Nonhuman Animal Paleopathology—Are We so Different?

Chapter 23

Nonhuman Animal Paleopathology—Are We so Different? Richard Thomas School of Archaeology and Ancient History, University of Leicester, Leicester, United Kingdom

Humans are a kind of animal that (like all kinds of animal) has been and continues to be profoundly reshaped by its interactions with other kinds of animals . . .. All history is animal history, in a sense. (Benson, 2011, p. 5).

. . .it is quite certain that if we restrict our observations of the processes of disease as they occur in man, our notion of them would be as crude as if we attempted to form conclusions as to his zoological position without reference to other species of animals. (Bland Sutton, 1890, pp. 12 13)

INTRODUCTION While the study of pathology in human skeletal remains has a long and illustrious history with established and increasingly refined theoretical and methodological foundations (e.g., Buikstra and Roberts, 2012), the same cannot be said for nonhuman (animal) paleopathology. The value of studying disease and injury in the skeletal remains of animals in the past was recognized by the early 20th century (Moodie, 1923a,b; Wintemberg, 1919); however, the subsequent development of the discipline was sporadic (Thomas, 2012). As recently as 2000, zooarcheological paleopathology was characterized as “an inchoate discipline studied by a relatively small number of analysts” (O’Connor, 2000: 98). Issues that complicate the pursuit of zooarcheological paleopathology have been explored in previous reviews (e.g., Bendrey, 2014; Siegel, 1976; Shaffer and Baker, 1997; Thomas, 2012; Thomas and Mainland, 2005; Upex and Dobney, 2012; Vann and Thomas, 2006). In brief, these can be grouped into three, interrelated problem areas:

1. Complications arising from the nature of zooarcheological material: most animal bone assemblages are comprised of disarticulated and fragmentary remains, which makes it difficult to place lesions within their biological context and to undertake differential diagnoses; the low frequency of pathological bones per site. 2. Inadequate clinical foundation: the etiology and pathogenesis of many lesions observed in archeological fauna are not observed clinically because the life expectancy of domestic livestock is considerably shorter in modern industrialized farming systems than it was in the past; the skeleton is examined less systematically than soft tissues during necropsy; many of the lesions observed in the zooarcheological record are considered incidental by veterinary pathologists because they do not affect animal health (or, critically, animal productivity) significantly, and thus feature rarely in veterinary literature; an absence of clinical evidence means that separating pathology from “normal” biological variation/adaptation can be difficult and may lead to speculative interpretation. 3. Complications arising from disciplinary practices: unsystematic recording, with an emphasis on diagnosis (over description) and a tendency to focus on “interesting specimens” (usually the most florid cases of pathology) devoid of biological and environmental context has hampered attempts to chart geographic and diachronic variation in prevalence; lack of integration with other kinds of archeological evidence; unfamiliarity with veterinary nomenclature, which is itself under constant revision. Over the past 20 years, however, things have begun to change with: improved understanding of the pathogenesis and archeological significance of skeletal lesions through analysis of known-history populations (e.g., Bartosiewicz et al., 1997; Bendrey, 2007; Darton and Rodet-Belarbi, 2018;

Ortner’s Identification of Pathological Conditions in Human Skeletal Remains. DOI: https://doi.org/10.1016/B978-0-12-809738-0.00023-5 © 2019 Elsevier Inc. All rights reserved.

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810 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains

Fabiˇs and Thomas, 2011; Levine et al., 2000; Niinima¨ki and Salmi, 2016; Rafuse et al., 2013; Taylor and Tuvshinjargal, 2018; Thomas and Grimm, 2011; Zimmermann et al., 2018); technological advances in imaging, biomolecular, and histological approaches (e.g., Bathurst and Barta, 2004; Bendrey et al., 2007; Martiniakova´ et al., 2008; O’Connor and O’Connor, 2005; Rothschild et al., 2001; Tourigny et al., 2016); publication of protocols to encourage systematic recording (e.g., Bartosiewicz et al., 1997; Vann and Thomas, 2006; Thomas and Worley, 2014); the development of theoretical frameworks to aid interpretation (e.g., Binois, 2013; Thomas, 2017); increased rigor in differential diagnosis (e.g., Binois, 2013; Lawler et al., 2016); systematic spatial and temporal analyses of lesion frequencies (e.g., Bartosiewicz, 2002, 2008, 2018; Fothergill, 2016, 2017; Thomas, 2008); and epidemiological modeling (e.g., Fournie´ et al., 2017). While human skeletal analysis and zooarcheology have followed different trajectories, particularly since the emergence of the New Archaeology in the 1960s, this separation is an artifice of practice and reflective of the dominance of anthropocentrism within archeology. It is telling that while humans hold their own disciplinary specialism within paleopathology, all other species are lumped together anonymously as “animal.” This disciplinary schism is less evident in the earliest publications within paleopathology. As far as Moodie (1923b: 11) was concerned, paleopathology encompassed “all evidences of disease and injury prior to the opening of recorded medical history, which begins with the writings of Hippocrates, Aristotle, Aetius, Alcemon, Democritus, Empedocles, and other early writers on biological and medical subjects.” Evidencing this wider consideration, Moodie’s (1923a) volume encompass the description of pathology in plants, crinoids, molluscs, fish, reptiles, birds, mammals, and humans. In this chapter, I will review the relationship between human paleopathology and its nonhuman animal counterpart by identifying areas of commonality and departure, before considering the benefits and challenges of closer integration. Whilst pathology can be observed in a wide range of animal tissues, my focus will center on other mammals, rather than the animal kingdom more broadly, as this is where the closest parallels and relevance reside. Before exploring these synergies and differences, however, a brief review of the kinds of archeological questions upon which nonhuman animal paleopathology can inform is necessary.

