Cretaceous Research 54 (2015) 34e49

Contents lists available at ScienceDirect

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

Plant taphonomy and palaeoenvironment from the Upper Cretaceous

of Isona, Tremp Basin, southern Pyrenees, Catalonia, Spain

ronique Daviero-Gomez a,
Sheila Villalba-Breva a, c, Josep Marmi b, *, Bernard Gomez a, Ve

ndez-Marro

n d
Carles Martn-Closas c, M Teresa Ferna
Laboratoire de G

eologie de Lyon: Terre, Plan
etes, Environnement, Universit

e Lyon 1, 69622, Villeurbanne, France
 de Paleontologia Miquel Crusafont, C/Escola Industrial 23, 08201, Sabadell, Catalonia, Spain
Institut Catala
c
Departament d'Estratigraa, Paleontologia i Geoci
encies Marines, Universitat de Barcelona, c/Mart i Franqu
es s/n, 08028, Barcelona, Catalonia, Spain
d
Instituto de Geociencias (CSIC, UCM), Facultad de Geologa, C/Jose Antonio Novais 12, 28040, Madrid, Spain
a

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 14 March 2013
Accepted in revised form 22 October 2014
Available online 30 December 2014

Lower Maastrichtian plant fossils were newly collected from the base of the Xull unit, Tremp Group, in
Isona, Tremp Basin, southern Pyrenees. They consist of conifers, monocot and eudicot angiosperms.
Palynology shows algal oospores, fern spores, gymnosperm and angiosperm pollen grains.
Sedimentological and taphonomic analyses indicate that this assemblage is parautochthonous and
allochthonous. The plant fragments probably came from a nearby exposed area and were deposited in
the distal oodplain. These taphonomic data shed light on the exact location and sampling biases of the
palaeobotanical collection from Isona gathered by Vicente i Castells in the 1980s.
Angiosperms and ferns dominated the plant community, which grew inland in freshwater environments. Cheirolepids inhabited coastal freshwater lakes in the same basin (the Posa unit) and in the
geographically close basin of Vallcebre.
2014 Elsevier Ltd. All rights reserved.

Keywords:
Palaeobotany
Taphonomy
Latest Cretaceous
Pyrenees
Spain

1. Introduction
The southern Pyrenees are well known for their abundance of
vertebrate remains, especially of dinosaurs, collected from the
Upper Cretaceous continental facies (Riera et al., 2009, and references therein). Plant fossil assemblages from the Pyrenean Maastrichtian have been mainly studied from a palynological point of

dus (1972), Porta et al. (1985), Ashraf and Erben (1986),
view. Me

dus et al. (1988), Lo

pez-Martnez et al. (1999), Mayr et al.
Me

ndez-Marro

n et al. (2004), Torices et al. (2012) and
(1999), Ferna

Villalba-Breva et al. (2012) focused their research in the Ager,

and Tremp basins. Their palynological
Vallcebre, Coll de Nargo
analyses showed a high diversity of plants, including freshwater
green algae, hornworts, spike mosses, ferns, cycads, ginkgoes, and
monocot and eudicot angiosperms. In most localities spores are far
more abundant than gymnosperm and angiosperm pollen grains.
Nichols and Johnson (2008) gave a state of the knowledge of the
palaeobotany and palynology of the Pyrenean Maastrichtian,
highlighting that this sporomorph assemblage differed from assemblages from other localities near the K-Pg boundary. Plant
* Corresponding author. Tel.: 34 93 7261769.
E-mail address: josep.marmi@icp.cat (J. Marmi).
http://dx.doi.org/10.1016/j.cretres.2014.10.007
0195-6671/ 2014 Elsevier Ltd. All rights reserved.

meso- and megafossil assemblages from the Maastrichtian of the

Pyrenees have been studied in the Vallcebre (Marmi et al., 2008,
2010, 2012; Riera et al., 2010; Villalba-Breva et al., 2012) and
Tremp (Vicente i Castells, 2002; Marmi et al., 2014) Basins.
In 1979, the geologist Enric Sunyer discovered a locality with
abundant plant fossils near the village of Isona (Vicente i Castells,
2002). In the early 1980s, the naturalist Joan Vicente i Castells
and collaborators from the Institut d'Estudis de la Natura del
s Nord (Santa Coloma de Gramenet) collected around 400
Barcelone
plant fossils in the village surroundings, mostly from the so-called
Isona Sud locality. Although it constitutes one of the few known
examples of uppermost Cretaceous plant assemblages in Europe
(Herman and Kvacek, 2007, 2010), these plant fossils were poorly

n and Die

guez, 1992; Vicente i
described until recently (Barro
Castells, 2002). Although Marmi et al. (2014) partially revised
these collections, still little is known about the sedimentology,
taphonomy and collecting biases of the Isona localities. Thus,
Vicente i Castells (2002) did not provide accurate information for
the locations and geology of the plant beds.
The present study reports the results of a palaeontological
excavation carried out in a plant fossil locality near Isona in 2010.
Our aim is: (1) to determine the geographical and stratigraphic
settings of the beds corresponding to the Isona Sud locality

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

sampled by Vicente i Castells and collaborators; (2) to characterize

the depositional origin of the plant beds; and (3) to suggest the
fossil plant ecology and environment based on comparative sedimentological and taphonomic analyses.
2. Material and methods
Based on the lithological features of the specimens of the
ncies
Vicente i Castells collections housed in the Museu de Cie
Naturals de Barcelona, we located their locality Isona Sud along a
trail to the south of the village of Isona. In 2010, the excavation
undertaken yielded 64 slabs with 116 plant fossils and 4 samples
were collected for palynology in dark grey lutites. The studied
section had the following base coordinates: N42 060 43.800
E01020 37.6200 . The lithostratigraphy and sedimentology of this
section were characterized from facies analysis. Bed-by-bed sampling was carried out in parallel with taphonomic analysis to
remove specimens from different points along the same bed. All
plant fossils were collected regardless of the size and preservation
state to avoid any collection biases. The whole set of specimens has
been taken into account to determine the plant fossil composition
and to quantify the number of leaf fragments by size, morphology
and taxa. Most (92 of 116) were fragments of unidentied plant
fossils. Only 24 plant fossils could be described in detail, measured
and identied. Specimens were prepared under an Olympus SZX10
stereomicroscope and photographed with a Canon EOS 60D integrated digital camera tted with a Canon 100 mm macro lens, at the

