Amphibians and Squamates from the Late Pleistocene (Rancholabrean) Clark Quarry, Coastal Georgia
Dennis Parmley1, *, Joshua L. Clark2, and Alfred J. Mead3
1Department of Biological and Environmental Science, Georgia College & State University, Milledgeville, Georgia 31061. 2College of Coastal Georgia, Brunswick, Georgia 31520. 3Department of Biological and Environmental Science, Georgia College & State University, Milledgeville, Georgia 31061. *Corresponding author.
Eastern Paleontologist, No. 7 (2020)
Abstract
Pleistocene amphibians and squamates are reported from a coastal Georgia site known as the Clark Quarry, located in Glynn County. Although dominated by skeletal remains of giant bison and Columbian mammoths, the site has yielded a relatively diverse microfossil collection of amphibians and squamates to include two species of frogs, two salamanders, one lizard, and at least (conservatively) 10 snake species, for a total of 15 genera. Collectively, these microfossils provide noteworthy information about the taxonomic diversity and paleoecology of the Late Pleistocene paleoherpetofauna of southeast Georgia. The paleofauna suggests the presence of an aquatic habitat in a parkland during the time of deposition.
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No. 6 2020
Proboscideans from
US National Park
Service Lands
Jim I. Mead, Justin S. Tweet,
Vincent L. Santucci, Jeffrey T. Rasic,
and Sharon E. Holte
Eastern Paleontologist
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Cover Photograph: Mammuthus columbi skull fragment with in situ large tusks from Waco Mammoth
National Monument (WACO), Texas (photo courtesy NPS).
Eastern Paleontologist
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2020 No. 6
1
2020 EASTERN PALEONTOLOGIST 6:1–48
Proboscideans from US National Park Service Lands
Jim I. Mead1*, Justin S. Tweet2, Vincent L. Santucci3,
Jeffrey T. Rasic4, and Sharon E. Holte5
Abstract - Proboscideans (Mammalia, Proboscidea) are an ubiquitous part of North American vertebrate
faunas throughout the Miocene, Pliocene, and Pleistocene. Here we discuss the fossil record
of proboscideans found on public lands administered by the National Park Service (NPS), which is
comprised of 419 units. At least 276 of these units contain some aspect of fossil heritage for the USA.
We present 63 NPS units and affiliated areas that have records documenting fossil proboscideans. The
geological and paleoecological diversity preserved and represented in these 63 units record fossils
from Arctic to tropical and steppe to rainforest environments. This is an invaluable data set that has
yet to be fully recognized. The information presented here, much of which has not been published, is
intended as a compilation to support researchers.
Introduction
Since 1916, the National Park Service (NPS, US Department of the Interior) has been
entrusted with the care of our national parks, and as their mission, they are to preserve unimpaired
the natural and cultural resources. The official emblem contains the arrowhead,
mountains, Sequoia tree, and bison which represent aspects of our natural and cultural resources
and exemplify the overall mission. The National Park System covers more than 85
million acres and is comprised of 419 sites with at least 19 different designations, including
130 historical units, 87 monuments, 61 national parks along with a number of other types of
units. In addition, the NPS recognizes, but does not manage, National Natural Landmarks
(NNL), National Historic Landmarks (NHL), and other affiliated sites. These preserved and
recognized localities represent a significant expanse of geographic area of the USA and embody
a tremendous aggregate of natural and cultural heritage to conserve and understand.
The geological and paleoecological diversity preserved in these lands and represented in
their fossil record (from Arctic to tropical, and steppe to rainforest) is noteworthy and an
invaluable data set that has yet to be fully recognized and understood (see overview about
fossils on federal lands in Liggett et al. 2018).
On March 30th, 2009 President Barack Obama signed the Paleontological Resource Preservation
Act into law authorizing five federal land managing agencies, including the NPS, to
understand, preserve, and conserve their fossil resources (see discussion in Santucci 2017).
At least 276 NPS units contain some aspect of our fossil heritage. Here we concentrate on
the vertebrate resources preserved or recognized by our nation’s National Park System (administrated
and affiliated) and focus on one iconic group of mega-mammals, the proboscideans.
To help better understand the evolution of Proboscidea in North America, we provide
1*The Mammoth Site, 1800 Hwy 18 ByPass, Hot Springs, SD 57747, jmead@mammothsite.org;
Desert Laboratory on Tumamoc Hill, University of Arizona, Tucson, AZ 85745; and East Tennessee
State University Natural History Museum, Johnson City, TN 37614. jmead@mammothsite.org; 928-
853-6393. 2National Park Service, 9149 79th Street South, Cottage Grove, MN 55016. justin_tweet@
nps.gov. 3National Park Service, Geologic Resources Division, 1849 “C” Street, Washington, DC,
20240. vincent_santucci@nps.gov. 4National Park Service, Gates of the Arctic National Park and
Preserve, and Yukon-Charley rivers National Preserve, 4175 Geist Road, Fairbanks, AK 99709. 5The
Mammoth Site, 1800 Hwy 18 ByPass, Hot Springs, SD 57747. sharonh@mammothsite.org.
Manuscript Editor: Robert Feranec
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here in one outlet a sometimes hard-to-assemble resource for researchers and interested
public who want to further understand these mammals from all NPS and affiliated lands
(Fig. 1). Proboscideans are found as fossils throughout the continent, both east and west of
the Mississippi River. Our approach is to view the distribution of fossil proboscideans from
NPS administered and affiliated lands (from here on collectively referred to as NPS Units)
on a continental-wide scope. The information presented here is intended as a compilation to
support researchers, to provide a preliminary examination of the holdings on NPS Units so
that more detailed studies can be planned. It is not our intent to present all the actual data
in any detailed account, but do indicate which NPS Units contain proboscidean remains.
Information about these remains is not always published or readily available to researchers.
We also present in a historiographic approach the discovery, recovery, and issues associated
with these remains. We hope that this document will help spur further analyses into better
understanding this often poorly understood group of land mammals.
Proboscideans
Proboscideans (Mammalia, Proboscidea) are a ubiquitous part of North American
vertebrate faunas throughout the Miocene, Pliocene, and Pleistocene. There were still
members of three families in greater North America during their waning phase in the
late Pleistocene: Gomphotheriidae (Gomphotheres), Mammutidae (Mastodonts), and Elephantidae
(Mammoths).
The phylogenetic understanding of Proboscidea is almost purely based upon the paleontological
record (Shoshani and Tassy 2005, Tassy 1996). Indigenous proboscideans in
North America have been extinct since the end of the Rancholabrean North American Land
Figure 1. Map of the USA with location of National Park Service Units (as defined in text) that have
in situ specimens or records of proboscidean fossils. Numbers refer to NPS Unit names listed alphabetically
in the text.
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Mammal Age (LMA; end of the late Pleistocene) and the beginning of the Holocene. The
precise timing and the ultimate cause(s) of the extinction are still hotly debated and heavily
scrutinized. The intent here is not to assess the extinction cause or its fin alization.
Proboscideans were the largest land mammals in North America during the late Cenozoic.
They originated in the Paleocene in northern Africa and southern Asia along the Tethys
Sea (Lambert and Shoshani 1998). At approximately 16 Ma (Barstovian LMA, Ba1), the
arrival of Zygolophodon (Vacek 1877; Mammutidae) in North America marks the Proboscidean
Datum. By 17–15 Ma North America was experiencing the Middle Miocene Climatic
Optimum which permitted a rich, warm temperate mixed mesophytic forest with numerous
conifers over much of the northern continent and the incipient spread of grasslands. The
woodland-grasslands of the Great Plains region of today graded west into a montane coniferous
forest and mixed mesophytic deciduous forest biome (see discussion in Woodburne
2004). At approximately 15 Ma the continental climate began to deteriorate into a cooling
trend—a possible response to a global shift in the oceanic circulation likely caused by a
major expansion of the East Antarctic Ice Sheet (Flower and Kennett 1993). By the late
Barstovian (Ba2, ~14.6 Ma) the first arrival of Gomphotheriidae occurred in the Western
Hemisphere (Tedford et al. 2004).
During the remaining Miocene and through the Pliocene, Proboscidea of North America
included numerous members within two families: Mammutidae and Gomphotheriidae (see
discussions in Lambert 1996, Saunders 1996, Shoshani and Tassy 2005). These proboscideans
promptly dispersed across North America and ultimately south into South America
during the Great American Biotic Interchange of the late Pliocene (Blancan LMA). Climate
continued cooling through the Pliocene culminating in the Pleistocene (beginning ca. 2.58
Ma), defined by its multitude of glacial maxima (20+), interglacial warm episodes, faunal
dispersals, and evolutionary developments. The first occurrence of Elephantidae in North
America during the early Pleistocene marks the beginning of the Irvingtonian LMA with the
arrival of the Elephantidae, Mammuthus (Brookes 1828). The current and most conservative
estimate for the oldest Mammuthus in North America is 1.3 Ma. However, the earliest
arrival date is still debated (see discussion in Bell et al. 2004).
Proboscideans and our National Park System
Sixty-three NPS Units have a record of proboscideans (Table 1), although not all specimens
have been placed into a retrievable numerical dataset system. The summary below
provides a synopsis about each park unit or affiliated site and the proboscidean remains
that are known from or are curated for the locality. The number accompanying the NPS
Unit listed below (text and Table 1) refers to the location on Figure 1. The great majority
of NPS proboscidean records are from the late Pleistocene. Specimens diagnostic to the
species level are almost all Mammut americanum (Kerr 1792 fide Osborn 1936, American
mastodon; although see below Mammut pacificus, Dooley et al. 2019, Pacific mastodon),
Mammuthus columbi (Falconer 1857, Columbian mammoth), or Mammuthus primigenius
(Blumenbach 1799, Woolly mammoth). Many of the specimens have been only provisionally
identified. Specific and authoritative taxonomic identifications should be reassessed and
studied in more detail. Researchers interested in viewing fossils referred to below should
contact the park unit manager even though the actual specimens may be cataloged off-site
at a nearby institution or regional repository, and in some cases widely distributed across
the USA. The NPS requires that all recovery of fossil specimens be undertaken through an
approved research and collecting permit.
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Table 1. List of localities from NPS Units (as defined in text) that contain a record of
proboscideans. See text for details about identification details for reported proboscideans.
Abbreviations: Ameb, Amebelodon, Eubel, Eubelodon; Gom, gomphothere undefined;
Gompho, Gompotherium; Mam, Mammuthus; Mamut, Mammut; Mast, mastodon undefined;
Probo, Proboscidea undefined; Rhyn, Rhynchotherium; Stego, Stegomastodon;
Tetra, Tetralophodon.; Zygo, Zygolophodon.
Map Locality Name NPS Abbreviation Proboscidean Recorded
1 Agate Fossil Beds AGFO Mam
2 Aleutian World War II ALEU Probo
3 Amistad AMIS Mam
4 Anza-Borrego Desert ANBO-CA Gompho, Stego, Mam, Rhyn
5 Arches ARCH Mam
6 Ashfall Fossil Beds ASFO-NE Eubel
7 Bent’s Old Fort BEOL Mam
8 Bering Land Bridge BELA Mam
9 Big Bend BIBE Mam
10 Big Bone Lick BIBO-KY Mamut
11 Big Cypress BICY Mam
12 Big Thicket BITH Mam
13 Blackwater Draw BLDR-NM Mam
14 Canaveral CANA Mast
15 Cape Krusenstern CAKR Mam
16 Channels Islands CHIS Mam
17 Colonial COLO Mamut
18 Colorado COLM Probo
19 Death Valley DEVA Prob
20 Denali DENA Mam
21 Florissant Fossil Beds FLFO Mam
22 Gates of the Arctic GAAR Mam
23 Glen Canyon GLCA Mam
24 Golden Gate GOGA Mam, Mamut
25 Grand Canyon GRCA Mam
26 Great Sand Dunes GRSA Mam
27 Hagerman Fossil Beds HAFO Mamut
28 Hopewell Culture HOCU Mast
29 Independence INDE Mamut
30 John Dave Fossil Beds JODA Ameb, Gomph,
Tetra, Zygo
31 John Muir JOMU Mam
32 Joshua Tree JOTR Mam
33 Klondike Gold Rush KLGO Mam
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1. Agate Fossil Beds National Monument (AGFO), Nebraska
Cook (1914) reported that a Mammuthus ? columbi tooth was recovered from “Niobrara
River gravels” from the Cook Family ranch at Agate in 1906. This fossil discovery was within
the vicinity of what is now AGFO and potentially within monument boundaries, but the precise
locality is not known. The whereabouts of the specimen is also unknown; our queries
to the American Museum of Natural History and the University of Nebraska State Museum
were unsuccessful in locating the mammoth tooth. Robert Hunt (University of Nebraska State
Museum, Lincoln, NE, December 2018, pers. comm.) noted that the Niobrara gravels of Agate
occasionally have fossils and originate from the Hartville Uplift to the west.
Table 1. Continued.
Map Locality Name NPS Abbreviation Proboscidean Recorded
34 Kobuk Valley KOVA Mam
35 Lake Mead LAKE Mam
36 Lake Meredith LAMR Mam, Gom
37 Lava Beds LABE Mamut
38 Lehner Mammoth-Kill Site LEMA-AZ Mam, Mamut
39 Mammoth Cave MACA Mamut
40 Mammoth Site of Hot Springs MASI-SD Mam
41 Mississippi MISS Mam
42 Mojave MOJA Mam
43 Montezuma Castle MOCA Prob
44 Murray Springs Clovis Site MUSP-AZ Mam
45 Natchez Trace NATR Mast
46 Navajo NAVA Mam
47 New Jersey Pinelands PINE Mamut
48 Nez Perce NEPE Mam
49 Niobrara NIOB Stego, Mam
50 Noatak NOAT Mam
51 Olympic OLYM Mam
52 Padre Island PAIS Mam
53 Potomac Heritage POHE Mamut
54 Rancho La Brea RALA-CA Mam, Mamut
55 Salinas Pueblo Missions SAPU Mam
56 Santa Monica Mountains SAMO Mam
57 Tule Springs Fossil Beds TUSK Mam
58 Valley Forge VAFO Mamut
59 Vicksburg VICK Mam, Mamut
60 Waco Mammoth WACO Mam
61 White Sands WHSA Mam
62 Wupatki WUPA Mam
63 Yukon-Charley Rivers YUCH Mam
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2. Aleutian World War II National Historic Area (ALEU), Alaska
Proboscidean fossils are recorded in the photographs taken by US military servicemen in
the mid-1940s (Fig. 2). The remote islands are the home of the Unangax (Aleut) people for
8,000 years, and during World War II, the islands became a fierce battleground. There apparently
is no precise locality information for the fossil remains featured in the photographs.
Pleistocene deposits with fossils are rare to almost nonexistent on the Aleutian Islands. The
closest credible record of a mammoth to this region is on St. Paul Island which was part of
the Bering Land Bridge (see BELA #8 and Graham et al. 2016 for more discussion).
3. Amistad National Recreation Area (AMIS), Texas
Proboscidean fossils have been found at Bonfire Shelter in Mile Canyon near Langtry,
southwestern Texas. Although the rock shelter is just outside of the boundary of AMIS
by a few tens of meters (set at 348 m [1144 ft] elevation), the lower canyon is included
in the recreation unit, thus the record is included here due to its importance to the overall
understanding of the region. Dibble and Lorrain (1968) were the first to mention the
proboscidean remains, which they cited as “Elephas”. Bement (1986) identified the fos-
Figure 2. Clifford McGinnis
with mammoth remains he
found on Amchitka Island
(ALEU), a volcanic island
of the Rat Island group in
the Aleutian chain (photo
supplied by NPS by Robert
H. McGinnis 1944–1946;
memo to file).
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sils as Mammuthus sp. based on molar remains associated with the skull and mandible.
Additional skeletal remains include a fragmented lumbar vertebra, rib fragments, pelvis
fragments, tibia, and an unidentifiable long bone fragment. Taxa from the same stratigraphic
unit include Bison, Camelops, Capromeryx, Equus, and others (Bement 1986).
Most of the distinction of the locality and its contents comes from the Bison and cultural
remains. The site is thought to represent a bison jump and processing area, but this has
been contested (see discussion in Bement 2007). The importance here is that Mammuthus
was recovered from this steep-sided canyon adjacent to the Rio Grande River, a region
where its Pleistocene fauna are not fully understood. Lands abutting the canyons to the
north include the southern end of the Edwards Plateau and the Southern High Plains, areas
well-known to contain proboscidean remains.
4. Anza-Borrego Desert State Park National Natural Landmark (ANBO-CA), California
ANBO-CA in southern California is the largest of California’s state parks, and was designated
a National Natural Landmark in 1974. It has an outstanding stratigraphic record of the
past 10 million years, including both marine and terrestrial rocks and fossils. Among the fossil
resources is a significant record of proboscideans. McDaniel and Jefferson (2006) reported
more than 80 proboscidean localities within the state park. Most of the finds are fragmentary,
but at least four taxa at the genus or species level can be distinguished: the gomphotheriids
Gomphotherium sp. (Burmeister 1837) and Stegomastodon sp. (Pohlig 1912) and the elephantids
Mammuthus columbi and M. meridionalis (Nesti 1825, Southern mammoth; McDaniel
and Jefferson 2006; although see the supplementary information from Lister and Sher 2015
and Lister 2017 for a different assessment of the M. meridionalis specimens). Records of
Mammut and Cuvieronius (Osborn 1923) have been re-evaluated as representing other taxa
(McDaniel and Jefferson 2006, Murray 2008). An additional genus, Rhynchotherium (Falconer
1868) was not included by McDaniel and Jefferson (2006) but was tentatively accepted by
Murray (2008) on the basis of an unpublished specimen. Paleontological work began in what
is now the state park in the 1930s, conducted by the American Museum of Natural History
(McDaniel and Jefferson 2006). The Natural History Museum of Los Angeles County and Imperial
Valley College also made large collections from the 1950s to the 1980s (McDaniel and
Jefferson 2006); these specimens, apart from type specimens, are now reposited at ANBO-CA
(Cassiliano 1999).
The proboscidean record of ANBO-CA begins in the Miocene, approximately 10 Ma,
with tusk and cheek tooth fragments from the Borrego Buttes attributed to gomphotheres.
Gomphotherium sp. is represented in the Blancan LMA Desguynos Formation and Diablo
Formation, at approximately 4.4 Ma and 3.7–3.6 Ma respectively. Two taxa are present
in the younger Hueso Formation: Stegomastodon sp. and Mammuthus meridionalis, the
latter showing the beginning of the Irvingtonian LMA here at approximately 1.2 Ma (a
1.4 Ma record is a mistake resulting from a typographical error; Murray 2008). Additional
Irvingtonian-age mammoths have been found in the Ocotillo Conglomerate and
Bautista beds. Both M. meridionalis and M. columbi are known from numerous specimens,
including a skull and partial skeleton representing the most complete specimen of
North American M. meridionalis. These specimens show an overlap of the two species
from approximately 1.1 to 0.9 Ma (McDaniel and Jefferson 2006). Potential evidence of
butchering of this specimen (e.g., Miller et al. 1988, 1991) has been reinterpreted as biteand-
glide marks left by large carnivorans (McDaniel and Jefferson 1997, 1999). Presumed
Rancholabrean-age proboscideans have been found near the park in the Salton Trough
(McDaniel and Jefferson 2006).
