North American Minnows, with Emphasison The

QL 638 .C94 M3 UNIVERSITY OF KANSAS MISCELLANEOUS PUBLICATION 1 989 MUSEUM OF NATURAL HISTORY No. 80

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198 HARVAF UNIV iY

Phylogenetic Studies of North American Minnows, with Emphasis on the Genus Cyprinella (Teleostei: Cypriniformes)

Richard L. Mayden

UNIVERSITY OF KANSAS LAWRENCE June 1, 1989 .C Ml

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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)

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 the systematics of ments of relationships may be present. North American minnows. Systematic studies of other North Ameri- Cyprinella is one of many recognizably can genera include Hubbs and Black (1947) monophyletic (sensu Hennig) groups of the and Niazi ( 1 960) for Pimephales; Illick (1956) North American cyprinid fauna. Presendy Cyp- for many cyprinids; Sisk (1961) for Notropis, rinella contains 27 species, distributed on the Uyeno (1961) for Gila; Reno (1966, 1969) for Atlantic Slope from the St. Lawrence River some Notropis and Hybopsis; Barbour and south to the Gulf Coast, west to the Great Miller (1978) and Jensen and Barbour (1981) Plains, and southwest to the Rio Nazas of for Algansea; Buhan (1970) for the genera central Mexico. Previous systematic works on Richardsonius, Clinostomus, and Semotilus; the group were mainly alpha in level, with the MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY exception of Gibbs, who supported species or between species {analostana and chloristia) species group relationships with characters. or subspecies {venusta and spilopterd) or none Species level problems have been reasonably at all. Whether these narrow areas of introgres- well resolved through efforts by Bailey and sion reflect conspecificity remains to be tested. Gibbs (1956), Gibbs (1955, 1957a, 1957b, The present study is a review of the genus 1957c, 1961, 1963), and Howell and Williams Cyprinella, with osteological descriptions of (1971) for eastern members of this group, the included species and hypotheses of their herein regarded as the whipplei clade. The phylogenetic relationships. Further, the rela- southwestern U. S. and Mexico clade, herein tionships of Cyprinella to other North Ameri- referred to as the lutrensis clade, has been can cyprinids are presented. In doing this, thoroughly studied by Contreras-Balderas some previously recognized subgenera of the (1975,1978),MinckleyandLytle(1969),Lytle large and unnatural genus Notropis are ele- (1972), Hubbs and Miller (1978), Chernoff vated to generic rank. These changes and oth- and Miller (1982), and Matthews (1987). ers are done so that species relationships are In spite of these efforts, some taxonomic consistent with formal classification. Because problems remain in Cyprinella. Within the of a growing interest in the ichthyological wide-ranging red shiner, Cyprinella lutrensis, community to recognize Cyprinella as a dis- several Mexican forms referred to by Contre- tinct genus rather than a subgenus of Notropis, ras-Balderas (1975, 1978) as subspecies of C. and the necessity to insure that the classi fication lutrensis are better recognized as separate is consistent with the phylogenetic relation- species and some may represent undescribed ships of the species, I have recognized Cyp- forms. Cyprinella lepida of the Edwards Pla- rinella as a distinct genus. In doing so, Luxilus, teau of Texas is considered a distinct species Lythrurus, and Pteronotropis are also recog- and the number of forms of Cyprinella lutren- nized as distinct genera because of their sister sis present in drainages in the U. S. deserves or close relationships to the genus Cyprinella. additional investigation. Although a first at- Relationships presented here are hypotheses tempt at the latter problem has been made by that remain to be tested with additional, and Matthews (1987), additional study is recom- perhaps different data sets. Further, some of mended. In the primarily eastern whipplei clade the above listed areas of concern in the genus interesting questions remain concerning the Cyprinella could not be addressed in this specific status of the present subspecies of spi- analysis. These areas of concern, as well as loptera and venusta, and the species analostana others, should be investigated independently and chloristia. All of these species have very to corroborate or falsify hypotheses presented narrow regions of hypothesized introgression here. ACKNOWLEDGMENTS

During my tenure as a graduate student in discussions with Gloria Arratia, Reeve M. biology, many people have been generous Bailey, Herbert T Boschung, Brooks M. Burr, with their time and money to further my edu- Donald G. Buth, Barry Chernoff, Miles Co- cation. I particularly wish to thank Marvin E. burn, Frank B. Cross, William L. and Sara V. Braasch, Brooks M. Burr, Donald G. Buth, Fink, Carter R. Gilbert, Robert E. Jenkins, Frank B. Cross, Lawrence M. Page, and E. O. Lawrence M. Page, J. D. Stewart, Walter Rain- Wiley for their assistance in initiating my both, and E. O. Wiley. My thanks to all of these career in ichthyology, for sharing research people. ideas, and for constant support over the years. I am grateful to the following individuals My research efforts with cyprinid sys- and their institutions for the loan and/or dona- tematics have been greatly furthered through tion of materials pertinent to this study: Reeve NORTH AMERICAN MINNOW GENUS CYPRINELLA

M. Bailey, Robert R. Miller, William L. Fink, ate students. I hold these experiences with the and Douglas W. Nelson, University of Michi- deepest appreciation. I wish to express my gan Museum of Zoology (UMMZ); Frank B. thanks to my close friends Richard Albano, Cross, E. O. Wiley, and Joseph T. Collins, Uni- Thomas Berger, Brad Bergstrom, Fernando versity of Kansas (KU); W. L. Minckley and Bird-Pico, Pacha Burrowes, Jonathan Dean A. Hendrickson, Arizona State Univer- Campbell, David Cannatella, Kevin Cum- sity (ASU); Steve Platania and William Palmer, mings, Fernando Cervantes, Richard Cloutier, North Carolina State Museum (NCSM); Rafael de Sa, Sharon Dewey, Barb Stein, Linda Lawrence M. Page, Illinois Natural History Dryden, Sushil Dutta, Linda Ford, Darrel and Survey (INHS); Brooks M. Burr, Southern Lynne Frost, Owen and Wendy Gorman, Peter Illinois University at Carbondale (SIUC); Gray, Dave and Ann Hillis, Thor and Elaine David A. Etnier, John Randy Shute, and Bruce Holmes, Rafael Joglar, Steve Johnson, Brad H. Bauer, University of Tennessee (UT); Livezey, Mike Morris, Becky Pyles, Mike Edward Esmond, University of Maryland Retzer, John Sauer, Kate Shaw, John Sim- Center for Environmental and Estuarine Stud- mons, J. D. Stewart, Tom Titus, Steve Walsh, ies; Herbert T. Boschung and David L. Nie- Kim Wollter, and Chris Wozencraft. land, University of Alabama Ichthyological I also wish to thank Brooks M. Burr, George Collecuon (UAIC); Henry W. Robison, South- Byers, Frank B. Cross, Carter R. Gilbert, and ern Arkansas University; Neil H. Douglas, E. O. Wiley for their review of this manuscript. Northeast Louisiana University; John S. Further, I am deeply indebted to Ed Wiley for Ramsey, Auburn University (AUM); Robert many hours of discussion of the theory and Schoknecht, Cornell University (CU); Royal application of phylogenetic systematics and D. Suttkus (RDS), Tulane University (TU); for support while at the University of Kansas. Brian W. Coad, National Museums of Canada, I am very grateful to Darrel Frost for his help Museum ofNatural Sciences; Tyson R. Roberts with taxonomic name changes and many pro- and William N. Eschmeyer, California Acad- vocative discussions in systematics. A very emy of Sciences; Robert F. Martin and Doyle special thanks to John Simmons for preparing Mosier, Texas Natural History Collection the black and white prints of the dorsal fins and (TNHC); Edward F. Menhinick, University of breeding tubercles from less than perfect nega- North Carolina at Charlotte; John L. Harris, tives, and to Linda Trueb for the use of photo- Arkansas State Highway and Transportation graphic equipment and advice with the illus- Department; RobertE. Jenkins (REJ) and Noel trations. I would also like to thank Richard Burkhead (NB), Roanoke College; James J. Cloutier, Brian Danforth, and Kate Shaw for Long, Oregon State University; Douglas E. executing figures 50A-C and Brooks M. Burr, Stephans, Kentucky Department of Fish and Lawrence M. Page, Robert R. Miller, and Wildlife Resources (KFW); Camm C. Swift, William N. Roston for supplying color slides Natural History Museum of Los Angeles of species of Cyprinella which were invalu- County (LACM); Carter R. Gilbert, Florida able in this study. I am especially thankful to State Museum, University of Florida (UF); Joseph T Collins, Robert Mengel, and E. O. William J. Matthews (WJM), University of Wiley of the University of Kansas, Museum of Oklahoma Biological Station; Robert Karl Natural History for their tremendous amount Johnson, Field Museum of Natural History of assistance in facilitating the publication of (FMNH); and curators of Museum National this manuscript. This research was partially d'Histoire Naturelle, Paris (MNHN). supported by the University of Kansas Gen- Some of the most enjoyable and important eral Research Fund (No. 3338), National Sci- aspects of my graduate career include the ence Foundation (research grant to R. L. many personal and academic benefits I have Mayden, BSR 861443 1), and Sigma Xi. I also gained through friendships with fellow gradu- thank Bernard Kuhajda and Michelle Mayden MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY for their technical assistance with the manu- Finally, I wish to thank my family for years script. This is contribution number 113 from of moral and financial assistance while endur- the University of Alabama Aquatic Biology ing a graduate career. To these people I dedi- Program. cate this publication.

METHODS

The phylogenetic techniques used in this outlined by Hubbs and Lagler (1964), Contre- analysis to determine species relationships ras-Balderas (1975), Howell and Williams follow Hennig (1966) and Wiley (1981). De- (1971), Hubbs and Miller (1978), and Chern- termination of character polarity involved off and Miller (1982). In some accounts, data outgroup comparison and ontogeny, where are summarized from some of these publica- possible. When two characters occurred within tions. Precaudal and caudal vertebrae were Cyprinella the character found in outgroups distinguished on the basis of haemal spines. was considered primitive within Cyprinella. The first caudal element was the first element The alternate character was considered de- with a well-developed haemal spine. The pre- rived and used to determine species relation- caudal vertebra at the dorsal fin insertion was ships. The primary outgroups used in deter- that element below the anterior extent of the mining species relationships within the genus first pterygiophore. In each account pore counts Cyprinella included species of the genera presented for cephalic -lateral is canals are for Luxilus and Lythrurus. Justifications for using one side of the head; numbers listed in paren- these subgenera as outgroups are provided theses for meristic characters represent modes. below. Peak or high males in discussions of tubercu- Characters used in this analysis included lation patterns and development of tubercles osteology, tuberculation, and body and fin refers to breeding males with maximum tu- pigmentation patterns. Osteological charac- bercle development. Gut coiling terminology ters were taken from cleared and double stained follows Kafuku (1958). Where scale counts specimens and dissarticulated materials pre- are presented numbers refer to the number of pared with methods outlined by Dingerkus scale rows; modes are in parentheses. and Uhler (1977) and Mayden and Wiley Terminologies and origins of bones follow (1984). Osteological characters were drawn Harrington (1955), Patterson (1977), and with the aid of a camera lucida. Whenever Coburn (1982). Nomenclature of the caudal possible, interspecific comparisons of mor- skeleton follows Nybelin (1963) and Buhan phologies were made between individuals of (1972). Exceptions include: the urodermals of similar standard length to avoid possible prob- the latter two authors are here considered lems associated with allometry. All body uroneurals, following Patterson (1968); lat- lengths referred to in the text and figures are eral process of the parhypural in B uhan (1972) standard lengths (SL). Tuberculation charac- and Gosline (1960) is considered the hypu- ters were taken from breeding males with rapophysis after Nursall (1963). The follow- maximum tubercle development. Males early ing anatomical abbreviations are used through- in tubercle development were also examined out the text and in illustrations for identification to assess ontogenetic characters in tubercle of structures: patterns. Coloration characters were taken from freshly collected specimens or color photo- AIT, tubercle of maxilla for insertion of Al graphs of freshly preserved individuals in branch of adductor mandibulae muscle breeding condition. ABHL, anterior basihyal ligament Within species accounts, meristic and ACH, anterior ceratohyal morphometric data presented follow methods AP, ascending process of premaxilla NORTH AMERICAN MINNOW GENUS CYPRINELLA

APT, autopterotic KE, kinethmoid ART, anguloarticular LA, lacrymal ASP, ascending process of maxilla LE, lateral ethmoid BB,basibranchial LG, lacrymal groove BH, basihyal LOF, lateral occipital foramen BO, basioccipital LP, lateral process BPT, basipterygium M, mentomeckelian BR, branchiostegal MC, Meckel's cartilage CB, ceratobranchial MCO, mesocoracoid CH, ceratohyal ME, mesethmoid CL, cleithrum MP, median process CLA, claustrum MPT, metapterygoid COR, coracoid MX, maxilla CP, coronoid process MXM, premaxillary process of maxilla DBHL, dorsal basihyal ligament N, nasal DE, dentary NA, neural arch DF, dilator fossa NC, neural crest DHH, dorsal hypohyal NPU, compound centrum DP, dorsal process of epibranchial 4 NS, neural spine DPT, dermopterotic OF, olfactory foramen EB, epibranchial OP, operculum ECP, ectopterygoid ORS, orbitosphenoid ENP, endopterygoid OS, os suspensorium EOC, exoccipital PA, parietal EP, epural PAL, palatine EPB, epiphyseal bar PBR, pharyngobranchial EPO, epiotic PCH, posterior ceratohyal FIC, carotid foramen PCL, postcleithrum FM, foramen magnum PDB, predorsal bone FR, frontal PE, preethmoid FV, trigeminal foramen PET, planum ethmoidale FVII, facial foramen PG, palatine groove H, hypural PH, parahypural HA, haemal arch PMO, posterior myodome opening HB, hypobranchial PMX, premaxilla HH, hypohyal POP, preoperculum HN, hyomandibular notch PPBO, pharyngeal process of basioccipital HS, haemal spine PPD, pharyngeal pad HYF, hyoideal foramen PR, posterior ramus of premaxilla HYO, hyomandibular PRO, prootic IC, cavum sinus impar PS, parasphenoid ICA, intercalarium PSOC, palatine socket IH, interhyal PTF, posttemporal fossa 1MB, intermuscular bone PTS, pterosphenoid 10, infraorbital bone PTT, posttemporal IOP, interoperculum PU, preural ISC, ischiac process QA, quadrate 1ST, isthmus of maxilla R,rib 6 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

RA, retroarticular STE, subtemporal fossa RAD, radial SY, symplectic RPM, rostral process of maxilla TRP, tripus SA, sesamoid articular UN, uroneural SCA, scapula UO, urohyal SCL, supracleithrum UOL, urohyal ligament SCM, scaphium UP, uncinate process SE, supraethmoid V, vertebra SEP, supraethmoid process of palatine VBHL, ventral basihyal ligament SO, supraorbital VHH, ventral hypohyal SOC, supraoccipital VO, vomer SOP, suboperculum VP, vomer process of palatine SPH, sphenotic MATERIALS EXAMINED

The following specimens were examined 6828 (8); KU 8809 (23), 19232 (10:5CS); TU in this revision. For each species the museum 23637 (42), 92770 (9:11CS). C. camura KU acronym and number is listed first, followed 12112 (2CS), 12215 (15), 12231 (9), 16019 by the number of specimens in parentheses. (14CS), 15734 (10), 15792 (IOCS), 19715 For some collections both cleared and double- (3CS), 20591 (20); LACM 39815-1 (IOCS). stained and alcoholic specimens were exam- C. chloristia KU 5267 (1), 5274 (4), 8882 ined from a single collection. Cleared and (15:12CS),8894(2),20O41(13CS);TU 28224 stained specimens are identified with the ab- (5). C.formosa ASU 64-0820 (4); KU 8307 breviation CS. Institutional acronyms are (47),8399(42:15CS),8391(15CS).C.£tf/ac- identified in acknowledgments. tura KU 10937 (10), 10882 (50), 10839 (49), Agosia chrysogaster KU 8084 (5:3CS), 9512 (9), 7866 (9), 11225 (9), 5205 (2), 7727 8085 (24:6CS). Aztecula sallei KU 5433 (3); (7),7765(15),9512(9CS), 16461 (2CS),9512 UMMZ 108625 (6CS), 182346 (6), 191695 (19), 19732 (13), 10882 (50), 14966 (14),

