CRANIOMANDIBULAR STRUCTURE AND FUNCTION IN MORMOOPID BATS

A Thesis

Presented to the faculty of the Department of Biological Sciences

California State University, Sacramento

Submitted in partial satisfaction of the requirements for the degree of

MASTER OF SCIENCE

in

Biological Sciences

by

Jeffrey B. Changaris

FALL 2017

© 2017

Jeffrey B. Changaris

ALL RIGHTS RESERVED

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CRANIOMANDIBULAR STRUCTURE AND FUNCTION IN MORMOOPID BATS

A Thesis

by

Jeffrey B. Changaris

Approved by:

______, Committee Chair Dr. Ronald M. Coleman

______, Second Reader Dr. Winston C. Lancaster

______, Third Reader Dr. Joseph Bahlman

______Date

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Student: Jeffrey B. Changaris

I certify that this student has met the requirements for format contained in the University format manual, and that this thesis is suitable for shelving in the Library and credit is to be awarded for the thesis.

______, Graduate Coordinator ______Dr. James W. Baxter Date

Department of Biological Sciences

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Abstract

of

CRANIOMANDIBULAR STRUCTURE AND FUNCTION IN MORMOOPID BATS

by

Jeffrey B. Changaris

Neotropical Ghost-Faced bats of the genus Mormoops (Order Chiroptera, Family

Mormoopidae) have a radically upturned rostrum, or snout, while the other mormoopid genus, Pteronotus, has only a slight upturning of the rostrum. This type of difference in morphology between closely related taxa is likely to be the result of some sort of specialization. Observation of Mormoops blainvillei, the Antillean Ghost-Faced bat, reveals that they can open their mouths very wide relative to the size of their heads.

Mormoopid bats are insectivorous with Mormoops blainvillei having a prey preference of large moths, but related species, such as Pteronotus quadridens, the Sooty Mustached bat, have a more varied diet with a large component of smaller hard-bodied beetles.

While there have been many studies on feeding ecology of phyllostomid bats, available research on mormoopids is limited and functional relationships between craniorostral shape and feeding mechanics have not been established. The objectives of this study were to quantify structural and functional differences in craniomandibular function between Mormoops blainvillei and Pteronotus quadridens by analyzing maximum gape, bite force, and masticatory muscle configuration and relate this to prey preference

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previously described in the literature. Results showed that Mormoops blainvillei had a much wider maximum active gape and reduced normalized maximum bite force than

Pteronotus quadridens which corresponded to the size and hardness of their respective diets. Mormoops blainvillei had a greater percentage of masticatory musculature allocated to wide gape than Pteronotus quadridens where the narrow gape chewing muscles were favored. Upturning of the rostrum reduces range of motion limitations at the craniomandibular joint which could have resulted in the ability to achieve wider gapes in Mormoops blainvillei.

______, Committee Chair Dr. Ronald M. Coleman

______Date

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ACKNOWLEDGEMENTS

First, I would like to thank Dr. Winston C. Lancaster without whose guidance, assistance, companionship, camaraderie, perseverance, and patience this endeavor would have never reached this successful conclusion; I am forever in your debt. I would like to thank Dr. Ron Coleman for assuming the leadership role as committee chairman once Dr.

Lancaster moved on in his worldly endeavors. I would also like to thank the rest of my committee, Dr. Rosalee Sprowls, Dr. Rodney Imamura, Dr. Joseph Bahlman, for their patience and valuable input, as well as the graduate committee, Dr. Jamie Kneitel, Dr.

James Baxter, and Dr. Robert Crawford. Thanks go to Dr. Tom Schultze for his mathematical advice, Drs. Rafael Diaz and Jamie Kneitel for their statistical advice, Dr.

Tim Davidson for his manuscript review and input, and Sulie Ober for her laboratory support.

Special thanks go to Dr. Armando Rodríguez-Durán for providing access to Mata de Plátano Field Station and Culebrones Cave in Puerto Rico, and Dr. Betsy Dumont for her bite force assistance and loan of the bite force meter used during field work. Partial funding from California State University, Sacramento for travel to Puerto Rico and equipment came from a Research and Creative Activity (RCA) Award granted to Dr.

Lancaster, Academically Related Activities (ARA), Student Academic Development

(SAD) awards and Delisle funding granted to the author.

I would also like to thank the following bat biologists for their support and assistance over the years and from whom I learned so much: Dr. Scott Pedersen, Dr. Gary

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Kweiznewski, Dr. Rick Adams, Dr. Allen Kurta, Ashley Rolfe, David Wyatt, and Linda

Angerer. Major thanks go to Justin Moore for his motivational participation and manuscript review. I would like to thank my colleagues, Marcie Woolsey, Dean Won,

Michael Sweet, Ken Kubo, Daniel Slutsky, Tom Peavy, Tom Landerholm, and D.J.

Larkey for their support. Lastly, I thank my friends and family, Sherry Kimbrow,

Danielle Delucchi, Stephanie Changaris, and Sherry Swim for their continued support.

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TABLE OF CONTENTS

Acknowledgements ...... vii

List of Tables ...... xi

List of Figures ...... xii

INTRODUCTION ...... 1

Hypotheses and Objectives ...... 6

METHODS ...... 7

Field Data Collection ...... 7

Gape Analysis ...... 13

Jaw Scaling ...... 18

Bite Force Analysis ...... 22

Jaw Musculature ...... 26

Statistical Analyses ...... 30

RESULTS ...... 33

Gape Analysis ...... 33

Bite Force Analysis ...... 38

Jaw Musculature ...... 38

DISCUSSION ...... 45

CONCLUSIONS...... 51

Appendix A ...... 52

Appendix B ...... 68

Appendix C ...... 72

Appendix D ...... 74

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Appendix E ...... 75

Literature Cited ...... 77

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LIST OF TABLES

Table Page

1. Total Number of Mormoopid Bats Analyzed by Species ...... 34

2. Total Video Frames and Bite Force Measurements Analyzed for Two Species of Mormoopid Bats ...... 35

3. Maximum Active Jaw Gapes for Two Species of Mormoopid Bats ...... 36

4. Maximum Bite Forces for Two Species of Mormoopid Bats ...... 39

5. Craniomandibular Muscle Architecture Measurements of Mormoopid Bats by Sex and Species...... 41

6. Craniomandibular Muscle Organization for Two Species of Mormoopid Bats ...... 42

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LIST OF FIGURES

Figure Page

1. Mormoopid Bats of Interest ...... 2

2. Rostral Orientations of Two Species of Mormoopid Bats ...... 4

3. Map of Puerto Rico showing the location of Culebrones Cave ...... 8

4. Active Gape Video Recording Apparatus ...... 10

5. Bat Position for Active Gape Video Recording ...... 11

6. Bite Force Measurement Apparatus...... 12

7. Video Gape Measurement Landmarks...... 15

8. Video Gape Jaw Relationships ...... 17

9. Jaw Scaling Measurement Landmarks...... 20

10. Landmark Relationships for Jaw Scaling ...... 23

11. Bite Force Meter Lever Properties ...... 25

12. Superficial Masticatory Musculature ...... 28

13. Deep Masticatory Musculature ...... 29

14. Average Maximum Active Jaw Gape Distance and Angle for Two Species of Mormoopid Bats ...... 37

15. Average Maximum Actual and Normalized Bite Forces for Two Species of Mormoopid Bats ...... 40

16. Masticatory Muscle Distribution in Two Species of Mormoopid Bats ...... 44

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INTRODUCTION

Feeding ecology in mammals is important for understanding species

diversification and encompasses many variables such as diet and feeding mechanics,

including cranial morphology and bite force. Structure of the feeding apparatus should relate to prey choice. Unique morphological structures usually reflect some sort of specialization (Wainwright and Price 2016). Mormoops, or Ghost-Faced Bats, have a unique skull shape that includes a radically upturned rostrum, or snout. It is readily observable that Mormoops can open their mouths much wider, relative to the size of their heads, than confamilial species of Pteronotus, the Mustached Bats, whose rostrum is only slightly upturned. This study will attempt to quantify structural and functional differences in the masticatory morphology between Mormoops blainvillei, the Antillean

Ghost-Faced bat, and Pteronotus quadridens, the Sooty Mustached bat, in relation to how wide they can open their mouths (gape) and how hard they can bite (bite force).

Mormoopidae is a Neotropical family of bats, Order Chiroptera, that diverged from the phyllostomid, or leaf-nosed bats in the latter Eocene, somewhere around 38 million years ago (Jones et al. 2005). Mormoopids consist of two genera, Mormoops and

Pteronotus. Several mormoopid species dispersed to the Greater Antilles from Central

America presumably during the Miocene (Dávalos 2006). Mormoops blainvillei

(Lancaster and Kalko 1996) and at least two species of Pteronotus, including Pteronotus quadridens (Rodríguez-Durán and Kunz 1992), are geographically isolated to the islands of the Greater Antilles in the Caribbean Sea as of the late Pliocene (Smith 1972). M. blainvillei and P. quadridens will be the model organisms of this study (Figure 1).

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Figure 1. Mormoopid Bats of Interest. Mormoops blainvillei (left) and Pteronotus quadridens (right).

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Much work has been done on individual components of feeding ecology for phyllostomid bats and while several studies have analyzed bite force and diet (Aguirre et al. 2002; Dumont and Herrel 2003; Santana and Dumont 2009; Santana et al. 2012;

Santana 2016), cranial morphology is also thought to play a major role in feeding mechanics (Dumont and Herrel 2003). The major morphological components of the feeding apparatus are the muscles of mastication, supporting bony structures, including the craniomandibular joint, and the teeth. Analysis of bite force and masticatory muscles are presented by Herrel et al. (2008) and Santana et al. (2010) for phyllostomid bats, while Nogueira and colleagues (2009) attempted to correlate cranial and mandibular bony morphology to published bite force data and diet using geometric morphometric analysis for phyllostomid bats. Freeman (2000) analyzed phyllostomid craniodental relationships.

However, functional relationships between craniorostral shape and feeding mechanics have not been established.

Research on feeding mechanics in the Mormoopidae is limited with only morphological characters and diet having been described. All mormoopid skulls exhibit an upturning of the rostrum which is only slight in Pteronotus but much more radical in

Mormoops (Smith 1972; Simmons and Conway 2001) (Figure 2). Pteronotus sp. appear to be seasonally opportunistic feeders whose diet includes beetles (Coleoptera) and moths

(Lepidoptera) while Mormoops preferentially feed on moths (Lepidoptera) (Simmons and

Conway 2001; Rolfe and Kurta 2013). In general, Lepidoptera tend to be consistently large, averaging around 19mm in length, based on studies of insectivorous bat diets for a similar neotropical insect assemblage from Esporitu Bolivia (Aguirre et al. 2003) while

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Figure 2. Rostral Orientations of Two Species of Mormoopid Bats. Mormoops blainvillei (left) exhibits a greater degree of rostral upturning than Pteronotus quadridens (right) relative to the basicranial axis. Right lateral views of skulls with pins glued along the basicranial axes to achieve a standardized orientation.

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Coleoptera tend to be highly variable in size but substantially harder than Lepidoptera

(Freeman and Leman 2007; Ober and Hayes 2008).

Both Mormoops blainvillei (Antillean Ghost-Faced bat) and Pteronotus quadridens (Sooty Mustached bat) have a similar skull length ranging between 12-14mm which is unusual because the overall size of M. blainvillei (6-11 g) is roughly twice that of P. quadridens (3-6 g) (Simmons and Conway 2001). The ability to achieve a larger gape could benefit M. blainvillei by allowing them to capture their prey of choice, moths, with a body length in excess of their own head length. However, wider gape could result in a tradeoff of bite force reduction because limits in jaw function accompany changes in jaw structure (Wainwright 1994; Wainwright and Price 2016). The question arises whether the difference in the rostral angle of the skull between the strongly upturned rostrum of M. blainvillei and the slightly upturned rostrum of P. quadridens functionally affects feeding mechanics. Bony morphology provides joint levers and surfaces for muscle attachment so maximum active gape (how wide can they open their mouth) and bite force (how hard can they bite) should relate to the configuration of the muscles of mastication. These muscles can be grouped into jaw abductors (openers) and adductors

(closers). Jaw adductor muscles can be further subdivided into two groups: those that operate better at narrow gapes, and those that operate well at wider gapes (Turnbull 1970;

Herring and Herring 1974; Freeman 1979).

This study attempts to quantify the difference masticatory musculature, maximum gape size, and bite force between M. blainvillei and P. quadridens. This research expands limited data available on mormoopid bats while elucidating how certain specific

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aspects of craniorostral morphology may affect feeding mechanics. Analysis of M. blainvillei and P. quadridens craniomandibular morphology and corresponding musculature should identify morphological similarities and differences such that corresponding function may be determined.

Hypotheses and Objectives

Hypothesis 1: It is predicted that Mormoops blainvillei has a greater maximum

normalized active gape than Pteronotus quadridens measured in millimeters between the

bases of upper and lower canine teeth and converted to angle in degrees at the

craniomandibular joint.

Hypothesis 2: It is predicted that Pteronotus quadridens has a greater normalized

bite force than Mormoops blainvillei measured in newtons at canine occlusion divided by

skull proportion in cubic centimeters where skull proportion = head length * head width *

head height.

Hypothesis 3: It is predicted that the Mormoops blainvillei jaw musculature

distribution will have greater jaw abducting (opening) and wide-gape adducting (closing)

muscle group proportions than Pteronotus quadridens as quantified by the proportions of

the physiological cross sectional areas of the abductor, wide-gape adductor, and narrow-

gape adductor muscle groups to the total jaw musculature.

The objective of this study is to quantify and compare feeding characters of gape,

bite force, and jaw muscle distribution between two species of mormoopid bats and

explain the differences in terms of their feeding ecology.

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METHODS

Field Data Collection

Bats were collected outside Culebrones Cave at the Mata de Plátano Field Station

near Arecibo, Puerto Rico (Figure 3) over an eleven-day period at the end of May, 2010 using harp traps (Kunz and Kurta 1988) at the cave entrance at dusk. Adults of each species, Mormoops blainvillei and Pteronotus quadridens, were captured to obtain a minimum sample size of twenty of each species with equal numbers of males and females. Bats were held individually in cloth bags until processed. Video recordings were made of individuals that were willing to display biting behavior using minimal gentle coercion (details below). Bite forces were measured for individuals willing to bite the bite force meter (details below). Individuals from which usable data were collected were sexed and measured when possible, then released. Multiple discreet videos were recorded per bat separated by a minimum resting period of one minute. Five bite force readings per individual were attempted separated by a one minute minimum resting period. Specimens were weighed using a digital scale (Ohaus HH320 320gx0.1g; Ohaus

Corp.; Parsippany NJ, USA) and measured for body length, forearm length, head width, head height, and head length using an electronic digital caliper (Pro-Max 0-150mm

±0.02mm; Fred V. Fowler Co, Inc; Auburndale, MA, USA). Handling of live animals adhered to the protocol for animal care and use submitted to and approved by the

Institutional Animal Care and Use Committee (IACUC) at California State University,

Sacramento.

Videos were taken of bats actively opening and closing their mouths. The camera

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Figure 3. Map of Puerto Rico showing the location of Culebrones Cave (A), approximately 7 km southwest of Arecibo. Image from Google Maps.

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was positioned 2.0 meters from and at the same height as a millimeter scale aligned in the

same plane as the craniomandibular joint of the biting bat (Figure 4). The bats were held

from behind and gently coerced as necessary to induce active biting (Figure 5). Videos included several maximal jaw openings from both left lateral and frontal views. Digital video was recorded using a Canon EOS 500D digital camera (Canon USA, Inc.; Lake

Success, NY, USA) equipped with a 32 gigabyte Class 6 secure digital (SD) memory card and a Canon EF 75-300mm F 1:4-5.6 zoom lens (Canon USA, Inc.; Lake Success,

NY, USA) mounted on a tripod. The lens was set to 200mm zoom (F5.6 ISO 1600) and

manual focus mode. Recording mode was set to high definition (HD) 1280x720 pixel

resolution at 30 frames per second (FPS). Prior to each recording, focus was manually

set using the graduations on the scale as the focal point. Videos were recorded in the

field. Available ambient light was used as the primary lighting source augmented by

headlamps and a handheld flashlight. Videos were downloaded to a laptop computer,

stored in QuickTime movie format (.MOV), named using specimen id and sequence

number, and logged.

Bite force was measured using a bite force meter containing a piezoelectric force

transducer (Kistler, type 9203, range ±500·N; Amherst, NY, USA) and a handheld charge

amplifier (Kistler, type 5995) accurate to 0.01 newtons (N) (see Dumont and Herrel

2003) (Figure 6). Adjustable distance between bite plates on the bite force meter was set

to its minimum distance of 2mm using the built-in micrometer. Metal bite plates were

wrapped in cloth tape to protect the bat’s teeth. Bite forces were taken bilaterally

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Figure 4. Active Gape Video Recording Apparatus. Camera positioned two meters from and level with a metric scale attached to a ring stand. Scale aligned in the focal plane of the camera for recording bat biting behavior. Photo by W. Lancaster.

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Figure 5. Bat Position for Active Gape Video Recording. Bat held in the plane of the millimeter scale and gently coerced to induce active biting behavior. Left lateral view of Mormoops blainvillei.

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Figure 6. Bite Force Measurement Apparatus. Bite force gauge with force transducer on wooden stand (right) connected to charge amplifier (left). Bite plates (right side of bite force gauge) were wrapped in cloth medical tape to protect bat teeth. Bite force applied at bite plates was measured at the force transducer (left side of bite force gauge) connected to the charge amplifier which displayed bite force in newtons (N). The caliper on the bite force gauge served as the fulcrum and was used to adjust distance between bite plates.

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between canines at minimum gape allowed by the bite force meter due to the small size

of subject skulls. The meter was zeroed prior to each reading.

Gape Analysis

Maximum gape was determined using two-dimensional kinematic analysis of

videos taken of bats displaying active biting behavior. Gape analysis was performed by

isolating and capturing individual video frames then using image analysis software to

obtain measurements. Maximum gape angle and distance were determined for each

individual. The average of the maximums were calculated for each species and compared

for statistical significance.

Videos were analyzed using QuickTime Player (v 7.7.3; Apple Inc.; Cupertino,

CA, USA). The display format was configured to have frame number in the lower left

corner and included a Movie Inspector window to provide file and frame information.

Video files were perused frame-by-frame and frames depicting maximal gape or detailed landmarks were selected and captured to include the millimeter scale, bat, and Movie

Inspector information using the Window Snip option in Windows Snipping Tool (v

6.0.6001; Microsoft Corp.; Redmond, WA, USA), and saved in Joint Photographic

Experts Group file format (.JPG). Files were named by appending the frame number to the video filename and recorded in a spreadsheet (Microsoft Excel v 2007; Microsoft

Corp.; Redmond, WA, USA) along with elapsed time; head orientation: lateral, frontal,

oblique, or posterolateral; and gape status: partial or open.

Measurements were obtained from manually digitized landmarks using Olympus

DP2-BSW/cellSens Dimension image analysis software (v 2.2; Olympus Americas

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Corp.; Center Valley, PA, USA). Images were calibrated using the millimeter scale in the frame which was digitized to provide the pixel to millimeter conversion factor. The pixel and millimeter values were recorded and the conversion factor was calculated as pixel/mm.

