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Comparative limnology of lakes in the Southern Rocky Mountain Trench, British Columbia Sparrow, Roger Arthur Hugh 1963

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COMPARATIVE LIMNOLOGY OP LAKES IN THE SOUTHERN ROCKY MOUNTAIN TRENCH, BRITISH COLUMBIA by ROGER ARTHUR HUGH SPARROW B.A., The University of B r i t i s h Columbia, 1955-A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of Zoology We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA SEPTEMBER, 1963 In presenting this thesis in p a r t i a l fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allotted without my written permission. Department of The University of British Columbia, Vancouver 8, Canada. Date ABSTRACT Within a r e s t r i c t e d geographical area of B r i t i s h Columbia, a detailed examination was conducted i n I960 and 1961 of 9-lakes having s i m i l a r morphometric and cl i m a t i c influences, but exhibiting a wide dissolved nutrient range (50 to 1460 ppm). Attempts were made to relate t o t a l dissolved s o l i d s as well as other chemical and physical indices, to b i o l o g i c a l factors i n d i c a t i n g or influencing lake productivity. Measurements were made of standing crops of plankton, bottom fauna and f i s h as well as hypolimnial oxygen d e f i c i t s , sedimentation rates and gross primary productivity. Standing crops of plankton appeared. relat,ed to oxygen d e f i c i t s and perimeter to area r a t i o s . Furthermore, t o t a l dissolved s o l i d content correlated with gross primary productivity estimates based on.Light and Dark Bottle techniques. Total dissolved s o l i d content of lakes was not correlated with standing crops of plankton, bottom fauna or f i s h . Ranking of selected physical or chemical indices of productivity f a i l e d to agree with ranking based on standing crops or other b i o l o g i c a l measurements of productivity. A number of possible factors which interact to modify the expression of primary correlates of productivity are discussed. - v i i i -ACKNOWLEDGMENTS The author appreciated the assistance of J.V. M a c k i l l , J.D. Varty and M. Teraguchi during the c o l l e c t i o n of f i e l d data. Encouragement by my wife during the course of this investigation i s also sincerely appreciated. Special thanks go to Dr. P.A. Larkin for suggesting the problem and to Dr. G.'G.E. Scudder and Dr. P.A. Larkin for valuable c r i t i c i s m of the manuscript. The guidance and c r i t i c i s m of Dr. T.G. Northcote throughout the study has been invaluable and i s therefore acknowledged with thanks. I am indebted to the Pish and Game Branch of the Department of Recreation and Conservation for use of equipment and a l l o c a t i o n of time for .collecting lifn'nological data. I am grateful to the Consolidated Mining and Smelting Company of Canada for the chemical analysis of water samples. - i i i -TABLE OP CONTENTS Page TITLE PAGE i ABSTRACT ' i i TABLE OF CONTENTS .>...» . . i i i LIST OF FIGURES v LIST OF TABLES v i ACKNOWLEDGMENTS i v i i i INTRODUCTION J . 1 HISTORICAL RESUME OF PRODUCTIVITY MEASUREMENTS. 2 MATERIAL AND METHODS • 3 DESCRIPTION OF THE STUDY AREA 8 Geographic Location..'..... 8 Geology. 8 S o i l s .. 11 Vegetation 12 Weather and Climate 14 Air Temperature... 14 P r e c i p i t a t i o n 16 Wind 17 PHYSICAL CHARACTERISTICS OF THE LAKES 19 Topography of the Drainage Area 19 Morphometry 22 Temperature 23 Transparency 27 Ranking of the Lakes by Physical Factors... 29 - i v -Page CHEMICAL CHARACTERISTICS OP THE LAKES . . . 32 Total Dissolved Solids and Al k a l i n i t y . . . . . . 32 Hydrogen Ion Concentration.... 32 Ion Composition... 34 Calcium and Magnesium.. . 34 Sulphate.......... 34 Dissolved Oxygen...................... I... 34 Ranking of the Lakes 1by Chemical Factors.. 37 BIOLOGICAL CHARACTERISTICS OF THE LAKES 39 Primary Productivity. 39 Flo r a . 41 Net Plankton........ 41 Bottom Fauna 47 Qualitative Composition 47 Quantitative Composition 47 Fish. 51 Hypolimnion Oxygen D e f i c i t i . . 53 Sedimentation Rates; 55 Ranking of the lakes by B i o l o g i c a l and other Factors Indicating Productivity....; 55 DISCUSSION .... 60 SUMMARY. ! 69 LITERATURE CITED 70 - V -LIST OP FIGURES Page Figure 1. Location of the 9 study lakes and weather stations within the Southern Rocky Mountain Trench of B r i t i s h Columbia.................. Figure 2. Location of the 9 study lakes i n the Southern Rocky Mountain Trench with respect to regional geology...... •. 10 Figure 3 . Bathymetric maps of the 9 study lakes with limhological stations as indicated (S). A l l - lakes have the same compass orientation'and scale of measurement - 21 Figure 4. Hypsographic curves of the 9 study lakes 24 Figure 5. Seasonal temperature (°C) d i s t r i b u t i o n with depth (ft) from one limnology station i n each of the 9 lakes i n I960 and 1961. Approximate period of ice cover i s indicated.. i 25 Figure 6. Secchi disk transparency i n r e l a t i o n to maximum depth ( f t ) of the 9 lakes during I960 and 1961 28 Figure 7. Variation of dissolved oxygen (mg/liter) with depth ( f t ) i n the 9 lakes during I960 and 1961 36 Figure 8. Relative abundance of benthos i n bottom samples taken from the 9 study lakes. Figure indicates an average of a l l bottom organisms taken from dredge samples on 6 sample dates"in'1960"arid'' 1961 48 Figure 9. Figure 10, Average number of a l l bottom organisms taken per square meter from bottom dredge samples. Numbers along the .ordinate represent the maximum depth of each 10 foot stratum sampled. Relationship of average summer (June-August, I960) plankton volumes (cc/m ) to mean hypolimnial oxygen d e f i c i t s (cc/liter/day) for the 9 study lakes i n I960. Figure 11. Relationship of the r a t i o of perimeter,to area to average plankton volumes (cc/m ) the 9 study lakes i n I960 and 1961 for 50 63 65 Figure 12 Relationship of t o t a l dissolved content (ppm) to gross primary productivity (nigC/m /day) as measured by Light and Dark Bottle techniques' during the period of 27 June - 8 July, 1961.., 67 - v i — LIST OP TABLES Page Table I Monthly and annual mean temperatures for the years I960 and 1961 and averages for the periods shown............... - 15 Table I I Monthly and annual t o t a l p r e c i p i t a t i o n for the year I960 and 1961 and averages for periods shown.... 18 Table I I I Morphometric data for lakes i n the Southern Rocky Mountain Trench.. 20 Table IV Mean epilimnion temperatures (°C) of the 9 lakes for I960 and 1961 26 Table V Ranking of the 9 lakes by physical factors.. 30 Table VI Chemical analysis of water samples for the 9 lakes i n the Southern Rocky Mountain Trench expressed as a mean and range for I960 and 1961 33 Table VII Ranking of the 9 lakes by chemical factors.. 38 Table VIII Gross primary productivity estimates for Light and Dark Bottle experiments i n the 9 lakes during June 27 - July 8, 1961... 40 Table IX The dominant forms of aquatic plants found i n the 9 lakes of the Southern Rocky Mountain Trench... 42 Table X Mean plankton volumes (cc/m^) for the 9 lakes examined i n 1960 and 1961 44 Table XI Mean dry weight of plankton samples obtained during I960 and 1961 45 Table XII Relative abundance of the dominant plankton genera i n the 9 lakes during I960 and 1961.. 46 Table XIII Number and weight of f i s h taken i n a g i l l net set i n each lake during June and July of 1961... 52 Table XIV Absolute areal and mean hypolimnetic oxygen de f i c i t s . . . ' . . 54 Table XV Sediment, volumes as measured from sediment co l l e c t o r s suspended at a specified depth i n each lake for a 10 day period'in'September, 1961... 57 - y i i -Page Table XVI Ranking of the 9 lakes by b i o l o g i c a l Table XVII Ranking of the 9 lakes by other factors i n d i c a t i n g productivity. 59 Table XVIII Ranking of the lakes by physical, chemical and b i o l o g i c a l c h a r a c t e r i s t i c s as well,as other factors i n d i c a t i n g productivity....... 62 INTRODUCTION Northcote and Larkin (1956) found t o t a l dissolved s o l i d s (T.D.S.) the best single indicator of standing crops of plankton, bottom fauna and f i s h , i n 100 lakes exhibiting a wide range of T.D.S. throughout B r i t i s h Columbia. However, for 33 lakes within the Southern I n t e r i o r Plateau region of the province, t o t a l dissolved s o l i d s (ranging between 85 and 558 ppm) showed no s i g n i f i c a n t r e l a t i o n s h i p with standing crops of organisms (Larkin and Northcote, 1958). Thus, although t o t a l dissolved., s o l i d content of lake waters apparently was related to standing crop l e v e l observations over the whole province, no such r e l a t i o n s h i p was apparent when lakes exhibiting a f a i r l y broad range i n T.D.S., but within a r e s t r i c t e d geographic area, were investigated. The purpose of this study was to examine in d e t a i l nine lakes a l l located within the Southern Rocky Mountain Trench having a wide dissolved nutrient range (50 to 1460 ppm) and to ..evaluate the importance of T.D.S. i n determining standing crop levels i n lakes where v a r i a t i o n i n climatic and morphometric factors modifying these le v e l s was minimal. HISTORICAL RESUME OP PRODUCTIVITY MEASUREMENT i Many studies of lake productivity have examined the p o s s i b i l i t y of using a single variable as an index of productivity i n comparisons of lakes. Various degrees of success have been achieved, but no single index has been p a r t i c u l a r l y s a t i s f a c t o r y , especially i n comparisons of lakes from a var i e t y of regions. Thienemann (1927) was the f i r s t to suggest the use of lake morphometry, and i n pa r t i c u l a r mean depth, as an indicator of productivity. Rawson (1930, 1952 and 1953) also'stressed the importance of mean depth on the production of plankton, bottom fauna and f i s h , as i t applied primarily to deep lakes. Strain (1931) and Hutchinson (1938) emphasized the measurement of hypolimnetic oxygen d e f i c i t as providing a means of c l a s s i f y i n g lakes. Oligotrophia lakes having small oxygen d e f i c i t s and eutrophic lakes having complete oxygen depletion i n the hypolimnion, were exceptions to th i s c l a s s i f i c a t i o n . Naumann (1932), Ohle (1934), Deevey (1940) and Moyle (1946), a l l considered the geology of the drainage•area or some portion of the nutrient cycle, as being of primary importance to the measurement of productivity among lakes. Other workers have used plant abundance-and d i s t r i b u t i o n as a useful index of production i n lakes. More recently, emphasis has centered around the use of C ^ or oxygen measurement by Light (LB) and Dark Bottles (DB), as a measure of photosynthetic a c t i v i t y or net and gross primary productivity. - 3 -MATERIALS AND METHODS Hand l i n e soundings were used to plot contour maps of Hiawatha, KlakhoHorseshoe and Bednorski lakes. A Kelvin-Hughes echo sounder provided information for the construction of contour maps of the others. Soundings were plotted on lake outlines, i .enlarged from B r i t i