RESEARCH FOCI WITHIN NONHUMAN ANIMAL PALEOPATHOLOGY The breadth of archeological questions that can be pursued through the identification and analysis of nonhuman

animal paleopathology has been highlighted in recent publications (Bartosiewicz and Ga´l, 2013). Shifts in emphasis are identifiable through time, however, tracking developments in (zoo)archeological theory more broadly (Thomas, 2012). An important theme has been the potential of animal pathology to contribute to our understanding of the economic exploitation of animals in the past. Considerable attention has been paid to lesions that might provide direct or proxy evidence for the use of animals for traction or riding, contributing to major debates within archeology, such as the timing and nature of the Secondary Products Revolution, the origins of horse riding, and the intensification of farming (e.g., Bartosiewicz et al., 1997; Brown and Anthony, 1998; de Cupere et al., 2000; Daugnora and Thomas, 2005; Groot, 2005; Higham et al., 1981; Issakidou, 2006; Izeta and Corte´s, 2006; Johannsen, 2009; Levine et al., 2000, 2005; Taylor and Tuvshinjargal, 2018; Telldahl, 2005; Thomas, 2008). Much of this research mirrors the approach taken by human paleopathologists using musculoskeletal stress markers to infer activity patterns in individuals and communities in the past (e.g., Hawkey and Merbs, 1995), with all the attendant challenges. Animal paleopathology has also been used to inform upon a wide range of subsistence and animal management strategies in the past, including: stock-keeping (e.g., Fothergill, 2017; Thomas, 2001); captivity (e.g., Nerlich et al., 1993; Zimmermann et al., 2018); herding (e.g., Niinima¨ki and Salmi, 2016); tethering (e.g., Darton and Rodet-Belarbi, 2018; von den Driesch, 1989: 651); breeding strategies and their consequences (e.g., Fothergill et al., 2012; Gordon et al., 2015); nutritional status (e.g., Albarella, 1995; Dobney and Ervynck, 2000; Dobney et al., 2004, 2007; Teegen, 2005); hunting techniques (e.g., Letourneux and Pe´tillon, 2008; Noe-Nygaard, 1974); and feather harvesting (e.g., Fothergill, 2016; Hargrave, 1970). Currently, zooarcheology is being advanced through engagement with ideas emerging out of animal studies and informed by posthumanist discourse (e.g., Allentuck, 2015; Armstrong Oma, 2010; Johannsen, 2009; Lorimer, 2006; Overton and Hamilakis, 2013; Russell, 2012; Sykes, 2012, 2014). These approaches acknowledge the complexity of the relationships between people and animals and stress consideration of the behavior and social lives of animals, as well as the people with whom they interacted. One zooarcheological manifestation of this perspective has been a call to employ a biographical approach to encourage consideration of the complexity of the life history (not just the death history) of animals (Morris, 2011). By way of example, MacKinnon and Belanger (2006) describe a range of pathologies in a small brachycephalic dog from Roman Carthage buried alongside a 10 15-year-old human. This dog exhibited

Nonhuman Animal Paleopathology—Are We so Different? Chapter | 23

congenital hip dysplasia, and pronounced degenerative joint disease was evident in the contralateral pelvis and femur (MacKinnon and Belanger, 2006: 41). Despite the permanently dislocated hip and advanced arthritis, which may have affected the animal’s mobility, the dog survived long enough to lose most of its teeth (potentially a consequence of its diet), suggesting that it had been cared for throughout its life (MacKinnon and Belanger, 2006). Dietary stable isotope analysis indicated a diet richer in protein compared with contemporary dogs (MacKinnon, 2010). In keeping with these concerns, and paralleling the development of the bioarcheology of care within human paleopathology (e.g., Tilley, 2015; Tilley and Schrenk, 2017), zooarcheological paleopathology is being employed to examine human attitudes to animals through the identification of maltreatment (e.g., Binois et al., 2013; Teegen, 2005), therapeutic intervention (e.g., Rozzi and Froment, 2018; Udrescu and Van Neer, 2005), and care giving (Atherton et al., 2012; Bartelle et al., 2010; Bellis, 2018; Thomas, 2017; Tourigny et al., 2016; von den Driesch et al., 2005: 227). While this brief review demonstrates the strength of animal paleopathology for informing upon multiple aspects of past human animal relationships, it should be apparent that the types of questions asked of the archeological evidence are often quite different from those asked within human paleopathology, though this is not the only point of departure.