Surface landmarks used for digitization were anterior alveolar margins of the

U upper and lower canine teeth (C and CL, respectively), and center of the eye (E) (Figure

U 7). Gape was recorded for each frame as the distance between C and CL. Objects drawn to facilitate measurements were expressed in pixels with resolution configured to two decimals. Circles were drawn with the 2-Point Circle tool, lines were drawn with the

Arbitrary Line tool, and line weight was configured to one point with short perpendicular helper end-lines to aide in object placement. Images were zoomed to 400% for accuracy without causing pixilation and measurement objects were drawn with different colors for clarity. The center of the eye (E) was located using a standardized small circle (25.13 pixel2) around the perimeter of the eye. The CU landmark was located and a line drawn

U U U connecting it to E (C E); CL was located and connected to C (C CL), and CL was connected to E (CLE) (see Figure 7). Adjacent frames of the movie file were continually referenced during landmark placement utilizing relative position of other facial features as aides. When possible on frontal and oblique orientations, contralateral landmarks were digitized and values were compared and reanalyzed as necessary until positioning was within the tolerance of one pixel. Pixel values were recorded and converted to mm using pixel/mm conversion factor. The file was saved as a Tagged Image File (.TIF) to retain the measurement information. All files were processed multiple times with results being

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Figure 7. Video Gape Measurement Landmarks. Distances were measured between the eye (E), anterior margin of upper canine tooth (CU), and anterior margin of the lower canine tooth (CL). The metric scale to the left of the frame was used to calibrate the measurements. Left lateral view of Mormoops blainvillei.

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updated until consecutive measurements were within one pixel and the existing values

were retained. The upper canine to eye distance (CUE) for an individual was determined

as the average of the individual CUE measurements from lateral and oblique orientations

only. Other orientations were excluded because their distances were understated as a

result of geometric perspective when not viewed in the horizontal viewing plane.

Normalization to allow interspecific comparison was achieved by converting gape

to gape angle. For purposes of this study, gape angle (G) is defined as the angle at the

craniomandibular joint (J) subtended by the maxillary segment (CUJ) and mandibular

segment (CLJ) (Figure 8). Because the canines are separated by the gape distance

U (C CL), a triangle is formed with the jaw segments and the law of cosines can be used to

determine any of the angles when the lengths of the sides are known. Maxillary and mandibular jaw lengths were calculated by multiplying CUE by the corresponding scaling

factor, based on intraspecific proportionality of the jaw lengths to the eye position

relative to the upper canine, as revealed in the next section (Jaw Scaling) for the two

species involved in this study. The law of cosines states that for given a triangle with

sides A, B, and C:

= + – 2 ( ) 2 2 2 or 𝐴𝐴 𝐵𝐵 𝐶𝐶 𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵 𝑎𝑎

+ ( ) = 2 2 2 2 𝐵𝐵 𝐶𝐶 − 𝐴𝐴 𝑐𝑐𝑐𝑐𝑐𝑐 𝑎𝑎 where a is the angle opposite side A. This calculation𝐵𝐵𝐵𝐵 provides the cosine of the angle

which is converted to the value of the angle by taking the inverse cosine. If G is the

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Figure 8. Video Gape Jaw Relationships. Gape angle (G) was calculated using the law of cosines with the lengths of the segments between the craniomandibular joint (J), anterior margin of upper canine tooth (CU), and the anterior margin of the lower canine tooth (CL) (see text). Left lateral view of Pteronotus quadridens skull. Pin glued to skull along the basicranial axis. The active gape will be smaller than the angle depicted here which is the maximum passive gape achieved without dislocation of the craniomandibular joint for this skull. The arc represents the path CL traces during jaw movement.

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U angle of interest and C CL is the side opposite, the resulting equation is:

( ) + ( ) ( ) ( ) = . 𝑈𝑈2 2× ( ) 2× ( 𝑈𝑈) 2 𝐶𝐶 𝐽𝐽 𝐶𝐶𝐿𝐿𝐽𝐽 − 𝐶𝐶 𝐶𝐶𝐿𝐿 𝑐𝑐𝑐𝑐𝑐𝑐 𝐺𝐺 𝑈𝑈 The Excel calculation for inverse cosine is ACOS𝐶𝐶 𝐽𝐽 which𝐶𝐶 computes𝐿𝐿𝐽𝐽 the result in radians so the DEGREES function is required to obtain the gape angle in degrees:

= ( ((( ^2) + ( ^2) ( ^2))/(2 ))). 𝑈𝑈 𝑈𝑈 𝑈𝑈 𝐿𝐿 𝐿𝐿 𝐿𝐿 U 𝐺𝐺 Maximum𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷𝐷 gape𝐴𝐴 was𝐴𝐴𝐴𝐴𝐴𝐴 determined𝐶𝐶 𝐽𝐽 for 𝐶𝐶each𝐽𝐽 individual− 𝐶𝐶 𝐶𝐶 as the greatest∗ 𝐶𝐶 𝐽𝐽of∗ the𝐶𝐶 𝐽𝐽 C CL measurements for that bat. The species average maximum gape was calculated by averaging the individual maximum gapes. Maximum gape angle for an individual was determined as the greatest gape angle for that bat. The species average maximum gape angle was calculated by averaging the individual maximum gape angles.

Jaw Scaling

Craniomandibular joint position was located using intraspecific jaw scaling factors derived from visible landmarks on the face. The distance between the eye and upper canine is directly proportional to the distances from the craniomandibular joint to the upper canine and the craniomandibular joint to the lower canine. These proportionality constants were used to calculate upper and lower jaw segment lengths in order to convert intercanine distance to an angle and are species specific for these two species.

Jaw scaling factors were derived using formalin preserved specimens from the

California State University, Sacramento Natural History Museum collection. Ten adults

(five males and five females) of each species, Mormoops blainvillei and Pteronotus

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quadridens, previously collected from Culebrones Cave in Puerto Rico were used (see

Appendix B for specimen identification numbers). Specimens were photographed and

dissected to expose the craniomandibular joint taking care not to disturb the position of

the eye. Manual measurements were also taken. Surface landmarks used were the same

as for gape analysis on live specimens: anterior alveolar margins of the upper canine (CU)

and lower canine (CL), and the center of the eye (E), with the addition of landmarks exposed by dissection of the annulus of the auditory bulla (O) and the craniomandibular joint (J) (Figure 9). All photographs and measurements were based on a right lateral orientation.

Dissections were performed in the lab at California State University, Sacramento.

All stages of dissection were photographed using an Olympus DP25 five megapixel digital color camera mounted via a 0.63x camera adapter (U-TV0.63XC) onto an

Olympus SZX10 binocular dissecting microscope equipped with a DFPL-4 0.5x, 0.06 aperture objective (Olympus Americas Corp.; Center Valley, PA, USA). A 24 inch tall boom stand with weighted base, 20.5 inch arm, and non-tilting boom stand adapter (SMS

16B; Diagnostic Instruments, Inc.; Sterling Heights, MI, USA) was used to provide necessary room for dissection. The microscope camera was attached to a laptop computer via a powered IEEE 1394 high-speed serial bus mini firewire cable (1394 6T4;

Olympus Americas Corp.; Center Valley, PA, USA) and controlled with Olympus DP2-

BSW/cellSens Dimension image analysis software (v 2.2; Olympus Americas Corp.;

Center Valley, PA, USA). Lighting was supplied by dual fiber optic illuminators

(TechniQuip FOI-150 150w; TechniQuip; Pleasanton, CA, USA). Software was

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Figure 9. Jaw Scaling Measurement Landmarks. Centers of the auditory bulla (O), and eye (E) were located using circles of specific sizes (see text). Additional landmarks of U anterior margin of upper canine tooth (C ), anterior margin of the lower canine tooth (CL) and craniomandibular joint (J) were located and connected as necessary for measurement purposes. Right lateral view of Mormoops blainvillei head with skin and musculus masseter removed.

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configured to metric and calibrated using a dial caliper (Mitutoyo 505-681 0-150mm

±0.03mm; Mitutoyo America; Aurora, IL, USA) for each objective and zoom setting. A small scale bar was configured to be displayed in the lower right corner of the display and images. Calibration was verified prior to processing each specimen using the dial caliper on the scale bar. Primary dissection instruments consisted of number 5 micro forceps (A.

Dumont & Fils; Autil, Switzerland) and a number 3 scalpel handle with number 11 scalpel blades (Feather Safety Razor Co., Ltd.; Osaka, Japan).

In dissections, the skin was removed from the right side of the head and around the mouth to expose the alveolar margins, mandible, rostrum, ear cartilages, and musculature over the cranium while leaving the eye in place. The external ear structures were removed exposing the annulus of the auditory bulla. Then the musculus masseter was removed and the craniomandibular joint exposed and opened laterally with care being taken not to disturb the position of the eye. A dissecting pin was inserted between the upper and lower incisors to provide approximately one millimeter of separation between the canine alveolar margins, and an insect pin was inserted into the anterior aspect of the craniomandibular joint to create a small gap between the mandibular condyle and the glenoid fossa to aide in visually locating the joint (see Figure 9). Several images were taken at each step of dissection.

Manual measurements between the exposed landmarks were taken using dial calipers (Mitutoyo 505-681 0-150mm ±0.03mm; Mitutoyo America; Aurora, IL, USA).

Digital measurements were obtained from the image files using the camera software

Olympus DP2-BSW/cellSens Dimension image analysis software (v 2.2; Olympus

22

Americas Corp.; Center Valley, PA, USA).

Software objects drawn to facilitate measurement were expressed in millimeters with resolution configured to two decimals. Circles were drawn with the 2-Point Circle tool, lines were drawn with the Arbitrary Line tool, and line weight was configured to one point with short perpendicular helper end-lines. Images were zoomed for accuracy

without causing pixelation and measurement objects were drawn with different colors for

clarity. A small circle, with an area of 0.70mm2 for M. blainvillei and 0.51mm2 for P.

quadridens, was drawn over the eye to locate its center (E), and a larger red circle, with

an area 1.50mm2 for M. blainvillei and 2.02mm2 for P. quadridens, was drawn over the

tympanic annulus of the ear to locate its center (O). The CU landmark was located and a

line with long perpendicular helper end-lines was drawn through E to the point either tangent to O where the helper end-line intersected O for P. quadridens or directly through

O for M. blainvillei (Figure 10). This CUO segment was subdivided with a line

U connecting C to E, and a line connecting E to O. CL and J were located and connected

by a line using the tangent point on the mandibular condyle for J, and a line was drawn to connect J to CU using the tangent point on the glenoid fossa for J (see Figure 10). The

TIF file was updated with the measurement information. All files were processed multiple times until consecutive measurements yielded consistent results.

Bite Force Analysis

Bite force measurements were converted to actual values by applying a conversion factor based on the configuration of the meter. The bite force meter operates as a first class lever system with the caliper serving as the fulcrum. The effort (E) is the

23

Figure 10. Landmark Relationships for Jaw Scaling. Circles of specific sizes were drawn over the eye (E) and auditory bulla (O). A line was drawn from the anterior margin of upper canine tooth (CU) through the center of E to the center of O for Mormoops blainvillei (left) or to the point on the circle tangent to O for Pteronotus quadridens (right). Right lateral views of head with skin removed. Pins glued to skulls along basicranial axis to achieve a standardized orientation.

24 compressive bite force of the bat at the bite point of the bite plates, the fulcrum (F) is the caliper, and the resistance (R) is the tensile force measured at the force transducer (Figure

11). In a first class lever,

( ) × ( ) = ( ) × ( ) or 𝐸𝐸 𝑁𝑁 𝐸𝐸𝐸𝐸 𝑚𝑚𝑚𝑚 𝑅𝑅 𝑁𝑁 𝑅𝑅𝑅𝑅 𝑚𝑚𝑚𝑚

= × 𝑅𝑅𝑅𝑅 𝐸𝐸 𝑅𝑅 where EF is the effort moment arm and RF is the𝐸𝐸 𝐸𝐸resistance moment arm (Hamilton et al.

2008). The mechanical advantage is the ratio of the effort moment arm to the resistance moment arm (Hall 2014). Because EF is longer than RF, the measured force was overstated and the mechanical advantage was a fraction, so the measured force was scaled by the ratio of RF to EF which was 0.6194 for this bite force meter.

Maximum bite force for each individual was determined as the maximum of all the bite force readings for that bat. Bats producing maximum measured bite forces of less than 2.0 N (1.24 N actual) were considered non-cooperative and therefore not included in the sample. The species average maximum bite force was calculated by averaging the individual maximum bite force values for the bats of that species. Skull proportion was used when available to normalize the maximum individual bite forces to newtons per cubic centimeter (N/cm3), where skull proportion = head length * head width

* head height, all in millimeters, and then dividing by 103 to convert to cm3. The species average maximum normalized bite force was calculated by averaging the individual maximum normalized bite forces for the bats of that species.

25

Figure 11. Bite Force Meter Lever Properties. Bite force meter functions as a first class lever system with the caliper serving as the fulcrum (F) along vertical fulcrum axis (AF). Compressive bite force is applied at bite point of bite plate (E) along vertical effort axis (AE) and measured as tensile force by the force transducer (R) along the vertical resistance axis (AR). The effort moment arm (EF) is longer than the resistance moment arm (RF) so the force measured at R is higher than the force exerted at E and has to be scaled down by RF/EF which is 0.6194 for this meter.

26

Jaw Musculature

Musculature analysis of the jaw was performed using formalin preserved specimens from the California State University, Sacramento Natural History Museum collection. One male and one female of each species, Mormoops blainvillei and

Pteronotus quadridens, previously collected from Culebrones Cave in Puerto Rico were dissected to remove main craniomandibular muscles from both sides of the skull. Jaw adductor muscles dissected consisted of musculus temporalis for wide gape, and m. masseter and m. pterygoideus medialis for narrow gape (Turnbull 1970; Herring and

Herring 1974). Jaw abductor muscles consisted of m. digastricus and m. pterygoideus lateralis (Kawamura et al. 1968; Turnbull 1970; Kallen and Gans 1972; McNamara

1973). M. pterygoideus medalis is also known as m. pterygoideus internus and m. pterygoideus lateralis is also known as m. pterygoideus externus (Turnbull 1970). For the purpose of this study, m. temporalis includes m. temporalis pars superficialis, m. temporalis pars medius, m. temporalis pars profundus, and m. temporalis pars suprazygomatica while m. masseter includes m. masseter pars superficialis, and m. masseter pars profundus, plus m. zygomaticomandibularis pars superficialis, and m. zygomaticomandibularis pars profundus (Herrel et al. 2008). Individual muscle pairs were processed to determine mass and fiber length which were used to calculate physiological cross sectional area (PSCA), a better predictor of muscle power than muscle mass (Lieber and Fridén 2000). PCSA proportions of muscle groups to total jaw musculature were calculated by individual and averaged for species.

Skulls were dissected down to the temporalis fascia which was then carefully

27

teased away and removed from the m. temporalis. Skin and connective tissue of the

rostrum, along with the superficial musculature of the neck region were dissected and

removed. Muscles of interest were carefully cleaned and teased away from their

attachments using microforceps and scalpel, and stored separately in 70% ethanol. The mm. digastricus were removed first from each of the specimens, followed by the mm. masseter, and mm. temporalis (Figure 12). The tongue and suprahyoid musculature were

then removed to expose the mm. pterygoideus groups. Lastly, the mm. pterygoideus

medialis and mm. pterygoideus lateralis were removed and processed (Figure 13).

Muscles were blotted dry, and mass was measured to the nearest 0.1mg using an analytical balance (Mettler-Toledo AB104-S 110gx0.0001g; Mettler-Toledo, LLC;

Columbus OH, USA). A small dissecting pin was included in each muscle measurement to make sure the analytical balance was above its minimum tolerance due to the small muscle masses being measured. Muscles were weighed five times each, both individually and in pairs, including the dissecting pin. The dissecting pin was weighed separately for each set of measurements and its mass subtracted off the recorded values for that set of muscles. Averages were calculated for individual and paired muscle masses for each specimen. Species average mass was calculated from the average bilateral mass values for each specimen.

In order to obtain fiber lengths, muscles were dissolved in 30% nitric acid for 24 hours and transferred to 50% glycerin (Herrel et al. 2008; Santana et al. 2010). Muscles were stained using Ponceau 2R acid fuscin, a variation of Masson’s Trichrome stain

(Flint et al. 1975), to differentiate connective tissue from muscle fiber. Fibers were

28

Figure 12. Superficial Masticatory Musculature. Right lateral view of Pteronotus quadridens head dissected to show musculus temporalis (T), m. masseter (M), and m. digastric (D). Pin glued to skull along the basicranial axis.

29

Figure 13. Deep Masticatory Musculature. Inferior view of Mormoops blainvillei head dissected to show musculus masseter (M) and m. pterygoideus medialis (PM) on the left side. M. pterygoideus medialis was removed on the right side to expose m. pterygoideus lateralis (PL).

30

teased apart using fine needles under a dissecting microscope. Ten intact fibers were

randomly selected and photographed for each muscle, both left and right sides.

Measurements were taken using Olympus DP2-BSW/cellSens Dimension image analysis

software (v 2.2; Olympus Americas Corp.; Center Valley, PA, USA). The greatest fiber

length measurement for each muscle, both left and right, were selected from the

individual measurements and then averaged to calculate the fiber lengths for that

specimen. Species muscle fiber lengths were calculated by averaging the specimen fiber

lengths for the bats of that species. Maximums were used to eliminate incomplete fibers

because individual fibers are very delicate and easily damaged during the preparation

process.

Physiological cross-sectional area (PCSA) was calculated as:

( ) × cos ( ) = ( ) × ( ) 2 𝑀𝑀 𝑔𝑔 𝜃𝜃 3 𝑃𝑃𝑃𝑃𝑃𝑃𝑃𝑃 𝑚𝑚𝑚𝑚 𝑓𝑓 where is the mass of the muscle in grams,𝜌𝜌 𝑔𝑔 and⁄𝑚𝑚𝑚𝑚 is the𝐿𝐿 pennation𝑚𝑚𝑚𝑚 angle of the fibers,

is the muscle𝑀𝑀 density in g/mm3, and is the fiber𝜃𝜃 length in mm (Leiber 2010). None of𝜌𝜌

𝑓𝑓 the muscles dissected had any noticeable𝐿𝐿 pennation so 0° was used for all muscles. The muscle density used was that of mammalian muscle, 1.056 g cm (Ward and Leiber 3 2005). ⁄

Statistical Analyses

Two-tailed independent t-tests with alpha<0.05 were used to compare sample

means and determine significance for hypothesis testing. The probability (p) value was

calculated using the Excel distribution function (TDIST) on the test statistic, t, where m is

31

the group mean, v is the group variance, and n is group size and degrees of freedom (df)

is n-2:

= 𝑚𝑚1 −+𝑚𝑚2 𝑡𝑡 𝑣𝑣1 𝑣𝑣2 � 1 2 = 𝑛𝑛( , 𝑛𝑛 , 2).

The first hypothesis that there𝑝𝑝 are𝑇𝑇𝑇𝑇 differences𝑇𝑇𝑇𝑇𝑇𝑇 𝑡𝑡 𝑑𝑑𝑑𝑑 in normalized gapes between

Mormoops blainvillei and Pteronotus quadridens was considered significant if the p

value was less than 0.05. Statistical analysis for jaw scaling used Pearson coefficient of correlation (r) with alpha<0.05 to determine significance. For the sample size of 10 with paired values, the degrees of freedom (df) is n – 2 = 8. The PEARSON function in Excel was used to calculate the r value, and the probability (p) value was calculated using the distribution function (TDIST) on the test statistic, t:

= ( (( )/(1 )) 2 𝑡𝑡 𝑟𝑟 ∗= 𝑠𝑠𝑠𝑠𝑠𝑠𝑠𝑠 𝑑𝑑𝑑𝑑( , , 2−). 𝑟𝑟

U U U The correlation between C E and C𝑝𝑝 J, 𝑇𝑇and𝑇𝑇𝑇𝑇 𝑇𝑇between𝑇𝑇 𝑡𝑡 𝑑𝑑𝑑𝑑 C E and CLJ were considered

significant for each species if the p value was less than 0.05.

