Digitized by the Internet Archive in 2009 with funding from University of Toronto http://www.archive.org/details/biologicalseries 17univ LOGICAL SERIES iy G (] W. A. CLEMENS: i ‘ - Host Dat Mae) hey, ry : ¥ ! . % . . € ; Ns “ak at ; ly ¥ yi} \ \ ) ny \ RY: PUBLISHED BY STUDY OF THE MAYFLY University of Toronto Studies COMMITTEE OF MANAGEMENT Chairman: StR ROBERT ALEXANDER Fatconer, LL.D., K.C.M.G. President of the University Proressor W. J. ALEXANDER, PH.D. Proressor W. H. E tis, M.A., M.D. PROFESSOR J. J. MACKENZIE, B.A., M.B. Proressor J. P. McMurricu, Pu.D. Proressor G. H. NEEDLER, PH.D. PROFESSOR GEORGE M. Wrone, M.A. General Edttor: H. H. Laneton, M.A. Librarian of the University AN ECOLOGICAL STUDY OF THE MAYFLY CHIROTENETES BY WILBERT AMIE CLEMENS, M.A. TABLE, OF, CONTENTS = ET ARR a a Se THE NympH: General habitat, Chemical analyses of the water, Temperatures, Relation to materials forming the creek bed, Relation to light, Relation to current, Food, Relation to the oxygen and carbon dioxide content of the water, Associates, Enemies, Severities of stream life, Protection, RemetamR Nira an ane ee ee wis chs, 5h 22 THE Supimaco: Emergence, Factors effecting the length of the subimaginal period, Enemies, 2 EYE dis gan RE STOR oe Re as ON TO TuHE Imaco: Transformation, Flight, Oviposition, Egg, Enemies, Protection, Length of life............ RMR e ee a as ido CUM PPh. es Lr JA Ye eek we eee BeeeTTEeIOH. PEATHS. 0.) 56 ti ok be ele ele wie aye aK nv pat “elt i ; AN ECOLOGICAL STUDY OF THE MAY-FLY CHIROTENETES INTRODUCTION Notwithstanding the numerous investigations which have been made upon the life phases of various Ephemeridae, our knowledge concerning the physical factors and their co-operation in relation to individual species is singularly small. It was for this reason that the writer, on the sugges- tion of Professor James G. Needham, undertook and carried out during the years 1913-1915 the detailed study of a single species in a very restricted area, the results of which are described below. The species chosen for study was Chirotenetes albomanicatus Needham, the nymph of which is common in and characteristic of the riffles and rapids of the streams of the Cayuga basin. The various stages of the insect are described by Needham (05). The description is illustrated by two plates and is accompanied by some data on the habits of the nymph and subimago and the food of the nymph. Morgan (’11) gives some brief observations made on this species in Fall Creek, and again (’13) gives further biological data with illustrations of gills and egg. The writer is greatly indebted to Professor Needham for invaluable advice and for many helpful suggestions freely given during the course of the investigation. THE NYMPH General Habitat.—The nymph of Chirotenetes albomant- catus inhabits all the larger streams in the vicinity of Ithaca, New York, but this study has been mainly confined to one stream, Cascadilla Creek. (Contour map, Fig. 1.) It was chosen chiefly because its smaller size made it much more accessible for observation and experimental work. The 6 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES creek has an approximate length of eight miles. It arises in an upland alder swamp and flows in a westerly direction, emptying into Cayuga Lake Inlet. The stream may be divided into three regions: (1) an upland portion approxi- mately six miles in length where the stream occupies a preglacial drift-filled valley; (2) a gorge portion one mile in length where the stream flows through a gorge which has been cut back in the Devonian shale from the edge of Cayuga valley; (3) a flats portion one mile in length where the stream flows through the sediment deposited at the head of Cayuga Lake by the tributary streams. At its source the stream winds among the alders, flowing over a bed of dark brown vegetable débris, and then, emerging into meadow land, it takes a winding course in the open, its banks bordered with shrubs and scattered trees. The average width is 18 feet. The descent averages 36 feet per mile, producing a moderate current over a bed of gravel and stones, with accumulations of stones and rubble* forming numerous riffles and rapids. Tributaries arising from springs in the surrounding hills increase the volume of the water. The nymphs of Chiro- tenetes occur very abundantly in the riffles and rapids of this upland portion. The creek has a much swifter current in the gorge, spreading out in broad sheets over the smooth rock bottom and tumbling over ledges. Chirotenetes nymphs are found under stones and rubble and occasionally on the rock bottom. After descending into Cayuga valley the stream takes a straight north-westerly course, passing over a bed formed of materials brought down from the heights. These materials form an intergrading series. The larger stones and pieces of rock occur at the base of the hill, and then follow in succession smaller stones and rubble, gravel, sand, silt. For the last 250 yards of its course the stream broadens out and becomes deep and sluggish because of the backing up of the water from the lake. The nymphs of Chirotenetes are abundant at the base of the hill but gradually decrease in *The term rubble is used to designate the flat fragments of shale and lime- stone as distinguished from the stones with rounded edges tending toward a spherical form. — aay, \ ee Kaw » AP a0) ce Ly Z S\ LVS \ topographic sheets of the United States Geological Survey. map from Fig. 1.—Contour CHEMICAL ANALYSES OF THE WATER OF CASCADILLA CREEK All results in parts per million -e__——————————————— EE n—aoer—er———— Date Temp. | Oxygen COs ape = (oe ne bonates 3 1914 Dect at 10° QA. 356 ) * 8 a0 T2e 220 fe) .O1 - 15 3° Tok 1.5 fo) ce 22 ae Lone 2.0 fe) 53.4 1915 Uabee = 1 ae i25 0 On| 2on0 107, “ 19 0° 12nA | 2a O-h5 2 Feb. 16 220 fe) 16.6 n 23 Ten] [ero 2.5 fo) 2255 5\ 3085 Mar. 5 0° 2h) as 0) (0) 17.6 a 19 On 10) (6) 2.5 fe) Bay OIL Apr 13 5° Oe) AO Oe 2l.Onla.OLn 2 20 Ave Wf May 12 14° 9.7 LO) oO 40.6 | .037 S 18 8° Tele Yi) (0) 38.7 June 17 22 8.0 Tent (0) 52.9 | .058 July 14 20 5e | Sey, 0 TEAG Dee a 15 ZO el Oa, 50) |p Water |e aittece: 21 7 9.8 (0) 9.8 | 44.6 af 30 28 ee ORO ROE | iair7elied Ons Aug. 10 19° 8.8 Soy] atisya(e) || ays} 7 .O4 Alb. .08 .O21 a125 O47 .142 118 . 101 .04 Oxygen 05 ° m No) 3.8 Nit- rites .OOI .006 .OOT .OOT OO! Chlo- rine No ° Alka- linity 108 119 120 103 Phos- phates trace trace Sul- phates Tron aie 34.8 33-3 42 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 9 numbers toward the mouth, disappearing when the slack water is reached. Chemical Analyses of the Water.—In order to obtain definite knowledge concerning the medium in which the nymphs live, chemical analyses were made at various times from December 1, 1914, to August 10, 1915. The results are given in tabular form. Comparison with analyses of the water of the neighbouring Fall Creek shows a very close correspondence. The amount of pollution is not excessive as in- dicated by the nitrogen determinations as free and albuminoid ammonia, nitrites, nitrates, oxygen consumed, chlorine and alkalinity. The oxygen content is high throughout the year, Hoot yo Hy Ao BRESSRRRE eee BERD SH a .ae ane SESRSSC areas as Co 8 EEE eee eet ttt eet 20 589° 08) 10. 30) WIG 9 pao Reiss 6) 28°. 13 DEC = TAWIS FEB. MCcH. APR. MAY JUNE JULY AUG. Fig. 2.—Relation of the oxygen content to temperatures of Cascadilla Creek. = oxygen in parts per million at 760 mm. pressure. -- eee e = temperature. doubtless as a result of the turbulent nature of the creek. Figure 5 shows the amounts of oxygen present reduced to 760 mm. pressure as compared with the amounts of saturation at the same temperatures and pressures. The carbon dioxide content is low, probably for the same reason that the oxygen content is high. Temperatures.—Records of the temperatures of the water in Cascadilla Creek were taken almost daily for over a year for the purpose of obtaining information regarding the fluctuation in temperature from day to day, the maximum 10 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES summer temperature, the length of the zero period, and the relation between the temperature and the oxygen content in a swiftly flowing stream. The records show that the water temperature fluctuates with the air temperature but never reaches the extremes of the latter. The highest temperature recorded during this period was 28°C. on June 25, 1914. The maximum air temperature on this day, as recorded by the Weather Bureau branch of the United States Department of Agriculture at Ithaca, New York, was 29.9° C. and the day before 32.2°C. The temperature remained at 0° C. from November 8, 1914, to March 28, 1915, except for.a rise to 3° C. during four days in,February, 1915. The relation of the oxygen content during the year to the temperature of the water is shown in Fig. 2. Relation to Materials Forming the Creek Bed.—Field observations show that the nymphs of Chirotenetes occur in. greatest numbers in those parts of the stream where there are deposits of rubble and stones in moderate to swift current. They occur most abundantly in the upland portion of the creek and so numerous at times are they in places that as many as sixteen nymphs of various sizes have been found beneath a stone of 3 x 4 inches, which would mean a nymph to each three-quarters of a square inch. Forty nymphs have been found beneath a piece of rock about one foot square. Frequently a flat shelving piece of rock may be lifted slightly and the nymphs observed clinging to the under surface in company with other may-fly nymphs such as Epeorus, Heptagenia, Ecdyurus and Baetis, stonefly nymphs, caddis worms and the water-penny. Needham (’05) reports some observations which he made in Fall Creek gorge. “I have observed the nymph, especially in those places where the creek bed is flat shelving rock over which the water streams in a thin sheet. In such places the flat, rocky floor of the stream is covered with a thin filmy growth of algae, with abundant nets of the caddis seine-maker, Hydropsyche; and the broken edges of the floor ledges are fringed with black masses of blackfly larvae, Simulium. Simulium and Hydro- CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES II psyche are fixed in their places, but Chirotenetes wanders about freely over the ledges, clinging securely even in the swiftest water, keeping of necessity head up stream, moving by short quick dashes effected by sharp strokes of its powerful tail fin and gill covers, moved synchronously. It is found in the stiller pools at the sides of the current, in which dwell other may-flies of the genera Caenis and Baetis; and also among the rocks in the current under which cling other nymphs of Heptagenia, Blasturus and Choroter pes.” Field observations appeared to indicate that the lower sides of stones and rubble contribute the preferred habitat of Chirotenetes. In order to test the matter, however, certain experiments were devised. These were based on the assump- tion that the nymphs would not remain in a situation which did not suit them if a more suitable situation were available. A wooden trough three feet long, one foot wide and ten inches deep was constructed and provided with wire netting at both ends, so as to allow water to flow through freely but prevent- ing anything in the trough from escaping. In the upper end of the trough was put a flat stone a foot square. The small crevices between the stone and the sides of the trough were filled in with fine gravel. Behind the stone for the remaining length of the trough course gravel was placed for a depth just equal to the thickness of the stone. In the middle of the trough there was placed a pile of rubble each piece about 314 inches square. At the lower end was put a pile of small stones, each stone about 1% inches in diameter. Just enough space was left between these groups of materials to allow a wire screen to be pushed down between them. The trough was then placed in the stream and arranged so that there was a current through the trough of 1 to 2%, feet per second according to the depth of the water. Then twenty-four almost mature Chirotenetes nymphs were put in the trough at the lower end. At the end of twenty-four hours the screens were carefully put in position between the groups of materials, the trough taken from the stream, the materials carefully removed and the nymphs in each section counted. Nineteen were found among the small stones, four 12 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES in the rubble, and none on the rock. A series of such experi- ments was tried; the arrangement and results of a typical set are given as follows: I 2 2 Rock Rock Rubble (0) (0) 5 29.4% Rubble Stones Stones 4 17.4% I 4.8% 12 70.6% Stones Rubble Rock 19 82.6% 20 95.2% fo) Ailes al a Dee a Ein BNO 1s Stones Stones Rubble a O oO 237% Rubble Rock Rock 10 77% O fe) Rock Rubble Stones (0) 16 100% 15 100% In all cases the nymphs were put into the trough at the lower end. In conducting the fifth test a blocking of the screen at the lower end of the trough one day caused an almost complete cessation of current, and when the trough was examined it was found that all the nymphs except one had migrated to the upper end, close to the screen where there was a slight movement of water. This appeared to indicate that current was a more important factor than material in the selection. of a habitat. The results of these experiments indicate that the nymphs have a decided aversion to open rock and a slight preference for rubble as against small stones. CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 13 Relation to Light.—It was thought that light might be an important factor in the choice of habitat. Wodsedalek (11) has shown by experiments that the nymphs of Heptagenia interpunctata Say are to a strong degree negatively photo- tactic. An experiment was devised to determine the reactions of Chirotenetes nymphs to light. A trough 51X4X3% inches, with wire screens at the ends, was divided into three equal compartments by means of cross pieces reaching from the upper edge to within an inch of the bottom. A similar cross piece was put at the lower end against the screen. At the upper end was a small Intake intake area where the water entered before flowing into the three remaining compart- ments. The trough was then put out in the stream under a small ledge where a current of Upper water could easily be sent through the trough and where conditions of light were normal. The amount of water was then regulated so that the surface just reached the lower edges Middle of the partitions. The bottom of the trough was rough, offering a foothold for the nymphs. Thirteen Chirotenetes nymphs were put in the lower compartment over which a close-fitting Lower cover was immediately placed. At the end of two and one-half hours not a single nymph had left the compartment. Then the cover Fig. 3. Di Bis: doused eames, Was taken off and placed over the upper com- mettion of “temste partment. Immediately the nymphs began alana to migrate up the trough. In four minutes all but four had disappeared into the upper compartment and seven minutes later the remaining four had disappeared also. Not one ventured beyond into the small open intake area. After waiting ten minutes the cover was removed and all the nymphs were driven back into the lower compartment again and the cover replaced. At the end of ten minutes all the nymphs were still in this part. Then the cover was removed and quickly placed over the middle compartment. At once the nymphs began to migrate and in two minutes all 14. CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES except one had disappeared into the darkened chamber, the last one following in forty-five seconds. The trough was left in this condition for seventeen hours and at the end of this time every nymph except one still remained in the middle darkened compartment. When the cover was removed the nymphs scattered. The cover was placed over the lower compartment again and when the trough was examined six and a half hours later all the nymphs were down in the covered area. These experiments were repeated many times. The nymphs thus show very strong negatively phototactic tendencies, The trough used in the experiment for habitat preference was then used to determine whether or not the nymphs would remain on the bottom of the trough if the empty area were darkened, in preference to moving where the stones and rubble were. Since the nymphs appeared to show no very decided preference as between stones and pieces of rock, the trough was divided into two areas only, one with stones and pieces of rock, the other without any materials, leaving the rough bottom of the trough as a surface to which the nymphs might cling. The empty portion of the trough was closely covered and the trough put out in the current, with the stones and rubble up stream. Eighteen Chirotenetes nymphs were put into the lower darkened compartment. When the trough was examined twenty-four hours later, all the nymphs were found in the upper area. The trough was then reversed and the nymphs put in at the lower end among the stones and rubble. Twenty-four hours later all the nymphs were still in this area. Judging from the results of these experiments and field observations it appears that current is the more important factor in determining the habitat of the nymph with light and materials as secondary but closely linked factors. Relation to Current.—Since the nymphs of Chirotenetes are current dwellers it appeared to be desirable to obtain more accurate data in regard to the velocity of the water in which the nymphs live. For use in a stream such as Cascadilla Creek where the water is turbulent, swift, and comparatively CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 15 shallow, it was necessary to have some instrument for current measurement which could be used in narrow places and give measurements at slight differences in vertical and horizontal ranges. Upon consultation with Professor E. W. Schoder of the Department of Hydraulics, Cornell University, a Pitot tube was suggested and an apparatus as shown in Figure 4 was constructed. This consists of two copper tubes 24 inches long and %¥/,, inch in diameter, fastened together and having the lower ends bent at right angles so that the open- ings extend in opposite directions. The copper tubes are connected by means of rubber tubing sixty inches in length and a quarter inch in diameter to two glass tubes each twenty-four inches in length and *5,, inch in diameter. The glass tubes are connected at the top and may be opened to the air by means of a stopcock. The glass tubes are attached to a board and between them is placed a scale. This instru- ment was rated in the canal of the Hydraulic Laboratory of Cornell University. With the Pitot tube a large number of measurements were made in Cascadilla Creek. The first measurement taken was in the middle of a small stream twenty-two inches wide and five inches deep flowing in a channel in the rock in the gorge. It was found that near the surface there was a velocity of 1.7 feet per second, while on the bottom the velocity was 1.0 foot per second. A stone with dimensions of about 12X10X2% inches was placed in the middle of the channel. Midway between the surface of the stone and the surface of the water the velocity was 1.9 feet per second. Fig. 4.—Diasram of On the surface of the stone the velocity was sae ee of encwith 1.5 feet per second; three-quarters of an inch eee ag “'b behind the stone and one inch above the bottom a ae there was no perceptible current; on the stream bottom behind the stone there was a slight current of not over 0.5 foot per second. A measurement in swifter water 16 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES showed a velocity near the surface of 4.3 feet per second while on the bottom only 2.1 feet per second. Many measurements were taken round stones in the creek, particu- larly round shelving stones, and a few results are here given. +b a 2& nn At a velocity 1.6 to 2.1 feet per second. 4c b e 2.0 sé 2 6 sé sé ae i, - very low and indications of slight eddy. ‘C c At a velocity 2.9 feet per second. sé b se 2.6 sé oe “é c “é 3 6 4c sé sé 4c d sé O sé “é ims . L 7 Size of stone = 12 X22 inches. At a velocity 1.0 to 1.5 foot per second. sé 44 ‘6 6c sé AGG xe) 3.5 by b&b dw mH © (SS) oe.o © CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 17 f velocity 2.8 foot per secon d. Ser: Cees “ee : a7 i 4.6 j % 1.0 k a too ¥mall to be measured with the Pitot tube. In a channel 7% inches deep and three feet wide, the bottom velocity was 1.5 feet per second; close to the surface the ‘velocity was 2.5 feet per second. A stone almost cubical in form was put in this channel. “ec “é “é d.« At a on top of stone velocity 2.9 ft. per second. ‘“ pb behind stone on bottom velocity 0.5 ft. per second. ‘““ ¢ behind stone 2% inches above bottom velocity .o ft. per second. “ d at side of stone velocity 2.8 to 3.1 ft. per second. xb a a ; Cx At a velocity 2.9 ft. per second. oe 3 3.0 to 3.4 ft. per second. te - 1.5 ft. per second to a reverse of 2.0 feet per second. There was a strong eddy behind this stone at times and the direction could be determined by turning the copper tubes. Numerous measurements taken round stones and rubble in the rapids and riffles of the stream have shown that nearly all are so placed that a current of greater or less velocity 18 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES flows beneath them. It is in this diminished current under- neath the stones and rubble that the nymphs of Chzrotenetes live. Many of the large shelving pieces of rock have a current underneath at one point and no current at another point. The distribution of seine-making caddis worms underneath a stone gives a very good indication as to the presence or absence of a current. Measurements were taken on July 1, 1915, over the smooth flat rock in the gorge where the water spreads out in a broad sheet and where blackfly larvae and seine-making caddis worms were very abundant. Two such measuremertts are typical. Water Three Inches Deep Close to surface — — 5.1 foot per second. Bottom Sm Sh ae ae ae Rt Water Four Inches Deep Close to surface — — 3.4 foot per second. Midway to bettom =" "= 9 3y0 7) ae ey Bottom = a = aii ee CL sf i At various times of high water in the creek measure- ments of the vertical distributions of velocities were taken. The results of three measurements are as follows: Inches below Velocity Velocity Velocity surface ft. per sec. ft. per sec. ft. per sec. 2 4.5 4-5 3.9 3 4.2 4.4. 3-7 a 4.0 Aes 36 5 3.8 4.1 3-3 6 207 3.8 3.0 7 3-3 2.9 8 yao | eat 28 9 3.0 10 3.4 2.9 27 Ua 12 Ree 2.4 2.6 13 14 3.0 2.4 25 15 2.9 22 16 De 2\92 1.7 | Bottom 17 hh ee .7 } Bottom 1.7 |} Bottom CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 19 On July 13, 1915, the smooth rock of the floor of the stream in the gorge was covered with a thin film of diatoma- ceous ooze, of Navicula and Synedra chiefly, just enough to make the rocks very slippery under foot. The water over the rock had a depth of one to two inches. The Pitot tube was placed on the bottom in the centre of an area of diatomaceous ooze. The velocity was 2.0 feet per second. The ooze was then cleaned off the rock over a large area and the tube placed in exactly the same position as for the first measurement and a velocity of 2.3 feet per second was obtained. There was thus a decrease in velocity of 13 per cent. due to the ooze. Numer- ous other measurements showed an equal or slightly smaller decrease. On rocks with a fine coating of algal growth and silt, measurements showed losses of as much as thirty per cent. Experiments were then conducted to determine in how swift a current the nymph of Chirotenetes could maintain itself. A wooden trough fifty-one inches long, four inches wide, and three and one-eighth inches deep was placed at the edge of a small waterfall so that water would flow through the trough at a depth of about two inches and arranged so that by raising or lowering the lower end the velocity of the water could be varied. The bottom of the trough was a rough unplaned board which supplied the nymphs with a foothold. Three almost mature Chirotenetes nymphs averaging 12 mm. in length were put in the trough about a third of the distance from the lower end where the velocity on the bottom was 1.4 feet per second. The velocity was gradually increased until all the nymphs let go their hold and the velocity of the water then measured. The greatest velocity which these nymphs could withstand was one where the velocity on the bottom of ‘the trough was 4.3 feet per second. With three nymphs 9 to 10mm. in length it was found that these could maintain their hold until a velocity of 4.8 feet per second was reached. A series of experiments was then carried out in order to observe the actions of the nymphs of Chirotenetes more closely in currents of various velocities and to compare their actions with those of the larvae and nymphs of other insects inhabit- 20 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES ing swift water. The procedure was as follows. The trough was arranged with a slight current and the forms to be experimented with were placed in the trough about a third of the distance from the lower end. The current was then increased, and the forms were observed and notes made as | to their actions. The velocity of the water was then measured | on the bottom in the middle of the trough and just off the bottom. The Chirotenetes nymphs used were of almost | mature form and fresh specimens were taken frequently so that the results might not be vitiated by fatigue. Nav 4 Cus Be Pex 2. idpco icy Expy Exp. 5 Bap. 6, + Bottom velocity .4 to .8 feet per second. Off bettony, =). 2.00 eee A Chirotenetes nymphs able to swim and crawl rapidly. Bottom velocity .9 to 1.0 feet per second. Off bottom PD POA Chirotenetes nymphs able to crawl but barely | able to hold their own against the current in swimming. Bottom velocity 1.4 to 1.8 feet per second. Off bottom BS | i Sasa Ne ae Chirotenetes nymphs able to crawl but carried back at once by the current when loosened — from foothold. Bottom velocity 1.0 to 1.9 feet per second. Off bottom 2.1.) 2 Ot aaa i Chirotenetes nymphs still able to crawl rapidly. Bottom velocity 2.2 to 2.4 feet per second. Off bottom 2. Qi a, a Chirotenetes nymphs able to crawl slowly. Bottom velocity 2.8 to 3.6 feet per second. Off bottom Ad 2 4 ‘ Chirotenetes nymphs able to, crawl slowly. Epeorus and Heptagenia may-fly nymphs, a water-penny, a large stone-fly nymph and a black-fly larva were all able to maintain their | CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 21 hold and move about. The large stone-fly walked up against the current with apparent ease. Exp. 7. Bottom velocity 5.5 to 5.7 feet per second. May-fly nymphs (Epeorus, Heptagenia and Chirotenetes), a small stone-fly nymph, a water-penny, a seine-making caddis worm, a black-fly larva and a fish-fly larva (Chauliodes) were put in the trough and the velocity gradually increased until the stated amount obtained. The stone-fly nymph walked up against the current with apparent ease; Epeorus clung securely but remained quiet; the black-fly larva maintained its hold but curled up and lay flat on bottom; the water-penny moved slowly backward; all the others lost their holds before this velocity was reached and did so in this order—Chirotenetes, Hepta- genta, fish-fly larva, caddis worm. The results of these experiments show that the nymphs of Chirotenetes clinging to the under surface of stones escape the main force of the current. The water in such places is moving very much less rapidly than nearer the surface, but it still brings constant and fresh supplies of food and oxygen to the nymph which may wander about in comparative security from many of the dangers which necessarily accompany life in swift water. When the nymph wanders out on the flat rock bed of the stream, it still is in a much reduced current, especially where the rock is covered with diatomaceous ooze or other algal growths. The results also show that the nymph is able to live in rather swift water but that it is scarcely so well equipped for a swift-water habitat as some of its associates with limpet-like forms of body. Neverthe- less the nymph of Chirotenetes does possess a form of body adapted to life in flowing water. The hard smooth chitinous covering reduces the friction of the water particles to a minimum. The head is well rounded. The thorax gradually widens and is followed by a depressed abdomen which 22 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES condition tends toward a limpet or Heptagenine form. The following experiments were devised to determine the mechan- ical or adaptive value of the Chirotenetes form for life in running water. A mass of grafting wax weighing 184 grams was moulded into the shape of a cone, the base of which was 5.7 cm. in diameter and the perpendicular 8.25 cm. A fine wire was put through the cone from the apex to the centre of the base. This wire was fastened to a small metal bar in the middle of the cone to prevent it from pulling out. The cone was attached to a 50-gram spring balance by means of a fine wire 33 cm. in length. When placed in a current of 1.65 feet per second the cone sank about 3 cm. below the surface of the water, and with the base upstream exerted a pull of 28 grams. The balance was held nearly horizontal and as close to the surface of the water as possible. There was considerable fluctuation in the amount of pull on the balance as a result of the unevenness of the current and probably to some extent to the imperfect form of the cone. However, it was found that the indicator of the balance remained at a certain point for a greater part of the time and also that this point was approximately the average of the highest and lowest points reached by the indicator. With the apex up stream the pull exerted by the cone, as nearly as could be observed, was 50 grams. The wax was then moulded into the form of a fish of the sunfish type and the pull exerted was 15 grams. A form of the trout type gave a pull of 6 grams with head up stream and 10 grams with tail up stream. A model of a Chirotenetes nymph gave a pull of 9 grams with head up stream and 16 grams with head down stream. In all these experiments the total amount of wax was used. It was found that the kind of edge fashioned at the base of the cone was of great significance. For example in a current of 1.2 feet per second a cone with a sharp edge to the base exerted a pull of 40 grams, whereas with the edges well rounded the pull was decreased to 10 grams. Another series of measurements in a current 1.5 feet per second gave the following results. A cone with a base 7 cm. in diameter with a sharp edge and a perpendicular of 11 cm. CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 23 gave a pull of 50 grams with the base against the current and a pull of 25 grams with the apex to the current. A cylinder 8.5 cm. in length and 5 cm. in diameter gave a pull of 18 grams. With one end pointed and directed toward the current, the pull was 6 grams, while with the blunt end to the current the pull was 17 grams. With both ends pointed forming a spindle the pull was 5 grams. A fish model and a Chirotenetes model each gave a pull of approximately 6 grams. Comparison of the amounts of pull exerted by the various models is open to the objection that the area of greatest cross section was not kept constant throughout. Nevertheless the results demonstrate in a rough manner that the nymph of Chirotenetes possesses a remarkably efficient form of body. This explains in part how it is that Chiroten- etes nymphs are found in association with the flattened limpet- like forms of the stream. Food.—May-flies are herbivores. They feed almost exclusively upon algae, from the minute diatoms to the higher filamentous forms. The nymph of Chirotenetes has developed a very remarkable specialization by means of which it avails itself of the suspended edible plant material carried along by the current. The inner sides of the fore femora, tibiae and tarsi are fringed with long hairs interspersed with shorter fine ones. These are supplemented by copious short hairs on the labrum, and on the maxillary palpi and by longer ones on the labial palpi. When the nymph takes up a position in the current head up stream, the forelegs are held ‘out in front of the head and flexed at the tibio-femoral joint so that the claws are almost contiguous on the surface of the rock. In this position the hairs of the forelegs and mouth parts meet and overlap in such a way as to form a straining apparatus. To demonstrate the use of this strainer a trough 51X4X3% inches was placed below an outlet from a tank and arranged so that a moderate current flowed through the trough at a depth of one and a half inches. A number of nymphs which had had no food for twenty-four hours were put into the trough at the lower end. They soon began to 24 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES crawl forward along the bottom. A mixture of silt, diatoms and other forms of algae was sent down in the current and observations were made by means of a reading glass. The strainers were soon loaded with the material and four nymphs were observed to feed upon it. The elongated fringed labial palpi were extended to sweep in the materials caught, while the maxillary palpi working laterally and the glossae of the labium working vertically pushed the food materials back to the mandibles. The legs were moved at times to bring food within reach of the mouth parts. Any materials caught and not wanted were expelled by moving a leg outward and allowing the current to wash them away. To determine what materials were available in the stream as food for the nymphs a plancton net was put out in the stream at various times throughout the year. The various forms identified are listed and the estimated abun- dance indicated by the numbers I, 2 and 3, indicating few, abundant, and very abundant respectively. A few quantitative determinations were made to ascer- tain the amount of food available for the nymphs. A small wooden trough four inches wide was placed in the creek in such a way that one end projected over the edge of a small waterfall. A current flowed through the trough to a depth of two inches and at an average rate of 4.0 feet per second. A plancton net of no. 12 mesh silk was hung at the lower end of the trough for sixty minutes. The catch was filtered through fine filter paper and then transferred to a measuring cylinder in 80 per cent. alcohol and allowed to stand for twenty- -four hours, and the volume then read. The catch amounted to 14 cc., a considerable proportion of which con- sisted of silt. Of this catch it was estimated that at least 40 per cent. was silt and inedible matter and another 10 per cent. of fine material which would not be caught by the hairs of the nymphal apparatus, which would mean that 7.0 cc. of edible material was taken in sixty minutes by the plancton net. The data at hand accordingly indicate that at a velocity of 4.0 feet per second there is delivered on an area of 5,160 square mm. in sixty minutes 7.0 cc. of food material. Now CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 25 s a re ee Sra) eked hee [hier Whsee hh gee (en. es Bl) el els) al ale] e] 4/2) 2/2] 2 2\2}/2/ 2/8) 8)/o]/a}<| 4/4) 4] < SS SS Se ee a Se SS ee Ss SS es eS Ee Ge Raat See ee aes ae Ship ee I Baier EES ere So) Eup (2 ee Tint |(y2 I I > See BS Fa aN ee Saas ih Srle2” 2 I BEA at eases |) 2 z ey ered! ayy | eee Pil | MRORGOMEISS - 20. cs os oes Qin 1 (Sal (a Dey sd Smesimion.. 2. 72.229 0.655. I I Gomphonema:..: 0... 2... I Tet bead Cabieat I el o7n) Bg I sn a Bewrasigi@ia.......-.-..-. I I Sa ee e |S as oe ARR Sai Scenodesmus............ ee) i a | EAU RARE I MEETS. s 3. Te logy | pemiastrume 25 22.5.-.-- A) IP es se: a) ‘Sie 6) ont see I Prorococcaceae....:.....- ce A | | Cyanophyceae........... cg ce Mee Re ee Se van Cras Mart I | SEE Vaan. 2 MOHERIA. ...2....-...-. 2 I 2 2 Sos | ibe wees fap og Wx Wedorzoninm .))'.)5 1)... 25... | I etn Gchiy.) 2S 2) ca ois «=! 3's 2 I pel I Me. a | 1 x | Microthamnion.......... RR RR 2 | Higher Plant Tissues....| 3 gh a2 (43 giz |a)3 ES EE ee ee I I I PMelophrys... 2)... +... I ae ee I {a0 I 2 ee Peas ee I 0 ee Ai I I SE eS I 1 I Espa 26 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES the plancton basket of a mature nymph is about 8 square mm. in area. If anymph lives ina current of 1.5 feet per second, then in twelve hours there should be delivered on the plancton basket .05 cc. of food material. The capacity of that portion of the alimentary canal from the mouth to the end of the mid- intestine of a mature nymph was then calculated as .0065 cc. This would mean that this portion of the alimentary canal would be almost filled eight times in twelve hours with food material. The results of other catches with calculations as just described are given as follows: Available food per Date 8 sq. mm. in 12 hrs. ‘at Times capacity of 1.5 ft. per sec. alimentary tract IQI5 June 25 .050 8 2 2S) .097 15 July 6 .OAI 6 After heavy rains I .048 7.5 After heavy rains Aug. 12 .066 10.2 These results are estimates but conservative ones. They show that there is a considerable abundance of food material coming down in the stream even when conditions are adverse, that is, following flood time. Doubtless the nymphs avail themselves also of algal growths on the stones to which they cling, so that the plancton catch (including particles of higher plant tissue and some animal material) may be considerably augmented. No calculations were made during the winter but during this season the creek is full of diatoms. All the stones and rocks are covered with a thick brown velvety covering of diatomaceous ooze, so that the food supply should be but little less than in the summer. Examinations of the stomach contents of nymphs were made throughout a year. Determinations were rather difficult because of the fact that the materials were so very finely ground up. Diatoms were found at all times, but were particularly abundant from September to April and included all the forms listed in the plancton catches. Particles of fila- CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 27 mentous algae were recognizable in most of the mounts but were difficult to identify. Cladophora and Mucrothamnion were identified and particles of a blue-green alga, probably Oscillatoria. Of the smaller green forms a few specimens of Closterium and Scenodesmus were found. Fragments of higher plant tissues were common. Of animal remains two rotifers were found in one individual, a small claw or mandible of some undetermined form in another, Simulium fan rays and remains of Chironomid larvae in several and pieces of chitin in a number. In all specimens examined there were considerable quantities of sand particles. Needham (’05) reports that for nine well-grown nymphs taken in Fall Creek (time of year not stated) “plant remains constituted in all cases fully half of the stomach contents.’’ The plant food consisted largely of higher plant tissues but mixed with this were Cyanophyceae, Chlorophyceae and diatoms. Of animal food, four had eaten Simulium larvae and Caents nymphs, seven had eaten nymphs of Ecdyurus maculipennis and one had eaten of a small platode and a young nymph of Chiro- tenetes. Morgan (’13) reports having found epidermis of roots, Zygnema, Gomphonema, may-flies and other insects in more than ten specimens. It appears that the nymph of Chiro- tenetes tends toward an omnivorous diet. No doubt a considerable number of small animals such as small Szmulium and Chironomid larvae and protozoans lodge on the nymph’s basket and are eaten as readily as the plant forms. Relation to the Oxygen and Carbon Dioxide Content of the Water.—Chemical analyses during 1914-1915 showed that the water of Cascadilla Creek was quite normal as regards the dissolved substances during that period. There was no evidence that the amount of pollution was harmful to the organisms of the creek. The effect on the oxygen and carbon dioxide content of a small amount of pollution in a stream flowing several feet per second is probably very slight. Analyses from December 1, 1914, to August 10, 1915, show a comparatively high oxygen content throughout the yearand a low carbon dioxide content—two factors of extreme importance to an aquatic organism. The samples for oxygen determina- 28 CLEMENS: ECOLOGICAL StuDY OF CHIROTENETES tion were taken from the creek in the gorge in bottles with a capacity of 250 cc. The bottle was allowed to stand un- corked for two minutes to allow entrained bubbles of air to escape and determinations were carried out at once. There is a possibility that a small quantity of entrained air still remained in the bottle but the results obtained do not represent the total amount of oxygen available, for the tumbling waters hold a large amount of entrained air which would be of extreme value tothe nymphs. In winter at 0° C. the results show the water to be almost saturated with oxygen. In summer, although the amount per million cubic centimeters has dropped considerably, still the water is frequently supersaturated. (Fig. 5.) The Ohio State Board \s 1 | ae an CORRE EE | WISER ft aaER a wae BERGNs PEEHOURSRRERDERSRERERROE ES SESEEEA UEP SRST OPERATES ERS PES BASRA Mar Gate 20 Swe) 48 |) 40 son 45 18.) Sees 43 Dect SANS FEB MCH = APR. may” JUNE JULY AUG. © Fig. 5.—Oxygen content of Cascadilla Creek compared with amounts of saturation of water at same temperatures and pressure. = oxygen content in parts per milllion. ------ = amounts of saturation in parts per million. of Health ('97) has reported water in Ohio supersaturated with oxygen during August, September and October. Shelford (13) states that the carbon dioxide content is probably the best single index of the suitability of water for fishes because carbon dioxide in excess has a toxic effect. Chemical analy- ses of the water of Cascadilla Creek show a low carbon dioxide content throughout the year, especially in midsummer CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 29 when during July and part of August the water was devoid of this gas. The nymph of Chirotenetes obtains its oxygen supply by means of tufts of tracheal gills. There are seven pairs attached to the posterior lateral margins of the abdominal segments one to seven, one pair attached to the bases of the fore coxae and one pair at the bases of the maxillae. An estimate of the gill area presented by the nymph was made by counting the gill filaments in a cluster and measuring the length and diameter of a single filament. The results show a gill surface of about 230 square mm. presented by a mature nymph. The nymph is thus well equipped for obtaining a good oxygen supply in that it possesses fore coxal and maxillary gill tufts which are of unique occurrence among the known may-fly nymphs of our fauna. The extensive gill equipment probably bears some relation to the active habits of the nymph. The following experiments were conducted to ascertain if the nymphs of Chirotenetes are dependent on current for their oxygen supply. On June 27, 1915, there was put into a large glass aquarium 2,000 cc. of water from Cascadilla Creek, containing 8.3 parts per million of dissolved oxygen and one part per million of free carbon dioxide at 18° C. Five nymphs were introduced into the water and on the bottom of the jar was placed a clean stone to which the nymphs might cling. On July 11 a rain added slightly to the amount of water in the aquarium. On July 13 a nymph died, and analysis of the water on July 14 showed 5.16 parts per million of oxygen and three parts per million of free carbon dioxide at a temperature of 21° C. The remaining nymphs died on July 20, having lived in the aquarium twenty-four days without food. Analysis on July 21 showed 9.9 parts per million of oxygen and no free carbon dioxide at a temper- ature of 15.1°C. Doubtless the nymphs had died of starvation. For purposes of comparison 5,000 cc. of creek water had been put into a large glass aquarium on June 17, 1915, with nonymphs. Analyses of this water were made as follows: 30 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES Carbon Dioxide Temperature Oxygen parts per million per million 1915 June 17 2052-6 8.01 1.5 a ahaa es 24 hours Tee 11H (0) 25 P Hhy, 24 7 days 11.0 9.68 5 July 14 27 days 21.0 7.49 25 eth ee 34 days Toa g.11 .O On July 21 three Chirotenetes, three Heptagenia and three Epeorus nymphs (may-fly nymphs from flowing water of Cascadilla Creek) were put into this jar. The next morning the three Epeorus nymphs were found dead. Of the remain- ing nymphs four were alive on July 30, the other two having transformed to subimagos. On August 16 one Chirotenetes nymph was still alive in the jar, having been able to live for twenty-six days in water which had been standing sixty days. Analyses after the addition of the nymphs gave results as follows: Carbon Dioxide Temperature Oxygen parts per million per million 1915 July 30 43, days 20.0°C 8.42 0 Aug. 10 54 days 17.0 8.00 TS Just before Cascadilla Creek enters the gorge section, a pond, known as Dwyer’s Pond, has been formed by a dam. Analyses of the water from Dwyer’s Pond failed to show very marked differences in oxygen or carbon dioxide content from that of the tumbling water in the creek in the gorge. The samples were taken in the pond just below the surface and close to a clump of sedges in a situation where may-flies would be likely to occur. That Chirotenetes nymphs do not live in the pond cannot be because of lack of oxygen there or excess of carbon dioxide, nor because of lack of current, since the results of the aquarium experiments show that the nymphs CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 31 are not dependent upon current for oxygen supply as are Epeorus nymphs apparently, for the latter die very soon after being placed in still water. Associates.—In the classification of the ecological com- munities of the stream, the nymph of Chirotenetes belongs in the strata under the stones to the Hydropsyche or riffle formation (Shelford, ’13). Its associates in Cascadilla Creek are as follows: | Ephemerida of the genera Heptagenia, Iron, Epeorus, Ecdyurus, Ephemerella, Leptophiebia and Baetis. Plecoptera of the genera Perla, Acroneuria, Neoperla and Pteronarcys. Neuroptera of the genera Corydalis and Chauliodes. Trichoptera of the genera Hydropsyche, Helicopsyche, Rhyacophyla, Leptocerus, Chimarrha and Polycentropus. Lepidoptera of the genus Eliophila. Coleoptera of the genus Psephenus (‘‘water-penny’’). Diptera of the genera Atherix, Chironomus, Diamesa, Tanytarsus, Tabanus, Eriocera and Tipula. Planarians. Hirudtinea. Mollusca of the genus Ancylus. On the smooth rock beds of the gorge, where the nymph of Chirotenetes occurs occasionally, are found larvae of Simulium, Blepharocera and Hydropsyche and Ephemerid nymphs of the genera Heptagenia, Iron, Epeorus and Baetts. Among the stones of the creek have commonly been taken the blacknosed dace (Rhynichthys atronasus), the young of the common sucker (Catostomus commersonit), Johnny darter (Bolesoma nigrum), the nigger chub (Exoglossum maxillingua), the satin-fin minnow (Notropis whipplit), small common shiners (Notropis cornutus) and the dusky salamander (Desmognathus fusca). Enemies.—Stomach examinations of some of the associates of Chirotenetes show that the two chief enemies are the large stone-fly nymphs, particularly Perla media, and the black-nosed dace Rhynichithys atronasus. Morgan (11) 32 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES reports having seen robins in Fall Creek gorge with the setae and abdomens of nymphs and subimagos projecting from the beaks. The period of moulting is an especially helpless time for the nymphs and at such time they are most liable to become the prey of enemies. Severities of Stream Life-—Life in swift water is beset with many difficulties and dangers. Probably the most trying season for the nymphs of Chirotenetes is flood time. Stones and rocks are moved by the force of the current and during the spring flood the out-going ice scrapes and scours the creek bottom. The movement of stones, rocks and ice over the rock bed in the gorge of the stream at a time of high water can be distinctly heard. The nymphs are in danger of being crushed or swept out in the current over waterfalls or into unsuitable situations. A pool about one hundred and fifty feet south of Fall Creek is usually flooded in spring by a portion of the creek being diverted through it. After such a flooding Chirotenetes nymphs have been found in the pool. Another illustration as to how the nymphs are carried about during flood time was afforded in the course of the habitat experiments. A heavy rain one night caused the water in the creek to rise and the trough set out for an experiment was carried about ten yards down ‘stream without being upset. The water was flowing over the trough to some extent and the stones, pieces of rock and gravel in the trough had all been shifted. The trough had contained thirteen almost mature Chirotenetes nymphs, but when it was examined only three of these remained, but twelve small Chirotenetes nymphs 7 to 8 mm. in length were found in the trough besides numerous Baetis nymphs and a number of Hydropsyche larvae. After heavy rains the nymphs have to contend with large quantities of silt. Samples of water have been taken from the creek when it was loaded with sediment, the water filtered and measured and the residue weighed after drying in the air of the laboratory for several days. The results obtained are as follows: CLEMENS: EcoLocicaL Stupy oF CHIROTENETES 33 Date | Amountof | Weight of | Parte, per Conditions 1914 June 28 2200 cc. 13.8 grs. 6200 Taken two hours after very heavy downpour of rain. Aug. 20 2400 1.0 415 Taken morning after a heavy night’s rain. Sept. 2 1900 3.4 | 1800 _—-| Taken three hours after heavy rain. IQI5 Jan: 7 400 Taken at time of a mid- winter thaw. July 5 2200 a3 590 Taken the day after heavy rains. Professor Chamot has stated that turbidities in Six-Mile Creek frequently exceed 6,000 parts per million. Such enormous amounts of sediment result from the hilly nature of the watershed and the nature of the soil. The soil of Cascadilla valley consists of silty, clayey and stony loams. Heavy rains cover the hillsides with rivulets which bring down to the stream immense quantities of sediment. The water becomes yellow-brown in colour and so loaded with silt that the earthy odour can be detected a long distance from the creek particularly in the vicinity of waterfalls. Flood time too carries out to the lake the plancton of the pools and ponds of the stream leaving the creek more or less deficient for some time of those suspended organisms upon which the nymphs depend to a large extent for food. This was shown by plancton catches taken before and after floods. Needham ('16) points out the dangers from ice particles and “anchor ice” during the winter period. Protectton.—The nymphs of Chirotenetes receive protec- tion from the stones, to the lower sides of which they cling. While the stones are in position, protection is afforded from the force of the current, from larger objects carried in the current, and where the space is small from larger enemies such as fish. The nymphs no doubt escape » 34 CLEMENS: ECOLOGICAL Srupy OF CHIROTENETES enemies by reason of protective colouring and agility. The chocolate-brown colour renders the nymph very incon- spicuous on the dark coloured rocks of the stream bed, and the ability to dart quickly from place to place by means of strong strokes of the abdomen and fringed setae is of decided advantage. Nymphs in a current of water in a trough have been observed to loosen their hold allowing the current to carry them down a short distance and then to catch the bottom or side of the trough again. Doubtless such a procedure is used as a means of escape. Regeneration.—The nymphs possess the power to regen- erate certain parts. Frequently it has been observed that a nymph lacking a leg will after moulting possess a small leg. How many instars are passed through before the leg attains normal size has not been determined. The ability to regen- erate a foreleg is of vital importance to the nymph. THE SUBIMAGO Emer gence.— There comes a time in the life of the nymph when some stimulus causes it to crawl up the side of a stone out of the water. Phe stimulus is probably supplied by a number of agencies of which the maturation of the sexual elements is doubtless the chief. The nymph crawls out just above the surface of the water. In a few seconds convulsive movements pass through the body, the head and thorax split along the mid-dorsal line and the body of the subimago slips out on to the stone. As soon as the wings are freed, they are spread, and at the same time the legs are extended so as to support the body of the insect. It soon takes a few steps or a little jump and the abdomen and setae are freed. After a few movements of the wings, legs and setae, the subimago flutters upward into the trees. Where no trees are near by they often flutter upward out of sight. The body of the subimago sometimes slips out on to the surface of the water and is carried down stream some distance standing on the surface film. A wave occasionally submerges the subimago or sweeps it away before it has freed itself from the old nym- CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 35 phal skin, thus ultimately bringing it to its death. Trans- formations occur in greatest numbers during the late after- noon and evening but some occur in the morning. The period of emergence is rather extended. The earliest observed emergence was on June 6 and the latest September 8, with the greatest numbers in June and July. Morgan (’11) reports emergings in great numbers in May, I9gI10. Factors affecting the length of the Subimaginal period.— The subimaginal period is one of quiescence. The subimagos remain quietly on the leaves and twigs of the vegetation bordering the stream. They take no food, the mouth parts being degenerate. This condition prevails normally from twenty-four to thirty-six hours, at the end of which time the subimago moults, and the may-fly takes on the adult form. A set of experiments was carried out to determine the effects of temperature, light and humidity on the length of the subimaginal period. Three bell jars were set up. In one was placed a tray of calcium chloride, in another a tray of water and saturated blotters. The third was used as a control. Hygrometers hung in the jars showed that in the first the relative humidity was about 32 per cent, in the second practically at saturation, and in the third 66 per cent. Subimagos as they emerged were put into small wire cages. A male and a female were put into each cage and the cages put under the bell jars. Two series are given as typical: Jar—low humidity (32%) Q@ emerged 12:25 P.M. Aug. 14, trans. 8:30 P.M. Aug. 15 = 32 hirsi'5 min. o' emerged 12:40 P.M. Aug. 14,—died. Jar—normal humidity (66%) 2 emerged 4:18 p.m. Aug. 14, trans. 9:00 P.M. Aug. I5 = 28 hrs. 40 min. o emerged 4:10 P.M. Aug. 14, trans. 9:00 P.M. Aug. I5 = 28 hrs. 50 min. 36 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES Jar—saturation (100%) 2 emerged 5:20 P.M. Aug. 14, trans. 1:00 A.M. Aug. 16 = 31 hrs. 40 min. co emerged 4:50 P.M. Aug. I4, trans. 10:45 P.M. Aug. 15 = 29 hrs. 55 min. o emerged 4:50 P.M. Aug. 14, trans. 11:15 P.M. Aug. 15 = 30 hrs. 25 min. Jar—low humidity (35%) @ emerged 5:15 P.M. Sept. 10, trans. 6:40 P.M. Sept. 12 = 49 hrs. 25 min. o emerged 5:15 P.M. Sept. 10, trans. 6:30 P.M. Sept. 12 = 49 hrs. 15 min. Jar—normal humidity (50%) 9 emerged 4:15 P.M. Sept. 10, trans. 4:10 P.M. Sept. 12 = 47 hrs. 55 min. 9 emerged 3:55 P.M. Sept. 10, trans. 4:50 P.M. Sept. 12 = 48 hrs. 55 min. Jar—saturation (95%) 9 emerged 4:25 P.M. Sept. 10, trans. 4:45 P.M. Sept. 12 = 48 hrs. 20 min. o' emerged 5:05 P.M. Sept. I0, trans. 5:05 P.M. Sept. I2 =48 hrs. 0 min. Other series brought out the same results, namely that individual variations were greater than the variations among the jars. Experiments were then conducted to determine the effect of darkness on the length of the subimaginal period. The exact time of emergence was obtained. The subimagos were transferred to wire cages some of which were put in a photographic dark-room and others on a window-sill in the laboratory in the bright light, at times partly in the sunlight. CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 37 Cage in light. o& emerged 4:10 P.M. Aug. 14, trans. 9:50 P.M. Aug. 15 = 29 hrs. 40 min. 2 emerged 4:45 P.M. Aug. 14, trans. 1:15 A.M. Aug. 16 = 29 hrs!) 20 cham Cage in dark room. o emerged 4:55 P.M. Aug. 14, trans. 12:20 P.M. Aug. 15 = 31 hrs. 25 min. 9 emerged 5:20 P.M. Aug. 14, trans. 12:30 P.M. Aug. 15 = 31 hrs. 10 min. Other experiments gave similar results, showing that darkness has no effect on the subimaginal period. During the time these experiments were being conducted, the temperature varied considerably and a decided lengthen- ing of the subimaginal period was noted when the temperature of the air lowered and a shortening of the period as the temperature rose. For example, on September 1 and 2, 1914, the temperature rose to 28.3° C. and the subimaginal period lasted 22 to 25 hours. On September 11, 1914, the tempera- ture dropped to 15.5° C. and the period lasted 48 to 49 hours. A difference of 12.8° C. had doubled the length of the subimaginal life. A number of subimagos were placed in a cage which was then placed in the ice box of an ordinary refrigerator where the temperature wes 8° C. The subimagos lived four days before transforming and some failed to transform. The length of the period thus varies greatly with the temperature but not with humidity or light. Enemtes.—The chief enemies of the subimagos are birds and spiders. Birds have been observed to fly out of the trees bordering the creek and catch the subimagos fluttering up- ward after emerging. All tree-inhabiting insect-eating birds doubtless feed upon the subimagos. Many subimagos are caught in spider webs. A fence along Fall Creek near Forest Home village usually has a great number of spider webs filling its spaces. One afternoon a large number of subimagos were emerging from the creek and a strong wind carried them 38 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES toward the fence. Every web along the fence had one or more Chirotenetes subimagos caught in the meshes. Heavy rains and winds are destructive to the subimagos, beating them down and wetting the wings so that transformation cannot be successfully carried out. Protection.—The only protection for the subimago con- sists in its dull colour and its quiescent habits. A subimago on the vegetation is usually difficult to detect. THE IMAGO Transformation.—At the subimaginal moult, the head and thorax of the subimago split along the mid-dorsal line, the wings are spread almost horizontally and with a few contractions of the body the adult form appears. When transformation occurs on a vertical surface, the body of the imago is bent down backward until the wings and legs are freed and then by movements of the wings the body is brought up until the legs are able to grasp the object above. The adult then walks away from the moulted skin. Flight——The males of Chirotenetes have a flight charac- teristic of the majority of may-flies. They appear over the stream usually after sunset, about twenty minutes before nightfall, in small swarms of thirty to fifty individuals. They are very graceful in flight, rising and falling in deep undulations of eight to fifteen feet, fluttering upward with — the body held obliquely and then falling slowly on expanded wings and spread setae, with the body horizontal. The — females do not take part in these flights but fly singly up and | down the creek in long undulations. What factors induce | the imagos to fly in the late evening have not been determined. | It may be that this time of day is the safest, having been — determined by natural selection, or it may be a negatively. phototactic tendency carried over from the nymphal stage. | When a female enters a swarm of males she is quickly caught - by a male flying up beneath her. The male places his long | forelegs over her prothorax and head and grasps her abdomen | with his forceps. The arching of the body of the male in order to grasp the body of the female with the forceps brings | CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 39 the penes in position to be inserted into the openings of the oviducts at the apex of the seventh abdominal segment. Copulation lasts twenty-five to sixty seconds. The couples do not rise high but remain at almost constant level, frequently making quick turns and occasionally sudden drops. The male in separating lets go his hold with the forelegs first and finally with the forceps. Apparently the male returns to the swarm while the female flies up and down the stream in long undulations and soon begins ovipositing. The mouth parts of the imago are degenerate and no food is taken during the short aerial life. The alimentary canal, however, is not degenerate but is filled with air and serves as a buoyant organ (Sternfeld, ’07). Ovtposition.—Preceding oviposition the eggs make their appearance from the openings of the oviducts and form a spherical mass which is apparently held in position by the bending forward of the sternal prolongation of the ninth abdominal segment. The eggs are held together by very fine strands of a viscid substance. The female flies over the water with long deep undulations carrying the greenish egg mass and dips to the flowing water so that the egg mass is carried away in the current. The eggs scatter somewhat in the water and adhere to objects by means of the viscid strands. The Egg.—The egg is almost spherical with a diameter of .2mm. It is greenish in colour when mature, with the surface divided into very small polygonal areas and slightly roughened. The egg complement consists of 1,900 to 2,000 eggs. It was found that at a temperature of 22.5 to 25° C. they hatch in about fourteen days, while at a temperature of 13° C. in twenty-five days. In the eggs ready to hatch the eyes and ocelli of the embryos can be seen moving up and down. After a considerable period of movement of the head in this way, a crescentric slit appears on the egg shell at the point where the head has been moving, and then the head pushes out through the opening. Soon the tips of the antennae are freed and extended. The pairs of legs follow —E in succession accompanied by considerable movement of the 40 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES body. At the end of ten minutes all the body is freed except the tip of the abdomen and the setae; and in two minutes more the nymph is able to shake off the egg shell. The nymphs at hatching are .8 to .g mm. in length, without gills, with forelegs unfringed and having eyes and ocelli of equal size. They moult six days after hatching and still show no signs of gills but the forelegs possess a few hairs along the inner margins. It was found that the eggs of Chirotenetes could be artificially fertilized. The testes of three males were put into water in a Syracuse watch glass and teased out. The eggs of a female were then put in the vessel and stirred about gently for a few minutes. The contents of the watch glass were left standing for an hour, then the water was poured off with the bits of tissue resulting from the dissections. Fresh water was poured over the eggs, and the watch glass covered and kept on the laboratory table. The eggs began to hatch eleven days later and continued to do so for nine days. Length of Life—How long the adults live under normal conditions and whether males return to swarm a second or | third evening has not been determined. Reared imagos kept out of doors in a large wire cage in which was put a leafy branch of a tree, lived four and one-half days. SUMMARY The results of the investigation are as follows: 1. The nymphs of Chirotenetes show a very decided © habitat preference for the lower surfaces of stones and rubble © as against smooth open sheets of rock, and a slight preference for rubble as against small stones. 2. The nymphs are negatively phototactic. 3. The nymphs live in a very much diminished current beneath the stones and rubble of the stream. 4. The nymphal form of body is well adapted for a more or less active life in running water. 5. By means of fringes of hairs and bristles on the forelegs the nymphs are able to strain out suspended organic | materials of the stream for food purposes. | ‘ * CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 41 6. The suspended plant and animal forms in the current are sufficient in amount to supply the food requirements of the nymph. 7. The water of Cascadilla Creek throughout nine months was neither excessively polluted nor contained excessive amounts of dissolved substances, so as to be harmful to the nymph. The oxygen content was high and the free carbon dioxide content low. 8. The nymph is not dependent on current for oxygen supply. g. Temperature hasa very marked effect on the length of the subimaginal period, while humidity and light have no effect. 10. The eggs of Chirotenetes can be fertilized artificially. It appears that food has been the factor determining the habitat of the nymph. With special equipments in bodily structure it has pushed out into the current and made use of the current to bring it food. Out in the swift water it has taken to the lower surfaces of the stones and rubble for shelter from the dangers accompanying life in swift water and for protection from enemies. While the activities of the nymph are concerned primarily with the acquisition of food, the activities of the adult are concerned with reproduction. The degeneration of the mouth parts, the inflation of the alimentary tract, the well developed wings, the elongated forelegs and large compound eyes of the male are modifications tending to insure the perpetuation of the race. 42 CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES LITERATURE CITED Morgan, Anna Haven. 1911. May-flies of Fall Creek. Ann. Ent. Soc. Amer., vol. IV, pp. 93-126. 1913. A Contribution to the Biology of May-flies. Ann. Ent. Soc. Amer., vol. VI, pp. 371-413. Needham, James G. 1905. May-fliesand Midges. N.Y.State Museum Bull. 86, pp. 17-36. Needham, James G., and Lloyd, John T. 1916. The Life of Inland Waters. Ithaca, N.Y. Ohio State Board of Health. 1897. Preliminary Report of an Investigation of Rivers and Deep Ground Waters of Ohio as Sources of Public Water Supplies. Cleveland, Ohio. Shelford, Victor E., and Allee, W. C. 1913. The Reactions of Fishes to Gradients of Dissolved Atmospheric Gases. Jour. Exp. Zool., vol. xiv, Pp. 207-266. 1913. Animal Communities in Temperate America as Illustrated in the Chicago Region. A Study in Animal Ecology. Geog. Soc. Chicago. Bull. 5. Sternfeld, Richard. 1907. Die Verkummerung der Mundteile und der Funktionswechsel des Darms bei den Ephemer- iden. Zool. Jahrb. Jena, Abt. fiir ae und Ent., vol. XXIV, pp. 415-430. Wodsedalek, J. E. 1911. Phototactic Reactions and their Reversal in the May-fly Nymphs Heptagenta interpunctata. Biol. Bull. vol, Xx, pp. 265-2758; CLEMENS: ECOLOGICAL STUDY OF CHIROTENETES 43 EXPLANATION OF PLATES PLATE I Fic. 1. Nymph of Chirotenetes albomanicatus Needham. 2. Pitot-tube as used in Cascadilla Creek. PEATE ib Upland portion, Cascadilla Creek. Upland portion, Cascadilla Creek. ae. oa PLATE III 5. Gorge portion, Cascadilla Creek. 6. Gorge portion, Cascadilla Creek. PEATEs TV 7. Cascadilla Creek after descent into Cayuga Valley. 8. Flats portion, Cascadilla Creek. Pears Vv g. Winter conditions, upland portion, Cascadilla Creek. 10. Flood conditions, upland portion, Cascadilla Creek. \ 3 *s ‘ ® ' ‘ i | f , * . eek. as ely b P TRGeR hone a, ; j A, ae Ry 7 , Aa Ri oe. i ie a NG Coe i y bt i ' * Hi 4 ‘ hy Y, AN rm p Le ae Saat i ae y a as * ” - * 6 v a 9“ 4, * f % ‘i > . 1 PLATE I Fig. 1—Nymph of Chirotenetes albomanicatus Needham. Fig. 2.—Pitot-tube as used in Cascadilla Creek - ~ i i ¥ ai é 4 “ : ; a ia ‘ PLATE II Fig. 4.—Upland portion, Cascadilla Creek PLATE III Fig. 6.—Gorge portion, Cascadilla Creek. PLATE IV Fig. 7.—Cascadilla Creek after descent into Cayuga Valley. Fig. 8.—Flats portion, Cascadilla Creek. PLATE V Fig. 9 —Winter conditions, upland portion, Cascadilla Creek. Fig. 10.—Flood conditions, upland portion, Cascadilla Creek. A. PU BY N. BRARY: PUBLISHED BY .* CAL SER -ISOPODA OF THE BAY OF FUNDY PRoFEssor W. J. ALEXANDER, Pu. De ProFEssor J. Ai MACKENzIE, B. Lae me. PRoressor J. P. McMurricu, Pa. Dy, ; Bric.-GEN. G.-H: MITCHELL, B. A. Se, fe a PROFESSOR G. H. NEEDLER, Pu. DL , PROFESSOR GrorceE M. Wrone, M.A. . General Editor: H. H. Laneton, M.A. Librarian of the THE ISOPODA OF THE BAY OF FUNDY BY N. A. WALLACE, B.A. (PREPARED FOR PUBLICATION BY A. G. HUNTSMAN, BIOLOGIST TO THE BIOLOGICAL BOARD OF CANADA) FOREWORD During the summers of 1912 and 1913 Mr. Norbert A. Wallace worked at the Atlantic Biological Station, St. An- drews, New Brunswick. At the beginning he spent some time studying the Argulidae and in addition determined the food of a number of fishes. Latterly he spent his whole time in investigating the Isopod fauna of the region, the dredgings which were being made during those seasons throughout the Western Archipelago and at St. Mary bay, Nova Scotia, furnishing him with the material for his investigations. He was most indefatigable and painstaking in this work, which he continued to some extent during the succeeding winters at the Biological Department, University of Toronto. His untimely death on December 3rd, 1914, cut short a brilliant career. Unfortunately he had only reached the stage where he was about to prepare the manuscript of his report, which was, therefore, in an incomplete state. His drawings of the species were not all completed, and he had made only preliminary drafts of the descriptions and accounts of distribution. It has been necessary for us to collate these and at the same time eliminate, add, or reconstruct freely. Where feasible his accounts are used verbatim. He had neither named the new species nor examined all the literature to make certain that they had not already been named and described. We have not been able to refer them to any of the described species and, therefore, believe them to be new. No attempt has been made, however, to examine critically the species of the genera to which they belong so as to determine their relation- ships with the described species. If Mr. Wallace had lived, he would undoubtedly have rendered this account much more complete. We hope that no serious mistakes have been made in its final preparation. The study embodied in this paper was carried on with the assistance of the Biological Board of Canada and of the Biological Department of the University of Toronto. 