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Ascertaining an agent: using tooth pit data to determine the carnivore/s responsible for predation in cases of suspected big cat kills in an upland area of Britain
Abstract
The identification of the involvement of a particular carnivore in the modification of bone assemblages concerns a number of fields of research including archaeological and palaeontological enquiry. Taphonomy provides a methodology by which bone assemblages can be analysed and interpreted and this is more often undertaken with archaeological or palaeontological assemblages. A taphonomic analysis is undertaken here in order to determine the perpetrator of predation attacks on domestic stock from a modern-day setting. Recently reported techniques using tooth marks preserved on bone surfaces made by known carnivores are successful at determining some class sizes of predators and are used here to determine the perpetrator(s). Although a class size of carnivore is readily identified by this methodology, a particular carnivore taxon is not. Tooth morphology and dental configuration are reported here as better criteria for identifying a particular taphonomic agent. Tooth pit dimensions are used here to identify the class size of carnivores involved, and tooth morphology and cusp spacing to suggest a medium sized felid and fox as taphonomic agents. The identification of the medium-sized felid may support observations and reports of alleged “big” cat kills in the area. The study has important implications for the interpretation of fossil sites where felids may have been involved in the modification of animal carcasses but are archaeologically invisible in terms of their fossil remains.
... Los animales carroñeros pueden dañar huesos en distintas formas. Los carnívoros como los canidos generalmente dejan 4 marcas en hueso: depresiones, surcos, perforaciones y hendiduras (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Andrews y Fernandez-Jalvo, 1997;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012). Las depresiones ocurren cuando el chupeteo o las mordidas pierden poder para atravesar hueso pero aún lo marcan; los surcos son más profundos que las hendiduras o canales en hueso, son causados por molares para llegar a la cavidad medular (con continua actividad mandibular) y son longitudinales al final del asta abierta; las perforaciones ocurren cuando el hueso es atravesado y forma un pequeño agujero; y las hendiduras ocurren cuando el diente arrastra la superficie del hueso, en general su largo es tres veces mayor que su ancho (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009). ...
... Los carnívoros como los canidos generalmente dejan 4 marcas en hueso: depresiones, surcos, perforaciones y hendiduras (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Andrews y Fernandez-Jalvo, 1997;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012). Las depresiones ocurren cuando el chupeteo o las mordidas pierden poder para atravesar hueso pero aún lo marcan; los surcos son más profundos que las hendiduras o canales en hueso, son causados por molares para llegar a la cavidad medular (con continua actividad mandibular) y son longitudinales al final del asta abierta; las perforaciones ocurren cuando el hueso es atravesado y forma un pequeño agujero; y las hendiduras ocurren cuando el diente arrastra la superficie del hueso, en general su largo es tres veces mayor que su ancho (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009). Wiley y Snyder (1989 reportaron que Canis lupus L. destruye los extremos de los huesos porosos más largos que los de los huesos compactos, causando perforaciones. ...
... Por el contrario, depresiones en diáfisis de 2 mm o menos de largo y 1,5 mm o menos de ancho, en epífisis de huesos largos, se asocian a carnívoros de tamaño medio como babuinos, chacales, osos y perros; mientras que depresiones más largas y anchas que 4 mm se asocian a carnívoros mayores como hienas, leones y lobos (Dominguez-Rodrigo y Piqueras, 2003;Pickering et al., 2004;Foust, 2007). Coard (2007) encontró que realizar un puntaje sobre el ancho de las heridas en hueso puede servir para identificar carnívoros y distinguir carnívoros grandes de pequeños. Estos harán marcas proporcionales a su tamaño, los primeros harán marcas de ancho pequeño o grande, y los segundos pueden hacer sólo marcas de ancho pequeño. ...
... Los animales carroñeros pueden dañar huesos en distintas formas. Los carnívoros como los canidos generalmente dejan 4 marcas en hueso: depresiones, surcos, perforaciones y hendiduras (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Andrews y Fernandez-Jalvo, 1997;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012). Las depresiones ocurren cuando el chupeteo o las mordidas pierden poder para atravesar hueso pero aún lo marcan; los surcos son más profundos que las hendiduras o canales en hueso, son causados por molares para llegar a la cavidad medular (con continua actividad mandibular) y son longitudinales al final del asta abierta; las perforaciones ocurren cuando el hueso es atravesado y forma un pequeño agujero; y las hendiduras ocurren cuando el diente arrastra la superficie del hueso, en general su largo es tres veces mayor que su ancho (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009). ...
... Los carnívoros como los canidos generalmente dejan 4 marcas en hueso: depresiones, surcos, perforaciones y hendiduras (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Andrews y Fernandez-Jalvo, 1997;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012). Las depresiones ocurren cuando el chupeteo o las mordidas pierden poder para atravesar hueso pero aún lo marcan; los surcos son más profundos que las hendiduras o canales en hueso, son causados por molares para llegar a la cavidad medular (con continua actividad mandibular) y son longitudinales al final del asta abierta; las perforaciones ocurren cuando el hueso es atravesado y forma un pequeño agujero; y las hendiduras ocurren cuando el diente arrastra la superficie del hueso, en general su largo es tres veces mayor que su ancho (Binford, 1981;Haynes, 1983;Haglund et al., 1988;Milner y Smith, 1989;Pickering et al., 2004;Coard, 2007;Delaney-Rivera et al., 2009). Wiley y Snyder (1989 reportaron que Canis lupus L. destruye los extremos de los huesos porosos más largos que los de los huesos compactos, causando perforaciones. ...
... Por el contrario, depresiones en diáfisis de 2 mm o menos de largo y 1,5 mm o menos de ancho, en epífisis de huesos largos, se asocian a carnívoros de tamaño medio como babuinos, chacales, osos y perros; mientras que depresiones más largas y anchas que 4 mm se asocian a carnívoros mayores como hienas, leones y lobos (Dominguez-Rodrigo y Piqueras, 2003;Pickering et al., 2004;Foust, 2007). Coard (2007) encontró que realizar un puntaje sobre el ancho de las heridas en hueso puede servir para identificar carnívoros y distinguir carnívoros grandes de pequeños. Estos harán marcas proporcionales a su tamaño, los primeros harán marcas de ancho pequeño o grande, y los segundos pueden hacer sólo marcas de ancho pequeño. ...
... Gifford-Gonzalez, 1991: 217 As marcas de dentes, associadas a outros aspectos comportamentais, são um dos indicadores tafonómicos mais informativos para realizar inferências acerca da formação dos registos arqueofaunísticos. Tratam-se de estrias sinuosas com secção transversal em U, i.e., fundo plano ou | ARKEOS 44 | 55 | ZOOARQUEOLOGIA E TAFONOMIA DA TRANSIÇÃO PARA A AGRO-PASTORÍCIA NO BAIXO E MÉDIO VALE DO TEJO | aplanado, morfometricamente variável, com uma propensão para a presença de bordos arredondados (Tabela 4) (Binford, 1981;Haynes, 1980Haynes, , 1983bMaguire et al., 1980;Bunn, 1981;Blumenschine, 1988;Selvaggio e Wilder, 2001;Domínguez-Rodrigo, 2002;Domínguez--Rodrigo e Piqueras, 2003;Landt, 2004Landt, , 2007Delaney-Rivera et al., 2009;Saladié, 2009 (Brain, 1969;Binford, 1978Binford, , 1981Maguire et al., 1980;Jones, 1983;Oliver, 1993;White, 1992;Elkin e Mondini, 2001;Landt, 2004Landt, , 2007Cáceres et al., 2007;White e Tooth, 2007;Blasco e Fernández Peris, 2009;Delaney-Rivera et al., 2009;Saladié, 2009;Fernández-Jalvo e Andrews, 2011;Andrés, 2012;Saladié et al., 2013), de suídeos (Greenfield, 1988;Domínguez-Solera e Domínguez--Rodrigo, 2008;Saladié, 2009), canídeos (Haynes, 1980(Haynes, , 1983b(Haynes, , 1985Binford, 1981;Stalibrass, 1984Stalibrass, , 1990Payne e Munson, 1985;Elkin e Mondini, 2001;Domínguez-Rodrigo e Piqueras, 2003;Coard, 2007;Campmas e Beauval, 2008;Delaney-Rivera et al., 2009;Yravedra et al., 2011;Andrés, 2012;Krajcarz e Krajcarz, 2012;Sala et al., 2014), felídeos (Moran e O'Connor, 1991Selvaggio e Wilder, 2001;Coard, 2007;Delaney-Rivera et al., 2009;Andrés, 2012), mustelídeos (Mallye, 2011;Mallye et al., 2008), roedores (Maguire et al., 1980;Brain, 1981;Shipman e Rose, 1983) e herbívoros osteófagos (Sutcliffe, 1973;Brothwell, 1976;Barrette, 1985;Cáceres et al., 2011). ...
