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STUDIA
GEOLOGICA
POLONICA
Vol. 128, Kraków 2008, pp. 5–96.
Geological Results of the Polish Antarctic Expeditions
Edited by K. Birkenmajer
Part XV
Krzysztof BIRKENMAJER1 and Anna Maria OCIEPA2
Plant-bearing Jurassic strata at Hope Bay,
Antarctic Peninsula (West Antarctica):
geology and fossil-plant description3, 4
(Figs 1–51; Tab. 1)
KRAKÓW 2008
1
2
3
4
Mailing address: Institute of Geological Sciences, Polish Academy of Sciences, Cracow
Research Centre, ul. Senacka 1, 31-002 Kraków, Poland. E-mail: [email protected]
Institute of Botany, Jagiellonian University, ul. Kopernika 27, 31-501 Kraków, Poland.
E-mail: [email protected]
Manuscript accepted for publication November 15th, 2007.
Key words: Hope Bay, Antarctic Peninsula, Jurassic, geology, fossil plant description.
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CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
A. Geological Part (K. Birkenmajer)
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Location . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Geological setting . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Basement of the Mount Flora Formation: The Trinity Peninsula
Group (?Permian) . . . . . . . . . . . . . . . . . . . . . . . . .
Botany Bay Group: The Mount Flora Formation (Jurassic). . . .
Antarctic Peninsula Volcanic Group: The Kenney Glacier
Formation (?Lower Cretaceous). . . . . . . . . . . . . . . . . .
Andean Intrusive Suite (Lower Cretaceous) . . . . . . . . . . .
B. Palaeobotanic Part (A. M. Ociepa)
Jurassic floras of Northern Antarctic Peninsula . . . . . . . . . . . . . .
The Polish Jurassic plant collections from Hope Bay . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic description. . . . . . . . . . . . . . . . . . . . . . . . . . .
Sphenophyta, Equisetales, Equisetaceae . . . . . . . . . . . . .
Pteridophyta, Filicales, Dicksoniaceae . . . . . . . . . . . . . .
Pteridophyta, Filicales, family unknown (Osmundaceae ?) . . . .
Pteridophylla . . . . . . . . . . . . . . . . . . . . . . . . . . .
Pteridospermophyta, Caytoniales, Caytoniaceae . . . . . . . . .
Pteridospermophyta, Corystospermales, Corystospermaceae . . .
Pteridospermophyta order and family unknown . . . . . . . . .
Cycadophyta, Cycadales . . . . . . . . . . . . . . . . . . . . .
Cycadophyta, Bennettitales (Cycadeoidales) . . . . . . . . . . .
Cycadophyta incertae sedis . . . . . . . . . . . . . . . . . . . .
Coniferophyta, Coniferales, Araucariaceae . . . . . . . . . . . .
Coniferophyta, Coniferales, Taxodiaceae . . . . . . . . . . . . .
Coniferophyta, Coniferales, Palissyaceae . . . . . . . . . . . . .
Coniferophyta, Coniferales, family unknown . . . . . . . . . . .
Comparison of the Hope Bay flora with Jurassic floras of Gondwana
and Laurasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Age of the Hope Bay flora based on fossil plants . . . . . . . . . . . . .
Palaeoclimatic aspects of the Hope Bay Jurassic flora . . . . . . . . . .
Palaeoenvironmental aspects of the Hope Bay Jurassic flora . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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PLANT-BEARING JURASSIC STRATA, ANTARCTICA
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Abstract
The fossil flora from Hope Bay, West Antarctica, is ranking among the richest Jurassic floras of
the world. So far, it was known mostly from loose blocks scattered at coastal plain below Mount Flora.
The present collections from the Mount Flora Formation, include the fossil plants assembled mainly
in situ (from the Flora Glacier Member), but also those obtained from loose blocks (derived from the
Five Lakes Valley Member and the Flora Glacier Member).
41 plant taxons were identified: 2 from Hepatophyta, 2 from Sphenophyta, 9 from Pteridophyta, 5
from Pteridophylla, 6 from Pteridospermophyta, 9 from Cycadophyta and 8 from Coniferophyta. The
taxons new for Antarctica include the species Coniopteris cf. simplex (Lindley & Hutton) Harris,
Equisetum cf. columnare Brongniart emend. Harris, Otozamites gramineus (Phillips) Harris, and the
genera Conites Sternberg emend. Cleal et Rees and Stachyotaxus Nathorst. New taxa include:
Crossozamia mirabilis sp. nov. Ociepa and Pagiophyllum arctowskii sp. nov. Ociepa (Schizolepidella
birkenmajeri sp. nov. Ociepa, 2007, has been described separately). A new combination Araucarites
antarcticus (Gee) comb. nov. Ociepa is presented.
The described flora confirms its Jurassic age. The paper presents also remarks on palaeoclimate
and environment of the Antarctic Peninsula volcanic arc during the Jurassic.
A. Geological Part
by Krzysztof Birkenmajer
INTRODUCTION
The plant-bearing beds at Hope Bay, Northern Antarctic Peninsula, were discovered in 1903 by J. G. Andersson (1906) during the famous Swedish Antarctic
1903–1904 Expedition led by Otto Nordenskjöld. The first complete description of
fossil plant collection by T. G. Halle (1913a) established its Jurassic age. Halle’s
collection was revised by Gee (1989b).
Further fossil plant collections were assembled by members of the British expeditions organized by the Falkland Islands Dependencies Survey and its successor –
the British Antarctic Survey, in particular by J. S. Bibby (1966), G. W. Farquharson
(1983, 1984), and P. M. Rees (in 1986/1987). They have been successively elaborated by Rees (1988, 1993a, b) and Rees and Cleal (1993, 1994). Sedimentological
aspects of the plant-bearing beds at Mount Flora, Hope Bay, were studied in particular by Farquharson (1983, 1984) and Elliott and Gracanin (1983).
During the 3rd Polish Geodynamic Antarctic Expedition (1987/8), leader Prof.
Aleksander Guterch, geology of the Hope Bay area was surveyed by K. Birkenmajer (1988, 1992, 1993a, b). It included, i.a., a detailed geological mapping, stratigraphic, sedimentological and tectonic studies, and collection of Jurassic plant
fossils. The latter were assembled both from loose blocks of clastic rocks scattered
on coastal plain below Mount Flora and, predominantly, from the beds in situ exposed above the Flora Glacier (at about 380 m a.s.l.), at northern rocky ridge of
Mount Flora. In this task, K. Birkenmajer was helped by W. Danowski.
Another collection of plant-bearing Jurassic blocks scattered on coastal plain at
Hope Bay, was assembled in 1991 by A. GaŸdzicki during the 4th Polish Geodynamic Expedition to West Antarctica (1990/1991).
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K. BIRKENMAJER & A. M. OCIEPA
LOCATION
Hope Bay is located at northern extremity of the Northern Antarctic Peninsula
(Fig. 1). This part of the Peninsula is also known as northern part of Graham Land
and Trinity Peninsula. Mount Flora, with exposures of plant-bearing Jurassic
strata, is situated between Kenney Glacier in the south, Buenos Aires Glacier in the
east, and Hope Bay in the west and north, about 2.5 km SW from the Argentine Base
“Esperanza”, and about 2 km SW from the abandoned British Base “D” (Fig. 2). In
horizontal plane, the mountain is crescent-shaped, dissected by a steep Flora Glacier which separates its two rocky ridges: eastern and western ones. The western
Fig. 1.
Position of Hope Bay in West Antarctica
11
Fig. 2. Surroundings of Mount Flora at Hope Bay. 1 – peaks and more important topographic
points; 2 – lakes; 3 – glacier front; 4 – rock exposures
ridge exposes a good section of the terrestrial plant-bearing strata, directly followed
by a column of stratified terrestrial volcanics (see Figs 3–7).
The W–E, NE–SW and NW–SE-trending faults delimit the exposures of the Jurassic plant-bearing beds to a triangle between Five Lakes Valley – Kenney Glacier
– Flora Glacier – Buenos Aires Glacier, with Mount Flora located in its southern
part (for new geographical names – see Birkenmajer, 1998). North of Mount Flora,
plant-bearing blocks of clastic rocks may be found scattered on coastal plain, in particular between Lake Boeckella and the Argentine Base “Esperanza” (Fig. 2).
GEOLOGICAL SETTING
The area of Hope Bay exposes four lithostratigraphic units of group rank (Birkenmajer, 1988, 1992, 1993a, b, 1994, 2001; Birkenmajer et al., 1995, 1997): the
Trinity Peninsula Group (Hope Bay Formation: ?Permian); the Botany Bay Group
(Mount Flora Formation: Jurassic); the Antarctic Peninsula Volcanic Group (Kenney Glacier Formation: ?Lower Cretaceous); and the Andean Intrusive Suite
(Lower Cretaceous).
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Basement of the Mount Flora Formation:
The Trinity Peninsula Group (?Permian)
Basement rocks. The basement of the Mount Flora Formation consists of metasediments of the Trinity Peninsula Group (TPG). This is a siliciclastic, predominantly turbidite complex more than 1200 m thick. The Trinity Peninsula Group
(“Trinity series” of Klimov, 1964: fide Tingey, ?1982; Birkenmajer, 2001), was
further described as the “Trinity Peninsula Series” (Adie, 1957), and the Trinity
Peninsula Group (e.g., Fleming & Thomson, 1979; Hyden & Tanner, 1981). At
Hope Bay, the TPG rocks are represented by the Hope Bay Formation (Hyden &
Tanner, 1981), redescribed and formally subdivided by Birkenmajer (1992) into
three lithostratigraphic members: (1) the Hut Cove Member (lower); (2) the Seal
Point Member (middle); and (3) the Scar Hills Member (upper).
(1) The Hut Cove Member consists of dark-coloured marine metaturbidite sequence of sandstones and shales (rhythmites), with intraformational shale-clast
slump breccias. The member is over 500 m thick (base not exposed).
(2) The Seal Point Member consists od dark-coloured marine metaturbidite sequence of sandstone and shale (rhythmites), with plant detritus, with subordinate
shale slump-breccia intercalations. It is 170–200 m thick.
(3) The Scar Hills Member consists of grey to rusty-weathered quartzitic sandstone rich in plant detritus, probably deposited as a submarine delta. The member is
over 550 m thick, its top is not exposed (see Birkenmajer, 1992).
Fig. 3.
Geological map of Mount Flora and vicinity (after Birkenmajer, 1993a)
Fig. 4.
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Hope Bay, Antarctic Peninsula: Mount Flora in the middle (phot. K. Birkenmajer)
Geological age. The age of the TPG at Hope Bay and elsewhere in Antarctic
Peninsula is under debate (see discussion in Birkenmajer, 1992, 2001). At Hope
Bay, the Hope Bay Formation may represent the Permian (?Upper Permian), while
elsewhere in the Peninsula it might also include the Triassic strata.
Botany Bay Group: The Mount Flora Formation (Jurassic)
The unit was discovered by Andersson (1906) during the Swedish (1901–1904)
South Polar Expedition led by O. Nordenskjöld. The name “Mount Flora plant
beds” has been used by Adie (1957). The present formal name is in use since 1980
(Caminos and Massambie, 1980 – fide Farquharson, 1984; Elliot & Gracanin,
1983). Closer descriptions of the unit were provided by Bibby (1966), Farquharson
(1983, 1984), Elliot and Gracanin (1983) and Birkenmajer (1988, 1993a, 2001).
This is a terrestrial clastic sedimentary wedge which grows from 230 m in the
north, to 270 m in the south (Birkenmajer, 1993a).
Much larger thickness values were given by Bibby (1966: 305.5 m), Elliot and Gracanin (1983:
366 m) and Farquharson (1983: 370 m). According to Birkenmajer (1993a), there is a repetition of
some strata by faulting, and the formation’s thickness cannot include the thicknesses of acidic sills
intruded in plant-bearing beds.
Angular unconformities divide the Mount Flora Formation from the underlying
Hope Bay Formation (TPG), and the overlying Kenney Glacier Formation (Antarctic Peninsula Volcanic Group).
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Fig. 5. Perspective view of Mount Flora, Hope Bay. TP – Trinity Peninsula Group (Hope Bay
Formation: ?Permian); MF1, 2 – Mount Flora Formation (Jurassic): MF1 – Five Lakes Member; MF2 –
Flora Glacier Member; KG1–KG4 – Kenney Glacier Formation (?Lower Cretaceous), successive
volcanic units; s – acidic sills; d – basic dykes (after Birkenmajer, 1993a)
Basal unconformity. It is generally accepted that an angular unconformity
separates coarse terrestrial clastics of the Mount Flora Formation from folded and
eroded marine siliciclastic deposits of the Hope Bay Formation (TPG) – see Adie
(1964); Bibby (1966); Farquharson (1983, 1984); Elliot & Gracanin (1983); Birkenmajer (1992, 1993a). This unconformity is evident from different attitudes of
strata of the two formations (Birkenmajer, op. cit.): the folded Hope Bay Formation
strata west and north of Mt Flora dip at 20–30°W, while those of the Mount Flora
Formation (Five Lakes Member – see below) – at 40–50°S (Fig. 3).
At western ridge of Mt Flora, basal beds of the Mount Flora Formation directly
contact with the middle member of the Hope Bay Formation (= the Seal Point Member). This attests to a long break in deposition between the two formations, during
which folding and subsequent erosion affected the older one. This unconformity
corresponds with the Trinity Phase folding (Birkenmajer, 1994a, b) of the Gondwanian (Smellie, 1981) vel Peninsula Orogeny (Smellie, 1991) that had occurred
probably close to the Triassic/Jurassic boundary, and was followed by intense erosion of the TPG deposits.
Subdivision of the Mount Flora Formation. Two lithostratigraphic units have
been distinguished in the Mount Flora Formation: (1) the Five Lakes Member; and
(2) the Flora Glacier Member (Birkenmajer, 1993a, 2001).
(1) The Five Lakes Member. This is the lower member of the Mount Flora Formation, maximum 170 m thick (Figs 3, 5–7). It begins with coarse polymict basal
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Fig. 6. Perspective view of Mount Flora, as seen from left morainic ridge (250 m a.s.l.) of the Flora
Glacier, Hope Bay. MF1, MF2 – Mount Flora Formation (Jurassic): MF1 – Five Lakes Member; MF2 –
Flora Glacier Member. Conventional signatures: shales – obliquely ruled; sandstones – stippled;
conglomerates and sedimentary breccias – heavy dots; clover marks main plant-bearing horizon; m –
moraine; f – faults; s – acidic sills; metres a.s.l. (after Birkenmajer, 1993a)
conglomerate consisting of sandstone and shale fragments derived from the the
Hope Bay Formation (TPG), grey marble, white and greenish granitoid, green volcanic rocks and red chert.
The basal conglomerate is followed by breccia-conglomerate-sandstone beds in
which three facies have been distinguished: (i) coarse clast-supported sedimentary
breccia and conglomerate; (ii) pebble to cobble lag-conglomerate; (iii) planar
cross-bedded sandstone. The clastic material of these strata was derived almost exclusively from the Hope Bay Formation. Plant remains occur frequently in all types
of clastic rocks (i–iii).
(2) The Flora Glacier Member. The Five Lakes Member is conformably followed by the Flora Glacier Member which is 60 m thick (Figs 3, 5–7). This is a terrestrial clastic unit consisting of two subunits: (i) the lower subunit, some 50 m
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Fig. 7. Hope Bay: Mount Flora Formation, lithostratigraphic-sedimentological column (after
Birkenmajer, 1993a)
thick, consists of grey to black sandstone-conglomerate beds alternating with siltstone and silty shale rich in plant detritus; pebble-lag concentrates form bases of
sandstone layers and channel fills; (ii) the upper subunit, 10 m thick, consists of
black shale with thin intercalations of finely cross-laminated sandstones, sometimes also of breccia-conglomerate. All these clastic rocks are very rich in plant remains.
Top unconformity. An angular unconformity separates the overlying stratified
volcanics of the Kenney Glacier Formation from the plant-bearing Mount Flora
Formation (Birkenmajer, 1993a, b, 2001) – Figs 5–7. The contact surface between
17
the two formations dips at about 20–25 degrees south, while the underlying beds of
the Flora Glacier Member are inclined some 45 degreees in the same direction. As
observed from a distance (from Five Lakes Valley), the angle of unconformity between the formations amounts to about 20 degrees (Birkenmajer, 1993a, p. 33).
Geological age. Based on rich plant remains, a Middle Jurassic age of the
Mount Flora Formation has been proposed by Andersson (1906) and Halle (1913a).
Stipanicic and Bonetti (fide Thomson, 1977: pp. 888–889) argued for a Late Jurassic or Early Cretaceous age. Rees (1988) proposed a Middle to Late Jurassic age,
later corrected by him (Rees, 1993a) to Early Jurassic.
There is no direct radiochronologic data available thus far from volcanics of the
Kenney Glacier Formation, and from acidic sill intrusions (possibly also tuffs) inserted in the Mount Flora Formation, to help decide whether it is of Early or Middle
Jurassic age. Birkenmajer (1993a) suggested that sills which intrude the Mount
Flora Formation might correlate in age with volcanic phase of the Kenney Glacier
Formation (“Early Cretaceous”), and with Jurassic acidic volcanics at Cape Dubouzet (northernmost tip of Trinity Peninsula).
At Cape Dubouzet, some 20 km NW from Mount Flora, occurs a comagmatic suite of rhyolitic
breccias, lava flows and ignimbrites more than 200 m thick, intruded by several tonalite-diorite
stocks, by a banded rhyolitic dome, and by dykes of andesitic composition (Hervé et al., 1991).
Radiometric ages: 158±22 Ma (Rb-Sr date: Middle Jurassic–Early Cretaceous) from a rhyolite; about
140 Ma (141±5 Ma and 147±6 Ma Rb-Sr dates: Jurassic–Cretaceous boundary) for contact metamorphic overprint in xenoliths contained in a tonalite; and about 100 Ma (Early/Late Cretaceous boundary) for the tonalite-diorite stocks (Hervé, 1992; Hervé et al., 1991; Hole et al., 1991; Loske &
Miller, 1991), indicate that the volcanic and plutonic activity at Cape Dubouzet spanned the Middle
through Late Jurassic and Early Cretaceous time.
Palaeotectonic setting and palaeoenvironment. The fining-upward terrestrial
clastic sequence of the Mount Flora Formation was deposited as an alluvial fan
bounded in the north by a high, probably fault-controlled morphological scarp of
the Trinity High – built of the TPG metasediments (Elliot & Gracanin, 1983; Birkenmajer, 1993a). Coarse to very coarse, chaotic, predominantly monomict, clastsupported sedimentary breccias of the Five Lakes Member are evidences of landslides which involved talus pediment aprons along the southern margin of the Trinity High. The landslides could have been triggered by seasonal heavy rainfall.
The debris-flow fan was being simultaneously dissected by an immature, probably braided, river system. Pebble-lag concentrates were laid down in shallow fluvial channels, while planar cross-bedded sands formed point bars at river meanders.
The alluvial fan deposition was followed by a short-lived inland lake (Flora Glacier Member, upper part). Quiet lacustrine deposition of dark silty clay was seldom
disturbed by turbidity currents which deposited fine, laminated and rippled sand
and silt. Occasional small-scale slumping produced slump-balls (Birkenmajer,
1993a).
Mode of occurrence of plant remains. The plant association in the Mount
Flora Formation mainly represents a thanatocoenosis. Fragmented pteridophyte
fronds, cycadophyte leaves, twigs and branches of conifers etc. were transported
18
from their original habitat to the site of deposition (alluvial fan) mainly by seasonal
running water, partly by landslides. Tree stumps in growth position are an exception: they occur in deposits of sheetflood (Elliot & Gracanin, 1983). The delta itself
was, however, barren of permanent vegetation cover, as is indicated by the lack of
palaeosol (regolith) and rootlet zones (Birkenmajer, 1993a).
The top shale thanatocoenosis (upper part of the Flora Glacier Member), with
very well preserved rich plant association derived from different habitats – from
dryer mountain slopes, from river floodplain, even from a tidal mangrove zone,
points to a lush vegetation cover of rain forest-type in a monsoon-affected part of
the Antarctandes adjacent to the Pacific Ocean (see part B of this paper). The thanatocoenosis had formed in much more quiet sedimentary environment than that of
the underling coarse beds, probably in a fresh-water lake with, at least temporarily,
stable water level. This is evidenced by the occurrence of water ferns, fish vertebrae, moreover scarce, probably fresh-water molluscs, and beetles’ elytra belonging to groups living in fresh water (Andersson, 1906; Halle, 1913a; Elliot & Gracanin, 1983; Zeuner, 1959).
Volcanic activity. Rhyolitic and related volcanic centres were located in the
Trinity High area at Cape Dubouzet, and along inner margin of the back-arc basin
(Hope Bay, Joinville Island), on both sides of the supposed Sobral-Joinville strikeslip fault (Birkenmajer, 1993a). From these centres could have derived some acidic
tuffs reported from upper part of the Mount Flora Formation by Farquharson
(1983) and Elliot and Gracanin (1983), but not the “ignimbrites” (up to 26 m thick)
of Farquharson (1984) which were recognized as sill intrusions related to a younger
(?Early Cretaceous) Kenney Glacier Formation volcanic phase (Birkenmajer,
1993a, b).
Antarctic Peninsula Volcanic Group: The Kenney Glacier Formation
(?Lower Cretaceous)
The Mount Flora Formation is unconformably covered by a stratified volcanic
complex some 200 m thick (Figs 3–7), consisting of predominantly acidic, rhyolite-dacite lavas, ignimbrites, agglomerates and tuffs, with subordinate chert and
shale intercalations, occasionally with petrified wood fragments (Birkenmajer,
1993b, 2001). This is the basal unit of the Antarctic Peninsula Volcanic Group, first
recognized by Andersson (1906) and studied in particular by Bibby (1966), Birkenmajer (1993b) and Birkenmajer et al. (1995, 1997). The age of the formation represents either the Upper Jurassic–Lower Cretaceous or Lower Cretaceous.
