14

Dicksoniaceae Bower
Genera 75-80

A tropical family of six distinctive, mostly Old World genera. Cystodium and Calochlaena are con-
fined to the Old World, and Thyrsopteris is endemic in the Juan Fernandez Islands. The spores are
consistently trilete and have a two-layered exospore. The prominent tubercles and equatorial
flange in Cibotium spores are also formed by the exospore. The pitted exospore surface, common
to some species of Dicksonia, may be obscured by granulate perispore deposit. The perispore is
usually two-layered, the outer one often includes rodlets, and the inner one forms the surface
contours in spores with plain exospore. The striate surface in Cystodium and the slightly papillate
one in Thyrsopteris are formed by the perispore. The exospore is especially elaborate in Calochlaena
and forms the coarse, tuberculate contours. The diversity of spores in the Dicksoniaceae and
differences in derivation of the contours do not support close relations between these genera.

A. F. Tryon et al., Spores of the Pteridophyta

© Springer-Verlag New York Inc. 1991
 Culcita 219

75. Culcita Presl

Figs. 75.1-75.10
Genus
A genus of two geographically discrete species. Aperture Trilete, the arms 'h to % the radius.
Surface C. macrocarpa-both faces with erect,
Range slender spines or rodlets, sometimes sparse (Figs.
The American Culcita coniifolia occurs from 2, 3, 6). C. coniifolia-shallowly tuberculate to
southern Mexico south to Peru, in eastern Bra- rugate distal face; the proximal nearly psilate,
zil, and in the Greater Antilles. Culcita macro- often with a slight ridge in equatorial area be-
carpa grows in southern Spain and Portugal, the tween proximal and distal faces (Fig. 7).
Azores, and Atlantic islands off North Africa. Structure Exospore of two layers, nearly plain;
perispore of two layers, forming the surface
Spores contours.
Nine collections examined from distant areas of
the range of both species. Comments
Size 42-53 JLm. Characteristics Spores of both species are
Shape Tetrahedral-globose, strongly depressed strongly angled and depressed between the ap-
between aperture arms, often with slightly erture arms (Figs. 1, 7). The relatively thin wall
prominent equatorial ridge. in the section of the whole spore envelops mem-

2 3

4 5 6

Figs. 75.1-75.6. Culcita spores, Xl,OOO, and wall sections. vesicles, X2,000. 4. Fractured sporoderm, the compact ex-
1-6. C. macrocarpa Pres\. 1-4. Pico, Azore Islands, Tryon ospore below, Xl 0,000. 5. Depressed areas between an-
& Tryon 7448. 1. Proximal face below, lateral above, with gles, sparse spines, Spain,Jermy 9899 (Z). 6. Dense, partly
sparse spines. 2. Detail of surface rodlets. X5,000. 3. fused spines, Spain, Molesworth-Allen 6065, Xl 0,000.
Fractured spore with thin wall enclosing membrane bound
220 Dicksoniaceae

7 8

9 10

Figs. 75.7-75.10. Culcita spores, X 1,000, and wall sections. Culcita spores. 9. C. macrocarpa, distal face, outer exo-
7,8. C. coniifolia (Hook.) Maxon. 7. Distal surface shallow- spore below perispore layers (E), the outer with spines,
ly tuberculate to low rugate below, equatorial ridge (arrow), Tryon & Tryon 7448, X50,000. 10. C. coniifolia, apertural
Venezuela, Steyermark 94026. 8. Fractured lateral wall, section with several canals, perispore of two layers, the inner
thick exospore below slightly irregular perispore, Colombia, showing artifactual lines (arrow), Venezuela, Steyermark
Killip & Smith 20690, XI 0,000.. 9, 10. Wall sections of 94026, X 18,000.

