fresh water algal succession

Fresh Water Algal SuccessionAuthor(s): Samuel EddySource: Transactions of the American Microscopical Society, Vol. 44, No. 3 (Jul., 1925), pp.138-147Published by: Wiley on behalf of American Microscopical SocietyStable URL: http://www.jstor.org/stable/3221462 .

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FRESH WATER ALGAL SUCCESSION

BY

SAMUEL EDDY

James Millikin University, Decatur, Illinois

Pond and stream succession in relation to forest and grassland development has been described by Cowles (1901). Transeau (1903-1905), Cooper (1913), and others have described bog succession in the same relation. Shelford (1913) and Allee (1911) have noted the sequence of animal life in the various stages of stream and pond development. However, very little seems to have been done on succession in relation to algae and the development of the aquatic habitat itself. Aquatic succession is just as complicated and as important as terrestrial succession. It has its own in- dividual climaxes apart from those which form the basis for certain terrestrial sequences. Aquatic succession is a fundamental primary succession establishing not only a basis for terrestrial succession but completing a sere of its own.

Primary succession (Clements, 1917) is that succession, the initial stage of which starts on a barren area. When the continental ice sheet receded, a vast area of barren till remained. Upon this the initial stage of primary succession started. Many small ponds and lakes established themselves in the basins and furrows between these ridges and banks, some of which remained until recent times as our typical prairie sloughs and swamps. Small streams trickled over the rough banks and ridges of glacial till. As these small streams aged, they eroded back and drained larger areas, en- larging both their channels and volume, forming small rapids and interrapid pools. The rapids remained and still remain ecologically young and represent the condition of a young stream. The more sluggish waters of the interrapid pools soon acquired lake or pondlike cpmmunities of organ- isms (Cowles, 1901). When the stream had worn its channel deep enough, oxbow ponds were formed by the channel cutting off and isolating some of the interrapid pools. These oxbow ponds with their lake communities then commenced or continued a typical lake succession ending in a flood plain forest. The shallow glacial lakes and ponds did not receive the eroding influence of the stream to give them a flood plain topography. Their aquatic climax ended in a grassland instead of a flood plain forest.

Stream succession may pass through three main or principal seres, the oxbow pond just mentioned, the deposition or sand bar sere, and a river sere. The deposition sere has been discussed by Cowles (1901) as sand bar succession. It ends in a flood plain forest as does the oxbow pond sere, both of these seres ending in terrestrial succession. The stream itself continues in an aquatic succession, reaching a climax in a large sluggish river. The

138




ultimate end of all streams is a base level stream where all rapids are de- stroyed and the interrapid pools become continuous. Ruthven (1905) states that a river in old age acquires the conditions of a pond and has much the same biotic communities. In relation to this sere, the oxbow pond and deposition seres are only minor seres. The principal sere of succession is in the stream itself. Just as in the climax forest, retrogression and restoration are constantly going on in various places. The banks and channels are constantly eroding and depositing. The stream community is just as much of a unit organic complex as the ecological equivalent in the forest or grassland.

SMALL STREAM SUCCESSION

In all primary hydrophytic succession the initial stage should start in a still or running body of water absolutely barren of life or the remains of previous organic occupation. According to Shelford (1913), a stream is always in the initial or youngest stage at its source and oldest at its mouth. Under these conditions the biotic communities in these habitats should represent the same equivalent ecological ages. In this relation a small stream near Muncie, Illinois, was studied in its various stages. The stream rose from a spring in glacial till near the prairie level and descended in a quarter of a mile through rapids and pools to Stony Creek sixty feet below the prairie.

The initial sere was represented by the water trickling from the glacial till down into puddles made by cow tracks. The water had a hydrogen ion concentration of pH 7.4 on November 1 and pH 7.2 on April 25. The hydrogen ion concentration did not seem to vary much from source to mouth, except that on April 25 it was pH 7.4 at the mouth due to the abundance of algae which were not established at the source. At the source, the bottom was covered by a brownish sediment of diatoms (mostly Synedra). The water had a greenish tinge both in November and April caused by large numbers of Euglena. Phacus, also, was abundant in November.

