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UBC Theses and Dissertations

Development of the Columbian black-tailed deer (Odocoileus hemionus columbianus) during the fetal period Ommundsen, Peter D. 1967

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DEVELOPMENT  OF T H E COLUMBIAN  (ODOCOILEUS  HEMIONUS  DURING  BLACK-TAILED  DEER  COLUMBIANUS)  THE F E T A L PERIOD  by  PETER B.Sc,  A  University  D.  OMMUNDSEN  of British  Columbia,  1964  THESIS SUBMITTED IN PARTIAL FULFILMENT THE  REQUIREMENTS FOR MASTER  in  OF  THE DEGREE  OF  OF  SCIENCE  t h e Department of Zoology  We  accept  required  THE  this  thesis  as conforming  to the  standard  U N I V E R S I T Y OF B R I T I S H COLUMBIA May,  19 67  In  presenting  for that  this  an a d v a n c e d the  study. thesis  degree  Library  Department  University  it  freely  of  representatives..  this  thesis  15,  for  permission.  Zoology  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a 19  67  for  the  Columbia  It  by t h e  requirements  Columbia,  I  reference  and  permission for extensive  or  by h i s  British  available  p u r p o s e s may be g r a n t e d  my w r i t t e n  May  f u l f i l m e n t of  scholarly  Department o f  Date  the  agree that  or p u b l i c a t i o n of without  at  in p a r t i a l  s h a l l make  I further for  thesis  copying of  gain  this  Head o f my  is understood that  financial  agree  shall  not  be  copying allowed  ABSTRACT  A  series  deer  (Odocoileus  From  the  data  probable tive A  growth  of  morphological  growing  structures  portion  of  orbital  region.  segments in  of  limbs  grow  patterns and  the  of  the  ears  relation grows  length,  skull  faster  show  in  between  deer  (0.  h.  than  body  diameter  and  the  to  be  measured. the  patterns of  the  of  rela-  fetus.  based  faster  on  exter-  fore  limbs.  growth were  than  the  inter-  The  neck  abdomen.  Greater  observed  cranial  postorbital  rates.  the  fastest-  The  and  growth than  the  length.  preorbital  similar  black-tailed  The  grows hind  differences  between  this  deer  (0.  virginianus  deer  and  Rocky Mountain  in  subspecies  borealis) mule  hemionus).  Some s o u r c e s technical  and  reconstruct  outlined,  appear  relatively faster  relative  the  to  development  to  the  Northern white-tailed  than  the  black-tailed  examined  morphology  d e v e l o p m e n t was  in  In  circumference  in  during  of  skull  the  Columbian  characteristics.  auricles  the  the  possible  changes  of  of  c o l u m b i a n u s ) was  i t was  occur  stages  The  fetuses  obtained  that  of  88  hemionus  sequence  series  nal  of  problems,  of  variation  individual  were  examined,  differences,  including  differences  between  the  sexes  regions Males of  and  appear  were  the  relationship  to  found  develop  at  larger  morphological development.  higher  correlation  with  between  the  size  same r a t e  than  females  Some b o d y  structural  change  and  form.  in  different  at  the  same  dimensions than  did  had  Not  all  animals stage a  others.  much  iv  TABLE  OF  CONTENTS  PAGE ABSTRACT TABLE  i i  OF  CONTENTS  LIST  OF  TABLES  LIST  OF  FIGURES  iv v i v i i  ACKNOWLEDGEMENTS  v i i i  INTRODUCTION M A T E R I A L S AND  1 METHODS  4  1.  Fetal Material  4  2.  C o l l e c t i o n of M a t e r i a l  5  3.  Methods  6  4.  Examination  5.  Method  of Preservation of External  of Designating  Form  Levels  7 of S t r u c t u r a l  Development 6.  Measurements  7.  Weights  7 8 15  RESULTS A.  16 Evaluation  of Methods  1.  Sources  2.  Evaluation  3.  E f f e c t s of Preservation  16  of M a t e r i a l of External  16 Dimensions  20  on R e s u l t s  23  V  PAGE B.  C.  General  Description  28  1.  V i b r i s s a e and Pigment  2.  Outline  of External  3.  Patterns  o f Growth  Variability  During  Patterns  28  Development  30 57  Development  1.  Differential  Development  2.  Development  3.  Sex D i f f e r e n c e s  4.  The R e l a t i o n s h i p  o f Twins  65  of External  Features  and T r i p l e t s  65 74 74  Between  Size  and Form  DISCUSSION  78 81  1.  Technical  2.  Fetal  3.  Patterns  4.  V a r i a t i o n i n Development  89  CITED  94  LITERATURE APPENDIX  Problems  i n Studying  Material of Development  Development  81 82 84  98  vx  LIST  OF  TABLES,  TABLE I.  PAGE Measurements after  II.  of  a Series  Preservation  Patterns  of Growth  Relationship  Y  Forehead-rump III.  Patterns  Comparison Forms  V.  VI.  Y  VII.  Degree  Saline  25  Period  of  the  where X Represents 62  i n the F e t a l  = b  X^  Period  of  the  where X Represents  Length  of F e t a l  63  Growth  Patterns  i n Three 64  of Development  between  Development of  i n Ten  Different 72  Relationship  gical  and  of Deer  Comparison Regions The  X^  before  Length  Calcaneus-hoof IV.  i n Formol  i n the F e t a l  = b  o f Growth  Relationship  of Fetuses  in a  Size,  C o r r e l a t i o n between  phological  Development  Sex  Series  and  and  Morpholo-  of Fetuses  the State Various  of  77 Mor-  Dimensions  79  vii  LIST  OF  FIGURES  FIGURES 1 - 1 8  PAGE Growth  Relative  t o Forehead-rump  - 35  Growth  Relative  to Calcaneus-hoof  3 6 - 3 8  Growth  of  39  - 41  Growth  42  - 44  Growth  19  Length  -  9  133  134  -  136  of the Trunk  137  -  139  o f t h e Limbs  I  Region  H"  116  -  t h e Head  Length  9  4  7  0  - I  4  2  ACKNOWLEDGEMENTS  I wish of  Graduate  this and  I Zoology, study the  am who  and  and  Dr.  I.  generous  help  indebted  to  collected  provided  McT.  for providing  for providing  Cowan, the  facilities  throughout Mr.  almost  advice  and  D.  C.  of  opportunity and  the  the  Faculty  to  undertake  experimental  animals,  study.  Thomas  a l l of  Dean  of  the  Department  the  fetuses  used  in  assistance  i n many  aspects  of  this of  project. I wish  with  thank  Studies,  project, for his  to  the for  to  thank  my  l a b o r a t o r y work, the  t i m e he  spent  wife, and on  Mr.  Arlene, S.  computer  W.  for  considerable  Borden  for his  programming.  help  advice  INTRODUCTION  The  Columbian  columbianus) North  is  an  Inlet,  California  (Cowan  and  of  form  This  i s an  Guiguet,  the  slope  of  deer,  much  period.  numerous w o r k e r s  less  According  a t t e n t i o n has to  1965;  to  Cowan,  of  Monterey 1956).  It  and  an  under-  i s important  to  i t s manage-  the  area  prenatal  development  (Cervus  elaphus),  roe  (Capreolus  Ziegler  and  systematic  Bischoff.  on  capreolus).  prenatal  Th.  investigation  deer  of  the  century  Schufeldt  (1891) p u b l i s h e d  deer  (Rangifer  tarandus)  fetuses,  A  development  number  Odocoileus.  of  of  and  the  those  (Dama  19th of  Pockels,  most  studies  on  roe  observations  on  rein-  Petrov  (1964)  studied  the  elk.  i n v e s t i g a t i o n s have been Cheatum  deer  made t h e  in his  i n 1843.  (1578-1657)  Fallow  During  growth  prenatal  Harvey  Bischoff probably  of  the  included  deer  intra-uterine  to  deer  dama) a n d  observations  given  William  red  century,  been  studied postnatal  (1936),  pregnant deer  have  Meyer  dissected  genus  Pacific  Columbia  in this  biology  investigation  hemionus  subspecies. Although  of  on  (Odocoileus  British  of w i l d l i f e  i t s reproductive  deer  strip  Rivers  important  ment.  narrow  from  standing  that  inhabits a  America  County,  black-tailed  and  Morton  (1942,  conducted 1946)  on  the  established  2  a  prenatal  growth  (Odocoileus the  fetal  ages  the  of  development  of  (Odocoileus  tics  were  the  subspecies and  fetus  of  age.  (1950) and  Hudson  the  (19 5 6)  Rocky  contributions  studies  of  studied  (1959)  pubon  Mountain  Developmental  Other  deer  characterized  Browman  study by  hemionus).  fetal  current  on  the mule  characterisprenatal  Segelquist  There  investigation treats  is little  the  embryonic  the  systems.  major  organ  minor fetal  smallest  period animal  discernible.  The of  available for  represented was  an  in  regarding  be  (19 66)  and  period  prenatal  features. to  i n which  adult with  (hoofs, a  as  of of  life. one  of  development  i s one  designated  study  animal  of  period  methods  considered  external  the  the  i n c l u d i n g the  fetal  arbitrarily  only  periods  can  organization,  was  This  the  period  differentiation  features  mm.  agreement  the d i v i s i o n s between  and  24.2  and  Armstrong  Hudson  a previous  differentiation  nal  same  life.  hemious  include  fetus.  Generally,  the  Northern white-tailed  (1966).  defining  with  the  the  embryo  r e l a t e d to  The the  fetal  the  deer  Nellis  of  r e s u l t s of  Odocoileus  for  virginianus borealis).  development  different lished  curve  of  growth,  In  this  include  most  of  ears,  crown-rump  of  case,  the  the  exter-  etc.)  were  length  of  3  The  objectives of  1.  To  describe during  2.  To  this  investigation  the changes  i n morphology  the development  describe relative  the changes growth  were:  of  the  i n size  that occur  that  occur  fetus. and  during  the patterns the  of  fetal  period. 3.  To  establish based  4.  To  on  evaluate  a  series  of  stages  of  external morphological sources  of  variation  development criteria.  in fetal  develop-  ment . 5.  To  outline  some t e c h n i c a l  procedures  i n studying  development. 6.  To  compare with  the f e t a l  that of other  development forms.  of  this  subspecies  4 MATERIALS  1.  A  total  a v a i l a b l e f o rstudy.  sets  of twins,  of  METHODS  MATERIAL  o f Columbian  These  consisted  and one s e t o f t r i p l e t s .  b l a c k - t a i l e d deer o f 23 s i n g l e t o n s ,  A group  d e t e r m i n a b l e s e x w a s m a d e u p o f 38 f e m a l e s Specimens  terms  1.  FETAL  o f 88 f e t u s e s  were  AND  of date  Animals  o f unknown  of collection,  o f 84  and 46  31  animals  males.  age a r e d i s t r i b u t e d i n time, i n as  follows:  c o l l e c t e d a t Goldstream,  Vancouver  Island,  British  Columbia. 1940 2.  Animals 1943  3.  Animals 19 64  January  One s i n g l e t o n  c o l l e c t e d a t Shellburn, April  One  and one s e t o f twins. British  singleton.  c o l l e c t e d a t Northwest  Bay, V a n c o u v e r  January  Four  February  Two s i n g l e t o n s  March  Two s i n g l e t o n s , one  Columbia.  singletons  s e tof  Island.  and one s e t o f t w i n s . a n d two s e t s seven  sets  of twins. o f twins, and  triplets.  May  Four  singletons  June  Two s i n g l e t o n s  19 65  January  Two s e t s  1966  April  Five  singletons  May  Two  singletons.  and e l e v e n sets and four  sets  o f twins.  of twins.  of twins and one s e t o f twins.  5  Known-age Columbia in  animals  were  Wildlife  January  19 66  Research  and  Known-age predetermined  2^-year-old breeding life  serious  20  long  lengthwise  from  the  December  the buck the  19 6 5 . or  and  The  tein  The  antlers the wide,  A  collected 19 6 6 .  available  removed  wall to  to  does  1 ^ - y e a r - o l d doe,  from  for a  until  Wild-  the buck used  the  once  was  introduce the  week  day in  does,  then  constantly while  about  every  second  pen,  to  extending  d i s t a n c e of  the buck  a  for  Columbia  b r e e d i n g pen  the breeding  animals  was  sacrificing  a partition  19 65, was  British  i n February  British  The  with one  procedure  the  were  exposed  into  of  does.  October  together, observe  obtained by  University  to  twins  collected  breeding.  center of  of  set of  of  MATERIAL  the  does were  week  University  1^-year-old buck were  feet  the  The  introduce  together  in  pen. Between  pen  12  from  last  separately  at  OF  were  after  injury  and  feet.  a  Unit.  prevent  eight  and  purposes  Research  feet  specimens  the  One  s e t was  COLLECTION  times  doe  from  Unit.  another  2.  at  obtained  20  feet  exposures,  long  supplemented  and  four  cereal  each feet  animal wide.  ration,  was The  kept deer  Buckerfields  in a were  U.B.C.  separate fed  a  3 657,  proand  6  the  food  consumption  of  each  animals  were weighed  at  one-week  doe  was  sacrificed  was  sacrificed  on  Fetuses 19 66  from w i l d  couver from of  the  28  of  18  February,  unknown  British  Coastal  Pacific  19 66  and  daily.  The the  The  1^-year-old  2^-year-old  doe  19 66. collected during  does were  shot  Columbia.  Western  Coastal  age  recorded  intervals.  January,  pregnant  Island,  the  on  a n i m a l was  The  Hemlock  and  Mesothermal  at  19 64,  Northwest  animals Coastal  Forest  19 65  Bay,  and  Van-  were c o l l e c t e d Douglas-fir  Region  Zones  (Krajina,  19 65) .  3.  Fetuses freezer, of  the  being  formol in  stored  in  had  their  penetration into  the  of  body  formol with  a  abdomens the  PRESERVATION  on  ice,  in  an  electric  several  hours  stored.on  i c e was  placed  dry  freezers  several f o r more  days. than  of  No five  the  in  death  electric  material months  before  saline. forehead-rump  length  opened w i t h  small  preservative.  cavities  dry  saline within  saline within  electric  Animals mm.  preserved  Material  or  stored  were  i n formol  doe.  freezers was  or  METHODS OF  of  some o f  a  Formol these  greater slit  to  s a l i n e was fetuses.  than allow  300 faster  injected  7  4.  Every  EXAMINATION  f e t u s was  given  and  a l l o b s e r v a t i o n s were  was  collected  such of  as  the  hair  general body integument,  patterns  structure auricles of  the  and  of of  and the  the  mouth,  external antler  regarding  form  on  FORM  external  data  and  proportions, vibrissae,  sheets.  preorbital  Information  characteristics  flexion,  thickness  emergence  colours, pigmentation and  examination,  of  gland,  the  c o n d i t i o n of  and  e x t e r n a l a u d i t o r y meatuses,  tongue  and  teeth,  of  brain  the  and  s t r u c t u r e of  skull the  bones,  genital  hair,  skin,  ears  c o n d i t i o n of  of  the  structure  external  condition of organs,  the  skin  nares, the  glands  limbs. w e r e made u s i n g  microscope  METHODS OF  For to  careful  recorded  form  hair  EXTERNAL  external morphological  eyelids  primordia,  dissecting  5.  a  emergence of  Observations a  OF  the  facilitate  with  LEVELS  purposes  this  OF  animals  appearance,  regard  without  the  (assessed to  STRUCTURAL  study,  d i s c u s s i o n s comparing  development between  10-power hand  lens  a maximum m a g n i f i c a t i o n o f  DESIGNATING  of  a  size).  and 84X.  DEVELOPMENT  s e v e r a l terms  are  used  extent  of  morphological  i n terms  of  morphological  8 Rank  i s a numerical  individual  compared  estimate  t o any o t h e r  Single-trait development to  Mean  the  level mean  rank  rank  o f the s i n g l e - t r a i t ranks  stage  o f development,  characteristics.  development  taken  device  of animals  series  on each  stage  fetus  uniformity  of results,  before  being  refers  with  regard  in a  measured.  This i s  set of  to a period rank,  At a  given  morphologi-  i n the total  conversely, i s  the relative  levels  of  group.  MEASUREMENTS  and c i r c u m f e r e n t i a l on d a t a  to the nearest each  rela-  individual.  development.  A  f o r comparing  and r e c o r d e d  recorded  of that  of the fetus.  of linear  ments were  tion  structural  o f an i n d i v i d u a l .  period during  6.  A  of  of the overall  a f e t u s has a s p e c i f i c A  sequence  a rating  level  to others,  estimate  development  i s a characteristic  merely  individual.  relative  i s a numerical  of structural  developmental  o f an  characteristic.  Stage  cal  o f development  i s the numerical  o f one i n d i v i d u a l  one s p e c i f i c  tive  of the l e v e l  animal I t was  sheets.  .1 mm. was  measurements  placed  A l l measure-  In order  placed  was  to  maintain  i n t h e same  on i t s r i g h t  posi-  side,  on  9  a flat  surface with  the examiner, from  i t s back along a s t r a i g h t  and i t s l e g s extended.  c u r l i n g backwards.  line,  The h e a d was  away  prevented  A l l measurements were t a k e n  r e f e r e n c e p o i n t s t h a t c o u l d be l o c a t e d  externally.  from  using No  dis-  s e c t i o n was r e q u i r e d . The  measurements u s e d w e r e c h o s e n  f o rseveral  1.  To d e t e r m i n e  2.  To p r o v i d e d a t a on t h e g r o w t h p a t t e r n s o f t h e s p e c i e s .  3.  To s t u d y t h e e f f e c t s  4.  To e v a l u a t e t h e u s e o f e a c h m e a s u r e d a t t r i b u t e of  t h e accuracy w i t h which each  purposes:  could be taken.  of preservative f l u i d . as an i n d e x  development.  5.  To a s s e s s  their  value i n f i e l d  6.  To p e r m i t  comparison  work.  of data with that of previous i n -  vestigators . All  o f t h e f o l l o w i n g measurements w e r e t e s t e d .  were v e r y d i f f i c u l t  Some  t o o b t a i n w i t h r e a s o n a b l e p r e c i s i o n and  were soon d i s c o n t i n u e d .  LINEAR MEASUREMENTS  T h e s e a r e measurements o f s t r a i g h t structural points, and  a meter  stick.  taken w i t h  a vernier  d i s t a n c e b e t w e e n two  caliper,  steel  calipers,  10  General  1.  Crown-rump  length  cephalon 2.  hump  Forehead-rump section  of  to  the  length the  tuberosity  i s the  of  from  p o s t e r i o r end  i s the  coronal the  distance  of  distance  and  the the  from  sagittal  top  of  the  mesen-  animal.  the  area  sutures  to  of  inter-  the  ischium.  Head 1.  2.  Head  length  i s the  lamboidal  crest.  Head h e i g h t  i s the  incisors 3.  to  Face height  Diameter taken  5.  at at  the  of  the  t i p of  distance  from  the  lateral  side  base  the  joining i s the  the  the  the  inner  of  straight  i s the  the  lips  distance  of  diameter  to  of  the  chin  to  the of  external  distance  the  of  corners  transverse  mouth w i t h  nose  the  two  the  a  eyes.  head  nares.  between  the  two  closed. between  the  right  and  entocanthion.  Diameter and  from  p o s t e r i o r end i s the  the  crest.  distance  Interocular breadth left  7.  nares  Mouth b r e a d t h corners  6.  line  from  lamboidal  i s the  projected 4.  the  distance  of  orbit  i s the  entocanthion.  distance  between  the  ectocanthion  11  8.  Bizygomatic front  9.  11.  Ear  t i p of  the  vertical  of  the  skull  directly  in  straight  d i s t a n c e between  plane  taken of  with  the  the  along  i s the  the  the  proximal  auricle  notch  held  and  along  head.  greatest width  held  d i s t a n c e between  notches.  ear,  i s the  auricle  i s the  ear  the  Preoptic length  the  of  the  auricle,  longitudinal  d i s t a n c e from  the  plane t i p of  taken of  with  the  the  head.  nose  to  entocanthion.  P o s t o p t i c l e n g t h i s the the  14.  left  the  the 13.  breadth  and  Ear width  i s the breadth  ears.  length i s the  the 12.  the  Biauricular right  10.  of  breadth  proximal  Cranial optic  length  notch  distance from  of  i s the  the  the  ectocanthion  to  ear.  diameter  of  the  orbit  plus  the  post-  length.  Trunk  1.  Total s p i n e to  2.  3.  the  length  t i p of  i s the  the  tail,  Precaudal  spine  length  crest  the  posterior  to  Shoulder  point to  cromion ischium.  process  d i s t a n c e from excluding  i s the end  p i n bone of  the  the  the  i s the  scapula  the  lambdoidal  sacrum.  d i s t a n c e from to  crest  hairs.  d i s t a n c e from of  lambdoidal  the  the  tuberosity  metaof  the  12  4.  S h o u l d e r p o i n t t o h i p bone i s t h e d i s t a n c e f r o m t h e metacromion process of the scapula t o the c r e s t of the i l i u m .  5.  W i t h e r s t o t a i l h e a d i s t h e d i s t a n c e f r o m a p o i n t mid-way b e t w e e n t h e v e r t e b r a l b o r d e r s o f t h e two s c a p u l a e t o t h e p o s t e r i o r end o f t h e sacrum.  6.  H i p bone t o p i n bone i s t h e d i s t a n c e between t h e c r e s t o f t h e i l i u m and t h e t u b e r o s i t y o f t h e i s c h i u m .  7.  Shoulder b r e a d t h i s t h e d i s t a n c e between the metacromion process of each s c a p u l a .  8.  Neck l e n g t h i s t h e d i s t a n c e f r o m t h e m e t a c r o m i o n p r o c e s s o f the scapula to the a t l a s .  9.  Transverse chest diameter i s the t r a n s v e r s e diameter of the chest d i r e c t l y behind the forelegs.  10.  S a g i t t a l chest diameter i s the depth of the chest i n the m i d - s a g i t t a l plane d i r e c t l y behind the forelegs.  11.  Dorsal t a i l of  12.  l e n g t h i s t h e d i s t a n c e b e t w e e n t h e t i p and h e a d  the t a i l .  Ventral t a i l tail  l e n g t h i s t h e d i s t a n c e between the t i p o f t h e  and t h e v e n t r a l j u n c t i o n o f t h e t a i l  and rump.  13.  H i p w i d t h i s t h e d i s t a n c e b e t w e e n t h e c r e s t s o f t h e two  14.  B i t r o c h a n t e r i c diameter i s the d i s t a n c e between t h e r i g h t and l e f t f e m o r a l t r o c h a n t e r s .  ilia.  13  Limbs  1.  Length of  o f humerus  the greater  olecranon 2.  Length to  3.  4.  5.  o f femur  of hind  of hind  calcaneus 7.  the  These structure  thread  the olecranon  process  between  of the foot from  the hoof  point  to theleg.  the greater  trochanter  t o the hoof  foot  the tubercle of  point.  i s the distance  length  from  from  the t i p of the  point.  i s the distance  to the distal  from  the t i p of the  edge o f t h e p r o x i m a l b e v e l  of  hoof.  CIRCUMFERENTIAL  the  of the  flexure.  t o the hoof  Calcaneus-hoof calcaneus  junction  l e gi s the distance  tibiofibula  Length  from  i s the distance  i s the distance  the patellar  Length  side  side  point.  of forefoot  Length  the proximal  to the posterior  i s the distance  the posterior  the 6.  of foreleg  Length  to  tuberosity  from  process.  t h e hoof  and  i s the distance  with  MEASUREMENTS  measurements concerned  vernier  a r e taken by passing  and then measuring  c a l i p e r s o r a meter  a thread  the length  stick.  Each  around  of the measure-  14  ment was t a k e n  three  times  a n d a mean o b t a i n e d  f o rthe final  result.  General  1-  Contour the  length  i s taken  external nares  from  t h e t i p o f t h e nose  between  to the t i p of the t a i l .  Head  1.  Head w i d t h over  i s taken  the cranial  from  the posterior  corner  dome t o t h e p o s t e r i o r  o f one eye,  corner  of the  other eye. 2.  3.  H o r i z o n t a l head the  smooth prominence  and  i n front  Vertical  head  cranial 4.  Mouth  Head  of  contour  across  above  the base of t h e eyebrows  i s t h e d i s t a n c e around t h e  the posterior  i s taken  between  corner  o f each eye.  t h e two c o r n e r s  of the  the lips. length  t h e nose  around  of the ears.  length  along  i s taken  of the forehead,  circumference  dome  cleft  mouth 5.  circumference  i s t h e contour  t o the lambdoidal  o f the head  from  the t i p  crest.  Trunk  1.  MedigA' neck between  circumference  t h e metacromion  i s taken process  at a point  mid-way  of the scapula  and  atlas.  15  2.  Chest  circumference  3.  Abdomen  i s taken  circumference  anterior  i s taken  to the umbilical  7.  The  total after  formol  w e i g h e d when  fresh  as w e l l  preservative to test  with  paper  than  500  balance. the  towels  gram  on  the  forelegs.  immediately  cord.  was  saline  obtained  f o r changes.  to  more  an A v e r y  .1 g r a m than  pan  f o r each  preservation.  as a t i n t e r v a l s  grams were w e i g h e d  nearest  at a point  500  after  Fetuses on  fetus.  being were  stored dried  weighing  a Mettler  less  electric  grams w e r e w e i g h e d  balance.  A l l  Some w e r e  A l l animals  b e f o r e b e i n g weighed.  Those weighing  behind  WEIGHTS  body weight  were weighed  in  directly  to  16  RESULTS  A.  EVALUATION  The  OF  METHODS  information  investigation  derived  i s probably  not  fetal  development  in wild  these  data  compared  past  or  may  be  future,  techniques  and  of  is  methods  sources  of  material a  sampling  sets  breeding  material  for  important  and  such  factors  well  as  may  the  of  one  an  to  of  twin buck  and  SOURCES  OF  fetuses to  two  the  size,  conditions the  age  given  case.  results  Before  to  This  the  evaluation  including  of  known  possible  accuracy.  age  they  obtained useful  development It  p a r i t y of  the  were  Cultivating  procedure.  of  of  investigators,  must be  prenatal  and  this  MATERIAL  of  under which  development  other  lack  does.  i n v e s t i g a t i o n on  in  representative  i n each  useful  error  used  b l a c k - t a i l e d deer.  those  of  time-consuming as  influence  Columbian  discussion  animals  completely  selected  1.  after  the  careful consideration  bias,  Two  from  are  fetuses  is  is possible the bred  parents, and  (Hafez,  an that as  reared, 1963).  17  Information  obtained  may b e u s e f u l  Behavior  that pen  i n the pen together  animals.  only  Consequently,  When 1^-year-old behavior  sniff  down. animals  antlers  Following  periods  the chasing would  Usually the entire were  originally  not left  first  was  planned  the other  changed  so that  t h e buck.  introduced  placed  to the  subside  aggressive be-  rush  I f allowed  and chase t o remain  and t h e animals  p a t t e r n would  then  of injury  would l i e  be repeated.  f o r three hours  due t o t h e r i s k  h e r back.  d e s c r i b e d b y Cowan a n d  t h e buck would  o f time.  together, the  along  exhibiting  and snort  together  unsatisfactory,  pen, t h e f o l l o w i n g p a t t e r n o f  begin  this,  of a l l three  towards  was  planned  i n the breeding  i n the pen w i t h  removed,  When  he would  the crouch  doe f o r v a r i a b l e  together,  aggressive  was a l l o w e d  i t was  to be  t h e doe and r u b h i s muzzle  typically  (1961).  judged  the procedure  observed.  a few minutes  havior,  the  always  program  The p l a c i n g  was s o o n  doe i n t h e b r e e d i n g  was  buck would  Geist  t h e buck,  here  conditions.  together  daily.  d o e was v e r y  one doe a t a time  Within  hours  described  to mating  of the breeding  three  the 2%-year-old  two  prior  procedure  such  a n d two does b e p l a c e d  f o rat least  animals as  t h e days  the outset  t h e buck  the breeding  i n helping to control  during  At  from  daily  The  a s was  t o t h e doe  from  18  the  a t t a c k i n g buck  first  week  together  of the project  f o r more When  she  "striking,"  a l . (19 6 5 ) .  to  remain  minutes  t h e doe,  Mating  behavior  The  remain  to the 2^-year-old  the animals  doe,  Thomas  were r a r e l y  pen f o r more  on 16 November  a heavy  dismounting. further  apparent allowed  than  i n the behavior to enter  allowed 10  times  thrust,  attempts  together.  does  her.  and t e r m i n a t e d  pushing  then  t h e buck,  rubbed  her forward  He  pen  or i f the  changed  i n a stationary  t o mount  patterns  the breeding  to the 1%-year-old  She s t o o d  attempted  before  t o t h e two  introduced  h e r 12  He  exposed  19 65.  immediately  remained  was  i f t h e doe e n t e r e d  b u c k was  copulate with  no  to  pattern described by  i n the breeding  h e was  days.  with  allowed  the  i n t h e pen, f r e q u e n t l y a t t a c k i n g t h e  i f t h e b u c k was  i n which  he  introduced  a behavior  difference  ferent  and  were r a r e l y  After  daily.  described  PM  t o avoid him.  minutes.  Consequently,  together  No  order  10  ran wildly  et  before  than  trying they  t h e b u c k was  typically  buck by  or from  on  dif-  doe a t  3:00  position  attempted  to  t h e t w e l f t h mount and  her back with  immediately  h i s muzzle  t o m o u n t h e r i n t h e 30 m i n u t e s  a n d made  that  they  19  At  5:00  was  introduced  and  after  and  gave  PM  o n 16 November  t o the buck.  a few attempts  a t mounting,  mount h e r i n t h e r e m a i n i n g  placed  9:00  PM  i n the breeding t o mate w i t h  Behavior  during  In  buck  using  t h e same  cycle  l e n g t h o f 12-14  the  on 4 December  l^-year-old  casion  and he  Behavior  16 November  2^-year-old  when  attempts  t h e b u c k was  to  again  d o e , b u t h e made  they were  no  outlined  No  d o e was  further  occurred, matings  twice away  doe c o u l d n o r m a l l y  at  assuming  a  oequrred, a l -  attempted  from  t o mount  h i m on e a c h  oc-  attempts.  not easily  i n estrus,  to the  previously, until  have  the buck  i f t h e does  exposed  when n o t i n t h e b r e e d i n g  1^-year-old  her  mating  d i d n o t make f u r t h e r  19 65,  copulated with  f o r a s u c c e s s f u l mating  days.  position,  o f them.  c y c l e s would  19 65,  he  19 65,  each  doe b u t she w a l k e d  of the deer  The  The  estrus  i n a stationary  doe  minutes.  after  procedure  two more  the 2^-year-old  made n o f u r t h e r  on 16 November,  least  though  either  to allow  not conceived  30  pen w i t h  the days  order  He  on 16 November  attempts  had  She s t o o d  a terminal thrust.  At  19 65,  s h e was be  pen  approachable, unusually  touched  b u t on  inactive.  on t h e head b u t  20  not  closely  very  docile  November pen. she O  approached  When  and could be thoroughly  19 65,  When  i n her pen.  examined.  i n estrus  she entered  readily,  was  The buck  often very  could  towards  t o 16  the breeding  but after  that  time  t h e pen on s e v e r a l o c -  u s u a l l y be approached  aggressive  s h e was  Prior  she o f t e n had t o be f o r c e d t o enter  c o u l d n o t even be f o r c e d t o enter  casions.  i n estrus,  and handled b u t  the investigator.  Fetuses  The and  twin  were on  1^-year-old  male  f e t u s e s were  obtained  from  28 F e b r u a r y  ments  d o e was s a c r i f i c e d  of these  t h e 2Jg-year-old  1966.  Morphological  animals  2.  are given  by  than  their  studying  others.  taken  The v a l u e  repeatability,  d o e when  19 66  and a female s h e was  fetus  sacrificed  d e s c r i p t i o n s and measure-  i n other  sections.  DIMENSIONS  on t h e f e t u s e s  a r e o f more  o f t h e measurements  and i f accurate,  by their  i s  determined  usefulness i n  growth. The  One  A male  E V A L U A T I O N OF E X T E R N A L  Some m e a s u r e m e n t s value  obtained.  o n 18 J a n u a r y  factor  degree  of accuracy  i s t h e shape  i s governed b y s e v e r a l  i n which  an animal  was  factors.  preserved.  21  Many of  fetuses  their  Even  were  bodies  preserved  were  i f a l lanimals  fixed were  i n unnatural i n stretched  held  positions or twisted  t h e p o s i t i o n i n which they were  influence  on t h e a c c u r a c y may  i n some  fixed  cases  be l e s s  affected  of a circumference  may  for distortion  i n another  of  reference  ment  used  using  skeletal  o n how  crest  could  length, dorsal often of to  assess.  spine  length.  greatly  t h e femur  on these  These  length,  The g r e a t e r  obscured  by soft  No  de-  specific  as t h e l a m b d o i d a l difficult  to locate. of  the total  to t a i l  trochanter  tissues.  I f t h e measure-  are therefore  include  withers  of  of reference,  them.  h e a d was  structures  seldom taken.  precaudal tail  the t a i l  t h e measure-  (by p a l p i t a t i o n ) , i t  the points  t i s s u e covers  animals  based  and were  soft  plane  i s the points  the point  accuracy.  be c o n s i s t e n t l y designated  and i n most  Measurements value  much  to locate  linear  i n one  When  structures,  structures  than  accuracy  the dimension.  great  some  plane.  