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 (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: (Atherinidae, 22 species), Poblana (Atherinidae, 3 species), Algansea (, 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 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 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 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" . 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.

2) Representatives of 12 of the 18 genera comprising the family Goodeidae have been cleared and stained in preparation for osteological studies.

3) Pregnant females of the goodeid genera Xenotoca, Neotoca, Allotoca and Goodea have been examined in d preliminary survey of the structure of the ovary.

4) Representative specimens of the genera Algansea, Xystrosus and Falcularius have been examined and preliminary data collected and tabulated.

5) A description of a new species of Algansea is in manuscript form.

6) The type specimens of Xystrosus popoche, Falcularius cha alae, Algansea dugesi, A. rubescens, A. stigmatura and A. affinis have been examined.

Proposed Studies

1) The Family Goodeidae

The distribution of the species within the Goodeidae reflects the known geological history of central Mexico. These fishes are widespread throughout the lakes and streams of the Lerma Santiago basin. They also occur in the Armeria and Ameca rivers, in a few smaller drainages between the Ameca and the Mezquital rivers and in the headwaters of the Balsas and Panuco rivers. As previously stated, one species is isolated in the head- waters of the Mezquital River in the state of Durango. This group is unique among Mexican cyprinodonts in that the species are viviparous. -4- The last revision of the family was that of Hubbs and Turner (1939). A history of the taxonomy of the group prior to that date can be found in their paper. The characters used by these authors included not only the usual counts and measurements, the degree of folding and length of the intestine, the morphology of the teeth and the strength of the ligamentous connection between the dentaries, but also the structure of the ovary and the morphology of the rectal processes of the embryos.

During the course of his studies on viviparity in fishes, C. L. Turner discovered that the ovaries of goodeid fishes were variable in structure and gave promise of being a valuable taxonomic tool. The organ itself is elliptical in shape and is divided longitudinally by a septum. This membrane may be complete in some species or incomplete. In the latter situation the dorsal and ventral septa may meet and intertwine in various ways but are not fused. The disposition of ovigerous tissue within the ovary is also variable. Turner also found filaments of tissue protruding from around the anal openings of the embryos within the ovaries. These structures, presumably nutritive in function, were variable in morphology and suggested a taxonomic use.

This early work (1932-1937) on the internal anatomy of goodeids was incorporated into the 1939 revision of the family. As a result of this emphasis on these reproductive characters, Hubbs and Turner's classifi- cation was considerably different from those previously used, and consisted of four subfamilies, 18 genera and 24 species. Preliminary examination of species represented in the Tulane Ichthyological Collection together with a reexamination of the data presented by Hubbs and Turner has led me to the conclusion that there are more names in this revision than can reason- ably be justified.

As a result of a personal communication with Dr. R. R. Miller and the dissection of Tulane specimens, it has become evident that the structure of the ovary may not be as reliable a taxonomic character as had previously been thought. The fact that the goodeid ovary is a compound structure con- sisting of fused right and left organs introduces a variable not usually associated with "conservative reproductive characters." The extent of variation must be determined before it can be used as a basis for taxonomic conclusions.

In a recently published paper, Mendoza (1965) has found considerable variation in the structure of the ovary of "Characodon" eiseni. Although he is unwilling to extrapolate his findings based on one species to in- clude the whole family, he suggests that a critical review of this charact- er is needed.

A brief survey of the genera recognized by Hubbs and Turner reveals that consolidation is definitely in order. Indeed, the closeness of some species groups is acknowledged by these authors. The subfamily Girardin- ichthyinae is a case in point. Based on characters described by Hubbs and Turner, Ilyodon and Balsadichthys would seem to be congeneric. Likewise, Skiffia, 011entodon and Neotoca. and Lermichthys also seem to be the same. The relationship of Hubbsina turneri to the last mentioned grouping remains at the present time unclear.

The largest subfamily, the , seems to be a miscellaneous collection of unrelated morphological types. As ovarian structure and other characters vary from genus to genus, there seems to be, at present, no al- ternative to a large number of species groups. Careful analysis of large

-5-- series of specimens may, however, reveal clues to relationships which have been overlooked in the past. Relationships which cut across the lines of the present classification may also be discovered.

The two remaining subfamilies, the Ataeniobiinae and Characodontinae, are both monotypic. Ataeniobius toweri is considered to be the most primi- tive goodeid because the embryos lack nutritive rectal processes. Characo- don lateralis was separated from species formerly placed in this genus on the basis of the structure of ovary and rectal processes. The relationships of both of these species will be reassessed.

