‘Cervified’: a method for the morphometric identification of red deer (Cervus elaphus), fallow deer (Dama dama), and roe deer (Capreolus capreolus) archaeological bones

‘Cervified’: a method for the morphometric identification of red deer (Cervus elaphus), fallow deer (Dama dama), and roe deer (Capreolus capreolus) archaeological bones | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article ‘Cervified’: a method for the morphometric identification of red deer (Cervus elaphus), fallow deer (Dama dama), and roe deer (Capreolus capreolus) archaeological bones Veronica Aniceti, Mauro Rizzetto, Francesco Giacalone This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6636241/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 16 Jan, 2026 Read the published version in Archaeological and Anthropological Sciences → Version 1 posted 11 You are reading this latest preprint version Abstract This paper presents a new method for differentiating postcranial bone remains of red deer ( Cervus elaphus ), fallow deer ( Dama dama ), and roe deer ( Capreolus capreolus ) from archaeological contexts. These species have a very similar bone morphology and partly overlapping size ranges, often preventing species-level identification in faunal research. Traditional methods, such as the use of diagnostic morphological criteria, or aDNA analysis, present practical and methodological limitations. In this study, bone biometric data from modern specimens, sourced from various institutions across northern and southern Europe, are analysed to determine size ranges and shape indices that can separate the three species, focussing on both adult and immature individuals. By comparing measurement values from archaeological remains with these biometric criteria, species identification can be attempted. The low-tech nature of this approach and the transparent, diagrammatic presentation of biometric results make the new method objective, highly accessible, and inclusive. In European and Mediterranean archaeology, the separation of fallow deer from other cervid species is crucial in the investigation of past biogeography, trading activities, hunting strategies, cultural values, and social status. red deer fallow deer roe deer method identification biometry archaeology Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 Figure 11 Introduction Red deer, fallow deer, and roe deer are among the most widespread large-sized wild mammals of Europe and, not surprisingly, they often played an important role in human society. These animals were hunted as a source of food and other materials, whether as part of subsistence or recreational activities, held significant religious and magico-medicinal meaning, and could represent, alive or on the table, a symbol of social differentiation. As such, deer has been the subject of extensive literature (e.g., Chapman & Chapman, 1975 ; Grant, 1988 ; Soderberg, 2004 ; Holmes, 2014 ; Schmidt, 2014 ; Miller & Sykes, 2016 ; Albarella & Aniceti, 2024 ; Almeida et al., 2024 ). In the last two decades, zooarchaeological research on this taxon has benefitted from methodological improvements and the application of new analytical techniques, from ageing tools (Bowen et al., 2016 ; Marín et al., 2024 ), to isotopic and genetic analyses (Skog et al., 2009 ; McDevitt & Zachos, 2014 ; Sykes et al., 2016 ; Baker et el., 2024a; Baker et al., 2024b ), to biometry (Carden et al., 2012 ; Sykes et al., 2013 ; Karastoyanova et al., 2020 ). This paper aims to contribute to a fundamental yet often overlooked aspect of cervid archaeological research, functional to many of the studies and techniques mentioned above, namely species-level identification of faunal remains. In zooarchaeology, the identification of the remains of red deer ( Cervus elaphus ), fallow deer ( Dama dama ), and roe deer ( Capreolus capreolus ) from European faunal assemblages has often posed significant challenges (Figs. 1–3). Bone morphological similarities and partial size overlaps, in addition to the fragmented nature of most faunal materials, can prevent species-level identification, limiting the interpretative potentials of the zooarchaeological evidence. The problem of identification of European cervid remains from historical times arises from the introduction and spread of the fallow deer in the Continent since at least Roman times and then, more consistently, during the Middle Ages (Davis & MacKinnon, 2009 ; Masseti & Vernesi, 2014 ; Baker et al., 2024a ; Baker et al., 2024b ). Indeed, this species partly overlaps in size with the red deer, which is larger, and, more rarely, with the roe deer, a smaller animal. The extent of such size overlaps in past animal populations is, however, often difficult to assess, due to substantial diachronic and spatial size variability. The factors determining deer size ranges and their fluctuations are multiple. Sexual dimorphism is especially strong in red deer and fallow deer (Post et al., 1999 ; McElligott et al., 2001 ; Milošević-Zlatanović et al., 2016 ), implying size overlaps of large male fallow deer and small female red deer. Climatic conditions also impact on deer size, cervids being larger in colder regions and periods (Toïgo et al., 2006 ; Apollonio & Chirichella, 2023 ), as do specific geographical contexts such as islands and isolated regions, where these animals are smaller (Lister, 1989 ; Mulville, 2010 ). Finally, anthropogenic factors such as hunting pressure, habitat reduction, forced relocation, and emparkment have also shown to lead to a reduction in size (Serrano et al., 2007 ; Karastoyanova et al., 2020 ). In faunal assemblages, the potential presence of immature individuals, whether identifiable or not as such, leads to even greater size overlaps and can further compromise species-level identification. Currently, the separation of red, fallow, and roe deer archaeological remains is attempted through different and sometimes complementary methodologies: Size analyses . When large assemblages are available and through comparisons with data from related sites, it is sometimes possible to identify size clusters that can be attributed to different species (Davis & MacKinnon, 2009 ; Aniceti et al., 2021 ; Albarella & Davis 1996 ). Large samples of cervid biometric values are, however, rarely available; in addition, size overlaps between species are frequent, for the reasons mentioned above, limiting the efficiency of species-level identification based on size alone. Morphological criteria . These have long been established for the distinction between red and fallow deer (Di Stefano, 1995 ; Lister, 1996 ). Their use, however, is inevitably biased by a certain degree of subjectivity and by the level of expertise of different researchers, and ideally requires the assistance of abundant reference material; in addition, not all criteria are equally diagnostic (Lister, 1996 ), implying that taxonomic separation is more difficult and less reliable on some zones. More importantly, the results of such attempts cannot be objectively scrutinised by other researchers. Ancient DNA ( aDNA ). Genetic analyses can be very effective when enough collagen is preserved. aDNA analyses are not used with the specific purpose of separating fallow deer and red deer remains but rather to investigate genetic structures and past zoogeographical dynamics in these species; in doing so, however, some specimens considered in these studies are shown to belong to the other species and excluded (e.g., Baker et al., 2024a ; Beglane et al., 2018 ). Despite their effectiveness, genetic analyses are nonetheless constrained by severe cost limitations and are destructive. Zooarchaeology by Mass Spectrometry (ZooMS), on the other hand, cannot currently separate between red and fallow deer remains (Buckley & Whitcher Kansa, 2011 ). In addition, this technique too involves destructive procedures and can be economically impacting when applied to large samples, despite being less time-consuming and more affordable compared to genetic analyses. This study introduces a new method for the separation of fallow deer from red deer and roe deer remains. The method consists in the use of biometric indices and size ranges, derived from modern cervid skeletons of known species, against which indices and raw measurement values from archaeological material can be plotted for comparison. Such approach implies a key set of advantages: It allows for more objective identification attempts, as these are represented graphically and therefore open to scrutiny; The separation potentials of size and shape are combined to maximize chances of reliable species-level identification; In some skeletal elements, diagnostic indices can also be used on immature individuals, when size alone is of no use, or on zones where no morphological criteria exist (e.g., long bone shafts); Lastly, the method relies on the use of the calliper and osteometric board and is therefore widely accessible and inclusive. The new method does not aim to replace the identification methods listed above, nor to dismiss any potential future improvements to such methods or the importance of an appropriate use of reference collections. Rather, it aims to provide additional support to the study of the history of cervids in temperate Europe and the Mediterranean; in phylogenetic studies of fallow deer in a specific period or region, for example, the use of an efficient, objective, and inexpensive biometric method may assist in the identification of a more reliable sample of fallow deer bones on which to focus funds for genetic analyses. This biometric method relies on an extensive dataset which, on the other hand, is not exhaustive of all regions and available modern cervid skeletons; however, the variability in size ranges covered and the consistency in the validity of shape indices across different regions ensure the applicability of the method to other regions, provided critical interpretation of the results is always used. Ultimately, more widespread and correct identification of cervid archaeological and palaeontological remains will contribute to the reconstruction of past biodiversity, human-animal-environment interactions, and socio-cultural and religious practices. The method here presented is the first output of the Cervified Project, conducted by the authors to investigate the potentials of biometric approaches in the species-level identification of archaeological European cervid remains, as well as in the study of past and present cervid bone morphometry. Materials and methodological approach Biometric data from modern red, fallow, and roe deer skeletal elements were collected between 2019 and 2024 from various institutions in northern and southern Europe (Table 1). The taxonomic and, when available, exact, or estimated age-at-death, sex, and provenance information were recorded for each specimen. The skeletal elements and zones to measure were selected based on the presence of measurable features, the likelihood of preservation in the archaeological record (i.e., denser and more robust elements/zones), and the probability of being recovered through hand-collection during excavation (i.e., larger elements). Measurements were taken on the bones of adult and immature individuals. In this study, individuals were considered immature when one or more epiphyses of the element being measured were unfused. The analysis of specific zones focused on specimens for which all epiphyses of that zone were fused or fusing; for selected zones, immature specimens were analysed in separate graphs. The measurements chosen to develop the biometric method include standard ones, as well as new ones created to translate morphological traits, some of which had been previously identified as diagnostic (Di Stefano, 1995 ; Lister, 1996 ) (Table 2; Open Resource 1). All measurements were taken using 15-cm Mitutoyo digital callipers with a precision to the tenth of a millimetre, with the exception of long bone greatest lengths for which an osteometric board was used, with a precision to the millimetre. Raw data for all selected elements are available in Mendeley Data ( https://data.mendeley.com/datasets/vn43dpvbzr/1 ), allowing researchers to reproduce the graphs and plot their own archaeological values against modern ranges. Absolute measurement values and relative values (i.e., shape indices) were grouped and analysed by species (three variables) and age (synthesized in two variables, adult and immature, as described above), without any further separation based on sex or geographical provenance. Sex and spatio-temporal differences are significant in defining potential size ranges (e.g., Sykes et al., 2013 ; Karastoyanova et al., 2020 ), but do not impact on the distribution of shape index values, and therefore have not been used in this presentation of the method but are discussed in a separated publication on European cervid morphometry (in prep.). The assessment of distribution and degree of separation of values from the three species primarily relied on data visualisation using scatterplots, often combining size ranges and shape indices (e.g., distal tibia: Bd vs Ddb/Dda); this approach aims to optimise the separation potentials of dimensional and morphological differences. For most zones, additional scatter plots with shape indices only are also presented. The results are summarized by presenting the shape index value means for each element, displayed in a linear graph, and expressed as LSI values using the red deer means as the standard; the values of each shape index are also displayed in notched box plots, describing and comparing the value distributions of the three species (Open Resource 3). The groups of shape index values were statistically tested using the software R. In detail, the statistical significance of differences between shape index values were calculated for comparisons of fallow deer with red deer and roe deer; the Mann-Whitney U test of significance was used. The resulting p values are reported in tables; for each element, the p values are also displayed graphically (Open Resource 4), providing a complementary visual summary. The Principle Component Analysis (PCA) was run for specific element zones using shape indices (Table 3), to provide an additional assessment of morphological differences among red deer, fallow deer, and roe deer. The exclusion of absolute size data from PCA analyses ensured that these latter focused on morphology rather than on the more straightforward, but often insufficient, separation potential offered by size. PCA is an unsupervised data reduction technique, capturing the maximum variance without using class labels in the computation. For each zone, it reduced all shape indices to two dimensions (PC1, PC2), that provided the best separation of the three cervid species. Multivariate outliers were identified and removed using the Mahalanobis distance method with a threshold of 0.975 (i.e., p < 0.025); this method measures data points' distance from the multivariate centre while accounting for correlations between variables. The inter- and intra-observer errors were estimated in order to assess the reliability of use of each measurement, some of which are new or adapted, as well as the repeatability of results. The recording protocol was shared with seven zooarchaeologists from the Sheffield Zooarchaeology Laboratory (UK) with different levels of expertise in biometric analyses; therefore, the total number of researchers involved, including the authors, was nine. Each researcher took the measurements listed in Table 2 on modern adult skeletons of red deer (ID: 1205), fallow deer (ID: 1207), and roe deer (ID: 1584) from the laboratory’s reference collection. To evaluate the consistency in data recording among observers, the Intraclass Correlation Coefficient (ICC) was employed, a standard statistical method for assessing reproducibility and inter-observer agreement in quantitative studies (Koo & My, 2016). For the assessment of intra-observer reliability, the authors measured the same three specimens repeatedly across three days at different times of the day. The inter- and intra-observer error results are reported and discussed in Open Resource 2. In general, very high ICC values were obtained, indicating an excellent inter-observer and intra-observer reliability for most measurements. Results Scapula The size ranges of the greatest length of the processus articularis (GLP; Fig 4, graph A) show partial overlap between fallow deer and red deer, particularly in the 42-48 mm range. This overlap indicates that, in this case, size alone can be insufficient for reliable species identification. By contrast, roe deer forms a distinct cluster of smaller values, clearly separated from those of fallow deer and red deer. The shape index ASG/GLP offers good morphological separation of fallow and red deer: fallow deer exhibits lower values, suggesting that this species’ scapulae have a proportionally shorter distance between the base of the spine and the edge of the glenoid cavity compared to red deer and roe deer. In fallow deer and red deer, there also seems to exist a negative allometry whereby larger specimens tend to have lower index values; such allometries follow different regression lines, favouring separation of the two species. In graph B, the same index is plotted against ASG/LG, resulting in a partial isolation of fallow deer values in the bottom-left part of the graph. In graph C, the smallest length of the collum scapulae (SLC) is plotted against its ratio to GLP. Fallow deer and red deer distributions show a positive allometry, a consequence of the fact that SLC size increase with age is greater than in other measurements; however, the former species presents lower SLC/GLP values, suggesting that the collum scapulae of fallow deer is proportionally narrower relative to the length of the processus articularis . The shape index value means in graph E summarise well the observations made above, highlighting the extent to which different indices allow to isolate fallow deer specimens; the results of the Mann-Whitney U tests fully support such separation potentials (Table 3). Also the results of the PCA confirm the good separation provided by biometrical analyses for fallow deer specimens (Open Resource 5: Fig 1). Humerus In Fig 5, graph A highlights some degree of overlap of Bd fallow deer and red deer values; HX/Bd values, however, tend to be lower in fallow deer, ensuring almost complete separation of the two clusters. Such lower values compared to red deer may reflect the separation criterion presented in (Di Stefano, 1995: Fig 5 therein), where it is suggested that, in fallow deer, the trochlea is directed laterally with a more acute angle. There is also a clear negative allometric relation between size and the shape index, at least in fallow deer and red deer, although such allometries rely on different regression lines. In graph B, two other shape indices (HTL/BT and HTL/Bd) separate well fallow deer and roe deer, as the two species present respectively the lowest and highest values in both indices; red deer values lie in between, therefore partly overlapping with fallow deer values. As both indices involve the diameter of the capitulum (HTL), it seems that this latter is, on average, proportionally smaller in fallow deer. The same explanation may apply to the lower HTL/HTC index values of fallow deer in graph C, which allow to overcome the only slight HTC size overlap with red deer. Two other shape indices employing distal end height measurement values (HTC/HX and HTL/HX) provide mild separation of fallow deer values from red deer and roe deer ones (graph D). Shape indices combining shaft and distal end measurements (graph E, SD/HTC vs SD1/HTC), and the combined plotting of shape indices of the shaft and distal widths (graph F, SD1/SD vs BT/Bd) provide the best morphological separations, although in the latter graph this applies to the separation of fallow deer and red deer specimens only. The smallest breadth of the shaft (SD) highlights a significant size overlap between fallow deer and red deer specimens but the fallow deer shape index (SD1/SD) values plot lower (graph G), indicating that this species’ humerus shaft is more tubular while the red deer’s is more projecting antero-posteriorly. To some extent, this characteristic applies to immature specimens as well (graph H), where size overlap is substantial; however, there seem to be a positive allometry along similar regression lines in fallow deer and red deer, which may compromise the effectiveness of morphological separation. In mature specimens, the greatest length from the caput (GLC) values displays substantial dimensional overlap between fallow deer and roe deer and, to a smaller extent, between fallow deer and red deer (graph I). Both graphs I and J highlight the greater slenderness of roe deer