Cretaceous Research 110 (2020) 104426

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

A new megaflora (leaves and reproductive structures) from the

Formation (Lower Cretaceous), Peru
Huancane
L.C.A. Martínez a, b, *, E. Pacheco Huacallo d, R.R. Pujana c, H. Padula e
a
nica Darwinion (ANCEFN-CONICET). Labard

Instituto de Bota en 200, CC22 - B1642HYD - San Isidro, Buenos Aires, Argentina
b
Facultad de Ciencias Naturales y Museo. Universidad Nacional de La Plata, La Plata, Argentina
c

Museo Argentino de Ciencias Naturales-CONICET, Avda. Angel Gallardo 470, Ciudad de Buenos Aires, C1405DJR, Argentina
d
Universidad Nacional San Antonio Abad Del Cusco, Cusco, Peru
e
Direccio
n General de Patrimonio e Instituto Histo

rico de La Ciudad Auto

noma de Buenos Aires, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: We describe fossil leaves and reproductive structures remains from the Lower Cretaceous of Peru. A great
Received 1 April 2019 part of the plant fossils are very well preserved, and they come from three different stratigraphic levels of
Received in revised form the Huancane
Formation in a single locality close to Cusco City. The fossil flora is dominated by Bra-
10 February 2020
chyphyllum, Cupressinocladus, Podozamites, leaves assigned to the Araucariaceae and Araucarites Presl.
Accepted in revised form 12 February 2020
Available online 19 February 2020
Other leaf fossil-genera found are: Weichselia, Sagenopteris, Sphenopteris, and Pachypteris. This fossil
assemblage is consistent with the age inferred for the Huancane
Formation, which was based on
biostratigraphic and isotopic studies. The plant assemblage suggests a forest canopy dominated by co-
Keywords:
Lower Cretaceous
nifers, with an understore composed by ferns (Weichselia) and putative pteridosperms. These new data
Fossil leaves increase the knowledge of the palaeofloristic composition of the Cretaceous flora of Peru, contribute to
Leafy shoots the fossil record of South America, and particularly to the palaeoflora near equatorial regions.
Reproductive structures © 2020 Elsevier Ltd. All rights reserved.
Huancane
Formation
Peru

1. Introduction
Cretaceous plants from Peru have been mentioned for over 110
years. However, the palaeobotany of Peru has received little interest
and only a few studies have been published. The earliest studies
with detailed descriptions of taxa were made at the beginning of
the 20th century by foreign researchers with fossils from Mesozoic
strata (Archangelsky, 1968). The first published works on Peruvian
plant fossils are from the Lower Cretaceous (Lisso
n, 1905;
Neumann, 1907; Zeiller, 1910; Salfeld, 1911).
Fossil plants such as Pecopteris (Brongniart) Sternberg, Zamites
Brongniart and “ferns” were mentioned by Lisso
n (1905) from
Lower Cretaceous strata in San Lorenzo Island (near the port of
Callao), but without descriptions. The first descriptions were made
by Neumann (1907), who studied fossil plants collected by Captain
Paul Berthon during a military expedition. Those fossils came from
* Corresponding author.
E-mail addresses: gesaghi@gmail.com (L.C.A. Martínez), edgarhuacallopacheco@
gmail.com (E. Pacheco Huacallo), rpujana@gmail.com (R.R. Pujana), horaciopadula@
hotmail.com (H. Padula).
Caleta del Paraíso (San Lorenzo Island) and Pin ~ onate (Lima City).
The taxa described by Neumann (1907) are: Equisetites Sternberg,
Otozamites Braun, Pecopteris, Weichselia Stiehler, Zamiostrobus
Endlicher, and Rhynchogoniopsis Neumann. Salfeld (1911) described
Taeniopteris Brongniart, Sphenopteris (Brongniart) Sternberg, Cla-
dophlebis Brongniart, Weichselia, Filicites Brongniart, Glossozamites
Schimper, Brachyphyllum Brongniart and Pagiophyllum Heer from
Morococha Lagoon. Later, Zeiller (1914) re-described the fossils
studied by Neumann (1907) and recorded Weichselia and Otoza-
mites, added descriptions of new taxa as Antevsia (Antholithus)
Harris, Cladophlebis, Sphenopteris, Klukia Raciborski, Podozamites
Braun and Ruffordia Seward; and reassigned some species from
Equisetites to Weichselia, and from Zamiostrobus to Cycadolepis
Saporta. Moreover, Zeiller (1914) did not find some of the taxa
among the samples of Captain Paul Berthon (Equisetites, Zamios-
trobus, and Rhynchogoniopsis) cited by Neumann (1907).
Berry (1922) and Lisso
n (1926) summarized all the data of the
Lower Cretaceous assemblages of Peru. Besides, Berry (1922)
described new material from Pin ~ onate (near Lima), Caleta del Par-
aíso (San Lorenzo Island) and Jatunhuasi (Junín Department) and
reassigned some of the fossils studied by Zeiller (1914). New taxa
recorded from these localities are: Taeniopteris Hooker,

https://doi.org/10.1016/j.cretres.2020.104426
0195-6671/© 2020 Elsevier Ltd. All rights reserved.
2 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Sphenopteris (Ruffordia), Filicites Schlotheim ex Brongniart, Clado-
phlebis (Klukia), Zamiostrobus, Thuites Sternberg, Brachyphyllum and
Antevsia (Antholithus) Harris (Berry, 1922). Berry (1945) described
some Cretaceous fossils from Colombia and listed the taxa that join
Weichselia from Peru [e.g. Equisetites, Rufforida, Sphenopteris,
Cladophlebis (Klukia), Otozamites, Cycadolepis, Podozamites, Thuites,
(Antevsia) Antholithus, Coniopteris Brongniart, Onychiopsis
Yokoyama, Thinnfeldia Ettingshausen, Sagenopteris Presl, Pter-
ophyllum Brongniart and Brachyphyllum]. Those were the last
Mesozoic megafloras with detailed descriptions recorded from
Peru (Archangelsky, 1968). Although the first studied Mesozoic
fossils showed a great potential and diversity, the palaeobotanical
studies in the region have been disperse and discontinuous since
then.
After almost a century without Peruvian Mesozoic megaflora
studies, a new assemblage was recovered 25 km west of Cusco City
and described in detail. Descriptions are based on observations of
many specimens collected in a single locality from the Huancane
Callialasporites, Cicatricosisporites australiensis and Appendicispor-
Formation (Fig. 1). The occurrence of some taxa provides some new
palaeofloristic information at the time of the deposition of the
Huancane
Formation.
2. Geological setting
The Huancane
Formation (Newell, 1949) is a sedimentary suc-
cession mainly composed of sandstones deposited in fluvial envi-
ronments. The Huancane
sandstone is especially important because
it has characteristics of a petroleum reservoir rock (Newell, 1949).
In Cusco Region, this formation crops out from Urcos to Ollantay-
tambo localities, on flanks of the anticline outcrops of the Vilcanota
Mountains overlying the basement or it occurs as tectonic scales
(Carlotto et al., 1995).
The sediments of the Huancane
Formation overlie with
disconformity or angular unconformity on other Mesozoic units
such as Hambutío Formation (KimmeridgianeBerriasian?), or on
the Mitu Basin (Upper TriassiceLower Jurassic), and are 30 up to
300 m thick succession of quartz sandstones, conglomeratic
Fig. 1. Location map of the Cusco Department and fossiliferous locality (+).
sandstones, conglomerates and mudstones (Carlotto et al., 1995,
2004, 2011; Sempere et al., 2002).
The Huancane
Formation was divided into lower and upper
units (Carlotto et al., 1995). The lower unit (10e50 m) has con-
glomerates, conglomerate sandstones and quartz sandstones with
foreset lamination and channel deposits with the finning-upward
trend. A calcareous level (Queqayoc) or black to red mudstones
are found towards the upper part of this section. The upper
member (70e150 m) is composed by of massive sandstones (Batty
and Jaillard, 1989; Carlotto et al., 1995).
Carlotto et al. (1995) suggest a BerriasianeValanginian age to
the Huancane
Formation. This is based on stratigraphic position of
this unit respect to underlying and overlying units which possess
biostratigraphic and isotopic studies (Newell, 1949; Mendívil and
Da
vila, 1994; Carlotto et al., 1995, 2004, 2011; Sempere et al.,
2002). Palynological studies of the Huancane
Formation suggest
an Early Cretaceous age based on the record of some taxa (e.g.
ites sp.) and absence of angiosperm pollen (Doubinger and
Marocco, 1976). The palynoflora recorded from the Huancane
For-
mation resembles those present in Lower Cretaceous of Patagonia
(Archangelsky and Gamerro, 1967) and Australia (Balme, 1957).
3. Material and methods
The fossils studied are represented by impressions of leaves, leafy
axes, and reproductive structures (cone scales). The plant fossils
come from three black mudstone layers of 0.5e2 m thick, close to
Chimur x Paucartambo road, at two kilometres NW of Caicay Town,
in the Paucartambo Province, Cusco Region, Peru (13
340 4700 S,
71
420 3800 W, Fig.1). The black mudstones are part of the upper strata
of the lower section of the Huancane
Formation (Fig. 2).
Three plant-bearing levels were identified. The fossils (97
samples) were collected in 2014 and 2015 by Edgar Pacheco Hua-
callo, Leandro C. A. Martínez and Horacio Padula, and deposited in
the Palaeobotany Collection of the Universidad Nacional San
Antonio Abad del Cusco (Cusco, Peru) under the acronyms CUZ-PB
1 to CUZ-PB 97. No cuticles are preserved, so the study is mainly
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426 3

focused on direct observations of the impressions. Each rock

specimen usually bears only one taxonomic unit.
When more than one sample is assigned to a taxonomic unit,
measurements in the descriptions are given as the range or the
weighted mean followed by the range in parentheses of the sam-
ples assigned to that taxonomic unit.
For suprageneric plant taxa, the systematics of Christenhusz
et al. (2011) and Schuettpelz et al. (2016) were followed.

