Plant macrofossils of the plants of the sediment sample from the Yuka mammoth cranium: (1) Hordeum sp., caryopsis; (2) Potentilla sp., nutlet; (3) Androsac e sp., seed; (4) Carex sp., nutlet; (5) Cyperaceae ( Carex , Scirpus ?), nutlet; (6) Tephroseris cf. atropurpurea , achene; (7) Batrachium sp., nutlet; (8) Poaceae, caryopsis; (9) Deschampsia sp., caryopsis; (10) Potamogeton vaginatus, endocarp. 

Contexts in source publication

Context 1

... and Alyoshina, 1972; Beug, 2004). Non-pollen palynomorphs (NPPs) were identi fi ed using descriptions, pictures and photo- graphs published by van Geel (2001). A total of 329 pollen grains and spores, which were taken as 100% for determining percentages of pollen taxa, were counted in the sample. The total number of palynomorphs, including the NPPs, was 481. The total number of NPPs was taken as 100% when calculating the percentages of individual NPPs. The plant macrofossil sample was washed through a 250- μ m mesh sieve and then air-dried. A total of 40 mL dry matter was examined and subjected to analysis using a Carl Zeiss Stemi 2000-C stereomicro- scope. Twigs, leaf remains, fruits, seeds, mosses and a variety of uniden- ti fi ed vegetative remains were identi fi ed from the sample, following Nikitin (1969). In total, 25 taxa of pollen and spores were identi fi ed in the sample. Herbaceous taxa (92%) dominated the pollen spectrum with 6% of the pollen originating from trees (Fig. 4). Among herbs, Artemisia pollen dominated (46%), together with indeterminable forb pollen (12.5%), and pollen from the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae, and Brassicaceae. Indeterminable herbaceous pollen related to very small (about 6 – 7 μ m) and probably not mature tricolpate forms. Arboreal pollen was represented primarily by Salix and Betula sect. Nanae and 2% of spores came from ferns and lycopods. The highest percentage of the NPPs was determined to be the remains of the green alga Botryococcus (up to 90%). The sample contained a mixture from the vegetative parts of the plants and the mineral fraction in a ratio of ca. 60:40 (before sieving). The vegetative parts of plants included small parts of herbaceous stalks and leaves. Other remains were represented by leaves and twigs of Bryales, seeds of herbs, 20 mammoth hairs, 13 ephippia of Daphnia , fi ve ostracod shells, seven chitin fragments of С oleoptera , two fragments of Oribatidae and a fragment of mammoth cranium. Plant macrofossils included 12 taxa (Fig. 5). Seeds belonged to the Potamogetonaceae — Potamogeton vaginatus (one endocarp); Poaceae — Deschampsia sp. (one caryopsis), Hordeum sp. (one caryopsis), Poaceae sp. (one caryopsis); Cyperaceae — Carex spp. (seven fragments of nutlets), Cyperaceae ( Carex , Scirpus ?) (one nutlet), Rosaceae — Potentilla sp. (three nutlets); Ranunculaceae — Ranunculus sp. (one fragment of nutlet), Batrachium sp. (one nutlet); Primulaceae — Androsac e sp. (one seed); С aryophyllaceae (one seed); Asteraceae — Tephroseris cf. atropurpurea (one achene). Among the Cyperaceae macrofossils, the nutlet, which combines di- agnostic features of Carex and Sciprus , was found. The absence of bristles at the base of the nutlet makes it similar to that of Carex ; however, bristles are not often preserved in fossil nutlets of Sciprus , Eriophorum and Eleocharis . The walls of this nutlet are thick and shape of nutlet is hemi- spherical, which is more typical for Sciprus than for Carex (Fig. 5). According to radiocarbon dating (34,300 + 260/ − 240 yr BP), the Yuka mammoth lived during the termination of the Kargin Interstadial. The presumed climatic optimum for the Kargin Interstadial in the Laptev Sea region occurred between ca. 44 – 32 kyr BP (see review in Wetterich et al., 2014). As mentioned above, the vegetation of the MIS3 optimum became mosaic and the earlier-prevailing tundra- steppe was combined with willow shrubs or relatively mesophytic communities that were spread throughout protected and wet places (Andreev et al., 2011). Plant macrofossil data from palaeorecords dating between 48 and 33 kyr BP from the Bykovsky Peninsula indicate that remains from typical steppe and meadow plants, e.g., Festuca , Kobresia , Hordeum , Linum , Silene and Potentilla , were numerous. Such plant communities suggest relatively warm summers. Macrofossils of taxa from permanently wet habitats were also found in the Middle Weichselian Interstadial, dating between about 48 and 35 kyr BP (Kienast et al., 2005). Paleobotanical data obtained in this study represent two sets of taxa (macro- and microfossils), whereas