(6CS), 192368 (6). Campostoma anomalum 16461 (2), 19732 (13), 12029 (12CS). C. gar- KU3946(1:4CS), 12581 (2CS), 13200 (3CS). mani FMNH 4368 (30); KU 5416 (17:8CS), C. oligolepis KU 7583 (17CS), 12502 (1CS), 20164 (51: IOCS). C. gibbsi KU 8790 (44), 12520 (2CS). C. ornatum KU 3251 (15). Cli- 15331 (10), 16904(10), 18892 (3:12CS),20266 nostomus elongatus KU 11322 (2CS). C.fun- (25), 20284 (1 52), 20472 (38), 20285 (47); TU duloides INHS 86752 (IOCS); KU 3262 (38), 40670 (32). C. leedsi KU 1 8985 ( 1 3CS); UAIC 10697 (4:6CS).CouesiusplumbeusKU 18881 3127.01 (6); UF8515 (15), 14535 (12),29099 (6:8CS). Cyprinella analostana INHS 74347 (30). C. lepida TNHC 5279 (63), 5634 (2), (15CS), 77855 (3CS); KU 5446 (1), 15367 7572 (50); TU 55189 (5: IOCS); WJM 375 (26), 19785 (10), 20400 (20); TU 25867 (31). (13). C. lutrensis KVS3SS (72), 10451 (5CS), C. bocagrande KU 20399 (1CS); UMMZ 15595 (274), 17293 (9), 17297 (10), 18779 208250 (1CS), 208265 (5). C. caerulea INHS (50), 19431 (8CS), 20592 (147); LACM 39833- 74805 (9CS); KU 18978 (2:8CS), 19666 (13); 1 (8CS); TNHC 1400 (15). C. nivea INHS LACM 38942-1 (IOCS). C. callisema KU 76888 (8CS); KU 5353 (1), 8883 (14), 18987 8830 (11), 8836 (9CS), 8842 (28: 11CS), 19653 (3:12CS); TU 29483 (6), 38616 (3). C. ornata (24); TU 26173 (17). C. callistia KU 5293 (1), ASU 9629 (6); INHS 84079 (10); KU 4997 8777(2), 16900(10), 18848(14:11CS), 18888 (45:10CS), 8405 (29:6CS). C. panarcys ASU (20), 19697 (15), 20166 (12CS), 20265 (3), 64-0853 (6); KU 4991 (5), 5404 (4), 5439 (3);

20282 (2), 20483 (36). C. callitaenia AUM TNHC 4341 (33); UMMZ 208212 (14CS).C. NORTH AMERICAN MINNOW GENUS CYPRINELLA proserpina ASU 3600 (5); KU 18852 18917 (15CS). Hybognathus nuchalis KU (15:10CS); TNHC 3262 (10:10CS), 9771 14123 (1CS). H. placitus KU 9766 (5CS), (170). C.pyrrhomelas INHS 76839 (4:16CS), 12597 (3CS). Hybopsis alborus INHS 75078 76978 (3CS); KU 5276 (2 1), 1 8989 (1 5); UAIC (6CS); KU 18973 (2: IOCS). H. amblops KU 2550.05 (10). C.rutila ASU 5982 (5:9CS), 64- 63 18 (IOCS), 16556 (6CS); TU 25401 (2CS). 0264 (12: IOCS), 64-0913 (5); KU 7347 (64). H. amnisKU 18847 (IOCS); TU 34008 (2CS). C. spiloptera KU 8628 (87), 3595 (17), 10088 H. bifrenatus KU 18977 (12CS). H. boucardi (11), 10353 (50), 10599 (8), 11356 (17:3CS), TU 55376 (3CS); UMMZ 191686 (7:8CS). H. 14561 (6CS), 17776 (IOCS); LACM 39819-1 calientis RDS 3408 (2CS);TU30695(5:15CS). (IOCS). C. trichroistia KU 5292 (4), 18853 H. dorsalis KU 4670 (98), 5382 (47), 12223 (IOCS), 20450 (31); LACM 39851-1 (12CS); (111), 16132 (103), 16237 (15CS). H. hypsi- UAIC 5550.04 (84), 5526.04 (36), 5528.04 notus KU 18968 (8CS), 20034 (4CS); REJ (148), 5557.04 (290), 5554.06 (238), 5551.04 1113 (1); REJ uncat. (20). H. labrosa CU (199), 5559.03 (237), 5559.07 (340), 5527.06 10922 (37); INHS 88319 (4:6CS); KU 20035 (190), 5556.06 (562), 5525.02 (41), 5534.06 (2CS). H. sp. cf. labrosa KU 20037 (3CS). H. (242), 5536.06 (349), 5532.05 (203). C. ve- UneapunctataKU 18885(2:8CS).//. longiros- nusta KU 8800 (118), 8810 (IOCS), 9413 (mKU 16871 (109), 17965 (10: 15CS);LACM (115), 9570 (89), 16318 (227), 17835 (28), 39853-1 (IOCS); RDS 3272 (1CS). H. ru- 17869 (9), 17591 (27), 19674 (2), 19698 (57); brifrons KU 8829 (1:8CS); TU 26159 (2CS). LACM 39831-1 (IOCS); RDS 3374 (1CS), H. sabinae KU 6237 (32: IOCS), 9480 (24),

3375 (1CS). C. whipplei KU 9040 (37), 1 1357 19673 (6). H. winchelli KU 16896 (25); TU (40), 14211 (15CS); LACM 39820-2 (IOCS). 3031 (2CS), 39705 (2CS); RDS 3384 (1CS). C. xaenura KU 8823 (8), 18994 (3:12CS), H. zanema INHS 76841 (6CS), 88320 (15CS); 19655 (8CS); TU 29458 (3). C. xanthicara KU 20036 (4CS).Iotichthysphlegethontis KU ASU 3642 (18:9CS), 5102 (6); INHS 84062 11935 (5CS). Lepidomeda mollispinis KU (2CS); UMMZ 179205 (25), 179827 (15), 11937 (3CS). Luxilus albeolus INHS 27231

179878 (6). Cyprinuscarpio KU 12440(1CS). (5:15CS); KU 3266 (5), 18972 (10); UAIC Dionda episcopa INHS 83145 (15CS); KU 4144.04 (58). L. cardinalis KU 12295 (48), 7427 (5:7CS), 16891 (5CS). D. ipni INHS 14418(147), 15015(85), 15281 (13CS), 16247 75342 (3), 75359 (3), 75369 (7). Ericymba (16),20737(23),20963(l).L.cmww«sINHS buccata KU 17764 (4:8CS). Erimystax cahni 83520(1);KU3281(26:14CS);UAIC4144.07 KU 19614 (3CS); REJ 737 (2). E. dissimilis (Ul).L.chrysocephalusKU786S(ll3), 12654 KU 15159 (1CS), 17235 (3CS); REJ 552 (2). (IOCS), 18440(1); LACM 39808-1 (5CS).L. E. insignis INHS 83108 (1:9CS); REJ 916 coccogenis INHS 75417 (1), 79254 (4CS); (20). E. monacha KU 19615 (9CS); KFW KU 8898 (7CS), 11468 (15CS), 12020 (26); 1102 (32); NB 783 (3); REJ uncat. (10); UL UAIC 6238.05 (26). L. cornutus KU 4035 7712 (19); UT 44.1031 (2CS). E. x-punctata (9CS), 7296 (26), 8686 (11CS), 12942 (38); KU2583 (3), 11279 (7), 16385 (1CS), 18439 LACM 39844-1 (3CS). L. pilsbryi KU 13009 (6CS). Exoglossum laurae KU 18922 (2CS), (5CS). L. zonatus KU 12648 (123: IOCS), 18923 (5CS). E. maxillingua KU 18925 21197(1CS).L.2oaiw//^INHS75135(15CS); (3:12CS). Extrarius aestivalis KU 9654 (1), KU 18995 (5: IOCS); UAIC 2553.03 (111). 2311 (10), 12110 (8CS), 14217 (10), 17035 Lythrurus ardens KU 4136 (27:15CS), 5718 (2CS). Gila atraria KU 11922 (1CS). G. caer- (40), 11557 (7); UAIC 5115.02 (3). L. atra- ulea KU 18904 (15CS). G. conspersa INHS piculus KU 18887 (IOCS), 18976 (3CS). L. 75307 (12CS). G. copei KU 11819 (1CS). G. bellus KU 14511 (15CS), 17377 (18); LACM orcutti INHS 76640 (19). G. robusta KU 3249 39811-1 (IOCS). L. fumeus INHS 87307 (5). Hemitrema flammea KU 18884 (IOCS), (IOCS); KU6244 (8CS).L. lirus INHS 75410 18931 (14). Hesperoleucus symmetricus KU (20); KU 18933 (15CS), 18986 (15); UAIC 8 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

647.07 (18), 653.05 (16), 4757.08 (14). L. N. chlorocephalus INHS 75049 (3: 16CS); KU roseipinnis KU 16857 (63: 12CS), 16909 (50). 18440(1), 18982(15).A>. chrosomus KU 16903 L. umbratilisKU1190(78),9547(4CS), 11275 (10),18849(10CS),18890(15);UAIC2574.01 (28), 15521 (15CS), 15550 (32), 15736 (9). (8), 4946.07 (7). N. cummingsae KU 5258 Macrhybopsisgelida KU 6862 (1), 9687 (2CS), (25:15CS); LACM 39832-1 (IOCS); UAIC 8111 (1CS), 1888 (2CS), 17360 (5), 19775 1618.02 (111). N. edwardraneyi KU 18891 (1CS).M. meeki KU 7326 (1CS), 9688 (3CS). (16CS). N. emiliae KU 14316 (18), 18855 M. storeriana KU 2173 (4CS), 12104 (5CS), (15:10CS), 18984 (12CS); LACM 39835-2 12126 (1CS), 14218 (14), 19896 (8CS); TU (3CS); RDS 4471 (2CS). N. girardi KU 3245 42770 (2CS), 45301 (2CS). Nocomis bigut- (127), 3959 (124), 8039 (14CS), 8572 (174). tatus KU 7295 (21). N. effusus KU 18932 N. greenei KU 10085 (114), 10766 (3), 12986 (IOCS). N. leptocephalus KU 12740 (1CS), (15CS). N. harperi KU 16849 (18), 19690 18858 (3:9CS). N. platyrhynchus KU 18926 (1CS); LACM 39812-1 (12CS); TU 2316 (15CS). N. raneyi INHS 74357 (5CS); KU (3:12CS),38202(2CS);UAIC941.08(37).M 19616 (5:4CS). Notemigonus crysoleucus KU heterodon KU 4590 (43), 10320 (1), 14255

1357 (2:4CS), 10775 (1CS), 13101 (4), 18938 ( 1 5CS); LACM 39850- 1 (6CS). A>. heterolepis (1CS), 19632 (4). Notropis aguirrepequenoi KU 3436(17), 11296(14CS), 12275(3), 18626 INHS 75330 (6CS);UMMZ 97399 (15: IOCS). (12).M/w&w/KU19039(2CS),19350(6CS).

N. altipinnis INHS 86955 (6CS); KU 18974 N. hudsonius KU 14257 ( 1 5CS), 20042 (4CS). (12CS), 19783 (10), 20040 (5CS); LACM N. hypsilepis KU 8807 (7:3CS); LACM 39845- 39837- 1 (6CS), 39837-3 (4CS); TU 7343 (163). 1 (7CS); UAIC 1235.04 (35). N. jemezanus N. amabilis KU 6008 (35), 15444 (12), 17023 KU 8066 (14: 12CS), 8073 (88), 8364 (10); TU (15:15CS).A>.0m^w«INHS77856(5:15CS); 11334 (&5).N.leuciodusKU 12022(1), 18850 KU 18975 (15), 19784 (2CS). N. anogenus (IOCS); LACM 39809-1 (IOCS); UAIC UMMZ 183967 (5CS). N. asperifrons KU 5998.05 (27), 6000.04 (7). N. lutipinnis KU 18886 (11:20CS), 20165 (1); LACM 39843-1 5314(15CS);LACM39818-1(14CS),39851- (9CS); TU 32744 (29). N. atherinoides KU 1 (2CS), 39852-2 (2CS), 39852-3 (2CS). N. 4138 (7CS), 10996 (4), 16494 (7), 18935 maculatusKU 15511 (13:12CS), 16906(25); (56:15CS);LACM39836-1(12CS);RDS4471 LACM 39846-1 (IOCS); UAIC 1599.01 (93). (4CS). N. atrocaudalis INHS 76980 (8), 87782 N. mekistocholas KU 20043 (1CS). N. nazas

(4); KU 6106 (2:12CS), 6073 (26), 6061 (4), INHS 84099 (5:15CS). N. nubilus KU 7286 20195 (14). N. baileyi KU 14510 (15CS), (14CS), 17189 (14), 19089 (16). N. 7 17375 (12); LACM 39849-1 (6CS). N. bairdi ortenburgeriKU 12962 (5: IOCS). A , oxyrhyn- KU 18473 (166), 18474 (34:11CS). N. blen- c/w.?KU2316(32),6028(3:15CS), 14288(9); nius KU 5130 (1CS), 8242 (36), 9640 (22), TU 19687 (8), 20228 (13). N. ozarcanus KU

9665 (32), 16045 (15CS). N. boops KU 6169 7664 (9CS), 7725 (28), 12656 (9), 15163 (16). (18CS), 14424 (115), 15873 (7), 19680 (10), N. perpallidus INHS 81047 (6CS). N. peter- 21195 (1); LACM 39820-1 (12CS), 39820-3 somKU 14072(56), 17161 (13L15CS), 17886 (3CS). N. braytoni TNHC 4457 (5:1 5CS); TU (9); LACM 39825-1 (2CS), 39825-2 (8CS), 87158 (15). N. buccula KU 14286 (8:12CS). 39838-1 (IOCS), 39841-1 (6CS). N. photo- N. buchanani KU 2313 (15), 3084 (4), 15031 genisWRS, 77121 (5:15CS); KU 5805 (6). A>. (23). N. candidus KU 18889 (10: IOCS); TU potteriKU 14222 (302:20CS); LACM 39839- 80042 (8:12CS). N. chalybaeus KU 9405 1 (IOCS). N. proem INHS 77857 (5:15CS); (58:15CS), 9568 (46), 19689 (4); LACM KU 16907 (5), 18877 (25). N. rubellus KU 39822-1-6 (26CS), 39829-1 (IOCS), 39855-1 7888 (53), 11030 (15CS), 12086 (35), 21196 (8CS). N. chihuahua KU 4999 (17CS), 8406 (3); LACM 39834- 1 (9CS), 39835- 1 (5CS). N.

(17). N. chiliticus INHS 74150 (7); KU 16902 rubricroceus KU 3290 (7CS); UAIC 5990.01 (10), 18981 (3: 12CS); LACM 39847-2 (IOCS). (10). N. saladonisFMNH 62160 (1 1).^. scab- NORTH AMERICAN MINNOW GENUS CYPRINELLA riceps KU 16911 (5), 18862 (2:13CS), 18990 18929 (1L6CS), 18930 (2); REJ 971 (1); NB

(5: IOCS). N. scepticus INHS 76886 (18CS); 746 (1). P. uranops KU 19619 (2CS); REJ 982 KU 5268 (6); LACM 39837-2 (IOCS). N. (2). Phoxinus cumberlandensis KU 18934 1 semperasper INHS 27260 (4CS); KU 18992 (8CS).P. eosKU 12255 (IOCS)./ , erythrogas- (1:9CS), 19617 (4CS). N. shumardi KU 9623 ter KU 5539 (36), 13198 (3CS). P. neogaeus (4), 14290 (14CS). N. simus KU 8067 KU 18882 (12CS). P. oreas KU 3259 (6CS),

(57:16CS); UMMZ 125064 (4CS). N. spec- 3275 ( 14). PimephalesnotatusKU 8003 (5CS), trunculus KU 5473 (9CS); TU 29229 (5), 12579 (3CS). P. promelas KU 12106 (4CS). P. 32807 (69); UAIC 2557.06 (8). N. sp. cf. tenellusKXJ 12127 (1CS). P. vigilaxKV 16388 spectrunculus SIUC 3536. (2); UAIC 2973.14 (1CS). Plagopterus argentissimus KU (48). N. stilbius KU 8770 (6: IOCS), 169 12 (5). 11932(3CS). Platygobio gracilis KU 3409 N. stramineus KU 8582 (233), 12288 (17), (5CS), 835 1 (13CS), 1 1950 (1CS), 14595 (35), 19011 (7CS), 21073 (12CS). N. telescopus 17146 (5CS). Pteronotropis euryzonus KU KU 11505 (1), 12653 (54), 15231 (19CS).M 19233 (IOCS). P. hypselopterus KU 16905 texanus KU 9612 (34), 17977 (69:15CS); (10), 17401 (IOCS), 19691 (5); LACM 398 14- LACM 39826-1 (3CS), 39826-2 (8CS), 39840- \(\2CS).P.signipinnisKJJ \6%5%{59\\5CS). 1 (6CS);RDS4471 (3CS).N. topekaKU4l9S P. welaka INHS 79413 (5:15CS); KU 16913 (50), 18682(19), 12289(17), 13136(7), 18759 (5CS), 18895 (10CS);LACM39817-1 (IOCS); (6), 19033 (15CS); UMMZ 163223 (1). N. TU 25785 (368). Ptychocheilus grandis KU tropicus TU 43597 (3: 12CS). N. uranoscopus 1 8920 (\2CS). P. umpquaeKVX 8908 (2:8CS). KU 18894 (18CS);TU 29893 (93). N. volucel- Rhinichthys atratulus KU 2779 (6), 4145 (2). lus KU 8456 (33), 9462 (16), 17025 (16CS); R. cataractae KU 11845 (16), 12264 (21). R. RDS4471 (6CS).N.xaenocephalusKU 18854 evermanni KU 18910 (15CS). R.falcatus KU

