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AOBPreview originally published online on August 6, 2007
Annals of Botany 2007 100(3):545-553; doi:10.1093/aob/mcm160
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© The Author 2007. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oxfordjournals.org

Early Cretaceous Angiosperm Invasion of Western Europe and Major Environmental Changes

C. Coiffard1, B. Gomez2,* and F. Thevenard1

1 UCB Lyon 1 et UMR 5125, Paléobotanique, 7 rue Dubois, F-69622 Villeurbanne, France
2 UMR 6118 du CNRS Géosciences, Université Rennes 1, Campus de Beaulieu, avenue du Général Leclerc, F-35042 Rennes, France

* For correspondence. E-mail bernard.gomez{at}univ-rennes1.fr

Received: 8 February 2007    Returned for revision: 26 March 2007    Accepted: 16 May 2007    Published electronically: 6 August 2007


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 

Background and Aims: At the beginning of the Late Cretaceous, angiosperms already inhabited all the environments and overtopped previously gymnosperm-dominated floras, especially in disturbed freshwater-related environments. The aim of this paper is to define what fossil plant ecology occurred during the early Cretaceous in order to follow the early spread of angiosperm taxa.

Methods: Floristic lists and localities from the Barremian to the Albian of Europe are analysed with the Wagner's Parsimony Method.

Key results: The Wagner's Parsimony Method indicates that (a) during the Barremian, matoniaceous ferns formed a savannah-like vegetation, while angiosperms composed freshwater aquatic vegetation; (b) during the Late Aptian humid phase, conifers increased, while matoniaceous ferns decreased, reflecting the closure of the vegetation; and (c) from the Albian, warmer and drier conditions induced the recovery of the matoniaceous ferns, while core angiosperms first developed in floodplains.

Conclusions: During the late Early Cretaceous (Barremian–Albian), angiosperms showed a stepwise widening of their ecological range, being recorded first during the Barremian as aquatic plant mega-remains and at the Cenomanian onwards occurred in all the environments.

Key words: Wagner's Parsimony Method, angiosperms, conifers, gymnosperms, ecology, environment, upper Lower Cretaceous, Europe


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 
Coiffard et al. (2004, 2006) used Wagner's Parsimony Method (WPM), a clustering method currently used in phylogeny (cladistics), to infer the relationships between the palaeobiocoenoses (floristic lists of fossil plant mega-remains) and the palaeoenvironments (biotopes), and showed that the angiosperms already had a wide ecological range during the latest Albian–Cenomanian. Nevertheless, the worldwide fossil record of angiosperm mega-remains extends back before the Albian: (a) Archaefructus, Yixian Formation, Liaoning, China (Sun et al., 2002); (b) several angiosperm assemblages, zones I and IIb of the Potomac Group, USA (Hickey and Doyle, 1977); (c) Araripa, Endressianthus, Klitzschophyllites, Crato Formation, Brazil (Mohr et al., 2006); and (d) Kachaikenia, Kachaike Formation, Argentina (Cúneo and Gandolfo, 2005).

Recent hypotheses for the origins and rise to dominance of angiosperms emphasize the roles of early angiosperm ecology and environments: (a) the ‘Paleoherb’ hypothesis – early angiosperms consisted of riparian weeds growing near stream-channel margins (Taylor and Hickey, 1992, 1996); (b) the ‘Wet and Wild’ hypothesis – early angiosperms were mostly aquatic herbaceous plants such as Archaefructus living in wetlands (Sun et al., 2002; D. L. Dilcher, pers. comm.); and (c) the ‘Dark and Disturbed’ hypothesis – early angiosperms were mainly composed of shrubs such as living Amborella and Austrobaileyales and inhabited disturbed understorey (Feild et al., 2004). In addition, the Cretaceous climate was less stable than previously thought (Francis and Frakes, 1993), and appears to be one of the main environmental factors in the evolution of early angiosperms. Temperature shifts, sea level fluctuations and ocean anoxic events (OAE in Fig. 1) occurred, partly related to volcanic activity (Fig. 1). The Barremian of Europe corresponded to a drought phase, as deduced by the presences of evaporites, carbonate-rich sediments, red beds, firm grounds, carbonate nodules, pyrite concretions, kaolinite depletion and low morphological diversity of the spore assemblages (Ruffel and Batten, 1990). The drought phase ended at about the Middle Aptian. It was followed by a wetter and 5 °C colder climate evidenced from the occurrence of detritic quartz deposits in the Vocontian Trough (Wortman, 2004) and oxygen isotopes in rudists (Steuber, 2005), respectively. The Lower Albian of Europe was a drier and semi-arid climate with a marked seasonality as recorded by the clay mineralogy (Mutterlose et al., 2003), and a further warming of about 10 °C was recorded from oxygen isotopes of foraminifers in the North Atlantic (Leckie, 2002; Price and Hart, 2002). The Upper Albian of Europe was a new wet period as indicated by clay mineralogy (Fenner, 2001).


