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Analysis of Late Pleistocene megafauna and puparia from the Lent dredging site, province of Gelderland (the Netherlands)

Published online by Cambridge University Press:  16 April 2021

David S. Douw Open the ORCID record for David S. Douw [Opens in a new window] ,
Belle E.I. van Rijssen ,
René H.B. Fraaije  and
Jonathan J.W. Wallaard
David S. Douw*
Affiliation:
Department of Earth Sciences, Utrecht University, P.O. Box 80115, 3508 TCUtrecht, the Netherlands Wereld van de Olifant, Binnenweg 4, 2132 CTHoofddorp, the Netherlands
Belle E.I. van Rijssen
Affiliation:
Department of Earth Sciences, Utrecht University, P.O. Box 80115, 3508 TCUtrecht, the Netherlands
René H.B. Fraaije
Affiliation:
Oertijdmuseum, Bosscheweg 80, 5283 WBBoxtel, the Netherlands
Jonathan J.W. Wallaard
Affiliation:
Oertijdmuseum, Bosscheweg 80, 5283 WBBoxtel, the Netherlands
*
Author for correspondence: David S. Douw, Email: d.s.douw@uu.nl
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Abstract

More than 900 vertebrate bones, ranging from Late Pleistocene to Holocene in age, have been identified in a collection that was recovered by a single dredging operation for the construction of artificial lakes near Lent (Nijmegen, province of Gelderland, the Netherlands). The Late Pleistocene assemblage comprises mainly Weichselian glacial fauna such as mammoths, reindeer and bison. Some Eemian fauna is represented as well, e.g. straight-tusked elephant. The abundance of certain species over others suggests that preservation bias had a considerable impact on this assemblage, while its time-averaged nature resulted in overrepresentation of certain species. A case study is here conducted on a fragmentary skull of a subadult woolly mammoth bull with embedded blowfly puparia. Some of these puparia are fully developed, indicating prolonged exposure of the mammoth carcass.

Keywords

mammalsmammothblowfliesWeichselianEemian
Type
Geo(im)pulse
Information
Netherlands Journal of Geosciences , Volume 100 , 2021 , e10
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Copyright
© The Author(s), 2021. Published by Cambridge University Press.

Introduction

Skeletal remains of Late Pleistocene mammals can be found across the Netherlands as they were preserved by fluvial sediments which now surface across the country (Doppert et al., Reference Doppert, Ruegg, Van Staalduinen, Zagwijn and Zandstra1975; Van Kolfschoten & Laban, Reference Van Kolfschoten and Laban1995; De Mulder et al., Reference De Mulder, Geluk, Ritsema, Westerhoff and Wong2003; Gouw & Erkens, Reference Gouw and Erkens2007; Laban & Van der Meer, Reference Laban and Van der Meer2011; DINOloket, 2020). At dredging sites, large numbers of such fossils have been collected and subsequently published (Mol et al., Reference Mol, Van den Bergh and De Vos1999; Van Kolfschoten, Reference Van Kolfschoten2001; Van der Jagt, Reference Van der Jagt2005; Van Kolfschoten et al., Reference Van Kolfschoten, Van der Jagt, Beeren, Argiti, Van der Leije, Van Essen, Busschers, Stoel and Van der Plicht2011).

Thousands of specimens have been collected during dredging operations at Lent near Nijmegen (51°52'38.6"N, 5°51'57.5"E; Fig. 1A). In total, 909 bones in this collection were considered suitable for identification. The faunal remains comprise 732 Holocene and 177 Late Pleistocene mammals. The Late Pleistocene fauna is analysed, with particular attention paid to fly puparia found embedded in skull fragments of the woolly mammoth, Mammuthus primigenius (Blumenbach, Reference Blumenbach1799), contained in the collections of the Oertijdmuseum at Boxtel (MAB10673).

Figure 1. Location of the dredging site and the abundance of Pleistocene megafauna fossils.

