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Late Pleistocene and Holocene Bison of the Colorado Plateau.

Perhaps no other extant animal in North America possesses the cultural, spiritual, ecological, economic, political, and natural history attributes that are emblematic of the North American bison (Bison bison; Artiodactyla, Bovidae, Bovini; Isenberg, 1997; Sanderson et al., 2008). Certainly, no other animal boasts the story of surviving the brink of extinction twice. This story does not stop at the North American plains subspecies (Bison bison bison), but it also applies to the North American woods bison subspecies (Bison bison athabascae) along with the European bison species (Bison bonasus; wisent). First, Bison survived the megafaunal extinction at the end of the Pleistocene approximately 11,700 calendar years Before Present (cal yr BP) and, in North America, the genus outlived mammoths (Mammuthus), mastodons (Mammut), horses (Equus), ground sloths (Megalonyx et al.), and other megafauna while coexisting with early Americans (Paleoindians; Koch and Barnosky, 2006). Second, Bison survived the threat of Americans of European descent, who purposefully hunted the bison nearly to extinction, during the late 1800s Common Era (Hornaday, 1889).

The most recent comprehensive analysis of Bison distribution in continental North America illustrates an apparent geographic 'gap' of bison records for much of the arid Southwest, especially on the Colorado Plateau (CP; Mc Donald, 1981; Fig. 1). In the western United States, there are well-documented occurrences of Quaternary-aged Bison in physiographic provinces neighboring the CP, such as the Great Basin (Jennings, 1978; Scott and Cox, 2008), southern and middle Rocky Mountains (McDonald, 1981), Wyoming Basin (McDonald, 1981), Arizona deserts (Agenbroad and Haynes, 1975; Wolff, 2013), Bonneville Basin (Madsen et al., 2001), and Rio Grande Rift basin (Harris, 2014).

In recent years, bison from Arizona's House Rock Valley in Arizona have dispersed onto the Kaibab Plateau on the North Rim of the Grand Canyon National Park (NP). Resource managers subsequently questioned the nativity of Bison on the CP and concluded, from lack of evidence and limited tangential environmental assessments (Reimondo, 2012; Huffer, 2013), that bison are not native to the area. These prior studies made no examination of specimens in collections but relied upon published literature. Thus, it is necessary to complete a comprehensive assessment of unpublished accounts of Bison remains from paleontological and archaeological sites that are preserved in local museum collections.

Here, we synthesize the paleontological and archaeological localities known to contain Bison from the Grand Canyon and the surrounding CP along with a few just off the plateau for adjacent references. Our data set includes published and unpublished accounts and unstudied museum specimens along with those records archived in database systems that have recently emerged. Our goal is to address two main questions: 1) when and 2) where did Bison occur on the CP, especially within the Grand Canyon NP and the Greater Grand Canyon Region (as defined in MATERIALS AND METHODS)? We address these questions by assessing, and in some cases describing, museum specimens of Bison from 74 localities on and around the CP (Table 1; Fig. 1). Details about each locality are in the thesis by Martin (2014); we provide only pertinent data herein.

BACKGROUND--Today, there are two, albeit disputed, subspecies of Bison (Cronin et al., 2013): 1) Bison bison bison are historically found in the Great Plains and elsewhere throughout much of North America, and 2) Bison bison athabascae are historically found north of 49[degrees]N latitude in Canada and Alaska. This project does not address subspecies or species but focuses on the animal at the generic level. In all, Bison bison, Bison antiquus, and Bison latifrons have been reported on the CP and in the surrounding provinces (Table 1).

The historical chronicle of bison in the Southwest is complex and confusing. Northern Arizona has scarce historical records, in general, but the Spanish reported a small herd of Bison in the 16th century in east-central New Mexico and adjacent to the CP (Reed, 1952). Native Americans in the 1200s AD and earlier created pictographs and petroglyphs on the walls of Kanab Creek near Kanab, Utah (among other places; Malotki and Wallace, 2011). However, the ideas, memories, and thoughts that inspired these renderings could conceivably have travelled great distances and well beyond the actual zoogeographic distributions, which brings into question the idea that the pictographs and petroglyphs represent local occurrences of bison. A few unrelated, documented occurrences confirm that bison frequented the CP near the Greater Grand Canyon Region in small but self-sustaining herds that most likely had relatively large home ranges to endure the low carrying capacity of the region (Seager et al., 2007).