macrofossils more re fl ect the local vegetation in the burial place of the Yuka mammoth, the pollen spectrum mainly re fl ects the regional vegetation. In general, macrofossil plant remains obtained from the sediment sample in the Yuka mammoth's skull are characteristic of the herbaceous taxa that are widespread on the modern Yakutian tundra (Table 1). However, seeds of P. vaginatus and Batrachium sp. as well as ostracod shells and ephippia of Daphnia re fl ect the existence of small freshwater ponds with stagnant or slowly moving water exactly at the site where the mammoth carcass was found. In addition, P. vaginatus re- quires warm summers and does not occur in modern vegetation north of the 12 °C mean July isotherm (Flora Sibiri, 1988a). One of the nutlets of Cyperaceae combines the characteristics of both Carex and Scirpus (Fig. 5). The genus Scirpus does not grow in the Eastern Siberian Arctic today. Scirpus lacustris and S. orientalis occur in the southern part of Central Yakutia and S. maximowiczii grows in the Yana-Indigirka lowland . Scirpus inhabits water meadows, the banks of ponds and landscape falls (Flora Sibiri, 1990). Deshampsia sp. and Ranunculus sp. are also indicators of wetland in the Yuka's burial place (Table 1). The high abundance of Botryococcus remnants in the pollen spectrum con fi rms the aquatic nature of the environment at the burial place of the Yuka mammoth. The fi nding of Carex nutlet fragments does not contradict the reconstruction of wetland in the studied sites; different species of Carex can be dominant in variety of steppe, tundra as well as wet intrazonal communities. Except for aquatic and wet-site plants, the macrofossil spectrum includes steppe elements such as Caryophyllaceae and Potentilla sp. The type of vegetation, including these macrofossils, resembles the plant community that is relic today, and persisted in var- ious parts of Metaberingia (Yurtzev, 2001). It is called ‘ mesic-xeric meadows enriched with steppe elements ’ , which are sometimes zoo- genic and have a sparse canopy of shrubs. The dominants of this community are grasses, and shrubs are represented by Rosa acicularis , Pentaphylloides (Potentilla) fruticosa , Salix glauca and Betula nana . This conclusion is con fi rmed by pollen data. Despite the dominance in the pollen spectrum of Artemisia , the Ranunculaceae and Poaceae are highly abundant; the percentages of Salix and Betula sect. Nanae are also signif- icant and are the most common arboreal taxa. The pollen spectrum from the sample studied is generally typical of pollen spectra from the late Kargin records of the Eastern Siberian Arctic. Six leading taxa in the spectra are Artemisia , the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae and Brassicaceae. Five taxa (ex- cluding Ranunculaceae) might characterise steppe-like vegetation, and we suggest that it is a regional (zonal) feature. However, the percentage of Artemisia (46%) is signi fi cantly higher than in contemporary pollen records from North Yakutia (e.g., in Bol'shoy Lyakhovsky Island it is up to 4% (Wetterich et al., 2014) and Kurungnakh Island and Bykovsky Peninsula it is up to 10% (Andreev et al., 2011). The overrepresentation of Artemisia might be explained by the existence of disturbed soils close to the studied site, where wormwoods grew as weeds. Indirect evidence of this is that a high abundance of Glomus chlamydospores, an indicator of soil erosion (van Geel et al., 2003 and pollen of the Chenopodiaceae, a very rare family in Arctic, mainly of ruderal origin (Arkticheskaya fl ora SSSR, 1966) was found. The results of pollen and plant macrofossil analyses con fi rm that the Yuka mammoth lived during the optimum of the Kargin Interstadial. Paleobotanical data presented in this study suggest that the vegetation of the MIS3 optimum in the Eastern Siberian Arctic became mosaic; zonal tundra-steppe might have been combined with mesic-xeric meadows enriched with steppe elements. The burial place of the mammoth could have been a small shallow freshwater pond with stagnant or slowly moving water. An explanation for the overrepresentation of Artemisia in the pollen spectrum is the existence of disturbed soils close to the studied site, where wormwoods grew as weeds. The research was partly supported by the Russian Foundation of Basic Research (12-04-98510). We thank Larisa Frolova for valuable ad- vice and we thank all our Russian colleagues who assisted during the fi eld and laboratory work. We also especially thank two anonymous re- viewers for very valuable scienti fi c comments and ...