AREAS OF DEPARTURE As Bartosiewicz and Ga´l (2013: 24) have emphasized, qualitative differences in the expression of bone lesions arise because of species-specific differences in skeletal morphology, such as the presence of horns or antlers, dental specialization, and differentiation of the pentadactyl limb. Size, life expectancy, behavior, and environment are also crucial factors. Fractures,for example, have a better chance to heal and thus be recognized archeologically in smaller species and animals that live to an advanced age (Bartosiewicz and Ga´l, 2013: 24). Interspecies differences in bone tissue composition also exist: for example, Aerssens et al. (1998) report lower bone density and fracture stress values in humans compared with dogs, pigs, cattle, sheep, chickens, and rats. As a consequence of physiological differences, some lesions affecting the skeletons of humans and other primates do not seem to be paralleled in other animals and vice versa. For example, porosities in the outer table of the cranial vault (porotic hyperostosis) and orbital roof (cribra orbitalia) appear to be almost exclusive to primates (e.g., DeGusta, 2010), early hominins (e.g., Domı´nguez-Rodrigo et al., 2012), and anatomically modern humans. Indeed, these are amongst the most frequent

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lesions observed in archeological collections of human remains (Walker et al., 2009). In humans, porotic hyperostosis has been strongly (although not exclusively) linked to anemia: the diploe¨ within the cranium expand, while the tissue of the inner and outer tables thins and becomes more porous. Given that other mammals experience anemia, the absence of these lesions in nonhuman mammals remains unexplained, although one possible case has been observed in dogs (Baker and Lewis, 1975). The reverse situation is exemplified by a diverse presentation of defects in the articular surfaces of other mammal bones, especially within the phalanges of cattle (Thomas and Johannsen, 2011). While “type 2” lesions bear resemblance to osteochondrosis in humans, and there may be a shared pathophysiology, some of the other defect types do not appear to be replicated in humans (Thomas and Johannsen, 2011). It is not only at the level of individual lesions where differences arise. Some diseases observed in human paleopathology are exceptionally rare in other mammalian taxa because of fundamental differences in physiology. Scurvy, e.g., is only observed in hominins, primates, guinea pigs (Cavia spp.), and the fruit-eating bat (Pteropus medius), because of their requirement for a continuous exogenous source of vitamin C (Brickley and Ives, 2008: 41). The similarity of lesions in humans and domestic guinea pigs (Cavia porcellus) was recognized by the early 20th century, and include: loosening of the teeth, especially the incisors and mandibular molars; enlargement of ribs adjacent to the costochondral junction; generalized osteopenia; reduced mechanical strength of long bones; and new bone formation, especially around the joints (Cohen and Mendel, 1918; Kipp et al., 1996). Notably, however, no archeological case of scurvy has been reported in domesticated guinea pigs, despite zooarcheological interest in this species (e.g., LeFebvre and deFrance, 2014; Sandweiss and Wing, 1997). Similar comments could be made in relation to the treponemal diseases (e.g., syphilis, bejel, yaws, and pinta), leprosy, and rheumatoid arthritis. While these diseases have exercised considerable interest amongst human paleopathologists, they infrequently affect nonprimate species. Rare exceptions include leprosy amongst red squirrels (Sciurus vulgaris), treponematosis in lagomorphs (e.g., Meredith et al., 2014; Verin et al., 2012), and a disease resembling rheumatoid arthritis in dogs and cats (e.g., Bennett, 1987; Hanna, 2005). Perhaps unsurprisingly, no zooarcheological cases of these conditions have been reported to date. The inevitable corollary is that there are many diseases that cause skeletal lesions in other mammals that are not paralleled in humans. This chapter is not the venue in which to review all these differences (see Craig et al., 2016 for a thorough review of skeletal lesions

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affecting domestic mammals). Instead, three examples suffice to make the point: G

G

G

Specific pathogens, e.g., canine distemper virus, bovine viral diarrhea virus, feline leukemia virus, which can cause bone lesions (e.g., growth retardation lattices) and, in the case of CDV, dental defects (i.e., enamel hypoplasia), in pups, young calves, and cats, respectively (Craig et al., 2016: 104 105; Dubielzig et al., 1981); Development disorders, e.g., canine craniomandibular osteopathy: the onset of the disease is usually between 4 and 7 months and it causes proliferative bilateral new bone formation, especially in the mandibles, occipital and temporal bones, although occasionally other bones of the skull and long bones are affected; this disease is more common in selected breeds of dog: it is autosomal recessive in terriers, for example (Riser et al., 1967); Diseases of unknown cause, e.g., hypertrophic osteodystrophy/metaphyseal osteopathy in young large/ giant breed dogs, which results in metaphyseal necrosis (Craig et al., 2016: 105, 106).

