Proposal for Support of Research on the Systematics and Evolution of the Fishes Endemic to the Mesa Central of Mexico
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PROPOSAL FOR SUPPORT OF RESEARCH ON THE SYSTEMATICS AND EVOLUTION OF THE FISHES ENDEMIC TO THE MESA CENTRAL OF MEXICO To: National Science Foundation From; Clyde D. Barbour Systematic Biology University of Utah Washington 25, D. C. 1. Institution.- Department of Zoology and Entomology University of Utah Salt Lake City, Utah 84112 (Telephone: 322-7586) 2. Principal Investigator.- Clyde D. Barbour, Assistant Professor of Zoology and Curator of Fishes 3. Title of Project Proposed.- The Systematics and Evolution of the Fishes Endemic to the Mesa Central of Mexico 4. Desired Starting Date.- September 1, 1967 5. Period of Time for Which Support is Requested.- Two years, beginning September 1, 1967 and terminating August 31, 1969. 6. Description of Research Proposed.- Introduction The Central Plateau is the largest physiographic region of Mexico. According to West (1964) it "extends from the present United States-Mexico border southward to the latitude of Mexico City." This area is enclosed by mountain systems on the east, west and south. Elevations are the low- est at the international border (less than 1000 m.) and slowly increase as one progresses southward. At its southern extremity the plateau sur- face may rise to more than 2000 m. The Mesa Central of Mexico is defined as the southern one-third of the Central Plateau. This tropical highland extends south from about the latitude of San Luis Potosi and Aguas Calientes to the Transverse Volcanic Axis that marks the southern limit of the plateau (West, a.. cit.). This region is drained principally by the westward flowing Lerma-Santiago river system although other rivers have cut back into the edges of the plateau, _ e.,_ the Panuco, Balsas, Armeria, and the Ameca. The uplifted and isolated nature of the Mesa Central has resulted in the development of .a highly endemic fish fauna. The family Goodeidae (Cyprinodontoidei, about 30 species) and the following genera are largely restricted to the present or former Lerma-Santiago drainage and those rivers that have cut back into this portion of the plateau: Chirostoma (Atherinidae, 22 species), Poblana (Atherinidae, 3 species), Algansea (Cyprinidae, about 5 species), Xystrosus and Falcularius (Cyprinidae, both monotypic), Evarra (Cyprinidae, 2 species) and Aztecula (Cyprinidae, 4 species). A major exception to this uniform pattern of distribution is the small enclave of Lerma-Santiago fishes (one goodeid and one ather- inid) found in and around the headwaters of the Mezquital River in the State of Durango. Studies on the fishes of this region were initiated by myself in 1962. Since that time I have personally made 95 collections on the Mesa Central and have accumulated a considerable body of taxonomic information concerning the genus Chirostoma (Barbour, 1966). My own observations and a survey of the pertinent literature have revealed that there are still challenging taxonomic and evolutionary problems to be solved in the other unique groups. Field work by Dr. Robert R. Miller and his students at the University of Michigan and by myself while I was at Tulane University has resulted in the accumulation of series of specimens for many of the species where formerly there had existed, at worst, only the type or type series. These facts make a taxonomic study of this fauna not only desirable but also feasible. Besides Chirostoma, the genera Poblana, Evarra, and Aztecula are being omitted from this proposal. Poblana is presently under study by my student, Mr. Robert Bolland, as a master's problem. Evarra and Aztecula are closely allied to the large and complex genus Notropis. While an attempt will be made to make comprehensive collections of these cyprinids, formal study is deferred to a later date. It is the conviction of most plant and animal systematists and eco- logists that the rapid exploitation and depletion of the natural environ- ment lend a new urgency to their studies. Although there are scientists who would debate this point, it is my belief that studies concerning physics, chemistry, et cetera can be put off to a future date without great loss. Atoms and molecules will always be around to study. The half-life of our natural environment, particularly our terrestrial and fresh-water habitats, is critically short. In an age of steel and concrete it is a truism to say what is here today can very easily be totally and irreplaceably gone tomorrow. Nowhere is this statement more clearly demon- strated than on the dry Mesa Central. The diversion of parts of the Lerma River for cities and agriculture has reduced the living space of aquatic animals and pollution resulting from raw sewage and increasing industri- alization renders more and more the remaining waters unfit for life. The recent introduction of the largemouth bass, Micropterus salmoides, into certain isolated lakes and Lake Chapala itself will certainly