1a Oe» i 4 a ay fe ISOPODA,.OF THE BAY. OF FUNDY INTRODUCTION In 1853 Stimpson reported in his ‘‘Synopsis of the Marine Invertebrata of Grand Manan”’ a total of nine species of Isopods from that island. These are in current _nomenclature Leptochelra filum, Gnathia cerina, Calathura branchiata, Cirolana polita, Chiridotea tuftsit, Idothea balitica, Edotea triloba, Jaera marina, and Jantra alta. Harger in his ‘‘Report on the Marine Isopoda’’, pub- lished in 1880, added six species to the fauna of the Bay of Fundy, which are - Ptianthura tenuis, Limnoria lignorum, Idothea phosphorea, Munna fabric, Munnopsts typica, and Bopyroides hippolytes. Mr. Wallace now adds twelve species, of which five are new to America although known from Europe, and three are new to science. Those merely new to the Bay of Fundy are Aega psora, Chiridotea caeca, Synidotea nodulosa, and Phryxus abdominals. 6 WALLACE: THE ISOPODA OF THE BAY OF FUNDY Those new to the American coast are Typhlotanats aequiremts, Pleurogonium rubicundum, Pleurogonium inerme, Pleurogonium spinosissimum, and Eurycope mutica. The species new to science are Typhlotanais mananensis, Leptognathia (?) psammophila, and Leptochelia profunda. It is significant that the species new to America are all very small and that those new to science are not only small, but belong to the difficult group of the Tanaidacea. Future additions to the list of Isopods will doubtless be in this same direction, namely, that of the minute forms, including parasites. ; In addition to the marine forms, Mr. Wallace has listed three species of terrestrial Isopoda from the shores of the Bay of Fundy. They are all well known species, but their distribution in eastern Canada has not been signalized. With these the list of species from the Bay of Fundy has reached the number of thirty. 2 CAN ey WALLACE: THE ISOPODA OF THE BAY OF .FUNDY 7 ACCOUNT OF THE SPECIES Family Tanaidae Typhlotanais aequiremts (Lilljeborg). Figure I. Sars, 1899, p. 21. The single specimen obtained—a female—was taken off Big Duck Island, Grand Manan, at a depth of 55 fathoms on muddy bottom. The body is elongated and very slender, slightly over seven times as long as broad, and much depressed. The cephalosome is slightly longer than it is broad and the anterior margin slightly ex- cavate on either side of a blunt median point. The first pair of antennae have the first article A longer than the other two com- bined. Thesecond pair of antennae are similar to those of the next species. The thorax is cylindrical in shape, having the first segment c the shortest, the second, third, fourth, and fifth subequal, and the sixth slightly shorter than the fifth, but longer than the first. Pick Tuiaonts eam a thee chee af the next x 240; (C), first antenna, x 240. species, while the other legs are much as in that species, the three posterior pairs having the basal article expanded and stout. The abdomen is composed of six segments, the last one being rounded posteriorly. The uropods are biramous, the peduncle of each is quite stout, and the two branches are 8 WALLACE: THE ISOPODA OF THE BAY OF FUNDY approximately equal in length, the outer being biarticulate and ending in a strong bristle, while the inner is uniarticulate, but gives an indication of subdivision in having a hair at about its middle. Typlotanais mananenstis, sp.n. Figure 2. A single female of this species was dredged outside Big Duck Island, Grand Manan, at a depth of 42 fathoms on a muddy bottom. The body is elongated and very slender, being nearly nine times as long as broad (about 1.8 mm.: 0.23 mam)./\) lhe head is about twice as long as broad and narrower in front. The anterior margin is produced in the mid-line to form a short, sharp rostrum. The first pair of antennae are short, conical, and very stout at their base. Each consists of three articles of which the first is very stout and a little longer | than one half of the whole antenna, the second is less than a quarter as long as the first and much narrower, and the third is one and one-half times as long as the second and narrower. The second pair of antennae are more slender than the first and are composed each of six articles, of which the first is very short and slender, the second somewhat dilated and a little longer than the first, the third a little longer and narrower than the second, the fourth twice as long as the third, the fifth a little over one-half the length of the fourth, and the sixth very short and almost inconspicuous. The first thoracic segment is united to the head, while the remaining segments are free. The second is the shortest, the third, fourth, fifth and sixth are subequal in length and about twice as long as the second, and the seventh is shorter than the sixth. The abdomen consists of six free segments, of which the first five are subequal in length, while the sixth ise) ttle longer than the others, approximately quadrilateral in shape, and very obtusely angular behind. The uropods are biramous, the peduncle consisting of a single, stout article, the outer branch of two subequal articles, WALLACE: THE ISOPODA OF THE BAY oF FUNDY 9 ™~.- Fic. 2. Typhlotanais mananensis, sp.n.; (A), first antenna, x 120; (B), second antenna, x 120; (C), gnathopod, x 120; (D), female, dorsal view, x 30; (E), second pleopod, x 120; (F), fifth pleopod, x 120; (G), seventh leg, x 240; (H), terminal segment, x 120; (I), uropod x 120; (J), mandible, x 480; (K), maxilla, x 480; (L), maxilliped, x 480. 10 WALLACE: THE ISOPODA OF THE BAY OF FUNDY and the inner branch of two articles, of which the first is three times as long as either of those of the outer branch, and the second one-half as long as the first. The gnathopods are very slender. The propodus is very poorly armed, the inner surface being only slightly roughened and having one short spine at the tip. The articles of the legs are all slender, except that the basal ones of the fifth, sixth and seventh legs are expanded and quite stout. Of the five pairs of well developed, biramous pleopoda, the first four are alike, while the fifth differs in that the inner edge of the inner ramus is slightly emarginate near the distal end. The mouth parts are normal. Leptognathia (?) psammophila sp.n. Figure 3. Females only of this species were found and in sand at two localities—near West Quoddy at a depth of nine fathoms, and at Woodward’s cove, Grand Manan, from low tide mark to a depth of two fathoms. The body is elongate and narrow, being more than six times as long as broad (about 3 mm.: 0.48 mm.). The head is longer than it is broad and wider behind than in front. The anterior margin of the head is produced medially into a very short, rounded, blunt rostrum, on each side of which is a shallow, angular excavation which lodges the peduncle of the first antenna. The latter, which is directed forwards, is com- posed of three articles, of which the first is long and stout, the second narrower than and only one-third as long as the first, and the third as long as the second. The second antenna is not so long as the first, and is composed of six articles. Of these the first is short and stout, the second one-third longer than the first, the third equal to the first, the fourth as long as the first three articles together and usually curved, the fifth one-fifth the length of the fourth, and the sixth very small and inconspicuous. Of the thoracic segments, the second (the first free one) is the shortest, the third, fourth, fifth and sixth are all sub- equal and each one and one-half times as long as the second, WALLACE: THE ISOPODA OF THE BAY OF FUNDY HII and the seventh approximately equal in length to the second. The thorax narrows to the seventh segment. The sixth segment of the abdomen is longer than any of the others, which are subequal, and is rounded posteriorly. \ i > F Fic. 3. Leptognathia (?) psammophila, sp.n.; (A), female, dorsal view, x 20; (B), first antenna, x 80; (C), second antenna, x 80; (D), mandible, x 320; (E), maxilla, x 320: (F), maxilliped, x 320; (G), gnathopod, x 40; (H), fourth leg, x 80; (I), pleopod, x 80; (J), uropod, x 80. Cc G The peduncle of the biramous uropods is stout. The outer branch consists of two subequal, short articles, while the inner 12 WALLACE: THE ISOPODA OF THE BAY OF FUNDY consists of two subequal, long articles, each of which is as long as the entire outer branch. The gnathopods are large and ovate. The propodus is produced at its extremity into along, narrrow thumb, which is armed on its inner side with two prominent teeth, the point being produced to form a third. The dactylus is long and narrow and produced into a sharp tip which on closure of the chela falls between the teeth at the end of the propodus and the one next to it. The outer surface of the dactylus has a number of short teeth or tubercles. The remaining legs are all ambulatory in character and have not the basal article appreciably expanded as in Typhlotanats. The fifth, sixth and seventh are furnished each with a couple of spines on the fourth, fifth, and sixth joints, but those of the second and third legs have only a few setae. The mandible has a cutting edge of two teeth and a_ finely serrated anterior border. The molar expansion is very slender. Each maxilla is tipped with a circle of stiff bristles. The maxillipeds have each a palp of four articles. The position of this species is open to doubt. In the degraded condition of the molar expansion of the mandible it agrees with the genus Leptognathia, but differs from it in having the first pair of antennae with only three instead of four articles. It may be necessary to erect a new genus for its reception, but we have considered the condition of the mandibles as of major importance, and have, therefore, placed it in the genus Leptognathia. Leptochelia filum (Stimpson). Figure 4. Stimpson, 1853, p- 43- This species occurs throughout the whole region at depths varying from 9 to 75 fathoms, chiefly on hard, rocky bottom among Boltenia ovifera, but also in old shells, mud or sand. The males are not so numerous as the females, there being as a rule about 8 to 10 females for every male. Records: Off Biological Station, St. Croix River, 10 to 15 fathoms; off Eastport, Me., 10 fathoms; off Cherry Island, WALLACE: THE IsOPODA OF THE BAy oF Funpy 13 Fic. 4. Leptochelia filum (Stimpson); (A), male, dorsal view, x 21; (B), first antenna, x 84; (C), second antenna, x 84; (D), chela of gnathopod, x 154; (E), gnathopod, outer surface, x 84; (F), pleopod, x 84; (G), uropod, x 84; (a), female, dorsal view, x 21; (b), second antenna, x 84; (c), pleopod, x 84; (d), uropod, x 84. 14 WALLACE: THE IsopODA OF THE Bay OF FUNDY Head Harbour Passage, 42 fathoms; off Spruce Island, 36 fathoms; Head Harbour, 9 fathoms; off Head Harbour Island, 27 and 70 to 75 fathoms; the Wolves, 16 to 30 fathoms; off Low Duck Island, Grand Manan, 34 fathoms; off Big Duck Island, 42 fathoms; off Three Islands, Grand Manan, 17 fathoms. Hake Bay, Grand Manan, 20 fathoms (Stimpson, 1853). As this species has never been described more thoroughly than in the brief account given by Stimpson, we will consider its structure somewhat fully. Female: The body is elongated and narrow, about five times as long as broad (214 mm.: 0.5 mm.). The head is longer than it is broad, and has the anterior end narrower than the posterior. The eyes are small, compound and distinct, and situated anterolaterally. The first pair of antennae have the first article long and stout, the second very short, and the third about twice as long as the second. The second pair of antennae are shorter than the first pair, and consist of five articles, of which the first is short and broad, the second a little longer and thinner than the first, the third shorter than the second, the fourth more than twice as long as the third, and the fifth half as long as the fourth. Of the thoracic segments, the second (first free segment) is about half the length of one of the four that follow it, which are subequal. The seventh is about as long as the second. The sixth segment of the abdomen is longer than any of the other five and is rounded posteriorly. The outer branch of the biramous uropods is composed of two very short articles, which are not clearly marked off from each other, but which usually have a stout hair at the joint between them. The inner branch is four times as long as the outer and is composed of four articles, of which the first two are subequal in length and short, and the last two also subequal in length but at the same time one and one-half times as long as the first two. F The gnathopods are large, the propodus being produced at its extremity into a long, narrow thumb, which is armed on WALLACE: THE ISOPODA OF THE BAY OF FUNDY I5 its inner side with a row of low, blunt teeth together with a sharp, prominent tooth at the very tip. The dactyl is long, narrow and sharply pointed. The other legs are ambulatory, with the terminal joint sharply pointed and recurved. The abdomen has five pairs of well developed biramous pleopods, not quite so large as those of the male and having shorter setae, which do not extend beyond the limits of the terminal segments. Male: The body is about three-fifths as large as that of the female and equally elongated (1.58 mm.: 0.3 mm.). The head is slightly longer than it is broad and narrows toward the front. The eyes are proportionately much larger than in the female, and occupy the sides of the head lateral to the first pair of antennae. The latter have the first article long and stout, the second nearly as stout but only a little over half as long, and the third only one-third as long as the second and narrower. The flagellum is nearly as long as the peduncle and is multi-articulate, consisting of five articles, which, with the exception of the exceedingly minute terminal one, are subequal in length. There is some indication of the separation of the first and fifth articles into two parts in the presence of numerous hairs. The second pair of antennae are shorter than the first pair and not as stout. Each consists of six articles, of which the first is short and stout, the second twice as long as the first, the third shorter than the second, the fourth three times as long as the third, the fifth twice as long as the third, and the sixth quite inconspicuous. The thoracic segments are similar to those of the female except that the fifth and sixth segments are a little longer than the third and fourth. The thorax narrows to the sixth seg- ment. The last abdominal segment is very obtusely pointed posteriorly, being nearly round. The abdomen is broadest in the middle and tapers to each end. The uropods are as in the female. 16 WALLACE: THE ISOPODA OF THE BAY OF FUNDY The gnathopods are similar to those of the female, but the thumb is not so well armed, and the inner surface of the propodus has a comb of long, sharp spines, usually 7 to 10 in number, which extend across the article parallel to the joint with the dactyl. Of the legs the second pair are longer than those following. The abdomen has five pairs of well developed, biramous pleopods, which are slightly smaller than those of the female, but have long setae extending beyond the terminal segment and, therefore, appearing in a view from above. Leptochelia profunda sp.n. Figure 5. Males and females of this species were obtained off Head Harbour Island at a depth of from 70 to 75 fathoms, from a bottom of sandy mud and stones on September 2nd, 1913. Female: The body is elongate and filiform, being about six times as long as broad (about 2.7 mm.: 0.48 mm.). The head (cephalosome) is a little longer than wide, and is narrower at its anterior end than posteriorly. The anterior margin is excavated at each side of a small median projection for the reception of the first pair of antennae. The eyes are proportionately smaller than in the male. The first pair of antennae are triarticulate, the first article being long, the second one-third as long as the first, and the third nearly twice as long as the second. The second pair of antennae are as in the male. The second (first free) thoracic segment is the shortest, the third, fourth, and fifth progressively longer, and the fifth, sixth and seventh progressively shorter. The gnathopods and the other thoracic legs are as in the male. The abdomen resembles that of the male. The pleopods have shorter setae than in the male, while the uropods may have only five seg- ments in the inner branch, in which case the first article gives an indication of subdivision in the presence of a bristle about its middle. Male: The body is long and filiform, being nearly six times as long as broad (about 2 mm.: 0.35 mm.). The WALLACE: THE ISOPODA OF THE BAY OF FUNDY 17 Fic. 5. Leptochelia profunda sp. n.; (A), male, dorsal view, x 30; (B), first antenna, x 120; (C), gnathopod, x 120: (D), second antenna, x 120; (E), pleopod, x 120; (F), uropod, x 180; (a), female, dorsal view, x 30; (b), second antenna, x 120; (c), pleopod, x 120. 18 WALLACE: THE IJSOPODA OF THE BAY OF FUNDY head is about as long as it is broad and is narrower in front. The anterior margin is slightly excavated on each side of a short, median, blunt point for the first pair of antennae. The eyes are very large and occupy the anterolateral angles of the head. The first pair of antennae have the first article long and stout, the second about three-quarters of the length of the first and nearly as stout, and the third one-third as long as the second and with a slight projection on the medial side. The flagellum consists of six articles. The second pair of antennae consist of a peduncle with five articles and a single, minute, flagellar article. The first three peduncular articles are short, the fourth is nearly as long as the first three together, and the fifth is three-quarters as long as the fourth. The second, third, fourth and seventh thoracic segments are subequal in length, the second being a little shorter than the others. The fifth and sixth segments are longer than any of the others and subequal. The gnathopods are quite stout, and the propodus of each has on its inner, medial surface a comb of strong setae arranged in a line parallel to the joint with the dactyl. The other legs, which are ambulatory, have the dactyl produced into a sharp point, that of the second leg being much longer than those of the following. The sixth abdominal segment ends posteriorly in an obtuse angle. The pleopods are like those of L. flum and have long setae projecting beyond the edges of the abdomen. The biramous uropods have the inner branch as long as the last three abdominal segments together and six-jointed. The outer branch is biarticulate and short, being equal in length to the first two articles of the inner branch. This species differs from L. savignyi in that (1) the outer branch of the uropods is biarticulate, (2) the propodus of the gnathopods is devoid of teeth, and (3) the articles of the peduncle of the first pair of antennae have different relative lengths. It differs from L. filum in that (1) it is larger, (2) the males are relatively more numerous, (3) there are more articles in the flagellum of the first pair of antennae, (4) the WALLACE: THE ISOPODA OF THE BAY OF FUNDY 19 inner branch of the uropod has six articles, and (5) the inner surface of the thumb of the propodus of the gnathopod is without teeth. Family Guathidae Gnathia cerina (Stimpson). Richardson, 1905, p. 59. This species was found in abundance throughout the Bay of Fundy and in Passamaquoddy Bay at depths of from five to fifty-five fathoms on sand or mud or among old shells, and on hard stony bottoms. Also the larvae were taken from the surface of the following fishes: winter flounder(Pseudo- pleuronectes americanus), cod (Gadus callarias), hake (Uro- phycis tenuis or U. chuss), and haddock (Melanogrammus aeglifinus). Records: St. Croix River, off Atlantic Biological Station, 15 fathoms; Gleason’s Cove, 1-5 fathoms; off Bald Head, Campobello, 20 fathoms; the Wolves, 16 to 35 fathoms; Grand Manan, off Fish Head, Duck Islands, Green Islands, and Southern Head, 12 to 55 fathoms. Eastport, 10 to 20 fathoms; off Head Harbour, 40 fathoms; Bay of Fundy, 25-30 and 60 fathoms (Harger, 1880). Off Cheney’s Head, Grand Manan, Io fathoms (Stimpson, 1853). Jn most cases males, females and larvae were obtained. Larvae alone were found living parasitically on fishes at the following localities: on flounder at Campobello Island; on cod and hake at Bliss Island; and on haddock in the St. Croix River and Passama- quoddy Bay. Family Anthuridae Ptilanthura tenuis, Harger. Harger, 1880, p. 406. Body very narrow and elongate, being ten times as long as broad (7.8 mm. long, 0.8 mm. broad.) Head a little broader than long with the anterior margin produced into a short and blunt rounded process which covers over the bases 20 WALLACE: THE ISOPODA OF THE BAY OF .FUNDY of the antennae. Eyes are small, rounded, and very distinct. The first antenna has the first two articles subequal in length, while the third is a little longer than the second. The fourth, or first article of the flagellum, is equal to the third, while the fifth is very minute. The second pair of antennae have the first article very short; the second is about three times as long as the first; the third article is about one-half as long as the second; the fourth and fifth are a little longer than the third and are subequal. The flagellum is composed of three articles and is covered with hairs. The first pair of antennae extend to the end of the peduncle of the second pair. The maxilliped has a palp consisting of two articles. Each maxilla terminates in a number of conspicuous, sharp teeth. The labium terminates in two rounded lobes. The first thoracic segment is longer than the head. The second and third are equal in length and each is a little longer than the first. The fourth and fifth are slightly longer than the third, and the sixth is a little shorter than the fifth, while the seventh is about half as long as the sixth. The abdomen is more than twice as long as the seventh thoracic segment. Its first five segments are coalesced, only sutures indicating the segments. The sixth segment is free and bears the uropods. The terminal segment is evenly rounded posteriorly. The peduncle of the uropods is as long as the superior branch, which is rather lanceolate in shape and acutely pointed posteriorly. The inferior branch is one-half as long as the peduncle and is rounded posteriorly. The first three pairs of legs are prehensile in character while the others, though with recurved dactyls, are ambula- tory in character. The colour is a mottled brownish-red and white. One specimen was taken in Seal Cove Sound, Grand Manan, at a depth of eleven and a half fathoms, fine sand bottom. Grand Manan (Harger 1880). WALLACE: THE ISOPODA OF THE BAY OF FUNDY 21 (Mr. Wallace believed this to be a species of Cyathura, judging by the partially fused condition of the abdomen, and by the presence of two joints in the palp of the maxilliped. There can, however, scarcely be any doubt but that this is the female of Harger’s species, and consequently any distinct difference between the genera Cyathura and Pitlanthura must be restricted to the second pair of antennae of the male. A. G. H.). Calathura branchiata (Stimpson). Richardson, 1905, p. 72. This form is very abundant in the Bay of Fundy, but not in Passamaquoddy Bay. Numerous specimens were - taken near the Wolves Islands, near White Horse Island, and off Head Harbour Island at depths of from twenty to seventy- five fathoms on muddy bottom or in mud with sand and gravel. Off Duck Island, Grand Manan, 20 fathoms (Stimpson, 1853). Off Head Harbour, 75-80 fathoms; between Head Harbour and the Wolves, 60 fathoms; Grand Manan (Harger, 1880). Family Cirolanidae Cirolana polita (Stimpson). Richardson, 1905, p. 99. Specimens of this species were not very numerous, being taken only near West Quoddy head, Maine and at Grand Manan at depths varying from low tide mark to forty fathoms. The nature of the bottom on all occasions was soft fine sand. Records: Inside West Quoddy Head, Maine, low tide and 9 fathoms; Whale Cove, Grand Manan, 30 to 40 fathoms; Woodwards Cove, Grand Manan, low tide and 2 fathoms. High Duck Island, Grand Manan, low tide (Stimpson 1853). The specimens agreed with the description given by Richardson (1905, p. 99) except that there was a considerable variation in the number of articles in the flagellum of the 22 WALLACE: THE JSOPODA OF THE BAY OF FUNDY second pair of antennae, the number ranging from 7 to 19, but usually there were 10. Family Aegzdae Aega psora (Linn.). Richardson, 1905, p. 168. Only three individuals of the “‘salve-bug”’ were seen, and none of these was taken in Passamaquoddy Bay. Records: From cod (Gadus callarias), near Campobello Island; from skate (Raza radiata) caught off North Head, Grand Manan; from skate (Raza stabuliforis) caught in St. Mary Bay, N.S. Family Limnoritdae Limnoria lignorum (Rathke). ‘Gribble’. Richardson, 1905, p. 269. This isopod is abundant all through the region on nearly all submerged pieces of wood and timber, which it destroys by burrowing into them. Bay of Fundy (Harger, 1880). Family Idothetdae Chiridotea caeca (Say). Richardson, 1905, p. 353. The animals of this species were not so abundant as those of the next species and occurred only in very shallow water (two fathoms or less in depth) and on fine sand bottom. Records: Inside West Quoddy Head, Maine, at low tide; Woodwards Cove, Grand Manan, at low tide and ata depth of about two fathoms. The specimens ranged in size up to a length of about 15 mm. Chiridotea tuftsit (Stimpson). Richardson, 1905, p. 354. This species was taken in abundance at depths of from 5 to 12 fathoms on sandy bottom. WALLACE: THE ISOPODA OF THE BAY OF FUNDY 23 Records: Duck Pond, Campobello, 5 fathoms; off West Quoddy Head, 9 fathoms; Seal Cove Sound, 12 fathoms; off Green Island, Grand Manan, 11 fathoms. At low water in Prince’s Cove, Eastport (Harger, 1880). Off Cheney’s Head, Grand Uanan, 10 fathoms (Stimpson, 1853). An examination of numerous specimens showed the number of articles in the flagellum of the second pair of antennae to be variable, ranging from 10 to 14. Idothea baltica (Pallas). Richardson, 1905, p. 364. This form is not very common in the Bay of Fundy nor in Passamaquoddy Bay. It was found usually at low tide among seaweed and eelgrass, and also on floating seaweed and at depths as great as 5 fathoms on gravel or sand bottom. Records: Minister’s Island, low tide; St. Croix River near Biological Station, at the surface on seaweed; Katy Cove, . low tide; Deep Cove, Campobello, low tide mark; Woodward's Cove, Grand Manan, low tide; Grand Harbour, Grand Manan, 2 to 5 fathoms; Little River, St. Mary Bay, N.S., in lobster pond. Grand Nanan, rare (Stimpson, 1853). Bay of Fundy (Harger, 1880). Idothea phosphorea (Harger). Richardson, 1905, p. 366. This is found more generally in the Bay of Fundy than is the preceding species, being taken in many places, at depths ranging from low tide mark to 15 fathoms, on mud, sand or sawdust, or on hard, rocky bottom. Records: Minister’s Island, low tide; Oak Bay, 5 to 9 fathoms; St. Croix River near Biological Station, 5 and 15 fathoms; Gleason’s Cove, I to 5 fathoms; Woodward’s Cove and Grand Harbour, Grand Manan, 2 to 5 fathoms; Lepreau ledges, low tide. Whiting River (Harger, 1880). St. An- drews region (MacDonald, 1912). The number of articles in the flagellum of the second pair of antennae was noted to be as low as 10 in what were appar- ently adult individuals. 24 WALLACE: THE ISOPODA OF THE BAY OF FUNDY Synidotea nodulosa (Kroyer). Richardson, 1905, p. 388. Only two specimens were seen. One, taken near Green Islands, Grand Manan, at a depth of 11 fathoms, was olive green in colour and corresponded with the description given by Richardson (1905, p. 388). The other, dredged in Seal Cove sound, Grand Manan, at a depth of 12 fathoms on a bottom of fine sand, was dark, yellowish-brown, excepting the fourth thoracic segment, which was distinctly red. It differed otherwise in that the flagellum of the second pair of antennae consisted of eight articles instead of six, the tuber- cles on the body were much sharper, and the abdomen was more sharply pointed. Edotea triloba (Say). Figure 6. Synonyms: Idotea triloba, Say, 1818, p. 425. Epelys trilobus, Harger, 1873, p. 571, et auct. var. Edotea triloba, Miers, 1883, p. 70 et auct. var. Idotea montosa, Stimpson, 1853, p. 40. Epelys montosus, Harger, 1873, p. 571, et auct. var. Edotea montosa, Miers, 1883, p. 72, et auct. var. Edotea acuta, Richardson, 1900, p. 228, and 1905, p. 395. This species was found to be very abundant throughout the region, and occurred in depths ranging from low tide mark to 15 fathoms, and on a variety of bottoms—mud, sand, shells, rock and sawdust. Records: Brandy Cove, off Joe’s Point, and off Navy Island, St. Croix River, 2 to 15 fathoms; St. Andrews Har- bour, 2 to 3 fathoms; Gleason’s Cove, I to 5 fathoms; off Eastport, 10 fathoms; Deep Cove, low tide and Duck Pond, Campobello, I to 5 fathoms; off West Quoddy Head, 9 fathoms; entrance to Head Harbour, 5 fathoms; Bliss Island, low tide; Grand Harbour, 2 to 5 fathoms and Woodward’s Cove, Grand Manan,, 0 to 2 fathoms. Grand Manan (Stimpson, 1853). Eastport; Whiting River; Seal Cove, Grand Manan (Harger, 1880). WALLACE: THE ISOPODA OF THE BAY OF FUNDY 25 The size reached 10 mm. The body is ovate, the length varying from two to two and one-half times the breadth, and the abdomen is usually about one-third the length of the entire animal. The head is about twice as broad as long with the lateral angles markedly produced and varying in form from rounded lobes to knobor horn-like projections, as described by Richard- son for Edotea acuta, all the intermediate stages being repre- Fic. 6. Edotea triloba (Say); (A). (B), and (C), individuals dredged off West Quoddy Head in 9 fathoms, sand botiom, Aug. 13, 1912; (D), individual dredged in Seal Cove sound, 11% fathoms, sand bottom, July 17, 1913. sented in a series taken from the one locality. The eyes are large and situated just behind the anterior projections. The top of the head has two conspicuous tubercles, one on either side of the median line, close together and near the anterior end. Most of the individuals showed a slight depression just behind the tubercles, but not nearly so well defined as figured by Richardson for Edotea acuta, although in relatively the same position (see Figure 6). 26 WALLACE: THE IJSOPODA OF THE BAY OF FUNDY The third and fourth thoracic segments are the longest and also the widest, being nearly equal in width. The epimera are firmly fixed toeach segment. The lateral margins of the thoracic segments vary from being sharply angular and, in the case of the anterior ones, having more or less knob-shaped projections, to being broadly rounded or nearly straight, the intermediate conditions found forming a perfect series. The thorax in some individuals was quite convex centrally while in others it was comparatively flat, although those with the greater convexity were more numerous. The lateral portions of the segments are flatter showing a slight elevation towards the edge. The abdomen is composed of a single segment with in- cisions of various depths near the base indicating a partly coalesced segment. The sides of the abdomen converge rapidly from a point a little below the middle, and the extrem- — ity varies from a triangular condition to a bluntly pointed one. Near the base is a large rounded eminence, which seems to be continuous with the convexity of the thorax; and separated by a deep depression from this tubercle is another elevation which extends nearly to the tip of the abdomen. After a careful examination of a large series of individuals from the Bay of Fundy, there was found to be a complete gradation in the characters typical of E. triloba, E. montosa and E. acuta, there appearing all conditions of variation from one to another as regards length and form of the lateral projections of the head, borders of the thoracic segments, and general shape of the abdomen. The differences in the relative lengths of the antennae were so slight as to be useless for distinguishing distinct species. There appears to be, therefore, no good reason for separating E. montosa and E. acuta from E. triloba. Family Janiridae Jaera marina (Fabr.). Richardson, 1905, p. 450. WALLACE: THE IsopoDA OF THE BAY OF FuNDY 27 This species is commonly found in tide-pools and under stones between tide-marks throughout the region. Grand Manan (Stimpson, 1853). Eastport; Dog Island (Harger, 1880). The size attained was as great as 5.5. mm. in length. A great variation in the number of articles in the flag- ellum of the second pair of antennae was found, one specimen having as many as fifty and yet in other respects agreeing with the typical individuals. Janira alta (Stimpson). Richardson, 1905, p. 475. Specimens were taken in the Bay of Fundy at depths of from 14 to 75 fathoms on bottoms consisting of hard rock, sand, mud or shells. None were found in Passamaquoddy Bay. Records: Wolves Islands, 20 fathoms; off Head Harbour Island, 70 to 75 fathoms; Head Harbour Passage (off Cherry Island), 42 fathoms; Whale Cove, Grand Manan, 20 to 30 fathoms; off Southern Head, Grand Manan, 14 fathoms. Grand Manan (Stimpson, 1853). Clark’s Ledge near East- port, low water; off Buckman’s Head; off Todd’s Head (Harger, 1880). Family Munnidae Munna fabricti, Kroyer. Figure 7. Richardson, 1905, p. 480. This was taken frequently in the Bay of Fundy, Passama- quoddy Bay and the St. Croix River, at low tide and at depths as great as 42 fathoms, on bottoms consistin g usually of sand or mud, but also on those of hard rock or of sawdust. Records: St. Croix River near Biological Station, 2 to 15 fathoms, Passamaquoddy Bay, off Tongue Shoal light, 5 fathoms; Head Harbour Passage (off Cherry Island), 42 fathoms; near North Head, Grand Manan, low tide mark; off Big Duck Island, Grand Manan, 42 fathoms. South Bay, Eastport, 12 fathoms (Harger, 1880). 28 WALLACE: THE ISOPODA OF THE BAY OF FUNDY The first pair of antennae reach nearly to the penultimate joint of the peduncle of the second pair. The flagellum has three or four joints, usually four. Cc E view, x 47; (B), seventh leg, x 47; (C), first Fic.7. Munna fabricii, Kroyer; (A), dorsal urface of abdomen of female, operculum raised, antenna, x 370: (D), uropod, x 370; (E), lower s x 185. WALLACE: THE IsopopA OF THE BAY oF FUNDY 29 The flagellum of the second pair of antennae is longer than the peduncle, but varies slightly. The uropoda are obliquely truncate, but usually show three processes from the free edge. The caudal segment in large specimens has the apical lamella distinctly serrated and a small hair in place of the lateral denticle, although the size varies. Small individuals often failed to show the serrations on the apical lamella. The colour was usually a dark dusky brown and often with black spots of pigment. Pleurogonium rubicundum (G. O. Sars). Figures 8 and 9. Sars, 1899, p. 113. This species of Pleurogonium is much more restricted in habitat than are the other two that occur in this region. It was found only in the vicinity of St. Andrews, never in the open Bay of Fundy. It was confined to shallow water at depths of from 2 to 5 fathoms on bottoms of sand, mud or sawdust. Records: Brandy Cove and near Joe’s Point, St. Croix River, 2 to 4 fathoms: outside Tongue Shoal light, Passama- quoddy Bay, 5 fathoms. Female (Fig. 9): The body is short and stout, being a little less than twice as long as broad. The head is not quite twice as broad as long and is rounded anteriorly. The eyes are entirely absent. The first pair of antennae project laterally from the head and have a peduncle of three articles, of which the first two are subequal, while the third is only about half as long as the others. The flagellum consists of three articles which are nearly subequal in length, the last article being surmounted by a stout bristle and a tuft of hairs. The second pair of antennae project dorsally and laterally from the head, and each is composed of a peduncle of six and a flagellum of seven articles. The first two articles of the peduncle are very small and inconspicuous from above, the third is long and has a decided protuberance on its outer side near the proximal end, the fourth is about one-half as long 4 ; | | 30 WALLACE: THE ISOPODA OF THE Bay OF FUNDY as the third and slightly expanded distally, the fifth article is one and one-half times and the sixth about twice as long as the fourth. G Fic. 8. Pleurogonium rubicundum (G. O. Sars); (A), male, dorsal view, x 30; (B), first antenna, x 240; (C), second antenna, x 240; (D), mandible, x 240; (E), gnathopod, x 240; (F), operculum, x 120; (G), uropod, x 240. The first four segments of the thorax with the head are nearly circular in outline, while the last three are markedly narrower and are directed posteriorly. The body is broadest at the level of third thoracic segment. The lateral edges of WALLACE: THE ISOPODA OF THE BAY OF FUNDY 31 the thoracic segments are rounded and from the middle of each projects a long, rod-like spine, those on the third segment being the longest. Fic. 9. Pleurogonium rubicundum (G. O. Sars); (A), female, dorsal view, x 30; (B), second antenna, x 240; (C), first antenna, x 240; (D), maxilliped, x 240; (E), gnathopod, x 240; (F), second leg, x 60; (G), abdomen, lower surface, x 60; (H), uropod, x 240. The abdomen is very narrow at the base but widens very much, and again narrows to end in an obtusely pointed tip. The sides of the abdomen are evenly curved and fringed with hairs. The operculum is a little narrower and more sharply pointed than the abdominal segment. 32 WALLACE: THE ISOPODA OF THE BAy oF FUNDY The uropods are terminal and biramous, the outer branch being extremely small, while the inner one is about three times as long and has a slight notch in its inner edge. The mandible has a slender molar expansion and a cut- ting part with four teeth. The palp of the maxilliped con- sists of five articles. The first thoracic leg on each side is strongly recurved for seizing, the carpus strongly armed with spines, and the propodus oval and dilated. The other legs are ambulatory in character. Male (Fig. 8): The body is short and oval being about twice as long as broad (1.2 mm.: 0.66 mm.). The head is a little broader and not as long as in the female. The eyes are entirely absent. The two pairs of antennae are practically the same as those of the female. The thorax does not show a distinct division into a_ rounded anterior portion and a narrower posterior part, but broadens gradually to the third and fourth segments and then narrows. ‘The lateral spines are not quite so pronounced as in the female. The abdomen, uropods, first legs, and mouth parts are similar to those of the female, but the operculum differs from that of the female in consisting of three parts, an unpaired medial and paired lateral parts. The medial portion is lanceolate in shape and bears a tuft of short, stout hairs at the ends of the expanded points and the lateral parts are oval and flat, and each has a palp-like pointed process on the inner edge. Pleurogonium inerme, (G. O. Sars). Figure Io. Sars, 1899, p. 114. This species is somewhat restricted to the inner bays, where it is fairly abundant at depths of from 2 to 10 fathoms on bottoms of mud, sand or sawdust, but a few were taken on hard bottom at a depth of 15 fathoms. It was taken in the open Bay of Fundy at the Wolves on one occasion. Records: Oak Bay, 5 to 9 fathoms; Brandy Cove, Joe’s Point reef, and opposite Robbinston, St. Croix River, 2 to WALLACE: THE IsopopA OF THE BAY OF FUNDY 33 15 fathoms; St. Andrews harbour, 2 to 3 fathoms; off Tongue Shoal light, Passamaquoddy Bay, 5 fathoms; Wolves Islands, 16 fathoms. Fic. 10. Pleurogonium inerme,‘(G. O. Sars); (A), male, dorsal view, x 40; (B) operculum, x 180; (a), female, dorsal view, x 40; (b), operculum, x 90. 34 WALLACE: THE ISOPODA OF THE BAy OF FUNDY Female: The body is short and stout, being a little less than twice as long as broad (1.5 mm.: 0.85 mm.). The head is about twice as broad as long and rounded anteriorly. Eyes are entirely absent. The first pair of antennae have a ped- uncle of three articles of which the first two are long and subequal, and the third one-half as long as the second. The flagellum consists of three articles, of which the first is about one-half the length of the last article of the peduncle, and the second and third subequal and one and one-half times as long as the first. The third article is surmounted by a stout bristle, which is as long as the last two articles together. The second pair of antennae have a peduncle of six and a flagellum of seven articles. In the peduncle the first two articles are very short and equal in length, the third is long and has a prominent protuberance on the outer side near the proximal end, the fourth is less than one-half as long as the third, the fifth is slightly longer than the fourth, and the sixth is twice as long as the fifth. The four anterior segments of the thorax are nearly circular, while the last three are decidedly narrower, the thorax being broadest at the third segment. The lateral edges of all the thoracic segments are rounded. The abdomen is much constricted at its base from which it broadens and then narrows to end in an obtusely pointed posterior extremity. The lateral edges are evenly curved and fringed with hairs. The operculum is not as broad as the abdomen and tapers to a sharper point. The uropods are terminal and biramous, the peduncle consisting of a single short article. The outer branch is - extremely small while the inner one is nearly three times as long. The first leg has the dactyl strongly recurved and is almost exactly the same as that of the preceding species, as is the case also with the other legs. The mandibles are much as in the preceding species, having a slender molar expansion and a cutting part with four teeth. The other mouth parts also are almost the same as those of P. rubscundum. WALLACE: THE Isopopa OF THE Bay oF FUNDY 35 Male: The body is small and oval, being only a little more than twice as long as broad (1.17 mm:0.55 mm.), resembling that of the young female. The head is about twice as broad as long and is rounded anteriorly. The two pairs of antennae are as in the female. The thorax does not show two distinct regions as in the female, and is widest at the level of the third and fourth seg- ments, which are subequal in width. The lateral edges of all the thoracic segments are rounded. The abdomen and uropods are practically the same as those of the female, except that the operculum consists of the usual three parts, of which the medial is lanceolate, with expansions near its middle, each expansion bearing a tuft of short strong bristles, and the two lateral flat and ovate, each having a palp-like pointed process on its medial side. The first leg and mouth parts are as in the female. Pleurogonium spinosissimum (G. O. Sars). Figure 11. Sars, 1899, p. 115. This species has a wider distribution than either of the other two species of the genus in our waters, and it was found as frequently in the open Bay of Fundy as in Passamaquoddy Bay. It occurred at depths ranging from 2 to 75 fathoms, chiefly on bottoms of sand and mud, but also in old shells, or on gravel or stones. Records: Oak Bay, 5 to 9 fathoms; Brandy Cove and opposite Robbinston, St. Croix River, 2 to 1 5 fathoms; off Head Harbour Island, 70 to 75 fathoms; Wolves Islands, 16 to 30 fathoms; off Low Duck and Big Duck Islands, Grand Manan, 34 and 42 fathoms; Grand Harbour, Grand Manan, 2 to 5 fathoms. Female: The body is about twice as long as broad (about 2.25 mm. :1.2 mm.). The head is twice as broad as long and rounded anteriorly. It is emarginate on each side of the Origin of the antennae. The eyes are entirely lacking. The peduncle of the first pair of antennae consists of three articles, of which the first is slightly longer than the second, and the 36 WALLACE: THE ISOPODA OF THE BAY OF FUNDY third one-half as long as the second. The flagellum consists of three articles, of which the first two are subequal, and the third somewhat longer and surmounted by a stout bristle. The second pair of antennae have a peduncle of six and a flagellum of seven articles. In the peduncle the first two articles are short and inconspicuous viewed dorsally, the third is long and has a distinct tubercle on its outer side near the proximal end, the fourth is one-third the length of the third, the fifth is slightly longer than the fourth, and the sixth is twice as long as the fifth. The first four segments of the thorax are subequal in length while the last three are short and nearly equal in length. The lateral edges of the first four segments are pro- duced each into an irregular angle, from the anterior side of which there projects a long, sharp, diverging, serrated process. | The angles of the fourth segment are not nearly so marked as those of the preceding three. Each of the last three segments has a long serrated spine on each side, just as in the anterior ones. The abdomen is very broad and sharply pointed poster- iorly. The operculum is very broad at its middle and is produced into a very sharp, acute tip posteriorly. The uropods are much as in P. rubicundum. The mouth parts and the legs (except the first pair) closely resemble those of the two preceding species, while the carpus and propodus of the first pair of legs are more strongly armed than in those species. The colour is usually brownish or reddish-brown. Male: The body is longer and narrower than that of the female, being nearly twice as long as broad (about 1.4 mm.: 0.6 mm.). : The head is much as in the female, as are also the anten- nae and mouth parts. The thorax differs from that of the female in that the angular processes from the lateral edges of the first four seg- ments are not nearly so pronounced and are almost entirely WALLACE: THE IsOPODA OF THE BAY OF FUNDY ae absent in the fourth segment. The legs, including the first pair, are like those of the female. Fic. 11. Pleurogonium Spinosissimum (G.O.Sars); (A), male, dorsal view, x 30; (B), operculum x 120; (a), female, dorsal view, x 30; (b), abdomen, ventral surface, x 60; (c), gnathopod, x 60, The abdomen and uropods resemble those of the female. The operculum, which varies much in both sexes, is in the male composed of the customary three segments, the central 38 WALLACE: THE ISOPODA OF THE BAY OF FUNDY part being lanceolate in shape and having an expansion in the middle bearing on each side a tuft of short, stout hairs, and the two lateral portions being flat and oval in outline, and each having a palp-like projection from its medial side, much as in the male of P. znerme. Family Munnopsidae Munnopsis typica. M. Sars. Richardson, 1905, p. 486. A single specimen was taken, swimming freely at the surface at the mouth of Harbour de Loutre, Campobello Island. The vertical currents resulting from the vigorous flow of the strong tides through the narrow and deep Head Harbour passage have apparently brought this deep water species to the surface. It had previously been dredged by ~ the United States Fish Commission expedition between Head Harbour and the Wolves at a depth of 60 fathoms. Eurycope mutica. G.O. Sars. Figure 12. Sars; 1899,.p. 140. Only five specimens in all were obtained, three of these in the St. Croix River off Robbinston at a depth of 15 fathoms, on hard, rocky bottom, and two near Tongue Shoal light, Passamaquoddy Bay, at a depth of 5 fathoms on muddy bottom. The body is short, oval and compact, being slightly less than twice as long as broad and having the greatest width at the middle (1.33 mm.: 0.75 mm.) The anterior end of the head is broad, rounded, and deeply excavated on either side for the reception of the antennae. The lateral margins of the head are produced into short, sharp processes. The superior antennae are short, and each has a very stout basal article. The second article is much more slender and not quite so long, the third and fourth are subequal and together equal to the second, and the flagellum is composed of four articles. The second pair of antennae consist each of a peduncle of five WALLACE: THE ISOPODA OF THE Bay oF FUNDY 39 articles and a multiarticulate flagellum twice the length of the peduncle, the entire antenna being twice as long as the body. The thorax consists of six distinct segments, the fifth showing indications of a subdivision by sutures which reach F Fic. 12. Eurycope mutica, (G. O. Sars); (A), female, dorsal view, x 40; (B), uropod, x 160 (C), first antenna, x 160; (D), second antenna, x 40; (E), mandible, x 160; (F), maxilliped, x 160; (G), first leg, x 160. half way across the body. The lateral margins of the seg- ments are not very sharply produced anteriorly. The ab- dominal segment is rounded posteriorly while its lateral margins are slightly coarctated. 40 WALLACE: THE IsOPODA OF THE BAY OF FUNDY The legs of the first pair are comparatively short, and have the propodal joint shorter than the carpus, and the dactylus very small. The natatory legs have the carpal joint oval in shape with a slight expansion on one side. The uropods are very small and biramous, each branch being uniarticulate. The outer branch is oval and about one-half as long as the inner, which is lanceolate in shape. The mandibles have cutting edges of sharp teeth and large, broad, molar expansions; the palp is large and curved. The maxillae are normal. Each maxilliped has a palp of five articles and an epignath which is short, broad and obtusely truncated at the tip. Family Bopyridae Phryxus abdominalis (Kroyer). Richardson, 1905, p. 500. This parasite was found only on specimens of Sptronto- caris pusiola, which were collected at the following places: — Minister’s Island, Passamaquoddy Bay, low tide; Head Harbour, Campobello Island, 5 fathoms; Grand Passage, off Dartmouth Point, St. Mary Bay, N.S., 15 fathoms; between North-west ledge and Brier Island, N.S., 32 to 36 fathoms. Bopyroides hippolytes (Kroyer). Richardson, 1905 p. 567. This species was taken at the following places :-—Digde- quash Harbour, 9 to 11 fathoms, on Spirontocaris polaris; south-east from Swallow Tail light, Grand Manan, “‘hake grounds”, on Spirontocaris spinus; High Duck Island, Grand Manan, low tide, on Spirontocaris pusiola. Bay of Fundy on Spirontocaris spinus and S. pustola. (Harger, 1880). Family Oniscidae Cylisticus convexus (De Geer). Richardson, 1905, p. 609. Specimens of this species were obtained from under WALLACE: THE IsopopA OF THE BAY OF FUNDY 41 decaying bark at Chamcook Mill near St. Andrews, and others were found by Dr. Philip Cox at Fredericton under stones. Porcellio rathket (Brandt). xtichardson, 1905, p. 617. A few specimens of this form were taken in dry sand and also in shaded woods near St. Andrews, also some by Dr. Cox at Fredericton. Porcellio scaber (Latreille). Richardson, 1905, p. 621. This species is common under driftwood above high tide throughout the region. 42 WALLACE: THE IsOpODA OF THE Bay OF FUNDY LITERATURE Harger, O., in Verrill, A. E. Report upon the invertebrate animals of Vineyard Sound, etc. Report U.S. Comm. Fish & Fisheries, 1871-72, pp. 293-478, pls. I-XXXVIII. 1873. Report on the marine Isopoda of New England and adjacent waters. Report U.S. Comm. Fish & Fisheries, 1878, Pt. 6, pp. 297-462, pls. I-x1m. 1880. Macdonald, D. L. On a collection of Crustacea made at St. Andrews, N.B. Contrib. Canad. Biol., 1906-1910, pp. 83-84. 1912. Miers, E. J. Revision of the Idoteidae, a family of sessile-eyed Crustacea. Journ. Linn. Soc., London, vol. XVI., pp. 1-88, pls. 1-111. 1883. Richardson, H. Synopses of North-American invertebrates, VIII: The Isopoda. Amer. Natur., vol. XXxXIV., pp. 207-230 and 295-309. 1900. Monograph on the Isopods of North America. Bull. U.S. Nat. Museum, No. 54, pp. Lii+727. 1905. Sars, G. O. Crustacea of Norway, II: Isopoda. 1899. Say, T. An account of the Crustacea of the United States. Journ. Acad. Nat. Sc., Philad., vol. 1, pt. 2, pp. 393- Aol and 423-433. 1818. Stimpson, W. Synopsis of the marine invertebrata of Grand Manan, etc. Smithson. Contrib. to Knowledge, vol. VI, Pp- 39-44. 1853. UNIVERSITY OF TORONTO TUDIES - _ BIOLOGICAL SERIES s . Y LIBRARY: PUBLISHED BY University of Toronto Studies COMMITTEE OF MANAGEMENT Chairman: Str RoBERT ALEXANDER FALCONER, LL.D Ke M.G. President of the University PRoressor W. J. ALEXANDER, Pu.D. Proressor J. J. Mackenzig, B.A., M.B. Proressor J. P. McMurricu, Pu.D. Bric.-GEen. C. H. MitcHeE Lt, B.A.Sc., C.B., COM Gs D5: 0: Proressor G. H. NEEDLER, Pu.D. PRoFESSOR GEORGE M. Wronc, M.A. General Editor: H. H. Lancton, M.A. Librarian of the University AN EGG OF STRUTHIOLITHUS CHERSONENSIS BRANDT | BY B. A. BENSLEY Professor of Zoology in the University of Toronto : i AN... EGG OF STRUTHIOLITHUS CHERSONENSIS BRANDT In February, 1919, the Royal Ontario Museum of Zoology, Toronto, acquired a fossil or semi-fossil egg, the features of which agree in all essential respects with the original and later descriptions of the type upon which Brandt’s species Struthiolithus chersonensis was founded. The specimen was presented to the Museum by the Reverend Harold M. Clark, of the Presbyterian Foreign Mission, formerly resident in Honan, China.* Remains of Struthiolithus chersonensis are of interest from several points of view. Some half dozen eggs are known with certainty, and three have been described or noted. The specimens indicate a struthious bird, even if not of the genus Siruthio, and evidently much larger than existing ostriches. No skeletal remains have been reported, though they must occur abundantly in the extensive Pleistocene deposits of eastern Asia. The continental distribution in- dicated by the points of discovery is not paralleled by any existing or fossil species, the former having a range which is African and Arabian (.S. camelus), while the latter so far have been reported from the Tertiary deposits of Samos and of the Siwalik Hills of India as skeletal remains not differing greatly from the modern species, and not of size relations suggesting Struthiolithus. At the same time the occurrence of Struthio- lithus remains in north-eastern China suggests a connection of the old world species with the nandu or three-toed ostrich of South America, though again there is no evidence of struthious birds in North America and accordingly no evidence of the utilization of the Behring Straits bridge as for the holarctic Tertiary mammals. *Vide Proc. and Trans. Roy. Soc. Can., vol. XIII, 1919. A second specimen also obtained by Mr. Clark has since been acquired, and has been presented to the British Museum, 4 B. A. BENSLEY Struthiolithus chersonensis was established by Brandt (’72) for the original egg, found in 1857 at Malinowka, Cherson Government, in southern Russia. The specimen was found floating in the water behind a mill weir, where it had evi- dently been washed out of the stream bed. Brandt’s de- scription of the egg includes the principal measurements, and he had also prepared a rough plaster mould. The specimen, however, was held for a price of one thousand roubles, and, not being sold, remained for many years in the possession of the owner or of his family, until finally it was accidentally broken into thirty-six pieces. At least some of the fragments ultimately found their way into the Petrograd Museum. The whole episode of the egg and its fate is reported by Brandt (85) in a subsequent article, the contents of which are by no means lacking in human interest. The ill-wind, in this instance at least, furnished an opportunity for examining the structure of the egg shell, on the strength of which Nathusius (’86) confirmed its ostrich-like nature, even going so far as to declare it generically inseparable from the existing Struthto. No further specimens were reported until 1898 when a second egg was brought to America and acquired for the Museum of Comparative Zoology of Harvard College. The specimen was found at a place some fifty miles south-west of Kalgan in Manchuria, and has been minutely described by Eastman (’98), who states that there were two of the eggs, probably in association, one, however, having been broken. In 1915 a third specimen was found in the province of Honan by a Chinese workman who saw it protruding from the bank of the Yellow River. A short note concerning its acquisition by the American Museum of Natural History is contained in the American Museum Journal. (17), the reference to which, together with particulars of the egg, were kindly furnished by Mr. Walter Granger aad \DrsiiF.\'Aa Lucas. The discovery of this specimen is referred to in Nature (18) in a review which has also a bearing on the present specimen. “Eggs of an extinct ostrich are already known from the surface deposits of northern China. One specimen from Yao Kuan Chang, fifty miles south-west of An Ecc oF STRUTHIOLITHUS CHERSONENSIS BRANDT 5 Kalgan, was obtained by Harvard University in 1898, and another specimen from the banks of the Yellow River in Honan was acquired by the American Museum of Natural History last year. Mr. Harold M. Clark of Wuan, Honan, now writes to the North China Herald that eggs of this kind are not uncommon in his neigh- bourhood, and are washed out of the river banks by floods. They seem to occur in the same manner as the eggs of Aepyornis on the shores of lakes in Madagascar. The Chinese eggs are about 7 in. in length, and thus scarcely larger than those of an average ostrich. No bones of birds which laid the eggs have hitherto been noticed in the same deposits.” Concerning the present specimen, Mr. Clark states that he obtained it from a native friend who is a travelling collector and dealer in curios. There is no exact information as to the geological setting. The egg was originally owned, and probably obtained, by a native workman in the village of Chwan Hu, in the extreme northern part of Honan, near the Pe Chi Li boundary. It was later borrowed by some rich relatives and taken to a neighbouring village of I Cheng, some ten miles distant, where it was exhibited at fairs. After purchasing the specimen, Mr. Clark heard of the existence of a very large egg in this village and was able to establish that it was the one in his possession. As to the common occurrence of these eggs Mr. Clark states that the newspaper report as quoted above is in- accurate. In addition to the second specimen acquired by him, and knowledge of the American Museum specimen, he has a fairly definite recollection of having been told of a fourth specimen which was formerly in an educational museum at Tungchow, near Peking, but was destroyed during the Boxer uprising of 1900. The specimen (Plate I) is complete except for a small perforation of about 15 mm. in diameter near one end. The longitudinal contour is evenly elliptical as in ostrich eggs generally, showing no indication of the polar inequality of the eggs of birds otherwise. Less than one-third of the surface, comprising an oval patch, is considerably eroded, while the balance is intact, giving the impression that the egg had been partly exposed for some time and had then suddenly been washed out of the earth. The intact portion of the surface is more closely, and in some respects more 6 B. A. BENSLEY finely pitted than in any ostrich eggs available for com- parison. Unfortunately the available descriptions of the eggs of the four existing species of African ostriches are widely divergent as regards the appearance and texture of the shell. The pits show a tendency to run together into short irregular depressions, due in part perhaps to the finer erosion of the surface. The interior of the shell is smooth, without encrustations and has a remarkably clean and fresh appearance. It shows a yellowish colour interspersed with dots of brown. The chief measurements are set forth below in com- parison with those reported by Eastman for the Harvard specimen and that of Brandt. US ee et ee ee Brandt Type Harvard sp. Toronto sp. eee eee eee IP ka Longitudinal axis..........-.-- 18.0 cm. 18.0 cm. 18.1 cm. DransverseyaxiSacneee to cae 1530 14.75 14.6 Ratiovaiaxese series ibe cuae 1.20 1.22 1.24 Major circumference........... 52.0 51.35 51.3 Equatorial circumference....... 46.0 46.45 45.75 Shrell thickness:e tea). Cat ke ones 2.65 2.20 1.98 Capaehby cry drei edb akan att 2075.0 1896 .90 1829.0 It will be seen from the data included above that the principal measurements are almost identical with those of the specimens previously described. There is a difference of some 67 c.cs. in cubic content of the Harvard and Toronto examples, but there are also undoubtedly differences of uniformity in contour in the latter and some indication of differences in the thickness of the shell, the measurements being made directly at the opening with a micrometer gauge. Concerning the weight of the shell in relation to its alteration, the figures for the Harvard and Toronto specimens are respectively 310.05 and 380. grams. It is evident both from the ordinary appearance of the egg of Struthiolithus and a comparison of the principal axes and cubic contents that the size is much greater than in the four existing species of African ostriches. Thus the An Eco oF STRUTHIOLITHUS CHERSONENSIS BRANDT 7 largest specimen of the latter quoted by Eastman (loc. cit.) gives longitudinal axis 16.38 cm., a difference of 1.62, while the cubic content is 1423.63, a difference of 473.27 c.cs. or of 405.37 c.cs. as compared with the Toronto specimen. 1872. 1885. 1886. 1898. 1917. 1918. LITERATURE CITED Brandt, A. Uber ein grosses Vogelei aus der Umge- bend von Cherson. Bull. de l’Acad. Imp. des Sci. St. Pétersbourg, T. xvili, pp. 158-161. 1873. Also Mélanges Biol., T. viii, pp. 730-735. 1872. Brandt, A. Uber das Schicksal des Eies von Struthio- lithus chersonensis. Zool. Anzeiger, Bd. viii, no. 191, ~pp. 191-2. 1885. Nathusius, W. von. Uber das fossile Ei von Struthio- lithus chersonensis Brandt. Zool. Anzeiger, Bd. ix, no. 214, pp. 47-50. 1886. Eastman, C. R. On remains of Struthiolithus cherso- nensts from northern China, with remarks on the dis- tribution of Struthious birds. Bull. Mus. Comp. Zool. Harvard, vol. xxxii, no. 7, pp. 127-144 (plate). 1808. The American Museum Journal, vol. xvii, no. 6, Bp. A429)! TQ17: Nature, vol. ci, no. 2525, p. 50. 1918. Vee A) ay) th 4 i rit yi h { at) HK Ny { Ait anal ba be 7 BYWAY alirbs Whe Mi te Swe NUN NENSIS Pret Wt ae oats ERSO CHINA O Yn on a e photograph Full si HONAN PROVINCE STRUTHIOI EGG OF ERSITY OF TORONTO O University of Toronto Studies COMMITTEE OF MANAGEMENT Chairman: Str ROBERT ALEXANDER FALCONER, LL Di. KC M.G, President of the University Prorgessor W. J. ALEXANDER, PH.D. Proressor J. J. Mackenzig, B.A., M.B. Proressor J. P. McMurricn, Pu.D. Bric.