4. Systematic palaeobotany

Class POLYPODIOPSIDA Cronquist et al., 1966

Subclass Polypodiidae Cronquist et al., 1966
Family Matoniaceae Presl, 1847
Weichselia Stiehler, 1857
Type species: Weichselia ludovicae Stiehler, 1857
Weichselia peruviana (Neumann) Zeiller, 1914
Fig. 3.1e3.5
Specimens. CUZ-PB 4, 5, 17, 19, 22, 24, 25, 26, 28, 29, 43, 44, 53, 54,
56, 71, 95 and 97.
Description. Fronds are bipinnate. Main rachis are 0.16 mm wide.
Pinna rachis are 0.05 mm wide with pinnae alternate. Pinnae have
pinnules from opposite, sub-opposite to alternate (Fig. 3.1). Pin-
nules are variable in size, typically 0.4 mm long and 0.24 mm wide
(Fig. 3.2e3.5). There is a great variation in the shape of the pinnules
(Fig. 3.1, 3.4). Pinnules are commonly entire, straight or sometimes
falcate. Margins are entire, with blunt, subacute to an acute apex.
Pinnules are attached at right angle from rachis with the whole
basis. Adjacent pinnules are in contact between them. The upper
surface of the pinnules is convex, with the main vein marked by a
furrow and the lower surface concave. Venation reticulate. Veins
are more marked on the abaxial surface of the leaves. The main vein
disappears into the net-worked venation before reaching the apex,
the secondary veins arise from the main vein and divide to form
polygonal vein meshes (Fig. 3.4, 3.5).
Fertile pinnules have 5 (3e6) sori on the abaxial side. Sori have a
diameter of 0.72 mm (0.56e0.94), and they are apparently covered
by an indusium. The indusia are constituted by a central button-like
structure (Fig. 3.2, 3.3).
Remarks. Weichselia was erected by Stiehler (1857) based on the
material of W. ludovicae from the Lower Cretaceous of Germany.
Some species such as W. reticulata (Stokes et Webb) Fontaine in
Ward emend. Alvin and W. negevensis Silantieva et Krassilov have
fronds separated into vegetative and reproductive fronds (dimor-
phic leaves). Reproductive fronds have modified fertile pinnae with
soral clusters (Daber, 1968; Alvin, 1971; Diez et al., 2005; Silantieva
and Krassilov, 2006; Sender et al., 2015). However, W. peruviana is
clearly distinguished by its monomorphic fronds, with sori on the
abaxial surface of fertile leaves.
GYMNOSPERMS
INCERTAE ORDINIS
Sphenopteris (Brongniart) Sternberg, 1825
Type species: Sphenopteris elegans (Brogniart) Sternberg, 1825
Sphenopteris sp.
Fig. 4.1, 4.2
Specimens. CUZ-PB 48 and 93.
Description. The two specimens are small and both correspond to
sections of ultimate pinnae. Pinnate fragments are 20 mm long and

Formation (lower section) and general view
Fig. 2. Sedimentological log of Huancane 7 mm wide. Rachis is 0.5 mm thick, with a groove on the adaxial
of fossiliferous locality. surface, with closely spaced pinnules typically departing at 15e30
.
4 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426

Fig. 3. Weichselia peruviana. 1. CUZ-PB 05. General view. Bipinnate frond. Scale bar ¼ 10 mm. 2. CUZ-PB 04. Pinnae with sori. Scale bar ¼ 5 mm. 3. CUZ-PB 22. Detail of pinnae with
sori and indusia. The rounded sori have sporangia arranged around a receptacle that is the stalk of a peltate indusium (arrows). Scale bar ¼ 10 mm. 4. CUZ-PB 53. General view. Scale
bar ¼ 10 mm. 5. CUZ-PB 19. General view. Scale bar ¼ 10 mm.

Pinnules are straight, 9.0 mm long, 2.5 mm wide; attachment is
decurrent, and occasionally the acroscopic margin is slightly con-
tracted. Pinnules are dissected in two to six narrow cuneate lobes;
apex is serrated, acute, margins are slightly straight. Veins of the
pinnules fork two or three times from the midrib or the base of the
pinnule to the apex (sphenopteroid venation).
Remarks. These specimens resemble those described by Zeiller
(1914). Although, the specimens described herein are similar to
Coniopteris Brongniart, which is distinguished by the presence of
polymorph fronds.
Order CAYTONIALES Gothan
Sagenopteris? Presl in Sternberg emend. Rees, 1993
Type species: Sagenopteris acuminata Presl, in Sternberg (1838)
(lectotypified by Cleal and Rees, 2003)
Fig. 4.3, 4.4
Specimens. CUZ-PB 12, 32 and 36.
Description. Narrow ovate to lanceolate leaflets of at least 39.1
(32.5e44.0) mm long (incomplete) and 16.5 (16.0e16.8) mm wide,
with a prominent midrib and reticulate venation. Veins are prom-
inent on the lower side, flush with the surface on the upper. Leaflets
have entire margins, acute base, and apex is absent in all specimens.
Midrib is moderately conspicuous in all specimens (ca. 1.2 mm).
Lateral veins arise from midrib alternately or suboppositely, in
angles between 60
and 40
, forming a reticulum, with meshes
elongated parallel with the veins about 2.69 (1.22e3.33) mm long
and 0.39 (0.18e0.50) mm wide.
Remarks. Although there are two incomplete specimens, the
prominent midrib and reticulate venation are two diagnostic
characters of Sagenopteris.
Order CORYSTOSPERMALES Petriella
Pachypteris Brongniart emend. Harris, 1964
Type species: Pachypteris lanceolata Brongniart, 1828
Pachypteris sp.
Specimen. CUZ-PB 08.
Fig. 4.5
Description. The leaf is pinnate, the apex is absent, with pinnae and
intercalary pinnules on the main rachis. Pinnae are inserted at wide
angles (80e90
), opposite to sub-opposite. Adjacent pinnae are not
superimposed, inserted on to the main rachis by decurrent bases.
Pinnae are rounded and entire in the apex, to lobed to the base. The
lamina of lowest pinnae continues downwards as a marginal wing.
Pinnae with a midrib of straight course and lateral veins diverging
with acute angles and forked dichotomously.
Remarks. Although it is a single and incomplete specimen, some of
those characters described in this fossil resemble Pachypteris (e.g.
intercalary pinnules, marginal wing, a single vein in each pinnule)
(Archangelsky, 1963; Harris, 1964).
Subclass PINIDAE Cronquist et al., 1966
INCERTAE ORDINIS
Family Podozamitaceae Nĕmejc, 1950
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426 5

Podozamites Braun, 1843 the axils of lateral leaves; they are slender and straight and diverge
Type species: Podozamites distans (Presl) Braun, 1843 from the axes at acute angles from 30
to 45
. Ultimate leafy-
aff. Podozamites branches are closely set. Leaves are decussate, borne in whorls of
Fig. 5.1 two, scale-like, rhomboid in surface view, with an acute apex.
Specimens. CUZ-PB 33 and 35 Branchlet facial leaves are rhombic in the exposed part, closely
adpressed except for the short acute tip, 1.5 (0.5e2.2) mm long, 1.3
Description. Podozamites is represented by two incomplete speci-
(0.6e1.7) mm wide. Facial leaves lack a marked midrib; however,
mens. Bases and apices are absent. Leaves have a minimum length
leaves are thickened along the midline and may be keeled.
of 8.5 cm and 0.8 cm in width. Leaves are bilaterally symmetrical,
Remarks. The description is based on 12 specimens of sterile branch
narrow, with entire margins and covered by 16e18 longitudinal
fragments. The fragments have a length from 2 to 10 cm. This fo-
veins. The two or four middle veins are closer than the other veins
liage is similar to some genera of the Cupressaceae (Harris, 1979).
and form a slight keel.
However, shoots of Watsoniocladus Srinivasan (Cheirolepidiaceae)
Remarks. The specimens are assigned up to genus level (aff. Podo-
have phyllotaxis characters like those of Cupressinocladus, but
zamites). The fossils described here are similar to those records
distinctly a cheirolepidiaceous cuticle morphology (Srinivasan,
from Lower Cretaceous of Peru described by Berry (1945). However,
1995).
all specimens lack leaf bases and apices, therefore they also could
be assigned to other similar taxa, such as Desmiophyllum Order ARAUCARIALES Gorozhankin
Lesquereux (1878), Eretmophyllum Thomas (1913) and Lin-
Incertae families
dleycladus Harris (1979).
Genus incertae sedis
Incertae families
Fig. 6.1e6.4
Brachyphyllum Brongniart, 1828 Specimens. CUZ-PB 1, 2, 59 and 65.
Type species: Brachyphyllum mamillare Brongniart, 1828
Description. The fossils are impressions of branches with leaves.
Brachyphyllum sp. Shoots are up to 7.5 cm long and 0.2 cm wide (Fig. 6.1). Shoots are
Fig. 5.2, 5.3 robust and bear simple leaves, opposite or sub-opposite, distichous
Specimens. CUZ-PB 14, 16, 21, 23, 30 and 58 and twisted to present adaxial surfaces uppermost, spreading 36

