provides
information about the neuromuscular and osteological mechanisms
associated with flight in a frugivorous bat, Artibeus jamaicensis
(Microciroptera, Phyllostomidae). The objectives of the project are to
describe in detail the musculoskeletal system of Artibeus, to
illustrate the kinematics of the normal wingbeat cycle, and to obtain
electromyograms during flight for the major muscles of the shoulder
region and arm.
Three factors make Artibeus jamaicensis desirable for
morphological analysis. First, Artibeus is a ubiquitous neotropical
bat whose basic biology has been well documented. The available
information regarding the foraging and flight habits of this bat
permits deduction of the demands placed on the bat. Second, Artibeus
1

2
jamaicensis is easily maintained in captivity during the duration of an
experimental period (Rasweiler, 1977). Third, Artibeus is placed in
the family Phyllostomidae (Jones and Carter, 1976), a family that has
not been employed in previous EMG studies.
The diet of Artibeus includes fruits, flower products, and leaves
(Goodwin and Greenhall, 1961; Villa-R., 1967; Heithaus et al., 1975;
Gardner, 1977; Bonaccorso, 1979). Insectivory is insignificant in this
species even though Tuttle (1968) observed Artibeus jamaicensis in
Mexico feeding on blackflies within a cave. The bats usually transport
relatively heavy fruits from their source to adjacent feeding roosts
except when too large to carry efficiently (Morrison, 1978a;
Bonaccorso, 1979). Thus the wings of Artibeus need to be capable of
producing lift at low speeds without a great deal of maneuverability.
The flight style of Artibeus differs from that of Desmodus,
Antrozous, and Myotis, three bats previously studied with
electromyography. The vampire, Desmodus, is characterized by its
direct, swift flight (Altenbach, 1979). Antrozous and Myotis are both
insectivores but each employs slightly different foraging behaviors.
Antrozous is a long-eared bat and either gleans insects from foliage or
preys upon large, slow-moving insect species at low altitudes. Myotis
myotis tends to also be an opportunistic gleaner, searching for and
obtaining insects from the surface of foliage or the ground (Findley,
1972). Desmodus, Antrozous, and Myotis all forage for extended

3
periods of time during the night. Antrozous was reported to engage in
continuous foraging on the wing for one to four hours during early
evening and later for several hours before dawn (O'Shea and Vaughan,
1977). Myotis lucifugus similarly feeds for several hours during
early evening, and again for several shorter periods before dawn
(Anthony and Kunz, 1977). I expect behavioral studies will show Myotis
myotis has a similar time-energy budget. Turner (1975) reported that
Desmodus spends one to three hours foraging per night and found that
the bats spent a mean of one hour per night searching for their prey.
Artibeus, in contrast, tends to spend relatively little time in flight
during the night (Morrison, 1978a; 1978b). Artibeus also is not very
maneuverable and is easily captured in nets. There are two brief
flights between the day roost and the food source: one at dusk to the
food and one shortly before sunrise back to the day roost (Morrison,
1978a_). Morrison found that day roosts were never very distant from a
source of ripe fruit. At night, Artibeus selects fruit at the source
tree and then carries the fruit to an adjacent feeding roost, often in
another tree or in a cave (Dalquest, 1953; Morrison, 1978a; Bonaccorso,
1979). After chewing the fruit and swallowing juice and edible parts,
the bat ejects the remaining fruit pulp and seeds before returning to
the source tree for another fruit. These feeding flights are repeated
many times during the night.
Artibeus jamaicensis transports fruit on average 270 m from the
parental trees to feeding roosts (Janzen et al., 1976).

4
Morrison (1978a) observed that the distance between several Ficus
insípida and Ficus yaponensis, two species of figs important as food
trees, and the adjacent feeding roosts ranged from 25-400 m (mean 175
m). It is not clear why these bats carry fruits away from the parent
tree. Janzen et al. (1976) speculated that branch morphology and odor
in Andira inermis evolved to insure dispersal of the seeds away from
the parent tree. Seed mortality resulting from fungal infections and
weevil predation was most severe beneath the parent trees but declined
with distance away from the tree. They were unable to demonstrate a
clear cause and effect relationship, but did speculate that bats were
intolerant of the odor associated with many ripe Andira fruit. As an
alternative hypothesis, several authors suggested that Artibeus carried
fruit to a distinct feeding roost to decrease predation (Fenton and
Fleming, 1976; Morrison, 1978a; August, 1979). Artibeus avoid visually
orienting predators such as owls and opossums, by feeding only on dark
nights or during dark periods of the night (Morrison, 1978b). Specific
cases of predation by Didelphis marsupialis upon Artibeus were reported
in Venezuela (August, 1979). In any case, Artibeus make a number of
short flights during the night to and from a ripe fruit tree.
The neuromuscular control of flight can be correlated with similar
mechanisms in operation in the control of terrestrial locomotion.
Analysis of terrestrial involved resolution of the locomotor activity
of each limb into successive strides or step cycles, and quantification

5
of the interaction of these events between the limbs (gait analysis)
(Wetzel and Stuart, 1976). Philippson (1905) described four stages in
the step cycle of dogs, beginning with foot liftoff: F or flexion
phase is when most joint angles in the limbs decrease; El is the final
portion of the "swing" or non-support phase of the stride; E2 is the
first portion of the "stance" or support phase and includes some
passive joint flexion as a result of the limbs yielding under the
animal's weight; E3 is the final portion of the "stance” phase during
which all joints are actively being extended and maximum thrust is
being generated by the limb. Neurophysiologists studying vertebrate
locomotion have long focused on events surrounding the two reversal
points in the stride. For example, Sherrington (1910) made the
observation that a graded alternation of activities occurred in the
flexor and extensor muscles and that this alternation could be invoked
in a model of reflexive control of the step cycle (cf. Wetzel and
Stuart, 1976).
As an analogy, Eermanson and Altenbach (1981) expressed the
chiropteran wingbeat cycle as being a quantifiable and repeated unit of
normal bat flight. Individual wingbeats exhibit little variation among
conspecific individuals performing the same locomotory tasks under
similar conditions. The chiropteran downstroke, or adductor phase, is
similar in basic form and timing to the E2-E3 phases of the Philippson
step cycle and produces components of both lift and thrust (Norberg,
1976). The chiropteran upstroke, or abductor phase, parallels the F-El

Figure 1.—The wingbeat cycle of Artibeus jamaicensis. Each image was
traced from a 16 mm movie film. About 8 msec elapsed between each image.
The downstroke begins with frame B, and ends at frame H. Upstroke
movements begin during frame I and continue through frame M. The flick
phase, not shown, occurs rapidly during the interval between frames L
and M when the wing is rapidly pronated.

7

8
phases of overground locomotion in its function as a recovery stroke.
The last portion of the upstroke often includes a thrust-producing
"flick phase" (Norberg, 1970, 1976). Except for the flick phase, it is
not practical to subdivide the upstroke or downstroke because of the
smooth arc followed by the wing during each phase (Figure 1).
Engberg and Lundberg (1969) demonstrated that cat limb muscles
exhibit characteristic activity profiles during overground locomotion:
extensor or flexor muscles produce a cumulative effect on the skeletal
system through overlap or coactivation, and muscles that span two
joints produce a complex function including both flexion and
extension. Engberg and Lundberg reported extensor activity prior to
foot touchdown during normal stepping. This important observation
indicated that muscle activity could be preprogrammed at the spinal or
supraspinal level, taking precedence over reflexive control of the step
cycle (Wetzel and Stuart, 1976). Subsequent studies demonstrated that
the EMG profiles for homologous mammals are conservative among mammals,
with subtle differences in timing related to differing functional
demands being placed upon the musculoskeletal system in various species
(English, 1978a_, 19781^; Gambaryan, 1974; Rasmussen et al., 1978;
Tokuriki, 1973a, 1973b).
Hermanson and Altenbach (1981, 1983) assembled EMG data for muscle
activity during flight in an insectivorous bat, Antrozous pallidus
(Vespertilionidae). Of 17 muscles studied, 11 muscles demonstrated
patterns of either adductor or abductor activity. Only six muscles

9
exhibited a complex bifunctional pattern of activity. The adductor
muscles demonstrated an onset of activity on average 15-25 msec prior
to initiation of the downstrolce. This pattern is similar to the El
extensor coactivation observed prior to foot touchdown in cats (Engberg
and Lundberg, 1969) and suggests that similar neural control mechanisms
are involved in regulating both the terrestrial step cycle and the
aerial wingbeat cycle. Despite gross anatomical modification during
the evolution of the chiropteral musculoskeletal system, the patterns
of limb movement and muscle activity remained relatively unchanged from
the basic mammalian condition.
This study presents the descriptive anatomy of the locomotor
system of A. jamaicensis, including the shoulder girdle, arm, and
forelimb portions of the wing. Descriptions are accompanied by
functional hypotheses for each muscle following a format employed in a
number of studies on the mammalian musculoskeletal system (Howell,
1926; Rinker, 1954; Klingener, 1964; Woods, 1972). Additionally, EMG
data are presented for 15 of the 17 muscles that were previously
analyzed in Antrozous pallidus (Hermanson and Altenbach, 1983). These
data are compared with simialr EMG observations upon terrestrial
mammals.

MATERIALS AND METHODS
Ten skeletons and ten alcoholic specimens of Artibeus jamaicensis
were used for descriptive study. Dissection specimens were preserved
in a solution of 10 percent solution of formalin for 10-14 days and
were later stored in 70 percent ethanol or 40 percent isopropanol.
Complete dissections performed on the shoulder, arm, and antebrachial
regions form the basis of descriptions and illustrations presented in
this study and for the design of electromyography experiments.
Terminology used in my study conforms to recommendations by the
International Commission on Veterinary Anatomical Nomenclature (Nomina
Anatómica Veterinaria, 1975). Although this procedure initially
appears to pose difficulty in making comparisons with previous studies
on chiropteran morphology, relatively few new terms are introduced.
Strict adherence to the terminology of the Nomina Anatómica Veterinaria
(NAV) will provide a consistent vocabulary for effective communication
about the subject and will facilitate communication with researchers
who have not mastered the specialized jargon of bat anatomy.
Several terms are used throughout this text to indicate direction
along the body axis or direction of movement at a joint. The terms
10

11
cranial and caudal have replaced anterior and posterior, respectively,
in all cases where the latter terms have been used in the chiropteran
literature. Cranial refers to the cranial end of the animal, or
indicates that a structure passes from a point on the body along a line
towards the transverse plane of the head. Caudal indicates the
opposite direction to cranial: to or towards a transverse plane
containing the caudal vertebrae. Medial and lateral are used to
indicate position or movement towards or away from the medial plane of
the body. Flexion and extension refer to a decrease or increase,
respectively, in the angle between two bones. Adduction is used to
denote movements of the wings, humerus, or clavicle towards the medial
plane on the ventral surface of the body. Abduction of the wings,
humerus, or clavicle is a movement away from the median plane of the
ventral surface of the body. Rotation of a bone about its long axis is
described as supination, or outward rotation of the top or lateral
surface of the bone, and pronation, an inward rotation of the top or
lateral surface of the bone. To facilitate comparison with terrestrial
mammals, the arm and antebrachium are described in the position of
maximal adduction. Thus these bones have cranial, caudal, lateral, and
medial aspects. The carpus and digits are described as if they are
held horizontally at the side of the body. These elements have
cranial, caudal, dorsal, and ventral (palmar) aspects. The apparent
inconsistency in terms and orientation between the anatomical position
of the arm and carpus is in fact the traditional and accepted format
for descriptive purposes.

12
Skeletons were measured with dial calipers in order to describe
shape and size of individual bones. This information is useful in
estimating position or attachments of soft tissues with respect to the
skeleton.
Muscle origins and insertions were described relative to named
structures or surfaces of individual bones: the extent of muscle
attachment to long bones is expressed as a percentage or proportion of
the total length of the bones. Fiber orientation was expressed with
respect to the long axis of a muscle, along a line fit by eye between
the lines of attachment.
Synchronized film and EMG records were obtained with a system
initially described in Altenbach (1972, 1979) and modified by Bermanson
and Altenbach (1981). A Hycam high-speed cine camera (Red Lakes
Laboratories, Inc.) photographed flight sequences at film speeds of
300-400 frames per second. Myopotentials were recorded from a bipolar
fine copper wire electrode implanted in the muscle belly. Implantation
was achieved by a simple microsurgical procedure. An incision was
first made in the dorsal integument superficial to the lumbar region
and a silicone plug assembly was sutured to the muscle and ligaments of
the paravertebral region. Electrodes were passed subcutaneously to a
second incision at the level of the specific muscle being studied. A
shielded cable surrounding the electrode assembly served as a tissue
ground. With the aid of a microscope, the electrode tips were
inspected to insure than less than

13
0.5 mm of the tip was bared. The tips were then placed in the muscle
belly with a 27 guage hypodermic needle, and both incisions were
sutured shut with 6-0 Ethicon nylon suture. Methoxyfluorane was used
as a short term anesthetic agent with excellent results. Data were
recorded shortly after the bat exhibited normal locomotor behavior and
within three hours of surgery. Extended delays between the time of
implantation and recording sessions resulted in less than satisfactory
results, apparently as a result of toxic reactions between the copper
electrode material and adjacent tissues. Inspection of one
implantation that had been in place for 24 hours showed significant
tissue necrosis in the muscle for a 2 mm diameter around the electrode
tip. Only one muscle was studied per experiment. Each implant was
visually inspected at the conclusion of each recording session to
verify that the electrode remained in place. Each animal was allowed
to recover for at least one week before being used in further
experiments.
Normal flight was studied with single frame analysis of films of
normal horizontal flight sequences. Single film frame images were
projected with an L and W motion analyzer (Model 224A) and were traced
to ascertain wingtip motion and joint position in the wing. Vertical
displacement of the wingtips was used to determine the turnover points
of adduction and abduction. The beginning of a downstroke was treated
as the beginning of a wingbeat cycle and was assigned a numerical value
of 0.000 (Figure 1, frame B). A complete wing cycle included values
between 0.000 and 1.00, with 1.00 representing the final frame of an

14
upstroke (Figure 1, frame M). The transition between downstroke and
upstroke (Figure 1, frame H) and periods of muscle activity are
expressed as a mean percentage of a complete cycle.
Estimation of aerodynamic parameters was accomplished by measuring
the dimensions of the wing and body of two fresh specimens of Artibeus
jamaicensis. The mass of each bat was obtained to the nearest 0.5 g.
Wing area was estimated by drawing the outline of the body and fully
outstretched wing of each bat on paper. The cut-out outline of the
airfoil was then weighed to the nearest 0.0005 g and then compared to
the mass of a .100 m^ square of the same paper to calculate actual
wing area. Wingspan was measured along a line between the two
wingtips. Aspect ratio is the ratio of length to width of the
airfoil. The equation for aspect ratio is b^/A (Norberg, 1981).
Wingloading is the ratio of body weight to wing area, W/A (Norberg,
1981).

FLIGHT AND AERODYNAMICS
Results
The temporal relationship between the downstroke and upstroke is
critical in the interpretation of EMG data. In part, this relationship
reflects whether or not the activity of a given muscle is coincident
with or occurs before or after a particular event, such as the
beginning of a downstroke. During all filming sessions, flight was
observed to detect any grossly abnormal flight behaviors. All flight
sequences exhibiting turning behavior or altitude loss were eliminated
from consideration. The wingbeat cycle of Artibeus appeared to be
conservative during slow horizontal flight, based upon unaided visual
observation as well as initial study of the film records. However,
analysis of the film revealed slight differences in the flight of
restrained and unrestrained animals.
During unrestrained flight in two bats, the downstroke encompassed
55.1 percent of the wingbeat. In contrast, bats that carried an
electrode and plug assembly exhibited a mean downstroke of 49.44
percent. During unrestrained locomotion, the bats averaged 10.08
wingbeats per second. Records analyzed from four EMG experiments
(clavotrapezius, spinotrapezius, clavodeltoideus, and infraspinatus)
demonstrated a mean wingbeat frequency of 10.45 wingbeats per second.
15

Figure 2.—Wing and body movements during slow forward flight in
Artibeus jamaicensis. The bat flew from left to right at 2.19
m/sec. The movements of the wingtip ( ), wrist (x), foot (o),
and ear (•) were plotted during two wingbeats. The interval
between each symbol was about 8.30 msec. There were two periods
when the ear was obscured by the position of the wing during a
wingbeat cycle.

17

Figure 3.—Wing and body movements during slow forward flight in
Artibeus jamaicensis. The bat flew from left to right at 2.85
m/sec. The movements of the wingtip ( ), wrist (x), foot (o),
and ear (•) were plotted during two wingbeats. The interval
between each symbol was approximately 5.10 msec.

19

Figure 4.—Wing and body movements plotted relative to still
air during slow forward flight in Artibeus jamaicensis. The
bat flew from left to right at a velocity of 2.15 m/sec.
(A) The movements of the wingtip are plotted relative to the
body. (B) The movement of the wrist is plotted relative to
the body.

21
A
B
5 cm

22
Table 1. Aerodynamic parameters of the wing in two fruit bats,
Artibeus jamaicensis. All measurements are recorded in
meter-kilogram-second units.
mass
0.04 kg
0.03 kg
weight
0.40 N
0.32 N
wing span (b)
0.39 m
0.32 m
wing area (A)
0.02 m2
0.02 m2
aspect ratio (b2/A)
6.79
6.55
wing loading (W/A)
16.74 N/m2
15.47 N/m2

23
Analysis of one sequence of unrestrained flight is provided to
illustrate airspeed and movement of several parts of the body and wing
(Figures 2-4). Plots of head movement exhibited forward velocity
ranging from 2.19-2.85 m/sec. At the beginning of the downstroke, the
W’ingtip was positioned above and caudal to the center of the body. The
w7ing was adducted, but moved with an initial caudal sweep of the
wingtip relative to the center of the body. After approximately 15
msec, the adduction of the wingtip exhibited an increasing cranial
component of movement. The wrist moved in synchrony with the wingtip
but travelled through a smaller excursion. The wingtip continued
moving ventrally and cranially through 62.5 percent of the wingbeat
(Figure 2, Symbols 2-12). The wingtip and wrist synchronously showed
abduction at the beginning of the upstroke. There was a cranial
movement of both elements for approximately 5 msec. Subsequently,
abduction of the wingtip and the wrist included a caudal component of
movement for about 26 msec until the beginning of the flick phase.
During the flick phase (Figure 2, Symbols 18-21) the outer portion of
the chiropatagium was rapidly pronated. Cranial movement of the
wingtip occurred during the flick phase, which lasted approximately 5
msec, or until the beginning of the downstroke. During the entire
upstroke, movement of the wrist was in a dorsal and caudal direction.
Thus, during the wingbeat cycle the wrist followed an elongated
elliptical path. The path of the wingtip was best characterized as a
tight and complex figure eight when viewed laterally.

