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Evolutionary relationships among Peromyscus from the Georgia Strait, Gordon, Goletas, and Scott Islands… Thomas, Barry 1971

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EVOLUTIONARY RELATIONSHIPS AMONG PEROMYSCUS FROM THE GEORGIA STRAIT, GORDON, GOLETAS, AND SCOTT ISLANDS OF BRITISH.COLUMBIA, CANADA  by  BARRY THOMAS B.A.  (1967), M.A. (1968)  C a l i f o r n i a Sta.te C o l l e g e at F u l l e r t o n  A Thesis Submitted i n P a r t i a l F u l f i l m e n t of the Requirements f o r the Degree of Doctor of Philosophy In the Department of Zoology  We accept t h i s  t h e s i s as conforming  tp the r e q u i r e d  standard  THE UNIVERSITY OF BRITISH COLUMBIA JUNE  1971  In  p r e s e n t i n g  an  advanced  the  I  f o r  s h a l l  agree  s c h o l a r l y  by  h i s  o f  t h i s  w r i t t e n  t h e s i s  degree  L i b r a r y  f u r t h e r  t h i s  at  the  make  t h a t  f r e e l y  p e r m i s s i o n  may  r e p r e s e n t a t i v e s .  f o r  p a r t i a l  U n i v e r s i t y  it  p u r p o s e s  t h e s i s  in  be  It  f i n a n c i a l  f o r  g a i n  Department  The  U n i v e r s i t y  Date  6>  o f  B r i t i s h  Canada  8,  -  2  i  -  7/  of  C o l u m b i a ,  B r i t i s h  C o l u m b i a  f o r  e x t e n s i v e  by  the  u n d e r s t o o d  p e r m i s s i o n .  V a n c o u v e r  o f  a v a i l a b l e  g r a n t e d  is  f u l f i l m e n t  s h a l l  Head  be  r e q u i r e m e n t s  r e f e r e n c e  c o p y i n g  that  not  the  o f  agree  and  o f  my  I  t h i s  o r  a l l o w e d  w i t h o u t  that  study.  t h e s i s  Department  c o p y i n g  f o r  o r  p u b l i c a t i o n  my  ABSTRACT  My s t u d y  i s d i r e c t e d towards understanding  of e v o l u t i o n t a k i n g p l a c e among p o p u l a t i o n s  the p a t t e r n  of Peromyscus  i n h a b i t i n g the i s l a n d s of B r i t i s h Columbia.  A  morphological  and r e p r o d u c t i v e  a p p r o a c h , as v e i l  as k a r y o t y p e  standard  i s o l a t i o n a n a l y s e s , have shown t h a t a t a x o n o m i c r e v i s i o n o f t h i s f a u n a l group i s d e s i r a b l e .  I have p r o p o s e d s u c h a  revision. A discriminant analysis u t i l i z i n g  morphological  measurements r e v e a l s t h e e x i s t e n c e o f two phenotypes w i t h i n the i s l a n d s surveyed.  distinctive Breeding  studies,  i n v o l v i n g 78 p a i r s o f i n s u l a r P e r o m y s c u s , i n d i c a t e t h a t r e p r o d u c t i v e i s o l a t i o n e x i s t s b e t w e e n t h e two morph g r o u p s . F u r t h e r m o r e , when g i v e n t h e c h o i c e  t h e l a r g e and s m a l l morph  g r o u p s p r e f e r t h e company o f t h e i r  own  Extensive p h a l l i c v a r i a t i o n occurs  type. among  island  p o p u l a t i o n s b u t no s x i g g e s t i o n o f t a x o n o m i c r e l a t i o n s h i p s are  discernible. K a r y o t y p e a n a l y s i s r e v e a l s t h a t t h e r e a r e two d i s t i n c t  karyotypic patterns.  The k a r y o t y p e  morphotype c l a s s i f i c a t i o n s . number o f m e t a c e n t r i c low number. identical  c o r r e l a t e s w i t h ,the two  The l a r g e morph has a h i g h  chromosomes and t h e s m a l l morph has a  The h i g h m e t a c e n t r i c  to that possessed  number k a r y o t y p e i s  by the m o r p h o l o g i c a l l y  similar  P e r o m y s c u s s i t k e h s i s known t o i n h a b i t t h e i s l a n d s t o t h e  north  of t h e . s t u d y group.  o f the the  t o v  major d i v i s i o n s i s m i n i m a l .  karyotypic  considered  r e m a i n as P. The  as P_.  species  s i t k e n s i s and  The the  d e g r e e and  subspecific  morphs i s  l a r g e morph s h o u l d s m a l l phenotype  should  r a p i d i t y of e v o l u t i o n wi,thin t h e s e requires  taxa.  has  been i n d i c a t e d .  the  a d j a c e n t m a i n l a n d and  a, b r o a d e n i n g i n t h e  Change i n s u b s p e c i f i c  Additional studies  m a i n l a n d g r p u p s a t the  be  scope  nomenclature  of P e r o m y s c u s u p o n  V a n c o u v e r I s l a n d are  b e f o r e m e a n i n g f u l r e l a t i o n s h i p s can i n s u l a r and  that  maniculatus.  Peromyscus p o p u l a t i o n s of som$  level.  each  I have p r o p o s e d  d i f f e r e n c e s b e t w e e n t h e s e two  s i g n i f i c a n t a t the be  Karyotype v a r i a t i o n w i t h i n  required  e x p r e s s e d between  subspecific  level.  TABLE OF  Title  CONTENTS  Page  i  Abstract  i i  Table of Contents  iv  List  vi  of Tables  i n Text  L i s t of Figures  i n Text  v i i .  L i s t of Data T a b l e s , Appendix l a n d 2  viii  L i s t of Tables, Appendix 3  x,  List  D i s c r i m i n a n t Function Data,  of P l a t e s , K a r y o t y p i c  Data, Appendix 4  Introduction  1  Population Density, Habitat Selection Food P r e f e r e n c e s  R e s u l t s and  and  of I n s u l a r Peromyscus  M a t e r i a l s and  •  xi  Methods  Discussion  11 13 14  The  southern  islands  14  The  northern  islands  18  Winter  25  Food p r e f e r e n c e s  25  Conclus ions Morphological  and  27 Behavioral Variation i n  Insular Populations  of Peromyscus  M a t e r i a l s and M e t h o d s Characters R e s u l t s and  measured  Discussion  32 33 35 36  A" The c o n t i n e n t a l  sample  B: The s o u t h e r n  islands  C: The n o r t h e r n  islands  B r e e d i n g T e s t s and C o m p a r a t i v e P o s t n a t a l D e v e l o p m e n t o f I n s u l a r and I n t e r - i s l a n d H y b r i d Peromyscus Materials Results  and M e t h o d s  and  Discussion  The P h a l l u s and i t s B e a r i n g upon t h e C l a s s i f i c a t i o n of the I n s u l a r Peromyscus of B r i t i s h Columbia Materials Results  and M e t h o d s .  and  Conclusions  De s c r i p t i o n o f p h a l l i Karyotypic  Variation i n Insular  Materials  Peromyscus  and Methods  Bone marrow method Tissue Results  and  culture  method  Discussion  C o n c l u s i o n s : A S y s t e m a t i c R e v i e w o f Some I n s u l a r Peromyscus A p p e n d i x 1 - Measurement Peromyscus  data f o r i n s u l a r  A p p e n d i x 2 -q Body, t a i l and b o d y / t a i l r a t i o histogram data Appendix 3 - D i s c r i m i n a n t  analysis  Appendix 4 - Karyotypes of i n s u l a r Peromyscus Appendix 5 - Specimens examined Acknowledgements Literature  Cited  data  LIST OF TABLES IN TEXT Trap success f o r beach and i n l a n d t r a p  sites  Keats Island Peromyscus  1970.  survey,  Body lengths f o r Peromyscus Strait region.  winter  of the  Georgia  Average t a i l and foot measurements f o r 1936 and 1968-1970 c o l l e c t i o n s .  the  Degree and d i r e c t i o n of morphological change w i t h time. Mean body measurements f o r Peromyscus c o a s t a l region of B r i t i s h Columbia. Male and female p a i r i n g  of  the  arrangement.  Comparative c r a n i a l measurements of Cox and T r i a n g l e Peromyscus and t h e i r h y b r i d young. Test crosses to determine ultimate conditions.  breeding  Mate p r e f e r e n c e s . P h a l l i c specimens  examined.  P h a l l i c measurements f o r Peromyscus.  several  species of  Divergence between i s l a n d samples u t i l i z i n g the combined baculum and d i s t a l t r a c t length measurements. Nomenclature for  chromosome  types.  C o l l e c t i o n and sample s i z e data f o r karyotype study. Karyotype c h a r a c t e r i s t i c s f o r the of the B r i t i s h Columbia r e g i o n .  the  Peromyscus  Proposed r e v i s i o n of the systematics of some i s l a n d Peromyscus.  Vll  LIST. OF FIGURES I N 1  Map  2  Map o f t h e G o r d o n , G o l e t a s , of B r i t i s h Columbia.  3 4  TEXT  of B r i t i s h Columbia  Map o f t h e Columbia.  8 and  Scott  Islands 9  Georgia S t r a i t region,  British 10  C o l l e c t i o n l o c a t i o n s f o r the c o n t i n e n t a l samples of Peromyscus m a n i c u l a t u s gambeli and P. m. rubidus..'  34  5  Changes i n b o d y l e n g t h o f P.m.  georgiensis.  43  6  Changes i n t a i l  georgiensis.  43  7  Changes.in o c c i p i t o n a s a l l e n g t h of Peromyscus m. g e o r g i e n s i s .  8.  l e n g t h o f P.m.  R e l a t i o n s h i p s o f i s l a n d P e r o m y s c u s ag by d i s c r i m i n a n t a n a l y s i s . "  44  determined 49  1  9  D i s p e r s a l route  of a n c e s t r a l Peromyscus.  10  P o s t n a t a l g r o w t h i n w e i g h t f o r some i s l a n d h y b r i d Peromyscus.  11  The  12  Phallic  13  Comparative glans  14  Comparative b a c u l a  15  S o m a t i c m e t a c e n t r i c chromosome numbers f o r some w e s t c o a s t p o p u l a t i o n s o f P e r o m y s c u s .  p h a l l u s of Peromyscus m a n i c u l a t u s . characters.  53 and  59 67 68  lengths. lengths.  75 75 86  1A i  C r a n i a l measurements f o r t h e P e r o m y s c u s m a n i c u l a t u s gambeli sample.  101  2A  S i g n i f i c a n c e o f v a r i a t i o n b e t w e e n mean c r a n i a l measurements i n P. m, g a m b e l i .  102  C r a n i a l measurements f o r P. jn.- r u b i d u s sample.  103  3A 4A 5A 64 7A 8A  S i g n i f i c a n c e o f v a r i a n c e b e t w e e n mean c r a n i a l measurements i n P. m. r u b i d u s . C r a n i a l measurements P. m. g e o r g i e n s i s ,  .  , a l l c o l l e c t i o n s of 105  Body measurements f o r t h e 1968-11970 collection.  106  Body measurements f o r t h e p r e - 1 9 5 2 collection.  107  C r a n i a l measurements f o r P f - t n . g e o r g i e n s i s c o l l e c t e d i n 1 9 3 6 , 1 9 4 5 , a n d 1968^ !  9A  104  108  C r a n i a l measurements f o r a l l c o l l e c t i o n s +,aken f r o m t h e n o r t h e r n i n l a n d s .  109  10A  Body measurements f o r a l l c o l l e c t i o n s .  110  11A  C r a n i a l measurements f o r t h e 1936 a n d 1952 n o r t h e r n i s l a n d c o l l e c t i o n . C r a n i a l measurements f o r t h e 1968-1970 collection only.  12A 13A 14A 15A 16A 17A  I l l 112  C r a n i a l m e a s u r e m e n t s , maJes o n l y , a l l collections.  113  C r a n i a l measurements, females collections.  114  only, a l l  R e p r o d u c t i v e and d e v e l o p m e n t a l d a t a f o r i n t e r - i s l a n d h y b r i d Peromyscus.  115  Reproductive and developmental f i e l d conceived l i t t e r s .  115  data f o r  I s l a n d Peromyscus p h a l l i c c h a r a c t e r i s t i c s  116  18A  Mean p h a l l i c measurements f o r i n s u l a r Peromyscus.  3.9A  S i g n i f i c a n c e o f v a r i a n c e b e t w e e n mean b a c u l a measurements.  118  S i g n i f i c a n c e o f v a r i a n c e between d i s t a l t r a c t measurements.  119  2PA  and r a t i o s  117  total  APPENDIX 2 Body and t a i l measurements and b o d y / t a i l r a t i o s , f o r each of the study i s l a n d s . FIGURE  histograms  ISLAND  21A  Histogram data f o r  f  ..  22A  Doyle Hurst  23A  •• • • •  Balaclava  24A  Nigei  25A  Vansittart  26A.  ..Hope  27A  Cox • .  28A  Lanz  29A  Triangle  :  30A  Savary  31A  Texada  32A  .  Thormanby  33A  Boven  APPENDIX 3 Discriminant  analysis data.  TABLE 21A 22A  Discriminant  function  coefficients.  The d e v i a t i o n s f r o m t h e c o l l e c t i o n l o c a l i t y on t h e d i s c r i m i n a n t a n a l y s i s .  based  PLATE 1  Karyotype o f Peromyscus m a n i c u l a t u s g e o r g i e n s i s , Boyen I s l a n d .  2  Karyotype o f Peromyscus i s o l a t u s , Cox I s l a n d .  sitkensis  3  Karyotype o f Peromyscus isolatus, Nigei Island.  sitkensis  4  Karyotype o f Peromyscus d o y l e i , Doyle I s l a n d .  sitkensis  5  Karyotype o f Peromyscus m a n i c u l a t u s g e o r g i e n s i s , Texada I s l a n d .  6  Karyotype of Peromyscus m a n i c u l a t u s g e o r g i e n s i s , Hope' I s l a n d .  7  Karyotype of Peromyscus s i t k e n s i s . saxamans, H u r s t I s l a n d .  INTRODUCTION  The  significance  of geographic  isolation i n speciation  i s c o n t r o v e r s i a l and t h e s e a r c h f o r a n s w e r s has l e d t o t h e r a i s i n g o f many more q u e s t i o n s t h a n have e v e r b e e n a n s w e r e d . P. J . D a r l i n g t o n (1957) m a i n t a i n s t h a t i n s u l a r p o p u l a t i o n s a r e e v o l u t i o n a r y d e a d ends and a r e i n s i g n i f i c a n t i n t h e i r total  c o n t r i b u t i o n t o the d i v e r s i t y of the world's  species.  M a y r (1966) h a s an o p p o s i n g p o i n t o f v i e w , h y p o t h e s i z i n g t h a t i s l a n d s a r e b r e e d i n g g r o u n d s f o r new s p e c i e s a n d t h a t t h e s e new s p e c i e s have g r e a t l y i n c r e a s e d t h e p r e s e n t d a y animal  diversity.  A n i m a l s have d e m o n s t r a t e d  the a b i l i t y to d i s p e r s e  a c r o s s most b a r r i e r s , b u t a s u c c e s s f u l c r o s s i n g does n o t n e c e s s a r i l y mean t h a t c o l o n i z a t i o n w i l l and W i l s o n , 1 9 6 7 ) .  result  (MacArthur  I n d e e d , t h e r e a r e numerous e x a m p l e s o f  easy access of animals t o i s o l a t e d areas y e t u n s u c c e s s f u l in colonization there.  Hyla arborea i s found  Europe b u t n o t i n England c a n be f o u n d  (Lack, 1969).  ( J . M. T a y l o r , p e r s o n a l c o m m u n i c a t i o n ) . three-fifths  Rattus  on t h e P h i l i p p i n e s , C e l e b e s , New  and New G u i n e a b u t n o t on t h e m a i n l a n d  across exulans  Zealand,  of A u s t r a l i a I r e l a n d has o n l y  of the b i r d fauna found i n England.  In  F e r n a n d o Poo 30 p e r c e n t o f t h e b i r d f a u n a i s endemic w h e r e a s i n Z a n z i b a r o n l y 3 p e r c e n t of the a v i f a u n a i s endemic, y e t b o t h o f t h e s e i s l a n d s a r e t h e same d i s t a n c e f r o m t h e m a i n l a n d  (Lack, 1969).  I t i s obvious  cross b a r r i e r s determines  t h a t more t h a n t h e a b i l i t y  t h e d i s p e r s a l and  to  e v o l u t i o n of  many a n i m a l s . Within small founding  colonies genetic d r i f t  abput major changes i n s h o r t p e r i o d s of t i m e . case  i s t h a t of the Darwin F i n c h e s  (Lack, 1947). b i r d s found  The  One  it  d i v e r s i f i c a t i o n o f one  hundred s p e c i e s  upon the Galapagos I s l a n d s c o n t r a s t s w i t h  Each i s l a n d  the  Caribbean that  i n nature.  s t u d y of the Peromyscus i n h a b i t i n g the i s l a n d s i n  the G u l f of C a l i f o r n i a  ( L a w l o r , 1971)  has  o b t a i n e d a degree  o f c o m p l e x i t y s i n c e 18 r e c o g n i z e d s p e c i e s i n h a b i t t h e region.  of  the  s i t u a t i o n conveys the i m p r e s s i o n  i s a l a r g e s c a l e experiment The  classic  on t h e G a l a p a g o s I s l a n d s  l a c k of s p e c i e s d i v e r s i t y i n the a v i a n f a u n a . o f Islands.  can b r i n g  My  Gulf  study d i f f e r s from t h a t of L a v l o r ' s i n t h a t the  P e r o m y s c u s p o p u l a t i o n s i n v e s t i g a t e d have b e e n c o n s i d e r e d all  of t h e  Guiguet, The remaining  species maniculatus  Cowan,  1945;  1952). i s l a n d s o f B r i t i s h C o l u m b i a a r e one  of the  few  N o r t h A m e r i c a n l o c a t i o n s where n a t u r a l p o p u l a t i o n s  of P e r o m y s c u s c a n be upset  (McCabe and  found  e x i s t i n g i n a s t a t e not  o r c h a n g e d by human i n t e r v e n t i o n .  I s l a n d s at the entrance c o l o n i z e d by man,  The, San  unduly  Juan  t o P u g e t Sound have l o n g b e e n  and most o f t h e l a r g e r i s l a n d s a l o n g  c o a s t have b e e n l o g g e d .  There a r e , h o w e v e r m a n y  i s l a n d s i n the n o r t h e r n r e g i o n w h i c h s t i l l pristine condition.  the  smaller  exist i n a  Numerous s u b s p e c i e s o f t h e s p e c i e s P. m a n i c u l a t u s  have  been d e s c r i b e d from the i s l a n d s of the c o a s t of B r i t i s h Columbia  (Cowan, 1935; H a l l , 1938; McCabe and Cowan, 1945;  Cowan and G u i g u e t , 1 9 6 5 ) .  S u c h an a r e a i s i d e a l f o r a n  i n v e s t i g a t i o n i n t o the systematics of c l o s e l y allopatric  populations.  related  McCabe and Cowan (1945) e x p r e s s e d  c o n c e r n f o r the need o f g e n e t i c i n v e s t i g a t i o n o f t h i s g r o u p when t h e y  complex  stated,  " I t i s almost u n t h i n k a b l e t h a t an a n c i e n t l y d e r i v e d s t r a i n o f c o m p l e x and d i s t a n t o r i g i n r P e r o m y s c u s m a n i c u l a t u s 1 c o u l d have m i g r a t e d f r o m somewhere b e y o n d t h e i c e c a p p e d a r e a and c o n t i n u e d t o g e t i t s e l f marooned on so s m a l l and i n a c c e s s i b l e , a range w i t h o u t l e a v i n g a g e n e t i c t r a i l over t h e course of i t s m i g r a t i o n . " Dice  (1948) p o i n t e d o u t t h a t s i g n i f i c a n t  differences  e x i s t i n t h e body measurements o f mice t a k e n from  locations  10 m i l e s a p a r t on t h e same i s l a n d .  Dice  In this  study  r e c o r d e d v a r i a t i o n s i n e a r l e n g t h o f a b o u t 0.7 mm., i  in tail  •  l e n g t h o f 6.3 mm.,  and i n b o d y l e n g t h o f 2.5 mm.;  these  d i f f e r e n c e s a r e g r e a t e r t h a n t h o s e e x i s t i n g b e t w e e n some established continental  subspecies.  B e r r y (1964), i n h i s  a n a l y s i s p f t h e s m a l l mammal f a u n a o f B r i t a i n , p o i n t s o u t t h a t t h e m a j o r i t y o f r e c o r d e d s u b s p e c i e s o f Apodemus, M i c r o t u s , and C l e t h r i o n o m y s  are i s l a n d p o p u l a t i o n s t h a t  have b e e n d e s i g n a t e d as new f o r m s . and B i s h o p two  Y e t t h e work o f Delany  (i960) notes t h a t the only r e a l d i f f e r e n c e i n  subspecies of Clethrionomys  tooth characteristic.  Hall,  i s a s m a l l and v a r i a b l e  i n h i s 1938 s t u d y o f t h e  Peromyscus of the G e o r g i a i s l a n d s , developed  a degree of  i s o l a t i o n -' t a i l geographic  l e n g t h formula which demonstrates c o n s i s t e n t  v a r i a t i o n i n t h e body t o t a i l  However, H a l l r e g a r d e d to  ratio.  t h e d i f f e r e n c e as t o o i n s i g n i f i c a n t  deserve s u b s p e c i f i c r e c o g n i t i o n . It  seems t h a t t h e s i g n i f i c a n c e  inter-island  (1953).  known b y g e o g r a p h i c  concept of nomenclature  scheme a l l i s l a n d p o p u l a t i o n s w o u l d be v e r n a c u l a r and n o t by a b i o l o g i c a l  Such a n a r r a n g e m e n t w o u l d s i m p l i f y t h e p r o b l e m  of n o m e n c l a t u r e b u t w o u l d i m p l y no p h y l o g e n e t i c Return  t o such a pre-Linnean  allopatric  pattern.  s y s t e m w o u l d , i n my o p i n i o n ,  s e r v e no sound b i o l o g i c a l p u r p o s e . taxonomists  One  t h e s u g g e s t i o n p u t f o r t h b y W i l s o n a n d Brown  Under t h e i r  trinomial.  l e f t to the  of the i n v e s t i g a t o r .  c o u l d perhaps abandon t h e c l a s s i c a l accept  of the variance i n  s m a l l mammal p o p u l a t i o n s i s a m a t t e r  whims and p e r s o n a l p r e f e r e n c e s  and  length  What i s n e e d e d i s f o r  t o d e r i v e some u n i v e r s a l f o r m u l a e  dealing with  p o p u l a t i o n s a n d t o a d h e r e t o them d e s p i t e  personal  dogmas. The  o b j e c t i v e of t h i s  systematics cranial  i s to r e - i n v e s t i g a t e the  o f sOme i s l a n d P e r o m y s c u s b y u s i n g  characters not previously considered  the taxonomic e v a l u a t i o n of t h i s group.  of t h e m o r p h o l o g i c a l these  standard  a n d b o d y m e a s u r e m e n t s , a e u r y t o p i c a p p r o a c h , as w e l l  as b y s t u d y i n g c r y p t i c in  study  The s i g n i f i c a n c e  v a r i a t i o n e x i s t i n g w i t h i n and between  island populations  i sa chief focal point.  The  d i r e c t i o n o f r a d i a t i o n , a n d where p o s s i b l e , t h e c l i n e s i n r e l a t i o n s h i p s , e x i s t i n g w i t h i n t h i s group a r e a s c e r t a i n e d  by both m o r p h o l o g i c a l  and k a r y o t y p i c c h a r a c t e r s .  Determination  of k a r y o t y p i c v a r i a t i o n i s of prime i n t e r e s t f o r I b e l i e v e t h i s feature holds great p o t e n t i a l  i n deciphering  probable  e v o l u t i o n a r y pathways o f Peromyscus i n i s l a n d p o p u l a t i o n s . The  complexity of karyotypes  i n t h e genus P e r o m y s c u s  i s w e l l documented i n t h e l i t e r a t u r e b u t t h e p a p e r s o f A r a k a k i and S p a r k e s  ( 1 9 6 5 , a , b ) , Hsu a n d A r r i g h i  Ohno e t a l ( 1 9 6 6 ) , and S i n g h a n d M c M i l l a n  (1966, 1968),  (1966) e s t a b l i s h  tha,t many p o p u l a t i o n s o f P e r o m y s c u s a r e k a r y o t y p i c a l l y Karyotypic  analyses  unique.  of c o n t i n e n t a l populations are complicated  by the u n c e r t a i n t y o f s p e c i e s b a r r i e r s , w h i l e t h e degree o f s y m p a t r y , h y b r i d i z a t i o n , and i n t e r - p o p u l a t i o n r e l a t i o n s h i p s are  i n , most c a s e s unknown.  At the present  time  chromosome  p o l y m o r p h i s m w i t h i n a n y one t a x o n c r e a t e s a s t u m b l i n g f o r the taxonomist. truei  block  The k a r y o t y p i c v a r i a t i o n i n P e r o m y s c u s  ( H s u and A r r i g h i ,  1 9 6 8 ) i s one o f t h e f e w e x a m p l e s :  where k a r y o l o g i c a l d a t a s u b s t a n t i a t e t h e c o n c l u s i o n s o f a n i n v e s t i g a t i o n b a s e d on g r o s s m o r p h o l o g y . concluded  i n h i s s t u d y t h a t P. t r u e i  genetic l i n e s . karyotype  breakthrough  then t h i s could represent a  insular  rodents.  i s l a n d environment provides p o p u l a t i o n s t h a t are  unquestionably and  of  i n t h e s t u d y o f d i s p e r s a l p a t h w a y s and t a x p n o m i c  r e l a t i o n s h i p s among t h e s e The  ,  confirmed by  I f a d i v i s i o n i n the karyotypes  i s l a n d m i c e c a n be f o u n d  (l95l)  i s composed o f two  T h i s has been s u b s e q u e n t l y  patterns.  Hoffmeister  interisland  a l l o p a t r i c , w i t h chances o f h y b r i d i z a t i o n crosses being  minimal.  The  d i s t r i b u t i o n of Peromyscus i s not c o n t i n u o u s a l o n g  beaches;  these  instead,, there are d i s c r e t e p o p u l a t i o n s which  e f f e c t are d u p l i c a t i n g the a l l o p a t r i c continental species;  s i t u a t i o n found  among  T h i s l a t t e r p o i n t i n c r e a s e s the  of f i n d i n g chromosome p o l y m o r p h i s m ,  in  chance  i f i t e x i s t s , and  should  p r o v i d e an e x c e l l e n t c h e c k u p o n t h e i n h e r e n t s t a b i l i t y t h e chromosome c o m p l i m e n t . karyotypic  change has  or a d d i t i o n .  In a non-Robertsonian  of  system  t o come a b o u t by i n v e r s i o n ,  any  deletion,  T h i s t e n d s t o c o m p l i c a t e and r e t a r d  recombina-  t i o n s more t h a n i n a, R o b e r t s o n i a n s y s t e m where t r i s o m y c a n and does e x i s t t o c o u n t e r a c t most d i s c r e p a n c i e s o r irregularities  i n t h e genome.  A l t h o u g h any g r o s s  e x i s t i n g between k a r y o t y p e s would tend to reduce and  the l e v e l  of f e c u n d i t y may  can take p l a c e .  difference recombination  be l o w e r e d , s u c c e s s f u l  I f t h e r e i s any  crosses  s p e c i a l adaptation to a  new  c o m b i n a t i o n and d i f f e r e n t k a r y o t y p e , t h e n s t a s i p a t r y , the. e v o l u t i o n of a new 1967),  s p e c i e s , t h r o u g h k a r y o t y p i c change  c a n become a  My  reality.  s t u d y i n v o l v e s two  groups t o t a l l i n g locations.  The  (Whi;te,  geographically isolated  13 i s l a n d s and  i n v o l v i n g 30  insular  different  i s l a n d s v a r y i n s i z e from those h a v i n g a  y a r d s of s u i t a b l e b e a c h ( i . e . D o y l e ) , t o t h o s e p o s s e s s i n g many m i l e s of good h a b i t a t group,  o f 9 i s l a n d s and  spectrum  The  The  24 s t u d y l o c a t i o n s , has  of P e r o m y s c u s m o r p h o t y p e s and  t h e l i t e r a t u r e by race.  ( i . e . Texada).  s i x s u b s p e c i e s and  northern a wide  i s represented i n  one  as y e t un-named  s o u t h e r n i s l a n d g r o u p has b u t one  morphotype,  few  Peromyscus m a n i c u l a t u s g e o r g i e n s i s H a l l . islands  The f o u r  have b e e n t r a p p e d i n s u c h a way as t o have  representative  sample f r o m most o f t h e o b v i o u s  t y p e s , s u c h as t h e b e a c h , f o r e s t  southern a  habitat  and g r a s s l a n d s , i n o r d e r  %o t e s t f o r i n t e r - p o p u l a t i o n v a r i a t i o n and t h e p o s s i b i l i t y of h a b i t a t - s p e c i f i c k a r y o t y p e s , i f s u c h e x i s t .  I'  1  .  138  '  !  '  126  1  —  ' 121  BRITISH COLUMBIA  12?  Fig.  2  The  Gordon, G o l e t a s ,  and  Scott Islands  of B r i t i s h  Columbia.  Fig.  3  The G e o r g i a S t r a i t  region, B r i t i s h  Columbia.  !— 1  o  POPULATION DENSITY, HABITAT SELECTION AND OF  FOOD PREFERENCES  INSULAR PEROMYSCUS.'. .  M a i n l a n d p o p u l a t i o n s of P e r o m y s c u s m a n i c u l a t u s been the f o c u s of i n t e n s i v e  i n v e s t i g a t i o n f o r many  have decades.  E x c e l l e n t references p e r t a i n i n g to p o p u l a t i o n d e n s i t i e s , h a b i t a t s e l e c t i o n and f o o d p r e f e r e n c e s o f p o p u l a t i o n s a r e t o o numerous t o l i s t , Blair Hall  (1942,  1950,  1951), Dice  continental  however the works of  ( 1 9 3 2 , 1939,  1940a,b,  1941a),  ( 1 9 3 8 ) , and H a l l and K e l s o n (1959). are. of m a j o r .  importance  .  i n b r i n g i n g a b o u t an u n d e r s t a n d i n g of t h e  c o m p l e x i t y o f t h i s genus on t h e m a i n l a n d . have y e t t o be  Island populations  i n v e s t i g a t e d to a comparable degree.  Taxonomic  i n v e s t i g a t i o n s of t h e P e r o m y s c u s f a u n a have b e e n made  on  many, b u t by no means a l l , of t h e i s l a n d s f o u n d a l o n g  the  c o a s t of N o r t h A m e r i c a .  The  s t u d i e s a r e t h o s e o f Osgood  most n o t a b l e o f t h e (1901, 1909), B u r t  island  (1932),  McCabe and Cowan ( 1 9 4 5 ) , D i c e  ( 1 9 4 9 ) , and F o s t e r  It  s t u d i e s whether the  i s not apparent  from these  m o r p h o l o g i c a l v a r i a t i o n e x i s t i n g between these p o p u l a t i o n s i s a l s o a c c o m p a n i e d by niche  (1965). tremendous  island  similar variation i n  criteria. D u r i n g t h e summers o f 1968,  1969,  and 1970  upon 14 i s l a n d s , a l o n g t h e c o a s t o f B r i t i s h Bowen, Thormanby, T e x a d a and  Savary  I trapped  Columbia.  (see f i g s .  1, 2, 3)  i n h a b i t e d by P e r o m y s c u s m a n i c u l a t u s g e o r g i e n s i s a w h i c h has b e e n c h a r a c t e r i z e d as h a v i n g l i t t l e  are  subspecies  morphological  variation  (Hall,  1938).  d i s t a n c e between the northern island  The  m o n o t y p i c p h e n o t y p e and  the l a r g e  island populations contrasts with  sample  ( f i g . 2) where g r e a t  the  morphological  v a r i a t i o n occurs between g e o g r a p h i c a l l y c l o s e i s l a n d p o p u l a t i o n s . The and  southern  Savary,  has  i s l a n d g r o u p o f Bowen, Thormanby, T e x a d a ,  a m i l d c l i m a t e w i t h ah a v e r a g e r a i n f a l l  l e s s t h a n 60 i n c h e s p e r y e a r .  of  Ground c o v e r , a l t h o u g h  fairly  dense i n c e r t a i n a r e a s , does n o t h i n d e r one's p a s s a g e and l i n e s can, be The  laid  i n most a r e a s w i t h o u t undue  trap  difficulty.  s e c o n d g r o u p of i s l a n d s i n v e s t i g a t e d i s c l u s t e r e d  a r o u n d t h e n o r t h e r n t i p of V a n c o u v e r I s l a n d . H u r s t , B a l a c l a v a , N i g e i , V a n s i t t a r t and  These a r e  Hope I s l a n d s .  Doyle, Although  t h e y a r e g e o g r a p h i c a l l y c l o s e t o g e t h e r , t h e y have a number of m o r p h o l o g i c a l l y d i s t i n c t p o p u l a t i o n s of Peromyscus. area r a i n f a l l  c a n be  as h i g h as 160  inches a year  and  In  this  the  v e g e t a t i o n t a k e s on a t e m p e r a t e r a i n f o r e s t a p p e a r a n c e . u n d e r g r o w t h i s so l u x u r i a n t t h a t p a s s a g e i s d i f f i c u l t  The  and  t h e l a y i n g o f t r a p s anywhere o t h e r t h a n t h e b e a c h i s a v e r y time these  c o n s u m i n g and and  Consequently,  on  o t h e r n o r t h e r n i s l a n d s I have t r a p p e d h e a v i l y u p o n  beach areas The  hazardous undertaking.  and  only a limited fashion inland.  l a s t g r o u p of i s l a n d s s t u d i e d i s composed o f  the  S c o t t I s l a n d s , l o c a t e d 5 t o 20 m i l e s t o t h e w e s t o f Cape S c o t t , at  t h e n o r t h e r n t i p of V a n c o u v e r I s l a n d .  well-known f o r t h e i r adverse 1912,  These i s l a n d s  weather c o n d i t i o n s .  In  are  October,  f o r e x a m p l e , t h e r o o f o f t h e l i g h t h o u s e on T r i a n g l e  I s l a n d , t h e w e s t e r n m o s t of t h e S c o t t I s l a n d s , was  blown o f f ,  two  sheds d i s a p p e a r e d ,  a n d a l l smoke s t a c k s were b l o w n down.  Winds o f up t o 105 m.p.h. have b e e n r e p o r t e d  for this  island.  ( C a r l e t a l , 1950 p p . 2 1 - 6 3 ) . All are  d e s c r i p t i o n s of winter  derived  habitats for island  Peromyscus  s o l e l y from the southern i s l a n d s , f o r only  here  d i d t h e weather c o n d i t i o n s p e r m i t access and t r a p p i n g opportunity  at this  season of the year.  MATERIALS AND METHODS I found the g r i d t r a p p i n g procedures f o r t h e census o f small animal populations  ( D i c e , 1941b; B o l e ,  to implement i n the m a j o r i t y of i s l a n d vegetation  s i t u a t i o n s . The  and g e o l o g i c a l c o n t o u r s were t o o f o r m i d a b l e f o r  t h i s type of procedure. along  1939) i m p o s s i b l e  areas of uniform  approximately  Consequently, I placed  a l l traps  t e r r a i n and ground cover,  f i v e yard  intervals.  w i t h p e a n u t b u t t e r and o a t m e a l .  and a t  The t r a p s were b a i t e d  Towards t h e end o f t h e  t r a p p i n g p e r i o d I u s e d a b a i t f o u n d t o be j u s t a s e f f e c t i v e , yet  easier t o apply.  This b a i t c o n s i s t e d of peanut o i l w i t h  d i s s o l v e d peanut b u t t e r can.  The t r a p l i n e s  (50:1  m i x ) a p p l i e d by a gun type o i l  c o n s i s t e d o f Sherman 8" l i v e  Museum S p e c i a l snap t r a p s .  Stomachs f r o m t h e snap  s p e c i m e n s were removed and p r e s e r v e d a n a l y s i s of t h e i r  t r a p s and trapped  i nformalin for  later  content.  I have c o n s i d e r e d  d e n s i t y t o be d i r e c t l y r e l a t e d t o t h e  number o f i n d i v i d u a l s c a u g h t a t a n y one l o c a t i o n d u r i n g t h e first  t h r e e , or fewer, t r a p n i g h t s .  By u s i n g t h e r e s u l t s  of the  only the f i r s t  three trap nights  ( t a b l e l ) the e f f e c t of  d e p l e t i n g p o p u l a t i o n numbers u p o n t h e p e r c e n t t r a p  i s m i n i m i z e d y e t t h e c h a n c e s o f o b t a i n i n g an a d e q u a t e i s maximized.  return sample  I found t h a t a l i g h t r a i n brought about the  maximum t r a p c a p t u r e s and a c l e a r s k y d u r i n g t h e n i g h t p r o d u c e d a l m o s t no r e t u r n s .  F o r t u n a t e l y , i n the n o r t h e r n  i s l a n d g r o u p e v e n i n g r a i n s a r e t o be e x p e c t e d t h u s g u a r a n t e e i n g comparable weather c o n d i t i o n s f o r the d u r a t i o n of the t r a p p i n g period.  A l l m i c e o f t h e s o u t h e r n i s l a n d g r o u p were  d u r i n g p e r i o d s o f good w e a t h e r and u s u a l l y w i t h o u t rains.  trapped night  A l t h o u g h constant weather c o n d i t i o n s occurred f o r a l l  samples w i t h i n each i s l a n d group the weather v a r i a t i o n s b e t w e e n g r o u p s were p r o b a b l y n o t c o n d u c i v e t o t h e p r o c u r e m e n t of  maximum t r a p r e t u r n s i n e a c h c a s e .  RESULTS AND The  DISCUSSION  Southern Islands:  Bowen, Thormanby, T e x a d a , and S a v a r y .  On Bowen I s l a n d , P e r o m y s c u s were f o u n d s c a v e n g i n g a l o n g the  intertidal  the  entire,  zone.  Peromyscus  extensive beach area.  occurred r e g u l a r l y This s i t u a t i o n  along  contrasts  w i t h the a l l o p a t r i c n a t u r e of beach p o p u l a t i o n s i n the northern i s l a n d groups.  The h i g h e s t d e n s i t y o f P e r o m y s c u s  occurs along the l i t t o r a l  zone o f t h o s e g e n t l y  sloping  b e a c h e s h a v i n g a t r a n s i t i o n zone o f d r y u n d e r b r u s h b e t w e e n the  b e a c h and t h e c l i m a x The i n t e r t i d a l  forest.  zone i s r i c h i n p o t e n t i a l  f o o d s o u r c e s and I was  soon a b l e t o r e c o g n i z e  invertebrate potentially  BEACH  INLAND  # TRAP # ANIMALS % TRAP NIGHTS CAUGHT SUCCESS  ISLAND  # TRAP # ANIMALS % TRAP NIGHTS CAUGHT SUCCESS  DOYLE  780  8  1.0  120  HURST  74  5  6.8  N.T.  BALACLAVA  320  9  2.8  N.T.  NIGEI  320  12  3.7  90  15  VANSITTART  160  12  • 7.5  60  0  0  HOPE  520  13  2.5  60  0  0  84  17  20.2  N.T.  LANZ  136  32  23.5  120  0  0  BERESFORD  N.T.  40  0  0  TRIANGLE  80  25  31.3  40  5  12.5  BOWEN  120  11  9.1  60  0  0  THORMANBY  180  5  270  12  4.4  TEXADA  240  26  10.8  450  41  9.1  SAVARY  20  0  0  160  . 18  COX  2.75  0  0  16.5  11.2  (N.T. a r e a n o t t r apped) Table  1  T r a p s u c c e s s f o r b e a c h and i n l a n d t r a p  sites.  good t r a p debris.  s i t e s b y t h e amount and f r e s h n e s s of t h e b e a c h Brand  and Rykman ( 1 9 6 8 ) , i n t h e i r s t u d y o f P e r o m y s c u s  g u a r d i a of the I s l a Lorenzo  Sur M e x i c o , had n o t e d a s i m i l a r  h a b i t a t p r e f e r e n c e and have s u g g e s t e d e x i s t i n g upon the beach d e b r i s .  t h a t P.  guardia  was  I have s u b s e q u e n t l y f o u n d  by  stomach c o n t e n t a n a l y s i s , t h a t the Peromyscus i n h a b i t i n g the i s l a n d s of B r i t i s h Columbia intertidal  invertebrates.  are i n d e e d s u b s i s t i n g upon Cliff  areas, v e t underbrush  open g r a s s pr l o g g e d a r e a s on Bowen I s l a n d y i e l d e d  zones,  no  Peromyscus, Thormanby I s l a n d has v e r y few b e a c h e s c o m p a r a b l e t o t h e t y p e on Bowen I s l a n d t h a t h a r b o r s P e r o m y s c u s . promising region by  subdividers,  preferred  on Thormanby has  long  most  s i n c e been t a k e n  and homes o c c u p y what I b e l i e v e was  Peromyscus h a b i t a t .  o f Thormanby. i s p r e c i p i t o u s  Since the r e m a i n i n g  i t was  p e r c e n t t r a p success a l o n g the i n t e r t i d a l  coastline find 2.75  z o n e , and t h e  success from the i s l a n d i n t e r i o r  to the s o r t of r e t u r n s  The  over  once t h e  not s u r p r i s i n g to  t h a t P e r o m y s c u s d e n s i t i e s were e x t r e m e l y l o w .  percent trap  The  are  4.4  comparable  t h a t I would expect from the s i m i l a r  h a b i t a t t y p e on t h e o t h e r i s l a n d s , and I t h i n k i t e m p h a s i z e s the importance  of t h e a b u n d a n t f o o d s u p p l i e s  found  on  the  beaches. T e x a d a . I s l a n d , the l a r g e s t i s l a n d t r a p p e d i n t h i s has  a desolate  c o a s t l i n e w i t h many s e c l u d e d b a y s .  p e r c e n t t r a p s u c c e s s compares f a v o r a b l y r e c o r d e d f o r t h e Bowen i n t e r t i d a l y i e l d .  t o t h e 9.1 The  The  study, 10.8  percent  Peromyscus  density  i n the  island interior,  9.1  percent,  was  t h a n t h a t r e c o r d e d f o r e i t h e r c e n t r a l Bowen o r Islands.  In three  nights  of t r a p p i n g  the  extent,  about 8 p e r c e n t ,  logging  crews.  The floral an  edge of t h e  i n among t h e  four  r o a d and  d i v e r s i t y u p o n T e x a d a , and  i s l a n d and  s l a s h l e f t by  the  Thormanby I s l a n d i s a r o c k y ,  on t h e  f o r e s t and  rocky  two  as  a  vegetated  i n h a b i t e d by p e o p l e , seems  previously studied  cliffs  utilized  sparsely  f o r Peromyscus c o l o n i z a t i o n .  c o a s t , , as  greater  this diversity is attracting  Bowen, where i t i s n o t  t o o wet  of  at a l e s s e r  d i f f e r e n c e , I b e l i e v e , i s a t t r i b u t a b l e t o the  source.  t o be  Thormanhy  at a density  abundance o f i n s e c t s t h a t c o u l d p o s s i b l y be  food  greater  i n a logged area  m i l e s w e s t of V a n a n d a , P e r o m y s c u s o c c u r r e d a b q u t 10 p e r c e n t a l o n g  far  A l o n g the  islands,  Texada  climax  a c t as b a r r i e r s b e t w e e n P e r o m y s c u s  populations. Savary I s l a n d i s the for  i t i s b o r d e r e d by  most u n u s u a l o f t h e  a wide, white  coast  arid a s h e e r , s a n d y c l i f f  area,  o n l y a few  Gaultheria old  forest.  was. h i g h e r  islands.  The  has  f e r n species  Potential subdividers  of e c o l o g i c a l t u r m o i l .  studied  on t h e w e s t .  intervening  has  density  replaced  the  have made numerous p a t h w a y s  of t h e  h e r e t h a n on any  I f t h i s higher  and  Sallal,  i n so d o i n g t h e y have c r e a t e d The  east  been logged  A l d e r , A l n u s r u b r a Bong,  s h a l l o n P u r s h , and  t h r o u g h t h i s a r e a and  population  s a n d b e a c h on t h e  hundred f e e t i n w i d t h ,  a dense g r o w t h of Red  southern islands  state  Peromyscus  of the  d e n s i t y was  a  previously the  r e s u l t of  c r o w d i n g due t o d i s p l a c e m e n t  of mice from other h a b i t a t s  t h e n t h i s w o u l d p e r h a p s e x p l a i n t h e e x i s t e n c e o f two i n t h i s apparently panmictic  g r o u p (Thomas, 1 9 7 0 ) .  were t r a p p e d  i n the climax f o r e s t , grass  The  Islands:  Northern  Doyle,  karyotypes  No m i c e  l a n d or marsh  areas.  Hurst, Balaclava, N i g e i ,  V a n s i t t a r t , Hope, C o x , L a n z , B e r e s f o r d a n d T r i a n g l e . D e v e l o p m e n t on t h e s e present  time  lighthouse  i s l a n d s has been l i m i t e d and a t t h e  o n l y two p e r m a n e n t i n s t a l l a t i o n s e x i s t , a  on B a l a c l a v a a n d a D e p a r t m e n t o f T r a n s p o r t  s t a t i o n on Hope.  Two I n d i a n v i l l a g e s , now i n d e c a y i n g  a l o n g w i t h a few s i n g l e  i s l a n d s were once t h e home o f many I n d i a n  Since  these  people  were n o t a g r i c u l t u r a l l y  made r e l a t i v e l y l i t t l e  ruins,  isolated dwellings, indicate that  these  Consequently,  radio  oriented  modification to their  t h e s m a l l mammal f a u n a  l a r g e l y u n a f f e c t e d by these  families. they  environment.  has p r o b a b l y  remained  e a r l y human i n h a b i t a n t s .  Doyle I s l a n d Doyle I s l a n d , l o c a t e d three m i l e s n o r t h of P o r t Hardy, i s a t w i n p e a k e d mound o f r o c k w i t h a maximum e l e v a t i o n o f l e s s t h a n 400 f e e t .  T h e r e a r e no s a n d y b e a c h e s a n d t h e b e s t  boat l a n d i n g approach i s the north or south of l a n d b e t w e e n t h e two r o c k p e a k s . grassed The  and p r o v i d e s  easy access  r e s t of the i s l a n d  ground cover collected  This saddle  t o both  saddle  i s densely  s i d e s of the i s l a n d .  i s s p a r s e l y f o r e s t e d , w i t h s a l l a l as  i n t h e more open a r e a s .  on t h i s  slope of a  island  The f e w P e r o m y s c u s  ( n = 8 f o r 780 t r a p n i g h t s ) were  found around the r o o t s of t r e e s under the s a l l a l  on t h e  northside  of the g r a s s y saddle  rocky c l i f f  area.  T r a p s p l a c e d upon t h e  f a c e s and a r o u n d t h e s t e p i s l a n d c o a s t  line  y i e l d e d no P e r o m y s c u s . H u r s t and B a l a c l a v a These a r e f a i r l y  s m a l l i s l a n d s , d e n s e l y f o r e s t e d , and  w i t h many s t o n y b e a c h e s .  On b o t h  of these  s e c l u d e d b a y s d i d n o t p r o d u c e any m i c e . an abandoned  Indian v i l l a g e  even though evidence,  The one g r a s s y  area,  on B a l a c l a v a , y i e l d e d no m i c e  i n t h e f o r m o f numerous r u n s made b y  M i c r o t u s , c o u l d be f o u n d . p e r t a i n beach areas.  i s l a n d s the q u i e t ,  P e r o m y s c u s were t r a p p e d  I t was upon t h e s e  only  upon  i s l a n d s t h a t the  p r e f e r r e d h a b i t a t f o r P e r o m y s c u s was more c l e a r l y d e f i n e d . The p r o d u c t i v e a r e a s were gentle  i n many r e s p e c t s s i m i l a r t o t h e  s l o p i n g b e a c h e s o f Bowen I s l a n d .  These b e a c h e s a l w a y s  had a c o n s t a n t l y r e n e w e d s u p p l y o f s e a weed d e p o s i t e d  along  t h e h i g h w a t e r mark, and y e t were n o t s u b j e c t t o h e a v y s e a spray. both  P e r o m y s c u s seem t o a v o i d t h e c o n s t a n t l y damp a r e a s  on t h e b e a c h e s and u n d e r t h e s a l l a l  Trapping  records indicate a preference  covered  for well  slopes. drained,  or s m a l l r o c k b e a c h e s . Nigei Island The w e s t end o f N i g e i I s l a n d gave s i m i l a r t r a p p i n g d a t a to  t h a t o f B a l a c l a v a and H u r s t I s l a n d s .  Here a g a i n t h e w e l l  d r a i n e d b e a c h e s w i t h f r e s h s e a weed d e p o s i t e d u p o n  gentle  s l o p e s t h a t l e a d up i n t o t h e f o r e s t e d a r e a p r o d u c e d t h e g r e a t e s t number o f P e r o m y s c u s . Logging  o p e r a t i o n s have c l e a r e d much o f t h e e a s t e r n end  of N i g e i I s l a n d . have grown up  c r e a t i n g a dense b u t  logged slopes. occurred of t h e  V a c c i n i a , Rubus and  A l o n g the  at high density  traps  set along  Peromyscus.  This  levels.  the  Vansittart ... T h i s and  f o r the  S i x t e e n and  food  high  captured  increase  over  the  and be  i s l a n d l o c a t e d midway b e t w e e n N i g e i  A l l t w e l v e mice c a p t u r e d d u r i n g  of the  dense f o r e s t and  rocky  cliff  ( t a b l e l ) upon the  sampling the a fair  rocky  cliff  i s one  area,  region.  and  Since  b e a c h i n c l u d e d 40  i n d i c a t i o n of the  240  7.5  cliffs.  the  The  avoided  the  160  the  trap  trap nights  percent trap  prefered  On  sandy, w e l l  steep rocky  flat  zone, the  the  distinctive region.  i s l a n d there  m i c e were f o u n d t o i n h a b i t t h e  with  percent  sources are b e l i e v e d to  s l o p i n g b e a c h b o r d e r e d by  All  a half  density.  i s a, v e r y , s m a l l  southwest corner  i s not  trail  the  road Peromyscus  Plant diversity  t r a p n i g h t s were caught, i n a v e r y  nights  ground cover over logging  s i d e of the  sallal  Island  Hope- I s l a n d s .  drained  ever present  i s more t h a n a f o u r f o l d  invertebrate  responsible  low  edge of t h e  beach h a b i t a t t r a p success r a t e . associated  the  habitat  of  success  density.  P e r o m y s c u s on V a n s i t t a r t were h e a v i l y i n f e s t e d  ectoparasites.  f l e a s , t i c k s , and  T h i s was  The  only  i s l a n d l o c a t i o n where  e a r m i t e s were so a b u n d a n t on e a c h mouse  t h a t t h e y must have, r e p r e s e n t e d their hosts.  the  other  a l a r g e p h y s i c a l " l o a d " upon  i s l a n d s sampled i n t h i s  remarkably p a r a s i t e - f r e e Peromyscus.  study produced  Hope I s l a n d T h i s i s the westernmost i s l a n d I t i s an I n d i a n r e s e r v e and, i t was  of the Gordon group.  although  i t i s now  deserted,  once t h e home o f s e v e r a l I n d i a n f a m i l i e s .  t h a t was  c l e a r e d f o r t h e I n d i a n v i l l a g e has  f o r M i c r o t u s townsendi adjacent brush Trapping  and  success  f o r e s t i n t e r i o r y i e l d e d no  The  grassland,  Peromyscus.  f o r a l l b e a c h e s on t h e e a s t and  on Hope I s l a n d , was  land  become a h a v e n  but not f o r Peromyscus.  coast produced v e r y poor r e s u l t s .  southern  B u l l H a r b o r , my  camp  site  h e a v i l y t r a p p e d d u r i n g t h e summer of  Even:though a l l c r i t e r i a p r e v i o u s l y determined necessary  The  as  1969.  being  f o r P e r o m y s c u s c o l o n i z a t i o n seemed p r e s e n t ,  with  t h e p o s s i b l e e x c e p t i o n o f a d a i l y i n f l u x o f f r e s h seaweed, few  p l a c e s around t h i s n a t u r a l harbor  produced Peromyscus.  I b e l i e v e t h a t o i l p o l l u t i o n from the f i s h i n g boats  and  camp has  extermina-  resulted  i n the  t i o n of most i n t e r t i d a l  c o n t a m i n a t i o n and life  and  Peromyscus of t h e i r prime f o o d M o s t of t h e of t h e  t h i s has  deprived  the  source.  s p e c i m e n s were t a k e n f r o m t h e w e s t  i s l a n d ; h e r e P e r o m y s c u s were f o u n d  rock b l u f f  ultimate  coast  l i v i n g around  a r e a s where t i d e p o o l s f o r m e d a t low t i d e .  s p e c i m e n s were t a k e n f r o m t h e d r i f t - w o o d zone above s a n d y b e a c h b u t none f r o m t h o s e pf the P a c i f i c  Ocean.  The  i n d i c a t e d t h a t open b r u s h of B u l l H a r b o r b a y ,  and  fish  A  few  the  r e g i o n s exposed t o the  summer o f 1970  the  spray  trap record  a r e a s , away f r o m t h e p o l l u t e d  c e r t a i n p o i n t s along the  shore  southern  c o a s t l i n e y i e l d e d Peromyscus w i t h a t r a p success  o f around  8 percent. The  remaining  o f Cape S c o t t .  islands  o f t h e n o r t h e r n sample o c c u r  Each i s l a n d  a c t i o n of the P a c i f i c  i sfully  and p r o v e d  r e a c h a n d seemed e v e n h a r d e r  west  exposed to the f u l l  t o be a d i f f i c u l t  to leave.  place to  Sox and L a n z I s l a n d s ,  l o c a t e d a b o u t s i x m i l e s w e s t o f Cape S c o t t , a r e s e p a r a t e d b y a channel  l e s s t h a n 400 y a r d s w i d e .  This channel  i s , however  a very e f f e c t i v e b a r r i e r since the f a s t f l o w i n g t i d e s  through  t h i s passage keep the water i n a constant s t a t e o f t u r b u l e n c e M i n k ( M u s t e l a v i s o n ) i n t r o d u c e d u p o n L a n z by t h e P r e d r i c k s e n b r o t h e r s i n 1938 o r 1 9 3 9 , i s now common u p o n b o t h Lanz I s l a n d s , w h i l e r a c c o o n ,  (Procyon  Cox a n d  l o t o r ) , has remained  u p o n Cox I s l a n d . A disassembled were p r o b a b l y  Sherman L i v e t r a p a n d a n u p s e t  t h e work o f t h e r a c c o o n .  robbing raccoons  Experience  line  with trap-  o n T e x a d a I s l a n d d u r i n g t h e 1970 t r a p s e a s o n  t o g e t h e r w i t h t h e i n c i d e n t s on Cox I s l a n d , i n d i c a t e raccoons  trap  are p o t e n t i a l predators  of Peromyscus.  and t h e o c c a s i o n a l s c r e e c h o w l w o u l d be t h e o n l y  that  Raccoons predators  o f P e r o m y s c u s on many o f t h e n o r t h e r n , i s l a n d s . Female P e r o m y s c u s were t r a p p e d more f r e q u e n t l y t h a n males i n areas  o f dense c o v e r .  Three o f t h e s e v e n  t r a p p e d away f r o m t h e b e a c h h a d d e l i v e r e d t h e i r in the trap.  I t seems t h a t g r a v i d f e m a l e s  females  litter  while  about t o give  b i r t h move away f r o m t h e b e a c h a n d s e e k p r o t e c t i o n o f t h e underbrush.  This apparent  need f o r p r o t e c t i o n a p p l i e s o n l y  f o r the p e r i o d of p a r t u r i t i o n f o r l a c t a t i n g females captured  along  the i n t e r t i d a l  vere  zone as f r e q u e n t l y a s any  o^her s e x o r age c l a s s . Beresford  Island  This i s l a n d ,  l o c a t e d two a n d a h a l f m i l e s  southwest of  L a n z , a p p e a r s a s a s m a l l 320 f o o t r o c k p i n n a c l e . and  Bad weather  heavy s u r f p r o h i b i t e d the s e t t i n g of t r a p s around the  h i g h t i d e mark a n d t h e o n l y a l t e r n a t i v e was t o s c a l e t h e r o c k and s e t t r a p s a c r o s s grass  t h e 200 f o o t w i d e f l a t  top.  The  ( P e s c h a m p s i a c a e s p i t o s a ) was dense o v e r t h e t o p o f t h e  i s l a n d and t h e t h i n s o i l burrows of sea b i r d s . recovered  l a y e r was honeycombed w i t h t h e  O n l y a y o u n g C a s s i n A u k l e t was  from our t r a p l i n e  a n d I must c o n c u r w i t h t h e  s t a t e m e n t made b y t h e p r e v i o u s 1950)  scientific  party  ( C a r l e_t a l . ,  t h a t no s m a l l mammal a p p e a r s t o i n h a b i t t h e u p p e r  reaches of t h i s i s l a n d .  I t i s unfortunate  sample c o u l d n o t be o b t a i n e d  t h a t a beach  f o r such a s m a l l and i n h o s p i t a b l e  h a b i t a t must e x e r t a t r e m e n d o u s s e l e c t i v e p r e s s u r e P e r o m y s c u s and t h e g e n e t i c  upon t h e  c o n s e q u e n c e s w o u l d be w o r t h y o f  investigation. Triangle This treeless sallal  Island i s l a n d i s u n i q u e among t h e i s l a n d s s t u d i e d . I t s slopes are densely  and salmonberry.  overgrown w i t h wind-pruned  P e r o m y s c u s were f o u n d t o be  abundant i n a l l h a b i t a t s (31.3 p e r c e n t Furthermore, these  trap  success).  m i c e were a c t i v e d u r i n g t h e l a t e  as w e l l a s a t n i g h t .  afternoon  Twenty t r a p s l a i d a t 3 p.m. A u g u s t 24,  when c h e c k e d t h a t same a f t e r n o o n a t a r o u n d 4 p.m., three a d u l t mice.  A t no  Peromyscus a t t h i s time  o t h e r l o c a t i o n have I of t h e day.  i n h a b i t i n g the Santa B a r b a r a r e p o r t e d t o be  diurnaJ  (J.  and  However, P e r o m y s c u s  Smith p e r s o n a l  T h i s d i u r n a l p a t t e r n i s p o s s i b l y due pressure  trapped  Channel i s l a n d s are  D.  yielded  to the  also  communication). l a c k of  the h i g h p o p u l a t i o n d e n s i t y .  P e r o m y s c u s f r o m T r i a n g l e I s l a n d a r e l a r g e and a t y p i c a l l y m a n i c u l a t u s - l i k e i n appearance. (as i n d i c a t e d by h a v i n g to  t h e maximum s i z e  u p o n t h e t o p of t h e of t h e  predator  The  somewhat  very  e x t e n s i v e t o o t h w e a r and  old  belonging  c l a s s ) male P e r o m y s c u s were t r a p p e d i s l a n d a t an e l e v a t i o n o f 690  s i x specimens c a p t u r e d  the f i r s t  only  feet.  n i g h t i n the  Five  apex  t r a p l i n e were m a l e s o f t h e maximum s i z e c l a s s ; t h e r e i s an apparent absence of these d i s t r i b u t i o n c a n be and  m a l e s on t h e b e a c h .  considered  n o t a chance o c c u r r e n c e  If  this  a natural biological  then t h i s i s the f i r s t  phenomenon reported  c a s e of s e n i o r c i t i z e n s e g r e g a t i o n among P e r o m y s c u s .  Under  l a b o r a t o r y c o n d i t i o n s P e r o m y s c u s f r o m T r i a n g l e I s l a n d behave d i f f e r e n t l y from those amicable  islands.  to other i n d i v i d u a l s from t h e i r  localities.  encounters  own  They a r e more and  other  P e r o m y s c u s f r o m T r i a n g l e a t t e m p t t o mate  minutes a f t e r being  my  o f most o t h e r  introduced to a strange  were n e v e r s e e n t o be  observations  non-Triangle  female.  consummated a t t h a t  suggest t h a t i n a l l cases  i t was  p a r t n e r w h i c h b r o k e away f r o m t h e  island just Although  time,  the interaction.  This  a m i c a b i l i t y i s s i m i l a r t o the  the Peromyscus from the California.  M i x e d sex  Santa Barbara Channel  Only i n t h i s  been observed l i v i n g  latter  of the  c a s e have m i x e d s e x  f o r m among  groups  conditions.  laboratory  northern' i n s u l a r Peromyscus.  Another-distinguishing Triangle  Islands,  i n harmony u n d e r l a b o r a t o r y  communal g r o u p s do n o t  population  c o n d i t i o n I n o t e d among  feature  of t h e  Peromyscus  I s l a n d i s t h e i r h a b i t of b u r r o w i n g b e n e a t h the  of s a w d u s t i n . t h e i r l a b o r a t o r y  cage.  A one  inch layer  wood s h a v i n g s i s , s u f f i c i e n t t o o b s c u r e them f r o m t h e glance.  on  I t seems t h a t t h e  l a c k of t r e e s has  s e m i - f o s s o r i a l mode of e x i s t e n c e which i s learned available  and  not  for inter-island  of  casual  brought about a  i n these animals,  i n h e r i t e d according  layer  a  t o the  trait data  crosses.  