Seeds of the extant Urtica kioviensis Rogow. (Urticaceae) and endocarps of the extinct Potamogeton sukaczevii Wieliczk. (Potamogetonaceae) were recorded in diverse plant macrofossil assemblages recovered from organic sediments exposed during excavations at Saham Toney, Norfolk, UK. Aminostratigraphical data show the sediments were deposited during the Ipswichian (Last Interglacial) Stage. Palynological data indicates deposition during the Carpinus pollen zone of the Ipswi- chian Stage—the latter part of pollen zone Ip IIb and Ip III. The records are noteworthy not only because they are the first in the British Pleistocene but also because of the geographical occurrences of these two species. Urtica kioviensis is absent from the British flora today and has a modern range in central and eastern Europe (only extending as far west as north–east Germany and Denmark), while the extinct Potamogeton sukaczevii has only been recovered from Late Pleistocene sediments in Belarus, Lithuania, Poland and western Russia. The presence of U. kioviensis along with other exotic species to the British Isles (e.g. Najas minor L. and Salvinia natans L., which today have central and southern ranges in Europe and in the case of S. natans occurs on other continents) may point to more continental conditions or warmer summer conditions during the second half of the Ipswichian Stage in southern Britain. No modern analogues occur in Britain for the assemblages recovered from Saham Toney. Evidence of colder winters or at least warmer summers at the time of deposition does not support the view that sea-level peaked in the Carpinus zone of the Eemian Stage (correlated with the Ipswichian Stage) associated with increased oceanicity. Southern Britain would have been under the influence of the Atlantic Ocean and a degree of oceanicity is supported by the presence of two thermophilous taxa, Hedera and Ilex, in the pollen spectra from Saham Toney. Alterna- tive explanations for the presence of these exotic species are that they were tolerating mild winters and cooler summers at the time of deposition or exploiting suitable micro-environments. The distribution of P. sukaczevii is probably an artefact of the distribution of expertise in the identification of Potamogeton fossil endocarps rather than having any palaeogeographic or palaeoclimatic significance. It is an extinct ancestor of the extant P. maackianus A. Benn, an eastern Asian pondweed. Its discovery in Britain encourages a reassessment of plant macrofossil assemblages from western Europe, which may lead to a consideration of the relationship between the Late Pleistocene vegetation of Europe and eastern Asia. Keywords Urtica kioviensis · Potamogeton sukaczevii · Ipswichian Stage · Palaeoclimate · Late Pleistocene · UK Introduction Investigations into the Pleistocene deposits at Saham Communicated by K.-E. Behre. Toney, Norfolk (52°35′56″N, 0°49′17″E; NGR TF 9125, 0197) were undertaken in 2007 during the construction * Michael H. Field firstname.lastname@example.org of an artificial fishing lake. Samples collected from this site have yielded a range of palaeoenvironmental infor- Faculty of Archaeology, Leiden University, van mation, including plant macrofossils of two angiosperm Steenisgebouw, Einsteinweg 2, 2333CC Leiden, species not hitherto known from the British Pleistocene The Netherlands 2 record: Urtica kioviensis Rogow. (Urticaceae) and Pota- School of Geography, Queen Mary University of London, mogeton sukaczevii Wieliczk. (Potamogetonaceae). The Mile End Road, London E1 4NS, UK Vol.:(0123456789) 1 3 Vegetation History and Archaeobotany site is some 33 km west–south–west of Norwich, within a This paper provides a summary of the site stratigraphy tributary of the river Wissey, which drains a catchment in and describes the new plant macrofossil records in advance central Norfolk and flows westward into the Wash basin of a full report on the multi-disciplinary investigations of (Fig. 1). The pre-Pleistocene geology of the area is Creta- this locality. ceous Chalk and the Pleistocene deposits locally consist The lithostratigraphy comprises up to 6 m of the organic of glaciogenic sediments of the Anglian Stage (MIS 12) sediments (beds 3–11), these underlie medium to coarse Lowestoft Formation and, along the course of the rivers, gravels with minor sand, clay and peat facies (beds 12–18) post-Anglian Stage fluvial sands and