take its toll of the native fauna. Indeed, one isolated population (out of three) of an undescribed subspecies of Chirostoma humboldtianum was already near extinction in 1963 because of this predator. As the area to be studied has undergone much volcanic activity in the recent past, many of the goodeids are highly restricted in their geographical distribution and may be considered to be endangered species. If the opportunity to study these fishes is not seized quickly, it may well be lost forever. Abstract The goal of the research described herein is to analyse systemati- cally the fishes peculiar to the Mesa Central. The research aims at the following specific objectives: 1) The family Goodeidae A. To ascertain the limits of specific variationiand so define the valid species. B. To reevaluate generic relationships. C. To determine, if possible, the position of the family within the suborder Cyprinodontoidei. -2- 2) The genus Algansea A. To ascertain the limits of specific variation and so define the valid species. B. To determine the relationship of this genus to other North American Cyprinidae. 3) The genera Falcularius and Xystrosus To determine the relationships of these monotypic genera to the other cyprinids of the Mesa Central. 4) To make a coherent and reasonable statement concerning the evo- lution of fishes on the Mesa Central of Mexico. Scope of the Problem Proposed The geographic scope of this project is limited by the distribution of the fishes under study. The fact that these distribution patterns have their center on the Mesa Central serves only to emphasize the highly en- demic nature of the fauna and does not reflect any arbitrary divisions on my part. The Lerma drainage and that part of the Santiago River above the waterfall near the city of Guadalajara have been designated a natural fish province of Middle America by Meek (1904), Regan (1907) and Stuart (1964). The philosophical approach to this problem is primarily taxonomic. Although much of my present thinking on the relationships of the fishes under consideration is based on morphological differences and similarities, I am aware that biological relationships are more than such morphological phenomena. It is for this reason that information on ecology and behavior is always sought in the field to augment other findings. I am also of the opinion that taxonomic schemes should reflect, whenever possible, evolutionary relationships. This emphasis on phylogeny inevitably leads to a general discussion of the problem of speciation and adaptive radiation as exemplified by the groups under study. In this regard, the proposed study transcends "straight" taxonomy. Wherever poss- ible, present distribution patterns will be correlated with past geologic events (Clements, 1963; Alvarez, 1966). My theory that character displace- ment is partially responsible for the diverse morphology found in the various species, particularly in the genus Chirostoma, will be investigated as will the idea that considerable evolutionary parallelism on the generic as well as on the specific level is also present. It is obvious that the research program described in this proposal cannot be completed in two years. At the end of this period the completed work will be evaluated and a new proposal submitted to the National Science Foundation or another granting agency. At the final conclusion of this study, it is hoped that a coherent statement regarding the taxonomy, distri- bution and evolution of the fishes of the Mesa Central can be made. Methods Fishes will be obtained chiefly by means of 10 ft. nylon common-sense seines. In the larger rivers and lakes, gill and trammel nets will be used to supplement the smaller seines. These large nets will be set from a canoe. -3- Size of specimens preserved is limited by the size of field con- tainers. For this reason, 3 and 5 gallon glass buckets are carried into the field as well as the standard 1/2 gallon jars. Ten percent formalin is used as the preservative. Detailed notes as well as photographs of color patterns will be made from live or freshly killed specimens in the field. If it is at all possible, a few specimens will be retained for observation in the laboratory. Basic data on habits and habitat which are routinely obtained are as follows: 1) general description and photograph of the habitat; 2) identification of the dominant plants in the habitat; 3) determination of width, flow, turbidity, temperature and bottom conditions of the aquatic habitat. The depth of capture and weather are also noted. Specific laboratory procedure to be followed is given below under the appropriate headings. Previous Work by_ the Principal Investigator The following progress has already been made toward the stated ob- jectives: 1) As a result of my dissertation work, I am familiar not only with the literature concerning the geology and the ichthyofauna of the Mesa Central, but have already spent a total of three months in the field. I have personally collected most of the species that I plan to study.