AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 143:155–160 (2010) Brief Communication: Additional Cases of Maxillary Canine-First Premolar Transposition in Several Prehistoric Skeletal Assemblages From the Santa Barbara Channel Islands of California Sabrina B. Sholts,1* Anna F. Clement,2 and Sebastian K.T.S. Wärmländer1,3 1 Department of Anthropology, University of California, Santa Barbara, CA 93106-3210 Institute of Archaeology, University College London, WC1H 0PY, UK 3 Division of Biophysics, Arrhenius Laboratories, Stockholm University, 106 91 Stockholm, Sweden 2 KEY WORDS dental anthropology; Native American; inbreeding ABSTRACT This article identifies and discusses seven new cases of complete maxillary canine-premolar transposition in ancient populations from the Santa Barbara Channel region of California. A high frequency of this tooth transposition has been previously documented within a single prehistoric cemetery on one of the Channel Islands. A total of 966 crania representing 30 local sites and about 7,000 years of human occupation were examined, revealing an abnormally high prevalence of this transposition trait among islanders during the Early period of southern California prehistory (5500–600 B.C.). One of the affected crania is from a cemetery more than 7,000-years-old and constitutes the earliest case of tooth transposition in humans so far reported. The results are consistent with findings by other studies that have indicated inbreeding among the early Channel Islands groups. Together with the normal transposition rates among mainland populations, the decreasing prevalence of maxillary canine-first premolar transposition among island populations across the Holocene suggests that inbreeding on the northern Channel Islands had all but ceased by the end of the first millennium B.C., most likely as a result of increased cross-channel migration and interaction. Am J Phys Anthropol 143:155–160, 2010. V 2010 Wiley-Liss, Inc. In modern humans, the most common form of tooth transposition is the positional interchange of the maxillary canine and neighboring first premolar (Mx.C.P1), which normally occurs in about 0.15–0.5% of the individuals within a single population (Sandham and Harvie, 1985; Peck and Peck, 1995; Burnett and Weets, 2001; Shapira and Kuftinec, 2001). These transpositions can occur on one or both sides of the dentition, and can be completely or incompletely expressed. In incomplete transpositions, the crowns may be transposed but the roots are not, whereas in complete transpositions the entire structure of each tooth is fully transposed (Shapira and Kuftinec, 2001). In both conditions, the canine is frequently rotated mesio-labially, and the first premolar is often tipped distally and rotated mesio-lingually (Peck et al., 1993). Unlike most other tooth transpositions, Mx.C.P1 transposition does not appear to be caused by external factors such as early dentofacial trauma or tooth loss, but rather displays a genetic etiology (Peck and Peck, 1995). Elevated frequencies of associated dental anomalies, bilateral expressions, and familial occurrences, together with significant differences in male/female prevalence rates, indicate a ‘‘multifactorial inheritance model’’ for Mx.C.P1 transposition that is more complicated than for a classic Mendelian recessive trait (Feichtinger et al., 1977; Svinhufvud et al., 1988; Zilberman et al., 1990; Peck et al., 1993; Ely et al., 2006). Because of its rarity and genetic basis, Alt (1997) lists Mx.C.P1 transposition among the most useful dental anomalies for identifying closely related individuals from archeological contexts. This trait has been documented at archeological sites in prehistoric Pakistan (Lukacs, 1998) and reported in high frequencies among ancient skeletal remains from New Mexico (Burnett and Weets, 2001) and Santa Cruz Island, located off the coast on Santa Barbara, California (Nelson, 1992). In the latter study, an Mx.C.P1 transposition prevalence of over 8% was found for 106 crania from a single prehistoric cemetery (SCRI-3), which led Nelson (1992) to suggest the presence of inbreeding among this island population. In this article, 966 prehistoric crania from 30 archeological sites on the northern California Channel Islands and adjacent mainland were examined for Mx.C.P1 transpositions, allowing Nelson’s (1992) original study to be expanded both spatially and temporally. As the sample represents a time span of more than 7,000 years and a geographical area of more than 100 km2, it is possible C 2010 V WILEY-LISS, INC. C Additional Supporting Information may be found in the online version of this article. This study is dedicated to the memory of Phil Walker. Grant sponsor: Leverhulme Trust. *Correspondence to: Sabrina B. Sholts, Department of Anthropology, University of California, Santa Barbara, CA 93106-3210. E-mail: sabrina.sholts@gmail.com Received 20 December 2009; accepted 22 April 2010 DOI 10.1002/ajpa.21343 Published online 17 June 2010 in Wiley InterScience (www.interscience.wiley.com). 