MSU LIBRARIES gig. RETURNING MATERIALS: PIace in book drop to remove this checkout from your record. FINES wiII be charged if book is returned after the date stamped beIow. al‘v - I (- Wok-l~t .‘l..‘. " ‘ 'IQo U o RELATIONSHIPS BETWEEN PRODUCTION AND ABUNDANCE OF RANID (Rana pipéenb, R. cateabeéana, and R. c£amiiunA) TADPOLES AND THE EFFECTS OF TADPOLE GRAZING ON ALGAL PRODUCTIVITY AND DIVERSITY AND ON PHOSPHORUS, NITROGEN, AND ORGANIC CARBON By Diana L. Neigmann A DISSERTATION Submitted to Michigan State University in partial fquiIIment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Fisheries and Niidiife 1982 '. §.. 001' . O o. . . (a “tag;- I C I . 5 (UP ‘a' I a O Co ‘v t (I! q. ABSTRACT RELATIONSHIPS BETWEEN PRODUCTION AND ABUNDANCE OF RANID (Rana pipicnb, R. catebbeiana, and R. clamitans) TADPOLES AND THE EFFECTS OF TADPOLE GRAZING ON ALGAL PRODUCTIVITY AND DIVERSITY AND ON PHOSPHORUS, NITROGEN, AND ORGANIC CARBON By Diana L. Weigmann The influence of stocking different densities and biomasses of tadpoles on their subsequent survival, growth, and production was as- sessed by culturing leopard frog (3. pipiens), bullfrog (3. catesbeiana), and green frog (3. clamitans) tadpoles at eutrophic Lake One and meso- trophic Lake Four of the Water Quality Management Project, Michigan State University and by rearing leopard frog tadpoles in the laboratory. In the laboratory, stocking density had no significant effects on indi- vidual growth rates of 3. pipiens, which were stocked at five different densities but similar total biomasses. Survival and growth of all species were consistently higher in the more productive waters of Lake One. Stocking biomass was inversely related to daily production of monocultured 3. pipiens at both lakes (r = - 0.91 and r = - 0.85 at Lakes One and Four, respectively). Monocultured tadpoles of bullfrogs exhibited higher productivity than those of green frogs; however, in mixed culture the production of 3, catesbeiana was invariably lower than . that of 3, clamitans. Tadpole production was related to the supply of ?--'---'-z Uus v‘ 'U U Q 0...- ~30- .O'OP-QA' In I l:. ‘ 231.":‘4' ‘B; .IA-u- .‘ .‘vb :. (I a; . . . ‘-"Q. ~ \ - .Y o I ' ' 0‘ "D- u‘. n c '1... :I 3:. '3. Neigmann food/demand by tadpoles (r = 0.93) in laboratory and field experiments. Ecological, growth, and assimilation efficiencies were calculated for tadpoles and compared with efficiencies of other aquatic consumers. The effects of tadpole grazing on nutrient cycling (phosphorus, nitrogen, and carbon) and on phytoplanktonic and periphytic algal communities (abundance, productivity, and diversity) were influenced by the trophic status of the lakes and the biomass of tadpoles stocked. All species of ranid tadpoles consumed periphyton primarily, as much as 87% of the periphytic carbon was daily diverted by tadpoles into respi- ration and tissue production or the accumulation of sestonic carbon. Grazing by tadpoles lowered the C-N ratio of the periphytic standing crop, elevated phytoplanktonic numbers and volumes, and altered the diversity of the phytoplanktonic assemblage. The feasibility of culturing ranid tadpoles and their potential impacts on nutrients and biotic communities in artificial environments and littoral zones of natural waters were evaluated. ACKNOWLEDGEMENTS I wish to express my special appreciation to Dr. Louis A. Helfrich for his encouragement and assistance during this study and for his editorial comments on the dissertation. I am also grateful to Dr. Thomas G. Bahr for financial support and use of the Water Quality Management Project as a research site, Dr. Stanley Zarnoch for assistance with the FORTRAN program for diversity calculations, Dr. Niles R. Kevern for his cooperation with scheduling and chairing the final defense, Dr. Robert Tafanelli for reviewing the Discussion section, and Dr. Frank M. D'Itri and laboratory associates for chemical analyses. In addition, I wish to acknowledge the contributions of Dr. Richard A. Cole, Dr. Marvin M. Hensley, Dr. Clarence D. McNabb. Dr. Richard w. Merritt, and Dr. Howard E. Johnson. Funds for this research were provided by the Office of Water Resources Research (U.S. Department of the Interior) through grants administered by the Institute of Water Research at Michigan State University. ii ‘l.-.. O.. a. I I DOV. .- 1.... I . .. 'r I. I \ It 0 A... . . D I: .. y 9 D E z' s... I l a l.. n “. \. \ I ‘ v I o D. " r a, V - b h h a . \ I I \ ‘ 4‘-.u ‘ “'- 2 ‘ TABLE OF CONTENTS Page INTRODUCTION ........................... 1 MATERIALS AND METHODS ...................... Zl The Field Study ...................... 2l Description of the Site ............... 2i Description of the Flow-through System ........ 23 Experimental Design ................. 28 Tadpole Procedure .................. 29 Dry Weight and C-N Analysis of Tadpoles ....... 3l Limnological Assays ................. 32 Dissolved oxygen and temperature ........ 32 Gross primary productivity and community respiration ................. 34 Nutrient and phytoplankton sampling ...... 34 Phytoplankton methods ............. 36 Carbon, nitrogen, phosphorus, and alkalinity analyses ................... 38 Analysis of periphytic biomass and C-N content ................... 38 Relative contributions of periphyton and phytoplankton to GPP and R .......... 40 The Laboratory Experiment ................. 4l Natural Food Sources, Algae ............. 41 A Comparative Food, TetraMin ............. 45 RESULTS ............................. 46 Environmental Factors ................... 46 Differential Effects ................. 46 Daily Measurements of Temperature and Oxygen ..... 46 Extremes of Dissolved Oxygen ............. 50 Alkalinity and pH .................. 54 Tadpole Survival, Growth, and Production .......... 57 Varying Stocking Density and Biomass (Field Experiments One and Two) .............. 57 Lake One .................... Lake Four ................... Comparisons of Lakes One and Four ....... Tadpole Species Composition (Field Experiment Three) ....................... Reducing Food and Space per Individual (Laboratory Experiment) .............. Algal concentrations, TetraMin, and tadpole density ................... Ingestion and conversion efficiencies ..... Effects of Tadpoles on Organic Material and C, N, and P Dynamics ........................ Tadpole Grazing Intensity and Periphyton (Experi- ments One and Two) ................. Lake One .................... Lake Four ................... Comparisons of Lakes One and Four ....... Dissolved, Detrital, and Phytoplanktonic Organic Carbon (Experiments One and Two) .......... Lake One . . . .‘ ................ Lake Four ................... Comparisons of Lakes One and Four ....... Impacts of Tadpole Composition on Phytoplankton and Periphyton (Experiment Three) ......... Comparisons of Lakes One and Four ....... Comparisons of Experiment Three with Experi- ments One and Two .............. Effects of Varying Tadpole Stocking Rates on Nitrogen (Experiments One and Two) ......... Lake One .................... Lake Four ................... CarbonzNitrogen ratios for organics ...... The Effects of Tadpole Composition on Nitrogen (Experiment Three) ................. Lake One .................... Lake Four ................... Effects of Tadpoles on Phosphorus (Experiments One, Two, and Three) ................ iv Page 57 62 65 67 75 75 81 85 85 85 90 93 94 94 98 100 101 101 104 105 105 110 113 114 114 118 118 Phytoplanktonic Composition and Diversity ......... 124 Overview ...................... l24 Experiment One (Lakes One and Four) ......... l24 Imported lake water ....... ' ....... 124 Tadpole effects ................ l25 Experiment Two (Lakes One and Four) ......... l32 Imported lake water .............. l32 Tadpole effects ................ l33 Experiment Three (Lakes One and Four) ........ l39 Imported lake water .............. l39 Tadpole effects ................ 140 Community Metabolism ................... l42 Effects of Varying Tadpole Stocking Rates on Gross Primary Productivity (GPP) and GPP/Autotrophic Biomass (Experiments One and Two) ......... l42 Lake One ................... l42 Lake Four ................... 145 Effects of Varying Tadpole Stocking Rates on Community Respiration (R), R/Autotrophic Bio- mass (R/B), and GPP/R (Experiments One and Two) ....................... l47 Lake One ................... 147 Lake Four ................... lSO Partitioning the GPP and R of PhytOplankton and Periphyton (Ancillary Tests in Experiment Two) . . l53 Lake One ................... l53 Lake Four ................... l56 Comparisons of Community Metabolism (Experiments One and Two) ................... l57 Effects of Tadpole Composition on Community Metabolism (Experiment Three) ........... l59 Lake One ................... l59 Lake Four ................... l6l Partitioning phytoplanktonic and periphytic GPP and R (ancillary test) ......... 163 ..-. no'l. ’u-DO' - b 0. g I I '; :09 h C. q I" c A .0 v 3'. '.. ‘ . A "u: ‘- '0’. .". .’ A .. ..- .5 ‘ v IQ...-. Iv-n I . .._‘I . 0 HI. |.I. .'-I I .' 1- l D l'l Page DISCUSSION ........................... l67 Abiotic Environmental Effects .............. 167 Influences on Algal Communities in Aquaria ..... 167 Influences on Tadpoles ............... 168 Relationships Between Plant Production and Tadpole Production ....................... 172 Apparent Impact of Tadpoles on Algae and Nutrients .................... 172 Relationships Between the Ratio of the Supply of Food/Demand by Tadpoles and Tadpole Production .................... 183 Ecological Efficiencies of Tadpoles ........ 190 Growth and Assimilation Efficiencies ........ 199 Potential Roles of Tadpoles in Natural and Wastewater Treatment Systems ................... 204 l_ITERATURE CITED ........................ 217 APPENDIX A ........................... 243 APPENDIX B .......................... 250 vi .0. '\ if v 0". 0" .qu I... :- . a .4. O . A § B an n ‘- 0 o .0 Q .\ 3. (- F . . v G- r r . on. u: 3. .u- :a .r. «iv -v (. P V v F a an I F I. I . . U ‘IU ' -. . l in a. an... . . . F .b- I 9 c O Au. .fiU r) on c o a» a 3 n as o . o . u r I My. \a a\ .‘i I A v I .\v . v P (4 by A.» 6 y a 3 o u A} A 3 I, I B... a I u .a as bus as c. ”J H. v .. a v o. u. I. . . P5 .61 war. r as. p15 :5 in. Pb» ‘- ~ A :4 F I. a... M. are a .fll .5 pi .- O. nus c~ : 3- .. 3G nu. a/s m . are o.. s.. n!.. Table LIST OF TABLES Physico-chemical characteristics of Lakes One and Four from May-October, l976. Values represent the range of measurements (mg/l) on samples at lake outlets (Chem- istry Laboratory of Michigan Institute of Water Research) ........................ Concentrations of carbon and nitrogen (:50) measured in tadpoles after laboratory and field experiments Average concentrations of nutrients (mg/d) and phyto- plankton (cells/ml/d) at the three supply rates and their availability to tadpoles after dilution with 3.8 liters of water (largest CV for N, P, C, and algae were 12.4, l3.4, 9.2 and l8.8%, respectively) ...... Alkalinity (mg/l) and pH measurements for Experiment Three at Lakes One and Four ............... Densities, biomasses (wet weight, m9). and lengths (snout-vent, mm) of leopard frog tadpoles in Experi- ments One and Two at Lakes One and Four ......... Information (wet weight, mg and snout-vent length, mm) on tadpoles (R, pipiens) that died during Experiments One and Two ....................... Total production (mg of wet weight, carbon, and nitrogen) and production per unit of biomass for leopard frog tadpoles in Experiment One (2l d, 36l degrees d) and Two (25 CONTROL STRUCTURE LAKE SYSTEM 22 INDEPENDENT SPRAY UNITS (REMOTELY CONTROLLED AND PROGRAMMABLE .1 GROUNDWATER .......... ., and EVAPO- 5735‘“ TRANSPIRATION TERRESTRIAL SYSTEM Figure 1. Diagram of the general facilities of the Water Quality Management Project, Michigan State University, East Lansing, Michigan. 23 system was described in detail by Burton et a1. (1979). Water from Lakes One and Four, representing two extremes in water quality and nutrient availability, was used in this study (Table 1). Lake One had a surface area of 3.2 ha and water elevation of 271.9 m. Emergent vegetation was comprised mainly of cattails (11233) and willows (53115); dense mats of Cladophora covered the surface of the lake during much. of the surrmer and fall. Lake Four had a surface area of 4.98 ha and a water elevation of 268.2 m. Little emergent vegetation occurred around the perimeter of Lake Four, but a luxuriant growth of submerged Elodea covered most of the bottom. Description of the Flow-through System During the early spring of 1976, two sheds (2.44 m = length, 1.22 m = width, and 3.66 m = height at peak) were constructed on each of the two lakes to house nine, 20 gal (75.7 liter) aquaria and the flow-through apparatus (Figure 2). The roof was green plastic and the sides of the sheds were plywood with aluminum screening. Black plastic was secured over areas of screen on the sides and door to minimize variations of incoming light. These structures prevented disruption of the experimental system by predators, high winds and rain. The trans- lucent roofs and black plastic coverings reduced illumination, so fluctuations in temperature and community metabolism were less extreme than would have occurred in systems not housed in sheds. A schematic diagram of the experimental flow-through system located on Lakes One and Four is shown in Figure 3. Water inlet hoses from each lake were submerged at a depth of 0.5 m (maximum) and secured in place in the epilimnion with floats and weights. Lake water, drawn into the sheds with battery-operated pumps, passed through a 0.5 mm 211 m~-- --ea e~-e. ee-ee e_-e_ eee.e-m~=.e .L.~-mee.= eec.=-ece.= e~.e-ee.c m_e.=-mee.c ~xe.e-eme.c eL.N-ee.= cec.=-e;e.c Pm.e-ek.e Ne.e-_e.o. ae_.e-_ee.= ee.m-me._ eee.e-~_e.e ec.e.ee.c Nec.c-me.o ~.e_-e.e e.e-m.e =.=..e=.e e.c_-v.e a.c.-~.e Le.e-ee.= me._-se.c _~._-ee.o oo.m-me.e e~.=-ee.c .e.c-.=.e. x.._-m..e _=.o ee.c-hc.e _o.e. .c.e-_=.=. .e.m-,m.m .e.c-_o.o. ~_.m-~c.c ec.c-_o.c :_.c-.s.e ne.c.__.e ...c-_o.c. m~.m-~e.= c~.o-mc.o x._-~e _~_-.e _~_-oe mm“ L. me.-c~ e. e.e ~_-e m.--m.a_ mm cw m.em-e. _.e_.e.~ e.m.-e.e L.e.,e.e ~..~-e.. ~.~_-e.e - - - - =_-n - - - - e-e - - - - em-e. eo.-e= ne_-em. we-ce .m~-~e_ me-nm . _ e _ e ”:5; i-.|-.( t '01.! t.ll)1!|t.1llt.'|i mm:cu m~._;n—.c cm._1m~.c am._1c~.c ~.c_-~.o cc.m1_¢.c n_.m-o_.c m.c-e_.v Nw.—-No.c Cem1c—— m.ea-~. ~.m—1~.c— m_-~ mic mmsam om_-_~— 'l! 1.! Al 11-1 I. llll =§_¢c::=: m:_czawcza-::dzc mchcamcz; -c;.L: Aczov-ecvxz £Z§3£E;_2c— .... ccaosa.z c.:caeo .muCH eecoea.z 1 e._tu.z :cccgamz 1 muaga.z ccecL._z . a_=§==< :o_umL=acm :caxxc N Au. oczueecaseb cocxxc te>_cmm_= cancau 0.:emco page» :cccau uvcacgc eo>_=mm.= :cchu _agc— ha,:w_aa—< page» .9" II! .111 ”mux<4 "mhzu2_zuaxu 1.10."): 21'.0i 1.1:" I. ail ." 0551-11 .Azecoewmz Leda: be w.=~,am:_ ec=_:g.z Lo xgodczosed xcuw.§oguv mae_a=c can. an ne_asem c: ..xaav m.:e=o;:¢cxs .c mace; ecu azarzgzcc ms:_e> .c\o_ .Lecceuc-xex acgu gzab tza ac: acaea .c mu..m.ccuue5~:u .au.ecge-o._m>zg ._ c.5e— 255 63:39.. 3: 3:52.... - o.-- QM1~ ~1_ ~-_ ~._ cm-.. aacfinuz o_aagod__. .uuzp conlmmn o-pm_m c~n1mae c_o-w~o wxm-com :eo103m u:u_mux .cdcb —M1c~ @o-~m mm-o_ “Klan cm1- om-_s 23.9.8; ~=_1no_ mo_-ma Naiam s——-mm o51c~ ma—lmm av_ca_;u m.c-—.o v m.alm.o N.o-_.c N.o ~.o-—.c ~.o :3;— ncdunodv eo.ouno.ov mc.c-m=.cv vo.c-mc.c _~.o1mo.ov mo.o1¢o.o u:_~ oc.o-mo.o. cc.c1mo.cv ~3.cino.cv ~—.o-oo.c no.o1mo.ov oc.a1mc.o vmuzuaznx 1 1 1 1 cm ~m_1c__ ago._:m sm-~m ocp1mm ©c130 .mumo cc-~m c¢1n~ 5:.ucn m.m-~.o m—uc— —.n-m.m _—-m.m o.m-m.~ c_1¢.m azfinnouam v — v — v _ "muxm m—v—(H .N mczmwm 27 a / /‘ fl H Figure 3. Diagram of flow-through system: A is an inflow hose for lake water, 8 is a trough with nine outlet hoses, C is a delivery hose to D (an aquarium, 75.5 liters), E is a screened, removeable standpipe, F is a drain pipe to G (catchment tray), and H returned water to the lake. 28 screen that eliminated macroinvertebrates and large filamentous algae. The water then flowed into a specially designed trough (l x 0.5 x 0.5 m), fitted with nine outlet hoses (2.54 cm dia) with control valves to regulate the filling rates of the nine aquaria (30.5 x 73.7 x 31.7 cm). Each all-glass aquarium had one hole (2.45 cm) drilled in the bottom that was fitted with a PVC adapter and a removeable standpipe (15.24 cm = height). The standpipe maintained 42.8 liters of lake water in each aquarium. The screened standpipes allowed excess water to flow out, but prevented the escape of tadpoles. Tubing attached to each aquarium . adapter returned water to a catchment basin then to the lake via an outlet hose. Experimental Design Three experiments were conducted during the spring, summer and fall of 1976. The first experiment started on 27 May, the second began on 16 July, and the third began on 13 September. The final dates were 19 June, 22 August, and 24 October, respectively. Experiments at both lakes followed a 4 x 3 block design with four treatments and three replicates/treatment. Lake water was sampled and characterized before introduction into the nine aquaria housed in each shed. In all experi- ments, these initial samples of lake water represented one of the four treatments. In the first two experiments, the nine aquaria provided for three treatments: three control aquaria contained no tadpoles, three aquaria stocked with a low density of tadpoles, and three aquaria stocked with a high density of tadpoles. One species, Rana pipiens, was used in the first two experiments. In Experiment Three, the treat- ments with three replicates were bullfrog (R, catesbeiana) tadpoles, 29 green frog (3. clamitans) tadpoles, andaicombination of bullfrog and green frog tadpoles. The tadpoles were placed in the respective aquaria on 30 May in Experiment One, 28 July in Experiment Two, and 27 September in Experiment Three. In the first two experiments, water from the lake was introduced and removed from aquaria daily (2410,,while in the third experiment, water remained in aquaria for 4811before aquaria were drained and refilled. Each morning, prior to removing water that had been in aquaria for 24 h or 48 h, oxygen (mg/l) and temperature (C) readings were taken in each aquarium. Additional oxygen and temperature measurements were made immediatebyafter aquaria had been filled with water from the lakes to determineii’initial differences among aquaria existed. Oxygen concentrations were determined with a YSI (Yellowsprings Instrument Company) oxygen meter, calibrated daily with Winkler analyses. Daily extremes in temperature were obtained from minimum-maximum thermometers in aquaria. Tadpole Procedure Eggs of the northern leopard frog (Rana pjpiens pipiens) were purchased from the Amphibian Facility, Ann Arbor, Michigan for Experi- ments One and Two. All clutches of eggs were thoroughly mixed, then placed in enamel pans and allowed to hatch at room temperature. Upon hatching, leopard frog tadpoles were redistributed and maintained in aquaria with aerated, dechlorinated water and fed TetraMin fish food (44.1% carbon, 7.7% nitrogen) until used in field experiments. Bull- frog and green frog tadpoles used as test animals in Experiment Three were collected from ponds at the Wolf Lake Fish Hatchery, near Grand liaven, Michigan. These larvae were kept in plastic pools (1.5 m) in 30 the laboratory and fed TetraMin fish food before they were tested in the field. Prior to each experiment, individual tadpoles were measured (snout-vent length, mm), weighed wet to the nearest 0.01 mg, and their stage of development was recorded (Gosner 1960). Gravimetric deter- minations were made by quickly blotting individual larvae with filter paper to remove excess moisture, then transferring them to tared aluminum pans with water for weighing. This technique was chosen since it combined accuracy with a minimum of stress to the animals. Tadpole numbers were the same and biomasses similar within a particular treat- ment, experiment, and lake. Sorted tadpoles were held for about 12 h; dead animals were replaced. Tadpoles were acclimated to the field temperatures before being placed in the reSpective treatment units. In all experiments, treatments with different densities and biomasses were used to assess the effects of grazing intensity on nutrient concentrations, community metabolism and algae. Stocking densities and biomasses of tadpoles, in conjunction with different water qualities, were also used to evaluate intra- and interspecific competition in tadpoles, as measured by survival, growth, and produc- tion. In Experiment One in both lakes, each low density treatment was stocked with 20 tadpoles/aquarium and each high density treatment had 40 tadpoles/aquarium (see Table 5 for biomass data). In Experiment Two, the low and high density treatments had dissimilar numbers and biomasses at the two lakes. At Lake One, there were 15 tadpoles/ aquarium in the low density replicates and 45 tadpoles/aquarium in the high density. At Lake Four in Experiment Two, 5 tadpoles were stocked in each low density aquarium and 15 were stocked in each high density 31 treatment (biomass data in Table 5). In Experiment Three at Lake One, there were three replicates/treatment: bullfrogs at lO/aquarium, green frogs at 20/aquarium, and 5 bullfrogs plus 10 green frogs/ aquarium. In the same experiment at Lake Four, there were 6 bullfrogs/ aquarium, 12 green frogs/aquarium, and in the mixed treatment, 3 bull- frogs and 6 green frogs/aquarium (see Table 8 for biomass information). At the conclusion of each experiment, length, weight and stage of development were determined and tadpoles were individually frozen for subsequent analyses. Linear regression analyses (log length-log wet weight) were applied on both the initial and final populations of tadpoles. Regressions were calculated for pooled samples and for individual replicates. The lengths of tadpoles that died during each experiment were put into appropriate initial and final regression equations; weight at time of death was estimated from these equations. Since dead tadpoles had been collected daily, weights could be adjusted to take into account the actual date that tadpoles died. A minimum and maximum value for dead biomass was calculated and used in estimating tadpole production. Dry Weight and C-N Analysis of Tadpoles A representative sample of 130 tadpoles, selected from the three experiments and the original TetraMin-fed laboratory animals, was oven-dried (60 C) according to the methods in Cummins and Wuycheck (1971). Tadpoles were defrosted and individually reweighed, since there were slight variations between frozen and prefrozen weights. Each larva was placed in a preweighed aluminum pan (previously ignited at 500 C for one h) and weighed to the nearest 0.01 mg. A series of 32 25 tadpoles were oven-dried for 24, 48, 72 and 96 h and weighed after each time interval to determine the appropriate drying time. Tadpoles were oven—dried for 48 h, then cooled in a desiccator with P205 for 24 h and weighed. This information was analyzed to determine the relationship between wet and dry weights of tadpoles. Regardless of species or treatment, there was a significant correlation (r = 0.98) between wet and dry weights (Figure 4). Seventy-five oven-dried tadpoles, selected from the three experiments, were thoroughly ground with a mortar and pestle. Duplicate analyses of organic carbon and nitrogen content of individual tadpoles were conducted in a Perkin-Elmer Elemental Analyzer (Model 240). The variation in the carbon and nitrogen content of tadpoles within a treatment was not greater than that among treatments in both lakes within a particular experiment. Twenty-two tadpoles were analyzed from Experiment One, 20 from Experiment Two, and 33 from Experiment Three. Statistical comparisons (t_tests) were performed on the mean carbon and nitrogen content in tadpoles from each of the three experiments. The percentages of carbon and nitrogen measured in samples of tadpoles from the three experiments are listed in Table 2. Although the ratio of carbon to nitrogen was similar in tadpoles from the three experi- ments, values for carbon and nitrogen were significantly different among all three experiments. Limnological Assays Dissolved oxygen and temperature: During each of the three experimental periods, four 24-h (or 48-h) measurements of dissolved oxygen and temperature were made at about weekly intervals. Dissolved 33 .mceawswpu .m use ocmwmnmmumu .m .mcmwapn mama Lo mm—oacmu Lee .3 Ego; is 3 8 a. o.. h p p - nwzmcowuerL pcmwmz Acuupmz . .nood . .o.o .nQo 3.6 126 inxwgu (01111616741 Mo unmgo wand 1010 e ......o .e mg:m_m 34 .”~.m Apo.m by e.__ Ame.N he m.Ne mm Ameee_5epe.um use mcmwmammucu .mv mwcsh ucwe_gmaxw Pum.m A—m.o Av m.o~ Amm.~ my o.mm om Amcmwmwm amv ozp ucwa_cwaxu .uo.e Aem._ av A.m Aem.e “V e.mm NN . .mee_m_m .mO mco acmepgqum upowm F a.e Am~.o av w.w Amm.~ av ..me a e_zeeeee Pno.m .mo.p AV p.o «mm.~ Av m.om mp couxcwpnouxga Amcmwmwm acumv NgoucgoaeA zuo zuocmhuz a zoma0.05) were obtained from all three magnifications. The results were converted and expressed as number of phytoplankton cells/m1. Volumetric determinations were made by measuring the dimensions of algal cells (generally 20 individual cells in each size category) throughout the sampling period with a Whipple micrometer (Nalewajko 1966, Bellinger 1974). Formulae for geometric solids, most closely resembling shapes of phytoplankton, were used to detennine the volume (um3) for individual cells, colonies, or filaments. The proportions of various cell sizes in each genus for each sampling date were deter- mined and average generic volume was estimated. One-way and two-way ANOVAS were used to determine significant differences among treatments and Student-Newman-Keul's test was employed to separate the means for numbers and volumes of algal cells in each treatment. Shannon-Wiener Diversity Index (H'), its variance (var H1), and the evenness (J') (Pielou 1966a, b) were calculated with a FORTRAN computer program. Hierarchical diversity was estimated by splitting H' into components that were additive (Pielou 1974). By this method, the generic diversity within each algal class, the class diversity, phylum diversity, and total diversity were assessed for both numbers and volumes of phytoplankton. Measurements of hierarchial diversity were calculated for individual aquaria on each sampling date and on 38 pooled treatments for each date, experiment, and lake. Carbon, nitrogen, phosphorus, and alkalinity analyses: Nitrogen and phosphorus analyses were performed on a Technicon Auto- analyzer (USEPA 1974) by the Institute of Water Research's (IWR) chemistry laboratory. Determinations of total phosphorus, ammonia- nitrogen, nitrite-nitrogen, nitrate-nitrogen and total Kjeldahl nitrogen were made on unfiltered water samples; dissolved phosphorus and Kjeldahl nitrogen were analyzed in filtered samples. Total and dissolved organic carbon concentrations were measured on a Beckman Single-Channel Carbon Analyzer as described by USEPA (1974). All samples were stored in the refrigerator prior to analyses and examined within 72 h of collection. Concentrations of particulate phosphorus and particulate organic carbon were obtained by subtracting the dissolved fraction from the total phosphorus and organic carbon. Total inorganic nitrogen was calculated by adding the concentrations of amnonia-nitrogen, nitrate-nitrogen and nitrite-nitrogen. Amnonia- nitrogen was subtracted from Kjeldahl nitrogen for organic nitrogen detenninations and dissolved organic nitrogen was subtracted from the total organic nitrogen to obtain the particulate organic nitrogen. Hydrogen ion concentrations were measured with a Corning Glass electrode pH meter; carbonate, bicarbonate, and total alkalinity were determined by acid titration (APHA 1971). Analysis of periphytic biomass and C-N content: A total surface area of 6,212.9 cm2 was available in each aquarium for periphytic growth. To monitor the production of periphyton, nine glass slides (5.1 x 7.6 cm) were attached to plastic strips and hung 39 on each side of an aquarium and eight slides were fixed on plexiglass plates on the bottom of each aquarium. A total of 44 slides were placed in each aquarium at the beginning of an experimental period. During the experiment, eleven slides (426.4 cmz) were removed for: algal identification, oven-drying, ashing, and C-N analysis at the end of each 24-h (48-h) dissolved oxygen and temperature profile. At the end of each experiment, the plexiglass plates (232.3 cmz) were collected and periphytic material was oven-dried and ashed. Slides for identification were stored in 10% buffered formalin, while slides used for determinations of periphytic biomass and C-N content were placed in plastic bags, air-dried, and stored in a desiccator with P205. Periphytic material was scraped from the surface of slides and placed in preweighed, aluminum pans (previously ignited in a muffle furnace at 500 C for one h). Periphyton was oven-dried (60 C for 24 h), desiccated (P205) for 24 h, and weighed to the nearest 0.01 mg. This oven-dried material was ignited in a muffle furnace, moistened with distilled water, oven-dried to a constant weight, desiccated, and reweighed. The oven-drying and ashing procedures were similar to those described in APHA (1971), Currmins and Wuycheck (1971), and Vollenweider (1974). Periphyton used for carbon and nitrogen analyses was oven-dried, weighed, and thoroughly ground with a mortar and pestle. The organic carbon and nitrogen content was measured in a Perkin-Elmer Elemental Analyzer. Estimates for accrual (mg/cmz/d) of periphyton were corrected to account for more extensive growth on the bottom of aquaria. The sampling procedure resulted in the bottom representing 20% and each side representing 20% (4 sides = 80%) of the total periphyton on each 40 date. Actually, the sides comprised 64% and the bottom of the aquarium was 36% of the total surface area. Information on the periphytic material from the plexiglass plates was used as a cor- rection factor. Relative contributions of periphyton and phytoplankton to GPP and_3; To distinguish between the separate contributions of periphyton and phytoplankton to the total oxygen production (GPP) and conmunity respiration (R) in aquaria, anCillary studies were conducted during the second and third experiments. On two different dates in Experiment Two and one date in Experiment Three, eight glass slides attached to plastic strips were placed on the bottom of each aquarium. The time allowed for periphytic growth on slides was 12 days in both separate tests in Experiment Two and 16 days in Experiment Three. At the end of each period allowed for periphytic growth, 18 wide-mouthed jars (0.95 liter) were filled with water from the appropriate lake. Glass slides were removed from each aquarium and one set of eight slides was placed in a jar filled with lake water (total of 18 jars with slides). These jars and jars with lake water only were capped; two jars, one with and one without slides, were placed in similar positions in each aquarium. Dissolved oxygen and temperature were measured in each jar (7-8 August, 21-22 August, and 22-24 October) concurrent with measurements on water in aquaria. The jars containing only lake water represented the contribution of phytoplankton (seston) to gross primary productivity and respiration, while the other jars with suspended glass slides were used to calculate the contribution of periphyton. 