brane-bound vesicles of varied size (Fig. 3). The Relationships The contours and surface depo-
exospore is overlaid by a thin perispore with sition of Culcita spores generally resemble those
slightly irregular surface (Figs. 4, 8-10). Studies of other genera in the Dicksoniaceae. The
on spores of the Dicksoniaceae show an ex- strong angles, with depressed areas between ap-
panded perispore in Culcita spores treated with erture arms, and the relatively unelaborated
sodium hydroxide (Gastony, 1981). surface suggest possible relations with Thyrsop-
Diversity Differences in the perispore and short teris. The equatorial ridges in well formed
spines of Culcita macrocarpa spores, compared spores of Culcita coniifolia suggest possible alli-
to the spineless perispore in C. coniifolia, are ances with Dicksonia and Cibotium. The spores
relatively slight. Density of the spines or rodlets supply some clues on affinities in the family, but
in C. macrocarpa spores varies within as well as the exact relation of the genus is unclear.
between collections (Figs. 2-6). Those of C. con-
iifolia lack rodlets or spines, and spores from Literature
Peru, figured by Gastony (1981), have a prom-
Gastony, G.J. 1981. Spore morphology in the Dick-
inent, spheroidal surface deposit. This variation
soniaceae, 1. The genera Cystodium, Thyrsopteris,
may represent different developmental condi- and Culcita. Am. J. Bot. 68: 808-819.
tions of the spores, or possibly reflect changes
resulting from geographic isolation.
 Calochlaena 221

76. Calochlaena (Maxon) Turner & White

Figs. 76.1-76.6
Genus
A paleotropical genus of five species, formerly Surface Coarse, more or less fused, basally
treated in Culcita. confluent tubercles.
Structure Exospore of two layers, the outer
Range forming the contours; the perispore an outer
Malesia, Java to the Philippines and New Guin- layer above a dark, granulate deposit (Figs. 5,
ea extending eastward in the Pacific to Samoa, 6).
and southward to eastern Australia and Tas-
mania. Comments
Characteristics, Diversity The spores are re-
Spores markably uniform and indistinguishable in the
Three species examined. three species (Figs. 1-3). The aperture is nearly
Size 35-43 Jim. obscured by ridges adjacent to the arms (Fig.
2). The basally connected tubercles, partly fused
Shape Spheroidal. in irregular ridges, have a pebbled microverru-
Aperture Trilete, the arms % the radius, usually cate surface (Fig. 4). The exospore of two layers
obscured by adjacent ridges. is of the blechnoid type with microverrucae,

3 4

Figs. 76.1-76.4. Spores of Calochlaena, XI ,000. 1. C. New Guinea, Brass 11589. 3,4. C. dubia (R. Br.) Turner
straminea (Labill.) Turner & White, irregular coarse tuber- & White, Australia Clemens, in 1949. 3. Coalescent tuber-
cles, the proximal face at right, Fiji, Degener & Ordonez cles. 4. Microverrucae on surface of tubercles, part of the
13834. 2. C. villosa (C. Chr.) Turner & White. The aper- aperture arm, lower right, XI 0,000.
ture partly obscured by prominent, coalescent tubercles,
222 Dicksoniaceae

5 6

Figs. 76.5, 76.6. Wall section of Calochlaena straminea, Sol- X 15,000. 6. Detail of perispore with dark, granulate de-
omon Islands, Brass 3402. 5. Apertural section with a posit below thin, compact layer (P) on exospore tubercle (E)
prominent exospore tubercle with microverrucate surface, of distal face, X 75,000.
adjacent to the slit; perispore (P) the thin surface layer,

formed on lobes of the outer exospore, covered The coarse exospore contours with microv-
by thin perispore (Fig. 5). errucae and thin perispore of Calochlaena
Relationships The spores support recognition spores are unlike the elaborate perispore for-
of Calochlaena as clearly distinct from Culcita. mation with rodlets and the unusual pitted
The spheroidal shape and prominent tubercu- structure, in the spore wall of some dicksonias,
late to verrucate surface differ from spores of as D. antarctica. However, the absence of pits in
Culcita and most other genera of the Dicksoni- spores of Dicksonia squarrosa (see Dicksonia, 80,
aceae. An alliance with Dicksonia, suggested on Figs. 13, 14) suggests possible connections be-
the basis of morphological assessment of Cal- tween Calochlaena and some elements in the
ochlaena, particularly the shoot morphology, genus.
does not include the spores (White & Turner,
1988). The spores of Calochlaena (as subgenus Literature
Calochlaena) are compared with those of the
Gastony, J.G. 1981. Spore morphology in the Dick-
dicksonias, as D. squarrosa Sw., as well as with soniaceae. 1. The genera Cystodium, Thyrsopteris,
species of Dennstaedtia (Gas tony, 1981). How- and Culcita. Am. J. Bot. 68: 808-819.
ever, the similarities noted between Calochlaena White, R.A., & M.D. Turner. 1988. Calochlaena, a
and Dennstaedtia in that work are considered to new genus of dicksonioid ferns. Am. Fern J. 78:
be superficial by White & Turner (1988). 86-95.
 Cibotium 223

