Vegetative and reproductive innovations of early land plants: Implications for a unified phylogeny

Philosophical Transactions of the Royal Society B: Biological Sciences

Volumn 355, Issue 1398, 2000, Pages 769-793

  Renzaglia, Karen Sue ^a   Duff, R Joel ^a,c   Nickrent, Daniel L ^a   Garbary, David J ^b  

^a SOUTHERN ILLINOIS UNIVERSITY   (United States)

^b ST FRANCIS XAVIER UNIVERSITY   (Canada)

^c UNIVERSITY OF AKRON   (United States)

Author keywords

Bryophytes; Embryophytes; Gametophyte; Land plant phylogeny; Morphogenesis; Spermatogenesis

Indexed keywords

BRYOPHYTE; ECOMORPHOLOGY; PALEOBOTANY; PHYLOGENY; REPRODUCTION;

BRYOPHYTA; EQUISETUM;

EID: 0034729785     PISSN: 09628436     EISSN: None     Source Type: Journal    
DOI: 10.1098/rstb.2000.0615     Document Type: Conference Paper

References (130)

1. Bartholomew, S. E. & Crandall-Stotler, B. J. 1986 The sporeling ontogeny of Monoclea gottschei subsp. elongata. Bryologist 97, 244-252.
2. Bartholomew-Began, S. E. 1991 A morphogenetic re-evaluation of Haplomitrium Nees Hepatophyta). Bryophytorum Bibliotheca, Band 41. Berlin: J. Cramer.
3. Bartlett, E. M. 1928 The comparative study of the development of the sporophyte in the Anthocerotaceae, with special reference to the genus Anthoceros. Ann. Bot. 42, 409-430.
4. Bateman, R. M. & DiMichele, W. A. 1994 Heterospory: the most iterative key innovation in the history of the plant kingdom. Biol. Rev. 69, 345-417.
5. Bateman, R. M., Crane, P. R., DiMichele, W. A., Kenrick, P. R. & Rowe, N. P. 1998 Early evolution of land plants: phylogeny, physiology and ecology of the primary terrestrial radiation, A. Rev: Ecol. Syst. 29, 263-292.
6. Beckert, S., Steinhauser, S., Muhle, H. & Knoop. V. 2000 A molecular phylogeny of bryophytes based on nucleotide sequences of the mitochondrial sequences of the mitochondrial nad5 gene. Pl. Syst. Evol. 218, 179-192.
7. Bell, P. R. 1994 Green plants: their origin and diversity. Portland, OR: Dioscorides Press.
8. Bernhard, D. L. & Renzaglia, K. S. 1995 Spermiogenesis in the moss Aulacomnium palustre. Bryologist 98, 52-70.
9. Bierhorst, D. W. 1971 Morphology of vascular plants. New York: MacMillan.
10. Blackmore, S. & Barnes, S. H. 1987 Embryophyte spore walls: origin, development, and homologies. Cladistics 3, 185-195.
11. Bold, H. C., Alexopoulos, C. J. & Delevoryas, T. 1987 Morphology of plants and fungi, 5th edn. New York: MacMillan.
12. Bopp, M. & Capesius, I. 1996 New aspects of bryophyte taxonomy provided by molecular approach. Bot. Acta 109, 1-5.
13. Bremer, K., Humphries, C. J., Mishler, B. D. & Churchill, S. P. 1987 On cladistic relationships in green plants. Taxon 36, 339-349.
14. Brown, R. C. & Lemmon, B. E. 1988 Sporogenesis in bryophytes. Adv. Bryol. 3, 159-223.
15. Brown, R. C. & Lemmon, B. E. 1990a Monoplastidic cell division in lower land plants. Am. J. Bot. 77, 559-571.
16. Brown, R. C. & Lemmon, B. E. 1990b Sporogenesis in bryophytes. In Microspores: evolution and ontogeny (ed. S. Blackmore & R. V. Knox), pp. 55-94. London: Academic Press.
17. Brown, R. C. & Lemmon, B. E. 1993 Diversity of cell division in simple land plants holds clues to evolution of the mitotic and cytokinetic apparatus in higher plants. Mem. Torrey Bot. Club 25, 45-62.
18. Brown, R. C. & Lemmon, B. E. 1997 The quadripolar microtubule system in lower land plants. J. Plant Res. 110, 93-106.
19. Brown, R. C., Lemmon, B. E. & Renzaglia, K. S. 1986 Sporocytic control of spore wall patterns in liverworts. Am. J. Bot. 73, 593-596.
20. Campbell, D. H. 1895 The structure and development of mosses and ferns (Archegoniatae). New York: Macmillan.