Context 2

... diversity and abundance of testate amoebae and ostracods (Bobrov et al., 2004; Wetterich et al., 2005). The modern climate of the Oyogos Yar coast is characterised by mean temperatures during the warmest month of approximately 4 °C, whereas the mean temperature during the coldest month is approximately − 30 °C (Rivas-Martinez, 1996 – 2009). The Oyogos Yar belongs to the southern arctic tundra province of Eastern Siberia, which is dominated by Alopecurus alpinus , Salix polaris and Carex bigelowii ssp. arctisibirica (Aleksandrova, 1980). The vegetation is also represented by Salix pulchra , S. reptans, S. sphenophylla, Vaccinium vitis-idaea, Arctous tetragona and Eriophorum vaginatum (Egorova et al., 1991) and by the mosses Aulacomnium turgidum , Dicranum elongatum , Tomethypnum nitens and Hylomium splendens (Andreev et al., 1987). The boreal or sub- arctic shrubs ( Alnaster fruticosa or Betula nana ) that are found to the south are completely absent from the study site today (Kienast et al., 2011). Two frozen sediment samples from the area of the skull condyles were collected for pollen and plant macrofossil analyses. This was the only place to obtain the samples since the Yukagirs washed all the mammoth remains, including the gut, with water from a pump. The intestinal contents were absent, and therefore, a sample for the analysis of microfossils and macroremains was collected from the skull. The sample was thawed, dried, and sieved through a 250- μ m mesh to remove coarse organic matter, which was later used for plant macrofossil analysis. The sample was sequentially treated with 10% HCl and 10% KOH and washed with distilled water. A 7- μ m mesh sieve was used to remove the fi ne-grained fraction. Pollen residues were mounted in glycerine and analysed using a Zeiss AxioImager D2 light microscope with 400× magni fi cation. The identi fi cation of pollen and spores was performed using a reference pollen collection and pollen atlases (Kuprianova and Alyoshina, 1972; Beug, 2004). Non-pollen palynomorphs (NPPs) were identi fi ed using descriptions, pictures and photo- graphs published by van Geel (2001). A total of 329 pollen grains and spores, which were taken as 100% for determining percentages of pollen taxa, were counted in the sample. The total number of palynomorphs, including the NPPs, was 481. The total number of NPPs was taken as 100% when calculating the percentages of individual NPPs. The plant macrofossil sample was washed through a 250- μ m mesh sieve and then air-dried. A total of 40 mL dry matter was examined and subjected to analysis using a Carl Zeiss Stemi 2000-C stereomicro- scope. Twigs, leaf remains, fruits, seeds, mosses and a variety of uniden- ti fi ed vegetative remains were identi fi ed from the sample, following Nikitin (1969). In total, 25 taxa of pollen and spores were identi fi ed in the sample. Herbaceous taxa (92%) dominated the pollen spectrum with 6% of the pollen originating from trees (Fig. 4). Among herbs, Artemisia pollen dominated (46%), together with indeterminable forb pollen (12.5%), and pollen from the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae, and Brassicaceae. Indeterminable herbaceous pollen related to very small (about 6 – 7 μ m) and probably not mature tricolpate forms. Arboreal pollen was represented primarily by Salix and Betula sect. Nanae and 2% of spores came from ferns and lycopods. The highest percentage of the NPPs was determined to be the remains of the green alga Botryococcus (up to 90%). The sample contained a mixture from the vegetative parts of the plants and the mineral fraction in a ratio of ca. 60:40 (before sieving). The vegetative parts of plants included small parts of herbaceous stalks and leaves. Other remains were represented by leaves and twigs of Bryales, seeds of herbs, 20 mammoth hairs, 13 ephippia of Daphnia , fi ve ostracod shells, seven chitin fragments of С oleoptera , two fragments of Oribatidae and a fragment of mammoth cranium. Plant macrofossils included 12 taxa (Fig. 5). Seeds belonged to the Potamogetonaceae — Potamogeton vaginatus (one endocarp); Poaceae — Deschampsia sp. (one caryopsis), Hordeum sp. (one caryopsis), Poaceae sp. (one caryopsis); Cyperaceae — Carex spp. (seven fragments of nutlets), Cyperaceae ( Carex , Scirpus ?) (one nutlet), Rosaceae — Potentilla sp. (three nutlets); Ranunculaceae — Ranunculus sp. (one fragment of nutlet), Batrachium sp. (one nutlet); Primulaceae — Androsac e sp. (one seed); С aryophyllaceae (one seed); Asteraceae — Tephroseris cf. atropurpurea (one achene). Among the Cyperaceae macrofossils, the nutlet, which combines di- agnostic features of Carex and Sciprus , was found. The absence of bristles at the base of the nutlet makes it similar to that of Carex ; however, bristles are not often preserved in fossil nutlets of Sciprus , Eriophorum and Eleocharis . The walls of this nutlet are thick and shape of nutlet is hemi- spherical, which is more typical for Sciprus than