In 1905, Charles J. "Buffalo" Jones brought bison to the Kaibab Plateau on the North Rim of the Grand Canyon NP (Hoffmeister, 1986) and crossbred them with Bos taurus to create a hardy livestock animal he called "cattelo." In 1906, when Congress established the Grand Canyon Game Preserve, they listed bison as one of the wildlife species that should be maintained on the Kaibab Plateau (unpublished report to U.S. Congress, Protection of Wild Animals in the Grand Canyon Forest Preserve). The Kaibab Plateau was described as "ideal for buffalo [Bison], deer and other wild game" and was "to be recognized as a breeding place therefore" (Wakeling, 2006:25). Most importantly, "the Preserve was created on 28 November 1906 by President Theodore Roosevelt and is still in effect. It predates the establishment of the [Kaibab] National Forest [in 1909], Grand Canyon National Park [in 1919], and the Arizona Game and Fish Commission [in 1913]" (Wakeling, 2006:25). In 1909, these bison were relocated east to the House Rock Valley (Marble Platform) because of the creation of the Kaibab National Forest when CJ. Jones moved all but 15-20 animals out of the area. The 15-20 remaining bison became property of James T. "Uncle Jim" Owens and, by 1927, the herd had increased to 98 individuals and was purchased by the State of Arizona via the Arizona Game and Fish Department. Over the past half-century, introduced 'purebred' Bison bison from Oklahoma and Montana have been added to the bison herd to improve its natural resilience (see discussions in Hoffmeister, 1986; Wakeling, 2006).

Throughout the Holocene, Bison abundances spatially varied over most of North America possibly because of hunting and intraspecific competition; yet, the population steadily increased in the Great Plains until European contact when their abundance probably reached their apex (Flores, 1991). This increase in bison might have occurred because of a few conditions including limited trophic-equivalent competition (Flores, 1991); reduced abundance and diversity of presumed predators since the Pleistocene extinction of the dire wolf (Canis dirus), American lion (Panthera atrox), and the giant short-faced bear (Arctodus simus; Flores, 1991); and cooler, wetter climate conditions, which are favorable for grass growth (Wisely et al., 2008; Craine, 2013; Craine et al., 2013). Thus, the observations made by early western explorers (Hornaday, 1889) are likely misleading or at least poorly representative of Bison abundance.

Furthermore, earlier in the Holocene, Bison were less abundant in the Great Plains and were intermittently present in the Southwest (Broughton et al., 2008). There are potentially multiple factors contributing to the seemingly poor fossil record of Bison during the Late Pleistocene and Holocene on the CP. Yet, the region is incompletely studied by Quaternary paleontologists and zooarchaeologists compared with the neighboring provinces, most likely because of its remoteness.

MATERIALS AND METHODS--The Colorado Plateau is an immense physiographic province between the Colorado Rocky Mountains and the Great Basin Desert (Blakey and Ranney, 2008). The Grand Canyon is defined as the geological gorge that incises the Colorado Plateau (Ranney, 2012) and is immediately surrounded by Grand Canyon NP. We define the Greater Grand Canyon Region as the ecosystem of Grand Canyon NP adjacent to the Colorado River corridor and the plateaus immediately rimming the Grand Canyon gorge. This includes the Coconino, Hualapai, Kaibab, Kanab, Uinkaret, and Shivwits plateaus and the Marble Platform. Thus, we also extend the Greater Grand Canyon Region to approximately 80 km beyond the borders of Grand Canyon NP to encompass the habitats of each plateau (Fig. 1).

Each Bison locality on the CP is listed in Table 1 and is numbered consecutively to correspond with the location shown in Figure 1. Undocumented and misidentified specimens from archaeological and paleontological localities are critical to our assessment. These specimens are archived in collections at the Museum of Northern Arizona (MNA), at the Grand Canyon NP, and at the East Tennessee State University Vertebrate Paleontology lab. The previously undescribed specimens consist predominately of dung and skeletal remains from cave, rock shelter, and packrat midden localities. We verified the identification of each specimen and, if necessary, corrected it using Balkwill and Cumbaa (1992), who improved upon identification guides by Lawrence (1951) and Olsen (1960). In an attempt to capture the full extent of variation, Balkwill and Cumbaa (1992) included every postcranial element and provide 192 quantifiable, osteological characters for comparing Bison, represented by 27 individuals of B. bison bison and B. bison athabascae, and Bos taurus, represented by 16 individuals of several breeds including Holstein, Ayrshire, Shorthorn, Longhorn, and Africander. Balkwill and Cumbaa (1992) described specimens of both sexes and of various ages to account for natural variation within Bison and Bos.

