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Paleocene flora from Seymour Island,

Antarctica: revision of Dusén's (1908)
pteridophyte and conifer taxa
a b c b
David J. Cantrill , Anne-Marie P. Tosolini & Jane E. Francis
a
National Herbarium of Victoria, Royal Botanic Gardens
Melbourne, Private Bag 2000, Birdwood Ave, South Yarra, VIC,
3141, Australia
b
Department of Earth Sciences, University of Leeds, Leeds, LS2
9JT, UK
c
Environmental Geoscience, Latrobe University, VIC, 3086,
Australia
Published online: 13 May 2011.

To cite this article: David J. Cantrill , Anne-Marie P. Tosolini & Jane E. Francis (2011): Paleocene
flora from Seymour Island, Antarctica: revision of Dusén's (1908) pteridophyte and conifer taxa,
Alcheringa: An Australasian Journal of Palaeontology, 35:2, 309-328

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 Paleocene flora from Seymour Island, Antarctica:
revision of Dusén’s (1908) pteridophyte
and conifer taxa
DAVID J. CANTRILL, ANNE-MARIE P. TOSOLINI AND JANE E. FRANCIS

CANTRILL, D.J., TOSOLINI, A-M.P. & FRANCIS, J.E., June, 2011. Paleocene flora from Seymour Island, Antarctica:
revision of Dusén’s (1908) pteridophyte and conifer taxa. Alcheringa 35, 309–328. ISSN 0311-5518.

The Paleocene flora from Seymour Island, Antarctica, first collected by Nordenskjöld on the Swedish South Polar
Expedition (1901–1903), was described by Dusén (1908) as having 87 leaf taxa making it one of the most diverse
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floras of this age in the Southern Hemisphere. The original descriptions of these leaf impressions included 37
pteridophytes and one conifer. Many species identified by Dusén were based on single fragmentary specimens. Major
new collections housed at the British Antarctic Survey (Cambridge, UK), together with the original collections held
at The Swedish Museum of Natural History, form the basis for a taxonomic revision of the flora. This paper
concentrates on the pteridophytes and conifers. The diversity is considerably reduced from Dusén’s estimates with
just three ferns (Cladophlebis aemulans, Cladophlebis seymourensis, Sphenopteris angustiloba) and two conifers
(Araucaria imponens, Elatocladus seymourensis) recognized.

David Cantrill [[email protected]], National Herbarium of Victoria, Royal Botanic Gardens

Melbourne, Private Bag 2000, Birdwood Ave, South Yarra, VIC 3141, Australia; Anne-Marie Tosolini*
[[email protected]], Department of Earth Sciences, University of Leeds, Leeds LS2 9JT, UK & Environmental
Geoscience, Latrobe University, VIC, 3086, Australia; Jane Francis [[email protected]], Department of
Earth Sciences, University of Leeds, Leeds LS2 9JT, UK. *Present address: School of Earth Sciences, The University of
Melbourne, VIC 3010, Australia. Received 21.2.2010, revised 15.2.2011, accepted 18.2.2011.

Key words: Paleogene, Antarctica, Antarctic Peninsula, Gondwana, polar, ferns, conifers, palaeobotany.

THE SWEDISH SOUTH POLAR EXPE- 2005). In contrast, the Paleocene flora from
DITION (1901–1903) led by Otto Nor- Seymour Island (Dusén 1908; Fig. 1) has
denskjöld discovered the first fossil leaf not been re-examined in any detail since it
floras in Antarctica of Jurassic (Hope Bay) was first described. This can be attributed in
and Late Cretaceous to Eocene (Snow Hill part to the difficulty of access to Seymour
and Seymour islands) age (Fig. 1). The Island compared with Hope Bay, but also to
Jurassic flora from Hope Bay (Halle 1913) the lack of documentation about the site
has received considerable attention (e.g. (Zinsmeister 1988).
Stipancic & Bonnetti 1970, Gee 1989, Seymour Island, situated in the Weddell
Birkenmajer 1993, Rees 1993a, b, Rees & Sea to the east of the Antarctic Peninsula,
Cleal 1993, 2004) owing to its importance in comprises deposits of Cretaceous to ?Oligo-
Southern Hemisphere Jurassic macrofloral cene age and is one of the richest sources of
biostratigraphy, and the considerable de- fossil material in the Antarctic. Although
bate over the age of the flora (Hunter et al. there is a considerable body of research into
the animal fossils, interest in the plant
fossils was only renewed in the 1980s. Fossil
ISSN 0311-5518 (print)/ISSN 1752-0754 (online)
Ó 2011 Association of Australasian Palaeontologists wood collected from Seymour Island was
DOI: 10.1080/03115518.2011.565214 initially described by Gothan (1908) with
 310 DAVID J. CANTRILL et al. ALCHERINGA
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Fig. 1. Locality map. A, Map to show position of Seymour Island in relation to Antarctic Peninsula.
B, Detail showing geology and sample localities on Seymour Island. C, Geological overview of the main
formations.
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 311

