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1981

Morphological variation of grizzly bear skulls from Yellowstone National Park

Harrie W. Sherwood

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Sherwood, Harrie W., "Morphological variation of grizzly bear skulls from Yellowstone National Park" (1981). Graduate Student Theses, Dissertations, & Professional Papers. 7381.

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D A T E i _ l M l _

MORPHOLOGICAL VARIATION OF GRIZZLY BEAR SKULLS FROM YELLOWSTONE NATIONAL PARK

By
Marrie W. Sherwood
B.A., U niversity of Colorado, 1974

Presented in p a rtia l fu lfillm e n t of the requirements fo r the degree of
Master of Science
UNIVERSITY OF MONTANA
1981

Approved Chairman,^BoaN oNExamfners D ^n, Graduate School

I s.- a 5-^ I

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UMI Number: EP38182

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ABSTRACT

  • Sherwood, Harrie W., M.S., Fall 1981
  • W ild life Biology

Morphological Variation of G rizzly Bear Skulls from Yellowstone National Park / O

  • /
  • /

’ I

D irector: P h ilip L. Wright ( j .A
Cranial characters of tooth eruption of the permanent dentition tooth wear, cranial sutures, and 24 cranial measurements were examined on 68 (44 males, 24 females) g riz z ly bear (Ursus arctos h o r r ib ilis ) skulls from Yellowstone National Park to investigate the cranial variations of g riz z lie s with respect to age, sexual, and individual variation and to examine the u t ilit y of using cranial characters of indicators of age and sex. Twenty four of the bear skulls were of known age whose ages ranged from cubs to 11 years old. The eruption of the g riz z ly bear dentition began sometime before the f if t h month of lif e and was completed by the seventeenth month, except fo r the canines and M3 which were

  • erupted by theth ird year of lif e .
  • Tooth wear increased with age

but was poorly correlated with agedue to considerable variatio n between individual bears of the same age class. The sequence of cranial suture closure occurred at approximately the same age fo r a ll g riz z lie s .

For taxonomic and s ta tis tic a l purposes of skull size, bear skulls were c la ssifie d into six age classes on the basis of tooth eruption and wear, bone fusion, changes in cranial structure, and chronological age. Female g riz z ly skulls attained adult size e a rlie r and were s ig n ific a n tly smaller than adult male g riz zly sku lls. Maximum female cranial size was reached between the ages of 6 to 11 years. Male g riz z lie s did not grow appreciably in length beyond 12 years of age, but continued to grow in breadth and height beyond 12 years of age. Age estimation by m ultiple regression, discrim inant analysis, and cranial sutures gave re lia b le results.

As age increased there was an increased difference in cranial size between male and female g riz z lie s . Skulls of adult male g riz z lie s averaged 13 percent larger than adult female skulls. Sex-related differences in size of the sa g itta l crest, mastoid breadth, zygomatic breadth, as well as discrim inant analysis were useful in determining the sex of unknown adult g riz z ly sku lls. Young bears of the same age-sex class were more variable in skull size than adult bears of the same age-sex class. Individual differences in the presence or absence of the a n te rio r premolars was o f l i t t l e taxonomic consequence.

ACKNOWLEDGEMENTS

Financial support fo r the study was provided by the Montana
Cooperative W ild life Research Unit (Montana Department of Fish, W ild life and Parks, U niversity of Montana, United States Fish and W ild life Service, and the W ild life Management In s titu te , cooperating).
I express my thanks and indebtedness to my committee members.
Dr. P h ilip W right, my major advisor, and Drs. Charles Jonkel and Bart O'Gara who s e lfle s s ly gave th e ir guidance in preparation and c ritic a l review of the manuscript.
I thank Dr. John Craighead, who generously loaned the use of his bear skulls with data fo r purposes of study.
I thank Jay Sumner fo r co lla tin g the data on the Craighead bear skulls and fo r the age estimates he provided on some of those specimens. I express my gratitude to Ken Greer (Montana Dept. Fish, W ild life and Parks) fo r use of state-owned bear skulls and his helpful suggestions and cooperation.
I appreciate the comments and suggestions o f my fellow graduate students, especially Steve A lbert fo r his s ta tis tic a l and computer assistance. Bob H o llis te r, Ph.D. candidate in Mathematics, provided assistance in s ta tis tic a l treatments.
I express my deepest appreciation to my w ife, Kathy, fo r her sharing and patient support.