Lyon 1. Measurements were taken using digital pictures
Universite
and the freeware Image J v. 1.40g (Rasband, 2008). Furthermore, a
camera lucida was used to create line drawings of some of the
leaves. Angiosperm leaves were described in accordance with Ellis
et al. (2009). The newly collected plant fossils were compared with
specimens of the Vicente i Castells collections, housed in the Museu
ncies Naturals de Barcelona (MGB). The samples for palynode Cie
logical analysis were prepared following established procedures of
crushing, treatment by hydrochloric (HCl) and hydrouoric (HF)

35

acids at the Universitat de Barcelona. The slides were examined

under a Leitz Laborlux D light microscope and the photomicrographs were taken with a Wild MPS 41/51 S at the Instituto de
Geociencias (CSIC-UCM) of Madrid.
Newly collected plant fossils are housed in collections at the
 (MCD), Isona, Lleida province, CataMuseu d'Isona i Conca Della
lonia, Spain. Palynological slides are housed in the collections of the
ncies Marines,
Departament d'Estratigraa, Paleontologia i Geocie
Universitat de Barcelona.
3. Geological setting
The Pyrenees are an Alpine fold-thrust belt that was formed as a
result of a collision between the Iberian and European continental
plates from the Santonian or Campanian to the Miocene (Teixell,
2004). The mountain belt consists of a Hercynian basement and a
sedimentary cover that developed from the Late Permian to the
Early Cretaceous in rift basins and later, from the late SantonianeCampanian to the Oligocene, in foreland and piggy-back babregas et al., 1986; Mun
~ oz, 1989; Teixell, 2004). As a
sins (Puigdefa
consequence, the Upper Cretaceous Southern Pyrenean Basin was
segmented into several sub-basins or depositional centres. These


, Tremp and Ager
are, from east to west: Vallcebre, Coll de Nargo
vol et al., 2000). The studied plant fossils were collected from
(Arde
the Tremp Basin.
The Tremp Basin forms a broad syncline located in the Montsec
thrust sheet (Fig. 1). The Upper Cretaceous basin inlling is subdivided into seven lithostratigraphic units that are mostly dominated by nearshore marine limestones to basinal marls (Caus and

mez-Garrido, 1989). By the Campanian, the sequence became
Go
regressive and followed an east-to-west direction trend. Its base is
dominated by the shallow-neritic to coastal cross-bedded calcarenites of the Areny Sandstone Formation (Mey et al., 1968). These
shallow marine rocks grade laterally to the east and upward to the
uppermost unit, the Tremp Group (Cuevas, 1992), known regionally
as Garumnian (Leymerie, 1862). It ranges from the late Campanian

Fig. 1. Geological map of the Tremp Basin with the location of the studied area.

36

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

to the Thanetian. It is mainly formed by lagoonal and coastalswamp lacustrine marls and limestones, intercalated in some
areas with lignites and covered by uvio-lacustrine red beds. The
Tremp Group is around 700 m in thickness (Cuevas, 1992), and the
plant fossils and palynological samples were collected from its
basal part, which corresponds to the Xull unit (Liebau, 1973). This
unit was later unied by Cuevas (1992) with the underlying Posa
unit (Liebau, 1973) in the La Posa Formation.
4. Stratigraphy and sedimentology
The type section of the basal Tremp Group was studied along a

u de la Posa church, which is
ravine to the north of the Mare de De
located to the north-east of the village of Isona (Fig. 2A). The base of
the section shows two intercalations of the nearshore marine
sandstones of the Areny Formation that contain abundant Radiolitella pulchellus Vidal rudists, other bivalves, and corals. This formation is related to deposition in coastal barriers (Mey et al., 1968;
Nagtegaal, 1972). The overlying succession is up to 130 m in
thickness, and shows alternating organic-rich marls, lignites with
cheirolepidiacean leafy twigs, and charophyte limestones with
some intercalations of marlstones, siltstones and sandstones. A
rudist horizon formed by Hippuritella castroi Vidal marks the top of
the unit. This succession corresponds to the Posa unit (Liebau, 1973)
and was attributed to deposition in a coastal-swamp environment
connected to shallow freshwater lakes (Nagtegaal, 1972), rich in
lower Maastrichtian charophytes (Villalba-Breva and MartnClosas, 2013). Finally, this unit is covered by a succession of grey
lutites with some intercalations of sandstones, variegated lutites
and limestones, corresponding to the Xull unit. These strata were
attributed to deposition in a lagoon laterally related to oodplains
with marshes and lakes (Daz Molina, 1987).
The plant fossils and palynological samples were collected from
a section representing the base of the Xull unit exposed at the
Isona Sud locality (Fig. 2B). Two lithofacies were dened. Facies 1
forms the thickest deposits in the section and comprises dark grey
lutites with abundant plant fossils and ferruginous nodules. This
monotonous succession is interrupted by facies 2, comprising up to
7-cm-thick laminated siltstones, which contain an abundant
accumulation of fossil shells and a few plant fossils (Fig. 2B). The
most abundant shells belong to the gastropod genus Melanopsis

russac. Less frequently, facies 2 includes bivalve shells of the
Fe
genus Corbicula Megerle von Mhlfeld. Both taxa are associated
with shallow, brackish to freshwater environments (Liebau, 1973).
Vertebrate remains are scarce, and consist only of isolated teeth
(probably of crocodiles). These materials (facies 1 and 2) are
attributed to the deposition of overbank, suspension-load sediment
in uvio-deltaic oodplains, according to criteria such as lithology,
colour, sedimentary structures and fossil content. The dark colour
and preservation of lamination indicate high organic matter content and show that they were deposited under anoxic standing
water, generally without bioturbation. The abundance of ferruginous nodules can be attributed to the high Eh and low pH conditions during early burial. The lithologies observed in the oodplain
deposits of Isona Sud are interpreted as reecting the distance to
the source area of sediment. Sediments close to the uvial clastic
supply were coarser than sediments deposited further away from
this point. As a consequence, alternating silts and muds characterized the distal oodplain. The presence in laterally equivalent
localities of thin sandstone beds corresponding to small channels,
which would provide some drainage to the oodplain, suggests
that the plant fossils were transported by a conned stream in the
oodplain. These deposits are early Maastrichtian in age, based on
the age of the underlying Posa unit, which was established from
two rudist horizons (Liebau, 1973; Riera et al., 2009) and

