ANTHROPOLOGICAL SCIENCE Vol. 114, 75–88, 2006

Maya skeletal remains from the Copan and El Puente sites in Honduras YUJI MIZOGUCHI1*, SEIICHI NAKAMURA2

1Department of Anthropology, National Science Museum, 3-23-1 Hyakunin-cho, Shinjuku-ku, Tokyo, 169-0073 Japan 2Copan Archaeological Project, Honduran Institute of Anthropology and History, Copan Ruins, Honduras

Received 31 March 2004; accepted 29 March 2005

Abstract The permanent teeth of two individuals from the 10J-45 compound in the Classic Maya site of Copan, Honduras, and the whole skeletons of two individuals from the El Puente site, a secondary Maya center of Copan, were morphologically observed and measured. Preliminary analyses of the well-preserved permanent teeth of the two El Puente individuals show that one is closest to Native South Americans and the other to Native North Americans. Although many human skeletal remains have already been excavated at these two sites, they have not fully been studied. In addition, many other archaeological sites in Honduras also remain to be investigated.

Key words: skull, permanent teeth, postcranial skeleton, Q-mode correlation, Native Americans

Introduction centers (Fash, 2001), and the El Puente site, a secondary Maya center (Nakamura, 1996). At the Copan site, some Many Maya archaeological sites have been found in the hundreds of skeletal remains have been uncovered up to the area of southeastern Mexico, Guatemala, Belize, western present (Maca and Rhoads, 2002). At least one series of Honduras, and western El Salvador. The most flourishing skeletal remains was excavated in 1938 and 1939, though period of the Maya civilization is called the Classic period, these were badly preserved (Longyear, 1940), and another which lasted from 250 to 900 AD. In this period the ancient was found between 1975 and 1985, and consists of at least Maya reached intellectual and artistic heights that surpassed 160 skeletons (Whittington, 1991). But, as far as the present anything found in the New World. Many city-kingdoms with authors know, no phylogenetic analysis based on morpho- their own ruling dynasties flourished in this period. Each logical characters or measurements has been carried out for kingdom had many populations and its rulers engaged in these materials, although several paleopathological or paleo- long-distance trade of exotic and rare objects such as jade demographic studies have been conducted (Storey, 1988, and marine shells. 1994; Whittington, 1991, 1992). In this report some basic However, at the end of the period these great cities of the data of ancient Maya skeletal remains are presented to accu- Classic Maya civilization declined and were finally aban- mulate information for use in future phylogenetic or other doned. The supporting populations never returned to the anthropological analyses. same place in the subsequent Postclassic period (900–1521 AD). This chain reaction of collapse and abandonment of the Classic kingdoms is referred to as the ‘collapse of Classic Materials Maya civilization’. Although various theories and hypothe- Of the four individuals observed and measured here, two sis have been proposed to explain this collapse (Culbert, are part of the remains unearthed at the Copan site between 1973), it is believed that some combination of factors, such 1999 and 2002, and the other two are part of the remains as external conflicts among the Classic Maya societies found at the El Puente site during the period 1990 to 1994. (wars), elite revolt against kings (Fash, 2001), climatic All these human remains were excavated under the direction change (Haug et al., 2003), and destruction of the environ- of S.N., as stated above. Both sites are located in the west- ment due to population growth, played a principal role in the ernmost area of Honduras (Figure 1). collapse. The identification numbers of the former two individuals The human skeletal remains reported here are four indi- are Copan 35-2000 (Burial #35-2000 from the 10J-45 com- viduals from the Classic period. They are part of some doz- pound of the Copan site) and Copan 36-2000 (Burial #36- ens of the remains which had been excavated by the middle 2000 from the 10J-45 compound of the Copan site), and of October, 2002, by a project team directed by one of the those of the latter two are El Puente 10-3 (Burial #3 from present authors (S.N.) at two sites of the Classic period in Structure 10 of the El Puente site) and El Puente 204-1 Honduras, i.e. the Copan site, one of the major Classic Maya (Burial #1 from Structure 204 of the El Puente site). The two individuals from the Copan site are thought to date back to * Corresponding author. e-mail: [email protected] around 550 AD on the basis of archaeological evidence phone: +81-3-5332-7170; fax: +81-3-3364-7104 (Nakamura, 2002, 2003), and the two individuals from the Published online 29 June 2005 El Puente site are of the Late Classic period, i.e. 600–900 in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.040331 AD (Nakamura, 1996).

© 2005 The Anthropological Society of Nippon 75 76 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE

and

Stature [in cm] = 1.96 (Lateral condyle–malleolus length of tibia [in cm]) + 93.752 − 2.5 ± 2.815.

In order to determine the affinities of ancient Mayan indi- viduals in relation to the comparative samples of various populations around the world, Q-mode correlation coeffi- cients (Sneath and Sokal, 1973) were calculated using the crown diameters of permanent teeth. The significance of the Q-mode correlation coefficients, strictly speaking, the z- transformed Q-mode correlation coefficients, was tested by the bootstrap method (Efron, 1979a, b, 1982; Diaconis and Efron, 1983; Mizoguchi, 1993). In order to estimate the bootstrap standard deviation of a z-transformed Q-mode cor- relation coefficient, 1000 bootstrap replications including the observed sample were used. The bootstrap standard devi- ation was obtained by directly counting the cumulative fre- quency for the standard deviation in the bootstrap distribution. Incidentally, it may be worthy of notice that the bootstrap method can be used for testing the significance of any statistic even if its distribution is different from a normal Figure 1. Maya region and principal archaeological sites. distribution. Statistical calculations were executed with the mainframe HITACHI MP5800 system of the Computer Centre, the Uni- El Puente 10-3 and 204-1 were observed and measured versity of Tokyo. The programs used are SDQCPC for R- for almost all major bones and permanent teeth. But, regard- mode standardization, BTPCA for estimating Q-mode corre- ing Copan 35-2000 and 36-2000, data were taken only on lation coefficients and for bootstrap tests of their signifi- the permanent teeth because the time available for observa- cance, and CLSTR for cluster analysis. All these programs tion was extremely limited. were written in FORTRAN by Y.M.

