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DECOMPOSITION of RAPTOR PELLETS by Jamesr DECOMPOSITION OF RAPTOR PELLETS by JamesR. Philips and Daniel L. Dindal Departmentof ForestZoology StateUniversity of New York Collegeof EnvironmentalScience and Forestry Syracuse,New York 139•10 Abstract Deco•nposingpellets from Great Horned Owls were studiedduring the summer and fall of 1973. After four months,pellets weighed 38-89 percent of their initial dry weight. Thirteen speciesof fimgi were isolatedfrom the pellets. Funnel extraction yielded3,665 invertebratesfrom 75 pellets.The most importantinvertebrates in pel- let decompositionappear to be the trogid beetlesand the tineid moths.Raptor pel- lets serve as a feeding,breeding, and sheltersite for many invertebrates,including predators,parasites, fi•ngivores, and saprovores. Introduction Indigestiblematerial is quite commonly regurgitatedby birds in the form of roughlyoval to cylindricalpellets. Such regurgitation is known to occur in 330 spe- cies in more than 60 families,according to a surveymade by The InternationalBird Pellet Study Group (Glue 1973). Among them are suchbirds as jays, flycatchersand thrushes,as well as hawks and owls (Simmons1973, Tucker 1944). Raptor pellets generallyconsist of fur, feathers,bone, scales, chitin, and other prey remains,densely packedand held togetherby mucussecreted from the digestivetract which driesand hardenssoon after egestion.In somecases, pellets of wax (HoneyBuzzard, Perhis api- vorus,Meinertzhagen 1959) or even soil (Kestrel,Falco tinnunculus,Davis 1975) may be found.Pellets are alsoejected by other vertebrates,including some mammals, rep- tiles, and amphibians(Hanson pers. comm., Dobroruka 1973). Pelletsprovide data on avian foodhabits and prey populations,but many other as- pects are of interestas well. Smith and Richmond(1972), Duke et al. (1973, 1975, 1976),and othershave examinedthe physiologyof pellet formationand regurgitation. Dobrokhotovand Litvin (1971)studied time-related dynamics of Kestrelpellet forma- tion by measuringpellet radioactivity after the raptoringested mice containing p32. Raptor pelletsmay alsoplay a role in the spreadof diseasessuch as sylvaticplague, since they are temporarilyinfective if diseasedprey has been eaten (Jellison1939). Plagueantigens can be detectedmuch longerthan the plaguemicrobe itself in pel- lets, and pellet analysishas been proposedas one of the best methodsof detecting epizooticplague infestations (Lobachev et al. 1971, Lobachevand Shenbrot1974). Pellet decompositionhas been given very little attention.Some authorshave re- ported that pelletsmay remainintact for many years(Brooks 1929, Presttand Wag- staffe 1973). Others have found that pellets are broken down in weeks or months (Wilson1938, Marti 1974).Warling (1962, 1963) studiedfungal succession on Kestrel 102 RaptorResearch 13(4):102-111 Winter 1979 Philips& Dindal--RaptorPellets 103 (Falcotinnunculus) pellets and isolated51 speciesof fungi and one alga. Hub•ilek (1974)isolated many fungalspecies from owl pellets.The invertebratefauna reported from pelletsis limited to a few insects(table 1). Therefore,the objectiveof this re- searchwas to elucidatethe invertebratemicrocommunity associated with decomposi- tion of owl pellets.The conceptof microcommunityas presentedby Dindal (1973a) wasassumed in the study. Table 1. Insect Fauna of Raptor Pellets Order References Family Coleoptera DermestidaeCarpet beetles Wallace, 1948 Dermestesspp. Simmons, 1971 SilphidaeCarrion beetles Hanson, pers. comm. TrogidaeSkin beetles Petersen, 1960 Trox erinaceousJ. Lec. Davis, 1909 Troxfoveicollis Harold Vaurie, 1955 Troxplicatus Robinson Vaurie, 1955 Trox scaber(L.) Davis, 1909 Troxspinulosus simi Robinson Vaurie, 1955 Trox striatus Mels. Vaurie, 1955 Trox variolatus Mels. Dillon and Dillon, 1972; Vaurie, 1955 Diptera AnthomyiidaeAnthomyid or latrine flies Fannia aerea (Zetterstedt) Aubrook, 1939 Itydrotaeaocculta (Meigen) Aubrook, 1939 ScatophagidaeManure-feeding flies Scatophagasqualida Meigen Aubrook, 1939 ScenopinidaeWindow flies Scenopinussp. de Joannis,1899 Hymenoptera BraconidaeBraconid parasitoid wasps Phaenocarparuficeps (yon Es.) Aubrook, 1939 Lepidoptera Tineidae Clothes moths Morton et al., 1977; Vernon, 1972 Monopisferruginella (Hubher) Hinton, 1956 Monopisrusticella (Clerck) Nurse, 1906; Buxton, 1914; Elton, 1966 Monopisweaverella Scott Elton, 1966 Tineapellionella (L.) de Joannis,1899 Tincolabisselliela (Hummel) de Joannis,1899 Trichophagasp. Moon, 1940 Trichophagatapetzella (L.) de Joannis,1899; Forbes, 1923; Baer, 1924;Davis (cited by Lutz, 1948) Methods Rats (Rattus norvegicus)were fed to five Great Horned Owls (Bubo virginianus). Pellets and pellet fragmentswere collectedas soonas possibleafter regurgitation. They were dried in an oven at 60øC for 24 hours.Dry weight and volume were 104 