SHORT COMMUNICATIONSFood caching by bears: A literature review and new observations for
Asiatic and American black bears
Food caching, or storing food for later consumption,
is a behavior that preserves surplus food (MacDonald
1976, Smith and Reichman 1984). Caching often occurs
during times of resource abundance to extend the pulseof food, while also limiting detection of food by competitors. Caching has been observed in many taxonomic
groups; in mammals, well-known examples include many
species of rodents caching nuts or seeds (Smith and Reichman 1984) and Arctic foxes (Vulpes lagopus) cachingexcess eggs for later consumption during a limited waterfowl breeding season (Careau et al. 2007). Many large
carnivores cache the remainder of ungulate or other large
kills that are too large to consume in a single feeding bout,either in a tree (e.g., leopards [Panthera pardus]; Balmeet al. 2017) or covered with soil or other debris for later
consumption (e.g., pumas [Puma concolor]; Hornocker1970). Although caching of food has been observed inmany carnivores, there is variation among species in how
and why caching is used.
Bears are carnivores that have evolved to consume
large amounts of food quickly during periods of foodabundance and most bear species hibernate during seasons of food scarcity. For example, bears are well-known
to exploit pulses of food, such as neonatal ungulates (Bullet al. 2001, Bertram and Vivion 2002), patches of ripening berries or insects (Bull et al. 2001), and spawning fish
(Barker and Derocher 2009). The general feeding strategy of bears is to consume as much food as possible ina patch; but there are times, such as with large prey, thatfood needs to be stored for consumption over multiple
days. Although it is rare for most bear species—with the
exception of polar bears (Ursus maritimus) and brown
bears (U. arctos) in some areas—to regularly kill large
prey, many bear species exploit the carcasses of adultungulates or other large prey they find or usurp fromother carnivores (e.g., Cristescu et al. 2014, Elbroch et al.
2015). The carcasses of large prey often need to be consumed over a period of multiple days, and caching thesecarcasses could help preserve them and limit losses toother scavengers.
Although caching is a common behavior for brown
bears, less is known about caching by other bear species.
The absence of systematic investigations into caching behavior in many bear species results in confusion about
its prevalence and importance. For example, Craighead
and Craighead (1972) have been cited by multiple authors as evidence of caching by American black bears

Maximilian L. Allen1,7 , Heiko U. Wittmer2 ,Akino Ingaki3 , Koji Yamazaki4 , andShinsuke Koike5,61

Illinois Natural History Survey, University of Illinois,
1816 S Oak Street, Champaign, IL 61820, USA
2
School of Biological Sciences, Victoria University of
Wellington, P.O. Box 600, Wellington 6140, New
Zealand3
Graduate School of Agriculture, Tokyo University ofAgriculture and Technology, Fuchu, Tokyo, Japan
4
Department of Forest Science, Tokyo University ofAgriculture, Setagaya, Tokyo, Japan
5
Institute of Global Innovation Research, Tokyo
University of Agriculture and Technology, Fuchu, Japan
6
Institute of Agriculture, Tokyo University of Agriculture
and Technology, Fuchu, Japan

Abstract: Food caching is a common behavior for many
mammals, but less is known about the prevalence and importance of food caching for some species. Here we report the first documented caching events by Asiatic blackbears (Ursus thibetanus, n = 5) in Japan and 3 additionalcaching events by American black bears (U. americanus)
in California, USA. We also performed a systematic literature review on caching by bears as a reference point
for future investigations. Caching was most frequently
reported for brown bears (U. arctos), and most cachingby bears occurred with large prey. Caching is most likely
used to protect large carcasses from spoiling or detection by scavengers, allowing bears to consume more of
the carcass. The lack of published studies on caching bybears may be due to the behavior being infrequently usedand difficult to record. We encourage an increase, butalso consistency, in future reporting, including specificdescriptions of caching behavior.
Key words: American black bear, Asiatic black bear,cannibalism, food caching, foraging strategies, scavenging, Ursus americanus, Ursus thibetanus
DOI: 10.2192/URSUS-D-20-00008.1
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email: maxallen@illinois.edu

