EVPP 550 Waterscape Ecology and Management – Lecture 11 Professor R. Christian Jones Fall 2007 Lake Biology – Benthos Profundal Benthos • Profundal habitat can be very challenging in lakes – Cold for most of the year due to summer stratification – Anaerobic in mesotrophic and eutrophic lakes – Poor food quality: no resident primary producers, all food is imported and “stale” Lake Biology – Benthos Profundal Benthos • A few groups have been able to adapt to this environment • Those which have can prosper if conditions are not too severe • Examples are chironomids (midges), chaoborus (phantom midges) and oligochaetes Lake Biology – Benthos Profundal Benthos • Chironomids have been extensively studied • Some species can maintain constant respiration even at low DO • Some can withstand no oxygen for up to 4 months at 10oC • Food supply of profundal chironomids is surface sediment particles ingested in bulk with algae and bacteria selectively assimilated Lake Biology – Benthos Profundal Benthos • Adaptations – Hemoglobin • Helps to bind and store limited amounts of O2 – Anaerobic glycolysis • Can split carbohydrates to produce energy with using oxygen • Similar to what happens in your muscles under strenuous activity • Accumulate an oxygen debt that must later be satisfied – Stop growth, become dormant Lake Biology – Profundal Benthos • Life History of Chironomus anthrocinus in L. Esrom – Egg mass deposited at night in May on lake surface near shore – Water currents spread the eggs throughout the lake as they sink to the bottom – By June, hatching occurs, food is abundant and larvae double in size by early July – Oxygen is depleted in summer and growth stops Lake Biology – Profundal Benthos • Life History of Chironomus anthrocinus in L. Esrom – Fall overturn brings oxygen to the bottom satifying the oxygen debt and allowing fresh growth to occur based on food still in the water column – Animals are quiescent during winter, but perk up again the following spring with onset of new food Lake Biology – Profundal Benthos • Life History of Chironomus anthrocinus in L. Esrom – Some have grown large enough to emerge after one year, but many need a little more growth and emerge the second year – Emergence occurs at the surface, mating occurs, eggs are laid, and adults die within a few days – Highly synchronized pop’n Lake Biology – Profundal Benthos • Chaoborus – Phantom midge – Alternates between plankton and benthos – Prey mostly on zooplankton and are preyed upon by fish – Migrates daily from sediment surface to photic zone – Under anaerobic conditions, may stop downward migration at the thermocline Lake Biology – Profundal Benthos • Oligochaetes – Development is unsynchronized – Burrow through surface sediment, digesting bacteria, mixing sediments, and recycling nutrients Littoral Zone • Portion of lake where photic zone includes the bottom The Littoral Zone Macrophytes • Macrophytes – Plants whose overall structure is visible to the naked eye – Distribution in lakes is subject to two basic constraints: • Water must be shallow enough for light to reach the bottom (= littoral zone as we have defined) • Physical stability sufficient to allow plants to grow to the bottom The Littoral Zone Macrophytes • Characteristics – General Morphology • 4 basic morphological types typically occupying “zones” of increasing depth • Emergent • Floating-leaved • Submersed • Unrooted The Littoral Zone Macrophytes • Emergent macrophytes – Occupy the transition zone between land and water – Rooted in sediment or saturated soils (anaerobic) – Shoots and leaves extend into the air so, like terrestrial plants, they must be self-supporting & get CO2 from air – Mostly angiosperms – Ex.: cattails, wild rice The Littoral Zone Macrophytes • Floating-leaved macrophytes – Root in sediment, leaves float on surface – Connections are via stems or petioles – 0.5 m < z < 3 m – Need to have some standing water, but limited by petiole or stem length – In case of water lilies, both root and stem are underwater and petioles (leaf stem) extends through water to surface leaves – A patch of water lilies may actually be one plant – Ex: yellow water lily (Nuphar), white water lily (Nymphaea) The Littoral Zone Macrophytes • Submersed Macrophytes – Whole plant is underwater – 0.5 < z < 10 m (angiosperms), up to 100 m for mosses, Chara – No supporting tissue, rely on turgor pressure and buoyancy to maintain erect form – Underwater leaves often finely dissected, but may be laminar – May have heterophylly (different underwater vs. surface leaves) – Ex: Myriophyllum (milfoil), Potomogeton (pondweed), Chara (stonewort), Isoetes (water fern) The Littoral Zone Macrophytes • Unrooted macrophytes – Floating • Lemna (duckweed) • Eichornia (water hyacinth) – Submersed • Ceratophyllum (coontail) The Littoral Zone Macrophytes • Taxonomy – Charaphytes (stoneworts) • Algal group related to green algae • Macroscopic form • Ex: Chara, Nitella – Bryophytes (mosses, liverworts) • Plants with some tissue and reproductive specialization, but no vascular tissue (xylem, phloem) • Ex: Sphagnum The Littoral Zone Macrophytes • Taxonomy – Ferns and Fern Allies • Plants with vascular tissue, but no flowers • Ex: Isoetes (submersed macrophyte found in soft water) • Ex: Equisetum (horsetail) (emergent macrophyte) The Littoral Zone Macrophytes • Taxonomy – Gymnosperms • Vascular tissue • Reproductive: “cones” • Ex: Bald Cypress (emergent) – Angiosperms • • • • • • • • Vascular tissue Flowers Ex: Cattail (Typha) Ex: Water Lilies (White and Yellow) Ex: Myriophyllum (milfoil) Ex: Hydrilla Ex: Potamogeton (pondweed) Ex: Vallisneria (water celery) Macrophytes – Factors Affecting Growth • Low oxygen levels around roots – Sediments are usually anoxic, but roots need oxygen or growth will be inhibited – Some species have vertical air tubes called lacunae which extend from the shoots down into the roots to help aerate – Root cells may be able to withstand oxygen debt Macrophytes – Factors Affecting Growth • Inorganic carbon supply – Low rate of diffusion of CO2 through bulky macrophyte tissue could lead to carbon shortage – Plants can also use CO2 and in very soft water, uptake can occur through roots Macrophytes – Factors Affecting Growth • Depth – Pressure – Vascular macrophytes do not grow to a depth of more than 10 m (representing 1 extra atmosphere of pressure) – This seems to be related to the effect of this extra pressure on the xylem and phloem – However, mosses have been found at up to 165 m and Chara to 64 m in Lake Tahoe, for example Macrophytes – Factors Affecting Growth • Depth - Light – Two effects: Light & Pressure – Water transparency is highly correlated with depth to which macrophytes can grow – Note that maximum depth of colonization is less than photic zone depth which is about double Secchi disc depth Macrophytes – Factors Affecting Growth • Depth - Light – One way that macrophyte communities respond to potential light limitation is to favor species that develop a canopy as opposed to those the grow near the sediment surface (rosettes) Macrophytes – Factors Affecting Growth • Nutrients – N&P can be taken up by roots and shoots – Relative importance of root vs. shoot uptake depends on sediment vs. water concentrations – Ex: Lake Wingra, WI • 73% of P by roots • 27% of P by shoots – Root uptake is then translocated to shoots to fuel growth Macrophytes – Factors Affecting Growth • Sediment Stability – Texture is important • Need fine particles: fine sand, silt or clay • Course sand, cobble, boulders are not good rooting medium – Stability is also important • If sand is moving, like on a beach plants will not become established Macrophytes – Patterns of Abundance & Production • Seasonal – In temperate areas, macrophytes are very seasonal in their growth – Maximum development in late summer – However, some dieback over much of the year – In fact, plants create and shed shoots continuously Macrophytes – Factors Affecting Growth • Productivity determination – Maximum standing crop • But this ignores biomass that was shed building up to maximum – C-14 approach • Measure C-14 uptake by actively photosynthesizing parts of plant – Cohort analysis • See previous page Macrophytes – Spatial Patterns • Within lake – Macrophytes generally cover only those parts of the right habitat (light, substrate, etc.) • Between lakes – Great differences between lakes Littoral Zone Periphyton • Characteristics – General Morphology • Algae: unicells, filaments, colonies • 2 general types of attachment – Adnate: cells in close contact with substrate, hard to dislodge – Loose: cells only loosely attached, easily dislodged – Taxonomy • All groups of algae represented, esp – Cyanobacteria, diatoms, greens Littoral Zone Periphyton • Factors affecting development – Substrate Availability • The amount of surface habitat obviously influences the abundance of periphyton • Could be fairly static like bottom area in photic zone or very dynamic like annual plant surfaces – Light • Have a very similar photosynthesis-light relationship as phytoplankton Littoral Zone Periphyton • Factors affecting development – Nutrients • Can periphyton get nutrients from their host substrate? • Results seem to suggest this is not a major factor • Label P in sediments, grow macrophytes, less than 5% of P in epiphytes comes from sediment • High correlation with lake water P Littoral Zone Periphyton • Patterns of Abundance and Productivity – Epiphytic periphyton vary both with depth and seasonally – These variations are a combination of: • Changes in the density of epiphytes on the macrophyte • Changes in the amount of macrophyte substrate available at different depths and times Littoral Zone Periphyton • Productivity would also need to take into account variations in light and P-I response Littoral Zone - Periphyton • • • • Resulting productivity could vary seasonally and from one year to the next Note day-to-day variation in production (light driven) Note different seasonal pattern (substrate availability driven) Note rough equivalence of 10 mg C produced per mg Chl a present per day Littoral Zone – Littoral Invertebrates • Characteristics – Include a much larger suite of organisms than found in the profundal benthos – Some of the dominant groups include: • Flatworms • Oligochaetes • Molluscs – Snails – Bivalves • Arthropods – Crustaceans – Insects Littoral Zone – Littoral Invertebrates • Characteristics – A wide variety of feeding strategies including: • Grazers/herbivores (due to presence of primary producers in the littoral zone) • Detritivores • Predators – Littoral grazers tend to focus on periphyton as it is much more digestable – Macrophyte production tends to get utilized as detritus Littoral Zone – Littoral Invertebrates • Characteristics – Type of predators • Lurking – – – – Dragonflies Sit in a concealed position Attack prey as they come by Concealed, but dependent on prey movement • Hunting – Water bugs – Actively search for prey – Often well-armoured, taste bad, and move quicky to avoid predators – Can capture both moving and stationary prey Littoral Zone – Littoral Invertebrates • Characteristics – Littoral zone can be an area of great physical and chemical complexity – Allows a very high diversity, but also presents some significant sampling problems – Heterogeneous distribution – Difficulties in capturing organisms within vegetation, rocks, logs, etc. Littoral Invertebrates – Patterns of Abundance • Seasonal and spatial patterns • Examine results from a study of littoral invertebrates in the tidal freshwater Potomac River – Organisms captured by dropping nets over 0.5 m2 of weedbed – Nets closed by diver at bottom and brought to surface where organisms were removed from vegetation and preserved • Study design – 3 bed types: open water, Hydrilla, mixed – 2 months (July, August) – 5 replicates each Littoral Zone – Littoral Invertebrates • Results – Macrophytes harbored much higher abundance of macroinvertebrates than open water – Taxa list was similar at all sites, but relative abundance differed both with plant type and month Littoral Zone – Littoral Invertebrates • Cluster analysis – Confirmed differences between veg and open water – Suggested that variation between months was more important that variation between plant types Littoral Zone – Littoral Invertebrates • PCA – Reinforced importance of plants – And the effect of month over plant type