24
Several measures are used in the description and comparison of the
aerodynamic properties of Artibeus jamaicensis. Based on two specimens
obtained in Haiti, the following means were obtained for these
parameters: mean wing area, inclusive of the body, 0.02 m^;
wingspan, 0.38 m; aspect ratio, 6.67; wingloading, 16.11 N/m^.
Discussion
The wing and flight characteristics of Artibeus jamaicensis are
generalized in comparison to many other bats. Analysis of the flight
kinematics and wingshape demonstrates that Artibeus is capable of slow
flight in the forest canopy but not the faster, more maneuverable
flight associated with the pursuit of insects among dense vegetation or
in open airspace.
The data obtained during EMG experiments on Artibeus represent a
departure from the mean downstroke times observed during the
unrestrained flight sequences. These EMG flight data are comparable to
downstroke phase relationships reported during EMG studies in Antrozous
where a range of 39.7 to 52.3 percent was reported for the downstroke
period (Hermanson and Altenbach, 1983). The downstroke of flying
animals lasts longer than the accompanying upstroke (Aymar, 1935;
Brown, 1963; Norberg, 1976; Brandon, 1979). Flying birds or bats can
be seen to oscillate in a vertical plane during flight, losing a slight
amount of altitude during each upstroke phase, and regaining altitude
during the subsequent downstroke (Eisentraut, 1936). The upstroke does

25
not provide sufficient lift to counteract the effect of gravity except
in certain species that employ a hovering mode of flight (Norberg,
1975).
The flight speeds observed in the present study are similar to
those observed in Plecotus auritus by Norberg (1976). In her study,
the bats were filmed while flying unrestrained within a room and flight
speeds ranged between 2 a 3 m/sec. Higher velocities were exhibited by
Noctilio albiventris (3.51-10.38 m/sec) and Tadarida brasiliensis
(4.33-9.AO m/sec) while flying in a wind tunnel (Brandon, 1979).
Leptonycteris sanborni, a facultative hovering bat, flew between 2.0
and 6.0 m/sec in a wind tunnel study. The higher speeds reported in
the wind tunnel studies were not duplicated in the present study.
Predictable flight, suitable for EMG analysis, was facilitated by
filming the bats shortly after takeoff and before they reached maximum
velocities. Also, it is not known if Artibeus is capable of flying up
to 4.47 m/sec as was generalized for many bats (Hayward and Davis,
1964).
The path of the wingtip during the wingbeat cycle changes with
increasing airspeed. During slow flight in Plecotus auritus, the
wingtip path was directed upwards and caudally relative to still air
during the upstroke (Norberg, 1976). At higher velocities (2-2.5
m/sec), the upstroke of Plecotus was directed almost vertically, and at
higher velocities (greater than 3.0 m/sec) exhibited a cranial and
dorsal movement relative to still air. The same tendency was present
in Artibeus flying between 2-2.85 m/sec. The wing was directed

26
caudally during the upstroke at low speeds (Figure 2) and more
vertically during the upstroke at higher velocities (Figure 3). If
wing movements are different at several airspeeds, these relationships
must be considered when evaluating EMG recordings. The difference in
wing movements may be correlated with differences in muscular activity.
Flying animals have been classified according to several
descriptors of wingshape including the aspect ratio (Saville, 1962).
Vaughan (1959) discussed the relationship between aspect ratio and
foraging habits. Struhsaker (1961) was the first author to describe
the aspect ratios of a diversity of chiropteran wings. Aspect ratios
were also provided for bats by Farney and Fleharty (1969), Findley et
al. (1972), Lawlor (1973), Smith and Starrett (1979), and Norberg
(1981). My calculation of an aspect ratio for Artibeus jamaicensis,
6.67, is slightly greater than but comparable to the value of 6.36
obtained by Norberg (1981). In general, bats with low aspect ratios
are common in heavily forested habitat, whereas high aspect ratio bats
are commonly associated with flight in open, uncluttered airspace
(Findley et al., 1972). For example, Vaughan (1959) compared the
high-altitude flight of Eumops perotis with that of swifts. These bats
fed high over the forest canopy at high airspeeds. Vaughan reported
that the aspect ratio of Eumops was 11.9, the highest value among the
three species that he studied. Intermediate aspect ratio wings are not
well correlated with any particular behavioral repertoire or foraging
environment (Norberg, 1981). Wing shape in Artibeus probably evolved

Figure 5.—High- and low-aspect ratio wingshapes. Tadarida
brasiliensis (Molossidae) exemplifies the long and narrow
outline of a high aspect ratio wing. In contrast, the wing
of Artibeus jamaicensis exhibits a lower aspect ratio,
characterized by a broad and stubby appearance.

28
Tadarida
Artibeus

29
in response to the needs of the bat in carrying large food items and
transporting infants. Artibeus females have been observed carrying
young weighing 10.0 g (personal observation). Low aspect ratio wings
promote increased lift at lower flight speeds (Findley et al., 1972),
permitting more careful scrutiny of potential food or of potential
feeding sites. Maneuverability does not appear to be characteristic of
the flight pattern of Artibeus in comparison with some more agile
bats. This was supported by repeated netting success while trying to
collect bats in mist nets (personal observation). In contrast, other
phyllostomids such as Micronycteris have low aspect ratio wings (Smith
and Starrett, 1979) and were frequently observed eluding my nets during
field work in Venezuela.
Wing loading is an estimate of the relationship between the weight
of a bat and the lift-producing potential of a wing. To fly slowly,
which would be adaptive to an Artibeus investigating a fig tree as a
potential food source, two strategies might be employed: the species
might exhibit low body weight or a greater airfoil surface area
(Findley et al., 1972). My data for wing loading in Artibeus were
slightly less than the 16.65 N/m^ reported by Norberg and greater
than the 15.94 N/m^ reported by Smith and Starrett (1979). Norberg
found that the wingloading of frugivorous microbats (this included most
phyllostomid fruit eaters) was intermediate between the insectivorous
Molossidae and Vespertilionidae. The molossids of equivalent weight
have high aspect ratios and must maintain high airspeeds to avoid a

30
stall (Vaughan, 1966; Norberg, 1981). In contrast, the vespertilionids
generally have a large wing area for a given body size (Norberg, 1981)
and fly at relatively lower airspeeds than similar-sized molossids
(Hayward and Davis, 1964; Brandon, 1979). Thus, the wing of Artibeus
represents a compromise between the two strategies suggested by Findley
et al. (1972). The bats have large body weights but have not
substantially increased wing surface area like vespertilionids have.
This combination leads to reduced maneuverability, different from the
butterfly-like flight observed in some vespertilionids. Also, I have
never observed Artibeus hovering. The bats tend to fly directly along
aerial pathways to and from a feeding tree, and fly past a tree several
times when investigating or searching for food.
In conclusion, the wing of Artibeus represents a generalized
condition relative to certain other bats. I suggest that this reflects
the heterogeneous nature of the airspace through which the bat must
travel every night. A high aspect ratio wing is not suited to travel
through the cluttered environment of most neotropical forests. Also,
at slow airspeeds, the larger lift production that accrues from a low
aspect ratio wing permits the bat to forage upon and to transport large
energy-rich foods. A secondary consequence of the large body mass and
low aspect ratio of Artibeus may relate to post-natal development and
maternal care of the young bats.

OSTEOLOGY
Results
Clavicle
The clavicle in five specimens had a mean length of 16.10 mm and a
mean diameter at the midshaft of 1.15 mm. The bone articulates
proximally with the manubrium by a synovial joint. Distally, the
clavicle articulates with the scapula both by a synovial joint at the
base of the coracoid process and by a ligamentous connection with the
tip of the acromian process. Viewed cranially, the bone is straight,
with slight lateral curvature near its distal end. Viewed laterally,
the bone has a slight caudal curvature at the distal end. The distal
one-third of the cranial border of the bone is expanded for insertion
of part of the subclavius muscle. In lateral view the distal end of
the bone lies cranial relative to the proximal end.
Scapula
The scapula is a thin, relatively flat bone positioned over the
dorsal surface of the cranial part of the ribcage. It articulates
proximally with the clavicle as described above. The scapula
31

Figure 6.—Lateral view of the thoracic and axillary skeleton. UF 16265.

rib 10
Lo
Lo

Figure 7.—Cranial view of the right pectoral girdle.
UF 16265.

35
craniomedial flange
acromion process
/
/
\ ✓
5 MM

Figure 8.—Dorsal view of the scapula. UF 16265.

acromion process
scapular
acromiom process
supraspinous
fossa
scapular spine
cartilaginous
exspansion
%
\
notch ,
\
caudal angle
glenoid fossa
infraspinous fossa:
'' craniomedial facet
-- caudolateral facet
5 mm
U)

38
articulates with the humerus at the glenoid fossa, which forms the
shoulder joint. The dorsal surface of the scapula is divided into two
major subdivisions by the scapular spine. Cranially, the supraspinous
fossa forms a small flat shelf immediately cranial to the scapular
spine. Caudal to the scapular spine is the infraspinous fossa which
is, in cranial to caudal sequence, divided into three subdivisions:
craniomedial facet, intermediate facet, and caudolateral facet. The
ventral surface of the scapula is the subscapular fossa, composed of
two concave facets. The cranial subscapular facet corresponds with the
contours of the combined supraspinous fossa and craniomedial facet of
the dorsum. The caudal facet of the subscapular fossa corresponds with
the contours of the combined intermediate and caudolateral facets of
the dorsum. There are two borders on the scapula. Medially, the
vertebral border lies in a sagittal plane extending from the caudal
angle to the scapular spine. Cranial to the scapular spine, the
vertebral border curves laterally and becomes continuous with the
craniomedial flange of the scapula. On the lateral aspect of the
scapula, the axillary border is convex, coursing cranially from the
caudal angle to the glenoid fossa. The infraglenoid tubercle is a
small projection from the axillary border 1-5 mm caudal to the glenoid
fossa. The supraglenoid fossa is slightly concave and lies between the
glenoid fossa and the dorsal base of the coracoid process. This
surface abuts with the greater tubercle of the humerus when the wing is
fully abducted.

39
Several processes are evident on the scapula of Artibeus. The
cartilaginous expansion is triangular and projects from the caudal
angle and curves ventrally. The acromion process projects cranially
and dorsally from the scaular spine, and curves cranially. A 2 mm long
flared part of the acromion process lies dorsal to the clavoscapular
articulation. The coracoid process extends cranially and ventrally
from the supraglenoid region of the scapula and curves laterally to
terminate 2 mm ventral to the glenoid fossa.
Humerus
The humerus averaged 31.90 mm in length (measured between the
articular surfaces) and 1.98 mm in diameter (along the craniocaudal
axis) at the midshaft. In cranial view the distal end of the shaft
curves slightly lateral. The shaft is unmarked. Both ends of the
humerus are modified to form articular surfaces and to accommodate
muscle attachments.
The shoulder joint is formed in part by the head of the humerus.
The head is a convex elliptical surface that articulates proximally
with the glenoid fossa. The greater tubercle lies lateral to the head
and projects 1 mm proximally. The greater tubercle provides attachment
for the suprascapular muscles and abuts with the supraglenoid fossa of
the scapular when the humerus is fully abducted. This lock occurs when
the humerus is abducted about 32 degrees dorsal to the horizontal plane
of the scapula. The lesser tuberosity also extends slightly proximal
to and lies on the medial aspect of the head. Several ridges extend

Figure 9.—Medial view of the left humerus. UF 16265.

41
5 MM

Figure 10.—Articular surfaces of the humerus. (A) Caudal
aspect of the proximal left humerus. (B) Cranial aspect of
the distal left humerus. UF 16265.

43
greater tubero
lateral ridge
A
head
e \
\ r
B
idge
ndyle
medial process
trochlea

44
distally from the lesser and greater tuberosities. From the lesser
tuberosity, the medial ridge extends 6 mm along the proximal medial
border of the humerus and serves as the point of attachment of the
teres major and latissimus dorsi muscles. A second indistinct ridge
crosses the proximal surface of the humerus and courses along the base
of the ventral surface of the pectoral ridge. This ridge is the site
of insertion of the pectoralis abdominalis. The lateral ridge is a
small indistinct ridge on the lateral surface of tlie humerus courses 7
mm distal to the greater tuberosity. This forms the dorsolateral
border of the origin of triceps brachii, lateral head. The most
prominent ridge from the greater tuberosity is continuous distally with
the pectoral ridge, a large flat projection on the lateral aspect of
the humerus. The pectoral ridge extends 8 mm along the lateral edge of
the humerus. The widest portion of the pectoral ridge, 2-4 mm distal
to the head, is termed the apex. The pectoral ridge provides insertion
sites for the pectoralis muscles medially, and the deltoid muscles
laterally.
At the elbow the radius articulates with the cranial surface of
the humerus. The ulna articulates along the distal and caudal surface
of the humerus. The capitulum and trochlea are fused cranially to
present a synovial surface for the humeroradial articulation. The
trochlea extends around the distal and caudal surface of the humerus to
provide an articular surface for the ulna. Medial to the trochlea and
composing one-third of the distal end of the bones is a medial

45
process. This region provides an attachment surface for several flexor
muscles of the antebrachium. On the lateral end of the capitulum is
the spool-shaped lateral epicondyle. Extending from the lateral
epicondyle proximally onto the shaft of the humerus is the short
epicondyle ridge. The lateral epicondyle and epicondylar ridge provide
an area of attachment for several antebrachial extensor muscles. The
elbow articulation is not a pure hinge joint. Slight rotation of the
radius relative to the capitulum is evident in fresh specimens of
Artibeus.
Radius
The radius is slightly curved along its length. In five
specimens, the mean length is 53 mm. At the midshaft, it is thickest
in the dorsoventral axis (1.8 mm) to resist bending forces associated
with the wingbeat cycle. The proximal articular region is thickened to
provide a surface for articulation with the humerus and to provide a
strong site for attachment of the biceps brachii. The distal articular
region is wide and flat. This provides a structural mechanism for
separation of the tendons of insertion of the antebrachial muscles and
increases the mechanical advantage of several of these muscles as they
cross the carpus.
The proximal articular surfaces is oriented cranially and includes
a large concave fossa for articulation with the capitulum. Laterally a
smaller elliptical concavity articulates with the trochlea. A ridge on
the caudal aspect of the radius courses 5 mm distal from the elbow

46
joint. A ridge on the medial aspect of the proximal radius courses 3
mm distal to the elbow joint and provides the site for insertion of the
biceps brachii.
The styloid process and pseudostyloid process are small bumps on
the distal articular end of the radius but are not greatly developed in
bats. The pseudostyloid process is on the medial side of the distal
articular surface, analogous in position to the styloid process of the
ulna in other mammals.
The axis of the distal articular surface is laterally rotated
about 20 degrees from the orientation of the proximal articular
surface. The distal articular surface presents a concave surface for
articulation with the proximal carpal bones. When the wing is
laterally outstretched, the radiocarpal joint permits flexion and
extension in a mediolateral plane and no movement in the dorsoventral
plane. This restriction braces the joint against displacement caused
by the force of the airstream.
Ulna
The ulna is reduced in bats to a length approximately one-half the
length of the adjacent radius. The proximal articular surface
comprises the cranial surface of the olecranon process and articulates
with the caudal extension of the trochlea of the humerus. An
interosseous membrane is found between the radius and ulna. The ulna
tapers towards its distal articulation with the radius and presents no
processes.

Figure 11.—Dorsal view of the radius and ulna.

48
o 1 c r a n o n
process
10 mm

49
Manus
The diminutive size of and the tightly bound joint cavities
between the bones of the carpus make it difficult to adequately
describe the bones of this region. A perspective on the carpal anatomy
can be obtained by studying Figure 12. The carpus of Artibeus includes
eight bones as is typical of many mammals.
The proximal row of carpal bones includes the scaphoid, lunar, and
triquetrum. The largest carpal bone, the lunar, articulates proximally
with the radius and distally with the trapezoid, trapezium, and
capitate. On the lateral aspect of the distal articular surface of the
radius, the triquetrum («cuneiform of Vaughan, 1959) is bound to the
radius and forms the caudoventral border of the canal traversed by the
antebrachial extensor muscles. The scaphoid is a small bone located
craniodorsally in the carpus along the cranial surface of the lunar.
The scaphoid, unlike the lunar and triquetrum, is not bound to the
distal surface of the radius.
The distal row of carpal elements includes the trapezium,
trapezoid, capitate, hamate, and pisiform. The trapezium articulates
with the proximal bases of metacarpals I and II, and with the
craniodistal surface of the lunar. The trapezoid is compressed
craniocaudally and articulates with the proximal base of metacarpal
II. The capitate («magnum of Vaughan, 1959) is a large bone that has
an hourglass shape when viewed dorsally. It articulates with the
proximal bases of metacarpals II and III. Lateral to the capitate, the

Figure 12.—Dorsal and ventral views of the carpus.

51
trapezium
5 mm

52
hamate (=unciform of Vaughan, 1959) articulates distally with the bases
of metacarpals IV and V, and proximally with the lunar and triquetrum.
The pisiform is a small rod-shaped bone on the palmar surface of the
carpus and is firmly bound to the ventral surface of the capitate and
hamate.
The distal part of the wing is supported by digits II through V.
The metacarpals diverge within the wing membrane and are the largest
elements in each digit. Movement at the carpometacarpal joints of
digits II through V is restricted to flexion and extension. Flexion of
the carpometacarpal joints brings the digits parallel to and into
juxtaposition with the radius. The metacarpophalangeal and
interphalangeal joints permit only palmar flexion or dorsiflexion.
These movements adjust the camber of the wing during the wingbeat
cycle. Digit II has one phalanx. Digits III through V have three
phalanges. Digit I, the pollex, is unique because of the range of
motion possible at the carpometacarpal joint. Besides flexion and
extension, the pollical carpometacarpal joint permits the extensive
rotation necessary for use of the pollex in several functions: the
pollex is a prehensile organ used during landing maneuvers or while
manipulating food. The second phalanx of the pollex bears a claw. The
proximal and second phalanges of the pollex are both free of the
leading edge of the dactylopatagium minus and the propatagium.