156 S.B. SHOLTS ET AL. MATERIALS AND METHODS Fig. 1. Map of the Santa Barbara Channel region, showing the archeological sites from which human remains with maxillary canine-first premolar transpositions have been identified. to monitor how the frequency of Mx.C.P1 transposition on the northern Channel Islands has varied throughout the Holocene and between different settlements. THE NORTHERN CHANNEL ISLANDS Until the early Holocene, a large landmass called ‘‘Santarosae’’ occupied the Pacific waters roughly 9 km from the southern California coastline, opposite the present day city of Santa Barbara (Orr, 1968). The archeological record of this landmass is long and rich, documenting 13,000 years of continuous human occupation and use (Rick et al., 2005). Around 7600 B.C. rising sea levels divided Santarosae into four separate islands, that is, the larger Santa Cruz (SCRI) and Santa Rosa (SRI) islands, and the smaller San Miguel (SMI) and Anacapa (ANA) islands (see Fig. 1), decreasing the amount of land and resources available for human subsistence and increasing water distances to the mainland shores (Porcasi et al., 1999). Although the initial colonization of the islands must have occurred via watercraft, as no land bridges from the mainland existed during the Quaternary (Rick et al., 2005), archeological evidence suggests that a fast and reliable means of cross-channel travel was not available until the invention of the oceangoing plank canoe by the historic Chumash around A.D. 500 (Arnold, 2007). The archeological record for the northern Channel Islands indicates trends of population growth and increasing settlement size across the Holocene. Kennett (2005) suggests that the earliest material remains on the islands derive from a single group of 30–50 people, and he estimates that Santa Rosa and Santa Cruz Island supported populations of 400–600 people each between 5000 and 1500 B.C. (Kennett, 2005). After 1000 B.C, King (1990) suggests that villages in the Chumash area grew two to five times their previous sizes. By the time of European contact in the sixteenth century A.D., a Chumash-speaking population of 3,000 people occupied all of the northern Channel Islands except for Anacapa (Glassow, 1977; Johnson, 1993:20). American Journal of Physical Anthropology A total of 966 crania were visually examined for the presence of Mx.C.P1 transposition. Because of poor preservation of the maxillary dentition, the presence or absence of the trait could not be determined in 94 of the crania, and these specimens were removed from the sample. For 42 of the crania in the sample, the trait was observable in only on one side of the maxilla. The final sample thus comprised 872 crania derived from archeological sites on Santa Cruz Island (n 5 442), San Miguel Island (n 5 32), Santa Rosa Island (n 5 276), and the coastal mainland of Santa Barbara County (n 5 117). The crania are held by the Phoebe A. Hearst Museum of Anthropology in Berkeley, California, the Santa Barbara Museum of Natural History, the University of California in Santa Barbara, the National Museum of Natural History in Washington, D.C., and the Natural History Museum in London, UK. The sample included male and female individuals of all ages, as determined by scores on tooth eruption, tooth wear, and sexually dimorphic cranial traits (Buikstra and Ubelaker, 1994). Each cranium, and associated mandible when present, were examined for the presence of tooth transpositions, defined as the positional interchange of two adjacent teeth within the same quadrant of the jaw, or the development or eruption of a tooth in a position normally occupied by a nonadjacent tooth (Shapira and Kuftinec, 1989; Peck et al., 1993). The transposition cases were categorized as either complete (i.e., transposition of crown and root) or incomplete (i.e., crown only) (Shapira and Kuftinec, 2001). As many archeological sites and assemblages in the Santa Barbara Channel region are not associated with reliable radiocarbon dates, specimens were classified according to a commonly used cultural chronology developed by King (1990) and modified by others (Erlandson and Colten, 1991; Arnold, 1992; Lambert, 1994; Kennett, 2005). This chronology divides Santa Barbara Channel prehistory into the Early (5500–600 B.C.), Middle (600 B.C.