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A Comparative Discussion of Trophic Preferences in Dung Beetle Communities

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A Comparative Discussion of Trophic Preferences in Dung Beetle Communities Miscel.lania Zooloqica 19.1 (1996) 13 A comparative discussion of trophic preferences in dung beetle communities F. Martín-Piera & J. M. Lobo Martín-Piera, F. & Lobo, J. M., 1996. A comparative discussion of trophic preferences in dung beetle communities. Misc. Zool., 19.1: 13-3 1. A cornparative discussion on trophic pre ferences in dung beetle cornrnunities. - Ava i la ble information on trophic preferences of dung beetles (Scarabaeoidea) in different biogeographic regions is reviewed. Trophic resource partitioning in a dung beetle (Coleoptera, Scarabaeoidea) community in the ((Parque Nacional de Doñana)), Spain, was also studied, using nine different kinds of wild and domestic vertebrate excrement as trap bait. Undifferentiated attraction to human and herbivore faeces was noted. Human and domestic ungulate faeces was colonized by a richer fauna than that of wild herbivores, which was not the specialized trophic adaptation of any species. Although polyphagy is the most common feeding behaviour, the excrement of carnivores and other omnivores was hardly colo- nized. This pattern differs from that of other biogeographic regions. Negligible importance of the trophic dimension on the structure of these communities may be due to the early presence of man in the Palaearctic Region. Nevertheless, human interference alone cannot have led to an absence of true polyphagy (undifferentiated attraction to al1 kinds of faeces). Further research is suggested, aimed at determining whether observed resource partitioning in dung beetles communities is a consequence of human colonization or is a pre-Neolithic evolutionary event. Key words: Dung beetles, Coleoptera, Scarabaeoidea, Trophic preferences, Resource par- titioning, Ecological-historical causes. (Rebut: 22 1 96; Acceptació condicional.. 29 V 96; Acc. definitiva: 18 VI 96) Ferrnín Martln-Piera & Iorge M. Lobo, Depto. Biodiversidad y Biologla Evolutiva, Museo Nacional de Ciencias Naturales -C.S./.C., dIosé Gutiérrez Abascal 2, 28006 Madrid, España (Spain). This work has been financed by Fauna Ibérica Project, D.G.I.C.Y.T. grant PB89-0081 @ 1996 Museu de Zoologia Martin-Piera & Lobo lntroduction (one of the rnost irnportant wildlife reserves in the Mediterranean area), The physical and chernical composition within the locality of El Rocío, Huelva, of herbivore faeces varies widely with UTM 29SQB2812 (Spain). the species (HANSKI,1987), and even with The sarnpling was taken in a clearing season within the sarne species, as a func- by rneans of 15 pitfall traps set out ran- tion of pasture quality (GREENHAM,1972; domly on 60 x 40 m grid, average dis- MATTHIESSEN,1982; RIDSDILL-SMITH, 1986). Still tance between traps of 10 m, left for greater variation occurs arnong the fae- 48 h (23-25 April, 1992). ces of herbivores, ornnivores and carni- The pitfall-traps were baited with vores. In rnany cases it has been shown approxirnately 1,000 g of fresh excrernent that dung beetles are attracted differ- (see LOBOet al., 1988; VEIGAet al., 1989), ently to different types of faeces (PAUL~AN,except in the case of lynx and fox faeces- 1943). It has been argued that trophic baited traps (250 g were used), dueto the choice could play a role in deterrnining scarcity of resources. the coexistence of species of a dung Nine kinds of excrernent were used, beetle cornrnunity, and their resource frorn vertebrates that still live in the re- partitioning. serve, ranging frorn herbivore (cow, horse, Few studies have been rnade on the deer and fallow deer); predorninantly her- trophic preferences of the Palaearctic bivore (wild boar); predorninantly carni- Region temperate biornes species, and vore (lynx, fox); to ornnivore (rnan, bad- these deal with the differential attraction ger). of the food resources of no more than Two traps were baited with each type four marnrnal species (LANDIN,1961; RAINIO, of faeces, except for those using faeces 1966; DESIERE& THOMÉ,1977; LOBO,1985; frorn lynx, fox and rnan (table 1). CARPANETO& PIATELLA, 1986; SANCHEZ-PINERO There is evidence in favour of a den- & ÁVILA,1991). With the exception of the sity of two traps per site being adequate works by NIBARUTAet al. (1980) and NIBARUTAto have a good representation of the dung (1982), no joint study, using the faeces of beetle cornmunity structure in the Medi- both wild and dornestic rnarnrnals, has terranean regions. Such a density ensures been rnade in this region. that about 53% of local spring species are