Occurrences of Bison were also recovered from the following online data sets: Neotoma Paleoecology Database (NeotomaDB,; Arizona's Cultural Resource Inventory (AZSite,; and Neogene Mammal Mapping Portal (NeoMap, wwwucmp. Several localities were in the legal format of Township and Range, which creates a large polygon instead of a point and is imprecise when using point data for other site localities. For each site that was in the legal format, we converted the data into the coordinate system by using a centralized datum in the Township and Range overlay (B. Clark, Earth Point: Township and Range--Public Land Survey System on Google Earth,; Earth Point Corporation, Kuna, Idaho) in Google Earth Pro (Google Earth Pro, version; Google Inc., Googleplex, Mountain View, California) using decimal degrees. The location is less accurate but more precise for geospatial assessment and analysis.

Each locality has a name and numbering system used by the archiving institution and maintained here. As part of this research, we directly radiocarbon dated Bison specimens from two sites, B:16:0461 and Grand Falls. For other sites, the age is first given as reported in the literature (if provided) or archival database. We converted the given age(s) to calendar years BP as a single mean geologic age for geospatial representation (Fig. 2). The calibration of the radiocarbon years was completed using OxCal Online Tool ( by employing the IntCal13 curve (Reimer et al., 2013). We assumed the average date of the calibrated age to be accurate, but it should be understood to be imprecise. We provide details including pertinent published references, archival databases, and dating sources for each site when available.

North American Quaternary Chronologies--There are a number of chronologies that are defined by specific metrics and cannot be easily integrated because of the inherent differences of paleontological and archaeological definitions; we define these below (Fig. 3). Each chronology used here is necessary for relating our various paleozoological sites that contain Bison remains. Furthermore, we implemented a climatic representation for understanding the environmental context of each period.

The Pleistocene is divided into biochronological units referred to as North American Land Mammal Ages based on faunal assemblage components. The Rancholabrean Land Mammal Age is defined by the first appearance of Bison south of 55[degrees]N latitude and begins approximately 160,000 years BP (Bell et al., 2004). Furthermore, the Rancholabrean Land Mammal Age terminates 14,000 years BP when the Santarosaean Land Mammal Age begins (see discussion in Barnosky et al., 2014, for Land Mammal Age divisions within the Holocene).

In addition to the mammal ages, archaeological stages were implemented based on North American human cultures in the Southwest (Polyak and Asmerom, 2001). The Pecos Classification was used in this study because it is culturally specific to the CP region and to the Southwest, and it often relates to the specimens we observed in museum collections. The later subdivisions of the Pecos Classification are within modern times. For this study, 850 years BP to today, which includes the formal historic, prehistoric, and protohistoric periods, is considered modern times to illustrate Bison nativity both precontact and postcontact with Europeans in the Southwest.

RESULTS--Herein, we report 74 sites located on or near the CP (with a few on the periphery) that contain Bison. We add 26 localities to the 48 reported in a recent study by Huffer (2013). Moreover, the previous study found that only 16 localities were described from the Holocene, of which 10 were on the CP (Huffer, 2013). In contrast to the Huffer (2013) study, we found 52 sites from the Holocene, of which 40 were on the CP.

Our assessment resulted in the summary data presented in Table 2. We present two sites of interest in and near Grand Canyon NP--B:16:0461 (specimen GRCA 69396) and Grand Falls (specimen MNA V8301; Table 1). We selected these specimens because they were previously insufficiently identified and only relatively dated.

Site B:16:0461, in the Mather Campground area of Grand Canyon NP, was a surface collection that had not been previously reported or identified. It is a lateral half of a proximal phalanx that is identified as Bison sp. because the lateral margin is curved (see Balkwill and Cumbaa (1992) and Martin (2014) for discussion). This sample returned a radiocarbon date (Beta 374436) of 50 [+ o -] 30 years BP (measured radiocarbon age of 100.5 [+ o -] 0.4 pMC).

The Grand Falls specimens represent two nearly complete postcranial individuals deposited 15 m apart in the same arroyo. In collections, these specimens' elements are mostly joined together with consolidant and adhesive. Here, we only describe the right astragalus because it was not treated with consolidant or adhesive. The medial tubercle of the astragalus is on a level with or above the line drawn across the proximal margin of the distal trochlea, as elaborated in Balkwill and Cumbaa (1992). The posterior surface of the astragalus is excavated and extends as far as the lateral margin. Grand Falls was described in the MNA computer database as an arroyo site near the Little Colorado River, which suggests that it is possibly of Pleistocene age. However, radiocarbon dating (Beta 374435) returned an age of 50 [+ o -] 30 years BP (measured radiocarbon age of 102.4 [+ o -] 0.4 pMC). Subsequently, we placed both B:16:0461 and Grand Falls at the beginning of the 20th century (Pueblo V).