... Gifford-Gonzalez, 1991: 217 As marcas de dentes, associadas a outros aspectos comportamentais, são um dos indicadores tafonómicos mais informativos para realizar inferências acerca da formação dos registos arqueofaunísticos. Tratam-se de estrias sinuosas com secção transversal em U, i.e., fundo plano ou | ARKEOS 44 | 55 | ZOOARQUEOLOGIA E TAFONOMIA DA TRANSIÇÃO PARA A AGRO-PASTORÍCIA NO BAIXO E MÉDIO VALE DO TEJO | aplanado, morfometricamente variável, com uma propensão para a presença de bordos arredondados (Tabela 4) (Binford, 1981;Haynes, 1980Haynes, , 1983bMaguire et al., 1980;Bunn, 1981;Blumenschine, 1988;Selvaggio e Wilder, 2001;Domínguez-Rodrigo, 2002;Domínguez--Rodrigo e Piqueras, 2003;Landt, 2004Landt, , 2007Delaney-Rivera et al., 2009;Saladié, 2009 (Brain, 1969;Binford, 1978Binford, , 1981Maguire et al., 1980;Jones, 1983;Oliver, 1993;White, 1992;Elkin e Mondini, 2001;Landt, 2004Landt, , 2007Cáceres et al., 2007;White e Tooth, 2007;Blasco e Fernández Peris, 2009;Delaney-Rivera et al., 2009;Saladié, 2009;Fernández-Jalvo e Andrews, 2011;Andrés, 2012;Saladié et al., 2013), de suídeos (Greenfield, 1988;Domínguez-Solera e Domínguez--Rodrigo, 2008;Saladié, 2009), canídeos (Haynes, 1980(Haynes, , 1983b(Haynes, , 1985Binford, 1981;Stalibrass, 1984Stalibrass, , 1990Payne e Munson, 1985;Elkin e Mondini, 2001;Domínguez-Rodrigo e Piqueras, 2003;Coard, 2007;Campmas e Beauval, 2008;Delaney-Rivera et al., 2009;Yravedra et al., 2011;Andrés, 2012;Krajcarz e Krajcarz, 2012;Sala et al., 2014), felídeos (Moran e O'Connor, 1991Selvaggio e Wilder, 2001;Coard, 2007;Delaney-Rivera et al., 2009;Andrés, 2012), mustelídeos (Mallye, 2011;Mallye et al., 2008), roedores (Maguire et al., 1980;Brain, 1981;Shipman e Rose, 1983) e herbívoros osteófagos (Sutcliffe, 1973;Brothwell, 1976;Barrette, 1985;Cáceres et al., 2011). ...
... Os dados obtidos indicam que pequenos carnívoros e omnívoros podem mais facilmente ser diferenciados de grandes carnívoros através de comparações do comprimento de mordiscos em tecido cortical; quanto aos sulcos, a sua largura permite distinguir entre pequenos e grandes carnívoros (cf. Domínguez-Rodrigo e Piqueras, 2003: 1386Coard, 2007Coard, : 1680. As duas variáveis estatisticamente significantes quanto às dimensões das marcas de dentes seriam o tamanho do carnívoro (pequeno, grande) e da carcaça. ...
... The types of damage left upon bone are likely affected by the dental patterns, tooth morphology, feeding habits, and bite strengths of the scavengers involved. Different families of carnivores, such as Canidae and Felidae, have different dental morphologies, but most carnivore species have broad similarities, such as mesodont crowns and secodont cusps; blade-like carnassials; pointed premolars; dagger-like canines; and small incisors (Coard 2007;Van Valkenburgh 1996). Van Valkenburgh (1996) studied the variation in carnivore tooth morphology while observing the feeding patterns of spotted hyenas (Crocuta crocuta), wild dogs (Lycaon pictus), lions (Panthera leo), and cheetahs (Acinonyx jubatus). ...
... Previous research has classified and defined tooth marks as pits, punctures, scores and furrows (Andrews & Fernández-Jalvo 1997;Binford 1981;Burke 2013;Coard 2007;Delaney-Rivera et al. 2009;Domínguez-Rodrigo & Piqueras 2003;Domínguez-Rodrigo et al. 2012;Haglund 1997;Haynes 1980a;Lyman 1994;Pobiner 2007;Pokines & Kerbis Peterhans 2007;Pokines & Tersigni-Tarrant 2017;Sala & Arsuaga 2018;Saladié et al. 2011;Selvaggio & Wilder 2001). Pits range from circular-to oval-shaped depressions that have a length that is less than 3× their width and do not penetrate the cortical bone. ...
Tooth mark and other gnawing damage modifications on bone from African carnivores have been extensively examined, but there are less data on North American carnivores, especially on Ursidae (bears). The present study examined gnawing damage by captive black bear (Ursus americanus) and grizzly bear (U. arctos) fed 55 proximal or distal femora from cattle (Bos taurus) in order to distinguish ursid gnaw damage characteristics. Tooth mark modifications examined include pits, punctures, scores, and furrows, while other gnaw damage modifications include crenellated margins, edge polish, scalloping, scooping, and crushed margins. Each tooth mark was processed through the open-source software ImageJ in order to obtain the area, perimeter, length, and width. Tooth pits had an average length of 3.5 mm and average width of 2.2 mm; scores had an average width of 1.5 mm. There was a statistically significant difference between ursid tooth pits and those created by various other scavenging species. Other common taphonomic effects included scalloping on the distal end of the femur, especially on the patellar articular surface; scooping on the proximal end of the femur, especially on the greater trochanter; and furrows, primarily on the distal end of the femur along the patellar articular surface and condyles. Cancellous scooping occurred in 35.2% of the entire sample, while scalloping occurred in 29.6% of the entire sample. These high percentages may be distinctive characteristics of ursid gnaw damage and therefore may help distinguish ursid scavenging from that of other carnivores.
... The relative amount of damage caused to a bone is dependent on the relation between the strength of the bone and the tooth morphology and jaw mechanics of the scavenging animal [12]. Recent studies have determined that the size of tooth marks in bone can assist in determining the general size of the scavenger but the particular carnivore taxon cannot be accurately identified through tooth mark measurements [13]. Bones can even be entirely consumed, leaving little to analyse, depending on the size of the scavenging animal [12]. ...
... Measurements of the tooth marks were not taken. It is suggested that the size of tooth marks in bone can assist in determining the general size of the scavenger but the particular carnivore taxon cannot be accurately identified through tooth mark measurements [13]. ...
Forensic anthropologists in Southern Africa are often confronted with post-mortem modifications of human skeletal remains caused by animal scavenging. This is troublesome as the post-mortem pseudo-trauma could be misinterpreted. This study aimed to describe the skeletal trauma caused by Southern African scavengers which are of forensic interest. The scavenging animals selected for this study included wild dog, spotted hyena, lion, leopard, black-backed jackal, caracal, and porcupine housed at the National Zoological Gardens of South Africa. Sparsely fleshed and articulated bovine front and hind limbs as well as ribs were placed in each animal enclosure and collected after 2 days for cleaning and analysis. Felids (lion, leopard, and caracal) gnawed away the greater trochanter of the humerus leaving deep, parallel furrows. Hyena caused massive trauma to bone with one-third of the tibia shaft surviving with jagged fracture edges. Porcupines left distinctive fan-like parallel scores and large, oval depressions with an eroded, polished appearance. Wild dogs and jackals did not leave any distinctive patterns. Light scavenging trauma is distinct from other bone taphonomy but cannot be used to determine the species. Heavy scavenging trauma patterns can be used to determine the general type of perpetrating animal; however, the exact taxa or species cannot be determined.
... The maximum diameters of the puncture on the frontal bone are 11.8 Â 10.6 mm (SOM Fig. S3), which we used to narrow the range of possible mammalian carnivores. While it is extremely difficult to determine specific taxa responsible for bites, it is possible to correlate bite size to carnivore group considering body size and behavioral characteristics (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009). Compared to experimental bite mark data from Domínguez-Rodrigo and Piqueras (2003) and Delaney-Rivera et al. (2009), the bite is in the size-range of a spotted or striped hyena, wolf sized canid, or lion sized felid. ...
... Compared to experimental bite mark data from Domínguez-Rodrigo and Piqueras (2003) and Delaney-Rivera et al. (2009), the bite is in the size-range of a spotted or striped hyena, wolf sized canid, or lion sized felid. We should note that in their experimental work, the authors provide data from cancellous and compact portions of long bones, which may not be analogous to crania, and the carnivores in these studies were fed bone as opposed to carcasses (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009). Because of this, and due to the similarities in damage patterns between what we observe and what Brain (1980) and Simons (1966) chronicle, a medium sized felid such as a leopard is the likely candidate for killing or scavenging this primate, though we do not rule out the possibility of the involvement of a larger bodied carnivore. ...
The Pleistocene hominin site of Makuyuni, near Lake Manyara, Tanzania, is known for fossils attributable to Homo and Acheulean artifacts (Ring et al., 2005; Kaiser et al., 2010; Frost et al., 2012). Here we describe the fossil primate material from the Manyara Beds, which includes the first nearly complete female cranium of Theropithecus oswaldi leakeyi and a proximal tibia from the same taxon. The cranium is dated to between 633 and 780 Ka and the tibia to the Pleistocene. The T. oswaldi lineage is one of the most important among Neogene mammals of Africa: it is both widespread and abundant. The size of the dentition, cranium, and tibia all confirm the previously recognized trend of increasing body size in this lineage and make their taxonomic assignments secure. The morphology of this specimen provides new insights into the evolution of this lineage through time, as well as its geographic variation and sexual dimorphism. The cranium also shows damage consistent with a mammalian carnivore, most likely a felid. The identification of this material as representing T. o. leakeyi agrees with the Middle Pleistocene age estimates for the MK4 locality in particular and the Manyara Beds in general.
... Carnivores are one of the most destructive biological agents, so they have been subject of research by several authors (e.g. Hughes, 1954;Sutcliffe, 1970;Haynes, 1980Haynes, , 1981Binford, 1981;Brain, 1981;Solomon and David, 1990;Cruz Uribe, 1991;Domínguez Rodrigo, 1999;Selvaggio and Wilder, 2001;Brugal and Fosse, 2004;Coard, 2007;Egeland, 2008;Kuhn et al., 2010;Domíguez-Rodrigo and Pickering, 2010;Yravedra et al., 2011;Gidna et al., 2013;Arriaza et al., 2016). ...
