Eemian Interglacial Period Research Articles

Whole genomes from the extinct Xerces Blue butterfly can help identify declining insect species

The Xerces Blue (Glaucopsyche xerces) is considered to be the first butterfly to become extinct in historical times. It was notable for its chalky lavender wings with conspicuous white spots on the ventral wings. The last individuals were collected in their restricted habitat, in the dunes near the Presidio military base in San Francisco, in 1941. We sequenced the genomes of four 80- to 100-year-old Xerces Blue, and seven historical and one modern specimens of its closest relative, the Silvery Blue (Glaucopsyche lygdamus). We compared these to a novel annotated genome of the Green-Underside Blue (Glaucopsyche alexis). Phylogenetic relationships inferred from complete mitochondrial genomes indicate that Xerces Blue was a distinct species that diverged from the Silvery Blue lineage at least 850,000 years ago. Using nuclear genomes, both species experienced population growth during the Eemian interglacial period, but the Xerces Blue decreased to a very low effective population size subsequently, a trend opposite to that observed in the Silvery Blue. Runs of homozygosity and deleterious load in the former were significantly greater than in the later, suggesting a higher incidence of inbreeding. These signals of population decline observed in Xerces Blue could be used to identify and monitor other insects threatened by human activities, whose extinction patterns are still not well known.

Whole genomes from the extinct Xerces Blue butterfly can help identify declining insect species.

The Xerces Blue (Glaucopsyche xerces) is considered to be the first butterfly to become extinct in historical times. It was notable for its chalky lavender wings with conspicuous white spots on the ventral wings. The last individuals were collected in their restricted habitat, in the dunes near the Presidio military base in San Francisco, in 1941. We sequenced the genomes of four 80- to 100-year-old Xerces Blue, and seven historical and one modern specimens of its closest relative, the Silvery Blue (Glaucopsyche lygdamus). We compared these to a novel annotated genome of the Green-Underside Blue (Glaucopsyche alexis). Phylogenetic relationships inferred from complete mitochondrial genomes indicate that Xerces Blue was a distinct species that diverged from the Silvery Blue lineage at least 850,000 years ago. Using nuclear genomes, both species experienced population growth during the Eemian interglacial period, but the Xerces Blue decreased to a very low effective population size subsequently, a trend opposite to that observed in the Silvery Blue. Runs of homozygosity and deleterious load in the former were significantly greater than in the later, suggesting a higher incidence of inbreeding. These signals of population decline observed in Xerces Blue could be used to identify and monitor other insects threatened by human activities, whose extinction patterns are still not well known.

QUATERNARY DEPOSITS AND ECOLOGICAL MONITORING OF THE ICE AGE PARK IN CARPATHIAN REGION, UKRAINE

An international geopark "The Park of Ice Age" located near the Starunia village in Bohorodchany district of Ivano-Frankivsk region is world-famous for being a paleontological site of Pleistocene fauna of the woolly rhinoceroses and mammoth (the Eemian interglacial period- 46-10 000 years ago according to radiocarbon dating C14) and for its mud volcano which is unique in the Carpathian region, This is a geological nature monument with an area of 60 hectares where abandoned ozokerite mine and oil and gas exploratory boreholes are presented. The detailed investigations carried out in the years 2004 - 2009 by Ukrainian-Polish expeditions were focused on discoveries of possible new sites of fauna fossils and human remains of European early modern humans (EEMH). The future discoveries at the paleontological site of Starunia are extremely important. They intend to carry out ecological assessment of soil, surface and stream waters, atmospheric air, snow, ashes of meadow grasses, radiological screening, and geo-radar sensing to find new locations of extinct mammals. The Starunia area meets all the requirements of a geopark according to UNESCO standards and concepts. The first research findings (1907) are connected with the remains of woolly rhinoceros, mammoth, horse, roe deer and other Pleistocene mammals which had been found in the ozokerite mine at the depth of 12 m near Starunia (Bohorodchany district, Ivano-Frankivsk region). In 1914 the scientists from Lviv (Ukraine) and Krakow (Poland) fully appreciated these discoveries and published a set of articles and a monograph. In 1929 the members of the expedition of Skill Academy (Krakow, Poland) found the remains of three more woolly rhinoceroses in the ozokerite mine at the depth of 17 m. Numerous bones of vertebrates (rodents), remains of mussels, a lot of species of insects, beetles, parasitic worms, fleas, butterflies, spiders, snails, vascular plants, mosses, seeds and branches of dwarf birch (Betula Nana), alder (Alnus glutinosa), and other fossil of tundra flora were also discovered. Polish scientists organized detailed investigations of flora and fauna around Starunia, their results were published in articles but the World War II suspended this process. The investigations around Starunia intensified the comprehensive study of stratigraphy, paleontology, paleogeography, geochronology and other aspects of the Quaternary Period. In 1932 there was established the International Geological Congress (the International Union of Quaternary Study) by INQUA in Leningrad. The main methodological principals of this study dealt with the investigations around Starunia.