(27
e45
) to the axis (Fig. 6.1, 6.2). Shoots usually show some ver-
Description. Vegetative shoots with helically arranged leaves.
tical striations (Fig. 6.2). Leaves are 3.0 (1.7e4.5) cm long and 0.4
Largest branch is 3 cm long. Leaves are small, 3.2 (1.7e5.8) mm long
(0.2e0.5) cm wide. Leaves appear to be thick, simple, triangular-
and 1.2 (0.5e1.8) mm in basal diameter. Leaves have an acute apex,
lanceolate, with the apex slightly acute, strongly decurrent base,
often projecting towards branch axis and accuminate. Venation was
entire margins and multinerved. Veins that enter the leaf from the
not observed.
shoot are parallel of equal prominences (Fig. 6.4). Some shoots
Comparisons. The taxonomic relation of Brachyphyllum is unclear at
show gradation from proximal broad-based, hemispherical and
the family level. Brachyphyllum is considered an unnatural assem-
small leaves to distal lanceolate leaves (Fig. 6.2).
blage by many authors. These leafy shoots are typical of some
Comparisons. These Cretaceous fossil leaves are comparable to some
Jurassic and Cretaceous conifers, and the majority of its fossil-
living and fossil leaves. Living conifers have foliage generally small
species have been assigned to the Araucariaceae or the Podo-
with a single vein, from needle-like to flattened. However, there are
carpaceae, whilst other species have been related to the Cheir-
some exceptions (e.g. Agathis Salisbury, Nageia Gaertner, and
olepidiaceae (Archangelsky, 1963; Harris, 1979; Stockey, 1982).
Phyllocladus Mirbel). The leaves described here resemble those
Remarks. The description is based on six specimens of sterile branch
present in the Araucariaceae, and in particular Wollemia rather than
fragments. The specimens are assigned up to genus level (Brachy-
Araucaria Jussieu or Agathis. Some of the characters shared between
phyllum) due to their lack of cuticle. However, these specimens are
the fossils and Wollemia are: robust shoots with simple distichous
similar in morphology to B. mucronatum Archangelsky and
leaves, opposite or sub-opposite and twisted to present adaxial
B. mirandai Archangelsky from the Lower Cretaceous of Patagonia
surfaces uppermost; leaves triangular-lanceolate with apex slightly
(Archangelsky, 1963). Some specimens of Brachyphyllum sp. from
acute, strongly and decurrent base, entire margins, and numerous
the Huancane
Formation have unusually long leaves that overlap
parallel veins (Jones et al., 1995; Pastoriza-Pin ~ ol, 2007). Besides,
the boundary of Pagiophyllum (Heer) Harris. These features were
Wollemia has a remarkable variety between juvenile and adult
mentioned by Harris (1979) in certain species of Brachyphyllum (e.g.
leaves; the juvenile leaves possess decussate to opposite phyllo-
B. crucis and B. muensteri). The sole distinction of Pagiophyllum from
taxis, gradation from proximal broad-based leaves to distal lance-
Brachyphyllum is its longer free leaf, and occasional specimens of
olate leaves (Chambers et al., 1998). The juvenile features of
both genera transgress the arbitrarily defined boundary (Harris,
Wollemia are recorded in some of the fossil leaves described here
1979).
(Fig. 6.2).
Incertae familis Comparisons among these leaves and other fossil taxa are also
complicated. The leaves described here are comparable with
Cupressinocladus Seward, 1919
Podozamites (Braun) emend Shi et al. (2018). However, they differ
Type species: Cupressinocladus massiliensis (Saporta) Seward (Thu-
from the fossils here described, because the leaves of Podozamites
jopsis massiliensis Saporta), typ. cons. prop.
are inserted on a slender axis. On the other hand, some Cretaceous
Cupressinocladus sp. Gondwana leaves are assigned to Podozamites, and placed in the
Fig. 5.4, 5.5 Araucariaceae (Cantrill, 1991).
Specimens. CUZ-PB 3, 6, 7, 9, 18, 20, 39, 46, 60, 61, 63 and 83. The Huancane
fossils share features with the Araucariaceae, and
their foliage resemble in some juvenile characters to extant Wol-
Description. The fragments have a length from 2 to 10 cm. Vegeta-
lemia, but the absence of cuticular features prevents making a more
tive shoots with decussate leaves, flattened, forming sprays of
precise taxonomic assignment.
branches and branchlets that do not overlap. The largest specimen
Remarks. The description is based on three specimens of sterile
is 10 cm long and bears 25 pairs of branchlets. Branchlets arise in
branch fragments. Fragments are not longer than 8.8 cm.
6 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426 7

Fig. 5. Conifer leaves. 1. aff. Podozamites. Scale bar ¼ 5 mm. 2. CUZ-PB 14. Brachyphyllum sp. Scale bar ¼ 5 mm. 3. CUZ-PB 16. Brachyphyllum sp. Scale bar ¼ 5 mm. 4. CUZ-PB 3.
Cupressinocladus sp. Scale bar ¼ 10 mm. 5. CUZ-PB 9. Cupressinocladus sp. Scale bar ¼ 10 mm.

Family ARAUCARIACEAE Henkel and Hochstetter, 1865
Araucarites Presl (in Sternberg, 1838)
Type species: Araucarites goeppertii Presl (in Sternberg, 1838)
Araucarites pachacuteci Martínez sp. nov.
Figs. 6.5, 7.1, 7.2
Holotype. CUZ-PB 41. Fig. 7.1.
Paratypes. CUZ-PB 10, 11, 31, 10, 69, 75, 81, 86, 88, 90, 91 and 96.
Derivato nominis. The species name was given in honour to
Pachakutiq Inka Yupanki (Quechua) was the ninth Sapa Inca
(1418e1471/1472?) of the kingdom of Cusco which he transformed
into the Inca Empire.
Diagnosis. Small detached ovulate bract-scale complexes. Cone
scales cuneate, ca. 10e20 mm long (from the base to the apex of the
ligule) by ca. 4e10 mm wide in the widest point. Straight and small
wings expanded laterally from each side of the central zone of cone
scale. Ligule of ca. 2e5 mm long and ca. 1e3 mm in diameter.
Ovuliferous scale clearly distinct, almost overlying the bract in
extension. Oval ovule in the proximal part of bract, of ca. 6e11 mm
long and ca. 2e4 mm wide.
Type locality and horizon. Chimur x Paucartambo road, 2 km NW of
Caicay locality, in Paucartambo Province, Cusco Region, Peru
(13
340 46.8000 S 71
420 38.4800 W); Type stratum: Huancane
Forma-
tion, BerriasianeValanginian (Lower Cretaceous).
Description. Small ovulate bract-scale complexes detached and
composed by an ovuliferous scale, which overlies a winged bract.
Bracts are cuneate, with margins diverging from the base to their
widest point. The bract has a length of 14.8 (11.0e18.5) mm from
the base to the apex of ligule. The base is narrow with a diameter of
2.8 (2.2e4.1) mm, and it expands distally to 5.8 (4.3e9.3) mm in the
widest point (ca. 60e65% of distance from the base). The distal part
of the complex is rounded and bears a ligule of 3.2 (1.9e4.7) mm
long and 2.0 (1.2e3.2) mm in diameter (Figs. 6.5, 7.1). Straight,
smooth and small wings are on each side of the central seed-
bearing zone. Ovuliferous scale is clearly distinguished from the

Fig. 4. Pteridosperms? 1. CUZ-PB 93. Sphenopteris sp. General view. Scale bar ¼ 5 mm. 2. CUZ-PB 48. Sphenopteris sp. General view. Scale bar ¼ 5 mm. 3. CUZ-PB 12. Sagenopteris sp.
General view. Scale bar ¼ 10 mm. 4. CUZ-PB 32. Sagenopteris sp. General view. Scale bar ¼ 5 mm. 5. CUZ-PB 08. Pachypteris sp. General view. Intercalary pinnules (arrows). Scale
bar ¼ 10 mm.
8 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426

Fig. 6. Araucariaceae leaves and Araucarites pachacuteci. 1. CUZ-PB 1. Araucariaceae incertae sedis leaves. General view. Scale bar ¼ 1 cm. 2. CUZ-PB 59. Araucariaceae incertae sedis
leaves. Scale bar ¼ 1 cm. 3. CUZ-PB 2. Araucariaceae incertae sedis leaves. Decurrent base (arrow). Scale bar ¼ 0.5 cm 4. CUZ-PB 2. Araucariaceae incertae sedis leaves. Detail of
parallel veins. Scale bar ¼ 1 cm. 5. CUZ-PB 96. Araucarites pachacuteci nov. sp. Scale bar ¼ 0.5 cm.