77. Cibotium Kaulfuss

Figs. 77.1-77.14
Genus
A genus of eight or nine species: two or possibly Aperture Trilete, the arms 213 to % the radius,
one in America, two or three in southeast Asia often less prominent than acljacent ridges.
and Maiesia, and four in Hawaii. Surface Plain and more or less depressed, es-
pecially on the proximal face (Fig. 2), with ir-
Range regular granulate deposit (Figs. 4, 7), or usually
with a prominent flange and coarse exospore
In America in southern Mexico and Central
ridges, especially on the distal face (Figs. 3, 6,
America; in Asia from Assam to southern
8).
China, Malesia from Java to New Guinea, and
Hawaii in the Pacific islands. Structure Exospore of two layers forming
coarse ridges and the flange (Figs. 9, 12); peri-
Spores spore of two slightly differented layers (Figs.
10-14).
Examined in seven species.
Size 47-80 /-Lm. Comments
Shape Tetrahedral-globose with extended an- Characteristics Prominently ridged spores with
gles and usually prominent equatorial flange. a well-developed equatorial flange are common

3 4

Figs. 77.1-77.4. Cibotium spores, X 1,000. 1. C. regale XI,OOO. 3,4. C. baromen (L.) Sm., China, Wang 76917.
Versch. & Lem., granulate, slightly ridged spores, proximal 3. Prominent equatorial flange; raised areas near aperture,
face at left, Honduras, Molina & Molina 14017, X I ,OOO. granulate proximal face, left; coarsely ridged distal face,
2. C. schiedei Schlect. & Cham., granulate, proximal face right. 4. Detail of granulate proximal face, X IO,OOO.
depressed between aperture arms, Mexico, Bourgeau 2378,
224 Dicksoniaceae

5 6

7 8

Figs. 77.5-77.8. Cibotium spores, X500. 5. C. splendens face with coarse parallel ridges, X500. 7. Abraded gran-
(Gaudich.) Krajina, coarsely ridged young spores, Hawaii, ulate outer perispore and exposed inner perispore,
Wilber 460, X500. 6, 7. C. cumingii Kze. var. arachnioi- X 10,000. 8. C. barometz, coarse, long ridges distal face, at
deum C. Chr., Borneo, Chew et al., in 1961. 6. Ridged areas left, granulate proximal face, at right, Burma, Kingdon- Ward
near aperture arms and prominent equatorial flange; distal 20433, X500.

to most species of Cibotium (Figs. 1, 3). The ex- verse the outer exospore and are obscured by
ospore ridges appear to be well formed late in perispore in mature spores.
wall development as shown by immature spores Relationships The prominent equatorial flange
lacking prominent ridges in SEM work on and ridges common to the Old World species
spores of Taiwan species of Cibotium (Liew & characterize Cibotium spores. The plain spores
Wang, 1976). of C. schiedei are regarded as a specialized form.
Diversity Spores of the American species Cibo- Those of the closely related C. regale (sometimes
tium schiedei consistently lacking ridges (Fig. 2) treated in the former species) with less promi-
and the less prominent ones in spores of the nent ridges, are considered forms intermediate
closely related, American C. regale (Fig. 1), con- to the latter and the strongly ridged spores of
trast with spores of Old World species. The Old World species. The plain spores may pos-
alignment of ridges in the latter may vary, as in sibly represent a neotenic form of the young
the long, contiguous ones in C. splendens (Fig. spores noted by Liew & Wang (1976).
5), and those of C. cumingii var. arachnioideum In the analyses of fossil spores and pollens of
(Fig. 6), with prominent parallel bands on the the Hawaiian Islands by Selling (1946, 1948),
distal face. Quaternary records of Cibotium spores were
The assessment of spore diversity in Cibotium considered to be an indicator of rain forest. The
based on species from three geographic re- spores occur in 80% of the samples, varying
gions, illustrates the different types of ridges, from trace amounts to 46% of the total. Cibo-
granulate deposit, and the presence of peri- tium was one of the most important constituents
spore (Gas tony, 1982). Pits shown in SEM fig- in the second of three major phases of the Qua-
ures in that work correspond to the open ends ternary vegetation, indicating a period of ex-
of the canals, shown here (Fig. 12). These tra- tended rain forest. Conditions were especially
 Cibotium 225

