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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:
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-
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ENGEL: CLASMATOCOLEA
33
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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
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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
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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. <?, 479A - c. <?, 811, 825 - c. <? as C.
koeppensis (JE). SOUTH GEORGIA: Koppenberg, Will 37 as C. koep-
pensis (M). FALKLAND IS. (selected). EAST FALKLANDS: Mt. Us-
borne Region, ridge between Mt. Usbornes 1 & 2, 685 m., Engel 2538B -
c. rf (MSC). WEST FALKLANDS: Mt. Adam, E. side of summit ridge,
670-700 m., Engel 3021 - c. <? (MSC). ARGENTINA. TERR. TIERRA
DEL FUEGO: Lapataia, Hatcher s.n. - c. <? (FH); W. side of B.
Lapataia, Crow 1507 - c. per., 7509, 1518 - c. per. (MSC); N. side of
L. Roca, Crow 1485 - c. <? (MSC); Ushuaia, Dusen 287 as Lophocolea
latissima (S); ibid., Dusen 303 as Lophocolea irregularis (G — c. per.,
NY); ibid., Baliza, near sea level, Roivainen 1206, 1206b — c. per. (F, H);
Ushuaia, R. Olivia, near sea level, Roivainen 1066 (F, H); ibid., Roivainen
50 FIELDIANA: BOTANY
1080b, 1082, 1089b (H); C. Garibaldi, 300 m., Roivainen 747b - c.
per. (H); I. Observatorio, Hassel de Menendez 3919 (BA, F). CHILE.
PROV. MAGALLANES: R. Azopardo, 300 m., Halle & Skottsberg 219
as syntype ofLophocolea monoica (UPS); Brunswick Pen. (cf. Engel, 1978).
PROV. AISEN: R. Aisen, Dusen s.n. as Lophocolea lenta (BM); Puerto
Aisen, Schwabe 45 /b p.p. as Lophocolea subaromatica (JE); Valle Frias,
Skottsberg 209 as Lophocolea humifusa (UPS). PROV. VALDIVIA: V.
Shoshuenco, Ruthsatz s.n. — c. tf (JE); SW slope of V. Quetrupillan,
1,450 m., Engel 11156 (F). PROV. CAUTIN: L. Quilleihue, 10.7 km. by
road E. of Puesco, ca. 7 km. by road W. of Chile-Argentina boundary,
1,050 m., Engel 11313 (F); Termas de Palguin, along R. Palguin, 730 m.,
Engel 11222 (F); Parque Nac. Villarrica, N. slope of V. Villarrica, 1,150 m.,
Engel 11180, 11190 (F); Pucon, "Halbinsel," Hosseus 363 A as syntype of
L. olens — c. per. + tf (JE); V. Llaima, Ruthsatz s.n. (JE); Parque Nac.
Conguillo, 1,050-1,120 m., Mahu 10686 - c. per., 70695, 10723, 10732,
10794, 10822 (hb. Mahu). PROV. MALLECO/PROV. ARAUCO: Cord.
Nahuelbuta, Parque Nac. Nahuelbuta, 44 km. W. of Angol, 1,300 m.,
Engel 12630 (F). PROV. SANTIAGO: C. Manquehue, Q. Agua del Palo,
940 m., Mahu 10592 - c. sporo. (F). ARGENTINA. PROV. CHUBUT:
A. Carbon, Halle 219 as syntype of Lophocolea monoica (UPS).
Subg. LACERIFOLIA Steph. ex Engel, subg. nov.
(Subg. Lacerifoliae Steph. Bull. Herb. Boissier 6 (7): 538. 1906 [= Spec.
Hep. 3:52], nom. nud.)
Folia caulina 2-3-lobis, apice ad 0.75 diviso; amphigastria caulina 0. 12-0.5-bifida;
oppositae <? bracteae connatae versus basin; praecursor coelocaulis praesens; peri-
anthii lobae bifidae.
Type species: Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle.
Leaves 2-3-lobed, divided to 0.75; underleaves 0. 1 2-0. 5-bifid; opposing c?
bracts fused toward base; coelocaule precursor present; perianth lobes
bifid.
Distribution. — Southern South America and Falkland Is.
Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. Figures 10-13,
Plates 3, 4.
For synonymy see under varieties.
Plants soft, prostrate, mixed with other bryophytes, occasionally in pure mats, grey-
green to pale brown to brown, sometimes a rich orange-brown with age; axes to 1.5
mm. wide.
Branches sparing to frequent, of lateral-intercalary and Frullania types (collections
varying in predominance of each); ventral-intercalary type not seen.
Stems 9-18 cells high, cortex in l-2(-3) rows of slightly to moderately thick-walled
ENGEL: CLASMATOCOLEA
51
FIG. 10. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. 1, Portion of main axis,
lateral view. 2, Main axis, ventral view showing underleaf and leaf insertion. 3a-c,
Underleaves. 4, Median leaf cells. 5, Underleaf segment. 6, Stem, cross section. Figures
1, 3a-b, 4-6, from lectotype of C. obvoluta, Davis, Chile, Prov. Magallanes, Cta. San
Martin; figure 2, from Engel 4386A, Chile, Prov. Aisen, F. Tempano; figure 3c, from
Engel 4381, Chile, Prov. Aisen, F. Tempano.
cells slightly to distinctly smaller than or ca. equal to size of medullary cells; medullary
cell walls thin; endophytic hyphae absent. Rhizoids in fascicles from stem near under-
leaf base.
Leaves with insertion distinctly recurved at ventral end; leaves strongly erect, with
dorsal lobes often connivent dor sally, leaves loosely to densely imbricate, conch iform
concave, obscurely to distinctly reniform, occasionally very wide-ovate; leaves 0.15-0.50-
bifid, dorsal lobe variable in size, medium- to wide-triangular, usually apiculate, plane or
with margins reflexed lending lobe slightly to distinctly canaliculate, rarely with margins
connivent toward base and then lobe here locally tubiform, the lobe entire or with 1-2
FIG. 11. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. 1-5, Leaves, illustrating
trend in apical and marginal characters. Figure 1, from lectotype ofLophocoleacookiana,
Spegazzini 14, Argentina, Terr. Tierra del Fuego, Pto. Cook; figure 2, from Engel
5581 A, Chile, Prov. Magallanes, Pto. Bueno; figure 3, from Engel 4381, Chile, Prov.
Aisen, F. Tempano; figures 4, 5, from lectotype of C. obvoluta, Davis, Chile, Prov.
Magallanes, Cta. San Martin.
52
$5-9
FIG. 12. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. 1, Longitudinal section
through sporophyte-bearing plant, showing axial proliferation of tissue; note position
of bracts and unfertilized archegonia. 2, Seta, cross section. 3, Perianth, ventral lobe
in middle. 4, Opposing antheridial bracts with stem(s), cross section. 5, Antheridial
stalk. 6, 7, Outer capsule wall cells. 8, Inner capsule wall cells. 9, Capsule wall, cross
section. Figures 1, 8, from Engel 4574 A, Chile, Prov. Magallanes, Pto. Eden; figures
2, 3, from Engel 5339, Chile, Prov. Magallanes, B. Wide; figure 4, from Engel 4277,
Chile, Prov. Aisen, Pto. Island; figure 5, from Engel 5890, Chile, Prov. Magallanes,
Pto. Charruca; figures 6, 7, 9, from Engel 6034, Chile, Prov. Magallanes, Pto. Gallant.
53
'•':.
54
ENGEL: CLASMATOCOLEA 55
pairs of commonly opposite teeth, the dorsal-basal portion of dorsal lobe or dorsal
margin immediately proximal to lobe entire or with a tooth or small canaliculate lobe;
ventral lobe variable in size, very wide-triangular, occasionally medium-triangular,
apiculate, plane, entire or with a single conspicuous tooth near base of ventral margin,
the lobe occasionally short and poorly developed; sinus in antical 0.25-0.45 of leaf , the
tissue at sinus base plane or reflexed, often toothed; dorsal margin of leaf straight,
plane or reflexed in distal portion, plane in proximal portion, not to long decurrent,
entire or with 1-2 straight or reflexed teeth or laciniae in distal portion; ventral margin
broadly rounded to subauriculate, with basal portion enlarged, extending vent rally beyond
stem and closely approaching or in contact with moderately spreading underleaves,
occasionally in contact with ventral margin of opposite leaf, the ventral margin entire
or with a few small teeth.
Leaf cells thin walled, trigones medium to large and bulging to occasionally coarse
and nodular; median leaf cells 19-42 p wide, (22-)24-50(-54) ^ long; cuticle smooth or
slightly roughened and appearing finely granular. Oil-bodies throughout leaf, occupying
small fraction of cell lumen, hyaline and glistening to tannish to "dirty white," of
grape-cluster type, with numerous, very small, barely protruding globules, the oil-body
surface appearing papillose; oil-bodies variable: subglobose to ovoid to elliptic to
fusiform to crescentic in shape, 2-3(-4) of leaf middle, 2-4 of leaf apex, 2-5(-8) of leaf
base, those of leaf middle 4-5 M wide, (5-)6-12 n long; oil-bodies throughout underleaf,
of about the same size as those of leaf middle.
Underleaves 3-4.3 X stem width, connate on both sides, rarely connate on 1 side in
axes with very loosely imbricate leaves, moderately spreading, approximate to loosely
imbricate, moderately convex (ventral view), moderately to very wide-ovate to suborbic-
ular (and slightly wider than long) to subcordate; underleaves bifid to 0.12-0.23, the
segments terminating in a small slime papilla, the segment margins entire; margins of
the lamina with 1 slime papilla or l-2(-3) small teeth terminating in a small slime
papilla, margins occasionally entire.
Plants dioecious; androecia terminal or intercalary on main axis, or short to long
lateral-intercalary or Frullania-type branches; bracts with basal portion strongly
saccate, the saccate portion fused toward base with opposite bract; bracts armed at apex
and upper portion of dorsal margin with 2-3 sharp teeth, or bracts armed with 2 small,
sharp lobes and a tooth on ventral margin near ventral lobe, the ventral margin other-
wise entire; lobule margin with an inrolled rounded lobe, the lobe entire or with a few
small teeth and/or slime papillae, occasionally with several low, rounded projections;
antheridia solitary, stalk uniseriate throughout or uniseriate and with scattered locally
biseriate areas.
Gynoecia on main axis or rather short to long lateral-intercalary or Frullania-type
branches; subfloral innovations absent or from below bracts of first, second, or inner-
most series, or bracteole of second series, the innovation may, in turn, produce a
perianth; coelocaule precursor present; bracts and bracteoles in 3 series, inserted on
coelocaule precursor; bracts of innermost series moderately convex to moderately con-
cave, elliptic to ovate, asymmetric and with dorsal margin more rounded than the
ventral, or symmetric; apex often rather narrowed, very short-bifid or divided to 0.25;
Opposite:
FIG. 13. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. • = var. obvoluta;
O = var. cookiana; O = both var. obvoluta and var. cookiana.
56 FIELDIANA: BOTANY
lamina margins with a few small to large teeth. Bracteoles of innermost series subequal
to bracts in size, free from bracts, concave (ventral view), ovate to elliptic; apex rather
narrow, very short bifid; lamina margins slightly to distinctly reflexed, entire or with a
few small to large teeth. Perianth strongly trigonous, narrowly elliptic to narrowly
ovate to short- or very long-subrectangular, with any 1, 2, or all sides infolded near
mouth, the perianth expanding slightly or moderately narrowing toward mouth, the
mouth bilaterally compressed due to infolded lobes; lobes broadly rounded, usually
obscurely to distinctly bifid, the segments acuminate, the mouth otherwise with several
teeth, mouth occasionally with several narrowly or broadly rounded small accessory
lobes.
Seta 7-12 cells in diameter, with 19-32 rows of outer cells with exposed walls moder-
ately to very thick-walled surrounding an inner core of scattered cells that are slightly
smaller than, or equal to, or slightly larger than outer row, the core cells with corners
thickened similarly to small to medium trigones. Capsule elliptic, the valves 0.74-1.82
mm. long, 30-72 /* thick, of (3-)4-6 layers, the outer row of cells thicker than all of
combined other strata or equal to a thickness of 2-3 X of interior strata; outer layer
with red-brown, nodule-like to spinelike thickenings that are often very weakly tangen-
tially dilated, with semiannular bands on some valves; exposed wall thickened; inter-
mediate and inner layers of cells subequal in thickness, the intermediate layers with
thickenings often distinctly extending onto tangential walls; inner layer of cells with
red-brown semiannular bands very common on some valves, sporadic on others, the
bands sometimes branched, often incomplete, the radial walls with nodulose thicken-
ings very common on some valves, moderately represented on others.
Spores (10-)11-12(-13) /*, light brown, exine with the light microscope appearing
minutely punctate, but under the SEM with a network of granulate to (more often)
short, closely packed, vermiform projections densely covered with nanogranules ;
spores averaging 1.1 X elater diameter. Elaters (8-)9-ll p wide, apical portions usually
with thick nonspiral walls, red-brown.
KEY TO VARIETIES OF Clasmatocolea obvoluta
1. Dorsal leaf lobe consistently canaliculate and/or ventral leaf lobe consistently with
a single conspicuous tooth on ventral margin; oil-bodies hyaline and glistening,
consistently 2 per cell in leaf middle and apex as well as in underleaf. .var. obvoluta
1. Dorsal leaf lobe plane or occasionally with margins slightly reflexed, the ventral leaf
lobe entire or occasionally with a single tooth on ventral margin; oil-bodies tannish
to "dirty white," 2-3(-4) per cell in leaf middle, 2-4 in leaf apex, (2-)3-4(-5) in under-
leaf var. cookiana
Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle, var. obvoluta.
Jungermannia obvoluta Hook. f. & Tayl. London J. Dot. / 30. 1845.
Lophocolea obvoluta (Hook. f. & Tayl.) Evans, Bull. Torrey Bot. Club
25:421. 1898. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle,
Rev. Bryol. Lichenol. 29:72. 1960. Lectotype (fide Engel, 1976):
Chile, Prov. Magallanes, I. Hermite, Cta. San Martin, Davis s.n.
(FHI).
Jungermannia obvolutaeformis De Not. Mem. Reale Accad. Sci. Torino
II. 16:220. pi. 8,f. 1-4. 1855, syn. fide Stephani (1906). Lophocolea
ENGEL: CLASMATOCOLEA 57
obvolutaeformis (De Not.) Mass. Nuovo Giorn. Bot. Ital. I. 17:223.
1885. Original material: Chile, "Prov. Valparaiso, Valparaiso,"
Puccio (non vidi; GE = O, NY = O, PAD = O, PC = O, RO = O,
TO = O).
Dorsal leaf lobe consistently canaliculate and/or ventral leaf lobe consistently with a
single conspicuous tooth on ventral margin; leaves usually 0.25-0.50-divided.
Differentiation. — Clasmalocolea obvoluta var. obvoluta is rather closely
related to C. trachyopa. The relationship is especially evident when typical
C. obvoluta is well developed and has strongly canaliculate dorsal lobes.
The following characters of C. obvoluta var. obvoluta will serve to imme-
diately distinguish it: (a) stems 9-18 cells high; (b) dorsal lobes medium-
to wide-triangular; (c) ventral lobes medium- to very wide-triangular, with
a single tooth on the ventral margin, but otherwise entire; and (d) under-
leaves nearly always connate on both sides and with apices 0.12-0.23-
divided.
Notes. — 1. Clasmatocolea obvoluta, like C. trachyopa, has a coelocaule
precursor or swollen region on which the perianth and three series of
bracts and bracteoles are inserted (fig. 15-1); see under C. trachyopa (p.
67) for further comments. In C. obvoluta, this phenomenon seems more
closely related to the occurrence of fertilization than in C. trachyopa.
2. In some collections, the outer capsule walls have semiannular bands
predominating but with nodulose thickenings rather common, whereas in
other collections the walls have nodulose thickenings predominating and
with semiannular bands very poorly developed.
Ecology. — Basically similar to that of var. cookiana, but seemingly with
a somewhat wider ecological amplitude. In forests, like the other variety,
it occurs on the floor and as a component of bryophyte mounds; var.
obvoluta, however, also grows epiphytic on shrubs (Pernettya base,
Berberis illicifolia, etc.) and tree branches. Particularly common as a
component of vegetation over the small cliffs that are common in southern
South American forests.
Phytogeography. — Tierra del Fuego and Patagonian Channels N. to
43° 57' S. (I. Guaitecas). Hassel (1977) records the species for South
Georgia; I have been unable to confirm or refute this report (see fig. 13).
Specimens seen.— ARGENTINA. TERR. TIERRA DEL FUEGO: I.
de los Estados, Spegazzini 21, 98, 99 as L. obvolutaeformis (VER); ibid.,
Spegazzini 21, 99 as L. obvolutaeformis (NY); ibid., B. Blossom, Spegaz-
zini 24 as L. obvolutaeformis (VER); I. de los Estados, B. Crossley, Hassel
de Mentndez 3880 (BA, F); I. de los Estados, M. Richardson, Spegazzini
159 as L. obvolutaeformis (VER). CHILE. PROV. MAGALLANES: Cabo
58 FIELDIANA: BOTANY
de Hornos, sin. coll. as L. fulvella (MICH); ibid., Hooker s.n. -• c. per.
(NY); Cta. San Martin, Hooker 19d (NY); ibid., Davis s.n. as syntype of
J. obvoluta (S); ibid., I. Hermite, Hahn s.n. — c. per. as L. obvolutaeformis
(PC, VER); I. Hermite, Hooker s.n. (NY); B. Tekenika, Skoltsberg s.n.
(S); I. Navarino, Angostura Murray, Spegazzini 244 as L. obvolutaeformis
(VER); I. Basket, Spegazzini 3 (NY); ibid., S. Darwin, Spegazzini 176*
as L. obvolutaeformis (VER); R. Azopardo, Halle 221 (S); I. Capitan
Arecena ("I. Clarence"), Hariot s.n. as L. obvolutaeformis (VER); I.
Clarence, S. shore of B. Pond, Engel 6264 A (MSC); I. Desolacion, Pto.
Angosto, Dusen 161 asL. lacerata(NY, S); I. Desolacion, Pto. Churruca,
head of Br. Lobo, Engel 5811 - c. c? (MSC); I. Desolacion, Pto. Chur-
ruca, N. side of Fo. Nassau, Engel 5890 - c. <?, 5924 (MSC); I. Desola-
cion, B. Tuesday, Engel 5648 — c. sporo., 57 13 A (MSC); B. Tuesday,
Naumann s.n. as L. obvolutaeformis (FH); Brunswick Pen. (cf. Engel,
1978); E. side of B. Borja, Engel 61 30, 6144C, 6153, 6185 (MSC); B. Borja,
Voyage of Albatross s.n. (FH); ibid., Voyage of Albatross s.n. as L. ob-
volutaeformis (NY); Can. Gajardo, Vo. Inga, Halle & Skottsberg 221 (S);
S. Skyring, Esto. Excelsior, Halle & Skottsberg 221 — c. per. (S); Can.
Smyth, 5m. coll. as L. trachyopa (PC); Pto. Mayne, Cunningham 169 (NY);
E. side of Pto. Bueno, Engel 5581 A (MSC); F. Peel, N. shore of Cta.
Amalia, Engel 5388 A (MSC); I. Chatham, N. shore of B. Wide, Engel
5296 A - c. per., 5322, 5339 - c. sporo., 5363 D - c. per. (MSC); Pto.
Alert, Engel 4923C (MSC); ibid., head of fiord W. of M. Markham,
Engel 4973 - c. <? (MSC). PROV. AISEN: N. side of F. Tempano,
Engel 4381, 4386 A, 4388C (MSC); Pto. Island, Cunningham 36 (BM);
ibid., Engel 4277 - c. cf (MSC); Pen. de Tres Montes, Pto. Barrosa [= Ot-
way Harbor], Challenger Exped. s.n. (BM, NY); Pto. Puyuhuapi, R.
Pascua, Schwabe 37b (JE); C. Tesoro, 800-900 m., Schwabe 41 1 a (hb.
Grolle, JE) PROV. CHILOE: I. Guaitecas, Dusen s.n. as L. trachyopa
(S); ibid., Dusen 363 as L. trachyopa (JE).
C lasmatocolea obvoluta var. cookiana (Mass.) Engel.
Lophocolea cookiana Mass. Nuovo Giorn. Bot. Ital. I. 17:224. pi. 16,
f. 11. 1885. Clasmatocolea cookiana (Mass.) Engel, J. Hattori Bot.
Lab. 36:156. 1973. Clasmatocolea obvoluta var. cookiana (Mass.)
Engel, Phytologia 41:311. 1979. Lectotype (fide Engel, 1973a):
Argentina, Terr. Tierra del Fuego, I. de los Estados, Pto. Cook,
February 1882, Spegazzini 14 (VER! - c. <?).
Lophocolea latissima Steph. Bih. Kongl. Svenska Vetensk.-Akad. Handl.
26 (III, 6): 42. 1900, syn.fide Engel (1973a). Original material: Chile,
Prov. Magallanes, S. Molyneux, 1 June 1896, Dusen 68 (FH!, G!).
ENGEL: CLASMATOCOLEA 59
Dorsal leaf lobe plane or occasionally with margins slightly reflexed, ventral leaf
lobe entire or occasionally with single tooth on ventral margin; leaves usually 0.15-0.25-
divided.
Differentiation. — The leaves of C. obvoluta var. cookiana bear some
resemblance to those of C. rigens, particularly when the latter are robust
and considerably wider than long. There should, however, be no confusion
between the taxa.
Note. — Stephani in his original description of Lophocolea latissima
states in reference to the leaf: "postice haud longe libera ut in Loph.
cookiana sed breviter inserta." The syntype of Lophocolea latissima and
lectotype of Clasmatocolea cookiana both have underleaves connate on
both sides with ventral leaf margins.
Ecology. — Rather common in mossy forests, where on the floor (often
mixed with Leptoscyphus horizontalis, Schistochila lamellata, Blepharido-
phyllum clandestinum, Megaceros endivaefolius, etc.) or on the sides or
apices of bryophyte mounds, often mixed or associated with Blepharido-
phyllum densifolium, Leptoscyphus aequatus, and Adelanthus lindenber-
gianus, etc. It also occurs on stream banks in moorland areas and over
soil near waterfalls. It occasionally occurs mixed with Sphagnum in
Sphagnum bogs.
Phytogeography. — Falkland Is.; Tierra del Fuego (including I. de los
Estados and western portion); Patagonian Channels N. to 42° 39' S.;
mainland West Patagonia N. to 40° 45' S. (see fig. 13).
Specimens seen.— FALKLAND IS. (selected). EAST FALKLANDS:
Port Stanley, Skottsberg 350 (S, UPS); Mt. Usborne Region, SE slope of
Mt. Usborne 2, ca. 455 m., Engel 2612C (MSC). ARGENTINA. TERR.
TIERRA DEL FUEGO: I. de los Estados, B. Crossley, Hassel de Menen-
dez 3882 (BA, F); I. de los Estados, Pto. Cook, Spegazzini 13 (G, VER);
ibid., Spegazzini 14 as L. obvoluta (NY). CHILE. PROV. MACAL-
LAN ES: I. Hornos, Harlot 155 (VER); I. Desolacion, Dusen s.n. (FH);
ibid., Dusen s.n. as L. obvoluta (FH); ibid., Pto. Angosto, Dusen s.n. (S);
ibid., Dusen 265 (NY); ibid., Dusen 401 as L. latissima (G); I. Desolacion,
Pto. Churruca, head of Br. Lobo, Engel 4877 B (MSC); I. Desolacion, B.
Tuesday, Engel 5695, 5714, 5742C (MSC); Brunswick Pen. (cf. Engel,
1978 sub C. cookiana); B. Isthmus, Savatier 205 (VER); E. side of Pto.
Bueno, Engel 5574 (MSC); Pto. Bueno, Cunningham 143 as L. obvoluta
(BM); head of F. Peel, Engel 5492 (MSC); E. side of I. Juan, Engel 5275C,
5282B (MSC); near E. shore of I. Pilot, Engel 4756 (MSC); W. side of
I. Grant, Engel 4711 (MSC); head of inlet, Pto. Charrua, Engel 4866 A
(MSC); Pto. Alert, Engel 4939C - c. per. (MSC); Pto. Eden, Engel
4532B - c. sporo., 4574 A - c. sporo. (MSC). PROV. AISEN: N. side
60 FIELDIANA: BOTANY
of F. Tempano, Engel 4391 A (MSC); Pto. Island, Engel 4291 - c. tf ,
4296 (MSC); Istmo de Ofqui, Pta. Leopardos, Hicken 62 as L. obvoluta
(JE); R. Aisen, Dusen 248 (NY). PROV. CHILOE: I. Guaitecas, Dusen
s.n. as L. trachyopa (FH, NY — cf , S — c. per.); ibid., Dusen 12, 363, 371
as £. trachyopa (S); I. de San Pedro, Dusen s.n. as L. obvoluta (FH); ibid.,
Dusen 576 as L. obvoluta (S); I. Chiloe, across Esto. Yaldad from village
of Yaldad, Cocauque area, sea level, Engel 11983 (F); I. Chiloe, E. of
Cucao near SW end of L. Cucao, Loncomilla (Alto de la Virgen), 75 m.,
Engel 12095, 12148B (F). PROV. OSORNO: Near small lake 10.3 km. by
road below Refugio Antillanca, 650 m., Engel 11553 (F).
Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle. Figures 14-17,
Plates 5-6.
Jungermannia trachyopa Hook. f. & Tayl. London J. Bot. 3:471. 1844.
Lophocolea trachyopa (Hook. f. & Tayl.) G. L. & N. Syn. Hep. 699.
1847. Clasmatocolea trachyopa (Hook. f. & Tayl.) Grolle, Rev.
Bryol. Lichenol. 29:73. 1960. Lectotype (nov.): Chile, Prov. Magal-
lanes, I. Hermite, Hooker (FH ! — c. sporo. + <? ).
Lophocolea arenaria Schiffn. in Naumann, Forschungsr. Gazelle 4
(4):13./>/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. + <? (MSC);
I. Desolacion, B. Tuesday, head of inner harbor, Engel 5716 (MSC);
Brunswick Pen. (cf. Engel, 1978); E. side of B. Borja, Engel 6174 - c.
sporo. (MSC); S. Skyring, Esto. Excelsior, Halle & Skottsberg 235 — c.
per. (S); NE corner of I. Tarlton, Engel 5106 — c. sporo. (MSC); S. side
of I. Madre de Dios, N. of I. Tarlton, Engel 5203 A - c. sporo. (MSC);
S. side of I. Madre de Dios, head of fiord E. of M. Roberto, Engel 5150,
5 151 A (MSC); Pto. Charnia, Engel 4805 - c. sporo. + cf, 4814 - c. tf,
4831 - c. sporo., 4836, 4850 - c. sporo., 4860 (MSC). PROV. AISEN:
Pen. de Taitao, Cabo Tres Montes, Pto. Barroso, Roivainen 1644b (H);
L. Risopatron, Schwabe 6/d(hb. Grolle). PROV. CHILOE: Pto. Ballena,
Engel 4167B - c. sporo., 4232 - c. sporo. (MSC); I. Chiloe, Cord. San
Pedro, near aserradero at San Pedro, R. Puidi, 320 m., Engel 11902 (F);
I. Chiloe, Cord. San Pedro, Butalcura, near R. Butalcura, 1 1 km. by road
from Ruta 5, 100 m., Engel 11812 (F). PROV. OSORNO: Around L.
Toro on road to Refugio Antillanca, 855-885 m., Engel 3969, 3979B,
ENGEL: CLASMATOCOLEA 69
3985 C - c. per. + <?, 4005, 4013, 4020 A - c. sporo., 4042C - c. per.
(MSC); below L. Toro on road to Refugio Antillanca, near R. Nauto,
825 m., Engel 4134 (MSC); 12.1 km. by road below Refugio Antillanca,
550 m., Engel 11534 (F); Agua Caliente, along R. Chanleufu, 4 km. from
Termas de Puyehue along road to Refugio Antillanca, 400 m., Engel
11468 — c. o", 11477 - c. per. (F); Anticura, near Salto del Indio, 19 km.
by road E. of Termas de Puyehue, 300 m., Engel 11669 — c. per. (F);
Huitrapulli, Aleucapi, 700-820 m., Schwabe s.n. — c. per. (hb. Schuster);
ibid., Schwabe 28 - c. <? (JE). PROV. VALDIVIA: W. slope of Cord.
Pelada, W. of El Mirador, between La Union and Pta. Hueicolla, 580-
680 m., Engel 12255, 14324 (F); E. slope of Cord. Pelada, E. of El Mirador,
between La Union and Pta. Hueicolla, 740-840 m., Engel 12466 — c. per.,
14432 (F); Chaihuin, Pto. Colun, ca. 500 m., Schwabe 6/g (hb. Grolle);
ibid., Schwabe 26, 26h (JE).
Subg. CLASMATOCOLEA
Leaves undivided, entire or variously armed ; underleaves of main axis and branches
conspicuous, not reduced, free or connate on 1 side; androecia narrower than sterile
portion of axis, the bract lobules entire, dentate or with 1 lacinium or lobe; bracteoles
free from bracts; perianth without scales or plicae, the lobes broadly rounded, undivided.
Distribution. — Pan-temperate, occurring in all three sectors of Southern
Hemisphere; in American sector north in Andes to Costa Rica, in African
sector north to Burundi, and in Australasian sector disjunct in Java.
Sect. PACHYCLASMATOCOLEA Engel, sect. nov.
Foliorum caulina laminae 2-4-stratosae in media-basis parte; foliorum et interdum
amphigastriorum margines conspicue incrassatae, 2-4-stratosae; antheridii pedicelli
4-seriati.
Type species: Clasmatocolea marginata (Steph.) Grolle.
Leaf lamina 2-4-stratose in median-basal area; leaf and sometimes underleaf margins
conspicuously swollen, the elevated border 2-4-stratose; antheridial stalks 4-seriate.
Distribution. — Endemic to Tasmania.
Clasmatocolea marginata (Steph.) Grolle. Figures 18-20.
Jungermannia marginata Mitt, in Hooker, Bot. Antarc. Voy. 3:222.
1859 non J. marginata Lehm. Nov. Minus Cogn. Stir. Pug. 5:11.
1833 (= Cololejeuned), nonJ. marginata Hook. f. & Tayl. London J.
Bot. 3:566. 1844 (= Raduld). Leioscyphus marginatus Steph. Bull.
Herb. Boissier 6 (3): 223. 1906 (= Spec. Hep. 3:23). Odontoschisma
marginalum (Steph.) Steph. Bull. Herb. Boissier 8 (8): 594. 1908
(= Spec. Hep. 3:378) = Clasmatocolea crassimarginata Grolle, Rev.
Bryol. Lichenol. 29:71. 1960. Clasmatocolea marginata (Steph.)
FIG. 18. Clasmatocolea marginata (Steph.) Grolle. 1, Main axis, ventral view. 2, Stem,
cross section. 3, 4, Portion of main axis, ventral view. 5, 6, Underleaf, cross section
through marginal area. 7, 8, Underleaf, longisection through median-basal area of
lamina. 9, Axis, dorsal view with Frullania-lype branch and associated half-leaf (HL).
10, Median leaf cells. 11, Underleaf segment, cross section. 12, Antheridial stalk (the
fourth row not in plane of view). Figures 1, 6, 7, 11, from Engel 14328, Tasmania, Mt.
Field Natl. Park, Mt. Mawson Plateau; figures 2-5, 9, 10, from lectotype; figure 12,
from Rodway, Mt. Wellington Plateau.
70
FIG. 19. Clasmatocolea marginata (Steph.) Grolle. 1-6, Leaves (1-3, not flattened,
4-6, flattened). 7, 8, Leaves, longisection through median-basal area. 9-16, Leaves,
longisection through apex. Figures 1-3, 7, 9-11, from Engel 14328, Tasmania, Mt.
Field Natl. Park, Mt. Mawson Plateau; figures 4-6, 8, 12-16, from lectotype.
71
72
FIELDIANA: BOTANY
FIG. 20. Clasmatocolea marginata (Steph.) Grolle. 1, Portion of axis, lateral view.
2-9, Underleaves. Figures 1-3, from lectotype; figures 4, 5, from Rodway, Mt. Wellington
Plateau; figures 6-9, from Engel 14328, Tasmania, Mt. Field Natl. Park, Mt. Mawson
Plateau.
Grolle, Nova Acta Leop. 25 (161): 73. 1962. Lectotype (nov.): Tas-
mania, Arthur's Lakes, 13/2/43, sin. coll. (Gunri) 1648 (NY!).
Plants very stiff, rigid, wiry, fleshy, erect, in pure tufts; axes to 12 cm. tall, to 3 mm.
wide, brown to brownish black, highly nitid.
Branches rather frequent, of Frullania- lateral-intercalary and ventral-intercalary
types, the intercalary branches a bit more common ; main axis and branches occasionally
becoming flagelliform, the flagelliform portions with leaves scaly and distant or with
leaves caducous.
Stems 14-15 cells high, cortex in 1-2 rows of smaller cells with thicker walls, the
epidermal row with the outer tangential and often also the radial walls brownish;
medullary cell walls thick; endophytic hyphae not seen. Rhizoids only sporadically
developed, in fascicles at underleaf bases.
Leaves rigid, fleshy, brittle, the margins appearing conspicuously swollen, the distal
(2-)4-6(-7) cell rows 2-4-stratose and forming a slightly to distinctly elevated border, the
margin swollen its entire length, but less conspicuously so toward the leaf bases; leaf
margin in cross section with the multistratose portion forming a knotlike swelling
sharply defined from the leaf lamina or the multistratose portion gradually merging
with the lamina, the marginal cells not appreciably differing in size from neighboring
cells, but with thicker walls; leaf lamina 2-4-stratose in median-basal area, the lamina
otherwise unistratose, except for occasional local bistratose areas; leaf insertion slightly
recurved at ventral end; leaves erect to spreading, contiguous to imbricate, moderately
to rather deeply adaxially concave, ovate to suborbicular, the apex broadly rounded to
subtruncate, undivided, the apex and margins entire; dorsal margin slightly to distinctly
rounded, plane, short to long decurrent on well-developed axes; ventral margin dis-
tinctly rounded, rather sharply so toward the base, plane.
ENGEL: CLASMATOCOLEA 73
Leaf cells with walls slightly thickened, trigones small; median leaf cells 24-38 n wide,
36-53(-60) n long: cuticle roughened.
Underleaves stiff, rigid, 0.65-1.5 X stem width, the margins (both lamina and segment)
unistratose or multistratose and then appearing slightly swollen, the marginal (l-)2-4 cell
rows bistratose, forming an inflated, sometimes knotlike, thick-walled group of cells;
underleaf lamina bistratose toward the base, the lamina otherwise unistratose, except for
occasional local bistratose areas; underleaves free, but in close proximity to leaf on
1 side, slightly spreading, distant to imbricate, plane or slightly convex, the segments
often curved toward stem (sometimes sharply so), the underleaves elliptic to subrectan-
gular to narrowly ovate; apices bifid 0.15-0.45; segments frequently of different size and
configuration, narrowly or broadly triangular, the tips acute or rounded, sometimes
acuminate, the segment margins entire; margins of the lamina entire or with one tooth.
Plants dioecious; androecia terminal or intercalary on leading axes; bracts nearly
obliterating view of dorsal surface of stem, with basal portion saccate, the distal portion
strongly concave, with apices appressed to bract immediately above; lobe margins with
an elevated border similar to leaves; lobule margin incurved, with a small tooth or
rounded projection, or entire; antheridia l(-2) per bract, stalk short, of 4 cell rows.
Gynoecia and sporophyte not seen.
Variation. — The lectotype represents a smaller facies of the species, with
less than optimal expression of characters. Among the notable differences,
robust plants have (a) a lesser tendency for axes to become flagelliform;
(b) leaves short decurrent, more often erect, more closely imbricate and
more deeply concave; (c) leaf borders more pronounced and composed
of a greater number of cells; (d) leaf bases composed of a greater number
of strata; (e) underleaves contiguous to imbricate and, on the whole, less
shallowly divided; and (f) underleaf borders more swollen and conspicuous.
Differentiation. — This species is at once distinct from all other members
of the genus in possessing swollen, elevated leaf and (often) underleaf
margins, which in section are 2-4-stratose. In fact, the only other leafy
hepatic with multicellular leaf margins known to me is Wettsteinia den-
sir etis (cf. Grolle, 1965). The species is also unique within Clasmatocolea
in (a) the median-basal polystraty of leaf and underleaf lamina; and (b)
the 4-seriate antheridial stalks. The collection of these features necessitates
a somewhat isolated position within Subg. Clasmatocolea; the closest
relative of the species appears to be C. humilis of Sect. Clasmatocolea.
Ecology-Phytogeography. — Rare endemic to Tasmania, where sub-
merged or semisubmerged over rock in lakes or cold, rather rapid moving
water of creeks at 950-1,300 m.
Specimens seen. — TASMANIA: Mt. Wellington Plateau, Rodway s.n.
(HO); Wellington Falls, Boston 349 (MEL); Brown's R., Oldfield (MEL);
Mt. Field Natl. Park, Mt. Mawson Plateau, W. of L. Dobson and S. of
L. Seal, 1,250 m., Engel 14328 (F); Ben Lomond Natl. Park, L. Youl, A.
Ratkowsky 79/4 (F); Cradle Mt. Region, Shadow L., ca. 950 m., Norris
27792 (F).
74 FIELDIANA: BOTANY
Sect. CLASMATOCOLEA
Intercalary branches basically similar to main axis, the branches not vermiform or
submoniliform (exc. C. moniliformis), usually of same width as main axis, often very
long; leaves and underleaves unistratose throughout, the margins not swollen or
elevated; leaf cuticle smooth or slightly roughened; antheridial stalks 1-2-seriate.
Distribution. — Pan-temperate; in American sector north in Andes to
Costa Rica, in African sector north to Burundi.
Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. Figures 21-24. Plates
7-10.
For synonymy see under varieties.
Plants fleshy and often soft to rather stiff" and wiry, prostrate to suberect, mixed with
other bryophytes, or in dense intertwined often pure mats, grey-green to olive-green to
light to deep brown, rarely light green, the stems commonly red-brown; axes to 7.3 cm.
tall, to 1.8 mm. wide.
Branches common, occasionally in irregular fascicles, commonly of lateral- and ventral-
intercalary type, the lateral-intercalary more prevalent; lateral-intercalary branches from
ventral portion of leaf axils or directly basiscopic to stem leaves; Frullania-0'/^ branches
occasional but more common in hygrophilous forms; branches occasionally with first
leaf or first pair of leaves retuse to short-bifid, especially on intercalary branches;
stolons very exceptional [only a single ventral-intercalary stolon seen (Engel 5304 A)].
Stems 8-14 cells high, the cortex in l-2(-3) rows of moderately to distinctly thick-
walled cells smaller than (or very rarely equal in size to) medullary cells; exposed wall of
outer cortical row moderately to very thickened; medullary cell walls thin to slightly
thickened; endophytic hyphae absent. Rhizoids in fascicles from stem near underleaf
base.
Leaves with insertion very slightly to distinctly recurved at ventral end; leaves
strongly erect, dorsally connivent, often lending axis a somewhat vermiform appearance,
the leaves occasionally moderately spreading, loosely to densely imbricate, conchiform
concave, orbicular to oblate to subreniform; apex broadly rounded to truncate, very
rarely broadly retuse, undivided, often moderately to distinctly inflexed, sometimes ca.
upper half of leaf moderately inflexed; apices and margins entire, or with a few sporadic
teeth, or regularly dentate; dorsal margin ± straight in proximal half, plane to sinuous,
rarely reflexed, not to rather long decurrent; ventral margin plane or moderately in-
flexed, broadly rounded, the basal portion often enlarged to =*= auriculate, often extend-
ing ventrally beyond stem and in contact with slightly spreading underleaf, occasionally
in contact with ventral margin of opposite leaf.
