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An external and cranial mophometric study of altitudinal variation in Microtus arvalis in Switzerland Prescott-Allen, Christine 1981

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AN EXTERNAL AND CRANIAL MORPHOMETRIC STUDY OF ALTITUDINAL VARIATION IN MICROTUS ARVALIS IN SWITZERLAND BY CHRISTINE/ PRESCOTT-ALLEN B.A. MCGILL UNIVERSITY, 1971 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN THE FACULTY OF GRADUATE STUDIES ZOOLOGY DEPARTMENT WE ACCEPT THIS THESIS AS CONFORMING TO THE REQUIRED STANDARD  THE UNIVERSITY OF BRITISH COLUMBIA APRIL, 1981 Q CHRISTINE PRESCOTT-ALLEN, 1981  In p r e s e n t i n g  this  thesis i n partial  f u l f i l m e n t of the  r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e of B r i t i s h Columbia, I agree that it  freely  the L i b r a r y s h a l l  a v a i l a b l e f o r r e f e r e n c e and s t u d y .  agree t h a t p e r m i s s i o n f o r extensive for  University  s c h o l a r l y p u r p o s e s may  for  financial  shall  of  The U n i v e r s i t y o f B r i t i s h 2075 W e s b r o o k P l a c e V a n c o u v e r , Canada V6T lW5v  Columbia  my  It is thesis  n o t be a l l o w e d w i t h o u t my  permission.  Department  thesis  be g r a n t e d by t h e h e a d o f  copying or p u b l i c a t i o n of this  gain  further  copying of t h i s  d e p a r t m e n t o r by h i s o r h e r r e p r e s e n t a t i v e s . understood that  I  make  written  ii ABSTRACT  Cranial  and e x t e r n a l measurements  o f t h e common v o l e , M i c r o t u s  arvalis,  amined t o d e t e r m i n e w h e t h e r t h e y c o r d a n c e w i t h e i t h e r (1)  i n 422 museum  specimens  from S w i t z e r l a n d were  varied with a l t i t u d e in  Bergmann's Rule,  or  (2)  t o sex  segregated  correlation  (1)  age g r o u p s .  In  n o t one o f t h e 108 t e s t s  r e v i e w i n g the b a s i c (2)  concepts of the R u l e ,  as a p p l i e d  a f f e c t growth i n M i c r o t u s  i n c l u d i n g c l i m a t e and c o m p e t i t i o n w i t h s y m p a t r i c  which might  have  T h e r e were s e v e r a l  i n f l u e n c e d the  - the nominate  subspecies  subspecies  Longchamps,  1841)  of M i c r o t u s  M. a . a r v a l i s  congenerics;  phenotypically c o n s i d e r e d by  o r M. a .  arvalis  the  Switzerland  1779)  and a  (Selys-  (Schinz,  1845).  f o r the e x i s t e n c e of  two  and d i s t r i b u t i o n a l l y d i s t i n c t s u b s p e c i e s  was  (1)  examining  study  in  (Pallas,  rufescentefuscus  the general appropriateness  v a r i a t i o n in Microtus  and (2)  computations.  c a l l e d e i t h e r M. a. i n c e r t u s  The l a c k o f e v i d e n c e i n t h i s  tion;  se-  arvalis,  r e f e r e n c e s i n the l i t e r a t u r e to  o c c u r r e n c e o f two s u b s p e c i e s  cussing  to  i n d i c a t i n g d e f i c i e n c i e s i n d a t a d e r i v e d from museum  specimens  montane  was  investigated  i d e n t i f y i n g t h e m a j o r a b i o t i c and b i o t i c  l e c t i o n p r e s s u r e s t h a t might  (3)  in-  significant.  homeotherms;  and  were  e a c h o f w h i c h had been d i v i d e d  The l a c k o f a d h e r e n c e t o B e r g m a n n ' s R u l e was by  ac-  subspeciation.  C o r r e l a t i o n c o e f f i c i e n t s b e t w e e n a l t i t u d e and s i z e c a l c u l a t e d on 32 d i m e n s i o n s ,  ex-  arvalis  i n terms of  r e v i e w i n g t h e l i t e r a t u r e on t h e  of  dis-  subspecia-  diagnostic  iii characters  and d i s t r i b u t i o n a t t r i b u t e d t o t h e montane  Two m a j o r c o n c l u s i o n s Bergmann's Rule definitions precepts taxonomic apply  should  were d r a w n .  to a m a j o r i t y of the cases  second c o n c l u s i o n i n which growth  is  large continuous, fraspecific  highly  is  f o r which i t i s  for species  like  it  and  is  the  found  intended.  Microtus  to The  arvalis,  v a r i a b l e and d i s t r i b u t i o n i s  the u s e f u l n e s s  populations  the g e o g r a p h i c documented.  was t h a t  - and u n t i l  firm  founding  to which i t a p p l i e s  at which i t a p p l i e s  that  a "rule" until  on a t l e a s t two o f i t s  - the groups of animals level  The f i r s t was  n o t be c o n s i d e r e d  are e s t a b l i s h e d  morph.  by  and  of formal r e c o g n i t i o n of  in-  questionable  until  v a r i a b i l i t y of the s p e c i e s  an o v e r v i e w  as a whole i s  of  well  iv  T A B L E OF CONTENTS  TITLE  PAGE  i  ABSTRACT  i  i  T A B L E OF CONTENTS  iv  LIST  OF T A B L E S  I N TEXT  v i  LIST  OF T A B L E S  IN APPENDIX  LIST  OF F I G U R E S  v i i  I N TEXT  x  ACKNOWLEDGEMENTS  x i  INTRODUCTION  1  MATERIALS  2  AND METHODS  1.  S k u l l Dimensions 1.1. C r a n i a l measurements 1.2. Mandibular measurements  2 2 10  2.  E x t e r n a l Dimensions 2.1. Measuring techniques 2.2. Technicians- involved  10 10 12  3.  Sexual  4.  Aging 4.1.  5.  A l t i t u d e and S i z e 5.1. Home r a n g e 5.2. Migration  i n the literature  Differences  12  t h e Specimens.. Aging techniques i n the l i t e r a t u r e  ,  12 12 14 15 15  RESULTS  18  1.  Skull 1.1. 1.2. 1.3.  Dimensions Paired structures Measuring consistency Brachyostism  2.  Sexual Differences 2.1. Sexual dimorphism 2.2. Sex r a t i o  3.  Aging 3.1. 3.2. 3.3.  4.  A l t i t u d e and S i z e  t h e Specimens Aging techniques applied... Changes i n i n d i v i d u a l d i m e n s i o n s Changes i n p r o p o r t i o n a l r e l a t i o n s h i p s Correlation  18 18 18 21 21 21 23 26 ...26 33 ..34 34  V  DISCUSSION...:^I.:.::...:.....:..............................37 1.  Bergmann's Rule 111. General review of Bergmann's 1.2. B e r g m a n n ' s r u l e and M i e r o t u s  2.  Subspeciation 2.1. Morphological variation 2.2. G e o g r a p h i c range 2.3. Nomenclature  rule arvalis  AO AO AA 5A 73 76 77  CONCLUSIONS  80  APPENDIX  81  LITERATURE CITED  125  vi  h1ST - OF•TABLES - IN-TEXT  T a b l e I:  L i t e r a t u r e S y n o p s i s o f Home Range S i z e  i n Microtus  arvalis Table  II:  Table  III:  16  Left-Right  Skull  Consistency  Differences  of Measuring  i n Microtus  Technique  20  Table  IV: L i t e r a t u r e S y n o p s i s of Sexual Dimorphism arvalis T a b l e V: L i t e r a t u r e S y n o p s i s o f S e a s o n a l Sex R a t i o arvalis T a b l e V I : Age and Sex A n a l y s i s of Capture  i n Microtus  arvalis  T a b l e V I I : L i t e r a t u r e S y n o p s i s o f Body L e n g t h Microtus a r v a l i s T a b l e V I I I : L i t e r a t u r e S y n o p s i s o f Body L e n g t h crotus arvalis Table  I X : Body L e n g t h s Microtus arvalis  from T a b l e s  a r v a l i s . . . . 19  in  Microtus  i n Microtus by Month  i n Laboratory i n Wild M i -  V I I and V I I I o f J u v e n i l e  T a b l e X: L i t e r a t u r e S y n o p s i s o f D a t e o f B i r t h and G r o w t h of Microtus Table  of Microtus  i n Body L e n g t h  (Dottrens,  T a b l e X V : Some M e t h o d s  25 27 28 29  31 arvalis  Specimens..33  Table X I I : Comparison o f C r a n i a l Measurements i n M i c r o t u s a r v a l i s Age C l a s s e s Table X I I I : P r o p o r t i o n a l R e l a t i o n s h i p s i n S k u l l Dimensions i n Two S p e c i e s o f M i c r o t u s  arvalis  24  Rate  arvalis  X I : Age and Sex A n a l y s i s  Table XIV: Trends  22  with Altitude  1962; Jones,  35 36  i n Microtus  1970; This  Thesis)  of Thermoregulation  37 42  Table X V I : L i f e H i s t o r i e s o f Three S p e c i e s o f M i c r o t u s 55 T a b l e X V I I : C o m p a r i s o n o f Body L e n g t h s w i t h A l t i t u d e i n M i c r o tus a r v a l i s ( D o t t r e n s , 1962; Jones, 1970; This T h e s i s ) . . 7 4 Table X V I I I : B r i e f Nomenclatural rufescentefuscus Table XIX: B r i e f incertus  Nomenclatural  History  History  of Microtus  of Microtus  arvalis arvalis  78 79  LIST•OF - TABLES-IN  APPENDIX  Appendix Table l a : Measuring Accuracy: Total Skull  Length o f  .  82  Appendix Table l b : Measuring Accuracy: Nasal Width  83  Appendix  84  Table l c : Measuring Accuracy: Rostrum Width  Appendix Table I d : Measuring Accuracy: Zygomatic Width Appendix Table l e : Measuring Accuracy: Least  85  Interorbital  Width  86  Appendix Table I f : Measuring Accuracy: B r a i n c a s e Width  87  A p p e n d i x T a b l e l g : M e a s u r i n g A c c u r a c y : P o s t Zygoma W i d t h  88  Appendix Table l h : Measuring Accuracy: Mastoid Width  89  A p p e n d i x T a b l e l i : M e a s u r i n g A c c u r a c y : F o r a m e n Magnum Height A p p e n d i x T a b l e l j : M e a s u r i n g A c c u r a c y : F o r a m e n Magnum  90  Width  91  Appendix Table l k : Measuring Accuracy: S k u l l  Height  92  Appendix Table 11: Measuring Accuracy: Condylobasal Length...93 Appendix Table lm: Measuring A c c u r a c y : B a s a l Length  94  Appendix Table I n : Measuring Appendix Table l o : Measuring Appendix Table l p : Measuring Length Appendix Table l q ; Measuring Length  95 96  Accuracy: B a s i l a r Length Accuracy: P a l a t a l Length Accuracy: Palatal Bridge  97 Accuracy: Palatal  Appendix Table l r : Measuring Accuracy: P a l a t a l Width Appendix Appendix  Foramen 98 Foramen 99  Table I s : Measuring Accuracy: Maxillary Length  Diastema  Table I t : Measuring Accuracy: Maxillary Length  T o o t h Row  100 101  vi i i Appendix  Table l u : Measuring l a r width  Accuracy: Maxillary  Outer  Appendix  Table l v : Measuring l a r Width  Accuracy: Maxillary  Inner  Appendix  Table lw: Measuring  Accuracy: Mandible  Appendix  Table l x : Measuring  A c c u r a c y : Jaw L e n g t h  Appendix  Table l y : Measuring Row L e n g t h  Accuracy: Mandibular  Appendix  Appendix  Appendix  Mo. . . . . . 102 Mo-  Length  103 104 105  Tooth 106  T a b l e 2 a : S e x u a l D i m o r p h i s m and . C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and T o t a l L e n g t h o f S k u l l , N a s a l W i d t h , Rostrum Width  107  T a b l e 2 b : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c e n t f o r A l t i t u d e and Z y g o m a t i c W i d t h , L e a s t I n t e r o r b i t a l Width, B r a i n c a s e Width  108  T a b l e 2 c : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and P o s t Zygoma W i d t h , M a s t o i d W i d t h , Foramen Magnum H e i g h t  109  Appendix  T a b l e 2 d : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and Foramen Magnum W i d t h , S k u l l Height, Condylobasal Length ...110  Appendix  T a b l e 2 e : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and B a s a l L e n g t h , B a s i l a r Length, P a l a t a l Length  Ill  T a b l e 2 f : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and P a l a t a l B r i d g e L e n g t h , P a l a t a l Foramen L e n g t h , P a l a t a l Foramen W i d t h  112  T a b l e 2 g : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and M a x i l l a r y D i a s t e m a L e n g t h , M a x i l l a r y T o o t h Row L e n g t h , M a x i l l a r y Outer Molar Width  113  T a b l e 2 h : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and M a x i l l a r y I n n e r M o l a r Width, Molar Width, Mandible Length  114  T a b l e 2 i : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and Jaw L e n g t h , M a n d i b u l a r D i a s t e m a L e n g t h , M a n d i b u l a r T o o t h Row L e n g t h  115  T a b l e 2 j : S e x u a l D i m o r p h i s m and C o r r e l a t i o n C o e f f i c i e n t f o r A l t i t u d e and Body ( P l u s Head) L e n g t h , T a i l L e n g t h , Hind Foot Length  116  Appendix  Appendix  Appendix  Appendix  Appendix  Appendix  T a b l e 2k: S e x u a l Dimorphism and C o r r e l a t i o n Coeff i c i e n t f o r A l t i t u d e and E a r L e n g t h , Weight  117  Appendix  T a b l e 3 a : Age D i f f e r e n c e s i n T o t a l L e n g t h o f S k u l l , Nasal Width, Rostrum Width, Zygomatic Width, Least I n t e r o r b i t a l Width, Braincase Width...118  Appendix  T a b l e 3 b : Age D i f f e r e n c e s i n P o s t Zygoma W i d t h , M a s t o i d W i d t h , F o r a m e n Magnum H e i g h t , F o r a m e n Magnum W i d t h , S k u l l H e i g h t , C o n d y l o b a s a l L e n g t h . . . . 1 1 9  Appendix  T a b l e 3 c : Age D i f f e r e n c e s i n B a s a l L e n g t h , B a s i l a r Length, P a l a t a l Length, P a l a t a l Bridge L e n g t h , P a l a t a l Foramen L e n g t h , P a l a t a l Foramen Width  120  T a b l e 3 d : Age D i f f e r e n c e s i n M a x i l l a r y Diastema L e n g t h , M a x i l l a r y T o o t h Row L e n g t h , M a x i l l a r y Outer Molar Width, M a x i l l a r y Inner Molar Width, M a x i l l a r y Molar Width, Mandible Length  121  T a b l e 3 e : Age D i f f e r e n c e s i n Jaw L e n g t h , M a n d i b u l a r D i a s t e m a L e n g t h , M a n d i b u l a r T o o t h Row L e n g t h , Body L e n g t h , T a i l L e n g t h , H i n d F o o t Length  122  Appendix  Table  123  Appendix  T a b l e 4: P r o p o r t i o n a l R e l a t i o n s h i p s w i t h C r a n i a l Measurements o f M i c r o t u s a r v a l i s  Appendix  Appendix  3 f : Age D i f f e r e n c e s  i n Ear Length,  Weight. Age i n  124  x  LIST-OF-FIGURES-IN-TEXT  Figure  1; S i t e o f C a p t u r e and Number of S p e c i m e n s in Switzerland  Figure  2: L a t e r a l  Figure  3 : Foramen Magnum M e a s u r e m e n t s  A  Figure  A; M a n d i b u l a r M e a s u r e m e n t s  A  Figure  5: D o r s a l  6  Figure  6; V e n t r a l C r a n i a l M e a s u r e m e n t s  Figure  7: N o n - j u v e n i l e Body L e n g t h s i n M i c r o t u s a r v a l i s ( D o t t r e n s , 1962; Jones, 1970; T h i s T h e s i s )  38  8: A l t i t u d e  51  Figure  by  Canton  C r a n i a l Measurements  ....A  C r a n i a l Measurements  Ranges o f  Three S p e c i e s  3  8  of M i c r o t u s  xi ACKNOWLEDGEMENTS  I which Dr.  thank lent  the following  the Microtus  F. B a u d  (Musee  persons  i n the s i x institutions  material:  d'Histoire  Naturelle),  (Zoologisches  Museum d e r U n i v e r s i t a t  (Zoologisches  Forschungsinstitut  Dr.  P. L u p s  (Station Muller  (Naturhistorisches  Federale  (Bundner  I  also  German a r t i c l e s , ings, Dr.  Verena  Harold  I Seal,  want  Nordan  advice  Dr.  J . Mary my l o n g  able  mentor,  i n guiding  direction.  D r . A. M e y l a n  f o rher t r a n s l a t i o n s of  f o r h i s map a n d s k u l l  thank  tolerance  t h e computer  draft.  - and  with  - gave  h i s time  analysis;  who s o c o n s i s t e n t l y  cheerful  and  you t o Dr. H i l a r y  who s o g e n e r o u s l y  my s u p e r v i s o r ,  draw-  D r . H. D e a n F i s h e r  special  me t h r o u g h  work w i t h  Koenig),  und Nationalparkmuseum).  Jurgensen  Virolle  a very  Hutterer  Agronomiques) and Dr. J .  humour and s u r p r i s i n g  distance  Bern),  f o rher typing,  t o express  Taylor,  D r . R.  f o r t h e i r comments on t h e f i r s t  my s t a t i s t i c a l  and  ed  Patrick  Wiestner  characteristic  Museum  Naturhistorisches  Marguerite  Zurich),  Claude  u n d Museum A l e x a n d e r  de R e c h e r c h e s  thank  D r . C.  encouragement  and t o support-  and  valu-  1 INTRODUCTION  In  S w i t z e r l a n d , t h e common v o l e , M i c r o t u s  found a t a l t i t u d e s r a n g i n g 2600 m e t e r s . ly  This  small country  in geographic 1962;  Jones,  on t h i s  To d a t e two m a j o r r e p o r t s  1970) and an a r t i c l e Also,  (Lehmann,  (Dottrens,  1967) have  appeared  r e f e r e n c e s t o v a r i a t i o n i n t h e common  have been made i n a t l e a s t two books  A l l e e and S c h m i d t ,  the statements  in a relative-  the vole a prime c a n d i d a t e f o r s t u d i e s  variation.  subject.  is  300 m e t e r s t o  2000+ m e t e r a l t i t u d i n a l r a n g e  makes  vole i n the Alps 1914;  from a p p r o x i m a t e l y  arvalis,  made a b o u t  1962).  (Gdldi,  However t h e d a t a p r e s e n t e d and  geographic  v a r i a t i o n i n Mj_ a r v a l i s  conflict. On t h e one hand t h e r e a r e t h o s e i n M. a r v a l i s which s i z e Schmidt,  e x e m p l i f i e s Bergmann's  increases  1962).  who c o n t e n d t h a t R u l e by s h o w i n g  On t h e o t h e r hand t h e r e a r e t h o s e who r e p o r t  Rule at a l l (Dottrens,  p a r t o f i t s range mann,  1967),  a cline in  w i t h a l t i t u d e ( G o l d i , 1 9 1 2 ; A l l e e and  t h a t a l t i t u d i n a l v a r i a t i o n i n M. a r v a l i s Bergmann's  variation  (Jones,  explains  1970).  e i t h e r does  1962) o r o n l y This  group  the v a r i a t i o n trends  not f o l l o w  follows i t in  (including  i n terms of  Lehsubspe-  ciation. This  study  Swiss specimens  analyses  c r a n i a l and e x t e r n a l d i m e n s i o n s  of Microtus  arvalis  whether t h e r e i s m o r p h o l o g i c a l to d e t e r m i n e whether such or,  i n an a t t e m p t t o e s t a b l i s h  v a r i a t i o n with a l t i t u d e ; i f so,  v a r i a t i o n conforms  a l t e r n a t i v e l y , suggests s u b s p e c i a t i o n ;  the reasons  why n o t .  in  t o Bergmann's  Rule  i f not, to explore  2 MATERIALS-AND•METHODS  Four  h u n d r e d and t w e n t y - t w o s p e c i m e n s  were e x a m i n e d . (Figure  1)  A l l the animals  b e t w e e n 1930  vole c o l l e c t i o n s of  of M i c r o t u s  were t r a p p e d i n  and 1978.  They  f o u r S w i s s museums,  cultural  s t a t i o n and one German museum  Two s e t s  o f measurements  (Appendix  common  1; page  81).  measurements t a k e n by  this  were p e r f o r m e d on a CDC 6500  c o m p u t e r a t t h e E c o l e P o l y t ! e c h n i q u e F e d e r a l e de 1.  the  a Swiss f e d e r a l a g r i -  were s t u d i e d ; e x t e r n a l  A l l computer a n a l y s e s  Switzerland  represent  t a k e n by t h e c o l l e c t o r s and s k u l l measurements author.  arvalis  Lausanne.  Skull-Dimensions A total  dibular  of  27 s k u l l d i m e n s i o n s  - 23 c r a n i a l and f o u r  - was r e c o r d e d f o r e a c h s p e c i m e n t o t h e n e a r e s t  millimeter  by means o f d i a l  illustrated  in Figures  2-6  calipers.  These  man-  .05  measurements  and a r e d e s c r i b e d as  are  follows:  1.1.  Cran_ia_l Mea;surement_ s_  1.1.1.  G r e a t e s ^ _ l e n g _ t h p _ f _ s k u _ l l ; maximum d i s t a n c e  from the  most p o s t e r i o r p r o j e c t i o n o f t h e o c c i p i t a l  condyles  to the  anterior  (Snyder,  1954;  vear, Figure  part of the curve of the i n c i s o r s  most Rose-  1969). 2  1.1.2.  Distance  AB  Page  Condy_lobasal_leng_th:  4 distance  from the a n t e r i o r  edge  of the median i n c i s i v e a l v e o l i t o the p o s t e r i o r p r o j e c t i o n of the o c c i p i t a l Figure  2  condyles.  Distance  AC  Page  4  FigureV.Site of capture and number of specimens by canton in Switzerland ^  FIGURE 2: A  LATERAL CRANIAL MEASUREMENTS  A FIGURE 3:  FORAMEN MAGNUM MEASUREMENTS  FIGURE -4: MANDIBULAR MEASUREMENTS B  C  D  5 1.1.3.  Nasal^ w i d t h :  ( D e B l a s e and M a r t i n , Figure  5  1.1.4.  maximum d i s t a n c e a c r o s s  Line A  Page  e a c h arm o f t h e c a l i p e r s  needlepoint  infraorbital Figure  6  distance across  i o r edge o f t h e m a x i l l a r y the  5  o r b i t s taken  processes  s  Page  dorsally  across  5  Line C  Page  bones ( R o s e v e a r , 1 9 6 9 ;  1974). 6  Z_yg_oma_ti£ width_: d i s t a n c e b e t w e e n t h e o u t e r  zygomatic arches at the widest  Glass,  1 9 7 4 ; D e B l a s e and M a r t i n ,  Figure  5  taken  o f t h e zygoma, and p l a c i n g  the f r o n t a l  Figure  1.1.7.  Line D Braincase  just  Page width:  points  5  1974).  6  maximum d i s t a n c e a c r o s s  1.1.8.  Figure  Page  Post_z^goma w i d t h :  diately  1.1.9.  Line F  minimum w i d t h  Maj;toid_ w i d t h :  cesses  (Glass,  Figure  5  Page  maximum w i d t h  Page  1974).  o f t h e c r a n i u m immeo f t h e zygoma.  6  1974).  Line G  of the  6  p o s t e r i o r t o t h e squamosal processes 5  the braincase  above t h e r o o t s o f t h e s q u a m o s a l p r o c e s s e s  Line E  edges o f  (Rosevear, 1969;  zygoma ( R o s e v e a r , 1 9 6 9 ; D e B l a s e and M a r t i n , Figure  anter-  i.nt_e£0£b_ital_w_id_th: minimum d i s t a n c e b e t w e e n t h e  1 9 7 4 ; D e B l a s e and M a r t i n  the  the respective  6  Glass,  1.1.-6.  against  t a k e n by  o f e a c h arm on t h e m a x i l l a b e l o w t h e r e s p e c t i v e  Line B e  the rostrum  foramen.  .__.__J_  1.1.5.  bones  1974).  Rostrum w i d t h :  backing  the nasal  6  across  the mastoid  pro-  FIGURE 5:  DORSAL CRANIAL MEASUREMENTS  1.1.10.  Fojramerj m a £ n u m _ h e i £ h _ t : maximum h e i g h t  o f the foramen  magnum. Figure  Line A  3  1.1.11.  Page  4  Foramen^ ma£num_w_id_thj_ maximum w i d t h o f t h e f o r a m e n  magnum. Figure  3  1.1.12.  Line  B  Page  4  S k u l _ l he _ig h t : d i s t a n c e  from the d o r s a l  f r o n t a l bones t o t h e v e n t r a l midline rectly  of the palatine  anterior to the mesopterygoid fossa  1.1.13.  Ba_sa_l . l e n g t h y  distance  midline  of the  bones d i -  (Rosevear, 1969).  from t h e a n t e r i o r edge o f t h e  median i n c i s i v e a l v e o l i t o t h e a n t e r i o r m i d - p o i n t on t h e lower e d g e o f t h e f o r a m e n magnum  (Glass,  1974; D e B l a s e and  Martin,  1974). 1.1.14. the  Ba^i_lar j^eng^h^ d i s t a n c e  from t h e p o s t e r i o r edge o f  median i n c i s i v e a l v e o l i t o t h e a n t e r i o r m i d - p o i n t on t h e  l o w e r e d g e o f t h e f o r a m e n magnum ( G l a s s , tin,  1974; D e B l a s e and Mar-  1974).  