Dung and Over: De
1989 Cranium, jrg. 6, no. 1, pag. 29-44, april 29
Pleistocene Dung and the Extinct Herbivores of the Colorado Plateau,
Southwestern USA
Jim+I. Mead and Larry+D. Agenbroad
Samenvatting
kan worden De auteur beschrijft hoe en welke informatie verkregen van fossiele mest van gro-
te herbivoren.
Op grond van de morfologie en de grote van de fractie ( drooggewicht in gr.) en soms op
kan welke het haarinhoud, van fossiele mest bepaald worden van herbivoor afkomstig is.
Bovendien kan uit de inhoud ervan conclusies getrokken worden over: de vegetatie, het dieet
van het dier, het seizoen, de ouderdom van het fossiel d.m.v. koolstofisotoop-datering, eventu-
ele veranderingen in het dieet, mogelijke oorzaak van uitsterven, de sexe van het dier dat de
in mest heeft achtergelaten en veranderingen het milieu.
overzicht locaties het Colorado Plateau waar is Er wordt een gegeven van (grotten) op mest
Mammuthus shastensis en Oreamnos gevonden van Bison sp., sp., Nothrotheriops (grondluiaard) harringtoni (uitgestorven berggeit).
De volgende conclusies worden getrokken over de dieetsamenstelling van de hiervoor genoemde
soorten van het Colorado Plateau:
de mammoet: verschillende plantensoorten, maar overwegend (1/3 deel van de totale mest- inhoud) grassen;
de uitgestorven berggeit: vnl. grassen en ook jonge scheuten;
de grondluiaard: vnl. jonge scheuten en daarbij ook grassen;
de bizon: merendeels grassen en daarbij jonge scheuten.
The vertebrate paleontologist is typically Although a wide variety of extinct megaher-
concerned with a plethora of isolated skele- bivores are known from skeletal remaines
tal remains from which to reconstruct (ANDERSON, 1984; KURTeN and ANDERSON,
extinct species or faunal communities. If 1980), only a few are recorded by preserved
lucky, the paleontologist will happen along dung and hair specimens. Here we review of
late land an articulated skeleton (THEWISSEN and the Pleistocene (Rancholabrean
mammal the FRANZEN, 1987), which permits an unusually age) dry cave localities of Colo-
detailed osteological examination of a parti- rado Plateau, Southwestern USA, that con-
cular species. The paleontologist will be able tain preserved dung deposits, and the variety
to describe the skeleton of an extinct animal of analyses that can be performed on the
than Because the fairly well, but more often not diet is unique remains. study of glacial-
at best hypothesized, based on tooth mor- age dung remains is in its infancy in North
phology, and the local plant community is America, we will examine only four of the
inferred from an associated pollen recon- best known species: Mammuthus
struction. (Proboscidea,Nothrotheriopsmammoth),
Water, the provider of life, is also the pro- shastensis (Xenarthra Megalonychidae, Shasta
motor of decay. Invariably the taphnonomic ground sloth), Bison (Artiodactyla, Bovidae
situation of an animal carcass is such that Bovini, bison), and Oreamnos harringtoni
Bovidae: stomach contents, skin, muscle, organs, and (Artiodactyla: Rupicaprini,
mountain hair/keratin decay away relatively quickly. Harrington's goat).
Even in arid regions of the desert South- Use of preserved dung for reconstrucution of
in west, USA or the Arctic, provide enough diet and environments began the arid
moisture to permit slow but certain decay of Southwest when EAMES (1930) examined
non-skeletal remains - unless protected. Two dung of extinct Shasta ground sloth from
environmental situations typically allow for New Mexico, and LAUDERMILK and MUNZ
soft-tissue preservation: identified macrobotanical fragments from
1) regions of the Holarctic permafrost (PO- similar dung from Gypsum Cave, Nevada
POV, 1948; VERESHCHAGIN, 1977; VERESH- (1934), and Rampart and Muav caves, Arizona
CHAGIN and NIKOLAEV, 1982), and (1938; LAUDERMILK, 1949).
the 2) xeric cave deposits of deserts MARTIN et al (1961) later identified pollen
(EAMES, 1930; MARTIN et al, 1961). from Shasta ground sloth dung in Rampart 30
Cave (western Grand Cannyon, southwestern- are rare. See MEAD and AGENBROAD (sub- most Colorado Plateau; fig. 1). IBERALL mitted 1988) for a more detailed field and
(1972; now, ROBBINS et al., 1984) went a laboratory guide to extinct and modern her- step further in paleoecological reconstruc- bivore dung of North America MURIE (1954)
when she used tions microhistological ana- provides the best overall field guide to dung lyses (plant cuticle and other remains) to identification of living North American mam- identity the diet of the extinct Harrington's mals (an equally good field guide to dung mountain goat form Stanton's Cave (northern identification of Australian native and intro- end of Colorado River in the Grand Canyon, duced animals is provided by MORRISON,
Arizona). HANSEN (1978) used the same 1981; an excellent book to observe how a technique to re-examine the Shasta ground dung guide is to be used). sloth dung from Rampart and Muav caves.