-Gen. C. H. Mitcuett, B.A.Sc., C.B.,C.M.G., D.S.O Proressor G. H. NEEpDLER, Pu.D. Proressor GEorGE M. Wrona, M.A. General Edttor: H. H. Laneton, M.A. Librarian of the University UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. | A PLAN FOR THE BIOLOGICAL INVESTIGATION OF THE WATER AREAS OF ONTARIO BY B. A. BENSLEY OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 AR si ne Nh 7A) A PLAN FOR THE BIOLOGICAL INVESTIGATION OF THE WATER AREAS OF ONTARIO During the past two years a considerable amount of attention has been given to the organization within the Department of Biology of the University of Toronto of a research branch to be devoted to the investigation of the water areas of Ontario with reference to existing biological conditions and economic problems. It has been proposed to carry on the work of this branch under the title ‘‘Ontario Fisheries Research Laboratory’’, some distinction being necessary to indicate its scope and outlook, as also to draw a line of demarcation between this and other interests of the Department, and being especially advisable in view of the work now conducted in marine biology. Moreover a special name is advisable in order to establish a basis for the better classification of publications which in this instance may be expected to appeal for the most part to those in- terested in general limnology or in conservation and fishery problems. Apart from the provision of apparatus and equip- ment the initial step in the formation of the new organization was the appointment of Professor ;\W. A. Clemens as Limno- biologist, an arrangement which renders possible full time service and supervision of field operations at any part of the year, and also provides for continuous investigation of the collected material and data. The first reports of a technical nature are now in course of publication. They refer principally to some special fisheries work on Lake Erie, but will be followed by a series of papers dealing with Lake Nipigon. The latter will be in a sense the products of the first field party, consisting of Professor Clemens and other members of the university and museum stafis who spent the greater part of the summer of 1921 in practical operations on the lake. The study of the scientific aspects of fishery questions is one for which the University Biological Laboratory is 3 4 BENSLEY: WATER AREAS OF ONTARIO especially fitted. Its position as part of the provincial educational system identifies its interest in matters touching the natural resources with that of the province at large. Nearly thirty years ago the desirability of a systematic biological survey of the provincial waters was urged by Professor Ramsay |Wright (’92), and in the interval the need of a thorough investigation of the factors underlying the distribution of aquatic organisms has become increasingly evident. Neither individual efforts nor the collective par- ticipation of members of the university staff in the work of the Georgian Bay Biological Station, which was maintained for some years in that region, led to the consideration of any measures upon which the hope of a continuous if not permanent organization for the biological investigation of provincial waters could be based. That the latter now ap- pears within the range of possibility is therefore a matter of some satisfaction. The following pages are devoted to a general discussion of the proposed line of investigation. This is mainly a matter of the relation of limnology at large to local conditions as represented by the waters of the Province of Ontario, with particular reference to general and economic problems. The principal topics considered are (a) the relation of the proposed investigation to the practical and administrative aspects of conservation, (b) the foundations of limnology, (c) the extent of the local fish fauna, and (d) the chief physical features of the provincial water areas, including, first, the provincial portions of the Great Lakes and, second, the land- enclosed areas, from the point of view of their limnological study and the nature of economic questions. Except in the case of the fishes, which occupy a central position in respect of the economic outlook, no reference will be made to particular groups of organisms. The distinction often made between theoretical and applied or economic considerations is generally recognized as purely a matter of convenience in description. Its value, if any, consists in denoting the purpose or outlook implied. There is, however, a relation between investigation of an ~ BENSLEY: WATER AREAS OF ONTARIO 5 economic kind or purpose and practical or administrative measures. In Canada, as elsewhere, various questions, partly of revenue, partly of control of exploitation of natural resources, wastefulness and many other matters having to do with natural wealth, gave rise long ago to regulative measures, and more recently to a public movement in the direction of conservation. More intricate perhaps than in other fields, because of the elusiveness of wild life and the obscure factors upon which its success depends, the problems of conservation as applied to living organisms were early recognized as involving scientific knowledge of an exact kind. On the other hand, the desire to arrive at what would usually be considered expert information has, in a great many instances, painfully evident in the literature of con- servation, given rise to confusion as to the extent and method of its application, resulting in the concealment of many sound practical ideas in a mass of information relating to general natural history. Measures of conservation as applied to the fisheries, including under the term also regulatory legislation, which has or ought to have a similar effect, consist broadly in striking a reasonable balance between forces of utilization and those of natural and articifial re- placement. In this process the administrative function is the first necessity; technical practice, as, for example, fish breeding and economic industrial processes, is next in order of urgency, while scientific investigation is accessory or contributory. Its value consists in the analysis of all factors physical, chemical and biological, and in drawing attention to particular factors the neglect of which may result in ineffective practice or futile legislation. The early development of fresh water biology in Europe, in many respects the foundation of similar work in America, established first of all the identity and distribution of the various species of aquatic organisms, and, second, the asso- ciation of these organisms with environmental conditions; to a certain extent also it carried out the classification of these conditions on the basis of their mutual associations. After nearly one hundred years of individual, and for the 6 BENSLEY: WATER AREAS OF ONTARIO most part isolated, effort on the part of early biologists, among whom will be found some whose names are foremost in the records of biological science, it gave rise, mainly through the efforts of Forel, Apstein, Zacharias, Zschokke and others, to an attempt at correlation of limnological results, best expressed by Zacharias (’91) and his co-workers in their compilation dealing with the fauna and flora of fresh water. The extensions of this earlier work not only resulted in a more thorough enumeration of species in local European areas, but also gave rise to the study of physical and chemical factors, environmental communities, plankton determination, and the formal and institutional recognition of limnology, as best exemplified by the complete limnological study of Lake Geneva by Forel (’92) and the establishment through Zacharias ('93) of the Biological Station of Plén after the pattern of the marine institution at Naples. American limnology, which naturally had a later origin, has the same foundation and in some respects a similar development. That it may be said to have only recently arrived at the stage of collective recognition is due to a variety of circumstances, including no doubt the great area to be covered in the classificatory study of the various groups, and the fact that the accumulation of information bearing in one way or another upon general limnology, as, for example, in the fisheries, natural resources in general, conservation and the like, has been almost from the beginning the particular interest of government bureaus, and of surveys and commissions under state or national auspices. Though the study of the water areas as entities, or complete physical- biological complexes, has not yet arrived at a stage of de- velopment commensurate with its great importance, its recognition is at least assured. As long ago as 1887 the principle was advocated by Professor Forbes (’87) and its application is observable in his later work (’08) in association with Richardson, Kofoid (’03) and others on the Illinois river system. ‘The same principle is exemplified in the work of others, some of whom owe their inspiration to the example of Professor Forbes, including that of Reighard (’94) on BENSLEY: WATER AREAS OF ONTARIO 7 Lake St. Clair, Ward (96, also ’18) on Lake Michigan, Birge and Juday (11) on the inland lakes of Wisconsin, the study of physical factors and their association by Need- ham and Lloyd (16), Shelford (13) and Adams (’13), and finally the recent work of Evermann and Clark (’20) on Lake Maxinkuckee, Indiana, in some respects the most complete study of an individual lake yet available in America. Not a small part in the collection of data bearing upon limnology has been taken by state organizations, by the establishment of temporary or permanent biological field stations (Ward ’98, Needham and Lloyd ’16) and the re- cognition of the state university as the natural centre of such investigation. On the other hand, while the results of investigation wherever conducted have at least some meas- ure of general applicability, it is unfortunate that the study of limnology has not proceeded along more uniform and better organized lines. The paucity of information con- cerning the water conditions of all the Canadian Provinces and certain of the States is in marked contrast to the ex- tensive information available in respect of certain States, notably Illinois and ‘Wisconsin. |What is chiefly significant, however, is the contrast between European and American efforts in general in view of the opportunities offered. If there is any one single outstanding fact concerning the im- portance of American water areas, it is the fact of the ex- istence of a mid-continental area of some 80,000 square miles represented by the Great Lakes of the St. Lawrence basin. Parts of a great international waterway, locally shared by one Canadian province, and eight states of the Union, the centre of an old established fishery, the Great Lakes constitute in all respects the outstanding fresh water area of the world. Had the example of Forel and of Zacharias been followed we should have had available on the Great Lakes either an extensively equipped limnological institution or at least some specific organization devoted to the study of the area in all its aspects. The question is not one of simply local concern, though it is evident from geographical considerations 8 BENSLEY: WATER AREAS OF ONTARIO that it is one in which the Province of Ontario is vitally interested. While various groups of aquatic organisms are directly or indirectly involved in questions of an economic kind, it is obvious that the fishes are the ones chiefly concerned. To what extent are they represented in Ontario? The commercially recognized fishes, including the more desirable species, lake trout, whitefish, ciscoes, tullibee, doré and sturgeon, as well as the coarser species, pike, carp, mullets and catfish, and the game fishes, principally speckled trout, small and large mouthed bass and maskinonge, are collectively a small, though very important part of the local fauna. The number of fish species, fresh water and marine, occurring in Canadian and Newfoundland waters, is stated by Hal- kett (13) to be 569. Evermann and Goldsborough (’07) place the number of fresh water species occurring in Canada as a whole at 145, representing 67 genera and 25 families. © In the first computation of the number occurring in Ontario (Wright ’92) reference is made to forty or fifty species, but the local records were at that time very incomplete, and the computation was doubtless based to some extent on the earlier reference compilation of Jordan and Gilbert (’83) later greatly extended and revised by Jordan and Evermann (96). The most complete index of Ontario fishes is that of Nash (’08), based partly on the foregoing compilation checked against collections from various points in the pro- vince. The total number is placed at 112, representing 69 genera and 26 families. Though this computation pre- cedes some other taxonomic work, notably that of Jordan and Evermann ('11), and though much is yet to be done in the matter of critical diagnosis of species from individual areas, it is probable that the number of species occurring in Ontario is in the neighbourhood of 125. While this re- presentation is fairly large as to species and apparently very inclusive as to families and genera as compared with Canada as a whole, it is probable that the number of species occurring at any one point is less than half the total. The number reported by Nash ('13) for the Toronto region is 49 and BENSLEY: WATER AREAS OF ONTARIO 9 by the writer (15) for Georgian Bay 48. These figures correspond fairly well to those given by Evermann and Clark (’20) for Lake Maxinkuckee (64) and other single areas (vol. I, p. 262). The total extent of water area in Ontario as estimated is in the neighbourhood of 80,000 square miles, probably a very conservative estimate when it is realized that a large part of the province is as yet only superficially surveyed. As compared with the total area of the province itself, 407,262 square miles, the submerged portion constitutes about one fifth. This relatively large extent of water area is accounted for in part by the provincial portion of the Great Lakes and by the occurrence within the province of the great Precambrian region, the rock depressions of which, left by the glacial period, now lodge an almost continuously con- nected series of overflow basins, ranging in size from the smallest ponds and muskegs to lakes of considerable or even of large size. Both geographically and from the point of view of their different economic associations, the Great Lakes and land-enclosed areas may best be considered separately. The former include as provincial waters the Canadian portions of Lakes Superior, Huron, together with Georgian Bay and the North Channel, St. Clair, Erie and Ontario, a total provincial area of some 38,000 square miles in extent, with some 1,500 or 2,000 miles of shore line forming the land boundary of the province on its south and west sides. The land-enclosed area, estimated at 41,385 square een eee eee ee Pee 'The figures reported are taken for the most part from the Atlas of Canada; published by the Department of the Interior, Ottawa, 1906. Those contained in the following table are from White (’05). Cf. also Map I. Physical Data, Great Lakes Ontario Maximum Lake Area Length Width Draina ge Depth Height sq. m1. m1. mt. Sq. mi. feet feet MEM oe lke. 7053 193 53 11,342 738 246 ae 9968 241 56 5,480 210 572 Se aes 503 26 24 41,160 180 575 MS see eek. . 22978 206 101 35,400 750 581 I i Soo See ss 32060 350 160 30,780 1012 602 10 BENSLEY: WATER AREAS OF ONTARIO miles, includes lakes and rivers distributed unequally over a land area which for convenience may be divided into five portions, each having a more or less definite facies, depending originally no doubt upon the geological foundation and the final effects of glacial action. These portions are:— (a) The south western peninsula, or that portion lying to the west of a line connecting the south end of Georgian Bay with the north shore of Lake Ontario to the west of Toronto. This portion comprises the older agricultural part of the province, is noteworthy for the paucity of water areas, and is underlain by strata of Silurian and Devonian age (cf. Map II)* covered by glacial soil deposits of consider- able thickness. It is also that portion of the province having the principal frontage on the Great Lakes. (b) A portion lying to the south of a line connecting the southern end of Georgian Bay with Lake Ontario in the region of Kingston. It forms the north shore of the larger part of Lake Ontario, contains some characteristic lakes, including Simcoe (300 square miles), Rice (27 square miles) and Scugog (39 square miles), is underlain by Cam- brian and Ordovician strata, and is transitional in many ways between the portion already described and the Pre- cambrian area to the north. (c) The northern Precambrian area of relatively great extent, including the Laurentian Highlands, and character- ized by its exposed igneous and metamorphic rocks with innumerable lakes distributed over its surface. The larger lakes include Nipissing (330 square miles) and Nipigon (1,730 square miles). (d) A north west portion containing the Lake of the Woods (1,851 square miles) and similar water areas lying beyond the height of land, and related both in aspect and origin (Lake Agassiz) to Lake Winnipeg and other lakes with the drainage system of which they are connected, though underlain by rock formations similar to the foregoing. *Prepared under the supervision of Professor W. A. Parks, University of Toronto. BENSLEY: WATER AREAS OF ONTARIO 11 (e) The Hudson Bay drainage area, a region character- ized by the broad coastal plain, and on the whole moderately inclined river basins of the Severn, Moose and Albany Rivers, and underlain in part with strata of Devonian age, an area hitherto, on account of its inaccessibility, only superficially considered in its relations to the province, but presenting as a matter of fact some 600 miles of provincial marine coast line. From the point of view of limnological investigation the Great Lakes offer almost unlimited opportunity. Except for the identification of aquatic organisms and, at least to some extent, their distribution, the most detailed information concerning the lake area refers to soundings and _ levels,! of importance in navigation, water supply and sewage disposal, of interest to municipalities, and practical matters relating to the fisheries? Information is desirable con- cerning a great variety of physical factors, depth and seasonal variation of water temperatures, suspended materials, dis- solved substances and absorbed gases, light penetration, substratum, as well as dynamic forces concerned in trans- portation of materials, water currents and circulation. Of especial importance are also environmental or ecological associations, in respect of which there are not only great local variations entering into the life cycle of organisms, but also marked general contrasts between individual lakes as complete environmental entities on a large scale. Thus even a superficial comparison of Lake Superior with Lake Erie in respect of their geographical position and physical characteristics reveals contrasts of geological formation, depth, transparency and temperature, the last-named 1Admiralty, Dominion and United States Charts, listed Department of Naval Service, Catalogue of Official Canadian Government Publications of Use to Mariners, Ottawa, 1920. Cf. Report of the International Waterways Com- mission, same reference; also Russell ('95), White (’15). “Reports of Dominion Department of Marine and Fisheries, Department of Naval Service, Commission of Conservation, United States Bureau of Fisheries; Ontario Department of Game and Fisheries, Geological, Natural History and Fish Commission publications of individual States. 12 BENSLEY: WATER AREAS OF ONTARIO difference doubtless involving several factors in addition to the more obvious ones of depth and maximum latitude difference amounting to some eight degrees. Furthermore the development of the Great Lakes is one of the most interesting chapters of glacial history, and one which will be found to bear very directly upon the origin and history of the fauna and flora, especially the origin of the northern fishes, their migrations, spawning periods, the effects of natural barriers and of natural and articifial means of com- munication. The study of the origin of the life of the Great Lakes is in fact one of the most interesting of general scientific questions. The significance of the provincial great lake area from the point of view of economic research is indicated both by the extent and value of the commercial fisheries and by the measures already adopted by the Dominion and Pro- - vincial Governments for the control of fishery operations and of the practical means of restocking. The annual value of the fisheries is subject to considerable variation, and it is necessary to examine the returns over a long period of years in order to form any real conception as to the possible relations to one another of the number of men engaged, gear, poundage of catch and probable supply. Taking the year 1918 as the last one for which figures for all the pro- vinces including Ontario are available,’ it is found for this year that the total value of the Ontario fisheries amounted to $3,175,111. As compared with the total for Canada of $60,250,544 this is approximately 5 per cent., a very creditable showing when it is realized that the latter figure includes the value of the Atlantic and Pacific marine fisheries, among the richest in the world. In relation also to the total for the fresh water fisheries of the Dominion, amounting in 1918 to $6,019,005, the Province of Ontario produced of this total an amount equal to 52.7 per cent., while the 1Fisheries Statistics, 1918, Dominion Bureau of Statistics, Ottawa, 1920. Cf. also Report of the Department of Game and Fisheries, Ontario, 1918; Toronto, 1919. BENSLEY: WATER AREAS OF ONTARIO 13 great lake fishery of the province alone produced 44 per cent. When it is considered that the Great Lakes are the mainstay of the commercial fishery, that this fishery has been culti- vated assiduously for very many years, and that it is first to be considered in all measures of Dominion or provincial legislation, it is evident that there should be available not only detailed information concerning the natural occurrence, migration, growth stages and dimensions, time of maturity, local variations of the spawning period, as well as concerning the natural associations and enemies of every species of direct market value, but also full particulars of all species directly or indirectly serving them as food. The land-enclosed water areas, though they at least equal, and in all probability greatly exceed the combined areas of the provincial portions of the Great Lakes, are so varied and irregularly distributed with reference to the topography of the country otherwise that they can scarcely be treated as having conditions in common. With the exception of the Lake of the Woods and Rainy Lake, which are boundary waters, they lie wholly within the province. They have a certain facies depending on their respective relations to the underlying substratum and rock formation, which in Ontario shows a marked contrast as between igneous and sedimentary, a contrast enhanced in every way by the extremes of soil formation and other effects of the glacial period. Some of the lakes as purely inland waters are of large size, notably the Lake of the Woods (1,851 square miles) and Nipigon (1,730 square miles), others though much smaller are nevertheless bodies of considerable magnitude, such as Rainy Lake (324 square miles), Nipissing (330 square miles) and Simcoe (300 square miles). Of the thousands of smaller lakes of perhaps less than 100 square miles in extent, some such as the Rideau waters are significant in the early development of the province, others are summer resort lakes of long standing, such as the Muskoka Lakes, while still others, like Lake Timagami are newer game fish areas of which hundreds have been made accessible within recent years through improvements of rail transportation. 14 BENSLEY: WATER AREAS OF ONTARIO Apart from any question of the utilization of the lake areas otherwise, as travel routes or for transportation of timber, the economic factors concerned are chiefly two in number, namely, in what way and to what extent are the lakes in general capable of utilization from the point of view of marketable fishes, and second, what restrictions .must be made, or practical means adopted, for increased utilization of the areas for game fishing, considering the great revenue to be derived from the presence in the province of the summer resident and sportsman. No limnological investigation of these collectively im- mense areas or indeed of any one of them has hitherto been attempted. Unlike the Great Lakes their waters have not been the object of hydrographic surveys even for determina- tion of depth. Except for ordinary geographical information, what is known about the physical character of the inland waters is chiefly a matter of canals and water power. Though > there have been a good many individual investigations dealing with particular groups of plants and animals, much more is known about the whole country from the standpoint of the traveller and sportsman than has been arrived at through scientific study. Every area which as a matter of development is subject to human interference, and this ‘nvolves the extreme of conditions in areas like the Muskoka Lakes, which are relatively of long standing occupation and therefore of game fish depletion, as opposed to others which with the improvement of means of ready access, always the deciding factor, are now beginning to be occupied, should be thoroughly known from the point of view of the factors entering into the conditions of existence within its limits. The limnological study of such areas, on account of their easy transitions from ponds and puddles to lakes of sufficient magnitude to present typical lacustrine con- ditions, ought also to yield important data concerning the general stages of lake formation in different situations, and the relation to this development of the groups of aquatic organisms. No comparative consideration of the annual yield of BENSLEY: WATER AREAS OF ONTARIO bS marketable fishes coming from the land-enclosed waters of Ontario is worthy of attention which does not first take into account the principles adhered to in framing legislation which would establish the respective spheres of the sportsman interested in game fish waters and the commercial fisherman. Recognizing the Great Lakes as primarily adapted for com- mercial fishing, and at least some of the inland northern waters as most attractive and eagerly appreciated by the game fisherman, the principle naturally became established in Ontario of the reservation and protection of inland areas from commercial fishing both in respect of sale and the opera- tion of ordinary types of gear. Fortunately in general this principle has been uniformly supported both in Dominion and provincial legislation. A first and very natural exception was made in the case of the Lake of the Woods and related waters, in part because of the predominance as in the Winni- peg area of fishes of a marketable type. A second exception of more recent origin was made in respect of Lake N ipigon and Lake Nipissing, though in general the inclusion of these areas was recognized as requiring close government super- vision. On the whole it will be observable both from the small values presented for the commercial fishes and the evidence of large sums of money brought into the country by summer residents and sportsmen that the element of the commercial fishery is only a small part in the total revenue accruing to the province from the existence of the land-enclosed area. The value of the fisheries of land-enclosed waters of the province was, in 1918, $503,858. This value amounts to about 16 per cent. of the total for Ontario as compared with 84 per cent. for the Great Lakes. Apart from the latter which give a decided advantage to the province, the total value of the fisheries of all land-enclosed waters of the Domin- ion amounted in 1918 to $3,347,600, of which the corres- ponding part of Ontario produced only 15 per cent. as com- pared with Manitoba’s contribution amounting to 55 per cent., while Saskatchewan almost equalled Ontario at 13 per cent. Furthermore, of the total for the land-enclosed 16 BENSLEY: WATER AREAS OF ONTARIO areas of Ontario more than half, or $285,169 was produced by the Lake of the Woods and related waters, which, as indi- cated above, are in a similar position to the Great Lakes as regards the commercial fishes. Lake Nipigon accounts for $128,647, or roughly one fourth, while the balance, amounting to only $90,042 includes the product of Lake Nipissing, valued at $32,294. The sum of the matter is that, excepting some four or more of the principal lakes, the com- mercial product for the province as regards the inland areas is negligible; the real significance of these areas lies elsewhere. So far as the economic aspects of the question are con- cerned the problems to be investigated in connection with the growth of fishes and related matters are exactly of the kind already outlined for the Great Lakes. But, in addition, the existence of a body of water both isolated in relation to other basins, and, because of its circumscribed boundaries, more naturally accessible to the commercial methods of fishing, gives rise to conditions which make the problem of balance and replacement more immediate. If the Great Lakes escaped total depletion before the advent of corrective measures, it was because the spaces were sufficiently vast in relation to the amount and efficiency of the gear employed to give the fishes most sought after some loop-hole of escape. In an enclosed area, even the size of Lake Nipigon, and the matter is more serious the smaller a lake becomes, it is evident that with continued fishing with modern gear, no large species, valuable or not, would have the least chance for survival, if not through the exercise of moderation, protection of small fish to sexual maturity, and practical propagation. More than this, the maintenance of the natural balance of all organisms is a matter of importance, whether as between predacious species and those feeding upon lower organisms, or as between spawn-destroying coarse fishes and others more desirable, or as between the various species of | minnows and similar small species and the larger fishes’ which depend on them for food. In fact an enclosed lake is a small and complete world in itself. Whatever happens in the way of modification of its living contents, practically | BENSLEY: WATER AREAS OF ONTARIO L7 in all cases a matter of human interference, the question of the balance of all organisms, and the balance between the poundage removed and that replaced by natural or artificial means is all important. Doubtless the closer study of con- ditions would also call into question the advisability of introducing into such area species not adapted either as young or fry to survive, or on the other hand give some point to the introduction of other species whose presence is desirable as marketable or game fishes or even to serve as food for such. Conservation of the game fishes is quite as much an economic problem as that of the commercial fishes. That it is not usually so recognized is perhaps on the whole a matter of gratification, since it is evident that the protection of game animals and the freedom of the great natural play- grounds are principles which from the beginning have been accepted by legislative authorities. On the other hand no attempt has been made to determine in what respects the invasion of the game country by the tourist and sportsman results in improved circulation of money and prosperity to a larger area, nor to compute what must be a very large amount expended by the non-resident, and contributing to the wealth of the province, if not directly to the provincial treasury through the sale of lands and licenses. It is evident on all hands that the matter of organizing this source of revenue, in a certain sense of capitalizing the natural re- sources, is one which is being seriously considered elsewhere in America and in Ontario is well worthy of attention. In the game fishes of Ontario we have a wide natural distri- bution of certain species, notably speckled trout and black bass, and also the occurrence in areas not open to commercial fishing of species, notably lake trout and doré, which deserve to rank as game fishes. The local abundance of these species in the smaller lakes, from most points of view of the game fisherman also the more desirable lakes, is a matter of funda- mental importance, since it is the deciding factor, or, if not, a very serious one, in establishing permanency of summer residence. The maintenance of the supply is also immediate 18 BENSLEY: WATER AREAS OF ONTARIO or urgent exactly in proportion as these lakes are frequented. Exact information concerning the complete life histories and the factors bearing upon their success or failure is es- sential for every occupied area. It is probable too that the communication of such information to the more permanent population would not only contribute to practical sentiment in the matter of conservation but also demonstrate the fact so generally neglected throughout the game country that the protection by the local inhabitants of the game animals is worth to them in money far more than their immediate utilization for food or sale. Another question with which scientific investigation will ultimately have much to do is that of artificial propa- gation. Already a hundred years ago, before the settlement of the country was yet solidly established, the better fishes of the Great Lakes were recognized as an important source both of food and profit. By 1850, after a period in which > the only restrictions upon the extent of capture were those imposed by primitive means of communication, the effects of excess and wastefulness were already evident. Not- withstanding the better organization of fishing administration from Confederation onwards (cf. Prince ’20), by 1880 de- pletion began to be evident in all the Great Lakes, and ques- tions of replacement became of increasing importance. Artificial propagation owes its inception in Ontario to Samuel Wilmot who began private hatching operations at New- castle in 1865, and who later as a Dominion officer established the entire hatchery system. The development of this system and the steady increase of the output from year to year’ has doubtless saved the commercial fishery of the Canadian portions of the Great Lakes from complete destruction. At the present time (cf. Map III) the Dominion system in- cludes eight hatcheries? in Ontario, the output of which is supplemented by that of hatcheries established by the 1Cf. comparative figures, Fisheries and Game, Report of the Commission of Conservation, Canada, Ottawa, 1911. 2Cf, Annual Reports on Fish Culture, Department of Naval Service, Ottawa BENSLEY: WATER AREAS OF ONTARIO 19 Provincial Government to the number of six,! while the general supply of the Great Lakes is enormously augmented by the operations of various United States hatcheries under the supervision of the Bureau of Fisheries. Fish propagation is a technical process, the more difficult because of its aquatic environment, and therefore demanding not only practical ability but also organization of method. The opportunity presented by a spawning period comes only once a year and if lost is irretrievable. The success of a single season’s operations depends on controllable elements of precise organization and on uncontrollable factors such as weather, scarcity of spawners, hatchery accidents and the like. If it were possible to effect some organization of scienti- fic work in relation to the hatcheries, an arrangement which would require considerable preparation, a contributory source of information would undoubtedly result which would be analogous in some ways to the usefulness of the agricul- tural experiment station to the practice of agriculture. Many sides of the hatchery question should be studied by investigators not bound to technical practice, having free time at the proper periods of the year, and provided with equipment adequate to the case; in other words, experimental hatchery work should be carried on with the same kind of equipment as that in regular use, but with additional facili- ties for scientific study. Bearing in mind that the object of propagation is not simply to make the fisheries hold out a few years longer, but to establish a balance between utilization and replacement, if not indeed an increased output, such as would be the aim of true pisciculture, it is evident that important sources of information are yet unavailable. The case of the hatchery and the commercial fishery is very much that of a mill in which the product is known but the extent or adequacy of the supply is unknown. In order to answer the question it will be necessary to know as nearly as may be practicable the percentage survival of fry and young at different stages and in localities differing both immediately 1Cf. Annual Reports, Department of Game and Fisheries, Ontario. Toronto. 20 BENSLEY: WATER AREAS OF ONTARIO and seasonally in respect of their physical characters. It will be necessary to know the true extent of depletion and of artificial increase by corrected figures indicating the poundage of the catch as modified by differences of the total amount of gear and its relative efficiency. There are, how- ever, other considerations. The recent developments of experimental embryology in relation to the behaviour and growth of aquatic eggs have shown to what extent these features are modified by physical differences in the surrounding medium, the constitution of the medium, absorbed gases and temperature. The investi- gation of such influences in its bearing upon the development of the eggs of desirable species under hatchery conditions would at least be of great scientific interest and most likely here and there of practical value. The same is true of the study of hybridization, the laws of which, under the influence of the recent work in genetics, are now much more clearly understood. In some of the fresh water fishes, notably Salmonidae, there are varietal, racial or environmental modifications which are almost baffling in their complexity, and which should now be thoroughly studied from a genetic point of view under experimental hatchery conditions. Finally the process of re-stocking which involves in part the utilization of young fish or fry, and to which reference has already been made, should be critically examined. While nominally a matter of assisting a natural replacement, the underlying idea is naturally expanded into that of the intro- duction into depleted waters of the same species from sup- posedly related waters; in which case it is a question whether the disturbance of the natural balance has not proceeded too far to ensure survival of the introduced stock, also whether as a matter of fact the physical conditions are similar, or if environmental differences are at the moment of introduction beyond the powers of adjustment of the organism. ‘The case is analogous when it is a matter of the introduction of foreign species. Artificial transfer to a hitherto un- occupied area may overcome for the introduced species a natural barrier, permitting it to attain a dominance over BENSLEY: WATER AREAS OF ONTARIO Pall native species perhaps more desirable, or it may readily happen that the desire to introduce a species of recognized merit from elsewhere overcomes the obvious reflection that the new environment is perhaps not suitable in any way, resulting in a process not very different in principle from liberating an infantile south sea islander on the shore of Hudson Bay. One of the recent pronouncements on this point is that of Evermann and Clark (’20, vol. I, p. 279). Concerning Lake Maxinkuckee these authors state :—" Four plants of lake trout aggregating 10,587 fish have been made in this lake. So far as we have been able to learn there is no evidence that any of these survived; there is no authentic record of the capture of a lake trout in this lake. If the physical and biological conditions obtaining in Lake Maxin- kuckee had been as well understood before the lake trout were planted, as they are now, those plants would not have been made.’ The records of failures as well as the some- times uninformed opposition to hatchery measures should be equally considered and a serious attempt made to bring together all facts and factors, practical and scientific, which bear upon the maintenance of natural balance in every ‘area of economic importance. LITERATURE CITED Adams, C. C., 1913. Guide to the Study of Animal Ecology. New York. Bibliography. Bensley, B. A., 1915. The Fishes of Georgian Bay. Con- tributions to Canadian Biology, 1911-1914, Fasciculus 2, Fresh Water Fish and Lake Biology. Biological Board of Canada; published as Supplement to Annual Report, Department of Marine and Fisheries. Ottawa. Birge, E. A., and Juday, C., 1911. The Inland Lakes of Wisconsin. Bull. Wis. Geol. Nat. Hist. Survey, 22, 27. 1911-1914. 22 BENSLEY: WAER AREAS OF ONTARIO Evermann, B. W., and Clark, H. W., 1920. Lake Maxin- kuckee, a Physical and Biological Survey. Department of Conservation, Indiana; publication No. 7. 2 vols. Indianapolis. Evermann, B. W., and Goldsborough, E. L., 1907. A foheele List of the qe Ry Fishes of Cana Proc. Biol. Soc. Washington. Forbes, S. A., 1887. The Lake as a Microcosm. Bull. Peoria Sci. Ass. Forbes, S. A., and Richardson, R. E., 1908. The Fishes of Illinois. Nat. Hist. Survey of Illinois; vol. 3, Ichthyo- logy. Urbana. Forel, F. A., 1892. Le Leman, Monographie Limnologique. 3 vols. Lausanne. 1892-1904. Jordan, D. S., and Gilbert, C. H., 1883. Synopsis of the Fishes of North America. U.S. National Museum, - Bulletin No. 16. Washington. Jordan, D. S., and Evermann, B. W., 1896. ‘The Fishes of North and Middle America. U.S. National Museum, Bulletin No. 47. Washington, 1896-1900. Jordan, D.S., and Evermann, B. W., 1911. A review of the Salmonoid Fishes of the Great Lakes, with notes on the Whitefishes of other regions. Bull. U.S. Bureau of Fisheries. Vol. 29, 1909. ‘Washington, 1911. Halkett, A., 1913. Check List of the Fishes of the Dominion of Canada and Newfoundland. Ottawa. Kofoid, C. A., 1903. Plankton of the Illinois River, 1, 2. Bull. Ill. State Lab. Nat. Hist., 6, 8. Nash, C. W., 1908. Check List of the Fishes of Ontario. Department of Education, Ontario, ‘Toronto. Nash, C. )W., 1918. Fishes; in Natural History of the Toronto Region. Canadian Institute, Toronto. Needham, J. G., and Lloyd, J. T., 1916. The Life of Inland Waters. Ithaca. Bibliography. Prince, E. E., 1920. Fifty Years of Fishery Administration in Canada. Proc. Amer. Fisheries Soc. BENSLEY: WATER AREAS OF ONTARIO a Reighard, J. E., 1894. A Biological Examination of Lake St. Clair. Bull. Mich. Fish. Comm., No. 4. Russel, I. C., 1895. Lakes of North America. Boston. Shelford, V. E., 1913. Animal Communities in Temperate America. Chicago. Ward, H. B., 1896. A Biological Examination of Lake Michigan in the Traverse Bay Region. Bull. Mich. Fish. Comm., No. 6. Ward, H. B., 1898. The Fresh Water Biological Stations of the World. Ann. Rep. Smithsonian Institution. Washington. Ward, H. B., and Whipple, G. C., 1918. Fresh Water Biology. New York. Bibliography. Wright, R. R., 1892. Preliminary Report on the Fish and Fisheries of Ontario. Ontario Game and Fish Comm. Toronto. Zacharias, O., 1891. Die Tier und Pflanzenwelt des Siiss- wassers. 2 vols. Leipzig. Zacharias, O., 1893. Forschungsberichte aus der Biologi- schen Station zu Plén, 1893-1903. Continued as Archiv fiir Hydrobiologie und Planktonkunde. UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 2 A STUDY OF THE CISCOES OF LAKE ERIE BY WILBERT A. CLEMENS OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO (REPRINTED FROM CONTRIBUTIONS TO CANADIAN BIo.ocy, BEING STUDIES FROM THE BIOLOGICAL STATIONS OF CANADA, 1921) TORONTO THE UNIVERSITY LIBRARY 1922 AStuUDY, OF THE, CISCOES?,. OF. LAKE ERTE By WILBERT A. CLEMENS, University of Toronto This study was carried out under the auspices of the Biological Board of Canada in response to a request from the Lake Erie Fishermens’ Association for an investigation of some of the problems in connection with the cisco fishing industry. In the request it was desired particularly that some information be obtained as to why smaller ciscoes in general are taken in the eastern end of the lake than in the western part. The major portion of the work having to do with the measurements of the fish and the taking of scales was carried out at various points on Lake Erie during the summer and fall of 1920, but shipments from various points were examined in Toronto during the years of 1919 and 1920. The author desires to express his appreciation of the assistance given by many fishermen, in particular by Mr. A. E. Crewe, who kindly provided accommoda- tion for the carrying out of the work during the summer of 1920, freely placed all the material of his catches for examination, and gave assistance in many ways. Other gentlemen who facilitated the work in supplying material and in other ways were: Messrs. Charles Ross, Roy Ross, Wilson S. McKillop, A. B. Hoover, C. W. Barwell, R. Kolbe, and W. D. Bates. IDENTIFICATION OF SPECIES For the separation of the species of shallow water ciscoes (subgenus T/hris- somimus) as described by Jordan and Evermannf (1911) it appears that three proportional measurements are more or less critical, namely, head in length, depth in length, and depth of caudal peduncle in head. Jordan and Evermann give the following proportions: *The word cisco is here used instead of herring for all members of the genus Leucichthys except for the tullibees, in accordance with the list of standardized names of North American fish as agreed upon by the U.S. Bureau of Fisheries, the Biological Board of Canada and the Canadian Fisheries Association. jJordan, D. S., and Evermann, B. W. 1911. A Review of the Salmonoid Fishes of the Great Lakes with notes on the Whitefishes of other Regions. Bull. U.S. Bureau Fish., Vol. xxix (1909). Document No. 737 (1911). i) ~I Species Head in length} Depth in length | Depth of caudal peduncle in head eae UREA OOD athena ste 4.33 4.3—4.6 3.0 ALC TE MOR: eo 2.9 1, onlaeies LEUSTA AO YORE TAN uA ULC: 4.5 3.7—4.2 oe Lede Pile BVH EC A NA me aa ie 4.4 3.5—4.0 Ne gos ES OR SeRStS Ga, Wetec men Miele clagisiag Lis 4 4.4 isi 3.3—3.5 ine 2.2 Accordingly for each fish examined the necessary measurements were made for the calculation of the above proportions. In addition the girth and the weight were determined and scales removed for age estimation. From June 14 to August 24, 1920, the ciscoes taken in twenty pound nets at the Crewe Bros. Fishery near Merlin were examined daily. In August and November the fish taken at Port Dover, Nanticoke, McKillop’s Fishery (near Port Maitland) and Dunnville were examined. The following species have been identified: (1) Leucitchthys sisco huronius (J. & E.), Lake Huron cisco. This species was readily distinguished by the long spindle-shaped body. The average proportions for 60 individuals were as follows: head in length 4.6, depth in length 4.8, depth of caudal peduncle in head 2.95. These figures are practically identical with those given by Jordan and Evermann for Lake Huron. This species is taken rather abundantly in the pound nets at Merlin but very few specimens were seen east of Long Point. (2) L. ertensts (J. & E.).. Jumbo cisco. This is the most abundant species taken in pound nets from Rondeau to Point Pelee. It also occurs in large numbers eastward to Long Point but appears to become very much less abundant beyond. It is noted for the large size attained as compared with the other species of the genus Leucichthys. The outstanding characters are (1) the deep body, (2) the more or less pronounced hump at the nape, (3) the deep caudal peduncle, (4) the relatively large scales. The average proportions for 150 individuals were: 4.41, 3.42 and 2.44. (3) L. artedi (Le Sueur). Lake Erie cisco or grayback. This species occurred in numbers at Merlin next in abundance to L. eriensis and appears to occur abundantly throughout the lake. It has been distinguished from the jumbo cisco by (1) the somewhat narrower peduncle, (2) the narrower body with usually little or no hump at the nape, (3) the smaller scales with less of the shiny appearance, (4) the much slower rate of growth as shown in the following table and also later in the discussion of the results of the scale exam- inations. 28 L. artedtz | L. eriensis Length Weight Age Date | Length | Weight] Age No. Date cm. Oz. Years|} No. Date c.m. oz. Years 600 July 8 20.7 6 Rane 934 July 8 20.8 ee a 3 ‘601 Ta ae oe a a eel se 602. 21.0 6 Ws). 937° 21.1 ian 3 603. 21.5 6 ce 938° 22.2 rine aieAe 604 21.0 6 5 239 22.6 8 aay 605 22 | 6 | 6 | 20 28 | 9] 2 606° 21.8 6 TRE a, July 9 23.7 AG CMD 607. July 9 22.8 vs bauer 949 26.5 ee a 609° 22.9 7 REG 943 21.5 ey 3 610 oe re eae ae ee Figure 1 shows a drawing of a scale from specimen No. 606, L. artedi and a drawing of a scale from specimen No. 235, L. eriensis. The average proportions for 50 individuals as they occurred at Merlin were 4.26, 3.7 and 2.86. These figures are somewhat different from those given by Jordan and Evermann and may be due in part to the fact that the young of L. eriensis are somewhat difficult to separate from this species, and in the selec- tion of the above 50 individuals rather extreme forms were chosen. There is an indication, however, that L. artedi is more closely related to the species of the other lakes than perhaps the figures of Jordan and Evermann show. (4) L. prognathus (Smith). Lake Ontario deep water cisco or longjaw. In both the pound nets and gill nets from Port Dover to Port Maitland a cisco occurs very abundantly whose exact identity and relationships have not been determined as yet. Dr. B. W. Evermann, to whom ten specimens were submitted for identification, refers them provisionally to the species prognathus pending further examination of these and additional specimens. The out- standing features of this form are the following: (1) the long mandible which usually projects beyond the upper jaw and in extreme cases almost hooks over it, (2) the relatively long bony snout, (3) the narrow caudal peduncle, (4) the shiny appearance of the scales, (5) the rather deeply forked caudal fin. Ina great many individuals the above characters are extreme as well as other features, as indicated by the following proportions, 4.0, 4.2, and 3.2. In other specimens the proportions are about as follows: 4.3, 3.75, and 2.85. The average for 148 individuals is 4.22, 3.88, and 2.85. However, Dr. Evermann states that L. prognathus varies greatly. Only a single longjaw was taken at Merlin during 29 the summer of 1920 on August 24, and it had the proportions 4.1, 3.3, and 2.8. A fisherman at Point Pelee has stated that he recalled having seen during one spring rather large numbers of small longjaws taken in the pound nets in that region. This would indicate a migration occurring during the winter or spring months when temperature conditions would be rather uniform throughout the lake. The longjaws examined at Dunnville and Port Dover early in November, 1920, were almost ready to spawn. Typically, members of the subgenus Cisco (Jordan and Evermann) are said to spawn in late summer but it would not be surprising to find the deep water forms in the shallower, warmer waters of Lake Erie spawning later than those in the other Great Lakes, especially in a mild fall such as occurred in 1920. Two females of L. johannae received from Wiarton, Georgian Bay, November 24, 1920, were found not to have spawned. The following table shows comparative measurements of certain characters of the longjaws in Lake Erie. Measurements are given in decimal fractions of body length. PORT MAITLAND PORT DOVER | 1032 | 1042 | 1037 oI eR! os Te | as | 25 | aes] 25 | 28 || 02s | ioe | 28 De ee TEAC Ur o | o Se cs 084) 079) 082 Eye SEM GM MN Cee Bait ay | 054| .058 060 oe | .06 | .055|| .066| .055| .065 ott bg eee eae Vek ma iid SEO EEK 063 “0631 .064 oes! os! .os7l| .058| .058| .058 NRE NY ha To | -08e| -oaal .09 | .08 | .087| .085|| 079) .08 oe eT eae nnn ene Snontitotacciput is. seer amie’. 52) Gd ak oe Gill TAKEES HR oie pee a iter 2 ae 45 a gg) Ta | a ae an Head in length UA Ce 3) Sane epee 4.0 |4.0 10. Wid 400k a ee ele Depth in length.:...+-----225--> it se yh hey 26 Ta ees ae oie a C.P. depth in head........-----++--: 3.2. |3.2°.13.2 "I3.0 ja. ae (5) L. harengus (Richardson). Georgian Bay cisco. A few individuals were taken which agreed in measurements and description with the Georgian Bay cisco. Jordan and Evermann report this species in Lake Erie and no doubt it occurs in small numbers. For purposes of comparison and for confirmation of the value of proportional measurements, specimens of L. ontariensis were obtained from Port Credit on Lake Ontario, and specimens of L. harengus from Wiarton and Midland on 30 Georgian Bay. The average proportions of 20 individuals of L. ontariensis were 4.5, 3.8, and 2.6. The average for 25 individuals of L. harengus were at and o. 1. The following table shows the measurements of typical individuals of the various species examined. Measurements are given in decimal fractions of body length. L. artedi \L.harengus| L. sisco |?L.ontart- L. ertensis|L.prognathus huronius ensts me Le 21 328 3 666 1026 1038 Se 24 22 21 .23 i 25 Degas. Beis 00), e 22 26 26 33 24 Caudal peduncle length. . it Di | J. ll .10 5 Bl | Caudal peduncle depth... .073 .074 . 084 085 095 .081 Be 062 051 054 057 057 065 - WE a re i" 057 053 051 055 .050 056 Say SPACE ses 155 > . 068 . 064 Tee . 068 . 067 . 065 Maxillary aS Ta See iM .079 074 066 081 .073 086 Bact EOVOCGIPUE. 2: 2: <.: Ri .16 .14 15 .16 15 im s1tt/ 1 to pectoral...... a sol -36 .35 36 36 a 33 ae to P-V distance.. Bi 2.25 2.2, 2.4 2.2 2.3 1.9 Pectoral length..... a ite .14 .14 .14 .16 .16 i 17 ae BBMOEME es Coe oops i ais 412 14 415 16 a 17 ‘er JISTS eae ¥ .14 1S 05 .16 wid Ry 18 ee 053 06 062 | 064 073 053 Anal LE ale | 88 .88 | 94 11 11 .13 Pa | 49 47 48 43 46 45 B tes BPRS de ss 4 cs 2 9-85-8 9-82-8 9-78-8 8-70-7 8-75-7 8-75-7 Podnia. She 4.2 4.6 i 4.6 4.3 4.4 ey 4.1 “ihe 4.7 4.5 3.8 3.5 3.1 4.2 cP. depth in head...... 3.2 3.0 | 2.7 2.8 2.4 3.0 1From Georgian Bay. *From Lake Ontario. 31 The following table shows proportions as given by Jordan and Evermann and those obtained for the ciscoes in Lake Erie with the exception of L. harengus and L. ontariensis. Jordan and Evermann Lake Erie Head in | Depthin |Depth C.P.| Head in | Depth in | DepthC.P. Species length length | in length length length in length 1. hie SAFARI AS ih 4.33 4.3-4.6 ao 4 14.2 4.3 3.1 Pe huronius........ eh 66 4.2-4.5 2.9 4.6 4.3 2.95 Lee Be MWeH anata a 4.5 3.7-4.2 2.66 24.5 3.8 2.6 ee a, 4.4 3.5-4.0 20-25 4.26 3.70 2.86 7 ee Be Nel nya 4.4 i 3.3-3.5 2.2 4.41 3.42 2.44 7 ORT a wa 4. AG 3.5 3.5 i 4.22 3.88 2.87 Fig. 1.—Drawings of scales of ciscoes. A, from specimen No. 606, L. artedi, showing 5 winter bands, the fish therefore being in its sixth summer. B, from specimen No. 235, L. eriensis, showing three winter bands, the fish therefore being in its fourth summer. 1From Georgian Bay. 2From Lake Ontario. 32 RATES OF GROWTH The scales were used in determining the rates of growth of the various species of ciscoes. The growth areas are usually well marked. Scales from approxi- nately the following number of fish of each species were examined: L. eriensis Toor ro SS 0SE RSE e eee p++ i =A STO Vv SH +) 10 15 20 25 30 35 40 45 Length in Centimeters Fig. 2.—Graph illustrating rates of growth of ciscoes in Lake Erie a=L. artedi, p=L. frognathus, 3=L. eriensis, sh=L. stsco hurontius. 30 E f Fig. 3 The length in centi- meters is from the tip of the snout to the base of the caudal fin; the girth just anterior to the dorsal fin. It is possible, therefore, that the In the majority of scales some of the wint The following table gives the data obtained 140; L. artedi 55; L. sisco huronius 55; L. prognathus 150. The results a bands were difficult to distinguish and there was evidence that in some cases at shown in Fig. 2. Considerable difficulty was experienced in estimating th curve for this species should lie to the left of the curve for L. erzensts. least one winter band was not recorded. for the three important commercial species in Lake Erie. rate of growth of L. sisco hurontus. shows the relation of age to weight. uedestsaGere He ‘ ae sates: HERE RE EEE Age in Years = L. artedi, p a Weight in Ounces 34 L. eriensts. Fig. 3.—Graph illustrating relation of weight to age of ciscoes in Lake Erie. prognathus, e Gog ¢’8T GIP II = Se ee 2a Se bela Seer: 4. GOP lees 2 2 Se G83 = etn GOP . 9 LT G68 ae SS es eee soe Sr SIT G°L1z 8 OT |.-$ 26 O'LT o’Lé eS ee OTT PIL 96 9k ee ZL 01 gSe Tor. 0°GE 0°9T g’ge ie 89 ¢'3 L 01 z || eo. GL € O01 G° Ss 6 Got 8 FI GSE © 1s £9 8°6 GS gg es 9°6 oi Gc'3 Pen €é1 62 ce eo | oF ¢'3 0°06 og oF aS ms oot | m2. GAL vit GZ PaaS GF 0's OL et tien = O'L 0°9T Sos O°L I 6. = G°0Z es eet | Sy Z 0ST [ae eee S eer = gees 9°9 0ST Pe = an 0'€ 0'8 a ee O's GL aa Sime! on | 2) Se ‘ul ‘Z0 = ee ‘ul = atin) “ul oe "Ww =. ‘ul ‘ZO a ate bee) | Saeay wy | wie | wsueq | wsue7 |] pny | asm | mBuey | Bue || pS | aySeM | wBueT | yuo | ady re ms | | eT nl snyjousord “'T UpajAD “'T sisuatda 'T The difference in weight between L. artedt and L. prognathus is partly due to the fact that the specimens of the latter species were examined chiefly in November and the females were then heavy with spawn. 35 SUMMARY 1. Three species form the bulk of the cisco catch in the Canadian waters of Lake Erie, namely, L. ertensis, L. artedi and L. prognathus. 2. L. eriensis is the dominant form westward from Long Point, and L. prognathus eastward from Long Point. This statement holds in general, for the former appears to prefer the shallower water while the latter is apparently a deep water form. However their ranges tend to overlap and their migrations at times take them into one another’s territory. For example, fishermen have reported occasional schools of longjaws as far west as Point Pelee, and, on the other hand, the jumbo is reported as abundant, at times, off Port Maitland. L. artedi occurs abundantly throughout the lake, but probably in greatest numbers west of Long Point. 3. L. artedi and L. prognathus have rates of growth and increases in weight which are practically identical, while ZL. eriensis increases about 1 1/3 times faster in length and two to three times faster in weight. 4. Examinations of the graphs and tables for rates of growth and increases in weight show that the optimum size for the taking of the jumbo cisco is from the fifth summer upward when they are at least 12 inches in length and 1 pound in weight. Whether the food supply would permit of this as the minimum size it is impossible to say. In regard to L. artedi and L. prognathus a minimum length of about 10 inches and a weight of about 6 or 7 ounces, when the fish ang in their sixth summer, would appear to be quite satisfactory. 5. Concerning the occurrence of smaller ciscoes in the eastern end of the lake, this much can be safely said: that in respect to gill net catches the fisher- men in the western portion of the lake secure a larger percentage of jumbo ciscoes and, therefore, get large fish, while the fishermen in the eastern end, particularly off Port Maitland, secure chiefly the smaller species, L. artedz and L. prognathus. The same facts apply to the pound net catches, with the addition that, since the young inhabit the shallow waters and the shallow water area east of Long Point is more limited, there appears to be a concentration of young ciscoes along the shore, particularly in Long Point Bay, and hence the young are apt to be impounded in large numbers in the pound nets. 6. No data were obtained as to the age when the various species spawn for the first time. Spawning is probably at the end of the third summer, and, if so, the six-ounce regulation protects the two species, L. artedi and L. prognathus in respect to being allowed to spawn once, but does not protect L. eriensis since it attains a weight of six ounces in its third summer. 7. The girth measurements were taken around the body just anterior to the dorsal fin, that is where the greatest girth occurs. The results show that the three inch gill net regulation is quite satisfactory for the species L. artedi and L. prognathus since they do not attain a girth of six inches until the sixth summer, but barely protects L. eriensis since this species attains a girth of six inches in three years. 36 8. In any undertaking for the artificial propagation of ciscoes in Lake Erie, at least for the region west of Long Point, particular attention should be given to L. eriensis, because of its rapid growth and its excellent qualities as a food fish. CONCLUSION This study has proved to be merely preliminary. The ciscoes of Lake Erie form a complex association and it has been impossible in this investigation to determine their inter-relations or to study the physical factors in relation to the various forms. Solution of the many difficult problems must await a thorough study of the physical conditions of existence in the various parts of the lake, such as distribution of temperatures, oxygen, carbon dioxide, currents, etc., and the relation of these factors to spawning, growth, movements of the fish, as well as to the production and distribution of their food organisms. 37 IV, yy rae iy DNV } mae ahs abs eAyl Tyr Nh rf veo mie enti Wy ne iA ‘ Sa V} party ik PNA Mi j i UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 3 THE FOOD OF CISCOES (LEUCICHTHYS) IN LAKE ERIE BY WILBERT A. CLEMENS AND N. K. BIGELOW OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO (REPRINTED FROM CONTRIBUTIONS TO CANADIAN BIOLOGY, BEING STUDIES FROM THE BIOLOGICAL STATIONS OF CANADA, 1921) TORONTO THE UNIVERSITY LIBRARY 1922 i LBBB BABA AAA AAA AAA THE FOOD OF CISCOES (Leucichthys) IN LAKE ERIE By WILBERT A. CLEMENS, AND N. K. BIGELow, University of Toronto. The results of the examination of the contents of the digestive tracts of 211 ciscoes (fresh-water herring) are presented herein. The bulk of the material was obtained early in June, 1919, and from July to November in 1920, from Lake Erie at various points along the north shore. The species examined were Leucichthys eriensis, the jumbo cisco; L. artedi, the Lake Erie cisco; L. sisco huronius, the Lake Huron cisco; and L. prognathus, the Lake Ontario deep water cisco (longjaw). These were taken at Merlin, Rondeau, Port Dover, Nanticoke, McKillop’s fishery (near Port Maitland), and Dunnville. In addition 19 individuals of L. harengus, the Georgian Bay cisco, from Wiarton, Georgian Bay, and 7 individuals of L. ontariensis, the Lake Ontario cisco, from Port Credit, Lake Ontario, have been examined for comparative purposes. The material from Merlin, Rondeau, Nanticoke and McKillop’s was obtained in pound nets while the material from all the other points was obtained in gill nets. The results are given in the following tables. In the table ‘‘ Unidentified species’’ are placed those fish whose identity was not determined. The figures indicate the relative abundance, namely: (1) that only a few individuals were noted; (2) that the organisms occurred rather abundantly; (3) very abundantly. SUMMARY. 1. An examination of the tables shows that the ciscoes are pre-eminently plankton feeders. The study practically covers the fishing season, and during that time at least, the free swimming crustacea form the bulk of the food of these fish. Of Canadian waters, Lake Erie produces more ciscoes than all the other Great Lakes combined. For example, in 1919 Lake Erie produced 7,425,713 lbs., while the remainder of the Great Lakes produced 4,022,711 Ibs. It is not improbable that the production of ciscoes is directly dependent upon the amount of plankton Crustacea produced. The numbers of these Crustacea which must abound in Lake Erie in order to support the millions of ciscoes, as well as the great numbers of white fish and young of many other species, is almost beyond the imagination. Comparative quantitative plankton studies ‘in the Great Lakes would, no doubt, afford considerable information as to the fish productive capacities of these lakes. 4] 2. It is doubtful if the various species of ciscoes show any preference among the entomostraca as food material. They doubtless take whatever forms occur in the waters they happen to inhabit. 3. In the great majority of alimentary tracts examined, Daphnia formed the great bulk of the contents, while other forms were represented by scattered individuals. In many cases Daphnia alone were present. This was particularly true of the jumbo and the Lake Ontario ciscoes. It appears, therefore, that Daphnia are very much the most important of the entomostraca as food organisms. Daphnia longispina occurred in all the material examined, as variety hyalina galeata. Daphnia ephippia were abundant in October in Lake Ontario and in November in Lake Erie. Occasional ephippia with three eggs were noted. 4. Of the Copepods Diaptomus sicilis and Limnocalanus macrurus were perhaps the most abundant forms occurring in the digestive tracts, although Epischura lacustris occurred frequently and occasionally in considerable numbers. Very often the oil globules of these forms gave the contents a bright red colour. 5. In the eastern end of Lake Erie one of the most important food organisms was Mysis relicta. As far as we are aware this is the first record of the occurrence of this form in Lake Erie. Its presence indicates at least an approach of con- ditions in the eastern end of this lake to conditions in the other Great Lakes. 6. Three individuals were found to have eaten small fish. In each case digestion had proceeded too far to allow of identification. All three ciscoes were taken in the eastern end of the lake, two were longjaws (L. prognathus) and the third, while not definitely identified, was probably also a longjaw. A fisherman near Point Pelee has stated that one winter he found that some ciscoes which he took through the ice, had eaten ‘‘ minnows.” 7. As is shown in the table for the longjaws (L. prognathus) these fish in June, 1919, had fed practically entirely upon Ephemeridae (Ephemera simulans), both adults and subimagoes. The importance of these insects as fish food is | thus further demonstrated. Moreover, there is no doubt that the transforma- tion of the nymphs to the subimaginal stage takes place at the surface of the water, as occurs in the closely related genus Hexagenia (Needham, 1920).* This means that the subimagoes, as well as the imagoes, were taken at the. surface of the water by the ciscoes. The projecting lower jaw of these forms is well suited to such surface feeding. 8. The following table, compiled from the food tables, shows the distribution of the food organisms in the lake. The outstanding points in the table are: (a) The absence of Mysis relicta from the western portion and the absence of Daphnia pulex and D. retrocurva from the eastern portion. Further investiga- tion, however, may show the presence of these species throughout the lake. (6b) Although only 48 gill net fish were examined, and the list of forms is, therefore, incomplete, yet the results are an indication of what would be expected in any large body of water, namely, that the shore waters contain a greater number of species of food organisms than the more open waters. The gill net *Needham, James G. 1920. Burrowing Mayflies of our Larger Lakes and Streams. Bull. U.S. Bur. Fish., Vol. XXXVI, 1917-18. 42 ae LL NN CE IM) a WESTERN PortTION EASTERN PorTION 87 pound net fish | 55 pound net fish: | 43 gill net fish from Merlin and from McKillop’s from Port Rondeau and Nanticoke Dover and Dunnville —, Epischura lacusiris.................. aapiomms seciles... 2... Limnocalanus macrurus.............. meer eda crysialiong.........2........ Diaphanosoma brachyurum........... Holopedium gibberum................ EE as “a “ec CS a ae ee Bosmina longirostris................. Eurycercus lamellatus ............... Be vtoraerare cies ly Ph Bepiodora kindtit................... oo & 3 y 3 8 ++ t+t+++4+4¢4+4+4+4+444 ++ zn ELS DNS) Sa + a ish were taken over 5 miles from shore while the pound net fish were taken vithin 2 miles of shore. (c) A comparison of the first two columns shows the possibility of there yeing a greater number of species in the western part of the lake than in the ‘astern end. 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DyMonpD OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 mE ROVISIONAL LIST OF ‘THE FISHES OF RAKE ERIE Considering the value of its fisheries, surprisingly little systematic study has been devoted to the fishes of Lake Erie. Our knowledge of the species occurring in the lake appears to have been gained incidentally in connection with investigations on some of the species of commercial import- ance and through surveys of neighbouring areas. Although Lake Erie is one of the most productive fishing grounds in the world, its productivity could no doubt be increased by proper management. Investigations with this object in view have been made from time to time, but for the most part these investigations have been confined to the species whose numbers it was desired to increase. No study of the problem as it affects the lake as a whole has yet been undertaken. When such a study comes to be made, information as to the species that occur in the lake, their distribution and relative prevalence will be of fundamental importance. The present list is intended to be a contribution towards such a study. It at least indicates how unsatisfactory is our present knowledge of the fish fauna of the lake, especially in Canadian waters. The list contains 91 species of fish and two species of lamprey. No species is included which has not been authen- tically recorded from Lake Erie. Except in the case of commercial species or those recorded in official reports, the authority for including it is given in the case of each species. Brief reference has also been made to the general range or habitat of the less common species found in the lake. In the preparation of this list, the publication ‘‘ Fishes of Ohio’’, Osburn (’01), has been found to be of great assistance. It includes not only the results of his own observations, but 57 58 DyYMOND: PROVISIONAL LIST OF FISHES also those of Rafinesque, Kirtland, Jordan, Henshall and others. Many species not elsewhere recorded for the lake are included as a result of Osburn’s observations in San- dusky bay in the summers of 1899 and 1900. I am also indebted to Professor Osburn for the identification of several specimens of Cyprinidae and of Cottus ictalops reported herein. . Other lists consulted contain only occasional references to the occurrence of species in Lake Erie. The expression ‘‘Great Lakes Region’’, often met with in describing the range of species, is unsatisfactory for the preparation of a list such as the present one. Even where a species is said to occur ‘‘in the Great Lakes’’, the reference is hardly less unsatisfactory, for the fauna of each of the lakes exhibits peculiarities of its own. Among the works consulted the following were found to be of most value: Jordan and Evermann (96), Forbes and Richardson (’08), Bean (’03), and Nash (08). Many other publications have been consulted but only those containing definite reference to the occurrence of species in Lake Erie are included in the list of literature cited. The basis of the present list was obtained by reference to two collections from Lake Erie made during the summer of 1920, one by Dr. ‘W. A. Clemens, University of Toronto, and the other by a party from the Royal Ontario Museum of Zoology. Dr. Clemens concerned himself principally with the ciscoes (lake herrings) but he secured specimens of all the species brought in by the fishermen. Most of his specimens, other than ciscoes, were taken at Merlin, Ontario. The Museum party was stationed at Point Pelee. Their primary object was to secure typical specimens of some of the chief commercial species from which to make casts and colour sketches for permanent museum exhibition specimens. Their specimens were also secured from fishermen. The species secured by Dr. Clemens were Ichthyomyzon concolor, Acipenser rubicundus, Lepisosteus osseus, Ama calva, Ictalurus punctatus, Ameturus nebulosus, Catostomus commersonit, Cyprinus carpio, Notropis rubrifrons, Hybopsis DyYMOND: PROVISIONAL LIST OF FISHES 59 storertanus, Hiodon tergisus, Dorosoma cepedianum, Coregonus albus, Leucichthys harengus, Leucichthys sisco huronius, Leu- cichthys artedi, Leucichthys eriensis, Leucichthys prognathus, Lucius lucius, Pomoxis sparoides, Ambloplites rupestris, Le- pomis pallidus, Eupomotis gibbosus, Micropterus salmozdes, Stizostedion vitreum, Stizostedion canadense griseum, Perca flavescens, Percina caprodes zebra, Roccus chrysops, A plodino- tus grunniens, Lota maculosa. The collection made by the Royal Ontario Museum of Zoology party included Ichthyomyzon concolor, Lepisosteus osseus, Ictalurus punctatus, Ameturus natalis, Ameturus nebulosus, Noturus flavus, Catostomus commersonu, Hybopsts storerianus, Hiodon tergisus, Coregonus albus, Pomoxts spar- oides, Stizostedion vitreum, Stizostedion canadense griseum, Roccus chrysops, Aplodinotus grunniens. During the spring and summer of 1921 a number of Lake Erie fishermen preserved for the Department of Biology specimens of the fish taken in their nets, especially the less common species. Mr. A. E. Crewe, of Merlin, Ontario, sent Ichthyomyzon concolor, Petromyzon marinus unicolor, Notropis rubrifrons, Hybopsis storertanus, and Cottus ictalops. Mr. W. D. Bates, of Ridgetown, Ontario, sent the following taken at Rondeau, Lake Erie: JIchthyomyzon concolor, Catostomus commersoniu, Moxostoma breviceps, Moxostoma aureolum, Hybopsis storerianus, Coregonus albus, Salmo gaird- nert, Pomoxis sparoides, Stizostedion vitreum, Perca flavescens, Roccus chrysops. From Mr. A. B. Hoover, of Nanticoke, Ontario, were received Carpiodes thompsoni, Catostomus commersoni, Moxostoma breviceps, Notropis rubrifrons, Hybopsis storerianus, Hiodon tergisus, Dorosoma cepedianum, Percopsis guttatus, Pomoxis sparoides, Ambloplites rupestris. The abbreviations used in the following pages refer uniformly to the following authorities :— Osburn (’01) “Fishes of Ohio”’ J. & E., Bull. 47 (96) ‘“The Fishes of North and Middle America.” Jordan & Evermann (’08) ‘‘American Food and Game Fishes.”’ 60 DyYMOND: PROVISIONAL LIST OF FISHES Jordan (’82) ‘‘Report on the Fishes of Ohio.”’ Forbes & Richardson (08) ‘‘The Fishes of Illinois.” Bean (’03) ‘Catalogue of the Fishes of New York.” Nash (’08) ‘Vertebrates of Ontario.” Clemens The collection made by Dr. W. A. Clemens from Lake Erie during the summer of 1920. R.O.M.Z. Party The collection made by a party from the Royal Ontario Mu- seum of Zoology stationed at Point Pelee during the summer of 1920. Crewe The collection received from Mr. A. E. Crewe as men- tioned above. Bates The collection received from Mr. 'W. D. Bates as men- tioned above. Hoover The collection received from Mr. A. B. Hoover as men- tioned above. The nomenclature adopted is that of Jordan and Ever- mann (’96) except in the case of Coregonus and Leucicthys, for which their later publication (’11) ‘‘ Review of the Sal- monoid Fishes of the Great Lakes’’ has been followed. ANNOTATED LIST OF SPECIES Ichthyomyzon concolor (Kirtland). Silvery Lamprey The only species of lamprey previously recorded from Lake Erie. Specimens secured by Clemens and R.O.M.Z. party in 1920, and by Crewe and Bates in 1921. Also recorded by Huntsman (’17) from Lake Erie. Petromyzon marinus unicolor (De Kay). Lake Lamprey A specimen, 21 inches long, was taken on November 8th, 1921 in Lake Erie at Merlin, Ontario by Mr. A. E. Crewe. DYMOND: PROVISIONAL LIST OF FISHES 61 Polyodon spathula (Walbaum). Paddle-fish ‘“‘A single example has been recorded from Lake Erie which it doubtless reached through the Wabash and Erie canal” Jordan and Evermann (’08). Common in the larger streams of the Mississippi valley. Acipenser rubicundus Le Sueur. Lake Sturgeon Less abundant than formerly but still of considerable commercial importance. Lepisosteus osseus (L.).!. Common Gar Pike Common. Lepisosteus platostomus Rafinesque. Short-nosed Gar Pike Rare. ‘Sandusky bay, one specimen’? Osburn (01). The specimen described by Bean (’03) is No. 3241 USS. Nat'l Mus. from Cleveland. Common throughout the Mississippi valley. Amaia calva L..1. Bowfin; Dogfish Common. Of little or no value as a food fish. Ictalurus punctatus (Rafinesque). Spotted Catfish Taken in larger numbers than any other catfish by the fishermen on the Canadian side of the lake, at least at Merlin and Point Pelee. ‘‘Taken most frequently [in our waters] in Lakes Erie and Ontario’ Nash (’08). Taken by Clemens and R.O.M.Z. party, 1920. Ranges south through the Mississippi valley and north at least as far as Winnipeg. Ameturus lacustris (Walbaum). Great Lake Catfish “Generally distributed throughout the Great Lakes and in deep rivers, but is more abundant in Lake Erie than any other of our waters’’ Nash (’08). Ameiurus natalis (Le Sueur). Yellow Catfish “In Lakes Ontario, Erie and Huron”’ Nash (’08). Com- monly taken in the nets off Point Pelee (R.O.M.Z. party). 1“ The time will doubtless come when thoroughgoing measures will be taken to keep down to the lowest practicable limit the dogfish and the gars—as useless and destructive in our productive waters as wolves and foxes formerly were in our pastures and poultry-yards.’’ Forbes and Richardson ('08). 