BIOLOGY JUl 1 9 1982 The person charging this material is re- sponsible for its return to the library from which it was withdrawn on or before the Latest Date stamped below. Theft, mutilation, and underlining of books or* reasons for disciplinary action and may result in dismissal from the University. To renew call Telephone Center, 333-#4DO UNIVERSITY Q£ liJ.IUaii.J.IBRARY AT URBANA-CHAMPAIGN L161— O-1096 FIELDIANA Botany Published by Field Museum of Natural History New Series, No. 3 A MONOGRAPH OF CLASMATOCOLEA (HEPATICAE) JOHN J. ENGEL October 20, 1980 Publication 1311 A MONOGRAPH OF CLASMATOCOLEA (HEPATICAE) FIELDIANA Botany Published by Field Museum of Natural History New Series, No. 3 A MONOGRAPH OF CLASMATOCOLEA (HEPATICAE) JOHN J. ENGEL Donald Richards Associate Curator of Bryology Department of Botany Field Museum of Natural History October 20, 1980 Publication 131 1 Accepted for publication Feb. 2, 1979 Library of Congress Catalog Card No. : 80-66382 US ISSN 0015-0746 PRINTED IN THE UNITED STATES OF AMERICA Dedicated to DONALD RICHARDS TABLE OF CONTENTS ACKNOWLEDGMENTS 1 I. HISTORY 3 II. SOME METHODS AND TERMINOLOGY 5 III. TAXONOMIC CHARACTERS AND MORPHOLOGY 7 Slime papillae 7 Branching 8 Stems 10 Endophytic hyphae 11 Rhizoids 11 Leaves 11 Leaf cells and oil-bodies 12 Underleaves 13 Androecia 13 Bracts 14 Bracteoles 15 Perianths 15 Setae 17 Capsules 17 Spores 18 Elaters 19 IV. COMPARATIVE MORPHOLOGICAL DIVERSITY AND EVOLUTION OF SPOROPHYTE AND GAMETOPHYTE 19 Artificial key to taxa based upon sporophyte material 20 V. ORIGIN OF CLASMATOCOLEA AND SOME EVOLUTIONARY TRENDS 21 VI. GENERIC RELATIONSHIPS OF CLASMATOCOLEA 24 VII. PHYTOGEOGRAPHY 29 VIII. CONSPECTUS OF TAXA 38 IX. CLASMATOCOLEA 39 X. KEY TO TAXA OF CLASMATOCOLEA 40 XI. EXCLUDENDA 190 REFERENCES 198 SCANNING ELECTRON MICROGRAPHS 204 INDEX OF TAXA . . .225 LIST OF ILLUSTRATIONS Figures \. Diagrammatic representation of suggested major evolutionary lines in Clasmatocolea 22 2. Pictorial representation of generic relationships in Lophocoleaceae 25 3. Reconstruction of continental blocs at the end of the Triassic (180 m. y. BP) 32 4. Reconstruction of Permian positions of Gondwanaland continents 33 5. Reconstruction of continental blocs at the end of the Jurassic (135 m. y. BP) 34 6. Reconstruction of continental blocs at the end of the Cretaceous (65 m. y. BP) 36 7. Clasmatocolea rigens (Hook. f. & Tayl.) Engel 45 8. Clasmatocolea rigens (Hook. f. & Tayl.) Engel 46 9. Map of Clasmatocolea rigens (Hook. f. & Tayl.) Engel 47 10. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle 51 1 1 . Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle 52 12. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle 53 13. Map of Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle 54 14. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle 61 15. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle 62 16. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle 63 17. Map of Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle 64 1 8. Clasmatocolea marginata (Steph.) Grolle 70 19. Clasmatocolea marginata (Steph.) Grolle 71 20. Clasmatocolea marginata (Steph.) Grolle 72 21 . Clasmatocolea humilis (Hook. f. & Tayl.) Grolle 75 22. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle 76 23. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle 78 24. Map of Clasmatocolea humilis (Hook. f. & Tayl.) Grolle 79 25. Clasmatocolea fasciculata (Nees) Grolle 92 26. Clasmatocolea fasciculata (Nees) Grolle 93 27. Map of Clasmatocolea fasciculata (Nees) Grolle 94 28. Clasmatocolea vermicularis (Lehm.) Grolle 98 29. Clasmatocolea vermicularis (Lehm.) Grolle 100 30. Map of Clasmatocolea vermicularis (Lehm.) Grolle 106 31 . Clasmatocolea gayana (Mont.) Grolle 113 32. Clasmatocolea gayana (Mont.) Grolle 114 33. Map of Clasmatocolea gayana (Mont.) Grolle 115 34. Clasmatocolea ctenophylla (Schiffn.) Grolle 120 35. Map of Clasmatocolea ctenophylla (Schiffn.) Grolle 122 36. Clasmatocolea crassiretis (Herz.) Grolle 125 37. Map of Clasmatocolea crassiretis (Herz.) Grolle 126 Vll 38. Clasmatocolea moniliformis Engel 129 39. Map of Clasmatocolea moniliformis Engel 1 30 40. Clasmatocolea minutiretis Engel & Grolle 134 41 . Clasmatocolea minutiretis Engel & Grolle 1 35 42. Map of Clasmatocolea minutiretis Engel & Grolle 136 43. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle 140 44. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle v 142 45. Map of Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. ...' 145 46. Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle 148 47. Map of Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle 152 48. Clasmatocolea tjiwideiensis (Sande-Lac.) Grolle 156 49. Clasmatocolea puccioana (De Not.) Grolle 158 50. Clasmatocolea puccioana (De Not.) Grolle 160 51 . Map of Clasmatocolea puccioana (De Not.) Grolle 163 52. Clasmatocolea navistipula (Steph.) Grolle 166 53. Clasmatocolea navistipula (Steph.) Grolle 168 54. Map of Clasmatocolea navistipula (Steph.) Grolle 170 55. Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle 174 56. Map of Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle 175 57. Clasmatocolea verrucosa Engel 180 58. Clasmatocolea verrucosa Engel 181 59. Clasmatocolea cucullistipula (Steph.) Grolle 184 60. Clasmatocolea cucullistipula (Steph.) Grolle 185 61 . Map of Clasmatocolea cucullistipula (Steph.) Grolle 188 Plates 1. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. X 5,000 204 2. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. X 10,000 205 3. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. X4,700 206 4. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. X 9,500 206 5. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. X 5,000 207 6. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. X 10,000 207 7. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 6,600 208 8. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 9,700 209 9. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 28,600 210 10. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X46,000 211 11. Clasmatocolea vermicularis (Lehm.) Grolle. X5,000 212 12. Clasmatocolea vermicularis (Lehm.) Grolle. X 10,000 212 13. Clasmatocolea gayana (Mont.) Grolle. X3,655 213 14. Clasmatocolea gayana (Mont.) Grolle. X 10,260 214 15. Clasmatocolea ctenophylla (Schiffn.) Grolle. X 5,000 215 16. Clasmatocolea ctenophylla (Schiffn.) Grolle. X 10,000 215 17. Clasmatocolea minutiretis Engel & Grolle. X5,400 216 18. Clasmatocolea minutiretis Engel & Grolle. X 10,750 216 19. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. X5,400 217 20. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. X 10,750 218 21. Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle. X5,240 218 22. Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle. X 10,000 219 23. Clasmatocolea puccioana (De Not.) Grolle. X 6,075 219 24. Clasmatocolea puccioana (De Not.) Grolle. X 12,100 220 Vlll 25. Clasmatocolea puccioana (De Not.) Grolle. X25,300 221 26. Clasmatocolea navistipula (Steph.) Grolle. X 5,000 222 27. Clasmatocolea navistipula (Steph.) Grolle. X 10,000 222 28. Clasmatocolea cucullistipula (Steph.) Grolle. X 3,100 223 29. Clasmatocolea cucullistipula (Steph.) Grolle. X 10,200 224 ACKNOWLEDGMENTS The specimens of Clasmatocolea that I personally collected were gathered during several different expeditions to austral areas. Grateful acknowledgment is made to the National Science Foundation (grants GA-1192 and GV-26615 to Dr. Henry A. Imshaug, Michigan State University), which supported much of the author's field work. The final stages in this project were funded by NSF grant (BMS76-03616) to the author, which also supported field work in Tasmania. I acknowledge this support with thanks. I thank Dr. Imshaug not only for the several opportunities of collecting in south temperate and subantarctic regions, but for stimulating my interest in the area. I wish to acknowledge the Richards Foundation for its continuing support of Bryology at Field Museum over a period of many years. This welcome assistance supported research on this project, made possible the neotropical field work, and aided in innumerable other ways. I should also like to extend my thanks to Mr. Donald Richards for his encourage- ment and interest in my research. Field work in Chile in early 1976 was supported by the National Geographic Society; this support is gratefully acknowledged. It is impossible to thank all the people who assisted in field and labora- tory work, but I would, however, like to mention some individuals. I should like to acknowledge the kind help of Dr. Edgardo Oehrens B. for facilitating much of the field work in the Valdivian region in Chile in 1976, and Sr. Margarita Barrandeguy for arranging field work on Isla Chiloe, Chile. I thank Sr. Luis D. Gomez P. for facilitating field work in Costa Rica in 1973 and 1975. I am indebted to Ann Hollmann for her criticism and very able assis- tance in several stages of the project; to Virginia Beatty for her advice, criticism, enthusiasm, and friendship; to Dr. William Beatty for assistance with the Latin diagnoses; and to June B. Wehausen for typing several portions of the manuscript. 2 FIELDIANA: BOTANY I am grateful to Mr. Yale Factor for preparing several of the illustrations. Mr. William Peterson and Zorica Dabich assisted with a few plates. I extend special thanks to my wife Karen, who has assisted in many and various ways in preparation of the manuscript. I am grateful to her for her continued support and encouragement. My sincere thanks to the individuals and institutions listed below for assistance and for loan of specimens: Dr. O. Almborn (LD), Dr. Frances- co Bianchini (VER), the late Dr. C. E. B. Bonner (G), Dr. A. Bresinsky (Univ. of Regensburg; formerly at M), Mr. J. F. M. Cannon (BM), Dr. John Child (hb. Child), Dr. D. M. Churchill (MEL), Dr. Howard Crum (MICH), Dr. Sean R. Edwards (MANCH), Dr. Patricia Geissler (G), Dr. S. R. Gradstein (U), Dr. R. Grolle (hb. Grolle), Dr. A. V. Hall (BOL), Dr. E. Hegewald (hb. Hegewald), Prof. Dr. H. Hertel (M), Dr. Henry Imshaug (MSC), Dr. Hiroshi Inoue (TNS), Dr. Suzanne Jovet-Ast (PC), Dr. T. Koponen (H), Miss B. H. Macmillan (CHR), Dr. Robert E. Magill (PRE), Dr. Manuel Mahu (hb. Mahu), Dr. F. K. Meyer (JE), Dr. John T. Mickel (NY), Prof. Dr. J. Miege (G), Dr. Roland Moberg (UPS), Dr. Elsa Nyholm (S), Dr. A. E. Orchard (HO), Dr. Donald H. Pfister (FH), Dr. David A. Ratkowsky (hb. Ratkowsky), Dr. Clark T. Rogerson (NY), Mr. R. Ross (formerly at BM), Dr. R. Santesson (S; formerly at UPS), Dr. W. Schultze-Motel (B), Dr. R. M. Schuster (hb. Schuster), Dr. George A. M. Scott (hb. Monash Univ.), Dr. A. J. Sharp (TENN), Prof. Dr. Carlo H. Steinberg (FI), Dr. Anna-Elise Torkelsen (O), Dr. A. Touw (L), Dr. J. Vana (hb. Vana), Dr. C. Vanden Berghen (hb. Vanden Berghen), Dr. Daniel M. Vital (SP). I. HISTORY The genus Clasmatocolea has been misunderstood for the majority of time since its inception. The confusion over the circumscription and taxonomic position of the genus persisted until the work of Grolle (1960). Clasmatocolea was described by Spruce (1885) in his classic Hepaticae Amazonicae et Andinae. Spruce placed Clasmatocolea between Lopho- colea and Chihscyphus and stated (p. 440), "These curious little plants come very near Lophocolea. ..." He included three species in Clasmato- colea— C. cuneifolia, C. fragillima, and C. heterostipa, the first was a transfer from Jungermannia, and the latter two were new. Schiffner (1893), in his treatment of Hepaticae in Engler and Prantl's Die Naturlichen Pflanzenfamilien, placed Clasmatocolea between Leio- scyphus and Apothomanthus, and then followed the latter with Lophocolea. Schiffner recognized the same three Clasmatocolea species as Spruce. Stephani added C. chilensis in 1900. In his Species Hepaticarum (1906), he presented the following sequence of genera: Leioscyphus-Southbya- Arnellia-Gongylanthus-Clasmatocolea-Lophocolea. In the genus Clasma- tocolea, Stephani included six species, and much of the subsequent con- fusion over the identity of Clasmatocolea can be traced to the particular species he included. Stephani transferred Jungermannia doellingeri Nees to Clasmatocolea and cited the species first in his generic treatment. Stephani's concept of Clasmatocolea included various taxa such as C. exigua, which he regarded as similar to Jungermannia doellingeri. Further, his version of C. heterostipa is doubtful, since according to Grolle (1956), a specimen of it is not represented in the Stephani Herbarium. Stephani included elements of several of these diverse species, including Clasmato- colea ( = Chonecoled) doellingeri, in his generic diagnosis. Of the six species that Sephani included in Species Hepaticarum, four — C. doellingeri, C. exigua, C. truncdta, and C. fragillima — were subsequently found to belong to other genera in four different families. As Grolle (1956, p. 288) points out, "Vielleicht hat Stephani die Spruce'schen Arten von Clasmato- colea nicht gut gekannt." 4 FIELDIANA: BOTANY After Stephani treated Clasmatocolea in Species Hepaticarum, this genus was only incidentally dealt with during the next 50 years. Verdoorn (1932) included Clasmatocolea in the Epigonanthaceae, Evans (1939), in the Harpanthaceae, and Miiller (1939-40), in the Plagiochilaceae. In 1943 Herzog transferred Lophocolea flavovirens Steph. to the genus, and in the same year Frye and Clark placed Clasmatocolea in the Lophocoleoideae, near Chiloscyphus. Frye and Clark included only C. doellingeri and C. exigua and published a generic diagnosis consisting of a mixture of Clasmatocolea sensu Spruce and Chonecolea doellingeri. Miiller (1951-58) placed Clasmatocolea in the Plagiochilaceae, and Herzog & Noguchi (1955) published C. innovata. Until the 1950's, then, the genus was a heterogenous and quite misunderstood one, consisting of eight species, five of which were later shown to belong to as many different families. Grolle (1956) revised the genus and regarded Clasmatocolea as having but one species, C. heterostipa, which he described in full and illustrated. At this time, Grolle thought Clasmatocolea was distinguishable by the two-layered capsule wall (differing from Lophocolea, which Grolle stated had a four- to five-stratose capsule wall) and the wide, entire perianth mouth. In this paper, Grolle excluded a number of species and described two new genera. He also considered relationships of Clasmatocolea to both Harpanthaceae and Lophocoleaceae, but refrained from a familial placement; at the same time he raised the question whether these two families were independent. Two years later Schuster (1958) placed Clasmat- ocolea in the Harpanthaceae. Grolle (1959), in his treatment of African Lophocolea, did not recognize Clasmatocolea, but placed C. heterostipa in the synonymy of Lophocolea subintegra, thus giving it an amphiatlantic distribution. In 1960, Grolle reelevated the genus to generic rank, excluded the features he earlier chose to differentiate the genus, and distinguished Clasmatocolea by concave leaves coupled with anatomy of the inner capsule wall layer. Grolle found the generitype, C. heterostipa, related not to Lophocolea, as he thought a year previously, but rather to numerous austral species, most of which were then called Lophocolea. For these reasons, Grolle (1960) transferred 18 species to Clasmatocolea. In this paper, he placed Clasmatocolea in the Lophocoleaceae, and Schuster concurred in 1966. After 1960, seven more taxa were transferred to the genus by Grolle (1962, 1966, 1969, 1971, 1972b) and two by Engel (1973a). It is interesting to note that since 1900 no taxa had been described in Clasmatocolea (s. str.) until Engel (1979a and 1980). ENGEL: CLASMATOCOLEA 5 Engel (1978) treated the Brunswick Peninsula taxa, therein reduced a number of species to synonymy for the first time, and presented a key to the species of that area. Taxa (including varieties) here included in Clasmatocolea (i.e., the genus in the strict sense) were described in a number of different genera. This wide array of genera underscores the degree of confusion that existed over the taxonomic position of these plants. Of the 20 names in Jungermannia, 14 later were transferred to Lophoco/ea, so that there existed a large pool of names under Lophocolea, with the remainder under a wide variety of miscellaneous genera. The basionyms are organized as follows: Genus of Number of Described Range of Percent basionym taxa mostly by publication of total Lophocolea 42 Stephani (25), Herzog (9), 1885-1958 50 Schiffner (3), Massalongo (3) Jungermannia 20 Hooker f. & Taylor (11), 1820-59 23.8 Lehmann (3), De Notaris (2) Clasmatocolea 7 Spruce, Stephani, Engel 1885-1980 8.3 Chiloscyphus 6 Various 1851-1915 7.1 Jamesoniella 1 Stephani 1911 1.2 Leioscyphus 1 Pearson 1922 1.2 Notoscyphus 1 Mitten 1877 1.2 Nardia 1 Stephani 1897 1.2 Alicularia 1 Herzog 1921 1.2 Conoscyphus 1 Pearson 1922 1.2 Blepharostoma 1 Gola 1923 1.2 Odontoschisma 1 Sim 1926 1.2 Isotachis 1 Stephani ex Herzog 1952 1.2 Total 84 II. SOME METHODS AND TERMINOLOGY All information pertaining to the vegetative plant parts, unless otherwise stated, was obtained from the main axis. The axis width was made by measuring the main axis, ventral side down, without altering the leaf direction, i.e., without spreading the leaves. Axes that could not be bal- anced for such a view were maintained in that position by laterally bolster- ing them. Underleaf vs. stem width was obtained by dissecting underleaves, which were then flattened, measured, and compared to the stem width. Leaf cell trigone size and terminology follows Schuster (1966). A few reagents were used. Endophytic hyphae were stained with the use of cotton blue. Ammonium hydroxide was used as a clearing agent 6 FIELDIANA: BOTANY for antheridia and setae when cells were obscured by enclosed foodstuffs. For all American taxa, capsules from several different collections were selected, and their spores were prepared for scanning electron microscopy. Spore micrographs were made from several of these different capsules and later compared. Further, under the SEM, numerous spores were examined to determine if spore variation was present in that capsule. The spore micrographs, found on pages 204 to 224, represent those typical of the species. Spores, in some cases, were prepared using a standard Critical Point Drying technique. Micrographs were taken using a Cambrid ge Stereoscan S-4 microscope located at Field Museum. Author citations follow the abbreviations in Sayre et al. (1964). Journals are abbreviated according to Botanico-Periodicum-Huntianum (B-P-H). Those authors and journals not in these references are abbreviated in a similar style. Herbarium citations follow Index Herbariorum (Holmgren & Keuken, 1974). The terms Magellanian and Valdivian are frequently found in phyto- geographical statements of species. These terms are defined in the section on phytogeography (pp. 29-30). The expression Patagonian Channels, how- ever, is not defined elsewhere. I have used this term to indicate the highly dissected portion of western Chile south of Puerto Montt to and including the Brunswick Peninsula (i.e., latitudes 41 ° 28' S. to 53° 30' S.). It includes the western portion of the mainland and the channels near Puerto Natales, S. Skyring, S. Otway, etc. The terms West Patagonia and Andean Patagonia are used according to the following definitions given by Skottsberg (1910, 1916) and Moore (1968): West Patagonia — The region including the west slope of the Andes to the Pacific Ocean, characterized by high precipitation and supportive of lush rain forests and magellanian moorland. Andean Patagonia — The eastern slope of the Andes from base to snowline. This region experiences moderate rainfall and supports a deciduous, comparatively dry forest. Hueck (1966) recognizes two forested regions that extend to Andean Patagonia, the Northern Nothofagus forests, which are composed pre- dominantly of two species of deciduous Nothofagus, N. obliqua and N. procera, and the Araucaria-Libocedrus zone. The localities are cited in the manner provided in the following gazet- teers: South Africa (Leistner & Morris, 1976); New Zealand (New Zealand Dept. of Lands and Survey, 1968, and U.S. Board on Geographic Names, 1954); and Peru, Australia, Brazil, Colombia, Chile, Argentina (U.S. Office of Geography, 1955, 1957, 1963, 1964, 1967, 1968, respectively). Abbreviations of localities follow the American Geographical Society ENGEL: CLASMATOCOLEA 7 index to map of Hispanic America (see American Geographical Society of New York. Index . . . , 1945, pp. 10-11) and are as follows: A. — Arroyo I. — lie, Isla, Island B. — Bahia, Bay L. — Lago, Lake Bo. — Boca La. — Laguna Br. — Brazo M. — Monte Cta. — Caleta Mt. — Mount, Mountain Can.