Long (LONG and MARTIN, 1974; LONG et Identification al., 1974) was the first to use multiple ra- diocarbon dating (beta particle technique) to Two analyses (morphology and size fraction; locate the last use of Rampart Cave by the and sometimes a third, hair content) are ground sloth. Analysis of dried dung on the used to identify the species of the herbivore greater Colorado Plateau began in earnest which produced the dung. Herbivores consu- when HANSEN (1980) provided the identifi- me large quantities of vegetation and much
from in of cations the fragmented remains Cow- the bulk is expelled as waste. Conse-
Utah boy Cave, (northern-most locality on quently, the dung is deposited as one large
It fig. 1). was this study that heralded a mass, or as a large number of fecal pellets new era of dietary and faunal community which are reasonably regular in size and studies in this arid region. shape, sometimes distinctly so. The dung,
when broken open, can be seen to be com-
Analyses posed of evenly cut twigs or chaff-like
vegetation. The size of the contents is often
a of indicative of of animals. Although variety analyses are currently a type or group conducted on preserved dung remains, it Overall morphology of a particular species of should be remembered that the study, in its dung can change somewhat with the moisture fullest spectrum, is just beginning. The use content of the ingested vegetation.
tool the in of dung as a for paleontologist is Proboscideans general produce an extre- endless. that Besides indicating a particular mely big bolus, larger than other dung mas- animal inhabited the area of the deposit, ses produced by other herbivores. Mammoth dung of extinct megafauna can be used: dung (fig. 2) is approximately 230 by 170 by
1) to determine some of the local plant com- 85 mm in size (DAVIS et al, 1984; MEAD et munity components, al., 1986c), full of graze remnants, and is
2) to directly reconstruct the diet identical in size and content to those bolu-
3) to identify the season(s) of occupation ses of the living elephants, Loxodonta and
direct 4) as a means to radiometrically date Elephas (WING and BUSS, 1970). Although the species the large size is a certain identifying cha-
determine the the 5) to possibly time of demise, racter, often boluses are trampled into the species smaller fragments (HANSEN, 1980) leaving
6) to define possible changes in diet through the identification to the size fraction ana- time, the up to time of extinction lysis. Fractional analysis of the dung con-
to examine of extinction 7) possible causes tents is expressed as dry weight in grams or
8) to identify the sex of species leaving the percent of dry weight according to size deposit (<0.25, 0.25 - 0.5, 0.5-1.0, 1.0-2.0,>2.0 mm
9) to environmental - identify changes to (fig.3). Lenghts and widths of grass stems
The name but a few uses. major items will and twigs in modern elephant dung measure be discussed here. up to 70.0 by 5.0 mm, similar to the 60.0 by
Analysis of dried dung is divided into three 4.5 mm observed in the mammoth dung from steps: Bechan Cave, Utah.
the 1) identification of producing species The Shasta ground sloth is a browser and
determination of the 2) contents (a large and produces dung boluses as linear-connected varied category) thick plates (MARTIN el al., 1961; MARTIN
3) paleoenvironmental and paleoecological 1975; fig.4). In addition to the distinctive reconstructions. The first step of identifica- shape, the twig contents are all clipped tion is the most critical analysis. Published short (HANSEN, 1980; SPAULDING and accounts and comparative collections of dung MARTIN, 1979). Size fractional analysis illu- 31
fig. 1 Map of the arid Southwest of the United States locating the Colorado Plateau
and the (stippled line) various caves containing late Pleistocene age dung deposits (dots).
fig. 2 A single bolus ofMammuthus from Bechan Cave, Utah. Note the coprophagus insect
made the holes when dung was still wet. 32
strates that Equus (horse) dung can be simi-
lar to that of the Shasta ground sloth (fig.
5). However, the size (width and length) of
the largest twigs of Nothrotheriops and My-
lodon (mylodont ground sloth, Chilean sam-
ple) is usually much shorter than that of
Equus, even when the horse eats woody
shrubs (fig. 6). Shasta ground sloth was the
first species of a North American extinct
megafauna to have a directly reconstructed
diet (MARTIN et al., 1985; THOMPSON et al,
1980).