recent additions by Torres (1985, 1990), The Paleocene flora described by Dusén
Francis (1986, 1991) and Torres et al. (1908) comes from the central part of
(1994a, b). More recently, a joint collabora- Seymour Island and is preserved in the
tion between the British Antarctic Survey Cross Valley Formation. Dusén (1908)
and Leeds University has seen significant described and named 37 leaf taxa, mainly
revisions and additions to the wood flora angiosperms but also including ferns and
(Poole & Gottwald 2001, Poole et al. 2000a, conifers. He also identified a further 50
b, 2003, Francis & Poole 2002, Cantrill & species illustrating these on the plates with-
Poole 2005a). Palynology, initially by Cran- out providing formal descriptions. The
well (1959), also saw a resurgence of interest combined total of 87 taxa makes the
in the 1980s and early 1990s (Askin & Seymour Island flora one of the most
Fleming 1982, Truswell 1983, Baldoni & diverse Paleocene floras in the Southern
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Barreda 1986, Dettmann & Thomson 1987, Hemisphere. Recent discoveries and re-
Zamaloa et al. 1987, Askin 1988a, b, 1989, examination of Paleogene localities in South
1990, 1992, 1994, 1997, Askin et al. 1991, America have revealed high levels of diver-
Jarzen & Dettmann 1991). Plant macro- sity compared with the Northern Hemi-
floras, although sparse, continued to be sphere (Iglesias et al. 2007), and Eocene
documented through this period but largely floras from Patagonia such as Laguna del
from the Eocene (Doktor et al. 1996, Hunco have exceptional diversity (Wilf
Gandolfo et al. 1998a, b, Upchurch & et al. 2003, 2005). Understanding the origins
Askin 1989) with the species described by of this diversity and how the vegetation
Dusén (1908) being included in wider evolved following the Cretaceous/Paleogene
revisions (e.g. Romero 1986, Tanai 1986, extinction remains a significant challenge
Gonzalez et al. 2007). Some of these papers that is limited by our current knowledge of
formed the basis for overall reviews of the the Antarctic floras. Reviews of Antarctic
Cretaceous and Paleogene vegetation his- wood floras through the Late Cretaceous
tory of the Antarctic (Dettmann & Jarzen and Paleogene suggest that some South
1988, 1990, Dettmann 1989, Dettmann et al. American diversity (e.g. Valdivian rainfor-
1990, Truswell 1991, Askin & Spicer 1995, est) had its origins in the Antarctic (Cantrill
Hill & Scriven 1995, Cantrill & Poole 2002). & Poole 2005a, Poole & Cantrill 2006).
Throughout this period of rediscovery, However, evidence from other sources such
the Paleocene fossil leaf site described by as leaf floras is yet to be explored. Indeed, as
Dusén remained forgotten. A review of Truswell (1991, p. 520) noted, the most
conifers from the Southern Hemisphere diverse floras from the Antarctic Peninsula
(Florin 1940) re-evaluated the conifers are from Seymour Island but, in order to
described by Dusén (1908), confirmed the determine how realistic the taxonomic
presence of the Araucariaceae (Araucaria assessments are, a ‘critical re-evaluation in
imponens) and reassigned other material to terms of leaf architecture’ is required. This
Acmopyle and Podocarpus sect. Stachycar- message has been reiterated by Gonzalez
pus (now Prumnopitys). The only other et al. (2007, p. 260) who pointed out that
research was based on a study of small ‘new specimens with better preservation are
new leaf collections that described Nothofa- required to confirm the affinities of the
gus, and one new unidentified leaf (Case Antarctic taxa described by Dusén with
1988). More recently, climatic analysis of extant Lomatia and also with Proteaceae.’
the flora has provided an overview of Fieldwork between January and March
angiosperm elements (Francis et al. 2006, 2001 allowed us to make a large collection
2008). from the original area where the material
 312 DAVID J. CANTRILL et al. ALCHERINGA