DISCUSSION--Much of what we know about bison on the CP is based on historical and modern studies. Although these are valuable resources, they are incomplete and do not explore the prehistoric record. We found that 13 of our newly identified 26 Bison localities (50%) were either previously not identified as Bison or were incorrectly identified as "unknown," "large mammal," or "Bos." Previous studies rarely identified Bison remains from in situ, pre-European contact because it was traditionally thought that Bison did not inhabit the CP. We are particularly intrigued that some researchers identified an in situ, pre-European contact faunal remain as "Bos taurus," knowing that this taxon arrived with the exploration of Spanish conquistadors, approximately 1540 Common Era. We hope to dispel this persistent line of circular reasoning by providing a thorough assessment of Bison on the CP since the latest Pleistocene.

Some hypothesize that the bison found in the Greater Grand Canyon Region were carrion from scavengers, bloat-and-float from upstream Colorado River, or goods traded by peoples. However, it would seem that the pre-Colombian cultural trading of Bison elements might not have been traded long distances but more locally (Cannon, 2001). Reed (1952, 1955) points out that the bison skeletal and artifactual remains found in a pre-Spanish context across the southern CP (Arizona) are likely due to trade connections; albeit, Reed does state that the record of bison ("cibola") provided by the Havasupai Indians in the Grand Canyon probably represents a local procurement and not a trade item (independently described in Garces and Coues, 1900:403, 406).

In addition, Bison are not known to frequent caves and are categorized as low frequenters of such shelters, with no more than 16.9% of all Bison remains found in caves across the contiguous United States (Jass and George, 2010). Thus, finding Bison remains in caves or rock shelters at 45 of 74 (60.8%) sites on the CP appears to be significant because either they were inhabiting the area or they were scavenged from a nearby location. For comparison, approximately 85.7% of bighorn sheep (Ovis canadensis) remains are found in caves (Jass and George, 2010).

CONCLUSION--The direct impact of this study is the production of the first comprehensive review of late Pleistocene and Holocene Bison on the Colorado Plateau. The results indicate that the geographic range of fossil and modern Bison can be extended to include the CP. Our data imply that Bison have been part of the CP from at least approximately 44,000 radiocarbon years ago though the latest Pleistocene (Rancholabrean) to the onset of the Holocene (~11,000 years ago). The past approximately 11,700 years is a critical time because it marks the beginning of the modern climate based on paleobotanical records (McClaran and Van Devender, 1995; Coats et al., 2008). Major ecological and faunal turnover occurred at or by approximately 11,700 years ago, yet the early Holocene climate was still colder than today. The fossil Bison record for the early Holocene is not well-reported on the CP, but there appears to be good evidence of Bison presence during the late Holocene, through the various cultural phases, and up to the time of European contact. The southern CP (south of the Grand Canyon) in Arizona appears to have been fairly devoid of bison; but, within the Grand Canyon (likely just north of the Colorado River) and north, Bison were likely present, albeit never likely abundant based on its record. Though Bison were not overly common, such as observed on the Plains, the fossil record clearly demonstrates that this iconic bovid played a role in the biotic communities over much, if not all, of the Colorado Plateau up to the time of European contact.

Submitted 7 July 2016. Accepted 19 December 2016.

Associate Editor was Troy Ladine.

We thank S. L. Swift and M. Carpenter for the countless hours contributed to data collection and improvement of this project. We thank the National Park Service (ZION 12396, Zion National Park Museum Collection) for the courtesy of abundant information and a specimen approved for destructive radiocarbon analysis (GRCA 69396) from the Grand Canyon National Park (research permit# GRCA-2013-SCI-0052), and C. Hyde and B. Holton for informative discussions about the House Rock Valley bison herd. We thank the Navajo Nation, who in conjunction with the Museum of Northern Arizona provided their specimen for destructive radiocarbon analysis MNAV8301, Loc. 1104 (Navajo Nation Minerals Department research permit date: 30 August 2013). We thank many staff members of the Museum of Northern Arizona for their assistance. S. Emslie and M. Stiger provided important information for bison presence in the Gunnison Basin. We thank the National Buffalo Foundation, the Dr. Ken Throlson American Buffalo Scholarship, (a crowd-sourcing platform), the Western Bison Association, the East Tennessee State University (ETSU) Don Sundquist Center of Excellence in Paleontology, and the ETSU Graduate School Graduate Student Research Grant for providing funds to JMM. We appreciate the continued support of The Mammoth Site and contribution of anonymous reviewers for much improvement of the manuscript.


