Palaeobotany

This volume covers the Prehistory of Ukraine from the Lower Palaeolithic through to the end of the Neolithic periods. This is the first comprehensive synthesis of Ukrainian Prehistory from earliest times through until the Neolithic Period undertaken by researchers who are currently investigating the Prehistory of Ukraine. At present there are no other English language books on this subject that provide a current synthesis for these periods. The chapters in this volume provide up-to-date overviews of all aspects of prehistoric culture development in Ukraine and present details of the key sites and finds for the periods studied. The book includes the most recent research from all areas of prehistory up to the Neolithic period, and, in addition, areas such as recent radiocarbon dating and its implications for culture chronology are considered; as is a consideration of aDNA and the new insights into culture history this area of research affords; alongside recent macrofossil studies of plant use, and anthropological and stable isotope studies of diet, which all combine to allow greater insights into the nature of human subsistence and cultural developments across the Palaeolithic to Neolithic periods in Ukraine. It is anticipated that this book will be an invaluable resource for students of prehistory throughout Europe in providing an English-language text that is written by researchers who are active in their respective fields and who possess an intimate knowledge of Ukrainian prehistory.

Malcolm C. Lillie (PhD 1998, University of Sheffield) is Professor of Archaeology at Umeå University, Sweden. Previously he was Professor of Prehistoric Archaeology and Wetland Science at the University of Hull, England. Malcolm undertook MSc and PhD research at Sheffield University, the latter under the supervision of the late Professor Marek Zvelebil. His main research interests are the Prehistory of Ukraine, in situ preservation in wetlands, prehistoric and wetlands archaeology in general and the analysis of prehistoric human remains.

Inna Potekhina (PhD 1992, Institute of Archaeology of the National Academy of Sciences of Ukraine, Kiev) is Head of the Bioarchaeology Department in the Institute of Archeology, National Academy of Sciences of Ukraine, Kiev and an Associate Professor at the University of 'Kyiv-Mohyla Academy', where she teaches a course of anthropology and bioarchaeology. Inna is an author of more than 150 articles and four monographs on the anthropology of the prehistoric populations of Ukraine and Eastern Europe, the genomic history of Mesolithic-Eneolithic Ukraine and palaeodemography and palaeodiet reconstructions.

Chelsea Budd (PhD 2016, University of Oxford) is a Postdoctoral Researcher in Archaeology at the University of Umeå, Sweden. Her main research interests are Early European Prehistory, isotope geochemistry and the application of statistical modelling techniques to archaeological research.

Extrafloral nectaries (EFNs) are commonly found in Passiflora L. Reports have been made on the occurrence of resin-producing structures morphologically similar to EFNs in the genus. The objective of this study was to characterise the morphoanatomy and development of leaf secretory structures in Passiflora amethystina and to use chemical and histochemical tests to detect the presence of sugars in the exudates. Samples of leaf blade and petioles in different developmental stages were collected and subjected to usual techniques using light and scanning electron microscopy. Secretion samples were analysed by high performance liquid chromatography. The concentration of total sugars in the secretion amounted to 39.67% for blade EFNs and 52.82% for petiolar EFNs. EFNs consist of a secretory, uni-or bistratified palisade epidermis, arising from the protoderm by means of anticlinal and periclinal divisions, glandular parenchyma originated from the ground meristem, and xylem and phloem elements formed from the procambium. Exudate accumulated in a subcuticular space formed outside the epidermal cells from where it was then released. Histochemical tests showed a positive reaction for neutral polysaccharides. The results confirm that the leaf secretory structures are indeed extrafloral nectaries, and these findings constitute important information for studies on the taxonomy and ecology of this species.

