Enigmatic Cranial Superstructures among Chamorro Ancestors from the Mariana Islands: Comparative Geographic Variation and a Proposal About Their Meaning

The Journal of Island and Coastal Archaeology ISSN: 1556-4894 (Print) 1556-1828 (Online) Journal homepage: https://www.tandfonline.com/loi/uica20 Enigmatic Cranial Superstructures among Chamorro Ancestors from the Mariana Islands: Comparative Geographic Variation and a Proposal About Their Meaning Gary M. Heathcote, Michael Pietrusewsky, Elizabeth Weiss, Vincent J. Sava, Bruce E. Anderson, Rona Michi Ikehara-Quebral, Michele Toomay Douglas, José M. Ramírez-Aliaga, Elizabeth A. Matisoo-Smith, Ann L. W. Stodder, Cherie K. Walth, Christopher A. King & Douglas B. Hanson To cite this article: Gary M. Heathcote, Michael Pietrusewsky, Elizabeth Weiss, Vincent J. Sava, Bruce E. Anderson, Rona Michi Ikehara-Quebral, Michele Toomay Douglas, José M. RamírezAliaga, Elizabeth A. Matisoo-Smith, Ann L. W. Stodder, Cherie K. Walth, Christopher A. King & Douglas B. Hanson (2019): Enigmatic Cranial Superstructures among Chamorro Ancestors from the Mariana Islands: Comparative Geographic Variation and a Proposal About Their Meaning, The Journal of Island and Coastal Archaeology, DOI: 10.1080/15564894.2019.1638470 To link to this article: https://doi.org/10.1080/15564894.2019.1638470 View supplementary material Published online: 18 Aug 2019. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=uica20 The Journal of Island and Coastal Archaeology, 0:1–44, 2019 Copyright # 2019 Taylor & Francis Group, LLC ISSN: 1556-4894 print/1556-1828 online DOI: 10.1080/15564894.2019.1638470 Enigmatic Cranial Superstructures among Chamorro Ancestors from the Mariana Islands: Comparative Geographic Variation and a Proposal About Their Meaning Gary M. Heathcote,1 Michael Pietrusewsky,2 Elizabeth Weiss,3 Vincent J. Sava,4 Bruce E. Anderson,5 Rona Michi IkeharaQuebral,6 Michele Toomay Douglas,7 Jos
e M. Ram
ırez-Aliaga,8 Elizabeth A. Matisoo-Smith,9 Ann L. W. Stodder,10 Cherie K. Walth,11 Christopher A. King,12 and Douglas B. Hanson13 1 Department of Anthropology, St. Thomas University, Fredericton, New Brunswick, Canada 2 Department of Anthropology, University of Hawaii at Manoa, Honolulu, Hawaii, USA 3 Department of Anthropology, San Jose State University, San Jose, California, USA 4 DPAA Laboratory, Joint Base Pearl Harbor-Hickam, Hawaii, USA 5 School of Anthropology, University of Arizona, Tucson, Arizona, USA 6 International Archaeological Research Institute, Inc., Honolulu, Hawaii, USA 7 Department of Anthropology, University of Hawaii at Manoa, Honolulu, Hawaii, USA 8 ~ del Mar, Chile Centro de Estudios Avanzados, Universidad de Playa Ancha, Vina 9 Department of Anatomy and Allan Wilson Centre for Molecular Ecology & Evolution, University of Otago, Dunedin, New Zealand 10 Office of Archaeological Studies, Museum of New Mexico, Santa Fe, New Mexico, USA 11 SWCA Environmental Consultants, Albuquerque Office, Albuquerque, New Mexico, USA 12 Scientific Consultant Services, Honolulu, Hawaii, USA 13 Formerly of The Forsyth Institute, Cambridge, Massachusetts, USA Received 20 June 2019; accepted 27 June 2019. Address correspondence to Gary M. Heathcote, 36 Brown Blvd., Unit #1A, Fredericton, New Brunswick E3A 0E4, Canada. E-mail: zinjman@gmail.com Color versions of one or more of the figures in the article can be found online at www.tandfonline.com/uica. Supplementary data associated with this article can be found in the online version. 1 Gary M. Heathcote et al. ABSTRACT Sublime expression of three ectocranial occipital superstructures (OSSs)—occipital torus tubercles (TOTs), retromastoid processes (PRs), and posterior supramastoid tubercles (TSPs)—is virtually restricted to Oceania, with epicenters in the Mariana Islands, Tonga, Mocha Island, and perhaps other Oceanic locales such as the West Sepik Coast of New Guinea. Enigmatic in etiology, OSSs are anatomically related to entheses for the trapezius, superior oblique (suboccipital), and sternocleidomastoid muscles, respectively. Our study focuses on Latte Period (950–250 BP) Chamorro ancestors of the Mariana Islands, contextualized with other skeletal samples from Remote Oceania, Near Oceania, and the Asian and American Pacific Rims. Frequent co-variation and pair-wise patterning of multiple markedly expressed OSSs distinguishes ancestral Chamorros from all other populations, but markedly expressed individual OSSs exhibit a broad network of pan-Pacific morphological affinities. The presence of markedly developed PRs and TSPs in archaic Javanese hominins indicates deep Southeast Asian origins for these morphs, but a Northeast Asian origin for tuberculated TOTs is suggested by their earliest presence in Late Pleistocene Okinawans and Neolithic Taiwanese. The central goal of this paper is to present and evaluate evidence that OSSs are informative of both Pacific population history and the life histories of “bone-forming” Pacific Islander and Pacific Rim individuals. Keywords occipital superstructures, bioarchaeology, osteobiography, Micronesia, Pacific INTRODUCTION The continuing potential for studies of comparative human skeletal anatomy to advance and test theories of Pacific Islander prehistory is not diminished by the recent growth and refinements of modern and ancient DNA studies. While molecular anthropologists and biologists are indeed at an advantage over morphologists in their ability to draw inferences about population historical relationships in a relatively straightforward manner, the study of osseous morphological variation—owing to the dynamism of bone tissue and the multiplicity of non-genetic and epigenetic factors that influence bone growth—has exclusive potential for inferring past behaviors. While all osseous morphologies have ultimate genetic underpinnings, they are commonly characterized as traits or complexes evincing 2 a more substantial genetic basis vs. those of greater non-genetic influence (e.g., phenotypes arising from plastic responses to functional biomechanical demands) (Churchill 1996). The former are considered essential to making sound inferences about evolutionary and historical relationships, while the latter are considered useful in inferring behavior and life history. Such a binary characterization of skeletal morphologies is recognized as simplistic and extreme, yet osseous characters are considered to array along a continuum of efficacy in making evolutionary vs. behavioral inferences (see Lieberman 1997). Herein, we reconsider this continuum, proposing that three particular morphoscopic traits of the posterior cranium—remarkably expressed in a minority of individuals in various Oceanic populations—may have the potential to at once inform us about VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin phylogenies of pan-Pacific populations and the life histories of a certain minority of their constituent “bone forming” members (see below). We begin with a review of previous studies, then present and discuss novel data on the spatiotemporal distribution and patterning of three enigmatic superstructures (Weidenreich 1940) of the posterior cranium: the occipital torus tubercle (TOT), retromastoid process (PR), and posterior supramastoid tubercle (TSP), collectively known as occipital superstructures (OSSs) (Heathcote et al. 1996). When present, TOTs, PRs, and TSPs occur at osteotendinous junction sites (entheses) associated with the upper trapezius, superior oblique (suboccipital), and sternocleidomastoid muscles, respectively (Figure 1). Arguably homologous to the better known musculoskeletal stress markers (MSMs) of the upper and lower limbs (e.g., Hawkey and Merbs 1995; Weiss 2004, 2010), minimal OSS expressions range from ectocranial swellings to raised rugosities or crests, while moderate to strong expressions appear as tubercles, tuberosities, and processes along a continuous morphological gradient. As a means of assessing the utility of OSSs in phylogenetic and osteobiographical inquiry, we focus on the patterning of OSSs among ancestral Chamorros of the Mariana Islands and the sharing of these cranial variants, by degree, patterning, and co-variation, with other Oceanic and Pacific Rim populations. A previous paper on the functional gross anatomy and microanatomy of OSSs among ancestral Chamorros presented an amendment to our working multifactorial model of OSS development, which proposed that among a significant minority of genetically predisposed individuals, motor behaviors trigger calcification of new fibrocartilage at enthesis sites associated with OSSs (Heathcote et al. 2014). That study presented a complication to this model resulting from a limited scanning electron microscopy (SEM) study by Timothy Bromage (New York University), who found widespread surface distribution of microstructures, appearing as OSS microforms, associated with vascular canals on both TOT superstructures and surrounding squamous bone. Such topographic Figure 1. Retromastoid process (PR), tubercle on the occipital torus (TOT), and posterior supramastoid process (TSP), in relation to musculature and anatomic features discussed in the text. Illustration on right shows these three posterior cranial superstructures on the cranium of a 40 to 50 year old male ancestral Chamorro from the Gogna-Gun Beach site, Guam (Burial No. 123). Permission to reprint granted by John Wiley and Sons. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 3 Gary M. Heathcote et al. distribution and vascular association of these microstructures is indicative of a systemic effect (Heathcote et al. 2014:1019). Notwithstanding the SEM results and implications, a role for mechanical traction in OSS development and expression continues to be supported by functional musculoskeletal considerations and bone remodeling at enthesis sites. Also, activityinduction is supported by independent findings of significant positive correlation between humeral robusticity and OSS expression in ancient Native Californians (Weiss 2010) and Mariana Islanders (Heathcote et al. 2012b). Our contention that mechanical induction serves as a trigger to, but is not the totality of, OSS development is theoretically aligned with the work of Rogers and colleagues (1997) regarding “bone formers.” Their study suggested that “observed variation in hyperostotic bone formation (enthesophytes and osteophytes) could be due to differences in individual ability to form bone in response to stress rather than differences in stress (Rogers et al. 1997:90).” In other words, genetic variation pre-conditions both individual responses and population variation in reactive bone formation at sites of musculoskeletal stress and injury. Earlier papers noted that marked expressions and co-occurrence of these OSSs appeared to be uncommon or rare in anatomically modern humans (AMH) outside of Oceania (i.e., geographic Micronesia, Polynesia, and Melanesia), New Guinea, and Australia. Further, within Oceania, markedly developed and co-varying OSSs appeared to be most frequent among ancestral Chamorros and Tongans (Heathcote et al. 2012a, 2012b; Sava 1996; Sava and Pietrusewsky 1995). Because this geographic circumscription proposal was overly dependent on negative evidence, we acknowledged its provisional nature. We have substantially expanded the comparative database on OSS frequency and expression in this paper, allowing better advancement and discussion of our proposal that OSSs have the potential to, at once, address questions about Pacific 4 population history life histories. and individual CHAMORRO ORIGINS AND DISTINCTIVENESS Chamorros are unique genetically, mirroring their morphological,1 archaeological,2 and linguistic3 distinctiveness (Vilar et al. 2013a, 2017). Concerning maternally inherited mitochondrial DNA (mtDNA), the vast majority of modern Chamorros (92%) possess haplogroup E lineages that are otherwise rare in Oceania, but well represented in Island Southeast Asia (ISEA) (Vilar et al. 2013a). Further, Chamorros have lineage branch tips belonging to haplogroup E1 and E2 clades that are unknown outside of the Marianas (Lum and Cann 2000). Indeed, a complete genome mtDNA analysis of 32 modern Chamorros found 16 unique haplotypes (Reiff et al. 2011). The Vilar team has concluded that the uniqueness and limited diversity of modern Chamorro E1 and E2 lineages suggest an old (Pre-Latte) introduction from founder groups from eastern Indonesia (Hill et al. 2007; Soares et al. 2008), with Sulawesi and the Moluccas being the likeliest source of Chamorro E1a2 and E2a lineages (Vilar et al. 2013a, 2017). Among the minority of modern Chamorros exhibiting mtDNA haplogroup B4 lineages, there is a paucity of pan-Pacific lineages (“Polynesian motif” and closely related subclades), so dominant in all other Micronesian and Polynesian populations; however, 7% possess one that is unique to the Mariana Islands of Rota and Guam, viz. a B4a1a1a lineage branch tip, provisionally known as the “Chamorro motif” (Vilar et al. 2013b). Ancestral derivation from Indonesia is suggested by Y chromosome data as well, with some connection to the Philippines (Tabbada et al. 2010). A Y-DNA haplogroup O1a1b branch tip, which appears to be more frequent in Chamorro males than counterparts