41 At the end of 24 (48) h, ten ml of formaldehyde were added to each jar. Attached material scraped from slides was returned to the jar. A known volume was filtered through a glass fiber filter and placed in an aluminum pan (both filter and pan were ignited in a muffle furnace and weighed). Sestonic material from lake water was filtered and all samples were oven-dried and ashed. Three replicates (300 ml) were filtered from jars with combined periphyton and phytoplankton. Calculations of gross primary productivity and total respiration, made on those jars containing seston only, were sub- tracted from the combined seston plus periphyton to determine the contribution of periphyton alone. Dry weight and ash-free weight of sestonic material were subtracted from combined measurements to obtain the dry and ash-free weight of periphyton. Sestonic values were corrected and converted to carbon, estimated as 50% of the ash-free dry weight (Vollenweider 1974). Amounts of carbon in periphyton were estimated from C-N analyses done previously. The Laboratory Experiment Natural Food Sources, Algae Tadpole growth, survival, and efficiency of conversion and assimilation of algae (analyzed as particulate organic carbon and Kjeldahl nitrogen) were assessed for different-size tadpoles of Rana pipiens fed three different concentrations of natural food, phytoplankton. Fifteen aquaria, representing five treatments of tadpole density with three replicates per treatment, received similar total biomasses but different total numbers of tadpoles. Initially, the amount of food and space per total tadpole biomass was similar 42 in all aquaria. The effects of reducing food and space per individual tadpole, simulated by the density treatments (increasing the number of tadpoles per aquarium), was evaluated in this laboratory experiment. Six weeks before the initiation of the feeding experiments, three containers (305 liters) were filled with equal portions of distilled water and tap water and enriched with modified Chu‘s Culture Medium (Chu 1942). Containers were stocked with either a mixture of phytoplankton from Lake One of the Water Quality Manage- ment Project, or Scenedesmus, or Pediastrum; pure strains of Scenedesmus and Pediastrum were obtained from the Indiana Culture Facility, Bloomington, Indiana. Cultures were illuminated 24 h/d and continually aerated. Phytoplankton were enumerated biweekly and Kjeldahl nitrogen, phosphorus, and organic carbon were measured weekly on filtered and unfiltered samples taken from the three cultures. Additional water and nutrient media were added three times per week to the cultures to maintain an abundance of phytoplanktonic growth. Mixed clutches of Rana pipiens eggs (Amphibian Facility, Ann Arbor, Michigan) were held either at 20-21.5 C or 10 C. Each day, some of the eggs developing at the cooler temperature were removed and placed at room temperature (20-21.5 C) to obtain populations of different-sized tadpoles. Individual tadpoles were measured (snout- vent length, 11111), weighed wet to the nearest 0.01 mg, and staged (Gosner 1960). The mean number of tadpoles in the 5 experimental densities was 9, 4, 3, 2, and l with a range of 70l.0-7l4.4 mg for initial total biomass per aquarium. All 18 aquaria (20.8 liters) were filled daily with 3.8 liters of aerated water, cleaned at the end of 24 h, and refilled 43 with aerated water. Tadpoles were supplied with different levels of algal food during the 24-day experiment. Initially, 500 ml of mixed algae were added to each aquarium and this level of feeding was continued for 14 days. The next higher addition of food material, 1000 m1 of mixed algae per aquarium, was provided for 6 days. During the final 4 days of the experiment, tadpoles were fed 1,500 ml of mixed algae per day. The food mixture of algae was obtained by combining predetermined amounts from each of the three cultures. At the lowest feeding level, 250 ml were taken from each algal culture. These samples were thoroughly mixed, then 500 ml were added to each aquarium. At the higher feeding rate, 500 ml were removed from each algal culture; these samples were mixed and 1,000 ml were added to aquaria. The highest food concentration, 1,500 ml per aquarium per day, was achieved by removing 600 ml from each algal culture and combining these samples before adding the prescribed amount of material to aquaria. To determine initial concentrations and avail- ability of nutrients provided at each feeding level, analyses of total and dissolved Kjeldahl nitrogen, organic carbon, phosphorus, and phytoplankton density were made on the remainders of the mixtures from the three algal cultures (Table 3). Water in each aquarium was stirred and then sampled at the end of each 24-h feeding period. Water sampled from stocked aquaria was passed through a 0.5 mm screen to remove tadpole feces and con- centrations of carbon, nitrogen, and phosphorus were analyzed. Un- stocked aquaria were also sampled and the changes in phytoplankton, organic carbon, nitrogen, and phosphorus that occurred over the 24-h period were noted. All tadpoles were weighed individually every 48 h zl4 .MAmemwmxw .memmgmaw p Aua.v-o.o~ maouo_c u_cu=uu mmmem.waw-. .m_a:o_a.a .a.:um~a.z .mmmmiwmm - Auc.v.-o.o. nsouawu ounczma m_—mu coacm u_ouuou .mmemwmbaww .mmdmammmw .amgwmmzewmm awwawwmmhww_mm( 1 Afio.:v _ mam—a :mogm conga .Aum.~m-o.n_. recumMWmmm .Axa.mm-m.o_. wmzmac:mmw - mam—a goose "a.Laaco c. vocoucaooco axe“ Lo meo.~coaoca mmaeo>u ==a c;u:ac_ en~.o- a.o “.8. c¢.~ m_.n om.~ we.” s.e\ae \xs.__ee__e>< A; An aozmov mm-cm. mm..¢m=._ «.mn m.o. m_.m __.~_ mo.~_ .m.n_ e . see\e. \9. .agap as .3 can. .... eme.e_~ e.e ..a ee._ .e._ mm.. oe.~ x.e\:e \s.__.ee_.e>< A: Av mucous -_-o~_v .Lm.m~m ~.e~ 3.8m mm.m SN.“ ve.e ~m.m u e see\ee \ma _odop as . .3 coo. .__ m_m.a¢ o.m a.m ee.o no.— m~._ ”8.. .ee\ce \»s___ee_.e,< A; an maggot . ua~-oamv em“ mam o.o. ..NN -.n mc.o .o.. -.e e .— >.e\ee \9. .auop Au .3 can ._ us_~ m—m>oa cue—so use such ouopau use _ouo» . can :u wmmn. ago» an ac.aoau caused—acuxsm coagau oven co wage: was; : asu.z pane—o x co.uaL:a _ ..x.es.eeeemee .nx.a_ wee ~.a ...n. ...N. new: ova—e gen .9 .a .2 Lee >9 axeaee_. Laue: .6 we»... a.« eu_: =o_.s._= en..e mu_oauou cu xu__.ns_.c>o L_agu s:e «each x_;;:m omega 3:“ an .:\.a\m__ouv =oux=cpaouxsa can Au\aiv mu:o_cu:= La neo.ucca=ou=oU oaacu>< .m u.;a~ 45 during the 24-d experiment. At the conclusion of this experiment, tadpoles were individually frozen after being weighed, measured, and staged. Determinations of dry weight and carbon-nitrogen content were done subsequently as described in the field experiment section. A Comparative Food, TetraMin Ten leopard frog tadpoles, encompassing the range of tadpole sizes employed in the algal feeding experiments, were selected from the same laboratory population of tadpoles. Each tadpole was placed in a plastic container (11.2 x 11.2 x 5.1 cm) with 500 ml of aerated water. Tadpoles were fed TetraMin, a highly nutritious food, for 14 days, concurrent with the last part of the previous laboratory experiment. Containers were cleaned daily and resupplied with water and TetraMin. TetraMin was provided in excess to tadpole daily requirements to assure maximum individual growth on this food source. Tadpoles were weighed every 48 h at the same time as algal-fed tad- poles and at the end of the experiment were processed like algal-fed tadpoles. This ancillary experiment provided information on indi- vidual growth rates of tadpoles of different sizes and stages to con- trast with results from the previous laboratory experiment when tadpoles were fed a natural food, algae. RESULTS Environmental Factors Differential Effects Tadpole exposure to the different physical and chemical characteristics of the lakes could have influenced the comparability of results among experiments. Environmental characteristics of Lake One and Lake Four are tabulated in Table l of the Materials and Methods section. These factors, along with temperature, oxygen, alkalinity, and pH measured in aquaria and described below, indicate the types and degrees of stress that tadpoles reared in water from the two lakes may have experienced. Daily Measurements of Temperature and Oxygen Within an experiment, temperatures usually varied less than 1 C among aquaria at both lakes (Figures 5 and 6). Therefore, tad- poles were exposed to similar thermal regimes within experiments at Lake One and Lake Four. The daily range in minimum-maximum temperature was 5-11 C in Experiment One and 4-15 0 in Experiment Two. In the first experiment, the mean daily temperatures increased throughout the duration of the experiment in response to increasing day length and average air temperatures (Figures 5 and 6). In Experiment Two, sunmer storms and intermittent cloudiness caused erratic variation in water temperatures (Figure 5). Mean daily temperatures declined in late:summer during the third experiment (Figure 6). The highest 46 47 EXP. LAKE! I 1"70 ExPI LAKE4 ”"0 9' «woo : so 2 4b 8 4b” 5 45"1 0 ’0 g < f e :80: URN U C )- 8!» . é " § '1' 2 a 70 on +17 4 IS a ' a {’2‘ ' 4,” a V 1P2: g 7’2. 3 ’ ’ hzo ’ ‘ a 1’" a a! 0'. M I"? a 010 MS 1 I‘ a 0': 3 4» '2 ”U. 1 '0 4» Q 2 V Y V V f Vi T 7 V B 2 v V T r v v v r v v ; Juno June «In vi ‘0 120 0040 :1 EXPZ LAKE 4 1'20 2 11100 g .00 5 o ll-i 0.0 8 04° .. 20 “’1 DISSOLVED OXYGEN (Mill 0 7: oi ) i ll ’27 026 025 ”a. ..z: 3 022 £ 021 3 (’20 ’ >19 5 "1. or! g "'9 0'5 3 “l4 ('3 .. 412 o" 4).” A A A - k A A A A A A L A A_ L A L A A 9* A_ A A A A A A LL A L44 1 A A A A A A A A_A TV 1 ‘7 W I Y I’ r V T V V V V r V V ' fiv f v V Y 7 I v t V v v v Y + r 1' j T f fi 25 3| 21 25 3. a 4.1, About Joly August Figure 5. Range of relative oxygen saturation, mean dissolved oxygen, and temperature for Experiments One and Two at Lakes One and Four (for dissolved oxy en, unstocked aquaria = -——3 low density = ---, and high density = °" . NOLLVURLVS i O. JUMVINIJA 48 .mmcpummg No ummgmws ucm pmmzoF Easy nmumwmm . - oz mucmsogzm . a 3 mm emu emu m gom avgmacm cw mm: swam: quwm z< op m comzamn mums mg .A.mwuummmoumpwmmgw new .--- u muwwwamncwxwe .111." mmoge_pza .cmmAxo vm>pomm_c Lomv Lao; umm meo.mmwmumwm «mesh acoswgmaxu Lam mgzumgmasmu ecu .cmmAxo um>Pomm_u cams .:o_umgaumm :mmxxo w>wum_mg we a cam o . .3300 34:32.5 35290 33.5....» o. N“ 3 u. N“ b b D P D b P P Sb h I 0 0 l“ 10 g. 353 nmxw o .r .93.. mmxu mi. N t \r 0.1 m t 4w c A N‘ ~11 o r E f n a: m oafi ..UI ...: M 0.. n r R 9.? ...... N.. E M . . w “T? E v # n 2: o_. m-:.- 0.11 1.- 8.:- .~ 410 m fa 0 L5 3 t: a 0 0 w tn. 1A.. 48. a :0. u as.0 ..m m 1' AV“. 9 u m ‘1 ..—N \' u A§.u Mao»: m m 0” I] o, m | T I 7.0a: ( M 00 IV L ( M . U h. w o. : m on : a on... 9. 8-1Y % 3.1: on. . 49 temperatures for Experiments One, Two, and Three were 28, 27, and 2l C and the lowest temperatures were 10, ll and O C, respectively. The mean daily temperature in Experiments One and Two was about 17 C, while in the third experiment, the temperature averaged 8 C. As with temperature, similar patterns of daily fluctuations in oxygen concentrations occurred among aquaria at the two lakes (Figures 5 and 6). However, oxygen concentrations frequently differed among the three treatments at Lake One, presumably because of differences in community metabolism since reaeration coefficients should haVe been the same. In Experiment One at Lake One, the un- stocked aquaria consistently had greater oxygen concentrations than aquaria with tadpoles, but oxygen was always above 8.8 mg/l in all aquaria (Figure 5). Throughout most of this experiment, the relative oxygen saturation exceeded lOO%. Since oxygen was sampled early in the day, the peak concentrations were probably higher than measured and all aquaria were supersaturated with oxygen throughout the experiment. In Experiment Two, measurements of daily oxygen fluc- tuated, but all treatments remained within l mg/l of each other at Lake One until the final week in August (Figure 5). At this time, oxygen dropped sharply in unstocked aquaria to about 2 mg/l or 20- 30% saturation. Oxygen levels also decreased in aquaria stocked with tadpoles, but remained at 60-80% saturation. In Experiment Three at Lake One, concentrations of dissolved oxygen were similar among treatments, invariably within 1 mg/l of each other (Figure 6). On only one date did oxygen decline slightly below 9 mg/l or 90% saturation. At Lake Four, little variation occurred among treatment 50 aquaria within an experiment; oxygen levels were within 0.5 mg/l of each other (Figures 5 and 6). The changes in dissolved oxygen measured within an experimental period were considerably less than at Lake One. The lowest concentrations of oxygen recorded for Experiments One, Two, and Three were 8.0, 6.9, and 8.3 mg/l; only in Experiment Three did the highest oxygen level exceed l2 mg/l. Relative saturation of oxygen remained above 60% and was generally 80-100%. Extremes of Dissolved Oxygen Unlike temperature, diurnal extremes in oxygen concentrations and percent saturation in aquaria were different at Lakes One and Four. Minimum-maximum oxygen concentrations and percent saturation, based on 24-h and 48-h analyses on four representative dates during the three experiments at Lake One and Lake Four, are provided in Figures 7-9. Oxygen concentrations in all aquaria at a lake were similar on the first sampling date in each experiment, before tad— poles were introduced. In the first two experiments, particularly at Lake One, the diurnal variation and differences among treatments increased through time. The growth of periphytic communities and tadpole feeding activities most likely influenced this increase in diurnal variation. In Experiment Three, extremes in diurnal variation of oxygen decreased with time and may have been related to declining air temperatures. At Lake One, oxygen varied 7 mg/l in aquaria stocked with tadpoles in Experiment One, 14 mg/l in Experiment Two, and 9.5 mg/l in Experiment Three (Figures 7 and 9). In contrast, the maximum change in diurnal oxygen for stocked aquaria at Lake Four was only 2.3 mg/l in Experiment One, 2.4 mg/l in Experiment Two, 51 20% DI SSOLVED OXYGEN (mg/l) EX? 2 LAKE 4 LAKE 4 l l v A I . C l 2 3 4 I. 2 (”up A. MIN-MAX MEASUREMENTS (24h) Figure 7. High and low concentrations of oxygen during four, 24-h periods in Experiments One and Two at Lakes One and Four (unstocked aquaria =-———, low density = ---, and high density = ~--). 52 LAKEI LAKEI 'lo SATURATION so LAKE 4 LAKE 4 r l 2 3 3' I 2 3 4 d d d SAMPLING DATES (24 Hr) Figure 8. High and low relative saturations of oxygen during four, 24-h periods in Experiments One and Two at Lakes One and Four (un- stocked aquaria = -——3 low density = ---, and high density = ---). % SATURATION DISSOLVED OXYGEN (mg/I) 53 24. EXPB LAKEI 13 . I; . I, I: I: I. I: 'I II '1 I. :3 I4 fi axes LAKE4 I? I. . I; I I: k I20 :2 1.. E .. :: fl I. . I. ‘- :2 I: .3 I3 I00 I: ~ ‘6 1: '1 b_: , . ‘8 .. e 8" 8" I I r I I I I r I 1 w ‘ 240 1 EXR 3 LAKE I zzo< 1§Wg I: I: . I: 2 200- 5 E I: : u I. I '5 . r709 '°°‘ : I I/ 5 5x23 LAKE4 . 2 : *“3 I40- ISO' l C 1 '20.. .‘0 I40‘ » j :2 2 ' I: I20- . l: _ 3 I000 I 1 I: Ioo- : q_ i . 3 IE . :: w. 80 I- 3 H T \ l6 . 5 I I I I I I I I ‘r I I I I I I 2 3 4 I 2 3 4 MIN - MAX MEASUREMENTS (24h) Figure 9. High and low concentrations and relative saturations of oxygen during four, 48-h periods in Experiment Three at Lakes One and Four (bullfrogs =-———, mixed-species = ~--, and green frogs = ---). 54 and 3.0 mg/l in Experiment Three (Figures 7 and 9). The daily extremes in relative oxygen saturation also differed at the two lakes (Figures 8 and 9). At Lake One, the aquaria with tadpoles were always at least 90% saturated and peak saturation was greater than 200% in Experiment One (Figure 8). In Experiment Two at Lake One, daily extremes in oxygen saturation ranged from 60 to 240% in aquaria stocked with tadpoles, while in Experiment Three, the extreme range was 75 to 185% (Figure 9). At Lake Four, the maximum daily range in percent saturation was about 80-120% in Experiment One, 55-85% in Experiment Two, and 85-135% in Experiment Three. Only during the first experiment at Lake Four did relative saturation of oxygen remain above 100% for part of the day throughout the experimental period (Figure 8). Alkalinity and pH Throughout this study, both lakes were moderately alkaline (Table 1). In Experiment Three, the total alkalinity in aquaria at Lake Four averaged 68% of values at Lake One (Table 4). The pH range in aquaria at Lake One was 8.9 to 9.8, while at Lake Four the range was 9.3 to 10.0. Since measurements were made during morning hours when some carbon dioxide from nighttime respiratory activities may have remained in solution, the peak pH may not have been attained until after CO2 was exhausted. The extent of diurnal change in pH caused by photosynthetic uptake of carbon dioxide was about one pH unit, based on the decline of oxygen fluctuations in Experiment Three. Some of the variation between lake and aquarium readings, especially at Lake Four, may have resulted from differences in sampling times. 55 c.o_ o.o_ o.c_ ..o. a.w w.w o.w c.m :; Av m.~m_ m.cn_ ~.m~. m.mn_ o.aep m.mo_ m.wc. o.~o_ .ouop Au ~.cm o.am m.no o.mm m.mc_ c.xn_ m.cv_ e.ee_ ouozoacoofia Ag ~.n~ ~.c~ ~.mo e.om m.m~ —:m~ m.m~ ~.m~ ouocoaccu Ac aka. concuuo e~-- Ac 3.. as 3.. ...: m4. 3.. 5a 3.. .... 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A“.._e-~.ooe. .m.>_m-m.n>mE AwaeIc. >._~m >.m~o . ..nam a.c_q m.c~. m.-. o.oom o.mme .5: _I=s_>_u=_ m Ana—m-camm> Am~_o~-am¢>_. A>mme-_am~> A¢N~8->NQ>. .mmuq.-cm___. .moae_-o~mo_. Ammo..-mccm. .>8~__-n8q>. AIaeoc. mcm> memo. comm comm ee-_ came. «awe. anew cI\III:a>> am I .z c~_ I oz aw I .z so I oz .m I .2 CN. I oz cm I .2 co I oz IIIs=_m m_>_~ «Imam cav__ ammIN mawam .maoom mmecm >>>m~ .I.o> mco m:9=_gmuxw _I=.L .I.u.=_ _I=_L .I.¢_:_ .I:_L .I_a_=_ _I=_L .._5_=_ m>2ux_¢uaxu >a_m=oo sa_= >>.m:ma zed ma_m=oo saw: »u_mcoo so; «sad ux magma no orb u=o ace w>:u§_cy;xu :_ mu_=;=ud ace. wcsacu_ ‘c .==_.~:z>.~=c=m~ m=ga=2_ ==a .A2= .HgaLQ: “a: .LJOL 2:: 0:6 V mmmwesama .angan:uc .m w~an~ 59 in Experiment Two had a slightly lower probability of survival than those in Experiment One. Environmental factors, measured during the two experiments, appeared similar so major influences by these factors could be discounted. The information on tadpole mortality summarized in Table 6 indicated that dead tadpoles in both density treatments had larger average sizes than tadpoles which survived to the end of Experiment One (Table 5). In Experiment Two, the average weight of surviving tadpoles was higher than that for dead tadpoles, but the average weight of dead tadpoles exceeded the mean initial weight. In both experiments, the coefficients of variation of weight were larger among dead tadpoles than among initial or final weights of living tadpoles, indicating that no one size group died at a consistently higher rate. In the first experiment, both large and small tadpoles appeared to have a higher mortality than medium-size tadpoles, based on the weight distributions (Appendix B, Figures Bl and 32), R2 values for length- weight regressions (Table 6), and average weights of dead tadpoles. The weights of tadpoles were more normally distributed among survivors than among those initially stocked (Appendix B, Figures Bl and B2). The relationship between length and weight (R2) for tadpoles increased in both experiments, suggesting that misfits died or the condition factor increased similarly among survivors. In the second experiment, final weight distributions were more skewed than initially (Appendix 8, Figures 83 and B4); however, the relationship between length and weight, indicated that smaller tadpoles were not thinner or in poorer condition than larger tadpoles. Overall, no consistent relationship emerged between tadpole size and survival, when the results from both 60 em.~ u>.~ >>._ um.. eo.mmacaem >8 mo.o m~.c m~.c ma.c N: xmmmw.o I a>o~._ I > x>>_m.c I 0¢o~._ I > xmomm.c I me>~._ I > chem.c I mom~._ I > Auea.oz was - gauged ass. :o_mmocoo¢ _e=_u no.~ no.~ N_.N ”8.. eo.mmocasz >8 .m.c ~m.a oN.o No.0 Na xmmm~.o I mNmN._ I > xsao~.o I aev~._ I > xmoem.c I m>>~.. 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N.NCm Hag e.~¢c_ m.oom_ N.ch~ :c\=o_du=cob; mmm.mxmm mack; coo;c moogu coogc a v::;...:: maogb__:m Hu¢x_z uwzzp hzu=_¢uaxw mzfiuzam-nox_a vac Amw_aauau ac .Ac cv;awc c.5cK .c NN. ou;;— .:;E_gc;x ;‘ =orgo v:c cog5__:av uc_um;m-o_==_m go. A: w :. mm;=._:o A_. 33:.._= c=e .=; __ .vccgb =egg: .9» , vac ~= .mmog _~:Q &c =°_~=Q_g~=.. :5 .32; :2 s zigzag 22...: .6 23 71 xee-.c . m-~._ v > xeemm.c + cko~._ n > - x\:c~.c * mum“. u » A~:a_ez so; - :da:o_ ca‘V :o_mmw;ao¢ _c¢..:_ xmmmm.o . Know._ u > m:¢.¢- Hug ..c.o- _a. e_~.o- c.o.o- m::.c- m\; NN.:- .c. Pm.c- _m. mm.ma no..- Nm.c- co\c:_uuatc;g cwaogu_z mm.c- Ho. _=.m- .m. mm.—_- mx.m- m_..- ca\:o_¢u:cogg ::;_eu oc.c- Ha. mm.o- _m. mm..- we.c- c_.:- v omgoou\:o\co_au:=ogg mn.c- ”c. N¢.~- Fa. cc.o_- om.m- co..- c\co\=o_uu=1ogg m.o.- gs. c.-- _m~ ..cmw- m._¢- m.:m- aa\:o_au:vog; g:ou oxmw um._ ~¢H No.— .aw Hm._ - M... co.mwmgamz >u mw.= HcH mo.c Hm. Km.o - mo.: m: xumcm.c + coxm.. u xmpmm.o + mc-.. memm.o ‘ o_-._ p > - x_c:a.c . comm._ " > Aaza.o3 mod - =.a=od mo‘v copmmmgamz —a:.u ll >->- maogu gouge woos; gouge w uccgb__:c mmo;~__:= ”vox_z 4 is: a is 72 $5 7: u~.c ”a. No.— - u~._ :o,mmogav= >9 Ko.o .c. om.= .mH pm.c - -.c z N x.N_m.o ‘ NNNN._ " > mu. x~o_m.o . c.¢~.. " > HEB . xo~o~.o + cm-._ u > - xmg Aagm.e: no; - guccea mad. :c'mmmgocz _e:.m «c.o Hug u~.o ”a“ v~._ - x=._ cc_mmmgmum >u No.o new mo.c an O®.O I C9.= N: «93.; .395 map... .3059 a mocha—:5 mooL5——:m "vwx_2 . a 45.25; m 93 _ 73 At both lakes, survival was high in all aquaria, regardless At Lake One, survivorship of bullfrogs was l00%, of treatment. 98% of the green frogs survived, and in the mixture of the two species, At Lake 100% of the bullfrogs and 93% of the green frogs survived. Four, 94% of the bullfrogs, 94% of the green frogs, and in the mixed- species culture, 78% of the bullfrogs and 100% of the green frogs The apparently lower survival of bullfrogs in the mixed survived. culture at Lake Four probably reflected the smaller number present more than an interaction with green frogs, since only two individuals died. At Lake One, biomass/aquarium and mean weight per individual ‘i ncreased in all treatments, despite the deaths that occurred (Table The range in the distribution of weight classes for bullfrogs 8) . reared in single-species culture broadened, but appeared more normally (1‘? stributed than initially (Appendix B, Figure 88). In mixed-species cu 1 ture, the final weight distribution of green frogs reflected a general shift of individuals into weight classes greater than the mea n (Appendix B, Figure 89). The weight distribution of smaller bu 1 1 frogs changed little in the mixed-species treatment, while the weights of larger bullfrogs were spread more than initially (Appendix B, F ‘igure 89). Green frogs reared in monoculture exhibited more QVOWth for small to medium-sized tadpoles than in the larger weight C133 ses (Appendix 8, Figure 810). Coefficients of variation based on 1:1. ha 1 weights decreased below initial values for bullfrogs and green Y‘OQS in mixed-speCies culture and green frogs in smgle-spemes cu‘ture. This coefficient increased slightly for bullfrogs reared in monoculture. The final relationship (R ) for length and weight 0" tadpoles was higher than that measured initially for green frogs in 74 single- and mixed-species treatments and for monocultured bullfrogs, indicating a higher, more similar condition factor among these tad- poles. Values for R2, based on initial and final length-weight relationships of bullfrogs in mixed-species culture, remained high and similar. At Lake Four, the initial and final distributions of weight for monocultured bullfrog tadpoles were alike, except for one indi- \Iidual that grew significantly (Appendix 8, Figure 8ll). In mixed- species culture, smaller bullfrog tadpoles appeared to be adversely affected by green frog tadpoles, as shown by their weight losses (Appendix 8, Figure 8l2). The final distribution of weight classes for green frogs in mixed-species culture was similar to that measured initially (Appendix 8, Figure Bl2). Only for green frogs in mixed- species culture was the initial and final average weight of individuals 5 imilar, in all other treatments the final weights of tadpoles de- creased (Table 8). Also, the relationship between length and weight CR2) decreased in all tests except for green frogs in mixed-species treatments (Table 9). In single-species treatment, a general shift 01“ green frog tadpoles into smaller weight classes occurred, except for the increased growth of one tadpole (Appendix 8, Figure 813). At Lake One, productivity was highest for bullfrogs, inter- mEdi ate in the mixture of the two species, and lowest for green frogs i“ monoculture. Values for tadpole production per unit of tadpole biomass (TP/TB) reaffirmed this sequence of results; bullfrogs had the highest TP/TB ratios (0.084), the mixed-species treatment was i ntermediate (0.058), and green frogs exhibited the lowest production per unit of biomass (0.045). The TP/TB for bullfrogs reared in 75 monoculture was twice that of green frogs in single—species treat- ments. In mixed-species culture, the TP/TB of bullfrogs decreased to about half the TP/TB measured for bullfrogs in single-species culture and was the same as that for green frogs in monoculture. The green frogs in mixed-species culture had higher TP/TB values than those in single-species culture, but not as high as monocultured bullfrogs. Apparently, an interaction occurred between bullfrog and green frog tadpoles that benefited green frogs. At Lake Four, values for production/aquarium/day and production per unit of biomass were negative in all treatments, indicating tadpoles lost weight (Table 9). Although production estimates were negative, the trends were the same as those observed at Lake One. Monocultured bullfrogs were the most successful in that they lost the least weight, the mixed-species culture was intermediate in weight loss, and green frogs in monoculture lost the most weight. In mixed-species culture, bullfrogs lost more weight than when reared in monoculture. Green frogs from the mixed-species culture had similar TP/TB ratios to bullfrogs ‘frwmisingle-species treatments. As at Lake One, green frog tadpoles rwaared with bullfrogs seemingly benefited from this association, while the production of bullfrogs was substantially lowered. Reducing Food and Spacejer Individual (Laboratory Experiment) Algal concentrations, TetraMin, and tadpole density; In field EExperiments at Lakes One and Four, both density and biomass were VaV‘ied in Experiments One and Two, so determining the separate effects of these factors on tadpole growth and survival was difficult. Liiboratory studies were designed to investigate the relationship 76 between the density of tadpoles and their growth when the food and space per unit of total tadpole biomass were initially the same. The largest initial difference in total biomass/aquarium among the five density stockings was l3.4 mg or l0% (Table 10). Since little variation occurred among the three replicates at each density (largest coefficient of variation = 4%), statistically significant differences were measured among the density treatments before the experiment began, even though actual differences in biomass were minor. The densities of tadpoles (9, 4, 3, 2, or l per aquarium), when converted to number per m2 or m3, were moderately high (l3 to l20 tadpoles/m2 or 263 to 2,638 tadpoles/m3) compared to densities estimated in natural systems, but similar to those employed in the field experi- ments at Lakes One and Four. Because tadpole biomass was held constant while density was varied, the mean size and stage of tadpoles at lower densities were greater than for tadpoles at higher densities (Table l0). After each algal supply rate (500, l,OOO, and l,500 ml), statistically significant differences were found between the total tziomass at the highest stocking density (nine tadpoles, A-l in Table 10) and all other total biomasses of tadpoles at the lower densities. The lower total biomass (A-l) was partially caused by tadpole death early in the experiment, resulting in the average number of tadpoles ‘il1 this treatment to be eight for the remainder of the experiment (Table 10). However, the average difference in total biomass (mg i 55‘3') between A-l and the mean of all other density groups was l98.7 1*; 30.8 mg at 500 ml, 255.7 1 35.3 mg at 1,000 ml, and 450.7 _+_ 98.8 ““E! at.l,500 ml of algal food. The biomass of tadpoles in the A-l Table lO. Daily changes in mean weight, biomass, and production per unit of biomass for tadpoles (wet weight in mg, coefficient of variation of weight, booy length in mm. stage) during the algal feeding experiment (* cnange 77 leopard frog _ and developmental in weicnt per unit of tatal weight per day). A-1 A-Z 71-3 4-4 A-S Initial Conditions No/aq 9 4 3 2 1 “Mt 78.6 176.5 233.7 57.2 706.5 (:50) (:27.3) (:95.0) (17.3) (153.5) (:0.7) cv 35.3: 14.2: 3.13 16.4% 0.1.: Slag 707.8 706.0 701.0 71 4 706.5 (:50) (14.3) (13.3) (+1.0) (11.2) (10.7) BL 7.8 10.3 11.5 13.6 17.5 Stages (Range) 3-(24-25) 3- 23-26) 3- 25-27) :«(27-29) S-29 3 50C ml after l4 0 Ni/aq 8 4 3 2 l 711: 158.5 360.3 494.3 725.0 1501.0 '150) (:58 3) (52.1) '+58.3) (184.5) (59.6) cv 431: 20.0: 11.9: 11.7: 1.8: E/ag 1268.0 1443.0 1483.3 1450.0 1501.0 (:50) (:}.2) (110.4) (:5.0) (333.5) (:19.5) 07' Ht/d 5.7 12.4 l8.l 26.3 55.7 ea/ac/a' 40.0 50.1 54.3 52.5 56.8 P/B/d 0.057 0.076 0.080 0.074 0 080 (0.073)* @ lOOO ml after 6 d Nz/aq 8 4 3 2 l 31‘ wt 227.6 513.9 690.8 1024.5 2129.0 (:50) (£101.31 (1110.0) (1127.0) (307.4) (152.0) cv 44.79. 21.43 18.4“: 10.57. 2.4: 8/40 1820.8 2055.7 2072.3 2049.0 2129.0 (:50) (:98) (17.3) (:11) (18.1) (:520) 37 'wt/d 11.5 24.1 31.8 49.9 104.7 ae/ac/d 92.3 96.3 95.4 99.3 104.7 P/B/d 0.073 0.071 9.066 0.069 0 070 78 Table 10 (cont‘d.). 4-1 A-Z A-3 A-4 A-s 0 1500 ml after 4 d N3/aq 3 d 3 2 1 7'0: 295.2 683.0 951.3 1367.7 2928.5 (1S0) (1132.5) (1149.4) (1226.6) (1183.4) (1119.5) cv 44.9: 21.9: 23.8: 13.4: 4.1: 6/64 2361.8 2732.1 2853.9 2735.5 2928.5 (:50) (130.6) (136.7) (165.3) 357.2) (319.5) zi'wt/a 16.9 39.9 63.3 85.8 199.6 ;8/aq/c 135.2 159.5 189.9 171.6 199.6 27870 0.074 0.082 0.094 0.084 0.094 5L 12.0 15.9 19.1 23.1 29.9 Stages (Range) 5-(26-29) 5-(28-30) 5-(29-31) 5-(30-32) 5-34 Total Biomass Comparisonsa 4-3 4-2 4-5 4-1 3-4 1. Initial 701.0 706.0 706.5 707.8 714.4 A-l 4-2 4-4 4-3 4-5 2. 0 500 ml 1268.0 1443.0 1450.0 1483.0 1501.0 A-l 4-4 A-Z 4-3 4-5 3. 0 1000 m1 1820.8 2049.0 2055.7 2072.3 2129-0 A-l 4-2 A-3 4-4 A-S 4. 0 1500 ml 2361.8 2732.1 2735.5 2853.9 2928.5 a t 05(4) 3 2.776. 79 treatment averaged 16, 14, and 19% lower than in the other density treatments at the three algal supply rates. The greatest difference in biomass should have occurred at the SOO-ml supply level, since it was during this period that tadpoles died. Instead, the largest average in total biomass was measured during the final feeding period, 1,500 ml. This could indicate that intraspecific competition was beginning to exert an affect on tadpoles in A-l, resulting in slower growth of certain individuals. The daily growth rates of individual tadpoles at each of the five densities were compared after each feeding level and over the total 24-d period. No significant differences occurred among the individual growth rates of tadpoles within each density treatment at any of the feeding intensities or over the whole experiment (Kruskal- Wallis nonparametric test, H = 1.0, X2.01(4) = 13.277). These results indicate that the density treatments had no identifiable effect on the growth rates of individual tadpoles exposed to the experimental con- ditions. The mean change in wet weight/d over the entire 24-d study (values for all three feeding intensities were incorporated) was A-l = 11.5%, A-2 = 12.0%, A-3 = 12.8%, A-4 = 11.8%, and A-5 = 13.1%. If :small, density-related effects occurred, they were relatively minor compared to other factors which influenced growth. In general, the largest tadpoles in each density treatment grew fastest, especially VV?hen individual growth rates were averaged over the 24-d study without '“eésgard for algal supply rates (Figure 10). However, the average weight gain of the smallest tadpole and of all tadpoles in the d‘i‘Fferent density stockings was sometimes greater than or similar to 1C?)at:of the largest tadpole during certain feeding intervals. This 80 .mzwxuoum >»_mcmv sumo cw m_oaveu Agog umumgmczv ummmgep new Agog nmumcmv umm_PeEm we :wmm u;m_m3 ucmmmcamg mean Lw::_ .zogcmz .couxcmpqouxcq cm» Am1< canoes“ _1< mucmEummeu xuwmcmn :_ meansac Lee o_ open» mmmv mmpoawmu eo agave; cw :_mm mmcgm>< .op mg:m_u new a a a m a e. 3m .26» Econ. © .689 e .608 © 10 <1- 10 N N . m e. n" N _ ID :1- m N IO :3 f0 ' ‘1‘ I 3.381533.