9 10

II 12

13 14

Figs. 77.9-77.14. Wall sections of Cibotium spores. 9, 10. Philippines, Elmer 22023. 12. Apertural section with prom-
C. barometz, Wang 76917. 9. Apertural section with raised inent raised area acljacent to aperture, X3,600. 13. Gran-
proximal ridges, X9,OOO. 10. Detail of granulate perispore ulate perispore with scales adjacent to exospore (E),
with diffuse outer layer on exospore (E), X 75,000. 11. C. X65,OOO. 14. Mature perispore with compact, thin layer
glaucum, granulate perispore above part of exospore (E), above diffuse inner part, X65,OOO.
Hort. Kew 44 (1969), X 75,000, (TL). 12-14. C. cumingii,
226 Dicksoniaceae

favorable for growth of Cibotium during a few equatorial flange clearly indicate it is not
periods, which characterize transitions toward Cibotium.
particularly wet conditions spread over the
postglacial. The three highest records of these Literature
spores correspond to the greatest accumulation Gastony, G.]. 1982. Spore morphology of the Dick-
of peat. soniaceae. 2. The genus Cibotium. Can.]. Bot. 60:
Fossil spores of Cibotiumsporites cumingii 955-972.
Huang, from Miocene deposits of Taiwan, are Huang, T.C. 1978. Miocene palynomorphs of Tai-
associated with the extant Cibotium cumingii wan. 3. Spores. Taiwania 23: 7-55.
Kze. (Huang, 1978). The large spore size and Kremp, G.O.W., & T. Kawasaki. 1972. See general
well-developed equatorial flange figured here literature.
support the alliance. A somewhat smaller spore, Liew, F.S., & S.C. Wang. 1976. Scanning electron mi-
Cingulatisporites taiwanensis Huang, from the croscopical studies on the spores of Pteridophytes.
same formation, with a less prominent equato- 8. The tree fern family (Cyatheaceae) and its allied
species found in Taiwan. Taiwania 21: 251-266.
rial ridge, is also regarded as possibly related to
Selling, O.H. 1946. Studies in Hawaiian pollen sta-
Cibotium, but this is less certain. tistics. 1. Spores of the Hawaiian Pteridophytes.
A spore from the Lower Paleocene in the Bishop Mus. Spec. Pub\. 37: 1-87.
central Urals was included in the compilation of Selling, O.H. 1948. Studies in Hawaiian pollen sta-
Kremp & Kawasaki as Cibotium gleichenioides tistics. 3. On the late Quaternary history of the
Agranovskaya (1972). However, the small size, Hawaiian vegetation. Ibid. 39: 1-154.
long aperture equal to the radius, and slight

78. Cystodium Hooker

Figs. 78.1-78.8
Genus
A mono typic genus of the paleo tropics, based Figs. 78.1-78.8. Cystodium sorbifolium (Sm.) J. Sm. spores,
on Cystodium sorbifolium (Sm.) J. Sm. X 1,000, and wall sections. 1-3,5. Papua, Brass 23810. 1.
Sinuous ridges slightly granulate. 2. Aperture nearly ob-
scured by granulate deposit. 3. Coarse tapetal deposit (ar-
Range row). 4. Fractured spore, exospore exposed below abraded
perispore, New Guinea, Floyd 5714. 5. Abraded perispore,
From Borneo to New Guinea, and adjacent with scales above exospore, X 10,000. 6-8. Papua, Brass
islands. 1087. 6. Apertural section of two-layered exospore, canals
traverse the outer one, without perispore, X 12,000. 7.
Section of outer exospore (Ee) with scales below perispore
Spores (arrow), more diffuse outer perispore above, X 65,000. 8.
Exospore (E) below two perispore layers, the inner one
Six collections examined. forms the prominent ridge, X40,OOO.
Size 44-58 #Lm.
Shape Globose.
Comments
Aperture Trilete, the arms '/2 to 3/4 the radius.
Characteristics The sinuous ridges are often
Surface Granulate deposit over low, sinuous nearly obscured by the granulate outer peri-
ridges. spore (Figs. 1-3, 8). The granulate material
Structure Plain exospore two layered; peri- forming the outer shell is readily detached from
spore two layered, the sinuous ridges formed by the exospore (Figs. 4, 5). This may separate at
the inner layer with scales in the inner part the inner scales formed below the perispore
(Figs. 6-8). (Fig. 5).
 Cystodium 227