Figure  6  1'.-. 1 . 1 5 .  Distance  AK  Page  Pal.at_a_l . l e n g t h y  8  distance  from t h e a n t e r i o r edge o f t h e  median i n c i s i v e a l v e o l i t o t h e a n t e r i o r edge o f t h e m e s o p t e r y goid  fossa.  1.1.16.  Pa.la.ta_l jar_idjg e_le_n£t_h: d i s t a n c e i  from t h e p o s t e r i o r  edge o f t h e r i g h t p a l a t a l foramen t o t h e a n t e r i o r edge o f t h e mesopterygoid Figure  6  1.1.17.  fossa  Distance  (Chicago Natural EI  Page  History  Museum, 1 9 6 2 ) .  8  Pa_la_ta_l . f o r a m e n l e n g t _ h j _ d i s t a n c e  from the a n t e r i o r  edge t o t h e p o s t e r i o r edge o f t h e r i g h t p a l a t a l foramen. Figure  6  Distance  CE  Page  8  FIGURE 6:  VENTRAL CRANIAL MEASUREMENTS  9  Palatal  1.1.18. of  the right  palatal Figure  palatal  foramen 6  _foramen w i d t h :  Line  foramen  Page  edge o f t h e a l v e o l u s the  alevolus  Martin, Figure  edge o f t h e l e f t  distance  from  the posterior  I"*" t o t h e a n t e r i o r e d g e o f  of the right  molar  row ( D e B l a s e and  1974). 6  1.1.20.  Distance  BF  Page  8  M a x i j ^ l a r ^ t _ o o t h _row jLengt_hj_ d i s t a n c e  from  the anterior  edge o f t h e a l v e o l u s  o f t h e M"*" t o t h e p o s t e r i o r e d g e  alveolus  of the last  cheektooth i n the right  (DeBlase  and M a r t i n ,  1974).  Figure  6  1.1.21.  Distance  FJ  Page  -  edge o f t h e a l e v o l u s  Figure  6  1.1.22.  G  Figure  of the M  edge o f t h e a l v e o l u s 6  1.1.23.  o f t h e M"*" o f t h e l e f t  Page  of the alveolus  Line  H  i s subtracted  1  of the right  of the M  remainder  from  the l a b i a l  row t o t h e molar  row.  8  Page  Mo_la£ w i d t h _ :  row  from  molar  Ma>^i_llary_ i^nnei; m o l a r ; w i d t j n : d i s t a n c e  lingual  width  Line  distance  o f t h e M"* o f t h e r i g h t  labial  molar  of the  8  M a x i _ l l a _ r y _ o u t e r ; mo_lar w i d t h :  edge o f t h e a l v e o l u s  edge  edge  8  of the right  of the  the l a b i a l  point.  Ma2<i.lla_ry_ d _ i a s t e m a _ l e n c ) t h :  1.1.19.  from  to the l a b i a l  at the widest D  distance  from  molar  the lingual  row t o t h e  of the l e f t  molar  row.  when t h e m a x i l l a r y  inner  molar  1  8  the maxillary outer  molar  width.  1.2.  Mand^bular_Measur_ement£  1.2.1.  Mand^b_le_leng_th: d i s t a n c e  the  right  lus  o f the r i g h t  Figure  condyloid process  4  1.2.2.  a w  AE  4  1.2.3.  4 f r o m t h e p o s t e r i o r edge o f t h e  e  o f the l a s t  Figure  Page  . l. 2.gth2_ d i s t a n c e  a l v e o l u s o f the r i g h t lus  t o t h e a n t e r i o r edge o f t h e a l v e o -  incisor.  Distance £  1^ t o t h e p o s t e r i o r edge o f t h e a l v e o -  cheektooth o f the r i g h t  Distance  BD  Page  Mand^bular_tooth_row_leng_th:  edge o f t h e a l v e o l u s o f t h e  the  a l v e o l u s o f the l a s t 4  1.2.4. ular  Distance  row l e n g t h  External Five  Page  _lengt_hj_ r e m a i n d e r when t h e mandib-  2.1.  e x t e r n a l body d i m e n s i o n s were a n a l y s e d : tail  length; hind  weight;  foot length;  The e x t e r n a l d i m e n s i o n s were p r o v i d e d  body  and e a r  b y numerous,  using unspecified  techniques.  depending b o t h on the measuring  a d o p t e d and on t h e p e r s o n p e r f o r m i n g  the technique,  s e t o f e x t e r n a l measurements may be u n r e l i a b l e . Measuring_ T e c h n i q u e s _ i n J e w e l l and F u l l a g a r  ing  from t h e jaw l e n g t h .  Dimensions  B e c a u s e measurements c a n v a r y  this  m o l a r row.  4  mostly u n i d e n t i f i e d c o l l e c t o r s  technique  from t h e a n t e r -  t o t h e p o s t e r i o r edge o f  i s subtracted  ( i n c l u d i n g head) l e n g t h ; length.  distance  cheektooth o f the r i g h t  Mandibu 1 ar_di_as_tema  tooth  2.  CD  m o l a r row.  4  ior  Figure  f r o m t h e p o s t e r i o r edge o f  t h e body and t a i l  t h e Lit_erat_ure  (1966) l i s t e d  five  methods o f m e a s u r -  l e n g t h : B r i t i s h Museum O l d ;  Hanging;  11 Morrison-Scott viders.  P i n s ; B r i t i s h Museum New;  They a l s o n o t e d t h a t t h e  most v u l n e r a b l e sinuous able  when h a n d l e d c a n  alone  (1965) l i s t e d  anus.  A l s o the  f i v e percent  s p i n a l column t a k e s s t r e t c h e d out  a  to a  vari-  t h r e e methods o f m e a s u r i n g t h e  (Dale,  the  l e n g t h may  californieus  interval  vary  tail  pelvis  or  considerably,  sanctidiegi  b e t w e e n d e a t h and  up  (Howell,  measurement  1940).  Even t h e h e e l to the or without others  tail  i n Microtus  1 9 2 4 ) , d e p e n d i n g on  hind  end  f o o t can  of the  be measured i n two  longest  i t . Conventionally  s i n e ungue ( A n s e l l ,  ment, but that  e v e n i t has i t was  or immediately  digit,  nail  ungue;  a most s t r a i g h t f o r w a r d m e a s u r e -  p o t e n t i a l hazards.  Cantuel  animal  while  (1940)  ad-  i t was  alive  i t s d e a t h as w e i g h t d r o p s q u i c k l y once  decomposition  begins.  of weight are  relatively  He  a r g u e d , however, t h a t p r e c i s e , given  w e i g h e d w i t h an e r r o r f a c t o r o f o n l y  o b t a i n the  the  e i t h e r i n c l u d i n g the  A m e r i c a n s measure cum  b e t t e r t o w e i g h an  after  ways - f r o m  1965).  W e i g h t m i g h t a p p e a r t o be  be  Di-  body measurement i s  - f r o m i t s j u n c t i o n w i t h t h e body o r w i t h t h e  w i t h the  vised  be  Morrison-Scott  degree. Ansell  to  head and  to e r r o r because the  f o r m and  and  same a p p r o x i m a t i o n  m e t e r measurement must be  measurements  t h a t a kilogramme one  m i l l i g r a m m e but  i n l i n e a r measurement, a  accurate  can  to a m i l l i m e t e r .  kilo-  to  2.2.  Jechnicians  I_nvo.lved  Even i f a l l t h e c o l l e c t o r s used  still  (1966:  be a s o u r c e o f v a r i a t i o n .  page  believe  50?) concluded  they  syncracies  are using  of procedure  that  f o r comparison".  3.  Sexual Of  tors.  Apart  from  render t h e i r  and  Fullagar  even  though  develop  they idio-  measurements un-  367 had b e e n s e x e d  t h e 55 s p e c i m e n s  f e m a l e s and 137  by t h e c o l l e c -  o f unknown s e x , t h e r e  were  males.  Aging-the-Specimens Ag_in£ J _ e c h n i q u e s _ i n t_he L i t e r a t u r e  4.1.  Aging lems,  Microtus arvalis  y e t an age a n a l y s i s ,  necessary  cause  age ( D o u t t ,  poses  no m a t t e r how  within  formidable prob-  rough  and  imperfect,  as t h e g r e a t e s t  a s p e c i e s c a n p r o b a b l y be  single  ascribed  1955).  S ku 11 c ha rac_t e r i s t _ i c_s  The skull  specimens  f o r any s t u d y o f v a r i a t i o n  of variation  4.1.1.  t h r e e major  approaches  characteristics  velopment applied  and t o o t h  include  wear.  because  i n which  to aging microtines  analysing  Aging  to Microtus arvalis.  pennsy1vanicus aid  "two w o r k e r s ,  involved  Differences  t h e 422 s p e c i m e n s ,  4.  to  that  Jewell  t h e same t e c h n i q u e s , may  suitable  is  a n a l y s e d had  t h e same t e c h n i q u e s , t h e n u m b e r o f i n d i v i d u a l s  could  230  o f the specimens  Snyder  (1954)  some d i s a p p e a r q u i t e  suture lines,  by s u t u r e l i n e s  In t h i s  early  through crest  cannot  de-  be  i t resembles Microtus found while  suture lines others  o f no  remain  13 apparent unlike  even  M. p e n n s y 1 v a n i c u s ,  manifest crests tooth  visible  with wear  with  changes  age.  Spitz  t h e common v o l e i n lambdoidal  does  not appear t o  or mastoid-exoccipital  (1974) s t a t e d t h a t  i s out of the question  1959 Lord  termining Louarn  Spitz  described  the dry weight  (1971)  technique  as t h e r e  (1974) t h e only  common  4.1.3.  vole.  aging  M. a r v a l i s  i s no  variation  by  However,  literature Within  (1966),  Le  I t has been c a l l e d  by  p r e c i s e method o f age d e t e r m i n a t i o n i n i t i s n o t p o s s i b l e t o a g e museum  most o f t e n u s e d  rodents  (D. C h i t t y ,  i t was d e c i d e d  study  by w e i g h t  on t h e d r a w b a c k s any g i v e n  may d i f f e r  A 40 d a y o l d m a l e , (Frank,  1952; Krebs  of this  age c o h o r t  Also with  when f o o d  1972) and i n pregnant  small  and Myers,  n o t t o a g e t h e museum s p e c i -  approach.  i n Microtus (Meunier  arvalis  and S o l a r i ,  f o r e x a m p l e , may be a s h e a v y  1957).  diminished  t o age l i v i n g  c l a s s due t o comments i n t h e  significantly  and, i n females,  weight  Martinet  ( 1 9 7 2 ) a p p l i e d t h e same  o f M. a r v a l i s .  i s the index  mens i n t h i s  animals  r a b b i t s by d e -  technique.  mammals, e s p e c i a l l y  weights  of aging  B o d y_we i g_h_t  Weight  1974).  and S o l a r i  However,  by t h i s  a process  o f t h e eye l e n s .  and M e u n i e r  t o specimens  specimens  ari,  Unfortunately,  Eye l.ens_  In  diet  individuals.  time.  4.1.2.  the  i n the oldest  weight  In captive  was u n f a v o u r a b l e females  litter  sexes, common  (Meunier  weight  1972).  as very o l d  v a r i e s i n both  pregnancy.  body  with voles,  and S o l -  was f o u n d t o  amount t o 5 3 . 2 % (1957) tions  of the mother's  weight  concluded t h e r e f o r e that every  (Frank, analysis  1957).  Frank  of vole  popula-  f o r age c l a s s i f i c a t i o n based on w e i g h t must l e a d t o  ous e r r o r s ,  and c o n c l u s i o n s  seri-  b a s e d on them w o u l d be s u b j e c t  to  question. A. 1.4.  Body_lenc_th  Body l e n g t h a l s o tus this  arvalis study  has  been u s e d as a method o f a g i n g  and i t was t h e benchmark were aged  since  aging  by w h i c h t h e s p e c i m e n s  errors  due t o p r e g n a n c y  be a v o i d e d and s i n c e more o f t h e s p e c i m e n s length than weight  (281  versus 2A5).  body  length  i n t h e common v o l e i s  tion  (Frank  and Zimmermann,  (1972) h y p o t h e s i z e d l i n k e d with weight with d i e t .  had d a t a on  However,  like  Also  Meunier  could  body  weight,  and  l e n g t h i n M. a r v a l i s  w h i c h made i t  s u s c e p t i b l e to  Solari  was  strongly  fluctuations  A d d i t i o n a l f a c t o r s a r e t h e s e a s o n o f b i r t h and  ulation cycling.  These  in  subject to i n d i v i d u a l v a r i a -  1957b).  t h a t body  Micro-  are discussed  pop-  on p a g e s 30 and A5 t o  A6  respectively. 5.  A l t i t u d e - and The f i n a l  Size  area of q u a n t i t a t i v e a n a l y s i s  t i o n s h i p b e t w e e n s p e c i m e n s i z e and a l t i t u d e .  i n v o l v e d the While  information  was p r o v i d e d on t h e a l t i t u d e s a t w h i c h a l l A22 s p e c i m e n s been t r a p p e d , two m a j o r p o i n t s what r e p r e s e n t e d t h e r e a l These were home r a n g e  had t o be c o n s i d e r e d  in  a l t i t u d e at which each a n i m a l  and m i g r a t i o n .  rela-  had  deciding lived.  Home r a n g e its  is  " t h a t area traversed  normal a c t i v i t i e s  young"  (Burt,  Spitz  (1970a)  o f M. a r v a l i s lives, Elton  1943;  of  food g a t h e r i n g ,  page  i n d i v i d u a l s were u s u a l l y  work by Fenyuk  t o wander  stacks  far; in fact  i n t h e same  stacks.  open p a s t u r e , kilometers, the s t a c k s  When, h o w e v e r ,  showed  very  in  were  little  their  flood.  (1938)  which  re-  inclination  far,  retaken  t h e y were r e l e a s e d  sometimes  on  as much as 2  a remarkable c a p a c i t y f o r Elton  1/2  finding  (1965)  r e s e a r c h c o n d u c t e d by V a r s h a v s k i i  w h i c h 51% o f t h e v o l e s  also  (1937)  marked and r e c a p t u r e d had moved a  t a n c e b e t w e e n 0 and 100 m e t e r s o n l y . home r a n g e  s u c h as a  When t h e y  i n w h i c h t h e y had l i v e d b e f o r e .  c i t e d M. a r v a l i s  vital")  126 o u t o f 131 were s u b s e q u e n t l y  t h e y wandered  and t h e y  t h e y showed  ("domaine  s t a b l e throughout  and S h e i k i n a  were marked and r e l e a s e d .  leased i n old straw  for  351).  except i n the case of a major event  common v o l e s  in  m a t i n g and c a r i n g  f o u n d t h a t t h e home r a n g e s  (1965) c i t e d  5.2.  by t h e i n d i v i d u a l  d a t a on M i c r o t u s  arvalis  Table  in dis-  I summarizes  the  f r o m T a b l e XVId (page  58).  Mi£ra_ti_0£i Many a l p i n e mammals make r e g u l a r  ward and downward  to avoid  ( A l l e e and S c h m i d t , trapped at  1000 m e t e r s  has m i g r a t e d t o t h i s higher  1962).  migrations  f o o d s c a r c i t y and s e v e r e Might  t h e n a M. a r v a l i s  i n October be, i n f a c t ,  lower a l t i t u d e  summer a l t i t u d e  seasonal  weather specimen  an a n i m a l  f o r the w i n t e r from  o f 2500 m e t e r s ?  It  is  up-  possible,  which its but  TABLE I: LITERATURE SYNOPSIS OF HOME RANGE SIZE IN MICROTUS ARVALIS SEX  LENGTH (METERS) 6.5 m i n J u l y (Dub, 1971a)  DIAMETER (METERS) 10 t o 20 m i n b r e e d i n g season ( F r a n k , 1957)  AREA (SQUARE METERS) 1/4 o f 146 t o 600 s q m ( N i k i t i n a , K a r u l i n and Z e n ' k o v i c h , 1972)  Females 5.8 m i n October (Dub, 1971a)  300 t o 500 s q m i n p l a n t a t i o n ( R e i c h s t e i n , 1960b)  S m a l l e r than 10 m ( S p i t z , 1963b)  Each f a m i l y o c c u p i e s o n l y a few square meters ( B e r n a r d , 1959)  Both 7.5 t o 9 m ( S p i t z e t a l , 1974)  12.9 m i n J u l y (Dub, 1971a)  A few square meters around t h e burrow ( S a i n t G i r o n s , 1973)  40 t o 100 m i n w i n t e r (Frank, 1957)  1/4 o f 828 t o 1008 s q m ( N i k i t i n a , K a r u l i n and Z e n ' k o v i c h , 1972)  Males 6.8 m i n October (Dub, 1971a)  1200 t o 1500 s q m i n p l a n t a t i o n ( R e i c h s t e i n , 1960b)  t—•  17 unlikely.  Research  physically  capable  indicates that of covering  of  V a r s h a v s k i i ' s (1937) marked  to  5000 m e t e r s ) ,  Rather  specimens  i t s migrations  (Regnier  and P u s s a r d ,  seasonal  i n nature  1926; B r i n k ,  due e i t h e r  at ploughing  1961;  Dub, 1 9 7 1 b ) o r t o c h a n g e s  two h a b i t s o f M i c r o t u s  storing  food  underground  no  by o t h e r  need In  to  distances (1%  travelled  2000  meters  a habit.  local  movements  1967).  These  tend  out of the (Cechowicz  i n food  Also  vole i s  t o very  and h a r v e s t i n g time  1955b).  great  t o make t h i s  to shifts  areas  faced  t h e common  relatively  i t does n o t appear  i trestricts  while  cultivated  and Pucek,  a v a i l a b i l i t y (Delost,  arvalis  - burrowing  - may m i t i g a t e t h e p r o b l e m s  a l p i n e mammals t o s u c h  t o be  an e x t e n t  that  and  of winter there i s  f o r migration. c o n c l u s i o n , t h e home r a n g e s o f M i c r o t u s  be r e s t r i c t e d  each  specimen  sent  i t s "true"  and m i g r a t i o n s  was t r a p p e d , altitude.  local.  arvalis  The a l t i t u d e  t h e r e f o r e , was c o n s i d e r e d  appear  at which to repre-  18 RESULTS  1.  Skull  1.1.  E i £d_S;t ruc_turejs a  r  Paired the  tooth  ured  structures  rows,  f o r each  To d e t e r m i n e  - were a l w a y s  whether  t h e r e was a n y  were measured  on b o t h  was r u n o n t h e d i f f e r e n c e  set of paired  structures.  of significance  meas-  There  sides  between t h e was no  differ-  (Table I I ) .  M e a s u r i n c j Consi_s;tencj£ The  author's measuring  sistency. times and  foramina, the diastemas,  and t h e jaws  25 s k u l l s  t-test  e n c e a t .001 l e v e l 1.2.  side.  difference,  a Student's  means  - the p a l a t a l  the mandibles  on t h e r i g h t  right-left and  Dimensions  Twenty-five  t e c h n i q u e was t e s t e d  intact  skulls  f o r 2 5 o f t h e 2/ d i m e n s i o n s .  mandibular  tionship  diastema  length  of other s p e c i f i e d  were each  for.its  measured  (As m a x i l l a r y  were based  dimensions,  molar  entirely  these  conthree width  on t h e  two were  rela-  not i n -  cluded) . The second  first  round  i n mid October  vember 1978. dimension  p l u s o r minus  coefficient  ly:  pages  son  of v a r i a b i l i t y  the  levels median  p l a c e i n June s  the standard error computed  The l a t t e r  among  computation  measurements w i t h  of variation  f o r each  was c a l c u l a t e d  (Appendix  ( D a v i e s and G o l d s m i t h , 1972). coefficient  1978; the  a t t h e b e g i n n i n g o f No-  o f the t h r e e measurements  of variation  82 t o 1 0 6 ) .  took  1978; t h e t h i r d  The a v e r a g e  the  mean  o f measuring  Tables  enabled  widely Table  f o r each  and  l a to compari-  differing I I I , based  o f t h e 25  skull  on  TABLE I I : LEFT-RIGHT SKULL DIFFERENCES IN MICROTUS ARVALIS NAME OF 7 MEASUREMENTS (LEFT VS. RIGHT)  MEAN (MM) OF MEASUREMENTS (N=25) * STANDARD ERROR  P a l a t a l foramen (L)  4.025 * .064  P a l a t a l foramen (R)  4.034 * .066  M a x i l l a r y diastema (L)  6.909 * .098  M a x i l l a r y d i a s t e m a (R)  6.898 * .097  M a x i l l a r y t o o t h row (L)  5.402 * .066  M a x i l l a r y t o o t h row (R)  5.417 i .066  Mandible length (L)  13.728 ± .154  M a n d i b l e l e n g t h (R)  13.703 i .155  Jaw l e n g t h ( L )  8.922 * .093  Jaw l e n g t h (R)  8.904 t .091  M a n d i b u l a r d i a s t e m a (L)  3.528 4 .049  M a n d i b u l a r d i a s t e m a (R)  3.479 ± .050  M a n d i b u l a r t o o t h row (L)  5.394 ± .071  M a n d i b u l a r t o o t h row (R)  5.425 ± .072  T-VALUE FOR DIFFERENCE BETWEEN THE MEANS  -.094  .076  155  ,110  .133  .670  -.003  20 TABLE I I I : CONSISTENCY OF MEASURING TECHNIQUE NAME OF MEASUREMENT  RANGE OF MEDIANS (MEANS  Condylobasal Total Basal  13.217 ^  Palatal  length  Basilar  length  Mandible  cv  23.617  0.5  5.400  0.5  <  width tooth  row l e n g t h  width  zygoma w i d t h  M a x i l l a r y diastema  ^=  x  dzz.  <^cv  length  Skull height Jaw  <  length  Mandibular  Post  x ^  0.0  width  Braincase  Mastoid  (CVS IN %)  length  length of skull length  Zygomatic  IN MM)  RANGE OF MEDIANS  10.167  1.0  0.833  1.0  length  Maxillary Palatal  tooth  foramen  M a x i l l a r y outer Least  length molar width  interorbital  Palatal bridge Rostrum  width  length ^—  width  Foramen magnum Nasal  row l e n g t h  x  ^—  ^  cv  height  width  Maxillary  i n n e r molar width  Foramen magnum Palatal  foramen  width width  5.467  11.0  /  21 dimensions,  summarizes  ly  (pages  82 t o 1 0 6 ) .  er  the dimension  the results  from  I t indicates  measured,  Appendix  that,  Tables  l a to  i n general, the larg-  t h e more c o n s i s t e n t t h e  measuring  technique. 1.3.  B r a c h y os_ti_sm Frank  tion,  called  tional  has d e s c r i b e d a homozygous r e c e s s i v e muta-  brachyostism,  variations  valis . edly  (1967)  causes  affected  h a d , among  and broadened  skull  was 6 8 % t o 7 3 % o f t h e c o n d y l o b a s a l it  propor-  other  f e a t u r e s , a mark-  i n which  the zygomatic  length.  ( I n normal  width  animals  i s 58% t o 62%.) The  specimens  brachyostism. tic  width  three  used  i n this  relationship  brachyostism,  fall  also  2.1.  Se><ual_Di m£r£hi sm A synopsis  within  that  the mutation  d i dthe  range.  As  the remaining  untested  speci-  L  of the l i t e r a t u r e i s presented  e x t e r n a l body  in skull  IV.  There  dimorphism  A slight  i s indicated,  a d u l t males. dimensions.  on s e x u a l  i n Table  measurements  s m a l l e r than  morphism  I n none o f t h e s e  zygoma-  Differences L  arvalis  f o r both  normal.  Sexual  crotus  were t h e r e f o r e t e s t e d f o r  had been t e s t e d and none had shown  i t was a s s u m e d  2.  being  length.  quarters of the t o t a l  mens w e r e  study  Of t h e 4 2 2 , 318 were m e a s u r a b l e  and c o n d y l o b a s a l  proportional  in  considerable  i n a l l parts of the skeleton of Microtus a r -  The a n i m a l s  shortened  which  with  i s less  i nMi-  dimorphism  adult  agreement  females on d i -  To q u a n t i f y p o s s i b l e d i m o r p h i s m  TABLE IV: LITERATURE SYNOPSIS OF SEXUAL DIMORPHI!iM IN MICROTUS ARVALIS DIMENSION  AGE GROUP  SEXUAL DIMORPHISM  8-30 days  Weight  No s i g n i f i c a n t d i f f e r e n c e ( D e l o s t ,  More t h a n 28 days  Weight  Female s m a l l e r ( R e i c h s t e i n , 1964)  "Adult voles"  Weight  Female s m a l l e r ( F r a n k and Zimmermann, 1957b)  0-15 days  Body  length  No s i g n i f i c a n t d i f f e r e n c e ( D e l o s t ,  More t h a n 15 days  Body  length  Female s m a l l e r ( D e l o s t , 1952; S p i t z ,  "Younger v o l e s "  Body  length  Female s m a l l e r (Cechowicz and Pucek, 1961)  45 days  Body  length  Female s m a l l e r ( S p i t z ,  60 days  Body  length  Female s m a l l e r ( D e l o s t , 1955b)  "Adult v o l e s "  Body  length  Female s m a l l e r ( F r a n k and Zimmermann, 1957b; S a i n t 1973)  "Younger v o l e s "  Tail  length  Female s m a l l e r (Cechowicz and Pucek, 1961)  "Younger v o l e s "  Condylobasal length  Female s m a l l e r (Cechowicz and Pucek, 1961)  "Adult v o l e s "  S k u l l dimensions  No s i g n i f i c a n t d i f f e r e n c e (Cechowicz and Pucek, 1961)  1952)  1952) 1963a)  1963a)  Girons,  23 among t h e  animals  temporarily vided  into  26  between  three  to  2k:  pages  107  to  the  reasons  differ  male  (61  versus males  43) he  other  hand  Stein  bryos  (560  versus  sex  ratio  males.  