The second hypothesis that there are differences in normalized bite forces

between Mormoops blainvillei and Pteronotus quadridens was considered significant if

the p value was less than 0.05 using a two-tailed independent t-test.

Statistical analysis of the third hypothesis that Mormoops blainvillei and

Pteronotus quadridens have different distributions of their jaw abductor and adductor

musculature is not being offered due to the nominal sample size. Dissection of

32 musculature required destructive sampling resulting in a small sample size of n = 4.

Individual variability is not taken into account in other bat jaw muscle studies which use a small sample size of one or two specimens per species (Herrel et al. 2008; Santana et al.

2010).

33

RESULTS

A total of 154 individual bats were processed in the field of which 112 provided usable data. An additional 24 museum specimens were analyzed for a total of 136 bats contributing data to this study (Table 1). Seventy one video recordings were made of 47 bats of which 43 individuals yielded 64 usable recordings. From these videos, 556 frames were captured and analyzed for an average of 13 frames per bat (range 5 to 34)

(Table 2). A total of 189 bite force measurements were obtained from 65 bats of which only 40 individuals provided readings that were above the minimum threshold of 2.0 N, reducing the total to 138 bite force measurements with an average of 3.5 per individual

(Table 2).

Gape Analysis

The average maximum active gape of 14.2mm for M. blainvillei was greater than

P. quadridens at 10.2mm (Table 3 and Figure 14) which is a statistically significant difference between the two species (two tailed independent t-test, t = 21.53, df = 41, p <

0.001). The normalized average maximum active gape angle of 78.2° for M. blainvillei was greater than P. quadridens at 65.1° (Table 3 and Figure 14) which is a statistically significant difference between the two species (two tailed independent t-test, t = 4.39, df

= 41, p < 0.001). The maximum active gape of 19.2mm, 100.3° for M. blainvillei was greater than P. quadridens at 13.7mm, 83.5° (Table 3). See Appendix A for detail summarized in Table 3 and Figure 14. For jaw scaling, the maxillary (CUJ) and

U mandibular (CLJ) jaw lengths were both strongly correlated to eye position (C E) in M.

blainvillei [Pearson coefficient of correlation: r = 0.99, df = 38, p < 0.001 and r = 0.99,

34

Table 1. Total Number of Mormoopid Bats Analyzed by Species. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens. Species Mobl Ptqu Total Bats video recorded 25 22 47 Bats measured for bite force 35 30 65 Jaw scaling dissections 10 10 20 Muscle analysis dissections 2 2 4 Total bats used 72 64 136

35

Table 2. Total Video Frames and Bite Force Measurements Analyzed for Two Species of Mormoopid Bats. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens. Species Mobl Ptqu Total Bats video recorded with usable frames 22 21 43 Total video recordings with usable frames 31 33 64 Number of frames captured 278 278 556 Average frames captured per bat 12.6 13.2 12.9 Total bats used for bite force measurement 20 20 40 Number of bite forces obtained 67 71 138 Average bite forces per bat 3.4 3.6 3.5

36

Table 3. Maximum Active Jaw Gapes for Two Species of Mormoopid Bats. Sample size is n, average values include standard error. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens. Avg Max Avg Max Gape Max Gape Max Gape

Species n Gape (mm) (degrees) (mm) (degrees) Mobl 22 14.16 ± 0.39 78.17 ± 2.48 19.23 100.26 Ptqu 21 10.21 ± 0.21 65.08 ± 1.71 13.65 83.52

37

Figure 14. Average Maximum Active Jaw Gape Distance and Angle for Two Species of Mormoopid Bats. Maximum gape distance (left) and maximum gape angle (right) with standard error bars, and sample size in parentheses. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens.

38

df = 38, p < 0.001, respectively] and P. quadridens [r = 0.99, df = 38, p < 0.001 and r =

0.99, df = 38, p < 0.001, respectively] (see Figure 10). See Appendix B for jaw scaling detail data.

Bite Force Analysis

Average maximum bite force of 1.74N for M. blainvillei was greater than P.

quadridens at 1.56N (Table 4 and Figure 15) which is a statistically significant difference

between the two species (two tailed independent t-test, t = 2.21, df = 38, p < 0.04).

However, the normalized average maximum bite force was greater for P. quadridens of

78.17N/cm3 than M. blainvillei at 65.08N/cm3 (Table 4 and Figure 15) which is a

statistically significant difference between the two species (two tailed independent t-test, t

= 2.74, df = 27, p < 0.02). Maximum bite force was greater for M. blainvillei (2.25N)

than P. quadridens (2.07N) while normalized maximum bite force was greater for P.

quadridens at 2.28N/cm3 than M. blainvillei at 1.79N/cm3 (Table 4). See Appendix C for

bite force detail data.

Jaw Musculature

Muscle masses, fiber lengths, and physiological cross sectional area (PCSA) of all

masticatory muscles were greater for Mormoops blainvillei than for Pteronotus

quadridens (Table 5). M. blainvillei had a greater overall PCSA of masticatory

musculature than P. quadridens (34.33mm2 versus 29.36mm2) and also a greater

percentage of total adductor (91.8%) to abductor musculature (8.2%) than P. quadridens

(91.0%, and 9.0%, respectively) (Table 6). M. blainvillei had a larger proportion of wide

gape adductor muscles (70.8%) than P. quadridens (68.9%), whereas P. quadridens had a

39

Table 4. Maximum Bite Forces for Two Species of Mormoopid Bats. Average values include standard error, and sample size in parentheses. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens. Avg Max Max Average Maximum Normalized Normalized Species Maximum Bite Bite Force Bite Force Bite Force Force (N) (N) (N/cm3) (N/cm3) Mobl 1.74 ± 0.06 (20) 0.65 ± 0.05 (20) 2.25 1.11 Ptqu 1.56 ± 0.07 (20) 0.95 ± 0.09 (9) 2.07 1.41

40

Figure 15. Average Maximum Actual and Normalized Bite Forces for Two Species of Mormoopid Bats. Maximum actual measured bite force (left) and maximum normalized bite force (right) with standard error bars, and sample size in parentheses. Skull proportion (skull height × skull width ×skull length) is used for normalization. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens.

41

Table 5. Craniomandibular Muscle Architecture Measurements of Mormoopid Bats by Sex and Species. Measurements of muscle mass (mass) in mg and muscle fiber length (fiber len) in mm used to calculate physiological cross sectional area (PCSA) for paired muscles (see methods for jaw musculature). Wide gape adductor muscles (wide gape close): musculus temporalis (temporalis); narrow gape adductor muscles (narrow gape close): m. masseter (masseter) and m. pterygoideus medialis (m. pterygoid); abductor muscles (open): m. digastricus (digastric) and m. pterygoideus lateralis (l. pterygoid). Sex is m for male and f for female n is number of specimens analyzed. Species Mormoops blainvillei Pteronotus quadridens Sex m f avg m f avg Specimen 180601-1 230510-42 (n=2) 260510-22 230608-10 (n=2) Muscle temporalis mass 93.2 79.2 86.2 67.4 74.6 71.0 fiber len 3.20 3.58 3.39 3.69 3.05 3.37 PCSA 27.64 20.97 24.30 17.30 23.16 20.23 masseter mass 16.0 13.9 15.0 13.0 13.2 13.1 fiber len 2.86 3.12 2.99 3.16 2.21 2.68 PCSA 5.32 4.24 4.78 3.89 5.68 4.78 m. pterygoid mass 4.7 5.0 4.8 3.3 2.2 2.8 fiber len 1.83 1.89 1.86 1.67 1.29 1.48 PCSA 2.43 2.49 2.46 1.89 1.63 1.76 digastric mass 11.0 9.0 10.0 8.0 9.3 8.6 fiber len 6.81 7.21 7.01 5.50 6.62 6.06 PCSA 1.53 1.18 1.35 1.37 1.33 1.35 l. pterygoid mass 2.4 3.5 3.0 2.2 1.9 2.1 fiber len 1.62 2.26 1.94 1.76 1.41 1.58 PCSA 1.40 1.48 1.44 1.18 1.29 1.24

42

Table 6. Craniomandibular Muscle Organization for Two Species of Mormoopid Bats. Values are for physiological cross sectional area (PCSA) in mm2. Wide gape adductor muscles consists of musculus temporalis; narrow gape adductor muscles consist of m. masseter and m. pterygoideus medialis; and abductor muscles consist of m. digastricus and m. pterygoideus lateralis. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens. Muscle Group PCSA (mm2) Species Mobl n=2 Ptqu n=2

Total (Adductors + Abductors) 34.33mm2 100.0% 29.36mm2 100.0% Abductors 2.79mm2 8.1% 2.59mm2 8.8% Adductors 31.54mm2 91.9% 26.77mm2 91.2% Wide Gape Adductors 24.30mm2 77.0% 20.23mm2 75.6% Narrow Gape Adductors 7.24mm2 23.0% 6.54mm2 24.4%

43 larger proportion of narrow gape adductor muscles (22.3%) than M. blainvillei (21.1%)

(Figure 16). See Appendix D for muscle mass data and Appendix E for fiber length data summarized in Tables 5, 6 and Figure 16.

44

Figure 16. Masticatory Muscle Distribution in Two Species of Mormoopid Bats. Percentages based on total physiological cross sectional area (PCSA) for paired muscles. Wide gape adductor muscles (wide closers) consist of musculus temporalis; narrow gape adductor muscles (narrow closers) consist of m. masseter and m. pterygoideus medialis; and abductor muscles (openers) consist of m. digastricus and m. pterygoideus lateralis. Mobl represents Mormoops blainvillei and Ptqu represents Pteronotus quadridens.

45

DISCUSSION

Coexisting species within a shared habitat that have similar feeding ecology

commonly exhibit resource partitioning, such as prey specialization, as a mechanism to

reduce competition. Morphological differences in feeding apparatus have been shown to

correspond to differences in prey selection in bats (Santana and Cheung 2016).

Masticatory structural and functional differences were examined for two coexisting species of insectivorous bats in relation to their dietary preferences.

Comparison of the two species of mormoopid bats in this study showed that the larger Mormoops blainvillei displayed greater active gape, overall masticatory muscle mass, and bite force than the smaller Pteronotus quadridens (see Tables 3, 4 and 5). A larger bat should have a larger skull because body size and skull size generally show positive allometry in bats (Nogueira et al. 2009). Larger skulls exhibit wider gape for the same gape angle, and larger jaw musculature supporting greater bite force (Herrel et al.

2008), both of which were true for M. blainvillei. Skull lengths between these two species were approximately the same suggesting their maximum gape and bite forces should have been comparable (Herrel et al. 2008); however, this comparison is complicated by the differences in skull shape. In the skulls of most species of bats, the jaw is in line with the condylobasal axis (Czarnecki and Kalen 1980), the standard metric for skull length, which scales directly with jaw length. The varying degrees of upturning of the rostrum in different mormoopid bat species changes this relationship and the jaw length is no longer a direct component of the skull length (Czarnecki and Kalen 1980).

M. blainvillei with a radically upturned rostrum has a longer functional jaw length, as

46

evidenced by a longer maxillary tooth-row averaging 7.5mm (Lancaster and Kalko

1996), versus P. quadridens averaging 5.9mm (Rodríguez-Durán and Kunz 1992), yet the

their skull lengths are essentially the same. Other researchers (Dumont and Herrel 2003;

Santana 2016) normalize gape by skull length or body mass but that would not work for the species studied herein based on the skull length issues previously stated, and many of

the specimens collected were pregnant females so body mass would have been

overstated. A better method than skull length might be jaw length measured from

mandibular angle to front of rostrum, but this measurement is not easily taken in the field

on live specimens. Forearm length is commonly used as a proxy for body size but this

has not been shown to have any direct relevance to craniomandibular size. Normalization

of gape is best achieved by gape angle, which is not dependent on individual variability

but not easily measured on live specimens. The modelling method developed for this

study worked well and is discussed below.

Maximum gape for M. blainvillei was 32% greater than P. quadridens while the

normalized gape (angle) in the former species was 18% greater than the latter. This

supports the first hypothesis that M. blainvillei has a greater maximum normalized active

gape than P. quadridens. Larger gape would improve the ability to take larger prey

(Freeman 1979) corresponding to M. blainvillei prey specialization of larger Lepidoptera

(Rolfe and Kurta 2013). The smaller gape of the insect generalist P. quadridens would allow it to feed on smaller Lepidoptera and Coleoptera, which are generally smaller

(Aguirre et al. 2003; Ober and Hayes 2008), but not larger Lepidoptera (Rolfe and Kurta

2013).

47

It is unlikely that active gapes recorded in this study are truly maximal. There is

no way to determine if induced gape is truly maximal, but maximal passive gape is a plausible limitation. The point of disarticulation of the craniomandibular joint would represent the maximum passive gape, which was shown in a preliminary study for these two species to have been greater than 15% larger for M. blainvillei and 18% larger for P. quadridens than the active gape values presented here (Changaris and Lancaster 2011).

Induced gapes were most likely the result of defensive biting behavior and may not represent feeding behavior or maximum gape. Weaker bite forces accompany wider gapes (Santana 2016) so unless wide gape was only being used as a defensive display and not for functional defense it is unlikely that the bat would open to a point that the ability to bite effectively would be compromised. Other limitations to active gape include restrictive soft tissue structures such as ligaments, joint capsule, muscle length/stretch, and skin/fur limitations, in addition to voluntary behavioral control. The act of yawning would likely elicit maximum active jaw opening (Terhune et al. 2015) which would be a challenge in itself to induce in bats, but a better predictor of maximum active gape.

The ability to locate the jaw joint on a live specimen based on eye and canine tooth position using kinematics is a powerful modelling tool. It would allow recording and analysis of biting behavior in a natural setting to determine functional maximum gape eliminating the inherent trauma and behavioral modifications that accompany catching and handling live specimens. For these two species, the maxillary and mandibular canine to craniomandibular offsets were found to be a function of maxillary canine to eye distance based on highly statistically accurate jaw scaling factors

48

(p<0.0001). These values, along with gape distance between canines allowed gape normalization by converting it to an angle. This, however, is not the same gape angle subtended by the maxillary and mandibular tooth rows, whose apex varies with gape due to the mandibular angle and ramus (Storch 1968). Future studies should examine this relationship to see if it extends to other bat species and potentially other mammals.

Kinematic analysis was limited by the equipment used. The inability to use blurry maximal gape frames probably resulted in understating maximum gape values. Faster video of at least 60fps (frames per second), preferably 120 fps or higher, would have increased the number of sharp, usable maximal gape frames than the 30fps used. High speed infrared video of bats in their native environment performing various behaviors including feeding, navigation, and socialization would be ideal, but current technology is limiting and such an implementation would be very costly if even possible at this time.

Even though bite forces showed statistical difference, the methodology used for obtaining bite forces did not appear to be reliable for smaller bats because they were generally non-cooperative when they were induced to bite, yielding apparently inconclusive results. While the bats readily showed biting behavior, they were reluctant to bite the bite force gauge which seemed to be too big for their small mouths. Other researchers did not experience this problem with phyllostomid bats, mainly frugivores,

(Aguirre et al. 2002; Dumont and Herrel 2003; Santana and Dumont 2009) but even using the same bite force meter in this study, I was unable to obtain consistently reliable results, or even replicate data for small phyllostomids. Freeman and Lemen (2010) also observed this issue with cooperativity in bats and other mammals. A less ominous gauge

49 that has bite plates that are closer together might provide more reliable results (Freeman and Leman 2008). M. blainvillei had a greater bite force than P. quadridens in this study, but anecdotal experience handing these two species suggested that P. quadridens was able to bite much harder than M. blainvillei, another reason to doubt the veracity of the bite force data collected. However, normalizing bite forces showed that P. quadridens had a 37.5% greater bite force than M. blainvillei which supports the second hypothesis that P. quadridens has a greater normalized bite force than M. blainvillei. Greater bite force is necessary for P. quadridens to process hard Coleoptera prey items (Freeman and

Leman 2007) that make up a large part of its diet (Rolfe and Kurta 2013) while the reduced bite force of M. blainvillei is adequate to process the softer prey items (Freeman and Leman 2007) of its specialized Lepidoptera diet preference (Rolfe and Kurta 2013).

Jaw adductor (closer) muscle distribution slightly favored the wide gape muscles

(musculus temporalis) in M. blainvillei and the narrow gape muscles (m. masseter and m. pterygoideus medialis) in P. quadridens, but the jaw abductor (opener) musculature (m. digastricus and m. pterygoideus lateralis) did not show any difference between the two species (see Table 6). The first part of the third hypothesis that M. blainvillei would have greater wide-gape adducting muscle group proportions than P. quadridens was supported but the second part that M. blainvillei would have greater jaw abducting muscle group proportions than P. quadridens was not. The assumption that there is no individual variation within species along with the small sample size limits the ability to draw conclusions from this data. Small sample sizes (n=1 or 2) are typical for bat jaw muscle studies (Herrel et al. 2008; Santana et al. 2010) mainly because it requires destructive

50

sampling, and is very time consuming due to tedious detailed dissection and processing.

Further analysis of the jaw musculature is warranted due to the small sample size.

Physiological cross sectional area represents force capability of the muscles so the

similarity of abductors makes sense because force required to open the jaw is minimal.

Longer abductor muscle fiber lengths found in M. blainvillei indicates higher velocity of contraction (Leiber 2010) which means faster opening speeds that could be beneficial for capturing large prey while airborne. Shorter fiber lengths in the adductor muscles of P. quadridens suggests greater force production and slower speeds of contraction (Leiber

2010) which supports their durophagous diet that includes hard-bodied Coleoptera

(Freeman 1979; Aguirre et al. 2003; Herrel et al. 2008; Santana and Dumont 2009).

Mormoops reduced bite force is consistent with greater gape which in turn supports their feeding specialization of larger, soft-bodied prey. The radically upturned rostrum of Mormoops could explain the greater gape capabilities by allowing for a greater range of motion before encountering structural limitations (see Figure 2).

Morphological differences supporting a feeding specialization might be attributable to selection pressures, suggesting Mormoops skull shape may have been an evolutionarily divergent specialization.

51

CONCLUSIONS

Mormoops blainvillei displayed wider normalized active gape, reduced normalized bite force, and muscle distribution that slightly favoring wide-gape muscles while Pteronotus quadridens displayed muscle distribution that slightly favored narrow- gape muscles. Wide gape could facilitate specialization on larger insects but reduced bite-force would likely limit prey to soft-bodied insects which is consistent with the

Lepidoptera prey specialization of M. blainvillei (Rolfe and Kurta 2013). The narrower gape of P. quadridens would limit prey size while the greater bite force would allow durophagy which is necessary to process hard beetles as part of the diet of an insect generalist (Rolfe and Kurta 2013). Muscle distribution to support the wider gape in M. blainvillei may be responsible for reduction in bite force production which would not be an issue while processing softer prey.

Higher speed video for kinematics, bite force data obtained with a simpler piezo- electric force transducer, and additional specimens dissected for muscle analysis would all serve to expand and enhance the data collected in this study. Analysis of jaw scaling relationships in other families of bats and potentially other mammals would provide a future opportunity to see if the jaw scaling proportionality would provide a powerful modelling tool for non-intrusive location of the jaw joint.

