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ARTICLE /ARTICLE

Paleoepidemiology of pre-Columbian and Colonial Panamá Viejo:

a preliminary study

Paléoépidémiologie à Panamá Viejo, l’état sanitaire des habitants précolombiens et coloniaux : une étude préliminaire

C.M. Rojas-Sepúlveda · J. Rivera-Sandoval · J.G. Martín-Rincón

© Société danthropologie de Paris et Springer-Verlag France 2011

Abstract Panamá Viejo, a coastal site in the Panamanian Pacific, was occupied from ca. AD 850 to 1200 and from the arrival of the Europeans in AD 1519 to 1671. This paper describes the bone anomalies observed and recorded in the pre- and post-contact skeletal series from the site.

The bone anomaly frequencies were calculated, taking their observability into account. A number of changes were found between the two periods, concerning MSM (Musculoskeletal Stress Markers), DJD (Degenerative Joint Disease), porotic hyperostosis and trauma, as well as changes that are classified as ‘cultural’. However, these differences were significant only in the case of porotic

hyperostosis and the‘cultural’anomalies. Further investiga- tions using an enlarged series should clarify the impact of European contact in this region.To cite this journal: Bull.

Mém. Soc. Anthropol. Paris 23 (2011).

Keywords Bioarchaeology · Gran Darién · Paleopathology · Pre-contact · Post-contact

Résumé Panamá Viejo, un site côtier sur le Pacific pana- mien, a été occupé entre le 850 et le 1200 après J.-C. et à partir de l’arrivée des européens en 1519 jusqu’au 1671 après J.-C. Nous avons observé et recensé les anomalies osseuses des séries ostéologiques issues du site de la période antérieure au contact européen ainsi que de la période postérieure. Leurs fréquences ont été calculées selon l’observabilité. Plusieurs changements osseux entre les deux périodes ont été repérés en ce qui concerne les MOA (marqueurs osseux d’activité), la MAD (maladie articulaire dégénérative), l’hyperostose poreuse, le trauma et les change- ments classés comme « culturels ». Cependant, des différences significatives ont été trouvées seulement pour les hyperostoses poreuses et pour les anomalies dites « culturelles ». D’autres études et notamment l’élargissement de la série devraient éclaircir l’impact du contact européen dans cette région.

Pour citer cette revue : Bull. Mém. Soc. Anthropol. Paris 23 (2011).

Mots clésBioarchéologie · Gran Darién · Paléopathologie · Précontact · Postcontact

Introduction

The site known today as Panamá Viejo is located on the Pacific coast of the Isthmus (Fig. 1) in the archeological area known as ‘Gran Darién’ [1], a zone with abundant natural resources and alternating dry and wet periods (1300 to 1500 mm of precipitation in nine months of

C.M. Rojas-Sepúlveda (*)

Unité d’anthropologie : adaptabilité biologique et culturelle, UMR 6578, faculté de médecine, secteur Nord,

université de la Méditerranée, CS80011, 51, boulevard Pierre-Dramard,

F-13344 Marseille cedex 15, France e-mail : clarosepul@gmail.com

Programa de Antropología, Facultad de Humanidades, Universidad del Magdalena,

Carrera 32 No. 22-08 Sector San Pedro Alejandrino, Santa Marta, Colombia

J. Rivera-Sandoval

Universidad Nacional del Centro de la Provincia de Buenos Aires, Av. Del Valle 5737, Olavarria, Provincia de Buenos Aires, Argentina

Fundación Erigaie, Calle 10 No. 3-76, Bogotá D.C. Colombia

J.G. Martín-Rincón

Departamento de Historia y Ciencias Humanas, Universidad del Norte,

Km 5 vía a Puerto Colombia, Barranquilla, Colombia Secretaría Nacional de Ciencia,

Tecnología e Innovación-SENACYT, Apdo, 0835-00590, Ciudad de Panamá, Panamá DOI 10.1007/s13219-011-0033-3

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the year). Tidal variations that also affect the nearby river frequently cause flooding in the Panamá Viejo site [2].

Panamá Viejo is considered to be the first Hispanic settle- ment in the South American Pacific and was used as a naval base for the‘conquistadors’and for warehousing the treas- ures looted from the Incas before they were shipped to Spain.

However, archeological excavations at the site since 1995 have shown that it was occupied by humans for over a thou- sand years before the first contact with Europeans [1,3–5].

Pre-Columbian coastal occupation of the Panamá Viejo arche- ological site may have lasted from AD 500 until the arrival of the Spaniards in AD 1519 [1,5,6].

