CALIFORNIA STATE UNIVERSITY, NORTHRIDGE THE GRAMMITIDACEAE: ANATOMY OF ADENOPHORUS, XIPHOPTERIS AND GRAMMITIS A thesis submitted in partial satisfaction of the requirements for the degree of Master of Science in . Biology by Mary Jane Teiman June, 1981 The Thesis of Mary Jane Teiman is approved: Dr. Andrew Statrett Dr. William A. Emhoden Dr. Kenneth A. Wilson, Chairman California State University, Northridge ii DEDICATION This thesis is dedicated to A.B.T., who never doubted me, or questioned the importance of this work. It is also affectionately dedicated to the Fern Wizard. iii ACKNffiiLEDGEMENTS I am deeply grateful to Dr. Kenneth A. Wilson, my thesis committe chairman, for his friendship, advice and sincere encouragement which not only made this work possible but enjoyable. I am especially grateful to him for introducing me to comparative plant morphology and to ferns. I should also like to acknowledge Dr. William A. Emboden for his help and useful suggestions during my work on this project and his constructive criticism of my thesis drafts. I wish to thank Dr. Andrew Starrett for his careful reading and thoughtful criticism of this manuscript. Thanks are due to my Parents, for their patience, understanding and support throughout my college career. To Jim Thor, a special personal thanks for his patience, his strength, and for his gentle prodding which often kept me going when my spirits got low. I thank my friends; Susan Crowther, Bill Perry, Betty Rose, Mark Crase, Pam Friedman, Jim Mauch and Joan Parker for their unwaivering moral support. iv TABLE OF CONTENTS Page Dedication iii• Acknowledgements iv List of Figures and Tables vi Abstract xi I. Introduction 1 II. Materials and Methods 4 III. AdenoEhorus 6 AdenoEhorus tamariscinus 6 AdenoEhorus haalilioanus 18 AdenoEhorus hymenoEhylloides 28 IV. XiEhOEteris 36 XiEhoEteris serrulata 36 XiEhOEteris trichomanoides 46 v. Grarnmitis 56 Grammitis marginella 56 VI. Discussion 66 Literature Cited 76 v LIST OF FIGURES AND TABLES Page Plate I. AdenoEhorus tamariscinus Figure 1. Habit 15 Figure 2. Frond 15 Figure 3. Rhizome scales 15 Figure 4. Cross section of the stele of the rhizome 15 Figure 5. Glandular hairs of rhizome epidermis 15 Figure 6. Cross section of the stele of the petiole 15 Figure 7. Adaxial epidermal surface 15 Figure 8. Abaxial epidermal surface 15 Figure 9. Cross section of lamina 15 Figure 10. Epidermal hairs 15 Figure 11. Paraphysis 15 Figure 12. Cross section of the stele of the root 15 Plate II. Adenophorus tamariscinus Figures 1 - 25. Serial cross sections through the stele of the rhizome 17 Plate III. AdenoEhorus haalilioanus Figure 1. Habit 25 Figure 2. Frond 25 Figure 3. Portion of the frond showing details of the venation 25 vi Page Figure 4. Rhizome scales 25 Figure 5. Cross section of the stele of the rhizome 25 Figure 6. Glandular hair of the rhizome epidermis 25 Figure 7. Cross section of the stele of the petiole 25 Figure 8. Adaxial epidermal cells 25 Figure 9. Abaxial epidermal cells 25 Figure 10. Cross section of the lamina 25 Figure 11. Epidermal hairs 25 Figure 12. Paraphysis 25 Figure 13. Cross section of the stele of the root 25 Plate IV. Adenophorus haalilioanus Figures 1 - 19. Serial cross sections of the stele of the rhizome 27 Plate V. Adenophorus hymenophylloides Figure 1. Habit 33 Figure 2. Fronds 33 Figure 3. Rhizome scales 33 Figure 4. Cross section of the stele of the rhizome 33 Figure 5. Glandular hair of rhizome epidermis 33 Figure 6. Adaxial epidermal cells 33 Figure 7. Abaxial epidermal cells 33 vii Page Plate VI. Adenophorus hymenophylloides Figures 1 - 21. Serial cross sections of the stele of the rhizome 35 Plate VII. Xiphopteris serrulata Figure 1. Habit 43 Figure 2. Frond 43 Figure 3. Portion of the frond showing details of the venation 43 Figure 4. Rhizome scale 43 Figure 5. Cross section of the stele of the rhizome 43 Figure 6. Cross section of the stele of the petiole 43 Figure 7. Adaxial epidermal cells 43 Figure 8. Abaxial epidermal cells 43 Figure 9. Vestigial hydathode on adaxial laminar surface 43 Figure 10. Cross section of the lamina 43 Figure 11. Epidermal hairs 43 Figure 11. Cross section of the stele of the root 43 Plate VIII. Xiphopteris serrulata Figures 1 - 29. Serial cross sections of the stele of the rhizome 45 Plate IX. Xiphopteris trichomanoides Figure 1. Habit 53 Figure 2. Frond 53 viii Page Figure 3. Portion of the frond showing details of the venation 53 Figure 4. Rhizome scale 53 Figure 5. Cross section of the stele of the rhizome 53 Figure 6. Cross section of the stele of the petiole 53 Figure 7. Adaxial epidermal cells, hydathode 53 Figure 8. Abaxial epidermal cells 53 Figure 9. Cross section of the lamina showing vein ending in a hydathode 53 Figure 10. Sclerified epidermal trichome 53 a) origin from abaxial surface b) origin from adaxial surface c) sclerified trichome Figure 11. Epidermal hairs 53 Figure 12. Paraphyses 53 Figure 13. Cross section of the stele of the root 53 Plate X. Xiphopteris trichomanoides Figures 1 - 23. Serial cross sections of the stele of the rhizome 55 Plate XI. Grammitis