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5. Arches National Park (ARCH), Utah
Agenbroad and Mead (1989) reported on a Mammuthus sp. mandible recovered from
Lower Courthouse Wash in ARCH (Fig. 3). A sample of the bone produced a uranium-series
age of 17,200 ± 800 (KU-88-A1) yr. B.P. Santucci (2000) reported that in the 1950s, Chief
Ranger Lloyd Pierson discovered the mammoth mandible eroding out of the sediments in a
small alcove. The sample has since been curated into the NPS museum collections. No other
bones were found by Agenbroad or Mead in the shallow alcove.
6. Ashfall Fossil Beds State Historical Park (ASFO-NE) and National Natural Landmark,
Nebraska.
This northeastern Nebraska locality was originally known as Poison Ivy Quarry and
contains predominantly articulated mammal skeletons that record the catastrophic impact of
a volcanic eruption (Voorhies 1985). Designated a NNL in 2006, the state park (not administrated
by the NPS) contains a number of sedimentary formations including Cretaceous,
Paleogene, and Neogene fossils (Tucker et al. 2014, Voorhies and Corner 1993). Over 40
volcanic tuff layers are recorded from the stratigraphic units in the park and surrounding
area. The locality is best known for its tremendous remains of the articulated rhino, Teleoceras
major, which are found in ash beds (Ash Hallow Formation; Tucker et al. 2014) and
date to 11.93 Ma (Perkins and Nash 2002). Of importance here is the underlying Valentine
Formation which is composed of fluvial sands, silts, and gravel and date to the Barstovian
Land Mammal Age, 14–13 Ma. In this lower formation numerous disarticulated remains of
the proboscidean, cf. Eubelodon (Barbour 1914; Gomphotheriidae) are known, reported in a
field guide list but not described (Tucker et al. 2014: Table 2). The location is administered
by the Nebraska State Parks and the University of Nebraska State Museum.
7. Bent’s Old Fort National Historic Site (BEOL), Colorado
A fragmented Mammuthus tusk was discovered at BEOL in 1964 in gravel at a borrow
pit. The discovery was made accidentally by heavy machinery, and some damage was
incurred. This specimen was identified as Mammuthus columbi by US Geological Survey
Figure 3. Edentulous mandible of Mammuthus from Lower Courthouse Wash, ARCH (photo
courtesy NPS).
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(USGS) geologist Glenn Scott (Tweet et al. 2015). Additional undescribed fragments of
mammoth tusks and teeth have been found at various places in the historic site (Scott et al.
2001, Tweet et al. 2015).
8. Bering Land Bridge National Preserve (BELA), Alaska
BELA protects a remnant of the Bering Land Bridge, the land mass exposed during
periods of lowered sea levels through the Pleistocene that connected North America (via
Alaska) with Asia (via Chukotka; autonomous federal subject of Russia) and provided an
important pathway for faunal exchanges between continents. BELA encompasses the northern
half of the Seward Peninsula and is one of the most remote units of the NPS system.
Along with other northern Alaskan NPS units (Cape Krusenstern National Monument and
Gates of the Arctic National Park) it defines the northwestern-most distribution of proboscidean
fossils in the NPS system.
Beginning as early as 1816, numerous vertebrate fossils, including Mammuthus sp.
(likely Mammuthus primigenius) have been observed and collected from eroding coastal
bluffs and river banks on the northern Seward Peninsula, most famously at Elephant
Point (and nearby Eschscholtz Bay), so named for its numerous well-preserved mammoth
finds (Buckland 1831, Hooper 1884, Kotzebue 1821, Maddren 1905, Seemann 1853:33),
which include rare examples of preserved mammoth soft tissue and hair (Quackenbush
1909:107–10). While Elephant Point lies ~80 km (50 mi) east of and outside the BELA
preserve boundary, the same frozen silt (yedoma) deposits continue into the preserve along
its northern and western edges, and the same excellent fossil preservation exists within the
preserve. Early research expeditions drawn to Elephant Point also noted scattered finds of
mammoth bones and ivory on lands that would later be included within BELA, specifically
in the vicinity of Good Hope Bay as well as the western coast of the preserve in areas surrounding
Shishmaref Inlet (Gilmore 1908, Maddren 1905, Moffit 1905:42, Quackenbush
1909:93). In 1885 Cantwell (1887), for example, noted the front half of the skull of a mammoth
lying on the tundra near Shishmaref Inlet. Quackenbush (1909:123) observed a small
mammoth tusk projecting from the surface of the ground near Imuruk Lake, in the eastern
interior portion of what would later become BELA.
Similar reports of isolated finds of mammoth bones, usually in detrital contexts, have
continued to accumulate over the past two centuries and through recent decades. USGS
collections by David Hopkins from 1967–1970 yielded at least 18 Mammuthus sp. skeletal
specimens from scattered locations across the Seward Peninsula that were primarily within
BELA at locations that include Good Hope Bay, Nuglungnugtuk River estuary, and Shishmaref
Lagoon (Repenning 1967, 1970). Santucci et al. (1995) reported a previously undocumented
juvenile mammoth specimen curated in the collections of the Colorado School of
Mines, but this specimen was not confidently identified in a recent inquiry (D. Gertenbach,
D. Schlegel, and S. Luallin, Colorado School of Mines, Golden, CO, March 2019, pers.
comm.). BELA museum collections include additional isolated proboscidean (likely Mammuthus
sp.) remains: a leg bone from a beach within the park, and five specimens of teeth,
tusks, and miscellaneous bones.
In 2012, NPS staff documented skeletal remains of Mammuthus primigenius in a shallow
lake context in the northern portion of the preserve (Rasic 2012) (Fig. 4). As with previous
finds from BELA these also derived from ice-rich silt permafrost deposits, but unlike these
isolated finds from secondary contexts, the “Mammoth Lake” locality contains multiple
post-cranial elements (at least 6) from a single individual and was found in near-primary
context (Rasic 2012). Additional elements are embedded in the muddy bottom of a shallow
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thaw lake and preservation of a substantial portion of a complete skeleton was indicated by
a geophysical survey of the site conducted in 2015 (Urban et al. 2016). As such this is one
of the more complete mammoth skeletons known from the region. Collagen from a tooth
and a vertebra yielded radiocarbon ages of 12,330 ± 50 BP (Beta-329841) and 12,430 ± 50
BP (Beta-331336), respectively (Rasic 2012).
Four late Pleistocene age Mammuthus primigenius specimens are known from an
inland dry cave context at the Trail Creek Caves locality in the western portion of the
preserve. This series of small caves was used intermittently by people for some 10,000
years (Lanik et al. 2017, Larsen 1968, Lee and Goebel 2016) and has both archeological
and paleontological components. Extensive excavations undertaken in 1949–1950 yielded
extinct late Pleistocene age fauna such as bison and horse from Cave 9 (Larsen 1968,
Pasda 2012), but not until 1985, in the course of excavations by NPS archeologists, were
mammoth specimens identified in a separate cave, Cave B (Vinson 1988, 1993). Recent
AMS dating of collagen from the 1985 collections, a scapula and a vertebra, produced
radiocarbon ages of 13,940 ± 70 (Beta-258437) and 13,770 ± 60 (Beta-258436) (Rasic and
Shirar 2009). These dates are favored over previous radiocarbon assays, one anomalously
young and not subsequently replicated, and one with a substantial standard error (Schaaf
1988:456, Vinson 1988:415). No human association with the mammoth remains at Trail
Creek Cave B is indicated.
Figure 4. Geologist Louise
Farquharson with a woolly
mammoth humerus recovered
from a shallow lake bed in
Bering Land Bridge National
Preserve in 2012. NPS photo
by J. Rasic.
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9. Big Bend National Park (BIBE), Texas
Proboscidean remains are known from BIBE but have not been reported on in any detail.
Pliocene-age silts and gravels from Estufa Canyon in BIBE have produced a mandible, rib,
and vertebrae from a long-jawed mastodon (Stevens 1993). Maxwell et al. (1967) mention
that two “elephant” teeth (presumed to be Mammuthus) were found in the Grapevine
Spring area. Wick and Corrick (2015) suggest this locality may be a Pleistocene-age cienega
(spring) deposit. A fragmentary tusk of a Pleistocene proboscidean was found in a wash in
the northern part of the park in 1987 (Reeder 1987).
10. Big Bone Lick National Natural Landmark (BIBO-KY), Kentucky
BIBO-KY (not administered by NPS), named for its voluminous megafaunal remains,
is also a State Historical Park in Boone County, Kentucky. Native Americans knew of the
bones and the tremendous amount of salt in the springs, and Europeans learned of the existence
of Big Bone Lick from them. A French-Native military expedition under Charles Le
Moyne (Baron de Longueuil) secured the first specimens for scientific study in 1739, and
a 1744 map of Louisiana marks the locality as the “place where they found the elephant
bones in 1739”. Meriwether Lewis visited the site in 1803 on his way to join the Corps of
Discovery and sent a box of specimens back to President Thomas Jefferson. Jefferson sent
William Clark in 1807 to Big Bone Lick to discover more about the local fossils, making
this the first organized paleontological expedition in the USA.
Although well known for many decades, the fossil deposit was not studied in detail
until the 1960s (Schultz et al. 1963, 1967). The potential occurrence of Clovis PaleoIndian
artifacts with Mammut americanum from BIBO-KY are discussed in Tankersley et
al. (2009). Specimens from the locality are archived in institutions including the Museé
National d’Histoire Naturelle (Paris), Academy of Natural Sciences of Drexel University,
University of Nebraska State Museum, Museum of Comparative Zoology, and the US
National Museum.
11. Big Cypress National Preserve (BICY), Florida
A partial molar of Mammuthus was found at the “Jetport Site” (a reference to the Dade-
Collier Training and Transition Airport) within BICY. This Pleistocene site also produced
a small number of horse, camel, and tortoise fossils. The mammoth specimen is curated at
the Florida Museum of Natural History.
12. Big Thicket National Preserve (BITH), Texas
A Mammuthus tooth and associated mandible fragment are within the archived collections
at BITH, Menard Creek Unit. The molar was removed in 1981 from the Trinity
River Pleistocene sediments. The mandible is archived at Texas Memorial Museum, Austin.
Pleistocene fossils at BITH are found mixed with reworked sediments of the Miocene-age
Fleming Formation (Kenworthy et al. 2007). Additional but not currently identified skeletal
remains have been retrieved from the same area as the molar (Fay 2009).
13. Blackwater Draw National Historical Landmark (BLDR-NM), New Mexico
Blackwater Draw includes one of the most significant archeological sites in North
America. Blackwater Draw Locality #1, the site that has come to exemplify the Clovis culture,
including the type Clovis points and abundant examples of artifacts in association with
mammoths and extinct bison (Hester 1972, Katz 1997). The excavation areas and the public
museum are owned and operated by Eastern New Mexico University. Located in eastern
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Roosevelt County, New Mexico near the border with Texas, the Blackwater Draw drainage
today is intermittent, but in the recent past the area was wetter and supported large mammals
such as bison, horses, and mammoth. At the close of the Pleistocene and into the Holocene,
early North American people butchered large mammals at a spring-fed pond perched above
the draw (Haynes and Agogino 1966, Hester 1972). The pond dried up approximately 8000
years ago (Hester 1972).
In the early 1930s, a gravel pit was established at the pond site. Mammuthus and Bison
bones were reported to E.B. Howard in November 1932 (Howard 1935), leading to the
first of a number of investigations. The active use of the site as a gravel pit for decades had
a significant influence on scientific investigations. Gravel is below the fossiliferous sediments,
necessitating removal of the sediments to get at the gravel (Hester 1972). Much
of the scientific work at the site can be described as salvage archeology and paleontology
(Katz 1997). The economic use of the site halted early efforts to preserve some part of it
(Hester 1972). Nevertheless, a number of significant finds have been made; beyond its
archeological reputation, Blackwater Locality #1 is the most prolific mammoth site in
New Mexico (Lucas and Effinger 1991). Material is assigned to either M. columbi or the
less specific M. sp., and comes from the gray sand and brown sand wedge units (Lucas
and Effinger 1991).
Cotter (1937) provided the first description of mammoth bones associated with artifacts
at the site. More spectacularly, in November 1962 the first of what turned out to be five
mammoth skeletons was found, associated with more than 100 Clovis artifacts (Hester
1972). One of the mammoths shows definite evidence of butchering and three others were
probably or possibly butchered (Hester 1972). They are in different states of articulation and
preservation, and probably died at different times; they may have been attacked in water,
where movement would have been more difficult (Lundelius 1972). Reportedly, additional
mammoths were present at the 1962 find but were lost to bulldozing (Katz 1997). Humans
may have been both scavenging and actively hunting (Katz 1997). Blackwater Draw Locality
#1 became a National Historic Landmark in 1982.
14. Canaveral National Seashore (CANA), Florida
CANA museum collections contain teeth and a small bone fragment described as mastodon,
recovered from Merritt Island National Wildlife Refuge, located adjacent to CANA (Tweet et al.
2009a). No formal description or their chronology exists for the specimens.
15. Cape Krusenstern National Monument (CAKR), Alaska
CAKR was created to preserve an extensive sequence of ancient beach ridges with a
rich archeological record spanning the last four millennia, but also contains fossil-bearing
Quaternary deposits in the form of ice-rich frozen silt. Pockets of large mammal fossil material
are known from these yedoma deposits and often accumulate at the base of coastal
bluffs. A site at Imik Lagoon is the best documented example of such a fossil locality and
was recorded by NPS archeologists in 1987 (McClenahan and Gibson 1990). Here, a diffuse
scatter of Pleistocene fossil bones was noted along a 3 km stretch of the lagoon shore above
and below the waterline and at the base of the adjacent bluff. Among the finds are at least 8
Mammuthus sp. bones and tooth fragments. None were found in situ, although the bone is
reported to have been unweathered and in good condition.
Six specimens of human-modified Mammuthus ivory, presumably representing recent/
Holocene use of fossil ivory, exist in the museum collections of CAKR (Elder et al. 2009).
Little else is known about these specimens.
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16. Channel Islands National Park (CHIS), California
The Channel Islands are a series of islands off the coast of southern California. The
northern Channel Islands (Anacapa, San Miguel, Santa Cruz, and Santa Rosa) plus Santa
Barbara Island of the southern Channel Islands are included in Channel Islands National
Park (49 km [31 mi] off shore today from the city of Santa Barbara). The islands have
a long history of ranching and exploration (see Schoenherr et al. 1999). During the late
Pleistocene glacial episode, with lower sea levels, the four northern islands coalesced to
form one large island, Santarosae, which was a mere 7.2 km (4.5 mi) from the coastal
mainland (Muhs et al. 2015). The islands have been prospected for paleontological and
archeological remains since the second half of the 19th century, yet much remains to be
discovered, analyzed, and published.
Mammoths were first reported from the islands by Stearns (1873).The islands are famous
for their endemic pygmy mammoth (Agenbroad 1998a, b; Roth 1996), originally
described as Elephas (Mammuthus) exilis (Stock and Furlong 1928, Pygmy mammoth).
Most of the remains are found as scattered and isolated skeletal elements but in 1994 a near
complete skeleton was recovered (Agenbroad 1998b, Agenbroad et al. 1999). Although
small in stature, M. exilis were highly variable in size (Roth 1993). The islands also have
mammoth skeletal remains attributed to the mainland-sized form, M. columbi (Pigati et al.
2017). There is a long debate on the possibility of human association with the pygmy mammoth,
developed at great length by Phil Orr in the 1950s and 1960s (Agenbroad et al. 2005,
Orr 1968). The diet of M. exilis has been reconstructed (Semprebon et al. 2016, Smith and
Desantis 2018).
Pygmy or dwarf proboscideans are not limited to the Channel Islands of California. They
have been recovered from various islands in the Mediterranean basin (Elephas falconerii,
Mammuthus creticus, and Palaeoloxodon spp.; Herridge and Lister 2012, Palombo 2007),
Wrangel Island (Russia; Mammuthus primigenius; discussion in Tikonov et al. 2003), and
various islands of South-East Asia (Elephas, Stegodon [Stegodontidae] Van den Bergh et al.
1996).
17. Colonial National Historical Park (COLO), Virginia
One of the earliest potential reports of proboscideans for a NPS unit comes from the Williamsburg
area. The Williamsburg-area mastodon was first reported in 1811 (Anonymous
1811). The details vary from report to report, but apparently the bones were found 10 km (6
mi) south (Anonymous 1811) or east (Mitchill 1818) of Williamsburg on the south bank of
the York River. It is unclear where the former location would be, but the latter is potentially
within COLO, in the vicinity of Bellefield Plantation and the mouth of Indian Field Creek.
Anonymous (1811) reported that the site was a few yards within high water near the home
of Gawin Corbin. The fossils include 2 tusks, 2 vertebrae, 1 pelvis, 1 femur, and partial mandibles
with 7 associated teeth (Mitchill 1818). Given the presence of molars, it is surprising
that Mitchill (1818) identified the specimens as mammoth, yet Hay (1923) reported them
as a mastodon (Mammut). Hay (1923) reported that the bones were probably destroyed in
the 1859 fire at the College of William and Mary. Clark and Miller (1912) refer this specimen
to the Pleistocene of the Talbot Formation (a now-obsolete name). See more historical
discussion in Tweet et al. (2014a).
18. Colorado National Monument (COLM), Colorado
A proboscidean tooth was found in 1965 in No Thoroughfare Canyon, but has apparently
been lost (Scott et al. 2001, Tweet et al. 2012b). No formal write-up exists.
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19. Death Valley National Park (DEVA), California and Nevada
Proboscideans are represented in the Pliocene and Pleistocene deposits of DEVA. The
Pliocene specimens are all tracks, found in the Copper Canyon Formation (Fig. 5). Some
of the key references for the Copper Canyon tracks include Scrivner (1984), Scrivner and
Bottjer (1986), Santucci (1998), Santucci and Nyborg (1999), and Nyborg (2011).
Pleistocene proboscidean skeletal remains are known from the “Rogers Beds” (formerly
called “Lake Rogers”), including a roughly “3-foot” tusk of a “mastodon” reported in an abstract
by Clements (1952:1324). Based upon known faunas from the region, Mammuthus is
more likely than Mammut. The “Rogers Beds” were long interpreted as lacustrine in origin
but have recently been investigated in more depth (Springer and Pigati 2018, Springer et al.
2018a) and have been reinterpreted as “spring-derived desert wetlands” such as are found
at TUSK (Tule Springs Fossil Beds National Monument, see below). These latter authors
have found abundant fossils including Mammuthus.
20. Denali National Park and Preserve (DENA), Alaska
Mammuthus fossils have been briefly reported from the Teklanika River area of eastern
DENA (Blong 2018, Guthrie 1985, Ten Brink and Waythomas 1978). In 2018, NPS staff
archeologist Jess Peterson discovered in situ in DENA a mammoth tusk preliminarily identified
as Mammuthus primigenius (Fig. 6). Subsequent excavation also yielded a M3 and
skeletal elements of other taxa. Work on the project is currently on-going.
21. Florissant Fossil Beds National Monument (FLFO), Colorado
Although FLFO is known for its plant and insect fossils from Eocene lacustrine shales,
in 1996 a Mammuthus columbi specimen was excavated from a Pleistocene gravel deposit
within FLFO (Meyer and Weber 1995). This specimen, a mandible with molars, has been
radiocarbon dated to 49,830 ± 3290 yr. BP (CAMS-22182; Veatch et al. 2004). Pollen as-
Figure 5. Proboscidean trackway in Copper Canyon, DEVA (photo courtesy NPS).