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Jeff M. Martin, * Rachel A. Martin, and Jim I. Mead

Wildlife and Fisheries Sciences, Texas A&M University, College Station, TX 77843 (JMM)

Ecosystem Science and Management, Texas A&M University, College Station, TX 77843 (RAM)

The Mammoth Site, Hot Springs, SD 57747 (JIM)

Don Sundquist Center of Excellence in Paleontology, East Tennessee State University Gray Fossil Site and Museum,

Gray, TN 37615 (JMM, RAM, JIM)

* Correspondent:

Caption: FIG. 1--Map of Bison localities (placement is approximate) on the Colorado Plateau (CP, heavy line) and surrounding region. Locality numbers refer to Table 1. Hashed line represents our definition of the Greater Grand Canyon Region (GGC, see Materials and Methods section) surrounding Grand Canyon National Park (GCNP, shaded area).

Caption: Fig. 2--Map of Bison localities placed in chronological context on the Colorado Plateau (CP, heavy black line) and surrounding region. Hashed line represents our definition of the Greater Grand Canyon Region (GGC; see text). Time unit is in thousands of calendar years ago BP (kya).

Caption: FIG. 3--Integrated North American Quaternary chronology. This time scale displays geological, paleontological (North American Land Mammal Ages: [Bell et al., 2004; Barnosky et al., 2014]), climatic (Climate Regime: Zhao et al., [2005]; Cohen et al., [2013]), and cultural (Pecos Classification: Polyak and Asmerom [2001]) units that are all important for relating Bison localities. Not all units are displayed to scale; some cultural units are enlarged for legibility. The relative temperature (solid line) is represented as a proxy derived from 818-Oxygen isotopes from the Greenland Ice Sheet Project (GISP2). Data from Marine Oxygen Isotope Stages (Anderson et al., 2000); and temperature from GISP2 (Grootes et al., 1993). Abbreviations: ka = kilo annum (thousand years ago); BP, Before Present; CE, Common Era; %, part per thousand.
TABLE 1--Summary table of Bison localities from the Colorado
Plateau region. Key refers to number and location in Figure 1.
Mean age in calendar years Before Present (cal yr BP) calibrated
by IntCal13 (Reimer et al., 2013). References other than primary
literature include GRCA collection (Grand Canyon National Park
collections), MNA (Museum of Northern Arizona), AZSite
(Arizona's Cultural Resource Inventory), NeotomaDB (Neotoma
Paleoecology Database), and NeoMap (Neogene Mammal Mapping Portal).
"--" indicates no data. Empty cells indicate no information is