influence  two s u r f a c e  may b e m o r e d i f f i c u l t  structure  may  may b e l o c a t e d w i t h  i s taken  pending  that  to evaluate  i s taken between  reference ment  factor  being  Circumferential  The d i s t o r t i o n  Another  positions.  may h a v e  o f t h e measurements.  measurements. compensate  portions  i n a s i m i l a r manner when  examined,  measurements  and  limited  spine  head and  o f t h e femur i s  Consequently  and b i t r o c h a n t e r i c diameter were  often  the length  difficult  Many take  with  circumferential  accuracy  measuring  thread  Although deviation those amount  comparison the  along  t h e same p a t h w a y  many m e a s u r e m e n t s animal,  specimens  of subcutaneous  differences  same  they  may  tissue  occasion. with  little  n o t be comparable t o  and muscle  and t w i s t i n g  due t o d i s t o r t e d  to  of passing the  may b e r e p e a t e d  n o t be p o s s i b l e between  animal  on each  due t o i n d i v i d u a l  i n stretching may  are d i f f i c u l t  due t o t h e s u b j e c t i v e p r o b l e m  o n t h e same  of other  measurements  variation  i n the  c o v e r i n g bones,  and  o f form.  Indeed,  different  dimensions  morphology  resulting  valid of  from  preservation. During changes be  taken  the growth  a r e o f such  period.  width  and v e r t i c a l  taken  on animals  on  magnitude  i n t h e same  fetal  animals  category  head  hair,  hair  certain  comparable  areas  f o r only  a r e crown-rump  circumference.  lacking  Obviously,  that  structural  In t h i s  with  of the fetus,  damage  can cause  keratinized  terminal portions of the hoofs  measurements.  length,  a r e not comparable  and must b e g u a r d e d  other  a portion of the head  t o those  taken  due t o t h e i n c r e a s e i n s u r f a c e t h i c k n e s s .  structural  involving  measurements can  C e r t a i n measurements  mensuration  Measurements  morphological  the hoof  against.  inaccuracies i n  The s o f t  non-  are easily  broken.  p o i n t s w e r e made b e f o r e  taking  23  It certain the  should  small  change  be  that  growth  cannot  3.  EFFECTS  studied the  preservation fetuses be  may  given  to  following  such  error  of  reflect  twisted  or  Changes  i n weight  fresh and /  for  two  fetus  in a  months  gained  negligible  P R E S E R V A T I O N ON  (such  as  Patten  t h a t may  occur  reviewing  and  changes  by  accuracy, to  large  RESULTS  and  Philpott,  19 21)  after  dimensions  of  c o n s i d e r a t i o n must  quantitative  changes  of  precision.  in tissue  p r e s e r v a t i o n , and  induced  great  compared  and  Weights  the  dimensions  equal  material i n formol  changes  stretched  then  be  illustrate  i n measuring,  Five  measured w i t h  c h a n g e s when  fetal  the  be  fluids.  after  examples  preservation probably  in various  although  measured w i t h  OF  changes  change  may  be  Numerous w o r k e r s have  that  s t r u c t u r e s may  during  dimensions  noted  that  data.  occurred  saline.  after  These  data  preservative fluid,  induced  by  fixation  in  The  human a  position.  group  weighed  of  again  experienced  3.19%. The  s i x animals after  t h a t were weighed  remaining  a mean w e i g h t  range  of  loss  was  loss from  i n formol of  6.59%.  2.11%  to  when  saline One 14.44%.  24  S i x t e e n animals were weighed when f r e s h and weighed again a f t e r two years i n formol s a l i n e .  A l l l o s t weight,  with  a mean l o s s of 10.86% and a range of 1.9 7% to 15.87%. There appears  to be no r e l a t i o n s h i p between forehead-  rump l e n g t h and amount of weight change or between f r e s h weight per u n i t l e n g t h and amount of weight l o s s .  However, these data  suggest t h a t the l e n g t h of time a f e t u s i s p r e s e r v e d should be c o n s i d e r e d when e v a l u a t i n g the s i t u a t i o n i n the f r e s h  Changes i n l i n e a r and c i r c u m f e r e n t i a l  animal.  dimensions  Table I i l l u s t r a t e s the changes i n 27 measurements a f t e r formol s a l i n e p r e s e r v a t i o n . using a t - t e s t .  These changes were e v a l u a t e d  25 TABLE MEASUREMENTS BEFORE  AND  AFTER  OF A  I  S E R I E S OF  PRESERVATION  FETUSES  I N FORMOL  SALINE SIGNIFICANCE  MEASUREMENT  FETUS  #  BEFORE  AFTER  OF  CHANGE  Forehead-rump length  Head  Face  length  height  Diameter  at  Interorbital  Diameter  of  Bizygomatic  NOTE:  nares  breadth  orbit  breadth  Fetuses  1  78.4mm  74.5mm  2  81.2  76.2  3  205.0  192.5  4  204.0  187.5  1  28.6  25.5  2  27.0  25.9  3  68.3  68.0  4  66.5  67.8  3 4  28.7  32.5  26.8  31.9  1  5.3  5.6  2  5.2  6.0  3  15.4  15.9  4  15.3  16.0  1 2 3 4  10.9 10.9 31.8 31.0  12.0 13.2 31.0 31.2  1  5.  5.7  2  4,  4.5  3  11. 11.4  11.2  4 1 2 3 4  16.7 15.4 34.8 33.8  15.3 15.7 35.5 35 .2  numbered  one  months b e f o r e b e i n g three  = not  *  =  0.01<P<0.05.  **  =  P<0.01.  ns .  ns  ns ns.  ns. ns  11.4 ns ns  a n d two w e r e p r e s e r v e d remeasured.  and f o u r were p r e s e r v e d  ns.  ns .  significant.  Fetuses  f o r two  f o r four  numbered  and one h a l f  mon  26 TABLE  I  (continued) SIGNIFICANCE  MEASUREMENT Ear  length  Preoptic  length  Postoptic  Shoulder  length  to pin  Shoulder  Neck  FETUS  to hip  length  Transverse  OF  CHANGE  3  24,. 1mm  24,.  4  24,.0  25, .5  1  12..2  11, .4  ns.  2 3 4  11, .5 33. .3 31, .4  11, . 6 32 , .9  ns .  1  10. .4  9 .9 ,  2  10, .9  9,.7  3  31, .9  30, .5  4  30. .8  31, .0  1  45. .4  2  43. .4  39 ..1 4 1 . .1  1  33. .3  30. .6  2  33. .6  31. .5  3 4-  55. .0  40. .6  52..8  40. .5  3lTOTl  ns .  31, .4 ns. ns.  ns.  *  **  3  34..0  38..0  ns .  4  35. .4  37. .4  ns.  3  49 ..5 4 8 ..8  49 ..5  ns.  4  Hip  AFTER  chest  diameter  Ventral  BEFORE  chest  diameter  Sagittal  #  tail  length  width  50. .7  1 2  6..7  5..9  7..2  5..6  3  26. .0  22..8  4  26. .0  25. .2  3  30. .9  30. .6  4  32..2  33. .3  3 4  38.,2  38.,6  38.,4  38.,2  ns.  ns.  Biochanteric diameter  Length  o f humerus  ns.  ns.  27  TABLE  MEASUREMENT Length  Length  Length  Length  Length  Median  FETUS  of foreleg  of  forefoot  o f femur  of hind l e g  of hind  foot  1 2 3 4  (continued)  #  BEFORE  AFTER  28.8mm  27.Oram  27.4  26.7  93.2  89 .8  92.4  87.9  3  50.5  50. 6  4  49.7  49 .5  3  43.5  37.3  4  38.0  39.7  3  97.9  98.8  4  99 .9  97.0  3  73.8  71.2  4  72.2  71.4  3 4  82.5  82.5  71.0  75.0  3  136.5  141.0  4  141.0  141.0  3 4  146.0  143.0 144.0  SIGNIFICANCE OF CHANGE ns  ns  ns  ns.  ns  neck  circumf erence  Chest  I  circumference  ns  ns.  Abdominal circumference  139.0  ns  28  B.  GENERAL  DESCRIPTION  The patterns  general  o f development  1.  The  of the skin  hair  follicles)  development  provide  that  described  here.  VIBRISSAE  FOLLICLES  Vibrissae  sides  the  AND  vibrissae surfaces  lower  jaw.  patterns  r e s u l t i n g from  landmarks  Some  section.  PATTERNS  hair  special  may b e o b s e r v e d  from which  of the skin  ventral  useful  and  the  underlying  i n the study regions  of  coloura-  of the  vibrissae  as t h e f e t u s  grows a r e  VIBRISSAE  follicles  vibrissae  Submental  of vibrissae,  and t h e b a s i c  i n this  PIGMENT  ( i n many r e g i o n s  pigmentation  Mystacial  are discussed  of the fetus.  elevations,  of the fetus  V I B R I S S A E AND  appearance  tion  and  morphology  appear  emerge  occur  as s m a l l  vibrissae  on t h e d o r s a l  of the rostrum a r e found  white  or  i n the following  and l a t e r a l  regions:  anterior  and upper l i p .  on t h e a n t e r i o r ,  of the skin  spots  covering  lateral  the anterior  and end of  29  Interramal of  vibrissae  the lower  Malar the  Superciliary eyelid Ciliary  side  below  and never  found  on each  level  they  are located  the rami  of the ventral  half  of  more  side.  than  the  eyelids.  anterior  vibrissae  have  vibrissae  The v i b r i s s a e  of other  regions  i n number  and l e n g t h  as one c r i t e r i o n  imme-  to the eye.  of malar  of the  upper  of the head  and extend  interramal  of the  one p a i r  used  of development  above  t h e edges  on t h e s k i n  eyelid  three  increase  may b e  along  are found  than  directly  t o the eye.  lines  the lower  the vibrissae  grows,  form  more  found  on t h e s k i n  anterior  vibrissae  Never  between  o f the head.  vibrissae  vibrissae  diately  As  a r e found  and extend  Subciliary  on t h e s k i n  jaw.  vibrissae lateral  are found  been  have  been  a r e numerous.  as t h e  animal  f o r determining the  fetus.  PIGMENTATION  Inferior the  upper  Superior the ders  nasal  pigment  surface  of the nares  pigment.  i s present  l i p and v e n t r a l  nasal  dorsal  pigment  on t h e a n t e r i o r  anterior  extends  surface  of  nose.  between  the external  o f the nose  anterior  to the dorsal  border  nares  to the posterior of the inferior  from bornasal  30  The  superior  along The  the  The  the  ciliary  of  ment of  labial  i n any  line  a  series  example,  presence region of  gical  of  in  may  is a line  of pigment  that  extends  of pigment  be  used  OF  not  Balinsky  general,  poor  as  an  and  at the  intensity  indication  EXTERNAL  the as  site  of  of  the  pig-  level  DEVELOPMENT  general  a whole  of periods  for vertebrates.  features,  colour  development.  a series  are based  appears  fetus.  organism  importance  that  vibrissae.  absence,  OUTLINE  stages of  lists  here  stages).  extends  ciliary  or  the  f o r the  obvious  cribed  that  understanding of  development of  of pigment  line  the  2.  velopment  line  is a  of  development  An  is a line  lower l i p .  emergence  The  line  upper l i p .  inferior  along  labial  A  pattern  may  be  Witschi and  table  size  or  age  of  (1960) p o i n t s  criteria  readily (ages out  fetal  gained by (1956),  "normal  that  listed size the  studying  of  stages" stages  discernible  are  for indicating  The  de-  for  standard stages  t o e x p e r i m e n t a l work. m a i n l y on  of  des-  morpholo-  for only  and  is  age  extent of  two  are,  31  development.  He f a v o r s the use of e x t e r n a l m o r p h o l o g i c a l  p r o p e r t i e s f o r d e f i n i n g stages. The stages i n c l u d e d i n t h i s o u t l i n e of development are based o n l y on the m a t e r i a l a v a i l a b l e f o r examination. I, t h e r e f o r e , r e p r e s e n t s the s m a l l e s t f e t u s a v a i l a b l e .  Stage A more  f i n e l y r e f i n e d s e r i e s of stages w i l l p r o b a b l y be p o s s i b l e , when more specimens are c o l l e c t e d .  The d e s c r i p t i o n of each  stage of development i s based on what i s c o n s i d e r e d to be a r e p r e s e n t a t i v e animal.  These animals were chosen to i l l u s t r a t e  the s t a t e of development of the f e t u s a t a s e r i e s of i n s t a n t s d u r i n g growth and thus demonstrate the r e l a t i o n s h i p between d i f f e r e n t m o r p h o l o g i c a l changes as growth p r o g r e s s e s .  The  animals s e l e c t e d are spaced so as t o minimize the e f f e c t s of i n d i v i d u a l v a r i a t i o n o b s c u r i n g the g e n e r a l p a t t e r n .  The  s t r u c t u r e o f s e x - s p e c i f i c c h a r a c t e r s i s i n c l u d e d f o r b o t h sexes by r e f e r r a l to specimens c o n s i d e r e d t o be of comparable l e v e l s of development although of the o p p o s i t e sex to the chosen t y p i c a l specimens.  The measurements, however, are those of  the chosen t y p i c a l specimen. H a i r c o l o u r as d e s c r i b e d i s i n r e f e r e n c e to the g e n e r a l appearance of the pelage, r a t h e r than the i n d i v i d u a l h a i r s , many of which c o n t a i n s e v e r a l c o l o u r bands.  General  32  surface bands  hair  emerge  c o l o u r may during  thus  change  development,  i n a given  even  though  area  as  new  t h e same h a i r s  are  present.  STAGE  I.  Crown-rump  length,  8.3  mm.,  weight,  1.  In general  2 4 . 2 mm.,  1.5  form,  bizygomatic  grams. the animal  the letter  ventrally  and c a u d a l l y and l i e s  the  the ventral front  phalic,  "C."  i s flexed  resembles  with  breadth,  aspect  so t h a t i t  The muzzle  i s directed  i n apposition  of the thorax  hoofs.  Flexures  pontine,  cervical,  apparent dorsal,  and between  are the ceand  lumbo-  sacral . 2.  Regions  of the brain  telencephalon, The 3.  sent 4.  are very  forms  that  auditory covered  by  The nares  fold  of the ears  are directed  meatuses the  externally and  No  prominence.  eyelids  are pre-  i s evident.  are present ventrally.  as  auricles. but closed.  small  The e x t e r n a l  are closed, but not  are distinct  are the  myelencephalon.  a conspicuous  distinct.  and no p r e o r b i t a l  The a u r i c l e s folds  5.  mesencephalon  mesencephalon  The eyes  evident  completely  33  6.  The  mouth  7.  The  neural  plete  and  i s open  and  arches the  of  total  length.  8.  The  liver  prominence  9.  The  phallus  10.  The  hoofs  hind  foot  the  spinal  its  the  dew  i s less  than  are  not  com-  is distinguishable  i s very  claws  protrudes.  vertebrae  cord  i s l a r g e and  and  tongue  conspicuous.  curved are  for  caudad.  differentiated.  twice  the  length of  The the  tail. 11.  No  hair  or  12.  Vibrissae submental,  pigment follicles  II.  Forehead-rump 9.5 4.34 1.  mm.,  on  present  the  animal.  i n the  mystacial,  subciliary  and  areas.  length,  45.2  calcaneus-hoof  mm.,bizygomatic  length,  9.2  mm.,  breadth,  weight,  grams. In  general  form,  the  the  dorsal surface  The  muzzle  to  the  pontine 2.  are  i n t e r r a m a l , malar,  superciliary  STAGE  i s evident  and  flexure  region  and  the  lying  is directed  thorax  The  animal  of  the  along  hoofs.  curled, same  and The  most  of  plane.  lies  anterior  region of  the  distinct.  cerebral  mesencephalon  the  ventrally  front  i s not  i s not  forms  hemispheres a  is  conspicuous  evident prominence.  34  3.  The e y e l i d s open.  A  Pigment 4.  are p a r t i a l l y  distinct  preorbital line  i s visible  i n the region  i s  direction  cover  and completely  meatuses.  The meatuses  The nares  are closed.  6.  The s i d e s  o f t h e mouth  but does  not protrude  The n e u r a l plete  from  arches  externally partially noticeably T h e abdomen thorax.  by  i s open.  cord  may  f o r much  i n the thoracic i s more  The area  ventral audi-  membranes, The  tongue  t h e mouth.  be  are not  com-  distinguished  f o r i t sentire length. enclosed  i n a  are closed.  of the vertebrae  and t h e s p i n a l  iris.  the external  are closed  t h e f r o n t o f t h e mouth  present.  of the  are folded  5.  8.  b u t t h e eyes a r e  The a u r i c l e s o f t h e ears  tory  7.  formed  The c o r d i s  of i t slength,  most  region.  v e n t r a l l y extended  of the l i v e r  than the  i s distinct  ex-  ternally. 9.  The p h a l l u s genital of,  i s curved  swellings  but slightly  male,  b u t more  caudad.  are evident, posterior  anterior  Two  distinct  one on  each  to the phallus  i n the female.  side i n the  35  10.  One  pair  of  teats  phallus  on  each  11.  No  or  pigment  12.  Vibrissae follicles  hair  submental,  I I I . Forehead-rump 25.5  mm.,  14.5  grams.  1.  The  i s evident are  of  Age,  63  the  present  in  animal.  the  mystacial,  subciliary  and  mm.,  bizygomatic  length,  17.7  breadth,  mm.,  weight,  proportions  resemble  days.  body  an  on  malar,  74.5  calcaneus-hoof  those  the  areas.  length,  general  to  side.  interramal,  superciliary  STAGE  is situated lateral  form  adult.  and  The  body  axis  is  somewhat  curved. 2.  In  general  tal  region  steep nent are  The  of  distinct.  regions  eyelids  are  pigment  of  fold  to  feature  as  is  the  is quite  angle  evident 3.  