In the austral summer of 2006/7 the ANDRILL MIS (ANtarctic geological DRILLing-McMurdo Ice Shelf) project
recovered a 1285 m sediment core from beneath the Ross Ice Shelf near Ross Island, Antarctica in a flexural
moat associated with volcanic loading. The upper ~600 m of this core contain sediments recording 38 glacial/
interglacial cycles of Early Pliocene to Pleistocene time, including 13 discrete diatomite units (DUs). The
longest of these, DU XI, is ~76 m-thick, and has been assigned an Early to Mid-Pliocene age (5–3 Ma). A
detailed record of the siliceous microfossil assemblages in DU XI is used in conjunction with geochemical and
sedimentological data to subdivide DU XI into four discrete subunits of continuous sedimentation. Within
each subunit, changes in diatom assemblages have been correlated with the δ18O record, providing a temporal
resolution up to 600 yr, and allowing for the construction of a detailed age model and calculation of associated
sediment accumulation rates within DU XI. Results indicate a productivity-dominated sedimentary record
with greater proportions of hemipelagic mud accumulating during relatively cool periods. This implies that
even during periods of substantial warmth, Milankovitch-paced changes in Antarctic ice volume can be linked
to ecological changes recorded in diatom assemblages.

In the austral summer of 2006/7 the ANDRILL MIS (ANtarctic geological DRILLing-McMurdo Ice Shelf) project
recovered a 1285 m sediment core from beneath the Ross Ice Shelf near Ross Island, Antarctica in a flexural
moat associated with volcanic loading. The upper ~600 m of this core contain sediments recording 38 glacial/
interglacial cycles of Early Pliocene to Pleistocene time, including 13 discrete diatomite units (DUs). The
longest of these, DU XI, is ~76 m-thick, and has been assigned an Early to Mid-Pliocene age (5–3 Ma). A
detailed record of the siliceous microfossil assemblages in DU XI is used in conjunction with geochemical and
sedimentological data to subdivide DU XI into four discrete subunits of continuous sedimentation. Within
each subunit, changes in diatom assemblages have been correlated with the δ18O record, providing a temporal
resolution up to 600 yr, and allowing for the construction of a detailed age model and calculation of associated
sediment accumulation rates within DU XI. Results indicate a productivity-dominated sedimentary record
with greater proportions of hemipelagic mud accumulating during relatively cool periods. This implies that
even during periods of substantial warmth, Milankovitch-paced changes in Antarctic ice volume can be linked
to ecological changes recorded in diatom assemblages.

Biogeographic histories cannot solely rely on cladistic reconstructions. First, a cladogram may be an insufficient representation of evolution, in particular at low taxonomic levels (here: intrageneric evolution). Diversification and speciation in plants is complex, rarely resulting in a single unambiguous tree-like genealogy. Second, the modern distribution of a monophyletic group (here: genus Fagus) may be highly incomplete. For instance, important areas such as Central Asia and Iceland no longer include the taxon. Third, incorporating fossil taxa, including potential ancestors of modern taxa, into a cladistic phylogeny may be intrinsically difficult. All three issues apply to the Northern Hemispheric genus Fagus, the beech trees. In this study, we use the exceptionally complete fossil record of Fagus to trace its spatiotemporal unfolding throughout the Cainozoic. Fossil taxa are placed in a phylogenetic framework following Darwinian classification by identifying common ancestries and overall patterns of similarity. These lines of evidence are then combined to put up a holistic biogeographic history of Fagus in the Cainozoic, which is discussed in the light of general patterns of morphological and molecular differentiation and diversification. Based on the fossil record, Fagus evolved in the Northern Pacific region. Earliest fossils are known both from Northeast Asia and north-western North America (possibly extending to Axel Heiberg Island) in the Middle Eocene. The range expansion of Fagus into Central Asia and Europe during the Oligocene was accompanied by a first differentiation into a continental Eurasian and northern Pacific lineage. In the Miocene, Fagus is found throughout the Northern Hemisphere up to very high latitudes (e.g., Iceland). By the end of the Miocene several lineages had been established, including putative ancestors of modern taxa (e.g. F. evenensis, the ancestor of subgenus Engleriana). Overall, the Miocene taxa are characterized by conspicuous morphological plasticity reflecting potentially unhindered gene flow. In post-Miocene times, Fagus underwent range reduction and extinction in high latitudes, Central Asia and western North America. The disruption of a former homogeneous area led to the shaping of modern species. The methodological approaches used here for Fagus take into account the particular nature of data sets palaeobotanists are usually confronted with.