62 DyYMOND: PROVISIONAL LIST OF FISHES Ameiurus vulgaris (Thompson). Long-jawed Catfish “Taken in Lake Erie’? Jordan (’82). ‘‘Occasionally taken in the Ohio river, but is more abundant in Lake Erie”’ Bean (’03). ‘‘Lower jaw more or less projecting; in other respects scarcely distinct from A. nebulosus, with which it may intergrade”’ J. & E. Bull. 47 (’96). Ameiurus nebulosus (Le Sueur). Common Bullhead Commonly taken by fishermen at Merlin (Clemens) and at Point Pelee (R.O.M.Z. party). Said by Bean (’08) to be the most abundant catfish in Lake Erie and its tributaries. Ameiurus melas (Rafinesque). Black Bullhead ‘Sandusky bay at Black Channel’’ Osburn ('01). ‘‘Gen- erally speaking, it is not distributed so far to the northward or eastward as our other abundant bullheads through the Great Lakes of Ontario, Erie and Michigan” Forbes and Richardson (08). ‘‘Variable, much resembles A. nebulosus but smaller, with shorter, deeper anal and especially shorter pectoral spines”’ J. & E. Bull. 47 (96). Leptops olivaris (Rafinesque). Mud Cat Osburn (’01) quotes McCormick of 1892 as follows, referring to the occurrence of this species in Lake Erie, ‘‘Quite rare; I have seen but one specimen fresh, though I have noticed heads on the beach.”’ A fish of the Mississippi valley and Gulf States. Noturus flavus Rafinesque. Stone Cat ‘‘Lake Erie at Sandusky, frequently thrown up dead on the beach by the waves; not noticed in Sandusky bay” Osburn (’01). Secured at Point Pelee by R.O.M.Z. party. Schilbeodes gyrinus (Mitchill). Tadpole Stone Cat Osburn (’01) reports having observed it to be common among decaying vegetation in shallow water in Sandusky bay in 1896. Schilbeodes miurus (Jordan). Mad Tom “Sandusky bay’”’ Osburn (’01). Carpiodes thompsoni Agassiz. Drum; Lake Carp ‘Lake Erie at Toledo” J. & E. Bull. 47 (96). ‘‘Common in Lake Erie’”’ Nash (’08). Taken by Hoover, 1921. ‘ DyYMOND: PROVISIONAL LIST OF FISHES 63 Catostomus catostomus (Forster). Northern Sucker ‘Quite abundant in Lake Erie’”’ Jordan (’82). ‘Occurs in the Great Lakes and northwest to Alaska in clear, cold waters. It is very common in Lake Erie’’ Bean (’03). Catostomus commersonit (Lacépéde). Common Sucker Common at Merlin (Clemens) and Point Pelee (R.O.M.Z. party). Bates and Hoover collections, 1921. ‘‘ Most abund- ant of all the suckers in Ontario waters’’ Nash (08). Osburn (01) speaks of it as ‘‘one of the commonest species’’, in Ohio. Catostomus nigricans Le Sueur. Hog Sucker Said by Nash (’08) to be found in Lake Erie. Erimyzon sucetta oblongus (Mitchill). Chub Sucker “Sandusky bay’”’ Osburn (’01). ‘‘Great Lakes region to Maine and the Dakotas, south to Virginia . . gradually passing southward into the typical sucetia.” ,. & E. Bull. 47 (96). Minytrema melanops (Rafinesque). Striped Sucker “Found in the Great Lakes and south . . iia\ dita Pennsylvania it is limited to Lake Erie and the Ohio valley” Bean (’03). Reported from Lake Erie by Nash (’08). Moxostoma anisurum (Rafinesque). White-nosed Sucker Reported by Jordan (’82) from Lake Erie as M. carpio. ““Great Lakes region; not very common, but widely dis- tributed”’ J. & E. Bull. 47 (’96). Moxostoma aureolum (Le Sueur).! Common Mullet; Redhorse “Sandusky bay”’ Osburn (’01). ‘‘Inhabits the Great Lakes and the region northward . . . . common in Lake Erie’’ Bean (’03). ‘‘ Formerly abundant in the waters of the Lakes from the St. Lawrence to Lake Superior, but owing to persistent netting during the spawning season it has become comparatively scarce’’ Nash (’08). Taken by Bates, 1921. “Until very recent years this has been recorded as two species, the short- headed, small-mouthed form as M. aureolum; and the more ordinary form as M. macrolepidetum duquesnit (Le Sueur). This matter is cleared up by Jordan and Evermann (Bull. 47, U.S. Nat. Mus.). It is very probable that some collectors have confused the short-headed form with M. breviceps (Cope).’’ Osburn (01). 64 DyYMOND: PROVISIONAL LIST OF FISHES Moxostoma breviceps (Cope). Short-headed Mullet ‘Abundant in Lake Erie”’ J. & E. Bull. 47 (96). ‘‘Seems to be confined entirely to Lake Erie so far as our province is concerned”’ Nash (’08). Taken by Bates and Hoover, 1921. Placopharynx duquesnit (Le Sueur) Osburn (’01) quotes McCormick of 1892 as follows, ‘‘Lake Erie, common with other mullets’’. A southern species. Cyprinus carpio L. Carp Very abundant, especially at Rondeau. Carassius auratus (L.). Goldfish Found by Turner (’20) along the shore of Middle and South Bass Islands, Lake Erie. Reported by fishermen at Point Pelee, Ontario, but no specimen taken. Pimephales notatus (Rafinesque). Blunt-nosed Minnow “Sandusky bay’’ Osburn (’01). Forbes and Richardson (08) say this species seems to find a satisfactory place of residence in streams of any size or lakes or ponds of any description. Semotilus atromaculatus (Mitchill). Creek Chub Essentially a creek species but recorded by Osburn (’01) from Sandusky bay. Leuciscus elongatus (Kirtland). Red-sided Shiner Ue speaking, a brook species though Dr. Kirtland, who described the species, records it from Lake Erie’’ Osburn (’01). Opsopeodus emilie Hay ‘Lake Erie” J. & E. Bull. 47 (96). Southern in general range. Abramtis crysoleucas (Mitchill). Golden Shiner Sandusky bay’’ Osburn (’01). Frequents sluggish waters. Notropis anogenus Forbes Taken at Put-in-Bay by Ward (’19) in connection with a study of fish parasites. Notropis cayuga Meek Osburn (’01) reports it as common in Sandusky bay. Notropis heterodon (Cope) ‘‘Sandusky bay’’ Osburn (’01). DyYMOND: PROVISIONAL LIST OF FISHES 65 Notropis blennius (Girard). Straw-coloured Minnow “Sandusky bay’’ Osburn (’01). Notropis hudsonius (DeWitt Clinton). Spawn-eater; Spot- tailed Minnow; Smelt “‘Lake Erie, near Sandusky, abundant’’ Osburn (’01). ‘‘Abundant in the Great Lakes’ J. & E. Bull. 47 (’96). “‘Abundant in the Great Lakes and at the mouths of the rivers opening into them’’ Forbes and Richardson (’08). Essentially a minnow of the larger rivers and lakes. Notropis whippli (Girard). Silver-fin “Sandusky bay’’ Osburn (’01). Characteristically a minnow of clear streams. Notropis cornutus (Mitchill). Shiner; Dace “Sandusky bay’’ Osburn (’01). ‘‘Almost everywhere the most abundant fish in small streams”’ J. & E. Bull. 47 (’96). Notropis athertnoides Rafinesque ‘““Exceedingly common in Lake Erie’’ Jordan (’82). “Through the Great Lakes . . . . It moves and feeds in large schools, thousands being frequently seen together near the surface’’ Forbes and Richardson (’08). Notropis rubrifrons (Cope). Rosy-faced Minnow “Sandusky bay and Lake Erie at Sandusky”? Osburn (01). Secured by Clemens at Merlin, 1920; also by Crewe and Hoover, 1921. ‘Delights in the clear waters of rapid streams’’ Forbes and Richardson (’08). Rhintchthys cataracte (Cuvier & Valenciennes). Long-nosed Dace Although fond of clear, swift waters, Jordan (’82) reports it as found in the tributaries of Lake Erie and even in the lake itself. Hybopsis dissimilis (Kirtland). Spotted Shiner “Lake Erie”’ J. & E. Bull. 47 (96). Great Lakes region, west and south. Hybopsis storerianus (Kirtland). Lake Minnow “Abundant in Lake Erie’”’ Jordan (’82). ‘‘Lake Erie near Sandusky’’ Osburn (’01). Common at Merlin (Clemens) and Point Pelee (R.O.M.Z. party) 1920. Taken by Crewe, 66 DyYMOND: PROVISIONAL LIST OF FISHES Bates and Hoover, 1921. ‘‘Lake Erie to Nebraska abundant in the larger streams, especially in lowa”’ J. & E. Bull. 47 (96). Anguilla chrysypa Rafinesque. American Eel “According to Kirtland the eel did not formerly inhabit the Lake Erie drainage, but if not, it has found its way there through the canals’? Osburn (’01). Hiodon tergisus Le Sueur. Moon-eye ‘Taken in considerable numbers in the fishermen’s nets. Of little commercial importance. Clemens and R.O.M.Z. party, 1920. Hoover, 1921. ‘‘Great Lakes and the Missis- sippi valley; north to Assiniboine river’’ J. & E. Bull. 47 (96). Dorosoma cepedianum (Le Sueur). Gizzard Shad Common. ‘Cape Cod to Mexico; abundant southward : permanently resident (var. heterurum) every- where’ in the Mississippi valley in the larger streams; also introduced into Lake Michigan and Lake Erie” J. & E. Bull. 47 (96). Clemens, 1920. Hoover, 1921. Pomolobus chrysochloris Rafinesque. Blue Herring; Saw- belly Originally confined to the Gulf of Mexico and Mississippi valley. ‘‘Introduced through the canals into Lake Erie and Lake Michigan”’ J. & E. Bull. 47 (’96). Coregonus albus Le Sueur. Lake Erie Whitefish ‘‘This species is the common whitefish of Lake Erie. It is very close to Coregonus clupeaformis, the whitefish of the other lakes, differing only in form and colour. Compared with the latter, the Erie whitefish has a smaller head, higher nape, more angular form, and the colour is almost pure olive-white, without dark shades or dark stripes along the back. The flesh is softer, containing more fat. All these differences may be correlated with the fact that Lake Erie is shallow, and its southern shore is fed by warm, shallow, muddy, or milky rivers’’ Jordan & Evermann (11). Leucichthys harengus (Richardson). Georgian Bay Cisco Clemens (’22) records this species as occurring sparingly in Lake Erie. DyYMOND: PROVISIONAL LIST OF FISHES 67 Leucichthys cisco huronius (J. & E.). Lake Huron Cisco “Occasionally enters Lake Erie’’ Jordan & Evermann (11). Clemens (’22) found it fairly abundant especially in western portion of the lake. Leucichthys artedi (Le Sueur). Lake Erie Cisco ‘“‘Abounds in Lake Erie especially in its southern parts Jordan & Evermann ('11). Clemens (’22) reports it as abundant in Canadian waters. Leucichthys eriensts (J. & E.). Jumbo Cisco “Inhabits especially the north shore of Lake Erie, where it is extremely abundant”’ Jordan & Evermann (’11). Clemens (’22) found it very abundant, particularly west of Long Point. On account of its great abundance, size and quality, the most important cisco of Lake Erie. Leucichthys prognathus (Smith). Lake Ontario Cisco; Long- jaw Clemens (’22) found this species very abundant east of Long Point. Typically inhabits the deep water of the eastern end of the lake. Leucichthys macropterus Bean A specimen remarkable for the development of its fins has been described by Bean (’16) as L. macropterus. Salmo gatrdnert Richardson. Steelhead Trout A specimen was taken at Rondeau, Lake Erie, on July 6, 1921, by Mr. W. D. Bates, of Ridgetown, Ontario. This species is propagated by the U.S. Fish Commission and has been introduced into Lake Superior. Cristivomer namaycush (Walbaum). Great Lake Trout The Annual Reports of the Department of Game and Fisheries, Province of Ontario, show that approximately two thousand pounds are taken annually from the Canadian waters of Lake Erie. It is found more especially at the eastern end of the lake. Umbra limi (Kirtland). Mud Minnow Osburn (’01) found it abundant in the ‘‘ Black Channel”’ in Sandusky bay. Usually met with in ponds and creeks with a soft muddy bottom. ” 68 DYMOND: PROVISIONAL LIST OF FISHES Lucius vermiculatus (Le Sueur). Little Pickerel Taken by Osburn (’01) in Sandusky bay. An inhabitant of the Ohio and Mississippi rivers and streams flowing into Lake Erie and Lake Michigan from the south. Lucius lucius (L.). Common Pike Still of considerable commercial importance in Lake Erie, although less abundant than formerly. A cosmopolitan species of the northern hemisphere. Lucius masquinongy (Mitchill). Maskinonge Nash (’08) gives the range of this species in Ontario as follows: ‘‘In the St. Lawrence about the Thousand Islands, in the waters of the Trent valley, Lake Scugog, Lake Simcoe, and many of our inland lakes, but I have no record of its cccurrence in any of the Great Lakes except Lake Erie and the Georgian Bay, where it is quite common.”’ Fundulus diaphanus menona (Jordan & Copeland). Killifish Osburn ('01) found it common in Sandusky bay. Eucalia inconstans (Kirtland). Brook Stickleback ‘““The Great Lakes from Ontario to Superior’’ Forbes and Richardson (’08). Percopsis guitatus Agassiz. Trout Perch According to Osburn (01), McCormick of 1892 found this species to be common in Lake Erie. Forbes and Rich- ardson (’08) report it as common in the Great Lakes but rare south of them. Nash (’08) says it ranges all through the Great Lakes and their tributaries north to Hudson Bay. Hoover took it at Nanticoke, 1921. A phredoderus sayanus (Gilliams). Pirate Perch Reported for Lake Erie by Osburn (’01) on the authority of Henshall in 1889. ‘‘Occurs in Lake Erie’’ Bean (’03). ‘Through the Great Lakes at least as far east as Lake Erie” Forbes and Richardson (’08). Apparently confined to the Great Lakes and southward. Labidesthes sicculus Cope. Silversides “Very abundant on sandy bottom in shallow water in Sandusky bay’’ Osburn (’01). ‘‘Found in Lake Ontario, DyMoNnbD: PROVISIONAL LisT oF FISHES 69 Lake Erie and the Detroit river’’ Nash (’08). ‘‘In all the Great Lakes’’ Forbes and Richardson (’08). A species of southern distribution. Pomoxis annularis Rafinesque.! Crappie Jordan (’82) reports it as rarely taken in Lake Erie. “Occurs rarely in Lake Erie’? Nash (’08). Pomoxis sparoides (Lacépéde). Calico Bass “Tn the Great Lakes in large numbers’’ Jordan (’82). Taken in 1920 at Point Pelee (R.O.M.Z. party) and at Merlin (Clemens). Bates and Hoover, 1921. See footnote under preceding species. Ambloplites rupestris (Rafinesque). Rock Bass “Sandusky bay’’ Osburn (’01). Forbes and Richardson (08) state that this species has been taken from Lakes Huron, Erie and Ontario, but that it lives by preference in clear waters flowing over a rock bottom. Clemens, 1920. Hoover, 1921. Chenobryttus gulosus (Cuvier & Valenciennes). Warmouth “Lakes Michigan and Erie seem to mark its most northerly distribution . . . . everywhere a fish of the bayous, mud-bottomed ponds, and lakes, and lowland streams’ Forbes and Richardson ('08). Lepomis megalotis (Rafinesque). Long-eared Sunfish “Sandusky bay”’ Osburn (01). “‘In Lakes Erie, Huron and Michigan’”’ Forbes and Richardson (08). Abundant southward. ‘Tt is worthy of note that in Chippewa Lake, which drains into the Ohio river system, this species (P. annularis) was found exceedingly abundant, but none of the next species (P. sparoides) were taken, while in Summit Lake, with very similar surroundings, but draining into Lake Erie, only P. sparoides was taken. These lakes are but a short distance apart, the former being near Medina, the latter at Akron.’’ Osburn (’01). “‘ A tendency to geographical separation is shown by the fact that annularis is the more abundant southward . . . . and sparoides northward—the latter, indeed, also ranging somewhat the farther to the north.’’ Forbes and Richardson (’08). 70 DyYMOND: PROVISIONAL LIST oF FISHES Lepomis pallidus (Mitchill). Blue Sunfish “Very abundant in Sandusky bay”’ Osburn (’01).‘' Occurs abundantly in some parts of Lakes Ontario and Erie’’ Nash (08). ‘In the Great Lakes from Ontario westward, ranging thence to the south and west’’ Forbes and Richardson (’08). Taken by Clemens at Merlin, 1920. Jordan and Ever- mann (’08) say that it is the sunfish of the lakes, whether large or small, but it is decidedly more abundant in the smaller ones. Eupomotis gibbosus (L.). Pumpkin Seed ' In Sandusky bay . . . .. itiis the most abundant sunfish’? Osburn ('01). ‘‘In Lakes Huron, Erie, Ontario and Champlain’? Forbes and Richardson (’08). Clemens took it at Merlin, 1920. Micropterus dolomieu WLacépéde. Small-mouthed Black Bass “Sandusky bay’’ Osburn (01). Less common in Lake Erie than the next species. Prefers running water. Micropterus salmoides (Lacépéde). Large-mouthed Black Bass ‘Sandusky bay’’ Osburn (’01). Clemens took it at Merlin, 1920. ‘‘Prefers lakes, bayous and other sluggish waters”? Jordan and Evermann (’08). Stizostedion vitreum (Mitchill). Pike Perch Very abundant and of great commercial importance. Three colour phases occur and have received distinctive vernacular names, viz. gray pickerel, yellow pickerel and blue pickerel. The significance of these colour phases is not understood. Stizostedion canadense griseum (DeKay). Sauger Abundant in Lake Erie; of much less commercial im- portance than S. vitreum. Perca flavescens (Mitchill). Yellow Perch Very abundant. One of the most important commercial species in the lake. DYMOND: PROVISIONAL LIST OF FISHES 71 Percina caprodes (Rafinesque). Log Perch ‘‘Lake Erie at Sandusky’’ Osburn (’01). To the north- ward this species is represented by the following variety. Percina caprodes zebra (Agassiz). Manitou Darter Said to be the common form in the Great Lakes. ‘‘Lake Erie at Sandusky’’ Osburn (’01). Clemens took it at Mer- lin, 1920. Cottogaster copelandi (Jordan). Copeland’s Darter According to Osburn (’01), Henshall of 1889 took this species in Lake Erie at Put-in-Bay. ‘‘Great Lakes region, from Lake Champlain to Lake Huron and south”’ J. & E. Bull. 47 (96). Cottogaster shumardi (Girard) ‘“‘It occurs also in the Great Lakes and has been reported from Erie and Michigan’’ Forbes and Richardson (’08). Diplesion blennioides (Rafinesque). Green-sided Darter According to Osburn (’01), McCormick of 1892 found this species to be not uncommon in Sandusky bay. Forbes and Richardson (08) report it ‘‘from Lakes Erie and Ontario’”’ and south. Boleosoma nigrum (Rafinesque). Johnny Darter “Sandusky bay’’ Osburn (’01). Typically a darter of the creeks and small brooks. Ammocrypta pellucida (Baird). Sand Darter “Taken also in the lake [Erie’’] Osburn (’01). Reported by Jordan and Evermann (’96) from Lake Erie to Minnesota, abounding in clear sandy streams. Etheostoma flabellare Rafinesque. Fan-tailed Darter Found by Turner (’20) along the shore of Middle and South Bass Islands, Lake Erie. Boletchthys fusiformis (Girard) “Rather common in shallow water in Sandusky bay’’ Osburn (’01). Roccus chrysops (Rafinesque). White Bass Abundant. A commercial species of minor importance. te DyYMOND: PROVISIONAL LiIsT oF FISHES A plodinotus grunniens Rafinesque. Sheepshead Quite an important commercial species, occurring fairly abundantly and growing to a good size. Cottus ictalops (Rafinesque). Miller’s Thumb Found by Turner (’20) along the shore of Middle and South Bass Islands, Lake Erie. Taken by Crewe at Merlin, 1920. Lota maculosa (Le Sueur). Burbot; Ling. Very abundant. In the Great Lakes region it is con- sidered of little value as food. Lake Erie fisherman destroy those taken in the nets because they believe it destructive to other fish. In some places it is esteemed as food. LITERATURE CITED Bean, T. H., 1903. Catalogue of the Fishes of New York. Bull. 60, N.Y. State Mus. Bean T. H., 1916. Description of a New Cisco from Lake Erie, ‘Proc: ‘Biol. Soc. "Wash., ‘vol! Xe) paseo. Clemens, W. A., 1922. A Study of the Ciscoes of Lake Erie. Contributions to Canadian Biology, 1921, No. IV, The Biological Board of Canada. Forbes, S. A., and Richardson, R. F., 1908. The Fishes of Illinois. Nat. Hist. Surv. Ill, vol. III. Ichthyology. Huntsman, A. G., 1917. The Lampreys of Eastern Canada. Ottawa Nat., vol. XX XI, pp. 23-27. Jordan, D. S., 1882. Report on the Fishes of Ohio. Rep. Ohio Geol. Surv., vol. IV. Jordan, D.S., and Evermann, B. W., 1896. The Fishes of North and Middle America. Bull. U.S. Nat. Mus., No. 47, 1896-1900. Jordan, D.S., and Evermann, B. W., 1908. American Food and Game Fishes. Doubleday, Page & Co., N.Y. DyMOND: PROVISIONAL LIST OF FISHES 73 Jordan, D.S., and Everman, B. W., 1911. A Review of the Salmonoid Fishes of the Great Lakes with notes on the Whitefishes of other Regions. Bull. U.S. Bur. Fish., vol. X XIX. Nash, C. W., 1908. Vertebrates of Ontario. Dept. Educ., Toronto. Osburn, R. C., 1901. The Fishes of Ohio. Special Paper No. 4, Ohio State Acad. Sci. Turner, C. L., 1920. Distribution, Food and Fish Associ- ates of Young Perch in the Bass Island Region of Lake Erie. Ohio Jour. Sci., vol. XX, pp. 137-151. Ward, H. B., 1919. Notes on the North American Myxo- sporidia. Jour. Paras., vol. VI, pp. 49-64. Ny hy th ty Wed he th vat Wy ‘i abs UNIVERSITY OF TORONTO SFUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 5 RATES OF GROWTH OF THE BLUE AND YELLOW PIKE PERCH BY F. B. ADAMSTONE OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 RATES OF GROWTH OF THE BLUE AND YELLOW PIKE PERCH (STIZOSTEDION VITREUM) IN, LAKE ERIE From material obtained during the summer of 1920, at Merlin, Ontario, on Lake Erie, a study has been made, under the direction of Dr. W. A. Clemens, of the rates of growth of the blue and yellow pike perch. These fish, together with the gray pike perch, are described as colour varieties of one species, Stizostedion vitreum Mitchell (Jordan and Evermann, 1898, American Fish Manual, 1903). This species is also known as the wall-eyed pike perch, pickerel or doré, the latter being sometimes restricted to the yellow variety. Apart from their difference in colour, these varieties are also distinguished by their great diversity in size. The blue is apparently the smallest, averaging less than one pound in weight, and occasionally attaining as much as five pounds. The yellow may reach 20 pounds and is often taken weighing from 5 to 10 pounds, and the gray, which is the largest, attains a maximum of 40 pounds, 10 to 20 being common. The specimens of blue and yellow pike perch obtained at Merlin agree in this divergence in size, but apparently none of the gray variety are caught there by fishermen. Measurements of length, girth, etc., were taken on the specimens of the blue and yellow varieties with a view to the later determination of their respective rates of growth. For this purpose scales from each fish were preserved for the determination of age. The scales were mostly taken from the middle of the sides of the body below the lateral line. In most cases the ear stones were also preserved for use in estimating age. By microscopic examination of the scales from a fish a fairly accurate estimate of the age can be made. Typical scales of both blue and yellow pike perch are shown in 77 78 ADAMSTONE: RATES OF GROWTH OF PIKE PERCH Figs. 1, 2, 3. From the illustrations it will be seen that the scales are ctenoid; that is, the exposed posterior portion is covered with a number of small spines or teeth. The anterior margin, which is embedded in a pocket of the epider- mis and is protected by the overlapping borders of those in front of it, has an entirely different appearance. Instead of being spiny it is fluted and the surface is marked by a large number of fine concentric lines, broken at intervals by radial lines, which pass inward towards the centre from the notches of the scalloped edge. The fine concentric lines are lines of growth, and each represents the addition of a ring a if Gp 2 z a ae Ae ie aes C ‘ go Aw a. << = t \ \ Fig. 1. Scale from a blue pike perch in its fourth summer. of material to the scale at the border where it is attached to the epidermis. At regular intervals on the scale the fine lines are laid down alternately closely set and more widely separated. The spacing is wide apart or close together according as the lines were added at a favourable or unfavourable season of the year. This arrangement of lines gives the scale the appearance of having wide light and dark bands when it is examined under the microscope. As shown in Fig. 1 the outer margin is marked by a light band and in the other scales examined a similar area was almost invariably present oo ADAMSTONE: RATES OF GROWTH OF PIKE PERCH 79 in this position. This apparently marks a summer’s growth since all the specimens were obtained in summer. The central portion of each scale is surrounded by a dark area which must necessarily represent a late summer’s growth and the first winter period, since the fish hatch in spring and the central portion of the scale would be formed during the first summer. This is shown by the illustration of the scale of the smallest blue pike perch in Fig. 2. By counting the number of dark bands present on a scale the age of the fish can be estimated. Thus, if there are three, as shown in Fig. 1, the fish must have lived three years and some fraction of the fourth summer. However, Fig. 2. Scale from a blue pike perch in its first summer. it is impossible to estimate the fraction of a year with any great degree of accuracy. Accordingly, in the present instance, the number of whole years has been determined and an average fraction added to each. Thus, assuming that in general these fish hatch in May, then fish caught in June of the following year would have completed, on the average, a growth period of one year and one month. Sim- ilarly, fish caught in July would have completed one year and two months, and fish caught in August, one year and three months. Therefore, the average growth periods of fish caught in June, July and August would be one year and two months. All the fish considered in this study were obtained during the months of June, July and August, and 80 ADAMSTONE: RATES OF GROWTH OF PIKE PERCH hence an average of two months has been added to the number of whole years indicated by the scale of any particular fish. Thus the estimated age of the fish whose scale is figured in Fig. 1 is 3 1-6 years and that of the yellow pike perch in Fig. 3 is 2 1-6 years. This arrangement, though quite ar- bitrary, is justifiable since the measurements of length, weights, etc., must also be averaged in the determination of the rate of growth. In a great many cases the ear stones of the fish were examined as well as the scales. These also show increase in le os (KG ———SS Fig. 3. Scale from a yellow pike perch in its third summer. size with age and alternate light and dark bands, much like the scales, but they are much more difficult to work with on account of their opacity. However, in a great many cases they could be used to check the results obtained with the scales. Blue Pike Perch The data obtained by measurements, and determination of age, of the specimens of the blue pike perch are given in the following table: ADAMSTONE: RATES OF GROWTH OF PIKE PERCH 81 No. Date Length Age in yrs.| Weight in Girth cm. inches OZ. in. 52 | Oct. 1920 7.4 2.9 Vu, | il July 7 15 BEG) 2% 2 3% 10 June 25 17 6.7 21% 3 334 1 June 24 17.2 6.8 214 214 356 if July 7 1820 tek 2% 2% 28 July 28 19.2 1S 3% 3 4% 48 Aug. 16 19.6 all a 3% 434 12 July 7 19.6 st ie 3 4% 15 | July 7 19.7 7.7 2 3 16 July 7 20.4 8 ui 3% 27 July 28 20.4 8 r 4 5 2 June 24 20.6 8.1 +3 + 43% 8 June 25 20:7 8.1 eh 4 4% ig | July 7 20.7 8.1 . 4 4% 13 July 7 21.0 8:3 x 4 4% 44 Aug. 16 SALE) 8.3 7 4 4% 38 | Aug. 13 21.0 ais - 5 456 32 July 30 22.0 SA < 4 434 30 July 28 22.1 8.7 v 5 5% 45 | Aug. 16 22.2 8.7 “- 5 4%, 46 Aug. 16 223 8.8 os 5 4% 37 Aug. 13 22.5 8.8 bs 6 5 34 July 30 22.8 9 a 5% 5% 19 | July 7 22.8 9 : 6 51% 14 July 7 23.2 9.1 o 6 5% 39 | Aug. 14 23.2 9.1 “ 7 51% 33 July 30 23.8 9.4 “ 6% 5M 29 July 28 23.8 9.1 5 6 5% 27 Aug. 16 24.5 9.6 . i 5% 43 Aug. 16 24.6 9.6 = 8 556 26 | July 28 25.2 9.9 7 8 556 31 July 28 25.5 10 4% 8 55% 40 Aug. 14 25.5 10 i 9 5K 20 July 7 26.7 10.5 9 5% 49 Aug. 16 D4 fai) 10.7 e 11 6% 26 Aug. 13 lial 10.9 7 12 6% 41 Aug. 14 27.6 10.9 °F 11 6% 25 July 28 28.0 11 5% 12 6% 21 July 7 28.0 il - 12 61% 9 June 25 28.6 inlets tod Lif 7% 42 | ‘Aug. 16 28.7 12/3 = 10 6% 4 | June 24 29.0 11.4 “: 17 7% 51 Aug. 16 29.6 LAYS 15 6% 50 Aug. 16 29.8 Ay, . 14 6% 6 June 25 3022 AED 6% ily 713% 5 June 24 30.5 12 a 17 7%, 22 | July 7 30.5 12 e 15 3 June 24 31.9 12.5 7% 17 7% if June 25 31.9 IPS ‘a 17 7% 23 July 7 33.0 13 3 16 7% ADAMSTONE: RATES OF GROWTH OF PIKE PERCH 82 AGE IN YEARS J 10 19 20 25 30 35 40 30 2 60 LENGTH IN CENTIMETRES Fig. 4. Graphs showing rates of growth of (a) blue pike perch, (6) yellowpike perch in Lake Erie. ADAMSTONE: RATES OF GROWTH OF PIKE PERCH 83 From the results given above the average length and weight, for each age, have been worked out and are given in the following table: LENGTH x Semen nee) et WEIGHT IN Oz. CENTIMETERS INCHES y% 7.4 2.9 2% 16.8 6.6 2% 3% 21.9 8.6 5 4% 26.4 10.4 10 5% 28.8 11.4 14 6% 30.4 L250 16% 7% 31.9 12.6 16% Using the averages of length and age given in this table, graphs have been drawn (Fig. 4a and 5a) showing the rate of growth of the blue pike perch. Yellow Pike Perch A similar determination of the rate of growth of the yellow pike perch has been made from the material collected for the purpose. The results of age determination for these specimens are tabulated below: No. Date Length Length Age Weight Girth cm. in. years Oz. in. 24 Aug. 16 14.7 5.8 2% 2 3% 20 July 9 15.1 5.9 " 2 21 atom 15.1 5.9 oe 2 13 June 25 15.6 6.2 2 12 ene) 16.3 6.4 . 2 38% 1 tae oe 16.8 6.6 or 2% 3% 11 St a 17.8 7.0 9 3 3% 2 cane! 18.2 7.3 <4 3 334 19 July 9 18.8 7.4 e 3 18 eae 18.8 7.4 ks 3 23 Aug. 16 21.0 8.3 * 5 434 14 June 25 21.0 8.3 re 4% 434 9 ey 21.0 8.3 i 4% 4 5 woe 21.0 8.3 > 4% 43 3 bbe 21.1 8.3 - 4% 434 22 Aug. 16 25.0 9.8 3% 9% 5% 4 June 24 28.6 11.2 Ks 11 6 7 «24 30.0 11.8 41% 16 6% 10 Lee 30.5 12.0 oa 14 6% 6 tee 30.7 12.0 eS 14 6% 25 Aug. 18 33.5 13.2 _ 1 lb. 6 7% 8 June 25 38.7 15.2 6% 2 Ib. 3 9% 16 BO 5.5 20.2 9% 4 lb. 14 12% 17 July 5 60.5 23.8 — on — c 3 oO iss) Oo > oo or i) or _ wf oO _ oO o — bo _ lor) AN 84 ‘¢ 31y SHONNO NI LHOIGM ‘dq eye] ur yosod ayid moyjad (¢g) ‘yos1ad ayid anyq (v) ‘ade 0} 7YS19M Jo uoTR]a1 Surmoys syderg ADAMSTONE: RATES OF GROWTH OF PIKE PERCH AGE IN YEARS ADAMSTONE: RATES OF GROWTH OF PIKE PERCH 85 The results given in the preceding table have been used to compute the following table of averages, from which curves illustrating the rate of growth of the yellow pike perch have been drawn in Figs. 4b and 5b. AGE YEARS LENGTH CM. LENGTH INCHES WEIGHT 2% 18.2 7.4 3% oz. 3% 26.8 10.6 10240 Ss 4% ShS2 1253 16% “ 6% 38.7 ses 2 bese Oz: 9% a as 20.3 aCe vil oh 12% 60.5 PANT Sane: > 13% | 63.5 25.1 tO) 12s A comparison of the two curves obtained brings out the fact that the rate of growth of the yellow pike perch is fairly uniform, whereas, in the case of the blue variety, there is a decided slowing up of the rate of growth at about the end of the fifth year. In both cases, however, there is a falling off in growth with age. In order to show this the lengths at the ages of 1, 2, 3 years, etc., have been obtained by interpolation on the curves, and from these figures the yearly increase in length is obtained. Age Blue Pike Perch Yellow Pike Perch Years | Yearly Yearly Length cm. | Increment cm. Length cm. Increment cm. 1 9.3 10.3 ies 2 16.2 ee 18 La 3 297 eg 24.4 aS 4 25.8 30 29.7 Ay 5 28.8 2.4 24.4 4.3 6 31.2 Ls 38.7 40 7 32.7 | ses Ao 3°8 8 46.5 afte = Se 9 50.2 3°3 10 53.5 24 a 56.9 39 12 60.2 33 13 62.5 +. In the case of the blue pike perch the decrease in rate of growth after the fifth year is not compensated by large 86 ADAMSTONE: RATES OF GROWTH OF PIKE PERCH increase in weight, as in the case of the yellow variety. Thus, in the fifth summer, the average weight of the blue is 10 oz., which increases to 16 1-3 oz., by the seventh summer, but this increase in weight is accompanied by a very small increase in length. In the case of the yellow pike perch, on the other hand, the slowing up of growth occurs about the end of the fifth year but the slow growth is more than compensated by the large increase in weight which then begins. From a consideration of the curves obtained, it would appear that the best time to take the blue pike perch is after the fifth year when they have obtained a length of about 28 to 30 cm. (11 to 12 inches) and weigh 14 to 16 ounces. Since the girth measurement posterior to the gill cover at this age is about 6 inches, this is approximately the size which would be taken in a 3-inch gill net. With the yellow pike perch, since they increase so rapidly in weight after the sixth year, it would appear that they should not be taken until they have reached a length of at least 15 inches and weigh approximately 2 Ibs. The study of the rates of growth of the blue and yellow pike perch shows that the former do not reach nearly so great a size as the latter, and amply confirms the opinion of fishermen that the blue are much smaller. Moreover, after the fourth year the rate of growth of the blue variety falls off very rapidly, whereas the yellow continue to grow uni- formly up to a considerable age. This peculiar difference in the rates of growth possibly indicates some basic physio- logical distinction between the two varieties which also possibly finds expression in their difference in colour. literature Cited Jordan, David S., and Evermann, B. W., 1898. The Fishes of North and Middle America, Bull. U.S. National Museum, Vol. 47, p. 1021. ; U.S. Commission of Fish and Fisheries, 1903. Articificial Propagation of the Pike Perch, Fish Manual (Revised edition) p. 165, Washington, D.C. UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 6 THE RATE OF GROWTH OF THE YELLOW PERCH (PERCA FLAVESCENS) IN LAKE ERIE BY W. J. K. HARKNESS OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 i i ui on pe ) ; 1 Wet yds Nay Be eK THE RATE OF GROWTH OF THE YELLOW PERCH (PERCA FLAVESCENS) IN LAKE ERIE As shown in the returns given in the annual reports of the Department of Game and Fisheries of the Province of Ontario, the catch of yellow perch (Perca flavescens Mitchell) in Canadian waters of Lake Erie has practically doubled in the past ten years. In 1919 the catch amounted to 1,096,935 pounds, valued at $87,755. In view of this increasing value of the yellow perch fishing industry in Lake Erie, it seemed desirable to obtain some definite information as to the rate of growth of this fish. The study was undertaken at the suggestion of Professor W. A. Clemens, and carried out under his direction. The material was obtained by Dr. Clemens, at the Crewe Bros. Fishery, Merlin, Ontario, during the summer and autumn of the year 1920. The age of the fish was determined by counting the seasonal growth areas on the scales. The scales found most satisfactory were those from the region about the middle of the body below the lateral line. An examination of a perch scale shows alternating light and dark areas. It has been taken for granted that the light area corresponds to a period of rapid growth which would take place during the summer months, while the darker area, where the lines are more crowded, represents a period of slower growth which no doubt occurs during the winter months. This inter- pretation of the areas on the scale is supported by the fact that the scales from perch taken in late summer show a distinct light area on the margin, while those from perch taken in early summer show a dark area on the margin. Figure 1 illustrates the characteristics of a perch scale, and the method used in age estimation. Table 1 shows the results of this study coérdinated. 89 90 HARKNESS: RATE OF GROWTH OF YELLOW PERCH ee e - AS SOS AS: . WON AN NS Fig. 1. Scale of a yellow perch at the beginning of its fourth summer. 91 HARKNESS: RATE OF GROWTH OF YELLOW PERCH Se ORE el ee a ee rie ees : 8°SI GL O'IT GPS FPS 6G v » AL 9°IT ere 8°01 G‘$Z P&Z Z's ¥ Teen) ¥'S OZ Z'6 9°9 6'6 C'1Z Ete Z's F a9 c'¢ 0°% Ls LG G's C61 PEST 0'ST ro iy POE. Pa GZ e'F Ig ea O'LT 8'9T 0'F% 8ST 1 See ei 0's 9°% t'P 9°9 O'FI PPI COP 0g Woke - 0°01 0 0 ewe | 0's z 0's PP eT i! ieak % ABIX 0} AVIA] uoljeatosqo Te9X 0} WIA} Zurjeurysy| JUIUTOINSeIT} “ON [e}OT Tequinyy WOrj WSeoIDUT) Aq aBeIaAy | sis ey} 07 [[® 1040 UOIF OSCOIDUT) AQ VBVIIAY | Aq 2BBLIIAY |JO adejuadIEg yenqoy pezyeuwnnsy] 1o14193s0d Soyoul ul poeyeuiys” o3V ysnf sayour yqsueT Aa ae ae ae ee 899UNO UI WYZIOM ul yyy uy [epnes jo aseq 0} ‘wo ul yyBuaT pourmexgy ys jo iaquiny ae ee ee ‘ada ANVI NI HOUAd MOTIAA sO HLMOUSD AO ALVa NO VLIVG ‘IT aIaVL RATE OF GROWTH OF YELLOW PERCH HARKNESS: 92 4 8 8 10 4 4 16 18 20 22 Fig. 2. Graphs showing rate of growth of yellow perch in Lake Erie. 24 26 HARKNESS: RATE OF GROWTH OF YELLOW PERCH 93 Fig. 2 is a graph showing the relation of age to length in centimeters, length in inches, girth in inches and weight in ounces. Embody (1915) gives the following table for the growth of yellow perch as compiled from data obtained on the fish growing in natural waters in the vicinity of Ithaca, N.Y. Length in in. Length in cm. Length in cm. to Age over all over all base of caudal fin 5 months 2-2.5 5-6.5 3.5-5 1 year 3-4 7.5-10 5.5-8 2 years 6-7 15-17.5 13-15.5 Spawn April Advanced fry May-June Fingerlings September The measurements given by Embody have been reduced to centimeters, and the last column shows the approximate lengths of these fish to the base of the caudal fin. These figures agree very closely with the data compiled from fish growing in Lake Erie as do also those of Pearse and Achten- berg (1920) who give the lengths of yellow perch taken in Lake Mendota, Wis., in the first summer as 2.9 cm. on July 7 to 6.1 cm. on August 24. If the fish are taken before they reach maturity and are thus prevented from spawning, the supply of that species will soon become depleted. This is amply exemplified by the disappearance of the trout from many of the Ontario streams, and the bass from the lakes. It is believed that the fish studied are a fair average of the total catch. By a study of table 1 and Fig. 2 it is seen that approximately 40 per cent. of all the perch caught were 2% years old averaging 14.0 cm. (6.6 in.) in length, 4.4 inches in girth and 2.6 ounces in weight. 24 per cent. of all the perch caught were 3% years old averaging 17.0 cm. (7.7 in.) in length, 5.1 inches in girth and 4.3 ounces in weight, an increase of 1.7 ounces over the 2% year old perch. 94 HARKNESS: RATE OF GROWTH OF YELLOW PERCH 18 per cent. of all the perch caught were 41% years old averaging 19.5 cm. (8.5 in.) in length, 5.7 inches in girth and 5.7 ounces in weight, an increase of 3.1 ounces over the 2% year perch, and only “5.2 per cent. of all the perch caught were 5% years old averaging 21.5 cm. (9.9 in.) in length, 6.6 inches in girth and 9.2 ounces in weight, an increase of 6.6 ounces over the 2% year old perch. Pearse and Achtenberg (1920), judging by the measure- ments made on individuals from a school of young perch which remained near the base of Picnic Point, Lake Mendota, during the summer of 1916, and by observations of the gonads of half grown perch at various seasons, believe that perch may become sexually mature in Lake Mendota at the end of two years of growth. No information was obtained as to the age when the yellow perch first spawns in Lake Erie waters, but it probably is the end of the third year. Yellow perch increase most rapidly in weight between the age of 3% and 5% years (Fig. 2). During these two years the perch in Lake Erie increase on an average about 5 ounces, which is more than their weight at 3% years of age (Fig. 2). It would appear from the data at hand, that no yellow perch should be caught which are less than 4% years old. In other words from the standpoints of conservation and of monetary returns this fish should be taken when about 8 to 10 inches in length. This would allow the fish to spawn at least twice, and this would tend to ensure an inexhaustible supply. This would also give the fisherman a larger supply of a much more satisfactorily marketable fish. Unless some protection is given there is danger of the yellow perch fishery in Lake Erie becoming depleted in a few years if the rate of increase in the amount of catch during past few years is maintained. HARKNESS: RATE OF GROWTH OF YELLOW PERCH 95 Literature Cited Embody, George C., 1915. The Farm Fishpond, Cornell Reading Courses, N.Y. State Coll. Agric., Cornell University, Ithaca, N.Y. Pearse, A. S., and Achtenberg, Henrietta, 1920. Habits of Yellow Perch in Wisconsin Lakes. Bull. Bureau of Fish. vol. 36, p. 297, 1917-18. Doc. No. 885, 1920. AU Baer UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 7 THE RATE OF GROWTH OF THE WHITE FISH (COREGONUS ALBUS) IN LAKE ERIE BY Joun H. Coucu OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 Pape hAteE. OF GROWTH OF .THE WHITE FISH (COREGONUS ALBUS) IN LAKE ERIE Fhe purpose of this investigation has been to obtain some definite information concerning the rate of growth of whitefish in Lake Erie. The study was undertaken at the suggestion of Dr. W. A. Clemens to whom the writer desires to express his appreciation of the kind assistance given. The specimens were procured from points along the north shore of Lake Erie (Kingsville, Merlin, Ridgetown and Nanti- coke) through the kindness of Messrs. B. Wescott, A. E. Crewe, W. D. Bates and A. B. Hoover. For purposes of comparison eight specimens of whitefish were obtained from Port Credit on Lake Ontario and two from Hudson Bay. The latter were collected by Rev. W. G. Walton on July 22, 1919, at Great Whale river. Identification The fish from Lake Erie are here referred to the species C. albus Le Sueur, while those from Lake Ontario to C. clupeaformis (Mitchell) following Jordan and Evermann (1911). By way of comparison detailed measurements were taken of three specimens from Lake Erie, numbers 101, 102 and 105; three from Lake Ontario, numbers 66, 67 and 68; and the two from Hudson Bay, numbers 131 and 132. All the specimens had been preserved in formalin and alcohol some time previous to the time the measurements were made. The results are shown in the following table: 99 RATE OF GROWTH OF WHITE FISH CoucH 100 a oe IT’ €1° €1° re 02° 1G" 61° 8T° st" : : GOT“ | SOT” 8ST" pa 61° 15 LT L0° L0° L0° CO™)-Sg0 ¢O° gO" G0" FO" FO" 60° 60° 60° 80° L0° [Ee Ets Le OT” Or° LZ" LZ° | GLZ° GS" £3" €Z" GS" | GES" 1G" 13° GZ 9% GZ 62 8Z ¥8 Sr €I ral! §I a! He PES | OE SIP AK] C&L Tél 2YVa)- 89 AVd NOSGNH TA ro 8T° Site ol” LT” LT” ST" ¢0° ¢0° v0} SEO" 80° 80° 60° Tike FG" 86° 6G" 1G" 96 06 L8 P8 al Il Il él OSE | c9OPF L9 99 OIAVLINO AVI ‘y8ua] Apoq jo suorjoe1y [eulOap se UaAIS sjUaUIOINSea, Gr 200 7) Sk i St" y ug] jeuy | a ee is Wy Cos f U JySIay [esi0g 1 a 2 ae ey) ie a Ug ySug] IAjaq OP 80 |RZe She ae y sug] [210}99q CO 0s SOF = “eb eLIXey S091 se0! “1 FO =| SO: Aq GDF) 80" i= OL Sb 60- yideap “g’9 SOF) S80" le. 60 20" yysugy ‘gq’ Re. 206=-c\ Vee 66. yidap Apog 06:—| vOE> | test (0a* peeHy 8% GZ 6 SIIYAeI [IID G8 18 G8 Sa[eOS rat v1 &1 shel [euy él IT €1 sAev1 [es10q G8E | 8ch | 8gg (wu) y38ua] Apog AY: GOT | eOt |} cor ‘ONuautIdeds a1aa AAV] LLL LC Coucu: RATE OF GROWTH OF WHITE FISH 101 The results for the Lake Erie and Lake Ontario fish agree closely with those given by Jordan and Evermann (loc. cit.) and Bensley (1915). For the Hudson Bay fish several slight though interesting variations appear. The caudal peduncle appears to be longer; the diameter of the eye greater; the length of the maxilla greater; the height of the dorsal greater; the lengths of the pectoral and pelvic fins greater, and the scales on the lateral line fewer. In spite of these differences there is a close resemblance to the Lake Ontario whitefish and it seems advisable for the present to refer these two fish to the species C. clupeaformis. Rates of Growth The rate of growth was determined by plotting curves between the age ascertained from the scales and the length and weight determined by direct measurement. The scales for determining the age were taken from the side of the fish, some from just below the anterior part of the dorsal fin, some near and including the lateral line, and some from just before the pelvic fin where the scales are large. The round even scales from the dorso-lateral region were found to be more satisfactory than those from the ventro-lateral region. The latter were larger but had radiate markings and ridges on them and the summer and winter areas were not so well defined. The vertebrae and otoliths of some specimens were preserved as a secondary means of determining the age but they were found to be much less reliable beyond three years of age. The method of determining the age from the scales is illustrated in figure 1. It is assumed that the areas with widely separated lines represent spring and summer periods when growth conditions are at their best. Conversely the closely spaced areas represent the winter months. In Fig. 1 the upper two scales (A and B) are from whitefish from Lake Erie aged two and five years respectively. The lower two (C and D) are from the whitefish from Hudson Bay, aged five and ten years respectively. All scales are enlarged to the same degree, i.e. fifteen diameters. The scales and likewise the fish themselves from Hudson Bay 102 CoucH: RATE OF GROWTH OF WHITE FIsH are smaller than those from Lake Erie, although the former are much older. This apparently furnishes a_ striking illustration of the effect of cold water on the growth of these fish. The rates of growth of the fish examined are shown in Fig. 2. The results indicate that during the first two or D Fig. 1. Scales from whitefish from Lake Erie and Hudson Bay. A and B from Lake Erie, third and sixth summers respectively. C and D from Hudson Bay, sixth and eleventh summers respectively. CoucH: RATE OF GROWTH OF WHITE FISH 103 40 -— GE IN YEARS, ad a ~ /0 20 30 40 50 Lenataw Centimerres. Fig. 2. Graphs illustrating rates of growth of whitefish from Lake Erie and Hudson Bay. O, whitefish from Hudson Bay. 0, . ““ Lake Erie. meek) “* Lake Ontario. 104 Coucn: RATE oF GROWTH OF WHITE FISH three years the fish grow quite rapidly in length, then gradu- ally the rate of growth lessens and the increase in length with age is much less noticeable. They do however, continue to increase in length until ten or twelve years of age and probably throughout their entire lives. The eight specimens from Lake Ontario are also shown on the graph although the number is too small to warrant a curve. However, they appear to have a rate of growth somewhat similar to that Oy ~ Aae tN YEARS HAC ae aan PEE EEE EEE EEE EEE EEE HEHE HEE EHH} / 2 fo) 4 ro) 6 vA WEIGHT iN Pounps Fig. 3. Graph showing rate of increase in weight with age, whitefish, Lake Erie. 8) of the Lake Erie fish. The two from Husdon Bay are also shown and an approximate curve projected through them. Here also it is strikingly demonstrated that the rate of growth in cold water is much slower than in such waters as Lake Erie. Since a fish is a ‘‘¢old-blooded” animal its body temperature in cold water is low, and it follows directly that its metabolic reactions are all depressed and hence that growth is retarded. CoucH: RATE OF GROWTH OF WHITE FISH 105 The fish were weighed at the same time that the scales were removed and the lengths determined. It is interesting to compare the rate of increase of length with age (Fig. 2) and the rate of increase of weight with age (Fig. 3). Since the conditions of the fish, such as the amount of fat and the development of the gonads, result in considerable differences in weight, the average of all the fish in each year was taken and this is used in this curve. It indicates that Per eae BE ES0 B80 BS SE SSSSRESReeeeeeeeee 2) oe SSESSeR Be SERSSREEE Cees BEE RE SEE SS ESE ERSRE ES Bee Sees eeeee a SaSgge 80 GSE CSeSRRGRUECEEEREe> | % fc Qs Ae <= z= tw i) rin GESR SESE ESEESS ARERR x oe rises BaaeEe: i 5 aneeene EEE RRER DES OF RRB RREHE SOS SSRREEE BESO s Tess eee ps Aocbssaaacessaaasesssaaeseeaaaeeeseseeeeeeaeeets fA ~! ry c?] ae yi 6 PA 6 Sidican IN Dearie Fig. 4. Graph showing relation between Iength and weight, whitefish, Lake Erie. the fish increase quite uniformly in weight up to four or six years of age, after which they increase rapidly relative to age and also to length. This is of the utmost importance from a commercial standpoint as the weight obviously bears a direct relation to food value. If left to the age of seven to ten years (40-50 cm.) the fish would have passed through that period when there is the greatest relative increase in weight. The wisdom of such a course is obvious. Another 106 Coucu: RATE OF GROWTH OF WHITE FIsH phase of this matter has been illustrated in Fig. 4 where weight and length have been plotted. In this graph each individual fish was recorded. It shows that at first the fish develops rapidly in length for a small increase in weight. Then at about 35 cms. the rate of growth in length decreases and the fish begins to increase in weight. This rapid increase in weight and comparatively slow increase in length continues at least up to ten years of age though probably soon after this age the rate of increase in weight falls off again. The curve corroborates the deductions drawn from Figs. 2 and 3. DATA—COREGONUS ALBUS—LAKE ERIE Speci- men Date Length | Girth Weight Age Sex No. cm. cm. No yy OPA Years 1 |May 1919 32.0 Vibs ye ioz 4 2 > 29.0 Ona 2 3 20.0 OO Be 1 ee . 25 June 6 37.0 Pam heme pt 5 Q very fat 26 va 46.5 tami | pave 10+ @ fat 27 . 33.0 1 pas 4 9 28 ve 34.3 Lats OKs 4 o fat 29 i 30.3 1 ay 3 of 30 as 34.8 teat wisi 6 of 3 a 18.5 10.1 1 32 i i re | 9.5 i) 33 i 15.7 8.8 1 34 s 15.6 8.8 1 50 |June 25/20] 16.8 10.1 OD raze 1 51 4 18.5 12.6 Ohad ini aay 1 Bo. dlejuly he (20.3. | 14e54 Oe eee 53 oa 16.1 be) OLy aoe 1 54 eyes 36.5 39.8 2 NO ie: 5 100 Dec. 6 39.6 Pe ian 6% Q large eggs 101 ae 42.8 4 ae 8 ae 9 ie) “se “ce 102 fi 55.8 Zo PO eee a Q spent 103 is 40 0 = Naslichiphe 40) 6% Q large eggs 104 vs 40.5 2 SAP ROW 7 2 spent 105 aM 38.2 PRESEN [aye 6 @ large eggs 106 es 39.3 Dien Mike on 7 o large testes 107 ‘4 40.6 Se mea th oe 7 3 partly spent 108 ae 39.6 2 ae 9 “ce 6 fot “se “ec 109 rt (has Tie | eae 6 of ‘% “f 110 i 40.5 yer anes 7% @ large eggs 111 * 38.1 Petes vbig Sie 7 o' fat 112 I 39.9 PARE tee Ub 8 o partly spent 113 > 41.8 Oh sakes 6% @ large eggs 114 t 37.8 p iaaat ik? > lire 6 @ partly spent 115 i 39.0 2 ON 4% Q A “@ 116 t 39.7 21, 10) ioe Qentirely “ 117 a 40.3 Ppt ates ono 2 7 OT ree 118 os 38.1 Deri h age ve 6 o partly spent 119 " 39.4 2 ee palutenes 6 @ large eggs 120 4 40.0 Mahe igs 4 Q@ spent CoucH: RATE OF GROWTH OF WHITE FIsH 107 DATA—COREGONUS CLUPEAFORMIS—LAKE ONTARIO No. Date Length cm. | Weight Age Years —S 1 3) ee | 61 Dec. 2, 1920 44.5 2% |b. 7% 62 ns 38.0 13% “ 4% 63 _ 39.3 Zee e 6% 64 . 37.0 Ca 5% 65 s 43.0 23% “ 7% 66 x 46.2 3 lb. 13 oz 7% 67 zh 39.0 ‘CN Se 4% 68 ie 41.8 Dh ean cities 6% DATA—COREGONUS CLUPEAFORMIS—HUDSON BAY No. Date [Length cm. | Age Years | 131 July 22 1919 | 30.2 10 Great Whale River 132 : 23.4 5 ag a iy Literature Cited Jordan, D.S., and Evermann, B. 'W., 1911. A Review of the Salmonoid Fishes of the Great Lakes with notes on the Whitefishes of other Regions. Bull. U.S. Bureau of Fish., vol. X XIX. Document No. 737, 1911. Bensley, B. A., 1915. The Fishes of Georgian Bay. Contrib. Canad. Biol. 1911-14, Fasc. II, Fresh Water Fish and Lake Biology. Biological Board of Canada, Suppl. 47th Ann. Rep., Dept. Marine and Fisheries, Fisheries Branch, Ottawa, p. 30-32. is ui H ‘ matt) r fh UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 8 REPRESENTATIVE CLADOCERA OF SOUTH-WESTERN ONTARIO BY N. K. BIGELOw OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 See en Let ed wh ee | REPRESENTATIVE CLADOCERA OF SOUTH-WESTERN ONTARIO During the past two years the author of this paper has made a special study of the Cladocera in various parts of the Province of Ontario. In view of their common occur- rence in practically all bodies of water and their fundamental economic importance, it is surprising that but little attention has heretofore been given to this interesting group in our province. Sars (1916) lists nine species found in samples of surface plankton collected during the summer of 1907 in the Go Home Bay region of Georgian Bay and submitted to him for examination by Professor E. M. Walker. The Cladocera of that list are as follows:— Holopedium gibberum Zaddack Sida crystallina (Miill.) Daphmella brachyura Liéven=(Diaphanasoma brachy- urum) Daphnia hyalina var. oxycephala Sars= (Daphnia longi- spina var. hyalina) Hyalodaphnia retrocurva var. intexta Forbes (Hyaloda- phnia = Daphnia) Ceriodaphnia scitula Forbes (probably =C. quadrangula Miill.) Bosmina longtrostris (Miill.) Polyphemus pediculus (L.) Leptodora hyalina Lilljeb. = (L. kindtit Focke) Birge (1918) mentions Chydorus latus Sars as “‘rare, Canada, near Lake Erie’’. Birge (1894) lists a number of Cladocera taken in the American waters of Lake St. Clair, Detroit River and the western end of Lake Erie. No doubt 111 12 BIGELOW: REPRESENTATIVE CLADOCERA these species also occur in the Canadian waters of these regions.” In the present paper the occurrence of forty-nine species of Cladocera is recorded. The classification and arrangement of these is based on Birge (1918). The material studied came from the following localities, which are indicated on the accompanying map (Fig. 1). 1. Georgian Bay. Several hundred samples of plankton collected during the years 1903, 1905 and 1907 from eight stations in the Go Home Bay region in the south-eastern part of Georgian Bay under the direction of Professor B. A. Bensley and Professor E. M. Walker were kindly placed at the writer’s disposal for examination. Also the stomach contents of 19 ciscoes (Leucichthys harengus) taken in the south-western part of Georgian Bay have been examined. These were obtained in November 1920 from Wiarton by Professor W. A. Clemens. In the locality records in the following part of the paper, Georgian Bay, unless otherwise stated, refers to the Go Home Bay region. 2. Port Sydney. This small village is situated on Lake Mary about 15 miles from the town of Huntsville in the Muskoka District. In the summer of 1919 plankton col- lections were made in various small lakes, ponds and pools within a radius of four miles of this village and also in the north branch of the Muskoka River. 8 Toronto. Plankton collections were made in Grena- dier Pond, in the lagoons on Toronto Island and in various small ponds and pools in the immediate vicinity of this city. These collections were made mostly in the spring and autumn of 1919 and the spring of 1920. 4. Lake Ontario. The stomach contents of seven ciscoes (L. ontariensis) obtained by Professor W. A. Clemens at Port Credit in November, 1920, have been examined. *Since the above was written an article has appeared: entitled “‘Notes on Canadian Entomostraca,”’ by A. Brooker Klugh, in the Canadian Field Naturalist, vol. XXXV, No. 4, pp. 72-73, 1921. In this paper twelve species of Cladocera are recorded from four localities in Ontario, all of which localities are other than those mentioned in the present paper. BIGELOW: REPRESENTATIVE CLADOCERA Mas 5. Bond Lake and Wilcox Lake. These two small lakes are situated about 20 miles north of Toronto. During the spring and autumn of 1919 and the spring of 1920 plankton hauls were made in these lakes as well as in several small ponds and pools in their immediate vicinity. 6. Lake Erie. The alimentary tracts of approximately 200 ciscoes from Lake Erie have been examined. Four species were represented, namely L. erzensis, L. artedi, L. sisco huronius and L. prognathus, and were obtained by Professor W. A. Clemens from various points along the north shore of Lake Erie in the spring of 1919 and the summer and autumn of 1920. Since ciscoes are pre-eminently plankton feeders, much interesting information concerning the dis- tribution of Cladocera was gained from this source. 7. Point Pelee. Samples of plankton were taken during the summer of 1920 from various types of permanent and temporary ponds and pools on this point of land, the southern- most point of Canadian soil. The drawings with which this paper is illustrated were all made from specimens collected in the various localities mentioned above. SIDIDAE Sida crystallina (Miiller) Georgian Bay, Lake Erie, Point Pelee. This species appears to seek the shallower water close to shore especially in the vicinity of aquatic plants and was not found in small ponds or pools unless they were immediately adjacent to larger bodies of water. It is a rather important form from the standpoint of fish food since it ranked third in the list of organisms most commonly found in cisco stomachs, Daphma and Leptodora respectively exceeding it in frequency. Forty-eight jumbo ciscoes taken from June 1 to August 2, 1920, at Merlin chiefly, but also at Rondeau, Lake Erie, had eaten it in considerable numbers. 114 BIGELOW: REPRESENTATIVE CLADOCERA Latonopsis occidentalis Birge Point Pelee. Fairly common June 18, 1920, in a small weedy pond near Lake Pond. Latona setifera (Miiller) Georgian Bay, Point Pelee (Lake Pond). Apparently uncommon in Georgian Bay since only a single specimen was taken. Five specimens were found in July and August in plankton swept from aquatic vegetation in Lake Pond (Point Pelee). Diaphanosoma brachyurum (Liéven) Georgian Bay, Lake Erie, Point Pelee. In some samples of plankton this species was fairly common and a few speci- mens were observed in two cisco stomachs (L. eriensis) from Merlin in July and August 1920. Diaphanosoma leuchtenbergianum Fischer Georgian Bay, Point Pelee. Less common than the preceding species. HOLOPEDIDAE Holopedium gibberum Zaddach Georgian Bay, Lake Erie. This species occurred very abundantly in most of the open water plankton from Go Home Bay and a few were taken from a cisco stomach (L. eriensis) from Lake Erie, July 1920. DAPHNIDAE Of the members of this family the genus Daphnia is the most widely distributed and the most abundant in point of individuals. Their immense numbers, their great economic importance as fish food and their amazing variations among individuals of the same species can only be fully appreciated by those who have made a study of fresh water plankton and the contents of fish stomachs. In a great many ciscoes BIGELOW: REPRESENTATIVE CLADOCERA 115 examined the entire alimentary tracts were found to be literally crammed with the remains of countless thousands of Daphnia. Of 205 of these fish examined 160 had eaten Daphnia to a greater or less extent. In the majority, Daphnia were by far the most abundant food organisms present and in 26 formed the entire food material. The only ciscoes which had not fed largely on Daphnia were some taken in the spring of 1919 when Ephemeridae formed the bulk of the food material, and a few others taken in the deep open waters where Mysis relicta and Limnocalanus macrurus were the dominant crustaceans. Daphnia pulex (de Geer) Port Sydney, Toronto, Lake Ontario, Bond Lake, Lake Erie, Point Pelee. Generally common everywhere in bodies of water of all sizes from large lakes to small temporary pools. However it was not found in the Georgian Bay plankton which is surprising in view of the large numbers of samples taken. Neither was it found in any of the cisco stomachs from the eastern portion of Lake Erie. It appears to be the commonest species in the western part of this lake, however, and apparently forms the bulk of the food of the ciscoes there during summer and autumn at least. The majority of the jumbo ciscoes (L. eriensis) taken during the summer and autumn of 1920 had eaten this species together with some Daphnia retrocurva which may be only a form of D. pulex, and all the individuals of this species taken during November 1920 had eaten D. pulex only. Daphnia retrocurva Forbes Georgian Bay, Lake Ontario, Lake Erie. Abundant in the plankton from Georgian Bay and very abundant in the cisco stomachs from Lake Erie. It appears to be a summer form in Lake Erie as ciscoes taken at Rondeau in June 1919 had eaten almost as many individuals of this species as of D. pulex. Many of these specimens showed astonishing variations in the shape and extent of the cephalic crest as shown in Figs. 1-4. One individual with a pronounced 116 BIGELOW: REPRESENTATIVE CLADOCERA vetrocurva crest had the same type of pecten on the claws of the post abdomen as is found in typical D. pulex. Forms approaching the D. arcuata type were also found. In spite of the amazing differences between typical specimens of D. retrocurva and D. pulex the indications point strongly to the conclusion that the two forms may be extreme vari- ations of one species. It is not improbable that a careful study of these forms at Rondeau during the month of June might establish their identity. Daphnia longispina (Miiller) Georgian Bay, Toronto, Lake Ontario, Bond Lake, Lake Erie. Very abundant, ranking next to D. pulex in numbers. It is an exceedingly polymorphic species. In the smaller lakes and larger ponds var. hyalina form typica was the most common phase. Form typica was found in Bond Lake and Grenadier Pond (Toronto), but in the latter body of water two remarkable variations occurred in May 14, 1920, as shown in Figs. 5 and 6. One had a protuberance over the region of the heart as is found in D. pulex var. minnehaha, while the other had a minute downward projecting crest on the head. In any large collection of Daphnia longispina it has been found that variations in form are not uncommon. In the larger lakes form galeata seems to be the pre- vailing form of this species. In his list of the Entomostra- ca of Georgian Bay, Sars mentions the Daphnia from Go © Home Bay region as D. hyalina var. oxycephala although at first inclined to consider it as D. galeata. The results of the present studies would indicate that the form he was dealing with was D. longispina var. hyalina form galeata according to the generally accepted nomenclature of the present time. It is much the commonest species in the southern part of Georgian Bay in the plankton and cisco stomachs examined. It occurred in nearly all of the plankton samples in varying numbers and abundantly in all of the stomachs of the nineteen Georgian Bay ciscoes (L. harengus) examined, along with Diaptomus and Epischura. Five out of seven Lake Ontario ciscoes (LZ. ontariensis) taken at Port BIGELOW: REPRESENTATIVE CLADOCERA LZ Credit in November 1920 had eaten this species almost entirely. In Lake Erie form galeata of D. longispina is the prevailing form but is much less abundant than either D. pulex or D. retrocurva. Jumbo ciscoes from Merlin, Ontario, taken during July and early August 1920 had eaten this species almost entirely. In Figs. 7,8 and 9 are shown some extreme variations such as are occasionally found scattered among ordinary speci- mens of this species. Simocephalus vetulus (Miiller) Toronto (Grenadier Pond), Bond Lake, Point Pelee. Fairly common in the above mentioned localities among aquatic vegetation and doubtless widely distributed. Scapholeberis mucronata (Miiller) Toronto, Lake Wilcox, Port Sydney. Commonest in small pools in these localities but also found in shallow water along the margins of lakes. Scapholeberis aurita (Fischer) Point Pelee. Found only about the middle of July 1920 in a small muddy pool in a pasture where it occurred spar- ingly in a plankton composed mostly of Certodaphnia reticu- lata and Moina affinis (Fig. 10). Ceriodaphnia reticulata (Jurine) Toronto, Bond Lake, Point Pelee. Only found in small pools or ponds but very abundant when found (Fig. 11). Ceriodaphnia lacustris Birge Georgian Bay, Toronto (Lagoon on Toronto Island). Not common. Ceriodaphnia quadrangula (Miiller) Georgian Bay, Point Pelee (Lake Pond). This is probably the Ceriodaphnia scitula mentioned by Sars in his list of Entomostraca from Georgian Bay. 118 BIGELOW: REPRESENTATIVE CLADOCERA Ceriodaphnia pulchella Sars Point Pelee. Taken during the months of July and August 1920 in some rather large ponds. Moina affinis Birge Point Pelee. This cladoceran was very common during the month of July 1920 in a small muddy pool ina pasture, along with Ceriodaphnia reticulata and a few Scapholeberis aurita. Most of the specimens were covered with a minute unicellular organism, a few of which are shown attached to the post abdomen in Fig. 12. This species also occurred in a few temporary rain pools along a roadway. BOSMINIDAE Bosmina longtrostris (Miiller) Georgian Bay, Toronto (Grenadier Pond), Bond Lake, Wilcox Lake, Lake Erie, Point Pelee. This is one of the most widely distributed of all our Cladocera. Some speci- mens from Go Home Bay appear to have the spine on the posterior ventral extremity of the valves longer than is usual in this species. This fact was noted also by Professor G. O. Sars in his examination of Go Home plankton collected in 1907. Bosmina longispina Leydig Port Sydney. The only specimens identified with cer- tainty as belonging to this species were two taken June 18, 1919, from the north branch of the Muskoka River. Eventu- ally this species may prove to be but an extreme variation of the preceding one as some specimens of Bosmina from Go Home Bay have been found to be intermediate in many respects between the two. MACROTHRICIDAE Macrothrix rosea (Jurine) Point Pelee (Lake Pond). A few specimens were taken in plankton swept from among weeds, August 22, 1920. BIGELOW: REPRESENTATIVE CLADOCERA 119 Illiocryptus spinifer Herrrick Toronto. Two brilliant red specimens of this species were found in December, 1918, creeping about among a thick growth of Chara in a small stream near Grenadier Pond, Toronto. Illiocryptus sordidus (Liéven) Point Pelee (Lake Pond). Common in August, 1920, in plankton swept from aquatic vegetation. The post abdomen of one of these specimens is shown in Fig. 13. Some individuals were very light yellow in colour and others were quite transparent. Acantholeberis curvirostris (Miiller) Georgian Bay, Port Sydney. Occurred sparingly in plankton collected from the shallower parts of Go Home Bay, but very abundantly in a small pond in the middle of a sphagnum bog situated about four miles south of Port Sydney. Drepanothrix dentata (Eurén) Georgian Bay. Apparently rare since it occurred in only a few of the many samples (Figs. 15-16). Streblocerus serricaudatus (Fischer) Bond Lake. Only one small specimen taken in plankton, November 9, 1919. Ophryoxus gracilis Sars Georgian Bay. Not uncommon in plankton collected from shallow waters (Fig. 14). CHYDORIDAE Eurycercus lamellatus (Miller) Georgian Bay, Toronto (Grenadier Pond), Bond Lake, Lake Erie, Point Pelee. Common in the weedy parts of ponds and lakes. 120 BIGELOW: REPRESENTATIVE CLADOCERA Camptocercus rectirostris Schoedler Georgian Bay, Bond Lake, Point Pelee (Lake Pond). Very common, November, 1919, in Bond Lake. Kurzia latissima (Kurz) Georgian Bay, Toronto (Grenadier Pond), Point Pelee. Only a few specimens were taken in each of these localities. Acroperus harpae Baird Bond Lake, Point Pelee. Especially common in Nov- ember, 1919, in Bond Lake. Acroperus angustatus Sars Bond Lake. Apparently this species is only a form of the preceding species since many forms intermediate between the two were taken on November 9 and 15, 1919, in Bond Lake. Leydigea quadrangularis (Leydig) Georgian Bay, Toronto (Grenadier Pond), Point Pelee. Only one or two specimens were found in each of these localities (Fig. 17). Alona guttata Sars Toronto (Grenadier Pond and also in smaller ponds and pools), Bond Lake, Point Pelee. Very abundant. Alona affinis (Leydig) Georgian Bay, Bond Lake, Point Pelee (Lake Pond). Occurred most abundantly in the plankton from the shallow weedy parts of Go Home Bay. Graptoleberis testudinaria (Fischer) Georgian Bay, Toronto (Grenadier Pond), Bond Lake, Point Pelee. Only a few specimens were found in each of these places. BIGELOW: REPRESENTATIVE CLADOCERA pwA Pleuroxus denticulatus Birge Port Sydney, Toronto, Bond Lake, Point Pelee. Never found in open water plankton. Pleuroxus trigonellus (Miiller) Port Sydney, Toronto (Grenadier Pond), Bond Lake, Point Pelee (Lake Pond). Several specimens were taken in November, 1919, in Bond Lake, but only one or two in each of the other localities. It appears to be an uncommon though widely distributed species (Figs. 18-19). Pleuroxus striatus Schoedler Point Pelee. Not uncommon during the summer of 1920 in certain ponds. Pleuroxus procurvatus Birge Toronto (Grenadier Pond and also smaller ponds and pools), Bond Lake, Wilcox Lake, Point Pelee. The only plankton in which this species was abundant was swept on June 1, 1919, from very shallow water along the margin of Wilcox Lake. Chydorus globosus Baird Georgian Bay, Toronto (Lagoon on Island), Point Pelee (Lake Pond). Only a single specimen was found in each of these localities (Figs. 20-21). Chydorus sphaericus (Miiller) Georgian Bay, Port Sydney, Toronto, Bond Lake, Wilcox Lake, Point Pelee. Most numerous in small weedy pools near Toronto. Variety punctatus was the only form of this species present in some plankton taken April 17, 1920, in Bond Lake. Chydorus faviformis Birge Georgian Bay, Bond Lake, Point Pelee (Lake Pond). Most abundant at Point Pelee, but only a few specimens were found in any of these localities (Fig. 22). 122 BIGELOW: REPRESENTATIVE CLADOCERA Chydorus barroisi ? (Richard) Toronto. Although this is supposed to be a southern species, a Chydorus which must be either this or an undescrib- ed species closely related was taken on October 19, 1919, in a small weedy pool near Bloor Street and Clendenan Avenue, Toronto. It showed the sharp teeth on the keel of the labrum distinctly but apparently lacked the spine on the postero- ventral margin of the shell. It appeared in company with a great many typical Chydorus sphaericus and a careful search not only through this sample of plankton but through collections made from the same pool many times since has failed to disclose another individual (Fig. 27). Anchistropus minor Birge Georgian Bay. Four specimens of this peculiar Clado- ceran were found in some plankton taken in August, 1905, in weedy shallow water (Figs. 23-24). Monospilus dispar Sars Georgian Bay, Point Pelee (Lake Pond). Occurred only in plankton swept from among weeds and not very abundantly (Figs. 25-26). Alonella excisa (Fischer) Port Sydney, Bond Lake, Point Pelee. Not very com- mon. Alonella nana (Baird) Port Sydney, Bond Lake, Point Pelee. This beautiful striated species was even less common than the preceding one. POLYPHEMIDAE Polyphemus pediculus (L.) Georgian Bay, Port Sydney (Muskoka River), Point Pelee. Fairly common. The specimens taken in June, 1919, from the north branch of the Muskoka River had a peculiar coloration. The tips of the last two pairs of legs as well as the ventral side of the caudal process were of a deep brilliant blue which contrasted strongly with the extreme transparency of the rest of the body. ———— BIGELOW: REPRESENTATIVE CLADOCERA 123 LEPTODORIDAE Leptodora kindtit Focke Georgian Bay, Lake Erie. Common enough in Lake Erie to constitute a most important item of food of ciscoes. Forty-three jumbo ciscoes (L. eriensis) taken from July 6 to August 2, 1920, had eaten Leptodora very largely along with a few Daphnia and Sida. In Georgian Bay Leptodora occurred in plankton from the deep open water. Some specimens taken September 12, 1905, contained the immature stages of some Trematode. In each case the parasites were quite conspicuous as they were in the very transparent part of the creature’s body lying alongside of the digestive tract where they contrasted strongly with their surroundings. It may well be that they were the early stages of a cisco parasite. The table that follows summarizes the distribution of the Cladocera as given in the preceding pages. It is evident from this table that the Cladoceran fauna as herein recorded is far from exhaustive for any of the local- ities studied but taken as a whole it may be considered representative of the south-western portion of Ontario. It is seen to be identical or nearly so with that of the northern portion of the United States as might be expected from the similarity of conditions in the two regions. During the course of these studies many puzzling forms were encountered especially in the family Chydoridae. For example, forms related to Chydorus, Alona, Alonella and Pleuroxus combining characters found in the anatomy of two or more of these genera were occasionally seen and the writer feels convinced that many surprises await the future student of these Cladocera. 