Large concentrations of sea shells, many ceramic artifacts and stone objects such as‘metates’,‘manos’(for grinding cereals) and knives have been found in pre-contact burial sites. However, shell, bone or metal objects are not frequent and they differ in style from the ceramics, indicating a foreign origin and, probably, the privileged treatment of some individuals [2].

Three types of burials have been identified in Panamá Viejo. Primary burials are where a complete fresh body was deposited and then decomposed without being disturbed until archaeological excavation; secondary burials are where a previously defleshed body was buried; reduced burials, finally, are defined as primary burials that were subsequently disturbed as the site was continually re-used for further burials [7].

Secondary burials in urns or bundles are the most frequent in the pre-Columbian contexts at Panamá Viejo (66%). They frequently include several individuals (44%), although single burials are the most common (56%) [4,5]. Before contact with Europeans occurred, the burial sites were usually adja- cent to domestic units [2], and individuals were buried with stone artifacts, ceramics, food and sometimes human skulls [4,5].

Reduced burial was frequent in Colonial times (84%), when individuals were mostly buried in churches (75%) and chapels, along with crucifixes and medals [4].

This paper presents the findings from a paleoepidemiolo- gical study of the Panamá Viejo osteological series, which is helping to reconstruct the lifestyles of early inhabitants of this coastal site.

Methods and materials

The study used bioarcheological methods [8–13], in which particular attention is given to relationships between human beings and the environment [9,10,14–16].

Sex determination and age estimations were made in accordance with conventional anthropological methods [12,17–21].

The bone‘abnormalities’or ‘anomalies’were observed macroscopically and recorded [8,9,11,14,22–26]. Abnormal Fig. 1 Location of the Panamá Viejo site /Localisation du site Panamá Viejo

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bone formation (periostosis and subperiosteal formation), abnormal bone destruction (osteolysis and osteoporosis), abnormal bone density, abnormal bone size and abnormal bone shape were observed as expressions of disease [11].

Care was taken to differentiate antemortem from postmortem bone anomalies [11]. The location of each abnormality was carefully described in each individual record.

It is very important to investigate multiple indicators when reconstructing health profiles from skeletal remains [14]: this paper presents six different indicators of health and disease. Four of them are specific indicators of stress:

nutritional deficiencies (porotic hyperostosis), some infec- tious diseases, trauma and degenerative pathologies [15].

The other two are non-pathological, but are potentially very important because they may reveal lifestyle features:

musculoskeletal stress markers inform us about physical activity; while so-called ‘cultural’ abnormalities (defined below) inform about behavior.

Porotic hyperostosis

Porotic hyperostosis is diagnosed by the observation of very porous lesions on the cranium, the roof of the eye orbits and the ends of long bones [11,14,15,24]. These lesions are produced by marrow proliferation caused by expansion of the diploe, which makes the outer layer of bone thinner, exposing the porous trabecular bone [14,15].

Porotic hyperostosis was recorded in this study because it has been shown to be a valuable tool for reconstructing the health profiles of past populations [11,14,15,24].

This indicator has been associated with anemia [14,15];

nutritional anemia is the main etiological factor in documen- ted prehistoric cases [14]. However, other causes such as infections, neoplastic or metabolic diseases can produce similar lesions [11]. Iron-deficiency anemia seems particu- larly prevalent [11,14,15,24,27]. This type of anemia may be caused by repetitive bleeding, parasitic infection, dietary deficiencies, diarrheal disease, iron absorption deficiencies and chronic inflammatory or infectious diseases [14,15,27,28]. The frequency of these lesions increased with sedentism, agriculture and increased population density [14,15,27,28]. They first appear in childhood but are present until adulthood [27,29].

Care must be taken when studying this indicator [30], because the presence of genetic hemolytic anemias, parasite exposure, hemoglobinopathies [31] and other associated pathological conditions such as inflammation, osteoporosis and the pseudopathologies [32] must be assessed. Wapler et al. have shown that only 44% of the cribra orbitalia cases observed in their study had histological features indicating changes due to anemia [32]. The real meaning of the presence of porotic hyperostosis is therefore yet to be clarified [33].

Although marine resources are rich in proteins and iron, a high frequency of anemia was found in one population, whose diet was based on seafood [27,29]. This can be explained by the phosphorus content of seafood, which can reduce iron absorption, as well as by other factors such as parasites and diseases like tuberculosis [33].

Although hereditary anemias (sickle-cell anemia and thalassemia) may produce porotic hyperostosis, the lesions seem much more pronounced and asymmetrical than those produced by nutritional anemia [14].

Anemia has highly negative effects on disease resistance, capacities for activity and cognition in individuals and populations. It may not be lethal, but in combination with other factors, it can contribute to poor health [14,15]

and predispose children to infection, which can increase mortality [33].

