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Podostemum Ceratophyllum and Patterns of Fish Occurrence and Richness in a Southern Appalachian River

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Podostemum Ceratophyllum and Patterns of Fish Occurrence and Richness in a Southern Appalachian River PODOSTEMUM CERATOPHYLLUM AND PATTERNS OF FISH OCCURRENCE AND RICHNESS IN A SOUTHERN APPALACHIAN RIVER by JANE ELIZABETH ARGENTINA B.S. Tufts University, 2000 (under the direction of Mary C. Freeman) ABSTRACT We estimated percent cover of Podostemum ceratophyllum Michx. and fish richness at twenty randomly selected shoals within a 42 kilometer reach of the Conasauga River to determine if changes in species richness or species presence are related to variable levels of P. ceratophyllum coverage. We also assessed fish behavioral shifts with changes in P. ceratophyllum coverage in an experimental study. P. ceratophyllum was correlated positively with increasing bed sediment sizes and decreasing canopy cover, and negatively associated with drainage area. Fish species richness peaked in the center portion of the study reach, declining further downstream, coincident with range contraction of six lotic fishes. Benthic insectivorous fishes showed significant preference for habitat with increased P. ceratophyllum coverage compared with low coverage. River-wide changes in P. ceratophyllum could influence fish productivity through lower insect biomass and abundances, although water quality may have a greater effect on fish abundance and persistence. INDEX WORDS: Conasauga River, Podostemum ceratophyllum, fish richness, shoal, turbidity, drainage area, benthic insectivores, experimental manipulation, aquatic macrophytes ii PODOSTEMUM CERATOPHYLLUM AND PATTERNS OF FISH OCCURRENCE AND RICHNESS IN A SOUTHERN APPALACHIAN RIVER by JANE ELIZABETH ARGENTINA B.S. Tufts University, 2000 (under the direction of Mary C. Freeman) ABSTRACT A Thesis Submitted to the Graduate Faculty of the University of Georgia in Partial Fulfillment of the Requirements for the Degree MASTER OF SCIENCE ATHENS, GA 2006 iii © 2006 JANE ELIZABETH ARGENTINA ALL RIGHTS RESERVED iv PODOSTEMUM CERATOPHYLLUM AND PATTERNS OF FISH OCCURRENCE AND RICHNESS IN A SOUTHERN APPALACHIAN RIVER by JANE ELIZABETH ARGENTINA Major Professor: Mary C. Freeman Committee: Byron J. Freeman Judy L. Meyer Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia August 2006 viv Acknowledgements This work is a combined effort of many people, with which I was able to complete this thesis. I’d really like to thank Mary Freeman, my committee Judy Meyer and Bud Freeman who inspire me to know more about our aquatic environments. Thanks to the Freeman lab, and the host of people who helped in the field and in the lab. Thanks to my family, and especially my Vinny, for all their love and support. Benjamin Petty, Chris Petty, Dr. Goode and Dalton Utilities who gave us access to the river, without which this work could not have been done. Funding was provided by the Georgia DNR under Section 6 of the Endangered Species Act. viv TABLE OF CONTENTS Page ACKNOWLEDGEMENTS……………………………………………………… iv CHAPTER 1. INTRODUCTION……………………………………………………….. 1 2. HABITAT AND LANDSCPE EFFECTS ON DISTRIBUTION OF A BENTHIC MACROPHYTE, PODOSTEMUM CERATOPHYLLUM, IN A SOUTHERN APPALACHIAN RIVER, AND ASSOCIATIONS WITH LOTIC FISHES…………………………………………………....…... 5 3. BENTHIC FISH RESPONSE TO EXPERIMENTALLY MANIPULATED DENSITY OF AN AQUATIC MACROPHYTE, PODOSTEMUM CERATOPHYLLUM......................................................75 4. CONCLUSIONS…………………………………………………...........101 APPENDICES A. SUPPLEMENTAL MATERIAL FOR CHAPTER 2………………103 B. SUPPLEMENTAL MATERIAL FOR CHAPTER 3………………107 1 Chapter One Introduction Species richness is shaped by the landscape and the riverscape at multiple scales in river systems. Benthic habitat structure and heterogeneity, in particular, can strongly influence local abundances of riverine biota. Aquatic plants are important habitat features for biota inhabiting low velocity areas such as sea grass beds (Heck and Wetstone 1977, Heck and Crowder 1991), lake edges (Werner and Hall 1988), and river margins (Newman 1991, Fritz et al. 2004). I have focused on a submerged aquatic plant, Podostemum ceratophyllum Michx., that is known to provide heterogeneous habitat to aquatic macroinvertebrates (Nelson and Scott 1962, Hutchens et al. 2004), and with which a number of benthic fishes have been reported to associate (Etnier and Starnes 1993, Marcinek 2003, Hagler 2006). P. ceratophyllum (Podostemaceae) is a completely submerged filamentous dicotyledon without true roots; it occurs in high gradient upland waters such as Appalachian streams and rivers (Hutchens et al. 2004, Hagler 2006), and shoal and riffle habitat of Piedmont rivers (Mullholland and Lenat 1992, Nelson and Scott 1962, Grubaugh and Wallace 1995, Marcinek 2003, Fritz et al. 2004). It characteristically