ill HIM l \ IHHHI NM 145 688 ”THS WATER SOILS IN RELATION TO LAKE PRODUCTIVITY Thesis {or the Degree of Ph. D. MICHIGAN STATE COLLEGE Eugene W. Roelofs 1941 TH WATER SOILS IN RELATION TO LAKE PRODUCTIVITY by F. nigh/I tG’J Eugene W3 Roelofa A THESIS Submitted to the Graduate School of Michigan State College of Agriculture and Applied Science in partial fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of 80113 1941 TABLE OF CONTENTS. Introduction ........................................... 1 Available plant nutrients in lake soils ................ 3 Fertilizer plots at Rose Lake .......................... 1? Greenhouse fertilizer tests ............................ 22 A comprehensive study of four lakes .................... 25 " Methods 25 Tabulation and treatment of data ............. 27 Data presented .................................... 31 Lake Fifteen ................................. 31 Lake Twenty-two .............................. 41 Little Wolf Lake ............................. 51 Tee Lake ..................................... 59 Discussion ............................................. 69 Bottom fertility .................................. 69 Organic matter content ............................ 73 The ecology of aquatic plants ..................... 76 Relative productivity of the four lakes ........... 84 Productivity as indicated by fish growth studies .. 85 Summary ................................................ 88 Literature Cited O...OOOOOOOOOOOOOOOOO0.0000000000000000 91 H” (U INTRODUCTION In June, 1958, an investigation of water soils in relation to the production of vegetation in lakes was begun by the Soils Section of the Michigan Agricultural Experiment Station. In September of that year, the project was formally organized with the following agencies cooperating: the State Department of Conservation, through the Institute for Fish- eries Research at Ann Arbor. agreed to cooperate in the study to the extent of maintaining the author on their payroll as a Fisheries Research Technician. and lending assistance by .making information in their files available. and by aiding otherwise in carrying out the investigation; the Conservation Institute at Michigan State College assumed the responsibility of providing field equipment and all field expenses; the Soils Section of the college provided laboratory, greenhouse. and office facilities. The author is grateful to Director An S. Hazzard of the Institute for Fisheries Research. Director L. R. Schoen- mann of the Conservation Institute, and Dr. C. E. Miller, Head of the Soils Section, for their part in making this study possible through the cooperation of their respective organizations. Gratitude is also expressed to Professor J. 0. Veatch, who originated the project and under whose immediate supervision the work was done. and to Dr. C. J. D. Brown of the Institute for Fisheries Research, who assisted in out- lining and supervising the study. The author benefited greatly by frequent conferences with Dr. C. H. Spurway of the Soils Section and Dr. H. T. Darlington of the Botany Depart- ment, who checked the identification of the aquatic plants. The growth studies of fish were made by Mr. W. C. Beckman. Er. E. L. Cooper assisted with the more laborious field work during the past summer. Biologists realize that any form of life in a lake, that is, the fish or the vegetation, is largely a product of environment. The objective of this investigation was to de- termine the importance of the soil factor in the environment, so that in subsequent studies of lake productivity or perhaps lake classification, the nature and extent of the various bottom types in a lake may receive due consideration in the light of their influence on the biota of that lake. AVAILABLE PLANT NUTRIENTS IN LAKE SOIIS * In a classification of soils a wide range of conditions must necessarily be accommodated, particularly with regard to moisture content. This range includes desert conditions on one extreme and swamp or marsh conditions on the other. In the latter group are embraced many soils which may be season- ally inundated and are at or near the saturation point during all seasons of the year. A complete classification should be still more inclusive so as to include those soils which are permanently inundated--occurring in lakes, ponds, and streams. Veatch (10) proposed an extension of the soil taxonomic system which included these lake—bottom soils and applied to them the term "hydrosol". The study of hydrosols, however, is not only the concern of the soil taxonomist but it has attracted the interest of the group whose objectives are to study and manage the aquatic life in the lakes--particularly fish and the "higher“ vegetation. The aquatic biologists, then, have recently given more atten- tion to hydrosols in conjunction with their more comprehensive studies of the lakes throughout the country. In our own state this work.has been carried out by the Institute for Fisheries Research, the research branch of the Fish Division of the State Conservation Department. They have made, during the last several years, reconnaissance surveys of many lakes in all parts of the state, continuing and expanding the work pre- viously suggested and initiated by the Land.Economic Survey. *Vlaterial under this heading has been previously published as a part of the thesis in the Michigan Agricultural Experiment Station Quarterly Bulletin, Vol. 22, No. 4, May 1940, pp. 247-254. At the outset of this investigation, it was believed that the relationship between the hydrosols and plant growth might be largely a matter of plant nutrient relationships, to the same extent as it is in agricultural soils and possibly more so since some other factors, particularly moisture con- tent, do not fluctuate as widely in hydrosols as they do in agricultural soils. Hence the nutrient content of hydrosols might assume proportionately greater importance. The purpose was, then, to determine the nature and extent of the nutrient supply and the way in which.differences in nutrient content are reflected by the nature and luxuriance of the plant asso— ciations. Another aspect of the nutrient question investigated involved the quantitative analysis of a group of aquatic plants for phosphorus and a comparison between the phosphorus content of the plant and that of the hydrosol in which.the plant was growing. Phosphorus in the plant material was de- termined by the standard Pemberton method (I) while all soil determinations were made by the Simplex'lethod designed by Dr. Spurway (9) of the Soils Department of Michigan State College. The object of this phase of the work was to learn something of the physiology of the aquatic vegetation and to determine the relationships between the content of a nutrient element in the plants and that of the soil. Phosphorus was selected because deficiencies of this element probably occur more generally in agricultural soils than do those of other elements.' In relation to hydrosols, Pond (6) states that “the primary cause of retarded growth of anchored plants is their inability to secure enough phosphorus and potassium, and possibly other elements". The results of these two phases of the work (the nutrient content of a variety of hydrosols and the phos- phorus content of aquatic plants and associated hydrosols) are reported here, while those phases requiring further and more intensive investigation will be reported later. .Detail- ed field methods will be omitted. Soil samples were collect- ed from various lake bottoms and brought into the laboratory to be tested. Data taken at each collecting station included the depth, pH, temperature, and turbidity of the water; a list of the vegetation; and evidences of the degree of wave action. The lake-bottom.soils have been divided into five broad types for the purpose of grouping the laboratory data. A brief description of the types used follows: 1. Sand. Soils included under this type are those whose outstanding characteristics are due to the sand content. Uh- less otherwise indicated a clean, compact, light-colored sand ‘will be implied when the term is used. Sand may be found in conjunction with.marl, peat, shells, or very finely divided particles of organic matter. It should be realized that in- numerable combinations of the various components may and do occur in nature. As mentioned above, however, as long as the characteristics due to the sand are dominant, the soils will be included under this heading--the modifications when present will be given in a column provided for that purpose. 2. Clay. This term is used to designate these soils which are composed of extremely finely divided particles of inorganic material. As such, clay is generally found as a bluish-gray, sticky, and rather homogeneous deposit. 3. Slime. Watersoils included under this heading are characterized by brown to black color, a semi-suspended con- dition of the components, and offering only a limited resist- ance to the passage of heavier objects such as sounding leads, anchors, etc. Here again, there are many possible intergrades between slime and the firmer deposits included under peat. Inclusions of all sorts may occur in slime. Some of the more common ones are marl, shells, sand, and clay. All degrees of decomposition of the plant fragments found in the slime also occur, giving a rather variable texture. 4. Peat. Included under this type are those soils which, like slime, are composed of plant remains. Unlike slime, how- ever, peat deposits are more firm and when secured with sampling devices, relatively firm.material is received. This soil varies with the nature of the parent plant material, environmental con- ditions under which decomposition has occurred, and the degree of decomposition. Three distinct divisions are recognized: (a) fibrous--including all those deposits whose composition is largely fragmented remains of the rooted aquatics, marsh plants, and bog plants such as mosses, cattails, and leather leaf, (b) woody--composed chiefly of woody material, and (c) gelatinous or pulpy--originating from semi-microscopic forms and from easily disintegrated rooted aquatics and characterized by a uniform fine composition and a rather firm but gelatinous con- sistency. Inclusions may involve marl, shells, clay, etc. 5. Earl. This type includes those watersoils whose dom- inating characteristics are due to the marl content. Inclusions or modifications are listed in a column describing the particular deposit under consideration. The series of tables (1 to 5 inclusive) gives the results of the rapid chemical tests made by the Simplex method, which was designed to give a rapid test for available plant nutrients in soils. Tests for the following nutrients are possible with this test: nitrates, nitrites, ammonia, aluminum, magnesium, manganese, phosphorus, potassium, calcium, iron sulphates, and chlorides. The tables presented give the amounts of some of the more important nutrients in the five broad bottom types. The nitrogen tests are omitted because when the sample is re- moved from the lake and allowed to remain in the laboratory, the state in which the nitrogen is present is undoubtedly changed and does not simulate its condition in the lake bottom. Hardness of the water is approximate and is express- ed as grains per U. S. gallon. An analysis of the data presented in these tables shows that lake soils vary widely in plant nutrient supply. It is significant that the extremely acid peat and the pure, highly alkaline marl bottoms are almost completely de- ficient in available plant nutrients, particularly phosphorus and potassium. Observations show that lakes having a propor- tionately large percentage of these types of bottoms are low Table 1. Available Nutrients in Sand. Soil (Air-dry) Sample Modifications Water Rooted No. or Inclusions Hard. , Plants* H P I Ca 1g F8 p‘ ppm ppm ppm ppm ppm 2 much dark organic matter 14 7.5 ... ... 