s h Columbia Department of Lands and Forests interim maps compiled from a e r i a l photographs. The volume between 2 depth contours was estimated by using the formula V=h (A-^  + k^) where h i s the distance between the 2 s t r a t a , and A-^  and A2 represent the areas of the respective upper and lower contours. The volume of the lowermost -portion of the lake was estimated using the area of the lower contour multiplied by the estimated mean depth within the area. Other parameters of lake morphometry and morphology, with the exception of the perimeter to area r a t i o , were calculated as suggested by Welch (1948). The transparency of the lake water, measured with a Secchi disk 20 cm i n diameter, was stated as the mean of the depth of disappearance and reappearance of the disk. A l l Secchi disk readings were taken on the shaded side of the boat. Notes on cloud cover and condition of water surface were recorded. Water samples for analysis of dissolved oxygen were obtained using a 1200 cc Kemmerer water b o t t l e . The sampler had been i n use for several years but had not been coated to prevent contact of water with the metal. Dissolved oxygen was determined using the unmodified Winkler technique. Corrections f o r a l t i t u d e and temperature were according to Truesdale, - 4 -Downing and Lowden as tabled i n Hutchinson (1957). No adjustment was made i n the t i t r a t i o n values to account for the volume of reagents added except for those used i n gross primary productivity c a l c u l a t i o n . A l l oxygen t i t r a t i o n s up to 27 June 1961 were made using 100 cc aliquots and t i t r a t e d with 0.025N sodium thiosulphate; thereafter 50 cc aliquots and 0.01N sodium thiosulphate were used. A n a l y t i c a l chemists of the Consolidated Mining and Smelting Company of Canada, KimberTey, B.C., analyzed water samples for dissolved nutrients u t i l i z i n g methods outlined i n Standard Methods (I960). A Yellow-Springs thermistor-thermometor, model 43TB, was used to record a i r and water temperatures. This instrument had been standardized with a long stem mercury thermometor calibrated to 0.1G. Temperatures were estimated to the nearest 6.1 of a Centigrade degree. U n t i l March 1961, a "Wisconsin" plankton net with a diameter of 11.2 cm was adopted for taking t o t a l v e r t i c a l plankton hauls. Thereafter, a 25 cm diameter net of the Wisconsin type was used. Both nets were f i t t e d with new No. 10 s i l k having 109 meshes per linear inch. Each time a lake was sampled', 3 t o t a l v e r t i c a l plankton hauls were taken at the deep water st a t i o n . A t o t a l of 18 samples was obtained from each lake and each was preserved i n a 5 percent formalin solution. Ward (1957) found that rapid random changes i n net-ef f i c i e n c y did not obscure differences i n crustacean a v a i l a b i l i t y between.stations oh Shuswap Lake. Ward also found that Wisconsin type nets were not subject to gradual changes i n ef f i c i e n c y during the period of use, nor were catches s i g n i f i c a n t l y different over - 5 -a range of hauling rates. Net plankton volumes were determined using a 24 hour s e t t l i n g period i n 15°0 cc centrifuge tubes calibrated to 0.1 cc. Because the formalin contained a flocculent p r e c i p i t a t e , a l l samples taken prior to Marbh 1961, were mixed with dilute o x a l i c acid and f i l t e r e d , before volume determination. Two plankton samples from t r i p l i c a t e v e r t i c a l hauls, were dried i n an oven at 60C for 48 hours or u n t i l a constant weight was obtained. Samples were f i r s t washed with d i s t i l l e d water. Weights were determined using a Mettler model K7 a n a l y t i c a l balance, sensitive to 0.03 grams. Bottom fauna samples were taken at 5 d i f f e r e n t periods of the year although none were taken during ice cover of the lake. Samples were discarded where the dredge f a i l e d to close completely. Although molluscs were found to be a minor part of the bottom fauna i n most lakes, shells were removed p r i o r to weighing. The t o t a l 1 number of dredgings i n each lake varied from 22 to 42, with duplicate samples being taken at each depth sampled. Bottom fauna dredging consisted of sampling with a 6 inch Ekman dredge at approximately the same position i n each depth zone, each time a lake was examined. The depth zone was chosen for ease of sampling, on i n i t i a t i o n of the study. Samples were washed using a 30 mesh per l i n e a l inch screen and sorted on a white enamel tray while the organisms were a l i v e . A f i v e percent formalin solution preserved the bottom fauna for l a t e r i d e n t i f i c a t i o n . - 6 -Only the common higher aquatic plants were noted as evident i n bottom dredgings and by examination of lake shoal areas. The f i s h fauna was sampled with standard sets of braided nylon experimental g i l l nets having stretched mesh sizes of 1 to 3lr inches at i inch i n t e r v a l s . G i l l nets were set at selected shoal areas for a 12 hour period during the night. A l l f i s h taken from the net were weighed with a spring balance and measured to the nearest 0.1 cm by using a measuring board. A 10 percent solution of formalin was used as a f i s h preservative. A sediment trap^ following the design of Patalas (personal communication), consisted of a 24 ounce ice box jar suspended at the lower l i m i t of the thermopline for a 10 day period. The l i d was designed to close when the trap was brought to the sur-face.: Formalin was placed i n the bottom of the jar prior to the ^ suspension of the sediment sampler i n the lake. Gross primary productivity was estimated using the 'flight and Dark Bottle'! method of oxygen evolution. Light and Dark Bottles, consisted of 300 cc BOD (biochemical oxygen, demand) bottles with the "dark'' bottles having been made l i g h t tight by covering with black ''Scotch Tape" as well as having several coats of black "Glyptol" paint. Paired Light and Dark Bottles were suspended at depths of 1, 3, 5 and 7 feet from metal hinges fastened to a weighted wire l i n e . The l i n e was i n turn suspended from a 1 square foot styrofoam buoy. Water samples were taken with a Kemmerer bottle from which 3 BOD bottles could be f i l l e d . One b o t t l e , termed a "blank," was used to determine the dissolved - 7 -oxygen at the start of the experiment. Bottles were shielded from sunlight u n t i l they could be suspended at their appropriate depth. Gross primary productivity was measured i n the 9 study lakes during the period of June 27th to July 8th, 1961. The LB and DB were suspended for a 12 hour period , from 0815 or 0915 to 2015 or 2115, depending upon the time required to f i n i s h setting the bottles at the various depths. T i t r a t i o n followed methods given by Strickland (I960). A l l oxygen deter-minations were done on the same day as the " i n situ!' oxygen measurements. The photosynthetic quotient was assumed to be 1 for gross primary productivity calculations. - 8. -DESCRIPTION OP THE STUDY AREA Geographic Location A l l of the lakes i n the study are located between longitude 115° 3 0 ' and 116° 30', and between 49 and 51 degrees of latitude. Figure 1 shows the location of the nine lakes i n the Southern Rocky Mountain Trench, B r i t i s h Columbia. Two main mountain ranges delimit the Southern Rocky Mountain Trench. The Rocky Mountains r i s e abruptly to the east of the v a l l e y f l o o r and a t t a i n an average height of 6,000 to 8,000 feet. The Columbia Mountains, and i n p a r t i c u l a r , the P u r c e l l Range, border the west side of the trench but r i s e less abruptly as r o l l i n g h i l l s . The P u r c e l l Range has an average a l t i t u d e of 6,000 feet. The f l o o r of the Rocky Mountain Trench varies between 2,460 and 2,800 feet elevation i n the study area. The lakes studied range between 2,757 feet and 3,921 feet elevation (Table I I I ) . Figure 1 gives the location of the 9 lakes i n the Southern Rocky Mountain Trench. A l l lakes are accessible throughout the year except Kiakho and New lakes, which are usually inaccessible for winter sampling because of heavy snowfall. Geology Figure 2 gives the position of the 9 lakes i n r e l a t i o n to the regional geology of the Southern Rocky Mountain Trench. Geological areas have been obtained from map 932A of the F i g u r e 1. L o c a t i o n o f the 9 study l a k e s and weather .. s t a t i o n s w i t h i n the Southern Rocky Mountain Trench of B r i t i s h Columbia - 10 -fl"fj"fl C A L C A R E O U S & ARGILLACEOUS SEDIMENTARY C A L C A R E O U S SEDIMENTARY Figure 2 Location of the 9 study lakes i n the Southern Rocky Mountain Trench with respect to regional geology - 11 -Department of Mines and Technical Surveys. Schofield (1915) shows Hiawatha, Kiakho and Jim Smith lakes situated i n the Aldridge formation (map 147A). The southern portion of Kiakho Lake l i e s within an area of granite and porphyritic granite. Schofield describes the Aldridge formation as consisting of argillaceous quartzites, purer quartzites and a r g i l l i t e s . Interlocking grains of quartz have been thoroughly cemented together and a small amount of feldspar i s present. Rice (1937) i n map 396A depicts Kiakho and New lakes within the Creston formation. Argillaceous quartzites and quartzites compose the Creston formation according to Rice. This formation-has quartz as the most abundant mineral-. Calcareous beds are not uncommon and carbonates occur in minor,quantities.: ' : Lazy Lake i s located i n an area of sedimentary sand and clay on map 932A. Horseshoe Lake and Enid Lake are located i n areas of stream and g l a c i a l deposits (map 932A) where the bedrock remains concealed. Schofield (1915) mentions a formation of s i l i c e o u s limestone-which surrounds Bednorski Lake (map 147A). L i l l i a n Lake i s likewise situated in. an area of Limestone (map 2070). Walker (1926) describes the area as the Mt. Nelson formation, composed of magnesian limestone and s l a t e . S o i l s Information i s available on s o i l c l a s s i f i c a t i o n i n areas surrounding some of the lakes. Kelley and Sprout (1956) state - 12 -that g l a c i a l t i l l and i t s derivates are the main soil-forming materials i n the Southern Rocky Mountain Trench. Enid and Lazy lakes and approximately one quarter of L i l l i a n Lake l i e within an area of the Wycliff S i l t Loam s o i l s (Kelley and Holland, 1961). The parent material of this s o i l i s a loamy, strongly calcareous t i l l containing gravel and stones. This s o i l type i s noted for i t s high t o t a l calcium content and the accompanying low t o t a l phosphorus. Bednorski Lake, the south end of Lazy Lake and a small portion of L i l l i a n Lake are situated within the Wigwam S o i l Complex. .This group of s o i l s have been developed on a l l u v i a l fans of limestone or non-calcareous sedimentary rocks (Kelley and Sprout, 1956). Elko S i l t Loam s o i l s comprise approximately one half of the s o i l s around L i l l i a n Lake. The amount of phosphorus i n this s o i