form,  the the The  lateral of  the  formed the  iris  distinct.  head high  i s elongate. however,  and  The forms  mouth.  The  muzzle  head.  The  cerebral  f r o n t a l s and  fron-  is  a  a promi-  hemispheres  parietals  are  plates. eyes and  protrude the  eyes  is visible.  noticeably. are  closed.  A preorbital  The The  4.  The a u r i c l e s from  of the ears  the sides  ventrally meatuses  o f the head  and l a t e r a l l y .  The nares  6.  The mouth i s c o m p l e t e l y  7.  8.  t h e upper  The area  that  The p e n i s rected  directed  10.  arches  of the  of the l i v e r  of the  are  anterior are  present  straight  externally but i s confluent  i n form  The s c r o t u m united  and i s d i -  i s evident  swellings  p o s t e r i o r and d o r s a l l y .  as two  posterior  The c l i t o r i s i s prominent  and  I t has begun  to  form.  are conspicuous,  hind  i s greater  tail.  by  thorax.  but medially  on i t s a d u l t  foot  enclosed  vertebrae.  i s distinct  The limbs  the  more  completely  o f t h e abdomen  i s mainly  the penis.  take  auditory  The l i p s  L a t e r a l membranes  i s almost  ventrally.  distinct to  closed.  l i p extends  l i p .  ventral plane  with  The e x t e r n a l  the l i p s .  neural  the  9.  The lower  The s p i n a l c o r d the  but are directed  are closed.  distinct.  between  unfolded  are closed.  5.  than  are p a r t i a l l y  than  slender twice  and long.  the length  of  The  37  11.  No  hair  or  12.  Vibrissae  pigment follicles  submental,  13.  IV.  Two  present  20.8  teats  mm.,  42.88  are  present malar,  on  the in  animal.  the  mystacial,  subciliary,  each  side  of  and  the  ventral  length,  103.2  mm.,  bizygomatic  length,  2 7.3  mm.,  breadth, weight,  grams.  The  c e r e b r a l hemispheres  crown The  of  region  and  the  forms  each  aspect side  an  are  abrupt  of  the  are  4.  The  a u r i c l e s of  and  medially  and  the  are  closed.  The  distance  The  the  iris  the  nares  are  than  is  ears  a u r i c l e s when  is less  but  separated  pigment  head  of  head  The  the  angle  p a r i e t a l s are  3.  of  prominent.  The  between  the  mouth.  f r o n t a l s and  dorsal  5.  on  abdomen.  calcaneus-hoof  frontal  2.  interramal, areas.  Forehead-rump  1.  are  superciliary  posterior  STAGE  i s evident  the  closed.  by  distinct the a  on  dorsal  distinct  the  borders gap.  visible.  are  external  directed dorsally auditory  between  f o l d e d back length  of  the flat  one  meatuses distal against  auricle.  tips the  38 6.  The mouth i s c l o s e d and t h e l i p s are v e r y w e l l developed.  The lower l i p i s l a r g e and  extends  n o t i c e a b l y more a n t e r i o r t o t h e upper. 7.  The submental r e g i o n of the lower jaw i s p o o r l y developed, g i v i n g the a n t e r i o r end of t h e snout a p o i n t e d appearance when viewed from t h e s i d e .  8.  The p e n i s i s s t r a i g h t and d i r e c t e d  anteriorly.  The scrotum i s i n the form of a s m a l l b i l o b e d s t r u c t u r e p o s t e r i o r t o the p e n i s . 9.  A n t l e r p r i m o r d i a a r e p r e s e n t on the d o r s o - l a t e r a l a s p e c t s of t h e head i n b o t h sexes.  10.  M e t a t a r s a l glands are p r e s e n t on the l a t e r a l  sides  of the h i n d f e e t . 11. STAGE V.  No h a i r o r pigment i s e v i d e n t on t h e a n i m a l .  Forehead-rump l e n g t h , 187.5 35.2 mm., 259 grams. 1.  mm.,  bizygomatic breadth,  c a l c a n e u s - h o o f l e n g t h , 66.5 mm., Age,  103  weight,  days.  The head i s e l o n g a t e , t h e r o s t r u m i s prominent, and the f r o n t a l r e g i o n does not form a s t e e p a n g l e t o the mouth.  2.  The f r o n t a l s and p a r i e t a l s a r e p a l p a b l e over most o f t h e d o r s a l r e g i o n of t h e head b u t a s m a l l  39  but of  distinct their  The  of  visible The  dorsal  closed  sides  space  form  head.  The  a u r i c l e s of  the  ears  head  overlap  than  half  one  folded does  and  reach  opposite  side.  the  the  the The  nares  are  The  mouth  i s closed.  The  the  muzzle  side.  positions  apparent. tinctly tongue  The  distal  the  edges  on  is  medially of  one  auricle.  the  distance  border  of  auditory  the  iris  a  of  The  greater When  one  auricle  meatus  auditory  a blunt  giving  of  the  meatuses  the  incisors of  the  Horizontal defined.  region the  ridge  and  is  canines  lower  lips  ridges  on  of  the  anterior  a p p e a r a n c e when  periodontal  insides  poorly  submental  developed,  The  serrated. are  length  closed.  is well  the  of  of  for  external  The  of  the  directed  distal  external  closed.  jaw  are  other  length  are  lower  the  each  medially,  not  from  pigment  eyelids.  the  of  protuberances  the  auricles  end  large  through  against  most  borders.  eyes  the  separates  viewed distinct. are are the  not dis-  40  7.  The p e n i s is  of a pendulous  and  the  9.  of the genital  the  skin.  foot  with  white  light  spots.  brown  much o f  times  Very  as  long  present.  sae  are evident.  STAGE V I . F o r e h e a d - r u m p  malar  The i n -  as l i g h t  brown  to  distinguish. The a n t e r i o r  are faintly  are just  length,  pigmented. Subciliary  beginning  and i n t e r r a m a l  2 2 6 . 5 mm.,  brown  s u p e r i o r and  on t h e head.  A few m y s t a c i a l  length,  spots.  i s present.  vibrissae  calcaneus-hoof  grams.  line  on t h e  i s present,  are d i f f i c u l t  of the hoofs  short  are evident  white  are present  superciliary  mm.,  three  pigment  The l i g h t  ciliary  are  436.80  obscuring  i s evident  lines  regions  vibrissae  emerge.  with  pigment  labial  proximal  and  of pigmentation  i n colour,  nasal  inferior  Some  than  The s u p e r i o r n a s a l  ferior  40.7  i s large  I t i s the prominent  region,  i s greater  areas  mid-brown  10.  The c l i t o r i s  tail.  Several  A  and t h e scrotum  labia.  The h i n d as  nature.  i s directed posteriorly.  feature  8.  i s directed anteriorly  vibrissae  and submental  bizygomatic 8 9 . 5 mm.,  to  vibris-  breadth,  weight,  41  1.  The  dorsal borders  proximity. sition  2.  is  present.  The  pigment  The  of  the  the  closed  a u r i c l e s of  folded  onto  distal  border  ternal  auditory  head.  The  the  p a r i e t a l s are  f r o n t a l s appear  anteriorly.  through 3.  The  of  A  iris  anterior  i s not  ears  dorsal  of  in  readily  close  appofontanelle  visible  eyelids.  the  the  distinct  to be  in  one  directed medially  aspect  auricle  meatus  external  are  of  of  head.  The  reaches  to  the  ex-  other  side  of  the  the  auditory  the  meatuses  are  and  slightly  opened. 4.  The  nares  5.  The  periodontal  6.  are  somewhat ridge  is distinct.  of  individual incisors  at  different  the  lips  The  scrotum  than  the  portion  are  regions  and  of  the  ridge.  The  positions  can  be  The  detected  insides  of  serrated.  i s pendulous The  canines  the  distinctly  penis. of  opened.  and  clitoris  genitalia,  the  distinctly occupies labia  longer a  smaller  being  clearly  visible. 7.  The with  superior white  nasal  spots.  pigmentation  i s dark  The  nasal  inferior  brown  pigmentation  42  is  light  labial line  8.  9.  brown w i t h  line  hoofs  are  are  evident  Vibrissae  areas.  are  skin  region, the  on  the  ventral  and  1.  Most but  of the  the  skin  rump,  surfaces  of  head  and  tail  hair  may  be  surface.  limbs  are  seen  skin  the  white. emerging  spots  submental,  ciliary,  and  covering  the  distal  region  region,  of  the  umbilical  108.5  light  the On  white  bizygomatic  abdomen, and  of  cord,  neck.  length,  i s very  labial  portions  mystacial,  2 6 9 . 5 mm.,  ventral  the  to  lateral  calcaneus-hoof  grams.  the  thoracic  and  730  superior  trunk.  posterior  anterior  mm.,  the  the  abdomen  inferior  Some d o r s a l  in  ventral  length,  the  proximal  of  ventral the  The  superciliary,  detectable  femur,  and  The  present  subciliary  Forehead-rump 46.0  the  malar,  Hairs  spots.  pigmented.  interramal,  the  VII.  brown.  on  are  olecranon  STAGE  i s mid-brown  is light  the  white  mm.,  brown  ventral close  over  weight,  in  much o f  breadth,  colour, the  side  medial of  the  examination  much o f  the  body  2.  The  anterior  3.  The  pigment  the  closed  The  auricles  and The  4.  onto  the  dorsal aspect  distal  border  the  open.  the  light  is  very  of  the  completely  notice-  reach  forward,  just  to  the  the  distal  tips  posterior  bor-  eyes.  i s closed.  The  The  tooth  incisors  horizontal  and  r i d g e and canines  ridges  on  positions  are  the  very  tongue  defined.  spots.  brown w i t h distinct.  auricles  margins.  The  distinctly white  can  head.  are  When f o l d e d  white  is  the  e x t e r n a l a u d i t o r y meatuses  superior nasal  with  auricle  of  medially  opposite  developing  are w e l l  one  directed  the  the  mouth  through  of  auricles  of  of  are  e x t e r n a l a u d i t o r y meatus  The  the  The  visible  eyelids.  distinct.  6.  i s not  folded  ders  of  iris  ears  ably  The  the  distinct.  the  side.  5.  of  is  of  cover  of  fontanelle  spots  of  pigmentation  i s dark  The  nasal  white  spots.  pigmentation  The  skin  of  the  ears  i s brown w i t h  pigmented.  the  ciliary  The  proximal  are  inferior  brown  portions of Both  apparent  on  dorsal surfaces  the  d o r s a l and the  line  skin  of  white  hoofs  are  lateral the  trunk.  44  7.  H a i r on t h e body s u r f a c e i s s p a r s e , b u t most r e a d i l y e v i d e n t on t h e s k i n c o v e r i n g t h e o l e c r a non r e g i o n , t h e l a t e r a l d i s t a l r e g i o n o f t h e humerus,  l a t e r a l proximal r e g i o n of the ulna, the  p o s t e r i o r l a t e r a l r e g i o n o f t h e femur,  lateral  r e g i o n o f the t i b i a , t h e l a t e r a l b o r d e r s o f t h e v e n t r a l s u r f a c e of the thorax, a t h i n m i d - v e n t r a l l i n e between t h e t h o r a x and u m b i l i c a l c o r d and the r e g i o n s o f t h e a n t l e r p r i m o r d i a o f t h e head. Dark-coloured  h a i r s a r e b e g i n n i n g t o emerge over  much o f t h e body s u r f a c e . 8.  White h a i r s a r e apparent on t h e s k i n c o v e r i n g the medial surfaces of the legs i n the regions of the humerus and r a d i u s , and on t h e r e g i o n s o f t h e femur and t i b i a , on t h e p o s t e r i o r v e n t r a l r e g i o n s o f t h e abdomen, on t h e v e n t r a l s u r f a c e s o f t h e a u r i c l e s o f t h e ears and on t h e v e n t r a l s i d e of the head.  9.  Areas o f t h e crown, r o s t r u m , no e v i d e n c e o f emerging h a i r .  f e e t and t a i l show  45  STAGE V I I I .  Forehead-rump length, 52.0  mm.,  calcaneus-hoof  850  grams.  1.  In general with but  2.  varies  i n density i s closed.  mid-brown  and  The  mid-brown tion nasal 3.  The proximal  4.  Faint white  spots in  on  are present  the region  on  of the animal  for  the following white of the t a i l ,  hind  legs  down  surfaces  on  to the l e v e l  i s  nasal and  the  superior  white  spots.  sides  brown. of the White  of the  legs  femur. grey  except  the ventral  medial  are  pigmenta-  the skin  i s light  i s  light  a r e dark  and t h e  surfaces  of the hock  of the pectoral  of the d i s t a l  line  of the trunk.  areas:  abdomen,  labial  brown w i t h  o f t h e humerus  face  spots  the l a t e r a l  remainder  level  white  sparse  areas.  line  inferior  surfaces  grey  o f t h e mouth  of the hoofs  The  medial  labial  are present  and l a t e r a l  is light  inferior  i s dark  borders spots  The  weight,  i s generally  the roof  with  pigmentation  dorsal  The  breadth,  mm.,  in different  i n colour.  i s mid-brown  123.4  Hair  the superior  ridges  bizygomatic  the animal  areas.  The mouth  mm.,  length,  appearance,  some w h i t e  brown.  5.  284.0  limbs  end o f t h e r a d i u s ,  surof the  joint,  down a  to the  ventral  46  surface  o f t h e head  ventral  surfaces of the auricles  posterior surfaces hoofs,  6.  i s sparsely  surface. present  coloured  hairs  The h a i r  on t h e s k i n  antler  pletely  band  over  pigment  gland.  posterior  hairs  areas.  as does  that  i s beginning  of the  regions.  i s s h o r t b u t dense A  t o appear.  a n d comdark A  throat-  i s evident. areas, Bands  the proximal  ring  i t s  and w h i t e -  the underlying skin.  on t h e f e e t .  distinct  of  on t h e w h i t e  appearance,  for specific  border  much  covering the olecranon region  o f t h e crown obscures  present  the  brown  o f brown h a i r  tarsal  region of the  haired  are present  face-band  Except  tles  regions of the  p r i m o r d i a and s u p r a o r b i t a l  The h a i r  brown  distal  o f dark  band,  on t h e d o r s a l  In general, dark-coloured  on areas  a golden  chin  (except f o r a  bands  the central  The animal  has  9.  claws, glands.  are  8.  of the auricles,  dew  f o r a grey  peripheral  metatarsal  body  7.  portion,  except  of hair  little  of golden  edges  borders  The a n t e r i o r border  very  brown  bris-  of the hoofs. the area  border  i s brown.  hair i s  of the  i s gold  Except  and  f o rthe  A  areas  mentioned,  distal is  10.  11.  STAGE  IX.  t o emerge  i s net p r e s e n t  No w h i t e  hairs  spots  the  on  Lateral of  mm.,  12 60  grams.  the front  the hind  regions  Hair  feet  where  from  the  feet.  of the rostrum  over  generally the  328.5 mm.,  calcaneus-hoof  skin  skin  dark  are  most  hair  of the body  142.5  may  mm.,  be  seen  surface.  s h o r t and n o t dense enough  or  areas.  white  free  i s closed.  the  mouth  have  The  inferior spots, brown  The p r o x i m a l  breadth, weight,  on t h e  The h a i r to  i s  obscure  Most o f the body  appears  white.  The mouth  dark  bizygomatic  length,  appearance,  i n most  grey  white  3.  on  foot.  trunk.  anterior  In g e n e r a l  is  of the hind  locate  hair.  5 6.9  2.  on  to  are evident emerging  Forehead-rump length,  1.  is difficult  to the calcaneus  beginning  hair  hair  areas  The  r i d g e s on  o f mid-brown  nasal pigmentation  the roof of  pigmentation.  i s mid-brown  and t h e s u p e r i o r n a s a l  with  pigmentation  i n colour. regions  of the hoofs  are dark  brown.  48  The  hair  of  the  appearance.  crown  Black  tion  of  hair  i s beginning  face. and  the  The  the  terior  skin end  mystacial  of  covered  by  the  of  the  is  on  the  auricles  Long white  tail  visible  the  gives  several on  hairs  just  hairs  are  the  ears.  this on  mm.  white  beginning present  on  area the  of  black  front  of  hair  of  is  submental band  of  surface of  dorsal  skin  surrounding  of  of the  obvious.  of  the  a  light  ventral  dark  trunk  brown  Small white are  no  of  the  hair  is  ends  visible.  trunk  and  appear-  surface  rump.  white  the  is  surfaces  Virtually  the  of  emergence  Pigmented  emerge  than the  the  long.  to  an-  eyes  site  are  hair  the  (other  to  the  lacks  covering  Hairs  border  area  ventral  the  meatuses  White h a i r s are  A  the  of  loca-  sides  ventral  at  at  eyelids  skin  vibrissae.  the  the  lacks  white  borders  along  rump  ance.  the  auditory  hair  on  brown the  an  lateral  head  visible  of  and  the  The  golden  present  brown h a i r .  superciliary  external  the  muzzle  breaks  Skin  visible the  bordering  light  hair  are  appear  vibrissae).  sides  head.  to  the  i t a  primordia  around  lateral  black  hairs  antler  skin  gives  spots.  of White  6.  Hair is  is distinct  surrounded  on  by  the  tarsal  a peripheral  glands. light  Each  gland  brown band  of  hair. 7.  Hair the  i s e v i d e n t on hind  gland.  The  border of  S T A G E X.  feet  of  hair  mm.,  1,002 1.  