humeri, whose shape index values plot on the bottom-left area of the graphs. Fallow deer and red deer shape index values are also well separated; in graph I, this is probably the result of the capitulum being proportionally smaller in fallow deer (as seen in graphs B, C), while in graph J such separation reflects the different proportion between shaft measurements (as seen in graphs F-H). The shape index value means (graph K) broadly support the observations made for graphs A-J. The overall slenderness shape indices see the fallow deer closer to the red deer, as do many of the distal end indices though with useful exceptions. Shaft morphology, on the other hand, differentiates to a good degree red deer from the two smaller species, as do most shape indices combining shaft and distal end measurements. The results of the Mann-Whitney U tests for the shape index measurements support the observations above (Table 4). The results of the PCA reflect well both the partial separation of fallow deer and red deer specimens, and the greater isolation of roe deer (Open Resource 5: Fig 2). Radius The value range of Bp indicates some dimensional overlap between fallow deer and red deer values, while roe deer values separate well from fallow deer’s (Fig 6, graph A). The BFp/Bp shape index values of fallow deer are on average lower compared to red deer’s, reflecting a more pronounced lateral bicipital tuberosity in the former species and ensuring complete separation of the two clusters; in addition, biometrical data from both species reveal a negative allometric relation between size and the shape index values, although following different regression lines and thus favouring species separation. BFp values also show some dimensional overlap between the two larger species but the shape index V/BFp produces lower values for the fallow deer, as the insertion for the lateral ulnar articulation is often less pronounced in this species (graph B) – possibly a reflection of character 6 from (Lister, 1996: Fig 2, Part 4 therein). The shape indices employing the proximal depth and widths present substantial overlaps, but fallow deer values run partly parallel to red deer’s and roe deer’s (graph C). The proportions between the smallest depth of the shaft (located close to the proximal end) and the proximal widths display a wider range of values for the red deer relative to the fallow deer, that does not seem driven only by differences in sample size, while they provide some separation between the two smaller species (graph D). Similarly, the shaft shape index (SD/SD1) does not separate fallow deer and red deer, which also present substantial size overlap, but provides some mild separation between fallow and roe deer (graph E). In mature specimens (both ends’ epiphyses fused or fusing), the robustness index SD/GL efficiently describes the greater slenderness of roe deer radii, separating them well from the cluster of fallow deer values despite a substantial size overlap; fallow deer and red deer values are well separated by size as well as by positive allometries along different regression lines (graph F). The shape index value means (graph G) broadly support the observations made for graphs A-F. Most indices highlight substantial morphological similarities between fallow deer and red deer, with the important exceptions of all indices involving measurement V and, to a lesser extent, of BFp/Bp. Roe deer, on the other hand, separates well from the fallow deer, displaying a greater overall slenderness and different average values for most proximal end and shaft indices. The results of the Mann-Whitney U tests for the shape index measurements support the observations above (Table 5). The results of the PCA highlight the partial separation of fallow deer from red and roe deer, as well as the efficiency of shape index using measurement V in separating fallow and red deer (Open Resource 5: Figs 3, 4). Metacarpus Biometric data show no substantial dimensional overlap in the absolute values of measurement a among the three species, only fallow deer and red deer values slightly overlapping in the range 14-16 mm; the shape index a/2 maximises the separation of the three clusters as the fallow deer index presents higher values (Fig 7, graph A). In both the fallow deer and the red deer, there seems to be a positive allometric relation between size and the shape index used, although along different regression lines. Two shape indices from the medial condyle, a/3 and 3/2 (graph B), also perform well in separating the three species: the resulting distribution assumes a clover-shaped pattern, with only partial overlap of red deer and fallow deer values, thus reinforcing the diagnostic value of the medial condyle’s biometrical proportions. In graph C, measurement b shows a similarly low degree of dimensional overlap between fallow deer and red deer, while the shape index b/5 enhances cluster separation, particularly between the two larger species; similar allometric relations to those in graph A are observed. Despite a slightly higher degree of overlap between fallow and red deer, the b/6 and 6/5 shape indices still provide good separation of the three taxa (graph D). Overall, metacarpus distal widths in fallow deer are proportionally less reduced relative to depths than in red deer and roe deer. In addition, the roe deer exhibits more pronounced proportional differences between the antero-posterior diameters of the verticillus and of the internal trochlea compared to the two larger species. Similar results were also obtained for the metatarsus. Proportions between condylar widths and the total width of the distal articular facet are assessed in graph E. Despite some overlap, fallow deer values tend to cluster in the top-right part of the graph, suggesting the condyles of this species are less spaced. Shape indices related to shaft morphology, particularly SD1/BFd and SD1/SD, indicate that fallow deer (and roe deer) has a proportionally flatter shaft compared to red deer (graph F). This pattern is consistent with findings from other postcranial elements, pointing to a characteristically flatter shaft profile in fallow deer. When the smallest breadth of the shaft (SD) is plotted against the shaft index SD1/SD (graph G), size overlap between fallow and red deer is substantial, but the two species are well separated by the combination of size and shape analyses. Similar observations can be made for immature specimens (graph H), where fallow deer continues to display lower shaft index values. In mature specimens, greatest length (GL) values for the metacarpus (graph I) show almost no dimensional overlap between fallow and red deer, and a much greater degree of overlap between roe deer and fallow deer; however, shape indices incorporating GL - such as SD/GL, SD1/GL, and BFd/GL (graphs I and J) - highlight the consistently greater slenderness of roe deer metacarpi. In addition, graph I indicates a negative allometric relation between roe deer metacarpus length and slenderness. In graph J, also the fallow deer separates relatively well from the red deer. The mean values of shape indices across species (graph K) summarise well the trends observed in the scatter plots. In detail, indices involving shaft measurements highlight the greater flatness of fallow deer bones, as do those describing condylar morphology, while roe deer bones are more slender. The results of the Mann-Whitney U tests support the separation potentials of most indices (Table 6). Also the results of the PCA confirm the excellent degree of separation of red deer, fallow deer, and roe deer specimens (Open Resource 5: Fig 5). Tibia The size values of the distal breadth of the tibia (Fig 8, graph A) display limited overlaps between fallow deer and red deer in the range 36-38 mm. The shape index Ddb/Dda (i.e., the proportion between the medial and lateral distal depths) separates well the fallow deer from the red and roe deer, although with some overlap between the two larger species. This separation indicates that the difference between the two depths is proportionally greater in fallow deer, although values from the two larger species seem distributed according to a positive allometric relation along similar regression lines, potentially compromising the diagnostic potentials of this shape index. Differences in the proportions of the tibia distal end measurement values are also visible in the shape indices of graph B, where both the proportions between the distal breadth and the two depths contribute to separate well the fallow deer from the two other cervids. The fallow deer shape index values run parallel to those of red deer and roe deer, indicating that in this species the medial depth is proportionally greater, and the lateral one proportionally smaller, than the distal breadth. In graph C, the smallest breadth of the shaft (SD) displays a much greater degree of size overlap between fallow deer and red deer values, the former species mostly plotting with the smaller third of the latter species. The proportion between the shaft breadth and depth (shape index SD1/SD), however, separates well the fallow deer from the roe and red deer. The shape index values of the first species plot lower, indicating that the fallow deer tibia shaft is consistently ‘flatter’ than that of the other two species; the proportionally larger breadth of the shaft in fallow deer also explains the considerable size overlap with red deer values. In addition, fallow deer and red deer values seem characterised by negative allometric relations along different regression lines. The same shape index also works on immature specimens (graph D), where fallow deer size values can overlap with both red and roe deer ones. Although complete adult long bones are rarely recovered from archaeological contexts, graph E shows that roe deer tibiae are overall much slenderer than those of the other cervids. SD being proportionally larger in fallow deer (graphs C, D) also provides some mild separation between this species and red deer, though with considerable overlap. The shape index value means (graph F), calculated as LSI values using the red deer means as standard, well describe the observations made for graphs A-E. The higher SD and lower SD1 mean values for the fallow deer reflect the flatter shape of the tibia shaft of this species (also visible in the SD1/SD mean value) displayed in graphs C-D. The lower value of fallow deer tibia Ddb (reflected in the lower mean value of Ddb/Dda) highlights the proportionally greater difference in the measurement values of the two distal end depths in this species, as shown in graphs A-B. The relatively high value of the roe deer GL and the low values of the roe deer robustness indices (SD/GL and Bd/GL) reflect the disproportion of length and width measurements displayed in graph E. The results of the Mann-Whitney U tests for the shape index measurements fully support the observations above (Table 7). Also the results of the PCA confirm the good degree of separation provided by biometrical analyses for the three species (Open Resource 5: Fig 6). Astragalus The size and shape index range analyses for the astragalus focus on fully ossified specimens only. The size ranges display some dimensional overlap between fallow deer and red deer values, while roe deer values are, as usual, better detached from those of the larger species. Both the shape indices GLm/GLl and Bd/H provide very little separation of fallow deer and red deer values, the latter plotting slightly lower, while the shape index values of the two smaller species overlap (Fig 9, graphs A, B). Graph B, however, highlights negative allometric relations between size and the values of the shape index of fallow deer and red deer, following different regression lines and thus possibly enhancing species separation potentials. In graph C, two other shape indices display substantial overlap of values from the three species but with the red deer cluster slightly skewed towards the bottom-left of the graph compared to fallow deer’s. The shape index value means (graph D) reflect these slightly different dimensional proportions between red deer specimens and the two other species, whose shape index average values are higher. The results of the Mann-Whitney U tests for the shape index measurements only partly reflect those from data visualization, sometimes producing highly significant differences that do not translate into actual visual separation (Table 8). The results of the PCA, on the other hand, confirm the little efficiency provided by shape index analyses alone in separating astragali from the three cervid species (Open Resource 5: Fig 7). Calcaneum The value ranges of measurements Bd and, to a lesser extent, c highlights some dimensional overlap between fallow deer and red deer, while size overlaps are negligible or absent, in the sample used here, between fallow deer and roe deer (Fig 10, graphs A, B). Combined with the shape indices Bd/Dd and c/d, however, size differences provide a good separation also between the two larger species, the fallow deer index values plotting higher in graph A and running parallel to red deer’s in a bottom-left to top-right direction in graph B, as a consequence of positive allometric relations following different regression lines. The separation potentials of the same shape indices are presented together in graph C. The higher values of the Bd/Dd shape index in fallow deer and roe deer reflect that the dimensions of the calcaneum distal end are proportionally more similar in these species, although the limited extent of such difference and different sizes make this difficult to observe by eye. Similarly, measurement values c and d are proportionally closer in fallow deer and roe deer compared to red deer. The value ranges of two other shape indices from the proximal end display substantial overlap but with the cluster of red deer values slightly skewed towards the top-right of graph D; this reflects different proportions between the lengths (c, d) and width (B) of the proximal end, roe deer’s and especially fallow deer’s being ‘slenderer’. The same shape indices are used to assess the distributions of values from immature specimens (i.e., with unfused distal end); despite the smaller sample sizes, the same observations made for graph D apply to graphs E. The shape index value means (graphs F) is in line with the observations made for graphs A-E, as are the results of the Mann-Whitney U tests (Table 9). The results of the PCA provide a better separation of fallow and red deer than that observed on scatter plots, with the results for immature individuals are similar (Open Resource 5: Figs 8, 9). Metatarsus The sample used in this paper does not highlight any substantial dimensional overlap of measurement a (width of the medial condyle) values among the three cervid species, with shape index a/2 maximizing the separation of the three clusters (Fig 11, graph A). Two other shape indices from the medial condyle (a/3 and 3/2, graph B) separate well the values from the three species, creating a clover-shaped distribution. Measurement b (width of the lateral condyle) values present a greater degree of dimensional overlap but the three shape indices work well in enhancing value cluster separations (graphs C, D), with a slightly greater overlap of fallow deer and red deer values in graph D. The consistency of the evidence from both condyles indicates that, in fallow deer, distal widths are proportionally less small than depths compared to red deer’s, while in roe deer the dimensional differences between the antero-posterior diameters of the verticillus and of the internal trochlea are proportionally greater than in the two larger species. In addition, similarly to the metacarpus, fallow deer and red deer values from both condyles display positive allometric relations between size and shape index, following different regression lines and thus favouring species separation (graphs A, C). Graph E plots the proportions between the condyle widths and the total width of the distal articular facet; despite substantial overlaps, the cluster of fallow deer values is slightly skewed to the top-right of the graph, suggesting that, in this species, the two condyles are proportionally closer to each other. In graph F, two other shape indices show that the smallest depth of the diaphysis (SD1) is proportionally smaller in fallow deer. The shaft index (SD1/SD) is also used, along with SD, in graph G: size overlap between fallow deer and red deer is significant but the value clusters from these two species remain well separated through the combination with the shape index; this latter suggests that, as in other elements, fallow deer metatarsus shafts are ‘flatter’ than in red deer and roe deer. Such characteristic is also detectable in immature specimens, despite some degree of overlap with red deer outlier values (graph H). In mature specimens, the metatarsus greatest length (GL) values show substantial dimensional overlap between fallow deer and roe deer, and minimal size overlap between the two larger species (graph I). However, all GL shape indices (graphs I, J) highlight the usual greater slenderness of roe deer specimens, while fallow deer and red deer values present positive allometric relations along different regression lines (graph I). In fallow deer, the smaller size of the shaft smallest depth (SD1), proportionally to all other measurements (as well as, to a lesser extent, the proportionally smaller BFd), shows up in graph J, with fallow deer values constrained in the bottom-left part of red deer value distribution range. The shape index value means (graph L) broadly support the observations made for graphs A-J. The fallow deer lower SD1 mean value compared to SD’s, translated in a lower SD1/SD index mean value in both mature and immature specimens, reflects the flatter shape of the metatarsus shaft of this species, as displayed in graphs F-H. The different dimensional proportions of distal widths and depths in the three cervid species is visible in the fallow deer and roe deer higher LSI mean values of some shape indices from both condyles. The relatively high value of the roe deer GL mean and the low values of all related robustness indices (SD/GL, SD1/GL, and Bd/GL) reflect the greater slenderness of this species’ skeletal elements. The results of the Mann-Whitney U tests for the shape index measurements fully support the observations above (Table 10). Also the results of the PCA confirm the excellent degree of separation of red deer, fallow deer, and roe deer specimens (Open Resource 5: Fig 10). Discussion The analysis of size and shape indices highlights a variable degree of success in the separation of red deer, fallow deer, and roe deer values from post-cranial elements. In mature specimens, size often provides good separation of the three species, from complete separation to mild overlaps. The use of biometrical data from populations of northern, central, and southern Europe, including Great Britain and part of the Italian peninsula, where red deer is smaller, allows to display a varying set of size ranges. As suggested in previous sections, however, such dimensional ranges are far from complete. The existence of smaller red deer than the specimens used here is of special concern when the proposed methodology aims to separate this species from the fallow deer, a slightly smaller animal. Iberian, southern Italian, south-eastern European, and north African red deer individuals, on average among the smallest representatives from continental mainlands (Mattioli & Ferretti, 2014 ; Becciolini et al., 2016 ), are missing in the dataset; similarly, specimens from extant populations inhabiting medium- and small-sized islands, where red deer underwent varying and often complex processes of size decrease (Vigne, 1988 ; Mulville, 2010 ), have not been sampled. In addition, red deer being an especially successful species due to its adaptability, its size fluctuates substantially to fit climatic change, geography, changes in habitat and dietary resources, and hunting pressure, both on long (Lister, 1984 ; Di Stefano et al., 2015 ) and much shorter chronological scales (Mattioli & Ferretti, 2014 ; Mitchell et al., 1977 ). In sum, wider ranges of red deer bone metric values than those displayed in this study should be born in mind, especially regarding the existence (past and present) of smaller red deer. As a consequence, a greater degree of bone dimensional overlaps must be considered in graphs that combine size ranges and shape index values. Nevertheless, the dimensional analyses presented here can at least suggest with which elements and measurements size overlaps are more likely to occur (e.g., the scapula SLC and GLP, the shaft measurements of all elements, in addition to the humerus and calcaneum Bd, the radius Bp, and the