— Canal P.— Peak C. — Cerro Pen. — Peninsula Cord.— Cordillera Pto.— Puerto Cr.— Creek Pta.— Punta Ens. — Ensenada Q. — Quebrada Esto. — Estero, Estuario R. — Rio, River F. — Fiordo S. — Seno Fo. — Fondeadero Sa. — Sierra G. — Gulf Vo. — Ventisquero Hr.— Harbor V.— Volcan NOTE: Add s to form plural. Several of the maps were made from tracings by the author. The south polar projection was adapted from a National Geographic Society map (1943). The nondetailed map of southern South America and the world projection map were adapted from the Goode Base Map Series, University of Chicago. The two different types of detailed maps of southern South America were adapted from several U.S. Naval Oceanographic Office charts. III. TAXONOMIC CHARACTERS AND MORPHOLOGY The genus Clasmatocolea has a great abundance of taxonomic criteria. For the most part, traits are expressed in several or all taxa, either charac- terizing a distinct group or the entire genus. However, at least in my experience, Clasmatocolea is unique — I have not previously encountered a genus of hepatics with this magnitude of single exceptions to a general pattern of morphology. There are, in fact, ten instances of a morphological feature present in only one species of the genus. Although I cannot account for this phenomenon, it does appear that there has been considerable experimentation in the evolutionary history of Clasmatocolea. These deviations will be pointed out in the following discussions of the various morphological characters. Slime papillae. — In Clasmatocolea, slime papillae are very common and 8 FIELDIANA: BOTANY widespread in their occurrence. They are frequently associated with under- leaves, bracts and bracteoles, terminating segments and armature, or sessile on the apex or lamina margins. I have not found slime papillae on leaves. In undivided underleaves, they are commonly present in the position of segments or teeth, representing vestigial lobes or teeth. In this connection I have seen, in several taxa, regular reduction sequences of underleaf apices, from segments to teeth to remnant slime papillae. The slime papillae on the underleaves serve to protect the apical meristem from desiccation, whereas those associated with bracts, bracteoles, and perianth lobes likely protect young gynoecia from desiccation during their develop- ment. Slime papillae are also common on lobules of androecial bracts where they prevent desiccation of the developing and mature antheridia as well as provide moisture important for survival of the spermatozoids, particularly since the androecia are so often intercalary in this genus. Branching. — Within Clasmatocolea, branching is of major taxonomic and phylogenetic significance. From the outset of this study I have been fascinated by the interrelationships and specialization of branching pat- terns in this genus. My study of branch types is based upon detailed examination of numerous different collections of each species. Of particu- lar interest is the association, to a rather significant degree, of generalized branching patterns with more primitive taxa and specialized branching with more derived species. In Table 1, I have arranged taxa with regard to branching patterns, from the primitive condition of general, less restricted branch types to the advanced one, with localization and restriction of branch types. If one compares this table with the "Conspectus of Taxa" (p. 38), which is arranged phylogenetically, there is considerable correlation (see also fig. 1, p. 22). For example, within Subg. Clasmatocolea, the very general branching pattern of C. marginata correlates with the polystraty and four- stratose antheridial stalks, features I regard as primitive in the genus. Other examples may be drawn from Sect. Clasmatocolea. Clasmatocolea moniliformis has almost entirely ventral-intercalary branching and, at the same time, has a specialized leaf insertion unique to the genus, plus consistently undivided underleaves. Clasmatocolea crassiretis has a deriva- tive branching pattern, being totally without terminal branching; the species has several other derivative features, a few of which are : (a) small, undivided underleaves; (b) terete perianths, which have virtually lost all trace of keels or wings ; and (c) nodular or maximally protuberant trigones. Clasmatocolea fasciculata is an interesting paradox in branching. Although the species has the most generalized branching mode, even possessing Acromastigum-typc branching (which to my knowledge is unique in ENGEL: CLASMATOCOLEA TABLE 1. Branching patterns in Clasmatocolea, arranged with taxa having more generalized, less restricted branch types (top) to taxa with localization and restriction of branch types. Taxon C. fasciculata C. marginata C. ctenophylla C. fulveUa C. humilis C. trachyopa C. strongylophylla C. rigens C. verrucosa C. vermicularis C. navistipula C. puccioana C. gayana C. obvoluta C. cucullistipula C. moniliformis C. notophylla C. crassiretis C. minutiretis C. tjiwideiensis Branch type Terminal Intercalary Frullania Acromastigum Ventral Lateral . tjiwideiensis 0 0 + + 0 0 = branch type absent; + = branch type rare; ++ = branch type occasional; + + = branch type common. Lophocoleaceae), it has gynoecia restricted to short, ventral-intercalary branches, a markedly derivative feature. Clasmatocolea minutiretis, which has no terminal branching and only occasional lateral-intercalary branch- ing, has several other derived/specialized features, among them the most reduced underleaves of the section, and the specialized, laterally fused androecial bracts. Other subgenera show equally intriguing correlations. Clasmatocolea cucullistipula (Subg. Plicaticalyx, see fig. 1), for example, has somewhat specialized branching, with predominantly Frullania-type and only occa- sionally ventral-intercalary type, but totally lacks the lateral-intercalary type. I regard this species as the most derived of the genus; it has, among several other specialized features: (a) contracted, plicate perianths that are often subterete toward the base and (b) androecia wider than the main axis. Clasmatocolea notophylla (Subg. Squamicalyx, fig. 1) commonly has 10 FIELDIANA: BOTANY intercalary branches of both kinds but lacks terminal branching; it also has several other specialized features, a few being (a) underleaves and bracteoles connate on both sides and (b) scaly, terete perianths without keels or wings. There are other forms of branch specializations within the genus. Clasmatocolea fulvella (Subg. Clasmatocolea), for example, has a system of short, abbreviated intercalary branches that are vermiform and sub- moniliform and appear quite differentiated from the main axis (fig. 43-1). Other patterns may be observed in Subg. Metaclasmalocolea. In one species, C. puccioana, the copiously produced lateral-intercalary branches are highly differentiated — being usually abbreviated and of limited growth and having underleaves inconspicuous, reduced, and scalelike (fig. 49). In one variety of the other species of this subgenus, C. navistipula var. navistipula, a flabellate branching pattern has developed (fig. 53). These two species fall adjacent to one another in Table 1, both having nearly identical branching types, with lateral-intercalary type common, ventral- intercalary rare or occasional, and terminal branching rare. Stems. — The stem anatomy is quite species specific and is usually a useful supplementary character. Clasmatocolea taxa may be divided into four groups based upon stem anatomy: Group 1. — Stems with moderate to distinct differentiation, having a cortex of l-2(-3) rows of thick-walled cells equal to or smaller than the medullary. The stems are 5-18 cells high. Eight taxa are in this group; examples are C. fasciculata (fig. 26-5), C. puccioana (fig. 49-4), and C. fulvella (fig. 43-15). None of the diminutive taxa of the genus fall here. Group 2. — Stems with moderate to distinct differentiation, having a cortex of one row of thick-walled cells larger than the medullary. All of the species are diminutive for the genus, and all have stems between four and seven cells high. All species referable here are of Sect. Clasmatocolea: C. strongylophylla (fig. 46-5, 6), C. crassiretis (fig. 36-10), and C. ctenophylla (fig. 34-14). Group 3. — Stems with virtually no internal differentiation and consisting of very thick-walled cells. The stems are 6-9 cells high. Here belong C. gayana (fig. 31-9), C. minutiretis (fig. 40-7), C. tjiwideiensis (fig. 48-10), and C. cucullistipula (fig. 57-3). Group 4. — Stems with the slightly to distinctly thickened cortical cells variable: smaller, equal to, or larger than the medullary cells and thus not fitting into any of the above groups. Species in this group commonly have a smattering of several large and small cells in both cortex and medulla. The stems are 5-10 cells high. Belonging here are species from four sub- ENGEL: CLASMATOCOLEA 11 genera — C. rigens (fig. 7-5), C. trachyopa (fig. 14-5), C. moniliformis (fig. 38-10), and C. navistipula (fig. 52-30). Endophytic hyphae. — Fungal hyphae, at least some of which are of basidiomycetes (verified by the presence of clamp connections), are present in a number of Clasmatocolea species. When present, the endo- phytic hyphae occur in both cortical and medullary cells of the stem. This feature does not seem to be of taxonomic significance. Rhizoids. — Within the genus Clasmatocolea, rhizoids are rather uniform, but several points of interest deserve mention. As in other Lophocoleaceae, rhizoids are in fascicles from the stem near the underleaf base. However, in addition to this position, C. tjiwideiensis also has them from the stem adjacent to the ventral base of leaves, and, in C. cucullistipula, the rhizoid field includes the median basal portion of the underleaf lamina (fig. 57-1). In C. gayana, rhizoids issue from a raised, flat, disciform rhizoid initial field (fig. 31-8 and p. 1 16 for discussion), a feature I find useful in assisting identification of the species. The rhizoid tips are simple or branched and are colorless, except in C. tjiwideiensis, where they are, remarkably, ma- genta.1 At least in some instances, the rhizoid tips branch in "response to contact with a solid substratum" (see Odu & Richards, 1976, for a discus- sion of this phenomenon in Lophocolea cuspidatd). Leaves. — Compared with many other groups of hepatics in which the major focal point of evolution seems to have been foliar structures, the leaf in Clasmatocolea is at best of moderate phylogenetic and taxonomic significance. Pronounced exceptions are taxa with divided leaf apices, namely those taxa in Subg. Protoclasmatocolea and Subg. Lacerifolia. Within these two allied subgenera, there is a trend that reaches a culmina- tion in degree of complexity in C. trachyopa. No other species in the genus, for example, has the intricacy of folding and armature; no other species has three-lobed leaves or accessory lobes at the leaf apex and margins (see figs. 14, 15). In the remaining bulk of taxa, i.e., those with undivided leaf apices, the leaf, though often distinctive for the species, shows remarkably little morphological diversity. Several taxa, in an array of diverse sections and subgenera, have leaves distinctly conchiform concave, oblate to wide ovate (and variations of these) in shape, entire, with the dorsal margin straight (at least in proximal half), and the ventral margin broadly rounded. However, within the undivided leaf group, generally when the leaf is armed it is of greatest taxonomic use. For example, the sinuate main axis 1 See Schuster & Engel (1977) for a discussion of the taxonomic value of claret-red (magenta) vs. colorless rhizoids in the genus Schistochila. 12 FIELDIANA: BOTANY leaves of C. fulvella (fig. 43-1), the distinctive position of the teeth in C. tjiwideiensis (fig. 48), the sharp, often strongly incurved teeth of C. minutire- tis (fig. 40), and notably, the dentition of the beautiful, diminutive, C. ctenophylla (fig. 34). Leaves of Clasmatocolea are unistratose throughout in all species save one — in C. marginata, polystraty has developed in the leaf margins and the median-basal portion of the leaves. Leaf cells and oil-bodies. — The form of the leaf cells in Clasmatocolea is somewhat variable. Leaf cells are thin to moderately thickened except in C. moniliformis where walls are distinctly thickened. Even though trigones for the most part are large and bulging to occasionally coarse and nodular, the deviations from this condition are not only taxonomically significant but possibly of some evolutionary significance. Minute trigones occur in the simple, most primitive member of the genus, C. rigens. Further, small trigones are also present in C. marginata, which likewise has several primitive features. The only other instance of trigones being absent or small is in C. vermicularis; this condition is useful, incidentally, in separat- ing this species from all other members of Sect. Clasmatocolea. In four, mostly unrelated, derivative taxa — C. crassiretis, C. notophylla, C. ver- rucosa, and C. cucullistipula — the trigones are massive, as well as nodular and protuberant to the extent that the cell lumen is bounded, except for a few narrow, thin-walled places, mostly by trigones (cf. figs. 36-9, 55-9, 57- 2). I regard this condition as derivative. The development and consistency of trigones shows a sharp divergence from the genus Lophocolea. It ap- pears that this trend has gone from absent or minute trigones to the mas- sive condition just discussed. The cuticle is mostly smooth or slightly roughened, but has been used as a criterion for Sect. Strongylophyllae, which has coarse, hemispherical- oblong papillae (fig. 46-7). Clasmatocolea verrucosa is unique in possessing huge, domelike verruca, one per cell. In general, cell size is of minor importance. In Clasmatocolea, although I have used cell size as an aid in separating "look-alikes," I have not used cell size as a key character. Oil-bodies are known for six of the species. In general, they are of the grape-cluster type, with the surface appearing granular or papillose or botryoidal, and are consistently 2 or 2-3 per cell in the leaf middle and occupy only a small fraction of the cell lumen. Clasmatocolea moniliformis is a noteworthy deviant, having oil-bodies occupying 0.20-0.75 of the cell lumen but with them absent from marginal rows of leaf cells. All data of oil-bodies are from personal observations, except that for C. vermicularis, which I have obtained from Gradstein et al. (1977). ENGEL: CLASMATOCOLEA 13 Underleaves. — The majority of species of Clasmatocolea have very characteristic underleaves that are of high taxonomic value. Many and various underleaf characters have been utilized taxonomically in the genus. The underleaves vary from the highly reduced, plane, deeply bifid individuals, which are maximally 0.6 X the stem width in C. minutiretis, to the large, cucullate, undivided structures, which are maximally 6.7 X the stem width in C. moniliformis. Many intermediates, with various combi- nations of characters, fall within these extremes. The underleaf width vs. the stem width, a comparatively stable feature, has frequently been used ; the following are some examples : (a) C. minutire- tis (which culminates a reduction sequence in Sect. Clasmatocolea) has underleaves only 0.35-0.6 X the stem width; (b) C. vermicularis has under- leaves normally slightly narrower than the stem, a very useful species criterion; and (c) C. moniliformis, with the very large underleaves men- tioned above. In this connection, mention should be made of the highly reduced, scalelike underleaves of the main axis and/or branches of Subg. Metaclasmatocolea; underleaves of C. puccioana and C. navistipula here provide critical criteria pertinent at the subgeneric level. Convexity and associated manifestations are a useful tool, helping to define many taxa, a few of them being (a) C.fulvella, with inflated, orbicu- lar cucullate branch underleaves (fig. 43); (b) C. moniliformis and C. notophylla, each with distinct cucullate underleaves (fig. 38); and (c) C. cucullistipula, with cucullate, auriculate underleaves (fig. 57). Very different from this condition are the plane underleaves of C. crassiretis (fig. 36) and o f C. minutiretis (fig. 40). Lamina armature is useful in circumscribing and identifying such species as C. fasciculata, with its single, large lacinium (fig. 25), and C. cucullistipula, with its single, large lobe extending above the dorsal stem surface (fig. 57). In nearly all species, the underleaves are either undivided or bifid to less than 0.5, but in C. minutiretis they are bifid to 0.7, and in C. rigens they are usually bifid to nearly the base. In the majority of taxa, the underleaves are free or connate on one side, but in four mostly unrelated species they are connate on both sides — C. obvoluta, C. tjiwideiensis, C. notophylla, and C. \errucosa (cf. fig. 55). Underleaves are usually transversely inserted, but the frequently oblique insertion in C. rigens is usually the first character I search for in identifying this species. Androecia. — Within Clasmatocolea, androecia are strongly differentiated from leaves. The bracts are distinctly saccate or ventricose in the basal 14 FIELDIANA: BOTANY half, whereas the distal portion is strongly concave. The androecial region, then, is markedly narrower than neighboring sterile portions of the axis, except in one species, C. cucullistipula, in which the androecial region is distinctly wider (fig. 58-8). The specialized, certainly derivative, laterally fused androecial bracts are exceptional in the genus and occur in three species (figs. 14-6 and 41-9). They are present in both species of Subg. Lacerifolia, where I have used the character taxonomically, not only as an aid to circumscribing the subgenus but in separating it from the allied Subg. Protoclasmatocolea. Fused bracts appear again in the quite unrelated C. minutiretis, a derivative species in Subg. Clasmatocolea. Compared to Lophocoleoid androecial bracts I have studied, the lobules in Clasmatocolea are large in proportion to bract size (see p. 83). In all species but one, the lobule is a flap of tissue formed by dilation and infolding of the dorsal base of the bract (figs. 28-15, 41-10). The exception is C. navistipula, which has conduplicately bilobed bracts (fig. 53-10, 11), a condition quite unusual for hepatics. The lobules in Clasmatocolea are usually incurved to involute, with the margin entire or sparingly dentate with few-celled teeth and/or with one lacinium or a single, commonly involute lobe; lobule margins nearly always have slime papillae. The armature of C. cucullistipula is somewhat unusual in having 1-3 laciniae or small lobes (fig. 58-9, 10). The antheridia are solitary, only rarely occurring two per bract. An- theridial stalks are uniseriate throughout or commonly with localized bistratose areas (figs. 12-5, 28-11); in no case are they bistratose through- out. Stalks are four-seriate in C. marginata (fig. 18-12). Bracts. — With the exception of size, the bracts do not deviate to any major extent from the leaves. Those species with divided leaves also have divided bracts (bifid, except in C. trachyopa, where often trifid), whereas those taxa with undivided leaves have undivided bracts. However, bracts are occasionally short-bifid in C. humilis and C. gayana, species in which the leaves are always undivided. With the conservative nature of the gynoecium in