52

APPENDIX A

Video Gape Frame Capture Measurement Data. Each line represents one frame with a summary line per bat. X-mm is length in millimeters measured from upper canine to eye (CUE). Y-mm is gape distance measured in millimeters from upper canine to lower canine U (C CL). Z-mm is length in millimeters measured from lower canine to eye (CLE) (see Figure 6). Average Canine to Eye (CUE) distance for each specimen is determined by averaging the X-mm values for that specimen excluding understated frontal views. Upper U U jaw (C J) and lower jaw (CLJ) lengths are calculated as C E × appropriate jaw scaling factor (see text). Gape angle (gape°) is calculated by the law of cosines using average CUE and jaw lengths (see text). Filename includes specimen id, species where Mobl is Mormoops blainvillei and Ptqu is Pteronotus quadridens, sex where M is male and F is female, and recording sequence number for that individual.

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 260510-2moblm 001.MOV 796 :26.53 frontal partial 3.6 8.5 9.9 41.68 260510-2moblm 001.MOV 797 :26.57 frontal open 5.6 10.2 11.3 50.38 260510-2moblm 001.MOV 798 :26.60 frontal open 3.9 8.2 10.4 40.38 260510-2moblm 001.MOV 883 :29.43 oblique partial 6.0 11.3 11.9 56.41 260510-2moblm 001.MOV 917 :30.57 lateral open 6.0 14.1 11.7 72.58 260510-2moblm 6.0 14.1 0.4879 0.5245 12.3 11.5 72.58 270510-2moblm 001.MOV 28 :00.93 oblique partial 6.0 5.1 8.9 24.75 270510-2moblm 001.MOV 501 :16.70 oblique partial 6.0 4.9 8.4 23.75 270510-2moblm 001.MOV 536 :17.87 oblique open 6.0 12.9 13.8 66.07 270510-2moblm 001.MOV 540 :18.00 lateral open 6.0 11.3 12.2 56.65 270510-2moblm 001.MOV 549 :18.30 oblique open 6.0 10.8 11.7 53.82 270510-2moblm 001.MOV 617 :20.57 frontal open 5.6 12.1 13.2 61.44 270510-2moblm 001.MOV 711 :23.70 oblique partial 6.0 12.0 12.8 60.80 270510-2moblm 6.0 12.9 0.4879 0.5245 12.3 11.4 66.07 270510-6mobm 001.MOV 270510-9moblm 001.MOV 371 :12.37 frontal open 5.4 11.0 14.5 57.55 270510-9moblm 001.MOV 379 :12.63 oblique partial 5.8 6.5 8.8 33.00 270510-9moblm 001.MOV 496 :16.53 oblique partial 5.9 4.7 8.0 23.60 270510-9moblm 001.MOV 513 :17.10 lateral open 5.8 10.7 10.3 55.49 270510-9moblm 001.MOV 514 :17.13 lateral open 5.9 9.7 9.7 49.94 270510-9moblm 001.MOV 837 :27.90 frontal open 5.7 10.8 10.8 56.52 270510-9moblm 001.MOV 838 :27.93 frontal open 5.3 11.3 9.8 59.47 270510-9moblm 002.MOV 92 :03.07 oblique partial 5.7 6.3 8.3 31.53 270510-9moblm 002.MOV 309 :10.30 oblique open 5.9 11.4 9.5 59.57 270510-9moblm 002.MOV 549 :18.30 lateral open 5.6 12.8 15.2 68.26 270510-9moblm 002.MOV 569 :18.97 frontal open 5.8 9.8 8.1 50.73 270510-9moblm 002.MOV 692 :23.07 oblique partial 5.5 7.7 8.8 39.00

53

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 270510-9moblm 002.MOV 755 :25.17 frontal open 4.9 12.1 13.6 63.90 270510-9moblm 002.MOV 841 :28.03 frontal open 5.5 10.7 13.4 56.00 270510-9moblm 002.MOV 856 :28.53 frontal open 5.5 13.5 13.7 72.21 270510-9moblm 002.MOV 859 :28.63 frontal open 5.0 13.1 12.9 69.93 270510-9moblm 002.MOV 1120 :37.33 lateral open 5.9 12.8 11.8 68.12 270510-9moblm 002.MOV 1131 :37.70 lateral open 5.8 15.1 13.7 82.78 270510-9moblm 002.MOV 1481 :49.37 oblique partial 4.5 5.1 8.0 25.64 270510-9moblm 5.8 15.1 0.4879 0.5245 11.8 11.0 82.78 270510-10moblm 001.MOV 694 :23.13 oblique partial 5.7 5.6 8.2 28.32 270510-10moblm 001.MOV 1041 :38.30 frontal open 5.8 15.6 13.3 86.53 270510-10moblm 001.MOV 1145 :38.17 lateral open 5.8 12.2 11.1 64.67 270510-10moblm 001.MOV 1149 :38.30 lateral open 5.8 11.3 12.2 59.66 270510-10moblm 001.MOV 1165 :38.83 lateral open 5.7 11.2 12.1 58.93 270510-10moblm 001.MOV 1372 :45.73 lateral partial 5.7 5.7 8.5 28.63 270510-10moblm 001.MOV 1651 :55.03 lateral partial 5.8 3.8 7.5 18.91 270510-10moblm 002.MOV 459 :15.30 frontal open 5.8 13.2 12.3 70.73 270510-10moblm 002.MOV 579 :19.30 lateral partial 5.8 4.4 7.8 21.95 270510-10moblm 002.MOV 853 :28.43 oblique partial 5.7 3.3 7.8 16.36 270510-10moblm 002.MOV 1978 1:05.93 frontal open 4.4 9.8 11.9 51.17 270510-10moblm 5.7 15.6 0.4879 0.5245 11.8 10.9 86.53 270510-41moblm 001.MOV 167 :05.57 lateral partial 5.6 6.0 8.6 30.84 270510-41moblm 001.MOV 211 :07.03 lateral partial 5.6 5.7 8.4 29.57 270510-41moblm 001.MOV 551 :18.37 frontal open 4.1 10.5 13.4 56.49 270510-41moblm 001.MOV 826 :27.53 lateral partial 5.6 6.7 8.7 34.96 270510-41moblm 001.MOV 967 :32.23 frontal open 5.6 12.8 13.3 70.20 270510-41moblm 001.MOV 1186 :39.53 frontal open 5.7 12.5 12.9 68.38 270510-41moblm 001.MOV 1212 :40.40 oblique partial 5.6 7.5 9.8 39.12 270510-41moblm 002.MOV 270510-41moblm 5.6 12.8 0.4879 0.5245 11.5 10.7 70.20 270510-42moblm 001.MOV 270510-42moblm 002.MOV 270510-43moblm 001.MOV 103 :03.43 lateral partial 5.5 5.6 8.4 29.90 270510-43moblm 001.MOV 123 :04.10 lateral open 5.4 12.7 12.1 71.56 270510-43moblm 001.MOV 125 :04.17 lateral open 5.5 12.5 12.7 70.26 270510-43moblm 001.MOV 133 :04.43 lateral partial 5.4 8.3 8.9 44.81 270510-43moblm 002.MOV 1888 1:02.93 lateral open 5.5 11.0 9.3 60.75 270510-43moblm 002.MOV 1890 1:03.00 lateral open 5.5 11.1 9.0 61.27 270510-43moblm 002.MOV 1926 1:04.20 oblique partial 5.5 4.2 7.6 21.95 270510-43moblm 5.5 12.7 0.4879 0.5245 11.2 10.4 71.56 270510-44moblm 001.MOV 10 :00.33 oblique partial 5.7 5.6 8.3 28.41 270510-44moblm 001.MOV 806 :26.87 lateral open 5.8 13.8 12.9 75.06

54

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 270510-44moblm 001.MOV 820 :27.33 oblique open 5.8 14.3 14.0 78.11 270510-44moblm 001.MOV 830 :27.67 lateral open 5.7 15.3 13.6 84.36 270510-44moblm 001.MOV 843 :28.10 post-lat open 4.9 15.5 14.6 86.25 270510-44moblm 001.MOV 1121 :37.37 oblique partial 5.7 7.7 8.9 39.48 270510-44moblm 001.MOV 1540 :51.33 frontal open 5.8 14.3 10.7 78.23 270510-44moblm 002.MOV 960 :32.00 frontal partial 4.8 2.4 6.4 11.33 270510-44moblm 002.MOV 1289 :42.97 lateral open 5.7 15.3 13.6 84.35 270510-44moblm 002.MOV 1292 :43.07 lateral open 5.8 14.1 12.1 76.69 270510-44moblm 002.MOV 1293 :43.10 lateral open 5.7 14.2 14.0 77.57 270510-44moblm 002.MOV 1309 :43.63 frontal open 5.7 15.4 12.7 85.23 270510-44moblm 002.MOV 1310 :43.67 frontal open 5.8 16.1 13.7 89.98 270510-44moblm 5.7 16.1 0.4879 0.5245 11.8 10.9 89.98 290510-4moblm 001.MOV 1760 :58.67 oblique partial 5.6 7.4 8.3 38.63 290510-4moblm 001.MOV 2250 1:15.00 frontal partial 5.6 7.4 7.5 38.73 290510-4moblm 002.MOV 293 :09.77 lateral partial 5.6 7.6 9.8 39.66 290510-4moblm 002.MOV 390 :13.00 lateral partial 5.6 7.0 8.7 36.47 290510-4moblm 002.MOV 1146 :38.20 lateral open 5.6 14.4 14.2 81.11 290510-4moblm 002.MOV 1165 :38.83 lateral open 5.6 14.0 12.5 78.18 290510-4moblm 002.MOV 1725 :57.50 lateral open 5.6 11.0 11.1 59.23 290510-4moblm 5.6 14.4 0.4879 0.5245 11.5 10.7 81.11 290510-40moblm 001.MOV 290510-40moblm 002.MOV 300510-03moblf 001.MOV 5 :00.17 oblique partial 5.6 5.7 8.7 29.66 300510-03moblf 001.MOV 10 :00.33 oblique partial 5.7 5.8 8.5 30.07 300510-03moblf 001.MOV 489 :16.30 lateral open 5.6 10.8 11.5 58.07 300510-03moblf 001.MOV 504 :16.80 lateral open 5.6 11.7 12.5 63.47 300510-03moblf 001.MOV 511 :17.03 lateral partial 5.6 6.5 9.0 34.02 300510-03moblf 5.6 11.7 0.4879 0.5245 11.5 10.7 63.47 300510-04moblm 001.MOV 24 :00.80 lateral partial 5.6 5.3 8.1 27.10 300510-04moblm 001.MOV 32 :01.07 lateral partial 5.7 4.7 7.8 24.05 300510-04moblm 001.MOV 269 :08.97 frontal open 4.3 10.3 12.1 55.00 300510-04moblm 001.MOV 485 :16.17 lateral partial 5.7 6.8 9.2 35.34 300510-04moblm 001.MOV 576 :19.20 lateral partial 5.6 5.3 8.4 27.36 300510-04moblm 001.MOV 606 :20.20 lateral open 5.7 13.9 13.5 76.74 300510-04moblm 001.MOV 610 :20.33 lateral open 5.7 14.5 13.1 80.83 300510-04moblm 001.MOV 612 :20.40 lateral open 5.7 15.5 13.7 87.75 300510-04moblm 001.MOV 614 :20.47 lateral open 5.7 14.8 13.7 83.12 300510-04moblm 001.MOV 615 :20.50 lateral open 5.7 14.3 12.4 79.47 300510-04moblm 001.MOV 616 :20.53 lateral open 5.6 13.8 12.9 75.92 300510-04moblm 001.MOV 630 :21.00 lateral open 5.6 13.0 13.1 71.10 300510-04moblm 001.MOV 632 :21.07 lateral open 5.7 12.6 12.4 68.70

55

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 300510-04moblm 001.MOV 633 :21.10 lateral open 5.6 13.0 12.8 71.09 300510-04moblm 001.MOV 673 :22.43 lateral open 5.7 15.0 14.0 83.87 300510-04moblm 001.MOV 674 :22.47 lateral open 5.6 13.2 11.5 72.19 300510-04moblm 001.MOV 708 :23.60 lateral partial 5.6 9.2 10.5 48.23 300510-04moblm 001.MOV 731 :24.37 lateral open 5.6 16.4 13.6 94.67 300510-04moblm 001.MOV 732 :24.40 lateral open 5.6 14.6 12.7 81.20 300510-04moblm 5.7 16.4 0.4879 0.5245 11.6 10.8 94.67 300510-05moblm 001.MOV 509 :16.97 oblique partial 5.7 4.5 8.0 22.90 300510-05moblm 001.MOV 564 :18.80 oblique partial 5.7 5.7 8.0 29.11 300510-05moblm 001.MOV 887 :29.57 oblique open 5.7 7.4 8.5 38.42 300510-05moblm 001.MOV 892 :29.73 oblique partial 5.7 6.0 8.6 30.66 300510-05moblm 002.MOV 15 :00.50 oblique open 5.7 10.1 9.9 53.47 300510-05moblm 002.MOV 482 :16.07 oblique open 5.6 9.5 10.5 50.01 300510-05moblm 002.MOV 511 :17.03 oblique open 5.7 12.3 12.1 66.34 300510-05moblm 002.MOV 534 :17.80 oblique open 5.7 12.6 11.8 68.41 300510-05moblm 002.MOV 564 :18.80 oblique open 5.7 11.2 11.1 59.93 300510-05moblm 002.MOV 574 :19.13 oblique open 5.7 11.5 11.7 61.44 300510-05moblm 002.MOV 661 :22.03 lateral open 5.7 10.4 11.2 55.03 300510-05moblm 002.MOV 672 :22.40 lateral open 5.7 13.4 12.5 72.97 300510-05moblm 002.MOV 696 :23.20 lateral open 5.7 13.4 12.4 73.47 300510-05moblm 002.MOV 697 :23.23 lateral open 5.6 13.2 11.9 72.28 300510-05moblm 002.MOV 698 :23.27 lateral open 5.7 14.0 11.7 77.12 300510-05moblm 002.MOV 699 :23.30 lateral open 5.7 13.5 12.1 73.61 300510-05moblm 002.MOV 709 :23.63 lateral open 5.6 12.2 11.8 65.98 300510-05moblm 002.MOV 737 :24.57 lateral partial 5.6 6.8 9.0 35.24 300510-05moblm 5.7 14.0 0.4879 0.5245 11.6 10.8 77.12 300510-06moblf 001.MOV 1715 :57.17 lateral partial 5.6 7.0 9.5 36.53 300510-06moblf 001.MOV 1717 :57.23 lateral partial 5.6 6.5 8.7 33.66 300510-06moblf 001.MOV 2083 1:09.43 oblique partial 5.6 5.8 8.9 29.91 300510-06moblf 001.MOV 2096 1:09.87 frontal open 5.6 13.3 13.2 73.25 300510-06moblf 001.MOV 2097 1:09.90 frontal open 5.6 14.6 13.0 81.56 300510-06moblf 001.MOV 2109 1:10.30 frontal open 4.8 14.5 12.0 81.16 300510-06moblf 5.6 14.6 0.4879 0.5245 11.5 10.7 81.56 300510-07moblm 001.MOV 625 :20.83 frontal open 5.7 8.7 11.4 44.74 300510-07moblm 001.MOV 645 :21.50 frontal open 5.8 14.5 10.3 78.94 300510-07moblm 001.MOV 669 :22.30 frontal open 5.7 12.9 10.9 69.00 300510-07moblm 001.MOV 807 :26.90 lateral partial 5.7 5.5 8.9 27.88 300510-07moblm 001.MOV 1291 :43.03 oblique partial 5.7 5.3 8.3 26.73 300510-07moblm 001.MOV 1449 :48.30 lateral partial 5.8 3.7 8.2 18.41 300510-07moblm 001.MOV 1531 :51.03 oblique partial 5.7 7.6 9.4 38.72 300510-07moblm 5.7 14.5 0.4879 0.5245 11.8 11.0 78.94

56

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 300510-08moblm 001.MOV 714 :23.80 frontal open 4.7 12.6 10.4 68.73 300510-08moblm 001.MOV 1012 :33.73 frontal open 3.6 8.5 10.6 44.56 300510-08moblm 001.MOV 1155 :38.50 lateral partial 5.6 3.2 7.6 16.04 300510-08moblm 001.MOV 1198 :39.93 lateral partial 5.6 4.1 9.1 20.89 300510-08moblm 001.MOV 1430 :47.67 lateral partial 5.6 2.4 7.5 11.46 300510-08moblm 001.MOV 1435 :47.83 frontal partial 5.1 3.8 8.5 19.18 300510-08moblm 001.MOV 1725 1:00.30 lateral partial 5.7 3.4 8.6 17.36 300510-08moblm 001.MOV 1809 1:00.30 lateral partial 5.6 3.2 8.2 16.04 300510-08moblm 001.MOV 1916 1:03.87 frontal partial 4.9 2.6 6.8 12.98 300510-08moblm 5.6 12.6 0.4879 0.5245 11.5 10.7 68.73 300510-09moblm 001.MOV 14 :00.47 oblique partial 5.5 4.1 7.6 21.55 300510-09moblm 001.MOV 19 :00.63 lateral partial 5.5 4.4 7.6 23.31 300510-09moblm 001.MOV 327 :10.90 oblique partial 5.4 3.8 7.6 19.72 300510-09moblm 001.MOV 334 :11.13 oblique partial 5.4 3.2 7.4 16.34 300510-09moblm 001.MOV 357 :11.90 oblique partial 5.5 3.3 7.6 17.28 300510-09moblm 001.MOV 535 :17.83 frontal open 5.5 15.1 11.2 88.62 300510-09moblm 001.MOV 536 :17.87 frontal open 5.5 16.6 15.8 100.26 300510-09moblm 5.5 16.6 0.4879 0.5245 11.2 10.4 100.26 300510-10moblf 001.MOV 199 :06.63 oblique partial 5.2 6.1 9.3 34.15 300510-10moblf 001.MOV 794 :26.47 oblique partial 5.2 2.5 6.8 13.08 300510-10moblf 001.MOV 1121 :37.37 frontal open 3.5 7.1 7.4 39.98 300510-10moblf 001.MOV 1194 :39.80 frontal open 4.6 7.9 9.9 45.08 300510-10moblf 001.MOV 1195 :39.83 frontal open 5.0 7.9 9.9 45.21 300510-10moblf 001.MOV 1242 :41.40 frontal open 5.0 9.7 9.8 55.91 300510-10moblf 001.MOV 1269 :42.30 frontal partial 4.2 6.8 8.9 38.41 300510-10moblf 001.MOV 1348 :44.93 oblique partial 5.2 5.2 7.5 28.91 300510-10moblf 001.MOV 1373 :48.13 oblique partial 5.2 4.2 7.5 23.10 300510-10moblf 5.2 9.7 0.4879 0.5245 10.7 9.9 55.91 310510-01moblf 001.MOV 175 :05.83 frontal open 4.1 8.5 7.3 47.09 310510-01moblf 001.MOV 637 :21.23 oblique partial 5.4 4.5 7.5 24.20 310510-01moblf 001.MOV 644 :21.47 oblique partial 5.4 3.4 6.3 17.84 310510-01moblf 001.MOV 648 :21.60 lateral partial 5.4 3.7 6.2 19.69 310510-01moblf 001.MOV 902 :30.07 oblique open 5.4 7.6 10.3 41.44 310510-01moblf 001.MOV 920 :30.67 frontal open 3.2 11.2 12.9 63.18 310510-01moblf 001.MOV 1256 :41.87 oblique partial 5.4 4.1 7.1 21.82 310510-01moblf 5.4 11.2 0.4879 0.5245 11.0 10.3 63.18 310510-02moblm 001.MOV 363 :12.10 oblique partial 5.2 6.9 8.2 38.46 310510-02moblm 001.MOV 385 :12.83 lateral open 5.3 9.8 11.2 56.04 310510-02moblm 001.MOV 391 :13.03 oblique open 5.3 13.9 13.1 83.49 310510-02moblm 001.MOV 392 :13.07 oblique open 5.2 13.7 12.9 82.36 310510-02moblm 001.MOV 401 :13.37 oblique open 5.3 12.5 10.9 73.89

