Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

A phylogenetic systematics study of the red algal family Dumontiaceae Lindstrom, Sandra Christine 1985

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1985_A1 L56.pdf [ 26.13MB ]
Metadata
JSON: 831-1.0096665.json
JSON-LD: 831-1.0096665-ld.json
RDF/XML (Pretty): 831-1.0096665-rdf.xml
RDF/JSON: 831-1.0096665-rdf.json
Turtle: 831-1.0096665-turtle.txt
N-Triples: 831-1.0096665-rdf-ntriples.txt
Original Record: 831-1.0096665-source.json
Full Text
831-1.0096665-fulltext.txt
Citation
831-1.0096665.ris

Full Text

A PHYLOGENETIC SYSTEMATICS STUDY OF THE RED ALGAL FAMILY DUMONTIACEAE By SANDRA CHRISTINE LINDSTROM B. A., Reed C o l l e g e , 1971 M. Sc., The U n i v e r s i t y of B r i t i s h Columbia, 1973 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (DEPARTMENT OF BOTANY) We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA JUNE 1985 © Sandra C h r i s t i n e Lindstrom, 1985 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of Bo T/f AJ Y  The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date D E-6 ( 3 / 8 1 ) i i ABSTRACT A c l a d i s t i c a n a l y s i s o f t h e r e d a l g a l f a m i l y D u m o n t i a c e a e s u p p o r t s t h e h y p o t h e s i s t h a t t h e a n c e s t o r of t h e f a m i l y was m u l t i a x i a l , w i t h narrow p o o r l y d i f f e r e n t i a t e d a x i a l f i l a m e n t s , s m a l l p i t c o n n e c t i o n s , narrow r h i z o i d a l f i l a m e n t s , and i r r e g u l a r l y b r a n c h e d a s s i m i l a t o r y f i l a m e n t s o f c y l i n d r i c a l , u n i n u c l e a t e c e l l s . S e c o n d a r y p i t c o n n e c t i o n s and a c u t i c l e were p r o b a b l y l a c k i n g . The t e r e t e o r s l i g h t l y f l a t t e n e d , s u b t i d a l , e p h e m e r a l gametophyte may have a l t e r n a t e d w i t h a c r u s t o s e t e t r a s p o r o p h y t e w i t h s m a l l , l a t e r a l , b a s a l l y a t t a c h e d c r u c i a t e t e t r a s p o r a n g i a . The l o n g , s t r a i g h t c a r p o g o n i a l b r a n c h e s o f s l i g h t l y d i f f e r e n t i a t e d c e l l s t e r m i n a t e d i n a c a r p o g o n i u m w i t h a l o n g , s t r a i g h t t r i c h o g y n e . S p e r m a t a n g i a o c c u r r e d i n d o u b l e w h o r l s a r o u n d s p e r m a t a n g i a l mother c e l l s . F o l l o w i n g f e r t i l i z a t i o n , t h e c a r p o g o n i u m d i v i d e d t r a n s v e r s e l y , and t h e c o n n e c t i n g f i l a m e n t s t h a t d e v e l o p e d a f t e r f u s i o n t o c e l l s o f t h e c a r p o g o n i a l b r a n c h were p r o x i m a l l y s e p t a t e b u t d i s t a l l y n o n s e p t a t e . The a u x i l i a r y c e l l o c c u r r e d p r o x i m a l l y on a f i l a m e n t s l i g h t l y d i f f e r e n t i a t e d from a s s i m i l a t o r y f i l a m e n t s . The c o n n e c t i n g f i l a m e n t d i v i d e d a d j a c e n t t o an a u x i l i a r y c e l l , and t h e d i s t a l segment c o n t i n u e d on t o c o n t a c t o t h e r a u x i l i a r y c e l l s . The p r o x i m a l segment, a f t e r f u s i n g w i t h t h e a u x i l i a r y c e l l , c u t o f f 2-5 g o n i m o b l a s t i n i t i a l s t h a t p r o d u c e d a compact c y s t o c a r p o f many, s m a l l c a r p o s p o r a n g i a . E n l a r g e m e n t o f p i t c o n n e c t i o n s i n t h e a u x i l i a r y c e l l b r a n c h d i d n o t e x c e e d 5 /xm, and f u s i o n s between c e l l s of t h e a u x i l i a r y c e l l b r a n c h o r between c e l l s o f t h e g o n i m o b l a s t were l a c k i n g . S p o r e s g e r m i n a t e d i n a d i p r o t o c e l l u l a r p a t t e r n . The h y p o t h e s i z e d a n c e s t r a l c o n d i t i o n i n t h e D u m o n t i