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Giant chromosomes, ecology, and adaptation in Chironomus Tentans Topping , Milton Stanlee 1969

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GIANT CHROMOSOMES, ECOLOGY, AND ADAPTATION .' IN CHIRONOMUS Tl'.NTANS by MILTON STANLEE POPPING B.A., U n i v e r s i t y o f K a n s a s , 1963 M.Sc. , U n i v e r s i t y o f B r i t : . s h C o l u m b i a , 1966 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR '.'HE DEGREE OF DOCTOR OF PHILOSOPHY i n t h e D e p a r t n e n t o f ZOOLOGY We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA O c t o b e r , 1969 In present ing t h i s t h e s i s in p a r t i a l f u l f i l m e n t of the requirements foi an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, | agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e for reference and study. I f u r t h e r agree tha permission for extensive copying of t h i s thes is f o r s c h o l a r l y purposes may be granted by the Head of my Department or by his representa t ives . It is understood that copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my wri t ten permiss ion . Department of Zoology  The U n i v e r s i t y of B r i t i s h Columbia Vancouver Canada Date . "28 October 1969 Chairman: Processor G.G.E. Scudder ABSTRACT E c o l o g i c a l adaptation has occupied a cen t r a l p o s i t i o n i n evolutionary theory for ove:: 100 years; however, s u r p r i s i n g l y l i t t l e i s know i n d e t a i l of i t s s i g n i f i c a n c e . This study was undertaken to determine i f chromosomal inversions of larvae of Chironomus tentans have any s e l e c t i v e significance i n nature and Lf they do, to define the r e l a t i o n s h i p between genetic adaptation, as r e f l e c t e d by the frequencies of the inversions, and the ec o l o g i c a l conditions to which the species mast be adapted. The procedure; used was (1) to define the habitats p o t e n t i a l l y a v a i l a b l e to C. tentans, (2) to define the e c o l o g i c a l conditions to which C. tentans i s adapted, and (3) to define the r e l a t i o n s h i p between inversion frequency and those e c o l o g i c a l conditions to which C. tentans has been found to be adapted. The envirmmental properties of 32 sal i n e lakes located i n south-central B r i t i s h Columbia were studied. The lakes d i f f e r e d i n t o t a l chemical concen-t r a t i o n , r a n g i i g from very fresh to four or f i v e times the concentration c f sea water. Measurements of temperature, conductivity of the water, t o t a l dissolved s o l i d s , and pH, and of concentrations of dissolved oxygen, sodivm, potassium, calcium, magnesium, carbonate, bicarbonate, c h l o r i d e , and sul f a t e were made during d i f f e r e n t seasons at d i f f e r e n t depths i n most of the lakes. Surface water samples were analyzed by a semi-quantitative technique for 28 trace elements. Selected samples of bottom muds were analyzed for substrate composition and percentage organic carbon content. Analysis of the environmental properties indicated that the chemical concentrations oi the lakes varied both seasonally and with depth. Sodium, magnesium, carbonate-bicarbonate, and s u l f a t e were the ions present at the greatest concentrations. i i The occurrence and abundance of larvae of C. tentans were determined with respect to the environmental properties of the lakes. C. tentans was found,to occur in only 17 of the 32 lakes studied and each of the 17 lakes had conductiviti.es within the range of from 500 to 4,500 micromhos. Further, C. tentans occuired only in the upper two meters of these lakes and was most abundant at. the shallower depths of i t s occurrence. Both occurrence and abundance of C. tentans were correlated primarily with chemical concen-tration, although other environmental factors also were found to be important. Analysis of the frequencies of certain inversions present in chromosome 1 indicated that, populations present in different environments may have significantly different inversion frequencies, that the frequencies did not vary seasonally and were extremely stable over time periods as long as eight years, and that the inversion frequencies of sub-divisions of populations livi n g within lskes do riot vary significantly, Although C. tentans was found to show considerable ecological adaptation to lake water chemistry, no obvious relationship between the frequencies of the inversions of chromosome 1 and lake water chemistry could hi detected. The studies of correlation of inversion frequencies with environmental properties and the results of a f i e l d experiment indicate th.it C. tentans is genetically adapted primarily to i t s co-occurrence with other chironomids. A lesser correlation with pH was also detected. Consequently, although genetic adaptation to environmental factors was found, that genetic adaptation was not to the tr.ajor differences which occurred between the lakes. The unique and major contributions of this study are the demonstration that the inversions of chromosome 1 have selective significance in nature and that not a l l ecological adaptation is accompanied by parallel genetic adaptation (as reflected by inversion frequency). The clear implication i s t h a t a l t h o u g h e c o l o g i c a l a d a p t a t i o n s may hs.ve e v o l u t i o n a r y s i g n i f i c a n c e , not a l l e c o l o g i c a l a d a p t a t i o n s do. E r r a t a Page 9- Line : 2 1 . For "polyehylene" read "polyethylene". Page 1 0 . Line 2 1 . For "Metier" read " M e t t l e r " . Page 11 . Line 1 1 . For "18 g of CaCl 2*2H 2 0 and 9 g of NaCl" read "18 g of CaCl2«2H 0 'and 9 g of . NaCl / 1". Page 1 2 . L i n e 1 2 . A f t e r LB add "which" t o read "LB 1 which was so a f f e c t e d " . Page 1 3 . L i n e 2. For "and the d r y i n g " omit "the" t o read "and d r y i n g " . Page 1 3 . Line 1 1 . For "Standard Methods" read "Standard Methods". Page 1 9 . Line 1 0 . For "yhe" read "the". Page 3 2 . Line .21 . For " f l o c o u l a n t " read " f l o c c u l e n t " . Page 3 5 . L i n e 2. For "ar" read "are". Page 4 1 . Line 2 2 . For "C. tentans." read " C. tentans". Page 4 2 . Line 1 8 . For "samples" read "sampled". Page 4 7 . L i n e 8 . For " l i e v e d " read " l i v e d " . Page 55- Add " 0 m " n e x t t o "NE OP CRESCENT" on r i g h t hand side of t a b l e . Page 6 5 . L i n e 2 7 . For"C. tentans" read "C. tentans". Page 7 2 . L i n e 1 1 . For " f l o c c u l a n t " read " f l o c c u l e n t " . Page 7 2 . Line 2 3 . A f t e r C. 'tentans omit "of popu l a t i o n s " . Page 8 3 . L i n e 18." For " i n v e r s i o n s " read " i n v e r s i o n " . Page 8 5 . L i n e 1. For"C. tentans" read "C. tentans". Page 9 6 . L i n e 5 . For "L" read " ] . " . Page 1 0 2 . Add "Dobzhansky, Th. 1 9 5 1 . Genetics and the O r i g i n of Species. 3 r d . Ed. r e v i s e d . Columbia B i o l o g i c a l S e r i e s . No. 1 1 . Columbia U. Press, New York." between second and 3 r d c i t a t i o n . iv TABLE OF CONTENTS PAGE Abstract .... i Table of Contents ... .... iv List of Tables v i List of Figures v i i i Acknowledgements' .... x I. Introduction 1 II. Study Area „ 5 III. The Lake Environments 7 A. Materials and Methods 7 1. Geographical, geological and climatic data 7 2. Morphometric data 7 3. Substrate composition 7 4. Organic carbon content of mud 8 5. Physico-chemical characteristics 9 a. Frequency and method of collection 9 b. Handling 9 c. Analytical methods .... 10 B. Results 13 1. The lakes and distribution . 13 2. Morphometry 14 3. Physico-chemical conditions 19 4. Substrate composition and organic carbon 32 C. Discussion 35 IV. Occurrence and Abundance of C. tentans 41 A. Methods of Determination of Occurrence 41 B. Results o 43 1. General distribution 43 2. Distribution in the study area 43 3. Abundance 48 C. Discussion 54 V. Inversion Polymorphism in Natural Populations of C. tentans with Respect to Environmental Factors 67 A. Introduction 67 B. Methods 69 V PAGE G. Results . o o ; . 70 1. Differences between lakes 70 2. Horizontal variation in lakes 72 3. Temporal variation 72 4. Effect of change in the biotic environment 76 D. Discussion 82 VI. DiscussiDn 95 VII. Literature Cited 101 VIII. Appendix I. Bathymetric maps of the lekes 104 IX. Appendic II. Chemical data collected from the lakes 131 X. Appendix III. Percentage of the total volume of each lake present in successive meter intervals of depth 152 v i LIST OF TABLES PAGE Table I Summaries of elevations of lakes and of •norphometric data „ 17 Table II Temperature, pH and dissolved oxygen in surface .and bottom waters of the lakes . . .. . 20 Table III Summary of average physico-chemical conditions present during different seasons 21 Table IV Comparison of trace elements present in samples of surface waters collected in May 1966 22 Table V Summary of the s t a t i s t i c a l significance of variation in certain physico-chemical properties . 23 Table VI Classification of the lakes according to the predominate anion(s) and caticn(s) present „. 29 Table VII Summary of partial correlation coefficients characterizing the relationships between certain physico-chemical properties of the lakes studied . . 31 Table VIII Substrate composition 33 Table IX Percentage of organic carbon present along sample transects in certain lakes and comparative measures at 0 m in other lakes 34 Table X . Seasonal variation in the conductivity of surface waters 38 Table XI Long term variation in the conductivity of surface waters 39 Table XII Occurrence of larvae of C. tentans with respect to water temperature, pH, minimum dissolved oxygen, and presence of hydrogen sulfide 50 Table XIII Occurrence of larvae of C. tentans in relation to percentage of organic carbon 51 Table XIV Observed abundances of larvae of C. tentans at sample lo c a l i t i e s and groupings used for the Kendall rank correlation coefficient analysis .-. . . 55 Table XV Correlation between the abundance of larvae of C. tentans and specific environmental factors .... 56 vii. PAGE Table XVI Percentage frequency, of inversions in chromo-somes 1 and 4 of C. tentans in 12 loc a l i t i e s in central British Columbia „ ............. 71 Table XVII Horizontal variation 73 Table XVIII Seasonal change in the percentage frequency of inversions in chromosomes 1 and 4 in populations occurring in Six Mile lake ana in Near Phalerope lake 74 Table XIX Changes in percentage frequency of inversions in chromosomes 1 and 4 over long periods of time ....... 77 Table XX Changes with time in the percentage frequency of inversions in chromosomes 1 and 4 after the introduction of eastern brook trout into Westwick Lake 78 Table XXI Summary of probabilities that differences in inversion frequencies observed during the sample periods are attributed to sampling error 81 Table XXII Summary of the partial correlation coefficients which describe the relationship between the frequency of 1 Rad and physico-chemical water properties 86 Table XXIII Comparison of partial correlation coefficients for the/relationship between the frequency of 1 Rad and certain environmental factors 88 Table XXIV Summary of step-wise multiple regression analysis ... 90 Table XXV Relative adaptive values of the karyotypes of 1 Rad and 1 Rade with respect to total environ-mental factors 97 Table XXVI Relative adaptive values of the karyotypes of 1 Rad and 1 Rade with respect to food 99 v i i i LIST OF FIGURES PAGE Figure 1 Climatic regions of British Coluuibia and selected climatic data 6 Figure 2 Lake l o c a l i t i e s in south-central British Columbia ... 15 Figure 3 Left. Distribution of lakes with regard to forest (shaded) and open grassland (clear). Right. Distribution of lakes.with regard to under-lying geology ,.„ 16 Figure .4 Relative composition of anions in and total ionic con-position of the lakes 25 Figure 5 Relative composition of cations in and total ionic composition of the lakes 26 Figure 6 Ionic diagrams showing examples of the six "chemical" types of lakes present in the study area 28 Figure 7 Conductivity as a function of ionic composition 40 Figure 8 World wide distribution of C. tentans 44 Figure 9 Occurrence of larvae of C. tentans with respect to chemical concentration of lake waters 45 Figure 10 Occurrence of larvae of C. tentans and of other species of chironomids with respect to depth ........ 46 Figure 11 Occurrence of larvae of C. tentans with respect to concentrations of the major ions 49 Figure 12 Occurrence of larvae of C. tentans in relation to substrate composition 52 Figure 13 Occurrence of larvae of _C. tentans at different depths in Westwick Lake during October 1966 in relation to larvae of other species of chironomids .. 53 Figure 14 Experimental analysis of survival of larvae of C. tentans in waters with different chemical concentrations at 5° C 59 Figure 15 Environmental factors potentially responsible for the absence of larvae of C. tentans from specific l o c a l i t i e s in lakes in which the species does occur . 63 Figure 16 Environmental factors potentially responsible for the absence of larvae of C. tentans from lakes 64 ix PAGE Figure 17 Drawings of chromosome 1: homozygDus for inversion 1 Rad, homozygous for inversion 1 Rade, and heterozygous for inversions 1 Rad and 1 Rade 68 Figure 18 Seasonal variation in frequencies of chromosomes 1 and 4 » 75 Figure 19 Localities sampled in Westwick Lake in order to analyze horizontal variation and to evaluate the effect of introduction of fish .. . 79 Figure 20 Scatter diagrams depicting the relationship between the frequency of the inversion of 1 Rad and the environmental factors indicated 87 X ACKNOWLEDGEMENTS I wish to express thanks to Professor^. A.B. Acton and G.G.E. Scudder for their assistance and cooperation throughout this study. I have benefited greatly from discussions with both. Also, thanks are extended to Professor C.V. Finnegan who has aided me: in many ways. The manuscript was read by Professors A.B. Acton, T.G. Northcotte, and G.G.E. Scudder. I am indebted to Professor G.W. Eaton who assisted in some of the s t a t i s t i c a l analyses of the data. I wish to express gratitude 'to the National Research Council of Canada for a Postgraduate Scholarship. I extend special thanks to m}- parents. 1 I. Introduction Since the times of Lamarck, Darwin, and Wallace, the problem of ecological adaptation has occupied a central position in evolutionary theory (Allee, Emerson, Park, Park, arid Schmidt, 1950). However, surprisingly l i t t l e infor-mation is available concerning how, in fact, genotypes respond to ecological conditions or about the relative importance of different environmental factors to evolution. Indeed, detailed knowledge of the relationship between distribution, abundance, and genotypic composition is v i r t u a l l y lacking for a l l animal species (see Ford 1964, 1965 for a review). While chromosomal polymorphism in various species of Drosophila has been studied extensively by Dobzhansky and others (Dobzhansky, 1951; Wallace, 1968) and a l i t t l e is known about the relationship between genetic differences among populations and the distributions of these populations, almost nothing is known about the ecological significance of these genetic differences, since the biology of most species of Drosophila in the wild is l i t t l e known. Alternatively, tho.'ie species for which good distribution and abundance data are available usually do not lend themselves to genetic analysis. However, those species which possess giant chromosomes present a unique opportunity for obtaining by direct observation genotypic information, unmodified by the environment except by natural selection. Chironomus i:entans Fabricius was selected for study (1) because i t contains giant chromosomes which make i t amenable to cytogenetic study, (2) because much work has already been done on i t s population genetics and inver-sion polymorphism (Acton, 1957, 1959,1962), (3) because the l i f e cycle is known and the larvae live in known and definable habitats and (4) because 2 • i t is relatively common in central British Columbia. C. tentans :".s a non-biting midge in which immature stages are aquatic and live in the bottom muds of lakes and sluggish streams. The short lived adults are aeria 7. and females deposit their eggs in water. The species over-winters as larvae. The larvae are characteristically blood red, contain hemaglobin, and possess giant chromosomes. The larvae have been described by Sadler (1935) and Acton (1955,1956), the pupal exuvia by Shilova (see Palmen and Aho, 1966),and variation in the adults by Palmen and Aho (1966). The l i f e history of C. tentans has been studied by Sadler (1935). The species has been reported from North America (Acton, 1962; Townes, 1945) and Europe (see Fittkau, 1967 and Palmen and Aho, 1966). Within central British Columbia, larvae of C. tentans occur in relatively small lakes which differ primarily by chemical composition and concentration. The fact that the larvae are relatively long lived in comparison to the adults increases the probability that most of the natural selection to which the species must adapt occurs in the larval stage. In addition, the fact that inversions are presumably adapted to specific ecological conditions (see Swanson, 1957 and Ford, 1964, 1965), together tfith the fact that several species of Chironomus show considerable adaptation to saline environments (Neumann, 1961) suggest the possibility that populations of C. tentans might show dif f e r e n t i a l genetic adaptation to the different chemical concen-trations of the lakes, which in turn could be reflected by different inversion frequencies. Of course, genetic adaptation might occur without any vis i b l e change in the banding patterns of the giant chromosomes, but i f John and Lewis (1966) are correct (i.e. , that chromosomal, rather than gene variations are the key to differences in populations) then one might expect inversion frequency to be affected by such an important environmental factor as lake chemical concentration. The purpose of this study was to define the genetic adaptation of C. tentans to ecological conditions. The procedure adopted was (1) to define the habitat conditions potentially available to C„ tentans, (2) to define the ecological conditions to which C. tentans is adapted, and (3) to define the relationship between inversion frequency and the ecological conditions to which _C° tentans i s adapted. Ecological adaptation was evaluated by determining the relationship between occurrence and abundance, and natural environmental conditions. The effect of environmental factors on occurrence was evaluated by comparing the means and/or ranges of those conditions which the species tolerates with those which i t does not. If a particular environmental factor exceeds the range of that factor which i s tolerated or i f the means of the conditions which are tolerated differ significantly from those which are not, then that factor or those factors might be responsible for the absence of the species. The effect of environmental factors on abundance was evaluated by determing the correlation between abundance and the environmental factors. Ecological adaptation was determined in order to ascertain what individual . factors might be important as agents of natural selection and to determine the specificity of adaptation of the species. Genetical adaptation was evaluated by determing the correlation between the frequencies of certain inversions and environmental factors The relationship between the genetics and ecology of C. tentans was evaluated by comparing those factors to which i t was adapted ecologically with those factors to which i t was adapted genetically. The relative impor-tance of the different environmental factors w.is evaluated by inspection of the degree of correlation between inversion frequency and the environmental factors. The relationship between the genetics and ecology of _C. tentans 4 was studied i n order to evaluate the fitness of different populations and the f i t n e s s of p a r t i c u l a r chromosomal inversions to complex environmental conditions. ' Although primary attention was directed to the chemical differences of the environment, other aspects of the environment were studied as well. The environmental variables which were considered are (1) chemical concentration, ( 2 ) chemical composition, (3) pH, (4) temperature, (5) dissolved oxygen concentration (and to a lesser extent hydroger. s u l f i d e concentration), (6) substrate composition, (7) food a v a i l a b i l i t y , a n d (8) co-occurrence with other species of chironomids. 5 II. Study Area The study was carried out in the Cariboo and Chilcotin areas of central British Columbia. Of the five study areas selected, the Clinton and Kamloops areas are located on the Thompson Plateau in the Thompson River drainage, while the Chilcotin, Gang Ranch, and Springhouse areas are located on the Fraser Plateau in the Fraser River drainage. The Thompson and Fraser Plateaus are pi ysiographic areas wMch occur in the more inclusive Fraser River drainage (Holland, 1964). The stud}' areas occur at elevations ranging from 700 m in the south-east to 1,100 m in the north-west on a broad plateau that was once covered by Pleistocene ice and is now overlain by glacial d r i f t . In general, the north-western half of the area (Fraser Plateau) is underlain by fl a t or gently dipping Miocene: or pliocene olivine basalt, while the south-eastern half (Thompson Plateau) is underlain by a diversity of rock formations- which include sedimentary anci volcanic, formations of Palaeozoic age intruded by granitic rock and f l a t or gently dipping early Tertiary (Eocene) lavas (Holland, 1964). The stud}1 areas occur in a relatively uniform climatic region known as the South West Interior Plateau (Figure 1) which is characterized by low precipitation, low average annual temperatures, and large fluctuations in seasonal and daily summer temperatures. About 70% of the precipitation f a l l s as rain. Summaries of temperature and precipitation data for the study area and surrounding areas are given in Figure 1. Generally, the vegetation of the study area is typical parkland or savannah and iv, characterized by open grassland and mixed stands of conifers and aspen (see Munro (1945) and Munro and Cowan (1947) for classifications and more complete discussions of the biota of this and surrounding areas). 6 Figure 1. Climatic regions of British Columbia and selected climatic data (taken from the British Columbia Atlas of Resources. B.C. Natural Resources Conference, 1956). Mr A N TEMPERATURE JANUARY IN DEGREES CENTIGRADE > o r i o ro -5 r i -IOTO-15 15 To-20 CLIMATIC R E G I O N S COAST WEST 1 OUTER 2 INNER 3 HEAD OF FIORD 1 SOUTHWEST 1 VALLEYS 2 PLATEAUS SOUTHEAST 1 VALLEY5 2 PLATEAUS CENTPAL 7 S p e c i f i c a l l y , the vegetation south of Six Mile lake and around Kamloops tends more to dry forest and d i f f e r s p r i n c i p a l l y i n the addition of sage brush. Each of the f i v e study areas i s i s o l a t e d from the other by forest. Therefore, one might expert d i s p e r s a l between the areas to be impaired, III. The Lake Environments A. Materials and Methods 1. Geographical, geological and c l i m a t i c data. Climatic data were taken from the B r i t i s h Columbia Atlas of Natural Resources, Geological and Geographical data were taken from C c c k f i e l d (1948) and from regional geology maps. Where ad d i t i o n a l references were used, t h e i r citat:.ons are given. 2. Morphometric data. In 1966, depth soundings were taken with a Furuno 200-kHf:/sec echo sounder. Bathymetric maps of these lakes were constructed using the soundings and are included i n Appendix I. Areas of each of the lakes were determined p l a n i m e t r i c a l l y from a e r i a l photos of known magnification and then extrapolated to drawn maps. Subseauently, areas of contours at meter i n t e r v a l s (i..e. , 1 m, 2 m, etc.) were determined p l a n i m e t r i c a l l y using the surface area of the 0 m contour taken from the a e r i a l photo as the point of reference and volumes of water l y i n g between the meter i n t e r v a l s were calculated therefrom. 