Araucarian leaves and cone scales from the Loreto Formation of Río de Las Minas, Magellan Region, Chile 1

Abstract

Cone scales and leaves of the Araucariaceae are reported from the Loreto Formation in Río de Las Minas, Punta Arenas, Chile. Two types of cone scales including one new species, Araucarites alatisquamosus are recognized. They are similar to Araucaria section Eutacta. Two types of leaves are assigned to Araucaria nathorstii Dusén, and one new type is distinguished in the specimens from the type locality of A. nathorstii. A narrow leaf type is identical to Dusén’s “Blätter der sterilen Zweige (leaves of sterile shoot)” of A. nathorstii, and a broad leaf type is identical to his “Blätter der fertilen Zweige (leaves of fertile shoot)”. Both types show characters of leaves of section Araucaria as suggested by earlier studies. A smaller leaf type differs from the original description of A. nathorstii, and consists of small appressed leaves with the obtuse apex and stomatal orientation that are the characteristic of the section Eutacta. The occurrence of leaves and cone scales similar to section Eutacta suggest the presence of araucarians close to this section in southernmost South America during the Eocene-early Oligocene, and provide evidence for the wide distribution and diversity of genus Araucaria relatives in South America during the Paleogene.

Full text

ARTICLE
Araucarian leaves and cone scales from the Loreto
Formation of Río de Las Minas, Magellan Region, Chile
1
Takeshi A. Ohsawa, Atsushi Yabe, Toshihiro Yamada, Kazuhiko Uemura, Kazuo Terada,
Marcelo Leppe, Luis Felipe Hinojosa, and Harufumi Nishida
Abstract: Cone scales and leaves of the Araucariaceae are reported from the Loreto Formation in Río de Las Minas,
Punta Arenas, Chile. Two types of cone scales including one new species, Araucarites alatisquamosus are recog-
nized. They are similar to Araucaria section Eutacta. Two types of leaves are assigned to Araucaria nathorstii Dusén,
and one new type is distinguished in the specimens from the type locality of A.nathorstii. A narrow leaf type is
identical to Dusén’s “Blätter der sterilen Zweige (leaves of sterile shoot)” of A.nathorstii, and a broad leaf type is
identical to his “Blätter der fertilen Zweige (leaves of fertile shoot)”. Both types show characters of leaves of section
Araucaria as suggested by earlier studies. A smaller leaf type differs from the original description of A.nathorstii,
and consists of small appressed leaves with the obtuse apex and stomatal orientation that are the characteristic of
the section Eutacta. The occurrence of leaves and cone scales similar to section Eutacta suggest the presence of
araucarians close to this section in southernmost South America during the Eocene–early Oligocene, and provide
evidence for the wide distribution and diversity of genus Araucaria relatives in South America during the Paleogene.
Key words: Araucaria, conifer, Eutacta, Paleogene, Patagonia.
Résumé : On rapporte la présence d’écailles de cônes et de feuilles d’Araucariaceae dans la formation de Loreto au Rio
de Las Minas, Punta Arenas, Chili. On trouve deux types d’écailles de cônes y compris celles d’une nouvelle espèce,
Araucarites alatisquamosus. Elles sont similaires a
`celles de la section Eutacta d’Araucaria. Deux types de feuilles sont
assignés a
`Araucaria nathorstii Dusén et un type nouveau se distingue chez les spécimens de la localité type d’A. nathorstii.
Un type de feuille étroite est identique a
`celui d’A.nathorstii de Dusén « Blätter der sterilen Zweige » (feuilles d’une pousse
stérile) alors qu’un type de feuille large est identique a
`celui du « Blätter der fertilen Zweige » (feuille d’une pousse fertile). Les
deux types présentent les caractères foliaires de la section Araucaria comme le suggèrent des études précédentes. Un type de
feuille plus petite diffère de celui originalement décrit chez A.nathorstii et il consiste en de petites feuilles apprimées dont
l’apex obtus et l’orientation des stomates sont caractéristiques de la section Eutacta. La présence de feuilles et d’écailles de
cônes similaires a
`celles de la section Eutacta suggère la présence d’araucariens proches de cette section dans l’extrême sud de
l’Amérique du Sud durant l’Éocène et l’Oligocène précoce, et donne la preuve d’une distribution et d’une diversité larges des
parents du genre Araucaria en Amérique du Sud pendant le Paléogène. [Traduit par la Rédaction]
Mots-clés : Araucaria, conifères, Eutacta, Paléogène, Patagonie.
Introduction
Araucaria Juss. (Jussieu 1789) of the Araucariaceae Henkel
et W. Hochstetter (1865) is a typical Southern Hemisphere
taxon distributed disjunctively in Australia, New Caledonia,
New Guinea, and South America (Farjon 2010;Debreczy
and Rácz 2011). This disjunctive distribution has been con-
sidered to be related to a break-up of the Gondwana super-
continent. There are 19 living species in four sections of
Received 2 March 2016. Accepted 4 June 2016.
T.A. Ohsawa. Graduate School of Science, Chiba University, 1-33 Yayoicho, Chiba 263-8522, Japan.
