Quaternary refugia of the sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill.): an extended palynological approach

Knowledge about the glacial refugia of the thermophilous European Castanea sativa Mill. (sweet chestnut) is still inadequate. Its original range of distribution has been masked by strong human impact. Moreover, under natural conditions the species was probably admixed with other taxa (such as Quercus, Fraxinus, Fagus, Tilia) and thus possibly represented by low percentages in pollen records. In this paper we try to overcome the difficulties related to the scarcity and irregularity of chestnut pollen records by considering 1471 sites and extending the palynological approach to develop a Castanea refugium probability index (IRP), aimed at detecting possible chestnut refugia where chestnuts survived during the last glaciation. The results are in close agreement with the current literature on the refugia of other thermophilous European trees. The few divergences are most probably due to the large amount of new data integrated in this study, rather than to fundamental disagreements about data and data interpretation. The main chestnut refugia are located in the Transcaucasian region, north-western Anatolia, the hinterland of the Tyrrhenian coast from Liguria to Lazio along the Apennine range, the region around Lago di Monticchio (Monte Vulture) in southern Italy, and the Cantabrian coast on the Iberian peninsula. Despite the high likelihood of Castanea refugia in the Balkan Peninsula and north-eastern Italy (Colli Euganei, Monti Berici, Emilia-Romagna) as suggested by the IRP, additional palaeobotanical investigations are needed to assess whether these regions effectively sheltered chestnut during the last glaciation. Other regions, such as the Isère Département in France, the region across north-west Portugal and Galicia, and the hilly region along the Mediterranean coast of Syria and Lebanon were classified as areas of medium refugium probability. Our results reveal an unexpected spatial richness of potential Castanea refugia. It is likely that other European trees had similar distribution ranges during the last glaciation. It is thus conceivable that shelter zones with favourable microclimates were probably more numerous and more widely dispersed across Europe than so far assumed. In the future, more attention should be paid to pollen traces of sporadic taxa thought to have disappeared from a given area during the last glacial and post-glacial period.Electronic Supplementary Material Supplementary material is available in the online version of this article at . A link in the frame on the left on that page takes you directly to the supplementary material.An erratum to this article can be found at     

Pattern of distribution of the American bullfrog <Emphasis Type="Italic">Rana catesbeiana</Emphasis> in Europe

Despite early reports of its presence, no recent data exist on the distribution of the American bullfrog in Europe, the causes of introduction, or the trends of populations. We monitored the European situation at two spatial scales. In SW France, we performed call surveys over 2,500 wetlands. We found bullfrogs over about 2,000 km², apparently the European area in which the strongest expansion of bullfrogs is taking place. In addition, we used questionnaires to investigate the situation at the continental scale. At least 25 independent introductions occurred in Europe; eradication attempts were successful three times, and bullfrog populations are present in five countries. Education programs and monitoring are necessary to reduce the rate of introduction and to start management action as soon as possible. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

An experimental test of the evolution of increased competitive ability hypothesis in goldenrod, <Emphasis Type="Italic">Solidago gigantea</Emphasis>

The mechanisms that allow introduced plants to become invasive are poorly understood. Here, we present a test of the evolution of increased competitive ability hypothesis, which holds that because specialized natural enemies may be absent from the introduced range, exotic plants may evolve to invest less in anti-herbivore defenses and thereby gain a competitive advantage over native plants. We grew Solidago gigantea plants derived from both the native range (North America) and the invasive range (Europe) in a common garden in the native range for 2 years. Half the plants were treated with insecticide to protect them from insect herbivores and the other half were exposed to insects that colonized the garden from nearby fields. Insect herbivore biomass was significantly higher on European plants than US plants in the first year but not the second. European plants were more heavily attacked by pathogens in both years of the study. When exposed to insect herbivores, US plants produced more seed than European plants, but when plants were protected from herbivores, seed production was equivalent between US plants and European plants. The presence of insect herbivores suppressed seed production of European plants much more than that of US plants, even though the level of herbivory experienced by European and US plants was similar in the second year, suggesting that the ability to tolerate herbivory was diminished in European plants. These results partially support the EICA hypothesis: plants from the introduced range were more susceptible to some natural enemies and benefited more from insect removal than plants from the native range. The prediction that European plants would perform better than US plants in the absence of insect herbivores was not supported. Electronic Supplementary Material Supplementary material is available for this article at     

Genetic diversity of the sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill.) in Central Europe and the western part of the Balkan Peninsula and evidence of marron genotype introgression into wild populations