100 ... tr. 6 - b 7 abundance of shells 13 9.0 tr. 2 200 7 tr. 5 - a 15 none 13 7.5 tr. ... 100 6 ... l - a 27 none 12 708 .00 so. 200 6 10 2 - b 44 110116 ll 7 .5 o o 0 tr. 200 8 o s o 2 - b 4:5 none 1]. 6.3 0.75 so. 000 5 so. 3 "' b 60 much organic matter 8.0 ... ... 150 24 ... l - a 61 varying from dark gray 7.0 tr. ... 125 8 ... l - a to black organic matter 63 §:::::el’able organic 8.0 175 16 3 - a 64 dark color 12 7.5 0.5 ... 175 8 ... 5 - b 65 brown color 12 7.2 0.5 ... ... 4 ... 10 - b 67 arl mixed in 12 7.0 0.5 ... ... l ... - b 68 Elack, compact 8 5.5 tr. 2O ... 6 ... - a 70 none 8 5.5 ... ... ... 4 ... ... 71 none 13 7.5 ... ... ... l ... 4 - b '72 none ' 11 5.0 tr. 100 '7 3 - a Range: 8 5.0 0 O 0 0 0 to to to to to to to 14 9.0 0.75 20 200 24 10 *The number indicates the number of plant Species growing on the corresponding soil while the letters indicate the luxuriance of growth: a, very luxuriant growth, b, intermediate, and c, poorly developed. This has reference to thrifti- ness or luxuriance of growth-~not density of stand. In the data, blanks indicate that no test was made, while the symbol (...) means lack of response to test. The foregoing procedures are carried out in tables 1 to 5, inclusive. Table 2. Available nutrients in clay. Soil (Air-dry) Sample Modifications Water ooted No. or Inclusions Hard p K Ca Mg Fe Plants PH ppm ppm ppm ppm 19le 5 none 14 , 8.0 tr. tr. 200 7 tr. 2 - b 9 none 13 8.2 ... ... 200 35 ... 1 - b 10 none 13 9.0 tr. ... 200 40 ... ... 13 none 13 7.5 tr. ... 200 24 tr. 1 - a 16 none 13 7.5 ' ... ... 200 10 ... ... Range: 13 7.5 o o o 7 'o to to to to to to to 14 9.0 tr. tr. 200 40 tr. Table 3. Available nutrients in slime. Sample Modifications Water Soil (Air—dry) . 1Rooted No. or Inclusions Hard P K‘ Ca M3 Fe Plants pH PPm ppm ppm PPm PPm none 14 8.0 tr. ... 130 ... ... 5 - b 6 none 14 7.5 tr. 2.0 175 6 tr. 5 - b 11 clay admixed 13 8.5 tr. ... 200 40 tr. 3 - a 12 marl and shells 13 8.5 tr. ... 200 24 ... 1 - a 14 none 13 7.0 tr. ... 125 10 tr. 1 - a 17 marl at depth of 2 ft. 13 7.5 ... ... 200 15 ... 4 - b 25 marl at depth of 4 ft. 7.8 tr. 5.0 200 10 tr. 6 - a 48 Green ”1°13 Shells 11 7.8 150 a 3 - b included 54 green color, very fluid 5 5.5 ... ... ... ... 2 - c 102 marl and shells 14 8.0 1.5 ... 200 7 ... 9 - a Range: 5 5.5 0 0 O 0 0 to to to to to to to 14 8.5 1.5 5 200 40 tr. Table 4. Available nutrients in peat. lO Soil (Air-dry) Sample Modifications Water No. or Inclusions Hard. pH P K Ca fig Fe Rooted ppm ppm ppm Ppm ppm Plants 11.5 gelatinous, green 13 7.0 tr. ... 125 3 ... 3 - a 20 fibrous, prolific Chara grthh’ PH Of water-8.5 13 5.5 0.5 ooo 125 14 ooo 5 - b 23 gelatinous, containing many small Shells 8o0 1.5 ooo 200 8 ooo 3 - a 41 fibrous, containing many small shells 18 7.6 ... ... 200 24 tr. 7 - a 47 gelatinous 11 7.0 0.5 ... 125 16 ... 4 - a 50 gelatinous 11 6.0 0.5 ... 125 ... 3 - b 52 gelatinOus 5 5.0 ooo ooo ooo ooo 3 " C 55 fibrous 5 5.0 oo- 0.. on. 8 .0. 2 - b 66 fibrous 12 6.5 tr. ... 150 20 ... 2 - b 69 fibrous 8 6.5 0.75 ... 150 10 ... 5 - a 100 fibrous, marl under shore but not under bottom 16 . 8.0 ... ... 200 24 ... 5 - b 101 gelatinous 16 7.7 ... ... 125 7 ... - 107 fibrous 8 5.5 ooo 2.5 1.25 14 ooo 7 " a. 130 fibrous, containing marl 8.5 ... ... 200 7 ... ... 5 5.0 0 0 0 3 0 Range: to to to to to to to 18 8.5 1.5 2.5 200 24 tr. Available nutrients in marl ll Table 5. Soil (Air-dry) Sample Mbdifications Watei Rooted No. or Inclusions Hard. PH P K Ca Mg Fe Plants ppm. ppm PPm ppm ppm 3 impure 14 9.0 tr. ooo 160 ooo ooo 2 "’ b 4 composed chiefly of very small shells l4 8'0 °“ "' 200 "° "' 2 _ b 8 none 13 8.5 ... ... 200 7 ... l - b 18 110118 13 8o5 ooo ooo 200 8 ooo ooo 19 impure, much organic 9 matter 15 8o0 ooo ooo MOO 24 ooo 2 " b 21 none 8.0 ... ... 200 16 ... 4 - b 22 considerable organic [flatter 708 ... ... 200 6 0.. 6 " a 24 numerous plant fibers 8.0 ... tr. 200 8 ... ... 26 none 8.0 ... ... 200 8 ... 2 - b 28 none 12 8.2 ... ... 200 4 ... 3 - c 4:0 impure 18 7.8 ... ... 200 24 ooo 4: - a 43 gelatinous peat admixed 9 8.5 ... ... 200 8 ... 5 - a 46 iYilpure 11 8.5 ooo ooo 200 8 ooo 4: - a 49 none 11 8.2 ... ... 200 8 ... 3 - c 51 none 1]. 8.5 ooo ooo 200 8 ooo ooo 9 7.8 0 0 160 0 Range: to to to to to to 0 18 9.0 tr. tr. 200 24 12 in productivity. not only with respect to vegetation but also with respect to plankton and fish production. Similar observa- tions are recorded by Raymond (7) and Welch (11). Among the other types of bottoms, there seems to be no correlation be- tween nutrient supply and the type and luxuriance of the plant communities. Certain plants prefer given types of soils. but the preference seems to be based on physical characteristics, such as firmness of the bottom and subjection to wave action, rather than on the chemical constituency of the bottom. Table 6 presents the results of the analysis of several different species of aquatic plants--giving the phosphorus con- tent as determined by the standard Pemberton method. The soils upon which the plants were found have been tested for phosphorus by the Simplex method and those data are also included in the table. The reason for selecting phosphorus has been previously mentioned. ‘ An examination of Table 6 indicates that there is a wide variation in the phosphorus content of aquatic plants. The per- centages of P205 range from 0.189 to 1.105. Different species of plants growing on the same soil absorb different amounts of phosphorus. In one case gotamogeton mm 11folius,uénacharis. Eyriophyllum. and Eggggrwere found grow- ing together and contained the following percentages of P205 respectively: 0.400. 0.479. 0.816. and 0.727. A single species of plant does not show a uniform phos- phorus content but varies widely when grown in different lake soils, although the phosphorus content of the plant shows no Table 6. that of associated soils. Comparison of phosphorus content of aquatic plants and 13 Phos- P205 . phorus i n Lake 8011 in soil plants Plant(s) Sample (ppm) (percent) N°° Pine L. . Barry Co. fibrous peat 0.0 .508 Brasenia P - 1 Pine L. Barry Co. fibrous peat 0.0 .351 Naias P - 2 Craig L. , , Potamogeton Branch Co. firm fibrous peat 0.5 .256 pectinatus P . 3 Craig L. fibrous peat Branch Co. no sample taken '227 Chara P ' 4 Fish L. green gelatinous , Potamogeton Oakland CO. peat 0'0 '534 amplifolius P ' 5 Fish L. slime too thin to .359 Brasenia P - 6 Oakland Co. get with sampler SQuare L. gray soft gelatinous 0 0 453 Potamogeton P 7 Oakland 00. peat and marl ° ' spp., Chara ' Pleasant L. green organic slime Oakland 00. and shells 0.0 .441 Potamogeton sp. P - 8 Pleasant L. Oakland 00. sand 0.75 .283 Potamogeton sp. P - 9 Pond on U.S. 10, ,. . 3 mi. use-J. of sand and clay 0.0 .762 MYI‘lOPhyllum: p .. 10 Clarkston Chara & algae Mona L. thin layer of fib- Lemna, Muskegon Co. rous peat over sand tr. 1'098 Spirodela P ' ll Pigeon L. semi-fluid fibrous F Ottawa Co. peat 0.75 .757 Chara P - 12 Cook L. . Livingston Co. brown fibrous peat 0.0 .833 Heteranthera P - 13 Bennett L. thin layer of black Livingston Co. slime over sand 1'5 '296 Chara P ' l4 Kawbawgam.L. rown gelatinous q . Marquette Co. peat 0.0 .363 Pontederia 410-A Kawbawgajn L. brown gelatinous 0.0 .496 Potasnogeton 410,4? Marquette Co. peat praelongus l4 Table 6. Continued. 51513;}. 1:235 Lake 5011 1; soil plants Plant(s) saggfe (Ppm) { percent ) ‘Ezizugtte Co. sand 0-0 .309 Isoetes 413-3 3::Zu2tte sand 0 - O . 553 SPaI‘ganium 413-A am 333.32%... mm 3:22;: Egfidford L. Ziifitifiius peat 0.5 .160 Chara 431-0 3:2:22 2:?dford L' ggigtiggus peat 0.5 .263 Potamogeton sp. 431-P 3:22;: 533de 1” Ziiitifiius peat 0-5 -521 Nuphar 43 -w Man 5W ‘fi2323eite Co. sand 0'0 '219 Scirpus SP. 453-8 fizzfgmggnLéo. :élgiifigis peat '304 Vallisneria 439'C EggggmggnLCo. gilgiiigis peat ’534 Zizania 439'R Eiiqi:§:: 36. dark sand 0.0 .262 Nuphar 445-L 8mm fi:::i:t::rgg.L. sand 0-0 ~491 ::;:?:§::°n 446-P fi:::i:t::rg:,L. sand 0.0 .791 Polygonum 446-8 81:22: £5. 2:23.332; 60 .400 23:32:33: 468% g::::: 36. Sifidgafiifiig 5.0 .479 Anacharis 468-3 Otsego L. sand, marl, 5.0 .816 Myriophyllum 468-M Otsego Co. and sawdust 15 Table 6. Continued. Phos- P205 . phorus i n n Sample Lake 8011 in 8011 plants Plant(o) H‘O. (ppm) (percent) Otsego L. sand, marl 9 m . ‘ Otsego Co. and sawdust 5’0 '7”? naias 468-N Buttonbush Pond clay and St. Joseph Co. organic matter tr. '526 Polygonum 504-A Buttonbush Pond clay and . . St. Joseph Co. organic matter tr. .760 Utricularia 504-3 Buttonbush Pond clay and m St. Joseph Co. organic matter tr. '788 iolypella 504'F Buttonbush Pond clay and t 004 Potamogeton St. Joseph Co. organic matter r. '“ natans 504‘“ Buttonbush Pond clay and "”0 Potamoaeton 04 St. Joseph Co. organic matter tr. ‘/0 obtusifolius 5 "P Buttonbush Pond clay and a ,L . St. Joseph Co. organic matter tr. 1'105 oagittaria 504-8 Prairie River L. 