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sociated with the remains were reported in Veatch et al. (2010). Significantly, this discovery
shows that the mammoth was inhabiting a high elevation (~2500 m; 8200 ft) prior to the
Last Glacial Maximum.
22. Gates of the Arctic National Park and Preserve (GAAR), Alaska
Gates of the Arctic is a large, roadless, remote, and mountainous park in northern Alaska,
which encompasses some 8 million acres of the Brooks Range Mountains and spans both
arctic and subarctic environments. One of the richest Pleistocene vertebrate fossil localities
in Alaska (Mann et al. 2013) lies immediately north of GAAR along north-flowing drainages
such as the Ikpikpuk River, which cuts through fossil-bearing and permanently frozen silt
deposits (yedoma) that in rare cases include mummified soft tissue (Guthrie and Stoker 1990).
Local Inupiaq residents are familiar with these abundant Pleistocene fossils and mammoth
ivory and bone has been used traditionally as a raw material for tools and crafts. Encounters
with mammoths and other giant animals are recounted in Inupiaq oral histories (Gubser
1965:33, Rasmussen 1932), and may be informed by fossil finds. Proboscidean specimens in
GAAR that are identifiable to the species level are all woolly mammoth (Mammuthus primigenius)
and represent the farthest north examples of the taxa within the NPS.
In 1885 Allen noted a well-preserved “os pubis” of a mammoth on the Alatna River, to the
south and outside of GAAR, but in the same fossil-bearing Quaternary deposits that extend
into GAAR. Observers noted the fresh appearance of the bone, which “to all appearances had
never undergone any process of petrification” (Allen 1887, Smith and Mertie 1930:252).
In 1964 a large segment of a single mammoth tusk was found protruding from the ground
surface during the course of archeological surveys on Fortress Creek in the northernmost,
Arctic foothills, portion of the park (Schlesier 1967:210). The undated specimen was excavated
and collected. No artifacts or demonstrable links to human activity were noted.
Figure 6. Excavation by Jess Peterson of the first mammoth tusk, Mammuthus primigenius, from
DENA, 2018 (photo courtesy NPS).
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In 2006 local residents found a complete woolly mammoth tusk on private lands within
the boundaries of GAAR. NPS researchers were permitted to sample it and AMS dating
of the collagen fraction yielded radiocarbon ages of 12,620 ± 90 BP (CAMS-131222) and
12,500 ± 60 BP (BETA-226405), demonstrating this to be one of the more recent mammoth
finds from Alaska. The specimen also has significance for its ecological setting—a
relatively high altitude, mountainous locale—distinct from most mammoth find spots in the
region, which are typically in lower elevation open terrain.
23. Glen Canyon National Recreation Area (GLCA), Utah and Arizona
GLCA holds some of the most unusual proboscidean remains in the way of dry-preserved
dung. The best-known and most studied late Pleistocene-age locality for mammoth
in GLCA is Bechan Cave (a Navajo word for “big feces”). All of the “dung localities” in
GLCA including Bechan Cave are rockshelters eroded into sandstone by incising river
channels. A test trench excavation by L. Agenbroad and J. Mead in the early 1980s established
that the Bechan shelter sediments contain a layer of over 300 m3 of dung, predominantly
Mammuthus dung (Agenbroad et al. 1989). The chronology was established using
radiocarbon dating (Agenbroad and Mead 1989, Martin 1987, Mead and Agenbroad 1992).
The identification of the large boluses and their coarse contents was established in detail via
morphology (Mead et al. 1986; Fig. 7), which has since been supported via aDNA (Karpinski
et al. 2017). Much of the attention of the dung has been on the dietary reconstructions
based on pollen, spores, and macro-botanical remains (Davis 1987, 1990, 2003, Davis and
Shafer 2006, Davis et al. 1984, 1985, Mead et al. 1984, 1986, Mead and Agenbroad 1989).
Other mammalian taxa also produced dung recovered from the layer (Kropf et al. 2007).
Additional, smaller sandstone shelters in GLCA have also produced dung of mammoth and
Figure 7. Dung of Mammuthus sp. from
Bechan Cave, Utah. A, overall morphology;
B, close-up showing the graze species, the
large number of the long plant fragments
illustrates the poor chewing abilities of
mammoths. The brown outer surface includes
bile acids that record DNA and other
biochemical data that has now been assessed
for aDNA (see text) (photo and permission
by S. L. Swift).
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other mammals (Mead and Swift 2012, Withers and Mead 1993). Mammoth bones have also
been reported from GLCA, but not in detail (Mammoth Alcove; Agenbroad and Mead 1989,
Tweet et al. 2009b). Santucci et al. (2001) and Tweet et al. (2009b) provided detailed overviews
about the work and contract reports conducted at Bechan Cave and other rockshelters
in GLCA that preserve mammoths and other Pleistocene taxa.
24. Golden Gate National Recreation Area (GOGA), California
Proboscideans are reported from two stratigraphic units at GOGA: the upper Pliocenelower
Pleistocene Merced Formation and the upper Pleistocene Colma Formation. Two
fragments of Mammut americanum were recovered from the Merced Formation at Fort
Funston (Fleishhacker Beach) in 1939 and 1975 (by T. Hall; see Henkel et al. 2015; Fig. 8).
P.E. Peabody collected Mammuthus tooth, tusk, and vertebra from the Merced Formation
at Mussel Rock; the Thornton Beach locality of the same unit has produced molar fragments
of Mammuthus. Hunter et al. (1984) suggested that the sedimentary convolutions in
the T2 unit of the upper Merced Formation between Wood’s Gulch and Fort Funston were
produced by mammoths and other large mammals. Hay (1927) reported observing a right
M2 of Mammuthus columbi in the collections of Golden Gate Memorial Museum of San
Francisco, collected from Cliff House Beach in 1908. Dooley et al. (2019) indicated that
the newly-recognized Mammut pacificus pertains to all mastodon specimens from the Sierra
Nevada west to the Pacific Coast and into southern Idaho, and that M. americanum is not
found in California. These specimens referred to from this park should be reassessed based
on the Mammut analysis of Dooley et al. (2019).
25. Grand Canyon National Park (GRCA), Arizona
The Grand Canyon contains the largest number of caves in the National Park System.
Due to the steep physiographic setting for most of this region, it is understandable that
remains of graviportal proboscideans are rare in this park. Caves along the Marble Canyon
Figure 8. Mammut molar from GOGA (photo courtesy NPS). Although originally identified as belonging
to M. americanum, analyses by Dooley et al. (2019) could alter this—see text.
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platform of the upper Colorado River area within the eastern Grand Canyon were systematically
prospected and select caves excavated in the early 1980s. Interestingly, Emslie (1987,
1988) reported a fragmented tooth plate of Mammuthus from the condor (Gymnogyps) nest
deposit in Sandblast Cave. Prior to this discovery, mammoths were not known from cave or
alluvial deposits within GRCA (Mead 1981). It appears that mammoths were not within the
steep corridor canyons within GRCA but were on the flat platforms adjacent to and forming
the rims to the river corridor and side canyons. Northwestern reaches of the Grand Canyon,
within the Grand Canyon-Parashant National Monument (PARA) and GRCA (portions of
the Shivwits Plateau) have yet to be prospected for paleontological remains in caves and
sinkholes that proboscideans could have visited, similar to those at Bechan Cave and others
in GLCA mentioned above.
26. Great Sand Dunes National Park and Preserve (GRSA), Colorado
Although mammoth remains from GRSA have not received official description or
chronology in a publication, Scott et al. (2001) mention that a Mammuthus femur has been
collected from the park.
27. Hagerman Fossil Beds National Monument and National Natural Landmark
(HAFO), Idaho
HAFO is internationally significant because it protects one of the world’s richest known
fossil deposits from the late Pliocene epoch, 3.5 million years ago, providing a window on
the time just before the Pleistocene. A multitude of publications detail the various vertebrate
and invertebrate fossils recovered from HAFO and surrounding region (see review Mead
et al. 1998, Sankey 2002). The only proboscideans (Mammut) from HAFO come from the
middle Pliocene Glenns Ferry Formation (Blancan; Gazin 1936, Ruez 2009).
HAFO also curates an unusual “occurrence” from Craters of the Moon National
Monument and Preserve (CRMO). The excavation of a circa 1920s–1930s trash dump
revealed a number of large, charred, partially-burned Mammuthus bones near the bottom
of the trash layer. H.G. McDonald, head of the excavation, indicated that the mammoth
remains were not in situ and may have been brought to the dump from a nearby gravel
quarry (mammoth remains are relatively common in the Snake River Plain gravel pits of
Pleistocene age sediments).
28. Hopewell Culture National Historical Park (HOCU), Ohio
HOCU (formerly Mound City Group National Monument) has no definite in situ proboscidean
records; however, Mills (1922:286) reported “mastodon tusk” fragments from
Mound 23. Unfortunately the specimens are apparently lost and no formal write-up of the
remains exists (Hunt et al. 2008).
29. Independence National Historical Park (INDE), Pennsylvania
There is a historic record of a Mammut tooth fragment from INDE, from an archeological
excavation of an area near Benjamin Franklin’s property. The specimen is of historic interest
because Franklin received mastodon fossils and wrote about them in his journal. This specimen
is from an archeological context and may have been owned and written about by Franklin.
A second historic occurrence of proboscidean fossil remains are associated with INDE.
During the early 1800s, Charles Willson Peale’s natural history museum was located on the
second floor of Independence Hall. Among the exhibits on display at Peale’s museum was a
mastodon skeleton, as shown in his 1822 self-portrait “The Artist in His Museum” (Fig. 9).
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30. John Day Fossil Beds National Monument and National Natural Landmark
(JODA), Oregon
JODA records the longest paleontological record for the Cenozoic in the NPS system
(55–7 Ma) (Fremd 2010). The Mascall Formation (lower-middle Miocene) and the Rattlesnake
Formation (upper Miocene) at JODA are known to produce proboscidean remains,
which are among the oldest such remains in the NPS (see overview in Fremd 2010, Kenworthy
et al. 2005). Based on Fremd (2010), The Rattlensake Formation contains both Amebelodon
(Barbour 1927) and Tetralophodon (Falconer 1857). A tooth from the Mascall Formation
(Fig. 10) is the anterior portion of a molar which lacks central conules in the medial sulcus
and, therefore can be attributable to Zygolophodon, although listed as “Proboscidea Indet.”
in Maguire et al. (2018). This formation has both Gomphotherium and Zygolophodon (Fremd
2010). Fremd et al. (1994) and Prothero et al. (2006) indicate that Gomphotherium is known
from the Mascall sediments but the study by Maguire et al. (2018) did not find evidence for
this taxon in the type area of the Mascall Formation. The fossils curated at JODA also contain
proboscidean remains from outside the park boundaries.
31. John Muir National Historic Site (JOMU), California
JOMU has a single tusk cataloged into its historic collection. John Muir collected the tusk
in Alaska in 1881 (Kotzebue Sound, Eschscholtz Bay), and discussed the find in his book “The
Figure 9. “The Artist in His
Museum”. Oil on canvas
by Charles Willson Peale,
1822. Courtesy of the Pennsylvania
Academy of the
Fine Arts, Philadelphia.
Sarah Harrison (The Joseph
Harrison, Jr. Collection)
(INDE).
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Cruise of the Corwin” (Muir 1917). The tusk is a 30 cm (12″) section at the alveolar end with
the other end sawed flat (V. Bones, JOMU, San Francisco, CA, December 2018, pers. comm.).
It appears to be Mammuthus primigenius.
32. Joshua Tree National Park (JOTR), California
Proboscidean fossils at JOTR are from the informally designated upper Pleistocene
“Pinto Formation” in Pinto Basin, in the eastern section of the park. The “Pinto Formation”
includes lacustrine, fluvial, alluvial, and eolian sediments deposited in a time frame
including at least 25,000–16,500 years ago. Fossils attributable to Mammuthus columbi and
Mammuthus sp. have been found since 2005, but have not yet been described in detail. See
also discussions in Scott et al. (2006) and Tocci et al. (2018).
33. Klondike Gold Rush National Historical Park (KLGO), Alaska
KLGO has curated an artifact pendant made of fossil Mammuthus ivory. It was a gift to
the NPS from the Moore family, descendants of the Moores who built the historic Moore
House (now KLGO property). Herman Kirmse, the first jeweler in Skagway, moved into the
Moore House about 1908. Kirmse made the pendant out of Alaskan mammoth ivory, but the
original sources of the fossil is unknown (D. Boettcher, KLGO, Skagway, AK, December
2018, pers. comm.).
34. Kobuk Valley National Park (KOVA), Alaska
There are several reports of Mammuthus from KOVA. Several specimens of Mammuthus
primigenius and Mammuthus sp. have been reported from the important and well-researched
stratigraphic section at Epiguruk Bluff along the south bank of the Kobuk River (Hamilton
et al. 1993). Although most of the bluff, including the documented mammoth fossils is located
just outside the park boundary, similar Quaternary alluvial deposits with scattered fos-
Figure 10. Zygolophodon sp. from JODA (photo courtesy NPS and Nicholas A. Famoso).
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sil material extend inside the park and are exposed along the Kobuk River. USGS geologists
working on the Kobuk River through the 1970s and 80s recovered numerous Pleistocene
vertebrate fossils, including several Mammuthus sp. specimens from these contexts (Repenning
1982). Elder et al. (2009) mentioned finds of a Mammuthus primigenius lower molar
from along the Kobuk River (from Epiguruk Bluff outside unit boundaries but important to
know); a fragment of M. primigenius tooth is in park collections. Giddings (1962:11), in a
description of the Onion Portage site on the Kobuk River, reported finding a “broken fishshaped
fishing lure of mammoth tooth”, a clear record of Holocene people making use of
Pleistocene-age fossils.
Along the lower and middle reach of the Kobuk River as far up stream as the village
of Shungnak—a portion of the river that spans KOVA—Cantwell (1887, 1889) observed
ice-rich Quaternary deposits in high bluffs along river banks that included many “elephant”
(Mammuthus sp.) tusks, teeth and bones. “Quantities of mammoth tusks were observed in
this clay and its debris where undermined by the stream”. He observed that in many of the
villages on the Kobuk River fossil ivory in use by local residents to carve ornaments and
tools, including large soup ladle.
A brief survey for archeological and fossil finds in Kobuk sand dunes has been conducted,
with an emphasis on Pleistocene age deposits. Work that followed identified bison and
mammoth paleontological remains, which remain undated and not published on (Stanford
et al. 1990).
35. Lake Mead National Recreation Area (LAKE), Arizona and Nevada
The first discovery of Mammuthus in LAKE was made by John Strong Newberry on the
1857–1858 J. C. Ives expedition to the Colorado River. Newberry found a mammoth maxillary
tooth at the base of a pyramidal hill (now dubbed “Elephant Hill”) on the east side of the
Colorado River (Hay 1927:45, Newberry 1861; whereabouts of specimen unknown). This
fossil is usually attributed to the Chemehuevi Formation (Longwell 1963), but may have
come from an underlying unit instead (Malmon et al. 2011). Numerous additional reports
and discoveries of mammoth specimens are reported from LAKE but are only briefly described;
further information can be found in the following: Agenbroad and Brunelle (1993),
Bonde et al. (2018), Hay (1927), Longwell (1963), and Malmon et al. (2011). LAKE also
curates late Pleistocene material, including Mammuthus, from the Glendale local fauna
which was collected just outside of the park along the south side of the Muddy River at
Glendale, Nevada (Scott et al. 2004).
36. Lake Meredith National Recreation Area (LAMR), Texas
LAMR has at least two records of proboscideans. Hunt and Santucci (2001) reported that
a now-lost proboscidean tooth was recovered from the Ogallala Formation in Cedar Canyon
near the eastern end of LAMR during the placing of a restroom pipe. The fossil was initially
identified as a Mammut tooth but was probably from a gomphothere instead. It came from
the medial sandy interval of the Ogallala Formation. In addition, a proboscidean humerus
tentatively identified as Mammuthus was found in a cliff of alluvium adjacent to an oil well
in an inset terrace just west of LAMR’s boundary with Alibates Flint Quarries National
Monument (Hunt and Santucci 2001).
37. Lava Beds National Monument (LABE), California
Many Mammut specimens have been recovered from lava tube caves (Fossil Cave
particularly) at LABE (Fig. 11). The park collections contain a number of these remains,
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some of unknown provenience (see details in Kast 2008, Santucci et al. 2001, Santucci and
Kenworthy 2009). These specimens should be reassessed based on the Mammut analysis of
Dooley et al. (2019).
38. Lehner Mammoth-Kill Site NHL (LEMA-AZ), Arizona
While Emil Haury (1953) excavated mammoths from the Naco Mammoth Site in
southeastern Arizona (Haury 1953), mammoth remains were also found exposed not far
away at the Lehner Ranch, Cochise County (Haury et al. 1959). Since the excavations in
the middle 1950s and subsequent field seasons in 1974 and 1975, a multitude of geological,
paleontological, paleoecological, and archeological research has been conducted at
the locality and other sites along the San Pedro River Valley (Mehringer and Haynes 1965,
Metcalfe et al. 2011). All research at this locality is decisively linked to the work at the
Murray Springs Clovis Site (# 44 below; Haynes and Huckell 2007) and other localities
in the valley. The Lehner locality contained the remains of Mammuthus columbi (Lance
1959, Saunders 1970) and Mammut (Mead et al. 1979) recovered from late Pleistocene
sedimentary units. The locality is administered by the Bureau of Land Management and
was designated a NHL in 1967.
39. Mammoth Cave National Park (MACA), Kentucky
Fittingly, MACA has proboscidean fossils. Ron Wilson (1981, 1985) reported finding
material of a mammoth or mastodon in 1979 within the Proctor section (aka Proctor Cave)
of Mammoth Cave (Santucci et al. 2001). Per an unpublished note by Wilson (in NPS paleo
archives) on fossils collected from MACA and reposited at the Carnegie Museum of Natural
History, the 1979 proboscidean material was only a possible metapodial fragment, with
Figure 11. Mammut americanum
molar (partial) located in
the UCMP collections from
LABE (photo courtesy NPS).
Although originally identified
as belonging to M. americanum,
analyses by Dooley et al. (2019)
could alter this—see text.
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additional proboscidean material collected in 1980 which consisted of 20 fragments of a
tusk. Later, Dan Fisher (University of Michigan) identified the tusk fragments as Mammut
americanum (Colburn 2017).
40. Mammoth Site of Hot Springs (MASI-SD), South Dakota
The Mammoth Site of Hot Springs (a National Natural Landmark and a 501 (c)(3) nonprofit
organization; not administrated by the NPS) is located in the southern Black Hills
of southwestern-most South Dakota. A hill on the southern side of the town was slated to
be removed for a housing project in 1974. Earth removers immediately uncovered skeletal
and ivory remains and halted excavation. Larry Agenbroad and Jim Mead were called in
to evaluate the fossils and declared in the summer of that year that Mammuthus remains
were in situ and abundant. The locality is a filled-in sinkhole measuring about 25 by 50
m (82 by 165 ft). Coring of the deposit indicates that at least 20 m (67 ft) of sediments
and mammoth bones fill the sinkhole, which contained a warm-water pond for much of its
existence. Steep-sided edges of the sinkhole made the pond a lethal trap and for the most
part just a mammoth-selective trap. Excavation in the 1970s and 1980s produced most of
the exposed and recovered skeletal remains. By the middle 1980s it was determined that
the site should be preserved and made into an active, nonprofit research and educational
institution. By 1986 the entire sinkhole was within a building, now with an attached museum,
archival lab, and preparation lab (see discussions in Agenbroad and Mead 1994).