The sweet chestnut (Castanea sativa Mill.) is a widely spread and important multipurpose tree species in the Mediterranean area, which has played an important role in human history. Natural events, such as glaciations, and human influence played significant roles in the distribution and genetic makeup of the sweet chestnut. In order to better understand how natural and human-mediated past events affected the current genetic diversity and structure of the sweet chestnut, we analysed populations from Central Europe and the western part of the Balkan Peninsula, utilizing ten polymorphic nuclear microsatellite markers. The study revealed the existence of three genetically and, to a large extent, geographically distinct and well-defined groups of sweet chestnut populations. Two not entirely separated groups of populations were detected in the northern part of the studied area and one in the southern. Our results indicate that the genetic structure of sweet chestnut populations in Central Europe and the western part of the Balkan Peninsula is the result of both natural colonization events and significant and lengthy human impact. Furthermore, it has been proven that the gene flow between cultivated/grafted trees’ and wild chestnut stands can influence their genetic structure. However, our results reveal that cultivated-to-wild introgression in the sweet chestnut is dependent on the close proximity of chestnut orchards and naturally occurring populations.     

Role of domestication in shaping <Emphasis Type="Italic">Castanea sativa</Emphasis> genetic variation in Europe

The genetic structure of sweet chestnut (Castanea sativa Mill.) across Europe was assessed using 73 inter-simple sequence repeat markers to screen 1,768 individuals from 68 stands distributed across 29 sites in five European countries (Italy, France, Spain, Greece, and UK). At each site, trees were sampled from three distinct management types (domestication levels): naturalized stands, managed coppice, and grafted fruit orchards. In more than a third of the orchards, nonlocal genetic material (grafted clones) were evident, showing (as predicted) large differences from the other two domestication levels for most of the within-population genetic diversity parameters estimated. Randomly generated linkage disequilibrium analysis revealed weak though significant differences in two-locus allelic correlations between naturalized stands and coppice, suggesting that long-term management techniques may influence the genetic makeup of the populations. Multivariate analysis revealed the existence of five distinct gene pools across the study area; three were located in Greece, one on the northwestern coast of the Iberian peninsula and a large gene pool covering the rest of the Mediterranean basin. The implications of the results are discussed in relation to developing conservation strategies for chestnut genetic resources in Europe.     

The spread of<Emphasis Type="Italic"> Abies</Emphasis> throughout Europe since the last glacial period: combined macrofossil and pollen data

In this paper we reconstruct the location of the last glacial refugia and postglacial spread of Abies throughout Europe based on combined pollen and macrofossil data. More than 208 pollen sequences available in the European Pollen Database (EPD) and 38 macrofossil sites are used to produce distribution maps encompassing a time span between 38000 and 5500 years B.P. The investigation excludes more recent periods, because these could be strongly influenced by human impact. The pollen data presented here confirm long-lasting refugial areas such as southern Italy and Greece already described in previous studies. The combined pollen/macrofossil dataset identifies the Pyrenees as a further important refugium. In addition the pollen data indicate potential refugia in south-east France and north-west Italy. Possible migration tracks of Abies are discussed by comparing the palaeobotanical evidence with isozyme studies on gene markers of recent fir populations.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00334-004-0049-4.     

First evidence of Mespilus germanica L. (medlar) in Roman Switzerland

The Mespilus (medlar) fruit tree, non-native in Europe, is generally believed to have been introduced to central Europe during the Roman occupation of the region. Archaeobotanical remains of medlar are generally rare, resulting in a patchy knowledge of its early distribution. We here report the earliest finds of Mespilus seeds of the 2nd century a.d. in Switzerland, which were discovered in the Roman vicus of Tasgetium in Eschenz. We summarize the archaeobotanical evidence of Mespilus fruit stones in central Europe during Roman times, which indicate a wide geographical distribution of Mespilus. In addition, we give an overview of Roman sources about the use of medlar fruit and glance at medieval evidence.     

Exploitation de la vesce commune (Vicia sativa L.) au Néolithique moyen dans le Sud de la France. Données carpologiques du site de Claparouse (Lagnes, Vaucluse)

Common vetch (Vicia sativa L.) exploitation during middle Neolithic times in Southern France. Archaeobotanical data from Claparouse (Lagnes, Vaucluse). Beginnings of common vetch cultivation are poorly documented. Domestication could have occurred in various places. Common vetch is recorded by rare seeds in pre-Neolithic and Neolithic sites but firm evidence of cultivation does not seem available before Roman times. The Neolithic site of Claparouse is providing evidence of utilization and probable cultivation as early as some 6000 years ago. Even if this pulse is nowadays only a fodder plant, it was more likely used as human food at that time. We do not know whether it was introduced in western Europe from the Near East or locally domesticated. To cite this article: L. Bouby, V. Léa, C. R. Palevol 5 (2006).     