described and illustrated by Dusén was with it. These sandstones are interpreted as
collected. Together with the existing collec- estuarine/shallow marine deposits of a
tions, over 1000 specimens were examined back-arc basin. Leaves represented in the
allowing us to make a more rigorous study Cross Valley Formation were derived from
of the morphological diversity present, and vegetation growing on the adjacent active
to better constrain the taxonomic bound- arc, and were subsequently transported,
aries erected by Dusén (1908). Dusén (1908) sorted and buried in nearshore sandstones
recorded 87 taxa in the Paleocene flora of of the back-arc basin.
Seymour Island, Antarctica, of which 37 The age of the Cross Valley Formation is
were pteridophytes and 3 conifers. Here we problematic; it contains few age-diagnostic
present a taxonomic revision of the pterdio- fossils and is bounded by erosional surfaces.
phytes and conifers described by Dusén The uppermost beds of underlying Sobral
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(1908) and a later paper will revise the Formation are dominated by Spinidinium
angiosperm component. sp. cf. S. lanternum, which, together with an
absence of Palaeoperidinium pyrophorum
and Trithyrodinium evittii, suggests a late
Geological setting and age early Paleocene age (Askin 1988a, Huber
Lithospheric extension and subsequent rifting 1988, Mohr 1990, Willumsen & Vajda 2010,
during the early phases of Gondwana break Willumsen 2011). The overlying La Meseta
up provided the tectonic mechanisms that Formation contains a rich palynoflora and
formed the Larsen Basin to the east of the the basal units can be assigned to palyno-
Antarctic Peninsula (Hathway 2000, Fig. 1). morph Zone 7 (predominance of Enigmadi-
This basin contains a series of megase- nium cylindrifloriferum) of early Eocene age
quences, of which the Early Cretaceous (Askin 1988a, 1990, Askin et al. 1991).
(Aptian) to Paleogene (?Oligocene) succes- These two units bracket the Cross Valley
sion in the James Ross Island region forms Formation, which is thought to be of early
the uppermost cycle (Hathway 2000). Three late Paleocene age (Wrenn & Hart 1988).
distinct groups are recognized within this This determination is supported by recent
package of sediments: the Gustav (Aptian to palynological work on the area (Greenhalgh
Coniancian/Santonian), Marambio (Santo- 2002). Fossil wood collected by the initial
nian to Paleocene) and Seymour Island Swedish South Polar Expedition (Gothan
(Paleocene to ?Oligocene) groups (Rinaldi 1908), together with recent collections (e.g.
et al. 1978, Elliot & Trautman 1982, Zinsme- Torres 1990, Poole et al. 2000b, Poole &
ister 1982, Elliot 1988, Macellari 1988, Sadler Gottwald 2001), are not diagnostic for age.
1988, Zinsmeister et al. 1989, Crame et al.
1991, Pirrie et al. 1997, Marenssi et al. 1998,
Porebski 1995, 2000). Material and methods
Leaf impressions occur in sandstones of The fossil flora locality (648160 1.9?S
the Cross Valley Formation, Seymour 568400 2.6?W), crops out over an area of
Island Group. The host sandstones are approximately 2 ha as a series of small (less
exposed on the hills to the south-southeast than 2 m high) hillocks of more or less
of Cross Valley that overlook the Weddell bedding parallel exposure. This makes
Sea. These can be referred to unit Tpcv 5 estimations of the stratigraphic thickness
(Silver Shales) of the Cross Valley Forma- difficult but it is thought to be less than
tion (Sadler 1988), a unit that has in places 10 m thick. The locality is situated 3.5 km
thin lenses of La Meseta Formation un- southwest of the Base Vicecomodoro Mar-
conformably overlying or in faulted contact ambio on Seymour Island (Fig. 1) within
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 313

sediments of the early late Paleocene Cross Family ?OSMUNDACEAE Berchtold & J.
Valley Formation. Original material col- Presl, 1820
lected by Nordenskjöld on the 1901–1903
Swedish South Polar Expedition (Dusén Cladophlebis Brongniart, 1849
1908) was examined in the Swedish Museum
of Natural History (prefix S). The collection Type species. Cladophlebis albertii Brong-
from the Swedish South Polar expedition is niart 1849
rather fragmentary with many of the
morphotypes recognized being represented Cladophlebis aemulans (Dusén) comb. nov.
by just one specimen; thus, making it (Fig. 2E–H, J)
difficult to evaluate morphological variation
in order to validate taxonomic boundaries. 1908 Polypodium aemulans Dusén, p. 12,
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Of the original 126 catalogued specimens pl. 3, fig. 10.

only 83 were available for study as some 1908 Taeniopteris blechnoides Dusén, p. 13,
appear to have been misplaced. pl. 3, figs 23, 24.
Other collections examined include ma- 1908 Taeniopteris crispissima Dusén, p. 14,
terial held in the Geological Collections of pl. 3, figs 19, 20.
the British Antarctic Survey (BAS). Materi- 1908 Alsophila antarctica Christ., Dusén,
al from this locality includes collections p. 14, p1. 3, fig. 11.
made by the Falkland Island Dependencies
Survey (FIDS), the forerunner to the BAS, Lectotype. Here designated as S132809_02
in 1944 and prefixed D512, 517, 523. In held in the Palaeobotany Collection of the
addition, material collected during recon- Swedish Museum of Natural History and
naissance work by Imogen Poole, DJC and illustrated by Dusén (1908 on pl. 3, fig. 10)
JEF in 1998–1999 prefixed DJ913 was and here as Fig. 2E.
examined. Fieldwork in January to March
2001 provided the opportunity to make Material examined. Five specimens as fol-
large collections from this locality. Three lows: S132809_02, S132834, S132845a,
separate localities from different parts of the S132845b, S132873b.
outcrop and from slightly different strati-
graphic levels were collected (DJ1111, Description. Large isolated pinnules that
DJ1112 and DJ1113; Fig. 1). range up to 57 mm long by 11 mm wide.
All specimens were examined with an Pinnules lanceolate to linear lanceolate with
Olympus SZH-10 stereo microscope and an acute apex (Fig. 2E). Venation comprises
photographed using Kodak Technical Pan a single midvein that persists to the pinnule
film. Negatives were scanned using a Nikon apex. Secondary veins arise acutely and arch
super CoolScan 9000 ED. Digital images of towards the pinnule margin branching once
macrospecimens were obtained with a Ni- near the midvein (Fig. 2E, J) or rarely twice.
kon D1 using low-angle incident lighting Six to eight veins per 10 mm of pinnule
from different directions to best enhance margin, pinnule margins entire and possibly
leaf venation features. Plates were compiled enrolled.
in Adobe Photoshop CS2.
Remarks. Dusén (1908) described four
specimens that he assigned to four taxa:
Systematic palaeontology Polypodium aemulans Dusén, Taeniopteris
Phylum PTERIDOPHYTA Schimper, 1879 blechnoides Dusén, T. crispissima Dusén and
Order ?OSMUNDALES Link, 1833 Alsophila antarctica Dusén. Each specimen
Downloaded by [Fudan University] at 10:48 29 April 2013