Methods Brief Description Observation, measurement, and statistical analysis were The basic observations and measurements of the individ- conducted by Y.M. Cranial and postcranial measurements ual skeletal remains are shown in Appendices 1 to 8. Appen- were taken after Martin and Saller (1957), except for nasal dix 1 gives the cranial measurements; Appendix 2, scores of height, which was measured according to Howells (1973). non-metric cranial characters; Appendix 3, preservation of The mesiodistal (MD) and bucco- or labiolingual (BL or LL) permanent teeth and degree of occlusal wear of the first diameters of a tooth crown were measured with sliding cali- molars; Appendix 4, measurements of permanent teeth; pers to the nearest 0.1 mm according to Fujita’s (1949) Appendix 5, scores of non-metric tooth crown characters; method. The mesiodistal diameter of the dental cervix was Appendix 6, postcranial measurements; Appendix 7, obser- measured after Mizoguchi’s (1985) definition. The criteria vations of postcranial bones; and Appendix 8, stature esti- of classification for non-metric tooth crown characters can mates. be found in Mizoguchi (1977). The degree of occlusal wear The results of sex and age determinations and some other on the crown surface was recorded according to Broca’s information are briefly presented below. grading system (Martin and Saller, 1957). As for the terminology of teeth, the following abbrevia- Copan 35-2000 tions are used. I, C, P, and M designate the permanent inci- Female (?), senile (?). Although many bone fragments are sor, canine, premolar, and molar, respectively; numbers 1, 2, preserved, only the permanent teeth were observed and mea- and 3 indicate the position of the tooth within a tooth class; sured. and U and L means ‘of the maxilla’ and ‘of the mandible’, The sex determination is based on a fragment of the ilium respectively. For example, UM2 means the maxillary second around the greater sciatic notch, the mastoid process, and the molar. mandible. The iliac fragment suggests that this individual is Stature was estimated using the formulae for living Native a female, but the mastoid process is large and the mandible Mexican males, originally developed by Genoves (1967) is not small, implying the opposite sex. The alveoli for the and corrected according to the comments by Bass (1971). following teeth are closed: the right and left UI1s, UI2s and The formulae are as follows. UCs, the right UM2 and UM3, the right LI1, the left LI1 and LI2, and the left LM3. The closed alveoli and Broca’s score Stature [in cm] = 2.26 (Maximum length of femur of 2 for the LM1 (Appendix 3) suggest that this individual is [in cm]) + 66.379 − 2.5 ± 3.417 not a young adult. Vol. 114, 2006 MAYA SKELETAL REMAINS FROM HONDURAS 77

Although a mandibular premolar had been preserved in the first molars and the erupted third molars (Appendix 3), addition to the permanent teeth shown in Appendix 3, the does not contradict this age estimation. Stature is estimated tooth was given for DNA analysis. This individual is consid- at 158.75 cm (Appendix 8). ered a sacrifice from archaeological evidence (Nakamura, Other observations and notes are as follows. The external 2002). surface of the squama frontalis is slightly depressed. This is considered to be evidence for an artificial anteroposterior Copan 36-2000 deformation of the skull. Although the left LP1 was Male (Buikstra, 2002), adult. Although some bone frag- extracted for DNA analysis, it was measured before extrac- ments are preserved, only the permanent teeth were tion (Appendix 4). A few lines of enamel hypoplasia are dis- observed and measured. cernible on the crown surfaces, especially of the maxillary Sex could not be determined by the present authors, anterior teeth and the mandibular canines. Dental caries are because the time available for observation was extremely not found in any of the observed permanent teeth. The limited. According to Buikstra (2002), however, this indi- medial side of the lowermost part of the body of the right vidual is identified as a male. Broca’s score of 2 for the UM1 femur is abnormally swollen. (Appendix 3) suggests that this individual is an adult. In addition to the twelve permanent teeth shown in El Puente 204-1 Appendix 3, some root fragments are also preserved. Fur- Male, adult. Both cranial and postcranial skeletons are ther, although the right UM3 was preserved, it was used for almost perfectly preserved, but the cranium lacks the lower DNA analysis. A relatively rare non-metric tooth crown halves of the nasal bones, the frontal processes of the maxil- character, i.e. a developed paramolar tubercle, was observed lae, the basilar and right lateral parts of the occipital bone, on the left UM2, in addition to the characters shown in most of the sphenoid bone, and the left condylar process of Appendix 5. the mandible (Figure 3). This individual is believed to have been one of the rulers, This individual was identified as male on the basis of mor- presumably the eighth or ninth, of the Copan Dynasty from phological characters of the skull and pelvis. Broca’s scores archaeological evidence (Nakamura, 2003). of 2 for all the M1s (Appendix 3) suggest that this individual is a mature male. But the left UM3 and the right and left El Puente 10-3 LM3s seem to be congenitally absent, though agenesis of the Male, around 20 years old. Both cranial and postcranial right UM3 could not be confirmed because of damage to the skeletons are almost perfectly preserved, but the cranium alveolar process. The estimated stature is 159.90 cm lacks the nasal and sphenoid bones (Figure 2). (Appendix 8). This individual was identified as male on the basis of mor- Other observations or notes are as follows. The external phological characters of the skull and the pelvis. No epiphy- surface of the squama frontalis is clearly depressed. This is seal lines are found on most limb bones, but both sternal and considered to be evidence for the artificial anteroposterior acromial ends of the clavicle are not fused with the shaft. deformation of the skull. Although the left LP1 was This suggests that this individual died at the age of around extracted for DNA analysis, it was measured before extrac- 20 years. Dental evidence, i.e. Broca’s scores of 1 or 2 for tion (Appendix 4), as with El Puente 10-3. A line of enamel

Figure 2. The skull of El Puente 10-3 (photographed by Hiroyuki Ikarashi). (a) Lateral view (right); (b) frontal view; (c) lateral view (left). 78 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE

Figure 3. The skull of El Puente 204-1 (photographed by Hiroyuki Ikarashi). (a) Lateral view (right); (b) frontal view; (c) lateral view (left). hypoplasia is found on the crown surfaces of the maxillary The second largest number of variables available was anterior teeth and the mandibular canines. Dental caries are twelve for El Puente 204-1 (Table 1). In order to assess not found in any of the observed permanent teeth. On the whether or not a classification based on these 12 variables labial crown surfaces of the right and left UI1s, a hemi- corresponds to that based on the above 22 variables, the spheric jadeite is inserted. On each of the labial crown sur- same analysis was conducted for El Puente 10-3, this time faces of the right and left UI2s, UCs and LCs, there is a small using 12 variables (Table 3). From Tables 2 and 3, it can be hole. This is also considered to be evidence of jadeite inser- seen that the first four samples closest to El Puente 10-3 tions. remain the same. Therefore, it may be considered that the results of a 12 variable analysis of El Puente 204-1 are also Preliminary Comparisons of Tooth Crown likely to be reasonable. From the 12 variable analysis, it is Diameters seen that the people closest to El Puente 204-1 are the early modern Native Americans in South Dakota, USA, or the Because of small sample size and artificial deformation of Classic/Post-Classic Southampton Islanders of Canada the well-preserved skulls, it is difficult to precisely deter- (Table 4). mine the phylogenetic origins of the present sample and Similarly, reliability of an 8 variable analysis of Copan their relations with other populations around the world. 36-2000 was examined by comparing results of the 22 and 8 However, we did carry out a preliminary analysis of affini- variable analyses of El Puente 10-3 (Table 2, Table 5). This ties using the crown diameters of permanent teeth. comparison showed that the results of the 8 variable analysis In Table 1, the crown diameters of the permanent teeth did not resemble that of the 22 variable analysis. Therefore, used for comparisons are shown for the three observed an analysis of Copan 36-2000 based on the same 8 variables ancient Mayan males and 36 comparative male samples (Table 6) is not considered to be reliable. from various regions of the world. Copan 35-2000 was From the above considerations, from dental measure- excluded from this analysis because the number of teeth ments, nothing should be said of the affinities of the Copan available was too small. Neither the MD and BL crown individuals. On the other hand, it is likely that people from diameters of the third molars nor the LL crown diameters of the El Puente site are phylogenetically closer to Native the anterior teeth were used, because these measurements South or North Americans than to other populations of the have not been reported for many of the comparative sam- world, as was expected. ples, especially when measurements were taken on plaster Incidentally, Shinoda (2003, n.d.), who analyzed the D- casts. loop region of mtDNA extracted from teeth of 13 Copan and The number of variables (or crown diameters) available El Puente individuals, including El Puente 10-3 and 204-1, varies among the Mayan specimens, as shown in Table 1. found a total of 13 mutations within 192 base pairs and iden- First, the interrelationship between El Puente 10-3 (for tified nine different haplotypes among individuals. He con- which the largest number of variables was available) and the firmed that all examined individuals could be classified into comparative samples was investigated using all 22 variables. haplogroup A or B. The frequencies of haplogroups A and B The sample closest to El Puente 10-3 was found to be Ticuna are very high in contemporary Native American populations from Colombia (Table 2), and the Q-mode correlation coef- of Mesoamerica including the contemporary Mayan people. ficient between the two was statistically significant. This Shinoda suggested that composition of mtDNA in the sam- seems reasonable from a geographical viewpoint. ple that he tested was similar to that of the present-day Vol. 114, 2006 MAYA SKELETAL REMAINS FROM HONDURAS 79 3 n Mat- 10 7 1 Ainu Prakash et Hokkaido 25 Mean 10 n Mizoguchi (2000), 32 Pre-Incan Peruvians Mean Matsumura (1995), an early Sharma (1983). 19 24 9 of the Postclassic period. n Native Mizoguchi (1993). 31 Surinamese Mayhall (1979), a sample from Igloolik. Mean e prehistoric Jomon period from all regions 2 8 n e protohistoric Kofun period from eastern Japan. Matsumura (1995). samples from various regions of the world 23 Ticuna (Colombia) Mean Mijsberg in Moorrees (1957). Mijsberg 7 Matsumura (1995), a sample 6 18 n es are listed in mm. Mexicans Mean Mizoguchi (1993). 22 n 6 Rosenzweig and Zilberman (1969). 30 Potter et al. (1981). Native (USA) early modern sample. 17 Matsumura (1995), a pooled sample of th averages in 36 comparative male Californians 12 Mean Matsumura (1995), a pooled sample of th n 5 14 Liu (1977). (1977). Liu Native (USA) Alvesalo (1985). (1985). Alvesalo 16 the left side were used. Mean valu Illinoisans 29 Mean n Mizoguchi (2000). 5 Matsumura (1995), an 11 5 from Yuendumu, Northern Territory. Northern Territory. from Yuendumu, (USA) Dakotans Mean Native South were not available, n 4 Moorrees (1957). Aleuts e ancient Maya males and crown diameter 4 (USA) Matsumura (1995). 10 Mean 3 n Selmer-Olsen in Moorrees (1957). Selmer-Olsen prehistoric Yayoi period from western Japan. prehistoric Yayoi 28 Japanese and Chinese samples. Islanders (Canada) Mean Southampton isons. If the right side teeth Taverne (1980). Taverne 9 2 n e permanent teeth of thre Townsend and Brown (1979), a sample Townsend 21 Inuit Mizoguchi (1993). (Canada) e Classic to Postclassic period. mple of the protohistoric Kofun period from western Japan. 27 Mean El 204-1 Puente (Male) ndard deviations obtained from El Bailit et al. (1968). 10-3 Harris and Nweeia (1980). 20 Puente (Male) 8 Sakai et al. (1971). Ancient Maya Matsumura (1995), a pooled sample of the 26 36- 13 2000 Table 1. Crown diameters of th Table Copan (Male) The right side teeth were used for compar 1 UI1UI2 —UC —UP1 7.9 —UP2 7.8 7.3UM1 — 7.4 8.1UM2 — 7.1 —LI1 9.6 6.3 8.7 — 10.7 —LI2 7.2 138 — 9.1 —LC 8.2 — 125 — 8.88 7.3LP1 10.8 5.0 — 7.36 119 6.8 11.1LP2 14 110 — 10.4 6.0 8.35LM1 17 105 116 7.31 — — 8.45 7.1 10.93LM2 — 90 35 7.00 — 6.8 7.29 38 97 — 43 9.90 5.3 7.2 8.03 35 8.3 11.2 88UP1 7.15 10.37 8.78 6.4 7.5 119 41 10.1UP2 81 6.65 7.4 7.39 7.0 9.9 77 — 53 124 10.00 27 5.30 7.2UM1 9.8 8.43 62 — 10.67 117 11.2 34 6.30 7.24 7.3UM2 11.7 51 8.88 9.6 109 11 11.7 7.62 39 6.89LP1 12.1 38 11.3 7.46 8.8 17 103 117 43 7.28 9.88 54 10.0 5.23LP2 10.64 12.21 11.4 36 8.42 39 — 7.30 12.9 62 6.09 83 7.35 39 41 9.00LM1 98 9.8 9.5 62 — 44 12.6 11.39 7.20 11.8 100 71 6.99 36 7.61LM2 — 75 7.01 9.65 50 11.08 11.56 5.40 118 7.0 11.8 9.3 10.0 123 43 91 6.22 8.55 72 7.17 60 8.0 7.57 12.36 94 9.33 71 73 47 9.00 10.5 104 11.19 29 109 7.48 8.1 37 69 7.28 9.4 11.98 81 10.23 10.76 11.88 7.34 77 41 7.13 37 9.15 33 43 11.9 5.45 9.1 41 6.33 8.15 77 11.34 42 8.0 11.5 7.36 80 34 42 32 11.37 7.44 11.3 41 9.31 31 8.73 51 11.36 8.5 10.39 11.74 11.2 7.42 64 35 45 111 7.20 42 9.85 44 7.45 114 35 7.20 81 9.13 23 5.69 109 12.22 42 11.24 8.21 27 61 11.19 76 6.68 