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Gametophytes of Homosporous Ferns B

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Gametophytes of Homosporous Ferns B THE BOTANICAL REVIEW VOL. 37 JULY-SEPTEMBER, 1971 No. 3 GAMETOPHYTES OF HOMOSPOROUS FERNS B. K. NAYAR 1 AND S. KAUR 2 INTRODUCTION ................................................................................................... 296 THE FERN PROTHALLUS .................................................................................. 298 SPORE GERMINATION .................................................................................... 299 PROTHALLIAL DEVELOPMENT ...................................................................... 303 MORPHOLOGY OF ADULT PROTHALLUS ....................................................... 307 Form of adult thallus ........................................................................................... 307 Prothallial trichomes ........................................................................................ 311 Sex organs ........................................................................................................... 316 VEGETATIVE PROPAGATION .......................................................................... 321 APOGAMY .................................................................................................................... 322 TAXONOMY ............................................................................................................ 323 Ophioglossidae ................................................................................................. 323 Marattiidae ..................................................................................................... 324 Osnmndaceae ................................................................................................. 325 Plagiogyriaceae .................................................................................................... 327 Schizaeaceae ............................................................................................................ 327 Anemiaceae .................................................................................................. 330 Parkeriaceae ................................................................................................ 331 Lygodiaceae ............................................................................................................ 332 Pteridaceae ....................................................................................................... 333 Cheilanthaceae .................................................................................................... 334 Adiantaceae ............................................................................................................ 336 Vittariaceae ..................................................................................................... 337 Cyatheaceae ......................................................................................................... 338 Dennstaedtiaceae ................................................................................................... 340 Hypolepidaceae ...................................................................................................... 340 Lindsaeaceae ................................................................................................... 341 Dryopteridaceae .................................................................................................... 342 Lomariopsidaceae ................................................................................................ 345 Oleandraceae ...................................................................................................... 346 Davalliaceae ........................................................................................................ 347 Aspleniaceae .......................................................................................................... 348 Thelypteridaceae .................................................................................................. 350 Grammitidaceae ..................................................................................................... 351 Blechnaceae ......................................................................................................... 353 Hymenophy]laceae .............................................................................................. 355 Loxsomaceae ................................................................................................ 359 Gleicheniaceae ............................................................................................. 360 1 Professor and Head of the Department of Botany, University of Calieut, Kerala (India). 2 National Botanic Gardens, Lucknow (India). 295 296 THE BOTANICAL 1AEVIEW Matoniaeeae .......................................................................................................... 362 Cheiropleuriaceae ................................................................................................... 363 Dipteridaceae ......................................................................................................... 363 Loxogrammaceae ............................................................................................... 364 Polypodiaceae .................................................................................................... 365 CONCLUSIONS ................................................................................................... 372 LITERATURE CITED .............................................................................................. 382 INTRODUCTION The pteridophytes have attracted the attention of botanists perhaps more than any other group of plants. This is mainly on account of the central position they occupy in the evolutionary history of the vegetable kingdom. Nevertheless they still remain one of the most controversial with respect to taxonomy and phylogeny. This is particularly so of the major group of pteridophytes, the ferns. This state of affairs is scarcely surprising when we consider that several major aspects of morphology and life history of most of the ferns are still little known, and that the ferns as a class do not offer any compact complex of characters (com- parable for example to floral morphology in Angiosperms) with which to assess relationships and interpret evolutionary progression. The search for a suitable complex of characters in this group is still progressing, and in this endeavor attempts are continuously being made to find new comparative criteria. Until recently (or perhaps even today) students of pteridophyte taxonomy and evolution were concerned only with the morphology of the sporophyte, the sporophyte being by far the dominant phase in the life history. The gametophyte of ferns has always been a neglected generation, ignored by morphologists and taxonomists alike. Even in the present century which has witnessed a tremendous upsurge in the study of morphology and taxonomy of the ferns, the gametophytic genera- tion has scarcely been looked upon as one worth serious consideration. Perhaps the unencouraging appraisal of the fern gametophyte by the greatest fern morphologist of the present century, Prof. F. O. Bower, discouraged attempts to use characters of the gametophyte in compara- tive studies. Bower, though acknowledging that gametophytes of the vast majority of ferns were unknown in his times and those of most of the others only imperfectly known, dismissed the gametophyte as morpho- logically unstable and thus undependable for use "in comparison with a view to classification of the ferns." He concluded "that its vegetative characters are deficient in stability and in variety of detail and conse- quently.., of minor importance for phyletic comparison" (Bower, 1923: 296). Though a wealth of information on the fern gametophyte has become available in the literature since Bower's days, no fern taxonomist has so far cared to re-examine the case of the fern gametophyte as an aid to comparative studies or paid much heed to the gametophyte. One NAYAR & KAUR: GAMETOPHYTES 297 reason for this lack of interest on the part of taxonomists could be that information on the gametophytes is too scattered in literature. Even as recently as 1949 the shadow of Bower's remarks could be detected in the assessment of the fern gametophyte by Prof. R. E. Holttum. Speak- ing of the gametophyte as an aid to taxonomy, he remarked: "Little has yet been published on the comparative morphology of the majority of ferns. There must be at least a considerable specialization to different habitats. In such a simple organism, such specialization may overshadow distinctive features that might indicate evolutionary lines." However, soon afterwards, Alma G. Stokey (1951), after a lifetime of work on fern gametophytes, most effectively presented a classical account dealing with the contribution of gametophyte morphology to the classification of the ferns. She proved beyond doubt that the fern gametophyte, though simple in structure when compared to the sporophyte, affords dependable criteria for taxonomic and phyletie studies. In the past 20 years since this publication, a vast amount of information has been added to our knowledge of the gametophytes of the ferns, more especially of the various groups of the advanced ferns, most of which were practically unknown to Stokey. This additional information serves materially to substantiate Stokey's conviction in the significance of gametophyte mor- phology for an understanding of the evolution and phylogeny of the ferns and in the characterization
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