Late persistence and deterministic extinction of “humid thermophilous plant taxa of East Asian affinity” (HUTEA) in southern Europe
Introduction
In the course of the stratigraphical and palaeontological study of the San Lazzaro section in central Italy (Fig. 1), recently assigned to the early Pleistocene (Baldanza et al., 2014), one of us (A.B.) found an endocarp of Sinomenium cantalense. The finding of this species, readily assignable to the humid thermophilous taxa of East Asian affinity, in an early Pleistocene section was the starting point for further collecting efforts to find evidence for the role of central Italy as a centre of refuge for such thermophilous taxa in the Plio–Pleistocene (Martinetto, 2001a). In this paper we adopt the definition of the Pliocene and Pleistocene of Gibbard et al. (2010), with the boundary fixed at 2.6 Ma, and we accept their indication for the chronologic boundaries of the four stages Zanclean, Piacenzian, Gelasian and Calabrian. Therefore, the terms middle Pliocene, late Pliocene and early Pleistocene used in previous works (among others, Ambrosetti et al., 1995a, Ambrosetti et al., 1995b, Abbazzi et al., 1997, Martinetto, 2001a) have a chronologic connotation which differs from that adopted here.
It is well known that many plant fossils found in the late Cenozoic of Europe belong to thermophilous genera or infrageneric taxa which do not grow in this continent today (Mai, 1989, Qian et al., 2006, Rodríguez-Sánchez and Arroyo, 2008). Such fossils are usually called “exotic elements” (Reid, 1920) and this concept corresponds more or less with “extinct plants” for the Plio–Pleistocene interval (Svenning, 2003). The climatic requirements are not considered in the definition of both exotic and extinct; however, several attempts have been made to assign the exotic (or extinct) elements to a few distinct plant groups that involve a climatic characterization and/or a phytogeographic aspect (Mai, 1989, Mai, 1991, Mai, 1995a, Grichuk, 1997, Grímsson et al., 2015). Examples of names which have been used include: “Palaeotropical flora/element”, “Arcto-Tertiary” or “Arctotertiary flora/element” (Engler, 1879–1882, Mai, 1989, Mai, 1991, Grímsson et al., 2015), “subtropical elements” (Mai, 1970, Zagwjin, 1990), “Mastixioideen” (Kirchheimer, 1957, Mai, 1964), “Boreotropical flora” (Wolfe, 1975), “Taxodiaceae group” (Bertoldi et al., 1994), “Tethyan plants (or Tethys flora)” (Szafer, 1961, Mai, 1989, Rodríguez-Sánchez and Arroyo, 2008), “Mega-mesothermic elements” (e.g. Popescu et al., 2010), and “humid subtropical elements” (Bertini and Martinetto, 2011). All these names leave some uncertainty as to what is included and what is excluded from the definition, firstly because the phytogeographic information, both past and present, is superimposed to, and variously interfingers with, the climatic one, and secondly because of the very difficult, not unambiguous, climatic characterization of the fossil-taxa (Kvaček, 2007, Grimm and Denk, 2012, Utescher et al., 2014). Also the modern reference models may be ambiguous, for example the qualitative term “subtropical” is used with very different temperature boundaries by Chinese (e.g. Hou, 1983) and Japanese authors (e.g. Kira, 1991).