bract, with a diameter of 4.88 (3.43e8.39) mm, which almost
overlies the bract in extension (Fig. 7.1). The ovule is located close to
the proximal part of bract, and it is defined by the presence of two
longitudinal grooves that delineate it from the ovuliferous scale.
The ovule has a length of 7.8 (6.7e10.9) mm and a diameter of 2.3
(1.8e3.4) mm.
Comparisons. These kind of ovuliferous complexes are abundant in
the fossil record. They are usually assign to the Araucariaceae and
compared with araucariaceous ovulate cones and cone-scale
complexes (Zijlstra and van Konijnenburg-van Cittert, 2006). The
genus Araucaria is divided into six sections based on foliar and
ovulate complex characters. Four of them have living species:
Eutacta, Intermedia, Bunya and Araucaria (or Columbea); whereas
Yezonia and Perpendicula sections only have fossil-species (Wilde
and Eames, 1952; Ohsawa et al., 1995; Pole, 1995; Dettmann and
Clifford, 2005).
The fossils studied here are characterized by cone-scale com-
plexes thinly winged with the seed embedded in the scale, and
apparently covered by its surface. This feature is typical of the
Eutacta section. On the other hand, bract-scale complexes of Bunya
and Intermedia sections are distinguished from our fossils by the
presence of wide wings, whereas those present in Araucaria section
have bract-scale complex without wings (Wilde and Eames, 1952;
Dettmann and Clifford, 2005).
Araucarites pachacuteci was compared with other fossil cone-
scale complexes such as Araucaria cutchensis (Feistmantel) Pant et
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Fig. 7. Araucarites pachacutesis (Holotype). 1. CUZ-PB 41. Scale bar ¼ 5 mm. l (ligule), b (bract), os (ovuliferosus scale), o (ovule). 2. CUZ-PB 75. Scale bar ¼ 5 mm.
Srivastava, A. lefipanensis Andruchow Colombo, Escapa, Cúneo et
Gandolfo, Araucarites alatisquamosus Ohsawa et H. Nishida,
A. antarcticus (Gee) Birkenmajer et Ociepa, A. aquiensis Fontaine,
A. baqueroensis Archangelsky, A. brodei Carruthers, A. charcotii
Harris, A. chilensis Baldoni, A citadelbastionensis Cantrill et Falcon-
Lang, A. cuneatus Ward, A. kujiensis Tanai, A. minimus Arch-
angelsky, A. minutus Bose et Maheshwari, A. parsorensis Lele,
A. philipsi Carruthers, A. rogersii Seward, A. roginei Crie
,
A. sehoraensis Bose et Maheshwari, A. stockeyi Tidwell et Ash;
A. vulcanoi Duarte, A. wollemiaformis Cantrill et Falcon-Lang,
A. wyomingensis Ward (Carruthers, 1869; Crie
, 1877; Fontaine,
1889; Ward, 1889; Seward, 1903; Harris, 1935; Lele, 1956;
Archangelsky, 1966; Pant and Srivastava, 1968; Bose and
Maheshwari, 1973; Baldoni, 1979; Tanai, 1979; Duarte, 1989, 1993;
Cantrill and Falcon-Lang, 2001; Tidwell and Ash, 2006; Birken-
majer and Ociepa, 2008; Ohsawa et al., 2016; Andruchow Colombo
et al., 2018).
Some Triassic species such as Araucarites nordenskioeldii Heer,
A. convexus Brik, A. carolinensis Fontaine, A. migayi Schvedow and
A. sydneyensis Walkom (Heer, 1874; Walkom, 1925; Brik, 1952;
Schvedow, 1958) were excluded from the comparisons because
those taxa were transferred to other fossil-genera (e.g. Lepacyclotes
Emmons and Cylostrobus Helby et Martin) (Retallack, 1997).
The species of Araucarites without wings (A. cuneatus, A. wolle-
miaformis) as well as those with well development wings
(A. lefipanensis, A. alatisquamosus, A. baqueroensis, A. brodiei; A.
chilensis, A. minimus, A. parsorensis, A. philipsi, A. stockeyi) are clearly
distinct from A. pachacuteci. Other species such as
A. citadelbastionensis and A. wyomingensis have thin wings, whereas
A. aquiensis and A. sehoraensis lack ligule, unlike A. pachacuteci. The
ovuliferous complexes of A. charcotii, A. cutchensis, A. rogersii and
A. vulcanoi differ in their larger size. The cone scales of A. antarcticus
and A. minutus are wider than A. pachacuteci.
The comparisons among A. kujiensis and A. pachacuteci are
difficult, because A. kujiensis is a compressed cone, probably
immature (Tanai, 1979), and have not got a wing characters to
9
establish accurate comparisons. However, the A. kujiensis is smaller
than A. pachacuteci.
Comparisons among Araucarites pachacuteci and cone scales of
araucarian affinity are resumed in Table 1.
5. Discussion
5.1. Fossil taxa
The fossil flora of the Huancane
Formation has nine taxonomic
units of pteridophytes, pteridosperms, and conifers.
The fossil conifers are the most abundant leaves, with the
following taxa: Brachyphyllum, Podozamites, Cupressinocladus,
Araucarites and other leaves of Araucariales affinity.
Among the conifer remains, Brachyphyllum is typical and
widespread in the Mesozoic. It could represent different families:
Araucariaceae, Podocarpaceae or Cheirolepidiaceae (Harris, 1979;
Anderson et al., 2007; Tosolini et al., 2013). Cupressinocladus is
also a common taxon found in Mesozoic and Cenozoic deposits
which is related to the Cupressaceae or Cheirolepidiaceae. Podo-
zamites is another taxon assigned to different families (Shi, 2018).
Some authors consider it to be the leaf of a voltzialean conifer
(Harris, 1935), of the Araucariaceae (Cantrill, 1991), of the Podo-
carpaceae (Kimura et al., 1988), or even as a taxon of a particular
family, the Podozamitaceae (Nĕmejc, 1950).
Some leaves incertae sedis (Fig. 6) described here have arau-
carian features, but could not be assigned with confidence to a fossil
or living taxon (e.g. Agathis Salisb. and Araucaria Juss.). However,
these leaves resemble in some juvenile characters those present in
extant Wollemia (Jones et al., 1995; Chambers et al., 1998; Pastoriza-
Pin~ ol, 2007; Tomlinson and Murch, 2009).
Furthermore, this new record from the Lower Cretaceous of Peru
is similar to other fossil shoots, which show affinities with Wollemia
from the Cretaceous of Australia (Chambers et al., 1998). Besides,
there are several records allied to Wollemia from the Cretaceous of
10 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Gondwana. Cretaceous seed cones such as Wairarapaia Cantrill &
Raine from New Zealand and Emwadea microcarpa Dettmann,
Clifford & Peters from Australia have a cone structure and seeds
similar to Agathis and Wollemia (Cantrill and Raine, 2006; Dettman
et al., 2012). Both fossil-genera are located as the stem of the aga-
thioid clade in phylogenetic analyses of extant and fossil Araucar-
iaceae (Escapa and Catalano, 2013). Pollen grains of Dilwynites
Harris have been recorded from the Late Cretaceous (Turonian) to
the Neogene in several localities of Australia, New Zealand,
Antarctica and South America (Macphail et al., 1995, 2013).
Currently, pollen grains of Dilwynites are supposed to be related to
Wollemia and Agathis (Macphail, 2007). All these records suggest
that Wollemia had a wider distribution across Gondwana in the
past. Another possibility is that these leaves correspond to a new
extinct taxon of the Araucariaceae with leaves similar to Wollemia.
Future studies with more fossils will surely clear up the assignment
of these fossil leaves.
In addition, the cone scales recorded from the Huancane
For-
mation and assigned to Araucarites pachacuteci have a prominent
distal spine that gives them a distinctly araucarian appearance and
confirms the presence of Araucaria section Eutacta. This section is
the most diverse today, with 15 species distributed in New Cale-
donia, New Guinea, Australia and Norfolk Island (Farjon and Filer,
2013). However, Mesozoic Araucaria had a broader worldwide
distribution, and they were especially diversified in South America,
with species assigned to Araucaria, Bunya and Eutacta sections
(Archangelsky, 1966; Kunzmann, 2007; Dutra and Stranz, 2009;
Archangelsky and Del Fueyo, 2010; Panti et al., 2012). On the
other hand, cone scales of Wollemia share with some species of
Araucaria section Eutacta the presence of obtriangular shape,
papery lateral wings, and prominent woody distal projection
(Chambers et al., 1998).
Focusing on the record of the bract-scale complex Araucarites
pachacuteci and the incertae sedis araucarian leaves, three hypoth-
eses arise. The first hypothesis is that Araucarites pachacuteci is
related to Araucaria section Eutacta. The second hypothesis is that
A. pachacuteci and the incertae sedis araucarian leaves are related to
the Wollemia linage. A third hypothesis is that the araucarian leaves
incertae sedis were part of a plant unrelated to the known sections
of the Araucariaceae.
Other fossils recorded from the Huancane
Formation are scanty
and fragmentary specimens, like those assigned to pteridosperms
(Pachypteris, Sphenopteris, and Sagenopteris). The distinctive and
most common fern found in the studied locality is Weichselia
peruviana.
5.2. Palaeoenvironment
Based on sedimentological evidence and the taxonomic
composition of the recorded flora, the Huancane
Formation was
deposited in terrestrial and fluvio-lacustrine environments, sur-
rounded by forest canopies dominated by conifers and with the
understore taxa composed by ferns (mainly Weichselia) and pteri-
dosperms. However, some of the fossils here studied might be
allochthonous. Many plant remains recorded are small and scanty
(e.g. Brachyphyllum, Sphenopteris and Sagenopteris), and they could
have been transported over relative long distances, easily frag-
mented and destroyed. On the other hand, the abundance of well-
preserved fossils such as Weichselia, Cupressinocladus, Araucarites
and araucarian leaves from the same levels suggests little transport
prior to burial or that they may came from a nearby source.
Therefore, the plants recorded may be a mixture from different
sources, and not necessarily be the dominant vegetation in the
studied locality.
Several authors (Seward, 1900; Gothan, 1910; Edwards, 1933;
Alvin, 1971) have discussed the ecology of Weichselia. Some of
them believed that Weichselia grew in an arid dune region because
most of the specimens occurred in sandstone and the morphology
of the plant suggests it is a xerophyte (Alvin, 1971). Daber (1968),
based on their sedimentological data, suggested that the Weichselia
reticulata from the Lower Cretaceous of Germany, grew in a mari-
time setting on wet ground, at the latest stage of dune develop-
ment, with the attainment of heath and meadow communities. The
specimens of Weichselia reticulata from Saudi Arabia (Late Creta-
ceous) probably grew in an arid dune region near the sea and its
remains were deposited by water in back shore lagoons and
swamps (El-Khayal, 1985). The stems of Paradoxopteris Hirmer, in
connection to Weichselia, also possesses xeromorphic features such
as sclerotic tissue, the development of a large-celled pericycle
probably for water-storage, and the presence of mucilage canals
(Edwards, 1933). However, the sedimentological evidence suggests