Figure 1
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  		FIG. 1. Main global changes proxies for the Mid-Cretaceous. 1, Steuber et al. (2005), rudists; 2, Stoll and Schrag (2000), foraminifers; 3, Leckie (2002) and Haq et al. (1988). OAE = Ocean anoxic events.

 
In Europe, recent palaeobotanical studies of early angiosperms focused on a few records of taxa which were probably aquatic: Bevhalstia pebja (Hauterivian–Barremian of England; Hill, 1996), Ranunculus ferrerii (Barremian of Spain; Blanc-Louvel, 1984) and water lily-like flower (Lower Cretaceous of Portugal; Friis et al., 2001). Despite the considerable Lower Cretaceous record of leaf mega-remains and the broad range and generally accurate stratigraphy already known from many countries (e.g. England, France, Germany, Portugal and Spain), these records were not analysed from a palaeoecological perspective. Here the WPM is used to test the most current hypotheses on early angiosperms and whether or not their rise to dominance during the Lower Cretaceous was driven by ecological and environmental factors.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 
Excluding the latest Albian–Cenomanian (100–93 Ma) data published by Coiffard et al. (2006), a new database was built for the Barremian–Albian (127–100 Ma). These datasets were independently studied because the two time intervals show completely distinct floras.

The database (Appendix 1) consists of a set of floristic lists of fossil plant impressions and compressions ordered according to a ‘presence/absence’ per locality matrix. It is compiled from the Barremian–Albian of Germany (Depape, 1965; Daber, 1968), Spain (Depape and Doubinger, 1960; Alvárez-Ramis et al., 1981; Alvárez-Ramis and De Lorenzo, 1982; Román-Gómez, 1985, 1987; Barale and Doludenko, 1993; Gomez et al., 1999, 2000, 2001, 2002), France (Carpentier, 1927), Poland (Reymanowna, 1965), Portugal (De Saporta, 1894; Teixeira, 1948) and the United Kingdom (Harris, 1981; Ross and Cook, 1995; Hill, 1996) and from personal data of B.G. Although many of the identifications were published over 50 years ago, no revision or re-examination of these floras exists. In addition, in several cases it is known that collections were lost or damaged. The present analysis is based entirely on the original authors. This database includes 38 localities and 115 taxa (of which 22 are angiosperm species). The localities were grouped according to the species content with the WPM.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 
The single most-parsimonious tree obtained from the floristic lists has a length of 164 steps, a consistency index of 0·4615, a retention index of 0·6733 and a rescaled consistency index of 0·4722. The tree is rooted in order to separate the Albian from the Barremian localities. The rooting allows for a hierarchy on the tree (e.g. the highest hierarchical level corresponds to the split between the Barremian and Aptian–Albian floras). The branches are subdivided into eight clades (1–8) and 12 groups (A–L). The groups are clades (Fig. 2) that contain localities sharing two to eight (usually six) taxa, whereas localities between two groups share two or fewer taxa (Table 1).


Figure 2
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  		FIG. 2. Consensus tree of the floristic lists from the Barremian–Albian of Europe showing 12 groups of localities [A, Aptian–Albian aquatic; B, Albian aquatic; C, Albian backswamps; D, Aptian-Albian channels; E, Aptian–Albian floodplains; F, Albian floodplains; G, Barremian channels; H, Barremian aquatic; I, Barremian backswamps; J, pre-Aptian (Valanginian?) floodplains; K, Barremian floodplains; L, Barremian salt marshes; numbers refer to clades cited in the text] and age of each locality (Wea, wealdian; Val, Valanginian; Ha/Ba, Hauterivian/Barremian; Bar, Barremian; Apt, Aptian; Ap/Al, Aptian–Albian; Alb, Albian).