Some diseases are present in both humans and other mammal populations, but the skeletal expression differs, either in frequency or in the morphology and distribution of lesions. A relevant example of this difference is provided by tuberculosis. Tuberculosis is a chronic, progressive, infectious disease primarily spread through inhaled droplet infection, although it can also be acquired intestinally through contaminated milk. A key feature of the disease is that it is capable of being transmitted within and between wildlife populations, domestic livestock, and humans (i.e., it is zoonotic). Two groups of genetically related mycobacteria are largely responsible for skeletal lesions in animals: Mycobacterium tuberculosis complex and Mycobacterium avium complex. The former can infect humans and a wide range of domestic and wild mammals occupying terrestrial and marine environments (e.g., Cooke et al., 1993; Cousins et al., 2003; CliftonHadley et al., 1993; Luke, 1958; Monies et al., 2000). While the disease has a wide host range, infection most commonly affects cattle, pigs, and carnivores, with Mycobacterium bovis forming the primary pathogen. Among domestic livestock, the spread of tuberculosis is facilitated in intensive husbandry regimes where there is regular close contact between infected humans and other animals. In free-ranging populations of cattle and wild cervids, the prevalence of individuals infected with tuberculosis is approximately 1% 5%, compared with 25% 50% in dairy cattle and farmed deer that are housed or penned in small paddocks (O’Reilly and Daborn, 1995: 4). In contrast to human paleopathology, the number of reported/potential cases of tuberculosis in zooarcheology

remains remarkably small (e.g., Bartosiewicz et al., 2018: 33 35; Bartosiewicz and Ga´l, 2013: 100 102; Bathurst and Barta, 2004; Rothschild et al., 2001; Wooding, 2010: 532). Robust clinical data regarding the appearance and frequency of skeletal lesions in nonhuman animal tuberculosis remain frustratingly poor (Lignereux and Peters, 1999; Mays, 2005; Wooding, 2010), but suggest superficial similarity with lesion formation in humans, affecting both axial and appendicular elements at loci of abundant hematopoietic marrow. However, there do appear to be species-specific predilection sites (e.g., the pelvis in pigs: Cohrs, 1967). Moreover, M. bovis infection in cats and dogs appears to be more proliferative, potentially capable of inducing hypertrophic osteopathy (e.g., Bathurst and Barta, 2004; Snider, 1971; Wooding, 2010: 66), although pulmonary neoplasia requires consideration in any differential diagnosis. In contrast, in ruminants, as in humans, bone resorption and lysis (e.g., osteomyelitis, osteoporosis, cavitation, destruction of articular surfaces) predominates (Bartosiewicz and Ga´l, 2013: 101; Wooding, 2010: 66). Rothschild et al. (2001: 306) have suggested that an undermined subchondral articular surface is “relatively specific for the diagnosis of tuberculosis” in bovids. Robust clinical evidence for this link remains to be demonstrated. The positive extraction, polymerase chain reaction amplification, and identification of M. tuberculosis complex DNA in an extinct bison (Bison cf. antiquus) metacarpal exhibiting this lesion from Natural Trap Cave, Wyoming, dated to c.17,000 BP, is tantalizing, but the possibility of environmental mycobacteria—either ancient or modern—must also be entertained. The zooarcheological infrequency of this disease may, in part, reflect the low frequency of skeletal involvement in animals infected with M. bovis: 0.5% 1% for cattle (Cohrs, 1967); 8% 9.5% for pigs (Nieberle, 1938); and 1.7% for the European badger (Meles meles) (Gallagher and Clifton-Hadley, 2000). However, it might also reflect the fragmentary nature of the zooarcheological record and the inconsistent approach to recording ribs and vertebrae, which are common sites for tuberculous lesions (Nieberle, 1938; Cohrs, 1967). Periosteal new bone formation on the visceral surface of ribs has been suggested as a potential indicator of tuberculosis in human skeletal remains (Roberts, 1999), but is rarely reported zooarcheologically, except when its presence has been sought explicitly (e.g., Thomas and Vann, 2015; Fig. 23.1). In some cases, differences between human and nonhuman animal paleopathology reflect the questions asked of our data. For example, while sharp force trauma is treated as paleopathology in human skeletal analysis (e.g., Appleby et al., 2015), in zooarcheology it falls under the remit of taphonomic analysis through the study of slaughter and butchery marks (Lyman, 1994) (Fig. 23.2). This

Nonhuman Animal Paleopathology—Are We so Different? Chapter | 23

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FIGURE 23.1 Periosteal new bone formation on the visceral surface of a medium-mammal rib fragment from a Roman site in England: a likely indicator of respiratory disease. Failure to examine elements such as these for pathology, represents a missed opportunity in zooarcheology.

