The morphometric relationship of Upper Cave 101 and 103 to modern Homo sapiens

Journal of Human Evolution 45 (2003) 1–18

The morphometric relationship of Upper Cave 101 and 103 to modern Homo sapiens Deborah L. Cunningham *, Richard L. Jantz Department of Anthropology, University of Tennessee, 252 South Stadium Hall, Knoxville, TN 37996, USA Received 8 February 2001; accepted 15 April 2003

Abstract Since the discovery of the Homo sapiens crania from the Upper Cave of Zhoukoudian in northern China (UC 101, UC 102, and UC 103), no clear consensus has arisen regarding their affinities with modern populations. We use linear craniofacial measurements to compare UC 101 and UC 103 to a worldwide sample of H. sapiens that includes Paleoamericans and Archaic Indians, and employ Mahalanobis distance analysis and associated unweighted, unrestricted canonical variate analysis for the comparisons. Analyses indicate that UC 101 has consistent affinities with Easter Island and European populations, whereas UC 103 has more tenuous similarities with Australo-Melanesian groups. Both fossils exhibit some similarities to certain Paleoamerican and Archaic Indian individuals, but rarely cluster together. Upper Cave 103 is more of an outlier to modern populations than is UC 101. The fossils are not representative of any group to which they have been compared, but may be part of the generalized population that was ancestral to Paleoamericans.  2003 Elsevier Ltd. All rights reserved. Keywords: Upper Cave; Zhoukoudian; Mahalanobis distance analysis; Canonical variate analysis; Paleoamericans; Archaic Indians

Introduction Since the discovery of the anatomically modern human skeletal remains from the Upper Cave, or Shandingdong, of Zhoukoudian in northern China, various researchers have found the fossils to be similar to widely disparate worldwide groups * Corresponding author. Department of Anthropology, University of Missouri-Columbia, 107 Swallow Hall, Columbia, MO 65211, USA. Tel.: +1-573-882-5407 E-mail addresses: [email protected] (D.L. Cunningham), [email protected] (R.L. Jantz).

(e.g., Weidenreich, 1938/39; Neumann, 1956; Wu, 1956; Stewart, 1960; Wu, 1961, 1992, 1994; Coon, 1962; Howells, 1983, 1989, 1995; Wolpoff et al., 1984; Habgood, 1985; Turner, 1985; Wu and Zhang, 1985; Kamminga and Wright, 1988; van Vark and Dijkema, 1988; Brace and Tracer, 1992; Kamminga, 1992; Thorne and Wolpoff, 1992; Wright, 1992, 1995; Wolpoff, 1995, 1996; Wu and Poirier, 1995; Cornell and Jantz, 1997, 1998; Brown, 1998, 1999; Cornell, 1998; Neves and Pucciarelli, 1998; Turner et al., 2000; Cunningham and Wescott, 2002). Howells (1995:43) calls the

0047-2484/03/$ - see front matter  2003 Elsevier Ltd. All rights reserved. doi:10.1016/S0047-2484(03)00064-2

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“Old Man” of Upper Cave (101) “everyone’s problem skull” because UC 101 is considerably larger and more robust than present day East Asians, and because there is disagreement as to whether or not this skull is morphologically Asian. Metrists tend to classify UC 101 as everything but Asian (Howells, 1983, 1989, 1995; Kamminga and Wright, 1988; van Vark and Dijkema, 1988; Kamminga, 1992; Wright, 1992, 1995; Cornell and Jantz, 1997, 1998; Brown, 1998, 1999; Cornell, 1998; Neves and Pucciarelli, 1998), whereas morphologists are inclined to categorize the skull as Asian (Coon, 1962; Weidenreich, 1938/39; Wu, 1956; Wu, 1961, 1994; Turner, 1985; Wu and Zhang, 1985; Wolpoff, 1995, 1996; Wu and Poirier, 1995; Turner et al., 2000). The remains of at least seven additional individuals were found with UC 101 (Wu, 1961), including two other complete crania (UC 102 and UC 103). These latter two specimens have been studied less extensively than the Old Man because of cranial deformation and fragmentation. Upper Cave 103 exhibits some postmortem protrusion of the vault along the coronal suture that is not severe enough to significantly affect cranial measurements. However, the cranial vault is significantly distorted in UC 102. For these reasons, we include only UC 101 and UC 103 in our analyses. Although Pei discovered the Upper Cave specimens in 1933 (Pei, 1939), it is Franz Weidenreich who described and interpreted the fossils (Weidenreich, 1938/39, 1939; Wolpoff, 1996). In his analysis of the three crania, Weidenreich (1938/1939:170-171) describes UC 101 as “primitive Mongoloid”, UC 102 as “Melanesoid”, and UC 103 as “Eskimoid” racial types. Based on these results, Weidenrech (1938/ 1939:170-171) suggests that, although one might expect older populations to be morphologically uniform, “man was split into different racial stems already in such an early stage of evolution as is represented by Sinanthropus and Pithecanthropus”, thereby contradicting the existence of “purer” races in the past. More recent metrical statistical studies of UC 101 have not classified this fossil as Asian. Although the group with which UC 101 is most closely affiliated varies greatly among studies, UC

101 is most often associated with Pacific and American populations (Table 1). Wolpoff (1995) is critical of multivariate analysis, arguing that it is inappropriate to perform a discriminant function analysis on a specimen from a population not represented in the data upon which the function is based. Wolpoff (1995:186) believes that “a multivariate analysis of measurements is not an anatomical analysis” and that multivariate clustering of principal components does not mean morphological similarity. He uses non-metric morphological analysis to suggest that UC 101 is “Mongoloid” but does not resemble living Asian groups because the specimen exhibits many “archaic characteristics” (Wolpoff, 1996: 723). He proposes that UC 101 belongs to a group from which some later Asians, including Eskimos and Amerindians, evolved. Other authors, using morphological criteria, concur with an Asian affinity for UC 101 (Table 2). In order to contribute to the discussion about the populational affinities of the Upper Cave fossils, we compare UC 101 and UC 103 to modern groups using cranial morphometrics. This study differs from previous studies in two ways: (1) we use a methodology not previously applied to these fossils, Mahalanobis distance analysis associated with complementary unweighted, unrestricted canonical variate analysis, and (2) we include measurements of Paleoamerican and Archaic Indian crania. Additionally, we have included UC 103 in the analyses, whereas most previous studies have focused only on UC 101 (but see Wu and Zhang, 1985; Habgood, 1985; Neves and Pucciarelli, 1991; Wright, 1992; Cornell, 1998; Cunningham and Wescott, 2002).

