Grazer species effects on epilithon nutrient composition

Summary 1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no-grazer treatment. 4. Traditionally, studies have focused on how grazer-induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.

[1]  D. Lodge,et al.  Multi-Trophic-Level Impact of Sublethal Interactions between Bass and Omnivorous Crayfish , 1995, Journal of the North American Benthological Society.

[2]  Patrick J. Mulholland,et al.  Top‐Down and Bottom‐Up Control of Stream Periphyton: Effects of Nutrients and Herbivores , 1993 .

[3]  S. V. Smith,et al.  C:N:P ratios of benthic marine plants1 , 1983 .

[4]  L. Solórzano DETERMINATION OF AMMONIA IN NATURAL WATERS BY THE PHENOLHYPOCHLORITE METHOD 1 1 This research was fully supported by U.S. Atomic Energy Commission Contract No. ATS (11‐1) GEN 10, P.A. 20. , 1969 .

[5]  G. Lamberti,et al.  Invading Crayfish in a Michigan Stream: Direct and Indirect Effects on Periphyton and Macroinvertebrates , 1996, Journal of the North American Benthological Society.

[6]  Patrick J. Mulholland,et al.  Seasonally shifting limitation of stream periphyton: response of algal populations and assemblage biomass and productivity to variation in light, nutrients, and herbivores , 2000 .

[7]  Helmut Hillebrand,et al.  BIOVOLUME CALCULATION FOR PELAGIC AND BENTHIC MICROALGAE , 1999 .

[8]  K. Keating A system of defined (Sensu stricto) media for daphnid (Cladocera) culture , 1985 .

[9]  J. Elser,et al.  ROSEMARY MACKAY FUND ARTICLE: Ecological stoichiometry of trophic interactions in the benthos: understanding the role of C:N:P ratios in lentic and lotic habitats , 2002, Journal of the North American Benthological Society.

[10]  A. Steinman,et al.  Effects of Herbivore Type and Density on Chemical Composition of Algal Assemblages in Laboratory Streams , 1987, Journal of the North American Benthological Society.

[11]  James J. Elser,et al.  THE STOICHIOMETRY OF CONSUMER‐DRIVEN NUTRIENT RECYCLING: THEORY, OBSERVATIONS, AND CONSEQUENCES , 1999 .

[12]  Margaret A. Nemeth,et al.  Applied Multivariate Methods for Data Analysis , 1998, Technometrics.

[13]  N. Grimm ROLE OF MACROINVERTEBRATES IN NITROGEN DYNAMICS OF A DESERT STREAM , 1988 .

[14]  Yosef Cohen,et al.  MOOSE BROWSING AND SOIL FERTILITY IN THE BOREAL FORESTS OF ISLE ROYALE NATIONAL PARK , 1993 .

[15]  G. Lamberti,et al.  Effects of N: P ratio and total nutrient concentration on stream periphyton community structure, biomass, and elemental composition , 2001 .

[16]  James J. Elser,et al.  Organism size, life history, and N:P stoichiometry , 1996 .

[17]  A. E. Greenberg,et al.  Standard methods for the examination of water and wastewater : supplement to the sixteenth edition , 1988 .

[18]  D. O. Hessen,et al.  Nitrogen and phosphorus excretion from the noble crayfish, Astacus astacus L., in relation to food type and temperature , 1992 .

[19]  G. Rhee Effects of N:P atomic ratios and nitrate limitation on algal growth, cell composition, and nitrate uptake 1 , 1978 .

[20]  Alan D. Steinman,et al.  Effects of Biomass, Light, and Grazing on Phosphorus Cycling in Stream Periphyton Communities , 1995, Journal of the North American Benthological Society.

[21]  S. McNaughton,et al.  Promotion of the cycling of diet-enhancing nutrients by african grazers , 1997, Science.

[22]  F. Friedl Nitrogen excretion by the fresh water pulmonate snail, Lymnaea stagnalis jugularis Say. , 1974, Comparative biochemistry and physiology. A, Comparative physiology.

[23]  G. Lamberti,et al.  Taxonomic and regional patterns in benthic macroinvertebrate elemental composition in streams , 2005 .