in ISEA, is estimated to have arisen 1,950 years ago. Modern Chamorro Y-DNA indicates paternal gene flow from Northeast and Southeast Asia, Oceania, VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin Europe, and Native America. European admixture, predominantly from the Western Mediterranean and Iberia, is found in 16% of male, but in no female lineages, reflecting male-biased gene flow. The 2–6% Native American admixture found likely derives from the contributions of seventeenth- and eighteenth-century Mexican mestizos (of mixed Native AmericanEuropean ancestry) to the Chamorro gene pool (Vilar et al. 2017). The mtDNA distinctiveness of Chamorros, together with their reduced genetic diversity (Vilar et al. 2013a), is consistent with a population history impacted by colonization of the Marianas by small groups from ISEA, coupled with subsequent long-term genetic isolation by distance4 and significant bottleneck effects (depopulation events) during Colonial times (Garruto 2012; Reiff et al. 2011; Vilar et al. 2013a).5 If Vilar and colleagues are correct in their stratigraphic deconstruction of Chamorro population structure, Latte and Early Historic Period Mariana Islanders likely included individuals who possessed ancestral mtDNA haplogroup E1 and E2 lineages that can be traced backwards to homelands of their most ancient ISEA ancestors, as well as haplogroup B4 lineages introduced by later ISEA migrants to the Marianas. The results of these mtDNA studies is supportive of a dual origin model of Marianas settlement, namely that between 3230 and 3085 cal. BP (2r), small founding populations from ISEA reached the archipelago (Rieth and Athens 2017), introduced a red-slipped pottery known as Marianas Red, and developed unique mutations to mtDNA haplogroup E1a and E2a lineages. Much later, around 1,000 BP, a second migration from ISEA introduced latte stone architecture and the practice of rice horticulture (see Hunter-Anderson et al. 1995), as well as a unique mtDNA haplogroup B4 lineage (Vilar et al. 2013a). Further, certain variants of the Y-DNA haplogroup O1a1b lineage may have been introduced by this later wave of ISEA migrants (Frank Camacho, personal communication 2018; Vilar et al. 2017). SKELETAL SERIES AND PROVENANCE In this study, 593 adult crania from Remote and Near Oceania (Green 1991), Asia, and the American Pacific Rim (Figure 2) were scored for the presence and degree of development at the TOT, PR, and TSP sites (Heathcote et al. 1996). Of these, 379 (63.9%) were excavated or collected from three of the Mariana Islands (Guam, Tinian, and Saipan) (Figure 3), and 38 (6.4%) originated from elsewhere in Remote Oceania, namely geographic Micronesia (Palau, Yap, Chuuk, the Marshalls, Kiribati, and Nauru), Melanesia (Fiji), and Polynesia (Tonga, Marquesas, and Rapa Nui). Of the remainder, 24 (4.0%) derive from sites along the Asian Pacific Rim (Taiwan and Cambodia), 58 (9.8%) from New Guinea, 85 (14.3%) from the North American Pacific Rim, and 9 (1.5%) from the South American Pacific Rim. The skeletal series studied, as well as their geographic provenances, sample characterizations (type), adult sample sizes by sex, investigators who scored the crania, present locations of the skeletal collections, and chronological dates or archaeological associations are presented in Table 1. Of the non-Marianas Pacific Islander crania, those from Remote Oceania are of uncertain temporal provenience, with the exception of the Tongan sample and an immediately post-Lapita burial from Fiji. The three Near Oceania samples from Papua New Guinea were dated approximately by historical inference, while all of the Asian and American Pacific Rim samples were dated chronometrically. Most of the Mariana Islanders studied lived during the Latte Period of Chamorro cultural history. Human burials are frequently found in association with latte sets in coastal sites. Since the pioneering archaeological work of Spoehr (1957), preEuropean history of the Marianas has been divided into two main Periods, Pre-Latte (beginning 3500 BP; but see Rieth and Athens [2017] who argue for a later date, based on recent refinement of radiocarbon chronologies) and Latte, the division between the two dating to around 950 BP. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 5 Gary M. Heathcote et al. Figure 2. Map of Oceania and the Pacific Rims showing the approximate locations (in bold) of the skeletal series studied by the authors. Locations in non-bold lettering are places mentioned in the text, regarding osteological, archaeological, genetic, and linguistic studies by other researchers. Latte refers to sets of megalithic columns (haligi) with capstones (tasa), which usually served as foundations for wooden house structures built in the quintessentially distinctive architectural style of the Marianas (Athens and Ward 2010; Morgan 1988:116–149; Russell 1998b:219–227). More recent, finer-grained cultural chronologies have been reviewed by Rainbird (2004). Here, we follow Moore’s (2002) construction: Early Unai (Early Pre-Latte), 3450–2950 BP; Middle Unai (flatergEarly Pre-Latte), 2950–2450 BP; Late Unai (Intermediate Pre-Latte), 2450–1550 BP; Huyong (Transitional), 1550–950 BP; and Latte, 950–250 BP. References herein to the Early Historic Period generally refer to 429–250 BP. Overlap of the latter with the Latte Period acknowledges that, while initial contact with Europeans was made in 429 BP (i.e., AD 1521 with Magellan’s contact), the indigenous settlement system and culture persisted until around 250 BP 6 (Moore 2002; Anderson 2005). see also Hunter- METHODS Age and sex estimations were based on standard anthropological techniques (e.g., Buikstra and Ubelaker 1994; Byers 2002; Ubelaker 1999). Crania of skeletally immature individuals are excluded from consideration here due to the difficulty in sex determination of sub-adults; also OSS formation, like other forms of bone formation at entheses, is most prominent during adulthood (Weiss 2010, 2017). When permitted by the structural completeness of crania and/or the presence of associated infracranial skeletal elements, individuals were assigned to the following age cohorts: Young Adults (20–30 years of age), Middle Adults (31–50 years of age), and Old Adults (more than 50 years of age). VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin Figure 3. Map of the Mariana Islands, showing the approximate locations of the skeletal series from Saipan, Tinian and Guam studied by the authors. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 7 8 Table 1. Adult crania from Oceania and the Asian and American Pacific Rims, scored for TOT, PR, and TSP superstructures. Geographic Provenance & Skeletal Sample VOLUME 0  ISSUE 0  2019 MARIANA ISLANDS Guam Agana (Hagåt~ na), von Luschan Collection Camp Watkins Road Project Chaot/Agana (Hagåt~ na) Sewer Project Duty Free Expansion Site Gognga-Gun Beach Number of Crania Scored2 F M T Scored by3 Present Location4 Dating and/or Archaeological Association Regional 1 2 3 GH AMN Dating uncertain 3 3 GH GUM Dating uncertain AMN records Collected from Agana, 1874–1878 PHRI (n.d.a) Selected 0 Selected 0 1 1 GH GUM Latte Period PHRI (n.d.b) Locality 1 1 1 RI GUM Latte Period Magnuson et al. (2000) Locality 20 18 38 GH with BA [reburied] Anderson (1992) Kurashina (1992) Regional 59 77 136 VS with GH GUM 1070 ± 70 to 290 ± 50 BP; dates derived from faunal and floral samples associated with a small subset of burials Undated, but an estimated 80% of the remains are from Latte Period sites along Tumon Bay. Others derive from coastal locales that include Agana Bay, Amantes Point, Jinapsan, and Tarague. References and Remarks GUM records Ikehara-Quebral (unpublished) Hornbostel (1924–1925) Thompson (1932) Pietrusewsky (1971) Graves and Moore (1985) Howells (1989) Sava (1996) Heathcote (2000) (Continued) Gary M. Heathcote et al. HornbostelThompson Collection aka Tataotao Manaina (the Body of the Elders) Sample Type1 Table 1. (Continued). Hyatt Hotel Selected 1 1 GH [reburied] Locality 1 1 2 GH [reburied] Merizo (Malesso) Locality 1 8 9 MP&MD GUM Historic Period Naton Beach, Latte Burials Locality 22 20 42 CW SWCA Latte Period Naton Beach, PreLatte burials Locality 17 23 40 CW SWCA Tumon Bay, Bordallo Donation Isolated 1 0 1 GH USM Early to Intermediate Pre-Latte Period, viz. Middle to Late Unai times, based on14C dates of burials. Dating uncertain Outrigger Hotel Ylig Bay Locality Locality 3 1 4 4 7 5 RI MP&MD [reburied] GUM Latte Period One Intermediate PreLatte, viz. Late Unai (1580 BP) radiocarbon date obtained; site possibly occupied from Pre-Latte to Post-European contact times. Isolated 1 0 1 DH [reburied] Early Historic Period Trembly with Tucker (1999) Ryan (2010) Douglas and Ikehara (1992) PHRI records Pietrusewsky and Douglas (unpublished) DeFant (2008) DeFant and Eakin (2009) Walth with Parr and Walborn (2013) DeFant (2008) DeFant and Eakin (2009) Walth with Parr and Walborn (2013) USM records Gift to USM from Governor Ricardo Bordallo Ikehara-Quebral (1998) Pietrusewsky et al. (2009) Hanson (1995) 9 (Continued) Occipital Superstructures across the Pacific Basin THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY Leo Palace Transitional to Early Historic; 1539 to 251 BP (for entire assemblage) Latte Period Saipan Achugao 0 10 Table 1. (Continued). Agingan Locality 1 2 3 GH NMM Latte Period VOLUME 0  ISSUE 0  2019 Locality 3 2 5 MP NMM Chalan Laolao, Beach Road Hazard Project Isolated 0 1 1 MP NMM Chalan Monsignor Guerrero Road Locality 4 4 8 MP NMM Associated cultural deposits range from Pre-Latte to Latte Period. Project area includes Japanese Period and postinvasion U.S. military camps. Dental and cranial features consistent with Chamorro ancestry. Bone sample submitted for radiocarbon dating. 890 to 500 BP Grand Mariana Resort, Anaguan (Garapan) Kalabera Cave Locality 1 13 14 CK LBG 860 to 790 BP Locality 0 1 1 GH NMM Dating uncertain Locality 0 2 2 GH FMN Pre-Latte, on basis of stratigraphy and pottery Laulau Rock Shelter Pietrusewsky (2009) Pietrusewsky and Douglas (2001a) Pietrusewsky et al. (2014) Perzinski and Dega (2016) NMM records Hornbostel (1924–1925) Cabrera and Tudela (2006) FMN records Spoehr (1957) (Continued) Gary M. Heathcote et al. Chalan Kanoa, Beach Road Sewer Project Spoehr (1957) Pietrusewsky (1971) Pietrusewsky (2006b) 9 9 18 VS NMM Prior to 429 BP Pietrusewsky and Douglas (1989) Pietrusewsky et al. (2014) Tinian Blue Site, Blue 1 Locality 2 1 3 GH FMN Latte Period Route 201 Project Isolated 0 1 1 MP RPT Route 202 Project Regional 1 4 5 MP RPT Remains likely from a prehistoric site, near the House of Taga, that was impacted several times during the Historic Period 941 to 339 BP FMN records Spoehr (1957) Pietrusewsky (2010) House of Taga Locality 4 10 14 GH NMM Latte Period Waterline Project Locality 6 8 14 MP RPT 750 to 310 BP ELSEWHERE IN MICRONESIA and REMOTE OCEANIA Palau Islands von Luschan Regional 4 5 9 Collection and Locality unknown GH AMN Dating uncertain Pietrusewsky and Douglas (2010) Pietrusewsky et al. (2014) NMM records Hornbostel (1924–1925) Thompson (1932) Spoehr (1957) Heathcote et al. (2012b) Pietrusewsky and Douglas (2001b) Pietrusewsky et al. (2014) AMN records Island provenances were not recorded (Continued) 11 Occipital Superstructures across the Pacific Basin THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY Table 1. (Continued). Oleai Locality 12 Table 1. (Continued). Yap Islands von Regional Luschan Collection Chuuk Islands von Luschan Collection VOLUME 0  ISSUE 0  2019 Marshall Islands von Luschan Collection Locality unknown Nauru von Luschan Collection Kiribati von Luschan Collection 5 8 GH AMN Dating uncertain AMN records Probably all from Yap Proper, but this is uncertain Regional 3 2 5 GH AMN Dating uncertain AMN records One male from Pulap; all others from Mortlock Islands Regional 1 1 2 GH AMN Dating uncertain Isolated 0 1 1 GH FMN Dating uncertain AMN records From Jaluit Atoll FMN records Isolated 0 1 1 GH AMN Dating uncertain AMN records Regional 0 2 2 GH AMN Dating uncertain AMN records One from Tautau; one (“probably Gilbertese”) had lived in Jaluit Isolated 0 1 1 MP UHM Associated with immediately postLapita ceramics; dated to 2700 BP Pietrusewsky et al. (1997) Petchey et al. (2010) Pietrusewsky and Douglas (2016) (Continued) Gary M. Heathcote et al. Fiji Waya Island 3 Table 1. (Continued). Tonga Pangaimotu Locality 5 6 VS with GH BPB Pre-European contact commoners McKern (1929) Sava and Pietrusewsky (1995) Sava (1996) Isolated 0 1 1 GH AMN Dating uncertain AMN records Isolated 0 1 1 GH FMN Dating uncertain FMN records Locality 8 10 18 MP TSP Burials associated with Tapenkeng Culture (5450 to 4950 BP), the earliest Neolithic sequence in Taiwan Tsang (2005) Pietrusewsky et al. (2013) Cambodia Vat Komnou Locality 2 4 6 RI UHM 2150 to 1750 BP Pietrusewsky and Ikehara-Quebral (2006) IkeharaQuebral (2010) NEW GUINEA West Sepik Coast, PNG Arop Locality 11 9 20 MD&AS FMN Samples consist of the remains of people presumed to have died in the mid to late 1800s. Skulls were obtained through trade, and not by grave robbing or plundering cult houses. Welsch (1998, 2000) Douglas and Stodder (2010 ) Stodder (2011) Marquesas Islands Locality unknown Rapa Nui Locality unknown ASIAN PACIFIC RIM Taiwan Nankuanli East 13 (Continued) Occipital Superstructures across the Pacific Basin THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 1 14 Table 1. (Continued). Sissano Locality Warapu Locality VOLUME 0  ISSUE 0  2019 NORTH AMERICAN PACIFIC RIM San Francisco Bay, California Ryan Mound Site Locality SOUTH AMERICAN PACIFIC RIM Mocha Island, Chile Concepcion Regional Museum Collection TOTALS 6 5 11 MD&AS FMN - ditto - 12 15 27 MD&AS FMN - ditto - 48 37 85 EW SJS 2180 to 250 BP Weiss (2010) 3 6 9 GH with EM &JR CMC Many burials derive from archaeological contexts that are PreColumbian (950 to 550 BP) Ramirez-Aliaga (unpublished) Matisoo-Smith and Ramirez (2010) 320 593 273 Welsch (1998, 2000) Douglas and Stodder (2010) Stodder (2011) Welsch (1998, 2000) Douglas and Stodder (2010) Stodder (2011) 1 Gary M. Heathcote et al. Isolated ¼ Single individual; Locality ¼ Sample recovered from a former village/occupation site or locality; Regional ¼ Sample is a regional composite assemblage of individuals from different localities; Selected ¼ Individuals selected non-randomly, from a larger sample. 2 F ¼ female; M ¼ male; and T ¼ total. 3 AS ¼ Ann Lucy Stodder; BA ¼ Bruce Anderson; CK ¼ Christopher King; CW ¼ Cherie Walth; DH ¼ Douglas Hanson; EM ¼ Elizabeth Matisoo-Smith; EW ¼ Elizabeth Weiss; GH ¼ Gary Heathcote; JR ¼ Jose-Miguel Ram