TABLE OF CONTENTS
Page

i i

ABSTRACT.......................................................................................................................

ACKNOWLEDGEMENTS.......................................................................................................

LIST OF TABLES...........................................................................................................

LIST OF FIGURES.........................................................................................................

i i i v1 v ii

LIST OF PLATES........................................................................................................... v iii

CHAPTER
I.

INTRODUCTION..................................................................................................

15

I I . MATERIALS AND METHODS................................................................................

Age V ariation............................................................................................

Secondary Sexual V ariation..................................................................

Individual V ariation..............................................................................

5
12 13

I I I .

RESULTS.............................................................................................................

14

Age V ariation............................................................................................

Permanent D entition............................................................................

Tooth Wear...............................................................................................

Cranial Sutures....................................................................................

Cranial Measurements..........................................

Characteristics of Growth................................................................

Estimation of Age.................................................................................

Secondary Sexual V a riatio n..................................................................

Cranial Measurements..........................................................................

Sexual Dimorphism................................................................................

Individual V ariation..............................................................................

Permanent D e ntitio n.............................................................................

Cranial Measurement.............................................................................

14 14 16 21 23 26 34 49 49 49 54 54 54

IV. DISCUSSION.......................................................................................................

57

Age V ariation.............................................................................................

Permanent D entition.............................................................................

Tooth Wear...............................................................................................

Cranial Sutures.....................................................................................

Cranial Measurements...........................................................................

Estimation of Age.................................................................................

57 58 58 59 60 61

iV
Page

Secondary Sexual V ariation..................................................................

Sexual Dimorphism................................................................................

Individual V ariation..............................................................................

Permanent D entition............................................................................

Cranial Measurement............................................................................

Taxonomy......................................................................................................

66 66

67 67

68

70

LITERATURE CITED......................................................................................................

APPENDICES
74
A. Description of tooth wear in a series of known andassigned-age

bears............................................................................................................

77 79 81
B. Description of suture closure in a series of known andassigned-

age bears....................................................................................................

  • C. Age variation of cranial measurements of g riz z ly bears
  • from

Yellowstone National Park....................................................................

D. Age group mean values of cranial measurement fo rsubstitution

  • in to discrim inant or regression equations....................................
  • 91

  • 93
  • E. Record of g riz z ly bears used in the study.........................................

LIST OF TABLES
Page

Table
1. Age d is trib u tio n of the known age and assigned age bear

sample............................................................................................................

6

2. The six age classes used in the study with delineation by

  • morphological characteristics and chronological age..................
  • 25

3. Maximum growth c la ssifica tio n of cranial measurement by age

class and sex.............................................................................................

27
4. Age variation in 5 cranial characters of g rizzly bears from

Yellowstone National Park................................................................. 29,30,3

5. Standardized discrim inant function coefficients of male and

female age groups.....................................................................................

38 39 39 43
6. Unstandardized discrim inant function coefficients and th e ir group means fo r males and female age groups

....................

6A. Discriminant functions evaluated at group means (group

centroids)...................................................................................................

7. Comparison of age estimates on seven unknown age g rizzly

bears from Yellowstone National Park,..............................................

8. Secondary sexual variation in 24 cranial measurements in adult (age class V and VI) g rizzly bears from Yellowstone

National Park.............................................................................................

50
9. Standardized and unstandardized discrim inant function coef-

fic ie n ts of sex on adult g riz z lie s ....................................................

53

  • 53
  • 9A. Discriminant functions evaluated at group means...............................

V I

LIST OF FIGURES
Page

Figure
1. 2.

Measurements of g rizzly bear sk u lls..................................................

11 15 18 22
F irs t and second year eruption sequence of the permanent

d e n titio n .................................................................................................

3. 4. 5.
Sequence of tooth wear in g rizzly bears from Yellowstone

Park...........................................................................................................

Sequence of suture closure in g rizzly bears from Yellowstone

Park...........................................................................................................

Representative growth curves by age class of 5 cranial characters of g rizzly bears from Yellowstone National

Park...........................................................................................................

32 41 42 52

6.

7.

Discriminant p lot of male age groups................................................

Discriminant plot of female age groups............................................