 


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  		TABLE 1. Typical taxa of each group occuring in more than half of the localities of a given group

 


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  		TABLE 2. Ages and palaeoenvironments of the localities studied (B, Belgium; D, Germany; F, France; P, Portugal; PL, Poland; S, Spain; UK, United Kingdom)

 
The grouping fits well with both environments and stratigraphy (Table 2). Living plant associations and their life environments are strongly correlated and, in the fossil records analysed herein, one given plant assemblage always occurs in the same sedimentological setting and vice versa. Thus, the analysis is also consistent in that assemblages reflect local biocoenoses. Group G (Fig. 2) does not correspond to a clade because it is mainly based on the absences of taxa. The taxa occurring in group G (e.g. Weichselia reticulata) can also be found in other groups.

Barremian
The floras from the Barremian localities (see Fig. 2, 1 G–L) are mainly composed of ferns Weichselia reticulata and Phlebopteris dunkeri. Three of them, Quedlindburg (Germany), Beare Green and Smockjacks Brickworks (UK), are placed at the base of the main branch and are clustered in group G (Fig. 2, G). G represents clay lenses generally included in coarse-grained sandstones, and are considered to be channel deposits. The remaining clades (Fig. 2, 2) consist of ferns Cladophlebis browniana and Onychiopsis psilotoides. Within clade 2, group K (Fig. 2, K) is characterized by a fern, Gleichenites nordenskioeldii, a bennettite, Zamites buchianus, and a conifer, Cyparissidium gracile. Laminated clays of K suggesting more quiet deposits and without any coaly (coal-like) remains may correspond to levee or floodplain deposits (Préjano, Spain; lower and upper part of Féron–Glageon white clays, France). Clade 3 (Fig. 2, 3) typically lacks Phlebopteris dunkeri. Group J (Fig. 2, J) shows ferns Pteridoleima spoliatum and Sphenopteris sinualoba. As in group K, the sediments in J indicate quiet depositions in levees or floodplains (Quinta do Leirião, Railroad 66, Portugal). Clade 4 (Fig. 2, 4) displays a conifer, Sphenolepis kurriana. Group H (Fig. 2, H) bears notably the aquatic angiosperm Montsechia vidalii collected from lithographic limestones deposited in freshwater lakes. Clade I (Fig. 2, I) contains a conifer, Pinites solmsii, and a Caytoniale, Sagenopteris variabilis. As far as is known, the sedimentology of I was not documented, but coaly lenses bearing Pinites solmsii (Daber, 1960) might indicate swampy conditions. The remaining Barremian localities, group L (Fig. 2, L), share a fern, Weichselia reticulata and a conifer, Frenelopsis hoheneggeri, and represent brackish water-related environments.

Aptian–Albian
Group A of the Aptian–Albian localities (Fig. 2, A) involves ferns s.l., Matonidium goepperti, Onychiopsis psilotoides, Isoetites choffati, and angiosperms, Quercus arnalensis and especially the aquatic Nymphaeites spp. Nevertheless, Casal da Quintã (Portugal) is older (Valanginian) and lacks angiosperms. The laminated clays (Casal da Quintã, Portugal) and the aquatic plants suggest freshwater lakes or ponds. Clade 5 (Fig. 2, 5) comprises numerous localities bearing a conifer, Brachyphyllum obesum. Group F (Fig. 2, F) includes ferns, Adiantum eximium, Onychiopsis psilotoides, Sphenopteris involvens, and angiosperms, Brasienopsis venulosa, Cissites spp., Menispermites spp. and Proteophyllum dissectum. As in groups J and K, F may correspond to levees or floodplains. Clade 6 (Fig. 2, 6) exhibits a conifer, Sphenolepis kurriana. Group D of the clade 7 (Fig. 2, 7 and D) bears Sphenolepis debile, and clay lenses (Ruña, upper part of Buarcos (Portugal)) or ferruginous crusts (La Cierva, Spain) generally included in coarse-grained sandstones (conglomerates in the case of Ruña, upper part of Buarcos and La Cierva) may be considered as channel deposits. Group E (Fig. 2, E) is composed of ferns, Cladophlebis browniana and Sphenopteris tenuifissa, and the bennettite, Pseudocycas tenuisectus. Laminated clays not associated with coaly deposits of E may indicate depositions in levees or floodplains. In addition, E and F differ in age, E being Aptian–Albian and F Middle–Upper Albian. Clade 8 (Fig. 2, 8) shows a fern, Onychiopsis psilotoides, and a cycad, Almargemia dentata. More specifically, Group B (Fig. 2, B) contains the aquatic angiosperms Choffatia francheti, Hydrocotylophyllum lusitanicum and Nymphaeites spp. B only consists of clay lenses (Cercal, Portugal and Pola de Siero, Spain), and was probably deposited in freshwater lakes or ponds. Although Groups A and B share aquatic taxa, they are clearly distinguished in the tree because A is Aptian–Albian and B is Middle–Upper Albian. Group C (Fig. 2, C) associates ferns Acrostichopteris nervosa and Laccopteris pulchella and gymnosperms Desmiophyllum latifolium and Frenelopsis hoheneggeri. Group C shows coaly laminated clays (Sao Mamede and Belas, Portugal) probably indicating swampy environments.