3. Substrate composition. Selected determinations of substrate composi-t i o n were made during October, 1967 i n some of the lakes. The substrate composition determinations! were made by c o l l e c t i n g raw mud 8 samples with a 15 x 15 cm Ekman dredge, returning them to the laboratory for separation into various size fractions by washing through seives of known pore size e.nd then drying at 105° C for 24 hrs. The mud samples were separated into the following six size fractions: (1) greater than 1.98 mm (granule); (2) 1.98 - 0.83 mm (very coarse sand and coarse sand); (3) 0.83 - 0.59 mm (coarse sand); (4) C. 59 - 0.42 mm (coarse sand and medium sand); (5) 0.42 - 0.15 mm (medium sand and fine sand); and (6) less than 0.15 mm (very fin i sand and s i l t ) . Analyses of substiate composition were performed for correlation with abundance and inversion genotypes of C. tentans Hence, samples were taken at meter intervals in depth from the shores to the mid depths of the lakes along single transect lines which were located so that they would pass through areas known to be inhabited by C. tentans. Comparative samples of substrate composition at 0 m depth were taken in some of the lakes not containing C. tentans. Organic carbon content of mud. During October, 1967, mud sample:; were collected with a 15 x 15 cm Ekman dredge for organic carbon analysis. After the samples were collected they were transferred to pint jars and transported to the laboratory. Muds were not refrigerated during transportation, but were dried immediately upon return. As in the collections for characterization of substrate composition, analyses of organic carbon were performed for correlation with the abundance, occurrence and inversion genotypes of C. tentans. The locations of the sample sites are as described for substrate composition. The percent organic carbon content of bottom mud was determined during a single season. Organic carbon content was determined by the method given in Diagnosis and Improvement of Saline and Alka l i Soils. 9 Agriculture Handbook No. 60. U.S.D.A. The samples were prepared by drying raw mud at 100° C and then proceeding as directed by the method given above. 5. Physico-chemical characteristics. Lake waters were analyzed in successive seasons for pH, dissolved oxygen, temperature, conductivity, total dissolved solids, sodium, potassium, magnesium, calcium, carbonate, bicarbonate, chloride, and sulfate. In addition, the presence of trace elements in surface waters was determined during a single season, a. Frequency and method of collection. In total, water'samples were collected from 32 lakes for analysis. During the spring and mid-summer of 1966, samples were collected from 28 of the lakes and of these lakes samples were taken again from five of the lakes during the winter of 1967. Samples were collected from over the deepest point of each lake at meter intervals. Samples were taken to within the nearest meter above the lake bottoms. The samples were collected with a two-liter Van Dorn bottle and then transferred to l i t e r polyethylene bottles. Ice samples were also collected for analysis during winter. Of the remaining four lakes, Clinton was sampled during mid-summer, 1966.and the remaining three were sampled during the spring of 1967. These samples were collected by immersing a l i t e r polyehylene bottle below the surface of the water at the margins of the lakes. Portions of the single samples collected from the four lakes and portions of the surface samples collected from the other lakes during the spring were analyzed for trace element. b. Handling. Temperature, pH, dissolved oxygen, carbonate and bicarbonate were determined in the f i e l d . The remainder of the 10 analyses were performed in the labcratory. The samples were stored in polyethylene bottles at 4° C anc the analyses performed as soon as possible. Most of the samples vere analyzed within four months, although some of the samples were kept for about a year before analysis. c. Analytical methods. Temperature profiles were determined by a YSI Yellow Springs thermister thermometer to + 0.1° C. oH was determined with a Radiometer expanding scale pH meter to the nearest + 0.03 pH units. Dissolved oxygen was determined by the standard unmodified Winkler method. The analyses "appeared" to work normally in lakes with conductivities less than 5,000 micromhos, but the r e l i a b i l i t y of these data cannot be attested to. Conductivities were determined in the laboratory with a line-operaued Radiometer conductivity meter at a standard temperature of 25° C, Some f i e l d measurements were also taken with a Solubridge conductivity meter. "otal dissolved solids were determined by evaporating 100 ml of lake water, which had been filtered through a 0.45 micron HA millipore membrane,over a steam bath and then drying the salt residue for 24 hrs at 105° C. The dried salt residues were weighed on a Metier electric balance to the nearest 0.0001 g. Hydrogen sulfide was estimated by a semi-quantitative method supplied by Hach Chemicals, Ames, Iowa. 11 Concentrations of sodium and potassium were determined by flame-emission photometry using a Zeiss flame-photometer. Both ions were determined with reference to commercial standards. Sensitivity of the flame photometer was adjusted so that f u l l scale deflection (100% absorbance) equaled one milliequivaleat. Since calcium, potassium, magnesium, and lithium do not interfere with sodium, determinations of sodium were made directly on samples of lake water diluted with d i s t i l l e d water. Concentrations were then calculated by multiplying by the appropriate dilution factor. Since sodium and calcium interfere with the emission of potassium, a modification of standard technique was employed. One ml of swamp solution containing 18 g of CaC^^H^O and 9 g of NaCl was added to lake water diluted in 25 ml volumetric flasks so that the concentrations of Ca and Na were at 50 mg each per 25 ml volume. Standard were treated in a similar manner so that corrections for volume displace-ment wov.ld not be required. These concentrations of sodium and potassium were determined experimentally to insure that flame emission was saturated with respect to sodium and potassium. The dilutions were made with d i s t i l l e d water. Cslcium concentrations were determined by spectrofluorometric analysis according to the technique described by in the Manual of Fluorometric C l i n i c a l Procedures by G. K. Turner Assoc. Magnesium and calcium were determined jointly by the method described by Richards (1954) and magnesium was determined by subtraction of the quantity of calcium determined by spectrofluorometry. Replicate determinations were made. Carbonate and bicarbonate concentrations were determined by 12 potentioraetric t i t r a t i o n of 25 ml undiluted lake water according tc Standard Methods (1965). Commercially prepared solutions of N-HC1 diluted to 0.02 N with glass d i s t i l l e d water were used as the titrant and end points were detected with an expanding scale Radiometer pH meter. The effects of d i s t i l l e d water on the pH of the titrant were negligible. Tests also were performed which demonstrated that the waters from the most dilute lakes could be held for one or two days without significant change :.n carbonate and bicarbonate and the most concentrated lakes could be held for a week or two without significant change. Chloride was determined by amperometric ti t r a t i o n using a commercial chloridometer. The t i t r a t i o n is affected by high concentrations of sulfide., but this applies only to the sample from 5 m in LB 1 was so affected. Sulfate was determined by the ion-exchange method described by Mackereth (1955). The crux of the technique is to be sure that the majority of the anions have been determined so that when their concen-trations; are subtracted from the concentration of total cations so that one can be sure that the remaining anions required to balance the concentrations of cations and anions can be safely~attributed to sulfate. Semi-qus.ntitative determinations of the trace elements indicate that carbonate, bicarbonate, chloride and sulfate are the most important anions. Trace elements present in the surface waters of the lakes were determined by Coast Eldridge, Vancouver, B.C., using semi-quantitative spectographic analysis. However, the possibility of horizontal variation in chemical properties cannot be ruled out. The samples used for these 13 determinations were prepared by evaporating water samples over steam and the drying the resulting salts at 105° C. The salts were then stored v.n glass v i a l s . Accuracy of the ionic analyses was evaluated by comparing the total m:" lliequivalents of anions to total milliequivalents of cations. If the analyses are accurate and essentially complete, the concentra-tions o'i anions should approximately equal the concentrations of cations. The equivalence of anions in the determinations reported here can l e determined by reference to Appendix II. In general, the analyses differed, by at most 5°L and usually less. Criteria for acceptance cf chemical analyses of waters given in Standard Methods (1965) indicate that this percentage of error is unacceptable. One possible explanation of these differences is incomplete determinations of the elements cr ions present; however, trace element data do not support this possibility. More li k e l y , the errors are due to faulty experimental technique (e.£. , errors in accuracy of determination compounded by multiplication by dilution factors). suits The lakes and distribution. The 32 lakes studied are distributed as follows: (a) Chilcotin Area: Near Phalerope, Phalerope, Box 20-21, Box 4, Opposite Box 4, Near Opposite Box 4, Near Opposite Crescent, Box 98, Box 17, Racetrack, Box 27, Rock; (b) Springhouse Area: Sorenson L. , Westwick L., Boitano L., Boitano NE, Rush, Sp6; (c) Gang  Ranch Area: GR2, GR 3, White L., LE 1, LE 2, LE 3, LE 4, LE 5, Long L.; (d) Clinton Area: Clinton, Six Mile; (e) Kamloops Area: Bower's L., 14 LB 1, LL 2, Lac du Bois. General locations of the gtudy areas and locations of each of the lakes studied are shown in Figure 2. The distribution of each of the lakes with respect to underlying geologic formations and with respect to open grassland and forest are given in Figure 3. With the exception of Six Mile lake, a l l the lakes are located in open grassland or at the interface between forest and open grassland. Morphometry. During 1967 and 1968 the summers were relatively wet and the water levels of the lakes unusually high. Data concerning morphometries of the lakes are given in Table I. For future discussion, the following general points are relevant. 1. Box 20-21 consists of two lake basins which are presently joined by high water. Data were collected from both halves of the lake. 2. Sorenson Lake and Westwick were formerly one lake (Westwick Lake). Sore.nson Lake was formed when the Al k a l i Lake Road was built through Westwick Lake in the 1940's. During high water, water w i l l flow from Westwick Lake into Sorenson Lake via a conduit. 3. Boitano NE is a small lake formed by a dam which lies across a drainage ditch inlet into the north end of Boitano Lake. 4. From east to west and in order of increasing chemical concentration, 2 Localities of lakes denoted by the work "Lake" or "Lac" in their names are listed in the British Columbia.Gazetter.. Other names used have been assigned by other researchers and myself and would be of no use in locating the water bodies. 15 Figure 2. Lake l o c a l i t i e s in south-central British Columbia. A. Chilcotin Area. B. Springhouse Area. C. Gang Ranch Area. D. Clinton Area. E. Kamloops Area. 16 Figure 3. Left. Distribution of lakes with regard to forest (shaded) and open grassland (clear). Right. Distribution of lakes with regard to underlying geology. The figure was adapted from Cockfiel 1 (1948), Campbell and Tipper (1!>66) and Tipper (1959, 1963). Geological formations are identified as follows: PALAEOZOIC 1 Permian, Cache Creek Group. Chert, a r g i l l i t e , limestone, greenstone, breccia, and conglomerate. Lower Permian, Cache Creek Group. Basic volcanic flows, tuff, ribbon chert., limestone and a r g i l l i t e . Upper Permian, Cache Creek Group. Marble canyon formation, massive limesrone, limestone breccia and chert, minor a r g i l l i t e , tuff, andesltic and basaltic flows MESOZOIC I I I I I 1^  ' l I I 1 t l ~ , 1 1 1 1 . Il ' l ' l ' l 1 Triassic (middle or upper). Limestone, basalt, related tuffs and breccias, a r g i l l i t e , greywacke, and conglomerate. Upper Triassic, Nicola Group. Greenstone, andesite, basalt, tuff, agglomerate, breccia, minor a r g i l l i t e , limestone, and conglomerate. Jurassic (?). Chert and pebbles conglomerate, greywacke. Jurassic (?). Shale and grit. Jurassic and later. Coast intrusions, granite, grandodiorite, and gabro. ' CENOZOIC Kamloops Group, Coldwater beds. Conglomerate, sandstone, shale, and coal. Kamloops Group, Meiocene or earlier. Rhyolite, andesite, basalt, associated tuffs and breccias, and agglomerates. Kamloops Group. Tranquille beds. Conglomerate, sandstone, shale, tuffs, and thin coal seams. •> 8 n ° n . I " I - I N ' h- + -H + + Miocine. Shale, sandstone, tuff, diatomite, conglomerate, and breccia. Tertiary. Miocene, and'/or Pliocene. Plateau lava, olivine basalt, basalt:, basalt andesite, and related ash and breccia. Pleistocene and recent. T i l l , gravel, clay, s i l t , and alluvium. 17 Table I. Summaries of elevations of lakes and of morphometric data (A= area i n hectares, V= volume i n thousands of cubic meters, z m = maximum, depth i n meters, z = average depth i n meters, and z : z m = volume development r a t i o ) . MORPHOMETRY DATA ELEVATION LAKE (m) A V z z : z. CLINTON GR-2 1,095 15.37 127.3 1.5 0.8 0. 53 LB-2 899 3. 06 65.6 2.5 1. 1 0.44 LB-1 • 884 5. 11 150.4 5. 2 2.9 0. 56 LONG LAKE 1,037 33. 53 735.5 4.5 2. 2 0.49 BOX 4 945 17. 19 348.4 4.5 2.0 0.44 BOWER'S LAKE 701 3.55 46. 1 2. 5 1.3 0. 52 LE-1 1,037 2.19 22. 1 2.3 1.0 0.43 LE-2 1,037 4.80 44. 7 2.3 0.9 0.39 PHALEROPE 945 30. 84 787. 6 6.2 2.6 0.40 BOX 20-21 945 46. 52 1,283.2 5.4 2.8 0. 52 WHITE LAKE 1,037 127.68 6,416.4 15.5 5.0 0.32 LE-5 1,037 2. 19 4.9 0.7 0.2 0.29 BOITANO LAKE 975 80.68 2,202.2 4.5 2.7 0.60 RUSH 975 19. 60 212. 7 2.5 1.1 0.41 LE-3 1,037 3. 13 22. 0 1.5 0.7 0.47 LE-4 1,037 6. 99 88.8 1. 5 1.3 0.84 SIX MILE 15.0 NR OP BOX 4 945 5.81 79.9 2.3 1.4 0. 61 BOX 89. 945 15.17 156.8 2.3 1. 0 0.44 ROCK 945 34.64 387.5 2.5 1.1 0.44 GR-3 NR PHALERO??E 945 5.06 64.6 3.0 1.3 0.43 WESTWICK LAKE 945 58.30 728.3 4.5 1.3 0. 29 SORENSON L\KE 945 2.5 LAC DU BOIS 869 29.63 1,380.2 8.3 3.9 0.47 NR OP CRESCENT 945 6.88 99.2 3.3 1.4 0.42 BOX 17 945 2.65 31.4 3.3 1.1 0.33 BOITANO NE 975 2.0 OP BOX 4 945 4.55 32. 8 2. 2 0. 7 0.32 RACETRACK 945 27.05 503. 0 6. 5 1.9 0. 29 SP 6 975 0.86 5. 1 1.5 0.6 0.40 BOX 27 945 4.30 23. 0 1.5 0.5 0.33 18 White L., LE 5, LE 4, LE 3, LE 2, LE 1, and Long L. a l l occur in the same drainage and are interconnected. Long L. is a closed Lake with no outlet and water from the other lakes empties into this lake. Since the summer of 1963 steps have been taken to divert Big Ba:: Creek drainage into the White Lake and through Long L. basin in order to freshen up the water contained in the basin so i t may be used for irrigation. If the plan is completed, the lifetime of this serLes as described is limited. The effect of diverting water into White Lake has been observed in the rise in water level of the entire series of lakes. LE 2 and LE 1 formerly were separate basins, but they are now joined with Long Lake. 5. Lac du Bois has an outlet through which water level within the lake can be regulated. Generally, a l l the lakes studied are shallow glacial type lakes and most occupy closed basins. Moreover, most of the lakes might be most usefully termed ponds; however, the term has no quantitative mean-3 ing. The lakes are "saucer shaped" and average 21 ha in area, 573,000 m in volume, and 1.6 m deep. Elevations', at which most of the lakes occur and morphometric measurements for each of the lakes on which soundings were made are given in Table I. Volume development ratios are given in the last column of the table and i t should be borne in mind that a ratio of 0.33 indicates a conical lake with the base of the cone corresponding in area to the area of the lake. A ratio of greater than 0.33, indicates a shallow lake with a f l a t bottom and ratios of less than 0.33 indicate deeper lakes or lakes deep relative to their surface areas. In the series of lakes studied, the volume ratios may be used to deduce shape, since only Long L. and Box 20-21 have more than a single deep basin in 19 the lake. The percentage of total volume of each lake which is present at meter intervals in depth in each of those lakes is given in Appendix III. Physico-chemical conditions. The general chemical properties of the lakes are summarized in Tables II-V. Vertical and seasonal variation in temperature, pH, and dissolved oxygen are given in Table II; seasonal variation in the average values of conductance, total dissolved solids (TDS), and the concentrations of the major ions are given in Table III. Trace elerrents present in the surface waters of the different lakes ace given in Table IV. An analysis of variance of the data given in Table II and III was made to determine yhe significance of variation in the chemical factors with respect to lakes, depths, and seasons and the interactions of each. The results of this analysis are given in Tabl= V. In general, seasonal variation in the physico-chemical properties is less pronounced than are the differences among and within the lakes. A surprising feature revealed by the analysis is the lack of any s t a t i s t i c a l l y significant differences in the concentration of dissolved oxygen. Conductivities of each of the lake waters have been measured as a function of concentration for periods ranging from two to ten years and the ranges of concentrations observed in the lakes are shown in Figure 9. The lakes form a continuous series of environments with conductivities ranging from 27-50,000 micromhos. Most of the conduct-i v i t y data used in the preparation of this figure were obtained by the measurement of conductivities of surface waters. Since "fresh-water" (e.g_. , runn-off or melt water) of low conductivity may layer on the top of more concentrated water, conductivity of these surface waters may exaggerate the range of variation shown by the remainder of the volume of any one of the lakes. 20 Table I I . Temperature, pH, and dissolved oxjgen i n surface and bottom waters of the lakes. DISSOLVED OXYGEN TEMP. (°C) pH (cc/1) zm — : LAKE (m) DATE iSURF. BOTT. SURF. BOTT. SURF. BOTT, CLINTON 8/VIII/66 8. 10 GR-2 1. 5 20/V/66 15. 6 25. 9 10.05 10.00 4/VIII/66 21. 1 20. 6 10. 20 10. 20 16/11/67 -2. 0 9. 70 LB-1 5. 2 23/V/66 14. 7 16. 4 8. 80 7. 15 5. 70 0. 00 6/VIII/66 27. 6 25. 3 8. . 70 7.80 5. 32 5. 94 LB-2 2. 5 23/V/66 15. 6 11. 7 9. 50 9.45 5.05 4.35 6/VIII/66 22. 8 21. 1 9. 20 9. 25 3. 84 2. 13 LONG LAKE 4. 5 20/V/66 14.4 7. 8 9. 00 9.40 5. 87 3/VIII/66 22. 2 15. 3 9. 20 9.45 5. 60 17/11/67 0.0 1. 3 8. 80 9. 25 BOX 4 4. 5 13/V/66 11.4 4. 4 9. 40 9.30 28/VII/66 20. 0 5. 6 9. 20 9.40 BOWER'S LAKE 2. 5 24/V/66 14.4 13. 9 8. 20 8. 15 7. 93 8. 25 7/VIII/66 22. 0 22. 0 8. 60 8.60 LE-1 2. 3 20/V/66 13.3 13. 6 9. 25 9. 00 5.54 5. 54 3/VIII/66 22. 5 22. 8 9. 30 9.40 4.87 4.65 LE-2 2. 3 20/V/66 12. 5 12. 8 8. 95 8. 95 5. 71 3/VIII/66 22. 2 22. 0 9. 30 9.30 5. 15 4. 87 PHALEROPE 6. 2 12/V/66 12. 2 3. 3 9. 20 9. 10 27/VII/66 18. 9 8. 9 9. 15 9. 15 5. 26 0. 00 BOX 20-21 5. 4 13/V/66 13. 6 4. 3 9. 10 9. 20 5. 50 0. 00 27/VII/66 20. 0 6. 4 9. 10 9. 20 4. 70 0. 00 WHITE LAKE 15. 5 21/V/66 11. 1 5. 6 9. 10 9. 20 5. 65 0. 00 4/VIII/66 20.3 5. 9 9. 25 9.40 8. 96 0. 00 16/11/67 0. 0 4. 0 9. 10 9. 15 3.42 0. 00 LE-5 0. 7 22/V/66 12. 0 • - 8. 40 6. 59 5/VIII/66 20. 0 • - 8. 70 3.30 BOITANO LAKE 4. 5. 10/V/66 14.4 8. 9 9. 00 8.80 4. 95 0. 00 31/VII/66 18. 9 17. 0 8. 85 8. 70 4.44 0. 00 RUSH 2. 5 l l/V/66 15.3 15.3 8. 70 8. 50 4. 50 0. 00 31/VII/66 20. 0 17. 8 8. 60 8. 60 3. 53 3. 92 LE-3 1. 5 22/V/66 11. 7 — • - 8. 50 5.49 • 5/VIII/66 19. 7 — 9. 20 7. 17 LE-4 1. 5 22/V/66 13. 1 12. 8 8. 70 8. 70 6.47 6.36 5/VIII/66 20. 3 20.3 9. 20 9. 20 8. 90 8.46 SIX MILE L5.0 17/V/68 • - 8. 00 7. 70 NR OP BOX 4 2. 3 14/V/66 12.8 7. 8 8. 90 8. 60 5. 85 1. 92 28/VII/66 20. 9 15. 9 9. 00 8. 65 8. 06 0.00 BOX 89 2. 3 17/V/66 13. 3 11. 1 8. 39 8. 58 6. 16 6. 05 29/VII/66 19. 2 19. 2 8. 70 8.80 4.42 0. 90 ROCK 2. 5 17/V/66 12.6 11. 7 8. 80 8. 90 6.39 6.32 30/VII/66 20.3 18. 3 9. 00 9. 10 4.14 2.86 GR-3 • - 4/V/67 — • - 8. 90 SORENSON LAKE 2. 5 4/V/67 — • - 8. 80 8.80 NR PHALEROPE 3. 0 12/V/66 14.4 14. 4 8. 60 8. 60 5. 57 4.44 27/VII/66 18. 9 16. 4 8. 10 8.10 1.04 1. 18 Table II ( c o n t . ) . DISSOLVED OXYGEN TEMP. (°C) pH ... (cc/1) LAKE (m) DATE SURF. BOTT. SURF. BOTT. SURF. BOTT. WESTWICK LAKE 4. 5 10/V/66 15. 6 15.6 8.90 8.90 . , 3.60 3.63 '26/VII/66 23.3 19.4 8. 80 8.80 1. 96 0. 22 18/11/67 0.9 3.6 8. 10 7. 90 1.02 0. 20 LAC DU BOIS 8. 3 23/V/66 14.4 7.8 8.50 8. 14 6.95 0.43 6/VIII/66 22. 2 10.0 8.60 8. 20 5.35 0.00 NR OP CRESCENT 3. 3 16/V/66 11. 7 11.1 8. 60 8. 70 5.48 5.64 29/VII/66 18.9 15.9 8.60 8. 70 0. 92 0. 00 BOX 17 3. 3 16/V/66 12.8 11. 7 8.30 8.30 6.38 5.53 29/VII/66 19. 2 15.3 8.80 8.60 5.43 0. 00 BOITANO NE 2. 0 10/V/66 16. 1 7. 70 26/VII/66 17.8 9.05 OP BOX 4 2. 2 14/V/66 11. 1 10. 6 8.60 8.50 4.15 2. 02 28/VII/66 20. 3 16.7 8.80 9. 00 5.71 3. 14 RACETRACK 6. 5 16/V/66 12. 2 11. 1 8.00 7. 90 5. 21 3.94 29/VII/66 18. 6 14.4 8.60 8. 00 5.40 0. 00 SP-6 1. 5 15/V/66 13.3 12. 8 7.70 7. 95 6.75 5.42 31/VII/66 24. 2 20.0 9. 00 8. 70 7. 22 5.71 BOX 27 1. 5 17/V/66 13.3 12. 8 6.40 6.40 6.39 5.59 30/VII/66 19.4 17. 2 9. 15 6. 20 4. 20 4.20 21 Table III. Summary of average physico-chemical conditions present during different seasons. H I L L I EQUIVALENTS PER LITER C o n d u c t i v i t y TQS Lake Date (umbos a t 25 C) ( n c i / l ) Na K <a Mg C 0 3 HCO^ C l SO C l i n t o n 5 / V I I I / 6 6 55,932 1 4 7 . 1 0 0 327. 50 18. 80 I 16 1 , 630. 45 52. 70 31.25 2,042. 70 GR 2 : :o/v/G6 32 ,010 29,v"S7 5 3 5 . OO 1 1 . "8 (•. 61 2. 43 416. 65 55. .98 126.14 4 / V I 1 1 / 6 6 40, 202 •;o, 3 7 0 712. 96 13. 86 0. 14 3 . 90 565. 77 70. 53 165.57 L6 / I1 /67 57,320 69,2-: a 1 , 312 . 00 25. 00 (.'. 29 9. 41 978. 00 162. .00 296.00 LB 1 ! 3 / V / 6 6 16,729 27,727 85. 01 9. 54 32 251 . 10 3. .35 11. .04 9. 24 325.55 6 /VI1I /6C- 17,033 28 ,000 90. 55 10. 59 .80 239. 72 2. .86 13. .13 9. 54 336. 25 LB. 2 >3/V/66 15,300 14,04 2 216. .30 9. . 4 4 Ii. .72 ... 93 64. .30 62. .30 16. 20 69. 60 6 / V I I I /66 17,067 • 15,288 225. 10 9. .10 II. .68 S. 56 65. .4 60. 90 15. 60 99. 10 1x3 ng L 2 0 / V / 6 6 12,694 1 ! , 661 1 58. .09 9. .80 . 56 12. 57 31. .84 34 . 33 21.75 96. 11 3 / V I 1 1 ,'G6 13,199 12,054 151. .24 9. . 04 0 . . 11 13. .95 29. .78 3 3 . .87 20. 78 96. 19 17/11/67 12,612 12,231 158. . 04 9 . 07 . 50 17. .38 29. .23 47, . 59 23. 35 29. 39 Box 4 1 3 / V / 6 6 11,3 56 9, 573 130. . 26 13. . 4 5 .73 2. .66 35. .99 53 . . 52 36.97 23. 71 2 8 / V I I / 6 6 12,275 9, 999 132 .07 13 . 34 •). . 1 2 3 . 25 63. . 14 54 . .32 37.63 27. 96 B o w e r ' s L 2 4 / V / 6 6 10,170 13,914 53 . 50 ' 5. . 4 5 2'.). .30 98, .90 3 . . 56 6.99 1 58. 58 7 / V I I I / 6 6 11 , 864 1 7 , 3 3 4 63 .85 5, .87 2 1. 97 122. . 59 0. .96 1. . 26 7.21 197. 83 LE 1 2 0 / V / 6 6 9, 157 8, 089 106. 84 - 3 / V I I 1 / 6 6 11,227 9, 658 119. 50 L E 2 2 0 / V / 6 6 7,923 6. 806 85. 77 3 / V 1 I I / 6 6 11,058 9, 558 116. 71 P h a l e r o p e 1 2 / V / 6 6 6, 459 5, 024 ' 71 . 35 ' 2 7 / V I I / 6 6 7,311 5, 696 7 5'. 93 Box 20-21 1 3 / V / 6 6 6, 412 4, 804 70. 25 2 7 / V I I / 6 6 6, 683 5, 078 69. 00 W h i t e L 2 1 / V / 6 6 . 5, 234 4, 334 58. 24 4 / V 1 I I / 6 6 5,371 4, 216 59. 59 16 /11/66 5, 384 4, 411 55. 14 LE 5 2 2 /V/66 3, 870 3, 058 40. 00 5 / V I I I / 6 6 4, 136 3, 185 38. 30 B o i t a n o L 1 0 / V / 6 6 4, 066 3 ,118 34. :o8 31/VI . I /66 4, 149 3, 160 33. 67 Rush l l / V / 6 6 3, 656 3, 196 31. 92 3 1 / V I I / 6 6 4, 254 3,251 34. 80 L E 3 : 2 / V / 6 6 3,435 2, 709 30. .00 5 /V111/66 3 ,915 3,237 35. 30 L E 4 2 2 /V/66 3, 167 2.481 27. . 50 5 / V I I I / 6 6 3,864 3,206 35. 30 S i x M i l e 1 7 / V / 6 8 2,899 2, 612 5. .41 Nr Op Box 4 1 4 / V / 66 2,372 2, 086 15. .42 >8 /VI1 /66 3 ,159 ' 2,913 16. .96 Box 89 17/V/66 1, 508 1, 004 13. .25 53 /V1I /66 1, 695 1,358 14. .28 Rock 1 7 / V / 6 6 1,404 1, 051 14. . 