- 3, 4n 1'1‘-.':‘-":"-:1;“?:1:.15;:?.-.'.Tr‘:8:,"~.;:’r~',-"~:-‘:,~ y-z- -:- :3 - a» e. a.“ _. a» a my mm m. m T.» ”...... mm m_ #5 aw mum ”H .w _ E s 1%) p/(MHM 1.3M vae 81 indicated that complete absence of growth or stunting did not con- sistently occur among the smaller tadpoles in the different density treatments. Generally,the average change in weight/biomass/day was sub- stantially higher for tadpoles fed TetraMin than for algal-fed tad- poles (Table 10 and Appendix 8, Table 81). The two larger and better developed tadpoles, T(9) and T(10), exhibited average changes in weight like the algal-fed tadpoles. A relationship appeared to exist between the stage of development and the growth rates of tadpoles with younger, smaller tadpoles growing faster than older, larger tadpoles. The growth rates estimated for tadpoles fed TetraMin, a highly nutritious, digestible food, were higher than those measured for algal-fed tadpoles in the laboratory or for tadpoles in the field experiments at Lakes One and Four. Ingestion and conversion efficiencies: The mean tadpole production/total biomass/day fluctuated during the periods when tadpoles were fed the three concentrations of algae (Table 10). At the first algal supply rate, the overall average TP/TB for all density treat- ments was 0.077. When food was increased to 1,000 ml, tadpole pro- duction averaged 0.070; at the final supply rate, 1,500 ml, mean daily tadpole production per unit of total biomass was 0.086. To some extent, these differences in tadpole production may have been influenced by \Iéit~iations in the supply of algae per unit of tadpole biomass in each stOcking replicate. The concentrations of nutrients and phyt0plankton VaPied over the duration of the study (Table 3). Although the amounts (D'F' algae provided to tadpoles increased through time, the actual 82 available food did not increase as anticipated (the second supply rate was designed to be 2X the first supply rate, and the third, 3X the first). The particulate carbon and nitrogen supplied in relation to the carbon and nitrogen measured in the tadpoles for A-l (9 tadpoles) and A-5 (l tadpole), the treatments with the extremes in tadpole density, are shown in Table 11. In both density stockings, the average daily supply of carbon in relation to the growing biomass (carbon) of tadpoles decreased over the experimental period (Table 11). The ratio of the supply of nitrogen to tadpole biomass measured as nitrogen remained high at each supply level; the ratio (supply of nitrogen/tadpole nitrogen) was slightly lower at 1,000 ml of algae than at the 500- or 1,500-ml supply rate. However, the initial supply of carbon and nitrogen at the next supply rate was invariably higher than that at the previous feeding level (Table 11). The consumption of both carbon and nitrogen per unit of tadpole carbon and nitrogen decreased from the initial (500 ml) to the final (1,500 ml) supply level. Values for conversion (G/I) and assimilation (A/I) efficiencies indicated that tadpoles became ilhcreasingly more efficient at using the amounts of phytoplankton Farcovided with the highest efficiencies estimated for tadpoles at the 7 ,SOO-ml supply rate (Table 12 and Appendix B, Table 82). The con- \reer~sion of assimilated material into growth (G/A) was greatest for '1'éi1rraMin-fed tadpoles (Table 12 and Appendix 8, Table 82); the G/A estimated for TetraMin-fed tadpoles were similar to values for tadpoles 'f:€3<1 1,500 ml of cultured phytoplankton. The average carbon and nitrogen COHtent of tadpoles fed TetraMin was significantly greater than for ‘t3'1tase fed algae (P<0.00l, Table 2), although the C-N ratios were similar 83 Table 11. Supply and consumption* of carbon and nitrogen n relation to the demand (measared as tad- pole biomass in C and N) for the highest (A l) and lowest (A-5) densities of leopard frog tadpoles. Supply of C,N/8iomass C,N Consumption C,H/Biomass C,N A-l A-S A-l A-5 @ 500 ml 1. Initial a) carbon (t) 87.6 87.7 70.4 70.6 b) nitrogen (3) 114.9 115.1 92.6 92.8 2. Final a) carbon (2) l .9 41.3 39.3 33.2 b) nitrogen (Z) 64.) 54.2 51.7 43.7 3. Average 3) carbon (%) 58.2 64.5 54.8 51. 'b) nitrogen (5) 89.5 34.6 72.1 58.7 3 1000 ml 1. Initial a) carbon 1’ ) 67.3 5‘ 2 7.3 40.0 b) nitrogen (3 ‘ 97.2 32 1 85.5 72.3 2. Final a) carbon (3 ) 47.2 40.4 3.0 23.2 b) nitrogen (% ) 67.7 57.3 59.6 50.9 3. Average a) carbon ( a) 57.5 18.8 40.1 34.1 0) nitrogen (3) 82.4 70.0 7 .5 51.6 @ 1500 m1 1. Initial a) carbon (1 ) 60.2 51.5 40.9 35.0 b) nitrogen (3 ) 100.5 85.9 68.8 58.9 2. Final 8) carbon (5) 46.4 37.5 31.5 25.4 b) nitrogen (%) 77.5 62.5 53.5 42.8 3. Average a) carbon (5) 53.3 44.5 36.2 30.2 b) nitrogen (i) 89.0 74.2 60.9 50.3 'Average tadpole conSumotion/d (:50): G 500 ml/d .49) mg Particulate K eldanl N/aq 2) 15.2 (12.01) mg Particulate Organic C/ 30 @ 1000 957d .0 (r .14) mg Part‘culate (jeldahl Hr’aq 2) 18.3 (31.46) mg Particulate Organic C/aq 0 1500 ml/d 1) 7.:7 {Ii-)5) mg Particulate Kjeldahl N/ao 2) 22.7 (12.58) mg 2articulate Organic C/aO 84 Table 12. GrOwth (mg C), assimilation (mg C), and efficiencies (carbon) of tadpoles (3. oioiens) fed algae or TetraMin. Growth/ Conversion Assimilation Assimilation Efficiency Efficiency Growth/d Assimilation/d (G/Az) (G/IS) (A/Ii) PHYTOPLANKTON A-l ”to ' 9, Nt]-3 ' 8 500 ml 1.22 3.17 38.5 .0 20.9 1000 ml 2.81 6.04 46.7 15.4 33.0 1500 ml 4.13 8.13 50.8 8.2 35.3 A-Z "to-3 ' 4 500 ml 1.53 3.48 43.9 10.1 22.9 1000 ml 2.94 6.24 47.2 16.1 34.1 1500 ml 4.37 9.35 53.8 21.5 4 .2 4-3 “t0-3 - 3 500 ml 1.66 3.59 45.0 10.9 24.3 1000 ml 2.91 6.2 46.9 15.9 34.0 1500 ml 5.80 10.37 55.9 25.5 45.7 A-4 N a 2 t0-3 500 ml 1.60 3.96 40.5 10.5 26.0 1000 ml 3.05 6.26 48.7 16.7 34.2 1500 ml 5.24 9.31 53.4 23.1 43.2 A-S N. a 1 “0-3 500 ml 1.73 3.84 45.1 11.4 25.3 1000 ml 3.2 6.67 47.9 17.6 36.4 1500 ml 6.09 10.35 55.7 2 .8 48.2 TETRAMIN 7(1) 1.55 1.41-2.38 72.3 - - (90.6-54.0) T(2) 4.33 4.56-7.83 74.9 - - (94.6-55.3) 7(3) 6.85 7.17-12.31 75.5 - - (95.5-55.6) 1(4) 5.01 5.53-9.18 72.6 - - (90.6-54.6) 1(5) 7 68 8 16-14 14 72.5 - - (90.8-54.7) 1(6) 8.24 9.46-15.66 63.9 - - (87.1-52.5) T(7) 9 19 11 05-17 98 67.1 - - (33.1-51.1) 7(8) 8.44 10.52-16.88 65.1 - - (30.2-50.0) T(9) 7 02 10.35-15 84 55.4 - - (66.5-44.3) T(lO) 11 36 11 37-17 35 62.3 - - {75.3-48 3) ¥ - indicates not measured. 85 in all tadpoles, regardless of the types of food ingested. Larger tadpoles fed TetraMin had slightly lower G/A and average daily weight gains than smaller tadpoles. Since tadpoles were fed excess TetraMin daily and density effects were excluded, this decrease in efficiency appeared related to developmental stage of the tadpoles. Effects of Tadpoles on Organic Material and C, N, and P Dynamics Tadpole GrazinggIntensity and Periphyton (Experiments One and Two) Lake One: Attached algae colonized all aquaria during the equilibration period before aquaria were stocked with tadpoles. The amount of organic material (oven-dried) averaged 1 g/aquarium in Experiments One and Two; no significant differences in amounts of periphytic algae and associated materials occurred in aquaria before tadpoles were introduced. After tadpoles were stocked in Experiment One, the daily accumulation of attached algae in stocked aquaria declined to about 30% of the accumulation rate in unstocked aquaria (Figure 11, Table 13 and Appendix B, Table 83). In Experiment Two, the tadpoles again significantly reduced the rates of periphytic accrual (Figure 11, Table 11 and Appendix B, Table 83). The daily accumulation of attached algae in aquaria with a high biomass of tad- poles declined to about 20% of that for unstocked aquaria. The aquaria with a low biomass stocking of tadpoles had accumulation rates about 30% of those measured for unstocked controls. In unstocked aquaria, Inean rates of periphytic accumulation were slightly higher in the first experiment than in the second. In both experiments, periphytic carbon invariably comprised a major portion of the daily total amounts of organic carbon, regardless of treatment (Figure 11). -o_a zo_ .mcan uwumzmca n mwcmzcm umxuoumczv :oncmu owuxggwcmq ucmmmcqmc mean cmccH um ozp ecu mco mucwewcmaxm Low fizzwcmzcm\me u mo—V :oncmu uwuchwcma use :oocmu owcmmco _mgoh NNum W vmx<4 mmxw 9-0 NTw mum run .11. o_.»=a_..a a 2:095 .20». uzomm_0mm_o Ao mwz_omm_o Ao moz,8mm_o Ao mwz.8mm_a Ao waz_w>_uumqmmc .mmocc :mmcm vcm mw_owam1umst .mmoc$—_:n acmmmcamc mean vmumcm ucm swam; mxm_ ucmmmcamg mean uwumzmcz .csou use mco mmxcm um awash acmswcmaxm mcwcau awcosca cw coagmu u_:oux:apaouzca ace uwuznawcma .mp mc=m_u v ux<4 n mxw _ 934 n mxu 3~1NNY9 8.10510. 31310. @N1umv1m avunumvuo. .~.1o=1o_ 31310.?N1N81m ;\ /9 My“ ”a“ .. ”We é We“ am - 1. zo._.v.z<._m0h>:n_ r N N wnuoan/(bwmoeuvo 601 \ /:.\ /_o. \ I$\ fl§\ I£\ .\ I§\ lfi\ Ig\ W 1 [am Is\ Ig\ \ /s\ ”s\ Iz\ \ - I:\ /§\ /3\ x mm“ wee yea -N zo»>:a_¢ua rm 104 (Table 16, Figure 13). Comparisons of the average daily changes in periphytic carbon at Lake Four indicated that the mixed-species culture and green frogs reared in monoculture depressed periphyton the same amount, resulting in lower periphytic accrual than measured for bullfrogs in single-species treatments (Table 16). However, considerable variation occurred among sampling dates for the three tadpole treatments at Lake Four with no observable trend in total periphytic carbon among the tadpole groups tested. Comparisons of Experiment Three with Experiments One and Two: Results from Experiments One and Two with 3, pipiens were compared with those of Experiment Three with 3, catesbeiana and B, clamitans to determine if all species had similar effects upon primary producers. Relatively high biomasses of bullfrog and green frog tadpoles were stocked in Experiment Three, compared to stockings of leapard frog tadpoles in Experiments One and Two. At Lake One, the mean accrual of periphyton averaged somewhat less in Experiment Three than the mean accrual in Experiments One and Two, presumably because of the high stock of tadpoles and the shorter days with less light and cooler temperatures. At Lake Four, the mean daily accrual of periphyton in Experiment Three was similar to or slightly higher than that measured in Experiments One and Two. Periphyton declined more than anticipated in Experiment Three at Lake One and increased more than expected at Lake Four, based on results from Experiments One and Two. Tadpole survival was higher in Experiment Three than in Experiments One and Two. At Lake One, tadpole production in Experiment Three was similar to production in Experiment One but lower than tadpole 105 production measured in Experiment Two. In summary, green frog and bullfrog tadpoles at Lake One produced more carbon than expected at the cooler temperatures and high biomasses of stocked tadpoles (see values for production/aquarium/degree d in Tables 7 and 9). Tadpoles at Lake Four in Experiment Three lost weight and may have died eventually of starvation as did 1e0pard frog tadpoles in Experiment One. However, bullfrog and green frog tadpoles at Lake Four lost less weight than expected, based on results from 3, pipiens treatments (see values for production/aquarium/degree d in Tables 7 and 9). The phytoplanktonic carbon increased in all aquaria in Experiment Three over levels measured in the imported lake water as it did in tadpole-stocked aquaria in Experiments One and Two, except for the high biomass stocking of R, pipiens tadpoles in Experiment Two at Lake Four. In Experiment Three at Lake Four, the increase in the abundance of phytOplankton over values for imported lake water was proportionally greater than previous increases for any aquaria with tadpoles in Experiments One and Two. In summary, bullfrog, green frog and leOpard frog tadpoles appeared to similarly affect the autotrOphic communities in their respective aquaria. All three species consumed mainly periphyton and associated settled materials and usually stimulated the accumulation of phytOplankton in the water column. Effects 9f_Varying Tadpole Stocking Rates on Nitrogen (Experiments One and TwO) Lake One: Total inorganic nitrogen imported in lake water was over three times higher in Experiment One than in Experiment Two (Table 17 and Appendix B, Table 85). In the first experiment, nitrate 106 88.8 88.8 88.8- 22 88 888888 888 8882888888: 28882882 mu.m 88.8 88.8 o 8888888888 888 m_o8888 28888 88.8 88.8 8 8 8888888 88.8 88.8 88.8 8 8888288888 88.888 88.88 88.888 88.82 88.888 88.8 88.888 8828888 888 888888888 88888 88.88 88.28 88.88 82.88 88.88 88.28 88 88 88888888 88.88 88.88- 88.88 88.88- 88.88 88.88- 88.88 88>888828 88.88 88.8 88.88 88.8 82.88 88.8 - 88.88 8288888 8888888 88.8 88.8 88.8 88.8 88.8 88.8 88.. 8828288 88.888 88.8 88.888 88.88 88.888 88.8. 88.888 8888822 88.8 88.8 - 88.8 88.8 - 88.88 88.8 - 88.8 8888882 88.888 82.8 88.888 88.82 88.888 88.88 88.888 2888888 888888888 828 8282888828 -- 828 8228 88888828 888888 88888828 888888 88888828 888888 8888 2888 28888882 82828888 2822 82828888 288 882888828 888828 8288828 <~m<=d< zH : em am>c mazo muz_0mmwo mm._m mm..- o_.~m mo.~- m_.~m ¢F.N- mo.om “_aBOPV oweaaco _m.o m_.o- Ne.o qo.o- _m.o mo.o- m¢.o accoae< mm._ mm.o- ap.F 85.0- _o._ mm.o- ma.o mocca_z me.o o m¢.o o me.c o m¢.o mo_cu_z ~¢.N «P..- mo.~ _m.o- mm._ No.o- m~._ “_aHOHV u_=amco:_ mzo pzmzfimmaxm -- m=OL m¥o uwz;a_cm¢ mm.mm qa.m No.o¢ _N.N Ne.mm ok.m m~.~a Because new u_=ameo=H _aooe mm.~ mm.m mm.P am.~ oo.~ cm.m _N.¢ u_=oummm mm.mm om.P- o_.mm mN._- ow.¢m mm.o- mm.mm um>P0mm_a mo.om mm.~ 8o.~m cm._ mm.om Pu.” mm.mm A_apohv u_=mmco m~._ co._ o_.~ _~.o m~.o mo.N _m.N a_=oas< m¢.o o me.o o m¢.o o me.o moccu_z m¢.c o me.c o m¢.o o m¢.o mu_cp_z _o.~ mo._ om.~ Pk.o Nc._ mo.m No.m A_aoo»v u_=amco=H o3» hzmszmaxu -- «no; m¥<4 auemoaxm amzopm omemoaxm ammoem ouhmoaxm gumOHm m¥<4 zoma zmoomhfiz wzuxoohm ram: wszQOFm 304 ouxu0pmz: o mozo uuz<=u z_OmmPo mm.m__ mm.om- 0N.MN_ mo.mm- mm.~mp mo.ee- Np.mm A_epoev u_=emco om.~ o“.m om.m we.“ om.m mm.o om.m_ necess< em._mm mm.o + Ne._mm m~.m - m~.-~ me.o - me.NN~ mpecu_z oe.o~ oo.P - 58.0N NN.. - _N..~ _m._ - oe.m_ ao_co_z om.mm~ mm.m + em...m mo.m - mm.mom om.e mp.mom Apeooev u_=mmeo=H “were Fzmzmamaxw -- uzo m¥o mwz;a_cma _eHOH em.o . mo.o . 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Ne.m um.oF+ mo.m~ euepzowecea em.¢m em.m . m~.m~ ¢F.m + Ne.Pm um.“ . oe.e~ ee>_ommwo Fm.om mm.e + mm.wm e_.N + mo.wm oo.m + mo._e Apeuohv uwcemco Fm.o o _m.o o uo.o ep.o + _m.o e_:oes< me.o o me.o o me.o o me.o epece_z me.o o m¢.o o me.o o me.o epwce_z No.P o so._ o mm._ e_.o + no.— Apeuopv u_:emco:H ummzp hzmszmaxm -- mac; mx<4 anemoaxm ommOHm ompmoaxm cumopm ouhmOme awashm m¥<4 zomu zuwoahmz hmoazH 4o eez<=e z.0mmeo _e._ ee.e- e_.P pm.o- eF.P .~.o- mm.o _eeoe ezo ezezmmeaxe -- mace weep oe.ep Ne.m- me.m_ me.o- _e.N_ em.~+ em.e_ eeeF=u_ecea _m.ee eN._+ em.ee me.o+ mm.om e_.e- me.ee ee>_Ommwo _m.me ep.N- N~.me o em.me me.o+ Ne.me _eeoe ozp ezezemeexe -- ezo ex_Omm_e ee.e~ o mm.m~ me.o- Ne.em ee.m+ ee.e~ _eeoe ezo Ezezeeeaxe -- ezo ex_0mm_e N«.. o _e.F eo.e- _e._ eo.o- mm._ _eeoe oak Ezezeeeaxe -- mace e¥o mwzwemmwa mm.m weeew mmmzh hzmzammmxm nu anon mx<4 mm.mm eee_=ewecea _e.~e ee>_0mm_e e~.___ _eeow mumzh hzmzwmmmxm an uzo m¥<4 omhmoexm ommOHm omhmoexu amePm omhmoexm ommOHm macaw zmmmw wwwumem omxwz mwommggam e eez<=e z“ ( 1 (D 124 PhytOplanktonic Composition and Diversity Overview Ranid tadpoles potentially can influence the composition, diversity, and abundance of phytoplankton, since these tadpoles are among the larger suspension feeding vertebrates in aquatic environ- ments of North America. The affect on the algal community may be direct, a result of their selection of particular algal taxa, or indirect, related to their influence on nutrient cycling or other physical and biological components. This portion of the field study was designed to elucidate the interactions between the phytoplanktonic community and different biomasses (densities) and Species of tadpoles at Lakes One (hypereutrophic-eutrOphic) and Four (mesotrophic- eutrophic). In general, the same genera of algae were encountered at Lakes One and Four on particular sampling dates; however, abundances and proportions of phytOplankton in major algal groups differed sub- stantially between the two lakes. Algal succession occurred at both lakes from May to OctOber, so no particular algal taxon was dominant during any one experiment. The genera and volumes of phytoplankton sampled at Lakes One and Four during this study are presented in Appendix B, Table 810. Experiment One (Lakes One and Four) Imported lake water: In Experiment One, total algal numbers and volumes averaged two to three times higher at Lake One than at Lake Four (Appendix 8, Tables 811-812). Initially, diatoms were most m! :5 A: 125 numerous with lower concentrations of cryptophytes and green algae in water imported from Lake One (Appendix 8, Table 812). During the experiment, diatoms decreased in abundance and green algae increased, which resulted in green algae comprising the largest proportion of algal numbers and volumes on the final sampling date in June. Algal succession also occurred at Lake Four, but blue-green algae numerically dominated the algal community at the conclusion of this study (Appendix 8, Table 813). Diatoms were dominant volumetrically at Lake Four throughout the experiment; however, cryptophytes, green and blue-green algae were relatively important on particular sampling dates. Tadpole effects: The major effects of tadpole feeding on the algal assemblages in Experiment One at Lake One are summarized in Table 22. In general, the activities of tadpoles stimulated phyto- planktonic growth; the highest total numbers and volumes of algae were invariably sampled in stocked aquaria (Table 22 and Appendix 8, Table 811). The effects of grazing intensity, simulated by low or high densities (biomasses) of tadpoles, were usually not statistically separable. Although both tadpole treatments influenced the algal community similarly, the magnitude of the response was typically greater in aquaria stocked with a high abundance of tadpoles. On 6 June, after tadpoles had been in aquaria for six days, all algal groups (green algae, cryptOphytes, diatoms, and blue-green algae) increased in abundance. In contrast, unstocked aquaria on this date had substantially lower concentrations of phyt0plankton (Table 22 and Appendix B, Table 811). Among the major phyla of phytOplankton sampled, blue-green algae and diatoms were consistently elevated by 1226 Table 22. Phytoplanktonic comparisons between unstocked (C for control) and tadpole-Stocked (L for low and H for high stock of 3. pipiens) aquaria in Experiment One at Lake One. 6 June 6 June La) 0 12 June 19 June Combined 3 Dates 6 June Combined 3 Dates (JO 0 6 June 12 June 19 June Combined 3 Dates m 6 June 12 June 19 June Combined 3 Dates 6 June 6 June 12 June Combined 3 Dates 6 June 12 June Combined 3 Dates ’3 June EXPERIMENT ONE--LAKE ONE Numbers of Greens F - 19.3" 3607(C) 7063(H) 10539(L) Volumes of Greens F - 11.07** 301484(C) 496537(5) 842028(L) Numbers of Diatoms F a 4.41 7233(C) 195271L1 26678(H) E = 4.37 d046437(§)7 5111234.7(L) 6764816.3(H) F . 3.J7** 9897(C) TSSOE(L), ZOJZQLHI Volumes of Diatoms F a 4.88 3595389.7(C1 4634885.7(H1 ? = 3.:a*~ 3475970(c) Number of Blue-greens 5692362.3(L3 F - 27.18*‘* 52(C) BSOS’L) 3544(Hl F a 4.93 2022(Cl 7236(L1 9229’H) F s 3.89 3896(C) 9408(L) lOl79(Hl F a 7.94** 1993(C) 6716(L) 7651FH) Volumes of Blue-greens F a 30.42*'* 1030(C) 540020.7fL} S7672§Lfll F = 6.70' 76405(C) 920241.7(L) l246660.7(Hl F = 14.58" 91167.7(C) 909720.7(H) lOllOll(L) F a l9.77*** 56201(C) 82357.8(L) 911035.8(9‘ Numbers of CryptOpnytes F 8 7.90' SEQ/C) 15921L1. 2069(H) Total Numbers of Algae F - ZS.S3** 15752(C) 26596(H) 3277S(L) F a 6.48* 25810(C)_ 48058(L1 62263(H) F ' 5.74** 26957(C) d39460.1 4814518) Total Volumes of Algae F . 8.65' 42549571Cl 579957818) 7738370(L) F = 4.32 1382065.3(C) 9n50150le 12223997.7(H) F 3 9.32**' 5175412.4(C) 3585894.2’L) 931636:(5L Numerical Diversity of Diatoms r = 22.98** 0.3643(Ll 9.36781”) 1.300(C) 1237 Table 22 (cont'd.). EXPERIMENT 0NE--LAKE ONE 11. Volume Diversity of Greens June F - 8.18* 0.9040(H) 0.9627(Cl 1.1533(L) 12. Volume Diversity of Diatoms 12 June F - 8.16' 0.4530(L) 0.5000(8) 0.8267(C) 19 June F - 27.99’** 0.3797(L), O.d737(H) 1.1650(C) Combined 3 Dates F - 3.09 0.4276(L) 0.4401(H) 0.7259’C) 13. Volume Giversity of Blue-greens 12 June F a ll.68** 0.3327’H) 9.1363(L) 0.5567(C) 14. Mean Sizes of Greens 12 June F a 4.39 70.3(H) 79.?(L1 83.5(C1 15. Mean Sizes of Blue-greens 6 June F a 194.46*** 16.5(C) 164.1(L1 162.7191 12 June F . 25.49" 37.8(C) 127.2(L) 135.1(Hl 19 June F . 15.95" 23.4(C1 39.4(H) 107.5(L2 Combined 3 Dates F a 15.82*** 28.2(C) 119.1(5) 122.7(L) 16. Scenedesmus Combined 3 Dates F . 10.06" 1178(H) 1648(L) 53d7(C) 17. Schroederia Combined 3 Oates F I 20.43*** 677(C) d145(L) 5194(8) 18. Oictyosphaerium Combined 3 Dates F a 7.07**' 306(C) 3306(L) 4090(5) 19. Nitzschia Combined 3 Dates F a 7.13" 8472(C) 170831L) 19OS7(H) 20. Coccoid Blue-green Cell Combined 3 Dates F a 20.76*** 235(C) 5156(L) 5546(8) ”F.05 (2.6) ' 5-‘4i “.05 (2.24) ' 3°‘0 .'F.0] (2’5) 3 10.90; F.01 (2’24) ' 5.61 "’F.001 (2.5) ’ 27-90‘ F.001 (2,24) ' 9-3‘ 128 the activities of tadpoles (Table 22). This pr0portional increase in blue-green algae and diatoms in stocked aquaria was not related to the decreased abundances of green algae and cryptOphytes, since values for these groups were similar to those in unstocked aquaria (Appendix 8, Tables 811 and 813). Rather, the numerical proliferation of diatoms and blue-green algae in stocked aquaria decreased the relative contributions of green algae and cryptophytes to the total algal assemblage. The mean size of blue-green algae was always greater in aquaria stocked with tadpoles, while green algae tended to be larger in un- stocked controls (Table 22 and Appendix B, Table 811). The lowest numerical and volumetric diversity of diatoms, volumetric diversity of blue-green algae, and except on 6 June, the lowest total numerical and volumetric diversity and evenness were measured in stocked aquaria (Table 22 and Appendix 8, Table 814). Green algae, Dictyosphaerium and Schroederia were significantly elevated and Scenedesmus was signif- icantly depressed in stocked aquaria (Table 22 and Appendix 8, Table 815). The diatom Nitzschia and a coccoid blue-green alga were more numerous in tadpole treatments than in unstocked controls (Table 22 and Appendix 8, Table 815). Chlamydomonas and small Anabaena tended to decrease in aquaria stocked with tadpoles (Appendix 8, Table 815). Concentrations of other algal genera varied among treatments and sampling dates with no distinctive patterns. In Experiment One at Lake Four, tadpole grazing stimulated phytoplanktonic increases, resulting in greater total algal numbers, total algal volumes, and mean sizes of algae than occurred in unstocked aquaria (Table 23 and Appendix 8, Table 812). The effects of grazing IZTQ Table 23. Phytoplanktonic comparisons between unstocked (C for control) and tadpole-stocxed {L for low and H for high stock of R. pipiens) aquaria in Experiment One at Lake Four. 1. Number of Greens 6 June F I 5.40* 2. Volume of Greens 6 June F I 6.55' 3. Number of Diatoms 6 June F I 11.89'* 12 June F I 7.42* 19 June F I 18.35" Combined 3 Gates F I 8.92'* 4. Volume cf Oiatcms 5 June F = 6.62‘ 12 June F I 7.36’ 19 June F I 18.29** Combined 3 Dates F I 28.16*‘* EXPERIMENT ONEI-LAKE FOUR 2698(61 3884(5) 5069(L) 14351§(C) 257266(H) 371645.3(L) 440(c) 918(L) 1698(H) 245(C) 4960(L) 7580(H) 1009(5) 22431L) 5102(H) 565(C) 2707(L) 4793(H) 145783.7(C) 292666.711}. 610393(H) 751071;) 1567038(L‘ 2403114(H) 318844(C) 709482(L) 1612232(H) 179911.7(C) 856395.6(L) 1541913{H) 5. Volume of Blue—greens 6 June F I 7.91' (.70 6 June F I 17.02" 12 June F I 6.68' Total Number of Algae 7. Total Volumes of Algae 6 June F I lS.77** 12 June F I 9.00* 19 June F I 18.84** Combined 3 Dates F I 12.63'** \ 8. Numerical Diversity of Greens 6 June F I 17.93" Combined 3 Oates F I 4.16* 1.005(c) 1.070(C) 9. Volume Diversity of Blue-greens 6 June F = 5.45' 0.394(H1 54980(C) 19mm) 219701.7(H) 4413(5) 7833(H) 83901Ll 3669(C) 9947(L1 11862(fll 148847.7(C) 933800(L) 1225962(H‘ 215826(C) 1806356(L) 2639837(Hl 1105529(Cl 15846491L) 2299105(H) 590494(C) 1441601.8(L) 2049387.l(H) 1.375(8) 1.453;;1 1.290(L) 1.301151 0.205(L) 0 5511C) 130 Table 23 (cont'd.). EXPERIMENT ONE--LAKE FOUR 10. Total Volume Diversity of Algae 12 June F . 22.78" 0.489(H) 0.525(L) 1.685(C) Combined 3 Dates F - 2.97 1.051(3), 1.205(L), 1.602(C) 11. Volume Evenness 12 June F I 20.68** 0.202(H) 0.248(L) 0.736(C) Combined 3 Dates F I 6.92** 0.419(H) 0.492(L) 0.580(C) 12. Mean Sizes of Greens 6 June F I 6.36‘ 53.2(C) 68.3(8) 73.3{L1 13. Mean Sizes of Total Algae 5 June F I 5.38* 101.7’C1 111.3(L) 156.5(H) 12 June F = 67.71*** 58.8(C) 135.0(Ll 222.5(8) 19 June F I 14.81** 80.6(C) 93.2(L) 147.5(H) Combined 3 Dates F I 18.35*** 81.5(C) 122.4(L) 174.3(H) ld. Scenedesmus Combined 3 Dates F I 7.64** 75(C) 508(H) 641(L) 15. Nitzschia Combined 3 Oates F - 3.19II 553(C) 2599(1) 4773(H1 16. Coccoid Blueogreen Cell Combined 3 Dates F I 4.51* 231(C) 719(L) 779(3) 17. Small Anacystis Combined 3 Dates F I 7.dl** 61(H) 470(L) 7891C) I F .05 (2,6) ' 5"“; F.0s (2,24) ' 3-40 .01 (2,6) ' ‘0'901 F.01 (2,24) ' 5-51 .001 (2,5)' 27°°°‘ F.001 (2.24)” 9°34 ti ti’f F in: w nuri COM ah- 131 intensity, simulated by low or high densities (biomasses) of tadpoles, influenced the algal community similarly, but the response was sig- nificantly greater in magnitude in aquaria with high numbers of tad- poles. Aquaria stocked with a low number (biomass) of tadpoles generally had algal abundances and volumes that were intermediate between those measured in unstocked aquaria and in aquaria with a high density (biomass) of tadpoles. The activities of tadpoles in both treatments elevated diatom numbers and volumes, total numbers of algae, and mean sizes of total algae over values in unstocked con- trols; however, these measurements were significantly higher in aquaria with a high stocking of tadpoles (Table 23 and Appendix 8, Table 812). The greater numbers and volumes of diatoms sampled in stocked aquaria were related mainly to significant increases in Nitzschia (Table 23 and Appendix 8, Table 815). The pr0portionately greater increases in numbers and volumes of diatoms in stocked aquaria lowered the relative contributions of all other algal groups, especially in aquaria with a high stock of tadpoles (Appendix B, Table 813). Numbers and volumes of green algae tended to be greater in aquaria stocked with tadpoles than in unstocked aquaria (Appendix 8, Table 812). In particular, one genus of green algae, Scenedesmus, was more numerous in aquaria with tadpoles (Table 23 and Appendix 8, Table 815). Tadpole grazing also influenced the numerical diversity of green algae, resulting in higher diversity values than those estimated in aquaria without tadpoles (Table 23 and Appendix 8, Table 816). After 6 June. the numbers and volumes of blue-green algae were lowest in aquaria with a high stocking of tadpoles. The total number of blue-green algae were typically lowest in unstocked :fiu 132 aquaria, while the volumetric diversity of blue-green algae tended to be highest in aquaria without tadpoles (Table 23 and Appendix 8, Table 816). Small Anacystis was least abundant in aquaria with tad- poles and a small coccoid blue-green alga was invariably elevated in stocked aquaria (Table 23 and Appendix B, Table 815). Throughout this experiment at Lake Four, activities of tadpoles decreased the volumetric diversity and evenness calculated for the total algal community (Table 23 and Appendix 8, Table 816). As at Lake One, on 6 June, the first sampling date after tadpoles were stocked, signif- icant increases were measured for many algal variables, including numbers, volumes, and mean sizes of green algae; volumes and volumetric diversity of blue-green algae; and total numbers, volumes and mean sizes of algae in stocked aquaria. Experiment Two (Lakes One and Four) Imported lake water: In July, when Experiment Two began, the total numbers, volumes, and mean sizes of algae were larger in the water imported from Lake Four than from Lake One (Appendix 8, Tables 817-818). During Experiment Two, algae bloomed at Lake One and after- wards, total algal numbers and volumes were much greater than at Lake Four, where no bloom occurred. Initially, the algal assemblage at Lake One was dominated numerically by green algae and volu- metrically by diatoms (Appendix 8, Table 819). Before the green and blue-green algal bloom in late August, cryptophytes, diatoms, and blue-green algae comprised a major proportion of total algal volumes and numbers on particular sampling dates. At Lake Four, the phyto- planktonic community changed from a blue-green algal assemblage to V1 133 one dominated volumetrically and numerically by cryptophytes (Appendix B, Table 819). Tadpole effects: Higher numbers of green algae, numbers and volumes of diatoms, and total algal numbers and volumes were con- sistently measured in stocked aquaria at Lake One in Experiment Two (Table 24 and Appendix 8, Table 817). In general, diatoms comprised the highest percentage of total algae in stocked aquaria, except at the end of August, when a bloom of green and blue-green algae occurred (Appendix 8, Table 819). At this time, numbers and volumes of diatoms were twice as high in stocked aquaria as in unstocked aquaria, but the proportionately greater increases in green and blue-green algae in stocked aquaria decreased the importance of the contribution by diatoms (Appendix 8, Tables 817 and 819). The diatoms, Nitzschia and Navicula, were elevated by tadpole activities throughout the experi- mental period (Table 24 and Appendix 8, Table 820). In Experiment Two, as in the first experiment at Lake One, the reSponses of the algal community to grazing intensity by tadpoles could not be separated statistically, except for differences in diatom volumes (Table 24). However, the magnitude of the reSponses of the algal community to tadpole grazing was typically greater when tadpoles were more abundant. On any particular sampling date in Experiment Two at Lake One, stocked aquaria had greater numbers, volumes and mean sizes of blue-green algae and volumes of green algae (Table 24 and Appendix B, Table 817). The volumetric diversity of green and blue-green algae tended to be higher in stocked aquaria, while total volumetric evenness and diversity were highest in unstocked aquaria on particular dates of Table 24. low and H for high stock of 3. pioiens aquaria in Experiment Two at Lake One. 134 Phytoplanktonic comparisons between unstocked (C for control) and tadpole-stocked (L for 8 August 15 August - 22 August 15 August 8 August 15 August Combined 3 Dates 8 August 15 August Combined 3 Oates 22 August 3 August 22 August 8 August 15 August 22 August 8 August 15 August 22 August Combined 3 Dates 22 August 15 August 4. 