Sexual dimorphism of g riz zly bear measurements.............................

8.

vn

L IST OF PLATES

Page

Plates

  • 1.
  • Progressive tooth wear d e tails of the upper and lower jaws

of 2 year, 5 year, 7 year, 10 year, 15 year, and 18 year

  • old g r iz z ly bears ...............................................................................................
  • 20

46
2. Comparative cranial size and suture closure of a 2 year,
5 year, 9 year, and 18 year old male g riz z ly bear ........................

3. Comparative cranial size and suture closure of a 2 year,
5 year, 9 year, and 18 year old female g riz z ly bear ...................
48

  • 58
  • 4. Individual variations in 4 old adult male g riz z ly skulls ............

V I I I

CHAPTER L
INTRODUCTION
As a prelim inary to the taxonomic study of a species, understanding the morphological variation w ithin that species cle arly is important. "Morphological variation w ithin an intrabreeding population of brown bears (Ursus arctos L.) is of three major types: differences between sexes, changes with age and individual variatio n. . . . Much of the past confusion obscuring the taxonomic status of th is species has resulted from attempts to compare specimens without adequate consideration fo r these variables." (Rausch 1963:33).
The great p ro c liv ity of brown bears to vary in size was recognized early by von Middendorf (1851). He concluded that the shape of the skull p ro file (vaulting of the p ro file ) was a mixture of age and individual variation. He understood that the v a ria b ility in cranial characteristics demonstrated that the Palaearctic and Nearctic forms of brown bears were conspecific.
The publications of Merriam (1896, 1914, 1918) and his "new species" cle a rly indicate his lack of understanding and consideration fo r v a ria b ility when he concluded that 86 species of brown bears existed in North America alone. Most species were id e n tifie d on the basis of few, sometimes a single, specimen. His conclusions, unfortunately, seriously hampered and influenced the work of contemporary taxonomists.
Rausch (1953, 1963) has greatly improved the understanding of the systematics of the brown and g riz z ly bears of North America. He demonstrated the significance of individual variation in cranial characteristics of bear skulls from the Alaska region and disclosed the erroneous assumptions upon which Merriam derived his species cla s s ific a tio n (Rausch 1953). He concluded that a ll North American g rizzlie s and brown bears belong in one highly variable species, Ursus arctos and recognized fiv e North American subspecies (lU.^. h o rrib il is Ord; IJ.a^. middendorffi Merriam; JJ.^. gyas Merriam; jJ.ja. richardsonii Swainson; and ]J.a^. californicus Merriam).
Rausch (1963) la te r studied a long series of local samples with regard to condylobasal length. He demonstrated that geographic variation was essentially c lin a l, decreasing in size north, east and south from a maximum on the Alaska Peninsula and Kodiak and Afognak Islands. As a re su lt, he diffe re n tia te d the North American brown and g rizz ly bears to two forms, the very large coastal and island form in Alaska and B ritis h Columbia (U.^. middendorffi Merriam) and the smaller form occupying the remainder of the geographic range of North America (]J.£. h o rrib il is Ord).
Kurtéh (1973) demonstrated that, with regard to width/length allom etry, there are two groups of bears possessing d iffe re n t growth patterns; the r e lic t, broad-skulled growth form that occupied the P acific coast of Beringia that now persists on Kodiak and Afognak Islands; and the narrow-skulled growth pattern of the form that inhabited inner and northern Beringia. He concluded that i t was possible to distinguish between only three subspecies of brown-grizzly bears now liv in g in the Nearctic region. The three subspecies he designated were: 1) Ursus arctos middendorffi Merriam 1896. — Kodiak and Afognak Islands, larger than the g riz z ly , on an average broader-skulled than mainland brown bears; 2) Ursus arctos d a lli Merriam 1896. — south coast of Alaska, west coast of B ritis h Columbia, larger than g riz z ly , on an average more narrow-skulled than U.^. m iddendorffi; and 3) Ursus arctos h o rrib ilis Ord 1815. -- a ll of Nearctic range except areas mentioned above, smaller than U.a^. middendorffi and U.a d a lli.
The problem of North American g rizzly bear taxonomy is further complicated by Hall (1981). In his la te s t edition of The Mammals of North America, he disregards the taxonomic conclusions of Rausch (1953, 1963), Kurtén (1973), Erdbrink (1953) and others by lis tin g 90 or so species-subspecies names of brown-grizzly bears; mostly those described by Merriam (1914, 1918). Thus disagreement among experts persists regarding the specific-subspecific status of the brown-grizzly bear complex of North America.
Contemporary taxonomy is characterized by a synthesis of morphological d e fin itio n combined with biological d e fin itio n , which takes ecological, geographical, genetic and other factors into consideration (Mayr, 1953). "The population, represented by an adequate sample, the series of the museum worker, has become the basic taxonomic unit- (Mayr 1953:13). With s ta tis tic s and computers, quantitative analyses of variation can be measured and compared in a standardized, precise, and meaningful way. Unfortunately, th is type of treatment on the systematics of g riz z ly bears has been lacking and thereby inhibited the advancement of g riz zly bear taxonomy. Manning's (1971) work on the geographic variation of polar bears (Ursus maritimus Phipps) provides an excellent example of how these techniques can be used to th e ir greatest benefit fo r taxonomic purposes.
G rizzly bear studies in Yellowstone National Park by Drs.
John and Frank Craighead from 1959 to 1967 (Craighead et a l. 1974) assembled information on the biology, ecology and population dynamics of the Yellowstone g riz z ly . As a part of th e ir studies, a unique series of g riz z ly bear skulls was collected with complete data on sex, age, and other biological inform ation.
Such a valuable series afforded an excellent opportunity to examine the morphological variation of g rizzly bear skulls w ithin a local, intrabreeding bear population. Additional g rizzly skulls from Yellowstone were loaned by Ken Greer of the Bozeman W ild life Laboratory (Montana Department of Fish, W ild life , and Parks).
The goal of my research was to determine the morphological characteristics of the central Rocky Mountain g riz z lie s , and to thereby provide baseline data on bears of th is area, in order that other studies could follow , and so that future comparisons could be made with other more northern g riz zly populations.
My specific objectives were to:
1) examine the cranial variation of g riz z ly bear skulls with respect to age, sex, and individual va ria tio n ;
2) describe the morphometric characteristics of the Yellowstone bear population; and
3) examine the u t ilit y of using cranial characters as indicators of age and sex.
CHAPTER LI
MATERIALS AND METHODS
Sixty eight g riz z ly skulls (44 males, 24 females) were