gravels and Holocene which were laid down under cold climate conditions during alluvium (Fig. 1). Excavation of these sands and gravels to the Devensian Stage (Weichselian Stage) (Fig. 1c, d). The construct the fishing lake resulted in recovery of fragments sedimentology of the organic sediments, which are predomi- of mammoth teeth from the basal gravels and the exposure nantly fine-grained, indicates deposition in slow-flowing to of underlying organic sediments. A series of trenches, sec- standing water with some terrestrial peat accumulation. tions and boreholes enabled the stratigraphy at the site to Aminostratigraphical data from Bithynia opercula con- be established and the deposits to be sampled for palaeoen- tained within the organic sediments (Penkman et al. 2011) vironmental analysis. dates them to the Last Interglacial Stage (Ipswichian Stage Fig. 1 a Western Norfolk; b The location of the site and cross section X–Y; c Cross section X–Y through the site including British Geological Survey borehole records (con- tains BGS materials, NERC 2016); d Summary stratigraphic log of beds 5–9 (overlying deposits, including fluvial sands and gravels are not shown) with the positions of the plant macrofossil and pollen samples (top, mid and base refers to position of subsamples within monoliths) and summary palynological data. B2 and B3 are adjacent sections, but shown in stratigraphic position 1 3 Vegetation History and Archaeobotany in Britain, Eemian Stage in continental Europe and MIS 5e The Urtica kioviensis Rogow. fossil seeds in the deep ocean record). The palynological information, together with molluscan and coleopteran assemblages, indi- Urtica kioviensis is monoecious and produces, after ferti- cates that full interglacial conditions developed during their lization, achenes (dry, one-seeded, indehiscent fruits). The deposition. The pollen spectra from these sediments indi- fruit wall is termed the pericarp if the fruit develops from cate the initial establishment of boreal woodland, followed a single ovary as in the case of U. kioviensis. The pericarp by deciduous Quercus/Corylus woodland, with later arrival has not preserved on the fossil specimens from Saham of Carpinus. Biostratigraphically, this vegetation pattern Toney and, thus, the specimens are called seeds. Samples is typical of the Ipswichian Stage and in particular pollen 32 (7 seeds), 37 (4 seeds) and 38 (4 seeds) yielded seeds of zones Ip I–Ip III (West 1980). U. kioviensis. The fossil seeds are ovoid and appear flat but The pollen spectra of beds 6–8 (Fig. 1d) show a domi- are slightly biconvex in side view (Fig. 2d, e). The greatest nance of trees and shrubs including Carpinus. It is not pre- width is in the centre of the seed. At the base is a broad sent in the underlying brown silt (bed 5), but increases to scar, while the apex tapers. The length of the Saham Toney levels between 5–10% in beds 6–8, while in bed 9 Carpinus seeds ranges between 1.85 and 1.58 mm, while the width is only present in small quantities (< 1%). Hedera is present ranges between 0.93 and 0.87 mm. Wolters et al. (2005) in beds 5 and 6, while Ilex is recorded in bed 9. The latter present additional biometric data for comparison. The fos- two thermophilous taxa have oceanic preferences as they do sil specimens are shown (Fig. 2d, e) alongside modern not tolerate cold winters (Iversen 1944). The pollen assem- material (Fig. 2g, h) for comparison of their shape. The blages recovered from beds 6, 8 and 9 allow correlation with modern specimens shown in Fig. 2 are achenes because the latter part of Ipswichian Stage pollen zone Ip IIb or the the seed is still encased in the pericarp. It was difficult to beginning of Ip III. remove the pericarps from the modern achenes as they closely adhere to the seeds. At the tip of the achene is a persistent, withered brush-like stigma. This feature has not preserved in the fossil specimens from Saham Toney. The new plant records The plant macrofossil assemblages also include seeds of the dioecious U. dioica (Fig. 2a, b). The clear size and Twenty-five sediment samples were collected from Saham morphological differences between the seeds of U. kio- Toney for plant macrofossil analysis. All plant macrofos- viensis and U. dioica from Saham Toney are illustrated in sils present in 200 cm of sediment from each sample were Fig. 2. Wolters et al. (2005) provide a useful morphologi- picked