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Identity and limits of Limbella tricostata (Musci: Amblystegiaceae) Christy, John A. 1985

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IDENTITY AND  LIMITS OF LIMBELLA TRICOSTATA AMBLYSTEGIACEAEl  (MUSCI:  by JOHN A.  CHRISTY  B.S., U n i v e r s i t y  Of O r e g o n ,  1978  A THESIS SUBMITTED IN PARTIAL FULFILMENT THE REQUIREMENTS FOR MASTER OF  THE DEGREE OF  SCIENCE  in • THE FACULTY  OF GRADUATE  D e p a r t m e n t Of  We  accept to  this  thesis  the required  THE UNIVERSITY  J o h n A.  Botany  as c o n f o r m i n g standard  OF B R I T I S H COLUMBIA  August  ©  STUDIES  1985  Christy,  1985  OF  In  presenting  requirements  this f o r an  Columbia,  I  available  for  permission  for  her  of  of  August  the  shall  reference  and  study.  I  extensive granted  by  this thesis written  copying the It for  Head of is  this thesis my  Columbia  gain  the  of  British  it  freely  agree for  Department  understood  permission.  make  further  financial  Botany  1985  of  of  University  Library  The U n i v e r s i t y of B r i t i s h 2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  Date:  fulfilment  the  w i t h o u t my  Department  partial  that  representatives.  publication allowed  be  in  advanced degree at  agree  p u r p o s e s may or  thesis  that  that  scholarly  or  by  copying  shall  not  his or be  i i  Abstract Investigation cultivation, Limbella  bud  geography  tricostata  Significant  of and  length, mobility  habitat  (Sull.)  differences  frequency, leaf  morphology,  CM.  karyotype,  isozymes,  confirmed d i f f e r e n c e s and  Sciaromium  fryei  were o b s e r v e d e x p e r i m e n t a l l y  insertion  angle, leaf  of s u p e r o x i d e d i s m u t a s e  chromosome orientation.  Chromosome numbers o f b o t h s u b s p e c i e s i s n=11.  hypnaceous pachyloma  peristome, (Mont.) C M .  dispersal  from  Tertiary."  L^  from  tricostata  C_  dispersal Tertiary.  from  and or  has a Bryum i s thought its  S o u t h . America tricostata  Hawaii  ssp.  ssp. to  subspecies  branch  tricostata.  tricostata  a  shoot  in  fryei  ssp.  considered  and  Williams.  Sc iaromium  tricostata  is  areolation,  between  type  to Hawaii  after no  America  after  Limbella L.  and from  a L.  long-distance  later  i s thought  tricostata North  sporeling  t o have d e r i v e d  antecedent  fryei  of  than the  t o have  mid-  derived  long-distance  i n t h e mid-  to  late  iii  Table  Abstract L i s t of T a b l e s L i s t of F i g u r e s Acknowledgement  of C o n t e n t s  i i ix x xiii  .  Chapter I INTRODUCTION  2  A. SCOPE, PURPOSE AND OBJECTIVES OF STUDY  2  B. STUDY MATERIAL  2  C. TAXONOMIC HISTORY 3 1.1 H a w a i i a n P l a n t s 3 1.2 O r e g o n P l a n t s 3 D. PROBLEMS IN NEED OF INVESTIGATION 4 1. Taxonomic P o s i t i o n o f S c i a r o m i u m f r y e i B a s e d on L i m i t e d Data 4 2. D i s p a r i t y o f H a w a i i a n and N o r t h A m e r i c a n Habitats 4 3. D i s t r i b u t i o n i n N o r t h America 5 E . QUESTIONS TO BE ANSWERED  6  F. METHODS USED TO ANSWER QUESTIONS 1 . 3 Methods Used 1.4 R a t i o n a l e f o r Methods U s e d  6 6 6  G. BIOSYSTEMATICS OF BRYOPHYTES  °  7  Chapter II HABITAT  10  A. INTRODUCTION  10  B . HAW A11 AN ISLAND HABI TATS .' 2.1 C l i m a t e o f H a w a i i a n H a b i t a t s 2.2 S o i l s o f H a w a i i a n H a b i t a t s 2.3 H y d r o l o g y o f H a w a i i a n H a b i t a t s 2.4 V e g e t a t i o n o f H a w a i i a n H a b i t a t s C. NORTH AMERICAN HABITATS 2.5 C l i m a t e o f N o r t h A m e r i c a n H a b i t a t s 2.6 S o i l s o f N o r t h A m e r i c a n H a b i t a t s 1. S u t t o n Lake S o i l s 2. Barview S o i l s 2.7 H y d r o l o g y o f N o r t h A m e r i c a n H a b i t a t s 1. S u t t o n Lake H y d r o l o g y 2. Barview Hydrology 2.8 V e g e t a t i o n o f N o r t h A m e r i c a n H a b i t a t s  10 10 13 14 15 19 19 21 21 22 23 23 23 24  i v  1. 2.  S u t t o n Lake V e g e t a t i o n Barview V e g e t a t i o n  24 29  D. DISCUSSION 2.9 C o m p a r i s o n of C l i m a t e s 2.10 C o m p a r i s o n o f S o i l s 2.11 C o m p a r i s o n of H y d r o l o g y 2.12 C o m p a r i s o n of V e g e t a t i o n  31 31 32 32 33  E. CONCLUSION AND  33  SUMMARY  Chapter I I I MORPHOLOGICAL ANALYSIS  35  A. INTRODUCTION  35  B. NEED FOR QUANTIFICATION OF DISCRETE MORPHOLOGICAL CHARACTERS  35  C. MORPHOLOGICAL SIMILARITY OF HAWAIIAN AND NORTH AMERICAN LIMBELLA 36 D. SIGNIFICANCE OF DIFFERENT BRANCHING PATTERNS 3.1 C o n t r o l  of B r a n c h i n g  IN LIMBELLA .36 37  E. METHODS 3.2 Q u a n t i f i c a t i o n o f B r a n c h i n g P a t t e r n s 3.3 P h e n e t i c A n a l y s i s  38 38 39  F. RESULTS 3.4 Q u a n t i f i c a t i o n , o f B r a n c h i n g P a t t e r n s 3.5 P h e n e t i c A n a l y s i s  40 40 43  G. DISCUSSION  46  H. SUMMARY AND  CONCLUSIONS  46  C h a p t e r IV KARYOTYPE ANALYSIS  50  A. INTRODUCTION 4.1 U t i l i t y o f K a r y o t y p e A n a l y s i s 4.2 C y t o l o g i c a l R e p o r t s f o r L i m b e l l a  50 50 52  Taxa  B. MATERIALS AND METHODS 4.3 Chromosome Squashes and D o c u m e n t a t i o n 4.4 V a r i a t i o n s i n M e t h o d o l o g y D i c t a t e d by S t u d y  52 52 Material 56  C. RESULTS 4.5 K a r y o t y p e A n a l y s i s  58 58  D. DISCUSSION  67  V  4.6 4.7 4.8 4.9  S i g n i f i c a n c e o f N=11 Chromosome Number S i g n i f i c a n c e o f Chromosome L e n g t h D a t a S i m i l a r i t i e s Between t h e K a r y o t y p e s D i f f e r e n c e s Between t h e K a r y o t y p e s  E . CONCLUSIONS  AND SUMMARY  67 67 68 68 69  Chapter V ISOZYME BANDING PATTERNS  70  A. INTRODUCTION 5.1 U t i l i t y o f E l e c t r o p h o r e s i s 5.2 E l e c t r o p h o r e t i c P r o c e d u r e 5.3 E l e c t r o p h o r e s i s o f B r y o p h y t e s  70 70 71 71  B. MATERIALS AND METHODS 5.4 E l e c t r o p h o r e s i s 5.5 Enzyme A s s a y s  73 73 75  C. RESULTS 5.6 I n c o n s i s t e n t 5.7 The Enzymes  76 76 77  Staining  D. DISCUSSION A. CONCLUSION  AND SUMMARY  80 81  C h a p t e r VI PHENOLIC CHROMATOGRAPHY  83  A . INTRODUCTI ON  83  B. MATERIALS AND METHODS  84  C. RESULTS  85  D. DISCUSSION E . CONCLUSION  ...88 AMD SUMMARY  89  Chapter VII EXPERIMENTAL  CULTIVATION  91  A. INTRODUCTION  91  B. U T I L I T Y OF MOSS CULTIVATION  91  C. CULTIVATION OF LIMBELLA 7.1 Need f o r L i v e M a t e r i a l 7.2 Need f o r C u l t i v a t i o n S t u d i e s o f L i m b e l l a 1. • Whole P l a n t C u l t i v a t i o n 2. Spore C u l t i v a t i o n 3. Fragment R e g e n e r a t i o n D. MATERIALS AND METHODS  93 93 93 '...93 94 ....94 94  7.3 Whole P l a n t C u l t i v a t i o n 1. C u l t i v a t i o n of Hawaiian P l a n t s 2. C u l t i v a t i o n of N o r t h American P l a n t s 7.4 S p o r e C u l t i v a t i o n 7.5 Fragment R e g e n e r a t i o n 1. Living Material 2. Dried Material 7.6 R e c i p r o c a l C u l t i v a t i o n 7.7 Shoot O r i e n t a t i o n E x p e r i m e n t  ;  95 95 96 96 97 97 99 99 101  E . RESULTS 101 7.8 Whole P l a n t C u l t i v a t i o n 101 1. Hawaiian P l a n t s 101 2. North American P l a n t s 103 7.9 S p o r e C u l t u r e s , 103 7.10 Fragment R e g e n e r a t i o n 105 1. Living Material 105 2. Dried Material 111 7.11 R e c i p r o c a l C u l t i v a t i o n 111 1. H a w a i i a n P l a n t s i n R e c i p r o c a l C u l t i v a t i o n ...112 2. North American P l a n t s i n R e c i p r o c a l C u l t i v a t i o n 112 7.12 Shoot O r i e n t a t i o n E x p e r i m e n t 112 F. DISCUSSION 7.13 A r e M o r p h o l o g i c a l D i f f e r e n c e s Between t h e Taxa Stable? 7.14 A r e V e g e t a t i v e F r a g m e n t s V i a b l e D i a s p o r e s i n Limbella?  11-3  G. SUMMARY AND CONCLUSIONS  119  Chapter  . -  113 119  VIII  TAXONOMIC TREATMENT  121  A. INTRODUCTION  121  B. TAXONOMIC TREATMENT  121  C. COLLECTIONS OF TAXONOMIC AND HISTORICAL SIGNIFICANCE .140 8.1 E a r l i e s t C o l l e c t i o n o f L i m b e l l a t r i c o s t a t a 140 8.2 Type C o l l e c t i o n o f N e c k e r a t r i c o s t a t a S u l l 141 8.3 F i r s t C o l l e c t i o n o f L . t r i c o s t a t a S p o r o p h y t e s ....141 D. RATIONALE  FOR TAXONOMIC DISPOSITION OF SUBSPECIES  E . CONCLUSION AND SUMMARY  142 143  C h a p t e r IX PHYTOGEOGRAPHY  144  A. INTRODUCTION B. PRESENT DISTRIBUTION  144  9.1 D i s t r i b u t i o n  of L i m b e l l a  tricostata  9.2 D i s t r i b u t i o n  of L i m b e l l a  tricostata  ssp. t r i c o s t a t a 1 44 s s p . f r y e i ...150  C. ORIGINS OF DISTRIBUTIONS 157 9.3 The O r i g i n Of H a w a i i a n P o p u l a t i o n s 157 1. R e l a t i o n of L. t r i c o s t a t a t o South American Limbella 157 2. D i d L i m b e l l a D i s p e r s e from S o u t h A m e r i c a t o Hawaii? 158 9.4 The O r i g i n o f N o r t h A m e r i c a n P o p u l a t i o n s 162 1. A r e H a w a i i a n and N o r t h A m e r i c a n L i m b e l l a Closely Related? 162 2. Was L i m b e l l a C a p a b l e o f L o n g - d i s t a n c e D i s p e r s a l from H a w a i i t o N o r t h A m e r i c a ? .....162 D. DISCUSSION 165 9.5 E v i d e n c e S u p p o r t i n g L o n g - d i s t a n c e D i s p e r s a l i n Limbella 165 1. Are L i m b e l l a Spores Capable of Long-distance Dispersal? 168 2. C o u l d L i m b e l l a Have I m m i g r a t e d I n t o New Habitats Successfully? 169 9.6 A l t e r n a t i v e H y p o t h e s e s C o n c e r n i n g D i s p e r s a l t o N o r t h America 172 1. Was D i s p e r s a l P r e g l a c i a l ? 172 2. I s N o r t h A m e r i c a n L i m b e l l a P a l e o e n d e m i c ? ....174 3. D i d L i m b e l l a D i s p e r s e from N o r t h A m e r i c a t o Hawaii? 174 4. A r e H a w a i i a n and N o r t h A m e r i c a n L i m b e l l a Vicariads? 175 5. Was N o r t h A m e r i c a n L i m b e l l a I n t r o d u c e d by Man? 1 76 E . CONCLUSION  AND SUMMARY  176  Chapter X SUMMARY AND CONCLUSIONS  178  A. REVIEW 1. 2. 3. 4.  178 178 178 178 179  OF EXPERIMENTAL EVIDENCE Morphological Differences Karyotype D i f f e r e n c e s Isozyme D i f f e r e n c e s Cultivation Differences  B. ANSWERS TO QUESTIONS POSED IN CHAPTER 1 10.1 A r e H a w a i i a n a n d N o r t h A m e r i c a n L i m b e l l a Conspecific? 10.2 What E x p l a i n s t h e G e o g r a p h i c a l D i s t r i b u t i o n Taxa? C. CONCLUSION LITERATURE CITED  179 179 of the 180 ...180 182  APPENDIX A - SEQUENCE OF SSP. TRICOSTATA COLLECTIONS, AS REPRESENTED IN HERBARIA 199 APPENDIX B - PLANT SPECIES L I S T , LIMBELLA HABITAT AT SUTTON LAKE , LANE COUNTY , OREGON . 200 APPENDIX C - PLANT AND ANIMAL REMAINS IN F I L T R A T E FROM WASHINGS OF SSP. FRYEI SPECIMENS  201  APPENDIX D - MORPHOLOGICAL CHARACTERS TRICOSTATA AND SSP. F R Y E I .  202  MEASURED IN SSP.  APPENDIX E - BRANCH BUD FREQUENCY DATA MATRIX. TRICOSTATA, 21-37 = SSP. FRYEI  1-20 = SSP. 203  APPENDIX F - DISCRIMINANT AND MAHALANOBIS DISTANCE ANALYSIS OF BRANCH BUD FREQUENCY DATA 204 APPENDIX G - PHENETIC DATA MATRIX. 13-24 = SSP. FRYEI  1-12 = SSP. TRICOSTATA, 205  APPENDIX H - DISCRIMINANT AND MAHALANOBIS DISTANCE ANALYSIS OF PHENETIC DATA 206 APPENDIX I - CORRELATION MATRIX OF PHENETIC DATA  207  APPENDIX J - ABSOLUTE AND RELATIVE CHROMOSOME LENGTHS. 154 = SSP. TRICOSTATA, 155-275 = SSP. FRYEI  1208  APPENDIX K - DISCRIMINANT AND MAHALANOBIS DISTANCE ANALYSIS OF ABSOLUTE CHROMOSOME LENGTHS 209 APPENDIX L - CORRELATION MATRIX OF ABSOLUTE CHROMOSOME LENGTHS APPENDIX M - RECIPES AND GELS AND STAINS  SOURCES *  APPENDIX N - NORTH AMERICAN  FOR ELECTROPHORETIC  HABITATS SEARCHED  210  STARCH 211  FOR LIMBELLA. 215  ix  List  of T a b l e s  1. D i a m e t e r s and a g e s o f P y r u s f u s c a and S a l i x hookeriana i n L i m b e l l a h a b i t a t a t S u t t o n L a k e , Lane C o u n t y , O r e g o n . 27 2. L i v i n g m a t e r i a l o f L i m b e l l a studies 3. D r i e d m a t e r i a l u s e d  used  in regeneration 98  in regeneration  study  4. D e v e l o p m e n t a l  stages  of s p o r e l i n g c u l t u r e s  5. D e v e l o p m e n t a l  stages  of s i n g l e  6. D e v e l o p m e n t a l  stages  of shoot  7. D e v e l o p m e n t a l regeneration  stages  o f 1-8 cm s h o o t  leaf  regeneration.  100 104 ...107  t i p regeneration  108  segment  8. R e l a t i v e s t a b i l i t y o f m o r p h o l o g i c a l c h a r a c t e r s i n c u l t i v a t e d bryophytes, as r e p o r t e d i n l i t e r a t u r e .  109 ...115  9. P h y s i c a l p a r a m e t e r s i n f l u e n c i n g e x p r e s s i o n of m o r p h o l o g i c a l c h a r a c t e r s , as r e p o r t e d i n l i t e r a t u r e .  117  X  List  of  Figures  1. T e m p e r a t u r e and stations  precipitation records.for  2.  and h a b i t a t , c r o s s i n g , Na  3.  4.  Ssp. t r i c o s t a t a at A l a k a i T r a i l K a u a i , Hawai i  three  Hawaiian 12  Kawaikoi Stream t r i b u t a r y Pali-Kona Forest Reserve, 18  T e m p e r a t u r e and p r e c i p i t a t i o n r e c o r d s f o r two c o a s t a l Oregon s t a t i o n s . A. - N o r t h Bend ( m o d i f i e d from Loy 1976). B. - Newport ( m o d i f i e d f r o m C o o p e r 1958) Ssp.  fryei  habitat,  Sutton  Lake, Lane County,  20  Oregon. 25  5.  Ssp. f r y e i on d e c a y i n g l i t t e r S u t t o n L a k e . S c a l e = 1 dm  and  living  tree  roots, 26  6.  B r a n c h bud f r e q u e n c i e s i n s s p . , per l e a f segment number  t r i c o s t a t a and  ssp.  fryei 42  7.  S c a t t e r p l o t of measurements of 28 m o r p h o l o g i c a l c h a r a c t e r s . T r i a n g l e = ssp. t r i c o s t a t a , c l o s e d c i r c l e ssp. f r y e i  = 44  8.  S c a t t e r p l o t of measurements of 7 m o r p h o l o g i c a l c h a r a c t e r s , a f t e r s e l e c t i o n by d i s c r i m i n a n t a n a l y s i s . T r i a n g l e = ssp. t r i c o s t a t a , c l o s e d c i r c l e = ssp. f r y e i . 45  9.  Leaf i n s e r t i o n angles p l o t t e d a g a i n s t longest basal l e a f c e l l lengths. T r i a n g l e = ssp. t r i c o s t a t a , c l o s e d c i r c l e = ssp. f r y e i 47  10.  L e a f i n s e r t i o n a n g l e s p l o t t e d a g a i n s t stem d i a m e t e r s . T r i a n g l e = ssp. t r i c o s t a t a , c l o s e d c i r c l e = ssp. f r y e i . 48  11.  S o u r c e s of laboratory  living plants experiments  of  ssp.  tricostata  Chromosome f i g u r e s of  ssp.  tricostata  13.  Chromosome f i g u r e s of  ssp.  fryei  14.  Ssp. t r i c o s t a t a chromosome a l i g n m e n t s . 1-10 C h r i s t y 5368. 11-14 - K a u a i , C h r i s t y 5337 Ssp.  fryei  for 54  12.  15.  used  chromosome a l i g n m e n t s .  ( Christy  ( Christy  1-6  5368  5505 ). -  ). 59 ...60  Kauai,  - Oregon,  62 Christy  xi  5499. 7-11 16. 17.  18.  - Oregon, C h r i s t y  5505  63  A b s o l u t e chromosome l e n g t h s , p l o t t e d a g a i n s t l e n g t h s , a l l chromosomes m e a s u r e d  relative 64  A v e r a g e a b s o l u t e chromosome l e n g t h s p l o t t e d a g a i n s t average r e l a t i v e l e n g t h s . T r i a n g l e = ssp. t r i c o s t a t a . Closed c i r c l e = ssp. f r y e i  65  G r a p h of a v e r a g e r e l a t i v e l e n g t h s of chromosome complements of s s p . t r i c o s t a t a and s s p . f r y e i  66  19.  Zymograms of s s p . t r i c o s t a t a and s s p . f r y e i . Dark z o n e s = enzymes s t a i n e d f o r ; w h i t e z o n e s = SOD a c t i v i t y . A l l g e l s run i n Ridgway b u f f e r s y s t e m 78  20.  Phenolic  21.  P h e n o l i c s p o t p a t t e r n s of s e q u e n t i a l l y - s p o t t e d on o n e - d i m e n s i o n a l c h r o m a t o g r a m s  22.  Means of a t t a c h i n g s h o o t segments t o shoot o r i e n t a t i o n experiment  23.  spot  patterns  on  two-dimensional  filter  25. 26.  paper,  Growth r a t e s of s p o r e l i n g c u l t u r e s , by d e n s i t y - 2 s p o r e l i n g s cm" . B. - 1 s p o r e l i n g c m . C. s p o r e l i n g s cm" . D. - 27 s p o r e l i n g s cm" -2  2  fractions 87 for 102  2  24.  chromatograms. 86  class. - 43  106  2  Shoot o r i e n t a t i o n e x p e r i m e n t . Ssp. s s p . f r y e i on r i g h t  tricostata  on  left, 114  Ssp. t r i c o s t a t a , d e n d r o i d p l a n t s . N o t e b r a n c h i n g ( C h r i s t y 5196 )  monopodial 129  Ssp. t r i c o s t a t a , t r a i l i n g a q u a t i c p l a n t s . B r a n c h e s s p r e a d t o show b r a n c h i n g p a t t e r n ( C h r i s t y 5245 ). tricostata,  upper b r a n c h  27.  Ssp.  leaves.  28.  Ssp. t r i c o s t a t a . A. c e l l s (both Fosberg  29.  Ssp. f r y e i , d e n d r o i d p l a n t s . N o t e m o n o p o d i a l C h r i s t y 5334 )  30.  Ssp.  fryei,  trailing  31.  Ssp.  fryei,  upper branch  32.  Ssp. f r y e i . A. - upper ( b o t h C h r i s t y 5338 )  - upper l e a f 10233 )  plants  leaf  (Fosberg  cells.  ( Christy  leaves  ( Frye  cells.  A.  B.  B.  ..130  10233  - basal  ). 131  leaf 132  branching ( 136  5338 )  1 37  s.n.)  138  - basal  leaf  cells 139  xi i  33.  S u r f a c e and e l e v a t i o n a l on H a w a i i , H a w a i i  distribution  of  ssp.  tricostata 145  34.  S u r f a c e and e l e v a t i o n a l on K a u a i , H a w a i i  distribution  of  ssp.  tr icostata 146  35.  S u r f a c e and e l e v a t i o n a l on M a u i , H a w a i i  distribution  of  ssp.  tricostata 147  36.  S u r f a c e and on M o l o k a i ,  elevational Hawaii  distribution  of  ssp.  tricostata 148  37.  S u r f a c e and e l e v a t i o n a l on Oahu, H a w a i i  distribution  of  ssp.  tricostata 149  38.  S i t e of e x i s t i n g s s p . Lane C o u n t y , Oregon  39.  T.C. F r y e ' s map and n o t e d e s c r i b i n g Sciaromium f r y e i W i l l i a m s  40.  Type l o c a l i t y of Sc i a r o m i u m Coos C o u n t y , Oregon  41.  S i t e s of h a b i t a t s Lake. B = Barview  42.  World d i s t r i b u t i o n  fryei  searched  population at Sutton  fryei  the  type  Williams,  Lake, 151  locality  of 153  Barview, 154  for ssp.  of L i m b e l l a  fryei.  SL  =  Sutton 156 159  Acknowledgement  I  e x t e n d warm and h e a r t f e l t  Schofield,  f o r the moral  time  Botany this due  at  UBC. I am a l s o  Department, thesis  t o K.M.  Cole,  expertise indebted  w i t h o u t whose  could  supervisor, support  given  W.J. H u n t e r ,  J.R.  Spence,  committee  members F.R. G a n d e r s  t o W.J. Hoe f o r a s s i s t a n c e  W i l f r e d B. extensive throughout  t o many members o f t h e UBC  assistance  and  encouragement  S p e c i a l thanks are  T.T. M c i n t o s h , K.W.  N.L. V o g t  and  so f r e e l y  n o t have been c o m p l e t e d .  Ritland,  due  t o my  and l o g i s t i c a l  bryological-bibliographical my  thanks  Nicholls,  K.  and R.E. de Wreede, and t o my  and B.A. Bohm. i n Hawaii.  Thanks  are  also  1  TO  D a v i d Dwight whose d i s c o v e r y  Baldwin  (1831-1912),  l a y unknown  f o r 105 y e a r s  and  Stephen Fowler C h r i s t y whose l o v e o f e n i g m a t a  (1910-1982),  l e d me  thither.  2  I.  A.  SCOPE, PURPOSE AND OBJECTIVES OF STUDY This  investigation  phytogeographical the  was c a r r i e d  questions  North  o f Sc i a r o m i u m  America  to determine  identities  phytogeographical  B.  fryei  (Oregon).  o u t t o r e s o l v e t a x o n o m i c and  r e g a r d i n g the  H a w a i i a n moss Sc i a r o m i u m  identity  taxonomic  limits  tricostatum (Sull.) Mitt. Williams  of  Seven a r e a s and l i m i t s  coastal  e x p l a n a t i o n of t h e i r  present  and t h e  northwestern  of i n v e s t i g a t i o n  of the taxa  of  were  used  and t o p r o v i d e  a  distribution.  STUDY MATERIAL Limbella  is  Amblystegiaceae. aquatic five up  INTRODUCTION  major  grows  dendroid  pleurocarpous  Limbella  to semi-aquatic Hawaiian  t o 0.5 m l o n g  fryei  a  tricostata  i n mountain  islands.  (Figs.  genus ssp.  in  25,26).  trailing  the f a m i l y  tricostata  is  s t r e a m s and i s common on t h e  Plants are t r a i l i n g  i n a dense s h r u b - c a r r  to occasionally  moss  Limbella in coastal (Figs.  or dendroid,  tricostata Oregon.  29,30).  ssp.  Plants are  3  C.  TAXONOMIC HISTORY  1.1  Hawai i a n  Plants  Limbella tricostata as  Sull.  Sc iaromium  considered  as  authors assigned  1.2  to  between  comb. nov.  described  and has been  (Sull.)  Mitt.  Hawaiian  as N e c k e r a  known  Until  Islands.  even t h o u g h t h e y had been  1980).  Seven  genera i n 3 d i f f e r e n t  recently  1971  Both  i t was  male  and  reported  c o l l e c t e d as e a r l y  synonyms were c r e a t e d  1889-1912, and o v e r t h e y e a r s  to 5 d i f f e r e n t  by  various  t h e t a x o n had  been  families.  Plants  Limbella  single  the  was  were known but s p o r o p h y t e s had n e v e r been  (Christy  Oregon  1854)  tricostatum  the l i t e r a t u r e , 1875  (Sull.) CM.  (Sullivant  endemic  female p l a n t s in  tricostata  was  tricostata described  collection  discovered  in  Sc i a r o m i u m  fryei  1978  ssp.  fryei  as Sc i a r o m i u m  (Williams (Christy  1933). 1980).  (Williams)  fryei A  Williams  Christy from  second p o p u l a t i o n Lawton  t o a synonym o f S_^ t r i c o s t a t u m .  (1971)  a was  reduced  4  D.  PROBLEMS IN NEED OF  1.  Taxonomic Limited  Lawton's  P o s i t i o n of Sc iaromium  tricostatum  Unfortunately,  would  s.n.,  have  3.xi.l933;  Lawton  would  decision and  on  she  did  solely not  2 s p e c i m e n s o f S_^  had  )  and  t o borrow number of are  2  and  morphological  At  that  time,  tricostatum  ( Neal  isotypes  of  S_^  additional material specimens.  superficially  habitat  as a  i n d i c a t e t h e number of  -- a p p a r e n t d i f f e r e n c e s i n  pattern  on  fryei  called  fryei•  or base h e r  Although  Hawaiian  similar  —  even  growth  form,  size,  for a re-evaluation  of  taxonomic r e l a t i o n s h i p .  2.  D i s p a r i t y o f H a w a i i a n and N o r t h A m e r i c a n  Habitat  conditions  most - r e s p e c t s  plants  for Limbella  strikingly  grow submerged  and  based  S v i h l a 2309  plants  microscopically  their  had o n l y  have  Oregon  was  she had b a s e d h e r o p i n i o n .  on a l i m i t e d  branching  Based  Data  s p e c i m e n s upon w h i c h WTU  fryei  (1971) d e c i s i o n t o submerge Sc i a r o m i u m  synonym of S^ grounds.  INVESTIGATION  dissimilar.  o r emergent  on r o c k s  of t h e e x i s t i n g Oregon  r o t t e n wood, and no  Accompanying dissimilar.  vegetation The  rocks of  i n H a w a i i and Oregon Hawaiian  the  plants  i n streambeds.  population are  Habitats  two  in  areas  H a w a i i a n p l a n t s a r e most common  usually  In c o n t r a s t ,  grow on wet  present  are in  peat, bark  the  habitat.  is  completely  between  1200-  5  2130  m  elevation,  Although  i t i s not  range  t o grow on  center  of  and  £L  their  fryei  raised  the  Lawton had  3.  (1971) fryei  of  a  80  that  plants  km  Sc iaromium  fryei  was  whether  proposed had  the  from the had  type  been  sea  edge  level.  of  their  from t h o s e o c c u p i e d Sc iaromium  other  in  the  tricostatum  apparent d i f f e r e n c e s , conspecific  as  population  of  usual  a rare  means by  introduced  answers  of  concerning  b e c a u s e the  in  be  critically.  by  man in  from an  Hawaii.  undisturbed  raised considerable I t was  used  which a l i e n  the  commercial  plants  introduced  are  distributional origin  was  that  f o r hobby or  n a t i v e moss f l o r a  the  doubt  also unlikely  I t seemed more l i k e l y  the  species  Oregon  1978,  locality,  areas.  element  the  introduced.  I n v e s t i g a t i o n of  been s t u d i e d  America  would  geographical  especially  the  t h e y were r e a l l y  that  been  tricostatum  Northwest. provide  with  second p o p u l a t i o n  habitat  purposes,  at  d i s p a r i t y in  D i s t r i b u t i o n in North  Discovery  new  different  habitats, coupled of  p l a n t s grow a t  for plants  substrata  range, the  Oregon  proposed.  Sc iaromium  into  uncommon  question  Lawton  the  whereas  and  so p o o r l y  status  of  that  the  Pacific  anomaly of  known and  S.  S_^ had  could f ryei, never  6  E.  QUESTIONS TO BE ANSWERED  The to  F.  problems o u t l i n e d  above r a i s e d  two  specific  questions  be a n s w e r e d by t h e r e s e a r c h :  1.  Were Sc i a r o m i u m  t r i c o s t a t u m and S^  2.  What e x p l a i n e d t h e i r  METHODS USED TO ANSWER  geographical  fryei conspecific?  distribution?  QUESTIONS  1.3 Methods Used In  order  compared  in  to  answer  s i x areas  morphology  (including  multivariate  analysis)  banding  patterns  experimental  the of  (3)  (5)  was  phytogeographical  karyotype  habitat  microscopy  morphology  (4)  chromatography  Evidence  synthesized  (1)  electron  phenolic  treatments  questions, the taxa  investigation:  scanning  cultivation.  investigation  above  to  from  (2)' and  isozyme  and  (6)  areas  of  taxonomic  and  these  formulate  were  t o answer t h e above q u e s t i o n s .  1.4 R a t i o n a l e f o r M e t h o d s Used Because W i l l i a m s ' seemed  to  differences, fryei  and  be  based  (1933) d e s c r i p t i o n  more on g e o g r a p h i c a l t h a n  and b e c a u s e S_^  of  Lawton's  tricostatum  seemed  (1971) less  Sc iaromium  fryei  on m o r p h o l o g i c a l  combination convincing  of  S.  upon r e -  7  examination study,  the problem,  methods  approach, to  of  in  in order  make  i t was  addition  d e s i r a b l e to apply,  to  the  traditional  decisions.  approach probably  will  other  taxonomic  methods have begun t o be  during  the  last  cultivation,  20  which  Biosystematics biologically  based"  synthesizing  data  in  contrast  morphology  —  morphological  of  bryotaxonomy,  a p p l i e d to include  methods  bryophytes  biochemistry,  numerical  taxonomy  collectively  i n b o t a n y were f i r s t  and  called  a p p l i e d to vascular  1982).  limited  1976;  Smith  1982;  BRYOPHYTES is  generally  approach from one  to  Anderson's  Wyatt  and  Stoneburner  biosystematic  methods  reasons  biosystematic  behind  why those  to  reviews  of v a s c u l a r p l a n t s .  1984),  [and  (Anderson  and of  on  elucidate]  differentiation"  of moss  1978,  above  solely  biosystematics 1963;  Stotler  1984;  i n c l u d e d a wider  i n bryotaxonomy, studies  based  its  Szweykowski 1984)  (Vickery  organisms  (1964) r e v i e w  1978;  "experimental,  t h e methods l i s t e d  taxonomy  leading  Other  an  systematics  or more of  processes  Koponen  d e f i n e d as  "a g r o u p of  to cytology.. 1978;  to  traditional  to c l a s s i f y  evolutionary  (Longton was  genetics,  which  the  cornerstone  T h e s e methods  with  systematics.  BIOSYSTEMATICS OF  the  the  physiology,  "biosystematics",  G.  years.  cytology,  experimental  plant  remain  Although  this  morphological  to o b t a i n a d d i t i o n a l i n f o r m a t i o n  taxonomic  in  Longton range  speculated  b r y o p h y t e s have  on  of the  lagged  8  Longton  (1982)  biosystematic piecemeal  techniques  to d i f f e r e n t  increasingly bryophyte  biological  of  biosystematic  a  the  solve  --  specific  isozyme  not  could  yet  often  approach  to  a  two  taxonomic  problems.  Szweykowski  a d v a n c e d by an  one is  or now  both i n v o l v i n g  such  Based  on  methods evidence  Krzakowa  1981;  (Szweykowski  and  Krzakowa  Newton  1981)  and  s e r o l o g i c a l i n v e s t i g a t i o n s (Szweykowski e t a l .  1981;  Zielinski  addition  to  of  the  B a s e d on plant provide  Zehr  isozyme,  redefine  investigation  biosystematic  data,  three  of  methods,  a  (Mendelak  (Szweykowski  (1980) u s e d  cytological sections  Limbella was  phenolic 1981;  r e l a t i o n s h i p s between c e n t r a l  have been c l a r i f i e d  phenolic,  investigations  limits  1981),  to morphological  terpenoid,  This  taxa  al.  cytological  1981),  from  al.  1977),  et a l .  1978,  to  et  et  Zielinski  of  groups."  bryophytes  three  1977;  array  applying  —  the  Szweykowski  European P e l l i a  1981;  be  studies  have used more t h a n  have become  selected  of  applied  systematic...  research  group  only  been  [using to  "various  studies  most e f f e c t i v e l y  I know of  (Krzakowa and  that  sufficiently  simultaneously]  methods  that  too  volume of p u b l i s h e d  large,  liverworts  all  comprehensive  biosystematic  relatively  have  but  techniques  Although  observed  b r y o p h y t e g r o u p s : our  systematics  application  two  astutely  1984).  evidence and  i n the  from  cultivation  genus  tricostata,  s i m i l a r attempt  In  Scapania.  using  to redefine  six the  taxon. the  taxonomy,  c o n t r i b u t i o n s of it  is  clear  a d d i t i o n a l information  biosystematics  that  to  biosystematic  w h i c h can  alter  or  vascular  studies  augment,  can but  9  not  necessarily  traditional  supplant,  morphological  systematic  analysis.  interpretations  based  on  10  II.  A.  HABITAT  INTRODUCTION This  chapter  characteristics  describes  of  the Hawaiian  Limbella.  Analysis  unexpected  physical  of  these  that  2.1  C l i m a t e o f Hawai i a n H a b i t a t s  climate, location air  Hawaiian  Islands  influenced a t 20°  between t h e two fryei  by  are  habitats  of  emphasizes  localities  originated  from  and  long-  tr icostata.  the  renowned  Pacific  north l a t i t u d e ,  topography  precipitation  a t any  mild causes given  Ocean  temperatures wide  and  where  i s greatest  on  the  between and  subtropical  the  islands'  warm t h e  year-round.  variation  place  i n mountains  precipitation  for their  b o t h of w h i c h  o c c u r s most commonly  the  floristic  ISLAND HABITATS  masses t o p r o d u c e  diverse  North American  d i a s p o r e s of s s p .  HAWAIIAN  and  characteristics  ssp.  B.  The  physical  and  similarities  s u p p o r t s the assumption distance dispersed  the  regional However,  i n temperature  and  islands.  Limbella  1200-2130 m  elevation,  temperatures  cooler  than i n  lowlands. Annual  montane  temperature  stations  on  and three  precipitation islands  —  records a l l  near  for  three  Limbella  11  localities Limbella below  -- a r e shown i n F i g .  precipitation  Data  of  show an  of  rainfall  not  restricted  Price  1972;  Carlquist  rain  from  both  is  1980).  weather  Mt.  Waialeale,  12  rainiest  site  known on e a r t h  m  the  precipitation  f o r m i n g between  always  1980;  shrouded  occurs  throughout point  vegetation  precipitation  ( B l u m e n s t o c k and  and  where  the  is  year Kauai,  reportedly  the  1972).  decreases  610-1520  m,  ( B l u m e n s t o c k and  In  At  islands,  on  1500-2100 m  on  visits  the  b e n e a t h a t r a d e wind  clouds.  the  systems.  development  condensation  D u r i n g my  by  to  At as  1972).  between  1983).  the  the annual r a i n f a l l  annual p r e c i p i t a t i o n  Whiteaker in  restricted  ( B l u m e n s t o c k and P r i c e  ( B l u m e n s t o c k and P r i c e  with cloud  weather  Blumenstock  annually,  little  rainfall  m,  the  and March  over  highest  2100  in  with  marked  1962;  variable  e l e v a t i o n s above  Carlquist  November  occurs,  still  over  layer  orographic,  a r e a s (Mink  fall  reduction  anticyclonic  in precipitation,  systems  but  or  In most windward m o u n t a i n s ,  receives  associated  a  where  24°C  coincides  i s widely d i s t r i b u t e d  distributed  Maximum  exceed  show  Between  t o mountainous  as 380 mm  mountains  s t o r m s of l o w - p r e s s u r e weather  time the r a i n f a l l  1C).  sites  high-pressure  increase  this  (Fig.  the  to October, which  slopes.  