groups  and  the  animals  was  run  96  of  the  32  i s discussed  on  tests  were  were d i -  f o r each  specimens  the  and  488)  the  felt  the was  there  (Appendix  on  difference a  Tables  conflict sex  month o f of  number o f  ratio  one  this  i n the  i n Microtus  2a  hand  Delost  f e t u s e s and accounted  wild  (461  arvalis,  (1955c)  newborns  f o r the  versus  and  concluded resulted  i n the (Table ratio  VI  the  subadult subadult  and  i n an  literature V).  The  are  There  of  to and  the  females  be  a  a  fall  em-  coupled females. shifts  in this  subdivided into  appears  On  female  seasonal  - 230  a d u l t females  males.  excess  on  were  280).  factor  specimens  difference sexes  capture.  that this  found  predomi-  r e p o r t e d a g r e a t e r number o f  data  Table  sex  of embryos,  (1957)  the  In  predominance in  the  On  caught  show a s i g n i f i c a n t 137  groups  significance  c o m p e t i t i o n among m a l e s Similarly  the  on  for i t .  a g r e a t e r number  with  of  367  specimens  117).  that  nance of  age  t-test  one  of  unsexed  Ratio  Researchers and  level  55  aging.the  In not  .001  Sex  of  the  remaining  A Student's  means. at  the  segregated  method  32.)  the  2.2.  sex  (The  to  difference  study,  s e t a s i d e and  dimensions. pages  in this  in  study  versus age  summer increase  TABLE V: LITERATURE SYNOPSIS OF SEASONAL SEX RATIO IN MICROTUS ARVALIS SEASON  SPRING/ START SUMMER  SEX RATIO Equal numbers ( R e i c h s t e i n , 1956) More females ( S t e i n , 1953) More females (AdamczewskaAndrzejewska and Nabaglo, 1977) More a d u l t females than a d u l t males (Lapshov, L o r b e r and Polegaev, 197A) More males (Dub, 1971b)  NUMBERS  333 v s . 221 Rapid r e c r u i t m e n t o f f e m a l e s c o u p l e d w i t h s l o w e r r a t e o f d i s a p p e a r a n c e o f females 35 v s . 21  Better survival  53 v s . 20  Onset o f r e p r o d u c t i o n and i n c r e a s e d s e x u a l a c t i v i t y o f males  More males (Grunwald, 1975)  SUMMER  END SUMMER  AUTUMN/ START WINTER  EXPLANATION BY AUTHOR(S)  ability  E a r l i e r achievement o f s e x u a l m a t u r i t y o f males  More females ( S t e i n , 1953)  6A8 v s . A39  More females ( S t e i n , 1953) More females ( R e i c h s t e i n , 1956) E q u a l numbers o r s l i g h t l y more males (AdamczewskaAndrzejewska and Nabaglo, 1977)  201 v s . 129  More females (Lapshov, L o r b e r and Polegaev, 197A) More males (Dub, 1971b)  135 v s . 100  Better survival  158 v s . 82  M i g r a t i n g component o f p o p u l a t i o n  F i g h t i n g among males d u r i n g b r e e d i n g season S i m i l a r recruitment o f females coupled with g r e a t e r d i s a p p e a r a n c e o f females  ability  TABLE V I : AGE AND SEX ANALYSIS IN MICROTUS ARVALIS BY MONTH OF CAPTURE AGE AND SEX MONTH GROUP OF CAPTURE  JUVENILES  SUBADULTS  ADULTS  FEMALES MALES  FEMALES MALES  FEMALES MALE  FEMALES MALES  -  -  4  -  -  6  2  -  -  -  2  1  2  2  June  1  5  -  8  2  1  July  4  1  25  13  24  17  12  August  1  9  28  8  13  6  September  6  4  13  13  11  1  October  4  4  32  30  10  1  November  1  7  3  2  2  December  _  3  6  2  _  January  -  1  1  February  -  1  -  March  -  5  April  -  May  1  3  26 3.  Aging  3.1.  i_9i £ J.ec_hrii_qu_es_A£p_li ed  3.1.1.  Specimens  Specimens  n  Tables vailable of  the  wi_th_body_  V I I and  on  VIII  mean b o d y  larger  work  They  i n which  sizes  and  present  are  the  lived  longer than  (1964) a t t r i b u t e d  laboratory  animals  same m a n n e r a s  corresponding practically 3.1.1.1. This als  M.  VII body  21  which  arvalis  food  i n the i n the  from  compared  field  two  i n the  specimens  of  Microtus  attained  and  longevity  conditions  field.  and  animals  Therefore, different  direct  a-  conditions.  laboratory  these  wild  because  greater size  is restricted had  IX  not  females  males  day  lists  VIII.  of  freedom do  not  the  environments  manner.  80.71  mm  (79.9  grouping).  mm  average was  Based  five  this,  of  first  from  breed-  about  three  ( T a b l e XVIm;  page  mean b o d y  l e n g t h s from  Table  day  o l d m a l e s and  i n the  youngest  l e n g t h from  substituted on  age  individu-  weeks  35  classed body  the  immature  able to reproduce  laboratory and  to sexually  reached  about  o l d females  The  yet are  from  lengths of animals  Table  mm  be  category  Table of  groups  cannot  weeks o l d and 67).  the  and  literature  Juven_i_les  - those  ing.  age  of the  laboratory  under  to advantageous  predators, noting that  grow i n t h e  segregated  v o l e s i n the  Gebczynska  from  a summary  lengths i n laboratory  Microtus arvalis.  agrestis  _leng_th_mea_sur_ement_s  any  these  for Dottrens specimen  wild  mean  groupings means  in  was  s m a l l e r than  w i t h a body  80  length  TABLE V I I : LITERATURE SYNOPSIS OF BODY LENGTHS IN LABORATORY MICROTUS ARVALIS AGE (DAYS)  FEMALE MEAN LENGTHS (MM)  0-1 0-2  26-28 ( D e l o s t , 1952) 23-30 ( D e l o s t , 1955a)  14  59.0  ( S p i t z , 1963a)  73.6 81.7  ( S p i t z , 1963a) ( S p i t z , 1963a)  15 21 28 30 35 36 42 44 51 56 60 70 84 90 120 126  MALE MEAN LENGTHS (MM) 28-30 ( D e l o s t , 1952) 26-29 ( D e l o s t , 1955a) 61.4 ( S p i t z , 1963a) 67.1 ( M a r t i n e t , 1966) 75.2 84.6  ( S p i t z , 1963a) ( S p i t z , 1963a)  90.1 82.8  ( M a r t i n e t , 1966) ( M a r t i n e t , 1966)  94.0  ( S p i t z , 1963a)  84.1 87.6  ( M a r t i n e t , 1966) ( S p i t z , 1963a)  94.0  ( M a r t i n e t , 1966)  86.9 90.4 86 85.8  ( M a r t i n e t , 1966) ( S p i t z , 1963a) ( D e l o s t , 1952) ( M a r t i n e t , 1966)  97.3 96 100.2  ( S p i t z , 1963a) ( D e l o s t , 1952)' ( M a r t i n e t , 1966)  92.7  ( S p i t z , 1963a)  95.1 101.8  ( M a r t i n e t , 1966) ( S p i t z , 1963a)  ( M a r t i n e t , 1966) ( M a r t i n e t , 1966)  102.5 106.9  ( M a r t i n e t , 1966) ( M a r t i n e t , 1966)  ( S p i t z , 1963a)  106.6  ( S p i t z , 1963a)  114.5 111.1 110.4 112.0 116.6  ( M a r t i n e t , 1966) ( S p i t z , 1963a) ( M a r t i n e t , 1966) ( M a r t i n e t , 1966) ( M a r t i n e t , 1966)  114.2  ( M a r t i n e t , 1966)  119.8  ( M a r t i n e t , 1966)  102.4 97.9 98.3  150 180  100.3 102.5  ( M a r t i n e t , 1966) ( S p i t z , 1963a)  210 240 270 300 360 390  110.0  ( M a r t i n e t , 1966)  105.2  ( M a r t i n e t , 1966)  110.4 107.3  ( M a r t i n e t , 1966) ( M a r t i n e t , 1966)  TABLE V I I I : LITERATURE SYNOPSIS OF BODY LENGTHS IN WILD MICROTUS ARVALIS AGE GROUP  FEMALE MEAN LENGTHS (MM)  MALE MEAN LENGTHS (MM)  Young(May)  85.8 (Lapshov, L o r b e r and Polegaev, 1974) 78.2 (Lapshov, L o r b e r and Polegaev, 1974)  87.2 (Lapshov, L o r b e r and Polegaev, 1974) 79.3 (Lapshov, L o r b e r and Polegaev, 1974)  Young(November). Juvenile Juvenile  Semiadult(May) Semiadult(November)  78.0 (Cechowicz and Pucek. 1961) S m a l l e r t h a n 80 ( D o t t r e n s , 1962)  94.1 (Lapshov, L o r b e r and Polegaev, 1974) 95.1 (Lapshov, L o r b e r and Polegaev, 1974)  91.4 (Lapshov, L o r b e r and Polegaev, 1974) 93.9 (Lapshov, L o r b e r and Polegaev, 1974)  Immature(750m) Immature(1360m) Immature(1940m) Subadult  Adult(May) Adult(November) Adult Adult Mature(750m) Mature(1360m) Mature(1940m)  ? MEAN LENGTHS (MM)  81.9 ( J o n e s , 1970) 80.4 ( J o n e s , 1970) 81.4 ( J o n e s , 1970) 80-99.9 ( D o t t r e n s , 1962)  107.4 (Lapshov, L o r b e r and Polegaev, 1974) 104.5 (Lapshov, L o r b e r and Polegaev, 1974)  109.9 (Lapshov, L o r b e r and Polegaev, 1974) 107.0 (Lapshov, L o r b e r and Polegaev, 1974) 105.1 (Cechowicz and Pucek, 1961) 100+ ( D o t t r e n s , 1962) 94.4 (Jones. 1970) 92.7 ( J o n e s , 1970) 101.1 (Jones, 1970)  CO  TABLE I X : BODY LENGTHS FROM TABLES V I I AND V I I I OF JUVENILE MICROTUS ARVALIS SEX AND AGE GROUP  LABORATORY OR WILD SPECIMENS  MEAN BODY LENGTHS (MM)  Females (21 days o l d )  L a b o r a t o r y specimens  73.6 ( S p i t z , 1963a)  Females (young)  W i l d specimens  78.2 t o 85.8 (Lapshov, L o r b e r and P o l e g a e v , 1974)  Both sexes ( j u v e n i l e )  W i l d specimens  78.0 (Cechowicz and Pucek, 1961)  Both s e x e s ( j u v e n i l e )  W i l d specimens  S m a l l e r t h a n 80.0 D o t t r e n s , 1962)  Both s e x e s (immature)  W i l d specimens  80.4 t o 81.9 ( J o n e s , 1970)  Males (35 days o l d )  L a b o r a t o r y specimens  82.8 ( M a r t i n e t , 1966)  Males (young)  W i l d specimens  79.3 t o 87.2 (Lapshov, L o r b e r and Polegaev, 1974)  30 smaller  than  80 mm  mens w i t h b o d y  was  l e n g t h measurements which  juveniles,  body  dard  o f 1.277  error  3.1.1.2.  time  fect  l e n g t h averaged mm  category  comprises  of capture  they  had  had r e a c h e d  levelled  their  The  e s t i m a t e s range  from  ( R e g n i e r and  mann, 1 9 5 7 b ) . which  each  voles  differ  Pussard,  animal  i s born.  from  average  those body  i f i t were  spring three six  1 0 0 mm  animal  of birth might  adopted,  to achieve  both  would  100+  arvalis was mm  of the adults  have reached born  t h e same s i z e ,  and  Zimmer-  born  (Table X). specimens  l e n g t h was  was  in common  1 0 2 . 4 6 mm.  a specimen  In  months  infe-  considered  d i d not matter.  animal  might  mark have  but i n the aging  be c l a s s i f i e d , c o r r e c t l y ,  lab-  reached  By u s i n g t h i s  t h e 1 0 0 mm  and 1/2  by t h e s e a s o n  o r more i n l e n g t h .  months, w h e r e a s an autumn  months  (Frank  individuals  (Table VIII)  therefore,  (Meunier  of spring  M.  are  arvalis.  1955b) t o 2  i n m a l e s and t h r e e t o f i v e  study,  the season born  Growth r a t e s  (Table VII) this  t h r e e months In t h i s  proach,  months  by  in ef-  Researchers  1926) t o 1 y e a r  length for wild  specimens  adult  stan-  rate  that  i n Microtus  1952 and  born  growth  an e x t e n t  size.  occurs  o f autumn  oratory  males.  whose  This i s complicated further  as mature o r a d u l t  about  animals  adult  (Delost,  classed  at  as  p l u s or minus a  t h e a g e o f 1 1/2  1972) t o 2 months  The  speci-  3e: page 1 2 2 ) .  t o such  full  i n a g r e e m e n t a s t o when t h i s  months  Table  those  not  Solari,  were c l a s s i f i e d  7 3 . 2 4 2 mm  (Appendix  F o r t h e 33  Adults_  This the  considered a juvenile.  an  apA  i n only taken system  as a d u l t s .  For  TABLE X: LITERATURE SYNOPSIS OF DATE OF BIRTH AND GROWTH RATE OF MICROTUS ARVALIS DATE OF BIRTH  GROWTH RATE  REFERENCE  January-April  Rapid growth r a t e  Martinet  February-June  Very q u i c k growth  S a i n t G i r o n s (1973)  March-May  F a s t growth r a t e u n t i l 3 t o A months o f age  M a r t i n e t and S p i t z (1971)  March-June  Quick growth r a t e  Spitz  March-June  Rapid i n c r e a s e i n weight up t o 47 g r  Reichstein  Summer  Slower growth r a t e  S a i n t G i r o n s (1973)  June-August  Intermediate  M a r t i n e t and S p i t z (1971)  June-October  Increase i n weight up t o 15 t o 22 g r ; remain u n t i l January/February a t these low r a t e s  Reichstein  July-August  Slow growth r a t e  Martinet  July-October  Slow growth r a t e  Spitz  Autumn  Slower growth r a t e  S a i n t G i r o n s (1973)  Autumn  Growth stopped d u r i n g w i n t e r a t 18 t o 20 g r ; i n s p r i n g r e a c h a d u l t weight  Wijngaarden (1960)  A f t e r second h a l f o f August  Cease growth, r e a c h i n g  S t r a k a and Gerasimov  September-October  Low p l a t e a u when 1 t o 2 months o l d ; grow a g a i n i n January  growth r a t e  stage o f s u b a d u l t  (1967)  (1967) (1964)  (1964)  (1967)  (1967)  (1971)  M a r t i n e t and S p i t z (1971)  32 the  69  specimens  sified a  as  adults,  standard  3.1.1.3.  error  (80  100  These  mm).  s t i l l  of  mm  be  The  average  or  semiadult  length mm  animals  in a period length  80  with  VIII)  mm  to  body  99.9  tal  length  of  3 . 9 2 5 mm  juveniles,  179  no  measurements  length  of  skull  body was  width.  gainst  the  of  range  specimens  belonged  to  an  which  clas-  or  page  minus  122).  altitude  but  significant  mm  8 9 . 4 5 3 mm Table  as  was  For  subadult  179  were  3e;  they  substituted  the  plus  than  growth.  classed  (89.9  than  (smaller  breeding  arvalis  (Appendix  larger  adults  of  grouping).  classified or  minus  page  a  122).  bod_y_le_ng_th mejaS_Jrem_en_ts 281  had  and  been group  length  adults. data,  at  length  result  aged  one  of  body  approximated  in  length  most  to-  length  compared  same d i m e n s i o n to  five  and/or  basilar  was  -  remaining  length  and/or  according  which  the  least  obtained  f o r the  measurements  For  - condylobasal  basal  The  body  69  length  measurement had  M.  9 2 . 8 9 mm  taken  and/or  zygomatic  capable  averaged  subadults  specimens with  than  measurements which  specimens,  and/or  281  3e:  were  plus  specimens  relatively  mm  length  body  422  of  was  Spec_ime_n_s wi_thout. the  be  for wild  3.1.2.  cranial  Table  includes smaller  would  error  141  which  1 0 5 . 4 6 4 mm  (Appendix  category  standard  33  measurements  averaged  more) and  (Table  subadults,  Of  length  1.202  or  body  Dottrens'  specimens as  body  intermediate  juveniles  for  body  Subadu_lt_s  This  would  with  athe and  closely  33 the  one from  came. 422  which  Table  XI l i s t s  specimens.  because and  the animal  the complete  I t includes  external  without  cranial  length  measurements  s e x and age a n a l y s i s  33 a n i m a l s  measurements  none o f t h e f i v e  body  that  could  n o t be a g e d  had n o t been p r o v i d e d measurements c o u l d  of the  f o r them  be  p e r f o r m e d on  them. T A B L E X I : AGE SEX  AND  S E X A N A L Y S I S OF  (AND NUMBER)  JUVENILES  MICROTUS A R V A L I S  SPECIMENS  SUBADULTS  ADULTS  ?  Females  (230)  17  128  72  13  Males  (137)  18  81  31  7  (55)  14  17  11  13  (422)  49  226  114  33  Total 3.2.  Chancjes Of  i n mean  classes  en d i m e n s i o n s only age  classes  were  length,  length,  width  ed  to 123). in this  with  outer  rnolar  palatal  A comparison  molar  classes.  The  change  width,  molar  dia-  width,  f o r a m e n magnum h e i g h t ,  (Appendix  least  Tables  between average  f o r some o f t h e c r a n i a l  elev-  three  mandibular  inner  and  between  c h a n g e among a l l  maxillary  width,  change ( a t  the juvenile  a significant  foramen width,  length.  both  and a d u l t  o r no s i g n i f i c a n t  zygoma w i d t h ,  foot  a significant  age between  showed  maxillary  post  study  21 showed  either  maxillary  and h i n d  Dimensions  and t h e s u b a d u l t  which  men magnum w i d t h ,  118  size  two age c l a s s e s  stema ear  Individual  t h e 32 d i m e n s i o n s ,  .001 l e v e l ) subadult  in  fora-  interorbital 3a t o 3 f : pages  measurements  dimensions  record-  and d a t a  34 from  the  XII. and  literature  I t s h o w s how how  little  3.3.  with  age  arvalis trens  dimensions  single  cranial  appears  in  dimensions  Table  can  be  there i s .  specimens  used  in this  found  ble  4:  page  age  i n M.  that  In  they  4•  the  The altitude  at  shown  The  age  i n the  four  significant  of  M.  sets  skull  Microtus that  Dot-  and  one  changes  change  nivalis  sets,  the  ( i n one  nivalis  coefficient was  f o r each  of  pages  - the  certain  (Appendix  Ta-  in three  of  with the  p r o p o r t i o n a l trends set  significantly)  (Table  XIII).  Size Correlation  capture  correlation  examined  trends  in proportional relationship  two  with  several  between  significant  those  i n M.  and  study  no  remaining  decreased  were  most  trends  matched  correlation  groups  2k:  had  increased  Altitude  these  shown t h e  124).  arvalis  whereas  had  arvalis  sets.  nivalis  proportional relationship of  reported  M.  in Microtus  sets  he  five  (1962) noted  Five  set  to  variaole  accord  i n the  dimensions.  age  equivalent  Chan£es_ in_Pro£or_tion_a_i R e l a t i o n s h i p s Dottrens  in  on  between  calculated the  even  to  the  107  to  117).  32  third  size  f o r the  dimensions. decimal  place  of  specimen  and  three  sex  segregated  I n no  case  was  (Appendix  there  Tables  2a  TABLE XII: COMPARISON OF CRANIAL MEASUREMENTS IN MICROTUS ARVALIS AGE CLASSES NAME OF CRANIAL MEASUREMENT  REFERENCE  AGE GROUPS MEAN (MM) i STANDARD ERROR (WHEN GIVEN) JUVENILES SUBADULTS ADULTS  Cechowicz and Pucek (1961) Dottrens This  (1962)  22.25 Condylobasal length  thesis  Dottrens  (1962)  M a x i l l a r y diastema length  6.1  (1962)  zygomatic w i d t h  11.9  This  (1962)  4  .10  22.189 ± .718 7;i  * .04  6.587 i .274  S k u l l height  6.9  Maxillary tooth row l e n g t h  thesis  ^-Presumably a l l D o t t r e n s ' a d d i t i o n s a r e s t a n d a r d  ± .09  * .07  * .16  23.939 * 7.7  .084  4 .07  7.285 ± .425  13.6  4  .04  12.694 4 .496  7.3  ± .05  14.6  4  .09  13.845 * .599  7.6  * .08  6.214 * .057  6.634 4 .260  7.028 4  .387  5.5  5.9  6.2  .04  4  .05  5.081 * .042 errors  25.1  13.69  11.570 4 .097  This t h e s i s  Dottrens  23.2  12.54  This t h e s i s  Dottrens  4 .07  5.904 4 .062  Cechowicz and Pucek (1961) (1962)  * .13*  20.346 4 .165  This t h e s i s  Dottrens  20.5  24.24  4  .03  5.396 4 .191  4  5.640 i .267  TABLE X I I I : PROPORTIONAL RELATIONSHIPS IN SKULL DIMENSIONS IN TWO SPECIES OF MICROTUS MICROTUS NIVALIS MEAN ( I N MM) ± STANDARD ERROR ( I N %) T-VALUE JUVENILES ADULTS DIFFERENCE (NUMBER) (NUMBER) (AND DF)  TRENDS WITH AGE  SETS OF CRANIAL DIMENSIONS  110.7 ± 1.2 (15)  122.0 4 1.0 (57)  9.19* (70)  (  68.6 i 0.6 (15)  63.1 i 0.5 (57)  -6.32* (70)  1  Interorbital ww ii dd tt hh Maxillary t o o t h row  I |  Interorbital width Jaw l e n g t h  I j  S k u l l height Zygomatic width  29.6 * 0.3 (15)  25.6 4 0.2 (57)  -6.90* (70)  59.4 4 1.0 (15)  54.5 4- 0.4 (57)  -5.33* (70)  58.9 4 0.5 (15)  57.0 * 0.4 (57)  -2.47 (70)  * Significant  a t .001 l e v e l  Maxillary diastema Maxillary t o o t h row  A  X1 I L O I U J - U J . u u ±  I I ^  Zygomatic width Condylobasal length  TRENDS WITH AGE  A T I , I  MICROTUS ARVALIS MEAN ( I N MM) i STANDARD ERROR ( I N %) T-VALUE JUVENILES ADULTS DIFFERENCE (NUMBER) (NUMBER) (AND DF) 116.8 i 1.1 (48)  128.2 i 14.8 (112)  6.48* (158)  61.9 * 1.5 (48)  55.6 * 6.6 (112)  -5.12* (158)  37.2 * 0.9 (44) .  33.7 * 3.9 (110)  -3.54* (152)  42.2 ± 3.5 (42)  47.5 4 12.8 (105)  2.82 (145)  52.7 4-2.4 (40)  55.1 4 1.3 (100)  4.07* (138)  1 | I »  ~~ A T  37 DISCUSSION  This  section  "ecogeographical second  was p r o m p t e d  two m a j o r  arvalis trends found  rule"  into  (Mayr,  papers  in this  size thesis  - the f i r s t  of subspecies.  with  altitude  described  1970).  i n each  i n Table  drawn i n  i n Microtus  1962; Jones,  a r e summarized  This  conclusions  on a l t i t u d i n a l v a r i a t i o n (Dottrens,  on t h e  105) o f Bergmann, t h e  by t h e q u e s t i o n a b l e  i n Switzerland i n body  two p a r t s  1956:page  on t h e i n f r a s p e c i f i c t a x o n  approach the  i s divided  The  paper and  XIV and F i g u r e  7.  T A B L E X I V : TRENDS I N BODY LENGTH WITH A L T I T U D E I N MICROTUS A R V A L I S (DOTTRENS, 1 9 6 2 ; JONES, 1 9 7 0 ; THIS T H E S I S ) REFERENCE  BODY LENGTH TRENDS WITH  Dottrens (1962)  Lowest Altitude Group  Mid Altitude Group  Jones (1970)  Lowest  Mid Altitude Group  This thesis  Lowest Altitude Group  Altitude Group  Dottrens tude  group  valis  concluded  belonged  (Pallas,  1779) whereas  belonged  fuscus  ( S c h i n z , . 1845).  his results  that  subspecies  =;  called  He s t a t e d  that  Highest Altitude Group Highest Altitude Group Highest Altitude Group  the lowest  alti-  M. a r v a l i s a r -  t h e mid and h i g h e s t  to a subspecies  an a l t i t u d i n a l c l i n e  > <  Mid Altitude Group  =  t o the nominate  groups  followed  from  ALTITUDE  M. a r v a l i s  altitude rufescente-  i f t h e v a r i a t i o n had  (Bergmann's R u l e ) ,  the specimens  38  F I G U R E 7: N O N - J U V E N I L E BODY LENGTHS I N MICROTUS A R V A L I S (DOTTRENS, 1 9 6 2 ; JONES, 1 9 7 0 ; THIS T H E S I S ) MM 108 106  90 L  M  ALTITUDE —  = DOTTRENS  GROUPS (LOW,  (1962)  H MEDIUM,  HIGH)  = JONES ( 1 9 7 0 ) —  = THIS  THESIS  39 in  h i s 1200  405  m to  m specimens  however, ferent  the  and  mid  from  t h a t the  tude  were e x p o s e d  and  those  i n the  Dottrens Jones  recorded  and  a necessity to  to  an  1200  be He  was  the  head  i n 1970  altitude 37  and  were  the  m  those  altiat  variance  described  XVII:  disparities.  On  and  supported  at  significantly  groups  Table  demands o f p l u s body  application  to  to  1500  c l i m a t e than  dif-  hypothe-  due  m to  a  analysed  page the  74).  of  the  size  at  the  he  were  hypothesis  mountain  Yet  contrary,  Dottrens' the  from,  by  a___ r u f e s c e n t e f u s c u s " w h o s e g r e a t  Dottrens  and  subspeciation to  flicting  the  to  of  a  size  is  environ-  higher elevations  Bergmann's R u l e "  (Jones,  62).  Since and  larger  point to  1970:page  meet  to  his If,  according  subspecies  at  Jones  his trend M.  have  variation  l e n g t h measurements  that  belonged  subspecies.  lengths differed  these  than  larger.  by  page  addressed  montane s u b s p e c i e s  may  montane  corresponding  h i s body  ment...the  of  a more m o d e r a t e  XIV;  been b i g g e r  groups  nominate  absence  t h e r e f o r e were  (Table  compatible  altitude  the  the  h i s mean b o d y  never  claimed  to  have  h i s 2000+ m s p e c i m e n s .  