[24]  F. P. Healey,et al.  Indicators of Phosphorus and Nitrogen Deficiency in Five Algae in Culture , 1979 .

[25]  J. Elser,et al.  Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere , 2002 .

[26]  A. Rosemond Interactions among irradiance, nutrients, and herbivores constrain a stream algal community , 1993, Oecologia.

[27]  A. Steinman,et al.  Productive Capacity of Periphyton as a Determinant of Plant‐Herbivore Interactions in Streams , 1989 .

[28]  W. D. Russell-Hunter,et al.  Variation in aufwuchs at six freshwater habitats in terms of carbon biomass and of carbon: Nitrogen ratio , 1974, Hydrobiologia.

[29]  U. Sommer,et al.  The nutrient stoichiometry of benthic microalgal growth: Redfield proportions are optimal , 1999 .

[30]  L. Krienitz,et al.  PHYLOGENETIC POSITION OF THE OOCYSTACEAE (CHLOROPHYTA) , 2000, Journal of phycology.

[31]  J. Urabe N and P Cycling Coupled by Grazers' Activities: Food Quality and Nutrient Release by Zooplankton , 1993 .

[32]  R. Sterner The Ratio of Nitrogen to Phosphorus Resupplied by Herbivores: Zooplankton and the Algal Competitive Arena , 1990, The American Naturalist.

[33]  V. Resh,et al.  Stream Periphyton and Insect Herbivores: An Experimental Study of Grazing by a Caddisfly Population , 1983 .

[34]  M. Bothwell Growth Rate Responses of Lotic Periphytic Diatoms to Experimental Phosphorus Enrichment: The Influence of Temperature and Light , 1988 .

[35]  A. Covich,et al.  Effects of an omnivorous crayfish (Orconectes rusticus) on a freshwater littoral food web , 1994 .

[36]  M. Vanni,et al.  Stoichiometry of nutrient recycling by vertebrates in a tropical stream: linking species identity and ecosystem processes , 2002 .

[37]  J. Elser,et al.  Elemental Composition of Littoral Invertebrates from Oligotrophic and Eutrophic Canadian Lakes , 2003, Journal of the North American Benthological Society.

[38]  H. Hillebrand,et al.  Effect of grazing and nutrient supply on periphyton biomass and nutrient stoichiometry in habitats of different productivity , 2001 .

[39]  Anthony Ricciardi,et al.  Extinction Rates of North American Freshwater Fauna , 1999 .

[40]  R. Hall,et al.  Exotic snails dominate nitrogen and carbon cycling in a highly productive stream , 2003 .

[41]  M. Vanni Nutrient Cycling by Animals in Freshwater Ecosystems , 2002 .

[42]  A. Covich,et al.  Predator avoidance by the freshwater snailPhysella virgata in response to the crayfishProcambarus simulans , 1991, Oecologia.

[43]  David B. Lewis Trade-offs between growth and survival : responses of freshwater snails to predacious crayfish , 2001 .

[44]  W. Dodds Misuse of inorganic N and soluble reactive P concentrations to indicate nutrient status of surface waters , 2003, Journal of the North American Benthological Society.

[45]  A. Steinman 12 – Effects of Grazers on Freshwater Benthic Algae , 1996 .

[46]  T. Burton,et al.  Growth and Abundance of the Crayfish Orconectes propinouus in a Hard Water and a Soft Water Stream , 1993 .

[47]  A. Covich,et al.  Predator-Avoidance Responses in Freshwater Decapod-Gastropod Interactions Mediated by Chemical Stimuli , 1994, Journal of the North American Benthological Society.

[48]  J. Elwood,et al.  Role of Nutrient Cycling and Herbivory in Regulating Periphyton Communities in Laboratory Streams , 1991 .

[49]  G. Schnell,et al.  Factors determining population density and size distribution of a freshwater snail in streams: effects of spatial scale , 1990 .

[50]  J. Elser,et al.  Effects of Caddisfly Grazers on the Elemental Composition of Epilithon in a Boreal Lake , 2002, Journal of the North American Benthological Society.

[51]  D. Jablonski Extinctions in the fossil record , 1994 .