Leaf cells thin-walled, trigones large to bulging to knotlike, occasionally confluent,
occasionally minute to medium in hygrophilous facies; median leaf cells ll-26(-30) n
wide; 12-32(-38) /* long; cuticle smooth or slightly roughened. Oil-bodies throughout
leaf, occupying small fraction of cell lumen, hyaline to very light grey, of grape-cluster
type, with numerous, small, slightly protruding globules, the oil-body surface appearing
coarsely papillose; oil-bodies subglobose to ovoid to elliptic to fusiform in shape,
consistently 2 per cell of leaf middle and apex, and 2-5 of leaf base, those of leaf middle
3-4 n wide, 5-8(-10) /* long; oil-bodies throughout underleaf, 2(-3) per cell, of same
size as those of leaf middle.
ENGEL: CLASMATOCOLEA
75
FIG. 21. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. 1, Main axis, dorsal view,
2, Main axis, ventral view. 3, Stem, cross section. 4, Median leaf cells. 5, Underleaf
segment, apical portion with slime papilla. Figures 1-3, from lectotype of C. humilis,
Hooker, Kerguelen; figures 4, 5, from syntype of C. humilis, Hooker, Kerguelen (S).
Underleaves l-4.3(-4.8) X stem width, free or connate on 1 side, transversely or
obliquely inserted, appressed to stem or slightly to strongly spreading (and then at
nearly right angles to stem); underleaves distant to imbricate, very slightly convex to
deeply convex (ventral view) and then with margins curved dorsally and often strongly
incurved, the underleaves occasionally ± cupulate, the basal portion often abruptly
convex; underleaves orbicular to oblate to ovate to elliptic to subrectangular, with
basal portion occasionally subauriculate; apices undivided and broadly rounded to
truncate, or bidentate to broadly retuse to bifid to 0.35, the apices very rarely only with
2 slime papillae; segments often of different size and configuration, occasionally apicu-
late, occasionally terminating in a slime papilla, the segment margins entire; lamina
76
ENGEL: CLASMATOCOLEA 77
margins with 1 slime papilla or tooth or lobe terminating in a slime papilla, the margins
occasionally entire.
Plants dioecious; androecia terminal or intercalary on main axis or short to rather
long lateral- or ventral-intercalary branches; bracts with basal portion strongly saccate
and distal portion slightly to strongly concave, with bract apices appressed to bract
immediately above or moderately spreading; lobule margin involute, with armature
variable: with 1 -several slime papillae and often few-celled teeth, or with 1 lacinium or
large, broadly rounded, involute lobe ; antheridia solitary, the stalk uniseriate throughout
or with 1-2 very local biseriate areas.
Gynoecia on main axis or on short to long lateral- or ventral-intercalary branches;
subfloral innovations absent, but occasionally with a branch below bract or bracteole
of first series; bracts in 3 series, those of innermost series plane to slightly concave,
ovate to obovate to wide-elliptic, plane or undulate; apices undivided, broadly rounded
to truncate, entire or sparingly dentate to laciniate, occasionally short-bifid, and then
with lobes occasionally apiculate; lamina margins occasionally recurved, entire or with
a few teeth. Bracteoles of innermost series free from bracts, concave to canaliculate
(ventral view), often appressed to the infolded ventral lobe of perianth, rarely reflexed, the
bracteoles orbicular to ovate to elliptic to obovate to subrectangular; apices undivided
and truncate or bifid to 0.25; lamina margins entire or with a few teeth or 1 lobe.
Perianth strongly trigonous, subclavate to subcampanulate to elongate-subrectangular, the
ventral side usually strongly infolded and lending perianth =*= bilaterally compressed,
the perianth sides straight or gradually expanding toward the ± bilaterally compressed
wide mouth; lobes commonly undulate, broadly rounded, often flaring and then broadly
rounded to auricula te just distal to keel, the lobes entire, occasionally repand, occasion-
ally regularly dentate to ciliate to laciniate; keels often with wings of a few to several
cells high, the wings occasionally sparingly dentate.
Seta 8-19 cells in diameter, with 30-70 rows of outer cells with walls moderate to very
thick, with occasional intermediate thickenings, the inner corners with thickenings
similar to large to bulging trigones; inner core of cells scattered, ca. equal to or slightly
larger than outer row, thin-walled, with thickenings similar to small to large trigones.
Capsule broadly ovate to elliptic, the valves 0.84-1.5 mm. long, 36-60/1 thick, of 4-6
Opposite:
FIG. 22. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. 1, Main axis, lateral view
of C. humilis var. suspecta; note leaf dentition. 2, Leaves. 3, Underleaves, showing
major different shapes. 4, Perianth mouth, ventral lobe in middle. 5, Main axis, dorsal
view of hygrophilous form; leaves removed to show leaf insertion and underleaf
form. 6, Main axis, ventral-lateral view of hygrophilous form. Figure 1, from Engel
5010, Chile, Prov. Magallanes, Pto. Alert; figure 2a, from Dusen 360, Chile, Prov.
Chiloe, I. Guaitecas; figures 2b, 3a, b, from syntype of C. humilis, Hooker, Kerguelen
(S); figure 3c, from Dusen 66, Chile, Prov. Valdivia, Corral; figure 3d, from Dusen 38,
Chile, Prov. Magallanes, Pto. Bueno; figures 3c, d, from Engel 5489, Chile, Prov.
Magallanes, F. Peel; figure 3e, from Dusen, Chile, Prov. Aisen, R. Aisen Valley;
figure 3f, from Dusen 360, Chile, Prov. Chiloe, I. Guaitecas; figure 3g, from Dusen 250,
Chile, Prov. Magallanes, Pto. Angosto; figure 3h, from type of L. magellanica, Naumann,
Chile, Prov. Magallanes, B. Tuesday; figure 3i, from Halle 222, Argentina, Terr.
Tierra del Fuego, L. Fagnano; figure 4, from Engel 5483C, Chile, Prov. Magallanes, F.
Peel; figures 5, 6, from Engel 577 IB, Chile, Prov. Magallanes, B. Tuesday.
FIG. 23. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. 1, Outer capsule wall
cells. 2, Inner capsule wall cells. 3, Capsule wall, cross section. 4, Antheridial stalk.
5, Sporophyte bearing axis. 6, 7, Perianth mouths, ventral lobes in middle. 8, Seta,
cross section. Figures 1, 2, 8, from Engel 6299, Chile, Prov. Magallanes, B. Pond;
figure 3, from Engel 4042B, Chile, Prov. Osorno, near L. Toro; figure 4, from Engel
4728, Chile, Prov. Magallanes, I. Grant; figure 5, from Engel 5472, Chile, Prov. Magal-
lanes, F. Peel; figure 6, from Engel 2331, Chile. Prov. Magallanes, Pto. Cutter; figure
7, from Engel 3319, Falkland Is., Weddell I., Waterfall Valley.
78
ENGEL: CLASMATOCOLEA
79
FIG. 24. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle (includes all varieties).
layers; outer row of cells 18-23 ^ thick, equal to thickness of 2.5-3 of interior strata;
outer layer with vinaceous or red-brown, nodule-like to =*= spinelike thickenings that
are often feebly tangentially dilated, senriannular bands absent; exposed wall of outer
layer thickened; intermediate and inner layers of cells subequal in thickness, the inter-
mediate layers with thickenings often feebly extending onto tangential walls; inner
layer of cells 4-6 n thick, with red-brown semiannular bands, the bands often incom-
plete, the radial walls with nodulose thickenings often present.
Spores 10-15 /u, light brown, exine with the light microscope appearing minutely
punctate, but under the SEM with densely granulate to short, wide vermiform projec-
tions covered with crowded, sometimes coalescing nanogranules, the intervening
hollows with nanogranules; spores averaging 1 .2 X elater diameter. Elaters 9-12 p
wide, apical portions often with thick nonspiral walls; elater walls light brown, 2 p in
diameter.
KEY TO VARIETIES OF Clasmatocolea humilis
1. Underleaves polymorphic in shape on any given axis, the apices usually undivided,
80 FIELDIANA: BOTANY
sporadically short-bifid; underleaves uniformly small for plant size, those in apical
portion (0.8-)1.0-1.3 X stem width. Endemic to Tristan da Cunha. .var. polymorpha
1. Underleaves =*= consistent in shape on any given axis, the apices undivided or (fre-
quently) bifid; underleaves not uniformly small for plant size, 1.0-4.3 X stem
width 2
2. Axes with leaves entire; perianth lobes entire or occasionally sparingly dentate
var. humilis
2. Axes with at least some of leaves sparingly to regularly dentate; perianth lobes
regularly dentate to ciliate to laciniate var. suspecta
Clasmatocolea humilis (Hook. f. & Tayl.) Grolle, var. humilis.
Jungermannia humilis Hook. f. & Tayl. London J. Bot. 3:468. 1844 non
J. humilis Kash. & Chopra, Liverworts of the Western Himalaya and
the Panjab Plains 2:88. pi. 25, f. 5-7. 1932 (= ^.Jungermannia sp.).
Mylia humilis (Hook. f. & Tayl.) Trev. Mem. 1st. Lomb. Sci. Lett.
III. 4:412. 1877. Solenostoma humilis (Hook. f. & Tayl.) Mitt.
Philos. Trans. 168 (extra vol.): 42. 1879. Nardia humilis (Hook. f. &
Tayl.) Berggr. New Zealand Hep. 7. 1898. Lophocolea humilis (Hook,
f. & Tayl.) Steph. Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26
(III, 6):40. 1900. Leioscyphus humilis (Hook. f. & Tayl.) Pears. Bull.
Misc. Inform. 1922:249. 1922. Clasmatocolea humilis (Hook. f. &
Tayl.) Grolle, Rev. Bryol. Lichenol. 29:72. 1960. Lectotype (nov.):
Kerguelen, Hooker s.n. (FH !).
Jungermannia palustris Hook. f. & Tayl. London J. Bot. 3:464. 1844,
syn.fide Engel (1978). Lophocolea palustris (Hook. f. & Tayl.) Besch.
& Mass. Mission Sci. Cap Horn 5:222. 1889. Original material:
Chile, Prov. Magallanes, I. Hermite, Hooker 19b (FH!, NY!, S!).
Lophocolea magellanica Schiffn. in Naumann, Forschungsr. Gazelle 4
(4):14,/?/. 4,f. 22, 23. 1890, syn.fide Engel (1978). Holotype: Chile,
Prov. Magallanes, I. Desolation, B. Tuesday, 2 February 1876,
Naumann s.n. (FH ! — c. per.).
Lophocolea haslatistipa Steph. K. Svenska Vetenskapsakad. Handl.
46 (9):45. /. 19 h, i. 1911, syn.fide Engel (1978). Lectotype (fide
Engel, 1978): Falkland Is., King George B., 22 November 1907,
Halle & Skottsberg 352 (UPS!).
Lophocolea incrassata Steph. K. Svenska Vetenskapsakad. Handl. 46
(9):46./. 17 c, d. 1911, syn.fide Engel (1978). Lectotype (fide Engel,
1978): Falkland Is., 1908, Skottsberg 644 (G!).
Lophocolea integerrima Steph. K. Svenska Vetenskapsakad. Handl. 46
(9):46./. 17 a, b. 1911, syn.fide Engel (1978). Lectotype (fide Engel,
1978): Chile, Prov. Magallanes, S. Skyring, F. de los Ventisqueros,
26 April 1908, Halle & Skottsberg 211 (UPS! - c. per.).
ENGEL: CLASMATOCOLEA 81
Lophocolea rotundifolia Steph. K. Svenska Vetenskapsakad. Handl.
46 (9): 52. / 19 r, s. 1911, syn. fide Engel (1978). Lectotype (fide
Engel, 1978): Chile, Prov. Magallanes, I. Atalaya, 25 May 1908,
Skottsberg 230 (UPS!).
Lophocolea catenulala Herz. Darwiniana 11:215. / 2. 1957, syn. nov.
Holotype: Argentina, Rio Negro, Parque Nac. Nahuel Huapi, M.
Tronador, 1500 m., 6 January 1937, Donat 182 (JE!).
Axes with leaves entire. Perianth lobes entire or occasionally sparingly dentate.
Variation. — This species exhibits considerable variablity, and I am
recognizing the following ecological variants :
Hygrophilous forms. — Plants lax, occasionally quite flaccid in texture;
leaves erect, closely imbricate; plants frequently growing in or near
water.
In extreme hygrophilous forms, the plants are comparatively large,
delicate, and often quite flaccid in texture. The underleaves are loosely
to closely inbricate, rather deeply convex (ventral view) and with
margins curved dorsally to strongly incurved ; the apices are undivided
to retuse. Examples of this extreme condition are Engel 2198, 5373,
5748, 5769, 577 IB, 5869, 6052 A, and 6447 (see fig. 22), as well as syntype
plants of Jungermannia palustris.
In the syntypes of Jungermannia palustris, which are quite flaccid in
texture, the ventral portion of the leaf is deeply adaxially concave with
the ventral margin incurved, whereas the dorsal portion of the leaf is
concave, often with the dorsal margin decurved. This condition pro-
duces, in the leaf middle, the "remarkable longitudinal obtuse fold,"
noted in Hooker & Taylor (1844). There is a trend in variation toward
this condition commencing in the more xeric forms of C. humilis with
the ventral margin only slightly incurved and the dorsal portion of the
leaf slightly concave. The trend then continues in aquatic forms,
especially in the very flaccid plants, to the condition found in the
syntypes of J. palustris. Jungermannia paluslris is a good example of the
hygrophilous facies of C. humilis.
Xerophilous forms. — Plants very rigid, stiff, never flaccid in texture;
leaves spreading, more loosely imbricate; plants growing on rocks and
in comparatively dry, exposed habitats.
The underleaves of C. humilis exhibit considerable variation. They
may be plane to very slightly convex (ventral view) as in the type plants
of Jungermannia humilis, to ± cucullate as in type plants of Lophocolea
magellanica. They are orbicular to oblate to ovate to elliptic to subrec-
tangular in shape. In undivided underleaves, the apex may be very wide
82 FIELDIANA: BOTANY
truncate or the underleaf may taper gradually to a narrowly rounded
apex; in the latter instance the underleaf margins occasionally are
incurved (dorsal view). Further, apices vary from unlobed (and broadly
rounded or truncate) to retuse to bifid to varying degrees, but not to
greater than 0.35. The unlobed apices may be entire or bidentate. It is
not unusual to observe considerable variation in underleaf apices on a
single axis. Underleaf margins are entire or with 1 slime papilla or with
one tooth or lobe in the middle third, but may be dentate or small to
large lobate in either upper, middle, or lower third of the underleaf
margin. Some of this diversity in underleaf shape and armature is shown
in Figure 22-3.
Differentiation. — Clasmatocolea humilis is related to C. fasciculata; see
under the latter for distinguishing characters.
Clasmatocolea humilis should be separated with care from those pheno-
types of Lophocolea austrigena with leaves concave (but never conchiform
concave) and underleaves occasionally plane. The following ensemble of
characters of L. austrigena will distinguish these plants: (a) underleaves
universally connate on one side; (b) underleaf apices entire to bidentate
to retuse; (c) underleaves occasionally strongly concave (recurved); and
(d) leaves sporadically weakly to rather strongly convex. Character (c)
sometimes is particularly evident toward axis apices.
In Australasia, care should be taken with certain facies of Lophocolea
gunniana that have sporadically convex (ventral view) underleaves and
adaxially concave leaves. Such plants are presumably an aquatic phase,
and confusion with C. humilis is very possible. In this case characters (a),
(c), and (d) listed above for L. austrigena also apply to L. gunniana and
may also be used to distinguish the latter from C. humilis. See also the
comments under C. notophylla.
I am including a discussion of Lophocolea otiphylla here because this
species, when of certain facies, will key to Sect. Clasmatocolea, and within
this group, it exhibits some relationship to C. humilis.
Lophocolea otiphylla has leaves that vary from distinctly convex to
plane to very slightly concave (with the slight concavity restricted to the
ventral-basal subauriculate portion of the leaf, which extends ventrally
beyond the stem) to distinctly concave. These differences are largely
dependent upon the orientation of the apex and ventral margin and not
upon any natural tendency for concavity in the ventral-basal portion of
the leaf lamina. Rather, the degree of concavity is correlated with the
degree the apex and ventral margin are inflexed. On the other hand, the
apex and both dorsal and ventral margins are occasionally all reflexed,
and in this situation, the leaves are distinctly convex, lending a definite
ENGEL: CLASMATOCOLEA 83
Lophocoleoid appearance to the plants. When the leaves are convex, the
plants, in general aspect, look quite like L. austrigena or L. elata, but
certainly not like any Clasmatocolea.
This species, as mentioned above, shows some relationships to C. humilis
and bears a rather striking resemblance to hygrophilous forms of that
species, sharing such features as size, texture, and color of plants as well
as form and size of underleaves. Examples of the hygrophilous form of
C. humilis are syntype plants of Jungermannia palustris (= C. humilis). It is
of interest that Hooker & Taylor (1844, p. 466) alluded to this connection
when stating that their Jungermannia otiphylla "approaches our Junger-
mannia palustris from the same country and from similar situations."
Two features of L. otiphylla necessitate, in my mind, placement in
Lophocolea, rather than Clasmatocolea. First, the leaves are frequently
plane or distinctly convex, and then totally unlike any Clasmatocolea
taxon, the leaves never have the distinct conchiform concavity so typical
of many of the species of Clasmatocolea. Second, the androecial bracts of
L. otiphylla are not strongly saccate at the base, but rather consist of a
small, inflated lobule of minimal proportion to overall bract size and a
lobe not much different from leaves. The bracts are thus quite leaflike
and hardly spicate. In my experience with the genus Lophocolea, the male
bracts tend to be more leaflike and nonspicate, very much like that of
L. otiphylla. In Clasmatocolea, male bracts are either strongly saccate
throughout or distinctly saccate in the basal portion, with the distal
portion strongly concave. Further, the lobule is quite large in proportion
to bract size, and the lobe is clearly differentiated from leaves.
Notes. — 1. There has been some confusion surrounding the names
Solenostoma humilis and Nardia humilis. Hooker & Taylor (1844) described
Jungermannia humilis for a Kerguelen plant with well-developed under-
leaves, and the plate in Taylor & Hooker (1847) clearly indicates the
presence of such structures. Mitten (1879) however, transferred the species
to Solenostoma and stated (p. 42), "Both S. humilis and J. inundata were
originally described as stipulate species, no amphigastria have, however,
been since found on the specimens. It is probable that the figure of the
supposed stipule of /. humilis may have been drawn from a fragment of
Leioscyphus turgescens." Berggren (1898), in his treatment of New Zealand
hepatics, believed his plants must be identical to those treated by Mitten
(he. cit.) but transferred the epithet to Nardia.
Stephani (1901) alleviated some of the confusion by stating he had seen
original material of Jungermannia humilis that indeed possessed con-
spicuous underleaves and belonged to the genus Lophocolea. Stephani
(loc. c//.), however, still used the name Solenostoma humilis for New
84 FIELDIANA: BOTANY
Zealand plants. Hodgson (1946) includes a discussion of this confusion.
All observed Berggren collections labeled Solenostoma humilis or Nardia
humilis from various localities in North and South Islands, New Zealand,
are Jungermannia inundata Hook. f. & Tayl. (15 specimens from LD and
1 1 from S).
2. Lophocolea magellanica is here regarded as a new synonym of Clas-
matocolea humilis. When considering the spectrum of underleaf variation
in C. humilis, the underleaves of type plants of L. magellanica exhibit an
extreme form. The underleaves are rather deeply convex (ventral view),
nearly orbicular in shape, and with apices truncate or shallowly and very
broadly lunate. The underleaf apices may or may not be bidentate. The
trigones of the type plants of L. magellanica are very large and bulging,
and the perianth mouth entire, both characters of which are found within
the variation of C, humilis.
3. The underleaves of Lophocolea hastatistipa are mostly bifid to 0.5 and
with a large lobe on one or both sides of the underleaf base. The under-
leaves, however, may be entire or 1 -dentate at the base and occasionally
are lobate on one side and entire on the other. Lophocolea hastatistipa fits
well within the variation circumscribed for C. humilis and is here reduced
to synonymy.
4. The underleaves of Lophocolea integerrima are occasionally undivided,
but more often bidentate with teeth of one to a few cells, or emarginate,
or at most retuse. If bidentate or entire, the underleaf apices are then ±
truncate. The underleaf margins are entire. Stephani (1911) states the
plants of L. integerrima are sterile. However, upon examination of plants
of syntype material, perianths were found to be present. Lophocolea
integerrima fits well within the variation pattern of C. humilis and is here
reduced to synonymy.
5. The syntypes of Lophocolea rotundifolia from "Azopardo-Tal" are
misdeterminations of Pachyglossa spegazziniana. Of the remaining syn-
types, the specimen from Isla Atalaya best fits the description and figures
in Stephani (1911) and is here designated the lectotype.
6. The Gasperi collections of Lophocolea magellanica cited in Gola
(1923) are actually specimens of Pachyglossa spegazziniana var. spegaz-
ziniana (fide specimens in FI !).
Ecology. — This variety has a somewhat wide ecological amplitude. In
forests on the floor, over rocks, cliffs, and rotted logs. Rather frequently
associated with streams — often on boulders, sometimes on logs over
streams. The variety also occurs in moorland areas where over rock
(mixed with Lophocolea sabuletorum) or in small pools.
ENGEL: CLASMATOCOLEA 85
The hygrophilous form seems to be restricted to cold, very high rainfall
areas. On Isla Desolation, in western Tierra del Fuego, it occurred in
pools of moorland areas on ridges and there commonly associated with
Isoetes sp. It also occurs submerged in small lakes in the Patagonian
Channel region.
Phytogeography. — Macquarie Is.; New Zealand, South I. (915 m.);
Antipodes; Australia, New South Wales (2,000-2,300 m.); Kerguelen;
Crozet Is.; Marion I. (15-130 m.); Prince Edward Is. (100 m.); Tristan da
Cunha Group, Gough I. (170-505 m.), Tristan da Cunha (170-650 m.);
Falkland Is. (sea level-595 m.); Tierra del Fuego (200-580 m.); Patagonian
Channels; Valdivian region (100-1,240 m.; N. to 36° 43' S.; and Andean
Patagonia in Parque Nac. Nahuel Haupi) (see fig. 24).
Specimens seen.— MACQUARIE IS.: Ashton 1112 as C. crassiretis (hb.
Grolle); ibid., Evans 6 (hb. Grolle). NEW ZEALAND. SOUTH ISLAND.
SOUTHLAND: Fiordland Natl. Park, head of Hollyford Valley, near S.
entrance to Homer Tunnel, Milford Rd., ca. 915 m., Schuster 55596a —
c. tf (hb. Schuster). NELSON: Nelson Lakes Natl. Park, Travers Saddle,
Simpson 34 12 p.p. as C. strongyhphylla (CHR). ANTIPODES: Hut Cove,
Godley s.n. as Lophocolea austrigena (CHR); ibid., Godley s.n. as C.
paucistipula (CHR). AUSTRALIA. NEW SOUTH WALES: Carruthers
P., 2,000-2,300 m., McVean 26449 - c. & , 26450 (hb. Grolle). KERGUE-
LEN: Sin. loc., Hooker s.n. as syntype of/, humilis (L, LD, S — c. cT);
Royal Sound, Eaton s.n. as Leioscyphus turgescens (BM, NY); ibid.,
Eaton s.n. (hb. Grolle); I. Longue, on margin of lake E. of le Kiosque (W.
of Port-Bizet), 125 m., Engel 8028 A (MSC). CROZET IS.: 1929, sin. coll.
(S); I. de 1'Est, Hebrard 39B2 (hb. Grolle); ibid., Ring & Raknes 69 (O).
MARION L: Sin. loc., Rand 3305 - c. per., 3394 (hb. Grolle); ibid.,
Huntley 171, 251k, 255, 942c (PRE); ibid., Hitntley 945e (hb. Grolle);
Tarn Valley, 130 m., Huntley 477 (F, hb. Grolle, PRE); Nellie Humps,
Bakker 207 jb (hb. Grolle); near weather station laboratory, 15 m.,
Bakker 236/b (hb. Grolle). PRINCE EDWARD IS.: Albatross Valley,
100 m., Bakker 619h (hb. Grolle, PRE). TRISTAN DA CUNHA: 600 m.,
Christopher sen & Mejland 984 - c. per. (O); S. of Round Hill, 650 m.,
Christophersen s.n. (UPS). GOUGH L: Upper Watersmeet, 170 m., Wace
539 as L. diversistipa (UPS); W. slopes of Gonydale, 505 m., Wace 598a
(UPS). FALKLAND IS. (selected). EAST FALKLANDS: Port Stanley,
sin. coll. (BM); ibid., Halle s.n. as Lophocolea otiphylla (UPS); ibid.,
Halle 222 as L. otiphylla (NY, S); ibid., Lechler s.n. as L. puccioana (S);
ibid., (Lechler) (FH, hb. Vana); ibid., Skottsberg s.n. as L. otiphylla
(UPS); ibid., Sapper Hill, Skottsberg s.n. as L. otiphylla (UPS); ibid.,
Skottsberg 217 as L. magellanica (NY); ibid., Skottsberg 222 as L.
86 FIELDIANA: BOTANY
otiphylla (NY, S); Port Harriet, Halle s.n. as L. otiphylla (UPS); ibid.,
Halle 222 as L. otiphylla (NY, S); headwaters of Mullet Cr., Engel 3161 B,
3167, 3182 (MSC); Mt. Usborne, Halle s.n., 230 as syntype of L. rotundi-
folia (UPS). WEST FALKLANDS: Hornby Mts., Skottsberg 222 as L.
otiphylla (S, UPS - c. cf ); Mt. Adam, basin E. of summit, 580-595 m.,
Engel 2996 A - c. per., 2998 A, 3000 (MSC); Weddell I., Halle & Skottsberg
226 as L. puccioana (S). ARGENTINA. TERR. TIERRA DEL FUEGO:
R. Olivia, Ushuaia, Skottsberg 231 as syntype of L. skottsbergii (LD, S);
I. de los Estados, Skottsberg s.n. as L. navistipula (G, S); ibid., Spegazzini
s.n. as Leioscyphus turgescens (NY); ibid., Pto. Cook, Skottsberg s.n., 45
as L. auslrigena (S); I. de los Estados, Pto. San Juan del Salvamiento,
Spegazzini 183 as Chiloscyohus notophyllus (NY); E. of M. Olivia, Sa. de
Sorondo, 580 m., Crow 1846, 1850 (MSC); S. side of Paso Garibaldi,
along Ruta Nac. No. 3, 400 m., Crow 1769 (MSC); Sa. Alvear, 380 m.,
Roivainen 854c (H); ibid., 370 m., Roivainen 874 (F, H). ARGENTINA/
CHILE: L. Fagnano, Halle & Skottsberg 205 as Lophocolea gottschaeoides
(S); ibid., Halle 222 as L. otiphylla (S, UPS). CHILE. PROV. MAGAL-
LANES: I. Saddle, sin. coll. 167 as L. palustris (PC); I. Hoste, sin. coll.
906 as L. otiphylla (PC); ibid., Hyades s.n. as L. palustris (FH); I. London-
derry, Exped. Aust. Belgica s.n. as L. palustris (FH); S. Skyring, Cabo
Leon, Hatcher 34-5 (F, UW-M); R. Fontaine, Halle & Skottsberg 4 as
L. otiphylla (S); ibid., Halle & Skottsberg 222 as L. otiphylla (NY); R.
Azopardo, Dusen s.n., 169 (S); ibid., Dusen 38 as Lophocolea triseriata
(UPS); ibid., Dusen 41, 42 as L. otiphylla (S); ibid., Dusen 56 (L, S, UPS);
ibid., Dusen 60 as L. puccioana (S); ibid., Dusen 117 as L. cookiana (NY);
I. Clarence, sin. coll. 64 as L. puccioana — c. per. -f <? (PC); ibid., "S.
Hoss," Harlot 64 as L. puccioana (PC); I. Clarence, B. Pond, Engel
6291 - c. per., 6293 - c. per., 6298, 6299 - c. sporo. (MSC); S. Otway,
Pto. Pomar, Halle & Skottsberg 225 as syntype of Lophocolea patulistipa
(UPS); I. Desolacion, Dusen s.n. as L. magellanica (FH); ibid., Dusen s.n.
(S); ibid., Dusen as L. puccioana (NY, S); ibid., Pto. Angosto, Dusen s.n.
(S); ibid., Dusen 169 (LD, NY, S); ibid., Dusen 172 as L. otiphylla - c.
per. (S); ibid., Dusen 176, 225, 249, 252, 371 (UPS); ibid., Dusen 250 (JE,
L, S, UPS); ibid., Dusen 268 as L. gayana (S); ibid., Dusen 373 as L.
puccioana (S); ibid., ca. 200 m., Dusen 180 (S); ibid., Dusen 384 as L.
puccioana (NY, S); Pto. Angosto, Dusen s.n. (LD); I. Desolacion, Pto.
Churruca, head of Br. Lobo, Engel 5822 B, 5869 (hygrophilous fo.) (MSC);
I. Desolacion, B. Tuesday, head of inner harbor, Engel 5691, 5708, 5748 —
c. c? (hygrophilous /<?.), 5769 (hygrophilous fo.), 577 IB (hygrophilous
fo.} (MSC); Brunswick Pen. (cf. Engel, 1978); E. side of B. Borja, Engel
6122 - c. sporo. (MSC); S. Skyring, F. de Los Ventisqueros, Halle &
Skottsberg 222 as L. otiphylla (NY, S, UPS); S. Skyring, B. Pinto, Halle &
ENGEL: CLASMATOCOLEA 87
Skottsberg 222 as L. otiphylla (UPS); I. Pacheco, Pto. Ramon, Skottsberg
144 as Jamesoniella oenops (UPS); Can. Smyth, Pto. Ramirez, Skottsberg
198 as syntype of L. diversistipa (LD, S, UPS); I. Newton, Dusen 56 (NY);
ibid., Dusen 117 as L. puccioana (NY); Pto. Bueno, Dusen 38 as L. otiphylla
(S); ibid., Dusen 42 as L. otiphylla (NY); ibid., Dusen 49 as L. puccioana
(FH, NY); ibid., Dusen 50 as L. puccioana (JE); ibid., Dusen 103 (NY);
head of F. Peel, Engel 5472 - c. sporo., 5489 (MSC); F. Peel, Skottsberg
222 as L. otiphylla (UPS); ibid., N. shore of Cta. Amalia, Engel 5384 -
c. sporo., 5423 — c. per. (MSC); Can. Messier, Cta. Rayo, Skottsberg 225
as syntype of L. patulistipa (S); ibid., Skottsberg 230 as syntype of L.
rotundifolia (UPS); E. side of I. Juan, Engel 5373 (hygrophilous fo.) (MSC);
S. side of I. Madre de Dios, head of fiord E. of M. Roberto, Engel 5 129 A,
5157 (MSC); Pto. Charrua, E. of Pta. Brown, Engel 4590 A, 4596 - c.
per. + c? (MSC); ibid., main cove S. of Pta. Brown, Engel 4633C,
4649 A - c. c?, 4651 (MSC); Pto. Alert, head of fiord W. of M. Markham,
Engel 4976 (MSC); B. Halt, Lechler s.n. (BM). PROV. AISEN: Pto.
Island, Engel 4338 A (MSC); B. Chacabuco, Halle 220 as L. navistipula
(BM, LD, S, UPS); ibid., Halle 226 as L. puccioana (S); R. Aisen, Dusen
s.n. (BM, JE, LD, S); ibid., Dusen 222 as L. puccioana (NY); ibid.,
Dusen 249 (FH); ibid., Dusen 250 - c. per. (NY); ibid., Dusen 949 - c.
per. (O); Pro. Puyuhuapi, Schwabe 43 as L. otiphylla (S); ibid., L. Risopat-
ron, sin. coll. 22 (JE); C. Tesoro, Schwabe 39a as L. rotundifolia (JE); ibid.,
960-1,000 m., Schwabe 39b, 39/cp.p., 40 p.p. as L. rotundifolia (JE). PROV.
CHILOE: I. Guaitecas, Dusen s.n. (hb. Grolle, JE, LD, S); ibid., Dusen
s.n. as L. puccioana (NY); ibid., Dusen 24 as L. puccioana (S); ibid.,
Dusen 360 (FH, NY, S); ibid., Bo. Chica, Dusen s.n. as L. puccioana (JE);
ibid., Dusen 375 as L. otiphylla — c. per. (NY); ibid., Dusen 384 as L.
puccioana (NY); I. Guaitecas, Melinca, Halle 210 (S); ibid., Dusen 352
(NY); Melinca, Dusen s.n., ?35a (S); I. Guaitecas, Pto. Low, Dusen s.n.,
336 (S); M. Corcovada, Halle & Skottsberg s.n. as syntype of L. rotundi-
folia (UPS); ibid., Skottsberg s.n. as syntype of L. rotundifolia (G); I.
Chiloe, Cord. San Pedro, near aserradero at San Pedro, R. Puidi, 320 m.,
Engel 11914 — c. per. (F); I. Chiloe, Aguas Buenas area, 4.7 km. E. along
Aguas Buenas road from Ancud-Quemchi road, ca. 100 m., Engel 12189 -
c. sporo. (F). PROV. LLANQUIHUE: L. Todos los Santos, Cayutue,
Wolffugel s.n. as L. rotundistipula (JE). PROV. OSORNO: near L. Toro
on road to Refugio Antillanca, 855 m., Engel 3961 B (MSC); Agua
Caliente, R. Chanleufu, 4 km. by road from Termas de Puyehue along
road to Refugio Antillanca, 400 m., Engel 11425, 11437 (F); Termas de
Puyehue, Schwabe 63/d p.p. as L. rotundifolia - c. <? (JE). PROV.
VALDIVIA: L. Puyehue, Mahu 7568 (JE, hb. Mahu); W. slope of Cord.
Pelada, W. of El Mirador, between La Union and Pta. Hueicolla, 680-
88 FIELDIANA: BOTANY
840 m., Engel 12333, 12341 (F); Los Ulmos, Mahu 7770 (hb. Grolle); R.
Futa, Schwabe 15 (JE); Corral, Dusen 66 (S). PROV. CAUTfN: Parque
Nac. Conguillo, 1,110 m., Mahu 10823 (hb. Mahu) PROV. MALLECO :
Parque Nac. Nahuelbuta, Mahu 6035 (JE, hb. Mahu). PROV. MALLECO/
PROV. ARAUCO: Cord. Nahuelbuta, Parque Nac. Nahuelbuta, 48 km.
W. of Angol, Aquas Calientes, 1,240 m. Engel 12610 (F). PROV. CON-
CEPCI6N: Talcahuano, Dusen 72 as L. otiphylla — c. per. (NY); ibid.,
Dusen 172 as L. otiphylla (S). ARGENTINA. PROV. Rfo NEGRO:
Puerto Blest, Dusen 504 (S). PROV. Rfo NEGRO/PROV. NEUQUEN:
L. Nahuel Huapi, Dusen s.n. as L. puccioana (FH). Patagonia: Sin. loc.,
Skottsberg s.n. as syntype of L. patulistipa (G); ibid., Skottsberg s.n. as
syntype of L. integer rima (G).
Clasmatocolea humilis var. suspecta (Mass.) Engel.
Lophocolea puccioana var. /3 suspecta Mass. Nuovo Giorn. Bot. Ital.
I. 17:228. 1885. Clasmatocolea humilis var. suspecta (Mass.) Engel,
Phytologia 41:309. 1979. Lectotype (nov.): Argentina, Terr. Tierra
del Fuego, I. de los Estados, M. Conegliano, March 1882, Spegazzini
770 (VER! - c. per. + <?).
Lophocolea multispinula Steph. Spec. Hep. 6:284. 1922. Original ma-
terial: Chile, Prov. Magallanes, I. Desolation, B. Tuesday, sin. coll.
(G!).
Leioscyphus patagonicus Pears. Bull. Misc. Inform. Kew 1922:250. 1 pi.
1922, syn. nov., non L. patagonicus Steph. Kongl. Svenska Veten-
skapsakad. Handl. 46 (9) :38. 1911 (= Leptoscyphus). Holotype:
Chile, Prov. Magallanes, S. Molyneux, 1 June 1896, Dusen s.n.
(BM! - c. cT; isotypes: BM! - c. &, hb. Grolle!).
Lophocolea hickenii Herz. Hedwigia 66:90. / 7. 1926, syn. nov. Holo-
type: Chile, Prov. Aisen, B. San Rafael, 26 February 1921, Hicken
68/a(JE\).
Axes with at least some of leaves sparingly to regularly dentate, the leaves with up
to 10 teeth. Perianth lobes regularly dentate to ciliate to laciniate.
Differentiation. — This variety differs from var. humilis by possessing
lea ves that are sparingly to regularly dentate. With regard to leaf armature,
at one end of the continuum are plants in which a few leaves on a given
axis have one or a few teeth, such as in syn- and lectotype plants of
Lophocolea puccioana var. /3 suspecta and original material of Leioscyphus
patagonicus. At the other extreme are plants in which the leaves are
regularly and often coarsely dentate, such as holotype plants of Lopho-
colea hickenii, Engel 4042B, 4050C, 12280, and 12406. There is a steady
ENGEL: CLASMATOCOLEA 89
continuum toward the development of this extreme and, interestingly,
those phenotypes with dense, coarse teeth tend to be copiously branched
and are very commonly corticolous. I have spent considerable time in
assessing whether the copiously branched, coarsely toothed facies might
better be treated as a discrete, if narrowly circumscribed, separate species.
However, the "cut-off point" in the definition of such a species would be
purely arbitrary, and I prefer to treat this complex as a rather plastic
one — both morphologically and ecologically.
Confusion of this variety with C. fulvella is possible when C. hwnilis
var. suspecta occasionally produces rather copious, short lateral-inter-
calary branches that are subvermiform or submoniliform and with branch
underleaves more closely imbricate than in main axes. In such cases, C.
humilis var. suspecta may be distinguished from C. fulvella by (a) main
axis underleaves at most moderately convex (ventral view), with margins
neither curved dorsally nor ± clasping the stem; and (b) intercalary
branch underleaves usually short-bifid with segments conspicuously evi-
dent. Clasmatocolea fulvella, on the other hand, has (a) main axis under-
leaves cupulate, with margins curved dorsally and ± clasping the stem;
and (b) intercalary branch underleaves usually undivided, distinctly cu-
pulate and appearing inflated. This facies of C. humilis var. suspecta
represents one kind of example of those phenotypes with coarse, dense
teeth (see discussion above).
Note. — Stephani (1922) states for the leaves of L. multispimda, "ceterum
ubique regulariter spinuosa," but was probably referring to leaves of the
smaller branches, which are more regularly and consistently dentate than
the leaves of the main axis or larger branches. The leaves of the latter are
entire or sparingly to irregularly dentate. The type plants are rather small.
Ecology. — This variety seems to prefer well-shaded niches. In the
Magellanian forests it occurs on Berberis illicifolia, etc., and in rocky
stream valleys. In the Valdivian region it occurs in forests from near sea
level to 840 m., and there on moist decorticated logs or on Nothofagus
bark. This variety, compared to var. humilis, is more often corticolous.
Phytogeography. — Marion I.; Tristan da Cunha; South Georgia; Pata-
gonian Channels; Valdivian region (N. to 39° 52' S. in Prov. Valdivia;
Andean Patagonia in Parque Nac. Nahuel Huapi); and on Mas a Tierra,
Juan Fernandez Is. (see fig. 24).