... However, sometimes evidences are not conclusive enough. For instance, the dimensions of pits or scores have long been used distinguish between carnivores (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012), but these studies have not managed to properly isolate the marks produced by each carnivore, and have rather established carnivore groups according to their size (e.g. large carnivores like hyenas or lions, and small carnivores like foxes or intermediate-small felids). ...
Before sedimentation, bones are exposed to an important amount of biostratinomic taphonomic processes. One of them is related to the action of carnivores, which is reflected in conspicuous tooth marks, such as pits, scores, punctures or furrowing. Different carnivores damage bone assemblages differently. Thus, several researches have tried to identify carnivore agency based on different parameters such as skeletal profiles, tooth mark frequencies and dimensions, breakage patterns, or more recently, taphotypes. Here we propose a new methodology based on the analysis of tooth scores to determine the carnivore type involved in bone modification. For this purpose, we have built 3D models of several tooth scores produced by wolves, lions, jaguars, foxes and hyenas using photogrammetric techniques. These models were later analyzed by means of Geometric Morphometrics and multivariate statistics. We show that although there is a high degree of overlap in tooth mark morphology, the combined action of tooth score dimensions and morphology enables the identification of some of the tooth scores made by lions from those of the other carnivores with a higher degree of confidence than any other inter-carnivore comparison.
see link: authors.elsevier.com/a/1V7T6,rVDBJ-KE
... So far, studies based on pit dimensions have only managed to identify groups of carnivores by size, e.g. large carnivores like hyenas or lions, and small carnivores like foxes or intermediate-small felids (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012;Saladié, 2009;Saladié et al., 2013;Bello et al., 2015). Recent studies such as those by Pante et al. (2017) or Arriaza et al. (2017) present new methods for the identification of carnivore tooth marks. ...
... Previous analyses focused on the external morphology of tooth pits, particularly the length and breadth of marks, measurements that enable the classification of carnivores according to their body size (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009;Andrés et al., 2012). Our analysis not only addresses these axes, but it also accounts for variation in internal pit morphology, which helps differentiate carnivores, and also includes a more detailed morphological description of their external areas. ...
Hominins and carnivores have shared similar habitats in Africa, evolving as direct competitors for the same prey and spaces. The overlap of their ecological niches has raised an important debate on the role that carnivores played in archaeological site formation. Different analytical techniques differentiate the action of carnivores and humans and identify the type of carnivore involved of which tooth mark analyses are prominent. However, available taphonomic studies present some limitations given the great overlap among tooth mark sizes from different carnivores, enabling only the distinction of size groups, not specific carnivores. In this work, we use a new technique combining three-dimensional (3D) reconstruction and geometric morphometrics (GMM) of tooth pits on equid and bovid bone created by different carnivores (including crocodiles, hyenas, jaguars, lions and wolves) in controlled settings. The 3D methodology we present isolates and differentiates tooth marks generated by different carnivores. We also test the applicability of the technique to ascribe tooth pits recorded on some hominin specimens from FLK Zinj and FLKNN 3 (Bed I, Olduvai Gorge). The tooth marks on the Olduvai OH8 and OH35 hominin fossils, previously assigned to crocodiles show that while OH8 tooth pit morphology falls in the range created by crocodiles, the pits on OH35 can not be interpreted as crocodile-inflicted marks.
... Another critical question about a faunal assemblage such as this one involves which carnivores impacted the faunal remains. Many authors have attempted to correlate the sizes of bite marks with specific carnivore taxa, and the general consensus is that more conservative categories such as "small carnivore" or "mediumsized felid" provide the most accurate means of evaluating bite mark data (Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009). Using experimental data and models from these previous studies in conjunction with the known set of carnivore taxa in the vicinity of Sch€ oningen during the Middle Pleistocene makes it possible to hypothesize about the specific carnivore agents that impacted the archaeological remains. ...
... There is no correlation between bone density and % Survivorship (r s ¼ À0.019, p ¼ 0.849), so there is no evidence for density-mediated attrition in the assemblage. Coard, 2007), so specimens were divided into categories to reflect these structural differences (Table 9). Length to breadth ratios were not calculated because of variation found by different authors for the same taxa (e.g., Selvaggio and Wilder, 2001;Domínguez-Rodrigo and Piqueras, 2003). ...
This paper presents the faunal remains from the new excavation area at the Lower Paleolithic site of Schöningen. The focus of the study is on the southern extension of the main find horizon (Spear Horizon South), which includes the layer that yielded the famous Schöningen spears (13 II-4). Taxonomic data corroborate previous studies, that hominins primarily hunted Equus mosbachensis, a large Pleistocene horse. Equid body part representation at the site suggests that the animals were hunted and butchered locally. There is no evidence for density-mediated attrition in the assemblage. Weathering damage is uncommon, though there is ample evidence that carnivores had access to the bone. Carnivore bite sizes were measured and compared to experimental data provided by previous authors. Based on relationships between bite size and carnivore behavior and body size, we conclude that the primary modifying agents were large carnivores (i.e., wolves or saber-toothed cats). Previous studies show that carnivores often had secondary access to the remains, after hominins. Cut marks are commonly arranged haphazardly on the bones. This may indicate that multiple hominins participated in the butchery of horse skeletons, or that they were butchered over the course of hours or days. Cut marks on axial elements are more "orderly," which probably reflects the physical logistics of orienting one's body in relation to a large carcass. These data differ from sites formed by Middle and Upper Paleolithic hominins, which might suggest that in later times, a system of organized meat provisioning was already in place. Taken together, the faunal evidence from the Spear Horizon South indicates that late Lower Paleolithic hominins using the site understood the behaviors of different prey species, hunted socially to take down large game, and successfully competed with large carnivores on the landscape for primary access to ungulate remains.
... Likewise, search efforts were impacted by scavenging and scattering, such that 80.43% of 92 searchers had participated in a search that did not result in the recovery of a whole set of human remains, even with the assistance of cadaver dogs (13). Young et al. (13) have also shown that where SIOs and searchers have knowledge of scavenging in this region, it is subjective, undervalued, based on potentially incorrect anecdotal evidence, or limited due to a gap in region-and species-specific forensic and archaeological literature (14)(15)(16)(17)(18), as well as poor access to academic research. This gap in scavenger knowledge leaves investigators and police specialist searchers at a disadvantage when faced with a crime scene involving suspected scavenging. ...
... SIOs and specialist searchers should treat all scenes of scavenged human remains as crime scenes until criminal activity has been identified or eliminated because it is not within their expertise or role to analyze and interpret skeletal remains but is instead that of forensic anthropologists, forensic archaeologists, and pathologists (12,15,25,31,32). Moreover, a physical search by specialist searchers for scavenged remains may also reveal further forensic evidence that require forensic examination and re-assessment of the objectives and search methods of the physical search (12). ...
Forensic investigations involving animal scavenging of human remains require a physical search of the scene and surrounding areas. However, there is currently no standard procedure in the U.K. for physical searches of scavenged human remains. The Winthrop and grid search methods used by police specialist searchers for scavenged remains were examined through the use of mock red fox (Vulpes vulpes) scatter scenes. Forty-two police specialist searchers from two different regions within the U.K. were divided between those briefed and not briefed with fox-typical scavenging information. Briefing searchers with scavenging information significantly affected the recovery of scattered bones (χ(2) = 11.45, df = 1, p = 0.001). Searchers briefed with scavenging information were 2.05 times more likely to recover bones. Adaptions to search methods used by searchers were evident on a regional level, such that searchers more accustom to a peri-urban to rural region recovered a higher percentage of scattered bones (58.33%, n = 84).
... Rodent tooth marks were recently noticed on seeds of the late Eocene freshwater aquatic floating plant Stratiotes; this is the oldest direct evidence of seed predation by rodents (Collinson & Hooker, 2000). The pattern and tooth marks made by different species of carnivores is becoming increasingly common as research focus and application (see Miller, 1969;Haynes, 1983;Selvaggio & Wilder, 2001;Domínguez-Rodrigo & Piqueras, 2003;Pickering et al., 2004a;Coard, 2007;Pobiner, 2007). ...
... Identification of the identity of the predators using measurements such as intercanine distances, tooth mark metrics, and tooth morphology and dental configuration can be a useful tool in focusing predator identification efforts in wildlife conservation and forensics (e.g. Dayan et al., 1990;Pasitschniak-Arts & Messier 1995;Lyver, 2000;Anthony et al., 2006;Murmann et al., 2006;Coard 2007). The inter-tooth distance approach is occasionally taken in the fossil record as well. ...
This paper reviews the evidence for tooth marks made by sharks, crocodiles, dinosaurs, rodents, and especially mammalian carnivores on modern and fossil bones. The ecological and taphonomic information revealed in tooth marks, including: predator identity, prey preferences, and feeding behavior and ecology are discussed, and a compilation of metric measurements of taxon-specific modern and fossil mammalian carnivore tooth marks from the published literature is also provided. Some recommendations intended to improve the scope and scale of future tooth-damage research are also presented.
... We also investigated the practicality of attempting to identify predators via DNA left on carcasses, for example in saliva (see for example Blejwas et al. 2006), by DNA extracted from scats (see for example Berry et. al. 2007) and by forensic veterinary examination of carcasses (see for example Coard 2007). ...
