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AN ECOLOGICAL STUDY OF SOME OF THE CHIRONOMIDAE INHABITING ASERIES OF SALINE LAKES IN CENTRAL BRITISH COLUMBIA WITH SPECIAL REFERENCE TO CHIRONOMUS TENTANS FABRICIUS by Robert Alexander Cannings BSc. Hons., University of British Columbia, 1970 ATHESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in theDepartment of Zoology We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, 1973 In presenting this thesis in partial fulfilment of the requirements for agree that an advanced degree at the University of British Columbia, Ithe Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia Vancouver 8, Canada Date i i ABSTRACT This thesis is concerned with astudy of the Chironomidae occuring in a saline lake series in central British Columbia. It describes the ecological distribution of species, their abundance, phenology and interaction, with particular attention being paid to Chironomus tentans. Emphasis is placed on the species of Chironomus that coexist in these lakes and a further analysis is made of the chromosome inversion frequencies in C. tentans. Of the thirty-four species represented by identifiable adults in the study, eleven species have not been previously reported in British Columbia, five are new records for Canada and seven species are new to science. The chironomid fauna of the lake series is divided into dominant associations whose existence seems to depend on salinity and productivity levels. Procladius bellus Ablabesmyia peleensis association pre- vails in the lowest salinities (40 to 80 jumho/cm conductivity) while in conductivities between 400 and 2800 jumho/cm Glyptotendipes barbipes Einfeldia pagana association dominates. In the most saline lakes (conductivity 4100 to 12000 Calopsectra gracilenta Cryptotendipes ariel association ischaracteristic. ACricotopus albanus - -a -jumho/cm) <ul style="display: flex;"><li style="flex:1">a</li><li style="flex:1">-</li></ul>Analysis of physical and chemical factors influencing the l i f e cycle of C. tentans indicates that conditions associated with high levels of organic carbon promote large numbers of larvae and greater emergence success. The results suggest that competition between C. tentans and other Chironomus species isreduced through spatial separation due to different preferences forsalinity or related factors. Furthermore, temporal separation among these and other abundant species such as G. barbipes and E. pagana occurs as a result of staggered generation times. The inversion frequency inchromosome 1 of C. tentans is negatively correlatedwith organic carbon levels and positively correlated with dissolved oxygen and the abundance of Glyptotendipes barbipes. Since the inversion frequency is lowest inhabitats where competing species are few and where C. tentans ismost successful, i tissuggested that the inversion governs amechanism reducing competition. Amajor contributionof this work istherevision ofthe distribution of many of the chironomid species under considera- In the past, l i t t l e research has been done on populations of chironomids ina saline lake series. The present this gap inentomological research, tion. study, inattempting to shows that f i l l aspecies' l i f e history and population structure can vary radically inclosely associated lakes of differing chemical and biological constitution. IV TABLE OF CONTENTS Page Title Page iAbstract <ul style="display: flex;"><li style="flex:1">i</li><li style="flex:1">i</li></ul>Table of Contents List of Tables List of Figures List of Plates Acknowledgements iv v i v i i i x i x i i IINTRODUCTION 1II THE LAKE ENVIRONMENTS A. THE STUDY AREA 44 B. THE PHYSICAL AND CHEMICAL PROPERTIES OF THE LAKES 8III SPECIES DIVERSITY AND THE CHIRONOMID COMPLEX IN THE LAKE SERIES 17 <ul style="display: flex;"><li style="flex:1">17 </li><li style="flex:1">A. </li></ul>MATERIALS AND METHODS <ul style="display: flex;"><li style="flex:1">1. Temperature Records </li><li style="flex:1">17 </li></ul>17 17 19 2. Biological Sampling Methods a) Larval Sampling b) Adult Sampling <ul style="display: flex;"><li style="flex:1">3. Rearing of Specimens </li><li style="flex:1">23 </li></ul>4. Preparation and Identification of Specimens 24 <ul style="display: flex;"><li style="flex:1">24 </li><li style="flex:1">5. </li><li style="flex:1">Analysis of the Data </li></ul>6. Storage of the Data for Further <ul style="display: flex;"><li style="flex:1">Study </li><li style="flex:1">24 </li></ul>26 B. RESULTS 26 30 1. Water Temperatures in the Lake Series 2. Chemical Data <ul style="display: flex;"><li style="flex:1">3</li><li style="flex:1">The Occurrence of Species in the </li></ul><ul style="display: flex;"><li style="flex:1">Lakes </li><li style="flex:1">30 </li></ul>4. 