It is the principal investigator's intention to arrive at a consist- ent and satisfactory classification of the family Goodeidae based not only on a reevaluation of known characters, but also on new studies.

Plan of Study

I External Morphology 1) Counts and measurements commonly used in ichthyological studies will be made on a large series of each species throughout their ranges to determine the limits of variation of these characters. At present there is almost no data of this type available. The methods of Hubbs and Lagler (1958), Gosline (1949) and Hubbs and Turner (op. cit.) will be followed. Limits of variation in the striking color patterns of these fishes will also be determined. An attempt will also be made to discover new characters.

2) Wherever possible, all size classes of each species will be examined for possible ontogenetic changes in characters, (i._ e._ teeth). II Internal Morphology 1) Large series of ovaries of each species will be grossly exa- mined according to the method of Mendoza (22. cit.) in an effort to establish the limits of variation of this character. Histological investigations will be initiated at a later date if deemed necessary.

2) Possible variation in the morphology of the embryonic anal processes (trophotaeniae) will be investigated according to the methods followed by Turner (1933, 1937) and Mendoza (1937, op: 3) The osteology of the family will be described.

While the approach outlined above is primarily that of "classical morphology," it is my firm conviction that the taxonomy and phylogeny of these fishes will never be accurately worked out until such basic pieces Jf information as the variation in the number of median lateral scales is determined.

2) The Genus Algansea

The 11 nominal species, including two undescribed forms, comprising the genus Al ansea are fine-scaled fishes which, like the goodeids, are widely distributed throughout the Lerma-Santiago river system. They range in size from about 3 inches to 15 inches in length.

-6- Beyond inclusion in standard ichthyological works (Jordan and Ever- mann, 1896; Meek, op. cit., Regan, op. cit., De Buen, 1946; Alvarez, 1950) the group has been generally neglected. As a result of a preliminary study, some interesting taxonomic relationships have come to light. Preliminary data collected from specimens housed in the Museum of Zoology of the Uni- versity of Michigan and Tulane University suggest that the genus can be divided into three species groups (Table 1).

The two undescribed species and A. barbata are small fishes which in- habit swift, headwater streams. Algansea affinis, A. dugesi and A. paratin- cella are found in various parts of the lower reaches of the Lerma River, some of the lakes and in the Santiago River below Lake Chapala. The system- atic status of all of these species is unclear as is their relationship to A. tincella which is found in the Valley of Mexico.

The species that are found predominantly in the lakes, and probably evolved in lacustrine habitats, include A. rubescens from Lake Chapala and A. lacustris (=A. tascorum) from Lake Pitzcuaro. The former species was described by Meik (1902) from the Santiago River just below the outlet of Lake Chapala at Ocotlan. As can be seen in the table, specimens from this population occasionally yield aberrant gill raker counts. It is suggested that these individuals represent hybrids with Xystrosus popoche (see below). Until the extent of the supposed hybridization is known, the status of the species cannot be resolved.

A thorough study of the genus based on a study of the osteology of all species as well as their meristic and morphometric characters is planned. Also, the relationship of Algansea to the other Mexican cyprinids will be determined.

3) The Genus Xystrosus

The monotypic genus Xystrosus is found only in Lake Chapala and the Santiago River above the falls. It is separated from Algansea, to which it is closely related, by the upturned position of the mouth in adults and the high number of gill rakers (Table 2). The fact that young Xystrosus popoche are impossible to distinguish from Algansea rubescens without counting rakers suggests that separation of these two genera is not warrant- ed. A similar situation is found in the subspecies of Siphateles bicolor, a found in the Lahontan system of Nevada and California (Snyder, 1917; Hubbs and Miller, 1943, 1948).

It is proposed to establish the systematic status of Xystrosus by means of the examination of further specimens from Lake Chapala and the Santiago River.

4) The Genus Falcularius

Like Xystrosus, Falcularius is a monotypic genus whose one species, X. chapalae, inhabits Lake Chapala and the Santiago River above the falls. It is closely related to Hybopsis alta and again the same problem occurs: the two genera are separated only with great difficulty. The status of this genus will be resolved by a meristic and morphometric study of the two species throughout their ranges.

-7- 7. Facilities.-

The Principal Investigator is a full-time, salaried member of the regular faculty of the University of Utah, a state institution with an enrollment of about 17,000 students. He has free use of the facilities in the Department of Zoology and the library.