Materials Dating the Upper Cave fossils has been problematic (Kamminga and Wright, 1988; Cornell, 1998), and has been summarized recently in Cunningham and Wescott (2002). Three different dates have been proposed, without consensus: (1) 10,000 BP (An, 1991), (2) 18,000 BP (An, 1991), and (3) 29–24,000 BP (Hedges et al., 1992).

Table 1 Metric analyses of Upper Cave 107

Howells (1983) Habgood (1985)* Kamminga and Wright (1988) Kamminga and Wright (1988)

Kamminga and Wright (1988) van Vark and Dijkema (1988) Howells (1989) Howells (1989) Neves and Pucciarelli (1991)* Brace and Tracer (1992) Wright (1992)** Howells (1995) Wright (1995) Cornell and Jantz (1997) Brown (1998; 1999) Cornell (1998)** Cornell and Jantz (1998) Neves and Pucciarelli (1998) Cunningham and Wescott (2002)*

Type of Analysis Discriminant Function PCA & Cluster Analysis

Resemblance

Plains Indians & Northern Europeans Australian (UC 101), Southwest Asian & Europe (UC 102), & East/SE Asian & Europe (UC 103) PCA Ainu, African & European populations KMEANS African, Australo-Melanesian & Easter Island populations when the data are divided into 2 groups; Caucasoid & Ainu when the data are divided into 3 groups Penrose Shape Sub-Saharan African, Australo-Melanesian & Easter Island populations Mahalanobis Distance South Australia, Norse, Ainu, Arikara, Santa Cruz & Easter Island populations PCA Elmenteita (Kenya, ca. 7,000 B.P.) C-scores Polynesian populations PCA Early South Americans, early & modern Australians (UC 102 & 103 only) Discriminant Function on C-scores N-E Amerindian, Pacific, Ainu, Europe, & Eskimo populations CRANID African, Australian & Melanesian populations DISPOP Arikara Giles-Elliot (1962) Discriminant Function African-American CVA & Mahalanobis Distance Polynesian, European & Amerindian populations Discriminant Function Eskimo Unweighted CVA & Mahalanobis Distance Polynesian, European & Amerindian populations (UC 101), Australo-Melanesian populations (UC 103) Unweighted CVA & Mahalanobis Distance Polynesian, European & Amerindian populations PCA Paleoamericans, South Pacific & African populations Mahalanobis Distance Easter Island (UC 101), Egypt (UC 102 - male), Blackfoot (UC 102 - female), Tasmania (UC 103)

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Author(s) and Date

*Also included UC 102 & 103. ** Also included UC 103.

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Table 2 Morphological analyses of Upper Cave crania Author(s) and Date Weidenreich (1938/1939:170) Wu (1956) Neumann (1956) Wu (1961; 1994) Coon (1962) Turner (1985) Wu and Zhang (1985) Wu and Poirier (1995) Wolpoff (1996:723) Turner et al. (2000)

Resemblance “primitive Mongoloid” Mongolian American Indian Mongoloid Mongoloid Sinodont Modern and fossil Chinese East Asian Mongoloid with “archaic characteristics” Sinodont

We compare UC 101 and UC 103 to a worldwide sample of H. sapiens crania from Howells’ database (1973; 1989), additional Plains Indians groups (Blackfoot, Cheyenne, and Pawnee), Paleoamericans, Archaic Indians, and the Liujiang specimen (Table 3). Paleoamericans are those ancient Americans older than 8,000 B.P. (Bamforth, 1988; Fagan, 1995; Table 4), and the Archaic period sequentially follows the Paleoamerican period (Table 5). While Pelican Rapids could be considered either late Paleoamerican or early Archaic due to its borderline date, we consider it an early Archaic specimen. The H. sapiens specimen from Liujiang, China is considered morphologically “Mongoloid” (Brown, 1992; Pope, 1992; Wu, 1994), and its age has been reexamined recently by Shen et al. (2002). These authors conclude that this fossil likely dates to w111– 139 ka, and is definitely not more recent than w68 ka (Shen et al., 2002). Liujiang is included only in Analysis 2 due to missing measurements (see below). Since UC 101 is a likely male and UC 103 a likely female (Weidenreich, 1938/1939; Wu, 1956; Coon, 1962; Wu and Zhang, 1985; Kamminga and Wright, 1988; Kamminga, 1992), we include both sexes in the reference samples.

Methods Mahalanobis distance analysis (D2) We generated Mahalanobis squared distances between each Upper Cave cranium and reference

group in order to estimate each fossil’s closest group affiliation. Mahalanobis distance analysis evaluates the squared distances of an unknown specimen from each known group’s centroid, and classifies the unknown specimen based on the shortest distance. This type of analysis is complementary to canonical variate analysis (CVA) (Albrecht, 1992) because the CV plots will show, in two dimensions, the graphical representation of the Mahalanobis distances. Mahalanobis distances were not calculated between the Upper Cave fossils and Paleoamerican or Archaic Indian samples, but these relationships are seen in the canonical plots. Canonical variate analysis In order to graphically represent the relationship of UC 101 and UC 103 to Paleoamericans, Archaic Indians, and modern groups, we performed four unweighted, unrestricted canonical variate analyses. Canonical variate analysis (CVA), first developed by Rao (1952), is a useful tool for describing differences among groups. The most traditional form of CVA is the weighted analysis in which each group’s influence on the among-groups covariance matrix is affected by its sample size as well as its mean values. However, when there is no justification in assuming that the groups differ in significance or a priori probabilities, it is inappropriate to define the amonggroup covariance matrix in terms of group sample sizes. Doing so results in the structure of the canonical variate axes being determined by the well sampled groups, while the inadequately sampled groups’ variation would contribute only slightly (Albrecht, 1992). We consider an “unweighted” analysis the most appropriate approach for including single crania because a group’s influence is contingent only on its mean values and not its sample size. Thus, all groups are equally important in the construction of the canonical variate axes (Albrecht, 1980, 1992). This allows the Zhoukoudian fossils to contribute to the multivariate space in which they are analyzed, and addresses Wolpoff’s (1995) criticism of traditional multivariate studies. Likewise,