13 3 0 / V I 1 / 6 6 1, 588 1,209 15.80 GR 3 1 0 / I X / 6 7 1, 500 1,241 12. .99 S o r e n s o n L 4 / V / 6 7 1, 500 1,031 4. .43 Nr Phalerope ".2 / V / 6 6 1. 182 862 7. .25 .'7 / V I I / 6 6 1 ,485 861 . 8. .85 Westwick L 10 / V / 6 6 1, 236 1, 003 4. . Gl . 2 6 / V I I / 6 6 1,338 93 5 4. .70 18 / 1 1 / 6 7 1, 770 1,375 6. .36 Lac du Bois 23/V/66 921 7 58 2. .85 6 / V I I I / 6 6 981 794 2. .42 Nr Op Cr 1 6 / V / 6 6 83 7 558 3 . .41 2 9 / V I 1 / 6 6 782 G38 3. .33 Box 17 1 6 / V / 6 6 740 471 3. .31 2 9 / V I I / 6 6 741 670 3 , .32 B o i t a n o NE 1 0 / V / 6 6 59 5 338 3 . .20 2 6 / V I I / 6 6 712 427 3 . ,80 Op Box 4 1 4 / V / 6 6 558 400 1. , 05 2 8 / V I 1 / 6 6 625 499 2. , 07 R a c e t r a c k 1 6 / V / 6 6 481 330 3. .08 2 9 / V I I / 6 6 49 5 4 14 3, , 24 Sp 6 1 5 / V / 6 G 245 111 0. .27 3 1 / V H / 6 6 2 5G 114 . 0. .41 Box 27 1 7 / V / 6 6 39 32. 0. .15 3 0 / V I 1 / 6 6 40 1 0. .02 7 . 19 ). 70 11. 39 21 . 00 25. 65 16. 91 75.36 7. 3 2 ), 18 13. S3 24. 16 23. 92 18. 74 72.02 5. 99 ). 69 12. 11 16. 42 25. 60 12. 60 39.49 7. OS 0. 14 13. 87 24. 07 29. 11 19. 67 73. 50 5. 21 ). 49 2. 96 13. 86 31. 87 23. 69 9. 08 5 - 29 0. 70 3 . 13 15. 86 32. 55 27. 10 13.60 5. 36 3. 74 2. 81 15. 7 5 33. 05 21. 90 7.93 4 . 85 ). 73 3. 06 15. 78' 34. 89. 22. 24 6. 05 5. 55 0 . 2] 9. 45 27. 99 35. 52 8. 45 4.06 4 . 32 0 . 16 9. 77 29. 78 32. 46 8. 64 2.26 4 . 54 3. 19 10. 07 29. 73 37. 84 9. 03 0. 11 3. 08 0 . 72 12. 28 2. 44 40. 30 7. 23 6. 08 3 . 18 3. 30 11. 79 5. 76 30. 12 7. 53 2.45 2. 98 0 . 73 11. .15 5. 02 17. 62 4. 20 22.59 3. 43 0 . 73 1 1 . 87 5. 10 17. 95 4. 73 23.07 2. 34 0 . 77 10. 92 2. 96 17. 52 3. 85 18. 04 3. 05 9. 71 12. 87 3 . 81 20. 09 4. .68 24. 03 2. 62 0 . 90 13. 87 4 . 40 27. 25 5. 16 5.91 2. 75 0 . 24 13 . 49 15. .84 24. 48 6. 86 6.48 2. 39 0. 74 . 1 1 . 94 3. 52 31. 00 5. 19 2.45 2. 65 0 . 36 12. 54 • 14. .30 25. 74 6. 65 2.98 0. 33 8. 56 •26. 34 3. 85 0. 53 31.02 1. . 52 1. 79 13. 51 1. .73 5. 52 3. 28 22. 34 1 . 53 1. 04 18. .79 2. .42 6. 48 3. 93 25. 44 1. 72 0 . 81 2. . 55 0. .66 13. 11 1. 25 2.73 1. .39 0 . 30 3. .30 2. .48 12. 55 1. 00 3.67 1. .00 0 . 85 1. ,47 3. . 18 12. 92 1. 85 0.03 0. .96 0 . 38 2. , 00 5. .46 11. .78 1. 28 0.32 0. .95 1. 52 6. . 28 1 . .07 12. .62 0. 61 1.95 0. . 69 1. 15 11. .95 1. .33 6. .38 0. 34 9.29 1. .30 0. 93 5. .20 1. .40 12. .02 0. .49 1.40 1. , GO 0 . 93 6. .30 -• - - - 16. . 73 1. .08 0.05 0. .81. 1. 30 11. .12 1. .39 7. .08 0. .11 7.26 0 . .85 0. 74 11 , .30 1. . 78 8. . 16 0. .35 7.15 0. ,90 1. 57 14. .99 — — 9. .40 0, .14 8. 19 0, , 70 1. 74 8. .35 1, .38 10. .30 0. . 19 0.85 0. . 54 0. 99 9. .38 1 . 72 1.0. .37 0. .44 1.11 0. . 65 1. 3 7 3. .90 0 .70 6, .37 0. .25 1 .07 0. . 54 0. 57 4. .90 0 .91 6. .47 0. .45 1.61 0. .69 1. 08 4 .35 8. .89 0 . 14 0. 10 0. . 52 0. 43 4. .96 2 .28 6 .49 0 . 13 0.23. 1. .31 0. 68 2 .33 _. 5. .01 0. . 10 2.80 0. . 57 0. 86 3 .31 1 . 76 5 .84 - • 0. 28 0. .60 0. 38 5 .13 0 .51 5 . 78 0. .15 0. .64 0. 38 5 .13 0. .37 5. .67 0. .33 0.40 0. .49 0. 76 1 .41 _. 5 .24 0 .26 0. 47 0. .40 0. 33 1. .90 0 .88 4 .67 0. . 15 0. 16 0. .10 0. 87 1 .46 - 2. .65 0 .09 0. . 76 0. 79 2 . 18 1 .01 2 .21 - • 0.04 0. . 12 0. 17 „ . . . _. 0 .39 0 .07 0. 13 0. .07 0. .06 0. . 16 0 .10 0. .18 0 .03 0.04 22 Table IV. Comparison of trace elements present i n samples of surface waters collected i n May 1966. T R A C E ELEMENTS (ppm) Lake Al Sb As Ba Be Bi B Cd Cr Co Cu Ga Au Fe Pb Mn Mo Nb Ni . S i Ag Sr T a Sn T i W V Zn C l i n t o n -- -- -- -- -- -- -- -- * -- — __ __ __ ' __ _. •__ » GR 2 - - — — — - - - - — — — — — — — * __ __ LB 1 -- -- — -- — -- -- -- * -- •-- — — * * -_ __ ._ . „ __ „ LB 2 - - - - - - - - — ' - - - - - - - - - - - - - - - - - - - - _.. __ __ __ ' __ __ . _ __ __ _ . . Long L 0. 80 — -- 0 . 24 — 0 . 40 -- 0. . 8 0 -- 0. . 56 -- --. 4 . 0 0 . 24 0 . 40 0 . 48 — * 24 . 0 0 . 24 0 . 56 — — 0.24 - - • -- — Box 4 7. 54 — * — 0. . 66 -- 18 . 9 0 . 47 -- 0 . 28 -- 7 . 5 0 . 09 0 . 9 4 — -- 0 . 0 9 - -Bower's L 6. 98 5. 58 • — * — 0. . 56 — — 11 _ 2 -- — 0 . 4 2 -- -- 14 . 0 0 . 4 2 9 . 77 — -- 3 . 9 6 — * T.E 1 * — 0. . 33 -- — • 16. .4 0 . 4 1 -- 0 . 66 -- -- 4 . 1 . 0 . 66 0 . 8 2 — — 2 . 4 6 - - 0 . 8 LE 2 6 . 14 — — 0 . 08 — — - — * -- 0. . 77 -- -- 6. . 1 -- 0'. 23 -- -- . 3 . 8 0 . 23 0 . 08 — -- 0 . 2 3 - - — — Phalerope 1. 57 -- -- * -- - -- -- 0. .32 .— -- 0. .9 * -- 0 . 03 -- -- 2 _ 2 0 . 9 4 0 . 94 — — 0 . 0 9 - -Box 20. 1 m 65 -- -- 0 . 33 * 0 . 33 -- 0. . 99 -- 0. . 66 — — 3 . 3 0 . 3 3 0 . 2 6 0 . 2 6 -- 0 . 03 3 . 3 0 . 3 3 0 . 99 0 . 2 6 - -Box 21 1. 61 -- -- 0 . 32 -- -- * -- -- 0. 32 -- -- 1. . 6 -- 0 . 03 -- ' — 1 . 6 0 . 2 6 0 . 9 7 — — 0 . 1 0 - -White L 3 . 98 -- -- 0 . 04 -- -- 0 . 12 — 0 . . 04 -- 0 . 20 -- -- 3 . 2 0 . 2 0 0 . 1 2 3 . 19 -- * 3 .2 0 . 2 0 1. 99 -- — 0 . 4 0 - - * — LE 5 2 . 14 * -- 0. 06 — -- 2. .4 • -- — 0 . 09 -- 0. .3 0 . 0 9 0 . 2 5 -- -- 0 . 2 5 — * — Boitano L 1. 53 -- -- 0 . 61 * -- * -- 0 . 92 -- 0 . 2 4 -- -- 0. .3 * -- 0 . 3 1 -- 0 . 2 4 0. .9 0 .1 5 0 . 9 2 -- -- 0 . 3 1 - - 0.9 — Rush T. ^ 12 -- • — 0 . 37 * -- •- -- 0. 19 -- 0 . 30 -- -- 1. . 1 * -- 0 . 3 7 -- * ' 3 . . 0 1. 12 1 .86 -- -- 0 . 3 7 - - 1.5 — LE 3 1. 63 -- -- 0 . 14 -- -- •- 0 . 03 -- 0 . 11 -- -- r, , 4 0 . 03 0 . 2 7 -- — 0. . 8 0 . 14 0 . 8 1 -- -- 0 . 2 7 — LE 4 1. 98 -- -- 0 . 03 * -- 0 . 20 — 0 . . 03 -- 0 . 07 — -- 0. , 7 . 0 . 2 5 -- 0 . 07 — — 1. . 2 0 . 1 2 0.49 -- * 0 . 1 2 — * — Six Mile 0 . 01 * -- * -- 0 . 05 — -- 0. . 2 — — * -- 0 . 01 0. ;6 -- 0 . 2 0 -- --. 0 . 0 1 — Nr Op Box 4 0 . 58 -- * -- -- * 0 . 06 — — 0. 1. — 0 . 02 0 . 02 — — 1. . 0 0 . 58 0 . 97 -- -- 1 .94 — 1.4 . — Box 89 2 . 1 2 — — 0. 21 -- 0 . 09 -- 0 . ,09 -- 0 . 11 -- -- 5. 3 0 . 11 0 . 0 5 0 . 3 2 — 0 . 01 1. 1 0 . 09 0 . 3 2 -- -- 0 . 1 1 - - * — Rock 0 . 84 — -- 0. 84 -- -- 0 . 00 -- 0 . . 1 1. — 0 . 05 -- — 1. 1 * * 0 . 08 — * 1. 1 0 . 1 1 0 . 53 — -- 0 . 0 7 0. 1 — GR 3 0 . , 01 0 . 04- -- * * 0 . 04 — -- 0 . 1 -- -- * -- 2. 0 -- 0 . 2 0 -- -- 0 . 03 — Sorenson L 0. . 01 * -- -- 0 . 01 -- 0 . 2 — -- * -- 0 . 01 0. 4 -- 0 . 4 0 — .-- 0.04 — Nr Phalerope 0 . .43 — * — -- 0 . 04 -- * 0 . 04 — 0 . 9 0 . 09 0 . 1 7 0 . 68 -- * 0 . 6 0 . 09 0 . 26 -- -- 0 . 0 1 — — — Westwick L 0. . 3 5 — — 0. . 12 — — 0 . 06 -- 0. . 04 -- 0 . 12 — — 0 . 6 0 . 01 -- 0 . 3 5 '-- * 0. 9 0 . 0 1 0 . 3 5 — -- 0 . 3 5 — 0.8 — Lac du Bois .0. . 3 8 -- -- * * — 0 . 04 --• * -- 0 . 08 -- -- 0 . 4 * -- 0 . 0 1 -- -- 0 . 5 0 . 06 0 . 3 8 -- -- 0 . 2 3 — 0. 6 Nr Op Cr 0, . 11 * -- 0 . 02 -- 0 . 3 -- — 0 . 04 -- -- 0 . 3 0 . 0 3 0 . 04 -- — 0 . 0 1 — Box 17 0. . 14 * -- 0 . 02 — — 0 . 2 * -- 0 . 0 4 -- -- . o. 1 0 . 01 0 . 05 — — 0.01 — Boitano NE 0. . 24 -- __ 0 . 03 * — * -- 0. . 01 -- 0 . 10 -- -- 0 . 3 0 . 24 0 . 0 1 -- -- 0. 7 * 0 . 17 -- -- 0.03 - - * --Op Box 4 0 . 0 4 * -- 0 . 01 — — 0 . 2 -- -- 0 . 03 --' -- 0 . 2 0. 01 0 . 01 — Racetrack 0 . 68 -- * — — 0. . 03 — . 0 . . 10 — 0 . 03 -- -- 6. 8 * 0 . 02 0 . 24 — * 0 . 3 • * 0 . 10 — — 0. 24 — 0.1 Sp 6 0 .11 0 . 0 3 * -- 0. . 01 — 0. . 0 1 — 0 . 09 -- -- 0 . 1 . 0 . 01 . 0 . 0 1 . 0 . 0 1 — 0. 01 0. 1 * 0.08 -- — 0.01 — * — 23 Table V. Summary of the s t a t i s t i c a l significance of variation in certain physico-chemical properties between the lakes, durir.g two seasons (spring and f a l l ) , at different depths ( O n , 1 m, and bottom), between the lakes during the.two seascns (L-S) , between the lakes a,; the different depths (L-D), and at different depths within the lakes during the two seasons (S-D). S t a t i s t i c a l l y significant variation implies only that the data analyzed for that source of variation cannot be considered to be homogeneous. The exact lakes, depths, or seasons which cause the observed hetero-geneity have not been identified. PHYSICO-CHEMICAL PROPERTIES SOURCE OF VARIATION CONDUC. TDS• pH TEMP 0 2 Na LAKE < 0.001 <0.'001 <0.001 <0.001 >0.05 <0.001 SEASON <0.001 <0.001 <0.001 <0.001 >0.05 <0.001 DEPTH < 0.001 <0.001 <0. 05 < 0. 001. > 0.05 < 0.001 L-S <0.Q01 <0.01 <0.001 <0.001 >0.05 <0.001 L-D < 0.001 <0.001 <0.05 <0.001 >0.05 <0.001 S-D < 0.001 < 0.05 >0.05 < 0.001 >0.05 <0.01 • K Ca Mg C03 HC03 Cl S04-<0.001 <0.001 <0.001 <0.001 <0.001 <0. 001 < 0.001 >0.05 <0.001 >0.05 <0.001 >0.05 <0.001 >0.05 <0.001 <0.01 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 >0.05 <0.001 <0.05 <0.001 > 0.05 <0.001 <0.001 <0.001 <0.001 <0.001 < 0.001 < 0.001 <0.001 >0.05 >0.05 >0.05 <0.01 <0.05 >0.05 24 In general, the lakes differ greatly in chemical concentration and osmotic pressure. In addition to possessing different total solute concentrations, the lakes also possess different concentrations of the major ions. The data given in Table III were derived by averaging values at the different depths in the lakes. The specific values of each factor, at each depth, during the different seasons are given in Appendix II. Of the eight quantitatively important ions, i t is relevant to note that the concentrations of a l l of them varied significantly between the lakes, tluring the different seasons, and in the different lakes during the different seasons. Significant seasonal variation was shown in. the concentration of sodium, calcium, carbonate and chloride. Significant differences in concentrations in the different lakes during the different seasons were shown by sodium, potassium, calcium, carbonate, bicartonate and chloride. Significant differences in concentrations at different depths during different seasons were shown only by sodium, potassium, bicarbonate and chloride. The measures of total chemical concentration (TDS and conductivity) showed significantly different values in a l l combinations of conditions. Clearly, the lakes indicate significant variation in chemical composition and concentration. In addition, i t is worth observing that Na and Mg are the major cations, while COg-HCO^  and SO^ are the major anions. Further, though not absent, chlorides were present only in relatively low concentrations. The concentrations of individual ions relative to total chemical concentrations of the lakes are shown in triangular co-ordinate plots in Figures 4 and 5 respectively. Data presented in Figure 4 indicate that three relatively distinct groups of anions occur in the lakes: 25 Figure 4. Relative composition of anions in and total ionic composition of the lakes. In this plot, the distance of a circ l e from the side opposite the angle marked by a particular anion is proportional to percentage concentration of that anion in that lake. The diameter of each cir c l e is proportional to the total ionic concentration of that lake. The circles, repre-senting the lakes, are coded according to the geographic area in which the lakes occur (i. a . , clear = Gang Ranch Area, solid = Chilcotin Area, stippled == Kamloops Area, and hatched Springhouse Area). 26 Figure 5 . Relative composition of cations ir. and total ionic composition of the lakes. In this plot, the percentage concentration of the cations and total ionic concentration of each of the lakes is indicated as described ir- Figure 4 . The same color codings are used to indicate the geographic areas in which the lakes occur. M a 27 (1) carbonate-bicarbonate, (2) sulfate, and (3) carbonate-bicarbonate and sul::ate. The carbonate-bicarbonate lakes show a slight trend toward replacement of the carbonate-bicarbonate by chloride; the sulfate groups occur in the Kamloops Area. Data presented in Figure 5 show that sodium and magnesium are the most common cations and that in general, magnesium is replaced by sodium as total lake concentration increases. Contradictions to this generalization are two concentrated magnesium lakes in the Kamloops Area. An attempt was made to classify the lakes according to major anions and cations in order to allow determination and evaluation cf their effects. Examples of each of chemical types of lakes are given in Figure 6. The dotted circles around each of the ionic diagrams are drawn so that i f a particular ion exceeds 50% of the total anionic or cationic concentration, the segment representing that ion w i l l cross the dotted c i r c l e . And, i f one of the anions or cations exceeds 50% of the total concentration, then the sum of the other three must be less than 507o and in likelihood the concentration of any one would be substantially less than 50%. In this manner the lakes were grouped according to the combination of anion and cation which exceeded 50% of the total concentra-tion. If the 50% level was not exceeded the lake was identified as a combination of the two most abundant cations or anions. This c l a s s i f i c a -tion and graphic representation has the advantage of being non-arbitrary and of allowing one to see at a glance the major anions and cations and the relative concentrations of each. The chemical categories into which each of the 32 lakes are classified are given in Table VI. As well as knowing the concentrations of various chemical constituents in the lake waters, i t is also relevant to know how these 28 Figure 6. Ionic diagrams showing examples of the s i x "chemical" types of lakes present i n the study area. In these diagrams, the segments representing the major ion are drawn so that their areas are proportional to the percentage concentrations of the ions. The dotted c i r c l e represents the l e v e l which a segment would reach i f the percentage concentration of the ion equals or exceeds 257» of the t o t a l ionic concen-t r a t i o n of the lake considered. a O 29 Table VI. Classification of the lakes according to the predominate anio-i(s) and cation(s) present. In the table,the lakes are listed according to their cherrical concentrations with Clinton being the most concentrated and Box 27 the least. COMBINATIONS OF PREDOMINATE IONS 1 pa 1 w H H < w < H S3 H w <! O Pd < o H f a P Q H P Q <C O < h O M H PQ <c n <! Q .-1 Q Pi O o o !=> O :<c (-I O r-H c o C O C O u pq C O C 3 u P Q M pd c o H w < !3 Pn O tJ 3 P S c o Pd H <: pn w H <J IS O P Q Pd H < O P Q < 3 Pi U < M O P Q CLINTON X GR-2 X LB-2 X LB-1 X LONG LAKE X BOX 4 X BOWER'S LAKE X LE-1 X LE-2 X PHALEROPE X BOX 20-21 X WHITE LAKE X LE-5 X BOITANO LAKE X RUSH X X LE-3 X LE-4 . • X SIX MILE X NR OP BOX 4 X BOX 89 X ROCK X GR-3 X NR PUALERO.'E X WESTWICK LUCE . X SORENSON L\KE X LAC DU BOI-5 X NR OP CRESCENT X BOX 17 • X BOITANO NE X X OP BOX 4 X RACETRACK . X SP-6 X BOX 27 . X 30 chemical components are related to one another. A correlation matrix of those data given in Tables II and III was computed and the partial correlation values are given in Table VII. In addition to the partial correlation coefficients, i t is instructive to consider the values of the squares of the correlation coefficients in order to see how much scatter about a straight line relationship Is present. If the relation-2 ship between the values is absolutely linear an "r " value of 1.0 would 2 be expected and i f correlation was minimal and scatter random the "r ,r value would be about zero. Consequently, although significant correla-tions occur between many of the chemical properties, only sodium is related to chloride, and magnesium to sulfate, in such a way that 90% of the difference in one can be accounted for by the quantity of the other (or so that scatter about a linear relationship is minimized). Conductivity and TDS, conductivity and sodium, and sodium and bicarbonate were related so that 80% of the variation of one was explained by the other. A l l the other significant relationships were a l l such that less than 80% of the variation in one of the pairs was accounted for by the other pair and consequently, would be of l i t t l e predictive use. The relationships in which 80% or greater of the, variation of one pair may be accounted for by the other may be summarized as follows: (1) that conductance is a good measure of total dissolved solids and sodium, (2) that sodium i s present in the same relative concentrations in most of the lakes and varies with concentration, (3) that sodium chloride is like l y the prime source of sodium and chloride, and (4) that magnesium sulfate is the prime source of magnesium and sulfate. In the most concentrated lakes (i.e. , Clinton, GR 2, LB 1, LB 2), the quantities and types of the trace elements could not be determined 31 Table VII. Summary of partial correlation coefficients characterizing the relationships between certain physico-chemical properties of the lakes studied. When r = 0.146,the probability that correlation is due to chance is 0.05. When r = 0.192, the probability of a spurious relationship is 0.01. TEMP CONDUC TDS 0 2 pH Na K Ca Mg CO3 HCO3 C l SO, TEMP CONDUC 0.227 TDS 0.241 0.919 0- 0 100 0.0£4 0.025 pH 0.166 0.437 0.202 0.049 Na 0.181 0.927 0.737 0.040 0.522 K 0.062 0.891 0.849 0.061 0.402 0.742 Ca 0.052 0.169 0.288 -0.024 -0.178 -0.009 0.135 Mg 0.177 0.446 0.755 -0.008 -0.223 0.121 0.494 0.402 C03 0.207 0.760 0.545 -0.005 0.472 0.919 0.471 -0.077 -0.062 HCO3 -0.097 0.720 0.555 0.034 0.550 0.719 0.787 -0.171 0.100 0.526 Cl 0.140 0.831 0.612 0.012 0.530 0.947 0.628 -0.056 -0.008 0.959 0.645 SO, 0.188 0.534 0.802 0.043 -0.148 0.208 0.612 0.424 0.966 0.047 0.192 0.031 32 owing tc blanking by excessive concentrations of sodium and magnesium. However, Bower's Lake appears to be rather different from the remainder of the lakes, since i t has exceptionally high concentrations of borDn, strontium, titanium, and vanadium. The highest concentrations of si l i c o n were observed in Long Lake and the highest concentrations of iron and aluminum were observed in Box 4. Trace elements were determined primarily to evaluate qualitative differences between the lakes and to determine i f large concentrations of elements existed in solutions in additior to the eight major ions given in Table III. On the whole, the lakes are qualitatively similar and the quantities of the trace elements are small. 4. Substrate composition and organic carbon. Substrate compositions of selectee lakes, expressed as a percentage of the dry weight of the sample are given in Table VIII. As would be expected, the data demonstrate a trend tc smaller particle size in the middle of the lake. In general, most of the lakes are characterized by soft material with a preponderance of smaller particles. Since volumes of the samples which were separated for percentage composition by weight vere not determined, no conclusions can be drawn about the relative density or compactness of the mud samples in nature. Qualitatively, except where the gravels were most pronounced, the mud bottoms were soft and flocoulznnt. Differences between lakes in organic carbon content in bottom muds is shown in Table IX. A trend toward increasing organic carbon conf.ent with increase in depth is apparent. 33 Table VIII. Substrate composition. Summary of percentage composition of size fractions of mud sample:; taken from different depths within different lakes. In addition, results are given for nine l o c a l i t i e s in Wentwick Lake, a l l of which occur at the same depth. PARTICLE SIZES (mm) SAMPLE 0.83- 0.59- 0.42- 0.15-LAKE DEPTH 1.98 1.98 0.83 0.59 0.42 0.15 (m) CLINTON . GR-2 -LB-2 0 15. 28 5.50 3. 27 9. 03 18. 27 48.64 LB-1 0 1.48 0.51 0. 52 2. 93 12.49 82. 04 LONE LAKE -BOX 4 0 1.30 2.42 3. 51 7. 08 20. 20 51. 59 BOWER'S LAKE 0 0. 50 0.54 1. 39 6.51 21.39 73.56 LE-1 T TT 0 - —;_ • PHALEROPE 0 7.45 7. 95 7.99 8. 51 25. 93 46.21 BOX 20-21 0 23. 62 8.41 5. 89 6,07 15. 15 40. 86 WHITE LAKE -LE-5 0 1.51 •' 1.03 1.48 14.45 13. 92 67.62 1 0. 28 0. 12 0. 60 3,10 21. 19 74. 72 BOITANO LAKE 0 45. 88 6. 27 5.43 6.86 11.32 24. 24 RUSH 0 7. 06 3. 25 4.62 11.48 30.06 43.53 NR OP BOX 4 0 1.89 2.34 2. 31 . 2. 16 9. 29 81. 98 1 3.29 1.30 1.40 4,53 19. 70 69. 75 2 0.29 0.89 4.05 3.99 12.53 78.22 LE-3 0 1.30 2,11 7.32 7. 71 13.27 68. 26 1 1.35 11.46 8.38 6.17 11.15 61.46 LE-4 0 0. 95 0.48 2. 19 3. 07 24.59 68.69 1 0.16 0. 65 12. 56 5.38 16. 65 64.57 SIX MILE 0 27. 65 4.04 5. 06 5.57 14.51 43. 18 BOX 89 0 42. 95 2.31 1.92 2.64 9.38 40.77 1 13. 29 7.35 5.95 8.29 19. 60 45.49 2 0.00 0. 00 1.03 3. 28 2.42 93. 26 ROCK 0 3. 73 3.49 2. 76 5.95 32. 11 51.94 1 4. 17 3.48 4. 64 5.02 9.80 72.86 GR-3 0 1,75 3.04 4.38 10. 89 27.44 52.47 NR PHALEROPE 0 9. 22 7. 72 14. 23 10.55 25.04 33. 21 1 14.91 10.63 3.05 2. 81 6.35 62. 22 WESTWICK LAKE SOUTH END 0 5. 21 8.45 1. 73 4.32 17.45 62.85 A 0 11.34 2.12 1. 59 3.87 17.99 59.31 B 0 ' 1. 23 4. 88 5. 02 7. 79 29.65 51.43 C 0 0. 57 10. 59 1.35 2.93 20.45 64.11 D 0 0.30 0.46 0. 96 4.07 29. 13 65.09 E 0 15. 16 4. 16 3. 26 8.05 30.85 38. 52 F 0 0.08 0.17 0. 37 2.03 15.98 81.36 G 0 1. 78 1. 07 1. 69 3.51 7.59 84.36 H 0 15. 51 4.35 3. 53 6. 10 15. 26 55. 26 SORENSON LAKE - • . LAC DU BOIS 0 16. 00 4. 95 4. 50 7.61 22.83 43.98 Table VIII (cont.). PARTICLE SIZES (mm) SAMPLE 0.83- 0.59- 0.42- 0.15-LAKE DEPTH 1.98 1.98 0.83 0.59 0.42 0.15 (m) NR OP CRESCENT 0 0. 21 0.29 0.61 1.70 17. 65 79. 51 1 0. 50 0. 76 2.76 1.85 7. 79 86. 31 2 3. 10 2.08 1.97 5.26 17. 94 69. 61 3 0. 43 0.20 0.81 0.88 8. 97 88. 67 BOX 17 0 0. 65 0.22 0.44 0.64 6. 44 91. 44 1 4. 79 1.38 1. 29 1.80 7. 38 83. 32 2 7. 04 4.84 4.86 6.81 27. 29 49. 12 BOITANO NE - — — — — OP BOX 4 0 7. 17 4.43 4. 76 10. 00 20. 33 53. 29 1 3. 08 3.48 3. 28 4.58 16. 41 69. 14 2 1. 88 5.67 1. 99 5.50 28. 25 56. 67 RACETRACK 0 1. 66 3.71 4.37 5.01 16. 37 68. 88 1 3. 61 4.27 4.12 6.34 14. 87 66. 76 2 1. 98 4.01 5.55 11.10 17. 08 60. 25 3 0. 56 0.67 1.25 2.07 13. 23 82. , 20 4 2. 21 7.30 11. 10 4.19 19. 65 55. 52 5 0. 23 0.29 0. 26 0. 76 3. 96 94. ,47 SP-6 0 3. 42 1.29 2.04 3.15 10. 73 79. 37 BOX 27 0 0. 45 0.79 1. 19 1. 77 8. 65 87. ,15 34 Percentage of organic carbon present along sample transects in certain lakes and comparative measures at 0 m in other lakes. Samples were collected in October 1967. DEPTH (m) LAKE CLINTON GR-2 LB-2 15. 68 LB-1 1-47 LONG LAKE BOX 4 . 2.47 ' BOWER'S LAKE 6.97 LE-1 LE-2 PHALEROPE 4.90 BOX 20-21 7. 22 WHITE LAKE LE-5 . 5.20 9.18 BOITANO LAKE 6.52 RUSH 8. 20 NR OP BOX 4 5.24 6.95 LE-3 6.83 7.31 LE-4 9.10 11.18 SIX MILE 8.09 BOX 89 1.92 1. 70 ROCK 1.46 7. 72 GR-3 14.44 NR PHALEROPE 4. 20 10.61 WESTWICK LAKE SOUTH END 12.25 A 8.51 B 11.27 C 6.48 D 3.07 E 5.41 F 6.32 G 15.45 H 8.74 SORENSON LAKE LAC DU BOIS 7.14 NR OP CRESCENT 9.83 7.33 BOX 17 4.27 13.17 BOITANO NE OP BOX 4 5.43 7.42 RACETRACK 4. 92 6.80 SP-6 7. 22 BOX 27 4.36 6.46 5. 