Andean Intrusive Suite (Lower Cretaceous)
The youngest group of magmatic rocks in the area of Hope Bay is represented by
intrusive gabbro to tonalite plutonic bodies of the Andean Intrusive Suite (Birkenmajer, 1993b, 2001; Birkenmajer et al., 1995, 1997) – Fig. 3. According to
Pankhurst (1982), the peak of this plutonic phase in Northern Antarctic Peninsula is
of late Lower Cretaceous age (100 Ma).
19
Small-scale melanocratic, mainly basaltic, dykes which intrude the plutons,
might be of mid-Cretaceous or younger (?Late Cretaceous) age.
B. Palaeobotanic Part
by Anna Maria Ociepa
JURASSIC FLORAS OF NORTHERN ANTARCTIC PENINSULA
Research on Jurassic floras of Antarctica is already over 100 years old. The first
discovery of Jurassic plant fossils was made by J. Gunnar Anderson, a geologist of
the famous Swedish Antarctic Expedition of 1901–1903 led by Otto Nordenskjöld.
Anderson and his two companions were left at Hope Bay, northern tip of Antarctic
Peninsula (vel Graham Land, vel Trinity Peninsula) completely unprepared for
wintering in the Antarctic, after the expedition’s ship Antarctic was crushed by ice
and drowned in a nearby strait, known since under the name of Antarctic Sound
(Nordenskjöld & Andersson, 1905). The brave Swedes, hoping in Hope Bay for the
rescue that fortunately came next year, not only survived the severe Antarctic winter crowded in a small shack built by them from loose stones, covered with seal
skins for the roof, their menu consisting mainly of penguin and seal meat, but had
assembled a unique collection of Jurassic plant fossils from blocks scattered on
coastal plain of Hope Bay below the fossil-bearing Mount Flora (Andersson,
1906). This was the first Jurassic flora ever collected from Antarctica, a milestone
in palaeobiosphere reconstruction history of Antarctica.
Andersson’s collection was elaborated in detail by T. G. Halle (1913a) who had
distinguished 24 genera with 59 species and 2 unnamed plant taxa. His monograph
is ranked among the classic works on Jurassic palaeofloras of the Gondwana.
Taxonomic revision of Halle’s monograph was undertaken by C. T. Gee
(1989b) who has likewise distinguished 24 genera but reduced the number of species to 43, of which 6 were new species.
New Jurassic plant collections were assembled from Hope Bay and Botany Bay
(NE Antarctic Peninsula) by members of the British Antarctic Expeditions: W. N.
Croft in 1946, G. W. Farquharson in 1979/1980, and P. M. Rees in 1986/1987 (Rees
& Cleal, 2004). A part of the Botany Bay collection was assembled in situ from
plant-bearing beds. The British collections, including some 2000 specimens, have
been determined by Rees (1993a–d) and Rees and Cleal (1993, 2004). Jurassic
wood fragments from Hope Bay have been elaborated by Francis and Poole (2002).
THE POLISH JURASSIC PLANT COLLECTIONS FROM HOPE BAY
The palaeobotanic part of the present paper is based on five collections of Jurassic plant-bearing rocks from Hope Bay, Antarctic Peninsula, assembled by K. Birkenmajer and A. GaŸdzicki – members of the Polish Antarctic Expeditions.
20
K. Birkenmajer’s Collections (1–4)
These collections, consisting jointly of 234 plant-bearing rock specimens, were
assembled in 1988 during the 3rd Polish Geodynamic Expedition to West Antarctica, 1987/8 (Birkenmajer, 1988, 1993a):
(1) 174 plant-bearing rock specimens (Collection S78 KRAM-P) were collected
on Mount Flora, western ridge, in situ, at c. 380 m a.s.l., from the highest plantbearing bed of the Flora Glacier Member (see Figs 5–7);
(2) 25 plant-bearing rock specimens (Collections: S79 and S82 KRAM-P) were
assembled on coastal plain north of Mt Flora, as loose blocks of shaly sandstone/siltstone/shale derived from the Flora Glacier Member;
(3) 4 plant-bearing rock specimens (Collection S80 KRAM-P) were assembled
on coastal plain north of Mt Flora as loose blocks of coarse-grained sandstone/fine
conglomerate derived from the Five Lakes Member;
(4) 40 plant-bearing rock specimens (Collection S81 KRAM-P) were assembled
on coastal plain north of Mt Flora as loose blocks of sandstone derived from the
Mount Flora Formation, undifferentiated.
The collections 1–4 have been donated by K. Birkenmajer to the W. Szafer Institute of Botany, Polish Academy of Sciences, in Cracow (ul. Lubicz 46, 31-512
Kraków, Poland) where they are being housed as the Collections numbered: S78, 79,
80, 81, 82 KRAM-P.
A. GaŸdzicki’s Collection (5)
(5) This collection was assembled in 1991 during the 4th Polish Geodynamic
Expedition to West Antarctica, 1990/91, from loose blocks of Jurassic plantbearing rocks scattered on coastal plain of Hope Bay (GaŸdzicki, 2003). The collection consists of 20 rock specimens derived from the Mount Flora Formation, undifferentiated. It is housed in the Institute of Palaeobiology, Polish Academy of Sciences in Warsaw (ul. Twarda 51/55, 00-818 Warszawa, Poland) as the Collection
ZPAL PAN.
Acknowledgements
The author of palaeobotanical part is deeply indebted to Prof. K. Birkenmajer and Prof. A.
GaŸdzicki for entrusting her with elaboration of their Jurassic plant collections from Hope Bay. Prof.
Danuta Zdebska (Institute of Botany, Jagiellonian University, Cracow), as the Promotor of the
author’s Ph. D. degree (botany), most kindly supervised her work with the fossil plants. The Referees,
Prof. Adam Zaj¹c (Institute of Botany, Jagiellonian University, Cracow) and Prof. Ewa Zastawniak
(W. Szafer Institute of Botany, Polish Academy of Sciences, Cracow), offered much constructive
criticism. Dr Maria Barbacka (Hungarian Natural History Museum, Budapest) is warmly thanked for
valuable discussions and for her hospitality while in Hungary. Prof. Johanna H. A. van Konijnenburg-van Cittert (National Natural History Museum “Naturalis”, Leiden) most kindly supplied the
author with some important but difficult to obtain papers.
21
SYSTEMATIC DESCRIPTION
This part of the paper includes systematic description of the whole Jurasic
plant-fossil collection (1–5 – see above) assembled by the members of the 1987/8
and 1990/91 Polish Antarctic Expeditions at Hope Bay, northern Antarctic Peninsula, exlusively of Hepatophyta which have been published separately (Ociepa,
2007).
Division Sphenophyta
Order Equisetales
Family Equisetaceae
Genus Equisetum Linné, 1753
Type species: Equisetum fluviatile Linné, 1753
The genus Equisetum is presently widely distributed from the tropics to the polar circle, most species occurring between latitudes 40 and 60 degrees North; it is
unknown from Australia, New Zealand and Antarctica. Presently, about 15 species
are known (Kramer & Green, 1990). The genus Equisetum is known since Jurassic
(Meyen, 1987): it includes well preserved specimens which do not significantly
differ from modern species; poorly preserved specimens are included to the genus
Equisetites. From other representatives of fossil horsetails, the genus Equisetum
differs in having fused leaves which form a leaf sheath.
Equisetum laterale Phillips, emend. Harris, emend. Gould 1968
(Figs 8, 9, 10B)
1905
1913a
1961
1965
1967
1968
1989b
2004
Equisetum phillipsii (Dunker) Brongniart; Ward, pp. 298-301; pl. 72, figs 1-11
Equisetites approximatus Nathorst n. sp.; Halle, pp. 6-8; text-fig. 1; pl. 1, figs 6-14
Equisetum laterale Phillips; Harris, pp. 20-24; text-fig. 5A, B (cuticle), C, D, G
Equisetites patagonica n. sp. Herbst; Herbst, pp. 29-31; pl. 1, figs 1, 3; pl. 2, figs 9, 10
Equisetites lateralis Phillips; Semaka & Georgesco, pp. 734-735, fig. 6
Equisetum laterale Phillips; Gould, pp. 157-168, figs 2, 3; pl. 1, figs 1-22; pl. 2, figs 1-18
Equisetum laterale Phillips, emend. Gould; Gee, pp. 157-158; pl. 1, figs 3-6
Equisetum laterale Phillips, emend. Gould; Rees & Cleal, pp. 8-10; pl. 2, figs 1-4
Material. A dozen or so fragmental impressions of diaphragms, a flattened stem
and a leaf sheath. Specimens: S82/5, 6 (positive); S82/6 (negative); S82/7, S82/12
KRAM-P; ZPAL Pl. 3/6, 3/7 (positive and negative); ZPAL Pl. 3/8, 3/11, 3/15,
3/16, 3/17.
Description. Shoot not branched, 11 mm broad, 52 mm long, with one node
preserved (Fig. 10B). Internode incomplete. Flattened, broken leaf sheaths are
30 mm long and 5–10 mm broad (denticulation inclusively). Denticulation consists
of teeth 4 mm long and 1–2 mm broad, about 20 per leaf. Teeth triangular,
sharpened at apex. Different size foramina of axial and carinal channels visible at
diaphragms. Diaphragm diametres – 3.5–9 mm, those of axial canal foramina –
1–3 mm. Carinal channels number from 15 to more than 30.
22
Fig. 8. Equisetum laterale Phillips
emend. Harris, emend. Gould, diaphragms, ZPAL PL 3/6: a – axial channel; b –
carinal channel. Scale bar 1 mm
Discussion. Attribution of the present specimens to the species Equisetum laterale is
based on length of internode which is over
52 mm long, its diameter and, first of all,
presence of diaphragms showing axial
channel foramen and carinal channels.
E. laterale has shoots narrower than
those of E. beani and E. columnare. In these
two species, the diaphragm is uniform, de-
Fig. 9. Equisetum laterale Phillips emend. Harris, emend. Gould, diaphragms. A – ZPAL PL 3/16;
B – ZPAL PL 3/11; C – ZPAL PL 3/7. Scale bar 1 mm
23
Fig. 10. A – Equisetum cf. columnare Brongniart, emend. Harris, stem with nodes and internodes,
S81/29 KRAM-P; B – Equisetum laterale Phillips emend. Harris emend. Gould, stem with nodes,
ZPAL PL 3/17. Scale bar 1 cm
void of either axial or carinal channels’ foramina (see Harris, 1961). E. laterale
shows smaller number of leaf denticulation as compared to Equisetites contectum
(see Gould, 1968).
Distribution. Known from Triassic to Cretaceous of Laurasia and Gondwana
(Gould, 1968; Jongmans & Dijkstra, 1970). In West Antarctica, it has been reported
from Jurassic terrestrial deposits at Hope Bay and Botany Bay, northern Antarctic
Peninsula (Gee, 1989b; Rees & Cleal, 2004).
Equisetum cf. columnare Brongniart, emend. Harris, 1961
(Fig. 10A)
1961
1967
1999
Equisetum columnare Brongniart; Harris, pp. 15-26; text-figs 4, 5E, F, I, J
Equisetites columnaris Brongniart; Semaka & Georgesco, p. 732, fig. 5
Equisetum columnare Brongniart; van Konijnenburg-van Cittert & Morgans, pp. 35-36;
text-figs 13B, 14
24
Material. Specimen S81/39 KRAM-P.
Description. Shoots not branched. Internodes 40–70 mm long and 26–30 mm
broad. Leaf sheaths c. 10 mm long and 27–31 mm broad, visible at nodes. 16–18
fused leaves visible in leaf sheaths. Leaves c. 2 mm broad. Leaf denticulation not
well visible.
Discussion. Sizes, especially width of shoots, are diagnostic features in Mesozoic
horsetails. In our specimens, width of shoots, 26–30 mm, indicates its attribution to
the species E. columnare. Wider shoots are known from E. beani – c. 100 mm,
while narrower shoots, c. 15 mm, from E. laterale (Harris, 1961). Our specimens
show few morphological details preserved: e.g., leaf denticulation is unclear and
diaphragm is missing. This does not allow for more certain specific determination.
Distribution. Triassic in Italy; Jurassic in England, Austria, France, Germany,
Poland, Romania, Italy, Caucasus, Siberia, China; Cretaceous in Siberia (Semaka
& Georgesco, 1967).
Division Pteridophyta
Order Filicales
Family Dicksoniaceae
Presently, it is a relictic family consisting of six genera of large to very large, often arborescent ferns, predominantly occurring in lower tiers of woods, in shrubs,
moreover in more open habitats, often in mountainous areas, first of all in the tropics, moreover in temperate-warm climatic zones (Kramer & Green, 1990). Fossil
Dicksoniaceae first appeared during Triassic (Skog, 2001), they were widely distributed during Jurassic (Taylor & Taylor, 1993): e.g., in the Jurassic deposits of
Yorkshire they were dominant (Harris, 1961).
Genus Coniopteris Brongniart, emend. Harris, 1961
Type species: Coniopteris murrayana (Brongniart) Brongniart, emend. Harris, 1961
The majority of palaeobotanists (e.g., Harris, 1961; van Konijnenburg-van Cittert & Morgans, 1999) attribute to the genus Coniopteris both fertile and sterile
specimens. They elaborated special keys to help determine fertile specimens of this
genus, and inculded to synonimic lists also those which are known from sterile fragments. This view is being also shared by the present author. On the contrary, Gee
(1989b) includes to the genus Coniopteris the fertile specimens only.
Coniopteris lobata (Oldham et Morris) Halle, 1913
(Figs 11A, 12A)
1862
1877a
1913a
Pecopteris (?) lobata n. sp. Oldham & Morris; Oldham & Morris, pl. 28, fig. 1; pls 29,
30
Pecopteris lobata Oldham et Morris; Feistmantel, p. 92, pl. 36, fig. 3
Coniopteris ? lobata (Oldham et Morris) Halle; Halle, pp. 22-24; text- fig. 5; pl. 1, fig.
27?; pl. 3, fig. 13?
25
Fig. 11. A – Coniopteris lobata (Oldham et Morris) Halle, S78/131 KRAM-P; B – Coniopteris cf.
simplex (Lindley et Hutton) Harris, S82/11 KRAM-P. Scale bar 2.5 mm
1913a
1988
1989b
2004
2004
Coniopteris cf. nephrocarpa (Bunbury); Halle, pp. 21-22; pl. 3, figs 11, 11a, b
Pecopteris lobata Oldham et Morris; Sengupta, pl. 13, fig. 36
Coniopteris meridionalis n. sp. Gee; Gee, pp. 160-161; pl. 1, figs 10-12
Coniopteris meridionalis Gee; Ociepa, pl. 1, fig. 1
Coniopterios lobata (Oldham et Morris) Halle; Rees & Cleal, pp. 16-19; pl. 5, figs 1, 2, 4
Material. Specimen S78/131 KRAM-P.
Description. Frond fragment (penultimate section), triangular in outline, narrowing from base to apex. Axis of penultimate section 0.5 mm broad. Pinnules 7 mm
long and 4 mm broad, distinctly separated, deeply lobed. Eight lobes on pinnules.
Lobes rounded or roundish. Pinnules slightly overlapping. In apical part of frond,
pinnules indistinctly separated, forming a single lobe. Midvein passing along
middle parts of penultimate and ultimate sections.
Discussion. The size of the specimen, triangular outline of penultimate frond
section and, first of all, rounded to roundish lobes of pinnules, are the features
typical of the species Coniopteris lobata.
Our frond imprint is nearly identical with that of Coniopteris? lobata presented
by Halle (1913a, pl. 3, fig. 13).
Gee (1989b) has described a new species Coniopteris meridionalis based on
specimens from Hope Bay. She included to this species the forms determined by
Halle (1913a) as C.? lobata and C. cf. nephrocarpa. To the first species, Halle included fertile specimens, to the second species – sterile ones. He also considered a
possibility that both belonged to the same species. Gee (1989b) created a new species C. meridionalis which included both taxons. Rees and Cleal (2004), while accepting her conclusion, have decided to return to the long-established species Coniopteris lobata. The present author follows their solution, as our frond fragment is
very similar both in outline and size to those presented by Oldham and Morris
(1862) and Sengupta (1988).
26
Fig. 12. A – Coniopterios lobata (Oldham et Morris) Halle, S78/131 KRAM-P; B – Coniopteris
murrayana (Brongniart) Brongniart emend. Harris, S78/6a KRAM-P. Scale bar 2.5 mm
Halle (1913a) was of the opinion that fronds of C.? lobata are generally similar
to those of Sphenopteris nauckhoffiana. However, as pointed out by Gee (1989b),
fronds of the latter taxon show the presence of shallow indentitions in particular
lobes of ultimate frond sections, a feature unknown in C. lobata.
By the presence of round to roundish lobes in ultimate frond section, our specimen resembles fronds of C. bella and C. murrayana. However, penultimate sections of fronds of C. bella are not triangular in outline, while fronds of C. murrayana – though having triangular penultimate frond sections, are obovate, and not
rounded to roundish, lobes as is the case with C. lobata (see Harris, 1961; Rees and
Cleal, 2004).
Distribution. North America: Hawkesbury series – Triassic. Asia: Caucasus –
Liassic, Fergans – Mesozoic. New Zealand: Waikato – Jurassic (Jongmans & Dijkstra, 1959b); West Antarctica: Hope Bay and Botany Bay, Antarctic Peninsula – Jurassic (Halle, 1913a; Rees & Cleal, 2004); India: Rajmahal Hills – Jurassic/Cretaceous (Oldham & Morris, 1862; Sengupta, 1988).
Coniopteris murrayana (Brongniart) Brongniart, emend. Harris, 1961
(Figs 12B, 13)
1913a
1961
1989b
Coniopteris hymenophylloides (Brongniart); Halle, pl. 3, fig. 27, 28, 28a
Coniopteris murrayana (Brongniart) Brongniart; Harris, pp. 158-164; figs 56-58
Coniopteris murrayana (Brongniart) Brongniart, emend. Harris; Gee, pp. 161-162; pl. 1,
fig. 9
27
Fig. 13. Coniopteris murrayana (Brongniart) Brongniart emend. Harris, S78/89 KRAM-P. Scale bar
2.5 mm
1999
2003
2004
Coniopteris murrayana (Brongniart) Brongniart; van Konijnenburg-van Cittert &
Morgans, p. 53; text-fig. 23E-F; pl. 3, fig. 2; pl. 20, fig. 1
Coniopteris murrayana (Brongniart) Brongniart; Meng et al., pl. 3, figs 5, 6; pl. 4, fig. 9
Coniopteris cf. murrayana (Brongniart) Brongniart, emend. Harris; Rees & Cleal, pp.
19-20, pl. 5, figs 3, 5
Material. Two imprints of sterile fronds, consisting of penultimate and ultimate
sections. Samples S78/6a, b (positive and negative), S78/89 KRAM-P.
28
Description. Rachis of composite frond 1 mm broad, axis of penultimate section
0.5–1.5 mm broad. Penultimate sections usually triangular, those of ultimate
sections – elongated, 5–8 mm long and 2–2.5 mm broad. Ultimate sections form an
angle of 30–40° with the axis, alternating or nearly opposite, slightly overlapping
(Fig. 12B). They are divided in 5–7 deeply incised lobes. Lobes obovate, rounded
or sharpened at apex, 1.5–2 mm long, 9.5–1.5 mm wide. Nervation not visible.
Discussion. Triangular outline of penultimate sections, their overlapping, obovate
shape of lobes, indicate attribution of the specimens to Coniopteris murrayana.
Imprint of frond shown in Fig. 12B is very similar to that illustrated by Rees and
Cleal (2004, pl. 5, figs 3, 5), determined by them as C. cf. murrayana. Imprint of a
frond shown in Fig. 13 is similar to those illustrated by Harris (1961, text-fig. 55A
and text-fig. 56C), determined by him as Coniopteris murrayana.
Our two frond imprints from Hope Bay differ from one another in size, and show
a variation in morphology at ultimate sections. Frond fragments of C. murrayana
from other locations are much larger, may show sections with 4–5-order branches,
and variation in size and shape of particular sections of the last order. Differences in
our specimens may have resulted from their original location in the fronds: e.g., at
apex or at base.
Triangular penultimate sections, and rounded lobes of Coniopteris murrayana
resemble those of C. lobata, however, in the latter species, the lobes are halfrounded or rounded, while in the former one they are obovate (cf. Halle, 1913a;
Harris, 1961; Rees & Cleal, 2004).
Fronds of C. murrayana are also similar to those of C. hymenophylloides. However, ultimate frond sections of the former species may overlap, a feature almost unknown in the latter one (see van Konijnenburg-van Cittert & Morgans, 1999).
Fronds of C. murrayana morphologically resemble also those of Sphenopteris
antarctica. However, lobes of ultimate section in C. murrayana are obovate, while
those of S. antarctica are triangular (Halle, 1913a).
Distribution. Europe: Yorkshire, England – Jurassic (Aalenian–Bajocian) (van
Konijnenburg-van Cittert & Morgans, 1999); China – Lower/Middle Jurassic
(Meng et al., 2003); West Antarctica: Hope Bay – Jurassic (Gee, 1989b; Rees &
Cleal, 2004).
Coniopteris cf. hymenophylloides (Brongniart) Seward, emend. Harris, 1961
(Fig. 14B–F)
1894
1894
1913a
1934
1961
1984
1989b
Dicksonia (Eudicksonia) heerii Raciborski, pp. 174-175; pl. 10, figs 7-11a, 14
Thyrsopteris (?) murrayana Brongniart; Raciborski, pp. 180-181; pl. 10, figs 15, 16; pl.