Although Cretaceous fossils (coal excluded) from Victoria, Australia, were first reported in the 1850s, it was not until the 1950s that detailed studies of these fossils were undertaken. Numerous fossil localities have been identified in Victoria since the 1960s, including the Koonwarra Fossil Bed (Strzelecki Group) near Leongatha, the Dinosaur Cove and Eric the Red West sites (Otway Group) at Cape Otway, and the Flat Rocks site
(Strzelecki Group) near Cape Paterson. Systematic exploration over the past five decades has resulted in the collection of thousands of fossils representing various plants, invertebrates and vertebrates. Some of the best-preserved and most diverse Hauterivian–Barremian floral assemblages in Australia derive from outcrops of the lower Strzelecki Group in the Gippsland Basin. The slightly younger Koonwarra Fossil Bed (Aptian) is a Konservat-Lagerstätte that also preserves abundant plants, including one of the oldest known flowers. In addition, insects, crustaceans (including the only syncaridans known from Australia between the Triassic and the present), arachnids (including Australia’s only known opilione), the stratigraphically youngest xiphosurans from Australia, bryozoans, unionoid molluscs and a rich assemblage of actinopterygian fish are known from the Koonwarra Fossil Bed. The oldest known—and only Mesozoic—fossil feathers from the Australian continent constitute the only evidence for tetrapods at Koonwarra. By contrast, the Barremian–Aptian-aged deposits at the Flat Rocks site, and the Aptian–Albian-aged strata at the Dinosaur Cove and Eric the Red West sites, are all dominated by tetrapod fossils, with actinopterygians and dipnoans relatively rare. Small ornithopod (=basal neornithischian) dinosaurs are numerically common, known from four partial skeletons and a multitude of isolated bones. Aquatic meiolaniform turtles constitute another prominent faunal element, represented by numerous isolated bones and articulated carapaces and plastrons. More
than 50 specimens—mostly lower jaws—evince a high diversity of mammals, including monotremes, a multituberculate and several enigmatic ausktribosphenids. Relatively minor components of these fossil assemblages are diverse theropods (including birds), rare ankylosaurs and ceratopsians, pterosaurs, non-marine plesiosaurs and a lepidosaur. In the older strata of the upper Strzelecki Group, temnospondyl amphibians—the youngest known worldwide—are a conspicuous component of the fauna, whereas crocodylomorphs appear to be present only in up-sequence deposits of the Otway Group. Invertebrates are uncommon, although decapod crustaceans and unionoid bivalves have been described. Collectively, the Early Cretaceous biota of Victoria provides insights into a unique Mesozoic high-latitude palaeoenvironment and elucidates both palaeoclimatic and palaeobiogeographic changes throughout more than 25 million years of geological time.

Charred botanical finds from the excavation of the Early Bronze Age city of Jericho (Tell es-Sultan),
one of the earliest urban centers of 3rd millennium BC Palestine, were collected during the 2015-2017
excavation seasons carried out by Sapienza University of Rome and the Palestinian MoTA-DACH.
Among other plant macro-remains, a round fruit was found in the subsidiary room behind the throne
room of Royal Palace G, next to a vase, in the burnt filling overlying the platform. It was identified as
a drupe of a dwarf palm, through classical archaeobotanical techniques and computed tomography
scan. Two dwarf palms were taken into consideration: the Mediterranean dwarf palm (Chamaerops
humilis L.) and the Mazari palm (Nannorrhops ritchiana (Griff.) Aitch. native to the Saharo-Indian
region), both with small, round/oval fruits, none of which currently grows in the area of Jericho.
A detailed analysis of iconography, archaeobotanical literature and herbarium samples of both species
stored in Rome (RO), Florence (FIAF) and Edinburgh (E), has allowed to identify the charred drupe as
Nannorrhops ritchiana. Its presence in the palace suggests the existence of an overland commercial
track to the south-east, across the desert of Saudi Arabia, which only recent excavations and other
finds have revealed.