Demographic History of the Brown Bear (Ursus arctos) on Hokkaido Island, Japan, Based on Whole-Genomic Sequence Analysis

Previous studies of the brown bear (Ursus arctos) on Hokkaido Island, Japan, have detected three geographically distinct subpopulations representing different mitochondrial lineages and shown that gene flow between subpopulations has occurred due to male-biased dispersal. In this study, we determined whole-genomic sequences for six Hokkaido brown bears and analyzed these data along with previously published genomic sequences of 17 brown bears from other parts of the world. We found that the Hokkaido population is genetically distinct from the other populations, keeping genetic diversity higher than the endangered populations in western Europe but lower than most populations on the continents. A reconstruction of historical demography showed no increase in population size for the Hokkaido population during the Eemian interglacial period (130,000–114,000 years ago). In a phylogenetic analysis of the autosomal data, the Hokkaido population formed a clade distinct from North American and European populations, showing that it has maintained genetic diversity independently from continental populations following geographical isolation on the island. This autosomal genetic similarity contrasts with the geographically separate mitochondrial lineages on Hokkaido and indicates the occurrence of male-driven gene flow between subpopulations.

Prospects for new sites of wooly rhinoceros and mammoths on the territory of paleontological site in Starunia, Carpathian region

An international geopark “Park of the Ice Age” located near the Starunia village in Bohorodchany district of Ivano-Frankivsk region is world-famous for being a paleontological site of Pleistocene fauna of the woolly rhinoceroses and mammoth (the Eemian interglacial period, 46–10 000 years ago according to radiocarbon dating C14) and for its mud volcano which is unique in the Carpathian region. This is a geological nature monument with an area of 60 hectares where abandoned ozokerite mine and oil and gas exploratory boreholes are presented. The detailed investigations carried out in the years 2004–2009 by Ukrainian-Polish expeditions were focused on discoveries of possible new sites of fauna fossils and human remains of European early modem humans (EEMH).The future discoveries at the paleontological site of Starunia are extremely important. They intend to carry out ecological assessment of soil, surface and stream waters, atmospheric air, snow, ashes of meadow grasses, radiological screening, and geo-radar sensing to find new locations of extinct mammals. The Starunia area meets all the requirements of a geopark according to UNESCO standards and concepts. The first research findings (1907) are connected with the remains of woolly rhinoceros, mammoth, horse, roe deer and other Pleistocene mammals which had been found in the ozokerite mine at the depth of 12 m near Starunia (Bohorodchany district, Ivano-Frankivsk region). In 1914 the scientists from Lviv (Ukraine) and Krakow (Poland) fully appreciated these discoveries and published a set of articles and a monograph. In 1929 the members of the expedition of Skill Academy (Krakow, Poland) found the remains of three more woolly rhinoceroses in the ozokerite mine at the depth of 17 m. Numerous bones of vertebrates (rodents), remains of mussels, a lot of species of insects, beetles, parasitic worms, fleas, butterflies, spiders, snails, vascular plants, mosses, seeds and branches of dwarf birch (Betula Nana), alder (Alnus glutinosa), and other fossil of tundra flora were also discovered. Polish scientists organized detailed investigations of flora and fauna around Starunia, their results were published in articles but the World War II suspended this process. The investigations around Starunia intensified the comprehensive study of stratigraphy, paleontology, paleogeography, geochronology and other aspects of the Quaternary Period. In 1932 there was established the International Geological Congress (the International Union of Quaternary Study) by INQUA in Leningrad. The main methodological principals of this study dealt with the investigations around Starunia.