57

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 310510-02moblm 001.MOV 402 :13.40 oblique open 5.3 13.7 12.6 82.15 310510-02moblm 001.MOV 403 :13.43 oblique open 5.3 13.3 12.3 79.73 310510-02moblm 001.MOV 423 :14.10 frontal open 5.3 13.0 11.9 77.20 310510-02moblm 001.MOV 424 :14.13 frontal open 5.3 13.6 11.5 81.85 310510-02moblm 001.MOV 425 :14.17 frontal open 5.3 15.1 12.5 92.84 310510-02moblm 001.MOV 438 :14.60 frontal open 5.3 13.6 11.7 81.94 310510-02moblm 001.MOV 461 :15.37 oblique partial 5.2 10.1 10.4 58.15 310510-02moblm 001.MOV 732 :24.40 lateral partial 5.3 5.8 8.3 32.43 310510-02moblm 002.MOV 93 :03.10 oblique open 5.3 12.2 12.5 71.78 310510-02moblm 002.MOV 95 :03.17 oblique open 5.3 11.6 12.4 67.90 310510-02moblm 002.MOV 148 :04.93 oblique partial 5.3 9.0 10.6 51.45 310510-02moblm 002.MOV 232 :07.73 oblique partial 5.2 6.8 8.8 38.08 310510-02moblm 002.MOV 372 :12.40 lateral open 5.3 10.2 10.8 58.94 310510-02moblm 002.MOV 393 :13.10 oblique open 5.3 12.5 13.4 74.11 310510-02moblm 002.MOV 396 :13.20 frontal open 5.3 12.6 13.2 74.62 310510-02moblm 002.MOV 400 :13.33 frontal open 5.3 12.2 12.8 71.98 310510-02moblm 5.3 15.1 0.4879 0.5245 10.8 10.0 92.84 310510-03moblm 001.MOV 487 :16.23 oblique partial 5.7 5.4 8.1 27.21 310510-03moblm 001.MOV 961 :32.03 lateral partial 5.7 9.1 10.1 47.70 310510-03moblm 001.MOV 970 :32.33 oblique open 5.7 10.3 10.6 54.32 310510-03moblm 001.MOV 1082 :36.07 lateral partial 5.7 6.4 9.5 32.87 310510-03moblm 001.MOV 1229 :40.97 lateral open 5.7 14.1 13.4 77.69 310510-03moblm 001.MOV 1230 :41.00 lateral open 5.7 13.2 12.9 71.78 310510-03moblm 001.MOV 1231 :41.03 lateral open 5.7 13.4 13.0 73.28 310510-03moblm 001.MOV 1232 :41.07 lateral open 5.7 13.9 12.4 76.56 310510-03moblm 001.MOV 1233 :41.10 lateral open 5.7 13.9 12.5 76.07 310510-03moblm 001.MOV 1234 :41.13 lateral open 5.7 13.5 12.5 73.87 310510-03moblm 001.MOV 1296 :43.20 lateral open 5.7 13.0 13.3 70.81 310510-03moblm 001.MOV 1304 :43.47 lateral open 5.7 14.4 14.1 79.71 310510-03moblm 001.MOV 1305 :43.50 lateral open 5.7 15.2 13.6 85.26 310510-03moblm 001.MOV 1307 :43.57 lateral open 5.7 14.5 13.9 80.29 310510-03moblm 001.MOV 1308 :43.60 lateral open 5.7 14.7 14.2 81.43 310510-03moblm 001.MOV 1331 :44.37 lateral open 5.6 15.6 13.0 87.58 310510-03moblm 001.MOV 1334 :44.47 lateral open 5.7 15.3 11.6 85.45 310510-03moblm 001.MOV 1344 :44.80 postlat open 5.5 15.8 11.3 89.07 310510-03moblm 001.MOV 1351 :45.03 postlat open 4.0 14.5 13.0 80.24 310510-03moblm 001.MOV 1362 :45.40 lateral open 5.7 15.4 13.8 86.45 310510-03moblm 001.MOV 1363 :45.43 lateral open 5.7 15.4 13.0 86.01 310510-03moblm 001.MOV 1364 :45.47 lateral open 5.7 15.0 13.6 83.76 310510-03moblm 001.MOV 1366 :45.53 lateral open 5.7 15.1 12.5 84.49 310510-03moblm 5.7 15.8 0.4879 0.5245 11.7 10.8 89.07

58

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 310510-04moblf 001.MOV 3 :00.10 frontal open 6.3 9.4 9.6 44.10 310510-04moblf 001.MOV 23 :00.77 frontal open 6.3 8.2 9.4 38.07 310510-04moblf 001.MOV 43 :01.43 frontal open 6.3 7.1 8.4 32.95 310510-04moblf 001.MOV 111 :03.67 frontal open 6.4 12.0 10.7 57.31 310510-04moblf 001.MOV 175 :05.83 oblique partial 6.3 6.9 8.2 32.11 310510-04moblf 001.MOV 236 :07.87 oblique open 6.3 9.9 11.7 46.87 310510-04moblf 001.MOV 238 :07.93 oblique open 6.3 10.3 11.9 48.97 310510-04moblf 001.MOV 254 :08.47 oblique open 6.3 11.4 11.2 54.43 310510-04moblf 001.MOV 262 :08.73 oblique open 6.4 11.0 11.3 52.37 310510-04moblf 001.MOV 264 :08.80 oblique open 6.3 11.9 11.9 56.74 310510-04moblf 001.MOV 270 :09.00 oblique open 6.3 12.6 12.9 60.94 310510-04moblf 001.MOV 271 :09.03 oblique open 6.2 13.3 13.1 64.31 310510-04moblf 001.MOV 272 :09.07 oblique open 6.3 12.5 11.1 60.39 310510-04moblf 001.MOV 275 :09.17 oblique open 6.3 11.5 11.6 55.10 310510-04moblf 001.MOV 276 :09.20 oblique open 6.3 11.9 11.7 56.96 310510-04moblf 001.MOV 294 :09.80 frontal open 6.3 14.5 13.1 71.25 310510-04moblf 001.MOV 296 :09.87 oblique open 6.3 12.8 11.9 61.66 310510-04moblf 001.MOV 305 :10.17 oblique open 6.3 12.5 12.4 60.24 310510-04moblf 001.MOV 330 :11.00 frontal open 5.4 11.2 10.6 53.41 310510-04moblf 001.MOV 340 :11.33 oblique open 6.3 11.0 10.9 52.32 310510-04moblf 001.MOV 442 :14.73 lateral partial 6.3 9.5 10.2 44.75 310510-04moblf 001.MOV 478 :15.93 lateral open 6.3 12.2 11.7 58.59 310510-04moblf 001.MOV 554 :18.47 lateral partial 6.3 8.1 9.7 37.83 310510-04moblf 001.MOV 652 :21.73 oblique open 6.3 12.3 12.8 59.09 310510-04moblf 001.MOV 653 :21.77 oblique open 6.3 11.9 12.3 56.91 310510-04moblf 001.MOV 656 :21.87 oblique open 6.3 10.2 10.7 48.40 310510-04moblf 001.MOV 740 :24.67 oblique partial 6.3 7.9 9.4 36.63 310510-04moblf 001.MOV 851 :28.37 lateral partial 6.3 6.6 8.4 30.46 310510-04moblf 001.MOV 1011 :33.70 oblique open 6.3 11.9 12.0 57.14 310510-04moblf 001.MOV 1221 :40.70 frontal open 6.3 12.2 13.6 58.73 310510-04moblf 001.MOV 1262 :42.07 frontal open 6.2 13.8 11.5 67.06 310510-04moblf 001.MOV 1265 :42.17 frontal open 6.3 14.1 14.0 69.09 310510-04moblf 001.MOV 1317 :43.90 frontal open 4.5 13.4 13.8 65.09 310510-04moblf 001.MOV 1320 :44.00 frontal open 4.7 14.0 14.7 68.21 310510-04moblf 6.3 14.5 0.4879 0.5245 12.9 12.0 71.25 310510-05moblm 001.MOV 28 :00.93 frontal open 6.3 9.6 8.1 45.02 310510-05moblm 001.MOV 118 :03.93 frontal open 6.3 13.4 13.1 64.80 310510-05moblm 001.MOV 173 :05.77 frontal open 6.1 13.1 12.4 63.07 310510-05moblm 001.MOV 186 :06.20 frontal open 6.0 13.8 14.0 66.81 310510-05moblm 001.MOV 191 :06.37 oblique open 6.3 12.4 12.1 59.32 310510-05moblm 001.MOV 198 :06.60 oblique open 6.3 13.2 10.9 63.46

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 310510-05moblm 001.MOV 206 :06.87 clipped open 6.2 14.9 14.9 72.89 310510-05moblm 001.MOV 216 :07.20 lateral open 6.3 15.4 13.9 76.08 310510-05moblm 001.MOV 217 :07.23 lateral open 6.3 14.0 13.4 68.01 310510-05moblm 001.MOV 233 :07.77 oblique open 6.4 14.0 12.2 67.64 310510-05moblm 001.MOV 296 :09.87 lateral open 6.3 13.6 12.0 65.57 310510-05moblm 001.MOV 305 :10.17 postlat open 4.3 13.2 13.0 63.61 310510-05moblm 001.MOV 347 :11.57 lateral open 6.3 14.6 14.5 71.42 310510-05moblm 001.MOV 351 :11.70 oblique open 6.3 14.8 13.3 72.40 310510-05moblm 001.MOV 389 :12.97 oblique partial 6.3 8.1 10.4 37.59 310510-05moblm 001.MOV 418 :13.93 oblique partial 6.3 8.4 10.8 39.19 310510-05moblm 001.MOV 424 :14.13 oblique open 6.3 13.2 12.8 63.66 310510-05moblm 001.MOV 427 :14.23 oblique open 6.3 13.3 13.2 64.24 310510-05moblm 001.MOV 521 :17.37 oblique open 6.3 12.8 12.8 61.18 310510-05moblm 001.MOV 768 :25.60 oblique open 6.3 13.6 13.3 65.83 310510-05moblm 001.MOV 783 :26.10 postlat open 4.6 13.9 14.8 67.44 310510-05moblm 6.3 15.4 0.4879 0.5245 13.0 12.1 76.08 310510-06moblm 001.MOV 234 :07.80 frontal open 5.9 12.8 11.9 66.27 310510-06moblm 001.MOV 240 :08.00 frontal open 5.9 12.9 11.8 67.34 310510-06moblm 001.MOV 330 :11.00 oblique partial 5.9 8.1 9.7 40.38 310510-06moblm 001.MOV 333 :11.10 oblique partial 5.9 6.8 8.9 33.54 310510-06moblm 001.MOV 382 :12.73 oblique open 5.9 12.1 12.7 62.44 310510-06moblm 001.MOV 384 :12.80 oblique open 5.9 13.3 12.7 69.23 310510-06moblm 001.MOV 398 :13.27 postlat open 5.4 13.4 15.4 70.00 310510-06moblm 001.MOV 399 :13.30 postlat open 5.6 13.3 15.1 69.23 310510-06moblm 001.MOV 402 :13.40 postlat open 4.8 12.7 15.0 66.09 310510-06moblm 001.MOV 411 :13.70 postlat open 4.6 14.3 13.2 75.81 310510-06moblm 001.MOV 412 :13.73 postlat open 5.4 15.9 14.3 85.91 310510-06moblm 001.MOV 484 :16.13 oblique partial 5.9 7.8 9.6 39.12 310510-06moblm 001.MOV 539 :17.97 lateral partial 5.9 6.2 8.7 30.46 310510-06moblm 001.MOV 782 :26.07 postlat open 4.6 13.0 14.4 67.50 310510-06moblm 001.MOV 836 :27.87 lateral open 5.9 11.7 11.6 60.11 310510-06moblm 5.9 15.9 0.4879 0.5245 12.1 11.2 85.91 260510-6-ptqum 001.MOV 1 :00.03 lateral partial 5.7 3.6 7.8 20.41 260510-6-ptqum 001.MOV 27 :00.90 lateral open 5.7 8.2 9.4 48.31 260510-6-ptqum 001.MOV 55 :01.83 lateral partial 5.7 4.4 7.2 25.20 260510-6-ptqum 001.MOV 128 :04.27 lateral open 5.7 6.5 7.8 37.92 260510-6-ptqum 001.MOV 172 :05.73 lateral open 5.7 7.7 8.1 45.44 260510-6-ptqum 001.MOV 249 :08.30 lateral open 5.7 8.1 8.7 48.13 260510-6-ptqum 001.MOV 251 :08.36 lateral open 5.7 7.1 7.6 41.40 260510-6-ptqum 001.MOV 264 :08.80 lateral open 5.6 8.3 7.4 49.29 260510-6-ptqum 001.MOV 663 :22.10 lateral open 5.7 8.4 8.8 50.02

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 260510-6-ptqum 001.MOV 709 :23.63 lateral open 5.7 7.7 9.0 45.44 260510-6-ptqum 002.MOV 35 :01.17 lateral open 5.7 7.8 8.0 46.28 260510-6-ptqum 002.MOV 104 :03.47 lateral partial 5.7 6.2 8.7 36.21 260510-6-ptqum 002.MOV 394 :13.13 lateral open 5.6 9.9 10.6 59.26 260510-6-ptqum 002.MOV 395 :13.17 lateral open 5.7 10.3 10.3 62.25 260510-6-ptqum 002.MOV 428 :14.27 lateral partial 5.7 2.5 7.2 13.72 260510-6-ptqum 002.MOV 441 :14.70 postlat open 5.1 7.3 8.3 42.58 260510-6-ptqum 5.7 10.3 0.5504 0.5933 10.3 9.6 62.25 260510-10-ptqum 001.MOV 260510-10-ptqum 002.MOV 767 :25.57 oblique open 5.7 7.4 7.5 43.45 260510-10-ptqum 002.MOV 1401 :46.70 oblique open 5.7 10.2 9.3 61.66 260510-10-ptqum 002.MOV 1403 :46.77 frontal partial 5.7 8.7 9.6 51.42 260510-10-ptqum 002.MOV 1456 :48.53 lateral partial 5.7 5.6 7.1 32.41 260510-10-ptqum 002.MOV 1476 :49.20 lateral open 5.7 6.1 7.6 35.21 260510-10-ptqum 002.MOV 1510 :50.33 lateral open 5.7 4.0 8.1 22.77 260510-10-ptqum 002.MOV 1618 :53.93 lateral open 5.7 5.6 8.3 32.10 260510-10-ptqum 002.MOV 1676 :55.87 oblique partial 5.7 4.6 7.2 26.32 260510-10-ptqum 5.7 10.2 0.5504 0.5933 10.3 9.6 61.66 260510-20-ptqum 001.MOV 528 :17.60 lateral partial 5.6 3.5 6.8 20.22 260510-20-ptqum 001.MOV 530 :17.67 lateral partial 5.6 3.5 7.2 19.99 260510-20-ptqum 001.MOV 533 :17.77 lateral partial 5.6 5.0 8.0 29.29 260510-20-ptqum 001.MOV 1063 :35.43 lateral open 5.6 7.7 7.9 45.95 260510-20-ptqum 001.MOV 1085 :36.17 lateral open 5.7 8.3 7.8 49.90 260510-20-ptqum 001.MOV 1162 :38.73 lateral partial 5.6 4.2 7.8 24.03 260510-20-ptqum 001.MOV 1279 :42.63 oblique partial 5.6 4.5 6.2 26.38 260510-20-ptqum 001.MOV 1419 :47.30 frontal open 2.7 9.0 8.2 54.16 260510-20-ptqum 001.MOV 1435 :47.83 frontal open 5.4 9.7 6.9 58.97 260510-20-ptqum 001.MOV 1436 :47.87 frontal open 5.1 10.6 7.5 65.17 260510-20-ptqum 001.MOV 1437 :47.90 frontal open 5.1 10.6 7.3 64.98 260510-20-ptqum 001.MOV 1439 :47.97 frontal open 3.0 9.5 8.3 57.37 260510-20-ptqum 001.MOV 1445 :48.17 frontal open 4.4 7.3 4.6 43.12 260510-20-ptqum 001.MOV 1452 :48.40 oblique partial 5.6 2.9 6.9 16.23 260510-20-ptqum 001.MOV 1566 :52.20 lateral partial 5.6 3.7 7.5 21.12 260510-20-ptqum 002.MOV 70 :02.33 lateral partial 5.6 3.6 6.9 20.40 260510-20-ptqum 002.MOV 269 :08.97 lateral partial 5.6 3.3 6.9 18.57 260510-20-ptqum 5.6 10.6 0.5504 0.5933 10.2 9.5 65.17 260510-21-ptqum 001.MOV 5 :00.17 lateral open 5.7 8.1 8.9 48.15 260510-21-ptqum 001.MOV 34 :01.13 lateral partial 5.7 4.5 7.4 25.91 260510-21-ptqum 001.MOV 365 :12.17 lateral open 5.7 9.7 10.3 58.45 260510-21-ptqum 001.MOV 370 :12.33 lateral open 5.7 7.0 8.7 41.02 260510-21-ptqum 001.MOV 536 :17.87 lateral open 5.6 8.4 8.8 49.65

