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The Ecology of Riffle Beetles (Coleoptera: Elmidae)

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The Ecology of Riffle Beetles (Coleoptera: Elmidae) 189 Article The ecology of riffle beetles (Coleoptera: Elmidae) J. Malcolm Elliott Freshwater Biological Association, The Ferry Landing, Far Sawrey, Ambleside, Cumbria LA22 0LP, UK. Email: [email protected] Received 12 May 2008; accepted 21 July 2008; published 5 November 2008 Abstract Riffle beetles in the family Elmidae are frequent members of the invertebrate community of running water. Over 80 species have been recorded in North America and 46 in Europe; this number decreases in the western and northern fringes of Europe with only 12 species in Britain, many rare, four in Ireland and three in Norway. The present review describes their habitat, food and predators, their life cycles, their dispersal, and human threats to their survival. All elmid species have aquatic larvae with five to eight instars, depending on the genus. Adults of a few species are terrestrial, but most are aquatic with plastron respiration. Most elmid species occur in well-aerated streams and rivers, but some also occur on wave-washed lake shores. A few species live in more unusual habitats, such as thermal pools and hot springs, subterranean habitats, and decaying wood. Little is known about the food of adults or larvae, but they appear to be collector-gatherers and scrapers that feed chiefly on algae and detritus. Adults and larvae are rarely taken by invertebrate predators, but are eaten by fish, especially salmonids. However, their proportion in the diet is always lower than that in the benthos, indicating low availability to the fish. To obtain a quantitative description of the life cycle, information is required on the number of larvae in each instar, the timing of oviposition, the number of eggs laid, the timing of pupation, and the number of adults. Few studies meet these criteria. However, three excellent studies from North America and a detailed study of the four commonest British species illustrate the variation in the life cycles of different species. Life tables, identifying critical periods for survival, are provided for the British species, these being the only such tables currently available for riffle beetles. The ability to disperse by flight varies among species and among individuals of the same species. Adults and larvae disperse downstream in the invertebrate drift, especially at night. In one study, drift densities of different life stages were related positively to their monthly losses in the benthos, but not to their benthic densities. Less information is available on their upstream–downstream movements on the substratum. There are ontogenetic shifts in diel drift periodicity and dispersal, and both relate to seasonal changes in drift density and critical periods for survival in the life cycle. Human threats to the survival of elmids include reduced oxygen concentrations, elevation of water temperature, extremes of flow, especially spates, and pollution, especially by soaps and detergents. Therefore, riffle beetles provide excellent indicators of water quality and perhaps also climate change. Keywords: Dispersal; drifting; feeding; habitat; life histories. DOI: 10.1608/FRJ-1.2.4 Freshwater Reviews (2008) 1, pp. 189-203 © Freshwater Biological Association 2008 190 Elliott, J.M. Introduction et al., 1996). Therefore, there appears to be a paucity of species in the western and northern fringes of Europe. Beetles in the order Coleoptera constitute the largest order All elmid species have aquatic larvae with anal gills and of insects with over 250 000 species worldwide. Complete five to eight larval instars, depending on the genus. Adults of or partial adaptation to life in fresh water has occurred a few species are terrestrial but most are aquatic with plastron independently several times during beetle evolution. There are respiration. Pupation always occurs out of the water. For about 5000 aquatic species, ranking them as one of the major most species, the final-instar larvae crawl out of the water groups of freshwater arthropods. Riffle beetles are in the small and pupate in the soil or under stones near the water. A few family Elmidae with over 80 species in North America (White species in streams with fluctuations in water level simply & Brigham, 1996) and 46 species in Europe (Berthélemy & wait in shallow water near the stream margin until the water Olmi, 1978). Early literature referred to the family as Helmidae. level drops and then pupate in situ (White, 1978; Seagle, In his pioneering re-organisation of the Coleoptera, Crowson 1980). The plastron of the adults is a remarkable structure (1956) placed them in the suborder Polyphaga, superfamily that acts as a physical gill. When adults first enter the water, Dryopoidea, family Elmidae. Steffan (1958, 1961, 1963) argued most of their body is covered by a thin layer of air which strongly for the family name Elminthidae and this name was functions as a gill, being in contact with air spaces under used in the key to British species (Holland, 