–A.D. 1150), and Late (A.D. 1150–1782) periods. The following sites are referred to by numbers assigned by U.C. Santa Barbara survey teams: SRI-2, 23, 241; SBA52, 260, 271, 278, 281 (Glassow, 1977). The other site numbers are those designated by the U.C. Berkeley Archaeological Research Facility (ibid.). To determine whether or not the observed trait frequencies were significantly different from the background trait frequency, chi2-tests were carried out using Yates’ 0.5 correction for sample size (Kachigan, 1991). As background frequencies in the range 0.15–0.5% have been previously reported for the Mx.C.P1 transposition trait (Peck et al., 1993; Burnett and Weets, 2001), the higher value (0.5%) was used in the chi2 statistics. RESULTS The results of this study are summarized by region in Tables 1 and 2. Among the 872 specimens in the sample, sex was determined as male for 354 crania, female for 370 crania, and ambiguous/indeterminate for 148 crania. Mandibles were associated with about half of the examined crania and expressed zero cases of dental transposition, which is consistent with previous results indicating that tooth transpositions occur predominantly in the maxillary dentition (Ely et al., 2006). Complete Mx.C.P1 transpositions were identified in seven specimens 157 TOOTH TRANSPOSITIONS IN THE CHANNEL ISLANDS TABLE 1. Total number of crania at the different sites (n) and the number of crania in which maxillary canine-first premolar transpositions were observed (Transp.), listed by time period and geographic region Early Period Region SCRI SMI SRI SBA All Site 3 83 100 103 131 147 159 No numberc Total Frequency No numberc Total Frequency 2A 2B 3A 6 24 31 34 35 40 41A 41X 50 60 76 78 147 156 No numberc Total Frequency 7 46 52 60 71 73 78 81 Total Frequency Total Frequency Housed ata PAHMA, PAHMA, PAHMA PAHMA PAHMA, PAHMA, PAHMA NMNH NHM SBMNH SBMNH SBMNH n Middle Period Transp. Transp. n Transp. 10 0 0 0 0 0 0 0 10 0 71 28 0 0 0 1 0 100 0 1 0 0 0 0 0 0 1 0 0 16 0 13 13 0 199 241 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 32 32 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 3 0 0 0 0 0 56 0 0 0 0 0 0 1 0 60 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 45 0 0 2 5 6 1 17 0 0 2 12 4 4 3 0 86 190 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 0 14 19 0 0 0 0 0 0 0 0 0 3 49 0 24 4 0 2 0 82 12 179 1 545 b 100 0 0 1 0 0 0 0 101 10 NMNH 0 0 SBMNH SBMNH SBMNH PAHMA PAHMA PAHMA PAHMA PAHMA PAHMA SBMNH SBMNH PAHMA PAHMA PAHMA PAHMA PAHMA PAHMA NMNH 0 0 25 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 26 PAHMA NMNH UCSB UCSB UCSB PAHMA UCSB SBMNH 0 0 21 0 0 0 0 0 21 1.0 0.0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0.53 0.0 0.56 0 0 0.0 0.0 0.0 8.1 0.41 0.0 7.7 148 Late Period n 0 0 0 1 0 0 0 0 1 1.2 3 0.55 a PAHMA, Phoebe A. Hearst Museum of Anthropology, Berkeley, California; NHM, Natural History Museum, London, UK; NMNH, National Museum of Natural History, Washington D.C.; SBMNH, Santa Barbara Museum of Natural History, California; UCSB, University of California in Santa Barbara. b Includes the nine cases of transposition previously identified by Nelson (1992). c Refers to specimens collected from various sites without documented numbers. (Table 2). On Santa Rosa Island: one female and one male from Early period site SRI-3 and one female from Late period site SRI-2. On Santa Cruz Island: one female from Early period site SCRI-3, one male from the Middle period site SCRI-83, and one female from Late period site SCRI-100. On the mainland: one specimen of undetermined sex from Late period site SBA-60 located within Santa Barbara county (see Fig. 1). Photos of these seven observed cases of Mx.C.P1 transposition are provided in Figures 2 and 3, and the specimens and their archeological contexts are described in detail in the online Supporting Information and summarized in Table 2. In Tables 