Late Pleistocene faunal abundance

One of the most common mammals found in Late Pleistocene deposits throughout the Netherlands is the woolly mammoth (Mol et al., Reference Mol, Van den Bergh and De Vos1999), and Lent is no exception (Fig. 2F further below). Mammoth remains make up 50.3% of the Late Pleistocene assemblage from this locality (Fig. 1B) and are represented mainly as fragmentary skulls, mandibles and pelvises. This high percentage is due in part to their colossal size, which increases the chances of preservation. Additionally, this taxon was abundant and widely distributed during the Late Pleistocene, having been well adapted to grazing and cold temperatures.

Figure 2. Late Pleistocene fossils found at the Lent site: A. dextral radius of a cave hyena (Crocuta crocuta spelaea); B. dextral humerus of a cave hyena; C. atlas of a wild boar (Sus scrofa); D. pelvic fragment of a wolf (Canis lupus lupus); E. dextral lower M3 fragment of a straight-tusked elephant (Palaeoloxodon antiquus); and F. sinistral upper M1 molar of a woolly mammoth (Mammuthus primigenius). Black bar represents 5 cm.

Wild horse Equus ferus Boddaert, Reference Boddaert1785 is also common (25.1%) at Lent, followed by steppe bison Bison priscus Bojanus, Reference Bojanus1827 (11.7%) and reindeer Rangifer tarandus (Linnaeus, Reference Linnaeus1758) (5.6%). Horse remains are abundant, likely because their ecological niche was relatively large compared to that of reindeer or steppe bison. Horses were able to cope with open steppe conditions as well as with more forested environments, while bison and reindeer were restricted to open steppe environments. This flexibility explains why horses are represented not only in interstadial and stadial faunas, but possibly even in interglacial ones (Von Koenigswald, Reference Von Koenigswald2003; Saarinen et al., Reference Saarinen, Eronen, Fortelius, Sepp and Lister2016). Furthermore, reindeer material is less likely to be preserved compared to other herbivores in the present collection, as their thin-walled humeri and radii were easily eroded or gnawed away by predators and scavengers (Van der Jagt, Reference Van der Jagt2005; Zimov et al., Reference Zimov, Zimov, Tikhonov and Chapin2012).

This collection contains only a few remains of the woolly rhinoceros, Coelodonta antiquitatis (Blumenbach, Reference Blumenbach1799), accounting for 2.8%. This low percentage likely represents the true number of individuals of this species that lived during the Late Pleistocene. Woolly rhinos are thought to have been solitary, with males having territories similar to extant forms (Garutt, Reference Garutt1999; Diedrich, Reference Diedrich2008; Becker et al., Reference Becker, Dini and Scherler2014). By contrast, mammoths, horses, steppe bison and reindeer lived in herds, and thus had considerably higher population densities compared to the woolly rhinoceros.

The geology at the site suggests these herbivores are to be associated with cold climatic conditions during the Late Pleistocene, and in particular with the Weichselian Glacial (DINOloket, 2020). They were occupants of the ‘mammoth steppe’, the cold, dry grass steppe that extended over much of the Northern Hemisphere during the Late Pleistocene (Zimov et al., Reference Zimov, Zimov, Tikhonov and Chapin2012).

Predators are not well represented in this assemblage. Of the three bones recovered, two belong to the cave hyena (Crocuta crocuta spelaea Goldfuss, Reference Goldfuss1823; Fig. 2A–B). The third is a heavily mineralised pelvic fragment of a wolf (Canis lupus, Linnaeus, Reference Linnaeus1758; Fig. 2D). Fossils of cave hyenas are not particularly rare in the Netherlands (Reumer et al., Reference Reumer, Mol and Borst2010), while the absence of skeletal remains of other predators suggests that these were too rare in the ecosystem for them to be represented in this assemblage, as predators often are. It is difficult to connect the presence of predators with a particular climatic condition as they can be found both in warm and cold periods of the Late Pleistocene (Von Koenigswald, Reference Von Koenigswald2003). Other predators missing from the present assemblage are notably cave lions, brown bears and cave bears (cf. Bocherens, Reference Bocherens2015).