'elatinous eat st. Josenh go find marl p 0.0 .656 Ceratophyllum 507-0 Prairie River L. gelatinous peat . St. Joseph Co. and marl 0'0 '659 Peltandra 507-P Silver L. sand, marl, and washtenaw Co. organic matter tr. '189 Chara 512-0 16 relation to that of the lake soil. Potamogeton natagg, for instance, contained 0.281 per cent P205 when growing on a soil containing 3.0 p.p.m. While another sample from a different lake contained 0.904 per cent P205 when growing on a soil containing only a trace of phosphorus. If aquatic plants obtain nutrients from the soil, they must have greater extracting powers than is generally supposed since the Simple: method is able to detect available phosphorus in concentrations of less than 0.5 parts per million, and in many cases plants showed a relatively high percentage of phosphorus when the cor- responding soil failed to show even a trace by this test. How- ever, if the nutrients were absorbed directly from the lake water, then a very minute amount of phosphorus in the lake water might supply the amount required by the plant for its metabolism. It is interesting, in this connection, to note that two floating plants, Lgmgaand Spirodela, contained a higher per- centage of phosphorus than all of the rooted plants, except for a sample of Sagittaria from.Buttonbush Pond in St. Joseph County. 17 FERTILIZER PLOTS AT ROSE LAKE Following some preliminary pot tests in the greenhouse, which were made to determine the feasibility of the use of commercial fertilizers, it was considered desirable to make some fertilizer applications on a larger scale and under more natural conditions. Through the cooperation of the Game Divi- sion of the Michigan Conservation Department, experimental plots were constructed in the floating bog surrounding Rose Lake in Bath Township, Clinton County, Michigan. The mat of vegetation here is about 18 inches thick and is underlain by varying depths of semi-fluid fibrous material of the same nature as that comprising the bottom of Rose Lake. Excava- tions were made during the winter, when the ground was frozen, to a depth of three feet. Wood cribs were placed in the pools to prevent the soft material from filling them in. Plots of two different sizes were made-~three of 15 feet square and nine which were ten feet square. The plots were placed 25 'feet apart. Before adding fertilizers to the plots, it was necessary to know the rate and amount of diffusion of salts which would take place in order to be certain that the application of ferti- lizer in one pool would not affect the adjoining pools. An experiment was established in the bog to determine this. A series of holes three inches in diameter were bored through the surface mat at the following intervals from the hole in which the salt was to be placed: 1, 3, 5, 10, 15, and 25 feet. 18 The salt (1500 grams of K31) was put in the end hole after a sample of the water and soil had been tested for potassium and found to be very low-~less than five parts per million. Tests were frequently made to determine the amount of diffusion. After two weeks, a slight increase occurred in the first hole-- one foot distant, while the others remained unchanged. Through- out the period of a year, tests have indicated that the salt did not diffuse to an appreciable extent and that applications of soluble salts in one plot would not affect the nutrient content of adjacent plots. The original nutrient content of the soil was: phosphorus-- 0.0 p.p.m.: potassiump-less that 5 p.p.m.; and nitrates--0.0 p.p.m. Equivalent amounts of vegetation were introduced into each pool. Plants used were two species of Potamogeton_and Utricularia vulgaris. The following treatments were made in duplicate, using two plots as controls: 500 pounds per acre metaphosphate; 2000 pounds per acre metaphosphate: 500 pounds per acre muriate of potash: 500 pounds per acre ammonium nitrate: and 500 pounds per acre metaphosphate, plus 1500 pounds per acre muriate of potash. These applications were made in May, 1939, and the experiments are still in progress. The plants were not harvested, but some data were ob- tained regarding their growth. Figures 1 to 4 inclusive show differences in growth in various pools. Within a week, following the fertilizer applications, algal growth.was enormous in the plots receiving a phosphorus 19 Fig. 1. One of the control plots. Water lilies have come in naturally. Plot receiving muriate of potash. vegetation is not as abundant as in the control plot. Fig. 3. A plot receiving 2000 pounds per acre of metaphosphate. Utricularia is very prolific. Duckweeds naturally intro- duced. +£‘ 1. _ I ' '. ' ' O. I fl- -1.” “t. I .‘ ‘ ..., ff. .1, “‘ 20r:1f¢1; ,‘vh. J‘;t- op . - "7.“ (WA “V?“ V / w _ '-’i . ‘, . " V (1': o. : - ' _ \ r‘ .I o :. ,_ .' 4 ' ' ‘1‘, . . A plot receiving both metaphosphate and muriate of potash. Vegetation about as abundant as in Fig. 3. 21 treatment. This condition persisted for two weeks, after which the algae were present, but not in such large quantities. The Uticularia in_these same pools grew very rapidly and at the end of the second season is still prolific--see Fig. 4. Duck- weeds and water lilies have been naturally introduced. Other treatments seemed to have little effect, since a noticeable difference in plant growth could not be detected. These results indicate that where definite nutrient de- ficiencies exist, the condition can be remedied by the addition of commercial fertilizers. However, in lakes the problem is complicated considerably. First, it is difficult, in a lake, to attribute the complete absence or even scarcity of vegetation solely to nutrient deficiencies because of the presence of num- erous other factors--probably chiefly physical in nature. The method of application would undoubtedly require modification. Diffusion of the nutrients into the water would occur more rapidly in a lake where water movement occurs. Experiments involving these problems should prove both interesting and valuable to the fisheries biologist. 22 GREEHHOUSE FERTILIZER TESTS. During the winter and spring of 1939-1940, a series of fertilizer tests were made in the greenhouse. The soils used in the tests were collected from the bottom of lakes throughout the state and were taken from areas which seemingly should be producing vegetation but were not. The soils were tested for plant nutrient content and the treatments made on the basis of the original fertility. The soil samples were placed in 2-gallon Jars, filling the Jar to a depth of three inches. The jars were then filled with distilled water. The plant used in this study was the common water-weed (Agacharis canadengig). This species was used because it grows rather rapidly under favorable conditions, and it develops roots readily so that anchorage is not a problem. The fresh weight of the plants was measured for each Jar. The plants were allowed to grow for three days, at the end of which time the few that had died were replaced. The fertilizers were then added. Dur- ing the growing period of ten weeks, the Jars were kept full by adding distilled water. Algal growths were treated with copper sulfate, but in many cases a treatment with sufficient strength to kill the algae damaged the plants to such an extent that they did not recover. At the end of 10 weeks, the plants were harvested and weighed. Table 7 presents the results from the experiment. It is interesting to note that some applications produced enormous 2:5“ increases in growth over the control. Others seemed to produce conditions very favorable to algal growth, with the result that the Anacharis_was either choked out or killed by copper sulfate treatments. In one case--soil 479-othe treatment seemed to in- hibit algal growth. From the practical standpoint, it may be important to point out that the majority of the large increases were ob- tained by using natural materials as fertilizers. The addi- tion of an acid muck to highly alkaline marl bottoms lowers the reaction and probably makes available some nutrients which are unavailable in a strongly alkaline condition. Additions of marl to acid soils seems to give the same response in plant growth by reversing the conditions of reaction and availability of plant nutrients. Earl and muck or peat are plentiful in the state, and if the productivity of lakes can be increased by artificial means, it seems that these two sources of fertiliz- ing material should prove valuable. It must be pointed out, however, that in some cases actual nutrient deficiencies may occur, and that in these cases, additions of commercial fertil- izers may prove to be the solution. .msmae an pso assess mpeeam**** .maonfigaom .Ho soapwoflamw poems execs swamp mass was pmop mafia: .opwafln Samsoganlz .mmepoa mo magnesium .opwnmmongoagunm .HnsEauda .Mosennz you 3356*... 3.33 3003qu £39 magnahanmh .psoa nagging ..naweiz Jessamine .csemunm ”memofimoap "flow .Hom maonaenm ... o.m Harcooa i ace 2.600 III: we gloom .muoom . m.e Honseoo 0.0 0.0 m.m mm .muumae . a .m .em cos n m .stoooa .1 m.e Hospeoo 0.0 0.0 m.e mavuaee m.s swoon .I. new Mloom In. m.m Hospeoo o.o o.H e.s 2 .muumom .Il o.w Hawoooa I'll: a.m Hospeoo o.e o.o e.e mnumee o.m swoon o.m zwoom o.m mnoom .IIIII. o.m auoom 1 o.m Hospeoo 0.0 0.0 m.a 2H.m--aam on. M .m .z .8 com m.m Hospeoo 0.0 0.0 a.a _sw-mam m.s swuoooa m.s Hospeoo .sp o.o m.a swuaae :1 o.m groom .msoom In. o.m Hospeoo m.s m.o n.s mm .muuame m.a HeToooa IIIIIII e.m Honpeoo o.o .sp s.m mauuomm a; Muoom .Zuoom . m.m Hospeoo o.o m.o m.s mm .awneom I 0.“. Eunoooa m.a Hospeoo o.e n.o m.s e .mnnoom moon meow .osa mead o .ome aooa ***mn msoa\.mpH sun sea Haom *Haom Ampnoavsonp opoowaasu ho .mp5 £9393 ......psofioohe M A no mg .389... tsetse assets Mm Essa .1. Sea 25 .A COMPREHENSIVE STUDY OF FOUR LAKES During the summer and fall of 1940, a detailed study was made on four lakes in the northern part of the Lower Peninsula onKichigan. The objective here was to study the natural distribution of vegetation in different lakes, and to determine the importance of water soils in the ecology of the larger aquatic plants. Lakes were chosen from this part of the state for two reasons: first, the Hunt Creek Experiment Station of the Institute for Fisheries Research, located at Lewiston, made an excellent headquarters since all of the lake conditions desired could be found within a radius of 10 or 12 miles: second, the problems of private ownership of surrounding lands and that of conflict with sportsmen and resort owners are less intense in the northern part of the state. Methods . The methods used in this investigation are essentially the same as those used for similar studies by'Wilson (12) on Sweeney Lake, Wisconsin, and Rickett (8) on Green Lake, Wis- consin. Transects were arbitrarily chosen on the basis of natural shoreline zones or types, assuming that opposite a uniform/shoreline, there is a corresponding uniform zone of off-shore vegetation. These transects extend from the shore through the zone of vegetation. In the lakes studied, the vegetation never extended to a depth exceeding six meters. 