The university has contributed materially to my research by providing two rooms, a total of approximately 710 square feet, for a collection and work area. This space is shared with two graduate students. Approximately one thousand dollars has been spent for tanks, cabinets, nylon nets and a dissecting microscope. Shelving is adequate for the space available.

With the completion of a new library scheduled for the fall of 1968, present plans call for the renovation of the old library building as a natural history museum. Approximately 1700 square feet will be devoted to ichthyological research.

8. Personnel.-

Principal Investigator (Clyde D. Barbour)

General.- The applicant is an ichthyologist who recently received his Ph.D. from Tulane University where he studied under Dr. Royal D. Suttkus. During his graduate career, he was occupied almost exclusively with a systematic study of the genus Chirostoma and with laying the groundwork for future studies on the fishes of the Mexican plateau. He has carried out extensive field work in Mexico.

Vital Statistics.- Born October 30, 1935, New York City; married in 1962 Caroline Andler of Newport, New Hampshire; one child.

Education.- Kent School, Kent, Connecticut, 1949-1954; College of William and Mary, Williamsburg, Virginia, 1954-1956; B. A. degree Stanford University, Stanford, California, June, 1958; Graduate School, Tulane University, New Orleans, Louisiana, 1961 to 1966; Ph.D. degree September, 1966.

Zoological Experience.- Laboratory Assistant, Virginia Fisheries Labora- tory, Gloucester Point, Virginia, summer 1956; Enlisted Scientific Per- sonnel, U. S. Army Medical Research Laboratories, Army Chemical Center, Edgewood, Maryland, 1958 to 1960; Laboratory Technician, Biophysical Laboratories, Harvard Medical School, Boston, Massachusetts, 1961. Museum Technician, Division of Fishes, U. S. National Museum, Smithsonian Insti- tution, Washington, D. C., summer 1962; Teaching Assistantships, Tulane University (General Botany, Animal Biology, Comparative Invertebrate Biology, Comparative Anatomy, Vertebrate Natural History), four years. Participant, Tulane Environmental Training Program, summer 1966.

Present Position.- University of Utah: Assistant Professor, Department of Zoology and Entomology, Curator of Fishes.

Address.- Department of Zoology and Entomolog, University of Utah, Salt Lake City, Utah 84112; or home address, 887 Third Avenue, Salt Lake City.

Telephones.- University of Utah, 322-7211; Department of Zoology and En- tomology, 322-7586; Barbour [office]322-7962, [home] 355-6547. Area Code 801. -8- Other Personnel and Curatorial Assistant

The field assistant and curatorial assistant will be graduate students from the Department of Zoology and Entomology.

9. Bibliography.-

1967 Some aspects of the ecology of the Mediterranean Gecko, Hemi- dactylus turcicus turcicus in New Orleans, Louisiana. (with Francis L. Rose) Submitted to The American Midland Naturalist.

Manuscripts in preparation:

Al ansea , a new cyprinid from central Mexico (with a va or Contreras).

The status of Moxostoma congestum (Catostomidae) in the Santiago River system of central Mexico.

The distribution, variation and osteology of Chirostoma jordani jordani (Woolman) and C. jordani mezquital Meek (Atherinidae).

Chirostoma humboldtianum avium, a new subspecies of the family Ather- iiiidae from western Mexico.

Chirostoma aculeatum, a new species of the family Atherinidae from the Mesa Central of Mexico.

The last three manuscripts in the above list represent portions of my Ph.D. dissertation. Submission of this material for publication will commence this spring when less time will be taken up with writing lec- tures and the preparation of classwork. Other parts of the disserta- tion will be published as soon as gaps in the known distribution of certain species are filled.

Literature cited. -

Alvarez, J. 1950. Claves para la determinaciOn de especies en los peces de las aguas continentales Mexicanas. Sec. Marina, Dir. Gral. Pesea e Indust. Conex. Mexico.

. 1966. ContribuciOn al conocimiento de los Bagres fOsiles de Chapala y Zacoalco, , Mexico. paleoeco1ogia/1, Dept. de Prehist., Inst. Nac. Antro. e Hist. Mexico.

Barbour, C. D. 1966. The systematics and evolution of the genus Chirostoma (Pisces; Atherinidae). Ph.D. dissertation, Tulane University, vi + 233 pp.

Clements, T. 1963. Pleistocene history of Lake Chapala, Jalisco, Mexico. In: Essays in Marine Geology in Honor of K. O. Emery. T. Clements, ed., pp. 35-49. Univ. Southern Calif. Press, Los Angeles.