a c e a e s u p p o r t s t h e o r i g i n o f t h e f a m i l y f r o m a s p e c i e s i n t h e H e l m i n t h o c l a d i a c e a e s e n s u l a t o . No e v i d e n c e was f o u n d t h a t any member o f t h e D u m o n t i a c e a e i s e i t h e r a d i r e c t a n c e s t o r o r d e s c e n d a n t o f any o t h e r f a m i l i e s o f t h e C r y p t o n e m i a l e s o r G i g a r t i n a l e s w i t h t h e p o s s i b l e e x c e p t i o n of t h e P e y s s o n n e l i a c e a e , P o l y i d a c e a e , and R h i z o p h y l l i d a c e a e . As a r e s u l t o f t h e c l a d i s t i c a n a l y s i s , t h e t r i b e s F a r l o w i e a e and Dumontieae a r e r e c o g n i z e d as d i s t i n c t l i n e a g e s i n t h e D u m o n t i a c e a e , and t h e g e n e r a D u d r e s n a y a and N e o d i l s e a a r e r e c o g n i z e d t o be p a r a p h y l e t i c . In a d d i t i o n , t h e f o l l o w i n g t a x o n o m i c c h a n g e s were made: The g e n e r a A c r o s y m p h y t o n and N e o a b b o t t i e l l a were removed from t h e D u m o n t i a c e a e . F a r l o w i a  i r r e g u l a r i s was r e c o g n i z e d t o r e p r e s e n t a d i s t i n c t genus. N e o d i l s e a i n t e q r a was r e i n s t a t e d i n D i l s e a , and N. i n t e g r a v a r . l o n q i s s i m a was r a i s e d t o s p e c i f i c r a n k . The d i s c o v e r y o f o l d e r s p e c i f i c e p i t h e t s f o r N e o d i l s e a a m e r i c a n a and W e e k s i a f r y e a n a n e c e s s i t a t e d new s p e c i e s c o m b i n a t i o n s f o r t h e s e t a x a . A new genus and s p e c i e s was d e s c r i b e d f r o m A l a s k a . The genus T h u r e t e l l o p s i s was s y n o n y m i z e d w i t h D u d r e s n a y a and t h e f o l l o w i n g s y n o n y m i e s were s u g g e s t e d : T h u r e t e l l o p s i s j a p o n i c a w i t h D u d r e s n a y a minima, F a r l o w i a c o m p r e s s a w i t h F. m o l l i s , P i k e a  r o b u s t a w i t h P. c a l i f o r n i c a , W e e k s i a d i g i t a t a w i t h W. r e t i c u l a t a and p o s s i b l y b o t h o f t h e s e s p e c i e s w i t h t h e s p e c i e s f o r m e r l y known as W e e k s i a f r y e a n a . i v TABLE OF CONTENTS ABSTRACT i i TABLE OF CONTENTS i v LIST OF TABLES . x LIST OF FIGURES x i ACKNOWLEDGMENTS xv CHAPTER I. INTRODUCTION 1 CHAPTER I I . MATERIALS, METHODS AND TERMINOLOGY 16 CHAPTER I I I . SYSTEMATIC SURVEY 27 G i b s m i t h i a h a w a i i e n s i s 27 I n t r o d u c t i o n 27 Observations 28 Comments 32 K r a f t i a dichotoma 33 I n t r o d u c t i o n 33 Observations 33 Dudresnaya v e r t i c i l l . a t a 37 I n t r o d u c t i o n 37 Dudresnaya c r a s s a 44 I n t r o d u c t i o n 44 Dudresnaya a u s t r a l i s 47 I n t r o d u c t i o n 47 Observations 49 Dudresnaya j a p o n i c a 62 I n t r o d u c t i o n 62 Dudresnaya hawai i e n s i s 64 I n t r o d u c t i o n 64 Observations ....... 65 Dudresnaya bermudensis 71 I n t r o d u c t i o n 71 Observations 73 Dudresnaya minima 75 I n t r o d u c t i o n 75 Comments 77 Dudresnaya colombiana 77 I n t r o d u c t i o n 77 Observations 79 Dudresnaya l u b r i c a 81 I n t r o d u c t i o n 81 Observations 82 Dudresnaya p a t u l a 85 I n t r o d u c t i o n 85 Observations 85 Dudresnaya georgiana 89 I n t r o d u c t i o n 89 Observations 89 T h u r e t e l l o p s i s peqqiana and T. japo n i c a 93 I n t r o d u c t i o n 93 Observations 96 Comments 101 Acrosymphyton 102 Pikea c a l i f o r n i c a ..104 I n t r o d u c t i o n 104 Observations 114 v i F a r l o w i a m o l l i s . . . 