Winter Guiguet. ( p e r s o n a l the w i n t e r  c o m m u n i c a t i o n ) has  speculated  Peromyscus m i g r a t e i n l a n d i n order  i s b e l i e v e d t o be  I s l a n d , Howe S o u n d , B r i t i s h C o l u m b i a , i n d i c a t e d  as  i n the  interior  It  i n the  northern  Keats  that  same d e n s i t y d u r i n g  summer months ( t a b l e 2 ) . of the  beaches  i n F e b r u a r y on t h e b e a c h of  Peromyscus i n h a b i t the beaches i n the  in  t o evade what  a d v e r s e weather c o n d i t i o n s upon the  However, a t r a p l i n e s e t up  winter  that  E a r l y May  i s l a n d s p r o d u c e d no  the  trapping mice.  seems t h a t t h e s e i n s u l a r P e r o m y s c u s have a d a p t e d t o a y e a r  round beach environment. Food  Preferences The  discontinuous  d i s t r i b u t i o n of P e r o m y s c u s  populations  ELEVATION  # Peromyscus CAUGHT " • •'  -  1  1 1  1  — ,'• 1  % ANIMAL MATTER I N GUT r •••  500ft.  -\S  02  15  300ft.  OS  22  25  100ft,  5^  22  20  beach  63  52  40  Table 2  K e a t s I s l a n d Peromyscus s u r v e y w i n t e r , 1970  along the to the  c o a s t l i n e of these  food  available.  i s l a n d s seems t o be  directly related  As p r e v i o u s l y s t a t e d , a number o f  s p e c i m e n s were s n a p - t r a p p e d on t h e b e a c h e s o f e a c h i s l a n d i n a l l cases the r e s u l t s of the stomach content same.  Eighty percent  of the stomach c o n t e n t s ,  sample, c o n s i s t e d of animal was  Colepptera  material.  The  a n a l y s i s were f o r the  spp.).  b e a c h e s w i t h d a i l y i n f l u x e s of f l o t s a m a r e more l i k e l y o f amphipods and  m a i n a t t r a c t i o n and  this  f o r the  this Those  to  i s b e l i e v e d t o be  means o f d i e t a r y s u p p o r t  harbor the  Peromyscus.  CONCLUSIONS The  w i n t e r o f 1968-1969 was  n a t u r a l animal the  severe  and  p o p u l a t i o n s were a d v e r s e l y  long l a s t i n g . affected  r e s u l t i n g low p o p u l a t i o n d e n s i t i e s a r e  by t h e  s p o r t h u n t e r r e t u r n s f o r the  P e r o m y s c u s were p r o b a b l y i n 1969 density.  Insular  approximate the minimal  and  population  t r a p p i n g l o c a t i o n s , d u r i n g t h i s p e r i o d of  w e a t h e r c o n d i t i o n s , may environment.  season.  documented  a f f e c t e d i n a s i m i l a r manner  t r a p p i n g r e c o r d may The  1969  The  1970  Many  and  clearly  severe  a l s o i n d i c a t e t h e most f a v o r a b l e t r a p p i n g r e s u l t s and  s i g h t i n g records  s u g g e s t a s i g n i f i c a n t l y h i g h e r d e n s i t y o f P e r o m y s c u s as w e l l as o f o t h e r mammals s u c h as m i n k , o t t e r a n d B a l a c l a v a and  Doyle I s l a n d s provided  and  trap success,  8 percent  of J u l y i n 1970.  deer.  a 10 p e r c e n t ,  r e s p e c t i v e l y d u r i n g the  the  summer  major p o r t i o n of  l a r v a e and a m p h i p p d s ( O r c h e s t r o i d e a  large populations  and  Hope, 12  percent  month  T h i s . i s c o n s i d e r a b l y above t h e t r a p  success  recorde'd  f o r 't^he •same, a r e a d u r i n g  the  W e a t h e r c o n d i t i o n s s i n c e 1968 1970  trap r e s u l t s probably  population and  level.  Goletas  island-groups  season ( t a b l e l ) .  have- b e e n m i l d , and  represent  If this  1969  a more n a t u r a l  i s t h e :case, t h e n u p o n t h e  100  beach.  by  This preferred h a b i t a t i s represented  r a n g e s t r e t c h i n g f r o m t h e w a t e r ' s edge up i n t o , the  Gordon  Peromyscus d e n s i t i e s , i n f a v o r a b l e  l o c a t i o n s , f l u c t u a t e f r o m 1 t o 10 a n i m a l s p e r  further  the  yards a  of  linear  t o an a r e a  no  f o r e s t canopy t h a n about t e n f e e t .  Several  m i c e have b e e n drowned when c a u g h t i n t r a p s i n a d v e r t a n t l y s e t b e l o w t h e n i g h t h i g h w a t e r mark. contents  t a k e n f r o m the, d r o w n e d a n i m a l s r e v e a l e d  P e r o m y s c u s had t e e n Bendell upon the  feeding  (1959) has  reduction  documented t h e  i n b i r t h r a t e c a n be  i s l a n d s ' the  of the  intertidal  of amphipods s h o u l d be t h e s e two detritus  years.  on t h e b e a c h e s and  this  t h e r e f o r e f a v o r -the study  populations increase  t h e mice- p o p u l a t i o n s  to t h a t d e s c r i b e d  by  Bendell  that  therefore  this  In  general  e f f e c t upon  zone and  found  is unlimited.  expected to remain f a i r l y  eff/ect of i n c r e a s i n g the and  has  weathe  the the  density  stable  I n d e e d , a d d i t i o n a l s t o r m s mean " t h a t  i s washed up  these  a t t r i b u t e d to  a i r t e m p e r a t u r e and  c o n d i t i o n s w o u l d seem t o have l i t t l e microclimate  that  e f f e c t s of weather  weather c o n d i t i o n s even though food  these northern  stomach  e x c l u s i v e l y upon amphipods.  f e c u n d i t y of. P e r o m y s c u s l e u c o p u s and  a 50 perce'nt bad  A n a l y s i s of the  should  over  extra have  o f a v a i l a b l e amphipods  i n Peromyscus d e n s i t y . are  f o r P.  the  showing a s i m i l a r leucopus,  except  In  cycle that  the  e f f e c t o f t h e w e a t h e r seems t o be  D p y l e I s l a n d , one considered  of the  t o be  few  favorable  environments.  t o P e r o m y s c u s , showed a  account f o r the The were n o t It  1969  of the  constant  favorable  supply, which I b e l i e v e  i n l a n d h a b i t a t , would  a f f e c t e d by t h e  density.  h  a v e  a  of  of the  beneficial e f f e c t i n maintaining  31 p e r c e n t  hard to imagine a higher  m i c e i n p l a g u e numbers.  was  t.he h i g h e s t  recorded  The  r e g u l a r , t h e n the  c o n t r o l , c a n be  known p r e d a t o r s  r u l e d out  upon T r i a n g l e I s l a n d .  of the  Predation* as t h e r e  f a c t o r s which regulate  l o a d on t h e s e p o p u l a t i o n s  no  P a r a s i t e s were  t h e s e mice were  of mice are  not  maintain  as a means  are  f r e e f r o m any parasite  of  islands  should  indeed  The  success  c o n s i s t e n t , weather  a problem t o the p o p u l a t i o n ; parasite.  of  impression  trapping  Scott Islands  s t a b l e d e n s i t y of P e r o m y s c u s .  of p o p u l a t i o n  one  d e n s i t y of Peromyscus  f o r any  I f these adverse yet annually  c o n d i t i o n s are  a  standards.  on T r i a n g l e I s l a n d , f o r t h e b e a c h zone gave t h e of h a r b o r i n g  1968.  Scott  p e r e n n i a l w i n t e r weather p r o f i l e , a l b e i t  . I t w o u l d be  a fairly  Peromyscus  severe w i n t e r  exposed nature  s e v e r e c o n d i t i o n s by human  studied.  probably  t r a p p i n g r e s u l t s i n d i c a t e d t h a t the  seems p r o b a b l e t h a t t h e  population  (8 p e r c e n t )  o b s e r v e d change i n p o p u l a t i o n  adversely  I s l a n d s may  extent  i n h a b i t i n g t h e more  A f l u c t u a t i n g food  is a characteristic  greater.  areas l a c k i n g a beach h a b i t a t  density that f l u c t u a t e d to a greater t h a p t h a t of p o p u l a t i o n s  somewhat  not  remarkably the solely  d e n s i t y d e p e n d e n t , f o r the V a n s i t t a r t I s l a n d of P e r o m y s c u s b e i n g population vectors  had  and  an  one  f o u r t h the  d e n s i t y of the  w o u l d o f f e r an  future  extreme t o l e r a n c e  The  The  c a u s e s of p a r a s i t e i n f e s t a t i o n s  r e m a i n unknown and  Triangle  Triangle  overabundance of e c t o ^ - p a r a s i t e s .  specific  study.  population  mice t e n d to c r e a t e  the  i n t e r e s t i n g problem f o r a and  a m i c a b i l i t y of  impression  t h a t an  the  intrinsic  s e l f - r e g u l a t o r y m e c h a n i s m , s u c h as t h a t p r o p o s e d f o r Microtus  ( C h i t t y , 1952;  Peromyscus The  K r e b s , 1970)  does n o t  regulate  r e s u l t s of the  stomach  populations.  b e a c h h a b i t a t and  the  a n a l y s i s e s t a b l i s h P e r o m y s c u s as an scavenger. Pacific  R i c k e t t s and  Tides"  ( p . 158)  of Orchestrojdea  m e n t i o n t h a t the  lessens  u t i l i z e d by  carnivorous material,  diet,  nocturnal  their vulnerability r i c h food  equally voracious  c o n t a i n i n g a high  The  readily  predator.  proportion  This  of  chitonous  region  be  one  relative,  that i s  pouch a f f o r d i n g p r o t e c t i o n to  ( H o r n e r e_t a l , 1 9 6 5 ) .  E x a m i n a t i o n of  stomach m o r p h o l o g y o f i n s u l a r P e r o m y s c u s i n d i c a t e s with  predation.  r o u g h c h i t i n o u s d i e t o f Onychomys a p p e a r s  encapsulated i n a fundic area  habits  to b i r d  s o u r c e was  c o r r e l a t e d w i t h a stomach g l a n d u l a r  sensitive  intertidal  i s somewhat a n a l a g o u s t o t h a t o f a c l o s e  Qnychomys. t o be  an  important  C a l v i n i n t h e i r b o o k "Between  They were unaware t h a t t h i s being  these  exception,  correlated with  no  special g a s t r i c adaptation  a h i g h l y c h i t i n o u s d i e t has  i n tl}is group of m i c e  T  The  this the  that, that  taken  can  place  stomach i s d i s t i n c t l y b i l o c u l a r  with  the glandular  region  f u l l y exposed t o t h e stomach  s i m i l a r t o the d e s c r i p t i o n g i v e n exception Island.  by Vorontsov  i n d i v i d u a l had a d i s t i n c t p a r t i a l  of the glandular  The Balaclava  invagination  region.  Peromyscus are o p p o r t u n i s t i c .  The d e n s i t y  seems t o be d i r e c t l y r e l a t e d t o t h e f l o r a l , habitat  (1957).  was n o t e d i n t h e s i n g l e s p e c i m e n t a k e n f r o m This  contents,  diversity.  On t h e i s l a n d s  ment i s t h e p r e f e r r e d t h r o u g h o u t the y e a r .  habitat  of Peromyscus  f a u n a l , and  s a m p l e d the b e a c h  an,d t h i s  environ-  seems t o be t h e c a s e  MORPHOLOGICAL AND  BEHAVIORAL VARIATION I N INSULAR POPULATIONS OP PEROMYSCUS.  I n t h e p r e s e n t s t u d y 13 i s l a n d s and  two  continental  p o p u l a t i o n s of Peromyscus m a n i c u l a t u s are a n a l y z e d  and  c o r r e l a t i o n s o f v a r i o u s m o r p h o l o g i c a l measurements a r e made to a s c e r t a i n the taxonomic  significance  i n s u l a r groups of Peromyscus. are p r e s e n t e d i n t h r e e A)  The  rubidus  The  of v a r i a t i o n  r e s u l t s of t h i s  within  analysis  sections:  Peromyscus m a n i c u l a t u s gambeli  and P.  maniculatus  sample f r o m t h e w e s t c o a s t o f t h e c o n t i n e n t a l  United  States. B) The  P. m a n i c u l a t u s g e o r g i e n s i s c o m p l e x f r o m Bowen,  Thormanby, T e x a d a , and C) The  Savary I s l a n d s , B r i t i s h  G o r d o n , G o l e t a s , and  a c o m p l e x P. m a n i c u l a t u s Hall  group.  upon t h e G e o r g i a S t r a i t I s l a n d s o f  a s s i g n e d a new  m a n i c u l a t u s of S a v a r y , The  S c o t t I s l a n d groups r e p r e s e n t i n g  ( 1 9 3 8 ) , i n a s t u d y o f t h e c o l l e c t i o n made by R.  Cummings i n 1936 Columbia,  Columbia.  subspecies t i t l e  A.  British  t o the Peromyscus  T e x a d a , Thormanby, and Bowen I s l a n d s .  a s s i g n e d t a x o n , P. nr. g e o r g i e n s i s , was  an a t t e m p t  convey the i m p r e s s i o n t h a t , d e s p i t e the degree of e x i s t i n g between these p o p u l a t i o n s , t h e y possessed s i m i l a r m o r p h o l o g i c a l c h a r a c t e r i s t i c s w h i c h were d i f f e r e n t from the a d j a c e n t m a i n l a n d v a r i a t i o n f o u n d among t h e s e  Peromyscus.  i s l a n d m i c e was  to  isolation very significantly The  shown t o be  slight a  clinal  i n v e r s e r e l a t i o n s h i p between the degree of  f r o m t h e m a i n l a n d and t h e t a i l  length.  isolation  McCabe and  Cowan (1945)  f o u n d among t h e i s l a n d s i m m e d i a t e l y t o t h e n o r t h o f V a n c o u v e r I s l a n d a complex of m o r p h o l o g i c a l l y d i v e r s e Peromyscus inhabiting adjacent islands t r e a c h e r o u s ocean water.  s e p a r a t e d by o n l y a few y a r d s  These a l l o p a t r i c , b u t g e o g r a p h i c a l l y  n e i g h b o u r i n g , P e r o m y s c u s r e p o r t e d l y show no  simple  m o r p h o - g e o g r a p h i c r e l a t i o n s h i p s u c h as r e p o r t e d f o r P. georgiensis.  F u r t h e r i n v e s t i g a t i o n comparing  the  and t h e r e l a t i o n s h i p b e t w e e n e n v i r o n m e n t a l and  e v o l u t i o n a r y r e l a t i o n s h i p s of these  The  insular  geographic  rodents.  c o n t i n e n t a l sample c o n s i s t i n g o f 183  skulls  g a m b e l i was  f r o m t h e Museum o f V e r t e b r a t e Z o o l o g y , B e r k e l e y , o t h e r s p e c i m e n s were e i t h e r l i v e  summers of 1968,  1969,  and  of t h e B r i t i s h C o l u m b i a  1970  o r came f r o m t h e  collection  P r o v i n c i a l Museum, V i c t o r i a . and a d u l t s t a t u s  Those  o f b r e e d i n g and  t h e r e f o r e u s e d as t h e p r i m a r y c r i t e r i o n f o r the  c l a s s i f i c a t i o n of a d u l t s t a t u s . that these a d u l t participating  was  e r u p t e d t h i r d m o l a r s and b r o w n p e l a g e ,  a r e a t the s t a g e where t h e y a r e c a p a b l e was  obtained  California.  i n t h e manner o u t l i n e d by F o s t e r ( 1 9 6 5 ) .  animals, having f u l l y  of  trapped during the  Only a d u l t specimens were u t i l i z e d determined  the  METHODS  P e r o m y s c u s m a n i c u l a t u s r u b i d u s and P. m.  All  m.  variation  factors could very w e l l permit a b e t t e r insight into  MATERIALS AND  of  F i e l d evidence  specimens are i n t h e i r  i n t h e i r f i r s t breeding  suggests  second y e a r  season.  and  this  Fig.  4  Collection locations  f o r the c o n t i n e n t a l  of P e r o m y s c u s m a n i c u l a t u s  gambeli  samples  and P. m.  rubidus  Characters  measured:  I n order  to a l l e v i a t e any  misunderstanding  r e s u l t i n g f r o m a d i f f e r e n c e i n measurement t e c h n i q u e o u t l i n e o f t h e methods u s e d i n t h i s s t u d y this  a brief  i spresented a t  point.  A) TOTAL L E N G T H . . . d e t e r m i n e d b y p l a c i n g t h e d o r s a l s u r f a c e o f the to  animal  f l a t upon a m e t r i c r u l e a n d a p p l y i n g enough  s t r e t c h the specimen s l i g h t l y  parallel  to the rule.  so t h a t t h e b a c k b o n e i s  M e a s u r e m e n t s were t a k e n f o r t h e d i s t a n c e  from t h e t i p o f t h e nose t o t h e end o f t h e t a i l the t e r m i n a l t a i l  disregarding  hairs.  B) T A I L LENGTH...the ruler  tension  b o d y was hung o v e r t h e end o f a m e t r i c  i n s u c h a way as t o f o r m a 90° a n g l e w i t h t h e t a i l .  Tail  l e n g t h t h e r e f o r e represents t h e d i s t a n c e from the f i r s t  free  c a u d a l v e r t e b r a t o the t i p o f t h e t a i l ,  the t e r m i n a l  hairs.  C) EAR LENGTH.. .measurements were t a k e n the n o t c h  and n o t i n c l u d i n g  from t h e bottom of  to the t i p of the ear.  D) HIND FOOT L E N G T H . . . t h i s i s t h e d i s t a n c e f r o m t h e h e e l t o the  end o f t h e l o n g e s t t o e a n d n o t t o t h e t i p o f t h e c l a w .  The r o c k y h a b i t a t o f most P e r o m y s c u s i n v e s t i g a t e d i n t h i s s t u d y was c o n s i d e r e d determination the d i g i t  t o be a s i g n i f i c a n t  of claw l e n g t h and f o r comparative a n a l y s i s  length without  more i l l u s t r a t i v e  t h e c l a w was c o n s i d e r e d  t o be t h e  measure.  E) BODY LENGTH...the l e n g t h from the t o t a l measurement.  f a c t o r i n the  value  obtained from s u b t r a c t i n g the t a i l  l e n g t h was u s e d as t h e b o d y  length  A l l m e a s u r e m e n t s were t a k e n t o t h e n e a r e s t h a l f m i l l i m e t e r and a l l d a t a were s t o r e d u p o n I.B.M. c a r d s f o r c o m p u t e r analysis. S k u l l dimensions  were t a k e n u s i n g H e l i o s n e e d l e  c a l i p e r s t o the n e a r e s t 0.1  RESULTS AND S e c t i o n A:  millimeter.  DISCUSSION The  C o n t i n e n t a l Sample  Osgood ( 1 9 0 9 ) , maniculatus  point metri  i n h i s a n a l y s i s of t h e  g a m b e l i , made t h e  following  Peromyscus  statement:  "Specimens from M o n t e r e y , the type l o c a l i t y , are a b s o l u t e l y i d e n t i c a l w i t h those from San D i e g o and t h e n o r t h e a s t c o a s t o f l o w e r C a l i f o r n i a , and t h e i n t e r v e n i n g r e g i o n i s i n h a b i t e d b y e x a c t l y t h e same f o r m . " This c r e a t e s the this Clark  i m p r e s s i o n t h a t no v a r i a t i o n e x i s t s w i t h i n  subspecies.  I n v e s t i g a t i o n s by  ( 1 9 4 1 ) , and  Dice  Sumner ( 1 9 1 8 , 1 9 2 6 ) ,  ( l 9 4 0 a , b ) , have shown t h a t e x t e n s i v e  v a r i a t i o n e x i s t s w i t h i n each s u b s p e c i e s , y e t s u b s p e c i f i c boundaries  are abrupt  are c o n s i d e r e d . tends  i f certain specific  characteristics  Sumner (1918) m e n t i o n s t h a t  towards continuous  g r a d a t i o n and  pigmentation  morphological  m e a s u r e m e n t s show a d i s c o n t i n u o u s d i s t r i b u t i o n . taxonomist  has t o j u d g e w h e t h e r t h e v a r i a t i o n i s o f  magnitude i n d i c a t i v e this  of s p e c i f i c  a l l o p a t r i c , and  a  or s u b s p e c i f i c l e v e l ,  j u d g e m e n t becomes e v e n more d i f f i c u l t  involved.  The  t o make when  especially insular, populations  are  and  I have s e l e c t e d P. m.  gambeli  and P. m.  r u b i d u s as  the  c o n t i n e n t a l s u b s p e c i e s u s e d as t h e c o n t r o l g r o u p s i n c e t h e y a r e among t h e most e x t e n s i v e l y s t u d i e d . a r e r e p r e s e n t e d b y a l a r g e and collection.  The  range  gambeli  r e p r e s e n t e d by s a m p l e s from f i v e geographic  subspecies  r e a d i l y a v a i l a b l e museum  of these subspecies i s a l s o a  e x t e n s i o n of the geographic P e r o m y s c u s m.  These two  r e g i o n sampled i n t h i s and P. m.  (n=85 and  r u b i d u s are  encompassing n e a r l y the t o t a l  study. each  98, r e s p e c t i v e l y )  areas of d i f f e r e n t l a t i t u d e s range  of each  southern  taken and  subspecies,  (fig.4). A n a l y s i s o f t h e 10 c r a n i a l m e a s u r e m e n t s ( t a b l e s 1A, appendix  l ) , show t h a t i n P. m.  gambeli  the males are  s i g n i f i c a n t l y d i f f e r e n t from the females.  Sexual  a p p a r e n t l y i s not pronounced i n e i t h e r subspecies according t o Dice maniculatus  (1949) any  i s s l i g h t and  when c o m p a r i n g  sexual difference  does n o t j u s t i f y  not  dimorphism and  i n P.  s e p a r a t i n g sexes  populations.  T h i s c o n t r a s t s w i t h t h e r e s u l t s r e p o r t e d by Sumner who  found  noticable 0.4-3.6 mm.  2A ,  t h a t s e x u a l dimorphism  i n P. m a n i c u l a t u s was  (1926), most  i n t o t a l body l e n g t h , w i t h t h e males b e i n g s m a l l e r than the females.  The  males are  also  r e p o r t e d t o have a h i n d f o o t l e n g t h s i g n i f i c a n t l y g r e a t e r t h a n t h a t of the female have g e o g r a p h i c and  (Sumner, 1 9 2 6 ) .  Other  s p e c i e s a r e known t o  g r a d i e n t s i n s e x u a l dimorphism  t h e r e f o r e any  (Cowan,  such v a r i a t i o n w i t h i n t h e i s l a n d  1956)  sample  c o u l d be  of, s i g n i f i c a n c e  i n e v a l u a t i n g the e f f e c t s of  an  i s l a n d e n v i r o n m e n t upon Peromyscus morphology. Comparison of the  two  c o n t i n e n t a l subspecies  using  the  t e n c r a n i a l measurements show t h a t s i g n i f i c a n t d i f f e r e n c e s e x i s t between subspecies  at the 1 percent  the t e n c h a r a c t e r s measured.  Sexual  level  d i m o r p h i s m i n P.  r u b i d u s , when e v a l u a t e d by c o m p a r i s o n o f t h e characters, exists level  ( t a b l e s 3A,  o n l y a t the 4A,  The  characters  i n t h i s study are a p p a r e n t l y not unduly  subspecific  m.  same t e n  5 p e r c e n t or g r e a t e r  appendix l ) .  d i f f e r e n c e s i n P e r o m y s c u s and  i n seven of  confidence utilized  i n f l u e n c e d by  can be r e l i e d u p o n t o  sexual  indicate  variation.  S i g n i f i c a n t v a r i a t i o n o c c u r r e d b e t w e e n sample means (both sexes  combined) f o r each l o c a t i o n a n a l y s e d ,  areas  2 (table  1 and  probably  1A,  appendix l ) .  a r e f l e c t i o n of the  O r e g o n , and  and  i n the  The  Formation  c o u l d be  Strait,  C o a s t a l W e s t e r n H e m l o c k Zone,  V a r i a t i o n b e t w e e n sample l o c a t i o n s  i n each r e s p e c t i v e c l i m a t i c  classification)  Georgia  i n the Mesothermal B i o g e o c l i m a t i c  ( K r a j i n a , 1969).  v a r i a t i o n b e t w e e n t h e two  southern,  California,  C o a s t a l D o u g l a s F i r Zone  t h e n o r t h e r n s t u d y g r o u p i n the  b o t h a r e c o n s i d e r e d t o be  in  This v a r i a t i o n i s  u n i f o r m i t y of n o r t h e r n  southern Washington.  study i s l a n d s are s i t u a t e d  except  zone i s m i n i m a l b u t  z o n e s (Csb a factor  and  climatic  C f b , Koppen  i n the d i s t r i b u t i o n  of  Peromyscus s p e c i e s . From t h i s a n a l y s i s o f t h e c o n t i n e n t a l p o p u l a t i o n s s e v e r a l i n f e r e n c e s c a n be made t h a t a p p l y t o t h e samples.  F i r s t , and  a n a l y s i s of  island  p e r h a p s t h e most i m p o r t a n t p o i n t , i s  t h a t t h e d e g r e e o f v a r i a t i o n , as d e t e r m i n e d b y t h e r e l a t i v e l y low a n d c o n s i s t e n t values,  standard  i s not being  d e v i a t i o n and standard  error  unduly i n f l u e n c e d by the small  s i z e o f some i s l a n d c o l l e c t i o n s .  Secondly,  sample  although  s i g n i f i c a n t d i f f e r e n c e s o c c u r w i t h i n each, g r o u p , s e v e n o f t h e ten c r a n i a l characters  s e l e c t e d have t h e p o t e n t i a l t o be  s i g n i f i c a n t l y d i f f e r e n t at a l e v e l i n d i c a t i v e of continental subspecies. Sample s i z e s f r o m e a c h o f t h e 13 i s l a n d s w e r e greater  generally  t h a n t h o s e f o r e a c h l o c a t i o n s a m p l e d i n t h e two  continental subspecies. d e v i a t i o n and s t a n d a r d  The s m a l l error values  and c o n s t a n t  standard  (the l a t t e r  consistently  l e s s t h a n 0.01 p e r c e n t ) f o r b o t h sample s e r i e s a l l o w s f o r direct  c o m p a r i s o n o f t h e mean a v e r a g e v a l u e s .  These d a t a  i n d i c a t e t h a t i n t r a - i s l a n d v a r i a t i o n i n c r a n i a l morphology i s comparable t o t h a t Several  studies  populations  f o u n d i n some c o n t i n e n t a l  (Lerner,  1954) have shown t h a t  have a g r e a t e r  are  that  C o n s e q u e n t l y , the v a r i a t i o n  f o u n d among t h e i s l a n d p o p u l a t i o n s  First,  inbred  degree o f v a r i a t i o n than  found i n out-bred populations.  regarding  populations.  r a i s e s two  questions  the s i g n i f i c a n c e of the observed d i f f e r e n c e s .  the p o s s i b i l i t y a r i s e s that the characters  measured  n o t t h e most s e n s i t i v e m a r k e r s o f t h e e v o l u t i o n a r y  changes t a k i n g p l a c e  i n this  t h a t even the s m a l l e s t population  group of r o d e n t s ,  island i s a "universe"  and i n b r e e d i n g  f a c t o r i n the evolution.  and  t o a mouse  e f f e c t s are therefore The f i r s t p o i n t  secondly,  not a  c o u l d n o t be  answered unless this  a l l characters are recorded  i s a near impossible  be a n s w e r e d b y  task.  r e f e r r i n g t o the  I f e a c h sample l o c a t i o n o f the considered  an  The  continental  e r r o r v a l u e s w o u l d r e f l e c t any  and  p e r h a p s some i d e a o f t h e minimum and  i n t h e i r morphological t h e y do  present profile  f o r each i n d i v i d u a l  s a m p l i n g of the  out-bred island  u n i t and  i s indeed  standard  the  can  of  characters,  ascertained.  that  The  same d e g r e e o f v a r i a t i o n  f o r the, t o t a l p o p u l a t i o n From t h i s  s m a l l ranges trapped  continental population  t h e r e f o r e the area a "universe".  This  f r o m s t u d i e s u n d e r t a k e n by B e r r y  be  deviation  maximum r a n g e  sample l o c a t i o n .  have assumed t h a t e a c h o f t h e the  populations.  discordance  i n b r e d u n i t c o u l d be  and  possibly  c o n t i n e n t a l subspecies  and  continental populations  compared,  s e c o n d p o i n t can  i s l a n d s i t u a t i o n t h e n the  c o n s t i t u t e s an  and  I  during  is still  o f e v e n the  as  an  smallest  conclusion gains  weight  (1964) w h i c h show t h a t  i n b r e d s t r a i n s of i s l a n d mice, o r i g i n a l l y taken from  the  same p o p u l a t i o n , have a h i g h e r d e g r e e of v a r i a n c e t h a n d i d the  original population.  populations  are  not  His suggestion  allopatric  l o c a t i o n and  populations  T e x a d a I s l a n d s were  exist.  No  on T e x a d a I s l a n d t h e  mice i s l a r g e r t h a n t h a t of the of the  island.  trapped  i n a r e a s where I have assumed significant  d i f f e r e n c e s o c c u r w i t h i n t h e s e s a m p l e s w i t h the exception:  island  inbred units.  N i g e i , Hope, T r i a n g l e and i n more t h a n one  i s that  cranial width  cranial one of the  mice i n h a b i t i n g the  beach  interior  The b o d y m e a s u r e m e n t s show a l a r g e r intra-population  samples  minimum sample  of  v a r i a t i o n b u t t h i s does n o t seem t o be  adversely influenced between  degree  b y t h e sample s i z e  i s comparable.  since the variance  That i s t o say, the  s i z e has b e e n e x c e e d e d  critical  i n most o f t h e c a s e s  analysed. On the b a s i s o f b o d y l e n g t h t h e i s l a n d s a m p l e s grouped i n t o  two c l a s s e s :  Bowen, Thormanby, Texada  a) t h e s o u t h e r n g r o u p , and S a v a r y I s l a n d s ,  offer  from Texada,  phenotype, the  S a v a r y , Thormanby and Bowen I s l a n d s  a unique o p p o r t u n i t y f o r the s t u d y of v a r i a t i o n i n  allopatric  populations.  The mean a v e r a g e o c c i p i t o n a s a l -0.4  mm.  length  fluctuates  and c o m p a r a b l y l o w l e v e l s o f v a r i a t i o n e x i s t i n  each of t h e o t h e r c r a n i a l measurements, ( t a b l e Such v a r i a t i o n s continental R.A.  are w e l l w i t h i n  subspecies.  Hall  between  m a i n l a n d and t h e t a i l  (1938) i n h i s s t u d y o f t h e  (table  3) a r e d i f f e r e n t  the b o d y / t a i l  a clinal  the degree of i s l a n d length.  