A. Yabe and K. Uemura. Department of Geology and Paleontology, National Museum of Nature and Science, 4-1-1 Amakubo,
Tsukuba, Ibaraki 305-0005, Japan.
T. Yamada. School of Natural System, College of Science and Engineering, Kanazawa University, Kanazawa 920-1192, Japan.
K. Terada. Fukui Prefectural Dinosaur Museum, 51-11 Terao, Muroko, Katsuyama, Fukui 911-8601, Japan.
M. Leppe. Instituto Antártico Chileno, Plaza Muñoz Gamero 1055, Punta Arenas, Chile.
L.F. Hinojosa. Laboratorio de Sistemática y Ecología Vegetal, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa,
Santiago CP 780-0024, Chile.
H. Nishida. Department of Biological Sciences, Faculty of Science and Engineering, Chuo University, 1-13-27 Kasuga, Bunkyo,
Tokyo 112-8551 Japan; Graduate School of Biological Science, University of Tokyo, Japan.
Corresponding author: Takeshi A. Ohsawa (email: asakawa@faculty.chiba-u.jp).
1
This Article is part of a Special Issue entitled “Mesozoic and Cenozoic Plant Evolution and Biotic Change,” a collection of research
inspired by, and honouring, Ruth A. Stockey.
Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink.
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Araucaria, including Eutacta Endl., Bunya Wilde and Eames,
Intermedia White, and Araucaria Juss. In addition, Section
Yezonia has been proposed for a fossil species A.vulgaris
(Stopes et Fujii) Oshawa, H. Nishida et Nishida, which has
Brachyphyllum-like shoots and Eutacta-like cone scales
(Ohsawa et al. 1995). The center of diversity for living
species is the Oceania region, where 17 of 19 species as-
signed to three sections, Eutacta,Bunya, and Intermedia,
are distributed. In South America, only two species of
section Araucaria (Columbea) are distributed in south
Chile, Argentina, and southwestern Brazil.
South America is considered to have played an impor-
tant role in the distribution of Araucaria (Del Fueyo and
Archangelsky 2002). Three of four extant sections have
been found as macrofossils such as leaves, shoots, and
cone scales. Section Araucaria is represented by A.cartellei
Duarte from the Lower Cretaceous of Brazil (Duarte
1993), Araucaria grandifolia Del Fueyo and Archangelsky
from the Lower Cretaceous of Argentina (Del Fueyo and
Archangelsky 2002), A.pararaucana Panti et al. (2007)
from the Paleogene of Argentina, and A.nathorstii Dusén
(1899) from the Paleogene–Noegene of Argentina and
Chile (Berry 1928;Doktor et al. 1996;Fiori 1939;Hünicken
1995;Menéndez and Caccavari 1966). Section Bunya is
represented by the seed cones Araucaria mirabilis (Spe-
gazzini) Windhausen (1931) from the Jurassic of Argen-
tina (Stockey 1978), although its relationship to section
Bunya was not supported by recent phylogenetic analyses
based on molecular and morphological data (Escapa
and Catalano 2013). Section Eutacta is represented by
cone-scales described as Araucarites species; A.minimus
Archangelsky (1966),A.baqueroensis Archangelsky (1966),
and A.chilensis Baldoni (1979) from the Lower Cretaceous
of Argentina and Chile, and Araucaria pichileufulensis
Berry (1938) from the Eocene of Argentina.
Araucarites Presl (in Sternberg 1820–1838) is a fossil genus
reported worldwide, and designated by Zijlstra et al. (2000)
as representing araucarian seed cones and cone scales. The
existence of section Yezonia in South America could be
inferred by the presence of the pollen cone Notopeuen
brevis Del Fueyo (1991) containing Araucariacites pollen
grains and attached to branches bearing Brachyphyllum
leaves. A Eutacta-type cone scale, Araucarites chilensis
Baldoni (1979) from the Lower Cretaceous of El Cóndor,
Santa Cruz, Argentina is also associated with Brachyphyllum
feistemantelii (Halle) Sahni, although the evidence of sec-
tion Yezonia is still insufficient in the Southern Hemi-
sphere.
In contrast to abundant and diversified macrofossil
records earlier than the Early Cretaceous, macrofossils
except permineralized wood are very rare from the Late
Cretaceous in South America (Panti et al. 2012). Only one
species represented by cone scales, Araucarites patagonica
Kurtz (1902) has been described from the Upper Creta-
ceous of Chile. However, it is designated as a nomen
nudum by Hünicken (1971) because of its poor preservation
and insufficient description. In the Paleogene, four spe-
cies, Araucaria araucoensis Berry (1922),A.pichileufulensis,
A.pararaucana, and A.nathorstii have been described
from Argentina and Chile. Most of them are assigned
to section Araucaria except for A. pichileufulensis, which
is attributed to section Eutacta. Some authors suppose
that sections Bunya and Eutacta had become extinct in
South America at the end of Mesozoic (Veblen et al.
1995;Del Fueyo and Archangelsky 2002). To understand
post-Cretaceous evolutionary history and biogeography
of the genus Araucaria and its sections in South America,
more fossil evidence is needed.