Konserwacja szkieletu słonia leśnego Palaeoloxodon antiquus (80–100 tys. lat) ze zbiorów Muzeum Okręgowego w Koninie – badania i praktyka konserwatorska

The article presents the exploratory and conservation-wise problematic aspects of the bones obtained from the skeleton of a forest elephant Palaeoloxodon antiquus (Falconer & Cautley, 1847), that lived during the Eemian Interglacial period 80–100000 years ago. This preserved skeleton, currently in the collection of District Museum in Konin, is particularly valuable due to being one of the largest skeletons in Europe that remained almost complete. The necessity of undertaking conservation processes has been caused by the poor state of its preservation – partly because of degradation of the materials applied during conservation in the 80s of the 20th century and later on, as well as numerous bone tissue damages; there also emerged a possibility of using materials and treatments improving value of the object, aesthetically and exhibition-wise. The publication presents the history of the skeleton and its conservation, the condition, the outline of its bone structure, and the results of instrumental research (ATR-FTIR) of derivative materials composition. Moreover, it shows the assumptions for and course of conservation and restoration works. The performed procedures allowed to stabilize the skeleton, reduce formation of new cracks and cavities, and to restore didactic and exhibition value, enabling the skeleton to be returned to a permanent museum exhibition. The works were twice awarded the 1st prize in the category – conservation and protection of cultural heritage in the prestigious competition Museum Event of the Year “Izabella 2018” and “Izabella 2019” organized by the Wielkopolska Museum Foundation under the patronage of the Marshal of the Wielkopolska Province.

Whole-Genomes From the Extinct Xerces Blue Butterfly Reveal Low Diversity and Long-Term Population Decline

The Xerces Blue (Glaucopsyche xerces) is considered the first known butterfly to become globally extinct in historical times and was notable for its iridescent blue colour with white spots in the ventral wings. The last specimens were collected in their restricted habitat, in the dunes near the Presidio military base in San Francisco, in 1941. To explore the phylogenetic relationships and demographic history of this iconic butterfly, we have sequenced at medium coverage four 80 to 100-year-old Xerces Blue and seven historic Silvery Blue (G. lygdamus) specimens, its closest living relative. We have subsequently compared their genomes to a novel assembled and annotated genome of the Green-Underside Blue (G. alexis). The phylogenetic relationships assessed with complete mitochondrial (mtDNA) genomes indicate that Xerces Blue diverged from the Silvery Blue lineage at least 850,000 years ago. We used the nuclear genomes to explore their past demographic history and found that while both species experienced a population growth during the Eemian interglacial period, Xerces Blue decreased to a very low effective population size in post glacial periods, a trend opposite to that observed in the Silvery Blue. The patterns of runs of homozygosity in the Xerces Blue reveal signs of background inbreeding in this species to a much greater extent than the Silvery Blue, while global heterozygosity shows an opposite trend. In addition, Xerces Blue carried a higher proportion of derived, putatively deleterious amino acid-changing alleles than the Silvery Blue. We conclude that Xerces Blue experienced a long population decline process prior to its final extinction.

Tectonism and volcanism enhanced by deglaciation events in southern Iceland

AbstractSouthern Iceland is one of the main outlets of the Icelandic ice sheet and is subject to seismicity of both tectonic and volcanic origins along the South Iceland Seismic Zone (SISZ). A sedimentary complex spanning Marine Isotopic Stage 6 (MIS 6) to the present includes evidence of both activities. It includes a continuous sedimentary record since the Eemian interglacial period, controlled by a rapid deglaciation, followed by two marine glacioisostasy-forced transgressions, separated by a regression phase connected to an intra-MIS 5e glacial advance. This record has been constrained by tephrostratigraphy and dating. Analysis of this record has provided better insights into the interconnectedness of hydrology and volcanic and tectonic activity during deglaciations and glaciations. Low-intensity earthquakes recurrently affected the water-laid sedimentation during the early stages of unloading, accompanying rifting events, dyke injection, and fault reactivations. During full interglacial periods, earthquakes were significantly less frequent but of higher magnitude along the SISZ, due to stress accumulation, favored by low groundwater levels and more limited magma production. Occurrence of volcanism and seismicity in Iceland is commonly related to rifting events. Subglacial volcanic events seem moreover to have been related to stress unlocking related to limited or full unloading/deglaciation events. Major eruptions were mostly located at the melting margin of the ice sheet.