   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 
Ecology and environments
Grime (2001) distinguishes three primary life history strategies for plants – ruderal, competitive and stress tolerant – which correspond to different types of habitats. Ruderal strategies are associated with disturbed environments that are located in stream channel margins or habitats subject to frequent flooding, fire or mass wasting. Competitive strategies are associated with mesic environments that are located in optimal water, nutrient and temperature-supplied habitats. Finally, stress-tolerant strategies are associated with stressed environments that are located in swamps, salt marshes and suboptimal growth habitats.

In Europe, during the Lower Cretaceous, examples of the ruderal strategy occurred amongst ferns. Examples of this are Gleichenites nordenskioldii (Gleicheniaceae) and Phlebopteris dunkeri (Matoniaceae), which show many analogies with their living relatives Gleichenia and Matonia and may have formed a dense fern thicket (Watson and Alvin, 1996); Weichselia reticulata (Matoniaceae) that displays a different organisation, unknown within the living relatives, with a trunk reaching 15 cm in diameter and exhibiting numerous xerophilous characters (e.g. sunken stomata, thick cuticles, etc.); and the Lower Cretaceous European Gleicheniaceae and Matoniaceae that were found nearly exclusively in channel deposits (Fig. 3), and often formed charcoals. The ruderal strategy of these fossil taxa is supported since they occupied environments disturbed by both floods and fires.


Figure 3
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  		FIG. 3. Block diagram showing the means of the numbers of species of the main component taxa (e.g. angiosperms) in each of the environments (n = number of localities per environment).

 
Competitors are mainly found in stable environments such as floodplains. They may include the dicksoniaceous fern Onychiopsis psilotoides that grew in a wide range of environments during the Lower Cretaceous (Fig. 3). Onychiopsis psilotoides shows xerophilous characters (Watson and Alvin, 1996), but there is no support for a tree habit similar to that in the living Dicksoniaceae. During the Barremian, the conifers occupied all the environments except the channels (Fig. 3), and may also have been competitors. The absence of conifers in the disturbed channel environments may be explained by the low competitiveness of trees under drought climates. Thus, Brachyphyllum obesum and Sphenolepis debile colonized these environments during the Aptian–Albian humid phase. Conifers also became widely distributed, and took in the flora and plant ecology a place comparable to the Barremian Matoniaceae.

Competitive and shade-tolerant strategies are not easily distinguished in the fossil record. Thus, at the macro-habitat scale, competitors can form the overstorey and species which tolerate shade comprise the understorey. Osmundaceae and Bennettitales were found mainly in stable environments (floodplains and swamps; Fig. 3). Such a distribution suggests a competitive strategy or more probably a shade-tolerant strategy, e.g. in Cladophlebis species, the conifers being more probably all tall trees producing shade. The shade-tolerant strategy of the Lower Cretaceous Cladophlebis species fits well with a humid understorey habitat of the living Osmundaceae.

The Albian shows the earliest occurrences of floodplain angiosperms in Europe (Fig. 3), and these angiosperms appear to have replaced Osmundaceae, and probably had a shade-tolerant strategy. They display characters of the eudicots (Cissites, Ranunculales?) and perhaps of the Piperales [e.g. Teixeira (1950) grouped Aristolochia daveauana De Saporta with Menispermites cercidifolius De Saporta]. The co-occurrence of eudicots and Piperales leaves is not surprising because flowers and fruits are known in beds of similar ages (Friis et al., 1995; Von Balthazar et al., 2005). These understorey angiosperm taxa show close affinities to Cissites parvifolius and Menispermites reniformis from the Potomac flora of Northern America. Only two equivalent taxa remained during the Cenomanian and grew in floodplain environments (Coiffard et al., 2006). In the Czech Republic ‘Aralia’ formosa and Hederaephyllum primordiale, and in Portugal Aralia calomorpha (of uncertain affinity) and Menispermites cercidifolius (‘Aralia’ formosa and Aralia calomorpha appear to be synonyms; the same applies to Hederaephyllum primordiale and Menispermites cercidifolius). Similar associations of A. calomorpha/M. cercidifolius and A. formosa/H. primordiale in the Albian and Cenomanian, respectively, persisted in floodplain environments.

