Late Pleistocene and Holocene Bison of the Colorado Plateau

Abstract

Fossils of Bison (bison) are scarce on the Colorado Plateau, especially in and around the Greater
Grand Canyon Region. Because of poor preservation and collection biases in the region, various resource
managers have erroneously designated bison a nonnative and human-(re)introduced species. This decision
directly impacts an extant herd of approximately 400 bison that periodically meander onto Grand Canyon
National Park lands from neighboring U.S. Forest Service and State of Arizona lands. We re-examined and
verified paleozoological museum specimens of this large mammal confirming a prehistoric and early historic
presence of Bison on the Colorado Plateau. Our findings indicate that Bison bison should be considered a
native species on the Colorado Plateau because they have a nearly continuous record of inhabitance in the
region. This record is supported by 74 Bison-bearing fossil, subfossil, and historical localities since the latest
Pleistocene.

Full text

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research
libraries, and research funders in the common goal of maximizing access to critical research.
Late Pleistocene and Holocene Bison of the Colorado Plateau
Author(s): Jeff M. Martin, Rachel A. Martin, and Jim I. Mead
Source: The Southwestern Naturalist, 62(1):14-28.
Published By: Southwestern Association of Naturalists
DOI: http://dx.doi.org/10.1894/0038-4909-62.1.14
URL: http://www.bioone.org/doi/full/10.1894/0038-4909-62.1.14
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and
environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published
by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of
BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries
or rights and permissions requests should be directed to the individual publisher as copyright holder.

THE SOUTHWESTERN NATURALIST 62(1): 14–28 MARCH 2017
LATE PLEISTOCENE AND HOLOCENE
BISON OF THE COLORADO PLATEAU
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: jeff.m.martin@exchange.tamu.edu
ABSTRACT—Fossils of Bison (bison) are scarce on the Colorado Plateau, especially in and around the Greater
Grand Canyon Region. Because of poor preservation and collection biases in the region, various resource
managers have erroneously designated bison a nonnative and human-(re)introduced species. This decision
directly impacts an extant herd of approximately 400 bison that periodically meander onto Grand Canyon
National Park lands from neighboring U.S. Forest Service and State of Arizona lands. We re-examined and
verified paleozoological museum specimens of this large mammal confirming a prehistoric and early historic
presence of Bison on the Colorado Plateau. Our findings indicate that Bison bison should be considered a
native species on the Colorado Plateau because they have a nearly continuous record of inhabitance in the
region. This record is supported by 74 Bison-bearing fossil, subfossil, and historical localities since the latest
Pleistocene.
RESUMEN—Los f´osiles de Bison (bisontes) son escasos en la meseta del Colorado, especialmente dentro y
alrededor de la regi´on del Gran Ca˜
n´on. Debido a la pobre preservaci ´on de los f ´osiles y parcialidad en las
colecciones de la regi´on, varios administradores de recursos naturales han denominado equivocadamente
bisontes como especies ex´oticas e introducidas por humanos. Esta decisi´on impacta directamente a la
poblaci´on actual de cerca de 400 bisontes que a veces traspasan a los terrenos del Servicio Forestal de E.E.U.U.
y del Estado de Arizona, hasta los terrenos del Parque Nacional del Gran Ca˜
n´on. Re-examinamos y verificamos
espec´ımenes paleozool´ogicos de museos de este gran mam´ıfero, confirmando la presencia prehist ´orica e
hist´orica temprana de Bison en la meseta del Colorado. Nuestros resultados indican que Bison bison debe ser
considerado como especie nativa en la meseta del Colorado por su casi continuo registro de habitaci ´on dentro
de la regi´on. Este record es respaldado por 74 f ´osiles y subf ´osiles relacionados con Bison, y por localidades
hist´oricas desde el Pleistoceno tard´ıo.
Perhaps no other extant animal in North America
possesses the cultural, spiritual, ecological, economic,
political, and natural history attributes that are emblem-
atic of the North American bison (Bison bison; Artiodac-
tyla, 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 Qua-
ternary-aged Bison in physiographic provinces neighbor-
ing 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 assess-
ments (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 archaeo-
logical localities known to contain Bison from the Grand
Canyon and the surrounding CP along with a few just off
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).
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 15