Makhnevskaya Ledyanaya Cave (Middle Urals, Russia): Biostratigraphical reconstruction

Pollen and fossil mammals of the terrigenous deposits in the Makhnevskaya Ledyanaya Cave (59°26′ N 57°41’ E), located in the northern part of the Middle Urals and on the border with the Northern Urals, were studied. The spore-and-pollen spectra are divided into four pollen zones. In all the spectra, pollen concentrations of tree species amount to 50–70%. The presence of pollen in thermophilic (Quercus robur-type, Tilia cordata-type, Ulmus sp., Carpinus betulus, Corylus avellana-type) and moderate boreal (Abies, Picea, Pinus, Betula, Alnus) species and a lack of any tundra flora elements indicate their interglacial nature. The following have been identified as small mammal fauna: 8 taxa of insectivorous mammals, 4 taxa of bats, 2 taxa of lagomorphs, 4 species of carnivores, 16 species of rodents, 1 species of odd-toed ungulates and 4 species of even-toed ungulates. The remains of cryoxerophilous (Dicrostonyx sp., Lasiopodomys gregalis) and steppe (Ochotona sp.) species are scarce in the studied fauna. Species typical of forests (genus Sorex, genus Craseomys, genus Myodes, Myopus schisticolor, Microtus agrestis) are dominant. Remains of Erinaceus sp., Talpa sp., Crocidura leucodon, Sciurus vulgaris, Hystrix brachyura, Dryomys nitedula, Apodemus flavicollis, Stephanorhinus kirchbergensis have been found. The morphology and dimensions of the teeth of Panthera spelaea fossilis, Hystrix brachyura, Dryomys nitedula, Arvicola amphibius, mice of genus Apodemus are described. The compositions of the palynocomplexes and mammal fauna as well as the SDQ values for m1 of Arvicola amphibius allow us to date the deposits from the Makhnevskaya Ledyanaya Cave to the Eemian interglacial period (MIS 5e).

Eemian Greenland ice sheet simulated with a higher-order model shows strong sensitivity to surface mass balance forcing

Abstract. The Greenland ice sheet contributes increasingly to global sea level rise. Its history during past warm intervals is a valuable reference for future sea level projections. We present ice sheet simulations for the Eemian interglacial period (∼130 000 to 115 000 years ago), a period with warmer-than-present summer climate over Greenland. The evolution of the Eemian Greenland ice sheet is simulated with a 3-D higher-order ice sheet model, forced with a surface mass balance derived from regional climate simulations. Sensitivity experiments with various surface mass balances, basal friction, and ice flow approximations are discussed. The surface mass balance forcing is identified as the controlling factor setting the minimum in Eemian ice volume, emphasizing the importance of a reliable surface mass balance model. Furthermore, the results indicate that the surface mass balance forcing is more important than the representation of ice flow for simulating the large-scale ice sheet evolution. This implies that modeling of the future contribution of the Greenland ice sheet to sea level rise highly depends on an accurate surface mass balance.

Past, present, and future geographic range of an oro-Mediterranean Tertiary relict: The juniperus drupacea case study

Juniperus drupacea is a unique relict species found in the mountains of southern Greece, southern Turkey, and western Syria and Lebanon. The aim of this study was to describe the natural range of this juniper by determining the current locations of its populations and to predict a theoretical range for the species based on current, past, and future climatic conditions. We used data from the literature, herbarium materials, and our unpublished field notes (about 500 georeferenced points in total) to determine the current natural distribution of J. drupacea (realized niche). To predict suitable conditions with the program MaxEnt, we used data from the WordClim database, which allowed estimation of the potential niche. The potential niche of J. drupacea was much wider during the Last Glacial Maximum (LGM) and was severely restricted during the Eemian interglacial period. Depending on the climate scenario, this species could become endangered in the future due to climate changes. Considering the relatively restricted geographic range of J. drupacea and the decreasing numbers of localities where it is found, conservation strategies should be adopted to allow for preservation of its genetic and morphological diversity.

Eemian Greenland SMB strongly sensitive to model choice

Abstract. Understanding the behavior of the Greenland ice sheet in a warmer climate, and particularly its surface mass balance (SMB), is important for assessing Greenland's potential contribution to future sea level rise. The Eemian interglacial period, the most recent warmer-than-present period in Earth's history approximately 125 000 years ago, provides an analogue for a warm summer climate over Greenland. The Eemian is characterized by a positive Northern Hemisphere summer insolation anomaly, which complicates Eemian SMB calculations based on positive degree day estimates. In this study, we use Eemian global and regional climate simulations in combination with three types of SMB models – a simple positive degree day, an intermediate complexity, and a full surface energy balance model – to evaluate the importance of regional climate and model complexity for estimates of Greenland's SMB. We find that all SMB models perform well under the relatively cool pre-industrial and late Eemian. For the relatively warm early Eemian, the differences between SMB models are large, which is associated with whether insolation is included in the respective models. For all simulated time slices, there is a systematic difference between globally and regionally forced SMB models, due to the different representation of the regional climate over Greenland. We conclude that both the resolution of the simulated climate as well as the method used to estimate the SMB are important for an accurate simulation of Greenland's SMB. Whether model resolution or the SMB method is most important depends on the climate state and in particular the prevailing insolation pattern. We suggest that future Eemian climate model intercomparison studies should include SMB estimates and a scheme to capture SMB uncertainties.