The memory of water: Archaeobotanical evidence of wetland plants from Modena (Emilia-Romagna,northern Italy) and palaeoecological remarks

Modena, founded by the Romans (183 BC), has always been conditioned by water in all its urban history. In the city, numerous archaeobotanical investigations have been carried out in order to reconstruct the natural landscape and human–environment interactions over time. During these investigations, four archaeological sites (two Roman and two medieval) have revealed deposits with a marked character of palaeobiocoenosis, largely resulting from the natural environment surrounding the sites, due to natural “seed rain”. These deposits are characterized by widespread evidence of plants related to water, constituting a valuable archive to investigate habitats which currently have become very rare and threatened, if they have not completely disappeared. The present paper aims to reveal the peculiarities of the Roman/medieval archaeocarpological floristic lists (through a comparison with the flora over the last two centuries in the area of Modena) and highlight the possible causes explaining the presence or the demise of several taxa, considering also the palaeoecological reconstruction of the environment in which they have been found.     

Landscape genetics structure of European sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill): indications for conservation priorities

Sweet chestnut is a tree of great economic (fruit and wood production), ecological, and cultural importance in Europe. A large-scale landscape genetic analysis of natural populations of sweet chestnut across Europe is applied to (1) evaluate the geographic patterns of genetic diversity, (2) identify spatial coincidences between genetic discontinuities and geographic barriers, and (3) propose certain chestnut populations as reservoirs of genetic diversity for conservation and breeding programs. Six polymorphic microsatellite markers were used for genotyping 1608 wild trees sampled in 73 European sites. The Geostatistical IDW technique (ArcGIS 9.3) was used to produce maps of genetic diversity parameters (He, Ar, PAr) and a synthetic map of the population membership (Q value) to the different gene pools. Genetic barriers were investigated using BARRIER 2.2 software and their locations were overlaid on a Digital Elevation Model (GTOPO30). The DIVA-GIS software was used to propose priority areas for conservation. High values of genetic diversity (He) and allelic richness (Ar) were observed in the central area of C. sativa’s European distribution range. The highest values of private allelic richness (PAr) were found in the eastern area. Three main gene pools and a significant genetic barrier separating the eastern from the central and western populations were identified. Areas with high priority for genetic conservation were indicated in Georgia, eastern Turkey, and Italy. Our results increase knowledge of the biogeographic history of C. sativa in Europe, indicate the geographic location of different gene pools, and identify potential priority reservoirs of genetic diversity.     

No loss of genetic divergence in Torymus sinensis following its release for Dryocosmus kuriphilus control in Europe

1. Arrival, establishment, and further dispersal of non-native natural enemies are considered essential for a successful biological control programme, while among the factors that may determine the success of such a programme, genetic diversity of the introduced population plays an important role in the establishment of a non-native species.
2. The Chinese parasitoid wasp Torymus sinensis Kamijo (Hymenoptera: Torymidae) was initially released in Europe in Italy to control biologically the Asian chestnut gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), and reduce the damage induced on sweet chestnut (Castanea sativa Miller). In the following years, T. sinensis was then released in numerous other European countries as a biological control agent of D. kuriphilus. Its presence has also been reported beyond the countries of release due to rapid natural dispersal.
3. To assess the post-release genetic diversity of D. kuriphilus, we screened T. sinensis populations from six European countries and tested the possibility of these populations suffering from frequently observed genetic effects that could threaten its successful establishment in Europe.
4. Our results exhibit that T. sinensis populations have suffered neither from the Allee effect nor from genetic bottleneck after their release and establishment in Europe, something that increases the possibility to effectively control D. kuriphilus in Europe.