8.46 35 7.39 83 35 25 7.54 77 9.54 12.09 69 8.62 10.97 8.49 12.45 49 10.64 40 45 41 68 7.61 67 26 6.98 76 9.81 7.12 27 12.01 42 7.42 42 10.56 9.49 28 28 33 69 5.60 11.91 43 7.82 66 6.40 8.53 27 7.60 11.76 26 10.79 28 11.87 10.58 7.35 66 71 9.62 49 40 9.00 8.40 33 75 86 7.23 34 28 7.03 12.43 71 73 9.92 11.16 52 7.48 35 11.31 45 28 7.35 77 9.52 25 79 5.33 12.34 8.01 10.26 34 11.32 6.36 10.77 8.34 28 74 7.58 46 28 27 35 7.62 35 9.65 74 8.10 8.47 25 11.93 80 43 10.24 11.00 33 24 7.24 10.68 25 45 38 7.27 35 6.91 51 77 7.17 9.59 12.02 19 42 5.29 11.68 10.61 35 73 48 8.13 27 24 6.24 7.56 49 7.24 21 6.87 27 9.45 76 11.72 35 11.45 8.50 29 29 10.66 75 78 9.12 7.32 29 27 6.45 27 11.50 11.99 32 11.06 35 7.12 27 9.53 71 78 5.47 29 34 8.25 29 6.26 34 7.19 11.69 26 10.80 60 11.61 75 29 9.67 35 8.48 38 72 11.41 7.30 50 11.54 10.44 29 28 35 57 27 7.39 9.38 72 5.00 12.19 8.15 28 10.51 60 34 5.72 10.74 35 7.69 63 9.56 27 12.13 8.60 22 10.32 53 35 24 6.71 22 33 63 28 6.85 9.25 11.58 50 10.81 35 24 7.86 30 6.81 11.26 34 10.64 9.78 28 33 29 8.19 27 34 34 11.36 49 29 9.63 27 7.89 10.92 60 9.04 47 8.26 29 10.97 57 32 8.76 29 10.28 8.08 34 34 8.58 33 62 64 7.58 8.05 33 34 Matsumura (1995), a pooled sa MD of: BL of: of Japan. modern sample. Matsumura (1995), a sample of th 15 sumura (1995). mean within-group sta al. (1979). 80 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE 22 n Chinese Mean 19 n Mean Mongolians 21 n Australian Aborigines Mean 20 n Nasioi Islands) (Solomon Mean 19 n Islanders New Britain Mean 18 n Javanese Mean 17 n 2 82 10.8 129 11.28 87 11.12 86 11.20 208 10.31 49 10.35 580 2 77 10.5 133 10.94 77 10.99 66 11.53 169 10.44 35 10.60 468 (Phil- 11 82 11.8 130 12.86 87 12.10 80 12.55 208 11.78 34 11.46 579 Tagalog ippines) Mean 16 n Table 1. (continued 2) Table Atayal (Taiwan) Mean 15 n Japanese (W Japan) Mean Kofun-period 14 n Japanese (E Japan) Mean Kofun-period 13 n Japanese (W Japan) Mean Yayoi-period Yayoi-period 12 n Japanese Mean Jomon-period 11 n Japanese Mean UI1UI2 8.61UC 7.11 1072UP1 1063 8.51 8.03UP2 7.40 7.10 108UM1 968 1157 6.96 10.56 106 8.81 7.55 6.90 7.44 1115 1111 57 10.28 153 68 6.46 50 8.58 190 7.59 8.17 10.68 183 7.17 19 77 65 7.10 68 23 8.84 10.60 7.47 8.15 67 7.44 26 30 34 27 7.11 10.85 27 7.54 8.7 8.19 34 37 7.1 33 151 33 7.16 10.8 151 8.33 7.4 8.1 152 36 6.76 87 10.0 152 152 94 7.0 8.6 6.89 7.75 7.0 147 82 129 84 131 6.56 9.14 7.5 8.0 7.59 32 85 133 136 45 9.00 7.76 7.0 8.66 7.71 68 76 66 129 79 9.36 7.55 7.27 8.60 206 7.54 81 77 75 203 8.40 7.63 7.05 8.25 6.99 23 200 192 25 89 8.65 7.21 7.87 6.98 459 7.13 47 42 429 191 7.28 7.99 6.62 545 507 52 6.82 460 UM2LI1 9.85LI2 1018 5.50LC 9.12 6.16 1046LP1 172 1043 5.27 7.12LP2 9.86 7.27 5.72LM1 61 932 7.31 60 1051 11.56LM2 91 5.44 6.73 1062 6.91 11.12 9.92 919 6.19 112 6.94 54 172 11.61UP1 27 841 7.24 71 190 7.38 10.80 5.52 210UP2 9.81 9.60 11.82 7.49 85 6.19 201 79UM1 17 9.46 1157 11.35 34 11.70 66 84 7.16UM2 32 7.38 5.61 9.27 1107 11.67 11.79 67 1115LP1 7.48 32 11.78 9.9 6.30 9.00 34 153 11.21 22 39LP2 975 189 36 8.07 7.28 184 9.74 114 31 11.91 7.42 11.45 35 12.06LM1 5.5 8.51 1057 9.52 11.39 7.56 10.97 9.24 175 36 77 38 38LM2 76 6.3 11.84 1070 7.79 153 10.58 72 77 35 12.10 38 9.77 11.3 925 7.0 8.33 151 173 61 7.0 5.08 11.23 9.65 10.0 10.7 855 35 31 132 11.77 193 8.35 7.2 150 10.47 5.74 218 95 156 12.31 10.73 125 31 129 11.33 9.77 8.76 29 207 95 83 152 6.77 10.08 6.77 10.24 57 36 10.73 12.10 5.5 9.69 79 35 86 94 6.75 8.24 86 96 77 11.5 11.28 6.2 11.7 70 35 128 36 8.61 81 10.31 35 10.9 10.79 9.7 7.2 114 133 47 150 7.3 5.65 11.6 61 39 12.28 9.6 11.40 11. 8.52 132 39 153 7.3 139 6.41 133 23 118 10.83 11.51 10.96 72 8.95 154 40 38 9.42 11.04 131 38 7.52 156 7.38 80 5.70 11.77 35 38 9.28 82 77 10.9 51 9.48 7.54 63 6.36 8.0 10.99 64 84 85 10.6 131 11.7 60 8.7 48 66 69 12.05 7.52 9.8 135 7.50 10.36 5.83 130 125 205 9.7 11.44 75 154 9.83 7.59 132 82 7.78 10.1 206 6.56 57 11.52 12.84 176 10.44 129 87 518 8.11 206 5.32 96 7.44 11.01 11.0 84 7.46 51 10.24 74 6.06 81 198 196 26 7.55 11.32 12.35 46 117 8.2 75 10.28 7.07 10.89 185 11.51 7.25 27 535 67 5.53 8.5 10.30 75 132 7.23 74 468 45 12.77 6.12 44 399 82 10.25 131 8.75 154 11.09 48 400 7.22 199 10.23 7.17 11.60 9.15 62 86 191 451 9.44 7.17 502 60 35 11.81 8.38 9.18 441 35 11.53 206 8.92 71 11.09 39 517 9.51 82 8.82 36 9.32 543 10.89 196 9.25 457 183 8.27 536 8.47 37 36 8.09 8.26 503 440 MD of: BL of: Vol. 114, 2006 MAYA SKELETAL REMAINS FROM HONDURAS 81 32 S.D. d.f. East Asians 31 n European Americans Mean 31 n Moroccans Mean n 30 (Israel) Bedouins Mean n 29 Finns Mean n 28 Lapps Mean n 27 527 9.34 267 10.0 93 10.03 35 10.15 52 9.92 140 0.55 269 Indians Mean 26 n Table 1. (continued 3) Table Pashtuns (Afghan.) Mean 26 n Tajiks (Afghan.) Mean 25 n Bhutanese Mean 24 n Tibetans Mean 23 n Neolithic Mean Thailanders 23 n Mean Thailanders UI1UI2 8.31UC 7.01 20UP1 8.09 29UP2 8.62 7.58 38UM1 6.96 7.15 15 41 10.44UM2 7.84 14 41 8.21 39 7.31 9.69 18 10.58 6.46 6.65 33 21 41 7.28 33 21 22 8.42 6.88 9.34 10.67 33 6.91 35 6.73 33 19 33 7.67 35 8.34 33 10.07 6.93 9.41 35 6.56 36 6.49 35 35 25 7.59 36 8.49 35 9.94 6.59 8.87 36 6.77 6.38 64 35 38 35 7.82 10.31 67 8.79 35 6.83 8.92 71 6.87 71 591 6.53 73 37 10.73 590 7.94 8.37 71 7.14 9.14 577 6.84 570 10.23 6.76 73 569 68 7.74 256 95 559 6.75 8.9 9.61 194 10.3 6.45 221 6.9 185 8.1 165 237 157 8.76 10.63 7.1 122 6.89 6.9 145 83 78 7.80 10.82 126 7.02 80 8.84 58 6.75 93 6.97 62 92 8.09 10.51 60 7.30 93 8.68 152 70 7.06 6.73 83 206 0.52 7.86 76 188 0.50 6.96 336 192 0.59 6.81 340 177 0.43 339 157 0.38 335 0.39 337 332 LI1LI2 5.56LC 6.20 25LP1 7.01 26LP2 5.39 7.39 38LM1 6.10 7.41 19 43 11.59LM2 7.09 21 43 10.98 5.20 34 7.12 24 11.89 5.56 39 7.09UP1 33 25 10.64UP2 6.35 21 33 25 9.63 5.22 7.00UM1 11.25 23 33 9.49 5.87 40 35 7.15 11.62 33UM2 30 9.98 39 6.94 35 11.71 42 5.12 9.48 33LP1 10.65 6.97 12.04 23 35 41 5.74 9.28LP2 36 6.85 19 35 35 8.29 10.08 11.56 6.73LM1 21 10.64 34 21 5.36 35 8.56 7.00 6.64 42 10.84 35LM2 18 37 6.02 9.15 27 7.22 6.93 46 54 10.51 33 38 37 8.11 9.82 10.97 8.37 6.93 67 33 11.10 33 44 5.80 34 8.47 9.18 6.88 35 25 49 11.16 25 10.52 72 6.17 580 10.18 9.05 22 6.97 11.22 26 35 73 11.11 5.88 35 583 6.96 23 5.36 35 63 577 68 9.20 11.14 8.86 6.21 35 7.13 33 10.95 5.98 10.49 578 8.01 11.26 8.79 7.19 35 30 38 76 574 33 7.94 6.82 228 532 123 10.64 11.27 38 23 35 564 6.72 10.51 8.88 8.25 5.6 11.3 35 219 10.33 11.39 35 68 6.1 6.74 9.01 226 254 72 35 155 10.47 133 7.62 10.98 35 68 7.1 172 11.27 232 10.8 71 5.45 10.04 8.86 8.10 10.66 7.2 573 26 37 6.09 150 11.24 75 8.95 10.36 7.3 565 528 33 167 75 34 83 7.69 7.03 11.25 10.96 77 10.49 10.11 231 557 8.91 143 7.03 51 8.10 5.60 73 59 258 11.6 10.34 91 6.10 39 8.72 7.21 61 203 63 68 93 10.96 7.32 11.6 11.14 574 168 7.12 208 92 9.96 51 11.54 10.40 9.3 7.27 8.04 117 566 5.46 54 81 532 94 5.47 9.5 7.54 246 147 560 78 0.51 7.39 84 11.55 10.07 10.91 248 252 6.92 11.63 135 10.8 9.20 7.75 70 330 128 0.34 226 259 6.96 35 0.36 9.42 257 137 94 62 217 11.62 0.61 10.5 7.30 326 235 10.77 8.0 327 0.38 11.37 60 237 171 9.41 0.40 53 8.6 161 77 75 339 163 11.33 10.44 9.71 0.42 341 10.84 83 143 7.96 0.52 156 329 39 76 8.49 8.77 93 63 335 0.57 10.79 10.53 9.21 51 184 8.10 232 55 127 170 0.52 8.78 84 10.37 0.47 0.52 335 71 137 7.71 332 327 0.49 8.33 232 246 181 0.50 0.49 339 330 MD of: BL of: 82 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE

Table 2. Five samples closest to El Puente 10-3 among 36 compara- Table 6. Five samples closest to Copan 36-2000 among 36 compara- tive samples from various regions in the world, suggested by the Q- tive samples from various regions in the world, suggested by the Q- mode correlations based on 22 crown diameters of permanent teeth1 mode correlations based on 8 crown diameters of permanent teeth1 Q-mode correlation Q-mode correlation Ticuna (Colombia) 0.3736* Thailanders 0.8970*** Native Illinoisans (USA) 0.3674 Ticuna (Colombia) 0.7535** New Britain 0.3347 Javanese 0.7081* Native South Dakotans (USA) 0.3063 Japanese 0.6259 Surinamese 0.2742 Atayal (Taiwan) 0.6144* 1 The 22 measurements used are as follows: MD crown diameters of 1 See footnote to Table 5. the maxillary and mandibular I1 to M2, and BL crown diameters of the * P < 0.05; ** P < 0.01; *** P < 0.001, by a two-tailed bootstrap maxillary and mandibular P1 to M2. test. * P < 0.05, by a two-tailed bootstrap test.

Mayan people, and that a direct genetic progression had con- Table 3. Five samples closest to El Puente 10-3 among 36 compara- tinued from pre-Columbian times. tive samples from various regions in the world, suggested by the Q- mode correlations based on 12 crown diameters of permanent teeth1 Numerous human skeletal remains have been excavated at the Copan and El Puente sites. But they have not yet been Q-mode correlation fully studied. In addition, in Honduras, many other archaeo- Native Illinoisans (USA) 0.4623* logical sites remain to be investigated. We hope that the Ticuna (Colombia) 0.4270* materials from these sites are exhaustively analyzed, so that New Britain 0.3956 a better understanding can be attained on the origins, evolu- Native South Dakotans (USA) 0.3894 tionary processes, migration routes, etc., of both American Mexicans 0.2866 and Asian peoples. 1 The 12 measurements used are as follows: MD crown diameters of UM2 and LC to LP2, and BL crown diameters of the maxillary and mandibular P1 to M2. Summary * P < 0.05, by a two-tailed bootstrap test. Results of observations and measurements of the cranium, postcranial bones, and permanent teeth of four individuals from two Mayan archaeological sites in Honduras are pre- Table 4. Five samples closest to El Puente 204-1 among 36 compara- tive samples from various regions in the world, suggested by the Q- sented as comparative data for future anthropological stud- mode correlations based on 12 crown diameters of permanent teeth1 ies. Q-mode correlation Native South Dakotans (USA) 0.6644*** Acknowledgments Southampton Islanders (Canada) 0.6100** Data of the human skeletal remains reported here were Lapps 0.5736** obtained in the process of preparing an exhibit called Inuit (Canada) 0.5456*** Native Californians (USA) 0.5350* “Maya: Kingdoms of Mystery” held in Tokyo in 2003 under 1 the auspices of the National Science Museum, Tokyo, Tokyo See footnote to Table 3. The Q-mode correlation coefficient Broadcasting System, and Asahi Shimbun. We would like to between El Puente 204-1 and 10-3 based on 12 crown diameters of permanent teeth is 0.0997 (bootstrap P = 0.7813). thank all participants for giving us the opportunity of * P < 0.05; ** P < 0.01; *** P < 0.001, by a two-tailed bootstrap observing the material, and two anonymous reviewers of the test. original manuscript.