( 10: IOCS), 1 8896 (20); LACM 39830- 1 (2CS), 18911 (3CS), 18912 (8CS). Richardsonius 39830-2 (8CS), 39848-1 (IOCS); TU 25956 egregius KU 11788 (3CS), 12442 (23). Semo- (11). OregomchthyscrameriKU 18905 (3CS), tilus atromaculatus KU 12094 (3CS), 12442 18906 (4CS), 18907 (3CS). Phenacobius ca- (23), 12094 (3), 16527 (6), 18921 (2CS). S. tostomusKU 18897 (7CS), 18898 (5CS); REJ corporalis KU 18856 (IOCS). S. margarita uncat. (2). P. crassilabrum KU 19618 (2CS); INHS 83643 (5CS), 19000 (IOCS). Yuriria Re 466 (2). P. mirabilis KU 7918 (5CS), alta INHS 75315 (20CS). 15151 (3CS), 16387 (1CS). P. teretulus KU

MONOPHYLY OF CYPRINELLA AND RELATIONSHIPS TO OTHER NORTH AMERICAN CYPRINIDS

Based on results of this study and those of other large monophyletic groups (Fig. 1). One Coburn (1982, 1983), many eastern North clade (Fig. 1, Suite 2) includes the genera American cyprinids belong to a single mono- Hybognathus, Exoglossum, Dionda, phyletic group, all sharing the derived charac- Campostoma, Nocomis, Platygobio, ter of an open posterior myodome (Fig. 1, Macrhybopsis, Extrarius, Phenacobius, and Suite 1). The genus Cyprinella is embedded in Erimystax (Figs. 1-3). A second clade (Fig. 1, this clade. Within this large group of cyprinids Suite4) includes species of the genus Hybopsis, with an open myodome is a large polytomy of Ericymba buccata, and species previously re- species and monophyletic groups. Included ferred to Notropis: H. boucardi, H. calientis, are species of Notropis (except Aztecula sal- H. dorsalis, H. longirostris, H. sabinae, H. lei), and the genera Cyprinella, Luxilus, alborus, and H. bifrenatus (Figs. 1, 4). The 1 Suite includes the Lythrurus, Pteronotropis, Oregonichthys, third clade (Fig. , 5) genera Clinostomus, Richardsonius, Yuriria, and three Pteronotropis, Cyprinella, Luxilus, Lythrurus, 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Notropl*

Fig. 1. Hypothesized phylogenetic relationships of North American cyprinids with an open posterior myodome. Other cyprinids examined in this study are not included in this clade. Numbered characters suites are discussed in text.

andNotropis. Within this third clade, the genus burnops (Fig. 5), the Notropis volucellus spe- Pteronotropis forms the sister group to the re- cies group (Fig. 6), the subgenus Hydrophlox, maining species and species groups (Fig. 1, and a monophyletic group inclusive of the Suite 7). Above Pteronotropis, a clade com- subgenus Notropis and the Notropis texanus posed of the genera Lythrurus, Luxilus, and species group. Cyprinella forms the sister group to the Notro- Species of uncertain relationships and in- pis clade, composed here of four major groups volved in this multichotomy between charac- whose interrelationships are unresolved at this ter Suites 1 and 5 in Fig. 1, but not illustrated time. These groups include the subgenus Al- are Notropis topeka, N. atrocaudalis, N.

vQ 9x vo o o o , A° -C ^ r° .<* •.* .* v^ NT *° ^

Cyprinids with an open posterior stramineus, N. chihuahua, N. procne, and N. MYODOME mekistocholas. All of these species have an (Fig. 1, Suite 1). open myodome, but lack the derived charac- The open floor of the posterior myodome ters for stem 5 (Suite 5). (Fig. 7), first noted and discussed by Coburn (1982, 1983), is unique to this group of cyp- Justifications for these particular statements rinids and is considered a synapomorphy of of relationship and the numbered character these species. The opening is oval to elongate suites presented in Fig. 1 are outlined below. and slitlike and is bounded by both the para- Because European and some North American sphenoid and basioccipital. Most species of species were not included in the present study, this group have the opening as juveniles and and because these results and those of Coburn adults. However, in the genera Nocomis, Cam- (1982, 1983) represent some of the first at- pos toma, and Dionda and in the species tempts at revealing higher relationships of Platygobio gracilis, Hybopsis boucardi, and these cyprinids, only a few formal taxonomic H. bifrenatus the opening exists only in juve- changes are proposed. However, in some cases niles and is fused shut in adults (Fig. 8). Only the existing classification is a poor representa- occasionally will an adult of these species tion of our knowledge of cyprinid interrela- retain the opening. The secondary closure of tionships, and necessitates change. For ex- this opening is considered derived, occurring ample, the obviously unnatural genera Notro- independently in Platygobio, Hybopsis pis and Hybopsis are each broken up into boucardi, H. bifrenatus, and the Nocomis- separate genera, elevating subgeneric names Campostoma-Dionda clade. Other North to generic level. Relationships of some species American cyprinids examined apparently never of Notropis remain problematical. For now, develop this opening in the posterior myo- these species are retained in "Notropis" but dome as juveniles or adults (Fig. 7A). with unknown affinities. A similar opening in the posterior myod-

Phenacobius Er imy s t a x

Fig. 3. Hypothesized phylogenetic relationships of species in the North American chub genera Erimystax and Phenacobius. Lettered character suites are discussed in text. Suite L corresponds to L in Figure 2. 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

,t> .

s* o* * * » ^

*" -*> -* 5, N « . * D .»

Fig. 4. Hypothesized phylogenetic relationships of species in the North American genus Hybopsis and the related genus Ericymba. Lettered character suites are discussed in text. This clade corresponds to group defined by suite 4 in Figure 1. ome was noted by Howes (1980) for the baril- ever, does not appear to be homologous with ine group of minnows. The open condition ob- that found in the clade of eastern North Ameri- 2 1 can minnows. In bariline the served by Howes (1 980: figs. , 22, 23), how- cyprinids open-

Lettered Fig. 5. Hypothesized phylogenetic relationships of shiner species in the subgenus Alburnops (genus Nolropis). character suites are discussed in text. NORTH AMERICAN MINNOW GENUS CYPRINELLA 13

N * ;s* N _^ .<* N

of in the volucellus Lettered character Fig. 6. Hypothesized phylogenetic relationships minnows Notropis species group. suites are discussed in text.

is located in ing is located posteriorly in the myodome and myodome the opening centrally is both is not bounded by the basioccipital posteriorly. the floor of the myodome and bounded The opening is only contained anteriorly by by the basioccipital posteriorly and parasphe- the parasphenoid. In the eastern North Ameri- noid anteriorly. can cyprinids with the opening in the posterior Chub clade (Fig. 1, Suite 2). Included in

PRO

in some taxa. Fig. 7. Basicranial region in some North American cyprinids illustrating open posterior myodome A) 48 lotichthys phlegathantis, 33 mm, KU 1 1935. B) Pimephales notatus, 43 mm, KU 8003. C) Macrhybopsis storeriana, mm, KU 12104. D) Cyprinella galactwa, 57 mm, KU 12029. Open posterior myodome designated by stippled pattern. Horizontal bar equals 1 mm. 14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 8. Developmental closure of the open posterior myodome in Campostoma anomalum. A) 26 mm, KU 3946. B) 43 mm, KU 3946. C) 70 mm, KU 13200. Distribution of opening in posterior myodome designated by stippled pattern. Horizontal bar equals 1 mm. this group are the genera and species Agosia, preethmoid. The primitive preethmoid-pala- Hybognathus, Exoglossum, Nocomis, Campo- tine articular surface on the palatine is a simple stoma, Dionda, Platygobio, Macrhybopsis and well-developed socket developed anter- meeki, M. gelida, M. storeriana, Extrarius omesially (Fig. 10A, B) into which fits the aestivalis, Phenacobius, and Erimystax (Fig. preethmoids. In the chub clade, the articular 2). All of these species share two derived char- surface is a groove that extends anteroven- acters of the palatine and one of the urohyal. trally to posterodorsally. It is unrestricted both Three types of maxillary processes of the pala- anteriorly and posteriorly (Fig. 9A-T). tine were noted among the cyprinids exam- Primitively the horizontal plate of the ined. In the primitive condition the maxillary urohyal is forked posteriorly (Fig. 1 1 A). In all process is directed laterally, perpendicular to taxa of the chub clade the posterior margin of the anteroposterior axis of the palatine (Fig. the urohyal is oval and smooth (Fig. 11E-H). 10E); this morphology occurs in most species Two derived characters support the mono- with an open myodome and in all outgroups. In phyly of all members of this chub clade, exclu- the species listed above the maxillary process sive of Agosia (Fig. 2, Suite A). In all species of the palatine is elongate, straight, and di- except Agosia, the endopterygoid and met- 12 rected anteriorly (Fig. 9A-T). apterygoid are laterally convex (Fig. A). The second derived characteristic of the The laterally convex endopterygoid and met- palatine for the chub clade involves the anter- apterygoid are unique to this group of cyp- omesial socket for articulation with the rinids. In all outgroups examined these bones NORTH AMERICAN MINNOW GENUS CYPRINELLA 15

Fig. 9. Dorsal Qtil) and mesial (right) views of palatines of some North American cyprinids with a derived anterior preethmoid articular morphology. Up is anterior. A) Agosia chrysogaster, 65 mm, KU 8084. B) Exoglossum laurae, 56 mm, KU 18922. C) Nocomis effusus, 36 mm, KU 18932. D) Campostoma anomalum, 64 mm, KU 13200. E) Aztecula sallei, 56 mm, UMMZ 191695. F) Hybognathus placitus, 51 mm, KU 12597. G) Dionda episcopa, 57 mm, KU 16891 .

H) Extrarius aestivalis, 32 mm, KU 12110. I) Platygobio gracilis, 75 mm, KU 17146. J) Macrhybopsis storeriana, 46 mm, KU 12104. K) Phenacobius mirabilis, 61 mm, KU 16387. L) P. teretulus, 68 mm, KU 18929. M) P. catostomus, 55 mm, KU 18897. N) P. crassilabrum, 53 mm, KU 19618. O) P. uranops, 86 mm, KU 19619. P) Erimystax monacha, 66mm, UT 44. 1031. Q)£. cahni, 61 mm.KU 19614. R)E.x-punctata,70mm,K.U 16385. S) E. insignis, 73 mm, INHS 83108. T) E. dissimilis, 55 mm, KU 17235. Horizontal bar equals 1 mm. 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 10. Morphological variation m palatmes of some North American cyprinids. Up is anterior. A) Notropis potteri (medial view), 50 mm, KU 14222. B) N. bairdi (medial view), 54 mm, KU 18474. C) N. chihuahua, 51 mm, KU 8406.

D)N. leuciodus, 62 mm, KU 18850. E) Dorsal (left) and medial (right) views of palatine of N. hoops, 56 mm, KU 6169. F) N.braytoni, 41 mm,TNUC 4451. G)N.shumardi,52mm,KU 14290. H)N. candidus, 68mm,KU 18889. T)N.stilbius, 57 mm, KU 8770. J) N. photogenis, 63 mm.INHS 77121. K)N.scabriceps, 67mm, KU 18862. L) N. amoenus, 60 mm, INHS 77856. M) N. sceplicus, 65 mm, INHS 76886. N) N. hypselepis, 53 mm, KU 8807. O) N.jemezanus, 48 mm, KU 8066. P) N. oxyrhynchus, 46 mm, KU 6028. Q) N. altipinnis, 45 mm, KU 1 8974. R) Pteronotropis signipinnis, 50 mm, KU 16858. S) P. euryzonus, 4Smm,KV 19233. T)P hypselopterus, 45mm,KU 17401. U) P. welaka, 50mm, KU 18895. Horizontal bar beneath A is 1 mm for A-D. Bars beneath other species represent 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 17 are strongly concave laterally, being dished- unique. Gut coiling also occurs in Notropis out posterior to the palatine-metapterygoid ar- nubilus and N. mekistocholas, but the gut coil- ticulation. The second derived feature of this ing is different in these taxa and, with refer- clade, exclusive of Agosia, is the elongate ence to higher relationships (Fig. 1), gut coil- maxillary process of the palatine. The process ing is independently evolved in these species. is short in Agosia (Fig. 9A) and long in other The pharyngeal pad of these two genera (Fig. members (Fig. 9B). The elongation of the 15) is oval and flattened and has an anteriorly anteriorly directed maxillary process of the directed process. Outgroups, except Hybog- palatine is thus considered a further nathus, typically have a triangular pad that is modification of the ancestral condition for the concave ventrally. Species of Hybognathus chub clade. have the derived oval pharyngeal pad and an- Above Agosia there are four monophyletic terior process (Fig. 13; Bailey and Allum, groups of unresolved relationships (Fig. 2, 1962), also supporting a close relationship of Suite A). Species of the genus Hybognathus this genus to Campostoma and Dionda. Fur- (Fig. 2, S uite C) share an expanded pharyngeal thermore, the anterior margin of the pad of process of the basioccipital not found in any most outgroups is blunt, lacking any type of other cyprinids (Fig. 13). Species of the genus process. Phoxinus erythrogaster, P. cumber- Exoglossum (Fig. 2, Suite B) share a derived landensis, and P. oreas have a small anterior condition of the dentaries and palatines. In process like that found in Nocomis and Di- these two species the dentaries are very deep onda. Species of Phoxinus, however, lack an and twisted anteriorly such that the rami are open posterior myodome. horizontal and parallel (Fig. 14). The palatines The synapomorphies of Nocomis (Fig. 2, of Exoglossum have a unique morphology. In Suite H) and species relationships are pre- these species a deep and wide supraethmoid sented by Mayden (1987). Synapomorphies process wraps around the preethmoids (Fig. for the genus Campostoma (Fig. 2, Suite G) 9B). include a coiled gut whose coils wrap around The clade inclusive of the genera Nocomis, the gas bladder (Burr, 1976) and the loss of a Campostoma, and Dionda (Fig. 2, Suite D) all supraethmoid process on the palatine (Fig. share the derived secondary closure of the 9D).SpeciesofthegenusD/<9Aufo(Fig.2,Suite opening in the posterior myodome (Fig. 8) and F) share a tubercle pad on the subopercle that very large cephalic tubercles. The closure of is developed only in breeding males. A similar, the myodome is discussed above. The cephalic but convergent structure occurs in the tubercles of these three genera are very large Phenacobius-Platygobio lineage (see below). and erect (Lachner and Jenkins, 1971a, 1971b; Because of the different locations of the pads Burr 1976) and are considered a synapomor- in the two groups and morphological dissimi- phy of this group. All other cyprinids exam- larities in the pads they are presently not con- ined have small cephalic tubercles compared sidered homologous and are thus convergent. with these three genera. The fourth lineage in the chub clade is one The genera Campostoma and Dionda (Fig. inclusive of the genera Platygobio, 2, Suite E) share a reduced or absent ascending Macrhybopsis, Extrarius, Erimystax, and process condition on the premaxilla (Fig. 15), Phenacobius (Fig. 2, Suite I). These species all a coiled gut (Burr, 1976; Hubbs and Miller, share: 1) a nuptial pad on the interopercle and 1978), and a derived pharyngeal pad on the ba- preopercle unique to this group, 2) a short and sioccipital (Fig. 15). The reduced or absent triangular basihyal (Fig. 16H-I), 3) a single ascending process on the premaxilla is appar- articular surface for the hyomandibular on the ently unique to this clade. All other cyprinids metapterygoid (Fig. 16E, F), and 4) enlarged examined have an obvious ascending process. sensory papillae on interradial membranes of The coiled gut of these two genera is not the pectoral fin. Coburn (1983) noted the sec- 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 11. Urohyals of some North American cyprinids. Up is anterior. A) Clinoslomusfundxdoid.es, 60 mm, KU 10697. B) Hybopsis amnis, 52 mm, KU 18847. C) H. winchelli, 51 mm, TU 39705. D) H. sabinae, 41 mm, KU 6237. E) Macrhybopsis storeriana, 94mm,KU 12126. ¥)M. meeki, 56mm,KU 9688. G) Agosia chrysogaster, 65 mm, KU 8084. H) Campostoma oligolepis, 85 mm, KU 12520. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 19 BO

MPT

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Fig. 13. Ventralviewof basicranial region of Hybognathus nucnalis (49 mm, KU 14123) illustrating expanded pharyngeal process of basioccipital. Up is anterior. Horizontal bar equals 1 mm.

ond and third derived characters for this group and suggested other derived features in addition to these. In all outgroups the metapterygoid has a double articular surface with the hyomandibular, one posteriorly and one posteroventrally (Fig. 16A-C). The basihyal of most other cyprinids is elongate and narrow