     

The Evolutionary History of the Transposable Element Penelope in the Drosophila virilis Group of Species

We have used phylogenetic techniques to study the evolutionary history of the Penelope transposable element in the Drosophila virilis species group. Two divergent types of Penelope have been detected, one previously described, clade I, and a new one which we have termed clade III. The phylogeny of some copies of the Penelope clade I element was partially consistent with the species phylogeny of the D. montana subphylad, suggesting cospeciation and allowing the estimation of the evolutionary rate of Penelope. Divergence times of elements found in different species are younger than the age of the species, suggesting horizontal transfer events. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Dmitri Petrov]     

Nutrition, aspects of land use and environment in medieval times in southern Germany: plant macro-remain analysis from latrines (late 11th–13th century a.d.) at the town of Überlingen, Lake Constance

From high medieval courtyards at überlingen, Lake Constance (Bodensee), 17 archaeobotanical samples from seven latrines (11th–13th century) were analysed for plant macrofossils. They contained small amounts of cultivated plants and many well preserved seeds and fruits of wild plants. The most numerous cereal finds were of Triticum spelta followed by Secale cereale. Recorded oil plants were Papaver somniferum and Linum usitatissimum. Cultivated and gathered fruits such as various Prunus species, Pyrus communis and Malus domestica, Rubus sp. and Fragaria vesca were eaten by the people of überlingen. Only one seed of Ficus carica was found which may have been imported; no other imported plants could be recognised. Agrostemma githago and Vaccaria hispanica were prevalent weeds. The presence of the latter probably indicates the cultivation of cereals on the steeper slopes above überlingen. It is shown that in central Europe, V. hispanica became a troublesome weed only around the 12th/13th century A.D. Grassland species—mainly from more or less wet locations—and ruderal plants of nitrogen-rich soils were well represented in the samples and characterize the surrounding area. Electronic Supplementary Material Supplementary material is available in the online version of this article at     

The history of rye cultivation in Europe

During recent years finds from several prehistoric and medieval periods have thrown new light on the history of the spread of rye. It is now proven that wild rye is indigenous to Anatolia and was already domesticated there by the early Neolithic at the beginning of agriculture. Secale migrated to Central Europe as a weed among other cereals, and single grains of weed rye have been recorded there since the early Neolithic. The number of finds increased during the Bronze Age and Iron Age, and the status of rye changed from weed to crop plant, probably in the course of the early Iron Age. This acculturation of Secale cereale in central and eastern Europe was obviously independent of the earlier one in Anatolia. The first stages towards deliberate cultivation happened unintentionally through harvesting close to the ground, so that the rye was permanently represented in the seed corn. From this point rye was able to take advantage of its competitive strength on poor soils and in areas with unfavourable climate. The start of rye as a crop in its own right during the pre-Roman Iron Age and Roman period presumably took place independently in different areas. The expansion of intensive rye cultivation occurred in the Middle Ages. However, new finds from north-west Germany, which are presented here, show that in this area rye has been cultivated as a main crop on poor soils since the Roman period. In two maps all rye finds up to 1000 A.D. are shown, which after critical consideration can be regarded as cultivated rye.     

The long history of Cannabis and its cultivation by the Romans in central Italy, shown by pollen records from Lago Albano and Lago di Nemi

The cores from the Albano and Nemi lakes, near Rome, were studied within the European Union funded PALICLAS project and provided high resolution records of the Late-glacial and Holocene. Pollen evidence of increasing human influence on vegetation was recorded in the Holocene parts of both diagrams, and the Cannabis (hemp) curve was one of the major signs. In this paper we present unambiguous pollen evidence from the Cannabaceae records for the cultivation of hemp in central Italy by the Romans. The oldest records of Cannabis and Humulus (hop) date from to the Late-glacial. Hop pollen values rise during the mid Holocene, while hemp pollen becomes more abundant from ca. 3000 cal B.P. onwards. The highest earliest hemp peak (21%) is dated to the 1st century A.D. This ‘Cannabis phase’, with the abrupt rise of hemp pollen soon after the rise of cultivated trees (Castanea, Juglans and Olea) is associated with the increase in cereals and ruderal plants. This unambiguous proof of cultivation by Romans around 2000 B.P. occurs as well as a long lasting pre-Roman presence of hemp in the area, which is natural and possibly also anthropogenic. Subsequent clear episodes of cultivation in the medieval period were found. Received February 4, 2002 / Accepted September 13, 2002 Correspondence to: Anna Maria Mercuri, e-mail: mercuri.annamaria@unimo.it     

The first archaeobotanical evidence of Spinacia oleracea L. (spinach) in late 12th–mid 13th century a.d. France

Macroscopic charred remains of Spinacia oleracea L. (Amaranthaceae) have been found in the Pyrenean village of Montaillou, France, in several contexts of a house dated to the end 12th–mid 13th century a.d. This is the first archaeobotanical record of this vegetable in France and the earliest European archaeobotanical material so far found. The paper presents the morphological criteria used for identifying the charred remains of the species. After a review of archaeobotanical finds in Europe, hypotheses on the economic status of this vegetable, which is unknown as a wild plant in Europe, are discussed with reference to medieval written and illuminated sources and to archaeological deposits. It appears that Spinacia was first introduced into France from Moorish Spain where it was cultivated at least since the 11th century. The French evidence of Spinacia thus represents a milestone in the history and geographic diffusion of this vegetable into temperate Europe.     