Specimens seen.— MARION I.: Huntley 727 (F, JE, PRE); Huntley
890/a (F, PRE); Huntley 937 a (PRE). TRISTAN DA CUNHA: 400 m.,
Christopher sen & Mejland 96 (O). SOUTH GEORGIA: Bay of Isles,
Rosita Hr., Skottsberg 222 (hb. Grolle). ARGENTINA. TERR. TIERRA
90 FIELDIANA: BOTANY
DEL FUEGO: I. de los Estados, "New Year Harbour," Spegazzini 84
as syntype of L. puccioana var. 13. suspecta — c. per. -f- tf (VER); ibid.,
M. Conegliano, Pto. Cook, Spegazzini s.n. as L. puccioana (NY). CHILE.
PROV. MACALLAN ES: I. Hermite, Harlot 129 as L. puccioana var.
suspecta (VER); Ens. Villarino, Hatcher s.n. as L. puccioana var. suspecta
(FH, VER); I. Capitan Arecena ("I. Clarence"), Pto. Hope, S. Magda-
lena, Spegazzini 234 as L. puccioana var. p. suspecta (VER); I. Desolacion,
Dusen 263 as L.fulvella (S); ibid., Pto. Angosto, Dusen 169 (UPS); ibid.,
Dusen 250 (L, LD); ibid., Dusen 268 as L. gayana - c. per. (NY); I.
Desolacion, Pto. Churruca, head of Br. Lobo, Engel 5810 (MSC); I.
Desolacion, Pto. Churruca, N. side of Fo. Nassau, Engel 5933 A, 5938 D
(MSC); F. Peel, Engel 5483C - c. sporo. (MSC); Cta. Rayo, Skottsberg
225 as syntype of Lophocolea patuli stipa (UPS); I. Chatham, N. shore of
B. Wide, Engel 5 304 A (MSC); S. side of I. Madre de Dios, head of fiord
E. of M. Roberto, Engel 5110 — c. sporo. (MSC); S. Molyneux, Dusen
s.n. (S); Pto. Charrua, Engel 4823, 4829 (MSC); Pto. Alert, Engel 4897 -
c. per., 5029 (MSC); ibid., head of fiord W. of M. Markham, Engel
5005 - c. sporo., 5070 - c. sporo. (MSC). PROV. AISEN: I. Magda-
lena, Schwabe 20/e as L. puccioana — c. per. -f tf (JE); ibid., Schwabe
20e-H (JE). PROV. CHILOE: Pto. Ballena, Engel 4197B (MSC), I.
Guaitecas, Dusen 352 as L. puccioana (JE); ibid., Melinca, Halle 210
(UPS); I. Guaitecas, Pto. Low, Dusen s.n. (JE); I. Chiloe, Pto. Quellon,
Halle s.n. (hb. Grolle); ibid., Halle 222 as Lophocolea otiphylla (UPS);
I. Chiloe, Aguas Buenas area, 4.7 km. along Aguas Buenas road from
Ancud-Quemchi road, ca. 100 m., Engel 12190 (F). PROV. OSORNO:
Antillanca, 700 m., Ruthsatz 59/30 (hb. Grolle); around L. Toro on road
to Refugio Antillanca, 855-885 m., Engel 4025 A, 4042B - c. sporo. + d",
4050C, 4063 A - c. per. + tf (MSC); Agua Caliente, R. Chanleufu, 4 km.
from Termas de Puyehue along road to Refugio Antillanca, 400 m.,
Engel 11480 - c. & (F). PROV. VALDIVIA: W. slope of Cord. Pelada
W. of El Mirador, between La Union and Pta. Hueicolla, 580 m., Engel
12280 (F); E. slope of Cord. Pelada, E. of El Mirador, between La Union
and Pta. Hueicolla, 740-840 m., Engel 12406, 12418, 12467 (F); R. Futa
in vicinity of Futa, 5.9 km. by road S. of junction of highway T-60 and
T-65, near sea level, Engel 10810 (F); Corral, Dusen 191 as L. puccioana
(NY); ibid. [Quitaluto], Hosseus 777 as L. ctenophylla (JE). ARGENTINA.
PROV. RIO NEGRO: Puerto Blest, Dusen 304 (NY); ibid., Dusen 504
(UPS). Patagonia: Sin. loc., Dusen s.n. as L. puccioana — c. per. (FH).
JUAN FERNANDEZ IS.: Mas a Tierra, Cordon del Barril, Skottsberg
249 as L. rotundifolia (JE).
ENGEL: CLASMATOCOLEA 91
Clasmatocolea humilis var. polymorpha Engel.
Clasmatocolea humilis var. polymorpha Engel, Phytologia 41 :309. 1979.
Holotype: Tristan da Cunha, above settlement, 620 m., 21 December
1938, Christopher sen & Mejland 52 (O!; isotype: S!).
Leaves erect, often connivent, with margins and apices entire; underleaves on any
given axis polymorphic: variable in shape and with apices often broadly rounded to
truncate to retuse, and only sporadically short-bifid, often with one lobe well developed
and the other represented as a tooth; underleaves distant, uniformly small for plant
size, those in apical portion (0.8-)1.0-1.3 X stem width, the underleaves generally
decreasing in size toward stem base, those toward stern base ca. 0.7 X stem width.
Ecology-Phytogeography. — Endemic to Tristan da Cunha, where over
dripping rocks at 620-1,200 m. (see fig. 24).
Specimen seen.— TRISTAN DA CUNHA : Above Burntwood, 1,200m.,
Christopher son 774 (hb. Grolle).
Clasmatocolea fasciculata (Nees) Grolle. Figures 25-27.
Jungermannia fasciculate. Nees, Hor. Phys. Berol. p. 46. pi. 5,/. 4. 1820
nonJ. fasciculata (Lindenb.) Hook. f. & Tayl. London J. Bot. 3:371.
1844 (= Plagiochild). Chiloscyphus fasciculatus (Nees) G. L. & N.
Syn. Hep. 190. 1845. Heteroscyphus fasciculatus (Nees) Schiffh.
Oesterr. Bot. Z. 60:172. 1910. Lophocolea fasciculata (Nees) S.
Arnell & Grolle in Grolle, Trans. Brit. Bryol. Soc. 3:584. 1959.
Clasmatocolea fasciculata (Nees) Grolle, Rev. Bryol. Lichenol. 29:72.
1960. Original material: South Africa, Cape Prov., Cape of Good
Hope, Ecklon (W, Lindenberg Hep. No. 4501 !).
Jungermannia bergiana Lehm. Nov. Minus Cogn. Stir. Pug. 4:43. 1832,
syn. fide G. L. & N. (1845). Original material: South Africa, Cape
Prov., Cape of Good Hope, Bergius 29 (W, Lindenberg Hep. No.
4499! &4500!).
Chiloscyphus fasciculatus var. /3 dregeana Nees in G. L. & N. Syn. Hep.
190. 1845, syn. fide Grolle (1959). Jungermannia dregeana Lehm. &
Lindenb. ex G. L. & N. Syn. Hep. 190. 1845, nom nud., pro syn.
Original material: South Africa, Cape Prov., Deutsch Kloof, 305-
610 m., Drege 1030 (W, Lindenberg Hep. No. 4497!).
Chiloscyphus fasciculatus var. y exarida Nees in G. L. & N. Syn. Hep.
190. 1845, syn. fide Grolle (1959). Original material: South Africa,
Cape Prov., summit of Table Mt., Ecklon (W, Lindenberg Hep. No.
4498!).
Chiloscyphus rabenhorstii Steph. Bull. Herb. Boissier 8 (1):51. 1908
(= Spec. Hep. 3:227), syn. fide Grolle (1959). Heteroscyphus raben-
FIG. 25. Clasmatocolea fasciculata (Nees) Grolle. 1, Portion of main axis, dorsal
view. 2, Portion of main axis with ventral-intercalary (vib) and lateral-intercalary (lib)
branches, lateral-ventral view. 3, Underleaves of main axis (semiflattened). 4, Leaves of
main axis (semiflattened). 5, Teeth of apices and margins of main axis leaves. Figures
1, 3a, e, 4a, b, from type of C. fasciculata, Ecklon, South Africa, Cape of Good Hope;
figures 2, 3d, 4c, 5b, c, e, from Esterhuysen 24164, South Africa, Cape Prov., Waterfall
Kloof; figures 3b, 4d, 5a, from type of Chiloscyphus fasciculatus var. /3 dregeana,
Drege, Deutsch Kloof; figure 3c, from Brunnthaler, South Africa, Table Mt.; figure
3f, from Esterhuysen 27356, South Africa, P. Formosa; figure 3g, from Drege 44,
South Africa, Cape of Good Hope; figures 4e, 5d, from Esterhuysen 25380, South
Africa, Cape Prov., Kromrivierkloof; figure 5f, from Esterhuysen 25128, South Africa,
Table Mt.
92
ENGEL: CLASMATOCOLEA
93
FIG. 26. Clasmatocolea fasciculata (Nees) Grolle. 1, Bracts and bracteole. 2, Perianth
mouth. 3, Gynoecium on ventral-intercalary branch; leaves and underleaves of main
axis removed. 4, Portion of main axis underleaf segment with a slime papilla. 5, Stem,
cross section. 6, Median leaf cells. Figures 1, 2, after Grolle (1959); figure 3, adapted
from Grolle (1959); figure 4, from type of Chiloscyphus fasciculatus var. /3 dregeana,
Drege, Deutsch Kloof; figures 5, 6, from type of C. fasciculata, Ecklon, South Africa,
Cape of Good Hope.
horstii (Steph.) Schiffn. Oesterr. Bot. Z. 60:172. 1910. Original
material: South Africa, Cape Prov., Table Mt., February 1884,
Rabenhorstf. (G!).
Lophocolea vermicularis Herz. Memoranda Soc. Fauna Fl. Fenn.
27:100. /. 44 f-i. 1952, syn. fide Grolle (1959). Holotype: South
Africa, Cape Prov., Table Mt., ca. 600-800 m., 16 April 1922,
Rolfes 22 (JE!).
Isotachis capensis Steph. ex Herz. Memoranda Soc. Fauna Fl. Fenn.
27:101. / 45 a-f. 1952, syn. fide Grolle (1959). Original material:
South Africa, Cape Prov., Table Mt., Rehmann s. n. (JE, "A. Reh-
mann: Hepaticae austro-africanae. N°. 33a"!; Cape Prov., Table Mt.,
Jelinek (JE, "Exped. Novara"!); Cape Prov., sin coll. (with Adelan-
thus unciformis) (non vidi).
o.
94
ENGEL: CLASMATOCOLEA 95
Plants soft, spongy, prostrate, mixed with other bryophytes or densely caespitose,
occasionally cushion-like, grey green, becoming brownish with age; axes to 4.7cm.
tall, 1.8 mm. wide (to 2.5 mm. wide in axes with spreading leaves).
Branches frequent, occasionally in irregular fascicles, usually of lateral and ventral
intercalary type, but Frullania-o^e quite common, rarely of Acromastigum-type;
branches usually with first 1-3 pairs of leaves bifid, especially on intercalary branches.
Stems 11-15 cells high, cortex in 1-3 rows of moderately thick-walled cells smaller
than medullary cells, the exposed wall of outer row moderately to very thickened;
cuticle smooth; endophytic hyphae absent. Rhizoids, when present, in fascicles from
stem near underleaf base.
Leaves with insertion very slightly to distinctly recurved at ventral end; leaves con-
nivent dorsally, lending axis a somewhat vermiform appearance, occasionally rather
strongly spreading or connivent in region of axis apices and spreading in region of axis
base; leaves loosely-densely imbricate, conchiform concave, wide-ovate to oblate to
occasionally ± reniform; apex broadly rounded to truncate to broadly retuse, undivided,
occasionally inflexed, the leaves entire or with a few teeth on apex or median-upper
ventral margin, rarely dentate on dorsal margin; teeth wide-triangular and sometimes ±
apiculate, or narrowly and often sharply triangular, occasionally terminating in a uni-
cellular row of several cells, the teeth commonly inflexed; dorsal margin plane to
sinuous, short to long decurrent; ventral margin very often inflexed, broadly rounded,
with basal portion enlarged, often extending ventrally beyond stem and in contact
with slightly spreading underleaf.
Leaf cells thin walled, trigones large to bulging, rarely confluent; median leaf cells
16-26(-31) n wide; 18-32 /* long; cuticle smooth.
Underleaves 2-3 X stem width, free, transversely or obliquely inserted, approximate-
imbricate, moderately to deeply convex, occasionally with basal portion abruptly convex
(ventral view), the margins curved dorsally; underleaves reniform to wide-ovate to occa-
sionally moderately ovate; apices 0.2-0. 3-bifid, the segments often apiculate, terminating
in a slime papilla, segment margins entire or few dentate, with armature terminating in
a slime papilla; margins of the lamina usually with I large often apiculate lacinium vari-
able in position, the margins otherwise entire or sparingly dentate-ciliate-laciniate.
Plants apparently dioecious ; androecia not seen.
Gynoecia on short ventral-intercalary branches; subfloral innovations absent; bracts
in 2 series, those of innermost series oblong-obovate, apical portion often sharply
inflexed or reflexed ; apices broadly rounded to truncate to subretuse, entire or sparingly
small-dentate; margins straight or reflexed, entire or with a few small teeth that often
terminate in slime papillae. Bracteoles of innermost series free from bracts, slightly
narrower than bracts, otherwise subequal to bract size, obovate; apices undivided or
short-bifid; margins plane or sharply inflexed or reflexed, entire or sparingly small-
dentate, the teeth often terminating in a slime papilla. Perianth inflated, obscurely
trigonous, wide-oblong, slightly narrowing toward a wide mouth; lobes broadly rounded,
often with portion of apex sharply reflexed, the lobes with numerous sharp teeth-laciniae
that occasionally terminate in a slime papilla; keel wings occasional, of few cells high.
Sporophyte not seen.
Variation. — The type material of C. fasciculata possesses both con-
nivent and rather strongly spreading leaves. This condition sometimes
occurs on a single axis, and the leaves then are connivent in the region
96 FIELDIANA: BOTANY
of the axis apex and spreading in the area of the axis base (see fig. 25-1).
Leaf armature of type plants is also variable — although many of the
leaves are entire, there are usually at least several on a given axis with
small to large, commonly inflexed teeth.
Differentiation. — I concur with Grolle (1959) that C. fasciculate, is most
closely allied to C. humilis. The following ensemble of features of C.
fasciculata will serve to distinguish it from that species: (a) the laciniate
underleaf lamina margins; (b) the restriction of perianths to short ventral-
intercalary branches; (c) the perianths that slightly narrow toward a wide
mouth, with the ventral lobes not infolded; and (e) the usually bifid
initial 1-3 leaves of intercalary branches.
Note. — Clasmatocolea fasciculata produces, on rare occasion, Acroma-
stigum-type branching. This, to my knowledge, is the first record of this
branch type in the Lophocoleaceae. The plastic branching found in this
species (ventral- and lateral-intercalary, Frullania- and Acromastigum-type
branches) marks this taxon as a rather primitive element within the genus.
Ecology-Phytogeography. — On wet rock in streams and on damp,
shaded south-facing slopes at altitudes of ca. 300-1, 200 m. The species is
endemic to Cape Prov., South Africa (see fig. 27).
Specimens seen.— SOUTH AFRICA. CAPE PROV. : "Kapland," sin.
coll. — c. young per. (JE); Cape of Good Hope, sin. coll. (W); ibid.,
Drege s.n., 44 (W); ibid., Tafelberg, ca. 900-1,000 m., Brunnthaler s.n. (hb.
Grolle, W); Table Mt., upper Disa Gorge, 760 m., Esterhuysen 25128 as
C. vermicularis (UPS); Table Mt., NE corner of Holy Hutchinson Water
Reservoir, Arnell 892 as /. capensis (G); Betty's B., Porter Nature Reserve
in kloof behind Botanic Garden, Magill 6295 (F); Riviersonderend Reeks
area, Riviersonderend, Wilman 556 as L. vermicularis (BOL); Rivierson-
derend Reeks, Schilpadkop area, 760-915 m., Esterhuysen 21102 as L.
vermicularis (BOL); Waterfall Kloof, off Elandskloof, off Du Toitskloof,
Esterhuysen 24164 (BOL); Kromrivierkloof, off Du Toitskloof, Esterhuy-
sen 25380 (BOL); Witteberg, Slanghoekberge, 915m., Esterhuysen 22268
as L. vermicularis (BOL); Mostertshoek, 1,220 m., Esterhuysen 24287
(BOL); foot of P. Formosa, Esterhuysen 27356 as L. vermicularis (UPS).
Clasmatocolea vermicularis (Lehm.) Grolle. Figures 28-30, Plates 11, 12.
Jungermannia vermicularis Lehm. Linnaea 4:361. 1829 non J. vermicu-
laris Hub. Hepaticol. Germ. p. 187. 1834 [= Lophozia sudetica (Nees)
Grolle]. Alicularia vermicularis (Lehm.) G. L. & N. Syn. Hep. 11.
1844. Nardia vermicularis (Lehm.) Trev. Mem. 1st. Lomb. Sci. Lett.
III. 4:400. 1877. Notoscyphus vermicularis (Lehm.) Steph. Bull.
Herb. Boissier 1 (2): 174. 1901 (= Spec. Hep. 2:35). Clasmatocolea
ENGEL: CLASMATOCOLEA 97
vermicularis (Lehm.) Grolle, Rev. Bryol. Lichenol. 29:78. 1960.
Original material: South Africa, Cape Prov., Table Mt., Ecklon
(FH!, NY!, S!).
Jungermannia flexuosa Lehm. Linnaea 4:361. 1829, syn. fide Stephani
(1901). Alicularia flexuosa (Lehm.) Nees in G. L. & N. Syn. Hep. 11.
1844. Notoscyphus flexuosus (Lehm.) Sim, S. African J. Sci. 12:20.
1916. Original material: South Africa, Cape Prov., Table Mt.,
Ecklon (G!).
Jungermannia subintegra Hook. f. & Tayl. London J. Bot. 3:477. 1844,
syn. fide Grolle (1960). Lejeunea subintegra (Hook. f. & Tayl.) G. L.
& N. Syn. Hep. 376. 1845. Lophocolea subintegra (Hook. f. & Tayl.)
Grolle, Trans. Brit. Bryol. Soc. 3:587. 1959. Original material:
Falkland Is., Hooker (FH!, NY!).
Jungermannia chamissonis Gott. & Lindenb. in G. L. & N. Syn. Hep.
668. 1847, syn. fide Grolle (1960). Leploscyphus chamissonis (Gott. &
Lindenb.) Mitt. Hooker's J. Bot. Kew Gard. Misc. 3:358. 1851.
Mylia chamissonis (Gott. & Lindenb.) Trev. Mem. 1st. Lomb. Sci.
Lett. III. 4:412. 1877. Leioscyphus chamissonis (Gott. & Lindenb.)
Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:445. 1885. Original
material: Chile, without specific locality, Chamisso (S! — c. cf).
Chiloscyphus nigrescens Lindenb. & Hampe in Hampe, Linnaea 24:640.
1851, syn. fide Grolle (1960). Original material: Costa Rica, (crateris)
Reventado, 3,050 m., Oersted (S!).
Notoscyphus variifolius Mitt. J. Linn. Soc., Bot. 16:188. 1877, syn. fide
Stephani (1901). Original material: South Africa, Cape Prov., Cape
of Good Hope, Eaton (NY!).
Clasmatocolea heterostipa Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15:441. pi. 20. 1885, syn. cf. Grolle (1959). Original material : Ecuador,
"Andes Quintensis," M. Pichincho, 2,700-3,400 m., Spruce (M! - c.
sporo., MANCH! - c. <?).
Nardia lindmanii Steph. Bih. Kongl. Svenska Vetensk.-Akad. Handl.
23 (III, 2): 25. 1897, syn. fide Grolle (1964). Alicularia lindmanii
(Steph.) Steph. Bull. Herb. Bossier 1 (5):481. 1901 (= Spec. Hep.
2:43). Notoscyphus lindmanii (Steph.) Schiffn. Hedwigia 51 :276. 1912.
Original material: Brazil, R. Grande do Sul, Porto Alegre, Lindman
42 (G! - c. sporo. + d", hb. Grolle!, M!).
Clasmatocolea chilensis Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 33. 1900, syn. cf. Grolle (1956). Lectotype (nov.):
Chile, Prov. Concepcion, Concepcion, 5 September 1896, Dusen 166
(G! — c. sporo.).
98
ENGEL: CLASMATOCOLEA 99
Lophocolea turbiniflora Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Hand!. 26 (III, 6):45. 1900, syn.fide Grolle (1960). Lectotype (nov.):
Chile, Prov. Valparaiso, Valparaiso, El Salto, Dusen 139 (G!).
Lophocolea flavovirens Steph. Kongl. Svenska Vetenskapsakad. Handl.
46 (9): 44. /. 19 d-g. 1911, syn. cf. Grolle (1956). Clasmatocolea
flavovirens (Steph.) Herz. Acta Horti Gothob. 15: 159. 1943. Lectotype
(cf. Grolle 1956): Chile, Prov. Chiloe, I. Guafo, Cta. Samuel, 25 July
1908, Halle 201 (UPS! - c. sporo.; isolectotypes: G!, JE!).
Lophocolea ligulata Steph. Kongl. Svenska Vetenskapsakad. Handl. 46
(9):47. /. 17 <?, / 1911, syn.fide Grolle (1960). Lectotype (nov.):
Chile, Prov. Chiloe, I. Guafo, (Cta. Samuel), Skottsberg (G! - c.
per.).
Lophocolea skottsbergii Steph. Kongl. Svenska Vetenskapsakad. Handl.
46(9) :53./. 20 f. 191 1, syn.fide Grolle (1960). Lectotype (nov.): Chile,
Prov. Chiloe, I. Chiloe, Ancud, Skottsberg s. n. (G! — c. per.).
Lophocolea boliviensis Steph. in Herzog, Biblioth. Bot. 21 (87):217. /
755. 1916, syn.fide Grolle (1960) non L. boliviensis S. Arnell, Svensk
Bot. Tidskr. 55:206. 1961. Lectotype (nov.): Bolivia, (Sillar, 1,800 m.),
1911, Herzog 2778 (G 17765!).
Lophocolea longissima Steph. in Herzog, Biblioth. Bot. 21 (87): 220. /
759 a. 1916, syn. nov. Holotype: Bolivia, (R. Corani, ca. 1,800 m.),
Herzog 4671 (G! - c. <?).
Alicularia grandistipula Herz. Biblioth. Bot. 21 (88):30./. 16. 1921, syn.
fide Grolle (1960) non A. grandistipula (Steph.) Horik. Hikobia 1 :30.
1950. Holotype: Bolivia, Cord, de Quimsacruz, near Mine Viloco,
4,400 m., 1911, Herzog 3183 (JE!).
Opposite:
FIG. 28. Clasmatocolea vermicularis (Lehm.) Grolle. 1, Portion of axis, lateral view.
2, Leaf. 3, 4, Underleaves. 5, Median leaf cells. 6, Stem, cross section. 7, Elater, apical
portion. 8, Underleaf margin with slime papilla. 9, Underleaf apices showing slime
papillae (some partially collapsed), roughed cuticle at lower right. 10, Seta, cross
section, lla-c, Antheridial stalks. 12, Outer capsule wall cells. 13, Inner capsule wall
cells. 14, Capsule wall, cross section. 15a-c, Antheridial bracts. 16, Perianth and sporo-
phyte-bearing plant, lateral view. Figures 1-6, from type of C. vermicularis, Ecklon,
South Africa, Cape of Good Hope; figures 7, 10, 14, 16, from Halle 185, Argentina,
Prov. Chubut, Estancia Miguens; figure 8a, from Esterhuysen E27342, South Africa,
near Joubertina; figures 8b, 9b, from Arnell 1698, South Africa, Gouna R.; figure 9a, d,
from Engel 3494, Falkland Is., Cheeks' Cr.; figure 9c, from Arnell 1695, South Africa,
Knysna; figures lla-c, 15b, from Schelpe 5140, Natal, Karkloof; figures 1-13, from
lectotype of C. chilensis, Dusen 166, Chile, Prov. Concepcion, Concepcion; figure 15a,
from Engel 3493, Falkland Is., Cheeks' Cr.; figure 15c, from Esterhuysen 25347, South
Africa, near Zondereinde.
FIG. 29. Clasmatocolea vermicularis (Lehm.) Grolle. 1, Axis, ventral view, with 2
Frullania-type branches. 2, Apex of gynoecial shoot with sporophyte, ventral view
(v = sporophyte valve; c = calyptra; pi = perianth lobe). 3, Axis, dorsal view.
4, Perianth mouth, ventral lobe in middle. 5-8, Axes with distal portion of leaves
removed, ventral view. Figures 1, 3, 7, from Engel 11632, Chile, Prov. Osorno, Salto del
Indio; figures 2, 4, from Engel 3939, Chile, Prov. Osorno, L. Toro; figure 5, from
Engel 11083, Chile Prov. Valdivia, Forestal Trafiin; figure 6, from Engel 12303, Chile,
Prov. Valdivia, Cord. Pellada; figure 8, from Engel 11240, Chile, Prov. Cautin, Salto
Palguin.
100
ENGEL: CLASMATOCOLEA 101
Odontoschisma variabile Sim, Trans. Roy. Soc. South Africa 15:77.
unnumbered plate on p. 77. 1926, syn. cf. Arnell (1958) non O. variabile
(Lindenb. & Gott.) Trev. Mem. 1st. Lomb. Sci. Lett. III. 4:419. 1877.
Jungermannia variabilis (Sim) S. Arnell, Bot. Not. 108:310. 1955,
nom. illeg. Original material: South Africa, without specific locality,
sin. coll. (non vidi).7
Lophocolea ovistipula Herz. Memoranda Soc. Fauna Fl. Fenn. 27:98.
/ 43 a-e. 1952, syn. cf. Grolle (1959). Lectotype (nov.): South Africa,
Cape Prov., Table Mt., ca. 600-900 m., 16 April 1922, Rolfes 68 (JE!).
Lophocolea subulistipa Herz. Memoranda Soc. Fauna Fl. Fenn. 27:99.
/. 44 a-e. 1952, syn. fide Grolle (1960). Holotype: South Africa,
Natal, Sandymount, 1922, Rolfes 136 p.p. (JE!).
Lophocolea elata f. aquatica Herz. Feddes Repert. Spec. Nov. Regni
Veg. 57:164. 1955, syn. fide Grolle (1960). Holotype: Colombia,
Eastern Cord., La. de Cunta, edge of Paramo de Santurban, 3,880 m.,
1927, Killip & Smith 17973 (JE!).
Lophocolea minima S. Arnell, Results Norweg. Sci. Exped. Tristan da
Cunha 3 (42):18./ 14 a-d. 1958, syn. fide Grolle (1960). Holotype:
Inaccessible I., NE of Blendon Hall, 3-4 m., Christophersen &
Me/land 2431 (Ol).
Plants rather soft, prostrate, but axes often sharply curved dorsally or ventrally,
usually pale green, occasionally brown, commonly so when dry, very rarely rose tinted
(several axes in a single collection — Engel 3494); axes to 2 cm. tall, 2.7 mm. wide.
Branches sparing, prostrate or arched, intercalary branches mostly ventral, rarely
lateral, terminal branches occasional, of Frullania-O^e.
Stems fleshy, 6-9 cells high, cortex in 1-2 rows of slightly thick-walled cells the same
size or slightly smaller than the medullary cells; endophytic hyphae occasionally
present in both cortex and medulla. Rhizoids, when present, in fascicles from stem near
underleaf base.
Leaves with insertion very slightly recurved at ventral end; leaves usually erect,
occasionally spreading, distant to imbricate, weakly to moderately to distinctly
concave, occasionally ± plane (especially in weak forms), ovate to oblong or subquadrate
or suborbicular (those on branches and depauperate stems small, often obcordate-
cuneate); apex broadly rounded to subtruncate to occasionally subretuse, very rarely
bifid (toward bases of poorly developed axes) ; margins and apices entire; dorsal margin
not or barely decurrent.
Leaf cells thin walled, trigones usually absent to small, occasionally medium; median
leaf cells 18-35(-42)M wide, 24-36(-48) M long; cuticle smooth or roughened and ap-
pearing finely granular. Oil-bodies colorless, very finely granulose-papillose, ellipsoid
to globose, consistently 2-3 per leaf cell, 7-13 X 6-8 /n.
7 I have been unable to locate original material of O. variabile Sim. It is not in PRE
or S; Arnell (1958) states he has seen the type, but does not indicate the herbarium in
which it is housed.
102 FIELDIANA: BOTANY
Underleaves highly variable in size, narrower than stem (normally slightly so), trans-
versely or obliquely inserted, usually free, occasionally weakly connate on 1 side,
weakly spreading (occasionally distinctly so), remote, slightly convex to slightly concave
(ventral view), narrowly ovate to lanceolate to subulate; apices undivided and acute
(less often narrowly rounded), or very often bidentate, or to 0.3(-0.5)-bifid; segments
directed toward stem apices or incurved, narrowly lanceolate-subulate or merely
reduced to a slime papilla, the segments terminating in a slime papilla, the sinus nar-
row; margins of the lamina entire, or with 1 slime papilla, or with 1 small tooth termi-
nating in a slime papilla.
Plants dioecious; androecia terminal or intercalary on leading axes or rather short
branches, bracts with basal portion strongly saccate, the distal portion strongly concave
or saccate in basal half, with distal portion moderately recurved; lobule margin with
2-several slime papillae that are often accompanied by 1 or 2 few-celled teeth, each
terminating in a slime papilla; antheridia solitary, stalks biseriate throughout, or
biseriate and with scattered uniseriate areas or uniseriate and with a few biseriate areas.
Gynoecia on leading axes; subfloral innovations occasionally present, originating
from below gynoecium, from below perianth, or from inside perianth; vestigial stem
perigynium present; bracts in 2-3 series, =*= appressed to perianth, the bracts of inner-
most series usually with lateral margins free, occasionally with one complete margin
fused with perianth, the bracts ovate to elliptic; apices =t truncate or retuse; margins
entire or with a single tooth on ventral margin, rarely with both dorsal and ventral
margins 1 -dentate. Bracteoles of innermost series free from bracts, the lateral margins
free, very rarely with 1 complete lateral margin fused with perianth; bracteoles ovate
or lanceolate, very rarely broadly obovate; apices undivided and acute or broadly
rounded, more often emarginate or bidentate or bifid to ca. 0.3; lamina margins entire
or often with 1 or both margins with a single tooth or slime papilla, rarely laciniate.
Perianth large, subterete, or obscurely to distinctly trigonous, clavate to campanulate;
perianth mouth wide and open, the lobes occasionally free for ca. 0.3 perianth length,
the three lobes broadly rounded, often undulate, often incurved, entire, rarely with the
lateral or just the ventral lobe bidentate or bilobed; keels occasionally winged, secondary
wings occasionally present. Calyptra large, often extending to perianth mouth.
Seta 5-10 cells in diameter, with 16-31 rows of outer cells surrounding an inner core
of scattered cells equal to or slightly larger than outer row. Capsule oblong to globose,
the valves 0.53-0.77 mm. long, 36-54 n thick, of 3-4 layers, the outer row of cells 17-24 /*
thick, equal to thickness of any 2 of interior strata; outer layer with red-brown, nodule-
like thickenings that are frequently feebly tangentially dilated, semiannular bands
absent; intermediate and inner layers of cells subequal in thickness, delicate, often
collapsing; inner layer of cells 6-11 n thick, with red-brown nodulose thickenings on
radial walls similar to those of outer layer, the thickenings often tangentially dilated,
semiannular bands often present.
Spores large, 15-20/u, yellow-brown, exine with the light microscope appearing
minutely punctate, but under the SEM with a network of wide, rather short, vermiform
ridges covered with often crowded nanogranules, the intervening hollows between
ridges sometimes with isolated nanogranules; spores averaging 1.9 X elater diameter.
Elaters 6-12 /x wide, slightly tapered toward the ends, apical portions often with thick
non-spiral walls, elater walls yellow-brown, 3-5 /x in diameter.
Variation. — Clasmatocolea vermicularis exhibits considerable variability,
especially with regard to color, axis stature, the size, concavity, and
ENGEL: CLASMATOCOLEA 103
configuration of leaves and the configuration, apices (especially the sinus
depth), and the disk marginal armature of underleaves. The leaf cells may
have trigones absent to medium. See Grolle (1960, pi. 3, but with erroneous
legend) for figures illustrating leaf and underleaf variation. As Grolle (he.
cit.) indicates, the variations listed above occur in all combinations and
without regard to geography.
The gynoecia of type material of Nardia lindmanii present an interesting
variant. The ventral margin of the bract nearest the perianth is completely
fused with the perianth. In addition, one complete margin of the bracteole
of the innermost series is likewise fused. The bracteole shape is also
deviant, being broadly obovate with apices undivided and broadly round-
ed. See Schiffner (1912) for a discussion of this species, which he transfers
to Notoscyphus.
Spruce (1885) and Stephani (1906) state the perianth mouth of C.
heterostipa is three-four lobed, whereas Schiffner (1893) states the perianth
mouth is two-four lobed in the genus. Clasmatocolea vermicularis (= C.
heterostipa), however, always has three lobes (fig. 29-2, 4). These lobes are
nearly always broadly rounded and entire and only rarely is the lateral or
just the ventral lobe bidentate (as in several New Zealand collections) or
more deeply divided (as in pi. 1, fig. 1, in Grolle, 1956).
Differentiation. — Clasmatocolea vermicularis may be separated from all
other Clasmatocolea taxa by the following ensemble of features: (a) the
ovate-oblong or suborbicular or subquadrate leaves that are at least
slightly adaxially concave; (b) the mostly narrowly ovate to lanceolate to
subulate underleaves that are narrower than the stem width; (c) the
underleaf apices that are usually bidentate or divided to at most 0.3(-0.5),
and with lamina margins entire or with 1 small tooth; and (d) the clavate
to campanulate, inflated perianths with mouths wide and open and with
lobes broadly rounded and nearly always entire.
Some phenotypes of C. vermicularis that do not possess distinctly
adaxially concave leaves may offer some confusion with Lophocolea
sabuletorum. Clasmatocolea vermicularis has leaves that exhibit at least a
slight concavity and have leaf apices broadly rounded or truncate, at most
subretuse, and never emarginate or dentate. Lophocolea sabuletorum has
at least some leaves slightly convex, and with leaf apices variable, i.e.,
broadly rounded to truncate or often retuse to emarginate to 1- or biden-
tate. Clasmatocolea vermicularis and L. sabuletorum not infrequently have
androecia, the antheridial stalk of which will immediately distinguish the
taxa, with C. vermicularis having at least locally biseriate stalks, and L.
sabuletorum, uniseriate stalks.
Jungermannia subintegra was described from the Falkland Islands and is
104 FIELDIANA: BOTANY
a synonym of C. vermicularis. This treatment is based upon examination
of a syntype of the former species and also upon the work of Grolle
(1960). Mizutani & Hattori (1959), however, treat the species as a synonym
of Lophocolea novae-zelandiae. There should be little confusion concern-
ing the disposition of /. subintegra, because the adaxially concave leaves
immediately place the plant within the genus Clasmatocolea.
If sterile South African plants are at hand, C. vermicularis may be
confused with the superficially similar Notoscyphus belangerianus (Lehm.)
Mitt. The two taxa may be easily separated by examining leaf prepara-
tions— N. belangerianus has a striate-papillose leaf cuticle and large to
knotlike trigones.
Notes. — 1. In a packet glued to the sheet with syntype plants ofJunger-
mannia subintegra are several plants of Lophocolea sabuletorum (leaf apices
frequently emarginate, antheridial stalk cells unistratose). These plants
should not be confused with those of the type.
2. Grolle (1960) has referred to Lophocolea ligulata as a "nomen mix-
turn," because the original specimen contains material of both C. vermicu-
laris (p.p. maj.) and L. sabuletorum (p. p. min.), and the original diagnosis
contains portions from both species. The description of the perianth was
based upon material of C. vermicularis, and that of the sterile structures on
material of L. sabuletorum. Grolle (loc. cit.) has typified the specimen as
one of C. vermicularis (because most of the material is of this taxon), but
he was in error, however, in stating Stephani originally described L.
ligulata as monoecious. Grolle has extracted several portions of Stephani's
description in Species Hepaticarum (vol. 6, p. 280, 1922); Stephani's
original description (1911) clearly states his new species is dioecious.
I have seen two Halle-collected specimens (c. per. + sporo.) of L.
ligulata from the Uppsala Herbarium and one collected by Skottsberg
(c. per.) from the Geneva Herbarium. Stephani (1922) effectively eliminates
the Halle collections from further consideration as lectotype candidates
because he lists only Skottsberg as the collector.
3. Of the eight syntype localities listed by Stephani (1911) for Lophocolea
skottsbergii, only one (Isla Chiloe, Ancud) is given for the species in
Stephani (1922). Stephani, in the latter publication, has effectively lecto-
typified the species. Among the plants of the lectotype (G), there are some
with suborbicular leaves and others with oblong leaves. Stephani notes
only suborbicular leaves in his original description. The syntype specimens
from Rio Olivia, near Ushuaia are of C. humilis.
4. Grolle (1960) lists Odontoschisma africanum (Pears.) Sim as a "pseudo-
synonym" of C. vermicularis. As discussed in Grolle (loc. cit.), the basio-
nym of O. africanum is Cephalozia denudata var. africana Pears, and is
ENGEL: CLASMATOCOLEA 105
actually a specimen of Odontoschisma. However, the description and plate
of O. africanum in Sim (1926) plus his herbarium specimens are actually C.
vermicular is. Arnell (1955b) utilized the information in Sim (he. cit.)
rather than that of the original description and type and mistakenly made
the combination Notoscyphus africanus (Pears.) S. Arnell (nom. illeg.).
Specimens of TV. africanus are actually C. vermicular is. Arnell (1963) later
recognized Odontoschisma africanum.
5. Herzog (1952, p. 98) states for the locality of Lophocolea ovistipula:
"Kapland: Tafelberg, an feuchten Steinen und Felsen, leg. L. Rolfes, IV.
1922". Among the specimens of L. ovistipula labeled "Herbarium Theodor
Herzog Jena" (JE) are three syntypes, all of which bear the note "Herz. n.
sp." Of the three, two (identical) are androecial specimens collected on
Devils-Peak, but bear no ecological data. The third, however, is labeled
"Tafelberg, feuchte Steine an Bachen" and is sterile. The third collection
has been selected as the lectotype; it matches closely with the locality,
plate, and description data, including the lack of sex organs.
6. The protologue and original material of Lophocolea boliviensis Steph.
(Stephani, 1916) contains elements of both C. vermicularis and Jungerman-
nia sp. Stephani's description and figure of the perianth are referable to
plants of Jungermannia, whereas the description and figure of the under-
leaves, plus the description of at least some of the other sterile features,
belong to Clasmatocolea. There were several problems involved with
typification, as illustrated in the following discussion of the specimens
studied.
Specimen 1: G 17765, leg. Herzog 2778, which by collection number is
referable to the specimen from Sillar, Bolivia. The plants are of C.
vermicularis and look much like Stephani's figure (a). However, the under-
leaves are not connate on both sides (as in the diagnosis and figure) but
only on one side, with the second underleaf margin in close proximity to
the leaf base. I did not observe leaves with the broad tooth as in the
Stephani figure. There was a single well-developed perianth present — it
was of C. vermicularis. No plants of Jungermannia were observed, but a
few reddish plants were seen.
Specimen 2: G 17766, leg. Herzog 2778, and thus also from Sillar. This
appeared to be a duplicate of G 17765, but in this case there were no
perianths, but rather some androecial individuals. Like the above, no
Jungermannia was found.
Specimen 3: JE collection of Herzog 3869 and labeled "Bei Altamachi,"
and thus is the second of two localities mentioned in the protologue. The
plants of C. vermicularis have underleaves free and not with both margins
£
O
106
ENGEL: CLASMATOCOLEA 107
in close proximity to the leaf bases. Intermixed were sterile plants of
Jungermannia sp., leaving me with the mystery of the origin of the perianth
data in the protologue.
The specimen of G 17766, being closest to the concepts embodied in
the protologue, was selected as lectotype.