However, recent research has brought a new and interest- ing perspective: Walker et al. have presented evidence sug- gesting that a vitamin-B12-deficient diet is more likely to be the key nutritional component in porotic hyperostosis and cribra orbitalia [30]. They argue that well-documented clini- cal studies have proved that maternal diets lacking foods of animal origin, and therefore vitamin-B12-deficient, have effects on nursing infants such as the development of mega- loblastic anemia [30].

Infectious diseases

The main skeletal response to many kinds of infection or inflammatory conditions is an elevation of the fibrous outer periosteal layer of the bone, or periosteal reaction [11,14,15].

Most prehistoric cases are nonspecific because lesions can be caused by a number of pathological conditions but the exact etiology is unknown [14,15]. As bone is constantly being remodeled, resorption of old bone and accretion of new bone are also common reactions [14]. Destruction and necrosis may also occur in response to infection [11,14,15].

Periosteal reactions may also be a response to trauma, and to endocrine-metabolic or circulatory dysfunction [11,12,24,29]. These reactions produced by infectious dis- ease may be distinguished from those produced by trauma, because the latter are small, localized and non-destructive, while the former are generalized, destructive and bilateral, and affect multiple long bones [14,24]. According to Good- mann and Martin, when a tibia presents periosteal reactions, other long bones should be carefully assessed [14]. If other bones are involved, the tibia is considered to display a reaction to systemic infectious disease. If only the tibia is affected, this should be classified as a localized response to trauma [14].

Specific infections such as tuberculosis and treponemal infections can be diagnosed through careful observation.

Tuberculosis produces a bone-destroying response in the

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vertebral column, the joints and the cranium. Treponemal infections, such as syphilis, produce saber shins, polydacty- litis and osteolytic lesions of the external nasal vault and the nasopalatal region [11,14]. However, because of the small size of the series studied, all infections, both specific and nonspecific, were taken together in this case.

Infectious disease combined with malnutrition is the main contributor to morbidity and mortality in human popula- tions, particularly during childhood [11,14,24,29]. The fre- quency of infectious lesions increases with sedentism and higher population density [14,24,29,34].

The study of infectious disease in skeletal remains raises a number of problems [35,36]. One of the most important is that only some infectious episodes leave diagnostic markers on the skeleton [14], because healing or death can occur before the bone reaction, potentially leading to underestima- tions of infectious disease frequencies in the past [11,14,24,29]. Chronic (usually non-lethal) conditions can therefore reveal aspects of lifestyle, such as nutritional adequacy, diet, waste disposal and hygiene, while virulent and epidemic infections do not [14].

On the other hand, the‘osteological paradox’proposed by Wood et al. argues that it is impossible to determine whether an absence of infectious bony reactions is accounted for people who were very resistant to infectious diseases, or by people who were very frail and died before any bony response was possible [36]. However it is worth studying infectious lesions from human skeletal remains in combina- tion with other indicators because they tell us about the living conditions of past population [37].

Trauma

Traumatic lesions are caused by physical force. Trauma is defined as any bodily injury or wound [29]. They are pro- duced by external impacts, either accidental or intentional, on the skeleton. Trauma frequency and profiles in the past are associated with culture and lifestyles [11,14,15,24,29].

Ortner has described four types of trauma: partial to com- plete breakage of a bone (fracture), abnormal displacement or dislocation of joints, disruption in nerve and/or blood sup- ply and artificially induced abnormal bone shape1or contour [11]. Deformation, scalping, mutilation, amputation, trepan- ning [11,24], crushing injuries, wounds caused by weapons or other devices and biomechanically induced pathologies such as exostoses, osteochondritis dissecans and spondy- lolysis are also classified as traumatic lesions [14].

Bone will always respond to any kind of fracture by form- ing new bone through the release of calcium salts to form the callous matrix [14,15].

Trauma can have functional repercussions, for example by disabling and deforming individuals and thus reducing their autonomy [14], or by causing complications such as misalignment, traumatic osteoarthritis, osteoporosis, non-union, joint fusion, traumatic myositis ossificans or infection, which may eventually cause death [11,24,29,38].

Trauma also has behavioral implications. For example, specific activities can predispose individuals to recognizable types of accidental or intentional trauma; for this reason, this indicator can be very useful for interpreting past lifestyles [14,15,29].

Some specific kinds of trauma must be carefully observed as they can give important insights into lifestyles. For exam- ple, ‘clay-shoveller’s fractures’, which are fractures of the transverse processes in the seven cervical and first thoracic vertebrae, are produced by combined trapezius and rhom- boid muscle action on bone, and may help to identify the occupations of people in the past [29]. Amputation, trepan- ning, intentional deformation and scalping can also reveal a great deal about a particular society [14,15,24,29].