0| 0 EXPERIMENT TWO-ILAKE ONE Numbers of Greens F I 6.26* 9976(Cl 12184(L) lSZO3(H) F I 33.01*** 6412(C) 17104(L) 21551(H) F I 7.68* 58862(C) 283697(L) 317121(H) Volumes of Greens F I 5.31* 865732(C)_¥4g 1820706.3(L) 2181011.3(H) Numbers of Diatoms F I 8.78* l3815(§) 13432(Ll 29148(H) F I 16.53** 22558(C) 68633(L) 123292{H) F I 3.59* 116128(Cj 37827(L) S8428(H) Volumes of Oiatoms F I ll.57** 7212504(Cl 7396200.3(Ll 1469TOOO(H) F I 24.02** 13979000(C) 43388666(L) 77581333(H) F I 3. 64* 8906340(C) 21080046(L) 34501262(H) Numbers of Blue-greens F I 5.32* 131470(C) 39225?(H) 412091(Li Volumes of Blue-greens F I 12.54*' 10635.7(6) 23471(L) 194477.7(H) F I 5.42* 1921038.3(C1 Total Numbers of Algae 6078900.3(H) 6383489.3(Lj_ F I 12.52** 28442(§)g 36885(L) Sl327(H) F I 7.63' 112839(C)7 215541(L) 284902(H) F - 5.5511 203103(c) 7245990) 734235(H) Total Volumes of Algae F I 16.SO** 15507867.4(C) l9366893.3(L), 2642847S(H) F I 27.68*** 16136822.3(C) 47343307.7(L) 82248394(H) F I 4.75 l3746452.3(C) 40147420.3(H) 44538425.?(L) F I 7.62" 15130380.7(C) 37091329.7(L) 49004247.1(fl) Numerical Diversity of Greens F - 5.12I 1.032(L) 1.084’H) ‘ 2971c: Volume Diversity of Greens F I 12.00** 1.328(C) 1.655(8) 1 ‘ASLL‘ 1135 Table 24 (cont'd.). EXPERIMENT THO--LAKE ONE 11. Volume Diversity of Blue-greens 15 August F I 8.35* 0.086(C) 0.339(L) 0.461(5) 12. Total Volume Diversity 15 August F I 4.92 O.799(H)___ O.900(L) 1.034(C) 13. Volume Evenness 15 August F I 9.69* 0.277(5) 0.308(L} 0.400(C) 14. Mean Sizes of Blue-greens 3 August F I 5.96' 7.9(C) 16.3(L) 83.3(H) 15. Nitzschia Combined 3 Dates F I 4.94' 7502(8) 1991¢(L) 31337(H1 16. Navicula Combined 3 Dates F I 4.86' 3137(C) l?O80(L) 25079(H) * “.05 (2,5) . 5.14; F.05 (2,24) - 3.40 " F-01 (2.6) ' ‘0'90‘ F.01 (2,24) ' 5-51 = 9.34 ”' F I 27.00; F .001 (2,6) .901 (2,24) the b1 “JG 0016 136 the study (Table 24 and Appendix 8, Table 821). Consistent trends in diversity and evenness components between stocked and unstocked aquaria were not obvious in Experiment Two at Lake One (Appendix 8, Table 821). Concentrations of the green algae; Scenedesmus, a small coccoid green alga, Chlorella, Ankistrodesmus, and Actinastrum, and the blue-green algae; small Anacystis, small Anabaena, and a coccoid blue-green alga were typically elevated in aquaria stocked with tad- poles (Appendix 8, Table 820). Tadpole grazing in Experiment Two at Lake Four consistently increased abundances of Nitzschia and numbers and volumes of all genera of diatoms over values measured in unstocked aquaria (Table 25 and Appendix 8, Table 818). Both the total number of diatoms and concentrations of Nitzschia were influenced by grazing intensity; aquaria with a high stocking of tadpoles elevated diatom numbers and concentrations of Nitzschia above levels measured in aquaria with a low stocking of tadpoles (Table 25 and Appendix 8, Tables 818 and 820). Diatoms comprised the largest percentage of total algal volumes in stocked aquaria and blue-green algae contributed the highest pr0por- tion to total volumes in unstocked aquaria (Appendix 8, Table 819). Different algal taxa were dominant in the three treatments on partic- ular sampling dates; however, the average values for numerical com- position of algae were similar in all aquaria in Experiment Two (Appendix 8, Table 819). The total number of algae was consistently lowest in aquaria with a high density (biomass) of tadpoles, while the mean size of all algae was lowest in unstocked aquaria (Appendix 8, Table 818). On particular sampling dates in Experiment Two, stocked aquaria had Taole 25. low and H for high stocx of 3. pipiens) aquaria in Experiment Two at Lake Four. 137 Phytoplanktonic comparisons between unstocked (C for control) and tacpole-stocxed (L for 15 August 15 August 3 August Combined 3 Dates 8 August 22 August Combined 3 Dates 22 August 22 August 8 August 22 August 15 AuguSt 22 August 16 August 22 August 15 August 15 August 10. 11. 12. EXPERIMENT TWO-ILAKE FOUR Numbers of Greens p . 14,25" 5444(11) 7079(L) Volumes of Greens F I 9.95* 479924.3(H) 568170(L) Numbers of Diatoms F I 10.72* 56(0) 133(L) F I 4.70* 199(C) 436(L) Volumes of Diatoms F I 9.21' 38243(C) IIIZTBIL) F I 87.52"* 1725’6(C) 548653 7(8) F I 5.66" 1454mm 432508 11;.) Numbers of Blue-greens 13704(C) 887225(C) 266(8) F I 12.60** 15497(H) 15894(L) 21146(C) Volumes of Blue-greens F I 20.97** 282122.7(L) 286966.7(H1 546721(C) Numbers of Cryptopnytes F I 7.0' Q(L) 95C) 22’?) Numerical Diversity of Diatoms F I 6.76' 0.5517(H) 0.9913(L) 1.1550(6) Numerical Diversity of Blue-greens F I 4.64 0.6227(§l_ 0.5813(Hl 0.9903(L) F I 9.33' 0.8953(L) 1.1400(H) 1.2753(QL Volume Diversity of Greens F I 12.41*’ 0.3324(C) 0.7947(H) 3 87S’(Li F I 6.59' 1.1623(C) 1.2573(L) 1 4970(H) Numerical Diversity of Total Algae F I 7.73* 1.1187(C) 1.1700(Hl 1 SETO'L) Numerical Evenness F I 3.98 0.493(C) 0.585'Lj O 68~ 4‘ Table 25 (Cont'd.). 1 353 22 August 22 August Combined 3 Dates Combined 3 Dates 22 August 22 August Combined 3 Dates 13. 14. 15. 16. 17. EXPERIMENT TWO-ILAKE FOUR Volume Evenness F I 5.95* Mean Sizes F I 6.10* F I 3.07 Mean Sizes F I 3.67 Mean Sizes F I 7.41’ Mean Sizes F I 5.20‘ Nitzschia F I 6.89** Of Of of of 0.607(L) 0.649(C) 0.695(9) Greens 56(H) 52.4(2) 136.3(L) 65.6(C) 78(8), 99.5(L) Diatoms 730.7(C) 822.3(H) 922.2(L) Blue-greens 16.7(Ll 18.5(Hl 25.6(C) Total Algae 46.7(C) 70.0(H‘ 33 57L, 128(C) 338(L) 481C”) ' F.05 (2.5) ' 5-14: F.05 (2,24) . 3.40 tiF .0: (2.5) *** F.001 (2,5) ' 27-00: 3.001 (2,22) . 9.34 I 10.90; F .01 (2,24) I 5.61 E. l39 greater numbers of cryptophytes, mean sizes and volumetric diversity of green algae, mean sizes and numerical diversity of total algae, and volumetric evenness of all algae (Table 25 and Appendix 8, Tables 818 and 822). Also, aquaria stocked with tadpoles had lower numbers and volumes of green algae, numbers and volumes of blue-green algae, numerical diversity of green and blue-green algae, and mean sizes of blue-green algae during the sampling period (Table 25 and Appendix 8, Tables 818 and 822). Concentrations of Ulothrix, a filamentous green alga, were lowest in stocked aquaria (Appendix 8, Table 820). Abun- dances of large Anacystis and small Anabaena were depressed in aquaria with a high stocking of tadpoles (Appendix 8, Table 820). Experiment Three (Lakes One and Four) imported lake water: In Experiment Three at Lake Four, con- centrations of phyt0plankton were the lowest measured in any experiment at either lake. At Lake One, algal numbers were 30-40 times higher and volumes were about 20 times greater than at Lake Four (Appendix 8, Tables 823-824). Blue-green algae contributed the highest propor- tion to total numbers at both lakes; green algae at Lake One and diatoms at Lake Four comprised the highest percentage of total volumes (Appendix 8, Table 825). A diverse green-algal assemblage was sampled at Lake One with Dictyosphaerium, Scenedesmus, Ankistrodesmus, Kirchniella, and a large coccoid green alga present as major genera (Appendix 8, Table 826). Chlamydomonas dominated the green-algal flora at Lake Four (Appendix 8, Table 826). At both lakes, Nitzschia comprised the major portion of the diatoms and Rhodomonas dominated the genera of cryptophytes. The blue-green algae at both lakes were 140 were primarily Agmenellum and Anacystis, but small Anabaena were numerous at Lake One (Appendix B, Table 826). At Lake Four, the mean total size of phytoplankton was twice that measured at Lake One and mean sizes of diatoms were consistently greater at Lake Four (Appendix B, Tables 323-824). Tadpgle effects: In Experiment Three at Lake One, comparisons among aquaria stocked with bullfrogs, green frogs, or a combination of the two species revealed no consistent trends that were statistically significant (Table 26). Total numbers and volumes of phytOplankton were invariably higher in the mixed-species culture than in single- species treatments (Appendix 8, Table 823). In the mixed-Species treatment, total numbers and volumes of blue-green algae were higher than other algal taxa in this treatment throughout the experiment (Appendix 8, Tables 823 and 825). In aquaria with monocultured green frog or bullfrog tadpoles, total numbers and volumes fluctuated during the experiment with no discernible pattern (Appendix 8, Tables 823 and 825). The lowest concentrations of Dictyosghaerium, a green alga, were measured in aquaria with a mixed culture of bullfrogs and green frogs (Appendix 8, Table 826). The blue-green algae, Agmenellum and small Anacystis, and the green alga, Ankistrodesmus, were most abun- dant in the mixed-species treatment (Appendix 8, Table 826). Aquaria stocked with a single-species culture of bullfrogs had the lowest concentrations of Scenedesmus and volumetric evenness of all algae (Appendix 8, Tables 826-827). The total numbers and volumes of diatoms and the numerical diversity calculated for all algae were highest in aquaria with monocultures of green frog tadpoles (Appendix 141 Table 26. Phytoplanktonic comparisons between aquaria with monocultured bullfrogs (B), mixed-species: bullfrogs and green frogs (M), and monocultured green frogs (G) in Experiment Three at Lake One. EXPERIMENT THREE-~LAKE ONE 1. Volumes of Diatoms 5 October F I 8.00* 546158.7(8) 743654.3(3) 987585.7(6) 2. Numbers of Blue—greens 12 October F I 3.73 259644(8)(( 275384(G) 3. Volumes of Blue-greens 12 October F I 3.52 4231996(G) 4325810.7(8) 4. Total Numbers of Algae 12 October F I 10.33* 127685f3) 458322(G) 5. Numerical Diversity of Greens 12 October F I 4.07 1.307(5) 1.355(5) 6. Mean Sizes of Total Algae (Volume/Number) 12 Octooer F I 9.79* 65.2(M) 71.4(61 515349(M) 1.399(8) 76.5(3) 5.14 ' F05 (2.5) ' 142 8, Tables 823 and 827). In Experiment Three at Lake Four, the treatments, species compositions of tadpoles, effected consistent differences in responses among the algal assemblage (Table 27). The highest numerical and volumetric diversity of all algae was measured in aquaria with mono- cultured green frog tadpoles (Table 27 and Appendix B, Table 828). Moreover, the average sizes of all algae tended to be lowest in single-Species cultures of green frogs (Appendix 8, Table 824). The relative pr0portions of algae in the different taxa were similar among all treatments, although blue-green algae contributed a greater pr0portion to the total algal assemblage in aquaria with monocultured green frogs (Appendix 8, Table 825). Single-Species cultures of bull- frog tadpoles generally had the lowest numerical and volumetric diversity of diatoms and abundances of Nitzschia estimated for any of the tadpole treatments (Appendix 8, Tables 826 and 828). Community Metabolism Effects of Varying Tadpole Stocking Rates on Gross Primary Productivity (GPP)—and GPP/Autotrophic8iomass (Experiments One andTTwo) Lake One: Initial measurements of GPP were similar for all aquaria in Experiment One and EXperiment Two before tadpoles were stocked (Figure 14 and Appendix B, Table 829). Tadpoles influenced the gross primary productivity in aquaria, but the response of the autotrophic community was small and not simply related to the intensity of grazing. In both experiments, average values of GPP/day fluctuated among sampling dates, apparently in response to factors other than tadpole grazing since unstocked and stocked aquaria responded Table 27. 143 Phytoplanktonic comparisons between aquaria with monoc bullfrogs and green frogs (M), and monocultured green frogs ( ultured bullfrogs (B). mixed-species: G) in Experiment Three at Lake Four. EXPERIMENT THREE-ILAKE FOUR 1. Numbers of Greens 12 October F I 11.59’* 402(M) 554(6) 722(8) 2. Volumes of Greens 12 October F I 4.62 44958.3(61 47509.7(M) 108518(8) 3. Numbers of Diatoms 5 October F I 3.71 7366(6) 11069(M 12527(8) 4. Volumes of Diatoms 5 October F I 4.16 1852566(G) 3115517(3) 3185959 7(8) 5. Volumes of Blue-greens 5 October F I 5.42* 150110.3(8) 210551.7IM) 359387.7(8) 6. Total Volumes of Algae 5 October F I 4.44 2090741.7(G)(( 3542571.A(M) 3717774.7(B) 7. Numerical Diversity of Diatoms 5 October F I 3.76 0.056(M) 0.056(8), 0.121(0) 24 October F I 8.89* 0.023(8) 0.053(5) 0.097(8) Combined 3 Dates F I 7.21** 0.029(8) 0.045(M) 0.087(G) 8. Volume Diversity of Diatoms 5 October F I 7.72* 0.127(M) 3.130(8) 0.229(0) 24 October F I 6.76* 0.053(8) 0.176(M) 0.359(0) Combined 3 Dates F I 7.38" 0.072(8) 0.132(M) 0.241(6) 9. Total Volume Diversity 24 October F I 4.05 0.324(8) 0.185(M) 0.624(0) 10. Numerical Evenness 5 October F I 3.70 0.428(M) 0 458(9) 0.506(8) 11. Volume Eveness 5 October F I 6.77' 0.201(M) 0.257(6) 0.303(8) 24 October F I 7.95* 0.113(M) 0.137(8) 0.212(6) Combined 3 Dates F I 6.25** 0.171(M) 0.195(8) 0.254(6) Combined 12. Mean Sizes of Greens 3 Dates F I 3.25 44.8(51 93.9(M) 162.2(8) ? F.95 (2,5) I 5.14; F.05 (2.24) I 3.40 '* F.01 (2.6) I 10.90; F.01 (2.24) I 5.61 144 .Acee eeeegm u mewxeeem new; use .meewcem _eucerLe; new: Lee H mcwxeeem zew .eee eeeecmc: n eweezee eexeepmcev Leek eee eco mexee we ezw ece eeo meeeeweeexm Lew Azeweeeee\amev u mewv new we mucesecemeez NNiD 0.10 m1 h 0N-N .e_ et=e_e 0 lllllllllllli g, M vwx<4 meu u owln I T U I N - 1 fl l0 _ ”.32.. N axe efe ere 9e eeb we @ I t 22838.. :25... Rome «9:: .80 r .32.. Ex“ M M T I j mnuonbb/(bm)noaavo 60| 145 similarly (Figure 14 and Appendix B, Table 829). In Experiment One, both biomasses of stocked tadpoles consistently depressed the GPP; the low biomass stocking reduced the total GPP by 20% and the high biomass stocking reduced it by 1 %. In Experiment Two, the overall effect of tadpole grazing was to stimulate the GPP (Figure 14 and Appendix 8, Table 829). The GPP in aquaria with a low stock of tad- poles averaged 21% higher than in unstocked aquaria, while in the high biomass stocking, GPP averaged 17% higher than in controls. A major temporal shift in primary production occurred toward the end of Experiment Two. If the experiment had not been terminated, GPP in unstocked aquaria may have continued to decline below levels of GPP in aquaria with tadpoles. In both experiments, tadpoles increased the GPP/autotrophic (phytoplanktonic and periphytic) biomass (Figure 15). Although the total standing crap of periphyton was significantly lowered by tadpole grazing, a higher rate of primary production was maintained by the remaining periphyton and phyt0plankton than occurred in unstocked aquaria. The feeding activities of tadpoles may have removed dying or less healthy plants and increased the availability of nutrients to autotrophs. Lake Four: In Experiments One and Two, GPP measurements were similar among all aquaria before tadpoles were introduced (Figure 14 and Appendix 8, Table 829). within an experiment, levels of GPP varied among sampling dates in aquaria, regardless of treatment. Apparently, environmental factors influenced the primary productivity similarly in aquaria representing the different stockings, since trends in variation of GPP were alike in stocked and unstocked aquaria (Figure 14 and 146 .Aeee eeeezm u xeeum saw; eee .Lee eeeweum u meweeeeu meew eaeeeew we xeeum 3ew .cee eeeezmce u ewaeeee eexeeemcev teem eee ego mexee we ezw eee eeo mueeswaeexm cw Ae\mev :eeeee me eeaemees mmeaewe ewzeecaee=e\aaw .m_ ecemwa mu._.In..mwn. LAKE 4 EXID 2 pipiens = 3. shaded bars). unshaded bars, low biomass stocking of Percentages of GPP and R attributed to periphyton based on ancillary studies in Experiment Two at Lakes One and Four (jars in un- stocked aquaria striped bars, and high stocking Figure l8. 1555 Table 28. Contributions of periphyton and phytoplankton to the total GPP and R (02 in mg/l) based on ancillary studies during Experiment Two. LAKE ONE LAKE FOUR EXPERIMENT Low High Low High THO Control Density Density Control Density Density A. 7-8 August 1) Total GPP (02) 19.6 19.1 17.0 3.5 3.8 2.8 a) Phytoplankton 3.1 3.0 4.4 0.6 0.4 0.2 (+0 08) (+9.32) (:0 98) (:9 17) (_o 30) (+0.10) 0) Periphyton 16.5 16.1 12 6 2.9 3.5 2.6 (:3 7) (+2.6) (*2 8) (:0.5) (:p.7) (:9.5) c) GPP in Carbon/ Autotropnic Biomass (C) . 0.09 0.16 0.26 0.15 0.22 3.22 d) Periohytic s of GPP 34 34 74 33 90 93 2) Total RESP (02) 15.3 13.8 11.0 2.2 2.3 1 a a1 Phytoplankton 2.2 2.1 , 3.2 0.4 0.1 0.9 (_0.24) (:9.32) (30.98) (_p.061 f:p.;7} {:9.23) a) Periphyton 1332 11.7 7.3 1.3 1.8 0.8 (12.4) (30.7) (:j.5) (:9.2) (:0.5) (_9.3) c) RESP in Carbon/ Autotrcphic Biomass (C) 0.07 0.12 0.17 0.09 0.13 0.20 d) Periphytic 1 of RESP 86 85 71 82 81 47 3) Ratio-oPeripnytic (C): Phytoplanktonic (C) 32:1 19:1 11:1 95:1 69:1 50:1 3. 21-22 August 1) Total GPP (02) 0 14.4 13.2 3 3 5.2 3 9 [-l.42] a) Phytoplankton 9.5 10.4 10.8 0.3 0.3 0.2 (:1 5) (:2 11 (:2 0) (30.03) (:9 04) (:0.03) b) Periphyton C 4.0 7.4 3.0 4.8 3.7 [-10.9:5.2] (+2.0) (+1.4) (+1.1) (+0 1) (:0.8) c) GPP in Carbon/ Autotrophic Biomass (C) 0 0.17 0.37 0.34 0.48 0.53 d) Periphytic : of GPP 0 28 41 91 92 95 2) Total RESP (02) 25.2 25.0 30.6 3.5 5.4 3.7 a) Phytoplankton 13.3 14.2 14.4 0.1 0.2 ’ 0.2 (11.2) (+1.3) (11.4) (+0.03) (19.06) (:0.051 b) Periphyton 11.9 10.8 16.1 3.4 5.2 2.5 (+2 4) (:5 4) (:3 3) (:1 2) (:0 1) (+0.8) c) RESP in Carbon/ - Autotrophic Biomass (C) 0.15 0.30 0.33 0.36 0.50 0.51 d) Periphytic % of RESP 47 43 53 97 96 95 3) Ratio--Periphytic (C): Phytoplanktonic (C) 9:1 1:1 2:1 17:1 21:1 15:1 T1 r+ De Su 1.1. 1a 9X1 156 aquaria, 2.5 times that of the low biomass stocking and 4.5 times values for the high biomass stocking. The GPP/R ratios were zero in jars from unstocked aquaria and 0.58-0.59 in jars from stocked aquaria. The GPP/ R estimates were similar to GPP/"R“ measurements for aquaria after respiration contributed by tadpoles had been deducted. On this date, high concentrations of detritus, green and blue-green algae, about four times greater than recorded on earlier dates,were imported in the lake water. This abundance of seston in aquaria and jars apparently de- pressed periphytic production and elevated the relative contribution of phytoplanktonic production. Tadpole stocking in ancillary experiments consistently elevated the GPP/R, GPP/B and R/B ratios over values in unstocked aquaria. A comparison of differences in periphyton and seston among the three treatments demonstrated that the low biomass stocking of tadpoles reduced the ratio of periphytic to sestonic material by 40-56% and the high biomass of tadpoles reduced the ratio by 75-80%. Lake Four: Ancillary experiments at Lake Four revealed that periphyton, rather than seston and associated phytoplankton, contributed substantially (>80%) to the GPP in tests (Figure 18 and Table 28). The importance of periphytic production to the autotrophic community appeared proportionately greater at Lake Four than at Lake One. The jars from aquaria with a high biomass stocking of tadpoles invariably exhibited a lower ratio of periphyton to seston and a higher GPP/R ratio than those from unstocked aquaria. As at Lake One, GPP/B and R/B were highest in jars from tadpole-stocked aquaria in both ancillary tests. In all jar experiments at both lakes, tadpoles appeared to g:- 1131 Le 157 increase the ratio of community metabolism to autotr0phic biomass, but the greatest effects were observed at Lake One. Comparisons of Community Metabolism (Experiments One and Two) In both experiments at Lakes One and Four, gross primary productivity and community respiration were related (r = 0.74, P<0.0l) in all aquaria, regardless of treatment (Figure 19). Measurements of GPP were higher at Lake One than at Lake Four; the mean GPP at Lake One was 5.9 times greater in Experiment One and 5.5 times greater in Experiment Two than at Lake Four (Appendix B, Table 829). In the first experiment, similar biomasses of tadpoles were stocked at Lakes One and Four. At Lake One, tadpoles depressed the GPP, while at Lake Four, the GPP was increased by tadpoles. In the second experiment, higher bio- masses of tadpoles were stocked at Lake One than at Lake Four. Tad- poles elevated the mean GPP at Lake One and depressed the mean GPP at Lake Four. In stocked aquaria, mean values of ”R" (community respiration minus tadpole respiration) were lower in both experiments at Lakes One and Four than community respiration in unstocked aquaria (Figure 16, Appendix 8, Tables 830-831). Similarly, GPP/"R" ratios were higher in the two experiments at both lakes than GPP/R in unstocked aquaria (Appendix B, Table 832). On all sampling dates in both experiments at Lake One, measurements of GPP/autotrophic biomass were higher in tadpole- stocked aquaria than in unstocked aquaria (Figure 15). Values for R/B and "R"/B for aquaria with tadpoles were higher than estimates in un- stocked aquaria on all sampling dates in Experiments One and Two at Lake One (Figure 17). These elevated values were a result of tadpoles influencing the autotrophic biomass proportionately more than metabolism. 158 ...o.ova .¢~.o umpnscm AL. acmwupwmmoo :o_uo.mcgou. Lao. ace mco mmxm. an mucme_equm omegg asp :. mucoaummeu ..m com a new new :mmzumn a_;m=o.um_me age .m. mg=a_m 2.08.23: m.~v\ No 9:. 20.22am“... E22228 000. 00m 00m 005 com .OOn 00¢ 00m CON 00. . q . d 4 . q q a d . K ...... on... 00. 100m \. 1 com \ . . 1 cow . vv\ 1 com . . a\. 1 com o \- 1. CON (Uta/$181!1 9217/30 6111) ado \. 1 com \ 1 com a\ o 1.unXu. 1C F‘- CIT. 1 M1“. ~\w 159 At Lake Four, GPP/B and R/B were more variable during both experiments than measurements at Lake One (Figures 15 and 17). In the first experi- ment at Lake Four, tadpoles decreased both the GPP/B and R/B below values measured in unstocked aquaria. In Experiment Two, stocked aquaria, especially those with a high biomass of tadpoles, increased the mean GPP/autotrophic biomass over values in unstocked aquaria. Mea- surements of "R"/B were lower than R/B for unstocked aquaria until the final sampling date in Experiment Two, when R/B declined in unstocked aquaria below R/B in tadpole-stocked aquaria. Effects of Tadpole Composition on Community Metabolism (Experiment 11ml Lake One: The largest differences in gross primary productivity and community respiration occurred among aquaria before tadpoles were stocked (Figure 20 and Appendix 8, Tables B33-B34). Measurements of GPP and R were positively correlated (Figure 19) and the magnitude of temporal variation for GPP and R was alike. Bullfrog tadpoles, green frog tadpoles, and the mixed culture of the two species affected the GPP and R similarly; tadpole activities appeared to dampen initial differences in GPP and R among aquaria (Figure 20 and Appendix 8, Tables 833-834). For the three groups of tadpoles, the difference in average GPP was 0.5-5%; for community respiration, the difference among treatments was 3-6%. Only minor differences were noted in GPP/R values among aquaria and GPP/R ratios always exceeded unity in Experiment Three (Appendix B, Table 835). Although monocultured green frogs reduced the periphytic biomass more than monocultured bullfrogs or the combination of the two species, values for GPP, R and GPP/R were similar among treatments (Appendix B, Tables 833-835). GROSS PRIMARY PRODUCTIVITY ////////////// mm'o \\\\\\\\\\\\\ //////////// 5369.35.53? \\\\\\\\\\\\\ /////////////// 9.0233323“? \\\\\\\\\\\\\\\\ ///////////// 3:303:3503030' \\\\\\\\\\\\ ‘ WW 0" Ow. 6' W? :zozozezozziogzoy” \\\\\\\\\\\\\\\\\\\’ W 9”? b...’ M? 2.2.2.2.222. .~\\\\\\\\\\\\\\\\\\\\\\ /////////////////////1 1:." 35" o o“??’3:9' ’ \\\\\\\\\\\\\\\\\\\‘ I I l l W 55:35..” \\\\\\\\\\\\\\\ / 000.3%. .9.9’. 0.0.0. 160 E t: - 5“ “““““ 33; “F \; n'é'l' 10-(10-12) 10122-24) 9'122'24) IO‘(3‘51 wnuonbo/(bwmoeuvo 601 EXP 3 LAKE 4 EXP 3 LAKE I Gross primary productivity and community reSpiration in Experiment Three at Lakes One and Four (bars represent monocultured bullfrogs, mixed-species culture, and monocultured green frogs, reSpectively). Figure 20. Cw 161 Differences in GPP/autotrophic biomass and R/B were greatest among aquaria on the first sampling date after tadpoles were stocked (Figure 2l). At the end of the third experiment, GPP/B and R/B were similar in aquaria with monocultures of bullfrogs or green frogs and in the mixed-species cultures. Although the differences were relatively minor, GPP/B and R/B were invariably highest in aquaria with mono- cultured green frog tadpoles and lowest in aquaria with single-species cultures of bullfrog tadpoles. Lake Four: At Lake Four, measurements of gross primary pro- ductivity for Experiment Three averaged 5-6 times lower than at Lake One (Figure 20 and Appendix B, Tables 833-834). The GPP and R were positively correlated (Figure 19) and varied similarly between sampling dates in all aquaria (Appendix 8, Tables 833-834). Measurements of GPP were consistently lowest in aquaria with monocultured green frogs, averaging 14% below values in aquaria with single-species cultures of bullfrogs. The lowest community respiration was also associated with green frogs in monoculture, averaging 10% lower than R values measured in single-species cultures of bullfrogs. Measurements of GPP and R were usually intermediate between the two single-species treatments (Figure 20 and Appendix B, Tables 833-834). These trends in GPP and R were found throughout the experimental period, but differences among tadpole groups were small. Single-species cultures of bullfrog tadpoles stimulated the primary productivity and accumulation of autotrophic biomass more than the mixed-species culture or monocultured bullfrogs. The GPP and R were elevated by the activities of monocultured bullfrogs to the extent \ any. Q00 162 433.58%? .39: :33 3:338:05 EB .9538 mmzmaméme .mmoctpzn 83:3 ¢ ux<4 n mxm .VNINNTQ 8703-0. 3-3-0. .8 23:23 \ 8. 29:: Emma .. .8 232.23 18 a8 .982 “5.0.9.3.. 988 .58 EB was $3.3 an 8...: «55283 L8 Ex 23 Emma mo 8.53. .8 PEPE .ude nmxm Avmlumvug $706-0. PH“ me in Del (3?? low and egg l63 that GPP/B and R/B were highest in these aquaria, despite the increased autotrophic biomass (Figure 2l). As at Lake One, ratios of GPP/B and R/B were similar for all tadpole groups at the end of the experiment. Bullfrog tadpoles in monoculture also maintained GPP/R ratios at slightly higher levels than were measured in the other treatments, but the increases in GPP/R were minor (6% was the largest difference among treatments). The lowest GPP/R ratios were estimated in aquaria with monocultured green frog tadpoles (Appendix B, Table B35). Partitioning phytoplanktonic and_periphytic GPP and R (ancillary ‘tgstl: To assess the relative contributions of periphyton and seston with associated phytoplankton to the gross primary productivity and community respiration, a 48-h ancillary experiment was conducted (Table 29 and Figure 22). At Lake One, the contribution of periphyton to the community respiration was similar for the three tadpole groupings, but the portion of the GPP attributed to periphyton fluctuated among treat- ments. Although the ratio of periphytic to sestonic carbon was lowest in jars simulating conditions in aquaria with monocultured green frogs, periphyton in this treatment generated the largest proportion of the GPP (Table 29 and Figure 22). Periphyton in jars from aquaria with monocultured bullfrogs contributed the least to the GPP and had the lowest GPP/R ratios. The highest respiration estimated for periphyton and phytoplankton (seston) occurred in jars simulating conditions in aquaria with a single-species culture of bullfrog tadpoles. At Lake Four, the periphyton in jars accounted for all of the GPP and 89% or more of the respiration in all treatments (Table 29 and Figure 22). The highest GPP/R ratios were observed in jars from 16i4 eN 9:3,: .q=.aq. ~.c ...qfl. e.o 9.: .u. u.=a..:u_.=..;. ".9. o...=;.caa--c..u= ...nwz .3 u 9.1.3:“... 2. «,3 massaging U__.__o¢5o~=< \coageu c. smug Au :o.x;;.caa A; =aux:a_;o.>:; Ac .Nc. .m.z .uso. ..c .o a o...g;..a. .2 Au. mm=§:.a u.::o..o.=< \coacou e. ..c .o :cuxga..u; A: :euxca_:ouxsa Ac .No..am .a.o. A N .. concave v~-~m .< a:_.:: >13.” xcu—__u:u :5 :3 1.33. :\?= ... Na. x 3.2. . me. . am. . mm. .... ..a. an Na ma N. a. a=.o ~..c no.a ma.= m=.¢ ...QH. .m.qu. ...qq. ...ce. .e.qu. e.. ~.~ a.. a.. =.~ .No.qH. ..c.c.. .c=.qu. .~.:fl. .m.qfl. ~.A. N... .... a... s... 3.. c.~ ¢.. o.~ ..N no. :3. cc. as .q o=.c mo.c 93.: o..c ao=.o ..~.ou. .mm.om. .ec.¢m. ..c.cn. ...qq. m.. o._ m._ o.= m.o ..e.=u..c-. .a=.ow..c-. .c:.qu..a-. .~.:u. .~.Qfl. a a c q.= n.o 3.. m.. ~.. c._ o.. ”mac. mace. sauce maoc.__:a moo.. mass. :moco :omcu a maacb__=m :m~.c a awash—.3: "sax.z "sm‘.x use. .x<. was ..<. mac..__:= IIIIIIIII- I . uuz:_ hz.z_z.;x. II. 'II. IIIIIIIIII .222: ......=:...:.: 2:0 _u.3. 2:. :. :o.4=a_;:.a:; ::u ::.x;;_cu; .3 n::..:a...::u .ca 9.33— 165 \\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\ \\ \\\\\\\\\\\\\ IO‘I22‘24) WWW” NMMWMMWMWKWWMWM mmmmwwwwm... ..wummmmm ..mmWMMMu 7///////////////////////////////////A /////////////////////////////////// W... . WWW... 79 .30 G3 mmuc o .._.> In Ema /IO"I22" \2\4I LAKE 4 LAKEI EDIP’IB ton based on Percentages of GPP and R attributed to periphy ancillary tests in Experiment Three at Lakes One and Four (bars represent monocultured bullfrogs, mixed-species culture, and mono— cultured green frogs, respectively). Figure 22. 166 aquaria with monocultured green frog tadpoles. This elevated GPP/R from aquaria with single-species cultures of green frogs resulted from lower respiration by autotrophs. DISCUSSION Abiotic Environmental Effects Since abiotic factors like temperature, light, and water quality can have major impacts on the production of algae and algal consumers, the potential influences of these factors on production were assessed. Water quality, as measured by the concentrations of nitrogen, phospho- rus, and carbon, best explained the significantly higher primary produc- tivity at Lake One. Extremes in certain abiotic factors like oxygen and ammonia were more prominent at Lake One; however, tadpole survival and growth were invariably higher at this lake. Apparently, food quality and availability had major impacts on the realized tadpole production at the two lakes and abiotic factors had few quantifiable effects. Influences on Algal Communities in Aquaria Temperature fluctuations in aquaria during single experiments at Lakes One and Four were similar and within ranges that commonly occur in shallow ponds. The average daily temperatures were alike in Experiments One and Two, about 17 C, whereas in Experiment Three, daily temperatures averaged 8 C. Differences in photoperiod and cloud cover also occurred among Experiments One, Two, and Three at both lakes. Despite the varia- tions in these factors, gross primary productivity (mean values for all aquaria, whether stocked or unstocked) varied only 18% among the three experiments at Lake One and as little as 5% among those at Lake Four 167 S?” -I 168 (Appendix B, Table 368). Variations in community respiration among the three experiments at both lakes averaged 20 to 30% (Appendix B, Table 363). Water quality, as indicated by concentrations of phosphorus, nitrogen, and organic carbon, differed widely among the three experiments at Lakes One and Four, but generally without significant impact on com- munity metabolism. Values for net primary productivity, phytoplanktonic biomass, total organic carbon, total nitrogen, and total phosphorus were compared with natural lakes of different trophic status (as summarized by Vollen- weider 197D, Likens 1975, Wetzel 1975) in Table 30. Based on this classification, Lake One would be categorized as eutrophic for net pri- mary productivity. Lake Four was less productive and values for primary productivity corresponded to those measured in oligotrophic waters. Since temperatures and light were similar in aquaria at the two lakes during each experimental period, the differences in the availability of nutrients must explain the disparate levels of primary productivity at the two lakes (Table 30). Daily fluctuations of oxygen in aquaria were much greater at Lake One than at Lake Four and were caused by the higher photosynthesis and respiration of the algal standing crop. Algal productivity at Lake Four was about 14% of that at Lake One. Both lakes were moderately alkaline and the pH fluctuated little during the period from May to October. Influences on Tadpoles In general, the environments in aquaria at Lake One were more stressful than those at Lake Four; however, tadpole survival and growth 169 upcmzco umxuoumca Low mmwmswpmm co ummmm H coo..