Radiocarbon analyses produced an age of approximately 26,000 yr. B.P. but that was
always assumed to be a minimum. In 2018 OSL (Optically Stimulated Luminescence)
analyses by two labs have shown the top of the sinkhole deposit dates to approximately
140,000 years ago and the lowest excavation produced an age greater than 190,000 years
old; there are more than 14 m (45 ft) of sediments and fossils below that level to be dated.
A minimum of 60 mammoth individuals have been recovered based on the number of
tusks (two per individual in elephants). Currently both M. columbi and M. primigenius (3
specimens) are reported from the deposit. The locality is a working research institution
with excavations occurring at various times throughout the year but especially during the
summer. Tours are given all year long to approximately 102,000 visitors per year.
41. Mississippi National River and Recreation Area (MISS), Minnesota
Several finds of isolated proboscidean fossils have been made in and around the “partnership
park” MISS (Stauffer “1945”, Tweet 2014). Stauffer produced an inventory of
Pleistocene finds in Minnesota through 1948 (hence the quotation marks around 1945, the
year usually given for the citation). In the report there are three finds that are most likely
from land now within MISS, and three potentially from within MISS.
The three records likely from MISS are: 1) a tooth and tusk found in 1878 in terrace gravel
along Coon Creek on the east bank of the Mississippi River attributed to Mammuthus columbi
(“Parelephas jeffersoni” of Stauffer “1945”); 2) a fragmentary Mammuthus primigenius molar
dredged from the Mississippi River near St. Paul in 1924; and, 3) the probable head of a mammoth
femur from the terrace gravel at Hasting. The Hastings specimen was probably found
in 1909 rather than 1923 as stated in Stauffer (Tweet 2014). Stauffer’s Coon Creek report is
broadly similar to a report in Johnson (1874) of tusk fragments and a tooth in drift 8 km (5 mi)
above St. Anthony Falls on the east bank of the Mississippi, but differs in many details, such
that it is improbable that they are the same unless Stauffer had a different information source.
Three other finds cited by Stauffer (“1945”) have such general provenance information
that it is impossible to know whether or not they were found in the river corridor: a proEastern
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boscidean tooth found at an unstated time in the glacial drift of Minneapolis; a Mammuthus
columbi molar from glacial gravel within Minneapolis, now at Yale (collected in 1895); and
another Columbian mammoth molar from glacial deposits near Hastings. As noted by Hay
(1924), the Yale specimen may be referred to obliquely in Lull (1908:198).
42. Mojave National Preserve (MOJA), California
Based on NPS records, three localities in MOJA have yielded fossils attributed to Mammuthus
sp. All three are near the southwestern boundary of the park; see discussions in
Jefferson (1991), Reynolds et al. (2003), and Tweet et al. (2016).
43. Montezuma Castle National Monument (MOCA), Arizona
MOCA has one of the oldest proboscidean records in the NPS. Fossil proboscidean tracks
(Fig. 12) are known from the middle Miocene–Pliocene Verde Formation within the monument,
at a locality known as “Elephant Hill” (#33 in Twenter 1962) (Tweet et al. 2008, Santucci
et al. 2014). The site has been mapped by McGeorge and Schur (1994). Although known
about earlier, Brady and Seff (1959) were the first to report these presumed proboscidean
tracks. They have been covered several times with soil to protect them, and are currently buried
(Protas 2002). Czaplewski and Davies (2007) questioned the identification of the tracks.
44. Murray Springs Clovis Site, NHL (MUSP-AZ), Arizona
Murray Springs (archeological site AZ EE:8:25 ASM; administered by the Bureau of
Land Management and designated a NHL in 2012) is a Clovis cultural site with multiple
activity areas in the San Pedro River Valley of southeastern Arizona (Haynes and Huckell
2007). In 1966 Vance Haynes and Peter Mehringer found multiple areas along the dry arroyo
exposure while working to understand the valley sedimentary deposits. As with the
nearby Lehner Mammoth Kill Site (#38 above), skeletal fossils were under and in contact
with a distinctive black mat. Among a number of megafaunal species, Mammuthus was
Figure 12. Proboscidean track at MOCA (photo courtesy NPS).
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abundant. All bone was highly leached of organics; enamel and dentine components of
teeth were typically well preserved. Amongst the many lithic artifacts recovered was a
bone shaft straightener constructed of mammoth bone (Haynes and Hemmings 1968: fig.
1, Hemmings 2007). An average age (of eight samples) of 10,900 ± 50 years B.P. (not corrected)
was determined for the Clovis occupation and utilization of mammoth and other
megafaunal species (Haynes and Huckell 2007). Multiple reports about local and regional
paleoecological and geological analyses are presented in Haynes and Huckell (2007).
Multiple skeletons and footprints of Mammuthus columbi were recovered (Saunders 1970,
Hemmings 2007).
45. Natchez Trace Parkway (NATR), Alabama, Mississippi, and Tennessee
Archeological investigations undertaken during the completion of the NATR uncovered
three fragments of proboscidean teeth (C. Smith, NATR archeologist and museum curator,
Tupelo, MS, February 2018, pers. comm.). Two are identified as mastodon, and the third
enamel fragment is identified as mammoth or mastodon. The small size of the specimens
and the presence of a small creek nearby suggest a detrital origin. No details or descriptions
exist for these remains.
46. Navajo National Monument (NAVA), Arizona
There are two non-specific references to Mammuthus finds in the vicinity of two of the
three units of NAVA in Agenbroad and Mead (1989). “#17 Tsegi” (Agenbroad and Mead
1989: Table 1) refers to Tsegi Canyon with a description of the sediments from Betatakin
Ruin to Marsh Pass at the canyon mouth of Laguna Creek at the community of Tsegi.
Original description of the canyon sediments is attributed to Hack (1942) but this author
does not mention proboscidean remains in the discussion of the Tsegi and Jeddito formations
in the canyon. However, Saunders (1970: JJS 3 site 50) does indicate one Mammuthus
find from the canyon mouth at Tsegi. Mammuthus remains are found throughout the
region around the various NAVA units, including in the canyon near Inscription House
(Agenbroad and Mead 1989, Agenbroad et al. 2013); unfortunately no formal description
of the NAVA specimen exists.
47. New Jersey Pinelands National Reserve (PINE), New Jersey
PINE is the first National Reserve and is administered as a partnership between the
NPS and the State of New Jersey. At least one Mammut find can be attributed to PINE. The
area around Pemberton, at the northwestern boundary of the unit, has produced a number
of Mammut specimens, but most can be placed outside of PINE. Rogers (1840) mentioned
bones including a skull, tusk, tibia fragments, and ribs found north of the road between
Pemberton and (New ?) Lisbon and 2.4 km (1.5 mi) from the latter, on the Forsyth property.
“New Lisbon” and “Lisbon” were used interchangeably at the time, and New Lisbon is well
within PINE.
A find mentioned by Heilprin and Leidy (1887), although superficially similar, is not
the same as Rogers’ (1840) mastodon because the former specimen was recovered in 1877
(Rhoads 1903, Hay 1923). Kitchell (1856) added another mastodon skull (found in 1855)
from John Ewens’s meadow. Rhoads (1903) included accounts of two mastodon specimens
from Pemberton, including Heilprin and Leidy’s specimen and a second skull from a swamp.
Hay (1923) reported several occurrences, although all of the finds where the locality can be
distinguished are outside of PINE. Hay did not include Rogers’ report, but did mention that
the Academy of Natural Sciences of Philadelphia (now the Academy of Natural Sciences
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of Drexel University) had two complete molars and two partial molars from Pemberton,
credited to G.C. Forsyth, which could be part of the 1840 find.
48. Nez Perce National Historical Park (NEPE), Idaho, Montana, Oregon, and Washington
NEPE is composed of 38 small units across four states (Idaho, Montana, Oregon,
and Washington), of which six are on NPS land. Mammuthus fossils have been found at
1 of the other 32 sites, Tolo Lake, Idaho. This site is managed by the Idaho Department
of Fish and Game (Kenworthy et al. 2005). In the fall of 1994, heavy equipment operators
unearthed mammoth bones, which prompted an excavation in the summer of 1995.
More than 400 well-preserved mammoth fossils, attributed to M. cf. M. columbi, were
recovered, representing at least 10 subadult to adult individuals. Among the fossils was
a ~95% complete skeleton, perhaps the most complete mammoth found in Idaho (Miller
et al. 1998). The excavations sampled a small fraction of the encasing lake bed, so it is
likely that there are more mammoth remains still in situ. The lake has since been refilled.
The primary repository is the Idaho Museum of Natural History in Pocatello, with some
specimens at the University of Idaho (Moscow) and the Grangeville Chamber of Commerce
Visitor Center (Kenworthy et al. 2005).
49. Niobrara National Scenic River (NIOB), Nebraska
Verifying the provenience of records in the Niobrara River corridor can be challenging,
because NIOB boundaries provided by various sources are inconsistent, and the history of
research goes back to the mid-19th century, when standards for reporting locality information
were more relaxed. Many of the oldest publications simply report “Niobrara Valley” as
the locality for various discoveries. An example of this issue can be found in Leidy (1858),
in which he named two species from the Niobrara Valley, Mastodon (Tetralophodon) mirificus
(now Stegomastodon mirificus, Leidy 1858) (“Bed F”, possibly the “Loup Fork Beds”)
and Elephas (Euelephas) imperator (now a synonym of Mammuthus columbi). Detailed
locality information is lacking, so it is unclear where along the valley these species were
found, therefore it is unknown if they are from NIOB.
Multiple reports of proboscideans of Miocene and Pleistocene age can be confirmed
within NIOB, which is one of the most productive NPS units for proboscideans. For example,
Voorhies (1990) provided specimen lists and other information for a number of
paleontological localities over roughly a quarter of what became NIOB in 1991, from approximately
Allen Bridge to just downstream of the Norden Dam site. In this section of
NIOB, Voorhies (1990) reported 12 Miocene sites and 8 or 9 Pleistocene sites (one site is
very close to the park boundary) that had yielded proboscidean fossils. The great majority
of these finds were tooth fragments, but they illustrate the pot ential of the corridor.
Reports of Miocene proboscideans in NIOB in the literature pertain to the Valentine
Formation. Almost all of the specimens appear to represent gomphotheres, although a tooth
and possibly a radius from the Norden Bridge site, low in the Valentine Formation, are attributable
to the mammutid cf. Zygolophodon sp. (Voorhies 1990). Per Holman (1977) and
Skinner and Johnson (1984), the Kuhre site (Cornell Dam Member) has a vertebra of cf.
Serridentinus (now Gomphotherium). Of the gomphothere remains, most are indeterminate
beyond Gomphotheriidae, although some Norden Bridge site specimens appear to represent
a large species of Gomphotherium comparable to G. osborni (Voorhies 1990). The most
significant Miocene proboscidean find from NIOB (Rockford or Rocky Ford site) is the
holotype of Tatabelodon gregorii (Frick 1933; a palate and partial mandible; now Gomphotherium).
Voorhies (1990) reported finding additional proboscidean remains at the site
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that may represent the same individual as reported by Frick (1933). These fossils are at the
University of Nebraska State Museum and include the neural arch of an axis, proximal ulna,
distal femur, metapodial, phalanx, and lunar.
The Pleistocene specimens (almost entirely tooth fragments) were found in unnamed
high terrace deposits at various locations overlooking the Niobrara River. The fossils all
appear to represent Mammuthus, but Voorhies (1990) could not assign any to species.
50. Noatak National Preserve (NOAT), Alaska
NOAT lies in northwestern Alaska and encompasses the northernmost extent of the Rocky
Mountain chain, the Brooks Range Mountains. It was established to preserve ecosystems and
resources of the Noatak River Basin, one of the continent’s largest mountain-ringed basins.
Mammuthus sp. (likely Mammuthus primigenius) fossils are moderately common in this area,
and appear to be concentrated in Pleistocene age valley bottom sediments where they are typically
found after having been eroded by river action and accumulated in secondary contexts.
USGS explorations in the early 1900s were the first to note fossil material in the Noatak River
valley. Smith and Mertie (1930:252) note “large pieces of mammoth tusk … found in the Quaternary
gravels about 20 miles [32 km] downstream from the mouth of the Cutler River” and
parts of a skull and thigh bones of mammoth along the Noatak River between the Nimiuktuk
and Kugururok Rivers. They also encountered Alaska Natives who reported “finding a good
deal of this old ivory on the Noatak” (Smith and Mertie 1930:252).
The use of fossil mammoth ivory has historic depth and occurs in prehistoric archeological
sites dating to the last millennia. Hall (1971), for example, reported 133 fragments of mammoth
ivory from the 16th century Kangiguksuk site near the confluence of Kangiguksuk Creek
and Noatak River. Twenty-six specimens show signs of cutting, including one large piece
from a transverse cut across a tusk. Among the remains were three pieces of mammoth ivory
fashioned into single or multi-pronged fish spear tips, another drilled to serve as a harpoon
socket, three fragments that may have been unfinished berry spoons, and five unidentified
worked pieces. A single unmodified mammoth tusk fragment and nine pieces of mammoth
tooth were also identified. Radiocarbon dating of other extinct Pleistocene fauna from Kangiguksuk
(Bison priscus) yielded ages beyond 30 kbp, demonstrating that the site occupants
obtained these materials long after the animal’s death (Rasic and Matheus 2007), which is
certainly also the case for the mammoth specimens. Hall (1971) reported that mammoth
ivory and other remains of extinct animals were frequently washed from the banks when the
Noatak River shifted its course, and that two tusks were found while he was in the field. The
Maiyumerak Creek Site near the confluence of Maiyumerak Creek and the Noatak River is
another Late Prehistoric age Inupiat site in the Noatak basin, in this case approximately 500
years old. At this locality the finds include a waterworn piece of mammoth ivory evidently
procured from a river gravel bar (Shirar 2009).
NOAT museum collections include a “poorly preserved mammoth tusk identified as
Mammuthus primigenius” (E. Devinney, Western Arctic National Parklands, Kotzebue,
Alaska, November 2009, pers. comm.) (Elder et al. 2009).
51. Olympic National Park (OLYM), Washington
The only proboscidean remains associated with OLYM are those curated in their park
museum. OLYM has five mammoth fossils in its collections, which include four specimens
of Mammuthus sp. molars and one tusk. Three of the molars have no locality information.
One molar is from a gravel pit on Morse Creek near Highway 101 and the tusk is from
Whiskey Creek, both near but outside of OLYM boundary (Fay et al. 2009).
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52. Padre Island National Seashore (PAIS), Texas
PAIS does not have in situ proboscidean fossils, but mammoth teeth are known to wash
into the park from Seven and One Half Fathom Reef a couple of miles offshore. Some specimens
have been entered into PAIS museum collections (Kenworthy et al. 2007).
53. Potomac Heritage National Scenic Trail (POHE), West Virginia, Virginia, Maryland,
and District of Columbia
Mammut fossils have been recovered from Cumberland Cave (Gidley and Gazin 1933,
1938), which hosts a middle Pleistocene (Irvingtonian Land Mammal Age) faunal assemblage,
rare for the eastern half of the United States. The site is northwest of Cumberland,
Maryland, adjacent to the former Western Maryland Railway along POHE. The site was
first brought to the attention of paleontologists in 1912 when it was encountered during
construction of the railroad. Mammut remains are uncommon in the cave, presumably
because the sinkhole entrance was too small for adult proboscideans to enter. Gidley and
Gazin (1938:70) reported four juvenile molars (“second, third, and fourth left, and third
right of the lower-jaw”) and a tibia without epiphyses, all consistent with Mammut americanum.
54. Rancho La Brea (RALA-CA), NNL, California
RLBR in southern California is a globally significant tar pit and asphalt deposit preserving
a rich and diverse assemblage of late Pleistocene animals and plants. The fossil locality
was designated a National Natural Landmark in 1964 and is managed in association with
the George C. Page Museum, Los Angeles County. The mammalian fauna from RALA-CA
exemplify the Rancholabrean LMA; the deposit is the type locality for the age. The Rancholabrean
LMA is defined by the first appearance of the bovid Bison in North America (Bell
et al. 2004).
The occurrence of fossilized remains of extinct fauna at RALA-CA was first noted by
Denton (1877). The composition of the mammalian fauna is dominated by the skeletal
remains of carnivores, representing approximately 90% of the identified elements (Marcus
1960, Merriam 1911, Stock 1929). Two extinct proboscideans have been documented at
RLBR including the Columbian mammoth (Mammuthus columbi) and the American mastodon
(Mammut americanum). Life-size reconstructions of both the mammoth and mastodon
are iconic displays exhibited in and around the on the grounds of Hancock Park at the La
Brea Tar in the Page Museum.
A relatively complete specimen of an adult male Mammuthus columbi was excavated
during the 2006 construction of an underground parking facility for the Los Angeles County
Museum of Art adjacent to the tar pits. This mammoth specimen nicknamed “Zed” was
determined to be between 48 and 52 years old at the time of its death approximately 37,000
years ago (El Aldi et al. 2015). Specimens from the Hancock Collection at RALA-CA were
used in the analysis and description of Mammut pacificus in Dooley et al. (2019).
55. Salinas Pueblo Missions National Monument (SAPU), New Mexico
Mammuthus remains have been found at two of the three parcels of SAPU. At the Abó
unit, eight mammoth tusk fragments were discovered and collected in 1987 by Fredrico
Sisneros, likely from north of the mission. These specimens are held at the Western Archeological
and Conservation Center (WACC) in Tucson (Thorpe et al. 2017).
The discovery of a mammoth at Quarai was first mentioned in Hibben (1941), who reported
that “a complete but disarticulated mammoth was excavated by the expedition near
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the ruins of Quarai”. The 1939–1940 excavations at Quarai, a joint Museum of New Mexico
and Work Projects Administration archeological and stabilization project, were documented
in detail by Hurt (1990), who stated only that during road construction “the bones of a mammoth
[were found]”. Until recently, these two passages were the only documentation of a
Quarai mammoth; all other reports (e.g., Morgan et al. 2001) lead back to these two sources.
Thorpe et al. (2017) contacted the Maxwell Museum of Anthropology at the University of
New Mexico, the Museum of Southwestern Biology at UNM, the New Mexico History
Museum, and the New Mexico Museum of Natural History and Science, but were unable to
find the fossils.
In October and November 2018, work in SAPU archives by Marc LeFrancois (SAPU
chief of facilities and resource management) uncovered photos of the mammoth excavation,
and an early draft of Hurt (1990) provided by Khaleel Saba (WACC) included a photo of the
discovery locality. On November 14, Marc and Ron Fields (SAPU integrated natural and
cultural resource specialist) were able to use the photo of the locality to place the locality on
the SAPU side of the entrance road into the Quarai unit. The SAPU staff have suggested that
the mammoth was uncovered but not excavated, and thus may still be in situ. Pleistocene
fossils, particularly those of mammoths, were reported as common near Punta de Agua (~1
km, less than a mile northeast of Quarai) by Hibben (1941).
56. Santa Monica Mountains National Recreation Area (SAMO), California
Proboscideans are known from Quaternary sediments in SAMO as mentioned in Koch
et al. (2004) and Tweet et al. (2012b), but have not been reported in detail in the literature.