26 Plant and soil samples were collected along the tran- sects with the Peterson dredge, which gives perhaps a better quantitative sample than do other available sampling devices. Samples were taken at depth intervals varying from one-fourth to one meter, depending on the degree of slope of the bottom-- the steeper the slope, the larger the interval. The entire sample, plants and soil, was discharged into a large pan. The plants were transferred to a wire bas- ket hanging over the side of the boat, washed, packeted in wax paper, and preperly labeled. .A sample of the soil was placed in glass jars provided for that purpose, and each jar labeled. These samples were taken into the laboratory for further study and treatment. See Fig. 5 for field equipment. In the laboratory, each plant sample was separated into species: each species sample was washed free from de- bris, drained and blotted to remove the excess water, and then carefully weighed. An index card was filled out for each species, recording the transect, collecting station within the transect, depth, and fresh.weight. Dry weights were not measured, chiefly due to lack of time and space. Ioreover, since the larger aquatic plants, with a few ex- ceptions, do not vary over eight or ten percent in water content, it seems that taking both fresh and dry weights is impracticable. If, for any lake, the total weed crOp is desired on a dry weight basis, one needs only to multiply the fresh weight figures by 12 or 13% since the plants average 87% or 88% water. Using this technique on the data from the Green Lake, Wisconsin, study,'the result is 27 probably not different enough to warrant the additional labor. In this lake, the total crop is given as 13,002,500 kg. fresh weight and 1,527,900 kg. dry weight. Twelve percent of 13,002,500 is 1,560,300. If considering only the emergent plants, the factor used would have to be increased because emergent plants contain a lower percentage of water. The soil samples were tested for reaction: and avail- able phosphorus, potassium, calcium, and iron. These tests were made with the Simplex Soil-testing System designed by Dr. C. H. Spurway (9). Briefly, this consists of making a .135 D. acid soil extraction from each sample, dividing the extracted solution into 1 cc. portions, and using colori- metric tests for each nutrient. The acid extraction is used as an arbitrary means of determining the readily soluble salts in the soil and it is believed that these salts are available to the plant. The results of the analyses were recorded an index cards, together with the information re- garding the collecting station and a description of the soil. A small sample of each soil was oven-dried at 110° C. for organic matter determinations. The hydrogen peroxide I method was used to determine the organic matter. Tabulation and treatment gfrggggz The discussion will be restricted to the methods in- volved in tabulating the data rather than a presentation of the results. In all computations, except that of the frequency of occurrence and relative abundance of various plant species 28 on different soil types, the method used was the same. The transects were divided into zones on the basis of depth of water. Zone 1 includes all readings between the shore and a depth of one meter (0 - l m.); zone 2, those between one and three meters (1 - 3 m.): and zone 3, from three meters through six meters (3 - 6 m.). Since the number of samples varied, the measurements from each zone were averaged. Plant weights, then,are ob- tained as the mean weight per sample. These means were converted into grams per square meter, kilograms per hectare, and, to give the maps (Figs. 8b, 12b, 16b, and 21b) more popular usage, into pounds per acre. Total yields were computed by multiplying grams per square meter by the area (in square meters), which was ob- tained from.a large base contour map. The average yield by zones (Figs. 9, 13, 17, 22, and 24) was determined on the basis of the total yield and the total area of the zone. Fertility and organic matter determinations of soils were averaged by zones; the mean reading was used in all calculations. In Tables 10, 13, 15, and 19, showing the frequency of occurrence and relative abundance of plant species on different soil types, the individual plant and soil samples were considered. - The derivation of the fertility figure or nutrient content warrants explanation. On the basis of some 3,200 29 tests, it was found that each of the nutrients occurred in quantities within a given range--this range varying with each.e1ement. In deriving a standard fertility figure, each element received equal weight or importance. Using a factor, based on the highest reading for that element, the nutrient could be properly proportioned. The following table illustrates: Standard laximum. Equiv. (ppm) Equiv. Nutrient Value (ppm) Factor Value Reading Value Phosphorus 2.5 8 20 1.5 12.0 Potassium. 20.0 1 20 6.0 6.0 Calcium 200 * 1/10 20 75.0 7 .5 Iron 40.0 t 20 15.0 7.5 Maximum.nutrient content -- 80 E.C. of sample 33.0 *In marl samples the calcium.content is greater than 200 ppm. Exact readings would require dilution of the soil extract--a process requiring more time than was available. Moreover, it is doubtful that calcium.in excess of 200 ppm. adds to the fertility of the bottom. The value of the figures obtained by this method and other possible methods will be discussed later. In tables dealing with plants, some species which occur in the list of plants for a given lake are omitted. In such cases there were not enough data to warrant the inclusion of those species which are omitted. When a species was observed but not taken in a sample, it was recorded as a trace (tr.). Fig. 5. Stern of boat with field equip- ment in place. 30 31 The outlines and contours of the maps in this manu- script were obtained from maps furnished by the Institute for Fisheries Research. Data Presented. The data will be presented for each lake individually, with a general discussion following. LakgpFifteen: ‘Situated in sections l4, l5, and 22 of Briley Township, Montmorency County. This lake occupies an 88 acre basin in the Thunder Bay River valley, the river flowing through the western part of the lake. The basin lies in a rather narrow strip of outwash plain situated between two moraines of the Port Huron system. The lake margin is swampy around practically the entire circumference (Fig. 8a). The swamp or bog is underlain by marl at a depth varying from three to fifteen feet. The lake is still a marl-producing lake--pebble marl being unusually abundant. The lake basin is characterized by a narrow shoal, which is terminated abruptly by a rapid ”drop-off". See Figure 8a for further hydrographic information. The water in Lake Fifteen is clear and relatively hard; it measures 18 grams calcium carbonate per U. S. gallon. Twenty-one species of plants (Table 8) are repre- sented in the lake. Table 9 shows the distribution of the flora with re- gard to depth. Zone 1 (the area between the shore and the 32 one-meter contour) contains 36.4% of the total vegetation. The plants comprising the bulk of the flora in Zone 1, in order of their abundance, are ghgggigp,, Nuphar_advena, and gygiophyllum heter0phyllum. In Zone 2 (the area between the l and 3-meter con- tours) g9 heterophyllum,.Chara_§p,,,Potamogeton amplifoliug, g. gggteriformis, and smaller amounts of other species com- prise 46.6% of the total crop. The vegetation in Zone 3 (the area between the 3 and 6-meter contours) accounts for 17.0% of the total crop; the chief contributing species, in order of their abundance, are E. heterophyllum, g. zosteriformig, and ghaga‘gp. The distribution of the flora with respect to bottom types is shown in Table 10. The relationship between yield of plants, the organic matter content of the bottom, and the nutrient content of the bottom is shown in Figure 9. The density of the vegetation and the bottom types occurring in the lake are given in Figures 8b and 8c res- pectively. 33 Fig. 6. View showing the uniformity of the east shore of Lake Fifteen. Fig. 7. A protected bay at Transacts A.& B. Picture taken in May. In July and August, vegetation here is abundant. LAKE FIFTEEN 1,100 ft. 30816: Baa!!! 7* 4 Swamp J Thunder Bay R. / I I /,‘ a / r/A\“ \ \ / /— /'\ \ \ / // \ Grass marsh Thunder Bay R‘ ,4 Spruce, Poplar . ~, G and tamrack , ' HT \‘ ‘\ )’ < \ (\W , .. \ ,Q> / \\ \\\| ‘ .‘\\\ I W ‘. \—————o IA / 1’ ‘ \’ / \ / /‘ 3 \ 9 / -\_———\“_/ / II/ A ' - x / (:4 é *:~ _/\ , B C \\“_,;I Spruce Swamp Fig. 88.. Map showing contours (in meters) and transects along which samples were collected. ...~ LEGEND 2000 to “000 EST] Less than 100 r‘7—-—7— LAZTJ m 100 to uoo #000 to 5000 047/153 m Moo to 1000 5000 to 16000 16000 to 32000 *:“rT“ ’37 A xiii/f x 12.2... 1000 t0 9000 Over 32000 Map showing vegetation density in lbs. oer .t‘ \ . a . .. O A ‘0 .V 9.2.9 L e... L 0. fizz... 0 .0 .e""' 0. 0:0. 0" 1.0,. ’. C '0 I I. .0. 0" 0;. ' e I '1 e 'e O I .0 e e e . D O. Fig. 8b. 36 LEGEND PP M D--delta denosits S--sand M--mar1 y PPT—nuloy neat Fig. 80. Map showing bottom types. Table 8. Plants Occurring in Lake Fifteen. Carex lasiocarpa Carex rostrata Ceratophyllum demersum Chara spp. Dulichium arundinaceum Iziyriophyllum heterophyllum Kaias flexilis Nuphar advena Nymphaea odorata Potamogeton amplifolius P. angustifolius P. filiformis var. borealis P. foliosus P. interior P. natans P. pectinatus P. praelongus P. zosteriformis Scirpus acutus Typha latifolia Utricularia vulgaris var. americana Sedge Sedge Coontail Muskgrass; Stonewort Three-way Sedge Water Milfoil Bushy Pondweed Yellow Water Lily White Water Lily Large-leaf Pondweed; Bass Weed Pondweed Pondweed Leafy Pondweed Pondweed Floating-leaf Pondweed Sago Pondweed Whitestem.Pondweed;'Muskie weed Flat-stemmed Pondweed Hardstem.Bulrus Common Cattail Bladderwort 38 Table 9. Total Weight and Percentage of Each Species by Zones. Lake Fifteen Zone 1. Zone 2. Zone 3. Total Species Height % Weight 92'. weight 3% Weight kg. kg. kg. kg. Carex lasiocarpa 529 1.7 529 Chara spp. 17,715 56.4 5,639 14.0 2,748 18.1 26,102 Kyriophyllum.heterophyllum 4,423 14.1 23,601 58.5 ,592 56.7 36,616 Naias flexilis 387 1.2 396 1.0 783 Euphar advena 5,663 18.0 1,175 2.9 6,838 Potamogeton amplifolius 1,638 5.2 4,185 10.3 41 0.3 5,864 P. angustifolius 291 0.9 263 0.7 554 P. interior 111 0.4 190 0.5 301 P. natans 62 0.2 120 0.3 182 P. pectinatus 278 0.9 278 P. praolongus tr. --- 644 1.6 644 P. zostoriformis 210 0.7 3,301 8.2 3,275 21.6 6,786 Scirpus acutus 71 0.2 71 Typha latifolia 26 0.1 26 Utricularia vulgaris 7 0.0 799 2.0 806 Total -------------------- 31,393 40,313 14,656 86,380 Percent of Grand Total in each Zone --------- 36.4 46.6 17.0 39 TableJIL. Frequency of Occurrence and Relative Yield of Certain Plant Species on Different Bottom Types.l Lake Fifteen Bottom Type. Species M PP S,M 'M,Pr PP,D,M S,M,PP Carex lasiocarpa 75* I b 25** Carex rostrata <1 25* 75* _ Ceratophyllum.demersum T 50* 50* Chara spp. —) 25** 1* 5* 9** l** 3*** i 18** 38*** Dulichium arundinaceum T 45* 55* myriophyllum.heterophyllum. 