In this study, particular attention was given to distinguish- ing fractures from post-mortem bone breakages [11,24,29].

Degenerative Joint Disease (DJD, Osteoarthritis)

The breakdown of the cartilage and lubricating systems of the articular surfaces of joints is a common and easily discerned condition [14]. When the subchondral bone is exposed, pit- ting, lipping, erosion and possibly eburnation may occur [14,15]. Thus, deformation of the joint is caused by two kinds of DJD processes, one of bone formation (osteophytes) and one of bone destruction (sclerosis, eburnation, porosity, cysts) [24,39–41]. Consequently, joint and bone degeneration are clearly recognizable in skeletal remains [29].

Although osteoarthritis is multifactorial and related to nutrition, genetics and viral infections [11,12,15,24,26,29, 40,42–48], it has been argued that it is primarily caused by biomechanical wear and tear and functional stress associated with activity patterns that habitually put strain on joint tissues [14,15]. For this reason, the study of osteoarthritis provides clues to the life of individuals in the past, because DJD is connected to the way humans relate to their environment [14], reflecting cumulative and repetitive motions (strain caused by repetitive use of the musculoskeletal system). It may therefore be considered as an indicator of lifestyles and work habits in prehistoric populations [14,15,29].

In this study, attention was given to discard septic and secondary (post-trauma) DJD [29,48]. Osteophytes, mar- ginal lipping and eburnation were recorded as diagnostic signs of DJD. However, pitting on its own was not included as an indicator of DJD because of its diagnostic weakness [41,49,50].

1In this study skull deformation was classified apart, as a‘cultural’

anomaly.

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Musculoskeletal Stress Markers (MSM)

A MSM or enthesopathy is an osseous change in the site where a muscle, ligament or tendon is inserted; conse- quently, it has been related to muscular hyperactivity [43,45,51–58]. However, the relationship between MSM and activity has to be studied with some care to avoid over- simplification [52,58].

Because MSM can occur in association with seronegative spondyloarthropathies, traumatic injuries and diffuse idiopathic skeletal hyperostosis-DISH [58], individuals with any of these conditions were discarded from the statistics.

The presence or absence of large MSM was recorded for all limb bones. In accordance with the methodology devel- oped by Hawkey and Merbs [53], individuals with MSM above 3 in the scale were included in the study. Although this methodology has been criticized [57,58], it makes it possible to see where an osseous change reveals muscular

overuse. This methodology has been widely applied [59–66], with very low inter-observer error [53].

‘Cultural’anomalies

The following were grouped together as ‘cultural’anoma- lies: deformation of the skull, cut marks showing possible defleshing and a very particular bone anomaly in the skull.

The latter is a very circumscribed area of extreme porosity in the scalp, delineated by marked temporal lines in the parie- tals. The upper temporal line provides an attachment to the temporal fascia and the lower temporal line indicates the upper limit of the muscular origin of thetemporalis, one of the muscles involved in mastication (Fig. 2).

The presence, absence or non-observability of each anomaly in the paleopathological data was recorded. Simple statistical analyses were performed, mainly based on frequency calculations. Prevalence values were calculated

Fig. 2 Culturalanomaly: extreme porosity in the scalp circumscribed by very marked temporal lines. Top: Parque Morelos, Burial C4, Individual 1. Bottom: Plaza Mayor, Burial 1, Individual 2 /Anomalie « culturalle » : microporosité intense sur la voute limitée par des lignes temporales très marquées. En haut : Parque Morelos, Tombe C4, Individu 1. En bas : Plaza Mayor, Tombe 1, Individu 2

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by taking only observable aspects into account [26,67]. The proportion n/N was used,‘n’ being the number of indivi- duals presenting the anomaly or indicator considered, and

‘N’the number of individuals with the relevant anatomical parts preserved [26,67].

Comparisons between groups were made by period, sex and age, and the significance of the differences between fre- quencies was calculated by applying the chi-square test.

When the sample was too small to apply the chi-square test, the Fisher exact test was used instead (P < 0.05).

The Panamá Viejo series comprises at least 65 individuals from the pre-Columbian period and 240 from Colonial times [4]. Unfortunately, the bone material was not in an optimum state of preservation, because of the acidity of the soil and repeated re-use of the burial sites [4]. For these reasons, of the total number of individuals analyzed, we were able to assess 52 from the pre-Columbian contexts for anomalies, and 64 for the Colonial period. The post-contact contexts, as mentioned above, produced reduced burials, making indi- vidualization difficult in many cases. Table 1 summarizes the distribution of the series by period and sex, while Table 2 shows the distribution by period and age.

Results

The proportion of individuals less than 15 years old in Pan- amá Viejo is high in the Pre-Columbian context compared to the Colonial period (Table 2), while the proportion of old individuals is low in both periods.