cyclonic  more e v e n l y  these  t i m e most  mountain  montane s i t e s advent  typical  At t h i s  higher  from  from A p r i l  winds  systems. to  In  i s most common, t e m p e r a t u r e s r a r e l y 10°C.  trade  1.  Mountain this  inversion  Price  1972;  peaks a r e almost  zone,  contributes  frequently  fog  appreciably  drip  from  to  total  Price.1972).  to h a b i t a t s  of L i m b e l l a  on Maui  and  Kauai  12  Figure  1 - Temperature  and  precipitation  Hawaiian A.  - Makawao, M a u i ,  [Kokee], elev.  466  Kaui, m.  elev.  for three  stations.  elev. 1097  records  640 m.  A l l graphs m o d i f i e d  m. C.  B.  -  - Mountain  from A r m s t r o n g  Kanalohuluhulu View, H a w a i i , 1983.  1 3  (June  1983,  May  temperatures  2.2  Soils  water  Hawaiian  sequence  are  basaltic varying  I s l a n d s are of  over  a  "hot  parent age  of  to  different soil  weathered,  the youngest  y e a r s a g o ) , and  the  developed The  soils  islands  the p r i n c i p a l  intensely  near  tectonic  formation  t o ' b e o f P l i o c e n e age  most is  Of  having  material.  and  locality.  15-22°C  igneous o r i g i n ,  spot"  Pacific  residual,  weathering  were  and  air  Habitats  northwesterly-shifting soils  temperatures  18-24°C.  of H a w a i i a n  The in  1984),  plate. from  which  has  the  Hawaiian  underlying the  amount  of  o c c u r r e d a t any  one  the  Kauai,  years),  considered  i s the o l d e s t  w i t h the b e s t - d e v e l o p e d s o i l s .  formed  d u r i n g the  has  least-developed soils  the  because  islands,  (5.7 m i l l i o n  formed  of  Most  are d i v e r s e ,  island, the  center  determines  existing  having  Pleistocene  and  Hawaii (450,000  (Macdonald  et a l .  1983). In  the mountains,  occasionally on  peat  i t i s f o u n d on  i n boggy  forests.  Swamp, a w e l l - c o l l e c t e d Foote  et a l .  "extremely occurring  soil  usually  A histosol  locality  ( p r o b a b l y pH  on m o u n t a i n t o p s , near  on M a u i ,  Olinda, was  series  of L i m b e l l a , underlain  wet  r o c k s , but  stream  beds  i n Kauai's was  Alakai  described  by c l a y  and  or  by  basalt,  4.5-5.0) and v e r y p o o r l y d r a i n e d ,  r i d g e s and another  described  grows on  in intermittent  (1972) a s mucky p e a t  acid"  association Limbella  Limbella  steep slopes. well-collected  by F o o t e  et  al.  A  latosol  locality (1972)  as  for a  14  deep, w e l l - d r a i n e d very  steep  latosol Wreede  soil  ground.  of moderately Sherman  group as a c i d  usually  occurs  usually  has  in  and  herbarium  s p e c i m e n s were d e s c r i b e d  perennial  streams. as  streams  For aquatic  described  been  or  the  youthful  perennial  soils  Surface  runoff  during  increase  r a p i d l y during  occur that  in  first-  and  are frequently  populations plants.  that  On r o c k s  which a b r u p t l y unable Maciolek  to  heavy  stream  flows  i s extremely v a r i a b l e .  rainfall  flash  In  (Stearns  The  second-order  exist  streams  measurements  raising  (Mink  greatest  volume  (sensu  from  soaking  pH 6.0-7.0 i n s m a l l  lakes  changes 1945)  Limbella  non-aquatic  (Fig.  these  Stearns  Horton  Limbella line  stream  1962;  as t e r r e s t r i a l ,  streams,  1966).  may be low, b u t may  flooding  10-15 cm a b o v e water  (1967) r e c o r d e d  o r emergent  t e m p e r a t u r e s as c o l d t o  rains, irregularly  perennial  pH  Most  i n d r y beds o f i n t e r m i t t e n t  of the i s l a n d s  1983).  otherwise  cease  obtain  aquatic.  intermittent, periodically  in  De  a r e h i g h l y p e r m e a b l e and p e r e n n i a l  moderate  a n d sometimes c a u s i n g Macdonald et a l .  ash.  intermittent  as e i t h e r a q u a t i c  i n streams  i n many p a r t s  1966;  and  considered  exposed  streams a r e r a r e  levels  of t h i s  a pH o f 4.5-5.5.  plants, c o l l e c t o r s described  The volume o f water  general,  soils  from v o l c a n i c  s w i f t , slow o r n o n - f l o w i n g , and water  warm.  on s l o p i n g t o  Habitats  streams,  in  texture  montane l a t o s o l s w i t h  of Hawaiian  Limbella  (1972)  (pH 5 . 0 ) , d e r i v e d  (1967) d e s c r i b e d  2.3 H y d r o l o g y  fine  forms  mats  2).  I was  streams, at  2040  but m,  1 5  and  running  streams at  this  e l e v a t i o n p r e s u m a b l y have a  similar  pH.  2.4  Vegetation  of Hawai i a n  Limbella  habitats  climatologically  and  Habitats  are  geographically,  edaphically  from H a w a i i , K a u a i ,  Maui, Molokai  90  (Chapter  to  over  12,000 mm  2130  precipitation.  (Appendix  A)  associated  with  can  be  at  9)  As  Limbella.  and  Limbella  Oahu, from  rule,  elevations  only  recent  However, v e g e t a t i o n way  of  the  the  760-  vegetation habitats  b e c a u s e most c o l l e c t i o n s where  of  collectors  a d e q u a t e d e s c r i p t i o n s of  in a general  i s known  from h a b i t a t s r e c e i v i n g  1200-2130 m e l e v a t i o n ,  vegetation  are is  uniform.  B a s e d on my recent  observations  herbarium  Limbella  i s one  communities (1983).  Styphelia  collections,  of  described  are  in part  Acacia  tameiameia  Molokai (  was  koa  by  most  Krajina  Gray,  (Cham.) F.  described  Bishop  because E ^ cumini species,  the  several Metrosideros  C h e i r o d e n d r o n , D r y o p t e r i s , and  forest"  f r o m Maui and  Common a s s o c i a t e s , d e p e n d i n g  elevation,  on  diverse.  and  a  have p r o v i d e d  described  from s i t e s fairly  m  topographically,  typical  s.n.,  Athyrium.  as  low  and  typical  data  habitat  p o l y m o r p h a Gaud. (1963) grazing  and  "mesic  from of  forest  Whiteaker  disturbance  S a d l e r i a cyatheoides  Muell.,  and  Kaulf.,  Coprosma, One  low-elevation  Eugenia  22.i.1967, BISH).  (L.) D r u c e and of  on  Kauai,  Schinus are  This  site  cumini-Schinus i s of  weedy,  interest  introduced  e l e v a t i o n s where n a t i v e p l a n t s have been  1 6  replaced 1911).  almost  completely  At s e v e r a l  Cryptomeria  by  an  alien  p l a c e s up t o 2000 m,  japonica  D.  Don  flora  exotic  (vid.  trees  Forbes  such  as  and E u c a l y p t u s a r e t h e dominant  spec i e s . Limbella along soil  the  most commonly  banks.  a n d wood.  irrigation On  near  Haleakala. bedrock, ferns  a l s o have been  d i t c h e s a n d wooden I  observed  one,  the p l a n t s  of  where  p o o l s o f water  Limbella  occurring  intermittent plants  stream  among  grew on damp s o i l  streambed.  There  in  was  perennial,  and M o h i h i  Streams,  abundant,  in  tens of meters m long  the L i m b e l l a green  to  shade  of  (b) on t h e lowermost accumulate  bedrock dense no  understory  evidence  of  sheets  of  overhanging exposures of  and (c) around  depressions.  rocks  slope  In  small  another  vegetation, the  were  present  i n the  of p o o l e d water.  In both  %,  fern  and  shrub  20-85 %-. In  0.5  roots,  man-made  NW  restricted  c a s e s , M e t r o s i d e r o s c a n o p y c o v e r was 75-80 cover  tree  from  Stream,  (a) i n the  and no  on t r e e s ,  and  populations in intermittent  was  r e c e d i n g waters remaining  streambeds  collected  Waikamoi  o r s h r u b s a l o n g t h e bank  bedrock,  in  flumes.  several  the headwaters In  rocks  I t occurs occasionally  Plants  Maui  streams  grows on  algae.  third-order  Na P a l i - K o n a F o r e s t  places  covering  of stream  (Fig.  streams  2).  on K a u a i  Reserve),  a l l submerged  i n streambeds  Limbella  is  shoots  up  to  4-20 m w i d e , most o f  i s a q u a t i c and i t i s o f t e n mixed M e t r o s i d e r o s canopy c o v e r  Kawaikoi  r o c k s and c h o k i n g  channel with undulating Here,  (e.g.,  ranges  with from  filamentous 0-30 %.  Perennial,  first-order  s t r e a m s on K a u a i  (e.g.,  near  foot  1 8  Figure 2 - Ssp. t r i c o s t a t a and h a b i t a t , K a w a i k o i Stream t r i b u t a r y a t A l a k a i T r a i l c r o s s i n g , Na P a l i - K o n a F o r e s t R e s e r v e , Kauai, Hawaii.  19  Pihea  Trail  Trail,  Na  densely  and  near  gauging  Pali-Kona Forest  shaded  station  Reserve)  on  great  i n h i g h e r - o r d e r streams.  NORTH AMERICAN  2.5  C l i m a t e of North American The  and  seasonal, the  of  winter  by  occurring  extended  and  p r e s s u r e weather precipitation temperature northeast and  below clear  on  or  (70  the  5°C and  km  snowfall  both  i s never  oceanic  Ocean, w h i c h  Precipitation  as  violent  is  in  July  and  records  population  for  of s s p .  in  Fig.  but  temperatures  In  winter,  i s rare.  year, the remainder  caused  by  which  storms.  High-  skies,  and  August.  Annual  N o r t h Bend  (11  fryei  at  population 3.  Summer  rarely  exceed  temperatures  Cooper  markedly  offshore,  cyclonic  and  moderates  and A p r i l ,  systems  n o r t h of the e x i s t i n g  coast,  i n one  Biomass  i n t h e summer m a i n t a i n c l e a r  precipitation  of the e x t i n c t  is  weather  negligible  fogs are frequent.  days  a l o n g t h e streambank,  between O c t o b e r  a t S u t t o n L a k e ) a r e shown  occurs cold  and  Newport  fryei  is  the P a c i f i c  occasionally  systems  is  gullies  Limbella  Oregon  temperatures.  p r e s e n c e of l o w - p r e s s u r e  form  narrow  Habitats  coastal  influenced  most  Koaie-Waialae  HABITATS  climate  Mediterranean, summer  occupy  submerged r o c k s i n t h e s t r e a m .  C.  of  or v e g e t a t i o n .  r o c k s i n s p r a y zones  aquatic as  often  by o v e r h a n g i n g c l i f f s  e m e r g e n t , on m o i s t  at foot  w i t h f o g on  Barview) of s s p . drought 20°C  seldom  (1958) r e p o r t e d  km  and fall  only  58 d a y s ,  47  rain  20  Figure 3 - T e m p e r a t u r e and p r e c i p i t a t i o n r e c o r d s f o r two c o a s t a l Oregon s t a t i o n s . A. - N o r t h Bend ( m o d i f i e d from Loy 1976). B. - Newport ( m o d i f i e d from C o o p e r 1958).  21  on  68,  cloud  Both  Cooper  variation general  and  at  being a  maximum  and  During  2.6  Loy  with  and  to were  S u t t o n Lake  Sutton  Lake was  (1976) o b s e r v e d  increasing  Sutton  Lake  14-19°C and  formed  peat  The  becoming  The  1 m-deep bed  pole.  less  composed o f  which  of  fine  was  fibrous  the  a b o v e and root  of  t h e method o f L a v k u l i c h  cm.  absent.  and May  1984,  14-16°C.  dune to  advances the  • east  developed  by p r o b i n g w i t h a a  Dried  depth The  on  of  1 m  c.  peat  4.8  10 cm  depth  4.7  a t 20 cm  dm,  readings  at the s u r f a c e depth.  layer  a n a l y z e d f o r pH  ( 1 9 8 1 ) , gave a v e r a g e  w a t e r ) o f 4.9  the  fibrous material is  samples  distilled  an  range 5  A b l a c k , mucky humic  30 ml  and  sand  Range  peat: at  1983  both  extreme,  fogs are  Limbella  below.  remains.  following  layer,  1-5  Coast  f i b r o u s at  the  g  inland:  medium- t o l i g h t - b r o w n p e a t ,  occupies  (5  top  km  and  contrast,  more  summer  i n June  In  a i r temperatures  determined  i s densely  sedge and  latitude.  when p o s t - g l a c i a l  population  of  precipitation  Soils  (Cooper  thickness  in  the only  Habitats  drained  adjoining  climatic  latitudes,  become  the c o o l i n g  b l o c k e d a stream which 1958).  little  o c c u r s o n l y a few  temperatures  of N o r t h American  1.  115.  increase  increases, visits  on  gradual  in conditions  temperatures  Soils  cloud  of d i f f e r e n t  minimum  precipitation  and  stations  of t e m p e r a t u r e s  an a b r u p t change  water  78  (1958) and  coastal  trend  cooling  f o g on  Sand,  humic gravel  22  and  rocks  underlain  are by  2.  origin  of many t h a t eustatic  occur  sea  the  late  gravel,  peat  podsols  develops at drainage soils,  Service  the  of  are  coast,  these  mostly  In F r y e ' s  evidence  or  with  isotype  (caddisfly)  an  larval  in  one  part  by  maxima and  (Allen  minima  and  Baldwin  ancient  sand w i t h  beach  some c l a y ,  underlain  by  marine  usually classified  al.  perennial  1969), pools.  marine  terrace  a pH  5.0  (UBC)  of I  case  f o r l o c a t i o n of  sandy m a r i n e t e r r a c e a t  is  iron-cemented  (Jenny et  similar  formed  late  1945).  Typically,  from a  terrace  composed of  t e r r a c e s are  vernal  The  glacial  remains,  (Griggs  oligotrophic  supporting  1944).  t e r r a c e s are  30-76 cm  forming  1970).  are  age.  a m a r i n e t e r r a c e of  Pleistocene  woody  age  d e p t h s of  are  trichopteran sand,  soils  described  Barview,  and  on  during  and  spodosols.  and  dune d e p o s i t s  s a n d s t o n e of E o c e n e  Pacific  and  The  Pliocene  or  ancient  Baldwin  changes  deposits,  sandy  and  occurred  the  Pliocene  layers  s a n d s t o n e of The  along  1981).  offshore  and  ( A l l e n and  level  Baldwin  and  peat  Soils  Barview p o p u l a t i o n  Pleistocene  1944;  The  marine s i l t s t o n e  Barview  The  during  absent.  Barview.  60  (Soil  found  the  type  hardpan impeding Blacklock km  S  of  Conservation  remains  constructed  as  of  a  entirely  of  locality  on  23  2.7  Hydrology  1.  of N o r t h  S u t t o n Lake  At  Sutton  surface,  Lake  even  Habitats  Hydrology t h e water  saturating  depressions  American  the  in dry  level  peat  and  i s 1-15 forming  summer months.  and  tree boles  as  as  the  well  trichopteran Probably  larval  the  percolation  i n the v i c i n i t y  presence cases  entire from  the  never  shrub-carr  pools  in scattered  2. It iron  Barview may  that  Frye at Barview is  supported  August typical  1922, of  the  i s further  part  of t h e y e a r  lake.  in of on  flooded depressions, snails  by  subsurface  winter  level,  f l o o d e d , except  and  p e r e n n i a l water.  Peak  lake  in  pools  Fissidens  bivalves,  impeded d r a i n a g e  c r e a t e d the  the  1922  fact  driest  coastal  case  of  and  irrigated  in  peat  but  runoff I  have  f o r enlargement  of  midwinter.  Hydrology  i n August by  rise  small  abundant  is  d e p r e s s i o n s , even  have been  hardpan  indicate  adjoining  a seasonal  the  small  community  presumably causes seen  of  below t h e  S c a t t e r e d occurrence  the a q u a t i c or h y g r o p h y t i c mosses F o n t i n a l i s branches  cm  (Christy  that  less  by  wet  that of  a  This  t h e p a s t u r e was the  subsurface  p a s t u r e " seen  ( W i l l i a m s 1933).  part  Oregon.  evidence  "more o r  caused  assumption  still  "Mediterranean"  Remains o f . t h e t r i c h o p t e r a n the h a b i t a t  1980).  was  wet  for  by  at  wet  in  summer larval least  24  2.8  V e g e t a t i o n of N o r t h American  1.  S u t t o n Lake V e g e t a t i o n  The by  S u t t o n Lake  mature  Salix  form a d e n s e (Bong.)  layer  occurs i n a shrub-carr  hookeriana Barratt  canopy  Carr.,  population  4-6  m  high  (Fig.  is  dominated  is  given  on t h e P a c i f i c  by C a r e x  ecologists  to  sunny  The  days  and  and  f u s c a Raf., Picea  in  the  obnupta  Bailey  John.  A  and D y r n e s s  and  species  of t h i s  formally  by  a t ground  level  from  plant  coverage  incident  twilight  )  and  with  Polypodium Based vegetation association  a  40 y e a r s a t dense  i s 70-98 %, e x c e p t f o r  (vid.  111  years at mm  DBH  c o r e s from 125 mm  (Salix).  epiphytic  observations  at  other  Appendix  this  appears  growth  Increment  of  glycyrrhiza on  o f 37.5  D.C.  light  conditions  a r e commonly m u l t i p l e - s t e m m e d ,  a mean age  kind  1973). i s 60-90 %,  o r decumbent.  The  scattered  complete  Pyrus  Carex  menziesi i  carr.  To d a t e , no h a b i t a t described  that  sitchensis  Pseudotsuga  and  Salix  leaning  1) gave  Pyrus  covered  varying  4).-  dominated  d e n s e l y - s h a d e d p o o l a r e a s where c o v e r a p p r o a c h e s  frequently  (  been  sunlight.  Pyrus  (Table  Coast  c o v e r of S a l i x  dappled  larger,  B.  (e.g., F r a n k l i n  Canopy on  i n Appendix has  Pyrus  and p e r i p h e r a l  L y s i c h i t u r n americanum H u l t e n & S t . list  and  Pinus c o n t o r t a Dougl.  (Mirb.) Franco are s c a t t e r e d herb  Habitats  0  %.  t h e stems 14  trees  DBH The  trees  bryophytes  are and  Eat-.  N),  t o be a s e r a i  but  Pyrus  sites /  long-lived  with Salix  similar /  community  Carex  26  Table 1 - D i a m e t e r s a n d a g e s o f P y r u s f u s c a and S a l i x h o o k e r i a n a i n L i m b e l l a h a b i t a t a t S u t t o n L a k e , Lane C o u n t y , Oregon.  PYRUS FUSCA  Diameter  (mm)  SALIX  Age ( y r )  Diameter  HOOKERIANA  (mm)  Age ( y r )  110  35  110  *  120  40  115  *  145  45  38  *  130  35  50  *  80  30  120  40  160  40  210  *  130  *  131  *  x=125  * Data  lacking;  x=37.5  increment c o r e s  X=110.6  not countable  x=40  in field.  28  dependent the  case  upon of S a l i x  Individual gaps  perennially  in  Spi raea,  hookeriana,  t r e e s appear the  canopy  the  Spi raea  common  bases of l i v i n g dead at on  vegetation.  maintained  and  Limbella  is  below, o r n e x t and  and r o t t i n g (Fig.  Rarely,  and t r e e s .  hummocks o f p e a t , branches. lowest to  substratum,  ssp.  Although  of  Limbella  as observed  Seedlings  and t h e  but  dense sods a t the living  stems,  to  0.5  mosses  Sutton  but  and l i t t e r f a l l . rhizoidal  plants are easily are  effective  f o r other  moss t a x a  sites:  elevated  t h e p l a n t s a r e wet and most o c c u r vegetation  stems  Fontinalis  elevated  fallen  m  observed  on s h r u b and t r e e  on s l i g h t l y  on  muck a n d p e a t  p l a n t s were  or hygrophytic occur  frequent,  and a t t h e edge o f  occurred  w h i c h has s t r o n g fryei  Salix  The community i s  plants climb  b a s e s o f s h r u b stems o r  of a e r i a l  tricostata,  plants,  plants  In a l l cases  level  Sods  sometimes  most  been  no P y r u s ,  stumps, and on l e a f y  5).  t o , the aquatic  Fissidens,  S a l i x , and  having  scattered  on decumbent  plants  creating  fires.  submerged t o d e p t h s o f 10 cm i n p o o l s Lake,  spray.  by gap p h a s e s u c c e s s i o n  P y r u s and S a l i x ,  shrubs  salt  years,  I observed  or e x t e n s i v e ,  t h e edge o f p o o l s of  dying  b u t dense s t a n d s  litterfall  stems  40  under a m a t u r e c a n o p y .  coastal  small  about  after  In t h e s h r u b - c a r r , forming  t o t o marine  eventually  reproduction  formerly  and p o s s i b l y i n  by young P y r u s ,  or  probably  soils,  that are f i l l e d  by t h e t a l l e r  therefore  proximity  to d i e after  overtopped Spiraea  waterlogged  a t the  In c o n t r a s t  connections  to  the  dislodged. seedbeds  (e.g.  o f C a r e x , L y c o p u s , P y r u s and S p i r a e a  for  Holcombe  vascular 1976).  a l l sprouted  from  29  sods  of  ssp. fryei  during  growth  chamber c u l t i v a t i o n  (Chapter  7).  2.  Barview  Frye's pasture  Veqetation  Barview h a b i t a t  land"  (Williams  1933).  work w i t h , a n d n o t f i n d i n g search is  for vegetation  no P y r u s /  visit the  Salix  (June  back  Sundberg,  unsuccessfully (Chapter 1980): often  carr  f o rssp.  "Because  different  Careful  comparison of  of  water,  The  (Voegtlin  litter litter  Elva  at  the  the habitat  habitat  possibly  At the  have  been  of  type  who  (S.D.  locality  i n 1956  (pers.  comm.  h a d been d e s c r i b e d a s  introduced  yielded  last  searched  I thought that  extraneous  to  There  my  there  was s u i t a b l e  this  and then d i e d litter  some  important  from b o t h l o c a l i t i e s  clues  were washed i n  funnel  f r o m a l l i s o t y p e p a c k e t s was removed  out."  i n my and  removed, and t h e wash water  The F r y e c o l l e c t i o n s  was  f o r t h e moss, and  plant  p a p e r w i t h a Buchner  thus obtained  chose  of  grew  The S u t t o n Lake c o l l e c t i o n s  the p l a n t s  I  time  and not s u i t a b l e  Limbella  1982).  information to  Lawton,  habitat etc.  wet  hookeriana e x i s t e d i n  Limbella  1980).  less  a t Sutton Lake.  of S a l i x  no  [Hawaiian]  i t might  t h r o u g h Whatman f i l t e r  but  but  i n streams with w a t e r f a l l s  concerning habitats.  flask  stand  fryei  that  the  Frye's c o l l e c t i o n s  tubs  i n Barview.  comm.  9 ) , doubted  an e n t i r e l y that  approximating that  of a house,  pers.  H a v i n g so l i t t l e  any p a s t u r e i n B a r v i e w ,  1983), a s m a l l  yard  i s known o n l y a s "more o r  and  filtered vacuum  were n o t washed, and  combined.  was t h e n  examined  30  and  s o r t e d under a d i s s e c t i n g  arthropods, all  t h e m a t e r i a l from  the dominant  profile  of  collection, "wet the  being of of  any  ssp. one  the  vegetation  the presence  can  occur  found  in at  Nevertheless,  Although  requirements, it  be  least  be  which  Cultivation  hardy  and  insolation, overstory  two  able would be  experiments  possibly  temperature  removal,  lethal  as  able and  his  evidence,  Presence  the  species  absence  indicate  very d i f f e r e n t open  site to  that  habitats,  "pasture".  once may  have  Frye's  1922  specific  (Schofield  habitat and  Crum  anthropogenic  habitat  to vascular p l a n t s  (Schofield  (Chapter  7)  showed  to s u r v i v e the humidity  l o n g as an  support  similarity  s e e d s and  t o have v e r y  survive  Frye  Trichoptera.  been c l e a r e d p r i o r  to  the  the o n l y  t h e o t h e r an  of d i s j u n c t i v e  good  this  Lake,  from  a  from  Based on  at Sutton  L i m b e l l a appears  typical  may  alteration 1969).  C).  i t i s p o s s i b l e t h a t the Barview  been a s h r u b - c a r r w h i c h had  numerous  litter  c l u e s which  dissimilar,  a c l o s e d , dense s h r u b - c a r r and  1972),  a few  of L i m b e l l a , water and  the p l a n t l i t t e r  fryei  visit.  Litter  c u l m s , Rubus p a r v i f l o r u s N u t t .  of  from  the h a b i t a t , g i v i n g  (Appendix  strikingly  Aside  Lake c o n t a i n e d  type.  though s c a n t y , y i e l d e d  h a b i t a t s are  grass  Sutton  v a s c u l a r p l a n t s of  pasture" description two  microscope.  radical  associated  a d e q u a t e water  Limbella  change i n  with  supply  to  sudden  remained.  31  D.  DISCUSSION  Comparison parameters  of  of h a b i t a t s  some s t r i k i n g  of  of L i m b e l l a  1 a s shown  for  stations Oregon,  similar,  both  summer p r e c i p i t a t i o n . Hawaiian because most  stations  show but  2 precipitation the  regions  The h i g h e r  and Oregon  seasonal  maintained.  showing total  The  as  Hawaiian  water  habitats  are  usually  f o r 10.5 months p e r y e a r and r a i n y Hawaiian  The  Oregon.  more  the  with  t e m p e r a t u r e s range  stations  b e i n g on a v e r a g e  5°C  at  the  to Limbella and  supplies  are  shrouded i n stations  are  similar seasonal  t h e Oregon of  greater  latitude.  10-25°C,  range of both l o c a l i t i e s  reduction i n  f o r 8 months p e r y e a r .  of  because  higher  from  Oregon  b o t h show  curve  pronounced  associated  seasonal  Hawaiian  stations  distributions,  slightly  Hawaiian  and Oregon  The  is  independent of  the  occurs almost d a i l y .  instead  well-watered  as  cloudy  fluctuation  a major  habitats are  long  minima  consequence  and r a i n  being  reveals  precipitation  cloud,  temperature  biotic  distribution  wet t h r o u g h o u t t h e y e a r , e s s e n t i a l l y  amount o f p r e c i p i t a t i o n ,  The  and  i n H a w a i i and Oregon  i s p r o b a b l y of s m a l l  both Hawaiian  remain  physical  of C l i m a t e s  Hawaiian  remarkably  above-mentioned  similarities.  2.9 C o m p a r i s o n The  the  Oregon  seasonal  Average from  i s identical warmer  stations  than  annual 5-20°C.  (15°C), the those  in  32  2.10 C o m p a r i s o n There latosols  is  no  Sutton  and i g n e o u s  However,  the  structurally histosols Olinda  of S o i l s  rocks  organic  similar  being c.  latosols  pH.  habitats, inundated wetting streams and  Hawaiian  of  ssp.  submerged  fluctuating  receive moisture  Limbella  both  humidity  peat  localities,  substratum.  may be c h e m i c a l l y and the  Alakai  Lake peat  significant  tricostata  Swamp  pH 4.9.  spodosols c.  similarity  may  The pH be  occurs  i n well-watered  i n p e r e n n i a l streams  or p e r i o d i c a l l y  streams. water from  In  levels,  addition  the frequent  rainfall,  a s s o c i a t e d with the c l o u d f o r e s t .  substratum water  plants are hydrated  to  by t h e  fog  drip fryei,  perennially  and a s s o c i a t e d h i g h h u m i d i t y .  i n some form  throughout  periodic  plants in intermittent  t h o u g h s u b j e c t e d t o d r y summers, i s h y d r a t e d waterlogged  the Hawaiian  Hydrology  in intermittent  high  histosols  The o n l y  tricostata  either  by  as  for  have a PH o f 5.0, t h e B a r v i e w  low  Limbella  serving  pH 5.0 and t h e S u t t o n  (Blacklock s o i l s ) .  Comparison  counterpart  t o t h e S u t t o n Lake p e a t ,  5.0  2.11  Lake  i s perennially  the year.  In  a v a i l a b l e and  33  2.12  Comparison  Although Hawaiian  of  there  and  incident walls  of  Limbella. high of is and  the  Dense  two  therefore  of  of  Hawaii  by  physiognomy  and  In stream  The  humidity  similarity,  floristic,  at  sites,  Oregon  the  vertical  dense  low-  reduce  air  favors  evidence  of  similarities  when s u c h  exists,  influencing light  habitats  more  where  that  physiognomic  open r i p a r i a n the  gullies  epiphytic vegetation,  Vegetative not  habitats  addition,  relative  of  the  dense t r e e canopy or  narrow high  of  the  i s s i m i l a r to  Hawaiian  stream g u l l i e s .  structural,  of L i m b e l l a although  level  i n Hawaii  Barview  may  SUMMARY  the  vast  geographic  physical conditions same as  North America.  climates,  saw  between  similar.  Despite  the  limited  habitats.  CONCLUSION AND  nearly  I  growths  counterpart  habitats,  correlation  the  f u r t h e r emphasizes the  humidity.  have been  Lake.  maintaining  humidity,  have no  E.  was  canopies  circulation,  floristic  vegetation,  Sutton  first-order  growing  no  H a w a i i a n h a b i t a t s of L i m b e l l a  light  of  is  Oregon  densely-forested shrub-carr  Veqetation  those at The  level  soil  pH  and  c o a s t a l Oregon a r e  sometimes p h y s i o g n o m i c a l l y Given these  sea  1200-2130  h a b i t a t s are  microclimates, and  at  d i f f e r e n c e s between  on  the  m  in  north  i n f l u e n c e d by hydrology. floristically  the  Hawaii  two are  Pacific  coast  similar  local  The  vegetation  disparate,  but  similar.'  similarities,  it  is  not  inconceivable  that  34  long-distance either growing  dispersed  locality. in  element  "quasi-temperate  latitudes"  diaspores  tricostata  " t r o p i c a l " H a w a i i , can  semi-temperate the  L_j_  Limbella  ssp.  could  tricostata,  be c o n s i d e r e d  of the Hawaiian  moss f l o r a ,  conditions  mountain  a l l u d e d t o by Moore  on  (1972).  survive in though  a temperate  or  illustrating  massifs  at  lower  35  III.  A.  ANALYSIS  INTRODUCTION If  discrete  specimens be  MORPHOLOGICAL  to  examination.  B.  identify  This between  differences proved  were  chapter  through  describes  Hawaiian  and  between  a n d Oregon,  i t should  careful  the p u t a t i v e  Oregon  morphological morphological  specimens,  measured q u a n t i t a t i v e l y , and which  how  the  characters  NEED FOR QUANTIFICATION OF DISCRETE MORPHOLOGICAL CHARACTERS (1984)  reviewed  b r y o t a x o n o m y and o u t l i n e d traditional  character  can  obvious  consistent, habitat  be  and  such  usually  differences  as  applications  Although  presence  and i l l u s t r a t e d  or  must  be  correlated  with  other  to  have  obvious  absence  without  differences  i f they a r e  in  i n s u p p l e m e n t i n g more  descriptions.  described  character  quantitative  t h e i r usefulness  morphological  morphological differences  use.  them  exist  t o be d i s t i n c t i v e .  Horton  less  differences  of s s p . t r i c o s t a t a from Hawaii  possible  differences  morphological  of  a  ambiguity,  quantifiable, character  meaningful  or  taxonomic  36  C.  MORPHOLOGICAL SIMILARITY OF HAWAIIAN AND NORTH AMERI CAN  LIMBELLA When L.  designating  tricostata,  morphological and  leaf  features,  times and  reveals  plants  trailing.  synonym no  to  significant,  it  was  and a n a l y z e  Leaf  (Vitt areas  necessary them  other  be i d e n t i c a l ,  both  by  Hawaii Christy  microscopic  than  size  of  s h o o t s and b r a n c h e s up t o 3-5  mosses  from  from  including  difference  of  significant  reiterated  length,  given and  are  To d e t e r m i n e o b j e c t i v e l y i f  characters  a  that  of specimens,  from O r e g o n .  appear  plants  was  develop  i n most a q u a t i c  addition,  as  indicated  opinion  little  than those  serration  inherent  This  comparison  Hawaiian  longer  (1971)  fryei  d i f f e r e n c e s e x i s t e d between s p e c i m e n s  Cursory  plants:  D.  Lawton  North America.  (1980).  Limbella  to  the polymorphism  Glime  either  these  areolation  1984).  In  dendroid  or  differences  measure  the  were  different  statistically.  SIGNIFICANCE OF DIFFERENT BRANCHING PATTERNS IN LIMBELLA  that  Close  i n s p e c t i o n o f H a w a i i a n a n d Oregon  their  branching  difficult pattern  to  patterns  describe  terminology.  fairly  regular  plants  a pinnate  differed.  satisfactorily Hawaiian  intervals or frondose  up  plants  specimens i n d i c a t e d  This using  difference  standard  exhibit  branching  branching  symmetrically  around  at  t o 1 cm d i s t a n t , w h i c h g i v e s t h e  habit, especially  i n aquatic  forms.  However, t h e b r a n c h e s a r e n e i t h e r d i s t i c h o u s n o r c o m p l a n a t e arise  was  the  stem.  In c o n t r a s t ,  but  i n Oregon  37  plants  the  plants  a  penicillate,  therefore  3.1  branches a r i s e in c l o s e l y - s p a c e d  nearly  Control  of  always  both  development dominance by  be  of  branching  from l a t e r a l  only  by  and  branch  Colbert (Bopp  bud  pattern  buds a p p e a r s  1977;  decapitation  determined  same  habitats  Quantification location  and  Each leaf  of  buds. t o be  1979;  1984)  branch  merophyte  to q u a n t i f y  ("leaf  b r a n c h bud  located  that  arise  strictly authors.  segment cells  directly 1931;  subsequently  leafy  and  below  from  shoot  growing  in  patterns. rely  on  c a l c u l a t i n g branch  as  i s the  product  apical  during  shoot  1966;  at  of  leaf  buds  bud bud  elongation.  1977).  of many divisions  As  such,  and  (von  Branches  therefore  erroneously  successive  one  intervals a  insertion  are  composed  (1)  cortical  regular  Kawai  stated  thus of  (3) the  these  is  m e r o p h y t e s and cell  taxa  composed  Schuster  speaking, " a x i l l a r y , A  be  l o c a t i o n must  same b r a n c h i n g  by  can  segment" a u c t . ) c o n s i s t s o f :  stem  parenchymatous  Lorch  stimulated  position  t h e r e f o r e , must  this  apical  number.  underlying  1911;  pattern,  branch  by  and  P r e s u m a b l y , bud  would e x h i b i t t h e  merophyte  (2) a s u b t e n d i n g  Schoenau  are  must  Although  inhibited  Bopp  1984),  location.  branching  I chose  f r e q u e n c y per  plants  differences  g e n e t i c a l l y c o n t r o l l e d , otherwise d i f f e r e n t  the  The  the  dendroid.  branches  (Hebant  shoot  appearance.  