charophyte assemblages (Feist and Colombo, 1983; Villalba-Breva

and Martn-Closas, 2013).
5. Systematic palaeontology
5.1. Plant megafossils
5.1.1. Conifers
Order Pinales
Family Cupressaceae
cf. Cunninghamites sp.
Fig. 3A
Material. MCD 5299-5304
Description. These specimens consist of six unbranched twigs
with spirally arranged leaves (Fig. 3A). They are 2.4e6.9 cm long
and 0.4e0.6 cm wide. The leaves are straight or curved distally
forming acute angles of 45.5 e57.8 with the stems. The free leaf
parts are narrow and needle-shaped. They are 9.3e10.8 mm
long and 0.4e0.6 mm wide. Macroscopically, the leaf margins
seem entire, but no cuticle could be removed to determine
whether or not microscopic teeth occur on the margin. The leaf
cushions are rhomboidal and are wider than the leaf lamina
(1.8e2.2 mm wide).
Discussion. Cretaceous conifers with spirally arranged, needlelike leaves are included within numerous form-genera based
on their gross morphology (e.g. needle curvature, presence of
keel(s), shape of transverse section). However, no cuticle could
be removed to ascertain the taxonomy of specimens in Isona. In
gross morphology, these specimens resemble Cunninghamites
Presl in Sternberg based on the following features: spirally arranged, linear-lanceolate, dorsi-ventrally attened leaves
diverging from the stem at acute angles, and branches showing
conspicuous rhomboidal leaf cushions (Bosma et al., 2012).
Several of these features are shared with other form-genera,
such as Elatocladus Halle and Geinitzia Endlicher. However,
Elatocladus differs from Cunninghamites in having the leaf base
contracted and lacking conspicuous leaf base cushions (Kva
cek,
1999). Geinitzia has smaller and usually falcate leaves with
clearly triangular transverse sections (Herman and Kvacek,
2010). Although Kunzmann (1999, 2010) suggested placing
Geinitzia in its own family (Geinitziaceae), Herman and Kvacek
(2010) and Halamski (2013) argued that Cunninghamites and
Geinitzia would be better regarded as members of the Cupressaceae. Specimens from the same locality assigned by Vicente i
Castells, (2002) to Elatocladus albertaensis Bell (e.g. MGB 38204,
38211) and Prepinus solmsii (Seward) Vicente i Castells comb.
nov. (e.g. MGB 38180) are similar to the specimens studied
herein. Marmi et al. (2014) identied these specimens from the
Vicente i Castells collections as cf. Cunninghamites sp.
Order Incertae sedis
Family Incertae sedis
cf. Podozamites
Fig. 3B
Material. MCD 5305
Description. This consists of a single specimen (Fig. 3B). The
medial part of the lamina and apex have an overall lanceolate
shape. It is 2.8 cm long and 0.7 cm wide. The leaf margin is
entire. The apex is acute. There are 19 equal veins running
parallel to the leaf margin and slightly converging to the apex.
Veins end just before reaching the apex margin.
Discussion. The genus Podozamites (Brongniart) Braun in
Mnster is known from the Upper Triassic to the Upper Cretaceous and is widely distributed geographically (Doludenko,

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

37

Fig. 2. Geological and stratigraphic setting of the studied area. A, Geological sketch of the area around Isona with the location of the studied section, Isona Sud. B, Stratigraphic type
section of the basal Garumnian in the Tremp Basin and detailed section of Isona Sud with locations of the slabs and pollen samples.

38

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

Fig. 3. Plant megafossils collected from the Isona Sud locality. A, Fragment of a leafy twig of cf. Cunninghamites sp.; MCD 5304. B, Middle and apical parts of a leaf of cf. Podozamites
sp.; MCD 5305. C, Fragment of a ribbon-shaped monocot leaf; MCD 5306. D-J, Eudicot leaves. D, Eudicot form 1; MCD 5310. E, H, Eudicot form 2; MCD 5317, MCD 5320. F, Eudicot
form 3; MCD 5318. G, Eudicot form 4; MCD 5319. I, Eudicot form 5; MCD 5321. J, Eudicot form 6; MCD 5322.