... One line of evidence that has some potential to advance our understanding is the forensic examination of prey carcasses, both livestock and wildlife. Reliable post-mortem predator identification from a prey carcass requires discovering the carcass while relatively fresh and skinning the carcass to look for puncture wounds, damage to bones, and other signs that could provide information on tooth dimensions and structure, for example (MacKay 2005;Saunders et al. 1995;Coard 2007; Internet Center for Wildlife Damage Management 2012; Texas Natural Resources Server 2012). However, even this level of evidence cannot prove the identity of the predator. ...
Summary
A review of information pertaining to the possible presence in Victoria of a wild population of
unknown species of ‘big cat(s)’ was undertaken. There are no records of ‘big cats’ in official
zoological databases managed by Victorian Government agencies, despite considerable fauna
survey, research and monitoring in the State over many decades. However, there are many
thousands of reports of ‘big cats’ in the files of community cryptozoological groups and
individuals.
We classified the types of evidence that have been used to support claims of the existence of ‘big
cats’ in Victoria into three broad classes based on the veracity of the evidence they provide. Most
reports involve sightings of an animal or signs of the presence of a large, unknown animal and are
inconclusive without further physical corroborating evidence. The clear conviction of some
observers about what they saw does not materially increase the veracity of the identification. Other
reports involve the finding of dead and partially eaten livestock or wildlife where the volume of
flesh removed and manner in which the flesh is removed are claimed to be beyond the capabilities
of known predators in Victoria. There are two possible explanations for such cases – that an
unknown predator is responsible, or that our understanding of the behavioural repertoire of known
predators is inadequate. The latter explanation seems most prudent at this stage.
The most parsimonious explanation for many of the reported sightings is that they involve large,
feral individuals of the Domestic Cat Felis catus. However, some evidence cannot be dismissed
entirely, including preliminary DNA evidence, footprints and some behaviours that seem to be
outside the known behavioural repertoire of known predators in Victoria.
Obtaining unequivocal evidence for the presence of ‘big cats’ in Victoria would require an
organised and structured program aimed at collecting DNA samples from faecal material or prey
carcasses, or the opportunistic collection of a number of ‘big cat’ carcasses of proven provenance.
Any such specimens need to be presented to Museum Victoria for incorporation into the State’s
mammal specimen collection so that they can be properly curated and are available to researchers
in future. [Note that more than one specimen would be required because the presence of a single
individual is not evidence of a self-sustaining population].
... Likewise, search efforts were impacted by scavenging and scattering, such that 80.43% of 92 searchers had participated in a search that did not result in the recovery of a whole set of human remains, even with the assistance of cadaver dogs (13). Young et al. (13) have also shown that where SIOs and searchers have knowledge of scavenging in this region, it is subjective, undervalued, based on potentially incorrect anecdotal evidence, or limited due to a gap in region-and species-specific forensic and archaeological literature (14)(15)(16)(17)(18), as well as poor access to academic research. This gap in scavenger knowledge leaves investigators and police specialist searchers at a disadvantage when faced with a crime scene involving suspected scavenging. ...
... SIOs and specialist searchers should treat all scenes of scavenged human remains as crime scenes until criminal activity has been identified or eliminated because it is not within their expertise or role to analyze and interpret skeletal remains but is instead that of forensic anthropologists, forensic archaeologists, and pathologists (12,15,25,31,32). Moreover, a physical search by specialist searchers for scavenged remains may also reveal further forensic evidence that require forensic examination and re-assessment of the objectives and search methods of the physical search (12). ...
Scavenger-induced alteration to bone occurs while scavengers access soft tissue and during the scattering and re-scavenging of skeletal remains. Using bite mark, dimensional data to assist in the more accurate identification of a scavenger can improve interpretations of trauma and enhance search and recovery methods. This study analyzed bite marks produced on both dry and fresh surface deposited remains by wild and captive red fox (Vulpes vulpes) and Eurasian badger (Meles meles), as well as domestic dog (Canis familiaris). The bite marks produced by foxes were distinguishable from those made by badgers and dogs based on ranges of mean length and breadth of pits. The dimensional data of bite marks produced by badgers and dogs were less discernible. Bone modifications vary due to a variety of factors which must be considered, such as scavenger species-typical scavenging behavior, scavenger species' dentition, condition and deposition of remains, and environmental factors.
© 2015 American Academy of Forensic Sciences.
... These remains show damage of medicolegal interest, such as injuries from firearms or knives, signs of combustion, or scavenging activity, which complicate the identification of perimortem trauma (Colard et al. 2015). Existing studies in the scientific literature about manipulation of cadavers and human remains by animals are used as theoretical support when researchers describe marks left due to animal consumption (Coard 2007;Erkol & Hösükler 2018;Haglund et al. 1988Haglund et al. , 1989. ...
La intervención carroñera en restos humanos por parte de perros (Canis lupus familiaris) puede dificultar el proceso de identificación y el análisis de traumas perimortem. Se han publicado numerosos estudios científicos acerca de los daños provocados por perros en restos óseos, como consecuencia de consumo postmortem o de ataques letales. Sin embargo, escasean aquellos relativos al análisis de vestimentas asociadas a los restos humanos. El objetivo de esta investigación fue identificar patrones de daños causados por perros domésticos en tejidos textiles de uso común. Para ello, se utilizaron 45 sacos, de 20 × 30 cm cada uno, elaborados manualmente con tres tipos distintos de tejidos textiles (15 por textil): bistrech, mezclilla y poliéster, con una densidad de 40,84; 57,95 y 31,46 hilos por cm², respectivamente. La muestra canina consistió en 15 ejemplares de diferente tamaño, edad y sexo, procedentes de la “Fundación Chile Mestizo” de Santiago (Chile). Se analizó, mediante análisis de la varianza, la relación entre tipo de textil y presencia de daños y, posteriormente, se calculó la frecuencia de daños observados por tipo de textil. Para ello se utilizó el programa estadístico Minitab 19. Según los resultados obtenidos, se identificaron cuatro tipos de patrón: punción y masticación, presentes en el 62% y 75% de los casos, respectivamente; perforación y el daño de hole and tear, con una frecuencia del 91% de los casos analizados. Con respecto a la relación entre tipo de textil y daños observados, se observó que la densidad y el peso del tejido están directamente asociados con el tipo de daño presente.
... Recent results of neotaphonomic and experimental studies of carnivore modifications are applied in analyses of fossil faunal assemblages, and in spite of interpretive disagreements may provide evidence about the timing of hominin and carnivore access to animal carcasses (e.g., Binford, 1981;Blumenschine, 1988Blumenschine, , 1995Selvaggio, 1998;Pante et al., 2012). Another (and complementing) application has been to identify the carnivore taxa responsible for tooth-marking of fossil mammal bones (Haynes, 1983;Sobbe, 1990;Pickering et al., 2004;Coard, 2007;Burke, 2013aBurke, , 2013bYoung et al., 2015;Arriaza et al., 2017;Aramendi et al., 2017;Mondini, 2017;Yravedra et al., 2017;Koungoulos et al., 2018). These studies examined the effects that living carnivores have on bones of small, medium, and large mammals, but modern observations of modifications specifically to proboscidean bones are unusual. ...
The paper provides a guide for identifying carnivore effects on proboscidean bones, which may partially or wholly reduce analysts’ variability in reporting frequencies of carnivore modifications in fossil proboscidean assemblages.
... In archaeological context, dogs may shape faunal assemblages through bone destruction and consumption of refuse (Munson, 2000;Atici, 2006;Russell and Twiss, 2017, but see Horwitz, 1990). Tooth-marked bone can be used as evidence of bone consumption, but identifying the mammalian predator responsible, based on tooth mark size and shape, has been met with mixed results (Selvaggio and Wilder, 2001;Domı́nguez-Rodrigo and Piqueras, 2003;Coard, 2007;Delaney-Rivera et al., 2009;Andr� es et al., 2012;Parkinson et al., 2015). It would be particularly difficult to distinguish tooth marks made by two canids of similar size. ...
Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between 40,000 and 15,000 years ago, with identification of the earliest dogs debated. We predict that these earliest dogs (referred to herein as protodogs), while potentially difficult to distinguish morphologically from wolves, experienced behavioral shifts, including changes in diet. Specifically, protodogs may have consumed more bone and other less desirable scraps within human settlement areas. Here we apply Dental Microwear Texture Analysis (DMTA) to canids from the Gravettian site of P�redmostí (approx. 28,500 BP), which were previously assigned to the Paleolithic dog or Pleistocene wolf morphotypes. We test whether these groups separate out significantly by diet- related variation in microwear patterning. Results are consistent with differences in dietary breadth, with the Paleolithic dog morphotype showing evidence of greater durophagy than those assigned to the wolf morphotype. This supports the presence of two morphologically and behaviorally distinct canid types at this middle Upper Paleolithic site. Our primary goal here was to test whether these two morphotypes expressed notable differences in dietary behavior. However, in the context of a major Gravettian settlement, this may also support evidence of early stage dog domestication. Dental microwear is a behavioral signal that may appear generations before morphological changes are established in a population. It shows promise for distinguishing protodogs from wolves in the Pleistocene and domesticated dogs from wolves elsewhere in the archaeological record.