ırez-Aliaga; MD ¼ Michele Toomay Douglas; MP ¼ Michael Pietrusewsky; RI ¼ Rona Ikehara-Quebral; VS ¼ Vincent Sava. 4 AMN ¼ American Museum of Natural History, New York City; BPB ¼ Bernice P. Bishop Museum, Honolulu; CMC ¼ Museo de Historia Natural de Concepci
on, Concepci
on, Chile; FMN ¼ Field Museum of Natural History, Chicago; GUM ¼ Guam Museum, Hagåt~ na, Guam; LBG ¼ Lower Base Garapan, Scientific Consultant Services Inc. temporary storage and curation station in Garapan, Saipan; NMM ¼ Commonwealth of the Northern Marianas (CNMI) Museum of History and Culture, Saipan, CNMI; RPT ¼ Re-interment Pit at the House of Taga site, Tinian. Administered by the Tinian Historic Preservation Office, Tinian, CNMI. Remains are available for further study, upon successful petition; SJS ¼ San Jos
e State University Anthropological Collection, San Jose, California; SWCA ¼ SWCA Guam Environmental Consultants, Tamuning, Guam; TSP ¼ Tainan Science Park, Tainan City, Taiwan; UHM ¼ Anthropology Department, University of Hawaii at Manoa; USM ¼ U.S. National Museum, Smithsonian Institution, Natural History Building, Washington, DC. Series presently located at GUM and NMM were formerly curated at the BPB, and were repatriated to Guam and the Commonwealth of the Northern Marianas in 2000 and 1999, respectively. Occipital Superstructures across the Pacific Basin Table 2. Five-point scale (0–4) for scoring developmental expression of TOT, PR, and TSP superstructures. Illustrations provided in Heathcote et al. 1996. Associated Morphology TOT: Occipital Torus Tubercles 0 Highest nuchal line (HNL) and superior nuchal line (SNL) are barely, if at all, palpable. 1 HNL and SNL are clearly defined (and palpable), but torus not present or is only incipiently developed. 2 Torus clearly developed, with increased rugosity at trapezius site, but there are no discrete tubercles present, i.e., there is nothing to "grab hold of". 3 Torus is well developed, with discrete ("grab hold of"), projecting tubercles at the trapezius site. 4 Torus is well developed, with massive, sometimes pedunculated, tubercles at the trapezius site. In a practical sense, a score of "4" (massive) is attained when the tubercle appears to be large enough to grasp between the thumb and index finger and so lift or suspend the cranium. PR: Retromastoid Process 0 Incipient, or no, discernible elevation inferior to the SNL at the superior oblique site. 1 Slight mounding at this site. The mounding is <3 mm elevation above the squamous occipital surface on the medial side. 2 Moderate mounding at this site. The mounding is 3–5 mm in elevation (as above). 3 Well developed mounding at this site, presenting as a truly retromastoid process. The process is 5.1–10 mm in elevation (as above). 4 Massive development at this site. The process is >10 mm in elevation (as above). TSP: Posterior supramastoid tubercles 0 Incipient, or no, elevation of bone immediately lateral and anterior to asterion. 1 Slight, palpable swelling present at this site. 2 Moderate development; the site appears mounded, rather than tuberculated. Elevation of the mounding is <3 mm, measured from the parietal squamous surface immediately superior to the site. 3 Well developed, discrete ("grab hold of") tubercles present. Tubercles measure 3–5 mm in elevation (as above). 4 Massive development. Discrete tubercles are conical and pedunculated, and measure >5.1 mm in elevation (as above). An illustrated protocol (Heathcote et al. 1996) was used to score continuously varying OSSs according to a fivepoint (0–4) scale (Table 2). Superstructures scored as “3” or “4” are referred to as marked or markedly developed OSSs; scores of “2” as moderate and “1” as slight developments; while “0” indicates no or incipient expression. Systematically produced data on measurement error are not available, but informal intra-observer precision testing indicates that a high degree of replication can be achieved with sufficient practice. Visual standards provided in our protocol likely reduced the magnitude of interobserver error, as suggested by Walker’s (2008) finding that illustrations reduced inter-observer imprecision in a study of morphoscopic skull sexing. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 15 Gary M. Heathcote et al. OSS development was evaluated in two ways. Data for 556 of 593 crania were produced through direct visual scoring of crania. The remaining 37—from Palau, Yap, Chuuk, Marshall Islands, Nauru, Kiribati, Marquesas, and Mocha Island—were scored from photographic images. Images of the Mocha Island crania were shared by JoseMiguel Ram
ırez-Aliaga and Elizabeth Matisoo-Smith, while photographic slides of the other samples were provided by Jaymie Brauer and Susan Rodriguez (American Museum of Natural History), following a protocol that employed both standard and modified anatomic views to allow maximum visualization of OSSs. Data were entered, screened, tabulated, and analyzed using Microsoft Excel 2007 and SAS for Windows, Version 9.2. PREVIOUS STUDIES OF OSSS In his theses, Hublin (1978c) provided an overview of “supranuchal tubercles” (i.e., TOT), PR, and TSP—among a number of other superstructures associated with the occipital torus (OT)—in archaic hominins and anatomically modern humans (AMHs) from Africa, East Asia, the Middle East, and Europe. The occipital torus is a variably marked transverse bony ridge along the squamous part of the occipital; it is sublimely expressed in Homo erectus (sensu lato) and variably encountered in AMHs (Lahr 1994). Hublin’s theses and other works (Hublin 1978a, 1978b, 1983, 1988, 1989) provide a foundation for this review. Regarding AMHs, previous researchers have generally not employed scoring standards to categorize posterior cranial superstructure development.6 Subjective attributions such as “weak,” “average”, or “strong” are generally employed without definition or illustration, making it difficult to establish concordances with our data. Experiential differences also impact interobserver comparability; for example, Hauser and De Stefano (1989:108) recommend that a PR of greater than 1 mm protrusion be classified as “strong”, suggesting unfamiliarity with the 16 exceptional degree of retromastoid process development within some Oceanic populations, for we would score such a PR as incipient or slight (Table 2). A further problem of inter-observer comparability is the frequent pooling of males and females in frequency tabulations by nineteenth- and early twentieth-century researchers; this skews inter-group comparisons, for sexual dimorphism in OSS expressions is virtually universal. Last, earlier workers frequently collapsed cases into vast geographic categories, nation states, or “races”, analytical units of dubious population biological meaning. These problems notwithstanding, comparative frequency data on occipital torus (OT), PR, and TSP expression are presented in Supplementary Data: Appendix Tables 1, 2, and 3, respectively. A minimum sample size of 20 was chosen as the criterion for inclusion in these tables. Regarding OT (see also TOT) prevalences, the proportion of “strong expression” scores that equate to TOTs is unknown, except for Hublin (1978c), who was explicit about OT cases bearing supranuchal tubercles (i.e., markedly expressed TOTs). Tubercles on the Occipital Torus (TOT) and Occipital Torus (OT) The earliest markedly developed TOTs known to us are seen in three Late Pleistocene AMH crania from Minatogawa, Okinawa (Suzuki 1982). Minatogawa I, II, and IV, an adult male and two adult females respectively, feature two “oval tuberosities” with connecting lineal ridges, each situated on a pronounced occipital torus. Judging from illustrations (Suzuki and Hanihara, 1982:plate 2.6), we would score all three individuals as having a marked (“3”) degree of TOT development. The former individual may date from around 18,000 BP, and the latter two may be somewhat younger, based on multielement analysis and relative chronology (Matsu’ura and Kondo 2011). Despite chronological age differences, these three individuals may belong to a single VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin morphological group (Matsu’ura and Kondo 2011), but this is disputed (Baba 2002; Brown 1999). In a broadly comparative overview for its time, Hagen (1880) reported that the greatest degrees of OT development in AMHs were found in aboriginal Australian, Australian (Torres Strait) Islander, and East Indian groups, followed by those from New Guinea, New Zealand, Hawaii, and Tahiti. Merkel (1871) and Joseph (1873) earlier reported high frequencies of OT in Papuans (of New Guinea) and Australians, and Matiegka (1906) later recorded a 14% frequency of “strong” OT in “Oceaneans.” Hublin (1978c) echoed these findings, reporting OT (all degrees of expression) in 61% of a combined aboriginal Australian and Tasmanian sample, followed by New Caledonians (46%), Japanese (40%), Greenlandic Inuit (13%), Parisians (11%), and Dogons (1.9%). Only Melanesians (New Caledonians) and AustralianTasmanians had nontrivial frequencies of “strong” OT; each sample included two cases with “strong and typical” supranuchal tubercles (TOTs) (Supplementary Data: Appendix Table 1).7 Retromastoid Processes (PR) Of the three OSSs considered herein, only the retromastoid process is substantially documented in the human paleontological record. The earliest hominin known to bear a prominently developed PR is the early Pleistocene East African Homo erectus (sensu lato) KNM-ER3733, who dates to approximately 1.7 MYBP (Lepre and Kent 2010). Leakey and Walker (1985:149) described this cranium as having a “bony spur,” more prominent on the left, just medial to the occipito-mastoid suture, along the lateral portion of the superior nuchal line. Later, Rightmire (1990) identified these structures as retromastoid processes. A cast of this specimen was examined by one of us (GMH), and the PRs were scored “3” (marked) and “1” (slight) on the left and right sides, respectively. PRs have also been attributed to at least four later archaic hominins from Africa and Europe, and one central European Upper Paleolithic individual, but none are markedly expressed.8 In contrast with African and European fossil hominins, PRs are much more frequently encountered among Early to Late Pleistocene hominins in Java (Supplementary Data: Appendix Table 4). In his classic study of the Ngandong (“Solo Man”) crania, Koenigswald (1951) described the retromastoid process as a “special character.” This was independently corroborated by Santa Luca (1980), who reported PRs (“weakly” and “strongly” developed) in six of seven Ngandong crania; Hublin (1978c:174), who found the PR to be a characteristic and “stable” feature throughout the fossil hominin record in Java; and Rightmire (1990), who reported varying degrees of PR development in seven of nine (78%) Javanese archaic hominins from Sangiran (n ¼ 3), Sambungmachan (n ¼ 1), and Ngandong (n ¼ 5).9 As Weidenreich (1943) made no mention of PR in his detailed work on Middle Pleistocene H. erectus skulls from Zhoukoudian, China, a north-south Asian distributional discontinuity for this superstructure is suggested during the Pleistocene.10 Among early studies of AMHs, Le Double and Dubrueuil-Chambardel (1905), Schlaginhaufen (1906), and Waldeyer (1909) reported that Pacific populations are generally distinguished by high frequencies of PR. Waldeyer’s paper included broad inter-regional comparisons, but his total frequencies appear inflated alongside those of other observers (Supplementary Data: Appendix Table 2).11 However, his qualitative attribution of prominent PRs to Australians, Papuans, and “other Melanesians” seems accurate, as it is echoed by contemporaries. Michelsson (1911) wrote of pronounced PR development in Papuans, and Le Double and DubrueuilChambardel (1905) considered PR to be practically a “racial