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Table 3 Sample size Population Ainu Andaman Anyang Archaic Indian Arikara Atayal Australia Berg Blackfoot Buriat Bushman Cheyenne Dogon Easter Island Egypt Eskimo Guam Hainan Liujiang Mokapu Moriori Norse North Japan Paleoamerican Pawnee Peru Philippine Santa Cruz South Japan Tasmania Teita Tolai Zalava´r Zulu

Group

M

F

Total

Reference

Ainu Andaman Island Continental East Asia Archaic Indian Americas Pacific Rim Australo-Melanesia Europe Americas Continental East Asia Sub-Saharan Africa Americas Sub-Saharan Africa Polynesia Egypt Eskimo Micronesia Pacific Rim Liujiang Polynesia Polynesia Europe Pacific Rim Paleoamerican Americas Americas Pacific Rim Americas Pacific Rim Australo-Melanesia Sub-Saharan Africa Australo-Melanesia Europe Sub-Saharan Africa

48 35 42 8 42 29 52 56 23 55 41 16 47 49 58 53 30 45 1 51 57 55 55 5 17 55 50 51 50 45 33 55 53 55

38 35 0 7 27 18 49 53 43 54 49 6 52 37 53 54 27 38 0 49 51 55 32 1 10 55 0 51 41 42 50 55 45 46

86 70 42 15 69 47 101 109 66 109 90 22 99 86 111 107 57 83 1 100 108 110 87 6 27 110 50 102 91 87 83 110 98 101

Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Key (1983), R.L. Jantz (unpub.) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) R.L. Jantz (unpub.) Howells (1973; 1989) Howells (1973; 1989) R.L. Jantz (unpub.) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) W.W. Howells (unpub.) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Key (1983), R.L. Jantz (unpub.) Key (1983), R.L. Jantz (unpub.) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989) Howells (1973; 1989)

1417

1223

2640

Totals

an “unrestricted” CVA is appropriate here since such an analysis does not assume that the unknown specimen must belong to one of the modern reference groups under consideration (Albrecht, 1992). Typicality probabilities The unrestricted approach makes use of “typicality probabilities” which evaluate whether the unknown individual is a typical member or an outlier of the population it has been classified

into by determining whether it falls within the multivariate normal distribution of that group (see Campbell, 1984 for a more detailed description). Low probabilities indicate that the specimen is unlikely to belong to the reference group. Unlike posterior probabilities inherent in the restricted approach, typicality probabilities do not have to sum to one (1.0) (Albrecht, 1992). Here, since the Mahalanobis D-squared figures follow a Chi-square distribution, typicality probabilities were drawn from a Chi-square table based on the Mahalanobis D-squared figure rendered from the

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Table 4 Paleoamericans Site 25CM2 25CM2 25FT– NSM* NSM* 21TR05

Specimen

Name

Sex

Analysis

Site Date

Reference

UN30631 UN30632 UN31444 2023 2064 Browns Valley

Wet Gravel Pit Wet Gravel Pit Lime Creek Wizards Beach Spirit Cave Brown’s Valley

F M M M M M

1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 3,4 3,4

11550 B.P. 11550 B.P. 11524 B.P. 9250 B.P. 9415 B.P. 9049 B.P.

Key (1983) Key (1983) Key (1983) Tuohy and Dansie (1997) Tuohy and Dansie (1997) Myster and O’Connell (1997)

*NSM = Nevada State Museum.

Table 5 Archaic Indians Site

Specimen

13MN2 #1 25DO9002 SI94-03 25MP2 B137 25MP2 B3138 25SF10 1B854 25SF10 852 32MO97 2–12B 32MO97 4–14D 32MO97 5–15E 32MO97 6–16F 32MO97 8–18H 871 NSM* UOFOREGON 11–110 210T3 MNWOMAN 14LV315 3103

Name

Analysis

Sex

Site date

Turin Dkd Dry Lake Dry Lake Gering Gering Bahm Bahm Bahm Bahm Bahm

1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 1,2,3,4 3,4 4

M M F F M F M F M F F M M F M

4720 B.P. 3770 B.P. 3250 B.P. 3250 B.P. 2000 B.P. 2000 B.P. 1900 B.P. 1900 B.P. 1900 B.P. 1900 B.P. 1900 B.P. 4480 B.P. 7000 B.P. 7840 B.P. 5579 B.P.