After the stream had descended about 20 feet, the flow of water in- creased until it had a depth of six inches. At this stage, filamentous algae (Vaucheria and Mougeotia) first appeared in November. Tetraspora and Endorina were also common. In April this stage was the place for the first appearance of Spirogyra. No other algae except diatoms were present.

At the 40 foot level, Diatoms and Spirogyra were abundant in Novem- ber. In April Spirogyra, Mougeotia, Diatoms, Cosmarium and Desmids were abundant. On November 1 at the 60 foot level, only a few feet from the mouth, Spirogyra and Diatoms were abundant in the small interrapid pools which had formed at this stage. In the rapids, Cladophora was abundant. In April, the same algae were abundant in the same places. A few small patches of Closterium and Apiocystis were scattered on the bottom of the pools under the rapids.

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SAMUEL EDDY

This illustrated the stages in a primary sere of the succession of a small stream from its origin until middle age. The youngest stages showed an absence or scarcity of filamentous algae. Diatoms and Euglana were dominant. A middle age stream has reached what might be termed the prime of succession or growth. This is the age of maturity where it can give birth to oxbow ponds and desposition seres. From this stage it continues into old age or the climax.

MIDDLE AGE STREAM SUCCESSION

Stony Creek represents an early middle age stage. It is mature enough to form ox bows and to deposit sand bars of recognizable size, but it yet retains many rapids which are a sign of youth. The rapids represent the youngest stages after the small stream. Cladophora, Oscillatoria and filamentous Diatoms were the only algae present. In the quiet sluggish pools between the rapids, Spirogyra was abundant. These pools were ecologically much older than the rapids. The Sangamon River near White Heath, Illinois, although about the same size as Stony Creek, was much older ecologically. The channel was worn smooth and free from rapids forming a sluggish stream of almost uniform pool composition. Spirogyra was very abundant here and Cladophora was very rare. This was more nearly a typical middle age condition. In both streams sand bars were being formed either as typical mid-stream bars or as banks deposited at the expense of the bank on the other side. Spirogyra grew almost to the water's edge of the bars. Water smartweed and sand bar willows (Salix longifolia) grew down into the water. These plants, by holding back the current, were causing more deposits of sand, which would be expected to result in time in the sand bar sere, ending with a flood plain forest as an edaphic climax.

The third or oxbow pond sere is illustrated in a group of small ponds nearby formed by the cutting off of a portion of Stony Creek by a railroad embankment. Here are three ponds, of the same age in years, but represent- ing different ages of ecological succession. The youngest and largest pond is two to three feet deep with a mud and clay bottom. Elodea was very abundant in the clear water, which has a hydrogen ion concentration of pH 7.6 to pH 7.4. Diatoms were not abundant in the Fall or Spring. Spirogyra, Ulothrix and Zygonema were abundant at both seasons.

The second pond represented a middle age pond. It was filled with mud from decaying vegetation until the water was only six to twelve inches deep. The water had a hydrogen ion concentration of pH 7.1 to 7.2. Bidens connata was intruding from the shores and had covered almost half of the pond. Chara and Hydrodictyon were very abundant in November, but their growth had not started in April. Diatoms, Oscillatoria, and Phacus were abundant in April.

140




The third and oldest pond was only a marsh. In a few years it would be developed into a typical flood plain subject to flood plain conditions.

Willows (Salix nigra) and buttonbush (Cephalanthus occidentalis) were invading from the shore. Swamp beggartide (Bidens connata) had spread fr.om shore to shore. The water was only about six inches deep over a deep mud deposit which composed the bottom. The hydrogen ion concentration was pH 6.9-7.0, showing a slight acidity. No filamentous algae were present in either season. A few Diatoms and Closterium seemed to be the only algae present. On the trunks of the willows were occasional patches of Pleuroccocus which indicated that the last stage of algal succession may overlap the earlier terrestrial stages of forest succession.