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 applicable.
Key Locality name
Database
reference
Mean age
(cal yr BP) Cited age Dating method References
1 The Neck
a
42SA8502 435 Common Era
(AD) 1235–
1415; AD 1425–
1655; AD 1425–
1655; AD 1485–
1795
Pollen and
charcoal
Osborn et al.,
1995
2 Ada Mesa
a
NA20657 Lots 50
&52
1,100 Georgetown/ San
Francisco AD
600–700+;
Tularosa AD
1100+
(Mogollon
Culture)
Radiocarbon,
lithics, and
ceramics
—
3 Alcove Spring 42SA8512 453 AD 1345–1650 Pollen and
charcoal
Osborn et al.,
1995
4 Awatovi
a
NeotomaDB
5910; NeoMap
2332
850 250–1,450 cal yr
BP
Montgomery et
al., 1949;
Lawrence,
1951;
Agenbroad and
Haynes, 1975
5 AZ J:14:356
a
225 0 to 450 cal yr BP Firepit charcoal AZSite
6 AZ P:8:3
a
1,325 1,200–1,450 cal yr
BP
Lithics and
ceramics
AZSite
7 Mather
Campground
B:16:0461 50 50 –30 yr Bison phalanx This study
8 B:16:105 GRCA 69396 1,050 927 cal yr BP;
1000 yr BP
(lithics)
Bison lipids on
ceramics
Downum et al.,
2011
9 Badger House NeotomaDB
1453; NeoMap
1268
814 650–1,100 yr BP
14
C on unknown Hayes and
Lancaster, 1975
10 Badger Spring MNA.Loc.112-0;
MNA Ariz
D:5:13;
NA10924
10,000 7,500–9,500 yr BP Lithic technology Hesse et al., 1999;
MNA collection
11 Battleship Rock 12,000 Faunal
assemblage
(Equus sp.)
GRCA collection
12 Beamer’s Cabin AZ C:13:0004 GC 555 410–700 cal yr BP Association of
ceramics
Jones, 1986;
Huffer, 2013
13 Bear Ruin AZ P:16:1 625 550–700 cal yr BP Ceramics Haury, 1940;
Agenbroad and
Haynes, 1975;
Lightfoot and
Feinman, 1982
14 Bechan Cave NAU QSP Site
872; GLCA
Accession 81
15,182 11,670 –300;
13,505 –580
yr BP
Mammuthus dung Mead and
Agenbroad,
1992; Kropf et
al., 2007
15 Bessie Bottom
Site
48UT1186;
NeotomaDB
4810
985 910 –80; 1,170
–60 yr BP
— McKern, 1988
16 vol. 62, no. 1The Southwestern Naturalist

TABLE 1—Continued.
Key Locality name
Database
reference
Mean age
(cal yr BP) Cited age Dating method References
16 Bison Alcove 42GR538 (ARCH
115);
MNA.Loc.
9144;
NeotomaDB
6290; NeoMap
2958
445 355 –60; 405 –
65 yr BP
Bison horn and
hoof
Mead et al., 1991
17 Black Mesa
a
D:11:2062;
D:7:0713;
NeotomaDB
6010 & 5980
1,302 885 –72; 1,673
–117 cal yr BP
— Leonard, 1989
18 Canyon Diablo
Dam
NA8793.Lot.1
#7136
975 Pueblo II — —
19 Catclaw Cave AZ F:2:1(ASM) 7,225 12,000 yr BP–
1500 AD
Bison bison with
assorted fishes
associated with
lithics and
ceramics
AZSite
20 Cement Creek
Cave
— 22,585 43,330 –
760;1,120 –40
yr BP
Marmota
flaviventris
Emslie, 2002
21 Charley Day
Spring
NA1898;
MNA.Loc. 149-
1
14,000 Rancholabrean Faunal
assemblage
MNA collection;
Lindsay and
Tessman, 1974;
Agenbroad and
Haynes, 1975
22 Cottonwood Seep
Spring
a
NA14674.Lot.955
and Lot.985 &
GSPS6
975 Pueblo II Lithics MNA collection
23 Cowboy Cave 45WN420;
NeotomaDB
9761; NeoMap
2048
14,519 11,020 –180;
13,040 –440
yr BP
Bison dung Hansen, 1980;
Lucias, 1980;
Mead and
Agenbroad,
1992
24 Coyote Creek NA14064 750 Tularosa phase of
Anasazi
(Pueblo III)
Lithics and
buildings
MNA collection
25 Dust Devil Cave NA7613.Lot.
1066.A9V6;
MNA.UT:V:
13:160
975 Kayenta culture
(Pueblo II)
Ceramics —
26 Ephraim (Witch’s
Knoll)
NeotomaDB:
7683
850 664 –6; 1020 –
22 cal yr BP
— Gillin and Allen,
1941
27 Fort Davy
Crockett
5MF605;
NeotomaDB
5210; NeoMap
1165
490 50 –1; 927 –3
cal yr BP
— Sharrock, 1966;
Walker, 1983
28 Furnace Flats AZ C:13:0010 GC 995 AD 641–1270 Association of
ceramics
Jones, 1986;
Huffer, 2013
29 Grand Falls MNA V8301
Loc.1104
50 50 –30
radiocarbon
years
Bison rib head This study
30 Gray Water Wash MNA.Loc.358-0 14,000 Rancholabrean — —
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 17