De Buen, F. 1946. Ictiograffa Continental Mexicana (I, II, III). Rev. Soc. Mex. Hist. Nat. VII (1-4):87-138.

-9- Gosline, W. A. 1949. The sensory canals of the head in some cyprinodont fishes, with particular reference to the genus Fundulus. Occ. Pap. Mus. Zool., Univ. Mich. No. 519, 21 pp.

Hubbs, C. L. and K. F. Lagler. 1947. Fishes of the Great Lakes Region. Cranbrook Inst. Sci., Bull. 26.

and R. R. Miller. 1943. Mass hybridization between two genera of cyprinid fishes in the Mohave Desert, California. Pap. Mich. Acad. Sci. 28:343-378.

and . 1948. The Great Basin with Emphasis on Glacial and Postglacial Times. II. The Zoological Evidence. Correlation between fish distribution and hydrographic history in the desert basins of western United States. Bull. Univ. Utah 38:18-166.

and C. L. Turner. 1939. Studies of the fishes of the order Cyprinodontes. XVI. A revision of the Goodeidae. Misc. Pub. Mus. Zool., Univ. Mich. No. 42, 80 pp.

Jordan, D. S. and B. W. Evermann. 1896-1900. The fishes of north and middle America. Bull. U. S. Nat. Mus. 47(1-4):1-3313.

Meek, S. E. 1902. A contribution to the of Mexico. Field Col. Mus. Pub. 65, 3(6):63-128.

. 1904. The fresh-water fishes north of the Isthmus of Tehuantepec. Field Col. Mus. Pub. 93, 5:1xiii + 252 pp.

Mendoza, G. 1937. Structural and vascular changes accompanying the resorption of the proctodaeal processes after birth in the embryos of the Goodeidae, a family of viviparous fishes. Jour. Morph., 61:95-125.

. 1965. The ovary and anal processes of "Characodon" eiseni, a viviparous cyprinodont teleost from Mexico. Biol. Bull. 129(2):303-315.

Regan, C. T. 1906-1908. Pisces. In: Biologia Centrali-Americana. 8:1-203.

Snyder, J. O. 1917. The fishes of the Lahonton system of Nevada and northeastern California. Bull. Bur. Fish. 35,1915:31-86.

Stuart, L. C. 1964. Fauna of Middle America. In: Handbook of Middle American Indians, Vol. 1. R. Wauchope and R. C. West, eds., pp. 316-362. Univ. Texas Press, Austin.

Turner, C. L. 1932. Unique reproductive features in killifishes of the Mexican Plateau. Anat. Rec. 54:113.

. 1933a. The unique nutritional organs in the embryos of the top of the Mexican Plateau. Science 77:93-94.

. 1933b. Nutritive processes in the embryos of the Goodeidae, viviparous top minnows of Mexico. Anat. Rec. 57:77. -10- . 1933c. Viviparity superimposed upon ovoviviparity in the Goodeidae, a family of cyprinodont teleost fishes of the Mexican Plateau. Jour. Morph. 55:207-51.

. 1935. The use of the embryonic rectal processes of the em- bryos in a revision of the classification of the family Goodeidae. Anat. Rec. 64(Supp.1):137-38.

. 1937a. The trophotaeniae of the Goodeidae, a family of viviparous cyprinodont fishes. In: Wistar Inst. Abs. :4.

. 1937b. The trophotaeniae of the Goodeidae, a family of viviparous cyprinodont fishes. Jour. Morph. 61:495-523.

West, R. C. 1964. Surface configuration and associated geology of Middle America. In: Handbook of Middle American Indians, Vol. 1. R. Wauchope and R. C. West, eds., pp. 33-83. Univ. Texas Press, Austin. TABLE 1. TOTAL NUMBER OF GILL RAKERS ON THE FIRST ARCH OF REPRESENTATIVE SPECIES

OF THE GENUS ALGANSEA

SPECIES 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32

A. sp: nov. Barbour 9 70 77 64 12 2

A. sp. nov. Miller 4 14 19 4 1

A. barbata 1 1 7 13 7 1 1 A. affinis 2 12 27 28 17 7 2

A. dugesi 1

A. paratincella 2 4 6 2

A. rubescens 1 1 3 2 2 2 (Sayula Basin)

A. rubescens 2 8 3 1 2 2 3 2 2 (Santiago River)

A. lacustris 2 3 2 3 1 TABLE 2. TOTAL NUMBER OF GILL RAKERS ON THE FIRST ARCH

OF XYSTROSUS POPOCHE

41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

1 1 1 3 3 1 6 3 4 4 2 1 2 2 2 2 2 Budget 1967-1968

% Time Salary Contribution This Academic of Project Year NSF U of U Total

1. Personnel

Principal Investigator 2 months (summer) 100 8700 1934 0 2153 Academic year 2.5 8700 0 220 220