The records consist of specimens in the collections of the Natural History Museum of Los
Angeles County (LACM), and at a minimum they include Mammuthus sp.
57. Tule Springs Fossil Beds National Monument (TUSK), Nevada
TUSK is a recent addition to the NPS, having been established in 2014. Proboscidean fossils
have been reported from the TUSK area since the early 20th century, when Josiah Spurr
(1903) noted the presence of “mastodon” (more likely mammoth) bones from between Corn
Creek Spring and Tule Springs. Archeologists became interested in the area in the 1930s due
to the possibility that human artifacts had been found with Pleistocene megafauna fossils,
but unequivocal evidence was not found (Harrington 1934). The development of radiocarbon
dating renewed interest in the area, and in 1962–1963 the Tule Springs Expedition crossed
the future area of TUSK with trenches, looking for artifacts and material that could be dated.
Although the archeological results proved to be disappointing, abundant fossils were encountered,
and the trenches permitted detailed stratigraphic work (Haynes 1967). Scientists from
the San Bernardino County Museum conducted extensive paleontological investigations from
the 1990s into the early 2010s, leading to the creation of TUSK.
Mammuthus columbi is the iconic animal of TUSK (Springer et al. 2018b). It is the most
abundant large mammal of the Tule Springs Local Fauna, although its relative abundance
may be somewhat overstated due to the size of adult mammoth elements and the relative
ease of identifying their remains, even when fragmented (Scott and Springer 2016, Scott
et al. 2017, Springer et al. 2017). Of definite interest is the recovery of neonate/juvenile
Mammuthus remains (Fig. 13). The host sediments, the Las Vegas Formation, were originally
interpreted as lacustrine, but have been re-identified as spring deposits (Springer et al.
2018c). Deposition occurred from approximately 573,000–8,530 years ago (Springer et al.
2018c), although the local fauna is restricted to approximately 100,000–13,000 years ago
(Springer et al. 2017).
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58. Valley Forge National Historical Park (VAFO), Pennsylvania
VAFO includes Port Kennedy Bone Cave, one of several notable Irvingtonian Land
Mammal Age bone caves in the eastern states. This cave, a sinkhole in the Cambrian Ledger
Formation, was encountered by quarrymen in 1870, with one of th e first recognized fossils
being a Mammut molar (Daeschler et al. 1993). Fossils were removed in 1870 and 1894–
1896, and were described by Cope (1871, 1895, 1896, 1899), Wheatley (1871), and Mercer
(1899). After this series of early publications the site sank into obscurity with its location
lost until investigations in the 1990s and 2000s (Daeschler et al. 1993, 2005). Proboscideans
at the cave are represented by Mammut americanum, notably “a high proportion of juvenile
and sub-adult individuals” (Daeschler et al. 1993:37).
59. Vicksburg National Military Park (VICK), Mississippi
Mammut and Mammuthus remains are common in the area in and around Vicksburg,
Mississippi. At least one occurrence of fossil proboscidean is potentially known from
VICK. Wailes (1854:284; referenced in Hay 1923:125) reported that mastodon fossils had
been found in the deep railroad cut at Vicksburg. As discussed in Kenworthy et al. (2007),
there is a large railroad cut within VICK for the Vicksburg and Meridian Railroad, near the
Railroad Redoubt, a Civil War site in the southeastern arm of the park. This cut exposes
loess deposits, known for Pleistocene fossils. It is possible, but not confirmed, that this is
Wailes’s proboscidean locality. VICK staff assisted in the initial study of a mastodon skeleton
discovered near the site of the Pemberton Square Mall, approximately 1.9 km (1.2 mi)
southwest of VICK (Knox and Pitts 1984).
Figure 13. Mammuthus columbi neonate/juvenile right mandible from TUSK (photo courtesy NPS).
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60. Waco Mammoth National Monument (WACO), Texas
WACO is a recently established national monument and is managed in partnership with
the City of Waco and Baylor University. Many of the fossil mammoth remains along the
Bosque River are left in situ for the public to see (Fig. 14); those fossils removed are curated
at the Mayborn Museum, Baylor University. As reported by the NPS, the mammoth fossils
represent the largest herd of Mammuthus columbi that died in a single catastrophic event
(Nordt et al. 2015; although see Wiest et al. 2016, Esker 2018). At least 26 mammoths are
known to occur in the deposit, along with a few other taxa. The age and cause(s) of death
are subjects of discussion, but there is an OSL analysis putting the time of death at about
66,800 ± 5k yr. B.P. (MIS 4 [Marine Isotope Stage]; Nordt et al. 2015).
61. White Sands National Monument (WHSA), New Mexico
Vandiver (1936) investigated a report of Mammuthus bones and teeth north of WHSA in
what is now White Sands Missile Range, but found only fragmentary remains. The proboscidean
record at WHSA today consists of many thousands of in situ tracks (ichnofauna megatracksite)
preserved in the gypsum sand and playa lake deposits, which were first encountered
at the missile range as early as 1981 (Santucci et al. 2007). Diverse and abundant late Pleistocene
tracks have been found within park boundaries, most notably human tracks associated
with Pleistocene megafaunal tracks of a ground sloth (Bustos et al. 2018). The proboscidean
tracks (Fig. 15) have been described recently in terms of their biological attributes, and using
new technologies including magnetometry and 3-D radar imaging for detecting and documenting
the tracks (Urban et al. 2018, 2019).
62. Wupatki National Monument (WUPA), Arizona
A single Mammuthus specimen is reported from the sediments immediately outside
the boundary of WUPA (Saunders 1970: a right tibia, site 52). Henderek et al. (2017) in-
Figure 14. Mammuthus columbi skull fragment with in situ large tusks from WACO (photo courtesy NPS).
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dicated that this specimen is most likely the tibia collected from the Navajo Reservation
west of Inscription Point in 1948 by D. Jones and J. Bean. This geographic feature is just
east of the location given by Saunders (1970). The fossil locality (MNA.LOC.1114 Inscription
Point West Area) is described as being located on the bank of the Little Colorado
River less than 1.6 km (1 mi) north of the WUPA boundary in undifferentiated Quaternary
alluvium.
63. Yukon-Charley Rivers National Preserve (YUCH), Alaska
Located in east-central Alaska along the Canada border, YUCH straddles a 185 km (115
mi) portion of the Yukon River and the entire Charley River basin. The Preserve’s enabling
legislation tasks the NPS with protection of an extensive geological and fossil record
(Santucci et al. 2011). Alluvial deposits along the preserve’s rivers are known to contain
Pleistocene fossils, which are typically found after having eroded and accumulated along
the base of stream banks and gravel bars. Placer gold mining in the 20th century, including
extensive industrial scale mechanical dredge mining, is another process that has exposed
and accumulated fossil material. Mining districts in YUCH include Fourth of July, Coal,
Ben and Woodchopper Creeks, all of which have yielded some mammoth specimens. A single
juvenile woolly mammoth (Mammuthus primigenius) molar was reported immediately
downstream from the mined area of Fourth of July Creek in 1986 (National Park Service
1989:106), and a molar and tusk of a young woolly mammoth were exposed by placer mining
in Ben Creek in western YUCH (National Park Service 1990).
Some of the earliest reported and most extensive finds have occurred at Woodchopper
Creek, roughly 80 km (50 mi) upriver from the Yukon River community of Circle. Gold was
first discovered here during the 1898 Klondike Gold Rush, and while little profitable gold
Figure 15. Proboscidean trackway
in playa deposit at WHSA (photo
courtesy NPS).
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was initially obtained at this locale, Woodchopper Creek achieved national fame when miners
encountered a mass of well-preserved extinct large mammal bones reportedly encountered
in a mine shaft 24 m (80 ft) below the ground surface The Mammuthus primigenius
specimens among them included the skull and lower jaw, with both tusks and all the molars;
pelvis; 1 scapula; 2 limb bones; 12 vertebrae; 15 ribs; and “some small bones” (Gilmore
1908:25, Quackenbush 1909:124) (Fig. 16).
NPS archives contain a photograph from the George Beck collection showing several
people including members of the Biederman family with a Mammuthus tusk near the Yukon
River, thought to have been taken in the 1930s somewhere in YUCH. The Biederman
homestead is today a private inholding in YUCH near Biederman Bluff.
Discussion and Conclusions
The National Park System covers more than 35,612,000 ha [~85 million ac], comprised
of 419 national parks and affiliated lands. These federal and affiliated holdings represent a
significant geographic area of the continent and embody a tremendous aggregate of natural
and cultural heritage. At least 276 NPS Units (as defined above) contain some aspect of the
North American fossil heritage, and here we have concentrated on those units and affiliated
lands that contain proboscidean remains. Proboscideans are a ubiquitous part of the
North American vertebrate faunas throughout the Miocene, Pliocene, and Pleistocene. Their
presence in greater North America included gomphotheres, mastodonts, and mammoths to
the end of the Rancholabrean, the most recent fully accepted LMA within the Pleistocene,
equivalent to approximately 11,000 radiocarbon years ago. Although these federal and affil-
Figure 16. Mammuthus primigenius bones
from Alice Creek (YUCH) on display in
1908 before they were purchased by the
Field Museum of Natural History, Chicago
(photo courtesy NPS).
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iated lands preserve vertebrate remains, it does not mean that all these fossils are recorded,
studied, or published about in any detailed, systematic form.
Within NPS units and affiliated localities that have fossils, we documented 63 that have
records concerning proboscideans. Proboscidean fossils from these units range from the
abundant skeletal remains, to the less common and less diagnostic track records, to rare,
dry-preserved dung. No frozen or mummified soft tissue remains have yet been encountered
on NPS or affiliated holdings, but there are strong prospects for such finds in park units in
northern Alaska where frozen contexts promote excellent preservation.
Based on data in Table 1, gomphotheres are the least represented group recorded from
NPS and affiliated lands. Of the NPS and affiliated localities, late Pleistocene (Rancholabrean)
localities are more commonly assessed than are older fossil sites; therefore, it can be expected
that there will be more records of mammoth and mastodon remains than gomphotheres. In
most cases the records that exist clearly point out that additional, more detailed analyses of
the remains are warranted to better understand generic and specific distributions.
This broad and admittedly superficial survey of proboscidean fossils from NPS and affiliated
lands highlights the considerable and mostly untapped research potential of fossil data
from these lands and museum collections. Admittedly, the distribution of the 63 NPS Units
shown in Figure 1 is a biased presentation of proboscidean fossil distribution across North
America. Certain areas, particularly east of the Mississippi River, are not well-represented
for various reasons.
Several conclusions come from our survey. 1) There are many NPS and affiliated units
that have field inventories of fossils bearing proboscidean remains. Our review here points
out that select, known-proboscidean fossil areas should be investigated further, in some case
intensely. This would be a first step toward management of these non-renewable resources.
2) The numerous unidentified and preliminarily described fossil occurrences should be
re-assessed by qualified vertebrate paleontologists to determine the precise taxonomy and
chronological assignment. 3) Specimens need to be entered into a museum collection management
system (ICMS, or MCMS) and made web-available. 4) This inventory and similar
inventories are intended to compile baseline paleontological resource data that is derived
from various sources of unpublished and often hard to obtain records, archives, field notes,
personal or email communications and other sources. Through this information collection
and data mining, it is often possible to demonstrate a more significant scope and distribution
of the fossil record being studied than would otherwise be understood through a review of
published literature. 5) As NPS and affiliated site holdings of proboscidean remains are verified/
identified and placed into a management system, collection managers should consider
having the information placed into a clearing house database system such as, but not limited
to, Neotoma Paleoecology Database, FAUNMAP, ePANDDA, iDigBio, and Paleobiology
Database. This would allow students, researchers, and interested public have access to the
wealth of information not easily discovered today.
Acknowledgments
The following NPS units and people are thanked for helping us with data records and photographs
for this article: Agate Fossil Beds National Monument (James Hill); Alaska Regional Office
(Kelsey Lutz and Amanda Lanik); Arches National Park (Peekay Briggs and Terry Fisk); Big Bend
National Park (Don Corrick); Colorado School of Mines (Dennis Gertenbach, Daniel Schlegel, and
Shellie Luallin); Canaveral National Seashore (Kristen Kneifl); Channel Islands National Park (Ken
Convery and Laura Kirn); Cogstone Resource Management, Inc.(Eric Scott); Colonial National
Historical Park (Dorothy Geyer); Death Valley National Park (Matthew Ferlicchi and Josh Hoines);
Denali National Park and Preserve (Denice Swanke, Phoebe Gilbert and Denny Capps); Eugene
Eastern Paleontologist
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O’Neill National Historic Site/John Muir NHS/Port Chicago Naval Magazine National Memorial/
Rosie the Riveter and WWII Home Front National Historical Park (Virginia Bones and Paul Scolari);
Hagerman Fossil Beds National Monument (Kari Prassack); Hopewell Culture National Historical
Park (Karen Beppler-Dorn); Independence National Historical Park (Karie Diethon); John Day Fossil
Beds National Monument (Nicholas A. Famoso); Klondike Gold Rush National Historical Park
(Deb Boettcher); Midwest Archeological Center (Linda Plock and Karin Roberts); Minnesota Historical
Society (Adam Scher); Natchez Trace Parkway (Lisa McInnis and Christina Smith); National
Natural Landmarks Program (Jeff Orlowski); Pennsylvania Academy of the Fine Arts (Philadelphia,
for permission to use Peale’s painting); Salinas Pueblo Missions National Monument (Ron Fields
and Marc LeFrancois); Santa Barbara Museum of Natural History (Jonathan Hoffman); Smithsonian
National Museum of Natural History (Diana Marsh); Tule Springs Fossil Beds National Monument
(Erin Eichenberg); US Geological Survey (Jeff Pigati and Kathleen Springer); University of Nebraska–
Lincoln (Robert Hunt); University of Nevada, Las Vegas (Aubrey Bonde); Waco Mammoth
National Monument (Raegan King and Lindsey Yann); White Sands National Monument (David
Bustos); Western Archeological and Conservation Center (Kim Beckwith, Brenda McLain, Tef Rodeffer,
and Khaleel Saba); Yukon-Charley Rivers National Preserve and Gates of the Arctic National
Park and Preserve (Jillian Richie).
Literature Cited
Agenbroad, L.D. 1998a. Pygmy (dwarf) mammoths of the Channel Islands of California. The Mammoth
Site of Hot Springs, SD. 27 pp.
Agenbroad, L.D. 1998b. New pygmy mammoth (Mammuthus exilis) localities and radiocarbon dates
from San Miguel, Santa Rosa, and Santa Cruz Islands, California. Pp. 169–175, In P.W. Weigand
(Ed.). Contributions to the Geology of the Northern Channel Islands, Southern California. American
Association of Petroleum Geologists. 196 pp.
Agenbroad, L. D., and J.I. Mead. 1989. Quaternary geochronology and distribution of Mammuthus on
the Colorado Plateau. Geology 17:861–864.
Agenbroad, L.D., and J.I. Mead (Eds.). 1994. The Hot Springs Mammoth Site: A decade of field and
laboratory research in paleontology, geology, and paleoecology. The Mammoth Site, Hot Springs,
SD. 457 pp.
Agenbroad, L.D., J.I. Mead, E.M. Mead, and D. Elder. 1989. Archaeology, alluvium, and cave stratigraphy:
The record from Bechan Cave, Utah. Kiva 54:335–351.
Agenbroad, L.D., and A.R. Brunelle. 1993. Recovery and analysis of the Virgin Mammoth, Lake
Mead National Recreation Area, Clark County, Nevada. Pp. 28, In Proceedings of the Southwestern
and Rocky Mountain Division of the American Association for the Advancement of Science,
68th Annual Meeting.
Agenbroad, L.D., D. Morris, and L. Roth. 1999. Pygmy mammoths Mammuthus exilis from Channel
Islands National Park, California (USA). Deinsea 6:89–102.
Agenbroad, L.D., J.R. Johnson, D. Morris, and T.W. Stafford. 2005. Mammoths and humans as late
Pleistocene contemporaries on Santa Rosa Island. Pp. 4–7, In D.K. Garcelon and C.A. Schwemm
(Eds.). Proceedings of the Sixth California Islands Symposium, Ventura, California, December
1–3, 2003. Institute for Wildlife Studies, Arcata, California. 508 pp.
Agenbroad, L.D., J.I. Mead, C.V. Haynes, Jr., and R.H. Hevly. 2013. Fossil fauna and flora from late
Pleistocene Cerros Negros locality, Pinal County, Arizona: With update of Mammuthus and all
Mammut localities from Arizona. Southwestern Naturalist 58:50–53.
Allen, H.T. 1887. Report of an Expedition to the Copper, Tanana, and Koyukuk Rivers, in the Territory
of Alaska, in the Year 1885. Government Printing Office, Washington, DC.
Anonymous. 1811. Curious discovery [elephant bones from York River, Williamsburg, Virginia].
Philadelphia Repertory 2:87–88.
Barbour, E.H. 1914. Mammalian fossils from Devil’s Gulch. Nebraska Geological Survey 4:177–190.
Barbour, E.H. 1927. Preliminary notice of a new proboscidean Ambelodon fricki, gen. et sp. nov. Bulletin
of the Nebraska State Museum 13:131–134.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
36
Bell, C.J., E.L. Lundelius, A.D. Barnosky, R.W. Graham, E.H. Lindsay, D.R. Ruez, H.A. Semken,
S.D. Webb, and R.J. Zakrzewski. 2004. The Blancan, Irvingtonian, and Rancholabrean Mammal
Ages. Pp. 232–314, In M.O. Woodburne (Ed.). Late Cretaceous and Cenozoic Mammals of North
America. Biostratigraphy and Geochronology. Columbia University Press, NY. 391 pp.
Bement, L.C. 1986. Mammalian faunal and cultural remains in the Late Pleistocene deposits of Bonfire
shelter, 41VV218, Southwest Texas. The University of Texas at Austin, TX. 73 pp.
Bement, L.C. 2007. Bonfire shelter: A jumping off point for comments for Beyerly et al. American
Antiquity 72:366–372.
Blong, J.C. 2018. Late-glacial hunter-gathers in the central Alaska Range and the role of upland ecosystems
in the peopling of Alaska. PaleoAmerica 4:103–133.
Blumenbach, J.F. 1799. Handbuch der Naturgeschichte. (6th Edition). Dieterich, Göttingen, Germany.
708 pp.
Bonde, A., V. L. Santucci, J. S. Tweet. E. Eichenberg, and B. Moore. 2018. Lake Mead National
Recreation Area: paleontological resources inventory. Natural Resource Report NPS/
Lake?NRR-2018/1618. National Park Service, Fort Collins, CO. 250 pp.
Brady, L.F., and P. Seff. 1959. “Elephant Hill”. Plateau 28:80–82.
Brookes, J. 1828. A catalogue of the anatomical and zoological museum of Joshua Brookes, Esq. Part
1. R. Taylor, London, UK. 76 pp.
Buckland, W. 1831. On the Occurrence of the Remains of Elephants, and other Quadrapeds, in the
Cliffs of Frozen Mud, in Eschscholtz Bay, within Bering’s Strait, and in other Distant Parts of the
Shores of the Arctic Seas. Pp. 331–356, In F.W. Beechey. Narrative of a voyage to the Pacific and
Beering’s strait, to co-operate with the polar expeditions: Performed in His Majesty’s ship Blossom,
under the command of Captain in the years 1825, 26, 27, 28, Volume II. H. Colburn and R.