Artiodactyls produce a variety of dung mor-
phologies. Bison dung will change from basi-
in cally circular plates aligned a row when
dry food is eaten, to the typical amorphous
of when pile ("chip") waste green or wet
vegetation is consumed (fig. 7). In conjuncti-
on with the distinctive shape, the size frac-
tion of the contents permit identification.
will Although graze versus browse ingestion
change the coar seness of the contents, the
overall fractional components and morphology
remain approximately the same.
Most artiodactyls produce relatively small
dung pellets. COE and CARR (1983) determi-
ned that with African there is a fig. 3 Size fractional analysis of various proboscidean dung. artiodactyls
fig. 4 Two views ofNothrotherops dung from Pampart Cave, Arizona, left view illustrating
the positioningof dung plates. Right view showing the morphology of the contents. 33
Grand Canyon was found to be, at adult
size, much larger and more cuboid in shape in comparison to the living mountain goat
(O. americanus), bighorn sheep (Ovis cana-
deer densis), (Odocoileus spp.), and prong- horn (Antilocapridae: Antilocapra americana).
the Dung from American moose (Alces alces)
and the muskox (Ovibovini, extinct and li-
ving genera) were found to be much larger
than those of Oreamnos. Only the living
wapiti (Cervus elaphus) produces (occasional-
ly) dung similar in size to that of the ex-
tinct mountain the goat, however, pellets are
never as cuboid as those that can be attai-
ned by O. harringtoni (fig. 8). The Grand
Canyon fossils, when found in direct associa-
skeletal tion with remains, are always with
Oreamnos never with harringtoni, any other
Most artiodactyl genus. of the deposits are in remote caves on cliffs, requiring entrance
by the best of agile climbers (MEAD et al.,
1986b). Many, but not all, of the living lar-
ge-pellet producers can be separated by dung
morphology (at adult size)(fig.8). The small
producers present some problems, however,
difficulty arises with the potential introduc- fig. 5 Size fractional analysis of dungfrom Nothrotheriops tion of extinct pellet-producing forms (e.g. and Equus. Navahoceros, ectinct mountain deer; Kurten
1975). Fractional size of the dung contents
the usually is same size, although the type
of forage will greatly determine the overall
size and shape. Richard Hansen (Colorado
State University; pers.comm.) has indicated
that the cuboid dung from the Grand Canyon
are best in caves identified, his view, as
Cervus; the extinct mountain goat, which is
30 % smaller in stature than the living spe-
should not cies, (COE and CARR, 1983) pro-
duce pellets twice as heavy as those of O.
americanus.
Hair is often found incorporated in the dung
of herbivores, being included with the in-
gestion of food by hide licking. Although
hair identification of extinct fauna is at
very preliminary stages in North America,
some haires have been identified (HANSEN,
1980; MEAD, 1983). Hairs of mammoth have
been identified from southwestern dung mats fig. 6 Width and length (mm) of the largest twigs in dung of (AGENBROAD and MEAD, 1987; HANSEN,
Nothrotheriops, Myolodin and Equus. 1980) and from frozen carcasses in Siberia
(SOKOLOV and SUMINA, 1982). steady increase in pellet with unit body weight gain. Whether this is true for all Contents pallet-producing ruminants is vet to be ter- mined. MEAD used the et al. (1986b) a ratio of After identifying species of animal re- the length and width of each pallet in com- presented by the dung, the most important parison to the pellet weight to distinguish analysis is the precise dating of the remains. the dung from a number of living and ex- Dung can be dated either indirectly (by its tinct North American artiodactyls (fig. 8). association with other dated remains) or
Dung of Harrington's mountain goat from the directly. MELTZER and MEAD (1985) have 34
the indicated that most reliable radiometric enough to be identified to species using a in determination for the extinction process dissecting microscope. Invariably seeds,
North America is the direct radiocarbon da- woody twigs, and leaves will be passed rela-
the ting of the concerned species. Once dung tively unaltered through the digestion tract,
been identified there is has to species, pro- especially in non-ruminants such as with the remain bably no better and no more direct mammoth, horse, and ground sloth (DAVIS et to date then the dung (and/or keratin). al., 1984, 1985; LAUDERMILK and MUNZ,
The fact that the material is preserved indi- 1934, 1938). Macrobotanical identification is
natural contamination the cates that virtually no probably most common dietary analysis exsists (usually contamination enters organic of Siberian mammoth dung (GORLOYA, 1982;
specimens via percolating water). Potentially KUPRIYANOVA, 1957; SOLONEV1CH and
minor contaminants that might pose a pro- VIKHIREVA-VASILKOVA, 1977; UKRAINTSE
if are easily removed during VA 1981, 1985; UKRAINTSEVA et al., blem, present, ,
normal sample preteatment (e.g. hydrocho- 1978).
