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Resource Partitioning in Coprophagous Beetles from Sheep Dung: Phenology and Microhabitat Preferences

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Resource Partitioning in Coprophagous Beetles from Sheep Dung: Phenology and Microhabitat Preferences Zool. Jb. Syst. 121 (1994), 171-192 Gustav Fischer Verlag Jena Biologisches Institut I (Zoologie) der Albert-Ludwigs-Universitat Freiburg, Freiburg, Deutschland Resource Partitioning in Coprophagous Beetles from Sheep Dung: Phenology and Microhabitat Preferences R WASSME S THOMA d an G SOWI R PETE With 6 Figures Abstract On a pasture in SW-Germany coprophagous beetles (Scarabaeidae, Hydrophilidae: Sphaeridiinae and Staphylinidae: Oxytelinae) from sheep droppings were studied conside- ring the following niche dimensions: season, size, type and water content of the dropping. 1. Phenological differences were found especially within endocoprid Scarabaeidae. Dominant Aphodius-species. were clearly separated from each other by their phenology, while nearly all Hydrophilidae from the genus Cercyon and paracoprid Scarabaeidae were most abundant in May. 2. Sheep produce different types of droppings, either compact lumps or small pellets. The latter are mostly deposited in groups. Since these pellets dry out quickly, but rehydrate during rainfall, a high variability of humidity conditions makes pellets unattractive to many dung bettles. 3. Pairwise niche overlap indices in coprophagous beetles from sheep droppings were calculated regarding two niche dimensions: dropping size and water content. Cluster trees showed a clear distinction between species according to these microhabitat factors. Especially Hydrophilidae tend to avoid small droppings which might be due to unstable humidity conditions there. Mean intrageneric overlap within genera with more than one species was equal to or even higher than mean intergeneric overlap rates. 1. Introduction a y b r diffe d shoul e biotop e on n withi s specie g cooccurin f o n utilizatio e Resourc limiting similarity according to the competitive exclusion principle (GAUSE 1934). f o n limitatio a d an s specie f o r numbe d limite a o t d lea d shoul y similarit g limitin s Thi y "Wh n questio e th s stand s thi o t y Contradictor . community e on n withi y diversit are there so many kinds of animals?" (HUTCHINSON 1959). Especially in invertebra- n exclusio e competitiv s specie f o s number h hig h wit s insect e lik a tax e t seemlese b s o evidentst . Thus validite th , GAUSE'f yo S principl thereupod ean e nth existence of competition has been discussed quite controversially (DEN BOER 1980, 1986; ABRAMS et al. 1986). Many studies focussed on additional explanations of the coexistence of competing similar species like high intraspecific interference competition (VANCE 1985), intraspecific aggregation (ATKINSON and SHORROCKS 1981), the role of predation on stable coexistence (CASWELL 1978, CONNELL 1971), 12* Zool. Jb. Syst. 121 (1994) 2 171 n i s diversitie h Hig . 1983) N SILVERTOW d an Y HARVE y b w (revie s event e chanc d an case of communities of insects and other taxa of small invertebrates with high numbers of species are said to be due to spatio-temporal heterogeneity of ephemeral t permi y ma s condition e unstabl e sinc , environment") y ("patch s microhabitat ecologically similar species to coexist (HORN and MACARTHUR 1972, HANSKI 1983, SHORROCKS 1990). Their theoretical models showed, that migration between and e quit n eve f o e coexistenc e possibl a o t s lead s patche r insula n withi n extinctio l loca e ar s resource h whic , detect o t s i y ecolog y communit f o l goa e On . species r simila partitioned by coexisting species, and how far these species are separated in niche space. This may help us to explain the extreme species richness of insect communities. y man y b d studie e wer s herbivore e larg m fro s excrement s decade t las e th e Sinc y patch a h suc n withi s specie f o s interaction t detec o t r orde n i s ecologist environment (ADAM 1986, BREYMEYER 1974, HANSKI and KOSKELA 1979, HOLTER 1982, LANDIN 1961, LUMARET and KIRK 1987; MADLE 1934, MOHR 1943, NEALIS 1977, OTRONEN and HANSKI 1983, RAINIO 1966, STEVENSON and DINDAL 1985). n compositio e th , completely d extracte e b n ca s insect l al s faece n chose a m fro e Sinc of this community