  • measured.
  • A ll of
  • the skulls were obtained as part of an extensive

  • in Yellowstone Park from 1959 to1971
  • population study of g riz z ly bears

(Craighead et a l. 1974). Ken Greer, Montana Department Fish, W ild life , and Parks, loanedseven skulls (4 males, 3 females) that were used to estimate age and sex.

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    Volume 1, Issue 1 A HEADS UP ON CRANIOSYNOSTOSIS Andrew Reisner, M.D., William R. Boydston, M.D., Ph.D., Barun Brahma, M.D., Joshua Chern, M.D., Ph.D., David Wrubel, M.D. Craniosynostosis, an early closure of the growth plates of the skull, results in a skull deformity and may result in neurologic compromise. Craniosynostosis is surprisingly common, occurring in one in 2,100 children. It may occur as an isolated abnormality, as a part of a syndrome or secondary to a systemic disorder. Typically, premature closure of a suture results in a characteristic cranial deformity easily recognized by a trained observer. it is usually the misshapen head that brings the child to receive medical attention and mandates treatment. This issue of Neuro Update will present the diagnostic features of the common types of craniosynostosis to facilitate recognition by the primary care provider. The distinguishing features of other conditions commonly confused with craniosynostosis, such as plagiocephaly, benign subdural hygromas of infancy and microcephaly, will be discussed. Treatment options for craniosynostosis will be reserved for a later issue of Neuro Update. Types of Craniosynostosis Sagittal synostosis The sagittal suture runs from the anterior to the posterior fontanella (Figure 1). Like the other sutures, it allows the cranial bones to overlap during birth, facilitating delivery. Subsequently, this suture is the separation between the parietal bones, allowing lateral growth of the midportion of the skull. Early closure of the sagittal suture restricts lateral growth and creates a head that is narrow from ear to ear. However, a single suture synostosis causes deformity of the entire calvarium.
  • Human Anatomy and Physiology- I BIOL2401- Lab Practical 2 Terminology