7. Nearly all collections of this species in New Zealand were thought to
be C. strongylophylla, with a few identified as C. paucistipula.
8. See Grolle (1960) for additional remarks on C. vermicularis; see also
his pi. 1. Refer also to Grolle (1956) for comments (sub C. heterostipa);
see also his pi. 1.
Ecology. — Frequently on shaded, damp earth or rocks of stream or river-
bed or banks. Here the plants are subject to inundation during wet
seasons or periods of flooding, but then become exposed or grow just
above water level during drier periods. Also, in either forests or exposed
sites on dripping rock walls, steep, often irrigated slopes of gulleys or
gorges, or clayey banks. Occasionally over calcareous sandstone (New
Zealand). Not uncommon on roadside banks. For notes on ecology in
Colombia see Gradstein et al. (1977, p. 392-393).
Phytogeography. — Pan-temperate; New Zealand, South I. (455-1,830
m.), North I. (300-1,500 m.), Great Barrier I. (275 m.); Tasmania (1,250
m.) ; Kerguelen ; Crozet Is. ; Marion I. (25-100 m.) ; Mascarene Is., Reunion
(1,550-1,570 m.); Madagascar (900-2,200 m.); South Africa, Cape Prov.
(215-900 m.), Natal (1,200-2,440 m.), Transvaal (795 m.); Rhodesia
(1,435 m.); Zaire (2,080 m.); Tanzania (1,500-1,725 m.); Burundi (1,340-
2,050 m.); Tristan da Cunha Group, Inaccessible I. (3-4 m.), Tristan da
Cunha (sea level-700 m.); South Georgia; Falkland Is. (sea level-75 m.);
Chile, rare in Tierra del Fuego, sporadic in Patagonian Channels, Valdi-
vian region (140-1,600 m.), Prov. Valparaiso (20-30 m.), Prov. Santiago
(1,200 m.); Argentina, Prov. Chubut, Prov. Rio Negro (1,800 m.), Prov.
Neuquen (820 m.); Juan Fernandez Is. (150-550 m.); Brazil; Bolivia
(1,800-4,470 m.); Peru (2,000-3,600 m.); Ecuador (2,700-3,400 m.); Co-
lombia (2,500-4,520 m.); Venezuela (3,500 m.); Costa Rica (1,500-3,050 m.)
(see fig. 30).
Specimens seen.- NEW ZEALAND. SOUTH ISLAND. OTAGO:
Bank of Matau Branch of Clutha R., near Kaitangata, Allison s.n. (hb.
Grolle); ibid., Allison H6413, H6818 as C. strongylophylla (CHR); Dune-
din, Signal Hill, Allison H1412 as C. strongylophylla (CHR); Old Man
Range, below The Old Man, ca. 1,220 m., Schuster 67524 as C. strongylo-
phylla (hb. Schuster); Pigroot, 3 mi. from Kyeburn, 455 m., Child 1683
as C. ? strongylophylla (hb. Child, F); Danseys Pass, near Naseby, ca.
108 FIELDIANA: BOTANY
940 m., Irwin 16 p.p. as C. slrongylophylla (CHR); Naseby, ca. 600 m.,
Allison H5443 as C. strongylophylla (CHR); St. Bathans, McConnochie
421, 432 as C. strongylophylla (hb. Monash Univ.). WESTLAND:
Fiordland, Olivine Mts., upper Dart Valley, ca. 1,830 m., Campbell s.n. as
C. zotovii (CHR); Regina Valley, 1,220 m., Teague s.n. as Lophocolea
austrigena (CHR). MARLBOROUGH: Pelonis R., Martin H601 as C.
paucistipula (CHR). NORTH ISLAND: Sin. he., Berggren 3701 as L.
paucistipula (LD). WELLINGTON: Wairarapa, near Tinui, Hewitt s.n.
(hb. Grolle); Cross Cr., Haast Road, ca. 460 m., Child 1896 as C. strongy-
lophylla (hb. Child, F); Kaimanawa Range, 1,500 m., Druce s.n. (hb.
Grolle). HAWKE'S BAY: Kiwi Valley, Wairoa, Hodgson 4153 (hb.
Grolle). GISBORNE: Rosie B., L. Waikaremoana, Hodgson 8387 (CHR,
hb. Grolle — c. c?1); Te Tara-a-Tu Pillar, Mt. Maungapohatu, Urewera,
1,065 m., Sainsbury s.n. as C. strongylophylla (CHR). SOUTH AUCK-
LAND: Taupo, Berggren 3704 as L. paucistipula (LD); Kaingaroa Plains,
near Rotorua, Allison H3718 as C. strongylophylla (CHR); ibid., Allison
328 — c. sporo. (hb. Grolle); plateau E. of Waiotapu Valley, 550 m.,
Allison H3566 as C. strongylophylla (CHR); ibid., 600 m., Allison 326 (hb.
Grolle); near Atiamuri, S. of Rotorua, Allison H387 — c. sporo., H470 —
c. per. as C. paucistipula (CHR); ibid., ca. 305 m., Allison H388, H3369 as
C. strongylophylla (CHR); ibid., 305 m., Allison s.n. as C. strongylophylla
(hb. Child, F); ibid., Allison H3369 as C. rotata (hb. Grolle); Cormandel
Pen., Matthews 37 as C. paucistipula (CHR). GREAT BARRIER ISLAND :
Hauraki G., Kiwiriki B., 275 m., Lloyd 219 as L. austrigena (CHR).
TASMANIA: Cradle Mt. Region, Plateau Cr. area, between Cradle
Plateau and Marions Lookout, NNW of Cradle Mt., 1,250 m., Engel
13974 (F); Cradle Mt. Region, Marigold Cr., W. of and draining into
Crater L., vicinity of Crater P. Lookout, 1,250 m., Engel 14108 (F).
KERGUELEN: Moseley s.n. (hb. Grolle). CROZET IS.: I. Possession,
Port Alfred, Hebrard 10 (JE); I. de 1'Est, Vallee du Naufrage, Hebrard 40
(JE); I. de 1'Est, Cap Nord, Hebrard 67 I a (JE). MARION L: Rand 3782
(hb. Grolle); SW of Met. Station, 25 m., Huntley 119c (hb. Grolle);
Macaroni B., 25 m., Huntley 916 (hb. Grolle, PRE); N. of Juniors Kop,
100 m., Bakker 18b (hb. Grolle). MASCARENE IS. REUNION: Plaine
d'Affouches, Hebrard 24j a (JE); Plaine des Cafres, 1,550-70 m., Gimalac
74. R. 8341, 74R. 8382 (JE). MADAGASCAR: Foret de Ranomafana, 60
km. N. of Fianarantsoa, ca. 900 m., Onraedt 71. M. 5061 (hb. Grolle); Soa-
vimbahoaka, above Tananarive, Onraedt 71. M. 5197 (hb. Grolle); Omka-
ratra, ca. 2,200 m., Cremers 2381 - c. per. (JE). SOUTH AFRICA. CAPE
PROV.: "Kapland," sin. coll. as Chiloscyphus vallisgratiae Gott. (JE);
ibid., sin. coll. as Jungermannia congesta (JE); Peninsula, Chapmans P., Ar-
ENGEL: CLASMATOCOLEA 109
nell 1127 as L. ovistipula (BOL); Peninsula, Constantianek, Arnell 169, 171,
181 (BOL); ibid., Arnell 170, 318 (UPS); Peninsula, Rhodes Road, near
Constantianek, 215 m., Garside 6685, 6693 (UPS); Table Mt., Constan-
tianek, Arnell 181 (UPS); Table Mt., Silverstream Ravine, 915 m., Ester-
huysen 25113 as L. ovistipula (UPS); Peninsula, Oranjekloof, Arnell 2160,
2165, 2185, 2210 (BOL); ibid., Arnell 2200 (PRE); ibid., Arnell 2183
(UPS); Tafelberg, 600-900 m., Rolfes s.n. as Lophocolea subrotunda (JE);
ibid., Rolfes s.n. as L. ovistipula fo. aquatica (JE); Devils P., Arnell 2215 as
L. ovistipula (BOL); ibid., 300-400 m., Rolfes 85 as syntype of L. ovisti-
pula — c. c? (JE); Groot Drakenstein, near Stellenbosch, Cholnoky s.n.
as L. subulistipa (UPS); near Kirstenbosch, Arnell 171 as L. ovistipula
(BOL); Caledonrivier, Riviersonderend Reeks, near Greyton, 610 m.,
Esterhuysen 21103 as L. subulistipa (UPS); Riviersonderend Reeks, near
Zondereinde, Esterhuysen 25347 — c. tf (UPS); Du Toitskloof, Ester-
huysen 19887, 25413 (BOL); Kromrivierkloof off Du Toitskloof, Ester-
huysen 25412 as L. subulistipa (UPS); ibid., Esterhuysen 25413 as L.
ovistipula (UPS); foot of Waaihoek P., 305-610 m., Esterhuysen 24151 as
L. ovistipula (BOL, UPS); Gouna Forest Station, Arnell 1695 (UPS); ibid.,
Gouna R., Arnell 1698, 1710 (UPS); N. foot of Tsitsikama Mts., near
Joubertina, Esterhuysen 27342 as L. subulistipa (UPS); Baviaanskloof,
Esterhuysen 24996 as L. ovistipula (BOL). NATAL: near Escourt, West
s.n. (UPS); Braco, Karkloof, 1,220 m., Schelpe 5140 (BOL, UPS - c. d");
Cathedral P. Forest Station, Catchment no. 1 Research Area, Magill
5650 (F); Mont Aux Sources, Koedoepas, 2,440 m., Sim 1613 as L.
subulistipa (UPS); Zululand, Ngome-Bosreserwe, 1,200 m., Oliver 7096
(F). TRANSVAAL: Farm Oshoek, E. of Wakkerstroom, Tolken s.n. (F);
Forst Ceylon, near Sabie, Cholnoky s.n. (UPS); Bonnet Junction and
Mac-Mac Falls, near Sabie, Cholnoky s.n. as L. ovistipula — c. tf (UPS);
5 mi. SW from Duivelskloof, 795 m., Schelpe 6031 as L. ovistipula (BOL,
UPS). RHODESIA: Umtali, near Odzani R. Bridge, 1,435 m., Schelpe
5650 as L. ovistipula (BOL, UPS). ZAIRE: Near Uvira, S. of Kwamwali-
mu, 2,080 m., Symoens 2153 (hb. Vanden Berghen). TANZANIA: Kili-
manjaro, near Kibo Hotel, 1,500 m., Esterhuysen 27263 as L. ovistipula
(UPS); Southern Highlands, N. foot of Mt. Kyejo, E. of Tukuyo town,
1,725 m., Pocs 67691 B (hb. Vana). BURUNDI: S. of Rutana, near Mu-
tambala, 1,340 m., Symoens 2762a (hb. Vanden Berghen); near Mu-
ramvya, 2,050 m., Symoens 2384 (hb. Grolle - c. per., hb. Vanden
Berghen). TRISTAN DA CUNHA: 620-700 m., Christophersen & Mejland
42, 51 as L. skottsbergii (O); 30-60 m., Christophersen & Mejland 198, 310
as L. strongylophylla (S); 650 m., Christophersen & Mejland 839 as L.
strongylophylla (S). SOUTH GEORGIA: Cumberland B., Skottsberg 231
110 FIELDIANA: BOTANY
as syntype of L. skottsbergii - c. tf (L). FALKLAND IS. (selected):
Hooker s.n. as Lophocolea grisea — c. a" (NY); Skottsberg s.n. as syntype
of L. skottsbergii - c. per. (G). EAST FALKLANDS: Port Louis, Halle
& Skottsberg 236 as Lophocolea vasculosa (NY); ibid., Skottsberg s.n. as
L. austrigena — c. per. (S); Port William Region, N. shore of Cape
Pembroke Peninsula, E. of Yorke Pt., Engel 2446 (MSC). WEST FALK-
LANDS: Fox Bay Region, near mouth of Cheek's Cr., ca. 12 m., Engel
3493 - c. cf , 3494 - c. <?, 3495 (MSC). ARGENTINA. TERR. TIERRA
DEL FUEGO: I. de los Estados, Pto. San Juan del Salvamiento, Spegaz-
zini 83 as Lophocolea grisea v. lava ? (NY); I. de los Estados, Pto. Basil
Hall, Spegazzini86 as L. grisea v. lava ? (NY). CHILE. PROV. MAGAL-
LANES: Brunswick Pen. (cf. Engel, 1978); S. Skyring, Cabo Leon,
Hatcher coll. (cf. Engel, 1973); S. Skyring, B. Pinto, Halle & Skottsberg
231 as syntype of L. skottsbergii (BM). PROV. AISEN: Cta. Conner,
Halle 201 as syntype of L. flavovirens (UPS); I. Magdalena, Schwabe
20/1 as Lophocolea minuta (JE); C. Tesoro, 1,000 m., Schwabe 39 c p.p.,
39/62 as Lophocolea tesorensis Herz., nom. hb. (JE). PROV. CHILOE: I.
Guafo, C. Samuel, Halle s.n. — c. per. (hb. Grolle); ibid., Halle 216 as syn-
type of L. ligulata — c. per. -f sporo. (UPS); I. Chiloe, Quemchi, Skottsberg
201 as syntype of L. flavovirens (UPS). PROV. LLANQUIHUE: Calbuco,
Schwabe s.n. (hb. Grolle); ibid., Schwabe 35 (JE); ibid., Schwabe 54 — c.
sporo., 777 as C. heterostipa (JE); ibid., Schwabe 125 as L. skottsbergii
(JE); Puerto Varas, Dusen s.n. as syntype of C. chilensis (BM, LD — c.
per.); ibid., Dusen 473 as syntype of C. chilensis (UPS). PROV. OSORNO:
L. Rupanco, Schwabe 51 as C. heterostipa (JE); L. Toro, on road to
Refugio Antillanca, 860 m., Engel 3977, 3989 - c. sporo. (MSC); valley
of R. Nauto near road to Refugio Antillanca, between La. El Encanto
and L. Toro, 730 m., Engel 4080, 4 105 A - c. sporo. (MSC); Termas de
Puyehue, Schwabe 111 (hb. Grolle). PROV. OSORNO near Prov. Valdivia
boundary: Anticura, Salto del Indio, 19 km. by road E. of Termas de
Puyehue along international highway, 300 m., Engel 11632, 11638,
11644 - c. sporo. (F). PROV. VALDIVIA: W. slope of Cord. Pelada,
W. of El Mirador on road between La Union and Pta. Hueicolla, 580 m.,
Engel 12303 — c. per. (F); Corral, Krause s.n. as Leioscyphus chamissonis
(S); ibid., Amargos, Hosseus 835 — c. <?, 846 as C. heterostipa (JE);
Panguipulli, 140 m., Hollermayer 662* as Leioscyphus chamissonis (JE);
Forestal Trafun, Engel 11083 (F); L. Pellaifa, Schwabe 9 p.p. as L. subin-
tegra fo. humilis (JE); SW of La. Los Patos, SW slope of V. Quetrupillan,
1,450-1,600 m., Engel 11133 (F). PROV. CAUTIN: Termas de Palguin,
along R. Palguin, 730 m., Engel 11316 (F); Salto Palguin, R. Palguin, road
to Termas de Palguin, 580 m., Engel 11240 (F); Pucon, Hosseus 170 as C.
ENGEL: CLASMATOCOLEA 111
heterostipa — c. tf (JE, M); Parque Nac. Conguillo, El Salto, 1,110m.,
Mahu s.n. - c. <? (hb. Mahu). PROV. MALLECO: Mininco, Schwabe
157 '/e (hb. Grolle); Parque Nac. Nahuelbuta, casa de Pincheira, 1,250 m.,
Mahu 6037 (JE, hb. Mahu); ibid., Mahu 6093 (hb. Grolle, hb. Mahu).
PROV. MALLECO/PROV. ARAUCO: Cord. Nahuelbuta, Aguas Cali-
entes, Parque Nac. Nahuelbuta, ca. 48 km. W. of Angol, 1,240 m., Engel
12614 (F). PROV. CONCEPCI6N: Conception, Dusen s.n. as syntype of
C. chilensis (BM); ibid., Dusen 157 as syntype of C. chilensis (G — c. per.,
S); ibid., Dusen 161 as syntype of C. chilensis (BM — c. per., FH — c. tf,
G — c. per., O); ibid., Dusen 346/2 as syntype of C. chilensis (M); Lota,
Skottsberg 159 as C.flavovirens (JE). PROV. VALPARAISO: Valparaiso,
(El Salto), Dusen s.n. as syntype of L. turbiniflora (BM — c. <? , LD); ibid.,
Dusen 139 as syntype of L. turbiniflora (BM — c. cf1, FH, JE, O, S — c. <?,
UPS); El Quisco, Q. Guallelemu, 30 m., Mahu 10327 '/a (JE); ibid., 20 m.,
Mahu 1 1374 (F), ibid., 20-30 m., Mahu 11020, 11391 (hb. Mahu). PROV.
SANTIAGO: Caleu, 1,200 m., Mahu 8611 (hb. Grolle, hb. Mahu).
ARGENTINA. PROV. CHUBUT: Colonia Diez-y seis de Octubre, Halle
204 as Lophocolea georgiensis (UPS); ibid., Halle 185 asL. austrigena — c.
sporo. (S). PROV. RIO NEGRO: L. Nahuel Huapi, C. de Los Hormigas,
R. Nirihuau, ca. 1,800 m., Hosseus 626a as Lophocolea patulistipa (JE).
PROV. NEUQUEN: L. Nahuel Huapi, L Victoria, 820 m., Sleumer 1722
as L. diversistipa (UPS). JUAN FERNANDEZ IS.: 150-200 m., Kunkel
334/5a (B); Mas a Tierra, head of Pangal Valley, Hatcher & Engel 18
(JE); Mas a Tierra, base of Piramide, Hatcher & Engel 534 (JE); Mas a
Tierra, near Port, de Villagra, 550 m., Hatcher & Engel 704 (JE). BOLIVIA.
DEPTO. COCHABAMBA. PROV. QUILLACOLLO: Cord, del Tunari,
above and SW of La. Marquina, 4,470 m., Lewis 2510, 2517 (F). PROV.
CHAPARE: R. Corani, ca. 1,800 m., Herzog 4719 as Chiloscyphus
parvistipulus — c. sporo. + <? (M). PROV. AYOPAYA: Altamachi, ca.
3,500 m., Herzog 3869 as syntype of L. boliviensis (JE). PERU. PROV.
OTUZCO: Casmiche, Hegewald 6005 (hb. Hegewald); ibid., Hegewald
6503 (hb. Hegewald, JE); C. las Gordas, between Quiruvilca and Agall-
pampa, 3,600 m., Hegewald 5970 (hb. Hegewald, F); ibid., between Los
Alisos and Quiruvilca, 3,600 m., Hegewald 5975 (hb. Hegewald). PROV.
CONTUMAZA: "Tambo de Lima," between Contumaza and Cascas,
2,000 m., Hegewald 7379 (F, JE — c. per.); at Tunnel between Contumaza
and Cascas, C. Luden, 2,400 m., Hegewald 7356 (F, JE). PROV. CA-
JAMARCA: Puente Bajo between Encanada and Celedin, 3,550 m.,
Hegewald 6596 - c. per. (hb. Hegewald, JE). PROV. CHACHAPOYAS:
Calla Calla, between Balsas and Leimebama, ca. 3,100 m., Hegewald
6938 (hb. Hegewald, JE). ECUADOR: R. Cusatagua, near Quito, Spruce
112 FIELDIANA: BOTANY
s.n. as Leioscyphus chamissonis (MANCH). COLOMBIA. BOYACA:
NW of Belen, Q. Minas, 3,800 m., Cleef2088a (U, F). CUNDINAMAR-
CA: Salto de Tequendama, ca. 2,500 m., Schaeck 72 Am. 33 (hb. Grolle).
VENEZUELA: Sa. de St. Domingo, 3,500 m., Oberwinkler & Poelt
HV69-10, HV69-25 (JE). COSTA RICA. PROV. CARTAGO: Ca. 10 km.
SSE of Tapanti along road on E. slope above R. Grande de Orosi, 1,500-
1,550m., Engel 8343, 8397 (F). PROV. SAN JOSE: La Palma area,
above R. La Hondura valley ca. 9 km. NE of San Jeronimo, ca. 1,575 m.,
Engel 8815, 8819 (F).
Clasmatocolea gayana (Mont.) Grolle. Figures 31-33, Plates 13-14.
Jungermannia gayana Mont. Ann. Sci. Nat. Bot. III. 4:349. 1845.
Chiloscyphus gayanus (Mont.) G. L. & N. Syn. Hep. 710. 1847.
Lophocolea gayana (Mont.) Mitt, in Seemann, Fl. Vit. 404. 1873.
Clasmatocolea gayana (Mont.) Grolle, Rev. Bryol. Lichenol. 29:72.
1960. Original material: Chile, Prov. Valdivia, Valdivia, Gay 44
(FH! - c. sporo., S!).
Lophocolea vinciguerreana Mass. Nuovo Giorn. Bot. Ital. I. 17:229.
pi. 18, f. 1-9. 1885, syn.fide Bescherelle & Massalongo (1889). Lecto-
type (fide Engel, 1978): Chile, Prov. Magallanes, I. Basket, Cabo
Desolation, June 1882, Spegazzini 317 (VER!).
Conoscyphus flaccidus Pears. Bull. Misc. Inform. 1922:252. / pi. 1922,
syn. nov. Holotype: Chile, Prov. Magallanes, Punta Arenas, March
1868, Cunningham 194 (BM!).
Plants fleshy and soft, prostrate, mixed with other bryophytes or in dense pure mats,
grey-green, highly nitid when dry; axes to 1.8 mm. wide, with apices and intercalary
portions of axes very commonly enlarged, the axes often with a somewhat flattened
appearance in ventral view.
Branches frequent, of lateral-intercalary type (commonly from ventral end of mero-
phyte) and Frullania-0'^ (collections varying in predominance of each type), ventral-
intercalary vegetative branches not seen.
Stems 6-9 cells high, cortex undifferentiated or more rarely in a single row of slightly
larger cells, cortex and medulla with usually distinctly thick-walled cells, rarely with
them only slightly thickened; endophytic hyphae absent. Rhizoids in fascicles from a
raised, flat, disciform rhizoid initial field confluent with median portion of underleaf base.
Leaves with insertion moderately recurved at ventral end ; leaves strongly erect and axis
then appearing narrowly channeled in dorsal view, occasionally locally connivent, rarely
connivent throughout axis, occasionally moderately spreading, densely imbricate, conchi-
form concave, wide-ovate to ± oblate; apex rather narrowly to very broadly rounded
to ± truncate, undivided, in moist condition the apical portion of leaves incurved, in
dried condition the apical portion of at least some leaves of main axis strongly incurved,
the apices with a few teeth or (commonly) entire; dorsal margin =*= straight, often
sinuate, often in part reflexed, not or slightly decurrent, with few to several often sharp
teeth and/or blunt projections, occasionally entire; ventral margin entire, very broadly
FIG. 31. Clasmatocolea gayana (Mont.) Grolle. 1, Main axis with perianth, ventral
view. 2, Perianth with 3 keelar wings and a short secondary wing, cross section through
median portion. 3, Median leaf cells. 4, Leaves. 5, Underleaf segment, terminal portion
with collapsed slime papilla. 6, Teeth of dorsal margin of leaf. 7, Underleaves. 8, Main
axis showing leaf and underleaf insertion, ventral view; note raised rhizoid initial fields.
9, Stem, cross section. Figures 1, 3, 4a, 5-7a, from type of C. gayana, "Chile, hb.
Montagne" (FH); figure 2, from Engel 4554, Chile, Prov. Magallanes, Pto. Eden;
figures 4b, 7b, from Engel 4225, Chile, Prov. Chiloe, Pto. Ballena; figure 8, from Engel
4473, Chile, Prov. Magallanes, I. Williams; figure 9, from type of C. gayana, Chile,
Valdivia, Gay 44 (S).
113
114
FIELDIANA: BOTANY
FIG. 32. Clasmatocolea gayana (Mont.) Grolle. 1, Outer capsule wall cells, note
semiannular band. 2, Inner capsule wall cells. 3, Capsule wall, cross section. 4, Median
apical portion of innermost bract. 5, Perianth mouth. 6, Perianth mouth, ventral lobe
in middle. Figures 1-3, from Imshaug 42959, Chile, Prov. Osorno, near L. Toro; figure
4, from Engel 4229, Chile, Prov. Chiloe, Pto. Ballena; figures 5, 6, from Engel 4554,
Chile, Prov. Magallanes, Pto. Eden.
rounded, the basal portion often extending across stem and closely approaching but
not in contact with ventral margin of opposite leaf, the ventral margin entire.
Leaf cells thin-walled, trigones bulging to nodular, rarely confluent; median leaf
cells 18-34 p. wide; 20-38(-42) p. long; cuticle smooth or roughened and appearing finely
granular, sometimes a few cells with smooth cuticle scattered among cells with roughened
cuticle.
Underleaves 1.5-2. 2(-2.8) X stem width, connate on 1 side, transversely or obliquely
inserted, slightly spreading, imbricate, slightly to moderately convex (ventral view),
ovate to subrectangular, 0.3-0.5-bifid; segments occasionally incurved, often long
apiculate, often with a uniseriate row of several cells, the segments terminating in a
slime papilla, the segment margins entire; margins of the lamina with l-2(-3) small to
large teeth, rarely with 1 lacinium, the armature terminating in a slime papilla.
Plants dioecious; androecia terminal or intercalary on main axis or short to long
lateral-intercalary branches (a single ventral-intercalary cf branch seen); bracts with
basal portion strongly saccate, with apices appressed to bract immediately above;
lobule margin involute, with several slime papillae and often few-celled teeth, or 1
lacinium, or 1 rounded involute lobe; antheridia solitary, stalk uniseriate.
ENGEL: CLASMATOCOLEA
115
50
FIG. 33. Clasmatocolea gayana (Mont.) Grolle.
Gynoecia on main axis or long lateral-intercalary branches; subfioral innovations
when present (rare) originating from below bracts or bracteole of first or second series,
usually lateral (1 ventral innovation seen); bracts in 3 series, those of innermost series
usually free, rarely connate with keelar wing on 1 side, slightly concave, particularly in
upper portion, obovate; apices undivided, broadly rounded or truncate, occasionally
with 2 feebly developed, rounded lobes, the apex rather densely denticulate with uni-
116 FIELDIANA: BOTANY
seriate teeth; lamina margins sinuate, entire or sparingly denticulate in extreme upper
portion, often with short, blunt projections of varying sizes. Bracteoles of innermost
series subequal to bracts in size, free from bracts, concave or canaliculate (ventral view),
especially in upper portion, margins in upper portion occasionally connivent, the
bracteole =±= appressed to the infolded ventral lobe of perianth, obovate; apices un-
divided or very short-bifid, broadly rounded, rather densely denticulate; lamina margins
entire or sparingly denticulate in extreme upper portion, often with short, blunt pro-
jections of varying sizes. Perianth nearly always present, strongly trigonous, subclavate to
elongate-subrectangular to elliptic, the ventral side strongly infolded lending perianth ±
bilaterally compressed, the perianth straight or gradually narrowing toward the ± bilat-
erally compressed mouth; lobes broadly rounded or ± truncate, often with several
short, blunt projections, rather densely denticulate by teeth that are (at least for a con-
siderable portion distally) uniseriate; keels with wings commonly large and conspicuous,
often dentate, occasionally sinuate, secondary wings occasional, of a few cells high.
Seta observed only in old, collapsed state. Capsule valves 0.83-0.91 mm. long, 34-42 p
thick, of 4-5 layers, the outer row of cells equal to thickness of ca. 2.3-2.5 of interior
strata; outer layer with red-brown, wide nodule-like to wide spinelike thickenings that
are often feebly tangentially dilated, a few semiannular bands present; exposed wall
moderately 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 incomplete, sometimes
branched, the radial walls with nodulose thickenings often present.
Spores 21-24 p., red-brown, exine with narrowly conical or subrectangular projections
with dilated, flattened, or rounded tips and with radiating bases interdigitating with one
another, the intervening wall with weak nanogranules ; spores averaging 2.3 X elater
diameter. Elaters 10-12 /* wide, spiraled to tips, walls light brown.
Differentiation. — This species is one of the easiest of Clasmatocolea
taxa to identify. Dried plants can be identified at once by the following
ensemble of features: (a) the very commonly swollen apices and inter-
calary portions of axes; (b) the nearly strictly corticolous habit; (c) the
very common presence of perianths; (d) the strongly incurved apical
portion of leaves; and (e) the highly nitid plants. Upon wetting the plants
the following combination of features is distinctive : (a) the dentate apical
but more commonly dorsal leaf margins; (b) the underleaves that are
connate on one side; (c) the presence of a raised, flat, disciform rhizoid
initial field confluent with the median portion of the underleaf base;
(d) the poorly differentiated stems, with cortical and medullary cells
usually distinctly thick walled; (e) the rather densely denticulate bract
apices, with the teeth uniseriate; (f) the rather densely denticulate perianth
mouths, with the teeth uniseriate, at least for a considerable portion
distally; and (g) the large, conspicuous, occasionally sinuate wings of the
perianth keels.
Notes. — 1. According to Schuster (1966), the presence of a well-defined
rhizoid initial disc occurs primarily in Porellaceae, Frullaniaceae, and
Lejeuneaceae. In C. gayana, this feature is a specialized one and is asso-
ENGEL: CLASMATOCOLEA 117
elated with the corticolous habit of the species. Rhizoids may or may not
issue from the rhizoid initial field.
2. Stephani (1900) described Lophocolea concava and listed Dusen
material from Monte Tronador in Chile and Puerto Blest in Argentina,
but the specimens from these localities bear no relationship to Stephani's
description. Stephani (1906), in his treatment of L. concava in Species
Hepaticarum, recognized the error and in effect typified the material by
citing only Dusen material from Tierra del Fuego. Lophocolea concava is
a synonym of Leptoscyphus horizontalis (fide Grolle, 1962). The material
listed under L. concava in Stephani (1900) is actually C. gayana (fide
material in S!). See also the discussion in Grolle (1962, pp. 43-44).
3. Pearson (1922) described a new species, Conoscyphus flaccidus, based
upon a specimen collected in 1868 by Cunningham at Punta Arenas
(Sandy Point), Chile, and named by Stephani as Lophocolea humilis. The
outside of the packet of the holotype specimen (BM) neither bears the
name Conoscyphus flaccidus nor an indication that Pearson saw the col-
lection. However, within the packet there is a small blue slip of paper
labeled "Conoscyphus flaccidus n. sp. Pearson M.S."
Ecology. — A corticolous species, for the most part in forests, occurring
on Nothofagus, Laurel ia, Drimys, Saxegothaea, Araucaria, Weinmannia,
etc. Like C. ctenophylla it usually occurs where there is a minimal amount
of competition with other bryophytes and commonly does not occur on
trunks or branches covered with large, thick masses of bryophytes (which
commonly frequent the very wet, west portions of southern South Ameri-
ca). The species is sometimes intermixed with other hepatics such as
Lepicolea ochroleuca, Frullania spp., Metzgeria spp., Radula sp., Plagio-
chila spp., Porella subsquarrosa, and Clasmatocolea cucullistipula. This
species does not seem to be strictly altitudinally zoned, e.g., it occurs in
Prov. Aisen (6-150 m.), Prov. Chiloe (near sea level-ca. 100m.), Prov.
Osorno (195-920 m.), Prov. Valdivia (10-1,000 m.), Prov. Cautin (730-
800 m.), Prov. Malleco (330-1,280 m.).
Phytogeography. — Tierra del Fuego (sparse); Patagonian Channels; N.
in Valdivian region (in West Patagonia N. to 37° 46' S.; East Patagonia
in Prov. Chubut and Prov. Rio Negro). Hassel (1977) records the species
for South Georgia; I have been unable to confirm or refute this report
(see fig. 33).
Specimens seen.— CHILE. PROV. MAGALLANES: I. Ghebel, Spe-
gazzini 306 as syntype of L. vinciguerrana — c. per. (VER); I. Desolation,
Dusen s.n. (S); ibid., Pto. Angosto, Dusen 255 as Leioscyphus turgescens
(LD), as L. gayana (NY, S); Brunswick Pen. (cf. Engel, 1978); E. side of
B. Borja, Engel 6135, 6145 (MSC); I. Newton, Dusen 17 (NY); Pto.
118 FIELDIANA: BOTANY
Charrua, Engel 4795 (MSC); Pto. Eden, Engel 4554 - c. sporo. (MSC);
SE point of I. Williams, Engel 4473 (MSC). PROV. AISEN: Pen. de
Taitao, B. San Rafael, Gusinde s.n. — c. per. (JE); ibid., Gusinde s.n. as
L. puccioana — c. per. (S); R. Aisen, Dusen s.n. as L. humilis (LD — c.
sporo., O); ibid., Dusen 230 as L. humilis — c. per. (JE); ibid., Dusen 249
(NY — c. per., S — c. sporo.); Puerto Aisen, Santesson M 318 as L.
puccioana (S); 32 km. E. of Puerto Aisen, R. Aisen Valley, Santesson M
326 (S); ibid., Santesson M 327 as L. puccioana — c. per. (S); I. Magda-
lena, Schwabe 20/1 - c. per. (JE); ibid., 6 m., Schwabe 33 Ib (hb. Grolle);
Pto. Puyuhuapi, Schwabe 4/b — c. per., 8/a p.p., 8/e, 23/b, 23/d — c. per.
(JE); ibid., R. Obscuro, Schwabe 21 /a (hb. Grolle); L. Risopatron,
Schwabe 6/c (hb. Grolle). PROV. CHILOE: Pto. Ballena, Engel 4225,
4229 (MSC); I. Guaitecas, Dusen 357 as L. humilis (S - c. per., UPS);
ibid., Bo. Chica, Dusen s.n., 368 as L. humilis (S); Melinca, Dusen 357 as
L. humilis — c. per. (S); I. Chiloe, Chadmo Central, just N. of Puente
San Juan along Ruta 5, 20.9 km. by road N. of Quellon, ca. 50 m., Engel
11960 (F); S. of Cucao near W. coast of I. Chiloe, C. Pirulil, 50 m., Engel
12086 — c. per. (F); I. Chiloe, 1.3 km. by road N. of junction of Ruta 5
and road to Delcahue, ca. 100 m., Engel 12158 (F); I. Chiloe, Lechagua
area, 5 km. by road W. of Ancud, near sea level, Engel 11712 — c. per.
(F). PROV. LLANQUIHUE: Puerto Montt, Claude Joseph 3203 as L.
ctenophylla (MICH); ibid., Alerce, Santesson M 557 — c. per. (S); Puerto
Varas, Dusen 461 - c. per. (FH, S). PROV. OSORNO: Antillanca,
920 m., Ruthsatz 56/9 (hb. Grolle); Refugio Antillanca, 195 m., Engel
3910B (MSC); around L. Toro on road to Refugio Antillanca, 855 m.,
Engel 4023 (MSC); near small lake 10.3 km. by road below Refugio
Antillanca, 650 m., Engel 11550 — c. per. (F); Agua Caliente, R. Chan-
leufii, 4 km. from Termas de Puyehue along road to Refugio Antillanca,
400 m., Engel 11459 - c. per. (F); Termas de Puyehue, Schwabe 64 (JE).
PROV. OSORNO/PROV. VALDIVIA: Forest reserve at Planta Hydro-
electrica Pilmaiquen, along R. Pilmaiquen, 2 km. by road W. of Entre
Lagos, ca. 100 m., Engel 11597 (F). PROV. OSORNO near Prov. Valdivia
boundary: Anticura, Salto del Indio, 19 km. by road E. of Termas de
Puyehue along international highway, 300 m., Engel 11664 — c. per. (F).
PROV. VALDIVIA: W. slope of Cord. Pelada, W. of El Mirador,
between La Union and Pta. Hueicolla, 580-840 m., Engel 12273 — c. per.,
12318 - c. per., 72345 - c. per. (F); E. slope of Cord. Pelada, E. of
El Mirador, between La Union and Pta. Hueicolla, 740-840 m., Engel
12422 — c. per., 12454 (F); Cord. Pelada, summit of El Mirador, between
La Union and Pta. Hueicolla, 1,000 m., Engel 12374 (F); Los Ulmos,
Mahu 7780 as C. humilis (hb. Grolle — c. per., hb. Mahu); E. slope near
ENGEL: CLASMATOCOLEA 119
R. Futa in vicinity of Futa, 10.5 km. by road S. of junction of highway
T-60 and T-65, 10 m., Engel 11019A (F); V. Shoshuenco, Ruthsatz s.n.
(JE); Corral, Dusen 91 — c. per. (NY); ibid., Krause s.n. — c. per. (S);
ibid. [Quitaluto], Hosseus s.n. — c. per. (PC); ibid., Hosseus 657, 657 B,
666, 667, 679 ex p. — c. per. (JE); ibid., Hosseus 662 — c. per., 668 p.p. -
c. per. (S); ibid., 430 m., Hosseus 641, 643A (JE); L. Rinihue, Enco,
Santesson M 673 — c. sporo., M 677 — c. per., M 679 — c. per., M 680 -
c. per., M 682, M 686 A — c. per., M 688 — c. sporo. as L. puccioana (S);
Valdivia, Herzog s.n. as L. puccioana (JE); Arique, [? Lechler] (FH — c.
per., NY, S). PROV. CAUTIN: Termas de Palguin, R. Palguin, 730 m.,
Engel 11334 — c. per., 11335 — c. per. (F); Parque Nac. Villarrica, N.
slope of V. Villarrica, 800 m., Engel 11202 - c. per., 772/2 - c. per. (F);
Pucon, V. Villarrica, Hosseus 209 — c. per. (JE); ibid., Hosseus 2 10 A (S).
PROV. MALLECO: Cord. Nahuelbuta, Parque Nac. Contulmo, 7 km.
by road E. of Contulmo, 330-360 m., Engel 12537 (F). PROV. MALLECO/
PROV. ARAUCO: Cord. Nahuelbuta, Parque Nac. Nahuelbuta, 48 km.
by road W. of Angol, 1,280 m., Engel 12598 (F). ARGENTINA. PROV.
CHUBUT: L. Puelo, sin. coll. lib (JE). PROV. RIO NEGRO: M.
Tronador, Dusen 577 as syntype of Lophocolea concava — c. sporo. (S);
Puerto Blest, Dusen 505 as syntype of L. concava — c. per. (S).
Clasmatocolea ctenophylla (Schiffn.) Grolle. Figures 34-35, Plates 15-16.
Lophocolea ctenophylla Schiffn. in Naumann, Forschungsr. Gazelle 4
(4): 12. pi. 3, f. 25-28. 1889. Clasmatocolea ctenophylla (Schiffn.)
Grolle, Rev. Bryol. Lichenol. 29:71. 1960. Holotype: Chile, Prov.
Magallanes, I. Desolation, B. Tuesday, 2 February 1876, Naumann s.
n. (FH! - c. per.).
Lophocolea cristato-spinosa Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 37. 1900, syn. nov. Original material: Chile, Prov.
Llanquihue/Prov. Osorno, L. Llanquihue, Dusen (BM!, FH! — c.
per., LD! — c. sporo., NY! — c. per.).
Plants delicate, prostrate, mixed with other bryophytes or pure in loose or dense,
compact mats, light green to pale grey-green, 315-490 /x wide.
Branches frequent, sometimes copiously developed, commonly of lateral-intercalary
and Frullania types, the former particularly common, ventral-intercalary branches
occasional.
Stems (4-)5-6(-7) cells high, cortex in 1 row of distinctly thick-walled cells larger than
medullary cells; medullary cell walls distinctly thickened; endophytic hyphae present in
both cortex and medulla or absent. Rhizoids in fascicles from stem near underleaf base,
long for plant size.