Prospect Location Pelican Rapids Lansing Man

Reference Fisher et al. (1985) Jantz (unpub.) Key (1983) Key (1983) Key (1983) Key (1983) Williams (1994) (as cited in Williams, Williams (1994) (as cited in Williams, Williams (1994) (as cited in Williams, Williams (1994) (as cited in Williams, Williams (1994) (as cited in Williams, Jantz (unpub.) Cressman (1940) Myster and O’Connell (1997) Bass (1973)

1997) 1997) 1997) 1997) 1997)

*NSM=Nevada State Museum.

unweighted analysis with degrees of freedom equal to the number of variables employed. Since most statistical packages do not allow for an unweighted CVA, we followed a two-step process recommended by Albrecht (1992). First, we obtained a covariance matrix using the weighted CV scores produced by SAS plus the fossil CV scores from the raw canonical coefficients. Second, we extracted the eigenvalues and eigenvectors from the covariance matrix and the principal component scores for each group and each fossil. Adjusting for size Since UC 101 is a male and UC 103 is a female, we included both sexes from the reference samples in the analyses. Assuming that size is the main sex

difference, it was therefore necessary to perform a size adjustment on the cranial measurements in order to distinguish principally shape differences. Although numerous methodologies for size adjustment are possible, Jungers et al. (1995) found that Mosimann’s ratio methods provide results superior to other methods (e.g., residual adjustments, c-scores, Burnaby’s method). For that reason, we adjusted for isometric size following the methodology of Darroch and Mosimann (1985), and pooled sexes without centering. The Darroch and Mosimann (1985) method yields dimensionless shape variables by dividing each variable by the geometric mean of all variables for that case. If such a size adjustment is not made, the sexes cannot be pooled without differences in size

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Table 6 Measurements Abbreviation 1,2

ASB AUB1,2,3,4 AVR1,3 BBH1,2 BNL1,2 BPL1,2 DKB1,2,3 DKR1 DKS1,2 EKB1,2,3 EKR1,3 FMB1,2,3 FMR1,3,4 FOL1,2 FRC1,2,3,4 FRF1,2,3,4 FRS1,2,3,4 GOL1,2,3,4 IML1,2 JUB1,2,3 MAB1,2,3 MDB1,2,3 MDH1,2,3,4 MLS1,2 NAR1,3,4 NAS1,2,3 NDS1,2 NLB1,2,3

Full name Biasterionic breadth Biauricular breadth M1 alveolus radius Basion-bregma height Basion-nasion length Basion-prosthion length Interorbital breadth Dacryon radius Dacryon subtense Biorbital breadth Ectoconchion radius Bifrontal breadth Frontomalare radius Foramen magnum length Frontal chord Nasion-subtense fraction Nasion-bregma subtense Glabello-occipital length Malar length, inferior Bijugal breadth Palate breadth Mastoid width Mastoid height Malar subtense Nasion radius Nasio-frontal subtense Naso-dacryal subtense Nasal breadth

accounting for most of the variation along the axes. Neves and Pucciarelli (1998) found that, when size was not corrected in a principal components analysis, UC 101 was always an outlier due to its large size.

Abbreviation 1,2,3

NLH NOL1,2,3,4 NPH1,2,3 OBB1,2,3 OBH1,2,3 OCC1,2 OCF1,2 OCS1,2 PAC1,2,3,4 PAF1,2,3,4 PAS1,2,3,4 PRR1,3 SIS1,2 SSR1,3 SSS1,2 STB1,2 VRR1,3,4 WCB1,2 WMH1,2,3 WNB1,2 XCB1,2,3,4 XFB1,2,3,4 XML1,2 ZMB1,2 ZMR1,3 ZOR1,3 ZYB1,2

Full name Nasal height Nasio-occipital length Nasion-prosthion height Orbit breadth Orbit height Lambda-opisthion chord Lambda-opisthion fraction Lambda-opisthion subtense Bregma-lambda chord Bregma-subtense fraction Bregma-lambda subtense Prosthion radius Simotic subtense Subspinale radius Zygomaxillary subtense Bistephanic breadth Vertex radius Minimum cranial breadth Cheek height Simotic chord Maximum cranial breadth Maximum frontal breadth Malar length, maximum Bimaxillary breadth Zygomaxillare radius Zygoorbitale radius Bizygomatic breadth

Paleoamericans (Browns Valley and Spirit Cave) and one additional Archaic Indian (Pelican Rapids), Analysis 3 reduced the measurement set to 34 (Tables 3–6; N = 2,638). Analysis 4 included only the neurocranium in order to include the Archaic “Lansing Man” (Tables 3–6; N = 2,639).

Analyses Distance analysis We performed four analyses including different variables and specimens. Analysis 1 was based on fifty-five of Howells’ (1973; 1989) cranial measurements, all of Howells’ (1973; 1989) modern crania, and the most complete Paleoamerican and Archaic Indian samples available (Tables 3–5; N = 2,635). Measurements are listed in Table 6. In order to include the Liujiang specimen, Analysis 2 excluded the ten cranial radius measurements, but otherwise was identical to Analysis 1 (Tables 3–6; N = 2,636). In order to include two additional

We conducted a distance analysis of the fossils using the methodology of Defrise-Gussenhoven (1967) that calculates an expected distance between any two skulls drawn at random from a single population. We used the modern reference samples to determine an expected distance between two skulls from the same population. This distance will be distributed as (2pⳮ1)1/2

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with a variance of 1, where p is the number of dimensions used. For a more detailed discussion of this methodology, see Jantz and Owsley (2001) and Cunningham and Wescott (2002).

Results Analysis 1 In Analysis 1, UC 101’s closest affiliation is with Easter Island, with a reasonable typicality probability of 0.134 (Table 7). All European populations, Arikara, and Tasmania are close behind, however the associated typicality probabilities are much lower. The next closest populations increase their D2 score and decrease their typicality probabilities more extremely. Upper Cave 103’s closest neighbors in multivariate space are the Australo-Melanesian populations, but these squared distances are almost twice those of the distance of UC 101 to its closest neighbor. Additionally, all associated typicality probabilities are less than 0.001, indicating that UC 103 is not a typical member of any of the Australo-Melanesian groups. Figs. 1–4 graphically illustrate the Upper Cave fossils’ proximity to the modern groups as well as to the Paleoamerican and Archaic samples, but because these ancient Americans were not included in the Mahalanobis distance analyses (Table 7), the CVA results below focus on the fossils’ proximity to these groups. In CVA 1, canonical axis 1 (CAN 1) explains 12.1% of the total variation and CAN 2 explains 11.2%, for a total of 23.3% of total variation1. Individuals with high scores on CAN 1 are characterized by a low interorbital prominence, 1