1
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2 SHORT COMMUNICATIONS(U. americanus), despite there being no mention ofcaching behavior reported anywhere in the manuscript.
Similarly, Svoboda et al. (2011) cite evidence of
caching by American black bears, but both studies cited(Elgmork 1982, Bertram and Vivion 2002) detail cachingby brown bears rather than American black bears. On
the other hand, the absence of caching observed during
mortality-site investigations of livestock has sometimes
been used to assign cause of mortality to American blackbears (Bertram and Vivion 2002). Here we report multiple instances of caching of ungulates by both American
black bears and Asiatic black bears (U. thibetanus). We
also performed a systematic literature review as a reference point for future investigations, on account of uncertainty regarding the prevalence of caching behavior bymost bear species.

Materials and methods
Study areas and field methods
We studied Asiatic black bears with Global Positioning System (GPS) collars in the Ashio–Nikko Mountainsof central Honshu Island, Japan. The study area has a wet
and cool-temperate climate. From 2006 to 2017, annual
precipitation averaged 2,236 mm and the annual mean
temperature was 7.2°C (Japan Meteorological Agency2018). Up to 1,600 m above sea level, the natural vegetation of this area is dominated by deciduous broad-leaved
forest composed of Quercus crispula, Q. serrata, Acerspp., and Fagus crenata (Furusaka et al. 2019). We havecaptured and fitted bears with GPS collars in the area
since 2003 and on average monitored 10 bears fitted withGPS collars each year. We randomly visited GPS location
clusters of collared bears after they had left the area, todetermine food items or record environmental conditions
associated with resting sites (Furusaka et al. 2019).
We also studied Asiatic black bears and other scavengers in Nikko National Park, central Japan. Precipitation averaged 1,631 mm/year and the mean annual
temperature was 7.7°C. Forest types included deciduous broadleaved forests, conifer plantation forests, andpatches of mixed forests, with an understory of bamboo
grasses in each forest type (Inagaki et al. 2020). In thisstudy, we deployed video camera traps on 49 sika deer
(Cervus nippon) carcasses to document scavenging activity. We obtained fresh deer carcasses from culled nuisance animals or animals killed through vehicle collisions
from June to November in 2016, 2017, and 2019, and secured each carcass to a tree with a wire rope. We monitored the carcasses using Ltl Acorn 6210 camera traps(Green Bay, Wisconsin, USA; Inagaki et al. 2020). We

programed cameras to record either 30-second videos ateach trigger with a 30-second refractory period or 30second videos at each trigger with a 1-second refractory
period.
We studied American black bears and other scavengers
in the Mendocino National Forest, California, USA. Climate in the Mendocino National Forest is considered
Mediterranean, with hot and dry summers and generally mild and wet winters. The mean annual rainfall was
1,320 mm and average daily temperatures ranged from
−1°C to 24°C (Allen et al. 2015a). The area is primarily
forested with conifers mixed with some deciduous trees,
but also contained chaparral and open grassland habitats. We deployed video camera traps on 100 Columbian
black-tailed deer (Odocoileus hemionus columbianus)
carcasses to document scavenger activity. We obtained
fresh deer carcasses from animals killed through vehicle
collisions from January 2010 to November 2012 (Allenet al. 2015b) and secured each carcass to a tree with a wire
rope. We monitored the carcasses using Bushnell ScoutCam camera traps (Overland Park, Kansas, USA), andprogramed the camera traps to record 60-second videos
at each trigger with a 1-second refractory period.
Literature review
On 8 February 2020 we performed a systematic literature search on Web of Science to find peer-reviewed documentation of caching among bears. We searched Web
of Science for the term “cach*” matched with the searchterm “ursus” and “bear” in English and Japanese. We
then read each entry and removed duplicate and mismatched publications, as well as those not from peerreviewed journals and studies involving captive animals.
The systematic search using Web of Science resulted in
few relevant studies (n = 5). We thus augmented our systematic review of the literature using snowball samplingby searching the references of papers that we reviewed
for additional articles (n = 11).