Species associated with warmer climatic conditions are few in the assemblage. Three molars of the straight-tusked elephant Palaeoloxodon antiquus (Falconer & Cautley, Reference Falconer and Cautley1846; Fig. 2E) and two heavily mineralised bones of wild boar (Sus scrofa Linnaeus, Reference Linnaeus1758; Fig. 2C) have been identified. The geology at the dredging site suggests that the remains of these animals date back to the Eemian and are possibly reworked (Laban & Van der Meer, Reference Laban and Van der Meer2011; DINOloket, 2020).

Fly puparia

In Pleistocene mammalian bones, blowfly puparia, typically the species Protophormia terraenovae Robineau-Desvoidy, Reference Robineau-Desvoidy1830, are occasionally found (Gautier & Schumann, Reference Gautier and Schumann1973; Vervoenen, Reference Vervoenen1991; Gautier, Reference Gautier1995; Verhagen & Mol, Reference Verhagen and Mol2009; Van der Plicht et al., Reference Van der Plicht, Post and Mol2012; Mähler et al., Reference Mähler, Wappler, Sanmugaraja, Menger and von Koenigswald2016). This Holarctic species of the Calliphoridae family is absent from the Netherlands at present (Erzinçlioğlu, Reference Erzinçlioğlu2009; Vanin et al., Reference Vanin, Turchetto, Galassi and Cattaneo2009; Verhagen & Mol, Reference Verhagen and Mol2009). This species breeds on the carcasses of vertebrates, on which their larvae feed and pupate (Roux et al., Reference Roux, Gers, Telmon and Legal2006; Erzinçlioğlu, Reference Erzinçlioğlu2009). Until the 1970s, remains of puparia in Pleistocene mammalian fossils were thought to be rare; subsequently, there have been more observed (Gautier & Schumann, Reference Gautier and Schumann1973; Vervoenen, Reference Vervoenen1991; Gautier, Reference Gautier1995; Verhagen & Mol, Reference Verhagen and Mol2009; Van der Plicht et al., Reference Van der Plicht, Post and Mol2012; Mähler et al., Reference Mähler, Wappler, Sanmugaraja, Menger and von Koenigswald2016).

A skull of a woolly mammoth (MAB10673) from Lent comprises 43 separate fragments (e.g. Fig. 2F), 15 of which contained puparia. Only five puparia (MAB11313; Table 1) were deemed to be sufficiently preserved for identification in accordance with criteria presented in the relevant literature (Gautier & Schumann, Reference Gautier and Schumann1973; Erzinçlioğlu, Reference Erzinçlioğlu2009; see Fig. 3). All five have been identified as P. terraenovae on the basis of length and morphology. Two of these have larger widths than is typical of this species (Erzinçlioğlu, Reference Erzinçlioğlu1988), but this is likely due to infilling sediment having flattened the puparia inside the skull fragments.

Table 1. Length, width and shape factor of the Lent puparia, compared with the data from Erzinçlioğlu (Reference Erzinçlioğlu1988)

* Estimated length because these puparia were incomplete. Shape factor calculated by dividing the greatest width by overall length.

Figure 3. MAB11313 Puparium 1 (unopened) ventral (A) and dorsal (B) view; Puparium 3 (opened) ventral (C) and dorsal (D) view. Black bar represents 1 mm.

These puparia were recovered from skull fragments close to natural body orifices such as the eyes (zygomatic arch), which are used by blowflies to enter the body, as has been documented for colonisation of human carcasses (Haskell et al., Reference Haskell, Hall, Cervenka, Clark, Haglund and Sorg1997). The larvae likely travelled further into the carcass to feed (Roux et al., Reference Roux, Gers, Telmon and Legal2006).