mind, perhaps the presence of at least some bifid bracts in taxa with undivided leaves is meaningful in a phylogenetic context, indi- cating a bifid-leaved ancestry. Almost without exception, taxa with armed leaves have armed bracts, whereas those with entire leaves have entire bracts. For the most part, the amount and magnitude of armature is expressed to a greater extent on bracts as compared to leaves (figs. 58-1 and 41-1). Armature is commonly ENGEL: CLASMATOCOLEA 15 expressed on the bract apex and upper margins and to a considerably lesser extent on their median and basal portions. Bracteoles. — The innermost series of bracteoles are conspicuous, con- siderably larger than, and, in most instances, clearly differentiated from, underleaves. These features of the bracteole at once set the genus Clasmat- ocolea apart from the genus Hepatostolonophora. Bracteoles are differentiated from underleaves by modification of several features, as outlined below: Size. — In nearly all cases, bracteoles are markedly larger than under- leaves. In fact, in seven instances, bracteoles are subequal to the bracts in size, even though these species have a considerable gap in leaf and underleaf size. Apical division. — In nearly all species with divided underleaves, the bracteoles are less deeply incised or even undivided. This reduction is often accompanied by an alteration in shape and magnitude of segments. Armature. — In nearly all species, the amount and magnitude of bracteole armature is significantly increased over that of the underleaves. Convexity. — Underleaves are, in ventral view, plane or convex, but in many instances, the bracteoles are, in ventral view, concave to canaliculate (see fig. 23-5). Connation. — With one exception, bracteoles are free from bracts even though, in many cases, the underleaves are connate with leaves on one side. Absence of bracteole connation is likely due to secondary meristematic activity in the region of the perianth, thus spacially dis- placing bracteole and bract. The exception noted above is C. notophylla, which has bracteoles (conspicuously) connate with the bracts on both sides. See plates of C. minutiretis (figs. 40, 41) for illustrations of most of these modifications. Perianths. — This structure, including position, shape, and lobe features, is of considerable taxonomic significance within the genus Clasmatocolea. Perianths are strongly trigonous to distinctly terete and have three broadly rounded, entire or variously armed, sometimes bifid lobes, the latter occurring only within the two most primitive subgenera (see discussion below). In general, as to be expected, those species with entire leaves will have entire perianth lobes, whereas those plants with armed leaves will have armed perianth lobes. Usually the character of armature will be expressed to a greater and more reliable extent on perianth lobes than on leaves. Several species may be immediately recognized by their taxo- nomically useful, characteristic perianth mouth and lobes; a few examples 16 FIELDIANA: BOTANY follow. Clasmatocolea vermicularis has perianth mouths wide and open and lobes entire2 and often deeply cleft (figs. 28-16, 29-2, 4); C. gayana has perianths with the mouth region bilaterally compressed and has lobes densely denticulate (figs. 31-1, 32-5, 6); C. ctenophylla has numerous, 1-2- celled, sharp teeth that are more or less equal in size (fig. 34-15, 16); and C. cucullistipula is unique in having perianths contracted at the mouth (fig. 58-1) and plicate toward the apex (fig. 58-1, 4, 5). The trigonous, "Epigoniantheae"-type of perianth is generally regarded to have arisen from a terete perianth or "urn-shaped cup" (see Schuster, 1966, incl. fig. 52, and Fulford, 1965). Within Clasmalocolea, the primitive condition is characterized by a sharply trigonous perianth with each of the three lobes bifid, reflecting the nature of the foliar structures from which they originated. Further, the points of fusion of the leaves are frequently delineated by the presence of wings (fig. 31-1, 2), which are not only a remnant, but a reminder, of the origin of the perianth from leaves. The combination of strongly trigonous perianths with bifid lobes and winged keels occurs only in the two most primitive subgenera — Subg. Protoclasmatocolea and Subg. Lacerifolia. The question then arises of the occurrence and significance of terete perianths in Clasmatocolea. In all of the more advanced taxa of the genus (i.e., all taxa except Subg. Protoclasmatocolea and Lacerifolia}, the perianth lobes are broadly rounded, the bifid condition having been lost. Further, in some taxa there has been a loss of distinct keels and, at the same time, of wings, and it is in these taxa that the terete perianth condition has developed. This leads me to the conclusion that the terete condition in Clasmatocolea is a secondarily derived one. Terete perianths occur in such derivative taxa as C. notophylla (Subg. Squamicalyx, fig. 55-2), C. cucul- listipula (Subg. Plicaticalyx, fig. 58-1, 6), C. strongylophylla (Sect. Strongy- lophyllae), and C. crassiretis (Sect. Clasmatocolea, fig. 36-3). A subterete condition also occasionally occurs in C. vermicularis (Sect. Clasmatocolea, fig. 28-16). It should be brought out that, for the most part, the occurrence of terete perianths is linked with a substantial reduction in size of the ventral lobe (fig. 36-3), another feature supportive of the derivative nature of the terete perianth. The reduction of the ventral perianth lobe mentioned above has some- times been accompanied by a suggestion of bilateral compression. This 2Grolle (1956) recognized only a single species in Clasmatocolea, C. heterostipa ( = C. vermicularis), and used the wide, entire perianth mouth of that species to assist in distinguishing the genus. Later, in Grolle (1960), however, the genus assumed a much larger size, and this character became meaningless. ENGEL: CLASMATOCOLEA 17 kind of condition is of some significance, for it suggests the connection to the Leptoscyphus type of perianth (see Schuster, 1966, p. 552) and supports, along with several other features, a relationship between Clas- matocolea and Leptoscyphus (see also p. 26). In the majority of Clqsmatocolea taxa, there is a well-developed and distinct perianth along with a true calyptra, without development of a stem perigynium. However, vestigial stem perigynia have independently developed in a wide variety of unrelated taxa. In this condition, there is a swollen region of axial proliferated tissue at the perianth base on which several or all of the bracts and bracteoles are inserted. Vestigial shoot perigynia are feebly developed in C. vermicularis and C. ctenophylla, more so in C. navistipula, and appear to reach an optimal extent of development for the genus in C. puccioana (fig. 50-1). With greater axial proliferation, along with formation of a shoot calyptra, a coelocaule precursor has developed in Subg. Lacerifolia (fig. 12-1). Here, the sporophyte "penetrates" to about the level of bracts of the first series or the leaves immediately below. The coelocaule precursor developed in Clasmatocolea is quite comparable to that in the genus Temnoma (see Schuster, 1967) and that referred to by Schuster (1966) as the '''Temnoma subtype" of axial protection of the embryo. More specifi- cally, the coelocaule precursor in Clasmatocolea is similar to less extreme forms of the Temnoma subtype; compare for example Figure 53-4 in Schuster (1966) for T. palmatum, and Figure 11-1 for C. obvoluta. As in Temnoma, there is likely minimal actual penetration of the sporophyte into axial tissue, but rather a proliferation of axial tissue upward in a cylinder around the developing sporophyte. The bracts and bracteole, as in Temnoma, are widely separated. Setae. — The seta exhibits no sharp internal differentiation when ob- served in transverse section. The epidermal layer is smaller than, equal to, or larger than the scattered, rather numerous internal cells, with little or no correlation of relative size and taxonomy. The number of cells in the epidermal row varies from 14 in C. cucuillstipula to as many as 70 in C. humilis. Other features also vary, such as the number of cells in seta diameter, as well as trigone-like thickenings of epidermal and inner cells. See the key to sporophyte material (p. 20). Capsules. — These structures are variable in shape, but in no case are acute or beaked capsule tips developed (such as those in Acrobolbaceae and Schistochilaceae). Capsule walls are 3-6-layered, often with some variation of layer number on the same valve. The outer layer is always thicker, equivalent to 1.8-3.9 of the interior strata. An extreme is reached 18 FIELDIANA: BOTANY in C. minutiretis, where the outer layer of cells exceeds the thickness of all the interior strata combined (fig. 41-7). The outer layer not only has nodule-like to spinelike thickenings on radial walls, but varying amounts of semiannular bands that extend from radial onto tangential walls (fig. 34-13). Only in C. vermicularis and C. humilis are semiannular bands lacking on the outer layer. The intermediate capsule wall layers are fairly uniform in wall features and possess tangentially dilated thickenings on radial walls. For the most part, the intermediate and inner layers are subequal in thickness. In a few species (C. navistipula and C. puccioand), the outermost intermediate layer is thicker than the innermost layer of the capsule. The inner layer of cells has semiannular bands extending from radial onto tangential walls. In several species, these bands are often incomplete. The radial walls often have nodulose thickenings interspersed with the bands. Except for the presence of semiannular bands in the outer layer, the type of capsule wall in Clasmalocolea fits the Jungermannia type outlined in Schuster (1966). Grolle used capsule wall anatomy as a generic criterion. Grolle (1956) thought C. heterostipa had a bistratose capsule wall that, along with the wide mouth and entire lobes of the perianth, was regarded as generically diagnostic. Grolle (1960), however, eliminated this feature and substituted for it the presence of "knotigen bis halbringformig" thickenings in the inner capsule wall layer as a generic criterion. The type of wall thickening found in Clasmatocolea needs evaluation and comparison with a wide array of other austral Lophocolea; this topic is currently under inves- tigation. Spores. — Within Clasmatocolea, there are two, narrowly circumscribed, yet very different spore types. Type 1. — Spores densely granulate, grading to short, wide, irregular vermiform ridges, which are covered (often densely so) with nanogranules (pis. 10, 25). 3 The great bulk of Clasmatocolea taxa have this spore type. Spores in this category are 10-18 ^t, reaching an upper limit in C. vermicu- laris, where they are 15-20 /z. This type of spore is surely primitive within the genus. In general, as to be expected, species with larger spore size have a greater spore-elater diameter ratio.4 For example, C. vermicularis has spores 15-20 n and a spore-elater ratio averaging 1.9:1, whereas C. obvoluta, with spores 11-12 /i, has a ratio averaging 1.1 :1. However, C. 3 1 have used the term "nanogranule" in the sense used by Sorsa & Koponen (1973) in their studies of Mniaceae spores. 4 In Clasmatocolea, as in many groups of Hepaticae, the direct correlation between spore size and spore-elater ratios is due to the relatively small elater width variation. ENGEL: CLASMATOCOLEA 19 rigens is an interesting exception to the rule, with spores 14-18 n and a spore-elater ratio averaging 2.4:1. Clasmatocolea obvoluta and C. puc- ciona, both with spores 11-12 n, form an interesting contrast: the former has a spore-elater ratio of 1.1 :1 and the latter, 1.5 :1. Type 2. — Spores with elongate, narrowly conical or subrectangular projections. Within this derivative spore type there is a trend in specializa- tion of the projections, culminating in strongly dilated, clawlike tips (pis. 28, 29). This type of spore is represented in but three species — C. cucul- listipula, C. ctenophylla, and C. gayana — belonging to two distantly related subgenera. Within these taxa, it appears that the presence of this spore type has ecological rather than phylogenetic relevance. The only instances of elongate projections of the exine are in corticolous members of the genus, and it is certainly possible that this spore type is useful in facilitating establishment of these plants. Two taxa in this group have the largest spores in the genus — C. gayana, with spores 21-24 /x, and C. cucullistipula, with spores 24-27(-30) /z, with the latter also having the largest spore-elater diameter ratio in the genus, 2.8:1. The greater surface area of the very large spore of C. cucullistipula may be of significance in anchorage and establishment of the spore in the unique and interesting niche of the spe- cies— loosely adhering to or semipendent from very small branches or twigs. The third species, C. ctenophylla, has much smaller spores, being 12-15 M. Scanning electron micrographs of spores of Clasmatocolea may be found on pages 204 through 224. Elaters. — I have not found elaters to be of phylogenetic or taxonomic value in Clasmatocolea. These structures in Clasmatocolea are bispiral and taper gradually toward the ends. Some species are spiraled to the tips, whereas others often have apice s with a thick, nonspiral wall, which does not seem to be of taxonomic significance. Elaters are 6-12 /* in diameter, with elater walls 2-5 /x wide. IV. COMPARATIVE MORPHOLOGICAL DIVERSITY AND EVOLUTION OF SPOROPHYTE AND GAMETOPHYTE From the beginning of this study I have been interested in determining if there are trends, relationships, or correlations of sporophyte characters that reflect evolutionary tendencies in the gametophyte. It is apparent that the few trends and character correlations present in the sporophyte generation reflect the stature of the gametophyte or ecology rather than any evolutionary trends in the gametophyte. For example, diminutive species such as C. ctenophylla, C. cucullistipula, and C. navistipula tend to 20 FIELDIANA: BOTANY have a reduced seta, both in number of cells in seta diameter and in epidermal row. It is apparent, after examining characters available from the sporophyte and the gametophyte, that morphological diversity and amount of evolution are vastly different in the two generations of Clasmat- ocolea. To underscore this, one need only to compare the preceding discussions of taxonomic characters found in the gametophyte and sporophyte generations. These findings are certainly not unexpected, for the persistent gametophytes and the ephemeral sporophytes have entirely different selective pressures acting upon them. See Schuster (1966) for a lengthy discussion of these factors. In an attempt to assess the overall taxonomic value of the sporophyte and the diversity of its characters, I have constructed a key, based solely on sporophyte material, but with spore and elater data added. To evaluate the variability of characters as well as to minimize risks that accompany inadequate sampling, a number of sporophytes of each species were studied whenever possible. For example, ten sporophytes of C. trachyopa were studied in detail. ARTIFICIAL KEY TO TAXA BASED UPON SPOROPHYTE MATERIAL 1 . Spores averaging more than 1 .5 X the elater diameter 2 2. Spore wall with elongate, narrowly conical, or subrectangular projections 3 3. Spores 12-15 y.; capsule wall 3-4 layers thick C. ctenophylla 3. Spores 21-27 /u; capsule wall 4-5 layers thick 4 4. Elaters 8-10 /x; outer capsule wall layer with semiannular bands very common C. cucullistipula 4. Elaters 10-12 ^; outer capsule wall layer with semiannular bands few C. gay ana 2. Spore wall granulate grading to short vermiform ridges 5 5. Spores averaging 2.4 X elater diameter C. rigens 5. Spores averaging 1 .9 X elater diameter 6 6. Capsule wall 3-4 layers thick ; outer capsule wall layer without semiannular bands, the wall equal to 2 of interior strata; capsule oblong to globose C. vermicularis 6. Capsule wall 4-5 layers thick; outer capsule wall layer with semiannular bands, the wall equal to 2.5-3 of interior strata; capsule broadly ovate to elliptic C. fulvella 1 . Spores averaging 1 .5 X or less the elater diameter 7 7. Spores averaging 1 . 1 -1 .2 X elater diameter 8 8. Outer capsule wall layer exceeding all of interior strata combined C. minutiretis 8. Outer capsule wall equal to 2-3 of interior strata 9 9. Seta with 30-70 rows of outer cells; outer capsule wall layer without semi- annular bands C. humilis 9. Seta (where known) with 19-32 rows of outer cells; outer capsule wall layer (of at least some valves) with semiannular bands 10 10. Capsule wall 36 /* thick, of 4 layers C. strongylophylla 10. Capsule wall 30-72 /x thick, of 4-6 layers C. obvoluta ENGEL: CLASMATOCOLEA 21 7. Spores averaging 1 .4-1.5 X elater diameter 11 11. Outer capsule wall layer equal to 1.8-2.5 of interior strata C. trachyopa 11. Outer capsule wall layer equal to 3-3.9 of interior strata 12 12. Seta with 15-16 rows of outer cells; capsule wall 36-42 n thick C. puccioana 12. Seta with 28-33 rows of outer cells; capsule wall 42-48 ^ thick C. navistipula V. ORIGIN OF CLASMATOCOLEA AND SOME EVOLUTIONARY TRENDS I believe that the genus Clasmatocolea arose from a simple member of the genus Lophocolea, characterized by such features as (a) convex, shallowly bifid, longer than wide leaves; (b) deeply bifid, rather large underleaves; (c) a plastic branching pattern without restriction of terminal or intercalary branches; (d) branches both like one another and like the main axis; (e) a simple, sharply trigonous perianth, with the three lobes bifid; (f) bracts and bracteoles bifid, much like the leaves; and (g) absence of any secondary pigments. Further, I postulate a Southern Hemisphere origin of the genus from an ancestor similar to a dioecious Lophocolea lenta (a monoecious taxon of southern Chile and the Falkland Islands), a species possessing many of the features listed above. From this pool of Lophocolea lenta-\ike plants, I believe there was an evolutionary line in which there had been an alteration of leaf concavity, seemingly a simple change, leading from a convex leaf to a concave one. Such plants might resemble Clasmatocolea rigens, also of the southern South American sector and the most primitive member of the genus. From this hypothetical prototype of Clasmatocolea, I believe early evolution occurred along two basic lines, both quite divergent from Lophocolea — one line toward Subg. Protoclasmatocolea and Subg. Laceri- folia, both with divided leaves, and the other toward the remaining groups, all of which have undivided leaves (see fig. 1). The Subg. Protoclasmatocolea-Lacerifolia line is an intriguing one, progressing from the basic, simple, shallowly bifid leaf of C. rigens, to species with an increasing elaboration and specialization of the leaf. This trend can be observed proceeding from C. rigens, to the still simple C. obvoluta var. cookiana, to the more complex C. obvoluta var. obvoluta, to the complicated, intricate leaves of C. trachyopa, which terminates the line. From the larger pool with undivided leaves, there is an offshoot com- prising Subg. Clasmatocolea (see fig. 1). This line, in turn, has several offshoots, each leading to a specialization of one or more structures, such as elaboration of branching and underleaves (Sect. Fulvellae) or elabora- • Subg. Plicaticalyx Subg. Squamicalyx Sect. Pachyclasmatocolea Sect. Fulvellae Sect. Strongylophyllae C. trachyopa C. obvoluta var. obvoluta C. obvoluta var. cookiana AJ ,cr Sect. Puccioanae • Subg. Metaclasmatocolea simple, Lophocolea-lenta Lophocoleoid ancestor FIG. 1. Diagrammatic representation of suggested major evolutionary lines in Clas- matocolea. 