ratio  1936  inverse  i s o l a t i o n f r o m the  T a i l and b o d y measurements  f o r each of t h e i s l a n d (figs.  5A, a p p e n d i x l ) .  the e s t i m a t e d t o l e r a n c e of a  Cummings c o l l e c t i o n e s t a b l i s h e d  relationship  yet  islands.  Southern Islands  D i s t i n g u i s h e d by a r e p o r t e d l y s i n g l e Peromyscus  comprising  and b) t h e  n o r t h e r n g r o u p , made up o f t h e r e m a i n i n g n i n e S e c t i o n Bt The  are  populations  29A-32A, a p p e n d i x 2) has  m o d a l v a l u e o f 1.0-1.1 f o r a l l i s l a n d s  i n t h i s group  m o r p h o l o g i c a l c r i t e r i a upon w h i c h the o r i g i n a l  a  The  taxonomic  e v a l u a t i o n v a s made i s n o t r e p r e s e n t a t i v e o f t h e p o p u l a t i o n o f P e r o m y s c u s upon t h e s e  i s l a n d s today.  study i s g e n e r a l l y l a r g e r i n t h e 1938 s t u d y  ISLAND  The sample u s e d i n t h i s  i n b o d y measurements  ( t a b l e 3, f i g s .  than that used  5,6,7).  Thormanby  Savary  3/4  1  2 1/2  2 3/4 m i l e s  BODY LENGTH (HALL,1938)  83.0(4)  82.6(14)  78.0(24)  69.2(25)  BODY LENGTH (1960-1970)  87.0(22)  81.0(28)  81.9(10)  85.1(37)  AVERAGE  86.4(26)  81.7(32)  79.1(34)  78.7(62)  DISTANCE OFF SHORE  (Sample s i z e Table  3  Bowen  Texada  i n brackets') Body l e n g t h s f o r P e r o m y s c u s o f t h e G e o r g i a  Strait  Region.  E i g h t s p e c i m e n s t a k e n f r o m T e x a d a I s l a n d i n 1 9 4 5 , when compared w i t h t h e 1936 a n d 1968 c o l l e c t i o n s , a r e i n t e r m e d i a t e i n o c c i p i t o n a s a l l e n g t h b u t l a r g e r i n many o f t h e o t h e r measurements.  The S a v a r y  I s l a n d sample has a s l i g h t l y  evolutionary pattern i n morphological data i n d i c a t e  skull  change a n d t o g e t h e r  that the p o t e n t i a l f o r s i g n i f i c a n t  f l u c t u a t i o n e x i s t s over a r e l a t i v e l y I have o b t a i n e d s k u l l measurements  different these  morphological  short p e r i o d of time.  of t h e 1936 c o l l e c t i o n t h a t  I o b t a i n e d o n l o a n f r o m t h e Museum o f V e r t e b r a t e  Zoology,  The U n i v e r s i t y o f C a l i f o r n i a a t B e r k e l e y . No c l i n e measurements.  c a n be e s t a b l i s h e d f o r a n y o f t h e s k u l l  I n d e e d , i f c h a n g e s a s l a r g e as t h o s e  recorded  island distance (mi) total P r e 1950 1968-1970 88  THORMANBY 2  SAVARY  BOWEN  4  1  <£ 2  86.4 83.0 87.0  81.7 82.2 81.0  79.1 77.7 81.9  TEXADA 21 78.7 69.3 85.1  _  84 80 '76 PRE 1950 72 68  IIi distance  Fig.  5  2 from mainland  2\  2\  i n miles.  Changes i n body l e n g t h o f P. m. g e o r g i e n s i s ,  island distance total P r e 1950 1968-1970  (mi )  THORMANBY 3.  4  SAVARY 1  92.3 87. 3 93.2  93.4 89.6 96.3  BOWEN 2\  94.0 92.9 96.0  TEXADA 2}  97.8 97.5 98.0  100  96 92 88  11  distance Fig.  6  Changes i n t a i l  .  2  2*  from t h e mainland  21 i n miles  l e n g t h o f P. m. g e o r g i e n s i s ,  Fig.  7  Changes i n o c c i p i t o n a s a l l e n g t h of P e r o m y s c u s m a n i c u l a t u s  georgiensis.  (i.e.  16mm.  expected  i n b o d y l e n g t h , 7mm.  are  of the group.  i s every  Differences i n morphological  incorporated into  arise.  biological  measurements  decade a r e v e r s a l phenomenon c a n be  f o r m u l a f o r naming  insular  determined.  i s l a n d s t h a t may  c h a n g e s t h a t have t a k e n facilities  clinal  1936 t h e r e have b e e n m a j o r e c o l o g i c a l c h a n g e s on  southern  southern  c a n be  s e p a r a t i o n and y e t  J u s t how t h i s  a working  p o p u l a t i o n s r e m a i n s t o be Since  of s p e c i e s  p o s s i b i l i t y t h a t i n the next  o f t h e s i t u a t i o n may  the  length)  c o u l d be e s t a b l i s h e d t o r e f l e c t t h e t r u e  of t h e m a g n i t u d e i n d i c a t i v e  there  skull  w i t h i n any 32 y e a r p e r i o d i t i s d o u b t f u l i f a n y  relationship status  i n total  c o r r e l a t e w i t h the  place i n s i t u .  have b e e n b u i l t  Houses a n d  supporting  i n c o n s i d e r a b l e numbers upon  islands i n recent years.  the n o r t h e r n u n i n h a b i t e d  morphological  I f t r a p p i n g success f o r  islands i s indicative  h a b i t a t f o r Peromyscus then  of the o p t i m a l  t h e l a n d a r e a most h e a v i l y  u t i l i z e d by P e r o m y s c u s has a l s o b e e n the f a v o r e d t y p e commercial development.  these  I n the s o u t h e r n  i s l a n d s the  sandy bays s u i t a b l e f o r dense p o p u l a t i o n s  of shallow  of Peromyscus  have l o n g s i n c e b e e n c l e a r e d o f s u r r o u n d i n g  v e g e t a t i o n and  housing  The  has been s u b s t i t u t e d i n i t s p l a c e .  P e r o m y s c u s have s u b s e q u e n t l y h a b i t a t and now Under t h e s e  occur  once-resident  been d i s p l a c e d from t h i s  only i n the surrounding  rocky  c o n d i t i o n s t h e change i n e n v i r o n m e n t a l  hillsides. pressures  w o u l d p e r h a p s have f a v o r e d a change i n gene and p h e n o t y p e frequencies. Foster  (1965) s p e c u l a t e d t h a t a r o c k y  s u b s t r a t e , because  it  n e c e s s i t a t e d more c l i m b i n g , f a v o r e d  balance. and  Horner  longer  A rocky  (1954) h y p o t h e s i z e d  tails  are  substrate  an a d v a n t a g e i n an  like  the  arboreal  t a k e n d u r i n g my  arboreal  a r e a t h a t the  considered  Horner.  ecologically disturbed  i s l a n d s , Thormanby, Bowen,  have l o n g e r  t h o s e P e r o m y s c u s f r o m the  Peromyscus  The  a p p l i e d as a t e s t o f The  samples the  Peromyscus from  t a i l s and  feet  analogous  t h e o r i e s o f F o s t e r and  Savary, should  for  habitat.  displaced  h a b i t a t r e f e r r e d t o by H o r n e r . s t u d y can be  tail  that larger hind  have b e e n f o r c e d t o i n h a b i t c o u l d be t o the  a longer  l a r g e r hind  the  and feet than  r e l a t i v e l y untouched Texada I s l a n d  populations. The  data  ( f i g . 6;  a d i s t i n c t increase i n h a b i t i n g the  t a b l e s 6A,  in tail  disturbed  length  islands.  7A,  appendix l ) i n d i c a t e  f o r t h o s e Peromyscus The  f o o t measurement,  h o w e v e r , seems to have r e m a i n e d r e l a t i v e l y u n c h a n g e d . data tends to substantiate tail,  an  favored,  under c e r t a i n  increment i n  were s a m p l e d i n two  the  and  circumstances.  a f u r t h e r means of t e s t i n g t h i s  populations that  idea that  o r g a n u s e d f o r b a l a n c e , i s an a d v a n t a g e ,  therefore As  the  These  hypothesis,  locations.  the  Texada  I have assumed  the f i r s t l o c a t i o n , a b e a c h h a b i t a t w i t h a  rocky  s u b s t r a t e , w o u l d show a s e l e c t i v e e f f e c t on appendage  length  if  context.  the  Foster  and  Horner h y p o t h e s e s are v a l i d  C o m p a r i s o n o f s k u l l measurements ( f i g . 7; show a r e v e r s a l o f the  general  trend  in this  t a b l e 8A,  appendix l )  of increment i n body  dimensions, f o r the s k u l l  s i z e s a r e s m a l l e r i n the beach-  d w e l l i n g mice than those o f the Peromyscus t r a p p e d i n t h e interior  of the i s l a n d .  appendix  2) a r e t h e same f o r b o t h o f t h e g r o u p s  mean a v e r a g e  The b o d y - t a i l r a t i o s  (figs.  30A-33A,  measured.  The  m e a s u r e m e n t s a r e l a r g e r f o r most o f t h e c h a r a c t e r s  m e a s u r e d and i f t h e r e i s s e l e c t i o n t h e n i t i s f o r a l a r g e r p h e n o t y p e and n o t f o r a s p e c i f i c a d a p t a t i o n hypotheses  appendage.  put f o r t h b y Horner  Hence, the  and F o s t e r  cannot  be f u l l y s u b s t a n t i a t e d b y my d a t a . Any  attempt a t r e d e f i n i n g the systematic r e l a t i o n s h i p s  Of t h e s e i n s u l a r r o d e n t s must u t i l i z e d a t a t a k e n f r o m t h e same l o c a t i o n o v e r a p e r i o d o f y e a r s , o t h e r w i s e s h o r t t e r m fluctuations  i n p o p u l a t i o n means o f t h o s e m o r p h o l o g i c a l  measurements used i n t h e taxonomic negate  evaluation w i l l  tend t o  a l l attempts a t d e r i v i n g a meaningful phylogenic  comparison.  S h o u l d s u c h a s t u d y be i m p o s s i b l e t h e n t h e  i n v e s t i g a t o r h a s no a l t e r n a t i v e t o c r e a t i n g a t a x o n o m i c  category  b r o a d enough t o a l l o w f o r t h e i n h e r e n t f l u c t u a t i o n i n population characters. 1936  1968  ISLAND  N  TAIL  FOOT  N  SAVARY  14  90  22.3  18  96  21.0  BOWEN  24  93  21.6  10  96  21.6  4  87  22.4  26  93  20.9  25  98  22.3  37  98  21.8  THORMANBY TEXADA *  .  TAIL  d i f f e r e n c e i n measurement t e c h n i q u e , d o e s n o t i n c l u d e length.  Table  4  FOOT*  claw  A v e r a g e t a i l and f o o t measurement f o r 1936 and 1968-1970 c o l l e c t i o n s .  Section  C:  The N o r t h e r n  Islands  I n t r a - i s l a n d v a r i a t i o n i n the northern comparable t o t h a t o b t a i n e d  islands i s  i n t h e s o u t h e r n sample.  Intra-  i s l a n d v a r i a t i o n s ( t a b l e s 9A, 10A, a p p e n d i x l ) o f t h e b o d y d i m e n s i o n s a r e i n many c a s e s l e s s t h a n t h e d e g r e e o f v a r i a t i o n s noted i n the southern p o p u l a t i o n s ,  y e t these d i f f e r e n c e s  have  b e e n s u f f i c i e n t f o r McCabe a n d Cowan (1945) and G u i g u e t (1955) t o d e s i g n a t e A discriminant  new s u b s p e c i e s on t h e n o r t h e r n analysis, using  fourteen  islands.  c h a r a c t e r s , has  been employed t o determine t h e i n t e r - r e l a t i o n s h i p s o f these insular populations. functions  ( a p p e n d i x 3) I c a t e g o r i z e d  as t o p r o b a b l e known p l a c e island  I n e m p l o y i n g t h e computed  i s l a n d of o r i g i n .  discriminant  each i n d i v i d u a l specimen  The d e v i a t i o n s f r o m t h e  o f o r i g i n a r e assumed t o r e f l e c t t h e a l t e r n a t e  population  which i s c l o s e s t taxonomically.  The d e g r e e  of i n t e r - i s l a n d r e l a t i o n s h i p i s shown g r a p h i c a l l y i n f i g u r e 8 with  t h e m a g n i t u d e o f t h e r e l a t i o n s h i p a p p r o x i m a t e d as a  p e r c e n t a g e of t h e t o t a l alternate  sample s i z e s h o w i n g a f f i n i t y t o t h e  island location.  I t i s obvious that the small  m i c e show a c l o s e a f f i n i t y f o r one a n o t h e r w h i l e the  l a r g e phenotype group a r e a l s o The  islands as  members o f  seemingly r e l a t e d .  d i r e c t i o n o f e v o l u t i o n i s n o t t h e same f o r a l l (table 5).  a general  I t i s not s p e c i f i c  increase  i n d i r e c t i o n , such  i n body s i z e f o r the l a r g e or s m a l l  mouse p h e n o t y p e s , o r f o r t h e i s l a n d g r o u p . I s l a n d sample e m p h a s i z e s t h i s  The  Triangle  l a c k o f d i r e c t i o n i n t h e mode  of e v o l u t i o n f o r t h e i r  s k u l l dimensions a r e apparently  i n c r e a s i n g i n s i z e a t t h e same t i m e t h a t t h e i r b o d y are  diminishing  proportions  ( t a b l e 5, t a b l e s 11A, 12A, a p p e n d i x l ) .  ISLAND  BODY LENGTH  DOYLE  1930 •  1970  HURST  SKULL LENGTH 6. Omm  1930 =  .QOmm  1970  6.0  .06  10.0  .04  NIGEI  1.0  .01  HOPE  6.0  .02  BALACLAVA  COX  1940 >- 1970  11.0  1940  1970  .03  LANZ  7.5  .02  TRIANGLE  4.5  .03  Table  5  Degree and d i r e c t i o n o f time.  m o r p h o l o g i c a l change  with  :  Sumner ( 1 9 1 8 ) d e s c r i b e d  i n Peromyscus p o l i o n o t u s  c o n d i t i o n where t h e f e m a l e s were  l a r g e r t h a n t h e males i n  t o t a l body l e n g t h b u t s m a l l e r i n f o o t s i z e . Peromyscus from D o y l e , H u r s t ,  The male  B a l a c l a v a , N i g e i , a n d Hope  I s l a n d s a r e l a r g e r t h a n t h e f e m a l e s i n t o t a l body but  all,  excepting  foot size.  Doyle  a  length  I s l a n d p o p u l a t i o n , have a l a r g e r  V a n s i t t a r t and T r i a n g l e p o p u l a t i o n s  have l a r g e r  f e m a l e s b o t h i n body s i z e a n d f o o t l e n g t h and i t i s o n l y t h e Cox a n d L a n z p o p u l a t i o n s  that follow the pattern  described  b y Sumne r . Body-tail ratios continuous north-south maximum o f 1.1.  (figs. cline  21A-33A,  a p p e n d i x 2) i n d i c a t e a  i n modal means f r o m a 0.8 t o a  S p e c i f i c modal v a l u e s  c a n be c o r r e l a t e d  w i t h the  g e o g r a p h i c a l d i s t r i b u t i o n of the i s l a n d s  Intermediate  b o d y - t a i l r a t i o ' groups are found  Balaclava Islands.  These two  d e t a i l e d a n a l y s i s i n the in this  sampled.  on Hope  and  p o p u l a t i o n s are d e s e r v i n g  final  of  e v a l u a t i o n of i n s u l a r e v o l u t i o n  area.  P e r o m y s c u s f r o m B a l a c l a v a , V a n s i t t a r t , Hope, Bowen, Thormanby, T e x a d a , and  Savary  a r e , i n a l l c h a r a c t e r s measured,  w e l l w i t h i n the range of v a r i a t i o n e s t a b l i s h e d f o r a c o n t i n e n t a l subspecies. There i s a s t r o n g r e l a t i o n s h i p between each o f the i s l a n d s w i t h the e x c e p t i o n of V a n s i t t a r t , which l i k e is morphologically distinct. t h i r t e e n i s l a n d s are these  i s l a n d s was  s t o c k s , one (fig.  such  The  affinities  as t o s u g g e s t  undertaken  by two  a s m a l l - b o d i e d and  seven  Boyle,  among t h e  that colonization  distinct  of  morphological  the o t h e r a l a r g e - b o d i e d form  9). I f t h e d a t a p r o v i d e d by Sheppe ( l 9 6 l ) and McCabe  and  Cowan (1945) h o l d t rue f o r the n o r t h e r n c o a s t o f  British  C o l u m b i a , t h e n i t i s p o s s i b l e t h a t the numerous  landslides  a r e l a u n c h i n g two  p h e n o t y p e s i n t o the o c e a n .  Ocean c u r r e n t s  a r e s u c h t h a t , a t c e r t a i n t i m e s o f t h e y e a r , any  raft  a d r i f t i n t h e n o r t h e r n r e g i o n w o u l d move s o u t h and good c h a n c e o f b e i n g w a s h e d up study  on one  set  stand  of the i s l a n d s i n  a my  area. Sheppe e s t a b l i s h e d t h e  500  f t . l e v e l as the e l e v a t i o n  where P e r o m y s c u s m a n i c u l a t u s have p e r s o n a l l y w i t n e s s e d along  t h e steep  i s r e p l a c e d b y P. o r e a s .  t h e r e s u l t s o f many l a n d s l i d e s  f j o r d s o f B r i t i s h C o l u m b i a t h a t have denuded  v e r t i c a l distances  i n e x c e s s o f 500 f t .  r e p o r t t h a t P. o r e a s a n d P. a u s t e r u s  McCabe a n d Cowan  occur a t sea l e v e l a t  Loughborough I n l e t , a f j o r d a p p r o x i m a t e l y of Vancouver.  Therefore  t h e two m o r p h o l o g i c a l l y  170 m i l e s  distinctive  that  f o r m s ( t a b l e 6) c o u l d be  of the observed dimorphic  island  Peromyscus s i t k e n s i s Merriam, another v e r y  f o r m o f i s l a n d mouse, i n h a b i t s t h e S i t k a , A l a s k a , The  north  i nboth s t u d i e s i t i s apparent  responsible f o r the founding populations.  I  large  region.  P. s i t k e n s i s m o r p h o t y p e , known b o t h a s P. s i t k e n s i s  provestensis  ( H a l l and K e l s o n ,  1959) a n d P. m a n i c u l a t u s  provestensis  (Cowan a n d G u i g u e t , 1 9 6 5 ) , i n h a b i t s t h e  geographically  intermediate  r e g i o n between A l a s k a and the  northern  study  islands.  possible  l i n k . i n the d i s p e r s a l o f the dimorphic a n c e s t r a l  This  l a r g e morphotype i s another  s t o c k t h a t I b e l i e v e t o be r e s p o n s i b l e f o r t h e s t u d y - i s l a n d colonization.  BODY  TAIL  FOOT  PEROMYSCUS SITKENSIS  219 mm  104 mm  26 mm  H a l l and K e l s o n  PEROMYSCUS OREAS  204 mm  109 mm  23 mm  Cowan and G u i g u e t  P . M . AUSTERUS  176  89  21  Cowan and G u i g u e t  LARGE MORPHOTYPE(A)  213  104  25  SMALL MORPHOTYPE(B)  183  95  21  Table  6  '  Mean b o d y m e a s u r e m e n t s f o r P e r o m y s c u s o f t h e c o a s t a l r e g i o n o f B r i t i s h Columbia.  128  126  124  BRITISH COLUMBIA Fig.  9  Dispersal  route  of  ancestral  Peromyscus  Two  cohorts would p r o v i d e a b i o l o g i c a l l y  meaningful  a r r a n g e m e n t t h a t i n d i c a t e s the e v o l u t i o n a r y p a t t e r n o f these  i s l a n d groups.  t o the p r o p e r c o h o r t s , and hypothesis such  t h e r e i s a need f o r a d d i t i o n a l  c a n be d e v e l o p e d  further.  I now  two  data before  have e v i d e n c e  to present  i n t e r a c t i o n s w i t h i n and b e t w e e n t h e s e  of that two  s i g n i f i c a n t f a c t o r s p e r t a i n i n g to the r e - e v a l u a t i o n  of the s y s t e m a t i c s of these  island  this  Cryptic characters,  as t h e b a c u l a and k a r y o t y p e , have p r o v e n t o be  the b r e e d i n g  as  taxonomic d e s i g n a t i o n a p p l i c a b l e to the  t a x o n o m i c h e l p , and  are  This d i v i s i o n creates a problem  mice.  forms  BREEDING TESTS AND OP  COMPARATIVE POSTNATAL DEVELOPMENT  INSULAR AND  INTER-ISLAND HYBRID PEROMYSCUS  Laboratory h y b r i d i z a t i o n between i s l a n d undertaken and  stocks  was  to a s c e r t a i n the c h a r a c t e r i s t i c s of s i z e  inheritan  the degree o f s t e r i l i t y between the v a r i o u s Peromyscus  m o r p h o t y p e s f o u n d u p o n t h e S c o t t , G o r d o n and G o l e t a s , Georgia Dice  Strait  island  (1940) f o u n d  groups. t h a t a complete range i n f e r t i l i t y  e x i s t s w i t h i n t h e genus P e r o m y s c u s . 1909)  Species groups  (Osgood,  e x h i b i t i n t e r g r o u p s t e r i l i t y whereas s u b s p e c i e s  most c a s e s  and  completely f e r t i l e .  T h i s s t u d y has  are i n  extended  the  i n v e s t i g a t i o n o f i n t e r p o p u l a t i o n f e r t i l i t y by a t t e m p t i n g c r o s s b r e e d P e r o m y s c u s t a k e n f r o m a l l o p a t r i c , and cases m o r p h o l o g i c a l l y d i s t i n c t i v e , Given the  c h o i c e , Peromyscus m a n i c u l a t u s  Howard, 1 9 4 4 ) .  Moore  (1965) f o u n d  have a s t r o n g  species  evidence  t h a t o l f a c t o r y c u e s were r e s p o n s i b l e f o r t h e  suggesting  isolation  inhabiting  c o a s t a l i s l a n d s of B r i t i s h C o l u m b i a show p r e or i n t e r g r o u p s t e r i l i t y  taxonomic r e v i s i o n w i l l  MATERIALS AND  be  two  insular  of  f r o m P.  f e r e n t i a l mating  I f . the  (Blair  P. m a n i c u l a t u s the  polionotus.  i n many  populations.  s e x u a l p r e f e r e n c e f o r members o f t h e i r own and  to  t h e n the case  forms  for  strengthened.  METHODS  P e r o m y s c u s were o b t a i n e d f r o m t h e  islands  (see  chapter  1  EH  Pi <  HH  8 •  S3 <H  , M P3 EH  2  X! o o  TRIANGLE  2  4  COX  3  2  S3 S3  M PH  <  O Ix!  i—i S3  1  1  2  2  LANZ HOPE  1  1  NIGEI  4  1  VANSITTART  1  1  EH CO PH  2  1  2"  m  PH >H  o  EH  O in EH  ej ,PH*L  4  3 1  HURST  1  1  DOYLE  1  1  TEXADA  4  THORMANBY  4  P.m.austerus ^<?TOTAL  Table  fH CD  -P  1 3  2.  2  S3  1  1  1  BALACLAVA  w  EH EH I—l W  >H PH S3  7  1 1  2 1  1 1  5  5  15 12 2  Male and f e m a l e p a i r i n g  arrangement.  EH  O o CM  for  l o c a t i o n d a t a ) d u r i n g t h e months o f A u g u s t ,  and  October  September  o f 1968 and May, J u n e , and J u l y o f 1 9 7 0 .  I n the  l a b o r a t o r y e a c h a n i m a l was m a i n t a i n e d i n i s o l a t i o n f o r t h i r t y days p r i o r  t o b e i n g p l a c e d w i t h a p r o s p e c t i v e mate. A l l  l i t t e r s born during t h i s control  i n t e r i m p e r i o d w e r e u s e d as t h e  group.  The a n i m a l s were m a i n t a i n e d i n r o u n d wash p a n s 5 i n c h e s deep a n d 12 i n c h e s i n d i a m e t e r . of  U n l i m i t e d f o o d i n t h e form,  P u r i n a L a b o r a t o r y Chow was p r o v i d e d a t a l l t i m e s a n d  o c c a s i o n a l supplements  o f f r e s h g r e e n s were o f f e r e d .  A  diet  s u p p l e m e n t o f mealworms was p r o v i d e d a t one p o i n t o f t h e experiment  i n an attempt t o s t i m u l a t e r e p r o d u c t i o n .  This  d i e t was m a i n t a i n e d f o r two months w i t h o u t any r e c o r d e d change i n the b r e e d i n g performance  o f the mice.  d i e t a r y s u p p l e m e n t was s u b s e q u e n t l y  This  expensive  dropped.  P a i r i n g was done a t random e x c e p t t h a t j u v e n i l e s were p l a c e d w i t h a mate o f t h e same age c l a s s .  The 78 p a i r s  ( t a b l e 7) w e r e m a i n t a i n e d a s b r e e d i n g p a i r s f o r a minimum of  16 m o n t h s .  D a i l y w e i g h t s w e r e t a k e n on e a c h  of the progeny  and t h e p r o g r e s s o f t h e y o u n g m o n i t o r e d f o r t h e f i r s t weeks o f t h e i r  RESULTS AND  life.  DISCUSSION  .  ' •  The l a b o r a t o r y g r o u p o f 78 p a i r s p r o d u c e d In  each case  three  the l i t t e r s  five  litters.  represented i n t e r - i s l a n d crosses,  h e r e a f t e r r e f e r e d t o as h y b r i d s .  None o f t h e i n t r a - i s l a n d  matings  conceived.  Thirteen field-conceived l i t t e r s  r e c o r d e d d u r i n g t h e t h i r t y day i s o l a t i o n p e r i o d a n d made up t h e c o n t r o l g r o u p f o r h y b r i d Litter for  these  comparison.  s i z e v a r i e d f r o m 1 t o 7 w i t h t h e means r e c o r d e d  each group  ( t a b l e 16A,  appendix l ) .  w e i g h t and g r o w t h r a t e was  Average  newborn  more c l o s e l y r e l a t e d t o t h e  s i z e t h a n to the female p a r e n t weight. was  were  also a f u n c t i o n of l i t t e r  Developmental  litter rate  s i z e and, c o n s e q u e n t l y ,  c o m p a r i s o n o f d a t a c a n be made o n l y b e t w e e n t h o s e l i t t e r s equal s i z e . the  This puts a n e a r l y impossible r e s t r i c t i o n  of  upon  a n a l y s i s o f t h e l i m i t e d amount o f d a t a g a i n e d i n t h i s  study.  However t h o s e i s l a n d s r e p r e s e n t e d by more t h a n  one  l i t t e r were p o o l e d and t h e a v e r a g e w e i g h t g a i n v a l u e f o r t h e average  litter  size plotted  (fig.  10).. I n t h i s manner some  c o m p a r i s o n s b e t w e e n some o f t h e i s l a n d P e r o m y s c u s p o p u l a t i o n s c a n be made. No c o r r e l a t i o n e x i s t s b e t w e e n the p a r e n t a l w e i g h t and' the  development  r a t e o f the young.  groups a n a l y z e d g r a p h i c a l l y  (fig.  I n the t h r e e  control  10) the s l o p e o f . t h e  a p p r o x i m a t e s .625 w h i c h i s i n a g r e e m e n t w i t h t h e  graph  previously  r e p o r t e d g r o w t h r a t e f o r P. m a n i c u l a t u s ( K i n g , 1 9 6 8 ) . (1949) and Dawson (1965) have b o t h r e p o r t e d t h a t r e a r e d Peromyscus  Dice  laboratory-  are m o r p h o l o g i c a l l y i n d i s t i n g u i s h a b l e  w i l d caught specimens  of t h e same age  class.  Consequently,  v a r i a t i o n i n t h e h y b r i d f o r m s i s assumed t o b e due other than the laboratory  environment.  from  to  factors  9  1 0  _ _ 0  1  2  3  4  5  6  7  ' 8  9  10  11  12  13  14  DATS  Fig.  10  Postnatal  g r o w t h i n w e i g h t f o r some i s l a n d and h y b r i d  Peromyscus  15  16  17  18  OCCIPITONASAL SKULL LENGTH BASAL LENGTH NASAL LENGTH DIASTEMA  ISLAND COX  .2.24(.07)  2.19(.08)  1.19(.04)  0.80(.03)  TRIANGLE  2.95(.07)  2.28(.08)  1.24(.05)  0.84(.03)  HYBRID  2.97(.Q8)  2.27(.Q8)  1.27(.Q5)  0.84(.04)  Table  8  C o m p a r a t i v e c r a n i a l m e a s u r e m e n t s of Cox and T r i a n g l e • P e r o m y s c u s and t h e i r h y b r i d y o u n g .  The  hybrid  mating classes  litters  ( t a b l e 16A,  f r o m T r i a n g l e , Cox, as d i d t h e The  one  and  P.  age  o f 25  (n=5)  and  the  appendix l ) .  austerus,  large  mice  p r o d u c e d one  b e t w e e n P.  m.  doylei  o f f s p r i n g that died at  no; g r o s s m o r p h o l o g i c a l  a tendency f o r the h y b r i d  eyes of the  the  Subsequent  deformation to  the  i n t e r n a l organ.  Comparison between the  to develop  litters  Vansittart islands.  days w i t h o u t ' e v e r h a v i n g o p e n e d i t s e y e s .  e y e s o r any  ( t a b l e s 15A,  h y b r i d and  litters  16A,  c o n t r o l group i n d i c a t e  to take a longer  appendix l ) . For  h y b r i d g r o u p opened a t a t i m e  l a t t e r part The  The  group  Lanz i s l a n d s produced h e a l t h y  l a r g e - s m a l l phenotype cross m.  r e s u l t o f two  s m a l l mice f r o m B a l a c l a v a a n d  autopsy'.indicated  the  are  of the  t o study' the b e h a v i o r a l ' i n h e r i t a n c e i s l a n d m i c e . ' P e r o m y s c u s m.  comparable  to  control  group.  c h a r a c t e r i s t i c s i n these  triangularis is fairly  t r i a n g u l a r i s have a n ' i n c o n s p i c u o u s l y  the  p r e s e n t e d a unique chance  among' t h e ' Peromyscus' o f B r i t i s h ' C o l u m b i a .  c o a r s e h a i r s and  example,  range e s t a b l i s h e d f o r the  t r i a n g u l a r i s / c a r l i hybrids  period  only  does  bicolored t a i l  with  pronounced s c a l e s but  Not  distinctive  i t has  the  habit  of  burying i t s e l f  i n the s h a l l o w sawdust l a y e r i n t h e bottom  of  the cage.  T h i s b u r r o w i n g b e h a v i o r i s b e l i e v e d t o be r e l a t e d  to  t h e s e m i - f o s s o r i a l mode o f e x i s t a n c e t h a t t h e s e m i c e a r e  f o r c e d t o adopt upon T r i a n g l e I s l a n d .  Cox I s l a n d  Peromyscus,  on t h e o t h e r h a n d , a r e n o t p r o n e t o b u r r o w i n t h e s a w d u s t ; i n s t e a d they c o n s t r u c t a surface, n e s t t y p i c a l t o t h a t of a l l other i s l a n d populations. carli yet  The h y b r i d s f r o m t h i s  c r o s s had t h e phenotype  triangularis/  of the Triangle parent (table  t h e y showed no t e n d e n c y t o b u r r o w ;  8)  The b u r r o w i n g t r a i t was  neither learned'or inherited.' ' Contrary to the laboratory breeding successes established for  the c o n t i n e n t a l Peromyscus  ( D i c e , 1 9 3 3 , 1968; Dawson 1965;  Rood, 1 9 6 6 ) , "those o f t h e i n s u l a r m i c e f r o m B r i t i s h are  Columbia  very poor. As a c h e c k o f t h e e f f e c t o f t h e l a b o r a t o r y e n v i r o n m e n t upon  the  b r e e d i n g s u c c e s s , s i x p a i r s of Peromyscus  (table  m a i n t a i n e d f o r t h r e e months ( F e b r u a r y - A p r i l , kept outdoors. first yeafl  litters  trap results  were c o n c e i v e d sometime  1970) i n c a g e s  i n d i c a t e d that the  i n early April  o f t h e same  An a d d i t i o n a l t h r e e m o n t h ' t e s t was u n d e r t a k e n u s i n g t h e  same animals.' the  I n the f i e l d  9) were  The same c a g e s and d i e t were m a i n t a i n e d b u t  a n i m a l s were-'brought. i n t o t h e l a b o r a t o r y and k e p t on a  16 h o u r l i g h t a n d 8 h o u r d a r k l i g h t  regime.  