DOMINANT PLANKTONIC ROTIFERS OF MAJOR WATERWAYS OF THE UNITED STATES Louis G. Williams1 US. Public IIealth Service, R. A. Taft Sanitary Engineering Center, Cincinnati, Ohio ABSTRACT Rotifers were the most numerous metazoans found in preserved plankton samples taken routinely at 128 sampling stations on the major rivers and Great Lakes of the United States. While rotifers of more than one genus often dominated a sample, the dominant form tended stations with high population densities had low species to be one species. Generally, diversity. Turbulence, silt, and some other cdaphic factors appeared to be more important than industrial or domestic wastes, during winter and early spring, in reducing the number of species of rotifers in the biota. During the summer and fall, differences in the abundances of dominant species appeared to be the best criterion for indicating differences in water quality or pollution. METIIODS INTRODUCTION AND MATERIALS Type and preservation Inadequate knowledge of the zooplankton of rivers is a major impediment to a better understanding of the trophic relationships governing biological productivity of the of the United States major wateiways (‘Williams 1962). Beach ( 1960) reported 34 species in 24 genera of rotifers in plankton from the Ocqueoc river system of Michigan and, like Chandler ( 1937)) found that rotifers in outlets of lakes and impounded waters gradually dccreasc in numbers downstream. These studies indicated that planktonic rotifer fauna of lotic environments are not qualitatively distinct from those of the lentic environment. The common river species have been reported, but no systematic attempt has been made to quantify them for the major rivers on a nationwide basis or to relate them to river environments. This report is based on data taken from semimonthly plankton samples at 88 of 128 stations of the National Water Quality Network of the US. Public Health Service during the last two years of a five-year study ending November 1962. Only those stations were included that had been in operation two or more years. Acknowledgment is given to Mr. Albert Katko, who, during 1961 and 1962, did much of the routine enumeration of the rotifers. of samples In 1958-1959, an extensive study of fresh and variously preserved plankton samples from 19 widely scattered sampling stations established that many of the delicate protoeoa and a few of the more delicate rotifers lost their identity when they died or when preserved with formaldehyde. After various preservatives and their combinations were tested, thimerosal (merthiolate) 0.007%, buffered with sodium borate and containing 0.1% iodine (potassium iodide and iodine) was found to be the least destructive to plankton and was adopted as the standard preservative. After the samples arrived at the counting laboratory in Cincinnati, iodine was occasionally added to them to enhance settling of dense plankton blooms or to prolong preservation. During the first 18 months oE this program, many planktonic forms arrived in a more readily identifiable condition when alive than when preserved with 5% formaldehyde. Major alterations of samples occurred between the time of collection and counting, preventing the use of live samples for quantitative mcasuremcnts. Although neither 5% formaldehyde nor thimerosal-iodine preservative proved to be completely bacteriostatic, the latter arrested ccl1 division in phytoplankton and zooplankton, and the samples were not significantly degraded by bacterial action if they were processed within a month. Since samples were processed routinely within l Present address: FWPCA, National Water @ality Research Laboratory, Duluth, Minnesota 55812. 