The different extant distribution of plant taxa that grew together in the Cenozoic of Europe have often been given considerable relevance in the analysis of palaeofloras (see Reid and Reid, 1915, Szafer, 1961, Mai, 1964, Mai, 1989, Mai, 1995a). However, in our opinion most previous analyses and descriptions of the floral change in the Plio–Pleistocene of Europe suffered from the lack of precisely defined categories whose chronological analysis would adequately point out timing and entity of the large Plio–Pleistocene mass extinction (Tallis, 1991, Svenning, 2003). Additionally, the descriptions of Plio–Pleistocene floral changes mostly relied on pollen data (e.g. Tzedakis et al., 2006, Postigo-Mijarra et al., 2009, Magri, 2010, Orain et al., 2013), particularly in Italy (Bertini, 2010, Combourieu-Nebout et al., 2015). However, by combining pollen and carpological records (Bertini and Martinetto, 2011) it was noticed that pollen assemblages mainly reflect anemophilous plants, whilst they do not accurately represent the assemblages of “subtropical humid forest” type (sensu Hou, 1983, and Bertini and Martinetto, 2011), which are very rich in entomophilous plants and were present in southern Europe right at the time when major extinction events are hypothesized (Martinetto, 2001a, Bertini, 2010, Bertini and Martinetto, 2011, Biltekin et al., 2015, Combourieu-Nebout et al., 2015). As recently confirmed by Goring et al. (2013), taxa that are pollinated by insect or animal vectors (entomophilous or zoophilous, respectively), and species with limited dispersal ability are rarely recorded in fossil pollen records. Some works on modern fruit and seed assemblages (e.g. Thomasson, 1991, Vassio and Martinetto, 2012 and references therein) indicate a less biassed representation of plant diversity, in particular for several enthomophilous (e.g. Actinidia, Frangula, Paulownia, Rubus, Sambucus) and herbaceous plants (e.g. Ajuga, Cyperaceae, Hypericum, and Potamogeton). For these taxa, the plant elements that enter the fossil record and allow species-level identification are fruits and/or seeds. Thus, the works which exclude carpological data definitely underestimate past plant species diversity and the extent of Plio–Pleistocene plant extinctions, and the focus of this paper will be on fossil fruits and seeds.
The analysis of the San Lazzaro material led us to reconsider the bulk of information accumulated for the Italian late Cenozoic fruit and seed assemblages in the last 30 years (in particular: Gregor, 1990, Martinetto, 1994, Martinetto, 1995, Martinetto, 1999, Martinetto, 2001a, Martinetto, 2001b, Martinetto, 2009, Martinetto, in press, Bertoldi and Martinetto, 1995, Mai, 1995b, Basilici et al., 1997, Fischer and Butzmann, 2000, Ghiotto, 2010, Martinetto et al., 2007, Martinetto et al., 2015). Consequently, we felt the need to introduce precisely defined categories, which would permit us to better appreciate the chronological steps of the dramatic southern European floral change in the Plio–Pleistocene. One of the necessary operations was to combine in a clear manner the modern phytogeography and the climatic requirements of several taxa. Therefore, we focused on geographical and ecological characteristics of modern relatives of fossil taxa: partly shared geographic range, minimum thermic tolerance and moisture requirement. Since the geographic area where most of the “exotic” taxa of the European late Cenozoic are still living today is definitely East Asia (Tralau, 1963, Martinetto, 1998, Qian et al., 2006, Manchester et al., 2009), we considered it to be important for the definition of the new categories.
The taxonomic similarity between Neogene European floras and modern East Asian ones is rooted at least into the Miocene (Mai, 1989). As known from various studies at global and regional scales, Cenozoic climates were generally warmer and more humid than at present, and were characterized by shallow latitudinal gradients (Utescher et al., 2011 and references therein). Several authors (Bruch et al., 2011, Liu et al., 2011, Xing et al., 2012, Jacques et al., 2013) pointed out that the climate was wetter and warmer than the present one during the Miocene in both central Europe and China. Even central and northern Eurasian areas, such as Kazakhstan (Bruch and Zhilin, 2006) and Siberia (Popova et al., 2012), were wetter and warmer during the Miocene, despite the relatively higher seasonality and continentality.
This climatic situation was probably suitable for the formation of latitudinal vegetation belts with a similar floristic composition in both western and eastern Eurasia (Mai, 1989, Mai, 1991, Kovar-Eder et al., 2008), and strong floristic affinities with East Asia have also been encountered for North American floras (Liu and Jacques, 2010, Huang et al., 2014). Several authors explained that the modern East Asian woody flora is richer than the European (and North American) one (e.g., Kubitzki and Krutzsch, 1996, Manchester, 1999, Wen, 1999, Tiffney and Manchester, 2001, Wen et al., 2010) mainly due to a minor impact of extinctions, even if several woody species got extinct also in East Asia during the Plio–Pleistocene (Momohara, 2015).