21. Capesius, I. & Bopp, M. 1997 New classification of liverworts based on molecular and morphological data. Pl. Syst. Evol. 207, 87-97.
22. Carothers, Z. B. & Duckett J. G. 1980 The bryophyte spermatozoid: a source of new phylogenetic information. Bull. Torrey Bot. Club 107, 281-297.
23. Crandall-Stotler, B. J. 1980 Morphogenetic designs and a theory of bryophyte origins and divergence. BioScience 30, 580-585.
24. Crandall-Stotler, B. J. 1984 Musci, hepatics and anthocerotes -an essay on analogues. In New manual of bryology, vol. 2 (ed. R. M. Schuster), pp. 1093-1129. Nichinan, Japan: Hattori Botanical Laboratory.
25. Crandall-Stotler, B. J. 1986 Morphogenesis, developmental anatomy and bryophyte phylogenetics; contraindications of monophyly. J. Bryol. 14, 1-23.
26. Crum, H. A. & Anderson, L. E. 1981 Mosses of eastern North America, vols. I II. New York: Columbia University Press.
27. Duckett, J. G. 1970 Sexual behaviour of the genus Equisetum, subgenus Equisetum. Bot. J. Linn. Soc. 63, 327-352.
28. Duckett, J. G. & Duckett, A. R. 1980 Reproductive biology and population dynamics of wild gametophytes of Equisetum. Bot. J. Linn. Soc. 79, 205-210.
29. Duckett, J. G. & Ligrone, R. 1995 The formation of catenate foliar gemmae and the origin of oil bodies in the liverwort Odotoschisma denudatum (Mart.) Dum. (Jungermanniales): a light and electron microscope study. Ann. Bot. 76, 405-419.
30. Ducken, J. G. & Renzaglia, K. S. 1988 Cell and molecular biology of bryophytes: ultimate limits to the resolution of phylogenetic problems. Bot. J. Linn. Soc. 98, 225-246.
31. Duckett, J. G., Renzaglia, K. S. & Pell, K. 1990 Desiccation causes the proliferation of multicellular hairs but not mucilage papillae in Cryptothallus mirabilis: a correlated light and electron microscope study. Can. J. Bot. 68, 697-706.
32. Duckett, J. G., Renzaglia, K. S. & Pell, K. 1991 A light and electron microscope study of rhizoid ascomycete associations and flagelliform axes in British hepatics with observations on the effects of the fungi on host morphology, and comparisons with mycorrhizae in the Ericaceae. New Phytol. 118, 233-257.
33. Duff, J. R. & Nickrent, D. L. 1999 Phylogenetic relationships among land plants using mitochondrial small-subunit rDNA sequences. Am. J. Bot. 86, 372-386.
34. Duncan, T. M., Renzaglia, K. S. & Garbary, D. J. 1997 Ultrastructure and phylogeny of the spermatozoid of Chara vulgaris (Charophyceae). Pl. Syst. Evol. 204, 125-140.
35. Fulford, M. 1956 The young stages of the leafy Hepaticae: a resumé. Phytomorphology 6, 199-235.
36. Garbary, D. J. & Renzaglia, K. S. 1998 Bryophyte phylogeny and the evolution of land plants: evidence from development and ultrastructure. In Bryology for the twenty-first century (ed. J. W. Bates, N. W. Ashton & J. G. Duckett), pp.45-63. Leeds: The British Bryological Society and Maney Publishing.
37. Garbary, D. J., Recnzaglia, K. S. & Duckett, J. G. 1993 The phylogeny of land plants: a cladistic analysis based on male gametogenesis. Pl. Syst. Evol. 188, 237-269.
38. Gifford, E. M. & Foster, A. S. 1988 Morphology and evolution of vascular plants, 3rd edn. New York: Freeman.
39. Gould, S. J. 1989 Wonderful life - the Burgess Shale and the nature of history. New York: W. W. Norton & Co.
40. Graham, L. E. 1993 Origin of land plants. New York: John Wiley.
41. Gunning, B. E. & Steer, M. W. 1996 Plant cell biology: structure and function. Sudbury, MA: Jones and Bartlett Publishers.