Key           Locality name                  Database reference

1     The Neck (a)                     42SA8502
2     Ada Mesa (a)                     NA20657 Lots 50 & 52
3     Alcove Spring                    42SA8512
4     Awatovi (a)                      NeotomaDB 5910; NeoMap 2332
5     AZJ:14:356 (a)
6     AZ P:8:3 (a)
7     Mather Campground                B:16:0461
8     B:16:105                         GRCA 69396
9     Badger House                     NeotomaDB 1453; NeoMap 1268
10    Badger Spring                    MNA.Loc.112-0; MNA Ariz
                                         D:5:13; NA10924
11    Battleship Rock
12    Beamer's Cabin                   AZ C:13:0004 GC
13    Bear Ruin                        AZ P:16:1
14    Bechan Cave                      NAU QSP Site 872;
                                         GLCA Accession 81
15    Bessie Bottom Site               48UT1186; NeotomaDB 4810
16    Bison Alcove                     42GR538 (ARCH 115); MNA.Loc.
                                         9144; NeotomaDB 6290;
                                         NeoMap 2958
17    Black Mesa (a)                   D:11:2062; D:7:0713;
                                         NeotomaDB 6010 & 5980
18    Canyon Diablo Dam                NA8793.Lot.1 #7136
19    Catclaw Cave                     AZ F:2:1(ASM)
20    Cement Creek Cave
21    Charley Day Spring               NA1898; MNA.Loc. 1491
22    Cottonwood Seep Spring (a)       NA14674.Lot.955 and
                                         Lot.985 & GSPS6
23    Cowboy Cave                      45WN420; NeotomaDB 9761;
                                         NeoMap 2048
24    Coyote Creek                     NA14064
25    Dust Devil Cave                  NA7613.Lot. 1066.A9V6;
                                         MNA.UT:V: 13:160
26    Ephraim (Witch's Knoll)          NeotomaDB: 7683
27    Fort Davy Crockett               5MF605; NeotomaDB
                                         5210; NeoMap 1165
28    Furnace Flats                    AZ C:13:0010 GC
29    Grand Falls                      MNA V8301 Loc.1104
30    Gray Water Wash                  MNA.Loc.358-0
31    Grobot Grotto                    NAU QSP Site 878;
                                         GLCA Accession 82
32    Hamblin Springs                  NeoMap 6329
33    Homolovi I;                      AZ J:14:3; MNA AZ
                                         J:14:8; NA952
34    Hooper's Hollow                  NAU QSP Site 873;
                                         GLCA Accession 82
35    Huntington Reservoir Sinkhole    NeotomaDB 5737; NeoMap 1953
36    Island in the Sky                --
37    Jim Walter's Cave                --
38    Keams Canyon                     --
39    Las Colinas                      --
40    Mammoth Alcove                   NAU QSP Site 875;
                                         GLCA Accession 82
41    Martinez Gravel Pit              NeotomaDB 6073
42    Marysvale                        NeotomaDB 7682
43    Mastodon Sinkhole                NeoMap 6315; 42EM231V
44    Mesa Verde site 866              NeoMap 1286; NeotomaDB 5320
45    Mesa Verde site 875              NeoMap 1284; NeotomaDB 5318
46    Kanab Creek                      NA8960. NA9074.Lot.2
47    Mt. Trumbull (a)                 NA9074 S-213; MNA.AZ.B:1:23
48    Oak Haven                        GLCA Accession 82;
                                         NAU QSP Site 881
49    O'Malley Shelter                 NeoMap 2774; NeotomaDB 6162
50    Pharo Village                    NeoMap 2848; Netoma 6191
51    Pine Springs                     NeotomaDB 4820; 48SW101
52    Point of Pines                   NeoMap 2236 & 2242;
                                         NeotomaDB 5866
53    Porcupine Cave                   NeotomaDB 7680
54    Red Horse Wash (b)               NA9528.Lot.4 & Lot.1 (NA 5164)
55    Ridge Ruin
56    Sand Dune Cave (b)               NA7523.Lot. 133:2183
                                         (MNA.UT.V:13 74)
57    Sandblast Cave                   NeoMap 4875; NeotomaDB 7710
58    Shrub Ox Alcove                  GLCA Accession 82;
                                         NAU QSP Site 882
59    Smith Creek Cave                 NeotomaDB 4684; 26WP46
60    Smith Tank Site                  CC:5:6
61    Smoking Pipe                     NeotomaDB 6363; 42UT150
62    Snake Rock Village               NeoMap 2851; NeotomaDB 6194
63    Snaketown
64    Spotten Cave                     42UT104, NeotomaDB 6358
65    Stanton's Cave (c)               C:5:3; NAU QSP Site 9121;
                                         GRCA Accession 4597;
                                         NeotomaDB 5747;
                                         NeoMap 2008 & 2004
66    Sudden Shelter                   NeoMap 2225;
                                         5856; 45SV6
67    Texas Creek                      NeoMap 1148;
        Overlook                         NeotomaDB 5197; 5RB2435
68    Upper Sand                       -Island
69    Walnut Canyon (a)                -
70    Walton Creek (a)                 NeotomaDB
                                         5202; 5RT11
71    Wetherill Mesa                   NeoMap 1282;
                                         5316; Site 1644
72    Whiskey Creek                    NeoMap 2438;
                                         5968; LA 4986
73    Wither's Wallow                  GLCA Accession
                                         82; NAU QSP
                                         Site 883
74    Zion                             ZION 12396

Key    Mean age                 Cited age
      (cal yr BP)