-OOm o~-N. o.-m .m.-mm .oo~-om. Lao. axe. omo.o.-oem mom.~-oae m.-. amm-mw. .OOm..-..m ago a... coo.m.x-oom coo.mx-om - - - 0.;Qoeuaacma.z - - om-m com A ooo.. x 0.;Qoepam ooo.Haoom omnofi . I . u_;qo.u=m-0mmz - - m-. v oom-oo. ooo..-cm~ 0.;QOLHOmaz coo-OmN o.-m - - - 0.;ao..0mas-om..o - - m-. v oo.-o~ oom-om 0.;aocpom..o omm-. v m-. v - om V cm v u...oc.om..o-m...= .me\z 95. .me\. as. ..\o as. .n\me\g as. .U\~e\u as. zmuomc.z mamozamoxa zomm=. >mv .Amumfi pmNamz use .mnmfi mcmxw. .ONoa mzumum u_;aogu “cosmewwu we mmxop _mgzam: saw: Lao. use one mmxm. Eo mcomwgmqeoo .om mpnoh In p-.- 1! la IS 170 were consistently higher at Lake One. Tadpoles at Lake One were sub- jected to extreme daily fluctuations in oxygen, especially in Experiment Two when oxygen ranged between 65 and 240% in relative saturation. The maximum range in relative saturation of oxygen (85 to 135%) at Lake Four occurred during Experiment Three. At both lakes, all aquaria had daily fluctuations in temperature and moderately high pH levels. Abiotic fac- tors, such as oxygen, pH, and temperature, may have influenced tadpole survival, the efficiency of conversion of food material into tadpole tissues, or other variables measured for tadpoles in this study. Yet, many of the shallow aquatic systems inhabited by tadpoles are stressful with fluctuating water levels; diurnally variable oxygen, temperature, and pH levels; and dense populations of aquatic plants. Variations in daily temperatures in aquaria during this study were similar to those measured in natural ponds inhabited by 3, catesbeiana, R, pipiens sphenocephala (18 to 36 C in June; Seale et al. 1975), and Bufo americanus (10.5 to 31.5 C in early July; Beisenger 1977). The influence of temperature in determining the growth rates of larval amphibians has been demonstrated in laboratory (Atlas 1935, Moore 1939, Ryan 1941a, b) and field studies (Calef 1973, Noland and Ultsch 1981). During single experiments at Lakes One and Four, temperature and photoperiod were alike at both lakes, so differences in growth and pro- duction exhibited by tadpoles were related to factors other than temper- ature and light. In the third experiment, average daily temperatures declined and the metabolism of tadpoles was probably lower than in the two earlier experiments. But values for growth per degree day, which should have minimized the temperature effects, were higher for individual tadpoles catesbeia tion) in bodies i reported 6% satur E. Clari commonly with oxy been obs: in oxyger oxygen t6 1.8 mg 02 Stubblefi L in ponds ‘ enriched 1 extreme (I On tadpole a]. 1973) and Under? and WM. 1964) Bro; H . “91.9mm, 171 tadpoles of R. pipiens in Experiments One and Two than those for R, catesbeiana and R. clamitans in Experiment Three. Minimum concentrations of oxygen (50 to 70% in relative satura- tion) in stocked aquaria were higher than concentrations in most water bodies inhabited by tadpoles. For example, Cole and Fischer (1977) reported mean dawn concentrations of about 30% relative saturation with 6% saturation (0.6 mg/l) near bottom in a pond with large populations of R, clamitans and R, pretiosa. Tadpoles of R. pipiens (=3, utricularia) commonly inhabit swampy areas where waters are less than 50% saturated with oxygen (Noland and Ultsch 1981) and tadpoles of 3, catesbeiana have been observed in littoral areas of ponds with daytime spatial variation in oxygen of 1 to 6 mg/l (Boyd 1975). In laboratory studies, critical oxygen tensions occurred at about 22% of relative saturation (1.7 to 1.8 mg Oz/l at 26 C) for unacclimated tadpoles of R. pipiens (Helff and Stubblefield 1931, Noland and Ultsch 1981). Little information is available on diurnal oxygen fluctuations in ponds with tadpole populations, but limnological data for shallow, enriched systems suggest the oxygen variation in such habitats may be extreme (Hetzel 1975). These fluctuations may have a moderate effect on tadpole mortality since ranid tadpoles develop lungs early (Just et al. 1973) and gape or gulp surface air (Wassersug and Seibert 1975). Moreover, tadpoles can alter their behavior (Whitford and Massey 1970) and undergo metabolic and physicochemical reorganization at the tissue and cellular levels in response to changing oxygen conditions (Barbashova 1964, Brown 1967, Rose and Drotman 1967, Seibel 1970, Bennett 1978). Weigmann (1972) and Weigmann and Altig (1975a, b) assessed the ability of tadpoles to acclimate to various oxygen concentrations and to employ 172 anaerobic glycolysis during hypoxia. Larvae of 3, pipiens were cultured successfully by Weigmann (1972) and Weigmann and Altig (1975a, b) under a cyclic oxygen regime with daily concentrations of oxygen ranging from 3 to 9 mg/l. In addition, leopard frogs were reared successfully from eggs into tadpoles at saturated (9 mg/l) and subsaturated (1.8 mg/l) oxygen levels. No large differences in pH occurred between Lakes One and Four during this study and diurnal fluctuations in pH within each lake were small because the moderately high concentrations of bicarbonate dampened variations. Total ammonia concentrations at Lake One invariably exceeded those at Lake Four, creating a greater potential for higher concentrations of toxic un-ionized ammonia at Lake One. The adverse effects of high ammonia concentrations (38 mg/l) on tadpoles in experi- mental ponds have been reported by Champ et al. (1971), but concentra- tions of this magnitude were not measured at either lake (the highest concentration of ammonia-nitrogen at Lake One was 5 mg/l). If pH and ammonia levels had differential impacts at the two lakes, tadpoles should have been more stressed by conditions at Lake One. Relationships Between Plant Production and Tadpole Production Apparent Impact of Tadpoles on Algae and Nutrients While nutrients were the key factors in determining the different levels of gross primary productivity at the two lakes, tadpoles played a major role in influencing the realized GPP in single experiments at each lake. The impact of ranid tadpoles on the aquatic environments in this study was a function of the trophic status of the lakes and the density and biomass of tadpoles stocked. This was obvious in the first 173 experiment when the same densities and biomasses of tadpoles were intro- duced at both lakes; the tadpoles at Lake One effected a maximum reduc- tion of 34% in the GPP, whereas those at Lake Four promoted a maximum increase of 50% in the GPP (see Appendix A for amended values of com- munity metabolism). During the second experiment, grazing pressure was initially lower at both lakes than in the first experiment, but lowest at Lake Four. In this experiment, tadpole grazing accounted for a maxi- mum increase of 15% in the GPP at Lake One and a maximum decrease of 25% in the GPP at Lake Four. Community respiration was consistently lower in the stocked aquaria than in the unstocked aquaria at Lake One, despite the added contribution of tadpole respiration. Conversely, community respiration was invariably elevated, as much as 54% higher in aquaria with tadpoles than those without tadpoles at Lake Four. Results from Lake One are similar to findings by Seale (1973, 1980) on tadpole populations in a small Missouri pond where tadpole grazing pro- duced either a 10 to 30% increase or reduction in the gross primary productivity, depending on the biomass of tadpoles present. Changes in the ratio of gross primary productivity to tadpole biomass were non- linear; the growth rates of algae were enhanced by low levels and depressed by high levels of tadpole biomass (Seale 1973, 1980). Based on these findings, data from Lake Four, particularly from the first experiment when the high abundances of tadpoles stimulated the GPP, were difficult to interpret. Leopard frog tadpoles grew poorly in this experiment, many died, and survivors were thin. Apparently, foraging activities and metabolism of the numerous tadpoles promoted this increased production by algae, but the mechanisms were not clearly discernible. Dc p'r al 174 Studies of other aquatic consumers of algae report either stimulation of primary productivity (Porter 1972, 1973, 1975, 1976, 1978, Lehman 1980, Lynch 1980) or more typically depression of primary productivity (Wright 1965, Hargrave and Geen 1970, Haney 1971, 1973, Porter 1977); as much as 100% of the phytoplanktonic production can be cropped daily by zooplankton. Grazers reduce autotrophic production mainly by decreasing the overall abundance of algae; however, extensive data exist on the influence of grazers on phytoplanktonic composition and diversity by which they may also affect primary productivity (reviews by Porter 1977, Porter and Orcutt 1980). Similar data for tadpoles are scarce; only Dickman (1968) and Seale (1973, 1980) have reported on algal-tadpole interactions. Dickman (1968) observed that caged tadpoles reduced the abundance of filamentous green algae and promoted the growth of diatoms and bacteria. Seale (1973, 1980) ana- lyzed changes in the ratio of chlorophyll a to phaeophytin during tad- pole grazing and proposed that tadpoles altered the structure of the phytoplanktonic community by decreasing the contribution of blue-green algae. In this study at Lakes One and Four, total phytoplanktonic numbers and volumes in stocked aquaria were generally increased two to four times greater than the abundances measured in unstocked aquaria. This proliferation of phytoplankton occurred in stocked aquaria at both lakes, whether algal concentrations in the lakes were high or low. Growth of phytoplankton may not account solely for these phenomenal increases in stocked aquaria since algae that had been ingested by tad- poles on the previous day may have been deposited as feces during the sampling period. Thus, algal cells in feces may have been resuspended and enumerated as living phytoplankton. All algae which were counted 175 and measured were readily identifiable and had intracellular structures intact, so phytoplankton appeared viable even if they had passed through the guts of tadpoles. I Although large increases in most phytoplanktonic taxa occurred in stocked aquaria, certain taxa increased more than others in relative abundance. One consistent trend recorded at Lakes One and Four throughout the experimental periods was significant increases in diatom numbers and volumes in aquaria with tadpoles. This happened whether or not diatoms were, at that time, a major component of the algal commu- nity. In particular, diatoms of the genus, Nitzschia, and to a lesser extent, Navicula, increased in stocked aquaria. Assuming that specific growth rates of the different algal taxa did not account wholly for the observed results and all genera were grazed in proportion to their abundances or non-selectively, then some algae were more resistant than others to passage through tadpole guts. Increases in phytoplanktonic numbers and volumes partially may be explained by the accrual of algal cells from the previous day's feeding, but the significant increases in abundance of particular genera indicate that algae were differentially digested and certain taxa may survive and perhaps benefit from gut passage. Porter (1973, 1976) reported that colonies of gelatinous green algae, like Sphaerocystis and Elakothrix, not only survive passage through zooplanktonic guts but incorporate nutrients, such as phospho- rus, into algal tissues during passage. More than 90% of these algal cells emerged from the grazers' guts intact and viable, resulting in a subsequent 63% elevation in the abundance of gelatinous green algae in the lake during the first 24 h after gut passage (Porter 1976). Labora- tory studies by Seale and Beckvar (1980) indicate that certain algal 176 genera (Chlorella) are less damaged than others (Anabaena) after pas- sage through tadpole guts and these taxa which are poorly digested will grow and reproduce when placed in algal nutrient media. Therefore, results of Porter (1973, 1976), Seale and Beckvar (1980), and those from this study suggest that certain taxa of algae gain a competitive advantage over others when grazers are present in aquatic systems. Although most evidence does not substantiate selective grazing on algae by tadpoles (Farlowe 1928, Savage 1951, 1952, Jenssen 1967), leopard frog tadpoles in this study may have consumed proportionately more of certain algal genera than others. This could be attributed more to certain algal characteristics and tadpole feeding behavior than to the active selection of algae by tadpoles. For example, diatoms have been shown to sink more rapidly than other algal taxa (Lund 1965, Uhlmann 1971) and diatoms may have settled in high concentrations on the bottom of aquaria. Also, diatoms appeared to dominate the periph- yton, adhering to the sides and bottoms of aquaria. Tadpoles in this study spent more time feeding on the sides and bottoms of aquaria than in the water column, thereby increasing the probability of their con- suming more diatoms. If diatoms were poorly digested by tadpoles and even benefited from gut passage or tadpole metabolites, then they would be expected to increase in relative abundance. The mechanisms under- lying the proliferation of certain algal taxa can only be postulated; however, the results are important because the composition of the phytoplanktonic community was altered by tadpole grazing. The observed increases in overall phytoplanktonic abundances in aquaria with tadpoles can be explained partially by the tadpoles' foraging primarily on periphyton and their influence on nutrient 177 cycling. Concomitant with the elevation of total numbers and volumes of algae in stocked aquaria was a decrease in the numeric and volumetric diversity of all algae. Since tadpoles promoted the proliferation of certain algal taxa more than others, they effected a lower evenness value for algae than was measured in aquaria without tadpoles. In addition, the data suggest that in early spring when blue-green algae were rare, as in Experiment One at Lake One, activities of tadpoles can enhance the growth of blue-green algae, particularly that of larger blue-greens. Apparently, the mean sizes of all algae may increase significantly with high grazing pressure, as occurred in the first experiment at Lake Four with high-stocked biomasses of tadpoles. Ranid tadpoles are typically categorized as suspension feeders; however, whether these suspensions are comprised of seston in the water column or are generated by materials scraped from substrates is influenced by characteristics of the aquatic habitat. Steinwascher (1978a, b, 1979a, b) proposed that the amount of time tadpoles spend scraping materials from substrates or filtering particles from the water depends on the quality and abundance of food. He investigated the effects of dispersed and concentrated food (Purina rabbit fodder) on the growth of three species of ranid tadpoles. Tadpoles exposed to clumped food resources spend more time scraping or rasping, whereas those exposed to less concentrated, dispersed food spend more time fil- tering. From these results, Steinwascher (1978a, b, 1979a, b) con- cluded that the energetic cost of scraping is greater than that of filtering for ranid tadpoles. Studies designed to analyze the filtering ability of tadpoles have shown that algal concentrations required for maximal ingestion are 178 7 high, greater than 2.0 x 10 um3/ml (Seale and Wassersug 1979, Seale and Beckvar 1980). The critical threshold or concentration of algae neces- sary for effective filtering by 3, sylvatica is 1.0 x 107‘um3/ml and for 6 5, catesbeiana, 1.0 x 10 um3/ml (Seale and Wassersug 1979, Seale and Beckvar 1980). Assuming these critical thresholds for filtering and ingestion by ranid tadpoles determined in the laboratory apply to tad- poles in nature, then ranid tadpoles may infrequently filter phyto- plankton since algal concentrations of this magnitude occur only during blooms in eutrophic waters. In this study, algal concentrations in water from enriched Lake 6 7 7 One were high, averaging 1.6 x 10 , 3.8 x 10 , and 2.2 x 10 umB/ml in Experiments One, Two, and Three, respectively. Algal concentrations in 6 water from Lake Four generally were lower (4.8 x 105, 2.5 x 10 , and 1.2 x 105 pm3/ml in Experiments One, Two, and Three, respectively) than required for filtering efficacy of ranid tadpoles. Of the 21 eutrophic lakes cited by Vollenweider (1970), only five had maximal algal concen- 7 trations averaging or exceeding 1.0 x 10 um3/ml. For a given lake to be categorized as eutrophic, Vollenweider (1970) suggested algal volumes 5 umB/ml. This concentration is much should approach or exceed 1.0 x 10 less than those proposed by Seale and Beckvar (1980) as critical thresh- olds for efficient filtering by ranid tadpoles. Few limnological studies have reported such high algal concentrations: maximum phyto- 7 planktonic concentrations of 2.6 x 10 umB/ml were reported by Baker and Baker (1976) in eutrophic Halstead Bay, Minnesota; in Lake Hashing- ton, enriched by sewage effluent, total algal volumes exceeded 1.0 x 107 um3/ml during only one month of a three-year study (Parker 1977); and phytoplanktonic concentrations in western Lake Erie averaged 179 1.4 x 107 um3/ml annually (Marcus 1972, Hocjik 1978). Although filter- ing may be less energy demanding than scraping or rasping materials from substrates, aquatic habitats where tadpoles occur may have low concen- trations of suspended particulates, below the critical thresholds required for efficient filtering. In this study at Lakes One and Four, periphyton was the major food consumed by ranid tadpoles. Results on tadpole production at Lake One indicate that tadpoles may benefit energetically from feeding off substrates in shallow-water zones of enriched environments. The pro- ductivity of periphyton in aquaria with tadpoles averaged 30 mg organic matter/dmZ/d at Lake One and at Lake Four, the periphytic accrual in unstocked aquaria averaged 1.3 mg organic matter/dmZ/d. Estimates at Lake One approach the highest productivities recorded for many aquatic environments (the highest productivities reported by Hetzel (1975) were in rich warm springs which averaged 73 to 97 mg organic material/dmZ/d). Measurements of daily periphytic growth at Lake Four are near the lowest productivities reported by Hetzel (1975); certain oligotrophic lakes had lower productivities, averaging 0.5 to 1.0 mg organic matter/dmZ/d. So periphytic accumulation rates were over 25 times higher at Lake One than at Lake Four, but overall GPP for the three experiments was only about ten times greater at Lake One. The tadpoles at Lake One greatly decreased the accrescence of periphyton and stimulated phytoplanktonic growth. At this lake, 78 to 87% of the periphytic carbon was diverted by tadpoles into tissue pro- duction or the accumulation of phytoplankton and detritus. A maximum of about 35% of the periphytic standing crop was converted daily into tadpole production and respiration. At Lake Four during the second 180 experiment, tadpoles reduced the periphytic standing crop by about 37% and more than 77% of the periphytic carbon was converted daily into tadpole biomass or respiration. .In the first experiment at this lake, the biomass of periphyton in stocked aquaria increased as though it were unavailable as a food source. Certain algae, like the diatom Cocconeis, are largely unavailable to grazers because they secrete a jelly-like substance that cements them to the substrate (Patrick 1948, 1970, Marcus 1980). Tadpoles, weakened by poor nutrition in Experiment One at Lake Four, possibly could not rasp such closely adhering periph- yton from the sides and bottoms of aquaria. Even in the second experi- ment at Lake Four, when tadpoles consumed attached algae, they removed a much lower proportion of the standing crop, but used it more effi- ciently than at Lake One. In these experiments, the influence of tadpoles on the peri- phytic communities exceeded their cropping of algal production. Tad- poles also altered the carbon-nitrogen ratios of the periphyton. At Lake One, C:N ratios of periphyton averaged 11:1 in unstocked aquaria, whereas the C:N ratio averaged 7.5:1 in stocked aquaria. Decreases in the C:N ratio reflect increases in the protein fraction of the algal biomass, generally by decreases in the proportions of cellulose (Russell-Hunter 1970). Other grazers of the littoral zone, like snails, have also been observed to increase the ratio of nitrogen to carbon of the aufwuchs and grow more rapidly when feeding on periphyton with low C:N ratios (McMahon et al. 1974, Hunter 1975). Tadpoles at Lake One apparently removed the unproductive, senescent portions of the periphyton and maintained the remaining algae in a healthy, actively growing state. Aufwuchs communities with low C:N ratios are generally 181 dominated by diatoms, bacteria, and blue-green algae, whereas those with high C:N ratios tend to have single-celled and filamentous green algae as dominants (Russell-Hunter 1970, McMahon et al. 1974). My cursory examinations of glass slides colonized by periphyton indicated that more diatoms and blue-green algae were present among the periphy- ton in aquaria with tadpoles than in unstocked aquaria; green algae appeared to be more common on slides from unstocked aquaria. Tadpoles and other consumers in the littoral zone of lakes and ponds may increase the rates of material cycling within this area, accelerate the transport of materials from the littoral to the pelagic zone, and increase the rates of transfer of materials to the sediments. These organisms appear capable of stimulating increased activity among autotrophs and heterotrophs at lower trophic levels, thus increasing the rate of decomposition processes and the metabolic activity of the system as a whole. Calef (1973) demonstrated that tadpoles of R. gyrgrg could generate high turnover rates of sediments and epibenthic algae. When the oligotrophic lake studied by Calef (1973) supported 31 tadpoles/m2 to metamorphosis, tadpoles at this density potentially could process 10 g/mz/d of dry sediments. Rapid gut passage times are characteristic of tadpoles (Savage 1952, Ueck 1967, Altig and McDearman 1975, Wassersug 1975) and most . invertebrates that feed on plants and debris (Berrie 1976). Large detrital particles may be broken down into finer detritus after passage through the alimentary canal of animal consumers, thereby accelerating a process that proceeds slowly by microbial activity. Surface sedi- ments are reused by benthic feeders, so all sediments in aquatic sys- tems may pass through the guts of indigenous populations several tinuas 182 a year (Brinkhurst and Chua 1969, Appleby and Brinkhurst 1970, Calow 1973, Berrie 1976). Both detrital and autotrophic pathways were influenced by the foraging and metabolic activities of tadpoles in this study at Lakes One and Four. Ranid tadpoles stimulated the accumulation of phyto- plankton and detritus and depressed periphytic growth, partially as a consequence of their effects on nutrient cycling, measured as the mobilization and relocation of organic carbon, nitrogen, and phosphorus. Under the extreme eutrophy characteristic of Lake One, tadpoles appeared to facilitate the movement of total phosphorus, mainly as particulates, and organic carbon, both dissolved and sestonic, into the water (which was removed daily from aquaria). Activities of tadpoles in the less enriched waters of Lake Four promoted the release of higher concentrations of dissolved phOSphorus and stimulated the storage of total organic carbon. Whether sestonic carbon decreased or increased in the water from aquaria at Lake Four depended on the extent to which it was consumed by tadpoles. Nitrogen dynamics were influenced by tadpoles at both lakes, but not in a consistent or predictable manner. Levels of inorganic and organic nitrogen measured in the water and the loss of nitrogen to the atmosphere either decreased or increased, depending on the abundances of tadpoles stocked and their subsequent production. In natural environments with similar grazing pressure simulated by the field experiments, tadpoles could consume over one- half of the periphytic standing crop daily. Nutrients not diverted into tissue elaboration and respiration by tadpoles could be exported to the pelagic zone as particulates and dissolved organics or recycled 183 within the littoral zone. Fecal material when sedimented would relocate nutrients to the benthic zone. Seale (1973, 1980) proposed that tadpoles controlled the rate at which nitrogen was assimilated by phytoplankton. She found that when tadpole biomass was highest, the total nitrogen ingestion rates and ammonia excretion rates by tadpoles equaled the nitrogen demand by phytoplankton. Studies on other algal consumers, mainly crustacean zooplankton but also fish, have shown that nutrient regeneration by these organisms can have an immediate and substantial impact on over- all nutrient budgets and on the nutrient demand of primary producers (Cushing and Nicholson 1963, Pomeroy et al. 1963, Marshall and Orr 1966, Hargrave and Geen 1968, Martin 1968, Butler et al. 1969a, b, Peterson and Lean 1973, Buechler and Dillon 1974, Ganf and Blazka 1974, Elwood and Goldstein 1975, Dugdale 1976, Jacobson and Comita 1976, Durbin et al. 1979). The population dynamics of algae and their pat- terns of species succession may depend just as much on the remineral- ization of nutrients as on algal mortality caused by grazers (Lehman 1980). For example, Lehman (1980) found that the epilimnetic zooplank- ton in Lake Washington supplied 10 times more P and 3 times more N to the surface-mixed layer during the summer months than entered the lake from all other sources. Algae in this lake wereF’limited (Edmondson 1970) and much of the calculated algal demand for P could be supplied by zooplanktonic populations during the summer (Devol 1979). Relationships Between the Ratio of the Supply of Food/Demand by_Tadpples and Tadpole Production At each lake, the biomass of leopard frog tadpoles stocked per aquarium was inversely related to their subsequent total production 184 (r = - 0.91 and - 0.85 at Lakes One and Four, respectivelY); however, when data from the two lakes were pooled, the relationship between stocked biomass and total production of tadpoles was much lower (r = - 0.52). The pooled data resulted in a poorer relationship between tadpole production and stocked biomass because patterns of tadpole sur- vival and growth were different at the two lakes. Tadpoles survived and grew better at Lake One, despite the potentially more stressful conditions there than at Lake Four. Higher food production at Lake One appeared primarily responsible for the better performance of tadpoles. If, as proposed, food availability was the major variable influencing tadpole performance at each lake, then the supply of food/ demand by tadpoles, measured as GPP/TB, in relation to tadpole produc- tion should best explain the findings at both lakes. The ratio of supply to demand was highly correlated with total daily production by tadpoles (r = 0.80) and the daily growth of individual tadpoles (r = 0.98) at the different densities and biomasses tested in field experi- ments. Both total (TP/d) and individual (TP/TB/d) production by tad- poles were greatest when the ratio of supply to demand remained high throughout the experiment, as at Lake One in Experiment Two. In order to examine the general applicability of this relation- ship between tadpole production and food supply/demand by tadpoles, results from laboratory studies were combined with those from the field experiments and reanalyzed. The values of TP/TB/degree d (A-1 and A-5, the two extremes of density) for R, pipiens reared in the laboratory on cultured phytoplankton and the TP/TB/degree d measured in field experi- ments in relation to the supply of food/demand by tadpoles were plotted in Figure 23. The correlation between the food supply, whether based 185 1.2. 1'1' r = 0.93 1.0. O = LAKE 1 O . o = LAKE 4 0.9« e = LABORATORY 0.8J O .7. 0.6. 0.5. (L4. o I TP/TB/DEGREE d x 10'2 1C... 0:6 ' ore j 130 2.6 2.8 1 I 0 0:2 0T4 FOOD SUPPLY/TAD POL E DEMAN D Figure 23. The relationship between tadpole production (mg C)/tadpole stocked biomass (mg C)/degree d and the supply of food (mg particulate organic C/d or GPP as mg C/d) per unit of stocked tadpole biomass (mg C) for all experiments at Lakes One and Four and the laboratory study (A-1 and A-5 at 500 ml, 1,000 ml, and 1,500 ml, respectively). 186 on particulate organic carbon measured in the laboratory or GPP in the field, per unit of demand (carbon biomass of tadpoles) and tadpole production remained high (r = 0.93), despite the different test con- ditions in the laboratory and field studies. Thus, tadpoles at the densities (laboratory, 13-120/m2 or 263-2368/m3 for 3. pipiens; field, 22-220/m2 or 117-1051/m3 for _R_. pipiens, 5. clamitans, and 5. catesbeiana) and biomasses (wet weight, 9-36 g/m2 or 186-770 g/m3 in the laboratory and 3-109 g/m2 or 54-573 g/m3 in the field) stocked effectively divided the food resources among individuals. The main response of leopard frog tadpoles to crowding or high densities was slow growth caused by the lower food availability per individual. Low densities of tadpoles with abundant food grew fastest in contrast to findings reported by Wilbur (1977) of a "pollution effect" which reduced growth of tadpoles. Biomasses of leopard frog tadpoles stocked did not influence survival except, indirectly, by lowering the amount of food available per individual. Whether the surviving biomass or yield of tadpoles would exceed or be less than the initial biomass depended on the stocking intensity (TB/GPP), expressed as the ratio of biomass initially stocked to the average gross primary productivity (Table 31). At both lakes, low stocking intensities resulted in higher ratios of surviving biomass of 3, pipiens to GPP than high stocking intensities. Neither exploitive competition, interference competition, nor social and behavioral interactions, as observed by other investi- gators (Rose 1965, Akin 1966, Pourbagher 1968, 1969, Wilbur 1977, Steinwascher 1978a, b, 1979a, b, Dash and Hota 1980), appeared to be operating consistently in Experiments One and Two and the laboratory study. 187 mcpxuoam mpoaumu Lmumm acmepgmaxm Lma mount mcwpqum moss» ms“ Low new came oumcmmmcxrnv mcwxooum mpoacca mgoema cwgzmams mwgmacm PPM cw new maesm>m mpmacm new megw:_ N H He.mH mm.eH om.e~ mm.e m~.e em.e aau\mmeeewm epeeeae mew>P>e=m o~.o~ me.m~ mm.- um.e me.e mm.e aem\mmeeepm apogee» emceeem mo.- mo.e~ m~.NN Ne.e am.e me.e emu _awewew\mmee°wm e_eeeee emceeem some zmuao omtz acadss=m womb zmumu .mmmma .mmmmmmmm “maze ezmzemmaxm we.“ NH.H mm.H ¢~.~ aee\mmeeewm apogeee mew>_>e=m NN.m No.“ mo.~ Rm.o aew\mmeee_m mPOQee» eaxoeem Nm.~ so.“ em.e me.H ego Pameeep\mmeee_m apogee» eexeoem ozp ezmzmmmexm me.m Ne.m mN.N mo.H eam\mmeeewm apogeee me_>w>e=m -.mfi mm.m mm.~ he.“ aeo\mmeeewm mPeeeeh eexeoem H~.- mfi.m NH.