314
DAVID J. CANTRILL et al.
ALCHERINGA
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 315

is an isolated pinnule in varying states of are known. Taeniopteris is also not suita-
preservation. The specimen described as P. ble for several reasons. Taeniopteris vittata
aemulans (Fig. 2E) is a single pinnule with is the type of the genus, a taxon that has
the apex and base preserved. The base been confused with Nilssoniopteris (Cleal
suggests that the lamina had a flange that et al. 2006). However, in the Southern
extends onto the rhachis. The specimen Hemisphere, this genus has been widely
described as Taeniopteris blechnoides is the applied to leaves of the Pentoxylales
mid-portion of a pinnule that is 45 mm long (Howe & Cantrill 2001). The Southern
with the margins poorly preserved and Hemisphere Taeniopteris also differ from
somewhat undulate. The specimen de- the material described here, in that they
scribed as T. crispissima is similar to T. are simple, shortly petiolate, leaves with a
blechnoides but the margins are broken and multistranded midvein and secondary
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strongly undulate (Fig. 2F, J). The final veins arising almost at right angles and
specimen described as A. antarctica is also generally occurring in higher densities.
incomplete but larger. This specimen has Fronds with pinnules that have a
some distinctive undulations either side of midvein, with secondary veins that openly
the midvein at the base of the pinnule (Fig. dichotomize, and arise more or less
2H) that were also illustrated by Dusén acutely, gently arching towards the
(1908), and might represent the attachment margin are placed in Cladophlebis (e.g.
points for sporangia. The case for separate Harris 1961). The pinnules described
taxa is poor given the range of pinnule here conform to these features and we
variation that occurs on a single frond in reassign this taxon to Cladophlebis aemu-
extant ferns. Polypodium aemulans could lans comb. nov. To date several hundred
easily be an apical pinnule, whereas the fossil taxa have been placed within Cla-
others are more typical of pinnules in the dophlebis, a genus that ranges from the
mid-part of a larger frond. The venation Triassic through the Cenozoic. Although
patterns are the same in all specimens both Cladophlebis has been used for sterile
in terms of vein density and pattern and foliage only, where foliage is fertile it
we regarded these four specimens as con- generally indicates affinity with the Os-
specific. mundaceae (e.g. Todites species described
The taxon was first described by Dusén by Harris 1961; Todites grahamii, Gee
(1908) as Polypodium aemulans and we 1989). However, no fertile material is
designate this as the basionym. Dusén available among the Seymour Island speci-
(1908) assigned the material to a range of mens so suprageneric placement remains
extant (Polypodium, Alsophila) and fossil uncertain. No further material assignable
genera (Taeniopteris). There is no strong to C. aemulans was present in the collec-
evidence to support placement within tions we examined, and this appears to
either Polypodium or Alsophila as neither be an extremely rare taxon within the
the sporangia nor the frond morphology deposit.

3
Fig. 2. Sphenopteris angustiloba (Dusén) comb. nov. and Cladophlebis aemulans (Dusén) comb. nov. from the Cross
Valley Formation. A–D, I, Sphenopteris angustiloba. A, Large pinna fragment, D523.1c; B, Apical portion of primary
pinna, note ribbed rhachilla, D512.1a; C, Pinna with inrolled pinnule margins, S132833; D, Terminal portion of
pinna, DJ913.19a; I, Details of poorly preserved secondary venation, S132833. E–H, J, Cladophlebis aemulans. E,
Isolated pinnule with tapering apex S132809_02; F, Pinnule with undulate margins, S132834; G, Pinnule fragment,
S132837b; H, Details of venation, S132837b; J, Details of venation from base of pinnule, S132809_02. Scale
bar ¼ 10 mm.
Downloaded by [Fudan University] at 10:48 29 April 2013