PRIMARY POND SUCCESSION

The study of Primary Pond Succession from the initial stage as left by the recession of the ice sheet is a difficult problem owing to the extreme scarcity of such habitats. MacDougall (1918) studied the succession of vegetation on glacial till under immediate post-glacial conditions in the strip lands near Danville. Here for the last thirty years, the coal company has been removing the top soil, leaving fresh areas of glacial till exposed each year. Also, each year fresh ponds are formed with bottoms of virgin glacial till. This gives almost the same conditions as that of the early stages of our recent prairie sloughs which originated as basins between ridges of glacial till. The ponds of the post-glacial period did not have as good reseeding conditions, or, as favorable climatic conditions as these present artificial ponds. No doubt, the close proximity of good reseeding areas greatly hastens the succession so that what takes place in the modern ponds in a few years probably repeats the succession of a few centuries in the glacial pond.

The initial stage was illustrated by a pond formed within the last year. The bottom was glacial till with no sediment apparent. The water was clear and without any apparent color. It had an hydrogen ion concentration of pH 7.6, showing considerable alkalinity. No filamentous or green algae, except a few Euglena, could be found. Diatoms, especially Synedra, were very abundant.

The next stage consisted of a large seven year old pond. This pond was unusually large, covering six or seven acres, and the size probably altered its comparative value. The water was clear with a slight greenish tinge, and had a pH of 7.3. The bottom was glacial till with considerable sediment. Much of this sediment seemed to be composed of Diatoms, which form an important part in pond deposition. Patches of Rhizoclonium hierglyphicum were abundant along the shores. This seemed to offer promise of developing into a flourishing algae habitat later in the season.

141



SAMUEL EDDY

The next stage was illustrated by two smaller ponds ten years old. The first had considerable deposits of sediment on the clay and contained numerous patches of Vaucheria. The other pond contained no filamentous algae, but the bottom was covered with a sediment of Diatoms. A few Oscillatoria were also present. The pH of the first was 6.8, and the latter had a pH 6.6. This acidity of the latter may have some influence on preventing the growth of filamentous algae. It would stand for old age as compared with the oxbow ponds.

The next stage of succession was represented by a shallow 23 year old pond. This seemed to represent the stage of a prairie lake which was semi- filled with rushes and aquatic vegetation. The bottom of this pond was covered with a thick layer of silt deposit. Large quantities of an aquatic sedge grew over the bottom and probably covered the surface later in the summer. The water had a pH of 7.6, probably due to the high dissolved oxygen content resulting from so much vegetation. No filamentous algae were to be seen. Diatoms were extremely abundant and were the only form of algae present.

The last aquatic stage of succession was represented by the remains of a pond 27 years old. This pond had filled with sediment until the bottom was covered to a depth of many feet with a heavy ooze. Owing to the fact that the strip lands are located on a flood plain, the flood plain forest was rapidly invading instead of the grassland, which is the edaphic climax of the typical prairie pond. The water was only a few inches deep and had a hydrogen ion concentration of pH 7.2. It was covered by a greenish scum of algae. This was composed of large quantities of Diatoms, Oscillatoria and Euglena. None of the higher filamentous algae seemed to be present, because of the large amount of decomposition to which the habitat was subject. The acidity was probably very great at times. Also, the temperature fluctuated widely with the air, because of the shallow depth and small volume of water. In the larger bodies with a more stable temperature, conditions in this respect seemed to be better suited for algae.

CONCLUSION

Primary succession of algae may continue in two major seres, a stream or a prairie pond sere. In both seres the initial stage is very similar, starting under barren conditions and containing unicellular algae' as pioneers. Ac- cording to Summerhayes and Elton (1923), the flora of the immediate post- glacial lakes and streams of Bear Island and Spitsbergen consist chiefly of unicellular algae. The few filamentous algae, such as Ulothrix and Phormidium, which are mentioned, are those types which grow under rather

1 Since this paper was prepared, 0. T. Wilson has published a series of experimental observations on marine algal succession, (Ecology, VI, 3, pp. 303-311). His observations have shown that a dominance of diatoms constitute the first stage of marine algal succession.