UK′37 temperature estimates from Eemian marine sediments in the southern coast of Hainan Island, tropical China

This study concerns high-resolution sea surface temperature (SST) changes for the Eemian interglacial period reconstructed from the southern coast of Hainan Island, China. The lower marine unit in the sediment core is composed of monotonous mud (31m thick), dated to be from the Eemian interglacial period or lower part of marine isotope stage 5 (MIS 5), determined by optically stimulated luminescence (OSL). The high sedimentation rate and the richness of organic matter and marine fossils provide good opportunity for the study of palaeo-environments in the northern margin of the South China Sea (SCS). Sediment grain-size, foraminifer, sediment color index and long-chain alkenone were used to reveal the near-shore sedimentary environments and temperature at millennial-scale. Specifically, a high-resolution UK′37-SST record was reconstructed for the Eemian interglacial period. The new evidence in northwestern margin of SCS shows that the maximum value of SST during the Eemian interglacial is approximately 29°C and the lowest is 26.5°C. The trend of decreasing SST from 29°C throughout the marine sequence is consistent with the variations in sediment grain size, reflecting the cooling trend of SST and sea-level lowering tendency during the last interglacial period. This new Eemian SST result agrees with other records in the region of SCS, showing a coherent descending trend, but spatial heterogeneity exists in timing and amplitude. The insolation changes might be the main force for initiating the SST evolution during the penultimate interglacial period.

Greenland ice sheet model parameters constrained using simulations of the Eemian Interglacial

Abstract. Using a new approach to force an ice sheet model, we performed an ensemble of simulations of the Greenland Ice Sheet evolution during the last two glacial cycles, with emphasis on the Eemian Interglacial. This ensemble was generated by perturbing four key parameters in the coupled regional climate-ice sheet model and by introducing additional uncertainty in the prescribed "background" climate change. The sensitivity of the surface melt model to climate change was determined to be the dominant driver of ice sheet instability, as reflected by simulated ice sheet loss during the Eemian Interglacial period. To eliminate unrealistic parameter combinations, constraints from present-day and paleo information were applied. The constraints include (i) the diagnosed present-day surface mass balance partition between surface melting and ice discharge at the margin, (ii) the modeled present-day elevation at GRIP; and (iii) the modeled elevation reduction at GRIP during the Eemian. Using these three constraints, a total of 360 simulations with 90 different model realizations were filtered down to 46 simulations and 20 model realizations considered valid. The paleo constraint eliminated more sensitive melt parameter values, in agreement with the surface mass balance partition assumption. The constrained simulations resulted in a range of Eemian ice loss of 0.4–4.4 m sea level equivalent, with a more likely range of about 3.7–4.4 m sea level if the GRIP δ18O isotope record can be considered an accurate proxy for the precipitation-weighted annual mean temperatures.

At what depth is the Eemian layer expected to be found at NEEM?

AbstractNo continuous record from Greenland of the Eemian interglacial period (130–115 ka BP) currently exists. However, a new ice-core drill site has been suggested at 77.449˚ N, 51.056˚Win north-west Greenland (North Eemian or NEEM). Radio-echo sounding images and flow model investigations indicate that an undisturbed Eemian record may be obtained at NEEM. In this work, a two-dimensional ice flow model with time-dependent accumulation rate and ice thickness is used to estimate the location of the Eemian layer at the new drill site. The model is used to simulate the ice flow along the ice ridge leading to the drill site. Unknown flow parameters are found through a Monte Carlo analysis of the flow model constrained by observed isochrones in the ice. The results indicate that the Eemian layer is approximately 60m thick and that its base is located approximately 100m above bedrock.