Bentley, London, UK. 452 pp.
Burmeister, H. 1837. Handbuch der Naturgeschichte. Zum Gebrauch bei Vorlesungen Entworfen.
Zweite Abteilung. Zoologie, T.C.F. Enslin, Berlin, Germany. 858 pp.
Bustos, D., J. Jakeway, T. M. Urban, V. T. Holliday, B. Fenerty, D. A. Raichlen, M. Budka, S. C. Reynolds,
B. D. Allen, D. W. Love, V. L. Santucci, D. Odess, P. Willey, H. G. McDonald, and M. R.
Bennett. 2018. Footprints preserve terminal Pleistocene hunt? Human-sloth interactions in North
America. Science Advances 2018:4:eaar7621.
Cantwell, J.C. 1887. A narrative account of the exploration of the Kowak River, Alaska. Pp. 21–52,
In M.A. Healy. Report of the cruise of the Revenue Marine steamer Corwin in the Arctic Ocean in
the year 1885. Government Printing Office, Washington, DC. 102 pp.
Cantwell, John C. 1889. A narrative account of the exploration of the Kowak River, Alaska. Pp. 49–74,
In M.A. Healy. Report of the cruise of the Revenue Marine steamer Corwin in the Arctic Ocean in
the year 1884. Government Printing Office, Washington, DC. 128 pp.
Cassiliano, M.L. 1999. Biostratigraphy of Blancan and Irvingtonian mammals in the Fish Creek-
Vallecito Creek section, southern California, and a review of the Blancan-Irvingtonian boundary.
Journal of Vertebrate Paleontology 19(1):169–186.
Clark, W.B., and B.L. Miller. 1912. The physiography and geology of the Coastal Plain province of
Virginia. Virginia Geological Survey Bulletin 4:13–222.
Clements, T.D. 1952. Lake Rogers, a Pleistocene lake in the north end of Death Valley, California.
Geological Society of America Bulletin 63(12, Part 2):1324.
Colburn, M.L. 2017. Under foot: The paleontology of Mammoth Cave. Pp. 175–189, In H.H. Hobbs,
III, R.A. Olson, E.G. Winkler, and D.C. Culver (Eds.). Mammoth Cave: A human and natural history.
Springer, NY. 275 pp.
Cook, H.J. 1914. Note on the occurrence of the mammoth in Sioux County, Nebraska. Nebraska Geological
Survey Bulletin 7(6):1–3.
Cope, E.D. 1871. Preliminary report on the vertebrata discovered in the Port Kennedy Bone Cave.
Proceedings of the American Philosophical Society 12:73–102.
Cope, E.D. 1895. The fossil Vertebrata from the fissure at Port Kennedy. Proceedings of the Academy
of Natural Sciences of Philadelphia 47:446–450.
Cope, E.D. 1896. New and little known Mammalia from the Port Kennedy bone deposit. Proceedings
of the Academy of Natural Sciences of Philadelphia 48:378–394.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
37
Cope, E.D. 1899. Vertebrate remains from Port Kennedy bone deposit. Journal of the Academy of
Natural Sciences of Philadelphia, 2nd series 11:193–286.
Cotter, J. L. 1937. The occurrence of flints and extinct animals in pluvial deposits near Clovis, New
Mexico. Part IV: Report on excavation at the gravel pit, 1936. Proceedings of the Academy of
Natural Sciences of Philadelphia 89:1–16.
Czaplewski, N., and K. Davies. 2007. Not tracks after all? The supposed gomphothere tracks at “Elephant
Hill”, Montezuma Castle National Monument, Arizona (Verde Formation). New Mexico
Museum of Natural History, Albuquerque, New Mexico, Cenozoic Vertebrate Tracks Symposium.
Daeschler, E., E.E. Spamer, and D.C. Parris. 1993. Review and new data on the Port Kennedy local
fauna and flora (Late Irvingtonian), Valley Forge National Historical Park, Montgomery County,
Pennsylvania. The Mosasaur 5:23–41.
Daeschler, E.B., M.C. Lamanna, and M. Carfioli. 2005. On the trail of an important Ice Age fossil
deposit. Park Science 23(2):31–34.
Davis, O.K. 1987. Spores of the dung fungus Sporomiella: Increased abuncance in historic sediments
and before Pleistocene megafaunal extinction. Quaternary Research 28:290–294.
Davis, O.K. 1990. Caves as sources of biotic remains in arid western North America. Palaeogeography,
Palaeoclimatology, Palaeoecology 76:331–348.
Davis, O.K. 2003. Fungal spores from coprolites: A means of detecting herbivore density. Palynology
27:223.
Davis, O.K., and D.S. Shafter. 2006. Sporomiella fungal spores, a palynological means of detecting
herbivore density. Palaeogeography, Palaeoclimatology, Palaeoecology 237:40–50.
Davis, O.K., L.D. Agenbroad, P.S. Martin, and J.I. Mead. 1984. The Pleistocene dung blanket of
Bechan Cave, Utah. Pp. 267–282, In, H.H. Genoways and M.R. Dawson (Eds.). Contributions in
Quaternary vertebrate paleontology: A volume in memorial of John E. Guilday. Carnegie Museum
of Natural History Special Publication 8. 538 pp.
Davis, O.K., J.I. Mead, P.S. Martin, and L.D. Agenbroad. 1985. Riparian plants were a major component
of the diet of mammoths of southern Utah. Current Research in the Pleistocene 2:81–82.
Denton, W. 1877. On the asphalt bed near Los Angeles, California. Proceedings of the Boston Society
of Natural History 18:185–186.
Dibble, D.S., and D. Lorrain. 1968. Bonfire shelter: A stratified bison kill site, Val Verde County,
Texas. Miscellaneous Papers No. 1, Texas Memorial Museum, Austin, TX. 138 pp.
Dooley, A.C., E. Scott, J. Green, K.B. Springer, B.S. Dooley, and G.J. Smith. 2019. Mammut pacificus
sp. nov., a newly recognized species of mastodon from the Pleistocene of western North America.
PeerJ 7:e6614.
El Aldi, J.J., Cherney, M.D., Fisher, D.C., Harris, J.M., Farrell, A.B., and Cox, S.M. 2015. Last Years
of Life and Season of Death of a Columbian Mammoth from Rancho La Brea. Pp. 65–80, In J.M.
Harris (Ed.). La Brea and beyond: The paleontology of asphalt-preserved biotas. Natural History
Museum of Los Angeles County, Science Series, Number 42, Los Angeles, CA. 174 pp.
Elder, W., V.L. Santucci, J.P. Kenworthy, R.B. Blodgett, and R.T.P. McKenna. 2009. Paleontological
resource inventory and monitoring—Arctic Network. Natural Resource Technical Report NPS/
NRPC/NRTR—2009/276. National Park Service, Fort Collins, CO. 98 pp.
Emslie, S.D. 1987. Age and diet of fossil California condors in the 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.
Esker, D.A. 2018. Strontium isotope ratios reveal disparate geographic origins for megafauna at Waco
Mammoth National Monument (TX, USA.). Journal of Vertebrate Paleontology, Program and
Abstracts 2017:123.
Falconer, H. 1857. On the species of mastodon and elephant occurring in the fossil state in Great
Britain. Part I. Mastodon. Quarterly Journal of the Geological Society of London 13:307–360.
Falconer, H. 1868. Palaeontological memoirs and notes of the late Hugh Falconer with a biographical
sketch of the author. C. Murchison (Ed.). Hardwicke, London, UK, 2 vol., 594 and 675 pp.
Fay, L.C. 2009. Geological resources inventory scoping summary: Big Thicket National Preserve, Texas.
National Park Service Geologic Resources Division, National Parks Service, Fort Collins, CO. 14 pp.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
38
Fay, L.C., J.P. Kenworthy, and V.L. Santucci. 2009. Paleontological resource inventory and monitoring—
North Coast and Cascades Network. Natural Resource Technical Report NPS/NRPC/
NRTR—2009/250. National Park Service, Fort Collins, CO. 162 pp.
Flower, B.J., and J.P. Kennett. 1993. Middle Miocene ocean-climate transition: High resolution
oxygen and carbon isotopic records from Deep Sea Drilling Project site 588A, southwest Pacific.
Paleoceanography 8:811–843.
Frick, C. 1933. New remains of trilophodont-tetrabelodont mastodons. Bulletin of the American Museum
of Natural History 59:505–652.
Fremd, T. J. 2010. Guidebook. SVP Field Symposium 2010. John Day Basin Field Conference. Society
of Vertebrate Paleontology. 48 pp.
Fremd, T.J., E.A. Bestland, and G.J. Retallack. 1994. John Day Basin Field Trip Guide and Road Log.
Society of Vertebrate Paleontology Annual Meetings 1994:1–80.
Gazin, C.L. 1936. A study of the fossil horse remains from the upper Pliocene of Idaho. Proceedings
of the United States National Museum 83:281–320.
Giddings, J.L. 1962. Onion Portage and other flint sites of the Kobuk River. Arctic Anthropology 1:6–27.
Gidley, J.W., and C.L. Gazin. 1933. New Mammalia in the Pleistocene fauna from Cumberland Cave.
Journal of Mammalogy 14:343–357.
Gidley, J.W., and C. L. Gazin. 1938. The Pleistocene vertebrate fauna from Cumberland Cave, Maryland.
United States National Museum Bulletin 171:1–99.
Gilmore, C.W. 1908. Smithsonian exploration in Alaska in 1907 in search of Pleistocene fossil vertebrates.
Smithsonian Miscellaneous Collection 51:1–38.
Graham, R.W.S. Belmechen, K. Choy, B.J. Culleton, L.J. Davies, D. Froese, P.D. Heintzman, C.Hritz,
J.D. Kapp, L.A. Newsom, R. Rawcliffe, E.Saulnier-Talbot, B. Shapiro, Y. Wang, J.W. Williams,
and M.J. Wooller. 2016. Timing and causes of mid-Holocene mammoth extinction on St. Paul
Island, Alaska. Proceedings of the National Academy of Science 113:9310–9314.
Gubser, N.J. 1965. The Numamiut Eskimos: Hunters of caribou. Yale University Press, New Haven,
CT. 384 pp.
Guthrie, R.D. 1985. Woolly arguments against the mammoth steppe: A new look at the palynological
data. Quarterly Review of Archaeology 6:9–16.
Guthrie, R.D. and S. Stoker. 1990. Paleoecological significance of mummified remains of Pleistocene
horses from the North Slope of the Brooks Range, Alaska. Arctic 43:267–274.
Hack, J.T. 1942. The changing physical environment of the Hopi Indians of Arizona. Peabody Museum
Papers 35(1):1–85.
Hall, E.S., Jr. 1971. Kangiguksuk: A cultural reconstruction of a sixteenth century Eskimo site in
northern Alaska. Arctic Anthropology 8:1–101.
Hamilton, T.D., G.M. Ashley, K.M. Reed, and C.E. Schweger. 1993. Late Pleistocene vertebrates and
other fossils from Epiguruk, northwestern Alaska. Quaternary Research 39:381–389.
Harrington, M.R. 1934. A camel-hunter’s camp in Nevada. The Masterkey 8:22–24.
Haury, E.W. 1953. Artifacts with mammoth remains, Naco, Arizona. American Antiquity 19:1–14.
Haury, E.W., E.B. Sayles, and W.W. Wasley. 1959. The Lehner Mammoth Site, southeastern Arizona.
American Antiquity 25:2–30.
Hay, O.P. 1923. The Pleistocene of North America and its vertebrated animals from the states east
of the Mississippi River and from the Canadian provinces east of longitude 95 degrees. Carnegie
Institution of Washington Publication 322, Washington, DC. 499 pp.
Hay, O.P. 1924. The Pleistocene of the middle region of North America and its vertebrated animals.
Carnegie Institute of Washington Publication 322A, Washington, DC. 385 pp.
Hay, O.P. 1927. The Pleistocene of the western region of North America and its vertebrate animals.
Carnegie Institute of Washington Publication 322B, Washington, DC. 346 pp.
Haynes, C.V. 1967. Quaternary geology of the Tule Springs Area, Clark County, Nevada. Pp. 15–104,
In H.M. Wormington and D. Ellis (Eds.). Pleistocene studies in southern Nevada. Nevada State
Museum, Anthropological Papers 13. Carson City, Nevada. 411 pp.
Haynes, C.V., Jr., and G.A. Agogino. 1966. Prehistoric springs and geochronology of the Clovis Site,
New Mexico. American Antiquity 31:812–821.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
39
Haynes, C.V., and E.T. Hemmings. 1968. Mammoth-bone shaft wrench from Murray Springs. Science
159:186–187.
Haynes, C.V., and B.B. Huckell (Eds.). 2007. Murray Springs: A Clovis site with multiple activity
areas in the San Pedro Valley, Arizona. The University of Arizona Press, Anthropological Papers
71, Tucson, AZ. 308 pp.
Heilprin, A., and J. Leidy. 1887. Report of the curators. Proceedings of the Academy of Natural Sciences
of Philadelphia 39:411–416.
Hemmings, E.T. 2007. Buried animal kills and processing localities, Areas 1–5. Pp. 83–137, In C.V.
Haynes and B.B. Huckell (Eds.). Murray Springs. A Clovis site with multiple activity areas in the
San Pedro Valley, Arizona. The University of Arizona Press, Anthropological Papers 71, Tucson,
AZ. 308 pp.
Henderek, R., V.L. Santucci, B. Rizner, J. Tweet, and J. Wood. 2017. Wupatki National Monument
paleontological resources inventory. Natural Resource Report NPS/WUPA/NRR—2017/1524.
National Park Service, Fort Collins, CO. 86 pp.
Henkel, C.J., W.P. Elder, V.L. Santucci, and E.C. Clites. 2015. Golden Gate National Recreation
Area: Paleontological resource inventory. Natural Resource Report NPS/GOGA/NRR-2015/915.
National Park Service, Fort Collins, CO. 78 pp.
Herridge, V.L., and A.M. Lister. 2012. Extreme insular dwarfism evolved in a mammoth. Proceedings
of the Royal Society B 279:3193–3200.
Hester, J.J. 1972. Blackwater Locality No. 1: A stratified early man site in eastern New Mexico. Fort
Burgwin Research Center Publication 8. Southern Methodist Universi ty, Dallas, TX. 238 pp.
Hibben, F.C. 1941. Evidences of early occupation in Sandia Cave, New Mexico, and other sites in the
Sandia-Manzano region. Smithsonian Miscellaneous Collections 99(23).
Holman, J.A. 1977. The Kuhre Quarry Fauna (Upper Miocene), north-central Nebraska. Michigan
Academy of Science 9:459–467.
Hooper, C.L. 1884. Cruise of the US Revenue Steamer Corwin in the Arctic Ocean in 1881. Report to the 48th
Congress, First Session. Senate Document 204. Government Printing Office, Washington, DC. 92 pp.
Howard, E. B. 1935. The occurrence of flints and extinct animals in pluvial deposits near Clovis,
New Mexico. Part I: Introduction. Proceedings of the Philadelphia Academy of Natural Sciences
87:299–303.
Hunt, A.P. and V.L. Santucci. 2001. Paleontological resources of Lake Meredith National Recreation
Area and Alibates Flint Quarries National Monument, west Texas. Pp. 257–264, In S.G. Lucas and
D.S. Scholle (Eds.). Geology of the Llano Estacado. New Mexico Geological Society, Socorro,
New Mexico. Guidebook 52. 340 pp.
Hunt, R., J.P. Kenworthy, and V.L. Santucci. 2008. Paleontological resource inventory and monitoring-
Heartland Network. Natural Resource Technical Report NPS/NRPC/NRTR-2008/132.
National Park Service, Washington, DC. 122 pp.
Hunter, R.E., H.E. Clifton, N.T. Hall, G. Császár, B.M. Richmond, and J.C. Chin. 1984. Pliocene and
Pleistocene coastal and shelf deposits of the Merced Formation and associated beds, northwestern
San Francisco Peninsula, California. Pp. 1–29, In Field Trip Guidebook 3. Society of Economic
Paleontologists and Mineralogists, Tulsa, OK. 126 pp.
Hurt, W.R. 1990. The 1939–1940 excavation project at Quarai Pueblo and Mission building: Santa Fe,
New Mexico. National Park Service Southwest Cultural Resources Center. Professional Paper 29.
Jefferson, G.T. 1991. A catalogue of Late Quaternary vertebrates from California. Part two: Mammals.
Natural History Museum of Los Angeles County, Technical Reports 7, Los Angeles, CA. 129 pp.
Johnson, A.E. 1874. Geological and archeological evidences of the antiquity of man. Bulletin of the
Minnesota Academy of Natural Sciences 1:101–141.
Karpinski, E., J.I. Mead, and H.N. Poinar. 2017. Molecular identification of paleofeces from Bechan
Cave, southeastern Utah, USA. Quaternary International 443:140–146.
Kast, S. 2008. Preliminary paleontological survey of Lava Beds National Monument, California.
Unpublished report to Lava Beds National Monument. National Park Service, Fort Collins, CO.
Katz, L. 1997. Late Pleistocene fauna at Blackwater Draw locality #1: An archaeological perspective:
New Mexico Museum of Natural History and Science Bulletin 11:135–143.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
40
Kenworthy, J.P., V.L. Santucci, M. McNerney, and K. Snell. 2005. Paleontological Resource Inventory
and Monitoring, Upper Columbia Basin Network. National Park Service TIC# D-259. National
Park Service, Fort Collins, CO. 74 pp.
Kenworthy, J.P., V.L. Santucci, and C.C. Visaggi. 2007. Paleontological Resource Inventory and
Monitoring, Gulf Coast Network. National Park Service TIC# D-750. National Park Service, Fort
Collins, CO. 106 pp.
Kitchell, W. 1856. Second annual report on the Geological Survey of the state of New Jersey for the
year 1855. True American Office, Trenton, NJ. 268 pp.
Koch, A.L., V.L. Santucci, and T.R. Weasma. 2004. Santa Monica Mountains National Recreation
Area paleontological survey. NPS Geologic Resources Division, Denver, Colorado. Technical
Report NPS/NRGRD/GRDTR—04/01. National Park Service, Fort Collins, CO.
Knox, S.G., and S. Pitts. 1984. Excavation of a mastodon at Vicksburg, Mississippi. Mississippi Geology
4(4):1–10.
Kotzebue, O. von. 1821. A voyage of discovery into the South Sea and Beering’s [sic] Straits, for the
purpose of exploring a north-east passage undertaken in the years 1815–1818, at the expense of
His Highness the chancellor of the empire, Count Romanzoff, in the ship Rurick, under the command
of the lieutenant in the Russian Imperial Navy, Otto von Kotzebue. Longman, Hurst, Rees,
Orme, and Brown, London, UK. 358 pp.
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 Research
67:143–151.
Lambert, W.D. 1996. The biogeography of the gomphotheriid proboscideans of North America. Pp.
143–148, In J. Shoshani and P. Tassy (Eds.). The Proboscidea. Evolution and Palaeoecology of
Elephants and Their Relatives. Oxford University Press, New York, NY. 472 pp.
Lambert, W.D., and J. Shoshani. 1998. Proboscidea. Pp. 606–621, In C.M. Janis, K.M. Scott, and L.L.
Jacobs (Eds.). Evolution of tertiary Mammals of North America. Cambridge University Press,
New York, NY.
