Ontogenetic shifts in drift periodicity and benthic dispersal in elmid beetles

1. There is a paucity of information on ontogenetic changes in the dispersal of benthic invertebrates, which is an important aspect of their ecology. This study quantifies ontogenetic changes in diel periodicity in drift, and in upstream–downstream dispersal on the substratum for Elmis aenea, Oulimnius tuberculatus, Esolus parallelepipedus and Limnius volkmari (Coleoptera: Elmidae). Three drift nets were emptied every 3 h over 24 h in each month (October 1965–December 1968) at two contrasting sites: one in a deep section with abundant macrophytes, the other in a shallow stony riffle. Comparisons of periodicity between life‐stages of the same species were limited to months when numbers in the drift were highest. Dispersal was evaluated in six experimental stream channels, placed above the stream, with initial numbers of each life‐stage varying from 20 to 80. 2. Drift numbers were always highest at night with few or no animals in the day samples. Ontogenetic shifts in diel periodicity were similar for all four species. Drift catches were similar throughout the night for the early and intermediate larval instars and for mature adults, but were highest in the early hours of the night with a gradual decline thereafter for later larval instars and immature adults. These patterns were unaffected by a severe spate, even though drift numbers increased considerably. 3. Dispersal was density‐independent; the number of dispersing animals was a constant proportion of the initial number for each life‐stage. The relationship between dispersal distance and the number of animals travelling that distance was well described by an inverse power function. Median and maximum distances (m day^?1) were estimated for each life‐stage. 4. Ontogenetic shifts in dispersal in the stream channels matched those shown in diel drift periodicity. For all four species, the later larval instars and immature adults showed little movement in either direction, whereas early and intermediate larval instars and mature adults dispersed predominantly upstream, adults travelling further than any other life‐stage. 5. Ontogenetic shifts in diel drift periodicity and dispersal were related to seasonal changes in drift density and critical periods in the life cycle. Such shifts have not been quantified in other stream invertebrates, but should be considered when evaluating the role of dispersal in their population dynamics and their colonization ability.     

Critical periods in the life cycle and the effects of a severe spate vary markedly between four species of elmid beetles in a small stream

1. The chief objectives were: (i) to describe quantitatively the life cycles of four species of Elmidae, Elmis aenea, Esolus parallelepipedus, Oulimnius tuberculatus and Limnius volkmari; (ii) to use life tables to identify critical periods for survival in the life cycle of each species; (iii) to evaluate the immediate and longer‐term effects of a severe spate on densities of the four species. Monthly samples were taken over 63 months at two contrasting sites in a small stream: one in a deep section with macrophytes abundant, and the other in a shallow stony section. 2. There were five larval instars for O. tuberculatus, seven for L. volkmari and six for the other two species. The life cycle of each species took 1 year from egg hatching (chiefly in June for E. aenea and O. tuberculatus, and July for the other species) to pupation in the stream bank and a further year before the adults in the stream matured and laid their eggs. Mature adults were present in most months, but were rare or absent in January and February and attained maximum densities in April for O. tuberculatus and May for the other species. 3. Laboratory experiments provided data on egg hatching and pupation periods and the number of eggs laid per female. Life tables compared maximum numbers per square metre for key life‐stages. Within each species, mortality rates between adjacent life‐stages were fairly constant among six cohorts and between sites, in spite of large differences in numbers. The only exception for all species was the high adult, but not larval, mortality during a severe spate. 4. Standardised life tables, starting with 1000 eggs, identified key life‐stages with the highest mortality, namely the early life‐stages for E. aenea (36% mortality), start of the overwintering period to pupation for O. tuberculatus (41%) and L. volkmari (51%), start of pupation to the maximum number of immature adults for E. parallelepipedus (41%) and between the maximum numbers of immature and mature adults for O. tuberculatus (41%). Therefore, critical periods for survival in the life cycle differed between species, presumably because of their different ecological requirements. Similarly, the effects of the spate on adult mortality, and hence egg production, varied between species, being most severe and long‐term for E. aenea and O. tuberculatus, less severe for E. parallelepipedus and least severe with a rapid recovery for L. volkmari. Possible reasons for these discrepancies are discussed, but more data are required on the food and microhabitat requirements of the elmids before satisfactory explanations can be found.     

Ontogenetic changes in the drifting of four species of elmid beetles elucidate the complexity of drift-benthos relationships in a small stream in Northwest England

1. This study aimed to quantify ontogenetic changes in the drifting of Elmis aenea, Oulimnius tuberculatus, Esolus parallelepipedus and Limnius volkmari (Coleoptera: Elmidae), and to relate their drift to benthic density. Monthly samples were taken over 39 months, using three surface nets at each of two contrasting sites in a small stream: one in a deep section with abundant macrophytes, and the other in a shallow stony section. 2. Most larvae and adults were taken in the drift at night with little variation between catches in the three nets at each site. Day catches were very low, often zero. No significant relationships could be established between mean numbers in the drift catches and benthic densities. 3. When night catches were converted to drift densities (number caught per 100 m³ of water sampled), the latter were positively related to monthly losses in the benthos, but not to benthic densities. A linear regression described the relationship, and equations for the different life‐stages within each species were not significantly different from the equation for all life‐stages combined. However, drift losses were only about 0.07% of total losses in the benthos. A severe spate in October 1967 increased the number of larvae and adults in the drift, but not drift densities, except for immature adults of E. aenea, O. tuberculatus and E. parallelepipedus. 4. Key life‐stages with the highest drift density were the earliest life‐stage soon after egg hatching for E. aenea, the start of the larval overwintering period for O. tuberculatus and L. volkmari, and mature adults during the mating season for all three species. Drift density for E. parallelepipedus was too low to identify a key life‐stage. These key life‐stages corresponded with critical periods for survival in the life cycle, as identified in an earlier study in the same stream. Mortality was high during these critical periods, hence the strong relationship between drift density and benthic losses. The latter relationship was very consistent for different life‐stages within each species, and partially supported the rarely‐tested hypothesis that drift represents surplus production in the benthos.