42. Hanson, D. T., Swanson, S., Graham, L. E. & Sharkey, T. D. 1999 Evolutionary significance of isoprene emission from mosses. Am. J. Bot. 86, 634-639.
43. Hébant, C. 1977 The conducting tissues of bryophytes. Bryophytorum Bibliotheca, vol. 10. Vaduz: Cramer.
44. Hedderson, T. A., Chapman, R. & Cox, C. J. 1998 Bryophytes and the origins and diversification of land plants: new evidence from molecules. In Bryology for the twenty-first century (ed. J. W. Bates, N. W. Ashton, & J. G. Duckett), pp. 65-77. Leeds: The British Bryological Society and Maney Publishing.
45. Hemsley, A. R. 1994 The origin of the land plant sporophyte: an interpolational scenario. Biol. Rev. 69, 263-273.
46. John, B. & Miklos, G. 1988 The eukaryote genome in development and evolution. London: Allen and Unwin.
47. Kenrick, P. & Crane, P. R. 1997a The origin and early diversification of land plants: a cladistic study: Washington, DC: Smithsonian Institution Press.
48. Kenrick, P. & Crane, P. R. 1997b The origin and early evolution of plants on land. Nature 389, 33.
49. Korall, P., Kenrick, P. & Therrien, J. P. 1999 Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. Int. J. Plant Sci. 160, 585-594.
50. Lang, W. H. 1907 On the sporogonium of Notothylas. Ann. Bot. 21, 201-210.
51. Lewis, L. A., Mishler, B. D. & Vilgalys, R. 1997 Phylogenetic relationships of the liverworts (Hepaticae), a basal embryophyte lineage, inferred from nucleotide sequence data of the chloroplast gene rbcL. Mol. Phyl. Evol. 7, 377-393.
52. Ligrone, R., Duckett, J. G. & Renzaglia, K. S. 1993 The gametophyte sporophyte junction in land plants. Adv. Bot. Res. 19, 231-317.
53. McCourt, R. M. 1995 Green algal phylogeny. Trends Evol. Ecol. 10, 159-163.
54. Maddison, W. P. & Maddison, D. R. 1992 MacClade: analysis of phylogeny and character evolution, 3.0. Sunderland, MA: Sinauer Associates.
55. Maden, A. R., Renzaglia, K. S. & Whittier, D. P. 1996 Ultrastructure of the spermatozoid of Lycopodiella obscurum (Lycopodiaceae). Am. J. Bot. 83, 419-429.
56. Maden, A. R., Whittier, D. P., Garbary, D. J. & Renzaglia, K. S. 1997 Ultrastructure of the spermatozoid of Lycopodiella lateralis (R.Br.) B. Ollgaard (Lycopodiaceae). Can. J. Bot. 75, 1728-1738.
57. Malek, O., Lättig, K., Hiesel, R., Brennicke, A. & Knoop, V. 1996 RNA editing in bryophytes and a molecular phylogeny of land plants. EMBO J. 15, 1403-1411.
58. Manhart, J. R. 1994 Phylogenetic analysis of green plant rbcL sequences. Mol. Phyl. Evol. 3, 114-127.
59. Manhart, J. R. 1995 Chloroplast 16S rDNA sequences and phylogenetic relationships of fern allies and ferns. Am. Fern J. 85, 182-192.
60. Mishler, B. D. & Churchill, S. P. 1984 A cladistic approach to the phylogeny of 'bryophytes'. Brittonia 36, 406-424.
61. Mishler, B. D. & Churchill, S. P. 1985 Transition to a land flora: phylogenetic relationships of the green algae and bryophytes. Cladistics 1, 305-328.
62. Mishler, B. D., Lewis, L. A., Buchheim, M. A., Renzaglia, K. S., Garbary, D. J., Delwiche, C. F., Zechman, F. W., Kantz, T. S. & Chapman, R. L. 1994 Phylogenetic relationships of the 'green algae' and 'bryophytes'. Ann. Mo. Bot. Gdn 8, 451-483.
63. Niklas, K. J. 1997 The evolutionary biology of plants. University of Chicago Press.
64. Nishiyama, T. & Kato, M. 1999 Molecular phylogenetic analysis among bryophytes and tracheophytes based on combined data of plastid coded genes and the 18S rRNA gene. Mol. Biol. Evol. 16, 1027-1036.
65. Ollgaard, B. 1987 A revised classification of the Lycopodiaceae. Opera Botan. 92, 153-178.
66. Oliver, M. J. & Wood, A. J. 1997 Desiccation tolerance in mosses. In Stress-inducible processes in higher eukaryotic cells (ed. T. Koval), pp. 1-26. New York: Plenum Press.