83 84 LOUIS TABLE 1. Population densities G. WILLIAMS of rotifers in the major waterways of the United States A Rotifers in semimonthly plankton samples, counts/liter, 1961 and 1962 Basin, river, or lake, and station Colorado River Basin Animas River, Cedar Hill, New Mexico Colorado River, Yuma, Arizona Parker Dam, Arizona Boulder City, Nevada Page, Arizona Loma, Colorado Southwest-Lower Mississippi River Basin Arkansas River, Pcndleton Ferry, Arkansas Ponca City, Oklahoma Coolidge, Kansas Mississippi River, New Orleans, Louisiana Delta, Louisiana West Memphis, Arkansas Cape Girardeau, Missouri Ouachita River, Bastrop, Louisiana ( 1962) Red River, Alexandria, Louisiana Index, Arkansas Dcnison, Texas 17.3 1.8 28.3 0.7 0 0.7 0.3 0.5 0.1 1.0 0 0 0.5 0.5 1.6 0.4 0 0 0.1 0.2 32.2 0 0 0 1.7 0.6 0.1 0 0 0 8.9 0 0.8 0 0 0.3 11.5 2.0 34.9’ 1.5 12.7 43.8 5.1 1.2. 1.0 3.4 2.8 127.4 62.5 33.8 18.0 1.6 75.0 0.1 0.1 0.8 0.4 1.4 6.8 21.4 0.5 15.9 0.2 0.2 0.5 0.7 1.6 1.8 0 0.5 0 0.5 0.1 1.1 0.6 0 0.4 0 0 0 0 0 0 3.3 1.0 0.1 0 0.8 0 12.5 79.6 1.4 1.4 1.2 4.0 4.0 97.1 27.9 9.2 3% 14.4 1.8 G 1.3 0.1 fi 188.7 3.7 68.8 0 16.7 11 96.9 0.5 74.1 65.8 5.5 45.2 191.2 ; 12 4 6 7 5 476’77 12:s 6.3 38.6 11.0 0’9 10 0:l 0.2 - 546 7’2 8:6 1.9 - 198 1’7 0:4 0 - 36 0’4 0:l 0 - 216 0’ 100.1 10.4 0.2 9.5 1.2 3.4 - 3 1 5.7 0.4 3.0 0.4 10.0 0.1 0.1 0 0 0 ; 8 1 0 0 2’8 11 5416 3.0 0.2 0.7 0.7 05 2’3 012 l:o 11417 0 1.0 0 0 _ - 07 3818 4.2 0 _ _ 04 310 0 0.1 _ 0 42.05 30.8 188:5 0 5.8 0 0.2 - :: 220.1 329.0 136.9 134.9 5.7 41.7 38.2 29.0 50’67 179 47.04 65 11.15 119’5 50 6 29 82.1 0.5 2.4 5.9 23.1 714.4 3.3 77.8 24.2 1.1 345.1 101.4 1.2 13 159:2 104:3 175:3 4.0 105.1 161.3 29.7 3.9 18.2 324.6 0.9 168.2 13 3 0 3 4 4 6 2 3 3 3 7 7 Pacific Northwest Basin Columbia River, Clatskanie, Oregon Bonneville, Oregon Pasco, Washington Wenatchcc, Washington Clearwater River, Lcwiston, Idaho ( 1962) Snake River, Wawawai, Washington Payettc, Idaho ( 1962 ) Yakima River, Richland, Washington ( 1962 ) Missouri River Basin Missouri River, St. Louis, Missouri Kansas City, Kansas St. Joseph, Missouri Omaha, Nebraska Yankton, S. Dakota Bismarck, N. Dakota Williston, N. Dakota N. Platte River, Henry, Nebraska (1962) Platte River, Plattsmouth, Nebraska ( 1962) S. Platte River, Julesburg, Colorado ( 1962) Big Sioux River, Sioux Falls, S. Dakota ( 1962) Upper Mississippi River Basin Illinois River, Peoria, Illinois Mississippi River, E. St. Louis, Illinois Burlington, Iowa Dubuque, Iowa St. Paul, Minnesota Red River, Grand Forks, N. Dakota (1961) (TABLE 1 continued 9 8 9 2.2 23.3 32.4 on page 86) 47:0 49.3 0 0 13.3 5.0 PLANKTONIC ROTIFERS OF UNITED Jcm STATES WATERWAYS 85 Among the metazoan plankters, cladocand copepods ranked next to the rotifers in abundance, but they were USUally present in such small numbers that they could not be adequately evaluated from a l-liter sample. Insect larvae were present only occasionally, and these forms, which make up a major portion of net plankton, were probably dcrivcd from the benthos. Bottom forms, which were more common in samples from shallow streams, probably represented displaced organisms when found Philoiha, with many in the plankton. benthic spccics, was found frequently in samples from shallolw streams. In all samples from the larger rivers, the most abundant forms were characteristically planktonic. crans T * ONE LITER MARK Collection, RUBBER STOPPE R L POLYETHYLENE DRAIN TUBE RUBBER CONNECTOR DETACHABLE VIAL IpIG. 1. Schem:&ic drawing of spccinlly designed graduated glass settling tube. two weeks, long-term preservation was usually unnecessary. !Study shotwed that the delicate protozoa could not bc counted or identified in prcserved samples, and attempts to include them in the routine plankton enumeration were abandoned. Some protozoa with protective testaceous walls, such as CoduneZZa cratera and Tintinnopsis cylindrata, were abundant in many samples and were well preserved. Because these species feed largely on bacteria, their relative density appears to bc an indication of the population of bacteria present. concentration, and counting of samples Three liters of water were taken directly from a river or lake or other continuously flowing source; for example, from the intakes of water treatment plants. The samples were shipped to the counting