121 I n t r o d u c t i o n 121 Observations 125 Comments 133 F a r l o w i a c o n f e r t a 1 35 I n t r o d u c t i o n 135 Observations 136 O r c u l i f i l u m d e n t i c u l a t u m 137 Di a g n o s i s 137 Observations 138 L e p t o c l a d i a binghamiae and L. peruviana 143 I n t r o d u c t i o n 143 Observations 147 Comments 152 Weeksia r e t i c u l a t a - 153 I n t r o d u c t i o n 153 Weeksia fryeana 156 I n t r o d u c t i o n 156 Weeksia t e m p l e t o n i i 160 I n t r o d u c t i o n 161 Weeksia h o w e l l i i 163 I n t r o d u c t i o n 163 Weeksia d i g i t a t a 165 I n t r o d u c t i o n . 165 Observations 165 Comments 169 Constantinea rosa-marina, C. simplex and C. s u b u l i f e r a .170 I n t r o d u c t i o n 170 v i i O bservations 183 Dumontia c o n t o r t a 192 I n t r o d u c t i o n ... 192 Observations . .. 200 Dumont i a simplex .205 I n t r o d u c t i o n 205 Observations And Comments .208 Dasyphloea i n s i q n i s 208 I n t r o d u c t i o n 208 Observations 209 C r y p t o s i p h o n i a woodii 212 I n t r o d u c t i o n 212 Observations 218 Comments 223 Hy a l o s i p h o n i a c a e s p i t o s a 225 I n t r o d u c t i o n 225 Observations 229 Fa r l o w i a i r r e g u l a r i s 233 I n t r o d u c t i o n 233 Observations ....235 Comments 238 D i l s e a and N e o d i l s e a 239 I n t r o d u c t i o n 239 D i l s e a carnosa .240 I n t r o d u c t i o n 240 Observations .248 D i l s e a c a l i f o r n i c a 250 I n t r o d u c t i o n .250 Observations 254 Comments 259 D i l s e a i n t e g r a 260 I n t r o d u c t i o n 260 Observations 265 Comments ..267 Neodi l s e a yendoana 269 I n t r o d u c t i o n 269 Observations 272 Comments 275 Neo d i l s e a c r i s p a t a and N. i n t e g r a var. l o n g i s s i m a 276 I n t r o d u c t i o n 276 Observations And Comments .278 Neo d i l s e a natashae 281 I n t r o d u c t i o n 281 Observations 281 Neodi l s e a americana .284 I n t r o d u c t i o n 284 Observations ...285 Neodi l s e a tenuipes 293 I n t r o d u c t i o n ..293 Observations And Comments 294 N e o a b b o t t i e l l a araneosa 295 CHAPTER IV. RESULTS AND DISCUSSION 297 Character A n a l y s i s 297 Character E v o l u t i o n 299 Taxonomic C o n c l u s i o n s 341 Biogeography 348 Some Furth e r Thoughts .352 Co n c l u s i o n .354 BIBLIOGRAPHY .358 TABLES 378 FIGURE ABBREVIATIONS 391 FIGURES 392 APPENDIX: REPRESENTATIVE SPECIMENS EXAMINED 574 X LIST OF TABLES I . Species C u r r e n t l y Recognized As Members Of The Dumontiaceae 378 I I . Comparison Of Chara c t e r s Of Four Species Of Weeksia Based On Abbott 1968 380 I I I . Comparison Of Spore And S p o r a n g i a l Dimensions In D i l s e a And Ne o d i l s e a 381 IV. C h a r a c t e r s And T h e i r Transformations Used In The C l a d i s t i c A n a l y s i s 382 V. Character S t a t e s Of The Species And Genera Used In The C l a d i s t i c A n a l y s i s 387 V I . C o n s i s t e n c y I n d i c e s For Ch a r a c t e r s In The C l a d i s t i c A n a l y s i s 388 V I I . Species Recognized As Members Of The Dumontiaceae By T h i s Study 389 x i LIST OF FIGURES 1. G i b s m i t h i a spp. 392 2. G i b s m i t h i a h a w a i i e n s i s .....394 3. G i b s m i t h i a sp .396 4. K r a f t i a dichotoma 398 5. Kraf t i a dichotoma 400 6. K r a f t i a dichotoma 402 7. Dudresnaya v e r t i c i l l a t a 404 8. Dudresnaya v e r t i c i l l a t a 406 9. A s s i m i l a t o r y f i l a m e n t s i n four s p e c i e s of Dudresnaya ..408 1 0 . Dudresnaya c r a s s a 410 1 1 . Dudresnaya c r a s s a 41 2 12. Dudresnaya a u s t r a l i s 414 13. Dudresnaya japonica 416 14. Dudresnaya h a w a i i e n s i s 418 15. Dudresnaya hawai i e n s i s 420 16. Dudresnaya bermudensis 422 17. Dudresnaya colombiana 424 18. Dudresnaya l u b r i c a 426 19. Dudresnaya p a t u l a ....428 20. Dudresnaya p a t u l a 430 21. Dudresnaya qeorgiana 432 22. T h u r e t e l l o p s i s peqqiana 434 23. T h u r e t e l l o p s i s peqqiana 436 24. T h u r e t e l l o p s i s peqgiana 438 