The r e s u l t s o f  t h e s e two  t h r e e month t e s t s were a l l n e g a t i v e .  environmental temperature  and  the c o n t r o l l e d  The  change i n  light  c o n d i t i o n d i d n o t i n d u c e b r e e d i n g i n any o f the t e s t  pairs.  Triangle x Balaclava T r i a n g l e x Lanz Cox x L a n z V a n s i t t a r t x Hope Texada x  Triangle  T e x a d a x Thormanby Table  9  Test crosses to determine  ultimate breeding  A d d i t i o n a l f a c t o r s i n f l u e n c i n g mate s e l e c t i o n w e r e u s i n g a two way  t e s t f o r female  s e l e c t i o n o f male  M a l e m i c e were p l a c e d . a t t h e end o f two box.  A g e n t l e f l o w o f a i r was  odor  choices.  The  for  p e r i o d s of f o u r hours  a c t i v i t y , was  recorded  (table  t h a t female  o f t h e Y c h o i c e box  a n i m a l s were m a i n t a i n e d i n t h e on f o u r c o n s e c u t i v e d a y s .  m o n i t o r e d f o r t h e 16 h o u r t e s t  10) as the p e r c e n t a g e  i n e a c h arm  companion.  arms o f a Y c h o i c e  m a l e s were a l t e r n a t e d f r o m s i d e t o s i d e e v e r y d a y . female  tested  p a s s e d o v e r t h e s e m a l e s so  a female p l a c e d i n the v a c a n t passage two  conditions.  of the c h o i c e  of t o t a l box.  that had  apparatus The  The and  activity for  FEMALE  MALE .  Triangle  2.  Texada  Hope  24$  Triangle  76%  Doyle  9%>  Texada 3.  10  Mate  7%  Cox  93% 44%  Cox  56%  91%°  Vansittart V a n c o u v e r <^ (mainland) ^ V a n c o u v e r  Table  MALE  FEMALE  16%  48%  Triangle  52%  •Cox  preferences.  I s l a n d f e m a l e s show a s t r o n g  preference  f o r a male t a k e n f r o m  the  same i s l a n d ( t a b l e 10, c a s e s n o s . 1, 2, 3, 4, 5.) a n d i f  one  i s not a v a i l a b l e then the choice  male f r o m t h e n e a r e s t  i s made i n f a v o r  g e o g r a p h i c l o c a t i o n ( t a b l e 1 0 , n o s . 4, 6)  W h e t h e r an o l f a c t o r y o r a n a u d i t o r y s e l e c t i v i t y has n o t been d e t e r m i n e d .  cue i s r e s p o n s i b l e  i n f l u e n c e upon t h e c r o s s b r e e d i n g  i n my l a b o r a t o r y  f o r this  However, i t i s o b v i o u s  t h a t a d e f i n i t e p a t t e r n o f s e l e c t i v i t y e x i s t s and t h i s have a s t r o n g  of a  could  performances  experiments.  , The l o w l e v e l  of h y b r i d i z a t i o n , t h e d e l a y e d  development  of t h e h y b r i d y o u n g , a n d t h e s e l e c t i v e b r e e d i n g g r o u p s a r e indicative island  complex.  chapters, able  of strong This  i s o l a t i n g b a r r i e r s that e x i s t w i t h i n the r e i n f o r c e s my v i e w , a l s o  that the current  stated i n other  taxonomic arrangements are q u e s t i o n -  and t h a t e v o l u t i o n b e y o n d t h e s u b s p e c i e s l e v e l i s  perhaps a r e a l i t y  i n this  case.  THE PHALLUS AND OF THE  I T S BEARING UPON THE  CLASSIFICATION  INSULAR PEROMYSCUS OF B R I T I S H COLUMBIA  W i t h i n t h e genus P e r o m y s c u s p h a l l i c m o r p h o l o g y h a s taxonomic importance a t the s p e c i e s l e v e l Eighteen  ( H o o p e r , 1958, 1 9 5 9 ) .  s p e c i e s i n v e s t i g a t e d b y Hooper ( 1 9 5 8 ) showed v a r i a t i o n  i n absolute  or r e l a t i v e  size  i n p a r t s of the p h a l l u s .  C l o s e l y r e l a t e d s p e c i e s h a d t h e most m o r p h o l o g i c a l l y phalli.  similar  I n t e r s p e c i f i c v a r i a t i o n h a s b e e n d e s c r i b e d as b e i n g  absent or very  s u b t l e , r e q u i r i n g a l a r g e sample and more  r e f i n e d procedures i n order t o f i n d  statistically  valid  c o r r e l a t i o n s between p h a l l i c v a r i a t i o n and the s u b s p e c i f i c taxa.  Phallic  d i f f e r e n c e s b e t w e e n s p e c i e s has b e e n t h e b a s i s  of the c r i t i c i s m  o f t h e Osgood (1909) c l a s s i f i c a t i o n o f t h e  genus P e r o m y s c u s (Hooper and M u s s e r , 1 9 6 4 ) .  Hooper ( 1 9 5 8 ,  p. 2 2 ) , u t i l i z i n g t h e p h a l l u s c h a r a c t e r i s t i c s o f P.  crinitus,  p l a c e s i t i n t h e subgenus P e r o m y s c u s and n o t i n t h e subgenus Haplomylomys as d i d Osgood. should probably  He a l s o c l a i m s t h a t  hylocetes  be removed f r o m t h e P e r o m y s c u s subgenus and  p l a c e d i n t h e Haplomylomys g r o u p .  I f the degree o f v a r i a t i o n  d e s c r i b e d i n p h a l l u s m o r p h o l o g y i s a c c e p t e d as s p e c i e s s p e c i f i c w i t h i n t h e subgenus P e r o m y s c u s t h e n c o m p a r a b l e v a r i a t i o n s b e t w e e n p o p u l a t i o n s , s u c h as t h e  inter-island  s i t u a t i o n e x a m i n e d i n my s t u d y , w o u l d j u s t i f y t a x o n o m i c  r e - e v a l u a t i o n of those  MATERIALS AND  METHODS  Specimens used  i n t h i s s t u d y were o b t a i n e d f r o m t r a p  b r e e d i n g colony deaths t h e p e n i s was p e n i s was  groups.  of a d u l t males.  removed and  The  d i s t a l p a r t of  s t o r e d i n 70 p e r c e n t e t h a n o l .  Further clearing  d a y s p r i o r t o t r a n s f e r r i n g f o r s t o r a g e i n 100  glycerine. KOH  The  p h a l l u s w a s . s t a i n e d on t h e t h i r d day  t r e a t m e n t by t h e a d d i t i o n o f a few d r o p s  A l i z e r i n Red  s t a i n t o t h e KOH  solution.  The  l a y e r , p o s s e s s i n g the s p i n e s c h a r a c t e r i s t i c phalli,  i s easily lost during this  consider that a pretreatment  i n v e s t i g a t i o n i s i n t e n d e d t o be  The  of the p r e p a r e d  specimens:  Distal  tract length-  of the  outer  tissue  of P e r o m y s c u s  t h i s problem.  I  now  prior The  n o t c a l i b r a t e d s i n c e the comparative  assumed t h a t e a c h p h a l l u s w o u l d d i s t o r t s i m i l a r manner.  percent  Alkaline  i n f o r m a l i n f o r t h e day  d i s t o r t i o n was  solution  c l e a r i n g p r o c e s s and  to a l c o h o l p r e s e r v a t i o n would a l l e v i a t e degree of p h a l l i c  of  of  was  o b t a i n e d by p l a c i n g i t i n 50 p e r c e n t aqueous g l y c e r i n e f o r two  The  c l e a r e d by s o a k i n g i t i n a 2 p e r c e n t s o l u t i o n  potassium hydroxide f o r three days.  and  in a  i n n a t u r e and  I  quantitatively  f o l l o w i n g m e a s u r e m e n t s were made on  each  the l e n g t h from the t i p to the p r o x i m a l p o i n t of  flexure.  Glans l e n g t h -  the  distance  from the  t i p to the  where t h e p r e p u c e a t t a c h e s  to  point  the  phallus. Glans diameter-  the w i d t h  of t h e  glans  at the  widest  point.  I n the  was  r e g i o n a r o u n d the. m e a t u s  the  specimens examined  this  urinarius. Baculum length--  the  greatest  straight line  l e n g t h of  the  baculum. Cartilage  t i p length-  the  translucent cartilagenous t i p  on t h e  distal  end  o f the b a c u l u m  measured to the n e a r e s t its  s m a l l s i z e and  standard  proximal  two  types,  ( t y p e B, i n the  end  o f t h e b a c u l u m was  sample  C,  D,  E)  l a p p e t v a r i a t i o n s ( f i g . 12, F, glans  a p p e a r s i n two  (fig.  10,  spines  H,  i t was  I).  forms:  I n order  to  large  the  The  distal  smooth and  t o determine the  of  o r o f a knob  as w e l l as two  G).  value.  c a t e g o r i z e d as t o one  Three v e n t r a l l a p p e t t y p e s a r e  ( f i g . 12,  due  resulting  of q u e s t i o n a b l e  e i t h e r t h a t o f a diamond ( t y p e A)  f i g . 12).  but  e r r o r t h i s measurement i s  b e l i e v e d t o be The  the  .01  was  recognized  dorsal t i p of  the  the b u l b o u s presence  necessary to observe f r e s h m a t e r i a l  and  of  types  Fig.  11  The  phallus  of Peromyscus  maniculatus  2.  VENTRAL LAPPETS  3.  DORSAL LAPPETS  4.  DISTAL PORTION OF GLANS  Bulbous  Fig.12  PHALLIC CHARACTERS  c o n s e q u e n t l y no p r e s e r v e d s p e c i m e n s a r e on d e p o s i t . p r o c e s s e d p h a l l i a r e on d e p o s i t the U n i v e r s i t y  i n The V e r t e b r a t e  Museum c  of B r i t i s h Columbia.  LOCATION  ISLAND  PEROMYSCUS MANICULATUS SUBSPECIES*  Doyle  50°127 N.W.  Hurst  50°127  saxamans  Balaclava  50°127  balaclava  Nigei  50°127  isolatus  Vansittart  50°127  **  Hope  50°127  balaclava  Cox  50°128  carl i  Lanz  50°128  carli  Triangle  50°129  triangularis  Texada  49°124  georgiens i s  Thormanby  49°123  georgiensis  *  The  doylei  Cowan and G u i g e t , 1965 Peromyscus from t h i s i s l a n d named a t t h i s t i m e .  Table  11  RESULTS AND  Phallic  have n o t b e e n d e s c r i b e d  or  specimens examined.  CONCLUSIONS  The g e n e r a l  morphology of the p h a l l u s  of the i n s u l a r  P e r o m y s c u s s t u d i e d does n o t d i f f e r s i g n i f i c a n t l y f r o m t h e  d e s c r i p t i o n g i v e n f o r the (Hooper, 1958). u r i n a r i u s and may  not  The  glans  tapers  terminate  c o n t i n e n t a l forms of t h i s i s bulbous around the  in a slight swelling.  thorns,  edge of t h e  meatus u r i n a r i u s .  Spines,  cover the p h a l l u s p r o x i m a l The  osseous rod w i t h a b l u n t d i s t a l  t i p and  communication) c o n s i d e r s  support t h i s view.  f u n c t i o n o f age.  v a r i a t i o n i n the  f o u n d i n a l l age  Hooper  category  D e s c r i p t i o n of Doyle  r e s u l t s d6  i s not  portion a  s t u d y were  adult pelage  and  class three, adult, i n  determination  ( F o s t e r , 1965,  p.  69).  Island  p h a l l u s i s one  Two  t h i r d s o f the  t i p to the  o f the  l a r g e s t present  sample d i d n o t  glans.  The  occur,  but  i n a l l cases the  baculum i s not  w i t h e i t h e r the 12)  c u r v e d and  island  are m e d i a l l y  cone and  terminates A, f i g . 12)  survey.  swollen p r o t r u s i b l e divided.  ridge v a r i a t i o n s ,  organ i s b i l a t e r a l l y  diamond ( t y p e  variation.  i n this  have t h e  v e n t r a l lappets  B o t h t y p e s o f d o r s a l l a p p e t s , the  fig.  terminal  phalli  The  The  (personal  terminal  in this  o r had  s u f f i c i e n t t o o t h w e a r r o r e g i s t e r as s c a l e f o r age  My  g r o u p s and  A l l specimens d i s c u s s e d  e i t h e r o f known a d u l t age  the F o s t e r  proximally  t h a t t h i s v a r i a t i o n i n the  The  fluted  slender  terminates  p o r t i o n o f the b a c u l u m i s a f u n c t i o n o f age.  or  resembling  to the  baculum i s a  i n e i t h e r a d i a m o n d o r knob s h a p e d s w e l l i n g .  o f the b a c u l u m was  meatus  to a smooth p r o t u s i b l e t i p w h i c h may  minute rose  not  genus  symmetrical.  a t the p r o x i m a l  end  or the b l u n t  (typeB ,  Hurst Island The  phalli  a r e t h e same a s t h o s e  from Doyle i n o v e r a l l  y e t the v e n t r a l l a p p e t s are of the non-divided g i v i n g the p h a l l u s a d i s t i n c t i v e  trait.  size  variety,  The d o r s a l l a p p e t s  are o f t h e r i d g e type and t h e organ i s s y m m e t r i c a l .  The  b a c u l u m i s s t r a i g h t and t e r m i n a t e s w i t h t h e d i a m o n d c o n f i g u r a t i o n . Balaclava Island A small p h a l l u s of uniform diameter imparts  a unique c h a r a c t e r to t h i s  along  i t s total  length  i s l a n d Peromyscus.  In  a l l b u t one o f t h e s a m p l e s t h e p r o t r u s i b l e p a r t o f t h e g l a n s terminates without  any s w e l l i n g .  Unlike the other  groups s t u d i e d a l l t h r e e v a r i a t i o n s o f the v e n t r a l  island lappets  o c c u r , w h e r e a s t h e d o r s a l l a p p e t s were a l l o f t h e cone t y p e . In  many c a s e s  in  a l l cases  t h r e e o r more cone l a p p e t s o c c u r p e r p h a l l u s a n d the p h a l l u s i s non-symmetrical.  The b a c u l u m i s  s t r a i g h t a n d t h r e e q u a r t e r s o f t h e sample have t h e b l u n t baculum  terminus.  Nigei Island P e r o m y s c u s f r o m N i g e i have a l a r g e p h a l l u s w i t h a r e l a t i v e l y longer glans s e c t i o n than those Islands.  The g l a n s t e n d s  from e i t h e r Doyle o r Hurst  t o be b u l b o u s  p r o x i m a l t o t h e meatus u r i n a r i u s . appears square, glans are f l a t . than t h a t found  i.e.  over  the c e n t r a l s e c t i o n  I n c r o s s s e c t i o n the glans  the d o r s a l and v e n t r a l s u r f a c e s o f t h e  The p r o t r u s i b l e t i p o f t h e g l a n s i s l o n g e r i n any o t h e r i s l a n d  sample.  f r o m t h e meatus u r i n a r i u s t o t h e d i s t a l  The mean d i s t a n c e  t i p o f the glans i s  0.34  cm  other  and  a v e r a g e s 0.08  group.  ventral  r e s t are  are  i n the  o f the  ratio  described f o r Hurst  The  sample a r e u n d i v i d e d  divided type.  of 7 : 1 , r i d g e t o cone. The  any  are b u l b o u s i n a l l c a s e s .  t h i r d s of the  partially  i n a l l cases.  Vansittart  longer than t h a t found i n  distal tips  l a p p e t s i n two  the  occurs  The  cm  The  dorsal  Bilateral  baculum i s o f the type  and  lappets  symmetry previously  Island.  Island  T h i s i s the  s m a l l e s t of a l l the p h a l l i  of the p h a l l i  p o s s e s s the nonbulbous g l a n s  l a p p e t s are  separable  lappets  of the  are  groups measured.  cone m o r p h o l o g y t h e  into a l l three  cone and  t i p and v e n t r a l  divisions.  ridge types.  Most  The  In those  dorsal  with  the  symmetry i s b i l a t e r a l .  Hope I s l a n d The to  pronounced decrease i n the p h a l l u s d i a m e t e r p r o x i m a l l y the meatus u r i n a r i u s p r o d u c e s a c o n d i t i o n o p p o s i t e  described for Nigei Island. terminates  a t the d i s t a l  In a l l cases,  t i p with a slight  the  divided.  swelling.  are  of the  o n l y f o u r of the n i n e t e e n - a p p e a r e d  Approximately ridge type.  t h r e e q u a r t e r s of the The  that  phallus  l a p p e t s on t h e v e n t r a l s u r f a c e a r e n e a r l y a l l o f t h e divided type,  to  The non-  partially  dorsal  b a c u l u m i s s t r a i g h t and  lappets terminates  w i t h t h e diamond s h a p e d h e a d . Cox This  Island l a r g e p h a l l u s type  appears v e r y t h i n .  The  t e n d s t o be  laterally  compressed  n o n - b u l b o u s t e r m i n u s t o the  glans  and  predominates i n t h e sample.  The v e n t r a l l a p p e t s  i n t o the n o n - a n d - p a r t i a l l y d i v i d e d c a t e g o r i e s lappets  a r e , w i t h h u t one e x c e p t i o n ,  One o f t h e b a c u l a shaped  appeared curved.  are s p l i t  and t h e d o r s a l  of the r i d g e v a r i e t y . A l l bacula  h a d t h e diamond  terminus.  Lanz I s l a n d The  p h a l l u s i s i n t h e same s i z e c l a s s a s t h a t o f Cox I s l a n d b u t  in this  case the tendency f o r l a t e r a l  compression i s greater.  N o n - d i v i d e d v e n t r a l l a p p e t s p r e d o m i n a t e a n d one t h i r d o f t h e dorsal lappets  a r e o f t h e cone t y p e .  The cone l a p p e t s  a non-symmetrical appearance t o the p h a l l u s . a s l i g h t c u r v e w i t h t h e b o t h ends v e n t r a d  give  The b a c u l u m h a s  t o the p h a l l u s  center  line. Triangle  Island  P e r o m y s c u s f r o m T r i a n g l e I s l a n d have t h e l a r g e s t p h a l l i . symmetrical  organ terminates  w i t h a bulbous t i p t o the g l a n s .  A l l v e n t r a l l a p p e t s , w i t h b u t one e x c e p t i o n , medially divided.  This  The e x c e p t i o n ,  are deeply  a type D ( f i g . 1 2 ) , i s  d i v i d e d but t o a f a r l e s s e r degree.  The b a c u l u m i s s t r a i g h t  w i t h the b l u n t v a r i e t y of t e r m i n a l s w e l l i n g . The  s o u t h e r n i s l a n d group i s r e p r e s e n t e d  i n this  study by  s p e c i m e n s t a k e n f r o m Texada and Thormanby I s l a n d s . The  phallus  i s e s s e n t i a l l y t h e same f o r b o t h p o p u l a t i o n  I t i s a s m a l l p h a l l u s b u t l a r g e r t h a n t h a t o f t h e Hope, B a l a c l a v a and V a n s i t t a r t p o p u l a t i o n s  that are, i n other  samples.  characters, quite similar.  The t e r m i n u s o f t h e g l a n s i s  e i t h e r bulbous or p l a i n , w i t h equal frequency.  The v e n t r a l  i  l a p p e t s a r e n e a r l y a l l u n d i v i d e d and t h e d o r s a l l a p p e t s a r e mostly  o f t h e diamond c o n f i g u r a t i o n .  Among t h e 95 p h a l l i  examined i n t h i s  study  66  have a b u l b o u s t e r m i n u s t o t h e g l a n s , 66 p e r c e n t type  d o r s a l l a p p e t s , 56 p e r c e n t  l a p p e t s a n d 82 p e r c e n t diamond c o n f i g u r a t i o n . lappets  have r i d g e  have u n d i v i d e d v e n t r a l  have a b a c u l u m t e r m i n a t i n g w i t h t h e The p r e p o n d e r a n c e o f u n d i v i d e d v e n t r a l  c o n t r a s t s w i t h t h e s t a t e m e n t b y Hooper (1958) t h a t  i n P. m a n i c u l a t u s i s uncommon. those  percent  the medially c l e f t v e n t r a l lappet  condition  The p h a l l i c measurements a r e c o n t r a s t e d  made b y Hooper ( 1 9 5 8 ) ;  HIND FOOT  N  they  are s i g n i f i c a n t l y  CARTILAGE GLANS BACULA T I P A  with different.  B  C  Hooper 1958 P.maniculatus  3  20  7.4  8.3  0.8  37  16  41  P.leucopus  5  21  7.8  9.0  0.8  37  24  43  P.polionotus  3  17  6.1  6.6  0.5  36  18  39  Island  samples  P.maniculatus  95  22. 5  8.5  9.7  1.25  37.8  22.4  43  Vansittart Is.  11  20  7.4  8.4  1.3  37  24.4  42  Legend A glans/foot ratio B diameter of glans/glans C baculum/foot r a t i o Table  12  length  ratio  P h a l l i c measurements f o r s e v e r a l s p e c i e s o f P e r o m y s c u s .  Texada Thormanby Triangle Nigei Hurst Doyle Cox Lanz Balaclava P i g 13  Hope Vans i t t a r i j  6.5  7.0  7.5  8.0  8.5  9.0  COMPARATIVE GLANS LENGTHS  9.5  1.00  1.05  Thormanby Texada Triangle  1.17 Doyle  I  1  Hurst Nigei  r  „  #  -7?~  1.29 1.28  Lanz Cox Hope  j  1  Fig  Balaclava  14  COMPARATIVE BACULAR LENGTHS  Vansittart  72 7.5 mean  8.0  3. 5  9-0  9. 5  1.00  1.05  2 standard f^j  "  range  1  e  ^  r  r  o  r  s  A f t e r D i c e and Ler.aas, 1 936.  1 .10  1. 1 5  The  phallus i s significantly larger  f o r P. m a n i c u l a t u s  (Hooper 1 9 5 8 ) .  (<.0l)  t h a n t h e mean  However, t h e  distal  t r a c t l e n g t h and t h e b a c u l u m l e n g t h show a d i r e c t to the s i z e of the a n i m a l . (figs.  The  apparent  13, 14) a r e a f u n c t i o n o f t o t a l  c a n n o t be  c o n s i d e r e d as an i n d e p e n d e n t  phylogeny  or taxonomy.  size  correlation  differences  animal size variable  and  indicative  R e g r e s s i o n a n a l y s i s of the  of  insular  d a t a i n d i c a t e s a 91 p e r c e n t c o r r e l a t i o n • b e t w e e n t h e b a s a l skull  l e n g t h , (used i n t h i s  a n i m a l s i z e ) and  t h e l e n g t h o f the b a c u l u m .  i s e x p r e s s e d as f o l l o w s : length +  s t u d y as an i n d i c a t o r  Basal skull  of  This  total  relationship  length=0.763 x baculum  (-0.658).  P. p o l i o n o t u s , b e l i e v e d t o be P. m a n i c u l a t u s  a recent derivative  c o m p l e x (Hooper 1958)  shows a c l o s e r  s h i p t o c o n t i n e n t a l P. m a n i c u l a t u s t h a n i s shown b y i s l a n d samples.  T h i s p r e s e n t s two  ( a ) t h e sample s i z e  (n=3)  not r e p r e s e n t a t i v e of the  of the  relationthe  p o s s i b l e hypotheses,  u s e d by Hooper i s i n a d e q u a t e  either and  s p e c i e s m a n i c u l a t u s , o r (b) t h e  i n s u l a r Peromyscus of B r i t i s h  Columbia  are not of the  species  maniculatus. The  f o r m e r h y p o t h e s i s seems t h e more p l a u s i b l e  i fi t  were n o t t h a t t h e V a n s i t t a r t m i c e , a l t h o u g h a p p r o x i m a t i n g c o n t i n e n t a l Peromyscus m a n i c u l a t u s from the p h a l l i c  dimensions  i n size, differ considerably  l i s t e d by Hooper.  d i f f e r e n c e s i n the glans diameter  the  I f the  i s a real difference  observed and  n o t an a r t i f a c t  o f t h e s m a l l sample  s i z e then the v a r i a t i o n  spans t h e r a n g e o f t e n o f t h e s p e c i e s Notable  exceptions  f r o m my i n s u l a r  to the g e n e r a l morphology  sample a r e a s f o l l o w s :  medially c l e f t v e n t r a l lappets phalli  c a t e g o r i z e d by Hooper.  D o y l e I s l a n d m i c e have  in a l l phalli  o f the mice from B a l a c l a v a , b e s i d e s  t h i c k e r t h r o u g h t h e g l a n s , have cone t y p e d o r s a l s u r f a c e , and a l l t h r e e types present.  E i g h t y - s i x percent  of t h e p h a l l u s  examined.  being  The  s h o r t and  l a p p e t s on t h e  of v e n t r a l lappets are  o f t h e T r i a n g l e sample have  b a c u l a r t e r m i n a t i n g p r o x i m a l l y w i t h t h e b l u n t end t y p e , a n d all  possess d i s t i n c t  phalli  cleavage  of the v e n t r a l l a p p e t s .  The  o f T r i a n g l e , T e x a d a , a n d B a l a c l a v a m i c e show a h i g h  percentage of the supposedly j u v e n i l e t r a i t ,  the blunt  baculum  terminus. Results  of a T - t e s t a n a l y s i s of the baculum and d i s t a l  t r a c t measurements ( f i g s .  19A, 20A, a p p e n d i x l ) p r o v i d e  method b y w h i c h t o e v a l u a t e island  r e l a t i o n s h i p s . The  samples are ranked i n t h e degree o f d i v e r g e n c e  b a c u l u m and d i s t a l are  phallic  tract lengths.  o b t a i n e d by t a l l y i n g  The d i v e r g e n c e  i n the baculum and d i s t a l  example  inter-  in  values  t h e number o f a l t e r n a t i v e i s l a n d  samples t h a t are s i g n i f i c a n t l y d i f f e r e n t a t the 5 level  another  t r a c t mean v a l u e s .  percent For  i f sample A i s s i g n i f i c a n t l y d i f f e r e n t i n b a c u l u m  l e n g t h f r o m s a m p l e s B, C, and i n d i s t a l t r a c t l e n g t h f r o m s a m p l e s x and z t h e n sample A w o u l d have a d i v e r g e n c e o f 4.  value  The p h a l l i conversely,  o f t h e s m a l l m i c e a r e t h e most d i v e r g e n t a n d ,  the p h a l l i  o f l a r g e mouse p h e n o t y p e s a r e somewhat  l e s s v a r i a b l e i n morphology. intermediate  The h y b r i d o f f s p r i n g  i n p h a l l i c measurements.  no b r e a k i n t h e c l i n e o f d i v e r g e n c e the  t o be a b l e n d i n g samples.  This  relationship  of a l l characters  There i s , however,  values  small from the l a r g e phenotypes.  are t r u l y  that  differentiate  The h y b r i d p h a l l u s a p p e a r s  f o u n d among t h e t e n i s l a n d  i s perhaps i n d i c a t i v e  of a close  phylogenetic  e x i s t i n g w i t h i n t h i s group o f r o d e n t s .  Hence t h i s  study  demonstrates that extensive  i n p h a l l i c morphology e x i s t s  among t h e i s l a n d  variation  Peromyscus.  I f t h e c r i t e r i a e s t a b l i s h e d b y Hooper were t o b e a p p l i e d i n this  situation  designated  a full  characteristics applicable  each of the i n s u l a r species.  The c l i n e  w o u l d be  i n some o f t h e  t e n d s t o r e f u t e p h a l l i c taxonomy  to t h i s situation.  differentiated  populations  No d i s t i n c t g r o u p s c a n be  a n d no c o r r e l a t i o n  between p h a l l i c  and m o r p h o l o g y a r e a p p a r e n t e x c e p t , i n p h a l l i c characters  as b e i n g  characters  as n o t e d , where t h e c l i n e s  a r e p o l a r i z e d b e t w e e n t h e l a r g e and  s m a l l mouse p h e n o t y p e s . DIVERGENCE VALUE  ISLAND  SAMPLE S I Z E  Hope and V a n s i t t a r t  30  19  Thormanby a n d T e x a d a  17  17  8  16  7  14  9  13  16  12  6  11  3  5  Balaclava Cox  -  Doyle N i g e i and T r i a n g l e Lanz Hybrid  (Triangle/Cox)  Table  13  D i v e r g e n c e between i s l a n d samples u t i l i z i n g t h e c o m b i n e d b a c u l u m and d i s t a l t r a c t l e n g t h measurements.  KARYOTYPIC VARIATION I N INSULAR  Karyology  PEROMYSCUS  c a n be i n s t r u m e n t a l i n t h e a n a l y s i s o f many  taxonomic problems.  This technique  i s , h o w e v e r , o n l y one o f  many c h a r a c t e r s t o be c o n s i d e r e d i n t h e o v e r a l l e v a l u a t i o n o f any group. t h a t made b y O r l o v structure  To make a g e n e r a l i z a t i o n ,  (1970),  that f i v e mutations  s u c h as  i n chromosome  i s s i g n i f i c a n t i n s p e c i e s d i f f e r e n t i a t i o n i s i n my  opinion unjustifiable.  I n the non-Robertsonian  chromosome e v o l u t i o n t h a t o c c u r s mutation  taxonomic  system of  i n a l l P e r o m y s c u s , one  can t h e o r e t i c a l l y present major problems t o the  meiotic  process.  An example o f t h e u s e s o f k a r y o l o g y the Peromyscus oreas karyotype  i s the a n a l y s i s of  c o m p l e x w h i c h i s composed o f two d i s t i n c t  g r o u p s (N. B r a d s h a w , p e r s o n a l c o m m u n i c a t i o n ) .  c o a s t a l P. o r e a s  a r e k a r y o t y p i c a l l y i d e n t i c a l t o t h e P. m a n i -  c u l a t u s , h a v i n g a l o w number o f s o m a t i c (F.N.), y e t are supposedly distinctive.  The  chromosome arms  m o r p h o l o g i c a l l y and b e h a v i o r a l l y  The s e c o n d o r e a s  group, found  inland,  isa  member o f t h e h i g h F.N. s e r i e s a n d i s p e r h a p s c l o s e l y  related  t o t h e l a r g e morph i n s u l a r r a c e s t h a t h a v e b e e n c h a r a c t e r i z e d in this oreas  study.  to f u l l  The s t u d y r e s p o n s i b l e f o r t h e e l e v a t i o n o f species  ( S h e p p e , 1 9 6 1 ) h a s a p p a r e n t l y n o t gone  f a r enough, s i n c e t h e m o r p h o l o g i c a l does n o t r e f l e c t t h e k a r y o t y p i c Lawlor  and b e h a v i o r a l v a r i a t i o n  situation.  ( 1 9 7 1 ) h a s shown c o n s i d e r a b l e k a r y o t y p i c v a r i a t i o n  among t h e P e r o m y s c u s i n h a b i t i n g t h e i s l a n d s o f t h e G u l f o f California.  These v a r i a t i o n s . a r e s p e c i e s s p e c i f i c and l i t t l e  i n f o r m a t i o n r e g a r d i n g s u b s p e c i f i c and i n t r a p o p u l a t i o n v a r i a t i o n i s recorded.  My s t u d y  seeks t o answer t h e f o l l o w i n g  three questions  using a l l o p a t r i c  closely related  Peromyscus.  populations  1) What i s t h e e x t e n t o f k a r y o t y p e w i t h i n these  insular populations  of a p p a r e n t l y  variation that  exists  o f P e r o m y s c u s and what  e l e m e n t s a r e t h e most f r e q u e n t l y i n v o l v e d i n t h e e v o l u t i o n of t h e k a r y o t y p e ? 2) Can t h e k a r y o t y p e in determining  be c o r r e l a t e d w i t h o r be i n s t r u m e n t a l  t h e s p e c i e s and s u b - s p e c i e s r e l a t i o n s h i p s ?  That i s t o say, can s t a s i p a t r i c  s p e c i a t i o n account f o r the  interisland variation? 3 ) Can d a t a  f r o m i s l a n d g r o u p s , s u c h as t h o s e  studied,  pro-  vide f u r t h e r i n s i g h t i n t o the r e l a t i o n s h i p s of c o n t i n e n t a l populations? Such c h a l l e n g i n g q u e s t i o n s  as t h e p o s s i b i l i t y  of i n t r a -  i s l a n d deme r e l a t i o n s h i p s and t h e p o p u l a t i o n d e n s i t y - g e n e t i c relationships  c a n n o t as y e t be a s c e r t a i n e d .  To u n d e r t a k e  s u c h a s t u d y w o u l d r e q u i r e e x t e n s i v e s t u d y upon e a c h  island.  My hope o f f i n d i n g a g e n e t i c p a t h w a y l e f t b y e m i g r a t i n g founding  s t o c k s has been abandoned.  c o a s t l i n e , the a l l o p a t r i c nature number o f i s l a n d t r a i l s  leaves  The c o m p l e x i t y  of the  o f t h e P e r o m y s c u s , and t h e  such a study  team i n v e s t i g a t i o n on a l o n g t e r m b a s i s .  to the realm of  MATERIALS AND METHODS All live  P e r o m y s c u s (n=326) were t r a p p e d u s i n g 8" Sherman  t r a p s and r e t u r n e d t o t h e l a b o r a t o r y a l i v e f o r a n a l y s i s .  Karyotypes  were o b t a i n e d f r o m bone marrow c e l l s  tissue cultures  of ear epidermal c e l l s .  method n e c e s s i t a t e s t h e s a c r i f i c e is  The bone marrow  of t h e specimen b u t i n t u r n  f a s t e r and i s o f e q u a l o r b e t t e r q u a l i t y t h a n t h e t i s s u e  culture technique. use  and f r o m  However, b r e e d i n g s t u d i e s r e q u i r e d t h e  o f t h e e a r c u l t u r e t e c h n i q u e so t h a t t h e a n i m a l s o f  known k a r y o t y p e s later  c o u l d be s e l e c t e d f o r m a t i n g  crosses at a  date.  Bone marrow method. T h i s i s a m o d i f i e d v e r s i o n o f t h e F o r d and Hamerton (1956) C o l c h i c i n e - h y p o t o n i c c i t r a t e t e c h n i q u e . Lilly  "Velbe," E l i  and Co., was s u b s t i t u t e d f o r t h e c o l c h i c i n e .  peritoneal  injections  o f a 0.125% V e l b e  Inter-  s o l u t i o n , 0.01  ml/g  body w e i g h t , were made one h o u r b e f o r e t h e a n i m a l was t o be sacrificed.  The bone marrow c e l l s were f l u s h e d o u t and  a s p i r a t e d i n - t h e manner o u t l i n e d b y H s u and P a t t o n ( B e n i r s h k e , 1969).  Then t h e c e l l  minutes i n a water the water  s u s p e n s i o n was i n c u b a t e d f o r e i g h t  b a t h a t 38° F.  bath r e s u l t e d i n inadequate  l a c k of metaphaseplates. P e r o m y s c u s seem t o p o s s e s s standard procedures Peromyscus fauna. in  F a i l u r e to incubate i n cellular  s w e l l i n g and  I n t h i s respect the northern a cell  l i n e more r e s i s t a n t t o t h e  t h a n does t h e s o u t h e r n  continental  Such r e s i s t a n c e t o t h e t e c h n i q u e  occurred  l a b o r a t o r y p o p u l a t i o n s o f P. m a n i c u l a t u s s o n o r i e n s i s  a f t e r t h e y had b e e n m a i n t a i n e d on a l a b o r a t o r y d i e t f o r s e v e r a l months (Thomas, 1 9 6 8 ) . the i s l a n d  No  s u c h change was  in  c a p t i v e stocks maintained i n the l a b o r a t o r y f o r  o v e r two y e a r s .  Bone marrow c e l l s  t a k e n from b o t h the o l d ,  l a b o r a t o r y r a i s e d a n i m a l s and f r o m t h e n e w l y r e q u i r e d t h e warm b a t h t r e a t m e n t .  t h e o s m o t i c and  caught  I believe that  c o n d i t i o n i s a f u n c t i o n of t h e m e t a b o l i s m affects  noted  and  mice  this  t h a t the  diet  o t h e r p h y s i c a l p r o p e r t i e s of the  cell. T i s s u e c u l t u r e method. The' b a s i c t e c h n i q u e f o r t h i s p r o c e s s has been by Hsu was  (Hsu, 1 9 6 9 ) .  utilized  facilities S o c i e t y and  A m o d i f i e d v e r s i o n o f t h e Hsu  in this  technique  study i n o r d e r t o t a k e advantage  made a v a i l a b l e t o me  by t h e C a n a d i a n  t h e L a b o r a t o r y o f Dr. H.  of the m o d i f i e d procedure 1) Wash t h e l e f t  outlined  i s as  Stich.  of t h e  Cancer  A brief  outline  follows;  e a r o f t h e a n i m a l w i t h 10%  alcohol prior  to  2 removing  a 10-15  mm  t e r m i n a l p o r t i o n of the  2) Cut t h e e a r t i s s u e i n t o i n 4 ml.  s m a l l fragments  organ.  and  suspend i t  o f 0.25$ T r y p s i n f o r 1 h o u r .  3) D e c a n t t h e t r y p s i n and wash t h e e a r f r a g m e n t s of growth media. 800  i n 2 ml.  C e n t r i f u g e t h e sample f o r 5 m i n u t e s  r.p.m. D e c a n t t h e wash  at  solution.  4) R e s u s p e n d t h e e a r f r a g m e n t s  i n 4 ml. of f r e s h media  and  u s i n g a P a s t e u r p i p e t t e p l a c e 1 ml.  i n t o each of 4 L e i g h t o n  c u l t u r e tubes c o n t a i n i n g p r e c l e a n e d  coverslips.  5) C u l t u r e i n a n i n c u b a t o r every  s e t a t 36  o t h e r day.  6) I n 7-10 d a y s a m o n o l a y e r o f c e l l s cover  C. c h a n g i n g t h e m e d i a  slip.  At this  in a hypotonic parts d i s t i l l e d  should  form over t h e  t i m e remove t h e c o v e r s l i p and submerse  s o l u t i o n o f one p a r t g r o w t h m e d i a and t h r e e water.  7) A f t e r 8 m i n u t e s o f p r e t r e a t m e n t  remove t h e c o v e r s l i p a n d ,  a f t e r d r a i n i n g , p l a c e i t g e n t l y i n Carnoy's f i x a t i v e . the  Remove  c o v e r s l i p a f t e r 10 m i n u t e s and a l l o w t o d r y t h o r o u g h l y .  8) . S t a i n f o r 6 m i n u t e s u s i n g A c e t o O r c e i n 9) C l e a r t h e c o v e r s l i p w i t h a b s o l u t e  stain.  a l c o h o l baths  before  m o u n t i n g upon t h e s l i d e .  CLASSIFICATION  ARM RATIO  NOTES  METACENTRIC  4/8-5/8  biarmed  SUBMETACENTRIC (SM)  5/8-6/8  biarmed  SUBTELOCENTRIC ( S T )  6/8-7/8  problem group  TELOCENTRIC OR ACROCENTRIC (A o r T)  7/8-8/8  no v i s i b l e arms  "rabbit  metacentric  Table  14  ears"  subtelocentric (problem group)  Nomenclature for  chromosome t y p e s .  acrocentric  RESULTS AND  DISCUSSION  W i t h i n the i s l a n d type groups.  sample t h e r e e x i s t two d i s t i n c t  The s a m p l e f r o m t h e s o u t h e r n i s l a n d s  G e o r g i a S t r a i t g r o u p have a l o w number o f b i a r m e d and  c o n s e q u e n t l y a low f u n d a m e n t a l  number.  karyo-  of the chromosomes  The sample  from  t h e n o r t h e r n S c o t t I s l a n d s have a h i g h number o f b i a r m e d chromosomes and a c o r r e s p o n d i n g l y h i g h f u n d a m e n t a l The f u n d a m e n t a l arms p o s s e s s e d  number  number.  (F.N.) i s t h e number o f chromosomal  by t h e s o m a t i c  i n t e r m e d i a t e i s l a n d group,  chromosome  complement.  t h e G o r d o n and G o l e t a s  The  islands,  are r e p r e s e n t e d by i n t e r m e d i a t e k a r y o t y p e s , i n d i c a t i v e  perhaps  of a complex g e n e t i c h e r i t a g e . The d a t a p r e s e n t e d  ( t a b l e 16) show t h a t some k a r y o t y p i c  v a r i a t i o n e x i s t s w i t h i n t h e s o u t h e r n g r o u p even t h o u g h  these  m i c e a r e so s i m i l a r p h e n o t y p i c a l l y t h a t t h e y a r e c o n s i d e r e d one s u b s p e c i e s  (Hall,  1938).  The m a j o r d i f f e r e n c e o c c u r s i n  t h e Texada p o p u l a t i o n where a r e d u c t i o n i n t h e number o f biarmed  chromosomes has t a k e n p l a c e .  chromosomes i n t h e s m a l l s i z e centric  One p a i r o f b i a r m e d  c a t e g o r y has u n d e r g o n e a p e r i -  i n v e r s i o n f o r m i n g an a c r o c e n t r i c  T h i s k a r y o t y p e was e x p r e s s e d  chromosome  pair.  i n 74 p e r c e n t o f t h e s a m p l e .  I n t h e r e m a i n i n g 26 p e r c e n t t h e k a r y o t y p e , t y p i c a l  of the  P. m. g e o r g i e n s i s i n h a b i t i n g t h e o t h e r i s l a n d s , was  expressed.  The gonosomes a r e a l s o s i t e s within this  subspecies.  chromosome t y p e s .  Savary  o f chromosome  I s l a n d possesses  Three of the f i v e animals  polymorphism two Y  karyotyped  ISLAND  NUMBER OF CELLS COUNTED  NUMBER OF ANIMALS EXAMINED  BOWEN THORMANBY TEXADA SAVARY  14 44 54 19  4 12 13 5  DOYLE HURST BALACLAVA NIGEI VANSITTART HOPE  5 4 5 81 68 85  3 1 2 18 13 20  COX LANZ TRIANGLE  13 20 44  6 5 9  464  112  Total Table  15  ISLAND  BOWEN THORMANBY TEXADA SAVARY DOYLE HURST BALACLAVA NIGEI VANSITTART HOPE COX LANZ TRIANGLE .  ISLAND GROUP  Georgia group  Straits  Gordon and G o l e t a s group  Scott group  C o l l e c t i o n . and s k a r y o t y p e st u d y .  Island  iple s i z e d a t a f o r t h e  CHROMOSOME . SOMATIC CHROMOSOMES F.N. NUMBER BIARMED ACROCENTRIC A* B* 48 48 48 48 48 48 48 48 48 48 48 48 48  28 28 26 28 36 32 30 32 24 28 36 36 36  0  0 0 0 4 0 0 6-8 4 0 4 4 4  18 18 20 18 6 14 16 8-6 18 18 6 6 6  74 74 72 74 86 78 76 84-86 74 74 86 86 86  _ P.m.austerus 48 28 18 74 — P . o r e a s ( c o a s t ) 48 28 18 74 P.sitkensis 48 40 6 86 A* s u b m e t a c e n t r i c s B* s u b t e l o c e n t r i c s , s u b j e c t t o misidentification. Table 16 K a r y o t y p e c h a r a c t e r i s t i c s f o r t h e Peromyscus o f t h e B.C. r e g i o n .  -  P.  Scott Is.  sitkensis  (20)  (20)  See  inset P.  map oreas  (14)  . m. g e o r g i e n s i s 7 j 3 and 14) P. P.  oreas  (13)  P.m.  P. m. r u b i d u s "03,14,15)  oreas  austerus  (18 and  (14)  P. m. gambeli I 13,14,15,16,17)  Hope (14)  Vans i t t a r t (14) Nigei (20) k 5 A K Balaclava (15) W ). Hurss t 6  Fig.  15  20)  Somatic m e t a c e n t r i c  ' Doyle °.. (20) )  chromosome numbers f o r some  w e s t c o a s t p o p u l a t i o n s of P e r o m y s c u s .  e x h i b i t e d a Y chromosome t y p i c a l o f t h e Texada and  Thormamby  p o p u l a t i o n s , a s m a l l a c r o c e n t r i c , w h i l e t h e o t h e r two  speci-  mens were i d e n t i c a l t o the Bowen t y p e i n h a v i n g a s m a l l t e l o c e n t r i c Y chromosome. h a r d l y be mainland  This polymorphic  c o n s i d e r e d t o be due populations.  P. m.  r e g i o n of B r i t i s h Columbia, a large subtelocentric  condition  sub-  can  t o t h e i n f l u e n c e of a d j a c e n t  a u s t e r u s of the S e c h e l t P e n i n s u l a 50 m i l e s n o r t h o f V a n c o u v e r ,  chromosome (Thomas, 1 9 7 0 ) .  The  has  P.  o r e a s p o p u l a t i o n s a r e a l s o t o be d i s c o u n t e d f o r b r i n g i n g about t h i s polymorphism  f o r t h e y , t o o , have a Y  t h a t i s u n l i k e e i t h e r o f t h e two tion.  found  chromosome  i n the Savary  I t i s p o s s i b l e t h a t the complex c o a s t l i n e of  Columbia, h a r b o r s numerous k a r y o t y p i c l i n e s  populaBritish  of Peromyscus  not  y e t r e c o g n i z e d as d i f f e r e n t and most l i k e l y n o t even s e e n a taxonomist.  The  i s l a n d p o p u l a t i o n s c o u l d have b e e n t h e  r e s u l t of f o u n d i n g c o l o n i e s t h a t a r e k a r y o l o g i c a l l y y e t f r o m t h e same g e n e r a l a r e a . t h a t S a v a r y I s l a n d was remaining islands other but not  varied  I t i s p o s s i b l e , f o r example,  i n v a d e d by two  s u c h r a c e s and  i n t h e g r o u p were i n v a d e d by one  the  or the  both.  A f u r t h e r c o m p l i c a t i o n of the k a r y o t y p i c a n a l y s i s when I f o u n d  by  t h a t i n some a n i m a l s v a r i a t i o n i n t h e  o c c u r r e d w i t h i n t h e same t i s s u e c e l l c e l l u l a r mosaicism  The  karyotype  occurrence  i s f u r t h e r c o m p l i c a t e d by a n o t i c e a b l e  f r e q u e n c y of heteromorphic s e r i o u s problem  line.  arose  chromosome p a i r i n g .  The  most  i n the a n a l y s i s arose from the d i s c o v e r y  of  that s i g n i f i c a n t v a r i a t i o n i n the biarmed/acrocentric r a t i o s e x i s t i n e v e r y sample.  I n t e r c e l l u l a r and i n d i v i d u a l k a r y o -  t y p i c v a r i a t i o n has b e e n n o t e d i n o t h e r P e r o m y s c u s p o p u l a tions is  (Ohno e t a l , 1966;  A r a k a k i and S p a r k e s , 1966) y e t i t  o n l y i n t h e w o r k o f K r e i z i n g e r and Shaw (1970) t h a t a n y  i n d i c a t i o n of t h e e x t e n t of t h e polymorphisms i s brought t o l i g h t . rufinus with this  i n Peromyscus  K r e i z i n g e r f o u n d one s p e c i m e n  three d i f f e r e n t karyotypes.  o f P. m.  The v a r i a t i o n i n  c a s e was due t o t h e m i s - m a t c h i n g o f one o r two chromosome  pairs. The v a r i a t i o n n o t e d i n my s t u d y i s o f a much g r e a t e r magnitude  and n e c e s s i t a t e s t h e f o l l o w i n g s t a t i s t i c a l  i n order to a l l e v i a t e the problem.  A l l r a t i o s f o r these  i s l a n d P e r o m y s c u s a r e b a s e d upon t h e modal a v e r a g e biarmed sampled.  to acrocentric  i n the k a r y o t y p e a r i s e from t h e  o f one p a r t i c u l a r g r o u p  chromosomes ( g r o u p B, t a b l e 32) a p p e a r as a c r o c e n t r i c t y p e s , d e p e n d i n g DNA r e p l i c a t i o n  r a t i o of  chromosomes f o u n d i n t h e p o p u l a t i o n  The d i s c r e p a n c i e s  d u a l phenotype  maneuver  o f chromosomes.  These  as b o t h b i a r m e d and  upon t h e t i m e a t w h i c h t h e  c y c l e was a r r e s t e d .  The s u b t e l o c e n t r i c  chromosomes a r e a p p a r e n t l y t i m e s e n s i t i v e t o t h e m i t o t i c a r r e s t i n g a g e n t and t h e " r a b b i t e a r s " ( t a b l e 14) u n d e r g o drastic  contractions.  Consequently, o p t i c a l a n a l y s i s of  g r o s s chromosome m o r p h o l o g y  i s r e n d e r e d d i f f i c u l t and r e s u l t s  i n t h e mis i d e n t i f i c a t i o n o f many o f t h e s e extreme c e n t r i c p a i r s o f chromosomes.  subtelo-  N i g e i I s l a n d has 8 p a i r s o f  these  s u b t e l o c e n t r i c chromosomes and  the morphotype r a t i o s occur w i t h equal The fig.  i s indecisive,  modal a v e r a g e  the r a t i o s  s u b t e l o c e n t r i c chromosomes ( g r o u p B,  study area.  38-8  t o the  However, P.  i n h a b i t a t and  and  t a b l e 32  40:6  and  The  sitkensis,  s i z e to those  the S c o t t and  i s t h e same as t h e Goletas  believe that this  islands  of the S c o t t  closely  islands, (Hsu,  sitkensis is  s i m i l a r morph s t u d i e d upon  ( t a b l e 16 and  s i m i l a r i t y may  this  a l a r g e mouse v e r y  b i a r m e d / a c r o c e n t r i c r a t i o f o r P.  this  40-6  and  l a r g e mouse p h e n o t y p e o f  a p p a r e n t l y does n o t h a v e t h i s g r o u p o f B chromosomes 1970).  of  frequency.  29), are s p e c i f i c  allied  the  appendix 5).  have taxonomic  I  significance  worthy of f u r t h e r i n v e s t i g a t i o n .  I t seems t h a t a  karyotype/  m o r p h o t y p e r e l a t i o n s h i p c o u l d be  i n f e r r e d f o r these  northern  P e r o m y s c u s s i n c e t h e h i g h P.N. m o r p h o t y p e and v i c e v e r s a . the c o n t i n e n t a l forms.  values  correspond  This i s not  the case  chromosomes (Thomas, 1 9 6 8 ) .  near r e v e r s a l  of the  large  f o r some o f  Peromyscus c a l i f o r n i c u s ,  l a r g e s t p h e n o t y p e s i n t h e genus P e r o m y s c u s , has acrocentric  to the  one  of  19 p a i r s  This represents  chromosome m o r p h o t y p e s f o u n d  the of a  i n the  B r i t i s h Columbia p o p u l a t i o n s . C e l l u l a r mosaicism, some p a i r i n g ,  i n t h e f o r m of h e t e r o m o r p h i c  i n c r e a s e s w i t h the i n c r e a s e i n F.N.  S c o t t I s l a n d P e r o m y s c u s [F.N.-86 ( t a b l e 16 and had  a frequency  chromo-  value.  appendix  o f mismatched* chromosomes a p p r o a c h i n g  c e n t of t h e s a m p l e .  The  low F.N.  g r o u p s t e n d t o be  12  more  The 5)] per  constantly  stable karyologically.  chromosome i s t h e more u n s t a b l e going frequent  inversions?  would i n f e r t h a t the h i g h most r e c e n t island  Can i t be t h a t t h e b i a r m e d  c o n f i g u r a t i o n and i s u n d e r -  I f this  i s the case, then i t  f u n d a m e n t a l number g r o u p i s t h e  form o f Peromyscus t o have e v o l v e d  complex.  This  within  this  i s not consistent with the g l a c i a l  r e f u g i a t h e o r i e s p u t f o r t h by McCabe and Cowan ( 1 9 4 5 ) and Foster  (1965).  I n b o t h o f t h e s e s t u d i e s P. s i t k e n s i s , a  member o f t h e h i g h F.N. g r o u p , i s a r e f u g i a l r e l i c t survived  t h e V a s h o n g l a c i a t i o n i n t h e n u n a t a k s o f t h e Queen  Charlotte The  Islands. k a r y o l o g i c a l e v i d e n c e s u g g e s t s t h a t T r i a n g l e , Cox,  L a n z , N i g e i and D o y l e P e r o m y s c u s s h o u l d cohort, studied.  be c o n s i d e r e d  one  d i s t i n c t f r o m t h e r e m a i n i n g i s l a n d s t h a t I have The r e m a i n i n g i s l a n d s a r e g e o g r a p h i c a l l y  Hope, V a n s i t t a r t , B a l a c l a v a miles  having  north  widespread.  and H u r s t a r e a p p r o x i m a t e l y 170  of t h e Georgia S t r a i t  i s l a n d group, y e t Pero-  myscus f r o m b o t h s e t s o f i s l a n d s a r e c h a r a c t e r i z e d b y a l o w f u n d a m e n t a l number ( t a b l e 1 6 ) . Balaclava  and H u r s t I s l a n d p o p u l a t i o n s  of Peromyscus  have u n d e r g o n e some e v o l u t i o n i n k a r y o t y p e ;  p e r h a p s due t o  genetic  This  immigration  o f t h e h i g h F.N. s t o c k .  i s not substantiated  point  by t h e b r e e d i n g s t u d i e s , f o r i t was  f o u n d t h a t t h e two k a r y o l o g i c a l s t o c k s do n o t i n t e r b r e e d  latter  i n the laboratory.  (F.N.=86 and F.N.=74) C o n s e q u e n t l y any i n f l u x  o f t h e a l t e r n a t i v e gene p o o l w o u l d n o t i n f l u e n c e t h e r e s i d e n t  population.  This  leaves  me  with  o n l y one  m u t a t i o n o f t h e a n c e s t r a l f o r m s has and  H u r s t I s l a n d s , and  taken place  c l o s e l y r e l a t e d t o and  i n a cohort  be  study  cohorts  g e n e t i c a l l y , s i n c e the m e i o t i c  insurmountable, i n d i c a t i n g that  s p e c i a t i o n has  therefore taken place.  w o u l d be  populations  into species  Robertsonian  (1968) i n h i s  rapidly.  therefore of a v e r y  differ-  groups based As  previously  t o proceed under a  t h a n a n o n - R o b e r t s o n i a n system of  e v o l u t i o n , and indicative  easier f o r meiosis  mutually  stasipatric  Nadler  s o l e l y on v a r i a t i o n of chromosome numbers. noted i t i s probably  be  problems would  of t h e k a r y o t y p e s o f S p e r m o p h i l u s t o w n s e n d i has  entiated allopatric  is  Texada,  maniculatus.  I t w o u l d a p p e a r t h a t t h e s e two  seem t o be  sitkensis,  Bowen I s l a n d P e r o m y s c u s s h o u l d  grouped w i t h Peromyscus  Cox,  considered  w i t h Peromyscus  w h e r e a s t h e Hope, V a n s i t t a r t , B a l a c l a v a , H u r s t ,  incompatible  species.  the T r i a n g l e ,  Doyle I s l a n d Peromyscus should  S a v a r y , Thormanby and  upon B a l a c l a v a  t h a t they are both i n c i p i e n t  R e l y i n g upon t h e s e k a r y o l o g i c a l d a t a L a n z , N i g e i and  alternative, that  the v a r i a t i o n i n the  chromosome island  samples  p l a s t i c animal group t h a t i s e v o l v i n g  CONCLUSIONS:  This  A SYSTEMATIC REVIEW OF SOME INSULAR PEROMYSCUS  s t u d y has shown t h a t t h e r e  of Peromyscus  i n h a b i t i n g the outer  Columbia coast. is  a r e two d i s t i n c t  i s l a n d s along  species  the B r i t i s h  The p r e f e r r e d h a b i t a t f o r i n s u l a r P e r o m y s c u s  t h e d r y , s t o n y b e a c h e s and t h e a d j a c e n t f o r e s t edge.  mice a r e l i t t o r a l  s c a v e n g e r s , s u b s i s t i n g upon, o r a t l e a s t  r e l y i n g h e a v i l y upon, b e a c h amphipods D u r i n g t h e summer months t h e r e of v e g e t a b l e m a t e r i a l . characteristic subspecies  These  No  i s very  and i n s e c t  larvae.  l i t t l e dietary  intake  i s l a n d - s p e c i f i c h a b i t a t or d i e t a r y  c a n be f o u n d t o c o r r e l a t e w i t h  the s p e c i e s  or  designations.  E c o l o g i c a l and e t h o l o g i c a l p e c u l i a r i t i e s draw a t t e n t i o n to the T r i a n g l e I s l a n d p o p u l a t i o n  of Peromyscus.  s i z e , a m i c a b i l i t y , and s e m i - f o s s o r i a l way for  coastal maniculatus.  of l i f e  The  are a t y p i c a l  Taxonomically the s e m i - f o s s o r i a l  mode c o u l d be i m p o r t a n t b u t i n t e r - i s l a n d b r e e d i n g have shown t h a t t h i s dominant t r a i t  and I am o f t h e o p i n i o n  that i t i s the l a c k  t h i s way  of l i f e  a l e a r n i n g b e h a v i o u r and n o t a t r u e e v o l u t i o n a r y morphological  island populations. and o t h e r  studies  c h a r a c t e r i s t i c i s n o t i n h e r i t e d as a  of a l t e r n a t i v e h a b i t a t t h a t b r i n g s  Extensive  large  adaptation.  v a r i a t i o n e x i s t s b e t w e e n some  Discriminant  morphological  and i s  a n a l y s i s of the c r a n i a l  d a t a d i f f e r e n t i a t e s two n e a r l y  exclu-  s i v e groups w i t h i n t h e Peromyscus samples. B r e e d i n g s t u d i e s and mate s e l e c t i o n c h o i c e s  substantiate  ISLAND Triangle  CURRENT NOMENCLATURE Peromyscus m a n i c u l a t u s triangularis  PROPOSED REVISION Peromyscus s i t k e n s i s isolatus  Lanz  carli  isolatus  Cox  carli  isolatus  Nigei  isolatus  isolatus  Doyle  doylei  doylei  Hurst  saxamans  saxamans  Hope  Peromyscus balaclava  Vans i t t a r t  N.A.  georgiensis*  Balaclava  balaclava  balaclava  Texada  georgiensis  georgiensis  Thormanby  georgiensis  georgiensis  Savary  georgiensis  georgiensis  Bowen  georgiensis  georgiensis  * s u b j e c t t o comparison s u b s p e c i es  Table  17  Proposed island  maniculatus  Peromyscus m a n i c u l a t u s eeorffiensis*  w i t h P. m. a u s t e r u s and P. m.  revision  Peromyscus.  interdictus  o f t h e s y s t e m a t i c s o f some  the  existence  of the  two  type groups, cohorts ence f o r t h e hybrid  A and  Morphological exception  These two  all  r e s u l t o f l i k e morph are  are  glandular  The  Balaclava  length.  The  i n t o any  o r e a s o r P.  The  between  along i t s  Doyle sample i s unique i n h a v i n g  any  biological  The  phallic variation  d i f f e r e n t i a t i o n of the groups.  island  4  two  group theory.  a d i s t i n c t k a r y o t y p e u n l i k e t h a t o f any maniculatus  ( f i g . 15),  but  i t i s the  maniculatus.  The coastal  same as  s i m i l a r s p e c i e s , P.  s m a l l morph i s k a r y o t y p i c a l l y a l l i e d  l a n d c o a s t a l P.  cohort  populations.  of u n i f o r m t h i c k n e s s  t h a t p o s s e s s e d by t h e m o r p h o l o g i c a l l y kensis.  The  distinct.  Karyotaxonomy f u r t h e r enhances the  P.  prefer-  Peromyscus are a t y p i c a l  c l e f t medially.  samples p r e c l u d e s  l a r g e morph has  Doyle  p h a l l i c v a r i a t i o n e x i s t s w i t h i n and  v e n t r a l lappets  populations  pheno-  crosses.  s t r o n g w i t h i n each  i n both cases  unique i n having a p h a l l u s  w i t h i n the  small  show d i s t i n c t  o f t h e V a n s i t t a r t and  island populations.  total  l a r g e and  ( t a b l e 17),  affinities  populations  Extensive  and  B  The  c o m p a n i o n s h i p o f a s i m i l a r morph p a r t n e r .  l i t t e r s were t h e  w i t h the  cohorts.  to the  Cowan ( 1 9 3 5 ) c l a s s i f i e d  sitmain-  the  l a r g e m o r p h o t y p e as P e r o m y s c u s s i t k e n s i s i s o l a t u s and  subse-  quently  and  changed i t t o P.  Cowan, 1 9 4 5 ) .  I t i s my  i s more r e p r e s e n t a t i v e taxonomic r e v i s i o n of  m a n i c u l a t u s i s o l a t u s (McCabe o p i n i o n t h a t the  of the  first  classification  l a r g e morph g r o u p and  that  a  these i s l a n d Peromyscus i s n e c e s s a r y .  Peromyscus s i t k e n s i s i s p h e n o t y p i c a l l y  s i m i l a r to the  large  morph g r o u p . type  Peromyscus s i t k e n s i s  t o t h a t f o u n d i n t h e l a r g e morph c o h o r t .  P. s i t k e n s i s groups. stand  i s also identical  Geographically,  i s f o u n d upon i s l a n d s t o t h e n o r t h  Landslides  o r i g i n a t i n g i n the northern  an e x c e l l e n t chance o f d e p o s i t i n g f o u n d i n g  upon t h e s t u d y  islands.  C o a s t a l P. o r e a s ,  i n karyo-  of the study r e g i o n would colonies  although  pheno-  typically  s i m i l a r t o t h e l a r g e morph g r o u p , i s k a r y o l o g i c a l l y  identical  t o t h e c o a s t a l P. m a n i c u l a t u s .  demonstrates the d i s p e r s a l patterns along American  The map ( f i g . 9) the western  North  continent.  I p r o p o s e t h a t t h e P e r o m y s c u s f o u n d upon T r i a n g l e , L a n z , Cox,  N i g e i , D o y l e , and H u r s t  species sittart,  P. s i t k e n s i s ,  Islands should  be c o n s i d e r e d t h e  and t h a t t h e P e r o m y s c u s o f Hope, V a n -  B a l a c l a v a , S a v a r y , T e x a d a , Thormanby, a n d Bowen  r e m a i n as P.  maniculatus.  Variation  e x i s t s w i t h i n e a c h o f t h e two c o h o r t s .  