MYOLOGY
Trapezius Group
Clavotrapezius and Acromiotrapezius
Form. The clavotrapezius originates by a fibrous attachment on
the dorsal surface of the seventh cervical vertebrae. Fibers of
acromiotrapezius arise from the dorsal surfaces of thoracic vertebrae
one through six. There is no apparent separation between adjacent
fibers of the clavotrapezius and acromiotrapezius. Insertion of the
clavotrapezius is by fibers on the distal 2.5 mm of the craniomedial
surface of the clavicle. The acromiotrapezius inserts separately from
the clavotrapezius: insertion is by fibers convergent upon the
anterior and dorsal surface of the acromion process and on a muscular
raphe caudal to the acromion process. Innervation of both muscles is
by cranial nerve XI (spinal accessory nerve).
Comparative aspects. The clavotrapezius and acromiotrapezius
muscles have been treated as two separate muscles in the Phyllostomidae
(Macalister, 1872; Walton, 1967), and also in the specialized vampire
bats (Altenbach, 1979). In contrast, the condition observed in many
other bats, including phyllostomatids, demonstrated fusion of these two
bellies (Strickler, 1978; Vaughan, 1959, 1970a). In the three species
he studied, Vaughan (1959) noted the fusion of the two bellies only in
53

54
Macrotus (Phyllostomidae). Macalister (1872; p. 138) most likely made
reference to the clavotrapezius of Artibeus sp. when he described "a
semi-detached upper slip passed from the two lowermost cervical spines
to the outer fifth of the clavicle.” The two muscles are
distinguishable in Artibeus only by a fascial division coursing from
the C7-T1 articulation to the distal end of the clavicle. Despite
their separate insertion, I consider the two muscles to be fused and to
share a similar function. The clavotrapezius and acromiotrapezius form
a flat sheet of parallel fibers, thickest at the cranial end
(clavotrapezius portion).
Functional aspects. The clavotrapezius and acromiotrapezius
exhibit a similar pattern of EMG activity during the wingbeat cycle.
Clavotrapezius has a major period of EMG activity during the transition
between the late downstroke and early upstroke. A second period of
activity occurs during the middle to late upstroke phase, but the
myopotentials have less than one-half of the amplitude observed during
the earlier EMG burst. Acromiotrapezius also has two periods of EMG
activity, but both exhibit similar amplitude and frequency. The first
period of activity in acromiotrapezius precedes clavotrapezius activity
by approximately 10 m/sec. Both muscles are anatomically positioned to
effect the upstroke by drawing the scapula medially (Vaughan, 1959;
Strickler, 1978; Altenbach, 1979). The primary activity of each muscle
contributes to this upstroke function. The secondary burst of activity
possibly stabilizes the scapula in response to the adductor activity of

Figure 13.—Dorsal view of the shoulder and arm of Artibeus jamaicensis.
Superficial muscles are exposed on the left, and deeper muscles of the
back and shoulder are exposed on the right.

occipitopollicalis
acromiotrapezius
spinodeltoideus
acromlodeltoideus
biceps brachii
teres major
latissimus dorsi
spinotrapezius
lavotrapezius
supraspinatus
clavodeltoideus
1
\ triceps brachii:
..-1
lat. head
long head
/
\
>
N
\
\
%
, ' infraspinatus
rhomboideus
Ln
On

Figure 14.—Activity patterns of shoulder and arm muscles in Artibeus
jamaicensis during slow flight. Black bars indicate observed periods
of high-amplitude activity. Unshaded bars indicate observed periods
of muscle activity that were consistently one-half of the amplitude
exhibited in high-amplitude sequences. Vertical lines are placed one
standard deviation from the means of activity onset and termination.
Small arrows indicate mean downstroke duration for each muscle
experiment.

Abductors
clavotrapezius
acromiotrapezius
latissimus dorsi
teres major
acromiodelt. (eran.)
acromiodelt. (caud.)
spinodeltoideus
triceps brachii (lat.)
triceps brachii (long.)
Adductors
clavodeltoideus
serratus ventralis thoracis
pectoralis
Bifunctional
spinotrapezius
supraspinatus
subscapularis
infraspinatus
l
downstroke
t
up
stroke J,
I 'I r r
'i r
IH
D-
-c
< •—r
>-
ID—1
IV—
h
É
fgXZSh
HZ8—1
KZZZH
h-CD—
SvTTT'
HD
rfPUftMff
t i . .i * i
b-raj
.00
.20
.40
.60
.80
1.0
Ln
00

59
Table 2.—Electromyographic data for activity patterns of the shoulder
musculature in Artibeus jamaicensis during slow flight. N= number of
wingbeats analyzed per muscle. Mean duration of the downstroke is
expressed as a percentage of the wingbeat cycle duration. Mean times
of muscle activity onset and termination are recorded in relation to
the wingbeat cycle from 0.00 to 1.00, and one standard deviation is
recorded.
Muscle
N
Downstroke
Duration
Onset
Time (S.D.)
Termination
Time (S.D.)
ABDUCTORS
Clavotrapezius
12
0.515
0.332
0.894
(.099)
(.030)
0.631
0.092
(.045)
(.047)
Acromiotrapezius
9
0.477
0.236
0.794
(.033)
(.080)
0.572
0.903
(.104)
(.104)
Latissimus dorsi
13
0.428
0.329
0.869
(.028)
(.035)
0.644
0.098
(.025)
(.025)
Teres major
Acromiodeltoideus
3
0.412
0.259
0.922
(.022)
(.046)
0.519
0.077
(.016)
(.045)
cranial
10
0.536
0.355
0.799
(.056)
(.041)
0.628
0.081
(.040)
(.055)
caudal
10
0.532
0.315
0.894
(.053)
(.051)
0.556
0.103
(.062)
(.065)
Spinodeltoideus
11
0.481
0.309
0.001
(.030)
(.115)
0.608
0.114
(.030)
(.080)
Triceps brachii
lateral
3
0.594
0.564
(.017)
0.741
(.053)
long
15
0.513
0.429
(.069)
0.714
(.074)

60
Table 2.—continued
Muscle
N
Downstroke
Duration
Onset
Time (S.D.)
Termination
Time (S.D.)
ADDUCTORS
Clavodeltoideus
19
0.544
0.853
0.481
(.059)
(.038)
0.135
0.617
(.027)
(.039)
Serratus ventralis
11
0.484
0.823
0.515
(.036)
(.058)
0.091
0.599
(.058)
(.109)
Pectoralis
7
0.526
0.781
(.027)
0.094
(.086)
BIFUNCTICNAL
Spinotrapezius
7
0.452
0.254
0.784
(.031)
(.020)
0.672
0.123
(.032)
(.047)
Supraspinatus
6
0.518
0.706
(.263)
0.142
(.084)
Infraspinatus
7
0.449
0.346
0.942
(.039)
(.059)
0.607
0.120
(.046)
(.039)
Subscapularis
10
0.450
0.420
(.060)
0.174
(.033)

61
the early downstroke, or causes active lengthening of the pectoralis
fibers during their early period of activity. Similar passive
lengthening, or "eccentric contractions," increased force production of
several locomotory muscles in other animals (Cavagna et al., 1977;
Heglund et al., 1979; Goslow et al., 1981). The biphasic activity
observed in Artibeus contrasts with the single EMG burst observed in
Antrozous (Hermanson and Altenbach, 1983). The heavier wingloaaing of
Artibeus may necessitate the extra muscular activity. In conclusion,
both clavotrapezius and acromiotrapezius show abductor activity, but at
slightly different times in the wingbeat cycle. The secondary activity
in each muscle stabilizes the scapula against the force of the
adductors.
Spinotrapezius
Form. The spinotrapezius originated by fibers on the dorsal
midline over lumbar vertebrae two through three. It inserts by an
aponeurosis on the vertebral border of the scapula, extending from the
caudal angle to the point 2 mm anterior to the scapular spine. The
spinotrapezius is a thin, parallel-fibered muscle. Based upon its
anatomy, the muscle tips the vertebral border of the scapula ventrally,
and retracts the scapula (Strickler, 1978). Innervation is by the
spinal accessory nerve.
Functional aspects. Spinotrapezius exhibits biphasic EhG
activity. Both periods of activity have approximately equal amplitude
and frequency characteristics. One period commences at 0.254 and

62
terminates at 0.672: This contributes to the overall abduction of the
wing along with the activity of clavotrapezius and acromiotrapezius. A
second period of activity commences at 0.784 and terminates at 0.123 of
the following wingbeat. Spinotrapezius is quiescent only during about
24 percent of the wingbeat, an observation similar to EMG data from
Desmodus (Altenbach, 1979), but in contrast to the shorter, single
activity period observed in Antrozous (Hermanson and Altenbach, 1983).
The extent of EMG activity throughout the wingbeat suggests a multiple
function for spinotrapezius as proposed by Vaughan (1959):
stabilization of the scapula and abduction of the wing and scapula are
likely functions.
Costo-spino-scapular Group
Serratus Ventralis Cervicis
The serratus ventralis cervicis was previously called the levator
scapulae in the chiropteran anatomy literature.
Form. The serratus ventralis cervicis originates from fibrous
attachments along the dorsolateral aspect of the transverse processes
of cervical vertebrae three through six. The muscle has a fleshy
insertion on the ventral surface of the craniomedial flange of the
scapula, and along the vertebral border of the scapula craniad of the
scapular spine. The muscle is composed of two parallel-fibered slips:
the cranial slip originated from cervical vertebrae three and four, and
the caudal slip from five and six. Innervation is by a branch of the
dorsal scapular nerve, and by cervical nerve five.

63
Comparative aspects. In many mammals, the serratus ventralis is
composed of a continuous sheet of muscle, including fibers that
originate from the cervical vertebrae (serratus ventralis cervicis) and
from the lateral surface of the ribcage (serratus ventralis thoracis).
The terminology and comparative anatomy are discussed with the serratus
ventralis thoracis.
Strickler (1978) noted that although the cranial origin normally
extends to cervical vertebra five to seven the origin of serratus
ventralis cervicis varies in bats. Phyllostomoids were characterized
by a relatively large insertion, usually extending along the vertebral
border of the scapula caudal to the scapular spine. Walton (1967)
observed a similar insertion in Artibeus lituratus extending along the
"coracoid border" and vertebral border of the scapula to a point
immediately posterior to the spine.
Walton (1967) described the serratus ventralis cervicis of A.
literatus to be composed of three large slips. Macalister observed
only two slips in Artibeus sp. (Macalister, 1872). I concur with
Macalister.
Functional aspects. Strickler (1978) concluded that a correlation
existed between degree of humeroscapular "locking" observed in some
bats and the size of the serratus ventralis cervicis muscle. He
observed that this muscle was relatively enlarged while the rhomboideus
was relatively small in vespertilionids and phyllostomatids. In
contrast, the serratus ventralis cervicis was small and the

64
rhomboideus large in emballonurids and pteropodids. These two groups
possess poorly developed humeroscapular locking mechanisms (Vaughan,
197(ta, 1970_b; Strickler, 1978). Strickler suggested that the major
role of serratus ventralis cervicis is to stabilize the medial border
of the scapula against rotation about its longitudinal axis. No EMG
data are available to test this hypothesis.
Serratus Ventralis Thoracis
Macalister (1872) observed and described the serratus magnus and
divided the muscle into superior and inferior portions. The same
muscle was described by Vaughan (1959) under the name serratus
anterior. He also divided the serratus anterior into two parts, the
anterior and posterior divisions. Subsequent authors followed his
precedent for bats (Walton, 1967; Norberg, 1970; Strickler, 1978;
Altenbach, 1979; Hermanson and Altenbach, 1981, 1983).
Serratus Ventralis Thoracis, Cranial Division
Form. The origin is a fibrous attachment on the dorsolateral
surface of the first rib, first intercoastal space, and cranial surface
of the second rib, medial to the origin of subclavius. The muscle has
a broad fleshy insertion along the ventral surface of the craniomedial
flange of the scapula, lateral to the insertion of serratus ventralis
cervicis. At the origin, the belly is small and rounded. Distally the
belly becomes wider and flatter, being thickest along the medial
border. The muscle is innervated by a branch of the dorsal scapular
nerve.

Figure 15.—Lateral view of serratus ventralis musculature.
In the upper figure, subclavius is intact. In the lower
figure, subclavius is removed to reveal the position of the
deeper serratus ventralis cervicis.

66
scapula
subciavius

67
Comparative aspects. The cranial division of serratus ventralis
thoracis is large in hovering bats and in slow-flying bats (Strickler,
1978).
Functional aspects. Vaughan (1959) proposed that the cranial
division functioned as a wing abductor by pulling ventrally on the
craniomedial edge of the scapula. The cranial division is large in
hovering bats, the glossophagines, and slow-flying bats, the
rhinolophids (Strickler, 1978). In both groups, the wingbeat amplitude
is large and may include extensive tipping of the lateral border of the
scapula ventrad during the downstroke. The cranial division is
situated to initiate abduction of the wing by arresting outward
rotation and by initiating inward rotation of the scapula. No EMG data
are available to test this hypothesis.
Serratus Ventralis Thoracis, Caudal Division
Form. The muscle originates by fibers along the dorsolateral
aspect of ribs two through 11, adjacent and dorsal to each
costochondral junction. A small, separate slip originates from the
caudolateral aspect of the first costal cartilage, caudal to the origin
of subclavius. The fibers of this small slip course caudally to insert
on the middle one-third of the axillary border of the scapula. The
fibers of the larger portion of the caudal division insert by fleshy
attachment along the caudal three-quarters to seven-eighths of the
axillary border and on the caudal angle of the scapula. The caudal
fibers inserting upon the caudal angle constitute the thickest portion

68
of the muscle. Fibers are arranged in parallel. The fibers
interdigitate at their origin with the scaleneus muscles cranially and
the external abdominal oblique muscles caudally. Innervation of the
caudal division is by the long thoracic nerve, coursing along the
lateral surface of the muscle.
Comparative aspects. Strickler (1978) found that the caudal
division is relatively large in most phyilostomids and small in the
Pteropidae and most of the Verpertilionidae. He felt that the function
and size of the muscle vary with respect to the development of the
humeroscapular locking mechanism.
Functional aspects. The shoulder locking mechanism described by
Vaughan (1959) provides a mechanical link between the action of the
caudal division upon the scapula and the resultant adduction of the
wing. Thus, contraction of the serratus ventralis thoracis was
hypothesized to power the downstroke. Electromyograms obtained from
Artibeus indicate that the posterior division has a major period of
activity that commenced before onset of the downstroke and ended during
the early downstroke. A second burst of activity is of shorter
duration and occurs during the early upstroke phase. Hermanson and
Altenbach (1981, 1983) reported a single period of activity for this
muscle in Antrozous during the transition between the upstroke and
downstroke. In both Antrozous and Artibeus, the pectoralis muscles are
active prior to the major burst of the caudal division of serratus
ventralis thoracis. The caudal division and pectoralis are coactive as
adductors throughout the early downstroke.

69
The second period of EMG activity occurs after the upstroke has
begun. This burst provides a force antagonistic to activity in
clavodeltoideus and latissimus dorsi. Clavodeltoideus activity tends
to protract the humerus, and thus advances the scapula craniad relative
to the thorax. Latissimus dorsi exhibits its most pronounced activity
as an abductor and pronator of the wing, causing retraction of the
scapula and arm.
English (1978a, 1978b) presented data for the serratus ventralis
cervicis and serratus ventralis thoracis of cats indicating biphasic
activity patterns. He concluded that serratus ventralis thoracis is
important in transmitting the weight of the body to the forelimbs
during the stance phase, an observation suggested by earlier anatomists
working only with cadaverous material (Davis, 1949; Gray, 1968). Of
greater interest was English's observation that coactivation of
cervical and thoracic portions of serratus ventralis during the swing
phase effected shoulder extension and repositioning of the limb for the
following step. Jenkins and Weijs (1979) found no significant
difference between EMG activity of different slips of serratus
ventralis thoracis in Didelphis. During walking, serratus ventralis of
Didelphis was active during the propulsive phase. These authors
commented upon a secondary period of activity during the swing phase in
Didelphis, but did not discuss its role. A secondary phase of activity
was not recorded in Antrozous (Hermanson and Altenbach, 1983).
Repositioning of the limb for the downstroke may be effected in

70
Antrozous during the upstroke by the force of the airstream on the wing
(Vaughan, 1959; Norberg, 1976) and by the action of the dorsal shoulder
musculature (Hermanson and Altenbach, 1983). The greater wingloading
in Artibeus may require an additional burst of muscle activity.
Rhomboideus
Form. The rhomboideus originates by fibers on the dorsal surface
of the transverse process of cervical vertebrae seven, lateral to the
cranial fibers of clavotrapezius, through thoracic vertebra six,
lateral to the proximal fibers of acromiotrapezius. The muscle fibers
course obliquely, caudodorsally through the interscapular region to
insert on the entire vertebral border of the scapula caudal to the
scapular spine. Innervation is by the dorsal scapular nerve.
Comparative aspects. The rhomboideus of the Phyllostomidae was
characterized by Strickler (1978) as being relatively small. In
contrast, both Macalister (1872) and Walton (1967) described the muscle
as robust in the Phyllostomidae. I concur with Strickler's
interpretation in part because of the muscle mass data presented in his
arguments, whereas the other authors relied upon subjective
interpretations. The romboideus is a thin, flat muscle in Artibeus.
Strickler (1978) hypothesized that either rhomboideus or levator
scapulae are well developed in bats depending on locomotor
requirements. In the Molossidae, he observed that the rhomboideus was
relatively large and adapted to rotating the caudal angle of the