Some close relatives of most European extinct species were already present in the warm temperate belt of East Asia before the Pliocene (e.g., Cathaya, Cephalotaxus, Craigia, Cryptomeria, Cyclocarya, Eucommia, Ginkgo, Glyptostrobus, Pseudolarix, Taiwania: Manchester et al., 2009) or possibly migrated there during the Pliocene (e.g., Hemiptelea, Rehderodendron: Manchester et al., 2009), and could survive the Pleistocene climatic crisis because of the presence of niches that were wet (atmospheric humidity) and warm enough, even in sites not related to rivers and swamps. Based on the concept of “physiological uniformitarianism” (Tiffney and Manchester, 2001) we can assume that the climatic tolerances of the living relatives of Neogene European taxa that survived in the humid and warm temperate to tropical areas of East Asia roughly correspond (maybe only in part) to those of the extinct European forms of the same genus, subgenus or section.
Svenning (2003) pointed out a deterministic effect in late Cenozoic plant extinctions and recognized three important groups of taxa for the analysis of the ancient European floras: 1) widespread taxa; 2) relictual taxa; 3) extinct taxa. In referring to extinct taxa, Svenning (2003) restricted his analysis to cool-temperate tree genera, but recently Eiserhardt et al. (2015) carried out an analysis on more thermophilous plants. Actually, several Plio–Pleistocene taxa occurring in Europe are more thermophilous than “cool-temperate” (Martinetto et al., 2015) so that we now consider it important to single out a new group of thermophilous taxa with a partly shared (as for eastern Asia) current distribution outside Europe and a common, definite climatic boundary. The thermophilous characterization of several European fossil-species is provided by the minimum Mean Annual Temperature (MAT) requirement of their modern relatives (Table 1).
Consequently, we define as “HUmid Thermophilous extinct European taxa of East Asian affinity”, in short HUTEA, those plant taxa which have well-documented fossil records in the late Cenozoic of Europe, which do not grow spontaneously in this continent and West Asia at present (unless as aliens), which do not tolerate a MAT below 8 °C and a Mean Annual Precipitation (MAP) below ca. 800–1000 mm/year, and which belong to genera or infrageneric taxa that presently grow in East Asia (Wang, 1961, Qian et al., 2006, Fang et al., 2009, Fang et al., 2011, Manchester et al., 2009, Grimm and Denk, 2012, Eiserhardt et al., 2015, Utescher and Mosbrugger, 2015).
We single out the 8 °C value of MAT because this is the boundary of the distribution of boreal (subarctic) and thermophilous (temperate) taxa in East Asia. The upper limit of fir and spruce forest and the lower limit of deciduous forest is 7.8 °C MAT in China (Fang and Yoda, 1989). Although the lower MAT limit of the thermophilous evergreen arboreal Fagaceae and Lauraceae (dominant tall trees of temperate broadleaved evergreen forests in East Asia) is between 9 and 12 °C (Hattori and Nakanishi, 1985, Fang and Yoda, 1989, Fang et al., 2011), we decided that adding 1 °C of tolerance would admit sporadic occurrences of thermophilous plants above the 9 °C MAT isotherm.
The focus on MAT for the definition of the HUTEA is justified by the large availability of data (Grimm and Denk, 2012, Utescher and Mosbrugger, 2015) for most of the plant genera documented by fossils in Europe, and by the determinant role of this parameter for plant extinction or survival in the late Cenozoic of Europe (Svenning, 2003, Eiserhardt et al., 2015). Conversely, we did not manage to gather precise values of minimum precipitation requirements for all the exotic Neogene plant taxa of Europe; nevertheless we consider important to include in the HUTEA definition a rule that excludes those plants which tolerate a low precipitation (below ca. 800–1000 mm/year). In fact it has been pointed out that the extinction of several Neogene taxa in Europe depended from a scarce tolerance not only of low temperature, but also of low precipitation (Svenning, 2003, Eiserhardt et al., 2015). The thermophilous genera that survived in southern Europe until the present time (e.g. Laurus, Olea) are mainly adapted to dry (Mediterranean) climate, whereas several thermophilous genera extinct in Europe are now growing in areas affected by the East Asian Monsoon that supplies higher precipitation to plants during the growing season. In East Asia the main evergreen forest formation, dominated by Fagaceae and Lauraceae, is called “lucidophyllous forest” and differs from the south European (Mediterranean) sclerophyllous forest formation by its less xeromorphic characteristics, such as larger shiny leaves, larger tree size and higher species diversity with many epiphytes and woody lianas (Kira, 1991).