Results
Asiatic black bear observations
We documented caching by Asiatic black bears on 5
occasions.
1. On 23 October 2006, a 10-year-old female Asiatic
black bear initiated a cluster of GPS points (elevation
1,210 m). On 26 October the bear left the area after staying
3 days at the location (Table 1), and we investigated thelocation. We found a mostly consumed juvenile (3-yrold) Asiatic black bear carcass cached with leaves undera Japanese oak (Quercus mongolica; Fig. 1a).
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SHORT COMMUNICATIONS

3

Table 1. Summary of the common variables across the 7 instances we documented food caching by American and Asiatic black bears (Ursus americanus, U. thibetanus), observed in California, USA, and Japan,
respectively.
American black bear
Date

Sex and age

HabitatFood cached

Hardwood–conifer
forestDouglas fir forest

Scavenged
black-tailed deer
Scavenged
black-tailed deer
Scavenged
black-tailed deer

14 Nov 2011

Adult

6 Apr 2012

Unknown

24 Sep 2012

Adult female

Hardwood–conifer
forestDate

Sex and age

Habitat23 Oct 2006
6 Aug 201321 Nov 201710-yr-old female
9-yr-old-male
Adult

14 Jul 201911 Aug 201910-yr-old female
Unknown

Duration of visit
1 day

Other scavengers

5 daysGray fox, American blackbear
Gray fox, ringtail, common
raven, woodrat sp.
American black bear

Food cached

Duration of visit

Other scavengers

Oak forest
Oak forest
Conifer plantation

3-yr-old black bearSika deer
Scavenged sika
deer

3 days3 days
1 day

Riparian forest
Conifer plantation

Sika deer
Scavenged sika
deer

1 day
5 daysUnknown
Unknown
Asiatic black bear,
Japanese marten,
mountain hawk-eagle,
raccoon dog, red fox,
wild boar
Unknown
4 total bears

6 days

Asiatic black bear

2. On 6 August 2013, a 9-year-old male Asiatic black
bear initiated a cluster of GPS points (elevation 1,270m) along Chuzenji–Lake shore line in Nikko area, andstayed at the same location for 3 days (Table 1). Wheninvestigating the location, we found a mostly consumed
sika deer carcass cached with soil and rocks under theroot of a fallen birch tree (Betula platyphylla).
3. On 7 November 2017, we set out an adult sika deercarcass at an elevation of 660 m in a conifer plantation
forest under a Japanese cedar (Cryptomeria japonica). A
raccoon dog (Nyctereutes procyonoides) discovered thecarcass first on 20 November at 2308 hours. An Asian
black bear discovered the carcass on 21 November at
0005 hours (Table 1), while it was snowing. The bear began to scavenge and cache the carcass with fallen leaves
at 0021 hours, as recorded on 5 minutes of video (Video
1 thumbnail; see Supplemental material). The bear waspresent at the site for ࣙ2 hours and 7 minutes and fedfor ࣙ55 total seconds (recorded on 16 videos) over the
next day, but we did not document caching behavior after the initial visit. Another Asiatic black bear, red fox(Vulpes vulpes), Japanese marten (Martes melampus),
wild boar (Sus scrofa), and mountain hawk-eagle (Nisaetus nipalensis) were also documented scavenging at thecarcass (Table 1).
4. On 14 July 2019, a 10-year-old female Asiatic black
bear initiated a cluster of GPS points where she stayedfor 14 hours (Table 1). When we investigated the cluster,
we found a cached sika deer carcass buried by grass in ariparian forest with vegetation dominated by Clethra barbinervis. The carcass remains included the broken skull
and foot bones.
5. On 9 August 2019 we set out an adult sika deer carcass at an elevation 650 m, in a conifer plantation forest(Cryptomeria japonica and Chamaecyparis obtusa). On
11August we visited the site and found that the deer carcass was moved 20 m up a slope and was cached with
undergrowth in an open area dotted with Morus australis
between the conifer plantation forests (Fig. 1b). Basedon camera footage, ࣙ4 bears (2 solitary bears, and amother with a cub) visited the carcass until the carcass
was completely consumed on 13 August, but other scavenger species were not recorded (Table 1).
American black bear observations
We documented caching by American black bears on
3 occasions.
1. On 11 November 2011, we set out a field-dressedblack-tailed deer carcass at an elevation of 1,254 m, under a blue oak (Quercus douglasii) with a canopy coverof 76%. A gray fox (Urocyon cinereoargenteus) discovered the carcass initially on 13 November at 2139 hours,
and made 4 short visits, before an American black bear
discovered the carcass on 14 November at 2255 hours.

