Prey Weights for Computing Percent Biomass in Raptor Diets
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PREY WEIGHTS FOR COMPUTING PERCENT BIOMASS IN RAPTOR DIETS by Karen Steenhof Snake River Birds of Prey Research Project Bureau of Land Management 3948 Development Avenue Boise, Idaho 83705 Researchershave been assessingthe relativeimportance of prey speciesin raptor dietsfor manyyears. Early in thiscentury, biologists attempted to documentthe numberof pestspecies consumedby raptors (e.g., Kalmbachet al. 1964). More recently, biologistshave studied raptor dietsto understandthe effectsof land usechanges and environmentalcontaminants (e.g., Cade et al. 1968). Frequency of an individual prey speciesin the diet is not alwaysdirectly related to its nutritional importance(Southern 1954, Morris 1979). Raptorsmay consumeseveral small itemsthat provide lessweight and energy than a singlelarge prey item. To accountfor this, raptor dietsare now usuallyreported in termsof biomass: frequencyof a prey item multiplied by its averageweight (e.g., McGahan 1966, 1967;Porter and White 1973; Smithand Murphy 1973; Marti 1974). Accuracyof a biomassestimate depends on the accuracyof the weight assignedto a prey item. To ensureaccuracy, weights for eachprey speciesshould be categorizedby age and sex when appropriate.An averageadult weightwill distortrelative importance of a prey speciesif raptorsare consumingjuveniles. Similarly, an averageweight will distort resultsif one sexof a sexuallydiomorphic prey speciesis more vulnerableto raptor predation.Unfortunately, few studieshave consideredsize classes in computing biomassin the diet. Prey weightscan rarely be obtaineddirectly from pellet remains,partially consumed prey, or decomposedfood items found in nestsor under perches. Snout-vent lengths may be reliableindicators of snakeweights (BLM unpublisheddata), and Morris (1979) and Hamilton (1980) reported a usefulrelationship between rodent jaw lengthsand bodyweights. Unfortu- nately,similar relationships are not availablefor mostprey species,and in mostcases, weights of freshlycollected animals or averageweights reported in the literature mustbe used.During studiesof raptor ecologyin the Snake River Birds of Prey Area in southwesternIdaho, I compiled information on weightsof 116 raptor prey speciestaken by 9 speciesof raptors (Table 1). These weightsmay be useful to others investigatingpredator-prey relations. When possible,I usedprey weights obtained in the areaby BLM researchproject personnel. Nestling raptors and Common Ravens(Corvus corax) of variousages were weighted by BLM researchersin the nests;live cottontails(Sylvilagus nuttallii); woodrats (Neotomaspp.), and Townsendground squirrels (Spermophilus townsendii) were weightedduring trapping activities by BLM contractorsfrom the University of Idaho; dead rodentscaptured in snaptraps were weighedby contractorsfrom Utah StateUniversity; and reptileswere weighedby L. Diller, Universityof Idaho. Weightsof prey speciesnot measuredduring the studywere obtained from publishedliterataure. In addition,C. Robbinsand M. Fuller kindly providedweights for severalbirds from bandingrecords, L.C. Stoddartprovided weights for black-tailedjackrab- 15 Raptor Research17 ( 1): 15-27 16 RAPTOR RESEARCH Vol. 17, No. 1 bits(Lepus californicus), and M.R. Browningprovided Say's Phoebe (Sayornis saya) weights from filesat the NationalMuseum. I calculatedweights for prey itemsthat couldbe identifiedonly to classor genusby usingthe mean weightof identified individualswithin that classor genus that were taken by raptors. Sizeclasses of prey were assignedeither at the time remainswere collectedor when they were analyzed.Neonates included very smallmammals just emergingfrom nestsor burrows. Most other youngof the year birdsand mammalsthat were smallerthan adultswere classedas juveniles.An intermediateclass was used for fledging-agebirds, secondyear marmots(Mar- motaflaviventris)and rabbitsless than approximately3 monthsold but older than 1 month. Adults included any fully grown prey, and an averageclass was used for any prey item that couldnot be aged.Averages were calculated using relative proportions of knownsize classes in raptor diets.Juvenile weights for prey speciesthat showlarge weightgains over a shortperiod of time (e.g., Canada Goose (Branta canadensis);badger (Taxideataxus), and mule deer (Odocoileushemionus)) were estimated by consideringthe typicalsize of a young animal available to raptors during the nestingseason. Because of large seasonalchanges, weights assigned to Townsend ground squirrelsdepended on the monthsground squirrelswere found in nests. This paper is a contributionfrom the Bureau of Land Management'sSnake River Birds of Prey ResearchProject. I thank M.J. Collopy and J.S. Marks for assistancein reviewingthe literature and G.W. Smithand