1. Shredders and Gougers Consume CPOM (> 1 mm mesh) - leaves, etc., and thus depend heavily on seasonal inputs of leaves to the stream Mouthparts usually “chewing” (e.g., many Trichoptera) Leaves entering the stream are first “conditioned” by bacteria and fungi, reducing their toughness and creating more digestible proteins and carbohydrates for the invertebrates Most shredders can’t digest cellulose, but Tipula harbors endosymbiotic bacteria capable of cellulose digestion Gougers consume wood, and are often found inside submersed snags (e.g., the chironomid Brillia); these are unusually slow growing taxa Their feces form an important constituent of FPOM Fam. Taeniopterygidae; O. Plecoptera Fam. Tipulidae; O. Diptera

2. Filter Feeders Consume suspended particles (FPOM), including phytoplankton, bacteria and seston (temporarily suspended particles) Frequently found below lakes and reservoirs Includes many Trichoptera, e.g., Hydropsychidae Blackfly (Simuliidae) larvae have cephalic fans which are held at the edge of the boundary layer and trap particles, which are removed by labral bristles and transferred to the mouth Flocculation of DOM by bacteria converts it to a form consumable by filter feeders

3. Filterers Fam. Simuliidae; O. Diptera Fam. Unionidae Fam. Philopotamidae; O. Trichoptera Fam. Hydropsychidae; O. Trichoptera

4. Deposit Feeders Also feed on FPOM, but by gathering it from the sediments; the FPOM includes bacteria, algae, detrital particles, of widely varying food quality Typically brushlike mouthparts Includes many mayflies, midges, crustaceans Fam. Chironomidae; O. Diptera (many are deposit feeders)

5. Scrapers/Grazers Specialists on periphyton on solid surfaces Mouthparts designed to shear attached algae from the substratum (e.g., scythe-like mandibles of the caddisfly Glossosoma; the radula of snails) Fam. Heptageniidae; O. EphemeropteraFam. Psephenidae; O. Coleoptera

6. Scavengers Many macrocrustacea (amphipods, isopods, crayfish) opportunistically consume animal, algal and plant material O. AmphipodaO. Isopoda

7. Invertebrate Predators This group is highly diverse, both in microhabitat and food specialization – many surface-feeding Hemiptera – sit-and-wait benthic predators (e.g., dragonflies) – fluid specialists (e.g., leeches), etc. etc. Some taxa are predaceous for only part of the life cycle (e.g., later instar Tanypodine chironomids, larval hydrophilid beetles)

8. Predators fam. Nepidae; O. Hemiptera fam. Gerridae; O. Hemiptera Fam Perlidae; O. Plecoptera) fam. Tabanidae; O. Diptera Fam. Hydrophilidae; O. Coleoptera

9. Predator Behavior Hunting Behavior – 1. stalking/active pursuit (e.g., perlid stoneflies) – 2. ambush (e.g., hemipteran family Nepidae) Prey detection Mechanisms – 1. tactile (e.g., the stream damselfly Calopteryx) – 2. Visual (e.g., most other damselflies) – 3. Chemical (perlid stoneflies search in an upstream, following prey chemical trails) Calopteryx visual damselfly

10. Prey Defenses Primary (operate regardless of predator’s proximity) – a. refuges (e.g., many Trichoptera, chironomids) – b. crypsis Secondary (behavioral responses) – a. Thanatosis (feigning death) (e.g., some Coenagrionid damselflies) – b. Secondary Compounds (e.g., many Coleoptera and Hemiptera) – c. Group Defense (e.g., Gyrinids may confuse predators) – d. Active (e.g., strigulating in the beetle Tropisternus; use of the scorpion posture in ephemerellid mayflies) Ephemerellid mayflies produce “scorpion posture” when threatened by stonefly predators

11. Macrophyte Consumers Leaf and stem feeders Many Lepidoptera, some chironomids, some beetles, some Trichoptera Fam. Chrysomelidae; O. Coleoptera Fam. Pyralidae; O. Lepidoptera

12. Fish predation Evolutionary effects on invertebrate behavior/morphology – (e.g., Trichopteran cases provide protection against fishpredators) Effects of fish on stream inverts much harder to detect than in lakes, owing to the effects of drift The Trophic Cascade – The basic idea is that the food web can be simplified to a food chain in some streams. An increase in size in a top trophic level may then reduce the size of the level below, increase the size of the level below that, etc.