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Appendix 2. Non-metric cranial characters of two adult individuals Appendix 1. Cranial measurements of two adult individuals 1 from the El Puente site (in mm)1 from the El Puente site Martin’s No.2 El Puente El Puente Characters El Puente 10-3 El Puente 204-1 10-3 (Male) 204-1 (Male) (Male) (Male) 1 Maximum cranial length — 165 Right Left Right Left 8 Maximum cranial breadth (153) 155 Supraorbital foramen 12 113 0 9 Minimum frontal breadth 95 98 Foramen of Huschke 0 0 0 0 10 Maximum frontal breadth 119 121 Hypoglossal canal bridge 0 0 — — 11 Biauricular breadth — 136 Mylohyoid canal 0 0 0 0 14 Minimum cranial breadth — (68) 1 20 Porion–bregma height — 116 1, Present; 0, Absent. 2 Two foramina, only one of which perforates into the orbit. 23 Horizontal circumference — 500 3 24 Transverse arc — 323 It is unknown whether or not the foramen perforates into the orbit 26 Frontal arc 122 111 because the orbit is damaged. 27 Parietal arc — 116 28(1) Lambda–inion arc — 64 29 Frontal chord 110 103 30 Parietal chord — 103 31(1) Lambda–inion chord — 62 43 Upper facial breadth 106 (108) 44 Biorbital breadth — 98 46 Middle facial breadth 106 95 47 Total facial height — 120 48 Upper facial height — 76 52 Orbital height — (35) 54 Nasal breadth 25 — NLH3 Nasal height — 52.5 57 Minimum nasal breadth — 10 60 Maxillo-alveolar length (52) — 61 Maxillo-alveolar breadth 65 64 63 Palatal breadth 41 (34) 63(2) Anterior palatal breadth 32 32 66 Bigonial breadth 97 — 69 Symphysial height 30 (29) 69(1) Mandibular body height 31 33 69(3) Mandibular body thickness 10 12 70(3) Mandibular notch height 10 13* 71 Minimum ramus breadth 33 34 71(1) Mandibular notch width (32) (36*) 1 When measurement could be taken on both sides, only the left side was measured. If the left side was not available, the right side was measured. The values in parentheses may not be accurate. Note that these two skulls are artificially deformed. 2 Martin and Saller (1957). 3 Defined by Howells (1973). * Measured on the right side. Vol. 114, 2006 MAYA SKELETAL REMAINS FROM HONDURAS 85