03 7.94 8.78 11.05 8.39 9.95 11.52 8.74 6.31 11.1 35 C. Discussion Since the e n t i r e study area was covered by Pleistocene i c e (Holland, 1964) the lakes which were studied ar at most about 10,000 years old. However, Hansen (1947) has concluded that temperatures were higher and p r e c i p i t a t i o n was lower i n the P a c i f i c Northwest some 4,000 to 8,000 year;; ago than today. Therefore, perhaps a better guess at the maximum age of the lakes i s about 4,000 years. Munro (1945) reported that some of the lakes which are considered i n t h i s study were dry during the 1930's and i t seems u n l i k e l y that -nany of the r e l a t i v e l y shallow lakes would have been present: during these warmer and dryer periods. The climate and physiography of t h i s area i s important since the types of lakes, as well as the aquatic and a e r i a l environmental conditions to which C. tentans i s exposed are dependent on them. The present c l i m a t i c conditions of the study area, 5-50 cm p r e c i p i t a t i o n with mean July temperatures of 10 to greater than 2 5° C and mean January temperatures of -10 to 0° C (Figure 1) r e s u l t i n semi-arid conditions within the study area and a "savannah type" b i o t i c community. This, together with the f a c t that greater than 70% of the p r e c i p i t a t i o n c a l l s as r a i n , i ndicates a marked s i m i l a r i t y between the area studied and p r a i r i e s areas studied by Rawson and Moore (1944). According to t h e o r e t i c a l r e l a t i o n s h i p s between p r e c i p i t a t i o n and temperature (see Cole, 1968), the c l i m a t i c conditions should allow for a net rate of evaporation of about 20-50 cm per year, which, i n view of the annual r a i n f a l l , would i n d i c a t e that closed basin lakes should occur i n the area. And, t h i s was found. According to a lake c l a s s i f i c a t i o n based on water temperature that was developed by F o r r e l and l a t e r modified by Whipple (Welch, 1956), most of the lakes studied are temperate lakes of two types: (1) Order 2, i n which the 36 temperature of the bottom water varies, but not far from 4 C, and in which the waters overturn during f a l l and autumn a;.id (2) Order 3, in which bottom waters have tenperatures similar to the surface water and are in continuous circulation except when frozen. The factors which result in these two types of temperate lakes appear to be depth of laks basin and lake chemical concentration 'Tables I and III). S t r i c t l y speaking, White Lake does not f i t in this classification since the lake is meromictic and does not overturn. The analysis of major ions showed that six main chemical types of lekes can be recognised and i f ionic composition, independent of total concentration, can act as a restrictive environmental factor, then one might expect to find differences in faunal composition or abundance, or in genetic constitution among the six types of lakes. Underlying rock formations and rocks present in overlying glacial d r i f t may be of particular significance in accounting, for the types of ions present in these lakes. The correlation analysis of ions showed that sodium and chloride were derived from sodium chloride and that magnesium and sulfate were derived from magnesium sulfate. Cummings (1940), in a study of saline deposits in central British Columbia, reported that sodium carbonate lakes are associated with relatively arid regions which are underlain by Tertiary basaltic flows, that magnesium sulfate lakes are associated with rocks of the lower Cach Creek series ( a r g i l l i t e , quartzite, and limestone) and that sodium sulfate is associated with areas underlain by greenstone and diorite. On the other hand, Cole (1926), in a study of sulfate deposits in western Canada, has proposed that leaching of: sodium sulfate from glacial d r i f t is the most lik e l y explanation of i t s presence in lake waters. The distribution of the B.C. lakes with respect to underlying geological formations is shown in Figure 3 and the findings are in general agreement with those of Cummings. Lakes with high concentrations of sodium carbonate occur over the Fraser Plateau 37 and magnesium sulfate lakes occur primarily in the Thompson Plateau area. Lakes less concentrated of the same chemical types occur sporadically through-out the study area. The lakes studied encompass a range of total concentrations varying crom extremely fresh to about four times the concentration of sea water (Table III). In addition, the data presented in Table. I l l and Figure 9, indicate that any single lake may show considerable change in chemical concentration, this being proportional to the total concentration of the lake. The lakes differ s i g n i f i -cantly with respect to chemical composition and concentration, and composition and concentration vary significantly with respect to season and depth. Since chemical composition and concentration have long been recognized as potentially important determinants of f l o r a l and faunal composition and abundance in inland waters (e.g., Hesse, Allee, and Schmidt, 1951; Allee, Emerson, Park, Park, and Schmidt, 1950), consideration of these factors and their variation is particularly relevant. Examples of seasonal variation in the chemical concentration of the surface waters of the lakes are given in Table X. It is apparent that concentrations rise from a minimum during spring to a maximum during late f a l l and winter. Further, i t is especially apparent that the range of variation within a lake is proportion to the total chemical concentration of that lake. Examples of long term changes in the chemical concentrations of the lakes are given in Table XI. Finally, chemical composition has a primary effect on conductivity (Figure 7). Since lakes with the same conductivity may differ by chemical composition, they may also differ in chemical concentration (and osmotic pressure). Therefore, i f concentration per se is a regulating factor, then 38 Table X. Seasonal variation in the conductivity of surface waters. Conductivities are expressed in micromhos/cm at 25° C. LAKES DATES GR-2 LONG L. BOX '4 WHITE L. BOX 20-21 BOITANO 7 April 1966 19,000 4,800 3,300 4,000 1,100 400 10 May 1966 30,100 9,110 6,613 5,090 4,680 3,900 7 June 1966 33,000 9,440 9,440 4,720 5,380 4,000 3 July 1966 40,000 10,450 11,017 • 5,068 6,220 4,120 28 September 1966 45,000 12,200 11,350 4,750 6,020 4,200 16 February 1967" 57,320 12,033 5,360 . 2 May 1967 27,500 8,400 6,850 4,780 5,000 4,480 3 June 1967 37,000 10,200 9,553 5,780 6,000 4,130 24 June 1967 47,500 11,000 10,603 5,080 6,530 4,350 23 July 1967 48,280 11,150 11,850 5,750 6,650 4,350 9 September 1967 60,500 12,600 13,300 5,380 7,250 4,500 7 October 1967 52,175 12,540 13,480 5,320 7,260 4,640 39 Table XI. Long; term variation in the conductivity of surface waters. Conductivities are expressed in mLcromhos/cm at 25° C. YEAR MAY LAKE 1961 1962 1963 1966 1967 RANGE GR-2 21,000 37,500 30,100 27,500 10,000 LONG LAKE 20,000 9,110 8,400 11,600 BOX 4 12,000 6,610 6,850 5,390 WHITE LAKE 4,900 7,500 5,090 4,780 2,720 BOX 20-21 5,750 4,680 5,000 1,070 BOITANO LAKE 4,000 4,500 3,900 4,500 600 CEMBER . > GR-2 60,000 50,000 45,000 60,500 15,500 LONG LAKE 29,000 12,200 12,600 16,800 BOX 4 20,000 10,000 14,000 11,350 13,300 10,000 WHITE LAKE 9,000 6,000 9,000 4,750 5,380 4,250 BOX 20-21 12,000 5,100 8,500 6,020 7,250 6,900 BOITANO LAKS 5,000 9,000 4,200 4,500 4,800 4 0 Figure 7. Conductivity as a function of ionic composition. Conductivity data used to draw this graph were collected from the lakes studied. The lake classification, described previously, was used to group the data according to the predominate ions in lakes. s a i 41 account must be taken of the ionic composition of the lake(s). In summary, a.series of environments ranging chemically from very dilute to relatively concentrated exist in the study area. The lakes studied form primarily six chemical types when classified according to their major anions and cations. Further, i t is apparent that both total concentra-tions and concentrations of individual ions may be altered at particular depths by redistribution of the ions due to mixing and freezing out (with concentrations alternately increasing in surface waters and lowering in bottom waters and then lowering in surface waters and rising in bottom waters). The range of variation in chemical concentretions of surface waters within lakes may be particularly pronounced, depending on the total chemical concentration of the lake. Further, chemical concentrations of the lakes undergo long-term cyclic fluctuations due tc long term changes in rate of evaporation. A l l of these conditions may interact to provide aquatic environ-ments which would differ either v e r t i c a l l y or totally with respect to their osmotic properties. Temperature, st r a t i f i c a t i o n of the lakes and subsequent depletion of oxygen in bottom waters (and in some cases corresponding increases in hydrogen sulfide concentration), and pH are other possible factors by uhich the lakes may differ. Preponderantly, differences in chemical concentration and composition appear to be the main attributes by which the lakes differ in a regular and predictable fashion. IV. Occurrence and Abundance of C. tentans A. Methods of Determination of Occurrence and Abundance Samples of mud containing larval chironomids were collected with a 15 x 15 cm Ekman dredge and sieved through a screen with a 0.56 mm mesh 42 s i z e and then p r e s e r v e d i n 107o f o r m a l i n . The l a r v a l chironomids were separated from the mud i n the l a b o r a t o r y u s i n g the sugar f l o t a t i o n t e c h n i q u e . N e i t h e r sampling e f f i c i e n c y nor s o r t i n g e f f i c i e n c y were determined. The occurrence o f C. tentam; was determined by thorough qual i ta t ive- , sampling and by a n a l y s i s of q u a n t i t a t i v e samples. The abundance o f C. t entans was es t imated by c o l l e c t i o n o f d u p l i c a t e samples taken at meter i n t e r v a l s i n depth (jL.e_^_, 0 m, 1 m, e t c . ) a long a s i n g l e t r a n s e c t l i n e . Samples were taken to the middle o f the l a k e . A c c o r d i n g to the method d e s c r i b e d by Mundie (1957)/ the accuracy o f the e s t i m a t e s o f abundance of C. t entans i s about 50%. Ten r e p l i c a t e . s a m p l e s were taken at each of four l o c a l i t i e s w i t h i n Westwick Lake and a n a l y s i s o f these samples i n d i c a t e d t h a t d u p l i c a t e samples would p r e d i c t s p e c i e s occurrence w i t h 99% accuracy . Occurrence o f p o p u l a t i o n s i n d i f f e r e n t lakes was o r i g i n a l l y determined i n order to e v a l u a t e chromosoiiie polymorphism i n r e l a t i o n to the environment ( w i t h emphasis p l a c e d on l a k e c h e m i s t r y ) . D i s t r i b u t i o n s w i t h i n l a k e s were determined i n order to i n s u r e t h a t the p o p u l a t i o n s l i v i n g w i t h i n the lakes were samples adequate ly f o r c h a r a c t e r i z a t i o n of: chromosome polymorphism f o r t h a t environment ( i . e . . , the sample should be r e p r e s e n t a t i v e o f the m a j o r i t y o f the l a r v a e l i v i n g w i t h i n a l a k e and .not j u s t a s m a l l p o r t i o n of the p o p u l a t i o n ) . T h e r e f o r e , s i n c e these data were not c o l l e c t e d f o r d e f i n i t i o n o f the ecology o f C. t e n t a n s , sampling regimes used i n the d i f f e r e n t l a k e s were not always e q u i v a l e n t . F u r t h e r , the c o n c l u s i o n s presented h e r e i n are drawn from c o r r e l a t i v e r a t h e r than e x p e r i m e n t a l d a t a and c a u t i o n should be used s i n c e s p u r i o u s r e l a t i o n s h i p s might r e s u l t from sampling e r r o r (see Mundie , 1957). 43 Larvae of different species of chironomids were identified by use of keys i n Roback (1957) and identifications of larvae of C. tentans were confirmed by cytological analysis of giant chromosomes. Survival of larvae of C. tentans in waters of different chemical concentrations at 5° C was determined by transferring larvae from lake water in which they were collected (3,000 micromhos conductivity) to different concentrations and dilutions o:: the same lake water. Five 4th instar larvae were placed in 10 ml of each concentration of lake water and ten replicates of each concentration were prepared. The larvae were not fed throughout the experiment and the experimental water was not changed. The experiment was performed in continuous darkness. The Kendall Rank correlation coefficient was used in the analysis of abundance as described by Siegel (1956). Results 1. General distribution. The distribution of the species which is relative continuous from near Moscow, U.S.S.R. to southern Alaska (Figure 8), suggests that C. tentans has a holarctic range; however, C. tentans ha;3 not been reported from Siberia. In British Columbia, the species is known only from the central interior region, which is characterized by semi-arid climatic conditions and savannah type vegetation. 2. Distribution in the study area. General features of the occurrence of larvae of.C. tentans in different lakes in the study area and within individual lakes are given in Figures 9 and 10, respectively. C. tentans occurs in 17 of the 32 lakes which were studied and the species is only a temporary resident in three of these (Boi.tano L. , Rush, and Box 17). The 44 F i g u r e 8. World-wide d i s t r i b u t i o n o f C. tent ans. 45 F i g u r e 9. Occurrence of larvae of C. tentans w i t h respect to the chemical c o n c e n t r a t i o n of lake waters (as measured by c o n d u c t i v i t y ) . The maximum varic'.tion i n c o n d u c t i v i t y observed i n each of the lakes i s shown by the h o r i z o n t a l bars. The occurrence of C. tentans i n the lakes i s i n d i c a t e d by the hatching w i t h i n the h o r i z o n t a l bars. • CLINTON • GR 2 • LB 2 . • LB I C ~ 1 LONG L. ~ 1 BOX 4 LZr BOWERS L. • LE I • LE 2 (ZD PHALEROPE — ! B Q X 20 -21 1 WHITE L. 0 LE 5 Wim BOITANO L. 0 LE 4 0 SIX MILE WA BOX 89 mm ROCK wm GR 3 ^ NR PHALEROPE WESTWICK L. 0 SORENSON L. 0 LAC DU BOIS 0 NR OP CR n BCX 17 0 O P BOX 4 ESS RACETRACK 1 = 3 SP 6 ] BOX 27 O O L U < WZZA RUSH ^ NR OP BOX 4 0 LE 3 IO IOO CONDUCTIVITY (millimhos at 25°C 46 F i g u r e 10. Occurrence o f l a r v a e o f C. tentans and o f o ther s p e c i e s o f ch ironomids w i t h r e s p e c t to depth. Surfaces e n c l o s e d by the heavy b l a c k l i n e s i n d i c a t e the abundance of C, t e n t a n s , w h i l e s u r f a c e s enc losed by the f i n e l i n e s i n d i c a t e the t o t a l abundance of other c h i r o n o n i d l a r v a e . The presence o f a v e r t i c a l b l a c k bar i n d i c a t e : ; t h a t C. tentans were c o l l e c t e d o n l y at t h a t depth. Abundances between i n t e g e r depth i n t e r v a l s were determined by e x t r a p o l a t i o n . 47 obvious implication of Figure 9 is that not a l l lakes are acceptable habitats for the larvae or .pupae. Qualitative observations made coneirning the presence or absence of C. tentans showed that, in general, the l>est indicator of the presence of the species was found to be the. marsh reed, Scirpus acutus. The reed has about the same distribution both between and within lakes as does C. tentans. When Scirpus was present, C. tentans was usually found in flocculent mud on the lakeward side of the reeds. The mud in which the species lieved was usually a dark brown or black, indicating reducing as opposed to oxidizing conditions. The mud was always flocculent as opposed to being firmly packed. C. tentans was usually found to be absent from lo c a l i t i e s in which the bottom was covered by heavy algal mats (e.g. , Lac du Bois, Box 17, Near Opposite Crescent, and at times Westwick Lake). The data presented in Figure 10 indicate that C. tentans does not occur at depths greater than two meters, although other species of chironomids do occur at these depths. Further, the greatest numbers of C. tentans occur at shallower depths. The occurrence of C„ tentans with respect to water chemical concentra-tion (Figure 9) indicates that larvae of C. tentans can occur in lakes in the conductivity range of 500 to 4,500 mlcromhos (measured at 25° C). Conductivity is used as a measure of chemical concentration since (1) the data are easy to collect, (2) the correlation analysis of chemical factors indicated a high correlation between conductivity and concentration, and (3) the lakes in which C. tentans larvae occur are not sufficiently concentrated for chemical composition to result in wide discrepancies between conductivity measured and total chemical concentration (see potential 4 8 e f f e c t i n Figure 7). Occurrence of larvae with respect to concentration of the eight major ions i s given i n Figure 11. In t h i s f i g u r e , data for each of the lakes have been combined and occurrence i s presented only with respect to the concentrations of i n d i v i d u a l ions. Larvae occur i n a l l of the s i x major chemical types of lakes (see Figures 6 and 9); however, they are only temporary residents i n the sodium s u l f a t e type of lake. No s p e c i a l s i g n i f i c a n c e should be attached to the temporary occurrence i n sodium s u l f a t e water since the only lake (Rush lake) of that type had a concentra-t i o n which approached the maximum tolerance of the species. The occurrence of larvae i n r e l a t i o n to c e r t a i n other water properties (temperature, pH, minimum dissolved oxygen and hydrogen s u l f i d e ) , food a v a i l a b i l i t y and substrate composition are shown i n Tables XII and XIII, and Figure 12, r e s p e c t i v e l y . In these two tables and one f i g u r e , data are grouped into categories associated with the presence of absence of -C. tentans. The co-occurrence of C. tentans with other species of chironomids i n a s i n g l e lake i s shown i n Figure 13. The species reported i n Figure 13 are also present i n the other lakes that contain C. tentans and comprise the bulk of a l l . o t h e r chironomids shown i n Figure 10. 3. Abundance„ Estimates of abundance were subject to large sampling errors (^ 507o) and the resultant v a r i a t i o n associated with these estimates precluded d i r e c t use of exact estimates to evaluate even p o t e n t i a l l y important r e l a t i o n s h i p s between abundance and environmental f a c t o r s . Therefore, ;some form of analysis was required i n which more emphasis could be placed on general trends and less on absolute numbers. The technique 4 9 Figure 11. Occurrence of larvae of C. tentans with respect to concen-trations of the major ions. The ranges of concentrations of ions present in the entire series of lakes are shown by open bars, while the ranges tolerated by C. tentans are shown by the hatched portions of the bars. I 1 1 ^ l i I I I I 1 I 1 ! i o o o o o o O o •7-- 'v 50 Table XII. Occurrence of larvae of C. tentat.s with respect to water temperature, pH, minimum dissolved oxygen, and presence of hydrogen sulfide. Conditions associated with the presence of G. tentans are compared to these conditions associated with i t s absence from specific l o c a l i t i e s and from entire lakes. Means and standard deviations of each factor are listed for spring and f a l l , 1966. TENTANS1 LAKEo ENVIRONMENTAL C. TENTANS C. TENTANS NON TENTANS' FACTOR PRESENT ABSENT LAKE TEMPERATURE (C' C) Spring 12.8 + 1.4 LI. 9 + 1.5 13.4 + 2.7 F a l l 19.5 + 1.4 17.5 + 1.9 20.0 + 3.8 • pH Spring 8.58+0.31 8.46+0.31 8.86+0.73 Fa l l 8.79 + 0.26 8.70 + 0.31. 9.00 + 0.57 Oxygen (cc/1) Spring 5.16 + 0.96 5.15 + 1.46 5.52 + 1.36 F a l l 3.95+2.77 2.48+2.36 5.02+1.44 H2S (ppm) Fa l l 0.02 + 0.07 1.08 + 1.92 0.00 51 Table XIII. Occurrence of larvae of C. tentans in relation to percentage of organic carbon. As in Tables XI and XII, means and standard deviations of the condition's associated with the absence of C. tentans are included for comparison. TENTANS' LAKES C. TENTANS C. TENTANS NON-TENTANS' . ... PRESENT . ABSENT .....LAKES PERCENTAGE ORGANIC CARBON 16.13+3.33 15.10+4.21 14.34+4.24 52 Figure 12. Occurrence of larvae of C. tentans in relation to sub-strate composition. As is Figure 11, conditions associated wil:h the presence of C. tentans are compared to conditions associated with i t s absence. Means and standard deviations are given for each of the three categories for the six sine groups of mud particles which were analyzed. PERCENTAGE COMPOSITION O O L P . 60 (30 OJ o o * * 00 en OJ 00 r o O O 3 00 oo o o O ...a „ O r — 3 o O 00 o Z > O D 3 ^ ^ o o m ^ m o o —I > o o . > m o o r n o o > 7^ o o o o 53 Figure 13. Occurrence of larvae of in Westwick Lake during larvae of other species C. tentais at different depths October 1.966 in relation to of chironomids. PERCENTAGE COMPOSITION 54 employed WE.S conversion of the interval estimates to ordinal rank values and the rar..k values were then analyzed by the Kendall rank correlation coefficient: test (Siegel, 1956). The crux of this analysis is the grouping of the data for conversion to ordinal rank values. Exact estimates of abundance for each locality (Table XIV. column 1) were grouped into categories which differed by about * 507o. Estimates grouped together in these categories were then considered to be samples drawn from the same population with variance being due to sampling error (Table XIV, column 2). Finally, the categories were assigned ranks as described in the Kendall rank correlation test (Table XIV, column 3) f-nd correlation coefficients ware calculated. The results of the Kendall rank correlation analysis between the abundance of larvae of C. tentans and environmental factors are given in Table XV. Analysis of those abundancas of larvae collected from 0 m in the different lakes indicate that conductivity, pH, potassium, magnesium, chloride, sulfate, three of tie six substrate fractions, and the amount of organic carbon available show correlation with abundance. Analysis of those abundances of larvae CDllected from different depths in the lakes indicated that conductivity, minimum dissolved oxygen, sodium, chloride, the number of larvae of other species of Chironomus, one of the substrate fractions, and the amount of organic carbon available are each correlated with abundance. Consequently, the differences in abundance with depth appears to be related primarily to minimum dissolved oxygen and to the numbers of larvae of other species of Chironomus. Discussion The world-wide distribution of C. tentans suggests that the general 55 Table XIV. Observed abundances of larvae of C. tentans at sample lo c a l i t i e s and groupings used for the Kendall rank correlation coefficient analysis. COLUMNS LAKES SAMPLE DEPTH (m) 1 ABUNDANCE (#/m2) 2 3 GROUTED ASSIGNED CATEGORIES . . . RANK LE-3 0 559 NR OP BOX 4 0 495 OP BOX 4 0m 1 1 - 215 BOX 89 0 86 LE-3 0 m 3.5 ROCK 1 22 OP BOX 4 1m 3.5 NR PHALEROPE 0 215 WESTWICK LAKE "C" 0 m 3.5 NR OP CRESCENT 0 43 NR OP BOX 4 0m 3. 5 1 43 2 22 NR OP BOX 4 1m 8 BOX 17 0 22 WESTWICK LAKE "A" 0 m 8 OP BOX 4 0 1,055 "B" 0 m 8 1 495 "H" 0 m 8 2 129 NR PHALEROPE 0 m 8 RACETRACK 0 129 1 65 RACE TRACK 0 m 16 2 43 OP BOX 4 2m 16 WESTWICK LAKE BOX 39 0 m 16 SOUTH END 0 86 WESTWICK LAKE "SE" 0 m 16 A 0 215 "G" 0 m 16 B 0 . 215 ' "A" 1 m 16 C 0 323 "G" 1 m 16 G 0 86 RACETRACK 1 16 H 0 237 2 m 16 WESTWICK LAKE NR OP CRESCENT ' 16 A 1 151 NR OP CRESCENT 1 m 16 C 1 22 D 1 22 WESTWICK LAKE "C" 1 m 25 E 1 22 "D" 1 m 25 G 1 65 "E" 1 m 25 H 1 22 "H" 1 m 25 ROCK 1 m 25 NR OP CRESCENT 2m 25 BOX 17 0 m 25 56 Table XV. Correlation between the abundance of larvae of C, tentans and specific environmental factors. Significance with P 0.05 is marked (*), with P 0.01 (**),.and with P 0.001 (***). ANALYSIS OF ALL ANALYSIS OF ALL ENVIRONMENTAL LOCALITIES LOCALITIES FACTOR AT ALL DEPTHS AT 0 m CONDUCTIVITY 0.276— 0.416** TEMPERATURE 0.212 0.354 pH 0.049 0.717 *** MINIMUM DISSOLVED OXYGEN -0.263** -0.054 SODIUM 0.277 ** 0.295 POTASSIUM 0.025 0.457 ** CALCIUM -0.019 0.242 MAGNESIUM 0.108 0.497** CARBONATE 0.158 0.322 BICARBONATE 0.085 0.215 CHLORIDE 0.380*** 0.564*** SULFATE 0.057 0.551 *** OTHER SPECIES OF CHIRONOMUS -0.313 ** 0.176 SUBSTRATE COMPOSITION 1.98 0.064 .0.086 1.98 - O.J53 0 . 3 3 2 ** 0 . 221 0.83 - 0.59 0.178 0.491 ** 0.59 - 0.^ 2 0.223 0.381 * 0.42 - 0.15 0.257 0.356 * 0.15 -0.109 -0,270 PERCENTAGE ORG/JSIIC CARBON AVAILABLE PER C. TENTANS 0.544 *** 0.736 *** 57 distribution of the species may be limited by temperature. The distribution of C. tentans in central British Columbia indicates the species is restricted to lakes that cccur in savannah vegetational zones (see also Acton and Scudder, 1969). Since central British Columbia is subjected to considerably lower temperatures than the southern extremes of the world-wide distribution of the spe.cies, but considerably higher temperatures than the northern range of the species, i t is unlikely that temperature alone is a particularly important factor within the study. However, temperatures in any specific area might exceed the maximum or minimum extreme tolerated by the species and it s effect cannot be completely disregarded. In general, the habitat observations on C. tentans in central British Columbia are in agreement with other qualitative observations reported in the literature. Sadler (193!) reported that larvae of C. tentans occur in pools, ponds, shallow warm lakes:, and sluggish streams. He also reported that the greatest abundance of larvae was found in stagnant water with a good deal of pollution. Palmen and Aho (1966) summarize other-literature which report similar observations. Therefore, i t appears that the habitat of the study area is roughly comparable to the "normal type" of habitat which the species enjoys elsewhere in i t s range. Within the study area, the analysis; of the lake environments suggested that the prime difference between the lakes was their chemical concentration and composition, and the restriction of C. tentans to lakes with con-ductivities in the range of 500 to 4,500 micromhos (Figure 9) certainly suggests that chemical concentration may act as a restrictive factor. 