1            435    Common Era (AD) 12351415;
                    AD 14251655;
                    AD 14251655;
                    AD 14851795
2          1,100    Georgetown/ San
                      Francisco AD 600-700+;
                      Tularosa AD 1100+
                      (Mogollon Culture)
3            453    AD 1345-1650
4            850    250-1,450 cal yr BP
5            225    0 to 450 cal yr BP
6          1,325    1,200-1,450 cal yr BP
7             50    50 [+ o -] 30 yr
8          1,050    927 cal yr BP; 1000 yr
                      BP (lithics)
9            814    650-1,100 yr BP
10        10,000    7,500-9,500 yr BP
11        12,000
12           555    410-700 cal yr BP
13           625    550-700 cal yr BP
14        15,182    11,670 [+ o -] 300; 13,505
                      [+ o -] 580 yr BP
15           985    910 [+ o -] 80; 1,170
                      [+ o -] 60 yr BP
16           445    355-60; 405 -65 yr BP
17         1,302    885-72; 1,673-117 cal yr BP
18           975    Pueblo II
19         7,225    12,000 yr BP-1500 AD
20        22,585    43,330 -760;1,120-40 yr BP
21        14,000    Rancholabrean
22           975    Pueblo II
23        14,519    11,020-180; 13,040-440 yr BP
24           750    Tularosa phase of Anasazi
                      (Pueblo III)
25           975    Kayenta culture (Pueblo II)
26           850    664 [+ o -] 6; 1020 [+ o -]
                      22 cal yr BP
27           490    50 [+ o -] 1; 927-3 cal yr BP
28           995    AD 641-1270
29            50    50 [+ o -] 30 radiocarbon years
30        14,000    Rancholabrean
31        27,384    18,528 [+ o -] 137;
                      33,540 [+ o -] 2836 cal yr BP
32         5,850    Holocene
33           575    Pueblo IV (AD 1300-1450)
34        22,783    22,783 [+ o -] 405 cal yr BP
35        12,668    12,668 [+ o -] 296 yr BP
36           975    Pueblo II
37        14,519    11,020 [+ o -] 180;
                      13,040 [+ o -] 440 yr BP
38        14,000    Rancholabrean
39           575    Hohokam (Pueblo IV)
40        20,082    16,630 [+ o -] 280 yr BP
41        61,655    13,341 [+ o -] 30;
                      110,000 cal yr BP
42           978    603 [+ o -] 37;
                      1379 [+ o -] 12 cal yr BP
43        14,000    Rancholabrean
44           750    778 [+ o -] 18;
                      856 [+ o -] 37 cal yr BP
45           975    856 [+ o -] 37;
                    927 [+ o -] 3 cal yr BP
46         1,075    850-1300 cal yr BP
47           975    850-1100 cal yr BP
48        11,958    9,180 [+ o -] 100;
                      11,690 [+ o -] 120 yr BP
49         6,739    7,100 [+ o -] 190 yr BP
50           681    760 [+ o -] 80 yr BP
51         8,560    7,695 [+ o -] 195 yr BP
52           573    543 [+ o -] 28;
                      595 [+ o -] 35 cal yr BP
53           542    510 [+ o -] 75 yr BP
54           975    Cohonina/ Anasazi/ Archaic
                      (Pueblo II)
55           750    Hohokam (Pueblo III)
56         5,650    Navajo or Basketmaker I/ III
57        13,704    13,110 [+ o -] 680 yr BP
58        15,003    12,690 [+ o -] 180 yr BP
59        27,267    Reddish-Brown Silt Zone
                      (12,600-35,000 yr BP)
60           675    1275 AD (Pueblo III)
61           621    640 [+ o -] 110 yr BP
62           736    1,500 [+ o -] 95 yr BP
63         1,433    835 [+ o -] 109;
                      1,799 [+ o -] 342 cal yr BP
64           684    730 [+ o -] 90 yr BP
65        14,191    5,760 [+ o -] 200;
                      17,300 [+ o -] 800 yr BP
66         7,458    6,310 [+ o -] 240;
                      7,090 [+ o -] 85 yr
67           458    430 [+ o -] 50 yr BP
68        12,000    13,000 [+ o -] 11,000 cal
                      yr BP
69           750    1150 [+ o -] 1220 AD
                      (Pueblo III)
70         1,672    1,730 [+ o -] 225 yr
71         1,237    1,237 [+ o -] 34 cal yr
72         1,540    600 [+ o -] 32; 2,310
                      [+ o -] 50 cal yr BP
                      (Pueblo III - Early
                      Basketmaker II)
73        13,900    12,010 [+ o -] 160 yr