The stress-tolerant taxa are mainly found in lakes and swamps, and in the freshwater-related swampy environments include Matoniaceae (Laccopteris pulchella and Matonidium goepperti). In the latter environments, a few gymnosperms, and especially Cycadales such as Almargemia dentata, were restricted (Fig. 3). Although one may question the occurrence of fossil Cycadales in wetlands, it is not so surprising when thinking of Ceratozamia, a living relative of Almargemia according to Brenner et al. (2003), which occupies cloud forests (Jones, 1993). One may assume that these Lower Cretaceous plants mostly tolerated anoxic and nutrient-depleted soils. Weichselia reticulata, Frenelopsis and Eretmophyllum/Nehvizdya [i.e. the use of Eretmophyllum (Thomas) emend. Harris et al. by Kvacek (1999) and Kvacek et al. (2005) versus Nehvizdya Hlustík by Gomez et al. (2000) is still in debate], which grew in brackish water-related environments, were mainly influenced by the water supply. Such a brackish ecology was recently suggested for one of the species of the genus Eretmophyllum/Nehvizdya, E. obtusum from the Middle-Upper Cenomanian of Bohemian Massif (Falcon-Lang et al., 2006).

As far as aquatic angiosperms are concerned, they diversified from the Barremian to the Albian: (a) a few Barremian taxa of unknown affinities (Montsechia vidalii); (b) Aptian–Albian taxa showing affinities to Nympheales (Nymphaeites) or monocots (Klitzschophyllites); and (c) Lower Albian showing the same taxa in addition to Choffatia francheti and Hydrocotylophyllum lusitanicum, the latter sharing affinity with the Piperales (Cantrill and Nichols, 1996). Klitzschophyllites shows a wide distribution, occurring in Tunisia, Egypt and Brazil (Mohr et al., 2006), while Choffatia is also found in Brazil, suggesting migration between the tropical areas of Northern Gondwana and the Iberian Peninsula. These Lower Cretaceous aquatic angiosperms were mainly constrained by water and carbon dioxide supplies and reproductive cycle.

Scenario for the angiosperm invasion of Europe
The Barremian flora (Fig. 4) consisted of matoniaceous thickets growing under warm and drought conditions. Nevertheless, conifers occupied part of the floodplain associated with possible shade-tolerant ferns (Cladophlebis) and bennettites (Zamites buchianus) and may have formed open woodland. The marshes bore a tree forest (e.g. Pinites solmsii), while angiosperms competed with charophytes in the freshwater environments (Martín-Closas and Gomez, 2004). This early angiosperm ecology fits well with the ‘Wet and Wild’ hypothesis (D. L. Dilcher, pers. com.). However, angiosperms were probably not only aquatic (hydrophytes) since at the Lowermost Aptian pollen grains referable to chloranthoids, lauraloids or magnolialoids already existed (Heimhofer et al., 2005).


Figure 4
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  		FIG. 4. Reconstruction of the vegetation changes during the Barremian–Cenomanian in Europe (blue, ferns; orange, gymnosperms excluding Pinales; purple, Pinales; green, angiosperms; B.R., braided river; F.S., freshwater swamp; F.P., floodplain; L., levee; E., estuary; S.M., salt marsh) (species numbers of pie diagrams from the Barremian to Albian are given in Fig. 3 and from the Cenomanian see in Coiffard et al., 2006).

 
The Late Aptian mild phase resulted in the closure of the vegetation by the conifers diversifying and spreading in all the environments (and so for the first time conifers grew in channel margins), whereas the matoniaceous ferns nearly became extinct (Fig. 4). Furthermore, although no simple-leafed, pinnately veined, fossil mega-remains referable to these groups were collected either from the Aptian or the Aptian–Albian of Europe, a quick diversification of the chloranthoid, lauralean or magnolialean pollen grains occurred (Heimhofer et al., 2005), and may be related to the migration of ombrophilous taxa toward upland habitats. Nevertheless, such leaves are known from the Aptian of the Potomac Group zone I usually collected in floodplain deposits and some of them (e.g. Ficophyllum) probably occupied part of the understorey (Upchurch, 1984). These co-occurrences of aquatic and understorey angiosperms during the Aptian may support the ‘Dark and Disturbed’ hypothesis. Thus, the freshwater aquatic environments should have been colonized briefly after the disturbed understorey in the early angiosperm ecological evolution.