71
scapula medially relative to an axis perpendicular to the dorsal
surface of the scapula. He speculated that this action is important in
terrestrial movements and not in the scapular movements associated with
the phyllostomid shoulder during flight.
Functional aspects. Based upon the attachments and orientation, I
deduce that rhomboideus pulls the caudal angle of the scapula
ventromedially. There are no EMG data with which to elaborate upon
this hypothesis.
Latissimus-subscapular Group
Latissimus dorsi
Form. The muscle originates from the superficial fascia overlying
thoracic vertebra eleven through lumbar vertebra four. At the origin,
the belly is thicker over thoracic vertebrae and grades into a thin
aponeurosis caudally over the lumbar region. Latissimus dorsi inserts
by a thin, 1 mm wide tendon 3-4 mm distal to the shoulder joint on the
medial ridge of the humerus. The tendon lies ventral to and on the
proximal edge of the fibrous insertion of teres major. Fibers in the
latissimus dorsi are arranged in parallel at the origin, but converge
distally on the tendon of insertion. Innervation is received from the
thoracodorsal nerve.
Comparative aspects. Strickler (1578) did not observe much
variation in the latissimus dorsi attachments of bats, except in the
Rhinolophidae. In several phyllostomid species studied, the origin

72
of latissimus dorsi ranged between thoracic vertebra 10 and lumbar
vertebra 5, but no correlation was established between the attachments
and taxonomy or locomotor modes. Strickler's muscle mass data
indicated that the muscle was largest in slow fliers, and average-sized
in hovering and moderately fast frugivorous bats.
Functional aspects. The action of latissimus dorsi for Artibeus
and for bats in general is confusing because of conflicting evidence
from several sources. Latissimus dorsi was described by Vaughan (1959)
as a pronator and flexor of the humerus, and also as an important
retractor during terrestrial locomotion (1970b). Strickler (1978)
added that the muscle might also serve in abduction of the wing during
upstroke movements. For Desmodus, Altenbach (1978) obtained EMG data
suggestive of a major role in humerus pronation and adduction, but not
in abduction. In Antrozous latissimus dorsi functions primarily as a
pronator during the downstroke (Hermanson and Altenbach, 1983).
The EMG data for latissimus dorsi in Artibeus are difficult to
interpret in comparison with the data obtained from other studies. In
17 wingbeats, latissimus dorsi exhibits a biphasic activity pattern,
with highest intensity EMG myopotentials observed during the late
downstroke to early upstroke transition period (onset mean was 0.329,
average onset of downstroke movements was 0.428). A second period of
activity, of shorter duration and with lower amplitude mypotentials,
occurs during the late upstroke and early downstroke. These patterns
are consistent in all wingbeats. These data contrast with the EMG

73
pattern reported for Antrozous (Hermanson and Altenbach, 1983).
Although the greatest activity for latissimus dorsi in Artibeus was
observed during the late downstroke, the muscle was quiescent in
Antrozous during the same period.
English (1978a_) observed variability in the EMG patterns of
latissimus dorsi in cats. Although the muscle was characterized by
English as monophasic and active predominently during the propulsive
phase (E1-E2), he observed inconsistent bursts of activity during the
swing phase (F). These inconsistent bursts are similar in timing to
the late downstroke activity observed in Artibeus. Jenkins and Weijs
(1979) reported monophasic activity in the posterior part of latissimus
dorsi of walking Didelphis, however, they noted a consistent biphasic
pattern for recordings made with electrodes located in the anterior
fibers of the muscle. Placement of electrodes in intermediate
positions resulted in "hybrid" EMG patterns: this region of latissimus
dorsi was monophasic during some step cycles and biphasic during
others. Tokuriki (1973a^ 1973b, 1974) observed two distinct phases of
activity in the latissimus dorsi of dogs at all gaits. One phase
occurred with other flexor muscles during the F phase, contributing to
lift the limb off the ground. A second and evidently more intense
period of activity occurred throughout the propulsive phase. In
terrestrial mammals, the activity of latissimus dorsi appears to be
biphasic, although the most intense activity occurs during a different
period of the locomotory cycle than observed in Artibeus.

74
Based on my EMG data and on comparative data for other mammals, I
conclude that latissimus dorsi in both Artibeus and Antrozous
stabilizes the shoulder and pronates the wing throughout the
downstroke. Primary EMG activity occurs during the transition from
downstroke to upstroke. Assuming a contraction time of 25 msec (Burke,
1978; Hermanson and Altenbach, 1981), the action of this muscle burst
pronates the wing during the first third of the upstroke. Analysis of
our film records indicates that the wing is continuously supinated
during the first two-thirds of the upstroke (Norberg, 1976; Altenbach,
1978), an action effected by the dorsal abductor muscles and by the
force of the airstream (Vaughan, 1959). The large burst of activity in
latissimus dorsi may therefore be necessary to counteract the
supinators and stabilize the shoulder during the early upstroke. The
temporal activity pattern of latissimus dorsi indicates that the muscle
is primarily a wing abductor.
Teres Major
Form. The teres major originates on the lateral and dorsal aspect
of the caudal angle of the scapula, and along the caudal 3 mm of the
axillary border of the scapula, by fibers. The origin does not extend
to the cartilaginous expansion at the caudal angle of the scapula. The
muscle inserts along the medial ridge of the humerus, 3-6 mm distal to
the shoulder joint. The insertion is primarily a fibrous attachment,
adjacent to but not conjoined with the tendon of insertion of the
latissimus dorsi. All myofibers are arranged in parallel. Innervation
is provided by several branches of the subscapular nerve.

75
Comparative aspects. Vaughan (1959) observed that the teres
major in Macrotus (Phyllostomidae) is larger relative to the latissimus
dorsi, than in Myotis (Vespertilionidae) and Eumops (Molossidae).
Strickler (1978) found that both the teres major and latissimus dorsi
exhibit positive allometry with respect to body mass. Triceps brachii
(long head) and biceps brachii (short head) were the only other muscles
in which Strickler observed positive allometry. He hypothesized that
whereas musculoskeletal specializations or lift provided by the
airstream permitted isometry to prevail in the primary adductors, and
abductors, respectively, the muscles associated with powering the flick
phase of the upstroke cannot be assisted by outside forces. Therefore,
a greater degree of positive allometry is exhibited in these four
muscles than in other muscles. Strickler did not include Artibeus in
his sample but did study other phyllostomids, including Carollia and
Phyllostomus, representative of locomotor styles broadly comparable to
Artibeus.
Functional aspects. Electromyographic data exhibit two phases of
activity. High-amplitude activity occurs during the late downstroke to
upstroke transition. A second burst of one-half amplitude activity
occurs during the late upstroke to downstroke transition. The
association of muscle activity with turnover periods of the wingbeat
cycle indicates these muscles play an important role in stabilizing the
shoulder joint. Previous investigators used dissection studies and
determined the role of teres major is responsible for pronation of the
wing and a flexion of the shoulder joint (Vaughan, 1959; Norberg,

76
1970, 1972; Strickler, 1978; Altenbach, 1979). Hermanson and Altenbach
(1983) reported EMG data for four wingbeats in Antrozous but made no
specific conclusions beyond the classification of teres major as a
bifunctional muscle. In Artibeus, the biphasic activity pattern is
observed but is significantly different from that observed in
Antrozous. The primary activity of teres major in Antrozous, during
the final stages of the downstroke, probably pronates the wing
synergistically with latissimus dorsi. Together, the two muscles
counteract the onset of supination effected by the abductor muscles and
airstream, and privide a smooth transition of wing movements between
the downstroke and upstroke.
Subscapularis
Form. The subscapularis fills the entire subscapular fossa. The
muscle originates along the periphery and on the elevated ridges of the
subscapular fossa, and from the ventral surface of the cartilaginous
expansion of the caudal angle of the scapula. The fibers exhibit a
multipinnate arrangement, with two main tendons converging to the
insertion on the lesser tuberosity of the humerus. The insertion is
composed of a broad tendinous attachment, and by fibers coursing
lateral of the tendon. Innervation is received from the two
subscapular nerves, derived from the dorsal division of the brachial
plexus.
Comparative aspects. The attachments of the subscapularis are
conservative in all bats (Strickler, 1978). The large size of the

77
subscapularis relative to body mass was noted by Macalister (1872) as
being largerr than observed in other mammals. Vaughan (1970a)
elaborated upon this point and provided data for the mass of
subscapularis relative to the total mass of the primary downstroke
muscles. This value ranged from 16.1 to 19.4 percent. Overall, the
relative weights of these primary downstroke muscles are consistent in
all bats. The data from Strickler (1978) are in agreement with Vaughan
except for in the Pteropodidae and for the phyllostomid genus
Desmodus. Strickler commented that pteropodids were unique among bats
in having small subscapularis muscles and small ventral muscle masses.
It was not established whether the small muscle masses represent a
primitive condition for chiropterans, or result from functional
constraints. In contrast, the large subscapularis muscle of Desmodus
appeared to be related to the terrestrial habits of the vampire bat,
for which there are increased demands for shoulder extension and wing
adduction (Strickler, 1978).
Functional aspects. Electromyography of the subscapularis in
free-flying Artibeus indicates that the muscle remains active through
approximately 75 percent of the wingbeat cycle. High-amplitude EMG
activity is observed through approximately 26 percent of the wingbeat
cycle and coincides with the second one-half of the upstroke and first
one-tenth of the subsequent downstroke. This activity precedes
myopotentials recorded from the other two downstroke muscles, serratus
ventralis thoracis and pectoralis. Subscapularis was discussed as a

78
component of the rotator cuff complex of primates and other mammals
(Tuttle and Basmajian, 1978) and is important in stabilizing the
shoulder joint. This stabilization includes the role of holding the
articular surface of the humerus within the glenoid fossa. Hermanson
and Altenbach (1981, 1983) concluded that subscapularis was primarily
responsible for fine control and stability of the shoulder joint in
Antrozous. They also observed that the expression of continuous versus
episodic activity in subscapularis was facultative and therefore
dependent upon the nature of the flight maneuver being attempted at any
given time. Kovtun and Moroz (1974) also used electromyography and
observed that subscapularis preceded pectoralis activity and ceased
activity 6-10 msec after mypotentials terminated in the "chest muscles"
of Myotis myotis. It was not clear whether their data were pooled from
several wingbeats or represented only one wingbeat.
The timing of subscapularis activity does not support Vaughan's
hypothesis (1959) that subscapularis is the third muscle among the
primary downstroke muscles to commence activity during a wingbeat.
However, the high-amplitude activity fit well with the temporal pattern
observed in downstroke muscles: myopotentials commence approximately
25 msec before wing adduction begins and terminates during the early
downstroke. Low-amplitude myopotentials during the early downstroke
and early upstroke indicate stabilization activity at the shoulder
joint. Low-amplitude activity at the transition between downstroke and
upstroke, and during the early upstroke stabilizes the shoulder against

79
forces produced by the abductor muscles during this period. Because of
these two periods of activity and the roles that the muscle carries
out, adduction and stabilization, the subscapularis is classified as a
bifunctional muscle and serves primarily to stabilize the shoulder
joint.
Deltoid Group
The deltoideus of bats has long been recognized as a tripartite
muscle composed of three parts: clavodeltoideus, acromiodeltoideus,
and spinodeltoideus (Macalister 1872). Because of functional
specializations associated with each of these parts, I will describe
and refer to each part specifically. All parts are innervated by the
axillary nerve.
Clavodeltoideus
Form. The clavodeltoideus originates by fibers along the
expanded, distal three-quarters of the clavicle. The muscle fibers
course dorsally, proximal to the apex of the pectoral ridge of the
humerus, and insert near the proximal insertion of acromiodeltoideus.
Fibers are arranged in parallel.
Comparative aspects. Vaughan (1959) pointed out that the
clavodeltoideus is separated from adjacent fibers of the pectoralis
only with great difficulty. Visualization of the axillary nerve is
necessary to correctly identify muscle bundles belonging to the

80
deltoideus group. Strickler (1978) illustrated some of the variation
in extent of origin that is possible within the order Chiroptera and
thus the need to undertake careful dissection of this region.
Functional aspects. The functional analysis of clavodeltoideus is
discussed along with the spinodeltoideus.
Acromiodltoideus
Form. The acromiodeltoideus has a fibrous origin along the entire
tip of the acromion process and proximally, along the base of the
acromion process as it intergrades with the scapular spine. Insertion
is by fibrous attachment along the lateral ridge of the humerus, from 1
mm distal to the greater tubercle along the proximal one-third of the
humerus. The myofiber architecture is parallel with some fibers
diverging distally along the insertion. A thin fascial plane bisects
the muscle into anterior and posterior portions and extends distally
from the middle of the acromion process.
Comparative aspects. The primitive condition for the origin of
the chiropteran acromiodeltoideus is restricted to the acromion process
(Strickler, 1978). In advanced families, the origin of the muscles has
extended medially along the scapular spine, a position that places
fibers farther from the shoulder joint and that increases the
effectiveness of acromiodeltoideus as an abductor and flexor of the
shoulder (Strickler, 1978). In Antrozous, fibers of acromiodeltoideus
and acromiotrapezius attach to each other along the dorsal scapular

81
ligament (Hermanson, 1978). The situation in Artibeus, in contrast, is
for both acromiodeltoideus and acromiotrapezius to insert upon a solid
mass of bone, adjacent to the acromion process.
Functional aspects. The action of the acromiodeltoideus is
discussed along with the spinodeltoideus.
Spinodeltoideus
Form. The spinodeltoideus originates by fibers from the dorsal
scapular ligament (sensu Hermanson and Altenbach, 1983), from the
vertebral border of the scapula caudal to the scapular spine, and over
the cartilaginous plate at the caudal angle of the scpaula. Insertion
is by an aponeurosis on the medial aspect of the dorsal ridge of the
humerus, deep to the insertion of the acromiodeltoideus and immediately
distal to the insertion of teres minor. The muscle is parallel-fibered
and forms a broad but thin sheet over the caudal scapular region.
Vaughan (1959) commented that spinodeltoideus was composed of two parts
in Macrotus.
Functional aspects. The deltoideus muscle of man is a single
muscle mass that was divided by anatomists into an anterior and
posterior portion. Each division can be correlated with differential
EMG activity while performing several actions (Inman et al., 1944).
Similarly, Hermanson and Altenbach (1983) presented functional
interpretations for all three heads in Antrozous and discussed these
data with respect to published data for several terrestrial mammals.
The acromiodeltoideus and spinodeltoideus EMG patterns for Antrozous

82
were in general agreement with the hypotheses of earlier descriptive
studies (Vaughan, 1959, Strickler, 1978): the muscles functioned as
abductors of the wing. The clavodeltoideus, however, vías active almost
in synchrony with the primary downstroke muscles. This observation,
and dissection studies (Norberg, 1972; Strickler, 1978), led Hermanson
and Altenbach to conclude that the muscle was an important humeral
protractor and also a humeral adductor.
During the propulsive phase in Antrozous, a lack of muscle
activity in acromiodeltoideus and spinodeltoideus contrasted with
observations of EMG patterns in the deltoideus of terrestrial mammals.
Deltoideus activity occurred during the stance phase in stepping cats
at several speeds of locomotion (English, 1978a). Biphasic activity
occurred in the deltoideus muscle in Didelphis: the largest amplitude
activity occurred during the swing phase, and a less intense period of
activity during the stance phase (Jenkins and Weijs, 1979). When the
acromioclavicular and spinal portions of the Didelphis deltoideus were
monitored simultaneously, the spinal portion exhibited intense activity
during the first one-third of the swing phase, while the
acromioclavicular portion was active during the final two-thirds of the
swing phase (Jenkins and Weijs, 1979). Recordings from the deltoideus
of Canis indicated continuous myopotentials throughout the step cycle
of walking, trotting, and galloping dogs (Tokuriki, 1973a, 1973b,
1974).

83
Data for the deltoideus of Artibeus provide contrasts and
similarities with the Antrozous observations. For example, the primary
(highest amplitude) EMG activity in the clavodeltoideus was evident
during the downstroke portion of the wingbeat. Myopotentials commenced
almost in synchrony with the pectoralis, contributing to wing adduction
but also to the anterior movement of the wing. This wing protraction
is an important aspect of the "Rudderflug” or rowing flight described
by Eisentraut (1936) and Norberg (1976). Low-amplitude activity of
brief duration occurs during the transition between downstroke and
upstroke and serves to stabilize the scapula. In contrast,
highest-amplitude activity in both the acromiodeltoideus and
spinodeltoideus occurred during the late downstroke and early
upstroke. These data therefore suggest an important role for
acromiodeltoideus and spinodeltoideus in wing abduction and agree with
observations in Antrozous (Rermanson and Altenbach, 1983). Both
acromiodeltoideus and spinodeltoideus were biphasically active. The
posterior part of acromiodeltoideus showed a low-amplitude phase of
activity roughly coincident with the primary burst of clavodeltoideus,
although of shorter duration and with less amplitude than in the latter
muscle. Spinodeltoideus exhibited low-amplitude activity during the
first one-third of the downstroke. These secondary EMG bursts in the
acromiodeltoideus and spinodeltoideus may stabilize the shoulder
against the powerful actions of the pectoralis.