populations living  trends  with,  would  measurements would  of  within  that the  The  their  continued  fact  and  highest  those  Bergmann's Rule  2 0 0 0 + m,  m group  s m a l l e r than  and  subspecies  separately sized  1500  data,  the  two  Jones  both  justify concepts  r e s o r t e d t o Bergmann's  identical are  c o n c l u s i o n s from  examined  here.  Rule con-  40 1.  Be r g m a n n ' s • R u l e  1.1.  G e n e r a l . Re _view_o£ Ber_gmannj_s_Rule_ Bergmann's Rule  scribe  parallelisms  i s one o f t h e " r u l e s " between m o r p h o l o g i c a l  ogeographic  features  (Mayr,  latitudinal  increase  i n average  (Keeton, (Mayr,  with  there  i s a  increasing cold  increase with  altitude  T h e r e . i s _ n o ag_reeme_nt_ on_whejthe_r_Be_rg_mann'_s R u _ l e _ i _ s t_o_homeot_he r m s _ o r  Some r e s e a r c h e r s Schmidt,  Schmidt,  (Dale,  1962; Keeton,  1967; Brown  homeotherms.  Others  i s considered  1.1.2.  Park,  Emerson,  1957; Ray, 1960; Case,  t h e homeothermic  1976) As  races  and r a c e s ,  by m o d e r n  recognized  ically,  race  1963).  The v a l i d i t y  continues  con-  Microtus  application  of  i n this discussion.  (1847) and o t h e r  as " p o p u l a t i o n s  formally  to  wh_ich_  appli.e_s.  to species  considered  and  Bergmann's Rule  f o r p o i k i l o t h e r m s as w e l l . only  Park  re  Bergmann  defined  1967; A l l e e ,  1956 and 1 9 6 3 ; A l l e e  T_h£ _ i.s_no_ a_gj;ee_me_n t. on_the_ta_x£n£mi_c_leve_l_a;t  Ber_gmarin_|_s_Rule_  equally  1940; Mayr,  (Ruibal,  i s a homeotherm,  Rule  incj_ude_s p_o_iki_ loth_erms_.  and L e e , 1969) r e s t r i c t  i t has a p p l i c a t i o n  arvalis the  size  and p h y s i -  1963).  re_strixt_ed  tend  variation  I t states that  body  1967) and a c o r r e s p o n d i n g  1.1.1.  and  1956).  i n b i o l o g y which de-  early  authors  b u t most o f t h e i r  authors  (Mayr,  or aggregates  subspecies"  was p r o b a b l y  species are  1956) - r a c e s  of populations  (Mayr,  1963;page  synonymous w i t h  o f Bergmann's Rule  t o be d i s c u s s e d  a p p l i e d the Rule  within  350). Histor-  subspecies  (Mayr,  i n homeotherms  at various taxonomic  being  levels  by  41 modern 1940; or  day b i o l o g i s t s : Cowles,  among  (Allee  1945);  races  within  closely  related  species (that  1962; Brown  and s u b s p e c i e s  species  i s , among  o r demes) ( P a r k , 1949; Mayr,  and Schmidt,  species  among  1956);  (Dale, subspecies  or both  and L e e , 1 9 6 9 ) . ( D e f i n i t i o n s  appear  i n the discussion  agreement  on_th_e_validity_  of  on s u b s p e c i a -  tion. ) 1.1.3.  There  is_no  of_the_major  p h y s i £ l £ g i c a l _ i n t e r p r e t a t i o n o.f_Ber£mann ' s^ R u ^ e ^ As stant  most h o m e o t h e r m i c  internal  physiological that  large  cause The  body  mension square  animals  producing  volume  interpretation a relative  is  more n e a r l y  energy  and S c h m i d t ,  t o body  1962; Mayr,  lose other face  McNab  (1971)  more h e a t factors areas  alluded  stated  v i a their being  than  that  1971).  of linear d i -  1949; A l l e e  of this  and  physiological  selective  advantage  the metabolic t o body  rate  weight  1963). are not convinced  by t h i s  large  of a species  simply  individuals  inadequacy"  argu-  small individuals ( a l l  because  small individuals.  to the "hopeless  (McNab,  s u r f a c e than  s u r f a c e than  equal)  the usual  i n homeotherms i s  1945; Park,  i s a further  Some r e s e a r c h e r s , h o w e v e r , ment.  ratio  reduction of the surface since proportional  con-  f o r t h e r m o r e g u l a t i o n be-  Proponents  there  of a  s u r f a c e i n c r e a s e s as t h e  (Cowles,  1963).  add t h a t  Rule  i n c r e a s e s as t h e cube  dimension  1962; Mayr,  in  less  the heat-radiating  of linear  (Allee  expend  maintenance  (Scholander, 1955),  small surface-to-volume  whereas  Schmidt,  temperature  require  e x p l a n a t i o n o f Bergmann's  of their  heat  animals  they  Scholander  have  larger  (1955:page  sur22)  of the surface-to-volume  ratio "the  reduction size  of the surface  importance surface factor Zeuthen  1.1.4.  at best...no  area,  absolute  i n arctic  size  declaring that  area  or relative,  that  i s o f minor  that  the size  i s i n a n y way a  phylogenetic  heat  of the  critical  adaptation".  While  the metabolic  rate  decreased  he s u g g e s t e d  that  t h e d e c r e a s e had  of animals,  t o do w i t h  i n conserving  of the animal  t a n g i b l e evidence  or northern  (1947) agreed  increasing nothing  hypothesis,  with  heat r e g u l a t i o n .  _There a r e s e v e r a _ l met_hods o f - t h e r m o r e g u l a t i o n - o t h e r -  t h a r i £U£f£C£-£0_3_v£lume_  ra_ti£ _red_u£t_ion j_n_h_ig_h _ l a _ t i t u _ d e _ o _ r £ l -  t_i_tud_e_h£m£o_the r m s . Table homeotherms  XV l i s t s regulate  some o f t h e w a y s i n w h i c h body  heat  production  a l p i n e and  and heat  loss  arctic (Hoff-  mann, 1 9 7 4 )  T A B L E XV; SOME METHODS OF A.  CONTROL OF HEAT L O S S ( I N S U L A T I O N ) 1. F u r o r f e a t h e r t h i c k n e s s 2. C o n t r o l o f p e r i p h e r a l c i r c u l a t i o n 3. E v a p o r a t i v e c o o l i n g  B.  CONTROL OF HEAT 1. 2.  C.  1.1.5.  Heat Heat  POSTURE  from from AND  PRODUCTION  increased metabolic rate basal metabolic rate BEHAVIOUR  J_here a r e d_iet_a£d_C£m£e_ti_to_r _1 inked_a_rgum£n_ts_t_ha_t  may_ £X£l_airi £e_rtai£ c a s e s McNab with  THERMOREGULATION  (1971)  latitude  £ f _ B £ r £ m a n n '_s  speculated  i n carnivorous  that  RujLe.  a correlation  mammals  may  reflect  o f body the size  size of  the  available  bution which  prey,  o f the prey they  must  tween body reflect  which  share  the prey  l e n g t h and l a t i t u d e seed  and t h a t  - a r e marked  gests  that  available  each  selection  food  sharing a limited  is  a special  case  similar  latitudes,  due m a i n l y  food  food  islands,  a n d moun-  which  sug-  an i n c r e a s e d s i z e  t o the reduced  r e s o u r c e . Bergmann's  of this  p a t t e r n s be-  of species,  trends reflects  particles  of  of other predators with  and o c c a s i o n a l  by a l o w d i v e r s i t y  o f these  by t h e d i s t r i -  i n g r a n i v o r o u s h e r b i v o r e s may  T h r e e r e g i o n s -" h i g h  tains  i s influenced  and t h e d i s t r i b u t i o n  differential  scarcity.  i n turn  g e n e r a l phenomenon"  of  necessity  Rule  usually  (McNab, 1 9 7 1 :  page 8 5 2 ) . 1.1.6.  J_here  tinued  i_s_n£t_S£f_fic_ie_n_t e v_ide_nce_t£ wa_rran_t  t_he c o n -  ap£licat_ion £f_th_e_te_rm __^r_ul_e"_t_o _the _ s i _ z e _ t r e n d d e -  OCjrijDed^ b_y_Ber£ma_nn. According mann's R u l e logical  of  "does  n o t depend  only  i fi ti s really  s p e c i e s of warmblooded of size  species"  (Mayr,  validity  o f Bergmann's  studied  enough  cases  mann's R u l e  as a  on t h e v a l i d i t y  true  1963;page  that  Rule  i n more t h a n  portions  By M a y r ' s  collapses.  rule.  i s an  50  percent  average of the  standards, the  Variation  species studied,  (or  Rule " i s  o f t h e range  s p e c i e s o f homeotherms  of those  of the physio-  Bergmann's  vertebrates there  319).  of Berg-  on t h e r e l i a b i l i t y  finding."  i n the cooler  i n enough  105) t h e v a l i d i t y  but merely  of the empirical  increase  in  (1956:page  interpretation,  regularity) valid  t o Mayr  has n o t been  n o r has i t been to qualify  found  Berg-  44 1.2.  Ber_gmannJ_s_Rule and M i c r o t u s _ a _ r v a l j L s According  "increase Bergmann Thus a  n  of  Rule,  i n the  the  any  of  arvalis)  variation lection affect  may  are  the  oe  Switzerland and  variation  would thus  no  skull  dimensions.  which of  M^  i n the  produce  for this  that  results the  common v o l e  to  normal  arvalis  undermine  i n the  variation  assertion  than  altitude  lack  of  to  in  altitudinal  and  of  biotic  events  so  study  in  those  that  in  arvalis  thorough  Alps  the  zone.  with  course  similar  i n the  zone  abiotic  any  with  glareolus nageri  of Microtus  i n data  the  alpine  alpine  showed  deficiencies  growth  i n the  s e v e r a l reasons  pressures,  i n the  (Evotomys  specimens  603)  i n accordance  phenomenon  larger  422  study  - from  the  thesis  Yet  (1962:page  c o l d e r zones,  familiar  e x t e r n a l and  There  Schmidt  A l p s . . . t h e meadow m i c e  in this  32  and  i n the  is a  valleys".  examined  Allee  size  Microtus  a  to  of  se-  this  altitudinal  is a  familiar  the  altitude  phenomenon. 1.2.1.  2 £iciencies_in  1.2.1.1.  I n a c c u r_a_t e _ a . l t i. t u de  As by  the  the  noted  altitude  also  tain south  i n the  collector  justment is  data_  e  at  was an  i n the  the  Swiss  site  the  of  specimen  source  A l p s , the  differ this  by  as  section, capture  f o r home r a n g e  important  meters  methodology  which  made  aspects  by '1500  at  l o c a l i t i es__  of  or  of  was  M.  taken  migration. On  vegetation limits  north-south  300  slope  represent  arva1is lived.  inaccuracy.  much a s  to  recorded  Slope, an  No  ad-  however,  average  moun-  on  north  and  to  start  with;  difference  i s up  to  meters  500  45 meters  (Huxley, 1967).  vorous  (Table XVIg:  ference. trapped  page  specified  on a s o u t h e r n o r n o r t h e r n  whether  t o Green  are  little  so,  t h e 27 c r a n i a l  affected  would  (1932)  and m a n d i b u l a r  n o t be s u i t a b l e  Green's  statement  as s k u l l  measurements  influences. dimensions  indices  B u t many m a m m a l o g i s t s  1940;  Snyder,  1954; D o t t r e n s , 1962; Rees,  and Hoffmann,  External variation  i n mammals  section,  1.2.1.3.  cycles  1970; Jones,  (W.B. D a v i s , 1 9 3 8 ; D a l e ,  have  1970).  dimensions  f o rprecise  and p o s s i b l e  arvalis,  i n which  been  1974).  like  there  (Table XVIh:  phenomenon  Myers,  with  1969; Claude, 1970;  used  t o gauge  size  As n o t e d i n this  analysis  i n t h e meth-  study  may n o t  d u e t o t h e num-  techniques involved.  affected  and  The mea-  by t h e f o l l o w i n g .  Pop ul. a t i o n_cy_c 1_ i n g _  Microtus  a decline  may  altitudinal  ( D a l e , 1940; D o t t r e n s , 1962; Delany  a c c u r a t e enough  also  i n this  1975).  the external  of collectors  surements  variation  measurements a r e a l s o  1964; Claude,  have been  this  body  an  were  measurements c o n t i n u e t o f e a t u r e i n  on g e o g r a p h i c  Robinson  measured  a p p a r e n t l y do n o t a g r e e  studies  tion  was  i n mammals  If this  f o r marking  their  ber  the animal  dif-  slope.  skull  by e x t r a n e o u s  cline.  odology  by t h i s  herbi-  Ques t _ i o na b _1 e_meas u rem en t_s  According  Healy,  i s primarily  6 1 ) , i t may b e a f f e c t e d  Y e t no c o l l e c t o r  1.2.1.2.  study  As M i c r o t u s a r v a l i s  page  i n small During  many m i c r o t i n e s ,  undergoes  i s a population build 62).  peak  up, a peak and  (A c o m p r e h e n s i v e  mammals c a n be f o u n d population years  popula-  review of  i n K r e b s and  i n M.  arvalis,  46 Stein  (1957) documented  Zimmermann adults.  adult  (1955) r e p o r t e d  The  change  microtine  cycle  uals,  by  not  an  an  in adult  i s caused  changes  increase body  by  i n age  increase  i n body  i n mandible  size  which  differing  and  length  among  accompanies  growth  s t r u c t u r e of  weight,  the  rates of  the  individ-  population  (Krebs,  1964) . Since over  the  a period  museum s p e c i m e n s of  specimens were been  40  taken  l a r g e r than  the  adults  and  most  from  of  the  times,  the  appear  similar  years at  in this  i t i s probable  normal.  This  were c o l l e c t e d a t  adults  1000  latter  group,  in size  being  to  the  meters  pressures  1.2.2.1.  Center_o_f  distr_ibut_ion_  representatives  doubtless  (Allen,  1906:page  Rodentia  s a i d to  thew,  also  378). be  of  1915), and  to  Microtus northern  subfamily  c e n t r a l Asian  genus M i c r o t u s (Dale,  the  whose 1940).  of  to  non  usual,  have  most  peak  of  phases  would  increase  center  arvalis  which  of  belongs  (Allen northern  Microtinae  origin  may  chance  distribution,  of  thus  adult  c o l l e c t e d at  species  origin be  the  peak  in  fact  group.  i t s original  Cricetidae considered  northern  origin  a given  i t s hypothetical center  most c a s e s  family  "of  were  l a r g e r than former  and  i f by  2500 m e t e r s from  some o f  times  means t h a t  Sej^ectj.o_n  toward  that  peak p o p u l a t i o n  1.2.2.  The  t h e s i s were c o l l e c t e d  (Hinton,  Holarctic distribution  to  origin  1962),  size  is in  dispersal"  1906),  described  in  the to  order the  (Mat-  to  be  of  and  to  the  indicates a  Boreal  47 Although ter  of  the  northern  If  origin  Allen's  encing tor  opinions and  were  d i s p e r s a l of  o r i g i n s of  i n body  broader  study  s i z e would to  restricted  a  hypothetical  arvalis,  origin  family  a better  band  and  of  order.  were  influ-  approach  rather  cen-  share  to  latitudinal  distances  narrow  i t may  hypothesis  have been a  south  to  the  subfamily,  common v o l e ,  north  on  Microtus  (1906) s i z e - c e n t e r of  changes  tudinal  found  i t s genus,  v a r i a t i o n i n the  involving  not  moni-  study  than  an  alti-  latitude.  1.2.2.2. _n£luence £f_c_ima_e_ Studies  on  latitudinal  ally  attempt  to  c o r r e l a t e the  such  as  frost  the  free  weather  mean o f days.  and  There  are  s i z e v a r i a t i o n usu-  measurements w i t h  temperature  These data  stations.  altitudinal  or  snowfall  or  climatic rainfall  are  usually  supplied  by  two  deficiencies inherent  data or  local in  this  approach. The exact are  first  locality  forced  nearest cond  or  to  and  surface,  or  from use  the  of  which  that  live  from  on  the  official  placed  which  restricted  to  the  ground  stations.  a t . two  thus  e l i m i n a t i n g the  chance  ed.  Then t h e  and  nature on  meters  the  i t , no  ( R u i b a l , 1957). to  are  for climate  the  influences state longer  of  level of  the  the  play  se-  growing  proximity cannot  of  the  be  de-  by  the  instrument of  the  shel-  ground,  position select-  ground, a  The  published  Meteorological above  the  young  exposed,  the  researchers  a v a i l a b l e from  climate  figures  are  growing  of  the  ters  vegetation  locality  are  station in  s p e c i m e n comes and  information  conditions that  i s r a r e l y a weather  each  climatic  animals  directly  network  there  most e q u i v a l e n t  i s that  plants  duced  i s that  and  significant  the role.  48 The  meterological  surrounding enced  by a p e r s o n  sometimes about  area,  s t a t i o n i s thus  representative  and t h e c l i m a t e  thus  walking  termed.  two m e t e r s  upright,  (Geiger,  The  official  meteorological  gross  climatic  not  be e v a l u a t e d .  attempt vole, by  t o gauge  Geiger's  standard  able  mens h a d b e e n  the  scale  f o r the  have  arvalis  could  been d e s i r a b l e t o o n s i z e i n t h e common that  s t a t i o n s would  of the ecoclimate  the data  n o t have  gathered  been  to which  suit-  the speci-  exposed.  may  i s an a n i m a l lives  gradient  overground  such  (McNab,  1957).  as the:pocket burrow  1971).  burrower  rufocanus) nests.  to which  animals  affect size variation.  i n closed  i s a superficial  (Kalela,  data  was n o t  so t h e i n f l u e n c e o f  (1973) work d e m o n s t r a t e d  (CIethrionomys in  of cases,  are d i f f e r e n t degrees  which  -latitude scale  of  will, i n  i n Switzerland  locality  the effects of climate  and t h e degree  sarius)  level  Burrowi^nci  There rial  close  i t would  i n any c a s e  as i t i s  agreed  on t h e s i z e o f M i c r o t u s  meteorological  indices  1.2.2.3.  While  network  even  i n the majority conditions  experi-  1973).  enough t o p r o v i d e  needed  this  substantially different conditions  be f o u n d  years  below  wider  i s that  o r human c l i m a t e ,  In the a i r l a y e r  general,  extensive  measured  of a  which  gopher  It,too, follows  (Geomys b u r a size  end o f t h e  as the grey-sided  overwinters  fosso-  A t one end o f  systems and f o l l o w s  At t h e o t h e r  such  c a n be  vole  u n d e r t h e snow b u t  a size-latitude  gradient  49 Somewhere i n t h e live nel  in a closed systems  also 68,  can  and  be  middle  burrow below  makes f o r a y s  surface  above  storage  qualify  as  of  For  fail  these  food  one to  decisive Mayr,  obey  animals  available  kinds  of  distinct  occurs  where or  or  they  59,  nests  60  and  underground  not  totally  As  relying  such,  Microtus  (Mayr,  depth  of  body  size  occur  the  on  "almost  the  soil  (W.B.  promay  consis-  1963:page or  but  arvalis  323).  amount may  be  Davis,  of the  1938;  i n v o l v e d can physiological;  Interaction sympatric  displacement. of  Wilson, i n the species  There  s t r o n g l y i n f l u e n c e d by  with  be  - they  - but  toward  related  the  second,  member o f even  assumed  to  be  anoth-  the  the  ecological, to  are  one  i t i n some c h a r a c t e r s .  are  the  species  'displace'  w h e r e one  morphological,  they  pair  extent  The behavi-  genetically  1956). form  of  competition  is implicit  i s no  competition,  closely  together  identical  and  be  p r e s e n t . Two  i t converges  (Brown  finition  can  more c h a r a c t e r s  nearly  characters oural,  variation  competitors  alone  being  lated  pages  Its  tun-  overwinters  snow c o v e r ,  the  not  _omp_tit_o_  i n one  based  amount o f or  n:  mammals w h i c h  i n determining  Geographic  of  the  chambers.  f and  Bergmann's R u l e "  i n winter  factor  for food,  burrowing  I t does  1963).  1.2.2.4.  er  the  storage  XVIe,  chambers.  arvalis.  i t does have e x t e n s i v e  common v o l e  ground  i n the  but  food  (Table  The  visions  tently  system,  underground  respectively).  i s Microtus  unanimity  yet  in this  between c l o s e l y theory  concerning  i t always  of  the  means t h a t  character  precise  two  re-  or  de-  more  50 individuals, sential live,  populations  environmental  to hide,  Wilson,  o r t o breed  Throughout  meters),  species altitudes  this  kind  less  arable  cy  XVIb  cake'  XVIc:  page  a n d more  specimens  distribution There  terspecific  do g i v e  view,  however,  or  o f these  three  5 6 ) , and t h e S w i s s specimens  were  i s an a c c u r a t e  reflection  vole or the  i s unknown.  an a n i m a l  taken.  However,  of cultivated  lands  with  altitude,  zones,  giving  of  way t o  i t i s possible that the  a true picture of the vole's  frequen-  i n Switzerland.  competition  from  congeneric  5 7 ) , and as t h e a v a i l a b i l i t y  rocky  i s , of course,  voles  (page  of the collectors  i s primarily  vole  1800 m e t e r s ) ,  ranges  o f t h e common  of h a b i t a t decreases land  (Brown and  meters).  diagram  distribution  (Table  supply  (up t o 1000  (above  t h e 4 2 2 M^_ a r v a l i s  'wedding  arvalis  pastures  museum  t o 1800  distribution  Microtus  an e s -  or a place to  one o t h e r  v o l e , M. a g r e s t i s  i n Table  which  the frequency  and  i s i n limited  at least  the f i e l d  data  from  this  frequency as  as f o o d ,  8 i l l u s t r a t e s the altitude  from  simultaneously  i n S w i t z e r l a n d , t h e common  with  1000 meters  Figure  of  such  o r t h e snow v o l e , M. n i v a l i s  (from  Whether  i t s range  - either  that  seek  1963).  i s sympatric  species  both  resource,  1956; Mayr,  probably  or species  museum  no way t o f i n d  among t h e t h r e e  collections  can provide  direct  of i n -  resident species of  o f M. a r v a l i s .  certain  evidence  clues.  A literature re-  Table  XVI compares  FIGURE 8: ALTITUDE RANGES OF THREE SPECIES OF MICROTUS  14 43 74 87 97  107  M. AGRESTIS  M. ARVALIS  M. NIVALIS  NUMBER OF SPECIMENS IN SWITZERLAND IN EACH ALTITUDE GROUP OF 400 METERS  the  life  histories  non  reproductive  productive There through tivity XVIh:  are three  rhythms  (Table  2 i^y. a  arvalis Brink,  agrestis (Brink,  a  t o q: p a g e s  categories  c  i  r  i s chiefly  1967; S a i n t  nm  nivalis.  Brink  snow  a principally  crepuscular  ski  (1957)  case  reflect  XVIc:  (Table  page 5 7 ) .  This  o f M. a g r e s t i s ,  as season,  Girons  after  and Janeau  field  or  reproduc-  predators.  periods  o f M.  (1973) c o n s i d e r e d t h e whereas  i t was  observations  (1975)  and a f t e r  and M a r s z a l e k  by  labelKowal-  labora-  (1974).  c o n t r a d i c t i o n s i n M.  plasticity.  have  t o one o r  ( E r k i n a r o , 1969)  activity  may  indicated a bio-  i n response  of competitors  these  arvalis  i n rhythm  o r i t may h a v e  t o change  by B i e n k o w s k i  a behavioural  change  diurnal creature,  a n d Le L o u a r n  M.  M.  c o n d i t i o n s , i t was m o r e a c t i v e  1933).  or nocturnal  (Delost,  In the w i l d ,  d i u r n a l than  (1967) and S a i n t  observations  cycling  or nocturnal  1973).  was d i v i d e d on t h e p e a k  led  (Table  daily ac-  s  Girons,  c o n d i t i o n , age, presence  vole  competition  displacement:  crepuscular  an a b i l i t y  v a r i a b l e s , such  Opinion  reflect  63), population  t o be m o r e  (D.H.S. D a v i s ,  plasticity,  55 t o 6 5 ) a n d n i n e r e -  may  character  X V I i : page  - 20  66 t o 7 1 ) .  