12* 2** 4* 12** 1* 2** 17** . 5o*** Naias flexilis 2* 9* 40* 4* ’ ‘ 45** Nuphar advena - 9* 40* 15* 1* 2* 33** Potamogeton amplifolius iv 2* I 6* 55* 1* L 37** P. angustifolius . 4* 1 [ 72* 1* 8* ' 15** P. interior 45* 33* 22** P. natans 16* 84* P. pectinatus . ‘ 67* ' 33*** P. praelongus . 3* 17* ‘ 80* . 2* 33* 4* P. zosteriformis . 40** 21*** Scirpus acutus 95* 5** ..... 1. .. ..‘d-M H .. Typha latifolia 100* Utricularia vulgaris 4* 55* 18* V 15* 10** 1 Frequency of occurrence on each bottom type stated in percent of total number of collections of given species. Relative yield: *, less than 300 gms./ sq. meter; **, 300 to 1000 gms./ sq. meter; ***, over 1000 gms./ sq. meter. Bottom types: Me-marl; PP--pulpy peat; S--sand; D--detritus. Example: 100% of the collections of Carex lasiocarpa were taken from a sand and marl bottom; 75% of the collections measured less than 300 gms. per square meter, 25% measured between 300 and 1000 gms./sq. meter. Fifi. 9. Graphs showing aVerage yield of plants (kg./ hectare-—————), percentage of organic mutt.r in bottom (.~_.__), and Intrien. content of bottom (___._) by zones in Lake Fifteen. r"O O ‘1 . 21' .5 / {3:500 ...-—-—-———- ’- 14000 123000 ,/ Zones 1 2 3 41 *IgéEDTwentyfpwg: Situated in section 22 of Vienna Town- ship, MOntmorency County. This lake is a good example of a "kettle" type, occupy- ing a basin of 109 acres in the Port Huron moraine. Its shores rise rather abruptly so that only a narrow wave-built terrace is found. Exception to the above occurs along the south side for a short distance and in the bay constituting the western end of the lake. The filling in of this large bay was undoubtedly hastened, or perhaps may be accounted for, by the existence of a large sawmill there in the lumbering days. The shoreline is characterized by fallen trees and an accumulation of logs, sticks, and all manner of debris. A list of the plants is given in Table 11. Table 12 shows the distribution of the flora with re- gard to depth. In Zone 1, the most abundant plants are, in order: 22飧,3 ., Nymphaea odorata, and Potamogeton EflPll' folius. In Zone 2, they are: Chap; gp., P. obtusifolius, and g. amplifolius, and in Zone 3, _I_’_. amplifoliusand g. obtusifolius comprise the total crop. Zone 1 produces 16.0% of the total crop; Zone 2 pro- duces 42.2%; and Zone 3 produces 41.8% of the total. Figures 12a, 12b, and 120 give the maps of the lake corresponding to those of Lake Fifteen. *The land surrounding Lake Twenty-two is owned by the Kneeland-Bigelow Lumber Company. Their permission for access to the lake is greatly appreciated. The relationship of plants to different bottom types is given in Table 13. Figure 13 shows the graphs or curves for yield, organic matter content of the bottom, and the nutrient content of the bottom by zones. Fig. 10. A swampy shoreline. 42 A rather steep, wooded shoreline, characteristic of the east and northwest sides of the lake. 43 LAKE TWENTY-TWO 600 Scale: in: Maple Fig. 12a. Lisp showing contours (in meters) and transects along which samples were collected. Fig. 12b. LEGEND [" ‘1. Less than 100 F1411; ' 100 to 400 400 to 1000 m m 1000 to 2000 16000 to 32000 Map showing vegetation density in lbs. oer A. 45 ( ) PF .5, U .1 {9 pp \ i j 5 ( ,3. “’" LEGEND 6 <5) D—~woody detritus /Q d--herbaeeous detritus ’0 S--send G-«gravel B~—bou1ders C--Chara marl PP—-eulny oeet Fig.12c. Map showing bottom types. 47 Table LL. Plants Occurring in Lake Twenty-two. Carex Lasiocarpa Chara spp. Dulichium.arundinaceum Eleocharis palustris Equisetum.fluviatile Eriocaulon septangulare MyriOphyllum.heterophyllum Nymphaea odorata Potamogeton amplifolius P. gramineus var. P. g. var. graminifolius f. myriophyllus P. obtusifolius P. pusillus Sagittaria latifolia Scirpus acutus Sparganium.eurycarpum Typha latifolia Sedge iuskgrass; Stonewort Three-way Sedge Creeping Spike Rush Horsetail; Securing Rush Pipewort Water Milfoil White Water Lily Large-leaf Pondweed; Bass Weed Variable Pondweed Variable Pondweed Pondweed Pondweed Duck Potato; Arrowhead Hardstem.Bu1rush Bur Reed Common Cattail I‘l‘flltliilIIIKIll Table 12. Total Weights and Percentage of Each Species by Zones 0' Lake Twenty-two 48 Zone 1. Zone 2. Zone 3. Species Total Weight <7. Weight % Weight 7. Weight kg. kg. kg. kg. Chara spp. 8,844 40.7 40,192 69.8 tr. 49,036 Eleocharis palustris 138 0.6 138 Equisetum fluviatile 316 1.5 316 Eriocaulon septangulare 803 3.7 803 Nymphaea odorata 5,412 24.8 tr. 5,412 Potamogeton amplifolius 4,762 21.9 7,086 12.3 36,074 63.3 47,922 P. gramineus var. gramini- " folius f. myriophyllus 200 0.9 297 0.5 497 P. obtusifolius 753 3.5 8,854 15.4 20,951 36.7 30,558 P. pusillus 1,164 2.0 1,164 Scirpus acutus 104 0.5 104 Sparganium eurycarpum 418 1.9 418 Total -------------------- 21,750 57,593 57,025 136,368 Percent of Grand Total in each zone ------ 16.0 42.2 41.8 Table 13. Frequency of Occurrence and Relative Yield of Certain 49 Plant Species on Different Bottom Types.l Lake Twenty-two. Bottom Type Species S S,D PP PP,C S,G Carex lasiocarpa 65* 35* Chara spp. 2* 6** 12** 80*** Eleocharis palustris 87* 13* Equisetum fluviatile 80* 15* 5* Eriocaulon septangulare 66* 4* 30** Hymphaea odorata 72* 18** Potamogeton amplifolius 2* 6* 5** 12*** 75*** P. gramineus var. gramin- 2* 7* 87* ifolius f. myriophyllus 4** P. obtusifolius 33* 57** P. pusillus 65* 35** Sagittaria latifolia 25* 75* Scirpus acutus 45* 24* 3* 22** 6** Sparganium.eurycarpum 85* 10** Typha latifolia 90* 10* l marl. For explanation of table, see Table 4. Bottom types: S—-sand; D--detritus; G--grave1; PP--pulpy peat; C--chara ‘ 13.1 . Graphs snowing QVQPnge yield (hf./ hcct, percentage organic matter (————-—), and nutrient contrnt (—-——-—7 by zones in lake Twenty-two. / v30 / / / // ... 10 70 I]: /'/ / “JO 5) / ,/ 50 In / / I, / (0 ” /'/ t I“ / 5000 /// 4000 , 3000 2000 H (.0110 S l to (A 51 Litt1e_Wolf Lake: Located in sections 34 and 35 of Albert Township, Montmorency County, and sections 2 and 3 of Greenwood Township, Oscoda County. This lake is unique in that it is composed of two basins, separated only by a narrow channel in which the water does not exceed a depth of 18 inches. The two basins differ widely, in that one is deep and contains little vegetation, while the other is shallower and rich in vegetation. The entire lake is 87 acres. Other information concerning the hydrographic aspect of the lake may be found in Fig. 16a. A rapid lowering of the water level--a phenomenon which has attracted considerable attention in recent years-~19 an in- teresting feature of this lake, as well as of many others in the surrounding, level outwash plain. In Little Wolf Lake, boat-docks have been left high on the shore as the water re- ceded. The outlet, which formerly flowed into Big Wolf Lake and several years ago was capable of accommodating a boat equipped with an out-board motor, no longer functions during any part of the year. The small bay at Transect K3, formerly a favorite bass-fishing spot for the local sportsmen, is now dry and has grown up to marsh grasses. Only two definite types of shorelines are present. A sand beach extends around nearly the entire lake. See Fig. 14. Exception occurs in the vicinity of Transect F, where a gravel pavement has been formed. See Fig. 15. The data for this lake have been tabulated in figures and tables corresponding to those of the two lakes discussed previously. Fig. 14. A wave-swept sand beach as found around nearly the entire lake. Fig. 15. Gravel pavement situated in the vicinity of Transect F. 52 53 .popooaaoo oael.neflas¢u goamh.wcoas anaconda» use . ._\ w n \//./x hunches adv announce madness mam asmH .mfim .--: ,. e ._ . \h, 1 . In .S as .eew \\. .. x .. e\. \ . .4 .oo snoooo .ou monoamauamm: m 1 see .. . .\\ x \n .1 \\ . / .e _ S. a a M ..t / _ . D/é ., e _ N ,. ._ x. .1» a / /, , \ .11 “a” \s / / .4 4. % “OHUOW \\\ // t 1 “x \\ ,, V uni he: Seas LEGEND é Sixéfi i1")! 11.. 2' . O shifi? 2' £82 Y 1000 to 2000 i a. 2000 to uooo O C) rvir'rr" . i: k * ; CU \\ ; .c* 4 : .+>. i t m ‘ . L... m L. o k. -4 >. ,‘Y' 69 e 06x 100 to 400 400 to 1000 4000 to 5000 3000 to 16000 per A. Man showing vegetation density in lbs. Fig. 16b. 54 55 .mmemp Seneca madsomm 062 uses measocnwm Humans: mamcasonnnm am>mamsnw cadmium mzuaap¢CIIo szomq .. 1‘ .fil e wit n./ x 4 be. 0 .W. 0 .0 Q -xx xx» .. n 0 x ./m ...... t /, \\\-\ala M a\ Al‘ .I Q \\ r r t m r. x x S .omH N‘ «K A") .uwm Table 14. Plants Occurring in Little Wolf Lake. Chara spp. Hyriophyllum.heterophyllum Naias flexilis Nuphar advena Nymphaea odorata Potamogeton gramineus var. P. praelongus P. natans Scirpus acutus TableIUS. Frequency of Occurrence and Relative Yield of Certain Muskgrass; Stonewort WaterIMilfoil Bushy Pondweed Yellow Water Lily White Water Lily variable Pondweed Whitestem Pondweed Floating-leaf Pondweed Hardstem.Bu1rush Plants on Different Bottom Typesl. Little WOlf Lake. 56 Bottom Type Species S,M S,G S,G,Mf S,B,G S,PP S,D S,M,D PP,M Chara spp. 16* 4* 16* 2* 2* 4* 11* 4* 33** 8** ."n ' t J r * 65* Lyriophyllum.heterophyllum 12 23** 54* Nymphaea odorata 33** 13*** 4** 6** 5** Potamogeton praelongus 87*** 1 Bottom Types: S--sand, M--marl, G--grave1, B--bou1ders, PP--pu1py peat, D--detritus For explanation of table, see Table 4. 57 Table 16. Total Weights and Percentage of Each Species by Zones. Little Wolf Lake Zone 1. Zone 2. Zone 3. Total Species Weight % Weight % Weight % Weight kg. kg. kg. kg. Chara spp. 2,254 22.1 1,341 2.5 245 3.5 3,840 myriophyllum.heterophyllum. 847 8.3 2,347 4.3 3,194 Naias flexilus 2,214 21.7 6,370 11.7 664 9.6 9,248 Nymphaea advena 3,862 37.7 3,862 Pbtamogeton praelongus 1,038 10.2 44,225 81.5 6,003 86.8 51,266 Total -------------------- 10,215 54,283 6,912 71,410 Percent of Grand Total in each zone --------- 14.3 76.0 9.7 t‘ufv c I~ .u) «16 «f 23000 100.0 Zones V 4 ... , .. I V - ' “‘71",'\"u wen! ~r.’ :o .1? )1. p \ ~"' -- 1d . x Y» . -L -_ . U- ki‘)llul LIAA\7oCl‘1k. dunks- LA;,{4 _ .L'ul I .1. ,0 ABC U 0 ‘r t’: ‘0 I'\ 1"" -(J 'v“‘;‘-. ~i . .1 v“.{ L‘L'r ‘ ‘3‘ ' ')-I<.L.1.o.1 c L/lL,L1.'L...