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The identification and distribution of two species of Peromyscus in southeastern Ontario Lanko, Joyce Laurian 1962

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THE IDENTIFICATION AND DISTRIBUTION OF TWO SPECIES OF PEROMYSCUS IN SOUTHEASTERN ONTARIO  by  JOYCE LAURIAN LANKO  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  MASTER OF SCIENCE i n the Department of Zoology  We accept t h i s thesis as conforming to the required standard  THE  UNIVERSITY  OF  MAY,  BRITISH COLUMBIA 1962  In presenting  t h i s thesis i n p a r t i a l f u l f i l m e n t of  the r e q u i r e m e n t s f o r an advanced degree a t t h e U n i v e r s i t y  of  B r i t i s h Columbia, I agree t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and f o r extensive  study.  I f u r t h e r agree that permission  c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y purposes may  g r a n t e d by the Head o f my Department o r by h i s  be  representatives.  I t i s understood t h a t copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n  Department o f  Zoology  The U n i v e r s i t y o f B r i t i s h Vancouver 3, Canada,  Columbia,  Date  May  7,  1962  permission.  11. ABSTRACT One  hundred and s i x t y - n i n e mice o f Peromyscus leucopus  noveboracensis and Peromyscus maniculatus g r a c i l i s were examined for  c h a r a c t e r s t o b e s t separate them.  from an area i n O n t a r i o where i t was  Each s p e c i e s was  obtained  known to be the o n l y one  An a d d i t i o n a l 12 mice of both s p e c i e s were o b t a i n e d  present.  from an a r e a o f sympatry.  The b e s t c h a r a c t e r s found to separate  them were, i n order of importance:  ear l e n g t h , i n t e r p a r i e t a l  l e n g t h , t a i l l e n g t h , s k u l l l e n g t h , and r o s t r a l l e n g t h . ear  Although  l e n g t h and i n t e r p a r i e t a l l e n g t h separated most o f the  i n d i v i d u a l s , t h e r e were s t i l l separated.  The two  i n d i v i d u a l s t h a t c o u l d not be  s p e c i e s were found to be completely s e p a r a b l e ,  u s i n g e i t h e r one of two  indices:  Ear length X t a l l length X i n t e r p a r i e t a l length skull length or,  Ear length X t a i l  length X i n t e r p a r i e t a l length  r o s t r a l length With s k u l l l e n g t h , P. 1. 2 . 5 7 - 4 . 0 9 and P. m.  noveboracensis has an index value of  g r a c i l i s has an index v a l u e o f 4 . 2 6 - 7 . 9 0 .  With r o s t r a l l e n g t h , P. 1.  noveboracensis has an index value of  7 . 3 9 - 1 2 . 5 0 and P. m. g r a c i l i s has an index value of 1 2 . 8 3 - 2 2 . 7 7 . Six  c r o s s b r e e d i n g experiments were attempted between P.  noveboracensis and f o u r subspecies of P. maniculatus. were kept t o g e t h e r f o r p e r i o d s r a n g i n g from 71 year.  No o f f s p r i n g r e s u l t e d .  The  1.  animals  days to over a  Three p a i r s of P. leucopus  and  four p a i r s of P. maniculatus were kept f o r the same p e r i o d s o f time i n the same room, as a c o n t r o l . uced two l i t t e r s , none.  Two  One  p a i r of leucopus prod-  another p a i r produced one,  p a i r s of maniculatus produced  and the t h i r d  one l i t t e r  pair,  each and the  iii. two  p a i r s , none. P. m. g r a c i l i s  was n o t a s e x c i t a b l e o r n e r v o u s a s P. 1.  n o v e b o r a c e n s i s and was t h e r e f o r e  e a s i e r t o handle.  A l t h o u g h t h e r a n g e s o f P. 1. n o v e b o r a c e n s i s and P. m. gracilis  differ,  t h e m i c e meet i n a zone o f o v e r l a p  occur sympatrically. of  they  C o r r e l a t i o n s were made b e t w e e n t h e r a n g e s  t h e m i c e and v e g e t a t i o n ,  food  preference,  tolerance,  water r e q u i r e m e n t , morphology,  behavior.  P. m. g r a c i l i s  temperature  c o l o r o f p e l a g e , and  was f o u n d t o o c c u r  i n c o n i f e r o u s and  P. 1. n o v e b o r a c e n s i s I n d e c i d u o u s v e g e t a t i o n . was  where  No c o r r e l a t i o n  f o u n d b e t w e e n t h e r a n g e s o f t h e m i c e and f o o d  preference,  temperature  tolerance,  of pelage.  C o r r e l a t i o n b e t w e e n t h e r a n g e s o f t h e m i c e and t h e i r  b e h a v i o r was  water requirement, morphology,  doubtful.  Preliminary  t e s t s were made o f t h e a b i l i t y  t o d i s c r i m i n a t e b e t w e e n i t s o d o r and t h a t Results in  showed t h a t  a mouse e n t e r e d  a chamber c o n t a i n i n g  containing  and c o l o r  o f the other  more o f t e n  was p r e f e r r e d  species species.  and s t a y e d  t h e o d o r o f i t s own s p e c i e s .  odor o f e i t h e r s p e c i e s  chamber w i t h o u t  o f one  longer  A chamber  to the c o n t r o l  odor.  We a c c e p t t h i s t h e s i s a s c o n f o r m i n g to the r e q u i r e d standard  iv. TABLE OF CONTENTS ABSTRACT  page  TABLE OF CONTENTS  i i iv  LIST OF FIGURES  v i i  ACKNOWLEDGMENTS  ix  THE IDENTIFICATION AND DISTRIBUTION OF TWO SPECIES OF PEROMYSCUS IN SOUTHEASTERN ONTARIO INTRODUCTION B r e e d i n g Experiments  1 5  Other S p e c i e s o f Peromyscus i n S o u t h e a s t e r n Ontario  6  D i s t r i b u t i o n o f leucopus and maniculatus  6  Summary o f the Problem THE TAXONOMIC SEPARATION OF PEROMYSCUS LEUCOPUS NOVEBORACENSIS AND PEROMYSCUS MANICULATUS GRACILIS  9  MATERIALS AND METHODS C o l l e c t i o n o f Specimens  11  P r e p a r a t i o n o f Specimens  12  F e a t u r e s S e l e c t e d f o r Measurement  13  Number o f Animals Used  15  Methods o f Data A n a l y s i s  15  Experiments i n I n t e r b r e e d i n g  16  Factors Influencing D i s t r i b u t i o n  16  Experiments i n Odor D i s c r i m i n a t i o n  17  THE SEARCH FOR GOOD CHARACTERS TO SEPARATE THE SPECIES RESULTS AND CONCLUSIONS  18  Weight  19  Hind f o o t l e n g t h  19  Ear  20  length  V.  THE  Body l e n g t h  20  T a i l length  21  I n t e r p a r i e t a l width  22  Interparietal length  22  Skull length  23  Rostral length  23  Baculum l e n g t h  23  The S e p a r a t i o n o f leucopus and maniculatus Using Morphological Characters  24  The C h a r a c t e r Index f o r the S e p a r a t i o n o f leucopus and maniculatus  26  Tests of Indices  29  T e s t s o f the I n d i c e s on Mice From an Area o f Sympatry  30  Summary o f the i n d i c e s  31  Results of Crossbreeding  32  DISTRIBUTION OF PEROMYSCUS LEUCOPUS NOVEBORACENSIS  AND PEROMYSCUS MANICULATUS GRACILIS  34  H i s t o r y o f D i s t r i b u t i o n of Peromyscus  34  Present D i s t r i b u t i o n o f Peromyscus i n O n t a r i o  35  C o r r e l a t i o n o f Ranges o f Mice w i t h V e g e t a t i o n  36  R e l a t i o n s h i p o f Ranges o f Mice to Food, Temperature and Water The Morphology and Pelage C o l o r o f the Mice as an Explanation of D i s t r i b u t i o n  38 40  Behavior o f the Mice as a Cause o f D i s t r i b u t i o n  41  I n t e r a c t i o n Between S p e c i e s as a Cause o f D i s t r i b u t i o n  43  I n t e r a c t i o n Between S p e c i e s by Odor  45  Apparatus  45  f o r T e s t i n g R e c o g n i t i o n o f Odor  R e s u l t s o f T e s t s w i t h the O l f a c t o m e t e r SUMMARY  47 50  vi. APPENDIX Table  Table  I.  Measurement d a t a p l o t t e d ( i n nuns.).  II.  " t "values  calculated  i n b a r diagrams  i n comparisons o f  body p a r t s . Table  III.  Table  IV.  Table  V.  LITERATURE  Data p l o t t e d Breeding  54i n b a r diagrams o f i n d i c e s .  data.  R e s u l t s o f odor d e s c r i m i n a t i o n experiments.  CITED  52  56 57 58 59  vii. L I S T OF  FIGURES  Fig. 1.  following  Histograms  .P. m. 2(b). and  4.  13  M e a s u r e m e n t s Made  2(a).  3.  Bar P.  of weights  g r a c i l i s by diagrams m.  sex  noveboracensis  Histograms  5(a).  P. m.  19 1.  o f P.  s e x and  noveboracensis 19  age.  o f e a r l e n g t h s o f P. g r a c i l i s by  Bar diagrams  .noveboracensis  s e x and  1.  1.  19  noveboracensis 20  age.  g r a c i l i s by  .and 6(b).  Histograms P.  m.  g r a c i l i s by  .noveboracensis 7(a). and 7(b).  Histograms P. m.  of t a i l  g r a c i l i s by  s e x and 1.  age.  20  noveboracensis 21  age. 1.  l e n g t h s o f P.  age.  22  H i s t o g r a m s o f i n t e r p a r i e t a l l e n g t h s o f P. 1. n o v e b o r a c e n s l s and P. m. g r a c i l i s by s e x and age.  22  Histograms  10(a).  Bar  10(b).  diagrams  Bar diagrams  noveboracensis Histograms and  P.  m.  and  P.  and P.  g r a c i l i s by  and  P.  and  P.  m.  P.  m.  1.  o f P.  s e x and  age.  l e n g t h s o f P.  g r a c i l i s by  s e x and  1.  age.  22  noveboracensis 23 1.  noveboracensis 23  l e n g t h s o f P. g r a c i l i s by  22  1.  age.  l e n g t h s o f P. sex and age.  of s k u l l and  s e x and  g r a c i l i s by  s e x and  1.  o f P.  widths  l e n g t h s o f P.  g r a c i l i s by  Bar diagrams  m.  s e x and  g r a c i l i s by  of i n t e r p a r i e t a l  of s k u l l  noveboracensls  m.  widths  of i n t e r p a r i e t a l  Histograms of r o s t r a l and P. m. g r a c i l i s b y  13(a).  m.  of i n t e r p a r i e t a l  noveboracensis  12.  1.  21  noveboracensis  11.  noveboracensls 20  l e n g t h s o f P.  of t a i l  noveboracensis  age.  g r a c i l i s by  s e x and  20  age.  noveboracensls  9.  m.  age.  20  o f b o d y l e n g t h s o f P. and P.  Bar diagrams  s e x and  1. 1.  o f body l e n g t h s o f P.  Bar diagrams  1.  s e x and  5(b). B a r d i a g r a m s o f e a r l e n g t h s o f P. .and P. m. g r a c i l i s by s e x and age. 6(a).  noveboracensis  o f h i n d f o o t l e n g t h s o f P. and P. m.  and  age.  of weights  g r a c i l i s by  1.  o f P.  and  H i s t o g r a m s o f h i n d f o o t l e n g t h s o f P. and P. m. g r a c i l i s b y s e x and age.  and  8.  page  1.  s e x and  age.  23  viii. Fig.  following  page  13(b). B a r d i a g r a m s o f r o s t r a l l e n g t h s o f P. 1 . . , n o v e b o r a c e n s i s and P. m. g r a c i l i s b y s e x and a g e . 14(a). Key t o measurement p o l y g o n s -and P. m. g r a c i l i s . 14(b). Measurement p o l y g o n s .and P. m. g r a c i l i s . 15.  o f P. 1 .  noveboracensis 26  noveboracensis  o f the index ( I ) , E.L. X T.L. X I.L., f o r SK.L. n o v e b o r a c e n s i s and P. m. g r a c i l i s b y s e x a n d  26  Histograms P. 1 . age.  16.  o f P. 1 .  23  Histograms P. 1 . age.  o f the index ( I I ) , E.L. X T.L. X I.L., f o r R.L. n o v e b o r a c e n s i s a n d P. m. g r a c i l i s b y s e x and  29  29  17(a).  B a r diagrams o f t h e index ( I ) , E.L. X T.L. X I.L., SK.L. f o r P. 1 . n o v e b o r a c e n s i s and P. m. g r a c i l i s b y s e x and a g e .  29  17(b).  B a r diagrams o f the index ( I I ) , E.L. X T.L. X I.L., R.L. f o r P. 1 . n o v e b o r a c e n s i s and P. m. g r a c i l i s b y s e x and a g e . 29  18(a). Map o f C o u n t i e s and D i s t r i c t s Ontario.  o f southeastern 35  18(b). Range o f P. 1 . n o v e b o r a c e n s i s i n s o u t h e a s t e r n . O n t a r i o , t a k e n p r i m a r i l y f r o m R.O.M.Z. t r a p p i n g records.  35  18(c). Range o f P. m. g r a c i l i s i n s o u t h e a s t e r n O n t a r i o t a k e n p r i m a r i l y f r o m R.O.M.Z. t r a p p i n g r e c o r d s .  35  19.  F o r e s t r e g i o n s o f s o u t h e a s t e r n O n t a r i o ( a f t e r Rowe,  1959). 20.  Diagram o f odor  37 d i s c r i m i n a t i o n apparatus  (olfactometer)A6  ix. ACKNOWLEDGMENTS T h i s work h a s b e e n made p o s s i b l e b y t h e g e n e r o u s h e l p o f many p e r s o n s I and  and i n s t i t u t i o n s .  am i n d e b t e d t o D r . J . F . B e n d e l l who s u g g e s t e d  gave g r e a t a s s i s t a n c e .  necessary  facilities  gave g e n e r o u s l y Mr. Ontario. of  D r . I . MoT. Cowan p r o v i d e d t h e  f o r the housing  o f h i s time  S. M. T e e p l e  and a d v i c e .  s u p p l i e d mice f r o m t h e K i n g s t o n a r e a o f  from  and  loaned  specimens from  t h e s t u d e n t s and s t a f f  Park,  o f t h e museum.  areas o f sympatry. a t the W i l d l i f e  Station,  support  Committee o n R e s e a r c h , am g r a t e f u l  made t h i s  Algonquin Biological  L a k e O p i n i c o n , p r o v i d e d h e l p and l o d g i n g .  Financial  I  D r . D. A.  Mr. R. H e p b u r n  and D r . S. Brown and t h e s t u d e n t s a t Queen's  Station,  specimens  t h e R o y a l O n t a r i o Museum o f Z o o l o g y and  p e r m i t t e d me t o u s e t h e f a c i l i t i e s Smith  and s t u d y o f t h e m i c e and  D r . R. L . P e t e r s o n and Mr. S. Downing l o a n e d  P. m. g r a c i l i s  the problem  f o r t h e s t u d y was p r o v i d e d b y t h e P r e s i d e n t ' s U. B. C.  t o a l l o f t h e above p e o p l e  study p o s s i b l e .  many o t h e r s who a l s o  My o n l y r e g r e t  who a s s i s t e d and  i s that  I cannot  mention  c o n t r i b u t e d t o the success o f the study.  1. THE IDENTIFICATION AND DISTRIBUTION OF TWO SPECIES OF PEROMYSCUS IN SOUTHEASTERN ONTARIO. INTRODUCTION The aims o f t h i s study were t o f i n d d i f f e r e n c e s between Peromyscus leucopus noveboracensis ( F i s c h e r ) and Peromyscua manlculatus g r a c i l i s (LeConte) t o separate them a c c u r a t e l y and t o determine whether these d i f f e r e n c e s c o u l d be used t o e x p l a i n their distribution. P._m.  g r a c i l i s and P. 1. noveboracensis are two s p e c i e s o f  mice o f the Family C r i c e t i d a e t h a t are extremely s i m i l a r i n appearance  1959;  and o f t e n occur s y m p a t r l c a l l y  Cameron, 1956;  Connor, I960;  ( R u t t e r , 1951;  Waters,  Klein,  I960).  C o n f u s i o n i n i d e n t i f i c a t i o n o f manlculatus and leucopus has l o n g been a t o p i c 'of c o n v e r s a t i o n and study (Downing, p e r s o n a l communication;  R u t t e r , 1951;  Waters, I960).  M I s i d e n t i f i c a t i p n o f these s p e c i e s extends throughout where they occur o r are thought t o occur.  the area  Waters (1962) c i t e s  an example o f m i s i d e n t l f i c a t i o n i n southern New England where H a l l and K e l s o n (1959) extended  the range o f P. m. g r a c i l i s  n o r t h e a s t e r n C o n n e c t i c u t on the b a s i s o f a s i n g l e Waters has examined t h i s specimen  into  specimen.  and has concluded t h a t i t i s  not g r a c i l i s b u t r a t h e r a young a d u l t P. 1. noveboracensis• The taxonomy o f these two s p e c i e s i s f u r t h e r confused by the wide range o f c h a r a c t e r i s t i c s t h a t have been used to separate them.  The mice are g e n e r a l l y d e s c r i b e d as f o l l o w s .  Peromyscus manlculatus g r a c i l i s (LeConte).  The deer mouse  2. Is  found  i n cold,  m o i s t p l a c e s , o r deep, m o s t l y c o n i f e r o u s  1909).  woods ( O s g o o d ,  I t i s o f medium s i z e ,  t h a n o r e q u a l t o t h e l e n g t h o f t h e body  1952).  The p e l a g e  color  with a t a i l  longer  ( B u r t and G r o s s e n h e i d e r ,  i s f u l v o u s t o cinnamon-brown o n t h e  s i d e s and g r e y i s h a c r o s s t h e s h o u l d e r s and t o p o f t h e h e a d . darker or  colored dorsal  i s lacking  are white.  than that  p r e a u r i c u l a r white h a i r s  (Osgood,  Peromyscus l e u c o p u s  thought  of leucopus.  a r e l o n g and n e a r l y p a r a l l e l - s i d e d .  1909).  but the t a i l  Many a n i m a l s  1952).  the base o f t h e t a i l  The  bicolor  palatine  (Fischer).  i n size  slits  (Standfleld,  1950).  The wood  leucopus  a r e narrower  Osgood except  (1909, that  P.  palatine  43)  i s less  than that  of g r a c i l i s .  a t t h e ends t h a n i n t h e m i d d l e .  1909).  and g r a c i l i s  found g r a c i l i s  i n a variety  t o be s i m i l a r t o  t h e b r a i n c a s e was narro;«;er, t h e n a s a l s less bulging i n front  foramen, t h e a n t e r i o r p a r t slits  s t r e a k extends to  a r e p r e s e n t (Osgood,  were l o n g e r , t h e m a x i l l a r i e s orbital  woods  to g r a c i l i s  The t a i l  and more s p a r s e l y h a i r e d  W o r k e r s have s e p a r a t e d l e u c o p u s ways.  and c o l o r  The d a r k e r d o r s a l  No p r e a u r i c u l a r w h i t e h a i r s  of  have  i s s h o r t e r t h a n o r e q u a l t o t h e body l e n g t h ( B u r t  and G r o s s e n h e i d e r ,  distinctly  o f as  1909).  