Crystal size variations in Eemian-age ice from the GRIP ice core, Central Greenland

Continuous measurements of ice crystal size have been carried out on an 80 m sequence between 2790 and 2870 m depth in the GRIP ice core from Central Greenland. The ice in this interval is at present considered to orginate from the Eemian interglacial period. The record reveals that the crystal size in ice older than 100,000 yr is highly dependent on climatic conditions at the time of snowfall. This dependence shows up as a strong correlation between δ 18O values and crystal size throughout the Eemian, as well as a negative correlation between crystal size and several soluble and insoluble impurities. Although high-resolution impurity records are available from selected parts of the Eemian ice, the study is not conclusive on which impurities are most effective in slowing grain growth. It is shown that the normal grain-growth process, commonly observed in the upper few hundred metres of polar ice sheets, does not yield grain sizes compatible with observed ones at this depth in the ice sheet, even in those parts of the Eemian ice where impurity drag effects are not present. Polygonization of crystals within the ice sheet and the nucleation and rapid growth of new grains at relatively high temperatures in the lowest part probably play an important role in producing the observed grain-size variations. The relevance of possible flow disturbances of the GRIP Eemian climatic record for the results presented is discussed briefly.

The role of the deep ocean in North Atlantic climate change between 70 and 130 kyr ago

THE suggestion1 that changes in North Atlantic Deep Water (NADW) production are linked through surface heat flux to the atmospheric temperature over Greenland is supported by earlier indications2,3 that NADW production decreased during glacial times, and by the subsequent finding4–6 that it declined during the Younger Dryas cool period at the end of the last glaciation. Changes in North Atlantic surface temperatures have been found7 to mirror high-frequency temperature changes recorded in Greenland ice cores over the past 80 kyr, but the connection to abyssal circulation has yet to be established, except for one or two isolated oscillations8,9. Here we present carbon and oxygen isotope analyses of benthic foraminifera in a high-resolution North Atlantic deep-sea sediment core for the period 70–130 kyr ago. These data allow us to reconstruct the history of NADW production, which shows a close correlation with Greenland climate variability for much of this time interval, suggesting that the climate influence of NADW variability was widespread. We see no evidence, however, for changes in NADW production during substage 5e (the Eemian interglacial period), in contrast with recent ice-core data10 which suggest severe climate instability in Greenland during this time period. Our results may support suggestions, based on data from a second ice core, that this apparent instability is an artefact caused by ice flow11. Alternatively, the Eemian climate instability may have had a different origin from the subsequent climate events.

COLEOPTERAN ASSOCIATIONS IN THE BRITISH ISLES DURING THE LATE QUATERNARY PERIOD

Summary1. The fauna of the climatic optimum of the Eemian Interglacial period contained thermophilous species and as such indicates a climate at least as warm as that today.2. The Chelford fauna dating from ca. 57,000 years before the present is comparable to that of Finland and Soviet Karelia. An analysis of the Carabidae suggests that within Scandinavia it is comparable with that which occurs at the top of the regio quercina.3. The Upton Warren fauna contains stenothermal arctic and boreal species and one Uralian species. The Carabidae suggest a comparison with the regio coniferina in Scandinavia.4. The Fladbury fauna has a large number of stenothermal arctic and boreal species and the Carabidae suggest a comparison with the top of the regio betulina in Scandinavia.5. The Colney Heath fauna contains stenothermal and stenotopic arctic and boreal species. The Carabidae suggest a comparison with the regio coniferina in Scandinavia.6. The St Bees fauna only contains a few remains of stenothermal boreal species and there is a considerable change in the percentage representation of the dominant species within the deposit.7. The stenothermal arctic element largely disappeared from the lowland fauna in a period of some 1600 years during pollen zone I.8. The thermophilous species immigrated into north‐west Europe in some 3000 years between pollen zones III and VII.9. Heliophilous species are well represented in Full‐ and Late‐glacial deposits. Their present restricted distribution is the result of Post‐glacial afforestation.10. Halophilous species have undergone a restriction in their range comparable to that of the heliophilous species.11. It is possible that certain observed distributional anomalies previously thought to result from glacial survival and subsequent extension of range may be the result of Post‐glacial survival in areas where there was only limited afforestation.12. During the late Quaternary era the changes in the distribution of the beetles parallel those of the angiosperms and gymnosperms.13. In terms of the Full‐, Late‐ and Post‐glacial periods the terminocratic grouping includes boreo‐alpine, northern oceanic, eurythermal and heliophilous species. The mediocratic grouping includes southern continental and oceanic eurythermal species.