giving  Branching  Quantification include  brush-like  groups,  by  hot, some  adjacent by  the  merophyte  38  features are  therefore  also the  are d i f f e r e n t i a t e d  arise branch  develop  primary developmental  in c l u s t e r s bud  directly  METHODS  3.2  Quantification  a bud  m e r o p h y t e number merophyte, branch  resuming the  the count  percent  using  pattern  rhizoids  therefore are  by:  (1)  from  shoot  encountered  the count  spiral  not  elongation.  that  counting base  each  to  shoot  (2) r e c o r d i n g  at zero  pattern  (4) r e c o r d i n g  r e a c h e d and  fryei  until  with  the  t h e next  merophyte  that next  bud  number  the c o u n t i n g t e r m i n a t e d . tricostata  were c o u n t e d .  at the  each  merophyte  frequencies  m e r o p h y t e numbers. to determine  discriminant  The  and  The  a t e a c h m e r o p h y t e number  by computer  stepwise  as  or (5)  at z e r o w i t h the next merophyte, e t c . , u n t i l  and  corresponding analyzed  spiral  the  occurrence  calculated,  and  rhizoids  same s i t e  present,  d u r i n g shoot  determined  m e r o p h y t e s ) of s s p .  occurring  cells  cell  o r b r a n c h was  m e r o p h y t e s ) of s s p . branches  the  encountered  (1134  cortical  (3) r e s u m i n g  s h o o t apex was  shoots  were  following  was  Although  and  of B r a n c h i n g P a t t e r n s  following  until  tissues.  development  each merophyte, a t the  the a p i c a l  frequencies  merophyte, apex,  from  by  E.  Bud  from  shoot  or s u r r o u n d i n g i t i f both a r e  secondarily  initiated  a t the time of  similarity  17 s h o o t s  (689  numbers o f buds o r were  number  plotted data  Twenty  totaled,  (=frequency) against  s e t (Appendix or  the  the E)  was  dissimilarity,  a n a l y s i s and M a h a l a n o b i s  distance  39  (D )  MIDAS p r o g r a m s .  2  values three  Discriminant  i n a x e s of d e c r e a s i n g a x e s c o n t a i n most of  reducing  and  containing  the  the  variance  it.  greatest variance  between t h e  represents  a  differences  variance.  simplifying  differences  between  overlap  between p a i r s of v a l u e s  between  specimens are maximized.  detect  (Sneath  3.3  Phenetic  Distance  ranks  and  is  1973;  computer,  using  and  stepwise  MIDAS p r o g r a m s .  because  they  decribed  2  measures  means showing  as  method  a  the  function  of  the d i f f e r e n c e s  i s most  Everitt  the  the because  taxa,  salient  should  sporophytes ssp.  stems and  few  useful  at to  1982).  tricostata f o r 28  (Appendix  28  G)  any of  ssp. up  characters  morphological  to d e t e c t  tricostata branch  ssp.  scores  The  g a m e t o p h y t e s i n enough d e t a i l  thick  that  d i s c r i m i n a n t a n a l y s i s and  (D )  of  distance  value  were s c o r e d  the  distance  shoots  axes  Mahalanobis  i n so d o i n g  The  the  greatest  measured and  of  thereby  r e v e a l the  distance  Dunn and  i n c l u d i n g types,  (Appendix D)  new  sampled.  specimens of  characters  scored  plots  or  Analysis  fryei,  between  set,  data  two  where d i f f e r e n c e s a r e more d i f f i c u l t  Sokal  Twelve herbarium ssp.  i n the  p a i r s of c h a r a c t e r  variance.  taxonomic  first  should  phenetic  greatest  lower  thus  character  U s u a l l y the  Scatter  specimens  generalized  analysis ordinates  fryei to c.  leaves, only  12  of  morphological analyzed  by  Mahalanobis were  chosen  f e a t u r e s of  significant  exist.  and  the  differences  O n l y g a m e t o p h y t e s were are  unknown.  6 cm  l o n g can  shoots  longer  Because have  very  than  8  cm  40  were s c o r e d . below  tips  apices.  of  main  Leaf  measured for  L e a v e s and  stem s e c t i o n s were m e a s u r e d  shoots  apical  or  angle  and  character  being  represented  means of  RESULTS  3.4  Q u a n t i f i c a t i o n of B r a n c h i n g counts  frequencies frequent  plants lesser buds  by  6).  ninth  up  were  the  two  (1951). 5-10  In  t o 20  most  peaks e v e r y  Measured  were values  measurements of  and  each  frequent  (30.0%) e v e r y  fourth, sixth no  and  cases,  individual  but  p o p u l a t i o n s as a whole a p p e a r e d  the  H a w a i i a n and  in  descending  most  lesser  peaks  every  A  first  ninth  more than  few  Oregon  merophyte,  merophyte.  plants exhibited atypical t o be  buds were in  13 m e r o p h y t e s . bud  uniform  with  A  few  In a  very  frequencies,  when  all  bud  averaged.  S t e p w i s e d i s c r i m i n a n t a n a l y s i s of bud measured  were  In c o n t r a s t , buds  few  f r e q u e n c i e s were  with  bud  buds  e l e v e n t h merophyte.  merophytes.  by  plants,  (26.4%),  separated  0.0001), 8  angle  revealed differences in  Hawaiian  were  plants  intact  Patterns taxa  f o u r t h merophyte  sixth,  separated  of  (Fig.  every  tooth  cm  scored.  F.  Bud  branches with  marginal  f o l l o w i n g t h e method of Wagner  each specimen  first,  largest  1.0-1.5  ( A p p e n d i x F)  indicated  Oregon p l a n t s were order  of  and  37  t h a t d i f f e r e n c e s between  statistically  significance,  (p < 0.001), 3 (p < 0.05)  f r e q u e n c i e s of t h e  at  significant,  merophytes  5 (p < 0.05).  1 (p < Based  on  41  these  4  characters,  tr icostata highly  and  ssp.  significant  the  Mahalanobis  fryei  was  (p < 0.0001).  distance  11.807 and was  between  ssp.  statistically  42  Figure  6 - B r a n c h bud f r e q u e n c i e s i n s s p . tricostata ssp. f r y e i , p e r l e a f segment number.  L e a f Segment Number  and  43  3 .5 Phenetic  Analysis  Stepwise  discriminant  morphological characters between were, (p  characters  exhibited  the  0.0001),  0.01), c o s t a cell  characters, and  ssp.  significant The  23  correlation be  of  was  indicated  The leaf  (p  0.05).  five  40.086  and Based  between  and  was  (p <  basal  leaf  on  ssp.  statistically  characters  significant.  matrix  identified  However,  (Appendix  correlated  were not  more  I)  examination  showed b a s a l  with  correlated  i t follows  that  of d i s c r i m i n a n t v a l u e lengths  addition  basal  leaf  leaf  A  the clear  7  five  scatter  distinction  characters the  also.  other  listed  plot  between t h e  characters distinction  being  the  lengths  length:width only  one  statistically  Longest  and  shortest basal  distinctive  may leaf  characters  in  above.  using taxa  all (Fig.  identified between  as  of  cell  cell  being  correlated characters  were t h e r e f o r e c o n s i d e r e d  t o the  5  highly  as  or  cell  these  tricostata  of  be  angle  length  Because d i s c r i m i n a n t a n a l y s i s tends to s e l e c t  significant,  5  characters  ratios. two  that  insertion  (p < 0.05)  <  28  differences  (p < 0.005), b r a n c h  insertion  the  0.0001).  remaining  highly  for  significant  Mahalanobis d i s t a n c e  (p <  H)  significance,  diameter at  data  Oregon p l a n t s .  ratio  fryei  statistically  to  stem  thickness  the  (Appendix  and  order  lengthrwidth  of  statistically  Hawaiian  in descending  <  analysis  the  by taxa  28  characters  7).  A  revealed  scatter  plot  little using  d i s c r i m i n a n t a n a l y s i s showed (Fig.  8).  44  Figure 7 - S c a t t e r p l o t o f measurements o f 28 morphological characters. Triangle = ssp. t r i c o s t a t a, c i r c l e = ssp. f r y e i .  <SCATTER V A R = 2 9 . 3 0 > SCATTER PLOTN= 24 OUT V29 614.98 •»  OF  24  29.V29  VS.  closed  30.V30  A  A  A A  A  •  38.535 30.474  54.657 4G.59S  70.779 S2.718  V30 78.840  45  Figure 8 - S c a t t e r p l o t o f measurements o f 7 m o r p h o l o g i c a l c h a r a c t e r s , a f t e r s e l e c t i o n by d i s c r i m i n a n t a n a l y s i s . Triangle = ssp. t r i c o s t a t a , closed c i r c l e = ssp. fryei.  ^SCATTER VAR=B,9> S C A T T E R PLOT N= 24 OUT V8 601.93  OF  24  8.V8  VS.  9.V9  553.31  456.07  -36.463  -24.467 -30.465  -12.472 -18.470  -.47647 -6.4742  11.519 5.5212  V9 17.517  46  G.  DISCUSSION The  bud  and  statistically  f r e q u e n c i e s of Hawaiian  bud in  consistent  counts,  confirmed  the  being  the  angle,  i s probably g e n e t i c a l l y  key  presented  c h a r a c t e r s were c o n s i d e r e d superfluous  for  insertion  the  plotted  experiments  their  SUMMARY AND  showed t h e s e  by  importance  than  10).  as  by  they  insertion Cultivation  c h a r a c t e r s t o be  c o n d i t i o n s and  were  9)  leaf  or  taxa.  length  (Fig.  (Fig.  two  g e n e t i c than  of b r a n c h  ssp.  fryei  which  is reflected  bud  stable  presumably  environmental  more  factors,  taxonomic c h a r a c t e r s .  frequency  revealed significant  tricostata  whereas buds o f  plotted  diameter  measure  cell  in  remaining  between t h e  leaf  effectively  stem  to  leaf  used  The  as  CONCLUSIONS  Measurement  ssp.  when  of e n v i r o n m e n t a l  controlled  corroborating  H.  7)  (Chapter  discrimination  more  against  under a v a r i e t y strongly  taxa  differences  were  8.  Chapter  time-consuming  and  mean  length,  length)  in  fixed.  (branch  cell  longest basal  measurable,  distinguished angle  convenient  a n g l e and  conveniently  3  in  too  on  by d i s c r i m i n a n t a n a l y s i s  significant,  longest basal leaf  dichotomous  Leaf  Oregon L i m b e l l a , b a s e d  7 characters identified  statistically  insertion  differences  t h e e x i s t e n c e of q u a n t i f i a b l e  branching p a t t e r n which Of  and  significant  by  ssp.  differences  the d i s t r i b u t i o n  most  often  fryei  i n ssp.  occur  of  i n bud  fourth every  and  distribution  branches.  every'  most o f t e n o c c u r  tricostata  Buds  of  merophyte,  first  47  Figure 9 - Leaf i n s e r t i o n angles p l o t t e d against longest basal leaf c e l l lengths. T r i a n g l e = ssp. tricostata, closed c i r c l e = ssp. fryei.  <SCATTER VAR = 1 , 2 > S C A T T E R PLOT N= 24 OUT VI 60.000  t*  OF  •  24  1.V1  VS.  • •  2.V2 •  50.000  00  c 03  a o  4G.6S7  A  A  u 0)  0) c  4-1  n)  A  4o. ooo  AA 2S.OOO  36.556 30.778  48.111 42.333  A  A 59.667  53.889  Longest basal leaf cell (pin)  71.222 65.444.  V2 77.000  48  Figure diameters.  10 - L e a f Triangle  <SCATTER> S C A T T E R PLOT N= V 1 SO.OOO  24 OUT  OF  24  i n s e r t i o n a n g l e s p l o t t e d a g a i n s t stem = ssp. t r i c o s t a t a , closed c i r c l e = ssp. fryei.  t.VI  VS.  2.V2  CD iH 60 (3  46.667  c o u, ts M-l CO  AA  4 0 . OOO  t-1  AA 165.00  261.67 213.33  358.33 310.00  455.00 406.67  Stem diameter (pm)  551.67 503.33  V2 600.00  49  merophyte. of  ssp.  I f and  tricostata  s p a c e d , and  those  in  branching  bud  frequencies  suggest  that  elongation  are  two  of  therefore  ssp.-  bud  fryei  clearly  by  more c r o w d e d .  e x h i b i t e d by  i n ssp.  formation  necessity  by  branches  more  widely  Such d i f f e r e n c e s  many s p e c i m e n s .  Average  tr icostata  and  ssp.  fryei  the  cell  during  shoot  apical  is genetically fixed. a n a l y s i s of  gametophytic  cell  length,  and  ssp.  congruent,  insertion  fryei.  characters,  ssp.  features.  angle  and  to  segregate  The  length  support  taxa.  in  and  ssp.  indicated basal  leaf  tricostata  measured were o v e r l a p p i n g  distinct  cell  and the  insertion  tricostata  basal  experiments  leaf  measurements  different  Other c h a r a c t e r s  showing t h a t  cultivation  phenetic  were s i g n i f i c a n t l y  morphological  characters  are  observed  Multivariate that  when s u c h buds p r o d u c e b r a n c h e s , the  the  was  or  larger  in  most  differences  in  leaf  confirmed  by  were use  of  these  two  50  IV.  A.  KARYOTYPE ANALYSIS  INTRODUCTION Among m o r p h o l o g i c a l l y  often  the  same  distinguishing confirming  and the  similar  therefore taxa,  similarity  taxa, are  although  based  on  analysis  This  in bryophytes  analysis  in  tricostata  ssp.  4.1  Utility  are  often  be  morphology.  In  the  same  (Koponen  number, 1978).  information  such and  an  useful these  critical  the  results  ssp.  in in  cases,  taxonomic  a p p l i c a t i o n s of  karyotype  of  such  tricostata  e x c e l l e n t r e v i e w of  an  and  L.  while  others  present  therefore being  "the  taxa  providing of  limited  analyses  the  additional  usefulness.  f o r more d e t a i l e d c y t o l o g i c a l  'count-  cluttering  little  have  variation  all-too-often  numbers  an  themselves  exhibit intraspecific  chromosome  reports  c o n t r i b u t i o n , by  b e c a u s e many d i v e r s e  (1981) d e c r i e d of  the  Chromosome numbers, t h o u g h sometimes  usefulness  Buck  reports  i s a need  provide  use  Analysis  or p h y l o g e n e t i c  limited  literature,"  there  Karyotype  taxonomic  and-run'  reports  tricostata  (1984) p r o v i d e d  of  taxonomic  are  fryei.  of moss c y t o l o g y .  important  they  reviews  and  Limbella  of  Ramsay state  chapter  little may  karyotype a n a l y s i s i s e s s e n t i a l to information.  of  chromosome numbers  Clearly which  can  51  provide why  more i n f o r m a t i o n  karyotype  received  taxa and  with  as  Ramsay  analysis  morphology  i n s u c h a way  karyotypes  possible. ranging  interpretive  new  (Koponen 1982,  is  an  attempt  as  to  make  Newton  1979;  dependent  on  Ramsay the  centromere. 1963;  photographs  activity  1982).  to describe  of  lengths  have  l e n g t h of complement, e x p r e s s e d between c l o s e l y - r e l a t e d  taxa.  Inoue  an  other  array  of  squashes, and  lengths,  to  chromosome shape, l e n g t h ,  and  by  (Inoue  on  length as  karyotype  ideograms  %)  their  and of  and  data,  relied  (relative  reviews,  with  chromosome  Derivation  centromere  1981)  has  1979;  included  correct identification  Lacking  Mendelak  chromosome  Newton  comparisons  have  f o r m u l a s which d e s c r i b e staining  1931),  closely-related  d r a w i n g s made from s u c h p h o t o g r a p h s ,  heterochromatic  is  1984).  Analyses  from  This  bryological  1978;  a l i g n m e n t s s h o w i n g chromosomes a r r a n g e d karyotype  (Lewitsky  a t t e n t i o n in recent  karyotypes  1980;  Karyotype  techniques  chromosome numbers.  a means of d i s c r i m i n a t i n g between  similar  Yamashita  just  a n a l y s i s , though h a r d l y  considerable  especially  than  Iwatsuki  such f o r m u l a s placement  some  workers  absolute = absolute to  1976;  and length  detect  of  is the  (Iverson relative : total  differences  52  4.2 C y t o l o g i c a l R e p o r t s To  forLimbella  my knowledge no t a x a  cytologically. information  Ramsay  of L i m b e l l a  (1984) c i t e d  have been  the paucity  investigated  of  cytological  f o r b o t h G o n d w a n a l a n d i c and H a w a i i a n moss  Under Sc iaromium M i t t . , o n l y been  Taxa  investigated  S^  lescur i i  cytologically,  (Sull.)  Crum  and  floras.  Broth.  Anderson  (1981) counts  reporting  a chromosome number o f n=11, b a s e d on  meiotic  (Fritsch  1982).  a  should  Sc iaromium l e s c u r i i  be a s s i g n e d  is  MATERIALS AND METHODS  4.3  Chromosome Squashes and D o c u m e n t a t i o n Squash p r e p a r a t i o n s different  not  plants,  of L ^  tricostata  using  were  gametophytic  shoot  Kauai p o p u l a t i o n s ,  Attempts  t o o b t a i n chromosome f i g u r e s from Maui  which had (Chapter  been  M a t e r i a l was o b t a i n e d maintained  study.  in  a  population;  from  d i d not  tricostata stain  growth  I t was n e c e s s a r y are  were f i x e d  properly  fryei  sporophytes  unknown,  easier  t o s e e and document b e c a u s e  c.  a p i c e s - (4 F i g . 11).  s p o r e l i n g s were  chamber  4-5  plants months  a n d a few immature  in  and c o u l d  t o use somatic  from  field-collected  7 ) . A l t h o u g h numerous s h o o t a p i c e s  sporophytes of ssp. material  1 Oregon  made  different  unsuccessful.  L i m b e l l a and  t o P l a t y l o m e l l a Andrews.  B.  150  has  the  field,  n o t be u s e d  tissues  and  mitotic  they  are less  because  this i n the ssp.  chromosomes a r e "sticky"  than  53  meiotic  chromosomes  Steel's modifications: material  in  dichlorobenzene a petri  (Lowry  1948;  Newton  (1978)  procedures  (1)  shoot  the  1-cm  growth  tips  p l a t e , and  stored  at  were were  chamber,  c r y s t a l s between 18°C  1979).  sheets  followed collected  placed of m o i s t  for 4 hr.  The  with  with from  living  a  filter  some  few  p-  paper  in  material  was  54  Figure  11 -  S o u r c e s of l i v i n g p l a n t s of s s p . used f o r l a b o r a t o r y e x p e r i m e n t s .  Na P a l i - K o n a Haena,  Hawaii  7.5'  Forest Reserve,  Kauai.  tricostata  Modified  t o p o g r a p h i c map.  A N 250  m  from  USGS  55  then  (2) f i x e d  glacial  acetic  distilled  hr  acid)  water  full-strength times  i n Carnoy  fixative  water  and  3 times i n d i s t i l l e d  (9) r i n s e d squashed  Slides  3 times  i n Hoyer's  were  (Beeks  by  pencil  at  water  5°C  to  Cover  Dried  P-5146,  (5)  in used  rinsed  solution  water  (after  and  3  with  slips  dissected  and Kuta  specimens  1983).  to squashing  the  end  of  and s p r e a d t h e c e l l s .  placed  were  directly  immediately with  from  the  chromosome f i g u r e s  an  Slides  and squashes  eraser  were r i n g e d  showing  aceto-  f o r 2 hr a t  (10)  Przywara  vigorously  separate  in  (Whitman's s t a i n )  in distilled  voucher  and s l i d e s  times  (8) s t a i n e d  pressure with a pencil  the t i s s u e s .  collections,  at  f a c e down on a b s o r b e n t p a p e r  applying  polish.  (Sigma  h e a t e d on a h o t m i c r o s c o p e lamp p r i o r  were t h e n p l a c e d  over  hydrate  1955) and t a p p e d r a t h e r  unsharpened  made  f o r 2 hr  3  1 part  (6) h y d r o l y z e d i n 5N HCL a t 18°C f o r  iron-haematoxylin-chloral 18°C  rinsed  immersed i n p e c t i n a s e  from t h e b o t t l e )  (7) r i n s e d  100% e t h a n o l :  f o r 3-8 h r a t 5°C (3)  (4)  in d i s t i l l e d  (3 p a r t s  nail  cultivated  are deposited  UBC. Photographs  under  o i l immersion  Sketches  of  the  photographed. tracing of  metaphase  figures  Interpretive  chromosomes  according  were  drawings  to  generated  chromosome  squashes  a t 1000X w i t h a L e i t z - compound  the photographs  interpretive  were  of  their  were  made  drawings  and i n k i n g made  by  at  the  with india arranging  using  the  lengths. MIDAS  cut-outs  were  Alignments from  the  left  to  plots  and graphs  program.  Absolute  Scatter  scatter  they  were made by  ink.  i n d e s c e n d i n g o r d e r from absolute  microscope.  time  of the f i g u r e s  were made  right,  1  56  lengths  (Appendix  discriminant Steel  J)  a n a l y s i s and  separation. HCL  were  Limbella, Hoyer's  and  an  d i d not  The  and  squashes, widely  in  my  determined most  the by  useful  One  simultaneously  only  nature  by  of  trial  and  produced,  (e.g.,  difficulty for storage  Smith  they  encountered of  fixed  fading.  schedule,  of  researchers,  material  at  As  a  are  not  as  to  be.  appear  material  in  part is also  hand.  The  taxon  result,  involved  vary  in  t i s s u e s of a g i v e n  first  Eight  chromosome  variation,  1978)  any  hydrate  s i g n of  general  error.  (a)  without  bryophyte  This  the p l a n t  of  Study M a t e r i a l  of  preferences  t o a l l mosses as  methodologies  (b)  slide  showed no  same  procedures.  of m e t h o d o l o g y  applicable  the  methods f o r s q u a s h i n g  identified  reviews  preparation  personal  the  Use  a dark p u r p l i s h - b l a c k .  preparations  following  certain  reflecting  staining.  a semi-permanent  nuclei  with  t i s s u e s of  aceto-iron-haematoxylin-chloral  the  though  chromosome  separating  medium  with  temperature  with  V a r i a t i o n s i n M e t h o d o l o g y D i c t a t e d by for  stepwise  treatment  improved  room  interfere  squashing  and  staining,  Methods  be  to  polish,  chromosomes  months a f t e r  4.4  prefixation  h y d r o l y s i s at  using  MIDAS p r o g r a m s .  2  f o r s o f t e n i n g and  a  nail  treatment.  stained  (D )  mitosis  appear  as  computer  a d d i t i o n a l t i s s u e - s o f t e n i n g a g e n t and  once r i n g e d w i t h further  and  essential  solution  provided  that the  retarded  Pectinase  by  Mahalanobis  (1978) r e p o r t e d  p-dichlorobenzene  5N  were a n a l y z e d  can  recent  universally  conflicting  either  fixative  57  or e t h a n o l .  Because Carnoy  refrigerated,  some  material  ethanol  field that  conditions.  researchers  1953;  should  H.P.  observed  T h i s was  negative  done, o n l y  storage  is  of  fixed  unavailable  under  to f i n d  poorly.  effect  comm. on  4-5  months  It i s probable  1984), a l t h o u g h  staining  unless  after  that  the  (Newcomer  McAdam  long  later  (1982)  storage  in  ethanol. There  softening (e.g.,  are and  numerous  cell  Bowers  conflicting  separation prior  1969;  Smith  recommended h y d r o l y z i n g f i x e d minutes  at  used heat and  recommend  n e v e r have been t r a n s f e r r e d t o e t h a n o l  Ramsay, p e r s .  no  deteriorates rapidly  i f refrigeration  the m a t e r i a l s t a i n e d very  material  70%  i n 70%  fixative  18°C.  as  the  Iwatsuki  (1982) f o u n d Limbella little  that  with  using  worked e x t r e m e l y flattening  1978;  Przywara  material  other  used heat  fixative  1N HCL on  to squashing.  in  at  and  alone  60°C  was  tissues.  pectinase  and  w e l l on  were a c h i e v e d .  used  to macerate t i s s u e s  that  different  taxa  (e.g.  2N  h y d r o l y s i s with  need d i f f e r e n t  1983),  6-30  Lowry  1948)  and  HCL  at  McAdam  18°C  Steel's 5N  Inoue  Hydrolyzing  HCL  t o 6N  had  (1978) at  spreading  emphasizing  treatments.  1983)  for  acid,  satisfactory  c o n c e n t r a t i o n s up  (Fritsch  workers  (1969) and  However,  L i m b e l l a and HCL  HCL  acetic  and  tissue  Kuta  sufficient.  (20-30 min)  softening  and  Ramsay  45%  Many  1-2N  researchers  s o f t e n i n g agent, while  (1976)  effect  methods,  Still  methods i n v o l v i n g  18°C, and  have  been  the  fact  58  C.  RESULTS  4.5  Karyotype The  fryei  Analysis  chromosome  i s n =11 .  somatic  number  McAdam  cells  (1982)  and  vary  from  can  preparation.  Cell  most  common  i n embryonic  shoot  tips.  from  one  counting  divisions  another and  tricostata,  in  a  I observed 11  of  (longest  3.5  the  constrictions  shortest  cell  failing  were  r e g i o n of  to separate acccurate  among a p p r o x i m a t e l y 25 c e l l s 12  &  150  (14 o f  ssp.  which  showed  13)  chromosomes  0.7  squash  abundant,  precluding  ; Figs.  The  chromosome  and  may  differential therefore  Mm,  thus  ssp.  dividing  given  the a p i c a l  photographed  plane.  how  any  never  "sticky,"  and  fryei  noted  and  were  extremely  urn), making a c c u r a t e  counts  i n many p r e p a r a t i o n s .  Although  features  l e a v e s from  and  tricostata  in  Consequently,  ssp.  good s e p a r a t i o n i n one  difficult  0-40  in Limbella,  squash  measuring.  ssp.  others  Many chromosomes were  preparations,  small  of both  bends  not  be  chromosomes,  which  as  to  placement  of  at  indentify them  but  (McAdam  these  regions 1982).  centromeres in  exhibit  centromeres,  all  staining  of  Limbella  resemble  centromeres  heterochromatic  reluctant  figures  on  of I  was  these  many c a s e s would  have  amounted t o g u e s s w o r k . Non-placement karyotype and  ssp.  of centromeres  formulas. fryei  precluded  Therefore, karyotypes  were compared on  identification of s s p .  t h e b a s i s of a b s o l u t e  of  tricostata and  59  60  Figure  13  - Chromosome f i g u r e s 5505 ) .  of  ssp.  fryei  ( Chri  61  relative show  lengths  of i n d i v i d u a l  chromosomes  right,  according  Absolute were  almost  congruent data  o f a l l 25 m i t o t i c to t h e i r  completely  lengths  taxa  and  of  were n o t i c e a b l e .  fryei  than  shorter  in  in ssp.  fryei  than  range of s i z e  15  left  to  1982).  two  the  ssp.  lengths). in  the  16);  larger  and  taxa  few nontr icostata  However, when t h e  the  data  set  were  18), d i f f e r e n c e s between t h e two 1-3  and  in  shows t h a t a v e r a g e chromosome broader  the  Chromosomes  tricostata,  14  Ramsay  l e n g t h s of  chromosome  (Fig.  (vid.  (Fig.  chromosome  each  graphed  lengths  chromosome  reflect  Figs.  p l a t e s a l i g n e d from  congruent  s e t (22 a d d i t i o n a l  averaged  absolute  and r e l a t i v e  records merely  relative  chromosomes.  were  longer  chromosomes  tricostata.  4-11  Also,  l e n g t h s of s s p .  c l a s s e s than  in  d i d those  fryei of  ssp.  were a l l Fig.  17  exhibited a tr icostata.  Stepwise d i s c r i m i n a n t a n a l y s i s of the a b s o l u t e  lengths  275 chromosomes m e a s u r e d ( A p p e n d i x  that differences  between  the  significant, 8  Hawaiian  and  i n descending  Oregon order  (p < 0.005) and 1 (p < 0.05).  the  Mahalanobis  distance  statistically  significant  were  correlated  highly  correlated  with  Despite each the  chromosome total  (17.5  am).  plants  statistically  of s i g n i f i c a n c e ,  a t chromosomes  B a s e d on t h e s e  between  with  LT  and  two  LF  characters,  was  Chromosomes  3.2912 and  6,7,  chromosome 8, and chromosome  and  9  2 was  1 (Appendix L ) .  d i f f e r e n c e s , the average a b s o l u t e i n e a c h complement was  average absolute  the  were  (p < 0.005).  chromosome  these  K) i n d i c a t e d  of  identical  length  of  (1.6 urn), and  complement l e n g t h was a l s o  identical  F i g u r e 14 - S s p . t r i c o s t a t a chromosome a l i g n m e n t s . 1-10 K a u a i , C h r i s t y 5368. 11-14 - K a u a i , C h r i s t y 5337.  O l !  tltlt"  ))))>>ttm  9  liliutii'ft*  >mtttu»t 1  )t>»Slftl»*»  Ulflftl AM*  )))UtlllM  63  F i g u r e 15 - S s p . f r y e i chromosome a l i g n m e n t s . 1-6 O r e g o n , C h r i s t y 5499. 7-11 - O r e g o n , C h r i s t y 5505.  1  2  ) ) ) ) ! • ! « • • •  7  )jj»|*»ftttl  9  J  )l»)ftllftt*  10 5  )»l>1ltlt#  . In  I)  I I}A | 4 « |  J>>l»t>W>t  11 UttitlM**  64  F i g u r e 16 - A b s o l u t e chromosome l e n g t h s p l o t t e d a g a i n s t r e l a t i v e l e n g t h s , a l l chromosomes m e a s u r e d . Square measurements  =  measurements i d e n t i c a l of  additional  to both  ssp.  taxa.  tricostata  Triangle = chromosomes  measured.  <SCATTER V A R - 1 , 2 > S C A T T E R PLOT N« 121 OUT »L 3.0000  OF  275  1.AL  VS.  2.RL  • I I I I •  •U 60  c  A  CD  A  AH A| I I |  I  A| I II  A 11  1 .9778  II  III  I A  I  ||  o CO <!  1.4667  • I  I  I  I • I A •  I  I  I  12.377 11.009  Relative  length  15.112 13.744  RL 16.480  65  F i g u r e 17 - A v e r a g e a b s o l u t e chromosome l e n g t h s p l o t t e d against average r e l a t i v e lengths. Triangle = ssp. tricostata. Closed c i r c l e = ssp. fryei.  <SCATTER V A R - 1 . 2 > S C A T T E R PLOT N » 11 OUT AL 2.80O0 +  OF  11  1.AL  VS.  2.RLORA  2.5889  1  c  cm  <u  4-1 3 rH O CO  &  (0  <u  •A  00 CO  A A  >  A  •  • A*  •  A  .90000 5.3300  7.6989 6.5144  8.8833  '0.06B  12.437 11-252  Average r e l a t i v e  14.806 13.621  length  RL0RA 15.990  F i g u r e 18 - G r a p h o f a v e r a g e r e l a t i v e l e n g t h s o f chromosome complements o f s s p . t r i c o s t a t a and s s p . f r y e i .  Chromosome Number  67  D.  DISCUSSION  4.6  Significance The  in  following  the  the  genus  two  Limbella.  use  in  first  Given  the  this  chromosome  mosses  (c.  doubling  36%),  numbers  Based  number  the  karyotype"  exhibited  postulated  that  lesser  numbers  "possibly remains  t o be  seen  Significance Although  karyotype  data  Inoue and 1981;  (Smith  from  t h e sample  formulas  size  makes  is  it  several  Ramsay  1982,  1976; 1984)  Inoue  1980; in  after Ramsay  the  McAdam  base number  n=11 (1982)  is  x=11,  fusions  and  fragments."  It  i s tenable.  small,  and to  the  S e v e r a l workers Yamashita  have e m p h a s i z e d  absence  compare  c o n c l u s i o n s can  and  most  Data  difficult  on chromosome l e n g t h . Iwatsuki  Crosby  s m a l l chromosomes and  of Chromosome L e n g t h  that  the  aneuploidy  chromosome  t h i s concept  little  showing is  similarity  the primary  arisen of  by  1978;  of  of  for pleurocarpous  Diplolepideae,  group  whether  complements o b j e c t i v e l y , the  x=6  n=11  reported  t o have a r i s e n  the  in that  elimination  far  are  o t h e r than  number  "remarkable  by  having  The  so  suggested  on  themselves  investigation,  o f t h e base number  1984).  chromosome number r e p o r t s  the m o r p h o l o g i c a l s i m i l a r i t y  t a x a have t h e same number.  common  4.7  Chromosome Number  r e p r e s e n t the  karyotypes,  taxonomic both  of N=11  be  the  of two  drawn  from  (Iverson  1963;  1980;  the u t i l i t y  Mendelak  of a b s o l u t e  68  and  relative  Absolute  chromosome  treatment (Newton these  chromosome  before  1979;  lengths  lengths  fixation,  Ramsay  variations  taxonomic  reportedly  s t a g e of  1984).  and  as  Use  of  provides  can  division  be and  relative  characters. affected squash  lengths  a discriminative  by  methods  minimizes  morphological  parameter.  4.8  Similarities Figures  the is  14  Between t h e and  chromosome  and  the  total  suggests  this  be  with  4.9  individual observed  18  that  the  numbers  Between t h e revealed  differences of  in  of  fact  This  and  are  chromosome  Figure  16.  Such  closely related,  that  exhibit  in  similarity  average  length, taxa  n=11  but  most  Diplolepideae  close  morphological  Pellia these  i n the  relative  lengths  s c a t t e r d i a g r a m s , Mendelak  megaspora  were s u f f i c i e n t different  Karyotypes  length  characters, on  the  Using  that  part  two  similarities  karyotypes.  values  differences  concluded  in  superficial  1982).  chromosomes.  chromosomes  two  identical  (McAdam  Differences Figure  the  assumed, g i v e n  chromosome  similarities  of  a v e r a g e complement  similarity cannot  show o b v i o u s  lengths  supported.by  length  15  Karyotypes  among t h e and  differences,  l a r g e s t and  P^ in  to d i s t i n g u i s h the  chromosome l e n g t h s ,  two  with taxa.  Inoue and  (1981)  smallest  endivi if o l i a concert  of  and other Based  Yamashita  69  (1980) c o n c l u d e d distinct  but  similarity  closely  i n the  a p p r o a c h was  E.  chromosomal  CONCLUSIONS AND  compromise clear  very  small  the  absolute  and  alignment  data  evidence  for that  became  of s s p .  but  exhibit  relative  lengths  pleurocarpous  Koponen  geographically similar  and N i l s s o n  The k a r y o t y p e s a  The  same  of L i m b e l l a .  formulae  t o some e x t e n t ,  tricostata  demonstrable  and s s p .  of some chromosomes.  Therefore,  isolated, "species pairs"  as in  fryei  so  far i s no  the  observed Mnium  in taxa  there  since  taxa among  (Lowry  1948;  1978). of s s p .  tricostata  and s s p .  fryei  indicate  c l o s e r e l a t i o n s h i p b u t show enough d i f f e r e n c e i n a b s o l u t e  relative additional  lengths  is  Both  t h e most common number  mosses.  it  differences  s e c o n d a r y p o l y p l o i d y has o c c u r r e d  morphologically  are  considerable  and l a c k of k a r y o t y p e  of L i m b e l l a  have t h e chromosome number n=11, reported  "show  pattern."  