TABLE 1—Continued.
Key Locality name
Database
reference
Mean age
(cal yr BP) Cited age Dating method References
31 Grobot Grotto NAU QSP Site
878; GLCA
Accession 82
27,384 18,528 –137;
33,540 –2836
cal yr BP
Bison dung Mead and
Agenbroad,
1989, 1992;
Withers and
Mead, 1993
32 Hamblin Springs NeoMap 6329 5,850 Holocene — —
33 Homolovi I; AZ J:14:3; MNA
AZ J:14:8;
NA952
575 Pueblo IV (AD
1300–1450)
Lithic technology
and ceramic
styles
AZSite
34 Hooper’s Hollow NAU QSP Site
873; GLCA
Accession 82
22,783 22,783 –405 cal
yr BP
Bison dung Mead and
Agenbroad,
1989, 1992;
Withers and
Mead, 1993
35 Huntington
Reservoir
Sinkhole
NeotomaDB
5737; NeoMap
1953
12,668 12,668 –296 yr
BP
Mammut bone
amino acid
Miller, 1987;
Gillette and
Madsen, 1992
36 Island in the Sky — 975 Pueblo II Carving styles —
37 Jim Walter’s Cave — 14,519 11,020 –180;
13,040 –440
yr BP
Associated Bison
dung in
Cowboy Cave
Lucias, 1980
38 Keams Canyon — 14,000 Rancholabrean — Hay, 1927;
Lindsay and
Tessman, 1974;
Agenbroad and
Haynes, 1975
39 Las Colinas — 575 Hohokam
(Pueblo IV)
— Johnson, 1974;
Agenbroad and
Haynes, 1975;
Teague and
Deaver, 1989
40 Mammoth Alcove NAU QSP Site
875; GLCA
Accession 82
20,082 16,630 –280 yr
BP
Mammuthus dung Mead and
Agenbroad,
1992
41 Martinez Gravel
Pit
NeotomaDB 6073 61,655 13,341 –30;
110,000 cal yr
BP
Bounding
formations
Lucas et al., 1988;
Smartt et al.,
1991
42 Marysvale NeotomaDB 7682 978 603 –37; 1379 –
12 cal yr BP
Unspecified
remains
Gillin and Allen,
1941
43 Mastodon
Sinkhole
NeoMap 6315;
42EM231V
14,000 Rancholabrean Faunal
assemblage
—
44 Mesa Verde site
866
NeoMap 1286;
NeotomaDB
5320
750 778 –18; 856 –
37 cal yr BP
— Anderson, 1966
45 Mesa Verde site
875
NeoMap 1284;
NeotomaDB
5318
975 856 –37; 927 –
3 cal yr BP
— Anderson, 1966
46 Kanab Creek NA8960.
NA9074.Lot.2
1,075 850–1300 cal yr
BP
— GRCA collections
47 Mt. Trumbull
a
NA9074 S-213;
MNA.AZ.B:1:23
975 850–1100 cal yr
BP
— MNA collections
48 Oak Haven GLCA Accession
82; NAU QSP
Site 881
11,958 9,180 –100;
11,690 –120
yr BP
Quercus gambelii Mead and
Agenbroad,
1989, 1992;
Withers and
Mead, 1993
18 vol. 62, no. 1The Southwestern Naturalist

TABLE 1—Continued.
Key Locality name
Database
reference
Mean age
(cal yr BP) Cited age Dating method References
49 O’Malley Shelter NeoMap 2774;
NeotomaDB
6162
6,739 7,100 –190 yr
BP
Charcoal Fowler et al.,
1973
50 Pharo Village NeoMap 2848;
Netoma 6191
681 760 –80 yr BP Wood Marwitt, 1968
51 Pine Springs NeotomaDB
4820; 48SW101
8,560 7,695 –195 yr
BP
Bone collagen Sharrock, 1966;
Frison, 1978
52 Point of Pines NeoMap 2236 &
2242;
NeotomaDB
5866
573 543 –28; 595 –
35 cal yr BP
— Stein, 1963;
Agenbroad and
Haynes, 1975
53 Porcupine Cave NeotomaDB 7680 542 510 –75 yr BP Ursus americanus
bone
Heaton, 1988
54 Red Horse Wash
b
NA9528.Lot.4 &
Lot.1 (NA
5164)
975 Cohonina/
Anasazi/
Archaic
(Pueblo II)
Ceramics MNA collections
55 Ridge Ruin 750 Hohokam
(Pueblo III)
Lithics Agenbroad and
Haynes, 1975
56 Sand Dune Cave
b
NA7523.Lot.
133:2183
(MNA.UT.V:13
74)
5,650 Navajo or
Basketmaker I/
III
— MNA collections
57 Sandblast Cave NeoMap 4875;
NeotomaDB
7710
13,704 13,110 –680 yr
BP
Associated
Oreamnos dung
Emslie, 1987,
1988; Mead
and
Agenbroad,
1989, 1992
58 Shrub Ox Alcove GLCA Accession
82; NAU QSP
Site 882
15,003 12,690 –180 yr
BP
Quercus twig with
Bison dung
associated
Mead and
Agenbroad,
1989, 1992;
Withers and
Mead, 1993
59 Smith Creek Cave NeotomaDB
4684; 26WP46
27,267 Reddish-Brown
Silt Zone
(12,600–35,000
yr BP)
14
C on unknown Bryan, 1979;
Mead et al.,
1982, 1992
60 Smith Tank Site CC:5:6 675 1275 AD (Pueblo
III)
Ceramics Woodson, 2007
61 Smoking Pipe NeotomaDB
6363; 42UT150
621 640 –110 yr BP Bison bison bone Billat, 1985
62 Snake Rock
Village
NeoMap 2851;
NeotomaDB
6194
736 1,500 –95 yr BP Wood Aikens, 1967
63 Snaketown — 1,433 835 –109; 1,799
–342 cal yr BP
Ceramics Haury, 1965;
Agenbroad and
Haynes, 1975
64 Spotten Cave 42UT104,
NeotomaDB
6358
684 730 –90 yr BP — Cook, 1980
65 Stanton’s Cave
c
C:5:3; NAU QSP
Site 9121;
GRCA
Accession 4597;
NeotomaDB
5747; NeoMap
2008 & 2004
14,191 5,760 –200;
17,300 –800
yr BP
Oreamnos
harringtoni
dung
Harington and
Euler, 1984;
Martin, 1984;
Mead and
Agenbroad,
1989, 1992
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 19