25. Pikea c a l i f o r n i c a .440 x i i 26. Pikea c a l i f o r n i c a 442 27. Pikea c a l i f o r n i c a 444 28. A x i a l s t r u c t u r e i n F a r l o w i a , O r c u l i f i l u m , Pikea, and L e p t o c l a d i a 446 29. Pikea c a l i f o r n i c a 448 30. Pikea c a l i f o r n i c a 450 31. Pikea c a l i f o r n i c a 452 32. Pikea c a l i f o r n i c a 454 33. F a r l o w i a m o l l i s 456 34. F a r l o w i a moll i s 458 35. F a r l o w i a moll i s 460 36. F a r l o w i a m o l l i s 462 37. F a r l o w i a m o l l i s 464 38. F a r l o w i a m o l l i s 466 39. F a r l o w i a m o l l i s 468 40. F a r l o w i a c o n f e r t a and F. m o l l i s 470 41. L e p t o c l a d i a binqhamiae, L. peruviana, and O r c u l i f i l u m  d e n t i c u l a t u m 472 42. O r c u l i f i l u m d e n ticulatum 474 43. O r c u l i f ilum denticulatum .476 44. O r c u l i f i l u m d e n t i c u l a t u m 478 45. O r c u l i f ilum denticulatum 480 46. L e p t o c l a d i a binqhamiae 482 47. L e p t o c l a d i a binqhamiae 484 48. L e p t o c l a d i a binqhamiae 486 49. L e p t o c l a d i a spp 488 50. Weeksia spp 490 51. Weeksia d i q i t a t a 492 x i i i 52. Weeksia d i g i t a t a 494 53. Weeksia d i g i t a t a 496 54. Weeksia spp 498 55. Constantinea spp 500 56. Constantinea s u b u l i f e r a 502 57. Constantinea spp ..504 58. Constantinea spp. 506 59. Dumontia c o n t o r t a 508 60. Dumontia c o n t o r t a 510 61. Dasyphloea i n s i g n i s 512 62. C r y p t o s i p h o n i a woodi i 514 63. C r y p t o s i p h o n i a woodii 516 64. C r y p t o s i p h o n i a woodii 518 65. H y a l o s i p h o n i a c a e s p i t o s a 520 66. F a r l o w i a i r r r e q u l a r i s 522 67. F a r l o w i a i r r e g u l a r i s . ... 524 68. F a r l o w i a i r r e g u l a r i s 526 69. D i l s e a and Neo d i l s e a spp 528 70. D i l s e a and Neo d i l s e a spp. 530 71. D i l s e a and Neo d i l s e a spp 532 72. D i l s e a carnosa 534 73. D i l s e a carnosa ......536 74. D i l s e a carnosa 538 75. D i l s e a carnosa 540 76. D i l s e a c a l i f o r n i c a 542 77. D i l s e a c a l i f o r n i c a 544 78. D i l s e a c a l i f o r n i c a .....546 79. D i l s e a c a l i f o r n i c a 548 x i v 80. D i l s e a i n t e g r a 550 81. N e o d i l s e a yendoana 552 82. N e o d i l s e a l o n g i s s i m a and N e o d i l s e a c r i s p a t a .554 83. N e o d i l s e a l o n g i s s i m a .556 84. N e o d i l s e a natashae 558 85. N e o d i l s e a natashae 560 86. N e o d i l s e a b o r e a l i s 562 87. N e o d i l s e a b o r e a l i s 564 88. N e o d i l s e a b o r e a l i s ........566 89. N e o d i l s e a b o r e a l i s 568 90. N e o d i l s e a tenuipes 570 91. Cladogram of the Dumontiaceae 572 XV ACKNOWLEDGMENTS I extend my s i n c e r e a p p r e c i a t i o n to a l l who c r i t i c a l l y read p a r t s or a l l of the t h e s i s , to a l l who pro v i d e d specimens f o r examination, and to a l l who helped with r e f e r e n c e s , t r a n s l a t i o n s , and the c o u n t l e s s other e s s e n t i a l t asks i n t h e s i s p r e p a r a t i o n . Included among these numerous people, I e s p e c i a l l y thank J . Boulogne, K. M. Cole, R. E. DeWreede, P. W. G a b r i e l s o n , L. Golden, M. W. Hawkes, M. H. Hommersand, Y. P. Lee, J . Maze, J . C. O l i v e i r a , R. F. S c a g e l , and L. J . Veto. 1 CHAPTER I. INTRODUCTION The Dumontiaceae c u r r e n t l y c o n s i s t s of 50 s p e c i e s i n 17 genera (Table I ) . P r e v i o u s l y d e s c r i b e d s p e c i e s which are c u r r e n t l y c o n s i d e r e d synonyms of recognized f a m i l y members are d i s c u s s e d i n the body of the t h e s i s . The f a m i l y name Dumontiaceae has been a t t r i b u t e d to two authors--Bory ( S i l v a 1980:82) and Schmitz ( K y l i n 1956:142). Bory (1828:197) c h a r a c t e r i z e d the " F a m i l l e des Dumontiees, Dumontiae" as, roughly t r a n s l a t e d , not at a l l d i s t i n g u i s h e d by the stem in i t s o r g a n i z a t i o n , c o n s i s t i n g f r e q u e n t l y of simple tubes, more or l e s s compound, with young i n d i v i d u a l s produced by the germination of spores being taken as branches, and most i n d i v i d u a l s the c o l o r of other marine F l o r i d e a e 1 , be i t p u r p l e or green. Bory a t t r i b u t e d four s p e c i e s i n three genera to t h i s f a m i l y , Dumontia f a s t i g i a t a . (Bory) Bory [=Nothogenia f a s t i q i a t a (Bory) P a r k i n s o n ] , Asperococcus l e s s o n i i Bory [=Adenocystis  u t r i c u l a r ! s (Bory) S k o t t s . ] , Asperococcus d u r v i l l a e i Bory [ = U t r i c u l i d i u m d u r v i l l a e i (Bory) S k o t t s . ] , and S o l e n i a compressa (L.) Bory [=Enteromorpha compressa (L.) G r e v i l l e ] . These s p e c i e s were c o l l e c t e d , probably in New Zealand, by the n a t u r a l i s t s on La C o q u i l l e . Although Bory d i d not s p e c i f i c a l l y mention Dumont i a i n c r a s s a t a (Mueller) Lamouroux, the type of the genus and hence the f a m i l y , he d i d not s p e c i f i c a l l y exclude i t . T h e r e f o r e the I n t e r n a t i o n a l Code of B o t a n i c a l Nomenclature c o n s i d e r s i t a ' S i l v a (1980:79) has d i s c u s s e d the use of F l o r i d e a e as a s u b c l a s s name (Schmitz 1892: 16, nom. d e s c i p t . ) . Since K y l i n , most authors have p r e f e r r e d the o r t h o g r a p h i c a l l y c o r r e c t name F l o r i d e o p h y c i d a e . I have used whichever name the authors themselves employed. 2 l e g i t i m a t e , v a l i d l y p u b l i s h e d f a m i l y name; however, had Bory not in c l u d e d the L a t i n f a m i l y name Dumontiae, which Schmitz (1889:453) c o r r e c t e d to Dumontiaceae, the famiy name would not have been v a l i d l y p u b l i s h e d ( N i c o l s o n , pers. comm., 27 June 1983). D i f f e r e n t n i n e t e e n t h century p h y c o l o g i s t s adopted v a r i o u s schemes that bear l i t t l e resemblance to our c u r r e n t l y accepted c l a s s i f i c a t i o n of the Rhodophyceae i n general and the Dumontiaceae i n p a r t i c u l a r . For example, J . Agardh (1851:233) i n c l u d e d Constantinea i n the T r i b e Kallymenieae of the Order G i g a r t i n e a e . Dumontia, ? H a l i s a c c i o n [ s i c ] , C a t e n e l l a , and Rhabdonia, were pl a c e d i n the T r i b e Dumontieae i n the Order Dumontieae (1852:348). Although h i s s u p r a f a m i l i a l c l a s s i f i c a t i o n v a r i e d from the one now g e n e r a l l y f o l l o w e d , by 1876 J . Agardh had brought C r y p t o s i p h o n i a , Pikea, ?Dasyphloea (as N i z z o p h l o e a ) , Dumontia, ?Ha l o s a c c i o n , F a r l o w i a , and D i l s e a (as S a r c o p h y l l i s ) together i n t o a s i n g l e taxon, which he i d e n t i f i e d as "Ordo VI. Dumontiaceae" i n " S e r i e s I I I . Nematospermeae, Subseries I." J . Agardh c l a s s i f i e d Dudresnaya ( i n c l u d i n g D. p u r p u r i f e r a , now Acrosymphyton purpuriferum) i n "Ordo V. Dudresnayeae," but he maintained Constantinea together with Kallymenia i n the T r i b e Kallymenieae, Order G i g a r t i n e a e . Schmitz (1883, see D a l l a s 1884 f o r E n g l i s h t r a n s l a t i o n ) , i n h i s epoch-making t r e a t i s e on p o s t - f e r t i l i z a t i o n development i n the F l o r i d e a e , i n c l u d e d comparative o b s e r v a t i o n s on three s p e c i e s i n the Dumontiaceae, Dudresnaya p u r p u r i f e r a , D. co c c i n e a , and Dumontia f i l i f o r m i s . He d e s c r i b e d i n some d e t a i l the