... The presence of tooth marks on leporid accumulations has been traditionally attributed to non-human predators. This assumption has changed over the past few years, with ethnoarchaeological and experimental research data showing that humans also make modifications during chewing (Coard 2007;Landt 2007;Lloveras et al. 2009b;Saladié et al. 2013;Sanchis Serra et al. 2011). However, identifying tooth marks caused by humans from other predators is not a straightforward matter, since tooth marks caused by human teeth are characterized by the removal of minimal to moderate amounts of cancellous bone tissue and are mostly associated with the fractures of long bone in order to remove bone marrow (Landt 2007;Sanchis Serra et al. 2011). ...
In Mediterranean Europe during the Pleistocene–Holocene, transition changes in the intensification of small prey exploitation by
humans are detected. In the NE of the Iberian Peninsula, these changes are mainly evidenced by an increase in the number of
rabbit remains, normally exceeding the 90% of the recovered animal specimens. The archaeological site of Balma del Gai (Moià,
Barcelona) provides one of the most significant archaeological records of this kind. Through zooarchaeological and taphonomic
analysis, our work’s aims are: to elucidate the human activity on leporids (rabbits and hares) in comparison to other predators, to
understand the different ways of handling and exploiting rabbit carcasses and to assess the importance of this small prey for the
Epipalaeolithic hunter–gatherers. Results on anatomical representation, breakage and bone surface modifications show that rabbit
remains recovered respond to an anthropogenic contribution. High proportions of thermo-altered bones, cut marks caused by
lithic tools and teeth marks are observed. All parameters indicate intense exploitation of rabbit fur and meat. This study shows
clear evidence of the importance of small prey for human subsistence during this period.
... The general consensus is that while no species can be identified reliably on the basis of tooth-mark size, marks from carnivores of different body weight classes [e.g. large (> 50 kg), medium (10-50 kg), and small (< 10 kg)] may be separated from each other (Andres et al., 2012;Dominguez-Rodrigo and Piqueras, 2003;Coard, 2007). Notably, there is a degree of overlap in medium-sized carnivore tooth-marks with those from large (Dominguez-Rodrigo and Piqueras, 2003) and especially small carnivores (Delaney-Rivera et al., 2009). ...
... Basically, bite marks caused by predatory animals include pits, punctures, scores, and furrows [33]. Pits are caused by collapsing of the bone under masticatory forces created by the tip of the teeth during chewing act, but they do not penetrate into the bone cortex [34][35][36][37]. Punctures are formed by greater force exerted by teeth on bone with resultant penetration of the tip of the teeth into cortical bone. ...
... Instead of Binford's system to understand the types of specific marks produced by scavenging predators, Haynes (1981 and1982) sought out to understand how entire carcasses were utilized. Instead of coding the types of modification present on the skeletal elements, Haynes (1982:275) Researchers in current carnivore modification studies have begun to ask how one could identify the specific predator that left the marks on the remains found at hominid sites in Africa (Coard 2007;Dominguez-Rodrigo and Piqueras 2003;Pickering, et al. 2004;Selvaggio and Wilder 2001). Arguments could be made that these researchers cannot discover the specific species consuming carcass materials from faunal remains. ...
... Other ancient taphonomic agents may also produce sub-rounded marks basically resulting from impact events due to lithic tool strikes (Shipman, 1981;Blumenschine and Selvaggio, 1988) or to batting the bone on to a stone or another type of anvil (an active percussion as Blasco et al., 2014 describe). Similarly, compressive forces exerted by tooth cusps during predation (Milner and Smith, 1989;Fern andez-Jalvo et al., 1999;Haglund et al., 1998;Domínguez-Rodrigo and Piqueras, 2003;Pickering et al., 2004;Coard, 2007;Andr es et al., 2012), or by penetration of sediment grains due to trampling (Andrews and Cook Dominguez-Rodrigo et al., 2009, 2010Marin-Monfort et al., 2014) may also mimic recent sub-rounded marks. ...
Different taphonomic processes throughout the history of a fossil assemblage may preserve, modify or destroy, particular palaeobiological traits, but these processes always increase taphonomic information of the past. Similarly, fossils are affected during later stages of taphonomic history, i.e. excavation, preparation, study and storage of fossils, known as sullegic and trephic phases. Tools used during excavation and preparation of fossils can damage them and produce marks on their surface. Some of these recent marks highly mimic taphonomic marks produced before excavation. Both modern and fossil marks lead to misinterpretations and erroneous conclusions when similarities are not clearly detected. In order to distinguish recent from ancient marks, several diagnostic criteria resulting from experimental work are described in this paper.
... Bien qu'insuffisantes prises isol?ment, les mesures des marques de morsure sont la m?thode la plus ? m?me d'?valuer la taille du pr?dateur (Coard, 2007;Delaney-Rivera et al., 2009;Dom?nguez-Rodrigo and Piqueras, 2003;Pickering et al., 2004;Selvaggio and Wilder, 2001). ...
... Actualistic data document the strong overlap of pit dimensions created by carnivores of different sizes, specifically tooth-marks on cortical bone [101][102][103]80,83,[85][86][87]. Even so, given most of these data, the presence on ThI94-UA28-7 of five pits larger than 4 mm in length and 2 mm in breadth and the size range and means of all pit dimensions indicate that they are attributable to a large carnivore (Fig 7A). ...
In many Middle Pleistocene sites, the co-occurrence of hominins with carnivores, who both contributed to faunal accumulations, suggests competition for resources as well as for living spaces. Despite this, there is very little evidence of direct interaction between them to-date. Recently, a human femoral diaphysis has been recognized in South-West of Casablanca (Morocco), in the locality called Thomas Quarry I. This site is famous for its Middle Pleistocene fossil hominins considered representatives of Homo rhodesiensis. The bone was discovered in Unit 4 of the Grotte à Hominidés (GH), dated to c. 500 ky and was associated with Acheulean artefacts and a rich mammalian fauna. Anatomically, it fits well within the group of known early Middle Pleistocene Homo, but its chief point of interest is that the diaphyseal ends display numerous tooth marks showing that it had been consumed shortly after death by a large carnivore, probably a hyena. This bone represents the first evidence of consumption of human remains by carnivores in the cave. Whether predated or scavenged, this chewed femur indicates that humans were a resource for carnivores, underlining their close relationships during the Middle Pleistocene in Atlantic Morocco.
... We took measurements of the dimensions of pits, punctures, scores and notches (maximal length and breadth) into account, as well as tissue location (cancellous bone or articular portions; cortical or median diaphysis; thin cortical bone or diaphysis extremity). While insufficient is taken alone, measuring the toothmarks (especially maximal dimensions) is the most accurate method for establishing the body size of the predator (Campmas & Beauval, 2008;Coard, 2007;Delaney-Rivera et al., 2009;Domínguez-Rodrigo & Piqueras, 2003;Faith, 2007;Pickering et al., 2004;Selvaggio & Wilder, 2001). We also measured the coprolites (length, breadth and thickness). ...
Study of faunal series resulting from recent excavations in two caves in North Atlantic Morocco (Grotte à Hominidés - GH - and Grotte des Rhinocéros - GDR - at Thomas I and Oulad Hamida 1 quarries, Casablanca) has yielded new evidence concerning the gathering and processing of ungulates carcasses
during the Middle Pleistocene in this part of North Africa. Preliminary taphonomic analysis of the
macrofauna indicates that the carcasses were mainly introduced in the caves by carnivores. Additionally,
marks generated by porcupines also occur. Dimensions and morphologies of tooth-marks and coprolites
suggest that carnivores of different sizes (mainly middle-sized canids, hyenids and felids), as well as
porcupines, used the cave. Cut-marks on the bones are absent at GH and scarce at GDR, despite their
association with lithic artefacts and human fossils. This raises the question of the relationship between
hominins and other competitors in these caves. The recurring question is to determine the modalities of
niche partitioning by the various predators and/or carrion-eaters as well as the mode of introduction of
artefacts and human remains.
... Unfortunately, there has thus far been little effort to present these data within the peerreviewed literature. Coard (2007), however, showed that bite marks present on Welsh sheep bones correspond to the dentition of a 'medium-sized felid' and hence provide support for the existence of exotic felids in the British fauna. ...
The alleged presence of non-native felid species in the British countryside – popularly, though in part erroneously, known as ‘British big cats’ or ‘alien big cats’ – is a long-standing and controversial topic, perennially of interest to both the mass media and amateur naturalists, and with little apparent acceptance from the technical zoological community. Nevertheless, a number of carcasses and captured live specimens have demonstrated the occasional presence within the region of escapees that potentially explain at least some ‘British big cat’ eyewitness records. We report here the existence of a probable Canada lynx, Lynx canadensis, shot in Newton Abbot, Devon, England, in or prior to 1903, and then accessioned to Bristol Museum and Art Gallery. The specimen (represented by extensive skeletal material and a stuffed taxidermy mount) is Bobcat-like in some respects but is identified as a Canada lynx on the basis of skeletal morphology with a high degree of support; attempts to extract DNA were unsuccessful. Stable strontium isotope analysis supports either a recent introduction from western Canada or long-term acclimation to the local area of Devon where it was collected. Although the specimen was undoubtedly an ‘alien’ (an escapee or release from a collection), it is significant as material evidence in demonstrating, for the first time, the presence of a wild-caught, feral, exotic felid dating to the early years of the twentieth century.
... Yet whoever I had encountered didn't behave in a canine manner – there was no noise, no confrontation or any engagement with Max. I put all of this to the back of my mind until I read Mrs Marshall's testimony in the following week's papers, but it wasn't until 2007 that I began to think seriously about what I (and Mrs Marshall and my other informants ) had experienced, when Ros Coard told me about her own findings (Coard 2007). Coard's research, which matched tooth pit data on carcasses with dental records for a range of carnivores, demonstrated that some of the animals examined had indeed been predated on by large felids. ...