noveboracensis  I t i s similar  The  The p a l a t i n e  mouse p r e f e r s d r y e r , more o p e n c o u n t r y o r d e c i d u o u s (Osgood,  I95O).  The s h a r p l y b i c o l o r t a i l i s  above and w h i t e b e l o w and i s g e n e r a l l y  b e i n g more d e n s e l y h a i r e d slits  i s not d i s t i n c t  i n t h e r e g i o n o f t h e rump ( S t a n d f i e l d ,  u n d e r p a r t s and f e e t brownish  s t r e a k i n most g r a c i l i s  The  o f the i n f r a -  o f t h e z y g o m a t a was  lighter,  were l o n g e r and more n e a r l y p a r a l l e l - s i d e d and  3. the molar  t e e t h were s m a l l e r .  (1945) r e l i e d m a i n l y on s k u l l c h a r a c t e r s t o s e p a r a t e  Rand leucopus  from m a n i c u l a t u s .  mouse s e v e n long  straight  and h a d  that  slits  i n the p a l a t e . and  L e u c o p u s was  shorter,  curved  Infraorbital found  f o r a m e n when v i e w e d  that  seven  slits  i n the  palate.  from  the  side.  cover  Standfield  s l i t s was  from leucopus.  a good  In the  former,  the o u t e r s i d e s o f the f o r a m i n a are p a r a l l e l  to each  curving  I n the l a t t e r ,  i n a b r u p t l y at t h e i r  a n t e r i o r ends.  is  a g r a d u a l c u r v i n g i n o f the s i d e s o f the foramina.  of  s e p a r a t i o n o f B u r t and are  Standfield  are d i f f i c u l t  (1952)  that  other, there  The  to use  methods because  subjective.  Burt difficult  and  Grossenheider  to t e l l  the b a s i s o f t h e i r Cameron usually less has  and  inches long  p l a t e does not  t h e shape o f t h e p a l a t i n e  means o f s e p a r a t i n g a l l m a n i c u l a t u s  they  a  (1948) s t a t e s t h a t g r a c i l i s d i f f e r s f r o m l e u c o p u s i n  the a n t e r i o r b o r d e r o f the zygomatic  (1950)  m a n l c u l a t u s was  one-half inches long, with a slender rostrum  a tapered rostrum  Burt  the  and  A c c o r d i n g t o him,  a tail  apart, but tail  admit  say t h a t  these mice  t h e y c a n be  are  separated  on  lengths.  (1956) s a y s t h a t t h e t a i l l e n g t h o f l e u c o p u s i s t h a n o n e - h a l f the t o t a l  length that  i s usually  l e n g t h and  that  manlculatus  g r e a t e r than o n e - h a l f the  total  length. Waters identifying skull  these  characters.  flatter is  ( I 9 6 0 , pp. 3 3 - 3 4 ) u s e d a more e l a b o r a t e s y s t e m o f  and  species.  H i s method d e p e n d s on a number o f  In manlculatus  l o n g e r , the temporal  more w i d e l y f l a r i n g  and  the tympanic p o r t i o n of the  bulla  Is  relatively  zygomatic  the m a x i l l a r y p o r t i o n l e s s  arch  widely  flaring,  the  curvature is  sphenoids are  i s greater.  more p r o t r u s i v e and  pronounced l a t e r a l "Relatively s p e c i e s on achieved against  few  In leucopus,  the  supraoccipital region  the  curve.  post  cranial  p a l a t i n e f o r a m i n a h a v e a more He  concludes  i n d i v i d u a l s could not  the  by  be  assigned  criteria.". s p e c i e s by  saying  This  graphing  that, to the  same  body l e n g t h t a i l -length  the  mice.  These  characters  i n c l u d e "whiteness" of the  a n t e r i o r p o r t i o n of  shape o f t h e  the  neck  zygomatic arch  when  a n t e r i o r p a l a t i n e foramina,  the p o s t e r i o r p a l a t i n e f o r a m i n a w i t h  respect  and  to  the  teeth. I n summary, a v a r i e t y o f methods have b e e n u s e d  l e u c o p u s and  maniculatus•  methods t h a t a r e  difficult  p r e p a r a t i o n o f the  o f mice u s e d are used  Most o f  in this  to i d e n t i f y  study.  That  The  above a r e  Some r e q u i r e  they  completely  c a n be  the  separate  subjective elaborate  applied.  separated  to  two  so many d i f f e r e n t  None  of  populations characters  the mice i n d i c a t e s the need f o r b e t t e r ,  more q u a n t i t a t i v e methods o f apparent  the  to use.  s k u l l before  t h e methods c o n s i d e r e d  separation i n order  t o end  the  confusion. similarities  measurements,  are  author  ( p e r s o n a l , communication) uses a v a r i e t y o f  shape o f t h e  p o s i t i o n of  could  correct  skull length zygomatic width  v i e w e d f r o m above,  cheek  the  a separation of  separate  region,  and  the b a s i s of these  Rutter to  flatter,  easily  and be  b e t w e e n l e u c o p u s and  reproductive  considered  seasonally polyestrous  as and  biology two  are  maniculatus i n appearance, so g r e a t  subspecies  breed  o f one  that  they  species.  from March o r A p r i l  to  Both  5. October  o r November ( B u r t , 1940;  Kelson,  1959;  Waters, I 9 6 0 ) .  be 2 2 - 3 5 d a y s .  Litter  same:  (young/litter):  4.26  (range 2 - 6 ) ,  4.10-4.72, (1959);  (1932);  Despite  -  4.14 ±.08,  5.20 ±.36,  5.1  these  similarities,  species.  where  they both occur.  Breeding  and K e l s o n 4.3  (range  4.04 ± . 0 3 ,  5 . 0 0 ( r a n g e 1-8),  1-8),  Svihla  Hall  and  (i960).  Connor  leucopus  and m a n l c u l a t u s a r e  I n these a r e a s they  can s t i l l Breeding  t h e y a l s o do n o t i n t e r b r e e d  1933,  1937a;  be  told  experiments  i n the l a b o r a t o r y  W a t e r s , I960).  Experiments  A number o f i n v e s t i g a t o r s  ( i n c l u d i n g m y s e l f ) have  i n t e r b r e e d i n g experiments  i n the l a b o r a t o r y  o f t h e two  (1937a)  species.  l e u c o p u s were  Dice  lnterfertile  f e m a l e h y b r i d s were  found  also f e r t i l e .  of various species of leucopus  the  conducted validity  subspecies of  and b o t h male (1931,  T h i s same a u t h o r 69 attempted  and m a n i c u l a t u s .  n o v e b o r a c e n s i s h y b r i d i z a t i o n s were f a i l u r e s . to h y b r i d i z e  to t e s t  t h a t 12  In the l a b o r a t o r y  f o u n d t h a t no o f f s p r i n g r e s u l t e d f r o m  attempted  to  T h e y do n o t I n t e r b r e e d i n n a t u r e , e v e n i n a r e a s  shown t h a t 1931,  (1952);  Hall  a p a r t by e x p e r t s ( H a l l and K e l s o n , 1 9 5 9 ) .  (Dice,  Jackson  (young/litter):  (range 3 - 7 ) ,  valid  have  the g e s t a t i o n  (1932);  Bendell (1959);  Beer e t a l ( 1 9 5 7 ) ;  Kelson (1959);  found  Svihla  5 . 0 0 (range 3 - 7 ) ,  manlculatus  5.33,  4.09 ± . 0 8 ,  Burt ( 1 9 4 0 ) ;  5.17 ± . 2 9  Connor ( I 9 6 0 ) ,  (1932)  and  s i z e s o f b o t h are approximately the  (1956);  Davis  Svihla  Hall  t o be 2 2 - 3 7 d a y s and t h a t o f m a n l c u l a t u s  period of leucopus  leucopus  1952;  Jackson,  and 1933)  hybridizations  A l lgracilis  Waters  X  (I960)  four noveboracensis with four g r a c i l i s  with  6. similar  results.  apparent pairs.  He a l s o  sexual i n t e r e s t As a c o n t r o l ,  F i v e F]_ and one F  2  observed  that  t h e r e h a d b e e n no  o r a c t i v i t y between t h e mice o f these  W a t e r s c r o s s e d 10 p a i r s o f n o v e b o r a c e n s i s .  litters  resulted.  My own e x p e r i m e n t s  will  be d i s c u s s e d l a t e r . Other  S p e c i e s o f Peromyscus i n S o u t h e a s t e r n O n t a r i o The  is  o n l y other form  o f Peromyscus i n s o u t h e a s t e r n O n t a r i o  P. m. b a l r d l l (Hoy and K e n n i c o t t ) ,  Saunders  t h e p r a i r i e d e e r mouse.  (1907, 1908, 1913) r e c o r d e d t h e e a s t w a r d movement o f  t h e p r a i r i e d e e r mouse a s i t b e g a n i t s r a n g e southern Ontario.  As i t i s s m a l l e r i n s i z e  f o r m s and t h e r e f o r e r e a d i l y consider  While  of leucopus  leucopus  t h a n t h e o t h e r two  s e p a r a t e d f r o m them, I s h a l l n o t  t h i s mouse i n t h e p r e s e n t  Distribution  extension across  study.  and m a n i c u l a t u s  and m a n i c u l a t u s  many r e s p e c t s , t h e y h a v e d i f f e r e n t  are exceedingly s i m i l a r i n ranges.  The s e a r c h f o r  d i f f e r e n c e s b e t w e e n t h e s p e c i e s was n o t o n l y t o s e p a r a t e but  t o f i n d how t h e d i s t r i b u t i o n  o f each  form  was  determined.  Many a c c o u n t s have b e e n g i v e n o f t h e d i s t r i b u t i o n mice.  i n O n t a r i o show t h a t  east o f the Great Lakes noveboracensis Kelson  g r a c i l i s occupies the area  up t o t h e b o r d e r o f Quebec, w h i l e  i s restricted  to southern Ontario.  (1959) g i v e d i s t r i b u t i o n maps o f b o t h  The indicates  general d i s t r i b u t i o n that  o f these  (1909) d i s t r i b u t i o n maps o f g r a c i l i s and  Osgood's  noveboracensis  them  they occupy  a r e a o f sympatry  o f leucopus  species.  and m a n i c u l a t u s  separate ranges.  which extends.from  H a l l and  They meet i n a n  approximately  44°45  l  to  7.  approximately 4 5 3 0 ' (Rutter, 1951). 0  Osgood (1909), and Cross and Dymond ( 1 9 2 9 ) noted a c o r r e l a t i o n between v e g e t a t i o n and d i s t r i b u t i o n In these species. K l e i n (I960) commented on t h i s tendency toward e c o l o g i c a l separation of leucopus and manlculatus. He could not f i n d reasons f o r t h i s separation but suggested that i t was  perhaps  due to d i f f e r e n c e s i n microclimate. Many attempts have been made to c o r r e l a t e other f a c t o r s such as d i f f e r e n c e s i n food preference, temperature t o l e r a n c e , and water requirement w i t h the d i s t r i b u t i o n of the two species. Kinds of food eaten by the mice were reported by Dice ( 1 9 2 2 ) , Cogshall ( 1 9 2 8 ) , Burt ( 1 9 4 0 ) , Hamilton (1941), Jackson ( 1 9 5 2 ) , Cameron ( 1 9 5 6 ) , W i l l i a m s ( 1 9 5 9 ) , Connor ( 1 9 6 0 ) , Getz (1961), and Howard (1961).  Foods eaten Included v a r i o u s seeds, nuts,  i n s e c t s , araehnlds, f r u i t s , f u n g i , green vegetation, r o o t s , and molluscs.  Being omnivorous, the mice apparently eat whatever  foods are most abundant and easy t o obtain.  No d i f f e r e n c e s i n  food preference are known f o r g r a c i l i s and noveboracensis ( K l e i n , I960).  Thus food does not appear to be a f a c t o r governing the  d i s t r i b u t i o n of these mice. Sealander (1951) suggested that the northward d i s t r i b u t i o n of small mammals might be l i m i t e d by winter temperature.  Klein  ( 1 9 5 9 ) f e l t t h a t the occurrence of maniculatus at higher  a l t i t u d e s than leucopus i n d i c a t e d that temperature play-s a great role i n their distribution.  He a l s o suggested (I960) t h a t  d i f f e r e n c e s i n s o i l temperature might also be a f a c t o r .  Howard  ( 1 9 5 1 ) , a n d Eskridge and U d a l l ( 1 9 5 5 ) found that both species  8. c o u l d withstand f r e e z i n g temperatures p r o v i d e d t h a t food was  sufficient  available.  L i n d e b o r g ( 1 9 5 2 ) found t h a t leucopus and maniculatus have a s i m i l a r r e a c t i o n to l i m i t e d amounts of water. s u r v i v e on 0 . 2  cc./day and a t 0 . 4  Chenoweth ( 1 9 1 7 )  cc./day,  Neither could  each l o s t  weight.  concluded t h a t although leucopus seemed to be  a b l e to adapt t o d i f f e r e n t moist environments, an important f a c t o r i n i t s d i s t r i b u t i o n .  e v a p o r a t i o n was  When leucopus  was  kept on a r e s t r i c t e d water d i e t (Chew, 1951)» most water c o n s e r v a t i o n was E v a p o r a t i o n was  e f f e c t e d by a r e d u c t i o n i n u r i n e volume. reduced and became the g r e a t e s t path of water  An average o f 3 9 $ o f the amount of water normally drunk  loss.  ad l i b i t u m was  r e q u i r e d i n order to m a i n t a i n normal weight.  l e s s water than t h i s was  g i v e n , the food i n t a k e decreased  and  the animal l o s t weight.  Dice (1922)  not  found t h a t water was  If  the f a c t o r l i m i t i n g leucopus and b a i r d l i to t h e i r r e s p e c t i v e environments.  Klein's (1959)  study o f water i n t a k e of leucopus  and g r a c i l i s showed t h a t b o t h used about the same amount of water: day;  leucopus: gracilis:  0.43 0.42  cc./gram body weight or 7 . 6  cc./gram body weight or 7 . 4  cc./individual/  cc./individual/  day. R e c e n t l y I o b t a i n e d a copy of the t r a p p i n g r e c o r d s o f Peromyscus i n O n t a r i o from the R o y a l O n t a r i o Museum o f Zoology. I s h a l l use these d a t a i n p r e f e r e n c e to a l l o t h e r because are the most up t o date r e c o r d s o f the two eastern Ontario.  they  s p e c i e s i n south-  In addition, I c o r r e l a t e d i s t r i b u t i o n with  9. Rowe's ( 1 9 5 9 ) new d e s c r i p t i o n o f the v e g e t a t i o n o f O n t a r i o . Numerous s t u d i e s have shown t h a t the form o f an animal i s r e l a t e d to i t s d i s t r i b u t i o n (Palmgren, Lack, 1 9 4 4 ;  Horner,  1954).  1932;  Robbins, 1 9 3 2 ;  I n t h i s study I s h a l l t r y t o  c o r r e l a t e the d i f f e r e n c e s I f i n d between the s p e c i e s i n form and c o l o r with t h e i r d i s t r i b u t i o n . I n t e r s p e c i f i c i n t e r a c t i o n o f some s o r t might e x p l a i n the d i s t r i b u t i o n and a l s o the f a i l u r e o f the two s p e c i e s t o i n t e r breed I n the l a b o r a t o r y .  S t u d i e s o f the b e h a v i o r and i n t e r a c t i o n  o f these s p e c i e s w i t h each other o r w i t h other animals have been made by D i c e ( 1 9 2 2 ) , Klein (1959),  Svihla (1932),  and Sheppe ( 1 9 6 1 ) .  King ( 1 9 5 8 ) ,  Foster (1959),  Sheppe concluded t h a t the  r e s t r i c t e d h a b i t a t d i s t r i b u t i o n of P. oreas and P. manlculatus In t h e i r zone o f sympatry competition.  may r e s u l t i n p a r t from  interspecific  S i n c e mammals operate i n a world o f odor ( a s  opposed to the h i g h l y v i s u a l world o f b i r d s ) , and s i n c e these p a r t i c u l a r mammals are n o c t u r n a l , i t seemed p o s s i b l e t h a t leucopus and manlculatus might r e a c t to one another by s m e l l .  The  avoidance o f one s p e c i e s by the other might e x p l a i n t h e i r d i s t r i b u t i o n and sexual i s o l a t i o n . conducted  T h e r e f o r e , experiments were  to t e s t the r e a c t i o n s o f each s p e c i e s t o i t s own and  to the o t h e r s p e c i e s ' odor. Summary o f the Problem Two very s i m i l a r s p e c i e s o f Peromyscus, P. 1 . and P. m. g r a c i l i s ,  occur i n s o u t h e a s t e r n O n t a r i o .  noveboracensis These s p e c i e s  occupy d i f f e r e n t ranges but o c c u r s y m p a t r l c a l l y where these  10. ranges overlap. the  species  The  problem Is to  i n order to  these d i f f e r e n c e s  find differences  s e p a r a t e them and  c o u l d be  used to  explain  between  to determine their  whether  distribution.  11. THE TAXONOMIC SEPARATION OF PEROMYSCUS LEUCOPUS NOVEBORACENSIS AND PEROMYSCUS MANICULATUS GRACILIS MATERIALS AND METHODS C o l l e c t i o n o f Specimens Most o f the specimens used i n t h i s study came from two areas i n O n t a r i o , one near K i n g s t o n and the other near Lake o f Two R i v e r s , A l g o n q u i n Park. and Downing ( 1 9 4 8 ) , around K i n g s t o n .  A c c o r d i n g t o maps o f Osgood ( 1 9 0 9 ) ,  o n l y P. 1 .  noveboracensis occurs i n the area  The mice from t h i s area f i t  the s p e c i e s (Osgood, 1 9 0 9 ;  Burt, 1 9 4 0 ;  the d e s c r i p t i o n f o r  Waters, I 9 6 0 ) .  c o l l e c t i n g o f mice near Lake o f Two R i v e r s ( W i l d l i f e  Extensive  Research  S t a t i o n , Lake Sasajewan) has y i e l d e d o n l y P. m. g r a c i l i s ( w i t h the e x c e p t i o n o f a s i n g l e specimen from the Joe Lake area, A l g o n q u i n Park, t e n t a t i v e l y i d e n t i f i e d as P. 1 . (Downing, p e r s o n a l communication)).  