Appendix 3. Preservation and degree of occlusal wear for the permanent teeth of four individuals from the Copan and El Puente sites Copan 35-2000 (Female?) Copan 36-2000 (Male) El Puente 10-3 (Male) El Puente 204-1 (Male) Tooth Right Left Right Left Right Left Right Left Preservation1 UI100001111 UI200001111 UC00 00 11 11 UP100011111 UP200111111 UM100111111 UM200111111 UM300011100 LI100001111 LI200001111 LC11 00 11 11 LP110001010 LP200001111 LM101001111 LM201111111 LM300111100 Degree of occlusal wear2 UM1 — — 23 23 11 22 LM1 — 2 — — 1 2 2 2 1 1, Preserved; 0, Not preserved. 2 Broca’s score (Martin and Saller, 1957). 3 The score for either or both of the right and left teeth. 86 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE

Appendix 4. Measurements of the permanent teeth of four individuals from the Copan and El Puente sites (in mm)1 Copan 35-2000 (Female?) Copan 36-2000 (Male) El Puente 10-3 (Male) El Puente 204-1 (Male) Tooth Right Left Right Left Right Left Right Left Mesiodistal crown diameter UI1 — — — — 7.9 8.0 — — UI2 — — — — 7.3 7.1 — — UC—— —— 8.18.1—— UP1 — — — 7.8 7.1 6.9 — — UP2 — — 7.4 (7.0) 6.3 6.5 — — UM1 — — — (10.3) 10.7 10.5 — — UM2 — — 9.6 (10.2) 9.1 9.0 10.8 — UM3 — — — 7.4 9.0 9.1 — — LI1 — — — — 5.0 5.2 — — LI2 — — — — 6.0 6.0 — — LC — (7.0) — — 7.1 6.9 8.3 8.3 LP1 6.3 — — — 6.8 7.22 7.5 (7.4)2 LP2 — — — — 7.2 7.1 — 7.0 LM1 — (11.1) — — 11.2 11.1 — — LM2 — — — (10.9) 10.1 10.0 — — LM3 — — 10.0 9.9 9.8 10.2 — — Labio- or buccolingual crown diameter UI1 — — — — 6.8 7.2 — — UI2 — — — — 5.6 5.8 — — UC — — — — 8.6 8.7 10.1 9.8 UP1 — — — 9.9 9.6 9.4 10.0 10.1 UP2 — — 9.8 9.6 8.8 9.3 9.8 10.1 UM1 — — — 11.2 12.1 11.8 12.9 13.0 UM2 — — 11.7 14.3 11.4 11.2 12.6 12.9 UM3 — — — 11.0 11.0 10.8 — — LI1 — — — — 5.4 5.5 6.1 6.1 LI2 — — — — 5.7 5.6 6.2 6.4 LC 7.8 8.0 — — 7.7 7.3 8.4 8.5 LP1 7.0 — — — 7.0 7.02 8.1 7.82 LP2 — — — — 8.0 7.6 9.1 9.0 LM1 — 10.9 — — 10.5 10.4 11.9 12.0 LM2 — 9.7 — 10.0 9.4 9.8 11.5 11.1 LM3 — — 9.8 9.9 9.6 9.7 — — Mesiodistal cervical diameter UI1 — — — — 5.4 5.6 7.7 7.4 UI2 — — — — 5.0 5.1 5.8 5.9 UC — — — — 6.1 6.1 6.9 6.4 UP1 — — — — 5.1 5.1 5.3 5.5 UP2 — — 5.5 5.2 4.7 4.6 5.5 5.2 UM1 — — 9.0 — 7.2 7.6 8.3 8.2 UM2 — — 8.1 — 7.2 7.0 7.9 7.9 UM3 — — — 6.1 6.8 6.9 — — LI1 — — — — 3.4 3.6 3.8 3.7 LI2 — — — — 4.0 3.8 4.5 4.2 LC 5.5 5.5 — — 5.6 5.5 6.2 6.2 LP1 4.4 — — — 4.9 4.82 4.8 4.92 LP2 — — — — 4.6 4.6 5.1 5.2 LM1 — (9.2) — — 9.3 9.2 9.6 9.6 LM2 — (8.1) 9.3 9.7 8.5 8.8 — 8.9 LM3 — — — 8.6 8.4 8.5 — — 1 The values in parentheses may not be accurate. 2 Measured before the tooth was extracted for DNA analysis. Vol. 114, 2006 MAYA SKELETAL REMAINS FROM HONDURAS 87