loric acid washes). Ruminants, however, finely chew the vegeta-
A directly radiocarbon dated and identified tion so that macrobotanical analysis of dung
dung remain is a potentially powerful paleo- pellets is rarely useful. For these dung, in
ecological tool. The examination of trace addition to those taxa just mentioned, mi- elements was one of the first analyses ex- crohistological remains and pollen are ex-
tracted form dung, unfortunatly little paleo- tracted for ecological information MARTIN
ecological information has so far been gained et al. (1961) and SPAULDING and MARTIN
from these experiments (CLARK et al., 1974; (1979) used the pollen data preserved in the
MARTIN et al., 1961; PROVOTOROVA and dung of the Shasta ground sloth to recon-
RYABIKOV, 1982). struct the diet and local plant community
also The contents of dung are described by (see MARKGRAF, 1985, for Mylodon examimining the macrobotanical, palynologi- from South America) THOMPSON et al. cal, and microhistological remains. Macrobo- (1980) found that pollen in Shasta ground
does tanical remains are those plant remains large sloth dung not accurately reflect the
7 Two Bison view of view fig. morphological types of dung. Left round plates. Right a
Note fragment of a large ”chip”. that fractional sizes are the same. 35
of actual total ingested dietary vegetation Bison sp., Mammuthus sp. (cf. M. columbi),
HANSEN (1978, 1980) pioneered the use of Nothrotheriops shastensis, and Oreamnos
microhistology for obtaining the diet of ex- harringtoni recovered from deposits on the
tinct megafauna in North America. THOMP- Colorado Plateau (fig. 1). Dung has been
SON al. the first demon- directly radiocarbon dated back >40.000 et (1980) were to to yr B.P. strate that both pollen and microhistological (years before present) for a number of
data should be gathered from dung, because species. Radiocarbon dating of dung indicates
that the Shasta they are recording different sets of data, ground sloth became extinct
± 50 each important to paleoecological interpreta- by 11,018 yr B.P., a weighted average
dates tions. O'ROUKE and MEAD (1985), MEAD et of (MARTIN et al., 1985). Similar ana-
and al (1986b), and MEAD et al (1987) used pol- lysis of dung keratin (horn sheath) re-
len and microhistological remains for recon- mains of Harrington s mountain goat indica-
extinction finale structing the diet and season of diet for tes an by 11,160 ± 125 yr
Harrington's mountain goat. B.P. (MEAD et al., 1986a). An equivalent is in analysis using mammoth dung progress,
although preliminary examination form just
Recontructions Bechan Cave indicates that the elephant of
the Colorado Plateau may have left or been
the best reconstructi- exterminated slightly earlier (12,490 ± 40 As one might assume, yr
ons are those that use multiple lines of evi- B.P., weighted average of 13 tandem accele-
dence for a conclusion or a set of conclusi- rator mass spectrometer samples; MARTIN,
1987))(this does not take bias into ons. It is not the scope of this paper to sampling
review all the conclusions and reconstructi- consideration).
With continued direct radiocarbon ons arrived form dung analyses, however, dating of all identified it some of the more important findings from dung species, should be pos-
will sible to determine when these mammals be- the Colorado Plateau be examined, espe-
cially using the microhistological data. came extinct on, or at least extirpated from,
the Colorado Table 1 lists the localities containing dung Plateau.
Mammoths on the Colorado Plateau are found
to have eaten a variety of plants (Table 2),
of although most the diet seems to be graze
species. Nearly one-third of the encountered
macrofossils form Bechan Cave mammoth
dung are sedge (Carex); 95% of the contents
from dung recovered from Cowboy were Spo-
robolus grass (Table 2).
Table 3 reports the large variety of plants
eaten by the extinct mountain goat. Both
graze and browse were important in the diet,
and differed by season and location relative
to the Colorado River (MEAD et al., 1986b,
ROBBINS et al. 1984). Table 4 lists the
plants eaten by the shasta ground sloth. Al-
though it was mainly a browser, the dung
Muav samples from Caves, near a spring,
indicate that graze items occasionally were
important to the diet, possibly during spring
growth. The diet of the bison is just begin- ning to be established for the Colorado Pla-
teau (Table 5). Although it appears to use
predominantly graze, the bison obviously will
eat certain browse species.