for Carex (Fig. 5). According to radiocarbon dating (34,300 + 260/ − 240 yr BP), the Yuka mammoth lived during the termination of the Kargin Interstadial. The presumed climatic optimum for the Kargin Interstadial in the Laptev Sea region occurred between ca. 44 – 32 kyr BP (see review in Wetterich et al., 2014). As mentioned above, the vegetation of the MIS3 optimum became mosaic and the earlier-prevailing tundra- steppe was combined with willow shrubs or relatively mesophytic communities that were spread throughout protected and wet places (Andreev et al., 2011). Plant macrofossil data from palaeorecords dating between 48 and 33 kyr BP from the Bykovsky Peninsula indicate that remains from typical steppe and meadow plants, e.g., Festuca , Kobresia , Hordeum , Linum , Silene and Potentilla , were numerous. Such plant communities suggest relatively warm summers. Macrofossils of taxa from permanently wet habitats were also found in the Middle Weichselian Interstadial, dating between about 48 and 35 kyr BP (Kienast et al., 2005). Paleobotanical data obtained in this study represent two sets of taxa (macro- and microfossils), whereas macrofossils more re fl ect the local vegetation in the burial place of the Yuka mammoth, the pollen spectrum mainly re fl ects the regional vegetation. In general, macrofossil plant remains obtained from the sediment sample in the Yuka mammoth's skull are characteristic of the herbaceous taxa that are widespread on the modern Yakutian tundra (Table 1). However, seeds of P. vaginatus and Batrachium sp. as well as ostracod shells and ephippia of Daphnia re fl ect the existence of small freshwater ponds with stagnant or slowly moving water exactly at the site where the mammoth carcass was found. In addition, P. vaginatus re- quires warm summers and does not occur in modern vegetation north of the 12 °C mean July isotherm (Flora Sibiri, 1988a). One of the nutlets of Cyperaceae combines the characteristics of both Carex and Scirpus (Fig. 5). The genus Scirpus does not grow in the Eastern Siberian Arctic today. Scirpus lacustris and S. orientalis occur in the southern part of Central Yakutia and S. maximowiczii grows in the Yana-Indigirka lowland . Scirpus inhabits water meadows, the banks of ponds and landscape falls (Flora Sibiri, 1990). Deshampsia sp. and Ranunculus sp. are also indicators of wetland in the Yuka's burial place (Table 1). The high abundance of Botryococcus remnants in the pollen spectrum con fi rms the aquatic nature of the environment at the burial place of the Yuka mammoth. The fi nding of Carex nutlet fragments does not contradict the reconstruction of wetland in the studied sites; different species of Carex can be dominant in variety of steppe, tundra as well as wet intrazonal communities. Except for aquatic and wet-site plants, the macrofossil spectrum includes steppe elements such as Caryophyllaceae and Potentilla sp. The type of vegetation, including these macrofossils, resembles the plant community that is relic today, and persisted in var- ious parts of Metaberingia (Yurtzev, 2001). It is called ‘ mesic-xeric meadows enriched with steppe elements ’ , which are sometimes zoo- genic and have a sparse canopy of shrubs. The dominants of this community are grasses, and shrubs are represented by Rosa acicularis , Pentaphylloides (Potentilla) fruticosa , Salix glauca and Betula nana . This conclusion is con fi rmed by pollen data. Despite the dominance in the pollen spectrum of Artemisia , the Ranunculaceae and Poaceae are highly abundant; the percentages of Salix and Betula sect. Nanae are also signif- icant and are the most common arboreal taxa. The pollen spectrum from the sample studied is generally typical of pollen spectra from the late Kargin records of the Eastern Siberian Arctic. Six leading taxa in the spectra are Artemisia , the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae and Brassicaceae. Five taxa (ex- cluding Ranunculaceae) might characterise steppe-like vegetation, and we suggest that it is a regional (zonal) feature. However, the percentage of Artemisia (46%) is signi fi cantly higher than in contemporary pollen records from North Yakutia (e.g., in Bol'shoy Lyakhovsky Island it is up to 4% (Wetterich et al., 2014) and Kurungnakh Island and Bykovsky Peninsula it is up to 10% (Andreev et al., 2011). The overrepresentation of Artemisia might be explained by the existence of disturbed soils close to the studied site, where wormwoods grew as weeds. Indirect evidence of this is that a high abundance of Glomus chlamydospores, an indicator of soil erosion (van Geel et al., 2003 and pollen of the Chenopodiaceae, a very rare family in Arctic, mainly of ruderal origin (Arkticheskaya fl ora SSSR, 1966) was found. The results of pollen and plant macrofossil analyses con fi rm that the Yuka mammoth lived during the optimum of the Kargin Interstadial. Paleobotanical data presented in this study suggest that the vegetation of the MIS3 optimum in the Eastern ...