The presence of non-endemic ‘Alien Big Cats’ (ABCs) in Britain is an oft debated topic in the press and the public sphere, especially in rural areas where agriculture is the dominant industry. Based on seven years of fieldwork in a farming community in rural Ceredigion, West Wales, this paper considers the ‘Big Cat’ phenomenon from the perspectives of informants for whom inexplicable livestock losses must be accounted for and ‘rationalised’. Moreover, in this ethnographic context, ABCs, like other transgressive non-human animals, have become particularly powerful metaphors for the humans who encounter them.
... These methods are of particular interest to establish the sequence of carcass access (Hominid only, Carnivore only, Hominid to Carnivore or Carnivore to Hominid to Carnivore), that has also been used to interpret the type of carcass acquisition by hominids (e.g. Blumenschine & Selvaggio, 1988; Selvaggio, 1994; Blumenschine, 1995; Capaldo, 1995 Capaldo, , 1997 Capaldo, , 1998 Domínguez-Rodrigo, 1997 Selvaggio, 1998; Selvaggio & Wilder, 2001;); (4) The metric values of tooth marks are used for actor identification (Lyman, 1994; Selvaggio, 1994; Andrews & Fernández Jalvo, 1997; Selvaggio & Wilder, 2001; Domínguez-Rodrigo & Piqueras, 2003; Coard, 2007; Delaney-Rivera et al., 2009). The results of tooth mark measurements between various modern carnivores show significant overlap. ...
Actualism has been a fundamental tool in taphonomy. The knowledge of accumulation patterning of modern faunal allows us to interpret the activity of different actors in the archaeological record and to reconstruct the behaviour of preterit animals and humans in which we are interested. However, until now, there are few works that include bone modifications made by bears amongst those made by carnivores.
Most data about bone modifications made by bears have been obtained from the archaeo-palaeontological record. In most of these assemblages, the presence of bears is related to their period of hibernation. Therefore, in these contexts, the changes documented on recovered bear bones are associated only with cannibalism.
In this paper, we present an actualistic study about modifications on bones made by modern brown bears. These animals can cause damage similar to those produced by other large carnivores. Generally, bear activity leaves slight damage, mainly on large-sized animal bones. However, on bones of small-sized animals and those of greater fragility, the bears can produce abundant damage. Though not usual, bears can break long bones and consume the complete epiphysis. This study suggests that bears have the potential to be agents of bone modification in fossil assemblages. Consequently, they should be considered as a possible agent of modification of faunal remains in the fossil record. Copyright
Animal scavenging by vertebrates can significantly alter human bodies and their deposition
site. For instance, vertebrate animals can cause postmortem modification to a body, alter perimortem
trauma, influence decomposition rates, disarticulate and scatter body parts or evidence, and affect
the identification of the deceased. Animal scavenging is a relatively common occurrence in forensic
investigations. Even so, studies on the subject are scattered and rare, with most focussing on
geographical areas outside of Europe. For that reason, we intend to collate the literature to provide an
account of forensically relevant vertebrate scavengers in Europe, their impacts on human remains, and
their implications for forensic investigations. Here, we provide an overview of forensic aspects where
the knowledge of animal scavenging is crucial, as well as an account of potential scavengers of human
remains in Europe and their typical alterations to soft tissue and, in particular, to bones. In addition,
we are the first to provide a guide for forensic practitioners to identify the presence of vertebrate
scavenging and subsequently inform outdoor search strategies for affected human remains.
This chapter focuses on the most common terrestrial mammalian and avian scavengers found in forensic cases in North America and Europe. It presents the general characteristics of scavenger species' scavenging behaviours and patterns, as well as damage and modifications to soft tissue and bones. Canid scavengers are widely discussed in the forensic literature as scavengers of human remains, in particular coyote, wolf, and domestic dog. Avian scavenger species commonly discussed in forensic and taphonomic studies, such as vulture and magpie. Procyonid, didelphid, phalangerid and peramelid scavengers have been recognised in forensic and taphonomic studies as scavengers of smallto medium-sized animal carrion. Crime scene investigations have been aided by the use of actualistic methods in forensic taphonomic research into the scavenging of human remains. Actualistic methods and bite mark analyses have been utilised in research and crime scene investigations in order to identify scavengers, interpret species-typical scavenging behaviour and scatter patterns.
Two children, two landowners and two zoologists were amongst visitors to the big cats stand at the 2012 Stroud Festival of Nature. Their comments and questions, summarised here, reflect different views on the parallel universe presented by big cats in Britain's landscape.
Scavenger-induced alteration to bone occurs whilst scavengers access soft tissue and during the scattering and re-scavenging of skeletal remains. Using bite mark dimensional data to assist in the more accurate identification of a scavenger can improve interpretations of trauma and enhance search and recovery methods. This study analyzed bite marks produced on both dry and fresh surface deposited remains by wild and captive red fox (Vulpes vulpes) and Eurasian badger (Meles meles), as well as domestic dog (Canis familiaris). The bite marks produced by foxes were distinguishable from those made by badgers and dogs based on ranges of mean length and breadth of pits. The dimensional data of bite marks produced by badgers and dogs were less discernible. Bone modifications vary due to a variety of factors which must be considered, such as scavenger species-typical scavenging behavior, scavenger species’ dentition, condition and deposition of remains, and environmental factors.
Vertebrate scavengers can modify surface deposited human remains which can hinder
forensic investigations. The effects of such scavenging vary between species and regions.
Published research into the effects of the scavenging of human remains is dominated by
work from North America with few studies covering Northwestern Europe. Forensic
investigators in Northwestern Europe are often left questioning on a basic level as to which
scavengers are active and how they might affect human remains. This paper presents the
results of a field study utilizing deer (Cervus nippon; Capreolus capreolus) as surface
deposits observed by motion detection cameras in a British woodland. The most common
avian and rodent scavenger species recorded included the buzzard (buteo buteo), carrion
crow (Corvus corone), wood mouse (Apodemus sylvaticus) and gray squirrel (Sciurus
carolinensis). The scavenging behaviors observed were affected by seasonality, rates of
decomposition and insect activity. Scavenging by buzzards, unlike carrion crows, was most
frequent during fall to winter and prior to insect activity. Overall, avian scavengers modified
and scavenged soft tissue. Rodents scavenged both fresh and skeletonised remains with
gray squirrels only scavenging skeletal remains. Wood mice were most active in winter and
scavenged both soft tissue and bone.
Teeth are generally the best-preserved elements among mammal fossil remains and are highly diagnostic characters. Consequently, much mammalian paleontological, systematic, and evolutionary research focuses on teeth, so it is important to understand how they vary and covary with other characters. Dental traits within populations of carnivores appear to be more variable than cranial traits, a pattern that results only partly from their usually smaller size. Furthermore, dental traits, although highly correlated with one another, are not highly correlated with cranial traits, which are also highly correlated with one another. Thus, teeth and cranial bones may be subject to quite different selective pressures and genetic/developmental constraints and may sug- gest different microevolutionary scenarios. Vestigial teeth show significantly greater variability than expected, reflecting the absence of stabilizing selection.
Livestock depredation by the snow leopard, Uncia
uncia, and the wolf, Canis
lupus, has resulted in a human-wildlife conflict that hinders the conservation of these globally-threatened species throughout their range. This paper analyses the alleged economic loss due to livestock depredation by these carnivores, and the retaliatory responses of an agro-pastoral community around Kibber Wildlife Sanctuary in the Indian trans-Himalaya. The three villages studied (80 households) attributed a total of 189 livestock deaths (18% of the livestock holding) over a period of 18 months to wild predators, and this would amount to a loss per household equivalent to half the average annual per capita income. The financial compensation received by the villagers from the Government amounted to 3% of the perceived annual loss. Recent intensification of the conflict seems related to a 37.7% increase in livestock holding in the last decade. Villagers have been killing the wolf, though apparently not the snow leopard. A self-financed compensation scheme, and modification of existing livestock pens are suggested as area-specific short-term measures to reduce the conflict. The need to address the problem of increasing livestock holding in the long run is emphasized.
Public attitudes towards snow leopard Panthera uncia predation of domestic livestock were investigated by a questionnaire survey of four villages in snow leopard habitat within the Annapurna Conservation Area, Nepal. Most local inhabitants were subsistence farmers, many dependent upon yaks, oxen, horses and goats, with an average livestock holding of 26.6 animals per household. Reported losses to snow leopards averaged 0.6 and 0.7 animals per household in two years of study, constituting 2.6% of total stockholding but representing in monetary terms almost a quarter of the average annual Nepali national per capita income. Local people held strongly negative attitudes towards snow leopards and most suggested that total extermination of leopards was the only acceptable solution to the predation problem. Snow leopards were reported to be killed by herdsmen in defence of their livestock. The long-term success of snow leopard conservation programmes may depend upon the satisfactory resolution of the predation conflict. Some possible ways of reducing predation losses are also discussed.