FIGURE 23.2 One specialist’s trauma is another specialist’s butchery: blunt and sharp force trauma in an Iron Age cattle skull from England, comprising an unhealed penetrating injury (the cause of death) and multiple cut marks on the frontal (evidence of skinning).

may begin to change, however, as more studies are undertaken in veterinary forensic pathology (e.g., Munro and Munro, 2008). There are also identifiable differences in research foci in human and zooarcheological paleopathology. For example, one can contrast the extensive attention paid to metabolic bone diseases in human paleopathology (e.g., Brickley and Ives, 2008) with the negligible attention paid in zooarcheology, despite the potential interpretative significance of osteodystrophies for informing upon the management of animals in the past. There have been

few published accounts of osteoporosis, rickets/osteomalacia, and fibrous osteodystrophy (e.g., Brothwell, 1995: 232; Leshchinskiy, 2009; Nerlich et al., 1993; Martiniakova´ et al., 2008), and a complete absence of published cases of metal poisoning and other forms of vitamin deficiency or excess. This difference is partly a taphonomic issue, since many of the conditions described above result in demineralization of bone or its replacement by soft tissues, rendering them more susceptible to postdepositional destruction. However, this issue also

814 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains

may reflect the paucity of systematic attempts to determine whether bones that qualitatively feel underweight reflect diagenetic factors or pathology: there is clearly more scope for systematic radiography and histological analysis (e.g., Horwitz and Smith, 1990; Martiniakova´ et al., 2008; Smith and Horwitz, 1984), although the large size of many faunal assemblages and the limited information concerning the normal range of bone densities for different species, breeds, ages, sexes etc., make this practically difficult. Beyond biology, fundamental differences between human and nonhuman animal paleopathology exist that reflect the composition of the archeological record. Faunal assemblages typically comprise disarticulated, isolated fragments of bone that represent waste generated from food preparation (i.e., disarticulation and butchery) and consumption, which may have experienced one or more redeposition events. Only rarely are complete skeletons preserved, and where they do it is most commonly in contexts associated with human internment and/or “ritual” (e.g., Baron, 2018; Lepetz and Van Andringa, 2003; Pluskowski, 2012). Unlike human osteology, therefore, where the nature and distribution of lesions can be assessed with regularity, pathologies in animal bone assemblages are encountered more commonly as isolated examples devoid of their biological context. Differential diagnosis in paleopathology primarily relies upon careful comparison of the morphology and bodily distribution of lesions in archeological specimens with clinical studies in which skeletal changes associated with specific diseases are documented. Conducting a differential diagnosis is a process of exclusion: ruling out all diseases that could potentially lead to the formation of similar lesions before a conclusion is drawn. The emphasis on bodily distribution is important, as bone can only form or destroy, thus limiting the variability of skeletal responses to pathology and potentially resulting in equifinality at the level of the individual lesion. Consequently, in animal bone assemblages it is often impossible to achieve greater precision than a broad nosological category, which, in turn, limits the interpretative potential. This problem is exacerbated by zooarcheological recording practices. In contrast to human osteology (e.g., Buikstra and Ubelaker, 1994), there are no minimum standards by which animal bone assemblages should be recorded. While general recommendations have been made (e.g., Baker and Worley, 2014) and a degree of conformity exists, making possible the creation of synthetic datasets (e.g., Arbuckle et al., 2014; Atici et al., 2013), zooarcheologists exercise considerable discretion in what to record and how. To a certain extent this is advantageous, permitting data to be collected to address focused research questions and avoiding a “paint by numbers” approach to recording. However, problems of comparability arise, particularly where a selective approach to recording is advocated due to