At least one bone anomaly was observed in 27 individuals of the 47 observable pre-Columbian skeletons (57%); in the Colonial contexts, at least one bone change was apparent in 32 individuals out of 64 (50%). Table 3 shows the frequen- cies and percentages of anomalies observed by period of time and by sex. At the top of the table are the frequencies for all the individuals observed; the bottom of the table shows frequencies for the adults only.

In both the Pre-Columbian and Colonial contexts, Musculoskeletal Stress Markers (MSM), Degenerative Joint Disease (DJD), trauma and infectious reactions are the most frequently observed bone changes (Table 3). In Pre-Columbian males, there are noticeably high frequencies of so-called ‘cultural’ bone changes (see methodology for more details).

MSM and DJD were more frequently recorded in the Colonial than the pre-contact sample. Trauma and porotic hyperostosis were more frequent in the pre-Columbian sam- ple. The frequencies of infectious reactions recorded were similar in both periods (around 15%).

These differences were tested with the Chi-square proce- dure, or the Fisher exact test when needed (Table 4). The first section of Table 4 compares the total sample for the two periods, showing a significant difference only in the compar- ative frequencies of‘cultural’anomalies. For the adults (bot- tom), significant differences appear between the frequencies of porotic hyperostosis and‘cultural’anomalies. The second section compares females and males within each period and shows no significant difference. The third section compares females from the two periods and males from the two peri- ods. Significant differences appear only in the frequencies of porotic hyperostosis and‘cultural’anomalies in males, when all are taken together (top), and also when only adult males are taken (bottom).

Discussion

Although a number of interesting features were observed in this study, poor preservation was a major issue (as is usually the case in bioarchaeological studies). The material analyzed is poorly preserved because of the acidity of the soil in this region of the Isthmus and disturbance due to repeated use of the same burial sites. These factors made observation of the bones difficult, and reduced the sample size. However, care- fully recording the presence, absence and non-observability of each anomaly and using the frequency calculation Table 1 Sex composition of the samples / Distribution des

échantillons par sexe

Pre-Columbian Colonial

n % n %

Female 4 7.7 12 18.7

Probable female 4 7.7 4 6.2

Male 23 44.2 11 17.2

Probable male 4 7.7 9 14.1

Indeterminate 17 32.7 28 43.7

Total 52 64

n: number of individuals.

Table 2 Age composition of the samples /Distribution des échantillons par âge

Pre-Columbian Colonial

n % n %

Total Younger than 15 years 18 34.6 10 15.6

Young (15-30 years) 17 32.7 30 46.9

Middle age (30-45 years) 14 26.9 8 12.5

Old (45+ years) 0 0 1 1.6

Adult (unclassifiable) 3 5.8 15 23.4

Total adults 34 65.4 54 84.4

TOTAL 52 64

n: number of individuals.

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method, which takes only observable elements into account [67], should help to reduce possible biases.

The proportion of male pre-Columbian individuals is higher than that of females. This could be due to sampling issues as the site has not been exhaustively excavated, but could also be explained by cultural reasons such as a proba- ble preference of the site for male burials, a practice already documented in other archaeological sites on the Isthmus [68–71]. However, it is very interesting to note that the most complex funerary contexts are related to women [5].

Individuals less than 15 years old seem very numerous in the pre-Columbian Panamá Viejo sample (35%), indicating high mortality in childhood. However, these figures are com- parable to those obtained for other pre-Columbian popula- tions from the north of South America, which range from 40 to 60% [72], and in a typical archeological sample where around 50% of individuals are less than 15 years old [11].

Although paleoepidemiological studies have major lim- itations [11,26,36,37,67], the indicators analyzed here have brought out interesting features that can help to reconstruct aspects of the lifestyles of the early inhabitants of Panamá Viejo.

MSM

Information on MSM in the north of South America is scarce [73,74]. Table 5 shows data reported for series in this geo-

graphical area, reflecting seemingly high levels of MSM at Panamá Viejo. However, another specific study on the pre- Columbian period has shown that the frequencies in Panamá Viejo are comparatively lower than in the other observed series from the north of South America [73], although the same study also shows that the size of the Panamá Viejo series is a major limiting factor in drawing robust conclu- sions [73]. A tendency towards an increase in physical activ- ities with the arrival of the Europeans was observed in Panamá Viejo, similar to that observed in other contact con- texts in the Americas [75]. In the pre-Columbian sample, female individuals show higher frequencies of MSM, possibly indicating a more strenuous life for women. How- ever, no significant differences were found. Further study of MSM profiles and body mass using an enlarged sample could clarify the differences between the two archaeological periods.