metapodials b), and which ones can instead be used more confidently to assist in taxonomic identification (e.g., the astragalus Bd and the metatarsus a). Shape indices present a varying degree of success in the taxonomic separation of the three species. When separation is provided, by one index or by a combination of two indices, this can either be in terms of isolation of the fallow deer cluster, when its values plot differently from both red deer’s and roe deer’s (e.g., the scapula ASG/GLP, the combination of humerus indices in graph E, the metapodials shape indices from both condyles, the tibia Ddb/Dda and SD1/SD), or in terms of isolation of one of the other species (e.g., the combination of humerus indices in graph F, the combination of metatarsus indices in graphs E and F). In the latter case, other measurements and shape indices can be plotted, if available, to facilitate or confirm species-level identification. An example is provided by the distal metatarsus, where, in addition to size, two shape indices per condyle allow to separate the three cervids: the width-depth proportions to separate fallow and red deer (a/3 and b/6), and the proportion of two depths for separating fallow and roe deer (3/2 and 6/5) (Fig. 11, graphs B, D). The different distributions produced by some shape indices reveal subtle morphological differences between species, which had not been previously observed; this may be a reflection of such differences being too slight to be detected by the eye, and/or being more easily missed due to the co-occurrence of size discrepancies. For example, one morphological trait that seems to characterise many fallow deer elements and zones is a distinct proportion between widths and depths, in this species often more in favour of widths: in most elements, the shaft shape index employing the smallest width and depth (SD1/SD) is lower in fallow deer, indicating that its shafts are ‘flatter’ in comparison to red deer’s and roe deer’s; similarly, in the metatarsus distal condyles of fallow deer, widths are proportionally less smaller than depths compared to red deer’s (Fig. 11, graphs B, D). A more visible morphological trait is the greater depth of the insertion for the lateral ulnar articulation for the radius in red deer (Fig. 6, graph B; probably a reflection of character 6 from Lister ( 1996 : Fig. 2, Part 4 therein)), which prompted the introduction and analysis of measurement V, or, in the same element and zone, the slightly less pronounced lateral bicipital tuberosity of red deer (Fig. 6, graph A). On the other hand, the biometrical translation of the diagnostic character from (Di Stefano, 1995 : Fig. 5 therein) on the distal humerus (whereby in fallow deer the trochlea is directed laterally with a more acute angle), achieved by taking multiple height and width measurements, has not been particularly successful, only one index (HX/Bd) possibly reflecting such character (Fig. 5, graph A). Complete bone robusticity indices involving greatest length measurements consistently isolate the roe deer, whose long bones appear far slenderer than the other two species’. Importantly for the aims of this study, shape index values from immature individuals (long bone shafts, proximal calcaneum) often follow similar distribution patterns to those observed in mature animals; as dimensional separation is more difficult with juvenile specimens, and the degree of ossification alone cannot be used to estimate adult size, these results may prove particularly relevant in the taxonomic identification of remains from younger cervids. In addition, the effectiveness of shaft shape indices (SD1/SD) in mature and immature individuals alike is also very important, as it suggests that it may now be possible to attempt species-level identification of fragmented cervid bones missing either or both ends (where diagnostic morphological criteria are described). In many instances, graphs combining size and shape index values reveal allometric relations between them. These are important to consider, as they contribute to assess the reliability and potentials of species-level identification. Using the cases reported above, for example, the ‘flatness’ of the tibia shaft (Fig. 8, graphs C, D), as well as the metatarsus’ (Fig. 11, graph G), increases (i.e., SD1/SD values become lower) with size in both fallow deer and red deer, although following different regression curves; this increased flatness is also observed for the distal metatarsus condyles described above, again with red and fallow deer values distributed along distinct curves (Fig. 11, graphs A, C). Similarly, the lateral angle created by the trochlea of the humerus, generally more acute in fallow deer than in red deer (Di Stefano, 1995 : Fig. 5 therein), becomes increasingly acuter with increased size in both species (Fig. 5, graph A), while the lateral bicipital tuberosity of the proximal radius becomes more pronounced (Fig. 6, graph A). These trends are the result of width measurements presenting a positive allometry (i.e., growing faster than other measurements). Other allometries are visible in the astragalus (Fig. 9, graph B) and the calcaneum (Fig. 10, graph B), in this case also involving the roe deer. The existence of allometries following different regression lines is especially useful in this study, as size and shape index overlaps are partly overcome by the different distributions, along similar ranges, of values from different species. The limited success of the c/d shape index of the calcaneum in taxonomic separation, for example, is overturned when the index is plotted against an absolute measurement, as its values increase with increased size along different regression lines for fallow deer and red deer. On the one hand, these trends highlight the importance of combining size and shape index values to maximise taxonomic separation; on the other, they warn against the acritical use of shape indices, as different shape index value distributions for differently-sized species may also be the result of allometric relations along similar regression curves: this could be the case of the tibia Ddb/Dda index values for the fallow and red deer, although in this case the regression trends remain unclear (Fig. 8, graph A). In this regard, it is also worth noting that, while the little efficiency of the astragalus’ shape indices is partly overturned by allometric relations following different regression lines (Fig. 9, graph B), the PCA of shape indices produced three largely overlapping clusters: despite the optimization of taxonomic separation offered by more sophisticated data processing, therefore, the use of the PCA turns out, in this case, to be less efficient that a critical combination of size and shape analyses in scatter plots. In sum, the results of the biometrical analyses highlight that: The size ranges for the three species produce different degrees of dimensional overlaps for different elements and measurements, roe deer size values often separating from fallow deer’s better than red deer’s; The selected shape indices separate the values from the three species with varying degrees of success, alone or in combination with size or other shape indices. Such distinct distributions are a reflection of both previously undetected and published morphological differences; Some shape indices remain effective when applied to the taxonomic separation of immature individuals, for which size overlaps are substantial and prevent identification; Shaft shape indices are often effective, extending species-level identification attempts to fragmented materials with no published diagnostic morphological traits; The value distribution of many shape indices, plotted against size, reveal the existence of allometric relations between them. In many instances, such allometries rely on different regression lines for different species, amplifying the taxonomic separation potentials of combined shape and size analyses. The results of PCA analysis of shape indices often reflect and sometimes reinforce the taxonomic separations provided by the comparisons of size and shape index value ranges presented in scatter plots, linear graphs, and box plots; however, it should be used alongside these latter analyses, to ensure a critical use of both size and shape data. Conclusions The biometrical analyses presented in this paper rely on data from nine modern reference collections and different European regions. They describe dimensional and morphological criteria that can be used to assist species-level identification of archaeological cervid bones. This is especially important for the separation of fallow deer remains; despite its known historical and ecological significance, indeed, this relatively rarer species has often been overlooked and sometimes its bones misidentified in European zooarchaeological research, leading to limited visibility in the archaeological record. Such underestimation biases our understanding of past human-cervid-environment interactions, calling for more accurate and objective approaches such as the one offered by this biometrical method. The obtained results show that size remains important evidence to use with caution, while the clustering of shape index values separately for the three species, despite the disparate origins of the modern skeletons, highlights the universal validity of the method when applied to materials from different regions and time periods. In addition, the many allometric relations between size and shape index values seem to support the existence of broadly similar morphometric characters; the often-different regression lines followed by such relations in each species prove useful in species-level separation. Only two published morphological diagnostic criteria (Di Stefano, 1995 : Fig. 5 therein; Lister, 1996 : Fig. 2, Part 4, character 6 therein) were translated biometrically; all other shape indices reflect more subtle differences and can be considered ‘new’ evidence. This characterises the method proposed here as an independent, complementary tool to use in addition to others (use of appropriate reference collections, assessment of morphological criteria). Equally important is the fact that the method is grounded in what are now frequently referred to - often with a somewhat dismissive connotation - as ‘traditional analyses’, as it consists of straightforward biometry. In a research landscape increasingly spellbound by techniques such as ZooMS, ancient DNA, and isotopic analyses, this study offers a timely and valuable re-evaluation of foundational methods. While scientifically advanced techniques undoubtedly expand the analytical potentials of faunal research, they often require substantial financial investment, specialised equipment and staff, and access to dedicated laboratory infrastructures - resources that are not always readily available to researchers. In addition, the (justified) sense of novelty conveyed when these techniques start being applied to certain materials, animal species, regions or periods, as well as the greater consideration implied by higher resource investments, may often lead to neglecting a proper integration of results with other lines of zooarchaeological evidence and within the broader archaeological and historical context. It has now become important to remind and value the primary role of methodologically sound and cost-effective ‘traditional analyses’, firmly grounded in archaeological reasoning and accessible to all. Declarations Author Contribution V.A.: Conceptualisation, Methodology, Formal analysis, Data Curation, Data processing, Writing - Original Draft, Review and Editing, Visualisation, Funding acquisition.M.R.: Conceptualisation, Methodology, Formal analysis, Data Curation, Data processing, Writing - Original Draft, Review and Editing, Visualisation, Funding acquisition.F.G.: Data curation, Data processing, Visualisation. Acknowledgement The authors are grateful to the institutions they visited for data collection: the former Department of Archaeology of the University of Sheffield (UK), Historic England (Fort Cumberland, UK), the Department of Natural History of the University-Museum of Bergen (Norway), the Natural History Museum (Austria), the Hungarian Natural History Museum (Budapest, Hungary), the Malcolm H. Wiener Laboratory for Archaeological Science of the American School of Classical Studies at Athens (Greece), the Department of Physical Sciences, Earth and Environment of the University of Siena (Italy), and the Natural History Museum of Venice (Italy). The data collection trip to Historic England also allowed to measure the modern deer from the private reference collection of Sheila Hamilton Dyer’s (Southampton, UK), who kindly made her specimens available. The authors are thankful to the many colleagues and friends who provided useful suggestions and comments: Umberto Albarella, Polydora Baker, Simon Davis, Kostantina Saliaris, and László Bartosiewicz. We would also like to thank (in alphabetical order): Umberto Albarella, Stephanie Baron, Marina Chorro-Giner, Steve Hollings, Phoebe Liu, Maeve Moorcroft, Mya Pearce-Ahdoud, Matt Rayner and Lenny Salvagno for their contribution in the estimation of the inter-observer error. A heartfelt thank you to Bjørn Reidar Olsson for providing alive deer photographs. Data Availability CERVIFIED Database is available in Mendeley Data: https://data.mendeley.com/datasets/vn43dpvbzr/1doi: 10.17632/vn43dpvbzr.1 References Albarella, U., & Aniceti, V. (2024). 'Unmaking' the deer in medieval Europe: Historical and archaeological evidence. European Journal of Archaeology, 27 (3), 1-19. https://doi.org/10.1017/eaa.2024.11 Albarella, U., & Davis, S. (1996). Mammals and birds from Launceston Castle, Cornwall: Decline in status and the rise of agriculture. Circaea, 12 (1), 1-156. Almeida, N. J., Guinot, C., Ribeiro, I., Barreira, J., & Basílio, A. C. (2024). Human-deer relations during Late Prehistory: The zooarchaeological data from central and southern Portugal in perspective. Animals, 14 (10), 1424. https://doi.org/10.3390/ani14101424 Aniceti, V., Albarella, U., & Sebastiani, A. (2021). The faunal assemblage from the manufacturing district at Spolverino: From the Roman Imperial period to Late Antiquity. Archaeofauna, 30 , 167-183. https://doi.org/10.15366/archaeofauna2021.30.009 Apollonio, M., & Chirichella, R. (2023). Deer and climate change: Impacts and perspectives. Animal Production Science, 63 (16), 1573-1582. https://doi.org/10.1071/AN22296 Baker, K. H., Gray, H. W. I., Lister, A. M., Spassov, N., Welch, A. J., Trantalidou, K., Rose, H., Schroeder, H., & Hoelzel, A. R. (2024b). Ancient and modern DNA track temporal and spatial population dynamics in the European fallow deer since the Eemian interglacial. Scientific Reports, 14 , 3015. https://doi.org/10.1038/s41598-023-48112-6 Baker, K. H., Miller, H., Doherty, S., Gray, H. W. I., Daujat, J., Çakırlar, C., Love, S., Zant, J. M., Cooper, A., Irving-Pease, E. K., Burger, J., Rose, H., Thomas, R., Ross-Piper, J., Bradfield, J., Linderholm, A., Boessenkool, S., Biebach, I., Schroeder, H., & Hoelzel, A. R. (2024a). The 10,000-year biocultural history of fallow deer and its implications for conservation policy. Proceedings of the National Academy of Sciences, 121 (8), e2310051121. https://doi.org/10.1073/pnas.2310051121 Becciolini, V., Bozzi, R., Viliani, M., Biffani, S., & Ponzetta, M. P. (2016). Body measurements from selective hunting: Biometric features of red deer (Cervus elaphus) from Northern Apennine, Italy. Italian Journal of Animal Science, 15 (3), 461-472. https://doi.org/10.1080/1828051X.2016.1186505 Beglane, F., Baker, K. H., Carden, R., Hoelzel, R., Lamb, A., Fhionnghaile, R., Vernesi, C., Mattingly, A., Gooley, R., Chechushkov, I., & Irving-Pease, E. (2018). Ireland's fallow deer: Their historical, archaeological and biomolecular records. Proceedings of the Royal Irish Academy: Archaeology, Culture, History, Literature, 118C , 141-165. https://doi.org/10.3318/priac.2018.118.01 Boessneck, J., Muller, H. H., & Teichert, M. (1964). Osteologische unterscheidungsmerkmale zwischen Schaf (Ovis aries Linne) und Ziege (Capra hircus Linne). Kuhn Archiv, 78 , 1-129. Bowen, F., Carden, R. F., Daujat, J., Grouard, S., Miller, H., Perdikaris, S., & Sykes, N. (2016). Dama dentition: A new tooth eruption and wear method for assessing the age of fallow deer (Dama dama). International Journal of Osteoarchaeology, 26 , 1089-1098. https://doi.org/10.1002/oa.2523 Buckley, M., & Whitcher Kansa, S. (2011). Collagen fingerprinting of archaeological bone and teeth remains from Domuztepe, South Eastern Turkey. Archaeological and Anthropological Sciences, 3 (3), 271-280. https://doi.org/10.1007/s12520-011-0066-z Carden, R. F., McDevitt, A. D., Zachos, F. E., Woodman, P. C., O'Toole, P., Rose, H., Monaghan, N. T., Campana, M. G., Bradley, D. G., & Edwards, C. J. (2012). Phylogenetic, ancient DNA, fossil and morphometric analyses reveal ancient and modern introductions of a large mammal: The complex case of red deer (Cervus elaphus) in Ireland. Quaternary Science Reviews, 42 , 74-84. https://doi.org/10.1016/j.quascirev.2012.02.012 Chapman, D., & Chapman, N. (1975). Fallow deer: Their history, distribution and biology . Terence Dalton Ltd. Davis, S. J. M. (1996). Measurements of a group of adult female Shetland sheep skeletons from a single flock: A baseline for zooarchaeologists. Journal of Archaeological Science, 23 (4), 593-612. https://doi.org/10.1006/jasc.1996.0056 Davis, S. J. M., & MacKinnon, M. (2009). Did the Romans bring fallow deer to Portugal? Environmental Archaeology, 14 , 15-26. https://doi.org/10.1179/174963109X400646 Di Stefano, G. (1995). Identification of fallow deer remains on the basis of its skeletal features: Taxonomical considerations. Bollettino della Società Paleontologica Italiana, 34 (3), 323-331. Di Stefano, G., Pandolfi, L., Petronio, C., & Salari, L. (2015). The morphometry and the occurrence of Cervus elaphus (Mammalia, Cervidae) from the Late Pleistocene of the Italian peninsula. Rivista Italiana di Paleontologia e Stratigrafia, 121 (1), 103-120. https://doi.org/10.13130/2039-4942/6402 Driesch, A. von den. (1976). A guide to the measurement of animal bones from archaeological sites . Peabody Museum Bulletin 1. Peabody Museum of Archaeology and Ethnology. Fernández, H. (2001). Ostéologie comparée des petits ruminants euroasiatiques sauvages et domestiques (genres Rupicapra, Ovis, Capra et Capreolus): Diagnose différentielle du squelette appendiculaire [Doctoral dissertation, University of Geneva]. https://archive-ouverte.unige.ch/unige:77284 Grant, A. (1988). Food, status and religion in England in the Middle Ages: An archaeozoological perspective. Anthropozoologica, Special Issue 2 , 139-146. Holmes, M. (2014). Making a fast buck in the Middle Ages: Evidence for poaching from medieval Wakefield. In K. Baker, R. Carden, & R. Madgwick (Eds.), Deer and people (pp. 200-207). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.20 Karastoyanova, N., Gorczyk, J., & Spassov, N. (2020). The natural history of the fallow deer, Dama dama (Linnaeus, 1758) in Bulgaria in prehistory and new evidence for the existence of an autochthonous Holocene population in the Balkans. International Journal of Osteoarchaeology, 30 (5), 616-628. https://doi.org/10.1002/oa.2886 Koo, T. K., & Li, M. Y. (2016). A guideline of selecting and reporting Intraclass Correlation Coefficients for reliability research. Journal of Chiropractic Medicine, 15 , 155-163. https://doi.org/10.1016/j.jcm.2016.02.012 Lister, A. M. (1984). Evolutionary and ecological origins of British deer. Proceedings of the Royal Society of Edinburgh Section B: Biological Sciences, 82 (4), 205-229. https://doi.org/10.1017/S0269727000003754 Lister, A. M. (1989). Rapid dwarfing of red deer on Jersey in the Last Interglacial. Nature, 342 , 539-542. https://doi.org/10.1038/342539a0 Lister, A. M. (1996). The morphological distinction between bones and teeth of fallow deer (Dama dama) and red deer (Cervus elaphus). International Journal of Osteoarchaeology, 6 (2), 119-143. https://doi.org/10.1002/(SICI)1099-1212(199603)6:23.0.CO;2-8 Marín, J., Saladié, P., Azorit, C., & Rodríguez-Hidalgo, A. (2024). Identification of age at death in red deer (Cervus elaphus) through the upper dentition: Eruption pattern, wear stage and crown heights. Journal of Archaeological Science, 163 , 105934. https://doi.org/10.1016/j.jas.2023.105934 Masseti, M., & Vernesi, C. (2014). Historic zoology of the European fallow deer, Dama dama dama: Evidence from biogeography, archaeology and genetics. In K. Baker, R. Carden, & R. Madgwick (Eds.), Deer and people (pp. 13-22). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.6 Mattioli, S., & Ferretti, F. (2014). Morphometric characterization of Mesola red deer Cervus elaphus italicus (Mammalia: Cervidae). Italian Journal of Zoology, 81 (1), 144-154. https://doi.org/10.1080/11250003.2014.895432 McDevitt, A. D., & Zachos, F. E. (2014). Genetic analyses of natural and anthropogenic movements in deer. In K. Baker, R. Carden, & R. Madgwick (Eds.), Deer and people (pp. 2-12). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.5 McElligott, A. G., Gammell, M. P., Harty, H. C., Paini, D. R., Murphy, D. T., Walsh, J. T., & Hayden, T. J. (2001). Sexual size dimorphism in fallow deer (Dama dama): Do larger, heavier males gain greater mating success? Behavioral Ecology and Sociobiology, 49 , 266-272. https://doi.org/10.1007/s002650000293 Miller, H., & Sykes, N. (2016). Zootherapy in archaeology: The case of the fallow deer (Dama dama dama). Journal of Ethnobiology, 36 (2), 257-276. https://doi.org/10.2993/0278-0771-36.2.257 Milošević-Zlatanović, S., Tomašević Kolarov, N., Vukov, T., & Stamenković, S. (2016). Correlation patterns in roe deer cranium: Sexual dimorphism across different habitats. Journal of Zoology, 300 (4), 291-304. https://doi.org/10.1111/jzo.12383 Mitchell, B., Staines, B. W., & Welch, D. (1977). Ecology of red deer: A research review relevant to their management in Scotland . Institute of Terrestrial Ecology. Mulville, J. (2010). Red deer on Scottish islands. In T. O'Connor & N. Sykes (Eds.), Extinctions and invasions: A social history of British fauna (pp. 43-50). Windgather Press. https://doi.org/10.2307/j.ctv13gvg6k.12 Payne, S. (1969). A metrical distinction between sheep and goat metacarpals. In P. J. Ucko & G. W. Dimbleby (Eds.), The domestication and exploitation of plants and animals (pp. 295-305). Duckworth. Payne, S., & Bull, G. (1988). Components of variation in measurements of pig bones and teeth, and use of measurements to distinguish wild from domestic remains. Archaeozoologia, 2 , 27-66. Post, E., Langvatn, R., Forchhammer, M. C., & Stenseth, N. C. (1999). Environmental variation shapes sexual dimorphism in red deer. Proceedings of the National Academy of Sciences, 96 (8), 4467-4471. https://doi.org/10.1073/pnas.96.8.4467 Salvagno, L., & Albarella, U. (2017). A morphometric system to distinguish sheep and goat postcranial bones. PLoS ONE, 12 (6), e0178543. https://doi.org/10.1371/journal.pone.0178543 Schmidt, K. T. (2014). Supplemental feeding and our attitude towards red deer and natural mortality. In K. Baker, R. Carden, & R. Madgwick (Eds.), Deer and people (pp. 270-279). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.27 Serrano, E., Angibault, J.-M., Cargnelutti, B., & Hewison, A. J. M. (2007). The effect of animal density on metacarpus development in captive fallow deer. Small Ruminant Research, 72 (1), 61-65. https://doi.org/10.1016/j.smallrumres.2006.07.007 Skog, A., Zachos, F. E., Rueness, E. K., Feulner, P. G. D., Mysterud, A., Langvatn, R., Lorenzini, R., Hmwe, S. S., Lehoczky, I., Hartl, G. B., Stenseth, N. C., & Jakobsen, K. S. (2009). Phylogeography of red deer (Cervus elaphus) in Europe. Journal of Biogeography, 36 (1), 66-77. https://doi.org/10.1111/j.1365-2699.2008.01986.x Soderberg, J. (2004). Wild cattle: Red deer in the religious texts, iconography, and archaeology of early medieval Ireland. International Journal of Historical Archaeology, 8 , 167-183. https://doi.org/10.1007/s10761-004-1135-0 Sykes, N., Ayton, G., Bowen, F., Baker, K., Baker, P., Carden, R., Hammon, C., Ingrem, C., Madgwick, R., Miller, H., Poole, K., Rowley-Conwy, P., Worley, F., & Evans, J. (2016). Wild to domestic and back again: The dynamics of fallow deer management in medieval England (c. 11ᵗʰ-16ᵗʰ century AD). Science and Technology of Archaeological Research, 2 (1), 113-126. https://doi.org/10.1080/20548923.2016.1208027 Sykes, N., Carden, R., & Harris, K. (2013). Changes in the size and shape of fallow deer — Evidence for the movement and management of a species. International Journal of Osteoarchaeology, 23 (1), 55-68. https://doi.org/10.1002/oa.1239 Toïgo, C., Gaillard, J.-M., Laere, G., Hewison, A., & Morellet, N. (2006). How does environmental variation influence body mass, body size, and body condition? Roe deer as a case study. Ecography, 29 , 301-308. https://doi.org/10.1111/j.2006.0906-7590.04394.x Vigne, J.-D. (1988). Les mammifères post-glaciaires de Corse: Étude archéozoologique . Gallia Préhistoire Suppl. 26. Éditions du Centre National de la Recherche Scientifique. Tables Tables 1 to 10 are available in the Supplementary Files section. Additional Declarations No competing interests reported. Supplementary Files Table1.docx Table 1. List of visited institutions with numbers of modern complete and partial skeletons measured for each species. * Includes specimens from the UK only. Table2.docx Table 2. List of measurements taken on each element. Italic: measurements also used in analyses focussing specifically on immature elements. +: newly introduced measurement. For visual descriptions of the new measurements see Open Resource 1. Table3.docx Table 3. p values from Mann–Whitney U tests for the shape indices of the scapula. A graphical representation of the results is available in Open Resource 4: Fig 1. Table4.docx Table 4. p values from Mann–Whitney U tests for the shape indices of the humerus. A graphical representation of the results is available in Open Resource 4: Fig 2. Table5.docx Table 5. p values from Mann–Whitney U tests for the shape indices of the radius. A graphical representation of the results is available in Open Resource 4: Fig 3. Table6.docx Table 6. p values from Mann–Whitney U tests for the shape indices of the metacarpus. A graphical representation of the results is available in Open Resource 4: Fig 4. Table7.docx Table 7. p values from Mann–Whitney U tests for the shape indices of the tibia. A graphical representation of the results is available in Open Resource 4: Fig 5. Table8.docx Table 8. p values from Mann-Whitney U tests for the shape indices of the astragalus. A graphical representation of the results is available in Open Resource 4: Fig 6. Table9.docx Table 9. p values from Mann–Whitney U tests for the shape indices of the calcaneum. A graphical representation of the results is available in Open Resource 4: Fig 7. Table10.docx Table 10. p values from Mann–Whitney U tests for the shape indices of the metatarsus. A graphical representation of the results is available in Open Resource 4: Fig 8. OnlineResource1.pdf OnlineResource2.pdf OnlineResource3.pdf OnlineResource4.pdf OnlineResource5.pdf Cite Share Download PDF Status: Published Journal Publication published 16 Jan, 2026 Read the published version in Archaeological and Anthropological Sciences → Version 1 posted Editorial decision: Revision requested 19 Jun, 2025 Reviews received at journal 12 Jun, 2025 Reviews received at journal 04 Jun, 2025 Reviews received at journal 27 May, 2025 Reviewers agreed at journal 16 May, 2025 Reviewers agreed at journal 15 May, 2025 Reviewers agreed at journal 15 May, 2025 Reviewers invited by journal 13 May, 2025 Editor assigned by journal 12 May, 2025 Submission checks completed at journal 12 May, 2025 First submitted to journal 10 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6636241","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":456146458,"identity":"011a2e9c-e45c-4a3c-8a65-0fbac2b16b28","order_by":0,"name":"Veronica Aniceti","email":"data:image/png;base64,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","orcid":"","institution":"Consejo Superior de Investigaciones Científicas, Institució Milà i Fontanals (CSIC-IMF)","correspondingAuthor":true,"prefix":"","firstName":"Veronica","middleName":"","lastName":"Aniceti","suffix":""},{"id":456146459,"identity":"756a53bd-734a-48e1-b69f-9603c024efc7","order_by":1,"name":"Mauro Rizzetto","email":"","orcid":"","institution":"Interdisciplinary Historical Food Studies research group (VUB-FOST)","correspondingAuthor":false,"prefix":"","firstName":"Mauro","middleName":"","lastName":"Rizzetto","suffix":""},{"id":456146460,"identity":"b90d7334-4675-4ed0-b224-38d1feafb414","order_by":2,"name":"Francesco Giacalone","email":"","orcid":"","institution":"Ludwig.guru","correspondingAuthor":false,"prefix":"","firstName":"Francesco","middleName":"","lastName":"Giacalone","suffix":""}],"badges":[],"createdAt":"2025-05-10 17:53:19","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6636241/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6636241/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s12520-025-02380-7","type":"published","date":"2026-01-16T16:29:40+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":82897179,"identity":"3d717f4a-c936-487b-a781-49e41c9a6368","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":6616143,"visible":true,"origin":"","legend":"\u003cp\u003eTwo female red deer (Norway). Photo credit: Bjørn Reidar Olsson.\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/4a4d62e9ef75a71460709d9d.png"},{"id":82897168,"identity":"e9728a24-5900-4a3e-b1a9-0b8e028db0e1","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":7280809,"visible":true,"origin":"","legend":"\u003cp\u003eA male fallow deer (United Kingdom). https://pixabay.com/ (royalty-free stock photos)\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/a6a1265e5b66d2b9e3e15bc8.png"},{"id":82897171,"identity":"3717152e-fb8f-4049-84ac-c830befdcc59","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":8805912,"visible":true,"origin":"","legend":"\u003cp\u003eA male roe deer (Norway). Photo credit: Bjørn Reidar Olsson.\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/a2f1b13dc76ee52e86d8a4eb.png"},{"id":82897176,"identity":"36911351-2ae5-40d2-aef7-e131aa974467","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":68533,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the scapula. A. GLP vs ASG/GLP; B. ASG/LG vs ASG/GLP; C. SLC vs SLC/GLP. The means of shape index values (D) from the three species are expressed as LSI values, using the mean values of red deer measurements and indices as the standard (box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 1).\u003c/p\u003e","description":"","filename":"4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/9d0cdfd36a8bd4453f64865a.jpg"},{"id":82897172,"identity":"0e474916-8376-46b0-afb2-b3a5cd5da7e0","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":386569,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the humerus. For the distal end: A. Bd vs HX/Bd; B. HTL/BT vs HTL/Bd; C. HTC vs HTL/HTC; D. HTC/HX vs HTL/HX. Graphs E (SD/HTC vs SD1/HTC), F (SD1/SD vs BT/Bd) combine distal end and shaft measurements. For the shaft: SD vs SD1/SD for adult (G) and immature (H) specimens. For overall proportions of complete mature humeri: I. GLC vs HTL/GLC; J. SD/GLC vs SD1/GLC. The means of shape index values (K) from the three species are expressed as LSI values, using the mean values of red deer indices as the standard; for the shaft shape index, the means of immature specimens (proximal end unfused, distal end unfused, fusing or fused) are also presented (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 2).\u003c/p\u003e","description":"","filename":"5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/11dca4b7f5cb53d2c42c9a5a.jpg"},{"id":82898415,"identity":"c318340f-af4d-4279-9b3b-9b86e4a8bc63","added_by":"auto","created_at":"2025-05-16 13:10:56","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":218608,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the radius. For the proximal end: A. Bp vs BFp/Bp; B. BFp vs V/BFp; C. Dp/Bp vs Dp/BFp. Graph D (SD1/Bp vs SD1/BFp) combines distal end and shaft measurements. For the shaft: E. SD vs SD1/SD for mature specimens. For overall proportions of complete mature radii: F. GL vs SD/GL. The means of shape index values (G) from the three species are expressed as LSI values, using the mean values of red deer indices as the standard; for the shaft shape index, the means of immature specimens (proximal end unfused, distal end unfused, fusing or fused) are also presented (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 3).\u003c/p\u003e","description":"","filename":"6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/343402f85e9403189d514656.jpg"},{"id":82898436,"identity":"7cd54a09-3d9d-4590-9d62-430e53b5a650","added_by":"auto","created_at":"2025-05-16 13:10:58","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":377930,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the metacarpus. For the distal end: A. a vs a/2; B. a/3 vs 3/2; C. b vs b/5; D. b/6 vs 6/5; E. a/Bd vs b/Bd. Graph F combines distal end and shaft measurements. For the shaft: SD vs SD1/SD for adult (G) and immature (H) specimens. For overall proportions of complete mature metatarsi: GL vs SD/GL (I) and SD1/GL vs Bd/GL (J). The means of shape index values (K) from the three species are expressed as LSI values, using the mean values of red deer measurements and indices as the standard (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 4).\u003c/p\u003e","description":"","filename":"7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/13b00153c14cfda62f4553c5.jpg"},{"id":82897189,"identity":"fe753fcf-306a-4e99-bdda-573b6101633e","added_by":"auto","created_at":"2025-05-16 13:02:57","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":194500,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the tibia. For the distal end: A. Bd vs Ddb/Dda; B. Dda/Bd vs Ddb/Bd. For the shaft: SD vs SD1/SD for adult (C) and immature (D) specimens. For overall proportions of complete mature tibiae: SD/GL vs Bd/GL (E). The means of shape index values (F; \u003cem\u003eSD1/SD\u003c/em\u003erefers to immatures) from the three species are expressed as LSI values, using the mean values of red deer measurements and indices as the standard (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 5).\u003c/p\u003e","description":"","filename":"8.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/207e6a29624ee7cfe9a270b4.jpg"},{"id":82899500,"identity":"4ea28818-2497-426f-92be-e34b0ea22700","added_by":"auto","created_at":"2025-05-16 13:18:58","extension":"jpg","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":109695,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the astragalus. For selected combinations of size ranges and shape indices: A. Bd vs GLm/GLl; B. H vs Bd/H. For two other shape indices: C. Dl/GLl vs Dl/H. The mean of shape index values from the three species are expressed as LSI values in graph D, using the mean values of red deer measurements and indices as the standard (box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 6).\u003c/p\u003e","description":"","filename":"9.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/04c1c1e8680e85aeac4d111d.jpg"},{"id":82898419,"identity":"f707a388-b011-410f-a725-757bd2b92018","added_by":"auto","created_at":"2025-05-16 13:10:56","extension":"jpg","order_by":10,"title":"Figure 10","display":"","copyAsset":false,"role":"figure","size":191407,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the calcaneum. For selected combinations of size ranges and shape indices: A. Bd vs Bd/Dd; B. c vs c/d. The two shape indices of graphs A, B are plotted together in graph C. For two other shape indices: D. B/c vs B/d. For immature specimens: E. B/c vs B/d. The means of shape index values (F) from the three species are expressed as LSI values, using the mean values of red deer measurements and indices as the standard (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 7).\u003c/p\u003e","description":"","filename":"10.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/28d0021d08efc8922b51e2e7.jpg"},{"id":82897207,"identity":"82c89311-ed0e-40f6-bca0-c173e89dccd7","added_by":"auto","created_at":"2025-05-16 13:02:57","extension":"jpg","order_by":11,"title":"Figure 11","display":"","copyAsset":false,"role":"figure","size":405481,"visible":true,"origin":"","legend":"\u003cp\u003eSize and shape index ranges for the metatarsus. For the distal end: A. a vs a/2; B. a/3 vs 3/2; C. b vs b/5; D. b/6 vs 6/5; E. a/Bd vs b/Bd. Graph F combines distal end and shaft measurements. For the shaft: SD vs SD1/SD for adult (G) and immature (H) specimens. For overall proportions of complete mature metatarsi: GL vs SD/GL (I) and SD1/GL vs Bd/GL (J). The means of shape index values (K) from the three species are expressed as LSI values, using the mean values of red deer measurements and indices as the standard (IM.: immature; box plots for the shape indices analysed in this study are available in Open Resource 3: Fig 8).\u003c/p\u003e","description":"","filename":"11.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/8edcbf92832206d8c0a8946c.jpg"},{"id":100614732,"identity":"767cce8e-9426-4397-8931-8bbaeddfdb77","added_by":"auto","created_at":"2026-01-19 17:23:34","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":23362810,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/e293eb23-4574-42e2-8be5-d649376e5585.pdf"},{"id":82898416,"identity":"c54ed1e3-e4f1-4962-a18a-8c34b7244151","added_by":"auto","created_at":"2025-05-16 13:10:56","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":15566,"visible":true,"origin":"","legend":"\u003cp\u003eTable 1. List of visited institutions with numbers of modern complete and partial skeletons measured for each species. * Includes specimens from the UK only.\u003c/p\u003e","description":"","filename":"Table1.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/eccabc0c99eec24dfa642f8d.docx"},{"id":82897169,"identity":"35836302-b67c-4064-b71d-eda26ae1de50","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":20070,"visible":true,"origin":"","legend":"\u003cp\u003eTable 2. List of measurements taken on each element. Italic: measurements also used in analyses focussing specifically on immature elements. +:\u003csup\u003e \u003c/sup\u003enewly introduced measurement. For visual descriptions of the new measurements see Open Resource 1.\u003c/p\u003e","description":"","filename":"Table2.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/384e92538334ba91426d5888.docx"},{"id":82897178,"identity":"9b1e6315-f31e-4e5e-b3ae-7c37680a9a3b","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"docx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":14547,"visible":true,"origin":"","legend":"\u003cp\u003eTable 3. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the scapula. A graphical representation of the results is available in Open Resource 4: Fig 1.\u003c/p\u003e","description":"","filename":"Table3.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/5405246dcbccdb7a6f0095bf.docx"},{"id":82897181,"identity":"4c7e8738-db39-40d6-b8fb-9802bf0773d6","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"docx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":16186,"visible":true,"origin":"","legend":"\u003cp\u003eTable 4. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the humerus. A graphical representation of the results is available in Open Resource 4: Fig 2.\u003c/p\u003e","description":"","filename":"Table4.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/2483f311349b0d49fc3885f8.docx"},{"id":82897180,"identity":"fb43a4d5-f5be-4e39-a247-e49cf5c71f70","added_by":"auto","created_at":"2025-05-16 13:02:56","extension":"docx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":15317,"visible":true,"origin":"","legend":"\u003cp\u003eTable 5. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the radius. A graphical representation of the results is available in Open Resource 4: Fig 3.\u003c/p\u003e","description":"","filename":"Table5.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/14ceb2d6c1476202c71c0e5d.docx"},{"id":82897191,"identity":"f938415b-3307-4091-a47d-e83cc13316e5","added_by":"auto","created_at":"2025-05-16 13:02:57","extension":"docx","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":16152,"visible":true,"origin":"","legend":"\u003cp\u003eTable 6. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the metacarpus. A graphical representation of the results is available in Open Resource 4: Fig 4.\u003c/p\u003e","description":"","filename":"Table6.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/4eb1f1830b737623d2b33bab.docx"},{"id":82897220,"identity":"6a648375-19a7-466e-9ce9-52a078da3ff9","added_by":"auto","created_at":"2025-05-16 13:02:58","extension":"docx","order_by":7,"title":"","display":"","copyAsset":false,"role":"supplement","size":14870,"visible":true,"origin":"","legend":"\u003cp\u003eTable 7. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the tibia. A graphical representation of the results is available in Open Resource 4: Fig 5.\u003c/p\u003e","description":"","filename":"Table7.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/19c1ab47c8e9149ed8e3bb54.docx"},{"id":82898422,"identity":"39eb50fe-ae61-468a-b843-c8018c6ef0b5","added_by":"auto","created_at":"2025-05-16 13:10:57","extension":"docx","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":14672,"visible":true,"origin":"","legend":"\u003cp\u003eTable 8. \u003cem\u003ep\u003c/em\u003e values from Mann-Whitney U tests for the shape indices of the astragalus. A graphical representation of the results is available in Open Resource 4: Fig 6.