Context 3

... limits can be identi fi ed as 44 – 32 kyr BP. Pollen records related to the early part of MIS 3 (50 – 40 kyr BP) are characterised by the dominance of Cyperaceae and Poaceae pollen, with some Artemisia and Salix , re fl ecting the tundra-steppe environment (Andreev et al., 2011). Higher pollen concentrations and a high abundance of Cyperaceae, Poaceae, Artemisia , Caryophyllaceae and Ranunculaceae, as well as the permanent presence of Salix pollen characterise the MIS3 optimum (Andreev et al., 2011; Wetterich et al., 2014). It appears that vegetation became mosaic; open grass-sedge associations combined with willow shrubs spread into more protected and wet places. Warmer summer air temperatures and moister climatic conditions that suggest the existence of small ponds during the MIS3 optimum are also con fi rmed by plant macrofossil records (Kienast et al., 2005), and the fi nding of green algae remains (Andreev et al., 2011; Wetterich et al., 2014) and a high diversity and abundance of testate amoebae and ostracods (Bobrov et al., 2004; Wetterich et al., 2005). The modern climate of the Oyogos Yar coast is characterised by mean temperatures during the warmest month of approximately 4 °C, whereas the mean temperature during the coldest month is approximately − 30 °C (Rivas-Martinez, 1996 – 2009). The Oyogos Yar belongs to the southern arctic tundra province of Eastern Siberia, which is dominated by Alopecurus alpinus , Salix polaris and Carex bigelowii ssp. arctisibirica (Aleksandrova, 1980). The vegetation is also represented by Salix pulchra , S. reptans, S. sphenophylla, Vaccinium vitis-idaea, Arctous tetragona and Eriophorum vaginatum (Egorova et al., 1991) and by the mosses Aulacomnium turgidum , Dicranum elongatum , Tomethypnum nitens and Hylomium splendens (Andreev et al., 1987). The boreal or sub- arctic shrubs ( Alnaster fruticosa or Betula nana ) that are found to the south are completely absent from the study site today (Kienast et al., 2011). Two frozen sediment samples from the area of the skull condyles were collected for pollen and plant macrofossil analyses. This was the only place to obtain the samples since the Yukagirs washed all the mammoth remains, including the gut, with water from a pump. The intestinal contents were absent, and therefore, a sample for the analysis of microfossils and macroremains was collected from the skull. The sample was thawed, dried, and sieved through a 250- μ m mesh to remove coarse organic matter, which was later used for plant macrofossil analysis. The sample was sequentially treated with 10% HCl and 10% KOH and washed with distilled water. A 7- μ m mesh sieve was used to remove the fi ne-grained fraction. Pollen residues were mounted in glycerine and analysed using a Zeiss AxioImager D2 light microscope with 400× magni fi cation. The identi fi cation of pollen and spores was performed using a reference pollen collection and pollen atlases (Kuprianova and Alyoshina, 1972; Beug, 2004). Non-pollen palynomorphs (NPPs) were identi fi ed using descriptions, pictures and photo- graphs published by van Geel (2001). A total of 329 pollen grains and spores, which were taken as 100% for determining percentages of pollen taxa, were counted in the sample. The total number of palynomorphs, including the NPPs, was 481. The total number of NPPs was taken as 100% when calculating the percentages of individual NPPs. The plant macrofossil sample was washed through a 250- μ m mesh sieve and then air-dried. A total of 40 mL dry matter was examined and subjected to analysis using a Carl Zeiss Stemi 2000-C stereomicro- scope. Twigs, leaf remains, fruits, seeds, mosses and a variety of uniden- ti fi ed vegetative remains were identi fi ed from the sample, following Nikitin (1969). In total, 25 taxa of pollen and spores were identi fi ed in the sample. Herbaceous taxa (92%) dominated the pollen spectrum with 6% of the pollen originating from trees (Fig. 4). Among herbs, Artemisia pollen dominated (46%), together with indeterminable forb pollen (12.5%), and pollen from the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae, and Brassicaceae. Indeterminable herbaceous pollen related to very small (about 6 – 7 μ m) and probably not mature tricolpate forms. Arboreal pollen was represented primarily by Salix and Betula sect. Nanae and 2% of spores came from ferns and lycopods. The highest percentage of the NPPs was determined to be the remains of the green alga Botryococcus (up to 90%). The sample contained a mixture from the vegetative parts of the plants and the mineral fraction in a ratio of ca. 60:40 (before sieving). The vegetative parts of plants included small parts of herbaceous stalks and leaves. Other remains were represented by leaves and twigs of Bryales, seeds of herbs, 20 mammoth hairs, 13 ephippia of Daphnia , fi ve ostracod shells, seven chitin fragments of С oleoptera , two fragments of Oribatidae and a fragment of mammoth cranium. Plant macrofossils included 