Models of Plio-Pleistocene hominid behavioral ecology often emphasize competition with large carnivores. This paper describes competition between modern humans and large carnivores in rural Uganda, including active, confrontational scavenging of carnivore kills by humans and carnivore attacks on humans. Information gathered from Ugandan Game Department archives (1923-1994) reveals that twentieth-century agropastoralists regularly tried to scavenge from leopard (Panthera pardus) and lion (Panthera leo) kills, and that these large carnivores have preyed on hundreds of humans in Uganda over the past several decades. Men were most often targets of carnivore attack, particularly while engaged in hunting-related activities. However attacks on men were less often lethal than attacks on women and children. Analyses show that lion attacks were more dangerous than leopard attacks. These data support recent contentions that hominids armed with even simple weapons can succeed in active, confrontational scavenging by chasing carnivores from kills. Hominids sharing East African habitats with large carnivores may have been regularly subject to attack.
The current extinction of many of Earth's large terrestrial carnivores has left some extant prey species lacking knowledge
about contemporary predators, a situation roughly parallel to that 10,000 to 50,000 years ago, when naı̈ve animals first encountered
colonizing human hunters. Along present-day carnivore recolonization fronts, brown (also called grizzly) bears killed predator-naı̈ve
adult moose at disproportionately high rates in Scandinavia, and moose mothers who lost juveniles to recolonizing wolves in
North America's Yellowstone region developed hypersensitivity to wolf howls. Although prey that had been unfamiliar with dangerous
predators for as few as 50 to 130 years were highly vulnerable to initial encounters, behavioral adjustments to reduce predation
transpired within a single generation. The fact that at least one prey species quickly learns to be wary of restored carnivores
should negate fears about localized prey extinction.
Population density in plants and animals is thought to scale with size as a result of mass-related energy requirements. Variation
in resources, however, naturally limits population density and may alter expected scaling patterns. We develop and test a
general model for variation within and between species in population density across the order Carnivora. We find that 10,000
kilograms of prey supports about 90 kilograms of a given species of carnivore, irrespective of body mass, and that the ratio
of carnivore number to prey biomass scales to the reciprocal of carnivore mass. Using mass-specific equations of prey productivity,
we show that carnivore number per unit prey productivity scales to carnivore mass near –0.75, and that the scaling rule can
predict population density across more than three orders of magnitude. The relationship provides a basis for identifying declining
carnivore species that require conservation measures.
From a detailed analysis of published and unpublished sources, we constructed a digitized three-dimensional, stratigraphically-controlled excavation grid of Zhoukoudian Locality 1 in order to assess the spatial relationships of the excavated materials. All 15 fossil Homo erectus loci were mapped on the grid. Meter cubes were used in excavation starting in 1934, and Loci H through O, established between 1934 and 1937, were mapped to within 1 m(3)vertical and horizontal provenience. Loci A through G, established between 1921 and 1933, were excavated in the northernmost part of Locality 1 by unmapped quarrying, but their stratigraphic levels were recorded. We could localize Loci A through G on the grid system by utilizing locations of remaining walls, stratigraphic sections, excavation reports, excavation maps, and photographs. Loci contained skeletal elements of Homo erectus individuals scattered over areas of the cave floor of up to 9 m in diameter. Scoring of taphonomic damage on the Homo erectus sample, as observed on casts and originals, demonstrates that 67% of the hominid sample shows bite marks or other modifications ascribed to large mammalian carnivores, particularly the large Pleistocene cave hyena, Pachycrocuta brevirostris. Virtually all of the remaining Homo erectus skeletal assemblage shows breakage consistent with this taphonomic pattern of fragmentation. Bioturbation by digging carnivores is the most likely explanation for a fragment of Homo erectus Skull XI discovered 1 m below its other conjoined portions in Locus L. Carbon on all the Homo erectus fossils from Locus G, a circumscribed area of 1-meter diameter, earlier taken to indicate burning, cooking, and cannibalism, is here interpreted as detrital carbon deposited under water, perhaps the result of hyaenid caching behavior. Locus G records the close stratigraphic and horizontal association of stone artifacts with Homo erectus and other vertebrate skeletal elements, an association that is seen at other loci as well. Layer 4 of the excavation contains equid cranial bone previously interpreted to have been burned while fresh. We here document that Locus B Homo erectus, including Skull I, is stratigraphically associated with this evidence, but at some 10-12 m distance. Even though the presence of wood-stoked fires and hearths is not supported by geochemical results, evidence of fire at Locality 1 in the form of burned bone is confirmed. Contextual relationships of fossil skeletal elements, relationships of carnivore damage and stone tool cutmarks on bone, and evidence of the burning of fresh bone associated with Homo erectus and stone tools support a model of transient hominid scavenging aided by the use of fire at the large hyenid den that became Zhoukoudian Locality 1. Although the original excavation catalogue from Locality 1, as well as a significant number of fossils and stone artifacts, were lost during World War II, catalogue numbers on the many surviving specimens can be used to locate fossils and artifacts within the three-dimensional grid provided in this paper.
It has been hypothesised that leopards were significant contributors to the bone accumulations of the Plio-Pleistocene hominid-bearing caves of South Africa. Interpretations of leopard activity in these fossil caves were previously based upon reports of modern leopard behaviour in areas of southern Africa that were lacking in caves. In 1991 a leopard lair with an accompanying bone accumulation was discovered in a dolomitic cave on the John Nash Nature Reserve, South Africa. All of the bones in this cave could be unambiguously attributed to the activity of one individual leopard over a 1-year period. The resulting bone assemblage indicates that, when available, leopards will preferentially utilise the deep recesses of caves to the exclusion of trees when feeding, and that the size of prey leopards are capable of capturing, killing and transporting has previously been underestimated. The implications this may have for understanding the accumulation of fossils in the hominid-bearing caves of South Africa are that bones derived form leopards consuming prey in trees probably did not contribute significantly to the assemblages, and further that it is not necessary to invoke sabre-tooth cat involvement for the larger animals found in these assemblages. This modern cave probably represents a more appropriate model for the accumulation of bones in the fossil caves of the Sterkfontein Valley, and the assemblage is being continually monitored to view any and all taphonomic alterations that are occurring.
Animal populations undergo repetitive cycles of rising and falling numbers. In a Perspective,
Ranta and colleagues
discuss the value of time-series analyses for examining the changing dynamics of animal populations. A large time series gathered
by the Hudson Bay Company based on the fur trade has provided invaluable data on the rise and fall of the Canadian lynx population
from 1821 to the present. Analysis of this time series reveals that climate as well as factors influencing birth and death
rates are important in regulating the lynx population (
Stenseth
et al.). In a separate study, mathematical modeling combined with fieldwork revealed the importance of predators in determining
the population fluctuations of the southern pine beetle (
Turchin
et al.)
Information on the number of carnivore taxa that were involved with archaeological bone assemblages is pertinent to questions of site formation, hominid and carnivore competition for carcasses and the sequence of hominid and carnivore activity at sites. A majority of early archaeological bone assemblages bear evidence that both hominids and carnivores removed flesh and/or marrow from the bones. Whether flesh specialists (felids) or bone-crunchers (hyaenas), or both, fed upon the carcasses is crucial for deciphering the timing of hominid involvement with the assemblages. Here we present an initial attempt to differentiate the tooth mark signature inflicted on bones by a single carnivore species versus multiple carnivore taxa. Quantitative data on carnivore tooth pits, those resembling a tooth crown or a cusp, are presented for two characteristics: the area of the marks in millimetres, and the shape as determined by the ratio of the major axis to the minor axis of the mark. Tooth pits from bones modified by extant East African carnivores and latex impressions of tooth pits from extinct carnivore species are compared to those in the FLK Zinjanthropus bone assemblage. Data on tooth mark shape indicate greater variability in theZinj sample than is exhibited by any individual extant or extinct carnivore species in the comparative sample. Data on tooth mark area demonstrate that bone density is related to the size of marks. Taken together, these data support the inference that felids defleshed bones in the Zinj assemblage and that hyaenas had final access to any grease or tissues that remained.
Tooth marks on bones have been used as a proof of carnivore involvement in carcass modification in archaeological assemblages. Recognition of the array of potential carnivores that may intervene in the consumption of carcass elements accumulated at archaeological sites may condition the way archaeologists reconstruct hominid–carnivore interaction and resource availability for both types of taphonomic agents. The development of techniques aimed at discerning carnivore taxa according to tooth mark location and size has proven problematic so far. The present work introduces new information, based on the use of tooth pit size, to determine the types of carnivores that have modified bone surfaces. It is concluded that tooth marks alone cannot be used to differentiate among specific taxa, unless the analysis of tooth pits is carried out taking into account their distribution and ranges of variation in large samples, together with other variables, such as the location of tooth marks according to bone section and element, and the anatomical distribution of furrowing. Even so, the attribution of specific bone damage to determined carnivores can only be confidently made when comparing small-sized versus large-sized carnivores.
1. The recent recovery of the wolf in southern Europe has not yet removed the risk of local extinction. Wolf populations are fragmented and often comprise fewer than 500 individuals. In North America, northern and eastern Europe, wolves feed mainly on wild herbivores. In southern Europe, this canid has apparently adapted to feed also on fruit, rubbish, livestock, small and medium-size mammals. 2. The main conservation problem lies with predation on domestic ungulates, which leads to extensive killing of wolves. The reintroduction of wild large herbivores has been advocated as a means of reducing attacks on livestock, but predation on the latter may remain high if domestic ungulates are locally abundant. 3. Our synthesis of 15 studies, published in the last 15 years, on food habits of the wolf in southern Europe, has shown that ungulates have been the main diet component overall. A significant inverse correlation was found between the occurrence (%) of wild and domestic ungulates in the diet. The presence of relatively few wild ungulate species was necessary to reduce predation on livestock. 4. Selection of wild and domestic ungulate prey was influenced mainly by their local abundance, but also by their accessibility. Feeding dependence on rubbish was local and rare. In Italy, the consumption of rubbish/fruit and that of ungulates was significantly negatively correlated. Diet breadth increased as the presence of large prey in the diet decreased. 5. The simultaneous reintroduction of several wild ungulate species is likely to reduce predation on livestock and may prove to be one of the most effective conservation measures.