- efi.m new _eeepev\mmeeewm apogeee emceeem mzo ezmzcmmexu x0 m H Q m mupkm Ia“: xoppm gnu “so; M¥ N .Lzou new mco moss; an mucmepgmaxm mugs“ asp Lee . eeo\mmmsown mpoaumu umxooum .A new mepwcw\mmmsown opaque“ umxooum .Amsv coaxed cw mew new mmmsopa amu\mmmsoma mpoaumu m=w>p>gzm new New wepewmeeeew me_xeeem .Hm e_ea» 188 In the third experiment, the biomasses of surviving tadpoles of bullfrogs and green frogs/GPP were greater at both lakes than antic- ipated, based on results from the two earlier experiments on the effects of stocking intensity on leopard frog tadpoles (Table 31). Since GPP was not appreciably elevated in Experiment Three at either lake, the high biomasses of surviving tadpoles in the third experiment can best be explained by the cooler water temperatures and concomitant reduced metabolic rates of tadpoles, which promoted increased survival despite the high stocking intensities. Tadpole production for the monocultures of bullfrogs and green frogs in the third experiment was related to food availability as in earlier experiments with leopard frog tadpoles (Figure 23). In the mixed-species culture, pooled production values for tadpoles of the two species corresponded to the ratio of food supply/demand; however, when the growth of 3, clamitans and B, catesbeiana was analyzed separately, green frog tadpoles performed better and bullfrog tadpoles performed poorer than expected, based on results from single-species cultures. Observations of tadpoles of these two species in natural habitats and information reported from laboratory studies have indicated that tadpoles of bullfrogs should grow better than those of green frogs when these species occur together. Steinwascher (1978a, b, 1979a, b) found that tadpoles of 3, catesbeiana reingest their feces, especially during low abundances of particulate food. Large tadpoles more efficiently exploit the feces of small tadpoles and outcompete them for this food source. Tadpoles of 3, clamitans are non-coprophagous and feed mainly on materials rasped from substrates, not particles in the water column as bullfrog tadpoles do (Steinwascher 1978a, b, 1979a, b). 189 In this study at Lakes One and Four, tadpoles of 3, catesbeiana, because of their larger size, could have had a slight competitive advan- tage over those of 3, clamitans (exploitive competition). As proposed by Steinwascher (1978a, b, 1979a, b), the clumped resource, represented by periphytic algae in this study, was reduced more by monocultured green frogs than bullfrogs; however, concentrations of particulate material, specifically phytoplankton, were not depressed more by bull- frog tadpoles. Tadpoles of bullfrogs and green frogs used the food resources similarly; when reared together, no measurable partitioning of food into suspended and attached components or differential resource utilization by tadpoles occurred in Experiment Three. Food, mainly periphyton and settled material, ingested by tadpoles of B, clamitans and 5, catesbeiana was like that used by 3, pipiens. Apparently, mono- cultured bullfrogs at both lakes used the available nutritional resources more efficiently than monocultured green frogs, since these tadpoles produced more biomass than those of green frogs. Data on algal production and nutrient concentrations from mixed-species cul- tures do not provide explanations for the higher production by green frog larvae, The smaller tadpoles of green frogs interfered with foraging and feeding by bullfrog tadpoles and the activities of bull- frogs increased the food available to green frogs. These results sug- gest that green frog tadpoles in natural environments may perform better in mixed-species populations with bullfrog tadpoles than in single-species populations. Bullfrog tadpoles, instead, may grow more rapidly when they occur in single-species populations. Tadpoles of bullfrogs and green frogs have not been reported to partition aquatic habitats by food, time, or space. Explanations for the different 190 performances of tadpoles of bullfrogs and green frogs singly and in combination will require additional investigations of these two species in natural habitats. Ecological Efficiencies of Tadpoles Net primary productivity (NPP), which is the excess of photo- synthetic production remaining after respiration has taken place, is a better indicator of the availability of food to grazers, like tadpoles, than gross primary productivity (GPP). Although at least 90% of the GPP of algal salt marsh communities may appear as net production (Pomeroy 1959), crops and natural vegetation, including phytoplankton, more typically respire about 20 to 40 per cent of the gross primary productivity (Hargrave 1969, Whittaker 1970, Wetzel 1975, Cole and Fisher 1978, Moss 1980). Respiration can account for as much as 70% of the GPP in certain plant communities (Kira and Shidei 1967, Golley 1970, Whittaker 1970). Accurately estimating net primary productivity of plants con- taining a few cells, which have a relatively short life span in aquatic environments, is difficult (Golley 1970, Barnes and Mann 1980, Moss 1980). In natural waters, estimates of algal respiration may be biased by the respiratory contributions of fungi, protozoa, microscopic zooplankton, and especially bacteria (F099 1980). At Lakes One and Four in this study, bacteria were not abundant in water samples or on membrane filters, so their respiration should not have exceeded about 20% of the total respiration. Leakage of organic compounds, like glycollate and peptides, can also influence estimates of net primary Productivity. F099 (1966, 1968, 1971, 1977) suggested that estimates 191 of NPP may have to be increased by as much as 40% to account for these extracellular compounds. However, the amount of organically combined carbon which is lost from algal cells relative to the amount of total carbon fixation is inconsequential in eutrophic waters, amounting to less than 1% (F099 1980). Ecological efficiencies (TP/NPP) of ranid tadpoles feeding upon algae were calculated by assuming that NPP was 20, 60, and 100% of the GPP (Table 32). Based on the correlation between respiration and gross primary productivity for the three experiments at both lakes, realized net production by autotrophs at Lakes One and Four probably ranged between 20 to 60% of the GPP. At both lakes, the highest total produc- tion by tadpoles and ecological efficiencies of tadpoles feeding on algae were observed in Experiment Two (Table 32). Values for ecological efficiency during this experiment were 6 to 32%, assuming leopard frog tadpoles fed only on live algae and NPP was 20 to 60% of the GPP. Based on these same assumptions, in Experiment Three at Lake One, all groups of tadpoles, single-species cultures of bullfrogs and green frogs and mixed-species cultures, were more efficient in using algae than leopard frog tadpoles at both lakes during the first experiment. However, the efficiencies of bullfrog tadpoles were almost twice those of green frog tadpoles feeding on live algae (Table 32). Ecological efficiencies for ranid tadpoles consuming algae are comparable to those of other grazers on primary productivity in aquatic systems. Energy transfers from primary productivity to secondary pro- ductivity for herbivorous benthos ranged from 1 to 14%; for herbivorous plankton, ecological efficiencies have been reported as 0.5 to 60%, but 192 m.m~ m.m m.m m.~m ~.mp m.e Am=a_eume eev & seeewesccm _.~_ e.o m.e o.~p o.m m.e Aeeeec. _eeoev & soee_o_ceu m.e_ m.mp m.e. N.m_ N.m_ ~.m_ eo_ge=eoee e_oaeep _esoe ~.mm_ o.mmm ~.o~m e.omp m.em_ m.m~m e_nap_a>e eeee Peeoe ..Nm P.~m ..Nm P.Nm ..Nm ..Nm “Lone? .ep_euma e.~m m..e_ ..mmm m.me m.e._ e..e~ su_>_se=eeee seee_ea pez uzo mxe eoee _eeoe m.mm m.mm m.am m.mm m.mm m.mm peeee_ Passageo m.mm o.mpp e.em_ c.5m _..P_ N.mmp snw>wpe=eeee seeeLee eez uzo m¥<4 - mzo Lzmzfizmaxm sow Roe woo. sow New goo. goshm Ia”: xoohm 3o; mezmzsmuaxm oofi x amw\aaz mcwsammm .Amcmwmnmmpmo .m.c:e .mcepwsmpu .m .mcmwm_m .mv mmpoaumu Lo mmwo:o_uw$wm pmuwmopoom .Nm mpnMH .maw mg“ we xoofi Lo .00 .ON m_ an: 193 m.m~ e.m P.m m.m_ N.e N.m Am=p_epee eev & seee_e_ecw m.e e.m m.m N.m N.N _.~ Aeeeece Feoepv N soeawescem m.. m.. m._ o._ o._ o.. eeLeeseeLe e_oeeas Pepop e.e~ m.em e.ee ~.e~ m.um m.me m_eepwe>e coo» Posse m._m m._~ m._N m._N m.PN m._m Beeee_ _ee_epeo _.m N.m. m.m~ e.m N.e_ o.- spw>wse=eoea seae_ee eez «so; m¥<4 - ozp Lzmzsmmexm N.~_- “.m- e.m- e.e m.~ m.o Amapsesme oev N seee_o_ecm N.e- m.~- ..N- m.o e.o m.o Aeeeece Feoeuv N seee_o_ccm e..- e..- m.P- m.o m.o m.o eowpeseeee apogee“ _eHOL m.mm m.em m.m~ m.mm m.me m.~e e_eas_a>a eeec _apeh o.m~ o.m~ o.m~ o.m~ o.m~ o.m~ eeeee_ _aeweueo m.m m.N~ m.ee m.e m.o~ m.mm sowswuezeeee seee_ee pez mace m¥<4 - uzo Lzmzsmmexm New New goo. sow Noe goes xuopm zufiz xooem 204 ooH x maw\¢az mhzmzmammxm .A.u.u:ouv mm mpnmh 194 n.5u m.Ni ©.Fi <¢.o: ¢¢.oi ¢¢.oi ~.m o.mp m.mm m.m N.F m.o o._ m.p ©.~ o.m¢ N.om_ m.mmm N.mi n.oi «.0: e_.oi up.oi ¢_.oi m.o ¢.mp ¢.Nm o.m N.F 0., N.N N.N N.~ m.m¢ c.—mp ¢.mpm o.oi «.0- P.oi eo.oi eo.oi co.o: m.© m.mp m.mm 5.0 m.m v.— N.m N.m N.m m.me P.Nmp _.mmm N xeeeLe_Lcm :owuuznoca mFoaumh >u_>wuu:uoca acmsweq amz anon mx<4 i mmmzh hzmzmzmaxm N seee_o_cem cowaoauoea mpoqumh »u_>_uo:voga xgasmcq umz mzo mx<4 i mmmzh hzmzfimmmxm Row Row Noop Row Row goop aom goo xoop mace» :mmem vocsupzuocoz mo_omamicmxwz mmogmppsn umeau_=oo:oz 00H x amw\¢¢z mhzmszmmxm .A.e.eeeev mm «_eee 195 estimated efficiencies of 17 to 20% appear most reliable (Brylinsky and Mann 1973, Moriarty et al. 1973, Gulati 1975, Lam and Frost 1976, Lehman 1976, Pederson et al. 1976, Coveney et al. 1977, Makarewicz and Likens 1979, Hughes 1980, McCauley and Kalff 1981). Values of ecological efficiency for planktivorous herbivores are influenced by the trophic status of the aquatic environment; ratios of herbivorous production to primary production are highest in oligo— trophic waters and decrease with increasing eutrophy (Pederson et al. 1976, McCauley and Kalff 1981). The quality, which includes algal size and manageability, chemical composition and nutritional value, and toxicity among other attributes, as well as the quantity, expressed as dry matter, organic material, carbon or energy per unit area, of primary productivity determines its utilization within food webs of aquatic systems (Boyd 1970, 1971, Iverson 1974, Schroeder 1977, Onuf et al. 1977, McNeill and Southward 1978). In eutrophic waters, nannoplankton (10 to 50 pm; Strickland 1960) are less abundant and many algal planktivores consume algae mainly in the nannoplanktonic range (Glicwicz 1967, 1977, Porter 1973, 1977, Nadin-Hurley and Duncan 1976). Because of their unmanageability and possible toxicity (toxin production varies among different strains of the same species; Carmichael and Gorham 1977, 1978), large filamentous blue-green algae, like Anabaena flos-aquae, are a low quality food for filter feeders. such as zooplankton (Porter 1973, 1977, Webster and Peters 1978, Lynch 1980, Porter and Orcutt 1980). Although these filaments are filtered and ingested at the lowest rates by zooplankton, they may be assimi- lated at rates and efficiencies comparable to other algae that are con- sidered good food sources (Arnold 1971, Lampert 1977). Algal 196 macroconsumers, like brown bullhead catfish, non-selectively graze large quantities of filamentous algae (Cable 1929, Rubec 1975); how- ever, the nutritional quality of different algal taxa affects the efficacy of digestion and assimilation by bullheads. For example, Gunn et al. (1977) have demonstrated that blue-greens (A, flos-aquae), which have a crude protein content of 48%, are assimilated more effi- ciently by brown bullheads than filamentous green algae like Spirogyra sp., with a crude protein content of 22% (A/I% for A, flos-aquae, 48 to 68%; and for Spirogyra sp., 24%). Thus, using total NPP rather than ingested NPP may produce inaccurate estimates of ecological efficiency for consumers. In this study at Lakes One and Four, ecological efficiencies of ranid tadpoles were based on net autotrophic production of both phytoplankton and periphyton. Since tadpoles foraged predominantly on periphyton, the tadpoles' use of attached algae, representing only a portion of the NPP, may more accurately depict their ecological efficiency. There- fore, efficiencies of tadpoles calculated for total NPP may be under- estimates. Most suspension feeders and scrapers in aquatic systems are omnivorous, not strictly herbivorous, since they consume protozoa, detritus, and bacteria as well as living plants. Because both bac- teria and detritus may be a major food source for consumers, estimates of NPP may not adequately represent energy availability and use. Even though the standing stock of bacterial biomass in an aquatic ecosystem may not be as great as that of phytoplankton, periphyton, and macro- phytes, the bacterial contribution to secondary productivity can be substantial because the bacterial biomass may be replaced several times 197 each day under optimal conditions (Sieburth 1976, Watson et al. 1977). The production of microorganisms sometimes exceeds the production of phytoplankton, especially in shallow water-bodies, where most of the macrophytic production becomes detritus and a high input of alloch- thonous material occurs. Pomeroy (1980) cites studies that demonstrate as much as 90% of plant production goes to detritus in many aquatic systems. Detrital material rapidly loses proteinaceous compounds and becomes less nutritious to consumers (Berrie 1976). However, the microbial biomass associated with the decomposing detritus may increase the detrital protein or nitrogen content, making detritus an important energy source (Odum and de la Cruz 1967, Kaushik and Hynes 1968, 1971, Hynes et al. 1974, Heal and MacLean 1975). Microorganisms may fulfill a dual role by converting detritus into microbial tissues which are directly ingested and by catalyzing the partial degradation of refrac- tory materials comprising detritus into substances that can be used by animal consumers (Barlocker and Kendrick 1974, 1975a, b, Berrie 1976). Because ranid tadpoles in natural habitats are omnivorous and may feed on bacteria and detritus, derived from plants, feces, or car- casses of other animals, ecological efficiencies were calculated for tadpoles consuming both algae and detritus (TP/(NPP + detrital carbon)). Like TP/NPP, the highest ecological efficiency of ranid tadpoles on algae with detritus was measured in Experiment Two at Lakes One and Four (Table 32). Ecological efficiencies of tadpoles presumed feeding on algae and detritus were lower than efficiencies calculated for tad- poles consuming living algae at both lakes, but especially at Lake Four where detrital carbon comprised a large portion of the available food. This was obvious in Experiment Two when ecological efficiencies 198 (TP/(NPP + detritus)) at Lake Four (3 to 5%) were less than half of those at Lake One (7 to 12%). Detritus appeared to be less nutritious as a food than living algae at both lakes, but least nutritious at Lake Four. Ratios of C:N for detritus were high at Lake Four where much of the detrital material probably was derived from macrophytes, rather than algae as at Lake One. Macrophytic tissues have unfavorably high C:N ratios (average C:N greater than 17:1, Russell-Hunter 197D, McMahon et al. 1974) and resilient structural elements (van Soest and Wine 1967, Boyd and Goodyear 1971, Polisini and Boyd 1974) that make them less likely to be grazed than most algae (average C:N less than 6:1, Wetzel 1975). The correlation between ecological efficiency (TP/(NPP + detritus)) and stocking intensity, denoted as the ratio of stocked tadpole biomass to NPP with detritus (TB/(NPP + detritus)), was lower (r = - 0.75) than the correlation obtained between stocking intensity and NPP without detritus (r = - 0.80). Although monitoring energy and material flow through producers and consumers will remain a major theme in ecology, present techniques for measuring energy transformations are oversimplifications of energy relationships in natural ecosystems (Steele 1970, Turner 1970, Edmond- son and Winberg 1971, Slobodkin 1972, Grodzinski et al. 1975, van Dobben and Lowe-McConnell 1975, Wiegart 1976, Cooke 1977). For these reasons, Pomeroy (1980) has suggested that energy transfers between producers and consumers should be measured as gross growth efficiency (growth/ingestion) and consumers should not be strictly categorized as, for example, herbivores, detritivores, or carnivores. 199 Growth and Assimilation Efficiencies Assessing energy flow in specific ecosystems by designating consumers as exclusively belonging to one of three categories--herbi- vores, omnivores, or carnivores--is misleading and inaccurate since many aquatic organisms are opportunists and more catholic in their dietary intake. However, this tripartite method of cataloging con- sumers as feeding primarily on plants, animals, or both has provided a conceptual framework for integrating anatomical structures and physio- logical processes in a diversity of taxa that feed on similar resources. For example, Welch (1968) reviewed reported values for growth and assimilation of aquatic herbivores, omnivores, and carnivores and observed that gross or ecological growth efficiencies (G/I or P/I) and assimilation efficiencies (A/I or P/I) were generally higher for carni- vores than for herbivores. Tissue growth efficiency or net growth efficiency (G/A or P/A), in contrast, appeared to be slightly higher for herbivores. Welch (1968) suggested this evidence indicated an obvious nutritional difference between carnivores and herbivores. Carnivores digest and absorb more of their dietary intake at a high metabolic cost, whereas herbivores ingest larger quantities of less digestible foods. According to Welch (1968), omnivores occupy an intermediate position, possibly obtaining more of their protein require- ments from animal tissues and their energy requirements from plant tissues. Recent data on aquatic consumers discussed below generally substantiates the relationships that Welch (1968) proposed between diet and efficiency. In this study at Lakes One and Four and in the laboratory, ranid tadpoles under semi-natural conditions appeared to convert 200 primary productivity and organic carbon into tissues as efficiently as invertebrates and vertebrates feeding on similar foods. Ecological growth efficiencies (TR/ingestion) for leopard frog tadpoles measured in the laboratory, represented as G/I% since mortality was negligible, averaged 10.2%, 16.3%, and 23.0% at the 500 ml, 1,000 ml, and 1,500 ml algal supply rates, respectively. By assuming that leopard frog, bull- frog, and green frog tadpoles in field experiments ingested 20 to 60% of the GPP, similar ecological growth efficiencies can be estimated. At Lake One, leopard frog tadpoles stocked at low levels had efficien- cies of 2.5 to 32.5% and those stocked at high levels had efficiencies of 0.8 to 29.3%. At Lake Four, tadpoles of R. pipiens in the low stocking had efficiencies of 1.5 to 18.5%, while those in the high stocking had ecological growth efficiencies that were negative to 25.5%. In the third experiment, gross growth efficiencies of bullfrog and green frog tadpoles averaged 1.2 to 6.7% at Lake One and were nega- tive at Lake Four. Assimilation efficiencies for leopard frog tadpoles fed algae in the laboratory were 23.8%, 34.4%, and 42.8% at the 500 ml, 1,000 ml, and 1,500 ml algal supply rates, respectively. In the laboratory, assimilation efficiencies were more than two times greater than growth efficiencies and perhaps would have shown a similar relationship in the field experiments. Tissue growth efficiencies (growth/assimilation) for leopard frog tadpoles supplied with phytoplankton were estimated as 42.6 to 53.9%, whereas those fed TetraMin had higher efficiencies, averaging 55.4 to 75.5%. The only data on tadpoles available for com- parison are those of Altig and McDearman (1975) for assimilation effi- ciencies and Nagai et al. (1971) for ecological growth efficiencies. As th (‘1'- (I) tat rat for aci and - fOOd attac 201 Assimilation efficiencies for R, catesbeiana ranged from 8 to 25% and those for R. heckscheri averaged 54% (Altig and McDearman 1975). Other tadpoles (Bufo woodhousei, Acris gryllus, and Gastrophryne carolinensis) tested by Altig and McDearman had higher efficiencies than the ranid tadpoles. Their efficiencies averaged 77 to 86% on a diet of Purina rabbit chow. Ecological growth efficiencies of 10 to 87% were estimated fOr Bufo vulgaris on different mixtures of grain, fish meal, amino acids, and dead tadpoles (Nagai et al. 1971). Comparisons of data for ecological growth, assimilation, and tissue growth efficiencies of aquatic consumers reported in the litera- ture are difficult, since efficiencies are influenced by the quantity and quality of the food supply, laboratory and field test conditions, age or stage of development of the organism, and many other abiotic and biotic factors. For example, rates of food passage through the guts of aquatic consumers like zooplankton and tadpoles are influenced by the quality and quantity of available food; organisms exposed to low levels of food process materials to feces less rapidly than those exposed to abundant food (Wassersug 1975, Porter 1978). Thus, consumers exposed to abundant food may feed superfluously and exhibit higher growth and assimilation efficiencies when foods of similar quality are less abun- dant (Corner 1966, Corner et al. 1967, Butler et al. 1970, Penderson et al. 1976, Porter 1977, Pomeroy 1980). Data from Lakes One and Four and the laboratory experiments indicate growth and assimilation effi- ciencies of tadpoles are influenced by the quality and quantity of food resources. When food was not highly concentrated as at Lake Four, ranid tadpoles appeared to utilize the available food, primarily attached algae, more efficiently than at Lake One. de C0171 whe 202 The efficiencies of invertebrates feeding on live plant material, detritus, bacteria, or a combination of these foods are highly variable; ecological growth efficiencies (G/I or P/I) were reported to range from 2 to 75% and tissue growth efficiencies (G/A or P/A) reportedly ranged between 6 and 99% (Richman 1958, Wright 1958, Corner 1961, 1972, Soro- kin 1966, Hargrave 1970a, b, c, Winberg 1971, 1972, Ladle et al. 1972, Lasenby and Langford 1972, Monakov 1972, Saunders 1972, Edington and Hildrew 1973, Edmondson 1974, Nilsson 1974, Otto 1974, Anderson and Grafius 1975, McCollough et al. 1979, Benke and Wallace 1980). Values averaging 7 to 15% for ecological growth efficiency, 45 to 70% for tissue growth efficiency, and 7 to 50% for assimilation efficiency appear to be most common. Gross ecological efficiencies of about 8% were measured for the common carp, Cyprinus carpio, feeding on detritus (Kevern 1966), whereas the grass carp (Ctenopharyngodon idellus) had gross efficiencies of 1.9% and net efficiencies of 14% when fed aquatic plants (Fisher 1970). Gross growth efficiencies for carnivorous fishes are higher than for omnivorous or herbivorous fishes, averaging 13 to 37% for salmonids (Warren and Davis 1967, Brett et al. 1969, LeBrasseur 1969, McCormick et al. 1972, Shelbourne et al. 1973, Brett and Shelbourne 1975, Biette and Green 1980) and 13 to 44% for other fresh-water and marine fishes (Ivlev 1945, Pandian 1967a, b, c, d, Birkett 1969, Edwards et al. 1970, Gerking 1971, Cowey and Sargent 1972, Klekowski et al. 1972, Wissing 1974, Nakashima and Leggett 1980). Assimilation efficiencies for carnivorous fishes ranged between 67 to 86% (Davies 1963, Wissing 1974) and the efficiency of protein assimilation is higher than 90% (Menzel 1960, Pandian 1967a, b, c, d, Birkett 1969). Fishes that are more onn gre tei fil pro pid: deti rang Droc (Bra in l and and and asse and tadp item so t tion Simi‘ Omnix Clenc nin Tad; haVe 203 omnivorous, like channel catfish, have assimilation efficiencies greater than 80% on plant protein and greater than 90% on animal pro- tein (Hastings 1969). Other catfish, like brown bullheads, assimilate filamentous algae at efficiencies betwen 24 and 67%, depending on the protein content of the algal taxa ingested (Gunn et al. 1977). Tila- pids (Sarotherodon mossambicus = Tilapia mossambica), which consume detrital material, have remarkably higher assimilation efficiencies, ranging from 37 to 46% (Bowen 1979). Tissue growth (net growth or production) efficiencies in carnivorous fishes range from 24 to 54% (Brett et al. 1969, Gerking 1971, Klekowski et al. 1972); average 91% in larval newts of Triturus helviticus and I, vulgaris (Avery 1971); and in the salamanders, Plethodon cinereus (Burton and Likens 1975) and Desmognathus ochrophaeus (Fitzpatrick 1973a, b), are 50 to 80% and 76 to 89%, respectively. Gross or ecological growth efficiencies of ranid tadpoles assessed in this study were higher than those of most invertebrates and herbivorous or omnivorous fishes. In contrast to invertebrates, tadpoles are larger and can consume a broader size range of dietary items. More importantly, tadpoles are immature and do not reproduce, so the energy they ingest and assimilate goes toward tissue elabora- tion, not reproduction. Assimilation efficiencies of tadpoles are similar to those of invertebrates and higher than efficiencies of omnivorous fish like the common and grass carp. Tissue growth effi- ciencies of tadpoles feeding on algae are higher than those of car- nivorous fishes and within the range of efficiencies of invertebrates. Tadpoles when fed a nutritious, easily digestible food like TetraMin have higher growth efficiencies than those of tadpoles feeding on in pel the com thes nitii in tl Delac 1963 Uoku trap tain 204 algae; efficiencies for TetraMin resemble those of herbivorous or omnivorous invertebrates measured under optimal conditions. Based on these comparisons, ranid tadpoles in protected environments effi- ciently process materials and convert them into tadpole tissues. Potential Roles ofATadpoles in Natural and Wastewater Treatment Systems In aquatic systems, tadpoles frequently inhabit shallow, inshore areas with fringing plant communities, commonly designated as littoral zones. Although limnological investigations were initiated in Europe over 100 years ago (Elster 1974), these concentrated on the pelagic area and its associated phytoplankton; only more recently, in the last 15 years, have other plant associations like fringing plant communities in lakes and ponds been studied intensively. Results from these studies have demonstrated the importance of fringing plant commu- nities comprised of macrophytes and microphytes to aquatic ecosystems in that they may (1) produce more carbon or calories annually than pelagic phytoplanktonic communities in the same water body (Straskaba 1963, Efford 1967, Hargrave 1969, Allen 1971, Pierczynska 1972, Dokulil 1973, Hutchison 1975, Talling 1975, Teal 1980); (2) serve as traps or sinks for nutrients during periods of the year, making cer- tain nutrients largely unavailable to pelagic autotrophs (Chamberlain 1968, Confer 1972, Lee et al. 1975, Prentki et al. 1979, Teal 1980); (3) release high concentrations of dissolved organic materials and phosphorus by leakage during photosynthesis (McRoy et al. 1972, Wetzel and Manny 1972, Titus et al. 1975, Twilley et al. 1977, Smith 1978, Carpenter 1980, Teal 1980) or during decomposition (Solski m pl to te mm per Con V83 205 1962, Nichols and Keeney 1973, Otsuki and Wetzel 1974, Carpenter and Adams 1978, 1979, Godschalk and Wetzel 1978a, b); and (4) inhibit growth of phytoplankters by reducing the availability of light through shading or by releasing allelopathic compounds (Strangeberg 1963, Boulder 1969, Brandl et al. 1970, Allen 1971, Dokulil 1973, Teal 1980). The influence of littoral zones on ecosystem metabolism extends beyond the primary and secondary production attributed to plants and animals of this area and may include contributions of nutrients and organic materials to augment production of phytoplankton, suspended baceteria, and pelagic and benthic consumers. Conversely, fringing plant commu- nities, at least for certain periods of the year, may accumulate nutrients and organic material, thus potentially decreasing phyto- planktonic and microbial production in the open waters of lakes. For tadpoles, littoral zones may serve as sources of abundant food with epibenthic and epiphytic communities comprised of algae, bac- teria, protozoa, and associated detritus. Vascular macrophytes may be a major component of fringing plant communities, but these generally are not grazed directly by tadpoles or other aquatic consumers (Savage 1952, Odum 1957, Gajecskara 1969, McMahon et al. 1974, Hutchison 1975). Teal (1980) has reviewed data on productivity of fringing plant com- munities that shows the contribution of microphytes, specifically periphytic and epibenthic algae, can vary from insignificance to being considerably more important than macrophytic production. Although vascular macrophytes when reduced to detrital material are more likely to serve as food for aquatic consumers (Barlocker and Kendrick 1975a, b, Berrie 1976), living macrophytes, as well as providing substrates for aufwuchs which are grazed, create a structurally complex habitat, 206 not found in the pelagic zone. Wiens (1970, 1972) has demonstrated that ranid tadpoles select particular habitats in the laboratory which are consistent with their ecological distributions in the field. These behavioral responses of tadpoles to habitat alternatives are geneti- cally controlled, not a learned response (Wiens 1970, 1972). The importance of structurally diverse environments in reducing the frequency of physical interactions among crowded leopard frog tad- poles has been investigated by John and Fenster (1975). They found that the growth of tadpoles reared at high densities was directly related to the frequency of physical encounters among tadpoles; how- ever, growth rates were not determined simply by physical space but, rather, by "psychological space" not mediated by vision. Crowded tad- poles grown in aquaria with partitions simulating a structurally diverse environment did not exhibit the "crowding effect" that the same density of tadpoles exhibited in aquaria without partitions. The importance of social and behavioral factors in affecting the growth of tadpoles has been established by Goetsch (1924), Adolph (1931a, b), Rugh (1934), Lynn and Edelmann (1936), Hodler (1958), Gromko et al. (1973), and Smith-Gill and Gill (1978). If, as suggested, social and behavioral factors limit tadpole production in the laboratory, these same factors may be relatively unimportant in natural habitats because of the structural complexity of littoral environments. Ranid tadpoles grazing in fringing plant communities swim little and use the abundant filamentous algae and macrophytes as physical supports to maintain their position in the water column. Ranid tadpoles do not possess the morphological adaptations of certain open-water tadpoles like hylids and xenopids (Porter 1972), so the 207 energy required in swimming or maintaining a stationary position in the water should be higher for ranid tadpoles than for pelagic tadpoles specifically modified for filtering seston. In addition to support, littoral vegetation may provide refugia for tadpoles from predation by more accomplished swimmers like fish. Although quantitative data of fish predation on tadpoles in lakes and ponds are scarce, predation has been cited as the primary factor controlling tadpole populations in nature by Herreid and Kinney (1966), Calef (1973), Licht (1974), Heyer et al. (1975), and Heyer and Muedeking (1976). Shallow, inshore areas are characterized by warmer temperatures than are deeper waters and tadpoles have been observed to orient to thermal gradients, choosing temperatures within the optimal range for their growth and development (Beisenger 1977). Rapid growth of tadpoles to large sizes has been shown by Heyer, McDiarmid, and