We are aware that other parameters (e.g. Warmth Index, Coldness Index; Kira, 1991) could be more appropriate to define a category such as HUTEA. Nevertheless, the minimum MAT requirement is an important factor determining the possibility for a plant taxon to overcome a climatic bottleneck. The climatic characteristics of the refugia might have been decisive for the possibility of a plant species to survive (Magri, 2010, Gavin et al., 2014) and obviously it would have gone extinct if its minimal thermal or humidity requirements would no longer have been present in the last refugium. In this respect, groups of taxa with similar requirements may be expected to go extinct together (Tallis, 1991, Grichuk, 1997, Eiserhardt et al., 2015), given that the MATmin of the refugia created a bottleneck for all of the plants growing there. However, some extinctions have certainly been controlled by complex and multiple factors. For example it has been suggested that Cedrus (Su et al., 2013) and Sequoia (Zhang et al., 2015) disappeared from China because of seed ecological aspects, triggered by climate change.
Three examples, concerning genera which do not tolerate a MAT below 8 °C (Table 1), may be useful to support the above definition of HUTEA: Toddalia is assigned to the HUTEA because it is distributed in the tropical–warm temperate zone of East Asia and in Africa, but not in Europe and West Asia (Gregor, 1979). Symplocos sect. Lodhra is assigned to the HUTEA because it is distributed in the tropical–warm temperate zone of East Asia, but not in Europe and West Asia (Fritsch et al., 2015). Rehderodendron is assigned to the HUTEA because it is distributed only in the “subtropical” zone (sensu Hou, 1983) of East Asia.
The genera Cathaya and Pseudolarix meet all the requirements to be classified as HUTEA, but they are excluded for their present highly relictual distribution, which may provide an inaccurate representation of their past climatic requirements, similarly as for Tetraclinis (Kvaček, 2007). Even if the distribution of Amentotaxus is similarly relictual, we temporarily decided to consider it a HUTEA. Azolla is not considered a HUTEA because it is a water plant rather independent from atmospheric humidity.
According to the above definition and to the data (e.g., minimum thermic requirements: MATmin) reported in Table 1, the following HUTEA have so far been documented for the late Cenozoic of Italy (Martinetto, 1995, Martinetto, 1998, Martinetto, 1999, Martinetto, 2001a, Martinetto, 2001b, Follieri, 2010, Martinetto et al., 2015): Amentotaxus, Cinnamomum, Craigia, Cyclea, Cyclocarya, Ehretia, Engelhardia, Eucommia, Glyptostrobus, Mallotus, Meliosma subgen. Kingsboroughia, Paulownia, Rehderodendron, Sabia, Sargentodoxa, Saurauia, Sinomenium, Stemona, Symplocos sect. Lodhra, Taiwania, Ternstroemia, Tetrastigma, Toddalia, Trichosanthes, Turpinia and Wikstroemia.
Other taxa documented in the late Cenozoic of southern Europe have the correct geographic distribution nowadays to be considered HUTEA (i.e. embracing East Asia and excluding Europe and West Asia), but they are not considered, because the modern representatives do tolerate a MAT below 8 °C (e.g. Actinidia, Alangium, and Ampelopsis: Table 1). These taxa will be named CTEA (“Cool-Tolerant extinct European taxa of East Asian affinity”) in this paper and belong to the somehow ambiguous [changing on the basis of the extent of territory considered] category of the “exotic” taxa (Reid, 1920; see the more precisely defined “category E” in Martinetto, in press).