DJD

There are few publications on the paleoepidemiology of DJD in South American osteological series [49,73]. Some studies on general paleopathology provide some data, but comparisons are difficult because the diagnostic criteria used are unclear [49,73]. However, it is worth comparing data even if only in a general way (see Table 5 for reported data [74,76–81]).

Table 3 Observed bone anomalies by period, by sex and by whole samples (top) and observed anomalies in adults only (bottom) / Anomalies osseuses par période, par sexe et pour les séries complètes (en haut) et anomalies exclusivement chez les adultes (en bas)

Pre-Columbian Colonial

Female Male Indet. Whole Female Male Indet. Whole

n % n % n % n % n % n % n % n %

MSM 2 28.6 5 19.2 0 0.0 7 14.9 5 31.3 7 35.0 7 25.0 19 29.7

DJD 3 42.9 3 11.5 2 14.3 8 17.0 5 31.3 4 20.0 5 17.9 14 21.9

Trauma 3 42.9 6 23.1 0 0.0 9 19.1 3 18.8 4 20.0 3 10.7 10 15.6

Infectious dis. 1 14.3 3 11.5 3 21.4 7 14.9 3 18.8 4 20.0 3 10.7 10 15.6

Porotic hyp. 1 14.3 7 26.9 1 7.1 9 10.6 1 6.3 0 0.0 1 3.6 2 3.1

Cultural 0 0.0 8 30.8 0 0.0 8 17.0 0 0.0 0 0.0 0 0.0 0 0.0

Pre-Columbian Colonial

Female Male Indet. Whole Female Male Indet. Whole

n % n % n % n % n % n % n % n %

MSM 2 40.0 5 23.8 0 0.0 7 21.9 5 33.3 7 36.8 7 35.0 19 35.8

DJD 3 60.0 3 14.3 2 28.6 8 25.0 5 33.3 4 21.1 5 25.0 14 26.4

Trauma 3 60.0 6 28.6 0 0.0 9 28.1 3 20.0 4 21.1 3 15.0 10 18.9

Infectious dis. 1 20.0 2 9.5 2 28.6 5 15.6 3 20.0 4 21.1 2 10.0 9 17.0

Porotic hyp. 1 20.0 6 28.6 0 0.0 7 21.9 1 6.7 0 0.0 1 5.0 2 3.8

Cultural 0 0.0 8 38.1 0 0.0 8 25.0 0 0.0 0 0.0 0 0.0 0 0.0

n: number of individuals. Indet: Indeterminate. MSM: Musculoskeletal Stress Markers. DJD: Degenerative Joint Disease. Infectius dis: Infectious disease. Porotic hyp: Porotic hyperostosis.

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Table4Chi-square,FisherandP-valuesforanomaliesobservedinthesamplesasawhole(top)andinadultsonly(bottom)/Chi2 etFisherpourlesdifférencesdefréquences entregroupes,pourlessériescomplètes(enhaut)etexclusivementpourlesadultes(enbas) Differencesbetween PeriodsSexesbyperiodPeriodbysexes Pre-Columbian andColonial Pre-Columbian Femalesand Males Colonial Femalesand Males PreColombian andColonial Females

PreColombian andColonial Males X2 p-valueX2 p-valueX2 p-valueX2 p-valueX2 p-value MSM3.30650.0690.62280.05620.8125-1.45790.2273 DJD0.40170.52620.09290.46970.65700.6816 Trauma0.23720.62620.0929-0.3185- Infectiousdis.0.01120.9158---0.6816 Porotichyp.0.13140.65160.4440.52570.0137 Cultural0.00070.1543--0.0201 Differencesbetweenadults PeriodsSexesbyperiodPeriodbysexes Pre-Columbian andColonial

Pre-Columbian Femalesand Males Colonial Femalesand Males Pre-Colombian andColonial Females

Pre-Colombian andColonial Males X2 p-valueX2 p-valueX2 p-valueX2 p-valueX2 p-value MSM1.83510.17550.58750.04520.8317-0.80680.3691 DJD0.02080.88520.06240.46200.34730.6889 Trauma0.98510.32090.3022-0.13130.7209 Infectiousdis.0.02670.87030.4885--0.3976 Porotichyp.0.0235-0.44120.44740.0212 Cultural0.00020.2805--0.0036 df=1.Italic:Fisherexacttest.Bold:Significant.MSM:MusculoskeletalStressMarkers.DJD:DegenerativeJointDisease.DJD:DegenerativeJointDisease.Infectiusdis: Infectiousdisease.Porotichyp:Porotichyperostosis.