(Fig. 16G). The nuptial pad and sensory papillae in this group are unique. Interradial sensory papillae of the pectoral fins are small in outgroups. Controversial in this group is the placement of monacha in the genus Erimystax. This species has also been closely allied to the genus Cyprinella. Some similarities do exist between E. monacha and some species of Cyprinella (see section below on Monophyly of Cyp- rinella). Whether or not these shared similarities are the result of common ancestry or convergence warrents further investigation. At present, I consider monacha to be a member of the genus Erimystax, based on the hy- Fig. 12. Pterygoid series of some North American pothesized synapomorphies listed above and cyprinids illustrating concave (A) and convex (B-E) for but for the chub lateral surfaces of endopterygoid and metaplerygoid bones. below not only Erimystax, Right is anterior. A) Notropis atherinoides, 53 mm, KU clade as a whole. 18935. Nocomis 53 mm, KU 19616. C) B) raneyi, The Macrhybopsis and Platygobio clade Erimystax insignis, 73 mm, FNHS 83 108. D) E. monacha, 2, Suite J) is a single pig- 66 mm, UT 44.1031. E) Phenacobius crassilabrum, 53 (Fig. supported by the caudal fin and a mm, KU 19618. Horizontal bar equals 1 mm. mentation character of 20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY urohyal character. In all of these species the gnathic ramus (Fig. 17), 3) a short ascending lower lobe of the caudal fin is darkly pig- arm on the preopercle and elongate horizontal mented except for the lowermost ray which is arm, and 4) the mesial edge of pharyngobran- depigmented. This color pattern is unique to chial 3 is notched. Outgroups and other these four species and is derived. In all of these members of the chub clade typically have a species the ventral horizontal plate of the more delicate fourth Weberian rib, a short urohyal is strongly reduced to a very thin coronoid process and long gnathic ramus, lamella (Fig. HE, F). Primitively, this plate is subequal arms on the preopercle, and smooth well developed in cyprinids. mesial margin of pharyngobranchial 3. The genera Extrarius, Phenacobius, and The Phenacobius-Erimystax clade (Fig. 2, Erimystax form a monophyletic group (Fig. 2, Suite L) is monophyleuc, supported by five Suite K), all sharing the following derived fea- derived characters. Species of this clade all tures which were also noted earlier by Coburn have 1) a narrow caudal skeleton (Fig. 18E,F), (1983): 1) a heavy transverse process on the 2) a small flap of skin with taste buds, extend- fourth Weberian rib, 2) a dentary with a high ing from beneath the anterior end of the lacry- coronoid process and a short ventrally deflected mal bone to the posterior ramus of the maxilla

Fig. 14. Modified jaws of Exoglossum. A-D)E. laurae (47mm,KU 18923). A) Lateral view of right lower jaw. B)Lateral view of premaxilla. C) Lateral view of maxilla. D) Ventral view of lower jaws. E) Ventral view of lowerjaws of Hemitrema flanvnea (39 mm, KU 18884). Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 21

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Fig. 15. Characters supporting the sister group relationship of Campostoma and Dionda. Viewof basicranial region and premaxilla of Campostoma anomalum (53 mm, KU 13200) (A and B) and Dionda episcopa (58 mm, KU 16891) (C and oval D) illustrating pharyngeal pad and weakly developed ascending process of premaxilla. Horizontal bar equals 1 mm.

(Jenkins and Lachner, 1971; Jenkins, pers. outgroups is straight. The maxillary flap of comm.), 3) a ventrally deflected posttemporal skin of these species is unique to this group and cephalic canal, 4) a broad platelike second in- in most outgroups the second infraorbital is fraorbital bone, and 5) reduced tuberculation not very broad. The reduced tuberculation on on the dorsum of the head. The caudal skeleton the dorsum of the head is derived since other of all outgroups examined is much broader species of the chub clade and other cyprinids (Fig. 18A-D). Erimystax dissimilis, E. insig- typically have considerable tubercle develop- nis, E. cahni, E. monacha, Phenacobius ca- ment in this area. tostomus, P. teretulus, and P. uranops all have Monophylyofthe genus Phenacobius (Fig. a much compressed caudal and precaudal 3, Suite M) is supported by four characters: 1) skeleton, relative to any Cyprinella and other loss of maxillary barbel, 2) heavy dentary, cyprinids surveyed.The posttemporal canal of maxillary, and premaxillary bones (Fig. 1 7), 3) 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

as a secondary loss. The dentary, maxillary, and premaxillary of Phenacobius is very large and heavy, more so than most outgroups examined (Fig. 17). In most members of this chub clade the anteriorly directed maxillary process of the palatine is narrow (Fig. 9). In all Phena- cobius the maxillary process is broader (Fig. 9K-0). This morphology is considered a further modification of the narrow ancestral condition of the chub clade. The pectoral fin tuberculation pattern in this group is different from other cyprinids examined (Fig. 19A). Species of Phenacobius have three rows of tubercles

per ray, with the anteriormost row being the longest and extending from the base of the ray to near the distal margin. The posterior two rows are gradually shorter. In most outgroups the tubercle rows are symmetrical. Basally, a single row is formed and branches into two to four rows distally. Within Phenacobius, P. mirabilis is the sister to other members. All species except P.

mirabilis (Fig. 3, Suite N) have short anterior arms on the urohyal with mesially directed

of Fig. 16. Metapterygoids (A-F) and basihyals (G-I) some North American cyprinids. Hyomandibular-metapterygoid articular processes are identified by arrows. Right is anterior for A-F. Up is anterior for G-I. A) Nocomis raneyi, 53 mm, KU 19616. B) Notropis atherinoides, 53 mm, KU 1 8935. C) N. volucellus, 40 mm, KU 17025. D) Phenacobius crassilabrum, 53 mm, KU 19618. E) Erimystax insignis, 73 mm, INHS 83108. F) E. monacha, 66 mm, UT 44.1031. G) Couesius plumbeus, 76 mm, KU 1 8881. H) Erimystax insignis, 73 mm,

INHS 83108. I) E. monacha, 66 mm, UT 44.1031. Cartilage is represented by stippled pattern. Horizontal bar equals 1 mm.

a broad anteriorly directed maxillary process on the palatine (Fig. 9K-0), and 4) a unique tuberculation pattern on the pectoral fins of breeding males (Fig. 19 A). Within the clade

supported by Suite I of Fig. 2 all species except and maxilla Fig. 17. Modified lower jaw (upper) 0ower) Phenacobius have a maxillary barbel. The oi Phenacobius crassilabrum, 53 mm, KU 19618. Right 1 mm. absence of a barbel in this group is interpreted is anterior. Horizontal bar equals NORTH AMERICAN MINNOW GENUS CYPRINELLA 23

UN 3

NPU

Fig. 18. Caudal skeletons of some North American cyprinids. Left is anterior. A) Lythrurus fumeus, 51 mm, KU 6244. B) Luxilus coccogenls, 72 mm, INHS 79244. C) Cyprinella panarcys, 43 mm, UMMZ 208212. D) Cyprinella xaenura, 71 mm, KU 18994. E) Erimystax cahni, 58 mm, KU 19614. F) Phenacobius uranops, 82 mm, KU 19619. Horizontal bar equals 1 mm. 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

palatine, one anteriorly and one posteriorly, that surrounds the preethmoids (Fig. 9). Phena- cobius crassilabrum and P. uranops (Fig. 3, Suite P) share three derived features. These include: 1) a narrow pharyngeal pad on the basioccipital (Fig. 20B), 2) a broad shelf connecting the anterior and posterior mesial processes of the palatine (Fig. 9), and 3) a second osseous articular surface dorsally on the metapterygoid (Fig. 16D). The narrow pharyngeal pad is unique in the chub clade. As mentioned above, Phenacobius plus Erimystax, Extrarius, Macrhybopsis, and Platygobio have only a single cartilage articular surface on the metapterygoid. This condition is reversed in P. uranops and P. crassilabrum. The second process appears to be different in these two species, in that the dorsal process is ossified and a

cartilage tip is not present. In all outgroups with two metapterygoid processes the dorsal

process has a cartilage tip. Monophyly of the genus Erimystax (Fig. 3, Suite Q) is supported by two derived characters: 1) a unique "stellate" barbel (Fig. 19C) and 2) an elongate basihyal. The barbel in Erimystax has large, fleshy processes extend-

ing off of the tip, producing a shape similar to a spiked mace (Fig. 1 9C).The barbel ofErimys- tax monacha is more intermediate between the "stellate" morphology as seen in other species of Erimystax and the morphology of other barbeled members of the chub clade. The processes of the barbel in E. monacha are smaller and fewer than are typical in other members of the genus. Other barbeled cyprinids have a relatively smooth texture to the barbel. Moore (1950) and Branson (1962) il- Fig. 19. Characters supporting the monophyly of the lustrated this derived barbel, but did not com- generaPhenacobius and Erimyslax. A) Scanning electron elon- micrograph of pectoral fin tuberculation in Phenacobius ment on its systematic significance. The mirabilis KU the (61 mm, 15151) illustrating graduated gate basihyal of these species (Fig. 16H, I) is rows of tubercles on each ray. B) SEM of barbel of hypothesized to be a reversal to the more Platygobio gracilis (79 mm, KU 8351) illustrating the from the derived trian- simple primitive barbel morphology. C) Barbel of primitive morphology with Erimystax cahni (7 1 mm, REJ737) illustrating the derived gular basihyal uniting these species the condition of the "stellate" genus. genera Platygobio, Macrhybopsis, Phena- cobius, and Extrarius.

processes (Fig. 20). Phenacobius catostomus, Within Erimystax, a trichotomy presently P. crassilabrum, and P. uranops (Fig. 3, Suite exists between E. monacha, E. x-punctata, and O) have two mesially directed processes on the a clade inclusive ofE. cahni,E. insignis, and E. NORTH AMERICAN MINNOW GENUS CYPR1NELLA 25

c M PPBO

Fig. 20. Osteological characters of the genus Phenacobius. A) Basicranial region of P. mirabilis (60 mm, KU 16387) illustrating pharyngeal pad with small lateral processes. B) Basicranial region of P. crassilabrum (53 mm, KU 19618). C) Ventral view of urohyal of P. mirabilis (same data as above). D) Urohyal of P. teretulus (68 mm, KU 1 8929). E) Urohyal of P. uranops (84 mm, KU 19619). F) Urohyal oiP. catostomus (55 mm, KU 18897). G) Urohyal of P. crassilabrum (53 mm, KU 19618). Horizontal bar equals 1 mm. dissimilis. The latter three species (Fig. 3, Genus Pimephales (Fig. 1, Suite 3). Four Suite R) share a narrowly developed maxillary species are included in this group. Pimephales process of the palatine (Fig. 9Q, S, T). This notatus and P. promelas form a monophyletic morphology is hypothesized to be derived for group, sister to a clade composed of P. vigilax this group, reduced from the broader condition and P. tenellus. Justifications for these rela- typical of other members of the genus and the tionships are presented by Mayden (1987). chub clade. Erimystax insignis and E. dissim- Genus Hybopsis and related species (Fig. ilis (Fig. 3, Suite S) both have dark and distinct 1, Suite 4). Included in this clade (Fig. 4) are blotches along the flanks. This derived pig- the species traditionally placed in the subgenus ment pattern is unique to these species in the Hybopsis (Jenkins and Lachner, 1971) (except clade of cyprinids with an open myodome. storeriana see above), plus Hybopsis boucardi, Erimystax monacha may represent the sister H. calientis, H. dorsalis, H. longirostris, H. group to other members of the genus if one sabinae, H. alborus, H. bifrenatus, and Eri- considers the well -developed "stellate" barbel cymba buccata. An alternative explanation for morphology to be a synapomorphy of all this group of cyprinids is convergent evolu- members, exclusive of £. monacha. tion. Many of these species are benthic dwell- 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ing forms with somewhat similar habitat characteristics. Thus, it is possible that this group- ing may be artificial, due to convergent evolution of morphological characteristics for a similar benthic lifestyle. All of the species of this group share a deep premaxillary process on the maxillary (Fig. 21B,C). Outgroups typically do not have a deep premaxillary process (Fig. 21 A). Some species of the genus Cyprinella also develop a similar, but convergent deep process. Species of the Hybopsis clade also share two derived conditions of the palatine. As described above, outgroups typically have a distinct perpendicular maxillary process on the palatine. In members of this clade, inclusive of Hybopsis, the maxillary process of the palatine is somewhat L-shaped with a long anterolateral^ directed process (Fig. 22A). This process is not considered homologous with that found in the chub clade. In the chub clade the maxillary process of the palatine is directed anteriorly (Fig. 9B),butin the Hybopsis clade the process is directed anterolateral^. This anterolateral process is an independent modification of the Fig. 21. Lateral view of maxilla of some North American of perpendicular maxillary process typical cyprinids illustrating the deep premaxillary process of the outgroups. Hybopsis clade. A) Notropis bairdi, 50 mm, KU 1 8474. The second derived character from the B) Hybopsis calientis, 42 mm, TU 30695. C) H. amnis, 42 mm, KU 18847. Horizontal bar equals 1 mm. palatine for this clade involves the palatine- preethmoid socket. As mentioned above, the primitive condition of the palatine socket is a clades. well -developed depression anteromesially on The monophyly of the dorsalis group (Fig. the palatine. In members of \heHybopsis clade 4, Suite B) was recognized by Coburn (1982) the articular surface on the palatine for the and is supported by two derived characters: 1) preethmoid is a deep, quasi-socket and groove, a deep anterior notch in the mesethmoid-su- extending anteroventrally to posterodorsally, praethmoid and 2) an elongate and straight much like members of the chub clade. In the kinethmoid. Most outgroups have only a shal- Hybopsis clade, however, the socket is differ- low notch in the anterior mesethmoid, a condi- ent, being restricted posteriorly by a mesially tion considered primitive. Some members of directed shelf on the palatine (Fig. 22B). A the volucellus species group also possess a similar palatine morphology is found in the convergent deep anterior mesethmoid notch. volucellus species group (Fig. 1) (see below). The kinethmoid of most outgroups is short and Within the Hybopsis clade, two less inclu- robust, unlike that found in the dorsalis group. sive monophyletic groups are supported; the Species united by Suite C (Fig. 4) share a dorsalis species group and a clade inclusive of derived, very deep maxilla (Fig. 23B-D). This the genus Hybopsis, H. boucardi, and H. cal- condition is nearly unique to this group. Out- ientis (Fig. 4). All other members are involved groups typically have a narrow isthmus on the in an unresolved polytomy with these two maxilla separating the ascending process from NORTH AMERICAN MINNOW GENUS CYPRINELLA 27

Fig. 22. Dorsal Qeft) and medial (right) views of palatines of some North American cyprinids with a derived anterior preethmoid articular morphology. Up is anterior. A)Hybopsis hypsinotus, 49 mm, KU 18968. B) H. amnis, 57 mm, KU 18847. C)//.amW.y,65mm,KU6318. D) H. winchelli, 54 mm, TU 39705. E)//. lineapunctata, 52 mm, KU 18885. F)Notropis heterolepis, 51 mm, KU 11296. G) Hybopsis labrosa, 55 mm, INHS 88319. H) //. zanema, 52 mm, INHS 88320. I) H. longirostris, 47 mm, KU 17965. J) //. sabinae, 41 mm, KU 6237. K) //. rubrifrons, 75 mm, KU 8829. L) Notropis maculatus, 50 mm, KU 1551. M) N. emiliae, 53 mm, KU 18855. N) N. ozarcanus, 45 mm, KU 7664. O) N. spectrunculus, 41 mm, KU 5473. P) N. volucellus, 41 mm, KU 17025. Q) N. aguirrepequenoi, 41 mm, UMMZ 97399. R) Hybopsis calientis, 4 1 mm, TU 30695. S) Notropis tropicus, 29 mm, TU 43597. T) Hybopsis boucardi, 5 1 mm, UMMZ 191686. Horizontal bar equals 1 mm. 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the premaxillary-rostral process of the maxilla. A similar, but convergent maxillary morphology is also seen in a few species of Cyp- rinella.