The  degree of v a r i a t i o n w i t h i n t h e c o n t i n e n t a l p o p u l a t i o n s of P. m. r u b i d u s  and P. m. g a m b e l i ,  together  t h a t e x i s t s between samples t a k e n makes i t m a n d a t o r y t h a t s u b s p e c i e s  with the v a r i a t i o n  one o r two d e c a d e s designations  enough t o encompass s u c h n a t u r a l l y o c c u r r i n g  apart,  be made b r o a d  morphological  fluctuations. Subspecific designations  a r e somewhat a r b i t r a r y and a r e  made w i t h t h e i n t e n t i o n o f d i s t i n g u i s h i n g m o r p h o l o g i c a l l y s i m i l a r groups of animals definable.  The i n s u l a r  that are usually geographically  s i t u a t i o n adds a n o t h e r d i m e n s i o n  to the problem of s u b s p e c i f i c nomenclature f o r i n t h i s all  populations  a r e a l l o p a t r i c and t h e c h a n c e s o f i n t e r b r e e d i n g  are v i r t u a l l y n o n - e x i s t e n t . subspecific t i t l e s I n my s t u d y less  Is i t practical  f o r insular  t h e r e f o r e t o have  populations?  the i n t e r - i s l a n d v a r i a t i o n  than t h e v a r i a t i o n  especially  i s i n some  found w i t h i n the i s l a n d  when t h e c h a n g e s o v e r a t w e n t y - y e a r  considered.  These p o p u l a t i o n s  subspecific  i n most c a s e s .  period are  s h o u l d be c o n s i d e r e d  Distinctive  cases  sample,  as c o n -  morphological  s u c h as i n t h e D o y l e I s l a n d P e r o m y s c u s , s h o u l d be in the f i n a l  case  traits,  considered  evaluation.  I h a v e i n d i c a t e d i n t a b l e 17 t h e s u b s p e c i f i c t a x a are  suggested  populations  by my s t u d y .  However t h e a d j a c e n t  ( P . m. a u s t e r u s )  mainland  and t h e n o r t h e r n V a n c o u v e r I s l a n d  P e r o m y s c u s ( P . m. i n t e r d i c t u s ) , the  that  have n o t b e e n c o n s i d e r e d i n  e v a l u a t i o n and t h e r e f o r e t h e s u b s p e c i f i c d e s i g n a t i o n s a r e  t o be c o n s i d e r e d ized  as o n l y r e l a t i v e  i n t h i s study.  characteristics  t o the i s l a n d  sample  util-  V a n s i t t a r t I s l a n d Peromyscus a r e i n a l l  very s i m i l a r  f o u n d 170 m i l e s t o t h e s o u t h .  to the georgiensis Tidal  subspecies  f l o w c o u l d have a c c o u n t e d  f o r t h i s d i s t r i b u t i o n and i n o r d e r t o c o n v e y t h e s i m i l a r i t i e s existing  among t h e i s l a n d  specific t i t l e  forms I c o n s i d e r t h a t t h e con-sub-  i s justified.  Doyle I s l a n d Peromyscus a r e m o r p h o l o g i c a l l y d i s t i n c t i v e (fig.  8).  lambdoidal  This i s l a n d  form i s c h a r a c t e r i z e d by a l a r g e  r i d g e , w h i c h i s u n i q u e among t h e i s l a n d  Peromyscus,  and  cone shaped v e n t r a l l a p p e t s  these characters subspecific  on t h e  e s t a b l i s h d o y l e i as  G u i g e t , 1965).  o t h e r member o f t h e the k a r y o t y p i c  are  cohort A  hybrids.  In t h i s  has  place.  The  f i g . 8) I  t o be four  upon t h e m e i o t i c  instance  I believe  v a r i a t i o n s a r e m i n o r and  are  n a t u r a l v a r i a t i o n found w i t h i n the  The  Balaclava  doylei the  s i g n i f i c a n t enough chromosomal p r o c e s s e s of  changes any  speciation  one  form.  encompassed neighboring  The  within populations.  I s l a n d v a r i a t i o n i s , however, s i g n i f i c a n t a t  level.  The  k a r y o t y p e v a r i a t i o n ( t a b l e 16)  thick d i s t i n c t i v e phallus among t h i s  Pero-  consider  stastipatric  the  subspecific  Island  than to  s m a l l morph c o h o r t i s m o r p h o l o g i c a l l y  karyotypic  sound  s i m i l a r i t y to  ( t a b l e 17,  v a r i a t i o n ( t a b l e 16)  a severe i m p o s i t i o n  The  ( f i g . 8)  D e s p i t e the  i n t e r - i s l a n d crosses.  taken  a biologically  c l o s e r t o Cox  myscus i n m o r p h o l o g i c a l d i m e n s i o n s  to hinder  Together  taxon.  H u r s t I s l a n d Peromyscus are  (Cowan and  phallus.  give  t h i s group a unique  o t h e r w i s e mono-phenotype  the  and  the  character  cohort.  T e x a d a I s l a n d P e r o m y s c u s d i s p l a y an a t y p i c a l k a r y o t y p e i n t h a t t h e y have o n l y 26 m e t a c e n t r i c chromosomes, b u t other characters  measured are  e x h i b i t e d by  o t h e r P.  the  m.  still  ogical variation.  This  range  georgiensis.  I t appears t h a t a t h r e s h o l d e s t a b l i s h e d a t s u c h a l e v e l as  w e l l w i t h i n the  the  of s u b s p e c i a t i o n  to allow  requires  the  must  for natural  recognition  be  morphol-  of b r o a d  taxa.  I n the case  of the i n s u l a r Peromyscus i t appears t h a t  c r a n i a l morphology, s p e c i f i c a l l y  the l a m b d o i d a l  o v e r a l l p h a l l i c m o r p h o l o g y ( b u t n o t any f e a t u r e ) , and level.  The  c r a n i a l and  the k a r y o t y p e ,  one  r i d g e , the  specific  phallic  are s e n s i t i v e at the s u b s p e c i f i c  s p e c i e s s h o u l d be d e f i n e d by a n a l y s i s o f other morphological  measurements t a k e n  s a m p l e s o b t a i n e d o v e r many y e a r s , as v e i l of t h e b r e e d i n g p r e f e r e n c e s  gross  from  as by c o n s i d e r a t i o n  of the a n i m a l s  studied.  Differ-  e n t i a t i o n o f i n s u l a r p o p u l a t i o n s o f P e r o m y s c u s must be w i t h the awareness t h a t n a t u r a l a n i m a l d y n a m i c and  c o n s t a n t l y changing  g r a p h i c v e r n a c u l a r ( W i l s o n and  populations  entities.  The  Brown, 1953)  done  are  i d e a of g e o -  being applied  to i n s u l a r p o p u l a t i o n s i s not s c i e n t i f i c a l l y a p p r o p r i a t e . Such a s y s t e m o f i s l a n d p o p u l a t i o n n o m e n c l a t u r e i s o n l y "precise"  i n the d e s i g n a t i o n of the c o l l e c t i o n l o c a l i t y , i t  does n o t c o n v e y any b e t w e e n any study  i d e a of the e v o l u t i o n a r y r e l a t i o n s h i p  of t h e n e i g h b o r i n g r a c e s .  I have found  in this  t h a t the S c o t t I s l a n d Peromyscus are d i s t i n c t from  Georgia  S t r a i t P e r o m y s c u s , a t what I c o n s i d e r t o be  the  s p e c i e s l e v e l , but w i t h i n the S c o t t group the Peromyscus m o r p h o l o g i c a l l y and k a r y o t y p i c a l l y v e r y c l o s e . each of t h e S c o t t I s l a n d p o p u l a t i o n s as would convey n o t h i n g t o the taxonomist review be  To  separate and  o f t h e l i t e r a t u r e t o a s c e r t a i n any  the  types  are  designate  entities necessary  d e s c r i p t i o n would  a t e d i o u s t a s k r e m i n i s c e n t of t h e p r e - L i n n e a n The  the  o f i n f o r m a t i o n I have u t i l i z e d  era.  in this  study  l e a d me  to the  evaluation  f o l l o w i n g r e v i s i o n as b e i n g t h e most m e a n i n g f u l  of t h e P e r o m y s c u s on t h e s e i s l a n d s . be  i s h Columbia.  P e r o m y s c u s s i t k e n s i s o c c u r s on N i g e i , Cox,  and  i s l a n d s o f f the  species  P e r o m y s c u s can  Hurst, Triangle,  f o u n d upon t h e  Two  Lanz I s l a n d s .  f o u n d on Hope, V a n s i t t a r t , B a l a c l a v a , Islands, species  i s of the P.  species  and  possessing  no  s . doylei.  and  karyotypically distinctive;  the  s u b s p e c i f i c t a x o n P. £.  saxamans.  little  have p l a c e d  Island,  them as P.  t o m a i n l a n d P. Island.  s h o u l d be  m.  m.  a  P.  m.  and diff-  separate although  phenotypically deserving small  m o r p h o l o g i c a l and  georgiensis,  a u s t e r u s and  are  Among t h e  considered  sub-  to a major  thus they are  i c a l v a r i a t i o n exists that a l l populations, of B a l a c l a v a  Lanz,  considered  osteological peculiarities,  Strait  The  Hurst I s l a n d Peromyscus,  r  m a n i c u l a t u s , so  morph,  Georgia  Cox,  D o y l e I s l a n d P e r o m y s c u s , due  e r e n c e i n c r a n i a l m o r p h o l o g y , must be s u b s p e c i e s , P.  the  Brit-  Doyle,  smaller  Peromyscus m a n i c u l a t u s .  _s. i s o l a t u s i n h a b i t s T r i a n g l e ,  Nigei Islands.  g r o u p , P.  The  c o a s t of  of  of  morph  karyolog-  except f o r  those  con-subspecific.  subject  to  I  comparison  i n t e r d i c t u s from Vancouver  APPENDIX 1  MEASUREMENT DATA FOR INSULAR PEROMYSCUS  o  + l"  Tien  POSTPALATAL LENGTH  DIASTEMA WIDTH  PALATINE FORAMEN  SHELF OF BONY PALATE  cn  O  +1  00 00  O  + 1 +Th1 Th"  Tien  cn  cn  cn  + 1* oo 00  CO  cn  cn  O  + 1•  + r  00  ON 00  CO  CM  cn O •  +1  Th  Th NO  vo  vO  N0  NO NO  cn  cn O  cn  cn  CM  +*l  +"l  o -H  o  O  O  in  ON Ti-  o  + •1 o  cn  en  cn  m  o  O  +\  in • cn  o  o  +] in cn  o•  +N£>1 cn  in  m O  O  + 1 +i ON oo o  in  •  CM  o  +i ON •  CM  o  o  +"l  O  rH  O  rH  CM  rH  o  O  +i  +i  m  cn  cn  cn  cn  m  cn  cn  cn  TP  -f'l  +TI- i  +i cn  +  "l  O  rH  CM i—1  i—1  rH  rH  O  in O  o  cn  +1  +1  r-t  ON  + l"  ON Th  CM in CO  00 rH  co +\  <  +1 rH  ON rH OO  O  + 1*  Tt-  00 Th  CM  CM  o  in  co  <  O  PH  *  O  +1  rH  in  00 Th  cn  cn  CM  +i n1 cn •  rH  O  B o  cn •  cn •  O  o  O  +i rH Th  kO  cu n3  e 0)  OO  T3  rH  rH  rH  in O •  h-  VD  O  t— o  ca  +'l  +'l  CM  in  O  rH  • PH  • CO  •  oo  o  •  +m 1 CM  o o  CO  1—1 o  <  Th  in Th  CM  CM  PH 00 r H  00 rH  •  +1 CM  Th  I—i pq co rJ «! o > . O •  ca  rH  CM  |z;  tr!  rl  rH  +cn1  •  o  00  CM in  O  O  ON 00  o  CO PH  CO  < o  1p—1 o  PQ  &  CO rH  CO  o  PQ  o  -P Pi  cn 00  cn ON  rH  r<  t>  •  Th  o  •H  rH  *  + r  +i CM  -P  rH  in O  O  Pl  <4H  in  rH  <P  ca  in  ON  a> PH  -P  rH  +1  rl  rH  rH  +001  V  rH  rH  +1  0  O + '\  rH  +cn1  + 1 ON  rH  rH  •  ca  O  rH  o  Th  m  rH  O  O •H Pl  i n*  rH  O  rH  o  in cn «  +i cn  vi -P  cn  + 1 +NO i NO  •  6 ca tl  +Th1  in O • +1 in ON •  o  CD  P  o•  cn  ON in •  B ca w  O  +i cn  +i  0)  rH FH  rH vD  O  O  +i +i rH Th  in  o•  +"l  rH  in cn  co  CxJ  O  i—i  + 1 +001  +1  + •1  in  rH  O  CM  +"l  cn cn  in ON •  cn  O  +'\  • +1  00  cn vo  O  o  o  + 1• t-  cn  Th  T)-  cn  o•  t-  Ttcn  cn  Th  CM  +1  ON 00  •  +cn1  cn  CM  *  + r  o  o  o  O  o  Th  O  •  cn  +1  cn  cn  •  cn  CM O •  +1  cn  ON  cn m  Tf  t>  + l  cn  m 00  cn O •  +i +i  Th  +1  CM  O  + r  00 00  +i  O  +i Th  iOn  +i  O  + 1•  CM vO  O  t~  r-l  O-  o .o -H +00•i  m  CM  O  O  +"l  t~  SAMPLE  00  cn cn  +*!  o•  O  SAMPLE SIZE  f  cn  +!  rH  OCCIPITONASAL LENGTH  +  O  + l"  Th  O  BASILAR LENGTH  O  +Th1  cn O  CM  BREADTH OF SKULL  +1  cn  O  •  *  cn  00 •  INTERORBITAL CONSTRICTION  •  O  cn  in  LENGTH OF NASALS  CM  CM  10  CM  10  MAXILLARY T00THR0W  w  -p Pl  o  +1  CM vO CM  •  Jz; co  1p—1  <J} r H « P o PH P pq  CO  CO CO  CO rH  <J  rH  ca  e rH  Th Th  00  o  Pi  W  < pq  o  ca  •H  Pi  ca  u  Oi M  ca  rt d) f>  ca  P caI a> S  < rH CD rH  ca  EH  MAXILLARY TOOTHROW  SI  DIASTEMA WIDTH  + +  PALATINE FORAMEN  + +  +  + +  + +  +  PI  -p  •H  X  -p  CO  PI  CD  + +  +  +  +  PHI  +  +  -p  CD  e  Pi  •H  CD rl PI CO  SHELF OF BONY PALATE  + +  + +  + +  + +  d  CD  CD  ' Pi  e  U  bo •H  PI o  PH  LENGTH OF NASALS  + +  + +  + +  + +  + +  + +  +  ci B  +  PI  PI CD  o  •H  PI  -P P)  INTERORBITAL CONSTRICTION  r&  + +  •H  +  CD CD  ?H  t*  -P co  •H  -P  ^  BREADTH OF SKULL  + +  +  PI CD  CD 0 CO  O •rl  -P c6  •rH  U  BASILAR LENGTH  + +  + +  +  + +  + +  +  + +  +  +  W W > t> H W m  r-H  (A >  HH  .  O CD O  PI  ct? O  OCCIPITONASAL LENGTH  EH EH  + +  + +  + +  + +  + +  + EH EH + <tj <J  + +  •rl «H •H  P!  60 •rl W  EH EH  <j <J  SAMPLE S I Z E \  o  > o  >  in  in  CM  > i—i  > i—i  > rH  >  >  >  CM  CM  CM  in  >  >  PH PH CO T-M—11—I  J>  'rl  ^J-  pi ri  &  CD C3 Cb I—I I—I S W 02 cd + W)+ +  CM  EH  +1*  LPi o  o +i tvo  CO O• + I rH LfN.  O  • +1  vO  O• +1  O  o  CM o + f  vO vO  •  co  O•  + 1  rH in •  CM  o  + f  + r  vO  + l"  vO ON  CO • o• +1  o o•  CO o• +1  rH ON •  co  o  • +1  + 1•  00  00  vO  •  O•  vO  +1  in VO O +1  in ON  o 00  in  CM  CM  00 ON  VO m  CO .J  s  CM o  o  rH O  +1  +1*  + f  CO  co  co  in co  CO o*  co o  O  . O  vO  ON  ON  CO o +'l  CO o +i CO vO  rH  O  O  o  +i  +i  +}  •  •  •  •  CO o  CM o•  in  in  in  rH O  ro O  CM O  • +1  CO  • vO  o*  +1  rH  o  •  rH  o  «  +1  vO  CO  O  • +1 00  O• +1  rH  o  •  + f  CO« '  ro O  CO  • +1  vO  ON  o + f  oo  in  in O +1  in ON  • + 1  w  PH  CM o • rH  CM O  • +1  ON ro •  +1  • +1  <  O + r  +1  vO  CM CM  in  o  O vO  CM  CM in  +1  o  co O  co O  + f  + f  + f  O  O  +\  +l"  o o  00  ON  •  •  CM o + f *  co O  O  + r  in vO  ON « rH t-  5H  + r  CD  + f  •  PH  CD  •  'A o  o +i  •H  -P  d  in  •H  t> CD  rrH  oo  00  o  o  + r  o  in ON  ON  ON  o  H  cti  PI cd -p  CM vO O  o  + f  + r  co in  in m  CM  CM ON iz; o o rH O CM CM< • rH & W O Pi O Jz; C3 o o Pi o. iz; PH H o CO rH M <Ji • o < Pi < iz; o  a  CM ON rH  o  02  o  CD  • +1  + r  co VO  CM  CM  <  iz; iz; M Pi  o o  PH  l—l iZ; (J W -4 PH  S o  O  CM O  PH  1—i  o  iz;  iz; i—i  PH  iz; rH M P Pi S3 -< P3  l—i  •J  < O  o  s  in  CQ  -P PI  CD  e 0  rH  Pi m «3  CD B ce Pi  o  a rH CD  o  O =H  +1  CD ON  fH  •H  O  in  -P  PI  on  O 1—1 I—1  m pi t> w !> S o  O  CO  CM ON  •H  CM  in  CM  rH Pi O  •  co O  + f  pi -p d  ON  CO  + l"  m  ON  co  +1  U  Pi  co  o  Pi  +1  + 1+vO1  in  43  VO co  co  vO O  • CM VO  o o  CNJ o  O  • + 1  •H  +1  in O  o  + r  CM o + f  tco  vO  • rH  00  O  •  rH  \o  o  co  •  Pi ^  •  in  CO  cr3  m  in  CO  E  O  rH in  in  •  o•  PH  m vO  + 1  CM  vO ro  in vO  «  +1  o  • +1  «  O  ON  rH VO  in ro  O  o  rH  • +1  •  ro  •  •  «  •  ON xr  •  + 1 rH ON  +1  CO  ON  »  +1  o  +i o  o  + f  ON  +1  CM  •  +1o «  CM  CM  +1  CM O•  O•  CM o  «  O  co  CM  •  CO  •  CM o + f  co co  •  rH  + r  > <  •  rH  o  in  in O  +o  rH  + 1 00  co ON  vO > vO  CO o  O  • +1  GE  SAMPLE  +1•  CO  oo  SAMPLE S I Z E  o  CO  O  OCCIPITONASAL LENGTH  ON •  co  •  BASILAR LENGTH  *  + 1+ 1 CM rH  • rH  BREADTH OF SKULL  hi  CO o•  O  in  INTERORBITAL CONSTRICTION  +1  -si-  CO  LENGTH OF NASALS  +1  + l'  CO  CM  SHELF OF BONY PALATE  rH O  CO  •  PALATINE FORAMEN  rH O  CO  O  DIASTEMA WIDTH  rH O  LT  o  OR  POSTPALATAL LENGTH  CM  O  T T  MAXILLARY TOOTHROW  !>  a  « PI 0j  CD  CO CD  rH  MAXILLARY TOOTHROW  + pl  POSTPALATAL LENGTH  +  + +  + +  + +  + +  + +  •H  + +  u El  DIASTEMA WIDTH  + +  + +  + +  +  + +  + +  +  + +  Pnl  -p x  Pl •H  VI -P Pl  -P Pl  PALATINE FORAMEN  + +  +  + +  + +  + +  + +  CO  •H  E  CD PS  CD rH  CO  0  ca CD  bD •H  SHELF OF BONY PALATE  «h PH ca  ca •H  E  Pl ca  Pl  u o  o  LENGTH OF NASALS  + +  + +  •H  +  Pl ca  -P &  E  •H  U -P V)  Pi CD  •H  INTERORBITAL CONSTRICTION  CD  >  -P  o  CD  42  0> W  Pl  o  •H  BREADTH OF SKULL  + +  + +  + +  -P  + +  ca SH  P W  BASILAR LENGTH  +  + +  +  +  + +  + +  + +  + +  > W  c3 !>  o o  Pi ca  o  OCCIPITONASAL LENGTH  + +  + +  + +  + +  + +  + +  + +  EH « '^EH  EH S  •H <+H •H  Pl  bD  •H CO  SAMPLE S I Z E Tt"  SAMPLE  *  O f C N J r n T j - t r N r n T h i P v T f i n t n  >  >  >  >  >  >  > CM  >  .  CM  CM  cn  g M CO  cn  Tt-  i  i M  +  ++  CD rH  ca EH  1  vfj en cn  o  +i  cn  •  o -H  00 vO •  CM  O  O  • + |  • +|  CM in  O in  CM o  CM o  +i r--  +i t~  •  •  •  cn  cn  •  •  m O  Tt  O  +i  +1  O  in O  rH • rH  • rH  O  + { t—  en  •  O  i—i  •  • +1 rH  • +1  •  •  in  +  CM  •  +1  in  cn  cn  •  rfen O -O  +i  +i in O  -si-  O  • rH  • rH  cn •  ON  O  •  +1  cn  {  00  en  • +1  CM  •  O  • +1  in  cn in  •  •  cn  •  m O  ON  . O  • rH rH  O ,  +1 CO  cn  •  t~ O  •  +1  ^  •  •  +1  O en  •  +i CM  en O  * +1 TT  +1  +1  +1  +1  O  rH V£>  CM  CM  CM  76  22  •  •  CM  *  t— CM o "• - •  +1  t—  MD *  CM  o o  •  O  •  O CD  cn  O  •  -P  +1 h-  O  o  fl o  •H  O  O  cn  m  rH * rH rH  m  m CM . H CM o. o +1* +f  M <H o  O  t-—  o  SI  en  in  o  CD •H W rl O CD W  oo  « rH  o  fl  •  rH • rH  O  w  TT  +1  en O  NO  TEXADA  CM  O  •  \D •  CM  r-i  +1  +1* Tt"  't-  •  O  + 1  +1* 00 vO •  t—  •  O  •  O  -n  rH  t— C M o  o +i  •  CM o +i CM  +i t-  •  in  o  •  cn  m  ON  +i t-  • •  t~-  o  +i  HON  +i t~  CM  NO  +1  o  CM o  • rH  . rH  cn  o  CM o  { + \D  O  cn  •  CM o  { + 00  oo  in cn  •  o  O  •  O  • +1  cn  in  rH  O  en  rH  •  +1  CM  O  rH • rH  •  SAMPLE  t-  CM  r— C M o  SAMPLE S I Z E  CM  O  O  CM  o  OCCIPITONASAL LENGTH  •  •  CM o +i  CM • rH  BASILAR LENGTH  ON  vO  ON  THORMANBY 19  BREADTH OF SKULL  rH • + 1  •  CM o +i  o  r-i r-t  r-t  INTERORBITAL CONSTRICTION  +i CM  cn  •  LENGTH OF NASALS  +i ON  r-l  •  +i ON  t—  cn  cn  o  O  NO  ' TJ-  cn  rH  o +\  rH  O  cn  -rt  CM  SHELF OF BONY PALATE  •  en  HON  •  PALATINE FORAMEN  •  o  •  DIASTEMA WIDTH  • +1  cn  •  POSTPALATAL LENGTH  rH  • +1 \0  O  oo  o  •  si) CM o  +1 rH  t>-  *  CM  TEXADA 30 INTERIOR  +  rH  • +1 vO  O  20  rH  •  TEXADA BEACH  rH  O  SAVARY  MAXILLARY TOOTHROW  o  •H  -P  ee  •rrl  > CD  m  -P fl CD  Ti  u  a fl  ca -P  • ui CD  a o  + 1  e  CD rl fl  in c$  • CD  S crj •H fl  ca rl  o  , CD  crj rl  in  CD  % fl ci CD  B  ca-  EH  Mean - s t a n d a r d d e v i a t i o n m i n i m u m a n d maximum  ISLAND  TOTAL LENGTH BODY LENGTH TAIL LENGTH  HIND FOOT LENGTH  N  DOYLE  210.4  + 14.9 109.6 ± 7.8 100 .7 ± 8 .0 177-223 95-118 82-110  22 .7 ± .7 22-24  9  HURST  205. 8 + 25.2 101.7 ± 1 4 . 5 102.5 i l 2 . 5 165-228 73-113 84-118  23 .2 ±1 .72  6  20-25 21 .5 ± .7 21-23  10  313.4 + 10. 2 111.7 + 6.3 101.7 + 6 2 194-234 87-110 101-125  24 .6 ± .9 23-26  19  95.8 + 7 1 85-110  21 .5 ± .9 21-23  15  95.2 + 5.4 79-104  21 .5 ± .9 20-24  34  COX  203. 6 + 23. 9 107.6 ± 9.1 101.0 + 7 2 125-234 88-112 90-125  24 2 £ .2 22-26  20  LANZ  212. 5 + 196-239  25 .2 ± .9 23.*5-27  29  TRIANGLE  223. 5 + 13.8 115.7 +10.0 109.9 +14 7 187-250 89-130 93-196  26 0 ± .9 24.*5-28  47  BOWEN  177. 9 + 166-200  9.3  81.9 ± 6.4 77-98  96.0 + 4 7 89-102  21 6 ± .6 21-23  10  °. 1 + 1 53-194  9.4  87.0 ± 4.3 78-96  93 2 ± 6 8 72-1 00*  20 9 ± .9 19*5-23  26  TEXADA  183. 4 + 162-201  8.5  85.1 ± 6.2 68-93  98.0 + 4.1 92-108  21 .8 ± .7 20.5-23 .0  37  SAVARY  177. 3 + 169-197  8. 0  81. 0 ± 3. 5 76-89  96.3 ± 5.4 90-110  21 0 ± .7 20-22  18  5.0  BALACLAVA  197. 1 ± 190-206  NIGEI  VANSITTART  197.0 + 186-210  HOPE  193. 3 + 11.9 165-225  THORMANBY  1 8  95.4 ± 7.4 101.7 i 4 .0 84-106 96-108  7.9 101.2 ± 2.5 96-106 98.2 ± 8.0 82-121  8. 5 111.0 ± 6.8 101.5 ± 5 .3 99-128 88-111  Mean ± s t a n d a r d d e v i a t i o n minimun and maximum  ISLAND  TOTAL LENGTH BODY LENGTH TAIL LENGTH  DOYLE  216.5' ± 205-235  8.6 111 .3 - 6. 5 105-2 3.4 102-123 101-112  23 .3 22-24  ±  .6  12  HURST  211-7 199-222  6-1  106.9 - 4. 9 104.8 - 3.8 100-114 95-110  23 -9 22-25  ±  -9  19  BALACLAVA  187.0 ± 165-204  7.6  90 .2% 81-101  5. 0  21.4 ± .8 20-23  37  NIGEI  212.6 ± 14.7 194-280  106. 9 99-118  5. 2 105.7 ± 1 4 . 4 90-178  24.9 24-26  40  VANSITTART  +  ±  •  ±  .  ±  •  HOPE  187-3 ± 171-204  7-0  88 . 7 - 6. 0 76-106  COX  214.3 ± 203-230  6.5  LANZ  96.4 - 5.1 83-106  +  •  98.5 - 4.0 91-116  N  HIND FOOT LENGTH  ±  . ±  .7  »  21.6 - .9 20-24  44  109. 7 - 5. 2 104.7 - 4.0 98-121 97-112  25.6 ± 1 . 1 21-27  23  205-0 ± 10.3 189-225  102. 2 ± 5. 8 102.8 ± 5.4 94-11392-115  25.6 ± .8 23-27  27  TRIANGLE  228.0 ± 211-239  9.4  118.1 ± 6. 3 109.9 ± 4.0 102-115 103-125  27.7 - .7 27-29  20  BOWEN  170.9 ± 154-193  7.5  78.0 + 3. 2 92.9 ± 7.1 73-85 80-116  21.6 ± .6  24  21-23  THORMANBY  170.3 ± 4.3 166-176  83. 0 ± 5. 0 87 .3 ± 3.5 79-90 83-91  22.4 ± .5 22-23  TEXADA  166.8 ± 156-184  6.7  69. 2 ± 4. 6 97.5 ± 4.5 61-78 90-108  22.3 ± .6 25 21-23.5  SAVARY  172.2 ± 160-190  7.3  82. 6 ± 4. 7 89.6 ± 3.3 75-92 85-98  22.3 ± .4 22-23  4  14  POSTPALATAL LENGTH  H/ O  •  + 1 VO  + 1 vO  + i  CO  CO  •  CO  + 1  • + i  *  in co  •  o  O  O  co  + f  + f  ON  ON  CM ON  rH  co  CM  + f 00 vO  rH  CM  + f  + 1  in  in  in  rH  iH  CO  CM  t-  •  •  + 1  + f  ON  •  •  in  o o  O  «  o  + 1 O  •  •  co  + 1  o  o  CM  + f rH ON  CO  •  CM  H/ O + 1  CO  DIASTEMA WIDTH  co  O  10  CO  + l vO  O  10  +1 00  «  rH  rH  10  TOOTIIROW  CM  o•  TO  MAXILLARY  o  •  00 00  O  + 1 CO vO  + f ON vO  CM  CM  tn  o  •  VO vO  + 1 o  r-  o  vO  co  CO  CO  CO  + f  + f  ON  ON  •  o  CM  •  o  in  CM  co  +1 ON vO  + f o  + 1 rH  CM  CM  o  •  O  00 vO ON  PI  in Nt ON  o  •  t>  VO  co ON  LENGTH OF NASALS  . O •  + f  + 1  in  in  CM  CM  CM  o•  +1  ro  o  o•  CO  + l"  + f LT\  O  rH  rH  co CO  VO  O rH  rH  ' rH O  CM  + 1  co  CO  co  rH  o  rH  t-  o  c-  co  o  o  + 1  + l"  o  o  + 1 rH rH  rH  rH  rH  •  co  CM  iH  O  o  + f t -  + l"  + 1 VO  ON  + 1 t-  + 1 ON  + f  + 1* 00  + 1*  CO  CO  CO  CO  rH  rH  CO  co  ro  CM  o• + i vO rH  O•  +1  o•  + 1 vO rH  + 1 vO rH  00 rH  rH  rH  rH  o +  f rH  O  O  O  + r  + l" CM  co  co  rH  rH  CO  in  O  +  + 1*  f  o  + i"  in  O• + 1  in co rH VO  o  + f  O  o o  CO ON  in rH  o o  CM  CM  CM  rH  CM  CM  CM  00 O  00  -<*  co  vO  vO O  o  + l"  ^  vO  CO i—1  o  + f vO VO CM ON  o + r  vO CM  00 rH  o  •  00  in  CM  + 1 vO vO CM  rH  o  + r  co  • + 1  o  vO  vO  CM  CM  CO  00  o o  rH  ro  CO  CO  ON  X w  < > <  *  *'  to  o  m *  PH  FH  *  O  +  PQ  +  CO •H  CO  0 PI  •H  &4 rl  0o bd El m  PHI  Pi o  •H  rl  o  -P  <H  + 1 •<* rH  + 1  •H r>  -p  rH 00  rH VO  O  + r  CM  O CM  0  o  CM  o  rl  crj  + f vO  Pi  O  -p  CM  co  o  o  + 1 rH VO  + 1  CM  CM  •  CO rH  •  ON  vO  O  in  <  o  X  K  EH  +  vO  in  00  VO CO O N ON ON  rH rH rH  £ •H  Pi •H •H  -p -p -p  pq  «  o o  o  o•  rH rH  <  0 o  rH  o  1--  a>  -P  o•  + 1  + l" vO  rH  in  + 1  + 1*  rH  CO  in  •  PI •H  •  CO  t-  o  r-T  O•  O + 1  + 1  o  o  CM  •  o  CM  SIZE  t>  o•  o  + 1 rH  o  CM  SAMPLE  +1  o  o  + f  O  O  OCCIPITONASAL LENGTH  CO  •  O  rH CO  BASILAR LENGTH  co  O  O  co  BREADTH OF SKULL  rH  o•  •  +1  o  CM  o o  rH  INTERORBITAL CONSTRICTION  t>  O  rH  in  O•  + 1  co  LfN  CM  CO  t-  O  •  +1  LT\ O  l> o  o  +1 rH LfN  AD  SHELF OF BONY PALATE  o  RM  PALATINE FORAMEN  CM  ed ed ed  rH  EH  +  o o o a> a>  0  o o o o o o  0  CO PI  0 E 0u pi  co  0  •H Pi  PI  o  + 1  0bo  O  cd rl  0)  '  00  >  Pi  0 E  EH  M A X I L L A R Y TOOTHROW  o P O S T P A L A T A L  + 1  o o  LENGTH  • •  o  + 1  WIDTH  O  •  + 1  cn D I A S T E M A  CM  ON  00  ON  •  cn  O  + 1  ON  cn  o  •  + 1  —cn—  O  + 1  rH  CM ON  O  •  ON VO  •  CM  o  CM  O  •  + 1  rH  CM + 1  •  O  + 1  cn  in  ON  •  CM  O  + 1 rH  cn  o  •  +1  TT  o +1  ON  •  ON  •  •  o  cn  Tt-  o  •  •  cn  + 1  vO ON  o  •  •  rH  O  •  TT  o  O  + 1  6  + 1  + 1  + 1  00  rH 00  Tt00  00  cn  CM  CM  rH  rH  00  +  cn  o  Pl  «  cn  o  rd  ON  vO  O  to  + 1  CS  rH CO •rl  Pl rl CD  43 -P  u  CM P A L A T I N E FORAMEN  o  •  + 1  CM  N0  cn  O  • -  O  NO  O  *  + 1  rH  CM  CM  +1  •  o  NO  in  rH  O  •  + 1  CM  m  CM  o  •  + 1  rH  in  O  •  + 1  rH v0  o  •  + 1  o  o  «  + 1  c  o  o P!  •  CD  + 1  43 -P  o  N0  NO  rH  CM  E CM S H E L F BONY  OF P A L A T E  LENGTH  OF  O  +1  cn  cn p  in O  Tt-  + 1  + 1  + 1  rH  O  INTER0RBITAL C O N S T R I C T I O N  O +\ rH •<*  CM  o BREADTH OF  + 1  S K U L L  CM CM  + 1  + 1  + 1  + 1  + 1  + 1  rH  + 1  CM  rH  + r  cn  TT  + 1  O o  cn o + 1  rH  CM CM  t-  CM  o  o  rH  rH  rH  rH  rH  O  O • +\ + 1 ON o cn  ON rH  rH rH  rH  rH  rH  ' rH  O  + 1  + f rH TT  •  o  TT  + 1  + 1  + 1  rH  rH  rH  AO  rH  TJCM  .O  cn "'cn" ' CM o O o  o  vO  ON  O*  cn o  + i  cn  + 1  rH Tt-  + ON 1  O  O  + 1 Tf  CM  CM  -P ce  t> CD  in O  VO  00  ON  CO  00  •  + ON 1 rH  CM  O  rH  + 1  + 1  vD t-  CO  •  + ON 1  ON rH  in  •  + in 1  CM  •  + TT1  O  •  + CM 1  O  CM  CM  CM  in  TT  00  O  O  + 1  + 1  rH  rH  O  + 1  in  •  +ON 1 rH  CM  o  o  + 1  + 1  00  VO  ON  vO  TINO  CM  CM  CM  CM  CM  CM  CM  ON  TT  in  oo VO  CM  CM  EH CO t>H  o  PH  rt  EH  <  < pq  o  CM CM ON  CO  C±3  1—i  rH CO  rg  t>  CM  in  PH  o  P3  o  CD  Pl O  I>  ON  + 1  CM  CM  CM  CD bo  cn  CM VO  to  o  c6 T3 Pl ca -p to  o +!  I—I EH  rl  o  m -P Pl CD  E  CD rl fl  CO  ca  CD E  ca  •H  P! ca rl  C_>  a rl  o o  CJ3  w'  rl  ON  EH  1—1  •  CM  + 1  ^  + 1  CM  + 1  in  O  CM  o  TT 00  cn  •  +0 10  o  < EH  r J  o  + 1  •H  CM  VO  •  P! o  •rl  o  VO  O  O CJ  + 1  t-  O  rH  rH  VD  o  i—I  cn  rH  O  O CD  •  rH  O  Pl o  •rl -P  , o o  rH  O  CO  rH  rH  o  44 ca -p  tCM  rH  in  S I Z E  + l  CD  rH  CM  SAMPLE  in o  o +\  cn O  Pl  in O  ON  rl  + 1  in o  rH  cn O  4  O  rH  ON  +  TT  rH  o  TT Tt-  in O  rH  + f  + 1  rH Tf  cn o  o  O  + 1  cn o  CM  O  + 1  cn  t-  rH  . rH  O C C I P I T O N A S A L  t-  + 1  co cn  +  rH  I>  LENGTH  o  o  o  o  rH  + 1 .  LENGTH  cn  cn  4  O  o  rH  O  B A S I L A R  vO  cn  rH rH  NASALS  + 1  cn  CM  CM o  rt bo pq Pl >H  rl  P3  EH  CD  ON  > ca  P I ca CD E  42 ca EH  Mean - s t a n d a r d d e v i a t i o n minimum and maximum  ISLAND  TOTAL LENGTH BODY LENGTH TAIL LENGTH  DOYLE  6.1 213-9 - 11 -8 110-6 7. 0 103.2 177-235 82-112 95-123  HURST  ±  HIND FOOT LENGTH 23.0 22-24  N  •7  21  210-3 ± 13 -0 105.6 ± 8- 0 104.3 6.8 165-228 73-114 84-118  23.7 ±1 .2 20-24  25  BALACLAVA  187-1 - 8 -2 91 -3 ± 5- 9 165-206 81-106  21.4 20-23  -8  47  NIGEI-  212.9 ± 13 .3 194-280  108.4 ± 6. 0 104.4 ± 1 2 . 4 87-108 99-125  24.8 ± .8 23-26  59  VANSITTART  197-0 ± 7-9 186-270  101.2 - 2. 5 96-106  95-8 ± 7.1 85-110  22.0 ± -4 21-23  15  HOPE  189-9 ± 165-225  9.8  92 •9 ± 8. 4 76-121  97.1 ± 4.9 79-116  21.6 20-24  -9  78  COX  209.3 ± 17.6 125-234  108.7 ± 7.3 103.0 ± 5 . 9 90-125 88-112  25 .0 ±1 .3 21-27  43  LANZ  208. 9 ± 10.2 189-239  106. 8 ± 7. 7 102.1 ± 5.3 94-128 93-116  25 .4 ± .9 56 23-27  TRIANGLE  224.8 ± 10.9 187-250  116. 4 ± 9. 1 104.9 ± 1 2 . 4 89-130 93-116  26.5 % -1 24.5-29  BOWEN  173.0 + 154-200  8.5  THORMANBY  178.6 ± 153-194  9.5  TEXADA  176.7 ± 11.3 156-201  SAVARY  175.1 ± 160-197  ±  i  ±  ±  ±  67  4. 6  94.0 ± 6.6 80-116  19.6 - .6 34 21-23  5  92.3 ± 6.7 72-100  21 .1 -9 19.5-23  26  61-V3 "  6  97.8 ± 4.3 90-108  22.0 ± -7 20.5-23  62  M  1  93.4 ± 5.6 85-110  21 .6 ± -9 20-23  32  79. 1 ±  75-98  8.0  97-5 5.3 83-108  ±  78- 9l 4  86  4  9  78  4  -  £  o  MAXILLARY TOOTHROW  o • +1  co  co  00  •  CO POSTPALATAL LENGTH  O • + 1 rH  Q  rH  co  DIASTEMA WIDTH  O +"l  o  00 rH  PALATINE FORAMEN  SHELF OF BONY PALATE  BREADTH OP SKULL  0CCIPIT0NASAL  LENGTH  SAMPLE S I Z E  CM o  + 1 00  ON  •  rH  +vOi o  + 1  CM ON  +"l ON trH  CO CO o +? 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Th  00  ON  CM  CM  CM  CM  O co  in rH  Th rH  Th co  o  co  rt sn  X O  o  H  cd  o  CD  m  -P Pl  CD  S  u pi  m  ca CD  E  ca •H Pi  +  I  . 0) tuO  ca  O  cr3  U  pq  o  a  CD  O  CM  pq  >  00  CM  CM  crj •H rH  CM  Th vO  Pl  O  CM  o  rH Cd  O  ON rH  o +i  CD  Th  rH  CM  o  m  CM  Th  <  EH EH l—l CO  o + 1  SI  <  0)  > crj Pl  S3  ca  i—1  s  rt EH  <! Th  CD  rH  ca  EH  SUBSPECIES*  AVERAGE ADULT WEIGHT  MEAN LITTER SIZE  balaclavae Hope I .  24.3gr.  4.5  2.07gr.  2  13-15  23  balaclavae 24.8 Balaclava I .  4.0  2.24  2  15-17  4  t r i a n g u l a r i s 33c? T r i a n g l e I . 302  4.7  carli Cox I .  28.1  4.0  carli Lanz I .  28.6  5.4  georgiensis Texada I .  27.4  3.3  ? 