84
Suprascapular Group
Supraspinatus
Forro. Supraspinatus originates by fibers along the cranial and
medial perimeter of the supraspinous fossa, deep to the belly of
acromiotrapezius. The muscle fibers are arranged in parallel. This
muscle has a fibrous insertion on the tip of the greater tubercle of
the humerus. Innervaton is by the suprascapular nerve.
Comparative aspects. The supraspinatus exhibits little variation
in its attachments for all bat species (Vaughan, 1959, 1970a;
Strickler, 1978). The regression of supraspinatus mass against body
mass suggests that the size of this muscle is average to relatively
large size in slow-flying bats, and smaller in faster-flying species
(Strickler, 1978).
Functional aspects. The mammalian supraspinatus is considered one
of the rotator cuff muscles of the shoulder: its primary role is to
stabilize the glenohumeral articulation in concert with the
infraspinatus, subscapularis, and teres minor (Hollinshead, 1974;
Tuttle and Basmajian, 1978). For bats, Vaughan (1959) described the
general action of the supraspinatus to be shoulder extension and
humerus supination, a description accepted by subsequent authors (cf.
Strickler, 1978). Vaughan (1959) suspected that supraspinatus was most
effective as a shoulder extensor in the phyllostomid Macrotus, as
compared to Eumops and Myotis. The shoulder joint of Macrotus allows

85
more freedom of movement than is observed in Eumops and Myotis. Both
of the latter species had bony locking mechanisms that restricted
movement of the humerus in both the dorsoventral and craniocaudal
directions. Thus, the humerus of Macrotus could be extended cranial
to a transverse plane through the shoulder, while the humerus of Eumops
could only be extended forward to this plane.
Electromyographic data for the supraspinatus is available for only
six wingbeats, primarily because the small size and deep position of
the muscle make the surgical implantation a traumatic proceaure for the
animal. The muscle exhibited a single, extended period of activity
commencing during the early upstroke, and continuing through the early
downstroke. During two of the wingbeats, supraspinatus exhibited two
periods of normal-amplitude EMG activity with a brief period of
quiescence interposed.
Biphasic EKG activity was observed in supraspinatus of Antrozous
(Hermanson and Altenbach, 1983). Biphasic activity was also observed
during the step cycle of Didelphis (Jenkins and Weijs, 1978), but not
during stepping in cats (English, 1978a_) or dogs (Tokuriki, 1973a,
19731), 1974). In several primate species, Tuttle and Basmajian (1978)
reported prominent activity of the supraspinatus during humeral
elevation and cautioned that the function of this muscle may be one of
acceleration of the humerus rather than as a simple stabilizer (i.e.,
rotator cuff). The variability observed in all of these studies may be
attributed to small sample sizes, at least for the two bat species

86
studied, and to an inconsistent pattern of locomotor movements studied
in the primate examples. In the latter study, movements of the animal
were not restricted to forward progression, but instead included
forward, lateral, and vertical movements. Support for a role in wing
supination is provided by EMG data indicating activity during the early
upstroke when the wing is supinated and abducted.
Infraspinatus
Form. The infraspinatus originates from the periphery of the
infraspinous fossa of the scapula: cranially from the spine of the
scapula; medially from the dorsum of the vertebral border of the
scapula; and laterally from the raised ridge of the infraspinous
fossa. All fibers insert upon a central tendon that courses through
the middle of the muscle. This tendon forms the sole insertion of the
infraspinatus upon the lateral ridge of the humerus. Fibers at the
medial end (nearest the vertebral border) are the longest in the muscle
and insert upon the central tendon at an acute angle. Fibers that
insert more distally on the central tendon are shorter and tend to
intersect the tendon at a larger angle than the medial fibers.
Innervation is by the suprascapular nerve, after that nerve provides a
branch to the supraspinatus and courses laterally under the tip of the
acromion process.
Comparative aspects. Strickler (1978) noted little variation in
the attachments of the infraspinatus in all bat species that he
studied. In contrast to the regression of muscle mass on body mass

87
obtained for supraspinatus, Strickler observed that the muscle was
larger in fast-flying species, but smaller in fast-flying frugivores.
No functional explanation was provided. Within families, there was no
consistent ratio observed between the mass of infraspinatus and
supraspinatus.
Functional aspects. Two phases of activity were evident in the
infraspinatus of Artibeus. The first period and the one exhibiting the
highest amplitude myopotentials commenced immediately prior to the
onset of downstroke movements, and continued during the first third of
the downstroke. A second, less intense period of activity was observed
in six wingbeats during the late downstroke and early upstroke. These
data were similar to observations made on the infraspinatus of
Antrozous (Hermanson and Altenbach, 1983). Although two distinct
phases were also observed in walking Didelphis (Jenkins and Weijs,
1979), only one period of activity was noted in cats (English, 1978)
and dogs (Tokuriki, 1973a^ 1973^b, 1974) at all gaits.
On the basis of anatomical position the infraspinatus is a
shoulder flexor, and a humerus abductor and supinator (Hermanson and
Altenbach, 1983; Strickler, 1978; Altenbach, 1979). The biphasic
activity pattern suggests an underlying stabilizing role at the
shoulder; the secondary EMG activity observed during the early
downstroke counteracts the action of the pectoralis and protects the
glenohumeral joint in association with other intrinsic muscles of the
shoulder. Because the wing of a bat is not capable of being

88
"feathered" during the upstroke like that of a bird, the role of the
abductors is accentuated in bats (Vaughan, 1S5S). Thus, the
infraspinatus exhibits strong activity during the non-propulsive
upstroke, and acts in concert with the more superficial spinodeltoideus
to abduct the wing.
Triceps Group
The triceps brachii is composed of three heads in most mammals.
Each head is innervated by branches of the radial nerve. Because of
the different origins of these heads, each is treated separately.
Triceps Brachii, Medial Head
Form. The medial head has a fleshy origin along the caudal
surface of the humerus, approximately 10-13 mm distal to the greater
tubercle. Muscle fibers pass medial to the radial nerve. Fibers
insert distally in a unipinnate fashion on the main triceps brachii
tendon. This is the smallest of the three heads of triceps brachii.
Comparative aspects. In most phyllostomids, the medial head
originates on the middle one-third of the ventral surface of the
humerus (Strickler, 1978). The attachments are conservative in most
chiropterans.
Functional aspects. The medial head of triceps brachii spans one
joint, the elbow. Vaughan (1959) suggested that the medial head
stabilizes the sesamoid bones found at the insertion of the triceps
brachii tendons in many bats. No EMG data are available for any bats
to corroborate or refute the hypothesis.

89
Triceps Brachii, Lateral Head
Form. The lateral head originates by fibers on the proximal
one-quarter of the humerus, primarily on the caudal surface adjacent to
the greater tubercle, surgical neck, and lesser tubercle. All fibers
pass lateral to the radial nerve. Central fibers insert directly upon
a central tendon of insertion. Peripheral fibers insert on either side
of this tendon at an angle of less than 10 degrees.
Functional aspects. The lateral head of triceps brachii crosses
one joint, the elbow joint, and is capable to effecting extension of
that joint. EMG data is discussed along with the long head of triceps
brachii.
Triceps Brachii, Long Head
Form. The long head arises from the axillary border of the
scapula, adjacent to the infraglenoid tubercle. The muscle is
pinnate. Fibers course distally to insert upon a central tendon in
common with the other two heads of triceps brachii. The main
insertional tendon contains a small sesamoid caudal to the elbow
joint. A synovial cavity is present deep to the tendon, permitting
movement of the tendon relative to the joint in this area. The
insertional tendon of the triceps brachii attaches to the caudal
surface of the olecranon region of the ulna.
Comparative aspects. The anatomy of the triceps brachii does not
depart significantly from the general pattern observed in other bats
(cf. Strickler, 1978): the long head spans the shoulder and elbow
joints, the lateral head spans only the elbow joint.

90
Functional aspects. In terrestrial mammals, the triceps brachii
functon as anti-gravity muscles principally by maintaining elbow
extension during standing or locomotion (Armstrong et al., 1982;
English, 1978b^; Jenkins and Weijs, 1979). EMG data for long and
lateral head of triceps brachii in Antrozous did not clearly support an
elbow extensor hypothesis (Hermanson and Altenbach, 1983). In A.
jamaicensis, muscle activity in the long head commenced prior to
initiation of the upstroke, and continued throughout the first one-half
of the upstroke. The lateral head also commenced activity prior to the
downstroke, but relative to the long head, EMG activity began later and
was of shorter duration. These temporal patterns are similar to
observations of triceps brachii activity in Antrozous (Hermanson and
Altenbach, 1983). In both bat species, the long and lateral heads of
triceps brachii exhibited activity patterns assoicated with wing
abduction. It is not clear how either portion of triceps brachii
effects elbow extension, a movement associated writh the downstroke
phase of the wingbeat cycle (Eisentraut, 1936; Norberg, 1976; Hermanson
and Altenbach, 1983). Elbow7 extension automatically contributes to
spreading the distal wing during the downstroke in many bat species
(Vaughan, 1966; Vaughan and Bateman, 1970). In Artibeus, the amount of
time between EMG onset and downstroke initiation was approximately 55
msec. Armstrong et al. (1977) and Hermanson and Foehring (1982)
demonstrated the existence of fast type histochemical profiles for
triceps brachii in Myotis and Tadarida, respectively. In several
mammalian muscles, fast type profiles are correlated with contraction

91
times of less than 35 msec (Burke, 1978). The effect of actvity in
triceps brachii is implemented during the upstroke, at first to abduct
the limb, and then possibly during the late upstroke, to extend the
elbow.
Pectoralis Group
Subclavius
Form. The subclavius originates along the lateral aspect of rib
one and the associated costal cartilage by fibrous attachments. The
muscle inserts on the distal seven-eighths of the ventrolateral surface
of the clavicle, by fibers. The muscle exhibits a parallel-fiber
architecture. Innervation is received from the upper pectoral nerves.
Comparative aspects. The attachments of the subclavius are
relatively similar in all bats with most differences limited to the
position of the insertion upon the clavicle (Strickler, 1978).
Struhsaker (1963) reported that the subclavius is relatively larger in
bats that have low aspect ratio wings. Altenbach (1979) noted the
robust nature of the subclavius in Desmodus, particularly with respect
to the insertion of the muscle upon nine-tenths of the length of the
clavicle. In Desmodus the ability to adduct the limb rapidly and
shoulder girdle is important in avoiding injury or predation. The
former situation could accrue while the vampire bat approaches the

92
fetlock joint of livestock and the potential predator steps back
suddenly or kicks at the bat. Also, lift off from the ground after
feeding is complicated by the added weight of the blood meal.
Altenbach (1979) felt that the subclavius, through its action on the
clavicle, could power the initial leap off the ground into flight.
Functional aspects. Vaughan (1959) deduced that the subclavius
pulls the clavicle ventrally and caudally, probably to steady the
clavicle against the forces of the dorsal abductor muscles. This
movement of the clavicle has been construed to contribute to wing
adduction in two ways. As previously mentioned, the clavicle may
adduct in synchrony with the wing during initial take-off maneuvers.
Also, adduction of the clavicle in synchrony with the wingbeat
contributes to effective force production in the major downstroke
muscle, the pectoralis, by reducing the overall excursion that this
latter goes through during the downstroke (Hermanson, 1981).
Pectoralis
The pectoralis muscle is divided into two parts in all bats
(Strickler, 1978), although the division is not usually visible on the
superficial surface of the muscle. The cranial division (clavicular
head) arises from the clavicle and parts of the manubrium. The caudal
division (sternal portion) arises from the sternum inclusive of parts
of the manubrium and the xiphoid process.

Figure 16.—Ventral view of the shoulder and arm of Artibeus jamalcensis.
Superficial muscles are exposed on the left; deeper muscles are exposed
on the right.

pectoralis %
biceps brachii:
long head
short head ....
serratus ventralis
thoracis
pectoralis, cranial div.
subclavius
subscapularis
s
X
^ triceps brachii:
lateral head
long head
'' pectoralis abdominalis

95
In many terrestrial mammals, the pectoralis includes a transverse
head located cranially and superficially, and a deep head originating
more caudally and coursing deep to the transverse head (Howell, 1926;
Rinker, 1954; Getty, 1975). The cranial division is equivalent to the
anterior division of Vaughan’s terminology (1959) and in bats may be
homologous to the transverse pectoralis of other mammals. The caudal
division is equivalent to the posterior division of Vaughan's
terminology. However, the variation observed within the chiropteran
pectoralis makes it difficult to consistently delineate the two
divisions. The pectoralis minor of humans and other higher primates
cannot be related with either part of the chiropteran pectoralis
complex (cf. Inman et al., 1944). I follow Vaughan's descriptions of
the myology of pectoralis, but have changed the terminology to avoid
the use of the terms "anterior" and "posterior," in accordance with
usage prescribed in the N.A.V.
Pectoralis, Cranial Division
Form. The cranial division of pectoralis arises from the cranial
end of the manubrium, and from the proximal two-thirds of the ventral
surface of the clavicle, by fibers. The insertion is by fibers along
the ridge extending from the greater tubercle to the pectoral ridge and
along the apex of the pectoral ridge superficial to the insertion of
the caudal division of this muscle. Innervation is by the cranial
pectoral nerves. The muscle fibers are parallel-fibered.

96
Comparative aspects. The cranial division is difficult to
separate from the adjacent clavodeltoideus in phyllostomids.
Consequently, Strickler (1978) combined the two muscles for the
purposes of description. Strickler reported that the combined relative
mass of the cranial division and the clavodeltoideus was large in
phyllostomid bats and particularly large in hovering species.
Functional aspects. The cranial division is discussed along with
the functional aspects of the caudal division of pectoralis.
Pectoralis, Caudal Division
Form. The caudal division of pectoralis arises by fibers along
the entire sternum from the keeled ventral portion of the manubrium,
from a raphe ventral to the mesosternum, and by fibers from the ventral
surface of the xiphisternum. Some deep fibers have a fleshy origin
from the ventral aspects of costal cartilages 2-4 and intercostal
spaces 2-3. At the caudal end of the muscle, some laterally placed
fibers attach to the abdominal fascia, ventral to the costal arch.
Fibers are generally arranged in parallel but converge laterally. The
insertion is both fibrous and aponeurotic along the distal one-half of
the pectoral ridge and along the cranial face of the humerus, 3 mm
distal to the pectoral ridge, distal to the insertion of pectoralis
abdominis, and cranially, deep to the insertion of the cranial division
of pectoralis.
Comparative aspects. The caudal division of pectoralis attains
its greatest relative size in the pteropodids and in Nycteris. In
these bats the caudal division of serratus ventralis thoracis is not

97
well developed to assist the pectoralis as an adductor of the scapula
(Strickler, 1978). Otherwise, the form of the muscle is conservative
among all bats (Vaughan, 1970a; Strickler, 1978).
Functional aspects. The pectoralis muscle of Artibeus was
analyzed in six regions to determine whether the muscle exhibits
homogeneous activity patterns. The origin extends along the entire
sternum, with fibers at the cranial end coursing at an angle of
approximately 60 degrees relative to the median plane, and caudal
fibers coursing approximately 40 degrees relative to the median plane.
Vaughan (1959) concluded that different regions of the muscle were
important in terms of the cranial or caudal components of force that
were exerted upon the humerus. In an analogous muscle, Herring et al.
(1977) demonstrated that different regions of the masseter muscle of
miniature pigs were active at different points in the chewing cycle, a
pattern they labelled functional heterogeneity. The pectoralis of bats
is similar in that the muscle is broad along the origin, with fibers
converging distally upon the insertion. Electromyograms in the
pectoralis of Artibeus exhibit a general gradient of activity: muscle
activity occurred earliest in cranial bundles of the muscle, the
cranial division. There was not a sequential cranial to caudal
sequence of EMG onset in regions 1-5 (see Figure 17 for identification
of elecrtrode locations). Region 3, the middle of pectoralis was
active next, followed in order by Regions 2, 1, 5, and 4. Based upon
the similarity in mean onset times, it is not possible to assess

Figure 17.—Electromyographic data for six regions in the
pectoralis muscle during slow flight. Electrode locations
are indicated in the upper figure, a ventral view of Artibeus
jamaicensis. Position 1 is in the cranial division of
pectoralis. Positions 2 to 6 are in successively more caudal
locations of the caudal division of pectoralis. In the lower
figure, unshaded bars indicate observed periods of normal-
amplitude muscle activity. Vertical lines are placed one
standard deviation from the means of activity onset and
termination.

99
I downstroke T upstroke
cranial 1
2
3
4
5
caudal 6
.00 .20 .40 .60 .80 1.0

100
Table 3.—Electromyographic data for activity patterns in six regions
of the pectoralis muscle of Artibeus jamaicensis during slow flight.
N= number of wingbeats analyzed per muscle. Mean duration of the
downstroke and time of muscle activity onset and termination are
expressed as a numerical value relative to the total wingbeat cycle
(mean + one S.D.).
Downstroke
N Duration
Onset
Time (S.D.)
Termination-
Time (S.D.)
Cranial division
1
12
0.494
0.749
(.022)
0.105
(.080)
Caudal division
2
9
0.533
0.801
(.048)
0.045
(.135)
3
17
0.496
0.792
(.050)
0.145
(.034)
4
7
0.526
0.785
(.031)
0.094
(.086)
5
9
0.526
0.877
(.039)
0.170
(.056)
6
7
0.542
0.837
(.036)
0.149
(.054)

101
activity heterogeneity within the pectoralis. A better determination
should be made with simultaneous recordings from all six regions in one
bat. In order to perform this experiment, a sufficiently light
electrode and plug assembly must be designed for use in bats.
The EMG data obtained for Artibeus corresponds with data for
Antrozous (Hermanson and Altenbach, 1981, 1983) and Eptesicus
(Altenbach and Hermanson, unpublished). A single phase of activity
began during the late upstroke and continued through the early
downstroke of the following wingbeat. Hermanson and Altenbach (1981)
suggested that pectoralis is the primary muscle powering the downstroke
of bats. Because of the high frequency and short duration of the
chiropteran wingbeat, the pectoralis exhibits electrical activity
during the late upstroke. The mechanical effect of the myopotential is
not realized until the transition from upstroke to downstroke. There
is a 22-25 msec delay between the onset of pectoralis activity and the
beginning of the downstroke. Mechanical studies of other mammalian
muscles demonstrated that a 25 msec delay between onset of EMG activity
and maximum tension production is expected during isometric
contractions of fast twitch muscles (Burke et al., 1974; Burke, 1978).
Chiropteran pectoralis muscle has been classified as fast twitch using
several histochemical staining techniques (Armstrong et al., 1977;
Hermanson and Foehring, 1982). Activity in the pectoralis terminates
during the first one-third of the downstroke. The muscle remains quiet
during the remainder of the wingbeat cycle.

102
Electromyograms obtained from stepping opossums (Jenkins and
Weijs, 1979) and cats (English, 1978a) indicated that pectoralis in
terrestrial mammals was active throughout relatively more of the
propulsive phase than in bats. In the present study, the propulsive
phase of terrestrial stepping is considered to be equivalent to the
downstroke of flying bats.
Pectoralis Abdominalis
Form. Pectoralis abdominalis lies deep to the pectoralis. The
origin of this muscle is along the ventral surface of the costal arch
and on the surface of the external abdominal oblique muscles, extending
from 7-12 mm lateral of the median plane. Insertion is by a flat
tendon on the ventral, medial surface of the humerus, proximal to the
apex of the pectoral ridges.
Comparative aspects. The pectoralis abdominalis has been
described in bats (Strickler, 1978) and rodents (Woods, 1972). The
muscle does not occur in domestic mammals (Getty, 1975; Evans and
Christensen, 1979). Macalister (1872) doubted synonomy with the
"lesser pectoral or pectoralis minor" of man because the pectoralis
abdominalis is innervated by the anterior thoracic nerve from the
"outer cord of the brachial plexus, not by the middle, which should
supply it if it were the lesser pectoral." The muscle was similar in
all chiropterans (Strickler, 1978).
Functional aspects. Vaughan (1959) concluded that the muscle was
not important in flight. He noted that it was best developed in bats
using extensive terrestrial locomotion in their locomotor repertoire.