Lance, J.F. 1959. Faunal remains from the Lehner Mammoth Site. American Antiquity 25:35–42.
Lanik, A., C. Hults, and R.B. Blodgett. 2017. A paleontological inventory of Arctic parks. Alaska Park
Science 16:29–34.
Larsen, H. 1968. Trail Creek: Final report on the excavation of two caves on Seward Peninsula,
Alaska. Acta Arctica 15.
Lee, C. and T. Goebel. 2016. The slotted antler points from Trail Creek Caves, Alaska: New information
on their age and technology. PaleoAmerica 2:40–47.
Leidy, J. 1858. Notice of remains of extinct Vertebrata, from the valley of the Niobrara River, collected
during the exploring expedition of 1857, in Nebraska, under the command of Lieut. G.K.
Warren, U.S. Top. Eng., by Dr. F.V. Hayden, Geologist to the expedition. Proceedings of the
Academy of Natural Sciences of Philadelphia 10:20–29.
Liggett, G.A., T. Childs, N.A. Famoso, S. Floray, H.G. McDonald, A.L. Titus, and E. Varner. 2018.
An overview of the contributions of federal land to paleontology, and a discussion of the U.S.
Department of the Interior Museum Program. Pp. 311–334, In G.D. Rosenberg, and R.M. Clary
(Eds.). Museums at the Forefront of the History of Philosophy of Geology, History Made, History
in the Making. Geological Society of America, Special Paper 535, Boulder, CO. 348 pp. DOI:
10.1130/2018.2535(21).
Lister, A. M. 2017. On the type material and evolution of North American mammoths. Quaternary
International 443A:14–31.
Lister, A.M., and A.V. Sher. 2015. Evolution and dispersal of mammoths across the Northern Hemisphere.
Science 350:805–809.
Longwell, C.R. 1963. Reconnaissance geology between Lake Mead and Davis Dam, Arizona-Nevada.
U.S. Geological Survey, Professional Paper 374-E. Washington, DC. 57 pp.
Lucas, S.G., and J.A. Effinger. 1991. Mammuthus from Lincoln County and review of the mammoths
from the Pleistocene of New Mexico. Pp. 277–282, In J.M. Barker, B.S. Kues, G.S. Austin, and
S.G. Lucas (Eds.). Geology of the Sierra Blanca, Sacramento, and Capitan Ranges, New Mexico.
New Mexico Geological Society 42nd Annual Fall Field Conference Guidebook.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
41
Lull, R.S. 1908. The evolution of the elephant. American Journal of Science 25(147):169–212.
Lundelius, E.L., Jr. 1972. Vertebrate remains from the gray sand. Pp. 148–163, In J. J. Hester (Ed.).
Blackwater locality no. 1. A stratified early man site in eastern New Mexico. Fort Burgwin Research
Center Publication 8. Southern Methodist University, Dallas, TX. 238 pp.
Maddren, A.G. 1905. Smithsonian exploration in Alaska in 1904, in search of mammoth and other
fossil remains. Smithsonian Miscellaneous Collections 49:1–117.
Maguire, K.C., J.X. Samuels, and M.D. Schmitz. 2018. The fauna and chronostratigraphy of the
middle Miocene Mascall type area, John Day Basin, Oregon, USA. PaleoBios 35:1–51.
Malmon, D.V., K.A. Howard, P.K. House, S.C. Lundtrom, P.A. Pearthree, A.M. Sarna-Wojcicki, E.
Wan, and D.B. Wahl. 2011. Stratigraphy and depositional environments of the upper Pleistocene
Chemehuevi Formation along the lower Colorado River. U.S. Geological Survey Professional
Paper 1786. Reston, VA. 106 pp.
Mann, D. H., P. Groves, J. L. Kunz, R. E. Reanier, and B. V. Gaglioti. 2013. Ice-age megafauna in
Arctic Alaska: extinction, invasion, survival. Quaternary Science Reviews 70:91–108.
Marcus, L. F. 1960. A census of the abundant large Pleistocene mammals from Rancho La Brea. Los
Angeles County Museum Contributions in Science 38:1–11.
Martin, P.S. 1987. Late Quaternary extinctions: The promise of TAMS 14C dating. Nuclear Instruments
and Methods B29:179–186.
Maxwell, R.A., J.T. Lonsdale, R.T. Hazzard, and J.A. Wilson. 1967. Geology of Big Bend National
park, Brewster County, Texas. University of Texas Bureau of Economic Geology Publication
6711. University of Texas, Austin, TX. 356 pp.
McClenahan, P.L. and D.E. Gibson 1990. Cape Krusenstern National Monument: An Archeological
Survey, Volumes I–II. Anchorage, Alaska, National Park Service, Alaska Regional Office, Research/
Resources Management Report AR-17, Anchorage, AK. 100 pp.
McDaniel, G.E., Jr., and G.T. Jefferson. 1997. A nearly complete skeleton of Mammuthus meridionalis
from the Borrego Badlands, Anza-Borrego Desert State Park, California. San Bernardino County
Museum Association Quarterly 44:35–36.
McDaniel, G.E., Jr., and G.T. Jefferson. 1999. Mammuthus meridionalis from the Borrego Badlands
Anza-Borrego Desert State Park, Borrego Springs, California. Final Report on the “Miller Mammoth”
for the Colorado Desert District Stout Research Center, Anza-Borrego Desert State Park,
Borrego Springs, CA. 74 pp.
McDaniel, G.E., Jr., and G.T. Jefferson. 2006. Mammoths in our midst: The proboscideans of
Anza-Borrego Desert State Park, Southern California, USA. Quaternary International 142–
143:124–129.
McGeorge, R., and C. Schur. 1994. Preliminary mapping of Elephant Hill, Verde Formation. Southwest
Paleontological Society, Mesa Southwest Museum, AZ. Unpublished report. On file at Montezuma
Castle National Monument, AZ.
Mead, J.I. 1981. The last 30,000 years of faunal history within the Grand Canyon, Arizona. Quaternary
Research 15:311–326.
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.V. Haynes, and B.B. Huckell. 1979. A late Pleistocene mastodon (Mammut americanum)
from the Lehner Site, southeastern Arizona. Southwestern Naturalist 24:231–238.
Mead, J.I., L.D. Agenbroad, P.S. Martin, and O.K. Davis. 1984. The mammoth and sloth dung from
Bechan Cave in southern Utah. Current Research in the Pleistocene 1:79–80.
Mead, J.I., L.D. Agenbroad, O.K. Davis, and P.S. Martin. 1986. Dung of Mammuthus in the Arid
Southwest, North America. Quaternary Research 25:121–127.
Mead, J.I., J.T. Sankey, and H.G. McDonald. 1998. Pliocene (Blancan) herpetofaunas from the
Glenns Ferry formation, southeastern Idaho. Pp. 146–155, In W.A. Akersten, H.G. McDonald,
D.J. Meldrum, and M.T. Flint (Eds.). And Whereas … Papers on the Vertebrate Paleontology of
Idaho honoring John A. White, Volume 1. Idaho Museum of Natural History, Occasional Paper
36, Pocatello, ID. 216 pp.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
42
Mead, J.I., and S.L. Swift. 2012. Late Pleistocene (Rancholabrean) dung deposits of the Colorado
Plateau, western North America. Pp. 337–342, In A. Hunt et al. (Eds.), Vertebrate coprolites. New
Mexico Museum of Natural History and Science, Bulletin 57.
Mehringer, P.J., and C.V. Haynes. 1965. The pollen evidence for the environment of Early Man
and extinct mammals at the Lehner Mammoth Site, southeastern Arizona. American Antiquity
31:17–23.
Mercer, H.C. 1899. The bone cave at Port Kennedy, Pennsylvania, and its partial excavation in 1894,
1895, and 1896. Journal of the Academy of Natural Sciences of Philadelphia 11:269–286.
Merriam, J. C. 1911. The fauna of Rancho La Brea. Part I. Occurrence. University of California
Memoir 1:192–21.
Metcalfe, J.Z., F.J. Longstaffe, J.A.M. Ballenger, and C.V. Haynes. 2011. Isotopic paleoecology
of Clovis mammoths from Arizona. Proceedings of the National Academy of Sciences 108
(44):17916–17920.
Meyer, H.W., and L. Weber. 1995. Florissant fossil Beds National Monument: Preservation of an
ancient ecosystem. Rocks and Minerals 70:232–239.
Miller, G.J., P. Remeika, J. Parks, B. Stout, and V.E. Waters. 1988. A preliminary report on halfa-
million-year-old marks on mammoth bones from the Anza-Borrego Desert Irvingtonian. San
Bernardino County Museum Association Quarterly 35:41.
Miller, G.J., P. Remeika, J. Parks, B. Stout, and V.E. Waters. 1991. A preliminary report on half-amillion-
year-old marks on mammoth bones from the Anza-Borrego Desert Irvingtonian. Imperial
Valley College Museum Society Occasional Paper 8:1–47.
Miller, S.J., W.A. Akersten, and R.M. Yohe, II. 1998. The Tolo Lake mammoth site, Idaho County,
Idaho. Pp. 110–120, In W.A. Akersten, H.G. McDonald, D.J. Meldrum, and M.E.T. Flint (Eds.).
And Whereas … Papers on the vertebrate paleontology of Idaho honoring John A. White, Volume
1. Idaho Museum of Natural History, Occasional Paper 36, Pocatello, ID. 216 pp.
Mills, W.C. 1922. Exploration of the Mound City Group. Certain Mounds and Village Sites in Ohio
3(4). Ohio Archaeological and Historical Society 31:245–408.
Mitchill, S.L. 1818. Observations on the geology of North America; illustrated by the description of
various organic remains found in that part of the World. Pp. 319–431, In G. Cuvier, Essay on the
theory of the Earth, with mineralogical notes, and an account of Cuvier’s geological discoveries,
by Professor Jameson. Kirk and Mercein, NY. 348 pp.
Moffit, F.H. 1905. The Fairhaven gold placers, Seward Peninsula, Alaska. US Geological Survey
Bulletin 247:1–85.
Morgan, G.S., S.G. Lucas, P.L., and A.P. Hunt. 2001. A review of Pleistocene vertebrate faunas from
northeastern New Mexico. New Mexico Geological Society, Socorro, New Mexico. Guidebook
52:265–284.
Muhs, D.R., K.R. Simmons, L.T. Groves, J.P. McGeehin, R.R. Schumann, and L.D. Agenbroad. 2015.
Late Quaternary sea-level history and the antiquity of mammoths (Mammuthus exilis and Mammuthus
columbi), Channel Islands National Park, California, USA. Quaternary Research 83:502–521.
Muir, J. 1917. The cruise of the Corwin. Houghton Mifflin, Boston, MA. 278 pp.
Murray, L.K. 2008. Effects of taxonomic and locality inaccuracies on biostratigraphy and biochronology
of the Hueso and Tapiado formations in the Vallecito Creek–Fish Creek Section, Anza-
Borrego Desert, California. Ph.D. Dissertation. University of Texas at Austin. 532 pp.
National Park Service. 1989. Draft environmental impact statement, cumulative impacts of mining,
Yukon-Charley Rivers National Preserve. Alaska Regional Office, Anchorage, Alaska.
National Park Service. 1990. Yukon-Charley Rivers National Preserve, final Environmental Impact
Statement, Cumulative Effects of Mining. Vol. 1. Denver Service Center, Denver, CO.
Nesti, F. 1825. Sulla nuova specie di elefante fossile del Valdarno all’Illustrissimo sig. Dott. Prof.
Ottaviano Targioni Tozzetti. Nuovo Giornale de’ Letterati 11(24):195–216.
Newberry, J.S. 1861. Part III. Geological report. In J. C. Ives. Report upon the Colorado River of the
West. U.S. 36th Congress, 1st Session, Senate Executive Document and House Executive Document
90(3).
Nordt, L., J. Bongino, S. forman, D. Esker, and A. Benedict. 2015. Late Quaternary environments of
the Waco Mammoth site, Texas USA. Quaternary Research 84:423–438.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
43
Nyborg, T. 2011. Age, stratigraphy and depositional environment of the Pliocene Copper Canyon
Formation, Death Valley, California. Ph.D. Dissertation. Loma Linda University, Riverside, CA.
300 pp.
Orr, P.C. 1968. Prehistory of Santa Rosa Island. Santa Barbara Museum of Natural History, Santa
Barbara, California.
Osborn, H.F. 1923. New subfamily, generic, and specific stages in the evolution of the Proboscidea.
American Museum Novitates 99:1–4.
Osborn, H.F. 1936. Proboscidea: A monograph of the discovery, evolution, migration and extinction of
the mastodonts and elephants of the world. Vol. 1. Moeritherioidea, Deinotherioidea, Mastodontoidea.
American Museum Press, NY. 802 pp.
Palombo, M.R. 2007. How can endemic proboscideans help us understand the “island rule”? A case
study of Mediterranean islands. Quaternary International 169–170:105–124.
Pasda, K. 2012. Seward Peninsula, Alaska: Trail Creek Caves 2 and 9 Revisited, The Skeletal Remains.
British Archaeological Reports International Series 2374. British Archaeological Reports,
Oxford, UK. 179 pp.
Perkins, M.E., and B.P. Nash. 2002. Explosive silicic volcanism of the Yellowstone hotspot: The ash
fall tuff record. Geological Society of America Bulletin 114:367–381.
Pigati, J.S., D.R. Muhs, and J.P. McGeehin. 2017. On the importance of stratigraphic control for
vertebrate fossil sites in Channel Islands National Park, California, USA: Examples from new
Mammuthus finds on San Miguel Island. Quaternary International 443A:129–1 39.
Pohlig, H. 1912. Sur une vieille mandibule de “Tetracaulodon ohiotocum” Blum, avec defense in situ.
Bulletin de la Société Belge Géologique 26:187–193.
Protas, J. 2002. A past preserved in stone: A history of Montezuma Castle. Western National Parks
Association, Tucson, AZ. 286 pp.
Prothero, D.R., E.E. Draus, and S.E. Foss. 2006. Magnetic stratigraphy of the lower portion of the
middle Miocene Mascall Formation, central Oregon. PaleoBios 26:37–42.
Quackenbush, L.S. 1909. Notes on Alaskan mammoth expeditions of 1907 and 1908. Bulletin of the
American Museum of Natural History 26:87–130.
Rasic, J.T. 2012. Mammoth Site Reconnaissance in Bering Land Bridge Preserve. Report on file,
National Park Service Fairbanks Administrative Center, Fairbanks, AK.
Rasic, J.T. and P. Matheus. 2007. A Reconsideration of Purported Holocene Bison Bones in Northern
Alaska. Arctic 60:381–388.
Rasic, J.T., and S. Shirar. 2009. Taxonomic Identification and Radiocarbon Dating of Large Mammal
Remains from National Park Service Collections at Trail Creek Caves, Seward Peninsula, Alaska.
Report on file, National Park Service Fairbanks Administrative Center, Fairbanks, AK.
Rasmussen, K. 1932. The Eagle’s Gift, Alaska Eskimo Tales. Doubleday, Doran and Company,
NY. 235 pp.
Reeder, W.G. 1987. Summary of vertebrate paleontological investigations, 1985–1987, Big Bend
National Park, Texas. Open file Report, Texas Memorial Museum, Austin, TX.
Repenning, C.A. 1967. Report on Referred Fossils, Shipment MAR-67-12M. Report on file, U.S.
Geological Survey Branch of Paleontology and Stratigraphy, Menlo Park, CA. 3 pp.
Repenning, C.A. 1970. Report on Referred Fossils, Shipment MAR-70-20M. Report on file, U.S.
Geological Survey Branch of Paleontology and Stratigraphy, Menlo Park, CA. 5 pp.
Repenning, C.A. 1982. Report on Referred Fossils, Shipment A-82-6M. Report on file, U.S. Geological
Survey Branch of Paleontology and Stratigraphy, Menlo Park, CA.
Reynolds, R.E., D. Miller, and K. Bishop. 2003. Land of lost lakes, the 2003 Desert Symposium
Fieldtrip. Pp. 3–26, In R.E. Reynolds (Ed.). Land of Lost Lakes. California State University Desert
Studies Consortium, Fullerton, CA. 68 pp.
Rhoads, S.N. 1903. The mammals of Pennsylvania and New Jersey: A biographic, historic, and descriptive
account of the furred animals of land and sea, both living and extinct, known to have
existed in these states. Privately published, Philadelphia, PA. 266 pp.
Rogers, H.D. 1840. Description of the geology of the state of New Jersey, being a final report. C.
Sherman and Co. Printers, Philadelphia, PA. 227 pp.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
44
Roth, V.L. 1993. Dwarfism and variability in the Santa Rosa Island mammoth (Mammuthus exilis): An
interspecific comparison of limb-bone sizes and shapes in elephants. Pp. 433–442, In F.G. Hochberg
(Ed.). Third California Islands Symposium: Recent Advances in Research on the California
Islands. Santa Barbara Museum of Natural History, Santa Barbara, CA. 661 pp.
Roth, V.L. 1996. Pleistocene dwarf elephants of the California Islands. Pp. 249–253, In J. Shoshani
and P. Tassy (Eds.). The Proboscidea: Evolution and Palaeoecology of Elephants and Their Relatives.
Oxford University Press, New York, NY. 472 pp.
Ruez, D.R. 2009. Revision of the Blancan (Pliocene) mammals from Hagerman Fossil Beds National
Monument, Idaho. Journal of the Idaho Academy of Science 45:1–143.
Sankey, J.T. 2002. Vertebrate paleontology and magnetostratigraphy of the Glenns Ferry and Bruneau
Formations (Plio-Pleistocene), near Murphy, southwestern Idaho. Pp. 52–100, In W.A. Akersten,
M.E. Thompson, D.J. Meldrum, R.A Rapp, and H.G. McDonald (Eds.). And Whereas … Papers
on the Vertebrate Paleontology of Idaho honoring John A. White, Volume 2. Idaho Museum of
Natural History, Occasional Paper 37, Pocatello, ID. 192 pp.
Santucci, V. L. 1998. Paleontological resource management of the Copper Canyon Fossil Track Site,
Death Valley National Park. Geologic Resources Division. National Park Service, Fort Collins,
CO. 9 pp.
Santucci, V.L. 2000. A survey of the paleontological resources from the National Parks and Monuments
in Utah. Pp. 534–556, In D.A. Sprinkel, T.C. Chidsey Jr., and P.B. Anderson (Eds.). Geology
of Utah’s parks and monuments. Utah Geological Association, Salt Lake City, UT. Publication 28.
Santucci, V.L., and T. Nyborg. 1999. Paleontological resource management, systematic recording, and
preservation of vertebrate tracks within Death Valley National Park, California. San Bernardino
County Museum Association Quarterly 46(2):21–26.
Santucci, V.L., W.P. Wall., and A. Mead. 1995. A preliminary survey of paleontological resources from
the Alaska Region national parks. Pp. 7–12, In V.L. Santucci, and L. McClelland (Eds.). National
Park Service paleontological research volume 2. National Park Service Natural Resources Technical
Report, NPS/NRPO/NRTR-95/16. National Park Service, Fort Collins, CO. 181 pp.
Santucci, V.L., J. Kenworth, and R. Kerbo. 2001. An inventory of paleontological resources associated
with National Park Service caves. NPS Geological Resources Division, Technical Report NPS/
NRGRD/GRDTR-01/02. TIC D-2231. National Park Service, Fort Collins, CO. 62 pp.