Mice from t h i s area f i t the  d e s c r i p t i o n f o r the s p e c i e s (Osgood, 1 9 0 9 ; 1952;  Waters, The  noveboracensis  B u r t and Grossenhelder,  I960).  specimens o f leucopus were trapped by Dr. J . F. B e n d e l l  and Mr. S. M. Teeple d u r i n g 1 9 5 6 , 1 9 5 8 , and 1 9 5 9 , and sent t o me i n 10$ f o r m a l i n .  T h i s group o f 1 2 0 a d u l t , subadult, and j u v e n i l e  mice c o n s i s t e d o f 59 males and 61 females.  I a l s o r e c e i v e d 17  live  specimens o f leucopus ( 1 2 males and 5 females) e a r l y i n  1961  from Mr. S. M. T e e p l e . I c o l l e c t e d 157 a d u l t and subadult P. m. g r a c i l i s ( 6 7 males  and 9 0 females) near Lake o f Two R i v e r s , A l g o n q u i n Park. C o l l e c t i n g was done by snap-trapping e a r l y i n the f a l l o f 1 9 6 1 . .  12. These animals were aged, sexed, weighed, and measured shortlya f t e r t r a p p i n g and were t r a n s p o r t e d t o the U n i v e r s i t y o f B r i t i s h Columbia  I n 10$ f o r m a l i n .  Seven l i v e g r a c i l i s (5 males  and 2 females) were brought back to the U n i v e r s i t y o f B r i t i s h Columbia  from A l g o n q u i n Park.  Specimens o f P. m. g r a c i l i s were o b t a i n e d from the R o y a l O n t a r i o Museum o f Zoology through the kindness o f Dr. R. L . P e t e r s o n and Mr. S. Downing.  These i n c l u d e d 61 male and 39  female mice t r a p p e d i n A l g o n q u i n Park. All  specimens mentioned  thus f a r were trapped i n areas  r e p o r t e d t o c o n t a i n o n l y P. 1. noveboracensls o r P. m. g r a c i l i s . Dr. D. A. Smith sent a c o l l e c t i o n o f 12 mice (6 males and 6 females) from areas near Cloyne, Frontenac County and C a l a b o g i e , Renfrew County.  I n these areas, the mice occur  sympatrically  and are t h e r e f o r e more d i f f i c u l t to s e p a r a t e . P r e p a r a t i o n o f Specimens A l l mice except those from the R o y a l O n t a r i o iviuseum o f Zoology and from Dr. D. A. Smith were prepared f o r study i n the f o l l o w i n g way. damp d r i e d .  Animals  i n f o r m a l i n were r i n s e d w i t h water and  F r e s h l y k i l l e d animals were used without f u r t h e r '  preparation.  The animals were aged, sexed, and weighed.  Specimens  of leucopus were weighed w i t h stomach contents removed, b u t time l i m i t a t i o n prevented a s i m i l a r procedure t o be f o l l o w e d f o r maniculatus. measured.  Body, t a i l ,  e a r , and h i n d f o o t l e n g t h s were  The baculum was removed, mounted immediately on a  g l a s s s l i d e under a p i e c e o f c e l l u l o s e tape o r embedded i n c l e a r glue, and measured.  The s k u l l was removed, p a r t i a l l y cleaned,  13. and s k u l l l e n g t h , r o s t r a l l e n g t h , I n t e r p a r i e t a l l e n g t h , and I n t e r p a r i e t a l width were measured. p a l a t i n e foramina was noted. for  c o l o r and t e x t u r e .  The shape o f the a n t e r i o r  F i n a l l y , the pelage was examined  Only s k u l l measurements were made on  specimens from the R o y a l O n t a r i o Museum o f Zoology and from Dr. D. A. Smith.  Body measurements o f these mice were taken d i r e c t l y  from accompanying  museum l a b e l s .  Features S e l e c t e d f o r Measurement Features were s e l e c t e d on the b a s i s o f ease o f measurement and use by other workers i n the s e p a r a t i o n o f the s p e c i e s (Osgood, 1909;  Dice, 1932, 1937(b);  Grossenheider, 1952).  These were:  B l a i r , 1941;  B u r t and  s k u l l length, r o s t r a l length,  ear l e n g t h , body l e n g t h , t a i l l e n g t h , baculum l e n g t h , and baculum w i d t h .  I a l s o measured I n t e r p a r i e t a l l e n g t h and width.  I n t e r p a r i e t a l l e n g t h and width have not been used e x t e n s i v e l y for  comparison o f Peromyscus,  b u t examination o f the s k u l l s o f  b o t h s p e c i e s l e d me to b e l i e v e t h a t these measurements ( e s p e c i a l l y i n t e r p a r i e t a l l e n g t h ) d i f f e r e d c o n s i s t e n t l y between the s p e c i e s . The f o l l o w i n g measurements were taken: T o t a l S k u l l Length ( F i g . 1 ( a ) ) : the s k u l l ;  The l o n g e s t l e n g t h o f  from the a n t e r i o r edge o f the n a s a l s to the p o s t e r i o r  edge o f the o c c i p i t a l s . R o s t r a l Length ( F i g . 1 ( b ) ) :  The d i s t a n c e from the a n t e r i o r  edge o f the upper i n c i s o r to the a n t e r i o r edge o f the f i r s t upper premolar. Hind Foot Length ( F i g . 1 ( c ) ) :  The d i s t a n c e from the back  of the h e e l t o the end o f the l o n g e s t  claw.  <  A  <  G  TOTAL  FOOT F.  SKULL LENGTH  LENGTH  INTERPARIETAL  LENGTH  H  MEASUREMENTS  Figure 1. A.  »  D.  B.  EAR LENGTH  LENGTH  G.  »  MADE  ROSTRAL E. BODY  LENGTH  C.  INTERPARIETAL LENGTH  H.  HIND WIDTH TAIL  14. Ear Length ( F i g . 1 ( d ) ) :  The  d i s t a n c e from the bottom o f  the notch of the ear to the t i p of the Interparietal  Width ( F i g . 1 ( e ) ) :  ear. The  shortest distance  a c r o s s the i n t e r p a r i e t a l bone, taken at the p o i n t where the m i d - s a g i t t a l suture meets the i n t e r p a r i e t a l bone. Interparietal  Length ( F i g . 1 ( f ) ) :  The  greatest length of  the i n t e r p a r i e t a l bone. Body Length ( F i g . 1(g)) and T a l l Length ( F i g . 1 ( h ) ) :  The  body and t a i l l e n g t h s were obtained by p l a c i n g the r e l a x e d animal on a p i e c e o f cardboard been r u l e d .  The  on which a s t r a i g h t  nose, base of the t a i l  were p l a c e d on the l i n e .  line  and t i p of the  had tall  P i n s were p l a c e d at the t i p of the  nose, base of the t a i l , and end o f the t a i l  ( e x c l u d i n g the  tip  the d i s t a n c e between  hairs).  The mouse was  then removed and  the p i n s along the s t r a i g h t Weight: gram.  The  l i n e was  measured.  mice were weighed to the n e a r e s t t e n t h of a  Leucopus specimens were weighed with stomach  removed.  tall  contents  Shortage of time d i d not permit maniculatus to be  weighed w i t h stomach contents Baculum Length:  The  taken out.  g r e a t e s t l e n g t h o f the baculum (os  penis). Baculum Width:  The  g r e a t e s t width of the b a s a l p o r t i o n o f  the baculum. These l a s t two  measurements are not shown i n F i g . 1 because  they are a p p l i c a b l e only to male  animals.  15. Number o f Animals Used The mice were aged by pelage c o l o r ( C o l l i n s , relative  amount o f t o o t h wear (Hooper, 1 9 5 7 ) .  1923) n d a  Of the 157  maniculatus t h a t I trapped, 25 could not be a s s i g n e d w i t h c e r t a i n t y t o e i t h e r the a d u l t o r subadult category and f o r t h i s r e a s o n were not used.  Of the remaining 132 specimens,  31  animals were l a c k i n g one o r more of the measurements taken ( u s u a l l y because o f a broken s k u l l o r m u t i l a t e d t a i l ) . animals a l s o were not used i n the p r e s e n t study.  These  Thus, out o f  157 specimens o f manlculatus, a t o t a l o f 1 0 1 specimens were used f o r a l l comparisons. Of the 1 2 0 leucopus, 3 animals were s u b a d u l t s and 2 were juveniles.  These numbers were h a r d l y s u f f i c i e n t  comparative study and were t h e r e f o r e o m i t t e d . 115  to use i n a  Of the remaining  animals, 4-7 were l a c k i n g one or more measurement.  Thus a  t o t a l o f 68 leucopus were used f o r a l l comparisons except f o r the  i n d i c e s where an a d d i t i o n a l 24 animals were i n c l u d e d .  Methods o f Data A n a l y s i s Histograms and bar diagrams were p l o t t e d o f a l l measurements (except baculum l e n g t h and width) and weights taken.  Each b a r  diagram shows the t o t a l range o f measurements by a h o r i z o n t a l line,  the mean by a v e r t i c a l l i n e , one standard d e v i a t i o n on  each s i d e o f the mean by an "open" r e c t a n g l e , and two standard e r r o r s on each s i d e o f the mean by a " s o l i d "  rectangle.  The " t " t e s t f o r comparisons of groups w i t h unequal numbers of  samples i n each ( L a r k i n , p e r s o n a l communication),  a t the 5 $  l e v e l o f s i g n i f i c a n c e was used to measure d i f f e r e n c e s between  16. the  species  i n the  s i z e of p a r t s .  l e n g t h , body l e n g t h , t a i l and  length,  interparietal length.  express the  Parts tested included  The  skull length, r o s t r a l  " t " value  was  a l s o used  study  o f t h e f o r m o f l e u c o p u s and  became c l e a r t h a t no  one  part  separated  the  animals  showed a g r o u p o f c h a r a c t e r s  to d i f f e r between the  species.  show t h e s e  species.  maniculatus,  However, e a c h s p e c i e s  To  length, to  amount o f d i f f e r e n c e b e t w e e n p a r t s o f e a c h  From t h e  ear  i t  well. that  tended  tendencies,  measurement p o l y g o n s o f a d u l t a n i m a l s o f e a c h s e x w e r e  constructed.  T h e s e p o l y g o n s c o n s i s t e d o f 8 a x e s on w h i c h t h e a v e r a g e m e a s u r e m e n t s ( i n mm.) represents  o f 8 b o d y p a r t s were p l o t t e d .  an a v e r a g e a n i m a l o f e a c h s e x  Experiments i n Breeding conducted.  periods  e x p e r i m e n t s b e t w e e n l e u c o p u s and  and  ranging  C.  r a t i o n #6-61) and  3 pairs  and  w a t e r was  The  were p r o v i d e d F o o d was provided  tops of the  for  a n i m a l s were with  sawdust  i n pellet  form  i n g r a v i t y water aquaria.  Influencing Distribution  C o r r e l a t i o n was and  maniculatus,  f r o m 71 d a y s t o o v e r a y e a r .  b o t t l e s i n s e r t e d through the w i r e Factors  m a n i c u l a t u s were  4 p a i r s o f m a n i c u l a t u s were k e p t t o g e t h e r  cotton b a t t i n g f o r nest m a t e r i a l .  (U. B.  s p e c i e s o f mouse.  Interbreeding  housed i n 2 g a l l o n g l a s s a q u a r i a and  and  S i x m i x e d p a i r s o f l e u c o p u s and  of leucopus,  Each polygon  m a n i c u l a t u s and  preference, behavior.  made b e t w e e n t h e d i s t r i b u t i o n o f vegetation,  leucopus  temperature tolerance,  food  water requirement, morphology, c o l o r of pelage, T h i s was  done f r o m r e c e n t  i n f o r m a t i o n i n the  and  literature.  17. Details  o f t h e method o f a n a l y s i s  Experiments  i n Odor  Experiments  were c o n d u c t e d  I d e s i g n e d and c o n s t r u c t e d  to test  Details  on  olfaction.  odor  later.  discrimination  For this part  o f t h e work,  an o d o r d i s c r i m i n a t i o n  o f t h e method o f a n a l y s i s  d i s c r i m i n a t i o n between the mice w i l l section  be p r e s e n t e d  Discrimination  b e t w e e n l e u c o p u s and m a n l c u l a t u s .  (olfactometer).  will  apparatus o f odor  be p r e s e n t e d i n a  later  18. THE  SEARCH FOR  GOOD CHARACTERS TO SEPARATE THE RESULTS AND  The  SPECIES  CONCLUSIONSr  measurements of P. 1.  noveboracensis and P. m.  gracilis  were compared to f i n d those u s e f u l i n the s e p a r a t i o n of species.  To do t h i s ,  f o r each sex and  the  the measurements of a p a r t were compared  a v a i l a b l e ages of a s p e c i e s .  The  measurements  were compared on the same s c a l e by histograms and bar diagrams (Dice and Leraas,  1936,  These data are presented  as m o d i f i e d by Hubbs and Hubbs, 1 9 5 3 ) . f o r weight i n F i g . 2 ( a , b ) ;  l e n g t h i n F i g . 3 and F i g . 5 ( a ) ; 5(b);  ear l e n g t h i n F i g . 4 and F i g .  body l e n g t h In F i g . 6 ( a , b ) ;  t a i l length i n F i g . 7(a,b);  i n t e r p a r i e t a l width i n F i g . 8 and F i g . 1 0 ( a ) ; l e n g t h i n F i g . 9 and F i g . 1 0 ( b ) ; F i g . 13(a);  hind foot  Interparietal  s k u l l l e n g t h i n F i g . 11  and r o s t r a l l e n g t h i n F i g . 12  and  and F i g . 1 3 ( b ) .  F i g u r e s to the l e f t o f each histogram and bar diagram i n d i c a t e the number of animals b e i n g  compared.  L e t t e r s to the r i g h t of  each histogram and bar diagram i n d i c a t e the s p e c i e s , sex, age  of the animals used (L  g r a c i l i s , cf^male,  P. 1.  noveboracensis, M=P.  9 = f e m a l e , A «adult, S = s u b a d u l t ) .  p l o t t e d i n the bar diagrams are presented The was no  and m.  Data  i n the Appendix (Table I ) .  e f f e c t o f f o r m a l i n p r e s e r v a t i o n on l e n g t h of body p a r t s  i n v e s t i g a t e d by Lanko ( I 9 6 0 ) .  T h i s p r e l i m i n a r y study found  s i g n i f i c a n t d i f f e r e n c e s between measurements taken b e f o r e  and a f t e r immersion f o r 30 days i n 10$ the use  of preserved  formalin.  m a t e r i a l would not a f f e c t any  Therefore, comparisons  made with mice t h a t had been measured when f r e s h l y k i l l e d .  19Weight The  weights  Leucopus tends  of leucopus  and  m a n i c u l a t u s were  t o be h e a v i e r t h a n m a n i c u l a t u s .  compared.  Although  s p e c i m e n s were w e i g h e d m i n u s s t o m a c h c o n t e n t s , t h e i r o v e r l a p and Fig.  even exceed  2 ( a , b ) shows t h a t w e i g h t g i v e s no  leucopus  and m a n i c u l a t u s .  l e u c o p u s were n o t for  the weights  male and weights  The  weights  significantly  from  those of a d u l t  Examination  of  separation of  The  female  same was  maniculatus  A d u l t leucopus  more f r o m  weights  o f a d u l t male and  female  d i d not d i f f e r s i g n i f i c a n t l y . differed  clear  different.  o f a d u l t male and  female maniculatus.  maniculatus did  those o f maniculatus.  leucopus  and  true  and  adult  subadult  maniculatus  Weights of subadult  those of a d u l t leucopus  than  they  maniculatus.  Hind foot l e n g t h Hind f o o t Peromyscus. best  l e n g t h s have been used  McCarley  (1954)  c h a r a c t e r t o s e p a r a t e P.  Examination  o f F i g s . 3 and  s e p a r a t i n g P. A d u l t male and  1.  female  though not maniculatus.  male and  l e u c o p u s and P.  5(a)  shows t h a t and P.  m.  The  gossypinus.  gracilis i s limited. significantly  Male a d u l t m a n i c u l a t u s have a  significantly ranges  longer, hind foot  than  from  the h i n d f o o t l e n g t h s o f a d u l t  slightly adult  o t h e r s are not.  differ  maniculatus.  Some p a r t s ( e g . body l e n g t h ) o f s u b a d u l t m a n i c u l a t u s than those of the a d u l t ,  in  of hind foot length of subadult  f e m a l e m a n i c u l a t u s were t h e same and d i d n o t  significantly  the  i t s usefulness i n  l e u c o p u s do n o t d i f f e r  the l e n g t h of h i n d foot. longer,  f o u n d h i n d f o o t l e n g t h t o be  noveboracensis female  to separate species of  are s m a l l e r  F o r example,  subadult  Figure 2 ( a ) .  Histograms  o f weights o f P. 1.  noveboracensis and P. m. g r a c i l i s by sex and age. F i g u r e s to the l e f t o f each histogram i n d i c a t e the number o f animals b e i n g compared.  L e t t e r s to the r i g h t o f each  h i s t o g r a m i n d i c a t e the s p e c i e s and age of  the animals b e i n g compared.  L = P. 1. noveboracensis M = P. m.  gracilis  A = Adult S =  Subadult  ( T h i s legend w i l l be used i n a l l histograms)  35 •  33  •I  L.  I..  I 22  31 •  22  I  I  L- 9  M. d" M. £  Jl  •  M.  d  (A)  (A) (A)  (A)  (S)  CS)  M. 9  26  d  UJ CD  L. d*9 CA)  68  M. C/9  53  M. d"9  48  IO  Figure 2(a).  15  J  I  20  WEIGHT  I  (A)  (S) I  25  (GMS.)  30  Figure  2(b).  Bar diagrams o f weights o f P. 1.  noveboracensis and P. m. g r a c i l i s by sex and age. The t o t a l range o f measurements i s shown by a horizontal line,  the mean by a v e r t i c a l  l i n e , one standard d e v i a t i o n on each side o f the mean by an "open" r e c t a n g l e ,  and two standard  e r r o r s on each s i d e o f the mean by a " s o l i d " rectangle. Figures  t o the l e f t o f each b a r diagram i n d i c a t e  the number o f animals b e i n g compared.  