8 ratio of to fig. Dung pellets of adult ruminants can otften be segregated by a weight
adult Oreamnus harringtoni width: length. Hachured area represents unquestionable dung. 36
Future directions analyses for paleoecology awaits the future
myriad of analytical techniques. We suggest
It is necessary to locate additional dung that those researchers working on the frozen
deposits and conduct all the above mentioned carcasses recovered from the permafrost and
tests on each species. If a particular deposit those working on the dried desert remains
contains, for example, three different layers begin working together to learn what effect
of dung, with bison, mammoth, and mountain the complexity of changing environments had
goat dung in each layer, then a sample of on the living and extinct megaherbivores of
each dung species should be tested from the last glacial.
each of detailed the three layers. Such ana-
ACKNOWLEDGEMENTS - Emilee lyses may begin to unravel the cause of ex- We thank
tirpation and extinction and when it may M. Mead for drafting the figures, providing have begun. the photography, and editing. Louella Coul-
The actual number of tests to be conducted ter, Bilby Research Center, Northern Arizona
remains on a dung is at the mercy of the University, provided typing services. There-
innovative scientist. We are beginning to sea Foppe, Composition Analysis Laboratory,
examine the biochemistry; Dr. Jerold Lowen- Colorado State University, provided the
stein (University of California) is searching microhistology analyses. Beta Analytic Incor-
for a way to extract a chemical signature porated, Florida, provided radiocarbon ana-
for each species of dung. This technique, if lyses.
succesful, will enable us to identify the spe-
cies belonging to the small pellet-producing
taxa and double-check identifications already LITERATURE
made. We might be able to identify extinct
species of deer ( Navahoceros) and pronghorn AGENBROAD, L.D. & MEAD, J.I., 1987: Late
(Stockoceros and Tetramervx), animals which Pleistocene alluvium and megafauna fung
probably produced dung pellets near identical deposits of the central Colorado Plateau. In:
to at least five other artiodactyls. Geologic Diversity of Arizona and its
Researchers at Johns Hopkins University Margins:
(Maryland) and San Diego Zoological Park Excursions to Choice Areas, by Davis, G.H. &
(California) are examining the possiblity of Van den Dolder, E.M., pp. 68-84. Field-Trip
determining the sex of the dung producer. Guidebook, the Geological Society of America With this it technique may be possible to 100th Annual Meeting, Phoenix, Arizona,
determine if a certain cave was used by one Arizona Bureau of Geology and Mineral in of sex or another, the form a birthing Technology Geological Survey Branch Special
area. Additional dietary contents (in dung Paper 5. and keratin remains) will be analyzed to see if there was a change in C3/C4 plant con- ANDERSON, E., 1984: Who's who in the
sumption during the late glacial. If there was Pleistocene: a mammalian bestiary.
such it will be In: a change, important to record Quaternary Extinctions: a Prehistoric
when it happened, and did it coincide with Revolution, by, Martin, P.S. & Klein, R.G.,
(cause) extinctions. pp. 40-89.
We have listed only four species of herbivo- University of Arizona Press, Tucson.
res known to produce dung which is preser-
ved in dry caves of alcoves on the arid Co- CLARK, F.E., O DEEN, W.A. & GELAU, B.E.,
lorado Plateau of the Southwest. Additional 1974: Carbon, nitrogen and 15N content of
species are preliminarily identified as be- fossil and modern dung from the lower
longing to Camelidae, Ovibovini, Caprini and Grand Canyon. Journal of the Arizona
Cervidae. It is unusual that dung be preser- Academy of Science, 9, 95-96.
ved at all, therefore, we must look for these
paleoecological remains in the unusual depo- COE, M.J. & CARR, R.D., 1983: The
sits. We must continue our research in the relationship between large ungulate body of Siberia permafrost and in the caves on weight and faecal pellet weight. African
the arid Colorado Plateau. We should be se- Journal of ecology, 21, 165-174.
riously examining the acidic, organic-preser-
ving bogs of northeastern Canada and Scan- DAVIS, O.K., MEAD, J.I., MARTIN, P.S. & dinavia. Caves in the arid lands such ad AGENBROAD, L.D., 1985: Riparian plants
Mongolia and northwestern China should be were a major component of the diet of
evaluated in terms of organic contents. The mammoths of southern Utah. Current
final determination of the usefulness of dung Research in the Pleistocene, 2, 81-82. 37
DAVIS, O.K., AGENBROAD, L.D., MARTIN, LAUDERMILK, J.D. & MUNZ, P.A., 1934:
P.S. & MEAD, J.I., 1984: The Pleistocene Plants in the dung of Nothrotherium from
dung blanket of Bechan Cave, Utah. Gypsum Cave, Nevada, Carnegie Institution
In: Contributions in Quaternary Vertebrate of Washington Publication, 453, 31-37.