character” of Papuans, adding that it is frequent among Polynesians and rare in Europeans. None of these researchers provided scoring criteria, THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 17 Gary M. Heathcote et al. but Waldeyer (1909:figures 5 and 6) illustrated a Papuan skull possessing what we would score as a markedly developed PR. Curiously, despite such qualitative characterizations by these authors, their reporting of “medium” and “strong” PR frequencies for Pacific Islanders (Melanesians and Papuans) ranges from only 1.8% to 5.9% (Supplementary Data: Appendix Table 2). Exceptional expressions of PR among Mariana Islanders, as well as Caroline Islanders (Yapese, Chuukese, and Pohnpeians), and Marshall Islanders, have been reported previously. In an early twentieth-century paper concerned primarily with craniometry, Schlaginhaufen (1906) stated that two thirds of Chamorro skulls examined showed weak to strong expressions of PR. His study was based on 23 crania from Saipan, collected by Georg Fritz in 1899 (Fritz 1986 [1904]). In a later study based on small samples of skulls from Yap, “Truk” (Chuuk), Saipan, and Rota, Arai (1970) noted that the “retromastoid eminence” was found in 8% of Japanese and 1.3% of Europeans, and characterized it as “frequent” in Micronesians. Most recently, Douglas et al. (1997) reported two males (2/15 or 13.3%) and no females (0/10) with markedly developed PRs in a sample of Latte Period Chamorros from Apurguan (a.k.a. Apotguan), Guam. In a survey of living males from Micronesia, Hasebe (1935) reported manually palpated PRs in 12% of individuals (18/ 150) from Pohnpei, 37% (26/71) from the Ralik, and 16% (27/167) from the Ratak chains of the Marshall Islands. Palpated PRs would necessarily be markedly expressed, as such processes and attached superior oblique muscles underlie three muscle layers (from superficial to deep), the trapezius, semispinalis capitis, and splenius capitis (Anderson 1983:figures 5-32 to 5-34). Posterior Supramastoid Tubercles (TSP) We are aware of only one fossil hominin that bears posterior supramastoid tubercles, Ngandong 1 from Java (Hublin 18 1978c:95), illustrated in Santa Luca (1980:22–24). Examination of a cast by Jean-Jacques Hublin (personal communication 1996), indicated a degree of expression intermediate between “2” and “3” (i.e., the site is mounded with only incipient tubercle development). As for earlier hominins, Kimbel and Rak (1985) made no mention of TSPs in their study of the asterionic region among Australopithecus afarensis, A. africanus, A. robustus, A. boisei, and early Homo crania. However, it must be conceded that the presence of nuchal crests in some of these specimens problematizes the identification of TSPs. The earlier literature on AMHs provides limited comparative data on posterior supramastoid tubercles of the peri-asterionic region. Jacob (1967) was the only previous researcher who scored TSP systematically; his study included 316 skulls from four regional groupings, “American Negroes”, Alaskan “Eskimos”, Central Europeans, and Central Javanese (Supplementary Data: Appendix Table 3). Scoring criteria were made explicit; for example, a “grade 3” (maximum score) TSP was defined as very distinct with an observable tubercle. However, his illustration of a “grade 3” TSP (Jacob 1967:63) equates with what we score as moderate expression, so it would appear that no markedly developed TSPs were encountered in his study.12 More useful is the early anatomical study and review of Matiegka (1906). Geographically broad in scope (Africa, Europe, Asia, the Americas, and Oceania), it included discussions on the distribution of the Processus asteriacus (asterionic process or AP), treated herein as a variant homologue of the TSP. Unfortunately, the absence of illustrations and ambiguities in the use of anatomical terminology frustrate data mining from his paper.13 Nonetheless, Matiegka makes it clear that he found the “best examples” of AP in skulls from Oceania. APs are described as “truly enormous” in a Chatham Islander, “most strongly developed” in a Maori, “relatively strong” in a Tasmanian, and “variably strong” in skulls from New Caledonia, VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin Hawaii, and Tierra del Fuego (which he typed as “Oceanic”) (Matiegka 1906:369). He also noted a “button like” protuberance at the AP site on a Fijian skull, as well as on a morphologically anomalous skull from southern Bohemia (Matiegka 1906:370). RESULTS Detailed information on intra-group patterning and distribution of OSS scores for Mariana Islanders is presented in Supplementary Data: Appendix Tables 5–9, while comparative data on the Oceanic and Pacific Rim groups we have studied are presented in tables below. Summary data on Mariana Islanders from isolated, locality, and regional sample types, dated mostly or exclusively to the Latte or Early Historic Periods (hereafter, “Latte Period”, will be used as a less cumbersome cultural period descriptor). In order to simplify comparisons and increase subsample sizes, scores for maximum degree of development at the three OSS sites (denoted XTOT, XPR, and XTSP) are featured in Supplementary Data: Appendix Tables 6–9. OSS frequencies for Latte Period Chamorros from three Mariana Islands are presented according to side (Supplementary Data: Appendix Table 5), Data: demographics (Supplementary Appendix Tables 6 and 7) and island (Supplementary Data: Appendix Table 8). Supplementary Data: Appendix Table 9 lists cases expressing co-variation of two or more marked expressions of OSSs, while Supplementary Data: Appendix Table 10 presents contrasting XTOT, XPR, and XTSP frequencies for Latte vs. Pre-Latte burials. Laterality of OSS expression in Latte Period Mariana Islanders Symmetry of development (L ¼ R) is the prevailing finding for TOT, PR, and TSP in both sexes. When expressed, asymmetry (L > R or R > L) is more common at the PR and TSP sites, especially among males (Supplementary Data: Appendix Table 5). Symmetrical expression at the TOT site is found in 95.7% of females and 90.3% of males, with each sex displaying a right side preference among the few cases with asymmetrical expression. Overall, sex differences are not statistically significant for TOT laterality. At the PR site, L ¼ R symmetry is found in 80% of females, but only 66.2% of males. Of the 19 asymmetrical females, greater development is found on the right side in 63.2% of cases, while 66.7% of asymmetrical males display greater expression on the left. Chi-square analysis found the latter sex difference significant (p  0.008). At the TSP site, highly significant (p < 0.001) differences in laterality is found between females and males, owing primarily to symmetry being much more frequent in females (81.4%) than males (58.1%). While males greatly exceed females in absolute frequency of asymmetry, right side preference is proportionately similar (61% and 55%) in females and males, respectively. OSS expression, by sex, in Latte Period Mariana Islanders All three traits are extremely sexually dimorphic, with highly significant (p < 0.001) differences in expression (Supplementary Data: Appendix Table 6). Marked expressions of XTOT, XPR, and XTSP are found in 2.5% (3/119), 1.8% (2/ 112), and 0.9% (1/110) of females, in comparison to 27.5% (42/153), 32% (50/156), and 15.8% (22/139) of males. OSS expression, by age cohorts, in Latte Period Mariana Islanders In contrast, association of OSS expression with age cohort is statistically insignificant among ancestral Chamorros (Supplementary Data: Appendix Table 7). Chi-square analysis was carried out after merging trait expression categories into not marked (0, 1, 2) vs. marked (3, 4) for males only, as few females had marked expressions of OSSs. In such merged cell analyses, tests of association found no THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 19 Gary M. Heathcote et al. statistically significant associations between dichotomous trait expression and age cohorts, for XTOT (p  0.92), XPR (p  0. 68), and XTSP (p  0.50). OSS expression, by island, in Latte Period Mariana Islanders OSS frequency breakdowns specific to Guam, Tinian, and Saipan, by sex, are given in Supplementary Data: Appendix Table 8. Examination of inter-island patterning is precluded for females, owing to the infrequency of marked expressions of OSSs. Males from Guam express marked developments at the TOT, PR, and TSP sites at frequencies of 31.0%, 35.0%, and 17.6%, while males from Tinian have similar frequencies of 29.4%, 33.4%, and 22.3%, respectively. In contrast, the sample from Saipan has a paucity of markedly expressed OSSs, with frequencies of 8.8%, 14.3%, and nil, respectively. Robust examination of inter-island patterning was precluded by small sample sizes for Tinian and Saipan, which rendered significance testing inappropriate due to cell sizes being <5 for 40 47% of male comparisons. OSS Co-variation in Latte Period Mariana Islanders Co-variation of markedly developed superstructures at two or more OSS sites is relatively common in ancestral Chamorros (Supplementary Data: Appendix Table 9), but examination of the patterning of such superstructure association is only permitted for males, given the paucity of marked OSS expression in females (Supplementary Data: Appendix Table 6). From a subset of 127 cases for which all three OSS could be scored, 38 (29.9%) express no markedly developed OSS, while 60 (47.2%) have marked developments at one, 19 (15.0%) at two, and 10 (7.9%) at all three OSS sites. Thus, 22.8% of ancestral Chamorro males display co-occurrence of at least two markedly developed OSSs. TOT–PR co-variation is the most common, occurring in 24/29 20 (82.8%) of these individuals. TOT–TSP and PR–TSP co-variation is far less frequent, occurring in 11/29 (37.9%) and 14/29 (48. 3%) of all co-varying cases, respectively. Nine (31%) of these cases express marked developments of all three OSSs. Marked co-variation exclusive to two sites is strikingly divergent, with 14 (48.3%) TOT–PR, but only one (3.4%) TOT–TSP, and four (13.8%) PR–TSP pairings. OSS expression in Latte Period vs. PreLatte Period Mariana Islanders Among Mariana Islanders, only Latte Period individuals—overwhelmingly, but not exclusively, male—express markedly developed TOTs and PRs. In contrast, TSPs are not exclusive to this more recent cohort, as two Pre-Latte individuals possess markedly developed TSPs. Among Latte Period males, prevalences of markedly developed TOTs, PRs, and TSPs are 27.5% (42/153), 32.1% (50/156), and 15.8% (22/ 139), while corresponding tallies for PreLatte males are nil (0/23), nil (0/13), and 15.4% (2/13), respectively (Supplementary Data: Appendix Table 10). These differences are highly significant (p < 0.001), significant (p  0.009), and insignificant (p  1.000) for TOT, PR, and TSP, respectively. Sum of Superstructures (SoS) expression in Oceanic and Pacific Rim groups Table 3 presents comparative measures of additive OSS development, Sum of Superstructures (SoS). SoS is calculated by summing each individual’s XTOT, XPR, and XTSP scores, which are categorized as Minimal (3), Medium (>3 < 7), or Strong (>6). For comparative purposes, skeletal series are merged into six regional groupings, contrasting Latte Period Mariana Islanders with samples from other regions of Remote Oceania, Near Oceania (Papua New Guinea), and the Asian, North American, and South American Pacific Rims (Table 1). VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin Table 3. Sum of Superstructures Rim Groupings2. Regional Groupings (SoS)1 Minimal Freq. % frequencies for Medium Freq. % Oceanic and Pacific Strong Freq. % Totals Females Mariana Islanders 90 86.5 13 12.4 1 1.0 104 Other Remote Oceania 11 91.7 1 8.3 0 0.0 12 Papua New Guinea 27 100.0 0 0.0 0 0.0 27 Asian Pacific Rim 6 100.0 0 0.0 0 0.0 6 North America Pacific Rim 37 80.4 9 19.6 0 0.0 46 South America Pacific Rim 0 0.0 3 100.0 0 0.0 3 Totals 171 26 1 198 Males Mariana Islanders 19 14.7 69 53.5 41 31.8 129 Other Remote Oceania 16 64.0 6 24.0 3 12.0 25 Papua New Guinea 24 82.8 2 6.9 3 10.3 29 Asian Pacific Rim 6 54.6 3 27.3 2 18.2 11 North America Pacific Rim 19 57.6 14 42.4 0 0.0 33 South America Pacific Rim 0 0.0 4 66.7 2 33.3 6 Totals 84 98 51 233 P 1 SoS categories ( XTOT þ XPR þ XTSP): Minimal 3; Medium >3 < 7; Strong >6. 