In this paper, we report newly collected araucarian
fossils from the late Eocene to earliest Oligocene Loreto
Formation which crops out along the river, Río de Las
Minas north of Punta Arenas, Magellan Region, Chile.
Two types of cone scales and three types of shoots and
isolated leaves have been recognized. All are assignable
to the genus Araucaria, including morphotypes compara-
ble with A.nathorstii previously described from the same
locality. The cone scales and one morphotype of shoot
and leaf have features characteristic of section Eutacta,
suggesting the possible coexistence of the section Eutacta
with other section of Araucaria until the late Paleogene in
South America.
Materials and methods
The fossils were collected from the Loreto Formation,
which is exposed along the river Río de Las Minas, near
Punta Arenas, Magellan Region, Chile (Fig. 1). The U–Th–Pb
SHRIMP age of 36.48 ± 0.47 Ma is derived from the anal-
ysis of zircons obtained from the uppermost part of the
Loreto Formation (Otero et al. 2012), thus the main part
of the formation is thought to have been deposited in the
late Eocene. This estimate is consistent with the age
based on dinoflagellates (H. Kurita, personal communi-
cation, 2015).
The Loreto Formation mainly consists of sandstones
and mudstones and several intercalated coal seams of
variable thickness. Most of the formation was deposited
in a shallow marine environment, because glauconites,
as well as abundant elasmobranch fossils, are contained
in the sandstones (Otero et al. 2012). Some deposits
were formed in brackish environments, as indicated
by herring-bone cross-stratification, typically formed by
tidal currents. The araucarian remains were densely ac-
cumulated in a carbonaceous mudstone bed intercalated
between sandstone beds of brackish origin, along with
various forms of angiosperm leaves including Nothofagus.
A leafy shoot of Araucaria nathorstii and some species of
Nothofagus have been described previously from this lo-
cality (Dusén 1899). Some species of silicified woods in-
cluding Araucarioxylon and Nothofagoxylon have also been
reported recently (Terada et al. 2006).
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Organically preserved compressions of vegetative shoots,
detached leaves and cone scales were collected by the
authors during repeated field studies at the locality in
2005, 2007, 2008, and 2016. The fossils were stored in
glycerin alcohol solution (glycerin–ethanol = 1:1, v/v)to
prevent rapid disorganization due to desiccation. Speci-
mens were photographed using a Pentax K-5 IIs camera
with a smc Pentax-D FA Macro 50 mm lens attaching a
polarizing filter. Cuticles were prepared using Schultz’s
solution (e.g., Kerp and Krings 1999) followed by diluted
sodium hydroxide or using commercial bleach (sodium
hypochlorite solution) until the cuticles were cleared. Ei-
ther lower or upper cuticles, or both, were removed, rinsed
with deionized water, and mounted in glycerine jelly. Cu-
ticular features were examined with an optical microscope
(Nikon Optiphot and Leica DMR) and photographed using
Nikon DS-Fi1 and Leica MC170HD microscope cameras.
All specimens examined are deposited in the National
Museum of Nature and Science, Tokyo, except for the
holotype specimen of the new species (specimen No.
SGO.PB 1577), which will be deposited in the Paleonotol-
ogy Section of the Museo Nacional de Historia Natural,
Santiago, Chile.
Results
Araucarian cone scales
Two types of morphologies were identified from abun-
dant specimens of detached cone scales. One is desig-
nated here as a new species of Araucarites, and the other
as Araucarites sp.
Systematic Palaeontology
Family Araucariaceae Henkel and W. Hochstetter, 1865
Araucarites Presl 1838
Araucarites alatisquamosus sp. nov. Ohsawa et H. Nishida.
Figures 2A(2)–2A(6).
HOLOTYPE:SGO.PB 1577, Museo Nacional de Historia Natu-
ral, Santiago, Chile.
PARATYPES:NSM PP-12151, NSM PP-12152, NSM PP-12153.
TYPE LOCALITY AND HORIZON:Loreto Formation, Río de Las
Minas area near Punta Arenas, Magellan Region, Chile.
Late Eocene – early Oligocene.
ETYMOLOGY:Species epithet, alatisquamosus originates from
a Latin alatus for “winged” and squamosus for “scaly”.
DIAGNOSIS:Fossil detached cone scales composed of cune-
ate bract, 32–33 mm long by 30–34 mm wide. Delicate
wings expanded laterally up to 5– 6 mm from each side of
woody central zone of cone scale. Bract spine short,
3 mm long. Single oval seed embedded in ovuliferous
scale, 18–24 mm long by 5–5.5 mm wide. Ovuliferous
scale surrounding seed, oval, 7.6–8.0 mm at the widest
portion. Ovuliferous scale tip rounded, about 2 mm from
distal end of seed.