\u003c/p\u003e","description":"","filename":"Table8.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/fbd62ab029dc82202df626c3.docx"},{"id":82898430,"identity":"62e6fc2c-a9bc-4f51-a888-39bb4308391b","added_by":"auto","created_at":"2025-05-16 13:10:57","extension":"docx","order_by":9,"title":"","display":"","copyAsset":false,"role":"supplement","size":14884,"visible":true,"origin":"","legend":"\u003cp\u003eTable 9. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the calcaneum. A graphical representation of the results is available in Open Resource 4: Fig 7.\u003c/p\u003e","description":"","filename":"Table9.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/d3b5353d8bfa31a64d65de58.docx"},{"id":82897223,"identity":"86eaeb9b-f110-4778-9222-944d3ab4c3dd","added_by":"auto","created_at":"2025-05-16 13:02:58","extension":"docx","order_by":10,"title":"","display":"","copyAsset":false,"role":"supplement","size":16264,"visible":true,"origin":"","legend":"\u003cp\u003eTable 10. \u003cem\u003ep\u003c/em\u003e values from Mann–Whitney U tests for the shape indices of the metatarsus. A graphical representation of the results is available in Open Resource 4: Fig 8.\u003c/p\u003e","description":"","filename":"Table10.docx","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/a1d00eb2b5235a9b4045788b.docx"},{"id":82897212,"identity":"34288cf2-1580-4ae5-8937-1522e4b49922","added_by":"auto","created_at":"2025-05-16 13:02:58","extension":"pdf","order_by":11,"title":"","display":"","copyAsset":false,"role":"supplement","size":744527,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource1.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/5968aa12d0f791f745022237.pdf"},{"id":82898431,"identity":"d4dc02f0-f862-4e09-b4e1-e45c8ecb8d94","added_by":"auto","created_at":"2025-05-16 13:10:58","extension":"pdf","order_by":12,"title":"","display":"","copyAsset":false,"role":"supplement","size":253576,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource2.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/e681dd6b61fd6baf45108cce.pdf"},{"id":82898425,"identity":"f0a6fd3a-4749-4a0d-9a39-99c86e1059d0","added_by":"auto","created_at":"2025-05-16 13:10:57","extension":"pdf","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":5764516,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource3.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/a8437ee809404a078675266d.pdf"},{"id":82898420,"identity":"80acfb78-55f0-4947-8e4f-acf0482032ee","added_by":"auto","created_at":"2025-05-16 13:10:56","extension":"pdf","order_by":14,"title":"","display":"","copyAsset":false,"role":"supplement","size":1328646,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource4.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/2be3439cf5a4076c47b05679.pdf"},{"id":82898421,"identity":"65a40d4e-7514-499f-a05f-44e111f7ce9c","added_by":"auto","created_at":"2025-05-16 13:10:56","extension":"pdf","order_by":15,"title":"","display":"","copyAsset":false,"role":"supplement","size":1416143,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineResource5.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6636241/v1/9e97cb6d21ca192ad7e24f3a.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"\u003cp\u003e‘Cervified’: a method for the morphometric identification of red deer (Cervus elaphus), fallow deer (Dama dama), and roe deer (Capreolus capreolus) archaeological bones\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eRed deer, fallow deer, and roe deer are among the most widespread large-sized wild mammals of Europe and, not surprisingly, they often played an important role in human society. These animals were hunted as a source of food and other materials, whether as part of subsistence or recreational activities, held significant religious and magico-medicinal meaning, and could represent, alive or on the table, a symbol of social differentiation. As such, deer has been the subject of extensive literature (e.g., Chapman \u0026amp; Chapman, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Grant, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Soderberg, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Holmes, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Schmidt, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Miller \u0026amp; Sykes, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Albarella \u0026amp; Aniceti, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Almeida et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the last two decades, zooarchaeological research on this taxon has benefitted from methodological improvements and the application of new analytical techniques, from ageing tools (Bowen et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Mar\u0026iacute;n et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), to isotopic and genetic analyses (Skog et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; McDevitt \u0026amp; Zachos, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Sykes et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Baker et el., 2024a; Baker et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e), to biometry (Carden et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Sykes et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Karastoyanova et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This paper aims to contribute to a fundamental yet often overlooked aspect of cervid archaeological research, functional to many of the studies and techniques mentioned above, namely species-level identification of faunal remains.\u003c/p\u003e \u003cp\u003eIn zooarchaeology, the identification of the remains of red deer (\u003cem\u003eCervus elaphus\u003c/em\u003e), fallow deer (\u003cem\u003eDama dama\u003c/em\u003e), and roe deer (\u003cem\u003eCapreolus capreolus\u003c/em\u003e) from European faunal assemblages has often posed significant challenges (Figs.\u0026nbsp;1\u0026ndash;3). Bone morphological similarities and partial size overlaps, in addition to the fragmented nature of most faunal materials, can prevent species-level identification, limiting the interpretative potentials of the zooarchaeological evidence. The problem of identification of European cervid remains from historical times arises from the introduction and spread of the fallow deer in the Continent since at least Roman times and then, more consistently, during the Middle Ages (Davis \u0026amp; MacKinnon, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Masseti \u0026amp; Vernesi, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Baker et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e; Baker et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e). Indeed, this species partly overlaps in size with the red deer, which is larger, and, more rarely, with the roe deer, a smaller animal. The extent of such size overlaps in past animal populations is, however, often difficult to assess, due to substantial diachronic and spatial size variability. The factors determining deer size ranges and their fluctuations are multiple. Sexual dimorphism is especially strong in red deer and fallow deer (Post et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; McElligott et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Milošević-Zlatanović et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), implying size overlaps of large male fallow deer and small female red deer. Climatic conditions also impact on deer size, cervids being larger in colder regions and periods (To\u0026iuml;go et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Apollonio \u0026amp; Chirichella, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), as do specific geographical contexts such as islands and isolated regions, where these animals are smaller (Lister, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Mulville, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Finally, anthropogenic factors such as hunting pressure, habitat reduction, forced relocation, and emparkment have also shown to lead to a reduction in size (Serrano et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Karastoyanova et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In faunal assemblages, the potential presence of immature individuals, whether identifiable or not as such, leads to even greater size overlaps and can further compromise species-level identification. Currently, the separation of red, fallow, and roe deer archaeological remains is attempted through different and sometimes complementary methodologies:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003eSize analyses\u003c/em\u003e. When large assemblages are available and through comparisons with data from related sites, it is sometimes possible to identify size clusters that can be attributed to different species (Davis \u0026amp; MacKinnon, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Aniceti et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Albarella \u0026amp; Davis \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). Large samples of cervid biometric values are, however, rarely available; in addition, size overlaps between species are frequent, for the reasons mentioned above, limiting the efficiency of species-level identification based on size alone.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003eMorphological criteria\u003c/em\u003e. These have long been established for the distinction between red and fallow deer (Di Stefano, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Lister, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). Their use, however, is inevitably biased by a certain degree of subjectivity and by the level of expertise of different researchers, and ideally requires the assistance of abundant reference material; in addition, not all criteria are equally diagnostic (Lister, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1996\u003c/span\u003e), implying that taxonomic separation is more difficult and less reliable on some zones. More importantly, the results of such attempts cannot be objectively scrutinised by other researchers.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003eAncient DNA\u003c/em\u003e (\u003cem\u003eaDNA\u003c/em\u003e). Genetic analyses can be very effective when enough collagen is preserved. aDNA analyses are not used with the specific purpose of separating fallow deer and red deer remains but rather to investigate genetic structures and past zoogeographical dynamics in these species; in doing so, however, some specimens considered in these studies are shown to belong to the other species and excluded (e.g., Baker et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e; Beglane et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Despite their effectiveness, genetic analyses are nonetheless constrained by severe cost limitations and are destructive. Zooarchaeology by Mass Spectrometry (ZooMS), on the other hand, cannot currently separate between red and fallow deer remains (Buckley \u0026amp; Whitcher Kansa, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In addition, this technique too involves destructive procedures and can be economically impacting when applied to large samples, despite being less time-consuming and more affordable compared to genetic analyses.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eThis study introduces a new method for the separation of fallow deer from red deer and roe deer remains. The method consists in the use of biometric indices and size ranges, derived from modern cervid skeletons of known species, against which indices and raw measurement values from archaeological material can be plotted for comparison. Such approach implies a key set of advantages:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003eIt allows for more objective identification attempts, as these are represented graphically and therefore open to scrutiny;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eThe separation potentials of size and shape are combined to maximize chances of reliable species-level identification;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eIn some skeletal elements, diagnostic indices can also be used on immature individuals, when size alone is of no use, or on zones where no morphological criteria exist (e.g., long bone shafts);\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eLastly, the method relies on the use of the calliper and osteometric board and is therefore widely accessible and inclusive.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eThe new method does not aim to replace the identification methods listed above, nor to dismiss any potential future improvements to such methods or the importance of an appropriate use of reference collections. Rather, it aims to provide additional support to the study of the history of cervids in temperate Europe and the Mediterranean; in phylogenetic studies of fallow deer in a specific period or region, for example, the use of an efficient, objective, and inexpensive biometric method may assist in the identification of a more reliable sample of fallow deer bones on which to focus funds for genetic analyses. This biometric method relies on an extensive dataset which, on the other hand, is not exhaustive of all regions and available modern cervid skeletons; however, the variability in size ranges covered and the consistency in the validity of shape indices across different regions ensure the applicability of the method to other regions, provided critical interpretation of the results is always used. Ultimately, more widespread and correct identification of cervid archaeological and palaeontological remains will contribute to the reconstruction of past biodiversity, human-animal-environment interactions, and socio-cultural and religious practices.\u003c/p\u003e \u003cp\u003eThe method here presented is the first output of the Cervified Project, conducted by the authors to investigate the potentials of biometric approaches in the species-level identification of archaeological European cervid remains, as well as in the study of past and present cervid bone morphometry.\u003c/p\u003e "},{"header":"Materials and methodological approach","content":"\u003cp\u003eBiometric data from modern red, fallow, and roe deer skeletal elements were collected between 2019 and 2024 from various institutions in northern and southern Europe (Table\u0026nbsp;1). The taxonomic and, when available, exact, or estimated age-at-death, sex, and provenance information were recorded for each specimen.\u003c/p\u003e \u003cp\u003eThe skeletal elements and zones to measure were selected based on the presence of measurable features, the likelihood of preservation in the archaeological record (i.e., denser and more robust elements/zones), and the probability of being recovered through hand-collection during excavation (i.e., larger elements). Measurements were taken on the bones of adult and immature individuals. In this study, individuals were considered immature when one or more epiphyses of the element being measured were unfused. The analysis of specific zones focused on specimens for which all epiphyses of that zone were fused or fusing; for selected zones, immature specimens were analysed in separate graphs. The measurements chosen to develop the biometric method include standard ones, as well as new ones created to translate morphological traits, some of which had been previously identified as diagnostic (Di Stefano, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Lister, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) (Table\u0026nbsp;2; Open Resource 1). All measurements were taken using 15-cm Mitutoyo digital callipers with a precision to the tenth of a millimetre, with the exception of long bone greatest lengths for which an osteometric board was used, with a precision to the millimetre. Raw data for all selected elements are available in Mendeley Data (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://data.mendeley.com/datasets/vn43dpvbzr/1\u003c/span\u003e\u003cspan address=\"https://data.mendeley.com/datasets/vn43dpvbzr/1\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), allowing researchers to reproduce the graphs and plot their own archaeological values against modern ranges.\u003c/p\u003e \u003cp\u003eAbsolute measurement values and relative values (i.e., shape indices) were grouped and analysed by species (three variables) and age (synthesized in two variables, adult and immature, as described above), without any further separation based on sex or geographical provenance. Sex and spatio-temporal differences are significant in defining potential size ranges (e.g., Sykes et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Karastoyanova et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), but do not impact on the distribution of shape index values, and therefore have not been used in this presentation of the method but are discussed in a separated publication on European cervid morphometry (in prep.). The assessment of distribution and degree of separation of values from the three species primarily relied on data visualisation using scatterplots, often combining size ranges and shape indices (e.g., distal tibia: Bd \u003cem\u003evs\u003c/em\u003e Ddb/Dda); this approach aims to optimise the separation potentials of dimensional and morphological differences. For most zones, additional scatter plots with shape indices only are also presented. The results are summarized by presenting the shape index value means for each element, displayed in a linear graph, and expressed as LSI values using the red deer means as the standard; the values of each shape index are also displayed in notched box plots, describing and comparing the value distributions of the three species (Open Resource 3).\u003c/p\u003e \u003cp\u003eThe groups of shape index values were statistically tested using the software \u003cem\u003eR.\u003c/em\u003e In detail, the statistical significance of differences between shape index values were calculated for comparisons of fallow deer with red deer and roe deer; the Mann-Whitney U test of significance was used. The resulting \u003cem\u003ep\u003c/em\u003e values are reported in tables; for each element, the \u003cem\u003ep\u003c/em\u003e values are also displayed graphically (Open Resource 4), providing a complementary visual summary.\u003c/p\u003e \u003cp\u003eThe Principle Component Analysis (PCA) was run for specific element zones using shape indices (Table\u0026nbsp;3), to provide an additional assessment of morphological differences among red deer, fallow deer, and roe deer. The exclusion of absolute size data from PCA analyses ensured that these latter focused on morphology rather than on the more straightforward, but often insufficient, separation potential offered by size. PCA is an unsupervised data reduction technique, capturing the maximum variance without using class labels in the computation. For each zone, it reduced all shape indices to two dimensions (PC1, PC2), that provided the best separation of the three cervid species. Multivariate outliers were identified and removed using the Mahalanobis distance method with a threshold of 0.975 (i.e., \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.025); this method measures data points' distance from the multivariate centre while accounting for correlations between variables.\u003c/p\u003e \u003cp\u003eThe inter- and intra-observer errors were estimated in order to assess the reliability of use of each measurement, some of which are new or adapted, as well as the repeatability of results. The recording protocol was shared with seven zooarchaeologists from the Sheffield Zooarchaeology Laboratory (UK) with different levels of expertise in biometric analyses; therefore, the total number of researchers involved, including the authors, was nine. Each researcher took the measurements listed in Table\u0026nbsp;2 on modern adult skeletons of red deer (ID: 1205), fallow deer (ID: 1207), and roe deer (ID: 1584) from the laboratory\u0026rsquo;s reference collection. To evaluate the consistency in data recording among observers, the Intraclass Correlation Coefficient (ICC) was employed, a standard statistical method for assessing reproducibility and inter-observer agreement in quantitative studies (Koo \u0026amp; My, 2016). For the assessment of intra-observer reliability, the authors measured the same three specimens repeatedly across three days at different times of the day. The inter- and intra-observer error results are reported and discussed in Open Resource 2. In general, very high ICC values were obtained, indicating an excellent inter-observer and intra-observer reliability for most measurements.