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 498N
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 picto-
graphs 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 trav-
elled great distances and well beyond the actual zoogeo-
graphic 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
TABLE 1—Continued.
Key Locality name
Database
reference
Mean age
(cal yr BP) Cited age Dating method References
66 Sudden Shelter NeoMap 2225;
NeotomaDB
5856; 45SV6
7,458 6,310 –240;
7,090 –85 yr
BP
Charcoal Lucias and
Colville, 1980
67 Texas Creek
Overlook
NeoMap 1148;
NeotomaDB
5197; 5RB2435
458 430 –50 yr BP Charcoal Creasman and
Scott, 1987
68 Upper Sand
Island Site
— 12,000 13,000–11,000 cal
yr BP
Carving styles Malotki and
Wallace, 2011
69 Walnut Canyon
a
— 750 1150–1220 AD
(Pueblo III)
Ceramics Starkovich, 2011
70 Walton Creek
a
NeotomaDB
5202; 5RT11
1,672 1,730 –225 yr
BP
Charcoal
14
C O’Neil, 1980
71 Wetherill Mesa NeoMap 1282;
NeotomaDB
5316; Site 1644
1,237 1,237 –34 cal yr
BP
Building
foundation
style
Hayes and
Lancaster, 1975
72 Whiskey Creek NeoMap 2438;
NeotomaDB
5968; LA 4986
1,540 600 –32; 2,310
–50 cal yr BP
(Pueblo III –
Early
Basketmaker
II)
— Heller, 1976
73 Wither’s Wallow GLCA Accession
82; NAU QSP
Site 883
13,900 12,010 –160 yr
BP
Mammuthus dung Mead and
Agenbroad,
1989, 1992
74 Zion ZION 12396 14,000 Rancholabrean Pleistocene
gravels
Smith and
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.
20 vol. 62, no. 1The Southwestern Naturalist

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 C.J. 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, intro-
duced ‘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 popula-
tion 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 prov-
inces, most likely because of its remoteness.
MATERIALS AND METHODS—The Colorado Plateau is an im-
mense 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 Paleon-
tology lab. The previously undescribed specimens consist
predominately of dung and skeletal remains from cave, rock
shelter, and packrat midden localities. We verified the identifi-
cation 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 quantifi-
able, 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 (Neoto-
maDB, www.neotomadb.org); Arizona’s Cultural Resource
Inventory (AZSite, www.azsite.asurite.ad.asu.edu/azsite/); and
Neogene Mammal Mapping Portal (NeoMap, www.ucmp.
berkely.edu/neomap/). 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, www.earthpoint.us.; Earth Point
Corporation, Kuna, Idaho) in Google Earth Pro (Google Earth
Pro, version 7.1.5.1557; 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
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 21

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 (https://c14.arch.ox.ac.uk/) 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
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).
22 vol. 62, no. 1The Southwestern Naturalist

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 558N 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).
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 d18-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.
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 23

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 Classifica-
tion 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 present-
ed 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
–30 years BP (measured radiocarbon age of 100.5 –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, radiocar-
bon dating (Beta 374435) returned an age of 50 –30
years BP (measured radiocarbon age of 102.4 –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 Garc´es 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,
TABLE 2—Temporal summary of Bison localities on the Colorado Plateau. ‘‘BP’’ is calendar years Before Present.
Years BP Pecos classification Geologic time Bison localities (n)
160,000–14,300 Preanthropogenic Late Pleistocene 14
14,300–10,000 PaleoIndian Latest Pleistocene 8
10,000–1,300 Archaic Early to late Holocene 15
1,300–850 Pueblo I–II Latest Holocene 14
850–650 Pueblo II Latest Holocene 8
650–400 Pueblo IV Latest Holocene 12
400–Present Pueblo V Latest Holocene 3
24 vol. 62, no. 1The Southwestern Naturalist