24.6 Vansittart I.  4.0  isolatus Nigei I.  2.0  30.5  AVERAGE ELEVATION NEW BORN OF PINNA WEIGHT I N DAYS  OPENING N.F1 OF EYES I N DAYS  14 2.40  2  15-20  38 3  16-17  10  2.30  3.5  14-18  8  2.83  3  2  *Cowan and G u i g u e t , 1965 Table  15A  R e p r o d u c t i v e and d e v e l o p m e n t a l conceived l i t t e r s .  SUBSPECIES SUBSPECIES MEAN c? * 2 * LITTER SIZE  data f o r f i e l d  AVERAGE ELEVATION NEW BORN OF PINNA WEIGHT IN DAYS  OPENING OF EYES IN DAYS  —  doylei Doyle I .  austerus mainland  2  3.4gr.  2  carli Cox I .  triangularis Triangle I .  7  2.4  4  21  carli Cox I .  isolatus Nigei I.  3  3.6  3  18  balaclavae ? 4 Hope I . Vansittart I.  -2.2  2  17  carli triangularis 5 Lanz I . Triangle I. *Cowan and G u i g u e t , 1965  2.9  2  20-21  T a b l e 16A  19  R e p r o d u c t i v e and d e v e l o p m e n t a l i s l a n d h y b r i d Peromyscus  data f o r i n t e r  ISLAND  N  PROXIMAL END OF BACULUM  B  A  n  DISTAL T I P OF GLANS  I  DORSAL LAPPETS  G  VENTRAL LAPPETS  F  0  D  CONE  RIDGE  4  5  0  0  9  DOYLE  9  4  5  3  H 6  HURST  2  0  2  1  1  0  2  2  0  0  BALACLAVA  8  6  2  7  1  8  0  4  2  2  NIGEI  9  0  9  9  0  2  7  6  3  0  VANSITTART  11  1  10  3  8  7  4  3  4  4  HOPE  19  0  19  19  0  5  14  15  4  0  COX  7  0  7  2  5.  1  6  4  3  0  LANZ  6  0  6  1  5  2  4  5  1  0  TRIANGLE  7  6  1  7  0  0  7  0  1  6  12  0  12  6  6  1  11  1  0  5  0  5  5  0  2  3  3  2  0  18%  82%  66%  34%  34%  66%  56%  22%  22%  TEXADA THORMANBY  T o t a l , & % o f , 95 Table  17A  The p h a l l u s c h a r a c t e r i s t i c s  of i s l a n d  Peromyscus  11  ISLAND  DISTAL TRACT LENGTH  GLANS  BACULUM CART LENGTH DIAMETER BONE TIP  HIND FOOT LENGTH  RATIO X 10QO GLANS/ BONE/ DIAM.GLANS/ FOOT FOOT LGTH.GLANS  N  DOYLE  1 .26 ( .04)  0.89 (.04)  0.20 (.02)  1 .07 (.04)  0.13 (.02)  23. 0  38.7  46. 5  224.7  9  HURST  1 .26 (.03)  0. 89 ' (.06)  •0.1 8 (.01 )  1 .05 (.04)  0.13 ( .02)  23. 7  37.6  44.3  202.2  2  BALACLAVA  1 .05 (.07)  0.76 (.04)  0.1 8 (.02)  0.87 (.06 )  0.13 (.01 )  21 .4  35.5  40.7  236.8  8  NIGEI  1 .26 (.04)  0.93 (.04)  0.20 (.04)  1 .03 (.03)  0.14 (.01)  24. 8  37.5  41 .5  21 5.0  9  VANSITTART  1 .03 (.07)  0.74 (.06)  0.1 8 (.02 )  0.84 (.06)  0.13 (.01 )  22. 0  33.6  38.2  243.2  11  HOPE  1 .05 (.06)  0.75 (.06)  0.17 (.03)  .88 (.04)  0.1 2 (.01 )  21 .6  34.7  40.7  226.7  19  COX  1 .25 (.06)  0.89 (.06)  0.20 (.02)  .99 (.04)  0.14 (.02)  25. 0  35.6  39.6  224.7  7  LANZ  1 .25 (.03 )  0.87 (.06)  0.19 (.04)  1 .00 (.03)  0.1 1 (.02)  25. 4  34.3  39.4  218.4  6  TRIANGLE  1 .32 (.11)  0.98 (.05)  0.23 (.03)  1 .07 (.07)  0.13 (.02)  26. 5  37.0  40.4  234.7  7  THORMANBY  1.13 (.02)  0.82 (.04)  0.22 (.02)  0.95 (.02 )  0.10 (.01)  20. 3  40.4  46.8  268.3  5  TEXADA  1.12 (.05)  0.83 (.04)  0. 18 (.02)  0.95 (.05)  0. 12 (.01 )  22. 0  37.7  43.2  216.9  12  HYBRID (Ttngl/Cox)  1.18 (.11)  0.84 (.07)  0.19 (.07)  1 .00 (.07)  0.01 (.01)  25. 0  33.6  40.0  226.2  3  0.19 0.97 0.12 by b r a c k e t e d v a l u e  23. 2  36.7  41 .9  228.3  AVERAGE 1.18 standard deviation Table  18A  0.85 indicated  Mean p h a l l i c measurements and r a t i o s f o r i n s u l a r  Peromyscus  + + ++ ++  ++ ++ 4~f  ++ ++  HYBRID  ++ ++ + +  ++ ++ ++ ++ ++ COX  ++ ++  ++ ++ ++ ++ ++ + HOPE  ++ ++ ++ ++  ++ ++ + ++  + + ++ HURST  + ++ THORMANBY ++ ++ TEXADA ++ DOYLE BALACLAVA  ++ NIGEI VANSITTART  LANZ ++ s i g n i f i c a n t  TRIANGLE  + significant Table  19A  Significance  o f v a r i a n c e between mean b a c u l a  at less  t h a n Vfo  a t 5%  measurements  oo  ++ ++  ++ ++  ++  ++ .++  + .++ ++  ++  ++ ++  ++ ++  ++ ++ COX  ++ ++ VANSITTART  TRIANGLE  Table.  20A  ++ ++ ++ HURST  + ++ ++ BALACLAVA  ++ THORMANBY ++ TEXADA DOYLE  NIGEI  HOPE  LANZ HYBRID  ++ ++  ++ ++ ++  + +  + ++ ++  S i g n i f i c a n c e of variance  ++ s i g n i f i c a n t a t l e s s + s i g n i f i c a n t a t 5%  between t o t a l  distal tract  length  than l f o  measurements  VO  APPENDIX  2  BODY, TAIL AND BODY/TAIL HISTOGRAM  DATA  RATIO  1 5 » OJ.  DOYLE ISLAND  10.01  0°0.  i i 11  70 20 . O4-  T A I L  Fl i M  111  IT-  ++  L E N G T H  15.0.  LO.Q  5-01  0«0.  ?0 ^  30°0l  20*04-  lO-Ol  126 E D D Y  /  T A I L  R A T I O  20 *Q.  15'-0..  HURST ISLAND  10 ' 0 -  5»CL.  0 = 0. I N N  tftft-  70  126 T A I L  20'0.  L E N G T H  15 • 0..  10.0_.  5»Q  0-0.  -ft  I I I I I I I I I I  I I I I II  70  J-cr.  B O D Y  40-0.  30«0„  20-0.  O'O.  o*: L  /  T A I L  R A T I O  15.0..  BALACLAVA  10.0.  5.O..  I I I In  70 20. OX-  T A I L  I l l I I l l I I]  12S  L E N G T H  15-0J  10.0..  5»0-  O.ojlj70 40'0  H  30.0-  20.0.  10'0.  B O D Y  I I I I I 1 I I I II /  T A I L  R A T I O  126  ISLAND  20°0__  15-0..  NIGEI ISLAND  10-0..  5-0..  0<0j 1 1 1 1 1 1 1 m  r  ft  H-  70  133 T A I L  L E N G T H  15-0..  10-0.  5-0..  0*0. 70 40*0^  1 I I I II I I I I I 12G E D D Y  /  T A I L  R A T I O  E0<0-  10 * OJ  0«0J  H  F  15-01  VANSITTART  10»0.  70 20 »0.  111111 n i n T A I L  r  T  in  +++ 1EG  L E N G T H  15-01  10-ol  5 " 0  11 I I 11 I 11 I 1 70 40.0.  B O D Y  /  T A I L  I I I I I I 1I 1 1EG R A T I O  30.01  E0« OJ  1 0 . OJ  o.oJ •1 .  c  ISLAND  15»0._  10 »0_.  5»n  •  70  EO'CL-  1EG  T A I L  L E N G T H  15«0..  10-0..  5»0 J.  0<0_ L2G  70 40'O.,.  3 0 '  0.  10*0..  O'OJ 0<5  B O D Y  /  T A I L  R A T I O  15-0.  COX  ISLAND  10-0..  0..  0-0.  l I I 11 I i 126  70 T A I L  L E N G T H  15-0.  10-0..  5-0.  O  • oft  -H-  70  B O D Y  126 /  T A I L  R A T I O  40»0_  30'0.  10-0.  0*0.  H  h  B O D Y  128  L E N B T H  15-0.  10-0-  5-0..  0>n  I ll I l I i 126  I I 1 I I I 70  01  20-04-  T A I L  L E N G T H  15-0.  10-0..  5-0..  4  0-0. 70 AO  B O D Y  /  T A I L  R A T I O  12G  -O-i  30 <0..  20.0..  10 «0-  0-0. 0»5  1-5  15»0._  iO-0..  5.0.  -ft  ft  70  IEB T A I L  L E N G T H  E0-0.L.  15.0.  lOCL.  5*0.  ft4  0-0. ft 70 40*0+  r IE  B O G Y  /  T A I L  R A T I O  3C  iO-0_.  0«5  1«5  15-OJ  10  .SAVARY ISLAND  -OJ  5-0j  4  0-0J 70 T A I L  L E N G T H  B O D Y  /  -fl-  I 1 I 1I 1 I 12G  15-OJ  10-OJ  5-01  0-OJ  70 4 0 -OJ  30.01  20.01  10'Oj  O-OJ  T A I L  R A T I D  1EG  B O D Y  L E N G T H  10»0J  5-01  O'OJ  MM 1E6  70 ?0*0J  T A I L  L E N G T H  15 "OJ  10.01  5-0j  0«0J  -H-  70 40 "Oj  30« OJ  3>0J  10 -0.  B O D Y  /  T A I L  R A T I O  1ES  15»0_.  10-0..  i-0._  O'O. 70  15G T A I L  L E N G T H  20-0+  15-0.  10.0..  5»0.  0-0.  -H-  70  126 B G G Y  4 0 . OJ  30*0..  E?0«0..  10.0..  0*0-  /  T A I L  R A T I O  B Q D Y  L E N G T H  15-OJ  10'OJ  5-OJ  0-OJ 70  12G T A I L  L E N G T H  20-OJ  15-01  10-OJ  5-OJ  0-OJ  ++ 70 B O D Y  4 0 - OJ.  30-Ol  20-01  10-0.  I I I fl  I I I I I I I I I I I | || 12G  /  T A I L  R A T I O  APPENDIX 3  DISCRIMINANT ANALYSIS DATA  APPENDIX 3 coefficients skull  length  basilar  Doyle  legend skull  breadth length of nasals  length i n t e r o r b i t a l constriction  l e n g t h of animal  body  length  shelf of diastema bony p a l a t e tail  Post pfalatine slit maxillary toothrow  length hind foot length  115.58285 388.15087 0.27753  1393.79638 135.38287 242.75891 1466.40918 16.12768  112.45332 406.14080 0.22558  1293.95312 206.02777 214.87585 1553.04370 16.49320  99.29972 305.24835 0.23036  1321.77954 161.97552 258.00543 1345.94604 15.21731  100.85078 429.90661  1279.41699 165.33013 228.24316 1490.66235  Island  c o n s t a n t ; -1155.21997 coefficient ' 8.15174 -83.03508 -212.10366 9.56216 0.76581 -0.91500 Balaclava  Island  c o n s t a n t ; -979.97265 coefficient 0.95112 -62.83573 -169.53933 7.32085 0.68174 -0.91393 Nigei  palatine slit  Island  c o n s t a n t ; -1125.14136 coefficient -10.95095 -41.16622 -226.40087 8.93644 0.63262 -0.65659 Hurst  DISCRIMINANT FUNCTIONS  Island  c o n s t a n t ; 1159.56103 coefficient -8.98146 _54.98657 -114.14604 5.09999  Vansittart  Island  c o n s t a n t ; -97.68335 coeffieient 16.83068 -61.70179 -148.75589 5.71217 0.49978 -0.56468 Hope  97.19392 303.12951 0.23543  1281.33203 175.89895 273.11566 1341.25610 15.30642  101.25082 409.82568 0.29151  1325.34765 174.05728 200.53790 1492.62720 18.19372  102.98414 417.34265 0.26982  1347.10400 160.96511 196.18627 1482.98535 19.0298  104.68608 389.93750 0.35650  1348.99634 185.62478 232.68136 1455.26855 19.99827  129.70477 349.13821 0.23716  1124.55566 163.08029 253.90820 1382.68359 15.67136  Island  c o n s t a n t ; -1154.35473 coefficient -0.98899 -62.37367 -159.94216 7.01263 0.61067 -0.73399 Lanz  1399.22436 113.95394 290.26428 1228.05469 16.36004  Island  c o n s t a n t ; -984.45080 coeffieient -1.24584 -64.11415 -133.79235 6.17992 0.61364 -0.88389 Cox  64.80624 279.70916 0.27029  Island  c o n s t a n t ; -1170.90210 coefficient -12.63926 -48.35587 -119.78697 5.24048 0.61158 -0.82431 Triangle  Island  c o n s t a n t ; -1238.74145 coefficient -6.99522 -62.34459 -121.41917 5.63358 0.58385 -0.65399 Bowen I s l a n d c o n s t a n t ; -941.91906 coefficient -34.08764 -28.06115 -145.81207 . 6.45590 0.55917 -1.09760  Thormanby  Island  c o n s t a n t ; -939.92675 coefficient -5.44735 -46.25228 -145.16732 6.13494 0.52232 -0.89130 Texada  1292.68823 144.65267 268.43298 1300.15503 15.17016  111.96975 341.904066 0.24885  1196.63550 177.41284 284.60461 1398.46069 15.61571  115.29104 288.40869 0.23160  1262.45849 168.59378 . 305.41015 1350.46411 15.83843  Island  c o n s t a n t ; -996.96435 coefficient -12.39646 -52.04350 -153.53219 6.93321 0.60903 -1.26903 Savary  92.12721 , 313.63641 0.25322  Island  c o n s t a n t ; -981.61413 coefficient -15.51098 -49.00769 -154.90383 6.84798 0.50345 -0.98707  Island  N  Discriminant analysis from expected.  deviations  Doyle  20  H u r s t (1)  Hurst  24  Cox (2)  Balaclava  42  Hope ( l l ) , Vansittart  Nigei  54  L a n z ( 1 0 ) , Cox ( 2 ) , H u r s t (2) T r i a n g l e (2)  Vansittart  14  none  Hope  75  Balaclava Thormanby  Cox  31  Hurst (4), Nigei T r i a n g l e (4)  Lanz  37  Cox ( 8 ) , N i g e i ( 2 ) , T r i a n g l e ( 7 ) , Hope ( 1 ) , H u r s t ( l )  Triangle  61  Nigei  Bowen  14  Texada ( 3 ) , Thormanby Balaclava ( l )  Thormanby  19  Savary  ( 2 ) , Hope  Texada  73  Savary  ( 6 ) , Bowen  Savary  19  Hope  Table  22A  T  Thormanby (2)  (5) Texada ( l )  ( 1 2 ) , Cox ( 4 ) , Bowen ( 3 ) , ( 8 ) , S a v a r y (2)  ( 2 ) , Lanz  ( 2 ) , Lanz ( 7 ) ,  ( 6 ) , Cox (5) ( 2 ) , Savary ( l ) ,  (2), Balaclava ( l ) ( 1 2 ) , Thormanby ( 3 ) ,  ( l ) , Bowen ( l ) , B a l a c l a v a  (2)  d e v i a t i o n s from the c o l l e c t i o n l o c a l i t y b a s e d upon d i s c r i m i n a n t a n a l y s i s . Sample s i z e s i n d i c a t e d by t h e b r a c k e t e d v a l u e . n  e  APPENDIX 4  KARYOTYPES OP INSULAR PEROMYSCUS  A A A AA*  A A *  KX X ft IflH * » «  j  ^  /)(! fi A I Y O A ; x Y f\ A A A A A A *» A c  » Peromyscus m a n i c u l a t u s  georgiensis  Bowen I s l a n d (-32C/62 11')  A t y p i c a l , s m a l l b o d i e d , l o w P.N. ( 7 4 ) , c o a s t a l m a n i c u l a t u s . T h i s k a r y o t y p e i s r e p r e s e n t a t i v e o f t h e Thormanby, S a v a r y , V a n s i t t a r t , and Hope I s l a n d p o p u l a t i o n s o f P e r o m y s c u s .  iuixi'utxtiiiu' |M«  Unih  Cox I s l a n d (148) Peromyscus s i t k e n s i s i s o l a t u s * T h i s k a r y o t y p e i s t y p i c a l f o r t h e l a r g e morph i n s u l a r  Mllixitxitt*** jf J Nigei  Island  (El)  Peromyscus  sitkensis  isolatus*  Doyle  Island  (ll)  Peromyscus  sitkensis  doylei*  x  revised  nomenclature  Peromyscus  *% M  ft *  HA  mm  mm  H  T e x a d a I s l a n d (229)  t i l  l  I  M  A » » #• A  /%  «  Jf  m)  Peromyscus m a n i c u l a t u s  Al  georgiensis  I SI II  fli  Hope I s l a n d (174D)  Peromyscus m a n i c u l a t u s  H u r s t I s l a n d (E19)  Pe r o m y s c u s s i t k e n s i s  georgiensis  saxamans  #  APPENDIX 5  SPECIMENS EXAMINED  APPENDIX  5  SPECIMENS EXAMINED U.B.C. The U n i v e r s i t y o f B r i t i s h C o l u m b i a V.P.M. The B.C. P r o v i n c i a l Museum, V i c t o r i a M.V.Z. The Museum o f V e r t e b r a t e Z o o l o g y , B e r k e l e y M male,P female UbC ubC UBC UBC VP I'M VPM VPM  013M 2 3 5M 016M 1196M 379 0M 378 7M 37 8 4M  UBC UBC UBC  VPM VPM VPM  012M 2 34M 015M 37 9 3. , 37 8 9M 37 8 6M  VPM VPM VPM  2 3 6M 2 7bM 1 19 5M 3 79 1M 376bM 3 76 5 i-'i  UBC UBC VPM VPM VPM VPM VPM VPM  12 06M . O20M 36 73M 36 7 0M 36 6 7M 36 64M 36 61M 36 5 8M  UBC UBC VPM VPM VPM VPM VPM VPM  019M 120 7M 3 67 2M 3 66 9M 3 66 6M 3 66 3M 3 66 0M 3657M  VPM VPM VP. . VPM  3833M 38 4 5M 3850M 3 82 7M 38 6 3M 38 6 0M 3 3 49M 1 2 0 3M 38 5 8M 3855M 0 3 0M  UBC  UBC UBC  DOYLE ISLAND  UBC UBC VPM VPM VPM VPM VPM VPM  U18M 022M 3674M 36 71M 3663M 366 5 M 3662M 3659M  HURST ISLAND  VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM  33Z 6M 38 3 3M 3846M 38 5 i M 38 6 6M 38 62M 38 5 9M 364 7M 384 1M 38 57M ' 38 5 3M  UBC UBC UBC UBC  02 6M 2 7 5M 02 4M 2 7 6M  BALACLAVA  ISLAND  VPM VPM VPM VPM VPM VPM VPM. UBC VPM VPM VPM UBC UBC UBC UbC  38 3 2M 38 3 9M 38 48M 38 2 9M 38 64M 38 6 1M 38 5 4M 1202M 3 86 5 M 3 8 56 M 38 5 2 M  VPM 'VPM UBC VP I-I VPM, UuC  023.-. 274M 0 3 1M 01 OM  UBC UBC UBC UBC  VP.-i  2 4 9M  0 29M 2 5 0M 0 2 7M  UBC UBC UBC UBC UBC UBC VP,-, VPw VPM VPFi V P i-i VP M VPM VPM VP;-, VPM VPf-i VPM UBC NIGEI  UBC UBC UBC UBC UBC  146M L 0 5M 1 3 9M  E01M 3 6 9 7M 3718M  a  3 7.2 M 3 7 2 2M 369 3 M 3 6 9 9M 37 0'nM 37U7M 3712M 3 7 1 5M 3 7 2 3M 37 2 6M E 0 8M  UBC UBC UBC UBC UBC UBC UBC  1 7 0M 1 3 4M 1 7 5M 1 2 8M 13IM  3708M 3 719M 3 7 2 9M 3 6 9 6M 36 92M 3 70 IM 37.0 5 M 3 7 0 9M 3713M . 3717M 37 2 4 M 37 2 7 M  UBC UuC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM UBC  L12M IH-GM  144M L16M L C 2 f3 69'nM 3 7 1 Ci-; 3 721M 3 8 44M 3 6 9 5M 3 6 9 IM • 3 703M 3 7C6M  in-;  37 3 7 14-M 3720M 3 7 2 5M E15M  UBC UBC UBC UBC UBC  007M 00 2 M 12 1 M 2 71M 2 5 2M 0 0 3M 2 5 8M 2 5 7M  UBC UBC UBC UBC UBC UDC UBC UBC UBC  2 7 9M  HOPE ISLAND  1 6 9M 130M 135M 125M 1 2 7M  UBC UDC  17 8M 124M  ULSC  132M  UBC UBC  1 2 9M U0 9M  UBC  OOH-M  UBC  1 2 3M 2 7 2M 2 5 5M 2 5 4M 2 7 0M 17 3M 2 6 9M 2 5 IM 3 7 7 7M  ISLAND  UBC UBC  13 8 M E 0 3M 137M 136M BUM  ISLAND  VANSITTART  UBC  N  UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM  UBC  006M 0 0 IM 2 3 7M 174M 2 5 3M 0 0 5M 2 4 8M 28 6M 2 4-7M 2'f6M  UBC UBC UBC UBC UBC UBC UBC VPM  VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM  UBC  3738M 3746M 3752M 375 9M 3764M 3772M 37 3 6M 37 5 5M 375 8M 377 0M 3773M 2 5 6M  VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM UBC  37 39M 3748M '3753M 3760M 376 5M 377 5M 37 3 7M 37 56M 37 61M 37 6 9M 37 6 8M 245M  VPM VPM VPM VPM VPM VPM VPM VPM VPM VPM UBC VPM  3741M 3 75 1M 37 5 4M 3 76 3M 3767M 3 776M 3 7 40M 3 757M 3 762M 3 771M 277M 37 4 5M  UBC  105M 5607M 5610M 5615M 5619M 5623M 5627M 102M 14 8M 165M  VPM VPM VPM VPM VPM VPM  5 604M 5 60 8M 5611M 5 6 16M 5 620M 5 62 5M 10 9M 10 3M 162M 15 7M  HOPE ISLAND  UBC VPM VPM VPM VPM VPM VPM  UBC UBC UBC  UBC  162M 5606M 5609M 5613M 5605M 5621M 5626M 105M 147M 055M 15 9M  VPM VPM VPM VPM VPM VPM  UBC UBC UBC  UBC UBC UBC UBC  COX ISLAND  UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM VPM  117M 04 2 M 113M 051M 55 8 0M 5585M 5591M 5602M 5582M 5587M 5594M 559 7M  LANZ ISLAND  UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM VPM  112M 0 50M 119M 118M 558 1M 5589M 5600M 5 57 7M 5 5 84M 5 5 92M 5595M 5 59 8M  UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM VPM  12 0M 0 5 3M 0 5 2M 04 1M 5 58 3M 5 59 0M 5 60 1M 5 57 8M 5 58 6M 5 59 3M 5 59 6M 5 60 3M  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM VPM  9222M 9216M 9232M 92 3 0M 9225M 9215M 9220M S06M 063M 069M 19 7M 16 5M 17 9M 066M 545AM 5AA9M 5442 M 5AA AM 5447M 545 3M 54 5 8M  rRIANGLE  VPM VPM VPM VPM VPM UBC UBC UBC UBC VPM LANZ  UBC UBC UBC  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM UBC  9221M 9 2 29M 92 2 3M 92 27M 92 24M 9218M 50 9M 508M 06 AM 18 8M 189M 192M 181M 0 6 8M 54 5 7M 5AA8M 5441M 5AA5M 54 5 0M 5A5 5M 190M  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM  92 17M 9233M 9 23 IM 9 2 26M 921AM 9219M 50 3M 18 0M 0 6 5M 19 IM 18 3M 19 8M 18 6M 0 7 0M 5 A5 2M 5 AA3M 5AA0M 5AA6M 5 A5 IM 5 A5 6M  VPM VPM VPM VPM VPM UBC UBC UBC UBC VPM  68819M 6 5947M 6 595AM 6 59 73M 33 8 AM 67 5 2M 033M. 87 5 AM 0 3 2M 68 817M  VPM VPM VPM VPM VPM UBC UBC UBC VPM  6 8 8 18M 65952M 65948M 6 5 97AM 3A5 IM 87 5 5M 0 3 5M 03 IM 68 816M  UBC UBC UBC  2 6 0M 2 2 0M 2 2 1M  ISLAND  68820M 6 594 6M 65972M 6 595 3M 33 8 3M 875 IM 8753M 034M ODJM 68815M ISLAND  262 M 218M 222M  THORMANBY  ISLAND  UBC UBC UBC  2 59M 219M 216M  UBC UBC UBC VPM  263M 261M 2A2M. 7 0 4 2 AM  UBC UBC UBC UBC  26 AM 15 2M 2A3M 15 3M  UBC UBC VPM  2A IM 2A0M 70A2bM  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM UBC UBC  01AM DO CM DOFM 015M 016M 1781M 17 8 5M 215M 2 2 8M 2 68M 2 11M 2 06M 2 33M 2 29M 2 3 8M 2 81M . 2 12M 70A08M 70A15M 70A0AM 70A12M 70A03M 70A21M 2 39M 178 8M  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM VPM VPM VPM UBC  DO AM DO DM DOGM 013M 0 0 3M 178 2M 1 78 6M 28 3M 2 3 0M 2 8 2M 20 9M 2 0 AM 23 2M 2 2 7M 2 8 AM 2 1AM 20 5M 70A11M 70A 1AM 7 0A0 7M 7OA13M 7 0 A 0 5M 7 0 A 0 2M 2 2 6M  VPM VPM VPM UBC UBC UBC UBC  703 8 3M 7 0 3 9 IM 7038IM D0 6M 00 5M 0 1OM M  VPM VPM VPM UBC UBC UdC UBC  70 3 8AM 70 3 8 9M 70 3 8 7M D0 7M G0 6M D0 3M M  THORMANBY ISLAND  UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC UBC VPM VPM VPM . VPM VPM VPM UBC  012M DOBM DOEM DOHM 213M DOIM 1783M 1769M 22AM 267M 210M 2 0 7M 2 8 0M 2 3 1M 2 2 5M 266M. 213M 202M 70398M 70A01M 70399M 70A16M 70A06M 70A22M 26 5M  TEXADA ISLAND  VPM VPM VPM UBC UBC UBC UBC UBC  70378M 70388M 70385M 00 9M 002M 00 8M D0 5M M  MVZ  5 4019M  MVZ  540  MVZ  54022M  MVZ  5 4 0 2 3M  20M  MVZ  54 0 21M 54U24M  MVZ  54025M  MVZ  54026M  •MVZ MVZ  MVZ  9 59 8 8M  MVZ  9 59 8 9M  MVZ  9 5 9 9 0M  i-IVZ  8 889 0M  MVZ  94363M  MVZ  9 4 36 4M  MVZ  94 36 7M  MVZ  94366M  MVZ  540  54018M  MVZ  54028M  MVZ,  54 0 2 9M  MVZ  54030M  MVZ  540 31M  MVZ'  54 0 3 2M  MVZ  5403 3 M  MVZ  540  34M  MVZ  5 4 0 3 5 M  MV  z  9 5 9 8 7M  17M  MVZ  120580M  MVZ  1 2 0 5 8 1M  MVZ  1205  MVZ  120583M  M V Z  120584M  MVZ  120 5 8 5M  MVZ  120586M  MVZ  120  MVZ  120 5 8 8M  MVZ  120569M  MVZ  120590M  MVZ  1 2 0 5 9 1 M  MVZ  99753M  MVZ  754M  MVZ  MVZ  101646M  MVZ  10 16 4 7M  MVZ  1 0 1 6 4 8 M  1016  5 0M  MVZ  10165  9 62 60M  MVZ  MVZ  101649M  MVZ  MVZ  1016 5 2M  MVZ  99  587M  99  3 2M  755M 1M  9 6 2 6 1M  MVZ  96262M  96 2 6 3M  MVZ  6719  MVZ  6 7191M  MVZ  67192M  MVZ  96 2 5 8M  MVZ  96259M  MVZ  135440M  MVZ  9 56 7 3M  MVZ  ,  MVZ  MVZ  95676M  MVZ  9 56 79M  MVZ  •  MVZ  0M  MVZ  9 5672M  9 5674M  MVZ  9 5 67 5M  9 5 6 7 7M  MVZ  9 5 6 7 8M  9 5 6 8 0M  MVZ  9 5 6 8 1M  MVZ  95662M  MVZ  9 56 84M  MVZ  9 5 6 8 3M  MVZ  9 568 6M  MVZ  . 9 568 7M  MVZ  9 5 6 8 8 M  MVZ  9 568 9M  MVZ  9 56 90M  MVZ  9 5 6 9 1M  MVZ  99579M  MVZ  99 5 8 0M  MVZ  9 9 5 8 1M  MVZ  99582M  MVZ  99583M  MVZ  99 58 4M  MVZ  99585M  MVZ  9 9 56 6M  MVZ.  99 5 8 7M  MVZ  99568M  MVZ  9 9 5 9 0M  MVZ  MVZ  99592M  MVZ  113147M  MVZ  11314  MVZ  113149M  MVZ  113  MVZ  1 1 3 1 5 1M  MVZ  11315  MVZ  11315  MVZ MVZ MVZ MVZ MVZ MVZ MVZ  63 6 2 9M 83 6 3 2M 3 1 5 8 5M 83 6 39M 8 3 o 4 4M 6 3 0ibM 83 819M  2M  Peromyscus m a n i c u l a t u s  MVZ  94393M  MVZ  MVZ  8 38 3 0M  MVZ  MVZ  8 3  MVZ  83335M  MVZ  MVZ  8 384 0M  MVZ  MVZ  6 3814M  M V Z  MVZ  8 3817M  MVZ  63  3 M  M V Z  Peromyscus m a n i c u l a t u s  150M  9 9 5 9 1M 8M  3M  gambeli  8 8 6 8 8 8 8  36 2 8M 36 31M 3b 34M 38 36M 38 4 3M 38 15M 3818M  rubidus  MVZ HVZ MVZ MVZ MVZ MVZ MVZ iMVZ MVZ MVZ MVZ f iV Z iMVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ  8 38 2 0M 03823M 8 38 2 6M 87651M 8 7613 3M 87656M 8 76 59M 68 963 M 81517M 13335M 8 3A8 7M 138850M 3676M 9 7.996M A7726M 9 9A7 0M 125833M 128680M 28833M 136189M 136193M 136197M  Peromyscus  MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ  maniculatus  8 382IM 8 38 2AM 8 38 2 7M 8 76A9M 8 76 5AM 8 76 5 7M 8 7660M 6 8 96AM 81518M 6 8 95AM 8 3A8 8M 138B51M 367 7M 9 799 7M 68 7 7 7M 99A71M 12583AM 28 8 3IM 2 88 38M 136191M 13619AM  rubidus  MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ M,VZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ MVZ  83 82 2M 83 8 2 5M 87650M 8 7 65 2M 87 6 5 5M 8 76 5 8M 68 96 2M 81516M 13 33AM 68 9 5 3M 83A89M 13U852M 3 86 3 2M 9 9 A 6 7M 9 9A6 9M 9 9 A 7 2M 128679M 28 83 2M 1361 88,-i 13 6 1 9 2 M 13 6 1 9 6 M  ACKNOWLEDGMENTS I am p l e a s e d t o l i s t  h e r e t h e many p e r s o n s who have  a s s i s t e d me i n some way i n c a r r y i n g o u t t h i s s t u d y . gratitude  goes t o t h e f o l l o w i n g v o l u n t e e r s  o f a d v e n t u r e j o i n e d me on one o r more o f my  My  who f o r t h e s a k e expeditions:  F a b i a n Anderson, B r a d l e y Ven H u i z e n , Bruce Hepburn, W a l t e r Wogee, L a r r y E v a n s , a n d D r . H. S t i c h and f a m i l y . all  o f t h o s e l i s t e d w o u l d j o i n me i n t h a n k i n g N.  for the loan of h i s excellent  I am s u r e Brearley  tent.  I am a l s o p l e a s e d t o t h a n k Don C o o k s h a n k a n d t h e proprietor of the Port o f whom were o f g r e a t  H a r d y B u i l d i n g S u p p l y Company, b o t h help  t o me d u r i n g  my s t a y s  i n Port  Hardy. I am i n d e b t t o D r . M i l l e r to Triangle  forhelicopter  I s l a n d , a s w e l l as t h e " R o s a l i e  and  the "Mabrisa" a l l f i s h b o a t s  the  further points  o f my r a n g e .  transportation  1", "Quest 2",  t h a t have t o w e d my b o a t t o The " I s l a n d E x p r e s s " t u g  d e s e r v e s t h a n k s f o r u n k n o w i n g l y t o w i n g o u r l o a d e d b o a t 30 miles  one c o l d a n d f o g g y n i g h t , much t o t h e d i s c o m f o r t  o f my  a s s i s t a n t W a l t Wogee. Mr.  T u l l y , Tony H o l l a n d ,  Annette K i l l e e n ;  "Zac t h e R u s s i a n " , L a r r y and  t h e crew o f t h e " M a b r i s a " , a n d t h e P o r t  H a r d y B u i l d i n g S u p p l y Company p r o p r i e t o r and h i s w i f e deserve s p e c i a l p r a i s e  f o r t h e numerous m e a l s t h a t were  o f f e r e d and g r a t e f u l l y d e v o u r e d . considered  There were t i m e s when I  t h a t my c o m p a n i o n a n d I made s e r i o u s  indentations  i n t o t h e f o o d s u p p l i e s of these k i n d people; know how An  deeply  g r a t e f u l we  were f o r t h e i r  a c c i d e n t A u g u s t 2 1 , 1969,  t h a n k f u l t o two  they w i l l  has  friendship.  l e f t me  shamefully  e x - N e w f o u n d l a n d f i s h e r m e n whose names were  not o b t a i n e d d u r i n g the c o n f u s i o n .  These two  men  rescued  a s s i s t a n t at n i g h t d u r i n g a p e r i o d of rough weather despite their fishing v e s s e l M. V. for  trip  h o s p i t a l have t o be  t o A l e r t Bay  we  in  Hospital.  The  i d e n t i f i c a t i o n , I was  I am v e r y much i n d e b t e d t o thanks  nurses  watching  The for  a b l e to cash a check  and  days, f o r t h i s  him.  must a l s o go t o Dr. ¥.  Z. L i d i c k e r f o r t h e  o f s p e c i m e n s f r o m t h e Museum o f V e r t e b r a t e Z o o l o g y , a l s o t o Dr. J . B r i s t o l P o s t e r and  Charles J . Guiguet  l o a n of specimens from the B r i t i s h  loan  Berkeley, f o r the  C o l u m b i a P r o v i n c i a l Museum,  c  Victoria. My  at the  commended f o r t h e i r u n d e r s t a n d i n g .  so d o i n g a v o i d s t a r v i n g f o r t h e n e x t few  My  rescue  were t r a n s f e r r e d  manager a t t h e l o c a l b a n k , h o w e v e r , d e s e r v e s w i t h o u t any  my  and,  c o m m i t m e n t s , b r o u g h t us t o t h e  R e a d y , where a t m i d n i g h t  the f i n a l  never  s i s t e r s , J a n e t who  C a r o l y n who  h e l p e d me w i t h the k a r y o t y p e s ,  h e l p e d w i t h the m a n u s c r i p t  preparation,  and  deserve  my a p p r e c i a t i o n . Thanks f o r t e c h n i c a l a d v i c e and Dr. -Ti C.  Hsu  and M r s .  D r . J . M a r y T a y l o r , my o f my  p a n e l ; Dr.  I . McT.  Stich.  I am  comments a r e e x t e n d e d t o especially grateful  s u p e r v i s o r , and t o the Cowan, D r .  G.  to  o t h e r members  S c u d d e r , and Dr. H. S t i  all this  o f whom have g u i d e d and p r o v i d e d c r i t i c a l study.  N.R.C. g r a n t  Financial  support  (A-3462) awarded  of t h i s  comment d u r i n g  p r o j e c t was  from  to D r . J . Mary T a y l o r .  an  LITERATURE  CITED  ARAKAKI, D. T. a n d SPARKES. 1965a. C y t o g e n e t i c s o f Peromyscus maniculatus (Deer mouse). Mamm. Chromosome N e w s l e t t e r , 18:150-151. 1965b. The Chromosomes o f Peromyscus m a n i c u l a t u s hollesteri. Mamm. Chromosome N e w s l e t t e r 17:79 BANKS, R. C. 1967. The Peromyscus g u a r d i a - i n t e r p a r i e t a l i s comples. J . Mamm., 48:210-218 BENIRSCHKE, K. ( E d ) . 1969 C o m p a r a t i v e Mammalian S p r i n g e r - V e r l a g New Y o r k I n c .  Cytogenetics.  BENDELL, J . F . 1959. F o o d as a c o n t r o l o f a p o p u l a t i o n o f w h i t e - f o o t e d mice P. l e u c o p u s novebronacensis (Fischer). C a n a d i a n J . Z o o l . 37:173-209 BERRY, R. J . 1964. The E v o l u t i o n o f a n I s l a n d P o p u l a t i o n o f the House m o u s e . E v o l u t i o n 18:468-483 BLAIR, ¥ . F . 1942. S y s t e m a t i c R e l a t i o n s h i p s o f Peromyscus and s e v e r a l o t h e r R e l a t e d G e n e r a a s Shown b y t h e Bacular. J . Mamm. 23:196-204 1950. E c o l o g i c a l F a c t o r s E v o l u t i o n 4 253-275.  i n S p e c i a t i o n o f Peromyscus.  1951. P o p u l a t i o n s t r u c t u r e , s o c i a l b e h a v i o r a n d environmental r e l a t i o n s i n a n a t u r a l population of the B e a c h mouse (Peromyscus p o l i o n o t u s leucocephalus) C o n t r i b . L a b . V e r t . B i o l . U. o f M i c h . 48:41-47 BLAIR ¥ . F . and ¥ . E . HOWARD. 1944. E x p e r i m e n t a l evidence o f s e x u a l i s o l a t i o n between t h r e e forms o f t h e c e n o s p e c i e s , Peromyscus m a n i c u l a t u s . C o n t r i b . L a b . V e r t . B i o l . U. o f Much. 26:1-19 BOLE, B. P. 1939. The q u a d r a t method o f s t u d y i n g s m a l l mammal p o p u l a t i o n s S c i . P u b l . C l e v e l a n d Mus. N a t . H i s t . 5:15-77 BRADSHAW, N. 1970. S e c t i o n o f C e l l B i o l o g y , The U n i v e r s i t y o f T e x a s , M. D. A n d e r s o n H o s p i t a l and Tumor I n s t i t u t e , ,... Houston, Texas. BRAND, L . R. a n d R. E . RYKMAN. 1968. L a b o r a t o r y l i f e h i s t o r i e s o f Peromyscus e r e m i c u s a n d Peromyscus i n t e r p a r i e t a l i s . V o l . 49 No. 3:495-501  BURT, W. H. 1932. D e s c r i p t i o n s o f h e r e t o f o r e unknown mammals f r o m i s l a n d s i n the Q u l f o f C a l i f o r n i a , M e x i c o . T r a n s . San D i e g o Soc. N a t . H i s t . 7:161-182 CARL, C. G., C. J . GUIGUET, G. A. HARDY. 1950. B i o l o g y o f t h e S c o t t I s l a n d group B r i t i s h Columbia. Report o f the P r o v i n c i a l Museum o f N a t u r a l H i s t o r y a n d A n t h r o p o l o g y , pp.21-63 CHITTY, D. 1952. 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