2 3

4 5

6 7
8
228 Dicksoniaceae

Diversity Sporangia of Cystodium form 32 rath- Dennstaedtia, such as D. magnifica and D. sa-
er than the normal complement of 64 spores. moensis. The spores suggest possible connec-
Reduction in spore number is exceptional in the tions with the Dennstaedtiaceae, but analyses of
Dicksoniaceae, although some genera in the Cy- other characters are needed to substantiate the
atheaceae characteristically have 16 spores per alliance.
sporangium. The coarse, granulate deposit on The genus was recognized in a distinct family
Cystodium spores varies with maturity and quan- based on the unique vasculature of the stipe
tity of residual material formed by the tapetum and other characters, including spores. It was
(Fig. 3). The assessment of the genus in studies considered to be allied with the Dicksoniaceae,
on spores of the Dicksoniaceae, includes a re- but with affinities possibly with the Dennstaed-
view of immature spores with exposed sinuous tiaceae (Croft, 1986).
ridges and fully developed ones that are densely
granulate (Gas tony, 1981). Literature
Relationships Cystodium spores somewhat re-
Croft, ].R. 1986. The stipe and rachis vasculature of
semble those of Metaxya (another monogeneric
the Dicksonioid fern, Cystodium sorbifolium (Cysto-
family) in their spheroidal shape and granulate diaceae). Kew Bull. 41: 789-803.
surface. In both genera the perispore is easily Gastony, G.]. 1981. Spore morphology in the Dick-
eroded. There are also similarities of the spores soniaceae. 1. The genera Cystodium, Thyrsopteris,
with genera in the Dennstaedtiaceae, especially and Culcita. Am.]. Bot. 68: 808-819.
those of Saccoloma, and some ridged spores of

79. Thyrsopteris Kunze

Figs. 79.1-79.7
Genus
An American, monotypic genus based on Thyr- Figs. 79.1-79.7. Thyrsopteris elegans Kze. spores, Juan Fer-
sopteris elegans Kze. nandez, XI ,000, and wall sections. 1,2,4,5. Meyer 9393.
1. Coarse tapetal deposit on papillate distal face. 2. Papil-
late proximal face. 3. Surface detail, tapetal residue Solbrig
Range et at. 3801, X 5,000. 4. Papillate or microverrucate surface
with granulate deposit, distal face, X5,OOO. 5. Fractured
Endemic to the Juan Fernandez Islands. wall, exospore below thin, papillate peri spore with residual
tapetal deposits, X 5,000. 6, 7_ Solbrig et al. 3801. 6. Aper-
tural section, irregular, papillate peri spore above exospore
Spores with many canals, X 15,000. 7. Detail, two-layered peri-
spore above exospore (E), X50,OOO.
Examined in seven collections.
Size 42-57 /-Lm.
Shape Tetrahedral-globose with prominent probably tapetal residue. The papillate surface
angles. as well as some of the finer granulate material
is perisporal (Figs. 1-4, 6,7). The surface was
Aperture Trilete, 213 to 3f4 the radius. described as microverrucate, and the perisporal
Surface Granulate deposit on papillate (mi- composition of this material was confirmed by
croverrucate) surface (Fig. 4). treating Thyrsopteris spores with sodium hydrox-
Structure Plain exospore of two layers; peri- ide (Gastony, 1981). The exospore surface is
spore of two layers forms the papillate surface relatively plain (Fig. 5).
(Figs. 5-7). Relationships The granulate surface of Thyrsop-
teris spores is similar to that in spores of Dick-
sonia antarctica and D. sellowiana, as well as the
Comments
more strongly contoured spores of Cibotium and
Characteristics, Diversity The irregular, coarse Cystodium. This supports the alliance of Thyrsop-
deposit above the finely papillate surface is teris with the Dicksoniaceae. However, the
 Thyrsopteris 229

