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In the Pecos River, Texas, and New Mexico

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In the Pecos River, Texas, and New Mexico GENETIC DIFFERENTIATION AMONG SUBPOPULATIONS OF THREE GAMBUSIA SPECIES (PISCES: POECILIMAE) IN THE PECOS RIVER, TEXAS, AND NEW MEXICO A Thesis Submitted to the Faculty of Baylor University In Partial Fulfillment of the Requirements for the Degree of Master of Science By Elisabeth Milstead Waco, Texas August, 1980 ABSTRACT Heterogeneity among subpopulations of three species of Gambusia was determined by electrophoretic analysis of allelic frequencies. The analysis included (1) all populations of G. nobilis, an endemic of the Pecos River drainage of New Mexico and Texas which occurs only in four widely separated spring systems; (2) five populations of G. affinis, a widely distributed species with a more continuous distribution in the Pecos drainage; and (3) four populations of G. geiseri, an endemic from the east side of the Edwards Plateau which has been introduced into three springs of the Pecos drainage on the west side of the Plateau. The average FsT values (Wright's standardized variance in allelic frequency) for Pecos River populations within the range of G. nobilis were 0.755 for G. nobilis, 0.160 for G. affinis, and 0.278 for G. geiseri. The relatively high value for G. geiseri compared to G. affinis may result from founders effect. Even when populations of G. geiseri and G. affinis from the opposite side of the Edwards Plateau from the Pecos are consid- ered, the heterogeneity of G. nobilis remains higher than that of the other two. The same result remains when a population of a second sub- species of G. affinis is included in the analysis. The average FsT value for G. nobilis is, in fact, higher than that previously reported for any species. In addition, of the three species of Gambusia, G. nobilis showed the highest divergence on the basis of overall genetic identity (Nei's measure, 16-17 loci). The data support the classical concept that gene flow exerts a dedifferentiating effect on subpopulations. The un- iii usually high genetic variance observed for G. nobilis may be character- istic of fishes restricted to desert springs. iv TABLE OF CONTENTS ABSTRACT ........................................................................................................................................... iii LIST OF TABLES ............................................................................................................................ vi LIST OF FIGURES .......................................................................................................................... vii ACKNOWLEDGEMENTS ........................................................................................................................ viii INTRODUCTION ..........1 , DISTRIBUTION AND HISTORY OF GAMBUSIA IN THE PECOS RIVER ..........14 MATERIALS AND METHODS ..........8 RESULTS .............................................................................................................................................. 11 DISCUSSION ...................................................................................................................................... 20 LITERATURE CITED ........................................................................................................................ 27 APPENDIX ........................................................................................................................................... 32 LIST OF TABLES Table Page 1. Number of alleles per locus in three species of Gambusia ... 9 2. Allelic frequencies at four loci in four populations of G. nobilis ..................................... 12 3. Allelic frequencies at eight loci in five populations of G. affinis ..................................... 13 4. Allelic frequencies at five loci in four populations of G. geiseri ..................................... 14 5. F and heterogeneity Chi-square values for polymorphic loci inST various combinations of populations of G. nobilis ...... 15 6. F and heterogeneity Chi-square values for polymorphic loci inST various combinations of populations of G. affinis ...... 16 7. F and heterogeneity Chi-square values for polymorphic loci inST various combinations of populations of G. geiseri ...... 17 8. Average FsT values for various fish populations ............ 