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March 16, 1973 CHIROSTOMA SWAINSON TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 18, Number 3 March 16, 1973 THE SYSTEMATICS AND EVOLUTION OF THE GENUS CHIROSTOMA SWAINSON (PISCES, ATHERINIDAE) CLYDE D. BARBOUR' Department of Biology, University of Utah, Salt Lake City, Utah 84112 ABSTRACT of C. jordani is noted. The evolution of Chirostoma is best explained by consider- The genus Chirostoma is composed of 18 ing the genus to be diphyletic; a Menidia- species and six subspecies not including C. like ancestor of the jordani group invaded compressum, thought to be extinct. Except the Mesa Central first, followed at a later for a few populations of C. ford ani near time by the Melaniris-like ancestor of the Durango City, Durango, it is restricted to arge group. The evolution of the species is the Mesa Central of Mexico where it makes summarized in a hypothetical phylogenetic up approximately 20 percent of the fish tree. fauna. The following names in current usage are herein synonymized: Otalia = Chiro- stoma; C. ocampoi, C. regani = C. hum- CONTENTS boldtianum; C. diazi = C. sphyraena; C. ocotlanae = C. /ucius. No subspecies of INTRODUCTION - - - - - - - - - - - - - - - - - 98 C. jordani are recognized. C. reseratum is METHODS - - - - - - - - - - - - - - - - - - - - - - 98 recognized as a subspecies of C. consocium. The name C. attenuatum Meek is reapplied ACKNOWLEDGMENTS - - - - - - - - - - - - - 99 to the populations in Lakes Patzcuaro and DIAGNOSTIC CHARACTERS - - - - - - - - 100 Zirahuen currently designated as C. bar- KEY TO SPECIES OF THE GENERA toni. Five individuals from the lake at Chirostoma AND Melaniris - - - - - - - 101 Balneario Cointzio, Michoacan are referred to C. charari De Buen. C. aculeatum is THE JORDAN! GROUP - - - - - - - - - - - - 102 described as new. Chirostoma jordani - - - - - - - - - - - - - - - 102 Chirostoma is divided into two species Chirostoma patzcuaro - - - - - - - - - - - - - 107 groups. The jordani group generally has high meristic values, the lateral line scales Chirostoma chapalae - - - - - - - - - - - - - - 107 with canals, scales with laciniate margins Chirostoma humboldtianum - - - - - - - - 108 and includes C. jordani, C. patzcuaro, C. Chirostoma consocium consocium - - - - 110 chapalae, C. consocium, C. humboldtianum, Chirostoma consocium reseratum - - - - 110 C. estor, C. grandocule, C. /ucius, C. sphy- raena and C. promelas. The arge species Chirostoma grandocule - - - - - - - - - - - - 112 group generally has low meristic values, Chirostoma cornpressum - - - - - - - - - - - 114 the lateral line scales with round pores Chirostoma estor estor - - - - - - - - - - - - - 114 (except for the last two species named Chirostoma estor copandaro - - - - - - - - - below) and scales with smooth margins and 116 includes C. arge, C. melanoccus, C. charari, Chirostoma lucius - - - - - - - - - - - - - - - - 118 C. riofai, C. bartoni, C. attenuatum, C. Chirostoma sphyraena - - - - - - - - - - - - - 118 labarcae and C. aculeatum. Chirostoma promelas - - - - - - - - - - - - - - 120 The similarity between C. arge and Me- THE ARGE GROUP - - - - - - - - - - - - - - - 120 laniris crystallina and the primitive nature Chirostoma arge - - - - - - - - - - - - - - - - - - 122 Chirostoma melanoccus - - - - - - - - - - - - 122 Present address: Division of Fishes, Mu- seum of Zoology, The University of Michigan, Chirostoma rio jai - - - - - - - - - - - - - - - - 122 Ann Arbor, Michigan 48104. Chirostoma charari - - - - - - - - - - - - - - - - 126 EDITORIAL COMMITTEE FOR THIS PAPER: DR. ERNEST A. LACHNER, Curator, Division of Fishes, National Museum of Natural History, Washington, D. C. 20560 DR. ROBERT R. MILLER, Curator of Fishes, Museum of Zoology, The University of Michigan, Ann Arbor, Michigan 48104 97 98 Tulane Studies in Zoology and Botany Vol. 18 Chirostoma attenuatum attenuatum - - 128 as having thin, translucent, pale flesh and Chirostoma attenuatum zirahuen - - - - 128 differences in the firmness of its scales and Chirostoma bartoni - - - - - - - - - - - - - - 129 bones. The same authors later decided ( Jor- Chirostoma labarcae - - - - - - - - - - - - - - 129 dan and Evermann, 1896-1900, Vol. 4, p. Chirostoma aculeatum - - - - - - - - - - 130 3158) that Lethostole was not separable PHYLOGENY OF Chirostoma - - - - - - - 131 from Chirostoma. Meek (1904) recognized PRIMITIVE AND DERIVED CHARACTERS 131 one genus and allocated the species among PRIMITIVE SPECIES WITHIN THE SPE- three subgenera: Chirostoma, Eslopsarum CIES GROUPS - - - - - - - - - - - - - - - 132 and Lethostole. Regan (1906-08) and Jor- POSSIBLE ANCESTORS AND THE ORIGIN dan and Hubbs (1919) could find no con- OF Chirostoma - - - - - - - - - - - - - - - - 133 sistent method by which the species could EVOLUTION WITHIN THE SPECIES be grouped