Summary • In regions where sheep are kept in fenced pastures and do not graze unattended in carnivore habitats, sheep losses vary greatly between sites and livestock farms. To assess the factors that may predispose farms to lynx predation in the French Jura, we compared sheep availability and environmental characteristics between pastures with and without attacks in a 1800-km2 study area. Nine lynx were radio-tracked in the same area for a total of 21 lynx years to estimate individual killing rates on sheep and to identify possible habitual livestock killers. • Depending on individual and year, lynx predation rate on sheep within lynx home ranges varied between 0 and 12·4 attacks 100 days−1. Predation rate on sheep was not related to sheep abundance nor sheep dispersion in lynx home ranges. Two individuals became habitual sheep killers during, respectively, their third and fourth year of monitoring. Other lynx that had access to the same flocks were only occasional sheep killers. No obvious causal factor (e.g. sex, reproductive status, physical debilitation) explained the differential individual propensity for lynx to kill livestock. • We found no difference in sheep availability between pastures with and without attacks, but strong differences in their environmental characteristics. In only 5·1% of 98 pastures > 250 m from a forest were sheep attacked by lynx. In 228 pastures adjacent or connected to forests by cover, 39·1% sustained attacks on sheep by lynx (P P P P = 0·01) and the presence of attacked pastures in their vicinity (P = 0·03). • These results suggests that lynx damage locally can be explained by a predictable set of habitat features that expose sheep on some pastures to risk, and by an unpredictable event, i.e. an individual developing regular predation on sheep. • In grazing systems like the Jura, where unattended sheep are distributed patchily and individual problem lynx may appear, removing lynx or lowering density without differentiating individuals will be insufficient to limit conflicts. Selective removals could temporarily reduce predation but the site effect implies that durable management can arise only through improved shepherding. This might include guard dogs in the few local sites at risk and providing shelter for sheep at night when attacks are on the increase.
ABSTRACT1. Literature on the wolf Canis lupus, brown bear Ursus arctos and lynx Lynx lynx is reviewed to determine if sufficient semi-natural habitat exists in the UK for a viable population of any of these species and to assess the potential risks to human safety, livestock and economically valuable wildlife. Public attitudes to the recovery and reintroduction of some other mammals are also briefly reviewed.2. The large home range sizes and low population densities of large carnivores mean that the Scottish Highlands is the only UK region with the potential to support a viable population. Human population density is also lower in the Highlands and the density of wild ungulate prey higher than in many parts of Europe where large carnivores survive.3. Attacks on people have been recorded in Europe for healthy bears and for rabid bears and wolves but there are no reports of attacks by lynx. Bears are more carnivorous in the north of their range than in the south and although wild mammals seldom appear to be important prey serious predation of livestock can occur. Livestock predation is also reported for the wolf and the lynx but they appear to prefer wild prey if available. However, mass kills of up to 100 or more sheep are occasionally recorded for wolves.4. Attitudes to reintroductions and carnivores generally tend to be favourable amongst the general public, but negative amongst those most likely to be adversely affected. Fears for human safety and significant livestock predation with bears and wolves, respectively, suggest that reintroduction of these species is unlikely to be acceptable in the foreseeable future. Reintroduction of the lynx may be feasible but habitat suitability and potential impact on vulnerable native wildlife need to be assessed. Socio-economic and legal issues also need to be addressed before such a reintroduction is considered.
Competing explanations of early human behavior concerning animal carcass acquisition and exploitation are currently some of the most debated topics in the study of human evolution. Various hypotheses depict hominids as either hunters and flesh-eaters, or as scavengers who mainly consumed marrow and brains. One of the main arguments advanced to support the scavenging hypothesis is that flesh-bearing medium-sized carcasses (weighing between 150 kg and 350 kg) at early sites could have been obtained from large felid kills. This paper presents the results of a preliminary study, in which I have analyzed lion-killed carcasses with respect to the availability and disposal of flesh and conspicuous carnivore-inflicted bone damage patterns, so as to have a reference that can be applied both to archaeofaunas and to actualistic experiments that try to model early human behavior. Bone damage made by lions overlaps the damage patterns caused by other carnivores, such as canids and hyenids, although it is not as intense. Scraps of flesh available after consumption are rare and show a typical anatomical distribution. The scavenging hypothesis is thus testable by comparing the distribution of cut marks on fossil archaeofaunas to the location of flesh in lions' kills. Comparisons between carcasses in different environments show that scraps of flesh can be obtained in open habitats. Carcasses consumed by lions in closed habitats are flesh depleted. The application of this referential framework to archaeological bone assemblages can help to identify hominid foraging strategies, and indirectly, trophic dynamics on savannas: scavenging in open habitats is only feasible in wet savannas with a slightly marked seasonality and lack of migratory biomass [M. Tappen (1992) Ph.D. Thesis, Dept. Anthropology, Harvard Univ., Cambridge, MA]; scavenging in closed habitats can only be made in semi-arid seasonal savannas whose biomass is subjected to migratory processes [R.J. Blumenschine (1986) B.A.R. Int. Ser. 283, Oxford].
The authors report a fatal case of a Persian leopard (Panthera pardus) attack in an animal sanctuary in Oklahoma. The victim was a 53-year-old Costa Rican woman who was attempting to feed the animal when she was attacked and killed. Autopsy, radiography, fingerprint analysis, microbiologic cultures, and dental impressions were used to evaluate the case. These simple techniques can be applied to similar cases involving wild and domestic animal attacks.
In addition to other methods for conservation of bite mark evidence, preservation of actual skin from deceased victims is often suggested. This study was undertaken to analyze the dimensional stability of such specimens. Utilizing a prefabricated template, marks approximating "bites" were made in postmortem skin of Miniature Hanford pigs, producing imprints with distinct margins and indentations. Tissue samples were stored in 10% formalin after affixing an acrylic support ring with cyanoacrylate adhesive and sutures. Measurements of the six tooth mark analogues and cross-arch dimensions were taken at intervals of up to 38 days. Data from these measurements indicate a wide range of amount and type of distortion in preserved tissue. Although some samples were dimensionally stable, there was both contraction and expansion of bite mark specimens, even within individual skin samples. It appears that standard techniques for storage and preservation of bite mark samples will not produce reliable dimensional accuracy.
There are many cases where animal populations are affected by predators and resources in terrestrial ecosystems, but the factors that determine when one or the other predominates remain poorly understood. Here we show, using 40 years of data from the highly diverse mammal community of the Serengeti ecosystem, East Africa, that the primary cause of mortality for adults of a particular species is determined by two factors--the species diversity of both the predators and prey and the body size of that prey species relative to other prey and predators. Small ungulates in Serengeti are exposed to more predators, owing to opportunistic predation, than are larger ungulates; they also suffer greater predation rates, and experience strong predation pressure. A threshold occurs at prey body sizes of approximately 150 kg, above which ungulate species have few natural predators and exhibit food limitation. Thus, biodiversity allows both predation (top-down) and resource limitation (bottom-up) to act simultaneously to affect herbivore populations. This result may apply generally in systems where there is a diversity of predators and prey.
Mammals in Carmarthenshire
- A Lucas
Lucas, A., 1997. Mammals in Carmarthenshire. Andrew Lucas, Dolgellau.
Snow leopard Panthera uncia preda-tion of livestock: an assessment of local perceptions in the Annapurna conservation area Nepal A tale of big game and small bugs Determination of predators responsible for killing small livestock
- M K Oli
- I R Taylor
- E Rogers
- E Ranta
- V Kaitala
- P Lundberg
Oli, M.K., Taylor, I.R., Rogers, E., 1994. Snow leopard Panthera uncia preda-tion of livestock: an assessment of local perceptions in the Annapurna conservation area Nepal. Biological Conservation 68, 63e68. Ranta, E., Kaitala, V., Lundberg, P., 1999. A tale of big game and small bugs. Science 285, 1022e1023. Roberts, D.H., 1986. Determination of predators responsible for killing small livestock. South African Journal of Wildlife Research 16 (4), 150e152.
Analysis of distortion in preserved bite mark skin), 573e576. Fig. 3. Measurement points for the carnassial cusp distances. The tooth shown is the upper carnassial of puma
- B R Rothwell
- M S D Thien
- A V Thien
Rothwell, B.R., Thien, M.S.D., Thien, A.V., 2001. Analysis of distortion in preserved bite mark skin. Journal of Forensic Science 46 (3), 573e576. Fig. 3. Measurement points for the carnassial cusp distances. The tooth shown is the upper carnassial of puma. R. Coard / Journal of Archaeological Science 34 (2007) 1677e1684
A review of wolf predation in southern Europe: Does the wolf prefer wild prey to livestock? Livestock predation by large carnivores in the Indian trans-Himalaya: conflict perceptions and conservation prospects
- A Meriggi
- S Lovari
Meriggi, A., Lovari, S., 1996. A review of wolf predation in southern Europe: Does the wolf prefer wild prey to livestock? Journal of Applied Ecology 33, 1561e1571. Mishra, C., 1997. Livestock predation by large carnivores in the Indian trans-Himalaya: conflict perceptions and conservation prospects. Environmental Conservation 24 (4), 338e343.