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4 SHORT COMMUNICATIONSVideo 2 Thumbnail. An American black bear (Ursus
americanus) caching a black-tailed deer (Odocoileus
hemionus columbianus) carcass in Mendocino National Forest.

Fig. 1. Documentation of caching by Asiatic blackbears (Ursus thibetanus) observed in the Ashio–
Nikko Mountains of central Honshu Island, Japan, including (a) the cache pile of a cannibalized Asiatic
black bear and (b) the caching a sika deer (Cervus
nippon) with vegetation.

Video 1 Thumbnail. An Asiatic black bear (Ursus
thibetanus) caching a Sika deer (Cervus nippon) carcass during a snowstorm.

During the initial visit of 13 minutes, the bear sniffedaround the carcass, but did not feed. It returned less thanhalf an hour later at 2346 hours and fed for 2 minutes. The
bear returned again on 15 November at 0014 hours, for avisit of 15 minutes where it investigated the bait and fed,
before caching the carcass with leaves. The bear returned
half an hour later and finished caching the carcass entirely
with leaves. We assume that the same bear returned for
multiple visits over the course of 6 days, and a female
with cubs also fed on the carcass for a few minutes.
2. On 28 March 2012, we set out a field-dressed blacktailed deer carcass in a Douglas fir (Pseudotsuga menziesii) stand at an elevation of 974 m with a canopy coverof 99%. A gray fox discovered the carcass initially on 1
April at 2202 hours, and a ringtail (Bassariscus astutus)
discovered the carcass on 4 April at 2206 hours. Bothspecies made multiple short visits to the carcasses, before an American black bear discovered the carcass on 6
April at 1836 hours. During its twelfth visit on 8 April,
the bear cached the carcass with needles and duff (Video
2 thumbnail; see Supplemental material for video), andsubsequently made 3 more short visits to the carcass overthe next 3 days. The carcass was subsequently visited bycommon ravens (Corvus corvax) and woodrats (Neotoma
sp.; Table 1).
3. On 24 September 2012, we set out a field-dressedblack-tailed deer carcass in at an elevation of 1,428 m,
under a ponderosa pine (Pinus ponderosa) with a canopy
cover of 89%. An American black bear discovered the carcass on 24 September and made multiple visits that night
until a female bear and her cub appeared at 2238 hours.
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SHORT COMMUNICATIONS
The original bear initially won a contest over the carcass,
but the female initiated another contest and usurped thecarcass from the original bear. The usurping female thenfed for 51 minutes, during which she cached the carcass
with pine needles and litter. The female and her cub fed
in an alternating pattern with what we assume was theoriginal bear until the carcass was completely consumed
on 29 September, but did not cache the carcass again.
Literature review
Most of the published studies of caching by bears had
small sample sizes (Table 2). Most reports were for brown
bears, with 2 studies each for polar bears and American black bears and our observations. Most studies reported caching based on observations during field investigations of clusters of bear activity or depredations.
However, one study used direct observations of predation and subsequent caching (Dean et al. 1986) and one
study used a mix of field investigations and direct observations over 45 years (Stirling et al. 2019), and we also
used camera-trapping at experimental ungulate carcasses.
The prey items that were reported cached were almost
always large prey (e.g., ungulates, marine mammals, orother large-bodied mammals), with the exception of fish(n = 1). Most studies had small sample sizes, and cachingmaterial was most often reported as vegetation and leaf
litter or soil, but polar bears always cached with snow
(Table 2).