Although it is desirable for the first few canonical axes to explain large percentages of the among-group variation, because the number of samples (including individual specimens of Archaic and Paleoamericans) is large, the amount of variation explained by each axis is inevitably reduced. Nonetheless, each axis still provides a good description of general patterns of craniofacial variation. The geographical groupings are accurate and the most ancient Americans (especially the Archaic Indians) are clearly differentiated from the modern reference populations. Therefore, we feel confident that the major differences in populations and individual specimens are shown in the CV plots, despite the fact that these plots explain between 23% and 32% of total variation.

high vaults, long and wide frontals, curved malars, narrow vault base, wide and short nasal apertures, long and wide mastoids, curved and long parietals, and subnasal orthognathism. Individuals with high scores on CAN 2 typically have wide upper faces, short malars, flat interorbitals, wide mastoids, narrow frontals, subnasal orthognathism, and flat parietals. Fig. 1 indicates that, in addition to the modern populations (Table 7), UC 101 is close to several Archaic Indians on CAN 1: specimen 871, Dry Lake (B3138), and Gering (852). It is also nearby the Lime Creek Paleoamerican. On CAN 2, UC 101 is close to one of the Bahm burials (2-12B), specimen 871, and Turin. On this axis, it is also close to the Wizards Beach Paleoamerican. On CAN 1, UC 103 is near several Archaic Indian individuals (Turin, Dry Lake [B3138], Prospect, and Gering [852]), as well as the Lime Creek Paleoamerican. Upper Cave 103 and UC 101 are not far apart on this axis. As is expected, based on the Mahalanobis distance analysis (Table 7), UC 103 lies farther away from most populations on CAN 2. Its closest neighbors are AustraloMelanesian populations and three Archaic Indians (Bahm Burials 5-15E and 8-18H, and Dkd). Analysis 2 As in Analysis 1, UC 101 is closest to European groups and Easter Island, with the addition of Tasmania, Arikara, and Blackfoot (Table 7). Its highest typicality probabilities are associated with Norse and Easter Island. Upper Cave 103 again shows its closest relationship with AustraloMelanesians, but, just as in Analysis 1, its squared Mahalanobis distances are twice as large as those associated with UC 101, and the associated typicality probabilities are less than 0.001. In CVA 2, CAN 1 explains 12.9% of the variation and CAN 2 explains 11.6%, for a total of 24.5%. High scores on CAN 1 mean individuals have flat nasals, long and curved frontals, narrow lower vaults, high and wide mastoids, curved malars, low interorbital prominence, short and wide nasal apertures, wide upper vaults, subnasal flatness, long parietals, and high vaults. Individuals with high scores on CAN 2 are characterized by narrow upper faces, long malars, narrow vaults,

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protruding nasal bones, long and curved parietals, and wide frontals. In Fig. 2, in addition to the modern groups, UC 101 clusters most closely with the Archaic Indians Turin, specimen 871, Prospect, both Gering crania, and Dry Lake (B3138), as well as the Lime Creek Paleoamerican on CAN 1. On CAN 2 it is closest to Turin, two Bahm burials (8-18H and 2-12B), specimen 871, and the Liujiang cranium. On CAN 1, UC 103 is closest to two Archaics (Dry Lake [B137] and Bahm [6-16F]), and the Wet Gravel Male Paleoamerican. It is not close to any population on CAN 2, but its nearest neighbors are the Archaics Dkd, one Bahm specimen (5-15E), and the Australo-Melanesian populations. Neither fossil is especially close to the Liujiang specimen (cf., Hanihara, 1994; Brown, 1998, 1999), except UC 101 on CAN 2. Analysis 3 Once again, UC 101 is closest to European and Easter Island populations, with the Ainu in close proximity (Table 7). The results for UC 103 continue the pattern of Australo-Melanesian populations (with the addition of Eskimo) as its nearest neighbors in multivariate space, this time with much smaller squared distances. However, only with the Tolai is a typicality probability greater than 0.001 present. In CVA 3, CAN 1 explains 12.5% of the variation and CAN 2 11.6% for a total of 24.1%. Individuals with high scores on CAN 1 have narrow lower vaults, flat and long parietals, flat nasions, general facial flatness, low cheek height, short upper faces, subnasal orthognathism, short nasal apertures, long mastoids, and long and wide frontals. High scores on CAN 2 are indicative of curved frontals, wide nasal apertures, retreating lateral orbital margins, narrow faces, flat nasions, narrow mastoids, and narrow vault base. Fig. 3 shows that on CAN 1, UC 101 clusters closely with the Archaic specimens Gering (1B854), Turin, and Prospect, and the Paleoamericans Wet Gravel Female and Browns Valley. On CAN 2, the fossil is closest to three Paleoamericans (Wet Gravel Female and Male, and Spirit Cave). On CAN 1, UC 103 plots closest to

9

the Archaic specimens Dry Lake (B3138), 871, and Prospect, while on CAN 2, it is closest to the Archaics Prospect and Dkd. Analysis 4 Analysis 4 emphasizes vault measures and is the only analysis in which UC 101 is most closely affiliated with Australo-Melanesian, modern Amerindian (Blackfoot, Santa Cruz, and Pawnee primarily), and Eskimo populations (Table 7). The highest typicality probabilities are associated with Australia, Tasmania, and Blackfoot. Upper Cave 103 is most closely associated with Tolai and Eskimo with extremely small typicality probabilites. In CVA 4, CAN 1 explains 17.5% of the variation and CAN 2 14.5% for a total of 32%. Individuals with high scores on CAN 1 are distinguished by projecting nasions, wide lower vaults, facial forwardness, long sagittal lengths, short mastoids, and flat frontals. High scores on CAN 2 indicate that individuals have flat and long frontals, high vaults, wide lower vaults, and short sagittal length. As can be seen in Fig. 4, on CAN 1, UC 101 is closest to three Paleoamericans (Browns Valley, Wet Gravel Male, and Lime Creek) and two Archaic Bahm burials (8-18H and 6-16F). On CAN 2, it is closest to the Archaic specimen from Turin, both Gering individuals, Prospect, Lansing Man, and Pelican Rapids, as well as the Paleoamerican Spirit Cave. On CAN 1, UC 103 is not close to any population. Its nearest neighbors are three Paleoamericans (Browns Valley, Wet Gravel Male, and Lime Creek) and one Archaic Bahm specimen (4-14D). On CAN 2, it is closest to two Bahm burials (8-18H and 6-16F), and the Lime Creek Paleoamerican. Distance between UC 101 and UC 103 Based on the modern reference samples included in this study, the expected distance between two skulls drawn at random from the same modern population is 10.392, with a variance and standard deviation of one (DefriseGussenhoven, 1967). The distance between UC