Leaves with insertion very slightly recurved at ventral end; leaves strongly erect,
the teeth of apical-dorsal portion intermcshing with those of opposite leaf; leaves densely
120
ENGEL: CLASMATOCOLEA 121
imbricate, conchiform concave, suboblate to wide-ovate; apex broadly rounded, un-
divided, often somewhat incurved ; apex and margins with 17-32 regularly spaced l-2(-4)-
celled sharp teeth that are ± equal in size; teeth with the tip cell usually thick-walled,
the tip cells of teeth toward base of dorsal margin often more elongated than other teeth ;
dorsal margin straight, entire, or sparsely toothed in basal portion; ventral margin
broadly rounded.
Leaf cells thin to slightly thick-walled, trigones large and slightly to distinctly bulging,
rarely medium; median leaf cells 17-26/i wide, 18-26(-30)M long; cuticle smooth,
occasionally with some cells with smooth cuticle scattered among cells with roughened
cuticle. Oil-bodies in all leaf cells except ultimate cells of teeth, occupying very small
fraction of cell lumen, very light grey, of grape-cluster type, with numerous, small
globules, the oil-body surface appearing finely botryoidal; oil-bodies variable: globose
to ovoid to elliptic to crescentic in shape, consistently 2 per cell of middle, apex, or
base of leaf, those of leaf middle 4-5 /i wide, 4-6 /* long.
Underleaves 1.4-2.1 X stem width, free, oppressed, slightly convex (ventral view),
ovate to subrectangular, bifid to 0.20-0.33; segments often apiculate, sometimes termi-
nating in a slime papilla, the segment margins entire or with 1-2 sharp, thick-walled
teeth near base of inner and/or outer margin ; margins of the lamina with several com-
monly 1-2-celled thick-walled teeth that sometimes terminate in a slime papilla, the
teeth occasionally somewhat larger, the margins rarely with only a few slime papillae.
Plants apparently dioecious; androecia not seen.
Gynoecia on main axis or short to rather long, lateral-intercalary branches ; subfloral
innovations absent or from below bract of first or second series; vestigial stem peri-
gynium present; bracts in 3 series, the bracts and bracteole of innermost and second
series inserted on stem perigynium; bracts of innermost series plane or slightly concave,
obovate, insertion rather narrow; apices undivided, broadly rounded to truncate,
apex and margins of distal portion of lamina with numerous l-2(-3)-celled thick-walled
sharp teeth, the teeth subequal in size; bracts entire toward base. Bracteoles of innermost
series subequal to bracts in size, free from bracts; bracteoles canaliculate, usually
appressed to the infolded ventral lobe of perianth, obovate to ± elliptic; the insertion
sometimes rather narrow; apices variable: ± truncate, or narrowly rounded, or with 1
broadly triangular lobe, or with 2 small lobes and then retuse, the apex (plus lobes
when present) and margins of distal portion of lamina with teeth as in bracts; bracteoles
entire toward base. Perianth strongly trigonous, subrectangular to narrowly ovate, the
ventral side convex or infolded, the distal portion of perianth =±= bilaterally compressed,
Opposite:
FIG. 34. Clasmatocolea ctenophylla (Schiffn.) Grolle. 1, Portion of main axis, lateral-
dorsal view. 2-6, Underleaves. 7, Teeth of apex and upper portion of ventral margin
of leaf. 8, Underleaf. 9, Leaf (flattened). 10, Portion of underleaf apex, tip a nearly
collapsed slime papilla. 11, Seta, cross section. 12, Median leaf cells. 13, Outer capsule
wall cells. 14, Stem, cross section. 15, Perianth mouth. 16, Perianth mouth, ventral
lobe in middle. 17, Capsule wall, cross section. 18, Inner capsule wall cells with bands
only sporadically represented. Figures 1-3, 7, 9, 10, 12, 14, 15, from holotype of L.
ctenophylla, Naumann, Chile, Prov. Magallanes, B. Tuesday; figures 4, 11, 13, 16, from
Engel 4714, Chile, Prov. Magallanes, I. Grant; figures 5, 8, from Engel 4021 A, Chile,
Prov. Osorno, L. Toro; figure 6, from Engel 4240, Chile, Prov. Chiloe, Pto. Ballena;
figures 17, 18, from Engel 5213, Chile, Prov. Magallanes. I. Madre de Dios.
•••
FIG. 35. Clasmatocolea ctenophylla (Schiffn.) Grolle. O = without specific location.
122
ENGEL: CLASMATOCOLEA 123
narrowing slightly in upper portion but expanding at immediate apex; perianth mouth =*=
bilaterally compressed, the lobes broadly rounded, with numerous, 1-2-celled, thick-
walled sharp teeth that are =*= equal in size; keels often with large, conspicuous, often
very expanded, sometimes in part auriculate, wings with numerous small teeth.
Seta 5-6 cells in diameter, with 18 rows of outer thin-walled cells surrounding an
inner core of scattered smaller cells with small trigone-like thickenings. Capsule valves
0.56-0.70 mm. long, 30-47 /i thick, of 3-4(-5) layers, the outer row of cells equal to
thickness of 1.8-3 of interior strata; outer layer with red-brown, wide nodule-like or
wide spinelike thickenings that are often weakly to strongly tangentially dilated,
complete and incomplete semiannular bands common; exposed wall thin to moder-
ately thickened; intermediate and inner layers of cells subequal in thickness, the inter-
mediate layer(s) with thickenings often considerably extending onto tangential walls;
inner layer of cells with red-brown semiannular bands very common on some valves,
sporadic on others, often incomplete, the radial walls with nodulose thickenings very
common on some valves, moderately represented on others.
Spores (II-) 12- 15 /*, light brown, exine with =*= hyaline, narrowly conical, rather
widely spaced projections with often weakly dilated, often =*= clawlike tips and with
radiating bases, the intervening wall has weak nanogranules; spores averaging 1.6 X
elater diameter. Elaters (6-)7-9 n wide, spiraled to tips or with apical portions with
thick nonspiraled walls, elater walls red-brown.
Differentiation. — This small species should, even in the dried condition,
offer no confusion with any other member of the genus. The following
features will serve to identify it: (a) the corticolous habit; (b) the copious
production of regularly shaped, ± equally sized, sharp, small teeth on the
leaf margins; and (c) the small axis size (315-490 n wide). The regular
dentition and small axis stature lend a beautiful, intricate appearance to
the plants. Clasmatocolea ctenophylla has spore ornamentation similar to
two other corticolous taxa in the genus — C. cucullistipula and C. gayana —
but the spores of C. ctenophylla differ in being ca. one-half the size.
Note. — At least one series of bracts and bracteoles of C. ctenophylla is
inserted on a vestigial stem perigynium and in this species the phenomenon
appears to be a response to fertilization.
Ecology. — A species of both deciduous and evergreen forests where
corticolous on a variety of substrates — Nothofagus pumilio, Drimys,
Podocarpus, etc., and occasionally on rotted logs and stumps. In the
Valdivian region (near sea level- 1,225 m.), it sometimes grows intermixed
with Xenocephalozia navicularis, a species, in general, with a similar
ecology. Clasmatocolea ctenophylla usually occurs where there is a mini-
mal amount of competition with other bryophytes and commonly does
not occur on those tree trunks that, in the range of this species, are so
frequently covered with large, thick masses of bryophytes.
Phytogeography. — Tierra del Fuego (I. Grande de Tierra del Fuego and
I. Desolacion); Patagonian Channels; N. in Valdivian region to 39°
52' S. (Corral, Prov. Valdivia) (see fig. 35).
124 FIELDIANA: BOTANY
Specimens seen— CHILE. PROV. MAGALLANES: Ens. Villarino,
Hatcher 52b (NY); I. Desolation, Dusen s.n. (FH); ibid., Pto. Angusto,
Dusen 417 as L. gayana (S); I. Desolation, Pto. Churruca, head of Br.
Lobo, Engel 5813 - c. per. (MSC); I. Desolation, B. Tuesday, head of
inner harbor, Engel 5701 (MSC); S. side of I. Madre de Dios, N. of I.
Tarlton, Engel 5213 - c. sporo., 5214 - c. per. (MSC); W. side of I.
Grant, Engel 4714 - c. sporo. (MSC). PROV. AISEN: I. Magdalena,
Schwabe 33/b 2 p.p. asL. multispinula (JE). PROV. CHILOE: Pto. Ballena,
Engel 4240 (MSC); I. Guaitecas, Melinca, Dusen 417 as L. gayana (NY);
I. Chiloe, Cord. San Pedro, Butalcura, near R. Butalcura, 1 1 km. by road
from Ruta 5, 100 m., Engel 11831 (F); I. Chiloe, Aguas Buenas area,
4.7 km. E. along Aguas Buenas road from Ancud-Quemchi road, ca.
100 m., Engel 12212 (F); I. Chiloe, Chepu, S. side of R. Chepu, near sea
level, Engel 11735, 11764 (F). PROV. OSORNO: Antillanca, (?) 1,225 m.,
Ruthsatz 58/4 (hb. Grolle); around L. Toro on road to Refugio Antillanca,
855 m., Engel 4021 A (MSC); 12.1 km. by road below Refugio Antillanca,
550 m., Engel 11 543 A (F). PROV. VALDIVIA: W. slope of Cord.
Pelada, W. of El Mirador, between La Union and Pta. Hueicolla, 580 m.,
Engel 12288 — c. per. (F); E. slope of Cord. Pelada, E. of El Mirador,
between La Union and Pta. Hueicolla, 840 m., Engel 12407 (F); Corral
[Quitaluto], Hosseus 116 (hb. Grolle, JE).
Clasmatocolea crassiretis (Herz.) Grolle. Figures 36-37.
Lophocolea crassiretis Herz. Trans. & Proc. Roy. Soc. New Zealand
65:354. 1935. Clasmatocolea crassiretis (Herz.) Grolle, Nova Acta
Leop. 25 (161): 69. 1962. Holotype: New Zealand, North Island, Te
Matawai, Tararuas, 3 June 1932, Zotov 135 (JE! — c. per.; isotypes:
CHR!, JE!).
Plants rigid, prostrate, in nearly pure mats or mixed with other bryophytes, yellow
brown to light brown, highly nitid when dry, submoniliform (especially in ventral view);
axes 0.5-0.6 mm. wide.
Branches sporadic, lateral-intercalary type common, from near ventral end of mero-
phyte, ventral-intercalary type sporadic, terminal branching absent.
Stems slender for plant size, rigid, wiry, frequently arced or wavy, commonly not
growing straight, 5-6 cells high, cortex in 1 layer of thick-walled cells larger than the
medullary cells, the outer tangential wall very thickened; medullary cell walls thick;
endophytic hyphae present in both cortex and medulla. Rhizoids occasionally present,
in tight fascicles from stem at underleaf bases.
Leaves rather fragile, the insertion slightly recurved at ventral end; leaves strongly
erect, opposing leaves connivent or closely approaching, the axis appearing ± narrowly
channeled in dorsal view, the leaves imbricate, deeply conchiform concave, appearing
inflated and billowed out, orbicular to oblate, the apex broadly rounded, undivided,
the apex and margins entire; dorsal margin =±= straight in proximal portion, the distal
FIG. 36. Clasmatocolea crassiretis (Herz.) Grolle. 1, Axis, dorsal-lateral view. 2, Por-
tion of axis, lateral view. 3, Perianth bearing axis, lateral view. 4, 5, Adaxial and abaxial
views of same leaf. 6, Apex of gynoecial shoot, ventral view. 7, Portion of main axis,
ventral view. 8, Antheridial stalk. 9, Median leaf cells. 10, Stem, cross section. 11-18,
Underleaves, in situ. Figures 1-5, 7, 9-16, 18, from holotype, Zotov 135, New Zealand,
North I., Tararuas; figure 6, from isotype (JE); figure 8, from Druce s.n., New Zealand,
North I., Maungapohatu; figure 17, from Scott s.n., New Zealand, Stewart I., Mt.
Anglem Track.
125
FIG. 37. Clasmatocolea crassiretis (Herz.) Grolle.
126
ENGEL: CLASMATOCOLEA 127
portion broadly rounded, the margin slightly undulate, decurrent; ventral margin
broadly rounded, the basal portion subauriculate and conspicuously extending ventrally
beyond stem.
Leaf cells with massive, nodular, or maximally protuberant, irregularly triangular to
rectangular trigones that are confluent or separated by narrow thin-walled places, the
cell lumen bounded mostly by the massive trigones; median leaf cells 1 3-20(-24) /* wide,
(16-) 18-26 M long; cuticle smooth.
Under/eaves rigid, 0.9-1.3 X stem width, often asymmetrically oriented, free, strongly
spreading, sometimes diverging from axis at 90°, distant, sometimes partially obscured
by ventral portion of adjacent leaf, plane, cuneate to suborbicular to oblate to ovate;
apices undivided, truncate to emarginate, commonly with a pair of slime papillae;
margins of the lamina entire except for an occasional slime papilla.
Plants dioecious; androecia intercalary on main axis, bracts densely imbricate and
with apices usually appressed to bract immediately above, the bracts considerably
smaller than leaves, deeply concave and strongly saccate at the base; lobule abbreviated,
the margin slightly inflexed, broadly rounded, entire; antheridia solitary, stalk uni-
seriate.
Gynoecia on leading axes or short to long lateral-intercalary branches; subfloral in-
novations absent; bracts in 2-3 series, the bracts of innermost series with insertion
rather narrow, the bracts erect and appressed (or nearly so) to the perianth, deeply
conchiform concave, rotundate to ovate, apices and margins broadly rounded, entire.
Bracteoles of innermost series free from bracts, plane, partially obscured by bracts,
cuneate to elliptic, apices broadly rounded (and sometimes repand) to shallowly emar-
ginate to bidentate, entire. Perianth inflated, terete, cylindrical to clavate to elliptical
(young perianths frequently campanulate), the sides slightly narrowing or straight or
moderately expanding toward the mouth, the mouth wide and open or partially closed
by incurving of some or all of lobes; lobes free for 0.15-0.25 perianth length, the dorsal
lobes broadly rounded, occasionally incurved, entire, the ventral lobe distinctly smaller,
narrowly rounded, incurved, entire; keels absent, wings usually absent (only one seen).
Sporophyte not seen.
Differentiation. — This species is quite comparable to C. strongylophylla
(C. turgescens), which also occurs in New Zealand, in (a) general aspect
and small stature of the plants; (b) underleaf shape variability; (c) general
shape of the inflated perianths, including the smaller, incurved ventral
lobe; and (d) the broadly rounded, slightly incurved and entire male bract
lobules. In fact, these characters serve to indicate that sections Clasmato-
colea and Strongylophyllae are allied and related through C. strongylo-
phylla and C. crassiretis.
Clasmatocolea crassiretis and C. strongylophylla should, however, not
be confused and may be separated as follows:
Character C. crassiretis C. strongylophylla
Leaf cuticle Smooth Coarsely papillate
Leaf trigones Massive, protuberent Large to bulging
Underleaves Strongly and stiffly Appressed to moderately
spreading, undivided spreading, commonly divided
128
FIELDIANA: BOTANY
Character C. crassiretis
Branching Strictly intercalary
Perianths Usually smooth, not winged
C. strongylophylla
Mostly terminal, occasionally
intercalary
Winged
The closest ally of C. crassiretis appears to be C. humilis var. humilis.
The two are separable as follows (note that several of the characters are
qualitative) :
Character C. crassiretis
Underleaves To 1.3 X stem width, free, plane
and not convex (ventral view),
undivided, never bifid
Branching Terminal absent, mostly lateral-
intercalary
Leaf cells Trigones massive, protuberant;
leaf cell lumen bounded
mostly by the massive trigones
Stem 5-6 cells high
Perianth Inflated, terete, the ventral side
not infolded; wings rare
Plant size Smaller, to 0.6 mm. wide
Habitat Often corticolous
C. humilis var. humilis
To 4.3 X stem width, free or
connate on 1 side, slightly to
deeply convex (ventral view),
sometimes cupulate, bifid or
undivided
Mostly ventral- and lateral-
intercalary, terminal occasional
Trigones large to bulging to
knotlike; leaf cell lumen
bounded mostly by cell wall
8-14 cells high
Strongly trigonous, the ventral
side usually strongly infolded;
wings common
Larger, to 1.8 mm. wide
Terricolous, saxicolous, or on
rotted logs
Ecology-Phytogeography. — Usually corticolous in Olearia scrub or wet,
mossy, Dracophyllum biforme-Olearia forests where associated with Pla-
giochila pleurota and Goebeliella cornigera; very rarely on ground. Known
from Macquarie Is.; Stewart I. (535-790 m.) and North I. (885-1,530 m.),
New Zealand (see fig. 37).
Specimens seen.— MACQUARIE IS.: Ashton s.n. (hb. Grolle). NEW
ZEALAND. STEWART I.: Mt. Anglem Track, 790 m., Mork s.n. as C.
turgescens (CHR); ibid., 535 m., Scott as C. turgescens — c. per. (hb.
Child, F); Mt. Anglem Track, Scott as C. turgescens (CHR). NORTH
ISLAND. WELLINGTON: Schormann's Track, E. slopes of Mt. Mines,
N. Tararua Mts., 10-12 mi. W. of Eketahuna, 885-1,005 m., Schuster
49174a - c. per. (hb. Schuster); ibid., Schuster 49160, 49 167 A (F, hb.
Schuster). GISBORNE: Maungapohatu, Urewera, 1,530m., Druce s.n. —
c. d" (hb. Grolle).
Clasmatocolea moniliformis Engel. Figures 38, 39.
Clasmatocolea moniliformis Engel, Phytologia 41:310. 1979. Holotype:
Chile, Prov. Magallanes, E. side of B. Borja, Engel 61 59 (MSC!).
cts
FIG. 38. Clasmatocolea moniliformis Engel. 1, Main axis, lateral view. 2, Axis, dorsal
view, with apical pair of leaves removed to show underleaves. 3, Axis, lateral view,
with 2 basal leaves removed on 1 side. 4, Portion of axis showing leaf and underleaf
insertion, ventral view. 5, Leaf and underleaf, lateral view. 6, Axis, cross section (1 =
leaves; ul = underleaves; s = stem). 7-9, Leaves; note ventral-basal subauriculate
projections. 10, Stem, cross section. 11, 12, Stalked slime papillae of underleaf apex. 13,
14, Underleaves, lateral and ventral views. 15, Median leaf cells. Figures 1-5, 9-15, from
holotype; figures 6, 8, from Engel 6231, Chile, Prov. Magallanes, B. Borja; figure 7,
from Engel 5677, Chile, Prov. Magallanes, B. Tuesday.
129
130
ENGEL: CLASMATOCOLEA 131
Plants rigid, fragile, prostrate, in nearly pure mats or more often mixed with other
bryophytes, golden brown, nitid when dry, capillaceous, distinctly moniliform; axes 315-
420 M wide.
Branches rather frequent, nearly exclusively of ventral-intercalary type, lateral-
intercalary type absent, Frullania-type very rare (only 1 branch seen).
Stems brittle, frequently arced or wavy, commonly not growing straight, 6-9 cells
high, with little internal differentiation, the cortex in 1-2 rows of thick-walled cells
slightly larger than, about the same size as, or slightly smaller than the medullary cells,
the outer tangential wall very thickened; medullary cell walls thick. Rhizoids not seen.
Leaves usually with marginal 1-3 hyaline cell rows, insertion not recurved at ventral
end, the ventral end of leaf inserted on stem at immediate base of underleaf or on basal
few cells of underleaf lamina; leaves erect but not connivent, the axis appearing chan-
neled in dorsal view, the leaves imbricate, distinctly concave, producing a sharp, elon-
gate bulge in lateral view, the distal portion of the leaf moderately incurved ; leaves sub-
squarrose, with the dorsal portion of the otherwise obliquely truncate, undivided apex
rather narrowly rounded, the sides of the leaf =*= parallel, the margins and apices entire;
dorsal margin ± straight, plane, long decurrent; ventral margin ± straight, considerably
shorter than the dorsal margin, the extreme base of ventral margin slightly broadened
and forming a subauriculate projection that sharply narrows toward the insertion, the
ventral-basal portion of leaf sheathing marginal parts of the underleaf base.
Leaf cells distinctly thick walled, trigones large, often bulging and often confluent;
median leaf cells 11-14(-17)M wide, 14-22 M long; basal cells very thick walled and
more deeply pigmented; cuticle smooth. Oil-bodies throughout leaf except in marginal
1-2 rows, obscuring ca. 0.20-0.75 of cell lumen, very light grey or off-white, of grape-
cluster type, the oil-body surface appearing coarsely granular; oil-bodies subglobose to
ovoid to elliptic in shape, occasionally somewhat irregular, 2-3 per cell of leaf middle
and base, l-2(-3) of leaf apex, those of leaf middle 3-5 /* wide, (3-)5-6(-8) M long; oil-
bodies of underleaf l-3(-4) per cell, of same size as those of leaf middle.
Under leaves 4.6-6.7 X stem width, free or base of underleaf lamina connate with leaf
on 1 side, the under leaves with apical portion oppressed to stem; underleaves approximate
to contiguous, cucullate, inflated, billowed out, frequently containing miscellaneous de-
bris, clasping stem, the margins curved dorsally and extending above dorsal stem surface;
apices undivided, broadly rounded or truncate, slightly reflexed, entire, but with a few,
commonly short-stalked slime papillae; margins of the lamina entire except for an
occasional slime papilla.
Androecia and gynoecia not seen.
Differentiation. — This species stands quite isolated in Sect. Clasmato-
colea, because of the following ensemble of features :
1) The leaf insertion does not recurve at the ventral end; the ventral
ends of a pair of opposing leaves are inserted on the stem in very close
proximity to the underleaf or may sometimes be inserted, on one side, on
the basal few cells of the underleaf lamina. Because of this kind of leaf
insertion, coupled with the ventral-basal subauriculate projections of the
leaves, the basal-lateral portion of the underleaves are sheathed on both
sides by leaf tissue.
2) Branching in C. moniliformis is unique to the section, for in this
132 FIELDIANA: BOTANY
species branches are nearly exclusively ventral-intercalary (only a single
Frullania-type branch was seen, and lateral-intercalary branches are
absent).
3) The cucullate underleaves represent an extreme condition within the
section, being only somewhat approached by some forms of C. humilis. In
such plants of C. humilis, however, the underleaf apices are not closely
appressed to the stem and are less than 4.8 X the stem width.
4) The distinctly moniliform appearance of the axes is unique to the
section.
Of the members of Sect. Clasmatocolea, the closest relative appears to
be C. humilis. Differences in axis size, branching modalities, leaf insertion,
and underleaves will distinguish these taxa.
Clasmatocolea moniliformis shows some relationships to the New
Zealand C. crassiretis, which also consists of small, stiff plants with
brownish pigments and strictly intercalary branching. Differences in
underleaves and branching modalities will easily separate these plants.
Clasmatocolea moniliformis may possibly be confused with the rather
distantly related C. cucullistipula, which has the same general stature and
appearance. The two taxa are separable as follows:
Character C. cucullistipula C. moniliformis
Leaf margins 1-4-dentate Entire
Underleaf margin With one large lobe Entire
Underleaf base Subauriculate Narrowing toward the base;
not auriculate
Branching Frullania-type predominant; Ventral intercalary;
ventral-intercalary Frullania-type very rare
occasional
Color Light olive green-light brown Golden brown
Habit Corticolous Saxicolous
Ecology-Phytogeography. — Only in the cold, very wet southernmost
channel region (S. of ca. 53° 30' S. in Prov. Magallanes), in both forests
as well as exposed outcrops and moorland areas. The species is saxicolous,
especially where there has been an accumulation of soil and where a layer
of bryophytes is present. It is frequently mixed with other hepatics such
as Cryptochila sp., Grollea, Krunodiplophyllum squarrosum, Metahygro-
biella sp., Pachyglossa sp., Pleurocladopsis simulans, and Riccardia sp.
(see fig. 39).
Specimens seen.— CHILE. PROV. MAGALLANES: I. Desolaci6n,
B. Tuesday, head of inner harbor, Engel 5677, 5778C (MSC); E. side of
B. Borja, Engel 6188, 6216C, 6231 (MSC).
ENGEL: CLASMATOCOLEA 133
Clasmatocolea minutiretis Engel & Grolle. Figures 40-42, Plates 17, 18.
Clasmatocolea minutiretis Engel & Grolle in Engel, Phytologia 41 : 309.
1979. Holotype: Chile, Prov. Magallanes, E. shore of I. Pilot, Engel
4785 - c. sporo. + <? (MSC!).
Plants rather soft, but with stiff, wirelike stems, suberect, the plants in dense, pure,
compact mats, pale brown to pale reddish brown, the stems a rich red-brown, the leaves
pale red-brown above, but with a well-defined strip of red-brown pigmentation near
the insertion, in ventral view the leaves with a small, local, concentrated patch of red-
brown pigment at the ventral-basal point of insertion, the leaves often mottled with
patches of red-brown ; axes to 1 .4 mm. wide, often with a somewhat inflated appearance
in ventral view, the axes often with several separated, intercalary, swollen and/or dimin-
utive areas; axis apices becoming long attenuated, with reduced but imbricate leaves,
the apices sometimes becoming subflagelliform.
Branches abundant, sometimes arising from a mass of entangled, old, dead, leafless
axes, the ventral-intercalary type very common, lateral-intercalary occasional, terminal
branches absent.
Stems 7-8 cells high, cortex very thick-walled, not markedly differentiated, in a single
row of red-brown pigmented cells with walls slightly thicker than those of the medul-
lary; medullary cells very thick-walled. Rhizoids sporadically developed, in dense
fascicles from a rather wide field at the underleaf bases.
Leaves with insertion oblique, very slightly recurved at ventral end; leaves strongly
erect and connivent or nearly so on robust axes, those on less well-developed shoots
either suberect but not connivent (and then with the axis appearing narrowly channeled
in dorsal view) or with the leaves moderately to distinctly spreading; leaves rather
closely imbricate, deeply conchiform concave, the ventral-basal portion appearing
billowed out and extending ventrally well beyond stem surface, the ventral-basal portion
often extending laterally and overlapping the ventral-basal region of opposing leaf and
thereby often partially or totally obscuring view of both stem and underleaves; leaves
orbicular to wide-ovate to oblate; apex often slightly undulate, broadly rounded to
subtruncate, undivided, the apex and ventral margin with several irregularly sized and
spaced, often incurved teeth along with low, rounded projections, the dorsal margin
entire; dorsal margin ± straight proximally, moderately rounded distally, the two areas
often separated by a rather sharp, humplike projection that generally occurs in the
median portion of the margin, the dorsal margin plane or slightly undulate, short
decurrent; ventral margin broadly rounded, incurved toward the base and thus accen-
tuating the sharply defined leaf concavity.
Leaf cells somewhat thick-walled, trigones large to knotlike; median leaf cells 14-24 /*
wide, 19-30/j long; cuticle smooth.
Underleaves reduced and rather inconspicuous but distinct and persistent, on robust
axes frequently partially or wholly hidden by ventral portion of adjacent leaves, 0.35-
0.6 X stem width, free, but in close proximity to leaf on one side; underleaves loosely
oppressed or more often slightly spreading, distant, plane, or slightly convex (ventral
view), narrowly lanceolate to narrowly long-triangular, bifid to 0.55-0.7, but in less robust
axes with lamina of only 1-2 cells high and then underleaves divided nearly to base;
segments cilia-like, straight or flexuous, with a uniseriate portion of up to 7(-9) cells
originating from a base 2 cells wide, the segments sometimes uniseriate throughout;
FIG. 40. Clasmatocolea minutiretis Engel & Grolle. 1, Portion of axis, dorsal view.
2-5, Leaves and within, underleaves. 6, Underleaf. 7, Stem, cross section. 8, Portion
of axis, lateral view; note underleaf (UL), and saccate ventral portion of opposing
leaves (OL) toward axis apex. 9, Median leaf cells. 10, Portion of axis, ventral view.
All from holotype.
134
FIG. 41. Clasmatocoiea minutiretis Engel & Grolle. 1, Gynoecium, ventral view.
2, Perianth mouth, ventral lobe in middle. 3, Seta, cross section. 4, Inner capsule wall
cells. 5, Outer capsule wall cells. 6, Diagrammatic representation of branching pattern.
7, Capsule wall, cross section. 8, Antheridial stalk. 9, Opposing antheridial bracts with
stem (S), cross section. 10, Male bract. All from holotype.
135
FIG. 42. Clasmatocolea minutiretis Engel & Grolle. Plus Tristan da Cunha.
136
ENGEL: CLASMATOCOLEA 137
segments terminating in a slime papilla, the segment margins entire; margins of the
lamina entire or with 1 short or long and ciliform tooth.
Plants dioecious; androecia terminal or intercalary on leading axes; bracts obliter-
ating or nearly so the view of dorsal stem surface, the bracts with basal portion strongly
saccate, the saccate portion fused toward base with opposite bract, the distal portion
strongly concave, with apices appressed to bract immediately above, the lobe margins
with several teeth; lobule margin incurved, irregularly broadly rounded, with a few low,
rounded projections and slime papillae; antheridia solitary, stalk uniseriate.
Gynoecia on leading axes or rather short branches as well as subfloral innovations;
vestigial stem perigynium present; bracts and bracteoles in 3 series, those of the inner-
most series inserted on the stem perigynium; bracts of innermost series plane or slightly
concave, orbicular to obovate; apices broadly rounded, undulate, with several coarse
teeth and laciniae, the armature often sharply incurved; dorsal margin entire proximally,
repand or more often armed with a few teeth and laciniae distally, the margin straight
or arcing toward the perianth, plane or becoming undulate distally in area of armature;
ventral margin much like that of the dorsal. Bracteoles of innermost series free from
bracts, rather strongly concave (ventral view), narrowly to broadly obovate, sometimes
orbicular; apices bifid to ca. 0.25, less often emarginate, the segments occasionally
canaliculate; lamina margins entire in proximal portion, the distal portion armed with
a few, often recurved teeth or laciniae, the margins sharply recurved (curved ventrally),
sometimes undulate distally. Perianth strongly trigonous, elongate-subrectangular to
narrowly subclavate, not or barely narrowing toward the small or ± closed mouth;
lobes broadly rounded, sometimes incurved, the ventral lobe and 1 or both side lobes in-
folded, the lobes coarsely armed with sharp or rounded teeth; keels occasionally with
wings of a few cells high.
Seta 7-8 cells in diameter, with 19-20 rows of outer cells with corners thickened
similar to medium-large trigones; inner core cells scattered, smaller than outer row,
with corners thickened similar to small to medium trigones. Capsule elliptic, the valves
0.75-0.84 mm. long, 30-34 p thick, of 3-5 layers, the outer layer of cells exceeding
thickness of all interior strata combined; outer layer with red-brown, nodule-like or
weakly spinelike thickenings that are often feebly tangentially dilated, a few semi-
annular bands present; exposed wall thickened; intermediate and inner layers of cells
subequal in thickness, the intermediate layers with thickenings, often feebly tangentially
dilated; inner layer of cells with red-brown semiannular bands common, the bands
often incomplete, the radial walls with nodular thickenings very common.
Spores 12-1 3 /a, light brown, spherical or subspherical, exine with the light micro-
scope appearing minutely punctuate, but under the SEM with granulate to short, irregu-
lar, wide vermiform projections that have nanogranules, the intervening hollows with
nanogranules; spores averaging 1.2 X elater diameter. Elaters 8-10/* wide, apical por-
tions occasionally with thick nonspiral walls; elater walls red-brown, 2-3 /* in diameter.
Differentiation. — Clasmatocolea minutiretis bears definite affinities to
C. humilis, yet must remain an isolated element within Sect. Clasmatocolea.
With C. humilis, C. minutiretis shares such features as leaf form, cell size,
and trigone characters, predominance of intercalary branching (but in C.
minutiretis, branching is strictly intercalary, just as in C. crassiretis and,
practically, C. moniliformis of the same section), and spore size and
138 FIELDIANA: BOTANY
surface features. Two features, however, rather isolate C. minutiretis in
the section.
First, underleaves represent an extreme form of reduction, being quite
inconspicuous and 0.35-0.6 X the stem width. The underleaves of this
species represent the culmination of an interesting trend in the section,
from large, robust structures in C. moniliformis (here to 6.7 X stem width)
and C. humilis var. humilis to those of C. humilis subsp. polymorpha to
C. vermicular is, and finally, with a considerable reduction in magnitude,
to C. minutiretis.
Second, opposing male bracts are fused toward the base, a feature
occurring only in C. trachyopa and C. obvoluta of the quite unrelated
Subg. Lacerifolia.
In dorsal aspect, plants of C. minutiretis resemble C. humilis var.
suspecta quite closely. A ventral view will immediately distinguish C.
minutiretis, with its reduced, inconspicuous underleaves that are at most
0.6 X the stem width and bifid to 0.55-0.7.
At first glance, C. minutiretis brings to mind C. navistipula var. navisti-
pula in general aspect of the plants, and in the reduced, inconspicuous
underleaves and profuse branching (see fig. 53). However, I regard the
similarity of the taxa as a superficial one and believe C. minutiretis belongs,
more naturally, in Sect. Clasmatocolea.
Ecology. — Over rock in mossy forests or shaded, rocky stream valleys;
sometimes mixed with C. humilis, less often with C. trachyopa.
Phytogeography. — Tristan da Cunha and southern South America,
occurring at ca. 50° 00-20'S. in Prov. Magallanes and in Prov. Neuquen,
Argentina (1,500 m.) (see fig. 42).
Specimens seen. — TRISTAN DA CUNHA: Above Burntwood, Chris-
tophersen & Mejland 803 (hb. Grolle). CHILE. PROV. MAGALLANES:
S. side of I. Madre de Dios, head of fiord E. of M. Roberto, Engel 5123,
5129B, 5156 (MSC); E. shore of I. Pilot, Engel 4750 - c. per., 4770 -
c. sporo. + d" (MSC). ARGENTINA. PROV. RIO NEGRO: Parque
Nac. Nahuel Huapi, [M.] Tronador, 1,500 m., Donat 182 (mixed with
holotype of L. catenulata = C. humilis} (hb. Grolle, JE).
Sect. FULVELLAE Engel, sect. nov.
Rami intercalares plane dissimiles quam principale axe, vermiformes et submonili-
formes, angustiores quam principale axe, plerumque copiosi, et breves, abbreviati,
habentes amphigastria imbricata propinquioria quam illis principale axe; amphigastria
rami intercalaris grandia, orbicularia, valde cupulata, inflata, plerumque indivisa;
folia caulina principalis axis sinuata ac incurvata, plerumque habentia aliquot grandium
dentium et laciniarum.
ENGEL: CLASMATOCOLEA 139
Type species: Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle.
Intercalary branches quite different from main axis, the branches vermiform and sub-
moniliform, narrower in width than main axis, commonly copiously produced and short,
abbreviated, with underleaves more closely imbricate than underleaves of main axis;
intercalary branch underleaves large, orbicular, distinctly cupulate, appearing inflated,
usually undivided; leaves of main axis sinuous and incurved, usually with a few large
teeth and laciniae; underleaves of main axis =*= cupulate, the apical portion commonly
either approaching or in contact with stem, the basal portion abruptly convex (ventral
view).
Distribution. — Southern South America and Falkland Is.
Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. Figures 43-45, Plates
19, 20.
Jungermannia fulvella Hook. f. & Tayl. London J. Bot. 3:464. 1844.
Chiloscyphus fulvellus (Hook. f. & Tayl.) Nees in G. L. & N. Syn.
Hep. 711. 1847. Lophocolea fulvella (Hook. f. & Tayl.) Mass. Nuovo
Giorn. Bot. Ital. I. 17:227. 1885. Clasmatocolea fulvella (Hook. f. &
Tayl.) Grolle, Rev. Bryol. Lichenol. 29:72. 1960. Lectotype (nov.):
Chile, Prov. Magallanes, I. Hermite, Hooker s.n. (FH! — c. per. +
sporo. + <f ).
Lophocolea homomalla Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 39. 1900, syn. fide Engel (1978). Lectotype (fide
Engel, 1978): Chile, Prov. Aisen, R. Aisen Valley, January 1897,
Dusen 253 (SI - c. per.).
Plants prostrate, but axes often sharply curved dorsally or ventrally, often in thick,
dense mats, pale green, often tinged with brown or pale brown throughout, nitid when
dry; axes to 2.7 cm. tall, to 2.8 mm. wide with leaves spread.
Branches very frequent, often with copious production of short to long lateral- or
ventral-intercalary branches that are narrower in width than main axis, the branches with
leaves and underleaves more closely imbricate and more concave than those on main axis,
giving the branches a narrow, vermiform, submoniliform appearance; ventral-intercalary
branches commonly somewhat laterally displaced and those of the lateral-intercalary
somewhat ventrally displaced, the ± vermiform, intercalary branches occasionally
repeatedly branched and may give rise to an enlarged axis like that of the main axis,
the latter of which may produce more vermiform axes; Frullania-/y/7e branches occa-
sional, appearing like the main axis.
Stems 7-13 cells high, cortex in l-2(-3 ventrally) rows of thick-walled cells the same
size or slightly smaller than the medullary, the cortex commonly with 1-2 rows of dorsal
cells the same size as the medullary cells and 2-3 rows of ventral cortical cells smaller
than those of the medullary; cuticle smooth; endophytic hyphae absent. Rhizoids in
fascicles from stem near underleaf base.
Leaves of main axis with insertion slightly recurved at ventral end; leaves loosely
imbricate, adaxially concave, especially in ventral portion, wide-ovate to =*= reniform;
apex broadly rounded and commonly incurved, undivided, usually with a few large teeth-
140
ENGEL: CLASMATOCOLEA 141
laciniae, occasionally entire; dorsal margin straight or slightly rounded, sinuate, occa-
sionally =*= reflexed, short to long decurrent, entire; ventral margin broadly rounded,
entire, or with a few large teeth, especially towards the apex. Intercalary branch leaves
ca. 0.5 the size of leaves, closely imbricate, more concave, more regularly toothed, the
teeth sharper.
Leaf cells thin-walled, trigones large, very often bulging, rarely confluent; median
leaf cells 12-25 n wide, (12-) 17-29 p, long; cuticle smooth to roughened and appearing
finely granular.
Underleaves 0.3-0.5 as long as wide, wider than stem, remote to contiguous, ± cucul-
late, the apical portion very frequently either approaching or in contact with stem, the
basal portion abruptly convex (ventral view) ; margins curved dor sally, ± clasping stem
and obliterating view of stem from lateral and ventral views; underleaves reniform to
subrectangular, occasionally suborbicular ; apices undivided and broadly rounded to
truncate or bidentate to refuse to short-bifid, the apices merely with 2 slime papillae or
with segments or teeth terminating in a slime papilla; margins of the lamina with 1-3
slime papillae or small to large teeth each terminating in a slime papilla. Intercalary
branch underleaves more cucullate (and appearing ± inflated) and imbricate, usually
undivided, less frequently ret use, rarely short-bifid.
Plants dioecious; androecia terminal or intercalary on short or rather long lateral-
or ventral-intercalary branches; bracts densely imbricate and with apices appressed to
bract immediately above, the bracts strongly saccate; lobule margin with several slime
papillae and often few-celled teeth, and a lacinium, or with 1 large, broadly rounded
involute lobe; antheridia solitary, commonly orange-brown, stalk uniseriate.
Gynoecia on short, abbreviated, ventral- or lateral-intercalary branches; subfloral
innovations absent; bracts in 2 series, those of innermost series subrectangular to
obovate to ± spatulate; apices truncate to broadly rounded, with few to numerous
teeth that often terminate in a slime papilla; margins entire. Bracteoles of innermost
series free from bracts, subrectangular to obovate to ± spatulate; apices undivided
and broadly rounded or retuse, with few to numerous teeth or merely with a few slime
papillae; margins entire or with a few teeth or slime papillae. Perianth trigonous,
elongate-subrectangular to subclavate to ± fusciform, not or barely narrowing toward
the wide and open, rarely closed mouth; lobes broadly rounded, rarely incurved, but the
ventral often infolded, the lobes with numerous small teeth that often terminate in a
slime papilla; keel occasionally with wings of a few cells high, the wings occasionally
dentate.
