Mineral grain availability and pupal-case building by lotic caddisflies: Effects on case architecture, stability and building expenses
Résumé
The retreat-making larvae of many lotic caddisflies build entirely new pupal cases with fine gravel and sand that they collect in the neighbourhood of the building place to fix it with silk to cobbles in swift flow (where finer sediments are generally rare). Previous field observations on Hydropsyche siltalai pupal cases illustrate that natural local resource limitations of the preferred grain fraction (2.5-3.15 mm) pro- duced chained effects across other grain fractions, as the alternative use of more grains in the 1.6-2 mm fraction (an unlimited resource) induced an increased use of more grains in the 0.315-0.5 mm fraction (another unlimited resource). To examine the implications of these observations for H. siltalai, we used (1) mesocosms to created minor deviations in the availability of the natural grain size composition of the building material of pupal cases at otherwise carefully replicated natural stream habitat conditions and (2) recently developed technologies to assess many case characteristics so far ignored in studies of caddisfly cases. When the preferred coarser grains (2.5-3.15 mm) were unavailable, more grains with intermediate size (1.25-2.0 mm) were used (and not other, still available coarse grains) and fewer lar- vae built cases in groups, thereby not only loosing the benefits (lower costs for grain transport and silk) but also avoiding potential disadvantages associated with grouped cases (more aggressive encounters with conspecifics for rare building material, less flow exposure and thus reduced water renewal in the pupal chamber). Unavailability of 2.5-3.15-mm and 0.315-0.5-mm grains caused a reduction of larvae building in groups, more use of grains with intermediate size, changes of several other grain character- istics (e.g. number, circularity) and considerable investment into silk to maintain the case resistance. Finally, grain availability deviating most from that observed in nature (no grains of 2.5-3.15 mm and 1.6-2.0 mm) caused dramatic responses, as mortality increased so that fewer pupal cases were built, using typically more coarse grains so that many cases had an elevated resistance against crushing forces; in addition, many males had a retarded development, whereas female development was unaffected. Thus, the response of H. siltalai to any of the three types of grain limitations differed, illustrating an immense diversity to respond to grain-size shortage.