RAPTOR RESEARCH Vol. 13 No. 4 measured.The dry pelletswere placedin 6-mm meshnylon bagsand frozen. On 3 July 1973, 90 "pellet bags"were set out in a maple-oakstand at our College'sForest Experiment Station, Lafayette Road, Syracuse,New York. The pellets varied in weight from 3.08 to 11.47 g (av. 6.06); lengthranged from 33 to 86 mm (av. 49.71) and width, from 20 to 34 mm (av. 25.11). Six pelletswere removedevery 8 daysfor 4 months.Five pelletswere placedin modifiedTullgren funnels for extractionof invertebrates.As the pelletsair-dried, des- iccationdrove out the invertebrates,which fell into a collectingjar containingan al- coholand glycerinsolution placed at the baseof the funnel.The remainingpellet was used for culturingof fungi and invertebrates.Portions of the pellet were in- cubated on moist filter paper in a petri dish in the dark at 25øC. When insects hatched,they were preservedin alcohol.As fungalgrowth became evident, pieces of the pellet were transferredonto mycobioticagar. The other half of the pellet was usedfor culturingof fungi by meansof the soil-dilutionagar plate method(Menzies 1965). Dilutions from 10-4 to 10-6 were used with preparationsof mycobioticand malt agars. Results and Discussion After 4 months'exposure to field conditions,pellets weighed from 38.03 to 89.53 percent of their initial dry weight, averaging62.87 percent (fig. 1). Decomposition rates were highly variable because of differencesin size, composition,micro- environment,and faunaland floral colonizationof pellets. PhysicalFactors. Weather is a major factor in pellet decomposition.Rain flattens the pellet, dissolvesthe mucus,washes hair away from bonesnear the surfaceof the pellet, and helpscreate entry holesfor invertebrates.Freezing and thawingare also important(Wilson 1938). Faunal Microcommunity.Fresh moist pellets attract flies, suchas Calliphoridaeand Anthomyiidae,which deposittheir eggson the pellets.Such flies do not appearto be attractedto dried or older pellets.Other flies may be foundon older, moistpellets; however,pellets may at times representislands of moistureon the soil litter surface and attract cryptozoans,such as sowbugsand earthworms.Snails and slugscan also be found on pellets,and the large slugLimax maximusoften leavesa trail of mucus on pellets it has visited nocturnally.Various transientspecies, such as ants, centi- pedes,millipedes, and spiders,also visit pellets,as well as occasionalhungry mam- mals (Wilson 1938, Fleay 1968) and industriouspack rats (Griffin 1976, Long and Kerfoot 1963). Pelletssupport large populationsof protozoaand nematodes,and even tardigradesmay be present. Pelletsserve as habitat for beetlessuch as Leptodiridaeand Staphylinidae.Various parasiticwasps find pelletsto be a sourceof hosts.Predatory, parasitic, fungivorous, and saprophagousmites all were found.Akimov and Shehur(1972) have shownthat the acarid mites Tyrophagusputrescentiae (Schrank), Glycyphagus domesticus (Deg.), and Rhizoglyphusechinopus (Fum. et Rob.)cannot digest keratin but feed on the lip- ids associatedwith keratinoussubstances such as feathers.However, other possible food sourcesare present,on feathersand pellets.Also, we have often found larvae, nymphs,and adultsof T. putrescentiae,on pellets,with alimentarytracts filled with hyphaeand fungal spores. The two mostimportant pellet invertebratedecomposers are the tineid mothsand Winter 1979 Philips& Dindal--RaptorPellets 105 the trogid beetles.The larvae of the Tineidae, or clothesmoths (fig. 2), eat hair and feathers,and we have foundover 60 of them in a singlepellet. Pelletssometimes ap- pear to be coveredventrally with a centimeteror more of debris.Actually this is a thin layer of soil and moth excretacovering many casesof silk constructedby the larvae.These cases range from 7 to 18 mm long by 2 mm wide, and the larvae feed and pupatewithin them. The open end of the caseis in the pellet and the casere- semblesa small "sock"sticking out of the pellet. In decomposedpellets, the silken network of thesemoth casesmay be all that holdsthe remainingbones together. Fly larvaeof the genusScenopinus prey on tineld larvae in pellets(de Joannis1899). The trogid beetlesalso feed on hair and feathers,and their larvae burrow into the soil beneaththe pellet. The burrowsmay be 30 mm deep by 2 mm wide, and they may be lined with hair for the upper 20 mm. Trogidbeetles also serve as transporta- tion for uropodineand macrochelidmites (Sixl 1971). Many mites attach themselves to insectsfor transportationfrom one site to another.This relationshipis known as phoresy.A mesostigmaticmite Macrochelespenicilliger (B.) hasbeen reported phoret- ic on Trox scaberin owl nestsin England(fig. 3). We have found it on Trox variola- tus in pelletsand rat carrionin central New York. In additionwe have foundnema- todes
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