Seta 8-10 cells in diameter, with 26-32 rows of outer thick-walled cells (with corners
thickened similar to trigones) surrounding an inner core of scattered cells that are
Opposite:
FIG. 43. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. 1, Portion of plant, dorsal-
lateral view. 2, Portion of plant with a few branches. 3, Portion of main axis, lateral
view. 4, Portion of main axis, ventral view. 5, Portion of intercalary branch, ventral
view. 6, Leaf of main axis. 7. Leaves of intercalary branch. 8-12, Underleaves of main
axis, flattened. 13, Median leaf cells. 14, Portions of main axis underleaf apices, each
with a slime papilla. 15, Stem of main axis, cross section. Figures 1-8, 10, 13, 14a, 15,
from lectotype of Jungermannia fulvella, Hooker, Chile, Prov. Magallanes, I. Hermite;
figures 9, 1 1, 14b, from Engel6323, Chile, Prov. Magallanes, B. San Nicholas; figure 12,
from Engel 2998B, Falkland Is., Mt. Adam.
142
FIELDIANA: BOTANY
FIG. 44. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. 1, Perianth mouth. 2, Seta,
cross section. 3, Outer capsule wall cells. 4, Inner capsule wall cells. 5, Capsule wall,
cross section. Figure 1, from Engel 3006, Falkland Is., Mt. Adam; figures 2-5, from
Engel 6125, Chile, Prov. Magallanes, B. Borja.
smaller, equal to, or slightly larger than the outer row. Capsule broadly ovate to elliptic,
the valves 0.91-1.19 mm. long, 41-53 /* thick, of 4-5 layers, rarely (and very locally) in
3 layers, the outer row of cells equal to thickness of 2.5-3 of interior strata; outer layer
with red-brown, nodule-like to ± spinelike thickenings that are often feebly tangen-
tially dilated, a very few semiannular bands occasionally present; exposed wall thick-
ened; intermediate and inner layers of cells subequal in thickness, the intermediate
layers with thickenings occasionally feebly extending onto tangential walls; inner layer
of cells with red-brown semiannular bands, the bands often incomplete, the radial walls
with nodulose thickenings often present.
Spores 13-17 jt, light brown, exine with the light microscope appearing minutely
punctate, but under the SEM with rather densely granulate to short vermiform projec-
tions covered with often crowded nanogranules, the intervening hollows with isolated
nanogranules; spores averaging 1.9 X elater diameter. Elaters 7-9 /u wide, tapering
gradually toward ends, apical portion often with thick, nonspiral walls, elater walls
light brown.
Clasmatocolea fulvella forms a transition of sorts between Sect. Clasmat-
ocolea, to which it is allied, and Sect. Puccioanae (Subg. Metaclasmato-
coled). Clasmatocolea fulvella is allied to Sect. Puccioanae because it
shares, with C. puccioana, a highly unique branching pattern within the
genus. The branches in these two taxa are copiously produced, with the
intercalary branches vermiform and submoniliform, narrow, smaller in
width than main axis, and usually short and abbreviated.
Variation. — The variability in position of ventral- and lateral-intercalary
ENGEL: CLASMATOCOLEA 143
branches in C.fuhella is worthy of note. In this species ventral merophytes
are very wide, which allows for branch development from the lateral
portions as well as from the middle of the merophyte. This condition,
which also occurs in the genus Calypogeia, explains the lateral displace-
ment of ventral-intercalary branches in C. fulvella (B. Stotler, in lift.).
The lateral-intercalary branches commonly are somewhat ventrally dis-
placed and occur close to the ventral merophyte in the following positions :
(a) very close to the ventral base of the leaves and not in close proximity
with an underleaf ; (b) at the very edge of the underleaf ; or (c) between the
underleaf and the ventral insertion of the leaf.
Differentiation. — Clasmatocolea fulvella is a distinctive taxon and should
not be confused with any other member of the genus.
The underleaves show little variability and possess characteristics very
distinctive of the species. The main axis underleaves are reniform to
suborbicular to subrectangular in outline, have apices very frequently
either approaching or in contact with the stem, and become abruptly
convex (ventral view) near the insertion. The margins are curved dorsal-
ly, ± clasping the stem and obliterating a view of the stem from lateral
and ventral views, lending the underleaf a cup-shaped appearance. The
underleaf apices are broadly rounded to truncate at the apex, which is
bidentate to retuse.
The branching is also distinctive for the species. The intercalary branches
are often copiously produced and are narrowly vermiform, ± submonili-
form, and considerably narrower than the main axis. Compared to the
main axis, the intercalary branch leaves are ca. 0.5 the leaf size, more
closely imbricate, more concave, more regularly and sharply toothed, less
sinuate, and with apices less incurved, whereas the intercalary branch
underleaves are considerably smaller, more closely imbricate, more cupu-
late (appearing ± inflated), and have apices less frequently retuse and
only rarely short-bifid.
The lateral-intercalary branches of C. fulvella are usually present, but if
absent, this species may be confused with C. humilis. Clasmatocolea
fulvella, when moistened, has main axis leaves sinuate and incurved and
has main axis underleaves ± clasping the stem, never spreading and with
apices very frequently either approaching or in contact with the stem.
Clasmatocolea humilis, on the other hand, when moistened, has main axis
leaves plane and has main axis underleaves not clasping the stem, occa-
sionally spreading and with apices not lying in contact with the stem. The
dentate to laciniate leaves of C. fulvella may be used as a supplementary
character to separate the species from C. humilis var. humilis.
Another condition, the reverse of the above, occasionally occurs — the
144 FIELDIANA: BOTANY
plants superficially appear to consist mainly of masses of branches, with
the distinctive combination of main axis and branches not apparent. In
this case the main axes should be searched for carefully. In the median
portion of a colony the main axes often are hidden and in various stages
of decay, likely resulting from the more vigorous growth of the younger
branches. The marginal portions of colonies exhibit well-developed,
vigorous main axes more typical of the species. In either instance, the very
distinctive characters of the branch leaves and underleaves discussed
above should be utilized to identify the species. This condition occurs
particularly where the species grows closely appressed to bark.
Clasmatocolea fulvella may be confused with some phenotypes of C.
humilis var. suspecta; see under the latter for comments to distinguish
these taxa.
Notes. — 1. In their original description of Jungermannia fulvella, Hooker
& Taylor (1844) make no mention of a perianth or sporophyte, but three
years later (Taylor & Hooker, 1847), they describe a perianth and include
a figure of a sporophyte-bearing plant in their plate of the species. Taylor
& Hooker (1847) are thus typifying the species as possessing both perianths
and sporophytes; I have designated as a lectotype a specimen in the Taylor
Herbarium (FH) that has both these structures represented.
2. There are two taxa that comprise the syntypes of Lophocolea homo-
malla — C. cucullistipula and C. fulvella. Stephani (1900, p. 40) states for
the Rio Aisen material, "in ramulis et truncis putridis." The material
collected "in ramulis" is C. cucullistipula, whereas that on "truncis pu-
tridis" is C. fulvella. The protologue resembles the C. fulvella material
more closely, particularly with regard to the comment on branching and
the description of the underleaves. Because perianths are described in the
original description, the lectotype must contain perianth-bearing plants.
Stephani in his Species Hepaticarum (1906, p. 78) states, "typhus in herb.
Stephani," so, with this in mind, I searched for perianths among the
Geneva material, but was unsuccessful. I therefore excluded Stephani
Herbarium material from consideration of a lectotype. I have studied
Dusen collected material of L. homomalla from a number of herbaria, but
have not seen specimens from Isla Guaitecas, Boca Chica, the second of
two localities mentioned by Stephani. Three of the four collections from
G were labeled merely "Patagonia," without a specific locality, and it is
quite possible that at least some of these were gathered at Boca Chica.
The Stephani Icones of L. homomalla (Icones Hepaticarum, Lophocolea
No. 90) is a mixture of information relating to both C. fulvella and C.
cucullistipula. The figure of a stem with an attached underleaf and pair of
leaves is of C. fulvella, yet within the leaf there is a note that the trigones
FIG. 45. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle.
145
146 FIELDIANA: BOTANY
are "magn. contig.," a feature more relevant to C. cucullistipula. The
figures of bracts, bracteole, and perianth mouth are much closer to C.
cucullistipula.
Ecology. — In the Magellanian region, very sporadic on rotted wood in
well-shaded forests (sea level) or over rock in more open subalpine areas.
In the Valdivian region the species seems more altitudinally zoned, occur-
ring 900-1,700 m. in forests. Here the species grows nearly exclusively on
rotted, decorticated wood in forests of the following types: Nothofagus
pumilio, Nothofagus scrub, Eitzroya, Araucaria, and Araucaria- Nothofagus
pumilio (in the latter two forest types the species is one of a few hepatics
growing on rotted, decorticated Araucaria wood, where it is quite common).
Phytogeography. — Falkland Is. (320-700 m.); rather common in Tierra
del Fuego (300-600 m.); N. to 36° 50' S. in West Patagonia; also in Parque
Nac. Nahuel Huapi of Andean Patagonia (see fig. 45).
Specimens seen.— FALKLAND IS. (selected). EAST FALKLANDS:
Stanley Region, summit of Mt. Kent, 455 m., Engel 2753, 2759 (MSC).
WEST FALKLANDS: Port Howard, Freezer Rocks, on E. slope of Mt.
Maria, 320 m., Engel 3143, 3150 (MSC); Mt. Adam, Halle & Skottsberg
226 as L. puccioana (UPS); Weddell I., Halle & Skottsberg 226 as L.
puccioana (UPS). ARGENTINA. TERR. TIERRA DEL FUEGO: I.
des los Estados, Spegazzini 63* (VER); ibid., Pto. Cook, Skottsberg
s.n. (S); ibid., Skottsberg s.n. as L. gayana (S); ibid., M. Conegliano,
Spegazzini 8 as L. puccioana (NY); I. de los Estados, Pto. San Juan del
Salvamiento, Spegazzini 51 (NY, VER — c. per.); Carbajal Valley,
base of Sa. de Sorondo, 300 m., Crow 1784 — c. sporo., 7797 — c. sporo.
(MSC); C. Garibaldi, SE of L. Escondido, near Rte. 3 from Ushuaia
to R. Grande, 600m., Schuster 5831 2a (hb. Schuster); Paso Garibaldi,
460-470 m., Roivainen 579, 580 - c. per. (F, H); Sa. Alvear, 370m.,
Roivainen 846 - c. sporo. (F, H). CHILE. PROV. MAGALLANES:
"S. part of Tierra del Fuego," Darwin 467 (FH — c. sporo., NY); Cabo
de Hornos, Antarc. Exped. as syntype of /. fulvella (NY); ibid., Davis
s.n. (FH, S); ibid., Hooker s.n. as syntype of J. fulvella (NY); I. Hermite,
Hooker s.n. as syntype of J. fulvella (NY, S); ibid., sin. coll. as L. cooki-
ana (PC); B. Orange, sin. coll. 104 (PC); R. Azopardo, Dusen 51 as
L. humilis (LD - c. per., UPS); ibid., Dusen 86 (NY, S); Cta. Gomez,
Halle & Skottsberg 207 as L. homomalla (S); I. Desolation, Dusen (NY,
S); ibid., Pto. Angosto, Dusen s.n. as L. humilis (S); ibid., Dusen 180 as L.
humilis (UPS); ibid., Dusen 263 (S); Brunswick Pen. (cf. Engel, 1978); E.
side of B. Borja, Engel 6125, 6232 A (MSC); Can. Gajardo, Cta. Inga,
Halle & Skottsberg 202 - c. per. + <? (S); B. Sholl, Discovery Exped.
982/1 (hb. Grolle); head of F. Peel, Engel 5502 - c. sporo. (MSC).
ENGEL: CLASMATOCOLEA 147
PROV. AISEN: R. Aisen, Dusen s.n. as syntype of L. homomalla (NY,
S); ibid., Dusen 253 as syntype of L. homomalla (S); ibid., Dusen 263
(NY); ibid., Dusen 287 as syntype of L. homomalla (FH); Pto. Puyuhuapi,
1,700 m., Schwabe 32 ex p. (JE); C. Tesoro, 1,000 m., Schwabe 39/c as
L. homomalla (JE); ibid., 900 m., Schwabe 40a (hb. Grolle - c. per. +
cf, JE). PROV. CHILOE: Pto. Quellon, Halle & Skottsberg 220 as L.
navistipula (UPS). PROV. OSORNO: Antillanca, 1,180 m., Ruthsatz 54/17
(JE); Refugio Antillanca, 1,160 m., Engel 3915B (F); 0.5 km. by road
below Refugio Antillanca, 1,000 m., Engel 11504 (F). PROV. VALDIVIA:
Cord. Pelada, summit of El Mirador, between La Union and Pta. Hueicol-
la, 1,000 m., Engel 12391 (F); V. Shoshuenco, Ruthsatz 49/16, 49/49 (hb.
Grolle); SW slope of V. Quetrupillan, 1,160-1,450 m., Engel 11159 - c.
sporo., 11162 - c. sporo., 11164 (F). PROV. CAUTIN: V. Lanin, 1,220
m., Ruthsatz s.n. (JE); L. Quilleihue, 10.7 km. by road E. of Puesco, ca.
7 km. by road W. of Chile-Argentina boundary, 1,050 m., Engel 11311
(F); V. Llaima, 1,260-1,280 m., Ruthsatz s.n. (JE); Parque Nac. Conguillo,
1,140-1,150 m., Mahu 10787, 10796, 10835 (hb. Mahu). PROV. MALLE-
CO: Parque Nac. Nahuelbuta, Pino Hueco, 1,400 m., Mahu 6055 (hb.
Grolle, JE, hb. Mahu). PROV. MALLECO/PROV. ARAUCO: Cord.
Nahuelbuta, Parque Nac. Nahuelbuta, W. of Angol, 1,280-1,300 m.,
Engel 12594, 12603 - c. sporo., 72527 (F). PROV. CONCEPClON:
Conception, Dusen s.n. as L. humilis (S); ibid., Dusen 180 as L. humilis
(NY). ARGENTINA. PROV. NEUQUEN/PROV. RIO NEGRO: Parque
Nac. Nahuel Huapi, Donat 27 ex p. as L. homomalla (S); ibid., 800 m.,
Donat 61 ex p. (JE); L. Nahuel Huapi, Puerto Blest, Donat 136/a (JE).
Sect. STRONGYLOPHYLLAE Engel, sect. nov.
Foliorum caulinarum cuticula habens grossas hemisphaericas hyalinas papillas.
Type species : Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle.
Leaf cuticle with coarse, hemispherical, hyaline papillae.
Distribution. — New Zealand and its subantarctic islands, Tasmania,
Java.
Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle. Figures 46,47,
Plates 2 1,22.
Jungermannia strongylophylla Hook. f. & Tayl. London J. Bot. 3:370.
1844. Alicularia strongylophylla (Hook. f. & Tayl.) G. L. & N. Syn.
Hep. 619. 1846. Leptoscyphus strongylophyllus (Hook. f. & Tayl.)
Mitt. Hooker's J. Bot. Kew Card. Misc. 3:358. 1851. Leioscyphus
strongylophyllus (Hook. f. & Tayl.) Mitt, in Hooker, Bot. Antarc.
Voy. 3:225. 1859. Mesophylla strongylophylla (Hook. f. & Tayl.)
f
FIG. 46. Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle. 1, Portion of main
axis, dorsal view. 2a-e, Leaves. 3a-m, Underleaves. 4, Underleaf segment, apical portion
with partially collapsed slime papilla. 5-6, Stem, cross sections. 7, Median leaf cells
showing coarsely papillose cuticle. 8, Perianth lobes, ventral lobe in middle. Figures
1, 2c, d, 3b, i, m, 4, 5, 7, from type of C. strongylophylla, Hooker, Campbell I. (S);
figure 2a, from Langridge 179, New Zealand, Harihari; figures 2b, 3e, j, b, from G.
Scott, New Zealand, S. of Dunedin; figures 2e, 3g, from Hooker, Auckland Is.; figure
3a, from type of C. strongylophylla var. /3 minor, Hooker, Auckland Is.; figure 3c,
from D. Scott 208, New Zealand, Godley R. Valley; figure 3d, from Wilson 2775,
New Zealand, Mt. Cook Natl. Park; figures 3f, k, 1, from type of/, turgescens, Hooker,
Auckland Is.; figures 3h, 8, from Schuster 48481, New Zealand, Otira R. Gorge.
148
ENGEL: CLASMATOCOLEA 149
Trev. Mem. 1st. Lomb. Sci. Lett. III. 4:398. 1877. Lophocolea strongy-
lophylla (Hook. f. & Tayl.) Hodgs. Trans. Roy. Soc. New Zealand
80:338. 1953. Clasmatocolea strongylophylla (Hook. f. & Tayl.)
Grolle, Rev. Bryol. Lichenol. 29:73. 1960. Original material: Camp-
bell I., Hooker (BM!, S! — c. cf + young per.).
Jungermannia strongylophylla var. /3 minor Hook. f. & Tayl. London
J. Bot. 3:370. 1844, syn.fide Grolle (1962). Alicularia strongylophylla
var. minor (Hook. f. & Tayl.) G. L. & N. Syn. Hep. 619. 1846.
Original material: Auckland Is., Hooker (BM!, L!).
Jungermannia turgescens Hook. f. & Tayl. London J. Bot. 3:375. 1844,
syn. fide Grolle (1962). Leptoscyphus turgescens (Hook. f. & Tayl.)
Mitt. Hooker's J. Bot. Kew Gard. Misc. 3:358. 1851. Leioscyphus
turgescens (Hook. f. & Tayl.) Mitt, in Hooker, Bot. Antarc. Voy.
3:225. 1859. Mylia turgescens (Hook. f. & Tayl.) Trev. Mem. 1st.
Lomb. Sci. Lett. III. 4:412. 1877. Lophocolea turgescens (Hook. f. &
Tayl.) Hodgs. in Martin, Trans. & Proc. Roy. Soc. New Zealand
78:495. 1950. Clasmatocolea turgescens (Hook. f. & Tayl.) Grolle,
Rev. Bryol. Lichenol. 29:73. 1960. Original material: Auckland Is.,
Hooker (LD! — c. per., S! — c. per.).
Lophocolea angulistipula Steph. Bull. Herb. Boissier 6 (9): 791. 1906
(= Spec. Hep. 3:91), syn. nov. Clasmatocolea angulistipula (Steph.)
Grolle, Rev. Bryol. Lichenol. 29:71. 1960. Holotype: Tasmania, Mt.
Wellington, St. Crispin's, 24 January 1899, Weymouth 1514 (G!).
Plants rather delicate, prostrate, usually creeping among other bryophytes, occa-
sionally caespitose, green to light or dull brown; axes 250-950 n wide without leaves
spread.
Branching sparing, mostly of Frullania-0'/^, occasionally of ventral-intercalary type,
lateral-intercalary type very rare.
Stems 4-7 cells high, cortex in a single layer of thick-walled cells usually moderately
to considerably larger than but occasionally ca. same size as medullary cells; cuticle
with low, rounded to hemispherical papillae or rarely striate ; endophy tic hyphae rare.
Rhizoids in fascicles from stem at base of underleaves.
Leaves with insertion narrow, slightly to distinctly recurved at ventral end; leaves
erect to slightly spreading, distant (especially in smaller plants) to loosely imbricate,
strongly adaxially concave, orbicular to reniform (-obovate); apex broadly rounded to
truncate to subretuse, undivided, often incurved, the apices and margins entire; dorsal
margin long decurrent.
Leaf cells thin to slightly thick walled, trigones large to bulging, occasionally confluent;
median leaf cells 10-20(-24) /* wide, 12-23(-28) » long; cuticle, including that of leaf
margin, with coarse, hemispherical-oblong, hyaline, usually juxtaposed papillae, very
rarely wide-striate, cuticle of basal cells often wide-striate.
Underleaves / to slightly over 2 X stem width, transversely or obliquely inserted,
free or weakly connate on 1 side, slightly to (often) strongly spreading, usually plane,
150 FIELDIANA: BOTANY
sometimes weakly convex (ventral view); underleaves polymorphous, often with lobes
of different size and configuration, occasionally undivided and then lanceolate or oblong
to =*= orbicular to cuneate with usually a truncate apex, bifid to 0.5; segments, when
present, directed laterally or toward stem apex, medium-triangular to lanceolate or
reduced to a rounded projection, tooth or slime papilla, or the segments often apiculate
and terminating in a slime papilla, the segment margins entire; sinus broadly rounded
to often lunate; margins of the lamina entire or with 1-2 slime papillae, very rarely with
a small tooth terminating in a slime papilla.
Plants dioecious; androecia intercalary on main axis or Frullania-type branches, the
entire bract strongly saccate, and with apices sometimes appressed to bract immediately
above; lobule margin slightly incurved, broadly rounded, entire or with 1 slime papilla;
antheridia solitary, stalk uniseriate.
Gynoecia on main axis, long ventral-intercalary branches or subfloral innovations;
subfloral innovations usually from below innermost bracteole; bracts in (2-)3 series,
those of innermost series with apices often more incurved than in leaves, the apices
and margins entire. Bracteoles of innermost series free from bracts, oblong to obovate
to subsquarrose ; apices emarginate or to 0.2-bifid; lamina margins with several slime
papillae; bracteoles occasionally not or hardly larger than the underleaves, and then
bracteoles undivided and elliptical. Perianth somewhat inflated, terete to obscurely
trigonous to bilaterally compressed, =*= clavate, expanding toward mouth, the mouth
somewhat to distinctly closed by incurved lobes; dorsal lobes broadly rounded, occasion-
ally emarginate, with a few slime papillae and rounded teeth of varying sizes, occasion-
ally entire; ventral lobe smaller and often more narrowly rounded, variable in size,
slightly convex, to concave, to folded inward, the latter thus giving the perianth a
bilaterally compressed appearance, the ventral lobe sometimes retuse, with a few slime
papillae and a tooth or entire; keels (especially the ventral) winged.
Seta not seen. Capsule wall 36 n thick, of 4 layers, outer row of cells equal to thickness
of 2 of intermediate strata; outer layer with red-brown thickenings, the thickenings of
blunt-tipped spines, semiannular bands common, complete or incomplete, somewhat
thickened, the thickenings occasionally nodule-like; exposed wall thin; inner layer of
cells with yellow or red-brown nodule-like or spinelike thickenings, semiannular
bands often present.
Spores 11-12 n, light brown, exine with the light microscope appearing minutely
punctate-short vermiculate, but under the SEM with a network of very wide, irregular,
often coalescing vermiform ridges covered with nanogranules ; spores averaging 1.2 X
elater diameter. Elaters 9-1 1 n wide, tapering slightly toward tips, apical portions often
with thick, nonspiral walls, elater walls red-brown.
Variation. — The perianths of C. strongylophylla vary from terete to
obscurely trigonous to bilaterally compressed. The ventral lobe is con-
sistently smaller (often strikingly so) than the dorsal pair and varies from
slightly convex to concave to distinctly infolded. In the latter instance the
perianths appear bilaterally compressed. This kind of variation is not
unusual within a single species in the Lophocoleaceae, i.e., in the genus
Leptoscyphus, which nearly always has strongly bilaterally compressed
perianths, a ventral lobe is commonly present that varies in stature and is
particularly well developed in some specimens of, for example, L. expansus
and L. horizontalis.
ENGEL: CLASMATOCOLEA 151
Differentiation. — The polymorphic underleaves of C. strongylophylla are
diagnostic, especially with regard to the lobes, which vary in configura-
tion, length, and overall stature, usually with one lobe of a different size
and configuration than the other. On a given axis there are commonly
bifid or bidentate underleaves among those that are undivided and lanceo-
late, with one lobe reduced to a tooth or slime papilla. Underleaves of
variable sizes and configurations can usually be demonstrated on a single
axis. The polymorphous underleaves, coupled with the coarse, hemispheri-
cal, hyaline, juxtaposed cuticular papillae, the large to bulging trigones,
the small axis size (250-950 n wide), and the perianths with a smaller
ventral lobe that is commonly slightly to distinctly infolded, will serve to
separate this species from all others of the genus.
Notes. — 1. The original material of Jungermannia strongylophylla de-
posited in the British Museum is somewhat flaccid and has rather poorly
developed cuticular papillae, whereas original material of the species
deposited in Stockholm (S) is not flaccid and has well-developed cuticular
papillae.
Original material of J. strongylophylla consists of brownish, very small
plants with distant to contiguous leaves that have a loose network of cells
with cuticular papillae ± hemispherical and rather closely juxtaposed.
On the other hand, original material of J. turgescens consists of light brown,
quite robust plants (0.63-0.95 /j. wide) with loosely imbricate leaves that
have a dense network of cells, with the cuticle possessing at most scat-
tered, ± hemispherical papillae or wide striae.8 Portions of leaves of the
latter may even have a smooth cuticle. I can, however, find no consistent
character to separate C. turgescens from C. strongylophylla, and I therefore
regard the original material of Jungermannia turgescens as an extreme in
a continuum with C. strongylophylla and concur with Grolle (1962) in
treating them as conspecific.
2. Stephani (1906) used the spelling Lophocolea angulistipula, although
the name written on the label of the type specimen is Lophocolea angu-
listipa. This is a case of Stephani changing his mind regarding the
spelling of a name to be published (cf. Engel, 1973b).
3. Reports of this species (sub Leioscyphus turgescens) from southern
South America and Kerguelen are erroneous. Those from Isla Desolation
are C. gayana, whereas those from Kerguelen are C. humilis. Although I
have not seen Rio Azopardo collections, I would not expect the species to
occur there.
8 The large cuticular papillae are sometimes (as in original material of J. turgescens)
nearly transparent and thus difficult to demonstrate and must be searched for with care.
J «0
H
FIG. 47. Clasmatocolea strongylophylla (Hook. f. & Tayl.) Grolle.
152
ENGEL: CLASMATOCOLEA 153
Ecology. — In New Zealand on steep, shaded, humid, or dripping cliffs
or rock faces in Nothofagus menziesii forests (with Blepharidophyllum
vertebrate and Lepidozia spinosissima, etc.) or tall subalpine scrub. Some-
times on wet, rocky, or sandy banks by roadsides where occasionally with
Bryum laevigatum. In Tasmania mostly above 900 m. and in subalpine
situations. It grows on soil over boulders or under protective cover of
scrubby vegetation, either on soil or on dripping cliff faces. Also in
Nothofagus gunnii-Eucalyptus coccifera forests where again it is associated
with more moist microhabitats, such as boulders of stream banks or lake
shores.
Phytogeography. — Campbell I.; Auckland Is.; New Zealand, South I.
(90-1,370 m.); Tasmania (sea level- 1,3 10 m.) (see fig. 47).
Specimens seen.— CAMPBELL I.: Mt. Beeman, Members of 1958
Expedition s.n. as C. turgescens (CHR). AUCKLAND IS.: Sin. coll. - c.
c? (BM); Erebus & Terror Exped. - c. per. (BM); Hooker s.n. (BM, L,
NY); Hooker Hills, Fineran s.n. (hb. Grolle). NEW ZEALAND. SOUTH
ISLAND. OTAGO: Waipori Road, S. of Dunedin, ca. 300 m., Scott
s.n. - c. per. (CHR); Waipori Road, W. of Dunedin, 490 m., Scott 15
(hb. Monash Univ.). Fiordland Natl. Park, track from L. Mackenzie to
Harris Saddle, Humboldt Mts., Schuster 67-504 (F, hb. Schuster); Fiord-
land Natl. Park, Humboldt Mts., N. of Earland Falls, on track to L.
Mackenzie, Schuster 67-3456B (F, hb. Schuster); Fiordland Natl. Park,
L. Mackenzie to Harris Saddle, above Hollyford Valley, Route Burn
Track, 90-155 m., Schuster 67-3904 (F, hb. Schuster). SOUTHLAND:
"Head of Hollyford R., Doubtful Sound area," Martin s.n. (hb. Grolle).
WESTLAND: Pleasant Flat on Haast Pass Road, Allison s.n. (CHR);
0.6 mi. N. of Haast Pass, 520 m., Schuster 53380a as C. turgescens — c.
per. + c? (hb. Schuster); Harihari, Langridge 179 as C. turgescens (CHR);
Arthur's Pass Natl. Park, Otira R. Gorge, W. of Arthur's Pass, ca. 300-
365 m., Schuster 48481 as C. turgescens — c. sporo. + c? (hb. Grolle, hb.
Schuster, TNS); ibid., Schuster 48484b as C. turgescens - c. <? (hb.
Schuster); Otira Pass, ? Jack s.n. as C. turgescens (CHR). CANTERBURY:
Mt. Cook Natl. Park, edge of Governors Bush, ca. 720 m., Wilson 2775
(CHR); mid Godley R. Valley, 1,370 m., D. Scott 208 (CHR). TAS-
MANIA: Kermandie R. at North Cr., ca. 50 m., Norris 29394 (F); Mt.
Field Natl. Park, A. Ratkowsky 78/171 (F); ibid., Beatties Tarn, 975 m.,
Engel 14509 A (F); Mt. Field Natl. Park, just below SE end of L. Fenton,
Lady Barren Cr., 1,000 m., Engel 13067 - c. per., 13075 A (F); Mt. Field
Natl. Park, just S. of L. Fenton and NE of Wombat Moor, 1,020 m.,
Engel 13268 (F); Mt. Field Natl. Park, Mt. Mawson Plateau, 1,200 m.,
A. Ratkowsky 78/75 (F); ibid., above and W. of L. Dobson and S. of L.
154 FIELDIANA: BOTANY
Seal, 1,250 m., Engel 13091 - c. sporo. (F); Mt. Field Natl. Park, Tarn
Shelf, below and E. of Rodway Range, 1,270 m., Engel 14355, 14368 (F);
Mt. Field Natl. Park, Rodway Range, between Rodway Ski Tow and K
Col area, 1,240-1,310 m., Engel 14393 (F); Mt. Field Natl. Park, L.
Belcher, ca. 900 m., Norn's 28619, 28621 (F); plateau region E. and SE of
Mt. Eliza summit, E. of L. Pedder, ca. 1,225 m., Engel 13713 — c. tf (F);
Mt. Sprent, A. Ratkowsky 78/172 (F); Gordon R. just E. of confluence
with Serpentine R., WNW of Strathgordon, 350 m., Engel 13831 - c. &,
13841 (F); Gordon R., Sir John Falls, just up river from Butler I., sea
level, Engel 14683 - c. sporo. (F); Mt. Rufus, A. Ratkowsky 79/15 (F);
Mt. King William I, 1,200 m., A. Ratkowsky 77/76 (F); E. slope of Black
Bluff just below summit, S. of Burnie, 1,250 m., Engel 15800, 15807 A -
c. per. (F); SW slope of St. Valentines P., SSW of Burnie, 920-1,100 m.,
Engel 15633 - c. per. (F).
Clasmatocolea tjiwideiensis (Sande-Lac.) Grolle. Figure 48.
Chiloscyphus tjiwideiensis Sande-Lac. Ned. Kruidk. Arch. 3:418. 1854.
Anthoscyphus tjiwideiensis (Sande-Lac.) Trev. Mem. 1st. Lomb.
Sci. Lett. III. 4:422. 1877. Conoscyphm tjiwideiensis (Sande-Lac.)
Schiffn. Consp. Hep. Arch. Ind. p. 124. 1898. Clasmatocolea tjiwideien-
sis (Sande-Lac.) Grolle, Rev. Bryol. Lichenol. 29:73. 1960. Original
material: Java, Crater Tjiwidei, Junghuhn (L no. 910. 288-134!).
Plants soft, spongy, delicate, apparently prostrate, yellow-brown or light brown;
axes 0.65-0.91 mm. wide.
Branches occasional, of ventral-intercalary type.
Stems rather slender for plant size, flexuous, sometimes golden brown, 6-9 cells high,
the cortex hardly differentiated, in 1-2 rows of moderately to distinctly thick-walled
cells ± same size as medullary cells; cuticle papillose; medullary cell walls moderately
to distinctly thickened, the thickenings of =*= same magnitude as those of cortical cells;
endophytic hyphae in both cortical and medullary cells. Rhizoids magenta, in dense
fascicles from stem very near underleaf base, occasionally from stem near ventral
base of leaf.
Leaves with insertion sharply recurved at ventral end ; leaves strongly erect, dorsally
connivent, rather loosely imbricate, distinctly conchiform concave, oblate to subsquar-
rose; apex broadly rounded, undivided, entire, sometimes repand; dorsal margin =±=
straight (at least in proximal ca. 0.75), often moderately wavy, often slightly recurved,
entire or with 1-2 sharp, often hooklike teeth in median and/or lower portion, the
margin conspicuously long decurrent; ventral margin very broadly rounded, the basal
portion nearly always with l-2(-3) few-celled, sharp or blunt teeth, the armature on that
portion of margin decending below stem, the margin decurrent, sometimes strongly so
and then connate with basal, adaxial surface of underleaf.
Leaf cells thin to moderately thick-walled, the trigones medium to large, the sides
straight to slightly bulging, with secondary cell wall thickenings occasional; median
leaf cells (25-)28-37 M wide, 28-38(-42) M long, cuticle, including that of leaf margin,
ENGEL: CLASMATOCOLEA 155
with hemispherical-oblong, hyaline, usually juxtaposed papillae, the cuticle of basal
cells papillose-striate to striate.
Underleaves 3.3-4 X stem width (when underleaf flattened), connate on 1 or both sides
with ventral-basal, adaxial portion of leaf lamina; underleaves narrow, slightly spreading,
distant to approximate, deeply convex to =*= cupulate (ventral view), ovate to suborbic-
ular; apices variable (usually so on same axis): undivided and 1- or bidentate or divided
to 0.45; segments, when present, somewhat incurved, the segment margins entire;
margins of the lamina 1-3-dentate, rarely entire, short to long decurrent.
Plants apparently dioecious; androecia not seen. Gynoecia on leading axes, only im-
mature individuals seen.
Differentiation. — The following combination of features will distinguish
this species from all others in the genus: (a) the coarse, hemispherical,
hyaline papillae of the leaves; (b) the strongly erect, dorsally connivent,
oblate to subsquarrose leaves with dorsal margin straight (at least for the
most part), and conspicuously long decurrent on the stem; (c) the 1-3
few-celled teeth on the basal portion of the ventral margin (and usually
situated on that portion of the margin extending ventrally beyond the
stem); (d) the relatively small stems that are 6-8 cells high and possess
very thick-walled cells, with very little, if any, cortical differentiation;
(e) the magenta-colored rhizoids (unique to genus); (f) the relatively large
underleaves that are rather deeply convex (ventral view) to =>= 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. <? (MSC); F. Peel,
Skottsberg s.n. as syntype of L. rotundistipula (UPS); I. Chatham, B.
Wide, Engel 5361B-2 - c. per. (MSC); E. side of I. Juan, Engel 5248 D
ENGEL: CLASMATOCOLEA 165
(MSC); W. side of I. Grant, Engel 4705 D (MSC); Pto. Alert, Engel
4913 — c. sporo. (MSC); ibid., head of fiord W. of M. Markham, Engel
4959 - c. sporo., 4990 A - c. per. (MSC); Pto. Grappler, Skottsberg s.n.
(hb. Grolle); Pto. Eden, Engel 4515, 4520 - c. sporo., 4522 - c. <? ,
4528 - c. & , 4532A - c. <?, 4533 - c. d\ 4541 - c. sporo. + &
(MSC); B. Halt, Cunningham s.n. (NY); ibid., Cunningham 95 as Chilo-
scyphus notophyllus (BM). PROV. AISEN: N. side of F. Tempano, Engel
4385 A - c. per., 4399B - c. <f (MSC); Pto. Island, Engel 4272B - c.
per., 4311 (MSC). PROV. VALDIVIA: Cord. Pelada, summit of El
Mirador, between La Union and Pta. Hueicolla, 1,000 m., Engel 12371,
12376 (F).
Sect. METACLASMATOCOLEA
Lateral-intercalary branches usually of unlimited growth; underleaves of both main
axis and branches reduced, those of main axis frequently with asymmetric segments;
male bracts conduplicately bilobed.
Distribution. — Southern South America.
Clasmatocolea navistipula (Steph.) Grolle. Figures 52-54, Plates 26, 27.
For synonymy see under varieties.
Plants delicate, spongy, prostrate to suberect, in very compact, pure mats, less often
mixed with other hepatics, whitish-pale green to pale grey-green, sometimes tinged with
red-brown; main axes 455-595 p wide.
Branches copious, lateral-intercalary type abundant, ventral-intercalary type occasional,
Frullania-/y/je rare; branches often arising from a leafless main axis, the branches very
often long and of unlimited growth, often exceeding length of main axis, which usually
remains quite short, the branches often in turn branched or repeatedly branched;
branches slightly to moderately narrower than main axis ; terrestrial plants with branch-
ing pattern very often rather narrowly to broadly flabellate; aquatic or subaquatic
plants with only a few branches.
Stems 6-10 cells high, cortex in 1-3 rows of moderately thick-walled cells larger than,
slightly smaller than, or ca. of same size as medullary cells; exposed wall of outer
cortical row moderately to very thickened; medullary cell walls increasing in size
toward stem periphery, the outermost cells subequal to cortical cells, the medullary
cell walls slightly to moderately thickened and with corners thickened similarly to
small trigones; endophytic hyphae absent or present in cortex or both cortex and
medulla. Rhizoids long for plant size, in ill-defined areas at and near underleaf bases,
but several scattered along ventral surface of stem, especially near the leaf insertion.
Leaves (main axis) often caducous or partially decayed, the insertion very slightly
recurved at ventral end; leaves erect, rarely dorsally connivent, contiguous to loosely
imbricate, conchiform concave, =*= reniform to oblate; apex broadly rounded to trun-
cate, undivided, the apex and margins entire; dorsal margin =*= straight in proximal half,
frequently slightly recurved, the ventral margin broadly rounded.
Leaf cells thin walled, trigones large to bulging to (mostly) nodular and protuberant,
often confluent; median leaf cells 13-25 p wide, 17-31 n long; cuticle smooth or slightly
166
ENGEL: CLASMATOCOLEA 167
roughened and appearing finely granular. Oil-bodies throughout leaf, occupying small
fraction of cell lumen, hyaline, glistening, of grape-cluster type, with numerous, bulging
globules of 1-2 n in diameter, the oil-body surface appearing coarsely botryoidal, the
segmentation distinct; oil-bodies globose to ovoid to elliptic in shape, consistently 2 per
cell of leaf middle and apex, 2-4 of leaf base, those of leaf middle 5 n wide, 9-1 1 n long.
Under/eaves often caducous, 0.46-1.7 X stem width, often covering only a fraction of
stem tissue, sometimes becoming very reduced and inconspicuous toward main axis
apices, the underleaves free or weakly connate on 1 side, moderately to rather strongly
spreading, distant, ± plane, polymorphous: narrowly triangular to ovate to elliptic to
subrectangular to cuneate; apices undivided or bifid; segments (when present) often of
different size and configuration, often widely spreading and separated by a lunate sinus;
segments or undivided underleaf apex terminating in a slime papilla; segment margins
entire or (rarely) with 1 tooth ; lamina margins entire or with 1-2 teeth or slime papillae.
Branch underleaves very reduced, very inconspicuous, strongly spreading, sometimes
recurved, variable in size and shape, some (especially those of 2nd, 3rd, and 4th order)
of only a few cells and then scalelike, undivided, and =*= narrowly triangular, whereas
others are somewhat larger and sublanceolate; underleaves of branches sporadically
large, and then are occasionally short-bifid and with lobes or teeth often asymmetric.
Plants dioecious; androecia terminal or intercalary on lateral-intercalary branches
of varying lengths; bracts with basal portion strongly saccate and distal portion strongly
concave, with the often subcrenulate apices appressed to bract immediately above;
bracts conduplicately bilobed, the lobule small, elliptic to narrowly ovate, attached for
varying lengths to adaxial surface of lobe, the lobule sometimes smaller and lanceolate;
antheridia l(-2) per bract, the stalk uniseriate or very locally biseriate.