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Table 5 Reported data from earlier American series /Données reportées pour des séries anciennes du continent

Series Date Reported data Reference

MSM

Ancón (Central Coast of Peru) 8% [74]

DJD

Highlands of Peru a rather common disease [76]

Mummies from Chile and Peru elevated frequencies

(young individuals)

[77]

Ancón (Central Coast of Peru) low presence [78]

Ancón (Central Coast of Peru) 76% [74]

Villa el Salvador (Central Coast of Peru) 79% male, 76% female [80]

Pre-Columbian Mochica 50% (56/112) [79]

Post-contact Mochica 75.6% (34/45) [79]

Muisca series (Andes in Colombia) 41.9% (31/74) [81]

Trauma

Ancón (Central Coast of Peru) 0% [78]

Ancón (Central Coast of Peru) 13% (15/116) [74]

Villa el Salvador (Central Coast of Peru) 10-20% cranial trauma [80]

Villa el Salvador (Central Coast of Peru) 22-29% post-cranial trauma [80]

Shamans from Peru and Chile 0% [77]

Male commoners from Peru and Chile 35% [77]

Female commoners from Peru and Chile 16% [77]

Santa Elena (Ecuador) BC 600 9% [83]

Guangala (Ecuador) BC 100 33% [83]

Ayalán without urn (Ecuador) AD 710 18% [83]

Ayalán with urn (Ecuador) AD 1230 13% [83]

Pecos Indians (USA) 4% long bones and hip fractures: [24]

Pecos Indians (USA) 7% cranial trauma added: [24]

North American pre-Columbian populations BC 4000-1000 10% postcranial factures [38]

North American pre-Columbian populations BC 1000-AD 1000 5% postcranial factures [38]

North American pre-Columbian populations AD 1000-1600 1-4% postcranial factures [38]

Central California 10% long bones fractures [82]

Central California 4% cranial fractures [82]

Central California 2% weapon wonds [82]

Central California 1% traumatic hip dislocation [82]

Infectious disease

Ancón (Central Coast of Peru) 36% (41/115) [74]

La Paloma (Central Coast of Peru) 8000-4500 BP 10-16% osteolysis [84]

La Paloma (Central Coast of Peru) 8000-4500 BP 22-25% periosteal reactions [84]

Peruvian shamans 9% osteolysis [77]

Peruvian males 20% osteolysis [77]

Peruvian females 18% osteolysis [77]

Santa Elena (Ecuador) BC 600 7% (9 bones/127 individuals) [83]

Guangala (Ecuador) BC 100 44% [83]

Ayalán without urn (Ecuador) AD 710 4% [83]

Ayalán with urn (Ecuador) AD 1230 14% [83]

Villa el Salvador (Central Coast of Peru) 43% periosteal reactions, males [80]

Villa el Salvador (Central Coast of Peru) 13% periosteal reactions, females [80]

North American mixed-economy populations 5% (59/1260) [34]

Pre-Columbian and postcontact Pecos Pueblo 0.8% (4 cases) osteomyelitis [24]

Pre-Columbian and postcontact Pecos Pueblo 2.6% (13 cases) periosteal reactions [24]

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The difference in frequencies between the two studies from Ancón [74,78], in the Peruvian Central Coast, has been attributed to the high proportion of young individuals in Kauffmann’s study sample [78].

According to the reported data (Table 5), the frequency of DJD found in the Panamá Viejo series seems comparatively low (17% for the pre-Columbian sample and 22% for the Colonial sample). This could be partially explained by the low proportion of old individuals and a moderate level of physical activity; but again, comparisons are difficult because of the sample size and poor preservation [73].

A tendency towards an increase in DJD (physical activi- ties?) with the arrival of Europeans was observed in Panamá Viejo, as also reported for other contact contexts in the Americas [75].

Trauma

Table 5 shows data from previous studies on trauma in the Americas [24,38,74,77,78,80,82,83].

The drop in fracture frequency reported by Steinbock’s study in 1976, cited by Merbs, may suggest changes in behavior when subsistence patterns shifted from hunter- gathering to agriculture [38]. Jurmain concluded that the high frequencies he observed compared with other studies were due to very intense interpersonal violence in the Cen- tral Californian population studied [82].

Our data from Panamá Viejo show a comparatively high frequency for both the pre-Columbian and Colonial periods observed. Trauma is more frequent in the group of pre- Columbian females, which could be associated with their occupations and with violent events (warfare?). However, the differences between sexes and periods are not signifi- cant. Further specific analysis of trauma in the Panamá Viejo series would help to better understand the lifestyle of the people who inhabited this site before and after the arrival of the Spaniards.

Infectious diseases

Table 5 shows the frequencies reported in previous studies on early populations from the Americas [24,34,74,77,80,83,84].

