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North American Minnows, with Emphasison The QL 638 .C94 M3 UNIVERSITY OF KANSAS MISCELLANEOUS PUBLICATION 1 989 MUSEUM OF NATURAL HISTORY No. 80 MCZ LIBRARY 198 HARVAF UNIV iY Phylogenetic Studies of North American Minnows, with Emphasis on the Genus Cyprinella (Teleostei: Cypriniformes) By Richard L. Mayden UNIVERSITY OF KANSAS LAWRENCE June 1, 1989 .C Ml UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum ofNatural History. The Mis- cellaneous Publications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion of the Director. Beginning with this number, Miscellaneous Publications are type set using Microsoft® Word and Aldus PageMaker® on a Macintosh computer. Conversion to current format by E. O. Wiley. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66045, U.S.A. Individuals may purchase separate numbers from the Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 6604-2454, U.S.A. The University of Kansas Museum of Natural History Miscellaneous Publication No. 80 June 1, 1989 Phylogenetic Studies of North American Minnows, with Emphasis on the Genus Cyprinella (Teleostei: Cypriniformes) By Richard L. Mayden Department of Biology The University of Alabama Tuscaloosa, Alabama 35487-0344 U.S.A. The University of Kansas Lawrence 1989 >38 University of Kansas Publications, Museum of Natural History 43 Editors: Robert M. Mengel and Richard F. Johnston Managing Editor. Joseph T. Collins Consulting Editor. E. O. Wiley Design and Typesetting: Kate A. Shaw and Joseph T. Collins Miscellaneous Publication No. 80 Pp. 1-189; 85 figures; 4 color plates Published June 1, 1989 Museum of Natural History The University of Kansas Lawrence, Kansas 66045-2454 U.S.A. MCZ LIBRARY JUL 5 1989 HARVARD UNIVERSITY Printed by University of Kansas Printing Service Lawrence, Kansas CONTENTS INTRODUCTION 1 ACKNOWLEDGMENTS 2 METHODS 4 MATERIALS EXAMINED 6 MONOPHYLY OF CYPRINELLA AND RELATIONSHIPS TO OTHER NORTH AMERICAN CYPRINIDS 9 Cyprinids with an Open Posterior Myodome 11 ChubClade 13 Genus Pimephales 25 Genus Hybopsis and Related Species 25 Pteronotropis-Notropis Clade 29 Genus Pteronotropis 30 Lythrurus-Notropis Clade 32 Lythrurus-Luxilus-Cyprinella Clade 32 Genus Lythrurus 34 Luxilus-Cyprinella Clade 34 Genus Luxilus 38 Genus Cyprinella 38 Genus Notropis 41 Subgenus Alburnops 42 Notropis volucellus Species Group 42 Subgenus Hydrophlox 43 Subgenus Notropis-N. texanus Species Group 44 Notropis texanus Species Group 44 Subgenus Notropis 44 SYSTEMATICS OF CYPRINELLA 45 Cyprinella Girard 45 Diagnosis 45 Composition 45 Distribution 45 Classification of Cyprinella 45 SPECIES ACCOUNTS 46 Cyprinella ornata 46 Cyprinella lutrensis 49 Cyprinella garmani 53 Cyprinella formosa 55 Cyprinella bocagrande 57 Cyprinella lepida 58 Cyprinella rutila 60 Cyprinella xanthicara 61 Cyprinella proserpina 63 Cyprinella panarcys 65 Cyprinella spiloptera 66 Cyprinella camura 68 Cyprinella whipplei 71 Cyprinella analostana 73 Cyprinella chloristia 76 Cyprinella venusta 77 Cyprinella galactura 80 Cyprinella pyrrhomelas 82 Cyprinella xaenura 84 Cyprinella caerulea 85 Cyprinella gibbsi 87 Cyprinella trichroistia 89 Cyprinella callistia 90 Cyprinella nivea 92 Cyprinella leedsi 93 Cyprinella callisema 95 Cyprinella callitaenia 97 OSTEOLOGY 107 Ethmoid Region 107 Mesethmoid 108 Lateral Ethmoid 108 Vomer 117 Preethmoid 119 Supraethmoid 119 Kinethmoid 120 Nasal 120 Orbital Region 120 Orbitosphenoid 120 Pterosphenoid 120 Frontal 121 Supraorbital 124 Infraorbital Series 124 Otic and Occipital Regions 126 Sphenotic 127 Prootic 127 Dermopterotic 130 Autopterotic 131 Parietal 131 Exoccipital 131 Epiotic 132 S intraoccipital 132 Intercalar 132 Basicranial Region 132 Basioccipital 132 Parasphenoid 134 Opercular Region 134 Opercle 135 Preopercle 135 Interopercle 136 Subopercle 138 Hyopalatine Region 138 Hyomandibular 138 Palatine 139 Ectopterygoid 142 Endopterygoid 142 Quadrate 142 Symplectic 143 Metapterygoid 143 Upper Jaw 143 Premaxilla 143 Maxilla 144 Lower Jaw 147 Dentary 147 Anguloarticular 147 Retroarticular 148 Sesamoid articular 148 Mentomeckelian 148 Hyoid Region 149 Interhyal 149 Ceratohyal 149 Hypohyal 151 Basihyal 151 Urohyal 153 Branchiostegals 154 Branchial Region 157 Pharyngobranchials 157 Epibranchials 158 Ceratobranchials 160 Hypobranchials 162 Basibranchials 162 Weberian Apparatus 162 Pars sustenaculum 163 Pars auditum 163 Vertebrae 164 Caudal Skeleton and Fin Supports 165 Neural and Haemal Arches and Spines 165 Epural 165 Pectoral