The mammoth host was likely a subadult bull, as the wear patterns observed on the lower dextral M2 suggest that it had reached the age of 24.5 ± 2 African Equivalent Years (Laws, Reference Laws1966), an estimate based on the procedure outlined by Haynes (Reference Haynes1993) for extinct proboscideans. Sex was determined by comparing the dimensions of the isolated occipital pars basalis to those of the analogous bone in the complete skull of an adult female (MAB11628; Van der Merwe et al., Reference Van der Merwe, Bezuidenhout and Seegers1995; Averianov, Reference Averianov, Shoshani and Tassy1996).

By applying modern forensic methods to the analysis of puparia, one can approximate the minimum temperature at the time of the mammoth’s death (Gilbert & Bass, Reference Gilbert and Bass1967; Vanin et al., Reference Vanin, Turchetto, Galassi and Cattaneo2009). Protophormia terraenovae can develop its eggs only in temperatures higher than 9°C (Nuorteva, Reference Nuorteva1987; Grassberger & Reiter, Reference Grassberger and Reiter2002). Reconstructions by Caspers & Freund (Reference Caspers and Freund2001) suggest that mean July temperatures during the Weichselian ranged from approximately 17°C during the early Weichselian to 3°C during the Late High Glacial stadials. This suggests that this mammoth died either during the early Weichselian or during an interstadial of the High Glacial when temperatures were high enough for egg development.

Three of the five puparia analysed were empty, suggesting that the flies fully developed and left their puparium (Nuorteva, Reference Nuorteva1987). Hence, the carcass was likely exposed to the surface long enough for them to become fully developed. Studies on the periods of development of extant P. terraenovae suggest that full development takes between 35 and 39 days (Marchenko, Reference Marchenko2001; Grassberger & Reiter, Reference Grassberger and Reiter2002). However, this varies according to region and temperature (Grassberger & Reiter, Reference Grassberger and Reiter2002). Other heat sources, such as the carcass itself (Henssge, Reference Henssge1988) and ‘maggot mass heating’, should be taken into account, as they can cause development times to be considerably shorter (Haskell et al., Reference Haskell, Hall, Cervenka, Clark, Haglund and Sorg1997; Erzinçlioğlu, Reference Erzinçlioğlu2009).

Conclusions

Late Pleistocene megafaunal remains from Lent document a considerable preservational bias in favour of mammoths and against species such as reindeer, while accurately representing the abundance of horses, steppe bison and woolly rhinoceros throughout climatic changes during the Late Pleistocene. The presence of Protophormia terraenovae puparia in the woolly mammoth bull skull MAB10763 suggests carcass colonisation occurred during the early Weichselian, or during an interstadial of the High Glacial, during which temperatures met the minimum requirement for egg development.

Acknowledgements

We wish to thank Werner Peters for collecting and donating the Lent material over a period of 3 years, with impeccable consideration for all types of fossil material at the site. Identification of the mammalian material was made possible thanks to Charles Schouwenburg and Dick Mol, while puparia identification was aided by comparative material from Lambert Van Es. Geological interpretations of the area were made possible thanks to Wim Hoek. The senior author thanks Wilma Wessels for supervision throughout this research project. Comments from Wilma Wessels, Charles Schouwenburg, Tom Giltaij, Jan van der Made and John Jagt on early drafts were extremely useful and essential in the writing process. Credit to Ruben Winter for taking the photographs for this project. We thank Dick Mol, Hans van Essen and an anonymous reviewer for pertinent comments and feedback. Special thanks to the MSB zand en grind (Beijer Bouwgrondstoffen, Boskalis and Spaansen collective) for allowing fossil collection at the dredging site with excellent hospitality.