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

1967). However, its taxonomic afnity with the Pinopsida is

uncertain (Herman and Kvacek, 2010). The specimen collected
from Isona is fragmentary, but it shows similarities in terms of
gross morphology and venation pattern with P. cf. lanceolatus
(Lindley and Hutton) Harris reported from the lower Campanian
of Grnbach, Austria (Herman and Kva
cek, 2010). Vicente i
Castells, (2002) assigned some specimens from the Isona Sud
locality to P. lanceolatus (MGB 38190-91) and P. marginatus Heer
(MGB 38192). However, these specimens were not included in
the revision by Marmi et al. (2014) because of the fragmentary
nature and bad preservation. Parallelodromous venation was
observed in all three specimens of the Vicente i Castells collections, but it was difcult to establish whether they belong to
Podozamites or represent some sort of monocot foliage because
the gross morphology was badly preserved.
5.1.2. Angiosperms
Clade Monocots
Monocot gen. et sp. indet.
Fig. 3C
Material. MCD 5306-5308
Description. They are represented by three isolated fragments of
ribbon-shaped leaves with parallelodromous venation (Fig. 3C).
They measure 1.6e4.6 cm long and 0.7e1.6 cm wide. The bestpreserved specimen shows 10 equally-spaced (0.6e0.7 mm),
parallel, single-order veins. The leaf margin is entire and lacks
additional structures such as spines.
Discussion. Monocot leaves with single order parallel, longitudinal veins were identied by Vicente i Castells, (2002) as Rhyzocaulon macrophyllum Saporta (MGB 38424, 38433e37,
38440e42) and Cyperites chavanesii Heer (MGB 38448, 38450).
These specimens were revised by Marmi et al. (2014) and
assigned to cf. Pandanites. Some leaves in the Isona collections
(e.g. MGB 38434, 38436, 38440) show narrower, transverse or
slightly oblique to longitudinal veins like in the specimen MCD
5308. This venation pattern is observed in other monocot fossil
leaves from the Upper Cretaceous of Europe such as in Typhacites rugosus Saporta from the upper Santonian of Fuveau,
France (Saporta, 1890). Kva
cek and Herman (2004; pl. IX, gs. 1,
2) and Herman and Kvacek (2010; pl. 21, gs. 7, 8) described a
specimen of monocot with parallel venation from the lower
Campanian of Grnbach (Austria). However, it differs from the
Isona specimens in having four vein orders and thick transverse
veins.
Clade Eudicots
Order Incertae sedis
Family Incertae sedis
Eudicot form 1
Figs. 3D, 4A-B
Material. MCD 5309-5311
Description. Three specimens consist of simple, linear leaves
(Figs. 3D, 4A-B). The leaf laminas are symmetrical and
untoothed. They measure 3.6e3.7 cm long and 0.5e0.6 cm wide.
The lamina base is symmetrical, acute and decurrent. The primary vein is wide, monopodial and pinnate. The secondary
veins are very thin and, judging from the small portion preserved, appear to be eucamptodromous to brochidodromous. A
single specimen has the petiole intact. The petiole is marginally
attached and measures 0.9 cm long and 0.08 cm wide.
Discussion. Specimens similar to Eudicot form 1 were assigned
to Eucalyptus angusta Velenovsk (e.g. MGB 38387), E. geinitzii
Heer (e.g. MGB 38388), Proteophyllum acus Velenovsk and

r (e.g. MGB 38383) and Nectandra prolica Berry (e.g.
Vinikla

39

MGB 38243) by Vicente i Castells, (2002). Marmi et al. (2014)

consider that specimens of E. geinitzii and N. prolica in the
Vicente i Castells collections represent a single species (Dicot
type 3). The shape and venation of Eudicot form 1 (Fig. 4B) is
very close to Salix assimilis Saporta from the upper Albian of
Nazareth, Portugal (pl. XXXVI, g. 8 and pl. XXXVII, gs. 2,3,6, 13
in Saporta, 1894). Eudicot form 1 leaves also resemble Dicotylophyllum proteoides (Unger) Herman and Kva
cek comb. nov.
and Dicotylophyllum sp. 7 from the lower Campanian of Grnbach (Herman and Kvacek, 2010) in being linear and entiremargined leaves.
Order Incertae sedis
Family Incertae sedis
Eudicot form 2
Figs. 3E, H, 4C, H
Material. MCD 5312-5317, 5320
Description. Seven specimens consist of simple, unlobed leaves
with symmetrical laminas (Figs. 3E, H, 4C, H). The lamina seems
elliptical to oblong and is 1.7e8.3 cm long and 1.6e5.4 cm wide.
In some specimens, the portion of the margin preserved is
toothed, with one or two orders of teeth. The sinus shape is
angular, and the tooth shape is proximally convex and distally
convex or straight. The primary vein is monopodial and pinnate.
The secondary vein framework is craspedodromous. The
attachment of the major secondary veins to the midvein is
excurrent.
Discussion. Eudicot form 2 is similar to specimens that Vicente i
Castells, (2002) assigned to Alnus corylifolia Lesquereux (e.g.
MGB 38280, 38314, 38315) and Betula stevensoni Lesquereux
(e.g. MGB 38332). According to Marmi et al. (2014), specimens of
A. corylifolia and B. stevensoni in the Vicente i Castells collections
might represent the same species (Dicot type 7) and belong to
the family Betulaceae. Grebenkia europea Herman and Kva
cek,
Viburniphyllum austriacum Herman and Kva
cek and Dicotylophyllum sp. 2 from the Grnbach Formation of Austria
(Herman and Kvacek, 2010) are characterised by elliptic to
oblong, symmetric leaves with a toothed margin and pinnate
craspedodromous venation. However, these species differ from
Eudicot form 2 based on the shape of marginal teeth and sinuses.
Order Incertae sedis
Family Incertae sedis
Eudicot form 3
Figs. 3F, 4D
Material. MCD 5318
Description. The single specimen consists of a simple, unlobed
leaf (Figs. 3F, 4D). The lamina shape is elliptical and symmetrical.
It measures 1.7 cm long and 1.0 cm wide. The apex angle is
obtuse and the shape convex. The primary vein is monopodial
and pinnate. The secondary vein spacing increases proximally.
The secondary veins are attached excurrently to the midvein at
uniform angles. There are simple agrophic veins.
Discussion. Leaves with monopodial venation are common in the
Vicente i Castells collections of Isona and from other Upper
Cretaceous localities in Europe (e.g. Grnbach; Herman and
Kva
cek, 2010). For instance, this feature was observed in Dicot
type 7 by Marmi et al. (2014), which was assigned to cf. Betulaceae. However, specimens belonging to Dicot type 7 have no
agrophic veins. Some important features, like leaf margin and
secondary vein framework, will be necessary for a precise
comparison.
Order Incertae sedis
Family Incertae sedis
Eudicot form 4

40

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

Fig. 4. Interpretative drawings of angiosperm leaves from the Isona Sud locality. A-B, Eudicot form 1; MCD 5310, MCD 5311. C, H, Eudicot form 2; MCD 5317, MCD 5320. D, Eudicot
form 3; MCD 5318. E, Eudicot form 4; MCD 5319. F, Eudicot form 5; MCD 5321. G, Eudicot form 6; MCD 5322.