stages from the formation of connecting f i l a m e n t s by the 3 f e r t i l i z e d carpogonium i n c o n j u n c t i o n with neighboring c e l l s (at l e a s t i n D. purpuriferum) to the f u s i o n of the segmented connecting f i l a m e n t with a remote a u x i l i a r y c e l l and the i n i t i a t i o n of sporangia from the remnant of the connecting f i l a m e n t a t t a c h e d to the a u x i l i a r y c e l l . Subsequently, Schmitz (1889) proposed a new c l a s s i f i c a t i o n of the genera i n the F l o r i d e a e . Although other f a m i l i e s experienced s i g n i f i c a n t s h i f t s , the Dumontiaceae remained u n a l t e r e d from J . Agardh's c l a s s i f i c a t i o n except f o r the i n c l u s i o n of Dudresnaya and Consta n t i n e a i n the f a m i l y and the removal of H a l o s a c c i o n . Schmitz (1889) i n c l u d e d the Dumontiaceae i n the Cryptoneminae, l a t e r changed t o the Cryptonemiales (Schmitz 1892, non v i d i ) a f t e r the G l o i o s i p h o n i a c e a e and G r a t e l o u p i a c e a e 1 (=Cryptonemiaceae/Halymeniaceae; see S i l v a 1980:82, 84) and before the Nemastomataceae (=Gymnophloeaceae, now i n the G i g a r t i n a l e s ) , R h i z o p h y l l i d a c e a e ( a s c r i b e d by Wiseman 1975, to the G i g a r t i n a l e s ) , Squamariaceae (=Peyssonneliaceae), and C o r a l l i n a c e a e (soon to be p l a c e d i n i t s own o r d e r ) . He p l a c e d the Cryptonemiales at the end of the F l o r i d e a e , a f t e r the Rhodymeniales. In h i s i n t r o d u c t i o n , he s t a t e d that the l i n e a r arrangement of the f a m i l i e s p r o v i d e d only a rough approximation of the many-faceted resemblances and r e l a t i o n s h i p s of these taxa. Schmitz and H a u p t f l e i s c h (1897:515) observed the same order of f a m i l i e s . In t h e i r n a r r a t i v e account of the Dumontiaceae, 'Family names f o l l o w those of the o r i g i n a l a u t h o r s . C u r r e n t l y accepted or p r o p o s e d - f o r - c o n s e r v a t i o n names are i n d i c a t e d i n parentheses a f t e r the f i r s t use of a name o n l y . 4 they c h a r a c t e r i z e d the f a m i l y by the many-celled, separate c a r p o g o n i a l and a u x i l i a r y c e l l branches s c a t t e r e d i n l a r g e numbers through the t h a l l u s and by d e t a i l s of p o s t - f e r t i l i z a t i o n development a l r e a d y d e s c r i b e d above. To the e i g h t genera a l r e a d y i n c l u d e d i n the f a m i l y , they added A n d e r s o n i e l l a Schmitz (=Leptocladia J . Ag. 1892). Er y t h r o p h y l l u m J . Ag., now c o n s i d e r e d a s p e c i e s of Kallymeniaceae, was i n c l u d e d under "Gattung u n s i c h e r e r S t e l l u n g . " Oltmanns (1904) moved the Cryptonemiales from i t s p o s i t i o n as the l a s t order i n Schmitz's l i s t to a p o s i t i o n between the Nemaliales and G i g a r t i n a l e s . Beginning i n 1923, K y l i n began p u b l i s h i n g a s e r i e s of d e t a i l e d s t u d i e s of r e p r o d u c t i v e s t r u c t u r e s and post-f e r t i l i z a t i o n development in s p e c i e s of red algae ( p r i o r to t h i s time each of h i s papers encompassed only a s i n g l e s p e c i e s ) . He adopted Schmitz's e m b r y o l o g i c a l b a s i s f o r c l a s s i f y i n g the F l o r i d e a e and s t r e s s e d as major c h a r a c t e r s c a r p o g o n i a l branch s t r u c t u r e , a u x i l i a r y c e l l p o s i t i o n , sporogenous f i l a m e n t behavior, and gonimoblast development. K y l i n (1923:133) concluded that the Cryptonemiales represented a lower developmental l i n e than the other three major d i p l o b i o n t i c o r d e r s because i t l a c k e d p r o c a r p s . S j o s t e d t (1926), who a l s