Declaration of interest

The authors declare adherence to Committee on Publication Ethics (COPE) guidelines and have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

References

Averianov, A.O., 1996. Sexual dimorphism in the mammoth skull, teeth, and long bones. In: Shoshani, J. & Tassy, P. (eds): The Proboscidea. Evolution and palaeoecology of elephants and their relatives. Oxford University Press (Oxford): 260–269.Google Scholar
Becker, D., Dini, M. & Scherler, L., 2014. Woolly rhinoceros from the Pleniglacial of Ajoie (Jura Canton, Switzerland): anatomical description and ecological implications. Abstract, Swiss Geoscience Meeting 2014, Fribourg. Available at: https://doi.org/10.5281/zenodo.18895.CrossRefGoogle Scholar
Blumenbach, J.F., 1799. Abbildungen naturhistorischer Gegenstände, viertes Heft, Tafeln. Dieterich (Göttingen): 31–40.Google Scholar
Bocherens, H., 2015. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews 117: 42–71.CrossRefGoogle Scholar
Boddaert, P., 1785. Elenchus animalium. Sistens quadrupedia huc usque nota, eorumque varietates 1: 1–38.Google Scholar
Bojanus, H.L. 1827. De Uro nostrate eiusque sceleto commentatio. Nova Acta Academiae Caesareae-Leopoldino Carolinae Germanicae Naturae Curiosorum 23: 413–478.Google Scholar
Caspers, G. & Freund, H., 2001. Vegetation and climate in the Early- and Pleni-Weichselian in northern central Europe. Journal of Quaternary Science 16: 31–48.3.0.CO;2-3>CrossRefGoogle Scholar
De Mulder, E.F., Geluk, M.C., Ritsema, I., Westerhoff, W.E. & Wong, T.E., 2003. De ondergrond van Nederland. Nederlands Instituut voor Toegepaste Geowetenschappen, TNO (Utrecht): 376 pp.Google Scholar
Diedrich, C.G., 2008. A skeleton of an injured Coelodonta antiquitatis from the Late Pleistocene of north-western Germany. Cranium 25: 29–43.Google Scholar
DINOloket, 2020. Netherlands Institute of Applied Geoscience. www.dinoloket.nl/en/subsurface-models, last accessed on 19 May 2020.Google Scholar
Doppert, J.W.C., Ruegg, G.H.J., Van Staalduinen, C.J., Zagwijn, W.H. & Zandstra, J.G., 1975. Formaties van het Kwartair en Boven-Tertiair in Nederland. Toelichting bij geologische overzichtskaarten van Nederland. Rijks Geologische Dienst (Haarlem): 11–56.Google Scholar
Erzinçlioğlu, Y.Z., 1988. The larvae of the species of Phormia and Boreellus: northern, cold-adapted blowflies (Diptera: Calliphoridae). Journal of Natural History 22: 11–16.CrossRefGoogle Scholar
Erzinçlioğlu, Y.Z., 2009. Fly puparia associated with the Condover mammoths. In: Lister, A.M. (ed.): Late-glacial mammoth skeletons (Mammuthus primigenius) from Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological significance. Geological Journal 44: 447–479.CrossRefGoogle Scholar
Falconer, H. & Cautley, P.T., 1846. Fauna antiqua sivalensis, being the fossil zoology of the Sewalik Hills, in the north of India 1. Smith, Elder and Company (London).CrossRefGoogle Scholar
Garutt, N., 1999. Skull pathologies in Coelodonta antiquitatis: implications about social behaviour and ecology. Deinsea 6: 175–186.Google Scholar
Gautier, A., 1995. Bovenpleistocene zoogdieren van Oudenaarde Donk (België), fossiele vliegenpoppen uit de Vlaamse Vallei en elders en nog een en ander over de Vlaamse Vallei. Cranium 12: 73–81.Google Scholar