23 vi LIST OF FIGURES Figure Page 1. Localities in the Pecos River drainage 5 vii ACKNOWLEDGEMENTS I would like to thank Drs. Anthony A. Echelle, Robert C. Gardner, and O. T. Hayward for the time and effort spent in serving on my committee and critically reviewing the manuscripts. I am especially grateful to Dr. Echelle for his advice and help during all phases of this study. Alice Echelle and D. Allen Rutherford offered invaluable assistance in making collections and executing the laboratory work. David Hillis and Doyle Mosier also assisted in making collections. Dr. John Patton, Dr. Clark Hubbs, Ken Asbury, Jim Duke, Mike W. Smith, and Mike Dean provided information and discussion on various aspects of this study. Jim Johnson of the U. S. Fish and Wildlife Service in Albuquerque was instrumental in the initiation of this study. Financial assistance was provided by the U. S. Fish and Wildlife Service, contract No. 14-16-0002-79-133, Sigma Xi, and the Baylor University Graduate School. The bulk of this manuscript was prepared while I was a student at the University of Oklahoma Biological Station. viii INTRODUCTION Fishes in desert river systems of North America often exhibit a dichotomy regarding opportunity for gene flow between subpopulations. Some species occur only in small, relatively stenothermal, springfed habi- tats which are discontinuously distributed. The various populations are therefore separated by large areas that are uninhabitable by the species. In such cases, the populations may have been effectively isolated from each other since pluvial periods of the Pleistocene (e.g., Miller, 1948). Other species thrive in situations that show great fluctuations in physi- cal and chemical features of the environment. Consequently, species of the latter group are more continuously distributed in tributaries and major streams, and should exhibit relatively high levels of gene flow between subpopulations. Good examples of this dichotomy include poeciliid fishes of the genus Gambusia in the Chihuahuan desert of the United States and Mexico (Hubbs and Springer, 1957). In this paper I compare electrophoretically detected genetic hetero- geneity in two species of Gambusia in the Pecos River system of Texas and New Mexico--one of which, G. nobilis, is a spring dweller, while the other, G. affinis, occurs in the same spring systems and yet is also widely dis- tributed in the Pecos River and its major tributaries. I also present an analysis of genetic heterogeneity in G. geiseri, an introduced (Hubbs, 1980) spring-dweller of the Pecos River system. Recently, there has been a number of reports of genetic heterogeneity among subpopulations of stream dwelling fishes. These include studies by 1 2 Avise and Felley (1979) and Avise and Smith (1974) on L2pomis macrochirus, Echelle et al. (1975, 1976) on two species of Etheostoma, Koehn (1970) on catostomids, Koehn et al. (1971) on Notropis stramineus, Merritt et al. (1978) on Rhinichthys cataractae, and Ryman et al. (1979) on Salmo trutta. Studies of populational variation in Gambusia consist of brief treatment of G. affinis in small local areas of central Texas and South Carolina, respectively, by Yardley and Hubbs (1976) and Yardley et al. (1974). Studies dealing with genetic variation in desert fishes include Turner's (1974) work on desert pupfishes and Koehn's (1970) studies of esterase variation in catostomids. However, Turner's study was not directly concerned with intraspecific populational variation, and Koehn's dealt only with esterase heterogeneity. To date there have been no comparisons of the genetic structure of desert species which exhibit the above described dichotomy in opportunity for gene flow. Contrary to the conclusions of Ehrlich and Raven (1969) and Endler (1977), it is intuitively apparent that gene flow should be a major cohesive evolutionary force contributing a dedifferentiating effect among subpopulations of a species. However, there have been few empirical tests in which proper controls were employed. In a brief review, Jackson and Pounds (1979) were able to cite only four studies with some controls (Metter and Pauken, 1969; Rees, 1970; Jackson, 1973; and Echelle et al., 1976), all of which support the concept that gene flow has a strong dedifferentiating effect on subpopulations of species. Thus, I began my study with the following prior hypotheses: (1) the strongly isolated populations of G. nobilis should dhow the greatest degree of differen- tiation; (2) because G. geiseri in the Pecos River represents a series of 3 recently introduced populations of a species with a highly restricted natural distribution, the various populations of G. seiseri should dhow the lowest level of differentiation; (3) even in the absence of natural selection (Rohlf and Schnell, 1971), differentiation in continuously distributed organisms may be expected to increase with geographic distance as a function of the "isolation by distance" phenomenon (Wright, 1943). Thus, subpopulations of G. affinis should dhow intermediate levels of differentiation within that segment of the Pecos River encompassed by the range of G. nobilis, i.e., greater differentiation than G. geiseri, lower than G. nobilis. DISTRIBUTION AND HISTORY OF GAMBUSIA IN THE PECOS RIVER G. nobilis, which is endemic to the
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