Gynoecia on rather short or abbreviated lateral- or ventral-intercalary branches; sub-
floral innovations absent; vestigial stem perigynium present; bracts and bracteoles of
innermost 2 or all 3 series inserted on the stem perigynium; bracts of innermost series
concave, ovate to ovate-wide subrectangular, and closely adhering to perianth; bract
apices rather narrowly to broadly rounded, undivided, entire; lamina margins some-
times recurved, entire or with 1 slime papilla. Bracteoles of innermost series free from
bracts, plane, ovate to elongate-subrectangular to narrowly triangular; apices poly-
Opposite:
FIG. 52. Clasmatocolea navistipula (Steph.) Grolle. 1, Lateral-intercalary branch,
with two young, secondary lateral-intercalary branches, ventral-lateral view. 2, 3, Ven-
tral and lateral views of same lateral-intercalary branch. 4, Leaf of long leading lateral-
intercalary branch. 5-27, Underleaves. 28, Median leaf cells. 29, Underleaf of large,
leading lateral-intercalary branch. 30, Stem, cross section. Figures 1, 4, 7, 10, 28, 29,
from type of L. navistipula, Dusen 155, Chile, Prov. Magallanes, I. Desolaci6n; figure
2, from Engel 2129-1, Chile, Prov. Magallanes, Pto. Cutter; figure 3, from Engel 6127,
Chile Prov. Magallanes, B. Borja; figures 5, 8, 16, from Engel 5357, Chile, Prov. Magal-
lanes, B. Wide; figures 6, 21, 30, from Engel 4251, Chile, Prov. Aisen, Pto. Island;
figures 9, 12, 13, 19, from Crow 1472, Argentina, Terr. Tierra del Fuego, Tierra Mayor
Valley; figures 11, 14, 15, from Engel 6175, Chile, Prov. Magallanes, B. Borja; figure
17, from Engel 4822, Chile, Prov. Magallanes, Pto. Charrua; figures 18, 20, from
Engel 4366, Chile, Prov. Aisen, F. Tempano; figures 22-25, 27, from Engel 2105, Chile,
Prov. Magallanes, La. El Parrillar region; figure 26, from Engel 6425, Chile, Prov.
Magallanes, between B. Bougainville and B. San Nicolas. (Figures 9, 12, 13, 19, 22-27
are of var. parceramosa).
168
ENGEL: CLASMATOCOLEA 169
morphous: undivided and truncate or acutely angular or narrowly rounded, or bifid to
0.30; lamina margins entire or with 1 small lobe. Perianth obscurely to moderately
trigonous, rather strongly inflated in median and basal portions, but with upper portion
somewhat bilaterally compressed (the perianths rather inflated throughout when young,
but often somewhat collapsing toward apex at maturity); perianth falcate-elliptic to
subcylindrical to ovoid to clavate, commonly narrowing toward the weakly opened or
closed, somewhat bilaterally compressed mouth; lobes often incurved or inrolled,
broadly rounded, the ventral considerably smaller, the lobes entire or with a few slime
papillae, occasionally sinuate.
Seta 6 cells in diameter, with 15-16 rows of outer cells surrounding an inner core of
scattered, smaller cells with corners thickened similar to medium trigones. Capsule
elliptic, the valves 0.88-0.98 mm. long, 42-48 M thick, of 4-5 layers, the outer layer of
cells 18-25 M thick, equal to thickness of 3 of interior strata or outer layer greater than
diameter of all interior strata combined; outer layer with very pale brown or hyaline
or red-brown, nodule-like to wide spinelike thickenings that are tangentially dilated,
a few semiannular bands present; exposed wall of outer layer moderately thickened;
intermediate strata with outermost layer thicker than inner layer of capsule, the inter-
mediate layers with thickenings often considerably tangentially dilated; inner layer of
cells (5-)6-8 n thick, with pale brown or red-brown semiannular bands very common,
the bands often incomplete, nodular thickenings occasional.
Spores 11-12/x, light brown, exine with the light microscope appearing minutely
punctate, but under the SEM with coarsely granulate to short, abbreviated, wide, vermi-
form projections that occasionally have nanogranules; spores averaging 1.5 X elater
diameter. Elaters 7-8 n wide, apical portions occasionally with thick nonspiral walls;
elater walls red-brown, 2(-3) M in diameter.
KEY TO VARIETIES OF Clasmatocolea navistipula
1 . Branching copious, in a narrow to broad flabellate pattern, Frullania-lype branching
absent; underleaves to 0.95 X stem width; plants on rotted stumps or logs or on
soil var. navistipula
1. Branching sparingly developed, not copiously produced and never in a flabellate
pattern, Frullania-type branching occasionally present; underleaves to 1.7 X stem
width ; plants of pools, pool margins, or Sphagnum bogs var. parceramosa
Clasmatocolea navistipula (Steph.) Grolle, var. navistipula.
Lophocolea navistipula Steph. Bull. Herb. Boissier 6 (7): 543. 1906
(= Spec. Hep. 3:57). Clasmatocolea navistipula (Steph.) Grolle,
Opposite:
FIG. 53. Clasmatocolea navistipula (Steph.) Grolle. 1, 2, Diagrammatic representation
of branching pattern of var. navistipula. 3-6, Diagrammatic representation of branching
pattern of var. parceramosa. 7, Perianth mouth, ventral lobe in middle (lobe to left has
3 slime papillae). 8, 9, Mature perianth, cross section through median portion. 10, 11,
Male bracts, adaxial view. Figure 1, from Engel 5357, Chile, Prov. Magallanes, B.
Wide; figure 2, from Engel 2129-1, Chile, Prov. Magallanes, Pto. Cutter; figures 3-6,
from Engel 6425, Chile, Prov. Magallanes, between B. Bougainville and B. San Nicolas;
figures 7, 9, from Engel 6127, Chile, Prov. Magallanes, B. Borja; figure 8, from Engel
1856, Chile, Prov. Magallanes, R. San Juan; figures 10, 11, from Engel 4822, Chile,
Prov. Magallanes, Pto. Charrua.
60
FIG. 54. Clasmatocolea navistipula (Steph.) Grolle.
parceramosa.
= var. navistipula; O = var.
170
ENGEL: CLASMATOCOLEA 171
Feddes Repert. 82 (1):88. 1971. Original material: Chile, Prov.
Magallanes, I. Desolacion, (Pto. Angosto), Dusen (FH! — c. <f , G!).
Jamesoniella difficilis Steph. Kongl. Svenska Vetenskapsakad. Handl.
46 (9): 17. /. 6 a. 1911, syn. fide Grolle (1971). Original material:
Chile, Prov. Magallanes, Pto. Grappler, Skottsberg (BM!, UPS!); I.
Atalaya, Halle and/or Skottsberg (non vidi); S. Skyring, B. Rodriguez,
Halle & Skottsberg (hb. Grolle!, S! UPS!); Canal Gajardo, V. Inga,
Halle and/or Skottsberg (non vidi); B. Arauz, Halle and/or Skottsberg
(non vidi); W. end of L. Fagnano, Skottsberg (UPS!).
Lophocolea subcapillaris Steph. Kongl. Svenska Vetenskapsakad. Handl.
46 (9): 54. /. 18 e, f. 1911, syn. fide Engel (1978). Lectotype (fide
Engel, 1978): Chile, Prov. Magallanes, R. Fontaine, 1 March 1908,
Halle & Skottsberg 234 (UPS!).
Branching copious, in a narrow to broad flabellate pattern, Fru/lania-typG branching
absent; underleaves to 0.95 X stem width.
Differentiation. — Clasmatocolea navistipula is a rather close ally of C.
puccioana, which also has lateral-intercalary branches profusely developed
and branch underleaves reduced and inconspicuous. Plants of C. navisti-
pula may be differentiated from the latter by the following features:
(a) the reduced main axis underleaves that often cover only a small
fraction of stem tissue; (b) the frequently asymmetric segments of main
axis underleaves (i.e., underleaves with one segment smaller than the
other; see below); (c) the often unlimited growth of the lateral-intercalary
branches; (d) the branch underleaves that are considerably more reduced
than those of the short, lateral-intercalary branches of C. puccioana;9
(e) the conduplicately bilobed male bracts; (f) the whitish-light green color;
and (g) the narrower axis width (455-595 /x). Further, the flabellate
branching pattern developed in the typical variety is lacking in C. puc-
cioana.
The androecia are unique within the genus. The bracts are condupli-
cately bilobed, with the dorsal, basal portion of the bract with a small,
elliptic to narrowly ovate lobule attached to its adaxial surface for varying
lengths. Sometimes the lobule is rather inconspicuous and in such cases
should be searched for with care.
Notes. — 1. The main axis, the underleaves of which I have used as a key
character, may not be obvious due to copious branching, and the main
axis should then be carefully searched for and identified by the compara-
tively large underleaves (0.46-1.7 X the main axis width) and the slightly
9 Underleaves of branches of both C. puccioana and C. navistipula may best be
observed by removing the leaves and viewing the branch with the light microscope.
172 FIELDIANA: BOTANY
wider axis width. The main axis can often be recognized by its age, for as
the copious branches become better developed, the main axis becomes
brown and looses its leaves and underleaves.
2. Stephani (1911), in his description and drawings of Lophocolea sub-
capillaris, was clearly not referring to the syntypes from Insula Pacheco,
which are misdeterminations of Hepatostolonophora abnormis (Besch. &
Mass.) Engel & Schust., and a Rio Fontaine plant was therefore selected
as a lectotype.
Ecology. — The two varieties of C. navistipula are ecologically distinct;
the typical variety grows on rotted stumps or logs and over soil. In the
Patagonian Channel region on well-decayed logs in forests; less commonly
on Pilgerodendron bark and in thick masses of epiphytic vegetation over
tree trunks (see Engel, 1978).
Phytogeography. — Tierra del Fuego ; Patagonian Channels ; in Valdivian
region N. to 36° 50' S. (Prov. Concepcion) (see fig. 54).
Specimens seen.— CHILE. PROV. MAGALLANES: Sin. he. [probably
W. end of L. Fagnano], Skottsberg s.n. as syntype of/, difficilis (G); Cta.
Barrow, Skottsberg 231 as L. subcapillaris (UPS); Brunswick Pen. (cf.
Engel, 1978); E. side of B. Borja, Engel 6127 - c. sporo., 6175, 6205
(MSC); S. Skyring, B. Rodriguez, Halle & Skottsberg 220 (UPS); I.
Chatham, N. shore of B. Wide, Engel 5357 (MSC); Pto. Charrua, head of
inlet, Engel 4822 - c. rf1 (MSC); Pto. Alert, at head of fiord W. of M.
Markham, Engel 4961 B - c. <f , 4965 - c. sporo. + <? (MSC); Can.
Messier, Port Grappler, Skottsberg 220 (S, UPS). PROV. AISEN: N.
side of F. Tempano, Engel 4366 (MSC); Pto. Island, Engel 4251 — c.
sporo. + cf (MSC). PROV. VALDIVIA: V. Shoshuenco, Ruthsatz 44/18
(hb. Grolle). PROV. CONCEPCION: Concepcion, Dusen 155 as L.
puccioana (NY). Patagonia: Sin. loc., Skottsberg 585, 807 as syntype of
J. difficilis (G).
Clasmatocolea navistipula var. parceramosa Engel.
Clasmatocolea navistipula var. parceramosa Engel, Phytologia 41:311.
1979. Holotype: Chile, Prov. Magallanes, Brunswick Pen., ridge
between B. Bougainville and B. San Nicolas, 9 October 1969, Engel
6425 (MSC!).
Branching sparingly developed, non-flabellate, Frullania-type branching occasionally
present; underleaves to 1.7 X stem width.
Ecology. — The two varieties of C. navistipula occur in separate and
distinct ecological niches; var. parceramosa occurs in pools, pool margins
or Sphagnum bogs.
ENGEL: CLASMATOCOLEA 173
Phytogeography. — Southern South America at I. Grande de Tierra del
Fuego and southern Patagonian Channels (Brunswick Pen.) (see fig. 54).
Specimens seen.— ARGENTINA. TERR. TIERRA DEL FUEGO:
Tierra Major Valley, near junction of Ruta Nac. No. 3 with Harberton
road (Ruta J), 240 m., Crow 1472 (MSC). CHILE. PROV. MAGAL-
LANES: Brunswick Pen., near E. shore of La. El Parrillar, ca. 365 m.,
Engel 2105 (MSC); Brunswick Pen., ridge between B. Bougainville and
B. San Nicolas, ca. 155 m., Engel 6444 A - c. per. + & (MSC).
Subg. SQUAMICALYX Engel, subg. nov.
Folia caulina indivisa, Integra; cellulae foliorum trigonae graves, protuberantes,
saepe confluentes; amphigastria caulina utrinque connata; bracteolae involucraliae
utrinque connatae; perianthium habens magnas squamas.
Type species: Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle.
Leaves undivided, entire; trigones massive, protuberant, often confluent; underleaves
connate on both sides, moderately to distinctly cupulate, undivided; bracteoles con-
nate with bracts on both sides; perianth with large scales.
Distribution. — Amphipacific; New Zealand and its subantarctic islands,
Tasmania, southern South America.
Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle. Figures 55, 56.
Jungermannia notophylla Hook. f. & Tayl. London J. Bot. 3:376. 1844.
Chiloscyphus notophyllus (Hook. f. & Tayl.) G. L. & N. Syn. Hep.
710. 1847. Lophocolea notophylla (Hook. f. & Tayl.) Hodgs. in Martin,
Trans. & Proc. Roy. Soc. New Zealand 78:495. 1940. Clasmatocolea
notophylla (Hook. f. & Tayl.) Grolle, J. Jap. Bot. 41:228. 1966.
Original material: Auckland Is., Hooker (FH— hb. Steph.!, L!, LD!,
NY!, S!).
Lophocolea tumida Steph. Bull. Herb. Boissier 6 (9): 791. 1906 (= Spec.
Hep. 3:91), syn. fide Grolle (1966). Holotype: Tasmania, Mt. La
Perouse, Oldfield (G! — c. per.; isotypes: FH! — c. per., JE! — c.
young per.).
Chiloscyphus cordifolius Rodw. Pap. & Proc. Roy. Soc. Tasmania
1915:106. 1916 (seors. 31 December 1915) (see also vol. 2 of Tas-
manian Bryophyta, p. 54. 1916), syn. fide Grolle (1966). Original
material: Tasmania, Adamson's P., 1,100 m., December 1913,
Rodway(hb. Grolle!, HO!).
Plants soft, fleshy, the leaves and underleaves fragile, easily torn; plants suberect,
mixed with other bryophytes or in rather dense pure tufts, whitish green to yellow-
brown to brown; axes to 8.5 cm. tall, 1.5 mm. wide, the apices usually swollen.
FIG. 55. Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle. 1, Portion of main axis,
ventral-lateral view. 2, Mature perianth showing scales (only the associated bracteole
shown), dorsal-lateral view. 3, Main axis, ventral view, showing underleaf and leaf
insertion. 4, 5, Leaves. 6-7, Underleaves, dorsal view. 8, Stem, cross section. 9, Median
leaf cells. 10, Main axis, lateral view, leaves of one side and one underleaf removed.
Figures 1, 4-9, from type of C. notophylla, Hooker, Auckland Is. (L); figure 2, from
isotype of Lophocolea tumida, Oldfield, Tasmania (FH); figures 3, 10, from Scott 1075,
Tasmania, Cradle Mt.
174
ENGEL: CLASMATOCOLEA
175
FIG. 56. Clasmatocolea notophylla (Hook. f. & Tayl.) Grolle.
Branches sporadic, of lateral- and ventral-intercalary type, the lateral-intercalary
branches ventrally displaced, between ventral portion of leaf and edge of underleaf.
Stems 6-10 cells high, cortex in a single row of moderately thick-walled cells smaller
than the medullary cells, the exposed wall of cortex very thickened; medullary cell walls
moderately thickened; endophytic hyphae absent. Rhizoids in fascicles from stem near
underleaf base.
Leaves with insertion slightly to distinctly recurved at ventral end; leaves usually
connivent or nearly so dorsally, closely imbricate, conchiform concave, very broadly
ovate to reniform, dorsal portion of leaf becoming moderately reflexed; apex broadly
rounded, undivided, often slightly to distinctly inflexed, the apex and margins entire,
although the margins sometimes with several slime papillae; dorsal margin broadly
rounded, plane to sinuous, rather long decurrent; ventral margin inflexed, broadly
rounded, basal portion =*= auriculate, extending ventrally beyond stem and in contact
with spreading underleaf, occasionally in contact with ventral margin of opposite leaf.
176 FIELDIANA: BOTANY
Leaf cells with massive, nodular or protuberant, irregularly triangular to rectangular
trigones that are confluent or separated by narrow thin-walled places, the cell lumen
bounded mostly by the massive trigones; median leaf cells 1 8-38(-42) ju wide, (23-)29-
49(-53) M long; cuticle smooth or roughened and then appearing finely granular.
Underleaves 3.1-3.6 X stem width, connate on both sides with ventral-basal adaxial
portion of leaf lamina or with ventral margin of leaf, the attachment hidden by ventral-
basal portion of leaves; underleaves moderately spreading, approximate to imbricate,
moderately to distinctly cupulate, =t orbicular to oblate, basal portion abruptly convex
(ventral view), and often with a flange of tissue marginal to insertion; apex undivided and
broadly rounded to truncate to broadly refuse, the apices with 2 slime papillae or with
(l-)2 teeth each terminating in a slime papilla; margins of the lamina with a few slime
papillae and/or 1-4 few-celled teeth each terminating in a slime papilla.
Plants ?dioecious; androecia not seen. Gynoecia (only 2 mature gynoecia seen) on
main axis; subfloral innovations from below bracteole of second series or absent;
bracts in 3 series, those of innermost series deeply concave, with a deeply concave
bulge in median-basal portion; apices undivided and broadly rounded, the apices and
margins entire. Bracteole (of mature gynoecium) of innermost series connate for ca. 0.3
its length on 1 side, short-connate on other side or (in juvenile gynoecium) 0.3 connate
with bracts on both sides, deeply convex (ventral view); lamina margins sparingly
dentate. Perianth with basal portion with numerous, large, irregular, commonly obovate,
sometimes bilobed scales attached to perianth for varying lengths, the scales occasionally
attached to one another; perianth subclavate, ± inflated and =•= terete in basal portion,
the upper portion somewhat bilaterally compressed, expanding slightly toward mouth;
perianth mouth =*= bilaterally compressed due to infolded lobes, the lobes broadly
rounded, incurved, entire. Seta present, but old, collapsed, and without a capsule.
Differentiation. — This species bears a superficial resemblance to some
plants of the hygrophylous facies of C. humilis. Clasmatocolea notophylla
may be distinguished by the following complex of characters : (a) leaf cells
with massive, nodular or protuberant trigones that are confluent or
separated by narrow thin-walled places; (b) leaves with dorsal margins
rounded; (c) underleaves connate on both sides, and frequently with the
ventral-basal adaxial portion of the leaf lamina; (d) underleaves consis-
tently undivided; (e) bracteoles connate on both sides, and on one side
(or both sides in one observed immature gynoecium) for ca. 0.3 its length;
(f) basal portion of perianth with numerous large (some extending to ca.
0.5 perianth length) obovate scales that are attached to the perianth for
varying lengths.10 In view of the fact that characters listed in (a), (e), and
(f) do not occur in any other Clasmatocolea taxon, C. notophylla stands
quite isolated and should therefore not be confused with any other
member of the genus.
Clasmatocolea notophylla shares some characters with C. tjiwideiensis
of Java, which also has (a) deeply convex (ventral view) underleaves that
are at least 3 X the stem width and sometimes undivided ; (b) underleaves
10 Unfortunately only two mature gynoecia were available for study. Characters (e)
and (f) thus require verification with the study of more copious material.
ENGEL: CLASMATOCOLEA 177
at times connate on both sides, and on one side, at least, with the ventral-
basal, adaxial portion of the leaf lamina; (c) the basal portion of the
ventral margins of leaves extending ventrally beyond the stem. However,
the highly unique characters of the gynoecium of C. notophylla isolate
this species and negate any real, close relationship to C. tjiwideiensis.
Notes. — 1. There has been considerable confusion regarding usage of
the name Clasmatocolea "Lophocolea" notophylla in New Zealand. Some
of the confusion can be traced to Hodgson (1953) and determinations by
her. Hodgson's concept of "Lophocolea" notophylla can be explained by
her statement (1953, p. 338), "Although there is no specimen of Taylor's
type in either the Hooker or Mitten Herbarium, with which to compare
the mainland plant, Lophocolea okaritana St., I believe that they are the
same species. . . ." In Hodgson (1953) the description and all observed
plants referred to by her as Lophocolea notophylla are actually Lophocolea
gunniana Nees (with one exception: the specimen from Regina Valley, leg.
Teague is Clasmatocolea cf. vermicular is). Lophocolea okaritana Steph.
(Spec. Hep. 3:85. 1906, type: New Zealand, Okarito, Kirk 592 G!), which
is listed as a synonym of C. notophylla in Hamlin (1972), is a synonym of
Lophocolea gunniana (fide Hamlin, 1973).
The confusion is compounded by Hodgson's (1953, p. 338) comment,
"Judging by the drawing in Bastow's Plates of Tasmanian Hepatics, Loph.
austrigena (Tayl.) Syn. Hep., of Tasmania, appears to be the same thing."
This may explain why many of the collections of Lophocolea gunniana
that I have studied are labeled Lophocolea austrigena, and further, several
had originally been determined as Lophocolea notophylla. Lophocolea
gunniana and L. austrigena (type — Cabo de Hornos, Chile) are actually
very closely related, but based upon my studies to present, L. austrigena is
restricted to Tristan da Cunha, Inaccessible I., the Falkland Is., and
southern South America. Hodgson (1962) alludes to the error as she
states for Lophocolea austrigena (p. 1 1 1), "Wrongly listed as Lophocolea
notophylla (Hodgson, 1953)."
2. There seems to be confusion regarding pagination of several of the
Rodway papers; the Rodway species I am citing here was published in the
Papers and Proceedings of the Royal Society of Tasmania. The title page of
the article bears the phrase "Issued separately (date)." Chiloscyphus
cordifolius Rodw., for example, appears on p. 106 of the Papers and Pro-
ceedings for the year 1915 (fascicle issued 24 February 1916). The title
page of the article, however, reads "Issued separately 31st December,
1915." The description was then republished in Pap. & Proc. Roy. Soc.
Tasmania for the year 1916, the fascicle of which appeared on 19 February
1917. This Rodway paper was also issued, separately paged, on 30 August
178 FIELDIANA: BOTANY
1916, appearing as vol. 2 of Tasmanian Bryophyta. The latter has been
widely distributed and is commonly cited.
Ecology. — In New Zealand in Dacrydium scrub (240 m.) or in Sphagnum-
Hypolaena-Dacrydium bogs (at 915 m. and associated with Anastro-
phyllum schismoides and Tetracymbaliella cymbaliferd) or on wet, peat-
covered, south-facing ledges of cliff faces (sea level). In Tasmania it grows
very loosely over ground or bryophyte-covered logs in Nothofagus-
Eucryphia, Nothofagus cunninghamii, or Nothofagus-Athrotaxis forests
(400-915 m.). It occasionally occurs over wet, peaty soil in subalpine
seepage areas under protective cover of Diselma archeri (1,225 m.). In
southern South America the species is restricted to the very wet, western
fringe of the Patagonian Channels in Prov. Magallanes and Prov. Aisen.
It occurs in pools and along streams in moorland areas and (rarely) on
the floor of mossy forests.
Phytogeography. — Amphipacific temperate; Auckland Is.; New Zealand,
South I.; Tasmania (400-1,225 m.); southern South America (Patagonian
Channels N. to 48° 04' S., Prov. Magallanes, Prov. Aisen) (see fig. 56).
Specimens seen.— NEW ZEALAND. SOUTH ISLAND. SOUTH-
LAND: Fiordland, Secretary I., ca. 240 m., Baylis s.n. as Lophocolea
austrigena (CHR); ibid., Baylis s.n. — c. per. (hb. Grolle); Fiordland
Natl. Park, Milford Sound, foot of Bowen Falls, Schuster 55546b (hb.
Schuster). WESTLAND: Omoeroa R., S. of Franz Josef, towards Weheka
Hills, Schuster 67-241b (hb. Schuster). WESTLAND/NELSON: Paparoa
Range, 915 m., Helms 244 as Leioscyphus turgescens (CHR). TASMANIA:
Adamsons P. Track, A. Ratkowsky 78/73 (F); near Manuka Flat on trail
to Adamsons P., ca. 300-400 m., Norris 26941 (F); plateau region E. and
SE of Mt. Eliza summit, E. of L. Pedder, ca. 1,225 m., Engel 13708 (F);
upper reaches of S. facing slope of Sentinel Range, between Lakes Gordon
and Pedder, 700 m., Engel 15114 (F); L. Gordon, Walters 79/17 (F); Mt.
Arrowsmith, on Lyell Hwy., 2.7 mi. W. of King William Saddle, W. of
Derwent Bridge, Surprise Valley, ca. 600 m., Schuster 50372 (hb. Schuster);
Cradle Mt.-L. St. Clair Natl. Park, Pine Valley, Cephissus Falls, NNW of
L. St. Clair, 850 m., Engel 14252 (F); Allans Cr., NE slope of Mt. Darwin,
off Kelly Basin Road (Crotty Track), 410 m., Engel 16529, 16584 (F);
Cradle Mt. Region, Cradle Mt., 915 m., Scott 1075 (hb. Monash Univ.);
Cradle Mt. Region, Dove L. Track, A. Ratkowsky 79/16 (F); Cradle Mt.
Region, Weindorfers Forest, along track from Waldheim Chalet to
Hounslow Heath Track, 975 m., Engel 14050 (F). CHILE. PROV.
MAGALLANES: Pto. Alert, Engel 5036, 5061 (MSC). PROV. AISEN:
Pto. Island, Engel 4268 A - c. young per., 4323 A (MSC).
ENGEL: CLASMATOCOLEA 179
Clasmatocolea verrucosa Engel. Figures 57, 58.
Clasmatocolea verrucosa Engel, Bryologist 83:220. /. 7-79. 1980.
Holotype: Tasmania, E. slope of Black Bluff just below summit, S.
of Burnie, 1,250 m., Engel 15764 (F; isotype: HO).
Plants soft, fleshy, the leaves and under/eaves fragile, easily torn; plants loosely creep-
ing to suberect, in rather dense, pure tufts, yellow-brown, becoming brownish with
age, highly nitid when dry; axes to 3 mm. wide.
Branches rather common, mostly o/Frullania type, occasionally of lateral-intercalary
type, the lateral-intercalary branches ventrally displaced, between ventral portion of
leaf and edge of underleaf ; ventral-intercalary branches very rare (only a few examples
seen).
Stems narrow for plant size, 11-12 cells high, the cortex poorly differentiated, in a
single row of slightly smaller cells; both the cortical and medullary cells moderately
thick walled, but the exposed wall of cortex very thickened ; cuticle smooth. Rhizoids
in tight fascicles from stem near underleaf base, the rhizoids hyaline and often adhering
to underleaf surface, the tips thick walled and often highly branched but nonseptate.
Leaves with insertion not or very slightly recurved at ventral end; leaves erect (but
not connivent dorsally) to moderately spreading, moderately conchiform concave,
broadly ovate to oblate; apex broadly rounded, undivided, often sharply inflexed,
lending a deeply concave, channeled appearance to that part of the leaf; apex often
repand, entire or with armature more sparing and smaller than that of margins; dorsal
and ventral margins armed with several irregular, often sharp teeth, the margins some-
times with a small, accessory, armed lobe near the base; dorsal margin straight or
slightly rounded, undulate, sometimes reflexed; ventral margin broadly rounded, but
often straight near base.
Leaf cells with massive, coarse, irregularly triangular to rectangular trigones that are
confluent or separated by narrow thin-walled places, the cell lumen bounded mostly by
the massive trigones; median leaf cells 24-30(-36) M wide, 31-41 ^ long; leaf cells on
their inner and outer surfaces each with a large, domelike, prominent, hyaline verruca,
the cells of the apex and margins of the leaves each with similar protuberances but with
them peripherally extended beyond the outer cell wall.
Underleaves reniform, in width 2.2-2.9 X their length (when flattened) and 4.4-5.1 X
stem width, connate on both sides, slightly spreading, imbricate, distinctly concave and
with margins appressed to leaves; apex undivided and broadly rounded to broadly refuse,
the apices with a few slime papillae along with a few teeth that commonly terminate
in a slime papilla, the apices sometimes edentate; margins of the lamina sometimes
inflexed, with a few slime papillae plus a few teeth and often laciniae, the armature
commonly terminating in a slime papilla.
Plants dioecious; androecia intercalary on main axis or on rather long Frullania- or
(occasionally) lateral-intercalary type branches; bracts with basal portion strongly
saccate, the entire bract strongly concave, and with apices appressed to bract immedi-
ately above; lobule margin incurved to involute, broadly rounded, sparingly dentate-
laciniate; antheridia solitary, the stalk uniseriate.
Gynoecia (only immature gynoecia seen) on main axes or innovations (plants some-
times repeatedly producing innovations each forming a perianth); subfioral innovations,
FIG. 57. Clasmatocolea verrucosa Engel. 1, Leading axis with Frullania-type branch.
2, Cross section through median portion of leaf. 3, Median leaf cells showing cuticular
verrucae. 4. Cells of ventral margin of leaf. 5, Cells of leaf apex. 6, Stem with portion
of leaf base, cross section. 7, Rhizoid tip. 8, Perianth showing scales, dorsal-lateral view
(only the associated innermost bracteole shown). 9, Bract of innermost series (not
flattened). All from holotype.
180
FIG. 58. Clasmatocolea verrucosa Engel. 1, Portion of main axis, ventral view (LIB =
lateral-intercalary branch). 2, Portion of main axis, dorsal view. 3, 5, 7, 8, Leaves of
main axis (not flattened). 4, 9, 10, Underleaves of main axis (flattened). 6, Antheridial
stalk. All from holotype.
181
182 FIELDIANA: BOTANY
when present, from below bract or bracteole of innermost or second series; bracts in
3 series, those of innermost series often closely appressed to perianth, the bracts rather
deeply concave, with a deep, vertically elongate, concave bulge in median-basal or
dorsal-basal portion; apex undivided, broadly rounded, irregularly dentate; lamina
margins rather abundantly and sharply dentate-laciniate. Bracteoles of innermost
series free or short connate on 1 side, rather convex (ventral view); apex undivided, with
several irregular, small teeth; lamina margins with several sharp teeth. Perianth with
basal portion with several large, vertical, irregular, ciliate to laciniate scales or lamellae
attached to perianth for varying lengths; perianth broad and stoutly ovate, inflated and
weakly trigonous in the basal portion (the angles blunt and broadly rounded), the upper
portion somewhat bilaterally compressed, slightly narrowing toward mouth; perianth
lobes broadly rounded, armed with several irregular, often sharp teeth, the ventral
lobe somewhat smaller and more narrowly rounded, the ventral lobe somewhat longi-
tudinally folded, rendering the perianth distally bilaterally compressed; perianth cells
with papillae similar to those of the leaves.
Sporophyte not seen.
Differentiation. — The manuscript of this paper was originally completed
and submitted with C. notophylla treated as an isolated taxon and placed
in its own subgenus. While subsequently working on my Tasmanian
collections, I became aware of the remarkable C. verrucosa, which, in all
cases, was mistaken for C. notophylla in the field. This field misidentifica-
tion is not surprising, for the two taxa have a similar facies and share the
following features: (a) plant size, although C. verrucosa is a bit larger;
(b) leaf shape, with apices undivided and broadly rounded; (c) leaf cells
with massive, coarse trigones that are confluent or separated by narrow
thin-walled places; (d) underleaves connate with leaves on both sides;
(e) underleaves large, deeply concave, and with apices undivided and
broadly rounded to broadly retuse; and (f) perianth with scales of varying
sizes. However, C. verrucosa may be immediately distinguished by (a) the
huge cuticular verrucae, one each per cell; (b) the predominance of
Frullania-type branching; (c) the armature of the dorsal and ventral
margins of the leaf; (d) the straight or at most slightly rounded dorsal
margin of the leaf; (e) the reniform underleaves that are 2.2-2.9 X wider
than high and 4.4-5.1 X the stem width; and (f) the armed perianth scales
and dentate perianth lobes.
The only other members of the genus with cuticular excrescences are the
unrelated C. strongylophylla and C. tjiwideiensis (both Subg. Clasmato-
coled), both of which have a papillose cuticle.
Note. — With the use of depth focusing, the cuticular verrucae may be
observed, one each per cell, oriented directly above the cell lumen and
not over the cell wall. This is the case in all leaf cells except those of the
marginal row. The marginal row has cuticular protuberances similar to
those of the interior cells, but differs in having the verrucae peripherally
ENGEL: CLASMATOCOLEA 183
extended beyond the outer cell wall. This feature lends a definite warty
appearance to the leaf, which is useful in identifying the plants. The
verrucae may be readily observed in profile on the leaf apices and margins,
utilizing the dissecting microscope.
Ecology-Phytogeography, — This subalpine-alpine species, found between
1,000-1.300 m., is endemic to Tasmania. It typically grows along creek
banks or pool margins, as well as under protective cover of rock ledges
or shrubs. One collection (Norn's 27866) occurred in the upper limits of a
Nothofagus forest.
Specimens seen.— TASMANIA: Mt. Field Natl. Park, Mt. Mawson
Plateau, above and W. of L. Dobson and S. of L. Seal, 1,250 m., Engel
13089, 14327 (F); Mt. Field Natl. Park, Rodway Range, between Rodway
Ski Tow and K Col area. 1,240-1,310 m., Engel 14416 - c. <? (F); W.
side of Mt. Eliza, 800-1,200 m.. Norris 31007 (F); L. St. Clair Region,
trail from L. Solitude to Little Hugel, 1,000-1,100 m., Norris 27866 (F);
Cradle Mt.-L. St. Clair Natl. Park, The Labyrinth, NNW of L. St. Clair,
1,130 m., Engel 14272 A (F); Cradle Mt. Region, Plateau Cr. area, between
Cradle Plateau and Marions Lookout, 1,250 m., Engel 13966, 13979 - c.
young per. (F); E. slope of Black Bluff just below summit, S. of Burnie,
1,250 m., Engel 15764 - c. per. (F).
Subg. PLICATICALYX Engel, subg. nov.
Folia caulina indivisa, parcc dcntata; amphigastria caulina cucullata. basi auriculata,
habcntia unum lobum basalc partc cxtcnsum ultra caulis dorsalcm paginam; androecia
latiora quam sterile parte axis; j bracteae lobuli l-3-laciniati ad lobati; perianthia
valde ovoidea, apice 4-5-plicato. ore contracto, lobi copiose dentati ad laciniatos;
sporae 24-27(-30) /j, cum anguste conicis prominentiis.
Type species: Clasmatocolea cucullistipula (Steph.) Grolle.
Leaves undivided, sparingly dentate; undcrlcavcs cucullate. auriculate at base, with
one lobe in basal portion of underleaf which extends beyond dorsal surface of stem;
androecia wider than sterile parts of axis; ^ bract lobules l-3-laciniate to lobate; peri-
anth stoutly ovoid, apex 4-5-plicate. the mouth contracted, the lobes copiously dentate
to laciniate; spores 24-27(-30) ju, the cxine with narrowly conical projections.
Distribution. — Southern South America.
Clasmatocolea cucullistipula (Steph.) Grolle. Figures 59-61, Plates 28, 29.
Lophocolea cucullistipula Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Hand!. 26 (111, 6): 37. 1900. Clasmatocolea cucullistipula (Steph.)
Grolle, Rev. Bryol. Lichenol. 29:71. 1960. Lectotype (nov.): Chile,
Prov. Chiloe, I. Guaitecas, (Melinca), April 1896, Dusen 376, ex hb.
Dusen (S!).
I ^XOdOQ
• — 'G>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. <? (MSC); SE point of I. Wil-
liams, Engel 4488 - c. &, 4496 - c. per. (MSC). PROV. AISEN: Pto.
Island, Engel 4293 — c. sporo. (MSC); Pen. de Taitao, B. San Rafael,
Gusinde s.n. (JE); R. Aisen, Dusen s.n. as syntype of L. cucullistipula
(NY); ibid., Dusen 538 as syntype of L. homomalla (G); I. Magdalena,
Schwabe 33a as L. filiformis — c. tf (JE); ibid., 6 m., Schwabe 33/b as C.
fulvella — c. per. (hb. Grolle); Pto. Puyuhuapi, Schwabe 4/b (JE); ibid.,
Schwabe 9//-7//as C. fulvella (hb. Grolle); ibid., Schwabe 23 /b p.p. as L.
navistipula (JE); ibid., Schwabe 50/b (JE); ibid., R. Obscuro, Schwabe
21 /a — c. tf (hb. Grolle); ibid., Schwabe 2J/b as C. fulvella — c. per.
(JE). PROV. CHILOE: I. Guaitecas, Dusen s.n. as syntype of L. cucul-
listipula (G — c. c?» LD — c. per., NY, O — c. per.); ibid., Dusen 343
as syntype of L. cucullistipula (FH, O); ibid., Halle s.n. — c. per. (G);
ibid., Bo. Chica, Dusen 367 (NY); I. Chiloe, 1.3 km. by road N. of junc-
tion of Ruta 5 and road to Delcahue, ca. 100 m., Engel 12175 (F); I.
Chiloe, Aguas Buenas area, 4.7 km. E. along Aguas Buenas road from
Ancud-Quemchi road, ca. 100 m., Engel 12215 - c. per. (F). PROV.
LLANQUIHUE: L. Todos los Santos, Cayutue, Wolff htigel s.n. as L.
cucullifolia (JE, S). PROV. OSORNO: Refugio Antillanca, 1,160m.,
Engel 3903 (MSC); around L. Toro on road to Refugio Antillanca, 855 m.,
Engel 3999 (MSC). PROV. VALDIVIA: E. slope of Cord. Pelada, E. of
El Mirador, between La Union and Pta. Hueicolla, 740 m., Engel 12478
(F); Cord. Pelada, summit of El Mirador, between La Union and Pta.
Hueicolla, 1,000 m., Engel 12398 (F); W. slope of Cord. Pelada, W. of
El Mirador, between La Union and Pta. Hueicolla, 680 m., Engel 12331 —
c. per. (F); Corral [Quitaluto], Hosseus 114, 613 ex p., 643, 657 ex p., 668A
(JE); ibid., Krause s.n. as Jungermannia moniliformis Angstr., nom. hb. (hb.
Vana); C. Lungoico, Schwabe s.n. (hb. Grolle). PROV. CAUTIN: Villarica,
Ruthsatz s.n. (JE). Patagonia: (All sin. loc. and all probable syntypes of
L. cucullistipula), Dusen 304 (G), Dusen 343 (FH, G), Dusen 367 - c. <?
(G); (sin. loc., probable syntype of L. homomalla), Dusen 376 (FH, G).
XI. EXCLUDENDA
1. Clasmatocolea abnormis (Besch. & Mass.) Grolle.
Leioscyphus (?) abnormis Besch. & Mass. Bull. Mens. Soc. Linn. Paris
1:629. 1886. Lophocolea abnormis (Besch. & Mass.) Steph. Spec.
Hep. 3:62. 1906. Clasmatocolea abnormis (Besch. & Mass.) Grolle,
Rev. Bryol. Lichenol. 29:71. 1960. Stolonophora abnormis (Besch. &
Mass.) Engel & Schust. Fieldiana, Bot. 36:114. 1975. Hepatostolono-
phora abnormis (Besch. & Mass.) Engel & Schust. in Engel, J. Hattori
ENGEL: CLASMATOCOLEA 191
Hot. Lab. 46:94. 1979. Type: Chile, Prov. Magallanes, I. Hoste,
Hyades (FH!, G!, hb. Grolle!) (c. per.).