Curatorial assistant 9 months 50 1080 0 1080

Salaries and wages charged to this project 3014 220 3234

2. Employee Benefits 301 22 323

3. Indirect Costs, 44% of wages and salaries 1326 97 1423

Travel (740) 36E3- Automotive expenses, 4CFCgt mi. ataR per mile 266 250 Living expenses, 20 days 140 140 One scientific meeting 300 300 (New York)

Equipment (1500)

Canoe (aluminum) 300 300 Uild stero microscope 1200 1200

Supplies (1000)

Supplies 1000 1000

Total expenses for 1967-1968 7831 339 8170

-14- Budget 1968-1969

% Time Salary Contribution This Academic of Project Year NSF U of U Total

1. Personnel

Principal Investigator 3 months (spring quarter) (Summer salary not requested) 100 9200 1533 1534 3067 Field assistant 3 months 100 900 0 900 Curatorial assistant 50 1080 0 1080 Salaries and wages charged to this project 3513 1534 5447 2. Employee Benefits 351 153 504 3. Indirect Costs, 44% of wages and salaries 1546 675 2221

Travel (3300) Automotive expenses, 15,000 mi. at 1 04 per mile 1500 1500 Living expenses, 90 days in the field 1500 1500 One scientific meeting (Ann Arbor) 300 300

Supplies (300) Supplies 300 300

Total expenses for 1968-1969 9010 2362 11372 Total budgeted expenditures 1967-1969 16841 2701 19542

Total percentage, cost participation, University of Utah: 5.2

-15- 10. Explanation of Budget.-

Salaries and Wages.- I plan to devote full time to this project during the summers of 1968 and 1969. Salary requested for the summer of 1968 is intended as summer salary and does not exceed the allowable 2/9. Salary is also requested for March, April and May of 1969 when a leave of absence will be taken from the University to collect in Mexico. This trip is scheduled to fill the gaps in collections presently in museums. Past experience has shown that it is pointless to try and collect on the Mesa Central during the summer months (the rainy season).

Wages are also requested for a field assistant and a curatorial assistant.

Travel.- Budgeted funds are chiefly for a three-month trip to Mexico during the spring of 1969. Mileage is estimated realistically for a four- wheel drive vehicle leased from the University at a set rate (10 cents per mile or $85.00 per month, whichever is greater).

Funds are also proposed for a trip to New Orleans in early September or December 1967 to examine and pick up the bulk of the collection of goodeids and cyprinids collected by myself during my graduate career.

Living expenses include food, lodging, field supplies and maintenance of camping gear, occasional purchase of specimens, insurance, documents and visas, and occasional procural of guides, pack animals, and native labor.

Supplies and Expenses.- Supply funds will be used chiefly for cura- torial supplies-ITanks, jars, preservative) and camping gear. Funds are also requested for a Wild M-5 microscope with drawing tube, lamp, swinging arm stand and accessory oculars. At present only one adequate dissecting microscope is available for myself and my two students due to increased en- rollment at the graduate level. This new microscope will make another in- strument available and greatly facilitate osteological study of cleared and stained specimens.

11. Other Sponsors.-

This proposal has not been submitted to any other granting agency. In the event that the proposal is unsuccessful, it may be submitted to another agency or, in an altered form, again to the National Science Found- ation.

Summary

Funds in the amount of $19,542 are requested for a study of the taxonomy and evolution of the family Goodeidae and the genera Algansea, Xystrosus and Falcularius, fishes which are endemic to the drainages of the Mesa Central of Mexico and contiguous waters. Knowledge of the ich- thyofauna of this interesting zoogeographic region has been hampered in the past because of a lack of specimens. Utilization of new material now available in collections coupled with intelligent field work to specifi- cally fill in gaps in these collections will make this investigation a highly fruitful one.

-16- Submitted by: Approved by:

C. D. Barbour, Assistant Professor Department of Zoology and Entomology

W. S. Partridge, Director, Office of Research Administration

A. C. Emery, Academic Vice President

March 15, 1967

-17-