DESCRIPTION:Detached cone scales consist of a cuneate
bract and an attached ovuliferous scale (Figs. 2A(2) and
2A(3)). The cone scale is winged, 32–33 mm long by 30–
34 mm wide at the widest portion including lateral
wings. The central part of the cone scale is woody and
thick. The delicate wings, which are often degraded, ex-
pand laterally up to 5–6 mm on each side of the woody
central zone. The bract has distal rounded corners from
which it tapers to form a distinct shoulder, and termi-
nates in a short apical spine up to 3 mm long (Figs. 2A(3)
and 2A(5)). A single seed is embedded in the ovuliferous
scale tissue (Figs. 2A(4) and 2A(6)). The seed is approxi-
mately 18–24 mm long by 5.0 –5.5 mm wide, and broadest
near the distal end, sometimes represented by a circular
imprint (Fig. 2A(4)). The ovuliferous scale, 7.6–8.0 mm at
widest portion, surrounds the seed, and its tip is rounded
(Fig. 2A(4)).
Fig. 1. Map showing the fossil locality (arrowhead) based on Google Earth (© 2016 CNES/Astrium, Cnes/Spot Image,
DigitalGlobe, Landsat, US Geological Survey and Google).
Ohsawa et al. 807
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COMPARISONS:The cone scale described here is closely sim-
ilar to Araucaria because of its woody, winged bracts, free
ovuliferous scale tip, with a single seed embedded in
ovuliferous scale tissue, which is retained at maturity
(Stockey 1982). Araucarites was designated as represent-
ing araucarian seed cones and cone scales by Zijlstra
et al. (2000), and the isolated cone scales with araucarian
affinities have been assigned to Araucarites (Zijlstra et al.
2000). Thus we assigned our specimen to Araucarites
Presl. Araucarites alatisquamosus sp. nov. is character-
ized by its large size, prominent but fragile lateral wings
that are narrower than the main seed body, and ovulif-
erous scale that has a rounded tip. In the genus Araucaria,
the ovuliferous scale is vestigial in section Araucaria (van
der Ham et al. 2010). Araucarites alatisquamosus differs
from cone scales of section Araucaria by having promi-
nent wings. The cone scales of section Bunya are charac-
terized by thick and rigid lateral wings and a prominent
woody tip of the ovuliferous scale (van der Ham et al.
2010). The lateral wings of Araucarites alatisquamosus are
thin and delicate in contrast to those of section Bunya.
The cone scales of section Intermedia are relatively large,
Fig. 2A. Araucarites alatisquamosus sp. nov. (2–6). (2) Cone scale of holotype specimen. White arrowheads on the left indicate
the edge of lateral wing. The right hand side of the structure was not preserved. SGO.PB 1577a. Scale bar = 10 mm. (3) Counter
part of Fig. 2. White arrowheads on the right indicate the edge of lateral wing. Black arrowhead points to the apex of scale,
the lower portion of which was curved and not preserved. SGO.PB 1577b. Scale bar = 10 mm. (4) Close-up of (3) showing seed
with circular imprint and ovuliferous scale (white arrow). Scale bar = 5 mm. (5) Cone scale of paratype specimen. Black
arrowhead indicates the apex of the scale. Two white arrowheads indicate distal rounded corner of scale. NSM PP-12152. Scale
bar = 10 mm. (6) Close-up of a paratype specimen showing seed and ovuliferous scale. NSM PP-12153. Scale bar=5mm.
Fig. 2A. (7 and 8) Araucarites sp. (7) Cone scale. Two white arrowheads indicate distal rounded corner of scale. NSM PP-12154.
Scale bar = 5 mm. (8) Close-up of cone scale in (7), showing the triangular tip of ovuliferous scale (white arrows). NSM PP-12154.
Scale bar = 2 mm. [Colour online.]
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with a narrow central woody portion and broad papery
wings that are broader than the central woody portion
(Wilde and Eames 1948;van der Ham et al. 2010). Araucarites
alatisquamosus is composed of a relatively wide central
woody portion and prominent lateral wings that are nar-
rower than the central woody portion. In these features,
Araucarites alatisquamosus is most similar to the cone
scales of section Eutacta.
A large number of fossil araucarian cones have been de-
scribed worldwide. They are assigned to either Araucarites
or Araucaria (Stockey 1994;Kunzmann 2007). In South
America and the Antarctic Peninsula, many species are
described from the Jurassic to the Eocene (Panti et al. 2012).
Among the described species, Araucaria pichileufulensis,
Araucarites baqueroensis,A.antarcticus (Gee) Birkenmajer et
Ociepa (2008) from the Jurassic of Antarctica, A.chilensis,
A.citadelbastionensis Cantrill and Falcon-Lang (2001) from
the Albian of Alexander Island, A.minimus, and A.phillipsi
Carruthers (1869) are characterized by prominent lateral
wings. Of these, A.antarcticus,A.chilensis,A.citadelbastionensis,
A.minimus, and A.phillipsi are much smaller than our
new species. Araucaria pichileufulensis and Araucarites
baqueroensis are most closely comparable with our new
species, sharing large cone scales with prominent lateral
wings. Araucaria pichileufulensis was established based on
leafy shoots and isolated cone scales. The cone scales are
kite-shaped with broad lateral wings, 27.5 mm long ×
30 mm wide including lateral wings. Araucaria pichileufulensis
is distinguished from Araucarites alatisquamosus by hav-
ing a prominent triangular tip rather than a rounded tip
on the ovuliferous scale and its slightly smaller size.