    Human Anatomy and Physiology- I BIOL2401- Lab Practical 2 Terminology

    Human Anatomy and Physiology- I BIOL2401- Lab Practical 2 Terminology Lab 12: Structure & Classification of Bone Microscopic Structure Sectioned Long Bone Osteon (Haversian) System epiphysis periosteum central (Haversian) canal diaphysis Sharpey's fibers perforating (Volkmann’s) canal epiphyseal line (disk or plate) medullary cavity osteocyte compact (dense) bone red marrow lacuna spongy (cancellous) bone yellow marrow canaliculi hyaline (articulating) cartilage endosteum trabeculae Lab 13: Organization of the Skeleton Axial skeleton vs. Appendicular skeleton Differentiate between left and right Lab 14: The Skull Cranium Frontal bone Temporal bone (2) Sphenoid bone frontal sinus squamosal suture sella turcica supraorbital foramen external acoustic meatus sphenoid sinus internal acoustic meatus optic canal mastoid process lateral pterygoid plate Parietal bone (2) styloid process medial pterygoid plate sagittal suture carotid canal coronal suture jugular foramen Occipital bone lambdoidal suture Ethmoid bone foramen magnum crista galli occipital condyles cribriform plate perpendicular plate sutural bone ethmoid sinus nasal concha Facial bones Maxillary bones Inferior nasal concha 1 maxillary sinus Mandible palantine process body alveolar fossa* ramus mandibular condyle Palatine bone coronoid process Zygomatic bone mental foramen alveolar fossa* zygomatic arch Lacrimal bone Hyoid Nasal bone Vomer bone Lab 15: Vertebral Column & Thoracic Cage Parts of vertebrae Costal (rib) (12 pairs) body types spinous process true (vertebrosternal) 7 pair
  • Lamar Institute of Technology DHYG 1301 Course Syllabus

    Lamar Institute of Technology DHYG 1301 Course Syllabus

    Lamar Institute of Technology DHYG 1301 Course Syllabus SUMMER II Taught by: Kristina M. Mendoza, RDH, DMD [email protected] MPC 211 (409) 839-2914 Orofacial Anatomy, Histology and Embryology (DHYG 1301) Summer II TABLE OF CONTENTS SCHEDULE ………………………………………………………………………….............2-4 COURSE DESCRIPTION …………………………………………………………………...6 COURSE GOALS ……………………………………………...………………………....…6 CREDIT HOURS………………………………………………………………………..…...7 CLASS MEETING TIME…………………………………………………………….….......7 INSTRUCTORS………………………………………………………………………...…....7 COURSE POLICIES …………………………………………………………………...........7-8 TEACHING METHODS…….……………………………………………………….…........9 REQUIRED TEXT……………………………………………………………………...........9 REFERENCES..………………………………………………………………………..….....9 COURSE REQUIREMENTS….…………………………………………………………......9 EVALUATION CRITERIA..……………………………………………………………...…9 GRADE SCALE..……………………………………………………………………….……9 CONTENT OUTLINE……………………………………………………………...............10-13 LEARNER OBJECTIVES………………………………………………………….............14-23 APPENDIX.…………………………………………………………………………………24 PARTICIPATION GRADE……………………………………………………………………………..…..25-26 STUDY GROUP….………………………………………………………………………………………....27 LABORATORY ASSIGNMENTS…………………………………………..............................28-51 GRADE COMPUTATION SHEET……………………………………………………….....…52 1 Orofacial Anatomy, Histology and Embryology (DHYG 1301) Summer II DHYG 1301: Orofacial Anatomy, Histology and Embryology Lecture Schedule: Monday - Thursday, 9:00 – 10:30 - Room MPC 112 Lab Schedule: MW and TR, 11:00 - 1:55 - Room MPC 127 Date Lecture Assignment Lab Assignment
  • Investigating Health in Medieval Uppsala an Osteological Study of Skulls from the Anatomical Collection of Museum Gustavianum

    Investigating Health in Medieval Uppsala an Osteological Study of Skulls from the Anatomical Collection of Museum Gustavianum