\ 1.11.011" (— -— —), £1301 Illiti‘l-nnt content (— —) 12* 2:01.05 in Little Joli“ Lake. |'-‘ (‘0 C23 58 59 13g Lake: Sections 2, 3, 10, and 11 of Greenwood Township, Oscoda County. This lake occupies a 210 acre basin in a relatively flat outwash plain area. Due to its peculiar shape, it has an unusually long shoreline as well as a wide variety of shoreline types. These vary from swampy margins to rather steep shores, and from extremely soft bottoms to gravel and' boulder shorelines. As a consequence, the plant zone is very irregular and variable, resulting in a greater number and variety of plant species than any of the other lakes. Due to the extent of the gradually sloping sand shore- line, a dense growth of sedges and bulrushes prevails. (See Figure 20.) As a result, this is the only lake in which Zone 1 produces a greater quantity of vegetation than do Zones 2 and 3. (See Table 18) The data are presented in the same manner as are those of the other three lakes. Fig.'18. A steep shoreline with coarse gravel bottom. Characteristic of entire east side of the main body of the lake. 60 Fig. 19. Fig. 20. A swampy margin-~at the north end of the lake. A gently sloping shoal with a zone of bulrushes and sedges. This type of margin characterizes the long narrow arm composing the southeast part of the lake. 61 *1: , b .L a . “" Oak, Aspen TEE LAKE A 00 t. . Scale= GEE-3;: ; Oak Man showing contours (in meters) and transects along which samples were collected. 21a. p2 Q LEGEND { ‘ " ' “ ‘ " "'v r'v_ 7,1 '.IN,.'51_"§' "7n ‘ I V. I- ’ ’l “I N ‘F *i‘ I Lees thehuioo 2000 to 4000 V /4 genera-WWW . ,/ / ‘ I A ‘ 100 to 400 4000 to 8000 V72 / ' ' -’“J i . 400 to 1000 8000 to 16000 '- ‘ gggyCYVJnvfl 1000 to 2000 16000 to 32000 f. e " ' 4.4... 0. 3,5/ , y .. ..‘e. ’ Q I - I t' . I. I . I a (I, :5. ’.. . .1" O Q \X K'1£.I(‘.(.)‘“ .. O ' 1' u' N 1K3, [yr/5173. ‘ ”>0 v I .- _ A ( ' ' ‘- 0 'V , , . . .’ '1 I ti, ~°' ’ ' xii-:53 .2415 . V ' . .v ’ i" \\ ,' f ’ff.Y-*7 "K' jébaJSQXXPL- d’.’ Fig. 21b. Map showing vegetation density in lbs. per A. ‘ LEGEND ‘\ // S--sand / b-«gravel 5 h--mar1 PP--yulpy peat FP--fibrous peat (f)~-ser.i-f1tid bottom F'P 213. Map showing bottom tLpes. - L). 64 65 Table 17. Plants Occurring in Tee Lake. Calla palustris Carex lasiocarpa Carer Pseudo-Cyperus Ceratophyllum demersum Chara spp. Cladium.mariscoides Dulichium arundinaceum Eleocharis palustris Equisetum.f1uviatile Eriocaulon septangulare Iris versicolor Lemna minor myriophyllum.heterophyllum Naias flexilis Nuphar advena Nymphaea odorata Potamogeton amplifolius P. angustifolius P. gramineus var. graminifol- ius f. longipedunculatus P. g. g. f. myriophyllus P. g. g. f. terrestris P.‘natans P. praelongus P. pusillus P. zosteriformis Sagittaria latifolia Scirpus atrovirens Scirpus acutus Sparganium.minimum Typha latifolia Water.knmn Sedge Sedge Coontail huskgrass; Stonewort Twig Rush Three-way Sedge Creeping Spike Rush Horsetail; Securing Rush Pipewort Blue Flag Lesser Duckweed Water Milfoil Bushy Pondweed Yellow'Water Lily White Water Lily Large-leaf Pendweed; Bass Weed Pondweed Pondweed Pondweed Pondweed Floating-leaf Pondweed Whitestem.Pondweed Pondweed Flat-stemmed Pondweed Duck Potato; Arrowhead Bulrush Hardstem.Bulrush Bur Reed Common Cattail 66 Table 18. Total Weights and Percentage of Each Species by Zones. Tee Lake Zone 1. 55 Zone 2. Zone 3. % Total Species ' Weight % ‘ Weight % ‘ Weight % Weight 0' i s. : Lg. 1 kg. Carex lasiocarpa 8,8161 6.9 1 i I 8,816 Chara spp. 549 0.4 E 77i 0.1 192 57.1 1 818 Cladium.mariscodies I 14,952 11.6 ‘ 3 f 14,952 Dulichium arundinaceum 275 0.2 ( t 275 Eleocharis palustris | 4,090) 3.2 : I 5 4,090 Eriocaulon septangulare 10,708? 8.3 i ! 10,708 Naias flexilus 15,207 10.5 < 13,845! 11.0 ' ' f 27,052 Nuphar advena 4,595 5.8 E v i , 3' 4,595 Nymphaea odorata 3,968 3.1 ' 3,968 Potamogeton amplifolius 2,069 1.6 3 63,420 50.3 i , 65,489 P. angustifolius 774; 0.5 g 574: 0.5 g * 1,348 P. gramineus var. gramini- ' ‘ folius f. myriophyllus i 1,274 1.0 54: --- 1,328 P. natans E 22,077,17.5 905; 0.7 1 22,982 P. praelongus i 10,945: 8.5 45,552; 55.9 ' 800 29.5 57,107 P. pusillus 1 155; 0.1 1,641 1.3 1,735 163.6 £ 3,531 P. zosteriformis ) 515 --- 514 0.2 g 555 Scirpus acutus i 27,014 21.0 . ' 27,014 Typha latifolia 2,730 2.1 2,730 Totals ------------------- 128,350 125,192 2,727 257,259 Percent of Grand Total in ‘ each zone --------- 49.9 -g[ 49.1 1.0 Table 19. Frequency of Occurrence and Relative Yield of Certain Plants on Different Bottom Types.l Tee Lake. 67 Bottom.Types Species S PP S,G S,PP Carex lasiocarpa 20* 5* 15* 25** 35** Chara spp. 48* 8* 44* Cladium.mariscoides 12* 4* 30* 54*** Eleocharis palustris 75* 12* 10** 3** Equisetum.f1uviatile 75* 8* 17* Eriocaulon septangulare 67* 4* 4* 25** Naias flexilis 18* 22* 4** 48** 8*** Nuphar advena 87* l3** Nymphaea odorata 65* 13* 22** 25* Potamogeton amplifolius 2* 33** 40*** P. angustifolius 2* 95* 7* P. gramineus var. gramini- folius f. myriophyllus 13* 87* 8* 20* P. natans 4** 22** 40*** 4* 15* 4* P. praelongus 56** 21*** 17* P. pusillus 33** 50*** P. zosterifonnis 95* 5** 17* 2* Scirpus acutus 43** 13** 25*** 1 Bottom types: S--sand; PP--pu1py peat; G--grave1. For explanation of table, see Table 4. l. nu C‘ 53L...) 6000 4500 3000 1500 Zones .i‘. Graph C‘ L.) lowing averafe yield (cr./ hect. percentage organic matter (-....-), and nutrient content (_____ x "I by zones 1‘0 Tee Lake. 68 69 DISCUSSION In addition to the presentation of the quantitative data, a discussion of some important ecological problems is warranted. These problems are reviewed in the light of, and with constant reference to, the data already presented. Bottom_Fertility: Before a discussion of fertility is undertaken, the term requires definition. The common definition is--the ability of a soil to produce vegetation, if seeded. Hence, the more vegetation a soil produces, the more fertile is the soil. In other words, soil fertility can only be measured in terms of its productivity. Fertility, then, embodies all of the factors responsible for plant growth. Fertility in lake soils, when considered from this viewpoint, becomes extremely complicated. There is no way to measure total fertility except by determining the plant yield, since fertility implies plant production. In such event, there must be a direct correlation between fertility and plant yield. In this study, the term "fertility" is used in a re- stricted sense. The phase of fertility studied was the nutrient content of the soils. As described previously, this included a determination of the supply, in readily soluble form, of four elements, namely, phosphorus, potassium, calcium, and iron. It was assumed that these are the most im- portant nutrients, and that if a correlation existed between nutrient content and plant production, it would be revealed 70 by this method. Further studies may show that other elements are important. Perhaps the minor soil elements such as boron, copper, zinc, and manganese have an active role in lake soil fertility. Pond (6) concludes, from a study of six aquatic plant species, that "the primary cause of the retarded growth of anchored plants is their inability to secure enough phos- phorus and potassium, and possibly other elements". Due to the quantity of data accumulated in this study, it became necessary to apply statistical methods. In order to do this, some means of expressing the nutrient content by a single figure had to be devised. This has been described previously. The validity of the method is subject to question. All the elements were given equal weight or importance. This was done because our knowledge of the physiology of aquatic plants has not progressed sufficiently so that we are able to say that one element is more important than any other. Another question which might arise in this connection is, "Does an increased nutrient content necessarily result in increased fertility?" It is possible that an increase of a given element beyond a certain point might prove toxic rather than beneficial. But in the soils tested, it is not likely that any element has been toxic. It is possible, however, that the depressing effect of the high calcium con- tent, in Lake Fifteen, on the solubility of the other nutri- ents has resulted in a much lower nutrient content figure than would otherwise be the case. 71 The chief objection to the method is that the figure itself gives no information regarding the individual constit- uents. Table 20 shows the mean nutrient content by zones, as well as the individual constituents. The table bears out the truth of the opening statement of the paragraph. Still another weakness is the fact that the figures from one lake can not be compared with those of another. For example, all of the nutrient content figures for Lake Fifteen are higher than those of the first two zones of Lake Twenty- two, and yet in the writer's opinion, the soils in Lake Twenty-two are better from the standpoint of nutrient content because they have enough calcium to supply the plants' needs, and they have a higher content of the other three elements. However, when considering one lake at a time, or comparing values within a single lake, the method is difficult to imp prove upon since, in every case, a higher figure means better conditions for plant growth from the standpoint of nutrient content. It is the writer's belief that this system, or any system embracing tests for several elements, can not be ap- plied in comparing different lakes because, as this study points out, lakes vary widely in total nutrient content from one another while soils within a given lake tend to vary much less markedly. See Table 20. An attempt was also made to determine whether or not the test for any single element might prove to be an index of the nutrient content. An examination of the data indica- ted that if such an index existed, it would be the phosphorus Table 20. Individual Constituents of Nutrient Content*. Lake Zone Phosphorus Potassium Calcium. Iron Nutrient PPma Ppm. ppm. ppm. Content L. Fifteen .14 1.1 200 0.0 24.5 .46 1.1 200 0.0 24.7 3 .80 3.4 200 0.0 29.8 Le Twenty-t0“) .74: 4.6 17.3 1.95 13.2 .85 4.4 47.7 1.77 16.9 1.80 11.0 58.0 3.00 32.7 Little VJOlf Lo 1 018 9.4 188.0 0.65 30.0 .04 6.5 200 0.0 26.8 .18 10.7 182.0 0.71 30.7 Tee L. 1.00 4.9 25.2 12.33 21.6 .72 5.2 39.9 13.57 21.7 .59 4.6 27.5 14.00 19.0 * See text for determination of nutrient content figure. 73 determination. Table 21 shows the correlation coefficients between phosphorus and nutrient content. While a significant correlation exists in the majority of cases, it is not likely that phosphorus could be used as an index. The difficulty again is that, while there is a correlation within a single lake, a phosphorus content of a given amount in one lake does not mean that the same amount in another lake has the same corresponding value. For example, in Table 20, a phosphorus content of .46 ppm. in Zone 2 of Lake Fifteen has a corres- ponding nutrient content value of 24.7, while in Zone 2 of Little Wolf Lake, a phosphorus reading of .04 ppm. has a corresponding nutrient content of 26.8. On the other hand, there seems to be a relationship between the readings or measurements within a single lake. However, a large number of measurements would have to be taken in order to determine whether or not, for a given lake, phosphorus could be used as an index. Hence, it would be advisable to assume that it could not be used, and to proceed with the more complete ex- amination as used in this study. OrganicHHatterIContent: Organic matter in a lake bottom is extremely important from the standpoint of carbon dioxide production. Brown (3) believes that the plants, by being rooted, show greater growth chiefly because they are held closer to the supply of carbon dioxide and can utilize it before it is taken out of solution by algae, marl precipitation, etc. Bourn (2) takes a median position between Pond and Brown by concluding that Table 21. and yield (N.C.