the karyotypes  sample s i z e  karyotypes  similar  because they  ontariense  SUMMARY  the karyotype  that  roseum and R_^  related,  used t o a n a l y z e  Although  are  t h a t Rhodobryum  to  justify  taxonomic c h a r a c t e r  using  this  to d i s t i n g u i s h  difference the  taxa.  as  and an  70  V.  A.  ISOZYME BANDING  INTRODUCTION  5.1 U t i l i t y  of E l e c t r o p h o r e s i s  Isozymes have p r o v e d biochemistry,  and  enzyme  structural  phylogenetics  by i s o z y m e s d u r i n g  staining  may  be  electrophoretic  interpreted  are  coding  f o r enzymes, and have been  calculate  interpreted  allele  genetic  1971,  identities,  phylogenetic method  1981,  genetic  diverse  similarity taxa  expressions  t o compare  of  genotypes  expressions  of a l l e l e s  analyzed locus  The d a t a diversity  between  taxa  a t gene  quantitatively polymorphisms, species  a r e used t o determine in  populations  (Gottlieb  1981).  and The  u s e f u l i n r e v e a l i n g both g e n e t i c d i f f e r e n c e s  between m o r p h o l o g i c a l l y genetic  as  Band  separation  d i s t a n c e s w i t h i n and between  1984).  comparisons  i s immensely  as  frequencies,  h e t e r o z y g o s i t y and g e n e t i c (Gottlieb  genetics,  plants.  Bands  to  of p l a n t  and s y s t e m a t i c s .  genes and may be used as a t o o l  individual  loci  u s e f u l i n the study  physiology,  patterns produced  of  PATTERNS  and  between  (Gottlieb  1984).  cytologically morphologically  similar  taxa,  and  and c y t o l o g i c a l l y  71  5.2 E l e c t r o p h o r e t i c  Procedure  Isozymes a r e d e t e c t e d which  proteins  (starch  in  crude  or p o l y a c r y l a m i d e ) ,  separated  on  the b a s i s  configuration. by  a  containing.a enzyme  in  (Shields  et a l .  enzymes c a n be d e t e c t e d of a c t i v i t y .  tissue allele one  is  simpler  Vries  et  a l .  that  of g e n e t i c  was  investigated  potential."  specific  enzymes  enzyme  sought,  reaction  that  to the product  1983).  In t h i s  the  of the way t h e  of  haploid  t i s s u e because o n l y  E a c h band u s u a l l y  one  represents  p r o d u c e two s u p e r i m p o s e d  variability  reviewed  (1983).  in  bands.  is  depleted  bryophtes,  and Wyatt  work s u b s t a n t i a t e d  polymorphism  dispelling  They p o i n t e d  by Cummins  Their  genetic  a genetically  s i z e and  of Bryophytes  populations  "are  may  and  bands w h i c h a p p e a r a t t h e  of d i p l o i d  per l o c u s .  that  plants,  1983; V a l l e j o s  on a g e l  charge,  each  f o r the  by t h e s t a i n e d  conclusions  vascular  for  stain specific  two l o c i  5.3 E l e c t r o p h o r e s i s  electrophoresis,  a  than  although  Investigation  one  for  in  current,  The i n t e r p r e t a t i o n o f zymograms  c a n be e x p r e s s e d  locus,  electrostatic  and s u b s t r a t e  and  are placed  to e l e c t r i c  a r e then assayed  catalyses,  reaction  subjected  solutions,  cofactor  electrophoresis,  tissue extract  of t h e i r  The g e l s  immersion  sites  by means o f g e l  as  high  the long-held group  out that  with  in as  belief  (1981) a n d de other  the  that  using  workers' bryophyte  reported that  limited  t h e same had been  for  bryophytes  evolutionary found  t o be  72  true  for  other  Lycopodium Both  "phylogenetic  lucidulum  monomorphic  mosses  and  Gliddon  and  and  and  as  review  Electrophoresis genotypic  by was  differences  populations  of  expected  to  observed  by  exhibit  and  active  w o r k e r s who  banding  is clonal.  pattern  —  i f at a l l —  on  the  L.  tricostata. reviewing  species,  the  If  alleles  parent"  that  ssp.  (1981,  the  ssp.  tricostata  fryei lack  only  ( G o t t l i e b 1984).  were  of  expected  that that  be  the  divergent based  derived  from  and  derivative  that  genetic  s m a l l , dependent  origin,  the  patterns,  differentiation  still  that  sporophytes  slightly  recently  very  recent  d e r i v a t i v e should  to  expected  emphasized are  were  sexually-reproducing  tricostata  1984)  s i n c e the  American  similar  also  was  (vid.  identifying  s t r o n g l y suggested  would be  fryei  d e r i v a t i v e i s of the  ssp.  ssp.  i t was  such s p e c i e s  time e l a p s e d  the of  of  North  c o m p a r i s o n s between p r o g e n i t o r  between  amount o f  However,  ssp. • f r y e i  Gottlieb  differences  propagation  from one  assumption  In  taxa.  of  and  the  for  1983). in  of  in  genetic  calculated  useful  investigated of  polyphemus.  phanerogams  polymorphism  Populations  vegetative  for  Populations  clubmoss  values,  been  Hawaiian  g e n e t i c a l l y monomorphic b e c a u s e  population  the  have  potentially  the  have been o b s e r v e d  V r i e s et a l .  genetic  bryophyte populations. t o be  de  as  Limulus  index  have been  between  Limbella.  other  loci  distances they  such  crab  Polymorphism  genetic  populations  1980  horseshoe  polymorphic  hepatics.  identities bryophyte  and  relicts"  "most  of  the  upon two  i f not a l l  present  in  the  73  B.  MATERIALS AND  5.4  METHODS  Electrophoresis Isozyme b a n d i n g p a t t e r n s  of  horizontal  of Helenurm  starch  maintained 1 cm  tips  of d i s t i l l e d  needed, plants  in  indicated  with  more  that  water  in a c h i l l e d  chilled  ml  The  higher  proportion  laborious  and  immersing  plants  grinding (Fahselt the  1980). thawing  rod i n 2 drops  i n the of  of  and  2 drops  of  the  A few powder  in ssp.  powder  drops of d i s t i l l e d to  make  a  were  was  of  a  slurry  distilled  223-  supernatant  much  improved.  made  grinding  simplified nitrogen  chilled  water  were  transferred  in liquid in  Whole  ( B i o Rad  fryei  task  above w e i g h t s  t i s s u e s to a  wicks  off)  The  i n t e n s i t y was  The  the  subsequent  blotted  a t 20,000 g.  stems  spot  extract  r e s u l t i n g p a s t e was  staining  five  of t h e m a t e r i a l .  tubing  time-consuming.  the f r o z e n  Initially,  polypropylene micro t e s t tubes  on w i c k s and  been  on a c h i l l e d  staining  ( e x c e s s water  f o r 15 min  plants  had  o n t o w i c k s and  grinding  The  Living  grinding 1981)  t h e methods  which  larger quantities  weight  mortar.  and c e n t r i f u g e d  absorbed  o r no  with crushed glass  1.5  was  et a l .  thorough  of 0.2-3.0 g wet ground  with a glass  Faint  (1984).  months.  then absorbed  gels.  following  material  f o r 1-4  (Szweykowski  the  then  to  chamber  were g r o u n d  water  enzyme a s s a y s  9501)  field-collected  the crude e x t r a c t s  inserted  to  L a y t o n and G a n d e r s  i n a growth  shoot  plate,  from  were o b t a i n e d by means  gel electrophoresis,  (1983) and  were p r o c u r e d  of L i m b e l l a  and  mortar  were t h e n  which  by  could  added be  74  transferred Freezing  to  did  the  not  micro  appear  test  to  tubes  alter  the  for  centrifugation.  staining  intensity  of  bands. Extracts two  gels,  Ridgway  buffer  handling  53  population plants (4)  w/v  and  11)  variation  these  to  14  g r o u p , and  A).  On  other  charged  gel  can  different  pairs  of  comparison  of  Hawaiian  of  gels,  of  individuals  their  49  of  with  M).  Gels  and  and  10  ease  % of  or  from  bands  of the  respective  Oregon p l a n t s  from 1  plants  sensu N e h i r a To  other  give  taxon  other,  taxon  the  Oelshlegel  giving  band m o b i l i t y  and  banding  populations,  on  together  run  next  (e.g.,  dissipation  were  small  1984)  detect  mobilities  sensu one  (3)  t a x o n were run  the  and  extracts Kauai  North American  f r o m one  using  different  cultivated  b e c a u s e uneven h e a t  ("bowing",  individuals  and  electrophoretic  distort  one  l o t 392),  from 4  material.  4 individuals  mobility  1973), p a i r s  plants  ("sporelings",  individuals  for comparison  Apendix  r u n s were p e r f o r m e d ,  cultivated the  and  on  1984).  mature  from s p o r e s  within up  (2)  in  enzyme r e s o l u t i o n  1 Maui p o p u l a t i o n ,  (Fig.  simultaneously  buffer  (Electrostarch  improve  Ganders  citrate  sources  i n d i v i d u a l Hawaiian  reciprocally  a  and  starch  added t o  grown  some g e l s  morpholine  electrophoretic  populations  19  %  ( L a y t o n and  (1)  were e l e c t r o p h o r e s e d  (recipes  12.5  Eight  as  with  s u c r o s e was  from:  Limbella  one  contained w/v  of  to  Fig.  from  the  appearance  of  and  Stahmann  alternated  with  a more d e p e n d a b l e  (e.g., F i g .  19  B).  75  5.5  Enzyme Gels  Assays, were  esterase  stained  (EST),  glutamic  phosphate dehydrogenase dehydrogenase  (IDH),  dehydrogenase  these  given  enzyme a s s a y s  (MDH,  PGI,  most  subsequent  i n t e n s e and  (SOD).  PGM,  runs,  regions  ("negative  tetrazolium  dyes,  staining"  because  exposed 6PGDH  negative  to all  achromatic  testing  staining  light.  Because  contained SOD  enzymes b e i n g of  bluish-purple  bands  i t inhibits  sought,  f o r two  color of SOD the  tetrazolium as  well  thus  being  auct.)  as by  SOD on  to  showed to  gels  i s e n h a n c e d when stains  f o r MDH,  salts,  all  dark-  of  dyes  and  enzymes w i t h a s i n g l e  a  assay.  as  blue their  Fridovich gels  PGI,  PGM  gels  are and  showed  s t a i n e d bands of t h e default  the  stained  formation  (Beauchamp  5  use  appears  tetrazolium  for  trials,  were s e l e c t e d  enzymes,  formazan which g i v e s g e l s s t a i n e d w i t h  The  initial  In c o n t r a s t  most  (PGI),  sources  best-resolved staining.  achromatic  1971).  After  and  consistently  of  malic  dehydrogenase  because they  typical  isocitric  isomerase  6PGDH, SOD)  bands  characteristic  glucose-6-  (LAP),  Recipes  i n A p p e n d i x M.  (DIA),  (HA),  6-phosphogluconate  staining  with  (GLDH),  aminopeptidase  dismutase  are  all  diaphorase  phosphoglucose  (PGM),  superoxide  enzymes:  dehydrogenase  leucine  stains  in  12  (G6PDH), h e x o s e a m i n i d a s e  (MDH),  phosphoglucomutase (6PGDH) and  for  labor-saving  other way  76  C.  RESULTS  5. 6 I n c o n s i s t e n t Banding However,  was  poor  and  apparent  bands  produced  was  of  6PGDH, PGI  plant PGM  the  of  intergrading  Because out  denaturation tricostata of  0.2-0.5  or  simply  often  from  or  populations  at  4°C,  c a u s e d poor slowly the  declined study,  a  null  ssp.  even  with  with  MDH,  if  the  existence  very  allele of  --  an  among faint I  a  and  series  group  bands t o  could  in  l e s s enzyme  to demonstrate a  electrophoretic  staining. in  distinct  of  clearly-  not  confirm  studied.  and  below  plant  determine  of  t h i s handicap  a l l extraction or  to  was  i n d i v i d u a l , while present  possible  no  show  inconsistent  intensities  B e c a u s e of  i n the  i n one  to  of  of was  It  g  However,  difficult  4  encountered  enough e x t r a c t  existence  staining  ranging  bands.  polymorphism  course  was  I t was  technical  intensity.  a minimum of  s t a i n i n g was  a band  the  extract.  individuals,  carried  of  it  tested.  from  difficulty staining  to y i e l d  polymorphism —  in  stained  consistent  and  systems  resulted  primary  runs t h a t  indicated  therefore  enzyme  somewhat ambiguous r e s u l t s f o r  material,  absence  others,  all  bands w h i c h were s c o r e a b l e .  and  apparent  The  necessary  dark-staining g  of  from t r i a l  material  in  resolution  5 enzymes t e s t e d .  obtaining  3.0  observed  band  difficulties, the  Staining  the fryei  it  is  procedures  unlikely  A l t h o u g h the growth thrived,  vigor  chamber yet  that  the  of  were heat ssp.  during  the  latter  also  77  showed v a r i a b l e s t a i n i n g . of  extraction  denatured Stahmann enzyme matter  by  or  assays of  of  this  study.  5.7  The  Enzymes  unsuitable  and  Limbella  would  scoring  appeared others  t o be  in  these  monomorphic and  were u s u a l l y  bands were the  tested,  and  tricostata invariant, activity,  can  PGM  ssp.  tricostata  2 migrating  the  and  SOD  it  and  origin  or  segregation  ssp.  and  zone showed  into 2 fractions.  buffer  and  cathodally-migrating  or and  systems  and  time-consuming  beyond the  6PGDH In  limits  precluded  some, b a n d i n g  i n both taxa, between t h e  while  in  taxa  (Figs.  the  taxa.  between  The  among the some fryei  5 enzyme  detail.  consisting  1 cathodally faster other  zone d i d not  Both  of  m o b i l i t y and  19  of  sometimes  smearing  Baur and  own  E).  anodally-migrating  appear  L.  3 zones  (Fig.  s e n s u O e l s h l e g e l and  f r a c t i o n s (sensu  systems  displayed their  showed e i t h e r much  ("tailing"  i n t o 3-4  in  pattern  anodally  showed s l o w e r m o b i l i t y and  oxidized  evident.  described  monomorphic  selection  (Oelshlegel  systems.  identical  best-resolved  be  anodally-migrating  segregated  The  PGI,  enzyme  be  have been a  Band m o b i l i t i e s u s u a l l y d i f f e r e d  to 3 l o c i SOD  .One  optimal  c o n f i g u r a t i o n s seemed t o d i f f e r  19 A-D). One  ranges  e r r o r w h i c h I j u d g e d t o be  Poor band r e s o l u t i o n of MDH, reliable  have been my  pH  C a r e f u l s e l e c t i o n of for  trial  e r r o r may  g e l b u f f e r s , s i n c e enzymes can  b u f f e r s of  1973).  The  between  Stahmann Schorr  to segregate  zone  1973)  1969). into  78  F i g u r e 19 - Zymograms o f s s p . t r i c o s t a t a and s s p . fryei. Dark z o n e s = enzymes s t a i n e d f o r ; w h i t e z o n e s = SOD a c t i v i t y . A l l g e l s run i n Ridgway b u f f e r s y s t e m . A. PGI;  - MDH;  Hawaiian  w i t h Oregon plants plants (with  14 H a w a i i a n p l a n t s plants  plants.  (R). (R). faster  D.  C.  (with - PGM;  - 6PGDH; E.  SOD  -  SOD  Sporelings  cathodally-migrating  SOD  14 Oregon Hawaiian  plants  mobility)  plants plants  Oregon  Kauai. bands.  3.  material. Sporelings  Horizontal  line  4  Hawaiian plants.  2.  Oregon plants 1.  Reciprocally-  from =  B.  alternated  (L),  a r e MDH);  (R).  (L) , 4 H a w a i i a n  a l t e r n a t e d w i t h Oregon  Hawaiian m a t e r i a l . from  faster  ( d a r k bands  mobility)  Reciprocally-cultivated cultivated  14  ( L ) , 4 Oregon  Maui.  origin.  4. Note  79  80  fractions.  Cathodal banding  electrophoretic only.  Fractions  tricostata it  runs and  was  et a l .  indistinct  was  two an  taxa: ssp.  they  rather  p a t t e r n s were not  never known  Because  consistently  as  a  single  as  a  separate  Pellia.  i n SOD  intense  within  zone  material  scoring  Following  negative  identical  ( C h a p t e r 7)  t o t h o s e of  influenced  of  band.  when  mobility  from b o t h K a u a i  showed  their  by* d i f f e r e n t  tricostata  ssp.  been a n a l y z e d p r e v i o u s l y ssp.  analyzed.  of  buffer.  this  between  greater  staining.  (mature band  plants)  patterns.  of b o t h t a x a parent  the  showed  stocks:  environmental  the  regimes.  DISCUSSION Limbella  was  i n ssp.  apparent.  more  tricostata  cultivated  segregated  showed bands of c o n s i s t e n t l y  usually  band p a t t e r n s i d e n t i c a l  D.  were  (sporelings)  Reciprocally  in  for in a l l  documentation  t h e same a p p r o a c h  obvious d i f f e r e n c e  of s s p .  Maui  each  bands loci  for  citrate  fractions  than  stained  here  grouped  (1981) used  SOD  not  more c l e a r l y  were  tricostata  and  Populations and  often  peroxidase  intepretation,  mobility  mentioned  to d i f f e r e n t i a t e  activity  Szweykowski  two  was  run on g e l s w i t h m o r p h o l i n e  a zone o f a c t i v i t y ,  There  is  were  difficult  enzymatic  activity  tricostata  Because s t a i n i n g  inconsistent, t h e two  is  taxa.  little SOD  tr icostata f o r isozymes,  the f i r s t  and and  Hawaiian  ssp. as  fryei far  bryophyte  o f 4 of the 5 enzyme systems can  banding,  be  said  about  however, was  genetic clearly  have  as to  is be  tested  variability resolved  and  81  consistently  monomorphic w i t h i n  in  band m o b i l i t y between  in  p l a n t and  be  1980;  monomorphic  in  1981).  Because  populations,  and  plant  that  populations  all  ssp.  Consistent  taxonomic c h a r a c t e r ,  of  the  their  tricostata  hepatic  species  conclusion  was  Pellia  additional  followed  in this  Limbella  showed bands i d e n t i c a l the  other  Gottlieb  A.  (1981) was  not  CONCLUSION AND  Banding  Krzakowa  patterns  it  ssp.  evidence  The  The  which they i n banding  is SOD  fryei useful  Szweykowski for retention  They and  supported cytological  same a p p r o a c h  reciprocally parent  in 5  e x h i b i t an  and  biochemical  to t h e i r  variation  was  cultivated  stocks,  not  grew, i n d i c a t i n g patterns  to that  cited  by  detected.  SUMMARY were  tested,  but  enzyme  s y s t e m e x h i b i t e d the  obtained  r e s u l t s were ambiguous  monomorphic w i t h i n  usually  islands,  the  to and  monomorphic  Krzakowa  1977).  of L i m b e l l a .  taxon with  environmentally-caused  of  Yeh  is  b o r e a l i s Lorbeer.  (Szweykowski and  of  was  (1981) as  evidence  those  1980;  SOD  populations  u s e d by  et a l .  with  study  been r e p o r t e d  El-Kassaby  from t h a t  monomorphic  d i f f e r e n c e i n band m o b i l i t y i s a  and  Szweykowski  has  from 2 d i f f e r e n t  consistently different  and  rarely  1969;  showed a d i f f e r e n c e usually  Yamazaki  probable  (1977)  only  is  Schorr  Limbella  population.  SOD  but  (Baur and  different  mobility  taxa  taxa.  a n i m a l t i s s u e s and  polymorphic  O'Malley  the  the  each taxon  best but  for  f o r 4 of  all  enzyme  5 enzymes.  r e s o l u t i o n , and  systems The  appeared  showed c o n s i s t e n t  SOD to  be  differences  82  in  mobility  difference two  between  the  taxa.  i n SOD m o b i l i t y was  different  islands,  and  mature p l a n t s .  The c o n s i s t e n t  a  difference  fixed  fryei the  two  genetic which taxa.  was  u s e d as  In  identical  ssp. in 5  identical differences  between s s p .  tricostata, populations  in both s p o r e l i n g s in m o b i l i t y tricostata  an a d d i t i o n a l c h a r a c t e r  to  the from and  indicates and  ssp.  distinguish  83  VI.  A.  PHENOLIC  CHROMATOGRAPHY  INTRODUCTION Literature  phenolics  in  were r e p o r t e d (1978), acid  documenting bryophytes  and  diglucoside In  flavonoids recently Suire  and  literature taxa  esters  contrast,  Asakawa  f o r compounds a n d / few  implications  abundant  and  1981)  and  Huneck  or  elucidating  Hepaticae  (vid.  ubiquity, old),  stability  McClure  characters  (obtainable  interspecific  environmental  (1978) e m p h a s i z e d  Most  screening structures.  phylogenetic  Markham e t a l .  in  and  1978;  1978; C a m p b e l l e t  the value plants,  from m a t e r i a l  differences,  (1978),  i n b r y o p h y t e s , and most o f  1980).  taxonomic  reviewed  (1984).  taxonomic and  al.  potential  Nilsson  been  chemical  ( 1 9 7 8 ) , Koponen a n d N i l s s o n  as  and  the occurrence of  has  Markham a n d P o r t e r  Markham e t a l .  Caffeic  i s i n t h e form o f s u r v e y s ,  of f l a v o n o i d o c c u r r e n c e s  1979; Markham  Nilsson  (1980).  by M a r t e n s s o n  documenting  p a p e r s have e x p l o r e d  t h e s e have c o n c e r n e d o n l y  derivatives  ( 1 9 7 8 ) , Markham and P o r t e r  (1979,  the subject  non-flavonoid  Cinnamic a c i d  (1978) and Zehr  literature is  of  (1967), Koponen and  were r e p o r t e d  Markham e t a l .  on  Relatively  i s scarce.  Zinsmeister  i n bryophytes by  occurrence  by M c C l u r e a n d M i l l e r  Mues  (1974).  the  up  relative  of  flavonoids  citing  their  to  years  85  resistance to  modification. and  Miller  (1967)  performed  the  first  major  84  flavonoid 0.1  g  survey  dry  flavonoids  mosses,  weight, in  flavonoids  34  mosses. they  of  them  acrocarps,  latter in  these.  20%  in  lacking  small  pleurocarps  were v e r i f i e d  1977;  Aeon e t a l .  1983).  ), w h i c h t h e y  Because t h a t they  verify be  or r e f u t e  provide  an  and  Bohm  ( Christy fryei  (  their  1979).  (Vandekerkhove  screened  by  McClure  s s p . t r i c o s t a t a (as  devoid  used enough p l a n t  of  flavonoids.  i t was p o s s i b l e  material  results.  If flavonoid differences  taxonomic  and s s p . character  to  to detect  i t worthwhile  tricostata  in  attempt  fryei,  this  with  which  to  could would to  the taxa.  plants  (1980).  METHODS were t r e a t e d  5209, 37.3 g; C h r i s t y 5334,  s a m p l e s were a l s o  tested  a t UBC.  following  t h e methods o f G o r n a l l  Two s a m p l e s o f s s p . t r i c o s t a t a  Christy  deposited  be  and Asakawa  I thought  additional  MATERIALS AND Dried  have  between s s p .  distinguish  B.  not  i n Limbella,  detected  to  that  pleurocarpous  workers  tricostata  found  occurrence  s t u d y was 18 y e a r s o l d , a n d b e c a u s e may  flavonoids  (70%  One o f t h e t a x a  found  they  detected  pigments e n t i r e l y or  Suire  by l a t e r  (1967) was L i m b e l l a  Sc i a r o m i u m  that  fid.  and  than  the  quantities  observations  Miller  species  acrocarpous  Their  and  70  samples a s s m a l l a s  Significantly,  in  either  very  Working with  screened  were more common  the  having  in  were  5245, 69.6 g) a n d one  46.0 g ) .  tested of s s p .  One-gram q u a n t i t i e s o f t h e s e  separately.  Voucher  specimens  were  85  100% run  Phenolics  were e x t r a c t e d  methanol.  The  on  based  two-dimensional solvents.  b-aminoethyl under the  n-butanol  from whole L i m b e l l a e x t r a c t s from t h e  chromatograms u s i n g  Phenolics  were d e t e c t e d  diphenylborinate  long-wave u l t r a - v i o l e t  light.  l a r g e r samples were l o a d e d  eluted  with  methanol column,  a s e r i e s of  plate.  and  visualized  phenolics  phenolic  activity  solvent  as  were t h e n  above.  Bands  phenols e l u t e d with plate  to detect  with and  0.1% viewed  and  were  and  increasing  collected  spotted  above.  i n the  from  the  s e q u e n t i a l l y on benzene  a  solvent  I n d i v i d u a l f r a c t i o n s with  streaked were  s o l v e n t s of  on  a  scraped  plate  and  f r o m the  m e t h a n o l , d r i e d down, and  activity.  benzene-  o n t o a Sephadex LH-20 column  P l a t e s were run as  were  from  dryness  chromatographic  hot  n-butanol e x t r a c t s  Fractions  to  spraying  C h e m i c a l Co.)  The  in  1 g samples  a q u e o u s - and  by  aqueous m e t h a n o l  concentration. evaporated  (Aldrich  plants  run  in  plates,  the  spotted  again  on  a  P r o c e d u r e s were t e r m i n a t e d  after  this  and  the  step.  C.  RESULTS Both the  t w o - d i m e n s i o n a l chromatograms  sequentially-spotted activity. after  Blue,  spraying Faint  disappeared purplish  fractions  yellow  but  and  21)  orange spots  bands  eluted  UV-absorbing  spots  and  visible run  typical  on  again of  20)  exhibited  were v i s i b l e  were c o l o r l e s s i n v i s i b l e  UV-absorbing when  (Fig.  (Fig.  phenolic under  UV  light. streak by  chromatograms  themselves.  f l a v o n o i d s were  No  detected,  86  Figure  20 - P h e n o l i c  spot  patterns  on t w o - d i m e n s i o n a l  chromatograms. A. fryei shading  - ssp. (Oregon,  tricostata Christy  (Kauai,  5336  = blue.  Aqueous-based solvent •  ).  C h r i s t y 5247 Light  shading  ).  B.  - ssp.  = orange,  dark  87  Figure A.  21 - P h e n o l i c s p o t p a t t e r n s o f s e q u e n t i a l l y - s p o t t e d f r a c t i o n s on o n e - d i m e n s i o n a l chromatograms. - ssp.  tricostata  t r i c o s t a t a (Maui,  (Kauai,  Christy  5334  ).  shading  = blue,  Christy  Unshaded striped  5245  Christy ).  = yellow,  shading  C.  light  > i—I  o XI  w CO  000«@o  ).  B.  -  ssp.  - ssp.  fryei  shading  = orange,  = yellow-blue.  A  c  5209  (Oregon, dark  88  nor  were any  D.  DISCUSSION  The ssp.  spots  fryei  other of  s u c h bands seen d u r i n g  than  the  most  flavonoids.  My  analyzed  observation  for  that  were c o r r e c t  follows:  acid 1978; et  al.  to  and  ssp.  of  than  structures  7  times  Miller  tricostata fryei  plant  was  (1967),  never  and  material  available  for L^  more  contains  has  McClure  no  before  Miller's yielded  in  no  0.3  tricostata.  They  compounds  of  variously-colored  i n b r y o p h t e s have been r e p o r t e d (a)  and  blue Nilsson  =  caffeic  1974)  (Koponen and  1980) 1968;  and  phenolics  in  g also L.  tricostata.  derivatives Zehr  McClure  that  least  up  2 blue-fluorescing  identities  spots detected  p r e s e n c e .of  and  t h e i r i d e n t i t y remains  compounds.  at  in reporting  Putative  (Martensson  these  reliable  ssp.  by  tricostata  information  is  tricostata  used  tricostata  spectra  obtained,  larger quantities  flavonoid  quantities  as  than  Limbella  the  r e s u l t s , using  their conclusion  flavonoids.  more  not  separation.  ssp.  However, b e c a u s e  were  material  column  of  indicated  chromatographic  substantiate  been  chromatograms  likely  compounds  unknown. plant  seen on  the  and  (c)  b l u e or  Nilsson  o r a n g e and  Karunen and  exception  of M a r t e n s s o n  and  Ihantola  (1977),  of  none  (b)  acid  yellow  Ihantola  1977;  Nilsson these  1978;  chromatographic  i n the  literature  diglucoside blue-green Mues and  =  esters cinnamic  Zinsmeister  = carotenoids Zehr  (1974)  1980). and  investigators  (Bendz With  the  Karunen  and  reported  the  89  structures their  of  the  compounds  conclusions McClure  phenolics readily species  are  entirely  Miller  (1967)  were o f  little  taxonomic  by  the  flavonoid  phenolics  interspecific  variation  by  McClure  non-flavonoid  phenolic  of  little  these  taxonomic  compounds."  phenolics ,  but  in ssp. because  tricostata  as  of any  tested.  Therefore,  E.  Analysis demonstrated compounds.  present, (1967).  of the L_^  non-flavonoid  because  are  show l e s s v a r i a t i o n  from  However,  be  Zehr  non-  than  (1980) f o u n d  that  t o o much  between s p e c i e s , r e n d e r i n g coupled  with  was  much  as ssp.  taxonomic  of  of  chromatographic  tricostata  within  I c o u l d not  significance patterns  of  non-flavonoid  ssp.  variation  fryei,  them  identification  patterns  from t h o s e  two  if  alterable,  greater  differed  the  that  "they  to  chromatographic  ssp.  discern  among t h e  were not  a  plants  u s e d as  a  taxa.  SUMMARY up  t o 70  g of  presence  of  tricostata  for phenols. substantiating The  and  tend  "unless  to d i s t i n g u i s h  that  value  Miller.  i t and  appears  environmentally  variation  apparent  CONCLUSION AMD  screened  and  it  flavonoids."  would  the're  between  the were  fryei  pattern  character  do  use The  reported  environment  t o s p e c i e s than  and  speculative.  and  altered  indicated  in question,  It  the  tricostata  several  ssp. is  chromatographic  ssp.  findings patterns  ssp.  unidentified  fryei  unlikely  and  had that  of  never  fryei  phenolic  before  flavonoids  McClure  and  of p h e n o l s v a r i e d  been were  Miller between  the  samples  tested,  rendering  them o f l i t t l e  taxonomic  use.  91  VII.  A.  EXPERIMENTAL  INTRODUCTION Cultivation  of  mosses  (1) h e l p d e t e r m i n e what of  CULTIVATION  gametophytes  remain  stable  morphological  i n "common g a r d e n " e x p e r i m e n t s c a n  factors control  and (2) i d e n t i f y under  different  which  describes  reviews  the  cultivation  taxonomic  plasticity  morphological  features  environmental conditions.  f e a t u r e s c a n be u s e f u l  chapter  morphological  taxonomic  Stable  characters.  experiments  in  This  Limbella  and p h y t o g e o g r a p h i c i m p l i c a t i o n s  and of the  results.  B.  UTILITY The  OF MOSS CULTIVATION  importance  contributions  and  1982; Wyatt  Richter for  1984).  a  under  controlled In  for  reviewed r e c e n t l y and S t o n e b u r n e r  Cultivation  experiments r e q u i r i n g  also  useful  tool  live  which  of  mosses  investigators  and  the  using  this  ( S m i t h 1978; L o n g t o n  1984;  Frahm  and  plants  (e.g.,  1982;  Nordhorn-  p r o v i d e s stocks of l i v i n g  w h i c h may  cytology)  material and  be u s e d t o compare p l a n t s  is  grown  conditions.  theory, c u l t i v a t i o n  taxonomic  identify  cultivaton  methods o f numerous  t e c h n i q u e have been Zander  of  purposes  o f mosses under should  morphological  enable  uniform the  characters  conditions  investigator are  to  genetically  92  controlled  and  which  characters  Genetically-controlled  characters  variety  r e g i m e s and  of c u l t i v a t i o n  populations, be  modified  habitat  by  i n the  P l a n t s may  different  become e t i o l a t e d reduced  Gametangial  production  may  conditions  different  taxa  c a n n o t depend studies  to  Cultivation mosses Zander  are  on  discern  so  Despite taxonomic  Stoneburner  tool,  may  or  widely to  in wild should  different  the  shown  develop  observed  i n the  field.  reduced  leaves,  rhizoidal  certain  working  with  synonymous t a x a ,  derived  from  and  in  general  maintain  than  cultivation Smith  acrocarps.  questions  of  (1978)  concerning  s t u d i e s and  such can  them.  pleurocarpous are  from c u l t i v a t i o n  one  cultivation  o r d i f f e r e n c e s between  limitations  by  development.  indistinguishable  When  putative  wider  Under  become  penetrating  limitations,  (1984).  a  often  1982).  information  as  stable  and  suppressed.  information  several  identified these  be  difficult  of  those  and  similarities  (1982) a s k e d  under  characters  regimes  exhibit  taxa,  methods d i f f e r  interpretation doing  similar  more  from  Zander  solely  a l s o be  Cultivated plants  branching  (sensu  morphologically  should  stable  of mosses i n v o l v e s b o t h t e c h n i c a l  and  internodes,  phenocopies  remain  altered.  field.  interpretive difficulties. quite  should  cultivation  practice, cultivation  appearances  environmentally  environmentally-altered  different  conditions  In and  whereas  are  by  experiments. be  and  a  powerful  Wyatt  and  93  C.  CULTIVATION OF  7.1  Need  for Live  Aside for  from  was  analysis.  Material a need t o m a i n t a i n  comparative  material  LIMBELLA  studies  also  under  needed  for  Because the l a t t e r  needed t o be accumulate  repeated  over  sufficient  data,  be m a i n t a i n e d  characters  phenetic that  fryei,  stability  before  segregate  the taxa.  under  reciprocal obtain  Limbella  conditions,  electrophoresis  and  live  karyotype  were t i m e - c o n s u m i n g and  a  of  period  time  in  e s s e n t i a l that  order  living  to  plants  of t h e e x p e r i m e n t s .  analysis  appeared  assessed  fryei  of  of L i m b e l l a  -  Whole P l a n t C u l t i v a t i o n  Because  ssp.  cultures  procedures  7.2 Need f o r C u l t i v a t i o n S t u d i e s  1.  controlled  i t was  f o r the d u r a t i o n  living  they  a  two  to d i s t i n g u i s h ssp. of  could  these  differences  variety  tricostata  be u s e d a s t a x o n o m i c c h a r a c t e r s  to  of  culture  needed  from be  provided  evidence.  morphological  to  C u l t i v a t i o n of s s p .  cultivation,  the needed  identified  tricostata conditions,  a potentially  useful  and  ssp.  including tool  to  94  2.  