12, figs 17-20
Coniopteris hymenophylloides (Brongniart); Halle, pp. 19-21; pl. 3, figs 23, 24, 29, 30
Coniopteris hymenophylloides (Brongniart); Edwards, pp. 92-93; text-fig. 2 (spores)
Coniopteris hymenophylloides (Brongniart) Seward; Harris, pp. 152-158; figs 53, 54
Coniopteris cf. hymenophylloides (Brongniart) Seward; Bose & Banerji, pp. 14-15;
text-fig. 7A-I; pl. 3, figs 1-5
Coniopteris hymenophylloides (Brongniart) Seward, emend. Harris; Gee, p. 159; pl. 1,
figs 7, 8
29
Fig. 14. A – Coniopteris cf. simplex (Lindley et Hutton) Harris, S82/11 KRAM-P; B–F – Coniopteris
cf. hymenophylloides (Brongniart) Seward, emend. Harris: B – S78/107 KRAM-P; C – ZPAL PL 3/9; D
– S78/40 KRAM-P; E – ZPAL PL 3/8; F – ZPAL PL 3/9. Scale bar 2.5 mm
1999
2002
2004
2004
Coniopteris hymenophylloides (Brongniart) Seward; van Konijnenburg-van Cittert &
Morgans, p. 53; text-fig. 23A, B; pl. 3, fig. 2
Coniopteris hymenophylloides (Brongniart) Seward; Wang, pl. 5, figs 4-6
Coniopteris hymenophylloides (Brongniart); Deng et al., p. 214; pl. 1, figs 1, 2
Coniopteris sp. B; Rees & Cleal, p. 22; pl. 8, fig. 2
Material. Several fragments of fertile frond impressions, together with small
fragments of sterile fronds. Some specimens show the presence of fertile frond in
penultimate and ultimate sections only. Specimens: ZPAL PL. 3/8 and 3/9 (fertile
fronds); S78/40, S78/104, S78/107 KRAM-P (sterile fronds).
30
Description.
Fertile fronds: ultimate fertile frond section alternating, elongated in outline,
with totally reduced frond blade. Axis 1 mm broad. Sori more or less opposite (Fig.
14F), reniform, 0.75 to 1.5 mm in diameter (Fig. 14E, F).
Sterile fronds: Penultimate sterile frond sections triangular in outline (Fig.
14D). Axis about 0.5 mm broad. Ultimate frond section rhomboid, with lobes
slightly rounded at apices. Lobes wider than axis, 3 mm long and 2.5 mm broad,
deeply incised: more than half distance from axis centre to a site where they would
terminate if not incised. Ultimate frond sections do not overlap. Nervation invisible.
Discussion.
Fertile fronds with totally reduced frond blade are characteristic of both Coniopteris hymenophylloides and C. simplex (see Harris, 1961), as is also the size of
sori. Attribution of our specimens to C. hymenophylloides is indicated by the occurrence (Fig. 14C, F) of both fertile (thus, probably belonging to) and sterile fronds
belonging to this species.
Sterile fronds showing such features as lobes shape in ultimate frond section,
their size, depth of incisions, and non-overlapping sections, are indicative of the
species C. hymenophylloides (see Halle, 1913a; Harris, 1961; Gee, 1989b; van
Konijnenburg-van Cittert & Morgans, 1999). Small sizes of our frond fragments do
not allow but tentative specific determination. They resemble also the species C.
simplex, however fronds of the latter species have narrower ultimate sections.
There is some similarity in outline of C. hymenophylloides fronds to that of C.
bella. However, the latter species shows distinctly rounded lobes in ultimate frond
section, and not roundish, as is the case with the former one (van Konijnenburg-van
Cittert & Morgans, 1999).
From C. burejensis fronds, the species C. hymenophylloides differs in having
deeply incised lobes in ultimate frond sections: these incisions run more than half
distance from the section’s axis centre to a theoretical site of section’s margin if not
incised. On the contrary, incisions in fronds of C. burejensis are smaller than half
this distance (Harris, 1961).
There is also a similarity of C. hymenophylloides fronds to C. murrayana. However, ultimate frond sections of the latter species very often overlap, a feature almost unknown in the former one (see van Konijnenburg-van Cittert & Morgans,
1999).
From C. lobata, fronds of C. hymenophylloides differ in outline of ultimate
frond sections: rhomboid in the latter species, and elongated in the former one.
Distribution. The species Coniopteris hymenophylloides was widely distributed in
Laurasia during Jurassic. It is also known from Jurassic of New Zealand (C.
hymenophylloides var. australis), Australia (New South Wales), West Antarctica
(Antarctic Peninsula) and India. The species is also recorded from Lower Cretaceous beds of China (Gee, 1989a; Jongmans & Dijkstra, 1959b).
31
Coniopteris cf. simplex (Lindley et Hutton) Harris, 1961
(Figs 11B, 14A)
1961
1999
2000
Coniopteris simplex (Lindley et Hutton); Harris, pp. 142-146; fig. 49; fig. 50a-g
Coniopteris simplex (Lindley et Hutton) Harris; van Konijnenburg-van Cittert &
Morgans, p. 53; text-fig. 13G, H; pl. 4, fig. 2
Coniopteris simplex (Lindley et Hutton) Harris; Barale et al., p. 685; fig. 9
Material. One imprint of sterile composite frond fragment, S82/11 KRAM-P.
Description. Frond fragment with three penultimate sections preserved, 7–16 mm
long and 4 mm broad. Sections branch off at 50° from axis which is 0.75 mm broad.
Axis of penultimate sections c. 0.5 mm broad. Ultimate frond sections rhomboid in
outline, 3 mm long and 2 mm broad; their lobes are roundish to elongate, very
narrow (maximum 0.75 mm broad).
Discussion. Small width of lobes makes our single specimen most similar to the
species Coniopteris simplex.
A possible occurrence of C. simplex in the Hope Bay material was suggested by
Harris (1961) who took into account the specimens determined by Halle (1913a) as
Sphenopteris leckenbyi. Gee (1989b), however, refrained from following his suggestion, as not all of the Halle’s collection was available to her for study. Our specimen is very similar to those presented by Harris (1961, fig. 49). A feature characteristic for C. simplex, as given by Harris and other authors (e.g., van Konijnenburgvan Cittert & Morgans, 1999), is a small width of lobes in ultimate frond section.
C. hymenophylloides has fronds most similar to C. simplex, however width of its
lobes is greater, as is also the case with other species belonging to the genus Coniopteris (see Harris, 1961; van Konijnenburg-van Cittert & Morgans, 1999).
Distribution. Europe: England (Yorkshire) – Jurassic (Aalenian) (van Konijnenburg-van Cittert & Morgans, 1999); Tunisia: Jurassic (Bathonian) (Barale et al.,
2000).
Family unknown (Osmundaceae ?)
Genus Cladophlebis Brongniart, 1849
Type species: Cladophlebis albertsii (Dunker) Brongniart
The genus Cladophlebis includes sterile fronds with characteristic shape and
nervation of pinnules. The pinnules have rounded or slightly sharpened apices,
with midvein passing along their middle part (Taylor & Taylor, 1993, fig. 16.38).
Such leaves may belong to ferns of the genera: Todites, Eboracia, Dicksonia, Klukia and Kylikipteris (Boureau, 1970).
Stratigraphic distribution of the genus Cladophlebis includes, first of all, Mesozoic, however Palaeozoic forms are also known: e.g., Permo-Carboniferous species C. royleri, C. tychtensis, C. yongwolensis, and Permian species C. mongolica,
C. nystroemi, and C. ozakii (Boureau, 1975; Jongmans & Dijkstra, 1959a).
32
Cladophlebis antarctica Nathorst, 1913
(Figs 15B, 16, 17B)
1913a
1913a
1934
1947
1971
1989a
1989b
2004
2004
Cladophlebis antarctica Nathorst; in Halle, pp. 14-15; pl. 1, figs 15-23
Cladophlebis antarctica Nathorst; in Halle (?), pl. 3, fig. 6
Cladophlebis antarctica Nathorst; Edwards, p. 94; text-fig. 4; pl. 4, fig. 5
Cladophlebis antarctica Nathorst; Frenguelli, fig. 5 (redrawn from Halle, 1913a)
Cladophlebis antarctica (Nathorst) Halle; Herbst, p. 268, pl. 1, fig. 10
Cladophlebis antarctica Halle; Gee, p. 41, fig. 2h
Cladophlebis antarctica Nathorst in Halle; Gee, pp. 166-167; pl. 2. fig. 15; pl. 3, fig. 24
Cladophlebis antarctica Halle; Ociepa, fig. 2
Cladophlebis antarctica Halle; Rees & Cleal, pp. 25-26; text-fig. 3c; pl. 6, fig. 2; pl. 7,
fig. 3
Material. Imprints of sterile frond fragments, consisting of penultimate frond
sections with pinnules. A larger fragment (Fig. 16) was preserved in a coarsegrained rock: its morphological features are invisible. A smaller fragment (Figs
15B, 17B) shows penultimate frond section with one complete and several fragmental pinnules preserved. Samples S78/113, S81/1a, b (positive and negative)
KRAM-P.
Description. Fragment of composite frond, 27 cm long and 21 cm broad, divided in
penultimate sections, with pinnules in ultimate sections. Axis 2 mm broad.
Fig. 15. A – Cladophlebis denticulata (Brongniart) Nathorst, emend. Harris, ZPAL PL 3/4; B –
Cladophlebis antarctica Nathorst, S78/113 KRAM-P; C – Cladophlebis grahamii Frenguelli, S82/13
KRAM-P. Scale bar 5 mm
33
Fig. 16. Cladophlebis antarctica Nathorst, S81/1b KRAM-P. Scale bar 2 cm
Penultimate frond sections 2–12 cm long and from 2.5 cm (at base) to 1 cm (at apex)
broad: they form an angle 60–80° with respect to rachis. These sections are almost
opposite, shifted with one another by 2–4 mm. Spaces between penultimate
sections amount to c. 5 mm.
Pinnules elongate, pointed at apices, with sitting base, not widened at base. Pinnules opposite or alternate, at an angle 60–90° with respect to axis. Their size diminish from base (18 mm) towards apex (5 mm), their width varies from 3 to 5 mm.
Length/width relation of pinnules at penultimate section base is always >3; this relation in higher situated pinnules is probably not much smaller (it is difficult to determine, as apices of such pinnules are not clearly visible). Pinnules slightly denticulated. Main nervation passes along mid-pinna, single lateral veins radiate from
the main one.
Discussion. Fronds of Cladophlebis antarctica resemble most those of C. denticulata. Fronds of these two species differ by width of axis: in our specimens it
34
measures barely 2 mm which is a diagnostic feature for C. antarctica (see Gee,
1989b), while in fronds of C. denticulata it reaches up to 7 mm (see Harris, 1961).
Penultimate frond sections in C. antartica are opposite or alternate, but alternate in
C. denticulata. Pinnules in C. antarctica are set at angle 60–90°, but in C.
denticulata – at angle 40–90° with respect to axis. Length/width ratio in pinnules of
C. antarctica exceeds 3, but in C. denticulata is smaller than 3. Pinnules in C.
antarctica are elongate, without a marked widening at base (Halle, 1913a; Gee,
1989b), while in C. denticulata they are triangular, with base widening apex-wise
(Herbst, 1971, pl. 1, fig. 9).
Fronds of C. antarctica are easily distinguishable from those of other species of
the genus Cladophlebis: axis in C. antarctica is narrow, while it is very wide in C.
grahami, that is also typical for C. denticulata (Frenguelli, 1947); pinnula apex in
C. antarctica is pointed, while it is rounded in C. oblonga (Halle, 1913a; Frenguelli, 1947; Gee, 1989b).
Rees and Cleal (2004) suggested that C. antarctica and C. denticulata might
represent morphotypes of the same taxon, i.e., C. denticulata. However, they have
refrained from redefinition of diagnosis of the latter due to poorly preserved material at their disposal. The present author disagrees with their suggestion, as both
species, C. antarctica and C. denticulata, are well separable, differing in important
features (see above).
The species Cladophlebis antarctica was first determined by Nathorst (in Halle,
1913a). Its diagnosis was first presented by Halle (op. cit.), however the author of
the species is Nathorst and not Halle: thus non “C. antarctica Halle” in Gee (1989a)
and Ociepa (2004). Conforming to the rules of Article 46.2 of The International Codex of Botanic Nomenclature (Greuter et al., 2000), the correct name of the species
would thus be: either Cladophlebis antarctica Nathorst, or Cladophlebis antarctica Nathorst, in Halle.
Distribution. Argentina: Río Autel, Llantenes Formation, Jurassic (Sinemurian)
and Tequetrén – Jurassic; New Zealand: Waikato and Matura – Jurassic; West
Antarctica: Hope Bay, Botany Bay and Orville Coast – Jurassic, and Mount
Goodman – Mesozoic (?), Alexander Island – Early Cretaceous (Halle, 1913a;
Jongmans & Dijkstra, 1967; Herbst, 1971; Gee, 1989a, b; Rees & Cleal, 2004).
Cladophlebis denticulata (Brongniart) Nathorst, emend. Harris, 1961
(Figs 15, 17F, G)
1876
1894
1905
1947
1947
1961
1971
Cladophlebis denticulata Brongniart; Nathorst, p. 19 (a short remark only)
Cladophlebis denticulata A Brongniart; Raciborski, p. 82; pl. 22, figs 3-4a
Cladophlebis denticulata (Brongniart) Nathorst non Fontaine; Ward, pp. 68-71; pl. 11,
figs 3-7 (non: 1-2!)
Cladophlebis grahami Frenguelli; Frenguelli, pp. 19-20, fig. 7 (redrawn from Halle,
1913a, pl. 2, fig. 8)
Cladophlebis denticulata (Brongniart) Fontaine; Frenguelli, pp. 17-19; fig. 4
Cladophlebis denticulata (Brongniart) Fontaine; emend. Harris, pp. 78-86, figs 25A-G,
26A, B
Cladophlebis denticulata (Brongniart) Fontaine; Herbst, p. 269; pl. 1, fig. 9
35
Fig. 17. A – Cladophlebis grahamii Frenguelli, S82/13 KRAM-P; B – Cladophlebis antarctica
Nathorst, S78/113 KRAM-P; C – Cladophlebis sp. B, S78/14 KRAM-P; D – Cladophlebis sp. A, S78/104
KRAM-P; E – Komlopteris indica (Feistmantel) Barbacka, emend. Rees et Cleal, S78/119 KRAM-P; F
– Cladophlebis denticulata (Brongniart) Nathorst, emend. Harris, S81/3b KRAM-P; G – Cladophlebis
denticulata (Brongniart) Nathorst, emend. Harris, ZPAL PL 3/4. Scale bar 5 mm
36
1980
1989b
2002
2004
Cladophlebis denticulata (Brongniart) Fontaine; Baldoni, p. 248; pl. 2, fig. D
Cladophlebis denticulata (Brongniart) Fontaine, emend. Harris; Gee, p. 168; pl. 2, fig.
14; pl. 3, fig. 25
Cladophlebis denticulata (Brongniart) Fontaine; Wang, pl. 1, fig. 3
Cladophlebis denticulata (Brongniart) Fontaine, emend. Harris; Rees & Cleal, pp. 26-28;
text-fig. 3D; pl. 6, fig. 4; pl. 7, figs 1-2
Material. Several fragments of penultimate sections of sterile fronds. Specimens:
S81/3a, b KRAM-P; ZPAL PL 3/4.
Description. Axis in penultimate frond part c. 0.8 mm broad. Pinnules 9–13 mm
long, 4–5 mm broad at base, their length/width ratio is <3. Pinnules situated
opposite to each other at an angle 50–70° with respect to axis. Pinnules sitting,
triangular in outline, pointed at apices; their margin is delicately folded, pinnule
base is asymmetric, slightly widening apex-wise. Main vein passing along pinna
centre from its base to apex; its lateral veins divide dichotomously.
Discussion. Cladophlebis denticulata is best characterized by: width of axis, angle
of penultimate frond section with respect to axis, pinnula shape and its length/width
ratio. These features characterize our specimens from Mt Flora attributed to
Cladophlebis denticulata. Fronds of C. denticulata are most similar to those of C.
antarctica, however they differ by pinnules’ shapes and sizes. Pinnules of C.
denticulata form an angle of 50–70°, while those of C. antarctica – of 60–90° with
respect to axis. Length/width ratio in our specimens is <3 in C. denticulata, while in
C. antarctica it exceeds 3. Pinnules of C. denticulata are triangular (Halle, 1913a;
Gee, 1989b), while those of C. antarctica are elongate. A feature typical for C.
denticulata, which is absent in C. antarctica, is the presence of apex-wise widening
at their base (Harris, 1961; Herbst, 1971, pl. 1, fig. 9): this feature is well
distinguishable in our specimens from Mt Flora (Fig. 15A).
The species of the genus Cladophlebis differ from one another by sizes and
shapes of their pinnules. Their apex in C. denticulata may be either blunt or pointed
(the latter feature is observed in our specimens), while that of C. oblonga is rounded
(Halle, 1913a; Frenguelli, 1947; Gee, 1989b); pinnules of C. grahami are slightly
narrowing at both sides of their bases (Gee, 1989b); pinnule bases in C. harrisi are
fused to form a winglet (van Cittert, 1966).
Rees and Cleal (2004) suggested that C. antarctica and C. denticulata may represent the same taxon (see discussion in C. antarctica).
Harris (1961) combined sterile leaves of Cladophlebis denticulata with fertile
leaves (with sporangia) of Todites denticulatus (Brongniart) Krasser in one and the
same species; he wrote, however (op. cit., p. 82), that when one has at his/her disposal only sterile leaves, the name Cladophlebis denticulata should rather be applied.
There are some inconsequences with synonymy of Cladophlebis denticulata. It
was Brongniart who, in 1828, had first described this species as Pecopteris denticulata (cf. Ward, 1905; Harris, 1961; Herbst, 1971). It is unclear, however, who first
changed its generic name to Cladophlebis. Harris (1961), followed by other authors
(e.g., Herbst, 1971; Gee, 1989b), believed it was Fontaine. In fact, Ward (1905,
37
footnote in p. 69) wrote that Fontaine was the first to describe a new species named
Cladophlebis denticulata, while such specific name, Cladophlebis denticulata
Brongniart, was already used by Nathorst (1876, p. 19). Nathorst, as the author of
this combination, is also mentioned by Jongmans and Dijkstra (1959a). Thus, a correct name for the discussed species would be: Cladophlebis denticulata (Brongniart) Nathorst, emend. Harris, 1961.
Distribution. Laurasia: numerous sites, Triassic to Cretaceous in age; New Zealand and West Antarctica – Jurassic; Africa: C. denticulata Brongniart form
atherstonei (Tate) Seward – Jurassic; Australia, India – C. denticulata Brongniart
var. australis Morris (this variety is sometimes treated as a separate species – see
Jongmans & Dijkstra, 1959a, 1967; Boureau, 1975).
Cladophlebis grahamii Frenguelli, 1947
(Figs 15C, 17A)
1913a
1913a
1947
1971
1989b
Cladophlebis (Coniopteris?) arguta (Lindley et Hutton); Halle, pp. 15-16; pl. 2, figs 1, 2,
5
Cladophlebis antarctica Nathorst; in Halle (?), pl. 1, fig. 24
Cladophlebis grahami Frenguelli; Frenguelli, pp. 50-52; pl. 3, figs 1-7; pl. 4, figs 1-3
Cladophlebis grahami Frenguelli; Herbst, pp. 269-270; pl. 1, fig. 2
Todites grahamii (Frenguelli); Gee, pp. 163-165; text-fig. 2; pl. 2, fig. 19; pl. 3, fig. 22
Material. One sterile frond fragment with penultimate and ultimate sections.
Specimen S82/13 KRAM-P.
Description. Frond bipinnate. Axis only partially visible, 1.5 mm broad. Preserved
fragments of penultimate pinnae are 4.5 cm long and 1–11.5 cm broad. They grow
at an angle of 50–60° with respect to second-order axis which is c. 0.8 mm broad.
Second-order axis with verrucae (Fig. 15C). Particular pinnules are 5.5–6.5 mm
long and c. 2.5 mm wide, their length/width ratio is 2.2–2.6. Pinnule outline ovate,
pointed at apex, cardiform, slightly narrowing at base. Pinnules delicately corrugated at margins, forming an angle of 66–70° with respect to axis. Pinnules form
pairs nearly opposite to each other, with the exception of base of the penultimate
section where a single pinnule occurs (Fig. 17A). Midvein runs along middle part of
pinnules, poorly visible single lateral veins propagate from both sides of it.
Discussion. Attribution of the analysed frond fragment to the species Cladophlebis
grahamii is based on the following features (see Frenguelli, 1947; Gee, 1989b):
pinnule pairs in penultimate frond section are opposite to each other; a single
pinnule appears at the beginning of each penultimate frond section; pinnule
length/width ratios amount to c. 2.5; pinnule basis cardiform, its margin delicately
corrugated. These features are absent from other species of the genus Cladophlebis.
Other species of this genus differ in size and shape of pinnules: the length/width
ratio in C. antarctica exceeds 2.5; pinnules of C. denticulata show base parts widened apex-wise; bases of C. harrisii fuse forming a winglet (Halle, 1913a; Harris,
1961; van Cittert, 1966; Gee, 1989b).
38
Halle (1913a) determined Cladophlebis-type fertile fronds from Hope Bay as
Todites williamsonii, Gee (1989b) disagreed with his conclusion and included his
specimens to T. grahamii, as a new combination.