All species united by Suite D (Fig. 4) share six derived characters: 1) mandibular canal attached to the retroarticular (Fig. 24B-D), 2) long interhyal, 3) enlarged first pair of hypobranchials, 4) elongate first basibranchial, 5) broad posterior parasphenoid (Fig. 25C, D), and 6) an oval orbit. The presence of the mandibular canal crossing the retroarticular, also discussed by Coburn (1982), and the broad posterior parasphenoid in these nine species are unique. Mostoutgroups from within and outside the clade defined by an open posterior myodome have a short interhyal, short first basibranchial, and a small first pair of hypobranchials. Thus, these three conditions are hypothesized to be derived for the group. As seen in live specimens, all species ofHybop- sis, except H. hypsinotus, have an oval orbit and eye. This characteristic is apparently unique to this group. The Hybopsis labrosa-H. zanema species group (Fig. 4, Suite E) is supported by four characters. These are: 1) enlarged and antrorse tubercles on the dorsum of the head, 2) enlarged and antrorse predorsal tubercles, 3) large and slightly antrorse tubercles on the caudal peduncle, and 4) heavily pigmented membranes of the dorsal fin. These four characters are not unique to this group. Similar structures and color patterns occur in the genus Fig. 23. Lateral view of maxilla of some North American Cyprinella, a group sometimes thought to be cyprinids illustrating deep isthmus in some species of related closely (Jenkins and Lachner, 1971). Hybopsis. Right is anterior. A) H. sabinae, 43 mm, KU The dorsal head tubercles and predorsal tu- 6237. B) H. boucardi, 57 mm, UMMZ 191686. C) H. 54 mm, KU 18885. D) H. hypsinotus, 49 bercles appear to be similar between these two lineapunctata, mm, KU 18968. Horizontal bar equals 1 mm. groups, but the caudal peduncle tubercles and the dorsal fin pigment patterns are different. The caudal peduncle tubercles of the C. tributed in all membranes and not just the pyrrhomelas and C. nivea species groups of posterior membranes as is typical of species of Cyprinella are located in the center of each Cyprinella thought to be allied to the H. labrosa scale. Those of the H. labrosa group may be group. The sister group relationship between single and centrally located, but are generally H. zanema and the undescribed "thinlip" chub two per scale, one dorsally and one ventrally. (Fig. 4, Suite F) is supported by a single The dorsal fin of the H. labrosa group also derived character. Both species have very differs from Cyprinella in having melanin dis- elongate anterior urohyal processes. Most NORTH AMERICAN MINNOW GENUS CYPRINELLA 29 outgroups, especially other Hybopsis, have short processes. The monophyly of the remaining species of Hybopsis (Fig. 4, Suite G) is supported by 1) a truncated posterior margin of the horizontal 1 1 later- plate of the urohyal (Fig. B , C) and 2) ally directed and pointed processes on the urohyal near the neck, before the urohyal forks anteriorly (Fig. 11B, C). The truncated horizontal urohyal plate rarely occurs outside of this group, occasionally seen in some species of No tropis. The lateral processes seen on the neck of the urohyal are also found in Macrhybopsis storeriana, some specimens of species oiPhenacobius, and about 40% of the specimens of Notropis hudsonius. Based on the distribution of other characters, the occurrence of these processes in all these groups is considered convergent. The Hybopsis amblops-winchelli-amnis clade (Fig. 4, Suite H) is supported by a posteriorly directed supraethmoid process on the palatine (Fig. 22). This character is unique to this group. Outgroups have an anteriorly directed supraethmoid process. Hybopsis lineapunctata is a member of this group, but the supraethmoid process in this species is oriented perpendicular to the axis of the palatine. This condition is hypothesized to be a modification of the posteriorly directed process since this species, //. amnis, and H. winchelli form a 24. Lower of some of North American monophyletic group (Fig. 4, Suite I) with Fig. jaws species cyprinids illustrating the positions of the mandibular unresolved relationships. These three species canal relative to the retroarticular bone. Right is anterior. share the derived condition of a well- having A) Macrhybopsis storeriana, 94 mm, KU 12126. B) developed depression on the palatine anter- Hybopsis amnis, 42 mm, KU 1 8847. C) H. hypsinotus, 49 KU 18968. H. 52 INHS 88320. omesially, between the supraethmoid and mm, D) zanema, mm, Horizontal bar equals 1 mm. maxillary processes (Fig. 22).

1 Suite Pteronotropis-Notropis clade (Fig. , 5). The clade of cyprinids supported by stem 5 the hyomandibular (Fig. 26A). Thus, the is composed only of certain species of the pre- smooth wing condition is hypothesized to be viously conceived genus Notropis. Although derived within the clade defined by an open variation is known to occur within some of the myodome. Exceptions to the smooth included groups, most of these species have a hyomandibular wing inside this clade include smooth and straight margin on the anterior Pteronotropis signipinnis, some species of the wing of the hyomandibular (Fig. 26B-E). genus Cyprinella (Fig. 26F), some members Almost all outgroups, below stem 5 and out- of the Notropis volucellus group, and some side the group defined by an open myodome, species in the subgenus Notropis (Coburn, have an obvious notch in the anterior margin of 1982). 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Genus Pteronotropis (Fig. 1, Suite 6). These based on four derived characters. All three four species (P. signipinnis, P. hypselopterus, species share: 1) a very broad lateral stripe, 2) P. euryzonus, P. welaka) all share a derived very bright orange, red, and blue colors, 3) a condition of the preethmoid socket of the pala- palatine with a long and rounded shaft (Fig. 10 tine. In the primitive condition the socket is R-T), and 4) mandibular tubercles with tips generally deep. In these four species the socket curled dorsally. These are the only species is shallow (Fig. 10 R-U). Other species with a found in this study to have a rounded palatine convergently shallow preethmoid articular shaft. Pteronotropis welaka and outgroups have surface include Lythrurus ardens, Notropis a more compressed shaft. Within the genus, scabriceps,N. altipinnis, Cyprinella trichrois- Pteronotropis euryzonus is the sister to P. tia, and C. gibbsi. signipinnis, and these two species form the Pteronotropis signipinnis, P. hypselopterus, sister to P. hypselopterus. The sister relation- and P. euryzonus form a monophyletic group ship of P. euryzonus and P. signipinnis is

Fig. 25. Parasphenoids of some North American cyprinids. Up is anterior. A) Notropis atherinoides, 53 mm.KU 18935. B)Cyprinellatrichroistia, 60mm, KU 18853. Q Hybopsis zanema, 52mm, JNRS 88320. D)//.amnw,57mm,KU 18847. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRfNELLA 31

Fig. 26. Right hyomandibular of some North American cyprinids illustrating smooth and notched anterior wings. Right is anterior. bellus A) Nocomisraneyi, 53 mm, KU 19616. B)Pteronotropiseuryzonus, 43 mm, KU 19233. C) Lythrurus ,

57 mm, KU 145 1 1 . D) Notropis atherinoides, 55 mm, KU 1 8935. E) Luxilus pilsbryi, 86 mm, KU 1 3009. F) Cyprinella gibbsi, 57 mm, KU 18892. Horizontal bar equals 1 mm. 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY supported by a uniquely derived, tightly packed Lythrurus, Luxilus, Cyprinella, Alburnops, row of tubercles along the lateral surface of Hydrophlox, Notropis, and the Notropis tex- each mandible. This "comb row" of tubercles anus species group, except the Notropis volu- is illustrated by Bailey and Suttkus (1952, cellus species group, lack this groove. Thus, plate IIB). The monophyly of the Pteronotro- the absence of the groove in these species pis clade is also supported by some allozyme supports the monophyly of this clade. Based data (Dimmick, 1987; W. Dimmick, pers. on other characters, the groove present in comm.). members of the Notropis volucellus species Lythrurus-Notropis clade (Fig. 1, Suite 7). group is hypothesized to be areversal, defining Included in this group are at least eight mono- the Notropis volucellus species clade. in this but not phyletic groups. These include the genera Other species included clade, Lythrurus, Luxilus, Cyprinella, and the sharing synapomorphies of stem 13 (Fig. 1), subgenQraAlburnops,Hydrophlox,andNotro- cannot at this time be placed in any of the pis of Notropis, as well as \h& Notropis volu- monophyletic groups. These include Notropis cellus and Notropis texanus species groups scabriceps, N. uranoscopus, N. asperifrons, and N. These are (Fig. 1 ). These groups can be divided, based on hypsilepis. species presently shared derived characters, into two major involved in a polytomy at stem 7 with the clade monophyletic groups, one including the genera containing the Lythrurus-Luxilus-Cyprinella Lythrurus, Luxilus, and Cyprinella. The other lineage and the clade inclusive of the subgen- clade includes the remaining taxa listed above. era and species groups of Notropis from Al- Two derived characters are hypothesized burnops through Notropis (Fig. 1). Their ge- to support the monophyly of this clade. One neric status is retained until such time that they involves the basihyal and the other a fleshy can logically be related to other groups. groove anterior to the lacrymal. These eight Lythrurus-Luxilus-Cyprinella clade (Fig. groups share one derived feature of the ba- 1, Suite 8). The monophyly of these three sihyal not found in any other cyprinids exam- clades is supported by three derived charac- ined. All have an ossified posteroventral proc- ters. Species of this group share 1) large ce- ess from the basihyal, directed toward and phalic tubercles, 2) submarginal tubercles on sometimes (in more derived clades) extending body scales, and 3) a triangular palatine. Most beneath the first basibranchial (Fig. 27). This outgroups within and outside the clade defined process and the anteroventral surface of ba- by an open myodome have very fine tubercles sibranchial 1 are joined by a ventral basihyal on the dorsal surfaces of the head. In these ligament. This basihyal process is not found in three genera, however, the tubercles are large, any other cyprinids. especially in the genera Luxilus and Cyprinella. This process was also noted and discussed Convergent enlarged tubercles occur in some by Coburn (1982) for many of these species, species of Hybopsis, Nocomis, Campostoma, but was not found in his study to include as Semotilus, and Dionda. Based on other char- many taxa as noted here. In members of the acters (Figs. 1, 2, 4), however, the develop- Lythrurus-Luxilus-Cyprinella clade the ba- ment of large cephalic tubercles is hypothe- sihyal projection is generally small, but in sized to be independent in these groups. In all other species (discussed below) it is quite outgroups examined, exczptNotemigonus, the are large. tubercles on flank scales of breeding males A groove extending up along the anterior developed near the edge of the scale (Fig. margin of the lacrymal, separating it from the 28A). In most species of these three genera, rest of the snout (Fig. 27), is found in most however, the tubercles are not at the edge of the species outside this clade and in cyprinids with scale, but are submarginal (Fig. 28B). These a closed myodome. All species of the clade species may have edge tubercles, but they also are defined by stem 7 (Fig. 1) inclusive of develop marginals. The only exceptions NORTH AMERICAN MINNOW GENUS CYPRINELLA 33

Fig. 27. Lateral view of basihyal illustrating presence or absence of posteroventral projection, and lateral view of head illustrating lacrymal groove in some eastern North American cyprinids. Left is anterior. A) Phoxinus erythrogaster, 52 mm, KU 13198. B) Couesius plumbeus, 65 mm, KU 18881. C) Hybopsisamnis, 57 mm,KU 18847. D)Notropistopeka, 50 19033. 6244. Luxilus 85 12648. mm, KU E) Lythrwus fumeus , 56 mm, KU F) zonatus, mm, KU G) Notropis asperifrons, 47 mm, KU 1 8886. H) N. altipinnis, 43 mm, KU 1 8974. I) Oudine drawing of head of Phenacobius mirabilis, illustrating lacrymal groove above lips. J) Outline drawing of head of Cyprinella spiloptera, illustrating absence of lacrymal groove. Horizontal pattern in A-H represents ligaments and stippled regions represent cartilage. Horizontal bar equals 1 mm. 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Luxilus pilsbryi,L. cardinalis, and L. zonatus. In these species the breeding males have edge tubercles only. As mentioned earlier, several morphological aspects of the palatine are important systematically in cyprinids. Most species within and outside the clade defined by the open myodome have a maxillary process. This process is obviously perpendicular to the axis of the palatine (Fig. 10; 29A, E, I, M) in these species. Three modifications of this primitive morphology occur in the open myodome clade. The first two have been discussed above. The Fig. 28. Distribution of breeding tubercles on lateral third derived condition of the palatine occurs body scales in some North American cyprinids. Left is anterior. A) rubricroceus, 55 mm, UAIC in the Lythrurus-Luxilus-Cyprinella clade. In Notropis 5990.01. B) Cyprinella camura, 60 mm, KU 10734. adults of these species the palatine is triangular when viewed dorsally (Fig. 29D, H, L, O). Ju- veniles of these taxa have the primitive later- and dorsolateral surfaces. The monophyly of is discussed Stein ally directed maxillary process (Fig. 29A, E, I, the roseipinnis complex by M). Species with this morphology do not have etal.(1985). the a well-developed maxillary process (Fig. 29). The sister group relationship between Thus, based on outgroup comparison, the tri- roseipinnis complex and the clade composed angular morphology is considered derived for of L. ardens andL. umbratilis is supported by this clade. Developmental data also support three characters. These are: 1) blue and red dark at this triangular shape as being derived. Adults breeding colors along sides, 2) a spot of species with the triangularly shaped pala- origin of dorsal fin, and 3) a very elongate tines have a well-developed maxillary process scapular process against the mesial surface of in L. as juveniles and a smooth transition from the the cleithrum (Fig: 31). The coloration with primitive to the derived morphologies is obvi- fumeus andL. lirus is pale or yellowish no ous with increasing body size (Fig. 29). blues or reds, as is also true for most out- Genus Lythrurus (Fig. 1, Suite 9). The groups, except the genera Luxilus, Cyprinella, monophyly of this genus was discussed by and the subgenus Hydrophlox of Notropis. Snelson (1972) and is supported by three de- Lythrurusfumeus, L. lirus, and outgroups lack rived features. These are: 1) small scales, 2) a dark spot at the origin of the dorsal fin. The reduced anterodorsal squamation, and 3) an scapular process is common to all cyprinids and enlarged urogenital papilla of breeding fe- with an open myodome, except Cyprinella in it is small or absent. In most males (Fig. 30). All three characters are unique Luxilus, which the to this group. The papilla of other breeding outgroups to Lythrurus, however, scapular shiners is never as enlarged as in species of process is shorter than that found in the roseip- umbratilis Lythrurus. innis complex, L. ardens, and L. Within Lythrurus a trichotomy exists (Fig. 31). clade 1 Suite between L.fumeus, L. lirus, and a clade com- Luxilus-Cyprinella (Fig. , 10). of these two posed of the sister species L. ardens and L. The sister group relationship umbratilis, and their sister group, the roseipin- subgenera is supported by ten derived charac- flank /i/scomplex (L. bellus,L. atrapiculus,L. roseip- ters: 1) broad and flat supraethmoid, 2) near the innis.) The L. ardens-L. umbratilis sister rela- scales taller than wide, especially tionship is supported by the presence in breed- lateral line, 3) cephalic tubercles large, 4) head in two rows ing males of dark vertical bars on the dorsal tubercles on dorsum of NORTH AMERICAN MINNOW GENUS CYPRINELLA 35 MP

to the Fig. 29. Ontogenetic transformation from the primitive palatine with a well -developed lateral maxillary process is anterior. derived palatine without a maxillary process, typical of the genera Lythrurus, Luxilus, and Cyprinella. Up A-D) Cyprinella garmani at 24, 30, 35, and 49 mm. E-H) C. camura at 18, 23, 37, and 45 mm. I-L) C. whipplei at 36, 45, 52, and 64 mm. M-O) Luxilus cerasinus at 36, 40, and 57 mm. Horizontal bar equals 1 mm. 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

occurring in some members of the Notropis

volucellus species group (Fig. 1). In this group, however, the elevated scales are confined to the lateral-line series. The enlarged tubercles characteristic of Cyprinella and Luxilus are found in other, unrelated cyprinids, as well as in the closely related genus Lythrurus. The larger tubercles of Cyprinella and Luxilus are interpreted as derived, representing increased modification of large tubercles in the genus Lythrurus. In Lythrurus and most outgroups the supraorbital and mandible tubercles are scattered, representing the primitive conditions for cyprinids with an open posterior myodome. These same outgroups also generally have suborbital tubercles and the dorsal cephalic tubercles in a scattered pattern throughout development (Fig. 32). Thus, the linear development of dorsal head tubercles (Fig. 32 B-E), the single row of supraorbital tubercles, the linear mandibular tubercles, and the absence of suborbital tubercles are hypothesized to be derived characters for these two genera. Reversed conditions are known for the mandibular and supraorbital tubercle patterns. In the genus Cyprinella some have scattered mandibular and su- Fig. 30. Genital papillae of breeding Cyprinella lutrensis species 46 and (above, mm, KU 17297) Lythrurusfumeus (below, praorbital tubercles and C. gibbsi develops tu- 44 mm, INHS 87307) illustrating enlarged for papilla bercles on several suborbital surfaces (see species of the genus Lythrurus. Both females were gravid section and collected with tuberculate males. Left is anterior. Phylogenetic Relationships below). Horizontal bar equals 1 mm. As discussed above, the scapula in most cyprinids within and outside the clade defined anteriorly, 5) mandibular tubercles in rows, 6) by the open posterior myodome has some de- suborbital tubercles absent, 7) supraorbital velopment of a posterodorsal process against

in 1 tubercles rows, 8) posterodorsal process of the mesial surface of the clei thrum (Fig. 3 ). In scapula absent or small, 9) occipital region of Cyprinella and Luxilus the process is absent or

cranium compressed, and 10) loss of pleomer- very small (Fig. 3 1). In Luxilus and Cyprinella ism (the correlation between maximum body the occipital region of the cranium is com- size and number of vertebrae). pressed, reaching an extreme in the lutrensis The broad and flat supraethmoid is unique clade of the genus Cyprinella. In lateral view to this group of cyprinids with an open myod- the occipital region of these two clades is more ome. Outgroups have a narrow supraethmoid nearly vertical than in outgroups, including the with concave margins and concave dorsal genus Lythrurus (Fig. 33). In outgroups the surface, producing a "butterfly" shape (see occipital region is more elongate (Fig. 33). Harrington, 1955). As noted by Coburn (1982) Thus, the compressed condition of these two scales of these two clades are taller than wide genera is considered derived. as measured from the focus of the scale. This Coburn (1982) discussed pleomerism in characteristic is not unique to this clade, also cyprinids and found that for all species except NORTH AMERICAN MINNOW GENUS CYPRfNELLA 37