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barren conditions. Diatoms are reported in great abundance in some lakes. In some material collected from the swift young mountain streams of North- western Canada, the writer found that Diatoms, Euastrum, and Cos- marium constituted most of the flora. Filamentous forms were rare and consisted chiefly of Ulothrix. Diatoms were abundant in the glacial streams to within a few feet of the ice. They were also very abundant in the swift waters of the Bow, Kicking Horse, and Athabasca Rivers.

In sterilized tap water cultures, exposed to air and sunlight, Diatoms appeared at the end of three weeks and Euglena and Desmids at the end of the fourth. In a small puddle formed by melting snow large quantities of Diatoms appeared the fourth day. Thus, in all probability, unicellular

Old age River- Climax

0. Lood Plain Forest Graessland, etc. (Climatic climax)

Slough Ox Bow Pond

Young Prairie Lake or Pond

Initial stage.

FIG 1. Diagram showing the relations of the different seres of algal succession.

algae, such as Diatoms and Euglena, are pioneers of the initial stage. Seed- ing conditions for these forms are generally much better than for the filamentous types. From the early appearance of Flagellates and Diatoms in sterile cultures and initial stages, it is very evident that the encysted forms of these species are very widely and readily dispersed. The higher types of filamentous algae are either not so readily dispersed, or do not possess so wide a range of adaptability and require more favorable conditions than offered by the initial stages.

143



SAMUEL EDDY

From the initial stages, the succession of algae in streams and ponds run a parallel course. Conditions steadily improve for maximum abundance of algae as the habitat approaches middle age. The conditions of an old river approach very closely to those of a late middle age pond. According to Kofoid'(1903), the waters of recent origin contain little plankton, but as the stream becomes older, lake conditions become more pronounced and plankton becomes very abundant in both stream and lake. The current of the Illinois River is not as rapid as that of smaller streams, and, consequently, contains many more plankton types of algae. Also, according to the same author, the Illinois River has a great abundance of limnetic algae and a smaller stream, the Spoon River, has almost all littoral species. In the Illinois River, according to unpublished data, both filamentous and unicellular algae are extremely abundant. This condition equals that of a middle age lake in which the same abundance might be expected.

In the middle age stage of stream succession, secondary seres of oxbow and sand bar succession may arise. These two seres may parallel each other in the first stages, but the former soon assumes the terrestrial aspect while the latter continues to follow the prairie pond sequence. The stream itself continues into a permanent old age or climax. This climax might be termed edaphic, but should more properly be considered as climatic, although not conforming to the same climatic formations as terrestrial communities. The small stream represents youth, the medium stream middle age, and the sluggish river with its large well-worn channel represents old age. Age varies according to physiographic conditions, and accordingly many streams age more quickly than others. After the young stages conditions rapidly become more favorable for algae. These conditions increase much more rapidly in the pond so that the algae communities of an old river resemble those of a middle age pond. Size and depth are the factors deter- mining the velocity of age in the pond. As the pond ages and vegetation creeps in, the plankton declines. Kofoid (1903) states that the amount of plankton produced by bodies of fresh water is, other things being equal, in some inverse ratio proportional to the amount of its gross vegetation of the submerged sort.

Finally the filamentous forms give way to a few unicellular forms. At the very last Pleuroccoccus on the trunks of the invading willows is usually the last trace of an all but completed algal succession.

The writer wishes to express his appreciation to Dr. W. B. McDougal and Dr. V. E. Shelford of the University of Illinois, and to Mr. R. E. Richardson of the Illinois Natural History Survey for many suggestions and criticisms.

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145 FRESH WATER ALGAL SUCCESSION

TABLE 1

Young Stream Succession November 8, 1924

Station Source Medium Medium Mouth

Level 6 ft. 20 ft. 40 ft. 60 ft.

Date Nov. 1 Apr.25 Nov. 1 Apr.25 Nov. 1 Apr.25 Nov. 1 Apr. 25

pH 74. 7.2 7.4 7.2 7.4 7.2 7.4 7.4

Euglena sp. X X X Diatoms (Synedra) X X X X X X X X Phacus sp. X Tetrospora X Eudorina X Zygonemales

(Mougeotia) X X Siphonales (Vaucheria) X Spirogyra X X X X X Cladophora X X Cosmarium X X Closterium X X Apiocystis X

TABLE 2

Middle Age Stream Deposition Succession (Sandbar), Stony Creek, Muncie, Illinois November 1, 1924

Locality Rapids Interrapids Sandbar

pH 7.6 7.4 7.4

Temp. 11?C. 11?C. 11?C.