The  present  volume,  fourth  in  a  series  of  five  devoted  to  the
Merovingian  hamlet  of  Develier-Courtételle  (Canton  of  Jura,
Switzerland),  presents  environmental  information  obtained
through a series of specialised analyses of material found at this
archaeological site and in its surroundings. The results of these six
distinct approaches are integrated in a final synthesis.
A description of the natural environment of the site enriched with
information from historical sources opens the volume (chap. 1).
Situated within the Delémont basin in north-western Switzerland,
in the central part of the Jura mountain range, Develier-Courtételle
lies  on  a  flood  plain  at  an  altitude  of  450 m.  The  site  covers  a
surface of 3,5 ha and lies on the banks of the brook “La Pran”, in
a lateral valley filled with quaternary sediments dating from the
Würm glacial maximum to the Late Holocene. The sedimentary
approach which follows this introductory chapter is based on a
description of the stratigraphy (chap. 2). The site is divided into
eight different domains and composite stratigraphic cuts are used
to  explain  the  extension  of  the  different  layers.  The  sediments
are  characterised  using  Passega’s  method,  i.e.  the  construction
of  C/M  patterns  of  the  alluvial  deposits.  This  analysis  permits
us  to  propose  a  reconstruction  of  the  palaeogeography  of  the
site, including some observations on the nature of the soils. The
palaeo-hydrological  and  climatic  fluctuations  occurring  locally
between 500 and 800 AD are also described. Additional relevant
data  are  provided  by  an  “off-site”  sedimentological  analysis
undertaken at Delémont, La Communance (chap. 3). It provides
a palustrine sequence dating from the 6th to the 11th century and
documenting the evolution of the local hydrology.
The  three  following  chapters  present  the  archaeobotanical
approaches. The pollen analysis documents the changing landscape
around  the  site  and  the  use  of  the  land  by  the  Early  Medieval
population (chap. 4). The study of seeds and fruit reveals the large
spectrum of plant species that were cultivated during this period
(chap.  5).  The  vegetational  landscape  surrounding  the  site  and
the human impact on it can be reconstructed from the remains of
wild plants. A geobotanical approach is used to deduce the spatial
distribution of woodlands and fields around the site during the
7th century by looking at modern vegetation (chap. 6). The last
analysis presented in this volume is the study of the faunal remains
recovered in and around the settlement which reveal the dietary
choices made by the inhabitants (chap. 7). The varying frequency
and age at death of the different species of domesticates say much
about animal husbandry during the Merovingian period. The rare
remains of hunted wild animals were also identified.
The concluding synthesis is divided into three thematic sections
(chap. 8). The climatic context and the evolution of the landscape
during  the  Early  Medieval  period  are  the  subject  of  the  first
section.  A  short  summary  of  current  knowledge  introduces
the  chronological  reconstitution  of  events  deduced  from  the
preceding analyses. At Develier-Courtételle, hydrological activity
is quite sustained before 550. Forest cover is still relatively dense
and actually expanded during Late Antiquity. From 550 to 750,