Girdle 166 Posttemporal 166 Cleithrum 167 Supracleithrum 168 Coracoid 168 Mesocoracoid 168 Scapula 168 Postcleithrum 169 Pelvic Girdle 169 PHYLOGENETIC RELATIONSHIPS 169 Cyprinella lutrensis clade 170 Cyprinella lutrensis-formosa-lepida species groups 171 Cyprinella formosa-lepida species groups 171 Cyprinella lepida species group 171 Cyprinella rutila-proserpina species pairs 171 Cyprinella rutila species pair 172 Cyprinella proserpina species pair 172 Cyprinella formosa species group 172 Cyprinella lutrensis species group 173 Cyprinella whipplei clade 173 Cyprinella whipplei-venusta-pyrrhomelas-nivea species groups 174 Cyprinella whipplei species group 174 Cyprinella whipplei-analostana-chloristia species group 174 Cyprinella analostana-chloristia species pair 174 Cyprinella venusta-pyrrhomelas-nivea species groups 175 Cyprinella venusta species group 175 Cyprinella pyrrhomelas-nivea species groups 175 Cyprinella pyrrhomelas species group 176 Cyprinella pyrrhomelas species pair 176 Cyprinella caerulea species complex 177 Cyprinella trichroistia-gibbsi species pair 177 Cyprinella nivea species group 178 Cyprinella nivea-leedsi-callisema-callitaenia species group 178 Cyprinella leedsi-callisema-callitaenia species group 178 Cyprinella callisema-callitaenia species pair 178 SUMMARY 179 LITERATURE CITED 179 INTRODUCTION The North American cyprinid fauna is di- Jenkins and Lachner (1971) for Nocomis and verse, comprising nearly 30 percent of the Hybopsis; Buth and Burr (1978) for Campos- total fish diversity north of Mexico. Our un- toma; Coad (1975) for many North American derstanding of cyprinid diversity is reasonably and Eurasian cyprinids; and Hubbs and Miller complete, but cyprinids are poorly known (1977) for the genus Dionda. In all of the phylogenetically. The present status of the above studies statements of relationship are systematics of this group has not changed generally presented. appreciably since 1977, when Carl L. Hubbs Coad (1975) and McAllister and Coad and Robert R. Miller stated "It is abundantly (1978) were among the first to examine higher obvious that much of the generic placement in relationships of North American cyprinids, American cyprinids is in a chaotic state . ." but some problems remain. Because the genus (Hubbs and Miller, 1977, p. 275). Many im- Notropis is so diverse, it was considered the portant studies of cyprinid species variation most primitive and was used to determine have been published in North America, again character polarities. This technique may seem aiding our understanding of species diversity, sound, but character evaluation may be flawed but not necessarily improving our knowledge given that Notropis may in fact be an ingroup. ofgenealogical relationships. Only a few stud- In the latter study hybridization frequency ies have developed hypotheses of species rela- data were used as an indication of genealogical tionships. Many have concentrated on Notro- relatedness. Although innovative, this meth- pis, the largest of the North American genera. odology is suspect because reproductive Suttkus (1950) examined the subgenus Pter- compatability may be a retained primitive onotropis, Gilbert (1964), Rainboth and Whitt characteristic (Rosen, 1979), and like any (1974),Buth(1978),ButhandMayden(1981), primitive morphological or genetic character, Menzel (1976, 1977), Dowling and Moore is not indicative of a close relationship. Fur- (1984), and Mayden (1988) dealt with the ther, frequency of opportunity to participate in subgenus Luxilus, Snelson (1972) and Stein et hybridization was not considered. Not until al. (1985) with Lythrurus, Swift (1970) with the late 1970's was the philosophy of phylo- Hydrophlox, Alburnops, and other species genetic systematics (sensu Hennig, 1966) groups, and Gibbs (1955, 1957a) and Contre- applied to species relationships of North ras-Balderas (1975, 1978) with Cyprinella. American cyprinids. Barbour and Miller's Our knowledge of Notropis diversity and sys- (1978) and Jensen and Barbour's (1981) study tematics has been gready aided by studies by of Algansea, Buth's (1978) analysis of Lux- Contreras-Balderas (1975, 1978), Hubbs and ilus, and Coburn's (1982) analysis of the Miller (1978), Chernoff and Miller (1981, subgenus Notropis, as well as other species of 1982, 1986), and Chernoff et. al (1982). In this genus and other North American genera, these revisionary studies hypotheses or state- were pioneering efforts into
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