Discussion
We documented 8 instances of black bears (Asiatic,n = 5; American, n = 3) caching large animal carcasses.
These are the first documented caching events by Asiatic
black bears, and only the third study to unequivocally
document caching by American black bears. The priorpublished literature on caching by bears was most frequently (75%) about brown bears, and caching may beless frequent for other bear species. For example, Stirling
et al. (2019) were only able to document 19 instances ofcaching by polar bears across 45 years of extensive fieldresearch and observations. The lack of published studies
on caching in most bear species may be because they areless often reported or of less interest to researchers, but ismost likely due to the behavior being infrequently usedand difficult to record (Stirling et al. 2019). Understanding the frequency of the behavior will help researchersand managers understand whether it plays an importantpart in their feeding ecology.
An important aspect of caching behavior for bears
is that it is most often used, or at least reported, for5large prey. For example, 6 of our cases were cachingof ungulates, and the other case was caching of a bear.
Similarly, in the prior literature, most cached prey were
ungulates, marine mammals, or bears; the exception
being the caching of multiple migrating broad whitefish(Coregonus nasus) by brown bears (Barker and Derocher2009). Two other reviewed studies documented brown
bears caching multiple individuals at the same site,
including ungulates (Cristescu et al. 2014) and domestic
sheep (Ovis aries; Elgmork 1982), but the caching ofmultiple individuals has not been reported for otherbear species. Caching of smaller prey may just not be
documented or reported as often as large prey, but it
appears to occur less frequently than with large prey.
This may be because caching is not needed as a resultof smaller prey being consumed more quickly, or thedifficulty of locating small prey compared with large
prey during field investigations.
That caching by bears is primarily for large food items(ungulates, livestock, and other large prey) likely indicates that the proximate mechanism for caching is thepreservation of food. Large prey take longer for bears
to consume, whereas the energetic costs of returning
to small prey that is cached can decrease the net energetic gain for bears. But large prey are also prone
to competition from scavengers and decomposers (e.g.,
Allen et al. 2015b). Covering food may decrease the rot
and consumption of large prey by decomposers, as well
as detection by scavengers. In most of the observations
we documented for black bears, they stayed in the vicinity of the large animal carcass for multiple days, often
staying until the carcass was completely consumed. The
effects of caching by bears for scavenger deterrence weremore equivocal. In Japan, one cached carcass we monitored with a camera trap was visited by 5 other scavengers,
whereas the other was visited by other Asiatic black bears.
Similarly, in the United States, 2 cached carcasses werevisited by other bears, whereas the other was visited by 4other scavenger species. In one instance, a pair of American black bears had 2 direct encounters where they fought
over the carcass.
Although there was a strong pattern of bears cachinglarge food items, other patterns were less clear. There
was little seasonal or geographic pattern, suggesting thatcaching occurs in most seasons and across the geographic
range of the bear species for which caching has been reported. Most reports were based on field investigationsof clusters of activity for collared bears, with 2 studies
also using direct observations, but most of our observations (5 out 8) were made using video camera traps. The
use of field investigations and camera-trapping are impor-

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6 SHORT COMMUNICATIONS
Table 2. The results of our review of peer-reviewed literature on caching by bears. We report the bear species,
location, caching events, type of food cached, and cache material.
Bear species
American blackbear
American blackbear
American blackbear
Asiatic blackbear
Asiatic blackbear
Brown bear

Location

Caching events

Food cached

Coahuila, Mexico

Not reported

Cattle (Bos taurus)

Michigan, USA1

Mendocino National
Forest, California,
USA
Ashio–Nikko
Mountains, Japan
Nikko National Park,
Japan
Alberta, Canada

3

White-tailed deer
(Odocoileus virginianus)
Columbian black-tailed
deer

Study

Not reported

Doan-Crider et al.
2017
Vegetation and litter Svoboda et al. 2011
Vegetation and litter This study2