can be documented accurately, which makes dung pats well suited for studies of patterns of resource partitioning. Dung is used as both s a g actin s specie y fl d an e beetl f o y variet a y b e resourc d foo d an t microhabita coprophagous decomposers or carnivorous predators. While most previous papers n o s focusse y stud s thi , strictu" u "sens s beetle g dun s a e Scarabaeida o t d confine e wer : Scarabaeidae : groups c taxonomi g followin e th m fro s beetle s coprophagou Geotrupinae, Aphodiinae and Onthophaginae; Hydrophilidae: Sphaeridiinae; d an e larva s a g dun n o d fee e Scarabaeida s Coprophagou . Oxytelinae : Staphylinidae e ar e Oxytelina . adults s a y onl s coprophagou e ar e Sphaeridiina e whil , imagines s a not strongly restricted to dung and use other decaying substrates, too. Within the - ("endoco t pa g dun e th n i y directl d lai e ar e Aphodiina m fro s egg , Scarabaeidae prid"), while members of Geotrupinae and Onthophaginae build brood chambers n desiccatio d avoi o t ) ("paracoprid" g dun h wit m the e provid d an t pa e th h beneat . 1930) R BURMEISTE ( Dung pats can be considered as discrete patches with a certain combination of a e influenc s character e thes f I . content r wate r o e siz , age e lik s character species-specific patch choice by dung beetles, species compositions between patches g dun w co h wit e don n bee s ha k wor l ecologica t mos e Whil . different e b d shoul (review by HANSKI 1990), faeces from sheep were considered only in a few studies d an T LUMARE , 1986 M ADA , 1974 Z OLECHOWIC , 1974 R BREYMEYE , 1961 N (LANDI KIRK 1987). One quite obvious feature of sheep dung is that sheep produce two different types of droppings: 1. compact lumps, which vary widely in size, or 2. small pellets, which are deposited in groups or isolated as single pellets. In contrast to cow dung, which develops a compact crust on its surface, sheep droppings are able to r wate , droppings l smal n i y especiall , Thus . rainfall g durin r o w de y b e rehydrat , 1961 N (LANDI e ag s it h wit d correlate e b r neithe n ca d an y widel s varie t conten g droppin d an t conten r wate , Therefore . size s it h wit r no ) 1987 K KIR d an T LUMARE size can be considered as important microhabitat factors for patch choice, while 172 Zool. Jb. Syst. 121 (1994) 2 f o l contro e th n i r paramete d goo a e b o t m see t no s doe ) age = ( e tim n expositio y extremel s varie e siz h patc s A . droppings p shee n i s communitie t insec f o n successio a n I . separation e nich n i e rol t importan n a y pla t migh t i , droppings p shee n i f o s bait g dun h wit s trap l pitfal d use ) (1984 N HOWDE d an K PEC t fores l tropica e nich a s a d regarde e b n ca e siz g dun t fac n i t tha , showed d an e siz t differen . beetles g dun r fo n dimensio d focusse s beetle g feedin g dun f o e structur y communit e th n o s studie s Previou s preference t macrohabita , 1987) K KIR d an T (LUMARE y phenolog n o y mostl (LANDIN 1961, NEALIS 1977), succession (HANSKI 1980a), and brood care strategies (BRUSSAARD 1983, BURMEISTER 1930, DOUBE 1990, HALFFTER and MATTHEWS 1966, o t s i y ecolog e beetl g dun l experimenta n i d metho n commo A . 1983) R KLEMPERE l equa f o s pat g dun w co d standardize g exposin y b e siz h patc f o e influenc e th e exclud weight in the field (HANSKI and KOSKELA 1977, HOLTER 1982, OTRONEN and HANSKI 1983, RAINIO 1966). This paper examines the extent to which coprophagous beetles inhabiting sheep dung are separated from each other in four niche dimensions: . content r wate d an , size g droppin , dropping f o e typ , season 2. Material and Methods 2.1. Field work and getting raw data r nea a are e larg a h 5 6 a n i l leve a se e abov m 0 40 t a s site y stud t adjacen o tw d selecte e W g durin t leas t a e pastur p shee a s a y extensivel d use s wa h whic ) (Southwest-Germany g Freibur the last two decades. For a detailed description of this locality see WASSMER and SOWIG ) May d an l (Apri g sprin n i y onl t presen e wer p shee , 1990 n i d perio y stud r ou g Durin . (1993) and autumn (September - November). Once in these months we randomly collected sheep droppings of different size and water content between 9:00 and 12:00 GMT. We yield a total of 698 sheep droppings with a dry weight of 10.67 kg as a whole.
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