Letters  to the r i g h t o f each b a r diagram i n d i c a t e the species  and age o f the animals b e i n g compared.  L = P. 1. noveboracensis M = P. m. g r a c i l i s A =  Adult  S = Subadult ( T h i s legend w i l l be used i n a l l b a r diagrams)  L.  3 3 »-  22  31  . M. d  »-  i-  , M.  3  n  22 •-  , M.  M.  2 6 »"  14  16  Figure  (8  2(b).  9  9  9  (A)  (A)  (A)  (S)  (S)  20  W E I G H T  22  24  (GMS.)  26  28  30  Figure  3.  P.  1.  by  sex  Histograms of hind  foot lengths  n o v e b o r a c e n s i s and  P.  and  age.  m.  of  gracilis  35  (A* 3 3  (A)  •  22 • • I I • • • • M. C / (A) 31  o-  •  o2 2  •  •  iiki.L I  6 8  M .  I  (A)  O  M .  L.  O*  ( S )  (A)  0*0^  25J  20-  15-  ioH H I W  IOI  I 18  Figure  3.  I  I  I 19  HIND  I 2 0  FOOT  I  .1.  '  21  LENGTH  I  M .  cfo  _l  2 2  l_  2 3  ( M M . )  20. hind  f o o t l e n g t h s a r e t h e same as t h o s e o f t h e a d u l t s .  the e a r l y  attainment of adult  size  i s necessary  Perhaps  f o r adeptness  in  climbing or running. Ear length E a r l e n g t h h a s b e e n u s e d a s an a i d t o s e p a r a t i n g t h e s e species.  Cameron  (1956) p o i n t e d o u t t h a t l e u c o p u s h a s s h o r t e r  ears than manlculatus.  My  data  ( F i g s . 4 and 5 ( h ) ) , show  l e u c o p u s d o e s h a v e s h o r t e r e a r s and t h a t shorter  than those o f manlculatus.  male and f e m a l e other. and  adult maniculatus.  hind was  significantly  N e i t h e r do t h o s e o f a d u l t male and f e m a l e maniculatus.  a d u l t m a n i c u l a t u s do n o t d i f f e r Hence  Perhaps the e a r l y  full  significantly  each  maniculatus  from  those o f  ear l e n g t h i s attained  attainment  early i n  of adult ear length,  significantly different  most n e a r l y s e p a r a t e d t h e s e  from  E a r l e n g t h s o f sub-  f o o t l e n g t h h a s some a d a p t i v e s i g n i f i c a n c e . t h e most h i g h l y  significantly  The e a r l e n g t h s o f a d u l t  l e u c o p u s do n o t d i f f e r  s u b a d u l t male and f e m a l e  life.  they are  that  like  Ear length  c h a r a c t e r found  and  s p e c i e s o f mice.  Body l e n g t h Body l e n g t h s were  compared t o a s s e s s t h e i r u s e f u l n e s s i n  s e p a r a t i n g t h e two s p e c i e s . body l e n g t h t h a n m a n l c u l a t u s . f o u n d b e t w e e n male and f e m a l e of either  species.  significantly  Leucopus tends No  t o have a s h o r t e r  significant differences  a d u l t s and male and f e m a l e  Body l e n g t h s o f s u b a d u l t m a n i c u l a t u s  different  from  those  of adult maniculatus.  a d u l t body l e n g t h i s n o t a t t a i n e d b y s u b a d u l t a n i m a l s ,  were subadults were Since  i t may  F i g u r e 4.  Histograms o f ear l e n g t h s of P.  noveboracensis and P. m. and  age.  g r a c i l i s by  1. sex  3 5  3 3  (A)  I • mM H  I 2 2  Ik •  li  •  L.  •  Q  •  (A)  •  •  •  •  •  M .  C  (A)  31  M  •  2 2  •  M  • M  II  M  11111  •  •  " M .  M.  6 8  I Ik J I H l  I L.  I H J k  d*<J  M .  d C)  C;  (A)  (S)  (S)  (A)  1..  M .  IOI  (A.S)  I  I  15  pisure 4 .  i  i  16  i  17  EAR  i  i  18  LENGTH  t  i  19  ( M M . )  i  i  2 0  i  i  21  Figure  5(a).  o f P. by  Figure  Bar diagrams of h i n d  foot  1. n o v e b o r a c e n s i s and P. m.  s e x and  5(b).  lengths gracilis  age.  Bar diagrams o f ear lengths  P.  1. n o v e b o r a c e n s i s and P. m.  by  s e x and  age.  of  gracilis  -i L .  35  33  L.  »-  Cf  9  -i M .  22  (A")  (A) O"  (A')  31  Figure 5(a).  i  •  26  »  20  19  18  35  22  1  HIND  I  21  FOOT  . L.  Cf  23  22  LENGTH  (MM.)  (A^  .  • M.  I 16  Figure 5(b).  I 17  EAR  I 18  I 19  LENGTH  Cf  (A)  (A) (S)  9  (S)  I 20  (MM.)  9  < M . Cf  < M. I 15  M.  I 21  t 22  Figure 6 ( a ) .  H i s t o g r a m s o f body l e n g t h s o f P . 1.  n o v e b o r a c e n s i s and age.  P . m.  g r a c i l i s by  sex  and  M.  22  •  •  •  k  .  26  M.  d  (S)  M.  9  (S)  66  C/9  L.  I  53  M.  9  (A)  (A)  (5*9  IO  5 M.  48  CH  J — J  70  Figure 6(a).  I 80  BODY  I  C?9 I  90  (S)  I IOO  LENGTH  IIO  (MM.)  (A)  Figure 6(b).  Bar  P.  1.  by  s e x and  d i a g r a m s o f body l e n g t h s o f  noveboracensis age.  and P.  m.  gracilis  35  i-  33  , L9  3  H  M . C5  22  31  CA)  H  H  22  3H  H  26  72  CA)  76  Figure  h  80  6(b).  M.  cf H  84  B O D Y  88  M.  M.  9  CS)  9 92  L E N G T H  (S) 96  (MM.)  IOO  102  CA)  21. not have as much adaptive ear l e n g t h .  s i g n i f i c a n c e as h i n d f o o t l e n g t h or  Although body l e n g t h s  o f a d u l t leucopus and a d u l t  maniculatus were found to be s i g n i f i c a n t l y d i f f e r e n t , the wide range o f measurement i n any one group ( f o r example, a d u l t female maniculatus) make i t u s e l e s s i n the s e p a r a t i o n o f these s p e c i e s . Tail  length I n v e s t i g a t o r s have o f t e n used t a i l l e n g t h as an a i d t o  s e p a r a t i n g these two s p e c i e s .  B u r t and G-rossenheider ( 1 9 5 2 )  admit  they are d i f f i c u l t t o t e l l apart, b u t g i v e the f o l l o w i n g as a f a i r l y good means o f s e p a r a t i o n :  P. 1 .  n.:  t a i l length  shorter  than 3 3 / 5 Inches ( 9 * 1 5 cms.);  P. m. £.:  than 3 3 / 5 inches  T h i s method, however, does n o t  completely  ( 9 . 1 5 cms.).  separate  t a i l length  the mice used i n t h i s study.  Of 1 2 0  noveborac ens1s, 9 have t a i l s l o n g e r than 9 . 1 5 cms.; g r a c i l i s (Algonquin and 9.15  Park),  o f 157  s h o r t e r than 9 * 1 5 cms.;  of 1 0 0 g r a c i l i s (R.O.M.Z. l o a n ) , 78 have t a i l s s h o r t e r than cms. Cameron ( 1 9 5 6 )  says t h a t the t a i l l e n g t h o f leucopus i s  u s u a l l y l e s s than one-half has  136 have t a i l s  longer  the t o t a l l e n g t h and t h a t manlculatus  a t a i l l e n g t h t h a t i s u s u a l l y g r e a t e r than one-half the  t o t a l length.  T h i s method does not completely  specimens used h e r e . l e s s than one-half (Algonquin  Park),  separate the  Of 1 2 0 leucopus, 119 have a t a i l  the t o t a l l e n g t h ;  length  o f 157 manlculatus  69 have a t a i l l e n g t h g r e a t e r than  one-half  the t o t a l l e n g t h ; and o f 1 0 0 maniculatus (R.O.M.Z. l o a n ) , 2 7 have a t a i l l e n g t h g r e a t e r than one-half  the t o t a l l e n g t h .  of the above methods i s s u b s t a n t i a t e d by my data.  Neither  Figure  7(a).  Histograms of t a i l  n o v e b o r a c e n s i s and age.  P.  m.  l e n g t h s o f P.  g r a c i l i s by  sex  1. and  35  33  L.  . -.1  L.  O"  J  (A)  (A)  M  22  o-  d  31  -j < o-  (A) M.  22  •  26  M/ Cf  M. 9  9  (A)  (S)  IS)  U. O or u  a 2 i  5  68  O  L. 0 * 9  (A)  5 M. 0 * 9 (A)  53  Ol  48  60  Figure 7 ( a ) .  •  M.  70  TAIL  eo  90  LENGTH  IOO  (MM.)  69  (S)  no  Figure 7(b).  Bar diagrams of t a l l  P.  1.  n o v e b o r a c e n s i s and P.  by  s e x and  age.  m.  lengths of gracilis  35  33  -i  h  H  t-  22  31  22  65  Figure  (A)  L. p  (A)  M . (3  »-  (A)  9  M. (A)  H  *M.  h  26  60  L.CJ  70  7(b).  M.  *•  75  T A I L  80  85  L E N G T H  90  (MM.)  9  (S)  (S)  95  IOO  22. Comparisons o f t a i l shown i n F i g . 7 ( a , b ) .  lengths of leucopus  No  significant differences  were f o u n d "between t h e s e x e s o f e a c h age Tail  tended  subadult maniculatus.  t o have s h o r t e r t a i l s  t h e r e was  Interparietal  species.  ments f o r b o t h  widths  widths  Interparietal  of leucopus  (Figs.  These authors found w i t h age  data (Figs.  were  Graphic presentation of  8 and 1 0 ( a ) ) , shows t h a t t h e m e a s u r e for this character  9 and  l e n g t h h a s b e e n u s e d by H o f f m e i s t e r  f o r comparisons that and  L e u c o p u s has  than manlculatus.  (1951)»  o f s p e c i e s o f Peromyscus.  the l e n g t h o f the i n t e r p a r i e t a l  changes  that  sexes.  i t does not d i f f e r between  1 0 ( b ) ) show t h a t i t d o e s n o t d i f f e r  c a n t l y between the sexes  length  and m a n i c u l a t u s  length  interparietal  very l i t t l e  and  value.  Cockrum ( 1 9 5 4 - ) ,  leucopus.  Although  t o h a v e some v a l u e i n t h e  s p e c i e s o v e r l a p too g r e a t l y  o f taxonomic  The  manlculatus  width  examined f o r p o s s i b l e d i f f e r e n c e s . interparietal  from  two.  Interparietal  t o be  Subadult  than a d u l t maniculatus.  c h a r a c t e r appears  s e p a r a t i o n of the  for  of each  are  length  g r e a t o v e r l a p i n l e n g t h o f t a i l between leucopus  manlculatus. t h i s  My  group  in tail  l e n g t h s o f a d u l t l e u c o p u s were s i g n i f i c a n t l y d i f f e r e n t  t h o s e o f a d u l t and  and  and m a n i c u l a t u s  and  ages o f m a n i c u l a t u s  a significantly shorter Interparietal  s e p a r a t i n g most a n i m a l s o f t h e two  and  the  signifisexes  of  interparietal  l e n g t h i s a good c h a r a c t e r species.  F i g u r e 8. P.  1.  Histograms  of i n t e r p a r i e t a l  noveboracensis  s e x and  age.  and  P.  m.  widths  gracilis  NUMBER O  cn  O 1  fO  t  n I  OF O  c *  CO  INDIVIDUALS n •  O  c i  n  O a  c .  n .  to  HC CD CO  r r  2  m  o»  >  CO  2  +o  2  > >  >  m  > a  09  H  2 2  3 +o >  F i g u r e 9. P.  1.  Histograms  of i n t e r p a r i e t a l  noveboracensis  s e x and  age.  and P.  m.  lengths of  gracilis  by-  5  O  35  O-  33  L.  L.  9  522  O  31 5  <  9  °  >  5  (A)  (A)  22  O"  M.  ..Liu  5  to  L  O*  .A  M.  9  d  M .  (A)  ( A )  ( S )  Q Z  26  9  M.  ( S )  u. O  cc hi CQ  Iz  5 68  L.  d"9  (A)  O  15  IO  5  lOI  O 6.5  Figure  9  .  7.5  M.  8.5  INTERPARIETAL  0*9  9.5  IQ5  LENGTH  (MM.)  C A , S ;  Figure of  10(a). P.  of  10(b). P.  diagrams  of i n t e r p a r i e t a l  1. n o v e b o r a c e n s i s and  s e x and  Figure  Bar  1.  s e x and  P. m.  widths  gracilis  by-  age.  Bar diagrams noveboracensis age.  of Interparietal and P.  m.  lengths  gracilis  by  33  f 22  H  -i M . C f  CA"J  (S)  31 »•  F i g . io(a).  22  >-  •i M . Cf  26  y  M. 9 ( S )  INTERPARIETAL  35  H  33  \-  -i L . Cf  CA)  - L. 9 ( A ) -• M . Cf CA)  22  M. 9 C A )  31 22  »-  M. Cf  CS)  M. 9 C S )  26  8  Fig. io(b).  WIDTH ( M M . )  INTERPARIETAL  IO  LENGTH  CMM.)  23. Skull length The  lengths of skulls  examined f o r p o s s i b l e  o f leucopus  and m a n i c u l a t u s  specific differences  Leucopus has a s i g n i f i c a n t l y  longer s k u l l  (Figs. than  11 and 1 3 ( a ) ) .  maniculatus,  a l t h o u g h t h e r e was g r e a t o v e r l a p o f measurement. maniculatus  tended  t o have s h o r t e r s k u l l s  No s i g n i f i c a n t d i f f e r e n c e s the sexes  of adult  maniculatus.  i n skull  leucopus,  Subadult  than adult  maniculatus.  l e n g t h were f o u n d  adult maniculatus,  Skull length i s useful  were  between  and s u b a d u l t  i n the s e p a r a t i o n o f the mice.  Rostral length C o m p a r i s o n s were made o f r o s t r a l maniculatus  (Figs.  longer rostrum  12 and 1 3 ( b ) ) .  than maniculatus.  l e n g t h s o f l e u c o p u s and  Leucopus tended  Subadult maniculatus had a  shorter rostrum than adult maniculatus. ences i n r o 3 t r a l leucopus,  No s i g n i f i c a n t  differ-  l e n g t h were f o u n d b e t w e e n t h e s e x e s o f a d u l t  adult maniculatus  and s u b a d u l t m a n i c u l a t u s .  a g r e a t e r d i f f e r e n c e between t h e s p e c i e s i n s k u l l in rostral  to have a  length.  This character i s useful  T h e r e was  l e n g t h than  as an a i d t o t h e  s e p a r a t i o n o f the mice.  Baculum l e n g t h The  l e n g t h o f t h e b a c u l u m h a s b e e n u s e d b y many a u t h o r s t o  d i s t i n g u i s h between genera  and s p e c i e s o f P e r o m y s c u s .  Blair  (194-2) f o u n d no marked q u a l i t a t i v e d i f f e r e n c e s b e t w e e n t h e b a c u l a of the maniculatus reported  and l e u c o p u s  s p e c i e s groups.  H o o p e r (1958)  t h e mean v a l u e f o r b a c u l u m l e n g t h i n l e u c o p u s  9.0 mms. and i n m a n i c u l a t u s , 8.3 mms.  t o be  B u r t (i960) q u e s t i o n e d t h e  F i g u r e 11.  Histograms of s k u l l  n o v e b o r a c e n s i s and P. and  age.  m.  l e n g t h s o f P.  g r a c i l i s by  sex  1.  35 a  33  •  22  I  •I  31 • •  22 •  26  (A)  •  H  ii  i I  I II  J L H J  48 • *  •  M . d"  L.  •  M.  9  MM§  25  11.  (A)  (S)  I •  •  24  (A)  • M. 9  •  •LMIJJ 1  9  (A)  m  • I iJ JL»^»M I MM  1  Figure  •  M. d" (S)  b — . Mm mi mm  68  53  J  J  » M  I • l a J h J• . • I •  U  a J k JL  •  SKULL  M.  1  1  26  d9 '  L.  <^9  (A)  m M. C J 9  (S)  1  27  LENGTH  '  (MM.)  28  '  29  F i g u r e 12.  Histograms of r o s t r a l  noveboracensis age.  and P.  m.  l e n g t h s o f P.  g r a c i l i s by  sex  1.  and  Figure 13(a).  Bar diagrams of s k u l l l e n g t h s o f  P. 1. noveboracensis  and P. m. g r a c i l i s by-  sex and age.  F i g u r e 13(b).  Bar diagrams o f r o s t r a l l e n g t h s o f  P. 1. noveboracensis sex and age.  and P. m. g r a c i l i s by  L.  35 33  M. Cf  H M. Cf  25  SKULL  35  H  33  '  (S) ,1 27  26  28  LENGTH  CMM.)  L.  7.4  •  78  F i g . 13(b).  M. Cf  (A)  H  ROSTRAL  8.6  (A)  (S)  C  8.2  cf  (A)  - M . O  31  26  9  M. Cf  C  y-  29  L.  i  22  (A)  (S)  M. 9  22  9  -t M.  22 •-  (A)  (A')  31  Figure 13(a).  (A')  - L. 9  '  22 -  24  Cf  9.0  LENGTH  M. 9  9.4  (MM.)  CS)  9.8  (A)  24. taxonomic value o f the baculum i n Peromyscus except f o r the s e p a r a t i o n o f subgenera.  He found the mean value f o r the l e n g t h  o f the baculum o f leucopus to be 9*5 mms. and  o f manlculatus to be 9 . 3 mms.  (range,  (range,  8.1-10.9 mms.),  8 . 7 - 9 » 8 mms.).  v a l u e s a r e so s i m i l a r t h a t t h e i r taxonomic u s e f u l n e s s  These  i s negligible.  My data show t h a t the l e n g t h s o f the b a c u l a o f a d u l t leucopus and maniculatus l i e i n the same s i z e range. 8.9 mms.  f o r 39 i n d i v i d u a l s (range,  averaged 8.1 mms. Burt's  Leucopus averaged  5 . 3 - 1 1 . 3 mms.), and maniculatus  f o r 19 i n d i v i d u a l s (range,  6.0-10.0 mms.).  statement (I960, P. 5 4 ) , t h a t (the baculum l e n g t h o f ) ,  "P. m. g r a c i l i s , leucopus."  f o r i n s t a n c e , I s w e l l w i t h i n the s i z e range o f  i s supported  by my data.  Baculum l e n g t h i s not a  good c h a r a c t e r f o r the s e p a r a t i o n o f these  species.  In summary, g r a p h i c comparison Bhows t h a t e a r l e n g t h and i n t e r p a r i e t a l l e n g t h a r e the b e s t c h a r a c t e r s to use i n the s e p a r a t i o n o f P. 1. noveboracensis and P. m. g r a c i l i s .  Neither  of these measurements d i f f e r s i g n i f i c a n t l y between subadult and a d u l t animals o f the one s p e c i e s .  T a i l l e n g t h i s the next b e s t  c h a r a c t e r , f o l l o w e d by s k u l l l e n g t h , r o s t r a l l e n g t h , and h i n d foot length.  Body l e n g t h and i n t e r p a r i e t a l w i d t h o v e r l a p too  g r e a t l y t o be o f any v a l u e . The S e p a r a t i o n o f leucopus and manlculatus u s i n g  Morphological  Characters Comparison o f p a r t s o f mice showed t h a t some measurements d i f f e r e d more than others between s p e c i e s .  To s e l e c t the  measurements most u s e f u l i n s e p a r a t i n g the s p e c i e s , " t " v a l u e s  25.  were c a l c u l a t e d to show the degree of d i f f e r e n c e between measurements.  No s i g n i f i c a n t d i f f e r e n c e s , at the 5$ l e v e l , were found  between the sexes of adult leucopus,  a d u l t maniculatus,  a d u l t maniculatus f o r a l l measurements compared.  The  and subparts  compared between species, and t h e i r " t " values, are arranged i n a decreasing order of d i f f e r e n c e i n the f o l l o w i n g t a b l e : Table 1.  