Sterile fronds of T. williamsonii differ from those of C. grahamii in a well distinguishable feature: the pinnules of T. williamsoni are not narrowing at base (Harris,
1961), while cardiform pinnules of C. grahami are slightly narrowing at base (Gee,
1989b). Cardiform base of pinnules is well recognizable in Halle’s (1913a) specimens determined as Cladophlebis (Coniopteris?) arguta. The latter specimens
were included by Frenguelli (1947) to the species C. grahamii, by Gee (1989b) to
Todites grahamii, and by Rees and Cleal (2004) – to T. williamsonii. It should be remarked that the Cladophlebis-type fertile fronds from Hope Bay determined by
Halle (1913a) as Todites williamsonii were included by Harris (1961) to synonimic
list of this species, while those determined by Halle (1913a) as Cladophlebis (Coniopteris?) arguta were not.
The specific shape of pinnule base was included in the diagnosis and illustrated
as Todites grahamii by Gee (1989b, text-fig. 2). The present author considers this
feature as diagnostic in distinguishing sterile fronds of Todites williamsonii (pinnule base not narrowing) from Cladophlebis grahamii (cardiform pinnule base
slightly narrowing).
Frenguelli (1947) incorrectly included to C. grahamii the fronds determined by
Halle (1913a) as C. denticulata. To the species C. grahamii Frenguelli correctly included Halle’s specimens determined as C. (Coniopteris?) arguta. However, his
reproduction of Halle’s specimen cited as “pl. 2, fig. 2” and described as C.
(Coniopteris) arguta, in Frenguelli’s paper depicts C. oblonga which in Halle’s
monograph was illustrated as “pl. 2, fig. 6”. C. oblonga is a well separable species
with characteristically rounded pinnule apex (Halle, 1913a; Frenguelli, 1947;
Herbst, 1971). The whole misunderstanding was clarified already by Herbst
(1971), and followed by Gee (1989b). However, Gee incorrectly included C.
grahamii of Frenguelli (1947, pl. 3, figs 1-7 and pl. 4, figs 1-3) in synonimies of
both Todites grahamii and Cladophlebis denticulata.
It should also be pointed out that correct specific name of Todites grahamii is:
“Todites grahamii (Frenguelli) Gee” and not “Todites grahamii (Brongniart) Seward”, because Frenguelli described only Cladophlebis grahamii, while a combination of Todites grahamii was created by Gee (1989b).
Distribution. Argentina: Paso Flores – Triassic (Rhaetian), Piedra Pintado –
Jurassic (Liassic), Patagonia – Jurassic; West Antarctica (Hope Bay) – Jurassic
(Jongmans & Dijkstra, 1959a; Gee, 1989b).
Cladophlebis sp. A
(Fig. 17D)
Material. Fragment of axis with two pinnules damaged at apex, S78/40 KRAM-P.
Description. Pinnules 5–6 mm broad and 10 mm long at preserved fragment.
Pinnules with rounded base attached to axis at midrib. Pinnule margin slightly
39
wavy. Pinnule midrib runs along its middle, singly furcated lateral veins propagate
from it.
Discussion. Pinnule shape and nervation of our specimen are typical for the genus
Cladophlebis (see Frenguelli, 1947; Harris, 1962; Gee, 1989b). Forms with single,
furcating lateral veins in pinnules belong to group A of Frenguelli (1947), which
includes Cladophlebis arguta, C. insignis, C. acuta, C. denticulata, C. antarctica,
C. toroyensis, C. grahamii, C. nebbensis, C. oblonga, C. atherstoni (presently most
frequently treated as a form of C. denticulata) and C. patagonica.
Our specimen, by its shape and size, resembles most C. atherstoni as figured by
Frenguelli (1947, fig. 12). Such features as wavy pinnule margin and rounded base
resemble those of C. grahami (see Gee, 1989b). Fragmentary preservation of our
specimen precludes its specific determination.
Cladophlebis sp. B
(Fig. 17C)
Material. Imprint of a single pinnule with midrib visible. S78/14 & 78/16 (positive and
negative) KRAM-P.
Description. Pinnule 13 mm long and 6 mm broad, its length/width ratio is 2.17.
Pinnule ovate, with blunt apex, with delicately denticulated margin, with rounded
base. It was probably attached to axis at midrib entry. Midrib passing mid-pinnule
from base to apex.
Discussion. Rounded pinnule base, length/width ratio close to 2.5, and ovate shape,
may suggest its attribution to the species Cladophlebis grahamii. Fragmentary
preservation of the specimen precludes its specific determination.
Pteridophylla
This is an artificial systematic group created by A. G. Nathorst, which includes
form-genera with fronds resembling those of ferns, but devoid of reproductive organs: thus, it is not possible to decide whether they belong to ferns or to pteridosperms (Boureau, 1975). Taxonomic units within this group are based on morphological structure of fronds.
Form-genus Sphenopteris Sternberg, 1825
Type species: Sphenopteris elegans (Brongniart) Sternberg
This is a form-genus created for fronds with characteristic ultimate sections,
usually sphenoidal, with pinnate nervation (Taylor & Taylor, 1993, fig. 16.34).
Stratigraphic age range of the genus Sphenopteris includes Devonian through Cretaceous, with maximum occurrence during Carboniferous (Boureau, 1975).
40
Sphenopteris antarctica Halle, 1913
(Figs 18A, 19A)
1913a
1989b
Sphenopteris antarctica sp. nov.; Halle, pp. 30-31; fig. 7d; pl. 3, figs 19, 21
Sphenopteris antarctica Halle; Gee, p. 172; pl. 3, fig. 20
Material. Several fragments of frond imprints. S78/18 and 78/21 (positive and negative),
S78/136, 78/173, 82/8 KRAM-P.
Description. Midrib of penultimate frond section faintly winged (Fig. 16A), with
longitudinal ridges. Midrib width 0.8–1 mm, winglet inclusively. Winglet c. 0.25
mm broad at both sides of midrib. Pinnules elongate in outline, at 55–60° angle with
respect to midrib, alternating, 6–8 mm long and 2–3 mm broad. Pinnule deeply
incised, lobes triangular, with slighly pointed apex, c. 2 mm long and c. 1 mm
broad; 6–7 lobes per each pinnule. Nervation preserved: a single vein detaches
from midrib at penultimate section and enters each section where it branches off
and enters each lobe (Fig. 18A).
Discussion. Sphenopteris antarctica was created by Halle (1913a) based on
material from Hope Bay. Fronds of this species resemble most those of S. pecten,
however midrib of penultimate section in S. antarctica is only 0.8–1 mm broad,
while that of S. pecten is 2 mm broad. Midrib of S. antarctica has a winglet at both
sides; it is c. 0.25 mm broad, while that of S. pecten is 0.75 mm broad. Lobes in S.
antarctica are smaller than those of S. pecten: they are maximum 2 mm long, while
lobes of the former species reach 2–5 mm in length. Lobes are triangular, slightly
pointed at apex in S. antarctica, while in S. pecten they are ovate with slightly
rounded apices.
Fig. 18. A – Sphenopteris antarctica Halle, S78/21 KRAM-P; B – Sphenopteris sp. A, S78/40
KRAM-P; C – Sphenopteris sp. C, S78/5a KRAM-P; D – Sphenopteris sp. B, S78/14 KRAM-P. Scale
bar 2.5 mm
41
Fig. 19. A – Sphenopteris antarctica Halle, S78/21 KRAM-P; B – Pachypteris cf. indica (Oldham et
Morris) Bose et Roy, emend. Bose et Banerji, S78/118 KRAM-P. Scale bar 2.5 mm
Fronds of S. antarctica resemble to some extent those of Archangelskya furcata
(see Rees & Cleal, 1993, text-fig. 2C), however lobes are triangular in S. antarctica,
while in A. furcata they are elongate, narrowing both apexward and baseward.
Fronds of S. antarctica morphologically resemble also those of Coniopteris
murrayana, however lobes are triangular and pointed at apex in S. antarctica, while
they are obovate and rounded at apex in C. murrayana.
It is of interest to note that S. antarctica has not been distinguished by Rees and
Cleal (2004) in their collections from Hope Bay and Botany Bay.
Distribution. West Antarctica, Antarctic Peninsula: Hope Bay – Jurassic (Halle,
1913a; Gee, 1989b).
Sphenopteris pecten Halle emend. Gee, emend. Rees et Cleal, 2004
(Fig. 20)
1913a
1989b
2004
Sphenopteris pecten sp. nov.; Halle, p. 35; pl. 4, figs 21, 21a
Sphenopteris pecten Halle; Gee, pp. 176-177; pl. 4, fig. 32
Sphenopteris pecten Halle; Rees & Cleal, pp. 31-32; pl. 8, fig. 4
Material. One specimen preserved as imprint of frond fragment consisting of
penultimate and ultimate frond sections. S78/137 KRAM-P.
Description. Midrib of penultimate frond section flattened, widely winged (Fig.
20), c. 2 mm wide including winglets; each winglet c. 0.75 mm broad. Ultimate
42
Fig. 20. Sphenopteris pecten Halle, emend. Gee, emend. Rees et Cleal, S78/137 KRAM-P, scale bar
5 mm (phot. M. Dziewiñski)
frond sections oblong in outline, 10–15 mm long and 2–6 mm broad. Lobes in these
sections are ovate in outline, narrowing apexward, rounded at apex. Lobes are
3–5 mm long and c. 1.5 mm broad, situated at an angle c. 45° with respect to axis,
nearly opposite to each other. Nervation visible: broad midrib passes along main
frond axis, branching off to enter pinnules of ultimate frond section (Fig. 20).
Discussion. The species Sphenopteris pecten distinguishes itself form other
species of the genus Sphenopteris by having widely winged midrib in penultimate
frond section. It is most similar to S. antarctica, however midrib in S. pecten is
much broader (c. 2 mm) than that of S. antarctica (c. 0.8–1 mm). Midrib in S. pecten
is wider winged, with wings c. 0.75 mm broad, as compared with S. antarctica
43
Fig. 21. A – Sphenopteris sp. C, S78/5a KRAM-P; B – Sphenopteris sp. A, S78/40 KRAM-P; C –
Sphenopteris sp. B, S78/57 KRAM-P. Scale bar 2.5 mm
where they are only c. 0.25 mm broad. Lobes of ultimate sections in S. pecten are c.
2–5 mm long, being larger than those of S. antarctica where they are maximum 2
mm long. These two species do also differ in shapes of lobes: they are ovate, with
slightlty rounded apex in S. pecten, but triangular, with slightly pointed apex in S.
antarctica.
Fronds of S. pecten somewhat resemble those of Archangelskya furcata (see
Rees & Cleal, 1993, text-fig. 2D), however midrib of S. pecten is strongly winged,
in contrast to that of A. furcata which is not winged.
Distribution. Australia, Queensland – Triassic (Jongmans & Dijkstra, 1965); West
Antarctica, Antarctic Peninsula: Hope Bay and Botany Bay – Jurassic (Halle,
1913a; Gee, 1989b; Rees & Cleal, 2004).
Sphenopteris sp. A
(Figs 19B, 21B)
Material. One fragment of ultimate frond section, S78/40 KRAM-P.
Description. Ultimate frond section with fan-like lobes (Fig. 19B). Lobes c. 2 mm
long and c. 0.6 mm broad.
Discussion. Our specimen resembles fronds of the genus Sphenopteris with
fan-wise ultimate frond sections, such as: S. herbstii (see Baldoni, 1980, pl. 2E), S.
metzgerioides (see Bose & Banerji, 1984, text-fig. 12A-B; Harris, 1961, text-fig.
71D), and S. bagualensis (see Menéndez, 1956, pl. 1, figs 1, 4, 5); the latter species
was included by Rees and Cleal (2004) to synonimic list of Pachypteris indica.
44
Our frond fragment resembles terminal frond parts of Ruffordia goepperti
which, morphologically, is a very variable species. Terminal lobes of its fronds
may be either broad (see Watson, 1969, pl. 4, figs 3, 4, 7) or narrow (see Watson,
1969, pl. 4, figs 1, 5). It is difficult to distinguish our species from the latter one.
The specimen of Sphenopteris sp. A differs from that determined as Sphenopteris sp. B by length and width of lobes, and from another one determined as
Sphenopteris sp. C in having well developed lobes.
Fragmentary preservation of our specimen preclude its closer determination.
Sphenopteris sp. B
(Figs 19D, 21C)
Material. Imprint of fragment of ultimate frond section. S78/57 KRAM-P.
Description. Frond section with deeply incised lobes descending midrib-ward
(Fig. 19D). Lobes 3–5 mm long and 1–1.2 mm broad. Single nerves enter the lobes.
Frond blade seems to be thick, with lateral nerves poorly marked on it.
Discussion. Our ultimate frond section resembles those of Sphenopteris herbstii, S.
metzgerioides and S. bagualensis (see discussion in Sphenopteris sp. A). It also
resembles that of S. nordenskjoeldii (see Halle, 1913a; Rees & Cleal, 2004);
however, fronds of S. nordenskjoeldii seem to be very thin, that causes clear
appearance of its nerves, while our frond blade seems to be rather thick that causes
its veins to appear faint.
From Sphenopteris sp. A, the investigated specimen differs in length and width
of lobes, which descend towards midrib, and from Sphenopteris sp. C – in well developed lobes.
Fragmentary preservation of our specimen precludes its closer determination.
Sphenopteris sp. C
(Fig 19C, 21A)
Material. Imprint of penultimate frond section, S78/5a KRAM-P.
Description. Midrib 1.2 mm broad. From it, laterally propagate incompletely
preserved ultimate frond sections divided in 2–3 poorly developed lobes. Preserved
lobe fragments are c. 2 mm long and c. 0.6 mm broad.
Discussion. Our specimen resembles analogous frond sections of Sphenopteris
herbstii, S. metzgerioides, and S. bagualensis (see discussion in Sphenopteris A). It
also resembles fronds of Pachypteris indica, the latter often showing poorly
developed lobes (see Bose & Banerji, 1984, text-fig. 20; Bose & Roy, 1967,
text-fig. 1B, C).
From forms determined as Sphenopteris sp. A and Sphenopteris sp. B, our
specimen differs in having poorly developed lobes.
Fragmentary preservation of our specimen precludes its closer determination.
45
Division Pteridospermophyta
Order Caytoniales
Family Caytoniaceae
This is the family of Mesozoic seed ferns. Its representatives have leathery,
palm-shaped leaves, divided in 3–6 segments with meshy nervation (see Taylor &
Taylor, 1993). Male microsporophyll – Caytonanthus, and female megasporophyll
– Caytonia, are known from widely scattered locations. From Gondwanaland,
Caytonanthus was first described from Botany Bay (Antarctic Peninsula, West
Antarctica) by Rees (1993a).
Genus Sagenopteris Presl, emend. Harris, emend. Rees, 1993
Type species: Sagenopteris acuminata Presl (singled out as such by Cleal & Rees, 2003)
About 70 species are known from Mesozoic strata, most commonly from Jurassic and Cretaceous (Jongmans & Dijkstra, 1964).
Sagenopteris nilssoniana (Brongniart) Ward 1900
(Figs 22–24)
1867
1900
1913a
1932b
1956
1989b
1993a
2000b
2004
Sagenopteris rhoifolia Presl; Schenk, pl. 12, figs 1-6; pl. 13, figs 4-10
Sagenopteris nilssoniana (Brongniart); Ward, p. 352; pl. 56, fig. 1; pl. 67, fig. 2
Sagenopteris paucifolia (Phillips) Ward; Halle, pp. 8-9; pl. 1, figs 1-5
Sagenopteris nilssoniana (Brongniart) Ward; Harris, pp. 5-10; text-fig. 1, 2A-F
Sagenopteris nilssoniana (Brongniart); Menéndez, pp. 329-333; fig. 4; pl. 5
Sagenopteris paucifolia (Phillips) Ward; Gee, pp. 177-178; pl. 4, fig. 33
Sagenopteris nilssoniana (Brongniart) Ward; Rees, pp. 35-36; pl. 1, figs 1-7; pl. 2, figs
1-11; pl. 3, figs 1, 2
Sagenopteris nilssoniana (Brongniart) Ward; Cantrill, pp. 189-190; fig. 2f, i, k
Sagenopteris nilssoniana (Brongniart) Ward; Rees & Cleal, pp. 32-36, pl. 9, figs 1-5
Material. Imprints of fragments of single leaflets of palmate leaves. S78/22, 81-1
KRAM-P; ZPAL Pl. 3/4, 3/5, 3/12, 3/18, 3/20.
Description. Leaflets obovate in outline, narrowing baseward, 9–14 mm broad;
preserved fragments are 20–59 mm long. Two types of leaflets occur: (i) with
entire margin; (ii) with deeply incised, irregular lobes 2–4 mm broad and 7–13 mm
long; incisions do not reach midvein.
Main vein passes along mid-leaflet; anastomosing lateral nerves radiate from it
(Fig. 22); there are c. 24 lateral veins per 1 cm.
Discussion. Shape, size, nervation type and, first of all, division of a part of leaflets
in irregular lobes, indicate attribution of our specimens to Sagenopteris nilssoniana.
Halle (1913a) determined Sagenopteris species from Hope Bay as S. paucifolia.
He stated that this species has leaflets similar to S. nilssoniana, however leaflets of
the latter are broader and lobate. Gee (1989b) retained his S. paucifolia taxon, and
pointed out that the feature important to distinguish leaflets of S. paucifolia from S.
nilssoniana is the dimension of nerve meshes. According to Harris (1932b, 1964),
leaves of these two species differ in cuticle features.
46
Fig. 22. Sagenopteris nilssoniana (Brongniart) Ward. A – ZPAL PL 3/18 (phot M. Dziewiñski); B –
S82/1 KRAM-P; C – ZPAL PL 3/4. Scale bar 5 mm
Rees (1993a), who had at his disposal a large collection of specimens of the genus Sagenopteris from Hope Bay and Botany Bay, wrote that a part of these showed
leaflets with entire margins, while the rest was lobate. These leaflets could be continuously sequenced starting with those with entire margin, through those with
shallow-incised, to those with deeply incised lobes. This was the reason for him to
include them all to a single species Sagenopteris nilssoniana.
S. mclearni has also lobed leaflets, however incisions between the lobes are
there very regular, contrary to those of S. nilssoniana which are irregular (see Rees,
1993a). S. undulata has leaflets very similar to those of S. nilssoniana. According
to Rees (1993a), the former may be a synonym of the latter.
47
Fig. 23. Sagenopteris nilssoniana (Brongniart) Ward. A – S78/22 KRAM-P; B – ZPAL PL 3/12.
Scale bar 5 mm
Distribution. Japan – Triassic; Europe, Greenland, Asia, South America and West
Antarctica – Jurassic (Jongmans & Dijkstra, 1964; Rees, 1993a; Cantrill, 2000b).
Order Corystospermales
Family Corystospermaceae
This is the family of Mesozoic seed ferns, known mainly from reproductive organs: male Pteruchus and female Umkomasia, occurring mainly in Gondwanaland,
and from associated leaves of the genera Dicroidium and Pachypteris (Taylor &
Taylor, 1993).
Genus Pachypteris Brongniart, emend. Harris, 1964
Type species: Pachypteris lanceolata Brongniart.
Pinnate leaves of the genus Pachypteris are known from Jurassic through Cretaceous at numerous sites on the whole Earth (Taylor & Taylor, 1993).
Pachypteris cf. indica (Oldham et Morris)
Bose et Roy, emend. Bose et Banerji, 1984
(Fig. 19B)
1913a
1913a
Sphenopteris (Ruffordia?) goepperti Dunker; Halle, pp. 25-26; pl. 3, fig. 9
Pachypteris dalmatica. F. von Kern ?; Halle, pl. 4, fig. 35
48
Fig. 24. Sagenopteris nilssoniana (Brongniart) Ward. A, B – various leaf fragments from same
rock sample, ZPAL PL 3/20, scale bar 5 mm
1956
1967
1984
1989b
2000a
2003
2004
Sphenopteris bagualensis sp. nov.; Menéndez, pp. 319-323; fig. 1 (stomatal apparatus);
pl. 1, figs 1-6 (1-5 – macerated leaf; 6 – cross-section of terminal leaf section); pl. 2,
figs 1, 2 (stomatal apparatus)
Pachypteris indica (Oldham et Morris); Bose & Roy, pp. 2-6; text-figs 1-4, 5A; pl. 1,
figs 3-5, 7; pl. 2; pl. 3, figs 1-16, 19-20 (cuticle)
Pachypteris indica (Oldham et Morris) Bose et Roy; Bose & Banerji, pp. 29-37; text-figs
17-20; pl. 8; pl. 9, figs 1, 5, 7-9; pl. 10, figs 5, 6 (cuticle); pl. 11, figs 8, 9 (cuticle)
Sphenopteris bagualensis Menéndez; Gee, p. 173; pl. 4, fig. 30
Pachypteris indica (Oldham et Morris) Bose et Roy; Cantrill, pp. 177-178; pl. 8, figs
1-3, 8
Pachypteris indica (Bose et Roy) Bose et Banerji; Prakash, p. 67; pl. 1, fig. 3
Pachypteris indica (Oldham et Morris) Bose et Roy; Rees & Cleal, pp. 64-66; pl. 18
Material. Impression of leaf fragment with ultimate sections preserved. S78/118
KRAM-P.
Description. Ultimate leaf sections are 10 mm long and 1.5 mm broad, lanceolate,
with entire margin, slightly narrowing towards pointed apex and towards base.
Sections radiate at angles 30° with respect to midrib. Nervation not visible.