l type i s comparable with the Wycliff s o i l s . The s o i l texture i s a loam, to s i l t loam and the amount of stones i n the surface varies. Around Horseshoe Lake the Elko S i l t Loam i s combined with the Saha S o i l s to form the Elko-Saha group. This s o i l type supports a mixture of grass and trees. The Saha s o i l s are gravelly with highly porous subsoil. Vegetation The p r i n c i p a l species of trees i n the southern part of the trench are ponderosa pine (Pinus ponderosa),-, western larch (Larix occidentalis) and Douglas-fir (Pseudotsuga menziesii). - 13 -Hiawatha and New lakes have wooded shore lin e s consist-ing of western larch and lodgepole pine (Pinus contorta). The drainage areas of both lakes were logged more than ten years ago. L i l l i a n Lake has a forested shoreline consisting of Douglas-fir and some deciduous trees such as trembling aspen (Populus  tremuloides). Kiakho and Jim Smith lakes have p a r t i a l l y open shore li n e s with lodgepole pine and western larch as the dominant coniferous trees. The Jim Smith Lake drainage was logged over whereas the Kiakho Lake drainage has been the s i t e of logging practices i n the l a s t 10 years. The eastern slope of the Kiakho drainage i s p a r t i a l l y open and composed of rock slid e s and semi-open grassland. The south and north ends of Jim Smith Lake are open to wind action and are composed of grassland or pasture. The shoreline i s free of trees around Enid, Horseshoe and Lazy lakes and p a r t i a l l y treed around Bednorski Lake. Douglas-f i r and ponderosa pine stands occur back from the shore of Lazy Lake. The drainage area was the s i t e of logging about 10 to 15 years ago. Only limited logging took place around Horseshoe Lake where Douglas-fir, lodgepole pine and ponderosa pine are the major species. Bednoi'ski Lake has Douglas-fir, western larch and lodgepole pine i n the drainage area, which was logged about 10 years ago. Deciduous trees such as aspen and alder are the most prominant about the southwest portion of the lake. Most of the lake shore i s open grassland and the north or west end of the lake i s cultivated land. The shores of Enid Lake have some Douglas-fir and aspen but a large part of the shore i s open grassland or sparsely treed. - 14 -Weather and Climate In the B r i t i s h Columbia Atlas of Resources, the Southern Rocky Mountain Trench i s c l a s s i f i e d as the Southeastern Interior subregion of the Interior c l i m a t i c region. The trench assumes a north-south d i r e c t i o n and for this reason a i r masses may assume a similar d i r e c t i o n of movement. Cold a i r may move into the trench area from the north or the region may be subjected to masses of warm a i r from the south during the summer. The l o c a l topography produces regional c l i m a t i c differences throughout the trench, compared with the dominant, modified Polar. Maritime a i r mass. Variations in temperature and pr e c i p i t a t i o n occur with changes of a l t i t u d e throughout the Rocky Mountain Trench. A i r Temperature Temperature records are available from The Climate of B r i t i s h Columbia for I960 and 1961 (B.C. Dept. of Ag r i c u l t u r e ) , for 1 inactive and 4 active weather stations, close to the lakes studied (Pig. 1). The monthly and annual mean temperatures for I960 and 1961 are given for the 4 active stations i n Table I. Also shown are the averages for the periods on record. '„••• The Cranbrook, Kimberley and Wilmer weather stations a l l have approximately the same a l t i t u d e . The average temperatures are lower as"one progresses north from Cranbrook to Wilmer. The greatest difference i s observed during the winter period. The I960 and 1961 temperatures for Cranbrook and Kimberley r e f l e c t the same pattern as the averages on record. TABLE I Monthly and annual mean temperatures for the yea (Degre Station Jan. Feb. Mar. Apr. May Cranbrook 1960 12 23 31 M. M. 3013 f t 1961 M. M. 35 41 53 Average 16 22 32 43 52 -Aberfeldie • . - • I960 16 26 33 44 50 2640 f t "1961 25 35 38 42 . 53 Average 21 26 33 43 53 Radium Hot Springs I960 9 21 28 41 46 3570 f t 1961 18 27 32 39 50 Average 15 21 29 40 51 Kimberley Airport 1960 11 22 30 43 48 3016 f t 1961. 21 30 35 41 52 Average 15 22 29 42 52 Wilmer (Inactive) Average 14 20 31 43 51 3100 f t s I960 and 1961 and averages-for. the. periods shown s F.) June July Aug. Sept. Oct. Nov. Dec. A, Y r 9 M. M. M. M. M. M. M. M. 63 66 68 49 40 25 17 M. 58 64 62 54 43 29 22 41 30 58 70 62 56 40 32 20 43 63 67 68 50 42 27 21 44 58 65 63 55 44 31 25 43 11 54 66 59 51 42 25 15 38 61 62 63 45 37 19 15 39 57 63 60 50 39 22 17 39 7 58 69 60 54 45 28 17 40 63 64 66 48 39 24 16 42 58 64 62 54 42 28 20 41 18 58 64 61 52 • 41 27 16 40 16 M. Missing 1. Annual 2. Years - 16 -Cranbrook and Aberfeldie have about the same l a t i t u d e (Pig. 1) but different elevations (Table I ) . The average temperatures are colder at Cranbrook which has the highest elevation. The effects of a l t i t u d e and lat i t u d e on a i r tempera-tures i s also reflected in. the growing season of lakes. Figure 5 shows that ice had not yet begun to form by November 6 on Lazy Lake or November 7 on Bednorski Lake, but Enid and L i l l i a n lakes were half frozen by November 15. A more s t r i k i n g pattern i s given for the date of ice cover removal. Bednorski, Horseshoe and Lazy lakes are.lower i n a l t i t u d e than Jim Smith, Kiakho or New lakes. Ice formed e a r l i e r and remained longer on lakes of higher elevation (Fig. 5). P r e c i p i t a t i o n Total annual p r e c i p i t a t i o n records i n the study area vary from 12 to 23 inches between stations. P r e c i p i t a t i o n i n the form of snow accounts for 35 to 85 inches per annum between regions. Approximately 30 percent of the. p r e c i p i t a t i o n i s i n the form of snow (Table I I ) . The month of June i s on the average the wettest month at a l l f i v e recording stations. March and A p r i l are the months usually having the lowest precipitation.. Table I I shows there i s approximately the same annual p r e c i p i t a t i o n at Cranbrook and Kimberley a i r p o r t s . A decrease in p r e c i p i t a t i o n at Wilmer i s evident when compared to the Crapbrook and Kimberley averages. An increase i n p r e c i p i t a t i o n i s reflected by changes of alt i t u d e between Wilmer and Radium - 17 -Hotsprings. The higher p r e c i p i t a t i o n at Aberfeldie when compared to Cranbrook, could be due to Aberfeldie being located near a route taken by thunderstorms (Kelley and Sprout, 1956). Enid and L i l l i a , n lakes, i n proximity to Wilmer, would be expected to have lower p r e c i p i t a t i o n than other lakes studied. Likewise, Bednorski , Horseshoe, Hiawatha and Lazjy lakes, because of their elevation being lower than New, Kiakho and Jim Smith lakes, would have lower p r e c i p i t a t i o n .Wind Wind records are available for the years 1938 to 1945 from the Cranbrook airport weather s t a t i o n . Kelley and Sprout (1956) give the average wind speed of 7.1 mph for south winds i n the trench regibn and 4.5 mph for the north winds. Chapman (1952) states that northwest and southeast winds are dominant at Windermere i n January and J u l y , but south and southwest winds are dominant at Cranbrook i n both seasons. This i s shown by records where 63 percent of the winds at Cranbrook are from the south, southeast and southwest. North, northeast and northwest winds account for 29 percent of the t o t a l winds at Cranbrook. - New, Lazy and Kiakho lakes, which have the i r maximum length as...the north-south axis, and Jim Smith Lake which has a southwest - northeast a x i s , would be expected to have the greatest effect from dominant winds at Cranbrook. 'Hiawatha and Kiakho lakes' are protected from excessive wind action as a result of h i l l s situated around the lakes. Horseshoe Lake with i t s open shoreline, i s susceptible to wind from any d i r e c t i o n . TABLE I I Monthly and annual t o t a l p r e c i p i t a t i o n for the year I960 and 1961 and averages for periods^ shown (Total p r e c i p i t a t i o n i s the sum of the r a i n f a l l plus the water equivalent of the snowfall, which i s snowfall divided by 10) (Inches) Station Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. Aberfeldie 196Q 1.81 2. 00 1. 41 0.78 2-. 39 1. 61 0. 44 1.64 1 .03 1 .90 4 .54 0.96 20 • 51 2640.ft 1961 2.01 2. 06 1. 54 1.46 2, 57 1. 64 1. 82 1.47 4 .28 1 • 73 1 .53 2.:34 24 .49 (58.8) 5 Average 3.30 1. 84 1. 04 1.02 1. 80 2. 73 1. 03 1.66 1 .50 1 .86 2 .13 2.32 22 .23 Cranbro ok 1960 0.76 0. 74 1. 35 M. M. M. M. M. M M M M. M. 3013 f t 1961 M. M. 1. 2Q 1.16 1. 40 1. 00 1. 41 0.79 3 .11 0 .89 1 ^34 3.45 M. (64.2) Average 1.58 1. 41 0. 81: 0.79 1. 21 1. 95 0. 83 1.07 0 .95 1 .24 1 .34 1.99 15 .17 Kimberley 1960 0.88 0. 86 0. 82 " 0.44 1. 11 0. 58 0. 34 2;.21 0 .55 0 .51 2 .72 0.80 11 .81 Airport 1961 0.84 2. 01 0. 94 0.62 1. 96 1. 16 2. 08 1.54 1 .98 1 .21 0 .83 1.81 16 .98 3016 f t Average 1.77 1. 22 0. 94 0.75 1. 27 1. 96 0. 74 1.36 0 .90 1 .21 1 • 36 1.64 15 .12 Radium I960 0.97 1.25 1.30 0.58 2.43 1.42 0.53 0.81 1.15 2.36 M. 1.58 M. Hot Springs 1961 2.38 1.80 0.35 2.00 2.38 1.70 2.48 1.49 2.64 1.06 0.95 1.25 20.48 3570 f t Average 1.59 1,36 0.90 1.85 1.92 3.00 2.34 1.75 2.30 1.94 1.56 2.08 22.59 (83.4) Wilmer (Inactive) Average 1.2-3 0.61 0.52 0.63 1.20 1.64 1.29 1.53 1.10 0.78 0.88 1.1-5 12.56 3100 f t (35.5) M. - Missing 1. - Periods of record are same as i n Table I 2. - Annual -3^: - Average snowfall i n inches - 19 -. PHYSICAL CHARACTERISTICS OF THE LAKES , Topography of the Drainage Area Table I I I l i s t s the drainage areas and water l e v e l f l u c t u a t i o n of the lakes examined. Hand l i n e soundings from a limnological station were used to record the water l e v e l . f l u c t u a t i o n , except i n Horseshoe and Lazy lakes, where, f l u c t u a -tions were measured between high and low water marks on the shore. Enid and Horseshoe lakes have no surface i n l e t s or outlets. The lake l e v e l of Horseshoe begins to r i s e during the early summer and f a l l s once again i n late summer. This lake has unusually large water l e v e l fluctuations since a difference of 21 feet was measured between low water of May 2, 1962 and the v i s i b l e high water mark. Hiawatha,' L i l l i a n , Jim Smith and New lakes have seasonal i n l e t s or outlets or both, which are normally dry by ' 3 late summer. Most of the creeks have volumes less than 0.5 f t /sec but the i n l e t flow of L i l l i a n Lake i s between 1 and 2 f t ' / s e c . Kiskho Lake, described by Stenton (I960), has a' drainage area of approximately 3 square miles. Two i n l e t creeks enter Kiakho Lake and i n 1958 Stenton found the flow of each i n l e t to be 0.5 f t /sec. The outlet i n the spring has a flow of 3 to 4 ft^/s e c but i n summer i s considerably l e s s . The permanent:inlet of Lazy Lake has a volume o f about 0.25 ft^/sec throughout the year and the stream f a l l s sharply from the mountains on the east side. Between September I960 and March 1961, a 4 foot f l u c t u a t i o n was recorded i n the lake l e v e l . However, TABLE I I I Morphometric d a t a f o r CD O ; si 03 CD CO f-l Q K E l e v a t i o n m ( f t ) 853 2799 S u r f a c e A r e a H e c t a r e Acre 1 0 . 