Degree of d i f f e r e n c e between body p a r t s of leucopus and maniculatus as shown by " t " values. -P a r t s Compared  " t " Value  Ear l e n g t h T a i l length I n t e r p a r i e t a l length Body l e n g t h S k u l l length Rostral length The  11  51.436 49-352 19.308 10.987 7-326 3.722  1" values i n Table 1 are a l l s i g n i f i c a n t at the  5$  l e v e l and represent comparisons of body p a r t s of adults of both sexes of leucopus w i t h those of a d u l t s of both sexes of maniculatus.  A complete summary of a l l " t " t e s t s c a l c u l a t e d i s  given i n the Appendix (Table I I ) . Table 1 I n d i c a t e s that leucopus and maniculatus are s i g n i f i c a n t l y d i f f e r e n t i n a l l comparisons f o r which " t " was the forms was values.  calculated.  The greatest divergence  between  i n those characters t h a t possessed the l a r g e s t " t "  Length of ear possesses the l a r g e s t " t " value but i s  not s a t i s f a c t o r y , when used alone, to separate leucopus and maniculatus because the ranges of measurement f o r the two overlap.  species  Hence, while ear l e n g t h separates the species  s t a t i s t i c a l l y , i n p r a c t i c e , i t does not separate a l l the i n d i v i duals.  The above t a b l e a l s o shows that these mice d i f f e r  26. s i g n i f i c a n t l y i n a number of c h a r a c t e r i s t i c s .  I f these  c h a r a c t e r i s t i c s are c o n s i d e r e d a t the same time, a t o t a l d i f f e r e n c e may  be seen between the s p e c i e s .  T h i s i s done i n  F i g . l 4 ( a , b ) , the measurement polygon. The measurement polygon expresses a number of c h a r a c t e r i s t i c s simultaneously.  Each polygon i s a " p i c t u r e " of an  average  animal, based on the mean v a l u e s of 8 measurements.  Examination  o f F i g . 14(b) r e v e a l s t h a t t h e r e i s c o n s i d e r a b l y more s i m i l a r i t y between polygons A and B and C and D than t h e r e i s between A and C or D or between B and C or D.  The polygons c l e a r l y show that  leucopus has a s h o r t e r average i n t e r p a r i e t a l width, ear l e n g t h , body l e n g t h , t a i l l e n g t h , i n t e r p a r i e t a l l e n g t h , and h i n d f o o t l e n g t h , w h i l e i t s average  s k u l l l e n g t h and r o s t r a l l e n g t h are  l o n g e r than comparable average v a l u e s f o r maniculatus. The Character Index f o r the S e p a r a t i o n of leucopus and  manlculatus  While polygons express t o t a l average f e a t u r e s , they are not a convenient way  to separate the s p e c i e s .  A l s o , I have no  measure of the goodness of f i t of a s p e c i e s to a polygon.  At  the moment, a l l that can be s a i d i s t h a t a mouse i s e i t h e r a leucopus or a manlculatus depending upon which polygon I t resembles  most.  Another way  of making use of a number o f t e n d e n c i e s i n each  s p e c i e s to s e p a r a t e them i s by the use o f the c h a r a c t e r index. The  term " c h a r a c t e r index" was proposed by Hubbs and  (1929), who  Whitlock  found the need f o r a simple mathematical e x p r e s s i o n  of several d i f f e r e n t i a l features.  These i n d i c e s have been used  s u c c e s s f u l l y i n f i s h s y s t e m a t l c s f o r the s e p a r a t i o n o f c l o s e l y  re 14(a).  Key to measurement polygons o f P. 1.  noveboracensis and P. m. g r a c i l i s . measurement ( i n mms.) E. L . =  ear l e n g t h  B. L.=  body l e n g t h  T. L . =  tail  One average  i s p l o t t e d on each r a d i u s .  length  SK. Lr= s k u l l l e n g t h R. L. = r o s t r a l l e n g t h I. L. =  interparietal  length  H. F. L.= h i n d f o o t l e n g t h I. ¥. = i n t e r p a r i e t a l ( T h i s legend  width  a p p l i e s to F i g u r e 14(b))  E.L  R.L. Figure 14(a).  M E A S U R E M E N T  K E Y  P O L Y G O N  Figure 14(b).  Measurement polygons o f P. 1.  noveboracensis and P. m. g r a c i l i s .  Average  measurements f o r each group o f animals are p l o t t e d on each r a d i u s . A.= a d u l t male P. 1. noveboracensis B.= a d u l t female P. 1. noveboracensis 0.= a d u l t male P. m.  gracilis  D^=" a d u l t female P. m.  gracilis  A.  PL.N.  C  RM.G.  cf  ( A )  B.  PL.N. 9  cf  C A )  D.  PM.G.  F i g u r e 14(b).  9  C A )  C A )  Measurement polygons o f P. 1. n. and P. m. g.  27. a l l i e d forms by S c h u l t z ( 1 9 3 7 ) , Hubbs and Kuronuma ( 1 9 4 1 ) , ( 1 9 3 5 ) used simple  and Hubbs e t a l ( 1 9 4 3 ) .  that the index,  Davenport  i n d i c e s to compare p r o p o r t i o n s of  Moreau and Southern ( 1 9 5 7 ) used the index, show g e o g r a p h i c a l  (1934),  S c h u l t z and Welander  variation i n shrikes.  animals.  tail/wing X 100,  Cameron ( 1 9 5 8 )  mean s k u l l height/mean r o s t r a l l e n g t h ,  to  found separated  b l a c k bears o f Newfoundland from those of the mainland. The  pros and  of s p e c i e s and  cons o f u s i n g c h a r a c t e r i n d i c e s f o r comparisons  subspecies  of animals are d i s c u s s e d by  ( 1 9 3 9 ) and Hubbs e t a l ( 1 9 4 3 ) .  Hubbs e t a l (P. 5 ) ,  Ginsburg  strongly  defend the use of c h a r a c t e r i n d i c e s by s a y i n g t h a t , "no  arithmetic  combination o f c h a r a c t e r s , however, can b r i n g out d i f f e r e n c e s which do not e x i s t ,  so l o n g as we  apply the i d e n t i c a l  index formula  t o b o t h types b e i n g compared.".  s t a t e (P. 6 ) ,  t h a t , "we  such v a l u e s as standard may  have been informed  They f u r t h e r  by  d e v i a t i o n , standard  character-  s t a t i s t i c i a n s that  e r r o r , and  the  like  l e g i t i m a t e l y be computed f o r an array of the i n d i c e s , p r o v i d e d  t h a t each o f the combined c h a r a c t e r s p r e s e n t s approximately The  normal frequency  in itself  an  distribution.".  main c r i t i c i s m a g a i n s t the use of c h a r a c t e r i n d i c e s i s  concerned w i t h the d e t e r m i n a t i o n (Ginsburg,  1939).  of s p e c i e s and  T h i s author p o i n t s out t h a t the degree of  d i f f e r e n c e between i n d i v i d u a l s of two c h a r a c t e r s s e l e c t e d and i n d i c e s should not, populations  subspecies  p o p u l a t i o n s depends on  on the manipulations  performed.  These  t h e r e f o r e , be used to determine whether  are subspecies  of one  s p e c i e s or are separate  the  two  species.  Ginsburg*s c r i t i c i s m s are v a l i d and worthy o f c o n s i d e r a t i o n . However, they do not apply to the problem I am  t r y i n g to s o l v e  28. with character i n d i c e s .  The animals used i n t h i s study are  known to be separate s p e c i e s on the b a s i s of t h e i r b r e e d i n g b i o l o g y and I wish o n l y t o be able to separate them by some q u a n t i t a t i v e r a t h e r than q u a l i t a t i v e method. Of the c h a r a c t e r s compared g r a p h i c a l l y and ear  statistically,  l e n g t h , i n t e r p a r i e t a l l e n g t h , and t a i l l e n g t h were found to  be the measurements d i f f e r i n g most between the two  species.  These c h a r a c t e r s were s h o r t e r i n leucopus than i n maniculatus. I f these t h r e e measurements were m u l t i p l i e d t o g e t h e r f o r each i n d i v i d u a l of each s p e c i e s , the r e s u l t a n t i n d i c e s f o r leucopus would be s m a l l e r than those f o r maniculatus. the d i f f e r e n c e between the two by another measurement t h a t was  To f u r t h e r i n c r e a s e  s p e c i e s , d i v i s i o n o f these i n d i c e s l o n g e r i n leucopus than i n  maniculatus would g i v e even s m a l l e r i n d i c e s f o r leucopus comparably  l a r g e r ones f o r maniculatus.  Both r o s t r a l l e n g t h  and s k u l l l e n g t h f i l l e d t h i s requirement and each was •part of a c h a r a c t e r index.  and  used as  The r e s u l t i n g i n d i c e s were:  I.  ear l e n g t h X t a l l l e n g t h X I n t e r p a r i e t a l l e n g t h s k u l l length  II•  ear l e n g t h X t a l l l e n g t h X i n t e r p a r i e t a l l e n g t h r o s t r a l length  Although each o f these i n d i c e s was  found to completely separate  leucopus and maniculatus, b o t h are g i v e n here because  specimens  c o l l e c t e d by snap t r a p p i n g are o f t e n damaged and one o f the necessary measurements, such as s k u l l l e n g t h , might be u n o b t a i n able.  In t h i s example, index I I c o u l d be used r a t h e r than  index I .  29. T e s t s of I n d i c e s I f an index i s to be o f value, i t should be e a s i l y and  separate a l l ages and  applied.  The  sexes of forms to which i t may  be  I n d i c e s were t e s t e d by c a l c u l a t i n g v a l u e s f o r a l l  animals used i n the study.  Histograms and bar diagrams o f the  v a l u e s f o r each s p e c i e s , sex, and age group were p l o t t e d 15,  calculated  16, 1 7 ( a , b ) ) .  (Figsi  A l s o i n c l u d e d i n the t e s t s (but not graphed)  were d a t a f o r mice from the Royal O n t a r i o Museum of Zoology from Osgood ( 1 9 0 9 , pp. 260-61, 263-64).  and  A summary o f data p l o t t e d  i n bar diagrams f o r each s p e c i e s , sex, and age group i s g i v e n i n the Appendix (Table I I I ) .  The  i n d i c e s f o r each s p e c i e s were  compared by " t " t e s t and found 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$  l e v e l f o r mice from a l l areas. I t has a l r e a d y been shown t h a t sex had no e f f e c t on measurements.  Although  15,  and 17(a,b) show t h a t age does not a f f e c t the v a l u e s of  16,  subadult mice tend to have s m a l l e r p a r t s , F i g s .  the i n d i c e s c a l c u l a t e d f o r a l l age groups examined. v a l u e s of the i n d i c e s are not a f f e c t e d by sex or age, each s p e c i e s can be presented i n T a b l e T a b l e 2.  Species leucopus maniculatus maniculatus leucopus maniculatus  combined.  Since  the  data f o r  Oombined d a t a o f the i n d i c e s are  2.  Means and ranges o f combined data of the c h a r a c t e r i n d i c e s o f leucopus and maniculatus.  N 92  101 13  10 10  Source L. Opin., P e r t h Rd. A l g . Park R.O.M.Z. Osgood Osgood  Index I Mean Range 3.41 2.57-4.09  Index I I Range Mean 10.45 7. 39-:12.50  5.62 6.57 3.49 5.75  17.16  4.26-7.14 5.39-7.90 5.69-5.83  19.85  12 .83--21.93 16 .24.-22.77  Figure  15.  Histograms  o f the index ( I ) ,  E.L.  X T . L . X I.L.« f o r P. 1. SK.L. and P. m. g r a c i l i s by s e x and  E. L . =  ear  T. L . =  tall  I. L. =  interparietal  SK.  skull  L.=  length length  length  length  noveboracensis age.  NUMBER  OF  INDIVIDUALS  Figure  16.  E.L. and  Histograms  X T . L . X I . L . , f o r P. 1. R.L. P. m. g r a c i l i s b y s e x and  E . L *= e a r T.  o f the index ( I I ) ,  L*= t a i l  length length  I . L.F I n t e r p a r i e t a l R.  L.= r o s t r a l  length  length  noveboracensis age.  Figure 17(a).  Bar diagrams o f the index ( I ) ,  E.L. X T.L. X I.L., f o r P. 1. noveboracensis SK.L. and P. m. g r a c i l i s by sex and age. (Legend as i n F i g u r e 15)  F i g u r e 17(b).  Bar diagrams o f the index ( I I ) ,  E.L. X T.L. X I.L., f o r P. 1. noveboracensis R.L. and P. m. g r a c i l i s by sex and age. (Legend as i n F i g u r e 16)  46 46  L.  •-  0*  L. 9  *~-  22 31 22  (A)  Fig.  46 46  , .  17(a).  M.  i  3  i  5.0  (A)  ( S)  (S)  _|  6.0  9  L  7.0  1  8.0  — i L. O* CA)  — . L. 9 CA)  '  2 2 31 22  i  -t M . Cf ( A )  c  '  . M . 9 CA)  •=  ^ M . Cf CS)  •26  I  I  65  9  CA)  E.LX T L X I.L.  K i r > r r v  i-  M.  M . Cf -  4.0  , INDEX-  M- Cf 1  i—  •  3.0  —»  *-  '—-  26  2D  CA)  85  IQ5  I  125  iMHCY F i g . 17(b).  INDEX*  -I  I  I  I  145  E.LX  I  165  TLX  p |_  I  I.L  M. 9  Cs)  I 8 [ 5 2 0 5  225  30. T a b l e 2 shows t h a t means and ranges of c h a r a c t e r i n d i c e s c a l c u l a t e d f o r leucopus d i f f e r from those c a l c u l a t e d f o r manlculatus.  11  1"  t e s t s and bar diagrams  ( F i g . 17(a,b) show t h a t  a l l leucopus i n d i c e s d i f f e r s i g n i f i c a n t l y from a l l manlculatus indices.  Hence, the c h a r a c t e r i n d i c e s completely separate  leucopus and manlculatus of a l l ages and sexes t h a t were examined. A mouse w i t h an index I of 2 . 5 7 - 4 . 0 9 i s leucopus and one w i t h an index I of 4 . 2 6 - 7 - 9 0 i s maniculatus.  A mouse w i t h an index I I  o f 7 . 3 9 - 1 2 . 5 0 i s leucopus and one w i t h an index I I of 1 2 . 8 3 - 2 2 . 7 7 i s manlculatus. Although Osgood (1909) measured ear l e n g t h s from d r i e d specimens,  the v a l u e s f o r i n d i c e s c a l c u l a t e d from h i s d a t a  w i t h i n the ranges of each s p e c i e s index d e r i v e d from my The  fall  data.  shrinkage of pinnae i s a p p a r e n t l y not s u f f i c i e n t to g r e a t l y  a l t e r the v a l u e o f the index. T e s t o f the I n d i c e s on Mice From an Area of Sympatrv The i n d i c e s worked to separate mice from areas and  samples  t h a t were chosen to r e p r e s e n t "good" leucopus and manlculatus. I f the mice are not good s p e c i e s , they might b l e n d i n areas of sympatry.  A l s o , i t i s i n areas of o v e r l a p of range  that  geography i s o f no a s s i s t a n c e i n the i d e n t i f i c a t i o n of a  specimen.  Thus, the f i n a l and perhaps most c r i t i c a l t e s t of the i n d i c e s was  to apply them t o mice caught i n the a r e a of sympatry  of  leucopus and maniculatus i n O n t a r i o . Twelve mice from areas o f sympatry  were sent t o me by Dr.  D. A. Smith o f G a r l e t o n U n i v e r s i t y , Ottawa.  Almost a l l were  captured two and o n e - h a l f m i l e s n o r t h of Cloyne,  Frontenac  31. County.  Two were captured f i v e miles northwest of Calabogie,  Renfrew County.  C a l c u l a t i o n of i n d i c e s I and I I f o r these mice  completely separated them i n t o two groups.  Four specimens were  c l e a r l y r e f e r a b l e to leucopus (index I : range 3.35-3.93; index I I :  range, 10.56-12.38) and 8 specimens were c l e a r l y  r e f e r a b l e to manlculatus (index I : 4.39-6.01; 18.61).  index I I : 13.36-  This t e s t again i n d i c a t e s that the Indices are v a l i d  methods f o r the separation of P. 1. noveboracensis and P. m. g r a c i l i s i n southeastern Ontario. Summary of the Indices Indices have been shown to be v a l i d t o o l s f o r separating c l o s e l y r e l a t e d , d i f f i c u l t to separate species.  Indices were  formulated t h a t completely separate P. 1. noveboracensis and P. m. g r a c i l i s on a q u a n t i t a t i v e b a s i s .  These i n d i c e s are:  I.  ear l e n g t h X t a l l l e n g t h X I n t e r p a r i e t a l l e n g t h s k u l l length  II.  ear l e n g t h X t a l l l e n g t h X I n t e r p a r i e t a l l e n g t h r o s t r a l length  These i n d i c e s completely separate the two species regardless of  sex and age.  For index I , values ranging from 2.57 to 4.09  are i n d i c a t i v e of leucopus, while values ranging from 4.26 t o 7.90are i n d i c a t i v e of manlculatus.  For index I I , values ranging from  7.39 to 12.50 are i n d i c a t i v e of leucopus, while values ranging from 12.83 to 22.77 are i n d i c a t i v e of manlculatus. The i n d i c e s are simple and easy to use q u a n t i t a t i v e methods for  the separation of P. 1. noveboracens1s and P. m. g r a c i l i s .  The i n d i c e s separated noveboracensis and g r a c i l i s from areas of sympatry and areas of a l l o p a t r y .  32. Results of Crossbreeding Crossbreeding experiments  b e t w e e n P. l e u c o p u s  m a n i c u l a t u s were a t t e m p t e d  i n the l a b o r a t o r y .  experiments  i n the Appendix  a r e summarized  o f t h e 17 l i v e  leucopus received  l o c a l maniculatus  R e s u l t s of the  (Table IV).  i n 1961  were p a i r e d  Columbia).  of leucopus produced  a total  Two c o n t r o l  of three l i t t e r s .  o f two l i t t e r s  Two m i x e d p a i r s  ( P . m. o r e a s f a i l e d  ( P . m. g.C?  produced  ), ( P . m.  P. 1. n . c f ) o f m i c e were p l a c e d i n s e p a r a t e c a g e s  X  o n F e b r u a r y 17,  The m i c e h a v e b e e n i n b r e e d i n g c o n d i t i o n b u t h a v e n o t  littered.  