Santucci, V.L., J.P. Kenworthy, and C.C. Visaggi. 2007. Paleontological Resource Inventory and
Monitoring—Chihuahuan Desert Network. National Park Service TIC# D-500. National Park
Service, Fort Collins, CO. 113 pp.
Santucci, V.L., and J.P. Kenworthy. 2009. Paleontological resource inventory and monitoring—Klamath
Network. Natural Resource Technical Report NPS/NRPC/NRTR—2009/208. National Park
Service, Fort Collins, CO. 84 pp.
Santucci, V.L., R.B. Blodgett, W.P. Elder, J.S. Tweet, and J.P. Kenworthy. 2011. Paleontological
resource inventory and monitoring: Central Alaska Network. Natural Resource Technical Report
NPS/NRSS/NRTR—2011/510. National Park Service, Fort Collins, CO. 124 pp.
Santucci, V.L., J.S. Tweet, D. Bustos, T. Nyborg, and A.P. Hunt. 2014. An inventory of Cenozoic fossil
vertebrate tracks and burrows in National Park Service areas. Pp. 469–488, In M.G. Lockley and
S.G. Lucas (Eds.). Fossil footprints of western North America. New Mexico Museum of Natural
History and Science Bulletin 62.
Santucci, V.L. 2017. Preserving fossils in the national parks: A history. Earth Sciences History
36:245–285.
Saunders, J. J. 1970. The distribution and taxonomy of Mammuthus in Arizona. Ph.D. Dissertation.
University of Arizona, Tucson, AZ. 115 pp.
Saunders, J.J. 1996. North American Mammutidae. Pp. 271–279, In J. Shoshani and P. Tassy (Eds.),
The Proboscidea: Evolution and Palaeoecology of Elephants and Their Relatives. Oxford University
Press, New York, NY. 472 pp.
Schaaf, J.M. 1988. Bering Land Bridge National Preserve: An Archaeological Survey (Vols. I and II).
Research/Resources Management AR-14, National Park Service, Alaska Region. Anchorage, AK.
Schlesier, K.H. 1967. Sedna Creek: Report on an Archaeological Survey on the Arctic Slope of the
Brooks Range. American Antiquity 32:210–222.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
45
Schoenherr, A.A., C.R. Feldmeth, and M.J. Emerson. 1999. Natural History of the Islands of California.
California Natural History Guides 61. University of California Press, Berkeley, CA. 491 pp.
Schultz, C.B., L.G. Tanner, F.C. Whitmore, L.L. Ray, and E.C. Crawford. 1963. Paleontologic investigations
at Big bone Lick State Park, Kentucky: A preliminary report. Science 142:1167–1169.
Schultz, C.B., L.G. Tanner, F.C. Whitmore, L.L. Ray, and E.C. Crawford. 1967. Big Bone Lick Kentucky:
A pictorial history of the paleontological excavations at this famous fossil locality from
1962 to 1966. University of Nebraska Museum Notes 33:1–12.
Scott, E., K.B. Springer, and J.C. Sagebiel. 2017. The Tule Springs local fauna: Rancholabrean vertebrates
from the Las Vegas Formation, Nevada. Quaternary International 443:105–121.
Scott, E., K.B. Springer, J.C. Sagebiel, and C.R. Manker. 2006. Planning for the future: Preserving
and interpreting paleontology and geology in Joshua Tree National Park. New Mexico Museum
of Natural History and Science Bulletin 34:159–164.
Scott, E., and K.B. Springer. 2016. First records of Canis dirus and Smilodon fatalis from the late
Pleistocene Tule Springs local fauna, upper Las Vegas Wash, Nevada. PeerJ 4:e2151; DOI
10.7717/peerj.2151.
Scott, E., C.J. Bell, M.C. Carpenter, J.I. Mead, L.M. Spencer, S.L. Swift, and R.S. White. 2004. Preliminary
report on late Pleistocene vertebrates from Lake Mead National Recreation Area, Mojave
Desert, southern Nevada. Geological Society of America, Abstracts with Programs 36(5):54.
Scott, R.L., V.L. Santucci, and T. Connors. 2001. An inventory of paleontological resources from
the national parks and monuments in Colorado. Pp. 178–202, In V.I. Santucci and L. McClelland
(Eds.). Proceedings of the 6th fossil resource conference: 2001 a fossil odyssey. Technical Report,
National Park Service, Geological Resource Division, Lakewood, CO. NPS/NRGRD/GRD/
GRDTR-01/01. 224 pp.
Scrivner, P.J. 1984. Stratigraphy, sedimentology and vertebrate ichnology of the Copper Canyon
Formation (Neogene), Death Valley National Monument. Thesis. University of California, Los
Angeles, CA. 334 pp.
Scrivner, P.J., and D.J. Bottjer. 1986. Neogene avian and mammalian tracks from Death Valley National
Monument, California; their context, classification and preservation. Palaeogeography,
Palaeoclimatology, Palaeoecology 57:285–331.
Seemann, B. 1853. Narrative of the voyage of H. M. S. Herald during the years 1845–51, under the
command of Captain Henry Kellett. Volume II. Reeve and Company, London, UK. 324 pp.
Semprebon, G.M., F. Rivals, J.M. Fahlke, W.J. Sanders, A.M. Lister, and U.B. Göhlich. 2016. Dietary
reconstruction of pygmy mammoths from Santa Rosa Island of California. Quaternary International
406B:123–136.
Shirar, S. 2009. Subsistence in seasonality at a late prehistoric house pit in northwest Alaska. Journal
of Ecological Anthropology 13:6–25.
Shoshani, J., and P. Tassy. 2005. Advances in proboscidean taxonomy and classification, anatomy and
physiology, and ecology and behavior. Quaternary International 126–128:5–20.
Skinner, M.F., and F.W. Johnson. 1984. Tertiary stratigraphy and the Frick Collection of fossil vertebrates
from north-central Nebraska. American Museum of Natural History Bulletin 178:215–368.
Smith, G.J., and L.R.G. Desantis. 2018. Dietary ecology of Pleistocene mammoths and mastodons
as inferred from dental microwear textures. Palaeogeography, Palaeoclimatology, Palaeoecology
492:10–25.
Smith, P.S., and J.B. Mertie. 1930. Geology and mineral resources of northwestern Alaska. US Geological
Survey Bulletin 815:1–351.
Springer, K.B., and J.S. Pigati. 2018. Desert wetlands in Death Valley National Park and the regional
hydrologic response to climate change during the late Quaternary. Geological Society of America
- Abstracts with Programs 50(6): paper 128-2.
Springer, K.B., J.S. Pigati, and E. Scott. 2017. Vertebrate paleontology, stratigraphy, and paleohydrology
of Tule Springs Fossil Beds National Monument, Nevada (USA). Geology of the Intermountain
West 4:55–98.
Springer, K.B., J.S. Pigati, E. Scott, V.L. Santucci, and M. Ferlicchi. 2018a. Desert wetlands record a
new Pleistocene vertebrate fauna from Death Valley National Park. Journal of Vertebrate Paleontology,
Program and Abstracts 2018:220–221.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
46
Springer, K.B., J.S. Pigati, and E. Scott. 2018b. The geology and paleontology of Tule Springs Fossil
Beds National Monument, Nevada. U.S. Geological Survey, Reston, Virginia. Fact Sheet
2018–3038.
Springer, K.B., J.S. Pigati, C.R. Manker, and S.A. Mahan. 2018c. The Las Vegas Formation. U.S.
Geological Survey, Professional Paper 1839. Reston, VA. 74 pp.
Spurr, J.E. 1903. Descriptive geology of Nevada south of the fortieth parallel and adjacent portions of
California. U.S. Geological Survey, Bulletin 208. Washington, DC. 236 pp.
Stanford, D.J., J.W. Jordan, E.J. Dixon, and M.A. Jodry. 1990. Archaeological Reconnaissance in the
Great Kobuk Sand Dunes, Northwest Alaska. Current Research in the Pleistocene 7:44–47.
Stearns, R.E.C. 1873. [no title: paraphrase of comments in minutes of Academy’s regular meeting,
Monday, September 1st, 1873]. Proceedings of the California Academy of Sciences 5:152.
Stauffer, C.R. “1945” [at least 1948 based on dates in the article]. Some Pleistocene mammalian inhabitants
of Minnesota. Minnesota Academy of Science Proceedings 13:20–43.
Stevens, M. 1993. Stratigraphy and vertebrate paleontology of the Late Tertiary sediments, lower Tornillo
Creek, Estufa Canyon areas, Big Bend National Park. P. 13, In V.L. Santucci (Ed.). National
park Service Paleontological Research Abstract Vol. 1; National park Service technical Report
NPS/NRPEFO/NRTR-93-11. 67 pp.
Stock, C. 1929. A census of the Pleistocene mammals of Rancho La Brea, based on the collections of
the Los Angeles Museum. Journal of Mammalogy 10:281–289.
Stock, C, and E.L. Furlong. 1928. The Pleistocene elephants of Santa Rosa Island, California. Science
68(1754):140–141.
Tankersley, K.B., M.R. Waters, and T.W. Stafford. 2009. Clovis and the American mastodon at Big
Bone Lick, Kentucky. American Antiquity 74:558–567.
Tassy, P. 1996. Who is who among the Proboscidea? Pp. 39–48, In J. Shoshani and P. Tassy (Eds.). The
Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives. Oxford University
Press, NY. 472 pp.
Tedford, R.H., L.B. Albright, A.D. Barnosky, I. Ferrusquia-Villafranca, R.M. Hunt, J.E. Storer, C.C.
Swisher, M.R. Voorhies, S.D. Webb, and D.P. Whistler. 2004. Mammalian biochronology of the
Arikareean through Hemphillian interval (Late Oligocene through Early Pliocene Epochs). Pp.
169–231, In M.O. Woodburne (Ed.). Late Cretaceous and Cenozoic Mammals of North America.
Biostratigraphy and Geochronology. Columbia University Press, New York, NY. 391 pp.
Ten Brink, N.W., and C.F. Waythomas. 1978. Late Wisconsin glacial chronology of the north-central
Alaska Range: A regional synthesis and its implications for early human settlements. National
Geographic Society Research Reports 19:15–33.
Thorpe, E.D., V.L. Santucci, J. Tweet, and D. Weeks, 2017. Salinas Pueblo Missions National
Monument: Paleontological resources inventory. Natural Resource Report NPS/SAPU/NRR—
2017/1436. National Park Service, Fort Collins, CO. 78 pp.
Tikonov, A., A.D. Agenbroad, and S. Vartanyan. 2003. Comparative analysis of the mammoth populations
on Wrangel Island and the Channel Islands. Deinsea 9:415–420.
Tocci, A.M., V.L. Santucci, J. Tweet, K.B. Springer, and E. Scott. 2018. Joshua Tree National Park:
Paleontological resources inventory. Natural Resource Report NPS/JOTR/NRR—2018/1813.
National Park Service, Fort Collins, CO. 112 pp.
Tucker, S.T., R.E. Otto, R.M. Joeckel, and M.R. Voorhies. 2014. The geology and paleontology of
Ashfall Fossil Beds, a late Miocene (Clarendonian) mass-death assemblage, Antelope County
and adjacent Knox County, Nebraska, USA. Geological Society of America Field Guide 36:1–22.
Tweet, J.S. 2014. Paleontology of Mississippi National River and Recreation Area. Natural Resource
Report NPS/MISS/NRR—2014/888. National Park Service, Fort Collins, CO. 210 pp.
Tweet, J.S., V.L. Santucci, and J.P. Kenworthy. 2008. Paleontological resource inventory and monitoring:
Sonoran Desert Network. Natural Resource Technical Report NPS/NRPC/NRTR—2008/130.
National Park Service, Fort Collins, CO. 148 pp.
Tweet, J.S., V.L. Santucci, and J.P. Kenworthy. 2009a. Paleontological resource inventory and
monitoring, Southeast Coast Network. Natural Resource Technical Report NPS/NRPC/NRTR-
2009/197. National Park Service, Fort Collins, CO. 220 pp.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
47
Tweet, J.S., V.L. Santucci, J.P. Kenworthy, and A.L. Mims. 2009b. Paleontological resource inventory
and monitoring—Southern Colorado Plateau Network. Natural Resource Technical Report NPS/
NRPC/NRTR—2009/245. National Park Service, Fort Collins, CO. 452 pp.
Tweet, J.S., V.L. Santucci, and T. Connors. 2012a. Paleontological resource inventory and monitoring:
Mediterranean Coast Network. Natural Resource Technical Report NPS/MEDN/NRTR—
2012/640. National Park Service, Fort Collins, CO. 150 pp.
Tweet, J.S., V.L. Santucci, T. Connors, and J.P. Kenworthy. 2012b. Paleontological resource inventory
and monitoring: Northern Colorado Plateau Network. Natural Resource technical Report NPS/
NCPN/NRTR-2012/585. National Park Service, Fort Collins, CO. 540 pp.
Tweet, J.S., V.L. Santucci, and T. Connors. 2015. Paleontological resource inventory and monitoring:
Southern Plains Network. Natural Resource Report NPS/SOPN/NRR—2015/971. National Park
Service, Fort Collins, CO. 332 pp.
Tweet, J.S., V.L. Santucci, and T. Connors. 2016. Paleontological resource inventory and monitoring:
Mojave Desert Network. Natural Resource Report NPS/MOJN/NRR—2016/1209. National Park
Service, Fort Collins, CO. 594 pp.
Twenter, F.R. 1962. New fossil localities in the Verde Formation, Verde Valley, Arizona. Pp. 109–114,
In R.F.L. Weber and H. W. Peirce (Eds.). Guidebook of the Mogollon Rim region, east-central
Arizona. New Mexico Geological Society, Socorro, NM. New Mexico Geological Society 13th
Field Conference. 174 pp.
Urban, T. M., J. T. Rasic, C. Alix, D. D. Anderson, S. W. Manning, O. K. Mason, A. H. Tremayne, and
C. B. Wolff. 2016. Frozen: The potential and pitfalls of ground-penetrating radar for archaeology
in the Alaskan Arctic. Remote Sensing 8:1007.
Urban, T.M., D. Bustos, J. Jakeway, S.W. Manning, and M.R. Bennett. 2018. Use of magnetometry for
detecting and documenting multi-species Pleistocene megafauna tracks at White Sands National
Monument, New Mexico, U.S.A. Quaternary Science Reviews 199:206–213.
Urban, T.M., M.R. Bennett, D. Buston, S.W. Manning, S.C. Feynolds, M. Belvedere, D. Odess, and
V.L. Santucci. 2019. 3-D radar imaging unlocks the untapped behavioral and biomechanical archive
of Pleistoene ghost tracks. Scientific Reports 9:16470.
Vacek, M. 1877. Über österreichische Mastodonten und ihre Beziehungen zu den Mastodon-Arten
Europas. Abhandlungen der Kaiserlich-Königlichen geologischen Reichsanstalt 7(4):1–45.
Van Den Bergh, G.D., P.Y. Sondaar, J. de Vos, and F. Aziz. 1996. The proboscideans of the South-East
Asian Islands. Pp. 240–248, In J. Shoshani and P. Tassy (Eds.). The Proboscidea: Evolution and
Palaeoecology of Elephants and Their Relatives. Oxford University Press, New York, NY. 472 pp.
Vandiver, V.W. 1936. Geological report on White Sands National Monument. Southwestern Monuments
Supplement for May 1936:381–400. Special Report No. 5. Department of Interior, National
Park Service.
Veatch, S.W., R.W. Graham, and H.W. Meyer. 2004. High elevation Mammuthus from the Florissant Fossil
Beds National Monument, Colorado. Abstracts with Programs, Geological Society of America 35:381.
Veatch, S.W., D.M. Jarzen, E.B. Leopold, and H.W. Meyer. 2010. Preliminary results of an examination
of palynomorphs associated with a mammoth molar from the Florissant Fossil Beds National
Monument, Colorado. Abstracts with Programs, Geological Society of America 42:564.
Vinson, D. 1988. Preliminary Report on Faunal identifications from Trail Creek Caves. Pp. 410–438,
In H.R. Melchior and B.M. Gilbert (Eds.). Bering Land Bridge National Preserve: An Archeological
Survey. Volume 1. National Park Service, Alaska Region, Anchorage, AK.
Vinson, D.M. 1993. Taphonomic Analysis of Faunal remains from Trail Creek Caves Seward Peninsula,
Alaska. Thesis. University of Alaska Fairbank, Fairbanks, AK. 230 pp.
Visaggi, C.C., J.G. Johnson, A.S. Johnson, J.P. Kenworthy, and V.L. Santucci. 2009. Paleontological
resource survey-Amistad National Recreational Area. Natural Resource Report NPS/NRPC/GRD/
NRR-2009/133. National Park Service, Fort Collins, CO. 84 pp.
Voorhies, M.R. 1985. A Miocene rhinoceros herd buried in volcanic ash. National Geographic Society
Research Reports 19:671–688.
Voorhies, M.R. 1990. Vertebrate paleontology of the proposed Norden Reservoir Area, Brown,
Cherry, and Keya Paha counties, Nebraska. Division of Archeological Research, University of
Nebraska, Technical Report 82–09, Lincoln, NE, 138 pp and 593 pp appendix.
Eastern Paleontologist
J.I. Mead, J.S. Tweet, V.L. Santucci, J.T. Rasic, and S.E. Holte
2020 No. 6
48
Voorhies, M.R., and R.G. Corner. 1993. An inventory and evaluation of vertebrate paleontological
sites along the Niobrara/Missouri scenic riverways project corridors. Internal National Park Service
report. University of Nebraska State Museum, Lincoln, NE. 88pp.
Wailes, B.L.C. 1854. Report on the agriculture and geology of Mississippi. Lippincott, Grambo, and
Co., Philadelphia, PA. 371 pp.
Wheatley, C.M. 1871. Notice of the discovery of a cave in eastern Pennsylvania containing remains
of post-Pliocene fossils, including those of Mastodon, Tapir, Megalonyx, Mylodon, etc. American
Journal of Science and Arts, Series 3, 1:235–237.
Wick, S.L., and D.W. Corrick. 2015. Paleontological inventory of Big Bend National Park: The place,
the people, and the fossils. Big Bend National Park, Division of Science and Resource Management,
Big Bend National Park, TX. 202 pp.
Wiest, L.A., D. Esker, and S.G. Driese. 2016. The Waco Mammoth National Monument may represent
a diminished watering-hole scenario based on preliminary evidence of post-mortem scavenging.
Palaios 31:592–606.
Wilson, R.C. 1981. First extinct vertebrates from Mammoth Cave, Kentucky. Pp. 339, In Proceedings
of the Eighth International Congress of Speleology. International Union of Speleology. National
Speleoological Society, Huntsville, AL. 802 pp.
Wilson, R.C. 1985. Vertebrate remains in Kentucky caves. Pp. 168–175, In Caves and Karst of
Kentucky. Kentucky Geological Survey, Special Publication 12. Kentucky Geological Survey,
Lexington, KY. 196 pp.
Withers, K., and J.I. Mead. 1993. Late Quaternary vegetation and climate in the Escalante River Basin
on the central Colorado Plateau. Great Basin Naturalist 53:145–161.
Woodburne, M.O. 2004. Global events and the North American mammalian biochronology. Pp.
315–343, In M.O. Woodburne (Ed.). Late Cretaceous and Cenozoic Mammals of North America.
Biostratigraphy and Geochronology. Columbia University Press, New York, NY. 391 pp.