gland  brown h a i r  surrounding  length,  the  as  the  an  central  by  a  gland  140  on  thin  i n n e r brown  mm.,  band  area.  bizygomatic  length,  seen  metatarsal  is lined  and  3 3 0 . 5 mm.,  clacaneus-hoof  In  general  appearance,  short but  appearance The  dense to  coat  the body  spots.  The  of  animal hair,  breadth,  weight,  ferior  nasal  apparent the  on  lines the  incisors  roof and  Most  head  of  golden  the brown  the  canines  membrane.  a velvety  i s dark  The  i s covered  appearance.  brown  s u p e r i o r and  mid-brown.  of  periodontal  giving  with  nasal pigmentation  spots.  are  i s covered  surface.  inferior  mid-brown w i t h w h i t e  of  the  superior nasal pigmentation  white  3.  It is easily  grams.  a  2.  feet.  far distally  metatarsal  dark  Forehead-rump 49.8  as  the  mouth. are  Grey The  distinct  in hair  that  Grey-appearing  areas  with is inare  positions under  the  gives  a  skin  is  50  visible  on  rostrum. the  A black  strip  p o s t e r i o r margin  tion ly  the anterior l a t e r a l  to the frontal  on  each  side  Black  sites  of the antler  l i p .  covered band  patches  i s present  upper  with  of hair  anterior ventral  of hair  r e g i o n where  of hair  primordia.  - golden  extentions  A  except  brown  are clearly  much  greater  for a  the hair  regions.  A  posterior  proximal  on  region  forming  of white h a i r  o f brown  the auricles.  The  that  is clearly  visible.  with  are of  of the  ears  and dark  auricle. skin  ventral surfaces  are sparsely covered  midof the  i s present  pigmented  cles  lateral  of the head.  black  edge o f each  each  submental  The v i b r i s s a e  covered  margins  of  and b l a c k  are  hair  white  two b l a c k  d o r s a l surfaces of the a u r i c l e s  pheral  of  margin  and one brown  than  each  at the  strip  The  with  over  d i s t i n g u i s h a b l e as t h e y  length  lateral  of the head i s  with  anterior extension.  head  band  the l a t e r a l  hair  pigmenta  i t extends  are present  ventral surface  white  from  of the superior nasal  along  The  of the  i s present  t o end i n a b l a c k  eye.  hair  regions  brown  are  brown at the  Perivisible  of the  pigmented  auriskin  5.  The d o r s a l a  dark  spots  brown  The  7.  velvet  hair  brown  covering of hair  extending  t h e humerus  and  and continuous  down  have  with  to the  skin  femur. surface of the t a i l with  the dark  i s  brown  of  trunk.  The m e d i a l the  surfaces of the trunk  colour of the dorsal  dark the  dense  of white  covering 6.  and l a t e r a l  surfaces of the legs  ventral  surface hair.  s u r f a c e o f t h e abdomen  of the t a i l The  down t o t h e  genital  are sparsely organs  and t h e  covered  are covered  feet, ventral  i n white  i n white  hair. 8.  The v e n t r a l are  9.  light  The f e e t giving feet  brown  and v e n t r a l  them  and l a t e r a l  i n s h o r t b u t dense  a dark  brown  appearance.  are a lighter  brown  colour than  proximal  neck  i n colour.  are covered  Long h a i r s  10.  thorax  giving  borders  a gold  appearance  hair  The  front  the hind cover  the  of the hoofs.  The m e t a t a r s a l glands a r e s u r r o u n d e d  by  long  of the  gold-coloured hairs.  glandular  area  feet.  is  visible.  The  skin  a border  of  central  52  11.  The  tarsal  hairs not  STAGE X I .  but  the  obscured  Forehead-rump 5 6.9  mm.,  1,345 1.  In  depression  these  long of  white  the  gland  is  hairs.  334  mm.,  bizygomatic  length,  Grey  The  The band  appearance,  covered  pigmented and of  with  skin  lateral  the  159.0  most  breadth,  mm.,  regions  weight,  is visible  and  ventral  of  i s golden  hair and  tinct.  Black  body  posterior to  the  to  the  posterior  branching areas  i s black  i s dark  over  the  of hair  the  black  surface of  eyes  are  the  brown.  i n c o l o u r and dorsal  and  the  i s very  present  dis-  over  the  primordia.  The  dorsal  are  covered  appearance, areas.  brown  covering the  rostrum  antler  the  nares.  nasal pigmentation  crown  of  hair.  superior nasal pigmentation  inferior  5.  by  length,  general  ends  4.  central  by  grams.  nares  3.  are bordered  calcaneus-hoof  are w e l l 2.  glands  surfaces of  the  with  hair  with  some g o l d e n  White h a i r  that  auricles gives  i s present  a  brown on  of dark  the  brown  i n the  the  ears  skin  central of  the  53  posterior  proximal  peripheral at  the  the  band  distal  distal  borders  of  brown  edges.  surfaces  covered with white on  the  with 6.  The of  the  skin  the  trunk  hair the  of  tail  proximal appears 7.  The from  the  vering the  The hair  is  that  is  is  visible skin  of  sparsely present  continuous  surfaces. the to  dorsal  golden  surface  brown  of  the  dorsal  whereas  the  hair  dorsal  covered  spots  ventral  is  A  surface  of  colour.  surface  on  the  the  tail  brown.  femur  appear  that  the  end  is  pigmented  with  white  spots  dorso-lateral surfaces  the  trunk  spots 8.  dark  of  skin  dark  distal  skin  White h a i r  covering  i t a  auricles.  auricles  dorsal  i s black,  end  trunk  hairs  the  the  of  the  give  brown  hair.  fold  of  non-white  The of  anterior  the  pigmented  The of  of  and are  humerus. long  to  the  The  enough  that skin  white  that  extend co-  hairs  some o f  of  the  confluent. surface  is quite  terior  regions.  scured  by  hair  of long  The and  the  abdomen  but  not  dense  g e n i t a l organs  the  teats  are  is  covered in  are  the not  distinct.  with posob-  54  9.  The m e d i a l of  the feet  long 10.  golden hair  border  appearance.  obscuring  The of  white  them  to the  hair  level  that i s  a mid-brown  Gold-coloured surfaces  lines  each  the central  i s pigmented  dorsal surfaces  lateral  gives  the proximal  the hoofs  the  with  down  dense.  of the feet  brown  almost  11.  are covered  gold band of h a i r  of  of the legs  and r e l a t i v e l y  The h a i r  A  surfaces  rows  of the  dark  The  brown  over  a n d much o f t h e way  of  hoofs.  metatarsal  region.  to  gland, keratin most  of  down t h e  surfaces.  tarsal hair,  gland  i s bordered  but the central  area  by  a  thick  i s not  white  mass  obscured  completely.  STAGE X I I .  Forehead-rump length, 63.5  mm.,  2,834 1.  calcaneus^hoof  mm.,  bizygomatic  length,  232.5  breadth,  mm.,  weight,  grams.  In general dense  2.  448  The  appearance,  covering of long  skin  on  the animal  and t h e b o r d e r s  covered  with  black  a  thick,  hair.  the l a t e r a l  rostrum  has  hair.  anterior  regions  of the  of the e x t e r n a l nares  i s  55  The  superior  inferior  nasal pigmentation  nasal pigmentation  is black.  The  i s black with  white  spots. The  incisors  Patches  of  and  dark  canines grey  the  roof  of  the  The  hair  of  the head  light  brown  dorsal  and  The  ventral  for  the brown  hair The of  hair the  Most the  border  ears  distal  are  the  mented  and  the  eyes  glands  on  and  surfaces of  hair the  is visible  of  white  lips.  surface in  colour.  auricles  of  brown h a i r . skin at  hair  auricles,  also.  No  is  the  anterior  long white  there  except  upper  pigmented  the  areas.  light  is visible  surfaces of  antler  Bands  the  the  proximal  Much  bands,  ventral  golden  peripheral  on  of  and  black  i s white  i s very  covered with  and  f o r the  band.  sides  brown  rostral  head  chin  lateral  borders.  skin  the  black  dorsal  except  anterior  surface of  ventral  present  golden  supraorbital  lateral  Some w h i t e  auricles  posterior on  band,  region of  visible. the  i s mostly  bordering the  the  are  mouth.  preorbital  of  erupted.  pigmentation  i n appearance  rostral  primordia  have  t i p of  and i s present but  pig-  56  8.  The  trunk  brown not  9.  hair  with  with  white  golden  spots.  length  white  that the spots  brown  and  light  The w h i t e s p o t s  have w e l l - d e f i n e d borders.  such of  i s covered  The pelage  appear  more  do  i s of  as  tufts  hair.  The h a i r  of the t a i l  has  a very  thick  woolly  appearance. 10.  The h a i r and  the metatarsal  by 11.  long hair  The  tarsal  general 12.  gland  distal  of and  due  brown  by hair  a thick  dark  obscured  colour. and t h e  woolly  completely  i s pigmented  surfaces  are covered  the t a i l rump  genital  has  i n appearance  are completely  i s obscured  area  brown  appearance.  enclosed  brown  i n kera-  except  f o r the  borders.  The m e d i a l feet  glands  are almost  which  i s light  of a l i g h t  gland  The hoofs tin,  13.  of the legs  of the legs  i n white  i s covered  are covered organs  hair.  i n white  a s f a r down The v e n t r a l hair.  i n dense white  The  hair  of t h e male a r e d i f f i c u l t  to the thickness  of the hair  as t h e side  abdomen  and t h e to  covering.  locate  57  3.  The dimensions during  age  tend (Reeve  studied,  to  and b  X,  sidered.  k  without  i s t h e growth  I t i s the value  Y t o X when k = 1 ) .  nificant. tionship  f i t  t o the transformed  The  p r o g r a m was w r i t t e n In  tionships,  made  addition  a standard equation  length  a basis  t o which  of  here  by  Y and X a r e t h e dimensions Y grows  i s unity  i n relation being  to represent  described. least  = log^gb  sig-  the rela-  Ih each'case,  squares  con-  (or the ratio  i s not biologically  i s used  log^Y  than  relative  t o the dimensions  of b  rather  an  straight  line  + k  x  log^n  •  i n FORTRAN I V .  to studying  specific  considered  of comparison  regional  have been  and calcaneus-hoof  a l l data  size  proportions  i s represented  at which  o f Y when X*  o f growth  a l l dimensions  forehead-rump  basis  specific  As  body-  t o time.  parts  ratio  The value  i n each pattern  7040 c o m p u t e r  patterns  reference  The a l l o m e t r i c equation  IBM  provides  1945),  Y = b X*, w h e r e  i s a constant  of various  of absolute  r e l a t i o n s h i p of growing  being  of  and Huxley,  a l l o m e t r i c equation  growth  by the following data.  t o be a function  may b e d e s c r i b e d The  the  relationships during  are represented  growth  absolute growth  size  P A T T E R N S OF GROWTH  to total  may b e c o m p a r e d .  rela-  related to both  length. body  growth  The  length;  I t also  former a  common  facilitates  58  comparison  of  caneus-hoof precise  these  length,  alternative  Table growth  hoof  I t has  to  a  a more  been  Cal-  accurate  and  with  presented  as  an  growth.  comparison  relative be  investigators.  correlate better  therefore  expressing  I I I may  other  i s probably  appears  structures  of  to  seen  of  the  equations  forehead-rump growth  for  length.  relative  to  calcaneus-  length.  described  should in  this  relative  than  obtaining  x  by  a  emphasized  +  symbols  of  a  (which  relative  (plus)  from  appear  The  "growth  synthesized  growth.  sign.  represent  a l l the  s e r i e s of  continuously  1-44 of  that  s e c t i o n have been  data  representations sented  be  dimensions  Figures  Two  of  in Table  It  the  those  II represents  of  Similarly,  and  rates.  method  to  however,  measurement  differentiation  the  data  same  i n the Each  centroid  actual points  after  distinct the  on  patterns" determining  fetuses,  individuals.  Appendix)  observation  are is  i s represented the  rather  line  graphic repre-  by  fitted  an  to  x.  the  data. In graphs  are  examining i n most  growth p a t t e r n s for of  future slope,  of  the  data  cases the  fetus  to  data  below  the  5%  level  fits  below  the  1%  level.  general  trends,  self-explanatory.  investigators. from highest  for  of  and  The  thus  tables  lowest  k  They  provide are  values.  basic  A l l the many  tables  illustrate  and the  information  constructed  s i g n i f i c a n c e and The  the  in  lines  are  f o l l o w i n g comments w i l l  order f i t the  significant aid  in  59  interpreting the In  t h e more  critical  relationships  p a t t e r n s t o be d i s c u s s e d have been the case  cannot  of multiple  comparisons,  b e compared w i t h  i n the data. A l l  evaluated using  the significance  validity, but are useful  t-tests. levels  as g e n e r a l  indicators.  Growth  of t h e head  In length, than the  relation  t o forehead-rump  the cranium  appears  the interorbital region cranium  This  does  n o t grow  i s i n marked  early  phases  other  skull  Growth  post-natal gains  rapidly  I n l e n g t h however,  significantly faster  the rostrum  more  more  than  the rostrum.  growth  during the  rapidly  than  any  area.  of the trunk  t o forehead-rump  grow r e l a t i v e l y f a s t e r (.OKP^.05). comparison, cumference  that  i n diameter  (.0KP<.05).  d i s t i n c t i o n from  of which  Compared  the  t o grow  l e n g t h and calcaneus-hoof  abdomen  When  than  appears  t h e neck  (P<.01).  the pelvic  i n circumference  calcaneus-hoof  the chest  l e n g t h , t h e neck than  In any comparison,  r e g i o n grows  as a b a s i s o f  relatively faster  (.01<P<.05) w h i c h  to  t h e abdomen  l e n g t h i s used  t o grow  appears  grows  there  faster  i s no  significantly faster  i n cirthan  evidence  i n width  than  60  does  the  bone  to  that  the  width  thorax. the  d i s t a n c e between  pelvis  of  the  Each relatively limbs  faster  length  of the  the  crests  of  relatively  faster  i n length  innominate  ilia,  i t  than  appears  in  limbs  of  the  faster  femur  ulna  growth  k  in  examining  values  growth differ  faster  neus -hoof  the  relatively  leg  Table  grows  and  limbs  hind  appear  components  relatively foot  the  I I and  Table  essentially  of  faster  grow  to  grow  the  fore  than  the  relatively  i n the  faster  length  as  two  than  calcaneus-hoof  other  and  The  seen  (P<.01).  than  the  The  bizygomatic  ears  grow  structure.  comparison,  length  i n c r e a s e s more r a p i d l y (P<.01).  diameter  of  be  same s e q u e n c e .  tables.  any  a basis  I I I , i t can  the  transverse chest  in position  than  hind  forefoot.  maintain  of  length  hind  and  the  general  forehead-rump  the  the  of  corresponding  the  length)  using  Thus  the  than  of  breadth  than  tibio-fibula  the  rates  components  the  From  of  the  the  and  Relative  grow  grows  (P<;.01).  humerus  (in  comparing  ( P < . 01) .  Growth  that  When  Indeed,  the  ears  The  calca-  total  length  61  Equations (1950), tailed ment  who  and  the  Rocky  compared tailed  studied  deer,  of  to  deer  have been fetal  Hudson  (1956),  Mountain  i n Table  from  development  the patterns of fetus  derived  mule  who  IV.  i n the studied  deer.  growth  the  of  These the  data  of  Armstrong  Northern the  fetal  equations  Columbian  whitedevelopare  black-  62  TABLE I I PATTERNS  OP GROWTH  Y = b  I N T H E F E T A L P E R I O D OF T H E  WHERE X R E P R E S E N T S FOREHEAD-RUMP  RELATIONSHIP LENGTH  k  Weight  .0001315  2.756  Ear  .002455  1.740  .03917  1.422  length  Calcaneus-hoof  length  Length  of hind l e g  .0588  1.413  Length  of foreleg  .09268  1.313  Length  o f femur  .05047  1.271  Length  o f humerus  .06839  1.213  .07745  1.174  .07129  1.110  .8921  .9 639  .2417  .9 619  .2011  .9519  . 6024  .9254  Hip  bone  Ventral Chest  t o p i n bone tail  circumference  Cranial  length  Preoptic Median Abdomen Hip  length  length  neck  circumference  circumference  width  Transverse  chest  Bizygomatic Interorbital  diameter  breadth breadth  1.432  .8719  .2463  .8699  .3992  .8408  .4608  .8179  .4506  .7763  63 TABLE I I I PATTERNS  OF GROWTH  Y = b X  I N THE F E T A L PERIOD  WHERE X R E P R E S E N T S  k  OF T H E R E L A T I O N S H I P  CALCANEUS-HOOF  LENGTH  Weight  .04764  2.020  Ear  .1292  1.2223  length  Length  of hind l e g  1.479  .9917  Length  of foreleg  1.852  .9224  Length  o f femur  Length  o f humerus  1.080  .8538  bone t o p i n bone  1.119  .8249  Hip  Ventral Chest  tail  length  circumference  Cranial  length  Preoptic Median Abdomen Hip  .9177  length  neck  circumference  circumference  width  Bizygomatic Transverse  breadth chest  Interorbital  diameter  breadth  .7713  .8916  .8129  8.050  . 