Spore  In  order  as p o s s i b l e to  culture  Cultivation to test  them.  Furthermore,  Fragment  Because  later  this  sporophytes  handicap  i t had  area,  vegetative  fragments  as propagula.  ssp.  tricostata  vegetative appeared  D.  or  (sensu  Nehira  —  specialized  i n ssp.  necessary  branches,  v e g e t a t i v e propagula  fryei,  colonized to  shoot  test  tips  The r e g e n e r a t i v e c a p a c i t y a l s o needed were  and b e c a u s e a  i n spite relatively  the v i a b i l i t y of  and  single  of such  fragments of  t o be a s s e s s e d t o d e t e r m i n e as  effective  leaves  propagules  whether  as spores  t o be.  MATERIALS AND METHODS  cultures  UBC.  i t was n e c e s s a r y  sporelings  successfully  i t was  fragments  Cultures  of  the  spores  Regeneration  extensive  —  t ricostata  in electrophoresis.  s u c h a s gemmae were unknown of  of s s p .  v e c t o r s of l o n g - d i s t a n c e d i s p e r s a l ,  1984) c o u l d be used  3.  the v i a b i l i t y  were m a i n t a i n e d  f o r n i n e months, e x c e p t  w h i c h were m a i n t a i n e d  a l l cultivated  material  f o r 13 months.  except  sporelings  Voucher  for  spore  specimens  were d e p o s i t e d a t  95  7.3  Whole P l a n t C u l t i v a t i o n Living  firstWet  and  material  ssp.  second-order  plants  were  refrigerated collected plants  of  as  in  as  Kauai  in  in  possible.  fluorescent  lighting  dark a t  and  a  100%  RH,  i n May  Sutton  600-800  collected 1984  Lake  E15  conditions  and  ssp.  fryei  was  i n May  1984.  The  chamber  16  2  11).  bags  growth  E i n s t m~ ,  from  (Fig.  plastic  Living  Conviron  of  was  "ziplock"  same manner a t  were p l a c e d  18°C  s t r e a m s on  sealed  soon the  tricostata  hr  approximating  under  light:  those  8 hr  in  the  field.  1.  C u l t i v a t i o n of H a w a i i a n  Plants  Six  collections  tricostata  12x12x25 cm All  perforated  of  plastic  t r a y s were immmersed  plexiglas  tank.  The  pump and  aerated  with  to  approximate  the  emergent water  changed  contamination of  the  contamination, chlorinated  —  water was an  depth  of  stone. those  a b o v e the  biweekly.  I  plants.  replaced  collection water  After  In an the  an  the  surface  of  tray.  Lauda  d i d not  T-1  immersed  field,  enabling  water.  months,  necessitated  effort  cm  were  to  dechlorinated  s l o w e d but  per  MGW  the  five —  in six  i n a 20x50x60  Plants in  e s p e c i a l l y Cyanophyta tank and  were p l a c e d  c i r c u l a t e d with  aquarium  tap water, which  proliferation.  t r a y s , one  in dechlorinated  shoots to protrude  was  cleaning  ssp.  algal frequent  reduce water  eliminate  The  algal with algal  96  2.  C u l t i v a t i o n of N o r t h A m e r i c a n  Seven  c o l l e c t i o n s of  transparent attempt were  plastic  to  ssp.  boxes  with  replicate field with  1 cm  were m i s t e d  with  dechlorinated  b o x e s was  7.4  Spore C u l t i v a t i o n Because  tricostata tricostata hr  and  following placed  c o l l e c t e d on  the  in  grown  conditions nutrient and  18°C  in  a  identical  to was  the  dishes  (Nehira  in  19  study.  Sporelings most d i s h e s ,  plants Water  in  incubator  two  solution tested  with  for  (1965)  and  with  autoclaved  The  sporelings  under c o n t r o l l e d chamber.  weeks.  Because  involved  g r o w t h were  The mixing  considerable  in dechlorinated  water,  observed,  dechlorinated  in s t i l l  48  extracted  growth  were grown u n d e r d i f f e r e n t  s p o r e l i n g s were grown  ssp.  38°C  Hatcher  filled  ssp.  of  S p o r e s were  1966).  the  changed every  replaced  an  boxes  unknown, o n l y  Sporangia  (1952) and  petri  nutrient  the  the  Maui were d r i e d a t  d i f f e r e n c e s i n subsequent s o l u t i o n was  the  Schelpe  s p o r e l i n g growth was  are  8 months.  those  and  cm  In  times weekly.  fryei  P s y c h r o t h e r m R-27  nutrient of  for  35x10 mm  3-5  lids.  b o t t o m s of  water,  cultured.  Knopf's s o l u t i o n  solution  b e c a u s e no  ssp.  be  the  i n 8x20x20  weeks.  K a u a i and  methods of  autoclaving  effort,  at  twenty  half-strength were  could  water  3-5  s p o r o p h y t e s of  stored  tight-fitting  of d e c h l o r i n a t e d  changed every  spores  were p l a c e d  conditions,  filled  the  fryei  Plants  aeration  water  all  i n week  regimes.  (standing)  and  In  water.  97  Some  dishes  plastic  submerged  box f i l l e d  aquarium current 41  were  stone,  with  weeks  while  with  obtained  by c o u n t i n g  Fragment  1.  aerated  with  an  tricostata  transferred  sporeling densities of 1  to  plants.  After  filter  paper  w a t e r , and grown  in the  petri  in petri  dishes  were  cm . 2  Living Material of  detached  fragments  ssp. from  consisted  2-3 mm  l o n g and  (Table  2).  diaspores, of  replicate  moist  of (1) s i n g l e  filter  segments  paper  during  leaves and  their  fryei  handling.  branches viability by  petri Filter  d e c h l o r i n a t e d water w h i c h was  The  shoot  1-7 as  cm  tips long  vegetative  type,  (Gemmell 1953; M i l l e r  type.  frequently  (2) a p i c a l  segregated  1981) i n 100x15 mm  d i s h e s per fragment  and s s p .  plants  ascertain  1980,  autoclaved weeks.  whole  (3) s h o o t To  tricostata  f r a g m e n t s were grown,  1976; S e l k i r k  two  a transparent  Regeneration  Fragments  with  water  distilled  subsets  in  were submerged and s u b j e c t e d t o  were  autoclaved  Average  sheets  others  a l l sporelings  dishes.  became  dechlorinated  and a e r a t i o n w i t h m a t u r e s s p .  moistened  7.5  experimentally  on  and  dishes, p a p e r was  4-5  Ambrose  with  2-5  moistened  replenished  every  98  Table  2 - Living  m a t e r i a l of L i m b e l l a studies.  ssp.  1.  single  leaves  2.  shoot  tips  3.  shoot  segments,  1-8  cm  tricostata  used  in  regeneration  spp.  50  50  20  20  120  80  fryei  99  2.  Dried  In an 1979a, ssp.  Material  effort  to  above.  were p l a c e d  Ages  and  recent In  a casual  ssp.  fryei  hr  25  filter  7.6  was  experiment  drying  paper a s  described  without cm  of  fryei nylon  were  in four  dechlorinated  c o n d i t i o n s of  ssp.  attached  and  placed  ssp.  with  15 cm  were  i n the  rubber  material  (2)  two  and  were  plastic  of  temperature  20°C f o r on  from t h e  moist  ssp.  placed  fryei  outflow  for  alone,  boxes  filled  the  cultivation  collections  tank  24  prepared.  of  bands t o a r o c k - f i l l e d  i n the  of  same c o n t a i n e r s  replicating  Portions  the  experiment.  placed  tricostata  8x12x20 cm  immersed  hr and  tricostata  water,  fryei•  window s c r e e n i n g ,  tricostata,  ssp.  fresh  drying  Only  drying  r e p l i c a t e s were  experimentallly  Some p l a n t s of immersion,  of  No  3.  f o r the  and  described  t r e a t e d , and  in Table  40°C f o r 24  above.  of c o l l e c t i o n s together  taxa  given  from e a c h t r e a t m e n t  Portions  10 weeks.  the  regeneration,  (1)  Cultivation  grown  of  (Zander  tricostata  t r e a t e d as  t o d e t e r m i n e how  affect  d r i e d at  fragments  p a p e r and  ssp.  a v a i l a b l e were used  Reciprocal  were  1  of  filter  ( i f known) a r e  collections  duration  and  on  identities  p r o c e d u r e s employed  and  d r i e d herbarium m a t e r i a l  1979b), e n t i r e s h o o t s or b r a n c h e s of fryei  most  regenerate  containing of  the  pump t o e n s u r e growth under c o n d i t i o n s of c o n t i n u o u s  with  ssp. bag  of  ssp.  circulating current.  100  Table  3 - Dried  Taxon  Limbella  material  used i n r e g e n e r a t i o n  Date  Drying  study.  Time  tr icostata  ssp.  tricostata  1983  ssp.  fryei  1983  "  "  48 hr @ 38 deg C H  it  1922  ?  L.  capense  1984  ?  L.  pachyloma  1976  ?  1981  ?  1980  ?  " Platylome-lla  lescurii  H  101  7.7  Shoot  Orientation  New  s h o o t s grown  petri to  dishes  either  Experiment from L i m b e l l a  exhibited  ssp.  two  different  tr icostata  or s s p .  movements were g e n i u n e  growth  by  material  shifting  of  the  segments were s t r i p p e d to  filter  Twelve 120  paper  replicate  shoot  under  60  of  paper,  b r a n c h e s and  each  conditions  fishing  taxon.  identical  peculiar  not a r t i f a c t s  the f i l t e r  monofilament  paper i n  each  line  The  to  these caused  1 cm  firmly  shoot  attached  (Fig.  d i s h e s were p r e p a r e d i n t h i s manner,  segments,  maintained  filter  To e n s u r e t h a t  r e s p o n s e s and on  on  orientations, fryei.  of e x i s t i n g  with  fragments  22).  containing  dishes  were  t h o s e of t h e s p o r e  cultures.  E.  RESULTS  7.8  Whole P l a n t  1.  Hawai i a n P l a n t s  Cultures a  growth  to  34°C  of both s s p .  f o r 12-18  in  occasionally  hr.  are often  tr icostata  decline  tricostata  chamber m a l f u n c t i o n a t w h i c h  temperatures ssp.  Cultivation  Glime  lethal  survived  vigor  and V i t t  severe a l g a l  4  ssp.  fryei  survived  time the temperature (1984) r e p o r t e d  to aquatic  the h i g h  after  and  mosses.  that  contamination.  which  high  L_^ t r i c o s t a t a  t e m p e r a t u r e but began a months  rose  coincided  Most o r i g i n a l  slow with  Hawaiian  1 02  Figure  A. line, -  holes  view.  22 - Means o f a t t a c h i n g s h o o t segments t o paper, f o r shoot o r i e n t a t i o n experiment.  sectional knotted at  view: ends.  bored i n p l a s t i c 6.  -  line  1 -  shoot  3 -  filter  disc  scored with  segment. paper.  with pencil.  hot  2 4 -  -  filter  monofilament  plastic  needle.  B.  disc.  5  Surface  103  growth new  turned  black  or  branches continued  t o be  culture  period.  I t was  caused  by  high  the  contamination. remained  the  2. The cm  produced  in t y p i c a l  throughout the  and  ssp.  basal  of  the  cultures of  1984).  Spore  tricostata  i n s e r t i o n angle in typical  stages  sporelings  Bryum t y p e ,  investigated sporelings produced  length  and  ssp.  6  months.  and  hyphae.  died,  Yellowing  (Frahm and basal  to  Nordhorn-  leaf  cell  length  of  the  ssp.  fryei.  Cultures  Developmental  the  was  algal  s h o o t s up  yellowed  fungal  Richter  7.9  cell  b r a n c h e s a f t e r 2-3  r e s u l t e d from n u t r i e n t d e f i c i e n c y  same as  in vigor  .  vertical  possibly  remained the  leaf  tricostata  lateral  proliferation  Leaf  decline  nine-month  n u t r i e n t d e f i c i e n c y or  produced v i g o r o u s  7 months, p o r t i o n s  the  However,  Plants  developed  by  were o b s e r v e d .  whether  temperature,  Oregon p l a n t s  accompanied  fungi  i n s e r t i o n angle  same as  which  No  unclear  North American  long,  After  Leaf  died.  typical  (Nehira resembled  by  are  mature  and  growth  shown i n T a b l e of  other  1984). the plants  i n the  squarrose  no  or v e r y  Sporelings  Amblystegiaceae the  flagelliform'  widely-spaced  with  4.  Throughout  (0.4-0.8x0.1-0.3 mm), leaves  rates  and  reduced  sporelings  culture period  branches  field,  were of  occasionally  e x h i b i t i n g very  often  the  small  wide-spreading  l i m b i d i a and  sparse  to  104  Table  Week  4 - Developmental  s t a g e s of s p o r e l i n g  Activity  1  germination  4  branched  1 1  shoot  13  shoots  16  limbidia  Observed  (a few d i d n o t g e r m i n a t e  protonemal  secondary  strong  visible,  unistratose;  a s i n mature  rhizoids  secondary  shoots arched,  a r e o l a t i o n and  plants  rhizoid initiation  a t merophytes;  branching of shoots  r h i z o i d a l attachment  substratum 30  week 9)  elongated  some m o n o p o d i a l 26  until  wefts  buds; s h o o t s d e v e l o p i n g w i t h  dentition 19  cultures.  some  aerial  to  1 05  chlorophyll. oxygen,  carbon  1984). with  The e t i o l a t e d  Figure  higher  developed those  23 shows r e d u c e d  sporeling  Aeration plant  at approximately but  In  general,  and l a r g e r l e a v e s  within  w h i c h were s u p p r e s s e d w i t h i n water.  filter shoot aerial  in  a l l sporelings  dishes  t h e same time a s d i d  shoot  elongation  was  t o improve g r o w t h r a t e s and  two weeks three  in dishes  crowded  s p o r e l i n g s produced of  larger  aeration,  weeks a f t e r b e i n g  shoot  features  returned  to  been t r a n s f e r r e d t o m o i s t  developed  more c h l o r o p h y l l , more  larger leaves,  lateral  branches  larger  and  more  shoots.  7.10 Fragment  1.  Regeneration  Living Material  Developmental are  rates  stature.  T h r e e weeks a f t e r h a v i n g  paper, tips,  because  were o f s m a l l e r  ( G l i m e and V i t t  elongation Plants  of s p o r e l i n g s appeared  stature.  still  shoot  densities.  morphologically  the p l a n t s  have been c a u s e d by  or n u t r i e n t d e f i c i e n c y  i n uncrowded d i s h e s ,  slower,  tips  dioxide  appearance c o u l d  shown  oribatid  i n Tables mite)  stages 5-7.  of the three A l g a l and  contamination  F u n g i were n o t o b s e r v e d .  basal  cell  leaf  A.  length  remained  S i n g l e Leaf  invertebrate  occurred  cultures.  regeneration  Leaf  stable  Regeneration.  in  treatments  (rotifer  and  some s h o o t segment  insertion  angle  i n a l l treatments.  Single  leaves  and  107  Table  Week  ssp.  5 - Developmental stages of s i n g l e regeneration.  leaf  A c t i v i t y Observed  tricostata 2  most p e l l u c i d o r brown,  dead  7  a few r h i z o i d s  from b a s e o f  produced  c o s t a and l i m b i d i a 10  a few s h o o t s p r o d u c e d and  limbidia  from base  on a b a x i a l  14  no c h l o r o p h y l l  left  17  secondary r h i z o i d s  side  of c o s t a of l e a f  i n any l e a v e s produced  by t h e few  shoots developed 19 ssp.  s h o o t s t o 3 mm fryei  2  some r h i z o i d s and  7 10  many  produced  from b a s e o f c o s t a  limbidia leaves  still  a l l l e a v e s brown,  green; l i t t l e dead  activity  seen  108  Table  6 - Developmental  Week  Activity  2  rhizoid  5  active  stages of shoot  Observed  production shoot  t i p regeneration.  prolific  development  and c o n t i n u e d  rhizoid production 9  s h o o t s t o 6 mm, from e a c h  13  secondary  many m u l t i p l e  shoots  fragment  rhizoid  initiation  a t merophytes  109  Table  7 - D e v e l o p m e n t a l s t a g e s of regenerat ion.  Week  ssp.  Activity  1-8  cm  shoot  segment  Observed  tricostata  3  rhizoid  i n i t i a t i o n a t m e r o p h y t e s and  points-of  breakage; a p i c a l  continued; 6  new  shoots  and 9  new  new 10-12  lateral mm  long,  erect-spreading  shoots  to  15 mm  leaf  long;  at  growth  shoots plagiotropism insertion rhizoids  evident  numerous,  some a e r i a l  ssp  13  shoots  etiolated-elongate  19  shoots  > 30  mm  fryei  2  new  3  shoots  to 5  4  shoots  t o 7 mm;  and  spreading  7  branching;  shoots  to  10  rhizoid  i n i t i a t i o n at  merophytes  mm  mm  leaves enlarging; leaf  insertion  orthotropism  evident  11 0  r e g e n e r a t e d p o o r l y and most d i e d few  developed  base  of  the  compared  and p r o d u c e d  costa.  2-10 weeks.  shoots  from  Development was slow  Only  a  the a d a x i a l  and s p a r s e  when  to the other regeneration treatments.  B. long  rhizoids  within  Shoot  included  T i p Regeneration. the a p i c a l  cell,  Shoot the  tips  2-3  mm  surrounding  embryonic  l e a v e s and t h e uppermost mature l e a v e s o f a s h o o t , and were common  in  vigorous  Limbella  regeneration  prolifically secondary tips did  by  rhizoids  single  culture,  The t i p s  exhibited  producing  rhizoids  from m e r o p h y t e s by week much  more r a p i d l y  13.  The  shoot  and p r o l i f i c a l l y  than  leaves.  Shoot  Segment R e g e n e r a t i o n .  segments o f b o t h production. branches  in  litter.  week 2, numerous new s h o o t s by week 5 a n d  regenerated  C.  fragment  taxa e x h i b i t e d  Entire  shoot  both produced  Rhizoidal  and  shoot  segments and segments s t r i p p e d o f  were  regeneration  treatments,  most  and p r o l i f i c a l l y .  rapidly  shoot  vigorous rhizoid  new b r a n c e s  attachents  Live  live  and r h i z o i d s  numerous.  Of  shoots developed  by week 3. the new  three growth  111  2.  Dried  None after  of  the  dried material  15 weeks of c u l t u r e .  fungal  In  green  material  and  week  had  duration  Failure symptomatic extensive  and In  of  aquatic  new  these experiments,  decomposers  of  dead  "destroy...potentially  dying  viable  (1976).  Reciprocal  turned  Cultivation  effect  brown leaf  of  regeneration,  and tips  4%  The  regenerate which  disruption  fungi  never  that plant  material  the dried  1.  appears in drying (Glime  the  to  be  undergo  and  Vitt  invaded s u c c e s s f u l l y  fungi  invade c u l t u r e s  material,  fragments"  of of  d e c o m p o s e r s by week  mosses,  drying  whereas 44%  growth.  to  I t appears or  on  the  growth,  new  irreparable cellular  cultures.  7.11  testing  fungal  material  regenerating  and Ambrose  and  of d r i e d most  usually  regeneration  1, a l t h o u g h a few  of d r y i n g  produced  brown l e a v e s  leaves  exhibited  4.  experiment  d r i e d a t 20°C  tested  by week  d r i e d a t 40°C p r o d u c e d  material  1984).  until  the c a s u a l  temperature  40°C  Most  decomoposers appeared  remained  at  Material  and  as r e p o r t e d  do by  as not  Miller  112  1.  Hawai i a n P l a n t s  After fryei  10 weeks, s s p .  culture  leaf  2.  growth cm)  produced  of e x i s t i n g  branches  was  by  ssp.  shoots.  assume t h e a p p e a r a n c e Chapter after of leaf  8.  which  the  normal  cell  c a s e s the  Shoot  tricostata  horizontal in  line  a l l  t o c.  in typical  and ssp.  was  limited  segments,  shoots  resumed.  t h e same a s  and  caused  branches  f o r a 5 mm  leaf  current,  new  to elongation (to 6  immersion  flagelliform produced  and  lateral shoots  described  segment o f  Except  insertion in typical  in  shoot,  for  leaves  a n g l e and ssp.  to  basal  fryei.  l e a v e s became s q u a r r o s e .  Orientation  monofilament  angle  C u l t i v a t ion  immersion  morphology  l e n g t h s remained  Orientations  ssp.  leaf  to  Initially, of  ssp.  orthotropic  insertion  t h e same as  P r o d u c t i o n o f new  form was  flagelliform  In a few  7.12  This  Leaf  i n Rec i p r o c a l  fryei  minimal.  under  slow.  Plants  10-weeks' e x p o s u r e  grown  numerous t e r e t e ,  l e n g t h s remained  American  Cultivation  plants  long.  Growth r a t e s were  North  After  produced  s h o o t s t o 3 cm  cell  tricostata.  tricostata  conditions  to p l a g i o t r o p i c basal  in Reciprocal  Experiment  exhibited  by  the  shoot  segments s e c u r e d w i t h  were t h e same as t h o s e of u n s e c u r e d produced 60°  plagiotropic  orientation,  cases orthotropic  shoots  whereas s s p .  s h o o t s of n e a r l y  segments,  ranging fryei  vertical  from  exhibited  orientation  1 13  (Fig.  24).  remained  F.  Leaf  Are  Morphological of  cultures  (which  evidence  that  Chapter are used  3 —  basal  leaf  D i f f e r e n c e s Between t h e  all  cultivation  included only two  angle  key of  reliably  to  modification.  ssp.  leaf  cell  length  therefore Stability  Stable?  except  the  ) provide  characters  and  basal  the  taxa.  identified  leaf  the  Other  t o some f o r m of  two  characters  spore strong  cell  in  length  e n v i r o n m e n t a l c o n d i t i o n s and  subject  of  Taxa  tricostata  insertion  distinguish  are  methods  morphological  s t a b l e under v a r i o u s  characters  can  be  morphological environmental  implies that  they  genetically fixed. Based  summarizes  in part  summarizes expression character different a given  stable  Wyatt  characters the  of may  reports  by  reported  of  observed  not  apply  in ssp.  taxa.  the  (1984),  relative  parameters  thought  characters. as  both  i t i s c l e a r that do  in other  Stoneburner  Table  8  stability  of  i n mosses under c u l t i v a t i o n .  morphological  authors,  never b e f o r e  and  physical  character  characters  on  literature  morphological  has  and  stable.  Results  be  angle  DISCUSSION  7.13  are  insertion  to  Because  9  influence the  same  and  unstable  by  rules regarding  stability  of  all  stable  taxa.  tricostata  I t appears that  been measured  to  Table  and  The ssp.  leaf  for i t s s t a b i l t y  two fryei  insertion in  stable may  not angle  comparative  1 14  F i g u r e 24 - S h o o t o r i e n t a t i o n e x p e r i m e n t . S.sp. on l e f t , s s p . f r y e i on r i g h t .  tr icostata  Table 8 - R e l a t i v e s t a b i l i t y of m o r p h o l o g i c a l characters in c u l t i v a t e d b r y o p h y t e s , as r e p o r t e d i n l i t e r a t u r e .  Character  Stable  branching bulbil  pattern  7  morphology  5  gametophyte leaf  morphology  -  alar cells  6,8  apex  7,12  auricles  6  cell  length  -  "  cell  shape  "  cell  wall  cell  width  "  11  12  6,  12  4,12 thickness  -  '  12  -  9,12  size  -  12  number  -  12  anatomy  -  12  length  8  7  width  4  6  decurrency  4,13  10  dentition  4,8,11  1,13  length  -  7,9  -  12  chloroplast " ?!  Unstable  costa  margin  rolling  morphology  7  oil  13  bodies  papilla  number  12  12  1 16  "  "  position  12  size  -  12  plication  -  12  shape  11,12  6,8,12  size  -  6,8,12  spacing  -  12  width  -  9  phyllotaxy  -  12  p r o p a g u l a abundance  -  12,13  protonemal  7  -  pseudostereids  -  2  rhizoid  -  12  "  "  morphololgy  abundance  seta p a p i l l a e  '11  sporophyte morphology  11  stem  -  length  References: Schets  (1) Agnew 1980  1958 (2) F l o r s c h u t z - d e Waard & W o r r e l l -  (3)  Forman  Smith  1977 (6) Lodge  1966  (9)  Zales  1973 (13) Zehr 1980.  Steel  7,12  1964 (4) Koponen  1960 (7)  1978  (10)  Longton Wigh  1967 (5) L e w i s &  1981  (8)  1972 (11) Wigh  Sonessen 1975 (12)  117  Table 9 - P h y s i c a l p a r a m e t e r s i n f l u e n c i n g e x p r e s s i o n of m o r p h o l o g i c a l c h a r a c t e r s , as r e p o r t e d i n l i t e r a t u r e .  Character  leaf "  "  H u m i d i t y , Water  apex awn  length  Temperature  2  -  -  3  3  -  cell  length  5  cell  size  7  wall "  Light  thickness  c h l o r o p l a s t number  2,7  -  2  -  2  -  -  length  4  4  -  thickness  7  -  -  dentition  2  -  length  2  -  7  -  shape  3,4  3,4  3  size  -  -  3  spacing  7  -  -  phyllotaxy  4  4  plant  1  1  size "  costa  "  "  "  II  margin  rolling  morphology  rhizoid  abundance  stem c e n t r a l s t r a n d length  2 7 7  118  Changes  i n humidity  than  caused  greater morphological  variability  changes i n l i g h t ( 6 ) .  References:  (1) Buch  1928  1964  (4) Lodge  1960  1975  (7) Zastrow  (2) Davy de V i r v i l l e (5) S e p p e l t  1934.  &  Selkirk  1927  (3) Forman  1984  (6)  Wigh  119  cultivation  7.14  Are  studies.  Vegetative  Based on Limbella are  are  dried  the  Fragments V i a b l e D i a s p o r e s  experimental  excellent diaspores below  20°C  and  Fragments d r i e d at h i g h e r not  regenerate.  plants  i n the  by  least  observations  G.  Shoot  of  or  remain moist  or  periods  for longer  most  most  rapid  commonly  effective  small  periods  propagules,  in  ( 1 9 5 3 ) , L o n g t o n and  do  from mature  and  broken  size.  only.  prolific  off  aquatic and  are  Single leaves  seem  contrast Greene  to  the  (1979)  and  CONCLUSIONS  provided  karyotype  of  ssp.  (1)  tricostata  live  a n a l y s i s (2) a means t o  and  vegetative  environmentally  Bryum  type.  viable  diaspores,  v i a b l e and  vegetative  their  for  fryei  for  nine  electrophoresis the  diaspores  and  and  viability  of  (3) a means t o  d i f f e r e n c e s between  or g e n e t i c a l l y  Living  ssp.  determine  f r a g m e n t s as  S p o r e s were c l e a r l y  and  material  d e t e r m i n e whether m o r p h o l o g i c a l are  for short  of  r o u t i n e l y broken  the  for their  Gemmell  Cultivation  spores  fragments,  they  fragments  (1980).  SUMMARY AND  months  temperatures  tips,  vegetative  l o n g as  remain d r y  exhibited  propagules  the  Selkirk  as  Limbella?  stream c u r r e n t s , e x h i b i t e d r a p i d r e g e n e r a t i o n  excellent t o be  Shoot  field,  regeneration. plants  evidence,  in  the  two  taxa  p r o d u c e d a s p o r e l i n g of  the  altered.  f r a g m e n t s were a l s o shown t o  efficiency  directly  p r o p o r t i o n a l to  be the  120  size few  of  the  days or Two  angle  fragment. months d i d  not  fragments d r i e d  leaf  i n s e r t i o n and  under a v a r i e t y cultivation,  and  morphological cultivation, plagiotropic consistently These  of  appeared to  orthotropic differences of  distinguish  leaf c e l l  regimes, be  with  produced  whereas  length,  including  genetically  varied  tricostata  orientation,  recognition  basal  cultivation  characters ssp.  longer  than  a  regenerate.  m o r p h o l o g i c a l c h a r a c t e r s which of  support  Vegetative  the  taxa,  were  stable  reciprocal  controlled.  Other  enviroment.  In  shoot  ssp.  g r o w t h of fryei  mostly  produced  shoots. are  ssp.  useful fryei  as  taxonomic a distinct  characters taxon.  which  121  VI11.  A.  TAXONOMIC  TREATMENT  INTRODUCTION Based on t h e f o r e g o i n g e x p e r i m e n t a l  documents t h e t a x o n o m i c d i s t i n c t i o n tricostata  and  subspecies,  citation  descriptions, presented. examined follow  ssp.  follows  o f synonyms,  Frahm  the terminology  Limbella  Hypnum s e c t . 4:37. Sciaromium 1869,  LEAVES  t ricostata ssp.  type  type  and  with a Z e i s s drawing  species  historical  notes are  and  (1984).  (1973)  a key t o  specimens,  specimens  Gradstein  of Flowers  Flora  82:466.  L i m b e l l a CM.,  specimens  Descriptions  Ireland  (1982).  tube.  1896.  Forschungsr.  "Gazelle" Bot.  1889. (Mitt.) Mitt., non Bot.  Leskea 8:7.  PLANTS p l e u r o c a r p o u s , long.  and  chapter  TREATMENT  (CM.) CM.,  Soc.  and  b i b l i o g r a p h i e s and p e r t i n e n t  Citation  TAXONOMIC  between  this  A generic description,  o f synonymy  F i g u r e s were p r e p a r e d  B.  fryei.  evidence,  J.  sect. 1864.  Linn.  Soc.  Bot.  Sc iaromium M i t t . , [= E c h i n o d i u m  12:571. J.  Jur.].  shoots d e n d r o i d or t r a i l i n g ,  smooth, a r r a n g e d  spirally  Linn.  t o 0.5 m  on stem, h o m o m a l l o u s  or  1 22  straight,  to  sometimes w i t h costa  3.5  mm  long,  b i s t r a t o s e regions  strong,  to  u.m t h i c k  or  excurrent,  percurrent  limbate,  limbidia strong,  rhomboidal;  sometimes basal  Note: L i m b e l l a 1983).  cells  CM.  of  Limbella,  non  just  defined  4-6 l a y e r s o f  forked;  margins  Echinodium Jur.  When  the )  acumen,  cells  name  by M i t t e n  Sc i a r o m i u m .  Echinodium  insertion;  usually  quadrate  to  oblong.  should  rank,  at  of  below  upper  Echinodium  Sc iaromium t o g e n e r i c  unistratose,  t o 75 /xm t h i c k a t b a s e , o f 2-4 l a y e r s  mostly  first  genus by two y e a r s . synonym  base,  i s the c o r r e c t generic  Sc i a r o m i u m,  or  sometimes  forked;  of L e s k e a , a c c o m o d a t e d o n l y  a  keeled,  lamina  at  stereids, usually disappearing  submarginal,  as  in  120  stereids,  of  concave,  ( 1864) and  Mitten he  be a s s i g n e d  in  ms.,  as a s e c t i o n  predates  be c o n s e r v e d  species  must  (Buck  that  or submerged  (1869)  elevated  described  (sensu  t o Limbella, the  n e x t a v a i l a b l e name.  KEY  1a.  TO SUBSPECIES  Plants  robust,  branches leaf  or  3-10 cm l o n g ;  cells  usually  leaves  t o (32)60-77 Mm  ecostate;  emergent  Hawaiian  mature s h o o t s  in  Islands  on r o c k , perennial  7-20(50) cm l o n g ,  lower  erect-spreading  35-45°,  long;  inner  rarely or  perichaetial  on s o i l  o r wood,  intermittent  L_j_ t r i c o s t a t a  ssp.  basal bracts  aquatic  streambeds; tricostata  1 23  1b.  Plants  slender,  branches cells  mature  shoots  1.5-4 cm l o n g ;  to  25-50(60)  4-8(13) cm l o n g ,  leaves  m  spreading  long;  inner  costate;  on wood, p e a t , o r b a r k ,  swampy  shrub-carr;  coastal  ( S u l l . ) CM.,  tricostata  3:81.  Sull.,  northwestern  Am.  in  North  dense  America  ssp.  fryei  1896.  Acad.  Arts S c i .  1854.  tricostatum  W i l k e s Musci Sciaromium  (Sull.) Sull., 13.  tricostatum  400.  U.S.  Expl.  Exped.  1859. (Sull.) Mitt.  i n Seem., F l . V i t .  1873.  Hypnodendron  tricostatum  Sauerb., Ber. 78:360. Lectotype  F l o r a 82:466.  Proc.  leaf  p e r i c h a e t i a l bracts  tricostata  LIMBELLA TRICOSTATA  Hypnum  50-60°, b a s a l  rarely aquatic,  (Oregon)  Neckera  lower  That.  St.  i n Jaeg.  Gall.  Naturw.  & Ges.  1877-  1879.  nov.: " U.S.  lecto,  ( S u l l . ) Jaeg.  Ex.  BM i s o l e c t o ,  Exploring  NY  Expedition  Ex.  Wilkes  isolecto).  12 ( ? ) , W20  1838-42," (FH [  United  States  ( ? ) , 1840-1841; "Hab.  forest  at eastern  base o f Mauna Kea, H a w a i i ,  Islands  (Sullivant  1854)].  Sandwich  N o t e : The s p e c i m e n a t FH was c h o s e n a s t h e l e c t o t y p e because depicted personal  i t by  is  clearly  the  same  specimen  as  that  S u l l i v a n t (1859) and i s from S u l l i v a n t ' s  herbarium.  1 24  Hypnum s u b t r i c o s t a t u m C M . , 4: 37.  1889.  nom.  nom.  Type: n o t s e e n . herbarium chosen  "Gazelle" Bot.  nud.  Sc iaromium s u b t r i c o s t a t u m 1898.  Forschungsr.  (CM.) Par., Ind.  Bryol.  1156.  nud. May have been d e s t r o y e d  (Schultze-Motel  because  with  1977).  No  no m a t e r i a l a s s i g n e d  the Miiller lectotype  to this  was  name was  seen.  Limbella  i n t r a l i m b a t a Card.,  Geneve  15-16: 176.  Hypnodendron (Card.)  that  material  Limbella  Broth.,  Jard.  Bot.  1912.  Nat.  P f l . ed.  May be a t PC. institution  assigned  leptolomacea  Sciaromium  Cons.  intralimbatum  Type: n o t s e e n . to  Annuair.  CM.,  leptolomaceum  1925.  Several  loan"requests  not  acknowledged.  were  to this  2, 11: 531.  name was  Flora  sent No  seen.  82: 467.  (CM.) Par., Ind.  1896. Bryol.  1155.  1898. Hypnodendron (3): Lectotype  leptolomaceum  1170.  no  Bescherelle:  Nat.  P f l .  1  1909.  nov.: " I n s u l a e Hawaiicae  [s.n.,  Limbella  (CM.) Broth.,  date], 1900."  limbatula CM.,  Hb.  C.  [Kauai?], Hillebrand Mull.,  (BM).  Flora  82: 467.  1896.  Herb.  Emile  125  Sc i a r o m i u m  1 imbatulum  (CM.)  Par., Ind.  Bryol.  1155.  1898. Hypnodendron  1imbatulum  1170.  (CM.)  B r o t h . , Nat.  P f l . 1(3):  1909.  Type: not s e e n . herbarium chosen  May  have  been  destroyed  (Schultze-Motel  because  1977).  no m a t e r i a l  with No  assigned  the M i i l l e r lectotype  to t h i s  was  name  was  seen.  Sc i a r o m i u m  flagellare  Ital.  Sc i a r o m i u m  1904:  23.  1904.  flexicaule  Broth.  Ital.  Sc i a r o m i u m Soc.  Broth.  1904:  23.  nom.  1904:  nom.  Broth.  23.  Bot.  Soc.  Bot.  Soc.  nud.  i n Lev., B u l l .  1904.  porotrichoides Ital.  i n Lev., B u l l .  nud.  i n Lev., B u l l .  1904.  nom.  nud.  PLANTS y e l l o w - g r e e n t o d a r k g r e e n or b l a c k i s h , sometimes  dendroid,  when submerged  5 m diameter, p r o l i f e r a t i n g 20(50)  cm  long,  (240)300-650(1200) central  strand;  layers  of  clusters 10 cm  round  by m o n o p o d i a l  to  transversely  urn d i a m e t e r , 14-30(48)  cortical  stereids;  layer  rhizoids  a t base of monopodial  long,  usually  sometimes  arising  trailing  elliptic cells  reddish  or  f o r m i n g mats up t o  branching.  