out to produce concentration data. The samples cal comparison of the fruits of these two species as well yielded diverse plant macrofossil assemblages. Five samples as others in the genus Urtica. contained remains of U. kioviensis and P. sukaczevii. These samples are from beds 6, 8 and 9; the shelly organic mud (samples 41, 38 and 37), grey shelly sandy silt (sample 33) and brown silt and clay (sample 32). Plant macrofossil data The Potamogeton sukaczevii Wieliczk. fossil from these sediment samples show that the water body was endocarps surrounded by damp woodland which included Alnus gluti- nosa. Where the drainage was better, Betula and Taxus bac- Preston (1995) comments that many terms have been used cata grew. Reed swamp occupied the margins of the water for modern propagules from species in the genus Pota- body which included taller plants such as Schoenoplectus mogeton including drupe, drupelet, achene, nut and nutlet. lacustris, Sparganium erectum and Typha. Where enough He concluded that none of these are entirely satisfactory light penetrated, shorter plants such as Lycopus europaeus, and preferred the term fruit. However, in a fossil context Mentha cf. aquatica and Solanum dulcamara grew in the it is often the case that not all the fruit is preserved. Aalto reed swamp. The presence of Cyperus fuscus and Lythrum (1970) recognized that the outer part of the fruit wall (the portula suggests that muddy areas also could be found at pericarp) is composed of exocarp cells below which is the water’s edge. The water body contained a diverse flora found a thin fleshy layer (parenchymatous mesocarp). of floating (e.g. Lemna and Salvinia natans), emergent (e.g. Often these do not preserve. What remains is the scleri- Potamogeton crispus and P. natans) and submergent (e.g. fied mesocarp sometimes termed the fruit stone, but also Ceratophyllum demersum, Najas flexilis, N. minor and N. called the endocarp. Here, Aalto’s example is followed and marina) plants. The aquatic component of the plant macro- the term endocarp is applied to the fossils. fossil assemblages indicates basic, mesotrophic conditions At Saham Toney, samples 41 (2 endocarps) and 33 (13 in still or slow-moving fresh water that contained little sus- endocarps) yielded endocarps of P. sukaczevii. Figure 3c, pended sediment. 1 3 Vegetation History and Archaeobotany Fig. 2 a, b Urtica dioica fossil seeds from Saham Toney (sample 33); viensis from Saham Toney (sample 32); g, h modern achenes of U. c the surface cells on a fossil specimen of a U. dioica from Saham kioviensis collected by E. Pechenyuk on the 19/9/1995 on the banks Toney (sample 33); d, e U. kioviensis fossil seeds from Saham Toney of the river Savala, Novokhopiorsk, Russia (these specimens were (sample 32); f the surface cells on a fossil specimen of an U. kio- kindly given to Field by E. Pechenyuk in 2000) d shows a fossil P. sukaczevii endocarp from sample (2006) comment that eastern European P. sukaczevii endo- 41 and, for comparison, a fossil P. sukaczevii endocarp carps often possess a wart at the base of the lid. This was kindly given to M. H. Field by Velichkevich in 1998 (from not evident in the Saham Toney specimens, but it may be Gołków, Poland; sample Q-11/64 A) is shown in Fig. 3a, that the base of the lid has been truncated (see Fig. 3c). b. The endocarps from Saham Toney are between 2.75 The style is in a central position and slightly tilted towards and 2.98 mm long and 2.25 and 2.86 mm wide. The length the dorsal side. The Saham Toney endocarps have a large measurement was taken from the base of the style to the stalk, flat sides with a small but deep central depression base of the endocarp. Velichkevich and Zastawniak (2006) in the shape of a comma whose tail points towards the describe the endocarps as broadly obovate in outline but stalk, an inconspicuous furrow running parallel to the it is clear in Fig. 3 that the endocarps are asymmetrical. dorsal margin along most of its length and pronounced, Like the Polish specimen in Fig. 3a and those illustrated large mamillate warts at the base of both sides of the in Velichkevich and Zastawniak (2006), the Saham Toney endocarp—all features consistent with Velichkevich and specimens have an irregularly sigmoid ventral margin (the Zastawniak’s (2006) description of the endocarp morphol- side opposite to the one where the lid is positioned). The ogy of P. sukaczevii. As with the eastern European fossils, upper two-thirds are very convex while at the base is a the Saham Toney endocarps have, in places, some of the straight section near the stalk. The lid, on the dorsal mar- fleshy mesocarp still adhering to the endocarp. The scleri- gin, is slightly curved, keeled and terminates just short fied mesocarp cells on the surface of the endocarp are very of the style base in a point. Velichkevich and Zastawniak characteristic (Fig. 3b, d). 