--Y.), phosphorus content and yield (P.~4Y.), organic matter and yield (0.M.-4Y.), phosphorus content and nutrient content (P.--N.C.), and organic matter and nutrient content (O.N.-—N.C.). Lake Lake Fifteen Lake Twenty-two Little WOlf Lake Tee Lake Zone N.C.--Y. .371** .572 .500 .738 .725 .693 None None None None None None P. '"[e .315** .499 .644 .699 .705 .569 None *** *** None None None 0 .111. ”-Ye .978 .789 .389** None None .480** .449** .609 .580 None .694 None *Correlation coefficients between nutrient content PO--IIOCO .742 .781 .546 .859 .913 .814 None *** *** .580 None None .675 .640 None None None None None None None None None None * The correlation coefficients are based on the averages from.measure- Each average represents ments in each transect in each zone. from 3 to 5 measurements. ** Correlation coefficients not statistically significant. *** Not sufficient data for statistical analysis. 74 O 0111. ...-PI 0 C e 75' "while a deficiency of carbon dioxide is found to be an import— ant factor limiting the growth of aquatic plants, --a constant supply of carbon dioxide does not eliminate the difference between plants rooted in soil and those merely anchored in solutions“. Organic matter determinations were made by the hydrogen peroxide method. While this method is considered unsatisfactory in agricultural soils work because it does not determine total organic matter but rather, the more easily decomposed organic matter, it is believed to be entirely satisfactory for use on lake soils. Since bacteria in a lake are concentrated at the bottom soil-water interface, and since organic matter is con- stantly accumulating, only the more easily decomposed organic matter is in a position to be decomposed. Hence a better picture of what might be termed the “effective organic matter“-- that available for carbon dioxide production--is obtained by the hydrogen peroxide method than by methods which give total organic matter. Organic matter not only functions as a source of carbon dioxide, but also as a source of nutrients. Statistical analyses. however, do not show a significant correlation co- efficient between organic matter and nutrient content. except in Zones 1 and 2 of Lake Fifteen. The reason for the correla- tion here is probably due to the fact that an accumulation of organic matter reduces the effect of the calcium ion in rendering the other elements unavailable. In other words. in a mixture of marl and organic matter, the nutrients are avail- 76 able from the organic matter whereas in the marl they are present in a relatively insoluble form. No reason for the lack of a significant correlation in the other lakes can be offered at present. T_h_e_ Ecology gfqfluatic giants: The study of plant distribution in lakes has received some attention in this country and in EurOpe. Our knowledge of the factors involved, however, is far from being complete. Any study along this line adds to the fund of general informa- tion and perhaps aids in correlating the results of the scattered workers. The present study dealt with plant distribution in re- lation to depth, bottom, type, bottom fertility, and organic matter content of the bottom. Figure 23 summarizes the distribution of plants in the four lakes with regard to depth. The total weights, as given in this figure, do not reflect the actual productivity of the bottom since the zones vary in size. When considered on a yield per unit area basis, the result is quite different. Figure 24 shows the average yield by zones, in the four lakes. Here it will be noted that in three of the lakes, Zone 2 is the most productive. There is also a wide variation between lakes. This variation is probably exaggerated, how- ever, due to the fact that nearly all of the plants in Lake Fifteen were coated with a fine marl or lime deposit which could not be washed off; consequently, the weights of these samples were affected considerably. Hence, Lake Fifteen is 77 e of Vegetation J Tron tables 5, 0, 10, a 13. Total weights and percentan 0. u-JI . Fig. zone 0 in each a f. X? F/////////////////////////////// A fl fl fl////// a WV///MV/////////VVV/V%/////////A/W///V///// mmlm W////V/////fl//fl//,/ flmmmm WflWWHWV///V//fl/////, % DMMWOVVVVVVOQOQOOOC/ eictt Percent .m w .., a D W 7//////////// a E O 0 0 w 0 O O O 8 7 6 A. IU Ou l T T l x .r i i w 4 i . i i. a J O O 0 O 0 O O C O we AU MW MW .0 AU “w no no a m m m m m m m m m. m m m w Wu.— H l (u 8 r/ 6 5 Au no 2 l 70 On Zone T80 1.31:0 0 Little ’olf L 22 Lake 1.8120 15 78 we .../w H ‘ _ .9 .U 6 t +5 Etut- ‘3 54b ‘fi ..l . x L‘} c.) \. 151i “a“ A (A V‘ 1 ' ~. . ix“; L 0"; COO QOO F" 900 000 >- ,. l coco 6000 L H OJ C 7 5 1 II: | VII 'Inh'. II ‘7» ‘-|| ’1 ‘1 ll: 3 1 .1 l I .. -_-._.1..,w.--,-m I on 1 gone C he .‘9 / £J no 3000 L ooo W. 79 not as productive as the graphs indicate, but it is undoubt- edly more productive than any of the other lakes. It is interesting to note that Lake Fifteen produces the smallest crop, and yet, on the basis of yield per unit area, is the most productive. The reason, of course, is that the plant zone is relatively small but the vegetation is dense where it is able to grow. In Lake Twenty-two, a very productive Zone 3 is found. The shape of the basin is believed to be largely responsible. A gradually sloping bottom is found around a good share of the lake; this is usually accompanied by greater growth in deeper waters. This phenomenon undoubtedly is an important factor in determining the total productivity of a lake. Table 21 shows the results of a statistical analysis pertaining to yield as influenced or related to the nutrient content and organic matter content of the bottom. In Lake Fifteen there is not a significant correlation in Zone 1 between nutrient content and yield, while there is in Zones 2 and 3. Wave action, which is severe in this lake, is likely responsible for inhibiting vegetative growth in Zone 1. Wherever organic matter has an opportunity to accumulate, however, vegetation gets a "foot-hold" and grows well. In Zone 5, of course, the organic matter content is high and rather uniform so a significant correlation does not exist. In Lake Twenty-two, there is a good correlation be- tween nutrient content and yield, whereas there is not between organic matter and yield, chiefly because the 80 organic matter content is rather uniformly high. In Little Wolf Lake, however, where there is a lower average organic matter content, there is a correlation between the organic matter and plant yield. From a consideration of the data presented, it can be pointed out again that the distribution of vegetation is the result of the interplay of many factors. Physical factors are, in the opinion of the writer, of primary importance. Wave action, where it is extreme, prevents plants from grow- ing abundantly, in spite of the possible available nutrients. Deeper water generally means a.more fertile bottom, from the standpoint of nutrient content. But again, light penetration and perhaps temperature inhibit plant growth. The consistency of the bottom.is important from.the standpoint of the rooting ability of the aquatics. Organic matter content, considered both physically and chemically, is important, and determines the nature and quantity of plant growth under certain con- ditions. In a lake with a sand or marl bottom, such as is found in Little'Wolf Lake and Lake Fifteen, respectively, an addi- tion or accumulation of organic matter in a region of suitable depth has a very noticeable effect on plant growth. This is undoubtedly due to a change in consistency of the bottom, the production of carbon dioxide, and the release of plant nutri- ents. On the other hand, if the organic matter reaches a certain limit, it seems that the further addition of organic 81 matter has little or no effect. This explains why, in Lake Twenty-two, there is no correlation between organic matter and plant yield. This same principle seems to apply in relation to nutrients. A deficiency of any one nutrient may retard plant growth, but addition or presence of a small amount of the same nutrient will give a substantial increase. Addi- tion beyond a given point will have no effect, until so much has been added that toxicity results. Referring back to the Rose Lake plots, the addition of 500 pounds per acre of phos- phorus fertilizer yielded excellent growth; the addition of 2000 pounds per acre yielded no better. This proves a phos- phorus deficiency, which was previously indicated by chemical tests, and its remedy by the addition of phosphorus. Another phase of the ecology of aquatic plants is the relation between various species of plants and their growth on different bottom.types. Tables have been presented for each lake to show the distribution of vegetation with respect to bottom type, and also with respect to depth of water. It may be pointed out that for most plants, there is one, or pos- sibly two, bottom types on which.the plant is found most fre- quently and in the greatest abundance. The same is true for depth--the majority of plant species tend to be concentrated in one depth zone. An examination of separate items reveals some inter- esting facts which are obscured in condensed and summarized data. It is found that a given type of bottom, Which a plant 82 occupies most frequently, is generally situated or located at the depth at which the given plant usually grows. This brings up the question, then, "Does the plant 'prefer' that type of bottom, or does that bottom Just happen to occur at the depth which the plant 'prefers'“? This question has received attention in the past. Lakes in England have been studied by Pearsall (5) and Misra (4). They have both pointed out the different kinds of vegetation associated with different bottom soils, and have worked out plant successions beginning from rock bottoms and continuing through the dominantly organic bottoms. In their discussion of plant succession, the general trend is to consider success- ion the result of chemical and physical changes in