1. Littoral Community: Larger Plants and Benthic Animals Wetzel Including, but not solely: Chs. 19, 20, 22 pp. 600-629, 665-725

2. Lacustrine Zonation and Terminology p. 42 class notes, Fig. 8-2 Wetzel Epilittoral – above water, not subject to spray Supralittoral – above water, subject to spray by waves Littoral –Eulittoral – between highest and lowest seasonal water levels, subject to disturbances of waves

3. Lacustrine Zonation and Terminology cont. Littoral cont. –Infralittoral – 3 regions based upon distribution of macrophytic vegetation Upper Infralittoral – emergent rooted vegetation Middle Infralittoral – floating-leaved rooted vegetation Lower Infralittoral – submerged rooted vegetation –Littoriprofundal – transitional zone of attached algae and bacteria; cyanobacteria important in this zone Profundal – free of vegetation

4. Lacustrine Zonation and Terminology

5. Benthic Algae Terminology Algae by attachment places: –Rocks: epilithic –Sediments: epipelic –Plants: epiphytic –Animals: epizooic –Sand: epipsammic

6. Benthic Animals Benthos defined – animals associated with substrata Distribution, abundance, productivity determined by ecological processes (1)Historical events that limit an organism reaching an environment (2)Physiological limitations of the organism at any life stage for a specific habitat (3)Availability of energy resources (food) (4)Ability of species to tolerate competition from endemic organisms

7. Esrom Lake diagrams from Berg, 1938 – distributions from 0.2-20m

8. Esrom Lake Distributions cont.

10. Benthic Animals cont. Major groups: Protozoa –Diverse group –Ciliate and flagellate protozoans important –Little known about productivity Porifera –Freshwater sponges –Few, but some occur in Lake Erie Hydrazoans –Freshwater hydras

11. Benthic Animals cont. Flatworms/Turbellarians Roundworms/Nematodes Horsehair worms/Nematomorpha Bryozoans Annelids –Oligochaets –Leeches Arachnids –Water mites/Hydrachnida Ostracods –microcrustaceans

12. Benthic Animals cont. Malocostracean Crustaceans –Mysids/Opossum Shrimp –Isopods/Sowbugs –Decapods/Crayfish, Shrimp –Amphipods/Scuds Mollusks –Gastropods –Bivalves Aquatic Insects

13. Benthic invertebrates as indicators of lake health Just as fish can be indicators, so can benthic invertebrates These organisms have high oxygen demands –As the lake becomes more eutrophic, decomposition in profundal waters increases –Decreases oxygen concentration, more tolerant organisms become more abundant in benthos –In Lake Erie, as lake became more eutrophic, lost Hexagenia sp. (burrowing mayfly); as lake became less eutrophic due to nutrient controls, Hexagenia has come back

14. Aquatic Macrophytes Net productivity does not overlap with max biomass due to high respiration and death –Fig 18-8 Macrophytes can significantly contribute to oxygen concentration –Fig 18-3

15. Types of Macrophytes Rooted emergent Rooted floating Floating Submersed

16. Macrophytes and Primary Productivity Table 18-10 Compare macrophytes to phytoplankton Macrophytes as productive as a rain forest? How much of this productivity is transferred to higher trophic levels?

17. Macrophyte interactions with zooplankton Two distinct possibilities have been proposed (1)Zooplankton avoid macrophytes due to mechanical interference (2)Zooplankton use macrophytes as refugia from predation

18. Zooplankton Avoidance of Macrophytes Pennak (1973) –Reasons for avoidance (i)Shadows cause zooplankton to avoid (ii)Macrophytes release substance that repels zooplankton Used a tank with divider to measure zooplankton preference Four treatments –Control – both sides empty –One side had vegetation –One side had plastic vegetation –One side vegetation, other side plastic vegetation Response was % Daphnia on vegetated side

19. Pennak cont. Results: –Shadowing effect was less important than repellent effect –However, repellent effect seemed to wear off after approximately sixteen days

20. Macrophytes as Refugia Lauridsen and Lodge (1996) Hypothesis: macrophytes can serve as both a refuge and an impediment for Daphnia depending upon the presence/absence of fish Response: % of Daphnia in central area of tank –Central area was near fish but had no macrophytes

21. Lauridsen and Lodge results Zooplankton also avoid macrophytes when no fish are present Zooplankton avoid the open area when fish or fish odor is present Therefore, behavioral avoidance of fish supercedes behavioral avoidance of macrophytes Consequently, macrophytes may serve as refugia in shallow lakes with fish where diel vertical migration (DVM) is not possible, but zooplankton still avoid macrophytes

22. Conclusions Spatial heterogeneity occurs and is of importance in lakes – i.e. Esrom lake Macrophytes are not just simple, non- interactive plants in the water Organisms specialize to all sorts of different habitats – kinds of benthic algae, different kinds of plants, etc.