Paleontology: a Volume in Memorial to John LONG, A. & MARTIN, P.S., 1974: Death of
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M.R. Carnegie Museum of Natural History,
Special Publications, 8, pp. 267-282. LONG, A., HANSEN, R.M. & MARTIN, P.S.,
1974: Extinction of the Shasta ground sloth.
EAMES, A.J., 1930: Report on ground sloth Geological Society of Amercia Bulletin, 85,
coprolite from Dona Ana Co., New Mexico. 1843-1848.
American Journal of Science, 20, 353-356.
MARKGRAF, V., 1985: Late Pleistocene
GORLOVA, R.N., 1982: Large remains of faunal extinctions in southern Patagonia.
plants from the stomach of the Shandrin Science, 228, 1110-1112.
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In: The Mammoth Fauna of the Asian Part of MARTIN, P.S., 1987: Late Quaternary extinc-
the USSR, Academy of Sciences of the tions: the promise of TAMS 14C dating.
USSR, 3, pp. 34. In: Nuclear Instruments and Methods in Phy- sics Research, B29, 179-186.
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fragments in the dungs of herbivores at 1985: Shasta ground sloth extinction: a test
Cowboy Cave. of the blitzkrieg model.
In: Cowboy Cave, by Jennings, J.D., In: Environments and Extinction: Man in
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D at l_ Rancholabrean,c CO CL 4-4 al. al. "ca directly O 13 «3 cn etat al.13 rH rH c reportOf 4-> 4-> 1988 et et ry 0) at recorded at cn 4-» etat D m 1981 1981 E i_ at etat et"aJ etat E Cl 2 2 •r- tn at 4-> •r- maximum/minimum at Late - -C 4-4 4-> ü> Preliminary this _J 13 "l3 > +-> l- <_ c in D! c. The - o (E Martin13 Martin LongO Emslie Robbins Mead Mead Mead Mead Davis LS ZT 21 z: z: o 1 -1 5! z: _J QC - 4it- LD5 LD6 c*7 9cn O 1 CM2 co3 800 co LR TR rH10 ?C' rH1 2 LR cn rH 00 ID *"• It* 2 2 2 1—4 2 8 co 10-14,19 cc 2,3,18 TR,4cc oc > O in15,16 17,20 LD15,16 TR,16QC TR,16 oz 5,15,6 t— N 7,8 cn e TR TR t— doudjd^duReference pTR -1 CO p TR TR 7 CN. 9 P TR age MajorJO pew ID Cn ID5 cn 2co LD5 LD5 1 1 6 -1 3CO 3co 3co 19 1 1 1 C\J 19 abrean 1,2 OO O ; O .r O ; ; ; ; ago 2 13 O O CO 00 00 § S rH +• + 1 +1 o -H -H -H +» in •H O Ranchol 13 cn O co O CO cn O oo years °-> • • - - >3 11,810±70; O 14,070+100; 24,190*2800! 30,600+1800;• >39,800;cn 13,505t580; 12,130±170; 13,040±440; 28,290±2100; >40,000; CO CO 2 late c |H A r—4 co co A rH rH CM -Q 1 1 1 1 -1 -1 1 i 1 • - 13 O O co O O O O O O O O O O O O O of LO O CO +1 CO co CM «—i ID CO rH co rH 1—4 2 CO +1 2 o O O O O O O O O O Radiocarbon■o O LD O O O O dung CO co OC ID O 00 00 co co LD r- O C0 00 LD CD O 10,650+220 10,035+250 A13,110+680;co 1—4 11,850+750 14,220+320 21,330+240 11,670+300 11,790±190 11,020+180 15,270±120 18,840±350 16,630±280 11,690+120 23,100+660 12,010+160 11,880+120 i—4 O O >11,0001—4 QC rH 1t- 1-H LD 00 LD CO CM LR LR USA. »—I < A -J rH co rH 1—4 rH CO containing pelletjailed a6je[large Xx Xx Xx Xx Xx Xx Xx Xx UMOU^UflUnknown >< X xX Xx X Xx Xx ?c- X harringtoniuioq-Bwiddvii x x Xx Xx x ?C- x southwestern eouiuveaoOreamnoa shelters X X X X X ahaatenaia x x x x edoidemoawoNNothrotheriopa Plateau, X Xx xX x Xx X Xx sandstone Mammuthua8ni{iTwtwvN xxx Xx Xx Xx xX Xx Xx X Xx and BiaonUOBIQ Colorado l_ at at 01 > 4-» 13 > Cave 'at Cave - o - - - - * 5 Shelter O - - the > Cave> in c oo O > r- g* CaveO C_> c at at > > u 4-4 4-4 Hollow Wallow"13 Limestone _o of Cave Cave Alcove O e at < c at at *-) in O BidaT- Kaetan -a Cave Cave Grotto CD Springs X o O "c J" o Caves t: IJ o Haven 1. 