Formation have grown
that fossils of Weichselia from the Huancane
in low lands close to coastal areas.
5.3. Age of the Huancan

e Formation
Newell (1949) originally proposed an Early or middle Cretaceous
age for the Huancane
Formation based on its stratigraphic position.
Later, biostratigraphic and isotopic studies suggested an Early
Cretaceous age (Newell, 1949; Doubinger and Marocco, 1976;
Mendívil and Da
vila, 1994; Carlotto et al., 1995, 2004, 2011;
Sempere et al., 2002). Some of the taxa described here indicate
an age not older than earliest Early Cretaceous, in accordance to the
age proposed previously. Weichselia was recorded from the Lower
Jurassic to the Upper Cretaceous. However, it had its wider
worldwide distribution (except Australia and Antarctica) during the
Early Cretaceous (Silantieva and Krassilov, 2006; Sender et al.,
2015). Sagenopteris has a similar history, with its first records in
the Triassic, but with a worldwide distribution, mainly in Jurassic
and Cretaceous strata (Edwards, 1933; Berry, 1945; Barale, 1979;
Diez et al., 2005; Taylor et al., 2006; Kujau et al., 2013; Pattemore
et al., 2015; Sender et al., 2015).
Weichselia, Sphenopteris, Sagenopteris, Podozamites and Brachy-
phyllum were also recorded in some other Cretaceous localities of
Peru (Neumann, 1907; Zeiller, 1914; Berry, 1922, 1945); whereas
other taxa are recorded for the first time (e.g. Cupressinocladus,
Araucarites and Pachypteris).
5.4. Comparisons with Early Cretaceous floras of South America
The Late Jurassic and Early Cretaceous floras of Gondwana are
generally interpreted as having certain uniformity because of the
wide distribution of certain taxa (e.g araucarian, podocarp, cheir-
olepidacean and taxodiacean conifers, caytonialeans, bennettita-
leans and ginkgoaleans) in Gondwana, and because of the absence
of important physical barriers to plant migration (McLaughlin,
2001). Besides, the global climate change throughout the Jurassic
and Early Cretaceous seems to have been minimal (Hallam et al.,
1993; Rees et al., 2000; Philippe et al., 2004). However, Rees et al.
(2000) divided the world into five climate zones (summer wet,
desert, winter wet, warm temperate and cool temperate zones),
and indicate an evident provincialism in Gondwana during this
time (McLaughlin, 2001).
South America has some well-studied Early Cetaceous floras
ranging from Patagonia to northern South America (Salfeld, 1911;
Berry, 1922, 1945; Lipps, 1938; Royo y Go
mez, 1945:; Bürgl, 1960;
Huertas, 1970, 1976; Shoemaker, 1982; Pons, 1983; Archangelsky,
2001; Lima et al., 2012; Del Fueyo et al., 2007; Martill et al.,
2007; Mohr et al., 2007, 2015; Monje Duss
an et al., 2016; Moreno
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426 11

Table 1
Comparison among Araucarites pachacuteci sp. nov. and cone scales of araucarian affinity (Araucaria and Araucarites).

Species Length (mm) Width (mm) Ligule (mm) Wings Ovule Period/Epoch

Length (mm) Width (mm)

Araucaria cutchensis 23e40 9e15 4 Present and developed 8 4e10 Jurassic eEarly
(Feistmantel) Pant and Cretaceous
Srivastava (1968)
Araucaria lefipanensis 16.7 (14.9e18.3) 15.2 (11.5e20) 2.1 (1.8e2.7) Present and well developed 9.9 (8.7e10.9) 4.9 (4.2e5.9) Late Cretaceous
Andruchow-Colombo et al.
(2018)
Araucarites alatisquamosus 32e33 30e34 3 Present and well developed 18e24 5e5.5 Late Eocene e early Oligocene
Ohsawa et H. Nishida
(Ohsawa et al., 2016)
Araucarites antarcticus (Gee) 7e18 5e15 1.5e3 Present and developed 6e10 1.5e5 Middle Jurassic
Birkenmajer and Ociepa
(2008)
Araucarites aquiensis Fontaine 50e35 30e15 Absent Present and developed ? ? Early Cretaceous
(1889)
Araucarites baqueroensis 20e30 20 10 Present and well developed 10e15 4e5 Early Cretaceous
Archangelsky (1966)
Araucarites brodiei Carruthers 15.4e21.2 11.6e23.2 Present Present and well developed ? ? Middle Jurassic
(1869)
Araucarites charcotii Harris 20 8 Present? Present and developed ? ? Late Triassic-Early
(1935) Jurassic
Araucarites chilensis Baldoni 11 8 1 Present and well developed 7 3.5 Early Cretaceous
(1979)
Araucarites citadelbastionensis 11e15 7e12 2 Present and thin 7 3 Early Cretaceous
Cantrill and Falconelang
(2001)
Araucarites cuneatus Ward 20 9 Absent Absent ? ? Early Cretaceous
(1889)
Araucarites kujiensis Tanai 10e12 ? Present ? 5e7 ? Late Cretaceous
(1979)
Araucarites minimus 7 8 3 Present and well developed 3.5e5 2e2.5 Early Cretaceous
Archangelsky (1966)
Araucarites minutus Bose and 10e15 8e11 1e2 Present and developed 8e10 3e6 Mid- Late Jurassic?
Maheshwari (1973)
Araucarites pachacuteci sp. 14.8 (11e18.5) 5.8 (4.3e9.3) 3.2 (1.9e4.7) Present and developed 6.7e10.9 1.8e3.4 Early Cretaceous
nov.
Araucarites parsorensis (Lele, 12 11 Present? Present and well developed ? ? Late Triassic
1956)
Araucarites philipsi Carruthers 11.6e15.4 11.6 Absent Present and well developed ? ? Early Jurassic
(1869)
Araucarites rogersii Seward 30 30 Present Present and developed ? ? Early Cretaceous
(1903)
Araucarites sehoraensis Bose 13e20 11e13 Absent Present and developed 10e12 6e9 Mid- Late Jurassic?
and Maheshwari (1973)
Araucarites stockeyi Tidwell and 11e12 8e12 1 Present and well developed 8 5e6 Jurassic
Ash (2006)
Araucarites vulcanoi Duarte 22 16 ? Present and developed 15 7 Early Cretaceous
(1989)
Araucarites wollemiaformis 35 13e20 12 Absent 7 8 Early Cretaceous
Cantrill and Falconelang
(2001)
Araucarites wyomingensis Ward 4e5 3e3.5 ? Present and thin ? ? Early Cretaceous
(1889)

nchez et al., 2007; Lindoso et al., 2018). Most of them were found
Sa
in the Andean region (Fig. 8) and there is a remarkable absence of
Early Cretaceous floras between Patagonia and Peru and only a few
in extra Andean regions.
The Early Cretaceous floras of Bolivia, Brazil, Peru and Colombia
share some taxa with that of the Huancane
Formation, such as
Weichselia, Brachyphyllum, Cupressinocladus and Araucarites
(Salfeld, 1911; Berry, 1922, 1937; Pons, 1983; Mussa et al., 2000; van
Waveren et al., 2002; Monje Dussa
n et al., 2016). It is worth
mentioning that some of the taxa from the Huancane
Formation are
also present in Early Cretaceous floras of Patagonia, such as Sphe-
nopteris, Pachypteris, Brachyphyllum and Araucarites (Archangelsky,
1963, 1966; Archangelsky et al., 2009; Del Fueyo et al., 2007;
Passalía, 2007a, b). However, cycads and bennettitals are absent
among the fossils of the Huancane
Formation. The abundance and
diversity of cycads and bennettitals characterize southern South
American and Antarctic Peninsula Early Cretaceous floras (Patago-
nia-Palmer sub-province) (McLoughlin, 2001), although some were
found in the Lower Cretaceous of Ecuador (Shoemaker, 1982). On
the other hand, Weichselia was not recorded in Patagonia. Thus, it
can be hypothesized that the latitudinal and/or climatic differences
might be the principal cause of the differentiation of these floras.
By the end of the Early Cretaceous (AptianeAlbian), the globally
warmer conditions and the appearance of angiosperms are the
most significant events that clearly distinguished other Early
Cretaceous floras of South America in Brazil (Crato and Codo
For-
mations), Colombia (Yaví Formation) and Argentina (Río Mayer,
Punta del Barco, Anfiteatro de Tico
, Piedra Clavada and Kachaike
Formations) (McLoughlin, 2001; Del Fueyo et al., 2007; Martill
12 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Fig. 8. Most studied/important South American palaeofloras (approximately with
more than five species or taxonomic units mentioned) from the Early Cretaceous. 1,
Kachaike Fm. (Del Fueyo et al., 2007; Pasalia, 2007a; b), 2, Baquero
group
(Archangelsky, 2001 and references therein), 3, Springhill Fm. (Del Fueyo et al., 2007),
4, Crato Fm. (Lima et al., 2012 and references therein; Mohr et al., 2015), 5, Paranaiba
(Lindoso et al., 2018), 6, Huallanca (Salfeld, 1911), 7, Lima surroundings (Neumann,
1907; Zeiller, 1914; Berry, 1922), 8, Puyango (Shoemaker, 1982), 9, valle superior del
Magdalena (Monje Duss
an et al., 2016), 10, Río Bata and surroundings (Royo and
Gomez, 1945; Bürgl, 1960), 11, Río Upín (Berry, 1945), 12, Villa Leyvas (Huertas, 1970,
1976; van Waveren et al., 2002; Moreno S
anchez et al., 2007), 13, Pa
ramo de Chita
(Lipps, 1938).
et al., 2007; Mohr et al., 2007; Archangelsky et al., 2009; Monje
Duss
an et al., 2016; Lima et al., 2012; Lindoso et al., 2018).
6. Conclusions
Some of the fossils studied are fragmentary or are very poorly
preserved. However, taxa such as Cupressinocladus, Araucarites and
Pachypteris are firstly recorded in Peru, strengthening the Early
Cretaceous age proposed for the Huancane
Formation by previous
geological and palynological data. The specimens of Weichselia are
the most frequent and well preserved, suggesting little transport
before burial and that they probably dominated the past vegetation
of the region.
Focusing on the records of the bract-scale complex Araucarites
pachacuteci and the incertae sedis araucarian leaves, they coincide
with the broader distribution and diversification of the Araucar-
iaceae during the Mesozoic, particularly during the Cretaceous.
Besides, the taxa of Huancane
suggest relationships with fossil
floras of Bolivia, Brazil and Colombia (e.g. Weichselia,
Brachyphyllum, Cupressinocladus and Araucarites), Patagonia (e.g.
Brachyphyllum, Pachypteris, Araucarites spp.) and Australasia
(Araucaria section Eutacta).
Finally, these new data from the Huancane
Formation expose
the fossiliferous potential of Mesozoic strata of Peru and contribute
to the fossil record of South America, and particularly to the
Cretaceous flora distribution near equatorial regions.
Acknowledgements
We would like to express our very great appreciation to R.
Barboni and L.M. Tejada Medina for providing us with useful
literature resources. The authors wish to acknowledge Eduardo
Koutsoukos (Editor) and the two reviewers (Dr Alexei B. Herman
and one anonymous reviewer) for their constructive comments
which allowed us to improve this paper.
References
Alvin, K.L., 1971. Weichselia reticulata (Stokes et Webb) Fontaine from the Wealden
of Belgium. Memorial Institute Sei. Nat. Belgique 166, 3e31.
Anderson, J.M., Anderson, M.H., Cleal, C.J., 2007. Brief History of the Gymnosperms:
Classification, Biodiversity, Phytogeography and Ecology. Strelitzia 20. National
Botanical Institute, Pretoria, p. 280.
Andruchow-Colombo, A., Escapa, I.H., Cúneo, N.R., Gandolfo, M.A., 2018. Araucaria
lefipanensis (Araucariaceae), a new species with dimorphic leaves from the Late
Cretaceous of Patagonia, Argentina. American Journal of Botany 105,
1067e1087. https://doi.org/10.1002/ajb2.1113.
Archangelsky, S., 1963. A new Mesozoic flora from Tico
, Santa Cruz Province,
Argentina. Bulletin of the British Museum (Natural History) Geology 8 (2),
45e92.
Archangelsky, S., 1966. New gymnosperms from the Tico
flora, Santa Cruz Province,
Argentina. Bulletin of the British Museum (Natural History) 13, 259e295.
Archangelsky, S., 1968. Palaeobotany and palynology in South America: A historical
review. Review of Palaeobotany and Palynology 7, 249e266.
Archangelsky, S., 2001. Evidence of an Early Cretaceous floristic change in Patago-
nia, Argentina. Publicacio
n Especial de la Asociacio