marginella Figure 1. Habit 63 Figure 2. Fronds 63 Figure 3. Rhizome scales 63 Figure 4. Cross section of the stele of the rhizome 63 ix Page Figure 5. Cross section of the stele of the petiole 63 Figure 6. Adaxial epidermal cells 63 Figure 7. Abaxial epidermal cells 63 Figure 8. Cross section of the lamina 63 Figure 9. Epidermal hairs 63 Figure 10. Cross section of the stele of the root 63 Plate XII. Grammitis marginella Figures 1 - 23. Serial cross sections of the stele of the rhizome 65 Table 1. Comparison of Sporophyte Characteristics known for the Grammitidaceae 73 X ABSTRACT THE GRAMHITIDACEAE: ANATOMY OF ADENOPHORUS, XIPHOPTERIS AND GRAMMITIS by Mary Jane Teiman Master of Science in Biology The anatomy of six species of ferns belonging to the family Grammitidaceae was examined using standard paraffin and staining techniques. Adenophorus tamariscinus, Adenophorus haalilioanus and Adenophorus hymenophylloides are endemic Hawaiian species, related anatomically by the presence of glandular paraphyses, glandular rhizome epidermal hairs, concolorous scales, differentiated cortex and solenostelic, dorsiventral rhizomes. Xiphopteris serrulata and Xiphopteris trichomanoides from Jamaica, are characterized by the presence of hydathodes, concolorous scales, and erect, dissected solenostelic rhizomes. Grammitis marginella from Jamaica, noted for its sclerified blade margin, which was found to be superficial in origin xi was also found to have branched sclerified epidermal hairs, and a dictyostelic rhizome. Branches, which have not been described in the family were found in 5 species; branches are extra-axillary in A. tamariscinus, A. hymenophylloides, X. trichomanoides and G. marginella, and rare and random in stelar organization in X. serrulata. Root propagules known in A. haalilioanus are reported for the first time in X. serrulata. This study represents a significant increase in the number of Grammitid ferns examined for the anatomical characteristics that are important for the determination of phylogenetic trends and for solving of taxonomic problems within the family. xii I. INIRODUCTION The fern family Grammitidaceae consists of approxi­ mately 500 species of pantropical distribution. These ferns are primarily small, specialized epiphytes, found in montane cloud forests with low temperatures. Ferns belonging to this family have presented taxo­ nomic problems both in the past and present. Prior to the acceptance of a separate taxonomic category for these ferns, they were frequently classified as members of the comprehensive family Polypodiaceae, but usually with a note about their apparently aberrant characters (Copeland, 1947, for example). Authors still differ in the numbers of genera included in the family (Ching,l940; Holttum,l947, 1949; Copeland,l951). Stokey (1951) feels the gametophytes present strong evidence for the distinction between fern families. The gametophytes of the Grammitidaceae have been extensively studied by Stokey and Atkinson (1958), and are character­ ized by a prolonged and extensive filamentous stage which is distinct in morphology. The filaments are bead-like, with the increase in filament length due to unequal division of the terminal cell. In the Grammitid ferns, there is a strong tendency for these filamentous gamet­ ophytes to propagate vegetatively by fragmentation, where 1 2 the breaking of the filament occurs at modified cross walls. These characteristics are not known for any of the Polypodiaceae, or for any other of the higher ferns (Stokey and Atkinson,l958). Detailed morphological and anatomical studies of the sporophytes of the Grammitidaceae are few. Papers by Nozu (1958-1960) on Micropolypodium, Scleroglossum and Gramrnitis represent some of the earlier work. Wilson and Rickson (1966) described in detail the anatomy of Adenophorus sarmentosus. More recently, Bishop has suggested new revisions of the genera Adenophorus (1974) and Cochlidium (1978) based on morphological and brief anatomical details. In this study, the anatomy of six members of the Grammitidaceae is described. Three species of the endemic Hawaiian fern genus Adenophorus were chosen for study in order to add information needed to determine relationships within this possibly artificial genus (Wilson,l964), as well as for characteristics necessary for family definition. Two groups within the genus have been proposed (Wilson,l9- 64; Bishop,l974). Adenophorus tamariscinus and Adenophorus hymenophylloides are representative of one group, Adenophorus haalilioanus was chosen as representative of the second group. For contrast, three Jamaican Grammitid ferns are included in this study. Two of these species, Xiphopteris serrulata and Xiphopteris trichomanoides were examined for features which may be useful in determining 3 their interrelationship. The third, Grammitis marginella, is one of fourteen species within the family that are characterized by the presence of a sclerified blade margin. II. HATERIALS AND HETHODS The plant materials used in this study were collected as follows: Adenophorus haalilioanus (Brack.) K.