123

4 5

6 7
230 Dicksoniaceae

strongly dimorphic leaves of Thyrsopteris and Konijnenburg-Van Cittert, pers. comm., 1987).
unusual chromosome number of n = 76-78 are However, the relationship based on granulate
exceptional features of the plants, suggesting surface of the spores is uncertain.
that it may be an isolated element within the
family. Literature
Fossil spores of six species of Coniopteris
Gastony, G.]. 1981. The spore morphology in the
Brongn. and Kylikipteris arguta (Lind. & Hut- Dicksoniaceae. 1: the genera Cystodium, Thyrsopter-
ton) Harris, are reported from the Jurassic of is, and Culcita. Am.]. Bot. 68: 808-819.
Yorkshire and associated with Thrysopteris (Van

80. Dicksonia L'Heritier de Brutelle

Figs. 80.1-80.18
Genus
A genus usually of wet, montane forests, of Figs. SO.I-S0.9. Dicksonia spores, X 1,000 except Figs. I, 2,
about 20 species, mostly of the Old World. X900; 6, 8, X600. 1. D. antarctica Labill., densely granu-
late distal face at left, aperture above, at right, Australia, Strei-
mann H.S. 3232, X900. 2. D. squarrosa (Forst.) Sw.,
Range granulate deposit on pitted exospore, New Zealand, Tomlin-
son & Braggins, in 1969, X900. 3,4. D. sellowiana, Hook.
The three American species include Dicksonia 3. Aperture obscured by granulate material, Brazil, Reiss 61,
sellowiana, wide ranging from Mexico and Cen- X 1,000. 4. Detail of granulate deposit, Peru, Dudley 13254,
XIO,OOO. 5. D. squarrosa detail, granulate surface, as 2,
tral America, south to the Andes, and in south- X 10,000. 6, 7. D. sciurus C. Chr., Papua,
east Brazil; D. steubelii Hieron. in Peru; and D. Brass 22422. 6. Compact rodlets bordering reticulate ridges,
berteriana (Colla) C. Chr. in the Juan Fernandez proximal face at left, distal at right, X600. 7. Detail of
rodlets bordering an areole, X5,000. S. D. archboldii Co-
Islands. In the Old World from Malesia south- pel., irregularly reticulate ridges below sparse rodlets, New
ward to Tasmania and New Zealand, and in the Guinea, Hartley 11260, X600. 9. D. grandis Rosen., detail
Pacific, eastward to Samoa. of particulate deposit and rodlets, New Guinea, Brass
11897, X5,OOO.

Spores
more diffuse heterogeneous material (Figs. 17,
Eighteen species examined. 18), partly coalescent in rodlets, as in Dicksonia
Size 44-68 ~m. sciurus (Figs. 6, 7). The aperture is usually some-
what depressed (Figs. 2, 6, 16). Granulate peri-
Shape Tetrahedral-globose, the lobes some-
spore often obscures the plain or coarsely
times prolonged, with lateral surfaces often de-
reticulate exospore as in D. archboldii and D.
pressed between aperture arms.
antarctica (Figs. 1, 12, 17).
Aperture Trilete, % the radius. Diversity Species with plain spores, as in Dick-
Surface Plain or coarsely reticulate, with gran- sonia antarctica and D. sellowiana have promi-
ulate deposit, and rodlets (Figs. 6, 7). nent lobes, with large depressions between aper-
ture arms, and dense granulate surface (Figs. 1,
Structure Exospore of two layers, the outer
3). The reticulate spores of D. sciurus, D. arch-
plain or with low ridges and often irregular pits
boldii, and D. grandis are well filled and less an-
(Figs. 12, 18), the aperture depressed; peri-
gular (Figs. 6, 8). The inner exospore surface is
spore of two or more layers in and above pits,
essentially plain, but short diverticulated ele-
forming more or less consolidated rodlets.
ments, adjacent to the inner layer at the aper-
ture, suggest that this formation may be more
Comments
complex than the usual blechnoid type of struc-
Characteristics The irregular perispore consists ture (Fig. 10). This appears to represent a more
of inner compact material (Figs. 5, 13-15) or primitive type of blechnoid exospore.
Dicksonia 231
232 Dicksoniaceae