1 3 Vegetation History and Archaeobotany Fig. 3 a, b Fossil Potamogeton sukaczevii endocarp kindly given to Field by Velichkevich in 1998 (from Gołków, Poland: sample Q-11/64 A) and a close up of its surface cells; c, d Fos- sil P. sukaczevii endocarp from Saham Toney (sample 41) and a close-up of its surface cells Compared to the Holocene, the climate of the Eemian Implications of these new records Stage was generally more oceanic in western and central Europe (Zagwijn 1996; Aalbersberg and Litt 1998). At the This paper reports on the first records of U. kioviensis and beginning of the Eemian Stage, differences between summer P. sukaczevii in the British Pleistocene. The Saham Toney and winter temperatures were at a maximum and Zagwijn organic sediments have been correlated with the Ipswi- (1996) interpreted the early Eemian finds of thermophilous chian Stage on aminostratigraphical, biostratigraphical and trees and aquatic plants as indicators of continental condi- lithostratigraphical grounds. Carpinus is present in beds tions with warmer summers. However, 2,000–3,000 years 6–8, which indicates sediment deposition during the mid- into the Last Interglacial Stage there was a change to a dle and latter half of the Last Interglacial Stage (either the more oceanic climate, which persisted throughout the rest latter part of pollen zone Ip IIb or Ip III). The presence of of the stage. Using the indicator species method Zagwijn P. sukaczevii, an extinct freshwater pondweed, which is (1996) reconstructed winter temperatures at Amsterdam to a characteristic component of Eemian plant macrofossil have been at a high of 3 °C in the Carpinus pollen zone assemblages recovered from Belarus, Lithuania, Poland and (E5). He noted that the winter temperature trend throughout western Russia (Velichkevich and Granoszewski 1996; Gra- the stage matched that of the rise and fall of sea-level, so noszewski 2003; Velichkevich et al. 2005; Velichkevich and that oceanicity during the stage reflected transgression and Zastawniak 2006), is also consistent with a Last Interglacial regression of the sea. Sea-level reached its peak during the Stage age. It should be noted that stratigraphically P. sukac- climatic optimum of the Eemian Stage (the Carpinus pollen zevii also occurs not only in the Eemian Stage but occasion- zone E5) and oceanic conditions prevailed (Zagwijn 1989). ally in the early Vistulian Stage, both in an interstadial and Palynological data from beds 6 and 9 at Saham Toney sup- during the glacial periods, such as at Horoszki Duże, Poland port the conclusion that during the Carpinus-phase condi- (Granoszewski 2003). The Vistulian Stage is correlated with tions were oceanic in southern England with the presence of the Devensian Stage in Britain and the Weichselian Stage in Hedera and Ilex, two taxa that Iversen (1944) showed were western Europe. 1 3 Vegetation History and Archaeobotany not tolerant of cold winters. This is unsurprising owing to particularly during the climatic optimum (in the Carpinus the proximity of the Atlantic Ocean. pollen zone E5) then it is possible that these species were Three species recorded from Saham Toney (Najas minor, tolerating milder winters and cooler temperatures during Salvinia natans and Urtica kiovensis) do not occur in the the growing season than they do today. Urtica kioviensis UK today (Stace 2010); they all have central or southern today occupies river valleys, growing in damp areas with European modern distributions (Fig. 4), with S. natans also Phragmites, Phalaris and Carex riparia reed swamp as well occurring in warm temperate or tropical areas in Africa and as Salix scrub and various Alnus plant associations (Wol- Asia (Mabberley 1997). If the Last Interglacial Stage (Ipswi- lert et al. 2003; Wolters 2005). Salvinia natans is a float- chian or Eemian Stages) climate conditions in western and ing aquatic pteridophyte that favours standing, fresh