74        14,000    Rancholabrean

Key           Dating method                       References

1     Pollen and charcoal              Osborn et al., 1995
2     Radiocarbon, lithics,            -and
3     Pollen and charcoal              Osborn et al., 1995
4                                      Montgomery et al., 1949;
                                         Lawrence, 1951;
                                         Agenbroad and Haynes, 1975
5     Firepit charcoal                 AZSite
6     Lithics and ceramics             AZSite
7     Bison phalanx                    This study
8     Bison lipids on ceramics         Downum et al., 2011
9     [sup.14]C on unknown             Hayes and Lancaster, 1975
10    Lithic technology                Hesse et al., 1999;
                                         MNA collection
11    Faunal assemblage (Equus sp.)    GRCA collection
12    Association of ceramics          Jones, 1986; Huffer, 2013
13    Ceramics                         Haury, 1940; Agenbroad and
                                         Haynes, 1975; Lightfoot
                                         and Feinman, 1982
14    Mammuthus dung                   Mead and Agenbroad, 1992;
                                         Kropf et al., 2007
15    --                               McKern, 1988
16    Bison horn and hoof              Mead et al., 1991
17    --                               Leonard, 1989
18    --                               --
19    Bison bison with assorted        AZSite
        fishes associated with
        lithics and ceramics
20    Marmota flaviventris             Emslie, 2002
21    Faunal assemblage                MNA collection; Lindsay and
                                         Tessman, 1974; Agenbroad
                                         and Haynes, 1975
22    Lithics                          MNA collection
23    Bison dung                       Hansen, 1980; Lucias, 1980;
                                         Mead and Agenbroad, 1992
24    Lithics and buildings            MNA collection
25    Ceramics                         --
26    --                               Gillin and Allen, 1941
27    --                               Sharrock, 1966; Walker, 1983
28    Association of ceramics          Jones, 1986; Huffer, 2013
29    Bison rib head                   This study
30    --                               --
31    Bison dung                       Mead and Agenbroad, 1989,
                                         1992; Withers and Mead, 1993
32    --                               --
33    Lithic technology and            AZSite
        ceramic styles
34    Bison dung                       Mead and Agenbroad, 1989,
                                         1992; Withers and Mead, 1993
35    Mammut bone amino acid           Miller, 1987; Gillette
                                         and Madsen, 1992
36    Carving styles                   --
37    Associated Bison dung            Lucias, 1980
        in Cowboy Cave
38    --                               Hay, 1927; Lindsay and Tessman,
                                         1974; Agenbroad and
                                         Haynes, 1975
39                                     Johnson, 1974; Agenbroad and
                                         Haynes, 1975; Teague
                                         and Deaver, 1989
40    Mammuthus dung                   Mead and Agenbroad, 1992
41    Bounding formations              Lucas et al., 1988; Smartt
                                         et al., 1991
42    Unspecified remains              Gillin and Allen, 1941
43    Faunal assemblage                --
44    --                               Anderson, 1966
45    --                               Anderson, 1966
46    --                                GRCA collections
47    --                                MNA collections
48    Quercus gambelii                 Mead and Agenbroad, 1989, 1992;
                                         Withers and Mead, 1993
49    Charcoal                         Fowler et al., 1973
50    Wood                             Marwitt, 1968
51    Bone collagen                    Sharrock, 1966; Frison, 1978
52                                     Stein, 1963; Agenbroad
                                         and Haynes, 1975
53    Ursus americanus bone            Heaton, 1988
54    Ceramics                         MNA collections
55    Lithics                          Agenbroad and Haynes, 1975
56    --                               MNA collections
57    Associated Oreamnos dung         Emslie, 1987, 1988; Mead
                                         and Agenbroad, 1989, 1992
58    Quercus twig with Bison          Mead and Agenbroad, 1989,
        dung associated                  1992; Withers and Mead, 1993
59    [sup.14]C on unknown             Bryan, 1979; Mead
                                         et al., 1982, 1992
60    Ceramics                         Woodson, 2007
61    Bison bison bone                 Billat, 1985
62    Wood                             Aikens, 1967
63    Ceramics                         Haury, 1965; Agenbroad
                                         and Haynes, 1975
64    --                               Cook, 1980
65    Oreamnos harringtoni dung        Harington and Euler, 1984;
                                         Martin, 1984; Mead and
                                         Agenbroad, 1989, 1992
66    Charcoal                         Lucias and
                                         Colville, 1980
67    Charcoal                         Creasman and
                                         Scott, 1987
68    Carving styles                   Malotki and
                                         Wallace, 2011
69    Ceramics                         Starkovich, 2011
70    Charcoal [sup.14]C               O'Neil, 1980
71    Building                         Hayes and
      foundation                       Lancaster, 1975
72    --                               Heller, 1976
73    Mammuthus dung                     Mead and
                                         1989, 1992
74    Pleistocene                      Smith and
      gravels                            Santucci, 2001; J. Hall,
                                         Zion NPS, pers. comm. 2013.

(a) Taxonomic reclassification from Bos taurus to Bison bison.

(b) Taxonomic classification from "Unknown" to Bison sp.

(c) Taxonomic reclassification from Equus sp. to Bison sp.

TABLE 2--Temporal summary of Bison localities on the Colorado
Plateau. "BP" is calendar years Before Present.

Years BP              Pecos             Geologic time

160,000-14,300   Preanthropogenic   Late Pleistocene
14,300-10,000    PaleoIndian        Latest Pleistocene
10,000-1,300     Archaic            Early to late Holocene
1,300-850        Pueblo I-II        Latest Holocene
850-650          Pueblo II          Latest Holocene
650-400          Pueblo IV          Latest Holocene
400-Present      Pueblo V           Latest Holocene

Years BP              Bison
                 localities (n)

160,000-14,300         14
14,300-10,000           8
10,000-1,300           15
1,300-850              14
850-650                 8
650-400                12
400-Present             3
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Author:Martin, Jeff M.; Martin, Rachel A.; Mead, Jim I.
Publication:Southwestern Naturalist
Article Type:Report
Geographic Code:1USA
Date:Mar 1, 2017
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