7 * 2 6 . 4 3 1 . 5 7 7 . 8 Maximum Depth m ( f t ) 6 . 1 * 20 1 1 . 6 38 Mean Depth m ( f t ) 2 . 7 * 8 .8 Volume m 3 X m5 ( A c r e f t ) 2 . 8 7 * 233 Maximum Length m ( f t ) 600* 1969 1152 . 3780 Maximum Width m ( f t ) 277* 909 503 1650 Shore Development 1.20* 1.63 Volume Development 1.32* P e r i m e t e r m ( f t ) 1400* 3246 4594 10650 P e r i m e t e r y 1 G 3 A r e a 3 . 9 9 * 3 . 1 4 D r a i n a g e A r e a H e c t a r e ( m i l e ) 2020 7 . 8 0 Water L e v e l " ( f t ) ' 2 L . 0 •H CO m o c T3 CD w 853 2757 9 . 6 2 3 . 8 13.1 43 8 .5 2 7 . 9 8 .19 644 503 1650 226 742 1.21 as si +j 05 aJ •H tc o si 05 •H •H T3 •H Si si •H E C O e •rl t-0 tS3 05 t-3 1.95 1-53 1.44 ^ l . 3 1 1.50 , 2 . 0 4 r . 5 0 1333 845 288 3228 2414 2359 3886 4372 2773 9460 10590 7920 7740 12750 4 .22 6 .28 7 . 2 8 3 . 7 0 3 .31 3 .38 3 .77 4105 124 15 .85 0 . 4 8 1.0 1.5 777 461 . 132 3 .00 1.78 0 . 5 1 1.5 1.5 2 . 0 1243 1362 4 . 8 0 5 .26 2 . 0 6 .0 CD 957 1097 942 963 1058 905 1195 3200 3600 3091 3160 3470 2970 3921 4 . 1 1 2 . 1 2 6 . 6 2 2 . 3 2 1 . 2 3 1 . 4 27 .6 1 0 . 1 2 9 . 9 6 5 . 8 ' 5 5 . 0 5 2 . 5 7 7 . 6 6 8 . 3 5 .2 9 . 8 1 1 . 6 8 .2 7 . 3 10 .7 10 .4 i ro o 17 32 38 27 24 35 34 2 . 6 4 . 7 ^ . 1 4 . 1 4 . 9 5 .2 3 .4 i 8,7 1 5 . 3 1 6 . 7 13 .5 1 6 . 2 17 .2 11 .1 1.08 5 .65 13 .57 9 . 1 8 1 0 . 5 3 16 .51 9 . 3 1 88 458 1100 744 854 1338 755 344 1006 777 701 846 1372 1052 1128 3300 2550 2300 2775 4500 3450 168 11? 777 594 320 297' 372 • 552 385 2550 1950 1050. 975 1220 1.17 2 . 3 4 1.76 1.44 U . 43 1.95 1.33 0 .98 2487 8160 2 .74 202 0 .78 2 . 0 F l u c t u a t i o n * - Low water data' 21 -Figure 3i Bathymetric maps of the nine study lakes with limnological stations as indicated (S); A l l lakes have the same compass orientation and scale of measurement. - 22 -the difference between the low water l e v e l and the v i s i b l e high water mark has been measured at 6 feet. Bednorski Lake has permanent i n l e t and outlet streams estimated at 1' ft^/sec i n June I960. The volume (0.25 ft^/sec) during the late summer to spring period i s reduced considerably. Morphometry Morphometric parameters are l i s t e d for the lakes i n Table I I I and bathymetric maps are given i n Figure 3. The areas of the lakes range from 10.1 acres (Hiawatha) to. 77.8' acres (Horseshoe). The lakes could be grouped into two sizes;'Hiawatha, Bednorski, Kiakho and Horseshoe at low water, as the smaller lakes, and the remainder as the larger size group. The 4 smaller lakes range from 10.1 to 29.9 acres and 5 large lakes have a range of 52.5 to 77.6 acres. The shoreline development of Hiawatha Lake most cl o s e l y approximates that of unity. Kiakho Lake, on the other hand, shows the. greatest departure (2.34) from a c i r c u l a r form. The perimeter to area r a t i o i s probably a better c r i t e r i o n than shore l i n e development when comparing lakes of differ e n t areas (Rawson, I960). With the exception of Hiawatha and Kiakho lakes, a l l other lakes have-similar r a t i o s . : . .. . The hypsographic curves (Fig. 4) show the features of shape'and area of the lakes at various depths. Bednorski and Jim Smith-lakes !have 68 and 73 percent of their surface areas 1: represented at one half of 'their maximum depths. New Lake has 24 percent of i t s surface area at 50 percent of maximum depth. The :remaining 6 lakes are similar i n having 38 to-53 percent of - 23 -their area at one half the maximum depth. Horseshoe Lake at low water l e v e l and Hiawatha Lake have mean depths which are about one half of the magnitude of other lakes. However, Table I I I shows a s i m i l a r i t y of mean depth with a range of 8.7 to 27.9 (Bedriorski Lake) for the 9 lakes studied. The 9 lakes have s i m i l a r maximum lengths ranging from 1,128 feet for Hiawatha to 4,500 feet for Lazy Lake. This distance i s the eff e c t i v e length of the lake over which the wind may act. The 9 lakes chosen for examination are similar i n many of their morphometric features including maximum depth, mean depth, volume development and maximum length. This i s especially true when considering the great differences to be found among lakes i n B.C. or even within the Rocky Mountain Trench. Temperature . . Figure 5 i l l u s t r a t e s the temperature at various depths of a l l lakes during the period of examination i n I960 and 1961. Except for Hiawatha Lake, a l l lakes have similar surface and bottom temperatures and a l l exhibit thermocline formation. Table IV shows that mean summer epilimnion temperatures for July 20 to August 5, I960 and August 30 to September 1 of 1961. Computation was according to the formula i n Welch (1948). Except for Hiawatha Lake, a l l lakes had simi l a r epilimnion temperatures. Epilimnion temperatures showed a greater v a r i a t i o n i n 1961 compared to I960. Lazy Lake had a thermocline located at approximately 25 feet, whereas other lakes except Hiawatha, had thermoclines at 10 to 15 feet (Fig. 5). P E R C E N T O F S U R F A C E A R E A Figure 4. Hypsographic curves of the 9 study lakes - 25 -I960 1961 Figure 5. Seasonal temperature (°C) d i s t r i b u t i o n with depth ( f t ) from one limnology s t a t i o n i n each of the 9 lakes i n I960 and 1961. Approximate period of i c e cover i s i n d i c a t e d - 26 -TABLE IV Mean e p i l i m n i o n temperatures (°C) of the 9 lakes f o r I960 and 1961 Lake Mean E p i l i m n i o n Temperature (°C) 20 J u l y - 5 August, I960 50 August - 1 September, 1961 Kiakho 21.2 17.6 L i l l i a n 20.6 18.9 Enid 21.4 18.3 New-.' 21.5 19.4 Hiawatha 23.8 20.0 Jim.jSmith 21. 4 19.9 Horseshoe 21.3 20.0 Bednorski 20.2 19.3 Lazy 21.5 21.3 - 27 -By mid-September of I960, the surface and ep i l i m n i o n temperatures of a l l lakes decreased considerably. However, Bednorski, L i l l i a n and Enid lakes s t i l l e x h i b i t e d a thermocline, but at a lower depth than during J u l y and August. Temperatures i n November were higher i n Lazy Lake than other lak e s . A near homothermous c o n d i t i o n e x i s t e d i n a l l lakes ( P i g . 5) during November, when temperatures ranged between 1.5 and 6.3 C. F i g u r e 5 shows the approximate time of i c e formation i n November. As Kiakho Lake was i n a c c e s s i b l e during March, no temperatures were recorded. New and Jim Smith l a k e s , being at the highest e l e v a t i o n , had i c e sheets 21 to 23 inches i n thic k n e s s . Hiawatha had 18 inches of i c e while the remaining lakes had 8 to 14 inches of i c e . Horseshoe and Lazy lakes were the warmest during the period of winter s t a g n a t i o n . Most lakes show a s l i g h t increase i n temperature near the bottom. The approximate time of i c e removal i s i n d i c a t e d f o r a l l lakes i n Pigure 5. Temperatures taken during the summer of 1961 can be compared with those of I960 (Figure 5). A l l lakes were w e l l s t r a t i f i e d i n the summer of 1961. New Lake was much warmer i n 1961 than I960. L i k e w i s e , Hiawatha, Jim Smith, L i l l i a n and Enid lakes were al s o warmer i n 1961. Lazy lake had temperatures i n 1961 that were s i m i l a r to those of I960, whereas, Bednorski and Kiakho lakes had temperatures that were colder i n 1961. Transparency The greatest transparency was observed i n L i l l i a n , New, NEW LILLIAN ENID HIAWATHA 4-JIM SMITH 2 0 -2 5 -2 0 -2 5 -r-LU 2 0 -2 5 -35 I D_ LU Q 1 0 -15 -10-2 5 -30 -3 5 -BEDNORSKI 1 S E C C M I X M A X D E P T H HORSESHOE LAZY KIAKHO 6. Secchi disk transparency i n r e l a t i o n to maximum depth ( f t ) of the 9 lakes during I960 and 1961 - 29 -Lazy and Jim Smith lakes ( F i g , 6 ) . Intermediate i n transparency were Hiawatha, Kiakho, Horseshoe and Bednorski l a k e s . Enid lake was outstanding by having the lowest v i s i b i l i t y of the nine l a k e s . E n i d , Hiawatha, Jim Smith and Bednorski lakes e x h i b i t e d l i t t l e v a r i a t i o n i n transparency measurement during the year. Ranking of the Lakes by P h y s i c a l Factors Ranking of the lakes i s based on a s c a l e of 0-9 w i t h the lake having the highest value being given 9 and the lake w i t h the lowest value given 0 (Reimers, Maciolek and Pister., 1955). Lakes with intermediate values between highest and lowest are ranked between 0 and 9 i n pr o p o r t i o n to t h e i r absolute values The f o l l o w i n g formula i s used f o r f a c t o r s d i r e c t l y r e l a t e d to p r o d u c t i v i t y . (9) Value f o r lake - minimum value f o r , „ a l l lakes Rank X = . - • Range of values f o r a l l lakes, For a f a c t o r i n v e r s e l y r e l a t e d to p r o d u c t i v i t y the formula i s : (9) Maximum value f o r a l l lakes - value f o r „ , „ lake Rank X = • - " • • • • Range of values f o r a l l l a k e s . Table V l i s t s some p h y s i c a l f a c t o r s which have been reported i n the l i t e r a t u r e as i n f l u e n c e s or i n d i c a t o r s of p r o d u c t i v i t y . With the exception of transparency, a l l are TABLE V Ranking of the 9 lakes by p h y s i c a l f a c t o r s . Mean Lake Mean Depth Perimeter Area Mean Transparency Ep i l i m n i o n Temperature Drainage Area Mean 'Rank Hiawatha 9.0 7.0 5.8 9.0 0 6.2 Kiakho 5.9 9.0 4.8 0 1.5 4.2 Horseshoe !8.9 2.5 5.1 4.3 4.3 5.0 New. 7.9 0 0 3.6 0.1 2.3 Jim'Smith 5.5 1.3 . 3.0 4.3 2.5 3.3 Bednorski 0 3-0 5.8 1.1 9.0 3.8 Lazy 5.0 2.0 1.6 7.2 2.9 3.7 L i l l i a n 5.3 1.9 1.3 1.1 0.8 2.1 Enid- 6.7 1.1 9.0 1.4 0 3.6 - 31 -d i r e c t l y r e l a t e d to p r o d u c t i v i t y . The mean rank i s not meant to imply that a l l p h y s i c a l f a c t o r s i n f l u e n c e p r o d u c t i v i t y e q u a l l y , but i n the absence of knowledge concerning'their i n d i v i d u a l i n f l u e n c e s , equal weight must be given to each f a c t o r . The mean ranks e x h i b i t a narrow range of values with the p o s s i b l e exception of Hiawatha Lake (Table V). Mean depths are s i m i l a r except f o r Bednorski Lake which has the highest mean depth and therefore the lowest rank. Kiakho and Hiawatha have perimeter to area r a t i o s much higher than o t h e r , l a k e s . L i k e w i s e , New and Enid lakes are outstanding by having the.:; highest and lowest transparency. A l i k e n e s s e x i s t s among mean e p i l i m n i o n temperatures except f o r the high temperatures of Hiawatha and Lazy l a k e s . Horseshoe Lake, and Bednorski Lake i n p a r t i c u l a r , r e c e i ved a high rank f o r the s i z e of drainage area. - 3 2 -CHEMICAL CHARACTERISTICS OF THE LAKES T o t a l Dissolved S o l i d s and A l k a l i n i t y The lakes studied v a r i e d from 50 (Hiawatha) to 1463 (Enid) parts per m i l l i o n i n t o t a l d i s s o l v e d s o l i d s (Table V I ) . The lakes were chosen purposely to incl u d e a wide range of T.D.S. The d i s s o l v e d s o l i d content showed seasonal and annual v a r i a t i o n , however values l i s t e d i n Table VI are averages f o r the summer and autumn of I960 only. Bednorski Lake was the only lake with T.D.S. higher i n summer than autumn. A l l lakes had summer t o t a l d i s s o l v e d s o l i d s higher i n 1961 than I960. Seasonal v a r i a t i o n s were greatest i n lakes of high t o t a l d i s s o l v e d content. A l k a l i n i t y increased with increases i n t o t a l d i s s o l v e d s o l i d s . Lazy lake was the only lake which showed a departure from the trend of i n c r e a s i n g a l k a l i n i t y with i n c r e a s i n g T.D.S. Hydrogen Ion Concentration A l l the lakes studied were a l k a l i n e w i t h only Hiawatha Lake ever having a pH i n the range of 7 to 8. Most lakes ranged from 8 to 9 and Enid Lake had pH values sometimes i n excess of 9. The pH was higher i n summer than autumn, or opposite to the T.D.S. r e l a t i o n s h i p . TABLE VI Chemical a n a l y s i s of water samples-for the 9 lakes i n the Southern Rocky Mountain Trench expressed as a mean and range f o r I960 and 1961 AUO, T o t a l Phen, v v S as % 5 T.D.S. ^  P H ( l ) A l k ( 3 ) A l k i y ^ SQ 4 MgO CaO S t 0 2 F e 2 ° 3 Hiawatha ; Summer 53 8.2 47 ,1.0 0.8 5.2 16.3 6.2 2.1 Autumn 58 8.1 (28-60) (0-2) (0-3) (4-6) (11-24) (2-12) (0-3) Kiakho Summer 94 8.8 , 73 4.5 N 1.0 12.1 32.5 14.4 2.7 Autumn; 116 8.2 (50-87) (3-6) (0-2) (10-15) (28-38) (2-27) (0-3) Horseshoe Summer 111 8.4 87 5.5 V 1.0 18.5 46.3 9.6 2.6 Autumn 115 8.4 (70-124) (5-6) (0-2) (13-36) (33-86) .