2 Chi-square test of regional associations are inappropriate as 61% and 33% of cells are <5 for females and males, respectively. Controlling for sex, the Cochran-Mantel-Haenszel General Association statistic is highly significant (p  0.0001). Only one female with Strong SoS expression is known from our study, an ancestral Chamorro from Saipan (Heathcote et al. 2014:1012). We found that females generally express a Minimal degree of additive superstructure development, but Mariana Islanders, North American (San Francisco Bay), and South American (Mocha Island, Chile) Pacific Rim samples are exceptional. Among female ancestral Chamorros (n ¼ 104), 12.4% and 1% are categorized as Medium or Strong in expression, respectively, while 19.6% and 100% of North (n ¼ 46) and South (n ¼ 3) American Pacific Rim samples, respectively, exhibit Medium expression. In contrast, 51 instances of Strong SoS expression are recorded in males, ranging across the Pacific from locations along the Asian Pacific Rim (two Neolithic Taiwanese cases) to the South American Pacific Rim (two cases from Mocha Island). Oceanic cases consist of 3 from the Sepik Coast of Papua New Guinea (PNG), 41 from the Marianas, and 3 from more easterly regions of Remote Oceania (1 from the Marshall Islands and 2 from Tonga). Cross-tabulations show that ancestral Chamorros (n ¼ 129) and the small Mocha Island sample (n ¼ 6) are distinguished from other Oceanic and Pacific Rim groupings by their low Minimal (14.7% and nil) and high Strong (31.8% and 33.3%) SoS scores, respectively. Pooled samples from PNG (n ¼ 29) and the Asian Pacific Rim (APR) (n ¼ 11) yielded Strong SoS scores in an intermediate range, viz. 10.3% and 18.2%, respectively, with both APR cases being from the Neolithic Taiwanese subsample (n ¼ 5). Smaller non-Marianas samples render cells in a (sex) stratified 6  3 contingency table too sparsely populated for either an exact or Chi-square analysis, but a highly significant (p < 0.0001) general association between OSS expression and regional grouping was found by the CochranMantel-Haenszel test, controlling for sex. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 21 Gary M. Heathcote et al. Table 4. Non-marked vs. Marked1 frequencies of individual OSSs for Oceanic and Pacific Rim Groupings2. FEMALES Regional Groupings Non-marked Freq. % MALES Marked Freq. % Maximum Tubercle on Occipital Torus (XTOT) Mariana Islanders 116 97.5 3 Other Remote Oceania 12 100.0 0 Papua New Guinea 27 100.0 0 Asian Pacific Rim 8 80.0 2 North America Pacific Rim 46 97.9 1 South America Pacific Rim 3 100.0 0 TOTALS 212 6 Maximum Retromastoid Process (XPR) Mariana Islanders 110 98.2 2 Other Remote Oceania 12 100.0 0 Papua New Guinea 27 100.0 0 Asian Pacific Rim 7 100.0 0 North America Pacific Rim 46 100.0 0 South America Pacific Rim 3 100.0 0 TOTALS 205 2 Maximum Posterior Supramastoid Tubercle (XTSP) Mariana Islanders 109 99.1 1 Other Remote Oceania 12 100.0 0 Papua New Guinea 27 100.0 0 Asian Pacific Rim 6 100.0 0 North America Pacific Rim 46 100.0 0 South America Pacific Rim 3 100.0 0 TOTALS 204 1 Non-marked Freq. % Marked Freq. % 2.5 0.0 0.0 20.0 2.1 0.0 111 22 28 11 34 4 210 72.6 88.0 96.6 78.6 97.1 66.7 42 3 1 3 1 2 52 27.5 12.0 3.5 21.4 2.9 33.3 1.8 0.0 0.0 0.0 0.0 0.0 106 22 25 11 33 6 203 68.0 88.0 86.2 91.7 100.0 100.0 50 3 4 1 0 0 58 32.0 12.0 13.8 8.3 0.0 0.0 0.9 0.0 0.0 0.0 0.0 0.0 117 23 26 11 34 4 215 84.2 92.0 89.7 100.0 100.0 66.7 22 2 3 0 0 2 29 15.8 8.0 10.3 0.0 0.0 33.3 1 XTOT, XPR, and XTSP scores of 0, 1, and 2 ¼ Non-marked; scores of 3 and 4 ¼ Marked (Heathcote et al. 1996). 2 Controlling for sex, the Cochran-Mantel-Haenszel General Association statistic is significant (p  0.002), highly significant (p  0.0001) and significant (p  0.04) for XTOT, XPR, and XTSP, respectively. Maximum TOT, PR and TSP expression in Oceanic and Pacific Rim groups Table 4 presents inter-regional comparisons for individual OSSs separately, dichotomizing XTOT, XPR, and XTSP frequencies into non-marked (scores of 0, 1, and 2) vs. marked (scores of 3 and 4) categories. Among females, only a single North American Pacific Rim case from Ryan Mound, San Francisco Bay (n ¼ 48), three ancestral Chamorros (n ¼ 109–116), and 22 two Neolithic Taiwanese (n ¼ 5–8), included in the APR grouping, bear markedly developed OSSs at any of the three locations. Ancestral Chamorro females are distinguished by having three, two, and one individuals with markedly expressed XTOTs, XPRs, and an XTSP, respectively, while two Taiwanese and one Californian female expressed markedly developed XTOTs (only). Among males, 52 markedly expressed XTOTs are pan-Pacific in distribution, as VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin follows: 3 Neolithic Taiwanese (APR), 1 PNG villager (Sissano), 42 Mariana Islanders, 3 Other Remote Oceanic (ORO) cases (one each from Yap, Tonga, and Rapa Nui), 1 coastal Californian, and 2 Mocha Islanders. The 27.5% prevalence of markedly expressed XTOTs in ancestral Chamorros (n ¼ 111) is surpassed by the small sample of Mocha Islanders (33.3%) and approached by the APR grouping (21.4%). As all three of the latter cases are Neolithic Taiwanese (n ¼ 10), that subsample prevalence is 30%. In contrast, markedly expressed XPRs in males are more restricted in distribution. Absent from the Americas, they are presently documented only along the Asian Pacific Rim and into Near and Remote Oceania, as follows: 1 case from Taiwan, 4 from the West Sepik Coast of PNG, 50 from the Marianas, and 3 from ORO (all from Tonga). Ancestral Chamorros are predominant among regional groupings in expression of marked XPRs (32%), followed by the PNG (13.8%) and ORO (12%) groupings; but since all three ORO cases are Tongan, its subsample prevalence is 60%. Further diversity of geographic distribution is found for marked XTSP expression, recorded in Oceanic and South American Pacific Rim male samples only, but absent among the two Asian Pacific Rim or Californian samples. Of 29 cases recorded, 22 were found among Mariana Islanders, 2 ORO islanders (one each from Tonga and Marquesas), 3 PNG villagers, and 2 Mocha Islanders. Marked expression of TSPs in ancestral Chamorros (15.8%) is surpassed only by the small sample of Mocha Islanders (33.3%). DISCUSSION Synchronic Marianas Archipelago OSS Expression Despite substantial OSS database expansion, a truly robust examination of intra-archipelago spatial patterning of Latte Period peoples awaits further amendments. In Guam, sparseness of data from sites and localities other than along Tumon Bay (Figure 3) inhibits examination of interregional or coastal-inland intra-island differences, and such paucity of data precludes biohistorical examination of early historic (Spanish mission) accounts, which make reference to manachang, lower caste groups who occupied inland and riverine locations. Writers of these documents suggest that the manachang were culturally and biologically distinctive, which would occur if they were socially and reproductively isolated from higher caste chamorri coastal occupants. It has been further advanced that the manachang might represent descendants of the original (PreLatte) people of Guam who were conquered and displaced by Latte Period ancestors of the chamorri (see Tolentino 2015 and references).14 Examination of inter-island OSS patterning is likewise handicapped by the small samples representing Saipan and Tinian, not to mention the absence of systematically produced data for Rota and Gani, the small islands of the Mariana Archipelago north of Saipan (Russell 1998a) (see Figure 3). These caveats aside, we note that the apparent higher prevalence and greater strength of expression of OSSs in Guam and Tinian vs. Saipan (Supplementary Data: Appendix Table 8) could reflect inter-island demographic interaction spheres—and attendant biobehavioral distinctions—during the Latte Period.15 Synchronic Marianas Archipelago OSS Co-variation (and Comparative Data) Ancestral Chamorros are most distinctive from other Oceanic populations in OSS co-variation patterning. Among males, covariation of two or more strongly expressed OSSs is a frequently encountered morphological complex, occurring in 22.8% of this cohort. Among these cases (see Supplementary Data: Appendix Table 9), 82.8% exhibit marked TOT expression co-varying with marked PR expression, THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 23 Gary M. Heathcote et al. suggesting quite frequent concordant use of the upper trapezius and superior oblique muscles. Co-variation of strongly expressed TOT with TSP and PR with TSP were less frequent but still substantial at 37.9% and 48.3%, respectively. Among this cohort with multiple markedly developed OSSs, 31% exhibit such expression at all three sites. Thus, TOT, PR, and TSP appear to be part of a morphogenetic and remodeling complex in which differential expression and co-variation of these OSSs reflects variable life histories—possibly related to longstanding roles in latte construction16—in combination with distinctive genomes. Of the non-Marianas crania from Oceania and the Pacific Rims, only five males manifest markedly developed superstructures at two or more OSS sites (Supplementary Data: Appendix Table 11)—one from Pangaimotu, Tonga (Sava 1996; Sava and Pietrusewsky 1995), two from Sissano, one from Warapu, along the West Sepik Coast, PNG (Douglas and Stodder 2010), and one from Mocha Island, Chile (Matisoo-Smith and Ram
ırez 2010). Within this residual grouping, TOT–PR covariation is not dominant as with Mariana Islanders. Small sample sizes disallow significance testing, but this stark difference suggests that a distinctive, if not unique, set of chronic activity patterns triggered the especially frequent marked TOT–PR co-variation among ancestral Chamorros. Diachronic Marianas Archipelago OSS Expression Comparisons While marked TOTs and PRs are found only in Latte Period burials, markedly expressed TSP cases are recorded in two Pre-Latte individuals (Supplementary Data: Appendix Table 10). The exclusivity of markedly developed TOTs and PRs among latte builders supports our working hypothesis on work-related mechanical induction of OSSs. These actions would be involved in latte house (guma latte) construction activities involving megalithic transport and emplacement, such as pulling on ropes to move sleds and bipods and 24 operating hoists, and using carrying poles in the yoke position (Heathcote et al. 2012b:53–54). In contrast, the presence of tuberculated TSPs in both Pre-Latte and Latte individuals suggests that chronic work activities not exclusive to (or perhaps other than) those involved in guma latte construction were involved in their mechanical induction. From these findings, we propose that a suite of consequential motor behaviors involving the TSP-related sternocleidomastoid muscle (SCM) was continuous from Pre-Latte through Latte Period times. As actions of this muscle include drawing the neck forward, lifting the neck when supine, raising the chest in forced breathing, and rotating and tilting the neck towards the shoulder (Heathcote et al. 2012b:54), we propose that participation in long-standing, long-distance seafaring traditions—from Pre-Latte through Latte times—could account for the marked expression of TSPs among individuals whose personal histories transcended cultural periods before and after the advent of latte construction and associated activities.17 Oceania and Pacific Rim Comparisons Latte Period Mariana Islanders have extremely high cumulative prevalences of markedly developed OSSs (SoS) compared to most populations sampled from Oceania and the Pacific Rim (Table 3). The only female known to us with Strong SoS expression is an ancestral Chamorro, and among 129 male ancestral Chamorros (for whom all three OSSs could be scored), Strong SoS is exhibited by 31.8% and Minimal by only 14.7%. Only the small sample from Mocha Island is Chamorrolike, with 33.3% Strong and nil Minimal SoS. The relatively well-sampled Mariana Islands are, thus, a demonstrated center of accumulative OSS expression. But pockets of Strong SoS expression recorded elsewhere—in small samples from Taiwan, PNG, Tonga, and Mocha Island and in a single individual from the Marquesas (Table VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin 3)—document pan-Pacific morphological affinities that, we propose, attest to population historical connections and shared motor behaviors. Ancestral Chamorros are somewhat more morphologically distinctive when examining inter-group differences of marked (“3” and “4”) vs. non-marked (“2” and less) maximum scores for each of the three individual OSSs (Table 4). Among females, markedly developed XPRs and XTSPs are exhibited only by ancestral Chamorros (two and one, respectively). This exclusivity does not apply to XTOTs, as two Neolithic Taiwanese and one indigenous coastal Californian join three ancestral Chamorro females in displaying veritable tubercles on their occipital tori. Among males, ancestral Chamorros display a high frequency of markedly expressed XTOTs (27.5%) surpassed only by two small Pacific Rim groups, viz. the Neolithic Taiwanese (30%) APR subsample and the South American sample from Mocha Island (33.3%). In contrast, ancestral Chamorro males have greater than twice the prevalence of markedly developed XPRs (32%) vs. other groupings in Table 4, though the small ORO Tongan subsample surpasses them, with 3/5 (60%) exhibiting marked XPRs. Finally, ancestral Chamorro male prevalence for markedly developed XTSPs (15.8%), while relatively high, is surpassed by Mocha Islanders (33.3%), Tongans (20%), and the single Marquesan (ORO) individual. Broad generalizations about interregional OSS patterning must be tempered with small sample size caveats, but a fuller picture of the geography of OSS expression is emerging. Marked expressions of XTOTs are now documented across the Pacific Basin for Neolithic Taiwanese, a PNG villager, Mariana Islanders, a Yapese, a Tongan, a Rapa Nui islander, a coastal Californian, and Mocha Islanders. Marked XPRs are known for crania from Taiwan, PNG, the Marianas, and Tonga, but are absent in North and South American Pacific Rim samples. Finally, markedly developed XTSPs are now known for Oceanic groups (PNG villagers, Mariana Islanders, a Tongan, and a Marquesan) and Mocha Islanders off the coast of South America, but not for Asian Pacific Rim or coastal Californian samples. Such diversity of distribution patterning of individual OSSs may signal a shared propensity to develop and grow these superstructures, interacting with different sets of mechanical induction demands related to motor activities involved in transportation, subsistence, and building technologies. Overview of Spatial and Temporal Patterning of OSSs Over 40 years ago, Hublin (1978c) noted that modern and fossil hominins from Oceania commonly exhibit occipital morphologies that are generally distinctive from those of African and European hominins. Modern Australians, Melanesians, and certain East Asians—compared to modern Afro-Europeans—were reported to have more strongly developed PRs, TSPs, and APs (conceptualized by Hublin as non-homologous to TSPs), and sometimes possessed supranuchal tubercles (TOTs) (Hublin 1978c:175). Our survey findings (Tables 3 and 4) extend the geographic parameters of these OSSs in AMHs to include the geographic regions of Micronesia and Polynesia, as well as the Asian and American Pacific Rims. In syntheses below about the geography of PR, TSP, and TOT expression, temporal depth and regional continuity are included. Tubercles on the Occipital Torus (TOT) Expression Like the external occipital protuberance (Hublin 1978c:171), TOT appears to represent an AMH apomorphy, albeit with a circumscribed geographic distribution. The presence of markedly developed TOTs in Late Pleistocene crania from Okinawa (Suzuki 1982) and in the APR subsample of Neolithic Taiwanese reported here suggests that TOTs represent a long-standing regional morph in coastal East Asia, that later spread to Near and Remote Oceania, then onward to the American Pacific Rims. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 25 Gary M. Heathcote et al. Regarding recent AMHs, with the exception of studies by Hublin (1978c) and Lahr (1995, 1996), we cannot determine what proportion (if any) of non-Oceanic skulls described as having “strong” OTs bear tuberculated developments (marked TOTs) at the upper trapezius attachment site. Hublin (1978c:44) identifies only one case of supranuchal tubercles in a cranium not from the Pacific region, a Greenlandic Inuit, but describes these tubercles as “weak or atypical”. Lahr (1995, 1996) reports finding Tubercles of Hasebe in small numbers of ‘East Asians, “Eskimos,” Australians’, and Sub-Saharan Africans, but illustrations of her “OT5” grade (Lahr 1996:350–351), said to be defined by “Tubercles of Hasebe” on the OT, show a lesser degree of expression than what we consider to be (minimally) marked (score of “3”). As well, Lahr’s “OT6” or “OT7” scores do not correspond to what we consider marked grades of TOT expression. Aside from our documentation of markedly developed TOTs among Mariana Islander, ORO and PNG groupings (Table 4), the only studies of AMHs from Remote and Near Oceania that explicitly mention Tubercles of Hasebe/supranuchal tubercles are those of Hublin (1978c) and Lahr (1994, 1995, 1996). Morphological equivalence of Lahr’s cases with ours is unclear, but Hublin’s (1978c) “strong and typical” supranuchal tubercles in crania from New Caledonia and Australia–Tasmania appear to be equivalent to our marked TOTs. Regarding native North Americans, we documented two cases (one female; one male) of tuberculated TOTs from the Ryan Mound site, San Francisco Bay, on the California Pacific Rim (Table 4). While early studies of aboriginal Americans (e.g., Hagen 1880; Matiegka 1906) reported relatively high frequencies of OT—including cases described as having strong degrees of development—none provide documentation of TOTs (Supplementary Data: Appendix Table 1).18 For South America, the situation is unclear, aside from the Mocha Islanders (Table 4). Marta Lahr (personal communication 1996) reported two “well-developed” cases with Tubercles of 26 Hasebe among an exceptionally robust group (n ¼ 29) of Fuegian-Patagonian crania, but in the absence of photographs we cannot ascertain concordance with what we score as marked TOTs.19 If Lahr’s two Fuegian-Patagonian cases do indeed bear tuberculated TOTs, the geographic circumscription of marked expressions of this superstructure would include southernmost continental South America,20 in addition to Australia, Near and Remote Oceania, and the North and South American Pacific Rims. Retromastoid Process (PR) Expression The earliest known non-trivial development of the retromastoid process is found in KNM-ER3733, an Early Pleistocene East African Homo erectus individual. We are not aware of such PR expression in approximate coevals elsewhere, nor in later archaic hominins from Africa, Europe, or Northeast Asia, but strongly developed PRs are commonly encountered in fossil hominins from Java, an epicenter of early expression and intra-regional temporal continuity (Supplementary Data: Appendix Table 4). Among AMH samples and subsamples from Oceania, marked degrees of PR development are expressed by 32% of ancestral Chamorro males (and 1.8% of females) (Supplementary Data: Appendix Table 6), 50% of Tongan males, and 13.3% and 7.4% of PNG males from the West Sepik Coast, respectively. Ancestral Chamorro males further align with Tongan and PNG males in the frequent co-variation of marked PRs and TSPs (Supplementary Data: Appendix Tables 9 and 11). The only non-Oceanic samples known to have non-trivial frequencies of “strong” PRs—excluding Waldeyer’s (1909) inflated frequency data—are both East Asian, namely a large sample of Japanese (2.8%) studied by Hublin (1978c) (Supplementary Data: Appendix Table 2), and the small subsample of Neolithic Taiwanese males (11.1%) included in our APR grouping (Table 4). Thus, the spatiotemporal distribution of markedly VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin developed PRs extends from later Pleistocene Indonesia to the northerly reaches of the Asian Pacific Rim, and later to Near and Remote Oceania.21 Posterior Supramastoid Tubercle (TSP) Expression The only fossil hominin known to us that exhibits an unmistakable TSP is Ngandong 1 (Hublin 1978c), a Middle to Late Pleistocene individual from Java. All other known cases occur in anatomically modern humans. Among previous researchers who have considered the Processus asteriacus (AP)/TSP in AMHs, only Jacob (1967) compiled frequencies. He studied three non-Pacific and one ISEA (Central Java) skeletal series (Supplementary Data: Appendix Table 3), but encountered no cases of markedly developed TSPs as we define them. Matiegka (1906) stands alone in describing tuberculated TSPs in AMHs unambiguously, as “button-like” APs in one skull from Fiji, and in one morphologically anomalous case from southern Bohemia. His review is otherwise diminished by anatomical ambiguity, leaving us unsure of whether attributed APs equate with tuberculated peri-asterionic processes or merely crests and ridges along the lateral third of the lambdoidal suture. We have established that the relatively well-sampled Mariana Islands were a center of marked TSP expression during the Latte Period and that, moreover, other samples from Remote Oceania, as well as PNG and the South American Pacific Rim22 approach or exceed ancestral Chamorros in their prevalence of marked TSPs (Table 4). We propose that such morphological continuity is evidence of phyletic connections, together with similar motor behaviors, across this region.23 CONCLUSIONS The present study documents a widespread albeit patchy distribution of markedly developed occipital superstructures across the Pacific Basin, spanning sites, locales, and regions from Australia, the Asian Pacific Rim, Near Oceania, Remote Oceania, and the American Pacific Rims. We propose that while ancestral Chamorros are morphologically distinctive in their multiple OSS co-variational patterning, historical connections to pan-Pacific populations are best attested to by a panPacific trail of individual markedly developed OSSs, namely tuberculated TOTs, PRs, and TSPs. Collectively, the geographic patterning of these three OSSs has fuzzy congruence with models and scenarios— based on archaeological, linguistic, and genetic evidence—of the migratory history of the Pacific,24 and contact with the Americas (Jones et al. 2011; Kirch 2010; Ram
ırez-Aliaga 2016). For needed temporal depth consideration, the bioarchaeology of Southeast Asia was reviewed, in order to consider ancestral contributions of archaic hominins to the first AMH settlers of Sunda and Sahul (see Tayles and Oxenham 2006).25 Regarding the widespread distribution of markedly expressed PRs and TSPs among AMHs across the Pacific Basin, coupled with the geographically restricted distribution of homologous superstructures in earlier hominins, such morphological continuity appears to reflect genetic continuity with Indonesian Homo erectus populations present in Southeast Asia by 1.6 MYBP (Oxenham and Buckley 2016).26 In contrast, markedly developed TOTs are first documented in Late Pleistocene crania from Okinawa; their later presence in Neolithic Taiwanese suggests a northeast Asian AMH origin and continuity for this OSS. We propose that a shared constitutional propensity for developing markedly expressed OSSs among both closely and more distantly related Pacific Basin populations reflects an ultimate genetic cause (and phylogenetic signal), and that culturally conditioned chronic motor behaviors account for penultimate induction of these superstructures among a varying minority of individuals in each population. Multiple lines of evidence support such penultimate THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 27 Gary M. Heathcote et al. mechanical induction. First, focusing on the only well-sampled population, ancestral Chamorros, strong sexual dimorphism is displayed as well as frequent co-variation of two or more marked OSSs. As these superstructures are anatomically related to entheses of posterior cranial muscles and their degrees of expression are associated with humeral robusticity, it is reasonable to infer that differential economic role activities, structured by gender and occupational specialty, triggered OSS development in a significant minority of genetically predisposed individuals. A strict activity-induction model of OSS development is not supported, however, for it would require congruent, overlapping geographic circumscriptions of markedly developed OSSs with motor behaviors implicated in their genesis (Heathcote et al. 2012b); and such an association (e.g., OSSs with megalith-building and/or traditional seafaring) simply does not exist at the global level. Further etiological complexity is indicated by SEM evidence of a systemic effect in OSS morphogenesis (Heathcote et al. 2014). Because of the apparent complex multifactorial etiology of OSSs, future inquiry into their multiple meanings will require contributions from many research fields. Phenomic and pleiotropic research (Freimer and Sabatti 2003; Karasik and Kiel 2010) would be ideal, including examination of genetic-morphological correspondences and developing and testing models of gene-environment (including mechanical) interactions. But such research is unlikely to receive funding, due to the nonclinical nature of OSSs. More realistically, future research can be carried out on the phylogenetic and osteobiographical signals of OSSs by proxy, through continued research into the origins and history of Pacific Islanders via a combination of ancient DNA (aDNA) and morphological studies. Guided by the dictum that studying the remains of ancestors directly, and not the biodiversity of their descendents, is the optimal way of addressing questions of ancestry (Matisoo-Smith 2011), aDNA research should be prioritized in all future 28 studies of skeletal remains. Regarding preLatte vs. Latte Period Mariana Islanders, blinded aDNA testing of skeletal and dental remains, according to each individual’s degrees of OSS expression, is essential to addressing questions about the role of megalithic building activities in triggering and mediating the expression of these superstructures. Presently, much of what we think we know about Chamorro origins and affinities is inferred from DNA testing of contemporary Chamorros. This is problematic, since ancestral Chamorros were likely more genetically diverse than modern descendants who are—after all—linearly related to survivors of severe population bottlenecks during Spanish Colonial times.5 Indeed, ancestral Chamorros may have possessed DNA haplogroups that are not represented among contemporary Chamorros, owing to diminution or extinction of lineages.27 As for future morphological research, databases on comparative cranial morphology need to be integrated with each individual’s total inventories of skeletal remains. This would facilitate, for example, case-matched study of the correlates and associations of OSS expression with infracranial metric, nonmetric, and entheseal (muscle marker) information relating to infracranial robusticity, muscularity, and inferred