10

Table 7 Mahalanobis distances and typicality probabilities, listed in order of D2 UC 101, Analysis 1

UC 103, Analysis 1 2

UC 103, Analysis 2 2

Typicality probability

D

Population

Typicality probability

D

Population

Typicality probability

D

Population

Typicality probability

66.7 71.2 71.8 72.6 73.2 73.6 77.9 79.1 79.8 81.7 83.7 84.2 85.2 86.9 87.5 88.1 88.1 88.4 89.5 89.9 90.6 91.2 91.6 91.9 92.4 95.3 97.6 98.8 113.3 114.8 117.2

Easter Island Norse Berg Talavar Arikara Tasmania Blackfoot Moriori Tolai Ainu Santa Cruz Atayal Phillipines Pawnee Guam Teita Anyang Mokapu N. Japan Buriat Egypt Zulu Australia S. Japan Hainan Peru Eskimo Bushman Andaman Dogon Cheyenne

0.134 0.069 0.064 0.056 0.051 0.048 0.023 0.018 0.016 0.011 0.008 0.007 0.006 0.004 0.003 0.003 0.003 0.003 0.002 0.002 0.002 0.002 0.001 0.001 0.001 0.001 <0.001 <0.001 <0.001 <0.001 <0.001

121.6 125.4 132.5 135.0 135.8 138.5 139.9 142.3 143.6 144.9 145.3 148.2 149.4 151.7 152.4 152.4 152.4 152.5 154.1 154.7 155.2 157.7 159.6 162.3 162.9 165.8 167.2 170.2 177.5 185.3 189.5

Tolai Tasmania Australia Ainu Zalavar Zulu Easter Island Norse Teita Mokapu Atayal Arikara Phillipines Egypt S. Japan Hainan Eskimo Anyang Berg Guam Moriori N. Japan Dogon Pawnee Andaman Bushman Peru Cheyenne Santa Cruz Blackfoot Buriat

<0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

57.4 58.5 60.6 61.1 64.5 65.0 65.8 70.3 70.9 71.5 73.1 73.2 74.4 74.6 75.1 75.7 76.8 78.6 79.1 79.3 80.1 82.7 82.9 82.9 83.3 84.6 90.5 90.6 95.2 99.2 102.0

Norse Easter Island Tasmania Zalavar Berg Arikara Blackfoot Tolai Ainu Moriori Santa Cruz Phillipines Guam Australia Atayal Egypt Pawnee Mokapu Teita Zulu Anyang Peru Buriat Hainan N. Japan S. Japan Eskimo Bushman Dogon Andaman Cheyenne

0.102 0.085 0.060 0.055 0.030 0.027 0.023 0.009 0.008 0.007 0.005 <0.005 0.004 0.004 0.003 0.003 0.002 0.001 0.001 0.001 0.001 0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

113.3 115.5 117.4 119.4 123.5 127.1 129.3 129.9 134.0 134.1 135.5 136.3 137.2 138.8 140.6 141.3 141.6 141.9 142.1 143.0 143.7 146.1 146.8 148.9 150.0 150.3 150.6 157.9 159.6 160.3 181.5

Tasmania Tolai Australia Ainu Zalavar Zulu Easter Island Norse Arikara Eskimo Atayal Teita Mokapu Egypt Moriori Guam Phillipines Anyang Hainan S. Japan Berg N. Japan Pawnee Dogon Peru Blackfoot Andaman Bushman Santa Cruz Cheyenne Buriat

<0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

Population

D

2

UC 101, Analysis 2

2

Table 7

(continued) UC 101, Analysis 3

UC 103, Analysis 3 2

UC 103, Analysis 4 2

Typicality probability

D

Population

Typicality probability

D

Population

Typicality probability

D

Population

Typicality probability

44.5 46.7 48.0 48.0 49.5 51.5 51.8 52.0 52.7 53.0 57.0 57.3 57.7 58.1 58.7 58.8 59.4 60.5 60.7 61.8 62.5 62.5 62.6 64.3 64.7 65.4 67.3 70.9 74.0 82.8 85.2

Zalavar Easter Island Berg Ainu Norse Pawnee Tasmania Moriori Arikara Blackfoot N. Japan Tolai Phillipines Australia Anyang Atayal Buriat Mokapu Santa Cruz Teita Guam Egypt S. Japan Hainan Eskimo Bushman Zulu Cheyenne Peru Dogon Andaman

0.107 0.073 0.056 0.056 0.042 0.027 0.026 0.025 0.021 0.020 0.008 0.008 0.007 0.006 0.005 0.005 0.004 0.003 0.003 0.002 0.002 0.002 0.002 0.001 0.001 0.001 0.001 <0.001 <0.001 <0.001 <0.001

56.4 70.8 70.9 72.0 72.8 73.1 73.8 74.7 76.1 77.4 77.8 78.2 78.6 78.8 79.4 79.8 81.4 81.9 82.7 84.2 88.6 88.9 90.4 90.6 90.8 91.2 91.4 92.5 94.5 96.2 107.3