which  1967) b u t i n l a b o r a t o r y  several  the  t o k: p a g e s  an a b e r r a t i o n i n c a p t i v i t y  rhythm  i n 29 c a t e g o r i e s  - JLiy_ JLy_ !!y.t . „  was r e p o r t e d  night  tory  species  page 62) and h a b i t a t s e l e c t i o n  1955b;  tive  XVII  XVIa  a c t u a l or p o t e n t i a l  M.  been  (Table  (Table  1.2.2.4.1.  at  of the three  As i n  nivalis  may  53 1.2.2.4.2.  P°£ A _iori__c_c_ing_ u  a  agrestis Britain and  was r e p o r t e d  (Summerhayes,  D.  Chitty,  Lindahl,  I960;  (Elton,  all  the countries  (The  Britain.)  are  cycling  1970); M.  that  dynamics  o f M.  arvalis  M.  only  M.  agrestis  where  M.  suppresses  a g r e s t i s when  1965).  or mainland  however, t h a t  arvalis  cycles i n  (Elton,  i n Scandinavia  proportions  implying  (Curry-  a n d on t h e c o n t i n -  and R u s s i a  (1967) s t i p u l a t e d ,  i s no i n f o r m a t i o n  population  not  cycling,  (Saint Girons,  1.2.2.4.3.  arvalis  t h e pop-  t h e two  species  ies  (Regnier 1967;  Tapper,  alpine  well  Janeau,  1976);  M.  ler,  apparently  i t  differentiated  among  the three  spec-  arvalis  (Corbet,  1975).  Indeed  (Bernard,  1967);  1966; B r i n k ,  although  some  to tolerate  1953) or r e q u i r e d  1955; Corbet,  o r wet g r a s s l a n d  a cultivated  1966; B r i n k ,  species  a g r e s t i s was n o t a b l e  cesses  i s subject  where  1926; E l t o n , 1965; Corbet,  1953; Corbet,  zone  nivalis  f o r France  a g r e s t i s a woodland  and P u s s a r d ,  (Bernard,  and  - M.  M.  i s  s_election  appears  of voles  except  on w h e t h e r  1973).  Habitat  Habitat  M.  i n Scandinavia  1973).  not occur  cycles i n  1 9 5 2 a n d 1 9 5 4 ; H. C h i t t y  and M y l l y m a k i , Girons,  population  sympatric. There  to  1966);  i n number t o p l a g u e  not occur,  Chitty,  o f c e n t r a l Europe does  Brink  increases  ulation  Corbet,  1965; S a i n t  common v o l e  does  1 9 4 1 ; D.  1956; Kanervo  ent  to undergo  1966; Ashby,  have  1967),  species  nivalis  1 9 6 7 ; Le  human  dense  1966; B r i n k ,  land M.  species  a  Louarn  suggested  that  cultivation  ground  Brink  rocky  cover  pro(Schind-  (1967) and  Saint  54 Girons  ( 1 9 7 3 ) commented  absent, Le were  M.  a g r e s t i s took  Louarn  still  France,  and  low  altitudes  i t would  be  due  to  forced  M.  establish  nivalis  nivalis  rather  the  than  nature  of  i t s outnumbering 1973).  were t o  vary  the 2.  mountains  the  and/or  a  and  M.  nivalis zone  arvalis.  species  the  in  Saint  of  latter  only  presence  the  This  arvalis  other  two  Microtus  i s able  i f M.  of  field  or  arvalis  be  to and  but  to  the  1953)  or  (Bernard,  i t appears  not  of  M.  a g r e s t i s and  due  (Bernard,  competition,  vole  absence  i n M.  may  species  reason,  result  the  i f M.  rare.  Microtus  as  of  altitudes  reverse.  Whatever  also  snow v o l e  absence  e t h o l o g i c a l changes  the  was  Mediterranean  more p r o l i f i c  lower or  arvalis  that  i n the  i t seems i t i s t h e  produces  Girons,  the  the  the  absent  general that  simply Saint  i f size  i t would  i n the  more  1953;  that  M.  do  common  so  in  vole.  Subspeciation It  i s not  infraspecific on  at  a g r e s t i s are  aggressive  into  itself  arvalis  to  (1975) remarked  at  that  w h e r e M.  i t s habitat.  found  (1973) wrote  In  i n areas  over  Janeau  Girons  M.  that  the  tion  a  this  variation  to  embark  and  and  1124)  single  which  nature  to  concept  (1974:page frame  relevant  can  definition  d i s c u s s i o n on  of  " i t i s probably really be  which  can  upon a h i s t o r y species.  be  accepted  of  According  meaningful  question by  to  taxonomists  debates  to  Scudder  species  to defini-  a l l organisms and  and  the  inherently impossible  precise, fully  applied without  subspeciation  in  TABLE XVIa: LIFE HISTORIES OF THREE SPECIES OF MICROTUS  ASPECT  BODY (PLUS HEAD) LENGTH (MM)  MICROTUS AGRESTIS  MICROTUS NIVALIS  MICROTUS ARVALIS  85-130 ( S a i n t G i r o n s , 1973)  82.5-122 ( S a i n t G i r o n s , 1973)  97-136.5 ( S a i n t G i r o n s , 1973)  95-133 ( B r i n k , 1967)  95-110 (Regnier and P u s s a r d , 1926)  117-140 ( B r i n k , 1967)  110-130 ( D i d i e r and Rode, 1939)  95-110 ( D i d i e r and Rode, 1939)  120-140 ( D i d i e r and Rode, 1939)  95-120 ( B r i n k , 1967) 100-111 ( M i l l e r , 1912)  TAIL LENGTH (MM)  25-46 ( S a i n t G i r o n s , 1973)  23-39 ( S a i n t G i r o n s , 1973)  44-71 ( S a i n t G i r o n s , 1973)  27-46 ( B r i n k , 1967)  30-35 ( R e g n i e r and P u s s a r d , 1926) 50-75 ( D i d i e r and Rode, 1939)  30-45 ( D i d i e r and Rode, 1939)  30-40 ( D i d i e r and Rode, 1939)  50-75 ( B r i n k , 1967)  30-45 ( B r i n k , 1967) 35-45 ( M i l l e r , 1912)  15- 21 ( S a i n t G i r o n s , 1973)  13-18 ( S a i n t G i r o n s , 1973)  16- 20.5 ( B r i n k , 1967)  14-18 ( R e g n i e r and Pussard,  17-20 ( D i d i e r and Rode, 1939)  15-16 ( M i l l e r , 1912)  18.3-21.5 ( S a i n t G i r o n s , 1973) 1926) 18.5-22 ( B r i n k , 1967)  HIND FOOT LENGTH (MM)  15- 18.5 ( B r i n k , 1967) 16- 18 ( D i d i e r and Rode, 1939)  19-22 ( D i d i e r and Rode, 1939)  TABLE XVIb: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS  MICROTUS ARVALIS  EAR LENGTH (MM)  8- 10 ( D i d i e r and Rode, 1939)  7-9 ( D i d i e r and Rode, 1939)  11.4-17.8 ( S a i n t G i r o n s , 1973)  9- 14 ( S a i n t G i r o n s , 1973)  8-12.3 ( S a i n t G i r o n s , 1973)  15-17 ( D i d i e r and Rode, 1939)  19- 52 ( B r i n k , 1967)  14-46 ( B r i n k , 1967)  38-50 ( B r i n k , 1967)  20- 47.6 ( S a i n t G i r o n s , 1973)  15-35 (Regnier and P u s s a r d ,  WEIGHT (GR)  MICROTUS NIVALIS  1926) 42-62 ( S a i n t G i r o n s , 1973)  30-35 ( D i d i e r and Rode, 1939) 16.2-51 ( S a i n t G i r o n s , 1973) 25-30 ( D i d i e r and Rode, 1939)  DISTRIBUTION  ALTITUDE RANGE (M)  A l l Europe-from S c a n d i n a v i a i n t h e n o r t h t o t h e Alps i n t h e south and from F i n l a n d i n t h e e a s t t o England, France, Pyrenees, P o r t u g a l i n t h e west and southwest ( D i d i e r and Rode, 1939)  G r e a t e r p a r t o f Europe and p a r t o f A s i a c h i e f l y between 45° and 55° n o r t h l a t i t u d e ( E l t o n , 1965)  S i e r r a de Gredos, S p a i n ; Pyrenees; A l p s ; C e n t r a l Appenines; p a r t s o f the Balkans; Carpathi a n s ; T a t r a ( C o r b e t , 1966)  In F r a n c e - t o 1800 ( S a i n t G i r o n s , 1973)  In France-to 2800 ( S a i n t G i r o n s , 1973)  In S w i t z e r l a n d - t o 1800 ( G d l d i , 1914)  In S w i t z e r l a n d - t o 2350 ( G d l d i , 1914) In S w i t z e r l a n d - t o 2436 ( D o t t r e n s , 1962)  In Alps-1000 t o 4000 ( S a i n t G i r ons, 1973) In Alps-1500 t o 2600 (Le Louarn and Janeau, 1975) In A l p s - t o 3500 ( A l l e e and Schmidt, 1962) In F r a n c e - t o 3600 ( D i d i e r and Rode, 1939) In S w i t z e r l a n d - 1 6 0 0 t o 2600 ( C o r b e t , 1966)  TABLE XVIc: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  HABITAT  MICROTUS AGRESTIS  MICROTUS ARVALIS  In B e l g i u m - f a l l o w l a n d ; f o r e s t e d areas ( B e r n a r d , 1953)  In B e l g i u m - c u l t i v a t e d l a n d ( B e r n a r d , 1953)  In B r i t a i n - p u r e g r a s s l a n d ( E l t o n , 1965) In B r i t a i n - c o v e r c o n s i s t s mainly o f g r a s s e s and i s dense a t ground l e v e l (Ashby, 1967) I n B r i t a i n - o p e n , g r a s s y area (Tapper, 1976)  In F r a n c e - g r a s s l a n d w i t h o r w i t h o u t shrubs (Le Louarn, S p i t z and D a s s o n v i l l e , 1970)  In France-woods and p l a n t a t i o n s ( R e g n i e r and P u s s a r d , 1926) In Scandinavia-open h a b i t a t s ( C u r r y - L i n d a h l , 1956) In G e n e r a l - g r a s s l a n d ; wet ground w i t h rushes and sedge ( C o r b e t , 1966). In General-moist a r e a s ; open woods ( B r i n k , 1967)  In M e d i t e r r a n e a n area-growths o f rushes (Rey, 1973) In G e n e r a l - p a s t u r e s ; meadows (Corbet, 1966) In G e n e r a l - p a s t u r e s ; meadows; f i e l d s ; c u l t i v a t e d land ( B r i n k , 1967)  MICROTUS NIVALIS In F r a n c e - a l p i n e and s u b a l p i n e meadows w i t h r o c k s l a r g e r than 20 cm (Le Louarn and Janeau, 1975) In General-above t r e e l i n e ; m i x t u r e l o o s e rock and g r a s s ; open mountain woodland ( C o r b e t , 1966) In G e n e r a l - h i g h mountains; rocky s l o p e s ; stoney meadows; open woods ( B r i n k , 1967)  TABLE XVId: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS For Females-27 m i n l e n g t h (Brown, 1966) For Males-36 m i n l e n g t h (Brown, 1966)  HOME RANGE SIZE  In G e n e r a l - g r e a t e s t d i s t a n c e a c r o s s seldom more than 913.5 m (D. C h i t t y , 1952) In G e n e r a l - s m a l l e r t h a n 210 sq m (Godfrey, 1954) I n G e n e r a l - s i z e and shape i n summer f l u c t u a t e s ; depends on g r a d u a l changes i n v e g e t a t i o n and c u l t i v a t i o n methods ( C u r r y - L i n d a h l , 1956)  MICROTUS ARVALIS For Females-10-20 m i n d i a meter i n b r e e d i n g season (Frank, 1957) For Females-300-500 s q m i n conifer plantation (Reichs t e i n , 1960b) For F e m a l e s - s m a l l e r t h a n 7 m i n l e n g t h (Dub, 1971a) For Females-1/4 o f 146-600 s q m ( N i k i t i n a , K a r u l i n and Zen'Kovich, 1972) For Maies-40-100 m i n d i a m e t e r i n w i n t e r ( F r a n k , 1957) For Males-1200 t o 1500 s q m i n conifer plantation (Reichs t e i n , 1960b) For M a l e s - s m a l l e r t h a n 13 m i n l e n g t h (Dub 1971b) For Males-1/4 o f 828-1008 s q m ( N i k i t i n a , K a r u l i n and Z e n ' K o v i c h , 1972) In General-each f a m i l y o c c u p i e s o n l y a few s q m ( B e r n a r d , 1959) In G e n e r a l - s m a l l e r t h a n 10 m ( S p i t z , 1963b) In G e n e r a l - d e c r e a s e i n s i z e with increase i n density i n summer ( S p i t z , 1970a) In General-a few s q m around burrow ( S a i n t G i r o n s , 1973) In General-7.5-9 m ( S p i t z e t a l , 1974)  MICROTUS NIVALIS  TABLE XVIe: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  RUNWAY SYSTEMS  MICROTUS AGRESTIS  MICROTUS ARVALIS  In Britain-extremely complicated, extensive systems (Summerhayes, 1941) In B r i t a i n - s u r f a c e systems (Tapper, 1976)  In B r i t a i n (Orkney I s l a n d s ) w e l l marked systems (Turner, 1965)  In General-prominent systems through grass (Corbet, 1966) In General-systems above ground but w e l l hidden (Brink, 1967)  In France-systems s u p e r f i c i a l (Saint Girons, 1973) TUNNERL SYSTEMS  In General-systems r a r e l y extensive (Corbet, 1966) In General-systems near surface (Brink, 1967)  MICROTUS NIVALIS  In General-systems connect tunnels on surface (Brink, 1967)  In Belgium-systems at depths varying with s o i l c o n d i t i o n s ; u s u a l l y a few cm from the surface (Dalimier, 1955) In B r i t a i n (Orkney Islands)short t o complex systems (Turner, 1965) In France-very deep systems (Saint Girons, 1973) In Poland-100-150 cm long systems at 35 cm depth (Cechowicz and Pucek, 1961) In General-extensive, complex, shallow systems (Corbet, 1966)  In France-entrances t o systems beside rocks (Le Louarn and Janeau, 1975) In General-systems with several openings (Brink, 1967)  TABLE X V I f : LIFE HISTORIES OF THREE SPECIES ASPECT  MICROTUS AGRESTIS In B r i t a i n - s t o r e s g r a s s seed (Baker and Ranson, 1932b) In F r a n c e - s t o r e s c e r e a l s , g r a s s , seeds o f composites ( D i d i e r and Rode, 1939) In General-has food s t o r a g e chambers ( B r i n k , 1967)  FOOD STORAGE  MICROTUS  MICROTUS ARVALIS In Belgium-2 k i n d s o f s t o r a g e chambers: autumn one f o r seeds, g r a i n s ; w i n t e r one for roots, bulbs, tubercles ( D a l i m i e r , 1955) In F r a n c e - o v a l s t o r a g e chamb e r s , 10-12 cm from s u r f a c e ; i n w i n t e r s t o r e s 2-3 kg o f rhizomes, b u l b s , t u b e r c l e s , roots o f wild p l a n t s (Regnier and Pussard, 1926) In F r a n c e - s t o r e s seeds o f c e r e a l s , lucern grass, apples D e l o s t , 1955b) In USSR-stores 2.5-3.3 kg o f f o o d such a s b u l b s ( G l a d k i n a and Chentsova, 1971) In G e n e r a l - s t o r e s food i n underground chambers ( B r i n k , 1967)  MICROTUS NIVALIS  TABLE XVIg: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  DIET  MICROTUS AGRESTIS  MICROTUS ARVALIS  In Britain-stems, leaves, roots of rough grasses of a l l kinds; c e r t a i n herbaceous plants (Ranson, 1934)  In Belgium-seeds of c u l t i v a t e d c e r e a l s , w i l d p l a n t s ; i n capt i v i t y small i n s e c t s ( D a l i mier, 1955)  In Britain-herbaceous vegetat i o n ; trees only attacked when vole numbers very high (Summerhayes, 1941)  In France-seeds s t i l l green; i n c a p t i v i t y , invertebrates (Saint Girons, 1973)  In B r i t a i n - g r a s s e s (Godfrey, 1953b and 1955) In B r i t a i n - g r e e n stems and leaves; not seeds and f r u i t (Elton, 1965)  In Switzerland-green p a r t s of p l a n t s , not the roots (Goldi, 1914)  In France-young shoots, green parts of plants (Saint Girons, 1973) In General-almost e n t i r e l y herbivorous: i n summer, stems, leaves of grasses, rushes, sedges; i n winter, roots, rhizomes, bark (Corbet, 1966)  In General-nuts; bark of l i v i n g t r e e s ; seeds of c l o v e r , lucern, wheat, oats, rye, barley, corn; potato tubers (Elton, 1965)  MICROTUS NIVALIS In France-seeds, r o o t s , flowers, leaves of a l p i n e p l a n t s (Didier and Rode, 1939) In France-roots, seeds of alpine p l a n t s e s p e c i a l l y b i l b e r r y (Saint Girons, 1973) In General-grasses, herbs, especially bilberry (Corbet, 1966)  TABLE XVIh: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  PEST STATUS  MICROTUS ARVALIS  In B r i t a i n - c h i e f agent of tree damage (Elton, 1965)  In most of Europe and European USSR-chief agent of damage t o crops; k i l l s t r e e s , shrubs by gnawing bark; destroys t r e e seeds (Elton, 1965)  In Bulgaria-source and t r a n s m i t t e r t o w i l d and domestic mountain grazers of p a r a s i t i c worms, espec i a l l y smaller f l u k e (Peshev, 1970)  In Belgium-cycles 1953 and 1959)  In France-does not c y c l e (Saint Girons, 1973)  In Scandinavia-chief agent of damage to ornamental p l a n t s , f r u i t trees, forest plantat i o n s , seed p l a n t a t i o n s (Kanervo and Myllymaki, 1970)  In B r i t a i n - c y c l e s (Summerhayes, 1941; D. C h i t t y , 1952 and 1954; H. C h i t t y and D. C h i t t y , 1960; Corbet, 1966) POPULATION CYCLING  MICROTUS NIVALIS  MICROTUS AGRESTIS  On Continent-cycles (Elton, 1965) In France-cycles (Saint Girons, 1973) In Scandinavia-cycles (Curry L i n d a h l , 1956; Kanervo and Myllymaki,1970)  (Bernard,  In Bulgaria-cycles (Straka and Gerasimov, 1971) In France-cycles ( S p i t z , 1967; Saint Girons, 1973) In Germany-cycles (Frank, 1957) In a l l the countries of cent r a l Europe and Russia-cycles (Elton, 1965)  ro  TABLE X V I i : LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS  MICROTUS ARVALIS  In C a p t i v i t y - 2 - 4 hour rhythm o f f e e d i n g a c t i v i t y and l o n g e r 24 hour rhythm; peak f o l l o w i n g s u n s e t ; more a c t i v e a t n i g h t (D.H.S. D a v i s , 1933)  DAILY RHYTHM OF ACTIVITY  In General-more d i u r n a l than M. a r v a l i s ( B r i n k , 1967) In G e n e r a l - a c t i v e day and night; 2 p r i n c i p a l times: s u n r i s e and sunset ( S a i n t G i r o n s , 1973) In G e n e r a l - a c t i v e a t n i g h t d u r i n g summer; a c t i v e d u r i n g day i n w i n t e r ( E r k i n a r o , 1969)  Low p i t c h e d p o l y s y l l a b i c (Frank, 1953b) Conspicuous c a l l 1966)  call  (Corbet,  In C a p t i v i t y - 8 - 1 0 p e r i o d s o f a c t i v i t y d u r i n g 24 hours (Durup, 1956) In C a p t i v i t y - 7 - 8 p e r i o d s o f a c t i v i t y d u r i n g 24 hours ( N i k i t i n a , K a r u l i n and Z e n ' k o v i c h , 1972) In General-mainly a c t i v e a t dusk ( B r i n k , 1967) In General-mainly n o c t u r n a l ( D e l o s t , 1955b) In G e n e r a l - a c t i v e day and n i g h t ; mainly a c t i v e a t n i g h t i n summer ( S a i n t G i r o n s , 1973) In G e n e r a l - a c t i v e a t n i g h t d u r i n g summer; a c t i v e d u r i n g day i n w i n t e r ( E r k i n a r o , 1969)  High m o n o s y l l a b i c (Frank, 1953b)  call  Less v o c a l t h a n M. a g r e s t i s (Corbet, 1966; B r i n k , 1967)  VOCALIZATIONS Low and c h a t t e r i n g c a l l ( B r i n k , 1967)  High c h i r p i n g squeak ( B r i n k , 1967)  Noisy when handled 1969)  More s i l e n t than M. a g r e s t i s when handled (Dienske, 1969)  (Dienske,  MICROTUS NIVALIS In C a p t i v i t y - b i p h a s i c p a t t e r n s o f a c t i v i t y ; mainly n o c t u r n a l ( B i e n k o w s k i and M a r s z a l e k . 1974) In G e n e r a l - a c t i v e mainly a t n i g h t ( K o w a l s k i , 1957) In G e n e r a l - a c t i v e mainly a t sunset (Le Louarn and Janeau, 1975) In G e n e r a l - a p o e a r s o f t e n by day ( B r i n k , 1967)' In G e n e r a l - d i u r n a l ( S a i n t G i r ons, 1973)  Abrupt and p e n e t r a t i n g c a l l ; u s u a l l y s i n g l e n o t e s ; i n breedi n g season a c o n t i n u o u s c h a t t e r i n g ( B r i n k , 1967)  TABLE X V I j : L I F E HISTORIES OF THREE SPECIES ASPECT  MICROTUS AGRESTIS In B r i t a i n - v e r y l o c a l i n movements (D. C h i t t y , 1952) In S c a n d i n a v i a - r e g u l a r autumn m i g r a t i o n from open f i e l d s t o f o r e s t borders and sometimes t o f o r e s t ; i n s p r i n g back t o summer q u a r t e r s , ( C u r r y L i n d a h l , 1956)  MIGRATION  MICROTUS MICROTUS ARVALIS In Czechoslovakia-manmade m i g r a t i o n : d u r i n g autumn c u l t i v a t i o n measures, r e s i d e n t voles emigrate t o adjacent h a b i t a t s (Dub, 1971b) In F r a n c e - l o c a l m i g r a t i o n s ( R e g n i e r and P u s s a r d , 1926) In F r a n c e - l o c a l m i g r a t i o n s : in spring to lucern, clover, g r a i n f i e l d s ; i n summer t o r y e and wheat f i e l d s ; i n w i n t e r t o t h i c k e t s , orchard borders ( D e l o s t , 1955b) In P o l a n d - s e a s o n a l m i g r a t i o n : d u r i n g and immediately a f t e r harvest, leave the c u l t i v a t e d f i e l d s ; end o f September when ploughing terminated, return m i g r a t i o n (Cechowicz and Pucek, 1961) In G e n e r a l - l o c a l m i g r a t i o n s ( B r i n k , 1967)  MICROTUS NIVALIS In F r a n c e - l o n g m i g r a t i o n s September t o A p r i l ; sedentary a f t e r A p r i l (Le Louarn and Janeau,1975) In P o l a n d - m i g r a t i o n s up mount a i n s i n summer ( K o w a l s k i , 1957) In S w i t z e r l a n d - p o s s i b l e v e r t i c a l seasonal migrations ( D o t t r e n s , 1962)  TABLE XVIk: L I F E HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  PREDATORS  MICROTUS AGRESTIS In France-weasels v i p e r s , noct u r n a l and d i u r n a l r a p t o r s ( D i d i e r and Rode, 1939) In F r a n c e - c a r n i v o r e s , v i p e r s , r a p t o r s ( S a i n t G i r o n s , 1973) In G e n e r a l - s t o a t s , w e a s e l s , p o l e c a t s , martens, f o x e s , c a t s , buzzards, eagles, harr i e r s , s h o r t e a r r e d owls, barn o w l s , snowy owls ( C o r b e t , 1966)  LIFE EXPECTANCY  MICROTUS ARVALIS  MICROTUS NIVALIS  In B e l g i u m - f o x e s , badgers, p o l e c a t s , ermines, weasels, g r a s s snakes, v i p e r s , r a p t o r s ( D a l i m i e r , 1955)  In F r a n c e - c a r n i v o r e s c o r v i d s , d i r u n a l raptors (Saint Girons, 1973) In F r a n c e - r e p t i l e s , c a r n i v o r e s (Le Louarn and Janeau, 1975)  In F r a n c e - w e a s e l s , martens, vipers, raptors (Delost, 1955b) In F r a n c e - c a r n i v o r e s , v i p e r s , r a p t o r s ( S a i n t G i r o n s , 1973)  A few months from weaning ( C o r b e t , 1966)  I f born February t o May: 2-4 months ( M a r t i n e t , 1967)  7-8 months ( L e s l i e and Ranson, 1940)  I f born March t o June: 2 months ( S p i t z , 1967)  F u l l y grown males o f 1 y e a r do n o t l i v e o v e r t o t h e next y e a r ( B a k e r and Ranson, 1933)  I f born J u l y t o October; 7-9 months ( S p i t z , 1967)  18 months maximum ( W a s i l e w s k i , 1956)  More t h a n 2 y e a r s ( D e l o s t , 1955b) 2-3 y e a r s ( R e g n i e r and P u s s a r d , 1926)  TABLE X V I I : L I F E HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  BREEDING SEASON  MICROTUS AGRESTIS  MICROTUS ARVALIS  MICROTUS NIVALIS  In B r i t a i n - M a r c h t o September (Baker and Ranson, 1933) In B r i t a i n - M a r c h t o October, ( B r a m b e l l and H a l l , 1939) In B r i t a i n - F e b r u a r y / M a r c h t o September/October ( A s d e l l , 1964) In B r i t a i n - f r e q u e n t l y an e a r l y end t o b r e e d i n g i n a peak year (D. C h i t t y , 1954)  In B e l g i u m - F e b r u a r y / M a r c h / A p r i l t o August/September/ October/November; b e g i n n i n g and end i n f l u e n c e d by i n t e r n a l f a c t o r s dominated by e x t e r n a l f a c t o r s such a s c l i m a t e , f o o d , population density (Bernard, 1964)  In F r a n c e - s p r i n g and summer ( S a i n t G i r o n s , 1973) In France ( i n h i g h A l p s ) - A p r i l t o August (Le Louarn and Janeau, 1975) In F r a n c e ( a t 1500 m ) - A p r i l t o September (Le Louarn and Janeau, 1975)  In F r a n c e - F e b r u a r y t o December; c a n be y e a r round ( S a i n t G i r o