67. Pahnke, J., Goremykin, V., Bobrova, V., Troitsky, A., Antonov, A. & Martin, W. 1996 Utility of rDNA internal transcribed spacer sequences from the inverted repeat of chloroplast DNA in pteridophyte molecular phylogenetics. In Pteridology in perspective (ed. J. M. Camus, M. Gibby & R. J. Johns). Kew: Royal Botanic Gardens.
68. Paton, J. A. & Pearce, J. V. 1957 The occurrence, structure and functions of the stomata of British bryophytes. Trans. Br. Bryol. Soc. 3, 228-259.
69. Pirozynski, K. A. 1981 Interactions between fungi and plants through the ages. Can. J. Bot. 59, 1824-1827.
70. Pirozynski, K. A. & Malloch, D. W. 1975 The origin of land plants: a matter of mycotropism. Biosystems 6, 153-164.
71. Proskauer, J. 1960 Studies on Anthocerotales. VI. On spiral thickenings in the columella and its bearing on phylogeny. Phytomorphology 10, 1-19.
72. Pryer, K. M., Smith, A. R. & Skog, J. E. 1995 Phylogenetic relationships of extant ferns based on evidence from morphology and rbcL sequences. Am. Fern J. 85, 205-282.
73. Qiu, Y. L. & Palmer, J. D. 1999 Phylogeny of early land plants: insights from genes and genomes. Trends Pl. Sci. 4, 26-30.
74. Qiu, Y. L., Cho, Y., Cox, J. C. & Palmer, J. D. 1998 The gain of three mitochondrial introns identifies liverworts as the earliest land plants. Nature 394, 671-674.
75. Remy, W., Gensel, P. G. & Hass, H. 1993 The gametophyte generation of some early Devonian land plants. Int. J. Pl. Sci. 54, 35-58.
76. Renzaglia, K. S. 1978 A comparative morphology and developmental anatomy of the Anthocerotophyta. J. Hattori Bot. Lab. 44, 31-90.
77. Renzaglia, K. S. 1982 A comparative developmental investigation of the gametophyte generation in the Metzgeriales (Hepatophyta). Bryophytorum Bibliotheca, vol. 24. Vaduz: J. Cramer.
78. Renzaglia, K. S. & Bartholomew, S. E. 1985 Sporeling development in Fossombronia cristula Aust. with special reference to apical organization and growth. Bryologist 88, 337-343.
79. Renzaglia, K. S. & Duckett, J. G. 1987 Spermatogenesis of Blasia pusilla, from antheridial initial through mature spermatozoid. Bryologist 90, 468-501.
80. Renzaglia, K. S. & Duckett, J. G. 1988 Different developmental processes underlie similar spermatozoid architecture in mosses, liverworts and hornworts. J. Hattori Bot. Lab. 64, 219-236.
81. Renzaglia, K. S. & Duckett, J. G. 1989 Ultrastructural studies of spermatogenesis in the Anthocerotophyta. V. Nuclear metamorphosis and the posterior mitochondrion of Notothylas orbicularis and Phaeoceros laevis. Protoplasma 151, 137-150.
82. Renzaglia, K. S. & Duckett, J. G. 1991 Towards an understanding of the difference between the blepharoplasts of mosses and liverworts, and comparisons with hornworts, biflagellate lycopods and charophytes: a numerical analysis. New Phytol. 117, 187-208.
83. Renzaglia, K. S. & Garbary, D. J. 2000 Motile male gametes of land plants: diversity, development and evolution. Crit. Rev. Pl. Sci. (In the press.)
84. Renzaglia, K. S. & Maden, A. R. 2000 Microtubule organizing centers and the origin of centrioles during spermatogenesis in the pteridophyte Phylloglossum. Micros. Res. Tech. 49. (In the press.)
85. Renzaglia, K. S. & Vaughn, K. C. 2000 Anatomy, development and classification of hornworts. In The biology of bryophytes (ed. J. Shaw & B. Goffinet), pp. 1-19. Cambridge University Press.
86. Renzaglia, K. S. & Whittier, D. P. 1993 Spermatozoids of Phylloglossum drummondii are multiflagellated. Am. J. Bot. 80, 111. (Suppl.)