and so recognized no subgenera. GROUPS - - - - - - - - - - - - - - - - - - - 137 Jordan, Evermann and Clark (1930) fol- CONCLUSIONS - - - - - - - - - - - - - - - - 138 lowed Meek (1904). De Buen (1945) dis- RESUMEN - - - - - - - - - - - - - - - - - - - - 139 tributed the species among three genera and LITERATURE CITED - - - - - - - - - - - - - 140 six subgenera. Alvarez (1950,1970) recog- nized two genera in his keys, Chirostoma and the monotypic Otalia. INTRODUCTION The objectives of this paper are to re- The Republic of Mexico, by reason of its define the species which have been included geographic position and geological history, in Chirostoma and to discuss their evolu- possesses a large and interesting fish fauna. tionary relationships. No attempt has been The coastal lowlands have served as avenues made, however, to diagnose the genus be- of migration for North and Central Amer- cause little comparative data is available for ican fishes and contain mixed faunas reflect- most of the other atherine genera. Patterns ing strong affinities for these regions. On of morphological variation within the family the other hand, the uplifted, central portion (Rosen, 1964) are not yet clear. This some- of the country, from the transverse volcanic what unorthodox presentation is also jus- axis northward, contains a large number of tified by the biogeographic history of Chiro- unique species. The fishes present in the stoma, a topic to be discussed in another basins of the north show clear but varying paper. degrees of relationship with the Rio Grande fauna. On the southern one-third of the METHODS pleateau, however, in the westward flowing Counts and measurements were made in Lerma-Santiago river system and contiguous accordance with Hubbs and Lagler (1958) fluvial and lacustrine basins, endemism in most instances. Where these methods reaches generic and familial levels. were in appropriate they were modified or Chirostoma is one of the unique groups in new characters used as described below. this evolutionary microcosm. The genus has The number of median lateral scales was been reviewed or revised several times, most considered to be the number of scales in a notably by Jordan and Evermann (1895; series extending from the scapular arch 1896-1900), Meek (1904), Regan ( 1906— dorsal and anterior to the insertion of the 1908), Jordan and Hubbs (1919) and De pectoral fin to the end of the hypural plate. Buen (1945). Through the years the num- The count includes scales which may be ber of species has steadily increased as well reduced in size just posterior to the head. as the confusion regarding their relation- The predorsal scale count was made along ships. Jordan and Evermann (1895) placed an imaginary line between the origin of the the species into two genera: Chirostoma, first dorsal fin and the head and included, characterized as having the snout slightly when present, the large scale which usually produced, small teeth, an oblong body and has its origin on the head, but lies on the a small first dorsal fin placed over the vent nape. Only those scales intersecting the line and Eslopsarum, being close to Chirostoma, by at least one-half a scale width were but with large and entire scales. Jordan counted. The interdorsal scales, the number and Evermann (1896-1900) erected a third of scales in a series lying between the scale genus, Lethostole, for C. estor characterized upon which the first dorsal fin membrane No. 3 Chiro stoma 99 inserts and the origin of the second dorsal Society (1935), the road atlas, Caminos de fin, were counted in the same manner as the Mexico, published by La Compania Hulera predorsals. Euzkadi (third edition, 1967) and from my The condition of the scale margins and own observations. Where more than one the type of lateral line pore were also found collection was taken from a specific locality, to be useful characters. The former are only one is shown on the map. Questionable either smooth (entire) or laciniate (toothed locality records are not plotted on the maps, or crenulate ). The openings in the scales of but are discussed in the text under Range. the lateral line system are either horizontally The following abbreviations appear in this elongate (canals) or round (pores). paper: ca. (circa), about; ft., feet; hwy., Nineteen morphometric and nine meristic highway; jct., junction; km, kilometer; mi., characters were examined in each of the miles; trib., tributary. Geographic directions species of Chirostoma over their known are indicated by the initial letters of the ranges and in Menidia berAna (Grand cardinal points of the compass, North, South, Isle, Louisiana) and Melaniris crystallina East, West or various combinations thereof. ( Rio Grande de Santiago at Hwy 15 crossing, Nayarit). Wherever possible body propor-
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