laboratory in Cincinnati, Ohio, where they were agitated and divided into three equal subsamples. The number of centric and pennate diatoms and genera of other algae in one portion were determined; in another, diatoms were identified to species (Williams 1964); in the third, metazoan invertebrates wcrc identified to genus. The portions for invertebrate and diatom analyses were settled in graduated glass settling tubes ( Fig. 1) . These tubes are 70 cm long and 5 cm diameter and taper near the bottom to about 2 cm diameter. To concentrate and clean the organisms for analysis, the sample was thoroughly agitated, poured into the settling tube, and allowed to settle for 18 hr m more. After settling, the water in the upper 50 cm was slowly drained from below until it was at the level of the stopcock inserted in the side of the tube. The rubber stopper and stopcock were then rcmovcd; a small, pointed polyethylene tube was inserted in the opening and the remaining supernatant water was drawn off by vacuum from the top of the water column to 86 LOUIS G. WILLIAMS TABLE 1 ( continued ) Rotifers in semimonthly plankton samples, counts/liter, 1961 and 1962 ____. Five most abundant genera of the network. Averages for 24 samples from May to November Basin, river, or lake, and station Western Great Lakes Basin Niagara River, Lake Erie, Buffalo, New York 9 Detroit River, Detroit, Michigan 13 Lake Huron, Port IIuron, Michigan 11 Lake Michigan, Milwaukee, Wisconsin 9 Lake Michigan, Gary, Indiana 15 Lake Superior, Sault Ste. Marie, Michigan 6 Lake Superior, Duluth, Minnesota 5 Ohio River Basin Allegheny River, Pittsburgh, Pennsylvania ( 1962 1 7 Kanawha River, Winfield Dam, West Virginia 7 Little Miami River, Cincinnati, Ohio 8 Ohio River, Cairo, Illinois 3 Evansville, Indiana 13 Louisville, Kentucky ( 1962 ) 8 Cincinnati, Ohio 10 Huntington, West Virginia 9 E. Liverpool, Ohio Wabash River, New Harmony, Indiana (1962) ; 106.3 22.1 14.1 63.7 45.3 12.0 8.5 10.9 104.2 100.4 1.2 25.5 6.2 0.7 33.1 12.5 1.2 82.0 45.0 0.9 38.6 22.4 0.3 4.2 8.0 0 15.2 3.0 29.0 23.8 21.2 11.8 104.3 84.2 84.0 68.1 74.9 79.0 14.1 4.2 1.2 0.2 0.6 3.8 7.0 119.6 16.4 43.3 17.9 126.0 29.8 125.0 - Northeast Basin Connecticut River, Northficld, Massachusetts St. Lawrence River, Massena, New York Hlldson River, Poughkeepsie, New York 148.1 35.0 21.0 1.4 127.9 0.4 27.7 0.8 22.4 North Atlantic Basin Delaware River, Philadelphia, Pennsylvania Martins Creek, Pennsylvania Schuylkill River, Philadelphia, Pennsylvania Susquehanna River, Conowingo, Maryland Sayre, Pennsylvania Potomac River, Great Falls, Maryland Williamsport, Maryland Shenandoah River, Berryvillc, Virginia ( 1962) Western Gulf Basin Rio Grande River, Brownsville, Laredo, Texas El Paso, Texas Alamosa, Colorado ( 1962 ) Sabine River, Ruliff, Texas 10 ; 11 6 10 8 12 z 8 10 4 Texas ; 7 Southeast Basin Apalachicola River, Chattahoochcc, Florida Escambia River, Century, Florida Savannah River, Port Wcntworth, Georgia N. Augusta, South Carolina Chattahoochec River, Columbus, Georgia Atlanta, Georgia Tombigbee River, Columbus, Mississippi ( 1962) (TABLE 1 continued 13 7 1 2 9 2 6 6.1 1.2 2.5 25.6 6.6 0.3 1.3 5.9 3.4 1.3 1.2 26.6 248.7 3.5 37.0 2.5 50.2 1.5 135.1 1.9 67.3 0.7 19.6 0.3 19.8 zo.0 17.0 25.3 - 3.0 0.9 0.1 11.1 2.2 6.0 0.6 - 6.8 0.2 4.2 43.8 45.1 47.2 4.3 - 65.4 29.5 2.1 22.6 3.0 0 17.6 230.0 3.0 63.7 7.6 33.4 - 34.2 3.7 11.3 207.2 127.1 143.7 185.1 - 31.8 8.2 8.4 0.2 128.5 201.5 7.6 55.0 76.5 0.3 6.5 1.4 3.6 0 44.5 - 35.4 1.2 4.3 64.7 3.4 2.6 0.9 - 6.0 0.3 0.7 33.3 0.5 0.3 0.6 - 1.5 0.2 0.2 5.1 0.1 0 0.1 - 2.8 55.1 0.9 3.3 36.7 66.6 9.1 119.9 5.4 9.8 1.4 5.6 0.2 1.9 - 128.0 22.5 2.5, 10.0 16.5 26.6 0 1.2 0.3 48.7 115.4 0 23.2 0.3 0 3.9 0.5 22.7 0.1 0 0.1 79.7 288.2 18.5 37.3 0.1 1.7 5.4 9.8 225.1 27.4 1.3 3.0 76.3 2.3 20.2 323.6 140.5 0 2.1 1.3 0 0.2 0.3 0 0.7 0.5 15.4 15.4 61.4 0 0.6 0.2 _ - 34.2 on page 88 > 23.8 129.6 342.2 0.3 0.9 4.7 0 0.2 0.8 y.2 282'3 07 477'3 45 0 _ 0:2 - 1:1 - PLANKTONIC ROTIFERS OF UNITED avoid disturbing