Australia Australia Eskimo Tasmania Teita Cheyenne Atayal Zulu S. Japan Ainu Phillipines Hainan N. Japan Guam Arikara Zalavar Anyang Easter Island Mokapu Norse Bushman Blackfoot Pawnee Egypt Andaman Peru Berg Moriori Dogon Santa Cruz Buriat

0.522 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

27.8 16.3 18.5 21.5 22.8 22.9 23.6 24.8 25.1 25.2 25.5 25.7 26.6 27.1 28.5 28.6 31.5 32.2 32.3 32.7 34.0 34.2 34.5 34.7 34.9 35.7 36.0 38.5 38.9 46.8 48.1

Tolai Tasmania Blackfoot Tolai Santa Cruz Eskimo Pawnee Moriori Zalavar Ainu Arikara Easter Island Cheyenne Norse N. Japan Berg Guam Mokapu Phillipines Atayal Bushman Teita Buriat Anyang Peru S. Japan Zulu Hainan Egypt Andaman Dogon

0.023 0.363 0.235 0.122 0.088 0.087 0.072 0.052 0.048 0.047 0.044 0.041 0.032 0.028 0.019 0.018 0.007 0.006 0.006 0.005 0.003 0.003 0.003 0.003 0.003 0.002 0.002 0.001 0.001 <0.001 <0.001

27.8 28.8 35.8 36.0 37.5 38.6 39.5 39.8 40.3 41.3 41.5 42.2 42.2 42.9 43.4 43.9 45.1 45.4 46.8 47.4 50.6 50.8 51.7 51.8 52.6 53.6 53.7 53.9 54.0 55.4 57.0

Tolai Eskimo Teita Atayal Ainu Australia S. Japan Tasmania Arikara Easter Island Zulu Hainan N. Japan Zalavar Cheyenne Anyang Phillipines Norse Guam Blackfoot Pawnee Berg Moriori Peru Bushman Dogon Santa Cruz Mokapu Egypt Andaman Buriat

0.023 0.017 0.002 0.002 0.001 0.001 0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

Population

D

2

UC 101, Analysis 4

2

11

12

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

Fig. 1. CVA 1, 55 variables, N = 2,635. WGF = Wet Gravel Pit Female, WGM = Wet Gravel Pit Male, LC = Lime Creek, WB = Wizards Beach, DL = Dry Lake, PL = Prospect Location.

101 and UC 103 is 14.291 which is nearly 4 standard deviations greater than expected if one assumes that the Zhoukoudian Pleistocene population exhibits the same amount of internal variation as do modern populations.

Discussion The Upper Cave fossils and modern populations Upper Cave 101 has much larger typicality probabilities associated with its populational classification than does UC 103, and could fall within the range of variation of some modern groups. Upper Cave 101 is most consistently affiliated with Easter Island and European populations with the highest typicality probabilities, and also shows similarity to Tasmania and some modern

Amerindian groups, especially the Arikara and Blackfoot (Table 7). Upper Cave 103 is consistently classified as Australo-Melanesian, albeit with extremely low typicality probabilities. This fossil is outside the range of variation of these robust Australo-Melanesian groups which are its closest neighbors in multivariate space. In Analysis 4, consisting of fifteen vault measurements, both fossils are classified as Australo-Melanesian with relatively high typicality probabilities, suggesting that the Zhoukoudian fossils and the AustraloMelanesians share more similarities in the vault than in the face. This is likely a reflection of the general robusticity of the neurocranium of both the fossils and these modern groups. Lahr (1995) argues that due to geographic and consequent genetic isolation, AustraloMelanesians remain closest to the morphology of

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

13

Fig. 2. CVA 2, 45 variables, N = 2,636. WGF = Wet Gravel Pit Female, WGM = Wet Gravel Pit Male, LC = Lime Creek, WB = Wizards Beach, DL = Dry Lake, PL = Prospect Location.

early robust modern humans, thus explaining why multivariate analyses consistently show that Australo-Melanesians are the closest relatives of fossil groups (e.g., Habgood, 1985; Kamminga and Wright, 1988; van Vark and Dijkema, 1988; Wright, 1992; see Table 1). She suggests that Australian and Melanesian populations have had a relatively independent evolutionary history during the late Pleistocene and did not undergo further derivations and specializations like that of the modern Asians. Paleoamericans and Archaic Indians Our analyses clearly link UC 101 and UC 103 to Paleoamericans from North America, lending support to the suggestion that these individuals were members of an unspecialized, pre-“Mongoloid” group that was involved in the peopling of the Americas (Neumann, 1956; Stewart, 1960; Turner,

1985; Brace and Tracer, 1992; Wu, 1992, 1994; Wolpoff, 1996; Cornell and Jantz, 1997; Neves and Pucciarelli, 1998). Previously, Neves and Pucciarelli (1998) found affinity between the Upper Cave fossils and Paleoamericans from South America. The results of recent studies of Paleoamerican skeletal morphology indicate that, like the Upper Cave crania, these earliest Americans do not exhibit typical “Mongoloid” morphology, but are most similar to an unspecialized, pre“Mongoloid” East Asian population (Neves and Pucciarelli, 1991; Lahr, 1995; Steele and Powell, 1992, 1993, 1994). Frequently, these first Americans resemble South Pacific modern populations (Neves and Pucciarelli, 1989, 1991; Steele and Powell, 1992, 1993, 1994; Neves et al., 1993, 1996; Powell and Steele, 1993; Munford et al., 1995; Jantz and Owsley, 2001). The Zhoukoudian crania also plot near several Archaic specimens. It is difficult to generalize

14

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

Fig. 3. CVA 3, 34 variables, N = 2,638. WGF = Wet Gravel Pit Female, WGM = Wet Gravel Pit Male, LC = Lime Creek, WB = Wizards Beach, SC = Spirit Cave, BV = Browns Valley, DL = Dry Lake, PL = Prospect Location, PR = Pelican Rapids.