Discussion. Our specimen resembles those from Hope Bay described by Halle
(1913a) as Sphenopteris (Ruffordia?) goepperti, revised by Gee (1989b) as Sphenopteris bagualensis. Rees and Cleal (2004), based on cuticle structure of their
specimens, attributed the latter species to Pachypteris indica. In our specimen from
Mt Flora, cuticle was not preserved; however, size and shape of specimen’s
terminal leaf sections, indicates a close similarity to the species Pachypteris indica.
Lack of preserved cuticle in our specimen precludes its closer specific determination.
Rees and Cleal (2004) consider Sphenopteris bagualensis as a synonym of
Pachypteris indica. Leaves of the latter species are similar to those of P. lanceolata,
49
Fig. 25. Pachypteris sp., S78/114 KRAM-P. Scale bar 2.5 mm
however their terminal sections, and cuticle character are different (Bose & Roy,
1967).
According to Bose and Roy (1967), leaves of Pachypteris indica are very similar to those of P. holdeni: these two species are distinguishable from one another by
their cuticle features only. Bose and Banerji (1984) treat P. holdeni as a synonym of
P. indica.
Leaves of Pachypteris indica resemble also those of P. specifica; however, the
latter species has wider and longer leaves in ultimate section with respect to the
former one.
Distribution. West Antarctica (Antarctic Peninsula): Hope Bay and Botany Bay –
Jurassic; Snow Island – Cretaceous (Aptian); Argentina: Bajo de los Baguales –
Jurassic; India – Cretaceous; Australia – Mesozoic (Menéndez, 1956; Cantrill,
2000a; Prakash, 2003; Rees & Cleal, 2004).
Pachypteris sp.
(Fig. 25)
Material. One fragment of bipinnate frond impression. S78/144 KRAM-P.
Description. Frond fragment elongated in outline. Midrib c. 0.5 mm broad.
Penultimate frond sections elongated in outline, narrowing apexward, 5 mm long
and 2 mm broad, situated at an angle of 60° with respect to midrib, alternating.
Pinnules of ultimate section are nearly obovate in outline (Fig. 25), 1.5 mm long
and 0.5 mm broad. Nine pinnules at penultimate frond section. Nervation not
visible.
Discussion. The above features are consistent with those of the genus Pachypteris
(bipinnate frond – see Harris, 1964). Our specimen is very similar to P. lanceolata
as determined by Harris (1964, text-fig. 56C). Both species are of small size, the
species illustrated by Harris fits morphological variation of P. lanceolata, while
our specimen is even smaller, its morphological variation being also smaller than
that established for the species. Therefore, our specimen was determined only at
generic level.
50
Order: unknown
Family: unknown
Genus: Archangelskya Herbst, emend. Rees et Cleal, 1993
Type species: Archangelskya protoloxsoma (Kurtz) Herbst
This genus, a representative of seed ferns, was created by Herbst (1964), its diagnosis was supplemented by Rees and Cleal (1993). Its systematic position is unclear, as both reproductive organs and details of epiderm structure are unknown.
Attribution to the seed ferns is based on its bipinnate frond blade, and thick
(leather-like) structure (see Gee, 1989b; Rees & Cleal, 1993).
Archangelskya furcata (Halle) Herbst, emend. Rees et Cleal, 1993
(Fig. 26)
1913a
1913a
1913a
1913a
1913a
1968
1989b
1989b
1989b
1989b
1993
2004
Scleropteris furcata sp. nov.; Halle, pp. 37-38; text-fig. 9; pl. 4, figs 3, 10, 11, 13-18, 19?
Scleropteris crassa sp. nov.; Halle, pp. 36-37; pl. 3, fig. 14; pl. 4, figs 4-9, 12?, 22?
Sphenopteris anderssoni sp. nov.; Halle, pp. 33-35; text-fig. 8; pl. 3, fig. 10?; pl. 4, figs
1, 2
Sphenopteris fittoni Seward; Halle, pp. 28-30; text-fig. 7a-c; pl. 3, figs 15-18, 22, 25
Sphenopteris nauckhoffiana (Heer); Halle, pp. 26-28; text-fig. 6; pl. 3, fig. 26
Archangelskya furcata (Halle) Herbst; Herbst & Antozegui, p. 187; fig 1E, F
Archangelskya furcata (Halle) Herbst; Gee, pp. 181-182; pl. 4, figs 38-40
Pachypteris crassa (Halle) Townrow; Gee, pp. 178-179; pl. 4, figs 34, 35
Sphenopteris anderssoni Halle, emend. Zeba-Bano; Gee. pp. 171-172; pl. 3, figs 26-28
Sphenopteris hoppetsvikensis sp. nov.; Gee, pp. 174-175; text-fig. 3
Archangelskya furcata (Halle) Herbst; emend. Rees et Cleal, p. 98, text-figs 2-6
Archangelskya furcata (Halle) Herbst; emend. Rees et Cleal, pp. 70-72; pl. 19, fig. 1; pl.
20, figs 1-4
Material. 27 fragments of frond impressions, 22 out of which in fine-grained rock
that helped preserve some details of its morphology; 5 specimens in coarsergrained rock, with less clear morphological features preserved. Penultimate and
ultimate frond sections preserved. Specimens: S78/6a, b (positive and negative), S78/36,
78/39, 78/51, 78/62, 78/66, 78/78, 78/80, 78/85, 78/97, 78/113, 78/135, 78/144, 78/153, 78/160, 78/163, 78/166, S81/25-28,
81/30, S82/3-4 (positive and negative), S82/9 KRAM-P; ZPAL Pl. 3/3, 3/13.
Description. Fronds elongated in outline. Midrib of penultimate section 0.75–1.5
mm broad. Ultimate frond sections elongate-lanceolate (Fig. 26A) narrowing
apexward, with the exception of sections which were probably located at frond
terminus (Fig. 26B, C), and which are clearly triangular in outline. Ultimate frond
sections 7–18 mm long and 3.5–6 mm broad, except for those which were situated
at frond terminus, and which are c. 10 mm broad. In basal part of ultimate frond
section, first lobe facing frond terminus is larger than that facing frond base (Fig.
26D). Lobes 2–4.5 m long and 1–2 mm broad. Larger lobes, probably derived from
ultimate frond section, are c. 5 mm long and 1.5–2 mm broad. Lobes oblong,
narrowing apexward and baseward; their margin is irregularly incised.
Fragments of fronds of Archangelskya furcata represent two types. The first one
includes fragments of ultimate frond sections, originally probably located at frond
51
Fig. 26. Archangelskya furcata Herbst, emend. Rees et Cleal. A – ZPAL PL 3/3 (phot. M.
Dziewiñski); B – S78/51 KRAM-P; C – S78/66 KRAM-P; D – S78/51 KRAM-P. Scale bar 5 mm
52
terminus, which are visibly triangular in outline. They are larger than those of the
second type and show well incised lobe margins (Fig. 26B, C). The second type includes oblique-lanceolate pinnae of ultimate frond sections, with poorly incised
margins (Fig. 26A, D).
Discussion. Attribution of our specimens to Archangelskya furcata is based on
their shape and size, as well as variably incised lobe margins (see Rees & Cleal,
1993). This is a polymorph species, that caused attribution of its fronds to different
species and genera (see the synonymy). A large frond fragment elaborated by Rees
and Cleal (1993, text-figs 2A, 3A) allowed to include them all to the same species
Archangelskya furcata.
Fronds of A. furcata have fainter dentated lobe margins than those of A. protoloxsoma (see Rees & Cleal, 1993). They show some similarity to those of
Sphenopteris antarctica, however lobes of ultimate frond sections of the latter are
triangular, while those of A. furcata are elongated.
Distribution. West Antarctica (Antarctic Peninsula): Hope Bay and Botany Bay –
Jurassic (Rees & Cleal, 1993); Argentina: Taquetrén, Chubut Province – Jura
(Herbst & Anzotegui, 1968).
Genus Komlopteris Barbacka, 1994
Type species: Komlopteris nordenskjoeldii (Nathorst) Barbacka
Komlopteris is a new genus separated by Barbacka (1994) from the genus
Pachypteris, based on morphological features and cuticle structure. Four species of
this genus are known: K. indica (Feistmantel) Barbacka, K. nordenskjoeldii (Nathorst) Barbacka, K. rotundata (Nathorst) Barbacka and K. speciosa (Ettinghausen) Cleal et Rees. They occurred in Jurassic and Early Cretaceous (Barbacka,
1994; Cleal & Rees, 2003). Investigations of female reproductive organs are in
progress; probably this genus belongs to a new, so far unknown group of seed ferns
(Barbacka, 2002).
Komlopteris indica (Feistmantel) Barbacka, emend. Rees et Cleal, 2004
(Figs 17E, 27)
1877a
1913a
1913a
1979
1986
1989b
1989b
1994
2004
Thinnfeldia indica sp. nov.; Feistmantel, p. 35; pl. 39, figs 1, 1a; pl. 46, figs 1, 2, 2a
Pachypteris dalmatica Kern; Halle, pp. 43-44; pl. 4, figs 23-28, 33?
Thinnfeldia constricta sp. nov.; Halle, pp. 45-46; text- fig. 10; pl. 4, figs 29-32, 34
Thinnfeldia indica Feistmantel, emend. Zeba-Bano; Maheshwari & Bose, pp. 145-148;
text-fig. 2; pl. 1, figs 1-6; pl. 2, figs 8, 9
Thinnfeldia indica Feistmantel; Maheshwari, pp. 14-19; pl. 1, figs 1-4; pl. 2, figs 1-6
Pachypteris hallei Frenguelli; Gee, p. 180; Pl. 4, figs 36, 37
Thinnfeldia constricta Halle; Gee, pp. 182-183; pl. 5, figs 44-46
Komlopteris indica (Feistmantel); Barbacka, p. 348
Komlopteris indica (Feistmantel) Barbacka; Rees & Cleal, pp. 67-68; pl. 19, figs 2-4
Material. Two specimens: the first represented by a single frond fragment, the
second one – by a composite frond fragment. S78/119 KRAM-P; ZPAL Pl. 3/4.
53
Fig. 27. Komlopteris indica (Feistmantel) Barbacka, emend. Rees et Cleal; ZPAL PL 3/4 (Phot. M.
Dziewiñski). Scale bar 5 mm
Description. Laves pinnate. Leaf midrib c. 2 mm broad. Pinnules elongatelanceolate, widest in 1/3 of length, slightly pointed at apex (Fig. 27), 25–30 mm
long and 7–10 mm wide. Leaf margins lobate. Particular pinnules overlap each
other, they form an angle of 25–30° with respect to leaf axis. Nervation dense: 6
veins per 1 mm (Fig. 27).
54
Discussion. Attribution of our specimens to Komlopteris indica is based on shape
and size of pinnules which show slightly pointed apex and lobate margin.
Komlopteris indica was first described by Feistmantel (1877a) as Thinnfeldia
indica. Long discussion on systematic position of the genus Thinnfeldia resulted in
a conclusion that this genus is a younger synonym of the genus Pachypteris (see
Doludenko, 1971; Doludenko et al., 1998). The long-established species Pachypteris indica is, however, morphologically different (cf. Bose & Roy, 1967), moreover it differs from Komlopteris indica in cuticle structure (Bajpai & Maheshwari,
2000).
Barbacka (1994) created a new genus Komlopteris based on both morphology
and micromorphology structure of pinnules. To this genus she has also included
specimens earlier described as Thinnfeldia indica. Morphological features characteristic of the genus Komlopteris, including pinnate leaves with elongatelanceolate leaflets, with pinnate nervation, are well recognizable in our specimens.
From other species of the genus Komlopteris, the species K. indica differs in size
and shape of leaflets (Barbacka, 1994, pl. 2): leaflets of K. indica are smaller than
those of K. nordenskjoeldii, but larger than ovate leaflets of K. rotundata. Both
mentioned species show the presence of entire leaflet margins, in contrast to lobate
leaflets of K. indica. Leaflets of K. indica are broader than those of K. speciosa
(Cleal & Rees, 2003).
Halle (1913a) determined some specimens from Hope Bay as Pachypteris dalmatica. This species has been included by Gee (1989b) to Pachypteris hallei, but
retained as Komlopteris indica by Rees and Cleal (2004). The present author agrees
with their solution, however with one exception: Halle’s (1913a, pl. 4, fig. 35)
specimen does not belong to K. indica but to Pachypteris indica. This is indicated
by leaflets’ size much smaller than those of K. indica, and by entire leaf margins being characteristic of Pachypteris indica leaves (cf. Rees & Cleal, 2004).
Distribution. West Antarctica: Hope Bay, Antarctic Peninsula – Jurassic (Rees &
Cleal, 2004); India, Rajmahal Hills – Upper Jurassic/Lower Cretaceous (Barbacka,
1994),
Division Cycadophyta
Order Cycadales
Cycads first appeared at the Carboniferous/Permian transition (Taylor & Taylor, 1993), they reached apogee in Mesozoic, now being represented by relic forms.
Presently, about 200 cycad species represented by 11 genera are known. They occur mainly in tropic and subtropic, less frequently in temperate-warm areas: in
South Africa, Australia, South and Central America, and in SE Asia (Jones, 1998).
Genus Nilssonia Brongniart, 1825
Type species: Nilssonia brevis Brongniart
Nilssonia, a Mesozoic genus, includes cycads with lanceolate leaves which are
55
Fig. 28. Nilssonia taeniopteroides Halle. A – apical part of leaf and rachis from adaxial side, S78/105
KRAM-P; B – middle part of leaf, adaxial side, S78/36 KRAM-P; C – basal part of leaf, S78/102
KRAM-P; D – basal part of leaf and convex rachis (arrowed) from abaxial side, S78/25 KRAM-P.
Scale bar 10 mm
either entire or divided in lobes of different size. Lamina shows a distinct rachis,
concave from adaxial and convex from abaxial sides. Lateral nerves, usually single,
radiate from main vein at angles close to 90° (Taylor & Taylor, 1993).
Nilssonia taeniopteroides Halle, 1913
(Figs 28, 29)
1913a
1989b
Nilssonia taeniopteroides n. sp.; Halle, pp. 47-50; figs 11a-c; pl. 5; pl. 6, figs 1-7
Nilssonia taeniopteroides Halle; Gee, pp. 184-186; pl. 6, fig. 49
Material. 38 imprints of various parts of lamina. No complete leaves were
preserved: usually, their apical part is missing.
Specimens with preserved apical part of leaf: S78/35, S78/105 KRAM-P.
56
Fig. 29. Nilssonia taeniopteroides Halle. A – S78/105 KRAM-P; B – S78/36 KRAM-P; C – S78/35
KRAM-P; D – S78/102 KRAM-P; E – S78/25 KRAM-P. Scale bar 10 mm
Specimens with preserved leaf base: S78/25, S78/102, S78/161 KRAM-P.
Other specimens: S78/7-9, S78/11, S78/20, S78/23, 24, S78/28, 29, S78/33, 34, S78/36, S78/38, 39,
S78/41, S78/43, S78/54, 55, S78/57, S78/71, S78/91, S78/103, S78/106, S78/116-118, S78/126, S78/132,
S78/143, S78/149, S78/162, 163, S78/168 KRAM-P.
Description. Leaves long, entire. Longest leaf fragments maximum 14 cm long,
total leaf length unknown. Lamina of changing width: 1.5–5 cm broad in the
middle, tapering towards pointed apex (Fig. 28A) and towards base, where it forms
a petiole (Fig. 28C, D). Lamina with well pronounced rachis which is depressed
from adaxial side (Fig. 28A, B), and convex from abaxial side (Fig. 28D: arrowed).
Rachis width 0.5–0.7 mm, growing towards petiole. 20–30 lateral nerves per 1 cm
of rachis. Laterial nerves single, radiating at angles 70–90° from rachis (Fig. 28).
Discussion. Characteristic features, such as rachis depressed from adaxial and
convex from abaxial sides, and single lateral nerves, indicate attribution of our
specimens to the genus Nilssonia (see Seward, 1917; Harris, 1964).
57
Entire leaves with Nilssonia-type nervation, i.e., rachis with lateral nerves radiating from it nearly at right angles, but with unknown anatomical structure, were included to the genus Taeniopteris. This genus might include both cycads and
bennettitales (Harris, 1932a). Apart from their anatomic structure, these two genera
are distinguishable from each other by rachis development: it is adaxially depressed
but abaxially convex in Nilssonia, while it is convex both adaxially and abaxially in
Nilssoniopteris (see Harris, 1964, 1969). The last feature is, moreover, characteristic for leaves of the genus Doratophyllum (Harris, 1932a).
Rees and Cleal (2004) had at their disposal collections of specimens which did
not clearly show the rachis character: thus they included these specimens to the species Taeniopteris taeniopteroides.
Entire leaves with pointed apex, density of lateral nervation, and its angle with
midrib, all indicate attribution of our numerous specimens to the species Nilssonia
taeniopteroides (see Halle, 1913a; Gee, 1989b).
Leaves of Nilssonia taeniopteroides differ from those of N. obtusa and N. polymorpha by leaf width: it measures 1.5 cm in N. obtusa, and 3 cm in N. polymorpha
(cf. Harris, 1913a). Leaves of N. revoluta are 0.3–0.5 cm wide only, those of N.
grandiflora are as much as 21 cm wide (Schweitzer et al., 2000). N. tenuinervis
shows much denser lateral nervation: 40–45 nerves per 1 cm (Harris, 1964). Leaves
of N. undulata, N. orientalis and N. thomasi show incisions at leaf terminus; in our
specimens it is pointed.
Distribution. Argentina: Cerro de las Cabras – Triassic (Rhaetian), Patagonia, Río
Genua – Jurassic (Liassic); West Antarctica: Hope Bay – Jurassic; China – Tien
Shan – Jurassic (see Halle, 1913a; Jongmans & Dijkstra, 1961; Gee, 1989b; Rees &
Cleal, 2004).
Genus Crossozamia Pomel, emend. Gao et Thomas, 1989
Type species: Crossozamia moreana Pomel
The genus Crossozamia includes fossil cycad megasporophylls, formerly described as the genera: Cycadospadix, Schimper, 1870; Cycadospadix Saporta,
1875; Norinia Halle, 1927; Cycalacis Barale, 1981, and Primocycas Zhu et Du,
1981. These names have been included by Gao and Thomas (1989) to synonymy of
the genus Crossozamia Pomel. Megasporophylls of these cycads are known since
Permian, being very similar to the present-day cycad megasporophylls (Gao &
Thomas, 1989), and of the genus Cycas in particular.
Crossozamia mirabilis sp. nov.
(Fig. 30)
Material. A single sporophyll leaf impression 35 mm long, its lower part showing
paired oval traces, probably after seeds.
Name derivation. From mirabilis – as a reflexion of its beautiful shape.
Diagnosis. Sporophyll leaf spathulate. Its upper part is 35 mm wide, triangular,
58
Fig. 30. Crossozamia mirabilis sp. nov. Ociepa, holotype, S78/99 KRAM-P (photo M. Dziewiñski).
Scale bar 5 mm
with rounded apex, dissected in stripes which are 1.5–12 mm long and c. 1 mm
wide. Bundles consisting of 2 or 3 stripes are separated by deeper incisions. Lower
leaf part 8 mm broad. Fanwise nervation, nerves reaching leaf margin; in lower part
of leaf – c. 6 nerves per 1 mm.
Holotype. Fig. 30: S78/99 KRAM-P.
Location and type stratum. Mount Flora, Hope Bay, Antarctic Peninsula (West
Antarctica), Mount Flora Formation, Flora Glacier Member (in situ) – see Figs 5–7
and Birkenmajer (1988, 1993a).
Geological age. Jurassic (?Early Jurassic).
Discussion. Our specimen resembles best sporophyll leaves of cycads of the genus
Crossozamia. This is expressed in fanwise nervation and the shape of sporophyll
leaf upper part divided in narrow stripes. However, the species of the genus
Crossozamia known thus far (see Gao & Thomas, 1989), have upper parts of leaves
divided in stripes of equal length, while in Crossozamia mirabilis they are of
variable length and form bundles of 2 to 3 stripes divided from each other by deeper
incisions. Such a feature is unknown from other species of the genus Crossozamia,
thus it favours creation of a new species.
Our specimen resembles to some extent sporophyll leaves of Cycadospadix
dactylota (Harris, 1932b, pl. 10, figs 1-2), attributed by Florin (1933) to Bennettitolepis dactylota. He was of the opinion that it represented a sepal-like scale of ben-
59
Fig. 31. Crossozamia sp., S78/131 KRAM-P. Scale bar 5 mm
nettitalean flower. His flower-scale is similar in length (4 cm), but much smaller in
width (0.7–1 cm), and divides at top in widely spaced stripes. This is at variation
with respect to Crossozamia mirabilis n. sp. which has closely spaced stripes.
Our specimen is similar in nervation and shape to sporophyll leaf of Palaeocycas integer (Florin, 1933, text-tab. 11, pl. 1, fig. 1), however the latter has an entire
leaf margin. In nervation and shape of sporophyll leaf, Crossozamia mirabilis sp.
nov. resembles that of Archaeocycas whitei (Mamay, 1976, fig. 3). However, upper
part of Mamay’s leaf has not been preserved; Mamay guessed that it was entire.
Our species resembles sporophyll leaves of the present-day cycad species: Stangeria eriopus, Macrozamia riedlei and Dioon edule (see Kramer & Green, 1900,
fig. 205D, E, F); however these species have entire, and not dissected at top sporophyll leaves.
Crossozamia sp.
(Fig. 31)
Material. A single sporophyll leaf impression, S78/131 KRAM-P.
Description. Entire length of sporophyll leaf unknown, the preserved part is 5 mm
long and 7 mm broad, with length/width ratio of 0.7. Leaf base not preserved. Leaf
triangular in outline, rounded at apex. Sporophyll leaf margin dissected in narrow
stripes which are 1.5 mm long and c. 5 mm broad; apices of leaf stripes not
preserved. Nervation fanwise, without midvein (Fig. 31); in lower part of the
sporophyll leaf, 5 nerves per 1 mm. No traces of ovules or seeds are preserved.