the settled organisms. By a series of steps, the organisms were carefully washed from the tapered portion of the tube into the detachable vial. E’our washings, with at least 20 min between settling, were necessary to insure a concentrated sample without loss of organisms. Most samples for invertebrato analysis coiuld be counted without further washing, but when ncccssary, the sediment was washed with distilled water to remove unwanted colloidal turbidity and suspensoids. Centrifugation, membrane filtration, and the use of plankton nets were tested as methods of concentrating and washing plankton from the samples, but were abandoned in favor of sedimentation. Studies of samples containing 4 ml or more of silt per liter contained few, if any, mctazoan invertebrates. The entire sediment from samplcs having less than 4 ml of silt was placed in a special microslide, 70 mm long, 50 mm wide, and 2 mm deep. Samples were analyzed as soon as possible after arrival to avoid deterioration. With aging, muddy river samples became increasingly difficult to count. The structural outlines of the organisms became less distinct, the organisms became entangled with silt and debris, and redispersing and washing the sediment was cxtrcmcly difficult or impossible. About 5% of the samples collected during the bloom period of rotifers-May to November-improved with different degrees of washing; about 95% of the samples required no washing. Highly silty conditions prevailed at many of the stations during the November 1961 to May 1962 period, and about 2% of the samples from this period were discarded bccause of excessive silt; data from this period h avc been excluded in the report of the five most abundant genera in Table 1. When the sediment was not excessive, counting was simplified by using the zooplanktcrs in the entire sedimented sample. Random distribution of the sediment in the counting chambe:r was accomplished by thoroughly agitating and then quickly pouring the quantity necessary to fill the counting chamber, The vials containing the washed sediment and STATES 87 T;VATEIW’AYS rotifers were decanted so that the final volume in each of the vials was equal to the volume in the counting chamber. Only a quarter of the chamber was counted when counts reached 200 in one-quarter of the chamber, and only a half was counted when counts reached 100 in one-half of the chamber. Since random distribution in the chamber cannot be obtained for those invertcbrates larger than most of the rotifers, the larger organisms were counted individually by rapidly scanning the uncounted portion of the chamber. The chambers were graduatcd for this partial counting. A compound microscope with a 10 X ocular and a 10 x objective was used, but grcatcr magnification was used to identify the smallest organisms and was essential to make specific identification of Keratellu, Pompholyx, and Gnstropus. Some organisms were difficult to identify because they were contracted. l%spccially designed tally sheets and codes to taxa and their densities wcrc developed to expedite routine counting, rc:porting, and analysis of data (US. Public IIcalth Service 1964). DISCUSSION Distribution AND RESULTS of common genera and species Pennak ( 1957, p. 229) states, “Any aquatic biologist who has spent much time counting quantitative limnetic zooplankton samples has been impressed with the fact that a sample usually contains one species of copcpod, one cladoceran, and one rotifer which are cxccptionally abundant and clearly dominant numerically over the other species in those groups.” In this study, rotifers of more than one genus often dominated the same sample, but each dominant genus strongly tended to have but one dominant species. Table 1 gives the number of dominant genera found at the 88 sclccted stations during the two-year period. For tho first two years of this study, the total numbers of rotifers were reported and no attempt was made to report their taxa. But during this period, 19 widely-scattered stations were selected and studied for rotifer species composition, In this study, 58 species, including a few unknown ones, . 