2 O* 2 o < a. 13 D 4-> at at +J in it c 13 at t. "at "at ra o O O 1 -X 4-> ation o Disappearing Muav Rampartl Sandblast Stanton's Stevens Three Tse'an Tse'an Bear Bechan Cowboy1 Grobot Hooper's Mammoth Wither's 2 BF OOak Ox herbivores o a z: OC l/> 00 oo h- h- \— co co ao O O □c z: o 5 Table ARIZONA UTAH| Loc_J < O 40 of percentage 1984; al., density. et Davis relative also of see 1%. 1985; percentage content. <_ al., 1980; 1987. amount, (continued) report. et macrofossil Presence. 2 this Davis Hansen, Martin, Trace * = = References: = = = T 1 2 3 X Table TR O +J 4J O Grotto(_ CU 9CT» QC 2CM »— +-> TR 89 O -OGrobot O on Cave> Based O 1 Tt— Th- >>CM2 1 1—T h-T Tt— r-H1 ir> 2CM »—» O 95CT» -§Cowboy O Plateau O X XX Colorado 3CO X XX X 4 Cave> Remains. O the T1— Th- 5 00 Tb- t— 1 c 1 CO O 3co ur> CM 13 12 18 11 IC 30 fH co on X3 Bechank CC 2 4 eTR CM *r Mammoth Macrofossil CX»94 the and co of 0 ö 0) co •I 1 R: co 9 3 § ■2 «-S Diets O ö neae £ co O +i 0J 5s neae Ü O stological co cx rQ <» •X <3 00 X s X X X X X X X X X X X X X X X X X Stanton'sStanton' Cave 3 2 2 C T 3 B Stevens Cave TRTR 14 14 T A T T T Tse'an KaetanKaetan CaveCave 1 10 T T Tse'anTse'an Bida Cave 1 T T 16 2A 52 Bear Ladder 2 6 OA (continued) Antennaria-Cirsium Boraginaceae ChrysothamnusChrysothammi8 3 Cryptantha AcaciaAcacia AdiantumAdiantwn Agave Aloysia Arenaria ArtemisiaArtemisia Aster AstragalusAstragalus Atriplex BaccharisBaaoharis BarkBark Cactaceae Cercocarpus Clematis ColdeniaColdertia Convulvus Cympteris DyssodiaDyssodia Table Reference: Browse s 1 X X X X X X X X X X X X X X X Stanton'sStanton CaveCave 3 on C 2 Based T T B 9 Stevens Cave TRTR 9 7 Plateau 6 T A 54 18 an 2 8 7 3 Colorado Tse'Tse'an Kaetan Cave 1 1313 the T T 7 T 1 4 2 T 3 4 10 19 on Tse'an BidaBida CaveCave Goat 3 6 9 3 2A2A Mountain Bear LadderLadder 2 OAOA the Remains. of Diets nardus Species: 3. Agropyron Andropogon AristidaArietida Bouteloua BromusBromua EleocharisEleodharis EnneapoganEnneapogan Equisetum FestucaFestuoa Hilaria Oryzopsis Panicum Phragmites SchedeSchedenardus Sporobolus Stipa Tridens Microhistological CarexCarex Poa Table Reference: Plant Graze 42 s X X X X X 3 X X X X X Stanton'sStanton' Cave levels C of below. 9 B 6 9 Stevens Cave TR Kaetan:x ABC and A cm 25 an density; T 1 5 2 T Cave 11 Tse'Tse'an Kaetan 11 density. density. levels an relative 50 T 1 2 2 1 T T 1 BidaBida Cave 2 T TseTse'an relative (X) relative of levels. 2A2A of all presence Bear 2Z Bear Ladder percentage of percentage 3 OA percentage Bida:x 1984; 1%. 