n Paleontolo

gica Argentina 7,
15e19.
Archangelsky, S., Del Fueyo, G.M., 2010. Endemism of Early Cretaceous Conifers in
Western Gondwana. In: Gee, C.T. (Ed.), Plants in Mesozoic Time: Innovations,
Phylogeny, Ecosystems (Ted Delevoryas Festschrift). Indiana University Press,
Bloomington, pp. 247e268.
Archangelsky, S., Gamerro, J.C., 1967. Pollen grains found in coniferous cones from
the Lower Cretaceous of Patagonia (Argentina). Review of Palaeobotany and
Palynology 5, 179e182.
Archangelsky, S., Barreda, V.D., Passalia, M.G., Gandolfo, M.A., Pra
mparo, M.B.,
Romero, E., Cúneo, R., Zamuner, A., Iglesias, A., Llorens, M., Puebla, G.G.,
Quattrocchio, M., Volkheimer, W., 2009. Early angiosperm diversification: evi-
dence from southern South America. Cretaceous Research 30, 1073e1082.
Baldoni, A.M., 1979. Nuevos elementos paleofloristicos de la tafoflora de la For-
macio
n Spring Hill, limite Jura
sico-Cret
acico subsuelo de Argentina y Chile
Austral. Ameghiniana 16, 103e119.
Balme, E.B., 1957. Spores and pollen grains from the Mesozoic of Western Australia.
Commonwealth Scientific and Industrial Research Organization, Coal Research
Organization. Technical Communication 25, 1e48.
Barale, G., 1979. Decouverte de Weichselia reticulata (Stokes & Webb) Fontaine
emend. Alvin, filicinee leptosporangiee, dans le Cretace inferieur de la Province
de Lerida (Espagne): Implications stratigraphiques et paleoecologiques. Geobios
12, 313e319.
Batty, M., Jaillard, E., 1989. La sedimentacio
n neocomiana (Jura
sico
terminaleAptiano en el Sur del Perú. In: Spalletti, L.A. (Ed.), Contribuciones de
los simposios sobre el Cret
acico de Ame
rica Latina. Buenos Aires, pp. A75eA88.
Berry, E.W., 1922. The Mesozoic flora of Peru. In: Mathews, E.B. (Ed.), Contributions
to the Paleobotany of Middle and South America, vol. 4. John Hopkins Univer-
sity Studies in Geology, pp. 44e65.
Berry, E.W., 1937. On the presence of the fern Weichselia in Colombia, South
America. Journal of the Washington Academy of Sciences 27, 458e461.
Berry, E.W., 1945. The Weichselia Stage in the Andean geosyncline. Johns Hopkins
University - Studies in Geology 14, 151e169.
Birkenmajer, K., Ociepa, A.M., 2008. Plant-bearing Jurassic strata at Hope Bay,
Antarctic Peninsula (West Antarctica): geology and fossil-plant description.
Studia Geologica Polonica 128, 5e96.
Bose, M.N., Maheshwari, H.K., 1973. Some detached seeds belonging to Araucar-
iaceae from the Mesozoic rocks of India. Geophytology 3, 205e214.
Braun, C.F.W., 1843. Beitra €ge zur Urgeschichte der Pflanzen. Beitr€ age zur Petre-
factenkunde 6, 1e46.
Brik, M.I., 1952. Iskopaemaya flora i stratigraphiya niznemezozo- iskikh otlozhenii
basseina srednego R. Ilek v zapadnom kazachstane (Fossil flora and stratigraphy
of lower Mezozoic deposits of the central Ilek River in western Kazachstan).
 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Trudy Geologiskhaya Instituta Nauchotekhniskoi Akademia Nauk S.S.S.R. 9,
3e73 (In Russian).
Brongniart, A.T., 1828. Prodrome d’une Histoire des ve
ge
taux fossiles. Dictionnaire
des Sciences Naturelles 57, 1e223.
Bürgl, H., 1960. El Jura
sico en infracreta
ceo del río Bata

, Boyaca
. Boletin Geologico 6,
169e211.
Cantrill, D.J., 1991. Broad leafed coniferous foliage from the Lower Cretaceous Otway
Group, southeastern Australia. Alcheringa: An Australasian Journal of Palae-
ontology 15, 177e190.
Cantrill, D.J., Falcon-lang, H.J., 2001. Cretaceous (late Albian) Coniferales of Alex-
ander Island, Antartica. 2. Leaves, reproductive structures and roots. Review of
Palaeobotany and Palynology 115, 119e145.
Cantrill, D.J., Raine, J.I., 2006. Wairarapaia mildenhallii gen. et sp. nov., A new
araucarian cone related to Wollemia from the Cretaceous (Albian-Cenomanian)
of New Zealand. International Journal of Plant Sciences 167, 1259e1269.
Carlotto, V., Ca
rdenas, J., Cha
vez, R., Gil, W., 1995. Sedimentología de la Formacio
n
Huancane
(neocomiano) de la regio
n de Cusco y su relacio
n con las
variaciones del nivel del mar. Bulletin de l'Institut Français d'Etudes
Andines 24,
1e21.
Carlotto, V., Ca
rdenas, J., Carlier, G., Dia
z-Martínez, E., Cerpa, L., Valderrama, P.,
Robles, T., 2004. Evolucio
n tecto
nica y sedimentaria de la Cuenca Mitú (Permo-
Tri
asico) de la regio
n de Abancay-Cusco-Sicuani (sur del Perú). XII Congreso
Peruano de Geología: Lima, Peru. Sociedad Geolo
gica del Perú 412e415.
Carlotto, V., Carde
nas, J., Carlier, G., 2011. Geología del Cuadr
angulo de Cusco 28-s -
1:50.000. INGEMMET. Boletín, Serie A: Carta Geolo
gica Nacional 138, 255.
Carruthers, W., 1869. I. On some Undescribed Coniferous Fruits from the Secondary
Rocks of Britain 1. Geological Magazine 6, 1e7.
Chambers, T.C., Drinnan, A., McLoughlin, S., 1998. Some Morphological Features of
Wollemi Pine (Wollemia nobilis: Araucariaceae) and their Comparison
to Cretaceous Plant Fossils. International Journal of Plant Sciences 159, 160e171.
Christenhusz, M.J.M., Reveal, J.L., Farjon, A., Gardner, M.F., Mill, R.R., Chase, M.W.,
2011. A new classification and linear sequence of extant gymnosperms. Phyo-
taxa 19, 55e70.
Cleal, C.J., Rees, P.M., 2003. The Middle Jurassic flora from Stonesfield, Oxfordshire,
UK. Palaeontology 46, 739e801.