88 LOUIS G. WILLIAMS TABLE 1 (continued) Rotifers in semimonthly plankton samples, counts/liter, 1961 and 1962 Five most abundant genera of the network. Avcragcs for 24 samples from May to November Basin, river, or lake, and station Tennessee River Basin Tennessee River, Chattanooga, Tennessee Bridgeport, Alabama Pickwick Landing, Tcnncssce ( 1962 ) Lenoir City, Tcnncssec ( 1962) 6 3 8 6 23.7 5.2 197.0 54.4 0.4 0.3 I 22.3 2.4 - 16.8 2.1 - 1.0 0.1 - 4.7 3.2 - 45.3 8.2 - California Basin Klamath River, Kcno, Oregon Truckec River, Farard, California 9 5 146.9 51.3 12.0 I 56.5 - 5.5 L 2.6 - 4.9 - 81.6 - 43.8 10.4 32.1 19.4 3.7 14.5 82.1 Average ( 1962) counts per liter were found among the dominants. An attempt was made to exclude washed-in or dislocated metazoa in the plankton samples by selecting the data so that only species occurring two or more times and having five or more individuals per liter were considcrcd as dominant. Keratella cochlea& was by far the most widely distributed and most abundant rotifer. Polyarthra vzdgaris was the second most abundant and widely distributed species. In this pilot study, the five most widely distributed and dominant genera in order of abundance wcro &Yatella, Polyarthra, Brachionus, Synchaeta, and Trichocerca. The kinds and numbers of genera of rotifers were recorded routinely beginning in 1959 and were reported to the Water Quality Network after 1 October 1961. From these many observations, it appears that the one-species domination of rotifer genera for any one sample is valid and provides the necessary information for any practical diversity study of the major rivers of the United States for water quality ecology. Populations of a few of the more dominant species of rotifers found at selcctcd stations from May to November over a three-year period, reported as the average 6.6 number of individuals per liter, arc as follows: Peoria, Illinois, on the Illinois River, Brachionus calyciflorus, 615; Columbus, Georgia, on the Chattahoochec River, Trichocerca Zongiseta, 512; and Chattahoochee, Florida, on the Apalachicola River, POZZJarthra vulgaris, 262. KerateZZa cochlearis averaged 181 per liter at stations on the Ohio River, but the numbers fluctuated with the phytoplankton counts during the rotifer season. Some factors affecting population density Most rotifers disappcarcd during winter and during periods of high stream flow. Rotifer populations were greatest from May to November, Keratekla cochlearis occurred at some stations at all seasons of the year, but in much lower numbers during the colder seasons. In the United States, rotifers begin to multiply earlier in the southern streams. Many species commonly found in rivers were also found at some of the Great Lakes stations, but some of the more common species of the Great Lakes, such as KeZZicottia longispina and Filinia Zon&&n, did not dominate typical river stations. KerateZZa hiemalis became common in the late fall at many cold water stations in the north. PLANKTONIC TABLE 2. Population densities ROTIFERS OF UNITED STATES by rank orders of 12 of 88 sampling and 1962 = 89 WATERWAYS stations May to November Rotifers Algae 25 26 26 26 27 28 28 29 30 30 3I 32 26 27 25 14 12 3 24 17 10 24 1961 - Twelve stations : Ohio River, Cincinnati, Ohio Ohio River, Evansville, Indiana Ohio River, Louisville, Kentucky Connecticut River, Northfield, Mass. Niagara River, Lake Eric, Buffalo, N. Y. Ohio River, Huntington, W. Va. Chattahoochcc River, Columbus, Georgia Mississippi River, Dubuque, Iowa Rio Grande River, Brownsville, Texas Mississippi River, St. Paul, Minn. Apalachicola River, Chattahoochee, Florida Illinois River, Peoria, Illinois All 88 stations: Median ranking Average ranking, lower 50 percent& Average ranking, upper 50 percentile - Repeated observations indicated that small rotifer populations at many stations were probably