the hydrological activity at the site becomes very intense, peaking
around 580 and 650. At La Communance, the hydrological balance
is at this time in surplus. From the beginning of this phase, forest
cover diminishes ; this clearance was caused by the intensification
of  agriculture  and  the  appearance  of  ironworking.  As  a  result,
erosion  cuts  ravines  into  the  slopes.  Several  factors  indicate  a
cool  and  possibly  wet  phase  affecting  the  whole  region  which
correlates well with data from other studies. The period between
700 and 830-890 appears to be a transition towards a somewhat
dryer climate, a decrease of hydrological activity persisting after
800  could  be  observed  on  the  site.  At  La  Communance,  the
hydrological balance becomes negative only after the middle or
towards the end of the 9th century. Forest cover does not vary to
any major extent. The climate continues to improve between 830-890 and 1000 : several swamps and marshes in the region appear
to have experienced a negative hydrological balance during this
time and superficial sediments undergo pedogenesis. The rupture
of the landscape equilibrium occurring during the later Middle
Ages  is  thought  to  be  due  to  the  increasing  impact  of  human
activity, in particular ironworking.
The second theme of the synthesis is the landscape surrounding
the settlement during the 7th century. A pictorial reconstruction
allows us to conclude that the divisions of the territory are largely
based  on  the  meanders  of  the  stream.  To  the  inhabitants,  the
advantages of being close to the water necessary for various craft
activities  clearly  outweighed  the  real  risks  posed  by  flooding.
The  marshy  surfaces  directly  abutting  on  the  settlement  were
good  pasture  but  not  useful  for  agriculture ;  the  fields  were
situated somewhat further away. The transition from cultivated
land  to  forest  occurred  gradually  and  gave  the  landscape  a
structured appearance. The forest, dominated by beech and pine,
is  transformed  by  the  selective  exploitation  of  plant  resources.
It too is used for pasture.
A  discussion  of  available  resources  and  land  use  closes  the
volume. The stream could not have been used for transporting
cargo. Used land surfaces included fields, pastures and grasslands.
Cereals and legumes were the primary crops, oily or fibrous plants
were of secondary importance and fruit and vegetables appear to
have formed only a relatively minor source of food. Supplying the
needs of the settlement seems to have been the principal goal of
these activities, a characteristic of subsistence agriculture. Textile
production  occurred  but  was  not  very  extensive.  The  forests
and hedges are a source of timber, but particularly of firewood.
Among the domestic animals, cattle are dominant and seem to
have mostly been raised for meat, a part of which may have been
exported. Goats and especially sheep are the next most important
group according to the bone counts. The pigs are in third position
but were actually in second place in terms of meat production.
Fowl,  mostly  chicken,  were  principally  kept  for  the  eggs.  The
diverse wild fauna was nutritionally a very minor factor but is a
good indicator of the natural environment surrounding the site.
The presence of deer and bear speaks of nearby biotopes not yet
disturbed by human activity.