o s t u d i e d r e p r o d u c t i v e s t r u c t u r e s of a number of higher F l o r i d e a e , p r o v i d e d another c r i t i q u e of t h e i r c l a s s i f i c a t i o n . He concurred with Oltmanns and K y l i n that the Cryptonemiales represented the most p r i m i t i v e and l e a s t d i f f e r e n t i a t e d group among the o r d e r s under c o n s i d e r a t i o n . He b e l i e v e d that the Dumontiaceae and G l o i o s i p h o n i a c e a e represented 5 p a r a l l e l p r i m i t i v e groups i n the Cryptonemiales and that from the Dumontiaceae were d e r i v e d , i n separate p a r a l l e l l i n e s , " a l l the other h i t h e r t o e m b r y o l o g i c a l l y b e t t e r known d i p l o b i o n t i c F l o r i d e a e , " i . e . , the R h i z o p h y l l i d a c e a e , Squamariaceae, and G r a t e l o u p i a c e a e i n the Cryptonemiales, the Nemastomataceae, Rhabdoniaceae (=Caulacanthaceae), and Rhodophyllidaceae (=Cystocloniaceae) i n the Nemastomales, the Rhodymeniales, the Sphaerococcales ( i n c l u d i n g Plocamium and Sphaerococcus) and the G i g a r t i n a l e s ( i n c l u d i n g only the G i g a r t i n a c e a e i n S j o s t e d t ' s system); only the G e l i d i a l e s , the Ceramiales, and G r a c i l a r i a were p l a c e d elsewhere. K y l i n (1928, 1930), l i k e S j o s t e d t , viewed the Cryptonemiales i n a narrow as w e l l as a broad sense. The Cryptonemiales sensu l a t o encompassed not j u s t the Cryptonemiales but a l s o the Nemastomales (an order proposed but not v a l i d l y p u b l i s h e d by K y l i n i n 1925 to c o n t a i n the Nemastomataceae, Rhabdoniaceae, and Rhodophyllidaceae), the Rhodymeniales, and the G i g a r t i n a l e s . K y l i n c o n s i d e r e d the C r u o r i a c e a e to be the outgroup of t h i s "superorder" and s e q u e n t i a l l y drew o f f the Nemastomales, the Rhodymeniales, the Cryptonemiales, and the G i g a r t i n a l e s . Within the Cryptonemiales, he showed two l i n e s . In one l i n e , i n which the sporogenous f i l a m e n t s f i r s t fuse with s p e c i a l n u t r i t i v e c e l l s i n the c a r p o g o n i a l branch and only then grow toward the a u x i l i a r y c e l l , the Dumontiaceae represented the s i s t e r group of the R h i z o p h y l l i d a c e a e , Squamariaceae, and C o r a l l i n a c e a e . He d i s t i n g u i s h e d the Dumontiaceae from these other f a m i l i e s by the f a c t that the c a r p o g o n i a l and a u x i l i a r y c e l l branches are 6 s c a t t e r e d throughout the t h a l l u s r a t h e r than o c c u r r i n g i n nemathecia. In the other l i n e , i n which no c e l l s of the c a r p o g o n i a l branch are c ontacted before the g e n e r a t i v e a u x i l i a r y c e l l , he i n c l u d e d the G r a t e l o u p i a c e a e , G l o i o s i p h o n i a c e a e , Endocladiaceae, G i g a r t i n a c e a e , and Kallymeniaceae. K y l i n (1932) dropped t h i s scheme i n favor of e n t i r e l y separate schemes f o r the Cryptonemiales and G i g a r t i n a l e s (the l a t t e r now i n c l u d i n g the f a m i l i e s of the Nemastomales p l u s the G i g a r t i n a c e a e ) . The 1932 scheme of the Cryptonemiales was e s s e n t i a l l y that of the 1930 p u b l i c a t i o n with the a d d i t i o n of the Tichocarpaceae and the Kallymeniaceae. In 1930, K y l i n s i n g l e d out the p r o p e r t i e s of the a u x i l i a r y c e l l as r e p r e s e n t i n g the most important c h a r a c t e r i n f l o r i d e a e n s y s t e m a t i c s , and i n 1932, he o u t l i n e d the c h a r a c t e r i s t i c s of the a u x i l i a r y c e l l that d i s t i n g u i s h each order of the F l o r i d e o p h y c i d a e . In p a r t i c u l a r , he d i s t i n g u i s h e d the Cryptonemiales