A failed invader in the North Atlantic, the case of Aglenus brunneus Gyll. (Col., Colydiidae), a blind flightless beetle from Iceland

The occurrence of an occasional pest of mouldy stored product residues, the blind flightless beetle Aglenus brunneus Gyll., in samples from the medieval farm at Reykholt in Iceland, along with several other strongly synanthropic beetles, is considered in relation to its fossil record. The species is dependent on man for its dispersal and survival and it probably had its primary habitat in the warm, decaying litter of the undisturbed forest floor in Europe. Now virtually cosmopolitan, it had been introduced to a remote site in the eastern desert of Egypt by the Roman period and was widespread in medieval northern Europe. The processes by which such an apparently stenotopic species could have invaded are discussed in relation to other evidence for anthropochorous dispersal.     

Archaeozoological records and distribution history of the osprey (Pandion haliaetus) in Central Europe

A review of the Holocene records of the osprey (Pandion haliaetus) in Central Europe indicates an immigration around 10,000 years BP at the latest. During the middle Holocene—9,000–5,000 years BP—the species seems to have been common in the south-western Baltic region and perhaps also in Switzerland, but scarce in the rest of Central Europe. After a complete lack of osprey remains for nearly 4,000 years up to medieval times, a few records again suggest a sporadic distribution of this species in the south-western Baltic. In early modern times, the osprey recolonised north-eastern Germany and, during the nineteenth and the first decades of the twentieth century, bred in colony-like densities. Because of relentless persecution and, later, contamination with pesticides, osprey populations collapsed. Recently, the Central European populations have been slowly increasing again. The significance of the subfossil findings is discussed and possible reasons for their scarcity are outlined.     

Chloroplast DNA polymorphism reveals little geographical structure in Castanea sativa Mill. (Fagaceae) throughout southern European countries

The distribution of haplotypic diversity of 38 European chestnut (Castanea sativa Mill.) populations was investigated by PCR/RFLP analysis of regions of the chloroplast and mitochondrial genomes in order to shed light on the history of this heavily managed species. The rapid expansion of chestnut starting from 3000 years ago is strongly related to human activities such as agricultural practice. This demonstrates the importance of human impact, which lasted some thousands of years, on the present-day distribution of the species. No polymorphism was detected for the single mitochondrial analysed region, while a total of 11 different chloroplast (cp) haplotypes were scored. The distribution of the cpDNA haplotypes revealed low geographical structure of the genetic diversity. The value of population subdivision, as measured by GSTc, is strikingly lower than in the other species of the family Fagaceae investigated. The actual distribution of haplotypic diversity may be explained by the strong human impact on this species, particularly during the Roman civilization of the continent, and to the long period of cultivation experienced during the last thousand years.     

Ectomycorrhizal characterization of an American chestnut <Emphasis Type="SmallCaps">(</Emphasis><Emphasis Type="Italic">Castanea dentata</Emphasis><Emphasis Type="SmallCaps">)</Emphasis>-dominated community in Western Wisconsin

Circa 1900, a farmer from the eastern US planted 11 American chestnut (Castanea dentata) seeds on a newly established farm near West Salem in western Wisconsin. These trees were very successful, producing a large stand of over 6,000 trees. Since this area is well outside the natural range of chestnut, these trees remained free from chestnut blight until 1987. In the West Salem stand, chestnuts are the dominant species of a mixed forest community, reminiscent of the chestnut–oak ecosystems of pre-1900 Appalachia. To identify putative mycorrhizal associates of chestnut in this unique forest, our approach was twofold: (1) an extensive fruiting body survey was conducted for four seasons that yielded approximately 100 putative mycorrhizal species and (2) a belowground molecular approach was used to generate DNA sequences of the internal transcribed spacer region from ectomycorrhizae. Unexpectedly, chestnut did not appear to be the dominant underground ectomycorrhizal-forming plant species. This study highlights the need to identify the plant host species when conducting belowground molecular-based surveys and provides preliminary identification of ectomycorrhizal fungi associated with a disjunct stand of American chestnut. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.