motor behaviors across different cultural and geographic landscapes. For the Marianas, such associative database expansion would allow for patterns of OSS variation and co-variation to be explored and interpreted according to island, region, interaction sphere, archaeological context, and temporality. Such associative morphological inquiry, if integrated with expanded and case controlled aDNA sampling, would allow for testing the efficacy of such exploratory OSS systematics to contribute to the construction of osteobiographical profiles and population histories of Oceanic peoples. Until that time, our proposal that markedly developed OSSs are morphological markers of ancestral and collateral genetic connections across the Pacific Basin, as well as indicators of similar VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin motor behaviors, is merely grounded in strong inference. Theoretically, this proposal aligns with the suggestion of Rogers and colleagues (1997) that there are differences between individuals and populations in the ability to form hyperostotic bone at entheses in response to mechanical stress, and that these differences (“bone formers” vs. others) relate to genetic variation within and between populations. Milford Wolpoff, and Yigal Zan. JeanJacques Hublin (Max Planck Institute for Evolutionary Anthropology) deserves special thanks for providing guidance, encouragement and resources for the beginning phases of this project. Shortcomings of this paper are ours alone. ACKNOWLEDGEMENTS GH’s research was supported by load allocations from the Dean of the College of Liberal Arts and Social Sciences, University of Guam. MP’s research involving Neolithic skeletons from the Tainan Science Park was supported by The National Science Council (Taiwan), and EA M-S and J-M R-A’s ongoing project “Redrawing the Polynesian Triangle: Did Polynesian settlement extend to South America?” is supported by UOO0926. The late DBH’s research was funded by a grant from The National Institute of Environmental Health Sciences, ES05064. For support and assistance at museum and laboratory work sites, we thank Jaymie Brauer, Susan Rodriguez, and Ian Tattersall (American Museum of Natural History); Toni Han and Lisa Armstrong (B.P. Bishop Museum); Glen Cole and John Terrell (Field Museum of Natural History); David Hunt, Carol Butler, Douglas Ubelaker, and the late Donald Ortner (Smithsonian Institution); Philippe Mennecier (Mus
ee de l’ Homme); and Alan Haun, Diane Trembly, Sue Goodfellow, and David DeFant (Paul H. Rosendahl Ph.D., Inc.). For German and French translation assistance, we thank Jurgen CarsonGreffe, Jean-Jacques Hublin, Marion Reyes, Peter Schuup, and Susanne Wilkins. Information retrieval assistance was provided by Chris Bellessis, Arlene Cohen, Joanne Tarpley Crotts, and Moses Francisco. Map design and production was provided by Marween Yagin, Graphics Specialist (Center for Instructional Support, University of Hawaii at Manoa). For information sharing and/or critical comments on drafts of this paper, we thank three anonymous reviewers, C. Loring Brace, Frank Camacho, Sara Collins, Vincent Diego, David Frayer, Hermann Helmuth, the late W. W. Howells, Rosalind HunterAnderson, Marilyn Knudson, Christopher € Knusel, Marta Lahr, Koji Lum, Alan Mann, Chris Meiklejohn, Janet Monge, Darlene Moore, Walter Neves, the late Nancy Ossenberg, Phillip Rightmire, Dirk Spennemann, Mary Torcat, Alan Walker, FUNDING NOTES 1. Apart from the high prevalence of markedly developed OSSs and frequent co-variation of two or more of same, ancestral Chamorros are also morphologically distinctive with regard to humeral robusticity and inferred upper body muscularity and strength. Based on a comparative study of a humeral shaft robusticity index (HSRI2)—which expresses the shaft’s minimum plus maximum diameters at midshaft as a proportion of maximum length—ancestral Chamorros are close to the extreme upper end of known variation for AMH and fossil hominins (see Heathcote et al. 2012a:157–168, 205–207; 2012b:55–59). 2. As examples, red-slipped pottery known as Marianas Red, derived from ancestral pottery traditions in ISEA, is unique to the four largest Mariana Islands and first appears around 3500 BP (Carson et al. 2013; Moore 2002) or somewhat later (Rieth and THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 29 Gary M. Heathcote et al. Athens 2017). Much later, prehistoric cultivated rice (Oryza sativa L.) and latte stone architecture become distinctive cultural features of life in the Marianas. Within Remote Oceania, prehistoric cultivated rice is unique to the Marianas, while stone latte structures are unique globally. Both domesticated rice and latte are first documented around 1000 BP (Hunter-Anderson et al. 1995), though it appears that rice was not added to the crop inventory of ancestral Chamorros to a significant degree until 400 years later, with the advent of the Little Ice Age (600 BP) (Hunter-Anderson 2012). 3. The mutually unintelligible Chamorro and Palauan languages belong to the Western Malayo-Polynesian (WMP) subgroup of the vast Austronesian family, and stand apart from the more than 450 other Pacific region languages in not being part of the Oceanic subgroup (Blust 2000). Historical linguistic relationships and branch classifications are contentious, with most formulations fitting into one of four claims (Reid 2014), viz. that Chamorro is (a) a Philippine-type language, probably most closely related to Ilokano and Tagalog (e.g., Topping and Dungca 1973); (b) most closely related to certain languages in Indonesia (e.g., Zobel 2002); (c) most closely related to some Austronesian languages in Taiwan (see Starosta 1995); or (d) descended from a protolanguage ancestral to all Austronesian languages outside of Taiwan, and not closely related to any other WMP language (see Blust 2000). Reid (2002) extends Blust’s (2000) hypothesis, contending that ancestral Chamorros spoke an Austronesian language that developed in the northern Philippines following the first migration out of Taiwan. Lack of close synchronic syntactic and phonological relationship of Chamorro to Philippine languages is interpreted to mean that Chamorro out-migrants left the Philippines before ProtoMalayo-Polynesian differentiated into dialects that later evolved into the Philippine languages of today. Zobel’s (2002) view of the linguistic position of Chamorro is in closest agreement 30 with genetic findings (Vilar et al. 2013a), viz. that Chamorro (and Palauan) are outliers within the putative Nuclear Malayo-Polynesian branch of the Austronesian family, which includes most of the languages of Sulawesi and the Greater Sunda Islands. Further, Zobel speculated that ancestors of Chamorro speakers reached the Marianas from Sulawesi. Blust (2000) and Reid (2002) argue that there is no phonological or syntactic support for this view. Virtually all formulations agree that Chamorro (like Palauan) is an outlier language that does not share a phylogenetic subgrouping, let alone mutual intelligibility, with extant Austronesian languages, due to its evolution in relative isolation from other languages deriving from ProtoMalayo Polynesian (Bellwood 1991). 4. Chamorro distinctiveness, biological, cultural, and linguistic, surely relates to the remoteness of the Marianas archipelago, separated from various proposed ISEA homelands by more than 2000 km (Petchey et al. 2016; see also Fitzpatrick and Callaghan 2013), perhaps the longest expanse of open sea traversed anywhere in the world (Craib 1999). From where the first visitors/settlers to the Marianas originated is a matter of intense debate. The northern Philippines was recently proposed by Hung et al. based on archaeological (2011), (ceramic), linguistic, and genetic considerations. Countering this proposal, Winter et al. (2012) made a case for origins further south in ISEA, based on ceramics, linguistics, and oceanographic considerations. More recently, Fitzpatrick and Callaghan (2013) employed seafaring simulation models and concluded that the Marianas were most likely settled from Halmahera, the Bismarcks, or northern New Guinea. Most recently, Montenegro et al. (2016) situated the Solomons and New Guinea as most likely homelands of the earliest visitors/settlers to the Marianas, based on a seafaring simulation study that employed shortest-hop trajectory data. 5. The latter include a series of depopulation events, beginning with VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin the late seventeenth-century SpanishChamorro wars and subsequent forced population relocations (Shell 2001; Underwood 1973) that set the stage for a severe mid-eighteenth-century population bottleneck. The total Marianas population of Chamorros, just before the first Jesuit mission in Guam (282 BP, i.e., AD 1668), is estimated to range from 24,000 to 100,000 (Shell 1999; Underwood 1973). By the time of the first Spanish census in 240 BP (i.e., AD 1710), the native population in Guam and Rota declined to under 4,000 (Shell 1999) and further to 1,700 by 192 BP (i.e., AD 1758), perhaps its lowest point in the post-contact history of the Marianas (Goetzfridt 2014). Besides direct and indirect mortality from the Spanish-Chamorro wars, the precipitous depopulation of the Marianas was due to introduced infectious disease epidemics and typhoons (Garruto 2012; Plato and Cruz 1967; Reiff et al. 2011), complicated and exacerbated by malnutrition, starvation, social disruption, and emotional trauma 2014; Leon-Guerrero (Goetzfridt 2015). Within this context of such social upheaval, birth rates would have been negatively impacted. While important in explaining the population structure of contemporary Chamorros, the impact of the eighteenth-century population bottleneck is moot to this study, for few (if any) of the ancestral Chamorros studied lived more recently than 250 BP (i.e., AD 1700) (Table 1). 6. While Valentin and colleagues (2005) referenced our protocol in describing OSSs in a single burial (dated to ̴ 1800–2400 BP) from Mangaliliu village in Vanuatu, no individual OSS scores were provided. The superstructures were described collectively as weakly developed, presumably <“2” according to our scoring scheme. 7. In a study of cranial variation across nine regional groupings, Lahr (1996) provided bar graphs indicating that strongly expressed occipital tori (scores of 5–7, according to her system) are most frequent in Fuegian–Patagonians, followed by Australians, then East Asians. She also recorded a zero frequency of pronounced OT samples of Europeans and Middle Easterners, and small prevalences in Sub-Saharan Africans, Southeast Asians, and North Africans. Lahr’s (1996) data are not included in Supplementary Data: Appendix Table 1, as frequencies and prevalences were not provided. 8. Regarding Middle Pleistocene European and East Africans, Hublin (1978c:138) described “trace” PR expressions on the occipital bone from Bilzingsleben (Germany), and Rightmire (1990:210, 217) identified “small, poorly defined” PR in the Petralona (Greece) cranium and a “low relief” outline of the superior oblique muscles in the Lake Ndutu, (Tanzania) cranium. The Spy 2 (Belgium) cranium is said to be distinguished from other Neandertal males in having a “distinct retromastoid process”, while the early modern Mladec 5 (Moravia) cranium is described as bearing a “prominent retromastoid process” (Frayer et al. 2006:232). Magnified images of Spy 2 and Mladec 5 (Wolpoff et al. 2001) reveal their PRs to be slight and moderate in development, respectively, using our protocol. 9. The poorly dated Sangiran individuals may range from Early to Middle Pleistocene, and probably antedate the late surviving H. erectus groups at Ngandong and Sambungmachan (who lived at least 100,000 BP and possibly <50,000 BP) by several hundred thousand years (Anton 2003; Bartstra et al. 1988; Swisher et al. 1996; see also Indriati et al. 2011). 10. This proposal finds support from Anton (2003; see also Baab 2011), whose work on morphological distinctions between Chinese and Indonesia H. erectus crania led her to conclude that intermittent isolation produced and maintained regional morphs in Southeast Asia. 11. This consistent discrepancy suggests that Waldeyer’s (1909) “weak” PR scores are equivalent to what others considered absent (see Supplementary Data: Appendix Table 2). Overreporting is further suggested by his THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 31 Gary M. Heathcote et al. claim of moderate total frequencies (14.3% and 20%) in two European series, in contrast to Le Double and Dubrueuil-Chambardel (1905) and Michelsson (1911), who reported low frequencies, ranging from 1.3% to 1. 4%, for three large European series. Waldeyer’s PR frequency ranges of 3. 7% to 27.6% for Asian and Native American groups are also exceptional, in contrast to ranges of nil to 4.7% for three Asian/New World groups reported by Michelsson (1911) and, later, Hublin (1978c). 12. Jacob reported that “grade 3” TSPs were not found in male or female African Americans and Central Javanese, nor among female Alaskan “Eskimos” and Central Europeans. Indeed, only five observed sides of such TSP expressions were recorded, in 3.5% (3/86) of Alaskan “Eskimo” and 4.3% (2/46) of Central European males. 