The HUTEA category already has a satisfactory climatic connotation, which we deem to be useful for an analysis of the climatic determinism in their extinction. Conversely, the CTEA category certainly contains a very heterogeneous mix of species with different climatic tolerances. In fact, Magnolia provides a good example of a cool-tolerant CTEA genus that contains several modern species (Grimm and Denk, 2012, Utescher and Mosbrugger, 2015), which are absolutely thermophilous and not cool-tolerant (tropical–subtropical). Similarly, a diversified climate tolerance has been also hypothesized for different European fossil-species (Mai, 1975). Given this situation, it is unsurprising that several CTEA would show a HUTEA-like extinction pattern. However, in this paper our attention has been focused on the species that show a delayed disappearance time in comparison to the HUTEA.
Finally, a few taxa which do not tolerate a MAT below 8 °C are not assigned to the HUTEA because of the modern geographic range: Coriaria, Datisca, Ficus, Laurus, Liquidambar, Morella, Ocotea, Olea, Platanus, Sideroxylon, Styrax, Tetraclinis and Visnea grow in southern Europe, North Africa (incl. Macaronesia) and/or West Asia. These taxa will be indicated as TEWA, Thermophilous European, West Asian and/or African elements, in this paper. Pterocarya, Parrotia and Zelkova are not assigned to the HUTEA nor to the CTEA or TEWA, because they grow in relict niches of south-eastern Europe and/or West Asia (southern shores of the Black Sea and Caspian Sea), commonly including sites with a MAT below 8 °C.
Finally, late Cenozoic south European taxa that today only survive in America are not many (Decodon, Dulichium, Leitneria, Proserpinaca, Sequoia, Taxodium) and will not be specifically dealt with in this paper.
It is apparent that the HUTEA and CTEA concepts have much to do with a change of geographic distribution between the Plio–Pleistocene and the present. The main aim of this work is to present new fossil data from Italy and an updated state-of-the-art regarding the timing of disappearance of HUTEA and CTEA species from Europe. Furthermore, we newly consider the possibility of deterministic extinctions (Svenning, 2003, Eiserhardt et al., 2015).
Section snippets
Geological setting
The post-Miocene, NW–SE oriented South Valdichiana Basin (Fig. 1), enclosed between the Meso-Cenozoic Rapolano-Mt. Cetona and Narnese-Amerina Apennine anticlines and bounded by extensional faults, occupies a wide area between south-eastern Tuscany and western Umbria, in central Italy. In the Pliocene–Pleistocene interval, the Narnese-Amerina ridge separated the mainly marine domain of South Valdichiana from the continental deposits of the Southern Tiberino Basin (Fig. 1, Fig. 2), whilst, during
Materials and methods
This work integrates the analysis of freshly collected material from the San Lazzaro section with the reinterpretation of the stratigraphic and palaeontological data from the sites Cava Toppetti II (Abbazzi et al., 1997, Argenti, 1999, Argenti, 2004, Martinetto, 2001a, Petronio et al., 2003, Sardella et al., 2003) and Dunarobba (Ciangherotti et al., 1998, Manganelli and Giusti, 2000, Manganelli et al., 2008, Martinetto et al., 2014).
In the San Lazzaro section, as well as in the neighbouring
The San Lazzaro section and its age constraints
The composite sedimentological and stratigraphic reconstruction proposed for the Fabro Scalo area integrates old observations (Baldanza et al., 2011, Baldanza et al., 2014, Bizzarri et al., 2015) and newly collected data. The general geological and sedimentological pattern, from the base to the top, is organized as follows (Fig. 3):
- –
about 10 m (cropping out) of structureless, mollusc-rich clayey and silty sand (offshore transition deposits); the lowermost layers are covered by recent alluvial
Discussion
Depending on the concepts of “physiological uniformitarianism” (Tiffney and Manchester, 2001) and deterministic late Cenozoic plant extinctions in Europe (Svenning, 2003) we singled out three groups of plant fossil-species occurring in southern Europe: CTEA, HUTEA, TEWA. The CTEA and HUTEA include several species extinct in Europe and belonging to supraspecific plant taxa with a modern distribution in East Asia. The HUTEA are the descendants of the “exuberant laurophyllous flora” (Kubitzki and
Conclusions
New data on early Pleistocene fossil fruit and seed assemblages from Italy allowed us to detect several extinct taxa that commonly went unnoticed in pollen analyses. The combined analysis of Pliocene and early Pleistocene occurrence data provided a detailed picture of the reduction of plant diversity in southern Europe. The possible explanation of the causes of plant extinction requires an excursion into deeper times: Several Neogene plants were mainly adapted to grow in thermophilous mesic
Acknowledgements
We thank G. Basilici for useful information and discussions about the stratigraphy of the Tiberino Basin and for help provided in the field for the positioning of the carpological sample within his Cava Toppetti II section. Thanks to A. Bertini for very useful information on pollen assemblages and for suggestions that improved the whole manuscript, which also profited from the valuable revision carried out by two anonymous referees that we wish to thank. We also thank A. Bruch, T. Denk, G.