(8-12) (2-4) New ; Summer 144 8.7 12"3 6 .0 0.5 25.9 44.6 14.4 2.1 Autumn 149 8.5 (86-162) (5-7) (0-1) (19-46) (34-64) (2-20) (1-3) Jim Smith Summer 168 8.5 145 9.0 n 35.7 49.7 15.9 2.6 Autumn 193 8.5 (104-187) (8-10) (0-3) (29-50 (33-5,5) (12.18) (1-4) Bednorski Summer 268 8.5 189 13.0 12.8 41.3 64.7 11.8 3.1 Autumn 259 8.6 (135-241)(12-14) (0-48) (10-50) (24-80) (10-16) (1-5) Lazy - Summer 300 8.9 132 11.0 35 73.9 41.3 9.7 2.7 Autumn 326 8.6 (98-164)(10-12) (20-37) (60-79) (34-66) ('6-11) (1.4) L i l l i a n Summer . 462 • 8.8 249 11.5 36 94.0 58.5 10.1 2.9 Autumn 478 . 8.7 (147-406) (9-14) (22-80) (65-105) (39-75) (9-11) (2-4) Enid ' Summer U 8 3 9.2 , 356 37 132 271.7 84.4 8.6 4.1 Autumn 1371 8.9 (280-560) (34-40) (79-195) (153-357) (16-236) (6-11) (2-7) ( l ) T.D.S. and pH are summer and autumn averages f o r I960 only 12) numbers i n parenthesis represent range of samples taken i n I960 and 1961 (3) A l k a l i n i t y (4) Phenolphthalein - 34 -Ion Composition Calcium and Magnesium The calcium concentration was high i n a l l lakes except Hiawatha (Table V I ) ; Ohle (1934) considered calcium as being high when i n excess of 25 m g / l i t e r . Bednorski, L i l l i a n and Enid lakes were the r i c h e s t i n calcium concentration. These lakes were also the lakes having marl formations on the lake bottom. The magnesium concentration was l e s s than the calcium concentration i n the lakes of lowest T.D.S. In lakes of high T.D.S., the q u a n t i t y of magnesium was greater than the q u a n t i t y of calcium. Large seasonal f l u c t u a t i o n s i n magnesium concen t r a t i o n were apparent i n lakes of high T.D.S. Enid and L i l l i a n had magnesium q u a n t i t i e s i n excess of 50 ppm while the remaining lakes had from 3 to 50. ppm. Sulphate Enid Lake had a sulphate concentration v a r y i n g from 79 to 195 ppm. The odor of H^S from lake bottom samples was very prevalent. The 5 lakes of lowest T.D.S. al s o showed the lowest concentrations of sulphate, with l i t t l e subsequent annual v a r i a t i o n . Lakes of high T.D.S. had high sulphate concentration w i t h l a r g e annual f l u c t u a t i o n s . D i s s o l v e d Oxygen Figure 7 i l l u s t r a t e s the oxygen c h a r a c t e r i s t i c s of the lakes during the summer and autumn of I960, and the spri n g and - 35 -summer of 1961. Hiawatha, Bednorski and Enid lakes i l l u s t r a t e examples of a clinograde oxygen curve. Summer oxygen values i n 1961 confirm r e s u l t s obtained i n I960. Kiakho Lake i s i l l u s t r a t e d i n Figure T as having a p o s i t i v e heterograde oxygen curve during J u l y and August of I960. However, i n September of I960 and J u l y of 1961, Kiakho Lake had a more t y p i c a l clinograde oxygen d i s t r i b u t i o n . Examples of orthograde oxygen d i s t r i b u t i o n are Horseshoe'* New, Jim Smith, Lazy and L i l l i a n l a k e s . New Lake i n June 1961 and Lazy Lake i n August of I960 show a tendency toward dlinograde oxygen d i s t r i b u t i o n s . Oxygen concentrations i n a l l lakes i n November of I960 are s i m i l a r w i t h more or l e s s equal amounts of oxygen at a l l depths. However, between lakes s t r i k i n g d i f f e r e n c e s i n oxygen curves are observed duri n g March of 1961 ( F i g . 7 ) . Clinograde oxygen d i s t r i b u t i o n s are evident i n Hiawatha, New, Jim Smith, Lazy and Enid lakes i n March 1961, while orthograde curves are found i n Horseshoe and L i l l i a n l a k e s . Bednorski Lake shows an i n d i c a t i o n of a p o s i t i v e heterograde oxygen d i s t r i b u t i o n . Between November of I960 and March of 1961 ( F i g . 7) a l l lakes except Horseshoe Lake, e x h i b i t marked oxygen decrease. This i s p a r t i c u l a r l y evident i n Enid and New lakes where a p a r t i a l winter k i l l of f i s h has been observed by the w r i t e r . Bednorski, Lazy and Jim Smith lakes have a marked oxygen decrease below 15 fe e t between November and March. - 36 -OXYGEN CONCENTRATION IN MILLIGRAMS PER o — IO -»-' o — 1 0 -2 0 -3oU' O to • 2 0 -30-<o-5 _ _ J O I 5 -i—m-i 1 ^ JUNE LLl L U 0_ L U Q O — IO-2 0 -3 0 -to -o — 10 -2 0 -3 0 -O ~~ IO-20-30-4 0 -o — 1 0 -2 0 -30-4 0 -• JUL * f IO-2 0 -3 0 -4 0 -O — IO-2 0 -3 0 - / 4 0 -• JUNE 5 J O IS *UGllST ' -T-m—i 1 1 • 1 1 i—w-i 1 5 10 15 5 10 15 • HIAWATHA .' *. f. P I KIAKHO -r-y-, , t 1 <;rn • 11 HORSESHOE r NEW ~ I 1 JIM SK-ITH • ~i 1 " T W 1 1 BEDNORSKI ~T~W 1 T SF. PT. LAZY LILLIAN ENID I ~ I 1 • Figure 7. Variation of dissolved Oxygen ( with depth ( f t ) i n the 9 lakes 1960 and 1961 - 37 -Ranking of the Lakes by Chemical F a c t o r s Chemical f a c t o r s were ranked by the same method as p h y s i c a l f a c t o r s . Low oxygen concentrations i n the hypolimnion have been used as an i n d i c a t o r of high p r o d u c t i v i t y i n the ep i l i m n i o n . Magnesium and calcium have s i g n i f i c a n c e i n p l a n t and animal ecology. High s i l i c a concentrations were assumed to i n d i c a t e a p o t e n t i a l f o r productivity,however low concentrations could i n d i c a t e high u t i l i z a t i o n . Table V I I gives the ranking f o r the 9 l a k e s , based on chemical f a c t o r s . The mean rank of lakes other than E n i d , l i e w i t h i n a somewhat narrow range '(,1.6 - 4.1). Enid Lake has a mean rank n e a r l y twice the value of the next highest l a k e . TABLE V I I Ranking of the 9 lakes by chemical f a c t o r s . Mean Mean Mean Mean Mean Bottom Oxygen Mean Lake T.D.S. MgO CaO S^O? (Summer) Rank Hiawatha 0 0 0 0 8.0 1.6 Kiakho 0.4 0.2 2.1 7.6 8.5 3.8 Horseshoe 0.4 0.4 4.0 3.2 5.5 2.7 New 0.7 0.7 3.8 7.6 3.6 3.3 Jim Smith 1.0 1.0 4.4 9.0 1.6 3.4 Bednorski 1.5 1.2 6.4 .5.2 6.0 4.1 Lazy 2.0 2.3 3.3 3-3 0.3 2.2 L i l l i a n 2.9 3.0 5.6 3.6 0 3.0 Enid 9.0 9.0 9.0 2.2 9.0 7.6 - 39 -BIOLOGICAL CHARACTERISTICS OF THE LAKES Primary P r o d u c t i v i t y Most d u p l i c a t e t i t r a t i o n s f o r gross primary p r o d u c t i v i t y estimates agreed w i t h the - 0.08 mg 0 2 / l i t e r l i m i t s set by N* S t r i c k l a n d (I960). Greatest disagreement i n oxygen t i t r a t i o n values occurred i n New, Lazy and L i l l i a n lakes (Table V I I I ) . At c e r t a i n depths the Dark B o t t l e oxygen concentration exceeded that of the L i g h t B o t t l e . This was evident i n Lazy and Enid lakes (Table V I I I ) . A l s o , oxygen concentrations were higher i n the Blank than i n the Light B o t t l e i n some lake s . Where.DB values exceeded LB values, they were not used f o r estimates of gross primary p r o d u c t i v i t y . Average gross primary p r o d u c t i v i t y estimates f o r the 9. lakes may be grouped i n t o lakes of low, moderate and high p r o d u c t i v i t y . Hiawatha, Horseshoe and Bednorski lakes have low primary p r o d u c t i v i t y (1L3-119 mg C/m /day), whereas Kiakho, Jim Smith, New and Lazy have moderate p r o d u c t i v i t y (286-395 mg C/m /day). High gross primary p r o d u c t i v i t y was observed i n Enid and L i l l i a n lakes (1040-1157 mg C/mVday). The minimum mean gross primary p r o d u c t i v i t y was observed i n Hiawatha Lake and the maximum i n L i l l i a n Lake (Table V I I I ) ; these values are i n the range reported i n the. l i t e r a t u r e . Eberley (1959) gave a mean production rate i n Myers Lake of 754 /<g 0 2 / l i t e r (282 mg C/m ) with d a i l y gross primary production r a t e s as high as 1377 /<g O g / l i t e r i n the metalimnion. Weber (1958) has given r a t e s of 910 mg C/mVday f o r West Okoboji Lake i n Iowa. - 40 -TABLE VII I Gross primary produc t iv i ty estimates for L igh t and Dark Bot t le experiments i n the 9 lakes during June 27 - Ju ly 8, 1961 Gross Primary Lake Depth Blank Light Dark Produc t iv i ty ( f t ) 0, 0, 0, •1. •1. 1. (mgC/mVday) Mean Hiawatha Kiakho Horseshoe New Jim Smith Bednorski Lazy L i l l i a n Enid 1 8.59 8.62 7.84** 292 3 8.42* 8.73 8.44 109 5 6.75* 8.61 8.48 49 7 8.74 8.54 8.53 4 1 8.41 8.43 8.34 34 3 7.87 7.88 7.89* 5 7.88 , 7.88 5.83 769 7- 8.27 8.29 a. 14 56 1 10.06 10.07 9.70 139 3 10.06 10.04 9.63 154 5, 10.03 10.11 9.96 56 7 10.40 10.40 Spoiled — 1 7.95 7.93 7.58 131 3 7.91 7.82 6.39** 536 5 7.87 7.93 7.76 64 7 7.98 7.83 6.29** 577 i 7.76 7.86 6.21 608 3 7.72 7.94 7.81 48 5 7.65 8.01 7.31 262 1 7.75 8.16 7.75 154 3 8.01 8.17 7.99 67 5 7.73 8.19 7.74 169 7 7.94 8.17 7.94 86 1 9.89 9.84 10.20** _ 3 9.30** 9.45 9.80** — 5 10.28 9.37 7.68 634 7 9.08 9.73 9.31* 157 1 7.46 8.22 4.49** 1399 3 6.62** 8.72** 6*91 679 5 7.57** 8.49** 8.17 120 7 6.77 8.59 2.11 243Q' 1 9.79 9.95 9.76 71 3 9.69 10.00 2.20* 2925 5 9.69 8.66 9.47 -7 9.70 9.80* 9.47 124 113 286 116 329 306 119 395 1157 1040 ** - Duplicate t i t r a t i o n s agree wi th in 0.06 and 0.10 m(* O p / l i t e r - Duplicate t i t r a t i o n s with difference greater than - 0.10 mg 0 2 / l i t e r 1. - o| mg / l i t e r - 41 -F l o r a E n i d , L i l l i a n , Lazy and Bednorski lakes had the highest T.D.S. of the lakes examined as w e l l as marl substrates w i t h Chara as the major aquatic p l a n t (Table I X ) . Enid Lake had a sparse r e p r e s e n t a t i o n of Chara with meagre areas of ass o c i a t e d marl sub s t r a t e . Bednorski, and i n p a r t i c u l a r Lazy Lake, had Chara w e l l represented as the dominant p l a n t . Hiawatha, New and Jim Smith lakes had the g r e a t e s t abun-dance of macrophytes, with Potamogeton, Nuphar and Chara as the major genera. These lakes had low mean depths and during the summer Hiawatha and New lakes had approximately one-half of t h e i r surface areas covered by.emergent aquatic p l a n t s . Potamogeton was evident i n the north and south ends of Kiakho Lake and then only sparsely d i s t r i b u t e d . .' t. Horseshoe Lake had a large, part of i t s substrate composed of g r a v e l . Substrate composition coupled with major water l e v e l f l u c t u a t i o n i s the probable reason f o r no major aqu a t i c plant d i s t r i b u t i o n i n Horseshoe Lake. T e r r e s t r i a l grasses formed part of the aquatic plant a s s o c i a t i o n at high water i n Horseshoe Lake. Net Plankton . Plankton samples obtained from t o t a l v e r t i c a l plankton hauls (#10 net) i n I960 and 1961, had the greatest volume during the summer months (Table X). An a n a l y s i s of variance of plankton volumes showed s i g n i f i c a n t d i f f e r e n c e s between lakes (p < 0.01) and between periods of sampling (p < 0.01). Using Duncan's - 42 -TABLE IX The dominant forms of aquatic p l a n t s found i n the 9 lakes of the Southern Rocky Mountain Trench Dominant Forms of Aquatic P l a n t s Lake Hiawatha Kiakho Horseshoe New Jim Smith Bednorski Lazy L i l l i a n Enid CtJ as o 6 C 3 o rH as +> ' r-t •H CD >5 u • bC Xl CD 5-1 ; o P. as ; e o CO as •H •H PL U r-l 3 o >. as •83 > +++ ++ + ++ +++ + +.