22 ENGEL: CLASMATOCOLEA 23 tion of the cuticle (Sect. Strongylophyllae). Another offshoot takes in the largest section of the genus, Sect. Clasmatocolea. There are some interest- ing trends within this section; some are isolated evolutionary experiments that are present in but a single species, whereas other trends are sequential, encompassing all taxa within the section. Some of these trends are listed below. Primitive character Underleaves large, conspicuous Underleaves divided Branching pattern generalized (see Table 1) Stem anatomy generalized Trigones absent or minute Rhizoid pads absent Leaf insertion simple (cf. C. humilis) Perianths trigonous Perianth keels present, wings well developed Ventral lobe of perianth large, like the lateral Spore exine with vermiform ridges Advanced character Underleaves small, inconspicuous Underleaves retuse or undivided Branching pattern specialized, with 1 or more types absent Stems with differentiation of a distinct cortex Trigones massive, the cell lumen bounded mostly by trigones Rhizoid pads present Leaf insertion specialized (cf. C. moniliformis) Perianths terete, at least toward the base Perianth keels and wings absent Ventral lobe of perianth reduced, smaller than the lateral Spore exine with elongate projections Another offshoot from the line possessing undivided leaves is Subg. Metaclasmatocolea, with specialized branching modalities and highly re- duced underleaves. There are fascinating trends in this line, with one sec- tion (Puccioanae) developing a complex system of lateral-intercalary branches and having underleaf reduction only on the branches. The other section (Metaclasmatocoled) has developed a flabellate branching pattern and underleaf reduction on all axes, including the main one. There are two independent and derivative lines, each quite apart from the main stream of evolution within the genus, but still within the un- divided leaf group (see fig. 1). One comprises Subg. Squamicalyx, with such derivative features as underleaves connate on both sides (including, on one side, connation with the leaf lamina in C. notophylla), massive trigones that nearly surround the cell lumen, and more or less terete perianths with large, curious scales. The second line, the monotypic Subg. Plicaticalyx, is the most derivative, isolated taxon in the genus. Nowhere else does one find such features as contracted, plicate perianths, andreocia wider than the main axis, 2-3 laciniate-lobate androecial bracts, or distinctly auriculate underleaves. 24 FIELDIANA: BOTANY VI. GENERIC RELATIONSHIPS OF CLASMATOCOLEA Clasmatocolea is a discrete, well-defined genus of Lophocoleaceae closely related to Lophocolea, Leptoscyphus, Evansianthus, Pachyghssa, Hepatostolonophora, and Xenocephalozia. These genera also independently relate to other genera in the family (see fig. 2). Several different evolutionary lines within Clasmatocolea show relation- ships to Lophocolea, a rather large genus with considerable species diversity in the Southern Hemisphere ; some of these are outlined below : 1 . Clasmatocolea rigens (Subg. Protodasmatocoled) approaches several bifid-leaved Lophocolea taxa, such as L. lenta and to a lesser extent L. leptantha and L. textilis, all from southern South America (see related discussion on p. 21). 2. Clasmatocolea vermicularis (Sect. Clasmatocolea) has some pheno- types quite reminiscent of L. sabuletomm of southern South America and the Falkland Islands (see p. 104). 3. Clasmatocolea humilis (Sect. Clasmatocolea) approaches the L. austrigena-gunniana complex and has a similar amphipacific distribution . There are certain phenotypes of all three of these species that cause considerable taxonomic difficulty between genera (see discussion on p. 82). 4. Clasmatocolea humilis also shows a relationship, but to a lesser extent, to L. otiphylla of southern South America (see p. 82). Although there are several points at which Clasmatocolea and Lopho- colea are taxonomically close to one another, evolution in the two taxa has diverged sharply. For example, in the C. rigens-L. lenta species pair, a divergent sequence can be observed beginning with C. rigens, progressing through the varieties of C. obvoluta, and culminating in C. trachyopa. The simple L. lenta, on the other hand, relates to Lophocoleoid species with distinctly convex leaves. A different situation exists for C. humilis and the L. austrigena-gunniana complex. Lophocolea austrigena is part of a trend sharply diverging from evolutionary developments seen in Clasmato- colea, proceeding from L. boveana to L. austrigena (and its ally L. gunniand) and culminating in the nearly isophyllous L. gottscheoides, which does not, by any means, suggest the genus Clasmatocolea. Divergent evolution of Clasmatocolea from the Lophocoleoid line seems to have involved mainly leaf characters. Leaves of Clasmatocolea are adaxially concave and are commonly conchiform concave, whereas those of Lophocolea are plane or convex. Concomitant with this, the trigones in Clasmatocolea are large to massive, and the cell size small; Lophocolea has comparatively much smaller trigones and larger cells. Apotho. Chilo. Cono. Evans. Hepato. Hetero. Leptophyll. Pachyg. Piga. Telra. Xeno. Genera Apothomanthus Chiloscyphus Conoscyphus Evansianthus Hepa tos tolonophora Heteroscyphus Leptophyllopsis Pachyglossa Pigafettoa Tetracymbaliella Xenocephalozia •primitive Lophocoleoid ancestry] Subgeneric units units in Lophocolea ten. com. = L. lenta complex Ot. = L. otiphylla sabu com. = L. sabuletorum complex aust com. = L. austrigena complex subgenera in Clasmatocolea Pro. = Protoclasmatocolea Lac. = Lacen folia Cla. = Clasmatocolea Mela = Metaclasmatocolea Sq. = Squamica/yx Pli. = Plicaticalyx FIG. 2. Pictorial representation of generic relationships in Lophocoleaceae. 25 26 FIELDIANA: BOTANY The number and significance of shared characters of Clasmatocolea and Leploscyphus indicate an affinity between these genera; these features are outlined below: 1) In several independent lines of Clasmatocolea, there is a tendency for reduction of the ventral lobe of the perianth. In these groups, as well as those that have a well-developed ventral lobe, a bilateral compression of the perianth is not unusual. Such bilaterally compressed perianths are not far removed from the Leptoscyphoid perianth. Further, one should recall that there are several species of Leptoscyphus in which a third ventral lobe of the perianth is present, even though perianths in these taxa are strongly bilaterally compressed and remain two-lipped. 2) Leptoscyphus has several species that have strongly conchiform concave leaves much like those of Clasmatocolea. 3) Large, bulging, knotlike trigones are commonplace in both genera. 4) The texture of the plants is quite similar in the two genera. The two genera, however, are quite independent of one another. Clasmatocolea lacks the ability to produce reddish pigments that is so widespread in Leptoscyphus. Further, trends toward perianth bilateral compression in Clasmatocolea notwithstanding, the shape of the perianth still may be nicely used to separate the genera — basically trigonous to terete with the ventral lobe well developed in Clasmatocolea and strongly bilaterally compressed, basically two-lipped in Leptoscyphus. Clasmatocolea is more remotely allied to the genus Evansianthus, which has a similar basic plant physiognomy, including concave leaves. Evansian- thus, however, is at once distinct from Clasmatocolea through possession of the following characters, which dictate a more distant relationship: (a) well-developed bilaterally compressed perianths; (b) Andrewsianthus- type branching; (c) presence of flagella; and (d) subterranean habit of growth. See Schuster & Engel (1973) for further notes on Evansianthus. Clasmatocolea is allied to the genus Pachyglossa, although evolution in the two genera seems to have gone in different directions. I regard C. marginata as pivotal in connecting these two genera. This species is erect, rigid, brown pigmented, has polystratose leaves and underleaves (at least in the basal region), and has well-developed intercalary branching — all of which are features of Pachyglossa. I found the apical portions of robust axes of C. marginata particularly striking, for they have much the facies of an overblown Pachyglossa (see fig. 18-1). The trio of genera — Clasmato- colea, Lophocolea, and Pachyglossa — are an intermeshed ensemble of taxa, each relating to the other two genera (see fig. 2). The genus Hepatostolonophora should be mentioned in this discussion, ENGEL: CLASMATOCOLEA 27 for in several respects it is involved in the Clasmatocolea-Pachyglossa connection. As outlined in Engel (1979b), Hepatostolonophora differs from Clasmatocolea as follows: (a) the presence of stolons; (b) the reduced, inconspicuous gynoecial bracteoles that are hardly modified from under- leaves; (c) the vertically oriented, subtransverse leaves (although some facies of H. paucislipula have succubously oriented leaves) ; and (d) the di- morphic leaf cells, with the majority possessing oil-bodies, although 10-12 percent of the cells lack them. In one way or another, one can state that all the genera discussed here are involved in a web of relationships. One ally of Clasmatocolea, however, forms an exception, and this is the genus Xenocephalozia, which seems to be connected to this complex only through kinship with Clasmatocolea. The deep concavity of the leaf, with the ventral portion incurved and flattened parallel to the ventral stem surface, as well as the strongly trigonous perianths with bifid lobes, remind one of Clasmatocolea, es- pecially of C. trachyopa. However, the narrow, transverse to weak in- cubous leaf insertion of Xenocephalozia is in my mind a fundamental difference negating any real, close relationship (see notes on p. 194). Clasmatocolea may be separated from all other members of the family by the key below. KEY TO LOPHOCOLEACEAE5 1 . Gynoecial bracts and bracteole not fused to form a tube, the perianth clearly visible, not completely enveloped by bracts 2 2. Perianths ± strongly laterally compressed, the mouth truncate, wide, basically 2-lipped. Plants often =*= brownish, often intensely so 3 3. Andrewsianthus-type branching present; leaves medially to basally polystratose ; flagelliform branches present; stem tissue with endophytic hyphae Evansianthus 3. Andrewsianthus-type branching absent; leaves unistratose throughout; flagel- liform branches absent; stem tissue without endophytic hyphae. .Leptoscyphus 2. Perianths trigonous to trigonous inflated 4 4. Perianths all or for most part restricted to strongly abbreviated lateral- and/or ventral-intercalary branches 5 5. Ventral margin of leaves and lateral margins of underleaves with a pouch Tetracymbaliella 5. Ventral margin of leaves and lateral margins of underleaves without a pouch 6 6. Androecia poorly differentiated, interterminal or calary °n leading axes; androecial bracts much like leaves except for a small, dorsal flap or lobule and an associated small, local pocket or concavity. .Chiloscyphus 6. Androecia spicate, usually on short, abbreviated lateral- or ventral-inter- calary branches; androecial bracts clearly differentiated from leaves, 5 A few portions of key adapted from Schuster (1963). 28 FIELDIANA: BOTANY lobule large and conspicuous, the entire bract base ventricose Heteroscyphus 4. Perianth normally on more or less long branches (branches rarely abbreviated and never consistently so) 7 7. Plants stoloniferous.6 Leaves (exc. in H. paucistipuld) transverse to sub- succubously oriented 8 8. Plants nearly isophyllous, the underleaves conspicuous and rigidly patent; leaves and underleaves polystratose at least at the base; female bracteoles =>= similar in size to bracts Pachyglossa 8. Plants clearly anisophyllous, the underleaves highly reduced, incon- spicuous, appressed to moderately spreading; leaves unistratose through- out; female bracteoles reduced, inconspicuous, hardly modified from underleaves Hepatostolonophora 1. Plants usually lacking stolons 9 9. Leaves moderately to deeply adaxially concave 10 10. Leaves strongly succubous, the insertion a long, strongly oblique line 11 11. Underleaves free or connate on 1 or both sides by a few cells; leaves (exc. C. marginatd) unistratose throughout; perianths unistratose, usually wide mouthed; plants occasionally with brown pigments, never with red Clasmatocolea 11. Underleaves distinctly and conspicuously connate on 1 side by several cells; leaves bistratose toward the base; perianth 2(-3)- stratose in median portion, abruptly contracted at the mouth; plants with brown or red-brown secondary pigments Apothomanthus 10. Leaves transverse, subsuccubous or subincubous, the insertion narrow 12 12. Leaf margins (and abaxial leaf faces) crenulate to mamillate with domelike cell wall protuberances; leaves Cephaloziella-ltke, at most slightly concave, without an incurved ventral margin, normally lacking any trace of teeth ; leaf insertion transverse to weakly succubous Pigafettoa 12. Leaf margins (and abaxial leaf faces) not crenulate to mamillate with domelike cell wall protuberances; leaves =*= Nowellia-likc, strongly concave, the ventral margin incurved and accentuating and defining a deep pocket-like concavity in ventral portion of leaf, the leaves with accessory teeth ; leaf insertion transverse to weakly incubous Xenocephalozia 9. Leaves convex, rarely plane, with apices decurved or deflexed 13 13. Leaves with lobes and marginal teeth caducous, often giving leaf apices a ragged appearance; leaf apices often with accessory teeth and laciniae Leptophyllopsis 13. Leaves entire or if lobed, then with lobes persistent; leaves with marginal teeth, if present, persistent; leaf apices never with a ragged appearance and not with accessory laciniae Lophocolea 6 Care should be taken in search of stolons before concluding that they are absent, because H. paucistipula does not have stolons uniformly present on all axes. ENGEL: CLASMATOCOLEA 29 1. Gynoecial bracts and bracteole fused to form a tube that encloses and hides the perianth Conoscyphus VII. PHYTOGEOGRAPHY The genus Clasmatocolea is among a large group of hepatic genera confined, or essentially so, to the south temperate and subantarctic regions of the world. The distribution patterns of Clasmatocolea species may be arranged in the categories below. Categories are provided even though species of Clasmatocolea do not fall within them. TEMPERATE Species occurring within the south temperate and sometimes subantarctic regions of the world. Species included here are regarded as temperate rather than subantarctic, because northward extension is not confined to higher altitudes (see Nontemperate, Subantarctic category below). Pan-Temperate Species occurring in temperate regions of South America, New Zealand, Tasmania-Australia, and South Africa. C. vermicularis Amphipacific Temperate Distribution occurring in temperate parts of the South Pacific in the Southern Hemisphere, i.e., temperate South America and the New Zealand sector. C. humilis C. notophylla Amphiatlantic Temperate Distribution mainly in temperate parts of the south Atlantic, i.e., temperate South America and South Africa. None American Temperate Species occurring within the south temperate regions of the American sector. Fuegian. — Species restricted to Tierra del Fuego, i.e., south of the Strait of Magellan. None Magellanian. — Species occurring in an area bounded by Cape Horn and 48° S. The northern boundary was affixed by Skottsberg (1916) to delimit the Magellanian and Valdivian regions and has been widely followed by various authors. This category includes the previous (Fuegian) one. C. moniliformis 30 FIELDIANA: BOTANY Magellanian-Valdivian. — Species essentially widespread in the South American temperate zone. They are represented in the Magellanian region and also the Valdivian region, which lies between 48° S. and 36° S. Skottsberg (1916, p. 13) places the northern boundary of the Valdivian region at 40° S., whereas Kuschel (1960) states that the zone occurs from 36° S. in the Andes, 37° S. in the coastal range, and 38° S. in the central valley. The taxa listed below are arranged according to their northernmost latitudinal range. A dagger indicates occurrence in the Falkland Islands, and an asterisk represents occurrence on the Tristan da Cunha Group. *C. minutiretis (41 ° 00' S.) C. cucullistipula (39° 16' S.) fC. obvoluta (40° 45' S.) C. gayana (37° 46' S.) C. puccioana (40° 07' S.) fC. fulvella (36° 50' S.) C. trachyopa (39° 56' S.) C. navistipula (36° 50' S.) C. ctenophylla (39° 52' S.) fC. rigens (33° 21' S.) South African Temperate Species occurring in the Cape Region. C. fasciculata Australasian Temperate Species occurring in New Zealand and/or Tasmania. A dagger indicates occurrence on New Zealand, an asterisk indicates occurrence on the New Zealand shelf islands, and a double dagger indicates occurrence on Tasmania. Species in temperate portions of Australia would ordinarily fall in this category, but the only Clasmatocolea taxon occurring in Australia (C. humilis) is amphipacific temperate, q.v. fC. crassiretis t*|C. strongylophylla }C. marginata %C. verrucosa NONTEMPERATE Subantarctic Species occurring on one or more subantarctic islands (as defined by Greene, 1964) of one or more sectors (e.g., American, African, or New Zealand) with northward extensions only at higher altitudes. See addi- tional notes in Engel (1978, pp. 35-36). None Antarctic Species occurring in the Antarctic Zone (as defined by Greene, 1964) with northward extensions into the subantarctic or temperate zones only at higher altitudes. None ENGEL: CLASMATOCOLEA 31 Malaysian Montane Species occurring in the mountains of Malaysia. C. tjiwideiensis Clasmatocolea vermicularis represents the other example of northward penetration beyond temperate latitudes, but this species is widespread in austral areas, being pan-temperate plus well distributed on several sub- antarctic islands. Its distribution northward is strictly at higher altitudes in the Andes and higher eastern African mountains. The exception to the strict south temperate-subantarctic range of the genus is C. tjiwideiensis of high-altitude Java, the only extra south tem- perate species. It appears that this species has been isolated for some time, for it has such features as magenta rhizoids (all other Clasmatocolea species having colorless rhizoids) and a unique pattern of leaf armature. Often the areas where a species or group does not occur are just as significant or interesting as where it does occur. Clasmatocolea is a cool- temperate, drought-intolerant genus, and much of its local, austral dis- tribution is explainable on this basis. The paucity of species in South Africa (two species, only one endemic) can probably be attributed to the area being both too warm and too arid for the genus. The same factors, but to a lesser degree, might explain the paucity of the genus in Australia, where none of the taxa confined to the Australasian temperate region (see above) occur. Only the amphipacific C. humilis occurs in Australia. There are no records of the genus in New Guinea at high altitudes, where one would expect the genus to occur. In fact, except for the Malaysian locality, Clasmatocolea does not occur north of Australia. The distribution patterns of Clasmatocolea illustrate that the genus is intimately associated with the concept of continental drift and the former presence of a large southern land mass. This supercontinent, Gondwana- land, originated from the breakup of Pangaea around the middle of the Triassic (200 m. y. BP) (Dietz & Holden, 1970a, 1970b) and consisted of South America, Africa, India, Antarctica, and Australia (see fig. 3). With the juxtaposition of these continents making up Gondwana, Schuster (1972a) and Raven & Axelrod (1974) emphasize the availability of a migratory route that involved the fringes of Antarctica. Further, these authors point out that this route was available for a very long period of time. The Gondwanaland configuration shown in Figure 3 was presented in Dietz & Holden (1970a) and is quite similar to those in Schopf (1970) and Smith & Hallam (1970), the former of which is shown in Figure 4. The Schopf as well as Smith & Hallam schemes are helpful for our pur- poses, because the configurations show the presumed positions of Tas- -I E § S° C- a. •3 g 'C C hH E « w o « J- P- •JC *i ON C! o ~" o «" c -0 -.