from the G i g a r t i n a l e s on the b a s i s of the accessory nature of the branch b e a r i n g the a u x i l i a r y c e l l . (The a u x i l i a r y c e l l i s s t i l l used as a major c r i t e r i o n f o r o r d i n a l c l a s s i f i c a t i o n . ) F r i t s c h (1945:656) questioned the importance of t h i s d i s t i n c t i o n between the Cryptonemiales and the G i g a r t i n a l e s . He c o n s i d e r e d a d e f i n i t i o n of the G i g a r t i n a l e s based on an i n t e r c a l a r y , v e g e t a t i v e a u x i l i a r y c e l l and that of the Cryptonemiales on an a u x i l i a r y c e l l l o c a t e d i n a s p e c i a l accessory branch to be a r a t h e r " t r i v i a l f e a t u r e upon which to base a major taxonomic s u b d i v i s i o n . " N e v e r t h e l e s s , he f e l t such a d e l i n e a t i o n served a u s e f u l purpose u n t i l c l e a r e r concepts of 7 a f f i n i t i e s c o u l d be obt a i n e d . T h i s d i s t i n c t i o n was a l s o q u e s t i o n e d by Drew (1957), who p o i n t e d out that t h i s c h a r a c t e r does not h o l d i n a l l cases. S e a r l e s (1968:1) wrote, " I t i s not yet c l e a r , however, whether these orders represent n a t u r a l assemblages or only convenient groupings." K y l i n ' s emphasis on c h a r a c t e r i s t i c s of the a u x i l i a r y c e l l i n f l o r i d e o p h y c i d e a n c l a s s i f i c a t i o n i n f l u e n c e d a t l e a s t a gen e r a t i o n of p h y c o l o g i s t s to concentrate on red a l g a l r e p r o d u c t i v e morphology i n order to r e f i n e the K y l i n i a n system. Papenfuss (1951) t r i e d to c l a r i f y the d i s t i n c t i o n s between the d i f f e r e n t types of a u x i l i a r y c e l l s . He b e l i e v e d the Dumontiaceae to have two kinds of a u x i l i a r y c e l l s , n u t r i t i v e and g e n e r a t i v e , and he d e s c r i b e d the Weeksiaceae (a segregate of the Dumontiaceae proposed by Abbott i n a then unpublished t h e s i s ) as having n u t r i t i v e a u x i l i a r y c e l l s which have become g e n e r a t i v e a u x i l i a r y c e l l s . (Papenfuss a p p a r e n t l y d i d not r e a l i z e t h a t the gonimoblast of members of the Dumontiaceae develops not from the a u x i l i a r y c e l l i t s e l f but from the connecting f i l a m e n t f o l l o w i n g establishment of a connection with an a u x i l i a r y c e l l , thereby g i v i n g the Dumontiaceae two kinds of n u t r i t i v e a u x i l i a r y c e l l s . ) Drew (1954:57) d i s t i n g u i s h e d four major types of carposporophytes found i n the Florideophycidae:' (a) carposporangia-producing gonimoblast not f u s i n g with other c e l l s of the gametophyte but a r i s i n g d i r e c t l y from the carpogonium, (b) carposporangia-producing gonimoblast f u s i n g with other c e l l s of the gametophyte but not t r a n s f e r r i n g the d i p l o i d nucleus to another c e l l , (c) primary gonimoblast t r a n s f e r r i n g the d i p l o i d nucleus to s p e c i f i e d c e l l s at some d i s t a n c e from the carpogonium 8 from which a r i s e carposporangia-producing gonimoblast, and (d) carposporangia-producing gonimoblast a r i s i n g from a s p e c i f i e d c e l l of the gametophyte r e l a t i v e l y near the carpogonium and to which the d i p l o i d nucleus i s t r a n s f e r r e d . Drew (p. 60) i n c l u d e d Acrosymphyton i n Group B ( " F i n a l l y , i n Acrosymphyton, the spore-producing l a t e r a l s develop from the gonimoblast i t s e l f as i n , t h e other cases but r i g h t at the p o i n t of f u s i o n with the c e l l of the gametophyte") and the r e s t of the Dumontiaceae i n Group C ("This type, as represented by Dudresnaya c o c c i n e a , has been we l l known s i n c e the tim