- lateral Fig. 31. Pectoral girdle of some Cyprinella and outgroups. A-D) Cyprinella xaenura, 7 \ mm.KU 18994: A view - of cleithrum with postcleithrum and scapula; B mesial view of cleithrum, dorsal postcleithrum, and dorsal mesocoracoid; - C - dorsal view of cleithrum; D mesial view of coracoid, mesocoracoid, scapula, and cleithrum. E-H) Lateral view of of of cleithra and scapula illustrating relative heights of ascending arm of cleithrum and length posterior process scapula. E) Notropis telescopus, 67 mm, KU 1523 1. F) Cyprinella analostana, 58 mm, INHS 74347. G) Lythrurus umbratilis, 56 mm, KU 15521. H) Luxilus cardinalis, 79 mm, KU 15281. Stippled pattern represents extent of scapular process. Horizontal bar equals 2 mm; bar below xaenura is for all species, except cardinalis. 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

linear tubercle Fig. 32. Tubercle patterns of dorsum of head in the genera Luxilus and Cyprinella and outgroup, illustrating the 56 5990.01 10 June 1980. pattern early in development for former two genera. A) Notropis rubricroceus, mm, UAIC , B) Early breeding male of Luxilus cardinalis, 64 mm, KU 20737, 13 April 1983. C) Peak breeding male of L. cardinalis, 85 mm, KU 20963, 9 May 1984. D) Early breeding male of Cyprinella proserpina, 41 mm, TNHC 9771, 8 May 1979. C. 977 1 8 1979. circles are tubercles viewed from above. E) Peak breeding male of proserpina, 50 mm, TNHC , May Open Horizontal bar equals 1 mm. those in the genera Luxilus and Cyprinella, sions and illustrations of osteological charac- there is a high correlation between maximum ters see the osteology section in this paper. The size and number of vertebrae. Thus, the lack of monophyly of Cyprinella is substantiated by correlation between maximum body size and the following derived traits: 1) anterior border number of vertebrae is suggested to further of trigeminal foramen formed by pterosphe- substantiate the sister group relationship of noid only, 2) preopercle wide medially and these two groups. anteriorly, 3) interopercle deep posteriorly, 4) Genus Luxilus (Fig. 1, Suite 11). Gilbert anterior hyomandibular wing with notch, 5) (1964) and Buth (1978) supported the mono- lateral shelf of quadrate wide, 6) symplectic phyly of this genus. From the present study broad, 7) frontals truncated anteriorly and three characters are hypothesized to support anterolaterally, 8) frontals slightly broadened the monophyly of Luxilus: 1) retrorse preorbi- laterally, 9) supraethmoid very broad, 10) tal tubercles (also see Gilbert, 1964), 2) clei- supraorbital canal on frontals with elaborate thral region with intense pigmentation (Gilbert, development of canaliculi in a dendritic pat- 1964), and 3) epibranchial 3 with an elongate tern, 11) metapterygoid articulation with in- and curled uncinate process (Fig. 34B, C). Ap- terhyal narrow, 12) ventral horizontal plate of parently, all three characters are unique to this urohyal narrow, 13) vomer with some neck genus. The preorbital tubercles of outgroups developed, 14) interorbital septum shallow, are erect, the cleithral region is never as heav- 15) lacrymal elongate, 16) occipital region ily pigmented as in this clade, and the uncinate compressed, 17) ascending wings ofparasphe- process of other cyprinids is short and straight noid broad, 18) insertion of Al branch of (Fig. 34 A). Coburn (1982) also noted a unique adductor mandibulae on maxilla placed ante- and derived scale morphology for species of rior to isthmus, 19) coronoid process of den- Luxilus. tary vertical, 20) posterior margin of gnathic 2 Genus Cyprinella (Fig. 1, Suite 12). The ramus tall, 2 1 ) fused pharyngobranchials and monophyly of Cyprinella is supported by 34 3 with rough dorsal surface, 22) anterior mar- osteological, behavioral, coloration, and tu- gin of epibranchial 1 with elongate process berculation characters. The hypothesized de- located medially, 23) ceratohyals broad, 24) rived characters are listed below. Only the neural complex and fourth neural spine tall, non-osteological characters are discussed in 25) caudal skeleton broad, 26) anterior mar- reference to polarity justifications. For discus- gins ofcleithra together straight and blunt, 27) NORTH AMERICAN MINNOW GENUS CYPRINELLA 39

PPBO TRP

and arches of some of Fig. 33. Lateral view of occipital regions, Weberian apparatus, and anterior vertebrae species is 54 6244. 57 Cyprinella and close relatives. Left anterior. A) Lythrurusfumeus , mm, KU B) Cyprinella galactura, mm, KU 12029. C) C. garmani, 50 mm, KU 5416. Horizontal bar equals 1 mm. 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dome. Only a few species observed in this study have antrorse cephalic tubercles. These include Lythrurus ardens, L. lirus, Cyprinella, Hybopsis zanema, H. labrosa, and Erimystax monacha. The latter three species have previously been thought to be closely related to Cyprinella. With reference to Fig. 1, however, antrorse tubercles have evolved at least three

times, once in E. monacha (Fig. 3), once in the ancestor of H. labrosa and H. zanema (Fig. 4),

and once in the ancestor of Cyprinella (Fig. 1).

Fig. 34. Third epibranchial and elongate uncinate process of the genus Luxilus and outgroups, illustrating the derived elongate and curled uncinate process of the genus Luxilus. A) Lythrurus ardens, 72 mm, KU 4136. B) Luxilus 80 12654. L. 67 chrysocephalus , mm, KU C) zonatus, mm, KU 12648. Horizontal bar equals 1 mm.

scapular bar pigmentbehind opercle and above pectoral fins, 28) spawning in crevices (not known for all species but hypothesized to be a synapomorphy), 29) pigmentation on scales above and below the lateral line in diamond-

shaped pattern (Fig. 35), 30) gular stripe present (Fig. 36), 31) number of pharyngeal teeth in secondary tooth row 1 or 0, 32) tubercles on dorsum of head antrorse, 33) tubercle pattern discontinuous between dorsum of head and snout, and 34) caudal-peduncle scales above and below the lateral line with scattered cen-

tral tubercles. The derived scapular bar and gular stripe pigment patterns are unique to this genus. The diamond-shaped pigment pattern on scales is not unique to this group. A similar, but conver- B gent color pattern occurs in Erimystax monacha, Notropis maculatus, Hybopsis labrosa. of zanema, and H. The development scales of Fig. 35. Pigmentation on lateral body Cyprinella of the head is the dorsum . lateral erect tubercles on proserpina, 42 mm, TNHC 977 1 A) Scales above Scales below lateral line. primitive for cyprinids with an open myo- line. B) NORTH AMERICAN MINNOW GENUS CYPRINELLA 41

A similar argument may be applied to the territories around substrates where some type evolution of the diamond-shaped pigment of crevice exists for deposition of eggs. This pattern described above. behavior is considered derived over the spawn- In most Cyprinella a hiatus exists between ing behaviors typical of other species within the snout tubercles and tubercles on the dor- and outside the clade with an open posterior sum of the head. A few exceptions are known; myodome. Although not known for all species Cyprinella proserpina, C.panarcys, C. ornata, of Cyprinella, the behavior is predicted to be a C. camura, C. galactura, C. pyrrhomelas, and synapomorphy for the genus. Outgroups typi- C. xaenura have no hiatus. In almost all out- cally deposit eggs in gravel substrates or re- groups the hiatus is absent between these two lease eggs in the open water column (Rabito tuberculate regions. Thus, the hiatus is hy- and Heins, 1985). pothesized to be a derived feature of Cyp- Erimystax monacha appears to be an ex- rinella and has apparently been reversed in ception to this generalization of outgroup some species of this group (see Phylogenetics spawning behaviors. Erimystax monacha has section below). Generally, central tubercles do been observed spawning in crevices like spe- not exist on caudal-peduncle scales of out- cies of Cyprinella (R. E. Jenkins and N. M. groups, especially LythrurusandLuxilus. Some Burkhead, pers. comm.) and a single hybrid members of the earlier discussed chub clade between this species and Cyprinella galactura and the Uybopsis clade, however, develop was reported by Burkhead and Bauer (1983). convergent central scale tubercles on the cau- Because of the similar spawning habits and dal peduncle. The central tubercles of mem- secondary sexual characteristics shared bers of Cyprinella, are considered derived for between E. monacha and species of the genus the group. Cyprinella, one may hypothesize a close rela- Where known, all species of the genus tionship between E. monacha and some spe- Cyprinella have a distinctive and derived cies of Cyprinella, principally members of the spawning behavior. All species are crevice whipplei clade. This is a reasonable hypothesis spawners (Rabito and Heins, 1985). To date, because secondary sexual characteristics and spawning habits are known for C. analostana, breeding behaviors shared by these species C. callitaenia, C. camura, C. galactura, C. appear to be homologous. However, results leedsi, C. lutrensis, C. spiloptera, C. venusta, from the present study indicate that it is more and C. whipplei. Males of these species defend parsimonious to include monacha in the genus Erimystax of the chub clade than in the genus Cyprinella. Pharyngeal dentition is variable in and between species groups having an open posterior myodome. However, because all species oiLythrurus and Luxilus modally have a 2, 4- 4, 2 pharyngeal dentition and species of Cyp-

rinella have 1, 4-4, 1 or 4-4 dentition, the reduction to a single tooth or no teeth in the secondary row is considered derived for the genus. Genus Notropis (Fig. 1, Suite 13). In- cluded in this group are the subgenera Fig. 36. Pigmentation of the gular region in the genus Alburnops ,Hydrophlox, and Notropis, and the Cyprinella. A) Pattern present in all species of the volucellus and texanus species groups. Notro- whipplei clade and most members of the lutrensis clade. and N. are also members of B) Pattern present in the rutila and proserpina species pis greenei harperi pairs. this clade, but are unassignable at this time to 42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY any of the above groups. All of these groups recognized including N. potteri, N. buccula, and species are in an unresolved polytomy, and N. bairdi (Fig. 5). This group is defined by except the subgenus Notropis and the texanus an enlarged lacrymal and an elongate palatine species group, which are sister clades (Fig. 1). (Figs. 10A,B; 37D-F). Other members of Al- All of these monophyletic species groups and burnops and close outgroups have shorter subgenera share a single derived character of palatines at a similar body size (Fig. 10C,D) the basihyal projection mentioned above. In and shorter lacrymals (Fig. 37A-C). Notropis these species the posteriorprojection of the ba- potteri and N. buccula are sister species based sihyal is well developed and rod like, some- on a greatly elongated lacrymal (Fig. 37E ,F). times extending below the first basibranchial. Other species relationships in this subgenus The basihyal projection is longer than that of are unresolved. the Lythrurus-Luxilus-Cyprinella clade and is Notropis volucellus species group (Fig. 1, thus considered a further modification of the Suite 15). Eight species are included in this presumably short ancestral condition. This clade. Notropis heterolepis, N. emiliae, and TV. conclusion is supported developmentally. maculatus form one monophyletic group, sis- Species with the elongate basihyal projection ter to another composed of N. buchanani, N. have a small process, like that observed in the tropicus, N. volucellus, N. ozarcanus,N. spec- Lythrurus-Luxilus-Cyprinella clade as juve- trunculus, and the undescribed "sawfin" shiner niles. The only exception noted is that juve- (Fig. 6). All of these species share two derived niles of Lythrurus umbratilis have a longer characteristics of the palatine. They all have a process than adults. As mentioned above, since long, deep, and restricted preethmoid socket no other cyprinids have this basihyal projec- anteromesially on the palatine and an antero- tion, the more elongate condition is considered laterally directed maxillary process (Fig. 22). evidence of the monophyly of this clade. Justification for this derived character of the Subgenus Alburnops (Fig. 1, Suite 14). palatine is presented under the discussion of Seven species are presently included in this the monophyly of the Hybopsis clade and is group: Notropis edwardraneyi, N. blennius, N. not repeated here. The conditions present in simus, N. girardi, N. potteri, N. buccula, and the Hybopsis clade and in the Notropis volu- N. bairdi (Fig. 5). Notropis orca (Chemoff et cellus species group are similar but not ho- al., 1982) may also belong to this clade. This mologous (Fig. 22). species was not included in my analysis. In all Within the Notropis volucellus species seven of these species the palatine is extremely group, N. heterolepis, N. maculatus, and N. flattened medially (Fig. 10). This morphology emiliae share five derived characters (Fig. 6, is unquestionably derived since almost all Suite B): 1) pharyngeal pad of basioccipital outgroups have a thick palatine. Other taxa narrow and without laterally flared edges, 2) with a compressed palatine include Notropis interopercle deep, 3) operculum with deeply amoenus, N. stilbius, N. photogenis, N. concave dorsal margin, 4) secondary tooth scepticus, N. candidus, N. shumardi, N. ox- row of pharyngeal arch elevated on a pedestal, yrhynchus, and N. jemezanus, all of the and 5) ascending arm of the pharyngeal arch subgenus Notropis, and/V. hypsilepis. Because flattened. These characters are discussed by some members of the subgenus Notropis have Coburn (1982). In close outgroups and other a flattened palatine and since the texanus spe- members of the volucellus species group the cies group is the sister to Notropis, these pharyngeal pad is triangular with laterally morphologies are presently considered con- directed wings, the interopercle is narrower, vergent in the two subgenera. This may not the dorsal margin of theopercle is only slightly prove valid when additional data are available concave, the ascending arm of the pharyngeal on subgeneric relationships. arch is rounded, and the secondary tooth row Within Alburnops, one species group is is not elevated. NORTH AMERICAN MINNOW GENUS CYPRINELLA 43

Species of the second clade within the Notropis volucellus species group (Fig. 6, Suite D) share three derived features. These are: 1) elevated lateral-line scales, 2) supraethmoid broadened anteriorly with expanded dorsolateral corners, and 3) very broad and posteriorly directed ascending wings of the parasphenoid. Other than the elevated flank scales in Cyprinella and Luxilus, the elevated lateral- line scale condition in these species is unique and derived. The broadened supraethmoid of these species (Fig. 38A,C) is also derived because a similar morphology is not frequently observed in outgroups. The broad and posteriorly directed ascending wings of the parasphenoid is derived for this group because close outgroups have narrow and laterally directed ascending wings of the parasphenoid. Notropis volucellus, N. spectrunculus, N. ozarcanus, and the undescribed species (Fig. 6, Suite E) all share a ventrally deflected canal on the lacrymal bone (Fig. 38B,D). No other species of this group and close outgroups have this deflected canal. The group including N. spectruculus, N. ozarcanus, and the undescribed species (Fig. 6, Suite F) is defined by two unique and derived features. All three species share two rows of large and erect tubercles on the leading ray of the pectoral fin of breeding males and a darkened spot on the leading 4-5 membranes and rays of the dorsal fin. Subgenus Hydrophlox (Fig. 1, Suite 16). This clade was diagnosed by Swift (1970) as having the derived features of bright red and/ colors in Fig. 37. Lacrymal bone and infraorbital series of species or orange breeding males, mainly 44 in the subgenus Alburnops. A)Notropis blennius, mm, concentrated on the body, lips, snout, clei- KU 16045. 48 mm, UMMZ 125064. C)N. B)N.simus, thrum, and fins. Included here are N. rubellus; girardi, 43 mm, KU 8039. D) N. bairdi, 53 mm, KU the rubricroceus species group which includes 1 8474. E) N. potteri, 49 mm, KU 14222. F) N. buccula, 36 mm, KU 14286. Horizontal bar equals 1 mm. N. rubricroceus, N. chiliticus, N. baileyi, N. chlorocephalus, and N. lutipinnis; and the 7 Notropis emiliae and N. maculatus share leuciodus species group including A , leuciodus, 7 , and three characters (Fig. 6, Suite C) that were N. nubilus, and A chrosomus. Monophyly discussed by Gilbert and Bailey (1972). These relationships in the leuciodus group are pre- of the include a long anterior arm of the pharyngeal sented by Mayden (1987). Monophyly arch, snout tuberculation, and crosshatched rubricroceus group is supported by Coburn share a derived pigment pattern on lateral scales. I agree with (1982). All of these species these authors that these are derived characters high number of scale radii. Relationships of suggestive of a close relationship. the three species groups of Hydrophlox (rubel- 44 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

share the derived features of an enlarged eye and a large vomerine process on the palatine. Subgenus Notropis (Fig. 1, Suite 19). Fourteen species are included in this clade,

which was diagnosed by Coburn ( 1982) . These include N. candidus, N. shumardi, N. N. atherinoides oxyrhynchus, N. jemezanus, ,