Gautier, A. & Schumann, H., 1973. Puparia of the subarctic or black blowfly (Protophormia terraenovae Robineau-Desvoidy 1830) in a skull of a Late Eemian(?) bison at Zemst, Brabant (Belgium). Palaeogeography, Palaeoclimatology, Palaeoecology 14: 119–125.CrossRefGoogle Scholar
Gilbert, B.M. & Bass, W.M., 1967. Seasonal dating of burials from the presence of fly pupae. American Antiquity, 32: 534–535.CrossRefGoogle Scholar
Goldfuss, A., 1823. Osteologische Beiträge zur Kenntniss verschiedener Säugethiere der Vorwelt: Fortsetzung. Ueber die Hoelen-Hyaene (Hyaena spelaea). Nova Acta Physico-Medica Academiae, Caesareae Leopoldino-Carolinae Naturae Curiosorum 3: 456–490.Google Scholar
Gouw, M.J.P. & Erkens, G., 2007. Architecture of the Holocene Rhine-Meuse delta (the Netherlands) – a result of changing external controls. Netherlands Journal of Geosciences/Geologie en Mijnbouw 86: 23–54.CrossRefGoogle Scholar
Grassberger, M. & Reiter, C., 2002. Effect of temperature on development of the forensically important holarctic blowfly Protophormia terraenovae (Robineau-Desvoidy) (Diptera: Calliphoridae). Forensic Science International 128: 177–182.CrossRefGoogle Scholar
Haskell, N.H., Hall, R.D., Cervenka, V.J. & Clark, M.A., 1997. On the body: insects’ life stage presence, their post-mortem artifacts. In: Haglund, W.D. & Sorg, M.H. (eds): Forensic taphonomy: the postmortem fate of human remains. CRC Press (Boca Raton, FL): 449–458.Google Scholar
Haynes, G., 1993. Mammoths, mastodons, and elephants: biology, behaviour and the fossil record. Cambridge University Press (Cambridge). 321-353.Google Scholar
Henssge, C., 1988. Death time estimation in case work. I. The rectal temperature time of death nomogram. Forensic Science International 38: 209–236.CrossRefGoogle ScholarPubMed
Laban, C. & Van der Meer, J.J., 2011. Pleistocene glaciation in the Netherlands. In: Ehlers, J., Gibbard, P.L. & Hughes, P.D. (eds): Developments in Quaternary Sciences 15: 247–260.CrossRefGoogle Scholar
Laws, R.M., 1966. Age criteria for the African elephant: Loxodonta a. africana . African Journal of Ecology 4: 1–37.CrossRefGoogle Scholar
Linnaeus, C., 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decimna, reformata. Laurentius Salvius (Stockholm): 11–56.CrossRefGoogle Scholar
Mähler, B., Wappler, T., Sanmugaraja, M., Menger, F. & von Koenigswald, W., 2016. Upper Pleistocene blow flies (Diptera: Calliphoridae) trapped in fossilized crania of large mammals discovered from gravel pits in the Rhine rift valley from Hesse (Germany). Palaeontologia Electronica 19: 1–10.Google Scholar
Marchenko, M.I., 2001. Medicolegal relevance of cadaver entomofauna for the determination of the time since death. Forensic Science International 120: 89–109.CrossRefGoogle Scholar
Mol, D., Van den Bergh, G.D. & De Vos, J., 1999. Fossil proboscideans from the Netherlands, the North Sea and the Oosterschelde Estuary. Deinsea 6: 119–146.Google Scholar
Nuorteva, P., 1987. Empty puparia of Phormia terraenovae R.-D. (Diptera: Calliphoridae) as forensic indicators. Annales Entomologicae Fennicae 33: 53–56.Google Scholar
Reumer, J., Mol, D. & Borst, W., 2010. The first Late Pleistocene coprolite of Crocuta crocuta spelaea from the North Sea. Deinsea 14: 15–18.Google Scholar
Robineau-Desvoidy, J.B., 1830. Essai sur les myodaires. Mémoires présentés par divers Savants à l’Académie Royale des Sciences de l’Institut de France 2: 1–813.Google Scholar