13. For example, Matiegka’s sometimes interchangeable use of AP and the German term for “asterial crest” is problematic, as the latter refer to folded bulges along the lambdoidal suture near asterion, features that we would score as an incipient, slight, or moderate TSP, at most. Drawing on his own research, Matiegka noted that Lapp, Samoyed, Chinese, Malayan, Javanese, Makassarese (southern Sulawesi), “Negro,” Kabyle, (northeast Algerian Berbers), and Egyptian mummy samples lacked APs, as well as noticeable ridges or crests in the asterionic region. In contrast, AP and/or asterial crests and ridges were reported for Native North and South Americans (e.g., Paiutes, and skulls from Florida, Santa Rosa Island [California], Chile, the Argentine pampas, and Tierra del Fuego). He reported a strongly developed asterial crest in 25% (4/16) of aboriginal Santa Rosa Islanders, a generally less developed crest in 10% of a larger cranial series from Bohemia and weakly developed processes in a small series of three Greenlandic Inuit (Matiegka 1906: 368–369). 14. Interpretation of the archaeological record bearing on the peopling of the Mariana Islands is contentious. For 32 example, Carson (2014:74) maintains that Early Unai archaeological sites provide evidence of a “formal population migration” to the Marianas, while Hunter-Anderson (2013) interprets these earliest Pre-Latte sites as bearing witness to temporary visitations by seasonal marine foragers, not settlers. Colonization was surely underway by Middle Unai times, however, as witnessed by recovery of 177 early human burials from the Naton Beach site in Guam. Combined calibrated AMS 14C age (expressed in 2r) for Conus shell bead necklaces from four of these individuals ranges from 2740 to 2280 BP (i.e., from Middle Unai to Late Unai Periods [DeFant 2008]). Beyond the debates about when the islands were settled is the question of in situ continuity and adaptive change vs. population replacement vs. intermediate models combining continuity with external influences from Pre-Latte through Latte Period times (see Hunter-Anderson and Butler 1995:28; Rainbird 2004:131–132; Vilar et al. 2013a). Partitioning the present Mariana Islands samples into Pre-Latte and Latte analytic units informs debate on settlement history (see Walth 2014), but cannot lead to resolution because interpretation of morphological differences between Pre-Latte vs. Latte samples is confounded by motor behavioral changes associated with the onset of latte house building and proposed physiological plastic skeletal responses to motor activities involved (see Heathcote et al. 2012b:54). Regardless of the degree to which PreLatte populations were ancestral to Latte Period Chamorros, the invention, development, and diversification of work involved in latte house construction complicates interpretation of Chamorro ethnogenesis, assuming the correctness of our multivariate model of OSS development, which includes mechanical induction as a co-factor. VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin 15. Further, the presence of latte construction specialists, unevenly distributed throughout the Marianas, cannot be ruled out. While undocumented for the Marianas, clans of specialist builders are known for Kiribati (Hockings 1989) in eastern Micronesia, a similar semi-sedentary society with an incipient political hierarchy (see Heathcote et al. 2012b:60–61). Alternatively, we cannot rule out that our pooled Saipan sample might be non-representative. Earlier studies (Hasebe 1935; Schlaginhaufen 1906) offer little help, for while markedly developed TOTs and TSPs of Saipanese Chamorro male crania were illustrated, frequencies were not reported. 16. An outline of proposed muscle actions related to latte construction, including attendant megalith transport and placement, is presented in Heathcote et al. (2012b:54). The actions of the TOT-associated upper trapezius include helping with neck extension and bending the neck from side to side, drawing the clavicle and scapula backwards, elevation and rotation of the scapula, and supporting the clavicle and scapula when heavy weights are born by hands when arms are down at the side. The superior oblique (suboccipital), anatomically related to the PR, bends the neck backwards and rotates it from side to side. The TSPassociated sternocleidomastoid draws the neck forward, raises the neck while supine, raises the chest in forced breathing, tilts the neck toward the shoulder, and rotates the neck. 17. While the historical record is mute on the physical labors involved in operating traditional Chamorro sailing canoes (proas), Gladwin (1970) provides such an account for Puluwatese crew members on a traditional outrigger sailing canoe. Puluwatese canoes are good proxies, due to design similarities to Chamorro proas (Hornell 1936). Thus, the work involved in maneuvering them must also be similar. Among Puluwatese navigators, routine labors include paddling, hoisting the sail, bailing water, steering with a big steering paddle, and adjusting the sheet (sail). The most rigorous work occurs when 18. 19. 20. wind changes necessitate tacking, when the sail is moved from one end of the canoe to the other. Illustrations of this process Gladwin (1970:104–105) show 11 steps from releasing the tack line at one end of the canoe to securing it to the end stay at the other end. Crew members engage in rig carrying, heaving, and pulling movements. A movement that especially loads the head and neck dynamically is overhead rig heaving with the head and neck forwardly flexed. Thus, sailing canoe voyaging would involve the SCM in ways with the potential to transform the TSP site (in predisposed individuals). Unpublished images of aboriginal Northwest Coast skulls from British Columbia, provided by Christopher Kn€ usel (Universit
e de Bordeaux), illustrate quite robust crania, but with only moderate (non-tuberculated) developments at the TOT (and PR) site. Unpublished images of two Fuegian Haush (courtesy of Walter Neves, University of S~ ao Paulo, Brazil) and published illustrations of FuegianPatagonian skulls (Hrdlicka et al. 1912:plate 38b) reveal that while one Patagonian and two Haush skulls bear exceptionally robust OTs, none bear TOTs. Should systematic surveys verify the latter, such a distribution would contribute to the debate on the peopling of the Americas. Intriguingly, Neves et al. (1999a) have demonstrated morphometric affinities between the Haush and three other Tierra del Fuegian groups with two Polynesian groups (Mokapu, Hawaii, and Moriori from the Chatham Islands) accessed global from Howells’s (1989) craniometric database. While outside the purview of this paper, Lahr (1995), Munford et al. (1995), Hernandez et al. (1997), Neves et al. (1999a, 1999b), Matisoo-Smith and Ram
ırez (2010), and Ram
ırez-Aliaga (2010, 2011), among others, provide alternative interpretations of distinctive FuegianPatagonian craniofacial morphology and the wider issue of pan-Pacific contact between Remote Oceanic and South American populations. THE JOURNAL OF ISLAND AND COASTAL ARCHAEOLOGY 33 Gary M. Heathcote et al. 21. These findings may bear on the “dual structure model” of Japanese population history (K. Hanihara 1991), which posits a more direct ancestral connection between the Jomonese and modern Ainu of Hokkaido, Sakhalin, and the Kuriles, as well as Ryukyuans (Hudson 1999), than to modern Japanese. According to K. Hanihara (see also Brace et al. 1989), modern Japanese of the main islands are a differentially admixed product of two ancestral groups: Neolithic Yayoi migrants from North East Asia and the longer-established Jomon autochthons of proximate Southeast Asian Paleolithic heritage. In comparative craniodental and craniometric and studies, respectively, T. Hanihara (1992) and Pietrusewsky (1992) offer support for the dual origin model. In contrast, Turner (1992) found evidence for Japanese admixture with Jomonese-Ainu, but also a close relationship between modern Japanese and Northeast Asians (“Sinodonts”), based on his study of dental nonmetric traits. Concerning affinities of prehistoric Japanese to Remote Oceanic populations, Brace (e.g., Brace and Hunt 1990) and K. Hanihara (e.g., K. 1993) found Hanihara et al. craniodental and craniometric support, respectively, for JomonPacific Islander population historical connections. Such close Jomon-Pacific Islander affinities are not supported by Pietrusewsky (e.g., 1996, 2006a), in independent craniometric studies. 22. Distribution may include Tierra del Fuego, if any of Matiegka’s (1906) attributed “strong” AP expressions in Fuegians are morphologically equivalent what we score as markedly developed TSPs. 23. If Swisher et al. (1996) are correct in their indirect dating of the Ngandong and Sambungmachan fossils to between roughly 27,000 and 53,000 BP, genetic continuity of a latesurviving population of Indonesian hominins with modern Pacific Rim and Oceanic populations would be problematic, given the advanced H. erectus morphology of these fossils (see Hawks et al. 2000). 34 24. Given the longstanding bias “towards all things ‘Polynesian’ … [in] scholarship on prehistory in the Pacific Islands” (Terrell 2011), a multitude of migration models have been developed concerning the spread of humanity into the non-Micronesian islands of Remote Oceania (see Bellwood 2011; Gosling et al. 2015; Greenhill and Gray 2005; Kirch 2010; Pietrusewsky 2012; Pietrusewsky and Douglas 2016; Sheppard 2011). Historically, treatment of the colonization of Remote Oceania has most often focused on Lapita expansion and the Polynesian diaspora (see Green 2003), and the competing settlement scenarios vary from one another along parameters of demographic viability, directionality, speed of migration, and the natural and cultural environments at the 2011:806), destination (Sheppard factors that surely apply to the peopling of Island Melanesia and Micronesia. 25. Remote AMH ancestors, including some who appear to have carried the genetic potential for developing marked expression of OSSs (through assimilation of Indonesian H. erectus and/or Late Pleistocene northeast Asian genes), were hunter-gatherers, probably of variable lineage origins stemming from several late Pleistocene out-of-Africa AMH colonization events (see Reyes-Centeno et al. 2014). By 50,000–60,000 BP, AMHs began colonizing Sunda (the extension of the SEA continental shelf that included the Malay Peninsula, Sumatra, Borneo, Java, and surrounding islands) and, perhaps somewhat later, Sahul (the great southern landmass which later became New Guinea and Australia) (Morley 2017), but earlier arrivals, dating to the Middle/Late Pleistocene boundary, are suggested by other studies (Corny et al. 2017). Whereas the first occupation of Near Oceania began during the Late Pleistocene, the first expansion into Remote Oceania began much later, during post-glacial Holocene times, from multiple origination points (Montenegro et al. 2016), initially out of southern China, Taiwan, the northern Philippines, and Indonesian archipelago (Ko et al. 2014; Lipson et al. 2014; Oxenham and VOLUME 0  ISSUE 0  2019 Occipital Superstructures across the Pacific Basin Buckley 2016). Southeast Asian voyaging populations—those who were not a part of the Lapita cultural complex—ventured southward, then east into western Micronesia (the Marianas, Palau, and, perhaps, Yap) (Clark et al. 2006; Fitzpatrick and Callaghan 2013; Hung et al. 2011). Contact with the Mariana Islands by sea-goers from ISEA occurred 3500 BP (Vilar et al. 2017) or somewhat later and, around the same time or somewhat earlier, there was movement of sister ISEA populations of Austronesian-speakers associated with the Lapita expansion into the Bismarck Archipelago (Rieth and Athens 2017), the Solomons, and eastern island Melanesia, eventually reaching Tonga and Samoa in Western Polynesia by 3000 BP (Petchey et al. 2010; Vilar et al. 2017). After a pause of approximately 1,000 years, navigators migrated to the furthest reaches of Remote Oceania (Rapa Nui, Hawai’i, and New Zealand [Bellwood et al. 2011; Kirch 2010]), as late as 800 BP (Hunt and Lipo 2006; Pietrusewsky 2012). 26. Such morphological continuity of these archaic humans to Southeast Asian, Oceanic, and Pacific Rim AMHs is supportive of the Assimilation Model of modern human origins, whereby “small but not insignificant anatomical contributions” from archaic to modern humans occurred (Smith et al. 2016:126), in this case, among more immediate ancestors of Oceanic and Pacific Rim AMH populations. 27. To date, aDNA analysis has been conducted (but not fully reported) on only one group from our study, viz. Latte Period remains from the Grand Mariana Resort site, Anaguan (Garapan), Saipan (Table 1). Perzinski and Dega (2016) report that their aDNA study was handicapped by poor biomolecular preservation. Geneticist John Dudgeon was able to partially sequence 13 individuals for ancient mtDNA, and all exhibited Haplogroup E variants with probable affiliation with ISEA E1 and E2 haplogroups. There was no evidence for the Chamorro motif or any other Oceanic B4 lineage (Dega et al. 2017). 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