Jamesoniella gracilis Gola, Nuovo Giorn. Bot. Ital. II. 29:164. 1923,
syn. fide Engel (1978). Original material: Chile, Prov. Magallanes,
B. Ainsworth, (Ghiacciaio), 27 February 1913, De Gasperi s.n. (FI!).
( Lophocoleaceae).
This species belongs in Hepatostolonophora; see Engel & Schuster
(1975) and Engel (1979b).
2. Clasmatocolea acutiloba Schiffn.
Clasmatocolea acutiloba Schiffn. in Schiffner & S. Arnell, Osterr. Akad.
Wiss., Math.-Naturwiss. Kl., Denkschr. 111:63. pi. 10, f. 112 a-g.
1964 = Cylindrocolea acutiloba (Schiffn.) Engel, comb. nov. Holo-
type: Brazil, Estado de Sao Paulo, "prope Lapa ad flumen Tiefe
juxta Sao Paulo," 800 m., Schiffner 2259 (W!). (Cephaloziellaceae).
This species appears to be an ally of Cylindrocolea rhizantha (Mont.)
Schust.
3. Clasmatocolea cuneifolia (Hook.) Spruce.
Jungermannia cuneifolia Hook. Brit. Jungerm. pi. 64. 1814. My I ins
cuneifolius (Hook.) S. Gray, Nat. Arr. Brit. PI. 1:694. 1821. Aplozia
cuneifolia (Hook.) Dum. Bull. Soc. Roy. Bot. Belgique 13:55. 1874.
Coleochila cuneifolia (Hook.) Dum. Bull. Soc. Roy. Bot. Belgique
13:106. 1874. Clasmatocolea cuneifolia (Hook.) Spruce, Trans. &
Proc. Bot. Soc. Edinburgh 15:440. 1885. Leptoscyphus cuneifolius
(Hook.) Mitt. Hooker's J. Bot. Kew Gard. Misc. 3:358. 1851.
Leioscyphus cuneifolius (Hook.) Steph. Bull. Herb. Boissier 6 (3): 218.
1906 (= Spec. Hep. 3:18). Anomylia cuneifolia (Hook.) Schust.
Amer. Midi. Naturalist 62:53. 1959. Original material: Ireland,
Bantry, Hutchins (non vidi). (Lophocoleaceae).
See Grolle (1962) for taxonomic treatment and illustrations and Grolle
(1956, p. 294).
4. Clasmatocolea doellingeri (Nees) Steph.
Jungermannia doellingeri Nees in G. L. & N. Syn. Hep. 104. 1844.
Clasmatocolea doellingeri (Nees) Steph. Bull. Herb. Boissier 6
(5): 391. 1906 (= Spec. Hep. 3:48 [sic, in errore pro 47]). Chonecolea
doellingeri (Nees) Grolle, Rev. Bryol. Lichenol. 25:295. 1956. Origi-
nal material: "In Brasilia, super truncos, in caespite Octoblephari
albidi (Hb. N. ex Hb. Doellinger.). Vidi etiam in Herb. Hampeano,
a Siewers e Brasilia adlata specimina." (non vidi). (Chonecoleaceae).
192 FIELDIANA: BOTANY
See Grolle (1956) and Schuster (1958) for taxonomic treatments,
including illustrations, and Grolle (1960, p. 87) for discussion of taxonomy
and type.
Specimens seen.— ARGENTINA. PROV. BUENOS AIRES: Sa. de la
Ventana, 540 m., Roivainen 369b (F, H). BRAZIL. San Cristobol, sin.
coll. - c. sporo. (M). ESTADO DE SAO PAULO: Municipio de Sao
Paulo, Institute de Botanica, Vital 1283 (F, SP); Municipio de Itirapina,
Vital 777, 136, 2060 (F, SP); Municipio de Pirassununga, Vital 1754 (F,
SP); Municipio de Rancharia, Vital 2577 (F, SP). UNITED STATES.
FLORIDA: Sanford, Rapp s.n. (F, L, M — c. sporo., TENN); Marion
Co., Ocala Natl. Forest, Juniper Springs, Schuster 31520 (TENN).
5. Clasmatocolea exigua Steph.
Clasmatocolea exigua Steph. Bull. Herb. Boissier 6 (5): 391. 1906
[= Spec. Hep. 3:48 (sic, in error e pro 47)]. Original material: North
America, Louisiana, Langlois (FH!) = Cylindrocolea rhizantha
(Mont.) Schust. Nova Hedwigia 22:175. 1972 (1971), syn. fide
Fulford (1976). (Cephaloziellaceae).
See Grolle (1956, p. 298) and Schuster (1958).
6. Clasmatocolea fiordlandiae (Hodgs.) Grolle.
Clasmatocolea fiordlandiae (Hodgs.) Grolle, Rev. Bryol. Lichenol.
29:72. 1960 = Lophocolea fiordlandiae Hodgs. Trans. Roy. Soc. New
Zealand 80:340./. 75. 1953. Original material: New Zealand, South
Island, Fiordland, Caswell Sound, 27 March 1949, Zotov 5235
(CHR! - c. (f ; hb. Grolle!). (Lophocoleaceae).
I believe the rightful place of this species is in the genus Lophocolea,
because the leaves are plane and not adaxially concave. The leaves,
however, sometimes have apices that are inflexed so that the leaves, then,
appear concave in the distal portions. Further, the species is more closely
related to taxa in Lophocolea rather than to any taxon assigned to Clas-
matocolea.
Calyptrocolea perssonii Schust. was erroneously placed in the synonymy
of "Clasmatocolea" fiordlandiae by Grolle (1972a). See Engel (1979b)
where the name is in varietal rank — Hepatostolonophora rotata var.
perssonii (Schust.) Engel; see also Engel & Schuster (1975).
7. Clasmatocolea fragillima Spruce.
Clasmatocolea fragillima Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15:440. 1885. Original material: Ecuador, M. Tunguragua, Spruce
(non vidi) = Leptoscyphus liebmanianus (Lindenb. & Gott.) Mitt.
ENGEL: CLASMATOCOLEA 193
Hooker's J. Bot. Kew Gard. Misc. 3:358. 1851, syn. fide Grolle
(1960). (Lophocoleaceae).
See Grolle (1960, p. 86) and Fulford (1976, p. 511), who synonymizes
the species with Leptoscyphus porphyrius (Nees) Grolle.
8. Clasmatocolea georgiensis (Gott.) Grolle.
Lophocolea georgiensis Gott. Ergebn. Deutsch. Polar-Exped. 2 (16): 453.
pi. 3-4. 1890. Clasmatocolea georgiensis (Gott.) Grolle, Brit. Antarc.
Surv. Bull. 28:86. 1972. Evansianthus georgiensis (Gott.) Schust. &
Engel, Bryologist 76:518. 1973. Lectotype (fide Grolle, 1972b): South
Georgia, 10 May 1883, Will 11 (M!). (Lophocoleaceae).
For treatment of this species, including illustrations, see Schuster &
Engel (1973).
9. Clasmatocolea innovata Herz.
Clasmatocolea innovata Herz. in Herzog & Noguchi, J. Hattori Bot. Lab.
14:39./ 7 a-g. 1955 = Phragmatocolea innovata (Herz.) Grolle, Rev.
Bryol. Lichenol. 25:299. 1956. Holotype: Taiwan, Karobetsu, 100 m.,
Schwabe 20 (non vidi}. ( ? Jungermanniaceae).
Grolle (1956), who provides illustrations and a discussion of the
taxonomy of the species, places the taxon provisionally in the Jungerman-
niaceae (s. str.). However, Schuster (1958, p. 16) and Varta (1975, p. 365)
express reservations regarding this placement.
10. Clasmatocolea navicularis (Steph.) Grolle.
Lophocolea navicularis Steph. Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 43. 1900. Clasmatocolea navicularis (Steph.) Grolle,
Rev. Bryol. Lichenol. 29:72. 1960. Xenocephalozia navicularis (Steph.)
Schust. Nova Hedwigia 10:25. 1965. Lectotype (fide Grolle, 1966):
Chile, Prov. Aisen, R. Aisen Valley, Dusen (non vidi).
Jungermannia chilensis Mont. Ann. Sci. Nat. Bot. III. 4:349. 1845 non
J. chilensis Lehm. Nov. Minus Cogn. Stir. Pug. 6:36. 1834 ( =
Porella), syn. fide Grolle (1966). Microlejeunea chilensis (Mont.)
Steph. Spec. Hep. 5:835. 1915. Original material: Chile, (Prov.
Chiloe), without specific locality, Gay (BM!, G!).
Drepanolejeunea stephaniana Mass. Atti Accad. Sci. Med. Nat. Ferrara
80: 9. / 1-4. 1906, syn. fide Grolle (1966). Original material : Patagonia,
Poyo-huapi, Spegazzini (non vidi).
Cephalozia trisetosa Steph. Bih. Kongl. Svenska Vetensk.-Akad. Handl.
26 (III, 6):48. 1900, nom. nud. pro syn. (Lophocoleaceae).
194 FIELDIANA: BOTANY
Differentiation. — The nearest relative of X. navicularis appears to be
the at least occasionally corticolous Clasmatocolea trachyopa, which also
has (a) distinctly divided leaves, with the ventral portion incurved and
flattened parallel to the ventral stem surface, lending a pouchlike con-
cavity in the ventral portion of the leaf; and (b) strongly trigonous peri-
anths with bifid lobes. Clasmatocolea trachyopa is distinct by the strongly
oblique, often highly complex and ornamented leaves, with the insertion
broad. Xenocephalozia, on the other hand, has transverse to weakly
incubously inserted, entire or sparingly armed leaves with the insertion
narrow. It is of interest, however, to note that depauperate axes of C.
trachyopa not only have subtransversely inserted leaves with a narrow
insertion, but lack some of the intricate ornamentation typical of well-
developed facies of the species; these plants look strikingly like the
spinose-leafed facies of Xenocephalozia.
Schuster (1965a, 1965b) thought Xenocephalozia belonged near Clasmat-
ocolea, although in the latter paper he emphasized the isolation of Xeno-
cephalozia. Grolle (1966) regarded the plant as a Clasmatocolea, but
thought the species unique enough to place it in its own subgenus — Subg.
Schusterocolea Grolle. Perhaps Xenocephalozia arose from an ancestor
much like C. trachyopa through alteration of leaf insertion, loss of orna-
mentation, and reduction in plant size. I believe, however, that the narrow,
transverse to weakly incubous leaf insertion of Xenocephalozia is a funda-
mental difference from Clasmatocolea and necessitates an independent
generic status.
Variation. — Leaves of weak forms of this species tend to be entire
margined and shallowly bilobed. The segments are short and rather wide-
triangular, with the apex narrowly rounded or sometimes obtuse, and
without development of a uniseriate row of cells. In fact, these phenotypes
sometimes have leaves retuse or with a broadly lunate sinus. Well-de-
veloped axes, on the other hand, tend to have leaves with several spinose
teeth on the dorsal margin and apices more deeply bilobed. In such plants
the segments are ± acuminate, often apiculate, and have a uniseriate row
of several cells. The former phase is illustrated in Schuster (1965b, f. II-8)
and in the Stephani Icones (Lophocolea no. 92a), whereas the latter is also
illustrated in the Stephani Icones (Lophocolea no. 92b) and in Massalongo
(1906), who described the plant as a species of Drepanolejeunea.
Ecology. — A corticolous species mainly of Valdivian forests, occurring
on tree or, less often, shrub trunks. It is comparable to C. gayana in
usually occurring where there is a minimal amount of competition with
other bryophytes and commonly does not occur on trunks or branches
covered or festooned with large, thick masses of bryophytes. Xenocepha-
ENGEL: CLASMATOCOLEA 195
lozia navicularis is sometimes mixed with other hepatics, such as C.
ctenophylla, C. cucullistipula, C. humilis var. suspecta, and Lophocolea
muricata.
Phytogeography. — Nearly exclusively a Valdivian species (sea level-
1,235 m.), occurring from R. Aisen (45° 25' S.) N. to Prov. Arauco
(37° 58' S.); also Andean Patagonia at 800 m. in Parque Nac. Nahuel
Huapi. Also known from one station in the Magellanian region — S.
Skyring, Cabo Leon (Engel, 1973c).
Specimens seen.— CHILE. PROV. MAGALLANES: Cabo Leon,
Hatcher 44-3 (UW-M). PROV. AISEN: R. Aisen, Dusen s.n. as syntype
of L. navicularis — c. tf (NY); ibid., Dusen 215 as Cephalozia trisetosa
Steph., nom. hb. (UPS); ibid., Dusen 225, 237 — c. cf as syntype of L.
cucullistipula (G); ibid., (Dusen) 237 as syntype of L. navicularis (G); ibid.,
Dusen 288 as syntype of L. navicularis (FH, G — c. per.); ibid., Dusen 301
as syntype of L. navicularis (BM); ibid., Dusen 419 as syntype of L.
navicularis (FH, JE — c. per., NY); Puerto Aisen, Schuster s.n. — c.
per. + tf (hb. Grolle). PROV. CHILOE: I. Chiloe, Cord. San Pedro,
Butalcura near R. Butalcura, 1 1 km. by road from Ruta 5, 100 m., Engel
11830 (F). PROV. LLANQUIHUE: R. Puelo, S. Reloncavi, Schwabe s.n.,
5d, 5i, lie — c. per. (JE); Puerto Montt, Dusen s.n. — c. <? (hb. Grolle).
PROV. LLANQUIHUE/PROV. OSORNO: L. Llanquihue, Dusen s.n. as
L. cucullistipula (LD). PROV. OSORNO: 12.1 km. by road below Refugio
Antillanca, 550 m., Engel 11543B — c. 3", 11548 (F); around L. Toro on
road to Refugio Antillanca, 855-885 m., Engel 3992B, 4009C, 4025B,
4052 (MSC); ibid., Imshaug 42963 - c. sporo. (MSC); Termas de Puyehue,
Schwabe 64 (hb. Grolle, JE); ibid., Schwabe 64 p.p. (JE); ibid., 700 m.,
Schwabe 68 p.p. (JE). PROV. VALDIVIA: L. Ranco, 80 m., Kubitzki 153
(hb. Grolle); W. slope of Cord. Pelada, W. of El Mirador, between La
Union and Pta. Hueicolla, 580-680 m., Engel 12259 - c. <? , 12264,
12317 (F); E. slope of Cord. Pelada, E. of El Mirador, between La Union
and Pta. Hueicolla, 740-840 m., Engel 12473 - c. per., 72477, 12414 (F);
near R. Futa in vicinity of Futa, 5.9 km. by road S. of junction of highway
T-60 and T-65, near sea level, Engel 10819 — c. per. (F); C. Lungoico,
500 m., Schwabe 25 f, HCHllO/c (JE); Forestal Trafun, Engel 11107,
11108 (F); R. Llancahue, Schwabe 10 p.p. (JE); SW slope of V. Quetru-
pillan, 1,160-1,235 m., Engel 11172 (F). PROV. CAUTfN: Termas de
Palguin, along R. Palguin, 730 m., Engel 11344 (F). PROV. ARAUCO:
Western foothills of Cord. Nahuelbuta, ridge S. of C. Lanalhue, SW of L.
Lanalhue, 200 m., Engel 12571 (F). ARGENTINA. PROV. RfO NEGRO:
Parque Nac. Nahuel Huapi, A. Frias, ca. 800 m., Donat 31 p.p. (JE).
196 FIELDIANA: BOTANY
Patagonia: Sin. loc., Dusen 215 - c. per., 288, 301, 419, 440, 441, 463
as syntype of L. navicularis (G); ibid., Dusen 288 — c. per. (FH).
11. Clasmatocolea paucistipula (Rodw.) Grolle.
Lophocolea paucistipula Rodw. Pap. & Proc. Roy. Soc. Tasmania
1916:46. 1917 (seors. 11 July 1916) (see also vol. 2 of Tasmanian
Bryophyta p. 46. 1916). Clasmatocolea paucistipula (Rodw.) Grolle,
Rev. Bryol. Lichenol. 29:72. 1960. Hepatostolonophora paucistipula
(Rodw.) Engel, J. Hattori Bot. Lab. 46:103. 1979. Original material:
Tasmania, near Cradle Mt, Rodway (hb. Grolle! — c. cf » HO!).
(Lophocoleaceae).
See Engel 1979b.
12. Clasmatocolea plan! foil a Steph., nom. herb.
Clasmatocolea planifolia Steph. in herb, et icon. ined. Cephaloziella
planifolia Steph. ex Grolle, Rev. Bryol. Lichenol. 29:89. 1960 non
C. planifolia Steph. Hedwigia 32:317. 1893. Cephaloziella novae-
caledoniae Grolle, nom. nov., Rev. Bryol. Lichenol. 29:208. 1960.
Cylindrocolea novae-caledoniae (Grolle) Schust. Nova Hedwigia
22:161. 1972. Holotype: New Caledonia, He du Pin, 1909, Le Rat
(M !). (Cephaloziellaceae).
See Grolle (1960) and Schuster (1972b) for taxonomic treatments; the
former author includes illustrations (pi. II). See also the note in Fulford
(1976, p. 401).
13. Clasmatocolea rotata (Hook. f. & Tayl.) Grolle.
Jungermannia rotata Hook. f. & Tayl. London J. Bot. 3:560. 1844.
Solenostoma rotatum (Hook. f. & Tayl.) Mitt, in Hook. f. Handb.
New Zealand Fl. p. 753. 1867, nom. inval. Solenostoma rotatum
(Hook. f. & Tayl.) Mitt, ex Bastow, Pap. & Proc. Roy. Soc. Tas-
mania 1887:234. 1888. Aplozia rotata (Hook. f. & Tayl.) Rodw.
Pap. & Proc. Roy. Soc. Tasmania 1916:81. 1917 (see also vol. 2 of
Tasmanian Bryophyta p. 31. 1916). Clasmatocolea rotata (Hook. f. &
Tayl.) Grolle, Oesterr. Bot. Z. 117:1. 1969. Hepatostolonophora
rotata (Hook. f. & Tayl.) Engel, J. Hattori Bot. Lab. 46:98. 1979.
Original material: New Zealand, Hooker (S!).
Aplozia alpina Rodw. Pap. & Proc. Roy. Soc. Tasmania 1915: 106. 1916
(seors. 31 December 1915) (see also vol. 2 of Tasmanian Bryophyta p.
32. 1916), syn. fide Grolle (1969). Clasmatocolea alpina (Rodw.)
Grolle, J. Jap. Bot. 41:226. 1966. Original material: Tasmania, Mt.
ENGEL: CLASMATOCOLEA 197
Wellington plateau, January 1915, Rodway s.n. (hb. Grolle!, HO!,
MANCH! - c. per.).
Lophocolea zotovii Herz. Trans. & Proc. Roy. Soc. New Zealand 65 : 354.
1935, syn.fide Grolle (1969). Clasmatocolea zotovii (Herz.) Grolle,
Rev. Bryol. Lichenol. 29:73. 1960. Holotype: New Zealand, North
Island, Tararua Mts., Mt. Hector, ca. 1,310m., 31 December 1933,
Zotov s.n., "Herb. P. R. S. H. 7499" (JE!; isotypes: CHR! - Hodg-
son hb. no. 478, hb. Grolle!). (Lophocoleaceae).
See Engel (1979b).
14. Clasmatocolea truncata Steph.
Clasmatocolea truncata Steph. Bull. Herb. Boissier 5:87. 1897 s Pedi-
nophyllum interruptum subsp. truncatum (Steph.) H. Inoue, J. Hattori
Bot. Lab. 19:33. 1958. Original material: Japan, Hokkaido, 1894,
Faurie 12606 (non vidi). (Plagiochilaceae).
See Hattori (1950), who removed the species from Clasmatocolea,
Inoue (1958) for illustrations, and Grolle (1956, p. 294).
Specimen seen. — Japan, Faurie 12602 (FH — hb. Steph.).
15. Clasmatocolea typhacella Steph., nom. nud.
Clasmatocolea typhacella Steph. in Grolle, Rev. Bryol. Lichenol. 25:301.
1956, nom. nud. pro syn. = Myriocoleopsis puiggari Schiffn. Hedwigia
81 : 235. pi. 1. 1944. (Lejeuneaceae).
See Grolle (1956, p. 301) and Gradstein & Vital (1975).
Specimen seen.— BRAZIL. ESTADO DE SAO PAULO: Iporanga,
Puiggari (FH— hb. Steph.!, M!).
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1. Clasmatocolea rigens (Hook. f. Tayl.) Engel. X 5,000; from Engel 1359 A.
204
2. Clasmatocolea rigens (Hook. f. & Tayl.) Engel. X 10,000; from Engel 1359A.
205
3. Clasmatocolea obvoluta (Hook. f. & Tayl.)
Grolle. X 4,700; from Engel 4574 A .
4. Clasmatocolea obvoluta (Hook. f. & Tayl.) Grolle. X 9,500; from Engel 4574 A.
206
5. Clasmatocolea trachyopa (Hook. f. &
Tayl.) Grolle. X 5,000; from Engel 5151 A.
6. Clasmatocolea tra-
chyopa (Hook. f. &
Tayl.) Grolle. X 10,000;
from Engel 5 151 A.
207
7. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 6,600; from Engel 5472.
208
8. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 9,700; from Engel 5472.
209
9. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 28,600; from Engel 5472.
210
10. Clasmatocolea humilis (Hook. f. & Tayl.) Grolle. X 46,000; from Engel 5472.
211
11. Clasmatocolea vermicu-
/orw(Lehm.) Grolle. X 5,000;
from Dusen 166.
12. Clasmatocolea vermicular is (Lehm.) Grolle. X 10,000; from Dusen 166.
212
13. Clasmatocolea gayana (Mont.) Grolle. X 3,655 ; from Engel 4554.
213
14. Clasmatocolea gayana (Mont.) Grolle. X 10,260; from Engel 4554.
214
15. Clasmatocolea ctenophylla
(Schiffn.) Grolle. X 5,000; from
Engel 5214.
16. Clasmatocolea ctenophylla (Schiffn.) Grolle. X 10,000; from Engel 5214.
215
17. Clasmatocolea minutiretis
Engel & Grolle. X 5,400; from
Engel 4785.
18. Clasmatocolea minutiretis Engel & Grolle. X 10,750; from Engel 4785.
216
19. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. X 5,400; from Engel 6125.
217
20. Clasmatocolea fulvella (Hook. f. & Tayl.) Grolle. X 10,750; from Engel 6125.
21. Clasmatocolea strongylophylla (Hook. f.
&Tayl.) Grolle. X 5,240; from Schuster 48481.
218
22. Clasmatocoleastrongylophylla (\\oo\i. f. &Tayl.)Grolle. X 10,000; from Schuster
48481.
23. Clasmatocolea puccioana
(De Not.) Grolle. X 6,075; from
Engel 4520.
219
24. Clasmatocolea puccioana (De Not.) Grolle. X 12,100; from Engel 4520.
220
A I
^^•i^^ ^•••^••••••••il^^B
25. Clasmatocolea puccioana (De Not.) Grolle. X 25,300; from Engel 4520.
221
26. Clasmatocolea navistipula (Steph.)
Grolle. X 5,000; from Engel4251.
27. Clasmatocolea navistipula (Steph.) Grolle. X 10,000; from Engel 4251.
222
28. Clasmatocoleacucullistipula(Steph.)Gro\\e. X 3, 100; from Dusens.n., syntype of
L. cucullistipula.
223
29. Clasmatocoleacucullistipula(Steph.)GTO\\e. X 10,200; from Dusens.n., syntype of
L. cucullistipula.
224
INDEX OF TAXA
Recognized taxa are in roman type, illegitimate or synonymous taxa
are italicized, and new taxa and new combinations are in boldface.
Alicularia flexuosa (Lehm.) Nees 97
Alicularia grandistipula Herz. 99
Alicularia grandistipula (Steph.) Horik.
99
Alicularia lindmanii (Steph.) Steph. 97
Alicularia strongylophylla (Hook. f. &
Tayl.) G.L. & N. 147
Alicularia strongylophylla var. minor
(Hook. f. & Tayl.) G. L. & N. 149
Alicularia vertnicularis (Lehm.) G. L. &
N. 96
Anomylia cuneifolia (Hook.) Schust.
191
Anthoscyphus tjiwideiensis (Sande-Lac.)
Trev. 154
Aplozia alpina Rodw. 196
Aplozia cuneifolia (Hook.) Dum. 191
Aplozia rotata (Hook. f. & Tayl.) Rodw.
196
Blepharostoma acanthifolium Gola 60
Calyptrocolea perssonii Schust. 192
Cephalozia rigens (Hook. f. & Tayl.)
Trev. 44
Cephalozia trisetosa Steph. 193
Cephaloziella novae-caledoniae Grolle
196
Cephaloziella planifolia Steph. (1893)
196
Cephaloziella planifolia Steph. ex Grolle
196
Chiloscyphus cordifolius Rodw. 173
Chiloscyphus fasciculatus (Nees) G. L. &
N. 91
Chiloscyphus fasciculatus var. /3 dregeana
Nees 91
Chiloscyphus fasciculatus var. y exarida
Nees 91
Chiloscyphus fulvellus (Hook. f. &
Tayl.) Nees 139
Chiloscyphus gayanus (Mont.) G. L. &
N. 112
Chiloscyphus nigrescens Lindenb. &
Hampe 97
Chiloscyphus notophyllus (Hook. f. &
Tayl.)G. L. & N. 173
Chiloscyphus rabenhorstii Steph. 91
Chiloscyphus tjiwideiensis Sande-Lac.
154
Chonecolea doellingeri (Nees) Grolle
191
Clasmatocolea Spruce 39
Subg. Clasmatocolea 69
Subg. Lacerfolia Steph. ex Engel
50
Subg. Metaclasmatocolea Engel 1 57
Subg. Plicaticalyx Engel 183
Subg. Protoclasmatocolea Engel 44
Subg. Squamicalyx Engel 173
Sect. Clasmatocolea 74
Sect. Fulvellae Engel 138
Sect. Metaclasmatocolea 165
Sect. Pa chy Clasmatocolea Engel 69
Sect. Puccioanae Engel 157
Sect. Strongylophyllae Engel 147
225
226
FIELDIANA: BOTANY
Clasmatocolea abnormis(Besch. & Mass.)
Grolle 190
Clasmatocolea acutiloba Schiffn. 191
Clasmatocolea alpina (Rodw.) Grolle
196
Clasmatocolea angulistipula (Steph.)
Grolle 149
Clasmatocolea chilensis Steph. 97
Clasmatocolea cookiana (Mass.) Engel
58
Clasmatocolea crassimarginata Grolle
69
Clasmatocolea crassiretis (Herz.) Grolle
124
Clasmatocolea ctenophylla (Schiffn.)
Grolle 119
Clasmatocolea cucullistipula (Steph.)
Grolle 183
Clasmatocolea cuneifolia (Hook.) Spruce
191
Clasmatocolea doellingeri (Nees) Steph.
191
Clasmatocolea exigua Steph. 192
Clasmatocolea fasciculata (Nees) Grolle
91
Clasmatocolea fiordlandiae (Hodgs.)
Grolle 192
Clasmatocolea flavovirens (Steph.) Herz.
99
Clasmatocolea fragillima Spruce 192
Clasmatocolea fulvella (Hook. f. &
Tayl.) Grolle 139
Clasmatocolea gayana (Mont.) Grolle
112
Clasmatocolea georgiensis (Gott.) Grolle
193
Clasmatocolea heterostipa Spruce 40,
97
Clasmatocolea humilis (Hook. f. &
Tayl.) Grolle 74
Clasmatocolea humilis (Hook. f. &
Tayl.) Grolle var. humilis 80
Clasmatocolea humilis var. polymorpha
Engel 91
Clasmatocolea humilis var. suspecta
(Mass.) Engel 88
Clasmatocolea innovata Herz. 193
Clasmatocolea koeppensis (Gott.) Grolle
44
Clasmatocolea marginata (Steph.) Grolle
69
Clasmatocolea minutiretis Engel &
Grolle 133
Clasmatocolea moniliformis Engel 128
Clasmatocolea navicularis (Steph.) Grolle
193
Clasmatocolea navistipula (Steph.)
Grolle 165
Clasmatocolea navistipula (Steph.)
Grolle var. navistipula 169
Clasmatocolea navistipula var. parcera-
mosa Engel 172
Clasmatocolea notophylla (Hook. f. &
Tayl.) Grolle 173
Clasmatocolea obvoluta (Hook. f. &
Tayl.) Grolle 50
Clasmatocolea obvoluta var. cookiana
(Mass.) Engel 58
Clasmatocolea obvoluta (Hook. f. &
Tayl.) Grolle var. obvoluta 56
Clasmatocolea paucistipula (Rodw.)
Grolle 196
Clasmatocolea planifolia Steph . 1 96
Clasmatocolea puccioana (De Not.)
Grolle 157
Clasmatocolea rigens (Hook. f. &
Tayl.) Engel 44
Clasmatocolea rotata (Hook. f. & Tayl.)
Grolle 196
Clasmatocolea strongylophylla (Hook.
f. & Tayl.) Grolle 147
Clasmatocolea tjiwideiensis (Sande-
Lac.) Grolle 154
Clasmatocolea trachyopa (Hook. f. &
Tayl.) Grolle 60
Clasmatocolea truncata Steph. 197
Clasmatocolea turgescens (Hook. f. &
Tayl.) Grolle 149
Clasmatocolea typhacella Steph. 197
Clasmatocolea vermicularis (Lehm.)
Grolle 96
Clasmatocolea verrucosa Engel 179
Clasmatocolea zotovii (Herz.) Grolle 197
Coleochila cuneifolia (Hook.) Dum. 191
Conoscyphus flaccidus Pears. 112
Conoscyphus tjiwideiensis (Sande-Lac.)
Schiffn. 154
Cylindrocolea acutiloba (Schiffn.) Engel
191
Cylindrocolea novae-caledoniae (Grolle)
Schust. 196
ENGEL: CLASMATOCOLEA
227
Cylindrocolea rhizantha (Mont.) Schust.
192
Drepanolejeunea stephaniana Mass. 193
Evansianthus georgiensis (Gott.) Schust.
&EngeI 193
Hepatostolonophora abnormis (Besch.
& Mass.) Engel & Schust. 190
Hepatostolonophora paucistipula
(Rodw.) Engel 196
Hepatostolonophora rotata var. pers-
sonii (Schust.) Engel 192
Hepatostolonophora rotata (Hook. f. &
Tayl.) Engel 196
Heteroscyphus fasciculatus (Nees)
Schiffn. 91
Heteroscyphus rabenhorstii (Steph.)
Schiffn. 91
Isotachis capensis Steph. ex Herz. 93
Jamesoniella difficilis Steph. 171
Jamesoniella gracilis Gola 191
Jungermannia bergiana Lehm. 91
Jungermannia chamissonis Gott. &
Lindenb. 97
Jungermannia chilensis Lehm. 193
Jungermannia chilensis Mont. 193
Jungermannia cuneifolia Hook. 191
Jungermannia doellingeri Nees 1 9 1
Jungermannia dregeana Lehm. &
Lindenb. 91
Jungermannia fasciculata (Lindenb.)
Hook. f. & Tayl. 91
Jungermannia fasciculata Nees 91
Jungermannia flexuosa Lehm. 97
Jungermannia fulvella Hook. f. & Tayl.
139
Jungermannia gayana Mont. 112
Jungermannia humilis Hook. f. & Tayl.
80
Jungermannia humilis Kash. & Chopra
80
Jungermannia marginal a Hook. f. &
Tayl. 69
Jungermannia marginal a Lehm. 69
Jungermannia marginata Mitt. 69
Jungermannia moniliformis Angstr. 190
Jungermannia notophylla Hook. f. &
Tayl. 173
Jungermannia obvoluta Hook. f. &
Tayl. 56
Jungermannia obvolutaeformis De Not.
56
Jungermannia palustris Hook. f. & Tayl.
80
Jungermannia Ipuccioana De Not. 157
Jungermannia rigens Hook. f. & Tayl.
44
Jungermannia rotata Hook. f. & Tayl.
196
Jungermannia strongylophylla Hook. f. &
Tayl. 147
Jungermannia strongylophylla var. /3
minor Hook. f. & Tayl. 149
Jungermannia subintegra Hook. f. &
Tayl. 97
Jungermannia trachyopa Hook. f. &
Tayl. 60
Jungermannia turgescens Hook. f. &
Tayl. 149
Jungermannia variabilis (Sim) S. Arnell
101
Jungermannia vermicularis Hub. 96
Jungermannia vermicularis Lehm. 96
Leioscyphus (1) abnormis Besch. & Mass.
190
Leioscyphus chamissonis (Gott. &
Lindenb.) Spruce 97
Leioscyphus cuneifolius (Hook.) Steph.
191
Leioscyphus humilis (Hook. f. & Tayl.)
Pears. 80
Leioscyphus marginatus Steph. 69
Leioscyphus patagonicus Pears. 88
Leioscyphus patagonicus Steph. 88
Leioscyphus strongylophyllus (Hook. f.
& Tayl.) Mitt. 147
Leioscyphus turgescens (Hook. f. &
Tayl.) Mitt. 149
Lejeunea subintegra (Hook. f. & Tayl.)
G. L. & N. 97
Leptoscyphus chamissonis (Gott. &
Lindenb.) Mitt. 97
Leptoscyphus cuneifolius (Hook.) Mitt.
191
228
FIELDIANA: BOTANY
Leptoscyphus liebmanianus (Lindenb.
& Gott.) Mitt. 192
Leptoscyphus porphyrius (Nees) Grolle
193
Leptoscyphus strongylophyllus (Hook. f.
& Tayl.) Mitt. 147
Leptoscyphus turgescens (Hook. f. &
Tayl.) Mitt. 149
Lophocolea abnormis (Besch. & Mass.)
Steph. 190
Lophocolea angulistipula Steph. 149
Lophocolea arenaria Schiffn. 60
Lophocolea austrigena (Hook. f. &
Tayl.) G. L. & N. 24, 82, 177
Lophocolea boliviensis S. Arnell 99
Lophocolea boliviensis Steph. 99
Lophocolea boveana Mass. 24
Lophocolea catenulata Herz. 81
Lophocolea concava Steph. 117
Lophocolea cookiana Mass. 58
Lophocolea crassiretis Herz. 124
Lophocolea cristato-spinosa Steph. 119
Lophocolea ctenophylla SchifFn. 119
Lophocolea cucullifolia Steph. 186
Lophocolea cucullistipula Steph. 183
Lophocolea debilis Steph. 44
Lophocolea diversistipa Steph. 159
Lophocolea elata f. aquatica Herz. 101
Lophocolea fasciculata (Nees) S. Arnell
& Grolle 91
Lophocolea filiformis Steph. 186
Lophocolea fiordlandiae Hodgs. 192
Lophocolea flavovirens Steph. 99
Lophocolea fulvella (Hook. f. & Tayl.)
Mass. 139
Lophocolea gayana (Mont.) Mitt. 112
Lophocolea georgiensis Gott. 193
Lophocolea gottscheoides Besch. &
Mass. 24
Lophocolea gunniana Nees 24, 82, 177
Lophocolea hickenii Herz. 88
Lophocolea hastatistipa Steph. 80
Lophocolea homomalla Steph . 1 39
Lophocolea humilis (Hook. f. & Tayl.)
Steph. 80
Lophocolea incrassata Steph. 80
Lophocolea integerrima Steph. 80
Lophocolea koeppensis Gott. 44
Lophocolea lacerata Steph. 60
Lophocolea latissima Steph. 58
Lophocolea lenta (Hook. f. & Tayl.)
G. L. &N. 21,24,48
Lophocolea leptantha (Hook. f. &
Tayl.) G. L. & N. 24
Lophocolea ligulata Steph. 99
Lophocolea longissima Steph. 99
Lophocolea magellanica Schiffn. 80
Lophocolea minima S. Arnell 101
Lophocolea multispinula Steph. 88
Lophocolea navicularis Steph. 193
Lophocolea navistipula Steph. 169
Lophocolea notophylla (Hook. f. &
Tayl.) Hodgs. 173
Lophocolea obvoluta (Hook. f. & Tayl.)
Evans 56
Lophocolea obvolutaeformis (De Not.)
Mass. 56
Lophocolea okaritana Steph. 177
Lophocolea olens Herz. 44
Lophocolea otiphylla (Hook. f. & Tayl.)
Mitt. 24,82
Lophocolea ovistipula Herz. 101
Lophocolea palustris (Hook. f. & Tayl.)
Besch. & Mass. 80
Lophocolea patagonica Beauv. 1 59
Lophocolea paucistipula Rodw. 196
Lophocolea puccioana (De Not.) Mass.
157
Lophocolea puccioana a* forma primi-
genia Mass. 157
Lophocolea puccioana var. /3 primigenia
(Mass.) Schiffn. 157
Lophocolea puccioana var. /3 suspecta
Mass. 88
Lophocolea rigens (Hook. f. & Tayl.)
Evans 44
Lophocolea rotundifolia Steph. 81
Lophocolea rotundistipula Steph. (1906)
159
Lophocolea rotundistipula Steph. (1911)
159
Lophocolea sabuletorum (Hook. f. &
Tayl.) G. L. & N. 24, 103
Lophocolea skottsbergii Steph. 99
Lophocolea stephanii Herz. 44
Lophocolea stephanii Schiffn. 44
Lophocolea strongylophylla (Hook. f. &
Tayl.) Hodgs. 149
Lophocolea subaromatica Herz. 49
Lophocolea subcapillaris Steph. 171
ENGEL: CLASMATOCOLEA
229
Lophocolea subintegra (Hook. f. &
Tayl.)Grolle 97
Lophocolea subulistipa Herz. 101
Lophocolea tesorensis Herz. 110
Lophocolea textilis (Hook. f. & Tayl.)
G. L. & N. 24
Lophocolea trachyopa (Hook. f. &
Tayl.) G. L. & N. 60
Lophocolea tumida Steph. 173
Lophocolea turbiniflora Steph. 99
Lophocolea turgescens (Hook. f. &
Tayl.)Hodgs. 149
Lophocolea vermicularis Herz. 93
Lophocolea vinciguerreana Mass. 1 1 2
Lophocolea zotovii Herz. 197
Mesophylla strongylophylla (Hook. f. &
Tayl.)Trev. 147
Microlejeunea chilensis (Mont.) Steph.
193
Mylia chamissonis (Gott. & Lindenb.)
Trev. 97
Mylia humilis (Hook. f. & Tayl.) Trev.
80
Mylia turgescens (Hook. f. & Tayl.)
Trev. 149
Mylius cuneifolius (Hook.) S. Gray 191
Myriocoleopsis puiggari Schiffn. 197
Nardia humilis (Hook. f. & Tayl.)
Berggr. 80
Nardia lindmanii Steph. 97
Nardia vermicularis (Lehm.) Trev. 96
Notoscyphus belangerianus (Lehm.)
Mitt. 104
Notoscyphus flexuosus (Lehm.) Sim 97
Notoscyphus lindmanii (Steph.) Schiffn.
97
Notoscyphus variifolius Mitt. 97
Notoscyphus vermicularis (Lehm.) Steph.
96
Odontoschisma marginatum (Steph.)
Steph. 69
Odontoschisma variabite (Lindenb. &
Gott.) Trev. 101
Odontoschisma variabile Sim 101
Pedinophyllum interruptum subsp.
truncatum (Steph.) Inoue 197
Phragmatocolea innovata (Herz.) Grolle
193
Solenostoma humilis (Hook. f. & Tayl.)
Mitt. 80
Solenostoma rotatum (Hook. f. & Tayl.)
Mitt. (1867) 196
Solenostoma rotatum (Hook. f. & Tayl.)
Mitt, ex Bastow (1888) 196
Stolonophora abnormis (Besch. & Mass.)
Engel & Schust. 190
Xenocephalozia navicularis (Steph.)
Schust. 193
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