One p a i r  o f leucopus  were p l a c e d i n s e p a r a t e c a g e s room, t o b e u s e d these  pair  to breed).  P. 1. n . 9  X  pairs  One c o n t r o l  o f e a c h o f t h e above s u b s p e c i e s o f l o c a l m a n i c u l a t u s  1962.  with  These mice have b e e n  f o r o v e r a y e a r and h a v e n o t l i t t e r e d .  a total  Four  ( P . m. a u s t e r u s , P. m. o r e a s , and P. m. s p p .  from Mandarte I s l a n d , B r i t i s h paired  early  and P.  as c o n t r o l s .  and one p a i r  of maniculatus  o n t h e same d a t e ,  i n t h e same  No l i t t e r s h a v e r e s u l t e d  from  pairs. The  negative results  of leucopus  o b t a i n e d by the attempted  and m a n i c u l a t u s  1937), and W a t e r s (I960). maniculatus,  chosen  support those of Dice The f a c t  litters,  indicates  apparently  inhibits  that  cages  something  (1931>  1933,  t h a t p a i r s o f l e u c o p u s and  a t random f r o m g r o u p s  same a r e a s and k e p t i n s i m i l a r  crossbreeding  o f mice caught  i n t h e same room,  i nthe  produced  other than l a b o r a t o r y c o n d i t i o n s  i n t e r b r e e d i n g b e t w e e n t h e two s p e c i e s .  33From my  work and  that leucopus w i l l  the experiments  o f o t h e r s , i t appears  n o t mate w i t h m a n l c u l a t u s  i n captivity.  b l o c k t o i n t e r b r e e d i n g may  be m o r p h o l o g i c a l , b e h a v i o r a l ,  physiological,  linked  The to  i n white mice  workers found  that  single  with sight or  ( B r u c e and  an e x t e r o c e p t i v e b l o c k  Parrott,  I960).  These .  i n t h e same  t h e i n t r o d u c t i o n o f a s t r a n g e male,  a male o f a d i f f e r e n t  or with the female.  be  or  smell.  or successive pregnancies  c o u l d be b l o c k e d b y  particularly near  perhaps  s m e l l o f a s t r a n g e male may  pregnancy  female  and  The  strain  from  These i n h i b i t o r y  t h e s t u d male,  e f f e c t s were a b o l i s h e d  i n mice from w h i c h t h e o l f a c t o r y b u l b s had b e e n removed. Perhaps  the females  mated w i t h a n i m a l s way.  The  odor  a role  leucopus  of the other species,  t o pregnancy.  leucopus  p r e s e n t my  results  own  reaction i n a later  and m a n i c u l a t u s On  a more o r  less might  s p e c l a t i o n i n mammals  in particular.  the r o l e o f odor  section of this  when  i n much t h e same  Hence, s p e c i e s o d o r  i n r e p r o d u c t i v e i s o l a t i o n and  i n g e n e r a l and  or manlculatus,  react  o f t h e s t r a n g e m a l e c o u l d be  permanent i n h i b i t o r play  of either  report.  I  i n interspecies  will  34. THE  DISTRIBUTION OF PEROMYSCUS LEUCOPUS NOVEBORACENSIS AND  PEROMYSCUS MANICULATUS  GRACILIS  D i f f e r e n c e s b e t w e e n P. 1.  noveboracensis  were i n v e s t i g a t e d n o t o n l y t o f i n d but  also  their  c o u l d be u s e d  History  to e x p l a i n  A t t e m p t s w i l l be made t o r e l a t e t h e  d i s t r i b u t i o n o f t h e mice w i t h e n v i r o n m e n t a l color o f pelage,  behavior,  of Distribution  and s p e c i e s  factors,  morphology,  interaction.  o f Peromyscus  Knowledge o f t h e h i s t o r y  of d i s t r i b u t i o n of species of the  genus Peromyscus i n e a s t e r n N o r t h A m e r i c a might p r o v i d e a t i o n f o r t h e i r present range. Wisconsin on  W a t e r s (I960) r e v i e w s  r e d i s t r i b u t i o n o f mice o f t h i s  t h e sequence o f e v e n t s  genus Peromyscus r e a c h e d  an explan-  the post-  g e n u s and s p e c u l a t e s  l e a d i n g t o s p e c i a t i o n and s u b s e q u e n t  development o f e x c l u s i v e ranges.  early Pleistocene.  gracilis  a method o f s e p a r a t i n g them  t o see i f d i f f e r e n c e s found  distribution.  a n d P. m.  He p r e s e n t s e v i d e n c e  t h a t the  the e a s t e r n U n i t e d S t a t e s d u r i n g the  I t i s probable  t h a t the frequent  s h i f t s and  a l t e r n a t i o n o f c l i m a t e and v e g e t a t i o n d u r i n g t h e g l a c i a l and interglacial to  shift  p e r i o d s o f the P l e i s t o c e n e a l s o  frequently, resulting  i n eastern North  elevations  i n highly variable  Peromyscus  s p e c i e s o f mice  America.  Waters proposes incipient  caused  t h a t toward t h e end o f t h e P l e i s t o c e n e , an  p o p u l a t i o n o f P. m a n l c u l a t u s  occupied  the higher  o f the southern h a l f o f the Appalachians,  i n c i p i e n t P. l e u c o p u s to the east, south,  population occupied  and west o f t h i s  area.  w h i l e an  the lower e l e v a t i o n s H i s data  indicate  35. t h a t a g e n e t i c divergence took p l a c e i n the i n c i p i e n t P. leucopus p o p u l a t i o n d u r i n g the l a t e P l e i s t o c e n e and r e s u l t e d i n forms w i t h e i t h e r s t r o n g l y or weakly b i c o l o r e d  tails.  Waters c o r r e l a t e s the northward movement o f maniculatus and leucopus w i t h b o t a n i c a l d a t a d e r i v e d from Braun ( 1 9 5 0 ) . A c c o r d i n g to t h i s author, spruce was the f i r s t to  f o r e s t component  move northward, f o l l o w i n g the r e t r e a t o f the W i s c o n s i n i c e  cap.  Oaks, a t f i r s t  spruce.  a s s o c i a t e d w i t h p i n e s , moved i n a f t e r the  Maniculatus moved n o r t h w i t h the expanding  b e f o r e leucopus.  spruce  forest,  Leucopus began to move northward w i t h the  expansion o f oak f o r e s t s , s p r e a d i n g throughout  the p o p u l a t i o n west o f the Appalachians  the midwest, t h a t e a s t o f the Appalachians  s p r e a d i n g a l o n g the A t l a n t i c c o a s t a l p l a i n .  Thus, maniculatus-  was a s s o c i a t e d w i t h c o n i f e r o u s and leucopus w i t h deciduous  forest.  The p r e s e n t r e l a t i o n s h i p . o f these s p e c i e s w i t h the v e g e t a t i o n of  O n t a r i o w i l l be d i s c u s s e d l a t e r . In  summary, Waters f e e l s t h a t the s p e c i e s and subspecies o f  Peromyscus are probably l i m i t e d to t h e i r p r e s e n t day ranges by c h a r a c t e r i s t i c s developed  through the l a t e P l e i s t o c e n e and d u r i n g  t h e i r p o s t - W i s c o n s i n movement northward. P r e s e n t D i s t r i b u t i o n o f Peromyscus i n O n t a r i o The p r e s e n t d i s t r i b u t i o n o f the two s p e c i e s has been o u t l i n e d by s e v e r a l authors (Osgood, 1909J  H a l l and Kelson, 1 9 5 9 ) .  For  t h i s study, I o b t a i n e d t r a p p i n g r e c o r d s o f Peromyscus i n O n t a r i o from the R o y a l O n t a r i o Museum o f Zoology.  These data are p r e s e n t e d  i n F i g . 18(a,b,c) and i n d i c a t e where Peromyscus have been trapped up t o 1960-61.  A l s o i n c l u d e d i n these f i g u r e s are d a t a I have  Figure 18(a).  Map  o f Counties and D i s t r i c t s o f  southeastern Ontario. 1. Essex  18. P e e l  35. Dundas  2. Kent  19.  York  36.  3. Lambton  20.  Dufferin  37. G l e n g a r r y  4. E l g i n  21. Ontario  38. P r e s c o t t  5 . Middlesex  2 2 . Durham  39. R u s s e l l  6. N o r f o l k  23.  40. C a r l e t o n  7. Oxford  24. P r i n c e Edward  41. Lenarck  8. B r a n t  25.  42. H a l i b u r t o n  9.  26. Grey  43. Muskoka  10. Welland  27.  44. Parry Sound  11. L i n c o l n  28. V i c t o r i a  45.  12. Wentworth  2 9 . Peterborough  46. M a n i t o u l i n  13. Waterloo  3 0 . Hastings  47. N l p i s s i n g  14. P e r t h  31.  48. Algoma  15.  3 2 . Frontenac  49.  3 3 . Leeds  5 0 . Timiskamlng  Haldimand  Halton  16. Huron  Northumberland  Bruce  Simcoe  Lennox and Addington  Stormont  Renfrew  Sudbury  17- W e l l i n g t o n 34-. G r e n v i l l e ( T h i s legend a l s o a p p l i e s to F i g u r e s 18(b) and 1 8 ( c ) )  Fi . B  18(a).  COUNTIES OF  AND  DISTRICTS  SOUTH-EASTERN  ONTARIO  Figure  18(b).  Range o f P. 1. n o v e b o r a c e n s i s i n  southeastern Ontario, taken p r i m a r i l y R.O.M.Z. t r a p p i n g r e c o r d s . indicate  The s t i p p l e d a r e a s ,  record(s) of animal(s)  identified  tentatively-  a s P. 1. n o v e b o r a c e n s i s .  b l a c k areas i n d i c a t e  from  The  counties or d i s t r i c t s  where n o v e b o r a c e n s l s h a s b e e n t r a p p e d .  F i g u r e 18(b).  RANGE IN  OF  R L.  NOVEBORACENSIS  SOUTH-EASTERN (ROMZ.  ONTARIO .  FILES)  Figure  18(c).  Range o f P. m. g r a c i l i s  eastern Ontario,  taken primarily  trapping records. record(s)  o f animal(s)  as P. m. g r a c i l i s . counties trapped.  The s t i p p l e d  f r o m R.O.M.Z. areas  tentatively  The b l a c k  or d i s t r i c t s  i n south-  indicate  Identified  areas  where g r a c i l i s  indicate has been  Figure 18(c).  RANGE IN  OF  R M. G R A C I L I S  SOUTH-EASTERN (R.OMZ.  ONTARIO  FILES)  36. o b t a i n e d from p u b l i s h e d f a u n a l surveys f o r p o r t i o n s o f O n t a r i o (Wright and Simpson, 1 9 2 0 ;  Snyder and L o g l e r , 1 9 3 0 ;  1931;  Snyder and L o g i e r , 1 9 3 1 ;  1941;  Jameson, 1 9 4 3 ;  Lanning, The  1954;  Clarke, 1933;  Rand, 1 9 4 5 ;  Snyder e t a l ,  Warburton, 1 9 4 9 ;  Brown and  Rutter, 1 9 5 9 ) .  d i s t r i b u t i o n s of P. 1.  noveboracensis  and P. m.  shown i n f i g . I8(b,c) are o n l y rough approximations (1)  Davis,  gracilis  because:  T r a p p i n g r e c o r d s f o r the whole of s o u t h e a s t e r n O n t a r i o are  Incomplete, and  (3)  (2)  Many specimens are o n l y t e n t a t i v e l y  identified,  T r a p p i n g r e c o r d s were mapped by c o u n t i e s and  districts,  not by s i t e of capture. Osgood's ( 1 9 0 9 , P. 1.  noveboracensis  p. 114,  f r o n t i s p i e c e ) d i s t r i b u t i o n maps of  and P. m.  gracilis  i n O n t a r i o showed t h a t  g r a c i l i s occupied the a r e a e a s t of the Great Lakes up t o the border of Quebec, while noveboracensis was Ontario.  Recent data ( R u t t e r , 1 9 5 1 ;  r e s t r i c t e d to  southern  R.O.M.Z. t r a p p i n g r e c o r d s ) ,  show t h a t the ranges o f the two. s p e c i e s have changed  slightly;  g r a c i l i s has moved i t s southern boundary northward, while noveboracensls has extended  I t s n o r t h e r n boundary f u r t h e r n o r t h .  C o r r e l a t i o n of Ranges of Mice w i t h V e g e t a t i o n The d i s t r i b u t i o n s of the mice can be roughly w i t h types of v e g e t a t i o n .  Osgood ( 1 9 0 9 ) ,  correlated  and Cross and Dymond  ( 1 9 2 9 ) noted t h a t g r a c i l i s p r e f e r r e d the c o l d e r , more moist p l a c e s , or deep, mostly  c o n i f e r o u s woods, while leucopus p r e f e r r e d the  warmer, d r y e r , more open country, or deciduous The  woods.  changes i n d i s t r i b u t i o n s of the mice, s i n c e 1 9 0 9 ,  can  a l s o be roughly c o r r e l a t e d w i t h changes i n types of v e g e t a t i o n .  37. S i n c e 1909,  c o n s i d e r a b l e s e t t l e m e n t has  Ontario  much o f t h e n a t i v e v e g e t a t i o n h a s  and  Settlement These  has  spread northward  and  taken place i n  conifer  b e e n removed.  f o r e s t has  c l e a r i n g s encouraged the growth o f deciduous  conifer-hardwood  subclimax.  G r a c i l i s has  northward, The  presumably f o l l o w i n g the e x p a n s i o n  leucopus  with deciduous  (1959).  He  found  f o r e s t was  maniculatus  t o be  with  of coniferous  extended  i t s range  o f hardwood  forest.  coniferous forest  recently  r e p o r t e d by  positively  a v o i d e d p l a n t s o f the o a k - c h e s t n u t  was  positively  r e g i o n and  associated with  forest region.  a v o i d e d p l a n t s o f the hemlock-white p i n e  -northern  f o r the  a s s o c i a t i o n of leucopus  r e g i o n s c a n be  shown by  and  maniculatus  with forest  ranges  the f o r e s t r e g i o n s o f s o u t h e a s t e r n O n t a r i o .  f o r e s t r e g i o n s o f O n t a r i o a r e w e l l known and r e d e s c r i b e d b y Rowe ( 1 9 5 9 ) . and  maniculatus 19)  (fig. lies  leucopus  lies  t h e i r ranges forest  a map  i n the  The  of  range of  region.  o v e r l a p c o i n c i d e s w i t h the zone.  The  The  been  leucopus Ontario  maniculatus  coniferous f o r e s t region while forest  their  have r e c e n t l y  of forest regions of southeastern  i n the deciduous  transition  comparing  Comparing the ranges  shows s e v e r a l c o r r e l a t i o n s .  primarily  forest  region.  Further evidence  with  region  Leucopus  a s s o c i a t e d w i t h p l a n t s o f the oak-chestnut  hardwoods f o r e s t  and  Klein  p l a n t s o f t h e h e m l o c k - w h i t e p i n e - n o r t h e r n hardwoods f o r e s t and  the  withdrawn i t s range  Leucopus has  a s s o c i a t i o n of maniculatus  been c l e a r e d .  t r e e s of  northward, presumably f o l l o w i n g the r e c e d i n g b o r d e r f o r e s t where i t p r e f e r s t o l i v e .  southern  that of  a r e a where  coniferous-deciduous  F u r t h e r , c o r r e l a t i o n w i t h the  principle  Figure  19.  Forest  r e g i o n s of  southeastern Ontario  ( a f t e r Rowe, 1 9 5 9 ) • 1.  Deciduous f o r e s t Niagara  2-10,  11.  12.  region  section G r e a t L a k e s - S t . Lawrence f o r e s t  2.  Huron-Ontario  3.  G e o r g i a n Bay  4.  Algonquln-Pontlac  5.  Middle Ottawa  6.  Upper S t .  7.  Algonquin-Fontiac  section  8.  S u d b u r y - N o r t h Bay  section  9-  Timagami  section section  section  Lawrence  section'  section  10.  Algoma  12.  Haileybury  Boreal  section  section  forest  clay  section  region  Missinaibi-Gabonga  section  region  Figure i  9  .  FOREST  REGIONS  OF S O U T H - E A S T E R N (AFTER  ROWE,  ONTARIO 1959)  38. trees  i n e a c h r e g i o n shows t h a t  associated with trees while  ulatus and  of f i g .  18(b)  distribution for gracilis Huron, a t the  s o u t h end  edge o f L a k e O n t a r i o . areas i n r e l i c  forest  reveals  a l o n g the  superficial  Perhaps  and  and  gracilis  Williams  and  w a t e r may  o f these mice.  Hamilton  (1959),  molluscs. exhibited.  No  apparent  Food  growth.  a l o n g the n o r t h e r n  i s present i n these  i s not  d i s t r i b u t i o n may Other  factors  foods eaten by (1922),  Getz  differences clearly  as  the  leucopus  and  Cogshall (1928),  (1952), (1961),  Cameron  (1956),  and Howard  of foods i n c l u d i n g fungi,  only  such  the ones g o v e r n i n g  Jackson  Water  seeds,  (1961).  nuts,  green vegetation, r o o t s ,  and  i n f o o d p r e f e r e n c e were  related  to the d i s t r i b u t i o n  of  mice. Temperature  be  The  Connor ( I 9 6 0 ) ,  arachnids, f r u i t ,  be  by D i c e  (1941),  species ate a v a r i e t y  Insects,  growth  t o F o o d . T e m p e r a t u r e and  not causal f a c t o r s .  m a n i c u l a t u s were m e n t i o n e d  (1940),  manic-  stands o f c o n i f e r o u s f o r e s t .  temperature,  distributions  these  that  a somewhat d i s c o n t i n u o u s  C o r r e l a t i o n s b e t w e e n v e g e t a t i o n and  Both  and  s o u t h e a s t e r n edge o f L a k e  o f Lake E r i e ,  R e l a t i o n s h i p o f Ranges o f M i c e  Burt  I conclude  birch,  i s associated with regions of coniferous forest  leucopus with regions of deciduous  food,  and  i s a s s o c i a t e d w i t h s u g a r maple  From t h e e v i d e n c e p r e s e n t e d h e r e ,  Examination  be  of maniculatus i s  s u c h as s p r u c e , p i n e , a s p e n ,  that of leucopus  beech.  the range  factors  t o l e r a n c e s of leucopus  influencing  and E s k r l d g e and U d a l l  and m a n i c u l a t u s  their distribution. (1955), f o u n d  might  However, Howard (1951),  that both  species could  39. withstand f r e e z i n g temperatures provided s u f f i c i e n t food was available.  Sealander (1952) showed that P. leucopus could '  endure temperatures from  +35 to -25°C.  Getz (1961) concluded  that temperatures had no obvious i n f l u e n c e upon the l o c a l d i s t r i b u t i o n of P. leucopus.  