Oscillatoria abd. Cladophora abd. Diatoms, (Matrix) Gomphonema? abd. Diatoms, (Motile) occ. Spirogyra abd. abd. Water smartweed abd. Willow (Salix longiflora) abd.



SAMUEL EDDY

TABLE 3

Ox Bow Pond Succession. Muncie Illirois

Age

pH

Temp.

Date

Spirogyra Ulothrix Zygonema Diatoms (Synedra) Closterium Diatoms (Pleurosigma) Phacus Oscillatoria Hydrodictyon Chara Tetraspora Merismopedia glauca

Young

7.6 7.4

18?C.

Nov. 1 Apr. 25

occ. abd. abd. abd. abd. occ. occ. rare

OCC.

OCC.

OCC.

Medium

7.1 7.2

18?C.

Nov. 1 Apr. 25

OCC.

OCC.

abd. abd.

abd.

OCC.

oCC.

OCC.

Old

6.9 7.0

18?C.

Nov. 1 Apr. 26

rare OCC.

TABLE 4

PRIMARY POND SUCCESSION STRIP MINE NEAR DANVILLE, ILLINOIS, MAY 2, 1925

No. 1 2 3 4 5 6

Age 1 yr. 7 yr. 10 yr. 10 yr. 23 yr. 27 yr.

Bottom Clay Clay Clay Clay Silt Mud

pH 7.6 7.3 6.8 6.6 7.6 7.2

Temp. 18?C 17?C 17?C 17?C 18?C 15?C

Diatoms (Synedra) abd. abd. abd. abd. abd. abd. Euglena occ. abd. Rhizoclonium

hierglyphicum abd. Diatoms, Amphora abd.

Navicula abd. abd. abd. abd. Pleurosigma abd.

Vaucheria abd. Oscillatoria occ.

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FRESH WATER ALGAL SUCCESSION 147

LITERATURE CITED

ALLEE, W. C. 1911 Succession in old Forest Ponds. Trans. Ill. Acad. Sci., Vol. IV, pp. 126-131.

CLEMENTS, F. E. 1916 Plant Succession, Carnegie Institution of Washington, pp. 512.

COOPER, W. S. 1913 Isle Royale. Bot. Gaz. Vol. 55 pp.

COWLES, H. C. 1901 Plant Societies of Chicago and Vicinity. Bull. Geog. Soc. Chicago. No. 1. 1901 The Physiographic Ecologyof Chicago and Vicinity;a Study of the Origin, Develop-

ment and Classification of Plant Societies. Bot. Gaz. 31, pp. 73-108; 145-182. KOFOID, C. A.

1903 The Plankton of the Illinois River, 1894-1899. Part 1. Bull. Ill. State Lab. of Nat. Hist. Vol. VI. Art. 11.

McDouGAL, W. B. 1918 Plant Succession on a Artificial Bare Area in Illinois. Trars. Ill. State Ac. Sci.Vol.

XI. pp. 129-131. RUTHVEN, A. G.

1906 An Ecological Survey of the Porcupine Mountains and Isle Royale, Michigan. Geol. Surv. Mich. Ann. Rep. 1905, pp. 17-55.

SHELFORD, V. E. 1913 Animal Communities in Temperate America. pp. 362.

SUMMERHAYES, V. S. and ELTON, C. S. 1923 Contributions to the Ecology of Spitsbergen and Bear Island. Jour. of Ecology, 11;

214-286. TRANSEAU, E. N.

1903 On the Geographic Distribution and Ecological Relations of the Bog Plant Societies of Northern North American. Bot. Gaz. 36, pp. 401-420.

1905-1906 Bogs and Bog Flora of the Huron River Valley. Bot. Gaz. 40. pp. 351-375; 418-448; 41, pp. 17-42.



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