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.
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 radiocar-
bon 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 MNA V8301,
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, Exper-
iment.com (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 provid-
ing funds to JMM. We appreciate the continued support of The
Mammoth Site and contribution of anonymous reviewers for
much improvement of the manuscript.
LITERATURE CITED
AGENBROAD, L. D., AND C. V. HAYNES. 1975. Bison bison remains at
Murray Springs, Arizona. Kiva 40:309–313.
AIKENS, C. M. 1967. Excavations at Snake Rock Village and the
Beaver River No. 2 site. University of Utah Anthropological
Papers 87:1–78.
ANDERSON, E. 1966. Animal remains. Pages 113–117 in Contri-
butions to Mesa Verde archaeology III: Site 866 and the
cultural sequence at four villages in the Far View Group,
Mesa Verde National Park, Colorado (R. H. Lister, editor).
University of Colorado Studies Series in Anthropology, No.
12, Boulder.
ANDERSON, R. S., J. L. BETANCOURT,J.I.MEAD,R.H.HEVLY,AND D. P.
ADAM. 2000. Middle- and late-Wisconsin paleobotanic and
paleoclimatic records from the southern Colorado Plateau,
USA. Palaeogeography, Palaeoclimatology, Palaeoecology
155:31–57.
BALKWILL, D. M., AND S. L. CUMBAA. 1992. A guide to the
identification of postcranial bones of Bos taurus and Bison
bison. Canadian Museum of Nature, Ottawa.
BARNOSKY, A. D., M. HOLMES,R.KIRCHHOLTES,E.LINDSEY,K.C.
MAGUIRE,A.W.POUST,M.A.STEGNER,J.SUNSERI,B.SWART Z,J.
SWIFT,N.A.VILLAVICENCIO,AND G. O. WOGAN. 2014. Prelude to
the Anthropocene: two new North American Land Mammal
Ages (NALMAs). The Anthropocene Review 1:225–242.
BELL, C. J., E. LUNDELIUS,JR., A. D. BARNOSKY,R.W.GRAHAM,E.H.
LINDSAY,D.R.RUEZ,JR., H. SEMKEN,JR., S. D. WEBB,AND R. J.
ZAKRSEWSKI. 2004. The Blancan, Irvingtonian, and Rancholab-
rean Mammal Ages. Pages 232–314 in Late Cretaceous and
Cenozoic mammals of North America: biostratigraphy and
geochronology (M. O. Woodburne, editor). Columbia
University Press, New York.
BILLAT, S. E. 1985. A study of Fremont subsistence at the Smoking
Pipe site. M.S. thesis, Brigham Young University, Provo, Utah.
BLAKEY, R. C., AND W. RANNEY. 2008. Ancient landscapes of the
Colorado Plateau: Grand Canyon. Grand Canyon Associa-
tion, Grand Canyon, Arizona.
BROUGHTON, J. M., D. A. BYERS,R.A.BRYSON,W.ECKERLE,AND D. B.
MADSEN. 2008. Did climatic seasonality control late Quater-
nary artiodactyl densities in western North America? Quater-
nary Science Reviews 27:1916–1937.
BRYAN, A. L. 1979. Smith Creek Cave. Pages 164–251 in The
archaeology of Smith Creek Canyon, eastern Nevada (D. R.
Tuohy and D. L. Rendall, editors). Nevada State Museum
Anthropological Papers, No. 17, Carson City.
CANNON, K. 2001. What the past can provide: contribution of
prehistoric bison studies to modern bison management.
Great Plains Research: A Journal of Natural and Social
Sciences 11:145–174.
COATS, L. L., K. L. COLE,AND J. I. MEAD. 2008. 50,000 years of
vegetation and climate history on the Colorado Plateau, Utah
and Arizona, USA. Quaternary Research 70:322–338.
COHEN, K., S. FINNEY,P.GIBBARD,AND J. FAN. 2013. The ICS
international chronostratigraphic chart. Episodes 36:199–
204.
COOK, C. W. 1980. Faunal analysis from five Utah Valley sites: a
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 25