Araucarites baqueroensis is represented by isolated cone
scales 20–30 mm long × 20 mm wide. Seeds are 10–
15 mm × 4–5 mm, and embedded in the ovuliferous scale
with a rounded tip (Archangelsky 1966). Cone scales of
Araucarites alatisquamosus are much larger than those of
A.baqueroensis.
Araucarites sp. (Figs. 2A(7) and 2A(8)).
MATERIAL STUDIED:NSM PP-12154.
NOTE:The fossil is a detached cone scale consisting of
a cuneate bract and an attached ovuliferous scale
(Fig. 2A(7)). The cone scale is 12 mm long × 12 mm wide at
the widest point including the lateral wings. The central
part of the cone scale is woody and thick. Delicate wings
expand laterally up to 3 mm on each side of the woody
portion of the bract. The bract has distal rounded corners
from which it tapers to form a distinct shoulder. The
apical spine is not preserved in the specimen we ob-
served. The seed is long and oval, 7.6 mm long × 1.6 mm
wide, and is broadest near the distal end (Fig. 2A(7)). The
free ovuliferous scale tip is distinct, triangular, and
1.8 mm long × 2.8 mm wide at base (Fig. 2A(8)).
COMPARISONS:Araucarites sp. is characterized by prominent
lateral wings and a triangular ovuliferous scale tip. As is
the case with Araucarites alatisquamosus, this Araucarites
sp. is similar to the cone scales of section Eutacta. Besides
the smaller size, Araucarites sp. differs from A.alatisqua-
mosus in the presence of a triangular ovuliferous scale
tip. Among the Araucarites species described from South
America and the Antarctic Peninsula, A.antarcticus,
A.chilensis,A.citadelbastionensis,A.minimus, and A.phillipsi
Carruthers (1869) are characterized by small size and
prominent lateral wings. However, cone scales of A.minimus
and A.phillipsi are much smaller than those of the
Araucarites sp. described here, and do not have the trian-
gular ovuliferous scale tip (Archangelsky 1966;Arrondo
and Petriella 1980). Cone scales of Araucarites antarcticus
are slightly larger than those of Araucarites sp., and the
seed is broad-oval in outline (Birkenmajer and Ociepa
2008), in contrast to the elongate oval seed of Araucarites
sp., Araucarites chilensis has cone scales as long as those of
the Araucarites sp. described here. However, its lateral
wings are not as wide as those of the Araucarites sp., and
the cone scale is no more than 8 mm wide (Baldoni 1979)
instead of the 12 mm in Araucarites sp. The Araucarites sp.
described here closely resembles A.citadelbastionensis in
size, and in having broad wings and triangular ovulifer-
ous scale tip. Considering that A.citadelbastionensis is de-
scribed from the Lower Cretaceous (as opposed to the
late Eocene – early Oligocene) of Coal Nunatak, Alexan-
der Island, Antarctica, the Araucarites sp. cone scales may
represent a different species. However, it is premature to
conclude whether or not Araucarites sp. is conspecific
with A.citadelbastionensis until further information has
been compiled for our Araucarites sp.
Araucarian leaves
Araucaria nathorstii was described by Dusén for
impression/compression shoots and leaves from the same
locality and formation as our specimens (Dusén 1899).
Specimens used for the original description show great
variation in form, and Dusén (1899) described them as
“Blätter der sterilen Zweige (leaves of sterile shoot)” and
“Blätter der fertilen Zweige (leaves of fertile shoot)”.
Leaves of his sterile shoots are lanceolate with fine lon-
gitudinal ridges (Dusén 1899, Plate XII, fig. 3), while those
of the fertile shoots are ovate, broadly lanceolate or
sometimes almost triangular with dense longitudinal
ridges (Dusén 1899, Plate XII, figs. 4–10). However, one
leaf specimen of his fertile shoot (Dusén 1899, Plate XII,
figs. 11 and 12) is smaller and has sparse longitudinal
ridges. In our specimens, we distinguished three leaf
types, some attached on shoots. These are a narrow type
comparable to the vegetative-shoot leaves of Dusén, a
broad type comparable with the fertile-shoot leaves, and
a smaller type similar to the small fertile-shoot leaf with
sparse ridges. The smaller type is quite different from the
two forms of A.nathorstii, and we conclude that it should
be distinguished from A.nathorstii. Here we report the
narrow and the broad types of A.nathorstii, and a new
leafy shoot that we describe as Araucaria sp.
Araucaria nathorstii Dusén 1899 (Figs. 2B(9)–2C(26)).
Ohsawa et al. 809
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MATERIALS STUDIED:narrow type, NSM PP-12155; broad type,
NSM PP-12156, NSM PP-12157, NSM PP-12158, NSM PP-
12159, NSM PP-12160, NSM PP-12161, NSM PP-12162, NSM
PP-12163, NSM PP-12164.