Sika deer, Asiatic blackbear
Sika deer

Vegetation, soil and This study
litterVegetation and litter This study

85

Ungulates

Not reported

Not reported

Broad whitefish

Vegetation and litter

Brown bear
Brown bear

Northwest Territories,
CanadaHokkaido, Japan
Hokkaido, Japan

1
2

Female adult brown bearSika deer

Soil and debris
Soil, litter, and snow

Brown bear

Hokkaido, Japan

Not reported

Sika deer

Soil and vegetation

Brown bear
Brown bear
Brown bear
Brown bear

Hedmark, Norway
Hedmark, Norway
Vassfaret, Norway
Lapland Reserve,
Russia2
5
16
Not reported

Moose (Alces alces)
None
Sheep
Moose, reindeer (Rangifer
tarandus), cattle

Vegetation and litter
Vegetation and litter
Vegetation and litter
Snow, or vegetationand litter

Brown bear

Cantabrian
Mountains, Spain
Denali National Park,
Alaska, USA
Yukon Flats, Alaska,
USA
Svalbard, Norway

Not reported

Livestock

Not reported

1

Brown bear

Sand and Gravel

Not reported

Moose

Soil

2

White-beaked dolphin
Snow(Lagenorhynchus
albirostris)
Bearded seal (Erignathus Snowbarbatus), ringed seal
(Pusa hispida), harp
seal (Pagophilus
groenlandicus),beluga(Delphinapterus leucas),polar bear

Brown bear

Brown bear
Brown bear
Polar bear

Polar bear

Norway, Greenland,
and Canada

3

Cache material

19tant because otherwise caching events would most likely
only be encountered by chance. Camera traps are usefulfor documenting behaviors that are difficult to document(e.g., Allen et al. 2019), and could be used for detailed
studies of caching behavior in the future. It seems thatmost caching involved covering or burying carcasses invegetation or soil without excavating a hole or depression, but it is difficult to be sure in some accounts fromthe prior literature.
Our Web of Science search performed poorly, missing approximately two-thirds of the records in the

Cristescu et al.
2014
Barker andDerocher 2009
Kadosaki 1983
Kadosaki and Inukai
2000
Okada and
Yamanaka 2001
Mysterud 1973
Mysterud 1980
Elgmork 1982
Semenov-TianShanskii
1972
Clevenger et al.
1994
Dean et al. 1986
Bertram and Vivion
2002
Aars et al. 2015

Stirling et al. 2019peer-reviewed literature that we were eventually able
to find. Past studies may also have used other terms
for caching, such as covering, which could lead torecords being missed. For example, there were vague
or potentially conflicting terms in the reports, including ‘bury’ versus ‘cover’ versus ‘excavate’ versus‘burial mound’ (in Japanese). We encourage future studies to use the term ‘cache’ to describe when bears
cover their food with vegetation, soil, or other material, and to include specific descriptions of caching
behavior.
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SHORT COMMUNICATIONS

Acknowledgments
We thank the Illinois Natural History Survey, and the
University of Illinois for their support. We thank Y. Sugimoto and the members of Asian Black Bear ResearchGroup who helped with fieldwork, especially S. Haneo
and C. Kozakai. We also thank Dr. Y. Sato and Dr. K.
Kobayashi for providing information. We thank N. Selva,
B. McLellan, and 2 anonymous reviewers for commentsthat greatly improved the quality of the manuscript.

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Received: April 13, 2020
Accepted: July 3, 2020
Associate Editor: N. Selva

Supplemental material
Video S1. An Asiatic black bear (Ursus thibetanus)
caching a Sika deer (Cervus nippon) carcass during asnowstorm.
Video S2. An American black bear (Ursus americanus) caching a black-tailed deer (Odocoileus
hemionus columbianus) carcass in Mendocino National Forest.

Ursus 32:article e10 (2021)
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Terms of Use: https://bioone.org/terms-of-use Access provided by International Association for Bear Research and Management