test of a subsistence model from the Sevier Fremont Area.
M.S. thesis, Brigham Young University, Provo, Utah.
CRAINE, J. M. 2013. Long-term climate sensitivity of grazer
performance: a cross-site study. PLoS ONE 8:e67065.
CRAINE, J. M., E. TOWNE,D.TOLLESON,AND J. NIPPERT. 2013.
Precipitation timing and grazer performance in a tallgrass
prairie. Oikos 122:191–198.
CREASMAN, S., AND L. J. SCOTT. 1987. Texas Creek overlook:
evidence for late Fremont (post A.D.1200) occupation in
northwest Colorado. Southwestern Lore 53:1–16.
CRONIN, M. A., M. D. MACNEIL,N.VU,V.LEESBURG,H.D.
BLACKBURN,AND J. N. DERR. 2013. Genetic variation and
differentiation of bison (Bison bison) subspecies and cattle
(Bos taurus) breeds and subspecies. Journal of Heredity
104:500–509.
DOWNUM, C. E., L. S. CUMMINGS,K.PUSEMAN,D.H.SORRELL,AND M.
M. VANCE. 2011. Archaeological excavations at Site B:16:105,
Grand Canyon National Park. Archaeological Report 1247,
Northern Arizona University, Flagstaff.
EMSLIE, S. D. 1987. Age and diet of fossil California condors in
Grand Canyon, Arizona. Science 237:768–770.
EMSLIE, S. D. 1988. Vertebrate paleontology and taphonomy of
caves in Grand Canyon, Arizona. National Geographic
Research 4:128–142.
EMSLIE, S. D. 2002. Fossil shrews (Insectivora: Soricidae) from the
late Pleistocene of Colorado. Southwestern Naturalist 47:62–
69.
FLORES, D. 1991. Bison ecology and bison diplomacy: the
southern plains from 1800 to 1850. Journal of American
History 78:465–485.
FOWLER, D. D., D. B. MADSEN,AND E. M. HATTORI. 1973. Prehistory
of southeastern Nevada. University of Nevada Desert Re-
search Institute Publications in the Social Sciences, No. 6,
Reno.
FRISON, G. C. 1978. Prehistoric hunters of the high plains.
Academic Press, New York.
GARC´
ES,F.T.H.,AND E. COUES. 1900. On the trail of a Spanish
pioneer: the diary and itinerary of Francisco Garc´es
(missionary Priest) in his travels through Sonora, Arizona,
and California, 1775–1776. F.P. Harper, New York.
GILLETTE, D. D., AND D. B. MADSEN. 1992. The short-faced bear
Arctodus simus from the late Quaternary in the Wasatch
Mountains of central Utah. Journal of Vertebrate Paleontol-
ogy 12:107–112.
GILLIN, J., AND G. M. ALLEN. 1941. Archaeological investigations in
central Utah. Papers of the Peabody Museum of Archaeology
and Ethnology, Harvard Museum 17:1–46.
GROOTES, P., M. STUIVER,J.WHITE,S.JOHNSEN,AND J. JOUZEL. 1993.
Comparison of oxygen isotope records from the GISP2 and
GRIP Greenland ice cores. Nature 366:552–554.
HANSEN, R. M. 1980. Late Pleistocene plant fragments in the
dungs of herbivores at Cowboy Cave in Cowboy Cave. Pages
179–189 in Cowboy Cave (J. D. Jennings, editor). University
of Utah Anthropological Papers, No. 104, Salt Lake City.
HARINGTON, C. R., AND R. EULER. 1984. Ungulate remains from
Stanton’s Cave: an identification list. Pages 69–75 in The
archaeology, geology, and paleobiology of Stanton’s Cave:
Grand Canyon National Park, Arizona (R. C. Euler, editor).
Grand Canyon Natural History Association, Grand Canyon,
Arizona.
HARRIS, A. H. 2014. Pleistocene vertebrates of southwestern USA
and northwestern Mexico. University of Texas at El Paso, El
Paso.
HAURY, E. W. 1940. Excavations in the Forestdale Valley, east-
central Arizona. University of Arizona Social Science Bulletin,
No. 12, Tucson.
HAURY, E. W. 1965. Food: animal. Pages 156–158 in Excavations
at Snaketown: material culture (H. S. Gladwin, E. W. Haury,
E. B. Sayles, and N. Gladwin, editors). University of Arizona
Press, Tucson.
HAY, O. P. 1927. On Pleistocene geology of western region of
North America and remains of vertebrates occurring in it:
Arizona. Pages 129–137 in The Pleistocene of the western
region of North America and its vertebrated animals (O. P.
Hay, editor). Publication No. 322B, Carnegie Institution,
Washington, D.C.
HAYES,A.C.,AND J. A. LANCASTER. 1975. Badger House
community, Mesa Verde National Park. U.S. Department of
the Interior, National Park Service, Wetherill Mesa Studies,
Publications in Archeology 7E.
HEATON, T. H. 1988. Bears and man at Porcupine Cave, Western
Uinta Mountains, Utah. Current Research in the Pleistocene
5:71–73.
HELLER, M. M. 1976. Zooarchaeology of Tularosa Cave, Catron
County, New Mexico. M.S. thesis, University of Texas at El
Paso, El Paso.
HESSE, I., W. PARRY,AND F. SMILEY. 1999. Badger Springs: a Late-
Paleoindian site in northeastern Arizona. Current Research
in the Pleistocene 16:27–30.
HOFFMEISTER, D. 1986. Mammals of Arizona. University of
Arizona Press, Tucson.
HORNADAY, W. T. 1889. The extermination of the American bison.
Smithsonian Institution, Government Printing Office, Wash-
ington, D.C.
HUFFER, D. J. 2013. A spatial analysis and zooarchaeological
interpretation of archaeological bison remains in the
Southwest and the wildlife management implications for
the House Rock Valley bison herd in Grand Canyon National
Park, Arizona. M.A. thesis, The University of Arizona, Tucson.
ISENBERG, A. C. 1997. The returns of the Bison: nostalgia, profit,
and preservation. Environmental History 2:179–196.
JASS, C. N., AND C. O. GEORGE. 2010. An assessment of the
contribution of fossil cave deposits to the Quaternary
paleontological record. Quaternary International 217:105–
116.
JENNINGS, J. D. 1978. Prehistory of Utah and the eastern Great
Basin. University of Utah Anthropological Papers 98:1–279.
JOHNSON, P. C. 1974. Mammalian remains from Los Colinas, a
classic Hohokam site in Arizona. M.S. thesis, University of
Arizona, Tucson.
JONES, A. T. 1986. A cross section of Grand Canyon archaeology:
excavations at five sites along the Colorado River. Publica-
tions in Anthropology No. 28. Western Archaeological and
Conservation Center, National Park Service, Tucson, Arizona.
KOCH,P.L.,AND A. D. BARNOSKY. 2006. Late Quaternary
extinctions: state of the debate. Annual Review of Ecology,
Evolution, and Systematics 37:215–250.
KROPF, M., J. I. MEAD,AND R. S. ANDERSON. 2007. Dung, diet, and
the paleoenvironment of the extinct shrub-ox (Euceratherium
collinum) on the Colorado Plateau, USA. Quaternary Re-
search 67:143–151.
LAWRENCE, B. 1951. Mammals found at the Awatovi Site: post-
cranial skeletal characters of deer, pronghorn, and sheep-
26 vol. 62, no. 1The Southwestern Naturalist