6757  2.183  .6742  1.749  . 6708  4.925  . 6514  10.424  .6137  1.790  .6111  2.734  .5938  .2726 2.579  .5891 .5504  64  TABLE COMPARISON OF F E T A L Y = b X  Length of hind l e g  Length  h  e  s  GROWTH P A T T E R N S  WHERE X R E P R E S E N T S FOREHEAD-RUMP  DEER  LENGTH  Odocoileus  Odocoileus  Odocoileus  hemionus  hemionus  virginianus  columbianus  hemionus  borealis  1.413 Y =.0588X  1.405 Y =.06637X  1.299 Y  =.09797X  Y  =.09745X  Y  =.7017X  Y  =.3134X  Y =1.592X"  Y  =.8654X*  Y =.2864X  Y  =.1986X  2  Y =.09268X  Y =.8921X  neck  circumference  Abdomen circumference  Bizygomatic breadth  *  Y =.09770X  .9639  t  circumference  Median  I N T H R E E FORMS OF  of  foreleg  C  IV  i  9  2  5  .9716  Y =.8754X  4  Y =.6024X  "  9  5  2  Y =.4132X  Q  J  Y =1.432X*  8  5  5  g  «l 7Q Y =.4608X"  2  .9908  Q  5  9  4  6  y  65  C.  VARIABILITY  DURING  Biological during  development  variation  order  representative  hair  covering  velopment 37  1.  sequence  each  Growth  later  observed  was  selected  a thick  coat  of  i n several  f o r study. very  a This  little  body  hair.  after  of development  between  regions,  regions  during  examining  a r e then  on t h e a n t e r i o r l a t e r a l  emerges  i n this  region  o n many r e g i o n s  h e r e when  observing  this  de-  a series  compared  region.  than  Progressive by  morphological  individuals with  o f changes  The r a t e s  of h a i r  Hair and  with  of  FEATURES  any v a r i a t i o n  i s o u t l i n e d as r e c o n s t r u c t e d  fetuses.  between  of animals  to those  OF E X T E R N A L  and assess  from  and causes  section.  i n different  ranging  i n d i v i d u a l s i s n o t uncommon  The extent  i n this  to detect  group  fetuses  The  of  and growth.  o f development  included  between  D I F F E R E N T I A L DEVELOPMENT  In rates  variation  a r e examined  1.  DEVELOPMENT  levels  skin  i s obscured  of hair  than  of t h e body.  o f development  t h e amount  later  sides  by hair  rostrum  on a d j a c e n t  Skin  may b e  growth  may b e r e a d i l y  present  of the  areas  readily  elsewhere.  distinguished  i n different  animals.  66  2.  Growth the  of  grows  a a  only  the  i s the  suitable colour  surfaces but  on  dorsal surfaces  of  the  auricles  of  ears  Not region  hair  of  indicator  change  the  p r o g r e s s i v e appearance of  occurs  auricles  development,  as  new  bands  of  but  as  emerge.  a p p e a r b l a c k when h a i r  become p r o g r e s s i v e l y more brown  or  hair  red  in  this  the  hair  The  dorsal  first  at higher  emerges, levels  of  development.  3.  Growth the  of  hairs  pelage  isolated  come m o r e dorsal  on  dorsal-most  The spaced  hair  dense  trunk  rows  hairs. and  from  mid-dorsal of  in this  first  emerge  the  As  region  a  the  white  trunk  spots  region  first  appears  development  e v e n t u a l l y obscure appears  of  black but  velvet-like  proceeds, the  as  trunk  between  a  well-  few  the  hairs  be-  underlying skin.  changes  covering to  to brown a  thick  as  The  the  woolly  covering.  4.  Growth  of  The appear hair  later  lies  hair  white than  flat  from  hairs the  along  the white  trunk  that  from  grow  dark-coloured the  back  of  the  spots  the  hairs  trunk of  animal,  the  spots  first  back.  When  relative  growth  the of  67  the of  trunk  proximity  spots at  with  higher  long  5.  spot of  of  white  Growth  of  hair  the  the  length  Growth the  of  in  which  genital  7.  and  but  thick  observing  Animals  may  be  the  degree  having well-defined  d i s t i n g u i s h e d from trunk  spots  those  composed  of  the  the  back  regions growth  of  the  hair  tail  the  is first  is black.  of  the  Part  fetus,  and  the  tail  but  back  emerging,  of  the  when  tail  the  reaches  is  hair  a  appearance.  on  medial  hair  and  tail  a brown  the  of  d o r s a l s u r f a c e of  the  surfaces  in this  long white  posterior  region  hairs  growth  of  hair  i n the  region  of  are  that  ventral  the  surface  of  legs  advances present  from  to  obscures  even  stages  animals the  male  organs.  Growth  In visible.  a  on  white  growth  sparse  exhibiting  the  these  i t has  abdomen  The  hairs  region  the  spots.  by  hair.  hair  during  estimated  development with  on  of both  proximal  certain  6.  of  tufts  colour  be  adjacent  levels  always b l a c k of  may  short white  When the  hairs  of  hair  early As  stages  growth  the  skin  proceeds,  the  of  the  tarsal  gland  of  the  tarsal  gland  gland  is  s u r f a c e becomes  easily covered  68  in short h a i r s .  Longer white h a i r s border the gland but do not  obscure the c e n t r a l r e g i o n .  Upon f u r t h e r development, the  p e r i p h e r a l white h a i r s h i d e the gland and e v e n t u a l l y the r e g i o n takes on a t h i c k w o o l l y  8.  appearance.  Growth of h a i r on the h i n d f e e t  H a i r growth on the h i n d f e e t i s slower than t h a t on the f r o n t f e e t .  The h i n d f e e t appear grey b e f o r e much h a i r  has emerged, take on a b l a c k appearance hair, of  when covered i n s h o r t  and become brown as the h a i r grows.  The c e n t r a l r e g i o n  the m e t a t a r s a l gland, although not i t s e l f  covered w i t h h a i r ,  becomes obscured by p e r i p h e r a l h a i r i n o l d e r animals. animal was  One  found to have h a i r growing from the c e n t r a l g l a n d u l a r  region.  9.  Development of the n a s a l pigmentation  The i n f e r i o r n a s a l pigmentation develops more s l o w l y than the s u p e r i o r n a s a l pigmentation.  It i s therefore lighter  i n c o l o u r than the s u p e r i o r n a s a l pigmentation d u r i n g much of the g e s t a t i o n , but i n the f i n a l  stages of development the  areas of pigment are the same b l a c k c o l o u r . v e n i e n t l y be s e q u e n t i a l l y grouped  as f o l l o w s :  Fetuses may  two con-  (a) those i n  which the i n f e r i o r n a s a l pigmentation i s mid-brown i n c o l o u r  69  and  readily  ment; dark the of  (b)  distinguishable those  brown  or  superior pigment  Most  grey  are  have white  but  spots i n the  less  p r e v a l e n t i n the  from  the  could and  superior  be  10.  of  those  Development  Incisors periodontal to  proceed  development  are  Using fetuses with trait  were  r e g i o n of  on  later  two  at  i n 7.1%  is from  areas  pigmentation. during  a l l levels examined,  White  pig-  of  spots  spots were  of  the  of  those  dewere  absent  animals  that  i n group  (b),  (c).  teeth  may  long before lateral  be  distinguished  they  sides.  erupt.  under  Eruption  Different  levels  of  the appears tooth  distinguished.  preceding data i n order,  o n l y one  orders).  the  n a s a l pigment  ( a ) , 22.2%  group  nasal  distinguish  nasal  animals  stages.  canines  arranged  their  37  of  to  i n which  series  readily  the  respect to rank  the  those  observed  the  membrane from  one  spots  i n group  and  difficult  (c)  superior  nasal pigmentation  have been  as  of  darker  inferior  n a s a l pigment  classified  28.5%  the  i n c o l o u r and  c o n f l u e n t as  White  velopment  the  n a s a l pigment;  animals  growth.  i n which  from  Thus  as  a basis  least  series  comparison,  developed  morphological the  of  t o most  region at  was  arranged  a  time in  the  developed, (single-  developmental  70  order  10 t i m e s .  fetus  was  can  be  assigned  compared  another  level  tics a  t o the rank  results  may b e s e e n  middle  could  on t h e b a s i s  rank value  t o each  V.  i n one  a  given  region  f o r development i n  fetus  i n accordance  of the preceding  When  of t h e ranks  i s evident  develop  further  illustrated  of  correlation.  rank  between  A  i t was  they  would  they  with  characteris-  impossible  i n a group,  were  be assigned  to assign a l l given i f they  to relate  i n a l l animals.  obtained  rates  using  presents  of  to assess  by the fact  of development  that  i n some  .57 i s f o u n d sides of of the  of the inferior  age.  problem  i s greater  features  nasal when  The use o f any  age i s o b v i o u s l y  there  measure  f o r t h e growth of  an i n t e r e s t i n g  structure to fetal  This i s  of the auricles  .21 i s o b t a i n e d  not a l l  Spearman's  a correlation  and t h e development  of characters  the table that  on t h e a n t e r i o r l a t e r a l  of only  situation  complicated  between  rates  and on t h e d o r s a l s u r f a c e s  This  combination  scanning  F o r example,  of hair  on t h e rostrum  attempting  from  by values  correlation  pigment.  just  a t comparable  t h e growth  rostrum  ears.  and  the animals  o f each  i n Table  regions  hair  assigned  that  be d i s t i n g u i s h e d . It  the  i n the series  of development  i t was  of ranking  of development  different  the  (rank)  region. The  the  The p o s i t i o n  than  limited,  variation i n others.  71  I t may be noted, however, t h a t a given f e t u s has been a s s i g n e d  often  t o a s i m i l a r rank f o r a number of c h a r a c t e r i s t i c s .  Animal number 15, f o r example, no lower than 9.5  i s ranked no h i g h e r than 19 and  f o r any area of development.  Thus l i m i t s  may  be s e t f o r the r e c o g n i t i o n of d i s t i n c t stages i n development, as was  outlined i n a preceding section.  sequence  The t r u e developmental  (age order) of the animals i n t h i s s e r i e s may  be approached w i t h i n l i m i t s . of development  likewise  A workable e s t i m a t e of the l e v e l  of each f e t u s t h a t may  approximate t r u e age i n  terms of s t r u c t u r a l development has been o b t a i n e d by  calculating  the mean rank of each animal. In r e f e r e n c e t o these r e s u l t s ,  i t i s i n t e r e s t i n g to note  t h a t s i z e alone cannot be r e l i e d upon as a p r e c i s e of age, as i s d i s c u s s e d i n another s e c t i o n .  indicator  72  TABLE COMPARISON OF  DEVELOPMENT  Ranks level Fetus No.  V OF  10 D I F F E R E N T  REGIONS  assigned  of  t o each fetus f o r d e v e l o p m e n t i n 10 r e g i o n s Mean  ab  *1  2  3  4  5  6  7  8  2 2  1.5  2  1.5  3  1 4  1.85 2 .05  9  10  Rank  1 2  3  2  2  3  1  1  2. 5 2. 5  1 2  1.5 1.5  3  3  4  4  2. 5  4  4  3  4  1  2  3.15  4  3  3  3  2. 5  3  3  4  3  4  3  3.15  5  5  5  5  5  6  5  5  5  6  5.0  6  3 6  6  6  6  6  5  5  5.8  7  7  25.5  7  9.5  12.5  6 7  6  7  6 7  19  8  11  9  10  8  9.5  12.5  17  9  10  15.5 15.5  9 10  14  10 10  11  9.5  12.5  17  17  7  15.5  8  11 8  9  9.5  12 .5  11 12  12  12  10  14  10 10  9.5 25  13  15  10  14  25.5  10  15  9 12  13 10  14 13 12  15  25.5  16  16  20  25.5  17 18  19 30  17  25.5  18  19 20  17  11.7 11.95  11.15  9  8  12 .5 17 1 2 . 5 13 12.5 17  11 10  11 8  15.5 25.5  13  19  15.5  13.5 1 3 . 65  25  12.5  17  10  9  15.5  14.35  21  25  12.5  8  16  18  15.5  17.05  21  25  12.5  17  15  12  15.5  17.95  21 21  25  12.5  10  17  11  25  12.5  18  19 15.5 30. 5 15.5  19  25.5  21  25  23  12 17  23  19  15.5  19.35 20.50  16 21  25  23  24  23  13  15.5  21.06  25 .5  21 21  25  21  30 18  25  23  23  30. 5 15.5  21.35  22  20  25  25  21  25  23  11 24  23  19  15.5  22.10  23 24 25  30  25  21  25  23  24  23  14  15.5  22.40  21 22  23 24 22  25 25.5  21 21  25 25  26  30  31  25  21  25  23 23 23  24 24 24  23 23 23  19 25.5 30. 5 15.5 19 28.5  23.10 23.15 25.05  27 28  30 30  31. 5 25.5 31. 5 25.5  23 32.5  24  23 32.5  30. 5 25.5 1 9 . 0 33.5  29 .45  * Numbers assigned  i n this  9.5  21 25 3 2 . 5 25  horizontal  to regions  row  described  32.5  correspond i n text.  30. 5 15.5  12.0 12 .45  18.15  25.90  t o t h e numbers  73  TABLE  V  (continued)  a s s i g n e d t o eacli f e t u s f o r o f d e v e l o p m e n t i n 10 r e g i o n s  Ranks level  Fetus 2  3  4  5  6  7  8  9  10  Mean Rank  No.  *1  29  30  31.5  25.5  32.5  25  32.5  32 .5 32 .5 3 0 . 5 27  30 31  30 30  31.5 31.5  25.5 25.5  32 .5 25 3 2 . 5 25  32.5 32.5  32.5 32.5  32.5 32.5  30. 5 28.5 30. 5 33.5  30.05 30.60  32  30  31.5  25.5  32.5  25  32.5  32.5  32.5  30. 5 33.5  30.60  33  30  31.5  25.5  32.5  25  30  31.5  25.5  32.5  25  32 .5 32 .5 32 .5 3 0 . 5 3 3 . 5 32.5 32.5 32.5 30. 5 33.5  3 0 . 60  34 35  30  31.5  32.5  30. 5  33.5  30.60  31.5  3 2 . 5 25 32 .5 25  32.5  30  25.5 25.5  32.5  36  32.5  32.5  32.5  30.60  37  30  31.5  25.5  32.5  32.5  32.5  32.5  30. 5 33.5 30. 5 33.5  25  29 .95  3 0 . 60  30.60  74  2.  A compare A  series  only  advanced  than  sidered of  the other.  member  the characters  stage  as  used  from  Fetuses  were  o f development  within  evaluated 19 s e t s  more  pair.  o f t w i n s , one  differed  b u t o t h e r s were  each  to  as a b a s i s f o r  advanced  animals  i n detail,  (have  a higher  with  respect  considered  more  level  examined  of development  was  con-  on t h e b a s i s  f o r assessment. i tnecessary to assign  that  an animal  to a  of i t s siblings.  SEX DIFFERENCES  examined  to detect  between males  compared w i t h r e g a r d  (mean r a n k ) , if  were  examined  of the set of triplets  no c a s e was  different  TRIPLETS  seven.  3.  rates  Some  to be at a d i f f e r e n t  In  were  t o be s t r u c t u r a l l y  i n a s many  AND  development  In a t least  one c h a r a c t e r i s t i c ,  Each  TWINS  o f t w i n s was  of structural  comparison.  appeared  mean r a n k ) to  OP  o f 12 c h a r a c t e r i s t i c s  single  member  o f 28 s e t s  the levels  maximum  any  DEVELOPMENT  sex and s i z e .  members o f o n e s e x w e r e  t o degree  and  possible females.  The  of morphological  The c r i t e r i o n usually  differences i n  larger  a d o p t e d was  specimens development such  that  t h a n members o f t h e  75  opposite  sex  of  equal  possible  sex-specific Twins were  of  a pair  animals  i s the  of  each  than  the  four  sets,  four  sets.  other,  male had greater  same  eight  s e t was  and  on  sets  the the  was  selected  to  an  extent  comparisons  both The in  forehead-rump  and  of  of  a  mixed  was  no  animal  following table.  of  of  15  pairs  more  than than of  sets  relationships  of  twins.  them.  exceeded were  member  female male sex,  case  in in  the  and  a  cases. necessary A  significant  considered  of  lengths.  one  the  mixed  sample  only fetuses that  made b e t w e e n  member  developed  the  length i n every  relatively  size  i n which  i n f o r m a t i o n , i t was  including  was  sex  more d e v e l o p e d  total  each  calcaneus-hoof  developed  l e n g t h i n 12  that  Size  structurally  more  than w i t h i n  c o u l d be  measurements  the  comparisons  obtain further  animals  parisons,  age.  female  other  such  developmental  m a l e was  comparisons  suggest  that  twins  the  greater calcaneus-hoof  To  would  assumption  of  forehead-rump  this  rates.  c o n s i d e r e d t o be  However,  a  growth  e v a l u a t e d on  were based Of  o r h i g h e r mean r a n k ,  To  simplify than  those  the  e v a l u a t e d and  of  make 40  developed  morphological  "larger" of  were  to  size  another other  are  comunless  animal.  represented  76  1.  In comparing in  which  than  a female  versely,  2.  One  was  no  male 3.  and  of a female  male  more  twins  VI) were  either  i n which  found  larger  singleton  developed  singleton  VI).  