58 The exact effect of concentration on the survival of the species was investigated by placing several larvee in waters of different chemical concentrations at 5° C (see Figure 14). The results of this experiment indicated that at 5° C water with conductivities ranging from 2.5 to 8,000 micromhos (at 25° C) could be tolerated equally well. When the experiment was terminated, the test chambers were removed to room temperature and chironomids in water- with conductivities in excess of 4,000 micromhos died within two days. These findings supported the f i i l d observations which indicate that C. tents.ns w i l l not tolerate waters with conductivities greater than 4,500 micromhos and that the maximum tolerance of the species may be modified by temperature. However, the experimental studies do not offer any explanation for the absence of the species from waters with conductivities of less than 500 micromhos. The absence of C. tentans from depths greater than two meters in lakes within the study area is in agreement with findings reported by Palmen and Aho (1966). However, C. tentens has been reported from consider-ably greater depths (see Palmen and Aho, 1966:230). Since particular significance is attached to the absense cf C. tentans below two meters observed in this study, i t would be interesting to know i f the reports of C. tentans at depths greater than two meters are attributable to mis-identification of the species as suggested by Palmen and Aho (1966). If C. tentans is restricted to depths less than two meters then the conclusions reported here would assume more general importance. Regardless, the data reported here indicate that conditions associated with depth exclude C. tentans from depths greater than 2 m in lakes within the study area. In addition, the absence of C_„ tentans from specific localities with depths less than two meters (see Figure 10) indicates that there are additional 59 Figure 14. Experimental analysis of survival of larvae of C. tentans in waters with different chemical concentrations at 5° C. The actual conductivities of the waters used at 5° C are given together with the conductivities of those same waters when measured at 25° C. PERCENTAGE MORTALITY — r o W A L n Q ~ 'v l C D G O O O O O O O O ~ L — i — I i 1 i I i 1 i I » I i I i I — Oo cy •o / 0 0 O J : — — f O L^J LP o o o o o . o o o ° — M w ^ (J 1 Cr~ OD r o u r O O O O O O O O n O O O . O O O O O O O O O O O O O o LP "21 n Cn — o — j 3 O 60 environmental conditions, not associated with depth, that can also exclude the presence of the species. The absence of C. tentans below two meters suggests that tempersture, dissolved oxygen, or other factors which vary regularly with depth might act to regulate the occurrence or abundance of the species and the absence of C. tentans at depths less than two meters suggests that co-occurrence with other species, food a v a i l a b i l i t y , substrate composition, and other factors which may vary within lakes, but which do not vary regularly with depth may also regulate the occurrence aad abundance of the species. It may be asserted that chemical concentration can be responsible for the differential distribution of C. tentans between lakes;, however, increase i i concentration with depth in the lakes in which C. tentans does occur is substantially less than the range of seasonal variation in concentration present in the surface waters and thus i t is unlikely that concentration is responsible for the differential depth distribution of the species. Of course, since chemical concentration results from differing amounts of individual ions, the effects of individual ions cannot be ignored. The occurrence of C. tentans with respect to concentrations of individual ions is given in Figure 12. As stated in the description of methods, data for each of the lakes were combined and the occurrence of the species with respect to concentration of individual ions in specific l o c a l i t i e s cannot be evaluated. Thus, caution should be used in inter-preting this figure since the data do not allow for the effect of the interaction of the ions, nor for the effect that total chemical concentra-tion might have on the tolerance of individual ions. However, since the 61 species occurs in each of the six chemical types of lakes, chemical composition is not considered to be a particularly important regulator of C. tentans. The analysis of the effect of temperature, pH, minimum dissolved oxygen, hydrogen sulfide, food a v a i l a b i l i t y , and substrate composition., involved comparison of the conditions which the species tolerated with those conditions associated with the absence of the species. In this analysis, the categories used for the comparison were "tentans' lakes" (conditions associated with the presence of the species), "non-tentans; lakes" (conditions associated with the complete absence of the species from the lake, aid "tentans absent" (conditiors associated with the absence of the species from particular l o c a l i t i e s within lakes in which the species was present in other parts). The potential importance of the environmental factor as a restrictive factor was then evaluated by comparing the means and standard deviations of each of the variables for the three categories. Any significant difference between the conditions associated with the presence and the absence of the species could account for the absence of the species, However, the lack of any significant difference suggests that the particular environmental factor might not be important or at least that judgement should be reserved. Palmen and Aho (1966) reported that: the occurrence of C. tentans in brackish water along the coast of Finland appears to be restricted chiefly by temperature and suitable substrate (i.e_., that the species avoids high temperatures and substrates composed of very fine muds). They also suggest that lack of sufficient food might also affect the occurrence of the species. However, the amount of food (as measured by percent of organic carbon available), substrate composition and temperature did not appear to be of any particular significance within 62 the study area. In fact, hydrogen sulfide proved to be the only environ-mental factor which showed any significant difference. However, since the presence of hydrogen sulfide is dependent on the absence of dissolved oxygen, the effact of the two cannot be clearly separated. In addition, the water properties discussed here and the amount of organic carbon w i l l vary seasonally and since these factors have not been evaluated during a l l seasons, i t remains possible that the quantity of any one of the factors might be limiting during a different season. The data reported for Westwick Lake concerning the distribution of C. tentans with respect to other species shows that while other species may be present at zero or one meter, the absence of C. tentans is inversel; related to the abundance of the three other species of Chironomus. This reciprocal relationship may indicate that C. tentans and the other species are merely responding in opposite manners to the same environmental factor; but i t also suggests that interspecific interactions might be responsible for the absence of C. tentans. In contrast to the evaluation of the general importance of environ-mental factors (as considered thus far), i t Is also interesting to know which of these individual environmental factors might be responsible for the absence of the species from specific l o c a l i t i e s and lakes. This infor mation is considered in Figures 15 and 16,respectively. In these figures, those values for individual environmental factors associated with specific l o c a l i t i e s that are outside the ranges tolerated by C. tentans are denoted by closed circles. The l o c a l i t i e s above two meters, at which C. tentans might be expected to occur, but from which the species is in fact absent, are characterised by having different substrate compositions and by having other species of the genus Chironomus present in greater numbers than are 63 Figure 15. Erwironmental factors potentially responsible for the absence of larvae of C. tentans >:rom specific l o c a l i t i e s in lakes in which the species does occur. The presence of an environmental factor outside the range of that factor tolerated by C. tentans i ; ; indicated by a dark circle. RACETRACK BOX 17 NROP CR WESTWICK WESTWICK D NR PHALEROPE ROCK BOX 89 r-n i OI NR OP BOX 4 r -m cr — O — to O to — -• O DEPTH (m) CONDUCTIVITY TEMPERATURE pH '© © © OXYGEN © © © ® @ H 2 S No CHEMICAL COMPOSITION . K CHEMICAL COMPOSITION . Co CHEMICAL COMPOSITION . Mq CHEMICAL COMPOSITION . co 3 CHEMICAL COMPOSITION . H C O 3 CHEMICAL COMPOSITION . Cl CHEMICAL COMPOSITION . so4 CHEMICAL COMPOSITION . > 198 % MUD PARTICLE SIZE (mm) @ © © 0 8 3 3 " 1-98 % MUD PARTICLE SIZE (mm) 0 . 5 8 9 - 0 - 8 3 2 % MUD PARTICLE SIZE (mm) 0 4 I 7 - 0 - S 8 8 % MUD PARTICLE SIZE (mm) ® © © O I 4 7 0 - 4 I 6 % MUD PARTICLE SIZE (mm) < O-147 % MUD PARTICLE SIZE (mm) % ORGANIC C ® © © 0 © 9 © OTHER SPECIES OF | CHIRONOMUS PRESENT 64 Figure 16. Environmental factors potentiall)' responsible for the absence of larvae of _C. tentans from lakes. As in Figure 15, the presence of an environmental factor which exceeds the range of that factor tolerated by C. tentans is indicated by a dark circle. BOX 27 CO cr LAC DU BOIS RUSH L. BOITANO L. WHITE L. 1 CD o X ro o 1 PHALEROPE i — m ro i — m • BOWERS L. BOX 4 LONG L. r~ CD I -IP I O C?) X3 CLINTON © ® © ® @ © ® m © © ® @ @ © CONDUCTIVITY ® @ TEMPERATURE ® ® ® @ © pH O X Y G E N H 2 S © © ® ® © © @ @ Na n ® i i ® ® ® ® © © © ® K X m © © Ca n > @ ® Mq 1— n @ ® © ® @ C 0 3 OMPOSI © © © © HCO3 OMPOSI ® @ ® @ ® ® © Cl —1 O © s o 4 z > 1-98 . 0 0-833-1 .98 MUD 1 SIZE ( 0 - 5 8 9 - 0 8 3 2 MUD 1 SIZE ( 0 - 4 I 7 - 0 . 5 8 8 PART mm) O . I 4 7 - 0 - 4 1 6 ICLE <O. I47 ICLE % ORGANIC C 65 found when C. t. en tans is present. That the amount of available food and the presence of hydrogen sulfide exceed the .range tolerated by C. tentans at only two l o c a l i t i e s suggest that these factors may not be of general importance. At depths greater than two meters, absence of C. tentans is associated with the presence of hydrogen sulfide, presence of other species of the same genus and to a lesser extent, lower concentrations of oxygen, lakes from which the species is entirely absent, or present only temporarily, are characterized by having concentrations of specific ions and total concentrations which exceed the ranges tolerated by the species. Temperature, substrate composition, and the amount of food available may also be responsible for the absence of C. tentans from different lakes, but again these factors do not appear to be of general importance in the study area. A potential source of error concerning the correlation between the abundance of C. tentans and environmental factors which should be under-stood, but which cannot be corrected for, is sampling error. Sadler (1935) reported that the head capsule diameter of 4t.h instar larvae varied from 0.71-0.74 mm,while the diameter of head capsules of 3rd instar larvae was less than 0.40 mm. Since a screen with a mesh size of 0.56 mm was used to separate the chironomids from raw mud samples, only 4th instar larvae would be collected quantitatively. Of course, i f populations of C. tentan;; occurring in the different lakes can be assumed to be composed of the same relative numbers of developmental stages and i f the populations over-winter as 4th instar larvae (the samples were collected during October; water temperature was about 5° C), then the assumption can be made that the estimates of 4th instar larvae, in fact, reflect differences in the abundance of C. tentans present in the lakes. Alternatively, i f the populations of G. tentans present in the different lakes are composed of the same relative numbers of developmental stages, then the assumption can 66 be made that the samples at least i n d i c a t e the r e l a t i v e abundance of tiie other species of chironomids sampled. No assumption can be made of tha accuracy with which d i f f e r e n t species were sampled. F i n a l l y , the abundance of C. tentans, and i t s r e l a t i o n to the r e l a t i v e numbers of other chiroaomids and food a v a i l a b l e , applies only to the Lime of year when the samples 'were c o l l e c t e d . The remainder of the environmental factors were either measured seasonally, so that maximum v a r i a t i o n could be accounted f o r , or the character (substrate composition) did not: vary. The r e s u l t s of the analysis of c o r r e l a t i o n between environmental factors and abundance of C. tentans indicate that abundance i s correlated with the amount of organic carbon a v a i l a t l e and with the concentration-:-. of i n d i v i d u a l ions. The decrease i n abundance of C. tentans with depth appears to be r e l a t e d to decreased dissolved oxygen and increasing chemical concentration. The low values of c o r r e l a t i o n between abundance and the environmental factors examined indi c a t e that no great importance should be a t t r i b u t e d to any of the r e l a t i o n s h i p s . However, any s i g n i f i c a n t c o r r e l a -t i o n values do i n d i c a t e p o t e n t i a l l y important environmental factors and do provide a basis for constructing an e f f i c i e n t experimental program for evaluating the e f f e c t of environmental factors on the abundance of C. tentans. In summary the l i f e h i s t o r y data reported by Sadler (1935) indi c a t e that older larvae, pupae, and adults may be subject to predation. The high mortality observed i n 4th i n s t a r larvae suggests that natural s e l e c t i o n may be p a r t i c u l a r l y important at t h i s stage of development. In ad d i t i o n , the environment may a f f e c t the rate of development (Sadler, 1935; Engelmann and Shappirio, 1965), which i n turn could a f f e c t the rate of response to, or adaptation to, forces of natural s e l e c t i o n . The e c o l o g i c a l data suggest that i n the study area, lake chemistry (either t o t a l concentration or differential concentrations of individual ions, or both) is the most important factor regulating the occurrence of C. tentans in different lakes, while oxygen content, hydrogen sulfide content and co-occurrence with other species of Chironomus may be of considerable importance in regulating the occurrence of C. tentans within lakes. Abundance in different lakes appears to be regulated by water chemistry, while differences in abundance within a lake appears to be related to chemical concentration and dissolved oxygen. In conclusion, the significance of the effect of any one environmental factor on the occurrence or abundance °f tentans can only be considered with respect to the extent of the range of distribution considered. Thus, temperature appears to be important Ln the world-wide sense, chemical concentration appears to be most important in a comparison between lakes, and dissolved oxygen and other species of Chironomus appear to be most important within lakes. Inversion Polymorphism in Natural Populations of C. tentans with Respect to Environmental Factors. ~~ ~ Introducti on The oroblems undertaken in this part of the study were (1) to describe inversion frequencies in the pcpulations inhabiting the 12 lakes which were studied intensively, (2) to determine i f the inversion frequencies of different populations do differ significantly, and (3) to determine what might be responsible for the differences in inversion frequences of different populations, i f these occur. The Inversions that were studied were 1 Rad and 1 Rade (see Figure 17), which occur on the right arm of the f i r s t chromosome, and inversions 4c, 4bc, and 4b which occur on the fourth chromosome. The frequency of the uninverted chromosome 4 was also measured. The locations of the inversions 68 Figure 17. Drawings of chromosome 1: homozygous for inversion 1 Rad, homozygous for inversion 1 Rade, and heterozygous for inversions 1 Rad and 1 Rade. CHROMOSOME I I Rod• I Rad KEYS, KEYS Rad I Rade • I Rade KEYS KEYS 1 Rade I Rad • I Rade 69 on the chromosomes are given by Acton (1959)„ The notation used to describe the inversions of.the 1st chrorrosome is modified slightly from that used oy Acton (1962) in that 1 RaRdRe is altered to 1 Rade and 1 RaRd is altered to 1 Rad. Inversion 1 Rade includes inversion 1 Rad and consequently both can never be present on the same chromosome of the pair. Moreover, since 1 Rad and 1 Rade are the only inversions present in any appreciable frequency on the right arm of Chromosome 1, and since either one or the frequency of the other is always present one inversion defines tha frequency of the other (i.ja. , 1 Rad plus 1 Rade equal 100), of the inversions). B. Methods Larvae of C. tentans were collected with an ordinary dip net from depths between 0 m and 1 m. Scoops of bottom mud and debris were collected and the mud was washed from the net by swirling the net under water: C. tentans larvae were then picked by hand from among the other chironomids and the remaining debris^ The twelve lakes studied for inversion poly-morphism covered the entire range of concentrations occupied by C. tentans in the study area. Samples of C. tentans were collected from at least a single locality in each of the twelve lakes for cytogenetic analysis. Two of the twelve lakes were sampled at more than one locality around the margin of each in order to evaluate horizontal variaticn in inversion frequencies (or isolation within lakes) and one of the lakes was also sampled at approximately monthly intervals from May to November 1967 in order to evaluate temporal variation in. inversion frequency within a single lake 70 at a single locality. Squa;:h preparations of the giant chromosomes were made directly from live specimens in the manner described by Acton (1955). In the experimental analysis of tha effect of biotic factors on inversion frequency, 175,000 eastern brook trout (Salvelinus fontinalis (Mitchill)) averaging 2.3 gm were introduced into Westwick Lake on May 23, 1968. Frequencies of inversions in C. tentans occurring in Westwick Lake were determined before and after the introduction of the fish. Soronson Lake (a sister lake to Westwick Lake) was maintained without fi,sh and was used as a control situation for Westwick Lake, St a t i s t i c a l tests employed consisted of standard chisquare contingency tests, multiple correlation analysis and stepwise multiple regression analysis. The program used for multiple correlation and step-wise multiple regression analyses were obtained from the UBC Step Program (Computing Center, University of British Columbia). Results 1. Differences between lakes. A summary of the frequencies of the inversions studied, the lo c a l i t i e s from which the samples were taken, sample sizes and collection dates are given in Table XVI. These data also are grouped according to the geographic area in which the population occurs and the averages for these areas are given. Inversion frequencies in populations present in the different lakes certainly appear to be different and the probability that the samples were drawn from the same 2 populations 'is less than 0.01 (X^ = 37. 00). Therefore, i t is concluded that inversion frequencies differ significantly in different lakes. 71 Table XVI. Percentage frequency of inversions in chromosomes 1 and 4 of C. tentans in 12 lo c a l i t i e s i i central British Columbia. CHROMOSOME 1 CHROMOSOME 4 DATE OF SAMPLE SAMPLE LOCALITY COLLECTION Rad Rade SIZE 4c 4b c 4b 4 SIZE CHILCOTIN AREA NR OP BOX 4 8 SEPT. 1967 79.4 20. 6 364 72.8 23. 6 0. 0 3.6 . 364 BOX 89 8 OCT. 1967 • 81.4 18. 6 220 66.8 27. 7 1.8 3.7 220 NR. PHALEROPE MAY-NOV. 1967 77.3 22. 7 1,168 69. 7 24.1 1.8 4.4 1,168 NR OP CRESCENT 2 MAY 1967 74.3 25.7 214 67.8 22.9 5. 1 4. 2 214 OP BOX 4 7 JULY 1967 74. 7 25.3 186 65. 1 30. 1 1. 1 3. 7 186 RACETRACK 8 OCT. 1967 78. 6 21.4 220 73: 2 22:7 ?-: 3 1. 8 220 AVERAGE 77.6 22.4 2,372 69. 2 30. 2 2.0 3.6 2,372. SPRINGHOUSE AREA WESTWICK LAKE 3 May 1967 73.6 26.4 1,220 76.4 20. 7 0.8 2. 1 1,230 SORENSON LAKE 3 May 1967 74.3 25. 7 1,066 79. 5 18. 2 0.7 1.6 1,068 AVERAGE 74.0 26. 1 2,286 78.0 19. 5 0.8 1.9 2,298 GANG RANCH AREA LE-5 9 SEPT. 1967 65.5 34.5 220 67. 7 23. 6 6.4 2.3 220 LE-3 9 SEPT. 1967 76.6 23.4 214 57. 9 28.5 4.2 9.4 214 GR-3 9 SEPT. 1967 65.0 35.0 220 66.4 26.8 2.3 4.5 220 AVERAGE 69.0 31.0 654 64.0 26.3 4.3 5.4 654 CLINTON AREA STY MJTR. 9 OCT. 1967 72.8 27. 2 600 7 26, 7 10. 8 7. 8 600 TOTAL AVERAGE AND TOTAL SAMPLE SIZE 74.5 25.5 5,912 68.2 24.6 3.1 4.1 5,924 72 2. Horizontal variation in lakes: The results of an analysis of horizontal variation in frequencies of inversions of C. tentans occurring in Westwick Lake and Sorenson Lake are given in Table XVII. Sorenson Lake and Westwick Lake were selected for this study because they represented, at least superficially, homogeneous and heterogeneous environments, respectively. The margin of Sorenson Lake was characterized by the presence of the marsh reed, S_. acutus, and C. tentans larvae around the entire lake, and substrate of uniform composition and consistency., In contrast, the margin of Westwick Lake was characterized by discrete clumps, of S_. acutus, discontinuous occurrence of C0 tentans and a substrate varying from flocculant mud to sand and firmly packed clay. Thus, i t was a surprise to find that the probability that the frequencies of inversions in chromosome 1 at six lo c a l i t i e s in Sorenson Lake were comparable, was 2 less than 0.05 (X^ = 13.6), while inversion frequencies of chromosome 1 in Westwick Lake and inversion frequencies of chromosome 4 in both Sorenson Lake and Westwick Lake showed no significant difference. 3. Temporal variation. The results of analysis by season of inversion frequencies for chromosome 1 and 4 of C„ tentans collected from Near Phalerope are given in Table XVIII and Figure 18. The results of the study conducted in Near Phalerope lake are vir t u a l l y identical to those obtained by Acton in Six Mile lake during 1959-1960 (also included in Table XVIII for comparison). Thus i t appears to be a general characteristic of populations of C. tentans of populations in central British Columbia that inversions of chromosome 1 do not vary significantly in frequency, while inversions in chromosome 4 undergo significant seasonal variation in their frequencies. 73 Table XVII. Horizontal v a r i a t i o n . ' Percentage frequencies of inversions i n samples collectec from several s i t e s within i n d i v i d u a l lakes. Chi-square values and p r o b a b i l i t i e s that the differences within lakes can be a t t r i b u t e d to sampling error are given. The s i t e s sampled i n Westwick Lake are shown i n Figure 19. LOCALITY SITE CHROMOSOME 1 SAMPLE Had Rade SIZE SORENSON LAKE A 75.3 24.7 194 B 63.7 36.3 124 C 75.3 24.7 162 D 73.9 26.1 188 E 73.0 27.0 200 F 81.8 18.2 198 AVERAGE 79.5 25.5 X 2 = 13.609 0.02 P 0.01 WESTWICK LAKE A 75.3 24.7 146 B 78.0 22.0 200 C 75.0 25.0 200 E 76.6 23.4 124 F 70.7 29.3 164 G 70.5 29.5 200 H 65.0 35.0 186 AVERAGE 73.6 26.4 X? = 10.610 0. 2 P 0. 1 CHROMOSOME 4 SAMPLE 4c ... 4DC 4b . 4 SIZE 80.4 17. 0 1.0 1.6 194 83. 9 12. 9 0.8 2.4 124 75.3 21. 6 0. 6 2.5 162 80.8 17. 6 0.0 1.6 188 78. 5 20. 0 1. 0 0.5 200 79. 0 18. 5 1.0 1.5 200 79. 5 18. 2 0.7 1. 6 x 2 15 = 9. 133 0.9 P 0.8 76.0 19. 9 1.4 2. 7 146 73.5 23. 5 2.0 1. 0 200 74.5 21. 5 1.5 2.5 200 80. 6 16. 4 0.0 3.0 134 68.3 27. 4 0.0 4.3 164 82. 5 17. 0 0.0 0. 5 200 79. 6 18. 3 0.5 1. 6 186 76.4 20. 7 0.8- 2. 1 x 2 X12 = 17.754 0. 2 P 0. 1 74 T a b l e X V I I l . S e a s o n a l change i n t h e p e r c e n t a g e f r e q u e n c y o f i n v e r s i o n s i n chromosomes 1 and 4 i n p o p u l a t i o n s o c c u r r i n g i n S i x M i l e l a k e and i n N e a r • P h a l e r o p e l a k e . C h i - s q u a r e v a l u e s and p r o b a b i l i t i e s t h a t the d i f f e r e n c e s can be a t t r i b u t e d to s a m p l i n g e r r o r a r e g i v e n . D a t a f o r the. p o p u l a t i o n l i v i n g i n S i x M i l e l a k e were o b t a i n e d f r o m A c t o n ( u n p u b l i s h e d d a t a ) . CHROMOSOME 1 CHROMOSOME 4 SAMPLE SAMPLE LOCALITY DATE Rad Rade SIZE 4c 4bc . 4b 4 SIZE SIX MILE Aug-Sept 1959 77. 1 22. 9 712 48. 0 30.4 12. 9 8. 7 712 April i960 76. I 25. 9 712 45. I 34. 4 14. 6 5. 9 712 May 1960 74. 0 26. 0 588 47. 9 29. 7 13. 9 8.5 588 AVERAGE 75. 8 24. 2 47. 0 31.6 13. 8 7.6 9 X3 = 1. 927 0. 95 P 0.90 X 2 = 28. 6 936 P 0. 001 NEAR PHALEROPE • May 1967 63. 9 36. 1 36 61. 1 38.9 0. 0 0.0 36 June 1967 84. 0 16. 0 188 77. 7 . 16.5 2. 1 3.7 188 July 1967 79. 0 21. 0 200 70. 0 21. 5 2. 5 6. 0 200 Aug 1967 77. 3 22. 7 150 56. 7 36. 7 0. 0 6.6 150 Sept 1967 75. 3 24. 7 194 74. 7 18.6 0. 