PAL

Fig. 38. Supraethmoid and lacrymal bone of Notropis volucellus (A and B; 40 mm, KU 17025) and N. spectrunculus (C and D; 41 mm, KU 5473). Horizontal bar equals 1 mm.

lus group, rubricroceus group, and leuciodus group) are unresolved. Subgenus Notropis-N. texanus species group (Fig. 1, Suite 17). This clade of Notro- pis is diagnosed by Coburn (1982). All these species share the derived condition of having a large endopterygoid which extends beyond the vertical plate of the quadrate (Fig. 39). In most outgroups the endopterygoid terminates at or anterior to the posterior terminus of the vertical quadrate blade. Notropis texanus species group (Fig. 1, Suite 18). This group is more inclusive than that described by Swift (1970). Eight species N. are included: N. petersoni, N. heterodon, Fig. 39. Pterygoid series of some species of Notropis short and relative to anogenus, N. altipinnis, N. texanus, N. illustrating long endopterygoids anterior fan of quadrate. A) Notropis volucellus, 40 mm, chalybaeus, N. boops, and N. xaenocephalus. KU 17025. B) N. texanus, 44 mm, KU 19797. C) N. All of these share four derived features species amoenus, 70 mm, INHS 77856. Horizontal bar equals 1 (discussed by Coburn, 1982): 1) pharyngeal mm. teeth with distinctly serrated edges, 2) large N. lapillar otolith, 3) black lateral stripe, bordered N. perpallidus, N. amabilis, N. amoenus, N. dorsally by a narrow depigmented stripe, and stilbius, N. photogenis, N. telescopus, 4) enlarged tubercles on the snout and lower ariommus, N. scepticus, and N. semperasper. jaw. Within this group relationships are pres- For a discussion of monophyly of this group ently unresolved, except for the N. boops-N. and species relationships within the subgenus xaenocephalus species pair. These two species see Coburn (1982). NORTH AMERICAN MINNOW GENUS CYPRINELLA 45 SYSTEMATICS OF CYPRINELLA

CYPRINELLA GlRARD by pterosphenoid only; preopercle wide medi- Cyprinella Girard 1857, Proc. Acad. Nat. ally and anteriorly; interopercle deep posteri- Sci. Philadelphia (1856) 8(5): 196-197. Type: orly; anterior hyomandibular wing with notch; Leuciscus bubalinus Baird and Girard, 1853 lateral shelfofquadrate wide; symplec tic broad; [=Leuciscus lutrensis Baird and Girard 1853], frontals truncated anteriorly and anterolater- by subsequent designation of Jordan and al^; frontals slightly broadened laterally; su- Gilbert, 1877:91. praethmoid very broad; supraorbital canal on Codoma Girard 1857, Proc. Acad. Nat. Sci. frontals with elaborate development of canali- Philadelphia (1856) 8(5): 194-195. Type: culi in a dendritic pattern; metapterygoid ar- Codoma ornata Girard, 1857, by subsequent ticulation with interhyal narrow; ventral hori- designation of Jordan and Gilbert, 1877:91. zontal plate of urohyal narrow; vomer with Moniana Girard 1857, Proc. Acad. Nat. some neck developed; in terorbital septum shal- Sci. Philadelphia (1856) 8(5): 199. Type: low; lacrymal elongate; occipital region com- Leuciscus lutrensis Baird and Girard, 1 853, by pressed; ascending wings of parasphenoid subsequent designation of Jordan and Gilbert, broad; insertion of Al branch of adductor 1877:91. mandibulae on maxilla placed anterior to isth- Photogenis Cope 1868, Proc. Acad. Nat. mus; coronoid process of dentary vertical; Sci. Philadelphia (1867) 19: 163-164. Type: posterior margin of gnathic ramus tall; fused Leuciscus spilopterus Cope, in Giinther, 1868, pharyngobranchials 2 and 3 with rough dorsal by subsequent designation of Cope in Jordan surface; anterior margin of epibranchial 1 with and Copeland, 1877:154. elongate process located medially; ceratohy- Erogala Jordan, in Jordan and Brayton, als broad; neural complex and fourth neural 1878, Bull. U. S. Nat. Mus. 12:20-21. Type: spine tall; caudal skeleton broad; and anterior Photogenis stigmaturus Jordan, 1877b margins of cleithra together form straight and [=Cyprinella venusta Girard, 1857], by origi- blunt surface. nal designation. Composition. The following 27 species Diagnosis. In addition to the synapomor- are included in the genus Cyprinella: C. ornata, phies listed under the previous section relating C. lutrensis, C. garmani, C. formosa, C. Cyprinella to other North American cyprinids, bocagrande, C. lepida, C. rutila, C. xanthicara, species of the genus Cyprinella may be distin- C. proserpina, C. panarcys, C. spiloptera, C. guished from other cyprinids by the following camura, C. whipplei, C. analostana, C. derived characters: scapular bar of pigment chloristia, C. venusta, C. galactura, C. behind opercle and above pectoral fins; spawn- pyrrhomelas, C. xaenura, C. caerulea, C. ing in crevices (not known for all species but trichroistia, C. gibbsi, C. callistia, C. nivea, C. hypothesized to be a synapomorphy); exposed leedsi, C. callisema, and C. callitaenia. margins of lateral scales taller than wide; pig- Distribution. Members of the genus Cyp- mentation on scales above and below lateral rinella inhabit rivers of eastern North America line in diamond-shaped pattern; gular stripe from the Atlantic S lope to central Mexico (Fig. present; fins of breeding males with milky- 40). All species occur east of the Rocky white deposition; number of pharyngeal teeth Mountains except C. ornata and C. formosa. in secondary row 1 or 0; tubercles on dorsum These species are found in parts of northwest- of head strongly antrorse; tubercle pattern ern Mexico. discontinuous between dorsum of head and Classification of Cyprinella. The classifi- snout; caudal-peduncle scales above and be- cation below is arranged in order of the phylo- low lateral line with scattered central tubercles; genetic relationships of the species, following anterior border of trigeminal foramen formed the phylogenetic listing convention of Nelson 46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

(1974) and Wiley (1979): C. camura (Jordan and Meek) C. whipplei Girard lutrensis clade C. analostana Girard C. ornata (Girard) C. chloristia (Jordan and Brayton) lutrensis species group venusta species group C. lutrensis (Baird and Girard) C. venusta Girard C. garmani (Jordan) C. galactura (Cope) formosa species group pyrrhomelas species group C.formosa Girard pyrrhomelas species pair C. bocagrande (Chernoff and Miller) C. pyrrhomelas (Cope) lepida species group C. xaenura (Jordan) C. lepida Girard caerulea species complex rutila species pair C. caerulea (Jordan) C. rutila (Girard) C. gibbsi (Howell and Williams) C. xanthicara (Minckley and Lytle) C. trichroistia (Jordan and Gilbert) proserpina species pair nivea species group C. proserpina (Girard) C. callistia (Jordan) C. panarcys (Hubbs and Miller) C. nivea (Cope) C. leedsi (Fowler) whipplei clade C. callisema (Jordan) C. spiloptera (Cope) C. callitaenia (Bailey and Gibbs) whipplei species group

SPECIES ACCOUNTS

Below are accounts of the 27 species in the and Brayton (1878), Jordan and Evermann genus Cyprinella. Each account includes an (1896),JordanandMeek(1884),Koehn(1965), abbreviated synonymy, including only genu- Lytle (1972), Matthews (1987), Meek (1904), ine synonyms for each species. For a more Minckley and Lytle (1969), Pflieger (1975), complete synonymy of each species see Gibbs Smith (1979), Trautman (1957; 1981), and (1955; 1957a, 1957b, 1957c; 1961; 1963), original descriptions of species. Patterns of Contreras-Balderas (1975), and various origi- breeding tubercles and melanophore distribu- nal descriptions. Each summary description tions in dorsal fins for species of Cyprinella includes meristic, coloration, and tubercula- and close relatives are illustrated in Figs. 41-48. tion characteristics, as well as distributional Species are illustrated in Plates 1-4. Species information, etymology, and an ecological Accounts below are arranged in order of the summary. Data presented below are original phylogenetic relationships of the species, fol- from this study and/or summarized from Bailey lowing the phylogenetic listing convention of andGibbs(1965),ChernoffandMiller(1982), Nelson (1974) and Wiley (1979). Contreras-Balderas (1975), Cross (1967), Denoncourt and Messersmith (1982), Forbes Cyprinella ornata (Girard) and Richardson (1 908), Garman (1 88 1 ), Gibbs Ornate Shiner (1955; 1957a, 1957b, 1957c; 1961; 1963), Plate I Girard (1857), Howell and Williams (1971), Hubbs and Miller (1978), Jordan (1877a, Codoma ornata Girard, 1857:195. [Orig. 1877b, 1877c; 1878a; 1878b; 1885), Jordan descr.; Type locality: Rio Chihuahua and tribu- NORTH AMERICAN MTNNOW GENUS CYPRINELLA 47 — -a n

Fig. 40. Composite distribution for the ranges of species in the genus Cyprinella. taries, near Chihuahua City, Chihuahua, shelf of dentary strongly developed; isthmus Mexico] of maxilla deep; vomer without neck; poste- Diagnosis. Anal rays 7-8; tall and dark rior margin of horizontal urohyal plate oval vertical bars along flanks; head blunt; base of and without notch; epiphyseal bar placed an- caudal fin with whitish patches; hiatus be- teriorly without modifications to frontals. tween dorsal head and snout tubercles absent; Description. A medium-sized Cyprinella, pigmentation on scales above and below lat- attaining a maximum SL of about 60 mm. eral line diffuse and scattered, not particularly Body robust and only slightly compressed, diamond-shaped; supraethmoid process of deepest predorsally. Head rounded to subcorn- palatine not strongly inflected above horizon- eal. Mouth terminal to subterminal, clinally tal; anterior hyoideal foramen of ceratohyal variable, jaws unequal. Snout blunt. Orbit located laterally; posterior hyoideal foramen generally small. Caudal peduncle robust and of ceratohyal located mesially; hypohyal fora- deep. Dorsal fin not expanded in breeding men small; sixth hypural absent; dorsal proc- males; origin over pelvic fin insertion. ess of epibranchial four short; dorsolateral 48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Dorsal fin rays 6-9 (usually 8); anal rays dark bluish-black; gular region with scattered 6-9 (7 in lower Conchos race and 8 in all melanophores; postanal stripe absent in all but others); pelvic rays 6-9 (7-8); pectoral rays Mezquital race, in which it may be present. All 9-16 (13-14); principal caudal rays 17; dorsal fins darkly pigmented on rays and membranes, procurrent rays 10 or 11, ventral procurrent except for narrow distal white band. Base of rays 10-11 (10). caudal fin with whitish patches. Exposed margins of lateral body scales Females, like males, but paler. Venter un- higher than wide. Lateral-line scale rows 37^47 pigmented, including gular and branchiostegal (41-43) in upper Conchos and 32^13 (36-38) regions. Intense black coloration over all of in others; predorsal scales 15-24 (19-21 in body not developed. Conchos and 16-18 in others); total predorsal Tuberculation.— In prenuptial males dor- circumferential scales 32^40 (34-37) in upper sum of head mostly covered with erect tu- Conchos and clinal in others (25-38, generally bercles located laterally in rows behind orbit 27-32, depending on drainage); predorsal and with clusters of erect organs between scales above lateral line 15-19 (17) in upper nares, above orbits, and between each orbit Conchos and 11-16 (13-15) in others, below and naris. Internarial cluster of tubercles rep- lateral line 15-19 (16-17) in Conchos or 12-1 8 resents continuation ofsnout organs with dorsal (13-15) in others; total caudal-peduncle scales tubercles. Snout tubercles erect and clustered, 16-22 (18-20) in Conchos or 12-20 (16-18) not necessarily in rows. Preorbital organs erect in others; scales above lateral line 5-10 (gen- and continuous with snout tubercles and single erally 7 in all but upper Conchos with 9), scales row of 5-6 erect supraorbital tubercles. Re- below 5-11 (7 in all except Yaqui and upper mainder of head devoid of tubercles. Conchos with 7-9). Body tuberculation weak compared to that Pharyngeal teeth 4-4, strongly hooked, and of other species of the lutrensis clade. Predor- with well-developed grinding surfaces. Total sal tubercles absent. Along flanks, 0-3 small, on each gill rakers 6-8 (8). Total vertebrae 36-39 retrorse organs developed subcentrally (37-38), precaudals 18 or 19, caudals 18-20 scale, as in other members of this group. Broad (19). Dorsal fin insertion above vertebrae 12- band of organs on lower portion of caudal 14(12). peduncle not developed. Each scale with 0-10 Lateral line complete to hypural plate. small, retrorse, subcentral tubercles; no cen- Supratemporal canal narrowly interrupted, tral organs present in specimens examined. pores 2. Supraorbital canal without interrup- Strongest development occurs below lateral tions and not joined with other canals, pores line. 7-8 (8). Infraorbital canal complete, pores All fins except pectorals with slight tu- 9-10 (10). Preoperculomandibular canal bercle development on first 2 rays distally. complete, total pores 11, mandibular pores 2. First ray of pectoral fin with 1 row of tubercles, Intestine simple, with single loop, Type I 1 organ per segment. Posterior rays 2-6 with 1 (Kafuku, 1958). Peritoneum silvery with speck- basal row, which continues onto posterior ling of melanophores. branch; anterior branch with 1 row. Each ray — 1 retrorse Ventral Coloration in life. Reproductive males with organ per segment. darkly pigmented, nearly black in peak condi- erect tubercles developed on paired fins. tion. Otherwise, dorsum dark, 9-14 lateral Distribution. Contreras-Balderas (1975) bars, variable in length, sometimes coalescing recognized five races of Cyprinella ornata.

to form lateral stripe, darkest posteriorly. Lat- These races are endemic to the upper Rio eral stripe obscured in peak males. Basicaudal Mezquital, Rio Nazas, upper Rio Conchos, spot present and usually separate from lateral lower Rio Conchos, upper Rio del Fuerte (not Rio stripe. Venter lightly pigmented, cream to white seen by Contreras-Balderas), and upper in some; other peak condition males mostly Yaqui of northwestern Mexico. A distribution NORTH AMERICAN MINNOW GENUS CYPRINELLA 49 map of the species, excluding Rio del Fuerte nize the genus Codoma for the ornate shiner populations, is provided by Contreras- and have considered the possibility that it is Balderas. related to the genus Pimephales. This conclu- Ecology. The biology of this species is sion is based on gross morphological similari- poorly known. It inhabits clear streams with ties, snout tuberculation, and spawning habits rubble or gravel substrate, where it frequents of the species. Results of this analysis are in riffle or fast-water habitats. Spawning has been agreement with Gibbs and demonstrate that C. observed by Minckley (pers. comm.). Appar- ornata represents the sister group to the re- ently this species spawns upside down under maining members of the lutrensis clade. rocks. Such behavior is known to occur in the cyprinid genus Pimephales (Cross, 1967), a Cyprinella lutrensis (Baird and to which C. ornata has been to genus thought Girard) be related because of these char- reproductive Red Shiner acteristics and gross morphological similari- Plate I ties (Miller, 1976). It is not unusual, however, for species of Cyprinella to spawn on the Leuciscus lutrensis Baird and Girard, undersides of objects in crevices (Pflieger, 1853:391. [Orig. descr.; Type locality: Otter 1965), where their adhesive eggs attach to any Cr., trib. to N. Fk. Red River, either Kiowa or substrate after spawned. Hendrickson et al. Tillman Co., OK] (1980) observed breeding C. ornata on 16 Leuciscus bubalinus Baird and Girard, June over bedrock and boulder riffles, defend- 1853:391. [Orig. descr.; Type locality: Otter ing areas around crevices and loose rocks on Cr., trib. to N. Fk. Red River, either Kiowa or the bottom. Ova were attached to the under- Tillman Co., OK. Hubbs and Ortenburger sides of the stones. Meek (1904) believed (1929), as first revisers, chose lutrensis over spawning to occur in early June. In May, 1987, bubalinus, as both were described on the same adult males from the Rio Conchos, Rio Yaqui, page] and Rio del Fuerte were observed defending Cyprinella beckwithi Girard, 1857:197. territories in riffle and raceway habitats around [Orig. descr.; Type locality: Sluices of Arkan- gravel and cobble substrate (Mayden, unpub. sas River, near Ft. Makee, KS] data). No eggs were found attached to any of Cyprinella gunnisoni Girard, 1857:197. the substrate materials. [Orig. descr.; Type locality: Cottonwood River, Etymology. The name ornata is Latin from ca. 5 mi NW Durham, Marion Co., KS (after ornare meaning ornamented or showy, refer- Gilbert, 1978a)] ring to coloration of breeding individuals. Cyprinella suavis Girard, 1857:197. [Orig. Comments. Contreras-Balderas (1975) descr.; Type locality: near San Antonio, TX] recognized five races of C. ornata, the Rio Cyprinella umbrosa Girard, 1857:197. Mezquital, Rio Nazas, upper Rio Conchos, [Orig. descr.; Type locality: Coal Creek, trib. lower Rio Conchos, and upper Rio Yaqui. to S outh Fork Canadian River, N of McAlester, Populations from the upper Rio del Fuerte Pittsburg Co., OK] were not examined. These "races" were found Moniana leonina Girard, 1857:199. [Orig. to differ meristically, morphometrically, and descr.; Type locality: Leon River, trib. to San in color and tuberculation patterns; and may Antonio River, Bexar Co., TX] eventually be recognized as distinct species. Moniana frigida Girard, 1857:200. [Orig. The relationship of the ornate shiner to descr.; Type locality: Salado Creek, just E San other minnows has been uncertain. Gibbs Antonio, Bexar Co., TX; Rio Sabinal, at Sabi- (1957a) considered the ornate shiner to be a nal, Uvalde Co., TX; Rio Medina, at Castrov- member of Cyrinella. Contreras-Balderas ille, Medina Co., TX; Rio Nueces, ca. 7 mi W (1975, 1978) and Miller (1976, 1978) recog- Uvalde, Uvalde Co., TX; Rio Frio, ca. 10 mi 50 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