Notes on leaf types
NARROW TYPE LEAVES:(Figs. 2B(9)–2B(11),2C(18)–2C(22)): One
isolated shoot, up to 55 mm in width (Fig. 2B(9)), is iden-
tified as the narrow-type of A.nathorstii. The stem is 8 mm
in diameter and shows rhomboidal and slightly curved
scars 3.7–4.4 mm in horizontal and 3.0–3.8 mm in verti-
cal dimensions (Figs. 2B(9) and 2B(11)). The shoot bears
densely packed imbricate leaves (Fig. 2B(9)). The leaves
are lanceolate, flat and not keeled, with an acute apex
and a decurrent base (Fig. 2B(10)), 23–31 mm long by 6.5–
8.1 mm wide at the widest point, tapering to the base, but
not dramatically contracted at the very base. The leaf has
entire margins and 7–9 longitudinal ridges, which prob-
ably correspond to the positions of veins.
These leaves are amphistomatic, but stomata are
sparse on the adaxial cuticle, where the epidermis is
composed of mostly elongate rectangular cells (Fig. 2C(18)).
The stomata are rare and arranged in indistinct rows. In
Fig. 2B. Araucaria nathorstii.(9–17).(9) Shoot of narrow type. NSM PP-12155. Scale bar = 10 mm. (10) Leaf of narrow type. NSM
PP-12155. Scale bar = 4 mm. (11) Close-up of stem showing leaf scars. NSM PP-12155. Scale bar = 4 mm. (12) Isolated leaf of broad
type with triangular outline. NSM PP-12158. Scale bar = 4 mm. (13) Ovate leaf attached to the stem. White arrowheads indicate
position of leaf attachment. NSM PP-12156. Scale bar = 5 mm. (14) Isolated leaf of broad type with ovate outline. NSM PP-12161.
Scale bar = 5 mm. (15) Shoot of broad type. NSM PP-12156. Scale bar = 10 mm. (16) Close-up of stem showing leaf scars. NSM PP-
12156. Scale bar = 5 mm. (17) Shoot of broad type. NSM PP-12157. Scale bar = 10 mm. [Colour online.]
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the abaxial cuticle, stomata are arranged in regular rows
with the apertures orientated almost parallel to the leaf
axis (Figs. 2C(19) and 2C(20)). The stomatal apparatus is
elliptical, monocyclic and usually has four subsidiary
cells (Figs. 2C(20) and 2C(21)). Stomata in the same row
are separated by 1–2 rectangular epidermal cells. Inters-
tomatal bands consist of 1–5 rows of elongate rectangu-
lar cells, excepting certain broad bands up to 15 cells
wide. Anticlinal walls of the epidermis are obviously sin-
uous (Fig. 2C(22)).
Fig. 2C. Cuticle of Araucaria nathorstii Dusén. (18–26).(18) Stomata of adaxial cuticle, showing four subsidiary cells. NSM
PP-12155. Scale bar = 50 ␮m. (19) Abaxial cuticle, showing elongate rectangular normal cells and regular stomatal rows.
NSM PP-12155. Scale bar = 100 ␮m. (20) Details of stomatal rows of abaxial cuticle. Stomatal apparatus in same row
separated by 1–2 regular epidermal cells. NSM PP-12155. Scale bar = 50 ␮m. (21) Stomata in abaxial cuticle showing four
subsidiary cells. NSM PP-12155. Scale bar = 50 ␮m. (22) Sinuous anticlinal walls of abaxial epidermal cells. NSM PP-12155.
Scale bar = 50 ␮m. (23) Stomata of adaxial cuticle, and four subsidiary cells. NSM PP-12164. Scale bar = 50 ␮m. (24) Adaxial
cuticle, showing three stomatal rows. Stomatal rows separated by 3–5 rows of elongate rectangular cells. NSM PP-12164.
Scale bar = 100 ␮m. (25) Abaxial cuticle, showing stomatal rows, with stomata of variable orientation. NSM PP-12164. Scale bar =
100 ␮m. (26) Close-up of (25). Stomatal apparatus monocyclic, with 4–6 subsidiary cells. NSM PP-12164. Scale bar = 50 ␮m.
Ohsawa et al. 811
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BROAD TYPE LEAVES:(Figs. 2B(12)–2B(17),2C(23)–2C(26)): Sev-
eral compressed shoots and isolated leaves are identified
as the broad-type of A.nathorstii. The shoot is up to 60 mm
wide, with the stem 9 mm in diameter (Figs. 2B(15)–
2B(17)). The stem shows rhomboidal and slightly curved
scars more than 6 mm in horizontal and 4.5–5.3 mm in
vertical dimensions (Fig. 2B(16)). The shoot bears densely
packed imbricate leaves. The leaves diverge at 55–72 de-
grees from the stem axis, and are ovate, concave, and not
keeled, with an entire margin, an acute apex, and a de-
current base (Figs. 2B(12)–2B(14)), and are 28–40 mm long
by 13–20 mm wide at the widest portion. The leaf base is
narrower than the widest part, 9–13 mm wide, and not
contracted. The leaves exhibit 25–30 dense vertical
ridges, which probably correspond to vein positions or
stomatal lines.