5 6.2 194 Oct 1967 76. 0 24. 0 200 67. 5 25.0 4. 0 3.5 200 Nov 1967 75. 0 25. 0 200 70.5 26.5 1. 5 1.5 200 AVERAGE 77. 3 22. 7 69. 7 24. 1 1. 8 4.4 x 2 -X6 _ 10 .123 0.25 P 0.1 X12 = 3 5 .810 P 0.001 75 Figure 18. Seasonal variation in frequencies of inversions of chromosomes 1 and 4. Percentage;; expressed along the oroinates have been converted by arc-sin transformation. The samples were collected from Hear Phalerope lake. qo D-8° c= cu o - 60 " S O 70 > c o B O D 7 0 y ( ) 0 % bO Z AO - 30 o O (D > cz O . I Rad • o - — - o - — o -o CHROMOSOME . 0 o-o-o o — o R a d e 1 1 1 i y il 1 May June July Aug Sept O c t Nov or *o. o 4 b c 'O ' p. 4 c O o o — - o ° CHROMOSOME 4 o o — 4 a 4 b May June July Aug Sept O c t N ov 76 Changes in inversion frequencies which might occur during longer periods of time are also of interest. Inversion frequencies in Westwick Lake and Six Mile lake were determined by Acton during 1959-1960 (Acton, 1962) and these same lakes were studied again in the present study. 'In addition, Westwick Lake and Sorenson Lake were sampled during 1967 and again in 1968. Thus change in periods of time as short as one year and a,3 long as eight years could be evaluated. The results of the analysis oc change in frequencies over longer periods of time are given in Table XIX. No significant change was observed in the frequencies of inversions in chromosome 1, but, in general, the inversions in chromosome 4 showed highly significant variation. Of course, the differences in frequencies of inversions of chromosome 4 might be due to seasonal variation since samples taken from Sorenson Lake during May 1967 were compared to samples taken during November 1968; however, samples collecte during the same season from Westwick Lake show the same significant variations., In addition, frequencies of the inversions of chromosome 4 in Six Mild lake showed the same pattern of difference whether the September :.959 data were compared to October 1968 data or whether the complete seasonal data from 1959-1960 were compared to October 1968. 4. Effect of change in the biotic environment. The effect of the introduction of fish into Westwick Lake on the inversions of chromosomes and 4 of C. tentans are given in Table XX together with the results of measurement: of inversion frequency in C. tentans occurring in Sorenson Lake (the control lake). The locations of the sample sites in Westwick Lake are shown in Figure 19. Qualitatively the pattern of change in inversion frequency in 77 Table XIX. Changes in percentage frequency of inversions in chromosomes 1 and 4 over long periods of time. Chi-square values and probabilities that changes can be attributed to sampling error are given. Data reported from Six Mile and Westwick Lake for 1959. were taken from Acton (1962). CHROMOSOME 1 CHROMOSOME 4 LOCALITY DATE Rad Rade SAMPLE SIZE 4c 4b c 4b SAMPLE SIZE SORENSON LAKE MAY ' 1967 NOV 1968 74.5 25.5 1,066 79.5 18.2 0.7 1.6 1,068 75.0 25.0 400 68.3 24.3 2.2 5.2 400 X 1 = 0.650 0.5 0.3 17.068 P 0.001 WESTWICK LAKE MAY 1967 MAY 1968 73.6 26.4 1,220 76.4 20.7 0.8 2.1 1,230 75.9 24.1 684 80.9 15.9 0.3 2.9 686 X t = 1.086 0.3 0. 2 X 3 = 4.036 0.7 0.5 SIX MILE SEPT., APRIL, MAY 1959 nPT 1Q£7 75.8 7 ^  9 24. 2 2,012 47. 0 A. 9 S 31.6 i n /. 13. 8 1 ^ 9 7.6 2,012 7 £ £nn WESTWICK LAKE MAY 1959 MAY 1967 X^ = 2. 252 0. 25 P 0. 1 X^ = 12.550 0.01 P 0.005 67.0 23.0 276 67.0 23.9 4.7 73.6 26.4 1,220 76.4 20.7 0.8 4.4 276 2.1 1,230 X^ = 4.857 0.05 P 0.02 X 3 = 30.868 P 0.001 78 Table XX. Chi-.nges with time in the percentage frequency of inversions in chromosomes 1 and 4 after the intro duction of eastern brook trout into Westwick Lake Sanples taken from Sorenson Lake during the same tine period are also considered ::or change in inversion frequency. Chi-square values and proba b i l i t i e s that these changes can he attributed to sampling error are given. CHROMOSOME 1 CHROMOSOME 4 SAMPLE SAMPLE SAMPLE DATE . SITE. Rad Rade SIZE,. 4c 4b c 4b 4 SIZE SORENSON LAKE (control s i t u a t i o n ) MAY 1967 A 75.3 24. 7 194 80. 4 17.0 1. 0 1.6 194 E 73.0 27. 0 200 78. 5 20.0 1. ,0 0. 5 200 AVERAGE 74. 1 25. 9 79. 4 18.5 1. 0 1.0 NOV 1968 A 75. 5 24. 5 200 66. ,5 26.0 2. 0 5.5. 200 E IL. s 25, 5 200 70. 0 22. 5 2. 5 5.0 200 AVERAGE 75.0 25. 0 68. 3 24.3 2. 2 5.2 X* = 0.650 0. 5 P 0. 3 X 2 = 17.068 P 0. 001 WESTWICK LAKE (experimental situ a t i o n ) MAY 1968 E . 75.0 25.0 344 80. 1 15. 9 0. ,6 3. 4 346 G 76. 7 23.3 340 81. 8 15. 9 0. 0 2. 3 340 AVERAGE 75.9 24. 1 80. 9 . 15. a 0. 3 2. 9 FISH INTRODUCED JULY 1968 G 81.4 18.6 188 75. 5 20. 2 0. 0 4. 3 188 OCT 1968 E 64. 5 35.5 200 72. 5 21. .5 1. 5 4. 5 200 G 71. 0 29.0 200 71. 5 21. 0 3: 0 4. 5 200 H 76.0 24.0 200 72. 5 . 20. 0 3. 5 4. 0 200 A T 7TT> T ) A ( - T O i i . v JLJ j . v n \ _ j iii n r\ c / \->. O r\ tr y . ~> 1 *\ / £. . C\ £. £.\J , rs O £. . 4. 3 NOV • 1968 E 70. 0 30.0 200 75. 5 22. 5 1. 5 0. 5 200 G 70. 0 30.0 200 76. 0 19. 5 1. 5 3. 0 200 AVERAGE 70. 0 30.0 75. 8 21. 0 1. 5 1. 7 X 2 = 12.737 0.01 P 0.001 x 2 -X18 " 33. 838 0. 02 P 0. 01 79 Figure 19. Localities sampled in Westwick L.ike in order to analyze horizontal variation and to evaluate the effect of introduction of fish. Sample transects used for the collection of data ^resented in Figure 13 are shown by the dotted lines. WESTWICK L SOUTH 0-5 KM. END 80 Westwick Lake was an i n i t i a l increase in the frequency of 1 Rad during July, followed by a decreas.e during October and November j (reciprocally an i n i t i a l decrease in 1 Rad, followed by an increase). The trend of change in frequency of inversions in chromosome 4 was for a decrease ia the frequency of inversion 4c, an increase in inversion 4bc, and increase in inversion 4 b , and an i n i t i a l increase in inversion 4 followed by a decrease. During the same period no change was observed in inversions in chromosome 1 in Sorenson Lake, although inversions in chromosome 4 varied significantly. The variation in inversions of chromosome 1 in Westwick Lc.ke was significant at the probability level of less than 0.01, while the change in the inversions of chromosome 4 was significant at the probability level of less than 0.02, but greater than 0.01. A summary of the probabilities of the differences observed during the four sample periods being due to change are given in Table XXI. The probabilities given in this table indicate that while the frequency of inversions in chromosome 1 during October differed significantly from that during July, i t did not differ from the i n i t i a l frequancy measured during May. However, by November the frequency was different from i t s i n i t i a l value. No significant, change was observed in the frequencies of inversions present in chromosome 4. The change in frequency of inversions of chromosome 1 in Westwick Lake was further analyzed by the method described by Wright and Dobzhansky (1946) in order to evaluate the significance of the change in frequency and to determine the relative adaptive values of the two inversions i f the change was significant. The time intervals at which cytogenetic data were collected from Westwick Lake corresponded roughly to one generation. The change in frequency of 1 Rad was highly significant (t = 23.5 with n = 5; 81 T a b l e XXI. Summary o f p r o b a b i l i t i e s t h a t d i f f e r e n c e s i n i n v e r s i o n f r e q u e n c i e s o b s e r v e d d u r i n g t h e sample p e r i o d s a r e a t t r i b u t a b l e t o s a m p l i n g e r r o r . CHROMOSOME 4 MAY 1968 0. 2 > P > 0 . 1 0. 0 2 > P > 0 . 01 0. 5 > P > 0 . 3 JULY 1968 0. 2 > P > 0 . 1 0 . 5 > P > 0 . 3 0. 9 > P > 0 . 8 OCT 1968 0. 1 > P > 0 . 05 0. 0 1 > P > 0 . 001 0. 1 > P > 0 . 05 NOV 1968 0. 0 1 > P > 0 001 0. 0 2 > P > 0 . 01 0. 9 5 > P > 0 . 9 SAMPLE SIZE 684 188 600 400 CHROMOSOME 1 82 P i s less than 0.001) and the regression r e l a t i o n s h i p between change i n in v e r s i o n frequency with respect to inversion frequency was s i g n i f i c a n t at the p r o b a b i l i t y l e v e l of P less than 0.05, but greater than 0.02 (t = 2.58 with n = 5). The r e l a t i v e adaptive values of 1 Rad and 1 Rade computed by least square s o l u t i o n were 0,66 and 1.49, r e s p e c t i v e l y . The average f i t n e s s of the e n t i r e population was 0.534. Together these values predicted a new equilibrium frequency of 1 Rad equal to 69.37o. This predicted equilibrium value was very nearly achieved by October. Discussion Before the r e l a t i o n s h i p between inversion frequency and environ-mental factors can be discussed, one muse consider how representative the genetic samples reported i n Table XVI are of the t o t a l p o p u l a t i o n s . l i v i n g within each of the lakes and how v a r i a b l e the inversion frequencies are within i n d i v i d u a l lakes. P r i o r to any s t a t i s t i c a l analysis of the r e s u l t s presented i n t h i s section, the p r o b a b i l i t y l e v e l of less than or equal to 0.01 was chosen as the l e v e l of si g n i f i c a n c e . This l e v e l of p r o b a b i l i t y may be unusually high for routine b i o l o g i c a l observations, but i t was so chosen to focus a t t e n t i o n on differences less l i k e l y to be at t r i b u t a b l e to sampling error. The accuracy with which a singl e sub-sample of the population represents the genetics of the e n t i r e population can be determined by analyzing the v e r t i c a l d i s t r i b u t i o n of larvae within lakes and by analyzing the h o r i z o n t a l v a r i a t i o n of inversion frequencies within lakes. The v e r t i c a l d i s t r i b u t i o n of C. tentans i n the lakes studied indicates that the larvae are most abundant at shallower depths and that the larvae 83 do not occur below two meters. These two facts, in turn, indicate that the samples taken for cytogenetic characterization of the populations in the different lakes (1) were taken from the areas of the lakes occupied by C. tentans (and more importantly that samples need not be taken from depths greater than 2 m) and (2) were ta'.-cen from areas of the lakes occupied by the greatest numbers of C. tantans. Thus, the samples were drawn from the largest portions of the populations and are at least representative of the majority of chironomids occurring in any particular lake. Furr.her, no significant horizontal variation in inversion frequency was found in either Sorenson or Westwick Lakes ( i . e_. , P less than 0.01). Therefore, a sample taken from anywhere around the margin of a lake should be representative of the entire population since the distribution of inversion frequencies within lakes is horizontally homogeneous. Seasonal variation in inversion frequencies of C. tentans has been studied in Britain (Acton, 1957) and in central British Columbia (Acton, unpublished data). In Britain, Acton studied temporal change in ten inversions and found slight variation in only one of the inversions (P less than 0.05, but greater than 0.02). He concluded that inversions frequencies of C. tentans did not vary seasonally, since variation in one out of ten cases might be due to chance. The study he performed in central British Columbia was done in Six Mile lake (a lake considered in the present study) and the sane inversions considered herein were studied. His conclusion was that the frequencies of inversions in chromosome 1 did not vary, but inversions in chromosome 4 did vary significantly. In view of the divergent results obtained from the populations studied in Britain and in Western Canada, and since Dobzhansky (1943) observed significant seasonal variations in the inversion frequencies of some populations of Drosophila pseudoobscura 84 but not in others, the possibility of seasonal variation was investigated once more using Near Phalerope lake. The results obtained in this study were vir t u a l l y identical to those obtained by Acton„ It was concluded with some certainty that inversions in chromosome 1 were stable, while those in chromosome 4 were concluded to undergo significant variation. The consequence of significant seasonal variation in the frequencies of inversions of chromosome 4 is that those frequencies of inversions of chromosome 4 reported in Table XVI cannot: be studied with respect to environmental factors, since the samples taken from the different lakes were not a l l collected during the same season. In light of this, only inversions in chromosome 1 w i l l be considered in the remainder of this discussion. The implication of the long term s t a b i l i t y of inversions in chromosome 1 is that the inversions are adapted to relatively stable environmental factors. Further, none of the frequencies, of inversions in chromosome 1 depart significantly from the values predicted by the Hardy-Weinberg principle. This result further emphasizes the sta b i l i t y of the inversions of chromosome 1. Clearly, the frequencies of the inversions are in equilibrium with their respective environments. Since; i t has been concluded that the sub-samples taken from the lakes are representative of total populations Living within any one lake and that the inversions of chromosome 1 are stable with respect to time the correla-tion between the differences in frequencies of inversions in chromosome 1 and the environments in which they exist can be considered. The lake environment data reported in section III and the ecological data reported in section IV both indicate that the primary character of the 85 l a k e s to which C. tentans must adapt , i s d i f f e r e n t i n c h e m i c a l c o n c e n t r a -t i o n s (whether o f t o t a l or of i n d i v i d u a l i o n s ) . F u r t h e r , the l a c k o f v a r i a t i o n i n i n v e r s i o n frequency w i t h i n lakes and the extreme s t a b i l i t y o f the i n v e r s i o n f r e q u e n c i e s i n the lake:) suggest t h a t g e n e t i c a d a p t a t i o n as r e f l e c t e d by i n v e r s i o n frequency p e r t a i n s to some environmenta l p r o p e r t y which i s r e l a t i v e l y u n i f o r m throughout i n d i v i d u a l l a k e s and which i s r e l a t i v e l y s t a b l e w i t h t i m e . Water c h e m i s t r y i s such a f a c t o r „ The r e l a t i o n s h i p s between the frequency of 1 Rad, c o n d u c t i v i t y , t o t a l d i s s o l v e d s o l i d s , sodium, p o t a s s i u m , c a l c i u m , magnesium, c a r b o n a t e , b icarbonate . , c h l o r i d e , and s u l f a t e are g i v e n i n Table X X I I . I t may be seen from t h i s t a b l e of p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s t h a t the frequency o f 1 Rad shows no s i g n i f i c a n t c o r r e l a t i o n w i t h any o f the aforementioned c h e m i c a l f a c t o r s . Hence, a l though occurrence and abundance appears tc be a f f e c t e d by these same c h e m i c a l p r o p e r t i e s , the i n v e r s i o n s do not i n d i c a t e g e n e t i c a d a p t a t i o n . However, i f c h e m i s t r y i s not i m p o r t a n t , then what i s ? I n order to s e l e c t o ther env ironmenta l f a c t o r s which might show a s i g n i f i c a n t : r e l a t i o n s h i p w i t h i n v e r s i o n f requency , s c a t t e r diagrams o f the r e l a t i o n s h i p between i n v e r s i o n frequency and those f a c t o r s were c o n s t r u c t e d and these diagrams are shown i n F i g u r e 20. From these d iagrams, c o n d u c t i v i t y , t o t a l d i s s o l v e d s o l i d s , p H , minimum d i s s o l v e d oxygen, numbers of A n a t c p y n i a  d y a r i , and t o t a l numbers o f a l l other chlronomids were s e l e c t e d f o r a n a l y s i s w i t h the frequency of 1 Rad. The c o r r e l a t i o n between frequency of Rad and the e n v i r o n m e n t a l f a c t o r s i s shown i n Table X X I I I . I n t h i s t a b l e , i t i s c l e a r t h a t 1 Rad shows a s i g n i f i c a n t r e l a t i o n s h i p w i t h the t o t a l numbers of o ther c h i r o n o m i d s . S i n c e at l e a s t one s i g n i f i c a n t c o r r e l a t i o n was found between i n v e r s i o n frequency and environmenta l f a c t o r s , the same data were ana lyzed by s t e p - w i s e m u l t i p l e r e g r e s s i o n i n order to q u a n t i f y the Table XXII. Summary of the p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s which describe the r e l a t i o n s h i p between the frequency of 1 Rad and physico-chemical water properties. With 12 degrees of freedom,r = 0.661 i s s i g n i f i c a n t at the l e v e l of P = 0.01 and r = 0.78 i s s i g n i f i c a n t at the l e v e l of P 0.001. 1 Rad CONDUC TDR Na K Ca Mg COo HC0 3 01 1 Rad CONDUCTIVITY -0.285 TDS -0.238 0. 990 Na -0.315 0. 876 0, .830 K -0.248 0. 808 0. . 742 0. 967 Ca -0.143 0. 143. 0. . 207 0. 266 -0. 391 Mg -0.186 0. 696 0. , 746 .0. 273 0. 175 0. 676 c o 3 • 0.021 0. 687 0. .657 0. 820 0. 779 -0. 293 0. 197 HC0 3 -0.461 0. 704 0. .615 0. 913 0. 937 -0. 332 0. 056 0. 662 Cl -0.232 0. 874 0. 840 0. 960 0. 936 -0. 250 0. 330 0.812 0. 854 s o 4 0.068 0. 495 0. . 581 0. 033 0. 076 0. 716 0. 922 0. 031 0. 240 0.096 87 F i g u r e 20. S c a t t e r diagrams d e p i c t i n g the r e l a t i o n s h i p between the frequency of the i n v e r s i o n 1 Rad and the e n v i r o n -mental f a c t o r s i n d i c a t e d . I n t h i s f i g u r e , the dark c i r c l e s i n d i c a t e e s t i m a t e s d e r i v e d from samples c o l l e c t e d from d i f f e r e n t l a k e s , w h i l e the open c i r c l e s i n d i c a t e r e l a t i o n s h i p s between samples c o l l e c t e d from a s i n g l e l a k e . The v a r i a n c e a s s o c i a t e d w.'.th the open c i r c l e s may be assumed to correspond to the v a r i a n c e which would be present i n any of the o t h e r l a k e s . 8 0 75 7 0 65 8 0 75 7 0 65 8 0 75 7 0 65 8 0 75 7 0 65 8 0 75 7 0 65 8 0 75 7 0 65 8 0 -75 • 7 0 -65 • 8 0 ; 75 " 7 0 -65 -Temperature C 0 o o -i 1 1 1 1 1— 17 18 19 2 0 21 22 © ' ' ' l i 1 3 £-5 9 -0 9 - 0 9 tneq /l Co "i—i 1—i 1 — i — i — r 1 2 3 4 5 6 7 8 —i 1 1— r— IO 2 0 3 0 4 0 e o I 1 1 1 1 IO 2 0 3 0 4 0 5 0 ° " 0 4 1 7 -% Substrate Composition Q 5 8 g m m • o o ~i—i—i—i—i—i—i—i—i—i—r I 2 3 4 5 6 7 8 9 IO It Conductivity umhcs. o o Total Dissolved Solids mq/l l 1 l 1 r~ O IOOO 2 0 0 0 3 C O O 4 0 0 0 O 1 —i 1 — -IOOO 2 0 0 0 3 0 0 0 . q/l Na O IO ~ 1 — 20 - " l — :so meq/'l K 4 0 O tq/l Mq O IO meq/l H C O 3 ; meq/'l 20 C! 30 meq/l C O 3 • ' 1 1 I 1 1 1 1 1 O 5 IO 15 I 2 3 n — T ~ 5 6 • " o meq/l SO4 % Substrate Compsition >l98mm ; o o 7 to bubstrate Composition , -„ 1 1 1 1— O 10 20 3 0 0/ c u . r lo substrate Composition „ m UflJZm l—1—T—1—1—1—r—1—1—1—r~ O I 2 3 4 5 6 7 3 9 IO e o o o 0 0 0-I47-/ 0 Substrate Compositio 1 Q ^\(=)TN j—1 . . . 1 IO 15 % Composition of • „ C. tentans —1 1 1 1 r~ IO 2 0 3 0 4 0 5 0 % Composition of 0 0 G. borbipes — 1 1 1 1 1 1 1 <~ ) 2 0 4 0 6 0 8 0 — 1 1 1 O 10 20 3 0 s % Composition of o other .Chironomus O o °/0 Substrate Composition < O I 4 7 mm 1 1 1 1——I r ~ — 1 — 30 4 0 5 0 6 0 7 0 8 0 9 0 e % Composition of A. dyari "I O IO % Composition of other ^ chironomids e 1 1 I I I 1 I 1 1 1 • O 2 0 4 0 6 0 8 0 , o 0 o % Orqanic C / 0 C. tentans -1—1—1—1—1—1—r~ 1 1—1—r~ O 20 4 0 60 ;30 l O O 1 • • • • 11 1 • • 1 • • • • 1 • • 1 1 1 1 • • • 1— O 5 IO 15 2 0 25 Table X X I I I . Comparison of p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s f o r the r e l a t i o n s h i p between the frequency of Rad and c e r t a i n environmental f a c t o r s . With eight degrees of freedom, r = 0.765 i s s i g n i f i c a n t at P = 0.01 and r = 0.872 i s s i g n i f i c a n t at P = 0.001. . 1 Rad CONDUC TDS pH MIN 0 2 ANAT0 OTHER 1 Rad CONDUCTIVITY 0. 349 TDS 0 274 .0. 991 pH 0 062 0. 564 0.616 MINIMUM DISSOLVED OXYGEN 0 342 0.407 0.481 0 605 # OF ANATOPYNIA -0 666 0, 809 0. 743 0 320 0.184 .. # OF OTHER CHIRONOMIDS 0 874 0.584 0. 517 0. 306 0.051 0.816 89 dependency of 1 Rad on the environmental factors. The results of this analysis are summarized in Table XXIV. When 0„01 is used as the probability level for selection of independent variables to be included in the regression analysis, i t is found the only environmental variable contribut-ing significantly to the frequency of 1 Rad is the total number of other chironomids. However, with the number of other chironomids being the only included variable, only about 767» of the difference of 1 Rad is explained. When 0.05 is used as the probability level for selection cf independent.: variables to be included in the regression analysis, the number of other chironomids and pH are found to contribute to the frequency of 1 Rad. When the two variables are included, 887o of the difference of 1 Rad is accounted for. However, the effect of the number of other chironomids is greater than that of pH. At this juncture, i t is of l i t t l e importance whether the probability level used for retention of independent variables :ls 0.05 or 0.01. The main point is that a biotic effect ( i . e. , the number of other chironomids) appears to be the primary factor upon which Rad is dependent, although a lesser dependency upon pH was observed. The inportance of a biotic factor as opposed to a chemical factor came as a complete surprise in view of a l l the environmental and ecological data. The foregoing discussion suggests that inversion frequency may be related to the numbers of other chironomids and to a lesser degree the pH of the lake water; however, the correlative studies just discussed do not allow any firm conclusions to be made about the effect of the environmental factors. Therefore, some experimental analysis of the effect of environ-mental factors is required. Acton (1955) conducted some simple laboratory experiments in which Table XXIV. Summary of step-wise multiple regression analysis. Rad is the dependent variable and conductivity, TDS, pH, minimum concentration of dissolved oxygen, number of Anatopynia,, and number of other chironomids are the independent variables considered. Using P = 0.05 as the probability level for selecting significant independent variables. RSQ = 0.8829 F probability = 0.002 Standard error of Y = 1.818 — Variables Coefficient of b Standard Error F- ratio F probability Const. 4.1892 25.4104 pH 7.0657 2.8509 6.1423 0.0470 Other 0.1608 0.0240 45.0430 0.0007 o Using P = 0.01 as the probability level for selecting significant independent variables. RSQ =0.763 F probability = 0.0023 Standard error of Y = 2.3945 Variables Coefficient of b Standard Error F-ratio F probability Const. 67.0539 1.9905 Other 0.1426 0.0300 22.5407 0.0023 91 he demonstrated that certain karyotypes of C. tentans in Britain are associated with differential survival of larvae maintained under "crowded" and "uncrowded" conditions. The implication of the experiment was that: the inversions contribute some selective value to the species. However, he cautioned that these laboratory tests did not unequivocally prove that the inversions have a selective significance in nature. Thus, not only is an experimental analysis required to conclude that an environmental factor is related to genotypic constitution, but the experimental analysis should be performed in natural conditions i f the environmental factor is to be concluded to have significance in nature. The problem of demonstrating that any genetic character has selective significance in nature has always been the stumbling block to the simple, clear, and direct demonstration of natural selection. The regression analysis indicated that a biotic effect (the number of other chironomids) showed the strongest relationship to inversion frequency and thereby suggested that a biotic effect might be the most promising for experimental analysis. The transportation of large enough populations of C. tentans or "other chircnomids" to create the "crowded" conditions of Acton (1955) would be a large undertaking which in turn would require continuous monitoring to demonstrate that the transferred populations did not simply emigrate elsewhere. However, one means of altering the environment which did seem reasonable was to introduce a predator. Predators are supposed to be a potent selective force (_e..g. , Slobodkin, 1961; Brooks, 1965; Brooks and Dodson, 1965) and they could be introduced in sufficient numbers to "guarantee" some effect, but the same time the numbers of predators could be regulated well enough so that not a l l the prey are k i l l e d . Consequently, predation was chosen as the "biotic effect" 92 to be analyzed experimentally. Of the potential predators of C. tentans, f i s h , in general, and the eastern brook trout, in particular, were chosen as the potential predator since they are known to eat chironomids, since their emigration from the lake could be controlled, and since they could be obtained in large numbers. Westwick Like was chosen as the location for the experiment since the lake had been sampled extensively for both genetic and ecological characterization and Sorenson Lake was studied as a control since i t was similar to Westwick Lake with respect to morphometry, water chemistry, and faunal composition, and since the enetics of the population living in the lake was known equally well. Further, since the two lakes occur very close together in the same basin they are subjected to the same climatic conditions. Originally, the effect of fish on the frequency of inversion 1 Rad was studied in order to demonstrate that the inversions considered herein did have selective significance in nature. Fortuitously, the use of fish also represented the use of a "biotic factor" and the results obtained represent a unique observation of natural selection under natural experi-mental conditions. The resultant data unequivocally demonstrate that the inversions have selective significance in nature. Furthermore, the selective value;; of the inversions were found to be rather large (i..e. , Rad = + 0.34 and Rade = - 0.49). The accuracy of these estimates are dependent on the accuracy with which the frequencies of inversions in the populations can be determined. 