time or financial constraints. Davis (1992), for example, developed a rapid recording method for animal bones from archeological sites that focuses attention on a limited suite of body parts, discounting ribs and vertebrae. While such elements are often highly fragmentary and can be difficult to identify to taxon, these bones are susceptible to lesions arising from systemic, bloodborne diseases (such as tuberculosis) because they contain a high proportion of cancellous bone. Failure to record these bones, therefore, immediately minimizes opportunities to differentially diagnose a suite of conditions (Fig. 23.1). Standardized, descriptive-led recording is essential given the low number of pathological animal bones that are recorded on archeological sites, typically less than 1% of the total number of identified specimens (NISPs; Siegel, 1976; Shaffer and Baker, 1997; Vann and Thomas, 2008). In part, this paucity reflects the high degree of reduction of animal bone assemblages, resulting from butchery, depositional, and taphonomic processes. However, this low number also reflects differences in life expectancy. Diseased wild animals are more likely to be predated or die prematurely, reducing the occurrence of chronic conditions that affect the skeleton (Bartosiewicz, 2016). Domestic livestock were less likely to die from natural causes: instead, they would have been slaughtered for meat (irrespective of their exploitation for secondary products during life) or sometimes for “ritual” purposes: given the correlation between susceptibility to pathology and age, it is no wonder that frequencies of pathology are much higher in humans than they are in livestock. Livestock exhibiting signs of illness may have also been earmarked for early slaughter and dumped/buried in locations away from the occupation sites that typically form the focus of archeological enquiry. This certainly adds complexity to the application of the “osteological paradox” (Wood et al. 1992) to animal bone assemblages because an absence of pathology in animals could reflect: G G

G

A healthy animal that was slaughtered; A diseased animal that was slaughtered before skeletal tissues became involved; A diseased animal that died before skeletal tissues became involved.

Even where lesions are identified and recorded, challenges exist for the zooarcheologist in quantifying pathology. As Bartosiewicz (2018: 187) has emphasized, in clinical epidemiology prevalence is calculated as the number of cases of a disease in the living population divided by the total number of individuals in that population. In human osteology, calculating prevalence is hampered by the fact that cemetery populations are not representative of the living population. The challenges are even greater in zooarcheology due to imprecise dating of assemblages (usually by association with material culture), unknown

Nonhuman Animal Paleopathology—Are We so Different? Chapter | 23

accumulation rates, uncertain depositional histories, and the questionable validity of minimum number of individuals determinations (Lyman, 2008). Conceivably, an urban animal bone assemblage could include many individuals from multiple herds raised in different environments over a period of tens or hundreds of years. Zooarcheologists attempt to overcome this issue by relying upon the proportion of pathological bones out of the total NISP as a crude proxy of prevalence (Bartosiewicz, 2018: 187). Such an approach facilitates diachronic and regional comparisons of changing frequencies and types of pathology. However, as with other zooarcheological phenomena (e.g., taxonomic diversity), a logarithmic relationship exists between the total NISP and the character being recorded (Lyman, 2008). Thus, relatively more pathological bone is likely to be encountered in larger assemblages (up to a NISP threshold that is geographically and diachronically dependent) (e.g., Bartosiewicz, 2018: 200). Consequently, equating the zooarcheological prevalence of pathology with clinical prevalence is fraught with difficulty.

AREAS OF COMMONALITY Notwithstanding the physiological, environmental, and behavioral differences between humans and other animals, a close connection exists with respect to disease histories that demands attention by both paleopathological practitioners. Following an examination of 25 unicellular microbial pathogens with the highest burdens on human health, Wolfe et al. (2007) established that more than half of the temperate diseases considered originated from domestic animals, while over half of tropical diseases came from wild (nonprimate) animal populations. Although the focus of such research has centered on pathogens transmitted from animals to humans (zoonoses), transmission occurs just as often in the other direction (reverse zoonoses) (Messenger et al., 2014), with intermediary hosts common in both pathways. Rather than being brought in from the wild, many major pathogens emerged in the Old World in the environmental conditions of domestication and urbanization, in which bacteria, viruses, protozoa, ticks, and parasites thrive, mutate, and jump hosts (Brothwell, 1991; Wolfe et al., 2007). The interconnectivity between human and animal health, pathogen behavior, and environment is exemplified by events in 14th-century Europe. Three consecutive failed crop harvests in the period 1315 17 occurred as a consequence of torrential rainfall, and constituted a major contributing factor to a human famine resulting in the death of 10% 15% of the European population (Jordan, 1996). Almost simultaneously, there were major outbreaks of