Discussion. By its shape, nervation type (fanwise, without midvein), and sporophyll leaf dissected in stripes, our specimen resembles those of the genus Crossozamia. The above features make it most similar to Cycadospadix pasinianus (see
Zigno, 1873–1885, tab. 42, fig. 3; Seward, 1911, pl. 7, fig. 18). However, the latter
is 3–4 cm long and 4–5 cm wide, its stripes are 12 mm long and 1.5 mm wide
(Zigno, op. cit.).
Our specimen is also similar to a leaf-scale type B of Rees and Cleal (2004): both
are triangular in outline, their length/width ratio amounts to 0.6–0.7, the leaf margins are similarly dissected in narrow stripes. Uncertainty about shapes of stripes’
terminations which are not preserved in our specimen, does not allow to state
whether or not they narrowed apexward, as is the case with sporophyll leaf of Cyca-
60
dospadix pasinianus of Zigno (1873–1885) and Seward (1911), and leaf-scale B
type of Rees and Cleal (2004).
A much poorer resemblance of our sporophyll leaf, as regards its triangular
shape and dissection of leaf in narrow stripes, may be discerned in sporophyll leaf
of Primocycas muscariformis (Zhu & Du, 1981, fig. 1): the latter has much lower
(0.33) length/width ratio, and is larger.
Our specimen resembles to some extent sporophyll leaves of Cycadospadix
hennocquei of Saporta (1873–1891, pl. 46, figs 1-5): both are triangular in outline
and their margins are dissected in narrow stripes. However, sporophyll leaf of C.
hennocquei is larger and shows a higher (1.2) length/width ratio.
Sporophyll leaves of Crossozamia minor (Gao & Thomas, 1989, pl. 1, fig. 2), of
C. spadicia (Gao & Thomas, 1989, pl. 2, fig. 1), and of our specimen, are similar in
size, however their shapes are different.
Our specimen resembles to some extent upper parts of sporophyll leaves of the
present-day genus Cycas (see Kramner & Green, 1990, fig. 205A, B).
Fragmentary preservation of our specimen precludes its specific determination.
Order Bennettitales (Cycadeoidales)
The order Bennettitales is represented by extinct plants which were widespread
on Earth from Triassic through Cretaceous, inclusively. Contrary to cycads, the
majority of Benettitales were monoecious. Morphologically, the leaves of Bennettitales are very similar to those of cycads, however, they differ from the latter in
anatomic structure of epiderm, and in structure and position of stomatal apparatuses in particular (Taylor & Taylor, 1993).
Genus Otozamites Braun, emend. Watson et Sincock, 1992
Type species: Otozamites obtusus (Lindley et Hutton) Brongniart
The genus Otozamites includes monopinnate leaves with characteristic pinna
shape: their pinna base is asymmetric, with an auricle at the acroscopic side (Watson & Sincock, 1992, text-fig. 4e, f). Acroscopic and basiscopic veins are curving
acropetally at pinna base (Person & Delevoryas, 1982, text-fig. 6B).
Otozamites gramineus (Phillips) Harris, 1969
(Fig. 32)
1969
1999
2004
Otozamites gramineus (Phillips); Harris, pp. 29-33; text-figs 12, 13
Otozamites gramineus (Phillips) Phillips; van Konijnenburg-van Cittert & Morgans, p.
86; pl. 14, fig. 1
Otozamites gramineus (Phillips); Deng et al., p. 215; pl. 2, fig. 8
Material. 2 impressions of leaves, their bases and terminations not preserved.
S78/44, S78/155 KRAM-P.
Description. Leaves monopinnate (bases and terminations not preserved),
3.5–4 cm long and 2.5–4.7 cm broad. Particular pinnae narrow-ovate, rounded at
61
Fig. 32. Otozamites gramineus (Phillips) Harris, S78/44 KRAM-P. Scale bar 10 mm
apex, their bases extended to form acroscopic auricles. Auricle protruding c. 2 mm
above the remaining part of leaf. Pinnae 30 mm long and 5 mm wide, alternating
(but not overlapping), situated at an angle of 60–70° with respect to leaf rachis.
Nervation fanwise, nerves single, acroscopic (Fig. 32) and basiscopic, curving
acropetally at pinna base. Nervation density: 4 veins per 1 mm.
Discussion. Size and shape of pinnae, their auricles and nervation type, indicate
attribution of our specimens to the species Otozamites gramineus.
From O. leckenbyi, the leaves of O. gramineus differ in having a well pronounced auricle at pinnae bases; the latter is faintly developed in the former species
(see Harris, 1969). Leaves of our specimens have pinnae smaller than those of O.
graphicus which are usually 10 mm long and 4 mm broad (Halle, 1913a; Gee,
1989b). Leaves of O. gramineus generally resemble those of Zamites wendyellisae
(Watson & Sincock, 1992), however pinnae bases of the former species are asymmetric, with well developed auricle, while those of the latter species are symmetric,
devoid of auricle.
Distribution. England, Yorkshire – Jurassic (Aalenian), van Konijnenburg-van
Cittert and Morgans (1999); China, Xinhe Formation – Jurassic (Bathonian/
Callovian) – Deng et al. (2004).
Otozamites linearis Halle, 1913
(Fig. 33)
1913a
1913a
1978
1989b
2004
Otozamites linearis n. sp.; Halle, pp. 61-64; text-fig. 15; tab. 7, figs 1-4, 8, 9, 9a, 11
Otozamites sp.; Halle, tab. 7, figs 10, 18
Otozamites linearis Halle; Baldoni, pp. 3-6; pl. 1, figs 2, 8
Otozamites linearis Halle; Gee, pp. 188-189; pl. 6, figs 50-52
Otozamites? linearis Halle; Rees & Cleal, pp. 46-48; text-fig. 6
62
Material. A single impression of monopinnate
leaf, apex not preserved. S78/31 KRAM-P.
Description. Leaf monopinnate, long and
narrow, gradually narrowing basisward. Entire
leaf length unknown; our fragmentary specimen
is 17 cm long, 0.5 cm wide at base, and 2 cm wide
in the middle. Rachis 1.5 mm broad.
Pinnae widely ovate in outline, rounded at
apices, with base widened to form a characteristic
auricle (Fig. 33). Auricle protruding c. 1 mm
above pinna. Pinnae differ in size depending on
their position within leaf: in middle part of leaf
they are 10 mm long and 4 mm broad, those near
leaf base are 4 and 3 mm, respectively. Pinnae not
overlapping (Fig. 33).
Pinnae nervation fanwise, nerves single. Acroscopic and basiscopic veins curving acropetally at pinna base. Nervation dense: 5–7 veins per
1 mm.
Discussion. Size and shape of pinnae, their bases
forming well developed auricles, and pinnae nervation-type, indicate attribution of our specimen
to the species Otozamites linearis.
Leaves of O. linearis morphologically resemble those of other species of the genus Otozamites
known from South America, such as: O. albosaxatilis, O. parviauriculata, O. sanctae-crucis
and O. walkomii. O. parviauriculata and O. walFig. 33. Otozamites linearis Halle,
komii differ from O. linearis in having poorly deS78/31 KRAM-P. Scale bar 10 mm
veloped auricle, which – on the contrary – is well
developed in the latter species (Menéndez, 1966;
Archangelsky & Baldoni, 1972). Pinnae of O.
linearis are broader than those of O. albosaxatilis: in the latter species, they are
6–9 mm broad (Herbst, 1965). Pinnae of O. sanctae-crusis are imbricate (Feruglio,
1951 – fide Archangelsky & Baldoni, 1972), while in O. linearis they are well separated from each other. Moreover, leaves of the discussed species differ from those
of O. linearis in details of their anatomic structure (Baldoni, 1978), unfortunately
not preserved in our specimen.
Leaves of O. linearis differ from those of O. gramineus in pinnae sizes: in the
latter species they are larger, 30 mm long and 5 mm wide (Harris, 1969).
Distribution. West Antarctica, Hope Bay and Botany Bay – Jurassic (Halle,
1913a; Gee, 1989b; Rees & Cleal, 2004); Argentina, Plaza Huíncul – Jurassic
(Baldoni, 1978).
63
Fig. 34. Zamites antarcticus Halle, emend. Archangelsky et Baldoni. A – ZPAL PL 3/2; B – S78/45
Genus Zamites Brongniart, emend. Harris, emend. Watson et Sincock, 1992
Type species: Zamites gigas (Lindley et Hutton) Morris, emend. Harris
The genus Zamites includes monopinnate, equally narrow leaves, having pinnae
with symmetric bases (Watson & Sincock, 1992), with acroscopic veins curving
basipetally and basiscopic veins curving acropetally at pinna base (Ash, 1975,
text-fig.7; Person & Develoryas, 1982, text-fig. 6A).
Zamites antarcticus Halle, emend. Archangelsy et Baldoni, 1972
(Figs 34–36)
1913a
1913a
1913a
1913a
1966
1972
1985
1986
1989b
Zamites antarcticus n. sp.; Halle, pp. 58-60; text-fig. 13; tab. 7, figs 23, 24, 28?
Zamites sp.; Halle, tab. 7, figs 19, 20
Otozamites hislopi (Oldham) Halle; Halle, pp. 65-67; tab. 7, figs 5, 7, 21?
Ptilophyllum (Williamsonia?) pectinoides (Phillips) Halle; Halle, pp. 69-70; tab. 7, figs
25-27
Ptilophyllum longipinnatum n. sp.; Menéndez, pp. 14-16; text-figs 25-26, 27-31 (details
of epiderm structure); pl. 6, figs 25-26, 27-28 (details of epiderm structure)
Ptilophyllum antarcticum (Halle) Seward; Archangelsky & Baldoni, pp. 236-246; pl. 3,
figs 1, 2; pl. 14, figs 1-7 (cuticle)
Zamites antarctica Halle; Anderson & Anderson, p. 170; text-figs 1, 2; pl. 221
Ptilophyllum antarcticum (Halle) Seward, emend. Archangelsky et Baldoni; Baldoni, pp.
81-82; pl. 1, 2 (cuticle)
Zamites antarcticus Halle, emend. Archangelsky et Baldoni; Gee, pp. 190-192; p. 7, figs
56, 57
64
Fig. 35. Zamites antarcticus Halle, emend. Archangelsky et Baldoni. A – S78/165 KRAM-P; B –
ZPAL PL 3/2; C – S78/45 KRAM-P. Scale bar 5 mm
2004
Zamites ? antarcticus Halle; Rees & Cleal, pp. 43-44; pl. 12
Material. 11 fragments of different size, showing impressions of monopinnate
leaves. S78/19, S78/36 (positive and negative), S78/45, S78/84, S78/86, S78/154, S78/165, S81/14,
S81/16, S81/31 KRAM-P; ZPAL PL 3/1, PL 3/2.
Description. Leaves monopinnate, long and narrow, narrowing apexward. Entire
leaf length unknown, preserved fragments are 18–80 m long and 16–24 mm broad.
Rachis 1.25–1.5 mm broad.
Pinnae equally narrow in outline, slightly rounded at apices, with symmetric,
slightly widened bases (Fig. 34). Pinnae 10–18 mm long and 1.25–2 mm wide, their
length/width ratio is 8–9. Pinnae alternating, set at an angle of 55–60° with respect
to rachis.
65
Fig. 36. Zamites antarcticus Halle, emend. Archangelsky et Baldoni. A – S78/86 KRAM-P; B –
S78/19 KRAM-P. Scale bar 5 mm
Acroscopic veins curving basipetally, basiscopic veins slightly arcuate, curving
acropetally at pinna base. Nervation dense: 5–6 veins per 1 mm.
Discussion. Attribution of our specimens to the genus Zamites is based on the
presence of symmetric pinnae bases with a characteristic venation: acroscopic
veins are curving basipetally, while basiscopic veins are curving acropetally at
pinna base (see Ash, 1975; Person & Develoryas, 1982; Watson & Sincock, 1992).
Seward (1917), followed by Archangelsky and Baldoni (1972), attributed
leaves of Zamites antarcticus to the genus Ptilophyllum, as the species P. antarcticum. The present author disagrees with their conclusion as – basing on emended diagnosis of the genus Zamites by Watson and Sincock (1992) – this genus is characterized by pinnae symmetric at bases (op. cit., text-fig. 4a-d), while pinnae of the
genus Ptilophyllum are asymmetric at bases (Watson & Sincock, 1992, text-fig.
4g, h).
Our specimens, the specimens descibed by Halle from Hope Bay, and numerous
specimens from Argentina (see synonimic lists in: Halle, 1913a; Menéndez, 1966;
Archangelski & Baldoni, 1972; Baldoni, 1986) all show symmetric pinnae bases.
Thus, their attribution to the genus Zamites is justified.
Pinnae shape, size, length/width ratio, and nervation density of our specimens
confirm their attribution to the species Zamites antarcticus.
66
Fig. 37. Zamites pachyphyllus Halle, S78/172 KRAM-P. Scale bar 2.5 mm
Pinnae of Z. antarcticus differ from those of other species of this genus in having
length/width ratio higher than in Z. anderssoni (l/w = 2.3–5.5), Z. pachyphyllus
(l/w = 1.5–2.4) and Z. pusillus (l/w = 2.75–3.2) – see Gee (1989b). Pinnae of Z. antarcticus are more densely nervated than Z. anderssoni, the latter showing only 3–4
nerves per 1 mm (Gee, 1989b).
Occurrence. West Antarctica: Hope Bay and Botany Bay – Jurassic (Halle, 1913a;
Gee, 1989b; Rees & Cleal, 2004), Sobral Peninsula – Late Jurassic/Early Cretaceous (Baldoni, 1986); Argentina: Santa Cruz – Late Jurassic/Early Cretaceous,
Chubut – Late Jurassic/Early Cretaceous (Baldoni, 1986); South Africa: Zululand,
Makatini Formation – Early Cretaceous (Anderson & Anderson, 1985).
Zamites pachyphyllus Halle, 1913
(Fig. 37)
1913a
1913a
1989b
2004
Zamites pachyphyllus n. sp.; Halle, pp. 60-61; fig. 14; tab. 7, figs 14-16
Otozamites abbreviatus Feistmantel; Halle, p. 67; tab. 7, fig. 17
Zamites pachyphyllus Halle; Gee, pp. 192-193; pl. 7, figs 58, 59
Zamites pachyphyllus Halle; Ociepa, pl. 1, fig. 3
Material. 3 fragments of poorly preserved imprints of monopinnate leaves, their
apices not preserved. S78/83, S78/114, S78/172 KRAM-P.
Description. Leaves monopinnate, long, narrow, tapering basisward. Entire length
67
and width of leaves unknown, the preserved fragments are 20–42 mm long and
7–12 mm wide. Rachis 2.5–3 mm broad.
Pinnae broadly ovate in outline, rounded at apex, with symmetric base, 4–7 mm
long and 2–4 mm wide. Their length/width ratio is 1.5–2.4. Pinnae alternating, attached at an angle of 60° to rachis.
Pinnae of the investigated specimens do not possess winglets. Their nervation is
poorly visible: acroscopic veins are curving basipetally, while basiscopic veins ar
curving acropetally at pinna base (Fig. 37). Nervation is dense: 4 veins per 1 mm.
Discussion. Poor preservation of our specimens does not allow to decide whether
or not their pinnae possessed winglets, a feature considered diagnostic (Gee,
1989b). However such features as: broadly ovate shape of pinnae, their size, and
lenght/width ratio, allow to attribute them to the species Zamites pachyphyllus.
Leaves of Z. pachyphyllus differ from those of Z. anderssoni, Z. antarcticus and
Z. pusillus in having broadly-ovate pinnae, the latter being elongated in Z. anderssoni and Z. antarcticus, or narrowly-ovate in Z. pusillus (Halle, 1913a; Gee,
1989b). Moreover, pinnae of Z. pachyphyllus show a smaller length/width ratio
than those of Z. antarcticus and Z. pusillus (Halle, 1913a; Gee, 1989b).
Distribution. West Antarctica: Hope Bay – Jurassic (Halle, 1913a; Gee, 1989b).
Zamites pusillus Halle, 1913
(Fig. 38)
1913a
1968
1989b
Zamites pusillus n. sp.; Halle, p. 56; fig. 12a; tab. 7, fig. 12
Zamites pusillus Halle; Herbst & Anzotegui, p. 188; fig. 1H
Zamites pusillus Halle; Gee, p. 193; pl. 7, fig. 60
Material. Six fragments of monopinnate leaves. S78/26, S78/42, S78/63, S78/72, S78/125,
S78/140, S78/161 KRAM-P (specimens S78/72 and S78/161 are represented by positive and
negative imprints).
Description. Leaves monopinnate, long, narrow, tapering apexward (Fig. 38).
Pinnae narrowly ovate in shape, with symmetric base and slightly rounded apex,
5.5–8 mm long, 2–2.5 mm broad. Their lenght/width ratio equals to 2.75–3.2.
Pinnae alternating, attached to rachis at an angle of 50–65°.
Acroscopic veins curving basipetally, basiscopic veins curving acropetally at
pinna base. Nervation dense: 5 veins per 1 mm.
Discussion. Attribution of our specimens to the species Zamites pusillus is based
on narrowly-ovate outline of pinnae, their size, and length/width ratio (Halle,
1913a; Gee, 1989b).
Narrowly-ovate shape of pinnae of Z. pusillus distinguish them from pinnae of
Z. anderssoni and Z. antarcticus, which are elongated, and from broadly-ovate pinnae of Z. pachyphyllus (Halle, 1913a; Gee, 1989b). Pinnae length/width ratio in Z.
pusillus is higher than that of Z. pachyphyllus (l/w = 1.5–2.4), but smaller than that
of Z. antarcticus (l/w = 8–9) – see Halle (1913a), Gee (1989b).
68
Fig. 38. Zamites pusillus Halle. A – S78/72 KRAM-P; B – S78/125 KRAM-P; C – S78/26 KRAM-P; D
– S78/42 KRAM-P; E – S78/140 KRAM-P; F – S78/63 KRAM-P. Scale bar 5 mm
Rees and Cleal (2004) included specimens determined by Halle (1913a) as
Zamites pusillus to the species Otozamites? latior. According to the present author,
this is incorrect, as both symmetric pinnae bases (cf. Watson & Sincock, 1992), and
type of their nervation – acroscopic veins curving basipetally and basiscopic veins
curving acropetally at pinna base (see Ash, 1972; Person & Develoryas, 1982) – indicate their attribution to the genus Zamites. Such features are common for both
Halle’s (1913a) and our specimens.
Occurrence. West Antarctica: Hope Bay – Jurassic (Halle, 1913a; Gee, 1989b);
Argentina: Taquetrén, provincia Chubut – Middle Jurassic (Herbst & Anzotegui,
1968).
69
Cycadophyta incertae sedis
Genus Conites Sternberg, emend. Cleal et Rees, 2003
Type species: Conites bucklandii Sternberg
The genus Conites includes mineralized stems or stem impressions of cycads
and bennettitales, with petiole bases, or scars after petioles, preserved (Cleal &
Rees, 2003). Previously, such fossils were classified as Bucklandia Sternberg,
1825, Clathrania Brongniart, 1828, Yatesia Carruthert, 1870, and Fittonia Carruthert, 1870. They were included to synonimic list of the genus Conites by Cleal
and Rees (2003).
Conites sp.
(Fig. 39)
Material. Single stem imprint with scars after petioles preserved. S81/15 KRAM-P.
Description. Entire length and width of stem unknown, preserved fragment is 9 cm
long and 5 cm broad. Numerous scars after petioles visible at the surface. Scars
rhomboid in outline, depressed, 5 mm long and 7 mm wide, arranged in regular
helical rows (Fig. 39).
Discussion. Lack of preserved anatomical structure precluded attribution of our
specimen to either benettitales or cycads.
Preservation of our specimen in form of imprint only does not allow to compare
it with the species of the genus Conites, such as: C. anomala (Seward, 1917; Watson & Sincock, 1992), C. millerana, C. yatesii (Seward, 1917) or C. pustulosa
(Harris, 1969), which are recognizable on their anatomic structure.
Our specimen resembles most the stem of C. indica (Seward, 1917, fig. 579).
The shape and size of its petiole scars are similar to those of Seward’s: in his specimens, the scars are also rhomboid in outline, 4–7 mm high and 7–10 mm broad.
Poor state of preservation does not allow a more detailed classification of our
specimen.
Class Coniferophyta
Order Coniferales
Family Araucariaceae
Araucariaceae represent a conifer family known since Late Triassic. Its greatest
diversification occurred during Jurassic. Presently, representatives of this family
occur as relics. Worldwide distribution of Araucariaceae was achieved during
Mesozoic (Taylor & Taylor, 1993). Presently, their distribution is restricted to
tropical and subtropical regions of Southern Hemisphaere, with the exclusion of
Africa; however, they may even reach warm-temperate climatic zones (Kramer &
Green, 1990). For many decades, only two living genera of this family were known:
Araucaria and Agathis, the third one – Wollemia – has recently been discovered in
Australia (Jones et al., 1995).
70
Fig. 39. Conites sp.: stem impression with scars after petioles, S81/15 KRAM-P. Scale bar 5 mm
Genus Araucarites Presl, 1838
Type species: Araucarites goepperti Presl
To the genus Araucarites have been included cuneate bract and ovuliferous
scales, known worldwide from Jurassic strata (Cleal & Rees, 2003).
Araucarites antarcticus (Gee) comb. nov.