Relationships The granulate spores of Dicksonia Figs. SO.10-S0.IS. Dicksonia spore wall sections. 10-12. D.
antarctica, Hort., Kew 77 (1969), Lugardon 378 (TL). 10.
antarctica and D. sellowiana (Figs. 1, 3, 4) suggest
Central region of aperture, inner exospore with arrows on
a close relationship between these two American parts of sheets (feuillets) in contact with outer exospore,
species. The reticulate spores of D. berteriana perispore (P) incomplete, X35,OOO. II. Oblique section of
pitted exospore, small pits (arrows) partly filled with peri-
from Juan Fernandez and D. arborescens l'Herit.
spore, X30,OOO. 12. Exospore with irregularly scattered
of St. Helena are similar to and support alliances pits (arrow), X 12,000. 13, 14. D. squarrosa, New Zealand,
with Old World species. Both granulate and re- R. Hebant in 1973 (TL). 13. Irregular perispore above low
exospore ridges, X6,OOO. 14. Detail of granulate perispore
ticulate spores in American and Old World
above exospore (E), X50,OOO. 15, 16. D. sellowiana Hort.,
plants suggest that both types may have migrated Kew 79 (1969), Lugardon 383 (TL). 15. Granulate peri-
to America from the region of greater species spore above exospore (E), X50,OOO. 16. Apertural region,
the apex of aperture slightly raised, perispore (P) detached
concentration in the Old World.
from exospore, X 15,000. 17, IS. D. archboldii, New Guin-
The exospore pits were initially considered to ea, Hartley 11260. 17. Prominent exospore ridge, below
be irregularities in wall development (Lugardon, perispore (P) with rodlets, XI 0,000. IS. Exospore pits
filled and overlaid by perispore, rodlet sections with dark
1974). In these sections (Figs. 11, 18), they ap-
central core (arrow), X 21,000.
pear similar to the pitted exospore in some
spores of the Cyatheaceae. However, Cyathea-
ceae spores have abundant, regularly disposed
pits, and the perispore is often differentiated in
discrete strata. 34-61 J,Lm (Van Konijnenburg-Van Clittert,
The reticulate exospore is useful in recogni- 1989). Fossil records of these spores suggest a
tion of fossil spores. Dicksonia is reported from formerly broader range of the genus.
the Jurassic and Cretaceous of Europe based on
stems and fertile pinnae. Spores of D. crocina Literature
Bolkh. are described from Jurassic deposits, at Bolkhovitina, N.A. 1953. Spores and pollen character-
Yakutsk, in the Soviet Union, and D. densa Bolkh. istic of Cretaceous deposits of central regions of the
from the Lower Cretaceous of Kazakhstan (Bolk- USSR, 184 pp. Trans. Ceo!. Inst. Acad. Sci. USSR,
hovitina, 1953, 1956). The reticulate surface of Re!. Ceo!. Ser. 61.
these fossils appears similar to that of recent spe- Bolkhovitina, N.A. 1956. Atlas of spores and pollen
cies. Smooth spores of Late Jurassic and Lower from Jurassic and Lower Cretaceous deposits of the
Cretaceous age, from the Bureja Basin in the So- Vilyui Depression, 188 pp. Trans. Ceo!. Inst. Acad.
Sci. USSR 2.
viet Union, included by Krassilov (1978), are
Krassilov, V. 1978. Mesozoic lycopods and ferns from
more difficult to ally with Dicksonia.
the Bureja Basin. Palaeontg. 166: 16-29.
Spores of Dicksonia kendalli Harris, and D. Lugardon, B. 1974. See general literature.
mariopteris Wilson & Yates, from the Jurassic of Van Konijnenburg-Van Cittert, J.H.A. 1989. Dickson-
Yorkshire were placed in the genus on the basis iaceous spores in situ from the Jurassic of Yorkshire,
of the reticulate surface and spore size, between England. Palyno!. 67:273-301.
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