20-40 txm w i d e , dark  Bot.  STEMS  7-  in section,  wide, w i t h composed brown,  small  of  i n dense  branches; branches + d i s t a n t ,  from e v e r y f o u r t h m e r o p h y t e .  3-6  3-  LEAVES  1 26  somewhat  c o n t o r t e d when d r y ; Upper b r a n c h  ovate-lanceolate, acuminate  or plane,  serrate  in  upper angle  projecting  cell  of  5-6  third of  urn,  rhomboidal,  5-36x3-18 Mm;  Mm,  at  walls  reduced  to  limbidia  or p o o r l y  elongate-triangular spreading cells  wide,  serrulate 80°;  10-40°,  mostly  incrassate,  percurrent  2-10  cells  30-  apices, to  pitted,  often  the leaves  to  squarrose,  leaves  brownish,  unistratose, wide,  1(2)  serrate  or  angle  of  t o 7 urn l o n g , d e s c r i b i n g  t o l o n g - e x c u r r e n t , depressed obovate i n of 6 l a y e r s  i n lowermost  transversely  7 5 ( 1 0 0 ) x 3 0 - 3 5 um a t b a s e , 1 0 ° ; upper  base,  acuminate-subulate,  cells  submarginal, absent  apices  at  7-77x3-18  produced,  margins  limbidia  acumen,  by  shortly  leaf  sparingly  plicate,  165-225(300)x40~75 Mm a t b a s e ,  below  to  oblong,  Lower stem  section,  well  base  subquadrate  wide-spreading  deltoid,  long  section,  cells  occasionally  developed;  with teeth  in  stereids;  basal  mm,  Mm  submarginal, d i s a p p e a r i n g  of  cells  to  68-120x28-48 Mm  elliptic  layers  auricles  7-20  percurrent  3-3.5x1.4-1.8 mm;  forming  t o base  costa  to  teeth  strong,  limbidia  branches  serrulate  serrulate  upper  0.4-0.8x0;1-0.3  absent  or  wide,  30-50(80)°;  insertion  Flagelliform  cells  in section,  (2)4  cuspidate tips,  mm;  usually  transversely  of  k e e l e d , 2.2-3.5x0.5-1.4  1-(2-) c e l l e d  costa  of s t e r e i d s ;  acumen,  50(75)x21-35  yellow.  with  depressed ovate  j u s t , below  1(2)  60-75°,  ends;  layers  excluding  concave,  unistratose,  describing  excurrent,  sometimes homomallous, e r e c t - s p r e a d i n g 3 0 - 4 5 ° ,  cuspidate,  margins  leaves ovate-oblong to  of  mostly  2(4)  leaves,  elliptic layers  oblong,  in of  of  stereids;  disappearing section,  stereids;  14-60x3-10 ixm; b a s a l  40leaf cell  1 27  oblong-vermicular, pitted,  often  yellow  Dioicous. branches;  absent;  inner  cells  or  bracts  bracts  1-2x0.8-1  mostly  archegonia  transverse  walls  for  mm,  mm  12-20, c o s t a  bracts  broadly or  base,  18-80x5-14  oblong  long,  on  weak  or  s e r r u l a t e at  long,  straight  smooth,  310-360 Mm  1.5-3  annulus  of  at  often  absent,  3-5  mm 2-3  to  hyaline  or p a l e  yellow  apical  cells  stem or  teeth  above,  2-16.  SETA  often  tortuose  endostome  margins  oblong  in  stereids.  deciduous; long;  lanceolate,  aquatic layer  sometimes when  dry; conic-  superficial;  sometimes  cribose  below, p a p i l l o s e a b o v e , t r a b e c u l a t e  hyaline  or  pale  yellow,  with  cm  SPORANGIUM  operculum  stomata  at  often  1.5-3.5  cortical  of  yellow-brown,  inner  incrassate, pitted,  rows  16,  branches;  0.8-1x0.5 mm,  u s u a l l y s h r u n k e n below mouth  mm  acute.  absent;  long,  r o s t r a t e , 0.5-1.5  at  cells,  central strand;  cells,  (xm,  15-30  limbidia  l a y e r s of  apex;  yellow-brown  of  vermiform  flexuose,  diameter, with  base, c r o s s - s t r i o l a t e  back;  s e r r u l a t e at  to cernuous, o b l o n g - c y l i n d r i c , asymmetric,  arcuate,  peristome  subulate,  upper p o r t i o n of  Antheridia or  w i d e , composed of  horizontal  acuminate to  32-1 05(1 50)x3-1 8  cells,  of  limbidia  paraphyses  mostly  upper p o r t i o n absent,  base,  distally,  walls  insertion.  apiculate  at  apex; c e l l s  Mm,  on  weak or  pitted,  45-55;  yellow-brown at  35-45 Mm  long,  ovate-acuminate, concave,  entire  plants,  insertion incrassate,  margins e n t i r e or  incrassate  1-1.5  mm  18-25, c o s t a  1-5(10) b r o w n i s h b a s a l  PERIGONIA bracts  2-2.5  incrassate,  insertion;  except  at  elongate-triangular,  v e r m i f o r m above,  walls  walls  brown.  PERICHAETTA  stem or  squarrose,  30-94x3-14 jum,  high  at  basal  128  membrane, green.  appendiculate. CALYPTRA  deciduous.  SPORES m i n u t e l y p a p i l l o s e , Mm,  4-5  naked, c y l i n d r i c ,  CHROMOSOME NUMBER  14-18 um,  cucullate,  early-  : n=11.  ILLUSTRATIONS: FIGS.  25-28.  Bartram  1933: F i g .  96 a - e .  B r o t h e r u s 1909:  Fig.  824 a - e ; 1924: F i g .  385 a - e .  Christy  1-6.  Sullivant  9B, F i g s .  1-13.  1859: P i .  1980:  Figs.  BIBLIOGRAPHY: Bartram 23;  1933:  1942: 330.  Brotherus  i n L e v i e r 1904:  1909: 1170; 1924: 438; 1925: 531; 1927: 23, 37.  1912:  176.  al.  1973:  291,  Christy 6.  305, 328.  1879:  360.  Miller 37;  132;  Hprmann  238,  1965:  1965: 552.  and M i l l e r  1978: 252.  1896: 466; 1897: 119.  1905:  Crosby  460.  Degener  240.  Jaeger  1967: 120.  and Miller  M i t t e n 1873: 400. Paris  Schultze-Motel  1898:  Sauerbeck 1954:  1155;  1904: 375;  1963: 102.  S m i t h 1967:  Sullivant  1962:  533; 1964: 168, 304, 460; 1967: 372; 1969: 761,  BISH  70.  M u l l e r 1889:  241.  SPECIMENS  et  Hoe 1967: 111, 121; 1974: 5f 1979: 65, 78,  McClure  et a l .  1980.  Cardot  1854: 81; 1859: 13; 1862: 46.  Wijk  et a l . 893.  EXAMINED: ( 4 9 ) , BM ( 1 3 ) , CAS ( 3 ) , COLO ( 5 ) , FH ( 2 5 ) , H-BR ( 1 6 ) ,  129  Figure  25 - S s p . t r i c o s t a t a , dendroid plants. m o n o p o d i a l b r a n c h i n g ( C h r i s t y 5196 ) .  Note  F i g u r e 26 - S s p . t r i c o s t a t a , t r a i l i n g B r a n c h e s s p r e a d t o show b r a n c h i n g p a t t e r n  aquatic plants. ( C h r i s t y 5245  Figure  27  - Ssp.  t r i c o s t a t a , upper F o s b e r g 10233 ).  branch  leaves.  1 32  Figure  28 - Ssp. tricostata. A. - upper l e a f c e l l s . - b a s a l l e a f c e l l s ( b o t h F o s b e r g 10233 ).  B  B,  1 33  M  ( 1 ) , MO  (3),  ( 5 ) , NY  YU  (1).  including herbaria  22  (30), S Note:  with  shown  ( 1 2 ) , UBC  32  (5),  specimens c o l l e c t e d  sporophytes,  will  be  US  (6),  for this  WTU  research,  distributed  to a l l  above.  LIMBELLA TRICOSTATA s s p . stat.  Lectotype  FRYEI  fryei nov.:  Williams,  (NY  ["Oregon,  Coos  northeast  of  ground  lecto,  35:  UBC  County,  on  i n more o r l e s s a t NY  was  comb.  et  (1933)  1933.  Frye  s.n.,  on e a s t  wet  WTU  Arago,  isolecto)  1-1.5  side  pasture"  c h o s e n as t h e  i t i s presumably  Williams  52.  isolecto,  Cape  Charleston  specimen  because  Bryol.  "Cape A r a g o , O r e g o n , "  8.viii.l922  N o t e : The  (Williams) Christy,  nov.  Sc i a r o m i u m  On  ( 9 ) , UC  of  (UBC,  miles highway.  WTU)].  lectotype  t h e same s p e c i m e n upon  based  his  which  description  and  illustration.  PLANTS y e l l o w - g r e e n t o d a r k g r e e n , trailing,  forming  monopodial elliptic  branching.  in section,  small central 3-7  layers  strand; of  sods  to  1m  STEMS 4-13  dendroid  diameter, cm  long,  cortical  stereids;  layer  rhizoids  sometimes  proliferating  round t o  165-480 am d i a m e t e r , 13-24  or  transversely  cells  wide,  25-75 um w i d e , composed dark  a t base of m o n o p o d i a l b r a n c h e s ; b r a n c h e s crowded,  cm  usually  arising  in clusters  with of  r e d d i s h - b r o w n , i n dense  clusters long,  by  from a d j a c e n t  1.5-4  merophytes.  134  LEAVES  somewhat  oblong  to  c o n t o r t e d when d r y ; Upper b r a n c h  ovate-lanceolate,  spreading  cuspidate,  concave,  1.5-3.8x0.1-1  unisratose,  1(2) c e l l s  keeled, wide,  serrate  50-60°,  angle  serrulate  to  cell  percurrent  to shortly  teeth  base  Mm  52-75(100)x35-48 apices,  section,  just  branches  base,  below  mm,  acumen, Mm  deltoid,  plicate,  3.5x1 mm;  margins  2-3(6) c e l l s  stem  of  disappearing section, apices basal  5-6  section,  of  below  upper  mostly  the  leaves  of  oblong,  layers  wide,  at  costa  stereids;  Flagelliform  quadrate  7-40x5-14  to  0.4-  a b s e n t or  40-60°,  to 2.5-  auricles  with teeth  t o 25  percurrent to shortly 115-130x40-48 limbidia  transversely  o f 2-4  of  insertion  forming  t o base  in  3-32x3-14 Min;  reduced  spreading  section,  acumen,  cells  elliptic  elongate-triangular  1(2) c e l l s  in  Leaf  submarginal,  3(4)  yellow.  leaves  60-75x25-28 nm a t b a s e ,  cells  in  to rhomboidal,  angle of 45-80°;  layers  well  10-35°;  base,  or s e r r u l a t e  e x c u r r e n t , d e p r e s s e d obovate base,  strong,  to squarrose, limbidia  unistratose,  wide, s e r r a t e  third  costa  limbidia  acuminate-subulate,  describing  i n upper  usually  ends;  often  produced,  Lower  plane,  60-80°,  transversely  at  pitted,  wide-spreading  developed;  or  o f 5-6 l a y e r s o f s t e r e i d s .  subquadrate  sparingly  occasionally  0.8x0.1-0.3  of  o b l o n g , 7 - 5 0 ( 6 0 ) x 3 - 1 4 Mm, w a l l s  mostly  incrassate,  Mm l o n g ,  at  Upper c e l l s  cell  margins  e x c u r r e n t , depressed obovate  30-55(70)x(8)20-32  stereids.  poorly  projecting  e x c l u d i n g c u s p i d a t e t i p , 15-60°;  disappearing  basal  by  mm;  acuminate  or s e r r u l a t e  14-25 Mm l o n g d e s c r i b i n g  with  leaves ovate-  layers  submarginal, elliptic  in  of s t e r e i d s ;  leaf  to oblong, Mm,  Mm a t  walls  7-36x5-18 M m ; at  insertion  1 35  incrassate,  sparingly  Dioicous. stems  or  inner  bracts  PERICHAETIA  branches.  squarrose, teeth  p i t t e d , often  incrassate,  archegonia  14-30,  elongate-triangular,  1.5-2.5x0.8-1  30-55.  CHROMOSOME NUMBER  mm,  on upper p o r t i o n o f  costate,  limbidia  acuminate  to .  absent;  subulate,  s e r r a t e o r s e r r u l a t e t o base  oblong-vermicular,  often  o r brown.  1.5-2.5 mm l o n g ,  Bracts  t o 18 Mm; c e l l s  mostly  yellow  22-112x3-22  Mm,  with walls  yellow-brown a t i n s e r t i o n , not p i t t e d ;  PERIGONIA  unknown.  SPOROPHYTES  unknown.  : n=11.  ILLUSTRATIONS: •FIGS Fig  29-32. I I , 1.  1933:  Grout Lawton  1934: P i . 1971: P i .  80, F i g s .  1-8.  158, F i g s .  Kawai 1968:  11-17.  Williams  F i g s 1-8.  BIBLIOGRAPHY: Andrews Grout  1945:102.  1934: 266.  1968: 130, 147.  Christy  1980. Crum  Hoe 1979: 65. Lawton  Ireland  1971: 287.  Wijk  & Steere  1958:  1971: 328. et a l .  41.  Kawai  1967:  372.  W i l l i a m s 1933.  SPECIMENS EXAMINED: BISH  ( 5 ) , NY  ( 6 ) , UBC  ( 7 ) , US .(5), WTU  specimens c o l l e c t e d f o r t h i s to the herbaria  research  will  shown above a n d on page 127.  ( 7 ) . N o t e : 11 be  distributed  136  '137  F i g u r e 30  - Ssp.  fryei,  trailing  plants  ( C h r i s t y 5338  ).  138  139  Figure  32  - Ssp. fryei. basal leaf c e l l s  A. - upper l e a f c e l l s . ( b o t h C h r i s t y 5338 ).  B,  1 40  C.  COLLECTIONS OF TAXONOMIC AND Appendix  A  outlines  HISTORICAL  the  historical  tricostata  collections,  which  largely  Hawaiian  bryology.  Hoe  (1979)  collections data 8.1  sequence  parallels  the  described  of  history  the  o f most of t h e s e w o r k e r s , and p r o v i d e d  ssp. of  important  biographical  f o r some. Earliest Collection L.  tricostata  James Macrae it  SIGNIFICANCE  was  appears  i n 1825  decribed  of L i m b e l l a  (St.  by  John  were  reportedly  among o t h e r s  (Mann  1866),  not  described  collected  ssp.  Macrae.  Maio,  Hawaii  ("Ins.  -Junio, ssp.  Vitiensis Hooker Mitten  first  1825";  BM  [Hb.  [Hb.  Hooker  (Mitten  them  1873).  for  from Hooker, by S u l l i v a n t  [Hb.  [Hb.  in  1854.  Mitten])  were p r o b a b l y  that  presumably a f t e r  Sandwich.  Mitten]).  —  Macrae's named  sheets the p l a n t s  Mitten L_j_  and  Macrae.  t i m e i n Seeman's  in reverse  indicating  Macrae  Kea].  T r a c e s of g l u e on t h e —  Candolle, tricostata  Ins.  [Mauna  the'first  before  bryologists.  H o o k e r ] , NY  and NY  L^  by  Macrae's  and De  that  ("Woahoo;  H o o k e r ] , NY  s p e c i m e n s a t BM  cited  Although  European  on Oahu  BM  h e r b a r i u m a t NY,  described  to  first  1979), 29 y e a r s  (1854).  sent  by  h e r b a r i u m a t BM match  specimens  Hoe  Owhyhee, ad montem Kaah  tricostata who  1978;  been c o l l e c t e d  i t i s surprising  tricostata  1825";  Mitten,  have  Sullivant  collections  was  to  tr icostata  by  Flora  in  the  i n the  obtained  the  t r i c o s t a t a had  been  141  8.2  Type C o l l e c t i o n The  or  type  1841,  specimen  presumably  Charles  Pickering, ("Wilkes  deposited  a t FH  H o o k e r ) and  label  NY  of  Ex.  the  The  specimens  base  Small). Ex.  of the  BM  type  Sullivant),  BM  All  Wilkes  in  Brackenridge  1840 or  additional Hawaii,  a r e numbered  specimens  exist,  (isolecto,  herb.  bear  handwritten  1838-1842," t h e o n l y  Expedition's  o f Mauna Kea,  a t FH and  collected  Dunlop  Three  (1854) s u p p l i e d  eastern  was  f o r the U n i t e d S t a t e s E x p l o r i n g  herb.  "U.S.  d a t a g i v e n f o r any  at  William  herb.  Sull.  tricostata  Expedition").  (isolecto,  Sullivant  tricostata  collectors  (lecto,  stating:  1975).  ssp.  by e i t h e r  Expedition  labels  of Neckera  specimens  (Sayre  d a t a : "Hab.  forest  Sandwich  "124," t h e  Islands."  NY  specimen  "W203."  8.3  First  of L.  tricostata  Although a l l relevant  literature  that  Collection  sporophytes  were c o l l e c t e d 1980).  ins.  Maui  label  207  ("In  ad  rupes  ("Maui,  from  by B a l d w i n  105  insula."  J.K.  as e a r l y  as  3 ("Insulae  specimens  The  herb  (Christy  specimens  with  Sandwich,  4500 p.,  [ B I S H ] ) , and  1920.", and  1956).  1875  Baldwin  The  stated  sporophytes  four  D u p l i c a t e from  Small,  (1979)  tricostata,  i n faucibus humidis,  [NY]).  (Miller  Hoe  years,  1875-1900+."  "Plantae Hawaiienses.  purchased Eaton  8  intervening  Baldwin  been d i s t r i b u t e d :  occid.,  [NY]), Baldwin and  by D a v i d Dwight  had  up u n t i l  were unknown f o r s s p .  During the  sporophytes  Sporophytes  1875."  Baldwin  a t NY D.C.  bear  206 the  Eaton,  were p r o b a b l y s e n t t o  specimen  at  BISH  was  1 42  obtained  from B a l d w i n ' s g r a n d d a u g h t e r  i n 1960.  Some  of  the three  is  n o t known by whom, and s p o r o p h y t e s were n e v e r  literature  D.  s p e c i m e n s a t NY were mounted on m i c a  slides,  but  reported  i t  i n the  u n t i l 1980.  RATIONALE FOR TAXONOMIC DISPOSITION OF SUBSPECIES It  term  is  well  "species"  species  (Wagner  in  resolution  to  Richards  and  criteria  bryophytes  (Steere  no f i x e d  1984),  ( G a n d e r s and N a g a t a  definition  1976;  known t h a t  according  studied  1984).  The  has been  However,  e x i s t s f o r the  investigators  specific  1947; A n d e r s o n  1978).  definition  to  reviewed  species  repeatedly  without  1970;  i s general  characters  My  sources  decision  subspecific  rank,  tricostata,  1.  (Szweykowski to  consistently-  include data  Sc i a r o m i u m  i n s t e a d of s p e c i f i c  Similarities  that  from  several  1978).  recognize  was b a s e d on two  Greene  agreement  expressed  which, h o p e f u l l y ,  being  of  1963; C r u n d w e l l there  define  group  question  s p e c i e s must be d e l i m i t e d by a s u i t e o f  biosystematic  must  the  bryophyte  L.  sporophytes  fryei  rank o r a s  a  at the  synonym  of  reasons:  of t h e taxa  are greater  than  their  di f ferences. Sciaromium which  are  fryei consistently exhibits distinct  regardless  of geography.  characters  expressed  from  Limbella  However, by  S_^ f r y e i  the  characters tricostata,  majority  are  shared  of by  143  L.  tricostata,  closely  2.  indicating  Cohesion  w i t h i n the  expressing  be  the  species.  genus,  and  because they  relatively but  ssp.  i s to  any  i s best  rank  as  by  defined  rank and  full  between  them.  not  be  i n so  fryei  and to of  one the  few  characters,  by  subspecific  (1978).  synonymy w o u l d o b s c u r e  Both  the  specific  relationship  SUMMARY  great  fryei  best  Sc iaromium f r y e i and  enough t o w a r r a n t i s recognized  expressing  Ample  type  found  for three  the  i s poorly  exists  documented.  Limbella  rank  for  S.  of  L.  as a s u b s p e c i e s  f o r both  synonyms of  literature  specific  relationship  material exists  l a r g e body of fryei  i t s constituent  member  expressed  Richards  d i f f e r e n c e s between  thus  are  clearly  more s i m i l a r  other  differ  relationship  Sc i a r o m i u m  not  clearly  their  taxa.  could  are  either  tricostata, two  Sc iaromium  than  are  taxa  i n a manner most  r e l a t i o n s h i p s between  tricostata  CONCLUSION AND  fryei.  constructed  another  tr icostata  two  genus.  Because  Limbella  Character  the  related.  A genus s h o u l d  E.  that  ssp.  between  taxa,  but  the types  tricostata.  for ssp.  A  tricostata,  1 44  IX. A.  PHYTOGEOGRAPHY  INTRODUCTION  The  Northern  Hemisphere  characterized  by  Interpretation  of  several  possible  dispersal  events.  This  Limbella fryei  these  chapter  tr icostata  its  a p p r o x i m a t e age  of  are  9.1  Distribution  Hawaiian 33-37; Hoe (Hawaii:  of  1974).  used  of  Although  Bishop & Herbst  and  often  from a b o u t distributed  abundant 110  where  different  w i d e l y as  and  analysis  related  L_;_  of  historical  tr icostata  observations putative  on  both  of  ssp.  long-distance  migratory  explain  the  paths. means  and  i s known from o n l y  the  ssp.  tricostata  tricostata  K a u a i , Maui, M o l o k a i  and  Oahu  rarely  low  as  s.n.,  2216.0, B I S H ) , most p l a n t s  disjunctions.  .modern d i s t r i b u t i o n s  tricostata  ssp.  Hawaii,  to  is  dispersal.  Limbella  tricostata  islands  and  Limbella  PRESENT DISTRIBUTION  by  and  the  tr icostata  Limbella  requires  mechanisms  followed  B.  Limbella  disjunctions  alternatives,  observations  of  transoceanic  describes  ssp.  i n some d e t a i l ,  dispersal, These  broad  distribution  occur  collected  as  11.viii.1966, between  i t occurs,  90  m,  BISH; M o l o k a i :  Hoe  1200-2130  ssp.  m.  Common  tricostata  i s known  herbarium c o l l e c t i o n s ,  duplicate  specimens.  (Figs.  many  Were i t  not  of  these  -44.5  1 46  Figure  34  -  S u r f a c e , and e l e v a t i o n a l d i s t r i b u t i o n t r i c o s t a t a on K a u a i , H a w a i i .  A  A  i  1  A  10 km N Contour interval 300 m  4200 3600 3000 • 'a  g  2400 •  U  «j  1800 •  A  D e g r e e s Longitude  A'  of  ssp.  147  Figure  35 -  S u r f a c e and e l e v a t i o n a l d i s t r i b u t i o n t r i c o s t a t a on M a u i , H a w a i i .  A  Contour interval 300 m 4200 -i 3600 1  B  D e g r e e s Longitude  of  ssp.  1 48  Figure  36 - S u r f a c e a n d e l e v a t i o n a l t r i c o s t a t a on M o l o k a i ,  distribution Hawaii.  MOLOKAI i  .  A  km N Contour interval 300 tn 10  4200 -I 3600 \ 3000 g  2400 •  T-t U  a  «  1800 •  A'  A D e g r e e s Longitude  of s s p .  ,149  Figure  37 -  S u r f a c e and e l e v a t i o n a l d i s t r i b u t i o n t r i c o s t a t a on O a h u , H a w a i i .  of  10 km N Contour interval 300 m  4200 l 3600 3000 2400 i 1800  A 1  D e g r e e s Longitude  ssp.  1 50  for  the  plants'  endemism, t h e  l a r g e s i z e and  latter  having  attracted  over  the y e a r s  (Appendix  A),  well  collected  as  been.  Certain  i t has  localities  native Metrosideros —  repeatedly  i n the  Maui  Alakai Trail  and  last 1967)  and  relatively  have  easy  botanists  (e.g.  Olinda  been  access  t h e most common  different  same a r e a s  not  have  Trail  i n Maui's K i p a h u l u V a l l e y  Limbella  of  the  ssp.  only  fryei  Hoe  to  occur  and  1978  Sphagnum locality  (Hoe  before  explored  t h e moss o c c u r s  in other  unexplored  occurs  above sea  9 km  (Fig.  124°6'W).  mires  m  ssp. of  level  fryei Limbella  tricostata  at Sutton  Lake,  Lane  from  the  n o r t h of F l o r e n c e and 38;  Township  17S,  I accidently  searching (Christy  with  1980).  "Cape A r a g o , O r e g o n " ,  l e c t o t y p e (NY).  slightly  the  never  population  while  as  doubt  areas  known l i v i n g  Ocean  44°3'N,  in  In  ( H a l e a k a l a C r a t e r , M a u i ) have  of L i m b e l l a t r i c o s t a t a  C o u n t y , O r e g o n , 8.9 Pacific  no  at P a l i k u  1946).  on  islands.  Distribution The  and  for  collected  by Hoe  Herbst  John  the  habitat  Pipeline  St.  as  to  15 y e a r s , c o l l e c t i o n s  regions  the  many  might  vid.  and  April  taxon  --  of  many i n d u s t r i o u s b o t a n i s t s  Kauai;  bryologically,  35;  forest  renown as a c e n t e r  on  shown  9.2  the  provide  rain  Limbella  Hawaii's  L a b e l s of t h e  Range  3.5  km  12W,  Sees.  26  and  d i s c o v e r e d the p o p u l a t i o n i n S.D.  Sundberg  Williams identical  coastal  (1933) c i t e d t o the  isolectotypes  more i n f o r m a t i v e : "Coos C o u n t y ,  for  Cape  the  label  (UBC,  type  data  WTU)  of are  1.5 1  Figure  38  Habitat  - Site Sutton  of e x i s t i n g s s p . f r y e i population L a k e , Lane C o u n t y , O r e g o n .  depicted  H e c e t a Head, O r e g o n  7.5'  by  wetland  topographic  symbolsi  Modified  map.  250  m  at  from USGS  1 52  Arago,  1-1.5 m i l e s  highway."  With  northeast the  (Lawton,  general  (Fig.  locality  in  the  approximately Point  comm. Frye's  community  segment  as the midpoint  Range  39).  Pacific  The a r e a  Frye's  facing  indicated  in  1950.  population  still  construction could to  not  Barview  population  unclear  existed  40;  from  Elva  me a d d i t i o n a l  extant.  details  along  an  Dr.  25S,  from  the  o f Coos Bay.  the note  locality  whether  (pers.  (pers.  r e g a r d i n g t h e Barview  was "now the  site  the Limbella  i t was d e s t r o y e d by  whether F r y e  Lawton  Fossil  1; 43°21'N,  a n d 2 km  i n 1950, a n d i f s o , whether he f o u n d still  the type  t h a t he v i s i t e d  Lawton  with c e r t a i n t y  of  the  Township  14W, S e c .  i n 1950, o r whether  of t h e house. recall  Barview,  that the type  back y a r d o f a h o u s e , " s u g g e s t i n g is  showing  places  (Fig.  the outlet  the  It  map  i s 15 m above s e a l e v e l  1950 n o t e  side  o f Cape A r a g o Highway, w i t h  o f t h e segment  Ocean, d i r e c t l y  east  1984)  of  14W, S e c . 36, T o w n s h i p 26S, Range  124°18'W).  on  a t WTU i s a map and n o t e i n  pers.  present  1-km  Charleston  isolectotype  Frye's handwriting locality  of  comm.  actually the ssp. comm.  locality:  In 1956 b e f o r e we went on a c o l l e c t i n g trip to Oregon Dr. F r y e t o l d me a l i t t l e more a b o u t t h e p l a c e where he c o l l e c t e d t h e moss. He s a i d t h e r e was a g r o v e o f t r e e s (a dozen or more, I t h i n k d o u g l a s f i r , ) b e y o n d and on the o p p o s i t e s i d e of t h e r o a d from where he g o t t h e moss. I am s u r e I found t h e p l a c e , the t r e e s were l a r g e r t h a n he h a d d e s c r i b e d a s i t was some y e a r s l a t e r . Some h o u s e s had been b u i l t i n t h e a r e a b u t t h e r e was p l e n t y of u n d i s t u r b e d g r o u n d a n d a s m a l l s t r e a m . I hunted a l l over t h e a r e a without f i n d i n g the moss.  1984) returned fryei  1980) gave  153  Figure  39  - T.C. locality  F r y e ' s map and n o t e d e s c r i b i n g t h e o f Sc iaromium f r y e i W i l l i a m s .  type  1.54  Figure Type north  40 - Type l o c a l i t y o f Sc i a r o m i u m f r y e i B a r v i e w , Coos C o u n t y , O r e g o n . locality  to  Charleston,  1  km  Oregon  was  on e a s t  south  of  7.5'  s i d e of highway,  Fossil  topographic  Point.  between'c.  Modified  map.  250  Williams,  m  1 km  from USGS  155  The 1980  Barview p o p u l a t i o n  (with  S.D.  S u n d b e r g ) and  u n c l e a r which Douglas letter,  as  t h e highway housing had  s e v e r a l such  impounded. were u n a b l e  1983-1984  hookeriana km  - Pyrus  of c o a s t l i n e  California  I  <Fig.  41;  populations,  However,  given  that  --  fact km  distant  populations  exist  discovered  eventually.  Few  such  and  any  habitats.  that  The  t h e many i s l a n d s a t  1973)  may  i n the  area  streams  p l a n t s , there  (almost  mouth  i n the  shrub of  contain promising  any  —  should  Columbia  Refuge, v i d .  sites  for  additional fryei.  have  been  i t is  likely will  dense,  occurring River  be  miry  concentrate  communities"  the  1600  northern  r e g i o n and these  Salix  along  ssp.  populations another  1980).  to  of  good  all  shrub-carrs)  rarity  one  is a  (Christy  finding  the  s i d e of  bryologically-trained  for Limbella  "dense, t a l l the  the  without  Columbia W h i t e - T a i l N a t i o n a l W i l d l i f e Dyrness  with  botanists explore  future search  her  covered  Columbia  somewhere  in  was  British  from  to  was  the a r e a  sites  two  It  west  obnupta  confirming  in  the  is extinct 43  A p p e n d i x N)  searches  referred  three  population  own  along  to r e l o c a t e the  searched  the 80  of  Because  other  carrs,  Most  southwestern  Limbella  documented  occurred  f u s c a - Carex  from  My  were u n s u c c e s s f u l .  d e v e l o p m e n t s , and  chance t h a t the Barview In  1983  stands  in that v i c i n i t y .  been  lost.  f i r g r o v e Lawton had  or c o m m e r c i a l  individuals  remains  on on  (e.g.,  F r a n k l i n and  investigation.  156  157  C.  ORIGINS OF  9.3  The  L.  DISTRIBUTIONS  O r i g i n Of  Hawaiian  1.  R e l a t i o n of L.  The  closest  pachyloma  northern Hawaii  15°  between  the  derived  from  exhibit habitat  relative  c l o s e r to the two  taxa,  South  rarely  branching  mostly  tr icostata  can  L^  branching  and  marked  differentiation  the  morphological  progenitors  --  to  the In  in aquatic in  ESE  leaves  length, forms  the  i s often  tricostata,  be  characters, polymorphic  heterotypical  always L^  of  species  h i g h l y v a r i a b l e , shoots  contrast, in  the  here to  two  is a 16  be  similarities  The  than  in  9000 km  sporophyte  fewer  cm,  serrate  when compared  no  12  m  being  on  L ^ pachyloma  1  with  simple.  size  limbidia,  Limbella  appears to  i s considered  L_j_ p a c h y l o m a .  p l a n t s are  0.5  Based  tricostata  the  exceed  population  Bogota, Colombia,  preferences.  than  tr icostata  equator.  areolation,  Although  longer  nearest  American  climatic  to South American  Limbella  the  Southern Hemisphere s p e c i e s synonyms.  of  Mountains near  similar and  tricostata  (Mont.) C M . ,  Andes  and  Populations  entire,  tricostata leaves  are and  ssp.  sharply  subpinnate.  The  especially  i t s larger  i t s South American c o u n t e r p a r t ,  parallels  differentiation  associated  Hawaii's native vascular  with  flora  — the  from p u t a t i v e c o n t i n e n t a l adaptive  ( C a r l q u i s t 1980).  radiation  of  1 58  2.  Did Limbella  In L.  order to understand  tricostata  L. pachyloma  and  must be  circumsubantarctic, and  D i s p e r s e from S o u t h  Andean  cordillera  the p h y t o g e o g r a p h i c  L_;_ p a c h y l o m a , examined.  occurring  extending northward and  of  p a c h y l o m a , o c c u r s between  Cape  Province.  originated  genus  oceanic the  as  Southern of  flora  islands,  undocumented  1972;  km  p o r t i o n s of South Sporophytes specimens,  but do  Based  my  ssp.  on  of exist  field  tricostata,  are almost  America  and  now  Zanten  universally  its  and  broadest  t h e genus  possibly  Antarcto-Cretaceous  p o p u l a t e d by L ^  and  P6cs  refugia  from  or S o u t h  on  be  during  the  1981).  Barring  the  these  islands,  the  attributed  populations  in  only  to  1500-6000  unglaciated  Africa. are  ( e . g . , Moreau  s.n.,  herbarium  pachyloma,  of s p o r e s over a t  L_;_ p a c h y l o m a  Limbella  Africa's  cool-temperate  ancient  o f L_^ p a c h y l o m a can  presumably  a subspecies  South  Exhibiting  long-distance dispersal  o f open o c e a n ,  in  the  Brazil (Fig.  completely g l a c i a t e d  e x i s t e n c e of g l a c i a l  modern d i s t r i b u t i o n postglacial  van  islands  1972).  were ohce c o m p l e t e l y o r a l m o s t (Moore  is discontinuously  possibly  Hemisphere,  of of  e l e v a t i o n s along  decidedly  the  (Moore  subantarctic  Pleistocene  relationship  distribution  1220-1820 m  is  Hawaii?  subantarctic  upper  nov.,  climates.  part  (Gondwanlandic) The  ( D i x . ) comb.  The  in  and  to  t h e c o a s t a l m o u n t a i n s of SE  capense  distribution  t h e major  at middle  Limbella  to  the  L_;_ p a c h y l o m a  on  42).  restricted  America  rare  in  3.iii.1941,  experience with L^  sporophytes are  herbarium  not  o v e r l o o k e d by c o l l e c t o r s  tricostata  infrequent who  do  (BA))  not  but take  Figure  42  - World d i s t r i b u t i o n  of  Limbella.  160  time  to s c r u t i n i z e  1984).  The  possible  rarity  a  derived  areas  repopulate  15°  1500-6000  diaspores  latitude  ( C o l o m b i a and  tricostata  ssp.  amount of  Hawaii).  tricostata  has  the  of  the  existing  volcanoes  i s l a n d s , now  located  northwest  existing of  Kure.  a n t e c e d e n t s of  the  Hawaiian  would have had  and  the  flora rates  investigators. ancient thought  to  It  Although  some  have  as of  was  be  a  well-  is as  the  at  lower  tricostata  could  the  distributional  extent  to  from  pachyloma,  level.  Kure,  ancient  adaptive  age.  proposed  it  is  oldest  guyots,  islands  more t i m e  to  most  Islands  that  early  land  have  recent  (or the  Stone  the  evolve,  alpine flora  vid.  are  supported  r a d i a t i o n would by  to  Many more  (1983) n o t e d  been assumed Leeward  and  Islands,  the  seamounts o r  (Skottsberg,  phytogeographers  occur,  L.  the m i d - T e r t i a r y  islands) that Hawaii's native evolved  which  Leeward H a w a i i a n  considerably  the  to  of m i d - T e r t i a r y only  Approximately  closest  e a r l y as  If these  t h a n has on  and  islands  d i s p e r s a l o f L_;_ p a c h y l o m a  two  sea  e v o l u t i o n and  slower  submerged  ssp.  H a w a i i a n b i o t a , McKenna  of  been c o n s i d e r a b l y  distant  diverged  eroded to  islands,  ancient  may  subantarctic  f o r such divergence  one  w h i c h have s i n c e m o s t l y  the  Given  that d i s p e r s a l occurred ancient  Vitt  therefore  from S o u t h A m e r i c a .  possible of  km  tricostata  separate  t i m e needed  is  Limbella  the  from l o n g - d i s t a n c e  i t s progenitor) and  and  G l i m e and  dispersal.  i t f o l l o w s that L^  been  9000 km  the  stock  (vid.  sporophytes  collecting,  pachyloma c o u l d  latitude,  (or  of  putative  populations  pachyloma  for long-distance  If L  have  of  artifact  equipped  from  individual  more is  1967). bridges  161  between N o r t h A m e r i c a 1955),  most  exclusively via  agree  o r M a l a y s i a and A u s t r a l a s i a that  the  from a n c e s t r a l  long-distance  immigrants  dispersal  importance of l o n g - d i s t a n c e Hawaiian  flora  was  South  paralleled  by  tortelloides Hawaiian  Andreaea  (S.W.  archipelago  continental in  the  geographical  Antarctic  areas.  origin  The  of  the  (1888), based  position  and  setosus  from  acutifolia  (Vitt  masses  occurring  & Hoe  on  its  is  Zealand f.  Australia,  1980), a l s o  &  New  var.  ( Z a n d e r and Hoe  1979).  may  (Schofield Wils.,  have  o c c u r s i n Hawaii  New  Crosby  sporophytes Hoe  collecting.  Based  on my  are  common and  rather  a l l t h e major  12,000 km  the  elevations  1974)  Crum  also  1972). southern  subantarctic  and Hoe on  1980).  