Roux, O., Gers, C., Telmon, N. & Legal, L., 2006. Circular dispersal of larvae in the necrophagous Diptera Protophormia terraenovae (Diptera: Calliphoridae). Annales de la Société entomologique de France 42: 51–56.CrossRefGoogle Scholar
Saarinen, J., Eronen, J., Fortelius, M., Sepp, H. & Lister, A.M., 2016. Patterns of diet and body mass of large ungulates from the Pleistocene of western Europe, and their relation to vegetation. Palaeontologia Electronica 19: 1–58.Google Scholar
Van der Jagt, I.M., 2005. Pleistoceen Woerden. Een archeozoölogisch onderzoek naar de ouderdom van de Cervidae, in het bijzonder Rangifer tarandus. MA Thesis. Leiden University (Leiden). 1–26.Google Scholar
Van der Merwe, N.J., Bezuidenhout, A.J. & Seegers, C.D., 1995. The skull and mandible of the African elephant (Loxodonta africana). Onderstepoort Journal of Veterinary Research 62: 245–260.Google Scholar
Van der Plicht, H., Post, K. & Mol, D., 2012. Over aasvliegen en een mammoetkalf uit de Eurogeul. Cranium 29: 14–19.Google Scholar
Van Kolfschoten, T., 2001. Pleistocene mammals from the Netherlands. Bollettino della Società Paleontologica Italiana 40: 209–216.Google Scholar
Van Kolfschoten, T. & Laban, C., 1995. Pleistocene terrestrial mammal faunas from the North Sea area. Mededelingen Rijks Geologische Dienst 52: 135.Google Scholar
Van Kolfschoten, T., Van der Jagt, I., Beeren, Z., Argiti, V., Van der Leije, J., Van Essen, H., Busschers, F.S., Stoel, P. & Van der Plicht, H., 2011. A remarkable collection of Late Pleistocene reindeer (Rangifer tarandus) remains from Woerden (The Netherlands). Quaternary International 238: 4–11.CrossRefGoogle Scholar
Vanin, S., Turchetto, M., Galassi, A. & Cattaneo, C., 2009. Forensic entomology and the archaeology of war. Journal of Conflict Archaeology 5: 127–139.CrossRefGoogle Scholar
Verhagen, A. & Mol, D., 2009. De Groote Wielen: er was eens… Wie woonden er in De Groote Wielen in de ijstijd? Norg, DrukWare, 1–192.Google Scholar
Vervoenen, M., 1991. Pleistocene vleesvliegen puparia uit hoornpitten van Bison priscus . Cranium 8: 57–58.Google Scholar
Von Koenigswald, W., 2003. Mode and causes for the Pleistocene turnovers in the mammalian fauna of Central Europe. Deinsea 10: 305–312.Google Scholar
Zimov, S.A., Zimov, N.S., Tikhonov, A.N. & Chapin, F.S. I, 2012. Mammoth steppe: a high-productivity phenomenon. Quaternary Science Reviews 57: 26–45.CrossRefGoogle Scholar
Figure 0

Figure 1. Location of the dredging site and the abundance of Pleistocene megafauna fossils.

Figure 1

Figure 2. Late Pleistocene fossils found at the Lent site: A. dextral radius of a cave hyena (Crocuta crocuta spelaea); B. dextral humerus of a cave hyena; C. atlas of a wild boar (Sus scrofa); D. pelvic fragment of a wolf (Canis lupus lupus); E. dextral lower M3 fragment of a straight-tusked elephant (Palaeoloxodon antiquus); and F. sinistral upper M1 molar of a woolly mammoth (Mammuthus primigenius). Black bar represents 5 cm.

Figure 2

Table 1. Length, width and shape factor of the Lent puparia, compared with the data from Erzinçlioğlu (1988)

Figure 3

Figure 3. MAB11313 Puparium 1 (unopened) ventral (A) and dorsal (B) view; Puparium 3 (opened) ventral (C) and dorsal (D) view. Black bar represents 1 mm.