93 That the inversion frequency was shifted, even though a l l of the evidence indicates that the.frequencies c-f the inversions are unusually stable, is emphasized. And, this change can be interpreted as a direct demonstration of natural selection. However, the exact effect of the fish on the inversion frequency is a matter of speculation. The fact that fish are known to eat chironomids and the fact that the greatest change in inversion frequency was observed between May and October when the chironomids would be moving around out of their tubes actively feeding or emerging, implies that predation might be the effect. However, analysis of fish stonach samples (Scudder, personal communication) did not indicate that the brook trout were eating C„ tentans. In fact, the number of head capsules recovered indicated that chironomids represent only a small part of the food eaten by the fish. This result is not especially surprising since even in areas of their greatest abundance, C. tentans larvae represent only about 37» of the total chironomid fauna. It must be concluded thai: there is no evidence that predation occurred. However, whether predation has or has not occurred is of l i t t l e significance and the point to be . emphasized is that the frequency of an inversion was altered from i t s equilibrium value and that this change was associated with the introduction of f i s h , while no change was observed in frequencies of inversions in Sorenson Lake during the same period of time. Since the only primary difference between Westwick and Sorenson Lake was the introduction of f i s h , i t can be concluded that fish are responsible for the change whether they are directly (via predation) or indirectly (via a secondary change; in the lake environment) involved. In summary, analysis of correlation, of inversion frequencies and environmental factors suggested that biotic factors are more important 94 than lake water chemistry as determinants of inversion frequency. Further, analysis of the effect of an experimental b i o t i c change on inversion frequency v e r i f i e d that inversion frequency i s indeed dependent on b i o t i c factors. However, the differences i n 1 Rid which cannot be accounted for by the variables considered as shown by regression analysis (see p. 90 ) indicates that s t i l l other environmental factors are involved. The analysis of the effect of b i o t i c factors on inversion genotypes represents a unique demonstration of natural selection i n nature. VI. 95 Discussion Although the importance of-natural selection has been appreciated for over a hundred years and although natural selection is now commonly believed to be responsible for a l l physiological, ecological, behavioral, etc. changes observed in aninals, surprisingly l i t t l e is known about natural selection i t s e l f . At the outlet of this study one of the important questions which was posed was i f the: inversions of chromosome 1 have selective significance in nature. The correlation of the numbers of other chironomids with the inversions 1 Rad and 1 Rade indicated that such a biotic effect might be of selective significance and the f i e l d experiment involving the change in the frequencies of 1 Rad and 1 Rade by the introduction of fish confirmed that a biotic factor could affect the genetics of the animal. The relative importance of certain environmental factors as agents of natural selection is another matter which was considered in this study. The relationship between inversion frequencies and (1) physical, (2) chemical, and (3) biotic properties of the lakes was considered and the latter was the only one showing a significant relationship with the genetics of C. tentans. This result is rather surprising since a relatively sedentary animal such as the larval stage of C. tentans might be expected to be closely adapted both ecologically and genetically to i t s immediate physical and chemical environment. The changes in the frequencies of the inversions 1 Rad and 1 Rade which were associated with the presence of fish allowed measurement of the relative adaptive values of the karyotypes Rad/Rad, Rad/Rade, and Rade/Rade with respect to the physical, chemical,and biotic conditions in Westwick Lake. The relative adaptive values of the karyotypes were 0.66 : 1 : 1.49, respectively. 96 Not only do these relative adaptive values indicate the magnitude of the selective values acting on the inversions, but they also make possible some theoretical enquiry into the fitnesses of the inversions to other environmental factors and situations. If i t may be assumed that the inver-sion L rad has the same genetic composition throughout the study area and that the same is true of 1 Rade, then the differences in the frequencies observed in the different populations residing in different lakes (Table XVI) should reflect the fitness of the two inversions to the complex environmental conditions present in the different lakes. With the relative fitness of the inversions in Westwick Lake serving as the reference point, the relative fitnesses of the same inversions in the other lakes can be calculated in the manner described by Svirezhev and Timofeeff-Ressovsky (1968). These theoretical adaptive values are given in Table XXV. The magnitude of the adaptive values for the three karyotypes indicate that their selective values would be large. Ford (1964) has argued that selective values might be rather large and evolution might be quick, while American evolutionists h&ve more often emphasized that evolution is a slow and gradual process requiring only small selective values. Alternatively, the relative adaptive values of 1 Rad and 1 Rade might also be determined with respect to some other environmental factor, , food. If the greatest density of numbers of 0. tentans associated with a unit of food is assumed to represent the maximum fitness (i.e., W = 1.0), then the numbers of larvae present per unit o>: food in the other lakes may be used to estimate the average fitnesses of populations present in the different lakes. Further, since p and q are known for each of the C. tentans populations occurring in the lakes and since W is estimated from abundance and food ava i l a b i l i t y data, the relative adaptive values of the 97 Table XXV. Re-.lati.ve adaptive values of the karyotypes of 1 Rad and 1 Rade. The relative adaptive values of the karyotypes present in the different lakes were calculated using the estimates obtained from Westwick Lake (after fish were stocked) as the reference value. This table compares the relative adaptive values with respect to the total environmental conditions present in each of the lakes. RELATIVE ADAPTIVE VALUES LAKES Rad/Rad Rad/Rade Rade/Rade WESTWICK LAKE (reference) WESTWICK LAKE (before fish) LE-5 LE-3 NR OPPOSITE BOX 4 SIX MILE GR-3 BOX 89 NR PHALEROI'E SORENSON UKE NR OPPOSITE CRESCENT OPPOSITE BOX 4 RACETRACK 0.65 0. 62 0.70 0. 63 0.53 0.63 0.70 0. 55 0. 59 0. 62 0. 62 0. 61 0.53 1.0 1.0 1. 0 1. 0 1.0 1.0 1.0 1.0 1. 0 1.0 1. 0 1. 0 1. 0 1.49 1. 73 1.33 1. 96 2. 22 1. 68 1.31 2.46 2. 02 1. 78 1. 78 1.81 2.14 98 karyotypes can te calculated using equations given by Wright and Dobzhansky (1946). The relative adaptive fitnesses of the karyotj'pes and the average fitnesses of the different populations relative to food are given in Table XXVI. • Although these two examples of theoretical consideration of the fitnesses of the: inversions 1 Rad and 1 Rade may have validity, no pretense is made that the. values calculated are correct. The correct relative fitnesses could only be established by experimentation using the appropriate environmental conditions. Rather, the point in generating these fitnesses was to learn i f the same inversions would ha^e rather different relative fitnesses when different environmental factor:; were considered or i f in fact the estimated fitnesses would be about the same. It may be seen that the two calculations yield different results. Finally, evaluation of the fitnesses of the. same inversions to different environmental conditions should be especially interesting in that i t represents the most direct method for quantitative comparison of the evolutionary significance of the different environmental conditions. Since, as stated in the introduction of this thesis, the problem of ecological adaptation has always occupied a central position in evolutionary theory (see Allee, Emerson, Park, Park, and Schmidt, 1950), then an analysi.3 of the ecology of an animal with respect to i t s genetics should be interesting. Ecological data collected for (L tentans indicated that both the occurrence and abundance of C. tentans in central British Columbia are related primarily to chemical composition and concentrations of lake waters. Further, the occurrence of C. tentans within individual lakes is related to co-occurrence with other species of chironomids, increasing concentrations of chemical in the lake waters and minimum dissolved oxygen concentrations. However, 99 Table XXVI. Relative adaptive values of the karyotypes of 1 Rad and 1 Rade. The relative adaptive values were calculated by using the assumption that the numbers of C. tentans per unit of food are a measure of average population fitness. The population with tho maximum numbers per unit of food was set equal to the theoretical average fitness of 1.0. AVERAGE POPULATION FITNESS RELATIVE ADAPTIVE VALUES LAKES W Rad/Rad Rad/Rade Rade/Rade NR OPPOSITE BOX 4 LE-5 BOX 89 WESTWICK LAKE NR OPPOSITE CRESCENT .RACETRACK 1.000 0. 119 0. 697 0.309 •0.087 0.353. 1. 00 •0. 35 0. 58 0.31 •0. 23 0. 22 1. 0 1.0 1. 0 1. 0 1. 0 1.0 1.00 -1.57 -0.82 -1. 64 -2. 56 -1.86 100 cytogenetic date collected indicated that the inversions in chromosome 1 are related primarily only to the numbers of other chironomids and to a lesser extent, the pH of lake waters. Thus, within the limits of the a b i l i t y to exactly characterize the inversion frequencies of any particular population, the ecological measures of adaptation do not agree with the genetic measures of adaptation as reflected by inversion frequencies. Since inversions: reflect only macro-genetic change, i t remains possible that genetic adaptation might be present at the level of gene l o c i which would account fcr a l l the different aspects of ecological adaptation which were observed ir. this study. However, i f John and Lewis (1966) are correct in their assertion that chromosomal variations rather than gene variations are responsible for the key differences in populations, then one might expect a closer accordance between genetic adaptation, as reflected by inversion frequencies, and ecological adaptation to major environmental differences. The inversions present in chromosome 4, which could not be dealt with in this study owing to their seasonal variation in frequencies, might be related, to some of the ecological adaptations shown by C. tentans; however, they do not appear to be related to the more stable environmental factors such as water chemistry. The implication of the lack of agreement between ecological and genetic adaptation is that not a l l ecological changes or differences are of evolu-tionary significance. That is to say that either not a l l of the ecological characteristics of an animal are inherited or that considerable p l a s t i c i t y is present in tie genetic control of ecological response. This result is surprising in view of the current belief that any ecological difference, no matter how minute, is of evolutionary significance. It cannot be denied that any ecological difference may be evolutionarily significant, but that is very far from indicating that a l l differences must be evolutionarily significant. 101 LITERATURE CITED Acton, A.B. 1955. Larval groups in' the subgenus Chironomus Meigen. Arch, f. Hydrobiol., 50: 64-75. Acton, A.B. 1956. The identification an! distribution of the larvae cf some species of Chironomus (Diptara). Proc. Roy. Ent. Soc. A., 31: 161-164. Acton, A.B. 1957. Cromosome inversions in natural population of Chironomus tentans. J. Genetics, 55: 61-94. Acton, A.B. 1959. A study of differences between widely separated populations of Chironomus (=Tendipes) tentans (Diptera). Proc. Roy. Soc. London, 151: 277-296. Acton, A.B. 1962. Incipient taxonomic divergence in Chironomus (Diptera)? Evolution, 16 (3): 330-337. Acton, A.B. and G.G.E. Scudder. 1969. The zoogeography and races of Chironomus (=Tendipes) tentans Fab. (In press). Allee, W.C., A.E. Emerson, 0. Park, T. Park, and K.P. Schmidt. 1950. Principles of Animal Ecology. W.B. Saunders Co., Philadelphia, x i i + 837 pp. American Public Health Association. 1965. Standard Methods for the  Examination of Water and Wastewater. 12th Ed. New York, xxxi _ 769 pp. Beermann, W„ 1955. Cytologische Analyse eines Camptochironomus-Aribastards. I. Kreuzungsergebnisse und die Evolution des Karyotypus. Chromosoma, 7: 198-259. Brooks, J.L„ 1965. Predation and relative helmet size in cyclomorphie Daphnia. Proc. Nat. Acad. Sci. , 53: 119-126. Brooks, J.L,, and S.I. Dodson. 1965. Predation, body size and composition of plankton, Science, 150: 28-35. Campbell, R„B. and H.W. Tipper. 1966. Bonaparte River, British Columbia. Geol. Surv. Can., Map 3. Cockfield, W.E. 1948. Geology and mineral deposits of Nicola map-area, British Columbia. Geol. Surv. Can., Mem. 249. Cole, G.A. 1968. Desert limnology. In: Desert Biology. G.W. Brown (Ed.)-Academic Press, New York. I: 423-486. Cole, L.H. 1926. Sodium sulphate of western Canada. Canada Dept. of Mines. Mines Branch No. 646. v i i + 160 pp. Cummings, J.M. 1940. Saline and hydromagnesite deposits in British Columbia„ British Columbia Dept. of Mines. kBull. 4. 160 pp. 102 Dobzhansky,. Th. 1943. Genetics of n a t u r a l p o p u l a t i o n s . IX. Temporal changes i n the composition of populations of Drosophila p;;eudoobscura 0 Genetics, 28: 152-186. Engelmann, W. and D.G. Sha p p i r i o . 1965. Photo p e r i o d i c c o n t r o l of the maintenance and t e r m i n a t i o n of Larva l diapause i n Chironomus tentans. Nature. 107: 548-549. F i t t k a u , E, J . 1967. Chironomidae. In: Limnofauna Europa. J . l i l i e s (Ed.). Gustav F i s c h e r Verlag. S t u t t g a r t , p. 346-381. Ford, E.B. 1964. E c o l o g i c a l Genetics. Methuen & Co. L t d . , London, xv + 335 pp. Ford, E.B. 1965. Genetic Polymorphism. The M.I.T. P r e s s , Cambridge, Massachusetts. 101 pp. Hansen, H.P. 1947. P o s t g l a c i a l f o r e s t succession, c l i m a t e , and chronology i n the P a c i f i c Northwest. Trans. Amer. P h i l . Soc. New S e r i e s . 37(1): 1-130. Hesse, R., W.C. A l l e e and K.P. Schmidt. 1951. E c o l o g i c a l Animal Geography. John Wiley & Sons, Inc. New York. x i i i + 715 pp. Ho l l a n d , S,S. 1964. Landforms of B r i t i s h Columbia. A physiographic o u t l i n e . B.C. Dept..of Mines and P e t r o l . Res. k B u l l . No. 48. 138 pp. John, B. and K.R. Lewis. 1966. Chromosome v a r i a b i l i t y and geographic d i s t r i b u t i o n i n i n s e c t s . Science. 152: 711-721. Johannsen, O.A. 1937. Aquatic D i p t e r a . IV. Chironomidae: subfamily Chironominae. C o r n e l l . U. A g r i . Exp. S t a t i o n . Mem. 210. 52 p.p. Mackereth, F.J.H. 1955. Ion-exchange procedures f o r the estimates of 0) T o t a l i o n i c c o n c e n t r a t i o n , ( I I ) C h l o r i d e s , and ( I I I ) Sulphates i n . n a t u r a l waters. M i t t . d. I n t e r n a t . V e r e i n i g . f..Limnologie. No. 4, 16 pp. Mundie, J.H. 1957. The ecology of Chironomidae i n storage r e s e r v o i r s . Trans. Roy. Entomol. Soc. 109: 149-232. Munro, J.A. 1945. The b i r d s of the Cariboo parklands, B r i t i s h Columbia. Can. J . Res. (D). 23: 17-103. Munro, J.A. and I. McT. Cowan. 1947. A review of the b i r d fauna of B r i t i s h . C o l u m b i a . B r i t i s h Columbia Prov. Mus. S p e c i a l Publ. No. 2. 285 pp. Neumann, D. 1961. Osmotische Resistenz and Osmoregulation aquatischer Chironomidenlarven. B i o l . Z e n t r a l b l . 80: 693-715. Palmen, E. and L. Aho. 1966. Studies on the ecology and phenology of the Chironomidae (Dipt.) of the Northern B a l t i c . 2. Ann. Zool. Fenn. 3: 217-244. 103 Rawson, D„S„ and J.E. Moore. 1944. The saline lakes of Saskatchewan. Can. J. Res. (D). 22: 141-201. Richards, L, A„, Ed. 1954. Diagnosis and improvement of saline and alkal soils. U.S. Dept. Agri. Handbook No. 60. v i i + 160 pp. Roback, S.S, 1957. The immature Tendipedids of the Philadelphia area, Monogr. Acad. Nat. Sci. Philadelphia. No. 9„ 152 pp + 28 plates. Sadler, W. 0.. 1935. Biology of the midge Chironomus tentans Fabricius,, and methods for i t s propagation. Cornell U. Agri. Exp. Station. Mein. 173 . 25 pp. Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences., McGraw-Hill Book Co. Inc. New Ycrk. x v i i + 312 pp. Slobodkin, L.B. 1961„ Growth and Regulation of Animal Populations. Biology Studies. Holt, Reinhart and Winston. New York. 184 pp. Svireshev, Yu. M. and N.W. Timofeeff-Ressovsky. 1968. Some types of polymorphism. In: Haldane & Modern Biology. K. R. Dronamraju. Ed. Johns Hopkins Press, pp. 141-164. Swanson, CP. 1957. Cytology and Cytogenetics. Prentice Hall, Inc. Englewood C l i f f s . , N.J. x + 596 pp. Tipper, H.W. 1959. Quesnel, British Columbia. Geol. Surv. Can., Map 12. Tipper, H.W. 1963. Taseko Lakes, British Columbia, Geol. Surv. Can., Map 29. Townes, H.K. 1945. The Neartic species of Tendipedini. Amer. Midi. Nat. 34: 1-206. Townes, H.K. 1952. Tribe Tendipedini (=Chironomini) in: Guide to the insects of Connecticut. State.Ceolo and Nat. Hist. Surv. Conn. Bull. 80. Welch, P.S. 1952. Limnology. McGraw-Hill Book Co., Inc., New York, x i + 538 pp. Wright, S. and T. Dobzhansky. 1946. Ger.etics of natural populations. XII. Experimental reproduction of some of the changes caused by natural selection in certain populations of Drosophila  pseudoobscura. Genetics. 31: 125-156. Appendix I. .5athymetric maps of the lakes. Maps of 28 of the 32 lakes studied are given. Trees present in Racetrack .ind Box 89 lakes are indicated by 105 LB i 106 108 110 LE I c> LE 2 I l l PHALEROPE 112 BOX-26-2 114 LE 5 OOI KM l _ — 1 116 RUSH L 117 NEAR OPPOSITE BOX 4 0-05 KM. L _ , I 118 LE 3 119 LE 4 120 BOX 89 121 ROCK 122 NEAR PHALEROP Oo KM. O I KM. i : _ J 125 NEAR OPPOSITE CRESCENT O Q 5 KM. 126 BOX I 127 OPPOSITE BOX 4 128 RACETRACK 129 SP 6 130 BOX Appendix II. Chemical data collected from the lakes. A l l conductivities listed in this appendix are expressed as micromhos/cm at 25° C. MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS H2S (m) (°C) (jumhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg COc HCO. Cl SO, 141.351 55,932 147,100 8.10 CLINTON 22 May 1966 375.0 16.21 2.28 1.352.0 5 August 1966 327.5 18.80 1.16 1,630.0 0.0 52.7 31.25 2,043.0 24.27 1.543.0 0 15.6 30,100 27,626 10.05 1 25.9 34,900 33,556 10.20 GR-2 20 May 1966 525.0 11.22 550.0 13.08 A. A11 o-i i o ^  IQfifi 0.54 2.42 379.0 57.1 116.23 0.0 0.72 2.45 473.6 54.3 141.13 0.0 0 21.1 40,000 39,207 10.20 1 20.6 40,508 42,128 10.20 Ice 39,930 42,100 9.90 1 -2.0 57,320 69,248 9.70 0. 0 0.0 705.0 725.0 13. 90 13.80 16 February 1967 585.0 10.50 1,312.0 25.00 0. 14 0. 14 0. 29 0.29 3.90 560.6 72.2 166.06 3.90 573.6 68.0 164.83 3.13 718.4 274.6 126.50 9.41 978.0 162.0 296.00 0. 0 0.0 0.0 0.0 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 2 H2S (m) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg CO3 HC0 3 Cl SO^ LB-2 0 15.6 13,400 . 11,912 9. 50 1 14.4 16,600 15,249 9. 50 2 11. 7 17,900 17,154 9. 45 0 22.8 17,119 15,256 9. 20 1 22. 2 17,034 15,292 9. 20 2 21.1 17,034 15,412 9. 25 0 14.7 11,700. 16,714 8.80 1 14. 7 13,780 19,135 8. 70 2 17.0 14,420 24,174 8. 65 3 17. 0 20,590 37,016 8. 40 4 18.6 34,150 66,592 7. 60 5 16.4 39,800 87,828 7. 15 23 May 1966 5.05 — - 200.0 8.00 0.76 3.91 ___ 230.0 10.50 0.76 4.35 - — 223.0 10.41 0.40 6 August 1966 3.84 0.0 228.8 9.50 1 - 0 0 3.72 0.0 221.8 8.90 0.44 2. 13 0.0 225.0 8.70 0.44 LB-1 23 May 1966 5.70 --- 62.5 6.79 2.12 6. 14 --- 65.0 7.62 2.04 4.13 --- 67.5 8.50 1.98 3.15 --- 107.0 11.90 2.84 0.0 --- 193.0 19.10 11.40 0.0 --- 253.0 24.50 14.10 4.30 57. 1 51. 1 13. 97 59. 78 5.25 66.0 71.5 17. 56 75.04 6.02 84.4 67.3 19. 62 84. 06 5.40 66.0 60.6 15. 39 96.48 5. 70 64.3 61.6 15. 71 100.84 5.60 68.0 58. 9 15. 97 102.28 150.30 4.3 8.0 6. 52 196.86 174.70 4.3 8.3 7. 21 228.37 196.14 4.6 9. 2 8. 25 252.20 319.23 2.1 11. 7 11. 19 410.41 615.98 0. 0 27.3 19. 91 889.22 794.64 0. 0 69.6 7. 82* 1 ,040.03 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC . TDS 0 2 H2S (m) (°C) (umhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C 0 3 HCO3 Cl SO LB-1 6 August 1966 0 27.6 14,746 23,056 8. 70 5. 32 0.0 77.5 8.60 1. 18 • 194.90 4.0 9.8 8. 27 264.96 1 26.7 14,746 22,584 8. 70 "5. 32 0.0 76.3 9.10 1. 20 189.50 3.5 10.2 8. 24 239.00 2 25.6 14.746 22,716 8.60 5. 15 0.0 76.8 9.00 1. 34 190. 20 3.5 10.3 8. 58 274.90 3 24.4 14,915 22,760 8.60 8. 01 0.0 76.8 9. 10 1. 50 192.00 3.4 10.4 8. 10 284.40 4 25.3 37,119 73,876 7.80 5. 60 4.5 212.5 24.80 6. 24 662.06 0. 0 38.6 20. 48 983.70 5 25.3 35,932 72,656 7.80 5. 94 5.0 200. .0 24.50 6. 10 638.46 0.0 38. 5 16. 83 817.00 I-1 0 0 00 LONG LAKE 20 May 1966 0 • 14.4 9,110 7,792 9. 00 5.87 102. 5 7.30 0.66 10.94 21.4 35. 2 14.51 67. 12 1 . 14.4 9,260 8,006 9.05 6.04 102. 5 7.33 0.60 . 11.00 20.8 25.7 14.98 70.12 2 13.1 16,410 15,537 9.40 6.04 210. 0 12.35 0.62 15.33 43.6 37. 9 28.95 129.55 3 10.6 17,280 16,673 9.40 4. 28 220. 0 13. 20 0. 76 15.37 45.1 41.5 30.89 130.46 4 7.8 24,180 24,157 9.40 0. 0 409.0 17. 92 0. 66 10. 29 68.6 . 57.1 45.39 166.04 3 August 1966 0 22. 2 11,525 10,452 9. 20 5.60 0.0 140. 0 8.10 0. 10 13. 80 23.8 31.4 18.50 100.60 1 22. 0 11,525 10,100 9. 20 5.43 0.0 125. 0. 7.50 0.10 13.80 24.0 31.4 18. 77 76.28 2 20. 9 11,695 10,248 9.30 4.48 0.0 126.5 7. 70 0. 10 13.80 25.6 31.0 19.18 78. 72 3 19.4 19,492 18,440 9.35 0. 73 0.0 225. 5 13. 70 0.18 14. 70 48.5 42.4 .. 34.88 137.77 4 15. 3 25 ,729 25,656 9. 45 0. 0 5. 0 3i9.u ia. /u 0. lb i4. bO 7 0, b 54. 8 4». /b 196.14 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 o H 0S (m) (°C) (umhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C0 3 HC0 3 C l SO LONG LAKE 17 February 1967 T r> o ? , LQL 2 ;036 8. 90 ---- 24. 17 1. 00 0. 13 2. 77 4. 92 6.24 3. 20 20. 75 0 0. 0 12^033 11,660 8. 80 150. 00 8. 75 0.47 17. 85 26.00 49. 50 23.00 45. 96 1 0. 0 12,033 11,811 8. 90 150. 00 8. 65 0.60 18. 04 27.84 49. 06 22. 24 25. 10 2 0. 2 12,361 11,890 9. 00 - — - 149. 50 8. 75 0.44 16. 86 27.60 44. 90 22. 60 21. 46 3 0.4 12,690 12,070 9. 05 160. 00 8. 90 0.40 16. 40 29. 76 42.44 22. 23 20. 18 4 1.3 23,740 25,026 9. 25 342. 50 17. 99 0.40 14. 34 73.60 55. 20 45.60 46. 51 CO BOX 4 13 May 1966 0 11. 4 6,610 5,283 9. 40 • - 70. 00 7. 90 0.62 1.41 22. 10 36. 70 21. 90 10.33 1 12. 2 11,510 9,430 9. 40 • - 120.00 12. 79 0. 76 2.57 25.60 54. 60 36.40 20. 