87. Renzaglia, K. S., Maden, A. R., Duckett, J. G. & Whittier, D. P. 1994 Monoplastidy in spermatogenesis of Lycopodium obscurum. Can. J. Bot. 72, 1436-1444.
88. Renzaglia, K. S., Rasch, E. M. & Pike, L. M. 1995 Estimates of nuclear DNA content in bryophyte sperm cells: phylogenetic considerations. Am. J. Bot. 82, 18-25.
89. Renzaglia, K. S., McFarland, K. D. & Smith, D. K. 1997 Anatomy and ultrastructure of the sporophyte of Takakia ceratophylla (Bryophyta). Am. J. Bot. 84, 1337-1350.
90. Renzaglia, K. S., Dengate, S. B. & Bernhard, D. L. 1998 Architecture of the spermatozoid of Selaginella australiensis. Am. Fern. J. 88, 1-16.
91. Renzaglia, K. S., Bernhard, D. E. & Garbary, D. J. 1999 Developmental infrastructure of the male gamete of Selaginella. Int. J. Pl. Sci. 160, 14-28.
92. Robbins, R. R. & Carothers, Z. B. 1978 Spermatogenesis in Lycopodium: the mature spermatozoid. Am. J. Bot. 65, 433-440.
93. Robert, D. 1974 Étude ultrastructurale de la spermiogenése, notamment de la différentiation de l'appareil nucléaire, chez le Selaginella kraussiana (Kunze) A. Br. Annales des Sciences Naturalles, Botanique, Paris, Série 12. 15, 65-118.
94. Roth, D. 1969 Embryo und Embryotheca bei den Laubmoosen. Eine histogenetische und morphologische Untersuchung. Biblioth. Bot. 129, 1 49. Pls. 1-9.
95. Roux, W. 1895 Gesammelte Abhandlungen zur Entwicklungsmechanik der Organismen. Leipzig: Engelmann.
96. Sack, F. D. & Paollilo, D. J. Jr. 1985 Incomplete cytokinesis in Funaria stomata. Am. J. Bot. 72, 1325-1333.
97. Schofield, W. B. 1985 Introduction to bryology. New York: Macmillan.
98. Schuster, R. M. 1966 The Hepaticae and Anthocerotae of North America, east of the hundredth meridian, vol. 1. New York: Columbia University Press.
99. Schuster, R. M. 1979 The phylogeny of the Hepaticae. In Bryophyte systematics (ed. G. C. S. Clarke & J. G. Duckett), pp. 41-82. London: Academic Press.
100. Schuster, R. M. 1984a Comparative anatomy and morphology of Hepaticae. In New manual of bryology, vol. 2 (ed. R. M. Schuster), pp. 760-891. Nichinan, Japan: Hattori Botanical Laboratory.
101. Schuster, R. M. 1984b Evolution, phylogeny and classification of the Hepaticae. In New manual of bryology, vol. 2 (ed. R. M. Schuster), pp. 892-1070 Nichinan, Japan: Hattori Botanical Laboratory.
102. Schuster, R. M. 1984c Morphology, phylogeny and classification of the Anthocerotae. In New manual of bryology vol. 2 (ed. R. M. Schuster), pp. 1071-1092. Nichinan, Japan: Hattori Botanical Laboratory.
103. Schuster, R. M. 1992 The Hepaticae and Anthocerotae of North America, vol. V. Chicago: Field Museum of Natural History.
104. Smith, D. K. & Davison, P. G. 1993 Antheridia and sporophytes in Takakia ceratophylla (Mitt.) Grolle: evidence for reclassification among the mosses. J. Hattori Bot. Lab. 73, 263 -271.
105. 2-concentrating mechanism and carbon isotope discrimination in lichens and bryophytes. Planta 198, 6-16.
106. Smith, E. C. & Griffiths, H. 1986b A pyrenoid-based carbon concentrating mechanism is present in terrestrial bryophytes of the class Anthocerotae. Planta 200, 203-212.
107. Smith, G. M. 1955 Cryptogamic botany. II. Bryophytes and pteridophyles, 2nd edn. New York: McGraw-Hill.
108. Soltis, D. E. & Soltis, P. S. 1998 Choosing an approach and an appropriate gene for phylogenetic analysis. In Molecular systematics of plants. II. DNA sequencing (ed. D. E. Soltis, P. S. Soltis & J. J. Doyle), pp. 1-42. Boston, MA: Kluwer Academic Publishers.