n s , 1973)  In France-March t o October ( D e l o s t , 1955b) In F r a n c e - c a n be y e a r round ( M a r t i n e t , 1967) In F r a n c e - F e b r u a r y t o O c t o b e r ; can be y e a r round (Le Louarn, S p i t z and G r o l l e a u , 1970) In F r a n c e - J a n u a r y / F e b r u a r y t o O c t o b e r ; l e n g t h v a r i e s dependi n g on c l i m a t i c , n u t r i t i o n a l , social conditions (Saint Girons, 1973) In Germany-February/March t o October/November; under f a v o u r a b l e c o n d i t i o n s c a n be y e a r round (Frank, 1957)  CTv  TABLE XVIm: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS For Females-21 days: i n l a b oratory (Baker and Ranson, 1933; L e s l i e and Ranson, 1940) For Males-6 weeks: i n laborat o r y ( L e s l i e and Ranson, 1940) For Males-6 t o 8 weeks: i n laboratory (Baker and Ranson, 1933)  AGE AT FIRST BREEDING  For Both-6 weeks (Corbet, 1966) For Both-in w i l d : i f born l a t e i n season a t t a i n puberty f o l l o w i n g spring (Brambell and H a l l , 1939)  MICROTUS ARVALIS For Females-15 to 20 days: i n laboratory (Bashenina, 1953) For Females-17 to 21 days: i n laboratory (Bernard, 1964) For Females-20 days or e a r l i e r : i n laboratory (Frank and Zimmermann, 1957a) For Females-20 to 25 days: i n laboratory and i n w i l d (Del o s t , 1956) For Females-21 days: i n l a b o r atory (Martinet, 1967) For Females-in Germany, i n w i l d : i f born during and a f t e r September do not reach sexual maturity i n same year (Frank, 1957) For Males-35 days: i n l a b o r a t o r y (Bashenina, 1953; M a r t i n e t , 1967) For Males-35 days: i n w i l d ; 40 days: i n laboratory (Delost, 1956) For Both-2 to 2 1/2 months (Regnier and Pussard, 1926) For Both-in France, i n w i l d : i f born February t o May reach sexu a l maturity q u i c k l y ; i f born a f t e r June, do not reach sexual maturity the year of b i r t h (Mart i n e t , 1967)  MICROTUS NIVALIS For Both-in w i l d : growth stops w i t h f i r s t snow but continues i n April/May when sexually mature (Le Louarn and Janeau 1975)  TABLE XVIn: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  NEST  MICROTUS AGRESTIS  MICROTUS ARVALIS  In B r i t a i n - n e a r l y a l l n e s t s found between May and September b u i l t above ground i n s l i g h t excavation i n s o i l ; g l o b u l a r i n shape, o f f i n e l y shredded dry g r a s s (Godfrey, 1953a)  In B e l g i u m - s p h e r i c a l n e s t s l i n e d w i t h dry g r a s s o r s t r a w ; sometimes i n summer s u r f a c e n e s t s under c o v e r such as hay s t a c k ( D a l i m i e r , 1955)  In F r a n c e - a t t h e s u r f a c e , sometimes under the s h e l t e r o f a rock ( S a i n t G i r o n s , 1973) In G e n e r a l - u s u a l l y c o n s t r u c t ed above ground; under a stone or l o g ; sometimes s l i g h t l y below ground l e v e l ( C o r b e t , 1966) In G e n e r a i - o f t e n above ground; o f g r a s s ( B r i n k , 1967)  MICROTUS NIVALIS In G e n e r a l - n e s t chambers o f hay and s t a l k s o f t e n j u s t below surf a c e ( B r i n k , 1967)  In B r i t a i n (Orkney I s l a n d s ) subterranean chambers l i n e d with roots, grass; surface n e s t s i n l o n g g r a s s and heather (Turner, 1965) In F r a n c e - s u r f a c e and underground n e s t s ; 8-10 cm i n d i a meter; o f d r i e d g r a s s on o u t s i d e , straw on i n s i d e ; 5-15 cm below ground i n summer, deeper i n w i n t e r (Regnier and P u s s a r d , 1926) In Poland-nest chambers w i t h 2-3 o u t l e t s ; 15 cm i n d i a m e t e r ; 7-12 cm h i g h (Cechowicz and Pucek, 1961) In General-underground n e s t chambers o r nest can be b u i l t above ground o f g r a s s and s t a l k s ( B r i n k , 1967)  <7> 03  TABLE XVIo: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS P o s t partum (Ranson, 1934; ( B r a m b e l l and H a l l , 1939; C o r b e t , 1966)  OESTROUS  MICROTUS ARVALIS  MICROTUS NIVALIS  Post partum ( S a i n t G i r o n s , 1973)  2 kinds of cycles: a t y p i c a l s h o r t c y l c e o r prolonged p e r i o d s o f an o e s t r o u s s t a t e (H. C h i t t y and A u s t i n , 1957)  Spontaneous ( B r a m b e l l and H a l l , 1939)  Spontaneous ( D e l o s t , 1955d; A s d e l l , 1964)  OVULATION Induced (Breed, 1967) Spontaneous and i n d u c e d (H. C h i t t y , 1957)  18-20 days ( D i d i e r and Rode, 1939) 20-21 days (H. C h i t t y , 1957) GESTATION PERIOD  21 days (Ranson, 1934; L e s l i e and Ranson, 1940; Corbet, 1966; S a i n t G i r o n s , 1973)  16-24 days (Frank and Zimmermann, 1957a) 16-25 days ( R e i c h s t e i n , 1964) 20 days (Regnier and P u s s a r d , 1926) 20-21 days ( D e l o s t , 1955b and 1955d) 21 days ( M a r t i n e t , 1967; S a i n t G i r o n s , 1973) Length o f g e s t a t i o n p e r i o d depends on s i z e o f l i t t e r - more embryos, s h o r t e r l e n g t h o f t i m e ( R e i c h s t e i n , 1964)  21 days ( S a i n t G i r o n s , 1973)  TABLE XVIp: L I F E HISTORIES OF THREE SPECIES ASPECT  MICROTUS AGRESTIS 3 l i v e b i r t h s per l i t t e r ; number d e c r e a s e s as age o f mother i n c r e a s e s ( L e s l i e and Ranson, 1940) 3-6 ( C o r b e t , 1966) 3- 8 ( S a i n t  G i r o n s , 1973)  4 a v e r a g e (Ranson, 1934) 4- 7 ( D i d i e r e t a l , 1939)  LITTER SIZE  L i t t e r s i z e may be g r e a t e r i n a peak y e a r than i n any o t h e r (D. C h i t t y , 1954)  - MICROTUS MICROTUS ARVALIS  MICROTUS NIVALIS  3-5: i n l a b o r a t o r y ( D e l o s t , 2-4 (Le Louarn and Janeau, 1955d) 3-7: i n w i l d ( D e l o s t , 1955a) 2- 7 ( S a i n t G i r o n s , 1973) 3-7 ( S a i n t G i r o n s , 1973) 3- 8: i n w i l d ( S t e i n , 1957) 3- 7 ( D i d i e r and Rode, 1939) 4.36 a v e r a g e : i n l a b o r a t o r y Frank and Zimmermann, 1957a) 4- 6 ( D a l i m i e r , 1955; Le Louarn, S p i t z and G r o l l e a u , 1970) 5- 6 ( R e g n i e r and P u s s a r d , 1926) L i t t e r s i z e depends on i n h e r i t a n c e , age and s i z e o f mother, season; m o d i f i e d by q u a l i t y and q u a n t i t y o f food (Frank, 1957) L i t t e r s i z e depends on q u a l i t y of f o o d , age o f mother ( B e r n a r d , 1964) L i t t e r s i z e depends on f o o d , s i z e and age o f mother, f r e quency o f l i t t e r s , l e n g t h o f d a y s , i n t e n s i t y o f l i g h t , population density (Reichstein, 1964) L i t t e r s i z e maximum (> 6) i n A p r i l / M a y , minimum (3) November t o F e b r u a r y ( S p i t z , 1967) L i t t e r size varies i n relation t o s e a s o n , age o f mother (Wojciechowska, 1970)  TABLE XVIq: LIFE HISTORIES OF THREE SPECIES OF MICROTUS ASPECT  MICROTUS AGRESTIS 12-16 days (Godfrey, 1953a)  MICROTUS ARVALIS  MICROTUS NIVALIS  More than 2 weeks (Dalimier, 1955)  14 days (Ranson, 1934; L e s l i e and Ranson, 1940; Clarke, 1955) 15 days ( S p i t z , 1970b) LACTATION PERIOD  Fewer than 21 days (Corbet, 1966)  15-18 days (Regnier and Pussard, 1926) 15-20 days (Delost, 1955d) 17 days (Wojciechowska, 1970)  NUMBER OF LITTERS  6-8 (Didier and Rode, 1939)  2 u s u a l l y ( S p i t z , 1967)  2 probably (Kowalski, 1957)  Succession of l i t t e r s bet, 1966)  3-5 (Dalimier, 1955)  2: f i r s t l i t t e r i n June, second l i t t e r mid J u l y t o mid August (Le Louarn and Janeau, 1975) 2-3 (Didier and Rode, 1939)  Several l i t t e r s ons, 1973)  (Cor-  (Saint G i r -  4-5 (Regnier and Pussard, 1926) Varies according t o q u a l i t y of food (Bernard, 1964)  72 evolutionists". necessary can  to e s t a b l i s h  be d i s c u s s e d .  definition species are  Mayr's  a r e "groups  species,  and d i f f e r i n g  in  1969:page  category  category  and need  t h e whole  Meylan  ate  subspecies  and  bears As  M.  contains  t h e same  mentioned  (1970) concluded  study  -  41).  recognized  from  the type  "A  that  subspecies  o f t h e range  other  of a  populations of i s the  i n the International i s , however,  in principle,  two s u b s p e c i e s  a. a r v a l i s  - a species  The s u b s p e c i e s  Subspecies n o t , even  that  populations of a  subdivision  a. r u f e s c e n t e f u s c u s  subspecies  species  groups".  1969).  of a c l a s s i f i c a t i o n  (1966) l i s t e d  S w i t z e r l a n d : M.  nominate  such  species  (Mayr,  taxonomically  Z o o l o g i c a l Nomenclature.  throughout  other  a geographic  s p e c i e s " (Mayr,  obligate  of this  of phenotypically similar  inhabiting  of  from  two o r more s u b s p e c i e s  taxonomic  subspeciation  26) b i o l o g i c a l  f o r the purposes  i s a polytypic  species,  lowest  which  species i s  of interbreeding natural populations  arvalis  an a g g r e g a t e  o f t h e term  within  (1969:page  reproductively isolated  contains  the  a context  i s adequate  Microtus  is  Y e t some d e f i n i t i o n  a non-  be  (Simpson,  Code  used  1961).  of Microtus  arvalis  ( S c h i n z , 1845) and t h e  (Pallas,  1779).  of the subdivided  (The nominhigher  taxon  name.) p r e v i o u s l y both  in their  papers  Dottrens  (1962) and  that the voles  from  Jones the lowest  altitudes  ( 4 0 5 m and 750 m r e s p e c t i v e l y ) b e l o n g e d  nominate mid  subspecies  and h i g h e s t  turn  whereas  represented  I t was t h e a b s e n c e  t o Bergmann's Rule  that  the voles  recorded.  discussion  section,  geographic  variation  than  the rule.  why t h i s  adherence  both  may  i n nature  other  than  be p a r t i c u l a r l y  true  i n Microtus  differentiation  (Durrant,  1954) and they  s u b d i v i s i o n s o f the range  diagnostic 1963). terms (1) from  Therefore,  demonstrate a l l other being  delimit  populations assigned  this  vole  variation  o f M. a r v a l i s  to a d e f i n i t e  morphological  enumerat  morphological definite  between t h e  and t h e e n v i r o n m e n t  any a n a l y s i s o f g e o g r a p h i c  morphological  o  arvalis.  inhabit  by c o r r e l a t i o n  characters  theory  of the species,' the  o f s u b s p e c i a t i o n i n t h e common  lation's (2)  morphological  largely  of this  s u b s p e c i a t i o n were  on t h e b a s i s o f t h e i r  determined  part  be m o r e t h e e x c e p t i o n  a r e named  distribution  f o r the size  t o Bergmann's  Subspecies  geographic  variation  researchers to  i n the f i r s t  i n h o m e o t h e r m s , may  Reasons  M. a.  of altitudinal  prompted  Y e t , as o u t l i n e d  from t h e  the subspecies  t o s u b s p e c i a t i o n as a p o s s i b l e e x p l a n a t i o n  trends  ed  a. a r v a l i s ,  altitudes  rufescentefuscus. according  M.  to the  variation in  i n Switzerland  must  sufficiently distinct to warrant  taxonomic  variation  (Mayr,  t h e popu-  category;  within a  and  definite  range. 2.1.  Mor_pho_og_ica_ Va_i_t._on_ Mayr  important much  (1942:page taxonomic  individual  36) c a u t i o n e d character  variation".  only  that  "absolute  size  i f i t i s not subject  The a v e r a g e  range  i s an to too  i n mammals a n d  birds  of purely  the  size  the  average  of different size  Individual could size al  individual  variation  p a r t s i s from  f o r the species variation  be much h i g h e r  as i t i s a very  as s o c i a l  (Saint  before  recognizing the v a l i d i t y  valis  based  Jones  (1970) based  these  Girons,  t h a t under  entirely  like  lengths  i n M.  arvalis  study.  To t h e r i g h t  cording  to Dottrens  morph.  The l a r g e r a n g e  demonstration  1973).  montane Table  recorded  s p e c i e s and t h e factors,  one must  Both  XVII  be v e r y  (1973)  careful  i n Microtus a r -  Dottrens  by D o t t r e n s ,  should  nutrition-  (1962) and  hypothesis  entire-  c o m p a r e s t h e mean  line  of  arvalis  Saint Girons  subspecies  o f the double and J o n e s  Microtus  of subspecies  on d i m e n s i o n s .  their  and i n  1906).  plastic  circumstances  on e x t e r n a l d i m e n s i o n s .  sive  (Allen,  a t t a i n e d by a d u l t s d e p e n d s o n n u m e r o u s  as w e l l  size  8 t o 1 5 t o 20 p e r c e n t  i n a species  warned  ly  i n general  Jones  and  body  this  a r e t h e means w h i c h a c -  apply  t o t h e montane  among t h e m e a n s r u l e s  o u t any c o n c l u -  o f t h e e x i s t e n c e o f a montane  morph.  T A B L E X V I I : COMPARISON OF BODY LENGTHS WITH A L T I T U D E I N MICROTUS A R V A L I S ( D O T T R E N S , 1 9 6 2 ; J O N E S , 1 9 7 0 ; T H I S T H E S I S ) REFERENCE  4 0 0 - 8 0 0 METERS' 1 2 0 0 - 1 5 0 0 METERS N MEAN BODY : N MEAN BODY LENGTH ( M M ) | LENGTH (MM)  Dottrens (1962)  17  97.8  Jones (1970)  39  94.4  60  92.7  This  85  92.9  36  91.2  thesis  |  ?  1 9 0 0 + METERS N MEAN BODY LENGTH (MM)  105.1 . II  ?  95.8  24  101.1  11  93.4  There in  subspecific  species  with  ameliorate ical  of  are at l e a s t  highly  disorderly  The  first  an  isted Mayr  trends of  Brown  like  (1953)  M.  of the degree  that  may  f o r formal (1963;page  appear  variation  and  called  as  within  possible  the  whole  accompanied  significance  apparent  remarked  of a species  crit-  i s known,  i f a need  any  After  still  ex-  classification. "the b e t t e r  by  of  W i l s o n , 1954) .  infraspecific  however,  might  theory.  names, and  and  of  t h e most  a picture  possible  (Brown  become  taxonomic 348),  arvalis,  systematic  latinized  i f implemented  i n the cases  individual characters  overview i t would  graphic  growth  a r e a o f modern  u n c o m p l i c a t e d by  concordance  particularly  i s t o t r y f o r as c o m p l e t e  some e v a l u a t i o n  such  variable  what W i l s o n and  variation  procedures which  classification,  and  species,  three  As  the  geo-  t h e more d i f f i c u l t i t  becomes t o d e l i m i t s u b s p e c i e s " . The  second  subspecies acters  are c l a s s i f i e d ,  which  ecological fication  i s t o expand  can  and  at the  be  beyond  highly  primarily  variable,  ethological species  the diagnostic  characters  level  and  may  characters  whereby  morphological char  to include  physiological,  which  are used  prove  a more  in  classi  stringent  test. The and  third  percent rules.  constitutes son, fic 75  i s to e s t a b l i s h  1943).  an  adequate  There  classification percent rule  There  working c r i t e r i a  i s no  absolute  on  criterion  sample as  sample  f o r taxonomic  purposes  are several  percent rules  used  - the  (Mayr,  50  percent rule  1969),  and  the  84  (Edwards,  in  to  what  (Simpsubspeci-  1954),  percent rule  size  the  (Simpson  and  Roe, 1 9 3 9 ) .  For  example,  mists  i n the case  judge a p o t e n t i a l  individuals viously er  Even t h e i r  differ  previously 2.2.  subspecies  subspecies  subspecies  rule,  on whether  some  75 p e r c e n t  1 9 5 4 ) ; o t h e r s o n wheth-  differ  f r o m 97 p e r c e n t  (Mayr,  1969).  range o f M i c r o t u s  arvalis  covers  ton,  1965).  Burt  continuous  the s u b s p e c i f i c  concentrated according Just  r a n g e s would be b e t t e r  as the o v e r a l l  continental distribution  continuous,  specimens  found?  variations  Goldi  arvalis continu-  (1914) r e p o r t e d t h e common v o l e was  included i n this  385 m e t e r s  o f M.  so i s t h e r e d i s t r i b u t i o n a l  up t o 2350 m e t e r s , D o t t r e n s  If  and b e h a v i o u r a l  were  t o geography.  i n Switzerland.  range from  understood  d e s i g n a t i o n s were d i s c a r d e d a n d e f f o r t s  on the m o r p h o l o g i c a l  i s more o r l e s s  found  latitude (El-  (1954) recommended t h a t c o n t i n e n t a l s p e c i e s  w i t h more o r l e s s  The  of a  the greater p a r t o f  E u r o p e a n d A s i a c h i e f l y between 4 5 ° a n d 55° n o r t h  ity  of i t s  Geog_a_h_c_Ran_e The  if  taxono-  o f the i n d i v i d u a l s o f a pre-  (Edwards,  of i t s individuals  recognized  i s not standardized.  o f t h e 75 p e r c e n t  f r o m 75 p e r c e n t  recognized  75 p e r c e n t  application  (1962) up t o 2436 m e t e r s .  study  represented  an a l t i t u d e  t o 2538 m e t e r s .  t h e r e were a montane morph a t what a l t i t u d e w o u l d Dottrens  (1962) s u g g e s t e d  a lower  t o 1500 m b e c a u s e h i s b i g g e s t n o n - j u v e n i l e that a l t i t u d e .  Jones  tane  began around  subspecies  (1970) s a i d  range  limit  i t be  o f 1200 m  s p e c i m e n s came  from  t h a t t h e r a n g e o f t h e mon-  1900+ m, b u t e v e n  Jones'  77 largest  group,  a t 1940  m,  was  s m a l l e r than  g r o u p a t 1200  m t o 1500  m.  in  showed no  size-altitude  this  study  evidence It  o f a montane morph i s interesting  the v a l l e y  (altitude  was  of  1500  m)  M.  arvalis  and  d e s c r i b e d by  summit o f S t . G o t t h a r d was  not  specified  m t o 2100  near  u n s p e c i f i e d but type  but  that  arvalis  of Urseren,  the  s u b a d u l t and  the  page type  largest  specimens and  d e s c r i b e d by  gave  no  74). locality  St. Gotthard,  Schinz  of  the  in  1845  U r i , Switzerland  i t c o u l d have been  locality  i n the  range  o f t h e montane s u b s p e c i e s  Selys-Longchamps Pass,  adult  correlation  (Table XVII:  to note  montane s u b s p e c i e s o f M. was  The  Dottrens'  i n 1841  was  near  U r i , Switzerland (again  of the  altitude  i t c o u l d have been i n the range  of  2000  m) .  To d a t e a d e f i n i t e  range  f o r a montane morph h a s  still  to  be d e f i n e d . 2.3.  Nomenclature Although  Dottrens  call  their  cus,  there i s another  of  M.  tories .  Lehmann  (1967) and  Jones  (1970)  montane s u b s p e c i e s M i c r o t u s a r v a l i s r u f e s c e n t e f u s -  arvalis  incertus.  (1962),  name by w h i c h a S w i s s  is called  Tables XVIII  i n the and  XIX  literature list  brief  montane  - Microtus  subspecies arvalis  nomenclatural  his-  TABLE X V I I I : BRIEF NOMENCLATURAL HISTORY OF MICROTUS ARVALIS RUFESCENTEFUSCUS  Hypudaeus  rufescentefuscus  Synonymized w i t h A r v i c o i a a g r e s t i s fusca Synonymized w i t h Pitymus subterraneus  subterraneus  Synonymized w i t h Microtus a r v a l i s rufescentefuscus  Type l o c a l i t y : v a l l e y o f Urseren, U r i , S w i t z erland  Schinz  ^600  F a t i o (1900)  m i n Swiss A l p s  (1845)  Belgium and n o r t h e r n France eastward through Switzerland t o Transylvania  Miller  France, Belgium. S w i t zerland, t o Yugoslavia and T r a n s y l v a n i a  E l l e r m a n and S c o t t (1946)  y  D o t t r e n s (1962) Meylan (1966) Lehmann (1967) Jones (1970)  1200 m i n Swiss A l p s  y 11900 m i n Swiss A l p s Referred to Microtus a r v a l i s  REFERENCE  DISTRIBUTION  NOMENCLATURE APPLIED  (1912)  Corbet (1978)  TABLE XIX: BRIEF NOMENCLATURAL HISTORY OF MICROTUS ARVALIS INCERTUS NOMENCLATURE APPLIED Arvicola incertus  Type l o c a l i t y : near summit o f S t . Gotthard Pass, U r i , Switzerland  Synonymized w i t h Arvicola arvalis fulva Synonymized w i t h Microtus incertus  Synonymized w i t h Microtus a r v a l i s incertus Synonymized w i t h Pitymus subterraneus i n c e r t u s  REFERENCE  DISTRIBUTION  Selys-Longchamps  (1841)  F a t i o (1869)  Mountains o f S w i t z e r l a n d and T y r o l from t h e c e n t r a l A l p s eastward  Miller  Mountains o f S w i t z e r l a n d and Tryol  R e g n i e r and P u s s a r d (1926)  Switzerland  (part), to Tyrol  (1912)  Ellerman  (1940)  Ellerman  and S c o t t  D o t t r e n s (1961)  (1946)  80 CONCLUSIONS  Size-altitude  variation  museum s p e c i m e n s o f The  l a c k of  result is  of  that  enced  geographic  so  many  species.  exist  nor  sively  date.  There of  size  trend  group(s)  of  organisms  level(s)  at  which  physiological these the to the  major  trend which  to  are  among  called which  revised  the  may  likely  rarely, of  been  are by  Microtus  influ-  a l l , in arvalis  on  every  major  - on  the  the  taxonomic  i t s regularity.  applies  appellation "rule"  nonUntil  a clear definition  should  the  taxon  i s shown t o be  does  conclu-  i t s p h y s i o l o g i c a l or  of  cut  diagnostic  Bergmann's Rule  and  the  clear  demonstrated  a majority  definition  thesis.  explanation  i f at  i t a p p l i e s , on  422  have been  expressed  biologists  resolved and  for this  i t s morphological has  i n the  common v o l e  subspecies  i n t e r p r e t a t i o n ( s ) , on  i s formulated,  trend,  only  i t applies,'on  conflicts  the  i n the  i t s distribution  the  A more  variation  neither  i s disagreement  examined  encountered  occur  I f a montane  character to  may  i n evidence  data.  factors that  in Switzerland,  aspect  variation  features  patterns  not  arvalis  d e f i c i e n c i e s i n the  morphological this  Microtus  morphological  by  was  (or adhere  discarded.  of taxa) to  APPENDIX  1  NAMES OF I N S T I T U T I O N S , T H E I R A D D R E S S E S , KEY THE NUMBER OF S P E C I M E N S OF MICROTUS A R V A L I S 1.  i .  i i .  2.  i .  Musee d ' H i s t o i r e N a t u r e l l e Geneve, CH-1211 , S w i t z e r l a n d Head,  Birds  and  Mammals  Department  i .  B u n d n e r N a t u r h i s t o r i s c h e s und N a t i o n a l p a r k m u s e u m C h u r , CH-7000, S w i t z e r l a n d D r . J. M u l l e r , D i r e c t o r 63 S p e c i m e n s  i .  i .  i i .  6.  Baud,  99  i i .  5.  F.  i i .  ii.  4.  AND  S t a t i o n F e d e r a l e de R e c h e r c h e s A g r o n o m i q u e s Nyon, CH-1260, S w i t z e r l a n d D r . A. M e y l a n , H e a d , V e r t e b r a t e Department 189 S p e c i m e n s  Dr.  3.  PERSONNEL FROM EACH  i .  i i .  