1 and 2, the seven new cases of Mx.C.P1 transposition identified in this study are combined with the nine cases previously reported by Nelson (1992) from Santa Cruz Island site SCRI-3. The total number of reported transposition cases for the Santa Barbara Channel area is thus 16. Of these, the Early period specimen 4037.1 from site SRI-3 constitutes the oldest reported case of human Mx.C.P1 transposition known to the authors (Fig. 2a). Nine of fifteen affected individuals (60%) were female (for one individual, sex was undetermined), which is consistent with the previously reported female bias of 61% for this trait (Peck and Peck, 1995). Unilateral transpositions are present in 7 out of 14 cases, or 50% (for two cases, the trait was observable on only one side), which is less than the unilateral bias of about 75% observed in other populations (Peck and Peck, 1995). Four of the seven unilateral cases are left-sided (all female) and American Journal of Physical Anthropology 158 S.B. SHOLTS ET AL. TABLE 2. The specimens in the sample in which maxillary canine-first premolar transpositions were observed Specimen 12-76 12-4217 182-27 12-4494 4037.1 4040.1 10035 3960a 3963a 3977a 3994a 3999a 10009a 10012a 10037a 10120a Site Period Sex R L SRI-2 SCRI-100 SBA-60 SCRI-83 SRI-3 SRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 SCRI-3 Late Late Late Middle Early Early Early Early Early Early Early Early Early Early Early Early F F n/a M M F F M F F F F F M M M x – n/a x x – x x – x x – x x x x – x x – n/a x x x x x – x x x x x a The seven new cases reported in this article are in bold, while the cases previously identified by Nelson (1992) are denoted by the superscript. three are right-sided (two females, one male), showing a 57% left-side bias. These findings are in line with the previously reported bias for left-sided unilateral occurrence, especially among females (Peck et al., 1993). Maxillary C.P1 transposition has furthermore been associated with elevated frequencies of carious lesions (Nelson, 1992; Burnett and Weets, 2001), congenitally missing teeth, and peg-shaped and/or unusually small incisors (Peck et al., 1993). For the Mx.C.P1 cases identified in this study, specimen 12-4217 displays maxillary third molar agenesis on both sides (Fig. 3b). Specimens 10035 (Fig. 2d) and 182-27 (Fig. 3c) display carious lesions on most or all of the teeth present, including those that are transposed. No peg-shaped teeth were observed, although Nelson (1992) noted one case of excessively worn or possibly unusually small incisors among the crania in which he identified Mx.C.P1 transpositions (i.e., specimen 10037). These results indicate that Mx.C.P1 transposition is not strongly correlated with anomalous incisor shape in the study sample populations. Almost two-thirds of the examined crania, that is, 545 out of 872, are dated to the Late period (Table 1). This may reflect preservation and excavation biases in favor of more recent material, as well as increases in the number and sizes of settlements from the Early to the Late period (Tainter, 1977; Glassow, 1980, 1993; Munns and Arnold, 2002). For the Santa Cruz Island specimens, the prevalence rates for Mx.C.P1 transposition are 9.9% (10/101) for the Early period, 1.0% (1/100) for the Middle period, and 0.41% (1/241) for the Late period (Table 1). On Santa Rosa Island, the rates are 7.7% (2/26) for the Early period, 0% (0/60) for the Middle period, and 0.53% (1/190) for the Late period. No cases of Mx.C.P1 transposition were identified in the skeletal assemblages from San Miguel Island, and only one case was found in specimens from the mainland (Tables 1 and 2). Even though the frequency rates for Middle period assemblages on Santa Cruz Island (1.0%) and Late period assemblages on the mainland (1.2%) are slightly higher than the 0.15–0.5% background frequency (Peck et al., 1993; Burnett and Weets, 2001), these rates are based on single transposition cases within each subsample and are therefore not statistically significant (Table 1). Instead, chi2-tests reveal that Mx.C.P1 transposition frequencies significantly higher than the background level are expressed only at the Early period American Journal of Physical Anthropology Fig. 2. Specimens exhibiting maxillary canine-first premolar transposition: (a) 4037.1 from site SRI-3, (b) 4040.1 from site SRI-3, (c) 12-76 from site SRI-2, and (d) 10035 from site