12 taxa (Fig. 5). Seeds belonged to the Potamogetonaceae — Potamogeton vaginatus (one endocarp); Poaceae — Deschampsia sp. (one caryopsis), Hordeum sp. (one caryopsis), Poaceae sp. (one caryopsis); Cyperaceae — Carex spp. (seven fragments of nutlets), Cyperaceae ( Carex , Scirpus ?) (one nutlet), Rosaceae — Potentilla sp. (three nutlets); Ranunculaceae — Ranunculus sp. (one fragment of nutlet), Batrachium sp. (one nutlet); Primulaceae — Androsac e sp. (one seed); С aryophyllaceae (one seed); Asteraceae — Tephroseris cf. atropurpurea (one achene). Among the Cyperaceae macrofossils, the nutlet, which combines di- agnostic features of Carex and Sciprus , was found. The absence of bristles at the base of the nutlet makes it similar to that of Carex ; however, bristles are not often preserved in fossil nutlets of Sciprus , Eriophorum and Eleocharis . The walls of this nutlet are thick and shape of nutlet is hemi- spherical, which is more typical for Sciprus than for Carex (Fig. 5). According to radiocarbon dating (34,300 + 260/ − 240 yr BP), the Yuka mammoth lived during the termination of the Kargin Interstadial. The presumed climatic optimum for the Kargin Interstadial in the Laptev Sea region occurred between ca. 44 – 32 kyr BP (see review in Wetterich et al., 2014). As mentioned above, the vegetation of the MIS3 optimum became mosaic and the earlier-prevailing tundra- steppe was combined with willow shrubs or relatively mesophytic communities that were spread throughout protected and wet places (Andreev et al., 2011). Plant macrofossil data from palaeorecords dating between 48 and 33 kyr BP from the Bykovsky Peninsula indicate that remains from typical steppe and meadow plants, e.g., Festuca , Kobresia , Hordeum , Linum , Silene and Potentilla , were numerous. Such plant communities suggest relatively warm summers. Macrofossils of taxa from permanently wet habitats were also found in the Middle Weichselian Interstadial, dating between about 48 and 35 kyr BP (Kienast et al., 2005). Paleobotanical data obtained in this study represent two sets of taxa (macro- and microfossils), whereas macrofossils more re fl ect the local vegetation in the burial place of the Yuka mammoth, the pollen spectrum mainly re fl ects the regional vegetation. In general, macrofossil plant remains obtained from the sediment sample in the Yuka mammoth's skull are characteristic of the herbaceous taxa that are widespread on the modern Yakutian tundra (Table 1). However, seeds of P. vaginatus and Batrachium sp. as well as ostracod shells and ephippia of Daphnia re fl ect the existence of small freshwater ponds with stagnant or slowly moving water exactly at the site where the mammoth carcass was found. In addition, P. vaginatus re- quires warm summers and does not occur in modern vegetation north of the 12 °C mean July isotherm (Flora Sibiri, 1988a). One of the nutlets of Cyperaceae combines the characteristics of both Carex and Scirpus (Fig. 5). The genus Scirpus does not grow in the Eastern Siberian Arctic today. Scirpus lacustris and S. orientalis occur in the southern part of Central Yakutia and S. maximowiczii grows in the Yana-Indigirka lowland . Scirpus inhabits water meadows, the banks of ponds and landscape falls (Flora Sibiri, 1990). Deshampsia sp. and Ranunculus sp. are also indicators of wetland in the Yuka's burial place (Table 1). The high abundance of Botryococcus remnants in the pollen spectrum con fi rms the aquatic nature of the environment at the burial place of the Yuka mammoth. The fi nding of Carex nutlet fragments does not contradict the reconstruction of wetland in the studied sites; different species of Carex can be dominant in variety of steppe, tundra as well as wet intrazonal communities. Except for aquatic and wet-site plants, the macrofossil spectrum includes steppe elements such as Caryophyllaceae and Potentilla sp. The type of vegetation, including these macrofossils, resembles the plant community that is relic today, and persisted in var- ious parts of Metaberingia (Yurtzev, 2001). It is called ‘ mesic-xeric meadows enriched with steppe elements ’ , which are sometimes zoo- genic and have a sparse canopy of shrubs. The dominants of this community are grasses, and shrubs are represented by Rosa acicularis , Pentaphylloides (Potentilla) fruticosa , Salix glauca and Betula nana . This conclusion is con fi rmed by pollen data. Despite the dominance in the pollen spectrum of Artemisia , the Ranunculaceae and Poaceae are highly abundant; the percentages of Salix and Betula sect. Nanae are also signif- icant and are the most common arboreal taxa. The pollen spectrum from the sample studied is generally typical of pollen spectra from the late Kargin records of the Eastern Siberian Arctic. Six leading taxa in the spectra are Artemisia , the Ranunculaceae, Poaceae, Caryophyllaceae, Asteraceae and Brassicaceae. Five taxa (ex- cluding Ranunculaceae) might characterise steppe-like vegetation, and we suggest that it is a ...