about the fossils’ relationship with the Archaic Indians, as the Archaics are spread widely throughout the CV plots, rarely clustering together or with modern groups. The Archaics are obviously a diverse group that span a significant time period. Whether or not they are the descendants of the Paleoamericans or the direct ancestors of modern Amerindian groups is unclear (Cunningham and Jantz, 2001). Although the genetic data support a mostly Asian origin for today’s Native Americans (Schurr et al., 1990; Wallace and Torroni, 1992), modern Amerindian ancestry remains complicated. The relationship of UC 101 and UC 103 Although the Upper Cave fossils have some morphological features in common (Wu, 1961; Wu and Poirier, 1995), UC 101 and UC 103 are almost always widely separated on canonical axes and

have a distance between them nearly 4 standard deviations greater than expected based on modern groups. They are most similar in Analysis 3 which reflects mainly facial and a few vault dimensions (Fig. 3). If these two individuals were contemporaneous, it is interesting that their crania are so divergent. Weidenreich (1938/1939) observed these differences and affiliated the fossils with separate modern populations, although claiming that they represented a single family. The results of our analyses indicate that ancient East Asians, represented by UC 101 and UC 103, exhibit more variation than do recent populations, but not more than Paleoamerican and Archaic Indian populations. The variation between these two fossil crania may suggest separate burial events and separate populations, although this seems unlikely given the archaeological details that are known about the Zhoukoudian Upper Cave. Alternatively, Pleistocene population structure may have been

D.L. Cunningham, R.L. Jantz / Journal of Human Evolution 45 (2003) 1–18

15

Fig. 4. CVA 4, 15 variables, N = 2,639. WGF = Wet Gravel Pit Female, WGM = Wet Gravel Pit Male, LC = Lime Creek, WB = Wizards Beach, SC = Spirit Cave, BV = Browns Valley, DL = Dry Lake, PL = Prospect Location, PR = Pelican Rapids, LM = Lansing Man.

different from that seen today, something previously noted in the literature (e.g., Gill, 1990; Brown, 1992; van Vark, 1994; Lahr, 1995; Sarich, 1997; Cunningham and Wescott, 2002). Implications for theories of modern human origins Wright (1992) finds morphological resemblance between the Zhoukoudian H. sapiens fossils and crania from Australia and Africa, and asserts that this supports the Out-of-Africa theory of modern human origins. This assumes, however, that the ancient Africans involved in the exodus resemble modern Africans. Other authors associate the fossils with modern East Asians (Wu, 1956; Wu, 1961; Coon, 1962; Wolpoff et al., 1984; Turner, 1985; Wu and Zhang, 1985, Thorne and Wolpoff, 1992; Wu and Poirier, 1995; Wolpoff, 1996; Turner et al., 2000), with some claiming support for

Multiregional evolution. As with other metric studies (Howells, 1983, 1989, 1995; Kamminga and Wright, 1988; van Vark and Dijkema, 1988; Brace and Tracer, 1992; Wright, 1992; Cornell and Jantz, 1997, 1998; Brown, 1998, 1999; Cornell, 1998; Neves and Pucciarelli, 1998; see Table 1), we find no strong affiliation between the Upper Cave fossils and modern East Asians, as Asian populations are always far removed in multivariate space from the Upper Cave fossils. In fact, the Buriat, considered by Brues (1977:255) to be the “type specimens . of the Mongoloid race” because of their “classic Mongoloid” morphology, is consistently the group farthest away from UC 103 (Table 7). It is difficult to claim support for either school of modern human origins based on our results. Since the Pleistocene population of East Asia was more generalized and heterogeneous than modern Asian populations, it is possible, given the time

16

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period involved, that the specialized craniofacial form of the modern East Asians developed from the generalized morphology of the Upper Cave fossils (Hanihara, 1994). These modern Asians became morphologically specialized relatively recently (Lahr, 1995), and ancestors and descendants may not cluster together in multivariate space. On the other hand, the generalized Upper Cave individuals may have been completely replaced by the more derived ancestors of modern Asians. However, this latter scenario does not necessarily support either school of modern human origins.

The variability in the Upper Cave group that was initially recognized by Weidenreich (1938/ 1939) has two possible explanations: (1) the fossils are not representative of the same local population, or (2) Pleistocene populations were more variable than modern groups. Based on archaeological and dental evidence (Turner et al., 2000), the first option seems unlikely. Additionally, the heterogeneity of Pleistocene populations has been previously recognized in both the Old (van Vark, 1994; Cunningham and Wescott, 2002) and New Worlds (Jantz and Owsley, 2001). Acknowledgements

Conclusions Our results suggest that two of the Zhoukoudian Upper Cave fossils (UC 101 and UC 103) show similarities with modern South Pacific, European, and ancient American populations, do not resemble any modern Asian group to which they have been compared, and depict more variability than is present in modern populations. While UC 101 could be a typical member of some modern groups, especially Easter Island and European populations, UC 103 is more of an outlier. The Zhoukoudian Upper Cave fossils are not representative of any group to which they have been compared, although their vaults are similar to those of the modern Australo-Melanesians that seem to possess a more primitive, robust cranial morphology. Both fossils frequently cluster with Paleoamerican crania. Numerous studies have shown the resemblance of Paleoamericans from North and South America to South Pacific modern populations (Neves and Pucciarelli, 1989, 1991; Steele and Powell, 1992, 1993, 1994; Neves et al., 1993; Powell and Steele, 1993; Munford et al., 1995; Neves et al., 1996; Jantz and Owsley, 2001). Our results support the contention that in the Pleistocene, East Asia was inhabited by people morphologically more similar to modern Polynesians and Australo-Melanesians than to any other modern population. This also supports the idea of a relatively short time depth for modern East Asian morphology (Lahr, 1995).

We thank Danny Wescott, Carol Ward, Terry Harrison and three reviewers for their critical comments and suggestions. D.L.C. also wishes to thank Andrew Kramer, Lyle Konigsberg, and John Philpot for their assistance with this project.

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