316
DAVID J. CANTRILL et al.
ALCHERINGA
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 317

Cladophlebis seymourensis (Dusén) comb. upper surface (Fig. 3B). Venation compris-
nov. (Fig. 3A–L) ing a single midvein, secondary veins arise
acutely and arch towards the margin,
1908 Polypodium nathorstii Dusén, p. 12, pl. dichotomizing sparsely (Fig. 3H, K).
4, figs 8–10. Pinnule margin generally entire (Fig. 3I, L)
1908 Pecopteris sp. 1 Dusén, p. 19, pl. 4, fig. 5. but can be lobed in terminal pinnules (Fig.
1908 Dryopteris antarctica Dusén, p. 15, pl. 3F).
4, fig. 6.
Remarks. Few of Dusén’s (1908) taxa are
1908 Dryopteris seymourensis Dusén, p. 15,
based on multiple specimens. Within the
pl. 4, figs 4, 19.
ferns he ascribed three specimens to Poly-
1908 Pecopteris sp. 7; Dusén, p. 19, pl. 4, fig.
podium nathorstii, two specimens to Dryop-
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18.
teris seymourensis, all other taxa were based
1908 Pecopteris sp. 2; Dusén, p. 19, pl. 4, fig. on single specimens and many represent
20. small fragments of pinnae or single pin-
nules. Within the collection in Stockholm,
Lectotype. Here designated as S132855 several specimens were on loan and un-
held in the Palaeobotany Collection of available for study, whereas others could
the Swedish Museum of Natural History not be located (either missing from the
and illustrated by Dusén (1908 on pl. 3, collection or not marked on the specimens).
fig. 19) and here as Fig. 3I. The lack of large intact fronds makes it
difficult to draw firm conclusions about
Material examined. Twenty-one specimens the number of taxa, but it is clear that
as follows: D512.1, D512.2, D512.4a, many taxa described by Dusén (1908)
D512.4b, D512.5, D523.1a, D523.1b, overlap considerably in pinnule morphol-
DJ913.2a, DJ913.2b, DJ913.2c, DJ913.11c, ogy and arrangement. The material as-
DJ913.196c, DJ1113.134, DJ1113.196, cribed to Dryopteris seymourensis, D.
DJ113.196a, S132818, S132819, S132824, antarctica, Polypodium nathorstii and Pe-
S122019_01, S132855, S132866. copteris sp. 1, 2 and 7 all have several
consistent features including: alternate
Description. Fronds at least once pinnate, pinnule arrangement, similar pinnule in-
pinnae alternate (Dusén 1908, pl. 4, fig. 5), at sertion angles, size, apparently concave
least 40 mm long by 10 mm wide, bearing nature of the pinnules and, where pre-
alternately small ovate to lanceolate pin- served, the venation pattern. Few signifi-
nules up to 13 mm long and 4 mm wide (Fig. cant differences exist. Some pinnules
3A–E), commonly with a decurrent flange appear to be relatively narrow (e.g. Fig.
extending down the rhachis from the basi- 3C, D) but this may be preservational, as
scopic margin of the pinnules (Fig. 3G, I, L). in these specimens the pinnules appear
Rhachis apparently four-angled with a dis- inrolled and the margins damaged. We
tinct ridge and pinnules inserted on the conclude that the range of variation seen

3
Fig. 3. Cladophlebis seymourensis (Dusén) comb. nov. illustrating the range in pinnule morphology. A, D523.1a,
terminal pinna with slightly lobed pinnule margins. B, Counterpart showing the recurved margins. C, D, Apparent
narrow pinnule form due to inrolling of the pinnules during preservation. C, DJ913.196c D, DJ1113.196a. E, Typical
widespread form S132866. F, Terminal pinna, D512.1. G, Portion of pinna with alternate pinnules, D512.5. H,
Enlargment of pinnule illustrating single midvein and faint dichotmizing secondaries S132866. I, S132855. J, Small
leafed form DJ1113.134. K, Details of venation in D512.5. L, D512.4b. Scale bars ¼ 10 mm.
 318 DAVID J. CANTRILL et al. ALCHERINGA

between these taxa is encompassed by that Sphenopteris angustiloba (Dusén) comb.

evident on a single dissected frond and nov. (Fig. 2A–D, I)
consequently synonomize those taxa that
could be examined into a single taxon. 1908 Lomatia angustiloba Dusén, p. 6, pl. 2,
Dusén used three names at specific figs 3, 4.
rank (Dryopteris seymourensis, D. antarc- 1908 Lomatia brevipinna Dusén, pp. 6–7,
tica, Polypodium nathorstii) for this mate- pl. 2, fig. 7.
rial, placing them in extant genera. The 1908 Asplenium antarcticum Dusén, p. 13,
lack of fertile material and anatomical pl. 4, figs 1–3, 13.
information makes it difficult to reconcile
1908 Asplenium sp.; Dusén, p. 13, pl. 4, fig.
the placement of this fossil material within
26.
extant genera. The placement of fossil
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material into extant genera based on 1908 Sphaenopteris sp. 1; Dusén, p. 19, pl. 4,
limited information was common practice fig. 15.
in the late 1800s and early 1900s. How-
ever, this can be misleading if accepted Lectotype. Here designated as S132833 held
uncritically, and it would be far better to in the Palaeobotany Collection of the
place the material within a fossil morpho- Swedish Museum of Natural History and
genus. The most appropriate genus based illustrated by Dusén (1908 pl. 2, figs 3, 4)
on the pinnule morphology is Cladophle- and here as Fig. 2C.
bis. Of the three names used by Dusén
(1908), the specific epithets for Dryopteris Material examined. Eighteen specimens as
antarctica and Polypodium nathorstii have follows: D512.1a, 523.1c, 523.1d, DJ913.1,
already been used within Cladophlebis. DJ913.11a, DJ913.11b, DJ913.18 DJ
Cladophlebis nathorstii Yokoyama 1894 913.19a, S132811, S132827_02, S132829,
was described by Nathorst (1890) and S132833, S132854_02, S132860, S132862,
Yokoyama (1894) from several localities S132870, S132872, S132874.
in Japan. Cladophlebis antarctica was used
by Halle for Jurassic material from Hope Description. Fronds at least once pinnate
Bay. This leaves only Dryopteris seymour- bearing alternate (rarely sub-opposite) pin-
ensis available for use, and on this basis nae up to 23 mm long and 4.7 mm wide
we make a new combination for the (Fig. 2A). Pinnae spaced from 2 to 5.5 mm
material described by Dusén as Cladophle- apart, sub-opposite to alternate, commonly
bis seymourensis comb. nov. dissected with a distinctive larger basal
pinnule on the acroscopic margin (Fig.
Family INCERTAE SEDIS 2D). Rhachis or rhachilla with distinct rib
(Fig. 2B) Pinnae inserted at 68 to 908
Sphenopteris (Brongniart) Sternberg, 1825 bearing sub-opposite to alternate pinnules.
Basal acroscopic pinnules larger and com-
Type species. Sphenopteris elegans (Brong- monly ovate (Fig. 2D) with one midvein or
niart) Sternberg, 1825 a single vein that dichotomizes. Most