N.C. Nydeggerfor assistancein tabulatingrodent weights.This compilation would not have been possiblewithout the efforts of M.N. Kochert, A.R. Bam- mann,J.H. Doremusand the many biologistsand technicianswho workedon the project.J.S. Marks, M.Q. Moritsch and M.N. Kocher reivewed draft manuscriptsand made helpful suggestions.J.A. Gessamankindly providedaccess to an unpublishedcompilation of literature on bird weights. Table 1. Weights of Prey SpeciesCaptured by Raptors Species Size Class & Sex Wt(g) N Reference MAMMALS: Shrew-unid. Average 6 (1) BLM Data (Sorexspp.) PallidBat Average 32 Burt & Grossenheider 1964 (Antrozouspallidus ) Bat-unid. Average 10 (2) Porter & White 1973 (Myotisspp.) Long-tailedWeasel Juvenile 85 Palmer 1954 (Mustelafrenata) Adult 178 Smith & Murphy 1973 Badger Neonate 2833 Estimated (Taxidea taxus) Coyote Juvenile 2043 Estimated (Canis latrans) DomesticCat Average 1800 Estimated (Felisdomesticus) Spring1983 Steenhof-- PreyWeights 17 Table 1. Weightsof Prey SpeciesCaptured by Raptors(cont.) Species Size Class & Sex Wt(g) N Reference Yellow-bellied Marmot Neonate 500 Armitageet al. 1976 (M armotafla viventr•) Juvenile 1000 ........ Intermediate 2346 (38) ....... 2530 (10) ........ 2280 (28) Adult 3222 (99) 39o0 (38) 2800 (61) ...... Average 1808 (147) BLM Data TownsendGround Squirrel Juvenile:April 79 (480) BLM Data (Spermophilustownsendii) Juvenile:May 120 (1282) " " 127 (646) .... 114 (636) " " Juvenile: 199 (1331) .... June-July 184 (751) " " 164 (580) " " Adult:April 205 (1188) " " 254 (440) " " 178 (748) " Adult:May-June 222 (750) " " 277 (285) " " 188 (465) " " Average:April 176 (3053) " " Average:May-July 177 (4501) " " White-tailed Antelope Squirrel Juvenile 4O Estimated (Ammospermophilusleucurus ) Adult 106 (12) Hall 1946 111 (6) 101 (6) Average 105 (40) BLM Data Groundsquirrel-unid. Juvenile 127 Calculated Adult 225 Average 181 LeastChipmunk Average 32 (108) Schreiber 1973 (Eutamias minimus ) Townsend PocketGopher Juvenile 100 Estimated (Thomomystownsendii) Adult 248 (4) Hall 1946 261 (3) "" 236 (1) "" Average 200 Calculated Great Basin Pocket Mouse Juvenile 10 Estimated (Per o gnat hus pa rvus ) Adult 17 (508) BLM Data Ord KangarooRat Juvenile 28 Estimated (Dipodomysordii) Adult 53 (31) Schreiber 1973 Harvest Mouse Adult 11 (43) Schreiber 1973 (Reithrodontomysmegalotis ) Deer Mouse Juvenile 10 Estimated (Peromyscusmaniculatus) Adult 19 (145) Schreiber 1973 18 RAPTOR RESEARCH Vol. 17, No. 1 Table1. Weightsof PreySpecies Captured by Raptors(cont.) Species Size Class & Sex Wt(g) N Reference GrasshopperMouse Adult 26 (76) BLM Data (Onychomysleucogaster) Mouse-unid. Juvenile 10 Calculated Adult 17 " Desert Woodrat Juvenile 75 Estimated (Neotomalepida ) Adult 124 (10) BLM Data • 137 (6) " " 9 105 (4) " " Bushy-tailedWoodrat Juvenile 155 (7) Martin 1973 OVeotomacinerea ) Adult 338 (32) " " ff 405 (16) " " 9 271 (16) " " Average 277 " " Woodrat-unid. Juvenile 195 (45) BLM Data OVeotomaspp.) Adult 326 (87) " " 335 (70) " " 275 (16) " " Average 281 "" Muskrat Juvenile 1065 Donahoe1966 (Ondatra zibethica) 10971032 "". Adult 129812561277 """ " Average 1171 (1895)(16) "" House Mouse Average 19 BLM Data (Mus muscutus) Montane Vole Juvenile 15 * (Microtusmontanus) Adult 50 " 9d' 406035 " Average SagebrushVole Average 30 Burt & Grossenheider 1964 (Laguruscurtatus ) Rodent-unid. Juvenile 10 Estimated Adult 5o Estimated Average 5o n Porcupine Adults 58oo Smith pers.comm. (Erethizondorsatum ) Black-tailedjackrabbit Fetus 20 (Lepuscalifornicus ) Neonate 100 Stoddartpers. comm. Juvenile 5oo Spring 1983 Steenhof- Prey Weights 19 Table1. Weightsof PreySpecies Captured by Raptors(cont.) Species Size Class & Sex Wt(g) N Reference Black-tailedJackrabbit Intermediate 1000 Stoddart pers. comm. (Lepuscalifornicus) Adult 2114 ..... 1885 2344 ..... Average 1536 Calculated Mountain Cottontail Neonate 100 BLM Data (Sylvilagusnutta!!ii) Juvenile 215 Intermediate 500 " Adult 650 (92) " " 590 (45) " " 720 (47) " " PygmyRabbit Adult 340 Burt & Grossenheider 1964 (Sylvilagusidahoemis) Rabbit-unid. Neonate 100 Calculated Juvenile 404 " Intermediate 1087 " Adult 1550 " Average 927 " Mule Deer Juvenile 3780 McGahan 1966 (Odocoileushemionus) PronghornAntelope Neonate 2700 Beuchner 1950 (Antilocapraamericana) BIRDS: Great Blue Heron Average 1905 (1) Poole 1938 (Ardeaherodias ) Canada Goose Juvenile 450 Estimated (Branta canadensis) Mallard Adult 1185 (3226) Bellrose 1976 (Anaaplatyrhynchos) 1248 (1809) " " 1107 (1417) " " Northern Pintail Adult 976 (556) Bellrose 1976 (Anaa acura) 1025 (390) " " 866 (166) " " AmericanGreen-winged Adult 316 (192) Bellrose 1976 Teal 322 (113) " " (Anaacrecca ) 309 (79) " " Blue-wingedTeal Adult 395 (164) Bellrose 1976 (Anasdiscors ) 463 (35) " 377 (129) " 20 RAPTOR RESEARCH Vol. 17, No. 1 Table 1. Weightsof Prey SpeciesCaptured by Raptors(cont.)