++ +++ +++ ++ ++ as rH at rH as cb Xl ft •H •rH >: 83 K H + + + + +++ - Abundant •¥+ - Moderate' + - Sparse - 43 -m u l t i p l e range te s t (Duncan, 1955), Horseshoe, Bednorski, Hiawatha and Kiakho lakes had s i g n i f i c a n t l y the g r e a t e s t standing crops of plankton. Of the 5 remaining lakes , Enid , Jim Smith and Lazy had s i g n i f i c a n t l y greater plankton volumes than New or L i l l i a n l a kes. Average dry weight of plankton from each lake gave a q u a n t i t a t i v e rank s i m i l a r to the rank based on plankton volume (Table X I ) . The major d i f f e r e n c e involved the order of Enid and Bednorski lakes i n the range of values. Enid Lake had the gr e a t e s t weight of plankton but had the s i x t h highest plankton volume. But Bednorski Lake ranked f i f t h by plankton weight and second by plankton volume. The other 7 lakes had approximately the same r e l a t i v e rank by plankton volume or weight. Gammarus l a c u s t r i s although o f t e n taken i n plankton samples i n Enid Lake, was omitted p r i o r to r e c o r d i n g plankton • volumes and weight. Daphhia sp. was represented i n plankton samples from a l l lakes and Diaptomus sp. from a l l but Lazy Lake (Table X I I ) . Ceriodaphnia sp. and pyclops sp. were other forms of zooplankton commonly observed. " Phytoplanktonic forms were i d e n t i f i e d to genus although most phytoplankton would be expected to pass through the number 10 net. Two f l a g e l l a t e s , Vdlvox sp and Ceratium sp. were the most common phytoplankters. Volvox sp. was moderately abundant i n samples from Hiawatha Lake. TABLE X Mean plankton volumes (cc/ra ) f o r the 9 lakes examined i n I960 and 1961 (Numbers i n parenthesis r e f e r to the range of t r i p l i c a t e t o t a l v e r t i c a l hauls) Mean Lake June-July 22 20 1960 July-Aug. 25 ; i i I960 Sept.-Sept. 12 22 I960 Nov.-Nov. 5 15 1960 March-March 14 28 1961 June-July T o t a l .27 .8 Volume 1961 ( c c V ) Hiawatha 1.8.2 (16.7-21.4) 51.1 (41.0-59.0) 9.4 ( 7.7-10.3) 10.3 (10.3) ( 1.8 1.3- 2.0) 13.6 17.6 (13.4-13.9) Kiakho 16.5 (11.5-19.5) 10.0 ( 6.9-11.5) 17.6 (17.2-18.4) 5.9 (5.9) - - - 12.3 12.4 (10.6-15.3) Horseshoe 28.5 (25.3-33.3) 8.9 ( 4.8-12.4) 11.9 ( 8.3-15.5) 10.6 ( 7.2-13.0) ( 6.7 3.8- 8.5) 7.4 12.3 ( 5.5-10.4) New 11.4 ( 8.6-13.3) 4.3 ( 4.0- 5.0) 2.9 (2.9) 5.6 ( 3.7- 7.4) ( 1.4 0.6- 2.8) 3-.,4 4.9 ( 3 . 1 - 3 . 5 ) Jim Smith 10.2 ( 6.9-16.7 16.2 (15.1-16.7) 10.1 ( 7.2-15.9) 4.0 ( 3.0- 6.1) ( 0.8 0.6- 0.9) 10.1 8.6 ( 4.9rrl3.4) Bednorski 49.4 (43.9-52.6) 18.3 (16.2-19.8) 13.7 (12.3-14.9) 7.8 ( 6.3- 9.9) ( 2.8 2.5- 2.9) 4.9 16.8 ( 4.6- 5.2) Lazy 26.7 (24.8-30.5) 16.1 (13.0-18.5) 13.9 (13.0-14.8) 5.1 ( 4.8-5.7) 0.2 (0.2) 7.2 11.5 ( 7.1- 7.3) L i l l i a n 5.1 ( 4.3- 6.0) 8.9 ( 6.O - 1 0 . 3 ) 5-0 ( 4.4- 5 . 3 ) 4.4 ( 3.5- 5.3) ( 0.9 0.8- 1.0) 3.8 4.7 ( 3.6- 4.0) Enid 4.1 ( 3.3- 4.4) 9.6 ( 8.0-11.5) 9.2 ( 8.0-10.3) 2.9 ( 1-1- 5.4) ( 4.1 3.9- 4.4) 5.9 6.0 ( 5.3- 6.4) TABLE XI Mean dry weight of plankton samples obtained during I960 and 1961. Lakes Dry weight Hiawatha Kiakho Horseshoe N § w J i m Smith Bednorski Lazy L i l l i a n Enid of Plankton : : — — — ;— • : — - '. •  mg/m3 1 3 1 . 99.1 100.4 22.9 41.3 75.7 68.3 33-3 158.1 kg/hectare 16.9 18.6 27.1 7.9 8.5 28.5 25.5 11 . 3 47.9 TABLE X I I R e l a t i v e abundance of the dominant "plankton genera i n the 9 lakes during I960 and 1961. Plankter Lake • H c o ca • H c cfl t 3 O • H CD o CO c • H E CO o pq CO I o -+» cO • H P l CO o r-i O o CO 3 o o cfl O Hiawatha +++ ++ + 4 + + Kiakho +++ ++ +++ + Horseshoe +++ ++'+ + ++ .+ + New +++ +++ ++ + Jim Smith ++ +++ + + Bednorski ++ ++ +++ +++ + Lazy- + +++ + L i l l i a n +++ ++ + Enid + +++ cfl o r-i O 4-? O CO • H Cfl >H • H O cfl 4-3. C f l E c CO Cfl r H 3 0) r H B r-i • H CO r H CO • r l +3 o , Q • H c t»c cfl > CO O cfl JH r H c 0] m 0) O o N o > + + +.+ + + + +++ - Abundant ++ - Moderate + - Sparse - 47 -Bottom Fauna Q u a l i t a t i v e Composition Figure 8 gives the q u a l i t a t i v e composition of bottom fauna i n the 9 lakes. Many of the same taxonomic groups of organisms were represented i n a l l l a k e s . Notable exceptions were the absence of amphipods from Hiawatha Lake and the presence of only chironomids and amphipods i n Enid Lake. H y a l e l l a azteca has been observed i n Hiawatha Lake but was not present i n bottom fauna samples. Chironomids were present i n a l l lakes and c u l i c i d s (Chaoborus sp.) were found i n a l l but Lazy and Enid lakes. Kiakho, Jim Smith, Bednorski and L i l l i a n lakes had H y a l e l l a azteca and Gammarus l a c u s t r i s represented i n samples, whereas Horseshoe, New Lazy and Enid lakes had one or the other of the amphipods, but not both. Ephemeropterans were found i n a l l lakes and molluscs i n a l l but Horseshoe Lake. Hiawatha Lake was the only lake without r e p r e s e n t a t i o n of Hydracarina sp. T r i c h o p t e r a , Coleoptera and Nematoda were a d d i t i o n a l groups c l a s s i f i e d as "others" i n Figure 8. Hemiptera were not represented and t h i s i s no doubt due to the l i m i t e d types of sampling. Q u a n t i t a t i v e Composition The r e l a t i v e abundance of each major i n s e c t group, c l a s s i f i e d on a numerical b a s i s , i s given i n Figure 8 f o r bottom fauna samples examined from the 9 lake s . HIAWATHA KIAKHO HORSESHOE NEW JIM SMITH BEDNOPSKI LAZY L ILL IAN ^ CHIBONOMIOAE CULICIOAE G A MMARUS H Y A L E L L A ENID ANNELIDA O D O N A T A OTHEWS Fxgure 8. R e l a t i v e abundance of benthos i n bottom samples , taken from the 9 study lakes. F i g u r e i n d i c a t e s an average of a l l bottom organisms taken from dredge samples on 6 sample dates i n I960 and 19ol - 4 9 -D i p t e r a n s , represented by chironomids and c u l i c i d s , were the most abundant bottom organism i n a l l lakes except Bednorski and E n i d , where amphipods were most numerous. Dipterans accounted f o r 93 percent of the t o t a l bottom fauna i n Horseshoe Lake and 30 percent i n Bednorski Lake. The remaining lakes had values between those of Horseshoe and Bednorski. Oligochaetes formed 22 percent of the bottom fauna numbers i n Hiawatha Lake. In Jim Smith and Hiawatha l a k e s , the only other n u m e r i c a l l y important bottom fauna components were Anisoptera and Zygoptera. D i p t e r a and amphipoda accounted f o r more than 78 percent of the benthic fauna numbers i n a l l lakes except Hiawatha, which had 59 percent. In Figure 9 the number or organisms are shown f o r each 10 f o o t stratum of the 9 lakes. The standing crops of bottom fauna by weight and number, showed a s i m i l a r trend. The d i s s i m i l a r -i t y was most evident i n Hiawatha, lazy, New and Jim Smith l a k e s , which; had moderate numbers of d a m s e l f l i e s and d r a g o n f l i e s represented. Comparisons of the number of bottom organisms per dredging i n the 0 - 2 0 foot r e g i o n , with the number i n the r e g i o n over 20 f e e t , gave a s i g n i f i c a n t " t " value (p < 0.05) i n Jim Smith, Bednorski and Lazy l a k e s . No s i g n i f i c a n t d i f f e r e n c e between depth zones was i n d i c a t e d i n other lakes. - 50 -HIAWATHA KIAK HO HORSESHOE NEW J I M SMITH BEDNORSKI LAZY L I L L I A N ENID Figure 9. ro 20 10 20 30 40 10 20 SO 40 SO IO 20 30 40 IO 20 30 20 E 30J 40 SO IO 20 30 40 IO 20 40S so E O IOOO 200 0 3 0 0 0 4 0 0 0 SOOO 6 0 0 0 AVERAGE NUMBER OF BOTTOM ORGANISMS PER SQUARE METER Average number of a l l bottom organisms taken per square meter from bottom dredge samples. Numbers along the ordinate represent the maximum depth of each 10 foot stratum sampled. - 51 -F i s h Rainbow trout (Salmo g a i r d n e r i ) , Eastern Brook t r o u t ( S a l v e l i n u s f o n t i n a l i s ) , and Cutthroat t r o u t (Salmo d a r k ! lewisi,) were the major species taken i n g i l l nets from the 9 l a k e s . Northern squawfish (Ptychocheilus oregonense) Were present i n Bednorski Lake and longnose suckers (Catostomus catostomus) i n Hiawatha Lake. Largescale suckers (Catostomus macrocheilus) have a l s o been observed i n Hiawatha Lake. Bednorski and Lazy lakes also had redside s h i n e r s (Richardsonius b a l t e a t u s ) represented i n g i l l net samples. Shapley (personal communication) stated that suckers had gained entry to Kiakho Lake but none were taken i n g i l l nets during t h i s study. Table X I I I gives the number and weight of the various species taken i n an overnight g i l l net set. No weights are given f o r the small number of redside shiners captured. Natural reproduction of t r o u t occurs i n Bednorski, L i l l i a n and Kiakho l a k e s . With the exception of Bednorski and Kiakho, a l l the lakes are stocked by the F i s h and Game Branch of the Department of 'Recreation and Conservation. Enid and New lakes are known to have part of t h e i r f i s h populations "winter k i l l e d " and such a phenomena was observed during the s p r i n g of 1961. Bednorski Lake had the greatest f i s h biomass (Table X I I I ) . I f the lakes were ranked on the catch of trout alone, Bednorski would place s i x t h out of nine. A l l the remaining lakes would maintain the same p o s i t i o n i n the rank order regardless of species separation. TABLE X I I I Number and weight of f i s h taken i n a g i l l net set i n each lake during June and J u l y of 1961 Number Weight Total Weight Lake Species of F i s h (kg) . (kg) Bednorski Raiflfbow t r o u t Northern Squawfish Redside Shiner 4 58 3 2.84 11.79 14.63 Horseshoe Rainbow t r o u t Eastern Brook trout 19 1 . 