the bottom. This is undoubtedly the case, but it seems that too much emphasis has been placed on the chemical nature of the bottom, and not enough/the physical nature of the bottom or the other components of the water soil profile as prOposed by Veatch (10), chiefly the nature of the aqueous horizon. This horizon constitutes a major part in the environment of aquatic plants; the lake bottom--accumulative horizon--serves, in some cases. merely for anchorage. The parent material--the original lake bottom--functions in plant distribution only when the accumu- lative layer is absent or very thin, so that roots penetrate through it'to the parent material. While most of the work is being done on the lake bottoms, the above conception of the soil profile must not be neglected because the chemical and physical nature of the water above 83 the bottom.are equally as important, in the ecology of aquatic plants, as is the type of bottom. There are few, if any, lakes in the Southern Peninsula of Michigan which are as primitive as some of the lakes studied in England by Pearsall and Hisra. There are few lakes, for in- stance, whose bottom is 70 percent rocky to a depth of thirty feet. In the lakes studied, however, almost all conceivable bottom.types are represented, ranging from a clean sand or marl to those Which are highly organic and semi-fluid in consistency. As pointed out above, most plants are found more fre- quently on a given type of bottom.than on others. But, it seems that plant distribution is affected more by physical factors than by those that are chemical, since a careful study of the data from.the four lakes fails to show any indication of a consistent "preference” on the part of any plant with re- gard to the chemical nature of the bottom as determined by the method used. The writer's inference is that physical factors, wave action in particular, determine to a large extent the distribution of bottom types in the portion of the lake in which vegetation can be produced. The bottom types, then, together with the active physical forces, determine the dis- tribution of the vegetation. Under conditions of extreme wave action, this force plays a more active role in plant distribution than does the type of bottom; under such condi- tions, 3 wide variety of bottom types will not be formed, except as the parent material varies. When conditions with respect to wave action are less pronounced and temperature. 84 depth, and light penetration are adequate, the character of the bottom.type occupies the determinant role in plant dis- tribution. Factors involved here are: nutrient content, when one or more plant food elements approach the point where they are either deficient or toxic; consistency of the bottom, determining the penetrability by roots; and organic matter content of the bottom, which affects the consistency and the nutrient content as well as the production of avail- able carbon dioxide. Relative productivity ‘pp_f__1_t_lge__p_f_g_u_r__l._a_k_e_§: As pointed out by Welch (11), an adequate index of pro- ductivity has not been determined. Total vegetation was con- sidered but did not prove satisfactory. Perhaps, however, a consideration of the yield per acre, figuring the entire lake, together with the percent of the lake which produces vegetation, would give a fair index to the productivity. The following table summarizes the information suggested above: 1.. 15 L22 fififiv ‘ Te: L. Area (acres) 88 109 87 210 % productive bottom. 31 48 54 52 ‘Yield/acre of lake (lbs) 1,220 2,750 1,780 2,695 ’“Shoreline develOpment 1.4 2.0 1.75 5.4 ’*The shoreline development is the ratio between the circumfer- ence of the lake and the circumference of a circle having the same area as that of the lake. This measurement is considered to be of value because it indicates the relative amount of shoreline and shallow water, which is important biologically. 1|...le .bi'b'llnl!!|...'b’uis.lflh51f N 85 From.these data it might be concluded that Tee Lake and Lake Twenty-two would be the most productive. Little Wolf Lake has a higher percentage of the bottom producing vegetation but the average yield is lower and the shoreline development is less. Lake Fifteen would be considered the least productive, in spite of its being most productive on the basis of the average yield of the vegetation zone. (Fig. 24) A On the basis of shoreline development, Tee Lake would probably rank higher than Lake Twenty-two. groductivity“35hindicatggwpwaighmgrowth‘studies: Productivity, to the fisheries biologist, is naturally measured by the production of fish. Hence, an attempt was made to determine the growth rate of the fish in the four lakes. Unfortunately, large enough samples of fish could not be taken in the fall of the year to make a satisfactory study. However, the data obtained will be presented since they at least may be indicative of the actual condition in the lake. See Table 22. Indications are that the perch in Lake Twenty-two are growing much faster than are those in any other lake. The data on bluegills is rather scant but it is interesting to note the difference in growth between the two basins of Little Wolf Lake. Although only one specimen was taken from any age group in the north basin, there were no speci- mens from.the corresponding groups in the south basin which reached the length of the north basin specimen. If this 86 condition exists generally, it reveals a contrast undoubtedly brought about by the difference in the two basins. since the north basin is shallow and rich in vegetation while the south basin is deep and contains little vegetation. It is doubtful that migration occurs through the shallow channel between the two basins. Here work on the comparison of the fish production in these four lakes would prove extremely interesting and valu- able. It would be an Opportunity to learn whether or not there is a direct relationship between the amount or extent of the vegetation and fish production. The data presented here may be indicative, and do agree for the most part with the pepular viewpoint, which is often fairly accurate. Lake Twenty-two is considered by far the best perch-fishing lake in the vicinity; Tee Lake is a popular bass and bluegill lake, and sizable catches are reported; Little Wolf Lake is fairly popular for its bluegills; Lake Fifteen attracts fewer anglers than do the others and the catches are reported to be very small. 87 .mpqomeamon omwaops onp moans mnesaoomm we Rosana one seesawsfi momonpnoawm a“ moasmHm any ms Haamosam kmvn.mma Leave.oma Amelo.maa Lem. n.nma scams mama mos Rev s.meH any s.mma xmv m.mm Anav n.mm Haamosam names endow “Hy mom AHV ska Adv mma Haammsam mamas apnea Lav sea may and Lav Hm scams saga ones mappfia Amy m.a¢a any o.ooa Haamosam Ana n.4mm xoavm.omm ”mam.nmm any ooa Assamesev any o.msm . Lav o.msa Asa m.emfl xmoassv nosed ospnhpsoaa exam Adv and Assam seasons mesa HH> He _> pH HHH HH H mmaomam a snag Amheowg macho owm Awesomeaaaas sa_aopspmv .mmqu asoh amp seam nmwm mo mapmsog Hobos .NN manna 88 SUMMARY A study has been made of water soils in relation to the productivity of lakes. Reconnaissance studies were first made on bottom soils from lakes throughout the state. Pot tests were made to determine the effect of fertilizing materi- als on plant growth; these were followed by similar tests on plots at Rose Lake. The final phase of the investigation con- sisted of a comprehensive study of the soils and vegetation of four lakes in the northern part of the Lower Peninsula. Conclusions reached as a result of the work are: 1. As shown by the Rose Lake plots, definite nutrient de- ficiencies may occur in lake soils, and these deficiencies can be remedied by the addition of fertilizer. ‘Modification of the method of application would probably be necessary for lake improvement work. 2. The greenhouse tests show that the addition of fertil- izing materials, in some cases, favors the growth of algae to such an extent that they interfere with the growth of the higher plants. It is doubtful that the problem.would be as acute in lake work as it is in the greenhouse, but there would be competition for nutrients between algae and the higher aquatics. 3. The study of four lakes shows a variety of conditions and an attempt has been made to interpret or understand the importance of various factors and their effect on lake pro- ductivity. 89 4. On the basis of the fertility tests, it is found that the average nutrient content of the bottom soils varies con- siderably between different lakes, and that the method used. can not be safely applied in comparing different lakes but is quite satisfactory in evaluating the comparative fertility of the soil samples from.the same lake. 5. There seems to be no single test which will serve as a satisfactory index to the nutrient content of the bottom soils. 6. Lakes vary in plant distribution; Zone 2 is generally the most productive of vegetation, however. 7. Plant distribution is governed largely by physical fact- ors, including shape of the basin, wave action, light penetra- tion, and consistency of the bottom. Under conditions of low nutrient content or low organic matter content, there is a direct relationship between these factors and plant yield, but when there is a relatively high organic matter content and nutrients are available in sufficient quantities, changes in these two factors do not result in corresponding changes in plant yield. 8. Bottom type distribution is determined to a great ex- tent by wave action; the character of the bottom, together with the other active physical forces, then control the plant distribution. The extent to Which the bottom type or other factors influence plant distribution varies locally. 9. Although there is no satisfactory index of productivity in lakes, from the standpoint of the fisheries biologist, indications are that a consideration of the shoreline deveIOp- ment; the percent of the bottom producing vegetation; and the average yield of vegetation, considering the entire lake, will give a fair index to the productivity. 90 1. 2. 3. 5. 7. 8. 10. 11. 12. 91 LITERATURE CITED 1955. ‘Methods of analysis. Assoc. of Official Chemists. Washington, D. C. ‘ Bourn, W. S. 1952. Ecological and physiological studies on certain aquatic angiosperms. Cont. of Boyce Thomp- son Institute for Plant Research. 4: 425-496. Brown, W. H. 1915. The relation of the substratum to the growth of Elodea. Philippine Jour. of Sci.. C. Bot. 83 1-200 Hisra, R. D. 1958. Edaphic factors in the distribution of aquatic plants in the English lakes. Jour. of Ecology. 26: 411-451. Pearsall, W. H. 1926. Dynamdc factors affecting aquatic vegetation. 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