§ 4) "O O 2 § K C8 '^ N t/5 fJ fl> 1 15 •^ j *- •a „ S 2 co" '•a II! *J '^3 4> N = W7 _- § g O £ •£ 32 ENGEL: CLASMATOCOLEA 33 \ SOUThL AMERICA FIG. 4. Reconstruction of Permian positions of Gondwanaland continents. Tasm = Tasmania; Aust = Australia; NZ = New Zealand; NC = New Caledonia; NG = New Guinea. (After Schopf, 1970.) mania and New Zealand. The dating of Gondwanaland differs somewhat — the Schopf model is Permian and thus earlier than the Triassic model of Dietz & Holden or the Jurassic model of Smith & Hallam. The dating of the supercontinent is less important than the possible migratory pathways available from southern South America along the fringes of Antarctica to Australia, New Zealand, and Tasmania, or, of course, the reverse. This migratory pathway is of fundamental importance in understanding the phytogeography of the far south. Returning to the Dietz & Holden models, there were two important events taking place by the end of the Jurassic or beginning of the Cretaceous (180-135 m. y. BP) (see fig. 5) — one is that there was some northward migration of the Antarctic-Australian bloc, and the other is the narrow rift (shown by the dotted line) between Africa and South America. These events continued through the Cretaceous, but at different rates, so that by the end of the Cretaceous or start of the "2 o §E 2 «8 I c c o o o • • 60 2 '1 C J 34 ENGEL: CLASMATOCOLEA 35 Tertiary (fig. 6), Africa was isolated, but the Australian-Antarctic-South American unit did not begin to separate until some time in the Cretaceous. The separation between South America and Antarctica was narrow and presumably a phytogeographically minor one, meaning that by the early Tertiary (60-65 m. y. BP), a major austral migratory pathway still existed between southern South America and Tasmania-Australia via the Antarc- tic bloc. New Zealand, however, separated from Australia-Antarctica 80 m. y. BP (Raven & Axelrod, 1972, 1974). The extant range of the most primitive groups would indicate that the genus Clasmatocolea originated in the southern South American sector of Gondwanaland. Of the two most primitive subgenera — Subg. Proto- clasmatocolea and Subg. Lacerifolia — all species occur in the southern South American sector. The genus may have later migrated from the American sector to other southern land masses. Southern South America is also the center of distribution of the genus — of the 20 species in the genus, 15 occur in southern South America. An alternative explanation might be that the genus originated somewhere in Gondwanaland, and that the most primitive groups, or their ancestors, had a fairly wide distribution but survived only in southern South America. It is entirely plausible to think that the primitive groups survived in an area where most of the species are found today and that southern South America had continuing environmental conditions necessary for survival. Clasmatocolea occurs in both forested and alpine areas, although I would not regard any species in the genus as an obligate alpine taxon. The forest-occurring species might be thought to be remnants of an early, cool-temperate flora that characterized the forested lowland areas of the Antarctic continent. Taxa that occur at higher altitudes and that are not uncommonly represented in the alpine-subalpine zone, on the other hand, might then represent remnants of a flora that characterized the more upland reaches of the Antarctic continent. In this connection, it is interest- ing to speculate on the history of the pan-temperate and amphipacific taxa. All species with these patterns — the pan-temperate C. vermicularis and the amphipacific C. notophylla and C. humilis — are not unusual in the subalpine-alpine zone in one or more sectors of their range. Perhaps, at least within the genus Clasmatocolea, the upland species had the greatest opportunity or capacity for dispersal over wide expanses, whereas taxa more characteristic of forests did not have this opportunity or capacity. An alternative explanation might be that the taxa presently restricted to one sector were never widely distributed or that their ancestors had a wider range but have since disappeared. In any case, species confined to 3 «-> ti 0 ° o Jl; 2 ca ~ Urtl c/i 55 .dS-e •£ O 73 n <« c O <-• J3 — i CS CJ •° •£ 5 C o .C •S i §15 d ° S'Sg r ( f ^ CT\ O O 2 -" "5 5 c g 3 fl . ^* *^ iS O 73 "o § ^§J a '3 .5 * 11 i— ^i r*t •rt » -g IS 2 5 ^ 36 ENGEL: CLASMATOCOLEA 37 one sector are, for the most part, forest species, with only an occasional occurrence in alpine-subalpine areas. A comparison of Clasmatocolea with other hepatic genera that are endemic, or nearly so, to south temperate-subantarctic regions provides some useful information. Besides strictly south temperate-subantarctic genera, also included here are those genera containing some species that are south temperate-tropical montane in the American or Australasian regions. This northward penetration in either the Andes or high-altitude Australasia in all cases is limited and in nearly all instances occurs in only one species in the genus. It should be mentioned that there is a rather large number of small genera endemic to a single south temperate-subantarctic region. These are outlined in Schuster (1969, cf. his category A-l on p. 50) and are not discussed here. Nearly all of these genera occur in southern South America or New Zealand-Tasmania-Australia, with a minimal number in South Africa. The genera occurring in more than one south temperate area, which are listed below, are all amphipacific in distribution, whereas none are amphiatlantic or pan-temperate. There is also a group of genera that is Gondwanalandic in origin, but with more extensive penetration into tropical latitudes. These genera usually have tropical species; Schuster (1969) enumerates these groups. In comparing Clasmatocolea to other endemic or near-endemic genera, only those genera occurring in more than one sector of the south tem- perate region are included. I have arranged the genera according to broad aspects of their ecology. The number of species in the genus is in pa- rentheses. A. Genera strictly alpine-subalpine Acrolophozia (2) Austrolophozia (3) ? Cephalolobus (4) Herzogobryum (6) Pachyglossa (4) Phyllothallia (2) B. Genera occurring all or mostly in forests Archeophylla (3) Austrolejeunea (2) ? Hyalolepidozia (2) Leptophyllopsis (2) C. Genera occurring in both alpine-subalpine and forested areas; oc- 38 FIELDIANA: BOTANY curring at lower altitudes on both the subantarctic islands and mainland areas Allisoniella (5) Balantiopsis (12) Blepharidophyllum (6) Clasmatocolea (20) Cryptochila (6) Gackstroemia (7) Lepidolaena (8) Pseudocephalozia (6) Hepatostolonophora (3) Triandrophyllum (3) Clasmatocolea, with 20 species, is the only comparatively large genus ot hepatics to have a near-pure south temperate-subantarctic range. Com- pared to Clasmatocolea, all of the genera in the above list are small or relatively so. Further, Clasmatocolea is the only pan-temperate representa- tive. The only instances of penetration of the genus northward beyond temperate latitudes involve strictly high altitudes. VIII. CONSPECTUS OF TAXA Subg. Protoclasmatocolea Engel, subg. nov. 1. C. rigens (Hook. f. & Tayl.) Engel Subg. Lacerifolia Steph. ex Engel, subg. nov. 2. C. obvoluta (Hook. f. & Tayl.) Grolle a. var. obvoluta b. var. cookiana (Mass.) Engel 3. C. trachyopa (Hook. f. & Tayl.) Grolle Subg. Clasmatocolea Sect. Pachyclasmatocolea Engel, sect. nov. 4. C. marginata (Steph.) Grolle Sect. Clasmatocolea 5. C. humilis (Hook. f. & Tayl.) Grolle a. var. humilis b. var. suspecta (Mass.) Engel c. var. polymorpha Engel 6. C. fasciculata (Nees) Grolle 7. C. vermicularis (Lehm.) Grolle 8. C. gayana (Mont.) Grolle 9. C. ctenophylla (Schiffn.) Grolle 10. C. crassiretis (Herz.) Grolle ENGEL: CLASMATOCOLEA 39 1 1 . C. moniliformis Engel 12. C. minutiretis Engel & Grolle Sect. Fulvellae Engel, sect. nov. 13. C.fulvella (Hook. f. & Tayl.) Grolle Sect. Strongylophyllae Engel, sect. nov. 14. C. strongylophylla (Hook. f. & Tayl.) Grolle 15. C. tjiwideiensis (Sande-Lac.) Grolle Subg. Metaclasmatocolea Engel, subg. nov. Sect. Puccioanae Engel, sect. nov. 16. C. puccioana (De Not.) Grolle Sect. Metaclasmatocolea 17. C. navistipula (Steph.) Grolle a. var. navistipula b. var. parceramosa Engel Subg. Squamicalyx Engel, subg. nov. 18. C. notophylla (Hook. f. & Tayl.) Grolle 19. C. verrucosa Engel Subg. Plicaticalyx Engel, subg. nov. 20. C. cucullistipula (Steph.) Grolle IX. CLASMATOCOLEA Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:440. 1885. Plants prostrate to suberect, occasionally erect or subpendent, various shades of green and brown, including red-brown; main axis of smaller species less than 0.5 mm. wide, of larger species to 3 mm. wide. Branching variable, ventral- or lateral-intercalary branches usually present (but not always together in one species); Frullania-type present in some taxa, absent in others; Acromastigum-type developed in one species; stolons absent. Stems with cortex in 1-3 rows of cells or with cortex hardly differentiated. Rhizoids colorless or magenta, from stem near underleaf base, rarely from stem near ventral base of leaf, the tips simple or branched. Leaves usually unistratose throughout, rarely with leaf margins 2-4-stratose and forming an elevated border, rarely with median-basal area 2-4-stratose, the leaves succubously oriented, insertion oblique, not or slightly to distinctly recurved at ventral end; leaves usually conchiform concave, occasionally moderately adaxially concave; apex undivided or 2(-3)-lobed, rarely with accessory lobes; margins entire or variously armed with teeth, laciniae or, exceptionally, accessory lobes. Leaf cells thin to moderately thick walled, trigones large to bulging to coarse and nodular, less commonly absent, small, or medium; cuticle smooth or roughened or with coarse, hemispherical to oblong, hyaline papillae or verrucae. Oil-bodies throughout leaf, rarely absent from marginal 1-2 rows of cells, of the grape-cluster type, with the surface appearing granular or papillose; oil-bodies consistently 2 or 2-3 per cell in leaf middle. 40 FIELDIANA: BOTANY Underleaves distinct, persistent, 0.35-6.7 X stem width, free or connate on 1 or both sides, insertion nearly always weakly to strongly arced, rarely straight; underleaves plane or convex (ventral view), sometimes cucullate, with apices undivided or bidentate to retuse to bifid to 0.7, rarely divided nearly to the base, the apex with teeth or segments usually terminating in a slime papilla; margins of the lamina entire or dentate to lacini- ate to 1-lobate or merely with slime papillae, the armature usually terminating in a slime papilla. Gemmae absent; asexual reproduction rare, by regeneration from cells of leaf apex. Plants dioecious (at least those seen); androecia terminal or intercalary on leading axes or on short, abbreviated branches, clearly differentiated from leaves; bracts (at least in basal half) strongly saccate, the distal portion usually strongly concave, the bracts sometimes with saccate portion fused toward base with opposite bract; bracts with the dorsal base dilated and forming an infolded flap or lobule; lobule large in proportion to bract size, the margin entire or dentate to laciniate or with one involute lobe, the lobule margin usually with slime papillae; bracts rarely conduplicately bilobed; antheridia solitary, rarely 2 per bract, stalk uniseriate throughout or with localized bistratose areas or 4-seriate. Gynoecia on main axis or elongated branches or on short, abbreviated ventral- or lateral-intercalary branches; vestigial stem perigynium or coelocaule precursor some- times present; bracts in 2-4 series, becoming progressively larger toward the perianth, those of innermost series with apices undivided to retuse or bifid, rarely trifid; margins entire or dentate to ciliate to laciniate. Bracteoles of innermost series conspicuous, considerably modified from underleaves, free, rarely connate on both sides; apices undivided or retuse to bidentate to bifid to 0.2(-0.5); lamina margins entire or dentate- laciniate-lobate. Perianth inflated, weakly to strongly trigonous to terete, particularly toward the base, the mouth wide, exceptionally contracted, 3-lobed; lobes broadly rounded, the ventral sometimes smaller and more narrowly rounded, the lobes entire or dentate to ciliate to laciniate, sometimes bifid; keels often winged. Seta in transverse section with epidermal cells not greatly differentiated from the scattered, numerous inner cells. Capsule wall of 3-6 layers, the outer row of cells larger, equal to thickness of 1.8-3.9 of interior strata, the outer layer with thickenings on radial walls, the thickenings nodule-like to spinelike, semiannular bands present or absent; intermediate layers with thickenings on radial walls; inner layer of cells with thickenings on radial and tangential walls, semiannular bands extending from radial walls, the bands sometimes incomplete, the radial walls often with nodulose thickenings. Spores light brown or red-brown, exine with granulate grading to short vermiform ridges that have nanogranules, or the exine with narrowly conical projections having dilated tips; spores averaging 1.1-2.8 X elater diameter. Elaters bispiral. Lectotype (fide Grolle, 1956): Clasmatocolea heterostipa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:441. 1885. X. KEY TO TAXA OF CLASMATOCOLEA 1. Leaves basically 2-3-lobed; perianth lobes bifid 2 2. Underleaves bifid from ca. 0.5 to usually near the base, often semiobliquely inserted; leaves usually oriented toward axis apices; opposing male bracts free; ENGEL: CLASMATOCOLEA 41 coelocaule precursor absent; spores averaging 2.4 X elater diameter. Subg. Protoclasmatocolea C. rigens (p. 44) 2. Underleaves 0.12- to slightly less than 0.5-bifid; leaves strongly erect, not oriented toward axis apices; opposing male bracts fused toward base; coelocaule pre- cursor present; spores averaging 1.1-1.4 X elater diameter. Subg. Lacerifolia. . .3 3. Dorsal lobe acuminate; leaves with dorsal and median (if trifid) or ventral (if bifid) lobes with several opposing small to large and spinescent teeth or short cilia and often a few laciniae; underleaves free or connate on 1 side, 0.25- to slightly less than 0.5-bifid C. trachyopa (p. 60) 3. Dorsal lobe medium- to wide-triangular; leaves with dorsal lobes entire or with 1-2 pairs of opposing teeth, the ventral lobes entire, at most with a single tooth at ventral base; underleaves nearly always connate on both sides, 0.12-0.23-bifid C. obvoluta (p. 50) Leaves unlobed; perianth lobes broadly rounded, undivided 4 4. Underleaves with one conspicuous lobe in basal portion of underleaf that extends beyond dorsal surface of stem, the underleaves cucullate, auriculate at the base; androecia wider than sterile portion of axis, the bract lobules frequently 1-3- laciniate to lobate; perianths stoutly ovoid, the apical portion 4-5-plicate; spores 24-27(-30) n, the exine with narrowly conical projections. Subg. Plicaticalyx .... C. cucullistipula (p. 183) 4. Underleaf armature, if present, not represented by a single basal lobe extending beyond dorsal surface of stem ; androecia narrower than sterile portion of axis, the bract lobules entire, dentate, or with 1 lacinium or lobe; perianths various but not stoutly ovoid (exc. C. verrucosd), the apical portion without plicae; spores to 24 » 5 5. Underleaves connate with leaves on both sides, attachment with either ventral margin or ventral-basal portion of leaf; leaf cell lumen, except for the abbrevi- ated thin-walled places, bounded mostly by the massive, protuberant often con- fluent trigones; perianth basal portion with large scales. Subg. Squamicalyx . .6 6. Leaf cells each with one prominant, domelike, hyaline cuticular verruca; leaves with dorsal and ventral margins irregularly dentate; underleaves reniform, 4.4-5.1 X the stem width; Frullania-type branching predominant C. verrucosa (p. 179) 6. Leaf cells with cuticle smooth or at most roughened ; leaves with margins completely entire; underleaves orbicular to oblate, 3.1-3.6 X the stem width; Frullania-type branching absent C. notophylla (p. 173) 5. Underleaves free or connate on 1 side, never connate with laminal portion of leaf [or if connate on both sides (C. tjiwideiensis), then with leaves dentate] ; leaf cell lumen bounded mostly by cell wall, the trigones consuming only fraction of boundary (exc. in C. crassiretis); perianth lacking scales 7 7. Underleaves of main axis and/or lateral-intercalary branches inconspicuous, reduced, scalelike, polymorphous, often of only a few cells. Leaves entire. Subg. Metaclasmatocolea 8 8. Underleaves of main axis well developed; lateral-intercalary branches copiously produced, of limited growth, usually abbreviated; main axis underleaves with segments of similar size and shape; male bracts not conduplicately bilobed; plants light brown-sienna; main axes 560-910 ^ wide. Sect. Puccioanae C. puccioana (p. 1 57) 8. Underleaves of main axis reduced; lateral-intercalary branches usually 42 FIELDIANA: BOTANY of unlimited growth ; main axis underleaves frequently with asymmetric segments; male bracts conduplicately bilobed; plants whitish-light green; main axes 455-595 /x wide. Sect. Metaclasmatocolea C. navistipula (p. 165) 7. Underleaves of main axis and branches conspicuous, not reduced or if inconspicuous and reduced (C. minutiretis), then leaves with several teeth. Subg. Clasmatocolea 9 9. Leaf and sometimes underleaf margins conspicuously swollen, the ele- vated border 2-4-stratose; leaf lamina 2-4-stratose in median-basal area; antheridial stalks 4-seriate. Sect. Pachyclasmatocolea C. marginata (p. 69) 9. Leaf and underleaf margins not swollen or elevated, unistratose; leaf lamina unistratose throughout; antheridial stalks 1-2-seriate 10 10. Leaf cuticle with coarse, hemispherical, hyaline papillae. Sect. Strongylophyllae 11 11. Underleaves plane to weakly convex (ventral view), 1-2 X stem width ; leaves entire ; rhizoids colorless C. strongylophylla (p. 147) 11. Underleaves deeply concave to ± cupulate (ventral view), 3.3-4 X stem width; leaves with ventral margin 1-3-dentate to- ward the base; rhizoids magenta C. tjiwideiensis (p. 154) 10. Leaf cuticle smooth or slightly roughened 12 12. Intercalary branches appearing quite different from main axis, the branches vermiform and submoniliform, narrower in width than main axis, commonly copiously produced and short, ab- breviated, with underleaves more closely imbricate than in main axis. Leaves of main axis sinuate and incurved; intercalary branch underleaves large, orbicular, distinctly cupulate, appear- ing inflated. Sect. Fulvellae C. fulvella (p. 139) 12. Intercalary branches not vermiform or submoniliform (exc. C. moniliformis), usually of same width as the main axis, often very long, with underleaves not conspicuously more closely im- bricate than in main axis (exc. in some plants of C. humilis var. suspecta, q. v.). Sect. Clasmatocolea 13 13. Leaves with 17-32 regularly spaced l-2(-4)-celled teeth sub- equal in size C. ctenophylla (p. 1 19) 13. Leaves entire or if armed, then with 1-10 teeth or laciniae, armature irregularly spaced, commonly several- or many- celled, and only rarely with teeth consistently 1-2-celled. . 