\u003c/p\u003e "},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eScapula\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe size ranges of the greatest length of the \u003cem\u003eprocessus articularis\u003c/em\u003e (GLP; Fig 4, graph A) show partial overlap between fallow deer and red deer, particularly in the 42-48 mm range. This overlap indicates that, in this case, size alone can be insufficient for reliable species identification. By contrast, roe deer forms a distinct cluster of smaller values, clearly separated from those of fallow deer and red deer. The shape index ASG/GLP offers good morphological separation of fallow and red deer: fallow deer exhibits lower values, suggesting that this species\u0026rsquo; scapulae have a proportionally shorter distance between the base of the spine and the edge of the glenoid cavity compared to red deer and roe deer. In fallow deer and red deer, there also seems to exist a negative allometry whereby larger specimens tend to have lower index values; such allometries follow different regression lines, favouring separation of the two species.\u003c/p\u003e\n\u003cp\u003eIn graph B, the same index is plotted against ASG/LG, resulting in a partial isolation of fallow deer values in the bottom-left part of the graph.\u003c/p\u003e\n\u003cp\u003eIn graph C, the smallest length of the collum scapulae (SLC) is plotted against its ratio to GLP. Fallow deer and red deer distributions show a positive allometry, a consequence of the fact that SLC size increase with age is greater than in other measurements; however, the former species presents lower SLC/GLP values, suggesting that the collum scapulae of fallow deer is proportionally narrower relative to the length of the \u003cem\u003eprocessus articularis\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eThe shape index value means in graph E summarise well the observations made above, highlighting the extent to which different indices allow to isolate fallow deer specimens; the results of the Mann-Whitney U tests fully support such separation potentials (Table 3). Also the results of the PCA confirm the good separation provided by biometrical analyses for fallow deer specimens (Open Resource 5: Fig 1).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHumerus\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn Fig 5, graph A highlights some degree of overlap of Bd fallow deer and red deer values; HX/Bd values, however, tend to be lower in fallow deer, ensuring almost complete separation of the two clusters. Such lower values compared to red deer may reflect the separation criterion presented in (Di Stefano, 1995: Fig 5 therein), where it is suggested that, in fallow deer, the trochlea is directed laterally with a more acute angle. There is also a clear negative allometric relation between size and the shape index, at least in fallow deer and red deer, although such allometries rely on different regression lines. In graph B, two other shape indices (HTL/BT and HTL/Bd) separate well fallow deer and roe deer, as the two species present respectively the lowest and highest values in both indices; red deer values lie in between, therefore partly overlapping with fallow deer values. As both indices involve the diameter of the capitulum (HTL), it seems that this latter is, on average, proportionally smaller in fallow deer. The same explanation may apply to the lower HTL/HTC index values of fallow deer in graph C, which allow to overcome the only slight HTC size overlap with red deer. Two other shape indices employing distal end height measurement values (HTC/HX and HTL/HX) provide mild separation of fallow deer values from red deer and roe deer ones (graph D).\u003c/p\u003e\n\u003cp\u003eShape indices combining shaft and distal end measurements (graph E, SD/HTC vs SD1/HTC), and the combined plotting of shape indices of the shaft and distal widths (graph F, SD1/SD vs BT/Bd) provide the best morphological separations, although in the latter graph this applies to the separation of fallow deer and red deer specimens only.\u003c/p\u003e\n\u003cp\u003eThe smallest breadth of the shaft (SD) highlights a significant size overlap between fallow deer and red deer specimens but the fallow deer shape index (SD1/SD) values plot lower (graph G), indicating that this species\u0026rsquo; humerus shaft is more tubular while the red deer\u0026rsquo;s is more projecting antero-posteriorly. To some extent, this characteristic applies to immature specimens as well (graph H), where size overlap is substantial; however, there seem to be a positive allometry along similar regression lines in fallow deer and red deer, which may compromise the effectiveness of morphological separation.\u003c/p\u003e\n\u003cp\u003eIn mature specimens, the greatest length from the caput (GLC) values displays substantial dimensional overlap between fallow deer and roe deer and, to a smaller extent, between fallow deer and red deer (graph I). Both graphs I and J highlight the greater slenderness of roe deer humeri, whose shape index values plot on the bottom-left area of the graphs. Fallow deer and red deer shape index values are also well separated; in graph I, this is probably the result of the capitulum being proportionally smaller in fallow deer (as seen in graphs B, C), while in graph J such separation reflects the different proportion between shaft measurements (as seen in graphs F-H).\u003c/p\u003e\n\u003cp\u003eThe shape index value means (graph K) broadly support the observations made for graphs A-J. The overall slenderness shape indices see the fallow deer closer to the red deer, as do many of the distal end indices though with useful exceptions. Shaft morphology, on the other hand, differentiates to a good degree red deer from the two smaller species, as do most shape indices combining shaft and distal end measurements. The results of the Mann-Whitney U tests for the shape index measurements support the observations above (Table 4). The results of the PCA reflect well both the partial separation of fallow deer and red deer specimens, and the greater isolation of roe deer (Open Resource 5: Fig 2).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eRadius\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe value range of Bp indicates some dimensional overlap between fallow deer and red deer values, while roe deer values separate well from fallow deer\u0026rsquo;s (Fig 6, graph A). The BFp/Bp shape index values of fallow deer are on average lower compared to red deer\u0026rsquo;s, reflecting a more pronounced lateral bicipital tuberosity in the former species and ensuring complete separation of the two clusters; in addition, biometrical data from both species reveal a negative allometric relation between size and the shape index values, although following different regression lines and thus favouring species separation. BFp values also show some dimensional overlap between the two larger species but the shape index V/BFp produces lower values for the fallow deer, as the insertion for the lateral ulnar articulation is often less pronounced in this species (graph B) \u0026ndash; possibly a reflection of character 6 from (Lister, 1996: Fig 2, Part 4 therein). The shape indices employing the proximal depth and widths present substantial overlaps, but fallow deer values run partly parallel to red deer\u0026rsquo;s and roe deer\u0026rsquo;s (graph C).\u003c/p\u003e\n\u003cp\u003eThe proportions between the smallest depth of the shaft (located close to the proximal end) and the proximal widths display a wider range of values for the red deer relative to the fallow deer, that does not seem driven only by differences in sample size, while they provide some separation between the two smaller species (graph D). Similarly, the shaft shape index (SD/SD1) does not separate fallow deer and red deer, which also present substantial size overlap, but provides some mild separation between fallow and roe deer (graph E).\u003c/p\u003e\n\u003cp\u003eIn mature specimens (both ends\u0026rsquo; epiphyses fused or fusing), the robustness index SD/GL efficiently describes the greater slenderness of roe deer radii, separating them well from the cluster of fallow deer values despite a substantial size overlap; fallow deer and red deer values are well separated by size as well as by positive allometries along different regression lines (graph F).\u003c/p\u003e\n\u003cp\u003eThe shape index value means (graph G) broadly support the observations made for graphs A-F. Most indices highlight substantial morphological similarities between fallow deer and red deer, with the important exceptions of all indices involving measurement V and, to a lesser extent, of BFp/Bp. Roe deer, on the other hand, separates well from the fallow deer, displaying a greater overall slenderness and different average values for most proximal end and shaft indices. The results of the Mann-Whitney U tests for the shape index measurements support the observations above (Table 5). The results of the PCA highlight the partial separation of fallow deer from red and roe deer, as well as the efficiency of shape index using measurement V in separating fallow and red deer (Open Resource 5: Figs 3, 4).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMetacarpus\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eBiometric data show no substantial dimensional overlap in the absolute values of measurement a among the three species, only fallow deer and red deer values slightly overlapping in the range 14-16 mm; the shape index a/2 maximises the separation of the three clusters as the fallow deer index presents higher values (Fig 7, graph A). In both the fallow deer and the red deer, there seems to be a positive allometric relation between size and the shape index used, although along different regression lines. Two shape indices from the medial condyle, a/3 and 3/2 (graph B), also perform well in separating the three species: the resulting distribution assumes a clover-shaped pattern, with only partial overlap of red deer and fallow deer values, thus reinforcing the diagnostic value of the medial condyle\u0026rsquo;s biometrical proportions.\u003c/p\u003e\n\u003cp\u003eIn graph C, measurement b shows a similarly low degree of dimensional overlap between fallow deer and red deer, while the shape index b/5 enhances cluster separation, particularly between the two larger species; similar allometric relations to those in graph A are observed. Despite a slightly higher degree of overlap between fallow and red deer, the b/6 and 6/5 shape indices still provide good separation of the three taxa (graph D). Overall, metacarpus distal widths in fallow deer are proportionally less reduced relative to depths than in red deer and roe deer. In addition, the roe deer exhibits more pronounced proportional differences between the antero-posterior diameters of the verticillus and of the internal trochlea compared to the two larger species. Similar results were also obtained for the metatarsus.\u003c/p\u003e\n\u003cp\u003eProportions between condylar widths and the total width of the distal articular facet are assessed in graph E. Despite some overlap, fallow deer values tend to cluster in the top-right part of the graph, suggesting the condyles of this species are less spaced. Shape indices related to shaft morphology, particularly SD1/BFd and SD1/SD, indicate that fallow deer (and roe deer) has a proportionally flatter shaft compared to red deer (graph F). This pattern is consistent with findings from other postcranial elements, pointing to a characteristically flatter shaft profile in fallow deer.\u003c/p\u003e\n\u003cp\u003eWhen the smallest breadth of the shaft (SD) is plotted against the shaft index SD1/SD (graph G), size overlap between fallow and red deer is substantial, but the two species are well separated by the combination of size and shape analyses. Similar observations can be made for immature specimens (graph H), where fallow deer continues to display lower shaft index values.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn mature specimens, greatest length (GL) values for the metacarpus (graph I) show almost no dimensional overlap between fallow and red deer, and a much greater degree of overlap between roe deer and fallow deer; however, shape indices incorporating GL - such as SD/GL, SD1/GL, and BFd/GL (graphs I and J) - highlight the consistently greater slenderness of roe deer metacarpi. In addition, graph I indicates a negative allometric relation between roe deer metacarpus length and slenderness.\u003c/p\u003e\n\u003cp\u003eIn graph J, also the fallow deer separates relatively well from the red deer.\u003c/p\u003e\n\u003cp\u003eThe mean values of shape indices across species (graph K) summarise well the trends observed in the scatter plots. In detail, indices involving shaft measurements highlight the greater flatness of fallow deer bones, as do those describing condylar morphology, while roe deer bones are more slender. The results of the Mann-Whitney U tests support the separation potentials of most indices (Table 6). Also the results of the PCA confirm the excellent degree of separation of red deer, fallow deer, and roe deer specimens (Open Resource 5: Fig 5).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eTibia\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe size values of the distal breadth of the tibia (Fig 8, graph A) display limited overlaps between fallow deer and red deer in the range 36-38 mm. The shape index Ddb/Dda (i.e., the proportion between the medial and lateral distal depths) separates well the fallow deer from the red and roe deer, although with some overlap between the two larger species. This separation indicates that the difference between the two depths is proportionally greater in fallow deer, although values from the two larger species seem distributed according to a positive allometric relation along similar regression lines, potentially compromising the diagnostic potentials of this shape index.\u003c/p\u003e\n\u003cp\u003eDifferences in the proportions of the tibia distal end measurement values are also visible in the shape indices of graph B, where both the proportions between the distal breadth and the two depths contribute to separate well the fallow deer from the two other cervids. The fallow deer shape index values run parallel to those of red deer and roe deer, indicating that in this species the medial depth is proportionally greater, and the lateral one proportionally smaller, than the distal breadth.\u003c/p\u003e\n\u003cp\u003eIn graph C, the smallest breadth of the shaft (SD) displays a much greater degree of size overlap between fallow deer and red deer values, the former species mostly plotting with the smaller third of the latter species. The proportion between the shaft breadth and depth (shape index SD1/SD), however, separates well the fallow deer from the roe and red deer. The shape index values of the first species plot lower, indicating that the fallow deer tibia shaft is consistently \u0026lsquo;flatter\u0026rsquo; than that of the other two species; the proportionally larger breadth of the shaft in fallow deer also explains the considerable size overlap with red deer values. In addition, fallow deer and red deer values seem characterised by negative allometric relations along different regression lines. The same shape index also works on immature specimens (graph D), where fallow deer size values can overlap with both red and roe deer ones.\u003c/p\u003e\n\u003cp\u003eAlthough complete adult long bones are rarely recovered from archaeological contexts, graph E shows that roe deer tibiae are overall much slenderer than those of the other cervids. SD being proportionally larger in fallow deer (graphs C, D) also provides some mild separation between this species and red deer, though with considerable overlap.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe shape index value means (graph F), calculated as LSI values using the red deer means as standard, well describe the observations made for graphs A-E. The higher SD and lower SD1 mean values for the fallow deer reflect the flatter shape of the tibia shaft of this species (also visible in the SD1/SD mean value) displayed in graphs C-D. The lower value of fallow deer tibia Ddb (reflected in the lower mean value of Ddb/Dda) highlights the proportionally greater difference in the measurement values of the two distal end depths in this species, as shown in graphs A-B. The relatively high value of the roe deer GL and the low values of the roe deer robustness indices (SD/GL and Bd/GL) reflect the disproportion of length and width measurements displayed in graph E. The results of the Mann-Whitney U tests for the shape index measurements fully support the observations above (Table 7). Also the results of the PCA confirm the good degree of separation provided by biometrical analyses for the three species (Open Resource 5: Fig 6).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAstragalus\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe size and shape index range analyses for the astragalus focus on fully ossified specimens only. The size ranges display some dimensional overlap between fallow deer and red deer values, while roe deer values are, as usual, better detached from those of the larger species. Both the shape indices GLm/GLl and Bd/H provide very little separation of fallow deer and red deer values, the latter plotting slightly lower, while the shape index values of the two smaller species overlap (Fig 9, graphs A, B). Graph B, however, highlights negative allometric relations between size and the values of the shape index of fallow deer and red deer, following different regression lines and thus possibly enhancing species separation potentials.\u003c/p\u003e\n\u003cp\u003eIn graph C, two other shape indices display substantial overlap of values from the three species but with the red deer cluster slightly skewed towards the bottom-left of the graph compared to fallow deer\u0026rsquo;s. The shape index value means (graph D) reflect these slightly different dimensional proportions between red deer specimens and the two other species, whose shape index average values are higher. The results of the Mann-Whitney U tests for the shape index measurements only partly reflect those from data visualization, sometimes producing highly significant differences that do not translate into actual visual separation (Table 8). The results of the PCA, on the other hand, confirm the little efficiency provided by shape index analyses alone in separating astragali from the three cervid species (Open Resource 5: Fig 7). \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCalcaneum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe value ranges of measurements Bd and, to a lesser extent, c highlights some dimensional overlap between fallow deer and red deer, while size overlaps are negligible or absent, in the sample used here, between fallow deer and roe deer (Fig 10, graphs A, B). Combined with the shape indices Bd/Dd and c/d, however, size differences provide a good separation also between the two larger species, the fallow deer index values plotting higher in graph A and running parallel to red deer\u0026rsquo;s in a bottom-left to top-right direction in graph B, as a consequence of positive allometric relations following different regression lines. The separation potentials of the same shape indices are presented together in graph C. The higher values of the Bd/Dd shape index in fallow deer and roe deer reflect that the dimensions of the calcaneum distal end are proportionally more similar in these species, although the limited extent of such difference and different sizes make this difficult to observe by eye. Similarly, measurement values c and d are proportionally closer in fallow deer and roe deer compared to red deer.\u003c/p\u003e\n\u003cp\u003eThe value ranges of two other shape indices from the proximal end display substantial overlap but with the cluster of red deer values slightly skewed towards the top-right of graph D; this reflects different proportions between the lengths (c, d) and width (B) of the proximal end, roe deer\u0026rsquo;s and especially fallow deer\u0026rsquo;s being \u0026lsquo;slenderer\u0026rsquo;. The same shape indices are used to assess the distributions of values from immature specimens (i.e., with unfused distal end); despite the smaller sample sizes, the same observations made for graph D apply to graphs E. The shape index value means (graphs F) is in line with the observations made for graphs A-E, as are the results of the Mann-Whitney U tests (Table 9). The results of the PCA provide a better separation of fallow and red deer than that observed on scatter plots, with the results for immature individuals are similar (Open Resource 5: Figs 8, 9).