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 260510-21-ptqum 001.MOV 641 :21.37 lateral open 5.6 8.5 9.6 50.62 260510-21-ptqum 001.MOV 731 :24.37 oblique partial 5.7 4.9 7.9 28.40 260510-21-ptqum 001.MOV 1268 :42.27 lateral open 5.7 6.6 8.0 38.26 260510-21-ptqum 002.MOV 78 :02.60 lateral open 5.7 7.8 9.3 45.84 260510-21-ptqum 002.MOV 108 :03.60 lateral open 5.6 8.5 9.2 50.35 260510-21-ptqum 002.MOV 553 :18.43 lateral open 5.7 5.3 7.3 30.77 260510-21-ptqum 002.MOV 819 :27.30 frontal partial 4.8 5.9 7.1 34.34 260510-21-ptqum 5.7 9.7 0.5504 0.5933 10.3 9.6 58.45 260510-23-ptqum 001.MOV 398 :13.27 frontal open 3.1 9.2 10.3 55.63 260510-23-ptqum 001.MOV 469 :15.63 frontal open 2.5 8.7 10.0 52.47 260510-23-ptqum 001.MOV 507 :16.90 oblique partial 5.7 6.3 8.5 37.08 260510-23-ptqum 001.MOV 663 :22.10 oblique partial 5.7 6.3 8.1 36.74 260510-23-ptqum 001.MOV 819 :27.30 lateral open 5.6 11.5 10.3 71.11 260510-23-ptqum 002.MOV 82 :02.73 lateral open 5.6 9.0 9.4 53.88 260510-23-ptqum 002.MOV 240 :08.00 oblique open 5.6 11.9 10.9 74.21 260510-23-ptqum 002.MOV 241 :08.03 oblique open 5.6 11.8 9.7 73.04 260510-23-ptqum 002.MOV 862 :28.73 lateral partial 5.6 5.6 8.7 32.99 260510-23-ptqum 002.MOV 934 :31.13 lateral open 5.6 11.4 9.8 70.60 260510-23-ptqum 002.MOV 941 :31.37 lateral open 5.6 10.9 9.7 67.19 260510-23-ptqum 5.6 11.9 0.5504 0.5933 10.2 9.5 74.21 260510-24-ptqum 001.MOV 294 :09.80 lateral partial 5.3 6.0 7.2 37.40 260510-24-ptqum 001.MOV 298 :09.93 lateral partial 5.3 3.9 6.8 23.51 260510-24-ptqum 001.MOV 302 :10.07 lateral partial 5.3 5.5 8.1 34.33 260510-24-ptqum 001.MOV 458 :15.27 frontal open 5.0 8.2 7.5 51.87 260510-24-ptqum 001.MOV 1199 :39.97 oblique partial 5.3 3.2 7.1 19.38 260510-24-ptqum 002.MOV 135 :04.50 lateral open 5.4 7.4 7.7 46.33 260510-24-ptqum 002.MOV 142 :04.73 lateral open 5.4 9.3 9.4 59.42 260510-24-ptqum 5.3 9.3 0.5504 0.5933 9.7 9.0 59.42 260510-25-ptqum 001.MOV 125 :04.17 lateral partial 5.7 2.2 7.3 11.78 260510-25-ptqum 001.MOV 205 :06.83 lateral partial 5.7 1.3 6.7 5.89 260510-25-ptqum 001.MOV 330 :11.00 lateral partial 5.7 2.8 7.7 15.47 260510-25-ptqum 001.MOV 731 :24.37 oblique partial 5.7 2.1 7.1 11.38 260510-25-ptqum 001.MOV 2043 1:08.10 frontal open 4.6 6.5 7.8 37.43 260510-25-ptqum 001.MOV 2151 1:11.70 frontal open 4.4 7.8 8.5 45.19 260510-25-ptqum 001.MOV 2283 1:16.10 lateral open 5.7 7.2 9.1 42.02 260510-25-ptqum 001.MOV 2338 1:17.93 frontal open 4.6 8.7 9.1 51.09 260510-25-ptqum 001.MOV 2444 1:21.47 lateral partial 5.8 7.1 8.5 41.11 260510-25-ptqum 002.MOV 462 :15.40 oblique partial 5.7 3.8 8.5 21.65 260510-25-ptqum 002.MOV 1312 :43.73 lateral partial 5.7 3.1 8.0 17.51 260510-25-ptqum 5.7 8.7 0.5504 0.5933 10.4 9.7 51.09 260510-40-ptqum 001.MOV 794 :26.47 frontal open 3.5 7.9 7.8 45.25

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 260510-40-ptqum 001.MOV 814 :27.13 frontal open 3.0 8.9 8.4 51.65 260510-40-ptqum 001.MOV 815 :27.17 frontal open 3.1 9.3 8.6 53.74 260510-40-ptqum 001.MOV 1191 :39.70 lateral partial 5.8 3.8 7.1 21.11 260510-40-ptqum 001.MOV 1193 :39.77 lateral partial 5.9 5.7 8.2 32.30 260510-40-ptqum 001.MOV 1207 :40.23 lateral open 5.9 6.9 7.9 38.88 260510-40-ptqum 002.MOV 135 :04.50 frontal partial 4.7 4.9 6.6 27.56 260510-40-ptqum 002.MOV 208 :06.93 lateral partial 5.9 1.5 5.8 7.49 260510-40-ptqum 003.MOV 73 :02.43 oblique partial 5.8 5.1 8.3 28.24 260510-40-ptqum 003.MOV 502 :16.73 oblique partial 5.8 4.8 7.8 26.95 260510-40-ptqum 003.MOV 510 :17.00 frontal open 2.3 8.5 8.7 48.93 260510-40-ptqum 003.MOV 834 :27.80 frontal open 2.4 7.7 7.9 44.23 260510-40-ptqum 5.8 9.3 0.5504 0.5933 10.6 9.9 53.74 270510-8-ptqum 001.MOV 5 :00.17 lateral open 5.7 8.7 9.5 51.60 270510-8-ptqum 001.MOV 26 :00.87 oblique partial 5.7 6.9 8.2 39.88 270510-8-ptqum 001.MOV 42 :01.40 frontal open 5.2 9.8 8.9 58.48 270510-8-ptqum 001.MOV 113 :03.77 lateral open 5.7 9.6 9.5 57.40 270510-8-ptqum 001.MOV 114 :03.80 lateral open 5.7 11.0 10.5 66.70 270510-8-ptqum 001.MOV 115 :03.83 lateral open 5.7 10.5 9.7 63.09 270510-8-ptqum 001.MOV 250 :08.33 frontal open 3.8 7.3 4.6 42.52 270510-8-ptqum 001.MOV 445 :14.83 oblique partial 5.7 5.1 8.3 29.36 270510-8-ptqum 001.MOV 481 :16.03 lateral open 5.7 8.2 8.3 48.44 270510-8-ptqum 002.MOV 71 :02.37 frontal open 2.5 9.0 10.7 53.43 270510-8-ptqum 002.MOV 146 :04.87 postlat open 4.5 10.1 10.5 60.67 270510-8-ptqum 002.MOV 177 :05.90 lateral open 5.7 9.5 9.8 56.26 270510-8-ptqum 5.7 11.0 0.5504 0.5933 10.4 9.6 66.70 270510-7-ptquf 001.MOV 27 :00.90 lateral partial 5.2 6.5 6.9 41.93 270510-7-ptquf 001.MOV 41 :01.37 lateral open 5.2 8.8 9.2 57.84 270510-7-ptquf 001.MOV 52 :01.73 oblique partial 5.1 2.6 6.7 15.60 270510-7-ptquf 001.MOV 58 :01.93 oblique partial 5.2 2.1 6.5 12.74 270510-7-ptquf 001.MOV 79 :02.63 oblique open 5.2 7.3 8.4 47.16 270510-7-ptquf 001.MOV 81 :02.70 oblique partial 5.2 4.8 7.5 30.54 270510-7-ptquf 001.MOV 180 :06.00 oblique partial 5.1 2.1 6.1 12.85 270510-7-ptquf 001.MOV 390 :13.00 oblique open 5.2 5.6 7.7 35.59 270510-7-ptquf 001.MOV 411 :13.70 lateral partial 5.3 2.2 6.6 13.29 270510-7-ptquf 001.MOV 663 :22.10 lateral open 5.3 5.7 8.5 36.31 270510-7-ptquf 001.MOV 666 :23.20 lateral partial 5.3 3.9 7.4 24.07 270510-7-ptquf 002.MOV 397 :13.23 oblique partial 5.2 2.8 6.3 17.46 270510-7-ptquf 002.MOV 420 :14.00 lateral partial 5.2 3.1 7.1 18.86 270510-7-ptquf 002.MOV 514 :17.13 frontal open 4.7 9.4 7.8 62.02 270510-7-ptquf 002.MOV 519 :17.30 frontal open 4.5 10.3 9.1 68.70 270510-7-ptquf 002.MOV 716 :23.87 lateral partial 5.2 4.4 7.5 27.84

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 270510-7-ptquf 5.2 10.3 0.5504 0.5933 9.4 8.8 68.70 270510-21-ptquf 001.MOV 32 :01.07 lateral open 5.4 7.2 8.9 45.05 270510-21-ptquf 001.MOV 38 :01.27 lateral open 5.4 9.9 9.5 63.38 270510-21-ptquf 001.MOV 44 :01.47 lateral open 5.4 9.2 9.4 58.59 270510-21-ptquf 001.MOV 76 :02.53 oblique partial 5.4 5.4 7.0 32.83 270510-21-ptquf 001.MOV 153 :05.10 oblique partial 5.4 3.0 6.6 17.84 270510-21-ptquf 001.MOV 159 :05.30 oblique partial 5.4 2.9 6.8 17.51 270510-21-ptquf 001.MOV 167 :05.57 oblique partial 5.4 3.0 6.7 17.57 270510-21-ptquf 001.MOV 219 :07.30 oblique partial 5.4 2.1 6.6 12.28 270510-21-ptquf 001.MOV 291 :09.70 lateral open 5.4 9.5 11.4 60.74 270510-21-ptquf 001.MOV 294 :09.80 lateral open 5.4 9.0 8.7 57.07 270510-21-ptquf 002.MOV 5 :00.17 oblique open 5.4 8.2 9.6 51.66 270510-21-ptquf 002.MOV 13 :00.43 oblique open 5.4 10.6 10.5 68.19 270510-21-ptquf 002.MOV 14 :00.47 oblique open 5.4 10.8 10.4 70.22 270510-21-ptquf 002.MOV 18 :00.60 oblique open 5.4 10.5 10.5 67.73 270510-21-ptquf 002.MOV 38 :01.27 oblique open 5.4 10.5 10.1 67.73 270510-21-ptquf 002.MOV 339 :11.30 lateral open 5.4 6.6 8.3 40.68 270510-21-ptquf 002.MOV 353 :11.77 lateral open 5.4 10.8 10.2 70.32 270510-21-ptquf 002.MOV 375 :12.50 lateral open 5.4 8.7 8.9 54.62 270510-21-ptquf 5.4 10.8 0.5504 0.5933 9.8 9.1 70.32 270510-24-ptquf 001.MOV 87 :02.90 oblique partial 5.5 4.4 7.7 26.12 270510-24-ptquf 001.MOV 96 :03.20 lateral open 5.5 8.5 9.6 53.10 270510-24-ptquf 001.MOV 162 :05.40 lateral partial 5.4 4.4 7.4 26.63 270510-24-ptquf 001.MOV 164 :05.47 oblique partial 5.5 2.9 6.7 16.90 270510-24-ptquf 001.MOV 327 :10.90 oblique partial 5.4 3.7 6.4 21.85 270510-24-ptquf 001.MOV 374 :12.47 oblique open 5.5 9.5 10.0 59.38 270510-24-ptquf 001.MOV 375 :12.50 oblique open 5.4 10.1 10.0 63.75 270510-24-ptquf 001.MOV 378 :12.60 oblique open 5.5 9.1 9.7 57.15 270510-24-ptquf 001.MOV 388 :12.93 oblique open 5.4 9.9 10.0 62.29 270510-24-ptquf 001.MOV 418 :13.93 oblique open 5.5 7.0 9.3 42.66 270510-24-ptquf 001.MOV 492 :16.40 oblique open 5.5 6.7 9.3 40.69 270510-24-ptquf 001.MOV 500 :16.67 oblique partial 5.4 4.2 7.9 25.27 270510-24-ptquf 001.MOV 535 :17.83 lateral partial 5.4 8.3 9.8 51.25 270510-24-ptquf 001.MOV 542 :18.07 lateral partial 5.5 8.1 9.3 50.24 270510-24-ptquf 001.MOV 652 :21.73 oblique open 5.4 2.3 6.7 13.09 270510-24-ptquf 001.MOV 787 :26.23 oblique open 5.4 8.4 10.3 52.21 270510-24-ptquf 001.MOV 855 :28.50 oblique partial 5.4 3.9 7.1 23.40 270510-24-ptquf 001.MOV 1241 :41.37 oblique partial 5.4 8.3 9.0 51.75 270510-24-ptquf 5.4 10.1 0.5504 0.5933 9.9 9.2 63.75 270510-45-ptquf 001.MOV 89 :02.97 lateral partial 4.9 4.3 7.1 29.07 270510-45-ptquf 001.MOV 98 :03.27 lateral partial 4.8 4.1 6.5 27.67

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 270510-45-ptquf 001.MOV 182 :06.07 oblique partial 4.8 1.6 5.8 10.06 270510-45-ptquf 001.MOV 522 :17.40 frontal open 3.6 8.1 9.5 56.89 270510-45-ptquf 001.MOV 606 :20.20 oblique partial 4.8 4.4 7.1 29.37 270510-45-ptquf 001.MOV 901 :30.03 lateral open 4.9 8.0 7.7 55.61 270510-45-ptquf 001.MOV 906 :30.20 lateral open 4.8 9.0 9.4 63.84 270510-45-ptquf 001.MOV 922 :30.73 oblique partial 4.8 1.7 6.3 10.46 270510-45-ptquf 002.MOV 14 :00.47 oblique partial 4.8 4.2 8.0 28.41 270510-45-ptquf 002.MOV 45 :01.50 frontal open 3.9 7.9 9.2 54.77 270510-45-ptquf 002.MOV 46 :01.53 frontal open 3.9 6.8 8.5 46.83 270510-45-ptquf 002.MOV 55 :01.83 oblique partial 4.9 5.4 7.5 36.89 270510-45-ptquf 002.MOV 93 :03.10 frontal open 4.3 9.6 7.1 68.46 270510-45-ptquf 002.MOV 94 :03.13 frontal open 4.6 9.4 7.4 66.56 270510-45-ptquf 002.MOV 95 :03.17 frontal open 4.6 9.9 7.4 71.11 270510-45-ptquf 002.MOV 116 :03.87 oblique open 4.9 8.7 9.9 61.08 270510-45-ptquf 002.MOV 172 :05.73 oblique partial 4.9 2.4 6.3 15.66 270510-45-ptquf 002.MOV 244 :08.13 lateral open 4.9 7.6 8.3 52.66 270510-45-ptquf 002.MOV 420 :14.00 oblique partial 4.8 5.6 7.1 37.87 270510-45-ptquf 002.MOV 438 :14.60 oblique open 4.9 9.1 9.4 64.76 270510-45-ptquf 002.MOV 442 :14.73 oblique open 4.9 7.6 8.6 53.01 270510-45-ptquf 002.MOV 447 :14.90 oblique open 4.9 8.7 8.8 60.96 270510-45-ptquf 002.MOV 636 :21.20 oblique open 4.9 9.5 9.8 67.92 270510-45-ptquf 002.MOV 637 :21.23 oblique open 4.9 9.6 9.8 68.30 270510-45-ptquf 002.MOV 755 :25.17 frontal open 3.8 9.1 8.7 64.13 270510-45-ptquf 002.MOV 758 :25.27 frontal open 4.3 9.3 8.7 66.33 270510-45-ptquf 002.MOV 772 :25.73 frontal open 3.7 9.0 8.4 63.69 270510-45-ptquf 002.MOV 1243 :41.43 lateral open 4.8 6.5 7.2 44.33 270510-45-ptquf 4.9 9.9 0.5504 0.5933 8.8 8.2 71.11 290510-2-ptqum 001.MOV 0 :00.00 oblique partial 5.1 4.0 6.9 24.86 290510-2-ptqum 001.MOV 29 :00.97 lateral open 5.2 8.6 8.7 56.38 290510-2-ptqum 001.MOV 88 :02.93 oblique partial 5.2 5.2 7.3 33.10 290510-2-ptqum 001.MOV 179 :05.97 frontal open 4.4 8.6 9.4 56.39 290510-2-ptqum 001.MOV 181 :06.03 oblique open 5.2 9.0 9.3 59.06 290510-2-ptqum 001.MOV 182 :06.07 oblique open 5.2 8.7 9.5 57.18 290510-2-ptqum 001.MOV 191 :06.37 frontal open 4.8 8.2 6.9 53.50 290510-2-ptqum 001.MOV 263 :08.77 oblique open 5.2 12.1 11.3 83.52 290510-2-ptqum 001.MOV 321 :10.70 frontal partial 4.7 5.3 6.8 33.50 290510-2-ptqum 001.MOV 330 :11.00 oblique partial 5.2 4.2 7.6 26.58 290510-2-ptqum 001.MOV 363 :12.10 lateral open 5.1 10.7 10.3 71.82 290510-2-ptqum 5.2 12.1 0.5504 0.5933 9.4 8.7 83.52 290510-9-ptquf 001.MOV 380 :12.67 lateral partial 5.5 3.2 7.1 19.00 290510-9-ptquf 001.MOV 1033 :34.43 oblique open 5.4 8.2 6.5 50.53

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U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 290510-9-ptquf 001.MOV 1041 :34.70 oblique open 5.5 9.6 9.1 60.58 290510-9-ptquf 001.MOV 1045 :34.83 oblique open 5.4 9.6 8.8 60.36 290510-9-ptquf 001.MOV 1054 :35.13 oblique partial 5.4 4.0 7.6 23.65 290510-9-ptquf 001.MOV 1207 :40.23 oblique open 5.5 8.2 9.1 50.62 290510-9-ptquf 001.MOV 1212 :40.40 oblique partial 5.5 7.3 8.7 44.80 290510-9-ptquf 001.MOV 1227 :40.90 oblique open 5.4 8.7 9.0 54.05 290510-9-ptquf 001.MOV 1236 :41.20 oblique partial 5.5 5.5 7.9 33.40 290510-9-ptquf 001.MOV 1260 :42.00 oblique partial 5.4 4.2 7.2 24.94 290510-9-ptquf 001.MOV 1313 :43.77 lateral partial 5.5 3.0 7.1 17.31 290510-9-ptquf 001.MOV 1327 :44.23 oblique partial 5.4 7.5 7.6 46.28 290510-9-ptquf 5.4 9.6 0.5504 0.5933 9.9 9.2 60.58 290510-11-ptqum 001.MOV 7 :00.23 oblique partial 5.4 3.3 6.9 19.98 290510-11-ptqum 001.MOV 27 :00.90 oblique partial 5.3 1.9 6.6 10.77 290510-11-ptqum 001.MOV 30 :01.00 oblique open 5.3 8.0 10.0 50.24 290510-11-ptqum 001.MOV 41 :01.37 oblique open 5.4 11.3 9.7 73.55 290510-11-ptqum 001.MOV 42 :01.40 oblique partial 5.4 6.9 8.5 42.47 290510-11-ptqum 001.MOV 135 :04.50 oblique partial 5.4 6.3 8.1 38.58 290510-11-ptqum 001.MOV 196 :06.53 oblique open 5.3 8.9 8.5 55.96 290510-11-ptqum 001.MOV 198 :06.60 oblique open 5.4 9.8 9.4 62.45 290510-11-ptqum 001.MOV 207 :06.90 postlat open 1.6 9.4 10.2 59.85 290510-11-ptqum 001.MOV 241 :08.03 oblique partial 5.4 9.9 10.2 63.36 290510-11-ptqum 001.MOV 423 :14.10 oblique open 5.4 9.7 9.9 61.66 290510-11-ptqum 001.MOV 552 :18.40 frontal open 4.9 10.5 9.6 67.93 290510-11-ptqum 5.4 11.3 0.5504 0.5933 9.8 9.1 73.55 290510-41-ptquf 001.MOV 4 :00.13 lateral partial 5.3 2.0 6.6 11.79 290510-41-ptquf 001.MOV 13 :00.43 lateral partial 5.4 2.6 7.0 15.15 290510-41-ptquf 001.MOV 16 :00.53 lateral partial 5.4 3.5 7.5 20.85 290510-41-ptquf 001.MOV 43 :01.43 lateral partial 5.4 6.3 8.0 39.22 290510-41-ptquf 001.MOV 62 :02.07 lateral partial 5.3 5.4 8.0 32.92 290510-41-ptquf 001.MOV 199 :06.63 lateral open 5.3 8.2 8.8 51.41 290510-41-ptquf 001.MOV 324 :10.80 oblique partial 5.3 6.1 7.9 37.44 290510-41-ptquf 001.MOV 523 :17.43 lateral partial 5.4 5.8 7.7 35.50 290510-41-ptquf 001.MOV 533 :17.77 frontal open 3.3 9.1 7.6 58.20 290510-41-ptquf 001.MOV 543 :18.10 frontal open 3.2 8.7 8.0 55.01 290510-41-ptquf 001.MOV 544 :18.13 frontal open 3.2 8.0 7.7 50.04 290510-41-ptquf 001.MOV 552 :18.40 oblique open 5.3 9.6 9.2 61.70 290510-41-ptquf 001.MOV 675 :22.50 oblique partial 5.3 5.3 7.9 32.60 290510-41-ptquf 5.4 9.6 0.5504 0.5933 9.7 9.0 61.70 300510-01-ptquf 001.MOV 101 :03.37 oblique open 5.0 9.5 9.6 64.70 300510-01-ptquf 001.MOV 105 :03.50 oblique open 5.0 7.8 8.2 52.12 300510-01-ptquf 001.MOV 109 :03.63 oblique open 5.1 7.7 8.0 51.88