The excursion will touch some of the sections where to collect plant fossil remains of Carboniferous, Permian and Jurassic deposits. To gather a fossil plant collection from the Miocene lacustrine sections is today very difficult because the artificial lake for water supply, such as the Omodeo Lake, has submerged the “Fossil Forest of Zuri” with its silicified trunks 12-15 m high and more than a meter in diameter and, furthermore, the stratigraphy of fill Miocene lacustrine basins is at present, for the most, hidden by the development of intensively cultivated fields. New findings of fossil plants are actually retrieved when canals for water distribution have been excavated in the interested areas. The “Foresta Fossile” of the Perfugas-Martin basin, and the “Foresta Fossile of Zuri” that are protected by a severe regional and national laws are, ironically, self-endangered.

A new stomatal proxy-based record of CO2 concentrations ([CO2]), based on Betula nana (dwarf birch) leaves from the Hässeldala Port sedimentary sequence in south-eastern Sweden, is presented. The record is of high chronological resolution and spans most of Greenland Interstadial 1 (GI-1a to 1c, Allerød pollen zone), Greenland Stadial 1 (GS-1, Younger Dryas pollen zone) and the very beginning of the Holocene (Preboreal pollen zone). The record clearly demonstrates that i) [CO2] were significantly higher than usually reported for the Last Termination and ii) the overall pattern of CO2 evolution through the studied time period is fairly dynamic, with significant abrupt fluctuations in [CO2] when the climate moved from interstadial to stadial state and vice versa. A new loss-on-ignition chemical record (used here as a proxy for temperature) lends independent support to the Hässeldala Port [CO2] record. The large-amplitude fluctuations around the climate change transitions may indicate unstable climates and that “tipping-point” situations were involved in Last Termination climate evolution. The scenario presented here is in contrast to [CO2] records reconstructed from air bubbles trapped in ice, which indicate lower concentrations and a gradual, linear increase of [CO2] through time. The prevalent explanation for the main climate forcer during the Last Termination being ocean circulation patterns needs to re-examined, and a larger role for atmospheric [CO2] considered.► A stomatal proxy-based CO2 record from the Last Termination is presented. ► The stomata based CO2 record is much more dynamic than ice core-based CO2 records. ► CO2 first increases abruptly before decreasing at cooling transitions and vice versa.

This is the first of a series of three papers synthesising the results of an interdisciplinary study of the chain of small forts built c. AD 40 on the southern bank of the Rhine between Vechten (prov. Utrecht / NL ) and the North Sea coast. The project focused on the reasons for the establishment of these military installations and on the efforts required to build and maintain them. These questions were addressed by a team of researchers from diverse backgrounds: archaeology, geomorphology, palaeobotany and zooarchaeology.
Whereas the project was formally restricted to the period c. AD 40-140 this first paper discusses the preceding era: the transition from the Late Iron Age into the early principate. The events from AD 40 onwards cannot be properly assessed without a basic understanding of what preceded: the pre-Roman landscape and habitation of the lower Rhine delta, Rome’s conquest of Gaul and its struggles with the Germanic peoples from the right bank of the Rhine.
The narrative is embedded in the historical sources and therefore essentially chronological, but there are several digressions on the landscape and its use, with the Rhine and its delta as a prominent element, and on military supply, which is of vital importance in a period of conquest. It appears that the political rivalry of the late Roman Republic was the key factor in the expansion to the Northwest, but once set off there was no way back. Each conquest induced new threats demanding further actions, as far as the natural landscape and the military supply network allowed.
Eventually the Roman troops advanced to the Elbe, and there remains little doubt that Augustus intended to turn most of the annexed territory between the Rhine and Elbe into a Roman province. Although this goal came within reach once Tiberius subjected all Germanic peoples between Rhine and Elbe in 8 BC , its full achievement was time and again frustrated. It is generally assumed that the abortion of the Roman military campaigns after AD 16 marks the definite abandonment of the imperial dream of a German provincial territory east of the Rhine, but the continuing tributary status of the transrhenane Frisii and the potential of the area for military supply – including cereals, meat, leather, metal ore, stone and army recruits – shed doubt on that conclusion.
At the death of emperor Tiberius in AD 37 Germania was not necessarily considered lost, and it may not be a coincidence that his successor Caligula crossed the Rhine when he needed a quick military success to secure his grip on the imperial throne.

Although progress has been made in recent years in reconstructing the environmental conditions at the Triassic/Jurassic Boundary (TJB), published records of atmospheric CO2 concentrations have been of low resolution and/or based on multi-taxon estimates. This is addressed here by reconstructing CO2 concentrations across the TJB using stomatal frequencies of four phylogenetically and ecologically distinct plant groups from two depositionally, geographically and taphonomically separate boundary sections in East Greenland and Northern Ireland, with stomatal proxy methods and regression analysis. The resulting CO2 records then are compared with an additional existing TJB record from a geological section in Sweden. The final results indicate that pre-TJB (Rhaetian), the CO2 concentration was approximately 1000 ppm, that it started to rise steeply pre-boundary and had doubled to around 2000–2500 ppm at the TJB. The CO2 concentration then remained elevated for some time post-boundary, before returning to pre-TJB levels in the Hettangian. These results are in very good accordance with published C-isotope, fire and leaf dissection records, and clearly indicate steeply rising and lingering CO2 concentration at the TJB.