a result of their intolerance to turbulence and silt. In most streams, dense lo,ads of silt arc associated with increased stream flow. Stations at which highly silty conditions were associated with extremely low rotifer populations were: Yuma, Arizona, and Loma, Colorado, on the Colorado River; Coolidge, Kansas, on the Atikansas River; all stations of the lower Mississippi River; Wawawaii, Washington, on the Snake River; St. Joseph, Missouri, Omaha, Nebraska, and Williston, North Dakota, on the Missouri River; El Paso, Texas, on the Rio Grandc River; and Cairo, Illinois, on the Ohio River. High densities of rotifers are generally associated with water of high clarity and littlle turbulence during the bloom period of :rotifers. These stations usually have average to high phytoplankton populations (Table 2). Some of the rotifer-rich stations are: Alexandria, Lousisana, on the Red River; Clatskanie and Bonneville, Oregon, on the Columbia River; Yankton, South Dakota, on the Missouri River; Peoria, Illinois, on the Illinois River, Burlington and Dubuque, Iowa, and St. Paul, Minnesota, on the upper Mississippi River; Evansville, Indi- 10 23 20 32 31 34 17 32 ana, Louisville, Kentucky, Cincinnati, Ohio, and Huntington, West Virginia, on the Ohio River; Northfield, Massachusetts, on the Connecticut River; Conowingo, Maryland, on the Susquehanna River; Brownsville, Texas, on the Rio Grande River; Chattahoochee, Florida, on the Apalachicola River; and Columbus, Georgia, on the Chattahoochee River. Many of these stations have high turbidities from November to May, when rotifers are practically absent, and so the plankton data for this period were excluded for the best ecological comparisons. Because the magnitude of rotifer polpulations was usually associated with the density of the phytoplankton populations (Table 2), exceptions are of ecological interest, In the downstream stretches of some large rivers, consumer populations tend to become more prominent where conditions toxic to them do not exist and may actually reduce the phytoplankton populations and their detritus (Williams 1964). This is important in stream management, because heavy blooms of algae and their detritus may be controlled by favoring conditions for consumer populations, such as rotifers. For instance, Cincinnati, Ohio, was noted for high phytoplankton populations while 90 LOUIS C. WILLIAMS Louisville and Evansville ( downstream ) had lower phytoplankton densities and relatively higher rotifer populations before Markland Dam became operational below Cincinnati in 1962. Rotifer populations fluctuate seasonally, and being small ( or absent) during winter, yearly averages can be misleading. For comparative purposes, it is best to study rotifer populations when populations are normally high, Table 1 shows yearly averages far lower than averages for the six months of high rotifer abundance, These data show that the Colorado and Missouri river basins are generally rotifer-poor, while the upper Mississippi and Ohio river basins are rotifer-rich. But many of the rivers have both rotifer-rich and rotifer-poor stations, One shallow water station having high species diversity and moderately high populations is Cedar Hill, New Mexico, on the Animas River. At this station, benthic rotifers were usually common in the plankton samples. Stations with both reasonably high counts and high species diversity are Clatskanie, Oregon, and Wenatchee, Washington, on the Columbia River; Grand Forks, North Dakota, on the Red River; Evansville, Indiana, on the Ohio River; Conowingo, Maryland, on the Susquehanna River; and Chattahoochee, Florida, on the Apalachicola River. The three stations with the highest populations of both phytoplankton and rotifers were: Peoria, Illinois, on the Illinois River; St. Paul, Minnesota, on the Mississippi River; and Brownsville, Texas, on the Rio Grande River (Table 1). Comparison of phytoplankton and rotifer data indicates that stations with high phytoplankton