References (190)
Sedimentary facies in an extensional and deep-lacustrine depositional system: the Pliocene Tiberino Basin, Central Italy
Sediment. Geol.
(1997)- et al.
Evidence of late Gelasian dispersal of African fauna at Coste San Giacomo (Anagni Basin, central Italy): Early Pleistocene environments and the background of early human occupation in Europe
Quat. Sci. Rev.
(2014) Pliocene to Pleistocene palynoflora and vegetation in Italy: state of the art
Quat. Int.
(2010)- et al.
Reconstruction of vegetation transects for the Messinian/Piacenzian of Italy by means of comparative analysis of pollen, leaf and carpological records
Palaeogeogr. Palaeoclimatol. Palaeoecol.
(2011) - et al.
Reevesia and Itea in the pollen flora of the Upper Neogene continental deposit at Sarzana (lower Magra valley, Northern Italy)
Rev. Palaeobot. Palynol.
(1994) - et al.
Anatolia: a long-time plant refuge area documented by pollen records over the last 23 million years
Rev. Palaeobot. Palynol.
(2015) - et al.
Precipitation patterns in the Miocene of Central Europe and the development of continentality
Palaeogeogr. Palaeoclimatol. Palaeoecol.
(2011) - et al.
Acinonyx pardinensis (Carnivora, Felidae) from the Early Pleistocene of Pantalla (Italy): predatory behavior and ecological role of the giant Plio–Pleistocene cheetah
Quat. Sci. Rev.
(2014) - et al.
The remarkable Middle Pliocene non-marine mollusc record from Ceresole d'Alba, Piedmont, north-west Italy: biochronology, palaeobiogeography and palaeoecology supported by fossil plants
Geobios
(2007) - et al.
The biogeographic history of beech trees
Rev. Palaeobot. Palynol.
(2009)
Vegetational patterns and distribution of relict taxa in humid temperate forests and wetlands of Georgia (Transcaucasia)
Biol. J. Linn. Soc. London
Conifer extinction in Quaternary Italian records
Quat. Int.
Systematics, biostratigraphy and paleoecology of the genus Toddalia Jussieu (Rutaceae) in the European Tertiary
Rev. Palaeobot. Palynol.
Contribution to the Late Neogene and Early Quaternary floral history of the Mediterranean
Rev. Palaeobot. Palynol.
Reliability and resolution of the coexistence approach — a revalidation using modern-day data
Rev. Palaeobot. Palynol.
Italian Cenozoic crocodilians: taxa, timing and biogeographic implications
Palaeogeogr. Palaeoclimatol. Palaeoecol.
Do extant nearest relatives of thermophile European Tertiary elements reliably reflect climatic signal?
Palaeogeogr. Palaeoclimatol. Palaeoecol.
Stable isotope record in mollusca and pedogenic carbonate from Late Pliocene soils of Central Italy
Palaeogeogr. Palaeoclimatol. Palaeoecol.
Sinomenium macrocarpum sp. nov. (Menispermaceae) from the Miocene–Pliocene transition of Gray, northeast Tennessee, USA
Rev. Palaeobot. Palynol.
The evolution of Miocene climates in North China: preliminary results of quantitative reconstructions from plant fossil records
Palaeogeogr. Palaeoclimatol. Palaeoecol.
Persistence of tree taxa in Europe and Quaternary climate changes
Quat. Int.
Paleontological and sedimentological record in Pliocene distal alluvial fan deposits at Cava Toppetti (Todi, Central Italy)
Boll. Soc. Paleontol. Ital.