In addition, the occurrence of Klitzschophyllites in the aquatic environments may reflect a north poleward migration from Africa and/or South America. The warm and semi-arid Albian climate coincided with the opening of the vegetation and the recovery of the matoniaceous ferns. The latter had a more restricted ecological range when compared with the Barremian, while conifers continued to dominate the channel environments (Fig. 4). From the Middle Albian, angiosperms lived in floodplains. Sender et al. (2005) described that floodplain angiosperms and Weichselia reticulata co-occurred in the Valle del Río Martín (Teruel, Spain), implying that angiosperms may have colonized the floodplains under semi-arid climate conditions before the Late Albian wet phase. These were mostly core angiosperms (in the sense of phylogeny, i.e. eudicots and eumagnoliids), and they could have been driven to a poleward migration by global warming. Angiosperms did not reach the disturbed margins of streams before the Cenomanian (Coiffard et al., 2006), and this strongly contrasts with the ‘Paleoherb’ hypothesis.


   APPENDIX 1
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 APPENDIX 1
ACKNOWLEDGEMENTS
 LITERATURE CITED
 
Data matrix
1, Rubielos de Mora (Gomez et al., 1999); 2, Uña (Gomez et al., 2001); 3, Sao Sebastiano (Teixeira, 1948); 4, Casal da Quintã (Teixeira, 1948); 5, Arnal (Teixeira, 1950); 6, Maceirinhas (Teixeira, 1950); 7, Leiria (Alvarez-Ramis and de Lorenzo, 1982); 8, Cercal (Teixeira, 1948); 9, Pola de Siero (Alvarez-Ramis and de Lorenzo, 1982); 10, Sao Mamede (Teixeira, 1948); 11, Almargen (Teixeira, 1948); 12, Belas (Teixeira, 1950); 13, Runa (Caixaria) (Alvárez-Ramis and Meléndez, 1971); 14, Buarcos, upper part (Teixeira, 1950); 15, Pimenteira (Teixeira, 1950); 16, Villadiego (Alvarez-Ramis et al., 1981); 17, Cierva (Alvarez-Ramis and Meléndez, 1971); 18, Ortigosa de Cameros (Depape and Doubinger, 1960); 19, Ohlos Amarelos (Teixeira, 1948); 20, Poussio de Galiota (Teixeira, 1948); 21, Torrinhas (Teixeira, 1950); 22, Buarcos, lower part (Teixeira, 1950); 23, Tavarede (Teixeira, 1950); 24, Beare Green (Harris, 1981); 25, Quedlinburg (Daber, 1968); 26, Smokejacks Brickworks (Ross and Cook, 1995; Hill, 1996); 27, Las Hoyas (Barale and Doludenko, 1993); 28, Montsec (Barale and Doludenko, 1993); 29, Montfaux, fossiliferous clays (Carpentier, 1927); 30, Bernissart (Carpentier, 1927); 31, Hanovre (Depape, 1965); 32, Quinta do Leirião (Teixeira, 1948); 33, Railway 66 (Teixeira, 1948); 34, Préjano (Román-Gómez, 1987); 35, Féron, lower part of the white clays (Carpentier, 1927); 36, Féron, upper part of the white clays (Carpentier, 1927); 37, Lipnik near Bielsko (Reymanowna, 1965); 38, Prznosza near Skrzydlna (Reymanowna, 1965).


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   ACKNOWLEDGEMENTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
LITERATURE CITED
 
The authors thank D. L. Dilcher, J. Kvacek and C. Martín-Closas for discussions, and J. Dinis for sedimentological and stratigraphic information on Portuguese localities. C.C. thanks A. Nel for his help in the development of the method. This article is publication number UMR5125-07.XXX, and is a contribution to Global Change IFB ‘Interactions biodiversité végétale–changements globaux à la transition Crétacé inférieur–supérieur d'Europe occidentale’. The research of B. Gomez was supported by projects ANR AMBRACE (No. BLAN07-1-184190) of the Agence Nationale de Recherches of France, CGL2005-00046/BTE and CGL2005-01121 of the Ministerio de Educación y Ciencia of the Spanish government and project 2005SGR-00890 of the Catalan government.


   LITERATURE CITED
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 APPENDIX 1
 ACKNOWLEDGEMENTS
 LITERATURE CITED
 

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