SCRI-3. Fig. 3. Specimens exhibiting maxillary canine-first premolar transposition: (a) 12-4494 from site SCRI-83, (b) 12-4217 from site SCRI-100, and (c) 182-27 from site SBA-60. sites of SCRI-3 (10/100 cases; X2 5 163; a \ 0.001) and SRI3 (2/25 cases; X2 5 15.2; a \ 0.001) on Santa Cruz and Santa Rosa Islands. Other time periods and other localities show normal trait frequencies. DISCUSSION AND CONCLUSIONS It has previously been argued that the high prevalence of Mx.C.P1 transposition among Early period Santa TOOTH TRANSPOSITIONS IN THE CHANNEL ISLANDS Cruz islanders was the result of inbreeding (Nelson, 1992). This scenario is supported by the results of McKern’s (1960) craniometric study of prehistoric human remains from Santa Cruz Island, where low morphological variability suggested ‘‘an inbreeding, homogenous population which probably lived in isolation for a relatively long period of time.’’ The results of this study support these conclusions, and further suggest that the early inhabitants of Santa Rosa Island were also subject to inbreeding, given the high Mx.C.P1 transposition frequency observed among crania from site SRI-3 (Table 1). The high trait frequencies on Santa Cruz and Santa Rosa Islands during the Early period suggest that both island populations descended from the same founding group, whose genetic predisposition for the trait was triggered by inbreeding. Being the largest and most ecologically diverse of the northern Channel Islands, Santa Cruz and Santa Rosa would have been able to support self-sufficient populations during the Early period. Together with small population sizes and the difficulties of cross-channel travel in simple canoes (Hudson et al., 1978), these factors appear to be a sufficient explanation for endogamy being practiced on the two islands during the Early period. Based on the low prevalence of Mx.C.P1 transposition observed for the Middle and Late periods, it appears that inbreeding did not continue on Santa Cruz and Santa Rosa Islands beyond the Early period. This could conceivably be the result of prehistoric population replacement, but population continuity on the islands is strongly supported by cultural (Rick et al., 2005; Corbett, 2007), osteological (McKern, 1955, 1960), genetic, and linguistic data (Johnson and Lorenz, 2006). Hence, the most plausible explanation for the decreasing prevalence of the transposition trait is a decline in consanguineous marriage on Santa Cruz and Santa Rosa islands after the Early period. Although the knowledge regarding marriage practices among the prehistoric Channel Island populations is limited, there is substantial evidence for the intensification of cross-channel interaction during the Middle period. Population expansion and a droughtlike climate increased the demand for mainland resources (Lambert, 1994), and the invention of improved watercraft during the late Middle period, that is, the ocean-going plank canoe (tomol), greatly facilitated transportation of both goods and people across the Santa Barbara Channel (Gamble, 2002; Arnold and Bernard, 2005; Arnold, 2007). Thus, population growth and increased cross-channel migration beginning in the Middle period provides a simple and straight-forward explanation for reduced inbreeding and a corresponding decline in Mx.C.P1 transposition rates among the Santa Cruz and Santa Rosa island populations. ACKNOWLEDGMENTS The authors thank Ray Corbett and John Johnson at the Santa Barbara Museum of Natural History, Tim White and Natasha Johnson at the Phoebe A. Hearst Museum of Anthropology, Dave Hunt and Chris Dudar at the National Museum of Natural History, and Margaret Clegg and Rob Kruszynski at the Natural History Museum in London for their assistance in providing access to the skeletal material used in this study. They also thank Michaela Huffman for her assistance with photography, and Mike Glassow, the associate editor, and two anonymous reviewers for their comments that improved the manuscript. The members and associates 159 of the Santa Ynez Band of Chumash Indians kindly supported Ms. Sholts in her research on the Chumash-affiliated skeletal remains held by the Hearst Museum. LITERATURE CITED Alt KW. 1997. Odontologische Verwandtschaftanalyse. Stuttgart: Gustav Fischer. Arnold JE. 2007. Credit where credit is due: the history of the Chumash oceangoing plank canoe. 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