, L., 1877. Recherches sur la ve
Crie
ge
tation de l'Ouest de la France a  l'e

poque ter-
tiaire. Annales des Sciences Ge
ologiques 9, 1e72.
Cronquist, A., Takhtajan, A., Zimmerman, W., 1966. On the higher taxa of Embry-
obionta. Taxon 15, 129e134.
Daber, R., 1968. A Weichselia-Stiehleria-Matoniaceae community within the Qued-
linburg Estuary of Lower Cretaceous age. Botanical Journal of the Linnean So-
ciety 61, 75e85. https://doi.org/10.1111/j.1095-8339.1968.tb00104.x.
Del Fueyo, G.M., Villar de Seoane, L., Archangelsky, A., Guler, V., Llorens, M.,
Archangelsky, S., Gamerro, J.C., Musacchio, E.A., Passalía, M.G., Barreda, V.D.,
2007. Biodiversidad de las Paleofloras de Patagonia Austral durante el Creta
cico
Inferior. Asociacio
n Paleontolo
gica Argentina. Publicacio
n Especial de la
Asociaciacio
n Paleontolo
gica Argentina 11, 101e122.
Dettmann, M., Clifford, T.H., 2005. Biogeography of Araucariaceae. In: Dargavel, J.
(Ed.), Australia and New Zealand Forest Histories. Araucarian Forests. Australian
Forest History Society Inc, Kingston, pp. 1e10.
Dettmann, M.E., Clifford, H.T., Peters, M., 2012. Emwadea microcarpa gen. et sp. nov.-
anatomically preserved araucarian seed cones from the winton formation (late
albian), Western Queensland, Australia. Alcheringa: An Australasian Journal of
Palaeontology 36, 217e237. https://doi.org/10.1080/03115518.2012.622155.
Diez, J.B., Sender, L.M., Villanueva-Amadoz, U., Ferrer, J., Rubio, C., 2005. New data
regarding Weichselia reticulata: Soral clusters and the spore developmental
process. Review of Palaeobotany and Palynology 135, 99e107.
Doubinger, J., Marocco, R., 1976. De
couverte d’une microflore wealdienne
(Ne
ocomien) dans la re
gion de Cusco (Sud du Pe
rou). Premie re datation des
gres Huancane
Jeanne. Sciences Geologiques 29, 79e89.
Duarte, L., 1989. Remains of the Lower Cretaceous plants from northeast of Brazil.
In: XI Congresso Brasileiro de Paleontologia, Curitiba, Brasil, vol. 1, pp. 219e233.
Duarte, L., 1993. Araucariaceae remains in the Santana Formation, Crato Member
(Aptian); northeastern Brazil. Anais da Academia Brasileira de Ciencias 65,
357e362.
Dutra, T.L., Stranz, A., 2009. Biogeografia, evoluç~ ao e ecologia da familia Araucar-
iaceae: o que mostra a Paleontologia. In: Fonseca, C.R., Souza, A.F., Leal-
Zanchet, A.M., Dutra, T.L., Backes, A., e Ganade, G. (Eds.), Floresta com Arauca
ria:
Ecologia, Conservaça ~o e Desenvolvimento Sustenta
vel. Holos Editora, pp. 15e33.
Edwards, W.N., 1933. On the Cretaceous fern Paradoxopteris and its connexion with
Weichselia. Annals of Botany 47, 317e341.
El-Khayal, A., 1985. Occurrence of a characteristic Wealden fern (Weichselia retic-
ulata) in the Wasia Formation, central Saudi Arabia. Scripta Geologica 79,
75e89.
Escapa, I.H., Catalano, S. a, 2013. Phylogenetic Analysis of Araucariaceae: Integrating
Molecules, Morphology, and Fossils. International Journal of Plant Sciences 174,
1153e1170.
Farjon, A., Filer, D., 2013. An Atlas of the World's Conifers. An Analysis of their
Distribution, Biogeography, Diversity and Conservation Status. Brill, Leiden, The
Netherlands.
Fontaine, W.M., 1889. The Potomac or Younger Mesozoic Flora. Monographs of the
United States Geological Survey 15, 1e377.
Gothan, W., 1910. Weichselia reticulata. In: H. Potonie
: Abbildungen and Bes-
chreibungen fossiler Pflanzenreste. Lief. VII, 126. Κ. preuss. geol. Landesanst.,
Berlin, p. 114.
13
Hallam, A., Crame, J.A., Mancenido, M.O., Francis, J., Parrish, J.T., 1993. Jurassic cli-
mates as inferred from the sedimentary and fossil record [and Discussion].
Philosophical Transactions: Biological Science 341, 287e296.
Harris, T.M., 1935. The Fossil Flora of Scoresby Sound East Greenland. Part 4.
Ginkgolaes, Coniferales, Lycopodiales and Isolated Fructifications. Meddelelser
om Grønland 112, 1e176.
Harris, T.M., 1964. The Yorkshire Jurassic Flora. II e Caytoniales, Cycadales & Pte-
ridosperms. British Museum (Natural History), London, p. 191.
Harris, T.M., 1979. The Yorkshire Jurassic Flora. IV e Coniferales. British Museum
(Natural History), London, p. 166.
Heer, O., 1874. Die Kreide-Flora der artischen Zone, gegründetauf die von den
schwedischen Expeditionen von 1870 und 1872 in Gro €nland und Spitzbergen
gesammelten Pflanzen. Kungliga Svenska Vetenskaps-akademiens Handlingar
12, 1e138.
Henkel, J.B., Hochstetter, W., 1865. Synopsis der Nadelho €lzer, deren charakter-
istischen Merkmale nebst Andeutungen über ihre Cultur und Ausdauer in
Deutschlands Klima. Verlag der J.G. Cottaschen Buchhandlung, Stuttgart.
Huertas, G., 1970. Sertum florulae fossilis Villae de Leiva II. Caldasia 10, 595e602.
Huertas, G., 1976. Sertum florulae fossilis Villae de Leiva. Caldasia 54, 17e23.
Jones, W.G., Hill, K.D., Allen, J.M., 1995. Wollemia nobilis, a new living Australian
genus and species in the Araucariaceae. Telopea 6, 173e176.
Kimura, T., Ghana, T., Mimoto, K., 1988. Discovery of a podocarpaceous plant from
the Lower Cretaceous of Kochi Prefecture, in the Outer Zone of Southwest Japan.
Proceedings of the Japan Academy 64, 213e216. Ser. B.
Kujau, A., Heimhofer, U., Hochuli, P.A., Pauly, S., Morales, C., Adatte, T., Fo € llmi, K.,
Ploch, I., Mutterlose, J., 2013. Reconstructing Valanginian (Early Cretaceous)
mid-latitude vegetation and climate dynamics based on sporeepollen assem-
blages. Review of Palaeobotany and Palynology 197, 50e69.
Kunzmann, L., 2007. Araucariaceae (Pinopsida): Aspects in palaeobiogeography and
palaeobiodiversity in the Mesozoic. Zoologischer Anzeiger 246, 257e277.
Lele, K.M., 1956. Plant fossils from Parsora in the South Rewa Gondwana Basin,
India. Palaeobotanist 4, 23e34.
Lesquereux, L., 1878. On the Cordaites and their related generic divisions, in the
Carboniferous Formation of the United States. Proceedings of the American
Philosophical Society 17, 315e335.


Lima, F.J., Saraiva, A.A.F., Saya ~o, J.M., 2012. Revisa ~o da Paleoflora das Formaço ~es
Missa ~o Velha, Crato e Romualdo, Bacia do Araripe, Nordeste do Brasil. Estudos
Geolo
gicos 22, 99e115.
Lindoso, R.M., Dutra, T.L., Carvalho, I.S., Medeiros, M.A., 2018. New plant fossils from
the Lower Cretaceous of the Parnaíba Basin, Northeastern Brazil: Southern
Laurasia links. Brazilian Journal of Geology 48, 127e145.
Lipps, T., 1938. Acerca de la flora subcreta
cica de Colombia. Estudios geolo
gicos y
paleontolo
gicos de la cordillera oriental de Colombia. Parte3. Departamento de
Minas y Petro
leos, Bogota
137e144.

n, C., 1905. Contribucio
Lisso
n a la Geología de la Isla de San Lorenzo en la Bahía de
Callao. Boletín de Minas. Industrias y Construcciones. Serie 2 1, 2e7.

n, C., 1926. Ape
Lisso
ndice a la obra “Edad de los Fo
siles Peruanos y distribucio
n de
sus depo
sitos en toda la República”. Boletín de la Sociedad Geolo
gica del Perú II
41e46.
Macphail, M.K., 2007. Australian Palaeoclimates: Cretaceous to Tertiary - A review of
palaeobotanical and related evidence to the year 2000. CRC LEME Spec 151, 266.
Open File Rep.
Macphail, M.K., Hill, K., Partridge, A.D., Truswell, E.M., 1995. ‘Wollemi Pine’ - old
pollen records for a newly discovered genus of gymnosperms. Geology Today
11, 48e50.
Macphail, M., Carpenter, R.J., Iglesias, A., Wilf, P., 2013. First Evidence for Wollemi
Pine-type Pollen (Dilwynites: Araucariaceae) in South America. PloS One 8 (7).
https://doi.org/10.1371/journal.pone.0069281 e69281.
Martill, D.M., Bechly, G., Loveridge, R., 2007. The Crato Fossil Beds of Brazil: Window
into an Ancient World. Cambridge University Press, Cambridge. UK, p. 625.
McLoughlin, S., 2001. The breakup history of Gondwana and its impact on pre-
Cenozoic floristic provincialism. Australian Journal of Botany 49, 271e300.
Mendívil, S., D
avila, D., 1994. Geología de los cuadra
ngulos de Cuzco y Livitaca.
INGEMMET, Boletín, Serie A: Carta Geolo
gica Nacional 52, 115.
Mohr, B.A.R., Bernardes-de-Oliveira, M.E.C., Loveridge, R.F., 2007. The macrophyte
flora of the Crato Formation. In: Martill, D.M., Bechly, G., Loveridge, R.F. (Eds.),
The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge
University Press, Cambridge, pp. 537e565.
Mohr, B.A.R., Bernardes-de-Oliveira, M.E.C., Loveridge, R., Pons, D., Sucerquia, P.A.,
Castro-Fernandes, M.C., 2015. Ruffordia goeppertii (Schizaeales, Anemiaceae) e A
common fern from the Lower Cretaceous Crato Formation of northeast Brazil.
Cretaceous Research 54, 17e26.
Monje Dussa
n, C., Martínez, C., Madrin ~a