These wide leaves are amphistomatic. In adaxial
cuticle, stomata are arranged mostly in regular rows with
the apertures orientated almost parallel to the leaf axis
(Figs. 2C(23) and 2C(24)). The stomatal apparatus is ellip-
tical, monocyclic, and composed of mostly four but up to
six subsidiary cells (Fig. 2C(23)). Stomata in the same row
are separated by 1–2 rectangular epidermal cells. Inters-
tomatal bands consist of 3–5 rows of elongate rectangu-
lar cells. The abaxial cuticle shows irregular rows of
more densely aggregated stomata with the apertures ori-
entated mostly parallel to, but sometimes oblique or per-
pendicular to the leaf axis (Figs. 2C(25) and 2C(26)). The
stomatal apparatus is similar in morphology to that of
the other cuticle observed. Stomata of the same row
are usually continuous and (or) sometimes separated by
1–2 epidermal cells. Adjacent stomatal rows are sepa-
rated by 1–2 rows of rectangular cells. Anticlinal walls of
epidermal cells are usually smooth and not sinuous.
Araucaria sp. (Figs. 2D(27)–2D(33))
MATERIALS STUDIED:NSM PP-12165, NSM PP-12166.
NOTE:Two compressed shoots (NSM PP-12165 and NSM PP-
12166) are 20–22 mm wide (Figs. 2D(27) and 2D(29)).
Leaves resemble the small leaf of Dúsen’s fertile branch
Fig. 2D. Araucaria sp. (27–33).(27) Shoot and leaves. NSM PP-12166 a,b. Scale bar = 10 mm. (28) Close-up of (27), showing leaf.
NSM PP-12166a. Scale bar = 5 mm. (29) Close-up of (27), showing sections of expanding leaves. NSM PP-12166b. Scale bar =
5 mm. (30) Adaxial cuticle consisting of quadrangular to irregular epidermal cells and few stomata (white arrowhead). Not all
the stomata are recognized owing to poor preservation. NSM PP-12165. Scale bar = 100 ␮m. (31) Abaxial cuticle, showing
irregular rows of stomata, separated by 1–3 rows of irregular or rectangular epidermal cells. NSM PP-12166. Scale bar = 100 ␮m.
(32) Abaxial cuticle showing variably oriented stomatal apparatus. NSM PP-12165. Scale bar = 100 ␮m. (33) Close-up of (32),
showing four subsidiary cells. NSM PP-12165. Scale bar = 50 ␮m. [Colour online.]
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of A.nathorstii (Dusén 1899, Plate XII, figs. 11 and 12). The
shoot bears densely packed imbricate leaves. The leaves
are ovate, appressed, concave, and have 7–9 rows of ab-
axial longitudinal ridges possibly corresponding to the
internal leaf veins (Fig. 2D(28)). They are not keeled, but
the middle ridge looks thicker than the lateral ones
(Fig. 2D(27)). The leaf apex is obtuse and the base is de-
current. The leaves are entire, 11–13 mm long × 6–7 mm
wide at the widest part. The leaf base is broad 2.5 mm
wide.
The leaves of Araucaria sp. described here are amphis-
tomatic, but the stomata are scarce on one adaxial cuti-
cle, where the epidermis is composed of quadrangular
to irregular cells (Fig. 2D(30)). On the abaxial cuticle
(Figs. 2D(31) and 2D(32)), stomata are arranged in irregu-
lar rows with the apertures orientated in varying direc-
tions to the leaf axis. The stomatal apparatus is elliptical,
monocyclic, and composed of mostly four but up to six
subsidiary cells (Fig. 2D(33)). Stomata of the same row are
separated by 1–3 irregular epidermal cells. Adjacent sto-
matal rows are separated by 1–3 rows of irregular or rect-
angular cells. Anticlinal walls of epidermal cells are
sometimes slightly sinuous.
Discussion
Araucaria nathorstii was described by Dusén (1899) from
the same locality where we made field collections. The
narrow type described here is comparable with Dusén’s
sterile shoot, and the broad type with his fertile shoot.
The broad type is the most typical form of A.nathorstii
which subsequently has been described widely from
southern South America and the Antarctic Peninsula
(Berry 1928;Doktor et al. 1996;Fiori 1939;Hünicken 1995;
Menéndez and Caccavari 1966;Panti et al. 2007;Troncoso
1986). The leaf of Araucaria sp. in this study is not identi-
cal with any of the leaves assigned to A.nathorstii, except
a small leaf figured in the original description as one of
the forms from a fertile A.nathorstii shoot.
In extant araucarian species, leaves often become
dimorphic, with individuals displaying differences
between the leaves produced by plants at juvenile and
adult stages (de Laubenfels 1972;Stockey 1982), and in
many cases the juvenile forms exhibit markedly nar-
rower leaves (Farjon 2010). In A.bidwillii Hook and
A.angustifolia (Bertol.) Kuntze, leaves vary a lot in length
and width according to branch positions (Farjon 2010).
Thus, the two leaf types of A.nathorstii might represent
leaves of different growth stages or of leaves borne in
different positions within the tree architecture. How-
ever, Araucaria sp. described here is quite different from
other two types in having an obtuse apex and frequent
stomata whose apertures are rather irregularly orien-
tated with respect to the leaf axis, and probably deserves
to be separated as a separate taxon.