124 BIGELOW: REPRESENTATIVE CLADOCERA Georg- | Port Te Wilcox Point ian Bay| Sydney |Toronto|Ontario| L. |L. Erie} Pelee Sida erystallina.). 4... ./.5 >: ot. ve ae Latonopsis occidentalis ..... ae Eatona serijer@aeeee i. 1) + a Diaphanosoma brachyurum . + zt 4 vy leuchtenbergianum + aA. Holopedium gibberum.......- si Daphnia pulex...........-- + an ae as a a ‘\ reirocurud......+-: ss ae at ‘\ longispima........ a +- ug a Be Simocephalus vetulus........ a a 2s Scapholeberts mucronata. .... ++ ae ae Scapholeberis aurita. . aa Certodaphnia reticulata . 4+ 4 ba JAGUSETIS.... )..« + + 4 quadrangula...| + a ih pulchella...... ni Motta afinis aceite. sss be Bosmina longirostris........| + ae a + a ti longispina........ + Macrothrix rosea.......---- a Illiocryptus spintifer. . Baha a8 sordidus.......- a Acantholeberis curvirostris...| | + + Drepanothrix dentata. . Seal) SF Streblocerus serricaudatus. ... - Ophryoxus gracilis.......... a Eurycercus lamellatus....... se + at 4 a Camptocercus rectirostris..... 3 Be + Kurzia latissima........-+- + + a2 Acroperus harpae.......-.-- ue | ¥ angustatus....... ae Leydigea quadrangularis.....) + + ae VA Ona REUULL LCE es aye dee + be ut POMEL NTT RSE) 4 22 = + a Graptoleberis testudinaria... . + as = aS Pleuroxus denticulatus...... =F + oe ~ . trigonellus....... + + 2 ay. oi SINGOLUS Woe = Sale bl: os ne procurvalus...... + + ae Chydorus PIOVOSUS. |. «' ar + ae sphaericus........ oe ar + + =e i‘ faviformis.........) + + ae Ni HITS, oc) SCR Paes is BORROUSI Cae tists oki e\ss =f Anchistropus minor......... + Monospilus dispar.......... + ; + Alonelia excisd............. + + Hs SON PECTS aap sesh 31. 3-0s Uso + + =f Polyphemus pediculus....... + + “bh Leptodora kindtit........... + + *Recorded by Birge ‘‘near Lake Erie.” BIGELOW: REPRESENTATIVE CLADOCERA 125 LITERATURE CITED Birge, E. A., 1894. A report on a collection of Cladocera mostly from Lake St. Clair, Michigan, 7m A Biological Examination of Lake St. Clair, by J. E. Reighard. Bull. Mich. Fish Commission, No. 4, Appendix I], p. 45. , 1918. The Water Fleas (Cladocera) ix Fresh Water Biology, Ward and ‘Whipple, New York, p. 676. Sars, G. O., 1916. Entomostraca of Georgian Bay, in Contrib. Canad. Biology 1911-1914, Fasc. II, Fresh Water Fish and Lake Biology, Biological Board of Canada. Suppl. 47th Ann. Rep., Dept. Marine and Fisheries, Fisheries Branch, Ottawa, p. 221. : ; ; i WG my, y vi y WW oe —— py fbr Mh WADE Nh } ain) VN tn 10) WAN ik , Ma ¥ { Hai Ni yi) SP ‘i \ \ t i ii ii K ty 4 Rh () Ni WGA ‘ —— ——= a ————— yee ae Sate adda nae eG “2 aS x/0 fort Sydiey £ AKE Go-Home Baylx ° Vv 60 Or y birarlorn \V26 =} HURON "Bons tene be fy LAKE Toronto, 7 NTA Trl es 6 Port Credit eee : Port Mallard, 4 Fort Dover g eee c R! E a= (2 Fondeau b 7 2Merlin rm Z LPoint Pelee 3 Fig. 1. Map of southwestern Ontario, showing localities from which Cladocera were obtained. As . — NS ae = PLATE I. CLADOCERA, ONTARIO Figs. 1-4. Daphnia retrocurva, Rondeau, Lake Erie, June, 1919; variations in cephalic crest. Figs. 5-6. D.longispina var. hyalina form typica, Grenadier Pond, Toronto; peculiar variations. Figs. 7-9. D. longispina var. hyalina form galeata, Georgian Bay; some extreme variations. PLATE IJ. CLADOCERA, ONTARIO Fig. 10. Scapholeberis aurita, end of post-abdomen; Fig. 11, Ceriodaphnia reticulata, end of post-abdomen; Fig. 12, Moina affinis, end of post-abdomen; Fig. 13, Illiocryptus sordidus, post-abdomen; Fig. 14, Ophryoxus gracilis, post- abdomen; Fig. 15, Drepanothrix dentata, antennule female; Fig. 16, Drepanothrix dentata, post-abdomen; Fig. 17, Leydigea quadrangularis, post-abdomen. ~ o PNaod w i 7 t u = < . ¥ « i. _~ ‘be, ass ; ¥ ad r wve ee n> = } : i " "@ ® y f ‘ ) = 7 : Pi a a4 i™ { tt oP t sh ¢ P . . 7 4 yf af A ju j 4 wiry i hs 4 Ue ge al ee Pa . ; at ih aoe A mm, ’ ; ‘ : 4 —s ae mS “ ‘ 4 : ue - t f A . F ; 0 i iB ; : Co f ‘ Re 3 : >’ a a tif , ee + EN ov = ' 5 1 ‘ ’ ’ . ; j ys n , 7 f , ‘ fh a ON = ' { . u 19 5 " ‘ \ V ‘ - r fe 4 ae ball ' ames AA AN AAA A MA AAMAS uu A PLATE III. CLADOCERA, ONTARIO Fig. 18, Pleuroxus trigonellus; Fig. 19, Pleuroxus trigonellus, post-abdomen; Fig. 20, Chydorus globosus; Fig. 21, Chydorus globosus, post-abdomen; Fig. 22, Chydorus faviformis; Fig. 23, Anchistropus minor; Fig. 24, Anchistropus minor, end of post-abdomen; Fig. 25, Monospilus dispar; Fig. 26, Monospilus dispar, end of post-abdomen; Fig. 27, Chydorus barroist? UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 9 BREEDING HABITS OF THE LAND-LOCKED SEA LAMPREY (PETROMYZON MARINUS var. DORSATUS WILDER) BY A. F. CovENTRY OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1922 f i i =} 7‘ i 7) be : ah i] Ni y i j i y i | Th) AVE ( ey { fi ty pov eee ey Wiel } ’ ! : { ay ae , ly j i q ' : f wr sf ve i) bh) y Love nom SUN Pio ens eae ere a he ety eg td i at " J}, ay ae ae eo a bs ayo mn abt ; mm, 1 ela 0 J } b i oh We AT ay i ‘ Cay pe , ies POM Od jie | ea AE Et he Bam : Mes) co i Meri ee yey 0 ‘ Ai i i ve Pee eet ty { ey rr pektlg Cie aS) Wien Ae eB ' i yy Ciara AL) ARTS Ta re fen AS iets belt i . 7. i Ory Ly ; ay, { i" at ae aay | YOt a) ey BREEDING HABITS OF THE LAND-LOCKED Sia LAMPREY (PETROMYZON MARINUS var. DORSA TUS WILDER) Nearly all that has been recorded concerning the life- history of the land-locked sea lamprey (Petromyzon marinus dorsatus Wilder) refers to the earlier description of Jordan and Fordice (1885) and the very inclusive study by Gage (1893) and his associates of the variety as it occurs in Cayuga and similar lakes of northern New York. For many years it has been known locally that a lamprey occurs in Lake Ontario at least some miles east and west of Toronto, where it is sometimes greatly in evidence, attaching itself tempor- arily to boats in motion or more especially to food-fishes and thus brought up by fishermen operating gill nets for whitefish and lake trout in the open waters. This lamprey has been confused to some extent with the silver lamprey (Ichthyomyzon bdellium Jordan). Itsidentity with P. marinus dorsatus s. unicolor was suggested by Bensley (1915), as a result of studies of the animal and of the surface wounds on fishes in relation to reports of fishermen as to occasional lamprey marks on whitefish in Georgian Bay; and confirmed by Huntsman (1917) as a result of a systematic review of the known records of lampreys from Ontario waters. So far as the writer is aware no young have been recorded hitherto from the Toronto area, though there is a single metamorphosed specimen of 13.5 cm. taken in a swamp near Whitby in the university collection. While it is evident from what is known of the glacial history of the Great Lakes Basin that a community of conditions is to be expected as between the New York lakes and Lake Ontario, and also between both and the Gulf of St. Lawrence, the relative isolation of the habitat previously described lends interest to its more open occurrences in 129 130 COVENTRY: THE LAND-LOCKED SEA LAMPREY Lake Ontario and also to the comparison of its breeding habits and development with that of the Cayuga lamprey. At the same time it will be recognized that the term ‘‘land- locked”’ as applied to the animal is largely a matter of descrip- tive convenience. The case is analogous in some respects to that of the land-locked sea salmon in its original occurrence in Lake Ontario. Of these and other species of fishes related as respectively fresh water and marine we know neither the nature of the original medium nor the environmental factors associated with their varietal and specific differences. On Friday, May 27, 1921, Mr. J. R. Dymond of the Department of Biology reported that large numbers of land-locked sea lampreys were in the Humber river, just west of the city of Toronto. The vast majority of those he saw, and they numbered hundreds, were in a small rocky pool immediately below the Lambton weir; many were swimming about, others hung on to the rocks from which they could be picked by hand. They appeared to be blocked in their ascent of the stream by the weir, which is about three feet high. Mr. Dymond did not see any nests on this date. It was then decided to attempt to follow the progress of the breeding season, and with this object in view frequent visits were ‘made to the river during June. The stretch of the river in which nests were found is about three-quarters of a mile long and terminated upstream by the weir already mentioned; no nests or adults were seen above this in spite of careful search. Downstream, nests were not discovered below the rapids at the Old Mill Bridge, below which point the river becomes deep with a muddy bottom for the rest of its course to Lake Ontario, a distance of about two miles. Between the weir and the bridge the river averages about thirty yards in width and two feet in depth at this season of the year, with occasional holes as much as six feet deep. The river bed is composed of clean gravel or shingle interrupted here and there by large flat slabs of solid rock, and the stream is broken up by frequent rapids, none steep enough to make real ‘white water’, although the larger are difficult enough to wade against. The general COVENTRY: THE LAND-LOCKED SEA LAMPREY 131 direction of the stream is from north to south and the valley is so wide that in spite of the high banks (80-100 feet) the sun reached the water by 8 o’clock in the morning (standard time). As was subsequently found during repeated wanderings along this stretch of the river, nests were excavated only in the more rapid regions and indeed only in the shallower parts of these. The majority were in not more than a foot of water and none were seen at a greater depth than two feet. Large stretches of gravel of apparently exactly the same physical qualities as in the rapids, but covered instead by comparatively slowly moving water, were completely neglected, while a few yards up or down stream nests were abundant. The nests are shallow depressions in the bed of the stream usually oval in outline, with the long axis in the direction of flow. They vary considerably in size, from about 12 inches by 18 inches to 24 inches by 30 inches. They are constructed usually by a pair of lampreys, but single animals have been seen working, and in one case at least two pairs occupied one large nest, all four individuals moving stones. The stones moved are in general not larger than an inch in diameter, and most of them are smaller than this; occasion- ally, and with great effort, stones as big as two inches across are dragged from the excavation. In no case were two animals seen combining to move one stone, although in a number of cases this would have enabled them to make a much more convenient nest; if a stone too large to be moved by individual effort lies within the ambit of the nest, it remains. The material is deposited on the downstream side of the nest so as to form a kind of parapet of which the curve conforms to that of the oval hollow. An animal, having attached itself to a stone, allows itself to drift tail first with the current, until its mouth is over the downstream edge of the nest, it then brings itself to a halt with a sharp movement of its tail and at once drops the stone in position. While choosing stones the animal’s movements are rather indeter- minate; it will often lift and drop immediately a number of 132 COVENTRY: THE LAND-LOCKED SEA LAMPREY pebbles before carrying one clear, and often it will remain stationary in the nest for minutes on end, either adhering to a large stone or lying at rest in the still pocket of water caused by the parapet. At the upstream edge of the nest there is nearly always a stone of markedly large size. No factor determining the site of the nest could be discovered other than the general ones of gravel of the right degree of coarseness and a sufficiently rapid current; given these, nests seemed to be impartially distributed in mid-stream or near the banks, at the upper edge of a rapid or down its full extent; in places they were so numerous as to suggest aeroplane photographs of heavily shelled fields. The actual process of laying was watched a number of times. The two animals concerned cease carrying stones and take up a position with their heads at the upper edge of the nest; this is achieved in one of two ways; either both attach themselves to the large stone already mentioned, or the female alone takes this position, the male clinging to the top of her head; at once after this the posterior halves of their bodies twist together for about a complete turn and simultaneously make very rapid flapping movements, so fast, indeed, as to be almost vibrations. During this process, which lasts only a few seconds, eggs may be seen pouring from the female as a number of small white specks, which become mixed with the very small stones and sand stirred up by the agitation of the parents’ bodies. As soon as this movement ceases eggs and sand together settle down at the bottom of the nest. The male and female then separate and resume their stone-hauling, often moving stones from points a foot outside the nests and placing them on the parapet, but after a few minutes the laying process is re- peated; how often this interruption and resumption of laying may occur was not determined, but certainly as many as four. In the large nest already mentioned as being the work of four animals one and the same male was seen to pair with each of two females, eggs from different mothers being mixed in the nest. The eggs when they are first laid stick so firmly to stones COVENTRY: THE LAND-LOCKED SEA LAMPREY 133 that any attempt to detach them usually destroys them; after about fifteen minutes, however, they do not adhere at all so closely and may be washed off with a gentle stream of water from a pipette; in the course of a day or two they lie loose among the pebbles. During the period over which nesting was watched the temperature of the water varied from 18° C. on June 4th, to 23° C. on June 21st and 27th, the temperature being taken between 8 and 9 o'clock a.m., standard time. The nesting season lasted approximately a month. As already noted abundant adults were seen on May 27th, but no nests. On June Ist nests were found in a rapid stretch about six hundred yards below the weir; the animals had apparently relinquished the attempt to go higher up- stream, since on this date none were seen in the pool, nor were any seen subsequently this far up. ‘The nest of June Ist had probably been provided with eggs the previous day, as those from which samples were taken all showed segmenta- tion stages. The last nestings observed were on June 21st and no adults were seen later than June 22nd. As to whether the parents die after spawning I cannot make a definite statement. I have, however, a strong feeling that they do not, and this opinion is based on the very small number of dead lampreys as compared with the abundance of the living seen during the nesting season. During June not more than a dozen or a dozen and a half dead were seen either in the nesting reach itself or in the slower stretch of water between this and Lake Ontario, while in the course of a few hours 150 living individuals were taken from the pool below the weir at the beginning of the season. If, then, the parents die at the end of the breeding season, they appear to return first to the lake, for it is hardly imaginable that so many individuals dying in so short a space of time and in so limited an extent of water should leave so slight a trace of the event. An attempt was made to determine the time that elapses before the young leave the nest, and although the data are not quite conclusive, a probable estimate can be made. 134 COVENTRY: THE LAND-LOCKED SEA LAMPREY There are two sources of possible error, first the necessity of disturbing the pebbles at the bottom of the nest among which the larvae are developing. The young after hatching are found one to two inches below the surface and in the undisturbed nest have presumably reached a position in which they are not affected by the flow of the stream; to get samples the overlying stones have to be turned gently aside with the point of a pipette until the larvae appear, when they are at once sucked up. In this process a number are inevitably dislodged from their crannies among the pebbles and swept away by the current, and it is not improbable that during the readjustment which must follow any such violent invasion of the colony new circulations of water are established, carrying still more of the young prematurely from the nest, which will thus be empty before its normal time. This error involves probably not more than two or three days. The other complication is the difficulty of deciding when a nest is quite empty. The larvae are often distributed in patches over different parts of the nest, and while the greater part of the nest may not reveal a single larva, finally a thickly populated area may be touched by the pipette. The danger lies in overlooking such spots, and this is the greater since thoroughly to search a nest is a tiring procedure, the most practicable way being to kneel by it and, using a water-glass for clearer observation, bend close over it while turning the stones. With practice, however, likely spots are soon recognized, and in this case again the total in- accuracy is probably not large, almost certainly within the limits of variation due to changes in temperature. The two errors, further, tend to balance one another. A nest called No. 3 may be taken as typical. It was first observed on June Ist, and the eggs were within a few hours of laying; on June 22nd no more larvae could be found in it. It should be mentioned here that specimens collected from this nest at various times and allowed to develop in the laboratory had by this time become quite transparent owing to the gradual disappearance of the shining white COVENTRY: THE LAND-LOCKED SEA LAMPREY 135 yolk, that they were taking solid food from the mud provided, that they could swim powerfully and fast, and that they always made vigorous efforts to get under small stones or into the mud in the vessel. They were thus properly equip- ped to start the new phase of their existence. It is certain that these laboratory-bred animals were normal since they were compared time and again with specimens brought freshly from the nest, and no difference could be found; they could be used, therefore, as a sort of indirect check on the likelihood of the nest’s being fully vacated. The young, therefore, left the nest after about three weeks: this result is confirmed by observations on other nests and may be accepted as the average length of time for such conditions as prevailed during the breeding season of 1921. That the young leave the nest voluntarily and by their own exertions can, I think, hardly be doubted, since during the breeding season no rain fell after June 4th, and the stream became steadily lower and consequently the current less likely to disturb the nests. The young hatch out of the egg in seven or eight days. I have not yet been able to discover where the next stage of life is spent despite careful examination of many samples of mud collected below the spawning reach at all depths down to 4 feet. It is worth remark that in all the nests examined a number of eggs failed to develop; the percentage was not accurately determined, but is estimated at 10 to 20 per cent. of the total. These eggs become a light brown colour and may be found in the nest throughout the whole period of ‘incubation.’ On examination they prove to be covered with a delicate felt-work of almost colourless fungal hyphae, and if any of these infected eggs are by accident introduced into a dish in which larvae are developing there is formed on the floor of the vessel a strong growth of fungus, in which the larvae, as they become active and attempt to burrow, entangle themselves and perish. The causes of this non-development were not discovered, but three possibilities seem available; (1) that some eggs are incapable of fertilization, (2) that the 136 COVENTRY: THE LAND-LOCKED SEA LAMPREY rather haphazard method of fertilization fails to ensure that a sperm reaches every egg, (3) that the eggs begin to de- velop, but owing to some weakness are at an early stage invaded by the fungal parasite. Further research is required to clear up this point. Finally it seems worth while to call attention to the combination of unattached eggs with the position of the nest in the fastest accessible water, a condition that would hardly be, a priori, expected. It is true that for the first few minutes of their free existence the eggs are attached, apparently just long enough for the stones to which they adhere to settle into a permanent arrangement under the current conditions obtaining in the nest; subsequently they lie scattered loosely among the pebbles. The parapet on the downstream side of the nest certainly creates an eddy of comparatively still water in the nest, but a very slight disarrangement destroys the effect of this, as many experiences while collecting have shown. The aetiology of so curious a laying habit presents some interesting problems. Iuterature Cited Bensley, B. A., 1915. The Fishes of Georgian Bay. Contrib. Canad. Biology, 1911-1914, Fasc. 2, Fresh Water Fish and Lake Biology. Biological Board of Canada; Suppl. 47th Ann. Rep., Dept. Marine and Fisheries, Fisheries Branch, Ottawa. Gage, S. H., 1893. The Lake and Brook Lampreys of New York, especially those of Cayuga and Seneca Lakes. The Wilder Quarter Century Book, p. 421. Huntsman, A. G., 1917. The Lampreys of Eastern Canada. Ottawa Nat., vol. XX XI, p. 23. Jordan, D. S., and Fordice, M. W., 1885. | Review of the North American Species of Petromyzontidae. Annals N.Y. Acad.Sci., vol..3, p. 279-296. Fig. 1. The weir at Lambton. Fig. 3. A characteristic reach of the R. Humber, showing a succession of rapids and the nature of the banks. UNIVERSITY OF TORONTO Sa Uilres BIOLOGICAL SERIES meets PUBLICATIONS OF He ONTARIO FISHERIES RESEARCH LABORATORY. X (hSLACIAL AND. POST. CEACTAL LAKES IN ONTARIO, sy A. PS IG@r reas: E UNIVERSITY L IBRARY: PUBLISHED BY - LIBRARIAN, 192 University of Toronto Studics COMMITTEE OF MANAGEMENT Chairman: Str RoperT ALEXANDER FALCONER, LL.D., K.C.M.G., President of the University Proressor W. J. ALEXANDER, Pu.D. Proressor J. P. McMurricn, Pu.D. Mme oes eH. AlircnELy, B.A.Sc:, C.B., C.A.G., D.S.O. oe eo al A SS mR fy ee io S30 i} EROPESSOR (Xe. Aran S\ Oi Roa s2vg) L fis Proressor Greorcre M. Wronc, LL.D. General Pdifor> WS. Wartacz, M.A, Pes Associate Librarian of the University - UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 10 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO BY A. P. COLEMAN - TORONTO THE UNIVERSITY LIBRARY 1922 “ELECTRONIC VERSIGR ,. AVAILABLE i a oe ee OR WN FY OO OO SIO oe Whe bISE OF ILLUSTRATIONS Map oF NIPISSING GREAT LAKES - . NIPISSING BEACHES AT BRULE - Map oF LAKE OJIBWAY - - - . Map oF LAKE JROQUOIS = = . TROQUOIS SHORE, SCARBOROUGH) - IROQUOIS GRAVEL BAR, EAst TORONTO . Map oF ADMIRALTY LAKE = = MAP OF MAXIMUM MARINE INVASION . DutcH CHURCH IN 1900 - : - . DutcH CuurcH IN 1915 - = - . MAP OF SHORE AND ISLAND AT TORONTO . SECTION ACROSS THE PALAEOZOIC BOUNDARY . ESCARPMENT AT HAMILTON” - - . MAP OF THE LAURENTIAN RIVER - . Map oF LAKE ALGONQUIN - - See Sa ee aes GEACIAL AND POST-GLACIAL LAKES IN ONTARIO Introduction The following paper has been prepared in collaboration with the Department of Biology of the University of Toronto, members of the staff of which are now engaged on a plan of investigation of the economic fishery problems of Ontario waters. Present conditions relating to the existence and dis- tribution of the fishes and other aquatic organisms obviously depend upon the succession of physical changes which have taken place during the past, but in the case of the Great Lakes and related waters the transition is especially import- ant, not only because of the enormous area affected but also because the most significant changes took place in the period immediately preceding the present one, and, centering in the northern continental region, involved great extremes of both temperature and physical modification of the land surface. It is intended to bring together scattered materials on the subject and to put on record a number of observa- tions thus far unpublished; but in the main to outline the Pleistocene history of the lakes as worked out by previous writers such as Gilbert, Spencer, Fairchild, Taylor, Gold- thwait, and Johnston. The account of Lakes Iroquois and Ojibway will be taken chiefly from the present writer’s previous publications. References will be made to climatic conditions and to the life in the waters and on the shores of these ancient lakes in so far as the evidence permits; and Special attention will be given to the comparatively little known lacustrine features of northern Ontario. The southern shores of most of these extinct lakes are almost as well known as those of the existing Great Lakes, but the north- eastern parts, where their waters met the waning ice sheet,’ are still only imperfectly worked out, largely because the region is sparsely settled and to a great extent covered with forests. 2 ody fad ire, f ’ re i ,% ‘We = ' 4 rye} ' thy f 4 Z 3 a >. ' re P . - ’ | J : ¢, - ' : : - 4? ‘ P ; MMe wl hi y Si ae igs 7% at Rae A The! P| ‘ aur . lownin “Hans tfeagly Iai t074 & OF ee) = 2 oc, Smee lite ait t aon it ae =e) J , wae... lage ty” 3 » ete ate ea rus i wt ea ne iw mpinen ipa i . 7 tT aa era! -oh °o ac) eras ivy wt Ma? le Lew ii? AEH Jal Jose sl i ‘ “y ey i) TI iit = rity tiie » vbte bre ede tk ig 10 ft) eevee “i at 7a : otf rir oe d “ ) = ;>iLw 7 ie ; a vy muro K d=, : et pepe of Dee coh bo 4G 01 ha Of yw oh theless alt loves > ; ~. ty ae na vad a) ardg sae cal 1 es ree or 2s eins Al Ay wiaity4 7 mit acta ily of Qu Oe y yi vr? ol Fe i < J Trutel : 5 riiw Lash < a 9 hick 7% 7 | vi 7: F -_ {nih se Jc bent’ nha an am Tid ere tie bone ribald oN sino oO) 20 isd ee enti ans an Pd "spied ody gs ] me aS iqani i nine i 2s 7. neveie Snead Pere meiboury tena & dy th —_— ioe imilton 2. Escarpment at I EARLY CONDITIONS IN THE GREAT LAKES REGION 7 and running waters. The crystalline Precambrian rocks have in general resisted these attacks best, and the thick beds of shale have suffered most. The result has been that the sedimentary rocks, which once encroached much farther northwards upon the granites and gneisses of the old Pre- cambrian continent, have, except where protected in some cavity of the ancient surface, been stripped off for a long distance south of the latest shore of the Palaeozoic sea. In this process the shales were removed most rapidly and the limestones resisted best, so that ultimately a broad unsym- metrical trough was carved out with a gently sloping surface of Archaean toward the north and an irregular set of cliffs of the strong Niagara limestone toward the south. The wall was by no means straight, and in places there were two or three lower lines of cliffs between the main elevation and the northern slope of granite and gneiss. This row of Sh a» > ONY sy NS AN \ \ ROA ONN \ Ay : SY, VESTER ARRAY ak AY MESS Aes QS RRA MV OWIV|_ MI RNAI SS 1. Section across the Palaeozoic Boundary cliffs facing the old land has played an important part in directing the drainage of the region and is still a marked physiographic feature, called in southern Ontario the Niagara escarpment. As the surface above the escarpment follows the gentle dip of the strata southwestward, the arrangement is sometimes called the Niagara Cuesta. The eating back of the sedimentary rocks must not be thought of as due to direct river action but rather to the slow decay of the shales under atmospheric attack, resulting in the undercutting of the’ cliffs, allowing slices of the overlying limestone to fall, the fragments being removed gradually by solution due to carbonic acid brought down by rain. The escarpment has been caused, then, by differential weathering. Low He BA y ‘ u Tt &; 4 * | 2 oe 4 re ‘ Te af o * ‘i ae ad set irpas rey) Dap; Tet? Ta box ole IQHIAD AGE to nit AL wiped nz cal osmegi ; : ; : a a nMinsawry: of > sealer =} : f. " sevid > D, { 7 v9 au 4 fp : : : # - Eye ber’ ped yo Fai EGF zor’ budiigte et wie Wonks wi y brn keane ee Le ‘of ' re a A va) gales oy is. wr i be Seen eh) oy site tat wht TO 8 nt: date ale Bin Dvrisl ery ‘singe ual , ye Fee we vis S/() i he yale ; 7 ye OL “y aM I wae \ ar a imerty lived adi 50 . = ine 4 pied ot vttnh . Jesse wis: 10) : sl nos agean ellis re its on Li gatwatty Oe att ot if oe ; | ohanamiehe o wn rerrat g i raPyORNa (ie 5 ah | ra § Iowa Penton 6 i tine OR wilt 14) fy sine ot phil) niieton A ‘a JiMNItEeD mpl 3! he ey u j Mit yi) Aaa ee EF se Aine 494 ca adT ites vwinle ; bmndu Whar of Dope ; he gh” feials wole nie “4g orf ati eines OY ni gett iu OF 4) Bretesnit any lyre, nl et sub sobtoloe vd’, ciinuheeh ae wh? Joomeeee tt nil a mr hh iw Ie ; a'% o - s Ps = =a" a of : ‘ =, : ty a “Ny diay fe'3. a ii See, 8 GLACIAL AND POsT-GLACIAL LAKES IN ONTARIO Some time before the beginning of the Glacial period the Great Lakes region stood much higher above the sea than at present, probably at least 1,800 feet. This must have had important effects upon the drainage, steepening the grade of the rivers; and if the uplift was greater in one direction than another, possibly turning the drainage into new directions. There were probably no lakes at this time, the valleys having slope enough to allow all the water to run off; but the directions of flow are not always very cer- tain, and different writers have expressed quite opposite views in regard to the matter. The Laurentian River J. W. Spencer, one of the first to discuss the Preglacial rivers, came to the conclusion that the whole region, except the Superior basin, was drained by a predecessor of the St. Lawrence, which he called the Laurentian river, because it flowed on or near the edge of the ancient Laurentian con- tinental mass, reaching the ocean through what is now Cabot straits between Newfoundland and Nova Scotia. Some American writers, on the other hand, consider that the elevation was more to the northeast and that the rivers crossed the escarpment by one or more gaps and joined the Mississippi, finally reaching the Gulf of Mexico. The latter idea has been worked out in detail by Grabau.? who thinks the main stream flowed southwestward, crossing the escarpment by a channel at Dundas. Since Grabau’s wori was done, evidence in the way of well records has been obtained showing that a channel connecting the Upper Lakes with Ontario extends at least to sea level. This and the great depth of the Ontario basin, reaching nearly 500 feet below sea level near its east end, strongly support Spencer’s theory, which will be outlined here. In Preglacial times eastern Canada extended to the edge of the continental shelf, 140 miles beyond the present ‘Evolution of the Falls of Niagara, Geol. Sur. Can., 1907, pp. 289, etc. *Bull., N.Y. State Museum, No. 45, Vol. 9, 1901, pp. 37, etc. rn mieeet > Ged. eal 7a Te alee. rn KA LAN 4 if 2 x ince , ads Yor prod pe sted amit : ie Sess gic degre MAD sodeed 0 ii eek ap ad A 46 vhingurds,.32eegae ; are Si)? I) yl 2G ‘ie bad t) Te a kis has eter ee pile HR Witrined - T cl eed vaste, | alt rt ty eeteton {doles arout J. ca righty “SAVE apes VERE be ras. ‘ores wait mil tae a’ yee JOM ene pa ty area sub ait, othe The ey eee oF, CIOs righ id B? ritiney 9a a PP. _ VERA Gi ver A}, es = 19 eee Silt one ‘ rh prt , ay “3 of eeu ee ae | YAR arid sn aie sod iioniiw eM: a ore iy ot ylos “seh Ter w) 7: eve. | eer quiload : bagohwavt, Tale | : - hah gey rid? alin’. : ed i} wan Scie mS, - ‘ ; hes sever ita é inhkotet eee an Me ‘ ny. 4 per sari a ‘Se cievepty hiner ean liek bol a on eel, oni Af i, ae , si | abd , ‘Ay ‘Yea tie id at = ws iat Lhe Veoa' oe ae 4 . Jie iS¥V F iJ ret ele 603 tee! rape. tHeKD vinci al mot is mH} mo 3B es mo Pe | boniisve ot Tee ‘aid tm mild ot eet ir S Ob Poni tained 2rd Inessny od Rye) Sollen, Olek /Veaae tame . bs 7 Le ‘y ss Det an OPE .ewl Au Jn? ote SE iy PORE (o To? } = ; ie — - . =~ = = i ~ i a i i Ms ; te 7 of i i ” 4 _ ra va f “ wre > med i »< yf - t ‘ : ot" i’ \eo)) em: ioe at ———EeoeEeEeEeeEeEe—— ES I EE SE ee Se a Se eons Sa ars Ot a Se bat THE LAURENTIAN RIVER 9 southeastern coast of Nova Scotia, and Newfoundland was: a part of the mainland. The old river channel then exca- vated can be followed by soundings to the edge of the en- larged continent, where the shallow water ends, and the bottom descends toward the depths of the sea. By the kindness of Dr. Huntsman the soundings on the chart have been examined, and it is found that the deepest point reaches 335 fathoms and the next deepest, farther up near Bird rock (Magdalen islands), 313 fathoms. The old river valley extended 840 miles beyond Quebec and at the edge of the continent is now from 1,878 feet to 2,010 feet below the sea level. Dr. Spencer made the depth 3,660 feet (611 fathoms), but this sounding is on the slope toward deep water and there are no shallower soundings on each side, so that his interpretation is probably a mistake. The rest of the great river channel, as given by Dr. Spencer, is mostly above sea level. It begins on his map in the deeper parts of the basins of Lake Michigan and Lake Huron, turns north from about the middle of Lake Huron to Georgian Bay and then bends southeastward along the foot of the escarpment to Barrie. A buried channel extends from Barrie to Toronto, after which the river kept to the deeper south side of the Ontario basin, and finally turned northeast along the present St. Lawrence valley. The buried channel between Georgian Bay and Lake Ontario was inferred by Spencer from wells sunk in drift for a town water supply at Barrie (280 feet) and at Richmond Hill (400 feet). Since his results were published the inference he made has been confirmed and strengthened by several other wells, one at Newmarket reaching 265 feet, another at Bradford reaching 330 feet before striking rock and one on Mr. Page's farm two miles west of Thornhill, where 650 feet of glacial and interglacial materials were passed through before rock was encountered. The well was finally sunk to 1,203 feet in solid rock, ending in Laurentian granite. As the Page well was drilled at a point about 650 feet above the sea, the old channel extends 246 feet below Lake Ontario to present sea level. 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Map of the Laurentian River Tributaries of the Laurentian River - If weassume that the Laurentian river existed, and that it threaded several now distinct basins, it is evident. that there must have been numerous tributaries joining it. Spencer has indicated several of them on his map, the first being the Huronian river, formed of two branches, one -coming from the depression of Saginaw bay, and the other running north from the southern portion of Lake Huron. In the Ontario basin he maps two tributaries, the small lea ok Bh si gens mAs 4) Mat vie ai tat! J eae ; ’ — , ' - = to vv ; i a ag ; gaviji’ ¥A Pre nava 1 rt ge } Poa : - of bk’ 14 ‘ breed ees 2 ial : rms 30.4.0 : be b& Avie ; . i ee vee) (SEH sir 2 “Sh mis “Weta oa ' Ts enerierey =f! x etait) jo Gena "i ‘Ti ia Ce Beer 1A) st ie wire var a ae ~e * dee write rar au eee at Tal 4 A y bi ‘an ai ‘i p ~ oguq i; , vet? i FTO . 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Sg ud aeiry. ‘ia OF iG + ty emewam tie ae oe a CS, ! meee ee PP Jol naae Lrowaoils! 9.40 AAli 94' urls? OFLE ir syns nak a adi oY wd am wit an Le Byes Am ys + foe ia. = Ay 27 : ‘> : u 14 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO The Lake Michigan valley probably drained partly into the Mississippi under Preglacial conditions; but the south-. ward outlet is now blocked by drift of much less thickness than that between Georgian bay and Lake Ontario. Lake Erie owes its shallow basin partly to the filling with drift of an old outlet toward Lake Ontario, but largely to differential uplift of its northeastern end. Lake Ontario, also, probably has a drift-filled outlet toward the east, but has been greatly deepened by differential raising of the outlet at the Thousand Islands. The two lower lakes illustrate more distinctly than the others the formation of basins from river valleys by the elevation of the outlets. They have been formed mainly by the action of epeirogenic forces, and damming has been of minor importance, while Lakes Huron and Michigan are held up by dams of drift materials. Precursors of the Great Glacial Lakes The glacial lakes of Ontario and the adjoining states, as¥ generally defined, include only those bodies of water formed during the retreat of the last, or Wisconsin, ice sheet, but it is evident that each of the earlier sheets which has covered the region must have dammed back the northward or north- eastward flowing waters in a similar way, and, in fact, that each ice advance must have formed lakes which gradually diminished in area as the ice front moved southwards until the whole of their basins was occupied; and afterwards a series of lakes expanding as the ice sheet retreated when the climate became temperate again’ How often the Great Lakes region was invaded by ice is not entirely certain, though the geologists who have studied the drift of Iowa distinguish five successive ice sheets as having covered parts of that state; while each of four interglacial intervals indicates a great retreat of the ice, if not its complete removal, betore the following sheet advanced. In Ontario there is positive evidence of one interglacial period of milder climate than the present, when es 2 ae | a ee “1 eae Ce ~ ¢ a / ak Fl ‘ Ad me rAL ‘sh; “= Se ure ’ — ; VA 7 bin ta, por hin’s peal nfl Mus: ‘cacti Dynegy it} slciepacert. Vee TIGA be Mpeg wiht ney twinit ) is ! ih sd REL yin? Apov aa Wyre “srs onic id st) tie W { a 23 “ro 9 ? . rel i ent FHA RP, ; ni al ae tidy a8 aaa tes un swoe Ati o ‘¥ ve t i) oe La » tte, Plev iG Leake Ale Orn " CNS, LEER IAMAL. yee | : a ie Hatlst ’ 4 dy yi’ 7s sold a i sult mation of . yond) aor f hfe { , ; ep pi pie ro by ily yd Qala Cpr tage Per sich Bet | pitorrcy. “Me nd | fi ytte H | Dah Wal: Came Salil oath — = S50. ope “Av i te ; ; Spi antete yninetie 0 han vices ‘ Fh sata ote oe ry fino ool wetneth ys | athlon). Saal oh ai ein DD ve 7 et le (ane : if 3a heer? rah. J t aan ’ 4 ‘ } wr “| } : 1 $s +x? is ' J \7 1 Ty mii rieey Ti 3 | < pric l ' . he ri ul a. pie Ah. Ganitgn ; vy, ShG 2 P iT; Dest , he ey THRO TROT I iaitoats na ” in i ® apr] 4 writ "Ohne le vy (ti cate ie ¥ eet peeks ft) myiK ial want ORM 7 ain Tose siti ned? Pate (ya 173 a “ = Be a neat Tr 1) " — ; oe, j yi é wie! 7 } Pr se } 4 eo ; * — aT) or ai A, o ant Oo : ; ae hoa 2 é rhe i Py iy sm i T g v7 7% if Pa i Lh ry & Pen 7’. Ay rh . ; eee nrc ae ee a ae en oe — cia Utila at oe a rm PRECURSORS OF THE GREAT GLACIAL LAKES Ly no ice sheet could have survived, and of another great re- treat of the ice when the Ontario basin at least was set free. These earlier glacial lakes, whether formed by an advancing or a retreating ice sheet, have left only very fragmentary evidence of their existence, and any attempt to map their boundaries is out of the question, though there can be no doubt of their existence. Later ice advances have in most cases completely buried or removed the proofs of such bodies of water. During the Toronto interglacial period there was a lake in the Ontario basin which stood at first 60 feet above the present lake, and later ros: to about 150 feet. An inter- glacial successor of the Laurentian river drained the Huronian valley into the lake just mentioned, forming a great delta at Toronto covering more than 100 square miles with sedi- ments having a thickness at Scarborough Heights of 190 feet. As the climate for part of this interglacial period was distinctly warmer than that of Toronto at present, as is shown by the trees of the time, ice cannot have served as a dam, and one must suppose that the waters were held up by a differential elevation of the outlet near the present Thousand Islands. Later the barrier was removed and the water fell to 40 feet below the present lake, as is shown by old valleys carved to that depth and then buried under the next boulder clay. This level may correspond to the supposed drift- filled outlet southeast of the Thousand Islands. The lake of the Toronto Formation had a rich fauna, including 41 species of shellfish, of which ten or eleven are unios, mostly forms now living in the Mississippi! Fish remains occur also, the only certain species being a large catfish, as de- termined by Professor Bensley from a spine. The evidence as to climate furnished by the aquatic life corroborates the conclusions drawn from the trees, suggesting temperatures corresponding to points 4 or 5 degrees farther south at the present time. No interlobate moraine had been deposited across the Preglacial channel before the Toronto Formation, ‘Guide book, No. 6, Geol. Congr. Tor., pp. 17-18. ~~ ees ir a me Ven ee eee ee ee oy a el 7 4 | m ~™ i aye ty sa 4 ) PRAMAS eed Ts TARAS iT bad santos An Fhety tone Ne Pan yb. rye ja ove bt siden M4 ooh a bani) * OAL. ea i ask ED: rrarbedei ee Dirkeeanlee Mig A ged oartsbclve. seme iio se Wehbe a eae ow bits ‘il cved .jotila Bora Sie) Ube: ee Hee TT. Winn ies wHito tales, Te aod alot eyooels , comp, ae, 19"ahtee gocher enh: moamrl ee “| 1 Yie tanh Stel <- biti eh anid Oar ie’ bee Rass , ees ¥ iy! oa pi iy ¥ al Ch at ' veal f ) . vi WAT? Ly | ais Nena ae” ae oanrer? fugen esi “He coal leer Tate Se eorceanel . ¢ w LA SGI BEND, 2 4 be wot wacky oe wy Hho nated age eam len ? tH in bd pr, ET TO sane ot pag qu biel ne soli pot aaa | Ls where tet out ee Ay bie hecoungn- ante abi ie &] as ot heen, ; ott uabow boiktaaie haan Sita eB SAT wt, lh onenae pesiet noes ‘dae brttel bosewett ons to 38h an do) dais ha ; i hfe wn Mowe TP oie 4 ; ieApert abe eee es tae be ont Lyn vite ean e Ruiud ieee eytiethys: of Fema. tice eotiepeL 4 ih? cosmic oll di bala jute wit it & mpet 07% cyitt Wy widsg nae, err ana iat alt as ote hedtae) eor'yoh dee “He ras iby poueeh al nai i oF la eb rin" ws nyio T ey eet chile is 2 Shan s eg ey) WO Roly ee, aes a, FO Ee ee 16 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO so that the St. Clair, Detroit, Niagara outlet had not yet come into existence. During the advance of the ice sheet which followed the Toronto interglacial time, a lake must have been formed in the Ontario basin, but no certain deposits belonging to it have been found. During the retreat of this ice sheet, which extended beyond Newcastle, fifty miles to the east, the glacial lake reached a level of 204 feet; and later the process was repeated twice more, as shown by beds of strati- fied sand and clay 25 and 36 feet thick respectively, between sheets of boulder clay, implying water levels 238 and 306 feet above Lake Ontario. A few small shells have been obtained from the second interglacial beds, belonging to species still living in the region; but the two upper sands seem to have been deposited in lifeless waters, probably with the ice front not many miles to the northeast. What took place in the basins of the other Great Lakes can in most cases only be inferred from our general knowledge of the advance and retreat of the great ice sheets which passed over the region; since lake deposits belonging to the successive interglacial intervals have not been certainly identified. Lake Agassiz The earliest of the glacial lakes, as usually defined, lay to the west of the region here specially considered and will be mentioned only briefly. Lake Agassiz, as it has been called, occupied parts of Saskatchewan, Manitoba, and Ontario in Canada, and parts of Minnesota and North Dakota in the United States. When at its greatest dimen- sions it covered 110,000 square miles between the watershed to the south and the retreating Keewatin and Labrador ice sheets, its northern shore being the margin of the Kee- watin sheet and its eastern that of the Labrador sheet. It was a comparatively shallow lake which drained south- eastwards by what has been called the Warren river into Minnesota river, a tributary of the Mississippi. When the two confluent ice sheets melted so far as to separate, a 26h eee - > 1 ive T erage ne? peal Tha ON a mo Barres bortentiy aie, Aeteyh® aha: ti s eet war den eetita) » y9ttte RT Pe Lae depretaiet etl D4 eticay f/f oii) hem = Di - ae ti teav tina Pid pel 63 td niente ; Trike Me ag tral n Carty aly nay Pete ee — 4 i 4 too > wth) v? 7 oh TE AR ‘treet te Bele: dati enya es ’ . in o 1% ld with Lh Levnin droid ar ee I y s ! §4 24) ; . if ISR. ivetlent any tian re ee wt P theg we te ors “al Hing lah Ay ae anes RAT a 2 na ak nia ae lpn sae Sa HOE 1. vestilteaah PR LAL is be rik csieaqaby el Cae! "os ‘rarpared suits then! alte lad (ont oe ior yids y iiieeths ‘MYO naa wt by a Sal aS Sa AS, Lag te ake Sik rad i aac fe ar Se 2 shel Aalst SI Nit aa ta nh 18 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO readily accounted for. The climate of Manitoba at the time was probably not much more rigorous than now, since the buffalo lived on the shores of the lake; and the fish and molluscs could not have multiplied as they did unless its waters were unfrozen for several months in the year. _ Early Glacial Lakes in the Basins of the St. Lawrence System Probably before Lake Agassiz was drained, the basins of the upper lakes of the St. Lawrence system began to be set free by the melting of the southern end of the ice lobes that occupied them. ‘The evidences of this are found mostly in the states to the south of the province of Ontario, so that it will not be necessary to describe in detail these early glacial lakes. They have been mapped by Leverett, Taylor, Goldthwait, and others in the United States, and to a minor extent by Spencer in southern Ontario. A full and excellent account of them is given by Taylor in Monograph LIII of the United States Geological Survey, to which those specially interested in the matter are referred. Numerous early stages of water have been recognized and more or less completely mapped in each of the three upper lake basins, but only the most long lived of them need be mentioned here. In the Superior basin Lake Duluth followed up the shrinking ice lobe and ultimately occupied about half of the present basin as well as flooding a strip of the present shores. It drained southwestward through the St. Croix river valley into the Mlississippi.t This lake, when at its greatest extent, covered part of the Thunder Bay region in Ontario. The southern half of Lake Michigan was occupied at first by a crescent-shaped lake which expanded northwards, and has been called Lake Chicago. It emptied toward the southwest by the valley of Desplaines river, a tributary of the Mississippi. The southern end of Lake Huron and the basin of Lake Erie had a very complicated history and numerous names 1U.S. Mon. LIII, p. 328, map op. p. 400. a, are , ar @.t& i) an hb i aa 2 ae pcitaiienihaslieeeasitialdapeh iia (MEIGAt leedininiio s6- ap voianeeaaiil 9 linumJon : Lio Fan ihe el to ecods often have are Mo. veg: war. Di! 1g) aie ov étr 68 bhions . oe a estes ouvey 1 aeons iat | | wold Yb 1.12 8) a 7 re tlodh oul ¥ eh) 2 eae r “ul ees aug : = fit ‘ iron. aed Oe , ae, ee 8 to! uM Vie ee) a oe at ia bricyegs ro nol ove t= aM “antzir. ret As ciel | ee it A eRe ae ii ee nye a aires? PAA hl. im 720'! GAA LOA “2 0 a ss rt fi LR Hie 4 si atgily Yes Nis OT ph i“ eel ike +, > ‘yh Ue ; ETT S.i/ “ahi «4 covt. sunlalgeo(t 3a ies > - a eT. © : a et : a . : Pe. PO aT fae a ne et te i PS Nh TE RRS Aa SESS CN a lakh Sl eT A al Mit ea SES. at le a LAKE ALGONQUIN 19 have been given to bodies of water following up the ice. Only one of these, Lake Warren, will be mentioned. Its— beaches extend into the province of Ontario and were studied first by Spencer.! He supposed that the region was covered by an arm of the sea and named this extension of the Gulf of St: Lawrence the ‘‘Warren Water.” Later it was recognized that it was a glacial lake having an outlet through what is now Saginaw bay into Lake Chicago. Spencer believed that this body of water extended far to the north and in- cluded the upper beaches around Lake Superior. He was followed in this by Lawson, who measured many beaches north of Lake Superior; and by the present writer in the same region and farther east. The more recent and more detailed work of Taylor and others in the United States has cut down greatly the dimensions of this lake, and the high beaches to the north are now known to belong to Lake Algonquin.? At a later stage the waters of the Huron-Erie basin sank below the level of the outlet to Lake Chicago, and Lake Lundy took the place of Lake Warren. Some of its beaches are found in southern Ontario. It is supposed tiat Lake Lundy emptied toward the east through the state of New York and the Hudson valley.’ Lake Algonquin—lIts Outlets The various lobes of the Labrador ice sheet were gradually shrinking, though with some halts and even re-advances, marked by the building of moraines, and at length the basins of the Upper Lakes were almost completely free, though to the north and northeast, beyond the present boundaries of Lakes Superior and Huron, ice still formed a part of the shore. At first all outlets toward the northeast were blocked, but the basins of the three great Upper Lakes were in com- munication, forming what was called by Spencer Lake 1Falls of Niagara, G.S.C., p. 287. 3U.S. Mon. LIII, pp. 392, 398. 3Tbid., pp. 399-406. Pe QS ire © y, ‘ 4 ~ % i ‘ ‘ asl bores Mr { ‘ S ) f * ' , ‘ + ret ry hia | ;7 } Hf ‘ j 4 ‘ ‘ 2 % z - f ia La ry @) co a ' 7 7 eniwidud atinw lo esibx oF ae ‘ ; : iors ; a ie peel “~P sit) ~~ Eig a? Trt ! reins i tig OO» ,F be PL air yonivatd Ad am by on ogihestiods 10't brenqe uit set Aaa } in att ' ak nottined Tipe ee olf t renal “ae nor") oe eer io bie lal fn by i RAVE ie te ee eee On con es fy alee) 21. lgbnatyes sa haeeaal scar tide ik p 4 } bt?i Vig nyt 4a wiih 1 iv TA} f be Jy Tr r ' yD Th } 7 eitads (hh oat ag / “i if lt to zee vit, ' ote! ae 9a eek i ows VE le Tee sou ae de OMAR: m7 my Jane wie TE ‘ one ¥ a si Bbaa beet f j ‘ Try ithe i ot tou dhe’ ‘ a/ 43 Lepr pay tins) pare me | Jb hon tirana i otolymep Jaonhe sal le ° _ > z » oe, A an , -s r > i," * se Ao “GEACTAL AND PosT-GEACKIAL LAKES. IN| ONTARIO Algonquin, and probably drained for a short time past Chicago into the Mississippi. Lake Erie was no longer dammed by the ice front and became partly a river valley, and partly a separate lake, through which for a time Lake Algonquin emptied into the Ontario basin. It may be that both of these channels functioned for a short time until the ice so far withdrew from the Georgian bay region as to allow a lower outlet past Kirkfield into the Trent valley. Affairs were in a very unstable condition with regard to outlets, and also as to the area and shape of Lake Algonquin. When the Kirkfield outlet was opened, the land in that region stood lower than either the Chicago or St. Clair channels, and the great lake must have been lowered to correspond, leaving the earlier channels dry. As the load of ice was removed by thawing toward the northeast, the land beneath the thinning ice sheet rose to correspond to the relief, and at length the Nirkfield or Trent outlet reached a level when the St. Clair channel and perhaps also the Chicago channel were occupied once more. There was a long continued two-outlet or three-outlet stage during which a substantial beach was built round the whole shore. Ulti- mately, the northeastward elevation closed the Wirkfield outlet, and the reinforced St. Clair river lowered its bed in drift deposits so far that the Chicago outlet, limited by a sill of rock, ran dry, and the whole drainage of Lake Algon- quin passed through the Erie valley and over Niagara Falls.} During the two- or three-outlet stage of Lake Algonquin, there seems to have been a long halt in the retreat of the ice and also in the elevation of the region toward the north. tn the southern parts of the basins, although the shore stoad higher than at present, the area occupied followed pretty closely the boundaries of the present lakes. Toward the north, northeast and east, however, Lake Algonquin ex- tended much beyond the present limits, a northern bay even including the basin of Lake Nipigon. Much of the land north of the Sault Ste. Marie and northeast of Georgian 1Ibid., pp. 407, etc. Caw , ( \ os°r ) mir iy 4 f Occ M J 4 ; } Te ‘ ’ ‘8 { iy ly ' { | 1 , >; ri } = - ie A} “1 Ad i} ti - a ‘ f ‘ i] } t ' \ ii 7 - >! | i 9, 4 j = h Wipe as {7 “Ad Ki ] $3" a} a ie if] i lo top lt “foie ‘. Mabel ‘og Pui nee i le tenon ris sl avin Iaieqanitte SS ee te (Me = 7 oo = i _ ‘th beat fl * . * . q , 5 ey 4 Te (Th fe alt Mba Va } ai) vleleeleriy hint ee cites ee = Wd ‘iairs ae Jail cat sei ; it soaks » Fe, nit alent el *y nbtew i panty Sole ewe THRACE Ae ee } 3 a8 vi oaln Tayo at net “wee cof See bs at gnivEsd » “getty Wal ime al unite abe ee tay 4 oft ors) Te oe cate Wed 1 att ‘! \ ries | ew, 8 ore Lk nétae role ome a ] ‘tieetl ats ri ofa Teete share ) > Voy ope tk an} oe a) fy of] Gam > te pol off aaigs vo aoay 4a 0 rie nage: or ig nal arma tl j sc ont to a yi rou St) Jee “wc od! To Sore a"? i (60 Us i ine oan es ado uinbuosjy oye] jo dey “fF al ‘ ~ CI irs 4 ; \ 6 Cia, \ Teg iii NIIONOOIV BWI TVIOWI9 40 =F 2 — | \ { uy , 4 if * By ' ¢ wees —) — = bs = a ate “ | me! { | f ¥,° ‘ 2 i j id ~ « 4 ri as A - i* foe! | a! ‘ | cu \ ; = yf) iH , mi i é % = til , ’ \ y is a - f s hi i “ i Ay 1 Ne ae “i S Ay * a P a " - ow >), ow 0 eeey = i ip ‘ - = o - } Le : 4 Dae * 4, yy ue 7 rs, i ay Ae cA A} i Pa ee ee y r a0 » ~_ 7 : 22 GLACIAL AND POST-GLACIAL LAKES IN "ONTARIO bay was flooded; so that the lake covered considerably more space than the three Upper Lakes. It had an area of perhaps 100,000 square miles, coming next to Lake Agassiz in dimen- sions and far surpassing it in volume of water, since parts of its basin had a depth of from 1,200 to 1,500 feet. The southwestern shores of Lake Algonquin have been carefully mapped, but the northern and northeastern are still only imperiectly known, since the region is largely forest-covered and _ roadless. Present Altitude of the Shore of Lake Algonquin A detailed account of the southern shores of Lake Algon- quin, accompanied by an excellent map, is to be found in Taylor's description of the Post-glacial lakes.!. It has been determined that along the southern half of Lake Michigan and the lower ends of Saginaw bay and Lake Huron the old shore is horizontal and rises 607 feet above the sea or 26 feet above the present lakes. The horizontal portion ends at what is called a ‘‘hinge line”’ crossing the lakes in a direction about 20° south of east and entering the province of Ontario at the village of Grand Bend, northwest of London. Beyond the hinge the shore rises as one goes north. At Kirkfield, for a long time on the main outlet of the lake, the shore is 883 feet above the sea, showing a deformation in the distance of 276 feet, about two feet per mile in the direction of tilt. The highest beach rises more rapidly farther north, reaching 1,007 feet at Huntsville, as deter- mined by Goldthwait, and 1,015 feet at Root river, six miles north of the Sault Ste. Marie, according to Leverett.2 Up to these points the position and elevation of the highest shore is considered to be somewhat definitely fixed. A number of shore deposits probably belonging to it are known still farther north, however. At Heyden station on the Algoma Central Raiiway, 13.3 miles north of the Sault, there is a gravel plain 1,082 feet above the sea, which almost 'Tbid., pp. 409-438. 2Ibid., p. 435. eh . 44 +h 1 oded’ dR § ts Fi a3 ae iat 2 a h \ i ; aes ee , 3 "i - k } , ~~ rT he | eon Le Ieee hh, ai as ye Le 4 1h i. LL tered ries fom), pity On eit fi ‘ial } «ff af 4, figelt a a Ls rt Mu at by To ony «faye t {Nast fy Mile bed Tet BATTS Ol “ast yt : nis cial ee aetid bey LP. ee sive A ot ail 1, Tool OY. oi) ingot a ai at heh ioe a) to hoa ae soit Pra tre aod oad 1916) Rr, : vol ke cmt ss AP this ; LGA ei ae me . | bie ‘ pd ahhnat” bath o ae y f tha! \ int Ga Ay “o swig ar » Prertior ane ig cine isin) abet Bh 0 loolles’ Ure yeh eae sign eA, es. st teat Tarmktetney only ae nolity a Heil S hel te tet otsdtios oats aad Pee f i eho K, ne 4 : : ig ve “" cel’ a a he ont gniionny hiked ha ran : veer 5 evel “es ned AT soll oda. 3.31 ane on to ay se 2 44 ’ 1 wt ’ rayne ay nest Td mh iste sit yal Hea, Veni” HN Cea Be de is qt coment £ UF ee pt (en mI ; m ' ae os ot ext tale» gobworde cia ny oioda tee ~ +5 ; 1. oyl ah ft a ~T HW oaoel. Denali , way SOA, weeks pierre a6. F971 vse e wi tae 2a soy CS buat |: *2a3snaiel @ gnibiouon 4 ovin tA DS taatiatel oils > Teen bere souk BD ta Shins chattels Spit ‘yh nde om 19 sani ior ats Woh’ rr ae m1 A, Sc long 27 § gai! ine tet: he 2 roa tf, VON st th fm, wie! jake val Aa i twse nr ie é deere) LAKE ALGONQUIN 23 certainly belongs to Lake Algonquin; and Goldthwait has_ found a probable Algonquin beach at Trout Creck, 5-4 miles north of Huntsville and 1,221 feet above the sea, repre- senting a rise of about four feet per mile between the two places. Northward of the hinge line the beach is split up into groups of successive beaches, showing that elevation Was progressing during the lifetime of the lake. The northeast shore of Lake Algonquin has never been traced continuously, though at many localities lake deposits or old beaches have been found which probably represent Algonquin water levels, a few of them belonging to the highest beach, but most of them corresponding to lower stages of the lake, after the Kirkfield or Trent outlet had been abandoned and the drainage followed the St. Clair river toward Niagara Falls. A number of these points are along railways, since the location engineer is much attracted by flat plains or level beaches; and many of the divisional stations, where yard facilities are required, have been located om lake deposits. In these cases the levels are definitely known. In other cases the old lake deposits have been measured by ancroid during canoe trips or walks across country, and the elevations are much less certain. _ As it will probably be many years before the wild region north of-the lakes is carefully surveyed, it is thought advisable to give these aneroid determinations in spite of their possible want of accuracy. It is not thought necessary, however, to give aneroid levels of old beaches near points where Wye level surveys have been made by Lawson and others, though - many such determinations were made. Water Levels in the Sudbury and Sault Districts Lake deposits are widespread in the Sudbury District, about 90 miles a little north of west from North Bay and 150 miles northwest of Kirkfield. The deposits include plains of stratified clay, mostly in the interior of the nickel basin, running from 820 feet at Coniston to 889 at Chelmsford (both railway levels), indicating moderately deep water : i's ae bigs 4 Peet May “eat 4 } 7 I wip > iis ' P a aye , ” ’ v 7. 5 <) t > Zz z @ = « ie j vid | pf iiiat :* it. iia Fh) To e “9 + 2 | 4 | ‘. a i % vs, Wee EP Ld inlet 2onclady ) The Ly anal rnd i ace fy, Chica rT hort a for “TS bee “ahi en wba nS + eee ied) eb Bi ae te, te nie Pa) irl yerrreh. wild Te leetpe Niwe tee wiolth OVE Laguna to delat? Thiol tien v2 se Seen ind: de otetia Jan Malssiile’ eA 4 naeitu Lac thai VG Bs: AS nla ldw ty nated sot 't Cireeolet Hib). lo we a love im a a(t Tee ka ee \ whee leon ud of lof: >). to TAB of noiz “im M ‘ lj ad “3 ii eihJ yt ite i rq Swe 4 wa tne HM) HOA ot aie) raite i is ‘ apart s tybtas “i peed: “jini al 7 . 0 , ; — ny & ; rry i “4 ‘ 16-' iti (oat (509) vel ottd aeaeee e whe a ; y.te- St) al * tre ha" rpinn” ot > (199 Aoi RAO pare ie a4 sh MM shige i iho “Say beets Sh spy aides wh ded sve es , > beer tre ey ay satecianalll ~cecwen idyaod ) 1g gaie nme 20h reat fet hoy hy als b snorted veh aban HAN wee Sy anon . vendbre oni.ni.§ secon hit a re (dio “non jeve Ts) alent, iit a eta agigebionT - bobo ea rae hey ¢ ms j civ ate anne iii afl soi jay elation PY aL. #1} i tat C 24 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO conditions in a bay of Lake Algonquin. To the north sand plains occur at Hanmer (968) and Capreol (1,003) stations, and there is a gravel terrace five feet above the track at Selwood Junction (1,089). Along the main line of the Cana- dian Pacitic Railway northwest from the Sudbury basin there are sand plains, succeeded by gravel plains at Windy Lake (1,221) and at Cartier (1,378). The last is a divisional station with many switches and tracks laid out on a gravel flat, which probably belongs to the highest Algonquin beach. All these figures are from railway levels, and are accurate. The highest sand and gravel plain observed in the district is at Meteor lake about 59 miles north of Chelmsford and 45 miles in a direction 2’-out 20° cast of north from Cartier. Aneroid readings make the level 1,420 feet, which would give a rise of one foot per mile from Cartier, probably under the true rate of change. The Meteor lake region is a pitted plain enclosing steep-walled kettles which were formed by the slow thawing of ice masses buried under lake shore deposits, at the margin of the retreating ice sheet. Aithough no continuous shore has been traced in the Sudbury District the widespread character of the lake de- posits and their position rising gradually to the north of Lake Huron, make it certain that Lake Algonquin when at or near its highest level extended at least 50 miles and probably 90 miles from the present coast of Lake Huron. On a former page reference was made to high level sand and gravel terraces aiong the Algoma Central Railway, extending for thirteen miles north of the Sault Ste. Marie. Many old lake beds have been found farther to the north, partly along the shore of Lake Superior on the route to Michipicoten, and partly inland beyond Michipicoten bay. The first point measured (by aneroid) is on the northeast side of Batchewana bay, thirty miles north of the Sault, where three terraces stand at 967, 1,047, and 1,152 feet respectively. The highest terrace is 70 feet above the highest one near the Sault, giving a rise of two and a half feet per mile. The next measurements were made near Brulé harbour, I ehh Mea dibate i anon gbwte row a fr i pe slin mad ) , if 7 tO hia of MU Oe r © : vt eine Boe me" Hdl ue > ahi riz 4] rid Ey Wat wiindl inet q la nel te l‘royrona nik im hu) jah See jive tapi ; | Le AL cag an a lon) ar eo 7 ) Seb LL ae j algk a a » nol ee a ont ants eat ik ‘ 194 jog pne Th oo 1 eqns: lo! aoe { teal lore ty wide ¥ R la 1 4 ety wi aedeg om} vi ole nits 1of i] a ae mnie ae inl’ wire’ Se +S ‘ toh 1 lvyyreb.3 it yar Sg sreh eh «act a" Ay A aaaR \. hee ( ee in ee oF to dinan eit nearie ‘enoigt pore: Woah apt ‘areata “0 Js bate hh iY Ned NS LAKE ALGONQUIN pay 92 miles in a direction a little west of north from the Sault. Here, on a path from Brulé harbour to the mouth of Old Woman bay a wonderful succession of bars is found having the following elevations: 857 feet} Shane or ‘ 23 4é 814 ‘* + Algonquin beaches rc eee ‘i ae 724 6 0 il 658 ‘is 636. “ 623 « {Nipissing beaches. G22 -:j* | The elevations up to 734 feet were determined by hand- level, the higher ones by aneroid. The beaches are practically continuous from $14 to 823, and from 842 to 867 feet. In fact there are only two wide gaps in the succession, one above what are considered Ninissing levels and one between 759 feet and 814. The levels given were measured from Brulé harbour up to a col between granite hills at 867 feet, but a similar succession goes down to Old Woman bay on the other side. They are all gravel or boulder beaches, and are very recent-looking up to 56 feet above Lake Superior, being free from trees or bushes, though covered with lichen. Twelve miles northeast of Brulé a branch of the Algoma Central railway runs from Michipicoten harbour to the Helen mine, its route being mainly over great delta plains of sand and gravel deposited by Michipicoten river in Lake Algonquin and later bodies of water. Somewhat east of the harbour, on a road leading from the Mission to Wawa lake, ten terraces or beach ridges were hand-levelled, run- sr \ é . , Mia y S ' } Te etoe ofi4 ttre: He torn igh ig Ae; noe se j EP da oY ei besoud. SU ol ae rirvien lai He mite be ts ye ty thai -sholtnyaledy " an ; be; , 4 ) : is Fr a ay | be Tat wre ‘ 47 Lf th — x = =? a9 * i) v i Balt , ‘ eo iy ” ony ' maya yen i591 Say “s Coe ' wel watt igo uel ety ‘Soot SOE ot SER mogk bak seman ‘He ea: Oe 7 at rh? nt ori | te 4 els ane eee ie Fett 1 VA Stivns hoe da teas) TH He ull, iui cawisd i mapa? fh ry ol cero 20n2 # d shied th nvnyg i KOKP MES? mt owed, Yao) fee ad 1 ub i sa ‘eB pie : v dent ao + yiikhh ye Goch) te pene uted tokonchlt lA sent ahi J ah ahr Sutras iv! Titra it anil. oe) nt progetto sf sel bya tae a ed lo Jeno Jadvretad.. sraticn 16. ail fra tot sae gat ino anita ae ysis: bv) ae maT + “ay! iY Ae 1» Oe 26 GLACIAL AND’ POST-GLACIAL LAKES IN ONTARIO ning from 623 to 810 feet. One at 681 feet is a broad sand terrace, perhaps the highest Nipissing beach, though the greatest break, of thirty feet, is just above 702 feet. Brient Station on the railway is on a terrace of stratified sand at 759 feet and Wawa Station, on a gravel terrace, is at 960 feet, while Wawa lake is at 988 feet. The lake is dammed by a bar at the railway level (960 feet), and was a bay of Lake Algonquin cut off by wave action. Higher beaches reach 1,042, 1,088, and 1,140 feet, as determined by aneroid. At Goudreau lake, twenty-eight miles northeast of Michipicoten harbour, a distinct bar was hand-levelled from a bench mark on a railway ‘‘try line,” working out at 1,330 feet above sea ievel.! Thirty miles northwest of the harbour a beach was found at 1,382 feet on Obatonga lake, and about ten miles farther northwest, on Pokay lake, there is a beach at 1,445 feet (both aneroid levels). This is the highest water level observed in the region, but at several C.P.R. railway stations east of Michipicoten there are lake deposits at about the same level. Pardee, 60 miles east, is on a sand plain (1,524 feet), and Chapleau, a divisional point 70 miles east, is on a sand plain (1,412), with a higher terrace including glacial kettles between it and Poulin station (1,499). Eighty miles east, at Nemegos (1,421), there is a sand plain, and near Winnebago, 100 miles east, there is a gravel plain and raiiway ballast pit at 1,447 feet. Winnebago is 80 miles northwest of Cartier and north of the Sault; and it is evident that, as usual, the railway has chosen for its route the sand and gravel plains and terraces of the old shore. The position of the railway roughiy outlines this part of Lake Algonquin at its highest stage. Algonquin Beaches Northeast of Michipicoten Bay The shore of Lake Superior between Michipicoten harbour and Heron bay, where the C.P.R. reaches it, is comparatively 1This beach was first put at 1,430 feet, but later it was found that the bench- marks were adjusted to 100 feet below the level of Lake Superior so as to allow for under-water work at the harbour. cheiat) Waele AlYet rea), Ota SATA ie imuk th to “syoot Og ot RS ae agply ny 9?" s) ‘AA 2 i: foetal or, 8 Le ‘rn adit ae indy MAT Heda a Sei ch VF ‘rail te ie et . rane ht uit. ae ef wits ay ne nuTTGe es ee Pa atm wre Te) “Oh fi 36% fal) ni : rhe wis wl” Rete ee per Pt: Vie lf kta, vate tii. owl ABQ) dowel oor ay aed Se cd A ahi itd o ‘wii ‘sp } Ji, sLiki} ang), oe Laster iad bt tare BT, C ese Oe vee jen a olel aT ean +) jong eer , S51 Ue : Lean 1 Wo v foal pad or ’ 1 Rab FORT are 103210 BO was ; . He RM satel eps a ees AN ae rd) oe ry hi tes ayy | URE, T in i . a. i ; Ie" nsan a pe ; 7. br pine / “ % 3 nel leas a recnigant ants: si idee: rv » omy Diige:s re + Jeng solar Ww iby gnoe nm soo DY g ~ lipratae ie ee iron Jainedy yollvbpt et eh ea ey MY 7 - Op] Gide nite o ” sail . uikig heats ee ; J nw un only boats a See (+ bas tama do, ewe anhin > ap) ala STE ARE ee imi inahve ‘ ey ait @ huley ening itt ‘Twite ott af mie 3 Pare ttlyt OcLe i? eset it at | ya) wus ehietiis: (| asthe rif, ner 19d 100 vioviicinqmope tieadann 71 A od) sae aad or ml? ay eee sito: tid 199) CROAT ale : ip 4m Ly ia J iyi ett i peo. eee slot aie Ba tp ueiinies ese tom he oe tenga ated ley Sova eld woe icivins a PA Sa ¢ ALGONQUIN BEACHES NORTHEAST OF MICHIPICOTEN BAY 27 little known, since it is not touched by any travelled route and can be studied only by using fishing boats. At the mouth of Dog river, fifteen miles west of Michipicoten harbour, beaches have .Jbeen measured by aneroid at 721, 851 and 962 feet; probably all belonging to Lake Algonquin. Farther to the west the shore is mountainous and too rocky and precipitous to provide much beach material, and the same is true of the shore bending northwest toward Heron bay. At Kilkenny, a small fishing harbour just at the bend, excellent gravel beaches were hand-levelled at 610, 618, and 644 feet, the highest one probably belonging to the Nipissing beaches. «As the highest 1.0ountains in Ontario rise within a few miles of the shore (Tip-top, 2,120 feet, is the cul- minating point), it is evident that there must have been a large promontory or island here in Algonquin times. The railway levels on the C.P.R. between Pardee and Heron bay are mostly below 1,200 feet, though some ‘‘sum- mits” reach nearly 1,500 feet; but no well defined water levels have been observed, except possibly the flat on which White river (1,225 feet), a divisional point, is located. It is possible that the ice front remained for a long time in contact with the mass of high land between Michipicoten and Heron bays. The region deserves a much more careful study than has yet been devoted to it. Michipicoten island, forty miles southwest of Michipi- coten harbour, displays, as might be expected, a fine series of beaches, which fall into distinct sets; the lowest from 617 to 624, the next from 667 running to 680, and a third from 730 to 780 feet. At 806 feet there is a well-marked gravel beach and a lake dammed by beach materials. There is a wide terrace at 835 feet including a lake, and a highest terrace at 897 feet.!. The beaches above 680 feet may be reckoned as belonging to the Algonquin series, which, how- ever, is not complete, the highest members being lacking. The measurements were by aneroid. ‘Bur. Mines, Vol. S, 1899, pp. 154-5. LS i¢ | Y Mea iba x. ht) DASE 4, PEST if. shies a Bo. » - ’ : (fal a en : ' ) Vat Vv I “h sp) pia 7 | Aals Ai i fe eu “ues ane yore vi) lines oe ers rae, Ah Y ie : ty i gaya : bY hig Pa ‘oy By. 16. (hetwens eel tertiya parte aad henihe orate ot TULL \, Gb da) yitigarelal ite Bile saeiee oi yt tie ie AOR rnc er othe wa seu" ntti and ty balsa PPD hh. -1 sil hig fag eh: BS ig wenliiritwedeediiam ses Ht) Satay aid to om ; Jw cuel Pokal y frivoct only Js deat ponder gevtelet Lana Bebe i) i boty yi bf renee sitoagh ; sine ait han Snipa voce ae weenie ‘by 2) ni