(Figs 40, 41)
1913a
1989b
2004
Araucarites cutchensis Feistmantel; Halle, pp. 72-74; fig. 16; pl. 8, figs 3-10
Araucaria antarctica n. sp.; Gee, pp. 196-197; pl. 8, figs 64-68a, 72a
Araucarites cf. cutchensis Feistmantel; Rees & Cleal, pp. 62-64; pl. 15, figs 1, 2
Material. 23 imprints of ovuliferous scales. S78/19, S78/33, S78/95, S78/107 (two scales),
71
Fig. 40. Araucarites antarcticus (Gee) comb. nov.: ovuliferous scales with traces after seeds. A –
S78/128 KRAM-P; B – S78/169 KRAM-P; C – S78/164 KRAM-P; D – S82/11 KRAM-P. Scale bar 5 mm
S78/108, S78/120 (two scales), S78/132, S78/140, S78/142, 143, S78/145, S78/150 (two scales),
S78/162, S78/164 (two scales), S78/169-171, S82/11 KRAM-P.
Description. Cuneate ovuliferous scales, 7–18 mm long (winglet not counted).
Upper margin of scales 5–15 mm broad, their lower margin 1–6 mm broad. Upper
scale margin elongated, forming a characteristic winglet (Fig. 40A) 1.5–3 mm long
and 1.5–2 mm broad. Traces after single seeds visible at scales; they are obovate in
outline (Fig. 40C, D), 6–10 mm long and 1.5–5 mm broad.
Discussion. Sizes of ovuliferous scales and the presence and size of winglets,
indicate their attribution to the species Araucarites antarcticus (= Araucaria
antarctica Gee, 1989b).
Ovuliferous scales of Araucarites antarcticus differ from those of A. cutchensis
in possessing a winglet, which is missing in the latter species (Bose & Banerji,
1984).
Fig. 41. Araucarites antarcticus (Gee) comb. nov.: A – S78/150 KRAM-P; B – S78/33 KRAM-P; C –
S78/169 KRAM-P; D – S78/128 KRAM-P; E – S82/11 KRAM-P; F – S78/164 KRAM-P. Scale bar 5 mm
72
Ovuliferous scales of A. antarcticus have winglets shorter than those of A. janaianis (winglet 3–8 mm long – Bose & Banerji, 1984) and those of A. wollemiaeformis (winglet 12 mm long – Cantrill & Falcon-Lang, 2001).
Ovuliferous scales of A. antarcticus are larger than those of Araucarites minutus
(scale height 0.7–1.5 mm, scale width 0.6–1.1 mm – Bose & Banerji, 1984), of
Araucarites minimus (scales 7 mm long and 3.5–5 mm wide – Archangelsky,
1966), and of A. citadelbastionens (scales 7 mm long and 3 mm wide – Cantrill &
Falcon-Lang, 2001).
Ovuliferous scales of A. antarcticus are smaller than those of A. baqueroensi
(scales 3 mm long and 2 cm broad – Archangelsky, 1966).
Gee (1989b) found that in the Hope Bay collections, both bract and ovuliferous
scales in fossil plants of the Araucariaceae family, differ from the species known so
far. This was the reason for creating a new species. Taking into account that these
scales were very similar to those of the present genus Araucaria, she included her
new species to this genus. However, with the recent discovery of the genus Wollemia, the third living genus of the Araucariaceae family, her solution does not
seem justified, as both the fossil bract and ovuliferous scales might represent either
the genus Araucaria or the genus Wollemia (cf. Chambers et al., 1998; Cantrill &
Falcon-Lang, 2001). This was the reason of creating a new combination: Araucarites antarcticus.
Occurrence. West Antarctica: Hope Bay – Jurassic (Halle, 1913a; Gee, 1989b;
Rees & Cleal, 2004).
Family Taxodiaceae
The family Taxodiaceae is known since Jurassic (Taylor & Taylor, 1993); it has
numerous representatives in fossil record. Presently, 9 genera with 16 species are
known, distributed predominantly in temperate-warm zones, mainly in Northern
Hemisphaere, with only one genus (Arthrotaxis) occurring in Southern Hemisphaere (Kramer & Green, 1990).
Genus Sphenolepis Schenk, emend. Harris, 1953
Type species: Sphenolepis sternbergiana (Dunker) Schenk.
The genus Sphenolepis includes foliated shoots with helically arranged small
scaly leaves. Female cones are small, roundish, terminally located (Harris, 1953).
The genus Sphenolepis was created in 1871 by Schenk. In 1881, Heer changed
its name to Sphenolepidium, arguing that the generic name Sphenolepis was earlier
used for a fish genus. Harris (1953) reinstated Schenk’s name Sphenolepis considering botanical nomenclature as independent of zoological one.
Representatives of the genus Sphenolepis are known from Jurassic and Cretaceous (Jongmans & Dijkstra, 1974).
73
Sphenolepis sp.
(Figs 42, 43)
1913a
1989b
2004
Sphenolepidium oregonense Fontaine; Halle, pp. 80-81; pl. 9, figs 5b, 9-11, 13
Pagiophyllum sp. A; Gee, pp. 198-199; pl. 8, figs 70, 71
Pagiophyllum sp.; Rees & Cleal, p. 59; pl. 16, figs 3, 4
Material. Seven imprints of cones, none complete, and more than 100 imprints of
foliated shoots. Their state of preservation is variable: morphological structure is
recognizable in a part of the collection, in other specimens, which are pyritized,
morphological structure is poorly visible.
Specimens with cones preserved: S78/2a, S78/24, S78/37, S78/66 (2 cones) KRAM-P.
Other specimens: S78/1b, S78/2a (positive and negative), S78/2b, S78/4b, S78/5a, S78/7, 8,
S78/10, S78/15, S78/19 (positive and negative made from S78/36) S78/20-25, S78/28-29, S78/31,
S78/33, S78/35-37, S78/39, S78/41-42, S78/45, S78/47-49, S78/51, S78/54, S78/58, S78/60, S78/64-66,
S78/68-71, S78/73, S78/85, S78/89-93, S78/95, S78/104-105, S78/108, S78/112, S78/114, S78/116, S78/119123, S78/126, S78/129-130, S78/134-136, S78/138-140, S78/142, S78/146-150, S78/157-158, S78/160-164,
S78/168-172, S81/4-5, S81/12-13, S81/20, S82/10-11 KRAM-P; ZPAL PL 3/8, 9.
Fig. 42. Sphenolepis sp. A – S78/2b KRAM-P; B – S78/2a KRAM-P; C – S78/24 KRAM-P; D – S78/37
KRAM-P; E – S78/138 KRAM-P; F – S78/66 KRAM-P; G – S78/135 KRAM-P; H – S78/66 KRAM-P.
Scale bar 2.5 mm
74
Fig. 43. Sphenolepis sp. A – S78/66 KRAM-P; B – S78/37 KRAM-P; C – S78/2a KRAM-P; D – S78/66
KRAM-P; E – S78/138 KRAM-P; F – S78/24 KRAM-P; G – S78/135 KRAM-P. Scale bar 2.5 mm
Description.
Cones. Cones loose, terminally located on foliated twigs. Due to incompletely
preserved cones, their shape is difficult to reconstruct: they probably were round in
outline. Preserved cone fragments are 5–8 mm long and 5–9 mm broad, their axis is
c. 0.5 mm wide, 3–5 scales are visible on cones. Scales attached to twig axis at angles 40–90°. Scales macelliform in outline, at cone apex some are bent axisward.
Cone apex moderately rounded (Fig. 42B) or blunt (Fig.42F), devoid of winglet.
Scales ribbed, two ribs visible onat each scale (Fig. B–H).
Foliated twigs. Twig fragments 5–75 mm long and 1–2 mm broad, total length
of twigs unknown. Some twigs bifurcate at angles 20–45°.
Leaves helically arranged, very dense, overlapping scaliform (Fig. 42A), narrowly-ovate in outline, with sharper or more blunt tip, 2–2.5 mm long and 0.5 mm
broad.
Discussion. Attribution of our specimens to the genus Sphenolepis is based on
occurrence of terminally located small cones, and thin twigs with helically arran-
75
ged, dense minute leaves which are narrowly-ovate in shape and possess a more or
less pointed tip (Harris, 1953).
Our specimens resemble those of the genus Arthrotaxites (Halle, 1913b) as refers to twig size, leaves arrangement and length, and terminal position of small
cones. However, scales of the genus Sphenolepis are devoid of winglet which occurs in scales of cones of the genus Arthrotaxites (Halle, 1913b).
Our specimens are very similar to twigs of the genus Pagiophyllum, especially
as refers to their leaf outlines. However, to the genus Pagiophyllum are included
twigs without reproductive organs (Seward, 1919), while our twigs show the presence of terminally-located cones.
By their size, shape and helical arrangement of leaves, our twigs resemble also
those of Nothodactrium warrenii (Townrow, 1967a), however cone scales of the
genus Sphenolepis have entire scale terminations, while those of the species N.
warrenii are partitioned in three segments (op. cit., fig 5C, D).
Our twigs and cones resemble best those of the species Sphenolepidium oregonense as described by Fontaine in Ward (1905, pp. 133-134, pl. 36, figs 3-8). The
similarity lies in shape and size of leaves and their arrangement on twigs; their
cones are similar in shape and size. However, Fontaine (in Ward, 1905) wrote that
preservation of his cones was poor. His figures do not allow to state whether his and
our specimens belong to the same species.
Gee, 1989b; Rees & Cleal, 2004).
Family Palissyaceae
The family Palissyaceae, represented by extinct coniferous plants, was created
by Florin (1958). Its representatives occurred during Triassic and Jurassic (Taylor
& Taylor, 1993).
Genus Stachyotaxus Nathorst, 1886
Type species: Stachyotaxus septentrionalis (Agardh) Nathorst
The genus Stachyotaxus is characterized by cones in which each scale contains
two seeds (Taylor & Taylor, 1993, fig. 21.33); the cones are located terminally on
twigs which are covered with equally-narrow leaves. The genus is known from Triassic and Jurassic strata (Rao, 1964).
Stachyotaxus sp.
(Figs 44, 45)
1913a
Conites?; Halle, p. 81; pl. 9, figs 12, 12a
Material. Two cone imprints with traces of seeds preserved, S78/149, S78/160
KRAM-P.
Description. Cones elongated in outline, 7–15 mm long and 7 mm broad, their axis
is 1 mm broad. 3–5 seed-scales present in cones. Seed-scales obovate, longitu-
76
Fig. 45. Stachyotaxus sp. A – S78/160
KRAM-P; B – S78/149 KRAM-P. Scale
bar 2.5 mm
Fig. 44. Stachyotaxus sp., cones with simple
sterile leaves. A – S78/160 KRAM-P; B – S78/149
KRAM-P. Scale bar 2.5 mm
dinally ribbed, with rounded termini (Fig. 45), bent towards axis. Scales 4–5 mm
long and 1–1.5 mm broad, set at an angle of 40–55° with respect to axis. Adaxial
part of seed-scales with traces of oval seeds, 2 mm long and 1 mm broad.
Sterile leaves occur at twig below cone. They are equally-narrow, decurrent,
3–3.5 mm long and 0.5 mm broad, set at an angle of 80–90° with respect to axis.
Discussion. Terminal location of cones, and equally-narrow sterile leaves on twigs
below cones, indicate their attribution to the genus Stachyotaxus. Traces of only
single, instead of double, seeds (see Taylor & Taylor, 1993) in our specimens are
probably an effect of overshadowing, the seeds being positioned one behind
another.
Gee (1989b) suggested attribution of some cones from Hope Bay to the genus
Stachyotaxus, but stated that her specimens have leaves different from those of this
genus. This is correct with respect to cones of the genus Sphenolepis, the leaves of
which located on twig below the cone are scaliform; however, both in our specimens and in Halle’s (1913a) Conites sp., below cones there occur equally-narrow
leaves that is a typical feature of the genus Stachyotaxus.
By their size and mode of arrangement of seed-scales, our specimens resemble
77
most those of the species Stachyotaxus sampathkumarani described from Jurassic
of India (Rao, 1964, pl. 1). However preservation state of our specimens, unclear
position of double seeds, and occurrence of single leaves at twigs below cones, allow to attribute them to the genus Stachyotaxus only.
Family unknown
Genus Brachyphyllum Lindley et Hutton, emend. Harris, 1979
Type species: Brachyphyllum mamillare Lindley et Hutton (designated as such by Harris, 1979)
The genus Brachyphyllum includes shoots of conifers with helically arranged
leaves of a characteristic structure: “leaf composed of a basal cushion tapering into
a small free part, length of free part (upper surface beyond leaf cushion) or total
height of leaf and cushion (outwards from shoot) less than width of leaf cushion”
(Harris, 1970, p. 4).
A part of the species belonging to the genus Brachyphyllum, e.g., B. mamillare
Lindley & Hutton, is being attributed to the family Araucariaceae (Stockey, 1982),
other species, e.g., B. crucis Kendall, to the family Cheirolepidiaceae (Watson,
1988). This is based on structure of the reproductive organs, and of cuticle structure
of leaves.
Brachyphyllum sp.
(Figs 46, 47)
1913a
1989
2004
Brachyphyllum sp.; Halle, pp. 79-80; pl. 8, figs 41, 42a; pl. 9, figs 14-16
Brachyphyllum sp.; Gee, pp. 199-200; pl. 8, fig. 69
Brachyphyllum sp.; Rees & Cleal, pp. 54-56; pl. 15, fig. 3
Material. Four impressions of shoots covered with leaves. S78/64, S78/115, S78/128,
S78/162 KRAM-P.
Descripton. Preserved fragments are 12–30 mm long and 3–6 mm broad, their
entire length is unknown. One of the shoots is branched (Fig. 47C). Leaves
helically arranged, rhomboid in outline, their length approximately equals width
which amounts to 1–2.5 mm. A ridge is marked on some leaves (Fig. 46).
Discussion. Rhomboid, helically arranged leaves, their length almost equalling
width, indicate attribution of our specimens to the genus Brachyphyllum. Shoots of
the genus Brachyphyllum differ from those of the genus Pagiophyllum in leaf
shapes: leaves of the former genus are nearly equidimensional, while those of the
latter genus are longer than wide (Harris, 1979; Gee, 1989b). The state of preservation, and the lack of cuticle in particular, do not allow but generic determination
of our specimens.
Genus Elatocladus Halle, 1913, emend. Harris, 1979
Type species: Elatocladus heterophyllus Halle
The genus Elatocladus includes conifer twigs with elongated, flattened leaves
78
Fig. 47. Brachyphyllum sp. A – S78/115
KRAM-P; B – S78/64 KRAM-P; C – S78/162
Fig. 46. Shoot of Brachyphyllum sp.: a ridge
is marked on some leaves, S78/115 KRAM-P.
Scale bar 2.5 mm
showing a single nerve (Harris, 1979). The most numerous species of this genus are recorded from Triassic
through to Cretaceous, there are also Middle and Upper
Carboniferous (e.g., E. kassagatschica) and Tertiary
ones (e.g., E. kerguelensis) – Jongmans & Dijkstra
(1973a).
Elatocladus confertus (Oldham et Morris) Halle, 1913
(Figs 48, 49)
1877a
1877a
1877b
1913a
1913a
1913a
1956
1976
1984
1989b
1989b
2000b
2001
2001
2004
Palissya conferta Oldham et Morris; Feistmantel, pp. 85-86; pl. 45, figs 4, 5, 8
Palissya, besides P. indica; Feistmantel, pl. 45, fig. 9
Palissya conferta Oldham et Morris; Feistmantel, pp. 21-22; pl. 8, figs 1-4
Elatocladus conferta (Oldham et Morris); Halle, pp. 86-87; pl. 8, figs 26-40
Elatocladus heterophylla n. sp.; Halle, pp. 84-85; fig. 18a, b; pl. 8, figs 12-14, 17-25
Elatocladus sp.; Halle, pl. 9, fig. 2
Elatocladus heterophylla Halle; Menéndez, pp. 327-328; pl. 2, figs 5, 6
Elatocladus conferta (Oldham et Morris) Halle; Maheshwari & Singh, p. 121; pl. 2, figs
14-15
Elatocladus confertus (Oldham et Morris) Halle; Bose & Banerji, pp. 85-87; text-fig.
50A-D; pl. 36, figs 1-5
Elatocladus confertus (Oldham et Morris) Halle; Gee, pp. 201-202; pl. 8, fig. 75
Elatocladus heterophyllus Halle; Gee, p. 200; pl. 8, figs 73, 74
Elatocladus confertus (Oldham et Morris) Halle; Cantrill, pp. 190-191; fig. 3b, c
Elatocladus confertus Halle; Cantrill & Falcon-Lang, pp. 139-140; pl. 7A, C-E
Elatocladus conferta Oldham et Morris; Thorn, fig. 6g
Elatocladus confertus (Oldham et Morris) Halle; Rees & Cleal, pp. 60-62; pl. 16, figs 5,
6; pl. 17, figs 1, 2
79
Fig. 48. Two types of leaf arrangement in Elatocladus confertus (Oldham et Morris) Halle. A –
S78/6a KRAM-P; B – S78/156 KRAM-P. Scale bar 10 mm
Material. More than 60 foliated twigs. S78/2a, 2b (positive and negative), S78/5a, 5b
(positive and negative), S78/6a, 6b (positive and negative), S78/15, S78/21, S78/27, S78/33,
S78/40, S78/46, S78/50, S78/53, S78/56, S78/59, S78/65, S78/69, S78/75, S78/79, S78/82, S78/88, S78/100,
S78/104, S78/108, 109, S78/112, 113, S78/118-121, S78/124, S78/127-129, S78/132, S78/134, S78/138-140,
S78/142, S78/146-148, S78/150, S78/152, 153, S78/158, S78/160,161, S78/169, S79/2-4, S79/8 (and its
negatives, not numbered), S81/15, S82/10 KRAM-P; ZPAL Pl 3/8, 3/15, 3/19.
Description. Fragmentary twigs, single or branching, 9–95 mm long, entire length
unknown. Two morphogroups based on shape and size of leaves may be distinguished.
Fig. 49. Elatocladus confertus (Oldham et Morris) Halle. A – S78/6a KRAM-P; B – S78/104
KRAM-P; C – S78/108 KRAM-P. Scale bar 10 mm
80
The first group (Figs 48A, 49A) includes twigs 6–10 mm long with planispirally
arranged leaves. Leaves are attached to axis at angles 70–100°; they are 3–11 mm
long and 1–2 mm broad. Leaves stiff, equally narrow, pointed at apices, with a single nerve passing from leaf base to apex.
The second group (Figs 48B, 49B, C) includes twigs 3–6 mm broad, with leaves
helically arranged, overlapping, set at angles of 20–30° with respect to axis, pointed
at apices, 4–6 mm long and 1 mm broad.
Discussion. Leaf shape and size and their arrangement in twigs (see Halle, 1913a;
Gee, 1989b; Rees & Cleal, 2004) indicate attribution of our specimens to the
species Elatocladus confertus. The first group of our specimens belongs to Elatocladus confertus (Oldham et Morris) Halle, the second one – to Elatocladus
heterophyllus Halle. Rees and Cleal (2004, pl. 17, fig. 1) found a specimen with
both types of foliage arrangement present on the same twig; this indicates that both
taxa belong to one and the same species – Elatocladus confertus. Such diversification in foliage structure is common in conifers at present. It is known, e.g., in
Dacrydium biforme from the Podocarpaceae family (Gaussen, 1973). In the
Cupressaceae family, juvenile leaves usually differ from those of mature ones, and
both may occur in the same individual; this phenomenon may also be induced
experimentally (Kramer & Green, 1900). In Xanthocyparis vietnamensis, a recently described new species of the family Cupressaceae, even three types of leaves
occur in mature individuals (Farjon et al., 2002).
Elatocladus confertus differs from other species of the genus Elatocladus in
shape and size of leaves. Leaves of E. confertus are smaller than those of E. zamioides, E. ramosus, E. planus and E. linearis (Townrow, 1967b; Harris, 1979; Cantrill
& Falcon-Lang, 2001) but larger than those of E. setosus (Harris, 1979). Leaves of
E. confertus are equally-narrow, while those of E. sidericus are ovate in shape (Harris, 1979). Leaves in the first group of twigs of E. confertus are arranged planispirally, but in the second group they closely overlap, while in E. laxus they are neither
planispiral nor overlapping (Harris, 1979).
Florin (1940) was of the opinion that twigs of the genus Elatocladus from Hope
Bay belong to the Podocarpaceae family. Cantrill and Falcon-Lang (2001) considered this attribution as probable, based on leaf cuticle structure (as known from
specimens derived from India): their investigations were conducted with the use of
light microscope only, while a scanning microscope examination would – according to them – be a proper one.
Occurrence. New Zealand: Mount Potts and Clent Hills, Canterbury – Triassic
(Rhaetian) – Jongmans & Dijkstra (1973), Mokaia, Gore, Southland – Early
Jurassic, Curio Bay, Waikawa, Southland – Middle Jurassic; West Antarctica:
Hope Bay and Botany Bay – Jurassic (Halle, 1913a; Gee, 1989b; Rees & Cleal,
2004), Alexander Island – Cretaceous (Albian) – Cantrill & Falcon-Lang (2001),
South Orkney Islands – Early/Middle Jurassic (Cantrill, 2000b); India: Rajmahal
Hills (Rajmahal stage), Bindrabun, Onthea, Murrero, Golapili, Madras, Nellore
Kavali Talug (Jabalpur stage), Sher River (Satpura Basin), S Rewah (Kota stage ?),
Chirakunt, Khatangi Hill Bihar – Jurassic; Australia, Victoria: Jeetho, Bena, Jeetho
81
Whitelaw, Jumbunna, Jumbunna East, Rainbow Creek, Moyarra, Woolmamai,
Burne’s Creek – Early Jurassic; Argentina: Río Atuel, Mendoza – Early Jurassic;
Central Asia: Basin of River Zeravshan – Jurassic (Jongmans & Dijkstra, 1973a).