goat with notes on Bos and Bison. Papers of the Peabody
Museum of Archaeology and Ethnology, Harvard University
35:1–43.
LEONARD, R. D. W. 1989. Anasazi faunal exploitation: prehistoric
subsistence on northern Black Mesa, Arizona. Southern
Illinois University Center for Archaeological Investigations
Occasional Paper, No. 13, Carbondale.
LIGHTFOOT, K. G., AND G. M. FEINMAN. 1982. Social differentiation
and leadership development in early pithouse villages in the
Mogollon region of the American Southwest. American
Antiquity 47:64–86.
LINDSAY, E. E. H., AND N. T. N. TESSMAN. 1974. Cenozoic vertebrate
localities and faunas in Arizona. Journal of the Arizona
Academy of Science 9:3–24.
LUCAS, S. G., T. WILLIAMSON,AND J. SOBUS. 1988. Late Pleistocene
(Rancholabrean) mammals from the Edith Formation,
Albuquerque, New Mexico. New Mexico Journal of Science
28:51–58.
LUCIAS, W. A. 1980. Bone and shell material. Pages 97–107 in
Cowboy Cave (J. D. Jennings, editor). University of Utah
Anthropological Paper, No. 104, Salt Lake City.
LUCIAS, W. A., AND J. K. COLVILLE. 1980. Osteological analysis of
faunal remains. Pages 157–170 in Sudden shelter (J. D.
Jennings, A. R. Schroedl, and R. N. Holmer, editors).
University of Utah Anthropological Papers, No. 103, Salt
Lake City.
MADSEN, D. B., D. RHODE,D.K.GRAYSON,J.M.BROUGHTON,S.D.
LIVINGSTON,J.HUNT,J.QUADE,D.N.SCHMITT,AND M. W. SHAVER
III. 2001. Late Quaternary environmental change in the
Bonneville Basin, western USA. Palaeogeography, Palae-
oclimatology, Palaeoecology 167:243–271.
MALOTKI, E., AND H. D. WALLACE. 2011. Columbian mammoth
petroglyphs from the San Juan River near Bluff, Utah, United
States. Rock Art Research 28:143–152.
MARTIN, J. M. 2014. Late Pleistocene and Holocene Bison of
Grand Canyon and Colorado Plateau: implications from the
use of paleobiology for natural resource management policy.
M.S. thesis, East Tennessee State University, Johnson City.
MARTIN, P. 1984. Stanton’s Cave during and after the last ice age.
Pages 133–137 in The archaeology, geology, and paleobiology
of Stanton’s Cave: Grand Canyon National Park, Arizona (R.
C. Euler, editor). Grand Canyon Natural History Association,
Grand Canyon, Arizona.
MARWITT, J. P. 1968. Pharo Village. University of Utah Anthro-
pological Papers 91:1–99.
MCCLARAN,M.P.,AND T. R. VAN DEVENDER. 1995. The desert
grassland. University of Arizona Press, Tucson.
MCDONALD, J. N. 1981. North American bison: their classification
and evolution. University of California Press, Los Angeles.
MCKERN, S. 1988. Archaeological investigation at the Bessie
Bottom site (48UT1186): late prehistoric animal procure-
ment in the upper Bear River drainage. Cultural Resource
Management Report, No. 45, Western Wyoming College,
Rock Springs.
MEAD, J. I., AND L. D. AGENBROAD. 1989. Pleistocene dung and the
extinct herbivores of the Colorado Plateau, southwestern
USA. Cranium 6:29–44.
MEAD, J. I., AND L. D. AGENBROAD. 1992. Isotope dating of
Pleistocene dung deposits from the Colorado Plateau,
Arizona and Utah. Radiocarbon 34:1–19.
MEAD, J. I., C. J. BELL,AND L. K. MURRAY. 1992. Mictomys borealis
(northern bog lemming) and the Wisconsin paleoecology of
the east-central Great Basin. Quaternary Research 37:229–
238.
MEAD, J. I., S. E. SHARPE,AND L. D. AGENBROAD. 1991. Holocene
Bison from Arches National Park, southeastern Utah. Great
Basin Naturalist 51:336–342.
MEAD, J. I., R. S. THOMPSON,AND T. R. VAN DEVENDER. 1982. Late
Wisconsinan and Holocene fauna from Smith Creek Canyon,
Snake River Range, Nevada. Transactions of the San Diego
Society of Natural History 20:1–26.
MILLER, W. E. 1987. Mammut americanum, Utah’s first record of
the American mastodon. Journal of Paleontology 61:168–183.
MONTGOMERY, R. G., W. SMITH,AND J. O. BREW. 1949. Franciscan
Awatovi: the excavation and conjectural reconstruction of a
17th-century Spanish mission establishment at a Hopi Indian
town in northeastern Arizona. Papers of the Peabody
Museum of American Archaeology and Ethnology, Harvard
University 36:1–361.
O’NEIL, B. 1980. Highway salvage excavations at Walton Creek
(5RT11), Routt County, Colorado. Southwestern Lore 46:3.
OLSEN, S. J. 1960. Post-cranial skeletal characters of Bison and
Bos. Papers of the Peabody Museum of Archaeology and
Ethnology, Harvard University 35:1–15.
OSBORN, A. J., J. BROWN,G.BURGETT,L.S.CUMMINGS,R.J.HARTLEY,
S. VETTER,J.WAT ERS,AND T. ZALUCHA. 1995. Aboriginal
adaptations on the Colorado Plateau: a view from the
Island-in-the-Sky, Canyonlands National Park, Utah. Midwest
Archeological Center Occasional Studies in Anthropology
33:245–309.
POLYAK,V.J.,AND Y. ASMEROM. 2001. Late Holocene climate and
cultural changes in the southwestern United States. Science
294:148–151.
RANNEY, W. (editor). 2012. Carving Grand Canyon. Second
edition. Grand Canyon Association, Grand Canyon, Arizona.
REED, E. K. 1952. The myth of Montezuma’s bison and the type
locality of the species. Journal of Mammalogy 33:390–392.
REED, E. K. 1955. Bison beyond the Pecos. Texas Journal of
Science 7:130–135.
REIMER, P., E. BARD,AND A. BAYLISS. 2013. IntCal13 and Marine13
radiocarbon age calibration curves 0–50,000 years cal BP.
Radiocarbon 55:1869–1887.
REIMONDO, E. L. 2012. Ecological impacts and management
implications of introduced bison in the Grand Canyon. M.S.
thesis, Northern Arizona University, Flagstaff.
SANDERSON, E. W., K. H. REDFORD,B.WEBER,K.E.AUNE,D.BALDES,
J. BERGER,D.CARTER,C.CURTIN,J.N.DERR,S.DOBROTT,E.
FEARN,C.FLEENER,S.FORREST,C.GERLACH,C.C.GATES,J.E.
GROSS,P.GOGAN,S.GRASSEL,J.A.HILTY,M.JENSEN,K.KUNKEL,
D. LAMMERS,R.LIST,K.MINKOWSKI,T.OLSON,C.PAGUE,P.B.
ROBERTSON,AND B. STEPHENSON. 2008. The ecological future of
the North American bison: conceiving long-term, large-scale
conservation of wildlife. Conservation Biology 22:252–266.
SCOTT, E., AND S. M. COX. 2008. Late Pleistocene distribution of
Bison (Mammalia; Artiodactyla) in the Mojave Desert of
southern California and Nevada. Science Series 41:359–382.
SEAGER, R., N. GRAHAM,C.HERWEIJER,A.L.GORDON,Y.KUSHNIR,
AND E. COOK. 2007. Blueprints for medieval hydroclimate.
Quaternary Science Reviews 26:2322–2336.
SHARROCK, F. W. 1966. Prehistoric occupation patterns in
southwestern Wyoming and cultural relationships with the
Great Basin and Plains cultural areas. University of Utah
Anthropological Papers 77:1–234.
SMARTT, R. A., S. G. LUCAS,AND D. J. HAFNER. 1991. The giant bison
March 2017 Martin et al.—Prehistoric Bison of the Colorado Plateau 27