5. Species Considered in Detail The Chironomid Complex and the Lake Series 33 85 a) The Cricotopus albanus -Procladius bellus -Ablabesmyia <ul style="display: flex;"><li style="flex:1">peleensis association </li><li style="flex:1">85 </li></ul>86 b) c) The Glyptotendipes barbipes -Einfeldia pagana association The Calopsectra gracilenta Cryptotendipes a r i e l association -88 C. DISCUSSION 1. 2. The Chironomidae andthe Lake Series 91 Chironomus tentans andthe Lake Series 102 a) b) Physical and Chemical Influences Biotic Interactions 102 106 IV CHIRONOMUS TENTANS AND SOME BIOTIC FACTORS AFFECTING CHROMOSOME INVERSION 113 113 115 121 125 A. B. MATERIALS AND METHODS RESULTS C. DISCUSSION CONCLUSION rV 129 140 Literature Cited Appendix vi LIST OF TABLES Page 10 TABLE IPhysical and chemical of the lakes. properties TABLE II Average water temperature in the lake series. 27 TABLE III The distribution of chironomids in the one meter depth zone in the lakes. 31 32 TABLE IV The distribution of larvae unidentified to species. TABLE VSummary of correlation coefficients describing therelationship between environmental factors andthe amount of emergence of certain species. 76 77 The correlation between emergence histogram dispersion and environmental factors. TABLE VI TABLE VII Summary of correlation coefficients describing relationships between environmental factors and species abundance. 78 79 80 81 82 TABLE VIII Summary of correlation coefficients describing therelationship between environmental factors and species per cent composition. TABLE IX Summary of correlation coefficients describing therelationship between environmental factors and the amount of emergence. TABLE XSummary of the correlation coefficients describing therelationship between environmental factors and thenumber of emergence peaks forvarious species. TABLE XI Summary of correlation coefficients describing therelationship between environmental factors and the times of major emergence in several species. VIX TABLEXII Summary of correlation coefficients describing therelationship between larval abundance, numbers of emerging adults, number of emergence peaks and emergence time. 83 84 TABLE XIII The developmental rates of C. tentans in various lakes. TABLE XIV The percentage composition of species in the lakes based on the total adult emergence, May - August, 1970. 90 Inversion frequencies inchromosome of C. tentans. 1TABLE XV 116 Summary of correlation coefficients describing therelationship between the frequency of 1 Rad and some environmental factors. TABLE XVI 117 118 119 120 TABLE XVII Summary of correlation coefficients describing therelationship between the frequency of 1 Rad andthe abundance of some chironomids. TABLE XVIII Summary of correlation coefficients describing therelationship between the frequency of 1 Rad and the per cent composition of some chironomids, Summary of correlation coefficients describing the relationship between the frequency of 1 Rad and some emergence variables. TABLE XIX v i i i LIST OF FIGURES Page 5FIGURE FIGURE 12 The study area; Springhouse region. The study area; water bodies in the Chilcotin region. 6FIGURE FIGURE 34Details of the emergence trap. 22 Daily temperature range in some of the lakes where Chironomus tentans is <ul style="display: flex;"><li style="flex:1">abundant. </li><li style="flex:1">28 </li></ul>29 FIGURE FIGURE 56 Cumulative day degrees measured at the mud surface at a depth of 1 meter in some of the lakes where Chironomus tentans is abundant. The emergence of adults of Procladius bellus (Loew) and Procladius freemani Sublette from the one meter depth zone of several lakes. 35 37 The emergence of adults of Procladius dentus Roback from the one meter depth zone of several lakes. FIGURE FIGURE 78The emergence of adults of Procladius clavus Roback and Ablabesmyia peleensis (Whalley) from the one meter depth zone of several lakes. 40 43 45 The emergence of adults of Cricotopus flavibasis Malloch and Cricotopus albanus Curran from the one meter depth zone of several lakes. FIGURE 9 The emergence of adults of Psectrocladius barbimanus (Edwards) from the one meter depth zone of several lakes. FIGURE 10 FIGURE 11 The emergence of adults of Crypto- tendipes ariel (Sublette) and Calopsectra gracilenta (Holmgren) from the one meter depth zone of several lakes. 47 ix FIGURE 12 Larval abundance and adult emergence of Perotanypus alaskensis (Malloch) in L. Lye, Boitano L. and L. Jackson. 