14 14. Underleaves consistently narrower than stem 15 15. Leaf margins entire; leaf cell trigones usually absent to small, occasionally medium; terminal branching occasional; opposing male bracts free. Perianths inflated, clavate-campanulate, distinctly or ob- scurely trigonous at the base, obscurely so toward the apex, mouth truncate, wide, the three lobes rotund, entire, occasionally emarginate C. vermicularis (p. 96) 15. Leaf margins with several, often incurved, teeth; leaf cell trigones large to knotlike; terminal branching ENGEL: CLASMATOCOLEA 43 absent; opposing male bracts fused toward base. . . C. minutiretis (p. 133) 14. Underleaves (0.9-)1 .0-4.3 X stem width 16 16. Underleaves with an associated, raised, flat, disci- form rhizoid initial field ; axis apices and intercalary portions very commonly enlarged; perianth keels with wings large and conspicuous, occasionally sinu- ate, the perianth mouth rather densely denticulate with teeth predominantly uniseriate C. gay ana (p. 112) 16. Underleaves without an associated rhizoid field; axes of approximately same width throughout; peri- anth keel wings, if present, never large and con- spicuous, never sinuate, perianth mouth entire, dentate or laciniate, never consistently denticulate with teeth predominantly uniseriate 17 17. Underleaf lamina margins usually with 1 large often apiculate lacinium, margins otherwise entire or sparingly dentate to ciliate to laciniate; perianths restricted to short ventral-intercalary branches, the perianths slightly narrowing to- ward mouth. Branches usually with first 1-2 pairs of leaves bifid C. fasciculata (p. 91) 17. Underleaf lamina margins with 1 tooth or lobe or entire, without 1 large lacinium; perianth posi- tion variable, on main axis or on short to long intercalary branches, but never restricted to short ventral-intercalary branches, the perianth with sides straight or gradually expanding to- ward the mouth 18 18. Underleaves to 1.3 X stem width, plane and not convex (ventral view); stems 5-6 cells high; plants often corticolous. Perianth terete, the ventral side not infolded C. crassiretis (p. 1 24) 18. Underleaves to 6.7 X stem width, slightly to deeply convex to cucullate (ventral view); stems 6-14 cells high; plants terricolous, saxicolous, or on rotted logs 19 19. Axes not moniliform, to 1.8 mm. wide, larger; branching with lateral-inter- calary type common, Frullania-type oc- casional; leaf cells thin walled; leaf in- sertion slightly to distinctly recurved at ventral end C. humilis (p. 74) 19. Axes moniliform, to 1.4 mm. wide, smaller; branching with lateral-inter- calary type absent, Frullania-iype very rare, ventral-intercalary type common; leaf cells distinctly thick walled; leaf 44 FIELDIANA: BOTANY insertion not recurved at ventral end. Underleaves cucullate, inflated, with apices closely appressed to the stem . . . C. moniliformis (p. 128) Subg. PROTOCLASMATOCOLEA Engel, subg. nov. Folia caulina bifida ad 0.15-0.3, plerumque longitudo magis quam latitudine et dis- posita versus axis apicem; cellulae foliorum trigonae minutae ad medias; amphigastria caulina saepe semioblique inserta, bifida e 0.5 ad plerumque prope basin; perianthiarum lobi bifidi. Type species: Clasmatocolea rigens (Hook. f. & Tayl.) Engel. Leaves bifid to 0.15-0.3, usually longer than wide and oriented toward axis apex; trigones minute to medium; underleaves often semiobliquely inserted, bifid from 0.5 to usually near the base; perianth lobes bifid. Distribution. — Falkland Is. and Dependencies, southern South America. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. Figures 7-9, Plates 1, 2. Jungermannia rigens Hook. f. & Tayl. London J. Bot. 3:461. 1844. Cephalozia rigens (Hook. f. & Tayl.) Trev. Mem. 1st. Lomb. Sci. Lett. III. 4:417. 1877. Lophocolea rigens (Hook. f. & Tayl.) Evans, Bull. Torrey Bot. Club 25:423. 1898. Clasmatocolea rigens (Hook. f. & Tayl.) Engel, J. Hattori Bot. Lab. 36:156. 1973. Original material: Falkland Is., Hooker (BM !). Lophocolea koeppensis Gott. Ergebn. Deutsch. Polar-Exped. 2 (16):453. pi. 2, /. 4-9. 1890, syn. fide Engel (1973a). Clasmatocolea koeppensis (Gott.) Grolle, Rev. Bryol. Lichenol. 29:72. 1960. Lectotype (fide Grolle, 1972b): South Georgia, Koppenberg, 10 February 1883, Will 35 (Ml). Lophocolea debilis Steph. Kongl. Svenska Vetenskapsakad. Handl. 46 (9):42./ 19 a-c. 1911, syn. fide Engel (1973a). Lectotype (fide Engel, 1973a): Chile, Prov. Magallanes, S. Skyring, Pta. Eulogio, 22 April 1908, Halle & Skottsberg 196 (UPS!; isolectotype : G!). Lophocolea stephanii Herz. Hedwigia 64:5. / 2. 1923, syn. nov., non L. slephanii Schiffn. Hep. Fl. Buitenzorg p. 181. 1900. Holotype: Chile, Prov. Valdivia, Valdivia, December 1911, Herzog (JE!). Lophocolea olens Herz. Arch. Esc. Farm. Fac. Ci. Med. Cordoba 7:16. / 5. 1938, syn. nov. Lectotype (AZOV.): Chile, Prov. Cautin, Pucon, "Halbinsel," Hosseus 140 (JE!). Plants rather fragile (especially small fades), prostrate, but axes often sharply curved dorsally, often in dense, thick mats, usually pale green, occasionally light brown, nitid when dry; axes to 2 mm. wide with leaves spread, small fades 200-460 /u wide. Branches mostly of Frullania-O^e, often slender and copiously produced, associated ENGEL: CLASMATOCOLEA 45 FIG. 7. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. 1, Portion of main axis, small facies of species, lateral view. 2, Leaf lobe, apical portion, roughened cuticle shown at lower right. 3, Leaf lobe. 4, Portion of main axis, lateral view. 5, Stem, cross section. 6, Median leaf cells. 7, Underleaf segment, the tip cell a partially collapsed slime papilla. 8, Leaves. 9, Underleaves. 10, 11, Underleaves, note oblique insertion. Figures 1, 3, 5-8a-b, d-e, from type of C. rigens, Hooker, Falkland Is.; figures 2, 8f, from Engel 1906, Chile, Prov. Magallanes, E. of Mina Loreto; figure 4, from Engel 1987, Chile, Prov. Magallanes, 8 km. W. of Punta Arenas; figure 8c, from Hatcher 4-13, Chile, Prov. Magallanes, near Fuerte Bulnes; figures 10, 11, from Engel 2008, Chile, Prov. Magal- lanes, B. Camden. half leaf usually undivided (but occasionally bifid), =«= triangular-ovate, apiculate, and with margins often sinuous; lateral-intercalary branches fairly common; ventral-inter- calary branches rare; intercalary branches occasionally short, reduced, and copiously produced. 46 FIELDIANA: BOTANY FIG. 8. Clasmalocolea rigens (Hook. f. & Tayl.) Engel. 1, Axis with perianth. 2, 3, Peri- anth lobes. 4, Outer capsule wall cells to symbolize pigmented (darkened) and hyaline (unshaded) thickenings, note few semiannular bands. 5, Capsule wall, cross section. 6, Inner capsule wall cells. Figures 1, 2, from Engel 2641, Falkland Is., Mt. Usborne; figures 3, 5, from Engel 1827, Chile, Prov. Magallanes, Pta. Santa Ana; figures 4, 6, from Engel 1815, Chile, Prov. Magallanes, Pta. Santa Ana. Stems 5-8 cells high, cortex in a single row of cells slightly smaller than, the same size as, or slightly larger than the medullary cells, exposed wall thin or moderately thickened, the cortex otherwise thin walled. Rhizoids in fascicles from stem near underleaf base. Leaves usually longer than wide (wide ovate leaves commonly wider than long), oriented toward axis apex and thus with dorsal margin longer than the ventral, the inser- tion distinctly recurved at ventral end; leaves erect, (distant-) loosely imbricate, adaxi- ally concave (a few convex leaves sporadically present, particularly on robust plants), oblong to medium-wide ovate; leaves bifid to 0.15-0.30, the lobes medium-wide triangular and usually apiculate, often incurved, often clawlike, the sinus narrowly-broadly rounded to lunate, occasionally triangular; margins entire or occasionally \ -dentate towards the apex, rarely 1-laciniate-lobate; dorsal margin often more broadly rounded than the ventral, often sinuous, commonly long decurrent; ventral margin often subauriculate at the base. Leaf cells thin-walled, trigones minute to medium; median leaf cells 14-34^ wide, (13-) 18-38 fj. long; cuticle smooth to roughened and appearing finely granular. ENGEL: CLASMATOCOLEA 47 FIG. 9. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. Underleaves usually slightly wider than stem, often obliquely inserted, moderately to distinctly spreading, remote to contiguous, bifid from ca. 0.5 to usually near the base; segments curved dorsally, particularly at the apices, the segments lanceolate-subulate, occasionally medium-triangular, often apiculate, terminating in a slime papilla, the seg- ment margins entire (-1-2-dentate-ciliate); margins of the lamina entire or usually 7-(to rarely 3) dentate-laciniate-subulate, the armature terminating in a slime papilla. Plants dioecious; androecia terminal or intercalary on main axis, Frullania-type, or lateral-intercalary branches; bracts erect and oriented toward axis apex as in leaves, strongly saccate, apical portion slightly spreading but with lobes incurved, the apices often appressed to bract immediately above (in small plants); lobule margin with several slime papillae and often few-celled teeth terminating in slime papillae, or with 1 lacinium, or 1 large, acute or rounded involute lobe; antheridia solitary, rarely 2 per bract, stalk uniseriate throughout, very rarely with a localized bistratose area. Gynoecia on main axis, rather long Frullania-type or lateral-intercalary branches 48 FIELDIANA: BOTANY and subfloral innovations; subfloral innovations nearly always present, originating from below gynoecium, below perianth or from inside perianth; bracts in 3-4 series; those of innermost series oblong-ovate ; apices bifid and with lobes incurved ; margins entire or few dentate-ciliate-laciniate. Bracteoles of innermost series free from bracts, 0.2-0.5-bifid; lamina margins slightly to distinctly recurved, entire-few dentate-laciniate. Perianths often produced, strongly trigonous, elongate-subrectangular to oblong or occasionally =*= campanulate, slightly expanding to slightly narrowing toward mouth, the mouth closed by slightly incurved lobes or in ± campanulate-shaped perianths by infolding of ventral lobe; lobes rounded, bifid, sparingly to usually densely dentate- ciliate-laciniate, very rarely subentire; keel wings common, of a few cells high, occa- sionally dentate. Seta not studied. Capsule valves 530-950 ^ long, 41-53 n thick, of 4-6 layers, outer row of cells about equal to thickness of innermost 3 strata ; outer layer with vinaceous or colorless, nodule-like or ± spinelike thickenings that are often feebly tangentially dilated, a very few semiannular bands occasionally present; exposed wall thickened; outermost intermediate layer of cells thicker than inner layer, the intermediate layers with thickenings feebly extending onto tangential walls; inner layer of cells with yellow- to red-brown, semiannular bands, the bands often incomplete, often forked, the radial walls with nodulose thickenings often present. Spores 14-18 /n, red-brown, exine with the light microscope appearing minutely punctate, but under the SEM with granulate to short, abbreviated vermiform projec- tions that have nanogranules ; spores averaging 2.4 X elater diameter. Elaters 6-7 /z wide, spiraled to tips, walls light brown. Variation. — The original material of Jungermannia rigens represents a not infrequently occurring very small fades of the species. This facies superficially resembles the genus Cephalozia, which likely accounts for its transfer to that genus by Trevisan de Saint-Leon (1877). The underleaves of this facies have margins entire or with a few small teeth and are not lacinate-subulate as is common in larger, more robust plants. Because this facies is part of a continuum with medium and larger individuals with increasingly larger underleaf armature, I do not recognize it taxonomically. Differentiation. — The combination of bifid leaves that are usually longer than wide and oriented toward axis apices (and thus with dorsal margins considerably longer than the ventral) and the medium-wide triangular, apiculate leaf lobes will serve to separate this species from all others of the genus. Further, in no other species are the underleaves bifid nearly to the base; underleaves in other Clasmatocolea taxa are, for the most part, undivided or bifid to at most 0.5. Larger plants of C. rigens resemble plants ofLophocolea lenta (Hook. f. & Tayl.) G. L. & N., but the former may be immediately distinguished by the possession of (a) at least some adaxially concave leaves on a given axis; (b) underleaves often obliquely inserted; and (c) dioecious inflores- cences. Lophocolea lenta, on the other hand, has (a) convex leaves; (b) ENGEL: CLASMATOCOLEA 49 underleaves consistently transversely inserted; and (c) monoecious in- florescences. Notes. — 1. In one specimen (Engel 1815), two mature sporophytes were observed arising from a single inflorescence. According to Schuster (1966), this phenomenon exists in the primitive orders Calobryales and Mono- cleales, but is of only isolated occurrence in other groups. 2. Lophocolea subaromatica Herz. (Arch. Esc. Farm. Fac. Ci. Med. Cordoba 7:15. 1938) described from Pucon (Chile, Prov. Cautin) may belong here; I have not seen type material of this species. Ecology. — In southern South America, seemingly characteristic of de- ciduous forests and boundaries between deciduous and evergreen Nothofa- gus forests. Rare in evergreen forest regions and then only in the drier portions. It is absent, for example, in the wet, western part of Tierra del Fuego as well as the wet western Patagonian Channels of Prov. Magal- lanes. In the Valdivian region on bark or over rocks in Nothofagus forests or mixed Nothofagus pumilio-Araucaria forests. On the South Sandwich Islands within the main crater of Bellinghausen Island, where around but away from fumaroles, and on boulders moistened, but probably not warmed, by steam (130-200 m.). Also on the slopes of the main cones, but again, away from fumaroles (75-80 m.). Phytogeography. — South Sandwich Is. (75-200 m.); South Georgia; Falkland Is. (335-700 m.); Tierra del Fuego (sea level-300 m.; vicinity of Ushuaia and R. Azopardo); southern Patagonian Channels (Brunswick Pen., S. Skyring, Pta. Eulogia); Valdivian region (730-1,450 m.; West Patagonia from 45° 25' S. to 38° 39' S., Andean Patagonia at 43° 30' S.); Prov. Santiago (see fig. 9). Specimens seen.— SOUTH SANDWICH IS.: Bellinghausen I., 75- 130 m., Holdgate 421 A, 430A, 816 as C. koeppensis (hb. Grolle); ibid., 75-200 m., Holdgate 431 A - c. /3, /. 20-24. 1890,^. fide Stephani (1906). Holotype: Chile, Prov. Magallanes, Punta Arenas, 7 February 1876, Naumann s.n. (FH! - c. per.). Lophocolea lacerata Steph. Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6): 41. 1900, syn. fide Stephani (1906). Original material: Chile, Prov. Magallanes, Pto. Bueno, 31 May 1896, Dusen 35 (NY!); Prov. Aisen, R. Aisen Valley, Dusen (FH!, G!, NY!, S! - 7 coll.). Blepharostoma acanthifolium Gola, Nuovo Giorn. Bot. Ital. II. 29:169. pi. l,f. 18-19. 1923, syn. fide Engel (1978). Original material: Chile, Prov. Magallanes, I. Laberinto, 5 February 1913, Gasperi s.n. (FI!). Plants somewhat hairy in appearance, prostrate, mixed with other bryophytes or in dense, interwined, often pure, thick cushions, pale grey-green or very pale green or light brown tinged, often becoming brownish throughout in older collections; main axes to 1.3 mm. wide. Branches copiously produced, often densely interwined, Frullania- and lateral-inter- calary type common, the latter especially so; ventral-intercalary type very rare. Stems 6-10 cells high, cortex in 1 row of slightly to distinctly thick-walled cells slightly smaller than, equal to, or larger than medullary cells, the cortex often with 1-2 rows of ventral cells smaller or equal in size to medullary cells, and with dorsal cortical cells larger than those of the medulla; medullary cell walls thin to slightly thickened; endophytic hyphae absent. Rhizoids in fascicles from stem near underleaf base. Leaves with insertion broad, narrow in weaker forms, oblique, not or slightly recurved at ventral end ; leaves strongly erect, approximate to densely imbricate, ventral-median FIG. 14. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. 1, Main axis with 3 lateral-intercalary branches (lib) and perianth on lateral branch. 2, Distinctly bifid perianth lobes, ventral lobe in middle. 3, Obscurely bifid perianth lobes, ventral lobe in middle. 4, Antheridial stalk. 5, Stem of main axis, cross section. 6, Opposing anther- idial bracts with stem(s), cross section. Figure 1 , from Engel 4850, Chile, Prov. Magal- lanes, Pto. Charnia; figure 2, from Engel 6249, Chile, Prov. Magallanes, B. Pond; figure 3, from Engel 4381, Chile, Prov. Aisen, F. Tempano; figures 4, 6, from Engel 4814, Chile, Prov. Magallanes, Pto. Charnia; figure 5, from Engel 5106, Chile, Prov. Magallanes, I. Tarlton. 61 -.'2-4.6-9 FIG. 15. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. 1, Median leaf cells. 2-4, Underleaves. 5, Apical portion of median lobe of leaf showing submoniliform uniseriate portions. 6-9, Leaves, abaxial (ab) and adaxial (ad) views (each number showing 2 views of same leaf); figures 1, 2, 6, 7, from type of C. trachyopa, Hooker, Chile, Prov. Magallanes, I. Hermite; figures 3, 5, from Engel 4831, Chile, Prov. Magal- lanes, Pto. Charrua; figure 4, from Engel 3985C, Chile, Prov. Osorno, L. Toro; figure 8, from Engel 4814, Chile, Prov. Magallanes, Pto. Charrua; figure 9, from Engel 4134, Chile, Prov. Osorno, R. Nauto. 62 ENGEL: CLASMATOCOLEA 63 FIG. 16. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. 1, Inner capsule wall cells. 2, Outer capsule wall cells, note 2 semiannular bands. 3, Capsule wall, cross section. 4, 5, Setae; cross sections of two different setae from same collection. Figures 1 , 3, 4, from Engel 5203, Chile, Prov. Magallanes, I. Madre de Dios; figure 2, from Engel 4850, Chile, Prov. Magallanes, Pto. Charrua. portion with a conspicuous, abrupt, concavity (the leaves often weakly concave in poorly developed axes), the leaves in abaxial view often with 2 elongate, sharp convexities that are proximal extensions of the canaliculate lobes; leaves obscurely to distinctly obtrapezoid or less often wide-ovate, basically 2-3-lobed, but often with 1-3 accessory lobes, the dorsal lobe acuminate, erect or incurved in a broad bowlike arc, with margins recurved and lending the lobe canaliculate, often with margins connivent toward base and then lobe here locally tubiform; dorsal lobe with several spinescent teeth or short cilia and often a few sharp laciniae (which may be toothed), the armature occurring in opposite pairs but commonly with a few interspersed solitary teeth, the dorsal-basal portion of dorsal lobe (or more rarely the dorsal margin immediately proximal to lobe) with a large tooth, or dentate lacinium, or dentate, canaliculate accessory lobe commonly situated at =*= right angles to the dorsal lobe, the armature nearly absent in depauperate axes, the uniseriate portions of dorsal lobe and lobe armature often submoniliform, the nonuniseriate portions often =*= crenulate; dorsal sinus 0.5-0.75 the leaf length, deeper than ventral sinus, the tissue at sinus base frequently recurved at the base and often with 1-2 spinescent teeth; median lobe (in basically trifid leaves, or ventral lobe FIG. 17. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. 