103
The muscle was also well developed in Antrozous (Hermanson, 1978) and
Desmodus (Altenbach, 1979). Both of these bats feed upon items located
on the ground.
Flexor Group of Arm
Coracobrachialis
Form. The coracobrachialis muscle is small and unipinnate,
arising from the lateral tip of the coracoid process deep to the short
head (coracoid head) of biceps brachii. Insertion is by a single
tendon on the anterior surface of the shaft of the humerus,
approximately two-thirds of the distance along the humerus. Fibers
insert along the tendon at an angle of approximately 18-22 degrees.
Comparative aspects. The muscle belly is small relative to the
adjacent long and short heads of biceps brachii (Strickler, 1978). The
coracobrachialis is not present in the Molossidae (Vaughan, 1959,
1970b). Innervation is by a proximal branch of the musculocutaneous
nerve.
Functional aspects. The function of coracobrachialis was
described by Vaughan (1959), Altenbach (1979), and Strickler (1978) as
a weak adductor of the wing. No EMG data are available for this
muscle.
Biceps Brachii
Synonymy. The biceps brachii includes two heads in bats. Vaughan
(1959) introduced the terms "coracoid head" and "glenoid head" to
parallel the "short" and "long" terminology used in human anatomy. I

104
will use the latter terms in accordance with usage proposed in the
N.A.V. Both heads are innervated by the musculocutaneous nerve after
the nerve emerges through the lateral surface of coracobrachialis.
Form. The long head of biceps brachii originates from the lateral
surface of the base of the coracoid process by a stout tendon. The
tendon courses laterally across the shoulder joint capsule anc emerges
through the bicipital groove, ventral to the pectoral ridge. Fibers
attach to the tendon or origin in a fusiform pattern but are generally
arranged in parallel throughout the belly. Distally, the fibers attach
to a broad tendinous sheet that narrows distally and courses to the
flexor groove on the anterior surface of the radius.
The short head of biceps brachii originates from the ventral tip
of the coracoid process. The fibers are arranged in parallel.
Distally, the fibers converge on a single tendon that inserts on the
flexor groove of the radius, separate from and proximal to the
insertional tendon of the long head.
Functional aspects. Although no EMG data is available for the
biceps brachii of A. jamaicensis, the anatomy of the muscle and its two
heads is sufficiently similar to the condition in Antrozous to permit
some comparison. Hermanson and Altenbach (1983) noted biphasic
activity patterns in both the long and short heads of biceps brachii.
These authors concluded that the long head initiated elbow flexion
during the upstroke. Later during the upstroke, a burst of activity

105
in the short head contributed to elbow flexion. Electromyograms also
demonstrated muscle activity throughout middle and later portions of
the downstroke in the short head, preceded by a burst of activity
during the early downstroke in the long head. Therefore, both heads of
biceps contributed to wing adduction, and also to elbow flexion.
Brachialis
Form. The brachialis is a small muscle located on the lateral
aspect of the arm. The origin is fleshy, along the dorsolateral
surface of the humerus and along the middle one-third of that bone.
Fibers of brachialis insert along the anterior surface of the radius,
proximal to the insertion of biceps brachii. The radial nerve courses
alongside the muscle in the arm, however, the innervation of brachialis
is by fibers of the musculocutaneous nerve.
Functional aspects. Although brachialis is an important elbow
flexor in most mammals (observation based on its one-joint condition),
it is weak and probably ineffectual in Artibeus and in other bats.
Antebrachial Extensor Group
The forearm muscles of the chiropteran wing include those muscles
originating on the distal end of the humerus and extending to insert
upon the antebrachial bones, the manus, and on the digits. These
muscles are small and difficult to see, precluding a thorough

106
comparative discussion. However, a description of their anatomy and
proposed function in Artibeus is included here.
The forearm muscles include a superficial group that originate
near or upon the lateral and medial epicondyles of the humerus.
Although these muscles probably stabilize the elbow joint, their
function is of greatest importance at the wrist or digital joints. At
the elbow joint, the brachial (triceps brachii and biceps brachii)
possess the greatest mechanical advantages and are primarily
responsible for elbow flexion and extension. The deep layer of forearm
muscles originate in general from the radius, ulna, or interosseus
membrane. These muscles produce wrist or digital movements. The
following descriptions provide a foundation for future studies of the
forelimb musculature in Artibeus and in other bats.
Extensor Carpi Radialis
The extensor carpi radialis is usually treated as two separate
muscles (cf. Rinker, 1954; Vaughan, 1959). I discuss the two heads as
a single functional unit both for simplicity and because the bellies
and proximal tendons are separable only with great difficulty in
Artibeus.
Form. The extensor carpi radialis longus has a tendinous origin
on the caudal surface of the epicondylar ridge. A small number of
fibers sweep proximally from this tendon along the epicondylar ridge.
Insertion is on the dorsal base of metacarpal I and on the

Figure 18.—Lateral view of the muscles of the elbow region.
The upper figure demonstrates the superficial muscles.
Deeper muscles are exposed in the lower figure by removal
of the common digital extensor and ulnaris lateralis.

108
biceps brachii
extensor carpi radialis
abductor pollicis longus
extensor pollicis brevis
supinator
5 mm

Figure 19.—Dorsal view of the muscles of the carpal region,
intrinsic muscles of the manus have been removed.
The

extensor carpi radialis longus
,, extensor carpi radialis brevis
5 mm
o

Ill
anterior surface of the base of metacarpal II. Innervation is by the
deep branch of the radial nerve adjacent to the elbow.
The insertional tendon of extensor carpi raaialis longus courses
along the anterior edge of the radius, and is separable only with
great difficulty from the adjacent tendon of extensor carpi radialis
brevis. Both tendons are crossed by the abductor pollicis longus
tendon on the distal one-third of the radius. The tendon splits
distally, just before coursing over the extensor grooves on the
distal end of the radius. The tendon of extensor carpi radialis
longus passes through the anterior extensor groove of the radius,
over the manus and to its insertion. In the carpal region, the
tendon is crossed by extensor pollicis brevis.
The extensor carpi radialis brevis arises by tendon, along with
the long head, from the caudal surface of the epicondylar ridge. The
muscle belly largely lies superficial and dorsal to the belly of the
long head, and the tendon of insertion courses alongside and
dorsolateral to the tendon of the long head. Over the distal radius,
the two tendons diverge: the tendon of extensor carpi radialis
brevis courses over the medial extensor groove of the radius, across
the carpus, and to the insertion on the dorsum of the trapezium and
base of metacarpal III.
Comparative aspects. Studies of other bats show little evidence
for variation in the attachments of extensor carpi radialis (Vaughan,

112
1970b; Vaughan and Bateman, 1970; Altenbach, 1979). The muscles are
quite similar in form to the same muscle in Alouatta (Schon, 1968)
and in rodents (Klingener, 1964). The two heads of extensor carpi
radialis are treated as a single entity by Evans and Christensen
(1979) for the domestic dog, although the insertions are similar to
those described in other mammals.
Functional aspects. The action of extensor carpi radialis is to
extend the váng. When the wing is closed, as during roosting or
terrestrial locomotion, digits II through V are held approximately
parallel with the radius. During the downstroke, the carpus is
extended primarily at the radiocarpal joint. The anatomy of both
heads of the extensor carpi radialis permits extension at the
radiocarpal joint, slight extension across the carpal joints, and
both dorsal extension and adduction of digits I and II. The latter
action braces the leading edge of the wing against the force of the
airstream. Norberg (1972) described the force-lever system of the
extensor carpi radialis longus tendon to demonstrate the action
relative to a fulcrum at the base of metacarpal II. The dorsal
extension and adduction of metacarpal II, in turn, is linked to digit
III by the dactylopatagium minus (the wing membrane between digits II
and III) and by a ligament between the phalanx of digit II and the
middle phalanx of digit III.

113
Supinator
The supinator originates by a short tendon on the lateral aspect
of the lateral epicondyle. The tendon of origin contains a small
sesamoid bone. The fibers insert at an angle on the tiorsal surface
of the proximal 15 mm of the radius. Innervation is by the deep
branch of the radial nerve. The belly of supinator is located
lateral to extensor capri radialis longus and medial to the common
digital extensor.
Norberg (1970), Vaughan (1970), and Altenbach (1979) described a
small sesamoid bone in the tendon of origin. The sesamoid was also
present in the tendon of Artibeus, and serves to increase the lever
arm of the muscle across the dorsum of the joint. The supinator is a
one-joint muscle. This muscle is a flexor of the elbow because
movement of the elbow is limited to flexion and extension (Vaughan,
1970b). In rodents and primates, the muscle is similar to the
chiropteran supinator (Woods, 1972). However, in these forms,
relatively more movement of the elbow is possible and the muscle
functions primarily as a weak supinator of the forearm.
Extensor Pollicis Brevis
Form. The extensor pollicis brevis originates by fibers on the
lateral interosseous surface of the proximal ulna, and by fibers
arising from the deeper abductor pollicis longus. The muscle belly
extends along the proximal 25 mm of the radius. The tendon passes
through the proximal and middle extensor retinaculum, and then

114
cranially across the carpus. Insertion of the tendon is on
metacarpal I, and on the base of the first phalanx of digit I.
Innervation is by the radial nerve.
Comparative aspects. No substantial variation was observed in
other chiropteran species. Woods (1972) argued that no extensor
pollicis brevis is found in hystricognathous rodents or in most other
mammals. However, in bats, the extensor pollicis brevis and abductor
pollicis longus diverge in the middle third of the antebrachium and
course separately across the ventral and dorsal surfaces of the
carpus respectively.
Functional aspects. The extensor pollicis brevis extends digit I
at the carpometacarpal and metacarpophalangeal joints. During slow
flight the pollex is flexed to increase the camber of the wing.
Position of the pollex is controlled by the relative tension produced
in the extensor pollicis brevis and in the pollical part of flexor
digitorum profundus. Activity in the extensor pollicis brevis
counters the downward force of the airstream upon the leading edge of
the wing.
Abductor Pollicis Longus
Form. The abductor pollicis longus originates by fibers on the
entire lateral interosseous surface of the ulna and adjacent
interosseous surface of the radius. The fibers are arranged in a
bipinnate fashion on a central tendon of insertion. Distal to the
interosseous membrane, the tendon courses over the cranial surface of

115
the radius, and across the superficial tendons of the extensor carpi
radialis. At the distal articular surface of the radius, the tendon
courses through a groove on the lateral side of the pseudostyloid
process. In the carpus, the tendon is attached to the ventral
surface of the scaphoid, and then continues to the ventral base of
metacarpal I. Innervation is by the deep branch of the radial nerve.
Comparative aspects. In most bats studied, the insertion of the
abductor pollicis longus is on the scaphoid (Vaughan, 1970b).
Although insertion on metacarpal one appears to be the usual
condition in mammals, Vaughan believed that a scaphoid attachment in
bats provided a mechanical linkage between abductor pollicis longus
and the base of metacarpal V. He argued that such a linkage
facilitated maintenance of the proper angle of attack in the
plagiopatagium during flight. In Desmodus, Altenbach (1979) observed
two insertions of the abductor pollicis longus that are similar to my
observations in Artibeus: one attachment was on the scaphoid while
the second attachment was on the surface of metacarpal one. Although
the insertion in Antrozous was normally on the scaphoid, the tendon
passed across the scaphoid and inserted directly on the thumb pad in
one specimen (Hermanson, 1978). No variation was observed in five
specimens of Artibeus.
Functional aspects. The abductor pollicis longus functions to
abduct the pollex, and thus provides a mechanism for positioning the

116
pollex. The flexor digitorum profundus simultaneously provices
positional control in the flexion-extension plane. Abduction of the
pollex occurs during the stance phase of walking. Fine control of
the pollex is also observed during head-up landing maneuvers. The
connection between the scaphoid and pisiform does not appear to
facilitate abductor pollicis longus control over digit V.
Lateral Digital Extensor
Vaughan (1959, 1970b) referred to this muscle as the extensor
digiti quinti proprius.
Form. The lateral digital extensor arises from an aponeurosis on
the lateral epicondyle and by fibers along the dorsal, cranial edge
of the ulna. The muscle attachments pass caudal to the center of
rotation of the elbow joint. A single tendon arises halfway along
the caudal aspect of the forearm and passes deep to the proximal
extensor retinaculum. Upon emerging from this sheath, the tendon
diverges from the adjacent tendon of the common digital extensor and
passes caudal to the carpus and the fifth carpometacarpal joint.
Eight to ten mm distal to the joint the tendon fuses with a branch
from the insertional tendon of the common digital extensor and
courses along the dorsal surface of the metacarpal, proximal and
distal phalanges of digit V. Extensor aponeuroses are formed over
the two distal joints, similar to those already described for the
common digital extensor. Innervation is by the deep branch of the
radial nerve.

117
Comparative aspects. Klingener (1964) assembled comparative
evidence that extensor digiti minimi (=lateral digital extensor)
represents the lateral (ulnar) side of the primitive deep extensors
of the antebrachium. In several rodent families, the insertion of
the lateral digital extensor inserts on digits four and five (Howell,
1932; Rinker, 1954; Woods, 1972). A similar insertion was noted in
primates (Schon, 1968; Howell and Straus, 1933). Vaughan (1959,
1970b) noted that the lateral digital extensor is absent in Eumops
and Hipposideros, and is only present on digit V in Hyotis.
Altenbach (1979) described essentially the same attachments in
Desmodus as I have observed in Artibeus. Norberg (1972) described
the insertion of the common digital extensor on digits III, IV, and V
in Rousettus. However, I believe she included the lateral digital
extensor as part of her common digital extensor. Norberg also
discussed and illustrated the tendon of common digital extensor
coursing from digit IV to the tendon of the lateral digital
extensor. I define the two muscles separately because the tendons
have separate bellies and the insertional tendons pass the carpus in
a different manner. The tendon of the lateral digital extensor
passes through the proximal flexor retinaculum while the tendon of
the common digital extensor passes through both the proximal and
middle retinacula.

118
Functional aspects. Vaughan (1959) and Altenbach (1979) agreed
that the muscle extends and deflects the fifth digit upward. The
passage of the tendon caudal to the fifth carpometacarpal joint
serves to abduct digit V, an important function in spreading the
chiropatagium during the downstroke.
Common Digital Extensor
Form. The common digital extensor crosses several joints,
passing from an origin on the distal humerus to digits III - V. The
origin of the common digital extensor is by two heads. The
superficial head arises by a broad aponeurosis on the lateral
epicondyle of the humerus, immediately lateral to the supinator. The
deep head arises from a thin tendon of origin that attaches to a
small circumscribed area on the lateral epicondyle. The muscle belly
lies superficially along the proximal one-half of the forearm,
gradually tapering to a distal tendon. This tendon splits along the
distal one-third of the radius into two tendons that pass through the
proximal and middle extensor retinacula (Fig. 19). Immediately
distal to the middle retinaculum, the lateral tendon turns caudad
across the carpus and metacarpal IV, giving off a small tendon to
digit V that fuses with the tendon of the lateral digital extensor,
8-10 mm distal to the carpal-metacarpal joint. The main tendon to
digit IV passes over the carpometacarpal joint and forms an extensor
aponeurosis over the metacarpophalangeal joint: central fibers of
the extensor hood insert on the base of the proximal phalanx while
lateral fibers continue distally. A second extensor aponeurosis

119
lies over the interphalangeal joint and the attachments are similar.
The medial tendon of insertion of the common digital extensor courses
across the dorsal surface of the carpus to insert on digit III in a
fashion similar to the tendinous attachments described for digit IV.
Innervation is by several branches of the deep branch of the radial
nerve. Distally, the deep branch runs along the deep surface of the
muscle belly. Topographically, the muscle bellies are superficial to
the extensor indicis, extensor pollicis brevis, and abductor pollicis
longus. Ulnaris lateralis lies caudal to the common digital
extensor.
Comparative aspects. There are usually four tendons of insertion
of the extensor digitorum (communis) of man although it is common to
find only three tendons, where the tendon located on the ulnar side
of the wrist gives off a small branch that fuses with the extensor
digiti minimi (=lateral digital extensor) (Hollinshead, 1974). The
latter condition is identical to my observations in A. jamaicensis.
The common digital extensor commonly inserts on digits II to V by
four tendons in rodents (Rinker, 1954; Klingener, 1964; Hill, 1937).
Woods (1972) observed a similar pattern of insertion but commented on
the variable interconnections present between the four tendons in the
carpus. Schon (1968) observed five tendinous insertions: one tendon
passed to each digit in Alouatta. The significance of this variation
is unclear.
Functional aspects. The function of the common digital extensor
in A. jamaicensis is to extend the caudal part of the carpus, digits

120
III to V, and to extend these digits at the carpometacarpal joints,
metacarpophalangeal joints, and interphalangeal joints. These
muscles probably are important to extend the digits and thus spread
the wing at the beginning of the downstroke, as proposed by Vaughan
(1959) and Altenbach (1979). If the common digital extensor is
active synchronously with the extensor carpi radialis longus, the
leading edge of the wing (digit II) is drawn craniad while the
trailing edge (digit V) is elevated and extended, thus spreading the
chiroptagium.
Extensor Indicis
Form. The origin of the extensor indicis is by fibers along the
craniodorsal surface of the ulna, 6-10 mm distal to the olecranon
process, and along the interosseous membrane and caudodorsal surface
of the middle third of the radius. The tendon of insertion passes
through the proximal and middle extensor retinacula, deep to the
compartment containing the tendons of the common digital extensor.
Extensor indicis inserts on the extensor process of metacarpal II.
The tendon continues along the craniodorsal surface of the metacarpal
for about 12 mm. The muscle belly is fused along an intermuscular
spetum with the belly of extensor pollicis brevis. Innervation is by
the deep branch of the radial nerve.
Comparative aspects. In Desmodus, an extensive insertion was
described by Altenbach (1979) to digits I through IV. Vaughan
(1970b) however, described a single insertion along the dorsum of
digit two on the extensor process on the base of the metacarpal.