intervening  acutifolia  once  spores  to reach Hawaii. of  U  1854;  appears  field this  Andreaea  rarity  (Sullivant  1965;  (Vitt  could  from  several  likewise  Guinea or the n o r t h e r n Andes), or  a minimum of  i n L_j_ p a c h y l o m a ,  and  known  Z e a l a n d and  high  nearly  fragilis  Hook.  t h e r e but s i n c e e x t i r p a t e d ,  tricostata  colonize  Tortella  (Hedw.)  New  Hook.  (e.g.  to t r a v e l As  distribution  moss  G r e e n e ) Zand.  U n l e s s as y e t u n d i s c o v e r e d a t  1954;  derived  from s o u t h e r n S o u t h A m e r i c a , a l t h o u g h i t  America,  islands  had  flora the  n o t e d by H i l l e b r a n d  isolated  the  dispersed  land  from  American-Hawaiian  Cyrtopus  originated  South  reaching  Gemmell  origin.  The  have  Hawaiian  dispersal  first  Hawaii's permanently igneous  native  (vid.  to  tr icostata  ssp.  Bartram  1933;  be  artifact  an  Miller of  experience i n Hawaii, sporophytes  no doubt  has e n a b l e d t h e s p e c i e s  i s l a n d s having s u i t a b l e  habitats.  to  Once  162  established  from  disseminates  readily  enabling  spores,  1.  tricostata  by v e g e t a t i v e  i t to populate  9.4 The O r i g i n  L^  of North  any r e a c h  American  ssp.  fragmentation of a given  tr icostata  (Chapter  7),  stream.  Populations  A r e Hawai i a n and N o r t h  American  Limbella Closely  Related? All  the  tricostata  having  other  members  them t o be  2.  fryei  closely  from  from H a w a i i .  clearly  ssp.  related,  —  that ssp.  ssp.  tricostata  Morphological  between t h e two t a x a of t h e genus —  indicate  fryei  after  features  long-  (Chapter  f e a t u r e s not e x h i b i t e d  t h a t Lawton  (1971)  considered  conspecific.  Was  L i m b e l l a Capable of L o n g - d i s t a n c e  Hawaii  to North  In t h e H a w a i i a n  Islands, L^  widespread, having  In  America,  North  are  characteristic  tricostata  unknown  of d i s j u n c t  ssp.  tricostata i s  s a t u r a t e d most a v a i l a b l e h a b i t a t s .  however, t h e p o p u l a t i o n s  small, highly localized  sporophytes  D i s p e r s a l from  America?  relatively  very  here  are  derived  dispersal  3) a r e so s i m i l a r by  presented  and s s p .  presumably distance  data  and  unisexual.  of s s p . Male  (Christy  1980).  These  bryophyte  populations  fryei are  plants  and  features are  thought  t o have  163  originated 1972).  from  for  the  occur,  and  ssp.  and  The 25°  Biogeographers  two  of  have  Some of t h e s e f o u n d e r  montane  biota  the P a c i f i c  Northwest  distributions liverwort endemic British  Lepidozia to  Hawaii  Columbia  Miller  Charlotte Frahm  (1980,  Hawaii, C a l i f o r n i a stabilized  concluded recent  sand  that  some  illustrated  and  j e t stream,  parallel  the  reported  (Evans  this  introductions"  3  1968)  locality  distribution from  the  Hawaiian  SSW  coast  once  t h e SE  Alaska-NW  of  the  known o u t s i d e  Sutton  1968).  Miller  from  Hawaii. from  occurring  Lake  i t s c e n t e r of d i s t r i b u t i o n  (of Queen  & Lac.  "probably r e f l e c t s  The  thought  Schofield  Bosch  of  bryophyte  t h e Oregon p o p u l a t i o n  km  Hawaii  Carlquist  hawai i e n s i s  Schofield  only  from  the  by  rainy  was  1914;  Herberta  Oregon —  dunes  4600  origin.  j e t stream t r a c k .  r e p o r t e d Campylopus a u r e u s and  early  of  as s e v e r a l  Lindenb.  later  fid. the  ssp.  t o have r e a c h e d  have r e a c h e d t h e c o o l  reported  1984)  majority  of  b u t was  Islands,  from  were of I n d o - M a l a y s i a n  sandvicensis  identity  a  elements  to  America.  have o c c u r r e d a s  that  j e t stream  archipelago  (1967)  questionable  on  biota  via this  appear  logical  p o p u l a t i o n s a r e s e p a r a t e d by  that  a l s o may  in North  Crum  f o r such d i v e r g e n c e t o  s p e c i e s are thought  It follows  and most  diverged  could  concluded  the N o r t h e r n Hemisphere 87).  has  and  latitude.  a n t e c e d e n t s of t h e H a w a i i a n  (1980:  fryei  to North America  Tertiary.  of open o c e a n  via  i s the s i m p l e s t  t h e amount o f t i m e needed  dispersal  as t h e l a t e  (Schofield  o c c u r r e n c e of L i m b e l l a  t h e e x t e n t t o which  tr icostata,  km  dispersal  Long-distance d i s p e r s a l  explanation Given  long-distance  —  and  relatively in  Indo-  164  Malaysia.  Claopodium  whippleanum  Scapania  ornithopodioides  Lindb.,  Anastrepta  rupestris wider  Hedw.,  and  Crum  that  based  disjuncts  highly  undercollected sporophytes, tricostata  Only American unknown,  disjunction Hoe  (Miller  1980).  1956;  North  Schofield  The s h o r t e s t  ocean.  Miller  of t h e s e  (1956)  America...perhaps  stated  sandvicensis derived  from  from A l a s k a by s u c h m i g r a t i n g b i r d s a s t h e Miller  d i d not  distribution whose  plausible  t o North America of the  North American  present  other  from  a that  Hawaii,  Hawaiian-North  populations are small  localized.  Because L i m b e l l a  dispersal  a l l exhibit a  have d i s p e r s e d  on t h e p a r a l l e l  American and  the n o r t h e r n hemisphere,  Unfortunately,  could  purpurea  increasingly  f o r h i s s t a t e m e n t , a n d i t seems e q u a l l y  taxa  Card.,  showing  respectively  and  from  carried  plover."  rationale  Pleurozia  A n a s t r e p t a o r c a d e n s i s and L e p i d o z i a  i n Hawaii  disseminules golden  Vitt  &  and A n d r e a e a  Northwest  (Hook.)  i s 3000 km o f open  Hawaiian  "arrived  in  Pears.,  Ren.  Schiffn.  although  1972;  disjunctions  these  orcadensis  distributions  Hawaiian-Pacific  (With.)  (Sull.)  -  i n both appears  sporophytes a r e not nearly  every  intermittent to  be  infrequent  population and p e r e n n i a l  perfectly  I  but  only  examined  had  streams  - ssp.  c a p a b l e of l o n g - d i s t a n c e  by means o f s p o r e s . f e m a l e p l a n t s have been p o p u l a t i o n s of ssp. and  dispersal  fryei.  is  fragmentation.  Cultivation  such  are e f f i c i e n t  fragments  found  i n t h e two  Sporophytes  apparently  experiments  limited  known  North  a n d gemmae a r e to vegetative  ( C h a p t e r 7) showed  that  agents of v e g e t a t i v e p r o p a g a t i o n .  165  At  Sutton Lake,  lying  on  detached  the substratum  beaver, Douglas  branches near  or branch  the parent p l a n t s .  s q u i r r e l s and  m i c e were seen  and  r a c c o o n s and mink p r o b a b l y a l s o  wind  and  water,  these  Limbella  dispersal.  colonized  4-5  the B a r v i e w  ha  DISCUSSION  9.5  Evidence Most  logical  occur  vectors. and  that  Zanten  that  o f wind  Zanten  plants,  and  bryophytes  Pocs  because  no  of has  extensive  P6cs,  long-distance species.  discussed,  seem t o be  (1981)  These  as  between  lines  genetically  As of  the o n l y  argue  of  the  against (1944) t o  this  kind  a u t h o r s a l s o argue  distributions,  several not  in Limbella  of c e r t a i n  record  been o v e r e m p h a s i z e d  are  fryei  how  only  and man  species disjunctions. and  with  vectors  i n b r y o p h y t e s , n o t e d by C a i n  for Bryopsida.  discontinuous  not  Together  unknown  distances previously  agents  Van  exists  agree  for generic s i m i l a r i t i e s  certainly  It is  birds,  shrub-carr  effective  f o r the d i s t r i b u t i o n s  the v a s t  t e c t o n i c s has  account  be  seen  was.  hydrochory  in vascular  generating  by van  account  dispersal  transport plate  at Sutton Lake.  involving  zoochory  the  Supporting Long-distance Dispersal  can  presumed  in  occur t h e r e .  may  be  S i g n s of  By v e g e t a t i v e means a l o n e , s s p .  bryogeographers  dispersal In c a s e s  animals  population  D.  segments c a n  a  major  disjunct stated  floras,  d e p l e t e d or  now  in  i t may  i n Chapter  evidence  that  factor  contending that  of  but 5 and  suggest  "evolutionary  166  failures"  (Crum  1972).  I f p l a t e t e c t o n i c s was  broad d i s j u n c t i o n s , d i f f e r e n t i a t i o n probably  would  have  occurred  If  this  were  available. bryophytes dispersal  conspecific i s the  most  likely the  explanation  (e.g.  in  distance  d i s p e r s a l must by  occurrence  constraints climatic  are  carried  westerly the  (24  species)  illustrates equatorial  the  the  and  disjunct  simplest  of  recent  be  and  the  c r o s s i n g both c l i m a t i c  archipelago  Australasia of  extremely  has (47  diaspores  different  P6cs•  that  "showing t h e  long-  circulation  and  of  (3)  Additional  hemispheric  direction  of  hazards  successful  of a i r masses of  Zanten  Given  f r e e z i n g ) and  1981).  review evidence  (1955) map  most  injection  a c h a n c e and  P6cs  (van  the  and  transport  unoccupied h a b i t a t ,  across  capacity  time  1981).  spores  may  prevailing zones  and  number of  moss  i n common w i t h " species)  South  graphically  to c r o s s c l i m a t i c  and  regions.  Despite is  survival  same a u t h o r s  Hawaiian  of  A more  successful  disparate  Gemmell's  s p e c i e s which the America  (2)  a i r streams," thereby  equator.  the  (1)  Zanten  and  many  related?  by  equator  "against  periods  d i s p e r s a l of mosses.  definition  the  the the  remains  improbable mixing  and  at  Nevertheless,  (van  the  zones  patterns  be  s u i t a b l e and  most  so  r a d i a t i o n , d e s i c c a t i o n and  arrival  populations  long  are  closely  for  explanation.  spores  atmosphere  ultra-violet  rare  very  required  i n t o the  a  or  the  why  for long-distance  constraints  diaspores  during  true,  feasible  Wind d i s p e r s a l of  between t h e  responsible  only  the  c o n s t r a i n t s of  logical  way  the  long-distance modern  wind d i s p e r s a l , i t  Hawaiian  bryoflora  167  originated. Hawaiian  Fosberg  vascular  varieties,  flora,  92.4%  to  30,000  ancient come,  years,"  arrival on  then  (1951)  element  i n the founder f l o r a  (1979)  to consist  Following Hoe  species,  (1948)  estimated that  original  founder  (including  species,  one  South American)  years.  Such  morphological  arrival an  the  ssp.  most e a s i l y the  plants  may  no  1979;  have  trace." American) Hoe  were  1979;  Vitt  time  frame  Hoe  endemic. Zander  and  from  225  derived  (Hoe  between  of  American  and Hoe as  1980).  Fosberg,  from  "an a v e r a g e of  every  25,000-35,000  South  Islands  was  had  postglacial little  effective.  differences  and N o r t h A m e r i c a n  by a r e c e n t ,  American  L_j_  pachyloma  i f long-distance dispersal  t h e few m o r p h o l o g i c a l  have  (Hoe  derived  of t h e  may  are  which  have  age  e s t i m a t e d by  of  t h e same  "an  (18.4% f i d .  was  had  --  immense t i m e span c o u l d a c c o u n t f o r t h e marked  Leeward  tricostata  Hoe  Stone  e v e r y 20,000  South  of w h i c h  establishment"  tricostata,  ancient  reasoning,  and  differences  and H a w a i i a n  analysis,  origin  modern moss f l o r a  successful  16.2%  flora  48.1%  7.1%  leaving  and  fid.  others  (including  t h e moss f l o r a  U s i n g Hoe's f i g u r e s and Hawaii's  extinct  native  species  species  "Countless  moss  the  million-year  t o be about  Hawaiian  of 233  Fosberg's  1979)  founder  e s t i m a t e d the American  native  1897  (96% endemic  t h e 5-10  become  that  and e s t a b l i s h m e n t  t o the northwest.  Fosberg  The  of  original  based  flourished,  1979).  estimated  consisting  from 407  successful  islands  1951)  o f w h i c h a r e endemic  1967), had d e r i v e d a v e r a g e of one  (1948,  time  ssp.  fryei  dispersal,  t o one  By  the  of  same  between  Hawaiian  c a n be  explained  after  which  to d i f f e r e n t i a t e .  time  Fosberg  1 68  (1951) l i k e w i s e  argued  that morphological  certain  Hawaiian  American  progenitors evidenced  1.  Are  vascular  plants  L i m b e l l a Spores  similarities  and  their  a relatively  putative  recent  Capable  of  1981)  described  between North  dispersal.  Long-distance  Di s p e r s a l ? Van  Zanten  capabilities bryophyte They  the  green  the green  was  the  initiated  sporangium.  s p o r e s and  to  have  indicating  chloroplast  development  Comparative  study  r u p t u r e of  contrast,  brown  release  from  spores  with  green  mosses,  brown s p o r e s w i t h more x e r o p h y t i c , d r o u g h t - t o l e r a n t  green  were  (1-3  d a y s ) and  spores  viable  appeared  to  apparently  and  were v i a b l e  be  up  t o 48  best  dispersal,  years.  suited  pleurocarpous  Pocs t o  d r o u g h t - i n t o l e r a n t , because  were u n a b l e  such h a z a r d s .  Zanten  conclude they  By  for rapid,  to s u r v i v e the  local  rigors  green  further  p o i n t e d out  that  year. (4-210 spores  dissemination of  and  long-distance  whereas brown s p o r e s a p p a r e n t l y c o u l d  They  1  to germinate  inference,  that  germinated  f o r o n l y 6 days to  were d r o u g h t - t o l e r a n t , s l o w e r  d a y s ) and  wind  l e d van  spores  rapidly Brown  Experiments  and  the  species,  species.  hepatics  in  development  only a f t e r  correlated  spores.  germination  before In  of  hygrophytic and  especially  "brown"  chloroplast  elongation.  relative  dispersal  begun  s t a g e of g e r m i n a t i o n  protonematal  the  long-distance  "green"  spores  color  first  c o a t and  spores  for  species producing  sporangio,  spore  Pocs  ("fitness")  considered  which  and  tolerate  smaller spores  (8-  169  12  jum d i a m e t e r )  than  larger  correlated the  were more r e a d i l y spores  with those  species  resistant  (10-20  of  /um) ,  wider  to d e s i c c a t i o n  time of  liberation,  tr icostata diameter.  ssp.  and  peristome  and  characteristics for  hygrophytic  capability Limbella distance (Chapter  that has  are  tricostata  observations  2.  was  and  that  spores  more  to  Limbella. and 14-18  lodge  on  growth. Zanten  rapid,  local  L.  the These  and  Pocs  dispersive  to long-distance d i s p e r s a l . make  been  of d r o u g h t  i t clearly of  ssp.  dried  within  t h e r e f o r e cannot  size  average  frequently  of  Cultivation  development  c o n s i d e r a b l e degree  and  protonematal  that  had  farther  L_^_ p a c h y l o m a  green  s t o r e d a t 20°C f o r 8 months, showed protonematal  in relation  both  are  ill-suited  which  produced  t h o s e d e s c r i b e d by van  features  7),  of  pleurocarps  dispersal.  spore  distributions,  interest  initiate typify  travel  freezing.  spores  of s s p .  and  that  distributions  tricostata  Spores  aloft  and  s p e c i e s of w i d e r  T h e s e o b s e r v a t i o n s a r e of At  borne  excellent 7  C o u l d L i m b e l l a Have I m m i g r a t e d  Van to  spores  f o r 48 h o u r s  and  germination  and  days,  tolerance. be a p p l i e d  c a p a b l e of l o n g -  tricostata  a t 38°C  Yet  indicating Zanten  and  a  Pocs'  Limbella.  I n t o New  Habitats  Successfully? In  order  immigrant successfully  for  long-distance  diaspores, invade  having a  suitable  dispersal  survived habitat  to  be  effected,  transport, unoccupied  must by  the  170  existing  vegetation.  accepted  observation  the  better-adapted sites'  aquatic  riparian  for  immigrant  competition rocks  and  (Conboy  where  or  Glime  Hawaiian mountain  sparse  s t r e a m s may  f o r new  The native  Oregon  flora  Hawaii. (e.g.,  at  In  cover,  the  surface mat  has  appear  indigenous The  is  an  substratum  suited  of  reduced  shifting  during  of  flooding  saxicolous  even now  habitat the  in  time  many s t r e a m s e x h i b i t  substrata  unoccupied  a l s o may  have "been u n o c c u p i e d  Limbella  diaspore  nearly  identical  and  bare,  and  & Mohr.  might  of  bare  encounter  at  is  The  much  Ochyra  Sutton  Lake  p e a t , and  little  Lake  covered  (Hedw.)  the from  Sutton  Oregon),  rest  praelonga  exposures  immigrant  the  by  introduction  to that  Park, L i n c o l n C i t y ,  (Hedw.) Web.  of by  a  and carr  i t would  competition  from  species.  o c c u r r e n c e of  instead  may  t i m e of  considerable  that  material  aquatic,  and  of K i n d b e r g i a  Climacium dendroides also  drift  in  saxicolous  appearing  e c e s i s c a u s e d by  leaving  habitats  shrub-carrs  discontinuous  occurs  The  one  with  diaspores.  D e v i l s Lake S t a t e  peaty  compete  have been u n o c c u p i e d a t  arrived,  bryophyte  be  generally-  often  hardly  may  continual  The  cannot  most  Limbella,  w a t e r and  the  i s minimal.  species,  1971).  diaspores  available  the  by  of  plant  and  competition  of  i s b a s e d on  immigrant flora,  weedy  because  and  an  habitat  abrasion  Limbella  that  assumption  native  disturbed or  This  of of  submerged  ssp. or  fryei  of  terrestrial  emergent  L ^ p a c h y l o m a and  have been a r e s u l t  on  L^  rocks  tricostata  climatic  and  peat  as  is  ssp.  geologic  or  sand,  the  usual  tricostata,  chance.  Within  171  the  narrow  climatic  strip  of  regime  streams  are  land  along  apparently  restricted  the  best  to  immediate suited  headland  to  were  already  bryophyte  flora  composed of common h o l a r c t i c  region  The  only  strip  today, p r e c l u d i n g  other  would  sheets. with  lotic have  sandy b o t t o m s and  gradient volume  and and  extending  t o 20  climate  abruptly  further  barrier  Schuster from  that as as  caused km  to  forest,  Van species  effectively  as  those  arrival,  penetrate  pattern  of  rocks,  As  just  dune  bearing also  no low-  d i u r n a l changes i n  influences, stated a few  tidewater  i n Chapter km  inland,  the  Limbella  P6cs  2, a  the  protected  b e c a u s e of  impeded  could  stated  disperse  i n e x p o s e d h a b i t a t s , as  hazards  of  native vegetation. gives  spores  ( 1 9 8 1 ) , however,  in sheltered sites species  that  topographically  t o become a i r b o r n e  of  survive  or  Z a n t e n and  of  P6cs' a s s e r t i o n .  tidal  inland.  riparian  diaspores  could  coastal  meandering,  S c h o f i e l d (1984) e m p h a s i z e d  likely  circulation.  they  spores.  narrow  rivers,  to  in  immigration.  (1979) and  closed  no  The  subject  or more  found  immigrant  the  virtually  by  species  aquatic  streams c r o s s i n g the  habitats. are  streams  well-developed  by  the rocky  These  low-gradient,  becomes more e x t r e m e  h a b i t a t s were l e s s wind  banks and  Hawaiian  salinity  and  typically  meandering,  up  a  penetration  rivers  These streams are  resemblance t o the  by  habitats a v a i l a b l e in been  L^_ f r y e i ,  areas.  probably  the  occupied  coastline,  transport The  world  ample c r e d i b i l i t y  t o van  and,  just long upon  dispersal Zanten  and  172  9.6 A l t e r n a t i v e H y p o t h e s e s C o n c e r n i n g D i s p e r s a l  1.  pachyloma  enough  fryei  Hawaiian p l a n t s Pleistocene  Coast  with  h a b i t a t s would  Range  two  Hawaiian  spores:  further  inland,  was  from s s p .  North  (2)  that  not  there.  tricostata,  America  been  prior  to  (1)  the  before  Plio-Pleistocene elevations,  such other  what  the  as  are  the  coastal plants  sea l e v e l  fluctuations  Today's the  Limbella  Pleistocene,  arrival was  time of  established  Range o r o g e n y , when t h e  coastline  during  the  late  was e s t a b l i s h e d d u r i n g t h e  today  older  d i d not  estimated  Coast  founder p o p u l a t i o n on  coastline  population  Pliocene  formed  Pliocene  during  the  founder  the  1981).  inundated for  during  modern  (Baldwin  possibilities  the  uplifted  of P l e i s t o c e n e  glaciation  w e s t e r n C a s c a d e Range  with  to  1973) b u t t h e  have  leaving  along  suggests  t o have o c c u r r e d  slightly  possibly dispersed  after cessation  associated  Miocene  f o r divergence  has d i v e r g e d  and D y r n e s s  form u n t i l  islands  populations  g l a c i a t i o n s , perhaps as e a r l y as the l a t e T e r t i a r y .  Oregon  (Franklin  among p o s t - P l e i s t o c e n e  on t h e s u b a n t a r c t i c  time has e l a p s e d  Because s s p .  The  America  Was D i s p e r s a l P r e g l a c i a l ?  An a b s e n c e o f d i v e r g e n c e of  to North  higher  Pliocene  remained  Coast  Range  m a r i n e t e r r a c e s , and  at  sea  level  during  orogeny. Although fossil  flora  bryophytes of t h i s  t o have p r e v a i l e d .  are  period, Conifers  not  represented  a c o o l i n g and d r y i n g and x e r o p h y t i c  i n the regional trend  Quercus  appears  associations  1 73  replaced late  the  temperate b r o a d - l e a f  Tertiary  Graham  and  1972).  favorable  extinctions  for  Limbella,  but  climate  —  bears  little  resemblance  Migration  from  the  a  be  few  of a t  least  100  deterioration  may  unisexual  or  kilometers  means,  population  at  populations, eventually  "presumably  barely  on"  Sutton  Schuster's formation archegonia the  because  so  of  correct, in  ssp. few  population  theory is  he  modern  c l i m a t e was  coast,  that  e x h i b i t e d by thought  the  that  ability  noted  produced climatic  production, reproduce  forming  solely ssp.  clonal  to evolve," taxa  hardly  favorable sex  of  to  have been  seems  of ssp.  of  be  barely  documented, If  gametangium  apparently  healthy  fryei  suggest  for antheridial  differentiation  which expand  depauperate.  suppression  occurrence  populations  less  E n v i r o n m e n t a l c o n t r o l of  populations  climatic the  does a p p e a r  would  persist  "depauperate p o p u l a t i o n s "  fryei  by  fryei  such  t h a t many  the  vegetative  r e g i o n a l l y , unable to s i g n i f i c a n t l y  Although  Lake  that  lack[ing]  c o n d i t i o n s as  'hang on'  range."  "hanging  He  become e x t i n c t , but  suboptimal  [their]  Lake.  the  inland  early populations  gametangium  populations  Sutton  that  s h o r e l i n e to  postulated  characteristics  seem  immmediate  western Cascade  (1984)  1968;  favorable.  unless  suppress  sterile  vegetative  km,  the  not  remembered  the  of  (Detling  time would  that along  ancient  Schuster  that  that  must  have been more  sporophytes.  the  to  numerous  forest  s h o r e l i n e would have n e c e s s i t a t e d a s t e p w i s e  dispersal  "may  it  sometimes o n l y  where c o n d i t i o n s may  under  were  C l i m a t i c c o n d i t i o n s at  inland  present  Mixed Mesophytic  would  production.  i n gametangia  seems  174  very  unlikely  antheridia  because  and  the  archegonia  early  developmental  a r e thought  stages  t o be i d e n t i c a l  of  (Sharma  1971). Based  on  localities,  numerous Hansen  pollen  (1947)  climate  of t h e n o r t h P a c i f i c  marine,  with l i t t l e  profiles,  concluded coast  variation,  that  from  "The  developed  further  undoubtedly  d i d o c c u r , t h e y were o f  during  While  the dry  coastal  postglacial  has p r o b a b l y been  even  inland.  many  essentially period  climatic  that  fluctuations  insufficient  magnitude  to  c a u s e a s y s t e m a t i c r e s p o n s e by t h e v e g e t a t i o n . "  2.  I_s N o r t h Amer i c a n L i m b e l l a  Paleoendemic derivation, would  status  of  to  be  considered  similarity  to L^ tricostata,  relatively  recent  required  fryei  and i t s  would present  relictual.  indicating  imply  ancient  distribution  I t s morphological  close  relationship  precludes the systematic  and  isolation  f o r paleoendemism.  Did Limbella  Did  ssp.  distance  The  the  to  pattern possibility  present  D i s p e r s e from N o r t h A m e r i c a  tricostata  dispersal  distribution  but  derivation,  3.  1980).  ssp.  systematic i s o l a t i o n ,  have  Paleoendemic?  originate  Hawaii  from  is parallelled  from  North  areal  fryei  cannot  distribution  by l o n g -  America?  by t h e M a d i i n a e  o f s u c h an o r i g i n  disparate  ssp.  t o Hawaii?  be  This  (Carlquist ruled  out,  o f t h e two t a x a  175  argues against population the  fryei  widespread existing  so  would a p p e a r  ssp.  fryei  an i m m i g r a n t  the  fryei  derived  population  Assuming  between  relictual,  having  been  more  sexually  established  a t l e a s t by means o f  arose  t h e y may have been the  possibility similarity  both  ssp.  t h e y may have been  they  Pacific of  of  from  and is  fryei  fryei, considered  long-distance  spores.  Vicariads?  tricostata population,  and s s p . n o t from one  in  the  hemisphere  somewhere  on  the  past. Limbella  mainland  N o r t h w e s t and t h e Andean c o r d i l l e r a . cannot  two t a x a  be  either  o r (2) a p o l y t o p i c  a possibility  ruled  and d i s s i m i l a r i t y  p a c h y l o m a would seem t o r e q u i r e  ssp.  The  seem t o p r e c l u d e  sympatric  now e x t i n c t , e x h i b i t i n g t h e f e a t u r e s and  by  Limbella  southern  sympatric  vicariance the  reproductive.  e x h i b i t s a l l the c h a r a c t e r i s t i c s  from a common a n c e s t r a l  that  of  true,  o f s p o r o p h y t e s would  that  that  were  recently  lack  implies  a n o t h e r , and t h a t  stock,  be  and  If this  A r e Hawaiian and North American  Vicariance  the progenitor  extensive.  and p o s s i b l y  d i s p e r s a l to Hawaii,  4.  L.  to  population  and  u n l i k e l y that  ) would be so r e s t r i c t e d ,  (relatively)  i n the past,  dispersal, any  I t seems h i g h l y  ( ssp. fryei  derivative  ssp.  of  it.  shared origin  but  the  between them a n d  (1) a common  which s t r e t c h e s  by van Z a n t e n and P6cs  out,  The  by s s p . of s s p .  ancestor, tricostata tricostata  the imagination  and  (1981) t o be " n e g l i g i b l e . "  176  5.  Was  North American  Anthropochory agent  speculated  Hawaii.  Frahm's  t h a t L_j_  (1980,  Campylopus a u r e u s  circumspect  in this  tricostata  would  intentionally commercial  too  in  regard.  between  between .them  indicate  they  tricostata prolific  are  closely  herbarium  were  not  causative Lawton  introduced the  were  more  u n l i k e l y that  North America  usual  from  occurrence  connection  confirmed  fragments Limbella  other  means by  L.  either  with  any  which  alien  fryei  ,  areas.  which  _in  but vitro  germinated  buds  ssp.  members  evidence  by  and  of  the  and  genus,  related.  infrequent  protonemata,  diaspores.  in  tricostata  and  and  spores,  vegetative  to  a  America  geographical  ssp.  :  was  been  concerning  brought  i n t o new  as  plants.  I t seems e x t r e m e l y  been  dissimilarity  Fecundity  had  Man?  SUMMARY  Similarities  sporophytes  of  North  hobby e n t e r p r i s e , t h e  CONCLUSION AND  Field  and  unintentionally  introduced  that  opinions  Hawaii  by  frequently  tricostata  1984)  have  or  or  p l a n t s are  E.  invoked  Introduced  to e x p l a i n troublesome d i s t r i b u t i o n s  (1971)  of  is  Limbella  and  showed them is therefore  showed only  rapidly  be  Limbella  undercollected.  cultivation  sporelings. to  that  and  of  ssp.  produced  Cultivation  excellent  of  propagative  amply endowed w i t h  dispersive  mechanisms. Postglacial islands  r e c o l o n i z a t i o n of  i n d i c a t e s that  glaciated  pachyloma  i s capable  circumsubantarctic of  long-distance  1 77  dispersal.  Occurrence  of  ssp.  Hawaiian  i s l a n d s i n d i c a t e s that  islands  by  occupy  short-distance  most  suitable  tolerances. native  dispersal long  ago  of  unisexual  Based tricostata  on  Hemisphere  by  and  slightly  to  and  could  I t has  principal  to c o l o n i z e a l l  a l s o been a b l e  its  entirely  limited  fryei from  from  that  is  distribution  typical  long-distance  evidence,  it  that  ssp.  fryei  dispersal.  Hawaii  could  have o c c u r r e d fryei  in  appears  derived  that  from  ssp.  of  ssp.  Southern  as e a r l y a s spores to  Tertiary.  i s t h e r e f o r e o f more r e c e n t  differentiated  ssp.  v i a long-  from  tricostata late  and  dispersal.  have o c c u r r e d  the  tricostata.  as  of d i s j u n c t  Introduction  i n t r o d u c t i o n of s s p .  the  long-distance  the d i s p e r s a l o c c u r r i n g  The  to  physiological  w i d e l y acknowledged  progenitors,  derived  been a b l e  a l l  from S o u t h e r n H e m i s p h e r e L i m b e l l a  Limbella  ssp.  is  ssp.  on  within  derived  long-distance  mid-Tertiary,  tricostata  it  foregoing  derived  North America  only  the  of  likely  dispersal,  tricostata  the  most  dispersal.  mid-Tertiary.  condition  populations  distance  flora  mainland  as t h e  i t has  habitats  Furthermore,  Hawaiian  tricostata  origin  L. and  178  X.  A.  CONCLUSIONS  REVIEW OF EXPERIMENTAL EVIDENCE Previous  which very  SUMMARY AND  shows similar,  1.  2.  chapters Hawaiian but with  described  and N o r t h  experimental  Differences  1.  Branching  pattern  2.  Leaf  insertion  3.  Stem  diameter.  4.  Branch  (bud f r e q u e n c y ) .  5.  Costa  thickness at i n s e r t i o n .  6.  Basal  leaf  angle.  length.  cell  length.  Karyotype Dif ferences  Chromosome length'.  Isozyme D i f f e r e n c e s  1.  Mobility  evidence  A m e r i c a n L i m b e l l a p l a n t s t o be  the f o l l o w i n g d i f f e r e n c e s :  Morphological  1. 3.  have  of superoxide  dismutase.  1 79  4.  Cultivation  Differences  1.  Geotropism.  2.  Reciprocal that  c u l t i v a t i o n experiments  observed  frequency cell  (2)  leaf  differences leaf  B.  ANSWERS TO  10.1  Are  length  Most Limbella  of are  the  evidence  of  Sc i a r o m i u m  differences recognizing  —  as  fryei  as  are  not  Sc iaromium  and  differences  most  i n d i c a t i v e of  within  the  indicates  genus  exist  outlined a  between  as  the  infraspecific  Limbella.  that  basal  and  thus  between above --  a  Conspecific? H a w a i i a n and  to  of  support L^  and  and  North  recognition  tricostata. specific  subspecies, both  taxa are  Oregon  (1971) v i e w p o i n t .  Hawaiian  enough t o w a r r a n t  fryei  (3)  superoxide  stable,  Lawton's  subspecies  great  (4)  bud  CHAPTER J_  conspecific, confirming  plants  were  North American L i m b e l l a  However, enough d i f f e r e n c e s American  and  (1)  genetically controlled.  QUESTIONS POSED IN  H a w a i i a n and  in  i n s e r t i o n angle  dismutase m o b i l i t y presumably  confirmed  In  similarities  accomodated  interspecific  rank.  The  i n a manner  relationships  180  10.2  What E x p l a i n s In  the Geographical D i s t r i b u t i o n  Chapter  explaining  9,  distribution  d a t e s of m i g r a t i o n geographical, Limbella dispersal,  Hawaii Hawaiian  North  supporting account occurred,  the  The  A  South  certain  is  to  with  other  any  other  degrees  certainty,  populations  bryophyte taxa argue  member  In  speciation  have  to  one  o f t h e genus,  as  t o North America would  but  i n L i m b e l l a , and  linkage. of  of  American-  p l a n t s a r e more s i m i l a r  dispersal  was l i m i t e d  route  t o H a w a i i , and from  t o H a w a i i c o u l d have happened  The Oregon  dispersal  v i a long-distance  American-North  be r e f u t e d  and Oregon  and  certainty,  putative  from S o u t h A m e r i c a  different  dispersal  Tertiary.  C.  America  spores.  in  either  mid-Tertiary,  because  be known w i t h  the Hawaiian-North American  for  A l t h o u g h r o u t e s and  and r e p r o d u c t i v e c o n d i t i o n  i t . Hawaiian than  North  route cannot  distributions  another  the  by  America.  