Context 4

... parts of plants included small parts of herbaceous stalks leaves. Other remains were represented by leaves and twigs of Bryales, seeds of herbs, 20 mammoth hairs, 13 ephippia of Daphnia, five ostracod shells, seven chitin fragments of Сoleoptera, two fragments of Oribatidae and a fragment of mammoth cranium. Plant macrofossils included 12 taxa (Fig. 5). Seeds belonged to the PotamogetonaceaePotamogeton vaginatus (one endocarp); Poaceae-Deschampsia sp. (one caryopsis), Hordeum sp. (one caryopsis), Poaceae sp. (one caryopsis); Cyperaceae-Carex spp. (seven fragments of nutlets), Cyperaceae (Carex, Scirpus?) (one nutlet), Rosaceae-Potentilla sp. (three nutlets); Ranunculaceae-Ranunculus ...

Context 5

... features of Carex and Sciprus, was found. The absence of bristles at the base of the nutlet makes it similar to that of Carex; however, bristles are not often preserved in fossil nutlets of Sciprus, Eriophorum and Eleocharis. The walls of this nutlet are thick and shape of nutlet is hemispherical, which is more typical for Sciprus than for Carex (Fig. ...

Context 6

... freshwater ponds with stagnant or slowly moving water exactly at the site where the mammoth carcass was found. In addition, P. vaginatus requires warm summers and does not occur in modern vegetation north of the 12 °C mean July isotherm (Flora Sibiri, 1988a). One of the nutlets of Cyperaceae combines the characteristics of both Carex and Scirpus (Fig. 5). The genus Scirpus does not grow in the Eastern Siberian Arctic today. Scirpus lacustris and S. orientalis occur in the southern part of Central Yakutia and S. maximowiczii grows in the Yana-Indigirka lowland. Scirpus inhabits water meadows, the banks of ponds and landscape falls (Flora Sibiri, 1990). Deshampsia sp. and Ranunculus sp. ...