The frequencies found in Panamá Viejo for the pre- Columbian and Colonial samples seem comparatively moder- ate to low (around 15% for both periods), which may reflect a low population density and less harsh environmental condi- tions than previously thought, at least before and at the begin- ning of the Spanish presence in the region.

Porotic hyperostosis

Data from several South American series are summarized in Table 5 [27,33,74,80,83]. Blom et al. have argued that anemia was related more to environmental factors such as parasites and chronic diseases than to diet [33]. They consider that El Niño had a substantial influence on the Peruvian Coast because it could weaken the populations living there [33].

Comparatively, the frequency from Panamá Viejo seems moderate to low. Since it has been considered as a marker of anemia, this disease would also have a low frequency at this site. If the new perspective proposed by Walker et al. [30] is accepted, our data would show that consumption of animal food was high in both periods. Although a significant differ- ence was found between males from the pre-Columbian period and males from the Colonial period, this difference may be the result of a cultural practice among some males of the pre-Columbian period (see paragraphs concerning‘cul- tural’anomalies). However, isotopic studies (in preparation) will provide more material for discussion of the nutritional status of the early inhabitants of Panamá Viejo, helping to clarify this point.

‘Cultural’anomalies

As mentioned in the methodology section, skull deforma- tion, traces of cuts indicating possible defleshing and a

Series Date Reported data Reference

Porotic hyperostosis

Ancón (Central Coast of Peru) 7% [74]

Peruvian coast more frequent than in the highlands [27]

Villa El Salvador (Central Coast of Peru) 23% cribra: (N=402) [33]

Villa El Salvador (Central Coast of Peru) 11% cribra males; 19% females [80]

Villa El Salvador (Central Coast of Peru) 23% porotic hyperostosis males, 6% females

[80]

Santa Elena (Ecuador) BC 600 0% porotic hyperostosis, 0% cribra [83]

Guangala (Ecuador) BC 100 23% porotic hyperostosis, 10% cribra [83]

Ayalán without urn (Ecuador) AD 710 8% porotic hyperostosis, 0% cribra [83]

Ayalán with urn (Ecuador) AD 1230 7% porotic hyperostosis, 2% cribra [83]

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very particular bone anomaly in the skull (a circumscribed area of extreme porosity in the scalp, between marked temporal lines) have been grouped together as ‘cultural’ abnormalities.

Concerning this particular bone anomaly in the skull, two things were observed: intense porosity and marked temporal lines. These two anomalies have been very frequently observed in association, suggesting the hypothesis that the porosity in the scalp could be a mechanical consequence of hyperactivity in the masticatory muscle [85]. In our study, this abnormality appears in four male skulls; three of these belong to a very particular archaeological context as they were among nine male skulls offered as grave goods to one young female, who shows evidence of a very physically demanding lifestyle (fractures, very high MSM, DJD). The presence of anomalies classified as‘cultural’has brought out significant differences between the pre-Columbian and the Colonial periods in Panamá Viejo, particularly in males, showing differences in lifestyles and practices that concur with historical data [4].

The study of these bone indicators has demonstrated its usefulness in obtaining new information about early popula- tions in the Panamanian Isthmus, despite the limitation of the small size of the series. The analysis of stress indicators from teeth and diet may help to clarify the picture regarding the state of health of the early inhabitants of the Panamá Viejo site.

Conclusion

Although the statistical tests failed to prove significant dif- ferences between subgroups of individuals for some of the anomalies considered, the percentages nevertheless show discernible trends. The differences were not statistically con- firmed, probably because of the small size of some groups, particularly females. These results are complementary to those initially presented [4], but our conclusions here have been refined. Some changes in lifestyle with the arrival of Europeans at Panamá Viejo have been confirmed. Neverthe- less, more specific studies must be carried out to shed more light on the history of contact in this region. A study of the profiles and frequencies of MSM, DJD and trauma could clarify our understanding of changes over time in physical activities and other behavior. The analysis of teeth anomalies could also help to produce more robust interpretations of these data. Finally, these results support the need to enlarge the series and consequently to extend the archeological exca- vations at Panamá Viejo. Given the characteristics of the soil in this region,in situanalyses of human remains by an expe- rienced bioanthropologist would be a positive step to avoid further losses of information.

AcknowledgmentsThis research study was financed by the Proyecto Arqueológico Panamá Viejo, the Patronato Panamá Viejo and SENACYT (Secretaría Nacional de Ciencia, Tecnología e Innovación, Panamá). Thanks from the first author to the French Ministry of Higher Education and Research, which sponsored the research grant. We are indebted to Dr. Tomás Mendizábal, who kindly revised our manuscript. The authors also thank two anonymous reviewers for their comments, which greatly improved our manuscript, as well as the corrector, I. Bossanyi who revised the English.

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