This paper atternpts to determine captured (confidence intervals at 95% are whether differences correlated with fae- 51.08-55.27%), these species representing ces type exist in an lberian dung beetle 86% of total abundance and 85% of to- cornrnunity. Results are cornpared with tal biornass (Lobo & Lurnaret, in prepara- those previously obtained in the sarne and tion). other biogeographic regions to: i) verify At the sarne time, sarnples were taken the irnportance of food preference results frorn one fresh cow-dung baited (1,000 g) in resource partitioning and ii) provide pitfall trap, set for 48 h in each of the material for a discussion, frorn a histori- following eight park habitats: inter-sand- cal point of view, of the extent to which dune troughs devoid of vegetation ('co- trophic preference may have conditioned rrales'); stationary dunes reforested with the present cornposition of these cornmu- pines; rnarsh; original holrn oak and cork nities. oak wood; rnarsh-holrn oak ecotone; scrub fringes of semi-permanent lagoons ('lu- cio~');strearn bank prirnary and reforested Material and Methods woods (table 2). The sarnples were examined to deter- The study was carried out in or near a mine local fauna diversity, and to obtain grove of holrn oaks on the northern an estirnate of the abundance and habi- edge of 'Parque Nacional de Doñana' tat distribution of each species. Miscel.lania Zoologica 19.1 (1996) 15 Results Whittaker's plots of species abundance data (WHITTAKER,1965) indicate: uneven The number of individuals of each species distribution of abundance in the horse collected with each type of bait is shown dropping community (fig. 3), giving low in table 1. As figure 1 shows, faeces bait eveness and diversity values; a more even type fell into three groups, according to distribution of abundance and greater the number of species and individuals diversity, in cow, fallow deer and human captured: group A, greatest richness and dung beetle communities; fewer species, abundance (cow, human and one of the none dominant, in deer and wild boar, and horse dung); group B, medium to high thus high eveness and diversity values, richness, lower abundance (three species along with an even species abundance of wild herbivore and another horse drop- figures. ping trap); group C, very poor both in Faunal similarity found in the differ- diversity and abundance (carnivores and ent types of faeces was cluster analysed, badger). Total biomass (computations based using percent dissimilarity (PD) and on length-body weight regressions; LOBO, UPGMA, flexible, weighted and unweigh- 1992, 1993) per group was also the great- ted centroid grouping strategies (LUDWIG est for group A (fig. 2). 81 REYNOLDS, 1988). Dung beetle species Number of Species 2 5 - - A ----- \ human \, B cow t \ 20 - / COW t * ,A¿ííGG=. i \ i--___ horseti , fallow deer t i --- -- ------_/-/' t wild boar i 15 - t deer t horse í ; t deer ,* kild boar A /,' lo-,' * N,' L/--.' 5- C \%S--, \ t iynx, o mer I I I I O 100 200 300 400 500 Abundance Fig. 1. Relationship between number of species and abundance for the 15 dung-baited pitfall traps belonging to nine different dung types. Relacíón entre el número de especies y la abundancia para 15 trampas pítfall cebadas con nueve tipos diferentes de excrementos. 16 Martín-Piera & Lobo Faeces H1 H2 C1 Q W1 WZ 81 82 D1 02 F1 F2 L Fo Hu Tot Nb Scarabaeus cicatricosus 110 O 1 O0010 O 1009140.09 100 O O O0000 O O000 10.07 Copris hispanicus O10 2 O O0000 O O000 30.18 Euoniticellus fulvus 30 11O 1 O00011 O00 1180.26 O 11 3 19O0 15 9100 O 1410.39 O01 O O O0000 O O000 10.07 O00 1 O O0000 O O004 50.03 O00 O O O0000 O 1000 10.07 Onthophagus furcatus 22 181 00002 2 3003240.49 10 21 34 48 6 8 O O 12 4 6 5 O O 52 206 0.37 210 o o 00010 o O001 50.22 299 55134 188 5 3 O O 6 74 38 77 O 023711160.42 97 11 58 82 3 7 O O 10 O 23 29 O O 63 3830.42 O 2 915 3 800 631719006880.44 O02 O O O0000 1 O000 3018 Caccobius schreberi 15 3 120 O 2 O O 6 2 11 130 018 910.50 Aphodius baraudi 402 5 3 40042 O 00011350.37 106 5 O 00010 4 5000220.27 O01 O O 10001 1 O000 40.50 100 O O 10000 O O001 30.25 O012 3 O O0000 2 0000170.11 113 3 O 20000 3 200 1160.44 7 O35 8 O O O O 1316 40 O 4 780.30 100 o o 10000 O O000 20.21 O01 o o 00000 1 O000 20.21 6 127 20 1 3 O O 4 2 12 130 0151040.38 O00 O O O0010 O O000 10.07 fhorectes hispanicus O00 O O O0001 O O000 10.07 Typhaeus momus 2 2 2 4 O 1 O000 O 21 113280.21 Total abundance 461 103 365 445 27 64 O O 61 104 174 201 1 1 470 2477 Total no. species 19 14 21 20 10 16 O O 16 13 19 17 1 1 21 35 Total biomass 4.6 1.6 2 9 4.3 0.5 0.6 O O 0.8 0.9 1.3 2.2 0.1 0.1 8.4 28.3 Miscel.lania Zooloqica 19.1 (1996) 17 Table 1.
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