64 ENGEL: CLASMATOCOLEA 65 in bifid leaves) =*= acuminate to narrowly triangular, erect or incurved in a broad bow- like arc, with margins often recurved lending the lobe canaliculate, the median lobe with characters of armature as in the dorsal lobe; ventral sinus, when present, 0.25-0.5 the leaf length, the tissue at sinus base plane or recurved, often with a tooth; ventral lobe (in trifid leaves) variable in size, much smaller than dorsal or median lobe, the margins plane or recurved and then lobe at most moderately canaliculate, the lobe entire or with a few pairs of opposite teeth, the uniseriate portion often submoniliform; dorsal margin of lamina usually =*= straight or occasionally rounded near base, recurved, entire or with a few teeth, short decurrent; ventral margin of lamina plane or if in- curved, then with ventral portion of leaf ± saccate, the ventral margin narrowly or broadly rounded, with a tooth or lacinium near base of lobe, the margin otherwise entire or with a few teeth or laciniae. Leaf cells thin to moderately thick-walled, trigones small to large and bulging; median leaf cells of disc 17-36 /x wide, 18-48(-53) /* long; cuticle smooth or slightly roughened and appearing finely granular. Underleaves 1.3-2(-3.5) X stem width, free or connate on 1 side, slightly to moder- ately spreading, approximate to densely imbricate, plane to moderately convex (ventral view) but not abruptly so, the margins occasionally curved ventrally, especially so toward base; underleaves ovate, rarely subrhomboid, with apices 0.25- to slightly less than 0.5-bifid; segments with uniseriate portion often =*= submoniliform, terminating in a slime papilla, the segment margins entire; margins of the lamina entire to 3-dentate- small laciniate, the armature often terminating in a small slime papilla. Branch under- leaves strongly spreading, deeply convex to ± cupulate. Plants dioecious; androecia terminal or intercalary on main axis or abbreviated to long lateral-intercalary or Frullania-lype branches; bracts with basal portion strongly saccate, the saccate portion fused toward base with opposite bract, the distal portion of bract with lobes and lobe armature as in leaves, or if bracts bifid, then with lobes plane, entire, and with ventral margin broadly rounded and entire or 1 -dentate; lobule margin involute, with several slime papillae and often few-celled teeth, or with 1 lacinium or broadly rounded involute lobe; antherida solitary, stalk uniseriate. Gynoecia usually on short, abbreviated lateral-intercalary branches, very rarely on main axis; subfloral innovations absent or when present (rare) from below innermost bract, the innovations very short and producing a perianth; coelocaule precursor present; bracts and bracteoles in 3 series, inserted on coelocaule precursor; bracts of innermost series slightly convex or concave, especially in upper portion, obovate to subspatulate to occasionally wide-ovate; apices 2-3-lobed, if bifid, then divided to slightly over 0.5, or if trifid (occasional), then dorsal sinus extending to slightly over 0.5 bract length and the ventral sinus to 0.3 bract length; dorsal lobe with margins consistently reflexed, occasionally strongly so and then lobe locally tubiform ; ventral lobe and, when present, median lobe occasionally with margins reflexed, occasionally strongly so and then lobe(s) locally tubiform; apex of lobes with several small to large and spinescent teeth or short cilia (and occasionally a dentate lacinium), the teeth and cilia in opposite pairs, the dorsal-basal portion of dorsal lobe (or more rarely dorsal margin immedi- ately proximal to lobe) usually with a small dentate, occasionally canaliculate lobe, the accessory lobe occasionally absent and then bract commonly ± bilaterally symmetric; lamina margins with several teeth, especially in upper portion. Bracteoles of innermost series subequal to bracts in size, free from bracts, slightly convex, or slightly to dis- tinctly concave (ventral view), the bracteoles elliptic to obovate to subspatulate; apices 66 FIELDIANA: BOTANY 0.2-0.4-bifid; segment margins plane or if reflexed, then with lobes moderately to strongly canaliculate, with several small to large and spinescent teeth or short cilia (and occasionally a dentate lacinium), the teeth and cilia in opposite pairs; lamina margins with several small teeth (especially in upper portion), often with a small lobe in upper portion. Perianth strongly trigonous, narrowly ovate to narrowly elliptic to elongate-subrectangular, commonly with 2 or all 3 sides wholly or in part infolded near mouth, but occasionally only with ventral side strongly infolded, the perianth gradually narrowing or straight or gradually expanding toward mouth, the mouth compressed due to infolded lobes, occasionally bilaterally so; lobes broadly rounded, usually ob- scurely to distinctly bifid, the segments narrowly acuminate and often with margins reflexed, occasionally strongly so and lobe locally tubiform, the segment margins armed with a few to several pairs of opposite teeth, the lobes otherwise with several teeth and elongate cilia; lobes occasionally undivided, and then either with a few small teeth, or copiously dentate to elongate-ciliate and with a few small, dentate, accessory lobes; wings often present, of several cells high and occasionally with a few teeth. Sporophyte with foot "penetrating" into axial proliferated tissue to level of ca. bract of first series or first leaf below. Seta (6-)7-9 cells in diameter, with 1 7-30 rows of outer cells with inner corners thickened similarly to medium to large and bulging trigones, the inner core of cells considerably smaller than or subequal to outer row in size, the core cells gradually increasing in size toward center, the core cells thin-walled, with corners thickened similarly to small to medium trigones. Capsule ± globose to elliptic to ovate, the valves 0.7-1.15 mm. long, 29-48(-52) ^ thick, of (3-)4(-5) layers, outer row of cells equal to thickness of 1.8-2.5(3) of interior strata; outer layer with vinaceous, nodule-like or spinelike thickenings that are often feebly tangentially dilated, a few to several semiannular bands present; exposed wall thickened; intermediate and inner layers of cells subequal in thickness, the intermediate layers with thickenings often considerably tangentially dilated; inner layer of cells with red-brown semiannular bands, the bands often nonpigmented in median portion, incomplete, the radial walls with nodulose thickenings often present. Spores 11-14/i, light brown, exine with the light microscope appearing minutely punctate, but under the SEM with a network of very wide, irregular, coalescing vermi- form ridges; spores averaging 1.4 X elater diameter. Elaters 6-11 n wide, apical por- tions often with thick nonspiral walls, elater walls red-brown or yellow-brown. Differentiation. — Clasmatocolea trachyopa is rather closely related to C. obvoluta var. obvoluta; for further notes regarding this relationship see p. 57. Clasmatocolea trachyopa represents the culmination of a trend in leaf complexity in Subg. Lacerifolia beginning with C. obvoluta var. cookiana, through C. obvoluta var. obvoluta to C. trachyopa. Specimens of C. trachyopa may be identified at once by the very distinc- tive general aspect of the plant (see fig. 14-1). The dorsal and median leaf lobes are usually canaliculate due to the recurved margins. As a result of these recurved margins, the opposing pairs of lobe armature are ventrally oriented and at right angles to the leaf lamina plane. Conspicuous also is the commonly dentate and canaliculate lacinium or lobe at the dorsal- basal portion of the dorsal lobe. This lacinium or lobe usually projects at right angles to the dorsal lobe but may arc toward the dorsal lobe. No ENGEL: CLASMATOCOLEA 67 other species of Clasmatocolea has the degree of leaf complexity observed in C. trachyopa. For example, both the development of a three-lobed leaf as well as accessory lobes at leaf apex and margins are unique to the genus. Notes. — 1. A problem of typification arises from the fact that the original material of Jungermannia trachyopa deposited in the Farlow Herbarium (hb. Taylor) is a mixture of two Clasmatocolea species — a few stems of C. obvoluta that are sterile and a comparatively large amount of C. trachyopa that has both perianths and sporophytes. The sheet on which the specimens and packet are mounted bears hand-drawn illustrations of (a) a portion of an axis with two leaves and an underleaf; (b) an underleaf; (c) a perianth; and (d) a perianth cross section. Although it is obvious that that portion of the original description dealing with the perianth and sporophyte in Hooker & Taylor (1844) pertains to C. trachyopa, I am uncertain whether the description of sterile gametophyte structures per- tains to C. obvoluta or poorly developed axes of C. trachyopa (which are present in the type collection). This is particularly the case because the description of the gametophyte matches the figure of the leaves and underleaf on the herbarium sheet, and this figure appears much like C. obvoluta. Further, a syntype of J. trachyopa deposited in the Stockholm Herbarium (S, ex hb. Lehmann) consists solely of a few stems of C. obvoluta with no C. trachyopa present. However, Taylor & Hooker (1847) have effectively typified the species because their description of Jungerman- nia trachyopa clearly refers to plants with 2-3-lobed, unequally spinose- dentate leaves, and the illustrations for the species clearly show spinose- dentate leaves. Both the description and illustrations exclude material of C. obvoluta. I have selected the material of C. trachyopa as lectotype, because the rules state [I.C.B.N. (1972, p. 76)]: "In choosing a lectotype, any indication of intent by the author of a name should be given preference unless such indication is contrary to the protologue" (see also I.C.B.N., Art. 70). 2. As mentioned above, some of the axes in the lectotype collection are depauperate. In these axes the leaves are only slightly concave, the margins of the dorsal and median lobes are only slightly reflexed, the lobes are entire or have one opposing pair of teeth, and the basal-dorsal portion of the dorsal lobe is entire or has one small tooth (i.e., the leaves lack the conspicuous lacinium or lobe that commonly projects at right angles to the dorsal lobe) (see fig. 15-6). The depauperate condition is rather rare. 3. Clasmatocolea trachyopa, like C. obvoluta, possesses a coelocaule precursor. In C. trachyopa there is a swollen region on which the perianth and three series of bracts and bracteoles are inserted. The sporophyte 68 FIELDIANA: BOTANY "penetrates" to about the level of the bracts of the first series or the leaves immediately below. 4. This species, like C. obvoluta, has androecial bracts with saccate portions fused toward the base with the bract on the opposing side of the stem (fig. 14-6). This condition totally obliterates a dorsal view of the stem. 5. All observed 1897 Dusen specimens labeled Lophocolea trachyopa from Islas Guaitecas are misdeterminations of C. obvoluta var. obvoluta or C. obvoluta var. cookiana. Ecology. — On rotted logs and stumps (often those covered with bryo- phytes) or occasionally corticolous on trees in forests, commonly where there is at least moderate amounts of shade. It sometimes occurs with Lepidolaena magellanica or Clasmatocolea humilis, but more often it is associated with Riccardia sp. Phytogeography. — Tierra del Fuego (including I. Grande de Tierra del Fuego); Patagonian Channels; Valdivian region (100-885 m.) N. to 39° 56' S. (Prov. Valdivia) (see fig. 17). Specimens seen.— ARGENTINA. TERR. TIERRA DEL FUEGO: I. de los Estados, Spegazzini 442, 99l (VER); ibid., Pto. Cook, Spegazzini 28i (VER); ibid., Spegazzini 45* (NY, VER); ibid., Spegazzini 320 (VER). CHILE. PROV. MAGALLANES: Cabo de Hornos, Hooker s.n. as syntype of /. trachyopa (NY) ; (western) L. Fagnano, Skottsberg 235 (S) ; I. Clarence, S. shore of B. Pond, Engel 6249 — c. sporo. + & , 6283 — c. sporo. + cT (MSC); I. Desolacion, Dusen s.n. as L. lacerata (S); ibid., Pto. Angosto, Dusen 256 as L. lacerata — c. per. (S); I. Desolacion, Pto. Churruca, N. side of Fo. Nassau, Engel 5942 — c. sporo. + = cupulate and (when flattened) ca. 3.3-4 X the stem width. This species appears to be related to C. strongylophylla of New Zealand and Tasmania, which also has (a) coarse, hemispherical papillae on the leaves; (b) long decurrent dorsal margins of leaves; (c) thick-walled stem cells that are of a comparatively few number of cells high ; and (d) poly- morphous underleaf apices — either undivided or distinctly bifid. Clasmat- ocolea tjiwideiensis may be distinguished from C. strongylophylla by (a) the connate underleaves; (b) the leaf armature; (c) the underleaves that are deeply convex (ventral view) to =*= cupulate and wider, being ca. 3.3-4 X the stem width; and (d) the larger median leaf cells. Notes. — 1. The underleaves are connate on one or both sides with the ventral-basal, adaxial surface of the leaf lamina. This feature is often a subtle one, and to observe it, an axis should be examined in both ventral and dorsal views while slowly and carefully rotating the axis. When connate on both sides, the attachment on one side is usually more obvious than on the other, depending on distance of the attached leaf. For this reason, where leaves are more distant on an axis, the attachment on one side disappears, and here the underleaves are connate on only one side ; the character should be studied on well-developed portions of the axis with leaves more closely imbricate. 2. This species has not been collected since the mid-nineteenth century. 511-14 156 ENGEL: CLASMATOCOLEA 157 Ecology-Phytogeography. — Endemic to Java, where at least sometimes corticolous; altitudes of ca. 1,830 m. are recorded. Specimens seen— JAVA: Mt. Salak, 1,830 m., Kurz s.n., 177 (L). Subg. METACLASM ATOCOLEA Engel, subg. nov. Folia caulina indivisa, Integra; amphigastria principalis axis vel rami laterales inter- calares inconspicui, reducti, squamiformi, polymorphi. Type species: Clasmatocolea navistipula (Steph.) Grolle. Lateral-intercalary branches often copiously developed, Frullania-type branching rare; leaves undivided, entire; underleaves of main axis and/or lateral-intercalary branches inconspicuous, reduced, scalelike, polymorphous, often of only a few cells. Distribution. — Southern South America. Sect. PUCCIOANAE Engel, sect. nov. Haec amphigastria caulina principalis axis bene manifesta, ilia ramorum lateralorum intercalarorum inconspicua atque reducta; rami laterales intercalarores copiose effecti, incrementi parvi, plerumque abbreviati. Type species: Clasmatocolea puccioana (De Not.) Grolle. Underleaves of main axis well developed, those of lateral-intercalary branches inconspicuous and reduced; lateral-intercalary branches copiously produced, of limited growth, usually abbreviated. Distribution. — Southern South America. Clasmatocolea puccioana (De Not.) Grolle. Figures 49-51, Plates 23-25. Jungermannia ?puccioana De Not. Mem. Reale Accad. Sci. Torino II. 16:221. pi. IX, 1-6. 1855. Lophocolea puccioana (De Not.) Mass. Nuovo Giorn. Bot. Ital. I. 17:227. 1885. Lophocolea puccioana a* forma primigenia Mass. Nuovo Giorn. Bot. Ital. I. 17:227. 1885. Lophocolea puccioana var. & primigenia (Mass.) Schiffn. in Naumann, Forschungsr. Gazelle 4 (4): 13. 1889. Clasmatocolea puccioana (De Not.) Grolle, Rev. Bryol. Lichenol. 29:72. 1960. Original material: Chile, "Prov. Valparaiso, Valparaiso," Puccio (non vidi; GE = O, NY = O, PAD = O, PG = O, RO = O, TO = O). Opposite: FIG. 48. Clasmatocolea tjiwideicnsis (Sande Lac.) Grolle. 1-2, Main axis, lateral view (note position of leaf dentition). 3, Main axis, ventral-lateral view. 4, Main axis, showing leaf and underleaf insertion, ventral view. 5, 6, Underleaves. 7-9, Leaves (flattened). 10, Stem, cross section, note irregularly shaped cell lumina. 11, Median leaf cells with cuticular papillae indicated by stipple; three secondary cell wall thickenings indicated. 12-14, Teeth from ventral-basal portion of leaf, with cuticular papillae indicated. Figures 1-3, 6-14, from type of Chiloscyphus tjiwideiensis, Junghuhn, Java; figures 4, 5, from Kurz s. n., Java. 33.4 158 ENGEL: CLASMATOCOLEA 159 Lophocolea diver sistipa Steph. Kongl. Svenska Vetenskapsakad. Handl. 46 (9):42./ 15 e, f. 1911, syn. fide Engel (1978). Original material: Chile, Prov. Magallanes, Pto. Gray, 7 June 1908, Skottsberg 198 (UPS! — c. per.); Pto. Ramirez, Halle and/or Skottsberg (non vidi); Pto. Cutter, 13 April 1908, Halle & Skottsberg 198 (UPS!). Lophocolea patagonica Beauv. in Steph. Spec. Hep. 6:572. 1924 ut nom. nov. pro Lophocolea rotundistipula Steph. Kongl. Svenska Vetenskap- sakad. Handl. 46 (9):52./ 20 a-e. 1911, syn. fide Engel (1978) non L. rotundistipula Steph. Bull. Herb. Boissier 6 (9): 793. 1906 (= Spec. Hep. 3:93). Original material: Chile, Prov. Magallanes, F. Peel, 16 June 1908, Skottsberg s. n. (UPS!). Plants prostrate, commonly mixed with other bryophytes, occasionally in ± pure tufts, light brown-sienna, stems often rather deep orange-brown; main axes 560-910 n wide. Branches copiously developed, somewhat less copious in very elongated axes; lateral- intercalary type in great abundance, much narrower in width than main axis, the branches short, of limited growth, subvermiform, sometimes =*= erect, delicate, usually not in turn branched, often originating from near ventral end of insertion of associated leaf, the lateral-intercalary branches sporadically becoming leading axes with their own copi- ously developed short, lateral-intercalary branches (this leading axis may, in turn, produce another leading axis); ventral-intercalary branches very rare (2 seen), like lateral-intercalary branches; Fru\\ania-type branches rare. Stems 9-12 cells high, cortex in 2-3 rows of very thick-walled cells smaller than, or ca. equal in size to the medullary cells, the exposed wall of outer row very thickened; medullary cell walls thin to slightly thickened, with corners thickened as in medium trigones; endophytic hyphae present in both cortex and medulla. Rhizoids frequently present, rather long, in fascicles from stem near underleaf base. Leaves with insertion distinctly recurved at ventral end; leaves erect to slightly spreading, occasionally connivent dorsally, loosely imbricate, conchiform concave, suboblate to subreniform to broadly ovate; apex broadly rounded to truncate, undivided, entire, exceptionally with a single large tooth; margins entire; dorsal margin ± straight in proximal half, sinuous; ventral margin broadly rounded, sometimes somewhat in- curved, usually extending ventrally beyond stem and nearly but usually not in contact with underleaf. Intercalary branch leaves 0.25-0.5 the size of the leaves, often slightly more closely imbricate than in main axis, entire. Opposite: FIG. 49. Clasmatocolea puccioana (De Not.) Grolle. 1, Main axis showing lateral- intercalary branching pattern. 2, Lateral-intercalary branches and adjacent main axis. 3, Median leaf cells. 4, Stem, cross section. 5-7, Leaf and underleaves from lateral- intercalary branches. 8-9, Leaf and underleaf from main axis. Figures 1 , 2, from Engel 4272B, Chile, Prov. Aisen, Pto. Island; figures 3, 8, from Skottsberg 198, Chile, Prov. Magallanes, Pto. Gray (syntype of L. diversistipa); figure 4, from Engel 4520, Chile, Prov. Magallanes, Pto. Eden; figures 5, 9, from Engel 4515, Chile, Prov. Magallanes, Pto. Eden; figure 6, from Engel 4385 A, Chile, Prov. Aisen, F. Tempano; figure 7, from Engel 5248D, Chile, Prov. Magallanes, I. Juan. 160 FIELDIANA: BOTANY FIG. 50. Clasmatocolea puccioana (De Not.) Grolle. 1, Gynoecium on abbreviated lateral-intercalary branch ; lower portion an oblique longitudinal section, upper portion with merely a portion of perianth removed ; note position of unfertilized archegonia (ua) and bracts. 2, Capsule wall, cross section. 3, Outer capsule wall cells. 4, Inner capsule wall cells. 5, Seta, cross section. 