modern d i s t r i b u t i o n s  against  and  have been  dispersal  similar  discussed.  possibilities  and e x p e r i m e n t a l e v i d e n c e s u g g e s t s t h a t  Hawaii  presumably  to  were  geological  would  phytogeographical  p r o b a b l y never w i l l  reached  dispersal  a l l  of t h e Taxa?  order that  to have  early  as  i n the l a t e  remained  small  to vegetative fragmentation.  CONCLUSION This  investigation  tricostata  and  information  about  Lawton's  (1971)  ssp.  was fryei  the f i r s t and  critical  revealed  t h e t a x a t h a n was a v a i l a b l e conclusion  that  study of s s p .  considerably  more  i n the l i t e r a t u r e .  t h e two t a x a were  conspecific  181  was c o n f i r m e d , subspecific  but enough  status.  w h i c h c a n be u s e d genus  Limbella.  t o many o t h e r their  led  Much  additional  to complete  a  were  found  information  much-needed  The methods u s e d  species  and  to  justify  was o b t a i n e d  revision  of  the  i n t h e s t u d y must be a p p l i e d  complexes i f b r y o l o g i s t s  classifications  phytogeographical  differences  attempt  h i s t o r y and e v o l u t i o n a r y  to  are  to  improve  elucidate  processes  that  t o d i f f e r e n t i a t i o n o f t h e B r y o p h y t a a s we know them  the have  today.  182  LITERATURE CITED Aeon, M., J . 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A t a x o n o m i c r e v i s i o n o f t h e genus P h i l o n o t i s f o r N o r t h A m e r i c a , n o r t h of M e x i c o . Ph.D. dissertation, Univ. of B r i t i s h C o l u m b i a , V a n c o u v e r . 166 pp. Z a n d e r , R.H. 1979a. Regenerated herbarium m a t e r i a l f o r b i o s y s t e m a t i c and c y t o l o g i c a l s t u d i e s . Bryol. 82: . 1979b. 643-644.  T e c h n i q u e s f o r s t u d y of P o t t i a c e a e .  . 1982. H e r b a r i u m and c u l t i v a t i o n Beih. Nova H e d w i g i a 71: 127-130.  methods  Taxon  28:  i n mosses.  & W.J. Hoe. 1979. G e o g r a p h i c d i s j u n c t i o n and heterophylly in T o r t e l l a f r a g i l i s var. tortelloides Sarconeurum t o r t e l l o i d e s T~. B r y o l . 82: 84-87. Z a n t e n , B.O. van & T. P6cs. 1981. Distribution of b r y o p h y t e s . Adv. Bryol. 1: 479-562. Zastrow, E. 1934. von Wasser und  323.  E x p e r i m e n t e l l e S t u d i e n iiber d i e Sumpfmoosen. Jena. [not seen]  and  (=  dispersal  Anpassung  Z e h r , D.R. 1980. An a s s e s s m e n t o f v a r i a t i o n i n S c a p a n i a nemorosa and " s e l e c t e d r e l a t e d s p e c i e s ( H e p a t o p h y t a ) . Bryophyt. Biblioth. 15: 1-140. Z i e l i n s k i , R. , W. P r u s - G l o w a c k i & M. Mendelak. 1981. Chemical v a r i a t i o n i n the c e n t r a l European P e l l i a taxa Pp. 25-27 in J . Szweykowski ( e d . ) , New P e r s p e c t i v e s Bryotaxonomy and B r y o g e o g r a p h y . Adam M i c k i e w i c z U n i v . Poznan, P o l a n d . 164 pp.  199  APPENDIX A - SEQUENCE OF SSP. TRICOSTATA COLLECTIONS, AS REPRESENTED IN HERBARIA. Date  Collector  Island  1825 1 840-1841 1851- 1871 1875- 1900 1876 1886 1895 1910 1915- 1919 1919 1 922 1 927-X 1 930 1930 1 931 1 932-1937  J . Macrae U. S. E x p l o r i n g E x p e d i t i o n w. H i l l e b r a n d D. D. B a l d w i n V. Knudsen F. L . C l a r k e A. A. H e l l e r J . F. Rock C. N. F o r b e s J . A. Kusche c. Skottsberg 0. Degener U n i v e r s i t y of Hawaii-Bishop Museum E x p e d i t i o n E . B. B a r t r a m E . C h r i s t o p h e r s o n & B. Hume F. R. F o s b e r g  1933 1 934 1 937 1 938 1 948 1 949 1 952 1 963 1 964 1 964-1965 1 965-X  M. C. N e a l R. D. S v i h l a G. E . O l s o n H a w a i i a n Bog S u r v e y F. F a g e r l i n d & R. S k o t t s b e r g P. R. Needham H. A. M i l l e r F. K. Sparrow H. 0. W h i t t i e r M. R. C r o s b y W. J . Hoe  1 966-1973 1 973-1977 1 975 1 977 1978 1983- 1 984  L. D. K. F. C. J.  1  2  1. 2. 3.  3  E. Bishop R. H e r b s t Nagata R. Warshauer W. S m i t h A. C h r i s t y  H a w a i i , Oahu Hawai i Kauai Maui Kauai Kauai Kauai Maui Maui, Molokai Kauai Hawaii, Kaui H a w a i i , K a u i , M a u i , Oahu Kauai K a u a i , Maui Oahu Hawaii, Kauai, Molokai, Oahu M a u i , Oahu Oahu Maui Kauai Molokai K a u a i , Maui K a u a i , Oahu Kauai Maui K a u a i , Maui, M o l o k a i K a u a i , Maui, M o l o k a i , Oahu Hawaii, Molokai K a u a i , Maui Molokai Hawai i Maui K a u a i , Maui  W. D. B r a c k e n r i d g e , C. P i c k e r i n g . H. S t . J o h n , E . Y. Hosaka, E . Hume, R. I n a f u k u , M a s u h a r a , D. D. M i t c h e l l , W. Wong. L . M. C r a n w e l l , 0. H. S e l l i n g , C. S k o t t s b e r g .  J . C. L i n d s a y  200  APPENDIX B - PLANT SPECIES L I S T , LIMBELLA HABITAT AT SUTTON LAKE, LANE COUNTY, OREGON. BRYOPHYTA Hepaticae F r u l l a n i a tamarisci ssp.nisquallensis (Sull.) Lophocolea heterophylla ( S c h r a d . ) Dum. Metzgeria conjugata Lindb. P e l l i a neesiana (Gott.) Limpr. P o r e l l a n a v i c u l a r i s (Lehm. & L i n d e n b . ) L i n d b . Radula bolanderi Gott. R i c c a r d i a c h a m e d r y f o l i a (With.) G r o l l e R. m u l t i f i d a ( L . ) S. G r a y  Hatt.  Muse i A n t i t r i c h i a curtipendula (Hedw.) B r i d . Dicranum sp. Drepanocladus sp. F i s s i d e n s sp. F o n t i n a l i s sp. Hooker i a l u c e n s (Hedw.) Sm. I s o t h e c ium s t o l o n i ferum B r i d . K i n d b e r g i a p r a e l o n g a (Hedw.) O c h y r a Leucolepis menziesii (Hook.) S t e e r e ex Koch Limbella t r i c o s t a t a ssp. f r y e i (Williams ) Christy N e c k e r a d o u g l a s i i Hook. O r t h o t r i c h u m l y e l l i i Hook. & T a y l . Rhizomnium g l a b r e s c e n s ( K i n d b . ) T. Kop. VASCULAR  PLANTS  Carex obnupta B a i l e y G a u l t h e r i a s h a l l o n Pursh Ledum g l a n d u l o s u m ( P i p e r ) Hitchc. L y s i c h i t u m americanum H u l t e n & S t . John P i c e a s i t c h e n s i s (Bong.) C a r r . Pinus- c o n t o r t a D o u g l . P o l y p o d i u m g l y c y r r h i z a D.C. E a t . Pyrus fusca Raf. S a l i x hookeriana Barratt S p i r a e a d o u g l a s i i Hook. V a c c i n i u m p a r v i f o l i u m Smith  201  APPENDIX C - PLANT AND ANIMAL REMAINS IN FILTRATE FROM WASHINGS OF SSP. FRYEI SPECIMENS. SUTTON LAKE Plant  Remains  bud s c a l e s ( s p e c i e s u n i d e n t i f i e d ) C a r e x o b n u p t a l e a f f r a g m e n t s and s e e d s L i m b e l l a b r a n c h e s and l e a v e s Pinus contorta needles Picea s i t c h e n s i s needles Pseudotsuga m e n z i e s i i needles Pyrus fusca flower p e d i c e l s S a l i x h o o k e r i a n a l e a v e s , bud s c a l e s a n d s e e d s Spiraea douglasii f o l l i c l e s Animal  Remains  b e e t l e s and b e e t l e l a r v a e b i v a l v e s (<4 mm d i a m e t e r ) insect frass i s o p o d (sow bug) mouse d r o p p i n g s o r i b a t i d mites s n a i l s (<3 mm d i a m e t e r , 2 s p e c i e s ) spiders c a d d i s f l y l a r v a l c a s e s (composed o f p l a n t Limbella leaves  material,  some o f  BARVIEW Plant  Remains  g r a s s culms ( s p e c i e s u n i d e n t i f i e d ) F r u l l a n i a fragment f o l i o s e l i c h e n fragment leaves (species u n i d e n t i f i e d ) Rubus p a r v i f l o r u s s e e d s (11) Animal  Remains  caddisfly  larval  case  (composed  of sand g r a i n s ;  fragmented)  APPENDIX D - MORPHOLOGICAL CHARACTERS MEASURED IN SSP. TRICOSTATA AND SSP. F R Y E I . Variable number 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20 21. 22. 23. 24. 25. 26. 27. 28.  Character  measured  shoot l e n g t h (cm) branch " , l o n g e s t (cm) l e a f i n s e r t i o n a n g l e (deg) " l e n g t h , s h o r t e s t (mm) " longest " w i d t h , a t w i d e s t p a r t (mm) " a p i c a l a n g l e , e x c l u d i n g mucro ( d e g ) " t o o t h l e n g t h , l o n g e s t (um) upper l e a f c e l l l e n g t h , s h o r t e s t (um) " " " , longest " " " width, s h o r t e s t " " " , longest " " " length:width basal " " l e n g t h , s h o r t e s t (um) " " " , longest " " width, s h o r t e s t " « " » longest " " length:width c o s t a w i d t h a t i n s e r t i o n (um) limbidia " number o f c e l l l a y e r s o f c o s t a c o s t a t h i c k n e s s (um) number o f c e l l l a y e r s o f l i m b i d i a l i m b i d i a t h i c k n e s s (um) stem d i a m e t e r , l o n g e s t d i m e n s i o n (um) number o f c e l l s i n stem d i a m e t e r w i d t h o f stem c o r t i c a l l a y e r (um) number o f c e l l s i n stem c o r t i c a l l a y e r f  APPENDIX E " BRANCH BUD FREQUENCY DATA MATRIX. TRICOSTATA, 21-37 = SSP. FRYEI.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37  1-20 = SSP.  17, 3 . 1 , 6 . 3 , 2 9 . 6 , 1 . 9 , 1 7 , 6 . 3 , 1 . 9 , 1 0 . 7 , . 6 , 1 . 3 . 3 . 8 , . 6 . 0 , 0 , 0 , 0 , 0 . 0 , 0 , 1 16.2,1.4,0,27,0,9.5.2.7,0,23,5.4.0,9.5,1.4.1.4,1.4,1.4,0,0,0,0,1 2 2 . 7 , 5 . 5 , 9 . 1 , 2 8 . 2 , 0 , 1 0 . 9 , 7 . 3 , 5 . 5 , 6 . 4 , 1.8, 1 . 8 , . 9 , 0 . 0 . 0 , 0 , 0 , 0 . 0 , 0 , 1 18.2,4.6,0,31.9,0,36.4,9.1,0,0,0,0,0,0,0,0,0,0,0.0,0,1 30.3,3.0,18.2,0,21.2,6.1,3,12.1,3,3,0,0,0,0,0,0,0,0,0,1 15.6,3.9,2.6,31.2,1.3,19.5,14.3,1.3,6.5, 1.3,2.6,0,0,0,0,0,0,0,0,0,1 2 8 . 8 . 1 . 3 , 1 . 3 , 18.8,0, 15,6.3,7.5, 1 1 . 3 . 1 . 3 , 6 . 3 , 2 . 5 , 0 . 0 . 0 , 0 . 0 , 0 . 0 . 0 , 1 23.1.3.1,3.1,20,0,24.6,12.3,1.6,6.2,3.1,0,0,1.6,0,0,1.6,0,0,0,0,1 16.3.0,0,30.6,4.1.28.6.6.1,2.4.1,2,6.1.0,0,0,0,0,0,0,0,0,1 17.1,9.8,0,24.4,0,29.3,7.3,0,12.2,0,0,0,0,0,0,0,0,0,0,0,1 6.4,6.4,0,34,0,12.8,34,0,6.4,0.0,0,0,0.0,0,0,0,0.0,1 2 2 . 9 , 2 . 1,0, 1 6 . 7 , 0 , 1 0 . 4 , 0 , 0 , 1 0 . 4 , 0 . 1 2 . 5 . 6 . 3 , 4 . 2 , 6 . 3 , 2 . 1 , 0 , 4 . 2 , 0 , 2 . 1,0.1 15.5,0,0,29.3.0,10.4,15.6,0.10.4,1.7,8.6,3.5,1.7,0,0,3.5,0,0,0.0,1 16.1,1.8,0.33.9,0,19.6,17.9,0,7.1,0,0,1.8,1.8,0.0,0,0,0,0,0,1 22.6,3.2,0,19.4,0,16.1,6.5,3.2,9.7,6.5,9.7,3.2,0.0,0,0,0,0,0,0,1 10,0,10,10,0,0,10,0,40,0,20,0,0,0.0,0,0.0,0,0,1 7,4.7,0,30.2,2.3,4.7,11.6,0,11.6,4.7,4.7,9.3,2.3,2.3,2.3,0,0.2.3,0,0,1 9.3,4.7,0,27.9,0,7,16.3,2.3.16.3,0,9.3,4.7,2.3,0,0.0,0,0,0,0.1 13,0.0,26.1,0,8.7,21.8,0.8.7,0.21.8,0.0,0.0,0,0,0,0,0,1 10.8,1.6,1.6.20,0,15.4,9.2,0,21.5,3.1,6.2,6.2,1.6,0,0,1.6,1.6,0,0,0,1 21.9,8.8,3.8,21,1.9,24.8,4.8,1,8.8,1.9,1,1,0,0,0,0,0,0,0,0.2 35.3,5.9,5.9,15.7,2,29.4,0,0,3.9,2,0,0,0,0,0,0,0,0,0,0,2 27.5, 17.5,17.5,12.5,0,22.5,2.5,0,0,0,0.0,0,0,0,0,0.0,0,0.2 25,5.8.9.6,15.4,0,21.2,9.6,0,9.6,3.9,0,0,0,0,0,0,0,0.0,0,2 44,8.8,5.9,5.9,0,26.5.2.9,0,0,0,5.9,0,0.0.0,0,0,0.0,0,2 29.4,11.8,0,29.4,0,17.7,0,0,5.9,0,0,5.9,0,0,0,0,0.0,0,0.2 37.5,6.3,6.3,18.8,0,31.3,0,0,0,0,0,0,0,0,0,0,0,0,0,0,2 29.4,5.9,5.9,17.7,0,41.2,0,0,0,0.0,0,0,0,0.0.0,0,0,0,2 33.3,6.7,6.7,13.3,6.7,33.3,0,0,0,0,0,0,0,0,0,0.0,0.0,0,2 20,0,0,30,10,30,0,0,10,0,0,0,0.0,0,0,0,0.0.0,2 21.4,0,0,21.4,0,57.1.0,0,0,0,0,0,0,0,0,0,0,0,0,0,2 35.7,0,7.1,14.3,0,42.9,0,0,0,0,0,0,0,0,0,0,0,0,0,0,2 23.7, 15.3,11.9,18.6,0,10.2,6.8,0.11.9,0.0.0.1.7,0,0,0,0,0,0.0,2 4 2 . 6 , 6 . 4 , 10.6, 1 4 . 9 , 0 , 1 4 . 9 , 2 . 1,2. 1 , 4 . 3 , 2 . 1 , 0 , 0 , 0 , 0 , 0 , 0 , 0 , 0 . 0 , 0 , 2 28.3,13.3,16.7,13.3,1.7,16.7,3.3,1.7,5,0,0,0,0.0,0,0,0,0.0,0.2 28,12,6,20,0,20,8,0,6,0,0,0,0.0,0.0,0,0,0,0,2 35.2,8,9.1,13.6,0,15.9,4.6,2.3.9.1,2.3,0,0,0,0,0,0,0,0,0,0,2  APPENDIX  F - DISCRIMINANT AND MAHALANOBIS DISTANCE BRANCH BUD FREQUENCY DATA.  <SEPARATE OPTIONS=DISTANCE V A R = 1 - 2 0 C A S E S = 1 - 3 7 SEPARATION OF STRATA CASES=CASE#:1-37  DISCRIMINANT  FUNCTIONS  SIGNIF  38.614 9.2239 5.5674 16.156 37)  .0000 .0047 .0246 . 0003  F-STAT  SIGNIF  3.1455 .37539 2.7408 .24276 .94346 .38514 2.6957 .76989 1.9544 2.3896 1.4418 1.4854 2.2165 .60693 2.2948 -O.  .0860 .5445 .1079 .6257 .3389 .5394 .1107 .7833 .1720 .1323 .2389 .2321 .1466 .9384 .1399 1.0000  CONSTANT 1 . V1 3.V3 5 . V5 8.V8 (N= 37 OUT OF NOT  INCLUDED  2.V2 4.V4 6 . V6 7 . V7 9.V9 1 0 . V10 1 1 . V11 1 2 . V12 1 3 . V13 1 4 . V14 1 5 . V15 1 6 . V16 1 7 . V17 18. V18 1 9 . V19 2 0 . V20  MAHALANOBIS DISTANCES V21 (1) (2)  STEP 0 1 2 3 4  <FINISH>  -2  (1) -3.6296 .43618 .11955 .51792 -.41251 20  BETWEEN STRATA  D-SOUARE  F-STAT  SIGNIF  11.807  24.799  .OOOO  FORWARD SEPARATION INCLUDE INCLUDE INCLUDE INCLUDE INCLUDE  -1  NO V A R I A B L E S 1.V1 8 . V8 3 . V3 5.V5  STRAT=V21>  STEP= 4  5.V5)  F-STAT  VARIABLE  N  (INCLUDE  ANALYSIS  . F-STAT  SIGNIF  35.542 14.321 7.2335 5.5674  .0000 .0006 .0111 .0246  (2) -16.942 .94163 .5988 1 1 . 3064 -2.0729 17  1 5 , 5 . 5 , 4 0 , 3 , 3 , . 8 , 5 0 , 14, 1 0 , 2 5 , 5 , 1 0 , 2 . 3 , 2 5 , 7 7 , 5 , 1 0 , 6 . 8 , 1 0 0 , 4 0 , 5 , 4 0 , 4 , 2 5 , 4 2 5 , 1 8 , 3 0 , 3 1 0 . 6 . 4 5 . 3 . 3 . 2 , . 8 , 4 0 , 1 4 . 7 , 3 2 , 7 , 1 4 , 1 . 9 , 2 2 , 7 0 , 3 . 8 , 8 . 4 , 9 0 , 6 0 , 5 , 4 4 , 4 , 3 2 , 3 8 5 , 18,20,3 1 2 , 3 , 3 0 , 3 , 3 . 2 , 1 , 4 5 , 2 1 , 8 , 2 0 , 4 , 12, 1 . 8 , 1 4 , 5 6 , 4 , 1 0 , 5 , 1 0 0 , 5 0 . 6 , 4 0 , 4 , 3 0 , 4 5 0 . 2 4 , 3 0 , 4 17,4,40.2.5,2.8, .6.20.7,7.18,3.7,2.5.7.32.3.7,3.9.75,30.5.36,4,25,260.18,45,4 18.5,6. 4 5 , 2 . 5 , 2 . 8 , . 8 , 4 0 , 1 4 , 1 0 , 2 8 , 3 , 12,2.5, 14,67,5,7,6.8,75,45.5,40,4,30,300,16,20,3 1 4 . 5 , 6 . 5 , 3 0 , 3 , 3 . 2 , 1.1,30, 1 0 , 7 , 3 6 , 3 , 1 2 , 2 . 9 , 2 2 , 6 0 , 3 . 12,5.5, 1 2 0 , 7 5 , 5 , 4 5 , 4 . 3 0 , 6 0 0 . 15,30. 15.9.5.40.3.3, .8,30,14,10,36,5, 10,3.1,18,74,3,8,8.4,95,55,6,48,4,35,500,25,30.3.5 1 1 , 3 , 4 0 , 2 . 5 , 2 . 8 , . 8 , 5 0 , 1 4 , 1 0 , 2 2 , 5 , 1 0 , 2 . 1 . 1 8 , 6 0 , 5 , 1 0 , 5 . 2 , 9 0 . 4 5 , 6 , 4 0 , 4 . 2 8 . 3 3 0 , 19,20, 3 7 , 4 , 4 5 , 2 , 2 . 5 , . 5 , 6 8 , 1 4 , 7 , 2 5 , 5 , 1 0 , 2 . 1 , 14, 4 5 , 5 , 1 0 , 3 . 9 , 6 8 , 5 5 , 5 , 3 0 , 4 , 2 5 , 2 4 0 , 1 4 , 2 5 , 3 15,6.5, 30,2.5,3,1,55, 14,8,22,5,14,1.6, 18,56,5,8,5.7,110,45,5,44,3,25,450,24,30,4 2 0 , 5 , 4 0 , 3 , 3 . 5 , 1 . 4 , 8 0 , 1 8 , 7 , 2 5 , 5 , 18, 1 . 4 , 1 8 , 6 6 , 5 , 1 8 , 3 . 7 , 8 5 , 4 5 , 5 , 2 8 , 2 , 2 1 , 4 1 5 , 2 0 , 2 0 , 3 1 3 , 3 , 3 5 , 2 . 2 , 2 , 2 , . 6 , 3 0 , 1 4 , 5 , 1 8 , 5 , 9 , 1 . 6 , 1 0 , 4 0 , 3 , 7 , 5 , 9 0 , 4 5 . 6 , 4 5 , 4 , 3 0 , 3 0 0 , 18,30,4 5.5,3.5, 5 0 , 3 , 3 . 8 , . 6 , 1 5 , 1 8 , 10,32,5,10,2.8,14,60,5.14,3.9,90.50,6,48,4,30,400,20.25,4 5, 1 . 5 , 5 0 , 2 . 8 , 3 . 5 , . 8 , 2 5 , 1 8 , 7 , 3 0 , 5 , 1 0 , 2 . 5 , 8 , 4 0 , 5 , 8 , 3 . 7 , 9 0 , 4 5 , 6 , 4 8 , 4 , 3 2 , 2 2 0 , 1 8 , 3 0 , 3 6 , 3 . 5 0 , 3 , 3 , 1,25, 1 6 , 7 , 2 2 , 5 , 1 2 , 1 . 7 , 1 0 , 40,3,14,2.9,1O0.7O,5,35,4,30.500,24,30, 3 9 . 5 , 4 , 6 0 , 1 . 9 , 2 . 5 , . 5 , 3 0 , 1 5 , 7 , 1 8 , 5 , 10, 1 . 7 , 7 , 2 7 , 3 , 1 0 , 2 . 6 , 7 5 , 5 0 , 5 , 3 8 , 4 , 1 9 , 4 0 0 , 2 2 , 2 5 , 4 6,4,60, 1.8,2.8, .8,55,25,7, 18,5, 10,1.7,7,36,7, 10,2.5,75,50,5,44,4,26,330,19,30,3 6,3,60,3,3, .8,60,22,7,18,5,10,1.7,8,40,3,10,3.7,75,55,6,40,4,30,420.23,30,4 5 , 2 , 5 0 , 2 , 2 . 2 , . 6 , 4 0 , 1 8 , 7 , 2 5 , 3 , 1 0 , 2 . 3 , 8 , 3 0 , 3 , 1 0 , 2 . 9 , 7 5 , 5 5 . 6 , 3 8 , 4 , 2 7 , 3 0 0 , 18,30,4 4.5.2.5,60,2.6,2.8, . 8 , 6 0 , 2 3 , 7 , 2 2 , 5 , 10,1.9,8,36,5,10,2.9,90,55,6,46,4,8,285, 16,30,3 4, 1 . 5 , 5 0 , 2 . 6 , 3 , . 6 , 4 0 , 1 4 , 7 , 2 2 , 5 , 14,1 . 5 , 7 , 3 2 , 5 , 8 , 3 , 8 3 , 4 5 , 6 . 4 3 , 4 . 2 8 , 2 7 0 , 2 0 , 3 0 , 3 1 0 , 2 . 5 , 6 0 , 1 . 5 , 2 , . 4 , 6 0 , 1 4 , 7 , 1 8 , 5 , 1 4 , 1 . 3 , 1 0 , 2 5 , 5 , 1 0 , 2 . 3 , 6 0 , 3 0 , 5 , 3 3 , 3 , 2 0 , 165, 1 3 , 3 0 , 3 1 2 . 5 , 2 , 5 0 , 2 . 5 , 3 , . 9 . 4 8 , 2 5 , 3 , 2 5 . 7 , 14, 1 . 3 , 8 . 5 2 , 3 , 1 0 , 4 . 6 , 5 2 , 4 8 , 5 , 4 0 , 4 , 3 0 , 3 3 0 , 2 0 , 3 0 , 4 8 , 2 , 5 5 , 2 , 2 . 5 , . 6 , 5 5 , 1 4 , 7 , 1 8 , 5 , 10, 1 . 7 , 9 , 3 0 , 5 , 9 , 2 . 8 , 7 5 , 4 5 , 6 , 3 8 , 3 , 1 4 , 3 1 5 , 2 2 , 4 0 , 4  206  APPENDIX H - DISCRIMINANT AND MAHALANOBIS DISTANCE ANALYSIS OF PHENETIC DATA.  DISCRIMINANT  FUNCTIONS  VARIABLE  2 3 18 22 25 N  CONSTANT . V2 . V3 . V18 . V22 . V25 (N= 24 OUT NOT  SIGNIF  7.3533 49.916 4.4482 10.085 7.8752 OF 2 4 )  .0143 .0000 .0492 .0052 .0117  1 .. V1 4. . V4 5 . . V5 6 . V6 7 . V7 8 .V8 9 . V9 10 . V 1 0 11 . V1 1 12 . V12 13,. V13 14 ,. V14 15, . V15 16 . V16 17. . V17 19,. V19 2 0 . . V20 21 ,. V21 23 . V23 24 . V24 26 . V26 27 . V27 28 . V28  F-STAT .52577 .32311 .63873 .29234 .94644 .34626 .42696 3.2223 .17066 3.2064 .38342 .37369 .16351 .44891 .25393 1.4643 .88095 .30836 .98508 2.0321 .94402 .50394 .45391  MAHALANOBIS DISTANCES  STEP O 1 2 3 4 5  18.V18)  F-STAT  INCLUDED  V29 (1)  (INCLUDE  STEP= 5 (1) -126.94 -7.5628 3.6675 -2.9503 2.7462 .15289 12  SIGNIF -2 -1 -1 -2  .9430 .8595 .8035 .9575 . 3443 .9854 .5222 .0904 .6847 .0912 .9846 .5491 .8998 .5119 .8753 .2428 .7702 . 5859 . 3349 .1721 . 3449 . 4874 .5095  -3  -3 -1 -1 -1  BETWEEN STRATA  D.-SOUARE  F - STAT  SIGNIF  (2)  40.086  39 . 3 5 7  .0000  FORWARD  SEPARATION  INCLUDE INCLUDE INCLUDE INCLUDE INCLUDE INCLUDE  NO V A R I A B L E S 3.V3 2.V2 25.V25 22.V22 18.V18  F-STAT  .  54.498 8.8556 10.155 5.3903 4.4482  SIGNIF  .0000 .0072 .0046 .0315 .0492  (2) -226.65 -11 .357 5.1518 -6.3923 3.7539 .20870 12  207  APPENDIX I_ - CORRELATION MATRIX OF PHENETIC DATA.  PO Kl M M CO  < t»  <  <  tn O I  a  Ul O I Ul <  M 10 O I  to  U M  to  O I O  <  <  <  <  < u O  Ul 10 10 M -4 01 Ul it  <  M 09  <  <  u  <  M M  -*  (0 a (0  <  <  tn  <  ts  < tn  Ul Ul <  u  <  <  o o U l O— I u— o U l o M & C O o u u o o t n t n C O O tn to <— o to 01 01 o 09 o Mt n o b fa o fa t o CO C O t -J b C o fa c n < o —O. •o cn — 01 r fa o cn .Ul n O o 91 • o cn t 0o9 a o o O o u i f a O t o -»J u o O O u* t oKJ O -J cn u O O O O to to w — to — — w — — — CJ o U o> oo • o UO 09 to u O o O I !* fa o OB fa fa o fa O -* fa o cn to fa to < M  O IUl U lc • oo  CD  —. fa  — Ul  —  Ul •  -J  Ul  u —  IO  Ul  —•  M  CD  .&  Ul  GO  U)  fa  Ul  u  -»  O  <  CJ  -Nl  1  U) (0  1  Ul  CJ  Ul Ul  < wb ro O o O •U l tU —•l — C J CD — oO O tC -fa  fO  O  u Ul  CO  -4  u  m  CJ  c n t—ol tfa oo fa o U o o tC oOl U • o CD O ~4 to o o _^  —  < to O — tocJfaO t too-w M— -4fa01 O — CO O u O CcOn tfan —co U—•O o -* < M i M f uo •too tn O Ul)o t>n o o o •o U tn ao 03 o o o O &  <O I i O I£k --u b O I O II CO CO O tn t oo  *J  u  b  o o o  m  < 03  CO  O CN  t 2 3 4 5 6 7 8 9  II  «D * I I  -I • Xo t i- • o i-1 z w u .J [14 ta o • m u! o c 2 in! n O c K  II  CJ  I D r» > i t—<I EH ID D < W  w  « <EH  p <EH < in o W CJ EH  D t-H  tJ EH t mo ft < to i 10 O  M n z w  a.  <  IO  1 1  13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55  3 , 16 . 48 2 . 3 , 12.64 1 . 7 . 9.34 1 . 7 . 9 34 1 6 , 8 . 79 1 5 8.24 8 . 24 1 5 8.24 1 5 1 3 7 . 14 7.14 1 3 0 .8 4 .40 2 5 15.06 12 .05 2 O 1 7 10. 24 1 7 10.24 1 7 10. 24 1 5 9.04 7.83 1 3 1 3 7.83 7 .83 1 3 0 7 4 . 23 O 9 5 . 42 12.76 2 5 11.74 2 3 11 .23 2 2 2 0 10. 20 2 0 10. 20 10. 20 2 O 1 7 8.67 6.63 1 3 1 2 6 . 12 1 2 6.12 1 2 6 . 12 2 8 15 . 39 13.74 2 5 1 7 9.34 1 7. 9 . 3 4 1 6 , 8 . 79 8 . 79 1 6 1 6 8 . 79 7 .69 1 4 1 3 7 . 14 1 1. 6 . 0 4 O 9| 4 . 9 5 2 2 . 12 .43 2 2 . 12 .43 2 2 . 12.43 2 0 , 11.3 1 7. 9.61 1 5, 8.48 1 3, 7.35 1 3. 7.35 1 3. 7.35 1 2. 6.78 0 8, 4.52  56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 1 10  2 1 . 12 . 28 2 0 . 11 . 7 0 1 8 . 10. 53 1 7. 9 . 9 4 1 6 , 9 . 37 1 . 5 . 8 . 77 1 5, 8.77 1 . 4 . 8 . 19 1 4 , 8 . 19 1 2, 7 . 0 2 O 9- 5 . 2 6 2 . 5 . 15.82 1 B. 1 1 . 39 1.5. .9.49 1 . 5 . 9.49 1 4. 8.86 1 3. 8.23 1 . 3 . 8 . 23 1 3. 8.23 1 . 3 . 8.23 1 1 . 6.96 0 8, 5.06 1 7. 11.33 1 7, 11.33 1 7 , 1 1 . 33 1 5. 10.0 1 3 . 8 . 67 1 3. 8.67 1 3 . 8 .67 1 2. 8 . 0 1 2. 8 . 0 1 1, 7 . 33 1 0 . 6 .67 2 5 . 13 .02 2 3 . 1 1 .98 2 0 . 10.42 2 0 , 10.42 1 9 . 9 . 38 1 7. 8 . 8 5 1 7. 8 . 8 5 1 4 . 7 .29 1 3. 6.77 1 3. 6.77 1 2, 6 . 2 5 2 0 . 12.9 2 0 . 12.9 1 7. 10.97 1 5. 9.68 1 3 , 8 .39 1 3 . 8 . 39 1 3 . 8 . 39 1 3 . 8 . 39 1 1, 7. 10 1 1. 7 . 10 0 9. 5.81  1 1 1 1 12 1 13 1 14 115 1 16 1 17 1 18 1 19 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165  2 5 . 14 .88 2 • 1 . 12.5 2 . 1 . 12.5 1 9 . 11.31 1 6, 9 . 5 2 1 • 3, 7.74 1 • 3 . 7.74 1 • 2, 7. 14 1 2 , 7 . 14 0 • 9, 5.36 0 •7, 4 . 1 7 2 • 3 , 12.5 1 . 9 , 10.33 1 8, 9.78 1 7. 9 24 1 7, 9 . 2 4 1 6, 8.70 1 5 , 8 . 15 1 5 , 8 . 15 1 6, 8.70 1 5 , 8. 15 1 3 , 7 .07 2 7, 13.5 2 7, 13.5 2 1 , 10.5 1 8 , 9.0 1 7, 8 . 5 1 7. 8 . 5 1 5, 7.5 1 5, 7 . 5 1 5, 7.5 1 5. 7.5 1 3, 6.5 2 3 , 13.86 1 8 . 10.84 1 7, 10.24 1 7. 10.24 1 6 . 9 .64 1 5, 9.04 1 5, 9.04 1 3 , 7 .83 1 3 , 7 .83 1 0, 6.02 0 9, 5.42 2 7, 15.98 2. 7. 15.98 1 .7 , 10.06 1 .5 , 8 . 8 8 1 5 , B.BB 1 4. 8.28 1 .3 . 7 . 6 9 1 .3 . 7 . 6 9 1 .2, 7. 1 0 . 8. 4 . 7 3 0 . 8, 4 . 73  166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 2 IO 211 212 213 214 215 216 217 218 219 220  3 . 5 . 15 70 2 . 7 . 12.11 2 . 7 , 12.11 2 . 2 , 9 . 87 2 .0 . 8.97 2 . 0 , 8 . 97 1 .7. 7 .62 1 .6 . 7 . 18 | 5 . 6 . 73 1 3 . 5 . 83 1 1 , 4 .93 3 3 , 15 . 14 2 7, 12 . 39 2 6 . 1 1 .93 2 1 . 9.63 2 0. 9.17 1 8 . 8 . 26 1 7 . 7 .80 1 6 . 7.34 1 4 , 6.42 1 3 . 5 . 96 1 3. 5.96 3 0 . 18 . 29 2 6 , 15.85 1 7 . 10. 37 1 5 . 9 . 15 1 3 . 7 .93 1 2. 7 . 32 1 2. 7 .32 1 1. 6 . 7 1 1 0 . 6 . 10 0 9 . 5 . 49 0 9 , 5 .49 3 0 . 15.54 2 7. 13 . 99 2 2. 1 1 .40 2 0 . 10. 36 1 8 . 9 .33 1 6 . 8 . 29 1 4 , 7.25 1 . 3 . 6 .74 1 . 3 . 6.74 1 . 2 . 6 . 22 0 8, 4 . 15 2 2. 14 .38 1 7, 11.11 1 . 7 . 11.11 1 . 4 . 9 . 15 1 . 3 . B .50 1 . 3 . 8.50 1 . 2 , 7 . 84 1 . 2 , 7.84 1 . 2 , 7.84 1 . 1 . 7 . 19 1 . 0 . 6 .54  22 1 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263.. 264 265 266 267 268 269 270 271 272 273 274 275  2 7 , 2 5. 2 5. 1 7. 1 7. 1 7. 1 6. 1 3. 1 2, 0 9, 0 9, 2 3, : i 7, i 2, 1 1 . 1 0, 0 9, 0 B, 0 8, o 8, 0 8. 0 7, 2 8. 2 0, 2 0. 1 8. 1 7, 1 3. 1 3. 1 2, 1 1 . 1 1 , O 9, 2 3. 2 0. 1 7. i 5, 1 4. 1 3, 1 3. 1 3. 1 2. 1 2, 1 o, 2 8. 2 2. 2 0, 1 7, 1 7, 1 3, 1 2. 1 0. 0 9. 0 9. 0 8.  14 .44 13 . 37 13 . 37 9.09 9.09 9.09 8.57 6 .95 6.42 4.81 4.81 19 . 0 0 14 .05 9.92 9.09 8 . 26 7 .44 6.61 6.61 6.61 6.61 5. 79 16 . 28 1 1 .63 1 1 .63 10.47 9.88 7 . 56 7 .56 6 . 98 6 . 40 6.40 5.23 14.20 12 . 35 10. 49 9 . 26 8 .64 8 .02 8 .02 8 .02 7.41 7.41 6 . 17 16.97 13.33 12 . 12 10.30 10.30 7 .88 7.27 6.06 5.45 5.45 4 .85  •2:03  APPENDIX  K - DISCRIMINANT ABSOLUTE  AND MAHALANOBIS DISTANCE ANALYSIS OF CHROMOSOME LENGTHS.  <SEPARATE OPTIONS=DISTANCE VAR=1-11 C A S E S = 1 - 2 5 SEPARATION OF STRATA CASES=CASE#:1-25  DISCRIMINANT  FUNCTIONS  8.V8)  VARIABLE  F-STAT  SIGNIF  CONSTANT 1 . V1 8.V8 (N= 25 OUT  15.675 10.224 25)  .0007 .0042  F-STAT  SIGNIF  NOT  OF  INCLUDED  . V2 . V3 . V4 . V5 . V6 . V7 . V9 10.V10 1 1 . V1 1  .69056 1 .2742 1 . 2663 .72730 .26019 .80111 .23459 .64168 - 1 .99985  MAHALANOBIS DISTANCES V12 (1)  STEP 0 1 2  (INCLUDE  (2)  STEP= 2 (1) -33.436 7.7619 35.736 14  .7953 .2717 .2732 . 4034 .6153 . 3809 .6332 .8025 . 3287  BETWEEN STRATA  D-SOUARE  F-STAT  SIGNIF  3.2912  9.6963  .0010  FORWARD SEPARATION  F-STAT  SIGNIF  INCLUDE INCLUDE INCLUDE  6.5439 10.224  .0176 .0042  <FINISH>  NO V A R I A B L E S 1.V1 8.V8  STRAT=V12>  (2) -34.309 13.454 25.044 1 1  240  APPENDIX L - CORRELATION MATRIX  OF ABSOLUTE CHROMOSOME LENGTHS  o CO aa 1C DC J1 CO IO < < < ^ 1 0 5 U I< < < < < < < CO G O CO ro V10  co I v  < >  r uo i  73  >  00  r-  ro CO  O A O O 73 a XI 73 m 73 r m > r-i > i i -t O m Z  2<> > 73  -i -4.  ui r ro oO c cn n ui -j 01 ro ro oo cn CO  <  IO  ro  <  CO  <  CO  UI  CO  CO  CO CO  CO  UI &  UI  CO -J  CO  UI  CO  CO  CO  CO  CO  CO  CD  .u .u n ui 00 o _h O cn c o ~j o o c nc -o oc oo o o cn co cn oO &  UI  uOi •~i ui  CO CO UI  01  CO  '-J  IO  ~4  CO  < 00 03 •  CO  03  03  CO  —  cn CO b O o o o r ro o a —  03  co  UI cn o ro 0 3o o O Ul o CO  CO  -  <  ^i o Oo O o o -J o o o o  CO CO  ucn i c 3 un i 09 0 0 9b o o o c n cn cn 03 o o  cn U l r o -I ro cn  <  UI  _»  UI  CO  CO CO -1 ' CO  CO  CO  UI  UI UI &  o o ui ui cn ucn i cn o O o ucn i o  CO  <n c•n c cn Ul cn ~j ro UI O cn O o -j -J  <  &  CO  CO  CO  O o o o  O O o o  o o o o  O Ul O co O CO 01  CO  •o. •o. O ro c 03 0 O cn nc n uuii0 O O O ~j • u r o -j ui ro O -*  CD  <  UI  ro —^ r o c n r o r cn ro oO Ol  CO  _*  II  ;a —  o o O uro i  21 1  APPENDIX  M - RECIPES AND SOURCES FOR ELECTROPHORETIC AND STAINS.  STARCH GELS  STARCH GEL RECIPES  MORPHOLINE CITRATE electrode distilled starch sucrose Source:  One %el  b u f f e r pH 6.1 water  modified  18 332 43.75 35  from C l a y t o n  Two g e l s ml ml g g  30 570 75 60  ml ml g g  60 540 75 60  ml ml g g  60 1 2.5 1  ml mg mg ml  50 50 100 100  mg mg mg ml  & T r e t i a k 1972  RIDGWAY c o n c e n t r a t e d g e l b u f f e r pH 8.5 d i s t i l l e d water starch sucrose Source:  modified  35 315 43.75 35  ml ml g g  from Ridgway e t a l . 1970  STAIN RECIPES DIAPHORASE (DIA) 0.2 M t r i s HC1 b u f f e r pH 8.0 2,6 d i c h l o r o p h e n o l i n d o p h e n o l NADH MTT Source:  Yeh & O ' M a l l e y  1980  ESTERASE ( E S T ) a-naphthyl acetate } d i s s o l v e d in b-naphthyl acetate } f a s t b l u e RR s a l t 0.2 M p h o s p h a t e b u f f e r pH 6.4  10 ml a c e t o n e  D i s s o l v e f a s t b l u e RR i n s m a l l amount t o remove i n p u r i t i e s . Source:  modified  from  GLUTAMIC DEHYDROGENASE  G o t t l i e b 1973 (GLDH)  of b u f f e r a n d f i l t e r  212  0.2 M t r i s HC1 b u f f e r L-glutamic a c i d NAD NBT PMS  pH 8.0  Source:  Shaw & P r a s a d  m o d i f i e d from  GLUCOSE-6-PHOSPHATE  60 2 1 1 1  DEHYDROGENASE  1970 and Hartmann e t a l . 1973  (G6PDH)  0.2 M t r i s HC1 b u f f e r d i s t i l l e d water 0.5 M MgC12 glucose-6-phosphate NADP NBT PMS Source:  m o d i f i e d from  ml g ml ml ml  .40 20 1 200 1 1.5 1 S i n g & Brewer  ml ml ml mg ml ml ml  1969 a n d Shaw & P r a s a d 1970  HEXOSEAMINIDASE (HA) 0.05 M p h o s p h a t e b u f f e r pH 4.0 d i s t i l l e d water 4-methylumbelli f e r y l - N - a c e t y l b-D-glucose aminide O b s e r v e under Source:  long-wave UV  50 ml 50 ml 100 mg  light.  m o d i f i e d from S i c i l i a n o  & Shaw 1976  ISOCITRIC DEHYDROGENASE (IDH) 0.2 M t r i s HC1 b u f f e r d i s t i l l e d water DL-isocitric acid 0.5 M MgC12 NADP NBT PMS Source:  pH 8.0  m o d i f i e d from A l l e n d o r f  40 20 250 1 1 1 1  ml ml mg ml ml ml ml  40 20 30 5 30  ml ml mg ml mg  e t a l . 1977  LEUCINE AMINOPEPTIDASE (LAP) 0.5 M t r i s m a l e a t e b u f f e r d i s t i l l e d water 1-leucyl-B-naphthylamide n,n d i m e t h y l formamide black K s a l t  pH 6.5  213  Dissolve Incubate Source:  1 - l e u c y l - B - n a p h t h y l a m i d e i n n,n d i m e t h y l 45 m i n , then add b l a c k K s a l t .  formamide.  Roose & G o t t l i e b 1976  MALIC DEHYDROGENASE  (MDH)  0.2 M t r i s HC1 b u f f e r pH 8.0 1 M Na L m a l a t e pH 7.0 NAD MTT PMS S o u r c e : m o d i f i e d from S i c i l i a n o  100 10 5 4 2  ml ml ml ml ml  40 20 1 2 2.5 1 1 1  ml ml ml ml ml ml ml ml  40 20 1 2 1.5 100 1 1.5 1  ml ml ml ml ml mg ml ml ml  40 20 20 1 1  ml ml mg ml ml  & Shaw 1976  PHOSPHOGLUCOSE ISOMERASE (PGI) 0.2 M t r i s HC1 b u f f e r pH 8.0 d i s t i l l e d water 0.5 M MgC12 glucose-6-phosphate dehydrogenase 0.018 M f r u c t o s e - 6 - p h o s p h a t e NADP MTT PMS S o u r c e : m o d i f i e d from G o t t l i e b 1973 PHOSPHOGLUCOMUTASE  (PGM)  0.2 M t r i s HC1 b u f f e r pH 8.0 d i s t i l l e d water 0.5 M MgC12 glucose-6-phosphate dehydrogenase 0.00017 M K2 a - D - g l u c o s e - 1 - p h o s p h a t e Na2 a - D - g l u c o s e - 1 - p h o s p h a t e NADP MTT PMS S o u r c e : m o d i f i e d from Roose & G o t t l i e b 1976 6-PHOSPHOGLUCONATE DEHYDROGENASE 0.2 M t r i s HC1 b u f f e r pH 8.0 d i s t i l l e d water 6-phosphogluconic a c i d 0.5 M MgC12 NADP  (6PGDH)  ml ml  MTT PMS Source: modified  from A l l e n d o r f  e t a l . 1977  215  APPENDIX N - NORTH AMERICAN HABITATS SEARCHED FOR Locality BRITISH COLUMBIA Vancouver, S t a n l e y Park, B e a v e r Lake V a n c o u v e r , U n i v e r s i t y of B r i t i s h C o l u m b i a Endowment Lands Ladner Marsh P i t t R i v e r Meadows B u r n a b y Lake Park WASHINGTON G r a y s H a r b o r Co. Ocean C i t y S t a t e P a r k Twin H a r b o r s S t a t e P a r k G r a y l a n d , Ewart S t . P a c i f i c Co. G r a y l a n d Beach S t a t e Park Oysterville I s l a n d Lake C r a n b e r r y M a r s h , 32nd P l a c e C r a n b e r r y Marsh, 55th S t r e e t OREGON C l a t s o p Co. C u l l a b y Lake C o u n t y Park T i l l a m o o k Co. Manzanita Manhattan Beach-Lake L y t l e Rockaway O l d - G r o w t h F o r e s t P r e s e r v e C h a m b e r l a i n Lake T i e r r a d e l Mar S e a r s Lake Neskowin L i n c o l n Co. D e v i l s Lake S t a t e Park E a s t D e v i l s Lake S t a t e Park S o u t h Beach S t a t e Park Lane Co. S u t t o n Lake Woahink Bog D o u g l a s Co. W i n c h e s t e r Bay Coos Co. B l u e b i l l Lake Second Creek F o u r t h Creek R e s e r v o i r B a s t e n d o r f f N a v a l S t a t i o n Bog B a s t e n d o r f f Beach Bandon C u r r y Co. G a r r i s o n Lake CALIFORNIA D e l N o r t e Co.  LIMBELLA.  Topographic  Map  North Vancouver Lulu  Island  «  92 G/3  92  II  G/6  P o r t C o q u i t l a m 92 G/7 New W e s t m i n s t e r 92 G/2 C o p a l i s Beach G r a y l a n d 7.5'  7.5'  O y s t e r v i l l e 7.5' Ocean Park 7.5' Cape D i s a p p o i n t m e n t  Gearhart  7.5'  Nehalem  15'  H  H  H  II  Tillamook Hebo 15'  15'  Cape F o u l w e a t h e r 15' E u c h r e M o u n t a i n 15' Y a q u i n a 15' H e c e t a Head 15' S i l t c o o s Lake 15' Reedsport Empire II  15'  7.5' H  Charleston H  II  II  II  Bandon  7.5'  15'  Langlois  15'  7.5  216  Yontocket Lake E a r l Dead L a k e C r e s c e n t C i t y South Humboldt C o . F r e s h w a t e r Lagoon Dry Lagoon S t a t e Park B i g Lagoon Mendocino C o . I n g l e n o o k Fen Lake C l e o n e  Smith R i v e r 7.5' Crescent C i t y 7.5'  It  Sister  TI  Rocks  7.5'  Orick 7.5' R o d g e r s Peak  7.5'  Inglenook 7.5' F o r t Bragg 7 . 5 '  PUBLICATIONS  1.  I l l u c t r c t i o n s i n : Honon, C. 1 9 7 5 . Guide t o t h e P l a n t s o f t h e W a l l o w a Mount Bins o f Nor the as t c m Oregon. S p e c i a l P u b l i c a t i o n s , Museum o f N a t u r a l H i s t o r y , U n i v e r s i t y Of O r e g o n . 411 p p .  2.  C h r i s t y , J . A . 1979. R e p o r t on n Survey o f Sphagnum-cent c l n l n g W e t l a n d s o f t h e Oregon C o n s t . Oregon N a t u r a l A r e a P r e s e r v e s A d v i s o r y C o m m i t t e e , E t c t e Lend B o a r d , Solera. 92 p p .  3.  . 355-358.  4.  _.- 1 9 8 0 . r e d i s c o v e r y o f £cicromiurn t r i c o c t c t u r o ( S u l l . ) K i t t . ( " L i m b e l l o t r i c o E f t . t a ( S u l l . ) B c r t r . ) i n N o r t h A m e r i c a . The E r y o l o R i s t 83521-523.  5.  . The  6.  1980. A d d i t i o n s  t o t h e noes f l o r a o f O r e g o n .  _ , J . H. L y f o r d L D. H. t'egner. B r y o l o g i o t B5: 22-36. _•  1983. Bibliogrcphy  The B r y o l o g i s t 83:  1 9 8 2 . C h e c k l i s t o f O r c g on D o c s e s .  o f Oregon c o c o e s .  Syccis  16: 43-52.  7.  Pvuching, A . E . & J . A . C h r i c t y . 1984. B r u c h i a f l e x u o s a tforth A m e r i c a . The B r y o l o g i c t 87: 68-69.  8.  C h r i s t y , J . A . 1984. New Sphagnum s p e c i e s from P a c i f i c N o r t h w e s t . o f t h e N a t i v e P l a n t S o c i e t y o f Oregon 1 7 ( 7 ) : 1 1 .  9.  F r e n k e l , R. E., W. H. K o i r £, J . A . C h r i s t y . 1985. V e g e t a t i o n o f Torrey Lake M i r e , c e n t r a l Cascade Range, Oregon. ( s u b m i t t e d t o M a d r o n o ) .  10. C h r i s t y , J . A., F. D . Bowers £> M. J . D i b b e n . Wisconsin. ( s u b m i t t e d t o The B r y o l o g i s t ) . 11.  (  edditions.  new t o w e s t e r n Bulletin  1 9 8 5 . F i v e mosses new t o  ,& 1985. A t l a s o f W i s c o n s i n b r y o p h y t e s : ( s u b m i t t e d t o M i l w a u k e e P u b l i c Museum P r e s s ) .  CBANT AWARDS  1983  Northwest Support  Scientific  f o r habitat  Washington, 1980  Oregon,  Mazamas  Research  Support  f o r survey  Association inventory northern  Research  of Limbella  Committee. tricostata  (Musci),  California.  Committee. of Sphapnim-cont a i n i n g  wetlands  of Cascade  Range,  Oregon. 1978  U n i v e r s i t y o f Oregon Undergraduate R e s e a r c h Committee. S u p p o r t f o r p u r v e y o f Sphajmum-cont a i n i n g w e t l a n d s o f O r e g o n  coast.  

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