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

Table 1
Palynological composition of samples from the Isona Sud locality.
ISP-1
Spores
Acanthotriletes sp.
Acylomurus sejunctus Phillips
Anapiculatisporites dawsonensis Reiser
and Willians
Anapiculatisporites sp.
Biretisporites potoniaei Delcourt and
Sprumont
Calamospora mesozoica Couper
Chomotriletes fragilis Pocock
Cicatricosisporites hallei Delcourt and
Sprumont
Cingulatisporites brevispeciosus Pf.
Concavisporites sinuatus (Couper) Krutzsch
Concavissimisporites subgranulosus (Couper)
Pocock
Cyathidites australis Couper
Cyathidites punctatus (Delcourt and Sprumont)
Delcourt et al.
Echinatisporis longechinus Kr.
Echinatisporis sp.
Foveotriletes sp.
Klukisporites scaberis (Cookson and Dettmann)
Dettmann
Laevigatosporites discordatus P.
Laevigatosporites haardti (R. Pot. and Ven) Th.
and P.
Laevigatosporites ovatus Wilson and Webster
Laevigatosporites sp.
Leiotriletes adriennis (R. Pot. and Gell) Kr.
Leiotriletes dorogensis Kedves
Leiotriletes sp.
Leptolepidites verrucatus Couper
ring
Matthesisporites plurituberosus Do
Patellasporites sp.
Polypodiaceoisporites sp.
Polypodiisporites sp.
Retitriletes austroclavatidites (Cookson)
ring et al.
Do
Retitriletes circolumenus (Cookson and
Dettmann)
Backhouse
Trilobosporites sp.
Trilites tuberculiformis (Cookson) Dettmann
Triplanosporites sp.
Undulatisporites sp.
Gymnosperms
Araucariacites australis Cookson
Cycadopites carpentieri (Delcourt and
Sprumont)
Singh
Cycadopites sp.
Pinuspollenites ruginosa (Stanley) Oltz
Pinuspollenites sp.
Angiosperms
Arecipites sp.
Echinotriporites sp.
cf. Interporopollenites sp.
Monocolpopollenites lutescens Kedves
Monocolpopollenites sp.
Monosulcites cf. borassioides Dutta and Sah
Monosulcites brevispinosus Sah and Dutta
Monosulcites sp.
Nudopollis sp.
Periporopollenites sp.
Platycaryapollenites ferrerri Porta et al.
Polyporopollenites sp.
Pseudoromeinipollenites sp.
Rugulitriporites sp.
Semioculopollis sp.
Subtriporopollenites constans P.
Subtriporopollenites sp.
Triatriopollenites sp.

ISP-2

X
X
X

X
X

41

Table 1 (continued )
ISP-1
ISP-3

ISP-4

X
X

Triporopollenites sp.
Trudopollis sp.
Algae
Lecaniella sp.
Oedogonium cretaceum Zippi
Ovoidites spriggi (Cookson & Dettmann) Zippi
Tasmanites sp.
Zygnemataceae aff. gelasinicysta

ISP-2

ISP-3

ISP-4

X
X
X
X
X
X
X

X
X
X

X
X

X
X
X
X
X
X
X
X

X
X

X
X
X
X
X
X
X
X

X
X
X
X

X
X
X

X
X
X

X
X
X
X

X
X
X
X

X
X

X
X

X
X

X
X
X

X
X
X

X
X

X
X
X

X
X
X
X
X

X
X
X
X

X
X
X
X

X
X
X
X
X
X

X
X
X
X
X

Figs. 3G, 4E
Material. MCD 5319
Description. A single specimen shows the lamina base (Figs. 3G,
4E). It measures 2.2 cm long and 2.4 cm wide. The lamina is
unlobed and the margin untoothed. The primary vein is
monopodial and pinnate. Two long secondary veins are excurrently attached near the lamina base. Secondary veins are
alternate.
Discussion. Some specimens in the Vicente i Castells collections
show entire margins, long basal secondary veins widely spaced
and intersecondary veins parallel to major secondary veins. For
instance, these features can be observed on the specimens
assigned to Ficus pealei Knowlton (e.g. MGB 38266 and 38267)
or Magnolia amplifolia Heer (e.g. MGB 38213) by Vicente i
Castells, (2002). These specimens were assigned to Dicot type
9 with uncertain taxonomic afnity by Marmi et al. (2014).
Order Incertae sedis
Family Incertae sedis
Eudicot form 5
Figs. 3I, 4F
Material. MCD 5321
Description. The single specimen is a simple, unlobed leaf with
symmetrical lamina (Figs. 3I, 4F). It measures 2.7 cm long and
1.6 cm wide. The margin is untoothed. It appears to have three
basal veins of which the midvein is wider and straight. Tertiary
veins are poorly preserved. Exterior tertiary veins arise from
basal secondary veins but it is difcult to ascertain if they are
looped or reticulate. Epimedial tertiary veins appear to be
reticulate.
Discussion. The Vicente i Castells collections include a number of
specimens with three basal veins very similar to Eudicot form 5.
For instance, MGB 38221 was assigned to Cinnamomum afne
Lesquereux, MGB 38291 and 38300 were assigned to Carya
heerii Ettingshausen and MGB 38390 to Eucaliptus geinitzii Heer
by Vicente i Castells, (2002). The later specimens were considered a single taxon (Dicot type 6) by Marmi et al. (2014).
Order Incertae sedis
Family Incertae sedis
Eudicot form 6
Figs. 3J, 4G
Material. MCD 5322
Description. The single specimen corresponds to the leaf apex
(Figs. 3J, 4G). The margin is untoothed. It measures 2.8 cm long
and 3.6 cm wide. The primary vein is monopodial and pinnate.
The secondary veins are excurrently attached to the midvein.
The secondary vein framework is difcult to ascertain because
the secondary vein ends are not preserved.
Discussion. Leaves with monopodial venation and entire margins are common in the Vicente i Castells collections of Isona.
However, additional features, such as secondary vein framework, are necessary to compare with given species or leaf
morphotypes.