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMetatarsus\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe sample used in this paper does not highlight any substantial dimensional overlap of measurement a (width of the medial condyle) values among the three cervid species, with shape index a/2 maximizing the separation of the three clusters (Fig 11, graph A). Two other shape indices from the medial condyle (a/3 and 3/2, graph B) separate well the values from the three species, creating a clover-shaped distribution. Measurement b (width of the lateral condyle) values present a greater degree of dimensional overlap but the three shape indices work well in enhancing value cluster separations (graphs C, D), with a slightly greater overlap of fallow deer and red deer values in graph D. The consistency of the evidence from both condyles indicates that, in fallow deer, distal widths are proportionally less small than depths compared to red deer\u0026rsquo;s, while in roe deer the dimensional differences between the antero-posterior diameters of the verticillus and of the internal trochlea are proportionally greater than in the two larger species. In addition, similarly to the metacarpus, fallow deer and red deer values from both condyles display positive allometric relations between size and shape index, following different regression lines and thus favouring species separation (graphs A, C).\u003c/p\u003e\n\u003cp\u003eGraph E plots the proportions between the condyle widths and the total width of the distal articular facet; despite substantial overlaps, the cluster of fallow deer values is slightly skewed to the top-right of the graph, suggesting that, in this species, the two condyles are proportionally closer to each other. In graph F, two other shape indices show that the smallest depth of the diaphysis (SD1) is proportionally smaller in fallow deer. The shaft index (SD1/SD) is also used, along with SD, in graph G: size overlap between fallow deer and red deer is significant but the value clusters from these two species remain well separated through the combination with the shape index; this latter suggests that, as in other elements, fallow deer metatarsus shafts are \u0026lsquo;flatter\u0026rsquo; than in red deer and roe deer. Such characteristic is also detectable in immature specimens, despite some degree of overlap with red deer outlier values (graph H).\u003c/p\u003e\n\u003cp\u003eIn mature specimens, the metatarsus greatest length (GL) values show substantial dimensional overlap between fallow deer and roe deer, and minimal size overlap between the two larger species (graph I). However, all GL shape indices (graphs I, J) highlight the usual greater slenderness of roe deer specimens, while fallow deer and red deer values present positive allometric relations along different regression lines (graph I). In fallow deer, the smaller size of the shaft smallest depth (SD1), proportionally to all other measurements (as well as, to a lesser extent, the proportionally smaller BFd), shows up in graph J, with fallow deer values constrained in the bottom-left part of red deer value distribution range.\u003c/p\u003e\n\u003cp\u003eThe shape index value means (graph L) broadly support the observations made for graphs A-J. The fallow deer lower SD1 mean value compared to SD\u0026rsquo;s, translated in a lower SD1/SD index mean value in both mature and immature specimens, reflects the flatter shape of the metatarsus shaft of this species, as displayed in graphs F-H. The different dimensional proportions of distal widths and depths in the three cervid species is visible in the fallow deer and roe deer higher LSI mean values of some shape indices from both condyles. The relatively high value of the roe deer GL mean and the low values of all related robustness indices (SD/GL, SD1/GL, and Bd/GL) reflect the greater slenderness of this species\u0026rsquo; skeletal elements. The results of the Mann-Whitney U tests for the shape index measurements fully support the observations above (Table 10). Also the results of the PCA confirm the excellent degree of separation of red deer, fallow deer, and roe deer specimens (Open Resource 5: Fig 10).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe analysis of size and shape indices highlights a variable degree of success in the separation of red deer, fallow deer, and roe deer values from post-cranial elements.\u003c/p\u003e \u003cp\u003eIn mature specimens, size often provides good separation of the three species, from complete separation to mild overlaps. The use of biometrical data from populations of northern, central, and southern Europe, including Great Britain and part of the Italian peninsula, where red deer is smaller, allows to display a varying set of size ranges. As suggested in previous sections, however, such dimensional ranges are far from complete. The existence of smaller red deer than the specimens used here is of special concern when the proposed methodology aims to separate this species from the fallow deer, a slightly smaller animal. Iberian, southern Italian, south-eastern European, and north African red deer individuals, on average among the smallest representatives from continental mainlands (Mattioli \u0026amp; Ferretti, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Becciolini et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), are missing in the dataset; similarly, specimens from extant populations inhabiting medium- and small-sized islands, where red deer underwent varying and often complex processes of size decrease (Vigne, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Mulville, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), have not been sampled. In addition, red deer being an especially successful species due to its adaptability, its size fluctuates substantially to fit climatic change, geography, changes in habitat and dietary resources, and hunting pressure, both on long (Lister, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Di Stefano et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and much shorter chronological scales (Mattioli \u0026amp; Ferretti, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Mitchell et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1977\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn sum, wider ranges of red deer bone metric values than those displayed in this study should be born in mind, especially regarding the existence (past and present) of smaller red deer. As a consequence, a greater degree of bone dimensional overlaps must be considered in graphs that combine size ranges and shape index values. Nevertheless, the dimensional analyses presented here can at least suggest with which elements and measurements size overlaps are more likely to occur (e.g., the scapula SLC and GLP, the shaft measurements of all elements, in addition to the humerus and calcaneum Bd, the radius Bp, and the metapodials b), and which ones can instead be used more confidently to assist in taxonomic identification (e.g., the astragalus Bd and the metatarsus a).\u003c/p\u003e \u003cp\u003eShape indices present a varying degree of success in the taxonomic separation of the three species. When separation is provided, by one index or by a combination of two indices, this can either be in terms of isolation of the fallow deer cluster, when its values plot differently from both red deer\u0026rsquo;s and roe deer\u0026rsquo;s (e.g., the scapula ASG/GLP, the combination of humerus indices in graph E, the metapodials shape indices from both condyles, the tibia Ddb/Dda and SD1/SD), or in terms of isolation of one of the other species (e.g., the combination of humerus indices in graph F, the combination of metatarsus indices in graphs E and F). In the latter case, other measurements and shape indices can be plotted, if available, to facilitate or confirm species-level identification. An example is provided by the distal metatarsus, where, in addition to size, two shape indices per condyle allow to separate the three cervids: the width-depth proportions to separate fallow and red deer (a/3 and b/6), and the proportion of two depths for separating fallow and roe deer (3/2 and 6/5) (Fig.\u0026nbsp;11, graphs B, D).\u003c/p\u003e \u003cp\u003eThe different distributions produced by some shape indices reveal subtle morphological differences between species, which had not been previously observed; this may be a reflection of such differences being too slight to be detected by the eye, and/or being more easily missed due to the co-occurrence of size discrepancies. For example, one morphological trait that seems to characterise many fallow deer elements and zones is a distinct proportion between widths and depths, in this species often more in favour of widths: in most elements, the shaft shape index employing the smallest width and depth (SD1/SD) is lower in fallow deer, indicating that its shafts are \u0026lsquo;flatter\u0026rsquo; in comparison to red deer\u0026rsquo;s and roe deer\u0026rsquo;s; similarly, in the metatarsus distal condyles of fallow deer, widths are proportionally less smaller than depths compared to red deer\u0026rsquo;s (Fig.\u0026nbsp;11, graphs B, D). A more visible morphological trait is the greater depth of the insertion for the lateral ulnar articulation for the radius in red deer (Fig.\u0026nbsp;6, graph B; probably a reflection of character 6 from Lister (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1996\u003c/span\u003e: Fig.\u0026nbsp;2, Part 4 therein)), which prompted the introduction and analysis of measurement V, or, in the same element and zone, the slightly less pronounced lateral bicipital tuberosity of red deer (Fig.\u0026nbsp;6, graph A). On the other hand, the biometrical translation of the diagnostic character from (Di Stefano, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e: Fig.\u0026nbsp;5 therein) on the distal humerus (whereby in fallow deer the trochlea is directed laterally with a more acute angle), achieved by taking multiple height and width measurements, has not been particularly successful, only one index (HX/Bd) possibly reflecting such character (Fig.\u0026nbsp;5, graph A). Complete bone robusticity indices involving greatest length measurements consistently isolate the roe deer, whose long bones appear far slenderer than the other two species\u0026rsquo;.\u003c/p\u003e \u003cp\u003eImportantly for the aims of this study, shape index values from immature individuals (long bone shafts, proximal calcaneum) often follow similar distribution patterns to those observed in mature animals; as dimensional separation is more difficult with juvenile specimens, and the degree of ossification alone cannot be used to estimate adult size, these results may prove particularly relevant in the taxonomic identification of remains from younger cervids. In addition, the effectiveness of shaft shape indices (SD1/SD) in mature and immature individuals alike is also very important, as it suggests that it may now be possible to attempt species-level identification of fragmented cervid bones missing either or both ends (where diagnostic morphological criteria are described).\u003c/p\u003e \u003cp\u003eIn many instances, graphs combining size and shape index values reveal allometric relations between them. These are important to consider, as they contribute to assess the reliability and potentials of species-level identification. Using the cases reported above, for example, the \u0026lsquo;flatness\u0026rsquo; of the tibia shaft (Fig.\u0026nbsp;8, graphs C, D), as well as the metatarsus\u0026rsquo; (Fig.\u0026nbsp;11, graph G), increases (i.e., SD1/SD values become lower) with size in both fallow deer and red deer, although following different regression curves; this increased flatness is also observed for the distal metatarsus condyles described above, again with red and fallow deer values distributed along distinct curves (Fig.\u0026nbsp;11, graphs A, C). Similarly, the lateral angle created by the trochlea of the humerus, generally more acute in fallow deer than in red deer (Di Stefano, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e: Fig.\u0026nbsp;5 therein), becomes increasingly acuter with increased size in both species (Fig.\u0026nbsp;5, graph A), while the lateral bicipital tuberosity of the proximal radius becomes more pronounced (Fig.\u0026nbsp;6, graph A). These trends are the result of width measurements presenting a positive allometry (i.e., growing faster than other measurements). Other allometries are visible in the astragalus (Fig.\u0026nbsp;9, graph B) and the calcaneum (Fig.\u0026nbsp;10, graph B), in this case also involving the roe deer. The existence of allometries following different regression lines is especially useful in this study, as size and shape index overlaps are partly overcome by the different distributions, along similar ranges, of values from different species. The limited success of the c/d shape index of the calcaneum in taxonomic separation, for example, is overturned when the index is plotted against an absolute measurement, as its values increase with increased size along different regression lines for fallow deer and red deer. On the one hand, these trends highlight the importance of combining size and shape index values to maximise taxonomic separation; on the other, they warn against the acritical use of shape indices, as different shape index value distributions for differently-sized species may also be the result of allometric relations along similar regression curves: this could be the case of the tibia Ddb/Dda index values for the fallow and red deer, although in this case the regression trends remain unclear (Fig.\u0026nbsp;8, graph A). In this regard, it is also worth noting that, while the little efficiency of the astragalus\u0026rsquo; shape indices is partly overturned by allometric relations following different regression lines (Fig.\u0026nbsp;9, graph B), the PCA of shape indices produced three largely overlapping clusters: despite the optimization of taxonomic separation offered by more sophisticated data processing, therefore, the use of the PCA turns out, in this case, to be less efficient that a critical combination of size and shape analyses in scatter plots.\u003c/p\u003e \u003cp\u003eIn sum, the results of the biometrical analyses highlight that:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003eThe size ranges for the three species produce different degrees of dimensional overlaps for different elements and measurements, roe deer size values often separating from fallow deer\u0026rsquo;s better than red deer\u0026rsquo;s;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eThe selected shape indices separate the values from the three species with varying degrees of success, alone or in combination with size or other shape indices. Such distinct distributions are a reflection of both previously undetected and published morphological differences;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eSome shape indices remain effective when applied to the taxonomic separation of immature individuals, for which size overlaps are substantial and prevent identification;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eShaft shape indices are often effective, extending species-level identification attempts to fragmented materials with no published diagnostic morphological traits;\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eThe value distribution of many shape indices, plotted against size, reveal the existence of allometric relations between them. In many instances, such allometries rely on different regression lines for different species, amplifying the taxonomic separation potentials of combined shape and size analyses.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eThe results of PCA analysis of shape indices often reflect and sometimes reinforce the taxonomic separations provided by the comparisons of size and shape index value ranges presented in scatter plots, linear graphs, and box plots; however, it should be used alongside these latter analyses, to ensure a critical use of both size and shape data.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe biometrical analyses presented in this paper rely on data from nine modern reference collections and different European regions. They describe dimensional and morphological criteria that can be used to assist species-level identification of archaeological cervid bones. This is especially important for the separation of fallow deer remains; despite its known historical and ecological significance, indeed, this relatively rarer species has often been overlooked and sometimes its bones misidentified in European zooarchaeological research, leading to limited visibility in the archaeological record. Such underestimation biases our understanding of past human-cervid-environment interactions, calling for more accurate and objective approaches such as the one offered by this biometrical method.\u003c/p\u003e \u003cp\u003eThe obtained results show that size remains important evidence to use with caution, while the clustering of shape index values separately for the three species, despite the disparate origins of the modern skeletons, highlights the universal validity of the method when applied to materials from different regions and time periods. In addition, the many allometric relations between size and shape index values seem to support the existence of broadly similar morphometric characters; the often-different regression lines followed by such relations in each species prove useful in species-level separation.\u003c/p\u003e \u003cp\u003eOnly two published morphological diagnostic criteria (Di Stefano, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e: Fig.\u0026nbsp;5 therein; Lister, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1996\u003c/span\u003e: Fig.\u0026nbsp;2, Part 4, character 6 therein) were translated biometrically; all other shape indices reflect more subtle differences and can be considered \u0026lsquo;new\u0026rsquo; evidence. This characterises the method proposed here as an independent, complementary tool to use \u003cem\u003ein addition\u003c/em\u003e to others (use of appropriate reference collections, assessment of morphological criteria).\u003c/p\u003e \u003cp\u003eEqually important is the fact that the method is grounded in what are now frequently referred to - often with a somewhat dismissive connotation - as \u0026lsquo;traditional analyses\u0026rsquo;, as it consists of straightforward biometry. In a research landscape increasingly spellbound by techniques such as ZooMS, ancient DNA, and isotopic analyses, this study offers a timely and valuable re-evaluation of foundational methods. While scientifically advanced techniques undoubtedly expand the analytical potentials of faunal research, they often require substantial financial investment, specialised equipment and staff, and access to dedicated laboratory infrastructures - resources that are not always readily available to researchers. In addition, the (justified) sense of novelty conveyed when these techniques start being applied to certain materials, animal species, regions or periods, as well as the greater consideration implied by higher resource investments, may often lead to neglecting a proper integration of results with other lines of zooarchaeological evidence and within the broader archaeological and historical context. It has now become important to remind and value the primary role of methodologically sound and cost-effective \u0026lsquo;traditional analyses\u0026rsquo;, firmly grounded in archaeological reasoning and accessible to all.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eV.A.: Conceptualisation, Methodology, Formal analysis, Data Curation, Data processing, Writing - Original Draft, Review and Editing, Visualisation, Funding acquisition.M.R.: Conceptualisation, Methodology, Formal analysis, Data Curation, Data processing, Writing - Original Draft, Review and Editing, Visualisation, Funding acquisition.F.G.: Data curation, Data processing, Visualisation.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eThe authors are grateful to the institutions they visited for data collection: the former Department of Archaeology of the University of Sheffield (UK), Historic England (Fort Cumberland, UK), the Department of Natural History of the University-Museum of Bergen (Norway), the Natural History Museum (Austria), the Hungarian Natural History Museum (Budapest, Hungary), the Malcolm H. Wiener Laboratory for Archaeological Science of the American School of Classical Studies at Athens (Greece), the Department of Physical Sciences, Earth and Environment of the University of Siena (Italy), and the Natural History Museum of Venice (Italy). The data collection trip to Historic England also allowed to measure the modern deer from the private reference collection of Sheila Hamilton Dyer\u0026rsquo;s (Southampton, UK), who kindly made her specimens available. The authors are thankful to the many colleagues and friends who provided useful suggestions and comments: Umberto Albarella, Polydora Baker, Simon Davis, Kostantina Saliaris, and L\u0026aacute;szl\u0026oacute; Bartosiewicz. We would also like to thank (in alphabetical order): Umberto Albarella, Stephanie Baron, Marina Chorro-Giner, Steve Hollings, Phoebe Liu, Maeve Moorcroft, Mya Pearce-Ahdoud, Matt Rayner and Lenny Salvagno for their contribution in the estimation of the inter-observer error. A heartfelt thank you to Bj\u0026oslash;rn Reidar Olsson for providing alive deer photographs.