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Seasonal Abundance and Reproductive Output of the Dung Flies

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Seasonal Abundance and Reproductive Output of the Dung Flies Bulletin of Entomological Research (2008) 98, 397–403 doi:10.1017/S0007485308005713 Ó 2008 Cambridge University Press Printed in the United Kingdom First published online 25 February 2008 Seasonal abundance and reproductive output of the dung flies Neomyia cornicina and N. viridescens (Diptera: Muscidae) R. Wall *, E. Anderson and C.M. Lee School of Biological Sciences, University of Bristol, Woodland Road, Bristol, BS8 1UG, UK Abstract The seasonal abundance and reproductive output of two common, but little studied, dung-breeding flies, Neomyia cornicina and N. viridescens, were examined in artificial cow pats in pastures in southwest England in 2001 and 2004. In 2001, the numbers of both Neomyia species increased slowly over summer to show a sharp seasonal peak in late August and early September. There was no significant effect of mean temperature, mean relative humidity or dung water content on abundance or seasonally de-trended abundance. High levels of aggregation were seen between pats and, when present, greater numbers of N. cornicina emerged than N. viridescens. Neomyia cornicina was present in 13% of 240 artificial standardized pats put out in 2001, at a median of 19 adults per colonized pat; N. viridescens was present in 8% of artificial pats at a median of three adults per colonized pat. In 2004, N. cornicina emerged from 46% of the 94 artificial pats put out at a median of three adults per colonized pat, while N. viridescens emerged from only 12% of pats at a median of one adult per colonized pat. Flies were also collected in 2004, using sticky-traps and hand nets. Again, free-flying N. cornicina appeared to be more abundant in the field than N. viridescens; 162 N. cornicina were caught compared to 44 N. viridescens over the same sampling period. The size of each adult female was recorded and ovarian dissection was used to determine the numbers of eggs matured. Female N. viridescens were significantly larger than the N. cornicina and matured significantly higher numbers of eggs. Gravid N. viridescens matured a mean of 37.1 (+16.9) eggs, whereas gravid N. cornicina matured a mean of 28.8 (+13.2) eggs. The reasons why the larger, more fecund, N. viridescens adults are less abundant in the field or emerging from pats than N. cornicina are unknown. Further work is required to identify the nature and cause of the mortality experienced by the larvae of these species and the ecological differences and functional specialisation which allows co-existence to be maintained. Keywords: cattle-dung, decomposition, Diptera, insect, mortality, Neomyia, oviposition (Accepted 24 October 2007) *Author for correspondence Fax: 00 44 (0)117 925 7374 E-mail: [email protected] Downloaded from https://www.cambridge.org/core. University of Athens, on 23 Sep 2021 at 21:25:59, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0007485308005713 398 R. Wall et al. Introduction species; egg-to-adult development for dung beetles may require several months, whereas the development of larval Dung is a rich resource. It can be highly abundant and flies may require only one to two weeks. During this period, relatively predictable in its occurrence compared with other larvae must also be able to survive high levels of predation patchily distributed resources, such as carrion, particularly within the pat. where large populations of grazing vertebrates are present. The dung-breeding flies Neomyia cornicina (Fabricius) and Herbivore dung is very similar to leaf litter in composition, N. viridescens (Robineau-Desvoidy) (Diptera: Muscidae) are consisting mainly of water and undigested plant material widespread and abundant in cattle pastures. Neomyia along with the products of metabolism, gut epithelial cells cornicina is widely distributed throughout the Holarctic, and a distinct array of micro-organisms (Marsh & Campling, Neotropical and Oriental regions, while N. viridescens is 1970; Greenham, 1972; Stevenson & Dindal, 1987; Aschen- thought to be restricted to the Palearctic. They can be seen born et al., 1989). As a result, the insect community that alighting on fresh droppings and are conspicuous iridescent inhabits this environment is particularly diverse, consisting green-coloured muscids, with a characteristic iridescent of as many as 400 species (Hanski, 1991; Skidmore, 1991). As frons and face. Adult females oviposit batches of eggs in well as high species diversity, the dung environment cavities near the surface of the pat and the egg batches of supports a large number of individuals. It has been several females may be aggregated in the same cavity estimated that a single cow can support an insect community (personal observation). Neomyia spp. are important compo- at least one fifth its own weight purely on the food left in its nents of the dung community but have also been used as faeces (Laurence, 1954) and