"
Fig. 4. Conifers from the Paleocene Cross Valley Formation. A–D, Araucaria imponens (Dusén) comb. nov. A,
DJ1111.139, isolated leaf with contracted base. B, S132021_01, lectotype. Note contracted base and lack of apical
preservation. C, DJ1111.14 large leaf. D, D512.2a leaf with well-preserved leaf apex. E–G, Elatocladus seymourensis
sp. nov. E, S132903, F, S132020, G, S132904. Scale bars ¼ 10 mm.
Downloaded by [Fudan University] at 10:48 29 April 2013

ALCHERINGA
ANTARCTIC PALEOCENE FLORA
319
 320 DAVID J. CANTRILL et al. ALCHERINGA

pinnules with a single midvein, and lacking Lectotype. S122021_01 designated here and
secondaries (Fig. 2I). figured by Dusén (1908 on pl. 1, fig. 16) and
here as Fig. 4B.
Remarks. Dissected foliage was assigned to
Lomatia angustilobia but re-examination of Material examined. Four specimens:
the specimens shows several with open D512.2a, DJ1111.14, DJ1111.139, S
branching venation pattern (Fig. 2I) sug- 122021_01.
gesting this material is better placed within
the ferns. Lomatia brevipinna is included Description. Detached leaves up to 90 mm
here also because of the same open branch- long and 20 mm wide, lanceolate to linear
ing venation enabling placement in the lanceolate with an acute apex (Fig. 4C, D)
pteridophytes. Dusén (1908) used Sphae- and tapering leaf base (Fig. 4A, B). Leaf
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nopteris for this foliage however, it is better bases commonly broken but where pre-
placed with Sphenopteris as the pinnules are served broad and probably adnate on the
terminated by a single vein. stem. Leaf margins entire and leaves with
Specimens of this taxon are more com- 12–23 parallel unbranched veins extending
plete than the other ferns with large frag- to the leaf apex (Fig. 4C).
ments bearing gracile pinnae preserved (Fig.
2A). The larger frond fragments allow a Remarks. Dusén (1908) described one leaf
determination of pinnule variation over the (S122021_01) and included an illustration
whole frond. Specimens such as that illu- and an enlargement of the leaf base. This
strated in Fig. 2B probably represent the leaf is 60 mm long but is missing the apex
terminal portions of primary pinnae as and 12 mm wide. The leaf apex (on other
indicated by the reduction in size of the specimens observed) is tapered and the base
secondary pinnae towards the apex. Within is broad. It is unclear if the leaf base is
the deposit are numerous fragmentary broken, but many studied specimens have
remains consisting of a few pinnules (Fig. broken leaf bases. Within the deposit these
2C). Arrangement of pinnules on the frond, leaves are always found unattached suggest-
together with venation strongly supports the ing that they were shed as units from the
placement into the genus Sphenopteris, with parent plant. This is in contrast to many
the new combination S. angustiloba. araucarian leaves from the Eocene of
Seymour Island that are preserved as iso-
Phylum PINOPHYTA Cronquist, Takhtajan lated fragments of axes with many leaves
& Zimmerman ex Reveal, 1996 still attached suggesting that the Paleocene
Order PINALES Gorozhankin, 1904 material belongs to a distinct taxon.
Family ARAUCARIACEAE Henkel & Foliage of this form has been described
W. Hochstetter, 1865 from the Cretaceous of Alexander Island
where several species were recognized
Araucaria de Jussieu, 1789 based on variations in leaf form (Cantrill
& Falcon-Lang 2001). However, the Alex-
Type species: Araucaria araucana (Molina) ander Island material is nearly always
K. Koch, 1872 attached to branch units and is distinct
from the taxon recognized here. Other
Araucaria imponens Dusén, 1908 (Fig. 4A–D) araucarian foliage recognized from the
Antarctic Peninsula includes Maastrichtian
1908 Araucaria imponens Dusén, p. 11, pl. 1, material from Vega Island (Césari et al.
figs 16, 17. 2001). From that assemblage, conifer
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 321

shoots with broad leaves up to 70 mm long 4E, F). Leaves linear lanceolate up to
were described in association with an 16 mm long by 2 mm wide. Leaf margins
araucarian cone. Isolate leaves were not entire, leaf apices acute. Leaves traversed by
recorded but this foliage type extends into a single midvein. Leaf bases decurrent on
the Eocene La Meseta Formation on axis with the free portion twisted and
Seymour Island. This points to a wide- slightly constricted where it departs
spread and persistent occurrence of this the shoot; inserted at between 48 and 688
family in the Antarctic Peninsula—an (Fig. 4G).
observation supported by extensive wood
records from Cretaceous (e.g. Falcon-Lang Remarks. This material was assigned to
& Cantrill 2000, 2001) and Cenozoic Phyllites by Dusén (1908) but reassigned to
(Cantrill & Poole 2005b) deposits. the conifers by Florin (1940) and placed in
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two genera: Acmopyle and Podocarpus.