66

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 300510-01-ptquf 001.MOV 306 :10.20 oblique partial 5.1 6.9 7.8 46.06 300510-01-ptquf 001.MOV 426 :14.20 lateral partial 5.1 4.2 7.3 27.49 300510-01-ptquf 001.MOV 456 :15.20 oblique partial 5.1 7.2 8.3 47.98 300510-01-ptquf 001.MOV 543 :18.10 lateral partial 5.0 2.4 6.7 14.74 300510-01-ptquf 001.MOV 574 :19.13 lateral partial 5.0 4.4 7.2 28.42 300510-01-ptquf 001.MOV 605 :20.17 lateral open 5.0 7.1 8.5 47.30 300510-01-ptquf 001.MOV 628 :20.93 oblique partial 5.0 3.8 7.3 24.47 300510-01-ptquf 5.0 9.5 0.5504 0.5933 9.1 8.5 64.70 300510-02-ptquf 001.MOV 258 :08.60 lateral partial 5.4 2.6 6.7 15.06 300510-02-ptquf 001.MOV 285 :09.50 lateral partial 5.5 4.1 7.1 24.25 300510-02-ptquf 001.MOV 289 :09.63 lateral partial 5.5 4.6 7.4 27.61 300510-02-ptquf 001.MOV 374 :12.47 lateral open 5.4 7.5 9.1 46.34 300510-02-ptquf 001.MOV 405 :13.50 lateral partial 5.4 3.8 7.1 22.41 300510-02-ptquf 001.MOV 602 :20.07 lateral open 5.5 8.7 9.6 54.34 300510-02-ptquf 001.MOV 855 :28.50 oblique partial 5.4 2.8 6.4 16.51 300510-02-ptquf 001.MOV 1063 :35.43 lateral open 5.4 6.9 8.8 42.00 300510-02-ptquf 5.4 8.7 0.5504 0.5933 9.9 9.2 54.34 310510-07-ptquf 001.MOV 253 :08.43 oblique open 5.4 9.9 10.5 63.08 310510-07-ptquf 001.MOV 254 :08.47 oblique open 5.4 10.2 10.1 65.29 310510-07-ptquf 001.MOV 257 :08.57 oblique open 5.4 9.9 10.3 62.94 310510-07-ptquf 001.MOV 398 :13.27 frontal open 4.6 10.5 9.1 67.36 310510-07-ptquf 001.MOV 495 :16.50 oblique partial 5.4 3.9 7.3 23.22 310510-07-ptquf 001.MOV 497 :16.57 oblique partial 5.4 3.6 7.1 21.38 310510-07-ptquf 001.MOV 548 :18.27 oblique partial 5.4 3.1 6.4 18.15 310510-07-ptquf 001.MOV 915 :30.50 frontal open 4.5 7.4 11.1 45.65 310510-07-ptquf 001.MOV 916 :30.53 frontal open 5.4 7.4 9.8 45.79 310510-07-ptquf 001.MOV 1344 :44.80 frontal open 4.9 10.6 9.2 68.29 310510-07-ptquf 001.MOV 1345 :44.83 frontal open 4.9 10.9 9.8 70.41 310510-07-ptquf 001.MOV 1346 :44.87 frontal open 4.9 11.3 10.0 73.29 310510-07-ptquf 001.MOV 1355 :45.17 frontal open 4.4 9.7 9.7 61.39 310510-07-ptquf 001.MOV 1379 :45.97 postlat open 3.2 8.8 8.6 55.60 310510-07-ptquf 001.MOV 1437 :47.90 lateral open 5.4 9.0 9.2 56.63 310510-07-ptquf 001.MOV 1442 :48.07 lateral open 5.4 8.8 8.6 55.71 310510-07-ptquf 5.4 11.3 0.5504 0.5933 9.8 9.1 73.29 310510-08-ptquf 001.MOV 380 :12.67 oblique partial 5.1 6.1 7.8 40.15 310510-08-ptquf 001.MOV 392 :13.07 oblique open 5.1 7.2 7.0 47.72 310510-08-ptquf 001.MOV 419 :13.97 oblique open 5.1 6.0 8.3 39.09 310510-08-ptquf 001.MOV 424 :14.13 oblique partial 5.0 4.9 7.8 31.43 310510-08-ptquf 001.MOV 443 :14.77 oblique partial 5.0 4.7 7.3 30.72 310510-08-ptquf 001.MOV 811 :27.03 lateral open 5.1 10.0 10.2 68.50 310510-08-ptquf 001.MOV 820 :27.33 lateral open 5.1 8.2 9.0 55.00

67

U U X- Y- Z- CLJ C J CLJ C J file (specimen/species/sex) frame time view state mm mm mm scale scale mm mm gape ° 310510-08-ptquf 001.MOV 1029 :34.30 oblique partial 5.1 5.9 8.2 38.69 310510-08-ptquf 001.MOV 1078 :35.93 lateral partial 5.1 4.9 7.5 32.07 310510-08-ptquf 001.MOV 1127 :37.57 oblique partial 5.0 6.1 8.2 40.02 310510-08-ptquf 5.1 10.0 0.5504 0.5933 9.2 8.5 68.50 310510-09-ptquf 001.MOV

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APPENDIX B

Jaw Scaling Measurement Data. Each line represents one picture taken during dissection with a summary line per specimen. All measurements are in millimeters (mm). The first line for each specimen contains manual measurements using a caliper, while subsequent lines are digital measurements using calibrated software. The summary lines contain measurement averages excluding manual measurements. The species summary line is an average of the specimen summary lines. CLJ is length measured from lower canine to jaw joint, CUJ is length from upper canine to jaw joint, CUO is length from upper canine to the center of auditory bulla, CUE is length from upper canine to center of eye, and EO is length from center of eye to center of auditory bulla (see Figure 8). Lower jaw U U U U U (C E/CLJ) and upper jaw (C E/C J) scaling factors are calculated as C E divided by C J and CLJ, respectively. File includes Species where Mobl is Mormoops blainvillei and Ptqu is Pteronotus quadridens, Sex where m is male and f is female, and Specimen Number for that individual, pic id is the digital number assigned to the image, state is either manual for manual measurements, early before jaw joint is exposed, late where all landmarks are visible, or final which contains averaged measurements.

U U U U U U file(species/sex/specimen) pic id state CLJ C J C O C E EO C E/CLJ C E/C J Moblm170608-9 manual 11.00 10.70 11.00 5.60 5.50 0.5091 0.5234 Moblm170608-9 204 early 11.64 5.70 5.94 Moblm170608-9 205 early 11.63 5.68 5.95 Moblm170608-9 206 early 11.64 5.68 5.96 Moblm170608-9 208 early 11.64 5.68 5.96 Moblm170608-9 209 early 11.65 5.63 6.02 Moblm170608-9 314 late 11.65 10.85 11.65 5.68 5.97 0.4876 0.5235 Moblm170608-9 315 late 11.64 10.83 11.64 5.70 5.94 0.4897 0.5263 Moblm170608-9 final 11.65 10.84 11.64 5.68 5.96 0.4876 0.5239 Moblm160608-7 manual 11.30 10.90 11.10 5.70 5.50 0.5044 0.5229 Moblm160608-7 224 early 11.52 5.64 5.88 Moblm160608-7 225 early 11.52 5.66 5.86 Moblm160608-7 228 late 11.52 10.61 11.52 5.62 5.90 0.4878 0.5297 Moblm160608-7 353 late 11.46 10.71 11.46 5.56 5.90 0.4852 0.5191 Moblm160608-7 354 late 11.45 10.60 11.45 5.60 5.85 0.4891 0.5283 Moblm160608-7 355 late 11.49 10.60 11.49 5.62 5.87 0.4891 0.5302 Moblm160608-7 356 late 11.51 10.72 11.51 5.65 5.86 0.4909 0.5271 Moblm160608-7 final 11.49 10.65 11.50 5.62 5.87 0.4894 0.5279 Moblm170608-7 manual 11.30 10.30 11.00 5.50 5.80 0.4867 0.5340 Moblm170608-7 218 early 11.53 5.60 5.93 Moblm170608-7 219 early 11.52 5.60 5.92 Moblm170608-7 220 late 11.53 10.73 11.53 5.63 5.90 0.4883 0.5247 Moblm170608-7 221 late 11.56 10.78 11.56 5.65 5.82 0.4888 0.5241 Moblm170608-7 222 late 11.52 10.72 11.52 5.62 5.90 0.4878 0.5243

69

U U U U U U file(species/sex/specimen) pic id state CLJ C J C O C E EO C E/CLJ C E/C J Moblm170608-7 357 late 11.52 10.74 11.52 5.65 5.87 0.4905 0.5261 Moblm170608-7 359 late 11.57 10.70 11.57 5.65 5.92 0.4883 0.5280 Moblm170608-7 363 late 11.52 10.70 11.52 5.61 5.91 0.4870 0.5243 Moblm170608-7 final 11.54 10.73 11.53 5.63 5.90 0.4877 0.5244 Moblm170608-8 manual 11.00 10.70 11.00 5.50 6.00 0.5000 0.5140 Moblm170608-8 212 early 11.51 5.62 5.89 Moblm170608-8 213 late 11.52 10.70 11.52 5.62 5.90 0.4878 0.5252 Moblm170608-8 214 late 11.54 10.70 11.54 5.61 5.93 0.4861 0.5243 Moblm170608-8 364 late 11.58 10.83 11.58 5.70 5.88 0.4922 0.5263 Moblm170608-8 365 late 11.70 10.85 11.70 5.73 5.97 0.4897 0.5281 Moblm170608-8 366 late 11.61 10.83 11.61 5.67 5.94 0.4884 0.5235 Moblm170608-8 367 late 11.66 10.88 11.66 5.68 5.98 0.4871 0.5221 Moblm170608-8 368 late 11.66 10.87 11.66 5.69 5.97 0.4880 0.5235 Moblm170608-8 final 11.61 10.81 11.60 5.67 5.93 0.4879 0.5241 Moblm180608-10 manual 10.90 9.90 10.80 5.50 5.30 0.5046 0.5556 Moblm180608-10 230 early 11.38 5.57 5.81 Moblm180608-10 232 early 11.37 5.56 5.81 Moblm180608-10 234 late 11.38 10.60 11.38 5.57 5.81 0.4895 0.5255 Moblm180608-10 235 late 11.37 10.60 11.37 5.55 5.82 0.4881 0.5236 Moblm180608-10 236 late 11.21 10.47 11.21 5.47 5.74 0.4880 0.5224 Moblm180608-10 371 late 11.36 10.65 11.36 5.54 5.82 0.4877 0.5202 Moblm180608-10 374 late 11.36 10.55 11.36 5.52 5.84 0.4859 0.5232 Moblm180608-10 375 late 11.36 10.59 11.36 5.53 5.83 0.4868 0.5222 Moblm180608-10 final 11.34 10.58 11.35 5.54 5.81 0.4884 0.5237 Moblf160608-6 manual 11.10 10.40 11.10 5.50 5.60 0.4955 0.5288 Moblf160608-6 246 early 11.50 5.61 5.89 Moblf160608-6 247 late 11.50 10.70 11.50 5.60 5.90 0.4870 0.5234 Moblf160608-6 248 late 11.52 10.70 11.52 5.62 5.90 0.4878 0.5252 Moblf160608-6 249 late 11.49 10.71 11.48 5.60 5.88 0.4874 0.5229 Moblf160608-6 final 11.50 10.70 11.50 5.61 5.89 0.4875 0.5239 Moblf180608-6 manual 11.10 10.10 11.00 5.50 5.40 0.4955 0.5446 Moblf180608-6 251 early 11.26 5.49 5.77 Moblf180608-6 252 early 11.26 5.48 5.78 Moblf180608-6 255 late 11.30 10.53 11.30 5.52 5.78 0.4885 0.5242 Moblf180608-6 256 late 11.22 10.44 11.22 5.47 5.75 0.4875 0.5239 Moblf180608-6 257 late 11.25 10.46 11.25 5.49 5.76 0.4880 0.5249 Moblf180608-6 final 11.26 10.48 11.26 5.49 5.77 0.4877 0.5240 Moblf160608-9 manual 11.50 10.30 11.30 5.70 5.50 0.4957 0.5534 Moblf160608-9 259 early 11.55 5.64 5.91 Moblf160608-9 260 early 11.56 5.65 5.91

70

U U U U U U file(species/sex/specimen) pic id state CLJ C J C O C E EO C E/CLJ C E/C J Moblf160608-9 261 late 11.54 10.75 11.54 5.63 5.91 0.4879 0.5237 Moblf160608-9 262 late 11.55 10.76 11.55 5.64 5.91 0.4883 0.5242 Moblf160608-9 263 late 11.52 10.73 11.52 5.62 5.90 0.4878 0.5238 Moblf160608-9 final 11.54 10.75 11.54 5.64 5.91 0.4885 0.5244 Moblf170608-6 manual 11.20 10.10 11.10 5.50 5.50 0.4911 0.5446 Moblf170608-6 272 early 11.50 5.61 5.89 Moblf170608-6 274 early 11.48 5.60 5.88 Moblf170608-6 275 late 11.54 10.74 11.54 5.63 5.91 0.4879 0.5242 Moblf170608-6 276 late 11.59 10.71 11.59 5.66 5.88 0.4884 0.5285 Moblf170608-6 277 late 11.55 10.76 11.55 5.64 5.91 0.4883 0.5242 Moblf170608-6 final 11.56 10.74 11.53 5.63 5.89 0.4869 0.5242 Moblf180608-8 manual 11.20 10.20 11.10 5.60 5.60 0.5000 0.5490 Moblf180608-8 265 early 11.26 5.50 5.76 Moblf180608-8 267 late 11.28 10.50 11.28 5.50 5.78 0.4876 0.5238 Moblf180608-8 268 late 11.32 10.53 11.32 5.52 5.80 0.4876 0.5242 Moblf180608-8 269 late 11.24 10.45 11.24 5.48 5.76 0.4875 0.5244 Moblf180608-8 final 11.28 10.49 11.28 5.50 5.78 0.4876 0.5241 Mobl 0.4879 0.5245 Ptqum220608-4 manual 9.60 8.70 9.70 5.40 4.40 0.5625 0.6207 Ptqum220608-4 282 early 9.92 5.45 4.47 Ptqum220608-4 283 late 10.00 9.23 10.00 5.49 4.51 0.5490 0.5948 Ptqum220608-4 284 late 9.90 9.15 9.90 5.44 4.46 0.5495 0.5945 Ptqum220608-4 final 9.95 9.19 9.94 5.46 4.48 0.5487 0.5941 Ptqum220608-9 manual 9.60 8.60 9.60 5.30 4.30 0.5521 0.6163 Ptqum220608-9 286 early 10.13 5.56 4.57 Ptqum220608-9 287 early 10.10 5.55 4.55 Ptqum220608-9 290 late 10.06 9.34 10.06 5.54 4.52 0.5507 0.5931 Ptqum220608-9 291 late 10.14 9.44 10.14 5.58 4.56 0.5503 0.5911 Ptqum220608-9 292 late 10.07 9.32 10.07 5.52 4.55 0.5482 0.5923 Ptqum220608-9 final 10.09 9.37 10.10 5.55 4.55 0.5500 0.5925 Ptqum230608-8 manual 9.60 8.70 9.80 5.50 4.30 0.5729 0.6322 Ptqum230608-8 300 early 10.20 5.61 4.59 Ptqum230608-8 302 early 10.20 5.61 4.59 Ptqum230608-8 303 late 10.21 9.43 10.21 5.62 4.59 0.5504 0.5960 Ptqum230608-8 304 late 10.20 9.45 10.20 5.61 4.59 0.5500 0.5937 Ptqum230608-8 305 late 10.20 9.48 10.20 5.62 4.58 0.5510 0.5928 Ptqum230608-8 final 10.20 9.45 10.20 5.61 4.59 0.5502 0.5939 Ptqum230608-32 manual 10.00 9.00 10.00 5.60 4.50 0.5600 0.6222 Ptqum230608-32 294 early 10.29 5.66 4.63 Ptqum230608-32 295 early 10.28 5.65 4.63

71

U U U U U U file(species/sex/specimen) pic id state CLJ C J C O C E EO C E/CLJ C E/C J Ptqum230608-32 297 late 10.30 9.55 10.30 5.66 4.64 0.5495 0.5927 Ptqum230608-32 298 late 10.26 9.53 10.26 5.64 4.62 0.5497 0.5918 Ptqum230608-32 final 10.28 9.54 10.28 5.65 4.63 0.5499 0.5925 Ptqum230608-4 manual 9.50 8.70 9.60 5.20 4.50 0.5474 0.5977 Ptqum230608-4 309 early 9.96 5.47 4.49 Ptqum230608-4 310 early 9.95 5.47 4.48 Ptqum230608-4 311 late 9.86 9.14 9.86 5.42 4.44 0.5497 0.5930 Ptqum230608-4 312 late 9.99 9.26 9.99 5.49 4.50 0.5495 0.5929 Ptqum230608-4 final 9.93 9.20 9.94 5.46 4.48 0.5504 0.5938 Ptquf220608-8 manual 9.60 8.60 9.80 5.30 4.50 0.5521 0.6163 Ptquf220608-8 318 early 9.90 5.43 4.47 Ptquf220608-8 319 early 9.93 5.46 4.47 Ptquf220608-8 320 late 9.83 9.14 9.83 5.41 4.42 0.5504 0.5919 Ptquf220608-8 321 late 9.88 9.17 9.88 5.43 4.45 0.5496 0.5921 Ptquf220608-8 322 late 9.79 9.07 9.79 5.38 4.41 0.5495 0.5932 Ptquf220608-8 final 9.83 9.13 9.87 5.42 4.44 0.5514 0.5941 Ptquf220608-10 manual 9.70 8.80 9.80 5.20 4.60 0.5361 0.5909 Ptquf220608-10 325 early 9.77 5.37 4.40 Ptquf220608-10 327 late 9.71 8.99 9.71 5.34 4.37 0.5499 0.5940 Ptquf220608-10 328 late 9.76 9.06 9.76 5.36 4.40 0.5492 0.5916 Ptquf220608-10 final 9.74 9.03 9.75 5.36 4.39 0.5502 0.5935 Ptquf230608-2 manual 9.90 8.90 9.90 5.50 4.40 0.5556 0.6180 Ptquf230608-2 335 early 10.34 5.72 5.62 Ptquf230608-2 336 late 10.37 9.61 10.37 5.70 4.67 0.5497 0.5931 Ptquf230608-2 337 late 10.23 9.56 10.23 5.63 4.60 0.5503 0.5889 Ptquf230608-2 338 late 10.26 9.54 10.26 5.64 4.62 0.5497 0.5912 Ptquf230608-2 final 10.29 9.57 10.30 5.67 4.88 0.5514 0.5927 Ptquf230608-6 manual 9.50 8.50 9.60 5.30 4.30 0.5579 0.6235 Ptquf230608-6 330 early 9.82 9.82 5.40 4.42 Ptquf230608-6 331 early 9.86 5.42 4.44 Ptquf230608-6 332 late 9.85 9.19 9.85 5.42 4.43 0.5503 0.5898 Ptquf230608-6 333 late 9.72 9.03 9.72 5.35 4.37 0.5504 0.5925 Ptquf230608-6 334 late 9.70 9.00 9.70 5.33 4.37 0.5495 0.5922 Ptquf230608-6 final 9.76 9.07 9.78 5.38 4.40 0.5514 0.5929 Ptquf220608-4 manual 9.60 8.70 9.70 5.30 4.40 0.5521 0.6092 Ptquf220608-4 344 early 9.93 5.46 4.47 Ptquf220608-4 345 late 9.91 9.20 9.91 5.45 4.46 0.5499 0.5924 Ptquf220608-4 346 late 9.90 9.18 9.90 5.44 4.46 0.5495 0.5926 Ptquf220608-4 final 9.91 9.19 9.91 5.45 4.46 0.5502 0.5930 Ptqu 0.5504 0.5933

