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Consumption of Fungal Sporocarps By CONSUMPTIONOF FUNGALSPOROCARPS BY YELLOWSTONE GRIZZLY BEARS DAVIDJ. MATTSON,U.S. GeologicalSurvey Forestand RangelandEcosystem Science Center,Colorado Plateau FieldStation, P.O. Box5614, Northern Arizona University, Flagstaff, AZ 86011-5614, USA, email: [email protected] SHANNONR. PODRUZNY, U.S. Geological Survey Northern Rocky Mountain Science Center, Interagency Grizzly Bear Study Team,Forestry Sciences Lab, Montana State University, Bozeman, MT 59717, USA, email: [email protected] MARKA. HAROLDSON, U.S. Geological Survey Northern Rocky Mountain Science Center, Interagency Grizzly Bear Study Team, ForestrySciences Lab, Montana State University, Bozeman, MT 59717, USA, email: [email protected] Abstact: Sign of grizzlybears (Ursus arctos horribilis) consuming fungal sporocarps (mushrooms and truffles) was observedon 68 occasions duringa studyof radiomarkedbears in the Yellowstoneregion, 1977-96. Sporocarpsalso weredetected in 96 grizzlybear feces. Mostfungi consumedby Yellowstone'sgrizzly bears were members of theBoletaceae (Suillus spp.), Russulaceae (Russula spp. and Lactarius sp.), Morchellaceae (Morchellaelata), and Rhizopogonaceae. Consumption of false truffles(Rhizopogon spp.) was indicatedby excavationsthat were deeper, on average(1.1 dm),than excavations for mushrooms (0.6 dm). Consumptionof sporocarpswas most frequent during September (7% of all activity), althoughmedian numbers of sporocarpsexcavated at feedingsites peaked during both August and September (22-23 excavations/site).Almost all consumption(75%) occurred on edaphicallyharsh sites typically dominated by lodgepolepine (Pinus contorta). At broadscales, consumption of sporocarpswas mostlikely where these types of lodgepolepine-dominated sites wereextensive or wherehigh-elevation sites supportingmature whitebarkpine (P. albicaulis) were rare. The number of sporocarpsexcavated at a feedingsite was greatest when cone crops of whitebarkpine were smalland in standswith abundant lodgepole pine. At finescales, consumption of fungiwas positively associated with lodgepole pine basal area and negativelyassociated with total ground vegetation cover. Becauseof thestrong association of sporocarpconsumption with lodgepole pine and its disassociationat broad scales with availability of whitebarkpine seeds, consumption of mushroomsand truffles by grizzlybears will likelyincrease in theYellowstone ecosystem with global warming. Lodgepole pine is predictedto increaseand whitebark pine to declinewith global warming. Ursus13:95-103 (2002) Keywords: foraging behavior, fungi, grizzly bears, habitat selection, mushrooms, sporocarps, Ursus arctos, Yellowstone Hypogeous and epigeous sporocarpsare often referred kistan (Roberts 1977), and Hokkaido (Odachi and Aoi to as truffles and mushrooms,respectively, and are com- 1987). In North America, heavy consumptionof fungi mon reproductivestructures of mycorrhizalfungi in tem- was observed among black bears in coastal Washington perate and boreal forests. These spore-producing state (Poelkerand Hartwell 1973). Among grizzly bears, structuresare eaten andefficiently digestedby omnivores morethan incidental consumption has been recordedonly such as humans and feral swine (Sus scrofa; Groot in the Yellowstone region (Mealey 1975, Mattson et al. Bruinderinket al. 1994). Moreover,fungal sporocarps 1991, Mattson 1997a). are present in the diets of numerousotherwise herbivo- Grizzly bearsin the Yellowstoneregion obtainmost of rous or frugivorous mammals including wapiti (Cervus theirenergy from seeds of whitebarkpine (Mattsonet al. Nelson and elaphus; Leege 1982, Collins and Urness 2001) or tissue of bison (Bisonbison) and wapiti(Mattson and tree 1983) squirrels(Sciurus spp., Tamiasciurusspp., 1997b),although there are marked differences in consump- and Glaucomysspp.; Hall 1981, Carey 1995, Steele 1998). tion of pine seeds andungulate tissue among bearsof dif- of animalsis Consumption fungi by consistentwith the ferentage andgender (Mattson 2000). Yellowstonegrizzly abundanceand nutritionalcharacteristics of mushrooms bears consume most pine seeds duringfall, primarilyby and truffles. Conifer forests of interiorNorth America digging whitebarkpine cones from ground-levelcaches can produce 10-17 kg of mushroomsper ha, dry weight, madeby red squirrels(Tamiasciurus hudsonicus; Mattson of 15-35% crude comprised protein(Mealey 1975, Fogel et al. 2001). Tissue is obtainedfrom ungulatesprimarily and This Trappe 1978). relatively high concentrationof during spring by scavenging winter-killed animals is protein complementedby a similarly high concentra- (Mattson 1997b). When pine seeds or carrionare scarce, tion of energy (4-6 kcal/g, dry weight), comparableto Yellowstone's grizzly bears eat diverse other foods, in- thatof most hardand soft fruits(Fogel and Trappe1978). cluding fungal sporocarps(Mattson et al. 1991). Based Some edible mushroomscontain as little as 13-18% total on field observationsof radiomarkedgrizzly bears, we fiber which dietary (Cheung 1997), equates to about 76- speculatedthat consumptionof sporocarpswas concen- 83% matter for and dry digestibility grizzly NorthAmeri- tratedin the fall and closely associated with pine trees in can black bears (U. americanus;Pritchard and Robbins forest standswith sparseground cover. 