Araucaria nathorstii has been assigned to section Araucaria
based on gross morphology and cuticle characters
(Dusén 1899;Menéndez and Caccavari 1966). The gross
morphology of the narrow and broad leaf types of our
specimens are congruent with those of the leaves of sec-
tion Araucaria. Although A.nathorstii is attributable to
section Araucaria, the absence of cone scales comparable
with that section has yet to be fully explained.
On the other hand, the obtuse leaf apex and oblique or
perpendicular orientation of stomata of this Araucaria sp.
do not conform to what is expected for sections Bunya,
Intermedia,orAraucaria (Stockey and Ko 1986). Frequent
stomata oriented obliquely or perpendicular to the leaf
axis are characteristic of section Eutacta (Stockey and Ko
1986), and some species of section Eutacta have adult
leaves with an obtuse apex (Farjon 2010). Thus, this
Araucaria sp. could be assigned to section Eutacta. Two
types of Araucarites reported here are also similar to cone
scales of section Eutacta in having thin lateral extensions.
The co-occurrence of shoots and cone scales similar to
section Eutacta suggests the presence of the section in
southernmost South America until the late Eocene or
earliest Oligocene.
Alternatively, it would also be possible that the fea-
tures compared here to characterize Eutacta are ple-
siomorphic in genus Araucaria or its extinct stem group.
Combined cladistics analysis of Araucariaceae by Escapa
and Catalano (2013) clarified monophyly of Eutacta. Al-
though only fossil permineralized cones were used for
their analysis, they revealed 5 of 62 scored characters as
unambiguous synapomorphies of the section. Among
the five characters none were related to cone scales, but
three were of leaves (No. 8, 45, and 47 in their score
sheet). The leaves of Araucaria sp. described here share
character 45 (rhomboidal or tetragonal leaves) and 47
(curved leaf apex). The lack of specimens showing cone-
scale/shoot attachment and the absence of unambiguous
synapomorphies for Eutacta cone scales bring up the pos-
sibility that our specimens could represent an extinct
basal lineage or a stem lineage between section Eutacta
and other sections. In either case, however, it is certain
that the extant section Eutacta is a remnant of an Eutacta-
like group, which would have been widely distributed in
the Southern Hemisphere before the earliest Oligocene.
Some reports of Cenozoic Eutacta from Australia and
New Zealand emphasize a need of more extensive analy-
sis including a wide range of fossil information to recon-
struct the phylogeny and biogeography of Eutacta as well
as the whole family (Hill and Bigwood 1987;Pole 2008).
The fossil record of section Eutacta is not abundant
from the Paleogene or later of South America. It has been
suggested that the fossil record of section Eutacta in
South America may extend up to the Miocene, based on
a leafy twig of Araucaria sp. from Matanzas, VI Región,
Chile, which was assigned to section Eutacta based
on external similarity to extant Araucaria heterophylla
(Troncoso and Romero 1993). Precise description of the
Matanzas specimen including cuticular anatomy, how-
Ohsawa et al. 813
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ever, is required to confirm this as a representative of
section Eutacta in South America.
Araucaria pichileufuensis from the Eocene of the Ven-
tana Formation of Río Pichileufú, Río Negro, Argentina,
is based on separate cone scales and leafy twigs (Berry
1938). Cone scales of A.pichileufuensis show prominent
lateral wings characteristic of section Eutacta. However,
the leaves of that species are acute, triangular, falcate,
and thick, about 7 mm long × 2.5 mm in maximum
width, and show intermediate characters between those
of sections Eutacta and Araucaria (Berry 1938). The Ven-
tana Formation, from which A.pichileufuensis originates,
is dated as about 47.5 Ma by
40
Ar/
39
Ar dating (Wilf et al.
2005); whereas, the Loreto Formation of Las Minas is
dated about 36.5 Ma by U–Th–Pb dating (Otero et al.
2012). Thus, the araucarian remains from Río de Las Mi-
nas are younger than those from Río Pichileufú, and they
are the youngest fossil record of the section Eutacta in
South America. Our study reveals the co-occurrence of
the relatives of sections Araucaria and Eutacta in Patago-
nia during the Eocene to early Oligocene, suggesting a
wider distribution of section Eutacta or its stem group,
and greater diversity of genus Araucaria in South Amer-
ica and the Antarctic regions during the Paleogene.
Acknowledgements
We thank Consejo de Monumentos Nacionales de
Chile and Corporación Nacional Forestal for permitting
us to collect fossil materials. Gar W. Rothwell and
Ruth A. Stockey, Oregon State University, kindly read an
earlier version of the manuscript and gave us helpful
comments. We also appreciate suggestions on scientific
names from H. Nagamasu. Comments and suggestions
from distinguished reviewers greatly improved this
work. This study was supported by KAKENHI (Grant-in-
Aid for Scientific Research) from the Ministry of Educa-
tion, Culture, Sports, Science, and Technology to H.N.
(14255007, 18405013, 24570112) and T.Y. (15H05233). This
paper is dedicated to Dr. Ruth A. Stockey, Professor
Emeritus of the University of Alberta, Canada, in honor
of her outstanding performance and contribution to pa-
leobotany for many years.
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