50 51 55 FIGURE 13 Larval abundance and adult emergence of Derotanypus alaskensis (Malloch) in Rock L., Sorenson L. and East L. FIGURE 14 Larval abundance and adult emergence of Einfeldia pagana Meigen in L. Jackson, Rock L. and Westwick L. FIGURE 15 Larval abundance and adult emergence of Einfeldia pagana Meigen in Near Opposite Crescent, Barkley L. and East L. 56 FIGURE 16 Larval abundance and adult emergence of Glyptotendipes barbipes (Staeger) in L. Jackson, Westwick L. and Sorenson L. 58 59 62 63 66 67 68 FIGURE 17 Larval abundance and adult emergence of Glyptotendipes barbipes (Staeger) in Rock L., Barkley L. and East L. FIGURE 18 Larval abundance and adult emergence of Chironomus anthracinus Zetterstedt in Boitano L., L. Jackson and Rock L. Larval abundance and adult emergence of Chironomus anthracinus Zetterstedt in Sorenson L., Barkley L. and East L. FIGURE 19 FIGURE 20 Larval abundance and adult emergence of Chironomus n.sp. in Barnes L., Boitano L. and L. Jackson FIGURE 21 Larval abundance and adult emergence of Chironomus n.sp. in Rock L. and Sorenson L. Larval abundance and adult emergence of Chironomus n.sp. in Barkley L. and East L. FIGURE 22 FIGURE 23 Larval abundance and adult emergence of Chironomus tentans Fabricius in L. Jackson, Westwick L. and Sorenson L. 74 FIGURE 24 Larval abundance and adult emergence of Chironomus tentans Fabricius in Rock L., Barkley L. and East L. FIGURE 25 Chironomid larval biomass and index of diversity forthe larval complexes at 1.0 m in the lake series FIGURE 26 Graph showing the relationship between .oxygen levels and organic carbon in the lakes. FIGURE 27 Salinity tolerances of the identified species of the one meter depth zone in the lake series. FIGURE 28 Examples of the spacing of emergence times of Chironomus tentans and three coexisting species. FIGURE 29 Seasonal variation in the frequencies of inversions of chromosome 1. LIST OF PLATES Page <ul style="display: flex;"><li style="flex:1">11 </li><li style="flex:1">Box 27 </li></ul>PLATE 1 A BBox 27;vegetation 12 13 14 15 16 21 PLATE 2 A BBarkley L. Barkley L.; Myriophyllum PLATE 3 A BNear Phalarope Near Opposite Crescent PLATE 4 A BL. Greer L. Jackson L. Lye PLATE 5 A BRound-up L. Barnes L. PLATE 6 A BBarnes L.; precipitated salts The emergence trap PLATE 7x i i ACKNOWLEDGEMENTS It is apleasure toexpress my gratitude to Professor G.G.E. Scudder who, asmy research supervisor, guided me through this work. The time and energy he spent aremuch appreciated. Dr. T.G. Northcote 1scriticism was invaluable during the also thank Dr. A.B. Acton for reading writing of thethesis. Ithe manuscript. Dr.M.S. Topping, whose PhD. thesis served as the basis for thepresent study, isespecially thanked forhis enthusiasm, support andpermission tousemuch ofhis unpub- lished data. Glen Jamieson, Tony Dixon andKen Bowler somehow put up with my innumerable questions about computer programming. Without their help thedata analysis would have resembled an infinite loop, orworse, would have crashed thesystem. Julian Reynolds, inbetween snickers, did a l l sorts of things to help. Iam indebted toDr. J.E.Sublette (Eastern New Mexico University) andDr. D.R. Oliver (Canada Agriculture, Ottawa) for their help with thedeterminations. Dr.A.M. Hutson (British Museum: Natural History) kindly supplied authentic specimens of Einfeldia pagana andPsectrocladius barbimanus for identification purposes. The research was carried outwhile inreceipt ofa National Research Council of Canada Postgraduate Scholarship and was further aided through an NRC grant toDr. Scudder. ERRATA 1. Where "Aphanozomenon" appears, read "Aphanizomenon". 2. Page 26, l i n e 18. " 3 1 ° i n L. J a c k s o n " s h o u l d read <ul style="display: flex;"><li style="flex:1">" 3 1 ° </li><li style="flex:1">i n B a r k l e y L . " . </li></ul>3. Page 61, l i n e 15. " u n i v o l t i v e " s h o u l d read " u n i v o l t i n e " . 4. Page 88, l i n e 17. "12000 umho/cm". 5. Page 1 0 2 , l i n e 18. " n o t r e a l t r e n d " s h o u l d read "no r e a l t r e n d " . 6. Page 126, l i n e 2. " e i g h t e e n s p e c i e s new to B . C . , t w e l v e s p e c i e s new t o Canada and seven s p e c i e s new t o s c i e n c e " . 1IINTRODUCTION This thesis is an ecological study of some of the Chironomidae inhabiting asaline lake series in the Chilcotin The chironomid complex of a saline lake series has never been thoroughly examined before. Rawson and Moore (1944) and Lauer (1969) and Cariboo regions of British Columbia. have mentioned chironomids in connection with work on saline waters, and others have recorded and studied various species in waters of differing salinities throughout the world (Remmert, 1955; Sutcliffe, 1960; Palmen, 1962; Bayly and Williams, 1966), but l i t t l e information has been gathered on how chironomid populations differ in a series of lakes of varying salinity. This type of study is particularly interesting since i t