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disease affecting animals in the first quarter of the 14th century. Sheep murrain (an unspecified infectious disease) was epidemic between 1314 and 1316, while a panzootic in cattle (probably rinderpest) was widespread between 1319 and 1322 (Jordan, 1996; Newfield, 2009). The scale of mortality was astonishing. In England and Wales, manorial accounts indicate that around 62% of cattle died of pestilence between 1319 and 1320 (Slavin, 2012), severely affecting the ability of farmers to undertake tasks requiring animal power (e.g., harvesting and transporting crops). Notably, a systematic and quantitative analysis of degenerative changes to the lower limb bones of cattle points to intensified use of the surviving cattle for traction (Thomas, 2008). It seems likely that the high mortality was, at least in part, a consequence of nutritional vulnerability amongst the cattle, as poor crop yields and spoiled hay affected the ability of farmers to adequately feed their animals (Jordan, 1996). Less than a generation later, the Black Death arrived, decimating the human population by between one third and a half (Benedictow, 2004). Clearly, the virulence of the strain of Yersinia pestis was instrumental, but preexisting nutritional stress caused by these earlier disasters may have been a predisposing factor. It is not just disease histories upon which there is shared intellectual ground between human and nonhuman animal paleopathologists. As I have argued elsewhere, “relationships between people and animals are closely entangled and bound up with the behaviors of both, human identities and conceptions of animal consciousness and/or moral considerations” (Thomas, 2017: 180). The links between animal abuse, child abuse, and domestic violence in contemporary society are well founded (e.g., Arluke et al., 1999; Lockwood and Ascione, 1997), and considerable potential exists for exploring these interrelationships in the past through the integration of human and nonhuman animal paleopathology. Beyond collective interests in the study of health, disease, and injury in the past, there are also many points of commonality in the foundational methods of human and nonhuman animal paleopathology. At the most basic level, the underlying cellular processes by which bone tissue responds to disease and injury are comparable across mammals. Consequently, the fundamental principles of describing lesions is identical. Indeed, the experience of human paleopathologists was explicitly acknowledged and drawn upon in the development of the first systematic methodology to record pathology in animal bone assemblages (Vann and Thomas, 2006; Vann, 2008). As Thomas and Worley (2014) note, systematic recording and reporting is essential because it: 1. Draws attention to pathologies that are absent, as well as those that are present;

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2. Highlights the full range of lesion manifestations, not just the “spectacular” cases; 3. Requires the calculation of lesion frequency, which in turn facilitates intra- and intersite comparisons and the identification of spatial and temporal patterns. As in human paleopathology, in the absence of quantitative recording schema (e.g., Bartosiewicz et al., 1997) individual lesions must be described in detail to capture shape, size, precise anatomical location, and morphology, especially with reference to the processes of bone formation and/or destruction and remodeling. Wherever possible, precise descriptive terminology should be employed (reflecting current clinical practice in veterinary medicine) and written descriptions should be supported by annotated illustrations (photographs, radiographs, and line drawings). This approach is vital for providing a robust foundation for differential diagnosis and overcoming the shared problem of separating pathology from taphonomic processes that result in pseudopathology. While the differential diagnosis of lesions in nonhuman animal paleopathology is challenging, for the reasons enumerated above, the intellectual process through which this is achieved is the same: requiring the exclusion of all potential causes of the lesions before a conclusion is drawn. The terminology of diagnoses should follow the precedent established in clinical literature, to recognize species-specific disorders and the fact that a terminological consensus does not exist between the two branches of medicine. Once a diagnosis is achieved, there is a shared requirement to indicate the strength of confidence in that determination. Appleby et al. (2015) have recommended the use of an adapted version of the Istanbul Protocol, a system of nomenclature ratified by the UN and widely used in forensic medicine for the identification of torture, for both human and nonhuman animal paleopathology. This system permits the degree of certainty by which a diagnosis is made to be qualified using one of five categories: 1. Not consistent: the lesion could not have been caused by the condition(s) described; 2. Consistent with: the lesion could have been caused by the condition(s) described, but it is nonspecific and there are many other possible causes; 3. Highly consistent: the lesion could have been caused by the condition(s) described, and there are few other possible causes; 4. Typical of: that the lesion is usually found with this type of condition(s), but there are other possible causes; 5. Diagnostic of: the lesion could not have been caused in any way other than by the condition(s) described (i.e., it is pathognomonic).

Some of the general principles of interpretation are equally relevant between human and nonhuman animal paleopathology (after Thomas and Worley, 2014): lesion frequencies within each species will vary as a result of age, sex, body mass, activity patterns/behavior, and inherited predisposition; lesions will, in general, increase in frequency with age; lesions occurring early in life may be obscured completely through remodeling; connecting bone lesions with symptoms (and pain) is difficult given the different ways in which these are experienced by individuals; pathology can differentially affect the preservation of bones negatively (e.g., osteoporotic bone) and positively (e.g., bones with sclerotic lesions). A strength of paleopathological investigations in both humans and other animals, is its ability to contribute meaningful archeological information at different scales of analysis. The following two examples have been selected for presentation here because they showcase how zooarcheological pathology can be used to inform upon millennia-spanning environmental change and individual human animal interactions. Systematic analysis of dental pathology in dire wolves (Canis dirus) from the Rancho La Brea tar pits, Los Angeles, revealed a decrease in fracture frequency from 7% to 2% between 15,000 and 12,000 BCE (Binder et al., 2002). This change was attributed to reduced competition caused by high prey abundance and/or low carnivore density in the early Holocene, lessening the risk factors for dental fractures (e.g., lower inter- and intraspecific aggression and reduced intensity of carcass utilization). Analysis of pathology in a 19thcentury dog burial from Toronto, Canada, revealed: systemic degenerative joint diseases; severe periodontal disease; a chipped tooth and associated abscess; a chronic ear infection (Fig. 23.3); and chronic infection of the left limb, probably secondary to a penetrating injury. When placed within a biological, archeological, and historical context, Tourigny et al. (2016) constructed a biography of the individual, reflecting upon its life history and its owners’ attitudes towards dogs.