42

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

5.2. Plant microfossils

All four palynological samples yielded diverse associations,
albeit not very abundant in number of grains (between 58 and 79
grains per sample). The sporomorph assemblage comprises 65
identied taxa (Table 1, Figs. 5, 6). In detail, sample ISP-1 consists of
31 taxa, including 21 of moss- (Acanthotriletes sp.) and fern spores,
as well as 4 gymnosperm and 6 angiosperm pollen taxa. Spores are
the most abundant (70%), trilete forms being dominant, including
Leptolepidites verrucatus Couper, Retitriletes circolumenus (Cookson
and Dettmann) Backhouse, Acylomurus sejunctus Phillips, Trilites
tuberculiformis (Cookson) Dettman, Matthesisporites plurituberosus
ring and Klukisporites scaberis (Cookson and Dettmann) DettDo
mann (Figs. 5B-C, N, P, 7). Angiosperms constitute 18% of the
assemblage (Figs. 6G, N, R, 7); pollen grains of conifers and Cycadales (Figs. 6J, 7) only 12%.
Sample ISP-2 consists of 28 taxa. Fern spores are abundant (55%)
(Figs. 5F-L, Q, 6A, F, 7). Gymnosperms constitute 28%, Pinaceae
being the most abundant (Figs. 6K, 7), while angiosperms are 17%
(Figs. 6D, T, 7).
Sample ISP-3 consists of 9% of phycomes of prasinophyte algae
(Tasmanites sp., Figs. 6U, 7). Fern spores are 41% (Figs. 5R, 6B, H, 7)
while gymnosperms are 16% (Fig. 7). Angiosperms are 34%, and
especially contain Arecaceae (Monocolpopollenites sp. Fig. 6I),
Juglandaceae (Playcaryapollenites ferreri Porta et al., Subtriporopollenites sp., Fig. 6O), Myricaceae (Triatriopollenites sp.) and
Betulaceae (Triporopollenites sp.).
Sample ISP-4 consists of 14% of freshwater green algae (Fig. 7). Fern
spores are 51%, trilete forms being dominant (Figs. 5D, 6C, 7).
Gymnosperms are 20% (Figs. 6L, 7), with Araucariacites australis
Cookson being the most abundant species. Angiosperms are 15%
(Figs. 6M, P-Q, S, 7).

which are lines of weakness (Fig. 8B). Experiments using leaves

placed in a revolving drum showed that the breakdown stages always followed the same pattern even in very tough leaves, and
secondary veins in particular are lines of weakness along which
early tears occur (Ferguson, 1985). Thus, it is likely that the plant
megafossils of Isona Sud were transported in suspension in the
water column of a conned stream (channel) before reaching the
oodplain. During this transport, a mechanical fragmentation and
size reduction of plant debris occurred as a result of collision and
joining with sharp objects. Furthermore, there are several leaves
with holes caused by arthropod feeding or fungal attacks (Fig. 8C),
and eif unhealed by the living plante these leaves usually sink
more rapidly than leaves without such damage. Up to 10-cm-long
stems were also collected, but no preferential alignment could be
determined (Fig. 8D). The variability in shape and size suggests that
there was no selection either in the supply or in the transport of
plant remains to the depositional setting. These data, along with
the lack of rootlet marks, suggest that the leaves and stems are
mostly allochthonous in the depositional setting. Although it is
impossible to precisely establish how far they were transported,
relative parautochthony could be estimated by taphonomic analysis. Thus, Eudicot form 2 represents the most abundant and wellpreserved remains, suggesting that the parent plants may have
grown along the river banks or close to the overow point and may
correspond to the riparian vegetation. Other angiosperm and
gymnosperm leaves are scarcer and less well-preserved, and were
possibly transported from further away.
6.1.3. Fossil diagenesis
Plant megafossils are preserved as compressions, which would
suggest that they underwent a relatively rapid burial. This resulted
in limited humication (Martn-Closas and Gomez, 2004, and references therein). In addition, they were collected in ne lutites and
siltstones that strongly limit percolation by oxidizing meteoric
water. However, a high degree of diagenetic carbonization occurred
and this hinders a detailed taxonomic identication.

6. Taphonomy and palaeoecology

6.2. Plant microfossils
6.1. Plant megafossils
6.1.1. Necrobiosis
In the plant fossil assemblage of Isona Sud, angiosperms are
more abundant and diverse than conifers. This is to be expected for
this period of plant history and also for litter deposited in oodplain
environments that display an over-representation of deciduous
plants and an under-representation of evergreen plants (Ferguson
et al., 1999). Also, abscission modes are different in angiosperms
and conifers. Many angiosperms shed their leaves by abscission at
the base of the petiole, whereas in many conifers, leafy shoots or
twigs fall when the stems break after drying for a few months or
years (Gomez et al., 2001, 2002). This would explain the differences
in anatomical connection between the articulated twigs of conifers
and the isolated angiosperm leaves.
6.1.2. Biostratinomy
All fossil leaves in the plant assemblage of Isona Sud are fragmented and the fragments are small and medium in size. Most of
the plant megafossils are unidentied and smaller than 2 cm2 (with
an average size of 0.69 cm2), while most of identied plant remains
are larger than 5 cm2. Most of the eudicot leaves show fragmented
laminas without anatomically connected petioles (Fig. 8A). The
point of attachment between the petiole and the lamina represents
a weak link, and biodegradation combined with tearing during
transport by water may cause their separation (Ferguson et al.,
1996). In addition, lamina tears are along the secondary veins,

The four samples are poor considering the abundance of sporomorphs, but they represent relatively high diversity. There is little
evidence of alteration during diagenesis, allowing for a taphonomic
(biostratinomic) analysis. They show contrasting taphonomic features. The assemblage of sample ISP-1 is dominated by very wellpreserved miospores and large and heavily-ornamented fern
spores. Fern spore-dominated assemblages are usually not transported far from their source (Traverse, 2008) and the assemblage is
probably parautochthonous.
In sample ISP-2, the percentage of pinaceous pollen grains is
higher than in any other assemblages (Fig. 7). Bisaccate pollen are
produced in large numbers and are easily transported for long
distances by water and wind due to the buoyancy of their structure.
Thus, the composition of this sporomorph assemblage is probably
less directly related to local vegetation and should be considered as
allochthonous.
Sample ISP-3 came from the bed that yielded the largest number
of plant megafossils. It is characterized by the occurrence of large,
lentoid phycomes attributed to prasinophytes (Tasmanites sp.).
These planktonic algae indicate either freshwater lacustrine conditions (Tappan, 1980) or a deltaic area of reduced salinity (GuyOhlson, 1996; Prauss, 1996; Martnez et al., 2005).
In sample ISP-4, the abundance of well-preserved fern spores,
the occurrence of conifer pollen, Araucariacites australis, with low
buoyancy (Peyrot et al., 2008, 2011), and the scarcity of bisaccate
conifer pollen suggest a relative proximity to the source area.