is well known that chironomids display extensive adaptation to <ul style="display: flex;"><li style="flex:1">a</li><li style="flex:1">wide variety of environments (Thienemann, 1954; Brundin, 1966) </li></ul>and are often able to thrive where many other animals cannot. The broad salinity tolerance of the Chironomidae gives them special prominence in saline habitats. This fact, in conjunction with their usual great abundance and wide diversity, makes chironomids useful organisms with which to study changes in the structure of species complexes that occur with variations in physical, chemical and biological conditions. The data obtained by Topping (1969) on this lake series in central British Columbia showed that in the dipteran 2Chironomus tentans Fabricius there was a significant correlation between the frequency of larval chromosome inversion 1 Rad and the total number of other chironomids present in the Since the selective value of inversions in wild pophabitat. ulations is not clearly understood, and as there have been few correlations of this sort, further investigation of this problem is considered valuable. Although Topping was able to show this correlation between inversion frequency and larval abundance, most emphasis was placed on the chemical composition of the environments and l i t t l e attention was directed to the biotic factors involved. Thus, one of the main aims of this thesis was to ascertain the effects of some biotic factors on the abundance of Chironomus tentans and the implications of these factors on the regulation of inversion frequency. In particular, i t was considered necessary to know more about the other chironomid species that coexist with C. tentans in this lake series, their numbers and their l i f e cycle characteristics. Only after determining the variations in the structure of the chironomid complex throughout the lake series is i tpossible to place the populations of C. tentans in proper perspective and to investigate the influence of the various species on C. tentans. I n i t i a l l y , attention is directed to the lake environments themselves; their physical and chemical characteristics are outlined. This information, in conjunction with extensive 3data collected on larval numbers and emergence patterns, is used in an examination of the major species of the one meter depth zone. Questions such as the effect of lake environments on the distribution and phenology of these species are discussed as are the possible interactions between the more dominant species present. The physical and chemical data are then integrated with information on chironomids and thus a description of a number of species associations is advanced. These associations, varying throughout the lake environments, form the basis for the examination of the relationship between C. tentans and the other species. With these data at hand the potential effect of the biotic factors on the chromosome inversion frequency in C. tentans may be more closely analyzed. Correlation coefficients are calculated to determine the types of interactions that may prove important In this respect. Aparticularly interesting problem that arises from the study of interspecific interactions is the apparent coexistence with C. tentans of two other very similar Chironomus species, C. anthracinus Zetterstedt and C^n.sp. (near a t r i t i b i a Malloch). Attention is focussed on this congeneric interaction and competitive exclusion (Hardin, 1960) is discussed in this context. 4II THE LAKE ENVIRONMENTS A. THE STUDY AREA The study was undertaken intheCariboo and Chilcotin areas of central British Columbia. The fifteen water bodies examined are situated intwo distinct butadjacent areas: the Springhouse area southwest ofWilliams Lake east ofthe Fraser River; and Becher's Prairie near Riske Creek onthe western (Chilcotin) side of theFraser (Figs. 1, 2). Those named as lakes canbe found on maps while theothers have <ul style="display: flex;"><li style="flex:1">names used for theconvenience of zoologists. </li><li style="flex:1">The water bodies </li></ul>include: a) Springhouse area: Sorenson Lake, Westwick Lake <ul style="display: flex;"><li style="flex:1">and Boitano Lake </li><li style="flex:1">b) Becher's Prairie (Chilcotin Area): Barnes </li></ul>Lake (Box4), Round-up Lake (Phalarope), Lake Lye (Box 20-21), Lake Jackson (Near Opposite Box 4), Lake Greer (Box89), Rock Lake, Near Phalarope, Near Opposite Crescent, Box 17, Barkley Lake (Opposite Box 4), East Lake (Racetrack) and Box27. FIGURE 1 The Study Area: The three lakes in the Springhouse region. Insets :The Fraser Plateau and i t s location in the province of British Columbia. 5

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