po,pulations generally have high rotifer populations, and the opposite relationship is also found, suggcsting a direct or indirect dependence of rotifers on the phytoplankton for food. Population densities were ranked for 88 stations from which there were two years of *data from May to November 1961 and 1962 for both rotifers and algae (Table 2). The average rank of the 12 stations with highest rotifer density was 28 and the same 12 stations averaged 24 when ranked for algal density, indicating that stations with high rotifer abundance also have high algal abundance. Some idea of the richness and variation of the rotifer fauna may be obtained from Table 1. Stations having 12 or more different genera of rotifers containing five or more individuals per liter from 48 semimonthly samples for two years are: Cedar IIill, New Mexico oa the Animas River; Wenatchee, Washington, Columbia River; St. Paul, Minnesota, Mississippi River; Grand Forks, North Dakota, Red River; Detroit, Michigan, Detroit River; Gary, Indiana, Lake Michigan; Evansville, Indiana, Ohio River; Sayre, Pennsylvania, Susquchanna River; and Chattahoochee, Florida, Apalachicola River. This list contains stations with enough algal abundance to support a rich rotifer fauna. A number of stations, such as Great Lakes stations at Duluth, Minnesota and Sault Ste. Marie, Michigan, did not have enough algae to support a dense population of rotifers and thus represent very clean stations. Several of the other stations with low populations of rotifers but with populations of phytoplankton sufficient to support much denser po,pulations of rotifers may indicate conditions toxic to the rotifers. Most of the stations are exclusively freshwater, but a marine influence is evidenced in a few coastal stations where brackishwater diatoms occur and where rotifer po,pulations are far below average. High chlorides at some stations on the Arkansas and Rio Grande rivers ( US. Public Health Service 1964) completely destroy the rotifer fauna and drastically reduce chloride-intolerant species of diatoms. Both phytoplankton and rotifer populations are much reduced by industrial chemicals in the lower Kanawha River (U.S. Public Health Service 1964 ) . SUMMARY 1. Rotifers were the most numerous metazoans found in plankton samples from 128 stations on the major rivers and Great Lakes of the United States. The ratio of rotifers to other metazoans was approximately 30 to 1. PLANKTONIC ROTIFERS OF UNITED 2. While more than one rotifer genus often dominated in a sample, each dominant genus tended to have only one dominant species at any one time. 3. Stations with a high population den- sity of rotifers with low spccics diversity were found widely. 4. There is no evidence to support the indicator-organism concept among the dominant rotifers present, regardless of what species compose the population, Rather, the presence of dominants appeared to be d&errnined by the principal cdaphic factors and their abundance by the amount of available food. 5. Stations with high rotifer populations were generally correspondingly rich in phytoplankton. STATES WATERWAYS 91 REFERENCES BIZACII, N. W. 1960. A study of the planktonic rotifers of the Ocqucoc River system, Presque Isle County, Michigan. Ecol. Monographs, 30 : 339-357. CIIANULER, D. c. 1937. Fate of typical lake plankton in streams. Ecol. Monographs, 7: 445-479. PENNAK, R. W. 1957. Spccics composition of limnetic zooplankton communities. Limnol. Oceanog., 2 : 222-232. U.S. PUBLIC HEALTH SERVICE. 1964. National Water Quality Network, Annual compilation of data from Oct. 1, 1961 to Sept. 30, 1962. U.S. Public Health Service Publ. No. 663 (1962 edition). Washington, D. C. 909 p. WILLIAMS, L. G. 1962. Plankton population dynamics. U.S. Public Health Service Publ. No. 663, suppl. 2. Washington, D. C. 93 p. -, 1964. Possible relationships between plankton-diatom species numbers and water quality estimates. Ecology, LE.5: 809-823.
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