-TRO‘MOLYA TAD { ee oy 7 nit erwin < (safe om tesa nerpigghé- ef } . We persiasicte On yan th ; Tt oe ieee P oui Peels ay ACH ‘indenel slash . mo raha es ‘ulotente fa hey: pred tpi POP, Ff) chit brent? Aine wie DBE % ‘nee oe eee 194: Oe sertt Beet nabaviva wai on ebeiie: coigualy estew ods ban ale Nus VA vril “oft do aoa vl yd ‘hone ) sis) Got CNet at oe ' ae wtf ec ast | ead 1 Les serail et te mete if | now ele Wal nf i nfs) te ely ile lew debt ae ; AOeTouTy itt i , bab budevotaan Oth 1a web sind 1a) Pere ojo nae Saees eer ont ey nds yeaa { jattos sblers any sea x shistigy ay Wah . Viet e a AY OAS B, és 7? ae ei tr 7 4 ae S ei: o Vey. ic a , “at i. r 4 eyes stk: ‘steiay es" et laval: ele sped 90a COSA a +e. Stal “cjunte Jenipote ae malt Pia c = eve jee lGet WG ISIS ino sae r wes ey : Plt ale re ad wy wrnttte 1} ; ITU hike lo ftst rie il (unt ernie seat aval om aa a." > i wen |v eer: * lo 1T2 de 7 - o eee int} >) = Settle sr nr ar end ine ' yes ity MODS sit, 105 ais - at at erpits aL wiag lives PALI’ of bee acl) bape figword wronhet Pus e's ¥ | te PO! ALGONQUIN RIVER 31 waters of the lake were divided between the Algonquin and the St. Clair rivers and finally passed entirely into the latter. The most careful study of the outlet has been made by Johnston,'! though it had previously been observed and described by Gilbert, Taylor and others. The route followed by the river was in the main that of the Trent Valley canal. A bay of Lake Algonquin, covering the present lakes Couchiching and Simcoe, extended east- ward from Nirktield to Fenelon Falls, where the Algonquin river began. It flowed through the basins of Sturgeon, Pigeon, Buckhorn and Stony lakes and then followed either the Otonabee or the Indian river valley to Rice Lake. at that time a bay of Lake Iroquois, which will be mentioned later. The old channel has been traced down the Trent valley to the Bay of Quinté and Lake Ontario, showing that Lake Algonquin survived Lake Iroquois for a period of time long enough to excavate a wide and deep channel to the present level of Lake Ontario if not below it. The present Trent river is far too small to have carved out the valley it occupies. The puzzling feature of this continuation of the Algonquin river channel so far below the level of Lake Ontario lies in the fact that between the draining of Lake Iroquois and the formation of Lake Ontario there came the marine episode when the basin of Ontario was invaded by an arm of the sea, which stood in the Bay of Quinté-region seventy-eight feet above the present Lake Ontario. The solution of the problem is probably to be found in the fact that when Lake Iroquois ended the sea was much lower than now, since a part of the water of the globe remained stored in the great ice sheets still remaining on the conti- nents. If we suppose that only half of the world’s ice caps had melted when the glacial tongue which held up Lake Iroquois at the Thousand Islands disappeared, the sea was 1G.S.C., Mus. Bull., No. 23, The Trent Valley Outlet of Lake Algonquin. 2U.S. Mon. LIII, pp. 410, etc. ee ee eee ee ee ee yh ns a vere j a ' a) f fs m1 ! | \ A i v.. " , 2 , s 3 al i 1 Ww j J K ‘ rh ’ 7h) TONAL : ee: a , let Hioopeeehy: ot rT et atk beatae Hib ey: v3 “aul is ; a Pe in aa Vea pet it) ety: mee yee bith. Bidens. main pred? ab", evan ani it AE Wes) ane i ‘aan bens. Wa pee We eer. ed res © bad’ tt ihe Ai ree we tiy cod oh era Prey » Tr een rth he dey ea “iti TAL ‘poverblta ei ik: fi Oe be: ek TO ee nena of /e Ls adypred eas Wei hae? pera: Ceetach ip ye, a a ") Pi catd ope alls i. deinen ob bk eet? 1 te CT ee iussonit bawoh. 3h Hotta ely ith enchhl Were ania 4 sh tod ARN et Colt Sib Aghbreh ‘3d 4 + corvieds od tiie, dally Jonpest aa 7 1 nih) are me i) v8 ah bah bases - filo ne ee 44) parce. eae * eats L eee ide ON 7 a, i ia 1 Leora BOA en lod boveaeie MA: ie » f Nestea ld ab De Pris" “ nord oll bir guetta ois at i ive olige-oed aan ea 1 aba of Maite y a Vein eryreny| r the Sates isis eae at i} (oO wieder yD bre akoupe Tt ential Yael Liban orhlok ~ ou es OStrIiert ahaa Nove TUT ‘ohms “alt Ves ae ma vd by aii BW, oni), ge Wels rieevion i ay inhi) (oLweee Lo ab . pe os one mln k 9 eh altyy My reqpere ee ae phisclonda ‘ eri we DT, Uae ee i bi log ; ee por: ite i Paris . aes nd VA st hci At) wo Daa oft iy geen ie teh Babes, wos oh bia athda Nee vist i ashe ee p iy omy poy _ksioay i ae Re ONT i hit is 4 ’ f, 4% ~ 32 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO 80 or 100 feet lower than now, according to computations which will be mentioned later. For most of its existence, according to Johnston, Algon- quin river flowed through the Indian river channel, one of the present connections between Stony lake and Rice lake. He infers this from the magnitude of the delta built at its mouth in the Rice lake bay of Lake Iroquois.! Ultimately the northward clevation of the region turned the water into the Otonabee channel, when an important delta was made in old Lake Peterboro, a little above Rice lake.2 The low marine stage probably occurred while the latter channel was in use, when the Algonquin river flowed down the Trent valley to the level of Lake Ontario. Life and Climate of Lake Algonquin At Tolleston near Chicago, in beds probably belonging to one of the earlier glacial lakes, many shellfish occur and also bones of several species of fish and of a duck, as well as leaves of oak (Quercus marceyana) and cones of spruce (Picea evanstont =canadensis?). The middle Tolleston, which F.C. Baker considers to correspond to the Algonquin beach, also contains fifteen species of molluscs. Another stage includes eleven species of heavy unios characteristic of the Mississippi, among them five which occur in the Toronto intergiacial beds on the Don. Except the spruce, which does not grow so far south as Chicago, all the species inhabit the same region now and indicate a climate like the present. One or two of the molluscs even suggest a climate warmer than the present.’ The Tolleston deposits were formed at or near the old outlet of Lakes Chicago and Algonquin in shallow water more than 400 miles south of the northern shore of ice, so the evidence is not conclusive as to temperatures in the lake as a whole. 1G.S.C., Mus. Bull., No. 23, pp. 11, etc. 2Bull. Geol. Soc. Am., Vol. 15, 1904, pp. 357-8. SIll. Acad. Sc., Vol. IV, 1911, Post-glacial Life of Wilmette Bay, Glacial Lake Chicago. —_ baa aah isi HtiOG ‘J : ' ? Mg “J Le Lae’)! LAs vh i 7 - i@ ' ian! aie.) arts ® % ef ‘ ’ utal itegellanolh, vente le Je Ae lals ld do -otureeeea SAD ‘aan viitaca Lakitibou Salud. }) 700..qc0) Sai poigen ih lo pathol: ie she Hartt mh. jee alk PMLA FITS raja “i OLOS Be 1 o* _* a1 (Pea dai ) = ced ‘ ae te aay j _ mae tots Seay ower bo re rave A) Lyvoeets a nincr ga pe exer endl. ai tayostl anaes o. int wtot aes « UE: eas es er te ee Losnnaks riuines Chae i nroie althil sahil ote Sd f cherenone, vasiaslorag onesarg 3 diol wy loninecawee of) noni int sole 0 1}. i wht holst Witt A i iT Ais re Ege not | ERGOT isthe at est ‘- yori wi. Hen i ah iM trivago ) ae Gi Denys 5 eh pnb. Ano; a, > oh Rae ames oem Y od Drea ne Op erage “4 my ) ecw! 2a ae { eioTi li uv at 1 a. Tat, ae 3 poe =f) » Mealilwy ey ood grown 4h ra HE ad ro nn? Oma pads Ta Gon! ee iipae 12 on OU LNs & OIF Mach Lam Wied Holes i wes HIT) ee llores ih. i te: Wiae by J 77 “> eet’? 4 rere “t te . z aa Fi ” ii ith it} - S eae onan ar, - aa pe pas. 4.73 oe att aid aes alti beni! Rares mr Wy 7 e LIFE AND CLIMATE OF LAKE ALGONQUIN 33 In the Algonquin beach near Jackson point on the shore of Lake Simcoe, Johnston has found fresh water shells in- cluding Lymnaca palustris, L. decollata and Sphacrium rhom- boideum,' all still living.in Georgian Bay. He has found them also at Roche’s point and Wilfred, and says: ‘‘The occurrence in the deposits of the highest shore-line of Lake Algonquin of fossil shells of mollusca similar to those living in Georgian Bay, and the fact that the fossil shells are only slightly reduced in size show that the temperature conditions of the water could not have been much more severe than at present. This is also borne out by the general absence of ice ‘ramparts’ on the abanuoned shores of Lake Algonquin and the rare occurrence of boulders in the lacustrine deposits of the lake.”’ In my own work fossils have been found in Algonquin sands 810 feet (aneroid) above the sea on Pic river, 35 miles above the C.P.R. at Heron bay, including fragments of Unio or Anodon, Sphaertum, Pisidium, Gontobasis, Lymnaea, Planorbis, Amnicola, Succinea, Valvata, and probably two or three other genera. Unfortunately the species were not determined, but all the genera still live in Lake Superior.’ Many years ago Robert Bell reported shells from Pic river 30 feet above the water and the same distance below the top of the bank, but it is uncertain whether the deposits belong to Lake Algonquin or the Nipissing Great Lakes. They include two Unios, an Anodon, and a Margaritana, with species of Lymnaea, Planorbis, Valvata, and Ammnzicola; ‘the whole being of a more southern type than the mollusca at present inhabiting the rivers and lakes of the neighbour- hood.’ Three miles above the Mission on Michipicoten river near its mouth there is a deposit of peat and logs six or eight feet above the water. An expert lumberman has recognized white pine, jack pine, white and black spruce, balsam, cedar, 1G.S.C., Mus. Bull., No. 23, pp. 15 and 16. Bur. Mines, Ont., Vol. 18, 1909, p. 153. 3G.S.C. An. Rep., 1870-71, p. 328. W i aA 7% Rwitical + ee hw] | 4) wares wilh tt ite. ‘7 ‘} { i '% >. FF bal de nik vcr tevin neiaho Ludden gam as elton vation ecatlgert io etintadt oh 1 aay ont tap @entel: Das eTav em, 5 ota wiubl bes BRAC) a nz rs ee COM mepiteh ys feu ee jtigeind spurge 3! aC 8 man Sls ah a a Pees ate ez Miro Ce “a : bp oo) Seed a i. rete Mot kes) eye Toga orto ayit ol eget weaytt a iwi Li aay ou rely nd hii, St rie: Higa, ‘ yi like wre cea iniuny e- lypatie rbieviimtea jonilosit 4 area a i linha: reed jus a y lou Pay ai + q yi) Oe » oeteebeveetnennid ely ratte , wire. } a. rs Wy at wi! fa (we ! ( 5 } tetas yh) a ot) Segal wiice eae rh o “ev ceey aia i egy wt (bien a j . be segh) aah’ | “vein 5 vailes gvergt (iS SEN) ej/02¢ SF SO SaAMWT LVAUD INISSIdIN ‘40 dVvwW 36 GLACIAL AND PosT-GLACIAL LAKES IN ONTARIO The Nipissing Great Lakes lasted long and formed a pronounced beach which has been traced almost all the way round their shores. In area they were only slightly larger than the present lakes, though there was an extension toward the east covering the valley of French river and Lake Nipis- sing and ending at North Bay, where the outlet river began. A detailed and excellent account of these lakes and their shores has been given by Taylor,! and it is not necessary here to give more than an outline of their history. Rise of the land toward the northeast gradually restored the use of the St. Clair channel and during most of their history the Nipissing Great Lakes had two outlets. Finally the rise toward the northeast turned all the water south- wards, the North Bay channel went dry and the present lakes came into existence. South of the ‘‘hinge line’? mentioned on an earlier page the Nipissing beach is horizontal, ten or twelve feet lower than the Algonquin beach, and about 15 feet above the present Lakes Huron and Michigan. North of the hinge the Nipissing shore is split up into a series of beaches closely following one another. At the North Bay outlet the highest Nipissing beach has been fixed at 698 feet by Goldthwait,? and to the south it becomes lower until at Sarnia near the hinge line it is only 597 feet. At Sault Ste. Marie it is 651 feet above sea level and on the north shore of Lake Superior it rises to its highest levels, given by Taylor as 703 feet at Jackfish bay and 710 (aneroid) at Peninsula harbour,’ the point farthest from the hinge line and a little north of the isobase of North jax. Probably the lower terraces along Pic river represent the highest known portion of the Nipissing shore. Delta materials are widely spread along the river valley; and one sand and gravel terrace twelve miles above the railway bridge and eight miles from Peninsula is at 718 fect (aneroid), 1U.S. Mon. LIII, pp. 447-462. *Ibid., p. 459. *Ibid., p. 460. a “=~ 1 | re a ed) AY SOR Is eo CcIya ‘saa ' * . _ hel ;- Ste \ihviea! (7 wae) 7h: hn) ‘yriget ae ct ’ fad nen 4 iid pend hil ava it aga l, My: é f ef tbutis jae tisay °/ vi) OG: at narra | fe b lnciwe: conleien mi are “4 tis Atengsty, ony inl Pas poss wel? sas. bbe en rie Oe villi ah quay cr ition | wotwer Cote er pate orn ira, 38 anh P SOA RegAT ae suns ae FoR!’ OF OP Wik. Al yn hy: ia wig oe -vatwilt: yea? 1 “ems ih Hilt 010 iv lip Love Thiel tient Soft t SAU "rent : +4 , ny atrraliy Pale Pacwians “+ lt Japa Gs 1 ay ded feet Outi), eae Eee fre . wit lia feild tenia Th Od oe ee bortend yok trie Ee } ; Pole a 1 OOF “on AS. pi Donning i endl sgnia’ * off a 1 eT 4b ssa yt ih ‘Hoa | ‘- murvin, seneel Cokes i .doewl ape n ay Te TEP a al iti all, Dis pies nade ern av _— a rong , r, ? =. 7 ee ee ; ic tAimedybles ) eer yon B00 4a! ve cs naive sruanl alt non sete lind. t rhe P sc deceie dk 100 of le ee bgt woe atet lo aie ict Aatde = anh SU ee ove Tegan tondtied jai ot ye cfige Zee: ot hax bVi Qo comedian One at ‘hon 3 me “me , i yoole aera oil. vis _sneale. atiinerqit, oft ey aera wae bon jwolley vewtt ods -ghol. pears y wicwler: an. wrod, aalit riod ook 8 At wos muvee ae a! aiuent ri) “| pers = » = tins wy — ee a ee at _ J - ae o ae ES * tee eo soa: A ee is ea Fmt v- rd £ opi 6. Nipissing Beaches at Brulé ~~ : 1 Sah nN be a‘ i 1 ae a " : wa apy.) . +, To P a, ‘sl No: ‘ war y +e “une! a “ hata” => ig DPR 04 oni \ * 7 aN a4 weg) i aa \ ' ' 7 ry] > ed AO Den Wi 6 vail eeelt - aE le a a ea eter ee te tod THE NIPISSING GREAT LAKES 31 ~ and no doubt belongs to the Nipissing shore,! while others a few miles up the river, somewhat higher though not measured, probably represent a continuation of it. A few other localities on the northeast shore of Lake Superior may be added to those mentioned by Taylor. The highest point after Pic river is probably at the Mission near the mouth of Michipicoten river, where terraces were measured by hand level, beginning at 623 feet above the sea (21 above Lake Superior) and continuing with com- paratively small gaps up to 702. An interval of thirty feet separates this series from the next, which may be considered to belong to the Algonquin set of beaches. According to the isobases shown on Taylor’s map, 702 feet is six or seven feet above the proper level. On the other hand, a broad sand terrace at 651 feet seems too low as compared with the isobases. Which should be taken as the highest Nipissing beach is uncertain, though the 702 feet level seems to the writer the more probable one. Near Brulé harbour, ten miles south of Michipicoten, very well formed beaches were hand-levelled with only small gaps from the present lake level up to 658 feet, which may be considered the highest Nipissing terrace. On Michipicoten island, just touched on the north by Taylor’s isobase of 676, aneroid readings show a first series of beach ridges up to 22 feet, where there is a sea cave, a second series up to 65 feet, where a terrace affords space for several houses, and above this a succession of faint stages up to 78 feet, followed by a gap before the next water level at 128 feet. robably the beaches at 667 and 680 feet belong to the Nipissing set and the higher ones to Lake Algonquin. Life and Climate of the Nipissing Great Lakes Shells have been found in Nipissing beach gravels at various places. Taylor mentions Unio luteolus, (= Lampsilis luteola), Sphaerium Striatinum, Lyninaea elodes, and Gonio- *Bur. Mines, Ont., Vol. 18, 1909, p. 153. i aE 0S aS aia ne Slate A Oe ne occa nen a Rate " . her a ¢ al Te ‘ =) % : ™* , , r i} <« 4 5 ; ; i rt Vs A@ j yb. a 7 i a iat 4 sire a a | - Vf : ‘ Hs # = = a a 38 GLACIAL AND PosT-GLACIAL LAKES IN ONTARIO basis depygis as occurring near Cheboygan, Mich.t These are common forms in the Great Lakes at present. Many years ago Dr. Chapman, professor of Geology in the University of Toronto, described shells from deposits on the banks of Nottawasaga river near Georgian Bay. At Angus on the west bank of the river 30 or 40 feet above Lake Huron he collected Unio (=£lliptio) complanatus, Cyclas similis (= Sphaerium sulcatum), C. dubia (= Pisidium virginicum), Ammnicola porata, Valvata tricarinata, V. piscinalis,, Planorbis trivoluus, P. cam panulatus, P. bicarinatis (=antrosus), Lymnaea palustris, and Physa ancillaria. This point is about 20 miles from Georgian Bay. About twelve miles from the mouth of the river Dr. Bigsby found two layers from four to six inches thick closely packed with unios. He adds JAelania (=Gontobasis) and Paludina (=Campeloma) to the list just given.?, Probably shells and wood found at Owen Sound belong to beach deposits of Nipissing waters also.3 To the species secured by these older geologists may be added the following shellfish obtained by the present writer from the banks of Nottawasaga river: Sphaerium rhomboi- deum, S. sulcatum, Pisidium noraboracense; Valvata sincera, Amnicola limosa, Goniobasis Livescens, Lymnaea desidiosa, Planorbis deflectus, P. parvus, Succinea avara, Polygyra monodon, and Helix (=Polygyra) tridentata. The collection includes nineteen species, seven of them mentioned in former lists. All of the species referred to above are still inhabitants of Lake Huron and suggest a climate similar to that of the present. A very interesting set of river deposits along the Niagara, from Queen Victoria Park and Goat Island to about the Whirlpool, is probably of the same age as the Nipissing Great Lakes at their latest stage when the drainage was partly through the St. Clair, Detroit, and Niagara channels. 2U.S. Mon. LIII, p. 452. 7G.S.C., Geol. Can., 1863, p. 910. 3Ibid., p. 912. t - ° ne 4 4 A M { a) e \ \ + a, ou . ' e. \ NM 4 ¢ vi + S i i] ; 5 Ay = ' + F - . 4 t = : i Zo sy y i Ve oe a Pu eee te at 7 AP ees’ 12 ‘ ‘ rT te - F ; a * + , P *) - al iy 7 | J sce Uf i J f i ot Beak 7 7 i 3 ! G, Sten 4 (irae , a) ; b 7 . i } 4 - A » + en oe ~ ‘i “yi eee or . 7 , \ | oe: ie: vy ie ee ee es LIFE OF THE NIPISSING GREAT LAKES 39 These are mentioned in the Geology of Canada (1863) as containing Planorbis bicarinatus (=antrosus), Physa hetero- stropha, Lymnaea caperata, L. stagnalis, Melania (=Gonto- basis) Niagarensis, M. conica, M. acuta, Paludina decisa (= Campeloma decisum), Amnicola limosa porata, Unio (= EI- liptio) gibbosus, U. (=E.) complanatus, U.(=Obovaria) ellipsis, U. rectus (=Eurynia recta), Margaritana (=Alasmidonta) marginata, Cyclas similis (=Sphaertum sulcatum) and a land snail Helix (=Polygyra) albolabris?' Miss E. J. Letson, in 1901, published a list of 31 shells from Goat Island with a description and figure of each species. Of these 24 are in addition to the species given above. They include Pleurocera subulare (=acuta), Goniobasis livescens, G. haldemant, Amnicola limosa, A. letsont, Bythinella obtusa (=Amnicola emarginata), Pomatiopsis lapidaria, Valvata - tricarinata, V. sincera, Campeloma decisum, Sphaertum stria- tinum, S. stamineum, Pisidium virginicum, P. compressum, P. abditum, P. ultramontanum, P. scutellatum, Alasmidonta calceola, Quadrula solida (= Pleurobema solidum), Q. coccinea (=P. coccineum). The present writer collected fourteen species of shells in Queen Victoria Park, a number of years ago. Of these five species are not included in the foregoing lists—Sphaertum solidulum, Unio luteolus (=Lampsilis luteola), U. clavus, (= Pleurobema clavum), U. occidens (=Lampsilis ventricosa) and Margaritana (=Alasmidonta) marginata. In addition cyprids and chara may be mentioned as well as bones of mammoth found by Hall many years ago. In all 36 species of shellfish have been obtained from the river sands and gravels near the Falls. Most of them still live in the river, but four of the species, Lampsilis ventricosa, Pleurobema solidum, P. clavum and Alasmidonta marginata, it is stated, are not found in the lakes but are Mississippi forms.’ They must have reached Niagara river in a round- 1G.S.C., 1863, pp. 913-14. *Bull. N.Y. State Mus., No. 45, Vol. 9, 1901, pp. 238-252. ‘Stockholm Geol. Congr., Changes of Climate since the last Glaciation, pp. 386-7. roe y eee errs Pes. ee eee oad in ig ; roe see gy a, nits ot al yon a oF | , * re ye ah} De Neen Hh BUT 30 ae (hIIRES wD YO TG" vt ae STG, fu. oe win sin 1 , UG ieeeina )- en Largiand none rvuda,) j my na rel {ehaey ny i] e 4 ASDADTD Ls ante ner natenitlh Ss thie ST, POY me Rat yy, im) Oral, ailnany aontek ojo ue niet | (ont Tr A jays ty! ‘—— a ais ie hye fhe 4 Sun eee Sete he he © haere Uh Oi yt pune tie ; iMiuiy APR Mey) bah WEN yi ba ee soete v Mead te ale yee ge OTH AE be oobi M4 h 7, ith coed act Hone eS ohiy a hee! noise Sanaa Mala ty eda paving lee 243 OF) se rd i { ‘ie Ne Pe? ie Tk? as | yee oS “i 43 Pan 1% re, py eT > ae 1 atid alee . ' = ay dn Hlboi alae, cheba OK. Uns jens ia reas ‘ Oyen fe Anon ie > i} } pee TABI Gh ao me ee bes! z PANE 3 me AG wanting un 8 ua cuhderaaal act wie babiiaay \ q if Neri bel tea 1s A Ser \ ee tosh eh Tews, na ts Kee vito (ote eit lay nun “beater tie od ae ie “ mo: PROV Yt ttelt a 5 > rey ra fe Seog icf vad Hedlbite ne ryt Ohh 431) oe “ile to 1 ¥ ritaty th Pin 7 ig, ia TO eG wel “eit ry) buh ae pve ly regniv’ ba haere OVE ae ; * , i 7 ns ‘ id é Lie & TE a) bf aq oer is 1 age mn itaed ia! mets fete wlagt) ie wean oes Bier? Uy Gate ae ar a: = y Lf i. ag f ' = e -— u aa oe I ’ ¢ "ie 40 GLACIAL AND PostT-GLACIAL LAKES IN ONTARIO about way going north from the Chicago outlet, when it was in operation, through Lake Michigan and then south through Lake Huron, St. Clair river. etc.. to Niagara Falls. Whether the Mississippi shellfish should be held to imply a milder climate than the present is uncertain. The three unios mentioned and four others occur jn the Toronto interglacial beds associated with trees now growing in Pennsylvania and Ohio, suggesting a climate 4° or 5° warmer than the present.} Twenty-one out of the forty interglacial molluses occur also in the lists from Niagara. As the last remnant of the Labrador ice sheet had dis- appeared from Ontario south of the Hudson bay watershed it is very probable that the Nipissing climate was as warm as the present, and it may have been warmer, as suggested by the unios. Lake Ojibway The earliest and highest levels of Lake Algonquin reached 1,400 or 1,500 feet north and northeast of Lake Superior, as shown on previous pages; and it is known that there are at least three passes across the watershed toward Hudson bay which are very much lower than this. From west to east these are the passes at the head of Paint river, northeast of Lake Nipigon, at 1,046 feet (railway level); at Long Lake 22 miles from the north shore of Lake Superior, at 1,040 feet (aneroid); and at Missinaibi 45 miles northeast of Michipi- coten bay, at 1,090 feet. It is evident that without some barrier Lake Algonquin at its highest stage would have extended indefinitely north of the watershed. This was pre- vented by the position of the ice front which stood at the divide or somewhat to the south of it, when Lake Algonquin began. Pitted plains in several places show the junction of ice and water. As the ice retreated the land rose toward the northeast, but not rapidly enough to prevent Lake Algonquin, when 1Geol. Congr. Mexico, 1906, Interglacial Periods in Canada, pp. 15, 16; also _ Geol. Congr., Toronto, 1913, Guidebook No. 6, pp. 15-18. - ee Fae eg So ee a nt NS ei =F Sa re RE Sa aie wa MSI SEC sie Aa Sy a i ne my o i Gud 7% Sey. o_o paw - ‘hae = ais A = ee i mT! YE Cra es) Fae. = i : £ 1H iy ett yan dit = yale ‘ af ' ry) ¥ if = « a ; q vid erate HAE at att y : itn six. , Heat an 1b ee Lee 4 i! : | af Q f ‘i wy cy int =) ae disludit chal hiaitte eat act ns ; ' A # rt a8 US es i raul = f 4 vy Aa + OF hin eRe ge meat te a Wahnyoee oe gue uO bata ai ean rh Ay abo oe yay ns weal etal onan olga he tate ae aniteray yi Sims | Z fds ye tt mae be Hh nea rie) oF Mt ) oh f = /. aft} sat Adele — . “vii ; “e nal eee Qe ae i ¢ = i Bite | ce ee at saottgi b ‘uci Lieut Arian aA ay th eae 4 stent eee epepta #aanth to ieen atte Ja i sence | . Veena 990 aoe rr “1 | tik Tae. me i, A? ie Lines Linea Act aye) My Oh fh ‘ ; er : (art wits 1 ie st heute ann eee ! oe este) pale iY lok ils | OMT om) gd 4 ating lartg ten is ent 29 7103 bysdeerti2 eee: beng tor | front ve sitet itt, oat) Basi i t4 ar is 4 intone 7 4 pens he. Val earopnt bali dels . 8 ee a a er Gy LAKE OJIBWAY 41 its waters stood at about 1,100 feet, from extending bays across the lowest portions of the watershed. These bays seem to have been of short duration, since no well-defined 7. Map of Lake Ojibway shore forms extend northward from the Algonquin beaches at these points. This was demonstrated for the Missinaibi col by Taylor, who decided, from lack of beaches north of the divide, that the Algonquin water, as it had been named I yawerd SR 4 * > af + a | - y ee a Mp SE ; € oh Py iets \< iy ; i Pi + ,i™/. y ‘ - Sat ’ " A a ‘ Aste Pon nt it at | ; on pea 1) beNt Lee ) ee j tid Hii: ae f ¥ / = f * f 7 : ta wigwed i} mat toga NA asnons tsi Mmolh, i dae beudwe Steel aaa ip) bev scent i oe andaten 1a Avei in oh iy eo: One " ge rotate wi i ° t “ ES RTE Cale aE al tele ie 42 GLACIAL AND Post-GLAcIAL LAKES IN ONTARIO by Spencer, was not a part of the sea but a separate ice- dammed lake.! The northward rise of the land presently lowered the Algonquin shore to 1,000 feet or less and the northern bays were cut off from the parent lake. Probably they formed small lakes between the retreating front of the ice and the watershed, and ultimately merged into a large body of water which the present writer has named Lake Ojibway, from the Indian tribe occupying the region.? The presence of a great body of fresh water in the position just suggested for Lake Ojibway is undoubted, since its deposits form the wide “clay belt” of northern Ontario, covering, according to explorations carried out for the Government of Ontario, an area of 25,000 square miles. The clay belt shows in many places beautifully stratified clay in annual layers from a half inch to an inch or two in thick- ness. The clay deposits have an undefined limit toward the north, but on the south pass into sandy shallow water materials and sometimes show beach gravels indicating the shore of the lake. Unfortunately, the region i. still only imperfectly explored and is largely forest-covered, so that up to the present the exact boundaries of the lake have not been mapped, and it may be long before this work is under- taken. Toward the west the clay belt ends northeast of Lake Nipigon; and toward the cast extends far into the province of Quebec. Whether the whole area of clay should be included in Lake Ojibway is doubtful. M. E. Wilson thinks that the Quebee clays were laid down in a separate body of water, which he names Lake Barlow} He is probably correct in supposing that a lobe of ice occupying the depres- sion of Lake Timiskaming separated the two bodies of water, at least for a part of their existence, and whether they united for a time when this ice lobe was melted remains uncertain. It is very desirable that some one should go over the ground more thoroughly than has yet been attempted, particularly 1Am. Geol., Vol. XVII, 1896, Piao. *Bur. Mines, Ont., Vol. 18, 1909, pp. 284-293. *G.S.C., Mem. 103, 1918, pp. 140-145. a ed be 4 { i ‘ « u ’ 4 gaan Tt) artes veto Aue ies MA abi 7 BS ; i A | re j \ e ‘ rie - a 11? * A he A ee a Sk ad , hats <5 sige 4] a? ‘ier yy Oe ibtine v0) A ES t Ais he feet , kaa bie 0 quiche tht St a Sagl HP OO4, b ae are ol Widetior in wis t q lt ee Ae vie; ih with: e2xi25 sit amed Leo ” buy ae ny (WAG ton a rf Leman hah swat. TO yi] wild at Nee eet bata cds ance toed Sa 7 ‘ iifa’ vo ' oi Hy | 309 eye) ore a " ‘ > . ~ * ba ‘wt * ty iba odd VOR) Seas ite ate cit arated Leg rat a ae ‘bi 0 | (3 iS fs Hons Ge) oon? > ly by Au aed Kone Yiott: aL, i 4 Laine, ooarternhae | | e arf ged iA Oe” a aa now? ote ass ene ’ Aypawt were tp" , ines Solty ont | . ; gush i ‘bh hod oot ae . nicl Wal oct eatwiaith ; ce 4 ' x f ay ong THAVRE: deine iad oat Do lovta ot yee .. eee age 1 Syrah RD: Pre tih ein pt aoe ny iA pel +f ote egrten on =i ¥ er ee tay chol & ox Joo ot oh) bétoaqeeie | 1 onde sedtexive en goaarelaee ty iis oomriveey bybioct fay wikel wst'as 3s ee ‘4 Laure ont eee aie bintvia ‘onto 5 ao omy sae ae viper {ee OPIS Bo a ges: adie ; ‘ CBR ae Pee = ait 7. be Aa ac: LAKE OJIBWAY _ 43 in the Timiskaming region, where a number of problems remain to be solved. The point of outlet of Lake Ojibway is not certainly known. It was believed by the writer, when the name was given to the lake, that the outlet was by the valley of Lake Timiskaming into the Ottawa, but if this lowest part of the southern edge of the clay belt was filled with ice some other channel must be sought for. The waters may have escaped southward along the edge of the ice, in which case the channel would be a shifting one, moving from west to east: or the outlet may even have been over the ice, at least for a time, so that no direct evidence of it would be left. Stratified clay is found on both sides of Lake Timis- kaming toward its northern end, reaching levels of 642 to 776 feet near Haileybury, New Liskeard, and Uno Park, and of 648 to 796 feet at Baie des Péres on the Quebec side. Farther south near the mouth of Montreal river sand and clay terraces were measured from 624 to 811 feet. The measurements were made partly with a hand level but mainly with an aneroid.t!. The clay beds were laid down probably in moderately deep water, but they are south of the -watershed, which is at 935 or 940 feet, and cannot be considered as belonging to Lake Ojibway, though they are just like the Ojibway clays. It seemed probable at first that they were deposited in an extension of the Nipissing Great Lakes, whose outlet is only about fifty miles south of Montreal river where it enters Lake Timiskaming; but the finding by Gilbert, Taylor, and others of a channel draining the Nipissing Great Lakes into the Ottawa river at Mattawa, only 488 feet above the sea, seems to make this impossible.? Johnston, repeating a measurement made by De Geer, puts the highest marine beach at 690 feet, near Kingsmere, eight miles northwest of the city of Ottawa, and suggests that a narrow arm of the sea extended to the head of Lake *Bur. Mines, Ont., Vol. 9, 1900, pp. 177-8. 2U.S. Mon. LIII, p. 448. > | > Ae i evi be, voaleenth’s) 22900 nop grein dali 704 i | | 1 7 ~ ih ] ¢q mot % i‘ Tyts ef « ae | 12 ‘ulgoges ite trad 26 BR av ' rN vu i i = , 3 Aj 14) r tHAS id “4 Lorena AY Al djitet. 16 sali, a Pipl «wand hi, of fella ene Stel wae + ott jo opbe aaa ay On eto ynighed wit yoo a “yott A ow 1) oe eee (Fs, 12 mi ogz t ia ra} wey. yu a 4 i alist sl A dion oft) of cen) Lane ae A “a “ae 7 shat. : re frets ctl 14 ee sequel eit oUt . tceorrey Le int 2901. 008 Ie a” HO lo a pi Le abana : a %, RB er 44 GLACIAL AND Post-cuaciaL LAKES IN ONTARIO Timiskaming; so that the clays were probably laid down in a northern tord of the Champlain sea. There is no evidence that the water was salt, since marine shells have not been found in the Timiskaming region. In accounting for the clay terraces above 690 fect it is wey fo reeall_ the fact that the, marine. water-level rises toward the north at the rate of three fect per mile. The suggestion just mentioned does not solve all the problems of the relations between Lake Ojibway and the Nipissing Great Lakes, for the North Bay outlet of the latter lake is given by Taylor as 698 or 700 feet, and since it is 50 miles north of the Kingsmere parallel, differential elevation would carry the marine level 140 feet above it. Apparently the North Bay outlet must have been opened considerably later than the highest marine stage at Ottawa, when elevation of the land had proceeded to the extent of more than 140 feet; and we must assume that ice still occupied the Nipissing-Timiskaming region when the highest marine level occurred at Ottawa. To return to Lake Ojibway, the retreat of the ice on the north of the watershed at length gave an escape for its waters toward Hudson bay, and the lake came to an end. It was probably the most variable and short lived of the glacial lakes of Ontario. When it ended Hudson bay reached much farther south than at present, at one point coming within’ 150 miles of Lake Superior, but there is no evidence that salt water encroached on the lake deposits. Estimates have been made of the time covered by Lake Ojibway by determining the annual layers of clay in its deposits. Paker states that the clay reaches twenty-six feet in a railway cutting near Ground Hog river and that the layers are from a half inch to occasionally three inches in thickness;! while M. E. Wilson estimates the clay beds of Lake Barlow as averaging less than twenty-five feet thick, and gives the maximum number of beds counted as 250. He suggests that this represents the number of years ‘Bur. Mines, Ont., Vol. 20, 1911, pp. 231-2. 4, 7 i i = 1 een a 7 J — ot 7 y Ba : a -" i 7. . Le [ ¢ ¢ 4 ‘ Fy oo ¥ re ' fa i ’ 6g a t ¥ \ : ‘< i ‘ : i9 Fe d i ~ 7 (es ‘ 4 Nc a ane t “ ~ p i . p« ad 5 : j ‘ if AY sort ¥ i hat | ie q id 7 a. 3 Fy ete i rp. FO afi’ vf LI ” a You: ®t ae alate =~ began. Pe te is y * _—— = it ot Wi wry ee , i} Tae *) ah hi cat + i ri : ‘ 1 / ’ ay teh e d * - io Danae ; . a oa =f LIFE oF LAKE OJIBWAY 45 during which a given point was covered by the lake.t W. A. Parks found a thickness of 40 or 50 feet on the shores of Night Hawk lake? and the present writer estimated the thickness near Matheson at about 50 feet. Railway engineers report a still greater thickness found in bridge work near Cochrane. If the layers average an inch in thickness, fifty feet would mean 600 years as the minimum duration of the lake. The fauna of Lake Ojibway must have been acquired from the bays of Lake Algonquin, which crossed the watershed toward the end of the history of the latter lake and the beginning of the former. Or'y one record is available as to its character. M. B. Baker reports Ammnicola porata, Lymnaea elodes, L. pallida, L. umbilicus, Planorbis bicarinata, Succinea obliqua, Valvata tricarinata, and a land snail, Helix striatella; but does not mention the locality where they were found. In all probability the Abitibi lakes and other smaller sheets of water which have succeeded Lake Ojibway in- herited their inhabitants from it and ultimately from Lake Algonquin. , The shellfish listed above do not indicate more severe conditions than the present. The northern shore of melting ice was perhaps largely covered with débris and seems to have had little effect in chilling the water of the lake, especi- ally on its shallow southern side. Waters of the Ontario Basin Lake Warren, one of the earlier stages of water, occupied the southern part of the Huron basin, the whole of the Erie basin, and extended along the south side of the Ontario basin; though most of the actual area of Lake Ontario was still filled with ice. The outlet of Lake Warren is considered to have been across Michigan to Lake Chicago and ulti- 1G.S.C., Mem. 103, 1918, pp. 141-5. *Bur. Mines, Vol. 8, 1899, p. 175. *Ibid., Vol. 20, 1911, p. 233. Val? te 74 wda ites adele 0 i) ule nore si Pease ma? 2 oe +P 7 y anh sity ‘J Tth 2 AO vue ay a : i) 5 re aA 7 a Tie an wv ok Sees My wt wher th “al aa odd tite ria ' Hon 4,1 bobo aie “it oe vue * wt way ale MW va ee shag re eg Dee's Mee | i. wwe «ei 1 al pd spray? Au pat cartes vt neat. my Ser. Gla ah! ia re + . rei 4,7 ih nome? cT ah) | “a Ss | isin] i oh ut ti Lia \ { ‘ oe ) : ole ye. = tal) oie : BD at FY: , oy in has « b deg Lie es it t} vind : i i 6 ‘ ‘ - 4 ie 1k i i Bao att I 1 ¥ 4 ey Me . > ~ ‘ Wiser ia 45 inf iy em Wate Ua, senate bch eh bee ‘i sees) bg idee IS Le iote badly ‘3 i ~cbipgi ze SET THE ’ rloaw/e et veges a ee ee cv Oe ters Cit ROB oA S hy af “yrqant Q\aniin y bie qi Gy eg solinae Ties ‘hse ; ity Orit 3 ‘matt aii tay fave “ob weal iis. miiot l?) 2a i eer 4 st) muy egies ~ + vie by om Pad oa Te nye, . i a | 14a. 20th a Parte Sronboiy ayeq jo dvyy ‘g -——--pu Sionboy, e407 NVINATIOD ‘d‘V i] S10ndoul AyWT mae “Chana « c ‘ ” vai Cbpya nosag” AL Bs the Rr ; 3 5 é t ¥ : 3 ¥ c z > t ~ ~ t Z ; 2 See rate ge °K Bee LE mee: qlee Sea ceite nee Bes Iroquois Shore, Scarborough LAKE [ROQUOIS 47 _mately to the Mississippi. As the Ontario lobe of ice with- drew, lower outlets were opened toward the east, at what has been called the Lake Dana stage, and drainage through the Mohawk valley to the Hudson began. Finally the waters were lowered still further and Lake Iroquois came into existence. An elaborate series of beaches and outlets has been worked out in the state of New York, especially by Fairchild, and those interested in the matter will find reports and maps of the different stages in Bulletins of the State of New York.! Lake Iroquois Much the most long-lived and interesting of the bodies of water occupying a part or the whole of the Ontario basin is Lake Iroquois, first described by Gilbert in New York and by Spencer in Ontario, the appropriate name having been given by the latter writer. This was the first of the glacial lakes to be defined, and it has furnished the criteria by which the others have been studied. The abandoned shores of Lake Iroquois were the first on which the differential eleva- tion of the beaches toward the north was demonstrated.? The shore of Lake Iroquois has been more completely and certainly worked out and mapped than that of most of the other glacial lakes, and the part within the province of Ontario has been studied in detail by the present writer.’ Lake Iroquois began as a narrow strip of water to the west and south of the ice lobe, but rapidly increased in area as melting proceeded until the whole Ontario basin was set free, when a considerably larger area was occupied than that of the present lake. The west half of Lake Iroquois follows closely the outline of Lake Ontario, but usually at a distance of from two to ten miles from the present shore. 1Bull. 106, 1907, Glacial Waters in the Lake Erie Basin; and Bull. 127, Glacial Waters in Central New York. Falls of Niagara, G.S.C., pp. 277, etc. *Bur. Mines, Ont., 13th An. Rep., 1904, pp. 225-244; also Bull. Geol. Soc. Am., Vol. 15, 1914, Iroquois Beach in Ontario, pp. 347-369. 48 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO At Scarborough Heights the Iroquois beach has been cut away for half a mile by the present lake. At Trenton in Ontario and at Sodus, New York, Lake Iroquois spread to the north and south far beyond the Ontario limits and reached a width of nearly 100 miles from the northwest to southeast. At the east end of the basin the northern shore is wanting for a distance of about 70 miles, where the ice barrier stood. With the exceptions just stated, the Iroquois beach is as complete as that of Ontario, though cut through at various places by rivers. From Quay’s Siding, about seven miles north of Port Hope, to Hemilton, and on the south side to Rome, N.Y., the beach :s continuous, but to the northeast of these points it is split up into several strands which tend to spread apart toward the north. - In reality the line from Quay’s to Rome is a pivot about which the water level swung, rising in a direction N. 20° E. and sinking in the opposite direction. Yo the southwest of Quay’s the earlier beach levels are buried more and more. deeply under later beach deposits. This is shown at the reservoir park in Toronto, where beach materials go down 70 feet, and still better at the Desjardins canal cut near Hamilton, where undoubted beach deposits reach a depth of 83 feet. It is evident that an important change of level was going on throughout the life of Lake Iroquois, the north- eastern end rising either continuously or at frequent inter- vals. The differential elevation of the northeastern end of the beach as compared with the southwestern amounts in Ontario to 460 feet. At Hamilton, the lowest point of the latest stage of the beach, the present level is 362 feet above the sea, and at an old island near West Huntingdon, about twenty-seven miles north of Belleville, the beach is more than 744 feet above sea level. Climate and Life of Lake Iroquois The beaches of Lake Iroquois have proved fossiliferous near Hamilton and at Toronto. The splendid gravel bar extending north from Hamilton toward Burlington has supplied many animal remains including mammoth, wapiti, '@ a Ardara) ef wa laneehe 7-0) CSA re i? ei . ; — ve tunnel wh Aainkse! etal td 71 ryt TQuge é 34 a wrty Se is ot ee wight Re ae ‘ pis DI cee a ec {ida oly We ee cre oe, > oor fluaint) “ ii} ig “wal JB P (ore teal a . a J ols a be ea Fae iMi4 oviik +) ; ie “4 J ma ; f 4 ae es . Srey wt, AL ee ee mt? to on‘. q 4 = . fr ioe 7 7 : t ae inet! oa | “rt 4 2 2 "1 "7 ,4 b | mn i {. le aia , ‘ : 4 ‘i> Sa! rl x rei vt wii prIzo bal: ; Pa st oe die he boa nrhtt) Fis Wigton) aad) Ms ahs 2 Oi ye athe JO sie 2a x hey iiath ob eae | tad atts oD) 1a ai iT at ‘ a Seat’ . ; | 3 ai) Lowes Pam a rin oe 1 aril Pumatieil ran $e iJ uee “ae ) "7 leuk: oleyyoy, eee age | sq Wo: ud ; i). dene *igvaeer ae 7 ire Law a Bis sh a rh oq arite- = . ont ri} ‘ne tua hy iM iy \ ee Pots T1458) r cle? tailte 1 ' gail a oyu Zar aot on wee Eten ise walt a) a, r oti a4 ed A) fab oyhty! leh sae i ; tore of) Le (Odi pootionth ae mt Oh reat zouk 208 Ube Meenas bee sf Siacve notumiatl ties? 2Oon forrasbb 4 1 [RR “) _ 1 vi mt 16.8 s\ ih op eucisiilicael heey eo vibes (TAS ser! tive. ebaolee ow | Deiat i hat | null Lemwwod cou liaglae wiicaw nog iia eke ae n ; i! } we ; % 4 53 = * . She ete =, EE i Fa OL ees = PN POP 2 GiGi pat ewes: alts = % Ss Sar a tee eT eee ae $e Le ee a 10. Iroquois Gravel Bar, East Toronto = MY i i a Pray ¥ v ¥ RD sy oi = 1 ¢ A { i wr { 7 ; t : ae et ’ 1 ay i> far i nT) ‘ ‘ > ey f ap NG 1, ER Ae" i A 1 iY TA ty & 7 ial 2c veel ane ADMIRALTY LAKE 49 -buffalo, and beaver.1. Mammoth ivory and bones are fre- quently found in gravel pits and brickyards west of the bar also. Fragments of wood determined by Penhallow as Larix americana and Picea (probably nigra) have been found 30 feet below the surface in Hamilton, suggesting a cooler climate than the present.2 At Toronto many horns of caribou and a few bones of elephants including a mammoth tooth have been found in gravel bars; and also a number of shellfish—Cam peloma decisum, Pleurocera, Sphaerium, and fragments of Unios, all still living in Lake Ontario. The mammals were animals that could endure a cold climate. At present the caribou ceases about 150 miles north of Toronto, and the mammoth had a heavy coat of hair. It is probable that the climate was somewhat colder than now, though not Arctic. As in the lakes previously described, the barrier of ice forming the shore toward the northeast did not prevent the waters from being inhabited. It may be that certain crumpled Iroquois sands near Toronto have been pushed by floe ice, but there is no evidence that icebergs floated on the lake, such as might be expected under glacial conditions. Admiralty Lake As the ice withdrew from Covey hill north of the Adiron- dacks, lower outlets than that of the Mohawk valley were opened, and Lake Iroquois came to anend.* The successive outlets were of relatively brief duration and only feeble records have been left in the way of beaches. One of the stages has been named Lake Frontenac, since it was held up by an ice barrier resting on the Frontenac axis of Pre- cambrian rocks, but no definite shore line has been identified with it.4 Formerly it was supposed that as soon as the ice with- drew from the Thousand Islands the Ontario basin was '1Geol. Can., 1863, p. 914. : *Bull. Geol. Soc. Am., Vol. 15, 1904, p. 366. *Fairchild, N.Y. State Mus. Bulls. 209 and 210. “Taylor, U.S. Geol. Sur. Mon., LIII, p. 445. t} i ye meat) [9C 1 if e q . a shee hic i iio) ool : rai te ia A yntemniue wut d ) PP 7 >) rah fick 7 i inde feut ' “ui RANG oe i+ act fe mand tt pagiteicia . 2 : ’ , - OTN sli vente B iG leary he hiumiint 08 iow 0 908 a 4 wae i) ae’ hen hs Why Ld sae a, ae : seni 3 | lis BT 4 yinne atu i £ “i eh E ¥ ‘* nie crtey't c] bes } eee he irom 4 y : AWVGIA bee ak ‘ Tw ing # oe att ote il 7 ios: aT ¥ e» ’ i% | ' ! A ue al +: Sad ll 7 ’ ‘ ~~ * J ty of JD - ie tad Wed yom “1 4 4% rat “7 part otnowl oe ited Gaeta . i ostg boo mae = ra thd «lovey reo? mon —_ het Ss eelt rio toms pa a C Steg} oT apa ite ylove ay “occu Jy ee a oy Th ivy f eb 1 : ides). melt “nek A Map Of Aomirarty Lake Ano Low Marine STAGE “A Seole Lcd 11. Map of Admiralty Lake ADMIRALTY LAKE 51 flooded by the sea. The continuation of the Algonquin river channel down the Trent Valley, as mentioned on a former page, makes this view untenable. The base to which the valley was cut must have been somewhat below the level of Lake Ontario, and soundings in the Bay of Quinte suggest a possible delta near the mouth of the river, which may have been formed by Algonquin river or perhaps by the present Trent river. Some soundings lower down the Bay of Quinte reach 100 feet, perhaps in an ancient deep channel now partially filled; but there is no certainty that Algonquin river extended much below the present water level. At that time the sea must have stood much lower than now, probably because a large umount of ice still remained unthawed upon the continents. It has been estimated by Drygalski' and Penck? that at the maximum of glaciation the amount of water withdrawn from the sea to form ice caps would lower it 150 metres. This is perhaps an over- estimate, and Daly’s later determination of from 50 to 60 metres scems more probable.? To what extent the world’s ice caps had shrunk when Lake Iroquois ceased to exist one can only guess. If we assume that only one half of the ice still remained, the ocean would still be 80 or 100 feet lower than now on Daly’s assumption and 250 feet according to the German estimates. The last beach in the Ontario region was probably formed much later, after most of the ice had melted, re- storing nearly the full volume of water to the sea. At Trenton, where the Algonquin channel reaches the Bay of Quinté (level of Lake Ontario), there are several beaches, the best marked, at 78 feet above the present lake or 324 feet above the sea, probably representing the highest marine shore. If the sea was 80 or 100 feet lower when the channel Was excavated, the basin must have been below the present 1Zeit. Geo. Erdkunde, Berlin, Vol. 22, p. 274. Morphologie der Erdoberflaeche, p. 660. 3Proc. Am. Acad. Sc., Vol. 51, p. 173. i roe | i + e Me me P< om wugik ’ Liyort, & rin t 7 i WY. i i ve * Pa e weer) ; se y , ‘7 * er he itis i i f ‘ iri : vi 7s i - ce ee iets ba | | Fi a * \ ot ot ny saa hie ih € a. | 4 i ; \d , ren 1" i } & ttt ‘ mn ‘tse, Bwon . a ® é, ’ yee 7 ae 8 & - LY oid pene eM = eek; yet) Ve a wy f } a i deal ¢ i i j — J 4 ; - 52 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO lake level and therefore must have contained an independent lake. Two other factors which probably modified the con- ditions of the time should be referred to. The great mass of ice still remaining to the north must have exerted some attraction on the water, but probably would not raise its level more than fifteen feet according to an estimate by Woodward,! so that the depression of sea level might be counteracted to this extent. On the other hand, the outlet at the Thousand Islands was probably blocked by drift materials left by the ice sheet Which had just vanished, presenting a barrier between the lake and the sea. At first this probably held up the water of the Ontario basin at least 25 feet, as shown by drift deposits at Gananoque; but later so easily attacked a barrier must have been removed by the strong river flowing east. The amount of tilting of the old water levels in the eastern part of the Ontario basin is two feet per mile, as will be shown later; and Trenton is 25 miles from the parallel of Kingston, where the outlet was, along ihe direction gf tilt. The water level at Trenton must therefore have been 50 fect below the present when the twenty or thirty feet of drift filling the channels among the islands were removed, and this must have been quickly accomplished by a river as large as the St. Lawrence. The body of fresh water occupying the Ontario basin when the Algonquin river reached the lowest point in its valley may be called the Admiralty lake from the beautiful Admiralty group of islands, between which its waters flowed seaward. Though its outlet must have been much the same as that of the present lake, the old shore of Admiralty lake is now below water, because of the northeastward uplift of the region, and its western end must have been about twenty- five miles east of Hamilton where the present lake has a depth of 220 feet. The St. Lawrence river was probably less than fifty miles long at this time. ee U.S. Geol. Sur., Bull. 48, 1SS88. Sica 4 “3 3 u i 7 GS ; a rion ky “4 iG ong ul T ieih’ olay: van’ é ; - . ’ i * te ' Bache n yd gehcrid beyaki wy , ae | - jt ‘at yay | : al jit wie Orns a , a7 vA a? iid hi Se. i? iY 9. trinket er ny v. ) Libre “pb i ic) cae 4 Ve att a a ‘ i yrs wot: eon ony 7 Lanpiak. ited. tO S425 ae) i} elt oa j i i { } Ps ver ‘ine 1 al , ¢ if) F fy ie ah by " we | i iri Kou § i 1 ek see alpen el overt lasre : i iid apa. wort! poe a a % SAO ri TRA fin.) ie tte at) OO ee ree: oa mp aye plod. Ay vil quence i EU la 3a Ath : + ee af? .oocpe ginal 1SViems Hie er ee Oe lsat B ‘ ile (he) pe at iat , dinates 1G Fe ast fe : > io He + = Gi sage svne aN gadtt Reb Iogear ou O€ ’ if Seyin ee Ol bo (ort Saare 4 Aude oy whe wl} ie ie AY yeh cep sag wih ve apt : Sirs » belo nn BM ty ye y a y third ast petit j ae pert wah tie ree isl + avert Wu hina aj bede Yh typi tls bh ot} teh: ound Dot 20) 10. 09mee py wen hae ae inane “iT re | ea « “i Vt'ecn Map Of GREATEST MARINE INVASION s 190 Mies Scole 1 12. Map of Maximum Marine Invasion Wn é ‘ a A pet mbes « nr rt? —— er " i. ‘ = — a tee ST : i * a t 12 40m 1 oe a La AoA “se maAM tea YAS fy | Se A a SO eae a (vet ak ihe ays panty ALY 7 GIRL EAR ee f ie} Wed ACY ib ; ai é ; ‘ h pee | ; . 17 if 3 ‘ ; : ; » - ' ——— — i mers, weilb mettle rin ls. ELL ey | % | ar \ eNie | f rk oa ‘ a —_, =) f nal = ¥ = a ._ ev Ry Aah! hi =v uv at) & 54 GLACIAL AND Post-GLACcIAL LAKES IN ONTARIO Marine Stages in Eastern Ontario As the climate grew milder the water removed from the ocean to form glaciers was returned to it and the level rose to correspond. All the lower part of eastern Ontario was flooded by what has been called the Champlain sea, really an extension of the Gulf of St. Lawrence. This is proved by many beds of clay, sand, and gravel charged with shells of Macoma, Saxicava, Mytilus, and other molluscs, though no continuous beach has yet been mapped across Ontario. The upper limit of the marine invasion has been determined in some places and it is certain that it rises toward the north.! East of Brockville there is a beach with shells of AMacoma rising 331 fect above present sea level, and marine shells occur also in the town itself; but west of Brockville none have been found. Fairchild has recorded higher marine terraces near Clayton in the state of New York, and has given the name Gilbert gulf to the extension of the marine water level into the Ontario basin.? Though no marine beaches have been traced past Brock- ville, it is almost certain that shore forms in the Bay of Quinté region represent the westward extension of these levels. In the state of New York such a shore occurs from point to point as far west as Oswego, where it is lost beneath Lake Ontario. On the north side of the lake a good beach hand-levelled at Waupoos, near the east end of the county of Ponce Edward, is at 340 feet. The best. beach near Trenton, as already mentioned, stands at 324 feet, and other beaches occur at Brighton, Colborne, Cobourg, and Port Granby five miles east of Newcastle. At the last point there is a terrace 274 feet above the sea, or 28 above Lake Ontario. Leyond Port Granby it has not been found, tMarine and Freshwater Beaches of Ont., Bull. Geol. Soc. Am., Vol. 12, 1901, pp. 129-146; also Late Pleistocene Oscillations of Sea Level, Johnston, Geol. Sur. Can., Mus. Bull. 24; and Champlain Sea in L. Ont. Basin, Mather, Jour. Geol., Vol. XXV, 1917, pp. 542-554. *Bull. Geol. Soc. Am., Vol. 17, 1907, p- 112; also N.Y. State Bull., Nos. 209-10, 1919. MARINE STAGES IN THE ONTARIO BASIN 55 probably because it has been cut away by the waves of the present lake.! From Waupoos to Port Granby there is an average decline of two feet per mile in the direction of tilt, and if this continued toward the southwest the old shore would sink beneath Lake Ontario a little west of Whitby. At the same rate of decline the ancient gulf would end about ten miles east of Hamiiton. As no fossils of any kind have been found on the shores of Gilbert gulf, while sea shells are widely distributed and often very numerous eastward from Brockville, one may conclude that its waters were kept fresh by the inflow of Niagasu river. The changes of water level succeeding Admiralty lake have been attributed, thus far, to the. continuous melting of the ice sheets restoring water to the sea. In reality, as Mather? and Johnston* have suggested, there was a com- plication due to the northward rise of the land brought about by relief from the load of ice. _As the sea was filling up by the melting of the ice there was also a tendency for the whole region to rise higher above sea level. Probably the rise of the land lagged considerably behind the removal of the load; so that the sea had time to leave its mark in shore deposits and high level beaches before the general elevation caught up with it. Ultimately, the Thousand Islands region rose above the level of the sea, and Gilbert gulf was replaced by Lake Ontario, the change being probably a gradual one. The northeastward rise continued till the lake stood 246 feet above the sea; but within historic times there appears to have been no further elevation. The outlet rose faster than the southwest end of the lake, so that all the rivers toward the west have their lower ends drowned, as illustrated by the depth of Niagara river near its mouth and as may be seen in the meanders occupied 1Bur. Mines, Ont., Vol. 13, 1904, p. 238, etc. 2Jour. Geol., Vol. XXV, pp. 542-554. 3G.S.C., Bull. 24. pt | , ; i : > «¢ : ‘ ’ é. su) achotem nisaott . fy aan ani ‘nmin ee haa rr | etia ref Ite Hh Silt a vibe ; ; bk to bao’ peli tabal “ - Vel tll ii ‘ig des pay a a i) eA, 1 eS : att i & i OF 4 { e ui be : hy, be un Aut’ bra rat ee 56 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO by dead water on the last two miles of the Humber. Pro- bably, too, the deep water behind Toronto island and Bur- lington beach near Hamilton is due to the continued building up of beach materials as the water was backed up toward the west. There is a great similarity between the present Burlington bar cutting off Hamilton bay and the splendid bar of Lake Iroquois rising 116 feet above the present water level and cutting off a vanished Dundas bay of the ancient lake. Both have been caused by differential elevation that continued for thousands of years, Life and Climrte of the Marine Sta ge Though the fresh water deposits of Gilbert gulf afford no evidence of life, the brackish and salt water beds farther east and north are very rich both in individuals and species. The marine forms include pelecypods, gastropods, sea acorns, starfish and sponges, the caplin and two other kinds of small fish, and seals, dolphins and whales. Concretions in the marine clay at Green’s creek near Ottawa have supplied a few feathers of unknown birds and some bones of a duck, as well as a chipmunk and several beetles, the latter extinct according to Dr. Scudder. Almost all of the aquatic animals discovered in the marine beds still survive in the Gulf of St. Lawrence, and Suggest a cooler climate than that of eastern Ontario at present. This may be accounted for by supposing that the entrances to the enlarged Gulf of St. Lawrence were broad- ened and deepened to the north and south of Newfoundland, allowing easier access for ice floes and bergs than at present. So large a body of cold water as the Champlain sea probably had a chilling effect upon the climate of its shores. A number of plants have been obtained, including sea weeds, marsh plants and several trees, such as poplars, yellow birch and sugar maple. All of the trees stil| live in the Ottawa valley, where most of the fossil species have been collected. Sir William Dawson believed that the plants indicate a somewhat cooler climate than that of Ottawa at present, but milder than that of Labrador. , } ' 1 PoP edie ue eM] t t he at “1 Cyr : 7 rma? f si 4 yey. atl gts pelt ey ih ern Yue seer iat heaven att ti ey eionspels oe i ey wae hs. — rw é tie. gma o1aey syh V°-80) sey BL; i] * 7 a) 1 Vie “y) yp Aware Oey s SRY oy , 404 ihe aa ” eck: sagen iy dan on ath nga ahd: i hie ahi waky 4! “it a) rast! iti 1 Vinh wr paren ay iw Say tal win beara € t meredith Vavlitie J8908 6 ai arf no trae ay r ye choca ieee | ioe iegie/a. 4 Sk oe Lh Ss ay a reals SAME iegiice Esa ahold: Suinae CLR alls Ba one 7, ee "1 : “gitod aves 1910 ona ae eltn ae p ¥ ty) Alege “onan a ‘ so} 1h quyregen haart Heth i 3 geld Je ye! as be he 2 an ; ren ee Eye a it paths ms rae ‘A iit ryt hue oltt ) . cobb Harpe Ses : fe bd iT in HAG | ald nll iavyilex isaizieal i ‘neti » git neartl rie ie bz eer cael lo if Slr dad at ity ry} on ‘ adiye aaytite ‘ vd dae cone : le a bing eibin tal pg hin ieee in aie whe ‘bens . { . ten Bs a sala and also on Soweska river, a branch of the Missanabie river, 128 miles southwest of James bay.’ None have been found, however, directly south of the bay; and it is probable that the ice withdrew from the southwest side of Hudson bay sooner than from the James bay region, allowing a narrow tongue of sea to come 1G.S.C., 1863, pp. 916, etc. *Can. Ice Age, 1893, pp. 18, etc. ‘Contributions to Pal. of Post-pliocene, Ottawa Naturalist, Vol. XI, pp. 22-26. 4G.S.C., Mem., 101, pp. 16-34. *Robert Bell, G.S.C., Vol. 1871-2, p. 112. 8J. Mackintosh Bell, Bur. Mines, Ont., 13th Rep., Part 1, p. 164. auf titiy (TOI ed: If oe a S iat in! az BY S % take ‘ Wad rave) carveal vi eae aca, eb lon OF 0! WANs WADERS at rf ? mn i, A Ve : " ad ’ - \ ok ee im ih ye WT] Ue te K a » % ol he os dl) spurt: Pickton lek a ae a) veriey TT S130 i? ety Ton a ive stermibes oF m2 “it pel a a 7h) wes 30 Virou oA) aniog sew Cy in BA: 3 lea an meant, aa © Aatey taksalsy ot3 To | Ment baniene levstens wl ie galindeen Wee cae anil: aides PE he ee ae a a yi ie Re ro “onag ¢ Jabs ‘ Me on, = = 4 et be ‘i Dawe “eee wv hn heir) OF ryt ary vabvocrokhisad 4 aie pals, pe i it Z gob IH % Ve LF Bis) m OY Sep ‘Aa te tt a 35h ts eth) Se ee COREY ‘Osr * rime | wa nen AE i) anoles cb Laren at sid daliet: oF) - ante tyr lf over Hit. GL. Ta See ave ths | sere NS ea): Tere Pits9 i] wh ted Sttledé 1) ahine ia noah 3 Psy be “ P ae P om eer Ad dd GailescieA - oy rie - Ly, 2 exc’) 6 Pies i 7) 5S GLACIAL AND POST-GLACIAL LAKES IN ONTARIO in between the watershed and the ice front. No gravel beaches nor shore cliffs have been described in the region and wave action was apparently only slight. It is generally held that the sea rose to 500 fect on the southwest side of Hudson bay; while Low gives its elevation at one place on the east side as 710 feet, and mentions the interesting fact that seals live in Seal lake nearly SOO feet above the sea and nearly 100 miles inland. They probably reached the lake when the sca was at its highest level. It is possible that the sea encroached even farther on the land, since the highest marine beaches usually do not enclose shells. Until careful study has been given to the Pleistocene features of the Hudson bay slope the exact southern limit of marine invasion will not be known. If we assume that it reached the 500 {eet level, White’s Relief Map of the Dominion shows its most southerly point to be on Abitibi river about 130 miles northwest of the head of Lake Timis- kaming, Turning now to sea levels south of the Hudson bay water- shed, we find that here also there are no definite shore lines, though there is reason to believe that marine waters reached much above the highest known deposits containing sea shells. De Geer and Johnston report a beach at Kingsmere near Ottawa at 690 feet, as mentioned before, and not far off Johnston has estimated the northward rise of one of the beaches as amounting to three feet per mile. This rate of deformation is more rapid than the two feet per mile found farther southwest near Lake Ontario, but is probably correct, since all the old beaches of the region rise more rapidly toward the northeast or north. We may apply this rate of rise to the fiord-lile bay reaching to the head of Lake Timiskaming. The distance north of the parallel of Kingsmere is 133 miles, which implies a rise of 399 feet. If this is added to the height of the beach at Kingsmere, 690 fect, it gives a sea level of 1,089 feet at _— 'G.S.C., Vol. IX, 1896, p. 13 L and p. 42 L, . : Leh Ww ‘aly ify t sistay ha ae ‘agtt UMA JA 4 wAwiatira «yet Til tis orn heath cp test: (VISE ee dois A pest Wa 7! "nt nod tylatt unineae | + 2 were) al jy 20g 4 T\y54)] la aye a (At vegies ta puel) UFa ia. 9 abe of ing) : KA ghwed aitnl | Agios snng 4 tervey spat) Abrinabste eotscre SN (risum brn. ae ; ti vit pie By we) mh ata 291 Hs bi 4 J } ailt FO wAP ut awe Daitsnencn? Ae alt: Curt rit ig ei, a/)] nou reat 12 ‘ a : ‘a . ey | t (ld [ ale OG) cervey, 2 on Hite vide iru se, OED) Ay a ota ; au. 10 ree jon Te H qj ( ij ure +e: » ah praofivoe Dah atl at tk: ; ‘Ete ta) toy sega hai: 1% ma a pee \ 4) + vod ooh Hy i renee N j , i oie pac tala mn vy 71 i pfey i wy i” me 7 s - ch “4 - wy aa. 12 ig i 7% i “wt BAe ¥ pom ete ened oe Lin eh eh i aS \ i ¢ i ’ A ! af ya Yyuilt os Qe sere 4 oath) Diget ai ehisal teeter? mint sed, x ‘| Ay" i 4 + acheter onthe a! ia) a ty ni A “At Titre es a) tek Wt 4 it vide eure Bet ar oe nite At? ob nea iekte - eh BOT Yo favo io ceo a ‘z: +89 38 et ' : -FAUNAL RELATIONS IN THE LAKES OF ONTARIO 59 the most northerly point reached by an arm of the Champlain sea. Now the watershed between Lake Timiskaming and Lake Abitibi is 935 or 940 feet above sea level, and the pass used by the T. and N.O. railway, some distance to the west, is at 1,0-14 feet. From these figures it is evident that if no obstruction intervened, the broad strait suggested by Dawson must have been a reality. However, the discrepancy between the highest probable sea levels on the two sides of the watershed is so great that the northern invasion of the sea must have occurred much later than the southern one, in fact after the whole region had risen some hundreds of feet because of the relicf from the load of ice. The southern side was the first to be unloaded and had already risen greatly before the northern side began to be set free. It may be considered certain, then, that the Hudson bay watershed was never submerged and that no strait turned Quebec and Labrador into an island. The isthmus connect- ing northern Quebec and northern Ontario was 130 miles wide at its narrowest point. Faunal Relations in the Lakes of Ontario There seems to have been no break in the continuity of the fresh water fauna between Lake Iroquois and Lake Ontario, in spite of the marine stage which has just been mentioned. Lake Iroquois inherited its inhabitants, through some short lived intervening stages, from Lake Warren, which had its outlet into the Mississippi. Ultimately, then, Lake Ontario has obtained its fauna from the Mississippi waters, which escaped glaciation because of their position south of the margin of the ice sheets during their different advances. Apparently some of the Mississippi forms, perhaps for climatic reasons, failed to reach Lake Ontario, or else have found conditions unfavourable in later times, since a number of the Mississippi shellfish which lived in the Niagara " J Fi : af ‘ vedboady aca] gett gears 13 LG plewiat 2 es va i An hye: {clog Zhen 7 Jip 7T suk neti) ae ) ree hag bce ly hd Cael AM ener *pediagt riaenteih fifo She Oe : ; boty a } dP meld ae Le 9; i 1 writs: Ong se rtore i iF Mit 30) = “ot no a yobs iin gat wale lo tro evel ies ‘Wh coh ; a eh? QAR al mod ‘ a on Te lund le Lote svat Be ott, tai ¥) , y f {s . hing heb oft i pc of GF 2 ' sod) cade pete ‘ aie tnd. bow vores. fer ; linikhe ae ir i. nO sain hie : digaise io) Sas “it o) doo aoe ent -fe3 poate mush 1 Yous inca aucia ote fH “38 it ainda tneigd ott jvythaiak aug 1s fi 4.1. tv) yep ane aA t in (‘eared rey cieniail oft oN i refers ied eM eer! nail 2h hoes sa iat Tae , a> mind outalsane ay “itd: port aruda oor oat Yo ie, 2g ~ aaron! lb lah if wih dos cae ‘yi ey ay onnia® oHe : Marte» | ba ie: Lala ‘ oie - pom t Taia) 1 rae ibeck : aE wth ni bovildlan’ ‘ » as A o - 2 "i " 7 See = me 60 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO river when the Falls was at the Whirlpool are not known from Lake Ontario. From the complicated history of the Great Lakes and their predecessors which has been outlined in this paper it will be seen that all of our Ontario Lakes, small and large, even to the north of the Hudson Bay watershed, have in one way or another had connections, and that ultimately the fluvial and lacustral faunas inhabiting them haye come from the Mississippi waters, sometimes, however, in very roundabout ways. The Mississippi and its tributaries supplied a harbour of refuge when the northern part of the continent was ice-covered and lifeless, and colonized the lakes and rivers which arose after the melting of the ice sheets. If the icc had reached the Gulf of Mexico, so that no place of refuge remained for fresh water life, our lake and river fauna must have been meagre indeed. Anadromous forms coming from the sea, like the salmon which once spawned in the streams tributary to Lake Ontario, might in time have become “land locked” varieties or species, but I am not aware that this has been the case. Probably the incoming of the Mississippi fauna filled all the positions available. The connection between the waters of the Great Lakes and Rainy Lake, Lake of the Woods and the Manitoba Lakes appears to be limited to the drainage of Lake Agassiz and of Lake Warren, etc., into the Mississippi. Here the mode of communication seems very devious, but sufficient in the lapse of hundreds or thousands of years to account for the identical or closely related species found in these distinct drainage systems. There are a few cases of lakes on the northern watershed having two outlets, one to the Great Lakes, the other to James bay; but I am not aware of any two-outlet lakes on the height of land between the Great Lakes and Scine river to connect up the St. Lawrence drainage system with that of Nelson river, so that the connections with the Missis- sippi must be considered to account for the species common bh ia | ~ — . i ; ’ v F ; i. . { ae «.2 be ¥ ij ole tTHA Jamaste - nyt ed uta oa : a ue ny i ae = Vids" et att t ae 7 ee: 1 wingar in ‘ y », “ yes pT j ee ‘ures wht in ry " ¥. year ub a . i“ am x eye yy be \ ms aris Ly Rin sich | Ta crcl) thee Oe by, anit wv 1. | i , ri — Lava e tf Te. ni I . Me i pA: me i 7 ¢ - Pais rape BBS oa ) wi i 7 ‘7 é V8 pct } ie >| nl “nt ) | ti}2 yy oe repty vical Tice! Hankesieallt ae wee ’ 7% z } seal I aeS 1 ae: ‘ \ ‘ we » aan iis ohare POP, 1S eur ~ : ‘ i eyrty SSS sont a roe ie one juzdl wha Vik ine det! 29) Daa bh! Semele ofp o ewes oes bude teil ack) Walt on ‘a 4 ie ay vidird esate, : i pl &, 4G bast : ahr i a wate |) rae es , i; the Syatt: nae Se ion Bie ‘ : r —_ ui ve omen Aa We = = 2 tf + + . jy = ra: atta 4 7. 7 2 Peieeery ot boxing RY i he fa We, dg of ‘i | / / athe hen) area View Gee ner ye al nn dy PLE “a “ye aber / if ! at} , a f | : bad tiny i gl breen vy rae vrs igi io ott lake uae) oe aa bing 4 AY Le wy) eile Be? ripe Jott tent yd} cams) ot nan «) Late Pee, iiie # . jovi eee ssyntvieals oon ith 2 ‘ede - est A: RTGS a TOO ates vl \oesates norragn ori Yoh spy ones “eh —- ; ls — - TEMPERATURES IN DEEP LAKES 61 to the two drainage systems, unless some mode of transport over land or through the air can be assumed. Temperatures in Deep Lakes A comparison of the fauna of the glacial lakes, so far as known, with that of our present lakes which have no perma- nent ice on their shores, shows little difference between them. Practically all of the species still live in our waters. This comes as a surprise when one recalls that the early lakes had a vast sheet of ice to the north forming their shores for many miles. Why should these subarctic waters be inhabited by the same species as live in cur temperate climate Jakes? The presence of caribou and fur clothed elephants and of black spruce and tamarack on the shores of Lake Iroquois suggests a colder land climate than the present at Hamilton and Toronto, as one might expect when one-third of the continent was still covered with ice; but the aquatic life shows no such difference. To explain this requires a consideration of the peculiar physical properties of water. Unlike other liquids water does not contract uniformly ,as the temperature is lowered to the point of consolidation. After contracting steadily to 39.2 degrees (Fahr.) water begins to expand and this con- tinues for seven degrees to the freezing point, when it changes to the solid state and suddenly expands about one eighth of its volume. Other liquids contract all the way and con- geal to a solid which is heavier than the liquid. For this reason cold water floats on warmer water and ice floats on cold water. If water followed the usual law when cooled below 39°.2 (4° Cent.) it would sink to the bottom instead of remaining at the surface, and at length the whole body of water would reach the freezing point of 32° (0° Cent.), when ice would be formed, and being heavier than water, would accumulate at the bottom. During a long winter the whole lake would be transformed into a solid block of ice, in which no life could survive. Some of the surface ice would thaw in summer, but in deep lakes the lower parts would be perpetually frozen. ry : mise | oe icy wt on A aN 4 7 " & P| F iy mii am wa ili win ade sh a TT, mr “1° 23% smog Va gd ted Ge une! | doula ean ‘olen unm on wein daby alicl ins Here ¢/ v/s pire oat ir! 1 altyik; ele ey pwolitw Wid: dr? Hide ests a A el vlvasorir yl) chess oy thw aaeee > | Oval) evitin Qe) of! | prio! HTT a a Koy i; # be T ff ne ed vat 5 rei) if Ax ectiane be al oe "4 . Pere ' inet 19,5 sine lla sw bie oo As 4 stl ir 1 dco asvieie lt poe A OH » iid 1 a a3 ih pep eer A TOLD it nti IOs L CHOSE Shel DT Mid aly sud ae y J 5 aoa bs | potda edie’ Aer ox) 22) ea ee niytd nea Worl netiabiloeiog hos hegqae oO) Sia eae & ui . papier deloy gren aon 5 ata $00 Leng? ne Reet vow 03 Tia Joong abi gph rit 7 sora i sip } ] Oy “>; bf8 19704 We yf Tt ott cow 2st Lec: ort Loews i raprvet di oy aie Bie : | slept iki Penal Jn nee 9 1. > bite noice Teetie wish ot ol vivre hnk wate Snot a get! CE * 5 xhtarled nies. tin ; be note ‘ie i 49] syntinel@é? Wh otroct ie 23Ing a pat Bane sa Steen (GEAGIAL AND POST-GLACIAL LAKES IX ONTARIO The anomalous physical property of water just mentioned permits the lake to cool to the point of greatest density, after which further cooling causes the water to grow lighter so that it remains at the surface and js finally changed into ice, which floats and forms a non-conducting protection to the water beneath. The importance of this for the inhabi- tants of lakes is manifests With winters as long and cold as those of our region the whole volume of water in a lake must reach the temperature of greatest density, and in deep lakes the lower portions must remain at this temperature all the year round. Temperatures at Different Depths in the Lakes Through the kindness of Dr. W. A. Clemens the following ‘data have been collected on this subject :! Lake Ontario—W. A. Clemens, Oct. 3, 1922. Depth Temperature Surface 67.6 Fahr. 10 fathoms 60:4. * 20 fathoms 47:0...