Il Pleistocene inferiore nel ramo sud occidentale del bacino tiberino (Umbria): aspetti litostratigrafici e biostratigrafici
Il Quaternario
La foresta fossile di Dunarobba (Terni, Umbria, Italia centrale): contesto litostratigrafico, sedimentologico, palinologico, dendrocronologico e paleomalacologico
Il Quaternario
Molecular phylogeny of Juglans (Juglandaceae): a biogeographic perspective
Tree Genet. Genomes
La biocronologia dei Roditori del Plio–Pleistocene dell'Umbria e l'evoluzione del genere Apodemus (Muridae, Rodentia) in Italia)
Plio–Quaternary mammal fossiliferous sites of Umbria (Central Italy)
Geol. Romana
Eine Frühdiluviale Flora im Mainzer Becken
Z. Bot.
Gelasian to Calabrian onland marine record: three case studies in the Mediterranean area. Proceedings of AIQUA Congress: Il Quaternario Italiano — Conoscenze e prospettive (Rome, February 24–25, 2011)
Il Quaternario
Lungo la costa del Mar Tirreno…due milioni di anni fa
The early Pleistocene shallow marine decapod crustaceans community from Fabro Scalo (western Umbria, central Italy): taxonomic inferences and palaeoenvironmental reconstruction
Neues Jb. Geol. Paläontol. Abh.
Sedimentologia della parte distale di un conoide alluvionale del Pliocene superiore (Bacino Tiberino, Umbria)
Il Quaternario
Pliocene lacustrine deposits of the Tiberino Basin (Umbria, central Italy)
Floodplain lake deposits on an early Pleistocene alluvial plain (Tiberino Basin, Central Italy)
Paleoenvironmental evolution in the Pliocene marine–coastal succession of Val Chiusella (Ivrea, NW Italy)
Boll. Soc. Paleontol. Ital.
Climate and vegetation in the Upper Valdarno basin (central Italy) as a response to Northern Hemisphere insolation forcing and regional tectonics in the late Pliocene–early Pleistocene
Ital. J. Geosci.
Ricerche paleobotaniche (palinologiche e paleocarpologiche) sulla successione “villafranchiana” del Rio Ca' Viettone
Il Quaternario
The rediscovered Hula painted frog is a living fossil
Nat. Commun.
On the meaning of the Amphistegina levels in the Plio–Pleistocene of the Orvieto area (Central Italy)
Plio–Pleistocene deltaic deposits in the Città della Pieve area (western Umbria, central Italy): facies analysis and inferred relations with the South Chiana Valley fluvial deposits
Il Quaternario
The latest continental filling of Valle Umbra (Tiber Basin, central Italy) dated to one million years ago by magnetostratigraphy
Il Quaternario
Lunulite bryozoans from Early Pleistocene deposits of SW Umbria (Italy): sedimentological and paleoecological inferences
Facies
Techniques
Early Miocene climate of Central Eurasia — Evidence from Aquitanian floras of Kazakhstan
Palaeogeogr. Palaeoclimatol. Palaeoecol.
Comparative skull osteology of Karsenia koreana (Amphibia, Caudata, Plethodontidae)
J. Morphol.
Late Pliocene palaeoenvironment and correlation of the Vildstejin floristic complex within Central Europe
Rozpr Cesk. Akad. Ved.
Flore carpologiche del Pliocene di Castelletto Cervo (Biella)
Boll. Mus. Reg. Sci. Nat. Torino
Canis etruscus (Canidae, Mammalia) and its role in the faunal assemblage from Pantalla (Perugia, central Italy): comparison with the Late Villafranchian large carnivore guild of Italy
Boll. Soc. Paleontol. Ital.
New well-preserved material of Lynx issiodorensis valdarnensis (Felidae, Mammalia) from the Early Pleistocene of Pantalla (central Italy)
Boll. Soc. Paleontol. Ital.
Re-defining Canis etruscus (Canidae, Mammalia): a new look into the evolutionary history of Early Pleistocene dogs resulting from the outstanding fossil record from Pantalla (Italy)
J. Mamm. Evol.
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