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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.
35
Fig. 1. Geological map of the Tremp Basin with the location of the studied area.
36
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
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
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.
39
40
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.
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
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.
45
7. Discussion
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
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.
47
48
r Europas.
Kunzmann, L., 1999. Koniferen der Oberkreide und ihre Relikte im Terta
Abhandlungen des Staatlichen Museums fr Mineralogie und Geologie zu
Dresden 45, 1e190.
Kunzmann, L., 2010. Geinitzia reichenbachii (Geinitz, 1842) Hollick and Jeffrey, 1909
and Sedites rabenhorstii Geinitz, 1842 (Pinopsida; Late Cretaceous) reconsidered
and redescribed. Review of Palaeobotany and Palynology 159, 123e140.
ognostique des Petites Pyre
ne
es et particulie
rement
Leymerie, A., 1862. Aperu ge
te
Ge
ologique de France 19,
de la montagne dAusseing. Bulletin de la Socie
1091e1096.
mez
Liebau, A., 1973. El Maastrichtiense lagunar (Garumniense) de Isona. In: Go
Angulo, J.A. (Ed.), XIII Coloquio Europeo de Micropaleontologa. Empresa
Nacional ADARO, Madrid, pp. 87e113.
pez-Martnez, N., Fern
n, M.T., Valle, M.F., 1999. The succession of
Lo
andez-Marro
vertebrates and plants across the Cretaceous-Tertiary boundary in the Tremp
Formation, Ager valley (South-central Pyrenees, Spain). Geobios 32, 617e627.
Marmi, J., Gomez, B., Martn-Closas, C., 2008. Presencia de macrorestos paractonos de Sabalites cf. longirhachis (Unger, 1850) Kva
uto
cek & Herman, 2004 en
licas del Creta
cico superior del Pirineo oriental. Revista Espan
~ ola de
facies para
Paleontologa 23, 7e14.
Marmi, J., Gomez, B., Martn-Closas, C., Villalba-Breva, S., 2010. A reconstruction of
the fossil palm Sabalites longirhachis (Unger) J. Kva
cek et Herman from the
Maastrichtian of Pyrenees. Review of Palaeobotany and Palynology 163, 73e83.
Marmi, J., Gomez, B., Villalba-Breva, S., Martn-Closas, C., 2012. Bergacarpon viladricii
gen. et sp. nov., angiosperm seeds and associated fruits from the early Maastrichtian of the eastern Pyrenees (Catalonia, Spain). Review of Palaeobotany and
Palynology 171, 83e94.
Marmi, J., Gomez, B., Martn-Closas, C., Villalba-Breva, S., Daviero-Gomez, V., 2014.
Diversied fossil plant assemblages from the Maastrichtian in Isona (southeastern Pyrenees). Review of Palaeobotany and Palynology 206, 45e59.
tations
Martn-Closas, C., Gomez, B., 2004. Taphonomie des plantes et interpre
oe
cologiques. Une synthe
se. Geobios 37, 65e88.
pale
lisis palinolo
gico de la
Martnez, M.A., Quattrocchio, M.E., Pramparo, M.B., 2005. Ana
n Los Molles, Grupo Cuyo, Jura
sico medio de la cuenca Neuquina,
Formacio
Argentina. Ameghiniana 42, 67e92.
hler, H., Tiedemann, R.,
Mayr, C., Thmmler, B., Windmaier, G., Altenbach, A.V., Ko
1999. New data about the Maastrichtian/Danian transition in the southern
~ ola de Micropaleontologa
Pyrenees (Ager Basin, Catalonia, Spain). Revista Espan
31, 357e368.
dus, J., 1972. Palynological zonation of the Upper Cretaceous in southern France
Me
and northeastern Spain. Review of Palaeobotany and Palynology 14, 287e295.
dus, J., Feist, M., Rocchia, R., Batten, D.J., Boclet, D., Colombo, F., Tambareau, Y.,
Me
Villate, J., 1988. Prospects for recognition of the palynological Cretaceous/Tertiary boundary and an iridium anomaly in non-marine facies of the eastern
Spanish Pyrenees: a preliminary report. Newsletters on Stratigraphy 18,
123e138.
Mey, P.H.W., Nagtegaal, P.J.C., Roberti, K.J., Hartevelt, J.J.A., 1968. Lithostratigraphic
subdivision of post-Hercinian deposits in the south-central Pyrenees, Spain.