water, central Europe were more oceanic than in the Holocene, while Najas minor also lives in slow-moving fresh water skoe, Russia, 18—Liškyava (Lishkyava), Lithuania, 19—Yanionis Fig. 4 a The distribution of fossil sites that have yielded endocarps of (Yanyonis), Lithuania, 20—Netiesos (Nyatesos), Lithuania, 21— P. sukaczevii (the position of each site is taken from Velichkevich and Zastawniak (2006) except for the positions of the Polish sites which Komotovo, Belarus; b The modern distribution of U. kioviensis (Jalas are placed according to Granoszewski (2003). The sites are: 1— and Suominen 1988); c The modern distribution of Salvinia natans Gołków, Poland, 2—Horoszki Duże, Poland, 3—Bedlno, Poland, 4— in Europe (Jalas and Suominen 1987). Mabberley (1997) notes that it Knyazhevodtsy, Belarus, 5—Murava, Belarus, 6—Cherikov, Belarus, also occurs in warm temperate or tropical areas in Africa and Asia; d 7—Loev (Loyev), Belarus, 8—Borkhov Rov, Belarus, 9—Cherny Dashed line shows the modern northern limit of Najas minor (Hultén Bereg, Belarus, 10—Zabolot’e, Russia, 11—Panfilovo, Russia, 12— and Fries 1986), who comment that N. minor is not native in America Koz’ya, Russia, 13—Ryasna, Russia, 14—Nizhnyaya Boyarshchina, and that the knowledge of its distribution is partly incomplete Russia, 15—Zharki, Russia, 16—Staroe Zarech’e, Russia, 17—Shen- 1 3 Vegetation History and Archaeobotany where light penetrates down into the water column. Alter- plant macrofossil assemblages is needed to assess the wider natively, these species may have exploited suitable micro- significance of these findings. Potamogeton sukaczevii has environments possibly being transported from further afield an affinity with the extant P. maackianus which is found in by wildfowl. eastern Asia today. Reassessment of previous work and new Until its discovery at Saham Toney P. sukaczevii had only studies on western European plant macrofossils may lead to been found from Late Pleistocene sites in central and eastern a better understanding of the relationship between European Europe (Fig. 4). It is probable that this distribution is an and eastern Asian vegetation through time. artefact of the distribution of expertise in the identification Acknowledgements The authors would like to thank Ralph Fickling of Potamogeton fossil endocarps rather than having any pal- who made and reported the initial fossil discoveries at the site, Gerda aeogeographic or palaeoclimatic significance. Potamogeton E. M. Lamers (Institute of Biology, Faculty of Sciences, Leiden Uni- sukaczevii is an extinct species most closely related to the versity) for her help using the SEM, and Nick Ashton, Wojciech Gra- noszewski, Steve Forden, Roger Jacobi, Nigel Larkin, Mark Lewis, extant P. maackianus A. Benn, which today is distributed Simon Parfitt, and Sylvia Peglar for their contributions to this work. on the Korean Peninsula, in Japan, throughout the Russian They are also indebted to three anonymous referees who took time to Far East, China, Taiwan, Indonesia, Philippines, Myanmar read the manuscript and provide constructive comments. Fieldwork and Vietnam (Ito et al. 2009; Zhang 2009). Potamogeton was undertaken as part of the Ancient Human Occupation of Britain Project funded by the LeverhulmeTrust. sukaczevii is the youngest species in the entire P. maacki- anus phylogenetic group (Field et al. 2000; Granoszewski Open Access This article is distributed under the terms of the Crea- 2003). Its occurrence in the British Isles during the Late tive Commons Attribution 4.0 International License (http://creat iveco Pleistocene is an opportunity to prompt a consideration of mmons.or g/licenses/b y/4.0/), which permits unrestricted use, distribu- the relationships between the European and eastern Asian tion, and reproduction in any medium, provided you give appropriate vegetation. However, much more work, including a reassess- credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. ment of western European plant macrofossil assemblages, is needed to achieve this. References Conclusions Aalbersberg G, Litt T (1998) Multiproxy climate reconstructions The occurrence of U. kioviensis together with Najas minor for the Eemian and Early Weichselian. J Quat Sci 13:367–390 and Salvinia natans, all of which are exotic to the UK today Aalto M (1970) Potamogetonaceae fruits. I. 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