1972). Berthélemy the elytra (hardened fore-wings). This plastron is facilitated (1979) reviewed the arguments for calling the family Elmidae by a dense coating of hydrofuge, hair-like setae that trap a or Elminthidae, and summarised the case for using the term film of air on the ventral surface of the adult. As oxygen is Elmidae. The latter term is usually used in the North American removed from this reservoir for respiration, it is replaced literature, and therefore it is logical that the same term is used by diffusion of dissolved oxygen from the surrounding on both sides of the Atlantic. water (Brocher, 1912; Harpster, 1944; Beier, 1948; Thorpe Eleven species in eight genera were recorded in Britain & Crisp, 1949; Thorpe, 1950; Hinton, 1976). Adult riffle in Limnofauna Europaea (Berthélemy & Olmi, 1978) and in beetles maintain a plastron indefinitely under considerable the British key (Holland, 1972). However, some of these water pressure and can thus remain submerged in species are rare. For example, Fowler (1889) predicted well-oxygenated water without having to re-surface. that Stenelmis canaliculata (Gyllenhål) would probably be The early European literature on the ecology of riffle recorded as British in the future, but it was many years beetles was largely qualitative and is summarised in the later that this species was found for the first time on the monograph by Berthélemy (1966). The North American shore of Windermere, the largest natural lake in England literature was included in the general review of the biology (Claridge & Staddon, 1961). Similarly, Oulimnius rivularis of the superfamily Dryopoidea, including the Elmidae, by (Rosenhauer), last found in the River Wey in 1870, was Brown (1987). The present review of the ecology of riffle rediscovered in the Sixteen-Foot Drain in Cambridgeshire beetles describes their habitat, food and predators, their life (Foster & Bratton, 1986). A twelfth species, Oulimnius cycles, their dispersal, and human threats to their survival. major (Rey), that occurs in continental Europe, has also Emphasis is placed on quantitative aspects of their been added to the British list (Parry, 1980). The four ecology, using examples from Europe and North America. commonest species in Britain are Elmis aenea (Müller), Esolus parallelepipedus (Müller), Limnius volckmari (Panzer), Habitat, food and predators and Oulimnius tuberculatus (Müller) (see distribution maps in Holland, 1972). These are also the only species of elmid Adults are small, brown or black, crawling beetles that beetles recorded in Ireland (Berthélemy & Olmi, 1978; range in size from 1 mm to 8 mm. Although essentially a Ashe et al., 1998), and only three of them (all except Esolus taxonomic account, Nilsson (1996) includes some ecological parallelepipedus) have been recorded in Norway (Ødegaard notes on riffle beetles and some beautiful illustrations of the © Freshwater Biological Association 2008 DOI: 10.1608/FRJ-1.2.4 Ecology of riffle beetles 191 northern European species. Excellent illustrations of the substances liberated into the decaying wood by microbial British species are provided in the monograph by Holland activity. They produce large quantities of woody faeces, (1972), who also provides body lengths, ranging from 1.35 indicating that only a small amount of the ingested food mm to 1.50 mm for the smallest, Esolus parallelepipedus, to is utilised. Xylophagous elmid larvae, especially Notriolus 3.75 mm to 4.75 mm for the largest, Stenelmis canaliculata. galstonius (Carter), N. quadriplagiatus (Carter), and Simsonia Adults live for several months in the wild but have been tasmanica (Blackburn), were the principal (keystone) kept alive in the laboratory for over three years (Brown, macroinvertebrate modifiers of wood in south-eastern 1973). They are often encrusted with so much inorganic Australian streams; their gauging of channels in the wood material that they have to be cleaned before identification facilitated colonisation by other macroinvertebrates and is possible. The larvae are elongate and hard-bodied with wood-decaying microorganisms and fungi (McKie & a shape in cross-section that is either triangular (e.g. genera Cranston, 1998). Density, biomass and production of six Elmis, Esolus and Oulimnius) or circular to semi-circular elmid species, including four Stenelmis spp., were much (e.g. genera Limnius, Riolus and Stenelmis). Larvae with higher on a submerged woody habitat than in an adjacent a triangular cross-section are usually lanceolate in shape channel bed in a river in Georgia, USA (Benke, 2002), but and are found under stones or among moss and the roots this study did not support a keystone role as suggested of plants, while those with a rounder cross-section often for xylophagous species by McKie & Cranston (1998). burrow in gravel or wood. The majority of elmid species are found in well-aerated As their common name implies, most elmids are found streams and rivers, but a few species in the genera Stenelmis in running water. However, a few species occur in more and Oulimnius also occur on wave-washed lake shores.
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