“ 30 fathoms — 406..." 50 fathoms oh fe 62 fathoms (Bottom) 39.2 « These records were obtained at a point about half way be- tween Toronto and the mouth of the Niagara River. Lake Ene—W. A. Clemens, off Merlin, Ont., Aug. 3, 1920 (bottom temperature), 5 and 6.6 fathoms 52°.5. Lake St. Clair—Prof. J. E. Reighard—A Biological Examina- tion of Lake St. Clair, Sept., 1893. 64°.4.te 69°.8 (little difference between bottom and top—not more thian’1°). Georsian Bay—A. T. Drummond, Can. Rec. Sey -Vol, IV. _ pp. 77-85, and Vol. V, pp. 13-19: Aug. 20, 1886, Bottom 31 fathoms 397.5 Bottom 47 fathoms 38°.25 ‘As most of the data are in the form of Fahrenheit degrees, with depths in fathonis, it has been decided to recast the others to make them comparable. KHizseAeea? YaesaA ? LAT Mir wp | era atonal we [_ - ye SPT sige) ary ‘ MM fen = ber ny tenia nual Ni » “vie vb Anse if wit tw 4 A> vif ae Ty } ek Ware 5% y joy } Aaya whi(ionr) yhoo dni —) td ain. £10) ae MSTibe ONE IS anil: z ik wand pelerey 5 W 5 aerion ret el i jo nme Vee j A : ag wit raced Gmriy> sh 3.F FT) “On i iy ‘ t fii befvil aye, iL i sie yhigreites ai ni Tate 16 OF +) he ont wolovrage . bo) prtlees ay csi mserygs oil wed? ' tT itrahoil op trabs: = ‘ ya? pr te j 7 ¥ i io ] t) u A a4 atric a 2s Retell aera bod 4 rena fh apentinl® 1 hanielals ‘ '> A yel 7 7 '. i ta Re } oy oh Rite Kiar ie Get : yordy ict futted nome Jou goa q i Pe pie AL 3 ba ‘ne V1.2 =’ sqet ? "OR eunitc 6} if motok iat 3¢.°RS afmoijal 7 mosIORL ey : Aeytty dre cegagely fete fat va tesla nid steed masahs aitein ot ay tar it = x a = - = — 7 _ TEMPERATURES AT DIFFERENT DEPTHS IN THE LAKES 63 Georgian Bay (Continued)--Pottom 42 fathoms 37°.75 July 10, ISS9, Bottom 70 fathoms 38°.75 Sept. S, 1SS9, Bottom 63 fathoms 39° Surface varied from 59°.75 to 6S° Fahr. July 27; 1888S, Surface GO*.2 10: fathoms 45°.7 20 fathoms 41°.4 35 fathoms 41° 66 fathoms 39°.5 In Parry Sound Harbour, 1890: May 2, Surface D022 Bottom, #2 fathoms 35°.7 Aug. 23, Surface 61.7 Bottom, 48 fathoms 39°.2 et. in, Surface SE BS Bottom, 57 fathoms 39° Lake Huron—Dr. Walter Koelz,! Department of Zoology, Univ. of Mich.: Sept. 12, 1917, Off Alpena, 65 fathoms Bottom 39°.2 sre Off Alpena, 15 fathoms Bottom 57°.2 ee oe Off Alpena, 60 fathoms Bottom 39°.2 neath corks Cheboygan, 35 fathoms Bottom 39°.2 wees JO19, Alpena Surface 60°.1 aAe tas Aipena 35 fathoms Bottom 41°.7 Lake Michigan—Prof. H. B. Ward-—Biol. Exam. in Traverse Bay Region—gives 98 records of surface and bottom temperatures taken during August, 1894: Highest Surface Temp., Aug. 15 70> Bottom Temp., Aug. 15, 6.2 fathoms 68° Lowest Bottom Temp., Aug. 16, 61.3 fathoms 39°.5 Lowest Bottom Temp., Aug. 18, 72.5 fathoms 39°.5 Surface Temp. on both dates 64°.9 Lake Superior—Dr. Walter Noelz: Aug. 24, 1921, Off Ontonagon Surface 64°.6 Aug. 24, 1921, Off Ontonagon 60 fathoms 39°.2 Aug.25, 1921, Off Ontonagon 34 fathoms 41° ~ Kind acknowledgment is hereby made to Dr. Koelz for permission to publish these records. f « ; : i i «tated ier vy) earth ss apeeetl Th Aen L Aa Fat A es <*) r 4 Tae meac® | Ni rat wor hath-gtlenel ane j sae een es : Lech wien lt OrA4 Fe s > ; ‘th he ey ivi c on Ai ray | R 6G. naire. ere GY " = etree ie Wi a Bor). steal eroe ii F ohio ga ina vorls De (ni) Al cwor ott St uiiorne) i Janae ivi rues he - ¥ rh J oa? pee ’ e ¢ ' a, | . r" ie "5 omol Std eee eee , Lidmaniar aaa aah ; ! cy ‘ oft orton at Ad ae 8 4 SenAM tl at rah B 1 ; rie eile rArs HARE y. worst ao ra ; Si: = , oY ‘§ io wubtél fn aver at Hox \ i Oe , ye enrol heA (orn lo eh werent BR-g ‘o~ Serrotnh, yoieboe "DY: él hy He | "A ariddicl 20. (OP ae jt oul iveelsh) SIO OE BUA: ‘ +r: a. UE “peed re} 3% Al aah a scons obo oe elab 1"30 4pa@linc LOR emroiish 09 Godel of °' amote) Dt do_iine fisildug ot wodeaborray pol thao rt she r - ‘ eed G4 GLACIAL AND PosT-GLACIAL LAKES IN ONTARIO Drummond in Can. Rec. Sc., Vol. IV, pp. 78, states that Hind found the surface of Lake Su xerlor on 30th July, at ~ a e - y noon, as low as 3%°.5 fifty miles from land. Lake Nipigon—Dr. W. A. Clemens: July 9, 1921 Slugtace. 7 E26 fuly 9; 1921 23.2. fathoms, 39°.5 (At deepest point, 67 fathoms, temp. probably would be 55 ed Aug. 29 Surface 627.8 Aug. 29 45 fathoms 40°.1 (At deepest point, 67 fathoms, temp. probably would be 39°.2) It will be seen that the data recorded are very unequally distributed, and it is probable that the bottom temperatures given by Drummond for Georgian bay and Lake Huron, feaching as low as 37°.75 and 35°.7 Fahr. are an error and should not be lower than the point of greatest density, 39°.2 (4° Cent.), or making correction for the expansion of mercury at a depth of 60 fathoms, 38°.5 Fahr. From the tables just given, it will be seen that the temperatures of deep water in the Great Lakes are about 39°.2, as might be expected in a region having cold winters. With reference to Lake Superior it may be mentioned that the present writer found a temperature of 40° at the surface near the north shore in July, 1899, and that Professor Ramsay Wright states that the temperature has been observed to be 39° Fahr. (=4° Cent.).1 Shallow lakes, such as Erie and St. Clair, no doubt get above this temperature in summer, but we must think of our deep water fish as living most of their lives in water constantly at 39°.2 (4° Cent.), though they may come up to shallower and warmer levels to spawn. Toward the close of the ice age, when the glacial lakes came into existence, it is probable that their waters also were at the temperature of greatest density, and that the salmon trout and whitefish would find conditions as con- —— ‘Rep. Ont. Game and Fish Commission, 1892, p. 425. iix¢ OQ 41 eal . eshte a sais Af es :¢ 2 a wine, Ost » A ie 2 sb) ee 4 ber, (eae sole tat i 1 eel ‘ . } ‘" rai he ' ai haa recite Vd Ghee rit ij } i ti he a1 wii 1 ty? a ’ , A! egravitis Ste it bs aid - _ " é ae ae ’ = nob shinee idedoky Ain? «ttenliel Waa 4 7) Le Pie we? 1 "Ok. eet i (wal Lin 13 ernie! 7 Ri “pntetr ert Sp heaty Ya | ix seify ohio ere ee Adi ae/ rat tian i] a4 i tS Ott awe hoe 49 sonnet ‘asl 8 Nee ire 11 inbat wy (foo: . tel hel OO ae 2 . ; ‘ 8 iy reue J zac {let os Gt a . ows of Agi i ay. ) eee a ol “9 (eS: Ss Sat | “1 tone oh oe darth 3 yous phe comme: AF ‘r “herp ted Ogata : ernie. reg Jd Reb a Te ry 304, 3 lnid? sean) 2a ee eT nip ae Bt peal AR 1@ Je on ae ‘ : nites CF rt roslt hb ! poe r é | 7 , a 4 ine iat) Ay iy’ dae 7g oo as. aes ae west - arid sgipt oi inclorgy te T we 7 siti fA sq feagtol) re acre > on exopibeds Omi bia ive -e9 ai = ae S Y : y i - » ACh at Tet a TIME RELATIONSHIPS AS SHOWN BY NIAGARA 65 “genial as in the cold depths of Lake Ontario or Lake Huron or Superior at present. The climate of these deep lakes may not have varied appreciably during the thousands of years since their basins were freed from ice; though the climate of their shores must have changed from subarctic, suitable for the reindeer and the hairy mastodon, to the present temperate conditions. Nevertheless, there are difficulties in accounting for the deep water fauna, since all the basins were filled with ice and the Mississippi south of the glaciated region has no lakes at 39°.2 in which the deep water fish could take refuge. It may be, however, that the Mississippi of glacial times had much colder water, most of it derived directly from the melting ice front, in which they found themselves at home in spite of its shallowness. The Mississippi must have been muddy, or at least milky, during the many thousands of years when it was draining the glaciers. Whether the animals of the clear lakes could accommodate themselves to muddy river water is a point for biologists to settle. The fauna specialized for the deep, clear, cold water of the Great Lakes must have found the conditions in the Mississippi much less favourable than the shallow water species. Time Relationships as Shown by Niagara It is of interest from several points of view to obtain an estimate of the length of time required for the various events outlined in the history of the Great Lakes region. The chronometer usually relied on to measure the time since the Glacial period is the cutting back of the gorge of Niagara from Queenston heights to the present falls, a distance of nearly seven miles. If the length of the gorge is known, and the rate of recession can be estimated, it seems a simple matter to determine the time since Niagara began its work, 1.e., since the ice retreated far enough to allow the river to plunge over the escarpment. The length is known and the rate of retreat of the falls since the first accurate survey, in 66 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO 1842, has been estimated by Spencer at 4.2 feet per annum! and by Taylor at 4.5 feet per annum.? By simple division the result is 8,000 or 9,000 years as a round number; but un- fortunately it cannot be assumed that the recession has been uniform, since it is known that the volume of water passing over the falls has varied greatly. At times the volume may have been greater than at present, since the natural drainage of the area was augmented by rapid melting of the ice sheet; but at other times the waters of the upper lakes were drawn off by lower outlets, through the Trent and the Mattawa channels, leaving only the drainage of the Erie basin, pro- bably 15 per cent. of the existing flow. There were, ther. two periods when the amount of water was sériously cut down so that the attacking power of the falls was greatly reduced. The times of increased and diminished flow, with other variations in conditions of less importance, are more or less completely recorded in the changing width of the ravine below the falls. The features shown in the gorge may be correlated with the known history of the lakes and their drainage channels, and thusatime table may be worked out for the cutting of its wider and narrower parts. This was attempted by Spencer, who studied the problem carc- fully and concluded that the total time consumed was 39,000 years. Taylor has repeated the calculation with more complete knowledge of the history of the lakes and makes a less positive statement of the age of the falls which he thinks lies between 20,000 and 35,000 years.* He divides the work into five stages, during two of which, when Lake Algonquin drained through the Trent valley, and when the Nipissing Great Lakes had their outlet through the Mattawa valley, the recession must have been slow because of the small amount of water passing over the falls. The most certain part of the process is naturally the latest, since present conditions have been operative; and he makes the 'Falls of Niagara, G.S.C., p. 342. *Niagara Folio, U.S. Geol. Sur., No. 190, p. 23, 1913. 3Evolution of the Falls of Niagara. ‘Folio 190, U.S. Geol. Sur., p. 24. o\ ‘ Pio } ghteOD ot eaval ya eee A Ae wh tein yy be wept teint ie wd god a impiewii ta (ithe a, Sos Dae a : } : 4 Tieng) fury _ = G ue , aks AS bebe. py! (miei a) dot i hese rt wyyerte ie af ’ ‘i ee wed 5) Le 44 1 al 11 von oegh ian Atal ite , Th co nan Oe . 7 j iit-1 anf J OUIPMTTOUA (ie Eres i ivy?) 4) vis ' ’ * : ‘ woe voerad ay ty u ; ly bate Yeu t elt ovdd oii Jae ly : 7 choo a wid uri EA ny > ; ooo to Ga rg 4 ods akg jalondae vik voll Py ) bby. ‘tan ere hy me : ra ul rT f *ernnogt. t | nie”, = fe | , al v4 i \ heat ; . i : , F ! i Ds ; i fi nine g ie | . /_ ire ? » Jag * f a Se ae ae i h é wid a : (hi rues oluronal te ny at ior Saag ae) i tay C08 vvwwr nest at A _ ; ira mle, yer Si avi ‘i Pot (yeni Bey cdl ete teuceds ogee Alene ee ded an ‘ti d de «toca aveel Wile) fi the i uF] ie) sd os iy gine Re 4 feist ue USO yey — On) esse i 216k | SVE STKO 60 sai Tres : wed: ike Aith Bt . 30) 64. oe prow: eh : - = 4 é i , Ay —" , e 13. Duteh Church in 1900 vw dete ey pt ange 14. Dutch Church in 1915 TimrE RELATIONS OF EVENTS IN THE ONTARIO BASIN 67 time required for the excavation of the Upper Great Gorge from 3,000 to 3,500 years. In the two time-estimates just given, the amount of water in the Niagara river determines the rate at which the falls has cut its way back from Queenston Teights, and this amount is estimated in accordance with the areas draining into the river. When the waters of the upper lakes flowed over the falls the work went on as rapidly as at present; but when the supply came from the Erie basin only it was very much slower. The excavation of the narrow parts of the gorge required probably four times as long a time as the broad ones. It wiil be noted that the Niagara chronology is drawn from the history of the outlet rivers of the upper lakes and provides a rough means of estimating the length of the different stages of these lakes, but has no reference to the bodies of water occupying successively the Ontario basin. After the water in this basin had fallen low enough to set at work the machinery of the falls the two systems followed separate lines of development, and it is not easy to correlate the events in the upper lake basins with those in the lower one. Time Relations of Events in the Ontario Basin The Ontario basin began to be freed from ice during the existence of Lake Warren, but the falls did not commence its work until the outlet past Rome into the Hudson was opened and probably not until the water fell to the early level of Lake Iroquois, which at its west end may have been as low as the present Lake Ontario, giving a much greater drop than that of Niagara as we know it. It is possible, however, that there were two or three separate falls over as many hard layers in the succession of rocks during the early parts of its existence. If one accepts the chronology of Niagara just given, the whole of the bodies of water mentioned in former pages as occupying the basin at different stages must have run ee : Lens ® 4 aT ie Soecntr its aA Ove tid i=? ~ oe 4) Devil ony } ‘yh i) fy he i ‘ Lae p, is i oe 1 5 a 141,-¢ 109 or 3 \ ou oe “- ‘s “’ a, re | autre DRONE 7 4 , : a we a =f . Hy iy Tahes a ; 7s + ee in y only att ee ae rep a ie | veel oe ach nigltyant wieweot) ov ae cad ee j ae ie a ree, 68 GLACIAL AND PostT-GLACIAL LAKES IN ONTARIO their course in from 20,000 to 35,000 or possibly 39,000 years. How is the time to be divided among them? Two of these bodies of water, Lake Iroquois and Lake Ontario, were much alike in area and have beaches of about the same maturity. The work performed on their shores shows that both must have been long lived; while the intervening stages, including the marine episode, are represented by much less perfectly formed beaches, sug- gesting a shorter time for wave action. It should be added, however, that the shores of Admiralty lake, which came just before the marine invasion, are probably mostly sub- merged under Lake Ontario and so out of reach. Lake Ontario is still at work and it is possibie to measure the results of its wave action. For this purpose the recession of the cliffs of glacial and interglacial deposits at Scarborough Heights east of Toronto is well suited. The first accurate survey of Scarborough was made in 1862 and the distance of the cliffs from seventeen fixed points was remeasured in 1912. Some of these lines came out at ravines which were — Ss ~< = = eee ee Lake, Ontario 246° aa anaes i RECESSION OF SCARBOROUGH CLIFFS 69 being rapidly cut back by streams and were left out of the computation, but the average of thirteen which seemed normal gives a recession of 81 feet in the fifty years.!. This works out to 1.62 feet per annum. Soundings show that shallow water extends for about 13,000 feet from the shore before the more rapid slope of the lake bottom toward greater depths; and it is inferred that the cliffs once reached this point. At the present rate of destruction a recession of 13,000 feet implies about 8,000 years. A computation of the time required to build Toronto island at the present rate of movement of sand along the shore suggests the same age, bit the problem is more com- plicated, and the results muc’: less certain. Spencer strongly. opposed this estimate and held that Lake Ontario has existed for only 3,500 years.2 So far as one can judge from his rather obscure statement, his reason for taking this as the age of the lake is that 3,500 years ago the northeast angle of Lake Huron was tilted, turning more water into the St. Lawrence drainage. By this he means, no doubt, the closing of the North Bay outlet of the Nipissing Great Lakes, inaugurating the present régime for the upper lakes. Taylor estimates the time during which the upper lakes have emptied through Niagara river at 3,000 to 3,500 years, as mentioned before, confirming Spencer’s view which one may agree is correct; but it is not apparent why this event in connection with the upper lakes should fix the age of Lake Ontario. There is good reason to believe that the basin has been greatly tilted since Gilbert gulf came to an end, as shown by the depth of the channel of the Niagara and by the 78 feet of water behind Burlington beach. No definite reason has been mentioned by Spencer why Lake Ontario should not have begun during the lifetime of the Nipissing Great Lakes. They were entirely post-glacial and their outlet through the Mattawa valley must have reached the Ottawa after the marine waters had greatly fallen, since 1Compte-Rendu, 12th Geol. Congress, Toronto, 1913, pp. 435-449. Am. Jour. Sc., Vol. LXIII, No. 257, 1917, p. 360. @=. 7 in ris) 1 iy Ovi ys 4 woETeN Me Had ify, sa sp 0(R cope OMe aeihs 200A ee, eet re - s) Se oi a aa je — a wee = : eae S fa AOR “7 cine er ~~ iss i - - va Pp 736 9 ih ir T,” Sued, Friar iy’ i] i ‘ci “5 \» HS roche eae | eek - eat “4 A rok. rer to ve) ey iiie ato ais) 4 its YT 7 + ' ? | | & AR! ae vet « i a ae Ail ue! 7 ith, ie ; ] ths Wel: ‘ont ish vie hada i re AMS On 24) j “ an “ ; ea ti fvri } , ‘a kos 4 a9 * ie Veh yy SL Wee ayicat wele = bt Dock one i *y oT | ‘Sigrte vf reste ph ior nade i ee ie Vi 2 mais ae | Eg : ag mit SL LTV “id he Oy To or Lact rhe | at as eee |" : ; : yi LO ae es: hb a** ri” av nite ¥ “ive eal hi ieee sat rae “iby! i t arly satents ip? iy ; oes sos d jiLaisioeya jal a TS x “Pruitt: oe pir rs ‘ i” t ey Tee Se vedo odd to iieee Hie ‘hairkeat Bans } } ates rh id atZ fi if iy) fad 8 eynels vg ‘ome i wah ae ba ean | Vs) Yyert Loeyy tics Rw nia he , Coe - et feabyy On © iSsinw whe . \ oy Health anh E191 onl) “4 iets r ive A TE ; ; ey = ‘ ; + ' ai 70 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO the point of junction is now only 488 feet above sea level. Mattawa is fifty miles north of the parallel of Kingsmere, and with a rise of three feet per mile would be 150 feet above it when the Champlain sea reached its highest level. The Kingsmere beach is at 690 feet, so that the highest stage of the sea at Mattawa must have been 840 feet, implying a sinking of 352 feet before the Nipissing Great Lakes ceased to exist. Mattawa is 150 miles north of the parallel of the Thousand Islands, and it is probable that while the outlet of Lake Ontario rose 246 feet to its present level Mattawa rose at least 488 feet, allowing for differential elevation toward the north, which would be only 1.6 feet per mile, about one half of the rate worked out for the highest beaches. As the Iroqucis shore is about as mature as the present Ontario shore one may assume that the two lakes lasted about the same length of time. If we suppose them equal in this respect and take Spencer’s estimate of 3,500 years as the age of each of them, only 7,000 years are accounted for out of the 20,000 to 39,000 years provided by the Niagara chronology. What happened during the other 13,000 or 32,000 years? Was the marine episode with its feebly developed beaches four times as long as the Iroquois or Ontario time? This seems highly improbable. and the natural conclusion is that Ontario has lasted at least 8,000 years, and that it was for a long time contemporary with the Nipissing Great Lakes. From the previous discussion of the correlation of events in the upper lakes region with those in the Ontario basin, it appears that during at least part of the time when the Trent outlet was in operation the water in the Ontario basin was at its lowest level, that of Admiralty lake; and that Lake Ontario was in existence and the marine waters had sunk 350 feet when the upper lakes were drained by the Mattawa valley. It does not appear, however, that the duration of Lake Iroquois or of the marine stage or of Lake Ontario can be exactly placed with reference to Lake Algonquin and the Nipissing Great Lakes. Lake Algonquin certainly lasted a ; ) . - ® ri 7 : (ie F - a : — ee ir \ P .- ¥ V2 o , 5 | J * - i py 2! r] = \ y p ¥ 7 i J 7t 7 fae = 4 € p q : 2 to Bea ' #4, > ie oer? “ss ‘ Z é p>; ‘. i Lhd SIE = il at ri 7 - CORRELATION OF EVENTS IN THE GREAT LAKES REGION 71 ~ long after Lake Iroquois had ceased to exist, and, on the other hand, Lake Ontario, which began during the life time of the Nipissing Great Lakes, has long survived them. Probably Lake Ojibway, north of the divide, was a contem- porary of the Nipissing Great Lakes for a part of their history, but came to an end long before they were trans- formed, by the northward rise of their outlet, into the present upper lakes. Correlation Table From the foregoing account of the lakes and arms of the sea which have occupied parts of the present St. Lawrence hydrographic region, it is evident that the relationships of the successive bodies of water in the different basins are very complicated. The following table is intended to make these relationships more clear: In the Ontario Upper Lake Hudson Bay | Eastern Ontario Basin Basins Slope Lake Duluth, Lake Chicago and ;|Lake Agassiz Lake Warren 25,000 | Niagara Falls and jLake Algonquin jbegins with St. Clair Outlet Trent Outlet into Lake Iroquois “LD! Se misleveta alee gata | CER DAL Sie. Sets oat en tis 17,000 |Lake Iroquois ends Lake Frontenac Admiralty Lake Trent Outlet into Early (Low) Admiralty Lake Marine Stage t. Clair Outlet Nipissing Great jLake Ojibway |Maximum Marine Lakes begin, Stage Mattawa Outlet re rr ed rr, Shs Sok Gib fi ae oe ce siviefin vend snes caneee 8,000 ;|Lake Ontario Marine waters begins \Two Outlet Stage} ;|Marine Stage fscdually retreat 3,500 Upper Lakes to Quebec Gilbert Gulf o ? ee =] (Ee ad Dull ss 6 90, © 6 a) elereiellia le\e ce wie sa 6 «uc allie es = & © v2 80 5 6 t..0 88 q ay ; oa | bs: / ' : ty i It Howe eaea.l 1 ts nl acl aD . * ee ord fo. wie (ees a.) lade as ped “eure rey: ; efit afd pl peal hin Sd 3 yl yr er ants aK. | oe Bho] al eal metad. ur jaa upihes a eaw.obivih of) Jo nebo breed Le’ ie hb se) ged Jas *yoietlag ts ‘4 anise wen “ail digl«l enol has tis oe naan en hate) 24hiye oo!) Jo arian “a : +6 ‘ rn t ih S. MOSTAVT a ; ae Pw beeP aa) ‘oh oh'4 her 7 “A i eTye ie ere, sO gle + s ‘ ; "9 HM ify OTt) a: ; ergs 2a ray =e) y 1; if 4 ran a best wl BT fhe as sia : d Ast) if isl ail ity Py k Usted aa et end Wo suo alle meals none # at OH why ra ad r nigefl . o> “-e pou pee ae odot i.) < ie | nes we — eas bp he fh 7] ‘dite @r : . sire eA ; ll eee otvtonpapyl mY . ’ ., teJ) wy hinlll enn awl nt - 7 , |\. atl ea : Cal par ee 1a “} wavehit } ytiel re ° ‘ vs . ‘ 7 155); & wnredAl tarot vieubangl Rate wight | \f | | =5 Hf ———_ 72 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO In the table just given the estimates of time are more accurate as they approach the present. That the Upper Lakes, Superior, Michigan, and Huron, began 3,500 years ago is fairly certain from the length of time required by Niagara to cut the upper great gorge. That Ontario began 8,000 years ago is made probable from the rate of wave erosion at Scarborough heights; but that Gilbert gulf, Admiralty lake and Lake Frontenac required altogether 9,000 years is merely a guess. The beginning of Niagara Falls and of Lake Iroquois is put at 25,000 years ago, while the various computations drawn from the Niagara gorge give from 20,000 to 39,000 years. The dates of the earlier lakes are, of course, still less certain. Lake Agassiz may have begun 45,000 years ago if Spencer’s estimate of 39,000 years for the work of Niagara is correct. On the other hand, Warren Upham suggests that 8,000 years are sufficient time to allow since the ice began to disappear from the Winnipeg region and that Lake Agassiz lasted only 1,000 or 1,500 years.' It may be that the late glacial and post-glacial chronology of eastern America will ultimately be put on an exact basis through the actual counting of the annual layers of clay deposited during the retreat of the ice and since that time, as DeGeer and his assistants have done in Sweden. Baron DeGeer with two assistants undertook last year to measure portions of the stratified glacial clays of eastern Canada and the United States for comparison with the results obtained in Sweden, and apparently they have been successful in correlating part of the American post-glacial record with that of Europe, though the work is not much more than begun.? Connections Between the Lakes It may be useful to summarize the connecting links in space and time between the various lakes referred to above, IGlacial Lake Agassiz, U.S. Mon. XXV, pp. 258-244. *Petermanns Mitteilungen, 67, 1921, June-July, p. 124. itt: 2 ee Mth sett ant i Accademia PuUOd.S Pires We HA Aue RRQ ih, sOtt=¢ Cy ete Te Nx ong nner nis Moe hie WATAEE BY ESAS | AP Ae org" Hts sont ot 77 nryi) r. vais tabs } ‘3 a? bibs eit alan denial? 3 _ oye? 19913 Tareas W3: ty whet oad) 5 «eRe Out ai a} ! wild wand Gud i xtdniad 7? a = i Ae te pe Vian 71 “3 thie d : ye Serge z Lk . i : bias F] iF Dea (VM: a tee oud 49 es > ivits >* , 3 vv, ' ASPs beng MB Looe L “ha ifs se i ares ve $44 " Vea. me: es ti vibe i ait tes vy SH Ady veermen it eet Bie7 aghe 2 7 cle Lad hey ey 9 OU Le ip stain aeial ile are’ i deepal ads ELD, ‘Shino d Ds foihile on 4 Si ae rrr (yet ten, | addy Cees 5290 1] yy bi, Y mer OU wee — tae, heal | Py sain ree 1s ie wht to “Wilt noes = fel ml aap in SmertIot ont ' an 47708 eg eee Honk det 0) Mesh 4d} oyu bata lodite - . Oat. nr ond Ate noe rnge Ta F ¥3. Sa | gure yode yioeee | pect } teat An Terie es’ ane ion ©) show ef ‘— . , _ Jnit-guircourioe od. annie rode? bondi walel endices CONNECTING LINKS BETWEEN THE LAKES 73 so as to show in brief form the possible migrations of the fresh water fauna. Lake Agassiz was probably connected with Lakes Duluth, Chicago, and Warren vid the Mississippi, since all of them drained into that river. It is possible, however, that Lake Agassiz came to an end before the other lakes began. Lakes Duluth, Chicago, and Warren became connected, as the ice lobes retreated northwards, and formed Lake Algonquin, which merged into the Nipissing Great Lakes, which at length passed into Lakes Superior, Michigan, and Huron. Lake Algonquin drained into Lake Iroquois vid Niagara and Algonquin rivers (Trent valley), and later into Admiralty lake during the early low stage of sea level when Algonquin river extended down to what is now the Bay of Quinté. Bays of Lake Algonquin extended across the Hudson bay watershed at three cols as the ice retreated and thus connected with Lake Ojibway and ultimately with Abitibi and other lakes of northern Ontario. It is probable that Lake Ojibway drained for a time into the Ottawa and thus into the Champlain Sea. The Nipissing Great Lakes drained in part through the Mattawa outlet into the Ottawa valley, then occupied by the sea, and in part z7@ Niagara into Lake Ontario. Lake Warren merged through various stages into Lake Iroquois, which drained through the Mohawk valley into the Hudson valley, then occupied by the sea. Lake Iroquois sank into Lake Frontenac, which was lowered tothe Admiralty lake, which rose to correspond to the rising sea level and merged into Gilbert gulf, the water remaining fresh although connected with the Champlain sea, an expansion of the Gulf of St. Lawrence. Gilbert gulf passed gradually into Lake Ontario, which began just above sea level as the land rose, so that the St. Lawrence river was at first very short but grew longer as the rise of the land continued until the sea level became stationary at Quebec. Anadromous fish must have slowly adjusted their instincts to the lengthening of the river. The Upper Lakes have had connections, often very long ; wi Pd at sonal ster rag? Ov eae ee + i Pas 43 ii reed be valier o)s2206, “fit” ine 1 NE “rey 7. io . oe oa tachi! wate T the 4 peer ay rh Lapeye NG on 2) blige t Wie Ui eran ciim bas ary, ots ie rte es ett. ws wR ei } hPa wr “I ovr am Muse eee) 7G jal veteitent hie ri rie ITI Lay) ting ME verte ie ORD outa Hh TY ine. Lair tial Erg wy iy VATROR yada nui Ye . ie Poni 2 gt r Ts) sh ober ferme 7 iis’. bn. .raretha Wg iomadite « isd aint bee i i . | MA A. cnt ‘ime Ae pis? ws ivallay ten Ey av nikpag i NS ary ‘ag a yeratily : on cde or gvoh bebaotts Lit nr nt: lene ty mca Ss tal a oes ™ 5 a5 : 1A ote a WW, i; | Osler ptadd ton i dstelhag ia ; hal hs Ns nie vig. ; ; i OTP eee re b \ nierntia “Down agtertt Ay Aaueatl aides shin? wye onl) tA] Lojqunon 198 6. lied sh Susehanad sha eal vine ath ne shail oe ol) 6d Bogaert a uriat aikn ora Side ee "ohn" ee 2 re relma = ant pint edie Deeg Te pave “Soicas) es to ent ‘ies Pa ae nat ate Nas yuievy 19s aw 130 ‘oon a a Ang aio bere iies Haat uty; ih adh eyelets) Fe xfouty. 48 J : i ania lesa Ott. 348 Orn Pin par ry Zn ae >. ytial ‘S19¥ > ‘sates FVKOIN tak! , ‘ : i ’ S, ‘i = an a c f ; = : - i iS te gee 74 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO and roundabout, with the Gulf of Mexico vi@ the Mississippi; with the Atlantic v7@ Algonquin river, Lake Iroquois, the Iri-Mohawk. river, and the Hudson channel; and with the Gulf of St. Lawrence vid the Mattawa outlet and the Cham- plain sea. None of these routes presented serious impedi- ments such as important vertical falls. A sea salmon could have made its way up any one of the three routes. At present the falls of Niagara form an impassable barrier to any ascent from Lake Ontario or the sea, but a few thousand years ago there was much less difficulty in ascending to the Upper Lakes by the Ottawa and Mattawa valleys. The sea, probably somewhat brackish, reached as far north as the Mattawa river, now 488 teet above the sea level. Above this the climb to Lake Nipissing is only 154 feet including rapids but no important falls. Any fish with good swimming powers could have made the ascent. Pleistocene Changes tn the Life of the Region As a sequel to the physical history of the Great Lakes region one may consider briefly the relations of its flora and fauna. Unfortunately the evidence as to the former plants and animals is very fragmentary. In the account of the different glacial and post-glacial lakes, lists have been given of the species tound in their shore deposits. All appear to be still living. The most frequent fossils are naturally shellfish and except in a few cases they still survive in the modern lakes. The few ex- ceptions are of Mississippi molluscs, which seem to have died out in these waters. The largest flora and fauna of post-glacial age are found in the marine beds, especially in concretions from the Leda clay of Ottawa. So far as known, none of the dozen plants, which include a few trees, differ from species of the present flora. Of the marine fishes and mammals none are extinct, so far as one can judge from the fragmentary remains pre- served. Shellfish are naturally the most frequent fossil forms. Dawson describes from Ottawa and Montreal Po s aa ou wtaCy va aea Wri ATO Tega Ma "1At , ; Ge Tis = ayth - CS . ¥ mupnep ts) ortigly | sale Hi: ot Lits . voree’ fh is: it PP ihe Sole ew? i i, one § Seer areas 2] ry oe Lyagare re ate) yur : Paty f ite) ano . lcm fe Cather Le agar “nif 9 pay OR Ke if as, con f | ; . io thé es Soe tit tons fay 1" onda bebe, inte! teanicw 7 T mined Beet ae ts: ws3)3 if) 4 si 19 1 yl e eles sew airtel (O ohh Aa 1 WE rit at a] ie Sea +3 yi UL ta it ) witht Webde oT =e yoy 8 ist 43 ote MWR, ot - iy ; es oS s i“ he b ere ry vn i thee! oak ayer ) 14 Lwin 4 Te ae ee debe “anPrrod 1 On tbe re ut biifem bp f p Need | is Re Per oT. wi . as 4h hh ete it! insite ont of Of. Ew 1 ecw vane q rh vied no {i iio) ae Sonohiys od see : | catnomg er eae eis . era Se. nloahy aes ai ota 400% ae ; Sel okt ys iy re nit a T ‘ > 4 Y enivil ie) oo: 36 —., ; yoo eee dd Ajdenty Vilna bi ; AT .jented cio eae atv: il nea hoi’ ee semis eee rea = stn yor, IRC Ede) atioit ‘ens ee olive , i Vibe 4) mul’ sagety oi to aon 9 yeaa: ane ang sl le eo el | voit Pres c=] rn ee et ee sith Sie ' rey 40H me VIIA ‘] wit} pak. Giuntl cseapewl fae aie (hee aneiett tna daeasie es PLEISTOCENE CHANGES IN THE LIFE OF THE REGION 75 142 species, of which he believes only two or three are ex- tinct,! though some others present varietal differences. On the other hand the only sponge mentioned no longer lives, and all the insects, four in number, are extinct, according to Dr. Scudder. ; The marine beds were laid down probably between 17,000 and 3,500 years ago, so that specific changes in molluscs seem to require a longer time than that, while the more highly developed insects, with their rapid succession of gencrations, are more quickly modified. The only other important Pleistocene flora and fauna for comparison with present day species are those of the Toronto interglacial beds;? from which 63 species of plants and 122 species of animals are on record. Of the plants 35 are trees, and all but three or four still exist, the extinct ones including Acer pletstocenicum, A. torontoniensis and perhaps another maple, and Gleditschia donensis, as determined by Penhallow. All are closely related to modern trees, the maples being apparently ancestors of our sugar maple. Among the 41 shellfish,. all are still living in North America, but apparently eight belong to more southern waters than Lake Ontario. The 72 insects are extinct with two exceptions, according to Dr. Scudder, as one would expect from the fate of the much later insects of the marine time. The two or three fish remains are too fragmentary for us to be sure of their species; and the six mammals are known only from separate bones or horns or tusks. The elephants, of course, are extinct, and the horn determined as cervalces by Bensley may be looked on as extinct also. Whether the large bear, known only from one of the bones of the head, the red deer, the caribou, and the bison belong to living or extinct forms cannot yet be decided. The Toronto interglacial formation probably dates back at least 500,000 years and the changes in the life of the 1Can. Ice Age, p. 279. 2Geol. Congr., 1913, Guidebook No. 6, pp. 15-25. | 4 ‘ ‘an 4D | > » ¢ 1 J ii ~ BOOTS aap i ogehy (tee Scotr etii : ‘| ‘ j » Ad rep he < weirs a) nh ye B t } FA ‘wee > 4 J if * ‘? LS oi ie. OF ) wre.) rare lnisrae ane mt} baie ; rays Shale wit rs m tat: sis eeey lity she te WE 7 . wit f 77 i i. ; elit ee eer. , oh yh G Fe is «hramegq 3, ge) @gnaha ohh, Artis os ee ese . a 4 waite “tj pt, dohiwes D4 vain rte vany wthhik a Ls May af rrrate Lpel he che wn? Awe nase hk 7 1 = U ae ba, hy n'y P isi ‘i ia stain’ “hoy _sonoas Ast 16: obdie web: Ipekeetety ee $ ‘by. Seid cont) be a I be | Met (Sis wel ia 1B ? Was eager wet ieniciin. pe uote, aor raat a wy i paicd asheesre 2 iva ann The ate e | — ipewn’ lead. ott Ms bd ican S ; fot gir MO SF. yf ante! Ray | mid, ese mh , no ae Hee x , : AT sella. { h ie 3h vie 3 b rilahn vk ed [ony ne LBCeReL fast y tone 5 r i a alt oy «) a eae, i 76 GLACIAL AND POST-GLACIAL LAKES IN ONTARIO region seem surprisingly small for so great a lapse of time. Apparently molluscs have the slowest rate of change and insects the most rapid; the mammals perhaps being modified almost equally rapidly. While no older Pleistocene deposits are known from Ontario there is reason to believe that the Aftonian beds of lowa are more ancient, since they occur between the Nebraskan and Kansan boulder clays and occupy the first of four interglacial intervals recognized in that state. If the Toronto formation comes about half way down in the Pleistocene, as seems probable, the Aftonian must be some hundreds of thousands of years older and its flora and fauna should be correspondingly more ancient in character. Among plants, pine, tamarack, oak, elm, ash, walnut, hickory, and sumac are reported, but unfortunately the species seem undetermined, so that one cannot be sure as to whether they are of still living forms or not. It may be observed that all of the genera except the walnut and the sumac are represented in the Toronto formation. Among animals only mammals have been found, and these have been determined mostly from isolated bones, jaws, and tusks; though the remains are more complete than those found at Toronto. Calvin describes and _ figures remains of two or three species of horses, a camel. two ground sloths, cervalces, two mammoths, and one mastodon.' All the mammals are extinct and have left no North American descendants, unless cervalces is an ancestor of the moose. It will be recalled that cervalces and either mammoth or mastodon occur in the Toronto formation, but that the other mammals may be of still living species. The number of extinct species of mammals is greatly increased in the older interglacial formation. 1Aftonian Mammalian Fauna, Bull. Geol. Soc. Am., Vol. 20, pp. 341-356. . a Ps prt es ong i } as oe q [ de - OUueAT 2th we re Dn ae Pe yA mae | TA —_—— | ves ¥ pre Law ri rit rie AP ime 5 i y : : “oa v1) ie MeiVe ails yea ah loud a ry ‘api ts 4 Tayi ee ba) kee ee ' y . : {bron gdt month a Mi: FPO Lota “kia 1 pobatisfe) Thy ORO F nent LEFT / P \ EP VWOTke vi | , ; ‘ rit ; “4 SL ' lobes oft ts | ny we ae Sey) elev Saehihs Al ‘aly 30 ; sew Tal note! Sante me artveal e pe Th eT LL ent ati oy. te. ot) preted! (or ae bd oi wig ett hee 4 Jive Var teo omuninhiy ' vt wate wartton ii ne OO POga ~¢l oh Meat ee nay one esi » | | ¢ ¥ ' Se oO ‘pee 4 ini ‘i mit etaee tk vf AMET Any rAd LN De, on bine Melis reer nok Any “a ct noid rudolph, eh aor i , ‘ rinea tte: e i Wiha | yey ieste : vioh. Wit. epee Gh ene sisal $0 “* oh aa at Rei ec oy tee 46 0 beh oe ah vist ‘eal a vant! ' ‘vue Porth, ck DO Le inf over hth Jee we fae) eo Laies, 22S HG CIE nists aki UNIVERSITY OF TORONTO STUDIES BIOLOGICAL SERIES ———— No.1: The gametophyte ot Botrychium Virginjanum, by Biel [ER ERE Ys cae ne alee einen eat ine ee No. 2: The anatomy of the Osmundaceae, by J. H. FauEn. Mo 32: On the identification of Meckelian and mylohy oid grooves in the jaws of Mesozoic and recent mam- malia, by B ARTHUR BENSLEY.....---- 220+ -+--- No. 4: The megaspore- -membrane of the Gy mnospernis, by Be AER BOUSON.. RAPE Ct Uae HOPES 1 cae Ae eet ae ae No. hs The homolog IES of the stvlar cusps in the 1 upper molars of the Didelphyv idae, by B. ARTHUR BENSLEY No. 6: On polystely in roots of Orchidaceae, by J. H. WHitrrE No.7: An early Anadidymus of a chick, by R. RAMSAY ReincINC oo - feee Sie SNe tae ee Ke. ee ge tts No.8: The habits and larval state of Plethrodon Ery- thronotus, by. W. Ey PIERSOL iio oe so es No. 9: Spawn and larva of Ambystoma Jeffersonianum, by ML eee RSI na ca fone pe oecaory se cy hss to No. 10: The colour changes of octopus vulgaris Lmk., by eRe Sra tree Gee iat ges Sa one dieid alo'> vite a's ts 0 OE No. 11: The North American Dragont flies of the genus Aeshna, Rey VAL BER Gc toes noha eie' 2 oe ee No L2: Ascidians from the coasts of Canada, by A. G. EUS Se en ee oa Ree AR Ee ee RORY Wo 13: A cont ribution to the inorphology and biology of insect galls, by UES Ts ols Genoese team a sen Sie No, 14: Fg ro Maturation, chromosomes, and spermatogenesis in Ee claps, by ROBERT CHAMBERS). 002. ss 28-42 Wo. 15: A new ce stode from A mia Calza L., by A. R. C OOPER No. 16: The egg-laying habits of Plethedon Cinerens, by Wort SO ae en ols gel elses cee he No. 17: An ecological study of the mayfly Chirofenetes, by hee eS SEN Tay ces Oe ana No. 18: The Isopoda of the Bay of Fundy, by N. A. WALLACE No. 10: Anegg ot Struthiolithus Crersonensts Bra indt, by B. A. aren Ne a oot ee ah hac leit wera RS ace ee RS Soe’ No. 20: Publications of he Ontario Fisheries : Research Lab- iene ON es wee ns Sag thse 8 ns oe No. 21: Publications of 7 the Onta io Fisheries Research Lab- oratory, N (Glacial and P ost-Glacial Lakes in Ontario), by A. P. COLEMAN......-.-----+-+: eens O.p. 6) Ya f O.})- University of Toronto Studies COMMITTEE OF MANAGEMENT Chairman: Str ROBERT ALEXANDER FALCONER, LL.D., K.C.M.G., ; President of the University PROFESSOR W. J. ALEXANDER, PH.D. ProFeEssor J. P. McMuraricu, Pu.D. a Bric.-Gen. C. H. MitcHELL, B.A.Sc., C.B., C.M.G, D.S.O. ag Proressor G. H. NEEDLER, Pu.D. 3 a ProFEssor GEORGE M. Wronc, LL.D. General Editor: W.S. WALLACE, M.A., Associate Librarian of the University UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 11 THE LIMNOLOGY OF LAKE NIPIGON BY WILBERT A. CLEMENS OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1923 j vo Art 4 " i Ve 4 Os pede Bat } THE LIMNOLOGY OF LAKE NIPIGON During the months of June, July and August, 1921, an intensive limnobiological investigation of Lake Nipigon was commenced by a field party from the Ontario Fisheries Research Laboratory, of the Department of Biology, Uni- versity of Toronto, under the direction of the writer. The purpose of the investigation was to carry out a thorough study of the biological and environmental factors, with special reference to the fish fauna and its economic and conservational aspects. Lake Nipigon was selected for investigation because of its isolation for a long period of time from the Great Lakes and, as far as known, from any other drainage system. Asa result, natural conditions have been undisturbed either in the drainage basin or in the lake itself, except by reason of the opening of the lake to restricted commercial fishing in recent years. In 1916, the Department of Game and Fisheries of the Province of Ontario opened the lake under supervision to commercial fishing for the purpose of augmenting the food supply during the war, and have continued the policy with some modifications to the present time. Statistics are available for only four years, 1916-1919, but during that time the following amounts of fish were removed: whitefish 2,511,614 lbs., lake trout 1,059,632 Ibs., pike perch 51,431 Ibs., sturgeon 21,810 lbs., other species, chiefly pike, ciscoes and northern suckers, 57,694 Ibs. In 1920, the following plants were made: white- fish 8,943,000 fry, lake trout 734,000 fry, and 240 parent black bass. It seemed desirable therefore to determine the conditions existing in a lake before the effects of such disturbances of natural conditions had become pronounced. An opportunity was thus afforded of studying the effects of commercial fishing in a circumscribed body of water and of 3 4 CLEMENS: LIMNOLOGY OF LAKE NIPIGON developing working plans for the best utilization of the productive possibilities of such a body. In view of the above considerations, the investigation during the first season followed three main lines:— (1) the identification of the species of fish, their distribu- tion, relative abundance, natural history, food and rates of growth; (2) qualitative and quantitative studies of the food organ- isms, confined largely to the plankton and the bottom fauna; (3) a study of the physical features, particularly tempera- tures, dissolved oxygen and carbon dioxide, colour and transparency, and the significance of these factors in respect of the biology of the lake. Some of the results obtained are presented in more or less preliminary form in this and the following five papers. Lake Nipigon, the largest of the many inland bodies of . water in Ontario, is situated in the northwestern portion of the province about 50 miles north of Lake Superior. It lies approximately between 87°35’ and 89°10’ west longitude and between 49°5’ and 50°30’ north latitude. The area as given by Wilson (1910) is 1769 square miles, of which about 1530 square miles is water surface, leaving an island area of about 239 square miles. The lake is roughly quadrangular in form, about 50 miles long and 30 miles wide, but very irregular in outline with numerous bays of various sizes and contours. The shore line is characterized by rocky projec- tions and headlands alternating with indentations. The latter, when open and exposed, are rocky or sandy, but where extended and protected, tend to become muddy with an approach to marshy conditions. The length of the coast line exclusive of smaller bays and coves is over 580 miles (Bell, 1870). The islands are numerous, particularly in the northwestern portion, and range in size from small rocky projections to large wooded areas. The elevation of the water surface is 852 feet above sea level (White, 1915). The drainage basin comprising about 6,000 sq. mi. is fairly well covered with tree growth consisting chiefly of balsam, spruce, poplar and birch, with a scattering of red pine, jack- CLEMENS: LIMNOLOGY OF LAKE NIPIGON 5 pine, tamarack, cedar and alder, and the major portion is included in the Nipigon Forest Reserve. Numerous catch- ment basins, ranging in size from small ponds to large lakes, \ Nipigon House <} , £ 350f- 4.8. Co- ote ¥) Champlain Pt (71 Fee Venn 4 Grano © Me (nT VRE 8 G sar v én ° Sar, Q © Aumaccor ff Bay et ae & Livingstone Fé. 402 440 EAST Sar Mungo Park Py Map of Lake Nipigon showing depths and the location of the three stations for the taking of plankton, temperatures and water samples. 6 CLEMENS: LIMNOLOGY OF LAKE NIPIGON Nipigon by a large number of streams. There is, on the other hand, but one outlet, the Nipigon River, by which the drainage is carried southward into Lake Superior, through a descent of 250 feet, over a series of rapids and falls. The uppermost of the latter is the Virgin Falls (35 ft. high) practically at the origin of the river, where the estimated low water flow is 5.500 cubic feet per second (Hydro Electric Power Comm. Report, 1907). The region surrounding the lake is typical of the Arch- aean area. It is rugged and hilly, with bluffs rising as high as 600 feet above the level of the lake. The present features of the district are the result of a long series of geological events whose sequence appears to be as follows. A trough existed on the Archaean surface which during a period of submergence received large deposits of sediments. (Wilson, loc. cit.) Elevation above sea level followed with extensive . erosion uncovering portions of the original trough. Diabase flows later invaded the area, but in a succeeding long period of erosion much of the diabase was removed as well as a large amount of the early sedimentary deposits. During this period, and possibly to some extent earlier, disturbances occurred in the earth’s crust resulting in elevations, block faulting, etc. It is probable that there were two outlets from the basin during this time, one from the southeast corner through the Pijitawabic canon (Orient Bay) and the other from the southwest corner by way of Black Sturgeon Lake. Then followed the glacial period, and when the ice finally retreated northward, the Lake Nipigon basin formed a very large bay of Lake Algonquin, the predecessor of Lake Superior (Coleman, 1922). With the development of the Great Lakes drainage system and the elevation of the region north of what is now Lake Superior, the modern Lake Nipigon was formed. A new outlet developed, the Nipigon river, the two preglacial outlets being blocked by glacial debris. There has been as yet no hydrographic survey made of Lake Nipigon. A line of large islands lying roughly in a north to south direction divides the lake more or less into east and west portions. The eastern portion contains the “J CLEMENS: LIMNOLOGY OF LAKE NIPIGON deeper water which lies well over towards the eastern shore. Soundings taken in the course of dredging and limnological operations gave the following depths:— 195 feet Pijitawabic Bay, opposite Macdiarmid village. 345 feet off mouth Blackwater river. 225 feet off mouth Sandy river. 225 feet off Mungo Park Point. 402 feet was recorded by McInnes (1894) two and a half miles south of Livingstone Point. This is probably the greatest depth in the lake. In the western portion the following records were obtained :-— 198 feet west of Kelvin island. 171 feet east of Champlain Point. 264 feet northeast of Grand Cape. 285 feet was recorded by McInnes just north of St. Paul island. It is thus evident that there is a very large body of deep water in Lake Nipigon and that the amount of shallow water is limited. The open shores are rocky or sandy and because of the size of the lake, subject to strong wave action. It is only in the deep, protected bays and shallow areas among the islands where conditions are such as to permit the growth of the larger aquatic vegetation with the accom- panying abundance of animal life. Because of the great extent of Lake Nipigon, the first season’s investigations were confined largely to the south- eastern portion. Three stations were established for the taking of temperature records, water samples and plankton. Station 1 was located in comparatively shallow water (depth of 28 yards) off the mouth of the Nipigon river; station 2 in the deepest water (depth of 63 yards) of Pijitawabik bay (Orient Bay) directly opposite the village of Macdiarmid; station 3 in the open water off Sandy river (depth of 63 yards). One series was obtained on August 29 off the mouth of the Blackwater river (depth of 100 yards). The temperature records were obtained with a standard- ized Negretti-Zambra deep-sea reversing thermometer. Water 8 CLEMENS: LIMNOLOGY OF LAKE NIPIGON samples were taken immediately after the temperature records with a modified Kemmerer water bottle (Birge, 1922). The water samples were put in 250cc. glass-stoppered bottles, packed in broken ice and taken to the laboratory where the analyses were carried out. The oxygen content was determined by a modification of Miller’s method (Pro- vincial Board of Health of Ontario, 1920). For the carbon dioxide determinations N/22 sodium carbonate with phenol- phthalein as indicator was used (Prov. Board of Health, loc. Git.) Fiche temperature records and the results of the analy- ses are given in Tables 1, 2 and 3, and from this data Figs. 1 to 6 have been eed Since no record of the temperature of the deepest water was obtained in this season it has been assumed, on the basis of the results obtained at the three stations, that the temperature was 4.0°C. Fig. 3 may be taken as a fairly accurate record of events. From June 17 to July 9 the sur- face waters were gradually acquiring heat and the thermo- cline was slowly increasing in thickness (Table 4). No record for late July was obtained for Station 3, but judging from the record for Station 2 on July 26, by the end of July the epilimnion had doubled in thickness and the thermocline therefore had shifted downward. During the morning of July 30a wind of considerable violence came up accompanied by some rainfall. The wind blew first from the southwest and gradually shifted to north- west reaching its maximum velocity about 2 p.m. No meteorological records are available for the Nipigon region, but judging from records obtained at various stations in Northern Ontario, it is probable that the wind attained a velocity at times of about 35 miles an hour. Captains on the fishing tugs stated that this was the worst mid-summer storm they had experienced on this lake. The temperature records on August 2 show the effects of the storm. The thermocline in the open part of the lake was obliterated. Apparently the three strata were set in motion and some of *All the solutions were prepared by Mr. George W. Lucas of the Department of Chemistry, University of Toronto, and the Provincial Board of Health. LIMNOLOGY OF LAKE NIPIGON CLEMENS 0% 98 49 €1T came 06 ro 8ST 6ST 0°9T L'l 88 69 GOL 209 | 38S % | *xO | ‘DO ‘dway SI jsn3ny TT O'T 6°0 9°0 “OD 68 G6 68 48 L8 Dee 8 °F Lok 19 tL GIT SFL 0°9 1°81 8°¢ Z6r Z 61 POL 8°¢ FOr "WS % | “XO | 'O ‘dwar ez Aint 0°0 68 0's pS 88 6'L G'¢ Lar 88 ey) 9°¢ Lg eal 06 0°8 g'¢ g-¢ 9'T 68 GZ 9 19 8°9 pra R22 +6 82 £°8 70D | 38S % | °xXO | ‘OD ‘dway gT eunf I aoRying spied ur yidaq ——_—$— a a a pc T NOILVLS Tt ATaVL CLEMENS: LIMNOLOGY OF LAKE NIPIGON 10 ov S9 8'0 98 0'°8 Uh a £9 |g 3 G8 082 09 ev og Te 48 6°2 9°F 88 0'°8 Og oer 8'P cP | tel & L8 82 [We i OF 8'P GE Jig! 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[oeuer spied ut LI ysn3ny I Isn3ny 92 Aint yidaq % NOILVLS (panurjuor) % ATAVL LIMNOLOGY OF LAKE NIPIGON CLEMENS 12 Go| 8 G9Q°L 9°F £9 P-Tet vs LL 18 88 th ae oO of Ej S88 0 ‘T | 06 6° T 9 9 ‘8 Vis 75 9°Z T 88 € og T [ZT | 28 9-2) G9 TeT—-68 672 8'°¢ L’9 I I} 06 v9 So gE=sh e065 S2L|-— 2-6 ZT 6 62 GC eT 6-Si- = tr S| 56 ¥Ol-—=—9-9 | Se Se ee: £9 GOL Jot | 66 €°9 99) '38S%| “xO Io'dweayz} Oo] 3es % | "xo |-5-dura y OO 17%S%\°xO | ‘DO ‘dwel] ‘p ‘dway T |e! vor | 82 vst 0 2% PT | 901 Ll wD 1d N CO 19 I QdeJAING — J] spied Z snsny 6 Aint €z aunf LT oun[ ul yydaq € NOILVLS € WTaVL CLEMENS: LIMNOLOGY oF LAKE NIPIGON STATION 3 (continued) OE UE hs Depth August 297 in yards | | ee Surface ibe 6.05 87 ey 5 16.8 8 10 15) 15 15 18 9 20 8 25 6. 30 6 40 5 45 00 i=) rs Ke) .5 6.40 70 1.6 100 4.2 5.70 62 2.2 ee {Off mouth Blackwater river. co i=) ee CLEMENS: LIMNOLOGY OF LAKE NIPIGON 14 eee O° $6 | F'EZI-91 b-O1 Gee 82| OP o7066~ 4 Per pe eae uoruumjod Fy ¢°6 -0'ST 91-FT OI-1Z 8% |0°6 -0'LI Pee ts ee oagusan O°ST-T “LT 1-0 12-3 2-0 | O'1T-S “21 G0 for popes ‘J ‘duay | ‘w yidaq $o) GUIS, | sur yidaq ‘9 ‘dway | ‘w yidoq 6z ysnsny Z jsnény 6 Aint 2 ounf re ae ae | Sa ae re te Mee SP Sie Seg Ta BR ee em | Se Se Es ae € NOILVLS €°¢9 -%'8 O9-2T 8°F 9 01 O9-FI [fa ae aes 09-01 GP GL |0'090°2 {°° *"'"** +s ‘uormuNfodAy 68 -0'FI LT-2I SOT ZI PI-IT So Rre Oh e “OLY 6 £660 (0520-0 ye su ooraui, O°FI-S LT oI-0 LS 4 er ah | IT-0 G°6I-G 02 S-0 GhOleg=0210 TaDeD ee uoruwryid | ‘DO dway | ‘w yidaq | ‘9 ‘dway | ‘w yydaq | ‘5 ‘dway | -w yidaq | ‘9p ‘dway | ‘w y3daq ZT snsny T jsn3ny 9z Ajnf 1261 ‘8 Ayn &@ NOILVLS ya TavL CLEMENS: LIMNOLOGY OF LAKE NIPIGON LS the warm surface waters were carried almost to the bottom of the lake. At Station 2 in the sheltered bay (Fig. 2) the thermocline was not obliterated but was decreased in thick- ness and pushed downward several metres. We have here a good example of the effect of the wind in distributing heat in a lake. Later on in August typical stratification was again established as shown by the record for August 29. The records in Figs. 1 and 2 coincide in general with that in 53 3 to depths of 30 and 60 metres * abn i = SERESSEEuD HA 40 Sone suneccassaeesas Seesessess ESccccuscccrscsseces sesuscasscssccesccssceese Fic. 1. Curves illustrating temperature records at Station 1, Lake Nipigon. DISTRIBUTION OF HEAT. In the absence of an hydrographic survey of Lake Nipigon, it is impossible at the present time to calculate the summer heat income and the work involved in the distribu- tion of the heat on the basis of mean temperature and _ re- duced thickness. Calculations have been made, however, for a column of water one square centimetre in area extending from surface to bottom. These figures are of some interest in indicating the distribution of heat and work for those particular points in the lake but do not furnish very accurate data which may be compared with that calculated for other lakes on the basis of mean temperature and reduced thickness (Birge, 1916). In Tables 5,-6 and 7 are given, (1) the temperatures at five-metre intervals as taken from Figs. 1, 2 and 3, and (2) the mean temperature for each five-metre stratum. The latter 16 CLEMENS: LIMNOLOGY OF LAKE NIPIGON figures have then been used for the calculation of the number of gram calories per square centimetre of surface received by each five-metre stratum above 4°C. using the formula (T—4)x500. These results are given in Tables 8, 9 and 10 and show: (1) that by the middle of June about a third of the total amount of heat of the season had been gained and that this heat was contained in the upper half of the lake, that is, in the upper 60 metres; (2) that by the end of the first week in July there had been a considerable gain in heat by the upper waters, con- fined largely to the upper five metres. F 41 48 57 c2 3 4 5 6 7 8 9 10 ff #12 13 14 15 16 17 18 19 20 21 ane Bege> os} Bengeaanaes sens Fic. 2. Curves illustrating temperature records at Station 2, Lake Nipigon. (3) that by the end of July there had been large gains in heat by the upper 20 metres. Unfortunately no tempera- ture records were obtained in late July in the open waters, but it is doubtful if at the end of July any appreciable amount of heat had penetrated below 60 metres. (4) that the effect of the storm on July 30 was to dis- 17 CLEMENS: LIMNOLOGY OF LAKE NIPIGON OF 9 69 GLY 9°P G08 9°9 oc '’s 6 °F 09 SI ¢'6 G08 wey) 08 “ST LST Lie rr 00 ‘OL 66 ST 6°ST GL 81 Gest L091 G6 ST 1261 Cl 61 o 91 OF 61 "| ‘wi-g 10d ‘9 ‘dway ‘| ‘wi-g sed ‘) ‘dway ‘dway ‘Ay ‘dwoy ‘Ay 8ST ysnsny e¢ Ain mMroNn A © 1D oO HoH 19 12 O Bint O19 1D KRawKRwo dt HH ere ‘| ‘wi-g red ‘9 ‘dway ‘dway ‘Ay z Ajnf ‘dua “Ay eT ounf Ors ogg 09°¢ 08'S cr 9 08 '8 ‘| ‘w-¢ sed | ‘d ‘duay ‘uw yydaq a SSE lh a SE SS ee T NOILVLS—SAYNLVAAd NAL G ATAVL 18 TABLE 6 CLEMENS: LIMNOLOGY TEMPERATURES—STATION 2 OF LAKE NIPIGON Se NOnNOBANNOHHY E¢ HHODDSONSODSHH = | ew CoH wowoow wow B so gee =| . 1919 O 1 a (~_NOMHNOTHOrMOWHXH SUR ewHH BOOSH HH HH Trlr ore on [anes ae : ONrFKA ee) os ~KCOARDDOAMNDMH ME aes RRONDOKROOH HS . Sees pia (MA DW eel ulmidcs Bl Alijies 1D SO 19 Sth Spies KrrNOADOTANOOOM SO EO REROODN SO 141916 | o | sens | & SMONNORMDMRAR S60 SSHOHONSOOER YH © ane Swanrownmod dH Q re oO No =) = : 0nSOM MOMS a JjPHeatotOtH aon EU Senroeonnwoidaist Ares ite yea Sis 1919 19 1D 19 19 o & COMWMMDHOMOWADN Bd 1D D O19 19 1d HOH SH OH SH SIS dll pat 0 ss dees a = a. NOCHONANNHAMN EU SrrF-oenndtdtttst st N = Aa. Eg SCONREANRNOOrFAN Hd -remnoddtonomr~ OH =H =f 20 oro OomM Mot H H oH sl > oO 9 |< a : | pias ; a, Vel ele el isis) | Ye) S) Ep | woSn and SHnB% a Aoroomwotadtd dt st | SCmHOoOMonMnsoHOoOMOW | Sar AN OO HH HD 1d Depth m. 4.0 4.15 4.2 4,20 4.30 4.40 4.75 4.87 5.3 5.35 60 TABLE 7¢ TEMPERATURES—STATION 3 August 29 Av. Temp. Temp. C. per 5-m. I. CLEMENS: LIMNOLOGY OF LAKE NIPIGON ly ga! 16.92 16.27 14.30 10.10 6.95 6.32 16.75 15.8 12.8 7.4 6.5 6.15 t a Yen (o) Yon) =) ~ 1S © wo 5. 52 5.40 5.27 5.15 5.05 4.97 4.90 4.80 4.70 4.62 4.57 4.50 4.40 4.30 4.20 4.10 4.00 5.45 5.35 5.2 5.1 5.0 4.95 4.85 4.75 4.65 4.60 4.55 4.45 4.35 4.25 4.15 4.05 4.00 19 August 2 Av. Temp. || Temp. C. | Av. Temp. || Temp. C.| Av. Temp. Temp. C. Depth m. per 5-m. 1. 12.6 10.4 15 t Yes) 2 4.85 4.65 4.6 So wD ~ N oe ip “D> 4.55 4.5 N uD i ao 4.45 4.4 4.35 4.3 22.0 12.5 A} 12.2 15.8 8.7 (feo) 6.27 5. 67 4.95 4.47 4,27 4.17 4.07 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 6.5 6.05 5.3 4.35 4.2 4.1 4.0 4.0 4.0 4.0 4.0 4.0 10.45 8.05 7.4 6.35 5.35 5.02 4.87 4.75 8.4 ee 10 15 20 25 30 35 40 45 50 55 AN La N i 4.25 4.2 N o~rnNn HHOANNA TS iMiottddt ddd HHH 4,2 4.15 4.1 > oS 4.05 4.0 S 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 7.1 5.6 5.1 4.95 4.8 4.7 1D 4.3 4.15 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 60 65 70 75 80 85 90 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 100 105 110 120 123.4 20 CLEMENS: LIMNOLOGY OF LAKE NIPIGON TABLE 8 CALORIES ABOVE 4° C.—STATION 1 Depth m. June 13 July 2 July 23 August 18 0-5 1685 5900 7635 6035 5-10 1060 3475 7375 5960 10-15 850 1125 5085 5900 15-20 775 435 2025 4300 20-25 725 | 5095 || 300} 11235 || 750 22870 || 2025 | 24220 25-30 675 | 675 || 260 260 || 375 375 || 1200 1200 5770 11495 23245 25420 TABLE 9 CALORIES ABOVE 4° C.—STATION 2 Depth m. June 14 July 8 July 26 August 1 August 17 0-5 | 2350 5800 8000 6850 6210 5-10 | 1760 2500 5825 6810 5750 10-15 | 1435 1275 2750 6485 5435 15-20 | 1210 925 1510 4485 3900 20-25 | 835] 7590 || 700] 11200|) 1069} 19145/| 2410 | 27040]} 2025 | 23320 25-30 | 560 525 800 1800 1260 30-35 | 500 400 635 1450 1010 35-40 | 450 300 525 1150 900 40-45 | 385 225 435 900 825 45-50 | 310] 2205 || 175] 1625/| 360] 2755]! 725 | 6025|| 775 | 4770 50-55 | 210 125 285 575 725 55-60 75| 285 || 100} 225] 200) 485/| 4351 1010/} 675 | 1400 10080 13050 22385 34075 29490 CLEMENS: LIMNOLOGY OF LAKE NIPIGON yA | TABLE 10 CALORIES ABOVE 4° C.—STATION 3 Depth m. June 17 July 9 August 2 August 29 0-5 3225 7450 6010 6460 5-10 2025 4100 5885 6135 10-15 1700 1800 5525 5150 15-20 1175 1135 4750 3050 20-25 675 | 8800 835 15320 | 3750 | 25920 | 1475 | 22270 25-30 510 475 2635 1160 30-35 435 235 1760 985 35-40 375 135 1210 850 40-45 250 85 785 760 45-50 75 | 1645 35 965 | 510 6900 | 700 4455 50-55 375 635 55-60 310 575 60-65 285 525 65-70 260 485 70-75 235 1465 | 450 2670 75-80 210 400 80-85 185 350 85-90 160 310 90-95 135 285 95-100 110 800 | 250 1595 100-105 100 200 105-110 85 150 110-115 60 100 115-120 35 50 120-123. 4 0 280 0 500 22 CLEMENS: LIMNOLOGY OF LAKE NIPIGON tribute heat practically to the bottom of the lake. The amount distributed to the deep water is relatively small, however, the great bulk being retained in the upper five metres. (5) that by the middle of August the lake had begun to E; 41 48 57 66. c2 4 a. GZ & 9 10 it 12 14 15 16 17 18 19 20 21 22 Fic. 3. Curves illustrating temperature records at Station 3, Lake Nipigon. CLEMENS: LIMNOLOGY OF LAKE NIPIGON 23 give up some of its heat. The loss in the upper 25 metres has probably been into the air, while the 25-50 metre area may have contributed of its heat to the underlying waters which apparently continued to gain heat for some days follow- ing the storm. It is probable that the summer heat income for Lake Nipigon in 1921 was in the neighbourhood of 30,000 gram calories per sq. cm. of surface. The mean summer heat income for Cayuga Lake, N.Y., over a period of five years was 29,480 calories (Birge and Juday, 1921), with the maxi- mum in one year of 30,910 calories. The two lakes have approximately the same maximum depth, Cayuga 133 metres, Nipigon 123.4 metres. Distribution of Heat The amount of work in gram-centimetres involved in the distribution of the heat contained in a column of water one square centimetre in area has been calculated for each metre stratum using the formula (I—D) CG, where (I—D) is the loss of density due to warming, C, the depth in centi- metres and G, the weight in grams. This is based on the principle stated by Birge (1916), that for the most part, the warming of the water in a lake is done by the transference of warm lighter water from the surface downward through water at its maximum density, by the agency of the wind. This involves an expenditure of work which can be calculated. The results here given are for each five-metre stratum and the temperatures used in the calculations are the means of the five-metre strata as given in Tables 5, 6 and 7. Further accuracy in this instance did not seem warranted. The results are shown in Tables 11, 12 and 13. The significant points are :— (1) that from the middle of June to the end of the first week in July practically all of the work was expended in the upper 10 metres and particularly in the upper five metres. (2) that during July, the work was expended to a greater depth but especially in the 5-10 metre stratum. August 18 156.75 406. 00 650. 00 490. 50 143.75 63.00 August 17 165. 00 380. 00 555.75 409. 50 143.75 70.00 54.45 49.40 47.30 48.00 45.05 43.50 1482.70 g. cm. | 1910.00 g. cm. 24 CLEMENS: LIMNOLOGY OF LAKE NIPIGON TABLE 11 DIRECT WORK—STATION 1 Depth m. June 13 July 2 July 23 0-5 13.50 150. 00 243.75 5-10 14.00 144. 00 608.00 10-15 14.95 26.00 490.75 15-20 18.00 5.40 112.50 20-25 19.55 3.45 20.70 25-30 21.00 2.80 7.00 101.00 g. cm.| 331.65 g. cm. TABLE, 12 DIRECT WORK—STATION 2 Depth m. June 14 July 8 July 26 August 1 0-5 | 25.50 144.75 265. 50 198.75 5-10 | 38.80 76.00 390.00 524.00 10-15 | 42.25 32.50 149.50 773. 50 15-20 | 42.30 25.20 63.90 531.00 20-25 | 25.30 18.40 40.25 207.00 25-30 | 14.00 12.60 29.40 140.00 30-35 | 13.20 8.25 23.10 107.25 35-40 | 13.30 5.70 17.10 79.80 40-45 | 10.75 2.15 12.90 55.90 45-50 | 7.20 2.40 9.60 40.80 50-55 | 2.65 2.65 7.95 29.15 55-60 0 0 2.90 17.40 235. 25 g. cm.|330. 60 g. cm.|1012. 10 g. cm./2704. 55 g. cm.|2011. 70 g. cm. CLEMENS: LIMNOLOGY OF LAKE NIPIGON 25 TABLE 13 DIRECT WORK—STATION 3 Depth. m. June 17 July 9 August 2 August 29 0-5 47.25 232.50 154. 50 177.75 5-10 50. 00 200. 00 396. 00 428.00 10-15 58.50 65.00 571.00 500. 50 15-20 40.50 36.00 594. 00 252.00 20-25 17.25 25.30 483.00 80. 20 25-30 11.20 9.80 294.00 60.20 30-35 9.95 3.30 161.70 49.50 35-40 7.60 1.90 89.30 43.70 40-45 4.30 0 43.00 38.70 45-50 0 0 19. 20 38.40 50-55 10.60 34.45 55-60 8.70 31.90 60-65 9.45 28.35 65-70 6.80 27.20 70-75 7.30 25.55 75-80 3.90 19.50 80-85 4.15 16.60 85-90 4.40 13.20 90-95 4.65 13.95 95-100 0 9.80 100-105 5.15 105-110 5.40 110-115 0 115-120 0 120-123.4 0 246.55 g.cm.| 573.80 g. cm. | 2865.65 g. cm. | 1900.00 g. cm. (3) that during the storm of July 30, the expenditure took place chiefly in the 10-20 metre area of the open portion of the lake, but in the 10-15 metre area of the more or less sheltered bay. In both cases considerable work was involved in distributing heat in the hypolimnion. It would appear that during the storm probably about 10,000 calories per sq. cm. of surface were distributed in the open waters of the lake requiring about 1500 g. cm. of work. 26 CLEMENS: LIMNOLOGY OF LAKE NIPIGON (4) that a considerable amount of work was expended in the deeper waters of the hypolimnion during August, since these waters continued to gain heat for some time following the storm. . In approximate terms then it may be said that in 1921, 2500 gram-centimetres of work per square centimetre of area were needed to distribute 30,000 gram-calories of heat per square centimetre of area in the waters of Lake Nipigon. This means, then, that our northern waters are essentially similar to those of more southerly distribution in the matter of summer heat income and its distribution. Where they differ, is in the length of the period from the time when the temperature begins to rise above 4°C. until a uniform tempera- ture of 4°C. is again reached in the autumn. This summer period is of short duration. For example, ice may appear in Lake Nipigon in November and the lake be entirely frozen over by the end of December. The lake may not be free of ice again until the middle of May and occasionally not until early in June. In contrast to this, Cayuga Lake rarely freezes over and when it does so, the period lasts but little over a month (Birge and Juday, 1914). Strong summer winds would therefore appear to be of special importance in the northern regions. It is probable that in a season of little wind the summer heat income for Lake Nipigon would be very small. It is evident that the summer or growing period for the organisms in Lake Nipigon is comparatively short and it is hoped that data in regard to the effect of this condition in causing a slow rate of growth for fish may be presented later. However, it does not appear that the productiveness of such a lake in respect of those fish which can tolerate a short summer period, such as the common white- fish, will be any less than that of a lake with a longer summer period, if other factors such as those of food, oxygen supply, spawning conditions, etc., are equal. That the short summer period is one of the factors contributing to the absence of certain species of fish from our northern waters is probable, in that doubtless some species are unable, in the short time available, to accumulate sufficient reserve material to carry CLEMENS: LIMNOLOGY OF LAKE NIPIGON 2h them through the long winter period and allow them to spawn successfully in the spring. Colour and Transparency A white wooden disk 20 cm. in diameter was used in the determination of the transparency and a U.S. Geological Survey standard colorimeter for the colour. The following records were obtained by Mr. George Geiger: Trans- No. Date Time Location parency Colour feet bh" Sept it > 10a.m. off Cedar Island | Zeca tna.) LOva.m. east of Shakespeare Island 1 Ear fawed 69) aubhee: 1a) 3.50 p.m. 2 mi. south of Echo Rock T a eas) 2. D.101, Sturgeon river 12.637 eet aL alin. Gull river 61 Bo eel iy 42 noon Gull Bay 31 Teaeete ol pm, Gull Bay 31 8 ‘“ 23 11.45 a.m. Open waters south of Cedar Is. L/iek | Saat f In No. 2, the water seemed to be slightly turbid with clay. In determining the colours the standard tube was used empty since no distilled water was available. Many other tests in the open parts of the lake gave colour values of about 7. The transparency of the open waters is thus high, com- paring closely with those of Cayuga Lake (Birge and Juday, loc. cit.). The colour is slight in the open waters but is very dark brown in all of the streams tested and observed and persists to some extent in the shore waters of the lake, particularly around the mouths of the streams. This dark brown colour, no doubt, has some relation to the dark colouration of many shore-water fish such as the pike perch, lake trout and some of the whitefish. 28 CLEMENS: LIMNOLOGY OF LAKE NIPIGON Dissolved Gases The results of the oxygen determinations (Tables 1, 2 and 3, and Figs. 4 and 5), for this season show: (1) that from the middle of June to the middle of August 5-6 58 60 62 64 66 68 70 72 74 76 7880 8&2 84 Yd Suneeeeaeaadaccuss (eddeuseecsecesuesseeescceeedcnrccestecccccccTsEeeee Pepa acess cossceccssssesses Fic. 4. Curves illustrating amounts of dissolved oxygen in cc. per litre, Station 3, Lake Nipigon. CLEMENS: LIMNOLOGY OF LAKE NIPIGON 29 60 70 80 20 100 110% Yd. M. OS OGSSESeR ORBRESeas as ireedt GURRRSRRE S57 288 H+ § COC etT EEE TT ATT po eS aes S88 Seeen-.'