Specimens  N a t u r h i s t o r i s c h e s Museum B e r n B e r n , CH-3000, S w i t z e r l a n d D r . P. L u p s , D i r e c t o r 52 S p e c i m e n s  Zoologisches Museum d e r U n i v e r s i t a t Z u r i c h , CH-8006, S w i t z e r l a n d D r . C. C l a u d e , D i r e c t o r 15 S p e c i m e n s  Zurich  Zoologisches Forschungsinstitut Alexander Koenig 5 3 0 0 B o n n 1, G e r m a n y D r . R. H u t t e r e r , D i r e c t o r 4 Specimens  Museum  und  APPENDIX TABLE l a : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS ( I N MM) 4 STANDARD ERROR: TOTAL LENGTH OF SKULL  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  GR321  20.200 4 .076  .655  2.  GR33  22.400 4 .058  .446  3.  GR472  22.000 4 .050  .394  4.  GR314  22.683 * .044  .337  5.  GR485  23.233 ± .044  .329  6.  GR364  22.317 4 .034  .259  7.  5248  23.533 4 .034  .245  8.  GR36  20.650 4 .029  .242  9.  GR297  21.950 4 .029  .228  10.  5252  23.050 4 .029  .217  11.  5246  24.600 4 .029  .203  12.  5245  24.650 4 .029  .203  13.  5251  25.550 4 .029  .196  14.  GR514  23.217 4 .017  .124  15.  GR223  23.683 4 .017  .122  16.  5247  23.617 4 .017  .122  17.  5249  24.117 4 .017  .120  18.  5307  24.083 4 .017  .120  19.  344  24.017 4 .017  .120  20.  5305  24.133 4- .017  .120  21.  5308  24.267 * .017  .119  22.  589  25.133 4 .017  .115  23.  586  25.767 4 .017  .112  24.  GR39  23.950 * .000  .000  25.  588  23.450 4 .000  .000  APPENDIX TABLE l b : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: NASAL WIDTH  COEFFICIENT OF VARIATION (IN PERCENT)  2.467 ± .102  7.119  5246  2.683 i .093  5.990  GR297  2.400 4 .076  5.512  GR485  2.367 * .060  4.398  5308  2.400 * .050  3.608  GR472  2.233 * .044  3.420  7.  GR39  2.367 4 .044  3.227  8.  GR36  2.383 * .044  3.205  9.  589  2.750 * .050  3.149  10.  5248  2.433 * .044  3.139  11.  GR364  2.333 4 .034  2.474  12.  5247  2.383 * .034  2.423  13.  5307  2.433 4 .034  2.373  14.  5305  2.433 * .034  2.373  15.  5245  2.483 4 .034  2.325  16.  5251  2.667 4 .034  2.165  17.  GR223  2.450 * .029  2.041  18.  GR33  2.217 * .017  1.302  19.  GR314  2.333 * .017  1.237  20.  588  2.433 i .017  1.185  21.  5249  2.517 4 .017  1.147  22.  GR514  2.583 * .017  1.118  23.  344  2.583 4 .017  1.118  24.  GR321  2.000 4 .000  0.000  25.  5252  2.150 * .000  0.000  1. 2. 3. 4. 5.  586  APPENDIX TABLE l c : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION 5307 1.  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) ± STANDARD ERROR: ROSTRUM WIDTH  COEFFICIENT OF VARIATION (IN PERCENT)  4.167 i .120  4.996  2.  GR472  3.850 4 .087  3.896  3.  5252  3.733 i .060  2.788  4.  5247  3.983 4 .060  2.613  5.  GR485  4.117 4 .060  2.528  6.  344  4.250 4 .058  2.353  7.  GR321  3.733 ± .044  2.046  8.  5248  3.867 4 .044  1.975  9.  5246  4.500 i .050  1.925  10.  GR36  3.983 4 .044  1.917  11.  GR297  4.017 4 .044  1.902  12.  GR223  4.067 4 .044  1.878  13.  GR514  4.167 ± .044  1.833  14.  5249  4.267 4 .044  1.790  15.  589  4.567 4 .044  1.673  16.  GR314  3.933 4 .034  1.468  17.  GR33  4.067 4 .034  1.420  18.  5245  4.183 4 .034  1.380  19.  GR364  3.850 ± .029  1.299  20.  586  4.483 4 .034  1.288  21.  GR39  4.200 4 .029  1.191  22.  5308  4.250 4 .029  1.177  23.  588  4.167 ± .017  0.693  24.  5305  4.283 4 .017  0.674  25.  5251  4.433 * .017  0.651  APPENDIX TABLE I d : MEASURING ACCURACY SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEAN OF 3 MEASUREMENTS ( I N MM) i STANDARD ERROR: ZYGOMATIC WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  GR39  13.450 i .076  .984  2.  5245  13.550 ± .076  .976  3.  5305  13.117 i .073  .959  4.  GR297  12.333 4 .067  .936  5.  5246  14.067 * .067  .821  6.  588  13.933 ± .060  .747  7.  GR321  10.817 ± .044  .706  8.  GR485  12.650 4 .050  .685  9.  5248  12.700 4 .050  .682  10.  GR314  12.367 * .044  .618  11.  GR514  13.483 4 .044  .567  12.  5247  13.233 4 .034  .436  13.  5249  13.583 4 .034  .425  14.  GR364  12.050 4 .029  .415  15.  GR472  12.200 4 .029  .410  16.  GR33  12.500 i .029  .400  17.  5307  13.550 4 .029  .369  18.  5251  14.250 4 .029  .351  19.  589  14.850 i .029  .337  20.  5252  12.267 4 .017  .235  21.  GR223  1 3 . 2 1 7 * .017  .218  22.  5308  13.683 4 .017  .211  23.  586  14.833 4 .017  .195  24.  GR36  11.200 * .000  .000  25.  344  13.000 * .000  .000  APPENDIX TABLE l e : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) * STANDARD ERROR: LEAST INTERORBITAL WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  589  3.100 4 .104  5.815  2.  586  3.317 4 .060  3.138  3.  GR297  3.200 4 .058  3.125  4.  GR485  2.967 4 .044  2.575  5.  5251  3.067 t .044  2.491  6.  5249  3.117 4 .044  2.451  7.  GR36  2.767 4 .034  2.087  8.  5248  3.033 4 .034  1.903  9.  5252  3.117 4 .034  1.853  10.  GR472  3.183 4 .034  1.814  11.  GR39  2.850 4 .029  1.754  12.  588  3.333 4 .034  1.732  13.  GR223  2.900 sr .029  1.724  14.  5246  2.950 4 .029  1.695  15.  5247  2.950 4 .029  1.695  16.  GR514  3.000 4 .029  1.667  17.  5308  3.100 4 .029  1.613  18.  5245  3.250 4 .029  1.539  19.  GR33  2.983 4 .017  0.968  20.  5307  3.017 4 .017  0.957  21.  344  3.017 4 .017  0.957  22.  GR314  3.217 4 .017  0.897  23.  GR321  3.100 4 .000  0.000  24.  GR364  3.000 4 .000  0.000  3.200 4 .000  0.000  25.. 5305  APPENDIX TABLE I f : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: BRAINCASE WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  GR33  9.950 ± .076  i.330  2.  GR314  10.083 t .073  1.248  3.  5246  10.717 ± .073  1.174  4.  GR39  10.167 i .067  1.136  5.  588  9.717  .060  1.071  6.  GR485  9.783 ± .060  1.064  7.  5249  10.200 i .058  0.980  8.  589  10.700 - i .058  0.935  9.  586  11.150 4 .050  0.777  10.  GR364  10.183 4 .044  0.750  11.  5251  10.883 4 .044  0.702  12.  GR297  10.033 4 .034  0.575  13.  5305  10.183 ± .034  0.567  14.  5308  10.417 i .034  0.554  15.  344  10.667 4 .034  0.541  16.  5247  9.850 4 .029  0.508  17.  GR472  10.150 4 .029  0.493  18.  GR514  10.550 4 .029  0.474  19.  GR36  9.317  .017  0.310  20.  GR321  9.333 4 .017  0.309  21.  5248  9.883 4 .017  0.292  22.  5252  10.083 4 .017  0.286  23.  GR223  10.133 * .017  0.285  24.  5307  10.483 * .017  0.275  25.  5245  10.783 * .017  0.268  4  4  APPENDIX TABLE l g : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: POST ZYGOMA WIDTH  COEFFICIENT OF VARIATION (IN PERCENT)  1.  GR364'  8.783 t  .093  1.830  2.  5308  9.933 4 .083  1.453  3.  GR36  8.783 ± .073  1.433  4.  GR485  8.883 ± .073  1.417  5.  GR297  9.033 ± .060  1.152  6.  5305  9.483 4 .060  1.098  7.  GR472  9.350 4 .058  1.070  8.  5248  9.500 ± .058  1.053  9.  5245  10.200 4 .058  0.980  10.  GR39  9.300 4 .050  0.931  11.  5307  9.700 4 .050  12.  586  9.900 4 .050  13.  5246  10.150 i  .050  0.853  14.  5247  9.317 i  .044  0.820  15.  5252  9.317 4 .044  0.820  16.  GR314  9.333 4 .044  0.818  17.  589  9.933 4 .044  0.769  18.  344  9.967 4 .044  0.765  19.  5251  10.167 ± .044  0.751  20.  GR223  9.133 4 .034  0.632  22.  GR321  8.650 4 .029  0.578  22.  588  9.050 4 .029  0.553  23.  GR514  9.700 4 .029  0.516  24.  GR33  9.333  *  .017  0.309  25.  5249  9.633 •4 .017  0.300  -  0.893 0.875  APPENDIX TABLE l h : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: MASTOID WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  589  9.500 4 .087  1.579  2.  344  9.967 4 .073  1.263  3.  5307  9.567 4 .067  1.207  4.  5308  9.967 4 .067  1.159  5.  5245  10.217 4 .067  1.130  6.  588  9.100 4 .058  1.099  7.  5248  9.667 4 .060  1.077  8.  5251  10.083 4 .060  1.032  9.  5249  10.083 i .060  1.032  10.  GR472  8.850 4 .050  0.979  11.  GR321  8.117 4 .044  0.941  12.  GR364  8.783 4 .044  0.870  13.  586  9.833 4 .044  0.777  14.  GR485  8.767 4 .034  0.659  15.  GR223  8.933 4 .034  0.646  16.  5305  9.667 4 .034  0.597  17.  5247  9.817 4 .034  0.588  18.  5246  10.233 4 .034  0.564  19.  GR314  9.150 4 .029  0.546  20.  GR36  8.833 4 .017  0.327  21.  GR297  8.833 4 .017  0.326  22.  GR33  9.083 4 .017  0.318  23.  5252  9.233 4 .017  0.313  24.  GR514  9.317 * .017  0.310  9.317 4 .017  0.310  25. • GR39  APPENDIX TABLE l i : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) i STANDARD ERROR: FORAMEN MAGNUM HEIGHT  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  586  3.233 4 .093  4.971  2.  GR472  3.167 4 .088  4.824  3.  GR314  3.633 4 .088  4.204  4.  GR485  2.900 4 .058  3.448  5.  GR364  3.117 4 .060  3.340  6.  5249  3.000 4 .058  3.333  7.  5248  3.267 4 .060  3.186  8.  588  3.367 4 .060  3.092  9.  5252  3.533 4 .060  2.946  10.  GR297  3.350 4 .050  2.585  11.  5246  3.167 4 .044  2.412  12.  GR39  3.317 4 .044  2.303  13.  GR321  3.317 4 .044  2.303  14.  5305  2.933 4 .034  1.969  15.  5308  3.033 4 .034  1.903  16.  5307  3.083 4 .034  1.873  17.  344  3.217 4 .034  1.795  18.  GR36  2.800 4 .029  1.786  19.  5247  3.283 4 .034  1.758  20.  5251  3.100 4 .029  1.613  21.  5245  3.100 4 .029  1,613  22.  GR33  3.200 4- .029  1.563  23.  GR514  3.500 4 .029  1.429  24.  589  3.650 4 .029  1.370  25.  GR223  3.400 ± .000  0.000  APPENDIX TABLE l j : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) * STANDARD ERROR: FORAMEN MAGNUM WIDTH  COEFFICIENT OF VARIATION (IN PERCENT)  1.  GR485  3.400 i .100  5.094  2.  GR223  3.650 i .104  4.940  3.  5248  3.500 * .087  4.286  4.  GR33  3.667 4 .088  4.166  5.  GR321  3.833 4 .088  3.985  6.  GR36  3.650 i .076  3.624  7.  GR39  3.517 i .073  3.578  8.  586  3.567 a .067  3.238  9.  5252  3.833 4 .067  3.012  10  5245  3.583 4 .060  2.905  11.  GR472  3.667 4 .060  2.839  12.  5305  3.600 4 .058  2.778  13.  GR514  3.833 i .060  2.715  14.  5251  3.567 4 .044  2.141  15.  5249  3.683 4 .044  2.074  16.  GR314  3.867 4 .044  1.975  17.  GR364  3.033 4 .034  1.903  18.  5247  3.333 4 .034  1.732  19.  589  3.667 4 .034  1.575  20.  588  3.450 4 .029  1.450  21.  5307  3.600 i .029  1.389  22.  GR297  3.750 4 .029  1.333  23.  5246  3.850 * .029  1.299  24.  344  3.567 4 .017  0.809  25.  5308  3.583 4 .017  0.806  APPENDIX TABLE l k : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: SKULL HEIGHT  COEFFICIENT OF VARIATION ( I N PERCENT)  5252  6.450 i .076  2.051  5246  6.850 i .076  1.931  5251  7.350 4 .058  1.361  5248  6.600 4 .050  1.312  5307  5.983 4 .044  1.302  GR36  6.650 4 .050  1.034  GR472  6.283 4 .044  1.276  8.  GR514  6.417 4 .044  1.216  9.  5308  7.183 4 .044  1.190  10.  588  5.583 4 .034  1.063  11.  GR321  5.683 4 .034  1.016  12.  GR364  6.033 4 .034  0.957  13.  GR223  6.333 4 .034  0.912  14.  5247  6.967 4 .034  0.829  15.  586  7.033 4 .034  0.821  16.  589  7.033 4 .034  0.821  17.  344  6.100 4 .029  0.820  18.  GR297  6.200 4 .029  0.807  19.  GR485  6.300 4 .029  0.794  .20.  GR39  6.500 4 .029  0.769  21.  5245  6.950 4- .029  0.719  22.  GR314  6.283 4 .017  0.459  23.  GR33  6.367 4 .017  0.453  24.  5305  6.883 4 .017  0.419  25.  5249  7.017 4 .017  0.411  APPENDIX TABLE 11: MEASURING ACCURACY SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEAN OF 3 MEASUREMENTS ( I N MM) i STANDARD ERROR: CONDYLOBASAL LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  5251  25.150 4 .050  .344  2.  GR485  22.983 4 .044  .332  3.  GR36  19.917 i .034  .290  4.  GR33  21.717 i .034  .266  5.  GR314  22.167 4 .034  .261  6.  5248  23.133 4 .034  .250  7.  GR39  23.467 4 .034  .246  8.  5308  23.633 4 .034  .244  9.  5252  22.350 4 .029  .224  10.  5247  23.100 4 - . 029  .217  11.  GR321  19.717 4 .017  .146  12.  GR297  21.517 4 .017  .134  13.  GR364  21.717 4 .017  .133  14.  GR514  22.817 4 .017  .127  15.  GR223  22.933 4 .017  .126  16.  588  23.133 4 .017  .125  17.  GR297  23.767 4 .017  .122  18.  5305  23.667 4 .017  .122  19.  5246  23.933 4 .017  .121  20.  589  24.517 4 .017  .118  21.  586  25.183 4 .017  .115  22.  GR472  21.145 4 .000  .000  23.  5249  23.800 4 .000  .000  24.  5307  23.600 4 .000  .000  25.  5245  24.150 4 .000  .000  APPENDIX TABLE lm: MEASURING ACCURACY SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEAN OF 3 MEASUREMENTS ( I N MM) 4 STANDARD ERROR: BASAL LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  GR33  20.617 ± .093  .780  2.  5307  21.933 i .088  .696  3.  GR472  20.150 4 .076  .657  4.  586  23.733 4 .073  .530  5.  5245  22.617 4 .067  .511  6.  5252  20.800 4 .058  .481  7.  5248  21.683 4 .060  .480  8.  GR321 .  18.450 4- .050  .469  9.  344  21.900 4 .058  .457  10.  5251  23.650 4 .058  .423  11.  GR314  20.800 4 .050  .416  12.  GR514  21.400 4 .050  .405  13.  588  21.700 4 .050  .399  14.  5249  22.250 4 .050  .389  15.  5246  22.750 4 .050  .381  16.  589  23.100 * .050  .375  17.  5305  22.167 4 .044  .345  18.  GR36  18.850 * .029  .265  19.  GR485  21.550 4 .029  .232  20.  5308  22.200 4 .029  .225  21.  GR297  20.333 4 .017  .142  22.  GR364  20.617 4 .017  .140  23.  5247  21.783 4 .017  .133  24.  GR223  21.867 * .017  .132  25.  GR39  21.950 4 .000  .000  APPENDIX TABLE I n : MEASURING SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: BASILAR LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  5246  21.500 4 .132  1.066  2.  5251  22.583 i .093  0.712  3.  5248  20.650 4 .076  0.641  4.  GR39  20.633 * .073  0.610  5.  GR364  19.817 4 .067  0.583  6.  586  22.750 4 .076  0.582  7.  GR36  17.883 4 .060  0.582  8.  GR321  17.700 4 .058  0.565  9.  5307  20.767 4 .067  0.556  10.  5252  19.800 4 .058  0.505  11.  5249  21.183 4 .060  0.491  12.  GR297  19.600 4 .050  0.442  13.  GR33  19.750 i .050  0.439  14.  5305  21.000 4 .050  0.412  15.  5308  21.100 4 .050  0.410  16.  GR472  19.383 4 .044  0.394  17.  588  20.717 4 .044  0.369  18.  5245  21.383 4 .044  0.357  19.  GR314  20.067 4 .034  0.288  20.  GR514  20.567 4 .034  0.281  21.  GR485  20.517 ± .034  0.281  22.  5247  20.717 4 .034  0.279  23.  GR223  20.800 4 .029  0.240  24.  344  20.967 4 .017  0.138  25.  589  21.883 * .017  0.132  APPENDIX TABLE l o : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: PALATAL LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  12.183 4 .060  .854  13.900 4 .058  .719  12.350 4 .050  .701  13.100 4 .050  .661  13.017 4 .044  .587  13.317 4 .044  .574  13.617 4 .044  .561  13.767 4 .044  .555  12.367 4 .034  .467  1.  GR364  2.  5246  3.  GR33  4.  GR485  5.  588  6.  5308  7.  5307  8.  5245  9.  5252  10.  GR514  12.567 4 .034  .459  11.  GR223  13.167 4 .034  .439  12.  5247  13.183 4 .034  .438  13.  5249  13.417 ± .034  .430  14.  5305  13.683 4 .034  .422  15.  GR472  12.200 4 .029  .410  16.  344  12.950 4 .029  .386  17.  5248  13.150 4 .029  .380  18.  GR321  11.117 4 .017  .260  19.  GR36  11.517 4 .017  .251  20.  GR297  11.883 4 .017  .243  21.  GR314  12.317 4 .017  .234  22.  GR39  13.117 4 .017  .220  23.  589  13.617 4 .017  .212  24.  586  14.217 •4 .017  .203  25.  5251  14.433 •4- .017  .200  APPENDIX TABLE l p : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION GR223  MEASURING  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: PALATAL BRIDGE LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  5.133 i .109  3.688  GR321  4.567 4 .093  3.520  5246  5.233 4 .073  2.404  GR39  4.933 4 .067  2.341  5305  5.417 4 .073  2.323  5251  5.700 4- .076  2.321  589  5.067 4 .067  2.279  GR33  4.450 4 .058  2.247  9.  GR364  4.500 ± .058  2.222  10.  5247  4.783 4 .060  2.176  11.  588  4.917 4 .060  2.117  12.  GR485  4.750 4 .050  1.823  13.  586  5.733 4 .060  1.815  14.  GR472  4.483 4 .044  1.704  15.  GR297  4.533 4 .044  1.685  16.  5249  5.317 t .044  1.437  17.  5308  5.317 i .044  1.437  18.  5245  5.117 4 .034  1.128  19.  5307  5.367 4 .034  1.076  20.  5252  4.700 4 .029  1.064  21.  344  5.000 4 .029  1.000  22.  GR36  4.283 4 .017  0.674  23.  GR314  4.667 4 .017  0.619  24.  5248  5.233 4 .017  0.552  25.  GR514  5.000 4 .000  0.000  APPENDIX TABLE l q : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION GR297 1.  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS ( I N MM) 4 STANDARD ERROR: PALATAL FORAMEN LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  3.650 4 .076  3.624  2.  5249  3.850 4 .076  3.436  3.  GR364  3.733 4 .073  3.371  4.  GR485  4.167 4 .067  2.771  5.  5248  4.300 4 .058  2.326  6.  GR472  3.850 4 .050  2.249  7.  GR33  3.950 4 .050  2.193  8.  GR223  3.917 4 .044  1.950  9.  GR39  4.133 4 .044  1.848  10.  GR514  3.817 4 .034  1.513  11.  344  3.817 4 .034  1.513  12.  5252  4.067 4 .034  1.420  13.  5245  4.283 4 .034  1.348  14.  GR314  3.850  .029  1.299  15.  5251  4.567 4 .034  1.264  16.  586  4.550 4 .029  1.099  17.  GR36  3.483 4 .017  0.829  18.  GR321  3.483 4 .017  0.829  19.  5308  4.017 4 .017  0.719  20.  5247  4.367 4 .017  0.661  21.  5307  4.417 4 .017  0.654  22.  5246  4.417 4 .017  0.654  23.  589  4.483 4 .017  0.644  24.  588  4.000 4 .000  0.000  25.  5305  4.200 4 .000  0.000  *  APPENDIX TABLE l r :  MEASURING  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: PALATAL FORAMEN WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  0.633 4 .060  16.434  5249  0.850 i .076  15.563  GR36  0.817 i .067  14.139  5308  0.833 4 .067  13.856  5246  0.917 4 .073  13.727  5251  0.883 4 .067  13.072  GR472  0.883 4 .060  11.783  GR321  0.750 4 .050  11.547  GR39  0.667 4 .044  11.456  5305  0.933 4 .060  11.152  11.  589  0.900 4 .058  11.111  12.  GR485  0.800 ± .050  10.825  13.  5252  0.717 4 .044  10.657  14.  GR223  1.000 4 .058  10.000  15.  5247  0.783 4- .044  9.750  16.  GR33  0.817 4 .044  9.352  17.  GR314  0.833 4 .044  9.165  18.  5245  0.867 4 .044  8.813  19.  5248  0.867 4 .044  8.813  20.  588  0.600 4 .029  8.333  21.  5307  1.033 4 .044  7.391  22.  GR514  0.700 4 .029  7.143  23.  GR297  0.850 4 .029  5.882  24.  344  0.817 ± .017  3.535  25.  586  0.833 4 .017  3.464  SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION 1. 2. 3. 4. 5. 6. 7. 8. 9. 10  GR364  APPENDIX TABLE I s : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) i STANDARD ERROR: MAXILLARY DIASTEMA LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  6.217 4 .093  2.585  7.233 4 .102  2.428  7.217 4 .073  1.744  6.333 4 .060  1.643  7.000 4 .058  1.429  7.967 4 .060  1.307  6.367 4 .044  1.200  6.433 4 .044  1.188  1.  GR36  2.  5245  3.  GR223  4.  5252  5.  5247  6.  586  7-.  GR297  8.  GR472  9.  5248  6.883 4 .044  1.110  10.  588  6.917 4 .044  1.104  11.  5249  6.917 4 .044  1.104  12.  5307  7.067  .044  1.081  13.  GR321  5.750 4 .029  0.870  14.  GR33  6.733 4 .034  0.858  15.  GR364  6.767 4 .034  0.853  16.  GR485  7.067 ± .034  0.817  17.  5308  6.550 4 .029  0.763  18.  GR39  7.050 4 .029  0.709  19.  344  7.100 i  .029  0.074  20.  5246  7.350 4 .029  0.680  21.  5251  7.550 4 .029  0.662  22.  GR314  6.483 4 .017  0.445  23.  GR514  6.883 4 .017  0.419  24.  589  7.617 4 .017  0.379  25.  5305  7.150 4 .000  0.000  APPENDIX TABLE I t : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION GR39 1.  MEASURING  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: MAXILLARY TOOTH ROW LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  5.333 4 .067  2.165  2.  5308  5.767 4 .067  2.002  3.  GR485  4.917 4 .044  1.553  4.  GR297  5.217 * .044  1.464  5.  5247  5.333 4 .044  1.432  6.  5307  5.617 4 .044  1.360  7.  5248  5.683 4 .044  1.344  8.  5245  5.867 4 .044  1.302  9.  GR33  5.117 4 .034  1.128  10.  GR314  5.317 4 .034  1.086  11.  GR514  5.467 4 .034  1.056  12.  5252  5.467 4 .034  1.056  13.  589  5.633 4 .034  1.025  14.  GR364  4.900 4 .029  1.020  15.  5246  5.733 4 .034  1.007  16.  GR223  5.100 i .029  0.980  17.  GR321  5.150 4 .029  0.971  18.  588  5.200 4 .029  0.962  19.  5251  6.133 4 .034  0.941  20.  5305  5.550 4 .029  0.901  21.  5249  5.750 4 .029  0.870  22.  344  5.750 4 .029  0.870  23.  GR36  4.917 * .017  0.587  24.  GR472  5.217 4 .017  0.553  25.  586  5.467 4 .017  0.528  APPENDIX TABLE l u : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION 5247 1.  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) * STANDARD ERROR: MAXILLARY OUTER MOLAR WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  4.117 ± .093  3.904  2.  344  4.017 4 .088  3.803  3.  5248  4.083 ± .060  2.549  4.  5245  4.183 4 .060  2.488  5.  GR364  3.533 ± .044  2.162  6.  5308  4.100 4 .050  2.112  7.  GR297  3.817 4 .044  2.001  8.  GR472  3.917 4 .044  1.950  9.  5251  4.217 4 .044  1.811  10.  5249  4.317 ± .044  1.769  11.  GR36  3.567 i .034  1.619  12.  588  3.967 4 .034  1.456  13.  GR321  3.650 4 .029  1.370  14.  GR39  3.650 4 .029  1.370  15.  589  4.267 i .034  1.353  16.  GR485  3.800 i .029  1.316  17.  5305  4.000 i .029  1.250  18.  GR33  3.683 4 .017  0.784  19.  GR314  3.717 4 .017  0.777  20.  GR223  3.883 4 .017  0.743  21.  GR514  3.917 4 .017  0.737  22.  5307  3.967 4 .017  0.728  23.  586  4.017 4 .017  0.719  24.  5246  4.167 4 .017  0.693  25.  5252  3.750 4 .000  0.000  APPENDIX TABLE l v : SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) i STANDARD ERROR: MAXILLARY INNER MOLAR WIDTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  5252  2.033 i .093  7.905  2.  GR36  2.067 4 .073  6.089  3.  5249  2.267 4 .073  5.551  4.  GR314  2.000 ± .058  5.000  5.  GR485  2.083 4 .060  4.996  6.  344  2.050 4 .058  4.878  7.  GR297  2.067 4 .044  3.696  8.  589  2.083 4 .044  3.666  9.  GR364  1.867 4 .034  3.093  10.  5248  2.033 ± .034  2.840  11.  GR39  2.067 4 .034  2.794  12.  GR321  1.800 4 .029  2.778  13.  5307  2.167 4 .034  2.665  14.  5251  2.217 i .034  2.605  15.  5305  2.233 4 .034  2.588  16.  GR514  2.200 4 .029  2.273  17.. GR33  1.833 4 .017  1.575  18.  588  2.017 4 .017  1.432  19.  586  2.017 4 .017  1.432  20.  GR223  2.083 4 .017  1.386  21.  5245  2.283 4 .017  1.264  22.  5308  2.317 4 .017  1.246  23.  5246  2.383 4 .017  1.211  24.  GR472  2.000 4 .000  0.000  25.  5247  2.300 4 .000  0.000  APPENDIX TABLE l w : MEASURING SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  ACCURACY  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: MANDIBLE LENGTH  COEFFICIENT OF VARIATION (IN PERCENT)  1.  