11.20 1.03 12.23 Kiakho Cutthroat t r o u t 33 7.84 7.84 L i l l i a n Rainbow t r o u t Eastern Brook t r o u t 6 28 0.60 4.70 5.30 Jim Smith Rainbow trout 6 4.50 4.50 Hiawatha Rainbow tr o u t Longnose Suckers 7 3 3.17 3.17 New Rainbow trout 8 1.93 1.93 Enid Eastern Brook t r o u t 2 0.84 0.84 Lazy Rainbow t r o u t Redside Shiners 3 3 0.68 0.68 - 53 -Hypolimnion Oxygen D e f i c i t In t h i s i n v e s t i g a t i o n , the hypolimnetic oxygen d e f i c i t was c a l c u l a t e d f o r a l l lakes except Horseshoe. Horseshoe Lake was omitted because,; of i t s extensive water l e v e l f l u c t u a t i o n at the time of summer temperature s t r a t i f i c a t i o n . Summer stagnation was taken as J u l y 25 to August 11 of I960. The period of stagnation was estimated on the b a s i s of the time of i c e cover removal i n i 9 6 i . ; Oxygen concentrations were not always taken at the point of thermocline formation. Therefore, the oxygen concen-t r a t i o n j u s t above the thermocline was used i n the c a l c u l a t i o n of d e f i c i t s . C a l c u l a t i o n of the oxygen d e f i c i t was based on the/use of the hypolimnion l a y e r , i n which the thermocline was i n c l u d e d . The upper plane of the thermocline, at the time of summer st a g n a t i o n , was 15 f e e t i n most l a k e s . The a r e a l absolute hypolimnetic oxygen d e f i c i t s (Table XIV) r e l a t e d to the lake s u r f a c e , show some v a r i a t i o n among lakes (0.0013 to 0.0270 cc/cm 2/day). The a r e a l d e f i c i t s r e l a t e d to the hypolimnion surface e x h i b i t a l a r g e r range (0.0054 to 0.0346 cc/cm / day). Mean h y p o l i m n i a l oxygen d e f i c i t s (Table XIV) ranged from 0.022 to 0.049 c c / l i t e r / d a y . The lakes w i t h the highest mean d e f i c i t s a l s o had the smallest volumes. Only Bednorski Lake would be c l a s s i f i e d as e u t r o p h i c , according to the l i m i t set by Hutchinson (1957) of 0.033 mg/cm /day. The reason f o r the low values of oxygen d e f i c i t was probably due to the depths of the lakes under c o n s i d e r a t i o n , as w e l l as the f a c t that photosynthesis probably takes place i n the hypolimnion. Deevey (1940) gave s i m i l a r low values f o r Oxygen d e f i c i t s .in shallow lakes i n Connecticut. - 54 -TABLE XIV Absolute a r e a l and mean hypolimnetic oxygen . d e f i c i t s D e f i c i t to D e f i c i t to Mean Lake Hypolimnign surface E p i l i m n i o n surface D e f i c i t cc/om /day cc/cm /day c c / l i t e r / d a y Hiawatha 0.0054 0.0024 0.049 Kiakho 0.0188 0.0084 0.040 New 0.0102 0.0026 0.029 Jim Smith 0.0057 0.0037 0.033 Bednorski 0^0346 0.0270 0.047 Iiazy 0.0061 0.0013 0.039 L i l l i a n . 0.0055 0.0023 0.022 Enid 0.0054 0.0019 0.031 - 55 -Sedimentation Rates Between August 30 and September 1 of 1961, sedimentation c o l l e c t o r s ( P a t a l a s , personal communication) were suspended i n each lake f o r a 10 day p e r i o d . Where p o s s i b l e , the depth of suspension coincided w i t h the lower l i m i t of the thermocline. Since none of the lakes examined contained l a r g e amounts of i n o r g a n i c matter i n suspension, the c o l l e c t o r s should trap organic d e t r i t u s passing through the. e p i l i m n i o n . No allowance can be made f o r m a t e r i a l which i s o x i d i z e d w i t h i n the e p i l i m n i o n . The so c a l l e d "plankton r a i n " would be c o l l e c t e d the sediment c o l l e c t o r i n p r o p o r t i o n to the amount to be deposited on the l a k e bottom. Table XV gives the sediment volumes obtained from a 10 day suspension of the c o l l e c t o r i n each l a k e . Lazy, Bednorski and Kiakho lakes are outstanding as having the greatest sedimentation r a t e s . The remaining lakes have s i m i l a r q u a n t i t i e s of sediment. Ranking of the Lakes by B i o l o g i c a l and Other F a c t o r s I n d i c a t i n g P r o d u c t i v i t y . The method of ranking b i o l o g i c a l and other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y was the same as used f o r p h y s i c a l and chemical f a c t o r s . Ranking of the 9 lakes by b i o l o g i c a l f a c t o r s or standing crops of plankton, bottom fauna and f i s h i s given as Table XVI. Bednorski Lake was assigned a high rank f o r plankton, bottom fauna and f i s h whereas New and Enid lakes ranked low f o r the same standing crop estimates. Lakes other than those already mentioned did not show consistancy i n rank f o r a l l 3 f a c t o r s . - 5 6 -Table XVII l i s t s the i n d i v i d u a l and mean rank f o r other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y . No general r e l a t i o n s h i p appears evident except perhaps that Hiawatha Lake was somewhat consista n t i n having a low rank by a l l 3 p r o d u c t i v i t y measurements. - 57 -TABLE XV Sediment volumes as measured from sediment c o l l e c t o r s suspended at a s p e c i f i e d depth i n each lake f o r a 10 day pe r i o d i n September, 196-1 Lake Sediment Volume (cc) Depth of Suspension ( f t ) New 0.3 33 Enid 0.2 18 L i l l i a n 0.3 32 Jim Smith 0.3 23 Hiawatha 0.4 14 Kiakho 1.2 23 Bednorski 1.9 30 Horseshoe 0.5 22 Lazy 3.3 35 TABLE XVI Ranking of the 9 lakes by b i o l o g i c a l f a c t o r s Quantity of Lake Plankton Hiawatha 9.0 Kiakho 5.3 Horseshoe 5.3; New 0.1 Jim Smi th 2.7 Bednorski 8.4 Lazy 4.8 L i l l i a n 0.0 Enid 0.9 Quantity Quantity of of Bottom Fauna F i s h Mean Rank 0.0 1.6 3.5 9.0 4.6 6.3 4.8 7.4 5.8 4.2 0.8 1.7 3.2 2.4 2.8 6.3 9.0 7.9 2.5 0.0 2.4 5.3 2.9 2:7 3\8 0.1 1.6 TABLE XVII Ranking of the 9 lakes by other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y Mean Hypolimnion Gross Primary Sedimentation Lake D e f i c i t P r o d u c t i v i t y Rates Mean Rank Hiawatha 0.0 0.0 0.6 0.2 Kiakho 4.1 1.5 2.9 2.8 Horseshoe 0.0 0.9 0.4 New 1.5 1.9 0.0 1.1 Jim Smith 0.1 1.6 0.3 0.7 Bednorski 9.0 0.0 5.0 4,7 Lazy 0.2 2.4 9.0 3.9 L i l l i a n 0.0 9.0 0.3 3.1 Enid 0.0 8.0 0.0 2.7 - 6 0 -DISCUSSION While i t may be true that mean depth, climate or some edaphic f a c t o r i s s i n g l y most important i n i n f l u e n c i n g p r o d u c t i v i t y i n a broad geographic area (Rawson (1955) and Northcote and L a r k i n , 1956), a multitude of f a c t o r s by t h e i r i n t e r a c t i o n modify the •. primary c o r r e l a t e s of p r o d u c t i v i t y when lakes w i t h i n a r e s t r i c t e d geographic r e g i o n are examined. Few studi e s have been undertaken using lakes which were morphometrically 'similar and which were s i t u a t e d w i t h i n a r e s t r i c -ted geographical a r e a . Reimers et a l . (1955) examined 10 lakes w i t h i n a r e s t r i c t e d region and found almost complete disagreement of p h y s i c a l and chemical f a c t o r s with standing crops of i n v e r t e b r a t e s . Trout growth was r e l a t e d to q u a n t i t i e s of d i s s o l v e d n u t r i e n t s i n the study by Reimers et a l . (1955). On the other hand, Rawson (I960) obtained good agreement of f i v e physical f a c t o r s (T.D.S. included) with the "score" of three b i o l o g i c a l f a c t o r s . The lakes examined by Rawson (I960) were widely scattered through-out a. l a r g e area of Saskatchewan and e x h i b i t e d greater morphometric d i f f e r e n c e s than lakes i n t h i s study. Ranking of the study lakes by p h y s i c a l , chemical and b i o l o g i c a l c o n d i t i o n s , as w e l l as other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y , i s given i n Table X V I I I . Complete agreement of physical, or chemical r a n k i n g w i t h b i o l o g i c a l r anking was not evident. There was al s o disagreement between combined p h y s i c a l and chemical ranks with e i t h e r b i o l o g i c a l f a c t o r s or other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y . Mean summer plankton volumes were r e l a t e d to mean h y p o l i m n i a l oxygen d e f i c i t s as evident from Fi g u r e 10. - 61 -Standing crops of plankton, bottom fauna and f i s h i n 9 lakes w i t h i n the Southern Rocky Mountain Trench f a i l e d to c o r r e l a t e with a wide range of T.D.S. A l s o , mean depth and ep i l i m n i o n temperatures were not c o r r e l a t e d with standing crops of organisms. However, a p l o t of perimeter to area r a t i o s versus mean plankton volumes ( P i g . 11) r e s u l t e d i n a r e g r e s s i o n of 0.636 (p 0.05-0.10). Lazy and i n p a r t i c u l a r Horseshoe Lake, have major seasonal water l e v e l f l u c t u a t i o n s which contributed to the r e s p e c t i v e low to moderate standing crops of bottom organisms. Gammarids were not represented i n bottom dredge samples i n e i t h e r l a k e . L a r k i n and Northcote (1958) mention the p o s s i b l e e f f e c t on l i t t o r a l bottom organisms r e s u l t i n g from appreciable water l e v e l f l u c t u a t i o n . Grimas (1961) stated that Gammarus sp. were h e a v i l y reduced w i t h water l e v e l f l u c t u a t i o n i n Lake B l a s j b n , Sweden. Rooted macrophytes would l i k e w i s e be in f l u e n c e d by water f l u c t u a t i o n s which may have accounted f o r t h e i r absence from Horseshoe and Lazy lakes. U-shaped lake basins w i t h narrow l i t t o r a l zones were c h a r a c t e r i s t i c of Jim Smith, Bednorski and Lazy lakes ( P i g . 4). These were the only lakes where s i g n i f i c a n t d i f f e r e n c e s were observed between the average num.ber of bottom organisms per dredging, above and below 20 f e e t i n depth. New and Hiawatha l a k e s , with l a r g e areas l e s s than 20 f e e t i n depth * had abundant aquatic p l a n t s . The importance of macrophytes to the production of bottom fauna has been s t r e s s e d by Rawson (1930), B a l l (1948) and more r e c e n t l y Wohlschlag (1950). I t was true that lakes with numerous aquatic p l a n t s had a greater TABLE XVIII Ranking of the lakes by p h y s i c a l , chemical and b i o l o g i c a l . c h a r a c t e r i s t i c s as w e l l as other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y P h y s i c a l Other Factors and " I n d i c a t i n g P h y s i c a l Chemical Chemical B i o l o g i c a l P r o d u c t i v i t y . Hiawatha 6.2 Enid 7.6 Enid 5.6 Bednorski 7.9 Bednorski 4.7 Horseshoe 5.0 Bednorski 4.1 Kiakho 4-0 Kiakho 6.3 Lazy 3-9 Kiakho 4*2 Kiakho 3.8 Hiawatha 3«9 Horseshoe 5.8 L i l l i a n 3.1 Bednorski 5.8 Jim Smith 3.4 Bednorski: 3«9 Hiawatha 3.5 Kiakho 2.8 L a z y . : - 3*7 New 3.3 Horseshoe 3.8 Jim Smith 2.8 Enid 2.7 Enid 3.6 L i l l i a n 3.0 Jim Smith 3.3 L i l l i a n 2.7 New 1.1 Jim Smith 3.3 Horseshoe 2.7 Lazy 2.9 Lazy 2.4 Jim Smith 0.7 New 2.3 Lazy 2.2 New 2.8 New 1.7 Horseshoe 0.4 L i l l i a n 2.1 Hiawatha H 6 L i l l i a n 2.5 Enid 1.6 Hiawatha 0.2 - 63 -o 10 2 0 30 4 0 so A V E R A G E S U M M E R P L A N K T O N V O L U M E C C C . / ^ I 3 ) F i g u r e 10. R e l a t i o n s h i p of average summer (June-August, I960) plankton volumes (cc/nr) to mean hy p o l i m n i a l oxygen d e f i c i t s ( c c / l i t e r / d a y ) f o r 9 study lakes i n I960 and 1961 - 64 -v a r i e t y of bottom fauna ( P i g . 8) than did lakes w i t h sparse aquatic f l o r a . Lakes w i t h abundant macrophytes d i d not have large standing crops of bottom fauna. The l u s h aquatic vegetation of New and Hiawatha lakes was r e s p o n s i b l e f o r oxygen l o s s during winter stagnation ( P i g . 