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eCERVIFIED Database is available in Mendeley Data: https://data.mendeley.com/datasets/vn43dpvbzr/1doi: 10.17632/vn43dpvbzr.1\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlbarella, U., \u0026amp; Aniceti, V. (2024). \u0026apos;Unmaking\u0026apos; the deer in medieval Europe: Historical and archaeological evidence. \u003cem\u003eEuropean Journal of Archaeology, 27\u003c/em\u003e(3), 1-19. https://doi.org/10.1017/eaa.2024.11 \u003c/li\u003e\n\u003cli\u003eAlbarella, U., \u0026amp; Davis, S. (1996). Mammals and birds from Launceston Castle, Cornwall: Decline in status and the rise of agriculture. \u003cem\u003eCircaea, 12\u003c/em\u003e(1), 1-156.\u003c/li\u003e\n\u003cli\u003eAlmeida, N. J., Guinot, C., Ribeiro, I., Barreira, J., \u0026amp; Bas\u0026iacute;lio, A. C. (2024). Human-deer relations during Late Prehistory: The zooarchaeological data from central and southern Portugal in perspective. \u003cem\u003eAnimals, 14\u003c/em\u003e(10), 1424. https://doi.org/10.3390/ani14101424 \u003c/li\u003e\n\u003cli\u003eAniceti, V., Albarella, U., \u0026amp; Sebastiani, A. (2021). The faunal assemblage from the manufacturing district at Spolverino: From the Roman Imperial period to Late Antiquity. \u003cem\u003eArchaeofauna, 30\u003c/em\u003e, 167-183. https://doi.org/10.15366/archaeofauna2021.30.009 \u003c/li\u003e\n\u003cli\u003eApollonio, M., \u0026amp; Chirichella, R. (2023). Deer and climate change: Impacts and perspectives. \u003cem\u003eAnimal Production Science, 63\u003c/em\u003e(16), 1573-1582. https://doi.org/10.1071/AN22296 \u003c/li\u003e\n\u003cli\u003eBaker, K. H., Gray, H. W. I., Lister, A. M., Spassov, N., Welch, A. J., Trantalidou, K., Rose, H., Schroeder, H., \u0026amp; Hoelzel, A. R. (2024b). Ancient and modern DNA track temporal and spatial population dynamics in the European fallow deer since the Eemian interglacial. \u003cem\u003eScientific Reports, 14\u003c/em\u003e, 3015. https://doi.org/10.1038/s41598-023-48112-6 \u003c/li\u003e\n\u003cli\u003eBaker, K. H., Miller, H., Doherty, S., Gray, H. W. I., Daujat, J., \u0026Ccedil;akırlar, C., Love, S., Zant, J. M., Cooper, A., Irving-Pease, E. K., Burger, J., Rose, H., Thomas, R., Ross-Piper, J., Bradfield, J., Linderholm, A., Boessenkool, S., Biebach, I., Schroeder, H., \u0026amp; Hoelzel, A. R. (2024a). The 10,000-year biocultural history of fallow deer and its implications for conservation policy. \u003cem\u003eProceedings of the National Academy of Sciences, 121\u003c/em\u003e(8), e2310051121. https://doi.org/10.1073/pnas.2310051121 \u003c/li\u003e\n\u003cli\u003eBecciolini, V., Bozzi, R., Viliani, M., Biffani, S., \u0026amp; Ponzetta, M. P. (2016). Body measurements from selective hunting: Biometric features of red deer (Cervus elaphus) from Northern Apennine, Italy. \u003cem\u003eItalian Journal of Animal Science, 15\u003c/em\u003e(3), 461-472. https://doi.org/10.1080/1828051X.2016.1186505 \u003c/li\u003e\n\u003cli\u003eBeglane, F., Baker, K. H., Carden, R., Hoelzel, R., Lamb, A., Fhionnghaile, R., Vernesi, C., Mattingly, A., Gooley, R., Chechushkov, I., \u0026amp; Irving-Pease, E. (2018). Ireland\u0026apos;s fallow deer: Their historical, archaeological and biomolecular records. \u003cem\u003eProceedings of the Royal Irish Academy: Archaeology, Culture, History, Literature, 118C\u003c/em\u003e, 141-165. https://doi.org/10.3318/priac.2018.118.01 \u003c/li\u003e\n\u003cli\u003eBoessneck, J., Muller, H. H., \u0026amp; Teichert, M. (1964). Osteologische unterscheidungsmerkmale zwischen Schaf (Ovis aries Linne) und Ziege (Capra hircus Linne). \u003cem\u003eKuhn Archiv, 78\u003c/em\u003e, 1-129.\u003c/li\u003e\n\u003cli\u003eBowen, F., Carden, R. F., Daujat, J., Grouard, S., Miller, H., Perdikaris, S., \u0026amp; Sykes, N. (2016). Dama dentition: A new tooth eruption and wear method for assessing the age of fallow deer (Dama dama). \u003cem\u003eInternational Journal of Osteoarchaeology, 26\u003c/em\u003e, 1089-1098. https://doi.org/10.1002/oa.2523 \u003c/li\u003e\n\u003cli\u003eBuckley, M., \u0026amp; Whitcher Kansa, S. (2011). Collagen fingerprinting of archaeological bone and teeth remains from Domuztepe, South Eastern Turkey. \u003cem\u003eArchaeological and Anthropological Sciences, 3\u003c/em\u003e(3), 271-280. https://doi.org/10.1007/s12520-011-0066-z \u003c/li\u003e\n\u003cli\u003eCarden, R. F., McDevitt, A. D., Zachos, F. E., Woodman, P. C., O\u0026apos;Toole, P., Rose, H., Monaghan, N. T., Campana, M. G., Bradley, D. G., \u0026amp; Edwards, C. J. (2012). Phylogenetic, ancient DNA, fossil and morphometric analyses reveal ancient and modern introductions of a large mammal: The complex case of red deer (Cervus elaphus) in Ireland. \u003cem\u003eQuaternary Science Reviews, 42\u003c/em\u003e, 74-84. https://doi.org/10.1016/j.quascirev.2012.02.012 \u003c/li\u003e\n\u003cli\u003eChapman, D., \u0026amp; Chapman, N. (1975). \u003cem\u003eFallow deer: Their history, distribution and biology\u003c/em\u003e. Terence Dalton Ltd.\u003c/li\u003e\n\u003cli\u003eDavis, S. J. M. (1996). Measurements of a group of adult female Shetland sheep skeletons from a single flock: A baseline for zooarchaeologists. \u003cem\u003eJournal of Archaeological Science, 23\u003c/em\u003e(4), 593-612. https://doi.org/10.1006/jasc.1996.0056 \u003c/li\u003e\n\u003cli\u003eDavis, S. J. M., \u0026amp; MacKinnon, M. (2009). Did the Romans bring fallow deer to Portugal? \u003cem\u003eEnvironmental Archaeology, 14\u003c/em\u003e, 15-26. https://doi.org/10.1179/174963109X400646 \u003c/li\u003e\n\u003cli\u003eDi Stefano, G. (1995). Identification of fallow deer remains on the basis of its skeletal features: Taxonomical considerations. \u003cem\u003eBollettino della Societ\u0026agrave; Paleontologica Italiana, 34\u003c/em\u003e(3), 323-331.\u003c/li\u003e\n\u003cli\u003eDi Stefano, G., Pandolfi, L., Petronio, C., \u0026amp; Salari, L. (2015). The morphometry and the occurrence of Cervus elaphus (Mammalia, Cervidae) from the Late Pleistocene of the Italian peninsula. \u003cem\u003eRivista Italiana di Paleontologia e Stratigrafia, 121\u003c/em\u003e(1), 103-120. https://doi.org/10.13130/2039-4942/6402 \u003c/li\u003e\n\u003cli\u003eDriesch, A. von den. (1976). \u003cem\u003eA guide to the measurement of animal bones from archaeological sites\u003c/em\u003e. Peabody Museum Bulletin 1. Peabody Museum of Archaeology and Ethnology.\u003c/li\u003e\n\u003cli\u003eFern\u0026aacute;ndez, H. (2001). \u003cem\u003eOst\u0026eacute;ologie compar\u0026eacute;e des petits ruminants euroasiatiques sauvages et domestiques (genres Rupicapra, Ovis, Capra et Capreolus): Diagnose diff\u0026eacute;rentielle du squelette appendiculaire\u003c/em\u003e [Doctoral dissertation, University of Geneva]. https://archive-ouverte.unige.ch/unige:77284\u003c/li\u003e\n\u003cli\u003eGrant, A. (1988). Food, status and religion in England in the Middle Ages: An archaeozoological perspective. \u003cem\u003eAnthropozoologica, Special Issue 2\u003c/em\u003e, 139-146.\u003c/li\u003e\n\u003cli\u003eHolmes, M. (2014). Making a fast buck in the Middle Ages: Evidence for poaching from medieval Wakefield. In K. Baker, R. Carden, \u0026amp; R. Madgwick (Eds.), \u003cem\u003eDeer and people\u003c/em\u003e (pp. 200-207). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.20 \u003c/li\u003e\n\u003cli\u003eKarastoyanova, N., Gorczyk, J., \u0026amp; Spassov, N. (2020). The natural history of the fallow deer, Dama dama (Linnaeus, 1758) in Bulgaria in prehistory and new evidence for the existence of an autochthonous Holocene population in the Balkans. \u003cem\u003eInternational Journal of Osteoarchaeology, 30\u003c/em\u003e(5), 616-628. https://doi.org/10.1002/oa.2886 \u003c/li\u003e\n\u003cli\u003eKoo, T. K., \u0026amp; Li, M. Y. (2016). A guideline of selecting and reporting Intraclass Correlation Coefficients for reliability research. \u003cem\u003eJournal of Chiropractic Medicine, 15\u003c/em\u003e, 155-163. https://doi.org/10.1016/j.jcm.2016.02.012 \u003c/li\u003e\n\u003cli\u003eLister, A. M. (1984). Evolutionary and ecological origins of British deer. \u003cem\u003eProceedings of the Royal Society of Edinburgh Section B: Biological Sciences, 82\u003c/em\u003e(4), 205-229. https://doi.org/10.1017/S0269727000003754 \u003c/li\u003e\n\u003cli\u003eLister, A. M. (1989). Rapid dwarfing of red deer on Jersey in the Last Interglacial. \u003cem\u003eNature, 342\u003c/em\u003e, 539-542. https://doi.org/10.1038/342539a0 \u003c/li\u003e\n\u003cli\u003eLister, A. M. (1996). The morphological distinction between bones and teeth of fallow deer (Dama dama) and red deer (Cervus elaphus). \u003cem\u003eInternational Journal of Osteoarchaeology, 6\u003c/em\u003e(2), 119-143. https://doi.org/10.1002/(SICI)1099-1212(199603)6:2\u0026lt;119::AID-OA265\u0026gt;3.0.CO;2-8 \u003c/li\u003e\n\u003cli\u003eMar\u0026iacute;n, J., Saladi\u0026eacute;, P., Azorit, C., \u0026amp; Rodr\u0026iacute;guez-Hidalgo, A. (2024). Identification of age at death in red deer (Cervus elaphus) through the upper dentition: Eruption pattern, wear stage and crown heights. \u003cem\u003eJournal of Archaeological Science, 163\u003c/em\u003e, 105934. https://doi.org/10.1016/j.jas.2023.105934 \u003c/li\u003e\n\u003cli\u003eMasseti, M., \u0026amp; Vernesi, C. (2014). Historic zoology of the European fallow deer, Dama dama dama: Evidence from biogeography, archaeology and genetics. In K. Baker, R. Carden, \u0026amp; R. Madgwick (Eds.), \u003cem\u003eDeer and people\u003c/em\u003e (pp. 13-22). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.6 \u003c/li\u003e\n\u003cli\u003eMattioli, S., \u0026amp; Ferretti, F. (2014). Morphometric characterization of Mesola red deer Cervus elaphus italicus (Mammalia: Cervidae). \u003cem\u003eItalian Journal of Zoology, 81\u003c/em\u003e(1), 144-154. https://doi.org/10.1080/11250003.2014.895432 \u003c/li\u003e\n\u003cli\u003eMcDevitt, A. D., \u0026amp; Zachos, F. E. (2014). Genetic analyses of natural and anthropogenic movements in deer. In K. Baker, R. Carden, \u0026amp; R. Madgwick (Eds.), \u003cem\u003eDeer and people\u003c/em\u003e (pp. 2-12). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.5 \u003c/li\u003e\n\u003cli\u003eMcElligott, A. G., Gammell, M. P., Harty, H. C., Paini, D. R., Murphy, D. T., Walsh, J. T., \u0026amp; Hayden, T. J. (2001). Sexual size dimorphism in fallow deer (Dama dama): Do larger, heavier males gain greater mating success? \u003cem\u003eBehavioral Ecology and Sociobiology, 49\u003c/em\u003e, 266-272. https://doi.org/10.1007/s002650000293 \u003c/li\u003e\n\u003cli\u003eMiller, H., \u0026amp; Sykes, N. (2016). Zootherapy in archaeology: The case of the fallow deer (Dama dama dama). \u003cem\u003eJournal of Ethnobiology, 36\u003c/em\u003e(2), 257-276. https://doi.org/10.2993/0278-0771-36.2.257 \u003c/li\u003e\n\u003cli\u003eMilo\u0026scaron;ević-Zlatanović, S., Toma\u0026scaron;ević Kolarov, N., Vukov, T., \u0026amp; Stamenković, S. (2016). Correlation patterns in roe deer cranium: Sexual dimorphism across different habitats. \u003cem\u003eJournal of Zoology, 300\u003c/em\u003e(4), 291-304. https://doi.org/10.1111/jzo.12383 \u003c/li\u003e\n\u003cli\u003eMitchell, B., Staines, B. W., \u0026amp; Welch, D. (1977). \u003cem\u003eEcology of red deer: A research review relevant to their management in Scotland\u003c/em\u003e. Institute of Terrestrial Ecology.\u003c/li\u003e\n\u003cli\u003eMulville, J. (2010). Red deer on Scottish islands. In T. O\u0026apos;Connor \u0026amp; N. Sykes (Eds.), \u003cem\u003eExtinctions and invasions: A social history of British fauna\u003c/em\u003e (pp. 43-50). Windgather Press. https://doi.org/10.2307/j.ctv13gvg6k.12 \u003c/li\u003e\n\u003cli\u003ePayne, S. (1969). A metrical distinction between sheep and goat metacarpals. In P. J. Ucko \u0026amp; G. W. Dimbleby (Eds.), \u003cem\u003eThe domestication and exploitation of plants and animals\u003c/em\u003e (pp. 295-305). Duckworth.\u003c/li\u003e\n\u003cli\u003ePayne, S., \u0026amp; Bull, G. (1988). Components of variation in measurements of pig bones and teeth, and use of measurements to distinguish wild from domestic remains. \u003cem\u003eArchaeozoologia, 2\u003c/em\u003e, 27-66.\u003c/li\u003e\n\u003cli\u003ePost, E., Langvatn, R., Forchhammer, M. C., \u0026amp; Stenseth, N. C. (1999). Environmental variation shapes sexual dimorphism in red deer. \u003cem\u003eProceedings of the National Academy of Sciences, 96\u003c/em\u003e(8), 4467-4471. https://doi.org/10.1073/pnas.96.8.4467 \u003c/li\u003e\n\u003cli\u003eSalvagno, L., \u0026amp; Albarella, U. (2017). A morphometric system to distinguish sheep and goat postcranial bones. \u003cem\u003ePLoS ONE, 12\u003c/em\u003e(6), e0178543. https://doi.org/10.1371/journal.pone.0178543 \u003c/li\u003e\n\u003cli\u003eSchmidt, K. T. (2014). Supplemental feeding and our attitude towards red deer and natural mortality. In K. Baker, R. Carden, \u0026amp; R. Madgwick (Eds.), \u003cem\u003eDeer and people\u003c/em\u003e (pp. 270-279). Oxbow Books. https://doi.org/10.2307/j.ctv13gvgms.27 \u003c/li\u003e\n\u003cli\u003eSerrano, E., Angibault, J.-M., Cargnelutti, B., \u0026amp; Hewison, A. J. M. (2007). The effect of animal density on metacarpus development in captive fallow deer. \u003cem\u003eSmall Ruminant Research, 72\u003c/em\u003e(1), 61-65. https://doi.org/10.1016/j.smallrumres.2006.07.007 \u003c/li\u003e\n\u003cli\u003eSkog, A., Zachos, F. E., Rueness, E. K., Feulner, P. G. D., Mysterud, A., Langvatn, R., Lorenzini, R., Hmwe, S. S., Lehoczky, I., Hartl, G. B., Stenseth, N. C., \u0026amp; Jakobsen, K. S. (2009). Phylogeography of red deer (Cervus elaphus) in Europe. \u003cem\u003eJournal of Biogeography, 36\u003c/em\u003e(1), 66-77. https://doi.org/10.1111/j.1365-2699.2008.01986.x \u003c/li\u003e\n\u003cli\u003eSoderberg, J. (2004). Wild cattle: Red deer in the religious texts, iconography, and archaeology of early medieval Ireland. \u003cem\u003eInternational Journal of Historical Archaeology, 8\u003c/em\u003e, 167-183. https://doi.org/10.1007/s10761-004-1135-0 \u003c/li\u003e\n\u003cli\u003eSykes, N., Ayton, G., Bowen, F., Baker, K., Baker, P., Carden, R., Hammon, C., Ingrem, C., Madgwick, R., Miller, H., Poole, K., Rowley-Conwy, P., Worley, F., \u0026amp; Evans, J. (2016). Wild to domestic and back again: The dynamics of fallow deer management in medieval England (c. 11ᵗʰ-16ᵗʰ century AD). \u003cem\u003eScience and Technology of Archaeological Research, 2\u003c/em\u003e(1), 113-126. https://doi.org/10.1080/20548923.2016.1208027 \u003c/li\u003e\n\u003cli\u003eSykes, N., Carden, R., \u0026amp; Harris, K. (2013). Changes in the size and shape of fallow deer \u0026mdash; Evidence for the movement and management of a species. \u003cem\u003eInternational Journal of Osteoarchaeology, 23\u003c/em\u003e(1), 55-68. https://doi.org/10.1002/oa.1239 \u003c/li\u003e\n\u003cli\u003eTo\u0026iuml;go, C., Gaillard, J.-M., Laere, G., Hewison, A., \u0026amp; Morellet, N. (2006). How does environmental variation influence body mass, body size, and body condition? Roe deer as a case study. \u003cem\u003eEcography, 29\u003c/em\u003e, 301-308. https://doi.org/10.1111/j.2006.0906-7590.04394.x \u003c/li\u003e\n\u003cli\u003eVigne, J.-D. (1988). \u003cem\u003eLes mammif\u0026egrave;res post-glaciaires de Corse: \u0026Eacute;tude arch\u0026eacute;ozoologique\u003c/em\u003e. Gallia Pr\u0026eacute;histoire Suppl. 26. \u0026Eacute;ditions du Centre National de la Recherche Scientifique.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables 1 to 10 are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"archaeological-and-anthropological-sciences","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"aasc","sideBox":"Learn more about [Archaeological and Anthropological Sciences](http://link.springer.com/journal/12517)","snPcode":"12520","submissionUrl":"https://submission.nature.com/new-submission/12520/3","title":"Archaeological and Anthropological Sciences","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"red deer, fallow deer, roe deer, method, identification, biometry, archaeology","lastPublishedDoi":"10.21203/rs.3.rs-6636241/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6636241/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis paper presents a new method for differentiating postcranial bone remains of red deer (\u003cem\u003eCervus elaphus\u003c/em\u003e), fallow deer (\u003cem\u003eDama dama\u003c/em\u003e), and roe deer (\u003cem\u003eCapreolus capreolus\u003c/em\u003e) from archaeological contexts. These species have a very similar bone morphology and partly overlapping size ranges, often preventing species-level identification in faunal research. Traditional methods, such as the use of diagnostic morphological criteria, or aDNA analysis, present practical and methodological limitations. In this study, bone biometric data from modern specimens, sourced from various institutions across northern and southern Europe, are analysed to determine size ranges and shape indices that can separate the three species, focussing on both adult and immature individuals. By comparing measurement values from archaeological remains with these biometric criteria, species identification can be attempted. The low-tech nature of this approach and the transparent, diagrammatic presentation of biometric results make the new method objective, highly accessible, and inclusive. In European and Mediterranean archaeology, the separation of fallow deer from other cervid species is crucial in the investigation of past biogeography, trading activities, hunting strategies, cultural values, and social status.\u003c/p\u003e","manuscriptTitle":"‘Cervified’: a method for the morphometric identification of red deer (Cervus elaphus), fallow deer (Dama dama), and roe deer (Capreolus capreolus) archaeological bones","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-05-16 13:02:50","doi":"10.21203/rs.3.rs-6636241/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-06-19T08:44:30+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-13T03:53:21+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-04T07:28:11+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-05-27T08:29:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"181953782295083304664352984460684780901","date":"2025-05-16T05:20:07+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"268873714386980650020758769727823733267","date":"2025-05-15T19:26:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"198236757670076820756388724480553044481","date":"2025-05-15T11:03:45+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-05-13T19:09:51+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-12T16:41:52+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-12T07:23:43+00:00","index":"","fulltext":""},{"type":"submitted","content":"Archaeological and Anthropological Sciences","date":"2025-05-10T17:41:04+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"archaeological-and-anthropological-sciences","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"aasc","sideBox":"Learn more about [Archaeological and Anthropological Sciences](http://link.springer.com/journal/12517)","snPcode":"12520","submissionUrl":"https://submission.nature.com/new-submission/12520/3","title":"Archaeological and Anthropological Sciences","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"dd494be7-8d82-4088-bf3b-a315570a9048","owner":[],"postedDate":"May 16th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-01-19T16:47:30+00:00","versionOfRecord":{"articleIdentity":"rs-6636241","link":"https://doi.org/10.1007/s12520-025-02380-7","journal":{"identity":"archaeological-and-anthropological-sciences","isVorOnly":false,"title":"Archaeological and Anthropological Sciences"},"publishedOn":"2026-01-16 16:29:40","publishedOnDateReadable":"January 16th, 2026"},"versionCreatedAt":"2025-05-16 13:02:50","video":"","vorDoi":"10.1007/s12520-025-02380-7","vorDoiUrl":"https://doi.org/10.1007/s12520-025-02380-7","workflowStages":[]},"version":"v1","identity":"rs-6636241","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6636241","identity":"rs-6636241","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}