Weigmann (1975) to increase their probability of survival, since they may have outgrown certain verte- brate and invertebrate predators. Results at Lake One from experiments which excluded grazing (unstocked aquaria) and simulated the effects of tadpole grazing in littoral zones with extensive standing crops of attached algae but without predation (stocked aquaria) were similar to those observed by Confer (1972). An open-water system comprised of ZOO-liter aquaria continuously supplied with water and radioactive phOSphorus (32F as H3P04) was designed by Confer (1972) to represent phosphorus circu- lation in small lakes with extensive littoral zones during summer stratification. As in aquaria at Lake One, Confer's (1972) micro- ecosystems approximated lakes with a large ratio of littoral area to open water and with abundant attached algae, without sediments or loss 208 of materials to the hypolimnion. In systems without grazers, both in Confer's (1972) aquaria and unstocked aquaria at Lake One, the net movement of phosphorus was out of the open-water phytoplanktonic community to the attached filamentous algae and associated organisms. Phosphorus was removed from the open water by incorporation into plant tissues of littoral algae, but more importantly, major amounts of phos- phorus were trapped as particles by the attached algae associated with the sides and bottoms of aquaria. Trapping of particulate phosphorus by fringing plant communities was not an artifact of the micro-ecosystem design, since this occurs in natural aquatic habitats (Chandler 1937, 1939, Chamberlain 1968). Grazers, whether snails or ostracods intro- duced by Confer (1972) or tadpoles stocked in experiments at Lake One, reduced the standing crop of attached algae and promoted the increase of phosphorus, especially in particulate form, in the open water. Snails and ostracods, like tadpoles, significantly increased the pro- liferation of phytoplankton in the epilimnion. Projections on the potential role of tadpoles in nutrient up- take and relocation in ecosystems with high standing crops of fringing plants can be made from observations at Lake One. In enriched systems where food is abundant and predators scarce, minimum densities of 100 tadpoles/m2 appear reasonable. At this density, small tadpoles, averaging 200 mg (wet weight) per individual, could remove over 150 mg C/mZ/d and about 4 mg P/mZ/d. Larger tadpoles, averaging one gram, when present at the same density as small tadpoles, could ingest periphytic algae at the rate of 750 mg C/m2/d and 21 mg P/mz/d. Sub- stantiation of these estimated rates of organic carbon and phosphorus removal by tadpoles is provided in laboratory studies by Seale (1973, 209 1980). She found that ranid tadpoles consumed filamentous algae (esti- mated C:N:P ratio of 40:7:1, Wetzel 1975) at the rate of 48 ug N/g (wet weight of tadpoles)/h. Estimates presented by Seale (1973, 1980) when used to assess nutrient removal by tadpoles in nature indicated that at densities of lOO/mz, ranid tadpoles averaging 200 mg could consume 144 mg C/m2/d and 3.4 mg P/mz/d and those averaging one gram could ingest 720 mg C/mZ/d and 17 mg P/mZ/d. Aquatic ecosystems with low primary production like Lake Four, where GPP averaged about 140 mg C/mZ/d and standing crops of micro- phytic fringing plants were correspondingly low, could support about 10 to 25 ranid tadpoles/m2. Nutrient removal by tadpoles stocked at these densities would be about 10 to 25% of those estimated for eu- trophic waters like Lake One. In comparison to Lake Four, oligotrophic Marion Lake which purportedly could support 31 (3, 323953) tadpoles/m2 (Calef 1973) had mean annual values for primary productivity of 180 mg C/mZ/d with epibenthic algal productivity averaging 109 mg C/mz/d (Efford 1967, Hargrave 1969, Gruedling 1971). Assuming differences in the biology of these two ranid species in Marion Lake and Lake Four were unimportant, the higher densities of tadpoles estimated in Marion Lake probably were related to the higher production by epibenthic algae, which averaged almost twice that of attached algae at Lake Four. Expected yields, as wet weight, carbon, and phosphorus, of ranid tadpoles can be assessed for productive waters like Lake One and less productive waters like Lake Four. From eutrophic systems, about 127 g/m2 (5 g C/m2 and 150 mg P/mz, assuming a C:P ratio of 35:1, Mackenthun 1969, Durbin et al. 1979) of live tadpoles could be harvested per month, if tadpoles (about 200 mg wet weight per 210 individual) were stocked at 20 g/m2 and these grew at the maximum rates measured in field experiments (TP/TB/d of 0.176 in Experiment Two). In ecosystems like Lake Four with tadpole production as in Experiment Two (TP/TB/d of 0.033), 5.4 g/m2 or 212.4 mg cm2 and 6.4 mg P/m2 could be harvested monthly when 5 g/m2 of tadpoles (averaging 200 mg/individual) were stocked initially. These potential rates of nutrient removal by tadpoles and harvest of tadpoles may not be realized in natural environ- ments, whether eutrophic or oligotrophic, since they are based on extrapolations of data obtained from studies of tadpoles in protected environments. But the similarities in estimates of nutrient removal by tadpoles in these different studies indicate that ranids may serve as "harvestable" nutrient concentrators in wastewater treatment facilities where abiotic and biotic variables can be controlled to maintain an "optimal" (as yet undefined) environment for tadpoles. If as shown in experiments at Lakes One and Four that ranid tadpoles consume attached microphytes primarily, then more efficient systems for culturing tad- poles can be devised than those simulated by aquaria in this study. For example, shallow systems with high ratios of substrate-to-volume, like modified raceways, could enhance the growth and production of attached algae to be used as food for tadpoles. Tadpoles are not the only consumers of algae and detritus in most water bodies; other organisms in the benthic, littoral, and pelagic zones may influence the quantity and quality of food available to tad- poles. Grazers like zooplankton which mainly filter seston in the pelagic area probably would not compete directly with populations of ranid tadpoles for food and, instead, may benefit from the activities of tadpoles. Evidence for this has been found in experimental systems 211 where higher survival and reproduction of cultured zooplankton occurred with the inclusion of tadpoles (Kurt Kearns, pers. comm.). An explanation for the better performance of zooplankton when cultured with tadpoles may be that grazing by tadpoles relocated nutrients to the open water and favored the accumulation of phytoplankton and detritus, making more seston available to filter-feeding zooplankters. Conversely, 200planktonic grazing and metabolic by-products hypotheti- cally may increase the availability of food to tadpoles, either, directly, by ameliorating growth of periphytic and epibenthic algae through increased water transparency and nutrient availability or, indirectly, by promoting the settling of high concentrations of ses- ton, as algae or detritus and fecal pellets with associated microbes. Results from general studies on the ecology of pelagic zooplankton and ranid tadpoles indicate the two taxa interact in a mutually beneficial manner and can be successfully cultured together. However, this evi- dence is circumstantial and additional data are required from experi— ments specifically designed to elucidate the interrelationships between pelagic zooplankton and ranid tadpoles. Other consumers in the littoral and benthic areas that graze substrates as their exclusive feeding mode could compete with tadpoles for limited food resources. Chydorid cladocerans (Meyers 1980), cer- tain insects (Cummins 1975, Merritt and Cummins 1978), and other inver- tebrates like snails (McMahon et al. 1974, Hunter 1975) inhabit fringing plant communities and consume foods similar to those eaten by tadpoles. Also, settled seston and epibenthic algae are ingested by benthic infauna (Brinkhurst 1974), amphipods (Hargrave 1970a, b, c, 212 Mathias 1971), isopods (Johannes 1964, 1968, Prus 1971), and other organisms (Berrie 1976). Tadpole interactions among conspecifics and congenerics may be as important in the performance of tadpole populations as competition between tadpoles and various invertebrate taxa which utilize similar food and space resources. Despite a plethora of investigations on intraspecific and interspecific competition among tadpoles (reviews by Steinwascher 1978a, b, 1979a, b), none of the theories proposed have been universally accepted by researchers. Frequently the ecologists who design laboratory or field studies of ranid tadpoles are concerned with testing mathematical models of competition and know little about the ecology or life history of the species they employ. Thus, results and conclusions based on these experiments concerning the presence or absence of competition may not apply to tadpole populations in dynamic and structurally complex natural habitats. Interpretations of findings from studies on the influence of intraspecific and interspecific competition on tadpole populations are further complicated by the conflicting results reported by different researchers testing the same species of anuran. For example, tadpoles of the wood frog (3, sylvatica) and leopard frog (R. pipiens) have been reared singly and in mixed-species cultures by Wilbur and Collins (1973), DeBenedictis (1974), and Smith-Gill and Gill (1978). Conclu- sions as to the importance of intraspecific and interspecific competi- tion in these anuran taxa differ widely among the three studies. Smith-Gill and Gill (1978) after conducting laboratory investigations (densities of both species corresponded to 500 to 16,000 tadpoles/m3) proposed 3, pipiens was more sensitive to densities of R, sylvatica 213 than conspecifics and the effects of R, pipiens on R, sylvatica were negligible. Similar studies by DeBenedictis (1974) in pond enclosures (tadpole densities of 165 to 1,6OO/m3) indicated that tadpoles of _R_. sylvatica exhibited lower growth and survival when cultured with R, pipiens, but tadpoles of R, sylvatica had no effect on the growth and survival of R. pipiens. Growth and survival by both wood frog and leopard frog tadpoles in the study by Smith-Gill and Gill (1978) were phenomenally high. At the highest densities of each species (8,000/m3) in mixed-species cultures, 84% of the leopard frog and 98% of the wood frog tadpoles metamorphosed. When species were reared singly at den- sities corresponding to 16,000/m3, 89% of the R, pipiens and 98% of the R. sylvatica metamorphosed (Smith-Gill and Gill 1978). DeBenedictis (1974) found tadpoles of both species grew and survived better in enclosures with conspecifics rather than with congenerics; however, maximum survival of either species in any density treatment was only 25% and typically was much lower. Smith-Gill and Gill (1978) observed an "Allee effect" result- ing in the highest growth of conspecific R, sylvatica at densities of 4,000 tadpoles/m3; growth at SOD/m3 and 16,OOO/m3 was lower for tad- poles of this species. In contrast, enclosure experiments in ponds by Wilbur and Collins (1973) produced evidence of intraspecific competi- tion and slower growth 0f.3- sylvatica at densities (5,000/m3) similar to those where an "Allee effect" was observed by Smith-Gill and Gill (1978). Results from these studies (Wilbur and Collins 1973, DeBene- dictis 1974, Smith-Gill and Gill 1978) and other studies reported in the literature (Steinwascher 1978a, b, 1979a, b) on ranid tadpoles indicate the difficulties encountered when attempting to apply 214 experimental findings to populations in nature. Discerning the effects of interactions among tadpoles is relevant to understanding the ecology of tadpoles in natural habitats and their potential per- formance in wastewater treatment systems. To achieve this, results and conclusions from previous studies must be reinterpreted and syn- thesized into hypotheses which, when tested, yield results applicable to tadpole populations in natural and managed environments. In a recent review, Odum (1980) contrasted the efficacy of terrestrial and aquatic methods of harvesting and demonstrated that harvesting natural populations of aquatic plants and animals is more costly and energy-demanding than harvesting terrestrial organisms. As a means of improving aquatic harvests of natural populations of plants and animals to meet increasing food demands by human populations, Odum (1980) suggested the selective harvesting of organisms belonging to lower trophic levels and designing techniques to concentrate organisms more efficiently to approach a net energy balance (energy expended during harvesting/energy obtained from the harvest). Although these improved methods may yield more food at a lower cost, aquaculture was proposed by Odum (1980) as the most promising technique to create a significant food production and processing industry, analogous to agriculture, with reasonably predictable and controllable yields. Dramatic increases in current aquacultural yields may be achieved by (1) devoting more area to culture, (2) employing more herbivorous species, (3) farming several species with different trophic positions in the same enclosure (polyculture), (4) using greater densities of organisms, and (5) increasing the feeding and breeding efficiency through genetic selection. Organisms selected for aquaculture should 215 meet certain criteria, such as having simple life histories so they can be reared through all life stages in captivity; be hardy, resis- tant to diseases, and adaptable to crowding; and occupy a low trophic position, either a herbivore or detritivore, which produces a high biomass per unit area without expensive, (animal) protein-based feed (Odum 1980). Results from this study at Lakes One and Four indicate that tissue production by ranid tadpoles in intensively managed wastewater treatment facilities may approach values reported for production of herbivorous fishes such as carp, tilapia, and milk fish, in fertilized ponds in Israel and the Far East (5,000 kg/ha/y, Bardach et al. 1972). However, the desirability of tadpole protein and monetary profits derived from the sale of tadpoles are likely not to be as high as that for fishes, since most humans would not accept tadpoles as suitable food. Indians in Mexico harvest and consume tadpoles, but ranid tad- poles, potentially grown at low cost in wastewater facilities, would be used primarily as a protein supplement for fish or livestock feeds. Evidence from Lake One demonstrates that tadpoles can be raised on natural crops of algae, are hardy, and adaptable to crowding. But ranids have complex life cycles and maintaining adults which require live animals or, at least, motile food (can be force-fed) is labor intensive and expensive (Culley and Gravois 1971, Culley and Meyers 1972, Culley 1973, Modzelewski and Culley 1974, Smalley 1978). Presently, a breeding program for bullfrogs is underway at Louisiana State University in indoor facilities where this species develops from -eggs into frogs (227 g) in 10 to 12 months (Culley 1973). To insure that supplies of eggs are available when required for scheduled 216 stocking in wastewater treatment facilities, clutches of ranid eggs could be purchased from culture facilities (Nace 1968, Hirschfeld et al. 1970, Culley and Gravois 1971, Priddy and Culley 1971), collected from ponds and lakes when natural populations of ranid frogs are spawning, or obtained from adults previously captured in the field, which later were injected with pituitary extracts or hormones to induce egg and sperm production (Nace 1968). 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Ecol. 56: 229-243. lu_::seou 8:8 888 :8 888888 888858888 8883 .u\8ewu_. w.~e\u msv 888888 888888 888888889 ..< 8.888 246 88.. 8... 88.. =8=\888 88.. 88.. 88.. 88888 ..888 8.888 8.888 888 88888888 ..88 8.88 8.88 8888.88888888 888 88888888 8.888 8.888 8.8.8 8888.88 .88888 8.888 8.888 8 ..8 8888.88 88888888 8..88- 8.888- 8.888- 888\88.888.8888 8.88. ..8.8 8.88. 8888888 8.888 8.888 8 888 .88 .8888. 888888 8.88888 .8.8.8. 888 8888 -- 88. .888.88888 88888.8 88888.8 88888.888 88.8 888 ...8.8888. .8 8.88. 247 mo.~ mo.p mo.o =m=\¢gc om.o om.o mm.o «\aao m.ee ~.mm ¢.- mam acmgoaa< «.mp p.m m.¢ . mumswummgmnca mac ucmgmag< o.opm m.m_m m.p_m mucmpmn szau< N.mom N.o~m m.mpm mucmpmn umuumaxm m.m¢ . m.Pe - m.~m . xmv\:owamgwammm m.~m m.om ~.mm xmn\aaw m.—~m m._~m m.F~m Ame peanuv :ongmu uwcmmgo pmwupcm «no; ux<4 .. uzo hzmszmmxu mm:oo van cuzocc .Nm mpnmh 252 Table 83. Comparisons of periphytic carbon between unstocked (C for control) and tadpole-stocked (L for low and H for high biomass of .3. pipiens) aquaria. I. EXPERIMENT 0NE--LAKE ONE A. Periphytic Carbon (mg/aq) 6 June 1976 F = 54.12*** 243.1(L) 261.1(H), 815.6(C) 12 June 1976 F = 30.53*** 403.4(L)' 468.2(H) 1422.8(C) 19 June 1976 F = 31.38*** 538.8(L) 618.4(fl) 2004.0(C) B. Production (mg C/aq/d) 75 Dates F = 11.14*** 25.8(L) 28.0(H) 95.5(C) II. EXPERIMENT 0NE--LAKE FOUR A. Periphytic Carbon (mg/aq) 4.69 65.7(Q) 82.5(L) 120.0(H) 6.14* 74.1(C) 107.7(L), 169.2(H) 12 June 1976 F 19 June 1976 11 II III. EXPERIMENT TWO--LAKE ONE A. Periphytic Carbon (mg/aq) 8 August 1976 F = 20.53** 435.1(H) 543.6(L) 1294.2(C) 15 August 1976 F = 16.92** 227.6(H) 324.1(L) 1660.3(C) 22 August 1976 F = 12.52** 302.4(H) 875.6(L) 2372.0(C) 8. Production (mg C/aq/d) 'XéDates F = 36.48*** 12.1(H) 28.0(L) 89.8(C) IV. EXPERIMENT Tw0--LAKE FOUR A. Periphytic Carbon (mg/aq) 22 August 1976 F = 5.85* 76.0(H) 153.2(L) 188.6(0) *F.05(2’5) = 5.14 **F 01(2’5) = 10.90 ***F.00](2’6) = 27.00; F.001(2’24) = 9.34 253 Tab1e B4. Concentrations of dissolved and toral organic carbon (mg/1 : SO) in imported lake wateg. unstocxed (control) aquaria, and stocked (3. pipiens) aquaria during Experiments One and we. Sampling Lake Low High Dates Readings Control Density Density LAKE ONE-oEXPERIMENT ONE A. DISSOLVEO ORGANIC CARBON 6-(5-6)-1976 7.0 7.5 7.4 7.5 (:0.06) (=o.23) (30.14) (:0.31) 6-(11-12)-1976 6.8 7.1 7.5 7.8 (:0.09) (:0.03) (:O.15) (:0.08) 6-(18-19)-1976 6.8 7.7 7.5 7.4 ($0.05) (20.13) (:0.20) (:0.17) 15 Dates 6.9 7.4 7.5 7.6 (:O.C4) (:O.10) (:0.04) (:0.06) 8. TOTAL ORGANIC CARBON 6-(5-6)-1976 7.9 8,0 8.1 8.2 (:0.04) (:O.23) (:0.12) (:0.03) 6-(11-12)-1976 8.1 7.5 8.1 8.5 (:0.07) (:0.26) (:0.08) (:0.06) 6-(18-19)-1976 7.8 8.3 8.3 8.4 ‘(:0.05) (:O.39) (:0.20) (:0.13) 75 Dates 7.9 7.9 3.2 8.4 (:0.06) (:0.12) (:0.05) (:0.05) LAKE FOUR--EXPERIMENT ONE A. OISSOLVEO ORGANIC CARBON 6-(5-6)-1976 5.9 6.1 6.1 6.0 (:0.15) (:0.19) (:0.15) (:0.23) 6-(11-12)-l976 6.6 6.6 6.9 6.8 (30.15) (:O.15) (:O.17) (:O.19)' 6-(18-19)-1976 7.8 7.1 6.8 6.7 (30.25) (:O.16) (:0.00) (:0.06) X5 Oates 6.8 6.6 6.6 6.5 (:O.33) (30.16) (:0.16) (20.12) 8. TOTAL ORGANIC CARBON 6-(5-6)-1976 7.7 6.5 6.6 7.0 (:0.52) (:O.26) (:O.l3) (:O.42) 6-(11-12)-1976 6.8 7.2 7.6 7.6 (:O.20) (:O.29) (:O.36) {:O.64) 6-(18-13)-1976 8.0 7.9 7.4 7.2 (:0.23) (:0.17) (: 34) (:0.38) I3 Oates 7.5 7.2 7.2 7.3 (:O.3é) ,:O.24) (:0.18) (:0.10) Table 84 (cont'd.). 254 Sampling Lake Low High Oates Readings Control Density Density LAKE ONE--EXPERIMENT TWO A. OISSOLVEO ORGANIC CARBON 8-(7-8)-1976 9.2 7.8 8.3 8.1 (:0.67) (:0.21) (20.35) (:0.13) 8-(14-15)-1976 9.0 8.3 8.9 9.2 (:O.48) (:0.08) (:0.33) (:0.32) 3-(21-22)-1976 9.4 8.9 9.6 9.5 (:O.70) (:0.30) (:0.20) (:0.17) XS Dates 9.2 8.4 8.9 8.9 (:0.07) (:0.18) (:O.22) (:0.24) 9 TOTAL ‘PGFNIC CAFEC. 8-(7-8)-1976 9.5 3.7 9.7 10.2 (:0 78) (:0.15) (:O.30) (:0.37) 8-(14-1S)-1976 11.0 10.9 13.1 14.9 (mm) (21.01) (:1.03) (=o.43) 8-(21-22)-1976 1 .6 14.5 14.7 17.7 (21.48) (:1.12) (:1.54) (:1.02) 15 Dates 13.0 11.4 12.5 14.3 (:1.63) (:0.98) (:0.36) (31.26) LAKE FOUR--EXPERIMENT THO A. OISSOLVED ORGANIC CARBON 8-(7-8)-1976 8.3 8.0 7.8 7.7 (:0.25) (:0.17) (:0.15) (:0.79) 8-(14-15)-1976 9.2 8.6 8.9 9.0 (:0.10) (:O.40) (:0.47) (:O.74) 8-(21-22)-1976 11 6 10.1 10.1 9.4 (:1 48) (:O.90) (:0.55) (20.10) X3 Oates 9.7 8.9 9.0 8.7 (:0.57) (:0.36) (:0.38) (:0.28) 3. TOTAL ORGANIC CARBON 8-(7-8)-1976 9.1 8.2 8.2 8.1 (:0.47) ($0.08) (:0.15) (:0.86) 3-(14-1S)-1976 10.1 10.4 10.4 9.6 (20.53) (:0.32) (:1.49) (:0.52) 8-(21-22)-1976 11.8 11.3 10.6 10.3 (:1.49) (21.47) (20.52) {:0.10) X; Oates 10.3 10.1 9.7 9.3 J (:0.46) (93.50) (53.441 {:0.38) 255 Table 85. Concentrations of dissolved organic, tota1 organic, and total inorganic nitrogen (mg/l i SD) in imported lake water, unstocked (control) aquaria, and stocked (B, pipiens) aquaria during Experiments One and Two. SAMPLING LAKE LON HIGH DATES READINGS CONTROL DENSITY DENSITY I. LAKE 0NE--EXPERIMENT ONE A. Dissolved Organic Nitrogen 6 June 1976 0.11 1.57 1.11 1.31 (10.01) (10.02) (10.003) (10.13) 12 June 1976 1.21 1.47 1.65 1.78 (10.09) (10.13) (10.15) (10.07) 19 June 1976 0.99 1.04 0.67 1.09 (10.03) (10.14) (10.17) (10.04) 73 Dates 0.77 1.36 1.14 1.39 (10.33) (10.09) (10.15) (10.11) B. Total Organic Nitrogen 6 June 1976 7.44 7.51 7.16 7.44 (10.003) (10.22) (10.17) (10.06) 12 June 1976 8.74 8.29 8.44 8.47 (10.003) (10.12) (10.11) (10.03) 19 June 1976 8.58 8.31 8.43 8.43 (10.07) (10.07) (10.01) (10.01) 'Xj Dates 8.25 8.04 8.01 8.11 (10.40) (10.15) (10.66) (10.51) Table 35 (cont'd.). 256 SAMPLING LAKE LOW HIGH DATES READINGS CONTROL DENSITY DENSITY C. Total Inorganic Nitrogen 6 June 1976 1.57 1.89 1.73 1.75 (10.02) (10.16) (10.14) (10.14) 12 June 1976 2.14 2.09 2.06 1.92 (10.09) (10.11) (10.11) (10.03) 19 June 1976 1.69 1.70 1.60 1.40 (10.09) (10.01) (10.15) (10.17) 73 Dates 1.80 1.90 1.80 11.69 (10.17) (10.08) (10.09) (10.10) II. LAKE FOUR--EXPERIMENT ONE A. Dissolved Organic Nitrogen 6 June 1976 0.56 0.63 0.60 0.62 (10.02) (10.02) (10.01) (10.01) 12 June 1976 0.62 0.67 0.67 0.71 (10.20) (10.03) (10.03) (10.04) 19 June 1976 0.66 0.66 0.66 0.69 (10.02) (10.02) (10.01) (10.02) '73 Dates 0.61 0.65 0.64 0.67 (+0.02) (10.01) (+0.01) (10.02) 257 Table 85 (cont'd.). SAMPLING LAKE LON HIGH DATES READINGS CONTROL DENSITY DENSITY B. Total Organic Nitrogen 6 June 1976 12 June 1976 19 June 1976 Y3 Da tes 6 June 1976 12 June 1976 19 June 1976 73 Dates 69 01) 78 02) 79 02) 75 .02) 0. (10. 0. (10. 0. .02) O. .03) (:9 Total Inorganic Nitrogen .62 0. .01) (10. .68 0. .01) (10. .81 0. .06) (1o. .70 0. .05) (10 .03 0. .00) (10. .03 0. .00) (10. .03 O. .00) (1o. .03 O .00) (102 03 00) 01 01) O9 01) 05 01) 64 03) 80 06) 81 75 .‘03 .03) .02 .01) .11 .02) .05 .67 .01) .76 .05) .77 .04) .73 .02) .04 .04) .03 000) .07 .03) .05 .01) Table 85 (cont'd.). 258 SAMPLING LAKE LON HIGH DATES READINGS CONTROL DENSITY DENSITY III. LAKE 0NE--EXPERIMENT TWO A. Dissolved Organic Nitrogen 8 August 1976 1.00 1.35 1.39 1.43 (10.02) (10.05) (10.01) (10.04) 15 August 1976 1.07 1.20 1.07 1.12 (10.07) (10.10) (10.10) (10.01) 22 August 1976 0.90 1.45 1.45 1.12 (10.13) (10.20) (10.12) (10.09) 73 Dates 0.99 1.33 1.31 1.22 (10.04) (10.07) (10.07) (10.06) B. Total Organic Nitrogen 8 August 1976 1.44 1.96 2.08 2.23 (10.04) (10.08) (10.04) (10.04) 15 August 1976 1.94 1.78 1.71 2.37 (10.10) (10.03) (10.28) (10.40) 22 August 1976 3.34 2.22 3.10 3.29 (10.12) (10.05) (10.28) (10.12) 73 Dates 2.24 1.99 2.30 2.63 (10.56) (10.07) (+0.23) (10.20) Table 85 (cont'd.). 259 SAMPLING LAKE LON HIGH DATES READINGS CONTROL DENSITY DENSITY C. Total Inorganic Nitrogen 8 August 1976 3.40 2.00 2.11 2.21 (10.02) (10.02) (10.07) (10.07) 15 August 1976 3.37 2.80 2.75 . 2.73 (10.02) (10.06) (10.06) (10.06) 22 August 1976 1.18 1.54 1.14 1.11 (10.02) (10.01) (10.11) (10.12) 73 Dates 2.65 2.12 2.00 2.027 (10.73) (10.18) (10.23) (10.24) IV. LAKE FOUR--EXPERIMENT TWO A. Dissolved Organic Nitrogen 8 August 1976 0.72 0.80 0.81 0.80 (10.01) (10.02) (10.02) (10.01) 15 August 1976 0.82 0.82 0.76 0.80 (10.02) (10.02) (10.02) (_0.01) 22 August 1976 0.84 0.82 0.89 0.87 (10.02) (10.01) (10.01) (10.01) '73 Dates 0.79 0.81 0.82 0.83 (10.03) (10.01) (10.02) (_0.01) Table 85 (cont'd.). 260 SAMPLING LAKE LOW HIGH DATES READINGS CONTROL DENSITY DENSITY B. Total Organic Nitrogen 8 August 1976 0.91 0.86 0.86 0.84 (10.01) (10.01) (10.02) (10.01) 15 August 1976 0.86 0.87 0.83 0.84 (10.01) (10.01) (10.04) (10.00) 22 August 1976 0.94 0.85 0.91 0.89 (10.01) (10.00) (10.02) (10.01) 1'3 Dates 0.90 0.86 0.87 0.86 (10.02) (10.01) (10.01) (10.01) C. 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Phytop1anxtonic diversities :10 nets) 1n 1moorted 1ake water, unstocked (control) aouaria. and stocked (10w and h1 n 'tocks of 180 are fro : 5 c . at Lake One. 9 ’ p 9 3000185) aquar1a .or Exper1menc One HIERARCHICAL DIVERSITY Number EXPERIMENT of ONE Genera Evenness LAKE ONE Greens Diatoms 31ue-greens Cryptopnytes Phy1a Tota1 (N5) (0') A. CONTROL 1) Initia1 Readings 28 May 1976 a) Number Efiversity X3 0.552 0.784 0 0.190 1.312 3 0.693 Poo1ed 3.671 0.967 0.154 0.551 0.774 0.372 b) Vo1ume Diversity 3 0.355 0.641 0 0.631 1.420 3 0.750 Poo1ed 0.306 0.350 0.592 0.527 0.323 0.396 2) 6 June 1976 a) Number D1versity 3 1.202 0.124 0 0.344 1.148 2n 0.433 Poo1ed 1.115 0.154 0.141 0.641 0.776 ' 0.256 b) Vo1ume Qfiversity x3 0.963 0.189 0 0.454 3.309 29 0.305 Poo1ed 0.923 0.313 0 577 0 512 0.543 0.215 3) 12 June 1976 a) Number 01' versity X3 1.279 0.536 0.777 0.396 1.934 0.668 Poo1ed 1.472 0.763 0.820 0.693 0.752 1.251 25 0.389 b) Vo1ume gyversity x3 1.137 0.827 0.551 0.399 1.764 2; 0.608 Poo1ed 1.420 1.011 0.716 0.10” 0.733 1.126 ” 0.349 4) 19 June 1976 a) Number Diversity 3 1.087 1.000 0.616 0 1.950 23 0.682 Poo1ed 1.183 0.843 0.424 0.090 0.870 1.310 0.418 b) Vo1ume Diversity 3 1.027 1.165 0.656 0 1.885 :3 0.559 Poo1ed 1.241 1.015 0.518 0.687 0.763 1.279 0.408 5. LOH DENSITY 1) Zn1t181 Readings 27 May 1976 a) Number giversity ‘3 9.956 1.171 3 0.245 1.056 1.853 11 0.775 b) Vo1ume Diversity 73 0.623 1.390 0 0.553 0 838 1 398 11 0.791 Tab1e 814 (cont'd.). 288 Greens Diatoms 81ue-greens CryptOphytes Phy1a Tota1 (N5) (0') 2) 5 June 1976 a) Number giver-sit)! x3 1.320 0.285 0.026 0.281 1.667 25 0.595 Poo1ed 1.500 0.360 0.100 0.466 0.718 0.954 0.293 b) Vo1ume giversity X3 1.153 0.450 0.004 0.552 1.315 25 0.465 Poo1ed 1.355 0.537 0.017 0.316 0.698 0.926 0.284 3) 11 June 1976 a) Number Diversity ‘ 1.324 0.362 0.596 0. 07 1.870 22 0.583 Poo1ed 1.300 0.541 0.851 0.451 0.729 1.070 0.346 b) Vo1ume I‘Jersfty 73 1.142 0.453 0.136 0.214 1.41:5 22 0 512 Poo1ed 1.145 0.357 0.371 0.329 0.658 0.953 0.308 4) 18 June 1976 a) Number D1versfty 3 1 96 0.364 0.719 0.110 1.904 22 0.594 900180 1.370 0.409 1.249 0.071 0.871 1.208 0.391 b) Vo1ume giversity x3 0.991 0.380 0.209 0.151 1.345 22 0.491 Poo1ed 1.016 0.582 0.531 0.660 0.768 1.043 0.337 C. HIGH DENSITY 1) 101:151 Readings 28 May 1976 a) Number giversity K3 0.806 0.511 0 0.168 1.284 0 0.631 Poo1ed 1.036 0.670 0.180 0.693 0.871 ' 0.396 b) Vqume gjversity X3 0.410 0.562 0 0 -99 1.402 9 0.688 Poo1ed 0.597 0.315 0.’35 0.695 0.364 0.439 2) 6 June 1976 a) Number Diversity .3 1.216 0.260 0.022 0.240 1.641 9n 0.616 Poo1ed 1.275 0.269 0.074 0.257 0.694 0.924 '7 0.309 b) Vo1ume ijersity X3 0.904 0.347 0.001 0.559 1 259 20 0.470 900180 0.946 0.430 0.004 0.537 0.698 0.354 0.235 3) 12 June 1976 a) Number giversity 13 1.392 0.360 0.331 3.583 1 312 a: 0.531 71301.20 1.481 0.445 0.610 0.593 0.627 0.546 “ 3 263 16618 814 (cont‘d.). 289 Greens Diatoms Blue-greens Cryptophytes Phy1a Tota1 (N5) (0') b) Vo1ume Diversity 3 1.336 0.500 0.033 0.212 1.439 25 0.502 Poo1ed 1.497 0.639 0.074 0.107 0.585 0.761 0.236 4) 19 June 1976 a) Number gfiversity X3 1.080 0.367 0.918 0 1.831 24 0.666 Poo1ed 1.171 0.607 1.293 0.713 1.062 0.334 b) Vo1ume inersity X3 0.932 0.474 0.463 g 1.382 24 0.504 P001ed 1.164 0.335 0 538 3 572 0.959 0.382 0. LAKE READINGS 1) In1t151 Pead1n05 23 May 1976 a) Number Diversity ’3 0.741 0.699 0 0.191 1.374 , 0.725 Poo1ed 3.973 0.924 0.203 0.538 0.879 ' 0 423 0) Vo10me gfiversity X3 3.436 0.617 0 0.618 1.440 8 0.76-1 Poo1ed 0.559 3.605 0.606 0.638 0.846 0. 07 2) 6 June 1976 a) Number giversity X3 3.577 0.463 0 0.194 1.276 3] 0 516 Poo1ed 0.699 0.889 0.142 0.738 0.916 0 382 b) Vo1ume inersity 3 0.709 0.316 0 0.506 1 584 11 0.764 Poo1ed 0.759 0.588 0.675 0.353 1.186 0.494 3) 12 June 1976 a) Number Diversity . 0.589 0.449 0.517 0.096 1.229 14 0.490 Poo1ed 0.531 0.792 0.231 0.042 0.525 3.750 3.284 b) Vo1ume 01versity 3 0.693 0.369 0.296 0.637 1.452 14 0.579 Poo1ed 0.636 0.678 0.636 0 551 0 556 0.723 0.274 4) 19 June 1976 a) Number giversity K3 0.150 0.502 0.437 0.250 0.506 10 3 22 Poo1ed 0.231 1.745 0.927 0.115 0.516 0.‘02 0.217 b) Vo1ume Diversity 'T3 0.236 3 8 0 ..3 0.251 3.308 1n 0.351 Pooled 0.293 5‘7 0. 3.592 0 557 0. 91 '” 0.219 290 I.‘ 3.: 3.3 .2: a; 2...... .33 $2.... :23 a. 2. 8. on ..N :2 No: 2.2. 22 .25.. 2-... ... .....d :23 .....fl 3.... :3 $3.... :33 2.3.3 mm 88 2.. 2 3. mm... 22 ...... 2.... as... 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SW. £2 95., 2.: Am 5.: $3 $3 8%.; 82m. 8mm: :33 : cm em = 2.: :2 E: a: £2 2% c-m 3 A83 333 .2... 83.: 8.; ES 2. m2 2.; 3. .32 BE 33. .33 £2 3. 3:.ka 35.83;: .m :3 A83 :3 :ad :3 :3 :3 Ed m; 2: E a: S a a. z 22 2.2, 2-2 ... a: :3 83 35 A85 :3 2...: 5.: c: 3 z: m .3 mm. 9.: : £2 25.. 2-: Am 83 2 3 :3 SN: $3 $3 3:: 3%; 2. a 2 _m 2 a: :2 .3 £2 22. 9m 3 ac: _ new; 3 . «can 3 .3.ch 35:8 3: :63. 5 3:3 3 _ mac: 35:8 8.2 :3: :3 9.2 :3: :5 . A 6.253 mg mg: 294 Tab1e 816. PhytopIanktonic diversities (in nets) in 1moorteo 1ake water. unszocked (control) aauar1a. and Stocked (10w and nign stocxs of 1eopard frog tadpoles) aquaria for Experiment One at Lake Four. HIERARCHICAL DIVERSITY Number EXPERIMENT of ONE Genera Evenness LAKE FOUR Greens Diatoms 81ue-greens Cryptopnytes Phy1a Tota1 (Ms) (J') A. CONTROL 1) Init1a1 Readings 28 Hey 1976 a) Number giversity K3 1.245 0.275 0 0.150 1.615 13 0.765 Pooled 1.555 0.485 0.451 0.475 1.383 3.422 5) Volume giversity x3 1.154 0.186 0 0..1. 1.345 73 0.683 Pco1ed 1.325 0.311 0. 3 0.776 1.219 0 525 2) 5 June 1976 a) number ‘giversity ‘3 1.005 0.188 0.311 0 1.749 15 0.730 Pooled 0.992 0.473 1.046 0.779 1.150 0.424 b) Vo1ume g1versity §; 1.285 0.244 0.551 g 1.827 15 0.765 001ed 1.320 0.610 0.572 0.996 1.252 0.462 3) 12 June 1976 a) Number giversity X3 1.158 0.157 0.653 0 1.621 14 0.119 Poo1ed 1.302 0.301 1.003 0.568 1.213 0.460 b) Vo1ume Diversity 73 1.097 0.068 0.897 g 1.685 14 0.736 Poo1ed 1.273 0.134 1.198 0.830 1.310 0.496 4) 19 June 1976 a) Number giversity X3 1.048 0 0.743 0 1.212 14 0.578 Pooled 1.389 1.017 0.514 0.901 0.341 b) Vo1ume Diversity Y} 0.884 0 0.583 0 1.294 14 0.538 PooIed 0.978 1.000 0.590 1.064 0.403 3. LOW DENSITY 1) Initial Readings 28 May 1976 a) Number giversity X3 1.386 0.392 D 0 311 1 550 1; 0 736 Poo1ed 1.229 0.559 0 537 0.561 1 121 3.167 b) Volume giversity 13 1.025 0.518 3 3 18 1.554 v 3.743 Poo1ed 0.912 0.356 3 79 3.913 1.301 3.543 Tab1e 816 (cont'd ). 295 Greens Diatoms 31ue-greens Cryptcphytes Phy14 7013‘ ("0) (4') 2) 6 June 1976 a) Number _inersity x3 1.458 0.038 0.772 0.198 2.018 15 0.838 Poo1ed 1.524 0.113 1.017 0.595 0.928 1.173 0.433 b) Vo1ume Diversity 3 1.414 0.066 0.206 0.071 1.794 15 0.742 Poo1ed 1.415 0.194 0.426 0.212 0.753 1.185 0.438 3) 12 June 1976 a) Number Diversity 3 1.257 0.132 0.735 0 1.720 17 0.570 Poo1ed 1.515 0.036 0.247 0.753 0.875 0.309 0‘. ‘Io'tume Zivers :1 ‘T3 1.244 0 310 9.549 3 0.525 17 0.242 PooIed 1 336 0. 3 0.856 0.506 0.559 0.197 4) 190une 1976 a) Number Diversity 73 1.145 0.012 0 803 0.079 1.338 13 0.545 Poo1ed 1.730 0.072 0.711 0.252 0.553 0.356 0.297 b) Volume giversity X3 1.045 0.015 0 511 0.188 1.199 13 0.495 Poo1ed 1.679 0.091 0.401 0.532 0.702 0.558 0.297 C. HIGH DENSITY 1) Initia1 Readings 28 May 1976 a) Number _D_iversity 3 1.256 0.480 a 0.224 1.713 12 0.816 PooIed 1.346 0.799 0.500 0.715 1 227 0.494 b) Vo1ume gfiversity 11;; 1.140 0 395 0 0.048 1.473 12 0.714 Poo1ed 1.085 0 869 0.308 0.772 1.314 0.529 2) 6 June 1976 a) Number giversity X3 1.375 0.123 0.591 0.213 1.944 18 0.756 PooIed 1.396 0.222 0.490 0.434 0.756 1.021 0.353 D) ‘O’OIUme Diversity .3 1.442 0.363 0.094 0 204 1.769 18 0.686 PooIed 1.707 0.768 0.046 0 568 0.831 1.152 0 397 3) 12 June 1976 a) Number giversity X3 1.248 0.014 0.710 g 1.294 is .540 Doo1ed 1.312 0.031 0.975 0.715 0 558 0.297 Table 816 (cont‘d.). 