larger  male.  a female  morphologically  "h" i n T a b l e  were  of equal  found  t o be  or greater  male  t w i n was  developed  Thus  twins  there  developed  found  male  appears  i n the specimens  more There  than  more  Consinglemale.  ("e" i n developed was  a more  no developed  t o be  more  more  developed  female  singles.  more  developed  female  twins.  female  that  singles.  than  morphological  "b," " c " a n d  was  female advancement.  "d" a r e a l l l a r g e r  female  singleton "f")  larger  a tendency  available.  female  larger  than  a structurally  singleton.  than  developed  than  a male  s t r u c t u r a l l y advanced  female  more  nance  ("g"  found  than  was  t o be m o r p h o l o g i c a l l y  i n which  larger  the table,  than  "e" i n T a b l e  a s t r u c t u r a l l y more  discovered  twins  singletons  No  than  one c a s e  twin.  Several  (In  s i t u a t i o n was  larger  instance  ("a" a n d  ("f") c o n s i d e r e d  V I ) was  female  singletons,  ( o f h i g h e r mean r a n k )  c a s e was  Table  and female  two m a l e s  developed  ton  male  twins.  Male  twins  Male  singles  Male Male  f o r male  singles  twins  tend  size  t o be  domilarger  are larger  are larger  are larger  than  than  than more  77  It than the  i s apparent  females  a t each  developmental  therefore that  stage  when  BETWEEN  DEVELOPMENT  larger  (least to  VI SIZE,  rank  developed  most  S E X AND  I N A GROUP OF  Developmental Fetus  t o be  circumstances.  RELATIONSHIP OF  tend  due c o n s i d e r a t i o n i s g i v e n t o  TABLE THE  males  LEVEL  FETUSES  Forehead-rump length  (mm.)  Calcaneus-hoof length  developed)  a.  Male  singleton  328.5  142.5  b.  Male  twin  321.0  147.5  c.  Male  twin  346.0  154.5  d.  Male  twin  335 .0  151.0  singleton  355.0  154.0  e. M a l e f.  Female singleton  6  309 .5  140.0  g.  Female  twin  7  334.0  150.5  h.  Female  twin  8  330.0  150.5  i.  Male  386.0  192.0  j.  Female singleton  10  366.0  176.0  k.  Male  11  448.0  232.5  1.  Female 11  436.0  216.0  twin  twin  singleton  (mm.)  78  4.  THE RELATIONSHIP BETWEEN SIZE AND  FORM  A number of animals were s t u d i e d i n order to estimate the degree of c o r r e l a t i o n between m o r p h o l o g i c a l changes and body dimensions.  For the purposes  of t h i s study, animals were segre-  gated w i t h r e g a r d t o sex, number i n the uterus (twin or s i n g l e ) , and year of c a p t u r e . u t e r i n e environment,  Any v a r i a t i o n caused by sex d i f f e r e n c e s , or environment  e l i m i n a t e d as w e l l as p o s s i b l e .  of the mother, was t h e r e f o r e  Fetuses i n each group were  arranged a c c o r d i n g to t h e i r estimated l e v e l of m o r p h o l o g i c a l development (mean r a n k ) .  They were a l s o ranked i n s i z e order,  i n terms of forehead-rump l e n g t h , b i z y g o m a t i c breadth, hoof  l e n g t h , ear l e n g t h and v e n t r a l t a i l l e n g t h .  calcaneus-  A l l are  measurements t h a t can be taken w i t h some degree of p r e c i s i o n , and thus are p o s s i b l y u s e f u l as i n d i c e s of development. M o r p h o l o g i c a l development was  compared t o each of these  a p p r a i s a l s of f e t u s s i z e u s i n g Spearman's measure of rank c o r relation.  In no group was  a perfect c o r r e l a t i o n obtained.  The  d i f f e r e n t values o b t a i n e d f o r the d i f f e r e n t measurements probably reflect,  i n p a r t , the accuracy w i t h which they may be taken.  h i g h e s t c o r r e l a t i o n of a l l was  The  obtained between calcaneus-hoof  l e n g t h and estimated l e v e l of s t r u c t u r a l development.  Indeed,  t h i s i s a measurement t h a t can be taken w i t h g r e a t e r p r e c i s i o n than any,  and p r o b a b l y r e s i s t s postmortem changes as w e l l as  any.  79 TABLE V I I DEGREE  OF  C O R R E L A T I O N BETWEEN  DEVELOPMENT  AND  Developmental Group  (least to  Male  twins  VARIOUS  DIMENSIONS  rank  developed  most  MORPHOLOGICAL  Size (smallest  rank  to largest)  developed) FR  CH  BZB  EL  VT  1  1  1  4.5  6.5  8  2  7  3  3  3  2  4  2  4 4.5  6.5  4  19 64 5 6  8 Correlation  with  developmental  rank  1  8  8  8  7  6  .55  .95  .19  .50  .31  5  4  1  1 3 Female  twins  1  3.5  4  2  3  2 1  4  2.5  3  2  1  2.5  5.5  4  3  5  5  19 64 5  3.5  6  8  7  8  7  5.5  6  7  8  6  .95  .83  8 Correlation NOTE:  with  5  developmental  8 rank  8 67  .82  .88  F R = F o r e h e a d - r u m p l e n g t h ; C H = C a l c a n e u s - h o o f l e n g t h ; BZB= B i z y g o m a t i c b r e a d t h ; EL=Ear l e n g t h ; VT=Ventral t a i l length.  80  TABLE  VII  Developmental (least  Group  to  Male  singletons  (continued)  rank  developed  most  Size (smallest  rank to  largest)  developed) FR  CH  BZB  EL  VT  1  2  2  4  4  2.5  2  1  1  1  1  1  3  3  3  2  2  4  4  4  4  5  5  5  5  5  5  3  3  2.5  .90  .90  .35  .55  .35  1966  Correlation  with developmental  rank  81  DISCUSSION  This ment is  investigation outlines  of t h e Columbian  small  T E C H N I C A L PROBLEMS  Preservation  Many rely  on d a t a  however, may  that  cause  of  obtained  changes  size  1925; J o u b e r t , most  not significant,  and  difficult i f fine  The sample  size  suggestive.  DEVELOPMENT  of f e t a l  material  I t i s well  i n various  a period great  of  known,  fluids  time.  v a r i a t i o n may  dimensions,  of animals.  investigators  populations  different  Similar variation  (e.g.,  Scammon  and  1956).  o f t h e changes they  to predict.  r e s u l t i n g from  are variable, For these  results are required,  preservation.  after  of different groups  i n wild  material.  the results that  by other  are  before  from  and d i f f e r e n t observed  growth  preserved  the reactions  Although  that  IN STUDYING  i n the dimensions  between  Calkins,  from  preservation  occur  been  fetus.  of the develop-  be considered  investigators studying  i t i s evident  has  aspects  material  Also,  animals  b l a c k - t a i l e d deer  and t h e r e s u l t s can o n l y  1.  some  difficult  reasons  fetuses  to  preservation explain  i t i s suggested  should  be  measured  82  Selection  of  The Todd, that  results  1925)  show  attempt  to  points  has  the  useful more is  for  formalin  this the  for  The  of  greater  soft  work  for  t i s s u e does  preservation  not as  for  studying animals ages.  the  development over  a  period  Environmental  trolled,  and  a  subcutaneous  and  most  i n v o l v i n g bony  These  greater  due  to  their  precision.  well  as  PETAL  defined resist  bone  factors  more and error  points.  caused  by  them.  MATERIAL  of  unknown  accurate  and  describe  time  are  Human  reference  resists  points  accuracy  changes  bony  precise  reference  tissue covering.  easily  (e.g.,  measurements  their  i s to  s e r i e s of  accurate  seem t o  most  of  external  to  growth: m a t e r i a l  Certainly  investigations  due  less  2.  Sampling  most  those  growth  field  other  dimensions between  probably  amount  and  s e l e c t i o n of  represent  studying  useful  soft  that  are  least  probably  Also,  of  limitations.  measurements with  measurements  as that  and  each  age  complete measure  reaches  influence  individual  records  a  method a  of  s e r i e s of  s e r i e s of  known  growth  can  can  obtained  be  be  live  conas  83  well to  as a s e r i e s o f p o p u l a t i o n  apply  t o most w i l d  intra-uterine  wild but also  were  unknown,  specimens  provide  absolute and  t h e e n v i r o n m e n t was  much  t o some  change  regarding  growth  rates  detected  may  a g e may  o f known  i t i s possible  of t h e specimens  season.  Nevertheless,  not be u s e f u l  i n studying  in  studying  relative  growth  become more m e a n i n g f u l i f age.  Using  to time.  Inferences  described. i n a very  may  then  then be  studying  one  of which  c e r t a i n dimen-  unknown-age m a t e r i a l  o f unknown-age m a t e r i a l ,  effective  specimens  t o d e t e r m i n e how  r e l a t i o n s h i p i s a s s u m e d when  i s attempting An  been  was n o t  here.  t h e r e l a t i o n s h i p between  (If this  has  specimens  i n relation  time.  which he  the population  o f unknown-age i n d i v i d u a l s  data  treated  o f unknown  a g e i s known,  sions  both  t o use f o r  specimens  Material related  i n t h e same  These  impossible  n o t c o n t r o l l e d , and a l l t h e  but are invaluable  differentiation,  study  method i s d i f f i c u l t  The ages  of a population  data.  growth,  Known-age  the  intra-uterine.  sampling  This  and n i g h  In t h i s  were n o t c o l l e c t e d  sequential may  populations  development.  only  means.  made  and  absolute  i s assuming  that  to prove.)  technique  f o r obtaining  The o c c u r r e n c e short  time by  known-age  of estrus  assessing  two  fetuses  i n t h e doe can be behavioral  84  situations: gator  series  i n t h e same p e n .  Browman  In contrast and Hudson  required  (1957),  distress.  controlled  t o such  of estrus  concerned  observed  material  3.  This sequence  period.  and water  study  Development,  processes  problems  by  much p h y s i c a l a n d under  carefully  may b e  maintained  consumption. The  i s a great  short advantage  age.  to reconstruct i n form  however,  occurring  of this  study  with  DEVELOPMENT  has attempted o f change  larger  of time i s  accordingly.  i n this  of accurate  a  placed  described  expenditure  are saved  as f o o d  a v a i l a b l e f o r study were  research  as t h a t  may b e k e p t  P A T T E R N S OF  and p a t t e r n s  biological  same s y s t e m ,  such  a minimal  factors  investi-  p o s s i b l y be obtained  i n t e r a c t i o n may b e r e g u l a t e d  obtaining  mens  could  to the  a n d d o e when  c o n d i t i o n s , and i n d i v i d u a l records  regard  duration  fetal  this  The animals  animal  t h e buck  t o a system  and t h e animals  emotional  Social  Using  o f known-age a n i m a l s  success.  in  of the isolated  and t h e i n t e r a c t i o n between  together  with  the reaction  and s i z e  during  i s a composite  i n the l i v i n g not living,  nature.  the probable  o f dynamic  animal.  a common  The p a t t e r n s  of  the  The s p e c i -  situation i n development  85  that  have  been  described  must  therefore  be  treated  as  purely-  inferential.  External  form  A  series  sequential stages of  are purely  material  with  stages,  available,  f o r other  the  period  eyelid  growth,  sequence  of  created  to outline  form of only  available  as  the  the fetus.  These  a convenient  means  f o r study.  When  a more d e t a i l e d o u t l i n e ,  fetal  species  development  more  including  sealing  emergence  i s s i m i l a r to  o f mammals.  are represented,  periderm  of h a i r  described  made.  pattern  described  been  the external  subjective,  can be  The  of  have  the material  becomes  fetal  stages  development  dealing  more  of  The  including and  typical  stages  in  the c h a r a c t e r i s t i c  periderm  demonstrates  that  resorption.  a cephalocaudal  The growth  gradient. It this a  is difficult  species  function  probably  of  to others, time.  inaccurate,  as  most  since  investigators.  between  fetal  Rocky  compare  Comparisons  between  the  to  Mountain  mule  studies based  technique's  However,  development  the f e t a l  express on  and  of  development  absolute  size  of p r e s e r v a t i o n  relationships will  i n the Columbian  deer  development  be  as  are  differ discussed  b l a c k - t a i l e d deer,  the Northern  white-tailed  86  deer.  I n f o r m a t i o n on  Armstrong  (1950)  In curred  at  deer.  For  tail in  at  the  the  and  the  latter  Hudson  a  smaller size example,  a  deer  at  a  at  168  i n the b l a c k t a i l  length  The  blacktail  by  229  mm.,  i n the  whitetail  by  2 35  mm.  Hair  mule  and  pectively. the  whitetail  Incisors  three forms.  difficult  and  No  difficulty  as  45.2  tailed Indeed, time  erupt  uncertain  in this  The  known-age  deer  and  to  mule  size,  served  i n the  rates,  techniques  specimens  at  (1959) i n mule  trunk by  on  the  mm.,  385  mm.  deer  252  mm.,  deer  sex  i n the  They of  oc-  other  327  mm.  mm.  and of  and mm.  evident and  appear  limbs mm.  are  110.7  spots  397  found  t o have  in in  the the  the b l a c k -  270  and  mm.  441  resmm.  determination  fetuses  fetuses  in  rather  smaller than  sexing blacktail  in  as  70  mm.  small  study. specimens deer  i f patterns of  than  226  encountered  mm.  from  evident i n the b l a c k -  length  white mule  appear than  are  103.2  appears  by  Hudson  was  of  forehead-rump mm.  i s taken  changes  metatarsal glands  forehead-rump  mule  forms  (1956).  many r e g i o n s s t r u c t u r a l  whitetail  tail,  two  available  fetuses  development  this  may  p r o b a b l y do  preservation not  account  faster  a r e more  explain  p r e c e d i n g examples. of  grow  suggest  some o f However,  and  that  than  closely the  blacktails. related  variation  different  incomplete  f o r a l l the  white-  to  ob-  growth  series  variation  of observed.  87 Greater features.  For  variation example,  whitetail  by  114  blacktail  by  12 6 mm.  in  the  46  mm.  in  the  deer  The  large body  mule  mm.,  at  the  The 59.2  in  mule  surface  deer  and  mm.).  In  summary  then,  similar  in  the  tend  to  occur  deer  than  in  at  a  the  exists  the  different  differences  by  size  forms. in  well  of  size  mm.  of  In  the some  absolute  the  in  the  in  the  "distinct" whitetail  i s reached hair  cases of  this  a the  appears  structural  Columbian  at  obvious  and  covers  development Most  rates  and  other  d i s t i n c t nor  after  deer.  in  in  of  apparent  110.7  neither  sequence  forms  smaller  other  first  "obvious"  much g r e a t e r  fontanelle  (300  is  development  s a g i t t a l s u t u r e becomes mm.  a  the  pigment  s a g i t t a l s u t u r e becomes  anterior  by  seen  nasal  in  blacktail until  plained  is  changes  black-tailed may  be  ex-  growth.  Growth  Careful of  the  data  on  consideration growth.  different  kinds  nutrition  must be  organs  respond  relationships in  comparing  of  to  When  tissues  in  considered. poor  described  must be studying the  given  fetus,  in  interpretation  allometric  Different  nutrition  to  the  growth  maternal  fetal  between weight  and  intra-specific populations.  length Most  plane  tissues  d i f f e r e n t ways.  of  between of  and  The  growth  may  be  useful  the  growth  88  patterns  described,  skeleton. ment  According  within  in  slowly  of  maternal  patterns fected of  one  or  kind  of  fetal  r a p i d l y growing  growth  the  than  If  of  would  Fetal  the  have been  compared  to  deer  and  Mountain  mule  Growth using chest,  testing has  the  been  abdomen,  relatively  In and  hind the  of  compared  s i x morphological  In b o t h  the  data  forelegs the  faster  legs  abdomen  significant.  and  of  on  the  of  and  same plane  then  the  little  sense w i l l  afbe  popu-  the  black-tailed  Northern,white-tailed examining  (1950) of  and  and  statisti-  Hudson  (1956).  forehead-rump the  length  cranium,  neck,  the  grows  legs.  and  mule  deer  than  faster  hind  the  high  the  than  neck  abdomen  the  legs  and  (P<.01)  forelegs  d i f f e r e n c e s i n growth between the  develop-  patterns.  namely  i n circumference  between  the  of  is  or  deer,  in this  after  hind  bone)  Columbian  basis  regions,  pattern  (low  to  in  d i f f e r e n c e s between  Armstrong  blacktail  the  and  the  deer  growth  l i k e w i s e be  growth  those  grow r e l a t i v e l y  whitetail,  applies  genetic  deer  cally  fetuses  doe  time-related  the  (e.g.,  here w i l l  growth p a t t e r n s  Rocky  tissue  this  in indicating  mainly  (1948),  sheep  described  nutrition  value  lations  reflect  to Wallace  nutrition).  