Family ?PODOCARPACEAE Endlicher, Impression foliage lacking cuticular infor-
1847 mation has been assigned to Elatocladus, a
form genus erected by Halle (1913) for
Elatocladus Halle, 1913 material from the Jurassic of Hope Bay.
Halle (1913) recognized four species based
Type species. Elatocladus heterophylla Halle, on differences in leaf form and suggested
1913 that this genus be used for foliage of the
dorsiventral type that lacked characters that
Elatocladus seymourensis sp. nov. (Fig. 4E– would permit placement in more peculiar
G) forms. This seems to imply more poorly
preserved material is to be placed in
1908 Phyllites sp. 4; Dusén, p. 17, pl. 3, figs Elatocladus, a practice endorsed by Harris
4, 6. (1979) in his emendation of the genus to
1908 Phyllites sp. 5; Dusén, p. 17, pl. 3, fig. include only gross morphological features.
5. Recent treatments of the Hope Bay flora
1940 Acmopyle; Florin, p. 32. suggest the four species recognized by Halle
1940 Podocarpus subgenus Stachycarpus (1913) encompass the range of variation
(now Prumnopitys); Florin, p. 32. seen in extant conifers where juvenile,
intermediate and adult foliage can be
Holotype. S132020. present (Rees & Cleal 2004).
The morphological characters seen in the
Etymology. Named after Seymour Island. two taxa recognized by Dusén (1908) are
also similar enough to account for the
Material examined. Three specimens: morphological variation within a species
S132903, S132020, S132904. on a single plant at varying places along a
branch. Venation and leaf shape are iden-
Diagnosis. Conifer shoots with helically tical, supporting assignment to one species
inserted bifacially flattened leaves up to (E. seymourensis), rather than two genera as
16 mm long and of uniform size for most of proposed by Florin (1940). Despite the poor
the shoot length. preservation, among the extant conifers, the
foliage is most similar to that seen in the
Description. Conifer shoots up to 6 cm long Podocarpaceae. However, the taxonomic
bearing helically inserted bifacially flattened assignments by Florin (1940) can not be
leaves so that they appear in two ranks (Fig. justified as they are based solely on leaf
 322 DAVID J. CANTRILL et al. ALCHERINGA

morphology without any details of the on the original descriptions by Dusén (1908)
cuticles. The placement of the material in have been made by several authors (e.g.
this case into modern genera is misleading Florin 1940, Gunckel Luer 1957, Van
and we have reassigned the material to a Steenis 1971, Romero & Dibbern 1985,
new species within the morphogenus Elato- Czajkowski & Rösler 1986, Romero 1986,
cladus. Tanai 1986, Case 1988, Zastawniak 1989,
Halle (1913) recognized four species, and Doktor et al. 1996). However, the majority
although synonymized by Rees & Cleal of these studies utlized collections from
(2004), they differ from the material de- younger Eocene localities, or focused solely
scribed here. The Seymour Island material on species within Fagus and Nothofagus, and
has larger leaves, up to 16 mm as opposed apparently did not review the original
to 11 mm, and the intermediate shoots that material held in Stockholm. Remarkably,
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the specimens are assumed to represent are only a single paper including small new
somewhat longer. Furthermore, the inter- collections has been published on these
mediate shoots of the material from Hope Paleocene leaf fossils in the past 100 years
Bay have very small leaves at the base that (Case 1988), despite the flora apparently
gradually lengthen in the mid-regions before being one of the most diverse in the
reducing in size again towards the inter- Antarctic (Truswell 1991). The extensive
mediate shoot apex. This gives the inter- new collection (housed at the British Ant-
mediate shoots a somewhat lanceolate arctic Survey), together with an examination
outline. In contrast, the material described of the original collection housed in the
here is more or less uniform in size for the Swedish Museum of Natural History, has
whole of the preserved shoot length. So, allowed us to revise the flora’s taxonomic
although the leaf insertion angles overlap diversity based on more complete specimens.
with the very broad ranges seen in the taxa As indicated by Dusén (1908) the poor
from Hope Bay, there are enough consistent preservation of the leaves makes it difficult
morphological differences to warrant recog- to determine venation, particularly of high-
nizing a separate species. er orders, and is it is acknowledged as being
The spreading foliage is typical of that difficult to determine taxonomic relation-
seen within the Podocarpaceae and although ships. However, over 1000 specimens have
we can not confidently place it within this been examined from Nordenskjöld, Falk-
family it is the most likely candidate. The land Island Dependency Survey, British
conifer pollen record from the Cross Valley Antarctic Survey archives and the current
Formation indicates the presence of only authors’ collections in 1999–2001, which
Araucariaceae and Podocarpaceae. Cupres- provided enough whole specimens with
saceous foliage and wood is known from well-preserved venation, or composite re-
younger (Eocene) sediments, where it is constructions of fronds, to allow confident
extremely rare. Interestingly, Elatocladus identification of the taxa.
seymourensis has not been re-collected by Dusén (1908) described a diverse assem-
subsequent expeditions and the taxon is an blage of 37 fern taxa. However, only eight
extremely rare component of the flora. were formally described (Polypodium aemu-
lans, P. nathorstii, Asplenium antarcticum,
Taenioptiers blechnoides, Taeniopteris cris-
Discussion pissima, Alsophilia antarctica, Dryopteris
A century has now passed since the initial seymourensis, D. antarctica). The remainder
collections from Cross Valley were described were either described and assigned to
by Dusén (1908). Taxonomic commentaries informal species of extant genera (e.g.
 ALCHERINGA ANTARCTIC PALEOCENE FLORA 323