21965 Greens Diatoms Blue-greens Cryptopnytes Phyla Total (Ne) (J') b) Volume Diversity 3 1.219 0.025 0.585 0 0.489 13 0.202 Pooled 1.345 0.065 1.011 0.470 0.535 0.185 4) June 1976 a) Number Diversity : 1.281 0 0 574 0 1.379 15 0.554 Pooled 1.770 0 799 0.665 0.837 0.302 b} Vo1ume giversity 3. l 329 0 0.513 g 0.394 15 9.359 P601ed 1 547 0.537 0.592 0.552 0.235 0. LAKE READINGS 1) 27 May 1976 a) Number giversity x3 1.448 0.199 0 0 1 010 1.890 9 0.860 bl Volume Diversity ‘7: 1.192 0.321 0 0 0.698 1.300 9 0.592 2) 5 June 1976 a) Number Diversity ‘T3 1.209 0 2 0 O 0.622 1.670 17 0.714 Pooled 1.331 0 6‘3 0.637 0.655 0.615 1.211 0.427 b) Vo1ume Diversity . 1.169 0.356 O 0.075 1.338 17 0.578 Pgoled 1.652 0.907 0.620 0.069 0.823 1.106 0.390 3) 11 June 1976 a) Number ‘inersity x3 0.928 0 124 0.476 0 1.710 13 0.331 Poo1ed 1.326 0.177 0.787 0.839 1.225 0.478 5) Volume giversity 3 0.577 0.177 0.515 g 7.507 13 0.719 Pooled 1.034 0.290 1.040 0.853 1.130 0.441 4) 18 June 1976 a) Number Diversity ‘73 0.882 0.028 1.069 0.210 1.885 16 0.752 Pooled 0.348 0.071 1.350 0.278 0.780 1.222 0.441 b) Vo1ume inersity X3 9.594 0.043 0.386 0.312 1.686 15 0.67. 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Phytoolanktonic diversities {in nats) 1n imported lake water. unstocked (control) aquaria. and stocxed (low and high stocks of leopard frog tadpoles) aquaria for Experiment Two at Lake One. HIERARCHICAL DIVERSITY Number EXPERIMENT of Genera Evenness LAKE ONE Greens Diatoms Blue-greens Cryotopnytes Phyla Total (N5) (0') A. CONTROL 1) Initial Readings 26 July 1976 a) Number giverslty x3 1.027 0.725 0.592 1.450 15 0.615 Pooled 1.309 0.352 0.992 0 0.540 0.981 0.362 5) Volume giversity x3 1.208 0.572 0.557 A 1.672 15 0.710 Pooled 1 7: 0.720 0.’16 ‘ 0.685 1.266 0.d68 2‘ 8 iugust 7976 a) Number giversity 23 1.190 0.792 0.465 0 1.881 72 0.664 Pooled 1.248 0.860 0.465 0.814 1.164 ‘ 0.377 0) Volume giversity A3 1.632 0.751 0.580 O 1.047 22 0.420 Pooled 1.854 0.768 1.029 0 687 0.854 0.276 3) 15 August 1976 a) Number giversity x3 1.631 0.770 0.104 0 1.020 19 0.393 Pooled 2.350 0. 7 0.074 0.592 0.708 0.241 bl Volume 21' versity x3 1.328 0.541 0.086 0 1.034 19 0.400 Pooled 1.737 0.682 0.065 0.498 0.672 0.228 4) 22 August 1976 a) Number Diversity ‘3 1.297 0.792 0.622 0 1.656 25 0.509 Pooled 1.216 1.003 0.593 0.433 0.577 0.179 b) Volume ijerslty X3 1.048 0.848 0.508 0 1.682 25 0.627 Pooled 0.889 1.089 0.606 0.583 0.729 0.227 8. Low DEVSITY l) lnitlal Readings 26 July 1976 a) Number ijerslty 13 0.324 0.673 0.473 1 1.179 ~5 0.605 Pooled 0.387 0.756 0.310 ‘ 0 514 0.742 3.257 :1 Volume _3_1' ver: 1 :y Va 1.040 0.605 0.7‘6 q 1 508 : 1.556 Ddolec 1.142 0.799 0.502 ‘ 2.525 ‘ 3:3 ‘ 3.349 316 Table 821 (cont'd.). Greens Diatoms Blue-greens Cryptophytes Phyla Total (No) (J') 2) 8 August 1976 a) Number Diversity 3 1.130 0.791 0.365 0.022 1.879 22 0.668 Pooled 1.396 0.776 0.397 0.051 0.814 1.151 0.372 0) Volume inersity X3 1.545 0.757 0.651 0.001 1.078 22 0.385 Pooled 1.803 0.774 0.984 0.003 0.644 0.748 0.242 3) 15 August 1976 a) Number giversity X3 1.857 0 7 0.199 g 1 .73 23 0.468 Pooled 2.086 0 7'9 0.090 0.689 3 55 0.273 b) Volume 0ivers‘ty T3 1.745 0.514 0.339 .3 0.900 23 0.308 Pooled 1.973 0.544 0.432 0.490 0.665 0.212 4) 22 August 1976 a) Number Diversity 3 1.03 0.807 0.346 0 1.460 28 0.522 Pooled 1.047 0.803 0.285 0.723 0.983 0.295 b) Volume giversity x3 0 764 0.739 0 314 0 1.800 28 3.574 Pooled 0.799 0.900 0 274 0 387 1.187 0. 56 C. HIGH DENSITY 1) initial Readings 26 July 1976 a) Number Diversity 3 0.900 0.682 0.693 0 1.365 15 0.566 Pooled 1 093 0.797 1.287 0.505 0.791 0.282 0) Volume Diversity 73 c 932 0 :12 0.447 ,3 1.493 15 0.620 Pooled 1 147 0.594 0.904 0.619 0.890 0.321 2) 8 August 1976 a) Number Diversity 3 1.051 0 743 0.839 0.010 1.713 22 0.598 Pooled 1.117 0 773 0.870 0.032 0.635 0.355 0.277 5) Volume Diversity 1‘; 1.311 0.741 0.198 0.0003 1.238 .2 0.432 Pooled 1.333 0.724 0 163 3.00 0.651 0.812 7 0.253 3) 15 August 1976 a) Number giversity (3 1.312 0.725 _ 0.159 3 1.411 21 0.187 Pooled 1.867 0.744 0 195 0 586 0.759 0.249 Table 321 (cont‘d.). 317 Greens Diatoms Blue-greens Cryptoonytes Phyla Total (N5) (0') 0) Volume inersity X3 1.655 0.510 0.461 0 0.799 21 0.277 Pooled 1.749 0.544 0.519 0.431 0.609 0.200 4) 22 August 1976 a) Number giversity X3 1.084 0.791 0.322 0 1.492 30 0.476 Pooled 1.209 1.056 0.354 0.528 0.697 0.205 0) Volume giversity X3 1.169 0.875 0 295 g 1 954 30 0.624 Pooled 1.047 1.197 0.345 0.699 O 7 0.238 0. LAKE READINGS 1) Initial Readings 25 July 1975 a) Number giversity X3 0.821 0 733 0.838 0 1.532 14 0.652 Pooled 0.779 1.040 0.876 0.686 1.012 0.383 b) Volume inersity X3 0.651 0.830 0.83 0 1.552 14 0.557 Pooled 0.638 1.220 1.202 0.700 1.018 0.386 2) 7 August 1976 a) Number _iversity x3 1.159 0.795 0.459 0.015 1.903 27 0.677 Pooled 1.467 0.869 0.773 0.042 0.880 1.213 0.368 b) Volume inersity . X3 1.185 0.803 0.185 0.001 1.092 27 0.389 Pooled 1.423 0.907 0.794 0.002 0.631 0.738 0.224 3) 14 August 1976 a) Number giversity x 1.447 0.776 0.053 0 1.512 21 0.553 Pooled 1 529 0.802 0.056 0.694 0.890 0.292 0) Volume Diversity 3 1.607 0.542 0.036 0 0.863 21 0.314 Pooled 1.673 0.550 0.084 0.468 0.583 0.224 4) 21 August 1976 a) Number _0_iversity X 0.997 0.572 0.262 0 1.392 29 0.459 Pooled 0.928 0.734 0.364 0.637 0.845 0.255 b) Volume Efiversity (3 0.681 0 855 0.101 0 1.70 33 3.550 Pooled 0.681 1 058 0.193 0 696 0.363 0.259 318 Table 822. Phytbolanktonic diversities (in nats) in imported 1aRe water, unstocked (control) aquaria, and stacked (low and nigh stocks of leooard Frog tadpoles) aquaria for Experiment Two at Lake Four. HIERARCHICAL DIVERSITY Number EXPERIMENT of TWO Genera Evenness LAKE FOUR Greens Diatoms Blue-greens cryptophytes Phyla TOCBI (”5) (J') A. CONTROL 1) Initial Readings 26 July 1976 a) Number 01 versity x3 0.813 O 0.393 0.096 0.725 12 0.326 Pooled 0.903 0.814 0.385 0.476 0.745 0.300 01 Volume Di vers 1' ty x3 0.576 0 0.070 0.006 0.516 12 0.322 Pooled 0.732 0.171 0.029 0.471 0.509 0.205 2) 3 August 1976 a) Number ‘giversity .3 0.893 0.442 0.558 3 1.350 15 0.397 Pooled 1.254 0.693 0.709 0.581 0.898 0.332 b) Volume 0i vers i ty ‘3 0.948 0.334 0.093 0 0.823 15 0.632 Pooled 1.150 0.578 0.162 0.632 0.763 0.282 3) 15 August 1976 a) Number Diversity X3 0.322 0.348 0.623 0 1.119 14 0.493 Pooled 0.384 0.948 0 703 0.447 0.537 0.204 6) Volume 21 vers 1 ty 3 0.333 0.277 0.085 0 1.149 14 0.570 Pooled 0.371 0.898 0.070 0.476 0.511 0.193 4) 22 August 1976 a) Number Diversity Y3 1.290 1.155 1.275 0.329 1.852 22 0.854 Pooled 1.566 1.846 1.235 0.257 0.538 3.933 0.302 0) Volume _D_i vers i ty X3 1.162 0.966 1.567 0.520 2.431 22 0.649 Pooled 1.405 1.730 1.521 0.600 0.672 1.041 0.337 8. LON DENSITY 1) Initial Readings 26 July 1976 a) Number _0_iversity X3 0.353 0 0.42 a 0.510 9 0.194 Pooled 0.534 0.361 ' 0.116 0.339 0.382 6) Volume giversity 13 3.286 0 0.106 3 0.333 3 0.307 Pooled 0.425 0.250 3.453 0.533 0.242 319 Table 322 (cont'd.). Greens Diatoms Blue-greens Cryptoohytes Phyla Total (N5) (0') 2) 8 August 1976 a) Number giversity '3 0.693 o 0.705 0 1.284 10 0.314 Pooled 0.845 0.922 0.471 0.650 0.282 b) Volume giversity X3 0.576 0 0.110 0 0.647 10 0.533 Pooled 0.373 3.208 0.448 0.504 0.219 3) 15 August 1976 a) Number Diversity ,3 0.515 3.333 0.4 O 3 2.2 1.531 23 0.585 Pooled 1.339 0.521 0. 7 3 637 0 588 0.929 3.296 b) Volume giversity x3 0.376 0.215 0.158 0.026 1.357 23 0.507 Pooled 1.523 0.418 0.278 0.079 0.713 0.917 0.293 41 22 August 1976 a) Number giversity ‘3 1.128 0.991 0.895 0.324 1.326 30 0.748 Pooled 1.712 1.759 1.048 0.117 0.668 1.072 0.315 b) Volume gyversity X3 1.267 0.563 1.577 0 511 2.251 10 0.607 Pooled 1.740 1.175 1 734 0.689 0.362 1.370 ' 0.403 C. HIGH DENSITY 1) Initial Readings 26 July 1976 a) Number giversity x3 0.613 g 0.665 O 0.958 14 0.326 Pooled 1.389 0.662 0.429 0.596 0.226 0) Volume giversity x3 0.599 3 0 232 0 0.718 14 0. 3 Pooled 1 429 0 0.537 3.57 0.217 2) 3 August 1976 a) Number Diversity 3 0.961 0.174 0.954 0 1.624 14 0.459 Pooled 1.305 0.333 1.102 0.593 0.554 0.568 0.329 0) Volume giversity X3 0.964 0.108 0.232 0 1 ‘8 12 3 105 Pooled 0.896 0.168 0.261 0.075 0 714 "36 0 ‘17 3) 15 August 1976 a) Number giversity ‘1 3.465 0.257 0 581 3 1 470 4 0.585 Pdoled 0 693 0.572 0.566 3.767 1 ‘4 3.397 320 Table 822 (cont‘d.). Greens Diatoms Blue-greens cryptophytes phyla Total (N5) (J') b) Volume Diversity 3 0.795 0.232 0.155 0 1.534 14 0.653 Pooled 1.037 0.562 0.334 0.857 1.068 0.405 4) 22 August 1976 a) Number Diversity 7} 1.388 0.552 1.140 0.315 1.992 24 0.784 Pooled 1.587 1.241 0.981 0.206 0.682 1.056 0.332 5) Volume §fiversity 13 1.197 0.515 1.643 0.543 2.255 24 0.595 Pooled 1.742 1.241 1.621 0.653 0.879 1.332 0.419 0. LAKE READINGS 1) Initial Readings 25 July 1976 a} Number 01 vers i ty 4, 0.300 0.187 0 . 0 0.328 12 0.154 Pooled 1.047 0.575 0 287 ‘ 0.456 3.576 0.232 5) Volume ‘giversity .3 0.710 0.114 0.145 0 0.331 12 0.151 Pooled 0.914 0.496 0.036 0.472 0.438 0.196 2) 7 August 1976 a) Number Diversity :3 0.971 0.297 0.789 3 1.658 15 0.702 Pooled 1.183 1.289 0.881 0.555 1.019 0.368 0) Volume giversity X3 0.939 0.221 0.395 g 1.441 75 0.705 Pooled 1.158 1.273 3.219 0.886 1.095 0.395 3) 14 August 1975 a) Number 0i versi ty A3 3. 23 0.905 0 794 g 1.409 22 3 ~03 Pooled 0.624 1.421 0 92 0 655 0.826 0 67 b) Volume _0_i versi ty X3 0.486 0.704 0.269 0 1.063 23 0.532 Pooled 0.539 1.190 0.360 0.886 0.687 0.222 d) 21 August 1976 a) Number giversity x3 1.707 1.313 1.604 0 020 ..351 23 0.404 Pooled 1.958 1.777 1.523 3 603 0.770 1.156 0.34~ 3) Volume Diversity I 1.15 1.375 1.133 0 225 .232 99 3.769 Pooled 1.280 1.562 1 274 3 245 0 736 3 96“ " 3.288 Table 323. 321 Numbers and volumes of phytoplankton (per m1 : $0) in imported lake water and aouaria with monocultured bullfrogs, a mixed-scecies culture. or monocultured green frogs in Exceriment Three at Laxe One. EXPERIMENT THREE LAKE ONE Greens Diatoms Blue-greens Cryptophytes Totals A. BULLFROGS 1) Initial Readings 22 September 1976 '7 Number 32606 617 417341 285 450850 (:8964.7) ‘(;;4.31 (:12313.1) (:87.1) (:15422.7) '7 v61ume 2957352.3 140246 5875316 21118.3 9004042.7 (:189512.71 (:22424.9) (:2121011.3) (:5451.9) (:2178492.5) 7 Size 91.3 227.3 14.1 74.1 29.0 2) 5 October 1976 x Number 152419 1900 273235 342 428896 (_18182.1} (:452 3: (:98979.2) (:57) (524413 9) ‘7 Volume io7ooooo.3 546158.7 4424416.? 25342.7 1569.. 8 4 (11555591.31 (:127950.8) (:9555405.21 (:4223.7) ’;49 1.9) ‘7 Size 59.7 287.5 16.2 74.1 35.5 3) 12 October 1975 x Number 159355 7070 259644 1615 427685 (:5256.9) (:853.7) {113543.1) (:?14.4) (:15476.3) 7 Volume 25663000 2581742.? 4328810.7 138024 32711577.4 (:845780.2) (:292698.4) (:252761.5) (:9880) (1552384.9) ‘7 Size 151.0 355.2 15.7 35.5 75.5 4) 24 October 1976 x Number 103745 2522 286841 2952 395251 (:1878.8) (3457.4) (:54824.9) (3230.4) (:52892.4) '7 Volume 24432333 598518.2 2828048.7 152cs3.3 28110953.2 (19945309) (11287059) (11031661 .31 (:17069.8) (134655751 ‘7 Size 235.5 256.4 9.9 74.1 71.1 Mean 01" 3 Dates- 7’Num5er 138840 3864 273240 .1336 417251 (324303.21 (3855.6) (333330.91 13272.5) (:18250.9) '7 Volume 20265111.1 1275473.2 3860425.4 105140 25506149.7 (:2768817.3) (:842072.9) (:50 72 .8) (+20880 11 (:2933937.7) 7 Size 145.0 330.1 14.1 78.7 51.1 5. MIXED: BULLFQOGS AND GREEN 74065 1) Initial Readings 22 Seotemoer 1976 7 Number 32351 5 433487 342 466854 (:1684.4) (:107.5) (:7553.51 (355.81 f;9125.5) '7 Volume 3314108 174179 3115125 25342.3 11523755.: {1246207.3) (:55304) (1114557.91 _4“77.21 {*270635.11 7'size 102.4 254.6 ‘5.’ ’4.‘ 24.9 2) 5 October 1976 1 Number 32551 3040 494624 :54 555539 I;=2259.21 1-219.1 (:50257.1: {:7‘ :1 (455479.7‘ Table 823 (cont‘d.). 322 Greens Diatoms Blue-greens CryptOpnytes Totals 7 Volume 10455418 743654.3 8291817.7 25997.3 l9517887.3 (:1704324.9) (:23893.9) (jfl193209.5) (35250.6) (:2882996.2) X'Size 118.1 244.6 16.8 74.2 33.3 3) 12 October 1976 Number 174038 6975 334297 1539 516849 ' (114391) (1785. 5) (127843.71 (:55 .8) (3683.6) 'Y Volume 25569666 2497942 5513913 114040 33695561 (3990466.7l (:284175.8) (:578087 31 {:4875.9) (:1047712.5) Y Size 146.9 358.1 16.5 74.1 65.2 4) 24 October 1976 x Number 100816 2869 315387 2623 421695 (:3097.6) (:18.3) {:41653.8) (:352.6) (_41405.21 ‘T Volume 21335009 330‘10 3 3393669 794339.? 23773719 (11530646.?) ’+25504 9) (:762385.9) (125129.91 (3946104.5) Y Size 241.6 289.’ 10.8 74.1 68.2 Mean of 3 Dates- ?'Number 121138 4295 381436 1.09 508373 (:14315.l) (:710.6) (135084) (:843) (:32283.8) 7 Volume 20125694.? 1357435.8 5733133.2 1.1792.3 27329055.8 (:2520103.3) (:297132.4) (:836234.6) (125418.21 (:2275210.2) 7 Size 166.1 316.0 15.0 74.1 53.3 C. GREEN FROGS 1) Initial Readings 22 September 1976 X Number 33537 494 412557 323 446912 (12156) (1933) (:5907.31 (:36) (:7843.61 7 Volume 3082443 115113 7775900. 3 23934. 3 10998391 (1335453) (:21481.8) (:85789.1) (35631.7) (:391077.3) Y'Size 91.9 254.6 18.3 74.1 24.6 2).; October 1976 X Number 102316 3753 442892 494 549455 (112073.21 (333.7) (1661237) (:1901 (168150.31 X Volume 11746213 987585.7 751765.7 36605 20342169.4 (:1657183.9) (:56990.5) (:1445961.1) (_13934.6) (:2882996.2) < Size 114.8 263.1 17.1 74.1 37.0 3) 12 October 1976 7 Number 173983 7298 275384 1657 458322 (:4535.61 (_717.4) (211349.43 (1204.4) {:3 5‘,51 x Jolume 25749000 2540554 4231996 .22309.3 32744353.3 {:j92610.2) I;93528.1, (1191668.8) (115142.31 (1741694.7) '7 Size 148.3 361.3 15.4 ‘4 1 71.4 Tab1e 823 (cont'd.). 323 Greens Diatoms 81ue-greens Cryptoohytes Tota1s 4) 24 October 1976 Y'Number 98562 2812 223004 2223 325601 (:5794.8) (:193.2) (35227.1) (:228.3) (:2712.5) Y'Vo1ume 23114666 768789.7 1598511.7 164749.3 25646716.7 (:935769.3) (:86274.4) (:16462.9) (:06745.8) (:1294052.4) 7 Size 234.5 273.4 7.2 74.1 78.5 Mean of 3 Dates- Y'Numoer 124954 4621 313750 1458 444793 (:12930.3) (1788.5) (:38427.3) (:275) (139271.71 Y'Vo1ume 20203293 1465643.1 4467424.5 108054.2 25244414.8 (:2221259.3) (:297890.8) (:961306.5) (321777.91 (:2052022.8) ‘7 Size 161.7 317.2 14.2 74.1 36.3 0. LAKE READINGS 1) Initia] Readings 32 September 1976 x Number 4771 2468 423251 380 473809 (30196.9) (:285.0) (:89966.3) 1:81.91 (313472.6) T Vo1ume 393944S.7 495902 9573054.7 28158 14036560.4 (:82445.8) (:19137.2) (:221894.1) (16073 3) (:121606.5) '? Size 82.5 200.9 22.6 74.1 29.6 2).; October 1976 7 Number 84077 2831 313336 513 400757 (:35908.5) (:114.7) (:21376.9) (_117.4) (:j1925.6) Y Vo1ume 8890906 727341.3 4798083.3 38013.3 14454343 (:1155834.1) (367264.21 (1466872.?) (1870 1) (:764018.9) '7 Size 105.7 256.9 15.3 74.1 36.1 3) 10 October 1976 x Number 149263 6347 281748 1852 439220 (16380.5) (1104.5) (330431.” (1199.3) (+28556.5) ‘7 Vo1ume 20105000 2417436.3 4737256 137999 27397691.3 (:661634.9) (:87217.7) (:554308.5) (:14765.7) (:983753.7) ‘7 Size 134.7 380.9 16.8 75.5 62.4 4) 22 October 1976 7 Number 98891 7244 238468 29 7 347530 (14946.5) (1512.1) (:19450.9) (2198.1) (:22212) 7'Yo1ume 20763333 -14551,.3 2197810 198563.? 25305224.3 (:295603.3) (:254535 1) (:501766.41 {127356.21 (1642000.1) 7 Size 210.0 296 2 9.2 57.3 72.3 Mean of 3 Dates- 7 Number 110744 5474 277951 1757 395836 ’111390.4) (:j77.5) ’;32528.5) f;§03.3) 1:}:95.2? ‘7 Vo1ume 16515968.7 1763431.6 “011049.9 121856.7 22335752.? (32224209.5) (1302507.5) ~49467‘ 5? (127525.9‘ --301721.5) 7 Size 149.1 322.1 11': ‘93 55.5 I32¥1 Tab1e 324. Numbers and voiumes of ohytop1ankton (per m1 : SO) in imported 1ake water and aquaria with monocu1tured bu11frogs, a mixed-species cu1ture. 0r monocultured green rrogs 1n Exoer1ment Three at Lake Four. EXPERIMENT THREE LAKE FOUR Greens Diatoms 81ue-greens Cryptoonytes Tota1s A. BULLFROGS 1) 1nit1a1 Readings 22 September 1976 x Number 462 89 3052 133 3736 (1126.2) (151.8) (1377) (_+_29) (_+_471.2) Y Vo1ume 26797 21747 64002.3 9855.3 122401.6 (:7089.3) (:12714.9) (:9309-41 (32150.5) (:17940.7) i’Size 58.0 244.3 21.0 74.1 32.8 2)): October 1976 7 Number 1330 12527 11862 152 25871 (:307.7) (:3931.9) (:5086.4) (150.3) (:fi208.3) 7 ‘Iqume 230606.7 3116517 359387.? 11263.3 3717774.7 (:81046.3) (:454449.2) (:14159.3) (:3724.9) (:500404.6) 7 Size 173.4 248.8 30.3 74.1 143.7 3) 12 October 1976 1 Number 722 10143 1380 63 12308 (137.7) (1701.3) (11214.5) (16.3) (11436.5 '7 Vo1ume 108518 2500411.7 42140.7 4693 2655763.7 (:23338.8) (:159134.11 (:24892.4) (:469.3) (:21soos.7) ‘7 Size 150.3 246.5 30.5 74.5 215.8 4) 24 October 1976 X Number 139 3496 823 6 4465 (127.6) (1142.6) (1130.9) (33.3) (1237.6) 7 Vo1ume 15001.7 887072 31452.3 469.3 934995.3 (:6336.9) (:57957.9) (27041.7) (3969.3) (:65021.63 Y Size 115.1 253.7 38.2 78.2 209.4 Mean Of 3 Dates- 7 Number 630 8722 4688 74 14214 (:193.9) (11475.2) (12014.7) (125.3) (:3494.5) ‘T Vo1ume 118375.5 2168000.2 144326.? 5475.3 2436177.9 (139518) (13510135) (:54457.1) (11912.7) (14352192) 7 Size 162.2 248.6 30.8 74.0 171.4 8. MIXED: BULLFROGS AND GREEN FROGS 1) In1t1a1 Readings 22 Seotember 1976 X Number 918 70 2705 133 3825 (3299.8) (122.9) (+287.7) ’147.8) (1552.5) 1 Vo1ume 40817 17886.7 54810.7 10794 1-3508.4 (34087) (35600.8) (:5005) ’+4009.7) 119679 71 7 Size 44.5 744.1 20.3 31.2 32.3 2).; October 1976 x Number -68 11069 5865 152 17954 f 12) (3574.1) ’ 2457.51 .:68.51 (3116.31 325 Tab1e 324 (cont’d.). Greens Diatoms 81ue-greens Cryptopnytes Tota1s x Vo1ume 79777.7 3185959.7 210551.7 55172.3 3542571.4 (35940.9) (3333852.1) (352743.4) (355543.2) (3357099.6) x Size 91.9 287.8 35.9 435.3 197.3 3) 12 October 1976 7'Number 402 9787 351 38 10588 (341.5) (31505.3) (3159) (321.9) (31535.4) ‘7 Vo1ume 47509.7 2444105.3 24587.3 2815 2519119.3 (315093.3) (3415593.9) (312715.5) (31525.5) (3422253.4) 7 Size 118.2 249.7 58.4 74.1 237.9 4) 24 October 1976 7 Number 351 3439 374 0 4574 (324.5) (345‘ 41 (3437.1) (3958.9) '7 Veibme 25575.7 ., 48.3 25354 9 925991 ($0455.?) (:1- 913) (19846.31 (111303911) 7 Size 71.1 254.4 29.0 9 198.1 Mean of 3 Dates- 7’uumber 543 8098 2357 53 11071 :+97.7) (31284.8) (31135.5) (330.91 (32184.41 ‘7 Vo1ume 50989 2168338.1 86904 22995.1 2329227.2 (39311.1) (3375519.0) (334794.1) (320052.61 (3414477.21 ‘7 Size 93.9 257.3 35.7 355.0 210.4 c GREEN FROGS 1) Initia1 Readings 32 September 1976 X Number 679 114 2951 146 3890 (3119.4) (322.9) (3233.5) (335.3) (3395.1) ‘7 Volume 45054 27950.3 58148 8447.7 139510 (35350.3) (38071.5) (35495.9) (32815.7) (321545.11 7 Size 55.4 245.3 19.7 57.9 35.9 2) 5 October 1975 Number 2454 7355 5283 133 15245 (+1382 9) (31289.5) (32545.91 (347.81 (33351) 7'Voiume 78209.7 1852566.3 150110.3 9855.3 2090741.5 (342173.7) (3297150.1) (351772.51 (_3543.11 (3401528.91 ‘7 Size 31.7 251.5 23.9 74.1 129.7 3) 12 October 1975 x Number 554 3771 5902 53 15290 (359) (3855.4) (34908.2) (310.91 (34902.31 7 Vo1ume 44958.3 2200153.3 301315.7 4593 3051121.3 (38305.9) {3204200.3) ‘;733572.i} '32045.51 (3199171.51 7 Size 31 1 250.3 735.3 74.5 199.5 Tab1e 824 (cont'd.). 3426 Greens Diatoms B1ue-greens Cryptopnytes Tota1s 4) g4 October 1976 x Number 177 3293 937 0 4407 (350.4) (3591.8) (3472.1) (39068.6) ‘7 Vo1ume 19927.3 757491.7 25472.3 0 312891.3 (311032.21 (3235795.3) (39371.1) (3257043.6) ‘7 Size 112.5 233.1 27.2 ,0 184.5 Mean of 3 Dates- 7 number 1055 5477 4374 55 11981 (3534) . (3950.3) (31818.8) (3251 (32577.3) ‘7 Vo1ume 47598.4 1505737.1 325533.1 4849 1984917.6 (315341) (3249085.4) (3255057.7) (31849.3) (3}19414.9) 7 Size 44.5 248.1 74.5 74.5 155.7 . 8175 92401355 1) Initia1 Readings 30 Seotember 1975 7 Number 177 38 3827 51 4093 (338.1) (321.7) (3113.7) (315.5) (3292) 7 Vo1ume 13481.4 35355 85737.7 3754.7 139329 (31952.9) (_20202.9) (31453.5) 131228.91 (312815.91 ‘7 Size 75.2 930.4 22.7 73.5 34.0 2) 3 October 1975 7 Number 101 70 1554 25 1750 (343.9) (339.8) (3335.9) (316.6) (3314.3) '7 Vo1ume 24159.7 52541.3 24921.7 1377.3 113500 (311051.81 (31113.41 (358.11 (31228.9) (3,1130 8) 7 Size 239.2 893.4 15.9' 75.1 54.5 3) 30 October 1976 7 Number 95 120 545 70 931 (318.3) (334.9) (3238.1) (316.5) (+207 3) 7 Voiume 7319 3 107518 7 11282 5152 131382 (31350 5) (373401.31 (33759.51 (31229: (_71048.4) 7 Size 17.0 895.8 17.5 73.7 141.1 4) 22 October 1976 x Number 145 101 554 25 833 (345.2) (312.5) (3215 3) (312.51 (3232 6) 7 Vo1ume 15254.3 92742.7 8103.2 1577 118987.7 (310391 4) (324552.2) (34799.51 (3929) (322413) 7 Size 111.4 918.2 14.4 75.1 142.5 Mean of 3 Dates- ? Number 111 7 925 '10 9.775 {~15_;) 13:4.4) ’+209.91 (310.7) {3159.3} 7 Yo1ume 15914.1 37534.3 14759 2 72.4 721290.? “+4814.71 (313125) T33562.9) :3‘93.31 :3305:.41 7 Size 139.5 303.4 15.0 74.‘ 103.1 A e6 .5... o.— —.v can 9.3 o...K e4: 9...:2, m6— o.n m6 9m 9% ed.” 52.. me .843: mcmocu Am 945. eeeebuc ~_-o_ .m c._ m.o ¢.— m.c n.c ~.c ..o m.c ae=_o> e._ e.c a.c 0.: —.c ..o ..o ..c genes: meaxgacaaxgu A: c.- «K c6 N6 ~.mm «.3 at? min 9.3—4; 9mm 9mm Ndm adv mg: 98 «IE 58 39.52 .mcmugoiwim Au ..mm o.ma a.am c.nx m.c e.e m.m m.m we:_c> o.v m.mv o.—o «.me ~.o ~.c m.c e.c . gcasaz 42548.; A; m.—m ~.m ~.~ ~.o m._m ~.~m c.mm N.cc as:_c> N.m ~.m_ m.e ..m o._~ e.w— ..m_ m.mm Loés:. mcmagw Ac 95o. guaouuc min AN ~.~ o.c ~.w o.m ~.c ~.c ~.c m.o oe:_o> n._ N.m m.n c.n ..c ..c ..a ..c Lyssa: meuzgzouaxgu Au 7] «0.. 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N. a. b can 5:. e. o .2: .23 .23 :3 .23 .23 .33 2:... m. .m .m N. no. am. new new .4: .c2. .cfi. .mm. .mfi. .9h. .mm. 2.3. e. m.~ mNN 0:. mm. ... 55. c.. .em. .3“. am: sea. ?em. 24: .ezd. .22. .m ea. co. eeN cm. aeN em. ... ...“. ..4u. .N... .54“. .45“. ..o... .45.“. .meq. cc. ... ON. ... use NN. ..m be. waits...“ woo... mac: :3... 35:—.... 3.5.3.. mac... 3o... :35 acct—=5 9.: :35 a acct—=5 9.5 :35 a 33:25 €82: 64...: «2:. ..<. .zc ..e. «E» act—:5 he m9:_:3.::_.x= 5.: 2.22% c5. ......_..3 ...... ......Er... ... 3m . __=\..._..: 5.5.... ...—:1 ...—....53. ... 5.2.3.7.: ...;— Lcnc..uc em.mm : o.e. 25.5.5; N.-=. .. 6.5. ca;b.uo m-m .m c.e. besee.eem mmn:~1.. ......Eaflmaflua U 5.5. Leceauc e..~m .e a... Leee.uc ~..c. .. b.a. 58.5.5: .-. .4 0:..— ..w...... as NNicmiC 2.2.823 = c.e. ceee.uo e.-.“ .e 6.5. 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S... .235 ..-... .. ...... ....n. .34. .2“. .2... .8... ...q. .3“. m. 2. .... .m. .... 2.; E. .... ...... .838 m-.. .. .2... ...... .3... ...... ...... .33 ...... .m... .m 3. 3. ... c... ... .... ...... o... ..3.E%M-..-c.b 3:35.55. .< <..=.c..... ... 3:38.. 39.. mac... :3... moot—.2. $5.20.. mac: 38.. :33 39.2.2. 8.... ..8.... a moot—:5 8.5 com... a 39:22. 69:: "8...: .. ......55. c... 2...... 7ao1e 327. 335 Phytoplanxtonic diversities (in nats) in imported 1axe water and aquaria witn mono- cultuged bullfrogs, a mixed-soecies culture. or monocultured green frogs in Experiment Three at Lake ne. HIERARCHICAL DIVERSITY Number EXPERIMENT of THREE Genera Evenness LAKE ONE Greens Diatoms Blue-greens Total (N6) (0‘) A. BULLFROGS 1) Initial Readings 22 Seotember 1976 a) Number 0.992 0.844 0.791 1.980 15 0.402 6) Volume 1.314 0.756 0.689 1.516 0.565 2) 5 October 1976 a) Number 1.053 1.025 0.717 1.170 19 0.415 b) Volume 1.179 0.941 0.669 1.586 0.601 3) 12 October 1976 a) Number 1 399 C 977 0 703 1.72d 22 0.590 b) Vo1ume 0.990 0.374 0.733 1.523 0.557 4‘ 24 October 1976 a} Number 1.412 0.47% 0.625 1.541 22 0.537 b) Volume 0.728 0.‘lS 0.969 1.200 0.418 Vean of 3 Dates- a) Number 1 288 0.825 0.682 1.178 2, 0.514 6) Volume 0 969 0.810 0. 9 1.505 ‘ 0.526 8. MIXED: BULLFROGS AND GREEN FROGS 1) Initial Readings 22 Seatember 1976 a) Number 0.884 0.684 0.904 1.377 13 0.40. b) Volume 1.217 0.711 0 707 1.533 0.573 2) 5 October 1975 a) Number 1.264 1.016 0 551 1.240 19 0.36: b) Volume 1 151 1 097 0.563 1.73 0.616 3) 12 October 1976 a) Number 1.307 0.935 0.747 1.666 20 0.584 b) Volume 1.013 0.784 0.703 1 655 0.580 4) 24 October 1976 a) Number 1.613 0.778 0.798 1 623 32 0.558 b) Volume 0 777 0.920 0.946 1.318 0.454 Mean of 3 Dates- a) Number 1 395 0.910 0.599 1 509 93 0.502 b) Volume 0.980 0.934 0.737 1 568 ‘ 0.550 C. GREEN FROGS 1) Initial Readings 22 SeDtember 1976 a) Number 1.046 0.387 0.905 1.103 73 3.404 5) Volume 1.28d 0.955 0.568 1.515 3.556 21 5 October 1976 a} Number 195 1.145 0.537 1 Z73 71 0.141 3) Volume 154 ‘ 180 0.648 1 319 " 0.530 Table 827 (cont’d.). 336 Greens Diatoms 81ue-greens Total (N5) (0') 3) 12 October 1976 a) Number 1.349 1.039 0.784 1.777 23 0.592 b) Volume 1 075 0.981 0.793 1 704 0.567 4) 24 October 1976 a) Number 1.563 0.849 0.515 1.524 79 0.538 b) Vo1ume 0.836 0.876 1.006 1.246 0.440 Mean of 3 Gates- a) Number 1.36- 1.011 0.645 1.525 2] 0.524 6) Volume 1.025 1.012 0.816 1.590 0.546 0. LAKE READINGS 1) Initial Readings 20 September 1976 a) Number 0.770 0.878 0.517 1.025 19 0.362 0) Volume 1.113 1.065 0.732 1.598 0.564 2; 3 October 1976 a) Number 1.208 1 075 3.381 1.115 31 0.391 b) Volume 1.242 1.243 0.318 1.760 0.671 3) 10 October 1976 a) Number 1.355 1.177 0.555 1 533 23 0.545 b) Vo1ume 1.072 1.226 0.684 1.781 0.595 4) 22 October 1976 a) Number 1.595 0.70 0.723 1.702 75 0.575 b) Vo1ume 0.334 0.946 1.109 0.992 7 0.499 Mean of 3 Dates- a) Number 1.386 0. 87 0 586 1.483 23 0 504 b) Volume 1.049 1 138 0.704 1.511 0.570 337 Table 828. Phytoolanktonic diversities (in nets) in imported lake water and aquaria with mono- cultured bullfrogs. a mixed-Sbecies culture, or monocultured green frogs in Experiment Three at Lake Four. HIERARCHICAL DIVERSITY Number EXPERIMENT of THREE Genera Evenness LAKE FOUR Greens Diatoms 31U8~9reens Total (N6) (0') A. BULLFROGS 1) Initial Readings 22 September 1976 a) Number 0.711 0 0.460 1.091 8 0.581 b) Volume 0.688 0.399 1.446 0.767 2) 5 October 1975 a) Number 1.186 0.05 0.341 1 60 21 0.506 b) Volume 1.165 0.130 0.371 9 1 0.303 3) 12 October 1976 a) Number 1.235 0.007 0.764 0.723 15 0.297 b) Volume 0.899 0.035 0.755 0.335 0.137 1) 24 October 1975 a) Number 1.127 0.023 1.036 0.775 15 0.384 b) Volume 0.829 0.053 0.926 0.324 0.144 Mean of 3 Dates- a) Number 1.256 0.029 0.380 0.953 17 0.396 b) Volume 0.964 0.072 0.857 0.490 0.195 8. MIXED: BULLFROGS AND GREEN FROGS 1) Initial Readings 22 Seotember 1976 a) NUMDEP 0.851 0 0.582 1.373 10 0.576 b) Volume 0.819 0.510 1.695 0.834 2) 5 October 1976 a) Number 1.381 0.058 0.924 1.169 20 0.428 b) Volume 1.406 0.127 0.768 0.549 0.201 3) 12 October 1976 a) Number 1.232 0.027 0.792 0.449 15 0.188 5) Volume 0.780 0.918 0.496 0.266 0.113 4) 24 October 1976 a) Number 1.013 0.053 0.884 0.942 14 0.412 b) Volume 0.946 0.176 0.320 0.458 0.199 Mean of 3 Oates- a) Number 1.209 0.045 0.867 0.853 17 0.343 b) Volume 1.044 0.132 0 695 0.421 0.171 C. GREEN FROGS 1) Initial Readings 22 Seotember 1976 a) Number 0.333 3 0 559 1 398 - 0.653 6) Volume 0.564 0.392 1 572 0.808 3) 5 October 1975 a) Number 1.122 0.121 .3 3 1.194 :3 1.458 3) Volume 1.145 0.229 0.597 0.599 0.267 338 Table 828 (cont‘d.). Greens Diatoms Blue-greens Total (N6) (0') 3) 12 October 1976 a) Number 1.242 0.042 0.582 0.736 17 0.319 b) Volume 0.995 0.134 0.593 0.519 0.212 4) 24 October 1976 a) Number 0.846 0.097 1.074 0.949 11 0.432 b) Volume 0.708 0.359 1.002 0.624 0.284 Mean of 3 Dates- a) Number 1.170 0.087 0.673 0 977 '6 0.403 6) Volume 0.949 0.241 0.764 0 514 0.254 0. LAKE READINGS 1) Initial Readings 20 September 1976 a) Number 0.762 0.462 0.136 0.460 10 0.237 b) Volume 0.441 0.260 0.189 1.055 0.534 2) 3 October 1976 a) Number 0 6.9 0.656 0.135 0 564 11 0.357 b) Volume 0 ’34 0.457 0.161 1 430 0.786 3) 10 October 1976 a) Number 0.737 0.333 0.168 0.259 9 0 581 6) Volume 0.506 0.273 0.194 1.157 0.630 4) 22 October 1976 a) Number 0 .52 0.631 0.695 1.568 12 0.790 b) Volume 0. 04 0.390 0.698 1.109 0.519 Mean of 3 Gates- a) Number 0.783 0.540 0.333 0.864 11 0.609 b) Volume 0.681 0.373 0.351 1.235 0.645 339 Table 829. Gross primary productivity as mg 0 /42.8 liters/24 h (2 SD) during Experiments One and Two at Lakes One ana Four. SAMPLING UNSTOCKED LOW HIGH DATES CONTROL BIOMASS BIOMASS Lake One--Experiment One 5-28-1976 167.5 (:10.6) 172.1 (112.3 158.9 (119.2) 6-6-1976 341.8 (:34.7 261.7' (:15.5 246.1 (:38.1 6-12-1976 804.9 (+56.1) 697.3 (174.4) 729.6 (154.2) 6-19-1976 337.9 (:174.3) 232.0 (:16.8) 261.4 (194.4) 73 Dates 494.7' (194.2) 397.0 (178.9) 412.4 (:81 7) Lake Four--Experiment One 5-28-1976 159.8. (+35.3) 132.8' (:5.9) 162.8 (+31.5) 6-6-1976 20.6 (34.4; 23.9 (32.5 19.1' 148.1; 6-12-1976 191.4 (:2.5 211.4 ($40.0. 240.1 (gTb.9 6-19-1976 38.3 (112.0) 51.4 (111.5) 36.0 (313.3) 73 Dates 83.5 (:27.3) 95.6 (:29.5) 98.4 (:71.1) Lake One-~Egperiment Two 7-26-1976 154.7 (112.7) 234.3‘ (352-4)" 173.8 (+63.1) 8-8-1976 990.7 (:57.1; 913.7 (188.8 854.5 (+T1o.9) 8-15-1976 532.4 (294-5 585.7‘ (151.7 547.8 Ti51.7) 8-22-1976 163.4 (188.6) 536.2 (292-1) 416.7 (:75.6) 75 Dates 562.2 (:121.7) 678.50 (168.6) 606.3 (:77.0) Lake Four--Experiment Two ' 7-26-1976 102.8 (+17.7) - 80.0 (:7.1)‘ 87.6 (:9.0) 8-8-1976 68.8 199.1) 57.6. (+2.8; 52.7 (+2.9) 8-15-1976 150.3 (+19.8) 114.63 (+18.2 108.4 (+14.6 8-22-1976 87.8 128.2) 80.9 '019.7) 76.0 .13.5) 73 Dates 102.3 (114.0) 84.4 (19.5) 79.1 (_+_9.2) TabTe B30. . Community respiration as mg 0 dur1ng Experiments One and Two at Lakes Ofie and Four. 340 /42.8 liters/24 h (1 SO) SAMPLING UNSTOCKED Low HIGH DATES CONTROL BIOMASS BI QMASS Lake One--Experiment One 5-28-1976 263.5 (351.7)' 264.6 (322.5) 235.5 (335.0) 6-6-1976 332.7 (340.9) 304.4 (326.8) 277.4 (3J2.9)1 6-12-1976 821.7 (+67.3) 727.2 (+72.8; 741.0 (346.7) 6-19-1976 413.2 (3789.3) 360.0 (36.6 379.9 (355.0 75 Dates 552.5 (396.0) 464.0 (370.0) 466.1 (371.8) Lake Four--E5periment One 5-28-1976 177.8 (+40.5) 134.28 (+8.8) 177.3 (339.6) 6-6-1976 32.6. (32.7) 57.0 (374.7) 106.8 (+14.0) 6-12-1976 171.8 (+7.1) 203.8 (3J3.0) 237.8 117.5) 6-19-1976 98.5 (321.7) 132.6 (320.5 122.0 (310.8) '73 Dates 101.0 (321.1) 131.1 (322.7) 155.5 (321.9) Lake One--§§periment Two 7-26-1976 131.4 (350.5) 181.0 (355.7) 135.8 (364.4) 8-8-1976 658.7 (312.5) 595.6 (393.7) 588.8 (360.1) 8-15-1976 615.4 (326.1) 611.8 (333.5) 568.0 (321.5) 8-22-1976 837.1 (351.8) 1001.7 (320.2 921.1 (350.4 73 Dates 703.8 (_67.9) 736.4 (372.5) 692.6 (361.8) Lake Four-~Experiment Two 7-26-1976 119.0 (39.7) 101.6 (39.8; 112.7 (+11.9 8-8-1976 47.0 (+7.3) 42.3 (35.3 42.6 (+5.8 8-15-1976 97.0. (+T6.3) 77.1 (+7.0) 76.4 (+T1.0 8-22-1976 115.4 T3§.6)‘ 102.2 (3T2.8) 97.1 ‘T33.4) 73 Dates 86.5 (3fl1.6) 73.9 (39.7) 72.0 (38.7) 341 Table 831. Community respiration (R) in unstocked aquaria and cor- rected respiration (”R" without tadpoTe respiration) in stocked aquaria as mg Og/42.8 Titers/24 h during Experiments One and Two at Lakes One and Fou . SAMPLING UNSTOCKED LOW HIGH DATES CONTROL BIOMASS BIOMASS Lake One--Experiment One 5-28-1976 263.5 (311.7) 264.6 (322. 5) 235.5 (+35.o) 6-6-1976 332.7 (340. .9) 251.7 (+18. 7 189.2 735.4) 6-12-1976 821. 7 (+67. 3 672.0. (+63. 4 661.1 (337.9 6-19-1976 413. 2 (3189. 3) 301.5 (34. 8) 311.0 (310.2) '73 Dates 522.5 (3151.5) 408.4 (3132.6) 387.1 (3143.5) Lake Four--Experiment One 5-28-1976 177.8 (+40.5) 134.2 (38.8) 177.3 (+39.6) 6-6-1976 32.6 732.7 14.4 (+4. .0) 18.0 Ti2.5; 6-12-1976 171 8 (37.1 167.5 (+9. 4 161.4 (39.0 6-19-1976 98.5 (321.7) 112.1 (3T5. 0) 79.2 (34.7) '73 Dates 101.0 (321.1) 98.0 (349.7) 86 2 (351 4) Lake One--Experiment Two 7-26-1976 131.4 (350. 5) 181.0 (365.7) 135.8 (364.4) 8-8-1976 658.7 (312.5) 558.6 (390.2) 526.5 (357.8) 8-15-1976 615.4 (326.1) 557.3 (+30.0) 489.9 (317.9 8-22-1976 837.1 (351.8) 927.3 (318.0) 825.0 (343.9) 73 Dates 703.8 (367.9) 681.1 (3111.8) 613.8 (398.3) Lake Four--§xperiment One 7-26-1976 119.0 (39.7) 101.6 (39.8) 112.7 (+11.9) 8-8-1976 47.0 (I7. 3) 38.3 (34.4 30.0 T33.0) 8-15-1976 97.0 (3T6. 3) 73.0 (+6.4 73.5 (38.8) 8-22-1976 115.4 T36. 6) 98.1 (311.9) 82.0 (33.0) '73 Dates 86.5 (311.6) 69.8 (317.3) 61.8 (318.7) 342 Am~.Fv A_N._V Lam.ov Aoo._v o... a... a... mm.c Na.o om.o maeee m to eeaz Amm.ov Amm.ov Apm.ov Amm.cv m~.o m~.o eL.o me.o mm.o mp.o em=m=< mm Ane._v Ahm._v AN.._V Amo._v Ne._ ae._ mm._ 83.9 em.o em.o em=m=< m_ AeL._. 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