1990). In this paperwe describefeatures associated with con- The few recordsof ursids trufflesor mush- consuming sumption of fungal sporocarpsby Yellowstone grizzly rooms are limited to eitherentries in tables summarizing bears. Because virtuallynothing is known about exploi- fecal contents or to short anecdotes. Most observations tation of mushroomsand truffles by bears, the come from analysis Eurasia,including the Alps (Couturier1954), reportedhere is exploratory.We evaluatedthe effects of EuropeanRussia 1931, Novikov et al. Pa- (Ognev 1969), forest stand structure,date, type of bear,and availability 96 Ursus 13:2002 of known high-qualityfoods (i.e., pine seeds and ungu- radius forest inventoryplot at the center of grizzly bear late carrion) using data collected during a study of activity. Treeswere tallied,identified as live or dead, and radiomarkedanimals, 1977-96. the diameterof each was measuredat 1.4 m above ground. Percentground cover also was ocularly estimatedfor all graminoids,forbs, and shrubswithin about 10 m of plot STUDYAREA center. Each site was describedin terms of habitattypes The approximately23,000 km2study area, from 43?30" describedby Mattsonet al. (1999). There were 10 forest to 45? 15"Nlatitude and 109?30"to 111?30"Wlongitude, habitat types, including 2 that characterized recently correspondedto the known range of Yellowstone's griz- burnedor harvestedsites (Table 1). zly bearpopulation. Most of the areaoccupied by grizzly Field crews described all feeding signs and collected bears was >2100 m elevation and consisted of remote all feces (scats) encounteredduring 1977-96. During mountainsand plateaussurrounded by valleys and plains 1986-96 descriptionsof feeding activity includedcounts more intensivelysettled or used by humans. Annualtem- and measurementsof excavations. For excavations of peraturesaveraged about 0?C. Monthlyaverages ranged fungal sporocarps,length, width, and depth (all in dm) from -2?C to 13?Cduring April throughOctober. Tem- were measuredand multipliedto estimateexcavated vol- peraturesrarely exceeded 27?C in most areas. Precipita- ume. Excavationsof sporocarpsby bearswere identified tion variedin amountand timing with elevationand region by several criteriaincluding the presence of sporocarps, and fell mostly as snow that reached accumulationsof the absenceof otherpotential foods, the presenceof feces 20-260 cm, depending on location, before melting dur- containingsporocarp remains, the presenceof othergriz- ing March-June(Dirks and Martner1982). zly bear signs, and excavationdimensions. Comparedto About 75% of the study areawas forested. Lodgepole excavationsfor sporocarpsby otheranimals such as squir- pine was the most common dominant tree species, al- rels and wapiti, those by grizzly bears were the largest, though whitebarkpine was abundantabove 2,500 m el- averaging about 1.6 dm in width and 2.0 dm in length. evation (Despain 1990). Forest structure varied Sporocarpsjudged to be the same type as those consumed considerablyduring the study primarilyowing to mortal- by the bear were described and identified using a field ity of trees caused by fire andby epidemic populationsof key synthesizedfrom Smith(1975) and Miller (1980). In mountainpine beetle (Dendroctonusponderosae; Despain some instances, sporocarpswere collected and keyed in 1990). The largest fire occurredin 1988 and burnedap- the lab. The field key emphasizedvisible featuressuch as proximately560,000 ha. Craigheadet al. (1995) provide color, pattern,shape, size, structuresof the cap or stipe, additionaldetails aboutthe study area. site type, and whether the sporocarpwas epi- or hypo- geous. The concurrentcollection of scats and informationon METHODS feeding activity maximized comparabilityof data from Grizzly bears were trapped,marked, and radiolocated these 2 sources at broad scales. We did not ascribe scat accordingto methodsdescribed by Knightand Eberhardt contents to specific feeding sites because foods found in (1985) and Blanchardand Knight (1991). A subset of scats were often not consumed at the site of collection radiolocations was visited and described according to given the 7-13 hr transittime of digesta (Pritchardand methods in Mattson(1997a, 2000). Field crews also de- Robbins
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