109. Soltis, P. S., Soltis, D. E., Wolf, P. G., Nickrent, D. L., Chaw, S.-M. & Chapman R. L. 1999 The phylogeny of land plants inferred from 18S rDNA sequences: pushing the limits of rDNA signal? Mol. Biol. Evol. 16, 1774-1784.
110. Southworth, D. & Cresti, M. 1997 Comparisons of flagellated and nonflagellated sperm in plants. Am. J. Bot. 84, 1301-1311.
111. Steinhauser, S., Beckert, S., Capesius, I., Malek, O. & Knoop, V. 1999 Plant mitochondrial RNA editing. J. Mol. Evol. 48, 303-312.
112. Stewart, W. N. & Rothwell, G. W. 1993 Paleobotany and the evolution of plants, 2nd edn. Cambridge University Press.
113. Taylor, T. N. & Taylor, E. L. 1993 The biology and evolution of fossil plants. Englewood Cliffs, NJ: Prentice-Hall.
114. van de Peer, Y, De Baere, R., Cauwenberghsm, J. & Wachter, R. 1990 Evolution of green plants and their relationships with other photosynthetic eukaryotes as deduced from 5S ribosomal RNA sequences. Pl. Syst. Evol. 170, 85-96.
115. Vaughn, K. C. & Harper, J. D. 1998 Microtubule organizing centers and nucleating sites in land plants. Int. Rev. Cytol. 181, 75-149.
116. Vaughn, K. C. & Renzaglia, K. S. 1998 Origin of bicentrioles in anthocerote spermatogenous cells. In Bryology for the twenty-first century (ed. J. W. Bates, N. W. Ashton & J. G. Duckett), pp. 189-203. Leeds: The British Bryological Society and Maney Publishing.
117. Vaughn, K. C., Campbell, E. O., Hasegawa, J., Owen, H. A. & Renzaglia, K. S. 1990 The pyrenoid is the site of ribulose 1-5-bisphosphate carboxylase/oxygenase accumulation in the hornwort (Bryophyta: Anthocerotae) chloroplast. Protoplasma 156, 117-129.
118. Vaughn, K. C., Ligrone, R., Owen, H. A., Hasegawa, J., Campbell, E. O., Renzaglia, K. S. & Monge-Najera, J. 1992 The anthocerote chloroplast: a review. New Phytol. 120, 169-190.
119. Vaughn, K. C., Sherman, T. D. & Renzaglia, K. S. 1993 A centrin homologue is a component of the multilayered structure in bryophytes and pteridophytes. Protoplasma 175, 58-66.
120. Waters, D. A., Buchheim, M. A., Dewy, R. A. & Chapman, R. L. 1992 Preliminary inferences of the phytogeny of bryophytes from nuclear-encoded ribosomal RNA sequences. Am. J. Bot. 79, 459-466.
121. Whittier, D. P. 1977 Gametophytes of Lycopodium obscurum as grown in axenic culture. Can. J. Bot. 55, 563-567.
122. Whittier, D. P. 1981 Gametophytes of Lycopadium digitatum (formerly L. complanatum var. flabelliforme) as grown in axenic culture. Bot. Gaz. 142, 519-524.
123. Whittier, D. P. 1983 Gametophytes of Ophioglossum engelmannii. Can. J. Bot. 61, 2369-2373.
124. Whittier, D. P. & Braggins, J. E. 1992 The young gametophyte of Phylloglossum (Lycopodiareae). Ann. Mo. Bot. Gdn 79, 730-736.
125. Whittier, D. P. & Peterson, R. L. 1984 Gametophytes of Botrychium lunarioides and their mucilage-coated rhizoids. Can. J. Bot. 62, 2854-2860.
126. Whittier, D. P. & Thomas, R. D. 1993 Gametophytes and young sporophytcs of Botrychium jenmanii in axenic culture. Int. J. Pl. Sci. 154, 68-74.
127. Whittier, D. P. & Webster, T. R. 1986 Gametophytes of Lycopodium lucidulum from axenic culture. Am. Fern J. 76, 48-55.
128. Wikström N. & Kenrick, P. 1997 Phylogeny of Lycopodiaceae (Lycopsida) and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. Int. J. Pl. Sci. 158, 862-871.
129. Wolf, P. G. 1997 Evaluation of atpB nucleotide sequences for phylogenetic studies of ferns and other pteridophytes. Am. J. Bot. 84, 1429-1440.
130. Wolf, P. G., Pryer, K. M., Smith, A. R. & Hasebe, M. 1998 Phylogenetic studies of extant pteridophytes. In Molecular systematics of plants. II. DNA sequencing (ed. D. E. Soltis, P. S. Soltis & J. J. Doyle), pp. 540-561. Boston, MA: Kluwer Academic Publishers.