72

APPENDIX C

Bite Force Measurement Data. Each line represents one individual bat processed in the field. Up to five bite force meter readings bf1, bf2, bf3, bf4, and bf5 are expressed in newtons (N). The maximum bite force meter reading (max) was converted to actual bite force (MAX), also in N, by applying the bite force meter conversion factor (bf factor) (see Figure 10). Head measurements for length (head L), width (head W), and height (head H), in millimeters (mm) taken with a digital caliper are multiplied together and converted to cubic centimeters (cm3) for the head proportion (LxWxH). Normalized bite force (norm bf) is MAX divided by LxWxH expressed in N/cm3. Specimen is a field assigned identifier containing date and sequence number, species is Mobl for Mormoops blainvillei or Ptqu for Pteronotus quadridens, sex is m for male or f for female. Shaded entries are those used for statistical calculations. specimen species sex bf1 bf2 bf3 bf4 bf5 max bf factor MAX head L head W head H LxWxH norm bf 220510-44 Mobl m 2.64 1.84 2.44 1.78 3.63 3.63 0.6194 2.25 19.3 13.1 13.9 3.51 0.64 300510-08 Mobl m 1.51 2.88 3.44 2.14 3.44 0.6194 2.13 16.2 10.8 12.0 2.10 1.01 310510-02 Mobl m 3.25 2.00 3.25 0.6194 2.01 16.0 11.1 13.0 2.31 0.87 290510-04 Mobl m 3.08 1.38 1.31 3.10 3.10 0.6194 1.92 310510-03 Mobl m 3.06 2.20 1.29 1.96 1.93 3.06 0.6194 1.90 15.2 10.4 12.0 1.90 1.00 210510-10 Mobl m 3.05 2.64 2.19 1.60 3.05 0.6194 1.89 220510-10 Mobl m 2.88 1.51 2.88 0.6194 1.78 18.8 11.8 13.0 2.88 0.62 210510-02 Mobl m 1.71 1.29 2.85 2.85 0.6194 1.77 16.9 12.1 13.0 2.66 0.66 310510-06 Mobl m 2.73 2.53 2.43 2.73 0.6194 1.69 16.1 10.5 13.1 2.21 0.76 300510-05 Mobl m 1.79 2.53 2.04 1.57 1.99 2.53 0.6194 1.57 16.5 10.4 12.4 2.13 0.74 260510-02 Mobl m 0.65 2.33 2.49 2.49 0.6194 1.54 300510-07 Mobl m 2.21 2.49 2.49 0.6194 1.54 16.4 12.3 12.9 2.60 0.59 220510-43 Mobl m 2.44 2.44 0.6194 1.51 19.2 13.3 13.0 3.32 0.46 310510-05 Mobl m 2.38 2.04 0.90 2.33 2.38 0.6194 1.47 15.9 9.9 12.3 1.94 0.76 300510-09 Mobl m 2.35 2.35 0.6194 1.46 300510-04 Mobl m 2.31 1.83 1.68 1.90 2.15 2.31 0.6194 1.43 16.8 11.6 12.0 2.34 0.61 210510-08 Mobl m 0.18 0.76 0.76 0.6194 0.47 18.8 11.8 13.0 2.88 0.16 300510-03 Mobl f 2.48 3.35 2.26 2.70 3.35 0.6194 2.07 15.1 10.5 11.8 1.87 1.11 300510-06 Mobl f 1.93 2.84 2.81 2.29 2.56 2.84 0.6194 1.76 15.4 13.0 12.7 2.54 0.69 220510-11 Mobl f 0.93 2.58 2.58 0.6194 1.60 17.8 11.9 12.3 2.61 0.61 300510-10 Mobl f 2.25 2.18 2.43 2.43 0.6194 1.51 15.9 12.1 12.0 2.31 0.65 220510-08 Mobl f 1.45 1.45 0.6194 0.90 18.8 11.8 13.0 2.88 0.31 310510-04 Mobl f 1.10 1.29 1.08 1.29 0.6194 0.80 16.1 9.6 11.2 1.73 0.46 310510-01 Mobl f 1.19 1.19 0.6194 0.74 15.7 12.2 12.6 2.41 0.31 210510-11 Ptqu m 3.35 3.26 3.35 0.6194 2.07 15.8 10.0 9.3 1.47 1.41 260510-06 Ptqu m 3.06 3.26 3.20 3.13 3.20 3.26 0.6194 2.02 260510-23 Ptqu m 2.09 2.11 3.09 2.26 2.56 3.09 0.6194 1.91 290510-11 Ptqu m 2.10 1.94 2.98 1.48 1.90 2.98 0.6194 1.85 220510-00 Ptqu m 2.86 2.86 0.6194 1.77 260510-20 Ptqu m 2.55 2.71 2.83 2.64 2.68 2.83 0.6194 1.75 290510-02 Ptqu m 2.21 2.51 1.91 2.51 0.6194 1.55

73 specimen species sex bf1 bf2 bf3 bf4 bf5 max bf factor MAX head L head W head H LxWxH norm bf 260510-21 Ptqu m 2.29 2.03 2.44 2.21 2.44 0.6194 1.51 260510-40 Ptqu m 2.39 2.31 1.50 2.23 2.39 0.6194 1.48 260510-24 Ptqu m 2.15 2.21 2.25 1.85 2.23 2.25 0.6194 1.39 13.7 8.6 10.7 1.26 1.11 290510-00 Ptqu m 2.20 1.89 2.20 0.6194 1.36 260510-22 Ptqu m 2.16 2.16 0.6194 1.34 210510-07 Ptqu m 1.85 1.85 1.85 0.6194 1.15 15.7 10.3 10.1 1.63 0.70 260510-25 Ptqu m 1.50 1.01 1.40 1.50 0.6194 0.93 16.3 10.9 9.6 1.71 0.54 270510-24 Ptqu f 1.49 1.99 2.34 2.96 2.60 2.96 0.6194 1.83 290510-41 Ptqu f 1.25 0.95 0.93 2.33 2.33 2.33 0.6194 1.44 300510-01 Ptqu f 2.31 2.10 2.04 2.15 1.91 2.31 0.6194 1.43 16.1 9.6 10.2 1.58 0.91 310510-09 Ptqu f 1.38 2.21 1.81 1.21 2.21 0.6194 1.37 15.5 8.3 9.2 1.18 1.16 310510-07 Ptqu f 2.06 2.09 2.09 0.6194 1.29 15.8 7.7 9.2 1.12 1.16 270510-45 Ptqu f 1.48 2.08 2.08 0.6194 1.29 270510-07 Ptqu f 0.96 2.00 1.53 1.13 2.03 2.03 0.6194 1.26 220510-00 Ptqu f 2.03 2.03 0.6194 1.26 300510-02 Ptqu f 1.86 1.36 1.84 1.36 1.86 0.6194 1.15 15.5 9.9 10.2 1.57 0.74 310510-08 Ptqu f 1.45 1.86 1.83 1.86 0.6194 1.15 16.0 9.2 9.5 1.40 0.79

74

APPENDIX D

Jaw Musculature Weight Measurement Data. Each line represents one set of muscle weight measurements, five sets per specimen, followed by an average line per specimen. Left, right, and both sides of muscles were weighed, and the two sides were summed for validation. All weights are in in milligrams recorded to the nearest 0.1mg. The museum specimen number (id) is an identifier containing date and sequence number, species is Mobl for Mormoops blainvillei or Ptqu for Pteronotus quadridens, sex is m for male or f for female. Bolded entries are those used for muscle architecture analysis (see Table 5).

temporalis masseter m. pterygoid digastric l. pterygoid left right both sum left right both sum left right both sum left right both sum left right both sum id species sex trial mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg mg 180608-4 Mobl m 1 45.5 47.4 93.1 92.9 7.8 8.1 16.0 15.9 2.5 2.2 4.7 4.7 5.4 5.8 11.1 11.2 1.3 1.2 2.5 2.5 2 46.0 47.2 92.7 93.2 7.7 8.2 15.9 15.9 2.5 2.4 4.8 4.9 5.3 5.3 10.6 10.6 1.3 1.2 2.5 2.5 3 46.2 47.3 93.4 93.5 7.8 8.3 16.1 16.1 2.4 2.3 4.7 4.7 5.4 5.6 10.9 11.0 1.2 1.1 2.3 2.3 4 47.2 47.0 94.7 94.2 7.7 8.5 16.1 16.2 2.5 2.2 4.6 4.7 5.8 5.7 11.3 11.5 1.2 1.1 2.3 2.3 5 45.3 47.3 92.3 92.6 8.0 8.2 16.1 16.2 2.5 2.2 4.6 4.7 5.4 5.8 11.1 11.2 1.2 1.0 2.3 2.2 180608-4 Mobl m avg 46.0 47.2 93.2 93.3 7.8 8.3 16.0 16.1 2.5 2.3 4.7 4.7 5.5 5.6 11.0 11.1 1.2 1.1 2.4 2.4 230510-42 Mobl f 1 37.6 40.6 78.0 78.2 6.8 6.8 13.6 13.6 2.4 2.8 5.2 5.2 4.3 4.6 8.8 8.9 2.0 1.6 3.6 3.6 2 38.3 42.6 80.9 80.9 6.9 6.7 13.8 13.6 2.3 2.5 4.9 4.8 4.6 4.5 8.9 9.1 2.0 1.6 3.5 3.6 3 38.6 40.4 78.9 79.0 7.2 7.1 14.3 14.3 2.4 2.5 4.9 4.9 4.5 4.8 9.2 9.3 2.1 1.5 3.6 3.6 4 39.2 40.1 79.0 79.3 7.3 7.1 14.2 14.4 2.2 2.8 4.9 5.0 4.6 4.5 8.9 9.1 1.8 1.6 3.3 3.4 5 38.3 40.2 79.0 78.5 6.9 7.0 13.8 13.9 2.3 2.6 4.9 4.9 4.7 4.4 9.0 9.1 2.0 1.6 3.6 3.6 230510-42 Mobl f avg 38.4 40.8 79.2 79.2 7.0 6.9 13.9 14.0 2.3 2.6 5.0 5.0 4.5 4.6 9.0 9.1 2.0 1.6 3.5 3.6 260510-22 Ptqu m 1 33.6 33.0 66.6 66.6 6.7 6.3 12.8 13.0 1.9 1.5 3.4 3.4 4.2 3.7 7.8 7.9 1.4 1.1 2.3 2.5 2 34.2 35.3 69.3 69.5 6.7 6.3 12.9 13.0 1.7 1.6 3.2 3.3 4.5 3.8 8.2 8.3 1.4 0.9 2.2 2.3 3 33.5 34.8 68.3 68.3 6.8 6.1 12.7 12.9 1.8 1.5 3.3 3.3 4.4 3.7 8.0 8.1 1.2 0.9 2.1 2.1 4 34.1 33.7 67.4 67.8 6.9 6.4 13.3 13.3 1.8 1.6 3.4 3.4 4.5 3.8 8.2 8.3 1.3 0.9 2.1 2.2 5 34.0 31.9 65.4 65.9 6.9 6.3 13.1 13.2 1.8 1.5 3.3 3.3 4.0 3.8 7.7 7.8 1.4 1.0 2.3 2.4 260510-22 Ptqu m avg 33.9 33.7 67.4 67.6 6.8 6.3 13.0 13.1 1.8 1.5 3.3 3.3 4.3 3.8 8.0 8.1 1.3 1.0 2.2 2.3 230608-10 Ptqu f 1 36.1 38.3 74.4 74.4 6.5 6.2 12.7 12.7 1.4 0.9 2.4 2.3 5.0 4.4 9.4 9.4 1.2 0.7 1.9 1.9 2 36.1 38.7 74.7 74.8 6.8 6.5 13.3 13.3 1.2 1.1 2.2 2.3 4.7 4.9 9.5 9.6 1.2 0.9 2.0 2.1 3 36.2 39.1 75.4 75.3 6.9 6.8 13.6 13.7 1.2 1.1 2.3 2.3 4.8 4.8 9.4 9.6 1.2 0.8 1.9 2.0 4 36.5 38.5 74.4 75.0 6.7 6.5 13.3 13.2 1.1 1.0 2.1 2.1 4.6 4.6 9.0 9.2 1.1 0.8 1.9 1.9 5 36.1 38.1 74.1 74.2 6.9 6.5 13.2 13.4 1.2 1.0 2.1 2.2 4.5 4.7 9.1 9.2 1.0 0.9 1.9 1.9 230608-10 Ptqu f avg 36.2 38.5 74.6 74.7 6.8 6.5 13.2 13.3 1.2 1.0 2.2 2.2 4.7 4.7 9.3 9.4 1.1 0.8 1.9 2.0

75

APPENDIX E

Jaw Musculature Fiber Length Measurement Data. There are two lines per museum specimen for each muscle, one for each side where L is left and R is right. Ten length measurements were taken (L1-L10) expressed in millimeters. The maximums were computed for individual muscles (max) and muscle pairs (avg max). The museum specimen number (id) is an identifier containing date and sequence number, species is Mobl for Mormoops blainvillei or Ptqu for Pteronotus quadridens, sex is m for male or f for female. Bolded entries (avg max) are those used for muscle architecture analysis (see Table 5). avg id species sex muscle side L1 L2 L3 L4 L5 L6 L7 L8 L9 L10 max max 180608-4 Mobl m temporalis L 3.02 2.92 2.90 2.86 2.86 2.98 3.04 2.93 3.01 2.91 3.04 3.20 R 3.02 2.84 2.71 2.81 3.27 3.28 3.23 3.35 3.20 3.32 3.35 230510-42 Mobl f L 3.12 3.19 3.33 3.53 3.50 3.49 3.59 3.35 3.22 3.64 3.64 3.58 R 3.00 3.40 3.36 3.01 2.91 2.86 3.25 3.21 3.51 3.11 3.51 260510-22 Ptqu m L 3.58 3.55 3.57 3.57 3.48 3.50 3.51 3.45 3.58 3.55 3.58 3.69 R 3.70 3.79 3.50 3.49 3.62 3.59 3.77 3.50 3.73 3.80 3.80 230608-10 Ptqu f L 2.75 2.77 2.71 2.89 2.96 3.01 2.96 2.86 3.00 2.83 3.01 3.05 R 2.95 3.07 2.71 2.76 3.09 2.87 3.06 3.08 3.06 2.98 3.09 180608-4 Mobl m masseter L 2.67 2.60 2.62 2.64 2.61 2.62 2.44 2.72 2.50 2.70 2.72 2.86 R 2.99 2.95 2.57 2.54 2.64 2.64 2.83 2.85 2.45 2.52 2.99 230510-42 Mobl f L 2.95 3.16 3.16 3.15 2.99 2.85 2.95 2.76 3.04 2.90 3.16 3.12 R 2.96 3.07 2.75 2.73 2.63 2.67 2.79 2.69 2.64 2.69 3.07 260510-22 Ptqu m L 2.84 3.04 3.12 3.14 3.14 2.98 2.72 2.99 3.13 3.16 3.16 3.16 R 3.03 3.07 3.06 2.98 2.89 3.15 2.90 2.92 3.00 2.98 3.15 230608-10 Ptqu f L 2.22 2.21 2.20 2.22 2.21 2.24 2.23 2.22 2.25 2.24 2.25 2.21 R 2.14 2.11 1.92 1.96 2.04 1.96 2.15 2.12 2.16 2.12 2.16 180608-4 Mobl m m.pterygoid L 1.85 1.87 1.84 1.81 1.78 1.81 1.72 1.84 1.78 1.78 1.87 1.83 R 1.73 1.70 1.66 1.60 1.78 1.59 1.61 1.43 1.57 1.77 1.78 230510-42 Mobl f L 1.72 1.60 1.59 1.48 1.69 1.69 1.78 1.82 1.86 1.85 1.86 1.89 R 1.92 1.87 1.73 1.83 1.77 1.86 1.86 1.83 1.80 1.83 1.92 260510-22 Ptqu m L 1.61 1.62 1.56 1.55 1.55 1.55 1.60 1.66 1.62 1.62 1.66 1.67 R 1.63 1.47 1.66 1.62 1.59 1.67 1.56 1.49 1.40 1.48 1.67 230608-10 Ptqu f L 1.15 1.20 1.16 1.29 1.28 1.26 1.29 1.25 1.23 1.22 1.29 1.29 R 1.04 1.17 1.17 1.13 1.29 1.21 1.23 1.24 1.15 1.16 1.29 180608-4 Mobl m digastric L 5.55 5.66 5.58 6.14 6.16 6.22 6.22 6.44 6.45 6.36 6.45 6.81 R 6.55 6.51 6.54 7.06 6.89 7.07 7.14 7.14 7.15 7.16 7.16 230510-42 Mobl f L 7.02 7.01 6.96 6.52 6.63 6.60 6.67 6.88 6.90 7.01 7.02 7.21 R 7.21 7.30 7.39 7.38 7.25 7.19 7.40 7.24 7.22 7.32 7.40 260510-22 Ptqu m L 5.93 5.82 5.69 5.66 5.60 5.52 5.55 5.66 5.82 5.90 5.93 5.50 R 4.95 5.02 5.07 5.05 5.01 4.64 4.95 4.78 4.86 4.75 5.07 230608-10 Ptqu f L 6.83 6.82 6.81 7.03 6.99 7.00 7.01 6.98 6.99 6.93 7.03 6.62 R 6.20 6.14 6.10 6.19 6.19 6.18 6.09 6.02 5.95 6.12 6.20

76

avg id species sex muscle side L1 L2 L3 L4 L5 L6 L7 L8 L9 L10 max max 180608-4 Mobl m l.pterygoid L 1.47 1.52 1.47 1.49 1.46 1.58 1.54 1.53 1.46 1.72 1.72 1.62 R 1.40 1.41 1.47 1.46 1.41 1.51 1.44 1.51 1.51 1.51 1.51 230510-42 Mobl f L 2.08 2.24 2.58 2.48 2.61 2.31 2.32 2.37 2.20 2.08 2.61 2.26 R 1.79 1.69 1.83 1.81 1.79 1.88 1.77 1.88 1.90 1.84 1.90 260510-22 Ptqu m L 1.74 1.72 1.69 1.67 1.73 1.70 1.76 1.68 1.75 1.76 1.76 1.76 R 1.72 1.70 1.76 1.56 1.55 1.64 1.71 1.61 1.61 1.69 1.76 230608-10 Ptqu f L 1.41 1.05 1.13 1.26 1.07 1.15 1.17 1.26 1.42 1.39 1.42 1.41 R 1.24 1.38 1.25 1.19 1.21 1.39 1.21 1.21 1.29 1.29 1.39

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