n, S., Escapa, I., 2016. Nuevos registros de
helechos y coníferas del Cret
acico Inferior en la Cuenca del valle superior del
Magdalena, Colombia. Revista Boletín de Geología 38, 29e42.
Moreno Sa
nchez, M., Go
mez, A., Castillo, H., 2007. Frenelopsis y Pseudofrenelopsis
(Coniferales: Cheirolepidiaceae) en el Creta
cico Temprano de Colombia. Boletin
de Geologia 29, 13e19.
Mussa, D., Carvalho, I.S., Martins, F.J.C., Zuccoloto, M.E., 2000. Paradoxopteris Hirmer
1927, o caule de Weichselia Stiehler 1857, presente no Cret
aceo da Bacía de S~ ao
Luís, Estado do Maranha ~o, Brasil, Revista Universidade de Guarulhos. Geo-
ciencias 5, 60e70.
Nĕmejc, J., 1950. The natural systematic of plants in the light of the present
palaeontological documents. Acta Musei Nationalis Pragae 6B, 1e83.
14 L.C.A. Martínez et al. / Cretaceous Research 110 (2020) 104426
Neumann, R., 1907. Beitr€ age zur Kenntniss der Kreideformation in Mittel-Perú.
Neues Jahrbuch für Geologie und Pal€ aontologie 24, 69e132.
Newell, N.D., 1949. Geology of the Lake Titicaca Region, Peru and Bolivia. Geological
Society of America 36, 1e302.
Ohsawa, T., Nishida, H., Nishida, M., 1995. Yezonia, a new section of Araucaria
(Araucariaceae) based on permineralized vegetative and reproductive organs of
A. vulgaris comb. nov. from the upper cretaceous of Hokkaido, Japan. Journal of
Plant Research 108, 25e39.
Ohsawa, T.A., Yabe, A., Yamada, T., Uemura, K., Terada, K., Leppe, M., Hinojosa, L.F.,
Nishida, H., 2016. Araucarian leaves and cone scales from the Loreto Formation
of Río de Las Minas, Magellan Region, Chile. Botany 94, 805e815.
Pant, D.D., Srivastava, G.K., 1968. On the cuticular structure of Araucaria (Araucar-
ites) cutchensis (Feistmantel) comb. nov. from the Jabalpur series, India. Journal
of the Linnean Society of London, Botany 61, 201e206.
Panti, C., Pujana, R.R., Zamaloa, M.C., Romero, E.J., 2012. Araucariaceae macrofossil
record from South America and Antarctica. Alcheringa: An Australasian Journal
of Palaeontology 36, 1e22.
Passalia, M.G., 2007a. A mid-Cretaceous flora from the Kachaike Formation, Pata-
gonia, Argentina. Cretaceous Research 28, 830e840.
Passalia, M.G., 2007b. Nuevos registros para la flora cret
acica descripta por
Halle (1913) en lago San Martín, Santa Cruz, Argentina. Ameghiniana 44,
565e595.
Pastoriza-Pin ~ ol, J., 2007. Wollemia nobilis: Araucariaceae. Curtis’s Botanical Maga-
zine 24 (7), 155e161.
Pattemore, G.A., Rigby, J.F., Playford, G., 2015. Triassic-Jurassic pteridosperms of
Australasia: speciation, diversity and decline. Boletin Geologico y Minero 126,
689e722.
Philippe, M., Bamford, M., McLoughlin, S., Alves, L.R.S., Falcon-Lang, H.J.,
Gnaedinger, S., Ottone, E.G., Pole, M., Rajanikanth, A., Shoemaker, R.E., Torres, T.,
Zamuner, A.B., 2004. Biogeographic analysis of Jurassic-Early Cretaceous wood
Assemblages from Gondwana. Review of Palaeobotany and Palynology 129,
141e174.
Pole, M., 1995. Late Cretaceous Macrofloras of eastern Otago, New Zealand: Gym-
nosperms. Australian Systematic Botany 8, 1067e1106.
Pons, D., 1983. Contribution a l'e
tude pale
obotanique du Me
sozoïque et du
Ce
nozoïque de Colombie. PhD. thesis. Universite
Pierre et Marie Curie, Paris,
p. 655.
Presl, C., 1847. Die Gef€ aßbündel in Stipes der Frane. Ko €niglich-kaiserliche Hof-
buchdruckerei von Gottlieb Haase, Prague, p. 48.
Rees, P.M., 1993. Caytoniales in Early Jurassic floras from Antarctica. Geobios 26,
33e42.
Rees, P.M., Zeigler, A.M., Valdes, P.J., 2000. Jurassic phytogeography and climates:
new data and model comparisons. In: Huber, B.T., MacLeod, K.G., Wing, S.T.
(Eds.), Warm climates in earth history. Cambridge University Press, Cambridge,
pp. 297e318.
Retallack, G.J., 1997. Earliest triassic origin of isoetes and quillwort evolutionary
radiation. Journal of Paleontology 71, 500e521.
Royo y Gomez, J., 1945. Fo
siles carboníferos e infracreta
cicos del Oriente de Cun-
dinamarca. Compilacio
n de Estudios Geologicos Oficiales de Colombia 6,
193e296.
Salfeld, H., 1911. Fossilien Pflanzen aus der obersten Jura bzw, Unteren Kreide von
Peru e in Hauthal, Reisen in Peru und Bolivien. In: Hautal, R. Zeitsch. Wiss.
Vero€ f. Gesell. Er &. Leipzig, vol. 7, pp. 211e217.
Schuettpelz, E., Schneider, H., Smith, A.R., Hovenkamp, P., Prado, J., Rouhan, G.,
Zhou, X.M., 2016. A community-derived classification for extant lycophytes and
ferns. Journal of Systematics and Evolution 54, 563e603.
Schvedow, N.A., 1958. Nekotorye predstaviteli triasovoj flory Vostochnogo Tajmyra
(Some representatives of the Triassic flora of Eastern Tajmyr). Sborn. Statej
Paleontology Biostratigraphy 7, 56e78 (in Russian).
Sempere, T., Carlier, G., Soler, P., Fornari, M., Carlotto, V., Jacay, J., Arispe, O.,
Ne
raudeau, D., C

nez, N., 2002. Late Permian-Middle
ardenas, J., Rosas, S., Jime
Jurassic lithospheric thinning in Peru and Bolivia, and its bearing on Andean-
age tectonics. Tectonophysics 345, 153e181.


Sender, L.M., Villanueva-Amadoz, U., Pons, D., Diez, J.B., García-Avila, M., Ferrer, J.,
2015. New reconstruction of Weichselia reticulata (Stokes et Webb) Fontaine in
Ward emend. Alvin, 1971 based on fertile remains from the middle Albian of
Spain. Historical Biology 27, 460e468.
Seward, A.C., 1900. La Flore Wealdienne de Bernissart. Me
moires Du Muse
e Royal
d’histoire Naturelle de Belgique 1, 1e37.
Seward, A.C., 1903. Flora of the Uitenhage Series. Fossil floras of the Cape Colony
Annals of the South African Museum 4, 1e22.
Seward, A.C., 1919. Fossil plants, p. 543. Cambridge.
Shi, G., Herrera, F., Herendeen, P.S., Leslie, A.B., Ichinnorov, N., Takahashi, M.,
Crane, P.R., 2018. Leaves of Podozamites and Pseudotorellia from the Early
Cretaceous of Mongolia: stomatal patterns and implications for relationships.
Journal of Systematic Palaeontology 16, 111e137.
Shoemaker, R.E., 1982. Fossil leaves from the Lower Cretaceous Ciano Formation
southwestern Ecuador. Palaeontographica B 180, 120e132.
Silantieva, N., Krassilov, V., 2006. Weichselia Stiehler from Lower Cretaceous of
Maktesh Ramon, Israel: New morphological interpretation and taxonomical
affinities. Acta Palaeobotanica 46, 119e135.
Srinivasan, V., 1995. Conifers from the Puddledock locality (Potomac Group, Early
Cretaceous) in eastern North America. Review of Palaeobotany and Palynology
89, 257e286.
Sternberg, K.G., 1825. Leipzig Versuch einer geognostisch-botanischen Darstellung
der Flora der Vorwelt Teil 4, pp. 1e48.
Sternberg, K.G., 1838. Versuch einer geognostisch-botanischen Darstellung der Flora
der Vorwelt, II (7-8). G.Ha€sse und So €hne, pp. 81e220.
Stiehler, A.W., 1857. Beitr€age zur Kenntniss der vorweltlichen Flora des Kreidege-
birges im Harze. II. Die Flora des Langeberges bei Quedlinburg. Palae-
ontographica 5, 71e80.
Stockey, R.A., 1982. The Araucariaceae: an evolutionary perspective. Review of
Palaeobotany and Palynology 37, 133e154.
Tanai, T., 1979. Late Cretaceous Floras from the Kuji District, Norheastern Honshu,
Japan. Journal of the Faculty of Science, Hokkaido University - Series 4: Geology
and Mineralogy 19, 75e136.
Taylor, E.L., Taylor, T.N., Kerp, H., Hermsen, E.J., 2006. Mesozoic seed ferns: Old
paradigms, new discoveries. Journal of the Torrey Botanical Society 133, 62e82.
Thomas, H.H., 1913. On some new rare Jurassic plants from Yorkshire: Eretmo-
phyllum, a new type of ginkgoalean leaf. Proceedings of the Cambridge Philo-
sophical Society 17, 256e262.
Tidwell, W.D., Ash, S.R., 2006. Preliminary report on the Early Jurassic flora from the
St. George Dinosaur Discovery site, Itah. The Triassic-Jurassic Terrestrial Tran-
sition. U.S. Bureau of Land Management, Bulletin 37, 410e413.
Tomlinson, P.B., Murch, S.J., 2009. Wollemia nobilis (Araucariaceae): Branching,
vasculature and histology in juvenile stages. American Journal of Botany 96
(10), 1787e1797.
Tosolini, A.M.P., McLoughlin, S., Wagstaff, B.E., Cantrill, D.J., Gallagher, S.J., 2013.
Cheirolepidiacean foliage and pollen from Cretaceous high-latitudes of south-
eastern Australia. Gondwana Research 27, 960e977.
van Waveren, I.M., van Konijnenburg-van Cittert, H., Burgh, J. Van Der, Dilcher, D.L.,
2002. Macrofloral remains from the Lower Cretaceous of the Leiva region
(Colombia) Villa de Leiva Bogota
Colombia. Scripta Geologica 123, 1e39.
Walkom, A.B., 1925. Fossil plants from the Narrabeen Stage of the Hawkesbury
Series. Linnaean Society of N.S.W. Proceedings 50, 214e224.
Ward, L.F., 1889. The Cretaceous Formation of the Black Hills as indicated by the
fossil plants. Annual Report of the United States Geological Survey 19, 527e712.
Wilde, M.H., Eames, A.J., 1952. The ovule and “seed” of Araucaria bidwilli with
discussion of the taxonomy of the genus. Annals of Botany 26, 27e47.
Zijlstra, G., van Konijnenburg-van Cittert, H., 2006. Proposal to Conserve the Name
Araucarites C. Presl (Fossil Gymnospermae, Coniferales, Araucariaceae) against
Araucarites Endl. (Fossil Gymnospermae, Coniferales). Taxon 49, 279e280.
Zeiller, R., 1910. Sur quelques plantes wealdiannes du Pe
rou. Comptes Rendus
Academie Science CL 1488e1490.
Zeiller, R., 1914. Sur quelques plantes wealdiennes recueillies au Pe
rou par M. le
Capitaine Berthon. Revue Ge
ne

rale de Botanique 25, 647e674.