title: A systematic review of animal predation creating pierced shells: implications for the archaeological record of the Old World.
abstract: BACKGROUND
  The shells of molluscs survive well in many sedimentary contexts and yield information about the diet of prehistoric humans. They also yield evidence of symbolic behaviours through their use as beads for body adornments. Researchers often analyse the location of perforations in shells to make judgements about their use as symbolic objects (e.g., beads), the assumption being that holes attributable to deliberate human behaviour are more likely to exhibit low variability in their anatomical locations, while holes attributable to natural processes yield more random perforations. However, there are non-anthropogenic factors that can cause perforations in shells and these may not be random. The aim of the study is compare the variation in holes in shells from archaeological sites from the Old World with the variation of holes in shells pierced by mollusc predators.
  
  
  METHODS
  Three hundred and sixteen scientific papers were retrieved from online databases by using keywords, (e.g., 'shell beads'; 'pierced shells'; 'drilling predators'); 79 of these publications enabled us to conduct a systematic review to qualitatively assess the location of the holes in the shells described in the published articles. In turn, 54 publications were used to assess the location of the holes in the shells made by non-human predators.
  
  
  RESULTS
  Almost all archaeological sites described shells with holes in a variety of anatomical locations. High variation of hole-placement was found within the same species from the same site, as well as among sites. These results contrast with research on predatory molluscs, which tend to be more specific in where they attacked their prey. Gastropod and bivalve predators choose similar hole locations to humans.
  
  
  CONCLUSIONS
  Based on figures in the analysed articles, variation in hole-location on pierced shells from archaeological sites was similar to variation in the placement of holes created by non-human animals. Importantly, we found that some predators choose similar hole locations to humans. We discuss these findings and identify factors researchers might want to consider when interpreting shells recovered from archaeological contexts.
authors.affiliation: Independent researcher, Poznań, Poland.
authors.forename: Anna Maria
authors.initials: AM
authors.lastname: Kubicka
authors.affiliation: Department of Cell Biology, Adam Mickiewicz University in Poznań, Poznań, Poland.
authors.forename: Zuzanna M
authors.initials: ZM
authors.lastname: Rosin
authors.affiliation: Institute of Zoology, Poznań University of Life Sciences, Poznań, Poland.
authors.forename: Piotr
authors.initials: P
authors.lastname: Tryjanowski
authors.affiliation: School of Medicine, University of Liverpool, Liverpool, United Kingdom.|Department of Archeology, Classics and Egyptology, University of Liverpool, Liverpool, United Kingdom.
authors.forename: Emma
authors.initials: E
authors.lastname: Nelson
keywords: Bivalvia; Gastropoda; Interspecies interactions; Jewellery; Predators; Scaphopoda; Shell beads