    Department of Archaeology and Ancient History Investigating health in medieval Uppsala An osteological study of skulls from the anatomical collection of Museum Gustavianum Varvara Papamargariti MA thesis 30 credits in Archaeology Spring term 2019 Supervisor: Michael Lindblom Co-supervisor: Sabine Sten English Park Campus Papamargariti, V. 2019. Investigating health in medieval Uppsala. An osteological analysis of skulls from the anatomical collection of Museum Gustavianum Papamargariti, V. 2019. Undersökning om hälsan i medeltida Uppsala. En osteologisk analys av kranier från den anatomiska samlingen av Museum Gustavianum Denna studie behandlar hälsan i det medeltida Uppsala genom osteologiska analyser på 32 skallar från Gustavianums anatomiska samling. Skallarna i fråga är del av en större grupp med mänskliga benrester som hittades vid Östra Ågatan år 1909. Gruppen består enbart av skallar; det finns ingen information om var de postkananiala skeletten finns. Studiens syfte är att undersöka sjukdom och hälsa i gruppen för att således kunna utöka informationen som finns om den medeltida hälsan i Uppsala. Den osteologiska analysen kompletteras med en diskussion om skallanas fyndplats i en möjlig relation till den närliggande medeltida Vårfrukyrkans kyrkogård. Benprover från fyra av skallarna i studien har använts för C-14 analys för att kunna fastställa den kronologiska kontexten då det inte finns någon ytterligare information om utgrävningen. C-14 analysen bekräftade den medeltida hypotesen. Den bekräftade tidsbestämmelsen tillsammans med den begränsade informationen om utgrävningsplatsen vid Östra Ågatan 37 möjliggör hypotesen att skallarna kan vara del av Vårfrukyrkans kyrkogård. Hälsoutredningen har baserats på kön och åldersbedömning likväl som på identifiering av de patologiska tillstånden som benen och tänderna befinner sig i.
  • Neural Manipulation 1 Preparation 1

    Neural Manipulation 1 Preparation 1

    Neural Manipulation 1 Preparation 1. Review this video 2. Read: Trauma: An Osteopathic Approach 3. Review these slides Important terms and structures important to study prior to NM1 attendance. Relationship of Spinal Nerves to Dura Brachial Plexus ~ Lumbar Plexus ~ Sacral Plexus Layers of the intracranial membrane system Cranial Suture Anatomy – coronal, sagittal, lamdoid Craniospinal Juncture Anatomy Pelvic Ligaments – sacrotuberous, sacrospinous Tentorium Cerebelli – anatomy of intracranial attachments Brachial Plexus Femoral Nerve Sciatic Nerve NM1 Preparation Quiz Answers at the end 1. The coronal suture is between what 2 bones? 2. The occipito-mastoid suture/jugular foramen is between what 2 bones? 3. The falx cerebri is mostly under what suture? 4. The attachments of the tentorium cerebelli are: a. frontal bone, petrouss part of temporal bone, occiput b. occiput, parietal, temporal, bones, anterior and posterior clinoid processes of sphenoid bone c. occiput, parietal and temporal bones, maxilla Neural Manipulation Neural Manipulation (NM) is a gentle hands-on therapy which helps to free up the nerves and the connective tissue around the nerves (dura mater), the bones around the brain (cranium) so that the nervous system functions better. (Barral & Croibier 2007) “Neural” refers to the nervous system of the body, which includes the brain, spinal cord, nerves of upper and lower extremities, trunk and head. Cranial Anatomy Frontal bone Coronal suture Parietal bones Sagittal suture Sutures The sutures provide for growth and elasticity of the newborn skull. The begin to close after 3-4 months, however the anterior fontanelle will not close until 20 months (average). They can be difficult to palpate after 6 months.
  • Anatomy and Physiology Model Guide Book

    Anatomy and Physiology Model Guide Book

    Anatomy & Physiology Model Guide Book Last Updated: August 8, 2013 ii Table of Contents Tissues ........................................................................................................................................................... 7 The Bone (Somso QS 61) ........................................................................................................................... 7 Section of Skin (Somso KS 3 & KS4) .......................................................................................................... 8 Model of the Lymphatic System in the Human Body ............................................................................. 11 Bone Structure ........................................................................................................................................ 12 Skeletal System ........................................................................................................................................... 13 The Skull .................................................................................................................................................. 13 Artificial Exploded Human Skull (Somso QS 9)........................................................................................ 14 Skull ......................................................................................................................................................... 15 Auditory Ossicles ....................................................................................................................................