6, Perianth mouth, ventral lobe in middle. Figures 1, 4, 5, from Engel 4541, Chile, Prov. Magallanes, Pto. Eden; figures 2, 3, from Engel 4520, Chile, Prov. Magallanes, Pto. Eden; figure 6, from Engel 5361B-2, Chile, Prov. Magallanes, B. Wide. Leaf cells thin to moderately thick-walled, the trigones large and moderately bulging to coarse and nodular, often confluent, commonly with leaf cell lumen, except for narrow thin-walled places, bounded mostly by the massive trigones; median leaf cells 1 1-20 n wide, 12-24 /* long; cuticle smooth. Underleaves 1.2-3.0 X stem width, free, moderately spreading, approximate, moder- ately convex (ventral view), elliptic to ovate to oblate; apices bidentate to broadly retuse to bifid to 0.2(-0.25), the segments of similar size and shape, very often apiculate, the segments or teeth at apex terminating in a slime papilla, the segment margins entire; margins of the lamina entire or median or median-basal portion with 1 small to large tooth or small lobe and occasionally a slime papilla, the margins rarely with 2 teeth, the armature terminating in a slime papilla. Underleaves of small, copiously produced intercalary branches, small, commonly inconspicuous, strongly spreading, often spreading to 90° or slightly to moderately recurved; Underleaves polymorphous, ovate to elliptic to ENGEL: CLASMATOCOLEA 161 subrect angular to =*= cuneate to obovate, the lamina often of only a few cells high; apices undivided or with a single tooth or lobe, sometimes bifid; lobes or teeth often asymmetric, terminating in a slime papilla, often with 1 lobe large and conspicuous and other lobe reduced to a tooth or slime papilla; lamina margins entire or with 1 tooth or slime papilla; underleaves of somewhat larger intercalary branches nearer to configuration of main axis underleaves. Plants dioecious; androecia on short, abbreviated lateral-intercalary branches, rarely terminal or intercalary on main axis or robust, well-developed lateral-intercalary branches; bracts with basal portion strongly saccate and distal portion strongly con- cave, the apices appressed to bract immediately above; lobule margin involute, with a rounded lobe possessing 1 tooth and/or a few slime papillae; antheridia solitary, the stalk uniseriate. Gynoecia on short to rather long lateral-intercalary branches; subfloral innovations absent; vestigial stem perigynium present; bracts in 3-4 series, the bracts and brac- teoles of innermost and third series inserted on the vestigial perigynium; bracts of innermost series slightly concave to slightly convex, broadly obovate to subrectangular; apices undivided, broadly rounded to truncate, entire; margins occasionally sinuous, occasionally reflexed, entire or with 1-few teeth. Bracteoles of innermost series sub- equal to or slightly smaller than bracts in size, free from bracts; bracteoles moderately concave to subcanaliculate (ventral view), broadly obovate to suboblate to subrec- tangular; apices undivided, broadly rounded, bidentate or with 2 slime papillae; lamina margins sometimes sinuous, with low, rounded projections or short, blunt lobes, the margins entire or occasionally with a few teeth. Perianth trigonous, narrowly ovate to elongate-rectangular, the ventral side slightly to strongly infolded nearly through- out, and then lending perianth =*= bilaterally compressed, or ventral side straight or slightly convex in median and basal portions and infolded in apical portion; perianth sides narrowing toward the =*= bilaterally compressed mouth; lobes broadly rounded, often with low, rounded projections, the lobes entire or with a few teeth; keels in apical region along with the adjacent portion of the lobe often sharply recurved or incurved ; keel wings sometimes present, of several cells high and sometimes with an auriculate appendage. Calyptra large. Seta 6-10 cells in diameter, with 28-33 rows of outer cells; median core cells larger than "epidermal" cells, the core cells becoming smaller toward periphery, with the outermost l(-2) rows of core cells ca. equal to or slightly smaller than "epidermal" cells; core cells with corners thickened similarly to small to medium trigones. Capsule narrowly elliptic to ovoid, the valves 0.74-1. 37 mm. long, of 36-42 p. thick, of 4-5(-6 locally) layers, the outer layer of cells equal to thickness of (2.5-)3.0-3.9 of interior strata; outer layer with vinaceous or red-brown, nodule-like or occasionally spinelike thickenings that are often feebly tangentially dilated, semiannular bands occasional; exposed wall of outer layer thickened ; intermediate and inner layers subequal in size or with outermost intermediate layer thicker than inner layer of capsule, the intermediate layers with thickenings often considerably extending onto tangential walls, semi- annular bands occasional; inner layer with red-brown semiannular bands moderately to very common, nodulose thickenings very common. Spores 1 1-12(-13) /*, light brown, exine with the light microscope appearing minutely punctate, but under the SEM with a network of short, wide vermiform ridges covered with sometimes coalescing nanogranules. the intervening walls with several, rather close nanogranules; spores averaging 1.5 X elater diameter. Elaters 7-9 n wide, apical portions often with thick nonspiral walls; elater walls red-brown. 162 FIELDIANA: BOTANY Variation. — The branch underleaves of C. puccioana and those of the main axis of C. navistipula are of comparable shape and tend to have similar variability of underleaf segments. Differentiation. — Clasmatocolea puccioana is closely related to C. navi- stipula; see the discussion (p. 171) under the latter for distinguishing the two taxa. Clasmatocolea puccioana shows some relationships to C.fulvella ; for a discussion, see that species. The primary distinctive features that will distinguish this species from all other members of the genus are those of branching patterns and branch characteristics; these are (a) the copiously developed lateral-intercalary branches that are much narrower in width than the main axis and that are short, of limited growth, subvermiform, delicate, and usually not in turn branched ; (b) the distinctive intercalary branch underleaves that are small, commonly inconspicuous, sometimes with a lamina of only a few cells high, strongly spreading, and with polymorphous apices and outline; and (c) the rarity of both ventral-intercalary and Frullania-type branching. Significant supplementary distinctive features are the 2-3-layered very thick-walled cortex and the light brown-sienna color. The quite charac- teristic color of this species can be used to identify it at a glance when the species is mixed with other Clasmatocolea taxa such as C. navistipula or C. obvoluta. Notes. — 1. Based upon the study of syntypes from all localities listed in Stephani (1911), Lophocolea diversistipa is a synonym of C. puccioana. In his original description, Stephani states that the plants are pale light green in color, and the branches are clustered. The specimens from Puerto Ramirez are misdeterminations of C. humilis, are light brown in color, and are sparingly branched. The specimens from Puerto Gray and Puerto Cutter, however, are C. puccioana and are light green to yellow-green in color and have numerous lateral-intercalary branches. Further, the Puerto Gray and Puerto Cutter specimens have underleaves with sinuses of the shapes given in Figures 15e and f of Stephani (1911), i.e., bifid to ca. 0.5 and widely triangular and bifid to less than 0.3 and rounded, respectively. Because these three characters are referable to plants identifiable as C. puccioana, Lophocolea diverstipa is included in its synonymy. Stephani (1922, p. 270) cannot be used for aid in typification, because he states the leaves are to 0.25-divided, a character not present in either C. humilis or C. puccioana. There is a Skottsberg collected specimen in G (No. 14135) labeled "Lophocolea diversistipa, Patagonia australis," that is actually Acrobolbus ochrophyllus. It is possible that Stephani utilized this plant in a portion of his description of Lophocolea diversistipa, because he states for the stem, "cuticla papillata," for the leaves "apice ••• FIG. 51. Clasmatocolea puccioana (De Not.) Grolle. 163 164 FIELDIANA: BOTANY ad 1/4 inciciso-biloba sinu recto lobis late triangulatis" and "cuticula dense minuteque papillata," all of which are characters of A. ochrophyllus. [Characters of the underleaves are seemingly those of L. diversistipa = C. puccioana, although they vary from those used in Stephani (1911).] It is possible that Stephani distributed that portion of the original material of Lophocolea diversistipa utilized for the protologue of this species in Stephani (1911), retaining only material of A. ochrophyllus (which may have been an admixture) and then using this plant for the description in Species Hepaticarum. 2. Clasmatocolea puccioana, like several other members of the genus, has a vestigial stem perigynium on which several series of bracts and bracteoles are inserted. The phenomenon in C. puccioana seems to be closely related to the occurrence of fertilization. 3. Reports of C. puccioana from the Falkland Is. are based upon mis- determinations of C. fulvella. Reports of L. puccioana in Stephani (1900, 1901) and Herzog (1939, 1954) are based upon misdeterminations of C. humilis. The record of L. puccioana in Arnell (1955a), plus many of Santesson's collections labeled L. puccioana are actually C. gayana. Ecology. — In forests where on fallen trees and in layers of vegetation over logs. The species also occurs on the sides and apices of rather exposed bryophyte mounds, where often mixed or associated with Blepharido- phyllum densifolium, Acromastigum sp., and Plagiochila sp. Also in moor- land areas where on cliffs or over soil. It is rather common in Sphagnum moors. Phytogeography. — Tierra del Fuego (I. de los Estados and wet, cold, southern Chilean islands); Patagonian Channels N. to 48° 04' S.; also at 1,000 m. in Cord. Pelada (40° 07' S., El Mirador, Prov. Valdivia) (see fig. 51). Specimens seen.— ARGENTINA. TERR. TIERRA DEL FUEGO: I. de los Estados, Pto. Cook, Skottsberg s.n. (hb. Grolle). CHILE: Sin. he., Poeppigs.n. as Chiloscyphus notophyllus (BM). PROV. MAGALLANES: S. Darwin, Spegazzini s.n. as Chiloscyphus notophyllus (BM); I. Chair, Spegazzini 253 (VER); I. Burnt, Cabo Desolation, Spegazzini 261 (VER); I. Basket, Spegazzini s.n. as Lejoscyphus turgescens (NY); S. Melville (not I. Melville), M. Skyring, Spegazzini 29b (VER); I. Capitan Arecena ("I. Clarence"), Harlot 7, 27 (VER); B. Tuesday, Naumann s.n. (FH); Bruns- wick Pen. (cf. Engel, 1978); E. side of B. Borja, Engel 6144B, 6163B - c. per. (MSC); E. side of Pto. Bueno, Engel 5589 - c. n>o<9° FIG. 59. Clasmatocolea cucullistipula (Steph.) Grolle. 1, Main axis, ventral-lateral view. 2, Median leaf cells. 3, Stem, cross section. 4, 5, Leaves (flattened). 6-8, Teeth of leaf apex. 9, Underleaf, dorsal view. 10, Underleaf, ventral view. 11, Underleaf (nearly flattened). 12, Axis with leaves removed, lateral view. All from lectotype of L. cucul- listipula, Dusen 376, Chile, Prov. Chiloe, I. Guaitecas. 184 S 11-13 FIG. 60. Clasmatocolea cucullistipula (Steph.) Grolle. 1, Axis with perianth. 2, 3, Por- tions of perianth mouth. 4, 5, Perianth, cross section through portion just below apex. 6, Perianth, cross section through median portion. 7, Portion of perianth mouth. 8, Androecial axis (note width compared to sterile portions). 9, 10, Male bracts showing lobule armature. 11, Capsule wall, cross section. 12, Outer capsule wall cells. 13, Inner capsule wall cells showing portion with semiannular bands sparsely developed. 14, Seta, cross section. Figures 1-4, 6,7, from Engel 4496, Chile, Prov. Magallanes, I. Williams; figure 5, from Engel 4293, Chile, Prov. Aisen, Pto. Island; figures 8-10, from Engel 5543, Chile, Prov. Magallanes, Pto. Bueno; figures 11-14, from syntype of L. cucul- listipula, Dusen s.n., Chile, Prov. Chiloe, I. Guaitecas. 185 186 FIELDIANA: BOTANY Lophoc olea filiformis Steph. Bull. Herb. Boissier 6:661. 1906 (= Spec. Hep. 3:77), syn. nov. Holotype: Chile, Prov. Valdivia, Valdivia, Hahn,e\hb. Jack (G!). Lophocolea cucullifolia Steph. Spec. Hep. 6:269. 1922, syn. nov. Holo- type: Chile, Prov. Chiloe, I. Chiloe, sin. coll., ex hb. Jack (G! — c. per.). Plants prostrate or suberect, or subpendent, in loose or dense, usually pure tufts, pale olive green-light brown to pale brown, nitid when dry, capillaceus, obscurely to distinctly moniliform (especially in ventral-lateral view); axes 350-490 n wide, the leaves and underleaves fragile, easily tearing. Branches rather common, Frullania-0^ predominating, ventral-intercalary type occasional, lateral-intercalary type absent. Stems (5-)6 cells high, cortex in 1 row of very thick-walled cells the same size as or slightly smaller than the medullary cells, the exposed wall very thickened; medullary cell walls very thickened; endophytic hyphae in both cortex and medulla or absent. Rhizoids in very tight fascicles from a region that includes both the stem immediately at underleaf base and the median basal portion of an underleaf, the rhizoids often tearing off with underleaf. Leaves with insertion distinctly recurved at ventral end; leaves strongly erect (axis commonly appearing channeled in dorsal view), occasionally connivent, loosely im- bricate, conchiform concave, =*= oblate to broadly ovate; apex broadly rounded to =±= truncate, undivided, with 1 or a few small teeth; dorsal margin ± straight in proximal half, entire or with 1 or a few small teeth, short decurrent; ventral margin broadly rounded, entire or with I or a few small teeth, the basal portion subauriculate, extending across stem and nearly in contact with or occasionally overlapping ventral margin of opposite leaf. Leaf cells thin walled, trigones massive, coarse and nodular, often confluent, commonly with leaf cell lumen, except for the narrow thin-walled places, bounded mostly by the massive trigones; median leaf cells ll-18(-22) ^ wide, 13-24 M long; cuticle smooth or slightly roughened and appearing finely granular. Underleaves 2.4-5 X stem width, free, apical portion oppressed to stem or ventral margin of leaf, the underleaves approximate, cucullate, =•= clasping stem, the margins curved dorsally and very often extending above dorsal stem surface, auriculate at the base; apices undivided, broadly rounded, occasionally retuse, occasionally with 2 slime papillae or 1-celled teeth terminating in slime papillae; margins in basal portion with 1 lobe that extends above dorsal stem surface, the lobe terminating in a slime papilla, the margins in median portion with a slime papilla or a tooth terminating in a slime papilla. Plants dioecious; androecia intercalary on main axis or long Frullania-type branches, wider than sterile portions of axis, densely imbricate, leaving a narrow strip of exposed stem tissue; bracts with basal portion strongly saccate, distal portion strongly concave, with apices appressed to bract immediately above, the apices with a few teeth; lobule margin with 1-3 erect or involute laciniae or small lobes that may or may not terminate in a slime papilla; antheridia solitary, stalk uniseriate. Gynoecia on main axis or long Frullania-lype branches or ventral-intercalary sub- floral innovations, often copiously produced; subfloral innovations originating from ENGEL: CLASMATOCOLEA 187 below bracteoles of first or second series or absent; bracts in 3-4 series, those of inner- most series deeply concave to subnaviculariform to wide-elliptic to wide-ovate; apices broadly rounded, with several teeth and laciniae or lobes, the laciniae and lobes some- times dentate; lamina margins with a few to several laciniae and rounded or sharp lobes and a few teeth. Bracteoles of innermost series ca. equal to bract size, free from bracts, deeply convex (ventral view), ovate; apices rounded, with several small teeth and some- times laciniae; lamina margins with a few to several irregularly shaped laciniae, rounded or sharp lobes, a few teeth, and a few slime papillae, the armature occasionally rather copious. Perianth subterete-obscurely to moderately trigonous, slenderly to rather stoutly ovoid, rather strongly inflated in median and basal portions, the apex 4-5-plicate, the ventral side strongly convex throughout, to moderately infolded throughout, or some- times infolded in apical and median portions with the basal portion convex; perianth sides gradually or abruptly narrowing toward the rather contracted mouth; lobes very broadly rounded, copiously dentate to laciniate, the laciniae commonly narrow and dentate. Seta 5 cells in diameter, with 14 rows of outer cells surrounding an inner core of scattered cells that are ca. equal to outer row, the core cells with corners thickened similarly to small trigones. Capsule valves 0.59-0.77 mm. long, 38-46 /* thick, of (4-)5 layers, the outer row of cells equal to thickness of 2.2-2.8 of interior strata; outer layer with red-brown, nodule-like or wide spinelike thickenings that are often moderately tangentially dilated, semiannular bands very common; exposed wall moderately thickened; intermediate and inner layers subequal in thickness or inner layer slightly less thick, the intermediate layers with thickenings often considerably tangentially dilated, semiannular bands occasionally present; inner layer of cells with red-brown semiannular bands sporadic to rather common, the radial walls with nodulose thicken- ings very common. Spores 24-27(-30) n, light brown, exine with =*= hyaline, narrowly conical, rather dense projections with radiating bases and dilated tips, the tips with finger-like projec- tions that are erect, laterally extended or recurved; spores averaging 2.8 X elater diameter. Elaters 8-10 ^ wide, spiraled to tips, elater walls red-brown. Differentiation. — This well-defined species, which exhibits compara- tively little variability, should offer no confusion with any other member of the genus. If sterile plants of C. cucullistipula are at hand, the following ensemble of features will serve to identify it: (a) the cucullate, free, un- divided underleaves that are auriculate at the base and that have a con- spicuous lobe in the basal portion extending dorsally beyond the dorsal surface of the stem; (b) the 1-4-dentate leaf margins; (c) the distinctive branching pattern, with Frullania type in predominance, ventral-intercalary type occasional and lateral-intercalary type absent; (d) the small axis size ( 350-490 n wide); (e) the characteristic stems that are (5-)6 cells high and that have both the poorly differentiated cortex and the medulla of very thick-walled cells; (f) the large and bulging leaf cell trigones; (g) the dis- tinctive and characteristic habit of loosely to densely intertwined, nearly pure tufts loosely adhering or semipendent from very small branches or twigs (this habit alone will distinguish C. cucullistipula from all other American taxa); and (h) the light olive green-light brown color. If an- FIG. 61. Clasmatocolea cucullistipula (Steph.) Grolle. 188 ENGEL: CLASMATOCOLEA 189 droecia are present, the unique combination of the androecia being wider than sterile portions of an axis and the frequently 2-3-laciniate to lobate bracts will further serve to distinguish the taxa. Perianths will provide the following additional ensemble of distinctive features: (a) the slenderly to rather stoutly ovoid shape; (b) the obscurely to moderately trigonous basal and median portions; (c) the 4-5-plicate apical portion with the contracted mouth; and (d) the copiously dentate and laciniate lobes. Further, the spores are 24-27(-30) M and larger than those of any other member of the genus. The spore wall is also unique. The above combination of characters, taken collectively, support an isolated position of the species within the genus. Indeed, with the number of unique features one could, with certain justification, argue for removal of the species from Clasmatocolea and for creation of a new, monotypic genus for it. However, I believe its natural affinities are with Clasmatocolea, where I choose to retain it. The species has some relationship with Sect. Strongylophyllae — on the one hand there is the branching pattern, stem anatomy, relatively small axis size, and large to bulging trigones of C. strongylophylla, whereas on the other hand, the cucullate underleaves and dentate leaves remind one of C. tjiwideiensis. However, I believe the species is most closely allied with Sect. Clasmatocolea. Note. — The syntypes of Lophocolea cucullistipula from the Rio Aisen Valley are misdeterminations of Xenocephalozia navicularis. Of the re- mainder of syntypes, all from Islas Guaitecas, Dusen 376 (S) best fits the original description and is here designated the lectotype. Ecology. — Corticolous; mostly on small branches and larger twigs of shrubs such as Pernettya or Berberis illicifolia or of trees such as Notho- fagus, Saxegothaea, or Fitzroya. When on twigs and small branches, the species nearly always occurs in pure tufts, because this is one of the few bryophytes of this region that has taken advantage of this kind of niche. When on tree trunks, however, it not uncommonly is intermixed with other hepatics such as Frullania spp., Clasmatocolea gayana, Porella subsquar- rosa, Lepicolea ochroleuca, Metzgeria sp., and Plagiochila sp. Less com- monly on bark of tree trunks. Ranging in altitude from near sea level in Prov. Magallanes to 1,000 m. (Cordillera Pelada) in Prov. Valdivia. Phytogeography. — Patagonian Channels (not S. of ca. 51° S. in Prov. Magallanes) and in Valdivian region N. to 39° 16' S. (Prov. Cautin) (see fig. 61). Specimens seen.— CHILE. PROV. MAGALLANES: Head of Pto. Bueno, Engel 5543 - c. per. + & (MSC); I. Chatham, N. shore of B. 190 FIELDIANA: BOTANY Wide, Engel 5368C - c. per. (MSC); E. side of I. Juan, Engel 5286 - c. per. (MSC); Pto. Alert, Engel 5033 - c.