High temperatures probably do  not a f f e c t the mice because of t h e i r nocturnal behavior. Perhaps the a v a i l a b i l i t y of food d u r i n g the winter d i f f e r s f o r each species and t h i s governs t h e i r r e s p e c t i v e d i s t r i b u t i o n s . No data are a v a i l a b l e on t h e i r a b i l i t y to survive t h i s c r i t i c a l p e r i o d under conditions of winter temperature, provided w i t h s e l e c t e d kinds and amounts of food. From the evidence above, I conclude that there i s l i t t l e d i f f e r e n c e i n temperature tolerance between leucopus and manlculatus.  Thus temperature, by i t s e l f , does not appear t o  be a f a c t o r determining t h e i r d i s t r i b u t i o n s . The water requirements of the two species might i n f l u e n c e their distribution.  Lindeborg (1952) observed that leucopus  and manlculatus d i s p l a y e d s i m i l a r r e a c t i o n s to l i m i t e d amounts of water.  N e i t h e r could survive on 0.2 cc./day and both l o s t  weight on 0.4 cc./day.  K l e i n (1959) showed that both species  used about the same amount of water:  leucopus, 0.43  cc./gram  body weight or 7.6 c c . / i n d i v i d u a l / d a y ;  g r a c i l i s , 0.42  cc./gram  body weight or 7.4 c c . / i n d i v i d u a l / d a y .  These studies Indicate  that water requirements f o r leucopus and maniculatus are not appreciably d i f f e r e n t . for their distributions.  This f a c t o r does not appear r e s p o n s i b l e  40. The of  Morphology  and Pelage  two  i n m o r p h o l o g y and c o l o r o f p e l a g e b e t w e e n t h e  s p e c i e s might be c o r r e l a t e d w i t h d i f f e r e n c e s  tions.  (1940)  Dice  and h i n d f o o t  suggested  that  l e n g t h w i t h type  u s u a l l y found  this  i n heavily  i n t u r n might  Horner  (1954)  demonstrated  climb a r t i f i c i a l also  forested  areas  that  mice,  (Pox,  f o r example,  1948),  tree  the long t a i l  was a n a d a p t i v e  structure  semi-arboreal existence.  Blair,  1950;  Linduska,  Horner  (1954)  c l a s s i f i e d b o t h P. 1 .  t h a n P.  1950;  Horner,  1954;  noveboracensis  b e t w e e n them i n c l i m b i n g p e r f o r m a n c e s . climbing glass drinking S i n c e no d i f f e r e n c e  tubes  of maniculatus  1940;  I960).  and P.  m.  difference  I observed both species  and cage w a l l s  i n the climbing a b i l i t i e s  over leucopus have been found,  leucopus, Burt,  Connor,  as b e i n g s e m i - a r b o r e a l and d e t e c t e d l i t t l e  longer t a i l  ability  t r u n k s and c r o s s gaps o f v a r y i n g w i d t h s .  a r e known t o b e s e m i - a r b o r e a l (Chenoweth, 1 9 1 7 ;  gracilis  and  Peromyscus i n t h e i r  A l t h o u g h P. m a n l c u l a t u s h a s a l o n g e r t a i l both  essentially  s e m i - a r b o r e a l P e r o m y s c u s were  than t e r r e s t r i a l  showed t h a t  which f a c i l i t a t e d  Longer-tailed  might  length  be c o r r e l a t e d w i t h s e m i - a r b o r e a l b e h a v i o r .  significantly better  She  i n distribu-  correlation of t a i l  of environment  a c o r r e l a t i o n with behavior.  are  to  as an E x p l a n a t i o n  Distribution Differences  be  C o l o r o f the Mice  I conclude t h a t  i s not causally  with equal  facility.  of maniculatus the s l i g h t l y  related  to i t s  distribution. C o r r e l a t i o n between s i z e distribution of either example,  the s l i g h t l y  o f o t h e r body p a r t s and t h e  s p e c i e s c a n o n l y be s u g g e s t e d . longer r o s t r a l  l e n g t h o f leucopus  For might  41. p r o v i d e more l e v e r a g e f o r e a t i n g that  a r e commonly f o u n d  conjecture in  the  and  large,  i n i t s range.  I have found  no  for birds  s u p p o r t i n g d a t a on  different  1944, 1947).  Lack,  pelage  this  color  function.  gracilis  soil  positively  surface.  Dice  color  colored.  however,  are  identical,  Gracilis,  so s l i g h t  i s o f importance  a l t h o u g h the former  B r i g h t e r p e l a g e may  have a  i n pelage that  m i g h t be reason,  factors  i t h a r d t o be  and  their distribution.  species,  convinced  that  forms.  maniculatus For  this  a t e v e r y o p p o r t u n i t y i n the  s p e c i e s were g o o d c l i m b e r s and  pelage  Distribution  i n b e h a v i o r between leucopus  t h e y were o b s e r v e d  become  colored coniferous  i n the d i s t r i b u t i o n o f these  influencing  to  protective  colored  c o l o r b e t w e e n t h e two  I find  tends  forests  with i t s l e s s b r i g h t l y  B e h a v i o r o f t h e M i c e as a Cause o f Differences  of  noveboracensis  colored.  Differences  color  c o l o r s of pelage of  m i g h t g a i n more p r o t e c t i o n i n t h e l e s s b r i g h t l y forests.  and  (1940)  (1947) o f i t s  f u n c t i o n I n t h e autumn when l e a v e s i n d e c i d u o u s highly  foods  gave f u r t h e r e v i d e n c e  The  are n e a r l y  more b r i g h t l y  o f the  t o be  c o r r e l a t i o n between s o i l  o f P e r o m y s c u s and  protective  i s an a d a p t a t i o n f o r t a k i n g  c o l o r o f mammals t e n d s  w i t h the  demonstrated  Both  Peromyscus  size.  pelage  correlated  this  i s only  (1947) c o n s i d e r s t h a t t h e marked d i f f e r e n c e i n s i z e o f  The  be  nuts  f o r f o o d h a n d l i n g have  1932;  (Robbins,  beaks o f the Galapagos f i n c h e s  and  s u c h as  literature.  been observed  of  foods  However, t h i s  Similar morphological differences  Lack  hard  readily  climbed the  laboratory. glass  42. d r i n k i n g tubes on g r a v i t y water b o t t l e s i n t h e i r  cages.  When I cleaned out cages, o c c a s i o n a l l y an animal would escape.  Escapees o f each s p e c i e s tended to behave d i f f e r e n t l y .  Maniculatus moved comparatively slowly and appeared t o be u n c e r t a i n about which d i r e c t i o n t o go. Leucopus r a n q u i c k l y as soon as i t had escaped from the cage.  Horner  (1954) a l s o noted t h a t  leucopus moved more q u i c k l y and u n h e s i t a t i n g l y than m a n i c u l a t u s . T h i s d i f f e r e n c e i n approach t o a new s i t u a t i o n c o u l d be r e l a t e d to the speed a t which the animal i s capable o f t r a v e l l i n g , o r t o some i n h e r i t e d f a c t o r r e l a t e d t o i t s normal h a b i t a t .  Leucopus  has been found to be the f a s t e r o f the two on the ground and Benton,  1954).  (Layne  These i n v e s t i g a t o r s found t h a t leucopus  could t r a v e l a t 8.46 f e e t / s e c o n d (range, 6.9-10.0 f e e t / s e c o n d ) , while maniculatus could t r a v e l o n l y 6.4 f e e t / s e c o n d (range, 6.2-6.6 f e e t / s e c o n d ) . The two s p e c i e s o f Peromyscus r e a c t e d d i f f e r e n t l y to handling.  P. 1. noveboracensis was d i f f i c u l t to c a t c h and would  r u n and jump r a p i d l y around the cage. e a s i l y approached of the cage door.  P. m. g r a c i l i s was f a i r l y  and o f t e n came out t o i n v e s t i g a t e the opening S v l h l a (1932) found leucopus t o be more  e x c i t a b l e and d i f f i c u l t t o handle than P. m. b a l r d i i .  Horner  (1954) and F o s t e r (1959) r e p o r t e d t h a t g r a c i l i s was more e a s i l y handled than b a l r d i i .  I found d i f f e r e n c e s i n the b e h a v i o r of  leucopus and m a n i c u l a t u s .  Leucopus was the more h i g h l y  excitable  and f a s t e r moving o f the two forms. Leucopus as mentioned  above, i s c o n s i d e r e d by many workers  to be the more e n t e r p r i s i n g , l e s s h a b i t a t - r e s t r i c t e d mouse  43. (Wilson,  I960).  1945; Perhaps  reflects  1955;  I960;  Connor,  the e n t e r p r i s e  related  and  W i r t z and  and g r e a t e r s p e e d  i t s wide c h o i c e o f h a b i t a t s .  enterprising be  Brand,  of  leucopus  Manlculatus i s less  s l o w e r o f movement t h a n l e u c o p u s  to i t s presence  Pearson,  and  t h i s may-  i n more homogeneous h a b i t a t s .  t h i s b e h a v i o r o f each form  i s cause  or e f f e c t  Whether  o f the v e g e t a t i o n  i n w h i c h t h e y o c c u r , i s unknown.  I n t e r a c t i o n B e t w e e n S p e c i e s as a Cause o f The  role  o f s p e c i e s i n t e r a c t i o n i n d i s t r i b u t i o n has  (1922),  examined by D i c e Klein  (1959)  suggest  and  m a n l c u l a t u s , no  areas.  (1961)  also r e s t r i c t  found  t h a t P.  m a n i c u l a t u s have more r e s t r i c t e d h a b i t a t of  sympatry  competition.  attributed,  Interspecific  present  distribution  sympatric species their  distribution  o r e a s and  P.  p r e f e r e n c e s i n the  t h a n where e a c h o c c u r s a l o n e .  h a b i t a t d i s t r i b u t i o n was  ecological  interspecific  not only prevent  b u t i t might  Sheppe  some  e v i d e n c e was  s u c h as  been  (I96I).  Sheppe  i n d e t e r m i n i n g the l o c a l  I n t e r a c t i o n may  interbreeding,  in local  and  played a part  the forms.  from  (1959),  that behavioral factors,  antagonism, of  Klein  r e p o r t e d t h a t a l t h o u g h t h e r e was  separation of leucopus to  Distribution  The  i n part,  i n t e r a c t i o n may  restriction to  o r may  area of  interspecific not  influence  distribution. Klein  (1959)  f o u n d no  keeping leucopus  and  initial  to each  c o u l d be  response  antagonism  manlculatus  mixed p a i r s  of both  together.  o t h e r was  found h u d d l i n g t o g e t h e r . species lived  or cannibalism r e s u l t e d  one  He  observed  of caution, but  I n my  that soon  from their  they  crossbreeding experiments,  amicably  and  no  antagonism  44. between the two was  evident.  Unfortunately, i n s u f f i c i e n t  numbers o f l i v e animals were a v a i l a b l e to t e s t  antagonism  between two p a i r s (one p a i r of each s p e c i e s ) d u r i n g the b r e e d i n g season.  Perhaps a t t h a t time o f the year, antagonism p l a y s  a l a r g e r o l e i n keeping the s p e c i e s a p a r t .  An experiment w i t h  one p a i r o f leucopus kept i n the same cage as two  successive  p a i r s o f l o c a l maniculatus, gave i n t e r e s t i n g o b s e r v a t i o n s . b o t h i n s t a n c e s , the female o f the p a i r o f maniculatus was dead.  found  The dead animals were not eaten b u t were m u t i l a t e d ,  the b l o o d - s p a t t e r e d cage gave evidence t h a t the dead had moved about b e f o r e they had succombed.  In  and  animals  These deaths o c c u r r e d  s e v e r a l months a f t e r b o t h p a i r s had been p l a c e d i n the cage and, a t no time was  antagonism  shown between the animals.  The  dead animals had been b i t t e n about the e a r s and the t a i l s were b a d l y chewed.  N e i t h e r the p a i r of leucopus nor the male  maniculatus showed evidence of h a v i n g fought. f o r t h i s b e h a v i o r has been found.  No e x p l a n a t i o n  B u r t ( 1 9 4 0 ) r e p o r t e d t h a t the  female leucopus h o l d s a d e f i n i t e t e r r i t o r y i n the b r e e d i n g season and i s a n t a g o n i s t i c to o t h e r leucopus females. maniculatus females were k i l l e d  i n mid-winter.  I n t r a s p e c i f i c antagonism was o n l y leucopus.  However, b o t h  e v i d e n t i n cages  containing  In one cage, f o u r males were d i s c o v e r e d s t a r t i n g  to e a t the warm c a r c a s s o f t h e i r female l i t t e r mate. animal had a b a d l y chewed t a i l  and neck.  The  dead  I n another cage, t h r e e  of seven male leucopus were found huddled i n the c o r n e r of the cage f a r t h e s t from the n e s t .  A l l had raw,  stubby, s w o l l e n t a i l s ,  and would not e n t e r . t h e n e s t even when prodded. leucopus were not i n j u r e d .  The o t h e r f o u r  Reasons f o r antagonism  i n leucopus  45. are unknown. Only one  i n s t a n c e of antagonism was  A female P. m. mates.  noted among m a n l c u l a t u s .  austerus k i l l e d and p a r t l y ate her t h r e e  litter  F u r t h e r i n v e s t i g a t i o n r e v e a l e d t h a t l a c k of food  prompted the  had  cannibalism.  I n t e r a c t i o n Between S p e c i e s by Odor I n t e r a c t i o n between the two  forms c o u l d be r e s p o n s i b l e f o r  t h e i r d i f f e r e n c e s i n d i s t r i b u t i o n and a b i l i t y to l i v e s y m p a t r i c a l l y , s t i l l r e t a i n i n g separate i d e n t i t y . t h e r e was  evidence  Moore (1961) r e p o r t e d t h a t  to support the r o l e of o l f a c t i o n i n the  i s o l a t i o n of P. maniculatus  and P. p o l i o n o t u s .  sexual  Because novebora-  c e n s i s and g r a c i l i s occur s y m p a t r i c a l l y , and are not known to i n t e r b r e e d where they do, a t i o n might be  i t seemed p l a u s i b l e t h a t odor d i s c r i m i n -  i n v o l v e d i n t h e i r apparent  sexual I s o l a t i o n .  To  t e s t t h i s , p l a n s were drawn up f o r an odor d i s c r i m i n a t i o n apparatus ( o l f a c t o m e t e r ) . Apparatus f o r T e s t i n g R e c o g n i t i o n of Odor C e r t a i n s p e c i f i c a t i o n s had t o be c o n s i d e r e d i n the d e s i g n of the o l f a c t o m e t e r .  Firstly,  i t should be as l i g h t - p r o o f  a i r - t i g h t as p o s s i b l e to prevent e x t e r n a l s t i m u l i from results.  Secondly,  affecting  i t should present the animal w i t h a number  of c h o i c e s , each of which could e q u a l l y be taken. method of r e c o r d i n g the animal's  T h i r d l y , some  movement w i t h i n the  must be I n c o r p o r a t e d i n the d e s i g n o f the apparatus. the suspected  and  apparatus Fourthly,  odor must be r e a d i l y o b t a i n a b l e and i n a form t h a t  c o u l d be used i n the  apparatus.  A f t e r c o n s i d e r a t i o n of these s p e c i f i c a t i o n s , a t e s t i n g  46. chamber was c o n s t r u c t e d ( F i g . 20).  The apparatus c o n s i s t e d o f  a c e n t r a l chamber with moveable doors c o n n e c t i n g to three r a d i a t i n g arms.  I t was c o n s t r u c t e d e n t i r e l y o f sheet metal and  was made as a i r - t i g h t and l i g h t - i m p e r v i o u s as p o s s i b l e .  The  t h r e e r a d i a t i n g , arms were spaced e q u i d i s t a n t l y t o g i v e the animals an equal o p p o r t u n i t y t o e n t e r any one o f them.  Three arms were  chosen because there were two s p e c i e s odors t o t e s t and the t h i r d could be used as a c o n t r o l . The problem o f r e c o r d i n g the a c t i v i t y o f an animal i n a metal c o n t a i n e r was s o l v e d by having a moveable j o i n t i n the middle o f each arm.  A p r o j e c t i n g p i e c e o f metal ( w r i t e r ) was  s o l d e r e d onto t h e end o f each arm.  The w r i t e r t r a c e d a r e c o r d  of the movement o f the animal i n the end o f the arm, on a p i e c e of smoked kymograph paper. S e l e c t i o n o f a source o f odor was a problem because Peromyscus have a number o f glands (eg. a n a l , p r e p u t i a l , c l i t o r a l ) which c o u l d be r e s p o n s i b l e f o r a s p e c i e s - s p e c i f i c o f o r and, almost n o t h i n g i s known about t h e i r f u n c t i o n .  Mammals are known to use  odor f o r sexual a t t r a c t i o n , t e r r i t o r y and home range marking, and p o s s i b l y f o r p r o t e c t i o n ( B o u r l i e r e , 1956, pp. 103-107, 225-  230).  Hedlger (1949, from B o u r l i d r e , 1956) r e p o r t e d the use o f  u r i n e and f a e c e s by the male pygmy hippopotamus i n marking i t s territory.  The ease o f o b t a i n i n g u r i n e and f a e c e s , and the  knowledge t h a t i t i s used by mammals i n the marking o f t e r r i t o r i e s and t r a i l s , made t h i s t h e choice f o r a source o f odor. The apparatus was s e t up as i n F i g . 20(B). cage l i n i n g was p l a c e d i n two of the three arms.  S o i l e d paper The t h i r d arm  was used as t h e c o n t r o l and contained only c l e a n paper cage  lining.  Figure 20.  Diagram  o f odor d i s c r i m i n a t i o n a p p a r a t u s  (olfactometer). A.  