121
In rodents, there is generally a single tendon to digit II
(Rinker, 1954; Klingener, 1964), although Woods (1972) noted a
collateral tendon coursing from the base of digit II to the terminal
phalanx of the pollex. Woods suggested that this collateral tendon
represented a vestige of the extensor pollicis brevis, a muscle
otherwise absent in these rodents. Both the extensor indicis and
extensor pollicis brevis are found in bats, representing the deep
layer of extensors on the pollical side of the forearm.
Functional aspects. The function of extensor indicis is to
extend the digit II and therefore to spread the chiropatagium for the
downstroke phase of flight (Altenbach, 1979). This muscle functions
as a synergist with the extensor carpi radialis longus.
Ulnaris Lateralis
The ulnaris lateralis is synonymous with the extensor carpi
ulnaris of domestic mammals (Evans and Christensen, 1979) and of
earlier bat descriptions (Vaughan, 1959).
Form. Ulnaris lateralis arises from the caudal surface of the
proximal 4 mm of the ulna. Fibers are arranged in parallel and
converge distally on a thin insertional tendon. The tendon courses
through the proximal and middle extensor retinacula, deep to the
common digital extensor. After crossing the dorsal surface of the
cuneiform, the tendon of ulnaris lateralis inserts in a groove on the
craniodorsal base of metacarpal V. Innervation is by the deep branch
of the radial nerve.

122
Comparative aspects. The proximal attachment of the ulnaris
lateralis is conservative in all bat species studied. The insertion,
however, exhibits variation among several species. In Macrotus and
in Desmodus the ulnaris lateralis inserts upon the base of metacarpal
five (Vaughan, 1959; Altenbach, 1979). In Antrozous, Plecotus,
Myotis, and Eumops the muscle inserts upon the dorsal surface of the
base of metacarpal three. The attachments are similar in other
mammals. The common position of insertion of ulnaris lateralis in
rodents is on metacarpal V (Woods, 1972).
Functional aspects. The insertional tendon of ulnaris lateralis
approaches the base of metacarpal V along a vector caudal to the
radiocarpal joint and the carpometacarpal joint of the digit V.
Thus, tension in the muscle will flex metacarpal V upon the radius.
Ulnaris lateralis is the only muscle of the antebrachium, innervated
by the radial nerve, to perform a flexor function. Vaughan (1959)
and Norberg (1970) claimed that the extensor carpi ulnaris was an
extensor of the metacarpal V in Macrotus and Plecotus, respectively.
They did not, however, provide a mechanical explanation for their
observations. The tendon of insertion courses sufficiently caudal to
the carpus to effect a weak flexion of the metacarpal. Because the
wing membrane connects the four lateral digits, flexion of digits II
through IV will also result when digit V is flexed. Flexion of the
wing occurs during terrestrial locomotion and roosting, and
transiently during the upstroke of flight.

123
Antebrachial Flexor Group
Flexor Carpi Radialis
Form. This muscle originates by fibrous attachments along the
medial surface of the proximal radius, and by fibers arising from the
surface of the pronator teres, exclusive of the proximal 8 mm of the
latter muscle. Flexor carpi radialis inserts by a thin tendon on the
ventral base of metacarpal II. The muscle is pinnate: the belly
extends about 20 mm distal to the elbow joint. Innervation is by the
median nerve.
Comparative aspects. The muscle is well developed in Macrotus
but is vestigial in Eumops and Myotis (Vaughan, 1959). In Macrotus,
the muscle inserts on the base of metacarpal III. The attachment of
the insertional tendon is on the base of metacarpal II in Rousettus
(Norberg, 1972) and Desmodus (Altenbach, 1979).
Functional aspects. Vaughan (1959) believed that the insertion
of flexor carpi radialis on metacarpal III provides the mechanical
arrangement necessary to flex the chiropatagium. Because of the
attachment of the wing membrane between the digits, flexion of
metacapal III would snychronously flex metacarpal II, the leading
edge of the wing. Similarly, flexion of the chiropatagium would be
achieved by the flexion of digits III through V by the flexor
digitorum profundus. The direct attachment of flexor carpi radialis
on digit II could provide additonal control of the leading edge of
the wing as well as control over the folding of the wing that is
necessary during terrestrial walking behavior (Altenbach, 1979).

Figure 20.—Medial view of muscles of the elbow region. The superficial
layer of muscles is exposed.

biceps brachii
pronator teres
flexor carpi
radialis
triceps brachii
—- ulnar n.
median n.
palmaris longus ''
flexor carpi ulnaris
ulnaris lateralis 7
5 mm

Figure 21.—Ventral view of the muscles of the carpal region,
intrinsic muscles of the manus have been removed.
The

abductor pollicis longus
flexor carpi radialis
, flexor digitorum
profundus
'' flexor carpi ulnaris
'' ulnaris lateralis
mm
127

128
The latter function may not be a specialized role or indication of
advanced structure. Attachment of the flexor carpi radialis on
digits II and III is common in many mammals (Woods, 1972).
Pronator Teres
Form. Pronator teres originates by a tendinous attachment on the
proximal aspect of the spinous process of the humerus. Insertion of
the muscle is by fibrous attachments along the ventral surface of the
radius, extending 24 mm distal from the elbow joint. Innervation is
by the median nerve.
Comparative aspects. The attachments of this muscle are similar
in all bats (Vaughan, 1959, 1970b). The muscle is only a weak
stabilizer of the elbow because of the limited pronation and
supination possible at this joint.
Functional aspects. The muscle is primarily a ventral stabilizer
of the humeroradial articulation. Some pronation of the radius is
possible because of the potential for rotatory movements of the
radial head on the capitulum of the humerus. In bats with elbow
movement restricted to flexion or extension, the pronator teres is
relatively small (Vaughan, 1959).
Palmaris Longus
Form. The origin of the palmaris longus is by a short tendon to
the spinous process of the humerus, and by fibers from the caudal
surface of a broad sheet of connective tissue on the proximal flexor
carpi radialis. The belly of palmaris longus extends 26-30 mm along
the radius. The insertional tendon bifurcates over the carpus. The
medial tendon courses to the ventral base of the metacarpal I.

129
The lateral tendon passes lateral to and inserts on the lateral
aspect of metacarpal I. Innervation is by the median nerve.
Comparative aspects. The palmaris longus has been described in
all bats studied except Myotis (Vaughan, 1955, 1970a). In Eumops,
the muscle inserts on the deep palmar fascia, on the thumb pad, and
on the surface of the abductor digiti quinti (Vaughan, 1959).
Extensive insertions were also described in Rousettus (Norberg,
1972), Plecotus (Norberg, 1970), and Desmodus (Altenbach, 1979). In
these bats, the palmaris longus attached upon digits I, II, III, and
V.
Functional aspects. The palmaris longus of Artibeus contributes
to flexion of the digit I along with the flexor digitorum profundus.
Ventral flexion of the pollex during slow flight increases the camber
of the middle wing sections and increases lift. This function is
most apparent in Artibeus, a bat that often flies slowly while
carrying large weight.
Flexor Carpi Ulnaris
Form. The flexor carpi ulnaris originates by two heads: one
head arises from a tendinous attachment on the distal tip of the
spinous process of the humerus, distal to the origin of the flexor
digitorum profundus and palmaris longus: a few fibers arise from the
proximal 8 mm of the ventral surface of the ulna. The two heads fuse
along the proximal third of the radius and the common belly extends
about 20 mm distal to the elbow joint. The muscle inserts by

130
a tendon on the palmar surface of the pisiform. Innervation of both
heads is by the ulnar nerve.
Comparative aspects. The attachments of this muscle are similar
in all species studied. Variation is observed primarily at the
proximal end of the muscle. The origin of flexor carpi ulnaris in
Macrotus includes attachments to both the radius and ulna (Vaughan,
1959). In Desmodus, the muscle originates from the ulna and the
surface of palmaris longus (Altenbach, 1979). Attachments of muscle
in Rousettus (Norberg, 1972) are identical to those of Artibeus.
Bats that are highly specialized for high-speed flight, such as the
molossids, usually possess a long projection on the distal aspect of
the spinous process. This bony modification, along with reduction in
the mass of muscle fibers inserting on the tendon of flexor carpi
ulnaris, permit an automatic mechanism for flexion of the manus when
the elbow is flexed during flight or roosting (Vaughan, 1959, 1966).
Variation in the origin of the flexor carpi radialis is common in
rodents (Woods, 1972). The primitive condition in mammals is for the
muscle to originate from the medial epicondyle, (Hill, 1937).
Functional aspects. On the basis of size and mechanical
advantage, I conclude that flexor carpi ulnaris is the major flexor
of the chiropatagium. The insertion of the flexor carpi ulnaris is
on the pisiform, a bone firmly bound to the adjacent carpal bones and
to the proximal base of metacarpals IV and V. Tension exerted upon
the pisiform is transmitted indirectly to the metacarpals and

131
folds the wing during roosting, walking, or during the middle
portions of the upstroke. The muscle probably acts synergistically
with the ulnaris lateralis.
Flexor Digitorum Profundus
Form. This deep digital flexor muscle originates by three
heads. The humeral head takes origin by a thin tendon from the
medial epicondyle between the origins of pronator teres and flexor
carpi ulnaris, and by tendinous slips from the deep surface of the
tendon of palmaris longus. The fibers are arranged in parallel. The
radial head arises by fibers along the ventral surface of the
proximal 22 mm of the radius. The fibers are unipinnate. The ulnar
head arises by fibers along the proximal 24 mm of the ventral surface
of the ulna. The fibers are unipinnate in their attachment upon the
main tendon of insertion. The three heads fuse along the proximal
third of the radius. The tendon passes deeply through the flexor
retinaculum of the carpus before bifurcating. One branch of the
insertional tendon attaches on the bases of phalanges 1 and II of the
pollex. The second tendon attaches on the ventromedial base of the
second phalanx of digit III. Innervation of the three heads of
flexor digitorum produndus is by the median nerve.
Comparative aspects. The flexor digitorum profundus of bats has
a reduced insertion on the digits relative to terrestrial mammals.
In many rodents, the muscle inserts by four heads on digits II
through V (Klingener, 1964; Woods, 1972). Innervation of the muscle
is by branches of the median nerve to all heads in cricetine (Rinker,

132
1954) and dipodoid rodents (Klingener, 1964). In hystricognathous
rodents, however, the ulnar head of flexor digitorum produndus in
innervated by the ulnar nerve while the remaining heads are
innervated by the median nerve (Woods, 1972). The innervation of all
three heads is derived from the median nerve in bats (Vaughan,
1959). The only branches of the ulnar nerve that I observed in the
proximal antebrachium in Artibeus provide motor innervation to the
flexor carpi ulnaris.
In other bats, the insertion of flexor digitorum profundus is on
digits I and III in Desmodus (Altenbach, 1979) and Macrotus (Vaughan,
1959). In Myotis, the insertion is on digits I, III, and IV; and in
Eumops the insertion is on digits I and V (Vaughan, 1959).
Rousettus, a megachiropteran, exhibits an insertion of the muscle on
digits I, II, and III (Norberg, 1972).
Functional aspects. Insertion on digit III provides a direct
line of pull for this muscle to flex the digit upon the radius and
thus fold the wing during roosting or terrestrial locomotion
(Altenbach, 1979). During the downstroke, flexion of the digit III
is important to prevent dorsiflexion of the digit and loss of wing
camber. Additionally, flexion of the chiropatagium during the late
upstroke reduces the amount of drag on the wing as it is returned to
a dorsal position for the ensuing downstroke.

ELECTROMYOGRAPHY IN FLYING BATS
Electromyographic data for 17 shoulder and arm muscles in
Antrozous indicated that muscles could be characterized as adductors,
abductors, or bifunctional muscles (Eermanson and Altenbach, 1983).
This classification paralleled the flexor, extensor, and bifunctional
categories proposed for terrestrial mammals by Engberg and Lundberg
(1969). Electromyographic data obtained in the present study do not
present such a clear functional classification for 15 of these
muscles. Data are not available for two muscles in Artibeus, the long
and short heads of biceps brachii. In Artibeus, two phases of activity
are demonstrated in 10 muscles, eight of which were uniphasic in
Antrozous (Hermanson and Altenbach, 1983). In this chapter,
differences in the pattern of muscle activity between Antrozous and
Artibeus are discussed, and data for both species are compared with
similar studies of locomotion conducted on terrestrial mammals.
There are several differences in the abductor muscle category
between Artibeus and Antrozous. In Artibeus, these muscles include the
clavotrapezius, acromiotrapezius, latissimus dorsi, teres major,
acromiodeltoideus, spinodeltoideus, and two heads of triceps brachii.
Two muscles are added to and one subtracted from the abductor category
in Antrozous (Eermanson and Altenbach, 1983). All of the abductors
133

134
except triceps brachii of Artibeus exhibit two periods of activity. A
primary EMG burst corresponds with the transitional period between the
downstroke and upstroke and is similar in timing to EMG activity
reported for abductors in Antrozous. In the biphasic abductors, a
secondary EMG burst, characterized by low-amplitude electromyograms,
occurred during the late upstroke and early downstroke, a period
appropriate for adductor function.
Latissimus dorsi and teres major both exhibit biphasic activity
cycles and are classified as abductors. Both exhibit single-phase
activity cycles in Antrozous and were classified as an adductor and a
bifunctional muscle, respectively (Hermanson and Altenbach, 1S83). In
Artibeus, a high-amplitude burst of activity in these muscles precedes
the beginning of the upstroke and coincides with the primary activity
of the other abductors. A synergistic function appears to exist
between these two muscles based upon their common insertion and similar
EMG profiles. Both contribute to arrest the cranial sweep of the wing
during the late downstroke and to power abduction of the wing during
the upstroke. The second burst of activity, immediately before and
during the early downstroke, provides the pronator action necessary to
position the wing for the downstroke. Similar to a mechanism proposed
in Desmodus, the combined action of latissimus dorsi and teres major of
Artibeus powers the flick phase, during which the wingtip is rapidly
pronated so that the leading edge of the wing faces ventrally.

135
The triceps brachii, long and lateral heads, exhibit uniphasic EMG
patterns in Artibeus. The activity of triceps brachii in Antrozous is
similar (Hermanson and Altenbach, 1983). In both species, the long
head commences activity prior to and continues for a longer duration
than the lateral head. Flexion of the elbow through the first
two-thirds of the upstroke was demonstrated in Plecotus by Norberg
(1976). A similar pattern of joint movement is present in Artibeus.
Thus, it is an enigma that triceps brachii is active primarily as an
abductor during a period of the wingbeat when the elbow is rapidly
flexed.
Spinotrapezius and infraspinatus both exhibit two periods of
activity in Artibeus. For both muscles, each period exhibits
electromyograms of approximately the same amplitude, thus it is
difficult to describe either as primary or secondary. I classify both
as bifunctional. Spinotrapezius is situated on the dorsum and is in a
position to draw the caudal angle of the scapula caudally and
medially. In Antrozous, spinotrapezius was uniphasically active and
was characterized as an abductor. Infraspinatus is located on the
infraspinous fossa and is situated to supinate the humerus and flex the
shoulder. Electromyograms in Antrozous were biphasic, indicating
activity during both the adductor and abductor periods (Hermanson and
Altenbach, 1983). Electromyography in Artibeus suggests that both
spinotrapezius and infraspinatus serve to abduct the wing during the
early upstroke. Coactivation of spinotrapezius and infraspinatus

136
with adductor muscles indicates that the interplay between these muscle
groups serves to smooth the transition from upstroke to downstroke, and
therefore protect the shoulder joint from disarticulation resulting
from the powerful activity of the adductors and the sudden forces
imparted to the wing when it first meets resistance from the airstream.
Three adductor muscles were identified in Antrozous, including
pectoralis, serratus ventralis thoracis, clavodeltoideus, and
latissimus dorsi (Hermanson and Altenbach, 1983). In Artibeus, only
three adductors are noted: clavodeltoideus, serratus ventralis
thoracis, and pectoralis. The activity of these three muscles is
similar to that observed in Antrozous except for the addition of a
low-amplitude period of activity in clavodeltoideus and serratus.
Pectoralis commences adductor activity slightly before serratus (0.781
versus 0.823) and both terminate activity during the early to middle
downstroke. The secondary burst of EMG activity in clavodeltoideus and
serratus occurs during the early upstroke, a period appropriate for
abductor activity. This secondary activity in both muscles probably
stabilizes the scapula against the forces developed by the primary
activity of the acromiotrapezius and clavotrapezius. Clavodeltoideus,
serratus, and subscapularis are the only ventrally situated muscles
active during the early upstroke.
Six muscles were studied in Artibeus that were classified as
bifunctional muscles in Antrozous (Hermanson and Altenbach, 1983).

137
Thej' included both heads of biceps brachii, supraspinatus,
infraspinatus, teres major, and subscapularis. No EMG data are
available for the biceps brachii of Artibeus. The spinotrapezius
represents an addition to the list of bifunctional muscles in
Artibeus. The spinotrapezius and infraspinatus of Artibeus exemplify
the bifunctional pattern. Activity is concentrated around the two
transition points of the wingbeat, the beginning of both the downstroke
and the upstroke. High-amplitude activity occurred during both of
these periods. The pattern in Antrozous was different for
spinotrapezius but was similar for infraspinatus (Hermanson and
Altenbach, 1983). Supraspinatus, a part of the rotator cuff complex
and a muscle that is situated in approximation with the infraspinatus
exhibits a different pattern of activity than was observed in
Antrozous. In Antrozous, two bursts of activity were confined to the
early upstroke phase (Hermanson and Altenbach, 1983). In Artibeus,
however, a single burst of activity begins during the middle upstroke
and continues throughout the early downstroke. The small sample size
(N=6) and the interruption in EMG activity observed during the burst in
two wingbeats suggest that the general pattern of activity in Artibeus
may not be significantly different from Antrozous. On the basis of
observed EMG activity, the spinotrapezius, infraspinatus, and
supraspinatus are active slightly before the major adductors, the
pectoralis and serratus ventralis thoracis. Their function is to
stabilize the shoulder joint and scapula against the larger forces
produced by these adductors.