occurs in fast-flowing waters where it attaches with hyphae to the surface of rocks. By increasing surface area, reducing scour, and increasing availability of organic matter and epiphytic periphyton, P. ceratophyllum enhances benthic habitat for aquatic invertebrates (Grubaugh and Wallace 1995, Hutchens et al. 2004), and may do the same for riverine fishes. In an observational study (Chapter 2), I have examined longitudinal patterns of occurrence of P. ceratophyllum and of fishes in relation to natural features and land use 2 in a biologically diverse southeastern Appalachian river. Specifically, I have evaluated relationships between P. ceratophyllum and habitat variables measured at three scales – local (i.e., habitat at the point of observation), shoal (channel slope, canopy cover) and landscape (drainage area, land use upstream and in the riparian buffer). I have also assessed relations between the longitudinal distribution of P. ceratophyllum in the Conasauga River and (1) fish species richness and (2) occurrence probabilities for lotic fish species, using survey methods that account for incomplete species detectabilities (Nichols et al. 1998, MacKenzie et al. 2002). In an experimental study (Chapter 3), I tested the influence of P. ceratophyllum on benthic fish habitat use. Benthic habitats provide protection from predators and refugia from high velocities for a broad range of aquatic animals. Benthic aquatic macroinvertebrate biomass generally increases in areas with complex habitats, including heterogenous bed sediments, woody debris and aquatic plants (Rabeni and Minshall 1977, Anderson et al. 1978, Allan 1995, Meyer et al. 1997). I expected that benthic insectivorous fishes would show a behavioral response to increased coverage of P. ceratophyllum and that this response may be due to increases in food availability. To test for a behavioral response by benthic fishes, I manipulated P. ceratophyllum cover and quantified fish abundances, as well as insect density and biomass, two weeks after the manipulation, in two experimental patches measuring about 30 m2. Together these observational and experimental studies provide new data and insights on factors controlling abundance of P. ceratophyllum and fish responses to macrophyte coverage at shoal and local habitat scales. 3 Literature Cited Allan, J. D. 1995. Stream Ecology. Kluwer Publishers. Dordrecht, The Netherlands. Anderson, N.H., J.R. Sedell, L.M. Roberts and F.J. Triska. 1978. The role of aquatic invertebrates in processing of wood debris in coniferous forest streams. American Midland Naturalist 100(1): 64-82. Etnier, D.A. and W.C. Starnes. 1993. The Fishes of Tennessee. The University of Tennessee Press, Knoxville. Fritz, K.M., M.M. Gangloff, and J.W. Feminella. 2004. Habitat modification by the stream macrophyte Justicia americana and its effects on biota. Oecologia 140: 388-397. Grubaugh, J.W. and J.B. Wallace. 1995. Functional structure and production of the benthic community in a Piedmont river: 1956-1957 and 1991-1992. Limnology and Oceanography 40(3):490-501. Hagler, M.M. 2006. Effects of natural flow variability over seven years on the occurrence of shoal-dependent fishes in the Etowah River. M.S. Thesis, University of Georgia, Athens, GA. Heck, K.L.J. and L.B. Crowder. 1991. Habitat structure and predator-prey interactions in vegetated aquatic systems. In: Bell, S.S., E.D. McCoy and H.R. Mushinsky (eds) Habitat structure: the physical arrangement of objects in space. Chapman and Hall. London, pp. 281-299. Heck, K.L.J. and G.S. Wetstone. 1977. Habitat complexity and invertebrate species richness and abundance in tropical seagrass meadows. Journal of Biogeography 4:135-142. 4 Hutchens, J.J., J.B. Wallace, and E.C. Romaniszyn. 2004. Role of Podostemum ceratophyllum Michx. in structuring benthic macroinvertebrate assemblages in a southern Appalachian river. Journal of North American Benthological Society 23(4):713-727. Marcinek, P.A. 2003. Variation of fish assemblages and species abundances in the upper Flint River shoals, Georgia. M.S. Thesis. University of Georgia, Athens, GA. Meyer, J.L., A.C. Benke, R.T. Edwards, and J.B. Wallace. 1997. Organic matter dynamics in the Ogeechee River, a blackwater river in Georgia, USA. Journal of the North American Benthological Society 16: 82-87. Mullholland, P.J. and D.R. Lenat. 1992. Streams of the southeastern Piedmont, Atlantic drainage. In C.T. Hackney, S.M. Adams and W.H. Martin (eds.), Biodiversity of the southeastern United States: aquatic communities. John Wiley & Sons, Inc., New York. pp. 193 – 232. Nelson, D.J. and D.C. Scott. 1962. Role of detritus in the productivity of a rock-outcrop community in a Piedmont stream. Limnology and Oceanography 7: 396-413. Newman, R.M. 1991. Herbivory and detritivory on fresh-water macrophytes by invertebrates - a review. Journal of the North American Benthological
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