Asplenium sp.) or simply illustrated and species: Cladophlebis aemulans (Dusén)

assigned to morphogenera Pecopteris (18 comb. nov. (incorporating Taeniopteris),
species) or Sphaenopteris (10 species). Many Cladophlebis sp. 1, and Sphenopteris sp. 1.
of the specimens assigned to different This considerably reduces the fern diversity
species were based on very fragmentary within this flora (Table 1).
material (just a few pinnules), or were single Dusén (1908) recognized just one conifer
specimens. The morphological variation (Araucaria imponens) based on a single leaf
seen in a single compound frond of an specimen. The new material confirms that this
extant fern encompasses the variation evi- conifer is present in consistent, but low
dent between many of the fragments as- numbers within the flora. Material described
signed to Pecopteris or Sphaenopteris. as Phyllites by Dusén (1908) was reassigned to
Morphometric studies of large frond frag- Acmopyle and Podocarpus subgenus Stachy-
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ments have recognized that this variation carpus (now Prumnopitys) by Florin (1940).
was often not considered when assigning The taxonomic assignments by Florin can not
material to fossil taxa in early studies be justified as they are based solely on leaf
(Rees & Cleal 1993). morphology without any cuticular details.
Based on the morphological variation The placement of the material in this case into
evident in extant ferns, and fossil studies, we modern genera is misleading and we have
reassign the pteridophytes to just three reassigned the material to a new species within

Dusén (1908) flora This paper

Ferns Ferns
Polypodium aemulans Formally described Cladophlebis aemulans
Taeniopteris blechnoides (Dusén) comb. nov.
Taeniopteris crispissima
Alsophila antarctica
Polypodium nathorstii Formally described Cladophlebis seymourensis
Dryopteris antarctica (Dusén) comb. nov.
Dryopteris seymourensis
Pecopteris (18 species) Illustrated only
Lomatia angustiloba Formally described Sphenopteris angustiloba
Lomatia brevipinna (Dusén) comb. nov.
Asplenium antarcticum
Asplenium sp.
Sphaenopteris (10 species) Illustrated only
Conifers Conifers
Araucaria imponens Formally described Araucaria imponens Dusén, 1908
Phyllites sp. 4 Described Elatocladus seymourensis sp. nov.
(Acmopyle Florin, 1940)
Phyllites sp. 5
(Podocarpus subgenus Stachycarpus,
now Prumnopitys, Florin, 1940)

Table 1. Summary of ferns and conifers, initially described by Dusén (1908), that are represented within
the Paleocene-aged Cross Valley flora from Seymour Island, Antarctic Peninsula, with comparisons to
species described in the current revision of the Dusén (1908) flora.
 324 DAVID J. CANTRILL et al. ALCHERINGA

the morphogenus Elatocladus. In summary, Natural History is thanked for giving

the conifers are rare in the flora but can be assistance and access to the Nordenskjöld
assigned to two taxa within two families: Collection. Reviewers are acknowledged for
Araucaria imponens Dusén, 1908 (Araucaria- helpful comments on the manuscript.
ceae) and Elatocladus seymourensis sp. nov.
(Podocarpaceae?).
The diversity of the Cross Valley Flora References
has been here revised to two fern families ASKIN, R.A., 1988a. Campanian to Paleocene palyno-
(three species): Cladophlebis aemulans (Du- logical succession of Seymour Island and adjacent
sén) comb. nov. (incorporating Taeniop- islands, northeastern Antarctic Peninsula. In Geol-
ogy and Paleontology of Seymour Island, Antarctica
teris), Cladophlebis seymourensis and Peninsula, R.M. FELDMANN & M.O. WOODBURNE,
Sphenopteris angustiloba; and two conifer eds, Geological Society of America Memoir 169,
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families: Araucaria imponens Dusén, 1908 131–153.

(Araucariaceae) and Elatocladus seymour- ASKIN, R.A., 1988b. The palynological record across
the Cretaceous/Tertiary transition on Seymour
ensis sp. nov. (?Podocarpaceae). The rigor- Island, Antarctica. In Geology and Paleontology
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within the Cross Valley Flora to enable a palynofloras of Seymour Island, Antarctica: im-
clearer picture of biodiversity within forests plications for origins, dispersal and palaeoclimates
growing in island arc settings on the of southern floras. In Origins and Evolution of the
Antarctic Biota, J.A. CRAME, ed., Special Publica-
Antarctic Peninsula during the late Paleo- tion—Geological Society of London 47, 107–119.
cene. Future papers will address the angios- ASKIN, R.A., 1990. Campanian to Paleocene spore and
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Funding of JEF, DJC and the postdoctoral A Perspective on Global Change, J.P. KENNETT &
D.A. WANKLE, eds, Antarctic Research Series
position of A-M.P.T. was derived from the American Geophysical Union, Volume 56, Wa-
NERC-BAS Antarctic Funding Initiative shington, 61–73.
grant GR3/G0001. All fieldwork to Antarc- ASKIN, R.A., 1994. Monosulcate angiosperm pollen
tica was supported by this funding initiative, from the Lopez de Bertodano Formation (upper
Campanian–Maastrichtian–Danian) of Seymour
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