138
Subscapularis is a difficult muscle to classify. In mammals, the
muscle forms the ventral component of the "rotator cuff," stabilizing
the shoulder joint and effecting pronation of the humerus (Hclliiishead,
1974; Tuttle and Basmajian, 1978). In bats, however, the subscapularis
attains the largest size relative to body mass observed in the class
Mammalia (Vaughan, 1959), an observation that led Vaughan to conclude
that the subscapularis functioned as a critical member of the
downstroke musculature as well as a shoulder stabilizer. In Antrozous,
subscapularis was found to be active throughout most of the wingbeat
cycle (Hermanson and Altenbach, 1981). A second study led these
authors to note a biphasic distribution of activity frequency.
However, Hermanson and Altenbach concluded that the muscle was
essentially active throughout the wingbeat cycle, as demonstrated by
the large variation in onset and termination times (Hermanson and
Altenbach, 1983). The muscle is not primarily an adductor, but
stabilizes the shoulder joint and provides for fine control of humeral
movements. The variation observed in level flight reflected the
facultative role of this muscle in the control of flight. In Artibeus,
subscapularis is active throughout the wingbeat cycle, and is quiescent
only during the third-quarter of the downstroke. This is similar to
data presented for Antrozous. High-amplitude activity occurred during
the last-third of the upstroke and during the early downstroke. This
high-amplitude activity must contribute to the dow'nstroke by pronating
the wing and maintaining the ventral orientation of the wing's leading
edge during the downstroke.

139
Different flight styles and wing shapes of Artibeus and Antrozous
may require differences in the basic neuromuscular program for flight.
Artibeus is a heavier bat than Antrozous. I converted Davis’s data
(1969) for wing loading in Antrozous into M-K-S system equivalents.
The mean wing loading in Antrozous was 12.67 N/m^, a value that
includes males and females throughout the annual cycle. Fall females
were found to exhibit relatively heavier wing loading values than
pregnant females or adult males. Estimates of wing loading in Artibeus
range from 16.60 N/m^ to 16.65 N/m^ (Eorberg, 1981). My
observations of wing loading values in two Artibeus females, 15.47 and
16.74 N/m^, represent extremes relative to these published accounts.
A range of males and females from different seasons is not available.
The wing loading values of Artibeus are larger than observed in
Antrozous, and indicates that the wing supports more weight per square
meter of wing surface. Additional muscular effort in the form of
secondary bursts of muscle activity, are necessary to control wing
position and shape with precision during the wingbeat of Artibeus. One
difference is the addition of a second phase of activity in eight
muscles of Artibeus that were uniphasic in Antrozous. The turnover
points of the wingbeat, at the end of the downstroke or upstroke, are
smoothed out by coactivation of abductors and adductors. Analysis of
the EMG profiles shows that the additional burst of activity found in

140
the eight additional biphasic muscles generally occur either at the end
of the downstroke or at the end of the upstroke. Because of the
elliptical path followed by the wingtip during the wingbeat cycle, the
transitions between the propulsive and non-propulsive phases are not as
clearly defined as they are in terrestrial stepping movements.
In review, a classification of the 15 muscles studied in Artibeus
includes eight abductors, three adductors, and four bifunctional
muscles. Four of the muscles are classified differently then they were
in Antrozous.
The classification of muscles as adductors, abductors, and
bifunctional muscles is an heuristic tool to analyze the contribution
of each group of muscles to the overall control of the wingbeat cycle.
The following discussion identifies the muscles, or muscle group, that
power specific portions of the wingbeat cycle.
The downstroke phase is preceded by increasing amounts of activity
in the adductor muscles. All three adductor muscles are active before
the end of the preceding wingbeat (by the .850 point). The adductors
remain active during the first one-fifth of the downstroke. Five of
the abductor muscles are active during the last one-fifth of the
upstroke or by the .900 point of the wingbeat cycle. Six abductor
muscles are active at the beginning of the downstroke (.000-.005).
Abductor activity decreases rapidly during the early downstroke and, at
.100, only two muscles, acromiodeltoideus (caudal part) and

141
spinodeltoideus are active. In all cases, the electromyograms of the
abductors during the upstroke to downstroke transition are of lower
amplitude than observed during activity associated with the end of the
downstroke. The spinotrapezius has a temporal pattern of activity that
is similar to the abductors, however, the electromyogram obtained at
the upstroke to downstroke transition is equal to the second burst
observed during the late downstroke. Therefore, spinotrapezius is
classified as a bifunctional muscle. Three additional bifunctional
muscles are active prior to and at the beginning of the downstroke:
they include the supraspinatus, infraspinatus, and subscapularis.
Spinotrapezius, supraspinatus, and infraspinatus are all situated on
the dorsal aspect of the shoulder girdle and should cause abduction of
the wing when active. Their contribution to the downstroke is
apparently one of joint stabilization. Subscapularis is situated
ventral to the shoulder joint and was proposed to be a major adductor
muscle (Vaughan, 1959). During the early downstroke, subscapularis
probably contributes to power the adduction of the wing.
The primary downstroke muscles in Artibeus include pectoralis,
serratus ventralis thoracis ventralis (caudal division), and
clavodeltoideus. These muscles cease activity early during the
downstroke and are quiescent during about four-fifths of the
downstroke. This absence of activity cannot be explained without
undertaking mechanical studies of the muscles. Rasmussen et al. (1978)
commented on the termination of EMG activity in all extensors of the

142
cat hindlimb prior to the end of the extensor phase (E3) during normal
stepping. These authors attributed the early termination of activity
to ’’unloading" of the limb, a relaxation of all extensor muscles,
before the subsequent flexion period begins. In Artibeus, no EMG
activity was recorded from abductors or adductors during the
second-quarter of the downstroke (.270-,470), except for the final
myopotentials observed in infraspinatus (until .120) and subscapularis
(until .174). The termination of adductor muscle activity some 30-40
msec prior to reversal of the wingbeat is puzzling and needs further
analysis.
The initiation of upstroke movements is characterized by
increasing activity in the abductor muscle group. Beginning with a
minimum of no muscle activity during the mid-downstroke, the .200 point
of the wingbeat cycle, the total number of abductor muscles showing
activity increases to seven by the last one-fifth of the downstroke
(the .400 point), and to a maximum of eight muscles by the time the
first upward movements of the wingtip are noted. The number of
coactive abductors decreases during the early upstroke, decreasing to
six muscles by the .600 point, and to two muscles at the .650 point.
Thus, the primary coactivation of abductor muscles occurs for about 15
msec before and 10 msec after the beginning of an upstroke. Two
adductor muscles are coactive with the abductors during the early
upstroke: the serratus ventralis throacis and the clavodeltoideus.
This coactivation of antagonists parallels the situation observed
during the transition between the downstroke and upstroke.

143
The most prominent pattern of muscular coactivation during the
chiropteran wingbeat cycle is the concentration of muscles, in absolute
numbers, that are active at the transitional points of the cycle. In
Artibeus, the maximal number of coactive muscles observed, 13, occurs
at the beginning of the downstroke (Figure 22). The frequency or
number of coactive muscles has a bimodal distribution, one peak being
at the beginning of the downstroke, the other at the beginning of the
upstroke when 12 muscles are coactive. Other intervals show decreasing
numbers of coactive muscles with none acting during the middle of the
downstroke, and a second minimum of two muscles during the upstroke.
Electromyograms of Antrozous during flight indicate a similar emphasis
of muscle activity around the transitional points of the wingbeat
cycle. The absolute number of coactive muscles is less than that
observed in Artibeus because the abductors do not develop secondary
periods of myopotentials (Hermanson and Altenbach, 1983).
The mode of support and propulsion in bats can be compared aptly
with those in terrestrial mammals. In terrestrial locomotion, several
muscles transmit the body weight to the limbs and therefore hold the
body off of the ground. Comparative anatomists recognized that the
serratus ventralis was ideally situated to function as a "sling” to
suspend the thorax between the forelimbs (Davis, 1949; Gray, 1968).
This "sling" hypothesis has been confirmed by electromyographical
studies which reveal intense activity of several regions of the
serratus ventralis thoracis during the stance phase in stepping

Figure 22.—Number of coactive muscles during the wingbeat
cycle: all muscles studied (o); abductors ( ); adductors
( ); bifunctionals ( ).

NUMBER OF ACTIVE MUSCLES
downstroke
upstroke
14
12
10
8
6
4
2
0
.00 .20 .40 .60 .80 1.0
4
2
0
WINGBEAT CYCLE

146
cats (English, 1978a) and opossums (Jenkins and Weijs, 1979). Jenkins
and Weijs (1979) suggested that the pectoralis transversalis and
rhomboideus also function to suspend the body between the forelimbs.
Flying mammals contend with similar problems during the propulsive
phase as the body must be supported between the two wings. Serratus
ventralis thoracis (caudal division), pectoralis and spinotrapezius are
all extrinsic muscles transmitting weight from the wing to the thorax
and are intensely active during the downstroke. Clavotrapezius and
acromiotrapezius also show activity, albeit low-amplitude activity,
during the propulsive or downstroke phase. All of these muscles appear
to transmit the weight of the body to the wings, or to stabilize
rotatory movements of the scapula along its long axis. No EMG data is
available for rhomboideus, however, in bats the rhomoboideus is a thin
sheet and may not be of great importance. Clavotrapezius and
acromiotrapezius are thick muscles and are ideally suited by virture of
their origin from the cervical vertebrae, 10-15 mm ventral to the
scapula to support the body between the forelimbs.
The intrinsic muscles of the shoulder include the supraspinatus,
infraspinatus, and subscapularis. Each of these crosses one joint, the
shoulder joint, and exhibit activity in bats during the transitional
periods between the upstroke and downstroke. The shallow profile of
the glenoid fossa provides poor stability for the shoulder joint except
for stability provided by the rotator cuff muscles. Compression of the

147
suprascapular nerve in horses, the sole motor nerve supply to both the
supraspinatus and infraspinatus, two of the four rotator cuff muscles,
results in subluxation of the joint during the stance phase of stepping
or during quiet standing (Rooney, 1969; Adams, 1974). In the opossum,
stability of the shoulder joint was provided by the almost synchronous
activity of supraspinatus, infraspinatus, and subscapularis during the
last half of the swing phase and throughout most of the propulsive
phase (Jenkins and Weijs, 1979). These muscles were active during the
El through E3 phases of the step cycle in cats (English, 1978a). In
both cases, activity during the El phase, or the first extension phase
prior to foot touchdown, indicates that the rotator cuff muscles are
also important in positioning the limb for the ensuing step.
Propulsion of the body is realized in a different fashion in bats
and in terrestrial mammals. During terrestrial locomotion, the body is
propelled forward through the forelimbs by action in two muscles,
caudal portions of latissimus dorsi and pectoralis (Tokuriki, 1973*1;
Jenkins and Weijs, 1979; English 1978a_). Kinematic analysis indicated
that the greatest contribution to propulsion is incurred by the
proximal part of the limb: movements of the scapula relative to the
thorax are of greater magnitude than are synchronous joint angle
changes in the elbow and carpal joints (Jenkins and Weijs, 1979).
Activity of the intrinsic limb musculature, therefore, contribute to
stabilization of the limb in order that the trunk musculature (i.e.,
latissimus dorsi and pectoralis) causes the body to pole vault

148
over the limb. Additionally, the pectoral limbs are thought to serve
primarily as weight-bearing organs while the pelvic limb provides most
of the forward thrust associated with locomotion in mammals (Manter,
1938). In bats the pectoral limbs provide both weight support and
propulsive forces. During the downstroke, the wing moves ventrally and
cranially. Because of its dorsal point of origin, the latissimus dorsi
lengthens during the propulsive phase, the downstroke. In Artibeus,
latissimus dorsi is classified primarily as an abductor. The muscle is
intensely active during the upstroke, and active as a wing pronator
during the downstroke. Thus, latissimus dorsi has a different role in
flying bats than is commonly required during stepping in mammals.
Vaughan (1959) and Altenbach (1979) noted the significance of a
relatively large latissimus dorsi in bats that spend large amounts of
time foraging on the ground. It would not be surprising to observe
primary propulsive activity in the latissimus dorsi during walking in
bats if EMG studies are conducted. The pectoralis is active as an
adductor in Artibeus. Because of the ventral origin of this muscle,
the muscle is ideally situated as a wing adductor. Shortening of the
muscle during the wingbeat may amount to only 10 percent of the
muscle's maximum length, permitting the muscle to operate within a
range of high efficiency of force production (Hermanson, 1981).
Pectoralis exhibits temporal patterns of activity that are similar to
those observed in terrestrial mammals.

CONCLUSIONS
Comparison of the activity patterns of homologous flight muscles
in two distantly related bats Artibeus jamaicensis and Antrozous
pallidus, indicates divergent activity correlated with their different
flight characteristics. These bat species are similar in the gross
appearance of skeletal and muscular systems despite their divergent
familial affinities. Their flight styles reflect adaptation to their
respective feeding behaviors. Artibeus is frugivorous and flies
directly to and from food sources at low to medium speed while moving
over short distances. The relatively heavy wing loading values
observed in Artibeus compromise their maneuverability. In contrast,
Antrozous feeds upon insects in aerial, arboreal, and terrestrial
niches. They are capable of slow, fluttering flight while foraging but
can travel long distances nightly during flights of three or four
hours. The temporal patterns of muscle activity recorded during EMG
experiments on the two species differ, just as do their flight styles.
These data, interpreted in light of the differences noted in the flight
style of both species, suggest the use of caution in speculation about
the neuromuscular control of flight in other bats that utilize
divergent or similar flight styles.
149

150
Electromyograms obtained from eight of the 15 muscles studied in
Artibeus exhibited a biphasic pattern not recorded in the same muscles
in Antrozous. These observations of biphasic muscles could relate to
increased stabilization roles encountered in Artibeus. I argued (see
preceding chapter) that the heavier wing loading of Artibeus, relative
to Antrozous, necessitates increased muscle activity at the turnover
points of the wingbeat cycle. This additional activity maintains a
smooth transition in changing the direction of wing movements. Most of
the muscles in Artibeus exhibited a high-amplitude period of activity
that is similar in timing to the single phase of activity that was
identified in the same muscles of Antrozous. Electromyographic studies
on other chiropteran species that are specialized for different flight
styles or that exhibit skeletal specializations may provide data useful
in evaluating these ideas.
Morphological studies conducted on locomotion in several
terrestrial mammals emphasize the facultative nature of stepping (cf.
Rasmussen et al., 1978). The variation in the neuromuscular control of
bat flight is also demonstrated by the data presented for the
individual muscles of Artibeus. It is possible to compare homologous
muscles in the wings of flying bats and to the limbs of other mammals
while stepping with the aid of the statistical means of activity onset
and termination. For example, bats support the body in a fashion
similar to terrestrial mammals: the body is supported between the two
pectoral limbs by the serratus ventralis thoracis

151
muscles, and partly by the pectoralis and trapezius muscles. The
recruitment of muscles during the propulsive phase of the locomotor
cycle is different in the two groups, however. In bats, the propulsive
or downstroke phase is powered primarily by the pectoralis and serratus
ventralis thoracis musculature. In cats and dogs, propulsion is
largely realized when the latissimus dorsi and caudal portions of the
pectoralis pull the body cranially relative to the fixed pectoral
limbs. Also, the activity cycle of the propulsive muscles is more
rapid in bats than in most terrestrial mammals. Correlation of these
functional data with further studies on the physiological
specializations of bat muscles will yield insights into the range of
functions possible for mammalian muscle tissue.
The pattern of muscle recruitment and activity in Antrozous is
simple relative to the complex pattern observed in many of the muscles
of Artibeus. The flight muscles of Antrozous can be classified as
abductors, adductors, or bifunctionals, a scheme that parallels the
extensor, flexor, or bifunctional classification proposed for
locomotory muscles in cats during overground stepping. The dichotomy
between abductors and bifunctionals is not clear in Artibeus EKG
recordings where the biphasic patterns are most common. A correlation
exists between the complexity of the muscle activity patterns and
musculoskeletal structure. Antrozous is a typical vespertilionid bat
and possesses a well developed shoulder locking mechanism. This

152
locking mechanism provides a skeletal constraint on wing movement.
Vespertilionids are derived chiropterans and are specialized for
flight. In contrast, Artibeus is a typical phyllostomid and possesses
a poorly developed shoulder locking mechanism. Phyllostomids are more
primitive chiropterans and exhibit generalized flight characteristics.
The more complex activity patterns of Artibeus, relative to Antrozous,
reflect the need for more gross muscular control of the wingbeat cycle,
particularly during the transition between the upstroke and downstroke.

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BIOGRAPHICAL SKETCH
John W. Hermanson was born in Quincy, Massachusetts, the son of
John and Mary Hermanson. He attended the University of Massachusetts,
at Amherst, and received the degree of Bachelor of Science in Zoology
in 1975. He studied in the Department of Biological Sciences at
Northern Arizona from August 1975 through August 1977. The degree of
Master of Science was awarded to him in 1978 after completion of his
thesis, "The forelimb morphology of the pallid bat (Antrozous
pallidus)." In September 1977, he entered the graduate program at the
University of Vermont to study with Dr. Charles Woods. He went with
Dr. Woods to the University of Florida in 1980 and enrolled there as a
graduate student in the Department of Zoology. During his tenure at
the University of Florida, his primary duties were as a graduate
teaching assistant in Anatomy in the College of Veterinary Medicine.
160

I certify that I have read this study
it conforms to acceptable standards of scholarly
is fully adequate, in scope and quality, as
degree of Doctor of Philosophy.
Associate
I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and
is fully adequate, in scope and quality, as a dissertation for the
degree of Doctor of Philosophy.
Associate Professor in Zoology
I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and
is fully adequate, in scope and quality, as a dissertation for the
degree of Doctor of Philosophy.
Donald A. Dewsbury
Professor in Psychology
I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and
is fully adequate, in scope and quality, as a dissertation for the

I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and
is fully adequate, in scope and quality, as a dissertation for the
degree of Doctor of Philosophy.
Ronald G. Wolff v
Associate Professor in Zodlogy
This dissertation was submitted to the Graduate Faculty of the
Department of Zoology in the College of Liberal Arts and Sciences
and to the Graduate School, and was accepted as partial fulfillment
of the requirements for the degree of Doctor of Philosophy.
Dean for Graduate Studies and
Research
April 1983

UNIVERSITY OF
FLORIDA