D. Benthic Invertebrates – most adaptations, wide phylogenetic diversity

        1. Morphology
            a. Flattened and streamlined - decrease resistance to flow; but is also an adaptation for living under rocks
            b. Suckers and hooks - allows to grasp rocks; hooks (tarsal claws)
            c. Tubes -- Chironomid larvae, sticky silk, attached to rock
            d. Ballast - help them to remain on bottom

        4. Behavioral responses to stream flow
            a. Current avoidance
            b. Drift (both a noun and a verb)
                    mostly at night
            c. There is also some movement upstream, but this is relatively slow
            d. Why drift?
                1) Proximate cause (cues) - light

                2) Ultimate or adaptive cause
                    a) None - Accidental -

• but this is counter to periodicity;
• there is some low level continuous accidental displacement

            e. Compensation for drift -- why aren't all the insects in the ocean?  Why are there any left in the streams?
                (1) Colonization cycle – upstream flight

                            Adults fly upstream; not tested until recently
                             -Arctic stream insects were labeled with ¹⁵NH₄⁺ by introducing it into the stream
                             -Collected adults upstream later in season when emerging
                             -Any insects above the ¹⁵NH₄⁺ emergence point with ¹⁵N had to have come from downstream
                            Average distance of flight upstream ~2 km
                            Average distance of downstream drift ~2 km
                            Therefore upstream flight of adults can compensate for drift

                    (2) Excess production hypothesis

                            Even if many drift, there are still a lot left
                            Better success of eggs deposited upstream – less competition
                            Difficult to assess because it is difficult to measure upstream production and combine these
                                    measurements with downstream movement

II. Stream ecology
    A. Feeding – functional group concept – ‘guilds’

        1. shredders - biters and chewers; take large food and produce small foods;
                                herbivorous or detritivorous (leaves and microfauna)
        2. scrapers - feed on aufwuchs (on substrates); specialized mouth parts to scrape material on substrates
        3. collectors - spin nets or use setae to collect organic matter; feed on fine particulate organic matter;
                                filter with nets, hairs; cephalic fans (black flies)
        4. predators - carnivorous; swallow prey whole or bite pieces or suck out contents
 

    B. detrital material - much of the food web in a stream is detrital; this detritus is broken up into categories by size
        1. CPOM - coarse particulate organic matter; >1 mm; leaves, wood, litter
        2. FPOM - fine particulate organic matter; 50 mm-1mm
        3. DOM - <~0.45 mm

    C. How do the guilds fit together?

    D. River continuum concept (Vannote et al. 1980)
        - Streams change as you go from the headwaters to the high order rivers
            1. predictable physical features and gradients

• stream velocity, suspended particulate load, width and depth of stream

• structure of community and function;
• P/R (production/respiration)

                -Why P<R at 1st order? --

• lots CPOM input to stream and it is being decomposed
• low production due to light limitation (stream is shaded)

• less CPOM; more collectors and scrapers (fewer shredders);
• more light leads to more production

• more FPOM, no scrapers because fewer algae;
• light-limited due to suspended sediments

        4. criticisms -
            i. oversimplified;
            ii. mostly holds for pristine rivers
            iii. relates only to macroinvertebrates
            iv. if low order streams are devoid of forest then they aren't shaded and don't have high CPOM loads

    E. Resource spiraling concept (Newbold et al. 1982)
        1. closed system (no inputs or outputs; have rate and pathways)
        2. open system (inputs, outputs; rates, pathways, residence time)
        3. open system with spiraling (downstream transport)
            a. rate
            b. pathway
            c. residence time
            d. downhill transport 'spiral length' :

• uptake length - downstream distance a released atom is transported until it is captured again;
• turnover length- downstream distance moved while in organisms

                Important for looking at distubances and the responses to disturbances because they propagate downstream

    F. Controls on lotic community structure -- What controls the biosystem?
        1. density dependent = 'biotic interactions'; function of how many organisms are around
            a. competition - for space
            b. predation
            c. parasitism
        2. density independent = 'abiotic factors'
            a. floods
            b. changes in substrate
            c. changes in temperature (e.g. freezing)
        3. Which mechanism dominates? Evidence for both
            a. Abiotic factors have clear influences
            b. Correlational evidence -- density dependent correlations of 1 species with another.
            c. Experiments
                1)  Have shown clear effects of grazers feeding on periphyton
                2)  Manipulation of insect predators in cages have demonstrated biotic density dependent control
                3)  Manipulations of fish predators in cages -- small biotic effects (when you change fish abundance,
                        the abundance of insects doesn't change much), although big behavioral effects

            d. Conclusions
                1) Evidence favors strong abiotic controls
                2) Importance of time scale, large abiotic factors (flood/freeze) reset the system frequently so that you don't get
                    enough time/high enough densities for important biotic effects in many streams
                3) In more stable conditions you get lots of biotic interactions
                3) Really not settled yet
                    a) continuum of regulation -- Peckarsky
                    b) long-term records -- to see how often resetting occurs
                    c) density dependent effects -- often subtle; behavioral

GOOD GENERAL STREAM ECOLOGY TEXT BOOKS

Allan, J.D. 1995. Stream ecology: structure and function of running waters. Chapman &  Hall.
Cushing, C. and J.D. Allan. 2001. Streams: Their Ecology and Life. Academic Press.
Giller and Malmqvist. 1998. The biology of streams and rivers. Oxford University Press.
 
 

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