Elatocladus sp.
(Fig. 50)
Material. Three fragments of foliated twig impressions.
S78/75, S78/156, S78/174 KRAM-P.
Description. Fragmentary twigs 18–38 mm long and
12–15 mm broad. Axis 1.5–2.5 mm broad. Leaves
planispiral, set at an angle of 60–70° with respect to axis.
Leaves delicate, equally-narrow in outline, pointed at
apices (Fig. 50), 6–9 mm long and 0.8 mm broad. A single
nerve passes from base to apex in every leaf (Fig. 50).
Discussion. Equally-narrow leaves with a single nerve
allow to attribute our specimens to the genus Elatocladus.
Our specimens resemble most the species Elatocladus
confertus: they have planispirally arranged leaves of
analogous length as in this species (cf. Halle, 1913a; Gee,
1989b; Rees & Cleal, 2004). From this species, our specimens differ in smaller width of leaves (barely 0.8 mm)
and in their delicate character.
For comparison with other species of the genus Elatocladus – see discussion under E. confertus. It is possible
that our specimens represent a third type of foliage characteristic of this species; however, they were found as
separate twigs, and not as parts of a twig with two other
types of foliage present.
Genus Pagiophyllum Heer, emend. Harris, 1979
Type species: Pagiophyllum circinicum (Saporta) Heer
Fig. 50. Elatocladus sp.
To the genus Pagiophyllum have been included coni- A – S78/156 KRAM-P; B –
fers with helically arranged leaves, with a characteristic S78/174 KRAM-P. Scale
structure: “leaf about as broad as its basal cushion, length bar 5 mm
exceeding width of cushion” (Harris, 1979, s. 23) and – in
transverse cross-section –leaf width greater than its thickness.
The genus Pagiophyllum includes twigs devoid of cones (Seward, 1919). Some
species of this genus are being included to the Cheirolepidiaceae family, based on
accompanied reproductive organs (Watson, 1988). The genus occurred from
Triassic through Cretaceous (Jongmans & Dijkstra, 1973b).
82
Pagiophyllum feistmantelii Halle, emend. Townrow, 1967
(Fig. 51)
1913a
1967c
1989b
2004
Pagiophyllum feistmantelli n. sp.; Halle, pp. 76-78; fig. 17; pl. 9, fig. 17-17b
Pagiophyllum feistmanteli Halle; Townrow, pp. 167-169; fig. 1G; fig. 2F, G; fig. 6D
(stomatal apparatus); fig. 7C, D; fig. 8A, B; fig. 9 (details of epiderm structure); fig.
11E (stomatal apparatus); pl. 1, figs C-D, E-F (details of epiderm structure)
Pagiophyllum feistmantelii Halle, emend. Townrow; Gee, pp. 197-198; pl. 8, fig. 72b
Pagiophyllum feistmantelii Halle, emend. Townrow; Rees & Cleal, p. 58; pl. 15, figs 4, 5
Material. An impression of branched twig, S78/27 KRAM-P.
Description. Twig doubly branched. Preserved fragment of main shoot is 42 mm
long and 8 mm wide. The 1st order branch is 35 mm long and 6 mm broad, shooting
off the main one at an angle of 60°. The 2nd order branches are 8–12 mm long and c.
3 mm broad, shooting off the 1st order ones at an angle of 40–60°. Helically
arranged leaves rhomboid to triangular in outline (Fig. 51). Triangular leaves are
clearly visible at last shoots. Leaves differ in size depending on position on
branched twig: they are larger at main shoot, and smaller at lateral ones, 2–3 mm
long and 1.5–2.5 mm wide.
Discussion. Shape and size of the leaves, their differentiated sizes depending on
position on twig, and angles of branching shoots of the 2nd/1st and 3rd/2nd orders
are consistent with those of the species Pagiophyllum feistmantelii (Halle, 1913a;
Gee, 1989b; Rees & Cleal, 2004).
Twigs of Pagiophyllum maculosum, P. insigne, P. kurrii and P. orinatum,
though generally similar to P. feistmantelii, have leaves larger than those of our
specimen. Our specimen has leaves similar in size to Pagiophyllum fragilis, however in the latter, leaves are always bent at apex (Harris, 1979); that feature is unknown from our specimen.
Gee, 1989b; Rees & Cleal, 2004); Australia: Tannymorel Colliery near Warwick,
and Clifton Colliery, Walloon Coal Measures – Jurassic; India: Jurassic; Argentina
– Middle Jurassic (Townrow, 1967c).
Fig. 51. Pagiophyllum feistmantelii Halle, emend. Townrow, S78/27 KRAM-P. Scale bar 10 mm
83
Pagiophyllum arctowskii sp. nov.
(Fig. 52)
Material. 13 imprints (on 6 rock specimens) of foliated twig fragments: 2 very well
preserved, the remaining ones preserved well or poorer, a part of specimens
pyritized. S78/76, S78/121, S78/133, S81/1, S81/21, 22 KRAM-P.
Name derivation. In honour of Dr Henryk Arctowski, an eminent Polish Antarctic
geologist and geophysicist, scientific leader of the famous Belgian West Antarctic
1897–1899 Expedition in Belgica under captain Adrien de Gerlache de Gomery.
Description. Twigs not branched. Twig fragments 6–35 mm long and 4–7 mm
broad. Axis 1 mm broad. Leaves planispiral, not overlapping, attached at right
angle to twig, triangular in outline, pointed at apex (Fig. 52), 2–5 mm long and
1.5–3 mm broad. Leaves apparently flat, with equally wide depression passing
mid-leaf from its base to its margin near apex (Fig. 52).
Diagnosis. Single twigs with planispiral leaves. Leaves not overlapping, set at right
angle with respect to twig axis. Leaves triangular in outline, with pointed apex, 2–5
mm long and 1.5–3 mm broad. Equally-wide depression passes from leaf base to its
margin, ending near apex.
Holotype. Fig. 52A, S78/76 KRAM-P.
Location and type stratum. Hope Bay, Mount Flora (Antarctic Peninsula, West
Antarctica), Mount Flora Formation, Flora Glacier Member – Figs 5–7 (see
Birkenmajer, 1993a, 2001).
Geological age. Jurassic (?Early Jurassic).
Discussion. Attribution of our specimens to the genus Pagiophyllum is indicated
Fig. 52. Pagiophyllum arctowskii sp. nov. Ociepa. A – holotype, S78/76 KRAM-P; B – S78/121
KRAM-P; C – S78/133 KRAM-P. Scale bar 5 mm
84
by its planispiral leaves, their length being larger than width (Harris, 1979). The
leaves are apparently flat, differing in this feature from the genus Geinitzia, which –
at cross-section – are as thick as wide (Harris, 1979).
Our specimens are generally similar to twigs of Geinitzia divaricata (Harris,
1979, pl. 3, figs 1, 2), however leaves of our specimens are triangular in outline,
while those of G. divaricata and are equally-narrow.
Our specimens resemble also twigs of Glenrosa pagiophylloides (Watson &
Fisher, 1984, figs 1-3): the similarity lies in outline and size of leaves, however
mid-leaf depression in P. arctowskii is equally-narrow, while that of G. pagiophylloides widens near leaf base. The latter feature is well visible in illustration by Fontaine (1893, pl. 42, figs 1A, 2A) who first described this fossil plant under the name
Sequoia pagiophylloides.
Our specimens resemble twigs of P. grantii (Bose & Banerji, 1984, text-fig.
52A, B), however leaves of the latter species are not planispiral; they are slightly
sickle-like and are smaller than those of P. arctowskii.
The features of P. arctowskii which were decisive for creating a new species,
and differ it from other species of the genus Pagiophyllum, were: planispiral
arrangement of non-overlapping leaves, and their triangular outline.
COMPARISON OF THE HOPE BAY FLORA WITH JURASSIC
FLORAS OF GONDWANA AND LAURASIA
Comparison with Gondwana
(1) The Hope Bay Jurassic flora here described is most similar to that of South
America (Tab. 1). Thirteen species are in common for Hope Bay and South America: Archangelskya furcata, Cladophlebis antarctica, C. denticulata, C. grahamii,
Elatocladus confertus, Equisetum laterale, Nilssonia taeniopteroides, Otozamites
linearis, Pachypteris cf. indica, Pagiophyllum feistmantelii, Sagenopteris nilssoniana, Zamites antarcticus and Z. pusillus.
(2) Seven species are in common for Hope Bay and India (Tab. 1): Cladophlebis
denticulata, Coniopteris cf. hymenophylloides, C. lobata, Elatocladus confertus,
Komlopteris indica, Pachypteris cf. indica and Pagiophyllum feistmantelii.
(3) Seven species are in common for Hope Bay and Australia (Tab. 1): Cladophlebis denticulata, Coniopteris cf. hymenophylloides, Elatocladus confertus, Equisetum laterale, Pachypteris cf. indica, Pagiophyllum feistmantelii and Sphenopteris pecten.
(4) Six species are in common for Hope Bay and New Zealand (Tab. 1): Cladophlebis antarctica, C. denticulata, Coniopteris cf. hymenophylloides, C. lobata,
Elatocladus confertus and Equisetum laterale.
(5) Only three species are in common for Hope Bay and Africa (Tab. 1): Coniopteris cf. simplex, Cladophlebis denticulata and Zamites antarcticus. Of these, only
one species (Z. antarcticus) is in common for Hope Bay and South Africa.
85
Table 1
Occurrence of Jurassic species from Hope Bay on different continents
Gondwana
Laurasia
India
Australia
New
Zealand
Asia
(India
North
Europe
exclusiAmerica
vely)
Coniopteris cf.
hymenophylloides
+
+
+
+
Coniopteris lobata
+
+
+
Species
South
America
Africa
Araucarites antarcticus
Archangelskya furcata
+
Coniopteris murrayana
+
Coniopteris cf. simplex
+
Cladophlebis antarctica
+
Cladophlebis denticulata
+
Cladophlebis grahamii
+
Elatocladus confertus
+
+
+
+
+
+
+
+
+
Komlopteris indica
Nilssonia taeniopteroides
+
+
+
+
+
+
+
+
+
+
+
Equisetum cf. columnare
Equisetum laterale
+
+
+
+
+
+
+
+
+
Otozamites gramineus
Otozamites linearis
+
Pachypteris cf. indica
+
+
+
Pagiophyllum feistmantelii
+
+
+
Sagenopteris nilssoniana
+
+
+
+
+
+
Sphenopteris antarctica
Sphenopteris pecten
Zamites antarcticus
+
+
+
Zamites pachyphyllus
Zamites pusillus
+
Comparison with Laurasia
(1) The Hope Bay Jurassic flora has eight species in common with Europe (Tab.
1): Cladophlebis denticulata, Coniopteris cf. hymenophylloides, C. murrayana, C.
cf. simplex, Equisetum cf. columnare, E. laterale, Otozamites gramineus and
Sagenopteris nilssoniana.
(2) The Hope Bay Jurassic flora has five species in common with North America
(Tab. 1): Cladophlebis denticulata, Coniopteris cf. hymenophylloides, C. lobata,
Equisetum laterale and Sagenopteris nilssoniana.
86
(3) Eight species are in common for Hope Bay and Asia (India exclusively) –
Tab. 1: Cladophlebis denticulata, Coniopteris cf. hymenophylloides, C. lobata, C.
murrayana, Elatocladus confertus, Equisetum cf. columnare, E. laterale, Nilssonia taeniopteroides, Otozamites gramineus and Sagenopteris nilssoniana.
Exclusively Antarctic and new species
(1) Five species of the Hope Bay flora: Araucarites antarcticus, Schizolepidiella gracilis, S. birkenmajeri n. sp. (Ociepa, 2007), Sphenopteris antarctica and
Zamites pachyphyllus have so-far been described from Antarctica only. (2) Two
new vascular plants here described include: Crossozamia mirabilis n. sp. and
Pagiophyllum arctowskii n. sp.
Remarks
The above data (Tab. 1) indicate that the Hope Bay Jurassic flora represents a
decidedly Southern Hemisphere one, in spite of having considerably numerous
species in common with Laurasia. This is a further proof for cosmopolitic character
of Jurassic floras.
From Laurasian Jurassic floras, the Hope Bay flora, besides the species known
only from Gondwana, differs in lacking representatives of the families Czekanowskiaceae, early Pinaceae, and Ginkgoaceae. Lack of Czekanowskiaceae and early
Pinaceae is characteristic for Jurassic floras of Southern Hemisphere (see Vakhrameev, 1991). In that hemisphere, Ginkgoaceae are rare. They seem to be missing
from Antarctica, perhaps with the only exception from Lassiter Coast and Behrendt
Mountains in Antarctic Peninsula, as based on a short note by J. M. Schopf (see
Laudon et al., 1983), however not supplied with palaeobotanic description and
illustration.
AGE OF THE HOPE BAY FLORA BASED ON FOSSIL PLANTS
Stratigraphic age ranges of fossil plants. Stratigraphic age ranges of the Hope
Bay flora indicate its Jurassic age:
(1) Of 25 species here described, nine are known from the Jurassic strata of West
Antarctica and elsewhere: Archangelskya furcata, Cladophlebis grahamii, Coniopteris murrayana, C. cf. simplex, Nilssonia taeniopteroides, Otozamites
gramineus, O. linearis, Pagiophyllum feistmantelli and Zamites pusillus;
(2) Five species are so-far known only from Jurassic strata at Hope Bay, West
Antarctica: Araucarites antarcticus, Schizolepidella gracilis, S. birkenmajeri (see
Ociepa, 2007), Sphenopteris antarctica and Zamites pachyphyllus;
(3) Two new species here described: Crossozamia mirabilis n. sp., and Pagiophyllum arctowskii n.sp., are so-far known only from Jurassic strata at Hope Bay,
West Antarctica;
(4) Five species range from Jurassic through Cretaceous: Cladophlebis antarc-
87
tica, Coniopteris cf. hymenophylloides, Elatocladus confertus, Komlopteris indica, and Pachypteris cf. indica;
(5) Four species are known from Triassic through Cretaceous inclusively: Cladophlebis denticulata, Coniopteris lobata, Equisetum cf. columnare and Equisetum laterale;
(6) Two species range from Triassic trough Jurassic: Sagenopteris nilssoniana
and Sphenopteris pecten. The genus Stachyotaxus (Rao, 1964) is known from Triassic through Jurassic strata.
Age conclusions. In his last age estimation, based on fossil plants, Rees (1993a)
suggested an Early Jurassic age of the Hope Bay flora. This seems to be supported,
i.a., by the presence of the genus Goeppertella known so-far from the Triassic and
Early Jurassic strata. A probably coeval flora from Botany Bay, Antarctic Peninsula, yielded Dicroidium feistmantelii: the genus Dicroidium has been reported sofar from Triassic deposits only.
The collection elaborated in this paper supports a Jurassic age of the Hope Bay
plant fossils, however without closer specification as to whether they represent
Early or Middle Jurassic stages.
PALAEOCLIMATIC ASPECTS OF THE HOPE BAY JURASSIC FLORA
Palaeoclimatic analysis based on worldwide distribution of 782 species of Jurassic plants has lead Rees et al. (2000, fig. 7A) to the conclusion that during Early
Jurassic, a warm-temperate climate reigned in the Antarctic Peninsula area. The
plant fossil collection here described allows to add some data as to climatic preferences of selected plant genera and families considered:
(1) The family Dicksoniaceae is presently known first of all from the tropics, its
worldwide distribution includes, however, also warm-temperate zones (Kramer &
Green, 1990); (2) The genus Pachypteris, a pteridosperm, is considered to be an index genus for subtropical climates lacking significant yearly temperature variations (Vakhrameev, 1991). The same can be said of Bennettitales, represented by
the genera Otozamites and Zamites (op. cit.);
(3) The cycads presently occur mainly in tropical and subtropical, more seldom
in warm-temperate areas (Jones, 1998). Generally, they are considered indices of
subtropical zone (Vakhrameev, 1991), however the genus Nilssonia – known from
Hope Bay flora – might range also to temperate zone (op. cit.);
(4) Occurrence of conifers of the Araucariaceae family in Jurassic and Cretaceous strata, is considered an evidence for a warm and stable climate without much
seasonal variation (Abbink, 1998). The present-day Araucariaceae occur first of all
in tropical and subtropical areas, less frequently in warm-temperate zone (Kramer
& Green, 1990);
(5) The present-day conifers of the Taxodiaceae family occur in warmtemperate zone (Kramer & Green, 1990);
(6) The conifers of the Podocarpaceae family are not represented in the investigated collections from Hope Bay. However, some authors (e.g., Florin, 1940; Can-
88
trill & Falcon-Lang, 2001) supposed that to this family might belong representatives of the genus Elatocladus, its twigs being frequent in our collection. Presently,
representatives of the family Podocarpaceae occur mainly in tropical and subtropical zones, however, there are also known species growing in warm-temperate and,
sporadically, even in cool-temperate zones (Kramer & Green, 1990).
The above data indicates that Jurassic palaeoclimate of the Antarctic Peninsula
area was most probably warm and stable, devoid of significant temperature variations. The Hope Bay flora lacks taxons known exclusively from the tropics, while
occurrence of representatives of the family Taxodiaceae, presently characteristic
for temperate climatic zone, allows to exclude a tropical palaeoclimate.
It is suggested that the Jurassic Hope Bay flora grew in a climatic zone transitive
between the subtropical and the warm-temperate ones. This conclusion is in agreement with palaeoclimatic reconstruction of the Jurassic period as presented by
many authors (e.g., Weissert & Mohr, 1996; Hallam, 1993; Ziegler et al., 1933;
Rees et al., 2000; Sellwood et al., 2000). Break-up of the large supercontinent Gondwanaland which commenced at the beginning of Jurassic and continued since then
to form separate continents divided by ever growing oceanic basins, probably was a
pre-requisite for appearance of monsoons (Parrish, 1990; Hallam, 1990; Rees et al.,
2000). The monsoon circulation in the Antarctic Peninsula area would produce intense precipitation during summer season, which disappeared during winter season
resulting in droughts (Rees et al., 2000).
PALAEOENVIRONMENTAL ASPECTS OF THE HOPE BAY
JURASSIC FLORA
During the Jurassic, the Hope Bay area was part of mountainous area of the Antarctandes foldbelt bordering on a wide expanse of the Pacific Ocean (see Birkenmajer, 1993a, 1994a). At least two types of palaeoenvironment might be distinguished there: A palaeoenvironment of meandering river valley with its wet floodplain; A palaeoenvironment of more dry mountain slopes.
The meandering fiver-floodplain environment. This environment was inhabited by representatives of the horsetails of the genus Equisetum, characteristic for
such environment (cf. Abbink, 1998), accompanied by ferns of the genus Cladophlebis. These ferns had very large leaves (cf. Deng, 2002). E.g., in Jurassic coalbeds of Romania, the species C. denticulata was the main coal-forming plant (van
Konijnenburg-van Cittert, 2002).
The small pteridosperm tree Sagenopteris nilssoniana (cf. Taylor & Taylor,
1993) grew on wet river banks (cf. Vakhrameev, 1991; Abbink, 1998), as also did
the cycad Nilssonia taeniopteroides (Vakhrameev, 1991).
The mountain slope environment. The mountain slopes were probably overgrown by two types of forests:
(i) A high forest consisting of trees belonging to the families Araucariaceae and
Taxodiaceae, and of representatives of the genus Elatocladus;
(ii) A low forest consisting of representatives of the Bennettitales. The Aracau-
89
riaceae and Taxodiaceae on the one side, and the Bennettitales on another, are considered to be characteristic for rather dry habitats (cf. Vakhrameev, 1991). Araucariaceae and Taxodiaceae are presently known also from the mountains (Kramer
& Green, 1990).
The present-day Podocarpaceae grow mainly in mountainous areas (Kramer &
Green, 1990); their probable representative in our collection could be Elatocladus
confertus.
The pteridosperm Archangyelskia furcata, probably a liana (Rees & Cleal,
1993), could be associated with taller trees.
The lowest tier of our forest was occupied by ferns of the Dicksoniaceae family.
Presently, representatives of this family are tree-ferns that often occupy lowest forest tier in mountaineous areas (Kramer & Green, 1990). During Jurassic, the Dicksoniaceae, with the exception of the genus Coniopteris – a small fern with groundcreeping rhizome (Deng, 2002) – could also live under similar conditions.
The genus Pachypteris, a small shrub (Abbink, 1998), is considered to be a representative of mangrove-like habitats (Vakhrameev, 1991); it could suggest a nearshore sedimentary environment of the Mount Flora Formation. On the other hand,
lack in our collections of representatives of the Cheirolepidiaceae family speaks
against ultra-dry habitats (op. cit.).
Present-day equivalents of the Hope Bay flora. Looking for contemporaneous plant formation equivalent to that from the Jurassic Hope Bay, we may point to
evergreen forests of temperate zone, such as those growing in East Asia (southern
Japan, Korea, China), which gradually pass into subtropical ones. Here belong also
deciduous-conifer forests which in New Zealand include Agathis australis from the
Araucariaceae family, the araucarian forests in south Brazil, and wet conifer forests
of California with Sequoia sempervirens – a representative of the Taxodiaceae
family.
Contemporaneous evergreen rain forest of temperate zone – the Valdivian forest
(Walter, 1976) growing in the Andes, may also resemble to some extent the plant
assemblage of the Mount Flora Formation. The Valdivian forest is a rich one, containing – besides flowering plants and representatives of the Cupressaceae family,
which do not occur in the Mount Flora assemblage – numerous Araucariaceae and
Podocarpaceae (op. cit.). This feature indicates a new trend in the West Antarctic
palaeofloras, expressed already in the Late Cretaceous ones which resemble the
Valdivian forest (Zastawniak, 1998).
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