GR485  13.167 * .088  1.160  2.  5252  13.567 i .088  1.126  3.  5251  14.783 i .093  1.087  4.  52A7  13.867 * .073  0.907  5.  GR31A  13.467 4 .067  0.858  6.  GR33  12.900 i .058  0.775  7.  GR297  13.000 4 .058  0.769  8.  586  15.233 4 .067  0.758  9.  GR36  12.100 4 .050  0.716  10.  5308  14.200 4 .050  0.610  11.  3AA  13.783 4 .044  0.554  12.  52A9  14.367 4 .044  0.532  13.  GRA72  13.133 4 .034  0.440  IA.  GR39  13.367 4 .034  0.432  15.  5307  13.967 * .034  0.413  16.  GR51A  13.800 4 .029  0.362  17.  588  14.000 * .029  0.357  18.  GR223  14.200 ± .029  0.352  19.  52A5  14.450 4 .029  0.346  20.  589  14.550 4 .029  0.344  21.  GR36A  12.683 4 .017  0.228  22.  52A8  13.383 4 .017  0.216  23.  5246  14.567 4 .017  0.198  24.  GR321  12.150 4 .000  0.000  25.  5305  14.050 * .000  0.000  APPENDIX TABLE l x :  MEASURING  ACCURACY  SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION  MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: JAW LENGTH  COEFFICIENT OF VARIATION ( I N PERCENT)  1.  5307  9.050 4 .104  1.992  2.  5251  9.533 4 .109  1.986  3.  344  9.417 i .088  1.622  4.  5308  9.533 4 .088  1.602  5.  5252  8.600 4 .076  1.538  6.  5247  8.700 4 .076  1.521  7.  5249  9.383 4 .073  1.341  8.  5248  9.167 4 .067  1.260  9.  589  9.367 i .067  1.233  10.  GR33  8.583 4 .060  1.213  11.  GR514  8.917 4 .060  1.168  12.  588  9.050 4 .058  1.105  13.  5305  8.750 i .050  0.990  14.  GR314  8.800 4 .050  0.984  15.  GR321  8.233 4 .044  0.928  16.  5246  9.417 4 .044  0.811  17.  586  9.833 4 .044  0.777  18.  5245  9.217 4 .034  0.626  19.  GR472  8.450 4 .029  0.592  20.  GR297  8.650 4 .029  0.578  21.  GR223  9.050 4 .029  0.553  22.  GR36  8.083 4 .017  0.357  23.  GR364  8.117 4 .017  0.356  24.  GR39  8.667 4 .017  0.333  25.  GR485  8.700 ir .000  0.000  APPENDIX TABLE l y :  MEASURING ACCURACY MEAN OF 3 MEASUREMENTS (IN MM) 4 STANDARD ERROR: MANDIBULAR TOOTH ROW LENGTH  COEFFICIENT OF VARIATION (IN PERCENT)  5251  6.100 4 .050  1.420  GR514  5.467 4 .044  1.397  GR297  5.233 4 .034  1.103  GR314  5.317 4 .034  1.086  GR36  4.700 4 .029  1.064  GR364  4.850 4 .029  1.031  5307  5.633 4 .034  1.025  8.  5248  5.767 4 .034  1.001  9.  5252  5.400 4 .029  0.926  10.  5305  5.450 4 .029  0.917  11.  586  5.500 4 .029  0.909  12.  5249  5.750 4 .029  0.870  13.  GR33  5.067 4 .017  0.570  14.  GR321  5.117 4 .017  0.564  15.  GR472  5.167 4 .017  0.559  16.  GR223  5.183 4 .017  0.557  17.  GR39  5.283 4 .017  0.546  18.  5247  5.367 4 .017  0.538  19.  5246  5.667 4 .017  0.509  20.  589  5.733 4 .017  0.504  21.  5245  5.833 4 .017  0.495  22.  GR485  5.000 4 .000  0.000  23.  588  5.300 4 .000  0.000  24.  5308  5.900 4 .000  0.000  25.  344  6.000 * .000  0.000  SPECIMENS IN DESCENDING MAGNITUDE OF THE COEFFICIENT OF VARIATION 1.  APPENDIX TABLE 2a; SEXUAL DIMORPHISM AND CORRELATION  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  MEAN OF MEASUREME NT (IN MM) * STANDARD ERROR  Total Length of Skull  Nasal Width  Rostrum Width  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  20.660 20.739 22.665 22.573 24.284 24.740  4 4 i 4  .288 .306 .979 . 110 i . 108 4 .162  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  17 18 128 81 72 31  15 13 109 74 64 25  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  17 18 128 81 72 31  16 18 122 78 70 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  2.369 * .045 2.408 * .072 2.561 .209 2.565 4 .024 2.810 * .034 2.745 4 .046  -.157 (32)  17 18 128 81 72 31  15 17 119 79 69 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.947 4 .053 4.015 * .058 .257 4.223 4.158 * .029 4.392 * .028 4.435 * .038  -.304 (30)  -.071 (26) .093 (181) -.538 (87)  -.021 (198) .236 (98)  .237 (196) -.192 (97)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000070 -.000028 -.000002 -.000004 000004 .000013  .000079 -.000202 -.000005 -.000020 .000005 .000013  .000180 -.000106 .000001 -.000018 -.000018 .000060  APPENDIX TABLE 2b^ SEXUAL DIMORPHISM AND CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  Zygomatic Width  Least Interorbital Width  Braincase Width  NUMBER OF SPECIMENS  NUMBER SEX MEASURABLE  AGE GROUP  MEAN OF MEASUREMENT (IN MM) i STANDARD ERROR  *  17 18 128 81 72 31  16 15 113 76 67 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  17 18 128 81 72 31  16 18 121 76 72 31  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.206 4 .050 3.219 % . 070 3.184 ± . 161 3. 178 .019 3.156 i .028 3.189 .031  17 18 128 81 72 31  14 12 112 75 65 29  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  9. 936 * . 102 9.833 * . 110 10.225 4 .351 10.161 i .043 10.496 4 .048 10.412 ir . 069  11.513 11.513 12.760 12.540 13.771 14.037  * *  .151 .181 . 716 .074 .074 . 119  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.001 .306 -,430  -.055 .034 -.154  .277 . 172 219  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000089 -.000049 .000000 -.000006 .000007 .000044  .000236 .000138 .000009 .000018 .000013 .000102  .000135 .000136 .000006 .000016 .000000 .00002 5  APPENDIX TABLE 2a  NAME OF MEASUREMENT  Post Zygoma Width  Mastoid Width  Foramen Magnum Height  SEXUAL DIMORPHISM AND CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NUMBER MEASURABLE  SEX  AGE GROUP  MEAN OF MEASUREMENT (IN MM) 4 STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  17 18 128 81 72 31  10 7 84 51 48 17  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  9. 185 4 . 133 9.050 * . 135 9.492 * .044 9.440 4 .043 9.564 4 .045 9.577 * .075  .341 (15)  17 18 128 81 72 31  13 12 105 69 63 22  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  8. 912 * 8. 792 4 9.388 4 9.396 49.598 •4 9. 766 4  17 18 128  10 11 100  F M F  Juvenile Juvenile Subadult  3.200 .088 3.305 4 .075 .026 3.219  81 72 31  65 56 23  M F M  Subadult Adult Adult  3.119 *• .028 3.266 4- .034 3.174 4 .046  NUMBER OF SPECIMENS  .099 .040 .397 .048 .045 .088  * *  . 141 (133) -.042 (63)  .284 (23) -.020 (172) -.454 (83)  -.399 (19) .407 (163) .376 (77)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000147 -.000296 -.000011 -.000044 -.000003 -.000094  -.000230 -.000135 -.000013 -.000025 -.000017 -.000022  .000063 -.000152 .000011 .000006 .000019 -.000007  APPENDIX TABLE 2d; SEXUAL DIMORPHISM AND CORRELATION  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  Foramen Magnum Width  Skull Height  Condylobasal Length  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  17 18 128 81 72 31  11 10 103 70 57 25  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.573 3.655 3.619 3.598 3.660 3. 612  17 18 128 81 72 31  13 14 111 74 66 28  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  17 18 128 81 72 31  14 13 109 72 64 25  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  * *  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.424 (19)  *  .072 .041 . 187 .028 .024 .033  6.162 i 6.204 4. 6. 678 * 6.574 4 7.029 4 7.093 4  .074 . 108 .339 .046 .052 .065  -.122 (25)  *  .311 .327 1.013 .125 . 106 . 150  .117 (25)  20.311 20.173 22.214 22.118 23.795 24.250  4  4  4-  4  *•  . 103 (171) .269 (80)  .280 (183) -.160 (92)  .093 (179) -.553 (87)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT .000134 -.000070 -.000000 -.000000 -.000002 -.000101  -.000317 -.000102 -.000010 -.000029 -.000016 -.000004  -.000064 -.000033 -.000003 -.000006 .000000 .000004  APPENDIX TABLE 2e; SEXUAL DIMORPHISM AND CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  Basal Length  Basilar Length  Palatal Length  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  NUMBER MEASURABLE  SEX  17 18 128 81 72 31  14 12 101 68 61 24  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  18.896 19.296 21.006 20.818 22.562 22.996  17 18 128 81 72  13 12 96 64 61  F M F M F  Juvenile Juvenile Subadult Subadult Adult  17. 808 *• .331 18.183 4 .394 19.932 4 . 101 19.738 . 120 21.419 * . I l l  31  24  M  Adult  21.829  17 18 128  13 14 119  F M F  Juvenile Juvenile Subadult  11.596 4- .206 11.679 * .181 12.704 * .591  81 72 31  74 69 29  M F M  Subadult Adult Adult  12.577 4- .0 71 13.618 4 .065 13.881 4 .105  NUMBER OF SPECIMENS  AGE GROUP  t .306 * .402 41.005 * .Ill * .111 4 .161  *  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.316 (24) .190 (167) -.513 (83)  -.294 (23) .199 (158)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000067 -.000044 -.000002 -.000005 .000004 .000003  -.000060 -.000042 -.000002 -.000004  -.471 (83)  .000005 .000006  -.116 (25)  -.000084 -.000037  . 180  .201 (191) -.477 (96)  -.000001 -.000008 .000009 .000038  APPENDIX TABLE 2 f ; SEXUAL DIMORPHISM AND CORRELATION  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  Palatal Bridge Length  Palatal Foramen Length  Palatal Foramen Width  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  17 18 128 81 72 31  13 13 119 72 68 28  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  4.827 t 4.785 % 5. 151 i 5.103 i 5.416 5.395 *  17 18 128 81 72 31  17 18 123 78 70 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.335 3.222 3.762 3.765 4.110 4.305  17 18 128 81 72 31  17 18 123 78 70 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  .  AGE GROUP  * * * * *  .906 .972 * .897 * . 910 * .931 * .920  .099 . 107 .332 .040 .063 .068  .099 .104 .319 .036 .035 .038  .035 .039 . 102 .013 .014 .022  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) . 113 (24) . 137 (189) .045 (94)  .266 (33) -.011 (199) -.721 (98)  -.427 (33) -.117 (199) .090 (98)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000140 -.000044 -.000007 -.000018 -.000011 .000025  -.000197 -.000085 -.000016 -.000023 -.000018 -.000154  .000151 -.000330 -.000025 -.000041 -.000051 -.000263  APPENDIX TABLE 2g: SEXUAL DIMORPHISM AND CORRELATION  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  MEAN OF MEASUREMENT (IN MM) t STANDARD ERROR  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  * .092 *• .121  17 18 128 81 72 31  17 18 122 79 71 30  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  5.859 5.997 6.592 6.584 7.250 7.378  17 18 128 81 72 39  17 17 127 80 72 31  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  5.079 5.041 5.395 5.386 5. 645 5.659'  * .069  Maxillary Tooth Row Length  17 18 128 81 72 31  17 17 126 81 71 27  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.891 3.985 4.015 4.041 4.151 4.152  *- .065  Maxillary Outer Molar Width  Maxillary Diastema Length  .391 4 .044 * .056 4- .078  *  .064 4 .263 4 .032 T .034 * .055  4 .075 * .211 *• .025 4 .030 4 .048  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.306 (33) .019 (199) -.279 (99)  . 140 (32) .031 (205) -.047 (99)  -.325 (32) -.113 (205) -.005 (96)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000188 -.000011 .000006 .000005 .000020 .000076  -.000143 -.000212 -.000022 -.000041 -.000017 -.000030  -.000029 -.000219 -.000014 -.000039 -.000023 -.000032  APPENDIX TABLE 2h; SEXUAL DIMORPHISM AND CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  Maxillary Inner Molar Width  Mandible Length  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  MEAN OF MEASURE MENT (IN MM) * STANDARD ERROR  17 18 128 81 72 31  17 17 125 78 71 27  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  2.077 4 2.124 * 2.082 4 2.137 * 2.170 *• 2.194 •i  .036 .043 . 149 .018 .023 .042  17 18 128 81 72 31  17 17 125 78 71 27  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  1. 818 •i 1.862 1. 930 * 1.905 * 1. 981 4 1. 957 *  .051 .068 . 180 .025 .032 .037  17 18 128 81 72 31  16 16 120 78 66 29  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  12.556 12.653 13.552 13.397 14.464 14.402  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.289 (32) -.336 (201) -.123 (96)  .178 (32) .125 (201) .094 (96)  .000020 .000012 -.000029 -.000032 -.000006 -.000000  -.000064 -.000274 .000000 -.000025 -.000016 -.000052  *• . 169 * .161  .147 (30)  .684 .058 * .080 *• . 123  .233 (196)  *  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT  .095 (93)  -.000099 -.000046 .000000 .000015 .000000 .000026  APPENDIX TABLE 2 i ; SEXUAL DIMORPHISM AND CORRELATION  NAME OF MEASUREMENT  Jaw Length  Mandibular Diastema Length  Mandibular Tooth Row Length  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  17 18 128 81 72 31  15 15 121 79 69 31  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  8.380 4 8.477 * 8. 825 8.791 9.317 9.295 i  .090 .080 .375 .041 .045 .072  -.294 (28)  17 18 128 81 72 31  14 15 121 79 68 29  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  3.321 3.373 3.438 3.456 3.670 3.705  •i  .055 .060 .202 .028 .034 .061  -.236 (27)  17 18 128 81 72 31  15 15 125 79 70 29  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  5.067 5.103 5.384 5.335 5. 641 5.591  * *•  .076 .070 .269 .030 .032 .048  -.130 (28)  * * *• *•  * *  .085 (198) .059 (97)  -.077 (198) -.119 (95)  .170 (202) .186 (97)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000124 -.000241 -.00000 7 -.000022 -.000008 .000024  .000061 -.000181 .000006 -.000008 .000011 .000013  -.000214 -.000182 -.000016 -.000036 -.000025 .000034  APPENDIX TABLE 2 j ; SEXUAL DIMORPHISM AND CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENTS  NAME OF MEASUREMENT  Body (Plus Head) Length  Tail Length  Hind Foot Length  NUMBER OF SPECIMENS  NUMBER MEASURABLE  SEX  AGE GROUP  MEAN OF MEASUREMENT (IN MM) 4 STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  17 18 128 81 72 31  15 17 106 72 46 22  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  73.267 73.058 90.019 88.667 104.065 108.364  17 18 128 81 72 31  15 17 106 72 .46 22  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  27.333 27.294 32.293 33.167 33.804 38.455  4 . 814 4 1.274 4 3.814 4 .477 * . 788 * 1.018  -.220 (176)  17 18 128 81 72 31  15 17 107 71 47 22  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  14.467 15.000 14.851 15.282 17.255 15.546  * 4 4 4 4 4  -.555 (30)  4 4  %  2.184 1.620 4.735 .630 1.706 1.024  . 133 .298 .859 .083 . 163 .205  .028 (30) .258 (176) -.432 (66)  .009 (30)  .898 (66)  -.527 (176) -.272 (66)  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT .000000 .000004 .000000 -.000000 .000000 .000002  -.000010 -.000009 .000000 -.000000 .000000 .000000  -.000045 -.000051 .000002 .000000 .000002 .000008  APPENDIX TABLE 2k; SEXUAL DIMORPHISM AND CORRELATION  NAME OF MEASUREMENT  Ear Length  Weight  NUMBER OF SPECIMENS  COEFFICIENT FOR ALTITUDE AND MEASUREMENTS MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  NUMBER MEASURABLE  SEX  AGE GROUP  17 18 128 81 72 31  10 15 67 44 34 16  F M F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  9.900 10.267 10.776 10.546 11.588 11.500  * * * * * i  17 18 128 128 81 72 31  14 12 97 97 67 36 19  F M F F M F M  Juvenile Juvenile Subadult Subadult Adult Adult  12.643 12.167 18.794 17.343 25.583 29.737  .958 .953 .466 .466 .484 4 .906 4 1.140  4-  * *4 *  . 180 . 300 . 129 . 132 . 208 . 354  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) -.376 (23) .233 (109) -.069 (48)  (gr) (gr) (gr) (gr) (gr) (gr) (gr)  .138 (24) >  3  3  5  (  1  6  2  )  CORRELATION COEFFICIENT FOR ALTITUDE AND MEASUREMENT -.000129 -.000049 .000000 .000005 -.000002 .000005  -.000025 -.000011 -.000000 -.000002  -.788 (53)  .000000 -.000004  APPENDIX TABLE 3a; AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  MEAN OF MEASUREMENT (IN MM) 4 STANDARD ERROR  20.705 * .017 22.647 4 .677 24.428 .086  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  Total Length of Skull  49 226 114  42 200 100  Juvenile Subadult Adult  Nasal Width  49 226 114  48 217 111  Juvenile Subadult Adult  2.382 .032 2.561 i . 154 2. 781 4 .255  4. 977*(263)  49 226 114  46 215 108  Juvenile Subadult Adult  3.999 4 .034 4.201 * .181 4.391 4 .218  4. 782*(305)  49 226 114  42 203 105  Juvenile Subadult Adult  11.570 .097 12.694 .496 13.845 4 .599  Least Interorbital Width  49 226 114  48 212 113  Juvenile Subadult Adult  Braincase Width  49 226 114  36 201 105  Juvenile Subadult Adult  Rostrum Width  Zygomatic Width  "'Significant  *  3.207 4 .033 3.181 4- . 117 3.159 4 . 195  9.929 *• .057 10.192 4 .261 10.485 * .379  11. 622*(240) 15. 704*(298)  7. 816*(326)  6.388*(321)  9.549*(243) 14.'152*(306)  906 (258) -1. 007 (323)  3. 970*(235) 5. 878*(304)  a t .001 l e v e l 00  APPENDIX TABLE 3b; AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  MEAN OF MEASUREMENT (IN MM) i STANDARD ERROR  Post Zygoma Width  49 226 114  25 145 71  Juvenile Subadult Adult  9. 168 9.468 9. 573  Mastoid Width  49 226 114  34 185 92  Juvenile Subadult Adult  Foramen Magnum Height  49 226 114  29 176 88  Foramen Magnum Width  49 226 114  Skull Height  Condylobasal Length  "'Significant  4  .066  * .302  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM) •3.873*(168)  *  .037  8.874 9.394 9.635  4  .069 .294 .039  6.968*(217)  Juvenile Subadult Adult  3.224 3.177 3.232  * *  .046 .184 .027  -.955 (203)  30 184 92  Juvenile Subadult Adult  3. 585 .037 3. 605 * . 153 3. 637 .018  .489 (212)  49 226 114  36 200 103  Juvenile Subadult Adult  6.214 T .057 6. 634 * .260 7. 028 * .387  6.375*(234)  49 226 114  40 196 100  Juvenile Subadult Adult  20.346 * .165 22.189 * .718 23.939 * .084  10.495*(234)  at .001 l e v e l  *•  4  * *  2.097 (214)  4.830 (275)  1.703 (262)  1.276 (274)  8.607*(301)  14.958*(294)  APPENDIX TABLE 3c; AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  Basal Length  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  49 226 114  40 183 96  Juvenile Subadult Adult  19.039 * .196 20.941 * .716 22.694 * .087  10.662*(221)  49 226 114  39 173 95  Juvenile Subadult Adult  17. 962 * .197 19.862 4 .724 21. 550 4 .091  10.635*(210)  49 226 114  39 209 109  Juvenile Subadult Adult  11. 669 * . 103 12. 672 4 .425 13. 705 -4 .525  9.290*(246)  Palatal Bridge Length  49 226 114  38 207 107  Juvenile Subadult Adult  4. 791 *• .055 5. 135 •4 .241 5. 397 4- .450  5.657*(243)  Palatal Foramen Length  49 226 114  49 218 111  Juvenile Subadult Adult  3. 2 64 *• .058 3. 767 •4 .230 4. 167 4- .264  9.004-(2 65)  Palatal Foramen Width  49 226 114  49 218 111  Juvenile Subadult Adult  .918 * .022 .903 * .074 .928 4 .114  -.838 (265)  Basilar Length  Palatal Length  "'Significant  a t .001 l e v e l  14.947*(277)  14.223*(266)  14.774*(316)  5.644*(312)  10.698*(327)  1. 878 (327)  APPENDIX TABLE 3d.- AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  Maxillary Diastema Length  49 226 114  49 218 112  Juvenile Subadult Adult  5. 904 4 .062 6.587 t .274 7. 285 4 .425  10.545*(265)  Maxillary Tooth Row Length  49 226 114  48 224 112  Juvenile Subadult Adult  5. 081 4: .042 5. 396 4 . 191 5. 640 * .267  6.922*(270)  Maxillary Outer Molar Width  49 226 114  48 223 108  Juvenile Subadult Adult  3. 902 4 .039 4. 021 4 . 154 4. 140 4- .237  3.140 (269)  Maxillary Inner Molar Width  49 226 114  48 219 108  Juvenile Subadult Adult  2. 037 •i .027 2. 095 4- . 116 2. 170 . 190  2.103 (265)  Maxillary Molar Width  49 226 114  48 219 108  Juvenile Subadult Adult  1. 867 * .034 1. 925 4 . 138 1. 970 4 .236  1.746 (265)  Mandible Length  49 226 114  45 214 105  Juvenile Subadult Adult  '•'Significant a t .001 l e v e l  12.652 * .103 13.505 * .447 14.452 * .619  14.298*(328)  7.396*(334)  4.317*(329)  3.564*(325)  1.751 (325)  7.883*(257) 12.294*(317)  APPENDIX TABLE 3e; AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  Jaw Length  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  49 226 114  44 216 110  Juvenile Subadult Adult  8.414 * 8.810 * 9.297 *  .047 .262 ,354  6.427*(258)  Mandibular Diastema Length  49 226 114  43 216 108  Juvenile Subadult Adult  3.326 * 3.446 *• 3.676 *  .031 . 156 .284  3.195 (257)  Mandibular Tooth Row Length  49 226 114  44 220 110  Juvenile Subadult Adult  5.091 * 5.363 *  .042 .190  5.849*(262)  5.618 *  .249  Body (Plus Head) Length  49 226 114  33 179 69  Juvenile Subadult Adult  73.242 * 1.277 89.453 * 3.925 105.464 * 1.202  15.221*(210)  Tail Length  49 226 114  33 179 69  Juvenile Subadult Adult  27.364 * .742 32.643 * 2.969 35.362 * .668  6.932*(210)  49 226 114  34 180 70  Juvenile Subadult Adult  14.882 *15.022 * 15.357 *  Hind Foot Length  - S i g n i f i c a n t at .001 l e v e l v  .206 .625 .128  10.942*(324)  7.689*(322)  7.934*(328)  16.358*(246)  4.293*(246)  .827 (212) 2.622 (248)  APPENDIX TABLE 3 f ; AGE DIFFERENCES NAME OF  NUMBER OF  NUMBER  AGE  MEASUREMENT  SPECIMENS  MEASURABLE  GROUP  49 226 114  27 111 50  Juvenile Subadult Adult  10.074 *• .185 10.685 * .941 11.560 4 - . 178  2.890 (136)  49 226 114  26 164 55  Juvenile Subadult Adult  12.423 * .666 ( g r ) 18.201 * 3.418 ( g r ) 27.018 4 .755 ( g r )  6.427*(188)  Ear Length  Weight  " ' S i g n i f i c a n t at .001 l e v e l  MEAN OF MEASUREMENT (IN MM) * STANDARD ERROR  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  4.748*(159)  12.017*(217)  APPENDIX TABLE 4: PROPORTIONAL RELATIONSHIPS WITH AGE IN CRANIAL MEASUREMENTS OF MICROTUS ARVALIS NAMES OF  NUMBER OF  NUMBER  AGE  MEASUREMENTS  SPECIMENS  MEASURABLE  GROUP  MEAN OF PROPORTION ( I N MM) 4 STANDARD ERROR  M a x i l l a r y Diastema M a x i l l a r y Tooth Row  I n t e r o r b i t a l Width M a x i l l a r y Tooth Row  Interorbital Jaw Length  Width  S k u l l Height Zygomatic Width  Zygomatic Width C o n d y l o b a s a l Length  *  T-VALUE FOR DIFFERENCE BETWEEN MEANS (WITH ITS DEGREES OF FREEDOM)  49 226 114  48 224 112  Juvenile Subadult Adult  .Oil 1. 168 1. 179 4 .162 1. 282 4 .148  .308(270)  49 226 114  48 224 112  Juvenile Subadult Adult  .619 4 .015 .554 4 .101 .556 4 .066  -2.847(270)  49 226 114  44 216 110  Juvenile Subadult Adult  .372 4 .009 .340 4 .061 .337 4 .039  -.224(258)  49 226 114  42 203 105  Juvenile Subadult Adult  .422 4 .035 .483 4 .098 .475 4 .128  2.306(243)  49 226 114  40 196 100  Juvenile Subadult Adult  .527 4 .024 .543 4 .091 .551 4 .013  .691(234)  4.099(334)  115(334)  -.272(324)  -.516(306)  .510(294)  125 LITERATURE  CITED  A d a m c z e w s k a - A n d r z e j e w s k a , K.A. a n d L. 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