Leidse Geologische Mededelingen 41, 21e228.
~ oz, J.A., 1989. The structure of the Pyrenees. In: Marzo, M., Puigdefa
bregas, C.
Mun
(Eds.), Alluvial deposits of the successive foreland basin stages and their relation to the Pyrenean thrust sequences, 4th International Conference on Fluvial
gic de CatSedimentology, Guidebook series, Excursion num. 10. Servei Geolo
alunya, Barcelona, pp. 7e13.
Nagtegaal, P.J.C., 1972. Depositional history and clay minerals of the Upper Cretaceous basin in the South-Central Pyrenees, Spain. Leidse Geologische Mededelingen 47, 251e275.
Nichols, D.J., Johnson, K.R., 2008. Plants and the K-T boundary. Cambridge University Press, Cambridge, 280 pp.
n, E., Comas-Rengifo, M.J., Barroso-Barcenilla, F., Feist-Burkhart, S.,
Peyrot, D., Barro
n de
2008. Palinologa del tr
ansito Cenomaniense/Turoniense en la seccio
~ a). Coloquios de Paleontologa 58, 101e161.
Puentedey (Burgos, Espan
n, E., Comas Rengifo, M.J., 2011. PalaePeyrot, D., Barroso-Barcenilla, F., Barro
oenvironmental analysis of Cenomanian-Turonian dinocyst assemblages from
the Castilian Platform (Northern-Central Spain). Cretaceous Research 32,
504e526.
de Porta, N., Civis, J., 1985. Palinologa del MaasPorta, J., de Kedves, M., Sole
rida, Espan
~ a). Problem
trichtiense del Barranco de la Posa (Le
atica regional.
giques 40, 5e28.
Revista dInvestigacions Geolo
Prauss, M., 1996. The lower Toarcian Posidonia Shale of Grimmen, Northeast Germany. Implications for the palynological analysis of a near-shore section. Neues
ontologie, Abhandlungen 200, 107e132.
Jahrbuch fr Geologie und Pala
bregas, C., Mun
~ oz, J.A., Marzo, M., 1986. Thrust belt development in the
Puigdefa
eastern Pyrenees and related depositional sequences in the southern foreland
basin. In: Allen, P.A., Homewood, P. (Eds.), Foreland basins. Blackwell, Oxford,
pp. 229e246.
Rasband, W., 1997e2008. ImageJ: Image processing and analysis in Java [version
1.40g]. http://rsb.info.nih.gov/ij/index.html.
2009. The end-Cretaceous dinosaur sucRiera, V., Oms, O., Gaete, R., Galobart, A.,
cession in Europe: The Tremp Basin record (Spain). Palaeogeography, Palaeoclimatology, Palaeoecology 283, 160e171.
Riera, V., Marmi, J., Oms, O., Gomez, B., 2010. Orientated plant fragments revealing
tidal palaeocurrents in the Fumanya mudat (Maastrichtian, southern Pyrenees): Insights in palaeogeographic reconstructions. Palaeogeography, Palaeoclimatology, Palaeoecology 288, 82e92.
49
Cretaceous, SE Netherlands) rediscovered. Bulletin de l'Institut royal des Sciences naturelles de Belgique. Sciences de la Terre 74, 89e96.
Van Itterbeeck, J., Markevich, V.S., Codrea, V., 2005. Palynostratigraphy of the
^ul Mare and Barbat Valleys
Maastrichtian dinosaur- and mammal sites of the Ra
(Haeg Basin, Romania). Geologica Carpathica 56, 137e147.
gic de la ora cret
Vicente i Castells, J., 2002. Estudi morfolo
acica dIsona (Pallasrs
s Nord, Se
rie Monogr
Juss
a). Institut dEstudis de la Natura del Barcelone
aca 2,
Santa Coloma de Gramanet, 223 pp.
Villalba-Breva, S., Martn-Closas, C., 2013. Upper Cretaceous palaeogeography of the
Central Southern Pyrenean Basins (Catalonia, Spain) from microfacies analysis
and charophyte biostratigraphy. Facies 59, 319e345.
ndez-Marro
n, M.T.,
Villalba-Breva, S., Martn-Closas, C., Marmi, J., Gomez, B., Ferna
2012. Peat-forming plants in the Maastrichtian coals of the Eastern Pyrenees.
Geologica Acta 10, 189e207.