an individual dung pat may indicator species in studies of the cow-dung invertebrate contain 1000 or more insect inhabitants (Laurence, 1954; community, since they are highly sensitive to the presence of Hanski, 1991). insecticide residues (Gover & Strong, 1995, 1996; Sommer The dung-colonizing insects play a particularly important et al., 2001; Iwasa et al., 2005; Lumaret et al., 2005). However, role in ecosystem function; the timely decay of plant and despite being abundant in the field and widely used in the animal remains is essential to the carbon and nitrogen cycles, laboratory, the basic ecology of these two species is not well soil fertility and the population dynamics of a wide diversity understood; N. viridescens in particular has been little of species at a range of trophic levels. The importance of studied. As a complicating factor, the nomenclature and these invertebrate colonizers in pat decomposition has been taxonomy of these species has been confused; the classic clearly demonstrated by exclusion experiments, where mesh study by Hammer (1941) in Denmark uses the name covers have been used to prevent colonization. Lumaret & Cryptolucillia caesarion. The name Orthellia caesarion has been Kadiri (1995) found that pats from which all insects were particularly widely employed (e.g. Stoffolano & Streams, excluded for one month, took 1.7 to 2.2 times longer to 1971) as has Orthellia cornicina (Wardhaugh & Rodriguez- completely disintegrate than uncovered pats. Holter (1979) Mendez, 1988). There is one report which uses the term also showed that selective exclusion of nocturnal colonizers, Orthellia viridis for flies collected in the south of France (Kirk, particularly Aphodius rufipes L., by covering pats overnight 1992). Whether the various Palearctic studies examined during the first week, resulted in only half of the only N. cornicina or did not differentiate N. cornicina from disappearance found in freely exposed pats. Exclusion of N. viridescens is unknown, but the latter is probable. The aim insects from cow pats for only two days following deposition of the present work, therefore, was to examine the seasonal resulted in a significant increase in the amount of dung that abundance and reproductive output of both N. cornicina and remained after 35 days (Lee & Wall, 2006a). Furthermore, N. viridescens in cow dung in pastures in southwest England several authors have shown that a range of insecticides and to attempt to identify possible ecological differences between anthelmintics administered to livestock and excreted in these two species. faeces may, under some circumstances, kill dung-colonizing insects and retard decomposition (Wall & Strong, 1987; Madsen et al., 1990; Sommer et al., 1992; Strong et al., 1996; Methods and materials Floate, 1998; Floate et al., 2005). Seasonal abundance While tropical dung communities dominated by dung- burying scarabaeid beetles can completely remove a dung The pattern of seasonal abundance was examined in an pat in a few hours (Anderson & Coe, 1974; Hanski, 1991), area of permanent grassland pasture on a farm located dung disappears much more slowly in temperate regions, approximately 20 km southwest of Bristol, UK. The pasture over periods of weeks or months, or even sometimes years used was grazed by a dairy herd of about 250 Holstein- given the occurrence of unfavourable winter or drought Friesian cows and a small number of sheep. The majority of conditions. This slower decomposition allows a complex the cattle were let out to pasture in mid-May and were community of insects to develop in and around the dropping rotated between contiguous fields until mid-October, when during its degradation. Even in temperate regions, however, they were brought back in for the winter. The cattle were not the successional changes in the dung usually make it treated with anthelmintics while maintained in the milking suitable for only one generation of most insects, so in- herd. dividuals must be able to disperse and colonize new pats Batches of ten artificially-constructed cow pats, formed during their life. Hence, selection pressures acting on adults from fresh dung, were placed out each week between the and larvae are very different. Adults must be highly mobile 21st May and 29th October 2001 to allow for insect and able to find new resource (dung) patches in which to lay colonization. When pats were required, fresh dung from eggs, while larvae do not disperse far, but must complete several cows was collected from the milking parlour during their growth before the dung is exhausted or no longer afternoon milking. This was thoroughly mixed to ensure suitable for their development. Clearly, these development uniform constituency and texture and used immediately. rates and selective pressure also vary with the colonising Using a hand-held spring balance, 1.5 kg of fresh dung was Downloaded from https://www.cambridge.org/core. University of Athens, on 23 Sep 2021 at 21:25:59, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
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