P l a n o f a p p a r a t u s as s e e n f r o m  B.  S i d e view  above.  of apparatus i n operation.  16' ODOR  DOOR  LEVER WRITING  CENTRAL  MOVEABLE  TIP  CHAMBER  JOINT  A.  P L A N  B.  E L E V A T I O N  ELASTIC BAND RING  STAND  KYMOGRAPH DRUM  D I S C R I M I N A T I O N  A P P A R A T U S  47., A mouse ( t e s t e r ) was t h e n p l a c e d 10-15  minutes  i n the c e n t r a l  t o a l l o w i t t o become  accustomed  chamber f o r to the apparatus.  E a c h e x p e r i m e n t was r u n a t t h e same t i m e o f day ( 5 : 3 0 P.M.) f o r t h e same l e n g t h o f t i m e ( o n e h o u r ) . more  than a s i n g l e To  start  and  No a n i m a l was u s e d f o r  experiment.  an e x p e r i m e n t ,  t h r e e kymographs " a t t a c h e d " t o t h e  e n d s o f t h e arms were t u r n e d o n and d o o r s l e a d i n g f r o m t h e central into  chamber t o t h e arms were Opened.  t h e ends o f t h e arms was r e c o r d e d as a " d i p " o n t h e kymograph  paper.  The i n i t i a l  movement  as a s e r i e s o f r a p i d v i s i t s would  E n t r a n c e o f t h e mouse  settle  o f t h e mouse was u s u a l l y r e c o r d e d t o e a c h arm.  G r a d u a l l y t h e mouse  down and t e n d t o r e m a i n l o n g e r i n one o f t h e arms.  Results o f Tests with the Olfactometer These  p r e l i m i n a r y e x p e r i m e n t s were r u n t o t e s t  o f t h e a p p a r a t u s and t o see i f mice other  species odors.  do r e s p o n d t o t h e i r  own o r  S e v e n e x p e r i m e n t s were done and t h e r e s u l t s  are g i v e n i n the Appendix is  the d e s i g n  (Table V).  A summary  o f the r e s u l t s  giv6n i n Table 3 .  Table 3.  R e s u l t s o f odor d i s c r i m i n a t i o n  Expt. No. Tester  1.  9P.l.n.  2.  P.m.£.  Odor Control cfand ?P.m.£. <*and ? P . l . n . Residual control R e s i d u a l cfand 9 P . m . £ . R e s i d u a l dand?J?.l.n.  experiments.  No. o f T i m e s - E n t e r e d Arm  % of T o t a l Time i n Arms  4 6 10 *  5.2 11.7 83.1 *  7 * 6 6  44.0 *  18.5 37.5  48. Table  3 (cont'd).  Expt. Tester No.  R e s u l t s o f odor d i s c r i m i n a t i o n experiments. No. o f Times E n t e r e d Arm  Odor  % of Total Time i n Arms  3.  cfP.m.£.  Control cfand $ P . m . £ . cfand 9 P . l . n .  28 33 * 23  44.0 * 25.0 31.0  4.  9P.S«£*  Control cfand 9P.m.£. Cfand 9 P . l . n .  31 * 25 30  62.6 * 15.9 21.5  5.  ?P.l.n.  Control  9P.m.£.  30 * 22 25  85.3 * 4.6 10.1  cfP.m.g.  6.  Cfp.l.n.  Control cfand 9 P.m.g_. cfand 9 P . l . n .  11 18 31 *  5.4 20.8 73.8 *  7.  £P.m.fi.  Control cfP.rn.g_. Oand 9 P . l . n .  20 31 * 22  4.0 79 ~. 3 * 16.7  *  denotes the g r e a t e s t number o f times any arm was entered and the g r e a t e s t % o f t o t a l time spent i n any arm. Although t h i s odor d i s c r i m i n a t i o n experiment was intended  to be only a p r e l i m i n a r y  study and few t e s t s were run,  the above r e s u l t s are worthy o f comment.  I believe  S i n c e the t e s t e r was  used only once, no b i a s was i n c u r r e d by h a v i n g the animal f o l l o w a l e a r n e d path.  The t e s t s were r u n e a r l y i n January and none  of the mice used were i n b r e e d i n g  condition.  T h i s r u l e s out  p o s s i b l e changes i n response to odor caused by animals b e i n g i n estrus.  The r e s i d u a l odor t e s t (no. 2) was conducted t o f i n d  out how e f f e c t i v e l y the apparatus had been cleaned.  In t h i s  test,  only c l e a n paper cage l i n i n g was p l a c e d a t the end o f each arm. I t was decided  t h a t although r e s u l t s showed some r e s i d u a l odor  49was p r e s e n t i n the arms, i t would not g r e a t l y a l t e r the f o l l o w i n g r e s u l t s p r o v i d e d each arm was c o n s i s t e n t l y used f o r the same odor. G e n e r a l l y , the t e s t e r e n t e r e d most o f t e n and/or stayed l o n g e s t i n the arm c o n t a i n i n g i t s own s p e c i e s odor (experiments no. 1, 2, 3, 6, 7).  E x c e p t i o n s t o t h i s are experiments 4- and 5  where the animals e n t e r e d more o f t e n and stayed l o n g e r i n the control.  No e x p l a n a t i o n i s o f f e r e d f o r experiment 4, hut i n  experiment 5» the animal appeared to be a v o i d i n g the arms which b o t h contained the odor of the o t h e r s p e c i e s .  Perhaps t h e r e  i s an a c t i v e avoidance o f the o t h e r s p e c i e s i n n a t u r e by odor? Another g e n e r a l i z a t i o n t h a t can be made i s t h a t the t e s t e r s appeared t o p r e f e r arms c o n t a i n i n g an odor t o the arm used as a c o n t r o l (experiments no. 1, 2, 6, 7).  Why t h i s i s so i s not  known. The p o s s i b i l i t i e s o f t e s t i n g s p e c i e s i n t e r a c t i o n by odor d i s c r i m i n a t i o n are only b e g i n n i n g t o be r e a l i z e d . thus f a r i n d i c a t e :  The r e s u l t s  (1) The apparatus works and (2) There seems  to be measureable odor d i s c r i m i n a t i o n i n the mice.  Continuation  of these experiments promises t o g i v e v a l u a b l e r e s u l t s  which  may h o l d the key t o s e x u a l i s o l a t i o n i n sympatrlc s p e c i e s and perhaps a c l u e to p a t t e r n s o f d i s t r i b u t i o n i n c l o s e l y species.  related  50. SUMMARY 1.  The aims o f t h i s study were:  (1) t o i n v e s t i g a t e  m o r p h o l o g i c a l d i f f e r e n c e s between two s i m i l a r  s p e c i e s o f mice,  Peromyscus leucopus noveboracensis and P. maniculatus  gracilis,  to f i n d a method o f s e p a r a t i n g them, and (2) t o determine whether m o r p h o l o g i c a l o r other d i f f e r e n c e s found between the two s p e c i e s c o u l d be used t o e x p l a i n t h e i r 2.  distribution.  Most mice used i n t h i s study came from areas i n O n t a r i o  where each was known t o occur a l o n e .  Samples o f leucopus came  from areas near K i n g s t o n and manlculatus from A l g o n q u i n Park. The only e x c e p t i o n i s the sample from the a r e a o f sympatry (Renfrew 3.  and Frontenac Counties).. The measurements o f 169 mice o f both s p e c i e s were a n a l y z e d  g r a p h i c a l l y and s t a t i s t i c a l l y t o f i n d a method t o separate them. The  c h a r a c t e r s e v a l u a t e d were:  weight, h i n d f o o t l e n g t h , e a r  l e n g t h , body l e n g t h , t a i l l e n g t h , i n t e r p a r i e t a l width,  inter-  p a r i e t a l l e n g t h , s k u l l l e n g t h , and r o s t r a l l e n g t h . 4.  No one c h a r a c t e r was u s e f u l i n s e p a r a t i n g the two forms.  From an e v a l u a t i o n o f the d i f f e r e n c e s i n s i z e o f p a r t s o f the mice, i n d i c e s were developed which completely separated leucopus and maniculatus o f a l l sexes and ages examined. were:  These i n d i c e s  ear length X t a i l length X i n t e r p a r i e t a l length s k u l l length ear  length X t a l l  or  length X interparietal length  r o s t r a l length 5.  The i n d i c e s completely separated a sample o f 12 leucopus  and manlculatus t r a p p e d i n the zone o f sympatry. 6.  B r e e d i n g experiments were conducted t o see i f the two  s p e c i e s would i n t e r b r e e d .  Four mixed p a i r s o f leucopus and l o c a l  51. m a n i c u l a t u s were k e p t t o g e t h e r f o r o v e r a y e a r and d i d n o t produce total  offspring.  of three l i t t e r s  produced  a total  No  litters  P.  1.  7.  8. species  An  oreas  mixed p a i r s  pair  of g r a c i l i s  same l e n g t h o f t i m e ,  d i s t r i b u t i o n o f P.  attempt  was  1.  manlculatus  failed  o f P.  m.  days.  to breed).  gracilis One  (controls),  failed  to  pair  and of  kept  reproduce. P.  m.  made t o e x p l a i n t h e d i s t r i b u t i o n  of  of f a c t o r s  temperature  of pelage,  noveboracensis  and  tolerance,  leucopus  water requirement,  M a n l c u l a t u s was  i n deciduous  found between the ranges  temperature of pelage.  tolerance,  each  s u c h as v e g e t a t i o n , f o o d  and b e h a v i o r .  c o n i f e r o u s and  rence,  one  on t h e b a s i s  l a t i o n was  found to  vegetation.  o f t h e m i c e and  water requirement,  morphology,  No  occur corre-  food  prefe-  morphology,  C o r r e l a t i o n between t h e i r ranges  and  and  behavior  doubtful. 9.  cause  I n t e r a c t i o n between the  of d i s t r i b u t i o n .  w i t h the odor  o f mice.  a c t i o n between mice by that  odor  o f i t s own  the c o n t r o l  s p e c i e s was  investigated  Experiments  were c o n d u c t e d  The  was  purpose  olfaction.  the mice responded  e n t e r e d more o f t e n and  to  two  ( P . m.  of  a  i n O n t a r i o vras r e d e s c r i b e d u s i n g r e c e n t d a t a .  preference, color  and  f o r the The  gracilis  was  three control pairs  litters  from  of leucopus produced  n o v e b o r a c e n s i s k e p t t o g e t h e r f o r 71  together  color  and  o f two  resulted  noveboracensis  in  Two-, c o n t r o l p a i r s  t o the odor  o f o t h e r mice.  a  i n an o l f a c t o m e t e r  to investigate  P r e l i m i n a r y work  as  inter-  suggested A mouse  s t a y e d l o n g e r i n a chamber c o n t a i n i n g t h e  species.  A  chamber c o n t a i n i n g o d o r was  chamber w i t h o u t  odor.  preferred  52. APPENDIX Table I .  Measurement data p l o t t e d i n b a r diagrams ( i n mms.). E a r L.  Body L.  T a i l L.  S k u l l . L. R o s t r a l L .  Range Adult d"P.l.n. Mean 1 S.D. 2 S.E. N=35  15.0-17.3 16.2 0.67 0.22  73.4-96.4 85.5 5-30 1.80  62.1-88.2 74.9 5.70 1.94  2 5 . 1 - 28.0 7.9-9.5 8.7 26.4 0.68 O.38 0.24 0.13  Range Adult 9 P . l . n . Mean 1 S.D. 2 S.E. N = 33  14.8-17-5 16.0 0.69 0.32  76.5-96.0 86.4 5.50 1.92  61.2-86.9 74.0 6.40 2.20  2 5 . 0 - 27. 9 26.4 0.69 0.24  7.9-9.1 8.6 O.32 0.11..  Range Adult d*P.m.g. Mean 1 S.D. N = 22 2 S.E.  17.0-20.9 18.8 0.85 0.38  80.6-97.4 88.6 4.60 2.10  76.2-97.2 86.8 5.80 2.60  24.6- 27. 0 25.9 0.61 0.28  7.6-9.1 8.5 0.40 0.18  Adult Range ?P_.m..g.. Mean 1 S.D. 2 S.E. N=31  17.0-20.4 19.0 0.76 0.28  72.5-101.0 69.5-105.5 89.2 86.8 6.80 8.90 2.40 4.00  24.3- •27.9 25.7 0.76 0.28  7.9-9.3 8.5 0.39 0.14  Subad.  Range c5p.rn.g_. Mean 1 S.D. N = 22 2 S.E.  17.4-20.1 18.6 0.73 0.34  75.0-86.0 82.5 3.02 1.38  66.8-97.0 81.8 6.80 3.00  24.0- 26. 1 25.3 O.54 0.24  7.7-8.6 8.2 0.21 0.10  Subad. Range ?P.m.g. Mean 1 S.D. N =26 2 S.E.  17.5-20.1 18.7 0.82 0.32  79-4-89.2 83.2 2.99 1.18  75.1-90.9 82.3 4.45 1.74  2 4 . 1 - •26.1 25.1 0.52 0.20  7.7-9.1 8.3 0.37 0.15  Table I.  (Continued)  Measurement d a t a p l o t t e d i n b a r diagrams ( i n  mms.)  I n t . L.* I n t . W. Weight (gms.) Hind Foot L. Adult Range ^ P . l . n . Mean 1 S.D. N =35 2 S.E.  6.8-9-0 7.5 0.51 0.17  2.1-3.4 2.9 0.33 0.11  i5.lO-3i.25 20.7 3.26 1.10  19.4-22.0 20.5 0.70 0.24  Adult Range ? P . l . n . Mean 1 S.D. N = 33 2 S.E.  6.8-8.3 7-5 0.42 0.15  2 . 0 - 3 . 6 13.16-25.46 2.8 19.3 O.36 1.04 0.13 O.36 •  18.4-22.0 20.3 0.76 0.26  Adult Range <?P.m.£. Mean 1 S.D. N = 22 2 S.E.  8.I-9.8 9.2 0.32 0.15  2 . 3 - 3 . 9 15-13-24.27 3.1 18.8 0.33 2.43 0.15 1.10  20.5-22.2 21.3 O.53 0.24  Adult Range ?P.m.£. Mean 1 S.D. N=31 2 S.E.  7.9-9.9 9-1 O.38 0.14  2.5-3-9 3.0 0.27 0.05  13.39-23.92 18.9 3.04 1.08  20.0-22.1 21.0 0.52 0.19  Subad. Range cfP.m.^. Mean 1 S.D. N = 22 2 S.E.  8.1-9.7 8.9 0.38 0.17  2.5-3.9 3.1 0.26 0.12  12.13-19-79 16.1 1.68 0.76  20.0-22.2 21.0 0.65 0.30  Subad. Range ?P.m.£. Mean 1 S.D. N=26 2 S.E.  8.I-9.8 9-0 0.36 0.14  2.5-3.3 3.0 0.25 0.05  13-18-18.81 16.2 1.49 O.58  20.0-21.8 21.0 0.49 0.19  i n t e r p a r i e t a l l e n g t h , i n t e r p a r i e t a l width.  54.  Table I I .  " t " v a l u e s c a l c u l a t e d i n comparisons o f body p a r t s Animals Compared  E a r L.  Body L.  T a i l L.  Ad. c f P . l . n . N=35  X  Ad. cfp.m.g. N =22  14.534  3.441 *#  14.399  Ad. d P . l . n . N =35  X  Ad.+Subad.S P.m.g. N=44  23-925  0.582 Nonsig.  19.458  Ad.9 P . l . n . N=33  X  Ad.9 P.m.g. N-31  20.532  3.881  19.328  Ad. 9 P . l . n . N=33  X  Ad. + Subad.9 P.m.g. N=57  0.221 Nonsig.  25.164  Ad.cf+^P.l.n. N = 68  X  Ad.cf+9P.m.g. N=53  51.436  10.987  49.352  Ad. cTp.l.n. N=35  X  Ad. 9 P . l . n . N=33  1.664  1.400  1.611  Nonsig.  Nonsig.  Nonsig.  Ad.cf P.m.g. N=22  X  Ad.$ P.m.g. N=31  0.071 Nonsig.  0.595 Nonsig.  0.000 Nonsig.  Subad.2 P.m.g. N= 26  0.415 Nonsig.  0.651 Nonsig.  0.471 Nonsig.  Ad.+ Subad. £ P.m.g. N = 57  1.967  Nonsig.  1.425 Nonsig.  Subad.cfp.m.g.' X N=-22  " ~  Ad.f Subad.d" P.m.g. N =44 **  p i 0.005  ***  P^ 0 . 0 0 1  X  9.492  1.778  Nonsig.  4HHfr  55.  Table I I .  (Continued)  " t " v a l u e s c a l c u l a t e d i n Comparisons o f body p a r t s . Animals Compared Ad.0P.l.n. N=35  X  Ad.cfp.m.g. N = 22  Ad.0* P . l . n . N = 35  X  Ad.9 P.l.n. N=33  S k u l l L.  R o s t r a l L.  Int. L./  1.840 Nonsig.  1.682 Nonsig.  19.051  Ad.-1- Subad.cf P.m.g. N=44  5.136 ##*  6.088  30.096  X  A d . 9 P.m.g. N =31  3.269  0.461 Nonsig.  23-278  Ad.9 P.l.n. N = 33  X  Ad. -4- Subad. 9 P.m.g. N=57  7.410 *#•»«•  4^738  34.782  Ad.c?-t-?P.l.n. N=68  X  Ad.d'+?P.m.£. N = 53  7.326  3-722  19.308  Ad.cTP.l.n. N=35  X  Ad.9 P . l . n . N=33  0.000 Nonsig.  0.787 Nonsig.  0.356 Nonsig.  Ad'. c f P . m . g . N = 22  X  Ad. 9 P . m . g . N = 31  0.684 Nonsig.  0.000 Nonsig.  0.484 Nonsig.  Subad.3 P.m.g. N = 22  X  Subad.9 P.m.g. N = 26  0.606 Nonsig.  0.940 Nonsig.  O.517 Nonsig.  Ad.4- Subad.& P.m.g. N=44  X  Ad. 9 Subad.? P.m.g. N=57  1.898 Nonsig.  1.445 Nonsig.  •0.085 Nonsig.  / ** ***  interparietal p i 0.005 P ^ 0.001  length  Table I I I .  Data p l o t t e d I n b a r diagrams o f i n d i c e s . E.L. Adult JP.l.n. N  = 46  Adult °P.l.n. N=46 Adult c?P.m.g_. N =22 Adult ?P.m.g. N=31 Subad. dP.m.g. N =22 Subad. ijlP.m.g. N = 26 *  X T.L. X I.L. SK.L.  E.L.  X T.L. X I . L . R.L.  Range Mean 1 S.D. 2 S.E.  2.66-4.09 3.45 0.37 0.125  7.98-12.47 10.43 1.52 0.446  Range Mean 1 S.D. 2 S.E.  2.57-4.01 3.37 0.34. 0.119  7.39-12.50 10.47 1.19 0.350  Range Mean 1 S.D. 2 S.E.  4.83-7.00 5-77 0.55 0.250  15.25-21.20 17.60 1.53 0.696  Range Mean 1 S.D. 2 S.E.  4.54-7.14 5.80 0.64 0.228  14.54-21.93 17.62 1.92 0.686  Range Mean 1 S.D. 2 S.E.  4.26-6.50 5.39 O.58 0.264  12.83-20.75 16.70 1.87 O.850  Range Mean 1 S.D. 2 S.E.  4.76-6.14 5.52 0.38 0.149  14.26-18.84 16.73 1.31 0.514  E.L.= ear length, T.L«= t a i l length, l e n g t h , SK.L.= s k u l l l e n g t h .  I.L.= i n t e r p a r i e t a l  57.  Table IV. Breeding Length o f Time P a i r e d 67 d a y s  data No. o f Litters  Pairs  2  P.l.n.c/ X P.l.n.3  No. i n Litter 4  5 73 d a y s  P.l.n.d* X P.l.n. 9  365+  days  P.m.o.C^ X P . l . n .  365+  days  P.m.a.cf x P . l . n . $  365f  days  P.l.n.C^ X P.m.a.£  365+  days  P.l.n.<^* X P.m.  1  5  1  3  spp.^  P.m.o. cT  365 + d a y s  P.m.a.  3 6 5 + days  P.m.sppJ^X P.m. spp.+  71+-  days  P.l.n. ^ X  P.l.n.^  71-1-  days  P . l . n . X  P.m.g.^  71+  days  P.m.g.C^* X P.m.g.? P.m.g.  4  9  365+ days  71+- d a y s  x  1  Jr.m.o.  3-  P.m.a.Tr:  . n. 4 -  -  P . l . n . = P. l e u c o p u s n o v e b o r a c e n s i s P.m.o.= P. m a n i c u l a t u s o r e a s P.m.a.= P. m a n i c u l a t u s a u s t e r u s P.m.g.= P. m a n i c u l a t u s g r a c i l i s P.m. spp.=P. m a n i c u l a t u s ( M a n d a r t e I s l a n d , B r i t i s h  Columbia)  58.  Table  V.  Results  Expt. No. Tester  o f odor d i s c r i m i n a t i o n experiments.  Odor  Minutes Spent i n Arm  % of Total Time i n Arms  4 6 10 *  1.2 2.7 19.1 *  5.2 11.7 83.1 *  6 6  0.7 1.6 * 1.3  18.5 44.0 * 37.5  Control o*and 9 P.m.g. Oand ^ P . l . n .  28 33 * 23 31 * 25 30  5.7 * 3-3 4.0 31.5 * 8.0 10.8  44.0 * 25.0 31.0 62.6 * 15.9 21.5  Control  30 *  42.3 * 2.3 5.0  85.3 * 4.6 10.1  0.5 2.1 7.4 *  5.4 20.8 73.8 *  1.8 36.2 * 7.6  4.0 79-3 * 16.7  1.  9P.l.n.  Control c£and 9 P.m.g. oand 9 P.l.n.  2.  cfp.rn.g_.  Residual control R e s i d u a l cfand 9 P.m.g. R e s i d u a l cfand 9 P . l . n .  3.  cf P.m.g.  Control <£and 9 P.m.g. "and 9 P.l.n.  4.  $P.m.g.  5.  ?P.l.n.  9P.m.g. eTP.m.g.  6.  cf P . l . n .  Control <£and 9 P.m.g. o a n d 9- P . l . n .  7.  9 P.m.g.  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