/// A- anes Perri ee ee a8 Eee =a58 nal PPA = seer auscetveevasrs aoe EEE 10 NEA CE CEE EEE EEE EEE EEE EE CEE EEE EEE Fic. 5. Curves illustrating percentages of saturation of dissolved oxygen, Station 3, Lake Nipigon. 30 CLEMENS: LIMNOLOGY OF LAKE NIPIGON there was a plentiful supply of dissolved oxygen, the content seldom going below about 85% of saturation. (2) that toward the end of August, the waters of the hypo- limnion showed a marked reduction in oxygen content, the bottom waters at a depth of 100 yards having but 62% of saturation. (3) that it is unlikely that the oxygen supply in the deep waters ever nears depletion and that in respect of dissolved oxygen, the lake is well suited to a deep-water fish fauna. The results of the free carbon dioxide determinations (Tables 1, 2 and 3) show:— (1) that at no time during the summer months was the amount of carbon dioxide large at any depth. (2) that in the early part of the summer the upper waters contained the larger amounts but as the season advanced this condition was reversed, doubtless correlated with the growth of phytoplankton in the upper waters and the increase in decomposition processes in the bottom waters. Conclusion In conclusion it may be said (1) that Lake Nipigon pos- sesses those features characteristic of large open bodies of water and does not appear to present any striking peculiar- ities in respect of physico-chemical conditions thus far investigated, except in regard to the short summer period; (2) that the peculiarities of its fauna and flora are the result of, at least the following three outstanding factors, (a) the short summer period, (b) the limited amount of shallow, protected water areas, (c) the isolation from the Great Lakes because of the falls and rapids in the single outlet. Literature Cited Bell, Robert, 1869. Report on the Geology of the North- east Side of Lake Superior and of the Nipigon District. Report for 1866-1869, Geological Survey Canada, pp. 313-364. Birge, E. A., and Juday, C., 1914. A Limnological Study of the Finger Lakes of New York. Bull. U.S. Bureau Fish., Vol. XXXII; document No. 791. CLEMENS: LIMNOLOGY OF LAKE NIPIGON al Birge, E. A., 1916. The Heat Budgets of American and European Lakes. Trans. Wisc. Acad. Sc., Arts and Letters, Vol. XVIII, p. 166. Birge, E. A., 1916. The Work of the Wind in Warming aeieake. Irans. Wisc. Acad. Sc., Arts and Letters, Vol. XVIII, p. 341. Birge, E. A., and Juday, C., 1921. Further Limnolo- gical Observations on the Finger Lakes of New York. Bull. U.S. Bureau Fish., Vol. XX XVII; document No. 905. Birge, E. A., 1922. A Second Report on Limnological Apparatus. Trans. Wisc. Acad. Sc., Arts and Letters, Vol. XX, p. 533. Coleman, A. P., 1922. Glacial and Pre-Glacial Lakes in Ontario. University of Toronto Studies: Publications of the Ontario Fisheries Research Laboratory, X. Department of Game and Fisheries of Ontario, 1916-1920. Annual Reports, Toronto. Hydro-Electric Power Commission of the Province of Ontario, 1907. Fifth Report, p. 14. Toronto. McInnes, William, 1894. Summary Report on the Sur- vey of Lake Nipigon. Geological Survey, Canada, New Series, Vol. VII, Part A, pp. 48-51. Provincial Board of Health of Ontario, 1920. Sewage and Water. Bull. No. 7, Div. Sanitary Engineering. White, James, 1915. Altitudes in the Dominion of Canada. Commission of Conservation, Canada, Ottawa. Wilson, Alfred W. G., 1910. Geology of the Nipigon Basin, Ontario. Memoir No. 1, Dept. Mines, Geol. Survey Br., Canada, pp. 1-152. UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 12 Pear sONaAlL LIST OF THE FISHES OF LAKE NIPIGON BY. JoHn R. DyMonD OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1923 A PROVISIONAL LIST OF THE FISHES OF LAKE NIPIGON The following brief account of the fishes of Lake Nipigon is based upon material obtained by the first field party of 1921. It contains a list of the species so far identified, with some reference to undetermined varieties, and is preliminary to a more comprehensive study and report, the material for which will, it is expected, be obtained during the summer of 1922. The most striking feature of the fish fauna of Lake Nipigon, in view of its size, is the small number of species which inhabit it. Asa result of the first season’s work there fewer than thirty species were found in the waters of the lake itself. This is in striking contrast to what is found in other large lakes. Dymond (1922) gives ninety-one species of fish and two species of lamprey as occurring in Lake Erie, while Bensley (1915) records forty-eight species from Georgian Bay. The small number of species occurring in Lake Nipigon is no doubt due in part to the physical conditions in the lake and in part to its geological history. Its waters have been isolated for long ages from the waters of the lower lakes, by means of falls sufficiently high to offer an insurmountable obstacle to the ascent of species from the south. Some species found in neighbouring lakes are absent from Lake Nipigon, which probably means that conditions in the lake are unsuitable for these forms. The Sturgeon, Acipenser rubicundus Le Sueur, is fairly abundant. During 1919, 17,595 pounds were taken from the lake. The largest specimen recorded weighed 99 pounds. The Long-nosed or Northern Sucker, Catostomus catos- tomus (Forster), and the Common Sucker, Catostomus com- 35 36 DyMoND: THE FISHES OF LAKE NIPIGON mersoniit (Lacépéde), are very abundant. The former, especially, is taken in considerable numbers in deep waters in gill nets set for whitefish and lake trout. The Common Sucker is found in shallower water. On account of the dist- ance from market it is not profitable to ship these species, and those taken in the nets are consequently destroyed. Only four species of Cyprinidae were found in the lake. In the order of their relative abundance they are, the Spot- tailed Minnow, Notropis hudsonius (De Witt Clinton), Lake Chub, Couesius plumbeus (Agassiz), Long-nosed Dace, Rhini- chthys cataracte (Cuvier and Valenciennes), and the Shiner, Notropis atherinoides Rafinesque. None of them appears to be at all common, and no specimens of any of them were identified in the stomachs of fishes. Leuciscus neogeus (Cope) is found in a number of small lakes in the neighbour- hood, but has not been taken from Lake Nipigon itself. This species has not been previously recorded from Ontario. Speckled Trout, Salvelinus fontinalis (Mitchill), of unusual size are taken on the line in the Nipigon river. The largest one taken in recent years weighed 143 pounds, but specimens of five pounds in weight are quite commonly secured. They are also found in the fishermen’s pound nets, but fish so taken are returned to the water. The Lake Trout, Cristivomer namaycush (Walbaum), is, next to the whitefish, the most important commercial species of Lake Nipigon. It is usually taken in gill-nets in deep water, but a very dark, slim form is commonly taken in pound nets in shallow water. The significance of the latter variety is to be further investigated. The Round Whitefish, Coregonus quadrilateralis Rich- ardson, isnot uncommon. It appears to remain in compara- tively shallow water. On account of its slimness, it is not taken in the fishermen’s nets. The Common Whitefish, Coregonus clupeaformis (Mitchill), is the most important commercial species in the lake. The fishermen recognize two types, viz., the deep-water and the shallow-water forms. In general, the latter is a darker, slimmer fish than the former, although both types are taken DyYMOND: THE FISHES OF LAKE NIPIGON 37 in deep water. The two extremes are connected by inter- gradations. The variation in colour, form, and other char- acters dependent on depth and other factors has not yet been fully investigated. The Ciscoes or Lake Herrings, Leucichthys spp., have not yet been definitely worked out, but there appear to be at least three varieties. The commonest form is a dark fish with black fins. It averages about 1} pounds in weight. A larger, lighter-coloured form, locally known as ‘‘Tullibee’’, is occasionally taken in the gill-nets in deep water. A much smaller variety with longer head and very pale fins appears to be confined to comparatively shallow water. The Pike, Lucius lucius (Linn.), is common in small bays along shore, and is occasionally taken in the fishermen’s nets. The Brook Stickleback, Eucalia inconstans (Kirtland), is much less common than the nine-spined Stickleback, Pygo- steus pungitius (Linn.). The latter is perhaps the most abundant of the smaller species inhabiting the lake, some specimens being taken in nearly all seine catches. It was also commonly found in the stomachs of Pike Perch, and less commonly in Lake Trout, Ling, Pike, and Whitefish. The Trout Perch, Percopsis omisco maycus (Walbaum), appears to be common. A few specimens were taken from the stomachs of predaceous species, and thousands were taken in July in a specially constructed bag net set in a stream a few hundred yards above where it enters the lake. The Small-mouthed Black Bass, Micropterus dolomieu Lacépéde, is believed not to occur naturally in Lake Nipi- gon. Small specimens of the year were taken in the seine near the foot of Orient Bay in July. They were probably the progeny of some parent fish planted the previous season. The Yellow Pickerel or Pike Perch, Stizostedion vitreum (Mitchill), is an important commercial species in Lake Nipigon. It is taken in pound nets in shallow water. The maximum size attained is about nine pounds. The Yellow Perch, Perca flavescens (Mitchill), was taken in considerable numbers in small bays by means of the seine. 38 Dymonpb: THE FISHES OF LAKE NIPIGON For some reason the species does not reach a very large size, the largest specimen secured being six inches in length. The only species of darter taken was the Tessellated Darter, Boleosoma nigrum (Rafinesque.) It was commonly taken in the seine in shallow water. A number of specimens of the Miller’s Thumb, Cottus ictalops (Rafinesque), and of Uranidea gracilis (Heckel), were taken about the middle of June in a specially constructed bag net set in a stream a short distance from where it enters the lake. Partly digested specimens, too fragmentary for definite determination, taken from the stomachs of ling may represent a third species of the family Cottidae. The Ling, Lota maculosa (Le Sueur), is very common in deep water, and large numbers are taken in gill nets set for Whitefish. Although this list is not extensive, it includes the principal ' commercial species characteristic of the Great Lakes. Its deficiency is mainly in Cyprinidae and other small forms. To what extent this deficiency limits the productivity of the lake by curtailing the food of the larger predaceous species cannot be estimated until other bodies of water, rich in the smaller species, have been investigated. Literature Cited Bensley, B. A., 1915. The Fishes of Georgian Bay. Contrib. Canad. Biol. 1911-14, Fasc. II, Fresh Water Fish and Lake Biology. Biological Board of Canada, Suppl. 47th Ann. Rep. Dept. Marine and Fisheries, Fisheries Branch, Ottawa. Dymond, J. R., 1922. A Provisional List of the Fishes of Lake Erie. University of Toronto Studies. Biological Series. Pub. Ontario Fisheries Research Laboratory, No. 4. UNIVERSITY OF TORONTO STUDIES PUBLICATIONS OF THE ONTARIO FISHERIES RESEARCH LABORATORY No. 13 THE PLANKTON OF LAKE NIPIGON AND ENVIRONS BY N. K. BIGELOW OF THE DEPARTMENT OF BIOLOGY UNIVERSITY OF TORONTO TORONTO THE UNIVERSITY LIBRARY 1923 THE PLANKTON OF LAKE NIPIGON AND ENVIRONS The following preliminary record of the more minute fauna and flora of Lake Nipigon and vicinity is the result of extensive collections made during the summer of 1921. The material studied was obtained from four sources :— (1) By means of a vertical closing net, plankton samples were taken from the open waters at all depths. Three stations were established, and visited regularly. Station I was located off the mouth of the Nipigon River; Station II, in the deep water of Pijitawabik Bay (Orient Bay), directly opposite the village of Macdiarmid; Station III, in the open waters opposite the mouth of Sandy River. The usual type of closing net was used, made of bolting silk, size No. 20, and vertical tows were made through five and ten yard intervals. The quantitative results will be presented in a later paper. (2) Surface tows were made in many places in the lake and in the lower portions of some of the larger streams. (3) Many interesting forms were secured by sweeping with a small, conical net among aquatic vegetation in streams, bayous, ponds, pools, cold springs, etc. (4) The stomach contents of many species of fish yielded much valuable data. Young fish, particularly the young of the common sucker (Catostomus commersonii) were found to be excellent collectors of small organisms. Several hundred small suckers were examined, and their digestive tracts were found each to contain from 30 to 50 or more different species of microscopic animals and plants. When from two to four centimetres in length, they were found to have subsisted almost entirely upon plankton. Slightly larger specimens taken a little later in the season were found to have fed upon the surface of the bottom ooze, where myr- iads of Rotatoria, Cladocera, Diatoms, etc., occurred. 41 42 BIGELOW: THE PLANKTON OF LAKE NIPIGON In the lists of organisms mentioned in this paper the following designations are used: A.=abundant; V.C.= very common; C.=common; F.=fairly common; I. =infre- quent; R. =rare. An asterisk after the name of an organism denotes that the species had been taken as food by the young suckers previously mentioned. ALGAE CLAass MYXOPHYCEAE The abundance of blue green Algae in Lake Nipigon and its vicinity was found to be insignificant in comparison with that of the diatoms; nevertheless, a considerable number of species were found to occur. The only genera found in the open-water plankton were Microcystis, Anabaena, and Aphanizomenon. The other genera occurred in creeks, bayous, ponds, and pools. ORDER COCCOGONEALES FAMILY CHROOCOCCACEAE Chroococcus limneticus Lemmermann I. Chroococcus turgidus Nageli * F. Coelosphaerium sp. I. Merismopedia elegans Braun * F. Microcystis aeruginosa Kiitzing * A. Although this species was not present in sufficient numbers to be the direct cause of lake bloom, it contributed to this formation by mingling with the more abundant Anabaena. Microcystis flos-aquae Kirchner * V.C. Aphanocapsa sp. * I. A phanothece sp. I. Gloeothece sp. I. Dactylococcopsts sp. * I. ORDER HORMOGONEALES FAMILY OSCILLATORIACEAE Lyngbya birget Smith * I. Oscillatoria sp. F. Myxophytes of this genus were found only in small pools. BIGELOW: THE PLANKTON OF LAKE NIPIGON 43 FAMILY NOSTOCACEAE Nostoc sp. Pale green colonies of a species of this alga were found attached to weeds in a sluggish creek near Orient Bay. The colonies were spherical, but only about 8 mm. in diameter. Anabaena circinalis Rabenhorst A. Anabaena lemmermanni Richter * A. Long strips of lake bloom in the latter part of July and in August were found, on microscopic examination, to be masses of spores of this species, from which the surrounding filaments had disin- tegrated. Masses of the same kind of spores were found occasionally in the digestive tracts of young suckers. Aphanizomenon flos-aquae Ralfs. R. FAMILY RIVULARIACEAE Rivularia sp. F. CLAss BACILLARIACEAE Of all the minute organisms found in Lake Nipigon diatoms were by far the most numerous. They occurred in prodigious numbers in the plankton, being often so very numerous that other organisms had to be specially searched for. They were also very abundant in the ooze and among aquatic vegetation. A large percentage of the food of bottom-feeding fish was found to be diatomaceous. The genera of diatoms found in open-water plankton in order of greatest abundance were Melosira, Asterionella, Tabellaria, Synedra, Stephanodiscus, and Fragillaria. FAMILY MELOSIRACEAE Melosira granulata Ehr. * A. During the month of June this diatom was so abundant that, if the net were towed behind a motor-boat at Station I or Station II for five minutes, it would contain at least half-a-pint of greenish- coloured plankton, which was found to be composed of about 95% of this particular organism. Quantitative methods of counting sometimes showed as many as 145,000 filaments 44 BIGELOW: THE PLANKTON OF LAKE NIPIGON per cubic centimetre. By the middle of July, Asterionella was much commoner on the surface, but Melosiva was still quite common in the deeper water. In the latter part of August, Melosira had become a rather uncommon species. FAMILY COSCINODISCACEAE Stephanodiscus sp. * C. FAMILY RHIZOSOLENIDACEAE Rhizosolenia sp. F. A diatom of this genus was fairly common in the plankton of Station II during the month of June. The species was very slender, and possessed very long indistinct terminal spines. In all probability it was a much commoner organism than appearances would indi- cate as its extreme transparency rendered it difficult to see. FAMILY NAVICULACEAE Pleurosigma sp. * I. Pinnularia sp. * V.C. Navicula sp. * V.C. Stauronets sp. * F. FAMILY CYMBELLACEAE Amphora sp. * I. Cymbella sp. * C. Cocconema sp..* V.C. Encyonema sp. * R. FAMILY GOMPHONEMIACEAE Gomphonema sp. * C FAMILY ACHNANTHACEAE Achnanthes sp. * C. FAMILY NITZSCHIACEAE Nitzschta sp. F. FAMILY AMPHIPRORACEAE Amphtiprora sp. * I. BIGELOW: THE PLANKTON OF LAKE NIPIGON 45 FAMILY SURIRELLACEAE Campylodiscus sp. * I. Cymatopleura sp. * C. Surirella sp. * V.C. FAMILY DIATOMACEAE Denticula sp. R. FAMILY MERIDIONACEAE Meridion circulare (Gren) C. Large brown masses of this species were found in a cold spring near Fairlock. FAMILY FRAGILLARIACEAE Synedra sp. * V.C. Fragillaria sp. * C. Asterionella formosa Hass. A. Next to Melosira this was the most abundant organism of the open-water plankton. It appeared to be commonest after Melosira had declined in numbers, but this may have been due, somewhat, to the fact that the thinning out of the latter rendered it more conspicuous. Plankton composed of Asterionella is very white in contrast to the green Melosira. FAMILY TABELLARIACEAE Tabellaria fenestrata Kiitzing * A. This was the third commonest diatom. It reached its culminating numbers in late summer, when Asterionella had greatly decreased and Melosira was scarce. Plankton composed of this species was white, but not so purely white as that composed of Astertonella. At the end of August the base of the stems of rushes standing in shallow water near the head of Orient Bay were found to be encased in a coating of whitish, slimy material which was found to be entirely composed of this Tabellaria. Apparently the long zigzag filaments had be- come tangled together and, being washed by the waves, had collected around the stems of the rushes. Several young suckers taken near this locality at this time had been feeding almost entirely upon this material as the contents of their 46 BIGELOW: THE PLANKTON .OF LAKE NIPIGON alimentary tracts averaged from 95% to 98% T. fenestrata. Tabellaria flocculosa Kiitzing * V.C. FAMILY EPITHEMIACEAE Epithemta sp. * V.C. Ceratoneis arcus Kiitzing F. One plankton tow from Sandy Bay early in June contained fair numbers of this species; otherwise it was scarce. CLAss HETEROKONTEAE ORDER HETEROCOCCALES FAMILY OPHIOCYTIACEAE Ophiocytium capitatum Wolle C. Masses of filamentous algae in a sluggish creek near the head of Orient Bay were found to contain numbers of this species. Ophiocytium arbuscula (Braun) C. This species was - found at the same time and place as the preceding. FAMILY BOTRYOCOCCACEAE Boitryococcus braunii Kiitzing * A. The only other organ- ism responsible for lake bloom besides Anabaena lemmermanni was this Botryococcus. It produced long strips of reddish colour in the latter part of July on the surface of the lake. ORDER HETEROTRICHALES FAMILY TRIBONEMACEAE Tribonema minor (Wille) V.C. This alga was found only in tiny pools along the railway in June. CLASS CHLOROPHYCEAE ORDER VOLVOCALES FAMILY VOLVOCACEAE Pandorina morum Bory F. Eudorina elegans Ehr. C. Volvox aureus Ehr. C. BIGELOW: THE PLANKTON OF LAKE NIPIGON 47 ORDER PROTOCOCCALES FAMILY PALMELLACEAE Sphaerocystis sp. F. Tetraspora lacustris Lemmermann I. Tetraspora sp. F. Long cylindrical masses of an alga of this genus occurred in pools along the railway near Mac- diarmid in the month of June. These colonies were about 2 centimetres in length by 1/5 as wide. FAMILY DICTYOSPHAERIACEAE Dictyosphaerium sp. FAMILY AUTOSPORACEAE Oocystis sp. * V.C. Tetraedron limneticum Birge R. Ankistrodesmus falcatus Ralfs. R. Quadrigula sp. R. Kurchneriella lunaris Mobius * I. Scenedesmus bijuga Lagerheim * I. Coelastrum cambricum Archer * I. Coelastrum proboscidium Bohlin R. Sorastrum americanum Bohlin I. Sorastrum spinulosum Nageli I. Actinastrum hantzschi Lagerheim F. FAMILY HyYDRODICTYACEAE Pediastrum duplex Meyen * V.C. Pediastrum boryanum Turpin * V.C. Pediastrum tetras Ehr. I. Pediastrum biradiatum Meyen R. None of the Algae belonging to the orders Volvoccales or Protococcales were very abundant members of the flora of the Nipigon region. The only plankton in which they were found was that of bays and creeks. ORDER ULOTRICHALES FAMILY ULOTRICHACEAE Ulothrix zonata Kiitzing C. Stones in shallow water near the margin of the lake were covered with a brown matting composed of this alga. 48 BIGELOW: THE PLANKTON OF LAKE NIPIGON FAMILY CHAETOPHORACEAE Chaetophora pisiformis Ag. C. Beautiful green masses of this species were attached to sticks and stones in a sluggish creek. Coleochaete sp. F. The flattened colonies of this alga were found on the under surface of water-lily leaves. ORDER SIPHONOCLADIALES FAMILY CLADOPHORACEAE Cladophora sp. A. The bottom of the lake for some dis- tance from the margin was found often carpeted with a species of this plant. It is rather important as food for sturgeon, whitefish, and other bottom-feeding fish. ORDER CONJUGALES FAMILY ZYGNEMACEAE Spirogyra sp. V.C. Several species of this alga occurred in ponds, creeks, and along the margin of the lake. Zygnaema stellinum Ag. C. This species was common in sluggish creeks near Orient Bay. Mougeotia viridis (Kiitzing) * A. This was the common- est species of the Conjugales, and was found in ponds, streams, and creeks, as well as at the margin of the lake. FAMILY DESMIDIACEAE Desmids occurred in considerable numbers in plankton swept from among weeds in sluggish creeks and bayous of the lake. They were especially numerous near the head of Orient Bay where the water is rather shallow. Weedy bayous occur along the bank and several sluggish creeks flow into the lake at this point, forming ideal conditions for their growth. Although many species of desmids were found they were never abundant enough to be an important constituent of the plankton. The following list contains only the recognized forms, besides which a great number of others occurred: Penium sp. * I. Closterium lunula Ehr. * I. BIGELOW: THE PLANKTON OF LAKE NIPIGON 49 Clostertum lineatum Ehr. C. Clostertum rostratum Ehr. V.C. Docidium baculum (Breb.) * C. Staurastrum coronulatum Wolle * I. Staurastrum orbiculare Ralfs 1. Micrasterias rotata Ralfs * V.C. Micrasterias furcata Ralfs F. Micrasterias truncata Ralfs F. Micrasterias americana Kutz F. Cosmarium octhodes Nord. * V.C. Sphaerozosma pulcrum Bailey I. Hyalotheca sp. * F. Desmidium swartzit Ag. * V.C. A ptogonum baileyi Ralfs * R. Gymnozyga sp. R. PROTOZOA CLASS RHIZOPODA As might be expected these organisms were found mainly in the ooze and in the digestive tracts of bottom-haunting fish which had fed upon this material. With one or two exceptions none were found in open-water plankton except an occasional adventitive individual ORDER GYMNAMOEBIDA FAMILY AMOEBIDAE Amoeba proteus Leidy F. ORDER TESTACEA FAMILY ARCELLIDAE Arcella vulgaris Ehr. * F. Arcella dentata Ehr. * R. Centropyxis aculeata Stein * V.C. Lecquereusia modesta Rhumbler * C. Difflugia corona Wallich * V.C. Diffiugia lobostoma Leidy * V.C. Although Dzfflugra is usually considered to be a bottom-haunting organism, one small round form, answering in every way the description of 50 BIGELOW: THE PLANKTON OF LAKE NIPIGON D. lobostoma, appeared in such numbers in the open-water plankton as to preclude the possibility of its being adventi- tious there. It was found most commonly on the surface, but occurred at all depths. Difflugia pyrtformis Perty * V.C. Difflugia acuminata Ehr. * V.C. Nebela sp. * R. ORDER TESTACEA FAMILY EUGLYPHIDAE Campascus ?* F. The test of an undetermined Sarco- dinid protozoan was occasionally taken as food by small suckers. This rhizopod resembles Campascus, but the processes of the shell are closer together and more terminal than in Campascus cornutus. Cyphoderia ampulla Ehr. * V.C. Cyphoderia ampulla var. papillata Wailes * I. Assulina seminulum Ehr. * F. Euglypha alveolata Dujardin * C. In all probability several species of Euglypha occurred. The list of rhizopods would probably be greatly extended if sphagnum bogs and other likely spots had been specially searched for them. CLASS ACTINOPODA ORDER APHROTHRORACIDA Actinophrys sol Ehr. F. ORDER CHALATHORACA Pompholyxophrys punicea Archer I. CLASS ZOOMASTIGOPHORA ORDER EUGLENIDA FAMILY EUGLENIDAE Trachelomonas hispida Stein I. Phacus longicaudus Ehr. I. BIGELOW: THE PLANKTON OF LAKE NIPIGON 51 CLASS PHYTOMASTIGOPHORA ORDER CHRYSOFLAGELLIDA Mallomonas sp. F. These organisms were commonest in a plankton tow, not on the surface, but at a depth of about 8 metres, near North Bay on July 19. Spongomonas sp. F. Long, slender, yellowish-brown colonies, often twisted and irregular, were found among weeds in a sluggish creek near Orient Bay during the latter part of August. Rhipidodendron splendidum Stein F. Small masses of the tubules of this flagellate were found at the same time and place as the colonies of the preceding species. Synura uvella Ehr. F. Colonies of these protozoa were fairly common in ponds and pools. Dinobryon sertularia Ehr. A. This species was sometimes abundant during the month of June in plankton from bays and creeks. Dinobryon bavaricum Imhof. F. Very typical example of this species occurred during the latter part of June in plankton from Station II. It was always found with the preceding species, but never was so abundant. Peridinium sp. F. Ceratitum hirundinella Miller A. This species was oc- casionally abundant during the month of July in plankton from bays and creeks. CLass CILIATA The ciliates included in the following list were mostly found in ponds and pools. Codonella is a typical open-water planktont and was found at all depths. Vorticellae were attached to colonies of Anabaena in the lake. ORDER HOLOTRICHA Coleps hirtus Ehr. C. Didinium nasutum Miiller F. Nassula ornata Ehr. R. Trachelius ovum Ehr. F. 52 BIGELOW: THE PLANKTON OF LAKE NIPIGON Urocentrum turbo Miiller C. Ophryoglena atra Ehr. I. Colpidium sp. F. Cyclidium sp. F. Paramoecium caudatum Ehr. C. ORDER HETEROTRICHA Stentor coeruleus Ehr. C. Halteria grandinella Miiller C. Codonella sp. V.C. OrDER HyYPOTRICHA Oxytricha sp. C. ORDER PEROTRICHA Vorticella sp. V.C. A number of species occurred. Ophrydium eichhornit Ehr. I. Two beautiful green col- onies of this ciliate were found attached to vegetation in - a creek. Pyxicola sp. I. CLASS SUCTORIA Podophrya sp. F. A pear-shaped species of this genus, with a long stalk, was often found attached to the legs and antennae of Limnocalanus macrurus and Mysis_ relicta. The crustaceans were often taken from a depth of 60 metres or more. ROTATORIA Although a great many species of the wheel animalcules were found, no one species was conspicuously abundant. Forming, as they do, a large percentage of the first food of young fishes, their economic importance is considerable. ORDER PLOIMA FAMILY NOTOMATIDAE Eosphora sp. F. In pools along the railroad track. Notomata aurita Miiller I. In weedy creeks. Diaschiza 1. Several sp. In weedy creeks. Cephalodella forficula Ehr. I. In weedy creeks. Monommata orbis Ehr. 1. In weedy creeks. BIGELOW: THE PLANKTON OF LAKE NIPIGON 53 FAMILY BRACHIONIDAE Brachionus capsiflorus Pallas R. Only one specimen of this species and genus was found during the season. It was a typical example of the form previously known as Brach- tonus bakeri with long, gracefully curved cephalic and pos- terior spines. It was found in a small pond close to Mac- diarmid village. FAMILY BRACHIONIDAE Platyias quadricornis Ehr. C. This rotifer was found throughout the summer among weeds and algae in small pools and sluggish creeks, but never in the lake. Keratella cochlearis Gosse * V.C. This proved to be the most widely distributed species in the region under discussion, and was found in all bodies of water examined except in temporary pools. In the lake it occurred at all depths from the surface down to a hundred metres throughout the season. Keratella quadrata Miiller F. This was an open-water form and was found from the surface down to a depth of 100 metres. It was by no means as common as the preceding species, three individuals being the greatest number noted in any one plankton haul. The rotifer was taken from the latter part of July until the end of August. The individuals had very large posterior spines, broad at their bases, and widely divergent. - Keratella serrulata Ehr.* 1. This species was found only in creeks and bays. The specimens showed the typical, prominent hexagons on the lorica and roughened projections on the egg carried, as well as on the lorica. Anuraeopsis fissa Gosse F. During the month of June this species occurred in a small pond close to Macdiarmid village. Notholca longispina Kellicott * V.C. Next to Keratella cochlearis, this was the commonest rotifer. It occurred throughout the summer at all depths from the surface down to 100 metres. It was also found in creeks and some of the larger ponds. 54 BIGELOW: THE PLANKTON OF LAKE NIPIGON Notholca striata Miiller * C. Although found occasionally in open-water plankton this species was commonest in bays and creeks. Nearly all small suckers taken during July and August near Orient Bay had eaten it. Besides typical Notholca striata, the forms of this species which were pre- viously designated as Notholca thalassia and Notholca acu- minata were frequently found. Notholca foliacea Ehr.* I. This species was taken a few times in plankton from Station II. A few were eaten by suckers in July and August. FAMILY MYTILINIDAE Mytilina mucronata Miller * C. This species was found in weedy creeks throughout the season. FAMILY EUCHLANIDAE Euchlanis deflexa Gosse * F. Euchlanis dilatata Ehr. I. Diplois propatula Gosse F. Lecane ohioensis (Herrick) * C. Lecane sulcata Gosse * C. Lecane luna (Miiller) * V.C. Lecane leontina (Turner) I. Monostyla bulla Gosse * C. Monostyla lunaris Ehr. * F. Monostyla quadridentata Ehr. R. Members of this family and the next are found in bays and creeks among weeds and are frequently fed upon by young fish. FAMILY LEPADELLIDAE Lepadella ovalis Miiller * F. Lepadella acuminata Ehr. * F. Lepadella ehrenbergit (Perte) R. A single individual was taken from the digestive tract of a young sucker. Colurella adriatica Ehr. * F. Colurella uncinatus (Miller) * I. Squatinella longispinatum Tatem I. This species was found in plankton swept from among weeds in the Pustagone BIGELOW: THE PLANKTON OF LAKE NIPIGON 55 River on August 23. A peculiar Squatinella was also found in this material which resembled a typical S. longis- pinatum except that it had a second long dorsal spine on the lorica some distance back of the first. This may be only a variation, as several quite normal individuals of this species occurred in the same material. FAMILY TRICHOTRIIDAE Trichotria pocillum Miiller * F. Macrochaetus collinsit (Gosse) I. Eight or nine individ- uals of this strange rotifer were taken from creeks and one from some plankton in the bay near Orient Bay village. The only other examples of this rotifer ever seen by the writer were from an entirely different environment, namely, in a small muddy pond in a cow-pasture near Cedar Rapids, Iowa. Scaridium longicaudum (Miiller) F. This rotifer was found among weeds in creeks. Scaridium eudactylotum Gosse R. This rare rotifer was found but once in plankton from among weeds near the mouth of the Pustagone River. Diurella tenuior (Gosse) * C. Diurella stylata Eyferth * I. Trichocerca cristata Harring F. Trichocerca cylindrica (Imhof) I. Trichocerca lata Jennings * C. Trichocerca longiseta Schrank F. Trichocerca multicrinis (Kellicott) I. Several rotifers of this family not satisfactorily determined are not listed. FAMILY CHROMOGASTRIDAE Chromogaster ovalis (Bergendae) C. In middle and late summer this rotifer was rather common in open-water plank- ton, but was never found at any great depth. FAMILY GASTROPODIDAE Gastropus stylifer Imhof * V.C. This wasa very common surface form in all parts of the lake throughout the season. It was also found in creeks and large ponds. 56 BIGELOW: THE PLANKTON OF LAKE NIPIGON Ascomorpha eucadis Perty R. One rotifer of this species was taken from weeds growing on the bottom of a pond. FAMILY SYNCHAETIDAE Synchaeta stylata Wierzejski V.C. Throughout the season this species was very common in the lake. It was occasion- ally taken at considerable depths, but was much commoner near the surface. When this species was numerous its spiny resting eggs were sometimes common in the plankton. FAMILY POLYARTHRIDAE Polyarthra trigla Ehr.* V.C. Increeks, ponds, and bayous as well as in the open waters of the lake, this species was common tbroughout the season. It was commonest in the latter part of summer. It is mainly a surface form and was rarely taken in deep water. FAMILY PLOESOMIDAE Ploesoma lenticulare (Herrick) * V.C. This also was a surface form occurring tbroughout the summer in all parts of the lake. Ploesoma hudsoni Imhof I. This species was found in similar conditions to the preceding, but was only seen about a dozen times. FAMILY ASPLANCHNIDAE Asplanchna sp. I. A large rotifer of this genus was occasionally found in open-water plankton. The specimens were in too poor a condition when examined to permit of specific determination. FAMILY TESTUDINELLIDAE Testudinella patina Hermann C. This beautiful discoid rotifer was common throughout the summer in weedy ponds and creeks. It was frequently attacked by some parasitic organism which filled its lorica with short cylindrical spores. Often the rotifer would be still alive when its shell had BIGELOW: THE PLANKTON OF LAKE NIPIGON 57 become so filled with these spores as to hide its internal organs completely from view. ORDER FLOSCULARIACEAE FAMILY FLOSCULARIDAE Floscularia ringens (Linn.) I. On August 3 this species was found attached to weeds in South Bay. Limnias melicerta Weisse I. The beautiful annulated tubes of this rotifer were swept from among _ weeds in a small creek near Orient Bay the latter part of August. FAMILY CONOCHILIDAE Conochilus unicornis Rousselet I. Occasionally clusters of this rotifer were taken in open-water plankton. Lacinularia sp. I. Masses of an undetermined species of this genus were attached to weeds in South and Orient Bays. ORDER COLLOTHECACEA FAMILY COLLOTHECIDAE Collotheca algicola (Hudson) I. This rotifer was found in colonies of Rivularia attached to weeds in a creek near Orient Bay. Collotheca ambigua (Hudson) F. This rotifer was found attached to weeds in sluggish streams. Collotheca cornuta (Dobie) I. This species occurred under conditions similar to the above mentioned form. Collotheca mutabilis (Hudson) F. This is a typical open-water planktont, free floating and never attached to weeds. It occurred in many parts of the lake, but always close to the surface. ORDER BDELLOIDA FAMILY ADINETIDAE Adineta sp. F. A species of this genus occurred in creeks and ponds and a similar, if not identical, one was taken from water contained in the leaves of pitcher plants. 58 BIGELOW: THE PLANKTON OF LAKE NIPIGON FAMILY PHILONINIDAE Rotaria rotatoria (Pallas) C. Rotaria citrina Ehr. F. Rotaria neptunia Ebr. R. Philodina roseola Ehr. 1. Dissotrocha aculeata Ehr. I. ARTHROPODA CLass CRUSTACEA ORDER COPEPODA FAMILY CENTROPAGIDAE Epischura lacustris Forbes C. This large copepod was found near the surface in the open water of the lake and in creeks and bays. Although not frequently found in the plankton, its abundance is attested by the numbers found to have been eaten by small fishes. The stomach of a young © small-mouth black bass (Micropterus dolomieu) contained 82 of these copepods, while 67 were found in another individual. These two fishes were 2.6 centimetres in length and were taken on July 19 at Orient Bay. Diaptomus sicilis Forbes V.C. This was probably the commonest species of Diaptomus in Lake Nipigon. It was found in all parts of the lake, often at considerable depths but not in the deepest water. Diaptomus minutus Lilljeborg C. This Diaptomus was not quite so common as the preceding form. It was not found until the middle of July, whereas D. sicilis occurred from the first of June. D. minutus was first found in creeks and shallow water, but as the season advanced it was taken in the surface waters of the more open parts of the lake. Quite often while making a series of vertical plankton hauls through various intervals from the bottom to the surface, D. minutus was found to occur in the first few metres below the surface after which D. sicilis would extend for a few metres more to be succeeded by Limnocalanus. Rather infrequently, late in the summer, both species of Diaptomus occurred together. BIGELOW: THE PLANKTON OF LAKE NIPIGON 59 Limnocalanus macrurus Sars V.C. This copepod was found in numbers in the deeper, cooler waters of the lake. It was common from 40 to 50 metres below the surface. Sometimes over 200 individuals would be taken in a 5-metre haul at these depths. Probably this gives a very erroneous idea of its actual numbers as it is a large active creature which would swim away from the net and not be caught in the same way as the smaller constituents of the plankton, such as diatoms and rotifers. Its numbers must be enormous indeed, judging from the stomach contents of whitefish and ciscoes. Four ciscoes were found to have their alimentary tracts packed with fragments of countless thousands of this crustacean. FAMILY CYCLOPIDAE Cyclops * V.C Specific identifications in this genus were not attempted, although several species doubtless occur. Although often eaten by many kinds of fishes, it never was found in great numbers in any of the stomachs examined. A peculiar dark-coloured, opaque cyclops was taken on August 30 from a small pond near Fairlock. FAMILY HARPACTICIDAE Canthocamptus stapylinus (Jurine) * C. Canthocamptus hiemalis (Pearse) * I. Canthocamptus minutus (Claus.) * F. FAMILY ERGASILIDAE Ergasilus sp. 1. A species of this copepod was found in the stomach of a ling which had been feeding upon the nine-spined stickleback (Pygosteus pungitius). Doubtless the parasite came from the latter fish. ORDER OSTRACODA Ostracods were of very great importance indeed as food for bottom feeding fish, as nearly all examined had eaten at least some of these crustaceans. A very few of the numerous species were identified. 60 BIGELOW: THE PLANKTON OF LAKE NIPIGON Cypria sp. C. 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Func ieee Vi (eve) aterivi ie Lene wi0}10q jo JoyoRieyy 6 6 9 Siretias favre. celrelcervertor a) rdalmeiye) ewtdl et lo/Te (i's Kerce] (er.vie netay, typ WitaO ITED Cg 8 o> Since (3994 ul) yidaq g Z T Peer aCe ae Tt las aa Ti a eat Tt CC Bn MCC eS Ceo Seiad tpi ah ibd) Ssurspoaiq (ANd HLYON) AVA AYALNIOW MOoLLuscaA IN LAKE NIPIGON ADAMSTONE 100 é‘ds ads aj1gvisDa UNjDIJaInNIS wmnjnoadagGnog UNINUIM WnUDLpaUt asuso0yppr —_—,,, UNSSadquMon UNIPISLT UNUDJUOUdaL WNLdaDYds ajrgDrave unjinprpuards unjnrdagnvg .,, UNINUIUM WNIUDIPaW WNIPIStT ” cI 91 OF I i I € ST tet a "tT" * -appaaamyds = a : =: = z a ee eres muon, *. T Zz oe oe G 9 tee a fret acrecnt iG oe *("TeA) DIDUIADIVAY 7 ; ; 1 : 7 = eee emma += pugouss DIDAD A o. g o. 8 GT Sac DeOeChicat yar Oar Tanya aCietia eee ‘ds = 2 : a ae ee + psomay mootuuy T ZI z it . ¢ P see eee SHEE CSS CAME SCH DCY = = 1 = aces eee re eee snpieny oo 2 z 4 Be Sp0l OO ea ireie 0h ue ee “SNSOAJUD SUQAOUD] T Se | a | a cee | eae een ra ae ire ee orm cee | ee “8 8" *STAQDIISS DAD DUuyUuaMsas FI pa ante (eC) | em es Pee EO eaitemensiens pungp3 = T T os heard Pos mGntist? Cacecanere te A ones type DaDUU'T 00T og GB OT O1Z OIT 09 cE OL |'******(-spA) ar0yg wo. dourIsIq POEs — |=PUes = PUES =| PINS || PN | PEN Pn =| PON | Png wW0}}0g JO JoOBIeYD 6 9 7 i 08 4G ST 6 ee es 7" (aay un) yydeq $ g z I Cc P g Z T erie) Me jakee)is- selene. 6—S:'0 0-6-6» oe * 3urspoiq IIXXX 8919S ‘CI YACNVXATV IXXX S212 dVO SOUD = Sem eee aro 95 —= a : ‘ . tine 3 Ladoys pe Sa een ‘Auf uo purs=a/, {yoor=V {Avpo=a4 {puvs=e@ tpnur=-} smojog *PULS dIBOIPUT SOUT] -YOOI oFeoFpUy SJOP -a404 ; “QUIT BIOYS 9Y? 5 eens sh PUL UdHL} OID STUIBpoip IVY SayI[LoO] oy} Surmoys uosidiN oye] jo UOI}AOd UsIISBIYINOS VY} Jo deyy 00.99 22 OR 56.29 4.50 f wa Ho ioe ” Gy a3 r ié 9g ‘Sf 7}?P®?d PAS a7 $944 102 ADAMSTONE: MOLLUSCA IN LAKE NIPIGON In the foregoing series of tables the results of all dredging operations are given. These were supplemented by a large number of shore collections. In all, some 235 samples were obtained with the dredge. Each sample represented an area of 81 square inches, and in the results that area is taken as a unit. The largest number of dredgings were made in water down to 21 feet in depth, a total of 144 having been secured in this range. The greatest depth from which specimens were brought up was 192 ft. In the detailed consideration of the Mollusca, no account is taken of dead specimens, but a very large number of them were obtained. DISTRIBUTION OF SPECIES Study of the data obtained makes it evident that, as regards distribution, the species of Mollusca form two distinct groups: 1. Species inhabiting very shallow water close to shore. 2. Species inhabiting deeper water at some distance from shore. The first group includes the various species of Lymnaea, Physa, and all but two of the species of Planorbis. Specimens of these were generally obtained in shore collections, and were nearly always found in rocky situations or in restricted areas having some peculiar environmental features. In most cases the species involved did not have a very general dis- tribution, but, taken as a whole, the group cannot be neg- lected. The following table gives the species of this division, as well as their habitats. The peculiar localization of some species is well illustrated by the two species Planorbis trivoluis and Planorbts crtsta. Each was found in only one situation where conditions were favourable. The former was obtained in a quiet, well- sheltered harbour at the north end of Alexander Island where the water was very calm. Here the species was very abun- dant because the environment was quite suitable to its fragile shell. The other species, Planorbis crista, was found in a small bay at the south end of the lake. Large numbers ADAMSTONE: MOLLUSCA IN LAKE NIPIGON 103 _of the minute spiny shells occurred on stones in very shallow water close to shore. Of the other members of the group, some were found in protected shallow bays, but the majority of them inhabited rocky, exposed shores, often where they were scarcely covered with water. Character Species of Remarks Bottom Lymnaea apicina.:........... Rock Exposed shores, water 1-3 ft. deep. ns emarginaia.......... ss “ So DGGE one ae = stagnalis (var.)...... aaa Common in water, 1-12 ft. deep. 7 Lg MOE ee Peps igs Na Rock Limited to one _ locality—very shallow water. * CXIICUOUSS: 212 412) c 121s. - ‘ Common in shallow water only. % SVSUTUS)§) a. es. . Frequent in shallow water. 5 femensss le... Mud Limited to one locality—water 4 ft. deep. Physaancilaria............. Rock Exposed shores. ME RUIDIN Pee ets a sh aa a * pS O(N a * Taken only in Pustagone River. SOD Se 3 os oa # & 5 =m shi.

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I bive oe oe ie ea Peer eek wee Oy alrel wan) 70) “aule ile) (sem (anes, DULADIDADA FT or z 6 1 iz 5 = Pe eee ee eee rere em pposo4sQ ¢ = : re - 2 = Fe Se ones oss -ppormua gy ¥Z I 9 9 oe I p ae 9 pater’ cet at Wyeast yom oe ue bet Un eee “‘DjanvyIosuy) OL Z ZI wt 6 OT € oe g inno eeere’ @ rol leeltalid vee e: pLjatnes ia) 8/28) ppodiyquy 9 o. z T Zz T a. oa ania eae | ere reter(e Kerra 6 iae) smh >" las ae “++ pdajqodna AT T oO o oe oe oo se T Ch Yhap es) feceex ar ak i Jolt bait Jeet dese nv VC Dante See tt EO "++ + p4ajgo0aj07 $e 9 9 L ji z z z Z mectetiat Yat it eet On Cent ti eee er a appraaMay gy € I se o. pa oe o. oe zZ So ndNelieste? 18/9) acm aljel im 0im_ (0) leLse "+ pdajqGoy vd [ ZSZ v 00r OF 8¢ ZS 4 II oes Sea a See eee apprmouo yd Gg 6 G OT 9 ¢ ¢ 9 pee | ee Co ee nosnpoyy = 09 Org 09% O1Z O91 OIT 09 Ol |= CEP) S10US 0s; eo ues ei pnw | PO | PU | PUN | PMN | OPPIN | PRIN | PRIN PO wio}}0g JO Ja}9eIeY GT 1Z 1Z ST 81 ral 9 SSS eee eee "***(qa9j uy) yideq g w 9 G P (3) Z T Scop see e shure ehe) (¢usiraxe S16 enereman die ‘Burspa1q s[e}OL ieee ne ion ee ee ee XHOT ses (spuvjs] Suowe apis seq) AVA AYALNDWN BOTTOM ORGANISMS 169 ADAMSTONE AND HARKNESS 1ee OF 121 HW 62 09 gp [cert tert estes ees bee ere a S110] T T BO en | nye eet, 1091 wie ar a byrey ©) see) eg ME Tet Ce EMS. oor butpard ib rigipe—cbilel eae” ewioke ets Déajqodna NT T T oe Ct yummie rel] eccrine GL Jel Je UP ec) SOM eS) or ee bat LO Tea ees PON eat ad et ety ol WO ret eae etm aos D4ajq0aj07 T i T eee ere aera ee ceca elceel stg m) Bow rwirenemet-eotene..0 nd 2 Gintyiys (amen ine tabarente « “p4ao0pv]) T . I Disa eee ce. te MN (Oe Me wel ee Sig lene moeais: 56 telapine ay Oa aie aa ois: relict: DIUIPHALET Zz I I seve wens capa 6ui6yMtueecs (cuteP ameleavicliatanere to) keaie’@aker els ia arallene DULepapapa FY 9 ; ; ae eae ee (85 5 1 Ce) A! og 1 1} 50} 48 1 1 .. |100 1 Pee. joo | 3 25 Cladocera (Opkryoxus gracilis, Chy- | dorus sphaericus) Gyrinid larvae 1; Haliplid beetles 5. 1 .. |100 1 0h ee 15 | .. {Algal filaments. 1 4] 25 35 | 35 |1 Dipterous larva. 1 Se Ne (1 ay v7 NM Ba 1 10 | 50 ye 25 | 10 jAlgal filaments; Cladocera (Alona sp.?). 1 60 40 1 60 5 Plecoptera nymphs 35. 1 50 50 1 5 | 60 a 35 1 .. | .. | 50 | 50 (pupae) lesa 1 Pail) we put eo A 25 | 25 |Diatoms (Surtrella, Navicula). 1 Be tu ean ies 4! 45 | 10 |Diatoms (Surtrella, Epithemia). 1 10)}) -3° 1°30: | 30 15 | 15 1 75 i PANG Wee 1 75 He Nh > 1 50 50 1 Bt (tated YO, wt SOs: 6 +/+ )4+ — + | + |Cladocera (Eurycercus lamellatus). App. Two genera of Ostracoda were represented, namely, Cypria and Candona. Three genera of Ephemeridae were represented, namely Ephemera, Heptagenia snd Tricorythus. All small dipterous pupae have been considered as Chironomidae, although in some cases the fragmentary state made identi- fication uncertain. 184 % THE Foop oF LAKE NIPIGON FISHES SMALL MOUTH BLACK BASS, Micropterus dolomieu Copepoda | Cladocera Insecta is : a ee Date | Length | Epischura = < @ |S 8 § = = No. | 1921 in cm. yar actgec Ss] ss Re Sikes e SR PSs i ey elas s2|S$s|/S8| & |se)se 3 SFatesesy 2 |e 1 |July 19) 2.6 95 5 ARNON he hs if a 2.6 66 2 : A 16 8 8 1 * 2.8 2 2 af 15 |, Ganhag i We 3 Baty cA Meihs ee 80 3 10 Mi 1 6 YELLOW PIKE PERCH, Siizostedion vitreum No. Date | Length Ciscoes Nine-spined Miller’s Fish remains 1921 | incm. sticklebacks | Thumbs Aurel race + 1 July 20) 5.6 + | neo Mn cd + 1 “ 28) 14.0 ue + 1 MZ) ATS 20 4 he 8 2S M70 6 + 1 eet 180 5 + 1 12) 29.5 1 1 ai 1 jAug. 8| 43.7 2 + 1 12) 44.0 1 He 1 |July 28) 47.3 ug 3 Ele 2 Sol Eb) 5020/02) LGFea0iem:) 1 - YELLOW PERCH, Perca flavescens | Copepoda | Cladocera Insecta | aN LEM Gene ae a oliee ad ay SS Cy x 3 | ee Date | Length] 8 3 sleletshs Ss 8 Fah a ae No.| 1921 | incm. |. 8] §] 2/S/S/S/S/8| 82/3] 2 of 2 o/ 8 ol 8 o 2] SB) S/S) SPS isl Si8 g-Sle e)8 s/84] esl. Ql] -8| S]s} slSySjS/S 5) $8 G8 S18 B)25) 38 QL AQ) OSAIQMIOMIO!N s]O/O 8/9 alS giQ alae 1 jAug. 15} 3.0 15). .|50 35 5 |June 27) 3.4 10) 78). Baya 2 atte 10 7 |July 30) 4.7- 62! 137 1 5.6 r 1 tunel) 7.8 +) +). .]. JE]. yl we Jeep oy oe 99? 1The Ephemeridae nymphs were Callibaetis sp.? 2The fish were the nine-spined stickleback (Pygosteus pungitius). Nee SB HB Bee ee ee eee ee Re H pace oR ee | THE Foop oF LAKE NIPIGON FISHES 185 fe oO 5 vet a= Suckers LING, Lota maculosa og o| < TRG Ciscoes aS an EO] 8 Aba Ae 1 1 1 i 1 2 2 1 Shi 1 1 1 2 8 wo: 30 and{ )40 cm. f a 3s | wn eS = i Fas 5 2S S = AOR Ty he oR ae eS 5 Sel =H 1 os 1S 180 == s a Ss [= n SA ls 7 = Sos en! fy AS (OS ~ : 3%| 95% | + | 2% if 1 5 if ial a 1 1 ere ++: 186 THE Foop oF LAKE NIPIGON FISHES LING, Lota maculosa (continued) o & BHl Staal) 2 Ue ents a : ed) ee S a ihe s No. Date te & 2 Ciscoes | 20 a 2 E © ag = Joga) (oh die Cl Ste | ein te ee 4 g Shieh Mn GIN auf Shy iia eg a o| MAIR i fe oie 1 |Aug. 22-23] 35-55] .. {1 (23 cm.) Lhe “1. 11 (20em.,) A ASM ee 1 4 ‘hi 5 ft) ae “ce 1 1 " on 1 MN 1 ¥ Wi Ane ilies 1 1] A Wy 1 {3 Mo ia 2a 1 ac 4c 2 + 1 ni 7 4 aie 1 hr nN 4 AIP stra 1 hy A 3 Nitta 1 6c fa] 1 ae 1 i "i 1 i + 1 ne i 1 1] + 1 te “ 6 ae 1 ve tis 2 1 .. [+ 1 is i a 38; + 1 ‘ A 5 1) + 1 “ “i 7 ub 1 “ 46 1 au 1 é tc 1 a 28 f vi Mt aN Pe BAR SOE, he Me We uy 3 Mi i Hy the CU Asay | ue 100%] .. A ie Aas niliiy y |. | 2. | 10%] 75%! 10%] 3% 1 |July 4 large | .. |3 (11 cm.) nt hy) A he Little need be added in amplification of the data given in the tables, but mention may be made of a few outstanding points. 1. Gill net records show that the northern sucker inhabits deeper water than does the common sucker. The food studies substantiate this fact, in that, as a rule, higher percentages of Pontoporeia occurred in the northern sucker than in the common. It appears also that the older individ- THE Foop oF LAKE NIPIGON FIsHEs 187 uals of the common sucker feed at greater depths than do the younger. 2. Somewhat similar conclusions may be drawn respecting the whitefishes. According to the gill net records, the round whitefish does not extend to as great depths as does the common whitefish, and its food, according to the table, is obtained in comparatively shallow water. The data do not show definitely that the older individuals of the common whitefish feed altogether at greater depths than do the younger. This would be expected where the data is not extensive, because large individuals are often taken in shallow water. It is evident that the common whitefish has serious competitors for food in the suckers, since the latter are bottom feeders and are very abundant in the lake. 3. The ciscoes, although very abundant, come into very little competition with other fish as regards food, in that they are open water plankton feeders, subsisting largely upon Mysis relicta and Limnocalanus. On the other hand they are fed upon extensively by the lake trout. 4, The outstanding item of food of the lake trout is ciscoes. In Lake Nipigon, where the operation of gill nets of 43 inch mesh only is permitted, the number of ciscoes taken is relatively small, and those which are taken are at present sent to the market as whitefish. In view of this fact and since the lake trout is of such great commercial importance, the feeding of the latter upon the ciscoes is not to be deplored. It has been a matter of some surprise that no whitefish have as yet been found in the lake trout stomachs, and if further investigation substantiates this condition, a very fortunate state of affairs will be shown to exist. 5. The food of the trout perch was evidently obtained in the river. 6. The importance of the nine-spined stickleback (Py- gosteus pungitius) in the food of the yellow pike perch is interesting and is possibly correlated with the small numbers of minnows occurring in the lake. 7. The chief competitor of the lake trout is no doubt the ling, since it apparently feeds largely upon ciscoes. The 188 THE Foop oF LAKE NIPIGON FISHES absence of fish of commercial value in its diet is important. It is interesting to note that five individuals had fed upon Mysis relicta. The amounts of the latter were so large as to preclude the possibility of their having been contained in cisco stomachs and, in fact, in three cases no other material could be detected. 8. The fish examined fall more or less definitely into the following groups as regards food:—(1) predaceous—lake trout, yellow pike perch, ling; (2) bottom feeders—sturgeon, northern sucker, common sucker, round whitefish, common whitefish; (8) open water plankton feeders—ciscoes; (4) shallow water plankton and insect feeders—young common suckers, minnows, young small mouth black bass, young yellow perch; (5) insect feeders—trout perch. UNIVERSITY OF TORONTO =» LUDIES BIOLOGICAL SERIES No. 23: A MUSKOX SKULL FROM IROQUOIS BEACH DEPOSITS AT TORONTO: OVIBOS PROXIMUS, SP. NOV., By B. A. BENSLEY THE UNIVERSITY LIBRARY: PUBLISHED BY THE LIBRARIAN, 1923 University of Toronto Studies COMMITTEE OF MANAGEMENT Chairman: Str ROBERT ALEXANDER FALCONER, LL.D., K.C.M.G., President of the University PROFESSOR W. J. ALEXANDER, PH.D. PRoFESSOR J. P. McMurricu, Pu.D. Bric.-Gen. C. H. MitcHEtxt, B.A.Sc., C.B., C.M.G., D.S.O. ProFeEssor G. H. NEEDLER, PH.D. PROFESSOR GEORGE M. Wrona, LL.D. General Editor: W.S. WALLACE, M.A., Librarian of the University SS ee ee A MUSKOX SKULL FROM IROQUOIS BEACH DEPOSITS AT TORONTO: OVIBOS PROXIMUS, SP. NOV. BY B. A. BENSLEY PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF TORONTO A MUSKOX SKULL FROM IROQUOIS BEACH DEPOSITS AT TORONTO: OVIBOS PROXIMUS, SP. NOV. The following description refers to a well-preserved though incomplete skull, consisting of the brain case and horn cores virtually entire, but lacking the face except its upper or naso-frontal portion. The specimen was secured by foreman J. Quinn of the York Sand and Gravel Co., East Toronto, and was brought to the Royal Ontario Museum by. Mr. H. G. Barter. It appears to merit description because of its relative completeness and characteristic nature as com- pared with most specimens, usually fragments, from the Toronto Pleistocene deposits; further because it fills a hitherto disconcerting gap in the faunal record of the Toronto beds; and finally because it exhibits some noteworthy differences of form as compared with previously described types. The area from which the skull was taken is part of the gravel bar formation of the Iroquois Beach deposit as it passes through Toronto. This deposit has been extensively studied by Coleman (’13). It is estimated as late glacial or postglacial and is relatively the youngest of the three char- acteristic Pleistocene beds, Don, Scarborough, and Iroquois, as differentiated at Toronto. Mammalian remains are scarce, though the fine exposures of the Iroquois Beach to the west of Hamilton, Ont., have yielded at least a number of fragments of caribou antlers. The sand and gravel pit mentioned is situated at a point immediately to the east of the present city limit of Toronto, about one-third of a mile south of Danforth Ave. and about half a mile east of Blantyre Ave. It is an excavation of perhaps twenty-five feet in depth, the extent of which is being constantly increased by the removal of material from the sides. An effort was made by Professor Coleman and the writer to determine the precise level at which the speci- 3 4 A MusKox SKULL: OVIBOS PROXIMUS, SP. Nov. men was found, but without success. It was apparently blasted out of the cliff-like wall of the pit, picked up, fortun- ately without damage, in the hoisting scoop, and was only discovered after having rolled off the sand separator screen together with gravel, boulders, and other course material. The edges and more delicate portions of the skull show some chipping as a result of the rough treatment, and the tip of the right horn core had been broken off and lost. Otherwise the skull appears to have been deposited originally much as at present shown, and bears no signs of having been imbedded in any deposit previous to its enclosure in the Iroquois. The specimen bears a general resemblance to the existing O. moschatus. It appears to differ, however, in a more compact and transversely quadrate shape. This is due in part to the very flat condition dorsally of the medial portions of the horn cores, and the fact that the posterolateral borders of the bases of these structures are filled out or buttressed out so that they are flush with the nuchal and lateral sur- faces of the skull. The basal exostoses are further concrescent in the median plane, concealing entirely the sloping portion of the back of the skull, and showing an apparent depth externally in the median plane of about 34 mm. The basal plane of the skull seems to be relatively broader, flatter or less compressed. The compact quadrate contour of the skull is best shown in the view of the occipital surface shown in Plate II, Fig. 3. The appearance is, however, unduly exaggerated by the imperfect condition of the mastoids, the edges of which, doubtless at least slightly convex in life, were apparently broken or worn off on each side before deposition. There is a resemblance in some respects to O. yukonensts as described by Gidley (’08), especially the dorsal contours of the horn core bases, the suggestion of a modified area on the dorsal surface of the skull in front of and more or less connecting the medial portions of the horn bases, and the general aspect at least of the attachments of the horn cores at the back of the skull, though Gidley does not describe the condition of the exostoses on this plane. On the other A Musxkox SKULL: OviBos ProximMus, Sp. Nov. 5 hand, the more expanded appearance of the basal plane in the present specimen, especially the shield-like form and parallel arrangement of the sides of the basioccipital in comparison with Gidley’s excellent figure, seems to indicate a greater deviation from O. yukonensis than is shown by O. moschatus itself. The characters of this specimen are obviously very close to those of O. moschatus and O. yukonensis, which latter is ‘considered by Allen (’13) as possibly falling within the limits of O. moschatus. The postglacial occurrence would suggest a form immediately antecedent to the existing species. On account of the difference noted above, it appears advisable as a provisional arrangement at least, to recognize the speci- men as the type of a species for which the name proximus is proposed. It is impossible at the present time, however, to obtain measurements that will place the type in proper position with reference to O. moschatus, the difficulty being mainly that no ratios appear to be available that can be exactly applied to the specimen and at the same time will give explicitly defined proportional measurements in the three principal axes. Concerning the general affinities of American species Gidley has pointed out that while Ovibos yukonensts is dis- tinctly ovibovine rather than symbovine, it possesses some features reminiscent of Symbos as re-defined by Osgood (’05), namely a rugosity of the frontal portion of the skull sug- gesting the anterior median extension of the exostoses characteristic of Symbos, and further a narrowing of the middle and anterior portions of the basioccipital as in the latter genus. In the present specimen an intermediate condition is suggested in a faint way by the appearance of the frontal area as described by Gidley, and by the posterior concrescence of the horn exostoses in the median plane which feature might be ascribed to O. yukonensis from the appear- ance of Gidley’s figure though he does not describe a con- crescence as actually present. On the other hand, the shield- like form of the basioccipital is extreme in the present speci- men, and thus apparently is relatively more removed from 6 A MuSsKOoXx SKULL: OVIBOS PROXIMUS, SP. NOV. the existing Ovibos than the latter from Symbos. Further, in the present specimen the horn cores, as they leave the skull, appear to be more depressed than in O. yukonensis, 1.e., they lie more nearly in a lateral position, not rising in a graceful curve from the dorsolateral surface of the skull. Referring to the relative depth of the skull as compared with other forms, the available measurements give the impression of dorsoventral compression. The present mas- toid width, 160 mm., has reference to a surface from which both mastoid angles and the paroccipital processes have been broken off. The original width, which was probably not less than 197 mm. The height of the back of the skull, to any point above the nuchal (occipital) crest, cannot be exactly stated, because the area is filled in with the horn exostoses. There is evidently a considerable variation in the ratio occipital depth to mastoid width in modern Ovzibos skulls, but Allen in his extensive memoir does not compare them. His measurements for single specimens of O. mos- chatus and Symbos cavifrons do not define the occipital height, but being doubtless comparable inter se establish depth ratios of .66 for O. moschatus and .90 for S. cavifrons, supporting his contention that the skull of Symbos is rela- tively deeper and narrower than that of Ovibos.* Speaking of Symbos apparently as a genus, however, he overlooks Osgood’s figures for Symbos tyrelli (loc. cit., p. 184), which indicate a much lower occiput than in S. cavifrons. Osgood states explicitly the limits of his occipital measurements. They may not be comparable with Allen’s, but are again doubtless comparable inter se. Expressed as ratios the height of the occiput in one-hundredths of the mastoid width are O. moschatus .68, Syynbos cavifrons .85, S. tyrella .59. Gidley (loc. cit., p. 683) refers to these measurements, adding for O. yukonensis figures yielding a ratio of .626. In the specimen at hand measurements as defined by Osgood yield a ratio of .61, the points of uncertainty being the extent of the exostosis of the back of the skull and the rela- * A similar proportion is indicated by measurements of a skull of S. cavifrons described by Case (’15). A Muskox SKULL: OVIBOS PROXIMUS, SP. Nov. 7 tive accuracy of the width measurement of the mastoid as reconstructed. There is at least some ground for the view that the present specimen has the skull relatively broad and shallow, like O. yuwkonensis in this respect, and that the breadth if not relatively greater than in modern Ovzbos is at least towards the extreme of the latter. Concerning the horn cores, Allen (loc. cit., p. 170) appears to have established that in O. moschatus the bases become approximated, but do not meet in even old animals. There is always a median portion of the skull left exposed. Further, as viewed from the occipital surface the horn cores, even if unsymmetrical to a slight extent, rise naturally from the top of the skull. In the specimen at hand the basal contour of the horn core is obscured and the whole surface squared off by the extended exostosis which not only fills up the natural concavity of the lateral borders but forms a bridge across the median line, interrupted only on its surface by the median furrow separating the horn cores in the mid dorsal line. Thus the two horn cores are perfectly distinct and separate in front, but the median extension of the exostosis apparently increases in depth posteriorly, until at the posterior end of the median groove it has a thickness of about 34 mm. The groove at this point is 26 mm. deep. The downward deflection of the horn cores is as in older males of Ovibos moschatus. Their original length is doubtful, since even the apparently intact one may have been worn down at the tip. The anteroposterior diameter near the median dorsal line shows nothing unusual as com- pared with modern Ovibos as described by Allen. The basal plane of the skull appears broad and flat. Comparison with a male Ovibos moschatus skull kindly loaned me from the collection of the Victoria Memorial Museum* shows several points of difference. Especially in the form of the basioccipital element, the specimen at hand shows a shield-like contour, accentuated by the parallel arrange- ment of the two sides rather than a convergence of them * The writer is indebeted to Director McInnes and Dr R. M. Anderson for the loan of this specimen. 8 A MusKkox SKULL: OVIBOS PROXIMUS, SP. Nov. when traced anteriorly. Doubtless the width of the basi- occipital cannot be properly expressed in terms of the trans- verse width of the skull at the same level since any expansion of the one is likely due to the same circumstances as have brought about transverse expansion of the other. It is noteworthy, however, that on a line cutting the basilar tubercles the ratio of the width of the basioccipital to the width of the skull measured to the borders of the zygo- matic processes of the temporal bones is .36 in the specimen as opposed to .31 in the specimen of O. moschatus. Allen gives some measurements for the anterior and posterior widths of the basioccipital in O. moschatus, but those of the width anteriorly are so small as to indicate that they were taken not on, but in front of, the sphenoccipital synchon- drosis. The posterior tuberosities of the basioccipital are massive, separated by a deep median groove or pit; replaced forwards by a conspicuous ridge in the manner described by _ Allen as characteristic of O. moschatus. The basilar tubercles representing the attachment of ,M. longus capitis are con- spicuously triangular, with their bases forming a straight line transversely raised above the surface of the bone. On account of the absence of the facial region and nearly all of the zygomatic arch, there is a little suggestion of the width of the basal portion of the skull in relation to the palatal aspect of the face. The external borders of the zygomatic processes of the temporal bones suggest a parallel arrangement as between two sides or a direct continuation forwards with less divergence than in Ovibos and much less than in Symbos, as described by Allen. Certain differences which may or may not be distinctive appear in the com- parison of the specimen with the Ovibos moschatus skull of the Victoria/ Memorial Museum already mentioned. In the latter of the basis cranii is almost straight while in the Toronto specimen the basisphenoid turned upward at a considerable angle. In the latter also the pterygoid lamina meets the base of the bone at a much sharper angle, so far as the posterior border of the process is concerned. The osseous bulla appears to be more compressed, and the A Muskox SKULL: OviBos Proximus, Sp. Nov. 9 alae vomeris extend not only over the front of the basis- phenoid, as described by Gidley for O. yukonensis, but pass upward and backward over a considerable portion of the medial surface of the pterygoid lamina. Partial concrescence of the horn cores posteriorly in the specimen described, if we presume that no concrescence of the horn cores exists in modern muskoxen, suggests either that the definition of Ovibos should not be based upon existing forms alone, or that forms showing concrescence should be generically separated. An interesting possibility is that O. proximus (and possibly O. yukonensis) are inter- mediate stages leading from the full concrescence of Symbos to the complete separation of modern Ovibos. There appear to have been two principal mechanical types of horn sup- port, one in which the horn deflection is upward as in Bison, the other in which the deflection is downward as in the Bootherium, Symbos and Ovibos group. Females and younger males of modern Ovibos suggest the stages of a downward deflection, which, from the point of view of apical leverage begins with a condition looking towards the mechanical type of the horn bases in Symbos where there is full con- crescence. But in Ovibos this type is modified and finally replaced by a hook or S-shaped arrangement by which the mechanical stress, both of leverage and direct support, is at points or in a plane parallel to the top of the skull. | Thus the separation of the horn cores in modern Ovibos may be a reduction phase following complete concrescence of the Symbos kind. PRINCIPAL MEASUREMENTS Nuchal plane: | Height, ventral border foramen magnum to MME Re da oe eee uaa, O4 nam. Height, dorsal border foramen magnum to TE ASS UN ae (ah a Height, ventral/surface of condyle to nuchal Height, ventral surface of condyle to top of EXOSLOSISH SNe, Chk PY al 2 Se ME aa Breadth, condyles...... By 8 9 ad Od Breadth, mastoid (imperfect)............. Breadth, mastoid (estimated)............. Horn cores: Anteroposterior diameter near median plane Anteroposterior diameter at point of leaving 14 011 A Ree Oe eee te hens ike eee ay Length, along dorso lateral surface in groove Length, along dorso lateral surface, posterior TVET ANE ee ee se Sl eee Height, exostosis, from level nuchal crest to median groove. . 1.) \2 2a) eee Height, exostosis, ventral edge to median STOOVE a ee ie ee Height, bottom of groove posteriorly to top Of Core sf on hye ee a Basal plane: Length, basioceipital. i... 4. iiss see Breadth, basioccipital, at posterior protu- berances.) 080. .)G0. AH) SUC Breadth, basioccipital, at basilar tubercles (maximum): .7 0 9 ee Breadth, skull at basilar tubercles, to lateral margins of zygomatic processes of tem- POT Aye DLT CR Dorsal plane: Breadth, skull, postorbital. 2.) ee 10 A MusKxox SKULL: OVIBOS PROXIMUS, SP. Nov. 156 146 A MusKox SKULL: OviBos PROxIMUS, SP. Nov. 11 1913. 1915. 1913. 1908. 1905. 1905. LITERATURE CITED Allen, J. A. Ontogenetic and other Variations in Muskoxen, with a Systematic Review of the Muskox Group, Recent and Extinct. Mem. Amer. Mus. Nat. Hist., New York, vol. I, pt. 4. Case, E.C. Ona Nearly Complete Skull of Symbos cavifrons Leidy from Michigan. Occ. Papers, Mus. Zool., Univ. Michigan, Ann Arbor, No. 18. Coleman, A. P. Toronto and Vicinity (Geology of). Ontario Bur. Mines; Guide Book No. 6. Toronto. Gidley, J. W. Descriptions of two new Species of Pleistocene Ruminants of the Genera Ovibos and Bootherium, with notes on the latter Genus. Proc. U.S. Nat. Mus., Washington, vol. 34, No. 1627. Osgood, W. H. Scaphoceros tyrelli1, an Extinct Rumi- nant from the Klondike Gravels. Smiths. Misc. Coll., Washington (Quarterly Issue), vol. 48, pt. 2, No. 1589. Osgood, W. H. Symbos, a Substitute for Scaphoceros. Proc. Biol. Soc., Washington, vol. 18. I 4 PLATE 1 3 figures x Both , left profile s, skull O. proximu O. proximus, skull, dorsal surface PreaAree lt Both figures x 3 Fig! 3. O. proximus, skull, nuchal surface a ae yl a ak a _ iP 7 ~ Etnies ev ee ‘ shy rea eS... » { ee ee BINDING ©7 77 aug 2919/4. University of Toronto. Biological series QH ag T68 (1917-23) no.17=-23 Biologica! & Medi | Serials PLEASE DO NOT REMOVE CARDS OR SLIPS FROM THIS POCKET UNIVERSITY OF TORONTO LIBRARY