of  by  more  however,  forelegs  the are  and  (P<.01). neck not  89  In not  most  significantly  bizygomatic the  cases  mule  patterns to  more  faster  increases  regions than  slowly  As variation  whitetail  variation  blacktail.  are The  (P<.01)  in  whitetail  appear  to  different  the  regions  length,  are  the  hind  Regions  the  median  VARIATION  be  to  IN  of  the  leg  that neck  of  grow  have  blacktail. the  whitetail  relatively  circumference  bizygomatic  In  breadth  (P<.01), (P<.01).  DEVELOPMENT  of  results,  fetuses  or  i n the  It  partially  skeletal  genetic  genetic  considerable  i n the  differentiation  is regional  field  the  another.  wholly  in regional out,  from  between  compared may  there  general  deer  faster  (.01<.P<.05) a n d  i s evident  (1958) p o i n t that  the  mule  variation  occurs  region  ferences  those  of  the  relatively  (.01<P<.05).  4.  one  of  corresponding  circumference  Individual  of  forehead-rump  i n the  abdomen  from  patterns  deer.  relation grows  growth  different  breadth  Several growth  the  rate  is possible  development that  attributed to rates.  system  control  influence)  of  individual of  this  genetic  dif-  As  Kraus  and  alone,  i t is  possible  (in addition  in prenatal  bone  to  Choi  the  growth.  90  Sex  ratio  The 38  females  sex to  males.  Hudson  (19 59)  by  deer.  Downing i n which  females  by  offered  the  year  may  matings this  Sex  (19 65)  f e t u s e s of  This of  100  females  produce that  that  the  predominantly predominantly few  bucks  87  the  males  was  ratio in  ob-  mule  in  white-tailed  in a ratio  of  24  nearly  first male  more  even  service  one  to  6  ratio.  a buck  while  offspring than  males  sex  of  offspring,  female  bred  to  to  sex  situation  p o p u l a t i o n of  suggestion  determinable  i s i n contrast  fawns were p r o d u c e d  produce  case,  the  described a  a breeding  He  each  subsequent  (assuming,  in  doe).  differences  The grows the  female  deer. same  She age,  the  female rank,  results  faster  criteria,  In  of  46  served  deer  ratio  of  (at least  fetus. reached found but  the  same  that  some  suggest  time  (1950),  These Donald  the  using weight  conclusion regarding  singletons male  that  than  were h e a v i e r  twins  Purser  fetus  were  found  (19 5 6)  and  than  age  as  white-tailed  females than  to be  d e t e r m i n a t i o n s were based and  male  d u r i n g development)  males were h e a v i e r  c u r r e n t study,  however.  at  study  Armstrong  female  singles.  this  observed  male  of  twins.  larger on  the  than  length  male  size  and  91  dominance  i n twin  fetus  be  may  supply 17  dency rump  of  fetal  mule sex  be  23,  number  31  Cheatum  and  mule  quadruplets  deer to  number  suggest  f o r some  environment. twins  and  dominate  t o be  that  element  Hudson  found  no  in either  the in  male  short  (1959)  examined  significant  weight  or  ten-  forehead-  Verme  different (during  twin  and  triplet  fetuses  56.3%  and  1.8%).  Hudson  of  67.7%,  suggested  deer  of  the  (19 65)  and  ratio  to  ovulation).  2:21:1  17:2:0 f o r a n i m a l s  variation  of  on  containing  12%  was  of  The  The  diet.  protein  related  size to  nutrients.  white-tailed  an  and  of  of  geoThe  on  1,250  deer  was to  November  singles,  excellent  Animals with  and  other population  ratio on  and  t h e more p r o d u c t i v e  during October  f o r animals  a poor  in litter  essential  subjected pregnant  prior  pellets  Morton  3:6:1.  nutrition  was  2.1%.  triplets  of  triplets  6.3%  that  planes  or  triplets  and  found  (1956)  twins,  populations studied  twins  was  singles,  in availability  singles, was  (41.8%,  2 3.8%,  variation of  single,  recoveries  (1946) h a v e  population 8:2:1.  of  one  deer  white-tailed  graphic  fed  compete  and  'size  reports  and  to  sex  length.  The  two  mixed  intra-uterine  for either  Litter  to  of  able better  i n the  sets  lambs  both  twins  diet, diets  digestible  were  92  calories  per  gram.  approximately  3.5  on  the low  in  O c t o b e r and  Twin  on  the high  pounds  of  ration  plane of n u t r i t i o n 1.5  pounds  were  daily  nutrition  per  diet  animal per  each  f e d 2.5  received  day.  Animals  pounds  per  day  i n November.  development  The was  Deer  found  degree  t o be  of  different  mixed  sex  studied.  twins  of w h i t e - t a i l e d  development  Variation  t o be  in  observed species  the  external  every pair  (1950)  fetuses  same w i t h i n  during  o f mammals. (Castle size  Hammond,  1958),  nutrition  deer  and between  maternal  Record,  Armstrong  to influence  genotypes  within  of  of  examined  and  characteristics  found  fetal 11  twins  sets  of  the degree  of  mixed of  each s e t .  general  Variation individuals  development  (Foote  the development include  Gregory, et al.,  m a t e r n a l age  sex  1948), of  development  populations.  These  and  (Wallace,  1953),  prenatal  Many f a c t o r s  have  of  in  1959;  G r e g o r y and Hunter,  (Dickinson  the fetus  the fetus  t h e m a t e r n a l and  1929;  parity  i s common  of  t h e mother  ( D o n a l d and  been  various  paternal  Castle,  1956;  et al.,  between  1931),  Joubert  1962),  maternal  (McKeown  Purser,  and  and  1956),  litter  size  (Wishart  et_ a _ l . , 1 9 5 5 ) , The describing useful only  a n d Hammond,  and ambient  current the basic  appraisal  1933),  temperature  placental  (Yeates,  o f development  of t h e causes  b e made i n f u t u r e  studies  of variation using  (Eckstein  1958).  i n v e s t i g a t i o n has mainly been pattern  size  concerned  of the fetus.  with A  i n development can  controlled experiments.  94  LITERATURE  ARMSTRONG,  R.  A.  white-tailed  1950.  Fetal  deer.  Am.  CITED  development  Midland  of the northern  Naturalist  43:650-666.  BALINSKY, B. I . 19 6 0 . An i n t r o d u c t i o n t o embryology. S a u n d e r s Company, P h i l a d e l p h i a . x i v + 5 62pp. BROWMAN,  L . G.  behavior C A S T L E , W. of  AND  o f penned mule  E . AND  size  P. HUDSON.  P. W.  1957.  deer.  GREGORY.  inheritance  Observations  J . Mammal. 1929.  The  W.  B.  on t h e  38:247-253.  embryological basis  i n the rabbit.  J . Morphol.  MORTON.  Techniques  Physiol.  48:81-103. CHEATUM,  E . L . AND  G.  H.  1942.  used  i n  determining  t h e p e r i o d o f t h e r u t among w h i t e - t a i l e d  of  State.  New  York  Trans.  Seventh  N.  Am.  Wildl.  deer  Conf.  337-342. CHEATUM, E . L . AND G. H. MORTON. 1946. Breeding season of w h i t e - t a i l e d d e e r i n New Y o r k . J . W i l d l . Mgmt. 1 0 : 2 4 9 - 2 6 3 . COWAN,  I . McT.  tailed  1956.  deer?  America.  The  What  I n W.  and where  P.  Taylor  Stackpole  a r e t h e mule  ( e d . ) , The  Company,  and b l a c k -  deer  Harrisburg,  of  North  Pennsylvania.  335-359. , AND of ,  VALERIUS  t h e genus AND  A.  Queen's G.,  T A Y L O R AND a  study  DONALD, H.  19 6 1 .  CHARLES GUIGUET.  Columbia. DICKINSON,  GEIST.  Odocoileus.  P.  between  G. W I E N E R .  AND twin  19 6 5 .  Printer,  J . L . HANCOCK,  involving  egg  A. F . lambs.  Aggressive behavior  J . Mammal.  1962.  T h e mammals  Victoria, G.  J . R. The  transfer.  PURSER.  1 9 5 6.  in  deer  4 2 : 5 2 2 - 5 2 6. of  B. C. HOVELL,  size  British  414pp. ST.  o f lambs  Animal.Prod. Competition  J . A g r . S c i . 48:245-249.  C. at  S. birth-  4:64-79. i n utero  95  DOWNING, R O B E R T L . 1965. An u n u s u a l s e x - r a t i o deer. J . W i l d l . Mgmt. 2 9 : 8 8 4 - 8 8 5 . ECKSTEIN,  P., T . McKOEWN AND  placental pig.  weight  J. Endocrinol.  F O O T E , W.  C ,  sequence  GREGORY, J. HAFEZ,  J . Animal AND W. Zool.  E . S. E .  of  12:108-114.  levels.  AND  L . E . CASIDA.  I I . E f f e c t s on f e t a l  E . CASTLE. basis  1959.  develop-  1931.  of size  Further  studies  inheritance  on  i n the rabbit.  59:199-211.  19 6 3 .  prenatal  Variation i n  i n the guinea  S c i . 18:463-474.  embryological Exptl.  1955. size  i n t h e y e a r l i n g ewe a s a f f e c t e d b y b r e e d a n d  of feeding  P. W.  the  to litter  A . L . P O P E , A . B . CHAPMAN  Reproduction ment.  R. G. RECORD.  according  i n white-tailed  Symposium  and p o s t n a t a l  on growth:  growth.  Physio-genetics  J . Animal  S c i . 22:  779-791. HUDSON, P A U L . 1956. The morphology o f t h e embryonic and f e t a l development o f t h e Rocky Mountain mule d e e r . M.S. T h e s i s . Montana S t a t e U n i v e r s i t y , M i s s o u l a . 61 p p . .  19 5 9 .  Fetal recoveries  i n mule  deer.  J. Wildl.  Mgmt.  23 : 2 3 4 - 2 3 5 . , AND in  t h e mule  HUNTER,  G. L .  J.  KRAJINA,  Embryonic  J . W i l d l . Mgmt.  and f e t a l  development  23:295-304.  The maternal i n f l u e n c e  1 9 5 6.  A  i n t h e sheep. V. J .  British  Western  KRAUS,  deer.  1956.  D. M.  ment  of  19 5 9 .  on s i z e  i n sheep.  A g r . S c i . 48:36-60.  JOUBERT,  of  L . BROWMAN.  19 6 5 .  Biogeoclimatic  Columbia. Columbia.  B E R T R A M S. AND  analysis  and develop-  zones  and biogeocoenoses  Department  (ed.),  Ecology of  o f Botany,  University  1:1-17.  SUNG C H I L  of the prenatal  22:231-242.  growth  I n V. J . K r a j i n a  North America.  British  Growth  study of pre-natal  J . A g r . S c i . 47:382-428.  CHOI.  growth  1958.  A  factorial  o f t h e human  skeleton.  96  McKEOWN,  T . AND  size J. MEYER,  R. G. RECORD.  on f e t a l  growth  Endocrinol. ARTHUR W.  Animalium fornia.  1936.  An a n a l y s i s  Stanford  Harvey.  University  Press.  E . L . CHEATUM.  J.  Wildl.  Mgmt.  C. H.  1966.  Lens weights  Wildl.  Mgmt.  30:417-419.  PATTEN,  B . M. AND  preparatory PETROV,  A . K.  REEVE, in  SCHUFELDT,  R. W.  SEGELQUIST,  JULIAN  upon  York.  fetuses.  o f embryos  20:393-413.  vnutriutrobnogo  Zh. 43:125-130. 1945.  growth. Essays  Press.  19 6 6 .  I n W.  1925.  Clark  of  fetal  N a t . S c i . 224-233. deer  embryos  by  30:414-417.  The r e l i a b i l i t y  subcutaneousbony  E. Le Gros  features  Acad.  white-tailed  Mgmt.  problems  1 2 1 - 1 5 6.  Proc.  Sexing  Some  on Growth and Form.  On t h e e x t e r n a l notes.  J. Wildl.  T. WINGATE.  Record  S. H U X L E Y .  (eds.),  1891.  C. A.  differences  i n New  deer  osobennosti  of allometric  and other  chromatin.  deer  The shrinkage  Anat.  Zool.  a t the Clarendon  reindeer  Cali-  (Some c h a r a c t e r i s t i c s o f i n t r a u t e r i n e  of elks.)  P. B. M e d a w a r  Oxford  Regional  o f mule  1921.  Nekotorye  losya.  the study  and  TODD,  R. P H I L P O T T .  E . C . R. AND  Generatione  University,  of white-tailed  to sectioning.  development  of b i r t h .  10:242-248.  19 6 4 .  razvitiya  of placental  xx+617pp.  1946.  breeding  J.  potential  o f t h e De  Stanford  in  NELLIS,  The i n f l u e n c e  t o sex and order  10:73-81.  of William  MORTON, G. H. AND  1953.  according  points.  Am.  o f measurements  J . Physical  based  Anthropol.  8:275-279. THOMAS,  J . W.,  R. M. ROBINSON AND  R. G. MARBURGER.  S o c i a l behavior i n a w h i t e - t a i l e d deer herd hypogonadal males. J . Mammal. 4 6 : 3 1 4 - 3 2 7. VERME,  LOUIS  tailed WALLACE,  J.  19 6 5 .  deer.  L . R.  Reproduction  J. Wildl.  1948.  birth in relation Sci. 38:368-401.  Mgmt.  The growth  studies  19 6 5 . containing  on p e n n e d  white-  29:74-79. o f lambs b e f o r e  to the level  of n u t r i t i o n .  and a f t e r J .Agr.  97  WITSCHI,  EMIL.  Saunders YEATES, of  N.  T.  high  84-89.  1956. Company, M.  Development of v e r t e b r a t e s . Philadelphia.  1958.  atmospheric  Foetal  B.  xv+588pp.  dwarfism  temperature.  W.  i n sheep-  J. Agri.  an  effect  S c i . 51:  98  APPENDIX  FIGURE  1  R E L A T I O N S H I P BETWEEN WEIGHT AND FOREHEAD-RUMP L E N G T H  FIGURE  EAR  2  R E L A T I O N S H I P BETWEEN L E N G T H AND FOREHEAD-RUMP  LENGTH  FIGURE  3  R E L A T I O N S H I P BETWEEN C A L C A N E U S - H O O F L E N G T H AND FOREHEAD-RUMP  LENGTH  FIGURE  4  R E L A T I O N S H I P BETWEEN L E N G T H OF H I N D L E G AND FOREHEAD-RUMP  LENGTH  FIGURE 5 RELATIONSHIP BETWEEN LENGTH OF FORELEG AND FOREHEAD-RUMP LENGTH  FIGURE RELATIONSHIP LENGTH  6 BETWEEN  OF FEMUR AND FOREHEAD-RUMP  LENGTH  FIGURE  LENGTH  7  R E L A T I O N S H I P BETWEEN OF HUMERUS AND FOREHEAD-RUMP  LENGTH  FIGURE RELATIONSHIP  8 BETWEEN  L E N G T H OF T H E INNOMINATE BONE AND FOREHEAD-RUMP  LENGTH  FIGURE 9 RELATIONSHIP BETWEEN VENTRAL TAIL LENGTH AND FOREHEAD-RUMP LENGTH  FIGURE RELATIONSHIP CHEST CIRCUMFERENCE  AND  10 BETWEEN FOREHEAD-RUMP  LENGTH  FIGURE 11 RELATIONSHIP BETWEEN CRANIAL LENGTH AND FOREHEAD-RUMP LENGTH  FIGURE RELATIONSHIP PREOPTIC  L E N G T H AND  12 BETWEEN FOREHEAD-RUMP  LENGTH  FIGURE RELATIONSHIP MEDIAN  NECK  CIRCUMFERENCE  13 BETWEEN  AND  FOREHEAD-RUMP  LENGTH  o o o o  111  10  FIGURE RELATIONSHIP ABDOMEN C I R C U M F E R E N C E  AND  14 BETWEEN FOREHEAD-RUMP  LENGTH  FIGURE RELATIONSHIP  15 BETWEEN  H I P WIDTH AND FOREHEAD-RUMP  LENGTH  10  FIGURE 16 RELATIONSHIP BETWEEN TRANSVERSE CHEST DIAMETER AND FOREHEAD-RUMP LENGTH  FIGURE  BIZYGOMATIC  17  R E L A T I O N S H I P BETWEEN B R E A D T H AND FOREHEAD-RUMP  LENGTH  FIGURE RELATIONSHIP I N T E R O R B T I A L B R E A D T H AND  18 BETWEEN FOREHEAD-RUMP  LENGTH  FIGURE RELATIONSHIP WEIGHT AND  19 BETWEEN  CALCANEUS-HOOF  LENGTH  FIGURE RELATIONSHIP E A R L E N G T H AND  20 BETWEEN  CALCANEUS-HOOF  LENGTH  o o  118  •CD  o  CD CD  Y =  .1292  X  1.2223  CD 'CD  CD  •H  -P QJ  Calcaneus-hoof Length i n  10  100  mm.  FIGURE RELATIONSHIP L E N G T H OF H I N D L E G AND  21 BETWEEN  CALCANEUS-HOOF  LENGTH  FIGURE 22 RELATIONSHIP BETWEEN LENGTH OF FORELEG AND CALCANEUS-HOOF LENGTH  120  CD CD  FIGURE 23 RELATIONSHIP BETWEEN LENGTH OF FEMUR AND CALCANEUS-HOOF  LENGTH  FIGURE RELATIONSHIP L E N G T H OF HUMERUS AND  24 BETWEEN  CALCANEUS-HOOF  LENGTH  FIGURE RELATIONSHIP L E N G T H OF T H E  INNOMINATE  25 BETWEEN  BONE AND  CALCANEUS-HOOF  LENGTH  FIGURE 2 6 RELATIONSHIP BETWEEN VENTRAL T A I L LENGTH AND CALCANEUS-HOOF LENGTH  FIGURE RELATIONSHIP C H E S T C I R C U M F E R E N C E AND  27 BETWEEN CALCANEUS-HOOF  LENGTH  FIGURE RELATIONSHIP CRANIAL  L E N G T H AND  28 BETWEEN  CALCANEUS-HOOF  LENGTH  o o o o  12 6  o o  Y  =  2.183  X  6742  CD  o  CD -rl-' •h  -P  C  0)  rH  (0  •h  C  (0  u  Calcaneus-hoof  10  100  Length  In  mm.  FIGURE 29 RELATIONSHIP BETWEEN PREOPTIC LENGTH AND CALCANEUS-HOOF LENGTH  FIGURE RELATIONSHIP MEDIAN  30 BETWEEN  N E C K C I R C U M F E R E N C E AND  CALCANEUS-HOOF  LENGTH  FIGURE 31 RELATIONSHIP BETWEEN ABDOMEN CIRCUMFERENCE AND CALCANEUS-HOOF LENGTH  FIGURE 32 RELATIONSHIP BETWEEN HIP WIDTH AND CALCANEUS-HOOF LENGTH  FIGURE RELATIONSHIP BIZYGOMATIC  B R E A D T H AND  33 BETWEEN CALCANEUS-HOOF  LENGTH  FIGURE 34 RELATIONSHIP BETWEEN TRANSVERSE CHEST DIAMETER AND CALCANEUS-HOOF LENGTH  FIGURE 35 RELATIONSHIP BETWEEN INTERORBITAL BREADTH AND CALCANEUS-HOOF LENGTH  FIGURE RELATIONSHIP INTERORBITAL  B R E A D T H AND  36 BETWEEN BIZYGOMATIC  BREADTH  1  I  1  0  I  1  0  ' I  0  1  0  0  0  •  i  I n t e r o r b i t a l Breadth i n mm.  * ll  I  1  0  0  0  0  FIGURE RELATIONSHIP EAR  L E N G T H AND  37 BETWEEN  BIZYGOMATIC  BREADTH  FIGURE  38  RELATIONSHIP  BETWEEN  P R E O P T I C L E N G T H AND  CRANIAL  LENGTH  FIGURE 39 RELATIONSHIP BETWEEN CHEST CIRCUMFERENCE AND ABDOMEN CIRCUMFERENCE  137  FIGURE 40 RELATIONSHIP BETWEEN TRANSVERSE CHEST DIAMETER AND HIP WIDTH  FIGURE RELATIONSHIP H I P WIDTH  AND  L E N G T H OF  41 BETWEEN INNOMINATE  BONE  FIGURE RELATIONSHIP LENGTH  OF F O R E L E G AND  42 BETWEEN LENGTH  OF  HUMERUS  FIGURE RELATIONSHIP L E N G T H OF H I N D L E G AND  43 BETWEEN L E N G T H OF  FEMUR  FIGURE RELATIONSHIP LENGTH  OF FEMUR AND  44 BETWEEN LENGTH  OF  HUMERUS  


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