13. The River Continuum Concept As stream order increases: – a. the influence of the riparian canopy diminishes as a light interceptor – b. production by periphyton and macrophytes increases – c. inputs of leaf litter decline – d. shredders are largely replaced by FPOM feeders – e. P/R increases, but then may decrease once more in very large rivers – f. plankton communities become sustainable in river systems Vannote et al. (1980)

14. Aquatic Insect Respiration Tracheal system – Usually consists of a spiracle leading to a trachea, then branching tracheoles Atmospheric oxygen obtained by – visiting the surface – Transporting a bubble underwater (“physical gill”). – e.g., most Hemiptera and Coleoptera Dissolved oxygen extracted from water – e.g., into gills or across the integument – e.g., mayflies, stoneflies, odonates, dipterans – Oxygen ultimately reaches “closed tracheal system” (no spiracles)

15. The Physical Gill – The bubble is brought down from the surface – As oxygen is drawn from the bubble through the spiracles into the insect, oxygen from the water diffuses into the bubble (greatly prolongs its use) Hemipteran Coleopteran

16. Oxygen from Plant Stems The beetle Donacia and its relatives tap into the roots of water lilies and other submersed plants

17. Hemoglobin as an adaptation to low- oxygen environments Enhanced ability to take up oxygen from oxygen-poor environments Midge (Chironomus) larva

18. Aquatic Insects Typically the immatures are found in water; the adults may either aquatic or terrestrial. The change from water to land during the life cycle may require considerable ontogenetic modification of body form. Two general and distinct types of life histories: Hemimetabolous: (Odonata, Ephemeroptera, Plecoptera, Hemiptera) egg  larva or nymph  adult Holometabolous: (Megaloptera, Neuroptera, Trichoptera, Coleoptera, Diptera, Lepidoptera) egg  larva  pupa  adult – The pupal stage may either be motile or non-motile, and is the principal means for reorganizing the body structure

19. Life History Stages A.Holometabolous (e.g., Trichoptera) B. Hemimetabolous (e.g., Plecoptera)

20. Mayflies are hemimetabolous Hexagenia mayflies typically spend a year as larvae, emerge as a pre-reproductive adult (“dun”), then molt again to reproduce (“spinner”). There is no pupal stage.

21. Holometabolous Life Cycle Whirligig beetles (fam. Gyrinidae) have morphologically very distinct larval, pupal and adult stages. Pupa

22. More primitive insect orders have more larval instars OrderTypical No. larval Instars Ephemeroptera15-25 Odonata10-12 Plecoptera12-22 Hemiptera5 Megaloptera10-11 Coleoptera3 Trichoptera5 Diptera4-7

23. Time to complete the life cycle univoltine – Species which require one year to complete the life cycle bivoltine – two generations per year Multivoltine – > 2 generations/yr Larger, or higher latitude, taxa may require more than one year to complete the life cycle, and two or more distinct size classes may thus co-occur in the same location. Larval hellgrammites (O. Megaloptera) may require 5-7 years before emerging to become adults Adult Megalopteran Hemimetabolous or holometabolous?

24. Overview of the Orders of Aquatic Insects covered in lab Ephemeroptera: Mayflies Plecoptera: Stoneflies Odonata: Dragonflies and Damselflies Trichoptera: Caddisflies Hemiptera: True Bugs Coleoptera: Beetles Megaloptera: Alderflies and Dobsonflies Diptera: True Flies A few examples are shown in the slides that follow; the complete list of taxa, with slides, is provided in the “Macroinvertebrates for Practicum” file posted on Blackboard