(Bison latifrons) from the middle Rio Grande Valley of New
Mexico. Southwestern Naturalist 36:136–137.
SMITH, J., AND V. SANTUCCI. 2001. Vertebrate ichnostratigraphy of
the Glen Canyon Group (Jurassic) in Zion National Park,
Utah. Pages 15–19 in The changing face of the east Mojave
Desert and abstracts from the 2001 Desert Symposium (R. E.
Reynolds, editor). California State University Desert Studies
Consortium, Fullerton.
STARKOVICH, B. 2011. Faunal analysis of Walnut Canyon, Arizona.
M.S. thesis, University of Arizona, Tucson.
STEIN, W. 1963. Mammal remains from archaeological sites in the
Point of Pines region, Arizona. American Antiquity 29:213–
220.
TEAGUE,L.S.,AND W. L. DEAVER. 1989. The 1982–1984
excavations at Las Colinas. Cultural Resource Management
Division, Arizona State Museum, University of Arizona,
Tucson.
WAKELING, B. F. 2006. Arizona bison genetics: verifying origins.
Pages 25–30 in Managing wildlife in the Southwest: new
challenges for the 21st century (J. W. Cain III and P. R.
Krausman, editors). A Publication of the Southwest Section
of The Wildlife Society, Tucson, Arizona.
WALKER, D. N. 1983. Faunal remains and subsistence practices at
5MF605, a fur trading post in northwest Colorado. South-
western Lore 49:6–29.
WISELY, S., M. STATHAM,AND R. FLEISCHER. 2008. Pleistocene refugia
and Holocene expansion of a grassland-dependent species,
the black-footed ferret (Mustela nigripes). Journal of Mam-
malogy 89:87–96.
WITHERS, K., AND J. MEAD. 1993. Late Quaternary vegetation and
climate in the Escalante River basin on the central Colorado
Plateau. Great Basin Naturalist 53:145–161.
WOLFF, S. E. 2013. Home on the range: biogeographic
distribution of bison in Arizona. PalArch’s Journal of
Vertebrate Palaeontology 10:1–11.
WOODSON, K. 2007. The Goat Hill Site and ancestral Pueblo
migrations into the Safford Basin. The Petroglyph, Newslet-
ter of the Arizona Archaeological Society 43:7–11.
ZHAO, M., C.-Y. HUANG,AND K.-Y. WEI. 2005. A 28,000 year U K37
sea-surface temperature record of ODP Site 1202B, the
Southern Okinawa Trough. TAO 16:45–56.
Submitted 7 July 2016. Accepted 19 December 2016.
Associate Editor was Troy Ladine.
28 vol. 62, no. 1The Southwestern Naturalist