15 Q 12 082 Q i n • - 1 00 1 7 15 0. 76 3.36 52. 80 61. 70 45. 70 36.45 3 8. 3 14,900 12^893 9. 20 •- 180.00 18.05 0. 76 2.79 56. 00 65. 50 49.00 32.75 28 J u l y 1966 0 20. 0 11,017 8,599 9. 30 --- 0. 0 119.30 11. 90 0. 14 3.10 35.60 45. 10 32.78 " 21. 10 1 20. 0 11,017 8,872 9. 30 --- 0. 0 117.50 11. 90 0. 14 3.10 36. 00 56. 90 33.86 26. 60 2 14. 4 13,051 10,797 9. 30 --- 5. 0 139.80 14. 10 0. 10 3.60 45.60 53. 20 40. 26 30.00 3 8. 6 16,102 13,712 9. 40 --- 5. 0 173.50 17. 90 0.08 3.30 61.90 65. 80 50. 73 39.00 4 5. 6 17,966 15,441 9. 40 --- 5. 0 206.30 20. 5 0. 06 2.80 74.60 70. 90 55.15 , 46.00 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS H 2S (m) (°C) (pihos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg CO-. HCOc Cl SO/ 0 1 0 1 14.4 9,820 13.9 10,390 22. 0 22.0 11,864 11,864 13,842 8.20 13,961 8.15 17,636 8.60 17,148 8.60 BOWER'S LAKE 24 May 1966 7.93 --- 55.00 5.50 31.40 98.65 0.0 3.70 6.95 158.83 8.25 --- 52.50 5.50 28.00 99.00 0.0 3.50 7.01 158.44 7 August 1966 o.O 64.00 6.00 27.20 119.70 0.96 1.40 7.00 193.40 0.0 63.80 5.80 22.00 124.40 0.96 1.20 7.35 200.60 LE-1 0 1 0 1 13.3 13.6 22. 5 22.8 8,960 9,270 11,271 11,186 7,889 8,214 9,370 9,928 9. 25 9. 00 9.30 9.40 5.54 5.54 4.87 4.65 20 May 1966 102.50 6.95 0.72 11.26 20.00 25.30 17.12 91.00 110.00 7.38 0.72 11.43 21.76 26.20 16.59 60.25 3 August 1966 0.0 120.00 7.43 0.18 13.80 24.16 24.03 18.84 75.30 0.0 118.80 7.20 0.18 13.84 24.16 23.77 18.61 68.81 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 2 H2S (m) (°C) (iimhos)' (rag/1) pH (cc/1) (mg/1) Na K Ca Mg C03 HC03 Cl S04 20 May 1966 0 12. 5 7,080 5,979 8. 95 5.71 --- 75.00 5.42 0.76 12. 73 14.60 25. 01 11.14 32.96 1 12. 8 8,840 7,675 8. 95 4.87 97.50 6.62 0.62 11. 45 18.40 26. 25 14. 19 46.59 3 August 1966 0 22. 2 11,017 9,627 9. 30 5.15 0.0 116.00 7.05 0.14 13. 88 24. 00 28. 96 18.16 76.79 1 22. 0 11,102 9,482 9. 30 0.0 0.0 117.50. 7. 10 0.14 13. 86 24. 16 29. 28 21.32 69.95 : PHALEROPE 12 May 1966 0 12. 2 4,380 3,164 9. 20 45.00 2. 92 0.32 2. 14 8. 90 18.40 15.24 6.61 1 11. 7 4,340 3,146 9. 15 41.25 3.30 0.46 1. 99 8. 20 19. 10 15.27 6.49 2 5. 6 7,840 6,167 9. 20 --- 90.00 6.51 0.66 3. 62 16. 90 35. 70 28.68 12. 05 3 3. 3 8,490 6,751 9. 20 96.25 7. 00 0.60 3. 95 16.80 39.10 31. 27 11.37 4 3. 3 8,640 .7,157 9. 10 103.75 7. 62 0.40 3. 72 23. 00 40.30 32.50 17.30 27 July 1966 0 18. 9 6,814 5,185 9. 15 5. 26 0.0 71.50 4. 70 0.60 3. 10 15.00 30. 50 24. 50 10.80 1 17. 8 6,864 5,362 9. 15 5. 26 0.0 71.50 5.00 0. 68 3. 00 14.20 31. 70 24. 91 12.70 2 17. 0 6,881 5,336 9. 15 4.65 0.0 70.00 4.90 0. 68 3. 00 14.40 31. 10 24. 91 13. 60 3 14. 4 7,797 6,113 9. 10 0. 50 3.5 80.90 5. 90 0.84 3. 30 17.60 30.80 28. 98 14. 80 4 8. 9 8,8i4 6,953 y. 15 0. 0 u yl.yu 6. 4u 0. /4 j. zti. uu 40. 3u 32. 50 17. 30 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUG TDS C>2 H 2S (m) (°C) (umhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C03 HC03 C l SO^  BOX 20 13 May 1966 0 13. 0 4,680 3,207 9. 10 5.88 . 47.50 3. 78 0.60 2. 29 . 10.30 24.00 14.81 8.09 1 13.3 4,680 3,300 9. 10 5.94 48. 75 3. 69 0.60 2. 20 10.20 23. 90 15.15 2.04 2 10.3 6,750 4,987 9. 10 4.71 58. 75 5. 40 0. 72 3. 32 16.20 35. 70 22. 90 3.88 3 6.7 7,490 5,810 9. 10 3.02 83.75 6. 45 0.86 3. 63 17. 90 40.50 24. 70 8.70 4 4.4 8,830 7,053 9. 20 0. 0 105.00 8. 00 0.66 3. 13 25.80 49.00 31. 20 10.56 5 4.2 11,310 10,012 9. 20 0.0 137.50 9. 60 0. 72 1. 41 37.90 59.00 41. 10 13.46 27 J u l y 1966 0 20. 0 6,220 4,713 9. 10 4.70 0. 0 64.80 4. 30 0. 70 3. 00 14.40 32.10 20/55 8.04 1 19. 7 6,220 4,674 9. 10 4.59 0.0 65.50 4. 20 0. 70 3. 00 15.70 31. 70 . 20.63 5.01 2 17.8 6.237 4.711 9. 1 0 4. 09 0. o 64 60 4(1 n p-4 2„ qn 13.40 3^  no 91 . OS 7. 99 3 15.0 6^ 610 4^ 947 9. 10 2.30 3.5 70.00 5. 00 0.86 3. 10 14. 90 34. 70 22.43 7.46 4 7.8 7,542 5,732 9. 10 0.0 5.0 80.60 5. 80 0.80 3. 20 19. 20 39.20 26. 15 9.68 5 6.4 8,644 6,902 9. 20 0.0 5.0 92.00 6. 50 .--• -- 23.40 44.60 30.15 13.36 BOX 21 13 May 1966 0 13.6 4,720 3,322 9. 00 5.50 -- 51.25 3. 78 0.66 2.23 10. 72 24.59 15.62 12. 29 1 13.6 4,545 3,224 9.10 5.94 48.75 3. 82 0. 72 2.12 10.88 23.61 15. 26 6. 91 2 • 8. 9 7,210 5,711 9. 10 4.43 82.50 5. 60 0. 72 3.64 17.04 29.55 24. 15 10.47 3 5.3 7,230 5,414 9.10 3.34 80.00 6. 28 0.92 3.48 16.80 40.40 25.45 7.65 4 4.4 8,700 6,840 9. 10 0.0 101.25 7. 82 0. 9.2 3.17 22. 56 48.05 32. 52 7.86 5 4.4 JL JL , O J . U 9 ,620 9. 20 G.G 156.25 9. 99 0. 66 1. 08 56.44 62. 00 41. 79 8. 53 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS ° 2 H 2 S (m) (°C) ( u m h o s ) (mg/1) pH (cc /1) (mg/1) Na K Ca Mg C0 3 HC0 3 Cl S.O^  BOX 21 27 J u l y 1966 0 19. 2 6,237 4,716 9. 15 1 19. 4 6,237 4,601 9.20 2 17. 2 6,254 4,721 9. 20 3 15. 0 6,576 4,984 9.20 4 7. 8 8,644 6,770 9.20 5 6. 4 9,831 8,054 9.20 0 11.1 5,090 3,966 9.10 1 11. 7 5,210 3,981 9. 20 2 11. 7 5,090 3,986 9. 20 3 11. 7 5,060 3,993 9.20 4 11. 7 5,130 4,012 9.30 5 10. 6 5,260 4,143 9. 20 6 8.3 5,560 4,433 9.30 7 7. 2 5,560 4,361 9. 25 8 6.7 5,710 4,518 9.30 9 5.8 6,210 4,926 9.30 10 5.6 6,670 5,351 9.30 11 5.6 7,700 6,306 9.30 12 . 5.6 6,950 5,558 9.30 13 5.6 7,700 6,306 9.30 14 5.6 : 8,680 7,120 9. 20 15 5. 6 « 300 7 i 9 20 4. 70 0.0 66.90 4.40 0.68 4.59 0. 0 62.90 4.50 0.68 4.09 0.0 62.60 4.50 0.68 2.30 3.5 66.30 4.80 0.68 0.0 5.0 85.60 6.60 0. 74 0.0 5.0 97.30 : 7. 50 WHITE LAKE 21 May 1966 5.65 _ _•_ 55.00 5.32 0. 20 5. 71 55.00 5.32 0. 12 5.82 57.50 5. 29 0.32 5. 71 57.50 5. 13 0. 20 5.71 52.50 5. 13 0. 12 5. 27 58. 25 5.47 0. 20 3.40 59.00 5. 78 0. 12 3. 29 59. 00 5.78 0.16 2. 08 57.50 6.38 0. 26 0. 0 65.00 7. 22 0.20 0. 0 75.00 7.15 0.26 0. 0 77. 50 6.96 0.26 0. 0 82. 50 7.72 0. 76 0.0 97. 50 7. 72 0.50 0.0 97. 50 9.10 0.46 n n r\ n o 2. 64 13.92 32. 72 20.52 4. 40 3. 05 14.40 32. 16 20.86 5. 90 3. 07 13. 92 32. 72 20. 11 7. 20 3. 26 15.84 33.68 21.48 5. 29 3. 30 21. 28 46.48 28.80 0. 44 — 27.84 50.80 32.63 6. 02 9. 44 26.60 33.70 8.38 7.89 9. 45 26. 10 33. 90 7.07 1.82 9. 30 26.00 34.10 8. 20 5.75 9. 32 25. 90 34.30 8.07 0. 92 9. 41 27.80 32.90 8.51 13.23 9. 51 28.40 34.30 8.38 4.52 9. 80 28.80 37. 10 8. 77 0.0 9. 86 29.40 37.10 8.90 0.85 9. 90 31. 20 37.00 9.09 0.0 9. 80 34.00 40. 20 10. 25 0.0 9. 29 36.60 43. 70 11.30 0.0 9. 29 36.90 43.90 11. 26 0.0 8. 66 39.90 44.60 11.83 0.0 8. 73 40.60 58.80 13.82 0.0 8. 57 42.40 72. 70 15.04 0. 0 /. r> on i /. an A n DEPTH TEMP CONDUC TDS H 2S (ra) (°C) (pmhos) (mg/1) pH ( c c / 1 ) (mg/1) MILLIEQUIVALENTS PER LITER Na K Ca Mg co 3 HCO3 C l so, 0 20.3 5,068 3,937 9.25 8. 96 1 20.3 5,085 4,281 9. 25 8.96 2 20.3 5,085 4,233 9.35 9.07 3 20. 0 5,085 4,121 9.35 8. 74 4 17.8 5,085 4,157 9.35 8.06 5 16. 7 5,085 4,177 9.35 5.60 6 15. 9 5,119 4,280 9.35 4. 06 7 13.9 5,254 4,365 9.35 1.74 8 10.3 5,661 4,728 9.35 0.0 9 8.9 5,780 4,793 9.40 0.0 10 7.8 6,119 5,093 9.40 0. 0 11 6.7 6,695 5 ,666 9. 40 0. 0 12 6.4 6,864 5,933 9.40 0. 0 13 6. 1 7,475 6,354 9.35 0.0 14 5.9 8,390 6,888 9.40 0. 0 15 5.6 8,390 7,559 9.40 0.0 I c e 1,619 1,148 9. 10 0 0.0 5,360 4,300 9. 10 3.42 1 0.0 5,360 4,273 9. 20 3.38 2 0.0 5,360 4,354 9.25 3.44 3 0.0 5,306 4,359 9. 25 3.46 4 0.0 5,306 4,746 9. 25 3. 29 5 0. 0 5,316 4,399 9. 25 3.00 4 J u l y 1966 0.0 57.50 4.10 0.0 57.00 4.10 0.0 57.50 4.00 0.0 57.00 4.80 0.0 56.80 4.00 0.0 59.50 4.20 0.0 57.50 4.20 0.0 60.00 4.20 3.5 67.00 4.70 5.0 67.50 4.70 5..0 71.00 5. 10 5.0 77.50 5.60 5.0 81.30 5.90 5.0 90.50 6.30 5.0 99.30 7.00 5.0 102.30 7.10 16 F e b r u a r y 1967 15.00 56.50 52.00 52.00 52.00 52.00 57.50 0. 18 0. 18 0. 10 0. 20 0. 14 0. 10 0. 16 0. 14 0.16 0.30 0.14 0.18 0.14 0.16 0.21 0. 18 9.90 9.90 9.90 80 80 80 60 30 9 9 9 9 9 9.80 9. 70 9. 60 9. 30 9.40 9.20 8.80 8. 70 28. 60 28.60 28. 20 29.30 29.40 29.04 29. 10 30.40 33. 00 33.10 35.00 40. 00 40.80 44.60 41.60 49.40 31.60 31.60 31. 70 30.80 30. 90 31.60 32. 20 32.40 34. 90 35.90 37. 70 40. 70 43. 10 51.00 59. 10 53. 70 8. 18 8.54 8. 18 8.13 8.74 8. 16 8.35 8.80 9.50 9.46 11.38 I I . 25 12.10 15.42 15.49 14.85 11. 90 4.86 0.0 0.08 2. 20 0.0 0. 0 0.0 0. 0 0.0 0. 0 0. 0 0.0 0.0 0. 0 0. 25 1.18 0.15 3. 30 9.60 8. 80 2. 76 0.0 4.32 0.17 10. 17 29. 76 37. 30 8. 76 0.0 4.50 0.17 10. 19 29.36 37. 64 8. 77 0.31 4.50 0.17 10. 14 29.44 37. 46 8.77 0.0 4.50 0. 18 10. 14 . • 29.28 37. 42 8. 63 0.0 4.50 0. 18 10. 04 29. 12 37. 78 8.65 0.0 4.37 0. 18 10. 08 29.12 37. 58 8. 75 0.0 CO VO MILLIEQUIVALENTS PER LITER pH (cc/1) (mg/1) Na K Ca Mg CO3 HCO3 Cl SO, WHITE LAKE 16 February 1967 (cont.) 6 0.0 5,316 4,285 9.25 2.49 56.50 4.47 0.18 10.08 29.92 • 39.08 9.94 0.67 7 0.0 5,338 4,295 9. 25 1.37 --- 52.00 4.57 0.18 10.08 29.36 37.64 9.60 0.15 8 0.3 5,272 4,349 9.25 0.0 56.50 4.55 0.18 10. 06 28.88 37.62 9.09 0.0 9 0. 7 5,316 4,359 9. 25 0.0 56.50 4.75 0.33 10.07 29. 28 37.32 9.38 0.0 10 1.0 6,104 4,969 9. 25 0.0 63,75 5. 25 0.30 9. 27 37.00 35.00 10.82 0.0 11 1.5 5,864 4,861 9.25 0.0 63.75 5. 13 0. 52 9. 24 32.84 41.56 10.59 0.0 12 3.5 6,269 5,193 9.25 0. 0 68.75 5.38 0. 19 9.25 36. 00 44. 00 11.65 0. 0 13 4.0 7,187 5,940 9.20 0.0 76.25 6.20 0.19 9. 07 36.80 52.10 12.57 0.0 14 4.0 7,187 6,439 9. 15 0.0 82.50 6.38 0. 22 8.78 38.80 55. 90 15.00 0.0 LE-5 22 May 1966 0 12.0 3,870 3,058 8.40 6.59 40.00 3.09 0. 72 12.28 2.44 40.30 7.23 6.08 5 August 1966 0 20.0 4,136 3,185 8. 70 3.30 0.0 38.30 3.18 0.30 11.79 5. 76 38.12 7.53 2.45 BOITANO LAKE 10 May 1966 0 14.4 3 .905 3.105 9.00 4. 95 34.50 2. 92 0.86 11. 08 5.00 17. 70 4.25 22. 74 1 13.3 3,850 3,051 9.00 4.90 34.00 2. 96 0.76 11. 07 5. 10 17.40 4.08 29. 17 DEPTH TEMP CONDUC TDS (m) (°C)_ (iimhos) (mg/1) MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0^ K^S (m) (°C) (pmhos) (mg/l) pH (cc/1) (mg/1) Na K Ca Mg C03 HC03 Cl SO 4 BOITANO LAKE 10 May 1966 (cont.) 2 13.3 3,850 2,958 9.00 4.90 34.00 3. 00 0.66 11.23 5.10 17.70 4.11 18.71 3 13.3 3,790 3,070 8.95 4.68 33.50 3. 07 0.72 11. 19 4.80 17.70 4.25 18.43 4 11.7 4,035 3,095 8. 90 4.68 35. 00 2. 81 0.86 11.17 5.00 17.80 5.17 20.59 C 8.9 4,190 3,210 8.80 0.0 38.50 3. 15 0.72 12.39 3.90 21.40 4.45 17.63 26 July 1966 0 18.9 4,119 3,223 8.85 4.14 0.0 33.80 3. 20 0.80 11.70 5.10 17.40 4.61 22.60 1 18. 9 4,102 3,220 8.85 4.09 0.0 33.30 3. 40 0.70 11. 90 5.10 17.60 4.72 23.10 2 17.8 4,119 3,193 8.85 4.26 0.0 33.80 3. 50 0.76 11.90 5.00 18. 50 4.74 22. 90 3 17.8 4,136 3,161 8.90 3.98 0. 0 33.80 3. 60 0.68 11.90 5.30 18. 20 4.83 23.70 4 17.2 4,136 3,142 8.85 3.86 0.0 34.10 3. 50 0.68 12.00 4.80 18.60 4.70 23. 10 _ •» —» r\ /-» r\ o / *"» r\ 1 o r\ -in on /. OA o i on o /. n n 0 J. / . Kl «4 , £- O • / \J \J • \J J-t. -r. RUSH 11 May 1966 0 15.3 3,550 3,773 8.70 4.50 30. 00 2.32 0.74 10.60 3. 00 17.00 3.65 16.86 1 15.0 3,490 3,715 8.70 4.65 30.50 2.32 0.76 10.66 3. 10 16.90 3.90 16.00 2 15.3 3,960 3,113 8.50 0.0 43.50 2.55 0.80 12. 96 2. 40 21.40 4.38 19.76 31 July 1966 0 20. 0 4,254 3,208 8.60 3.53 0.0 34.40 3. 10 0.64 12.80 4. 00 19. 90 4.86 23.80 1 ' 18.3 4,254 3,280 8.60 4.00 0.0 35.10 3.00 0.76 13.00 3. 70 20.20 4.51 24.01 2 . 17.8 4,254 3,309 8.60 3. y/ 0.1 35.30 3. 00 u. au 12. /u 3. 50 20.40 4. 59 24. 92 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 2 H2S (in) (°C) (pirihos) (mg/1) pH (cc/1) (mg/1) Na K. Ca Mp CO, HCOc Cl SO, LE-3 22 May 1966 0 11.7 3,435 2,709 8.50 5.49 --- 30.00 2.62 0.90 13.87 4.40 27.25 5.16 5.91 5 August 1966 0 19.7 3,915 3,237 9.20 7.17 0.0 35.30 2.75 0.24 13.49 15.84 24.48 6.86 6.50 0 13.1 3,100 2 ,498 8 . 7 0 6 .47 1 1 2 . 8 3 , 2 i U 2 , 4 / 0 8 . / U b . ^ b LE-4 22 May 1966 27.50 21. 5 U 5 August 1966 2.36 0.72 11.96 id. 4 1 v. io i i . y z 3.52 31.00 2,899 2,612 8.00 s i x MILE' 17 May 196 8 5.41 0.33 8.56 26.34 5.08 5. 26 0.0 1 0 6.19 0 20.3 3,864 3,117 9.20 8.90 0.0 35.30 2.58 0.24 12.71 14.40 26.16 6.65 3.80 1 20.3 3,864 3,262 9.20 8.46 0.5 35.30 2.70 0.44 12.43 14.24 25.48 6.65 2.45 3.85 0.53 31.02 DEPTH TEMP CONDUC TDS 0„ H0S (ra) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1) Na NEAR OPPOSITE BOX 4 14 May 1966 0 12.8 2,264 1,955 8.90 5.85 --- 13. 75 1,42 1.86 12.42 1.60 . 4.80 2. 90 23.09 1 12.8 2,320 1,939 8.90 5.74 13.75 1.50 1.62 12.72 1.80 4.70 2. 78 21.55 2 12. 2 2,830 2,961 8. 70 5.53 20. 75 1.83 2.28 19.25 1.80 7.80 4.15 26.00 2 7.8 3,660 3,379 8.60 1.92 24.00 2. 25 2.78 22.03 1.60 8.80 4.73 32.02 28 July 1966 0 20.9 3,136 2,782 9.00 8.06 0.0 17.00 1.40 1. 08 18.70 2.70 6.20 3.92 25.90 1 20.9 3,169 2,978 8.80 8.23 0.0 16. 90 1.60 1.02 18.80 2.30 6.60 3.94 24.40 2 17.8 3,186 3,024 8.80 5.10 0.0 17. 10 1.60 1.00 19.00 2.10 6.80 3.95 25.40 3 15.9 . 3,390 3,163 8.65 0.0 5.0 18.50 1.50 1.26 20. 20 1.80 8. 10 4.22 28.70 BOX 89 17 May 1966 0 13.3 1,525 1,096 8.39 6.16 _ « 13.25 1.74 0.86 2.53 0.32 13.70 1.27 2.19 1 • 13.0 1,490 1,059 8.60 6.22 13.25 1.69 0.76 2.59 0.96 12.60 1.28 3.53 2 11.1 1,510 1,125 8.58 6.05 13.28 1.74 0.80 2.50 0. 96 ,12.60 1.13 1.93 29 July 1966 0 19.2 1,695 1,329 8.70 4.42 0.0 14.30 1.50 0. 28 3.30 2.40 12.70 0.98 4.52 1 18.9 1,695 1,387 8.80 4.12 0.0 14.20 1.30 0.32 3.30 2.60 12.40 1.01 2.84 2 19. 2 1,695 1,375 8.80 0.90 1.0 14. 50 1.30 0.30 3.30 2.40 12.50 1.04 3. 15 MILLIEQUIVALENTS PER LITER K Ca Mg CO HC03 Cl SO, .MILLIEQUIVALENTS PER.LITER DEPTH TEMP CONDUC TDS 0 o H0S (m) (°C) (jimhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg CO3 HC0 3 Cl S 0 4 ROCK 17 May 1966 0 12. 6 1,395 1,049 8 80 6. 39 14.25 1.08 0. 84 1. 47 3 10 . 13 00 2. 23 0.0 1 12. 2 1,415 1,053 8 90 6. 34 14.00 0.94 0. 84 1. 47 3 20 12. 90 1. 59 0.0 2 11. 7 1,395 1,057 8 90 6. 32 -- 14.25 1. 12 0 96 1 41 3 40 12 70 1 52 0.34 30 July 1966 0 20. 3 1,593 1,184 9 00 4. 14 0. 0 15.80 0.99 0. 38 2. 00 5 40 11. 80 1. 26 0. 01 1 20 3 1,585 1,249 9 10 3. 15 0. 0 15.80 0.93 0. 36 2 00 5 50 11 80 1 30 0.45 2 18. 3 1,585 1,119 9 10 2. 86 0. 1 15.80 0.92 0 46 1 90 5 50 11 60 1 28 1.12 GR-3 10 September 1967 0 — -- 1,500 1,241 8 90 --•-- -- 12.99 0.95 1. 52 6. 28 1 07 12 62 0. 61 1.95 SORENSON LAKE 4 May 1967 0 — 1,500 1,031 8 90 — 4.43 0.69 1 15 11 95 1 33 6 38 0 34 9.29 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 o H S 2 2 (m) (°C) (umhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C03 HC03 Cl S0 4 NEAR PHALEROPE 12 Meiy 13oo 0 14.4 1,170 867 8. 60 5. 57 ---- 7.25 1. ,38 0. 92 5. 19 1. 30 11. 70 0.41 3. 61 1 14.4 1,189 853 8. 60 4. ,86 ---- 7.25 1. ,23 0. 96 5. 17 1. 50 12. ,20 0.42 0. 0 2 14.4 1,189 873 8. 60 4. 44 -- 7. 25 1. 31 0. 90 5. 28 1. 40 12. 30 0.86 0. 0 27 July 1966 0 18.9 1,492 865 8. 10 1. ,04 0. 0 9.00 1. ,70 0. 84 6. 30 0. 0 16. ,90 1.02 0. 0 1 17.8 1,475 854 8. 15 1. 23 0. 0 8. 75 1. 50 0. 94 6. 30 0. 0 16. 60 1.06 0. 0 2 16.4 1,492 850 8. 10 1. 18 0. 0 8.75 1. ,60 0. 94 6. 30 0. 0 16. , 70 1.26 ' 0. 30 WESTWICK LAKE 10 May 1966 0 15. 6 1,208 948 8.90 3. 60 --• - 4.63 0.79 1. 26 11. 06 1. 20 7. 40 0.10 7.82 1 15. ,6 1,282 1,170 8.90 3. 67 4.63 0.86 1. 36 11. 12 1. 20 7. 10 0.12 6.47 2 15. .6 1,189 1,159 8.90 3. 63 4.48 0.86 1. 22 11. 28 2. 30 6. 20 0.05 8.84 3 15. 6 1,189 1,066 8. 90 3. 31 -- 4.88 0,79 . 1. 22 11. 22 2. 30 6. 80 0.21 4.54 26 July 1966 0 23. ,3 1,356 969. 8.80 1. 96 0. 0 4.59 0.88 0. 70 11. 30 1. 90 8. 20 0.34 6. 59 1 21. 1 1,322 918 8.75 1. 68 0. 0 4.78 0.82 0. 76 11. 30 1. 90 8. 20 0.34 8.47 2 19. ,4 1,339 901 8.80 0. 22 0. 0 4.73 0.85 0. 76 11. ,30 2. 10 8. 00 0.42 5.44 December 1966 0 _ _ . 1.130 1,026 9.40 _ _ 4.54 0.62 0 .40 7 .34 3 .80 2 .98 0.11 8.33 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS Q>2 (m) (°C) (umbos) (mg/1) pH (cc / 1 ) (mg/1) Na K Ca Mg C 0 3 H C 0 3 C l S 0 4 WESTWICK LAKE 0 — •- 1 , 5 5 9 1 , 3 9 5 9. , 0 5 -• — 6 . 2 5 0 . 88 0 . , 5 1 1 3 . 1 5 3 . 20 5. 25 0 . 12 9 . 76 18 February 1967 Ice . _ 428 292 9. 30 _. . 1 . 4 1 0 . 13 0 . 49 3 . 23 3 . 10 2. , 0 0 0 . 0 8 1 . 63 0 0 . 9 1 , 7 7 0 1 , 4 7 9 8 . 10 - - — 6 . 3 8 0 . 90 1. 42 1 5 . 1 4 0 . 0 9 . 42 0 . 16 4 . 88 1 2 . 7 1 , 7 7 0 1 , 2 5 5 8 . 05 1 . 02 6 . 3 8 0 . 90 1. 44 1 5 . 1 4 0 . 0 9 . 42 0 . 13 1 0 . 70 2 3 . 6 1 , 7 7 0 1 , 4 4 7 7. , 9 0 0 . 20 6 . 2 5 0 . 90 2. 31 1 4 . 19 0 . 0 9. , 3 0 0 . 13 1 0 . 08 LAC DU BOIS j> ON 23 May 1966 1 A A r\ /. / > - T T - 7 r t-/ w _) r\ O . 50 u. y J ---- 2. D(J 0. 7 i i . 30 • 8. , 5b 0 . « 0 1 1 . 3 0 0. 18 0 . 0 1 1 3 . 9 944 . 761 8. 50 6. 78 ---- 2 . 5 0 0 . 6 7 1. 76 8 . , 1 8 1 . 52 1 0 . 0 0 0 . 17 0 . 0 2 1 3 . 3 732 718 8 . 65 7. 06 -- 2. 50 0 . 67 1. 92 8 . , 0 0 1. 60 9 . 90 0. 25 1 . 9 0 3 1 2 . 2 948 724 8 . 60 6 . 41 --• - 3 . 25 0 . 6 7 1. 84 8 . , 1 0 1 . 60 9 . 90 0 . 17 1 . 10 4 1 1 . 1 919 727 8 . 50 6. 08 ---- 3 . 0 0 0 . 7 5 1 . 76 8 . 20 1 . 52 1 0 . 0 0 0 . 20 0 . 4 3 5 1 0 . 0 948 782 8 . 70 3 . 91 ---- 3 . 0 0 0 . 67 1 . 54 8 . , 4 2 1 . 60 9 . 9 0 0 ; 12 1 . 8 7 6 9 . 2 948 760 8 . , 8 0 1. 58 _. .- 3 . 0 0 0 . 7 5 1. 64 8 . , 3 0 1. 60 1 0 . 0 0 0. 21 1 . 0 1 7 8 . 6 925 756 8 . 70 0. 60 3 . 50 0 . 75 1. 60 8 . , 3 2 1. 60 . 9 . 8 0 0 . 18 1 . 0 5 8 7. 8 1 , 0 9 4 842 8 . 14 0 . 43 3 . 50 0 . 71 2. 32 8 . , 55 0 . 0 1 1 . 61 0 . 26 0 . 21 6 August 1966 0 2 2 . 2 966 859 8 . 60 5. 35 0 . 0 2 . 50 0 . 4 8 0 . 92 9. , 8 0 1. 80 9 . 1 0 0. 27 1 . 4 4 1 2 2 . 2 966 764 8 . 80 5. 29 0 . 0 2 . 4 0 0 . 5 3 0 . 92 9. , 3 0 1. 90 9 . 0 0 1. 06 0 . 7 3 2 2 1 . 1 966 111 8 . 80 5 . 26 0 . 0 2 . 3 5 0 . 4 5 0 . 96 9. , 2 0 1. 90 8 . 9 0 0 . 43 0 . 8 0 DEPTH TEMP CONDUC TDS (m) (°C) (pmhos) (mg/1) LAC DU BOIS 6 August 1966 (cont.) 3 20. 9 949 819 8.75 4. 82 0.0 2.37 0. 55 0. 92 9. 20 1. 90 8.80 0.22 1.20 4 20.3 966 746 8. 75 3. 70 0.0 2.38 0. 58 1. 02 9. 10 1. 70 9. 00 0.21 1.39 5 17. 8 966 738 8. 75 1. 37 0.0 2.35 0. 53 1. 02 9. 10 1. 70 9. 00 0.19 1.02 6 13. 9 975 753 8. 45 0. 0 5.0 2.35 0. 63 0.96 9. 10 1. 20 9. 70 0. 19 1.37 7 11. 7 1,008 776 8. . 20 0. 0 5.0 2.34 0. 78 1. 32 8. 80 0. 0 10. 90 0.16 1.56 8 10. 0 1,025 826 8. ,20 0. 0 4.0 2.35 0. 75 1. 32 9. 00 0. 0 11. 10 0.27 1.42 NEAR OPPOSITE CRESCENT 16 May 1966 0 11. 7 1,048 569 8. 60 5. 48 ~ _ _ 3.40 0. 67 1. 46 3. 82 0. 72 6. 40 0.30 1.20 I II. 7 /30 541 8. 60 0. 85 _ _ _ 3.45 0. 64 1. 34 3. 92 0. 64 6. 40 O. 21 1.05 2 11. 7 722 571 8. ,70 5. 85 3.35 0. 64 1. 30 3. 96 0. 88 6. 30 0. 24 0.92 3 11. 1 798 561 8. 70 5. 64 3.35 29 July 1966 0. 64 1. 22 4. 00 0. 80 6. 30 0. 20 0. 96 0 18. 9 781 654 8. 60 0. 92 0.0 3.33 0. 56 0. 56 4. 90 0. 88 6. 70 0.45 1.90 1 18. 6 783 630 8. 55 0. 34 0.0 3.33 0. 54 0. 58 4. 90 0. 96 6. 60 0.45 1.30 2 17. 5 780 634 • 8. 60 0. 22 0.3 3.35 0. 54 0. 58 4. 90 1. 60 6. 00 0.45 1. 70 3 15. 9 788 623 8. 70 0.0 5.0 3.31 0. 57 0. 56 5. 00 1. 12 6. 60 0.43 1.60 MILLIEQUIVALENTS PER LITER °2 H 2 S pH (cc/1) (mg/1) Na K Ca Mg CO3 HCO3 Cl S0 4 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 2 E^S (m) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C0 3 HC03 Cl SO4 BOX 17 16 May 1966 0 12. 8 733 472 • 8.30 6. 38 3. 25 0. 67 1. ,06 4.33 0. 0 8. 90 0.13 0. 23 1 12. •5 734 466 8.30 6. 81 3. 35 0. 75 1. ,10 4.35 0. 0 8.90 0. 13 0. 0 2 12. 2 733 485 8.30 6. ,06 3. 35 0. 67 1. 08 4.36 0. 0 8.80 0.18 0. 0 3 11. 7 870 428 8.30 5. ,53 3. 30 0. 67 1. ,08 4.36 0. 0 8.90 0.18 0. 0 29 July 1966 0 19. 2 746 629 8.80 5. 43 0. 0 3. .33 0. 50 0. 38 5.00 2. 01 6. 20 0.14 0.09 1 19. 2 732 749 8.80 2. 30 0. 0 3. ,30 0. 52 0. 46 4.90 2. 72 6.40 0.14 0.24 2 16. 7 729 576 8.80 0. 45 0. 1 3. 31 0. 51 0. 48 4.90 2. 16 6. 90 0.07 0.46 3 15. 3 781 738 8.60 0. 0 4. 0 3. , 28 0. 57 0. 48 5.20 1. 28 8. 10 0.24 0.52 BOITANO NE 0 16.1 595 338 7.70 0 17.8 712 427 9.05 10 May 1966 3.20 0.49 0.68 2.33 0.0 5.01 0.10 2.80 26 July 1966 0.0 3.80 0.57 0.86 3.31 1.76 5.84 0.0 0.28 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0_ H„S (m) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg CO, HCO„ Cl SO4 BOX 17 16 May 1966 0 12. 8 733 472 ' 8. .30 6. 38 ---- 3. 25 0. 67 1. 06 4.33 0. 0 8. 90 0. 13 0. 23 1 12. 5 734 466 8. .30 6. 81 , ---- 3. 35 0. 75 1. 10 4.35 0. 0 ' 8. 90 0. 13 0.0 2 12. 2 733 485 8. ,30 6. 06 ---- 3.35 0. 67 1. 08 4.36 0. 0 8. ,80 0. 18 0.0 3 11. 7 870 428 8. ,30 5. 53 •- 3.30 0. 67 1. 08 4.36 0. 0 8. 90 0. 18 0.0 29 July 1966 0 19. 2 746 629 8, ,80 5. 43 0. 0 3.33 0. 50 0. 38 5.00 2. 01 6. 20 0. 14 0.09 1 19. 2 732 749 8. .80 2. 30 0. 0 3.30 0. 52 0. 46 4.90 2. 72 6. ,40 0. 14 0.24 2 16. 7 729 576 8. ,80 0. 45 0. 1 3.31 0. 51 0. 48 4.90 2. 16 6. 90 0. 07 0.46 3 . 15. 3 781 738 8. ,60 0. 0 4. 0 3. 28 0. 57 0. 48 5.20 1. 28 8. 10 0. 24 0. 52 BOITANO NE 10 May 1966 0 16. 1 595 338 . 7. 70 -- ---- 3. 20 0. 49 0. 68 2.33 0. 0 5. 01 0. 10 2.80 26 July 1966 0 17. 8 712 427 9, .05 . 0. 0 3.80 0. , 57 0. ,86 3.31 1. 76 5. ,84 0. 0 0. 28 DEPTH TEMP CONDUC TDS 0 2 H2S MILLIEQUIVALENTS PER LITER (m) (°C) (jumhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C03 HCO-j Cl S0 4 OPPOSITE BOX 4 14 May 1966 0 11. , 1 564 404 8. .60 4. ,15 ---- 1.85 0.60 0. ,93 3. 81 0. ,48 5. 90 0.15 0. 0 1 11. 7 . 562 393 8. .70 4. ,04 - --- 1.85 0.60 1. ,00 3. 85 0. ,56 5.90 0.15 0. 0 2 11. 9 . 564 411 8. .60 3. ,94 ---- 1.85 0.60 1. ,08 3. 80 0. 64 . 5.80 0.15 0. 0 3 10. 6 673 . 471 8. . 50 2. ,02 ---- 2. 05 0.71 1. ,32 4. 31 0. 48 7.00 0. 18 0. 40 28 J u l y 1966 • 0 20. 3 625 502 8. .80 •. 5. 71 0. 0 2.06 0.64 0. 38 5. 10 0. 0 5.90 0.32 0. 16 1 18. 9 624 488 9. ,00 4. 87 0. 0 2.10 0.62 0. 38 5. 20 1. 08 5. 20 0.35 0. 91 2 17. 6 615 627 9. .00 3. 53 0. 0 2.00 0.63 0. 32 5. 20 1. 02 5.30 0.37 0. 45 3 16. 7 625 537 9. ,00 3. 14 0. 0 2.00 0. 63 0. 34 5. 20 0. 88 5.50 0.40 0. 20 RACETRACK 16 May 1966 0 12. 2 481 349 8. ,00 5.21 3. ,05 0. 45 0. 76 1. .42 0. 0 5. 24 0; 16 0.13 1 11. ,9 481 338 8. ,00 5.00 3. , 10 0. 53 0. 76 1. .41 0. 0 5. 24 0. 46 0.0 2 11. 7 489 333 8. ,00. 4.95 --• 3. , 10 0. 49 0. 76 1. ,40 0. 0 , 5. 24 0. 21 0.48 3 11. 7 . 472 298 8. ,00 4.95 3. 15 0. 49 0. 76 1. .41 0. 0 5. 24 0. 20 1.63 4 11. 7 474 335 8. ,00 4.68 3. 05 0. 53 0. 76 1. .41 0. 0 5. 24 0. 20 1. 23 5 11. 4 519 317 8. ,00 4.57 3. 10 0. 45 0. 80 1. .37 0. 0 5. 18 0. 05 1.03 6 11. 1 645 272 7. ,90 3.94 3. .15 0. 45 0. 82 1. ,32 0. 0 5. 18 0. 03 0.61 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS °2 H 2 S ' — (m) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1) Na K Ca Mg C03 HC03 Cl S0 4 RACETRACK 0 18. 6 495 435 8.60 5. 40 0. 0 3. 10 0. 41 0.30 1. 90 0. 88 4.80 0. 14 0. 43 1 18. 6 495 407 8.60 5. 38 0. 0 3. 29 0. 42 0.34 1. 90 0. 96 4. 60 0. 16 0. 0 2 17. 8 495 406 8.80 2. 80 0. 0 3. ,26 0. 40 0.36 1. 90 1. 07 4.40 0. 16 0. 06 3 16. 7 497 398 8. 70 1. 90 0. 1 3. ,41 0. 38 0.32 1. 90 0. 88 4. 70 0. 11 0. 23 4 15. 9 505 415 8. 50 0. 50 0. 5 3. 26 0. 38 0.40 1. ,90 0. 64 4. 90 0. 21 0. 07 5 15. 0 508 394 8.30 0. 56 0. 6 3. 31 0. 38 0.40 1. ,90 0. 0 5.60 0. 14 ' 0. 0 6 14. 4 510 665 8.00 0. 0 2. 0 3. ,29 0. 38 0.40 1. ,90 0. 0 5. 70 0. 06 0. 23 SP 6 15 May 1966 o 0 1 13. 3 12.8 240 270 114 7. 70 7.95 6. /5 5.42 u. i/ 0. 27 u. i l 0.11 u. 8 b 0.92 i . <+/ 1.42 u. u 0.0 2.90 U . VO 0.11 0.0 0 1 24. 2 20.0 256 257 131 73 9.00 8. 70 7.22 5. 71 31 July 1966 0.0 0.41 0.1 0.41 0. 75 0. 78 0.80 0. 74 2. 20 2. 10 1.07 0.80 2. 12 2.40 0.0 0. 02 0.04 0.06 BOX 27 0 1 13.3 12.8 37 44 31 42 6.40 6.40 6.39 5. 59 17 May 1966 0.15 0.12 0.11 0.11 0.16 0.26 0.0 0.0 0.0 0.0 0.40 0. 29 0.06 0.11 0.13 0.15 MILLIEQUIVALENTS PER LITER DEPTH TEMP CONDUC TDS 0 2 H2S (m) (°C) (pmhos) (mg/1) pH (cc/1) (mg/1). Na K Ca Mg CO3 HC0 3 • Cl* SO 4 30 July 1966 0 19.4 41 9.15 4.20 0.0 0.02 0.07 0.06 0.16 0.12 0.14 0.02 0.01 1 17.2 33 6.0 6.20 4.20 0.0 0.02 0.07 0.10 0.12 0.0 0.39 0.10 0.17 Appendix III. Percentage of the total volume of each lake present in successive meter intervals of depth (i._e. , the percentage listed for 0 m indicates the percentage of the total volume of the lake present between 0 m and 1 m depth). PERCENTAGE VOLUME OF .LAKES AT SUCCESSIVE METER INTERVALS I N DEPTH LAKE 0 1 2 3 4 5 CLINTON G R - 2 1 0 0 . 0 L B - 1 3 1 . 1 2 8 . 0 2 1 . 3 1 4 . 8 4 . 7 0 . 1 LB - 2 u 5 . 9 3 1 . 1 J. u LONG LAKE 4 0 . 5 2 9 . 6 1 8 . 8 1 0 . 4 0 . 7 BOX 4 4 2 . 4 3 1 . 2 1 8 . 6 7 . 2 0 . 6 BOWER'S LAKE 5 9 . 8 3 5 . 6 4 . 6 L E - 1 7 0 . 3 2 8 . 3 1 . 4 L E - 2 7 7 . 1 2 2 . 5 0 . 4 PHALEROPE 3 2 . 8 2 4 . 2 1 8 . 9 1 3 . 0 7 . 2 3 . 8 BOX 2 0 - 2 1 3 1 . 7 2 4 . 8 1 9 . 6 1 4 . 5 8 . 8 0 . 6 WHITE LAKE 1 8 . 8 1 7 . 0 1 4 . 6 1 1 . 4 8 . 9 7 . 2 L E - 5 1 0 0 . 0 BOITANO LAKE 3 3 . 1 2 8 . 9 2 3 . 8 1 3 . 0 1 . 2 RUSH 6 6 . 2 3 0 . 6 3 . 2 NR OP BOX 4 6 1 . 3 3 6 . 1 2 . 6 L E - 3 9 1 . 8 8 . 2 L E - 4 6 0 . 5 3 9 . 5 S IX MILE BOX 89 6 7 . 9 3 0 . 0 2 . 1 ROCK 6 8 . 2 2 9 . 5 2 . 3 G R - 3 NR PHALEROPE 6 1 . 0 3 2 . 0 7 . 0 WESTWICK LAKE 6 1 . 2 2 8 . 2 8 . 8 1 . 7 0 . 1 SORENSON LAKE LAC DU BOIS 2 3 . 4 1 9 . 6 1 6 . 3 1 3 . 4 1 0 . 9 8 . 0 NR OP CRESCENT 5 3 . 4 3 2 . 9 1 3 . 3 0 . 4 BOX 17 5 9 . 4 2 8 . 3 9 . 8 2 . 5 10 11 12 13 14 15 BOITANO NE OP BOX 4 RACETRACK S P - 6 R O Y 9 7 8 3 . 1 4 1 . 3 94 . 1 1 6 . 8 2 4 . 4 5 . 9 L. 0 . 1 1 5 . 9 5 . 2 2 . 6 1 . 1 0 . 5 5 . 7 2 . 0 0 . 2 

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