7 ) . P a t a l a s (I960) mentioned lack of oxygen as a f a c t o r l i m i t i n g a v a i l a b l e areas f o r bottom fauna. Hiawatha Lake had a low standing crop of bottom fauna while New Lake was moderate f o r bottom fauna but had winter f i s h m o r t a l i t y r e s u l t i n g from low oxygen concentrations. Although few aquatic p l a n t s were present i n Enid Lake, a low standing crop of f i s h r e f l e c t e d the r e s u l t s of w i n t e r - k i l l . The l i m i t e d q u a l i t a t i v e bottom fauna of Enid Lake. ( F i g . 8) was due to oxygen stagnation. L a r k i n ( i n press) stated that a small surface to volume r a t i o may mean l e s s heat per u n i t of volume from c i r c u l a t i o n . This could have been the main reason f o r low epilmnion temperatures i n Bednorski and L i l l i a n lakes and high temperatures i n Hiawatha Lake. Rawson (1942),and Northcote and L a r k i n (1956) have shown a s i g n i f i c a n t r e l a t i o n s h i p between summer e p i l i m n i o n temperatures and standing crops of plankton. The highest mean e p i l i m n i o n temperature was associated with plankton abundance i n Hiawatha Lake and the lowest temperature with meagre plankton i n L i l l i a n Lake. Perimeter to area r a t i o s appeared r e l a t e d to mean summer plankton volumes ( P i g . 10). Length of growing season or the l a r g e volume per u n i t area may mask the e f f e c t s of " l i t t o r a l development!' on standing crops of organisms. The l i t t o r a l zone should e x h i b i t the greatest p r o d u c t i v i t y because of the 20 P E R I M E T E R / A R E A F i g u r e 11. R e l a t i o n s h i p of the r a t i o of perimeter,to area to average plankton volumes (cc/nr) f o r the 9 study lakes i n 1960 and 1961 - 66 -overlap of the productive upper and lower surfaces at t h i s p o i n t . B a l l and Hayne (1952) recorded s i g n i f i c a n t changes i n bottom fauna with removal-of the f i s h p o pulation. S i g n i f i c a n t l y lower- numbers of bottom organisms per dredging were present i n the l i t t o r a l zone of Bednorski an,d Lazy lakes. Since both lakes contain redside s h i n e r s which frequent the l i t t o r a l zone, the f a u n a l r e d u c t i o n may have r e s u l t e d from g r a z i n g by f i s h . The l i m i t e d measurement of f i s h biomass i n t h i s study i s somewhat influ e n c e d by the species complex w i t h i n lakes (Table X I I I ) and the occurrence of n a t u r a l reproduction i n some lakes and not others.' T h e 3 lakes having n a t u r a l reproduction of f i s h , namely Bednorski, Kiakho and L i l l i a n , ranked as 3 out of the 4 highest by f i s h biomass i n an overnight g i l l net set. The s i z e and nature of the drainage area (Table I I I ) was important toi the p r o d u c t i v i t y of Bednorski, L i l l i a n and Enid l a k e s . The l a r g e drainage area of Bednorski was composed of calcareous sedimentary deposits ( F i g . 2) which undoubtedly supplied the lake with water r i c h i n n u t r i e n t s . S o i l s surrounding Enid and L i l l i a n ' l a k e s were low i n phosphates and t h i s could have con t r i b u t e d to low to moderate plankton crops. Without more extensive sampling i t i s d i f f i c u l t to say whether faunal d e f i c i e n c i e s e x i s t w i t h i n the lakes s t u d i e d . The absence of Gammarus sp. from some lakes has already been mentioned. Major groups such as the Hemiptera were not taken i n dredge:samples. Furthermore, the presence of more than 1 f i s h species would account f o r d i f f e r e n c e s i n f i s h biomass. Gross primary p r o d u c t i v i t y (Light and Dark Bottle), estimates appeared c o r r e l a t e d with t o t a l d i s s o l v e d s o l i d s ( F i g . 12) - 67 -Fi g u r e 12. R e l a t i o n s h i p of t o t a l d i s s o l v e d content (ppm) to gross primary p r o d u c t i v i t y (mgC/irr/day) a s measured by L i g h t and Dark B o t t l e techniques during the period of 27 June - 8 J u l y , 1961 - 68 -but not with standing crops of plankton, bottom fauna or f i s h . The absence of lakes from the mid-range of T.D.S. leaves some doubt about t h i s c o r r e l a t i o n . Thus, although t o t a l d i s s o l v e d s o l i d s may be c o r r e l a t e d w i t h p r o d u c t i v i t y throughout B r i t i s h Columbia, w i t h i n the l i m i t e d geographic region of the Southern Rocky Mountain Trench, a number of f a c t o r s i n t e r a c t to modify primary i n f l u e n c e s of p r o d u c t i v i t y i n a most complex manner. - 69 -SUMMARY 1. Standing crops of plankton, bottom fauna or f i s h were not c o r r e l a t e d w i t h a wide range (50 to 1460 ppm) of t o t a l d i s s o l v e d s o l i d s i n 9 lakes w i t h i n a r e s t r i c t e d geographical region of B r i t i s h Columbia. 2. Average summer plankton volumes appeared to be r e l a t e d to mean h y p o l i m n i a l oxygen d e f i c i t s . 3. The r a t i o of perimeter to area of lakes showed a r e l a t i o n s h i p to average plankton volumes. 4. T o t a l d i s s o l v e d s o l i d content was c o r r e l a t e d with gross primary p r o d u c t i v i t y estimates as measured by Li g h t and Dark B o t t l e techniques. 5. Ranking of lakes by p h y s i c a l and chemical i n d i c e s of pro-d u c t i v i t y d i d not agree with ranking by b i o l o g i c a l or other f a c t o r s i n d i c a t i n g p r o d u c t i v i t y . 6. A number of f a c t o r s by t h e i r i n t e r a c t i o n were found to modify the expression of primary c o r r e l a t e s of p r o d u c t i v i t y w i t h i n a r e s t r i c t e d geographic r e g i o n of B r i t i s h Columbia. - 70 -LITERATURE CITED B a l l , R.C. 1948. R e l a t i o n s h i p between a v a i l a b l e f i s h food, f e e d i n g h a b i t s of f i s h and t o t a l f i s h production i n a Michigan l a k e . Mich. State C o l l . Agr. Exp. Sta. Tech. B u l l . , 206:59 p. B a l l , R.C, and p.W. Hayne. 1952. E f f e c t s of the removal of the f i s h population on the f i s h - f o o d organisms of a lake. Ecology,35: 41-48. Chapman, J.D. 1952. The climate of B r i t i s h Columbia. Trans. . F i f t h B.C. Nat. Res. Conf., 8-54 p. Deevey, E.S. 1940. L i m n o l o g i c a l studies i n Connecticut. V. A c o n t r i b u t i o n to r e g i o n a l limnology. Am. J . S c i . , 238: 714-741. Duncan, David B. 1955. M u l t i p l e range and m u l t i p l e F t e s t s . B i o m e t r i c s , 11: 1-42. E b e r l e y , W.R. 1959... The metalimnetic oxygen maximum i n Myers Lake. Invest. Indiana Lakes and Streams, 5: 1-46. Grimas, U l f . 1961. The bottom fauna of n a t u r a l and impounded lakes i n northern Sweden (Ankarvattnet and B i a s j o n ) . I n s t . Freshwater Res. Sweden, 42: 183-237. Hutchinson, G.E. 1938. On the r e l a t i o n between the oxygen d e f i c i t and the p r o d u c t i v i t y and typology of l a k e s . I n t . Rev. Hydrobiol. Hydrogr., 36: 336-355. 1957. A t r e a t i s e on limnology. V o l . I. Geography, physics and chemistry. John Wiley & Sons, New York 1015 p. K e l l e y , C.C., and P.N. Sprout. 1956. S o i l survey of the upper Kootenay and E l k r i v e r v a l l e y s . B.C. Dept. A g r i c . S o i l Survey r e p t . , 5: 99p. K e l l e y , C.C. , and W.-D. Holland. 1961. S o i l survey of the upper Columbia r i v e r v a l l e y . B.C. Dept. A g r i c . S o i l Survey r e p t . , 7: 107 p. L a r k i n , P.A., and T.G. Northcote. 1958. Factors i n lake typology i n B r i t i s h Columbia, Canada. Verh. i n t e r n . Ver. Limnol., 13: 252-263. L a r k i n , P.A. ( i n p r e s s ) . Canadian Lakes. Verh. i n t e r n . Ver. L i m n o l . , 1 5 . - 71 -Moyle, J.B* 1946. Some i n d i c e s of lake p r o d u c t i v i t y . Trans. . Am. P i s h . S o c , 76: 322-334. Naumann, E i n a r . 1932. Grundzuge der regionalen Limnologie. Die Binnengewasser, 11: 176 p. Northcote, T.G. , and P.A. L a r k i n . 1956. Indices of p r o d u c t i v i t y i n B r i t i s h Columbia lakes. J . P i s h . Res. Bd. Canada, 13: 515-540. •.. \ Ohle, W. 1934. Chemische and p h y s i k a l i s c h e Untersuchungen norddeutscher Seen. Arch. H y d r o b i o l . , 26: 386-464. P a t a l a s , K. I960. Stosunki termiczne i tlenowe oraz prezezroczy-stosc wody w 44 j e z i o r a c h o k o l i c Wegorzewa. :. Roczn. Nauk Rbln., 77, B, 1: 105-222. Rawson, D.S. 1930. The bottom fauna of Lake Simcoe and i t s r o l e i n the ecology of the lake. Univ. Toronto S t u d i e s , B i o l Ser* No. 34. Ontario P i s h . Research Lab., Publ. 40: 1-183. 1942. A comparison of some large a l p i n e lakes i n Western Canada. Ecology, 23: 143-161. 1952. Mean depth and the f i s h production of large l a k e s . Ecology, 33: 513-521. 1953. 'The standing crop of net plankton i n l a k e s . J . P i s h . Res. Bd. Canada, 10: 224-237. 1955. Morphometry as a dominant f a c t o r i n the p r o d u c t i v i t y of l a r g e lakes. Verh. i n t e r n . Ver. Limnol. 12: 164-175. I960. A l i m n o l o g i c a l comparison of twelve l a r g e lakes i n northern Saskatchewan. Limnol. Oceanogr., 5: 195-211. Reimers, Norman, J.A. Maciolek, arid E . P . " P i s t e f . 1955. L i m n o l o g i c a l study of the lakes i n Convict Creek B a s i n , Mono County, C a l i f o r n i a . U.S. P i s h and W i l d l . Serv., P i s h . B u l l . , 56: 437-503. R i c e , H..M.A. 1937. Cranbrook map-area, B r i t i s h Columbia. Geol. Surv. Canada, Dept. Mines and Resources, Ottawa, Memoir 207, 67 p. S c h o f i e l d , Stuart J . 1915. Geology of Cranbrook map-area, B r i t i s h Columbia. Geol. Surv. Canada, Dept. Mines and Resources, Ottawa, Memoir 76, 245 p. Stenton, C.E. I960. Ecology of the Yellowstone Cutthroat t r o u t (Salmo" G l a r k i i L e w i s i G i r a r d ) i n Kiakho Lake, B r i t i s h Columbia, M.Sc ."^Thesis, Dept. Z o o l . , Univ. B r i t i s h Columbia, 77p. - 72 — S t r i c k l a n d , J..D.H. I960. Measuring the production of marine phytoplankton. J . P i s h . Res. Bd. Canada, B u l l . , 122: 172 p. Strain, K.M. 1931. Feforvat n . A physiographic and b i o l o g i c a l , study of a mountain l a k e . Arch. Hydrobiol. , 22: 491-536. Thienemann, t A. 1927. Der Bau des Seebeckens i n seiner Bedeutung f u r den Ablauf des Lebens im Sec. Verhandl. Zool. Bot. G e s e l l . , 77: 87-91. Walker, J.P. 1926. Geology and mineral deposits of Windermere map-area, B r i t i s h Columbia. Geol. Surv. Canada, Dept. Mines and Resources, Ottawa, Memoir 148, 69 p. Ward,. P.J. 1957. Seasonal and annual changes i n a v a i l a b i l i t y of the a d u l t crustacean p l a n k t e r s of Shuswap Lake. I n t e r n . Pac. Salmon P i s h . Comm., Progr. Rept., 56 p. Weber, C.I. 1958. Some measurements of primary production i n Bast and West Okoboji l a k e s , Dickinson County, Iowa. Proc. Iowa Acad. S c i . , 65: 166-173. Welch, Paul S. 1948. L i m n o l o g i c a l Methods. B l a k i s t o n , P h i l a d e l p h i a . 581 p. Wohlschlag, D.E. 1950. Vegetation and i n v e r t e b r a t e l i f e i n a marl lake. Invest. Indiana Lakes and Streams, 3: 321-372. 

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