TOWARD CLOSER INTEGRATION Human and nonhuman animal paleopathology share common goals, namely to: G

G

G

Identify, record, and differentially diagnose skeletal lesions in the past; Interpret the significance of disease and injury within past populations, at different scales (individual, local, regional) within itsbiological, cultural, and environmental context; Understand the evolutionary history of diseases.

Nonhuman Animal Paleopathology—Are We so Different? Chapter | 23

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FIGURE 23.3 Basocranial view of a dog skull from 19th-century Canada, exhibiting reactive bone formation and destruction of the left tympanic bulla and external acoustic meatus consistent with chronic osteitis (see Tourigny et al., 2016). This advanced ear infection likely caused deafness and the dog may have walked around with his head tilted to the left and exhibited behaviors such as head shaking and scratching. Such details enable the biography of the animal to be reconstructed.

While the two disciplines share common goals, and many of the underpinning approaches to recording (e.g., lesion description, analytical methods, and the process of differential diagnosis) are compatible, differences do exist in anatomy, physiology, immune response, behavior, environment, life expectancy, and disease incidence and expression, which must be acknowledged, especially in the application of clinical evidence across taxonomic boundaries. As I demonstrated in the literature review at the beginning of this chapter, zooarcheologists are beginning to make great strides in addressing the first two goals. A particularly encouraging development has been the analysis of pathology in known-history collections to address some of the gaps in clinical knowledge (e.g., Bartosiewicz et al., 1997; Bendrey, 2007; Darton and Rodet-Belarbi, 2018; Fabiˇs and Thomas, 2011; Levine et al., 2000; Niinima¨ki and Salmi, 2016; Rafuse et al., 2013; Taylor and Tuvshinjargal, 2018; Thomas and Grimm, 2011; Zimmermann et al., 2018). The fragmentary nature of the archeological record makes the third goal the most challenging, and pessimism has been expressed in our ability to make a meaningful contribution (Bartosiewicz and Ga´l, 2013). Nevertheless, understanding the evolutionary history of disease is one arena where there is enormous potential for future collaboration. The value of comparative pathology was recognized in the late 19th century (e.g., Bland Sutton, 1890) and in the very first publications concerning paleopathology (e.g., Moodie, 1923a,b), but disciplinary specialization and an anthropocentric worldview have undermined this approach. The systematic analysis of skeletal lesions in humans and

other animals is essential for this work, but it needs to be supplemented with the direct measurement of diagnostic parameters (e.g., Horwitz and Smith, 1990) and, for infectious diseases, the direct detection of causative pathogens (e.g., parasites: Søe et al., 2018; and bacteria: Bendrey et al., 2007) (Mays, 2018). Ancient DNA analysis, in particular, has enormous potential to unpick the genetics of disease resistance in humans and other host animals. Crucially, paleopathological evidence needs to be grounded within an ecological/environmental (as well as biological) context, especially for infectious disease, to generate an understanding of the interplay between pathogen, host(s), and environment (e.g., Engering et al., 2013). The epidemiological modeling approach advocated by Fournie´ et al. (2017) is particularly exciting in this respect. This approach parallels the recommendations of contemporary movements that have taken an integrated systems approach to managing health risks (e.g., “One Health,” “Eco Health,” “Planetary Health”), which “can only be tackled by interdisciplinary working across the domains of human medicine, veterinary medicine and the life sciences” (Woods et al., 2018: 14). Such research demands a collaborative approach, not least because disease definitions, which are based upon clinical signs, pathology, and response to treatment, are constantly under review, especially with the widespread application of genetic/molecular evidence. Bringing an integrated paleopathological approach to these debates has the potential to make a major contribution to our understanding of the health consequences of domestication and the progressive intensification of agricultural systems—issues that have profound relevance in contemporary society.

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ACKNOWLEDGMENTS I am indebted to Jane Buikstra for inviting me to contribute to this volume and to Elizabeth Uhl for providing insightful comments on an earlier draft. Thanks to Dennis Lawler for drawing my attention to the potential case of porotic hyperostosis in dog. I am also hugely grateful to my colleagues Jo Appleby and Mara Tesorieri, and all the students who have taken the “Human and Animal Health and Disease” module, which has provided intellectual inspiration for many aspects of this chapter. The first quotation at the beginning of this chapter was reproduced in Woods et al. (2018, 1); I was alerted to the second quotation by Elizabeth Uhl. Figs. 23.1 and 23.2 were taken by Sian Holmes. Fig. 23.3 is reproduced with the kind permission of Eric Tourigny.

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