Fig. 5. Sporomorphs from the Isona Sud locality. A, Biretisporites potoniaei Delcourt & Sprumont; ISP-1. B, Acylomurus sejunctus Phillips; ISP-1. C, Matthesisporites plurituberosus
ring; ISP-1. D, Anapiculatisporites dawsonensis Reis and Willians; ISP-4. E, Cicatricosisporites hallei Delcourt and Sprumont; ISP-1. F, Concavissimisporites subgranulosus (Couper)
Do
Pocock; ISP-2. G, Cyathidites australis Couper; ISP-2. H-J, Klukisporites scaberis (Cookson and Dettmann) Dettmann; ISP-2. K, Anapiculatisporites sp.; ISP-2. L, Echinatisporis longechinus
Krutzsch; ISP-2. M, Echinatisporis sp.; ISP-1. N, Trilites tuberculiformis (Cookson) Dettmann; ISP-1. O, Patellasporites sp.; ISP-2. P, Leptolepidites verrucatus Couper; ISP-1. Q, Leiotriletes
verrucatus Copuer; ISP-2. R, Polypodiaceoisporites sp.; ISP-3. Scale bar 20 mm.

Fig. 6. Sporomorphs from the Isona Sud locality (continued). A, Leiotriletes dorogensis Kedves; ISP-2. B, Leiotriletes adriennis (R. Pot and Gell.) Krutzsch; ISP-3. C, Foveotriletes sp.; ISP4. D, Monosulcites sp.; ISP-2. E, Laevigatosporites ovatus Wilson and Webster; ISP-1. F, Laevigatosporites haardti (R. Pot and Ven) Th. and Pf.; ISP-2. G, Monosulcites brevispinosus Sah
and Dutta; ISP-1. H, Chomotriletes fragilis Pocock; ISP-3. I, Monocolpopollenites sp.; ISP-3. J, Araucariacites australis Cookson; ISP-1. K, Pinuspollenites ruginosa (Stanley) Oltz; ISP-2. L,
Cycadopites carpentieri (Delcourt and Sprumont) Singh; ISP-4. M, Polyporopollenites sp.; ISP-4. N, Rugulitriporites sp.; ISP-1. O, Subtriporopollenites sp.; ISP-3. P, Subtriporopollenites
sp.; ISP-4. Q, Subtriporopollenites constans Pf.; ISP-4. R, Nudopollis sp.; ISP-1. S, Pseudoromeinipollenites sp.; ISP-4. T, Pseudoromeinipollenites sp.; ISP-2. U, Tasmanites sp.; ISP-3. Scale
bar 20 mm.

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

45

7. Discussion

Fig. 7. Diagram showing the abundance of the most representative morphological

groups of sporomorphs in the studied samples.

Furthermore, the relative abundance of freshwater green algae in

this sample indicates dominant freshwater conditions during
deposition of the bed.

The plant megafossil assemblage of Isona Sud closely resembles

that studied by Vicente i Castells, (2002) and recently reviewed by
Marmi et al. (2014). Both have a large proportion of angiosperms
compared to gymnosperms (ratio of 4:1), with eudicot leaves representing the most abundant group. Some specimens probably
belong to the same taxa in the two assemblages (e.g. Cunninghamites). Several eudicot forms collected during the 2010 excavation share key morphological features with specimens in the
Vicente i Castells collections that were in some cases identied as
different taxa (e.g. Eudicot forms 1, 2 and 5). These afnities support a strong relationship between our specimens and those
collected by Vicente i Castells and collaborators in the early 1980s.
More importantly, the lithological features shared by the two collections also support that they represent the same beds. Furthermore, a signicant contribution of this study is to clarify the
preservation status of the Upper Cretaceous plants from Isona and
the sampling biases in the Vicente i Castells plant fossil collections
ncies Naturals de Barcelona. Clearly, the
housed in the Museu de Cie
previous view that the Isona ora consisted only of large and
generally well-preserved leaf laminas should be now abandoned.

Fig. 8. Plant taphonomic features from the early Maastrichtian of the Tremp Basin. A, Eudicot leaf without petiole, MCD 5312. B, Eudicot leaf showing tears following the secondary
venation; MCD 5298. C, Fragment of a Eudicot leaf showing insect or fungal marks; MCD 5323. D, Stem compressions without preferential alignment.

46

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

Fig. 9. Plant palaeoecological and palaeoenvironmental reconstruction during the early Maastrichtian in the Tremp Basin, inferred from data of Isona Sud (oodplain) and the
Barranc de la Posa (wetland) localities.

S. Villalba-Breva et al. / Cretaceous Research 54 (2015) 34e49

The plant fossil assemblage of Isona shares several features with

other Cretaceous oras of Europe. For instance, Cunninghamites is
reported from the Cenomanian of the Czech Republic (Kva
cek,
1999), the Santonian and Maastrichtian of the Netherlands
(Bosma et al., 2009; Van der Ham et al., 2004) and the CampanianeMaastrichtian of Poland (Halamski, 2013). Podozamites has
been reported from the Cenomanian of the Czech Republic and the
lower Campanian of Austria (Herman and Kvacek, 2007, 2010, and
references therein). The taxonomy of the Isona angiosperm leaves
is poor. However, as an example, Eudicot form 1 resembles Salix
afnis from the Albian of Portugal (Saporta, 1894) and Dicotylophyllum proteoides from the lower Campanian of Grnbach,
Austria (Herman and Kva
cek, 2010). According to these authors,
D. proteoides-like leaves are widespread in the Late Cretaceous of
the Northern Hemisphere.