1986; 1987; <_ 5; al., al., al., report; to et amount, (concluded) et et SymphoricarposSymphoricarpoe surface Robbins 3 Sphaeralcea Trace Rhus SarcobatusSarcobatus Sida SolanumSolarium Solidago Sphaeraloea TetradymaTetradyma Tragia Urtica Yucca This Mead Mead = Reference: References: = = = = 2 3 Table TR 1 T s X X X X X X XX X X X X X X X X Stanton'sStanton' Cave 3 X X X C 96 T B Stevens Cave TRTR ABC 54 3 A • an 1 6 2 T 1 1 T 1 Cave 1 22 Tse'anTse Kaetan an a 4 T 7 T T T 2 1 2 T Tse'anTse' BidaBid Cave 2A 24 Bear Ladder 2 16 OA 76 (continued) Glossopetalono Phlox-LeptodactylonPhlox-Leptodaetylon PhoradendronPhoradendron Pseudotsuga 3 Ephedra EriogonumEriogomun Euphorbia FallugiaFallugia Galium GutierreziaGutierrezia Hedeoma JuniperusJuniperua Laphamia Lesquerella LygodesmiaLygodeamia MentezeliaMentezelia OenotheraOenothera Opuntia PhysalisPhyaalia PinusPinua PopuluaPopulus Prunus P3eudot8uga Table Reference: 43 eau T T 4 1 T 3 4 1 T 16 Off Plateauat ColoradoColorado PI Bechan Cave TR 48 2TR 3 2 Mexico. CowboyCave 60 boluses. New 40 T T 1 T Rampart CaveCave 1 1 50 from southern 5 44 1 2 3 T 10 density. Cave, 5 2 C 5 3 Caves 12 representative; TRTR Shelter B 7 9 relative Muav ABC31 of 4 1980; A 4 sample 1%. 1974. <_ al., type type percentages 1978; 1980. al., et forb report. amount, (continued) type et are Ranunaulue-type Hansen, Hansen, Long Thompson Trace 4 ChrysothamnusChrysothamnue ForsellesiaForaeileaia FraxinusFraxinua Garrya Geranium Juniperus Oenothera-typeOenothera- PhaceliaPhaaelia ProsopisPro8opie PseudotsugaPeeudoteuga Prunus Ranunculus- Rosa-Roea- SphaeralceaSphaeraloea TidestromiaTidestromia UnknownUnknown YuccaYucca This = « = « ■ » References: 4 T Reference: 1 3 Reference: Numbers 2 Table TR 2 T 2 1 4 T Off Colorado Plateau 2 han 2 2 Microhistological Bechan Cave TR 46 Bee 2TR T T the 1 1 Cowboy Cave 2 36 on Only T 1 T T T T Rampart Cave 1 1717 Based T T T T T T 3 10 37 Sloth 5 3 2 5 Caveslves C 3 50 15 Ci Ground TRTR 9 B 2 Muav ABC31 14 16 27 16 Shasta A 2 2 6 44 10 60 1212 the pe ype of Diets Species:Species: ex Amelanchier-typeAmelanchiei*-type Chenopodium-tChenopodiwn-type 4. GramineaeGramineae AgropyronAgropyron AristidaAriatida BoutelouaBoutaloua MulenbergiaHulenbergia OryzopsisOryzopaia PhragmitesPhragmitea SporobolusSporobolua Tridena ArtemisiaArtemisia AtriplexAtriplex BumeliaBumelia CactaceaeCactaceae Bromus CarCarex FestucaFestuaa HilariaBilaria Tridens AcaciaAoacia Agave Table Remains. Reference: PlantPlant Graze Browse 44 2 2 2 MammothMammoth Alcove TR 47 30 17 s 1 T 3 T T 2 Hooper'sHooper Hollow TR 93 on Based 8B 1 Grotto 12 15 72 Plateau TR 2 9 2 5 2 Grobot A 3 3 74 density. Colorado chemistry. 1 1 2 4 5 T T Cowboy Cave 12 74 the relative relative on Bison percentage the Remains. percentage value. of type type 1980; 1% Diets report; _<_ Species: ex 5. Chenopodium-type AgropyronAgropyvon EquisetumEquisetum OryzopsisOryzopaia SporobolusSporobolua Chenopodium- Clematis- ChrysothamnusChryaothamrrue Ephedra Lupinus This Hansen, Trace; BromusBromua CarCarex Stipa Bark Ephedra Lupinue Moss PiceaPioea QuercusQuercue ïuoaaYucca Microhistological Reference: References: = = » BrowseBrowse 1 Table Plant Graze TR T