Appendix 5. Non-metric crown characters of the permanent teeth of four individuals from the Copan and El Puente sites1 Copan 35-2000 (Female?) Copan 36-2000 (Male) El Puente 10-3 (Male) El Puente 204-1 (Male) Character Right Left Right Left Right Left Right Left Shoveling UI1—————2 —— UI2 — — — — 1 1 — — Carabelli trait UM1——1?—22 11 UM2 — — — — 0 0 — — UM3 — — — — 0 0 — — Hypocone reduction UM1——000000 UM2 — — — — 0 0 — 0 UM3 — — — 32 22—— Protostylid LM1———————— LM2 — 1 — — 1 1 — — LM3———————— Sixth cusp LM1———————— LM2———————— LM3 — — 0 — — 0 — — Seventh cusp LM1 — — — — 0 0 — — LM2 — — — — 0 0 — — LM3 — — — — 0 0 — — Hypoconulid reduction LM1 — — — — 0 0 — — LM2———————— LM3 — — 3 3 3 3 — — Groove pattern LM1 — — — — 2 1 — — LM2 — 2 3 3 2 2 — — LM3——3—33—— 1 Expressivity, i.e. degree of development of a tooth crown character is shown here. All characters except groove pattern of the mandibular molar are classified into four expressivities or grades: 0, 1, 2, and 3. In the case of groove pattern, the number of expressivities is three: 1 (Y pattern), 2 (+ pattern), and 3 (X pattern). For detailed criteria of classification, see Mizoguchi (1977). 2 A bicuspid tooth. 88 Y. MIZOGUCHI AND S. NAKAMURA ANTHROPOLOGICAL SCIENCE

Appendix 6. Postcranial measurements of two adult individuals Appendix 7. Observations on the postcranial bones of from the El Puente site (in mm)1 two adult individuals from the El Puente site El Puente 10-3 El Puente 204-1 El Puente 10-3 El Puente 204-1 (Male) (Male) (Male) (Male) Martin’s No.2 Right Left Right Left Character Right Left Right Left HUMERUS HUMERUS 1 Maximum length 301 302 — 298 Deltoid tuberosity weak weak weak weak 5 Maximum mid-shaft 20 20 22 21 FEMUR diameter Pilaster absent absent moderate moderate 6 Minimum mid-shaft 15 14 16 16 Allen’s cervical fossa1 absent absent — — diameter Extended articular absent absent present present 9 Transverse diameter 39 38 — (38) surface of head2 of head Charles’ facet1 present present present — 10 Vertical diameter of 42 43 — 42 TIBIA head Inferior fossa absent absent absent absent RADIUS (squatting facet)1 1 Maximum length 234 235 223 226 Inferior facet absent absent present present 4 Transverse shaft 15 15 14 15 (squatting facet)1,3 diameter TALUS 5 Sagittal shaft diameter 11 10 12 11 Abnormal surface absent absent absent absent ULNA of neck1 1 Maximum length 247 250 — — Lateral talar facet4 absent absent present present 11 Dorsovolar shaft 12 13 12 13 diameter 1 For the definition, see Baba (1970). 12 Transverse shaft 14 14 16 16 2 This may be regarded as a variant of Allen’s cervical fossa (Baba, diameter 1970). 13 Upper transverse 18 18 18 18 3 Or called a lateral tibial facet (Aiello and Dean, 1990). shaft diameter 4 For the definition, see Aiello and Dean (1990). 14 Upper dorsovolar 22 20 23 24 shaft diameter FEMUR 1 Maximum length 421 420 (406) — Oblique length 416 417 — — Appendix 8. Stature estimates for two adult individuals 2 1 6 Sagittal mid-shaft 24 25 25 25 from the El Puente site diameter 7 Transverse mid-shaft 25 25 28 28 Long bone Stature diameter (in cm) (in cm) 9 Subtrochanteric 30 29 31 31 El Puente 10-3 (male) transverse diameter 2 Maximum length of right femur 42.1 159.0 ± 3.4 10 Subtrochanteric sag- 21 22 23 23 2 ittal diameter Maximum length of left femur 42.0 158.8 ± 3.4 Lateral condyle–malleolus 34.3 158.5 ± 2.8 11 Supracondylar sagit- 26 25 29 — 3 tal diameter length of right tibia Lateral condyle–malleolus 34.4 158.7 ± 2.8 12 Supracondylar trans- (41) 33 37 — 3 verse diameter length of left tibia 15 Vertical diameter of 29 28 30 31 Average 158.75 neck El Puente 204-1 (male) 16 Sagittal diameter of 22 21 25 25 Lateral condyle–malleolus 35.0 159.9 ± 2.8 neck length of right tibia3 18 Vertical diameter of 42 42 44 43 Lateral condyle–malleolus 35.0 159.9 ± 2.8 head length of left tibia3 19 Transverse diameter 42 42 (43) — Average 159.90 of neck 1 21 Epicondylar breadth 75 75 — — Estimated using the formulae for Native Mexican male stature 22 Length of lateral 61 62 — — while alive, originally developed by Genoves (1967) and corrected condyle according to comments by Bass (1971). 2 Martin’s No. 1 for the femur (Martin and Saller, 1957). TIBIA 3 1 Lateral condyle–mal- 343 344 350 350 Martin’s No. 1 for the tibia (Martin and Saller, 1957). leolus length 1a Spinomalleolar (349) 349 (355) 358 length 8 Maximum mid-shaft 29 28 33 32 diameter 8a Sagittal diameter at 29 30 37 — nutrient foramen 9 Transverse mid-shaft 19 19 22 21 diameter 9a Transverse diameter 21 20 23 — at nutrient foramen 1 The values in parentheses may not be accurate. 2 Martin and Saller (1957).