dus et al.
Dealing with palynology, Porta et al. (1985) and Me
(1988) studied the sporomoph assemblages from the Upper
Cretaceous of the Barranc de la Posa section in the Tremp Basin,
which includes the la Posa Unit and the overlying Xulli Unit.
Samples for sporomorphs were collected from the upper part of the
la Posa Unit and the lower part of the Xulli Unit. The distribution of
sporomorphs found by these authors differs signicantly from our
results. In the Barranc de la Posa, there are fewer spores, which in
no case exceed 40%, but there is a higher proportion of monocolpates, which correspond to different pollen forms in Isona Sud.
However the assemblage of Isona Sud is signicantly similar to the
sporomorph assemblage studied by Ashraf and Erben (1986) in the

, Barranc de la Munya and Barranc
Barranc de la Posa, Coll de Nargo
de Basturs sections of the Tremp Basin. In particular, the pollen of
Cycadales and Pinaceae and a signicant proportion of the spores
(47%) of the Isona Sud assemblage match those described from the
above-listed sections. The angiosperms of Isona Sud are less
abundant than in the Barranc de la Posa section, according to data
by Porta et al. (1985). In Isona Sud the Normapolles group is
identied, with Semioculopollis sp., Trudopollis sp. and
Pseudoromeinipollenites sp. In addition, pollen of Platycaryapollenites ferrerri, Rugulitriporites sp., Triatriopollenites sp. and
Subtriporopollenites sp. were found. These last four species constituted only a minor component in the assemblages reported by
Ashraf and Erben (1986). The planktonic algae Tasmanites sp. of
Isona Sud is very similar to specimens that Ashraf and Erben (1986)
assigned to Leiosphaeridia deandrei Madler.
The sporomorph assemblage of Isona Sud contains diverse trilete spores, while gymnosperm and angiosperm pollen are less
abundant. This resembles the assemblage from the Maastrichtian of
the Hateg Basin in Romania (Van Itterbeeck et al., 2005), while it
contrasts with the assemblages from the Netherlands, especially
from the Gulpen Formation (Kedves and Herngreen, 1980) and the
Bunde Borehole (Herngreen et al., 1986), which show predominant
Normapolles taxa. These differences may be palaeogeographically
driven, as Porta et al. (1985) already noted. The north European
assemblages belong to the Boreal realm of the Normapolles
Province, while the Pyrenean, the Ibero-Lusitanian and the Carpathian basins form the three subunits of the Mediterranean realm of
this province (Herngreen et al., 1996).
Angiosperm and gymnosperm pollen percentages are consistent in their abundance with the plant megafossil assemblage of
Isona Sud, but one may question why spore-producing plant
megafossils are lacking in the record. As has been often suggested
in the literature, ferns may be not preserved due to their generally
poorly cutinized leaves. Moreover, being generally understory
plants, ferns are often under-represented in leaf-litter assemblages. Finally, some ferns retain their marcescent leaves attached
to the stems and thus they stand little chance of entering the fossil
record.

47

Lastly, the plant assemblage of Isona Sud can be compared with

other lower Maastrichtian plant assemblages from the neighbouring locality of the Barranc de la Posa (Tremp Basin) and from
Fumanya (Vallcebre Basin). The assemblage of Fumanya is mainly
formed by leafy axes of the conifer Frenelopsis sp. and the palm
leaves, logs and rooting systems of Sabalites longirhachis (Unger) J.
Kvacek and Herman (Villalba-Breva et al., 2012). Less abundant
remains include cycadalean and monocot (probably pandanacean)
leaves. Abundant minute monocot (probably commelinid) seeds of
Bergacarpon viladricii Marmi, Gomez, Villalba-Breva et MartnClosas were also described (Marmi et al., 2012). Villalba-Breva
et al. (2012) interpreted the assemblage dominated by Frenelopsis and Sabalites as growing in the margins of coastal, freshwater lakes, while other leaves and seeds were allochthonous. In
contrast, the assemblage of the Barranc de la Posa is formed
exclusively by Frenelopsis leafy axes, which are found in lignites
and black marls rich in organic matter. According to Villalba-Breva
and Martn-Closas (2013) these lignite beds resulted from a parautochthonous accumulation of organic matter in a coastalswamp environment of variable salinity connected to shallow
freshwater lakes.
The differences between the plant assemblages of the Barranc
de la Posa and Isona Sud can be understood in terms of plant
palaeoecology and environmental factors. Thus, at the beginning of
the Maastrichtian in the Tremp Basin, Frenelopsis appeared to
inhabit wetlands close to the sea border, whereas some eudicot
angiosperms and ferns dominated the river systems as riparian
elements. The habitats of other plants represented by allochthonous remains are difcult to ascertain. However, they would be
located farther upstream (Fig. 9). These new data are useful for
completing and rening the previous works by Coiffard et al. (2006,
2007, 2012) and Coiffard and Gomez (2010), who analysed the rise
to dominance of angiosperms from the Barremian to the Campanian of Europe in a variety of ecosystems, from freshwater lakerelated wetlands to oodplains and coastal swamps. Even after a
long history of colonization of diverse environmental settings by
angiosperms, it is noteworthy that at the end of the Cretaceous,
cheirolepidiacean conifers still occupied the coastal swamps.
Acknowledgements
This study is a contribution to projects CGL2011-27869,
CGL2011-30069-C02-01/02, CGL2011e23948 and CGL2012-35199
of the Spanish Ministerio de Economa y Competitividad. Support
was also provided by projects CGL2009e11838/BTE and CGL2009
n, and
09000/BTE of the Spanish Ministerio de Ciencia e Innovacio
project 2014SGR251 of the Catalan government. Fieldwork at the
Isona Sud locality was authorized by the Department of the Culture

n
of the Catalan government. We acknowledge C. Coiffard, Q. Guzma

s for their collaboration in the excavation. The
and A. Garcia-Selle
research of SV-B was nanced with a travel grant from the Facultat
de Geologia, Universitat de Barcelona, and with a grant from la

gion Rho
^ne-Alpes. B.G. and V.D.-G. were funded by the CNRSRe
UMR5276. The English text was corrected by S. Keddie and C.
Evans (Serveis Lingstics, Universitat de Barcelona). We acknowledge the contribution of C. Wood and J. Kva
cek in improving the
original manuscript during the peer-review process.
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