NE Uvalde, Uvalde Co., TX] Description. A moderately large Cyprinella, Montanapulehe lla Girard, 1857:200. [Orig. reaching a maximum of about 75 mm SL. Body descr.; Type locality: Sugar Loaf Creek, trib. form geographically variable. Individuals from Poteau River, about 20-25 mi S Fort Smith, Mississippi River tributaries, Rio Grande AR, vicinity of Poteau, OK] proper, and Rio Grande tributaries (ribs San Montana complanata Girard, 1857:200. Juan, Pecos, and Conchos) have a very deep [Orig. descr.; Type locality: Rio Grande (or body with highly arched back, deepest at dor- tributary), Brownsville, Cameron Co., TX] sal origin, and are highly compressed. Head Montana laetabilis Girard, 1857:200. [Orig. large and deep. Snout pointed. Orbit small. descr.; Type locality: Hurrah Creek (trib. Pecos Mouth and gape width moderately large, River), ca. 10 mi NE Santa Rosa (35°05' maxillary extending to middle or anterior orbit. 23.7"N,104°43'27.3"W),GuadalupeCo.,NM] Caudal peduncle deep and robust. Dorsal fin Montana couchi Girard, 1857:201. [Orig. origin over or slightly posterior to insertion of descr.; Type locality: Rio San Juan, vicinity of pelvic fin. Individuals from GulfCoastal drain- China, Nuevo Leon, Mexico] ages (here considered C. /. suavis and C. /. Montana gibbosa Girard, 1857:201. [Orig. forlonensis) are much more slender and elon- descr.; Type locality: trib. to Rio Grande, gate with a narrower body, head, and caudal Brownsville, Cameron Co., TX] peduncle, larger orbit, and smaller mouth. Cyprinella billingsiana Cope, 1871:439. Dorsal fin not expanded in breeding males and [Orig. descr.; Type locality: Missouri River, inserted over pelvic fin insertion. These trends St. Joseph, MO] are discussed by Matthews (1987) for popula- Montana jugalis Cope, 1871:439^440. tions within the United States and by Contre- [Orig. descr.; Type locality: Missouri River, ras-Balderas (1975) for populations in Mexico. St. Joseph, MO] Values for meristic characters generally Hypsilepis iris Cope in Cope and Yarrow, increase from east to west and southward from 1875:653-654. [Orig. descr.; Type locality: upper Mississippi, through the Platte, Kansas, Rio Grande, at San Ildefonso, ca. 10 mi E Los Arkansas, upper Rio Grande, and Conchos Alamos, Santa Fe Co., NM] drainages. Meristics generally decrease east Cyprinella forbesi Jordan, 1878b:57-58. and south in the Neosho, Verdigris, Canadian, [Orig. descr.; Type locality: Illinois River, upper Red, and lower Red-Mississippi drain- Union Co., JL] ages. Meristics of populations from Gulf Notropis forlonensis Meek, 1904:70-71. Coastal drainages from the Sabine-Calcasieu [Orig. descr.; Type locality: Rio Forlon, trib. system westward are also variable. In the Rio Panuco, Forlon, Tamaulipas, Mexico] Sabine-Calcasieu system low circumferential Notropis lutrensis blairi Hubbs, 1940:6-8. and lateral-line scale counts predominate in [Orig. descr.; Type locality: Garden Springs addition to the lowest means of all drainages (=Monument Spring), trib. Pefia Colorado for caudal-peduncle and predorsal scales, but Creek (Maravillas Creek drainage), 12-13 mi overlap occurs. Variation in meristic features SSW Marathon, Brewster Co., TX. Elevation of populations west of the Trinity River show

3700 ft] no significant concordant variation other than Diagnosis. Anal rays 9; body generally slightly higher means for circumferential and deep; fins of breeding males red; chin bar caudal-peduncle scales. short; scapular bar well developed; flank scales Dorsal fin rays 6-12 (8); anal rays 8-10 (9); above lateral line with single central tubercle; pelvic rays 4-9 (8); pectoral ray s 8- 1 6 ( 1 2- 14); neck on ceratobranchial 1 bent mesially; pos- principal caudal rays 17; dorsal procurrent terior process of quadrate deep and heavy; an- rays 11-13 (12), ventral procurrent rays 10-11 terior notch on vomer deep; hypohyal foramen (10). small. Exposed margins of lateral and dorsolateral NORTH AMERICAN MINNOW GENUS CYPRINELLA 51 body scales higher than wide. Lateral-line ily pigmented cresents on each scale. Centers scale rows 28-38 (all 33-36, exceptforlonen- of scales powder-blue or sometimes pinkish. sis with 32-34); predorsal scales 13-21 (15-18 Crosshatched pattern obliterated just posterior in most, except Rio Conchos with 16-19); to opercle by dark blue wedge-shape scapular area total predorsal circumferential scales 22-33 bar and pink or reddish rectagular poste- (24-29); scales above lateral line 11-17 rior to bar. Venter white, silver, cream, or red. (13-15), below lateral line 8-15 (9-12); total Gular bar short and dark, extending from caudal-peduncle scales 12-20 (14-16, most symphysis to retroarticular-interopercleregion. with 14, except 14-18 in Rio San Fernando Side of head silver. Pectoral and pelvic fins red and 16-18 in Rio Conchos and upper Rio to reddish-orange, except for milky-white distal Grande); caudal-peduncle scales above lateral margins. Anal fin darkened basally by mela- line 5-9 (most with 7, except rios San Fer- nin, red medially, and white distally. Caudal nando, Conchos, and upper Grande with 7-9), fin reddish except for clear or whitish distal scales below lateral line 5-7 (most with 5, margin. Dorsal fin heavily pigmented with exceptRio San Fernando and upper Rio Grande interradial and radial melanophores, over a with 5-7 and Conchos with 6-7). background of light reddish tint. Pharyngeal teeth generally 4-4. Teeth The above description pertains to the most hooked and with well-developed grinding sur- frequently observed color pattern, but vari- to exist. an faces. Total gill rakers 8-11 (10). Total verte- ations are known As example, brae 34-36 (35), precaudals 16-18 (17), cau- Hubbs (1982) suggested that at least 5-10 dals 17-19 (18). Dorsal fin insertion above different forms of "lutrensis" could be distin- vertebrae 10-12 (11). guished from the Trans-Pecos Region ofTexas Lateral line generally complete to hypural alone. Characteristics and distributions of these plate. Supratemporal canal broadly interrupted, forms were not given. pores 2 per side. Supraorbital canal complete, One variation is what Matthews (1987) pores 8. Infraorbital canal complete, pores refers to as the "yellow-fin form." These indi- 10-11 (11). Preoperculomandibular canal viduals are geographically restricted, have a complete, total pores 10-12 (11), mandibular typical lutrensis body form, lack the red or pores 3. reddish-orange fin coloration, and have been Intestine with two simple flexures, forming collected with the"typical" red shiner "morph." S-shaped coil, Type I. Peritoneum silvery with Populations of this form are known from King- sparse speckling of melanophores. man, Jefferson, and Cowley Cos., KS and — Coloration consists Coloration in life. Females and nonbreed- Baron Co., MO. basically

ing males pallid. Dorsum olive, metallic blue, of bluish dorsum and sides. The head is not red or bluish-green turning to bluish-silver later- dorsally and the scapular bar is not set off from ally and white or cream ventrally. Fins lightly remainder of flank by depigmented or lightly colored; anal and caudal sometimes with light pigmented rectangular region posteriorly. Pec- orange or red; dorsum dusky; pectorals and toral, pelvic, anal, and caudal fins are consis- pelvics depigmented except for leading rays tently yellow. Another variant, in some Gulf with some melanin. Coastal populations, has a deep red venter, Breeding males brightly colored. Dorsum unlike breeding lutrensis observed elsewhere. brightblue, greenish, or bluish-green. Dorsum Tuberculation.— Dorsum of head of of head same, but may be bright red or rufous. smaller, younger males or prenuptial males Opercles, subopercles, and posterior pre- with moderately sized antrorse tubercles in opercles and interopercles generally pink or linear pattern. Later in development, linear light reddish orange. Flanks lighter, with pattern of 2 rows down midline obscured by powder-blue background overlain by a dark development of additional tubercles on oc- steel-blue crosshatched pattern forming heav- cipital region, obliterating linear pattern on 52 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY postorbital area. Two rows anterior to postor- single tubercle per segment. First ray of pecto- bit. Supraorbital tubercles moderately large, ral fin with a single row of tubercles and 2-3 about same size as dorsal head tubercles, con- organs per segment. Early in development, tinuing beneath each nare and connected with posterior rays 2-7 with single row per ray; snout tubercles. Generally, no preorbital tu- later single row turns to 2-4 rows on main bercles develop close to orbit. At anterior edge body of each ray, with several tubercles per of lacrymal, where supraorbital row connects segment. Distal to branching, pectoral fin rays with snout tubercles, 3-5 erect, moderately of early males with single row and 2-3 organs large tubercles are formed. Snout tubercles per segment; peak breeding males generally few and scattered, antrorse posteriorly and have 2 rows and several tubercles per segment. erect anteriorly, and separated from dorsal In extremely tuberculate males, row pattern head tuberles by wide hiatus. Except for a few obscured by thick mat of tubercles. Ventral erect, moderately large tubercles restricted to sides of paired fins with erect tubercles on dorsal one-third of opercles, no other tubercles rays. develop on head. Distribution. Native to the upper Missis- Predorsal scales generally with 3 moder- sippi basin as far north as southern Wisconsin ately large tubercles in transverse row, similar and western tributaries to the Mississippi river, in size to dorsal head tubercles or slightly including the Missouri, Arkansas, and Red smaller. Central tubercle sometimes slightly rivers. Along the Gulf Slope, the red shiner larger than lateral tubercles. At edge of each inhabits rivers from near mouth of Mississippi predorsal scale several retrorse organs (some River westward and southward to the Rio erect) present. Along flanks, scales with 1-2 Panuco, Mexico. The red shiner also occurs in centrally located antrorse organs, smaller than the Rio Grande basin, inclusive of rfos Con- predorsals by half. Around these, 0-5 erect chos, Salado, and San Juan. This aggressive submarginals may develop per scale, but are species has been introduced into many areas in generally absent. Rank scales also with 3-15 U.S., and where introduced may sometimes retrorse edge tubercles similar in size to sub- "swamp-out" native Cyprinella gene pools via marginals and about one-half size of central hybridization (Page and Smith, 1970). tubercles. Below lateral line, above and ante- Ecology. Considering the broad range of rior to pelvic fins, each scale with small edge this species and its widespread use as a baitfish, tubercles only. little is known of its biology. Detailed studies Caudal peduncle tuberculation complex, include those of Cavin (1971), Laser and beginning above and posterior to pelvic fins Carlander (1971), and Farringer et al. (1979). and extending to hypural plate. Maximum Other studies and general comments on its development above anal fin. Below lateral biology have been published (Saksena, 1962; line, 1-4 (almost always 3 in peak males) Koehn, 1965; Cross, 1967; Islam, 1972; Minck- moderately large, antrorse organs in center of ley, 1972, 1973; Miller and Robison, 1973; each scale. Marginal to these, 1-4 moderately Pflieger, 1971, 1975; Matthews and Hill, 1977). large and slightly antrorse organs developed. Basically, the red shiner is an aggressive spe- Along edge of scale, 3-8 small retrorse tu- cies, capable of inhabiting a variety of habitats bercles developed in a series. Scales above and environments, but is generally atypical of lateral line similar, but organs much smaller. highland regions of the central U.S. It does Tubercle development on dorsal fin does well in harsh and variable environments when not occur until male is in peak breeding condi- other species disappear (Cross, 1967) and has tion. Pelvic fins tuberculate, with a single row a high tolerance for low oxygen levels (Cavin,

1 It with or inter- per ray and 2 tubercles per segment. Dorsal and 197 ). prefers creeks moderate anal fins both with tubercle development on mittent flow (Paloumpis, 1958), but also fre- first ray; both with a single row per ray and quents large rivers over a sand, silt, or gravel NORTH AMERICAN MINNOW GENUS CYPRINELLA 53 substrate. It appears to be quite tolerant of logs, roots) while releasing eggs and milt. turbidity and siltation. Etymology. The name lutrensis is Latin for In Illinois and Missouri most red shiners Otter, in reference to Otter Creek, Arkansas, live 2+ years (Lewis and Gunning, 1959; where first discovered. Pflieger, 1975). Its reproductive cycle in Okla- homa and Texas spans from April to Septem- Cyprinella garmani (Jordan) ber, in Missouri from May to September, in Gibbous Shiner Kansas from to October, and in Illinois May Plate I. from May to August. Ovarian regression in Oklahoma and Texas occurs in midsummer Notropis garmani Jordan, 1885:813. [Sub- with increased temperature and aridity, but stitute name for preoccupied rubripinna Gar- some data on length frequency and ovarian man, 1881] recrudescence indicate two peak periods of Cyprinella rubripinna Garman, 1881:91. spawning activity. Most if not all individuals [Orig. descr.; Type locality: Lago del Muerto, breed during their second summer, with very near Parras, Coahuila, Mexico] few breeding during the first summer (Farrin- Diagnosis. Anal rays typically 10; body ger et al., 1979). Some data from Illinois red very deep; caudal peduncle very slender; anal shiners, however, suggests that 2-year-old fin placed anteriorly such that a vertical line individuals spawn in July and the 1 -year-old dropped from posterior base of dorsal fin inter- fish spawn later in August and September sects base of anal fin posterior to base of (Lewis and Gunning, 1959). second principal ray (Chernoff and Miller, Spawning locations vary. Males and fe- 1986); gular bar short; mandibular pores 4; males may congregate in pools below riffles or preorbital tubercles distributed over entire near brush, logs, or other debris where the eggs lacrymal area; dorsal head tubercles erect; are fertilized, settle to the substrate, and adhere pharyngeal pad of basioccipital flat. to various objects (Minckley, 1972; Smith, Description. A large Cyprinella for the 1 979). In some cases the red shiner may spawn lutrensis clade, reaching about 60 mm SL. over nests of sunfish (Lepomis cyanellus, L. Body deep and strongly compressed; dorsum humilis). Males defend territories near the convex, strongly arched, deepest at dorsal margins of the sunfish nests where the eggs are origin. Head moderately large. Snout blunt to released (Minckley, 1959; Pflieger, 1975). subconical. Mouth small, terminal, oblique; Spawning occurs most frequently, however, in maxillary extending to anterior margin oforbit. clean gravel riffles, or on submerged objects Caudal peduncle narrow and elongate. Dorsal (e.g., logs and tree roots), where the eggs are fin not expanded in breeding males and origin deposited in crevices (Pflieger, 1975; see also in both sexes over anal fin.

Minckley, 1972). Minckley (1972) also re- Dorsal fin rays 8-9 (8); anal rays 9-13 ported that spawning may occur in midwater (10-11); pelvic rays 7-9 (8); pectoral rays while the pair swims through the water col- 11-14 (12-13); principal caudal rays 17; dor- umn. The demersal, adhesive, and yellowish- sal procurrent rays 9-12 (11-12), ventral pro- white fertilized eggs are then spread over the current rays 10 or 11. substrate. Exposed margins of lateral and dorsolateral When single territories are maintained by a body scales higher than wide. Lateral-line male, a female swims into the area, eliciting scale rows 33-40 (35-37); predorsal scales display behavior by the male consisting of 16-24 (19-21); total predorsal circumferen- several "figure 8" movements before swim- tial scales 24-35 (29-30); scales above lateral ming near the female. Once the male is along- line 13-19 (15-16), below lateral line 9-16 side the female, the two move side by side (11-13); total caudal-peduncle scales 12-17 through or near the substrate (gravel, algae, (14); caudal -peduncle scales above lateral line 54 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

5-9 (7), below lateral line 4-7 (5). gans on dorsal operculum. Pharyngeal teeth 4-4, well hooked, narrow, Predorsally, 3 moderately large, erect or- but with moderately well-developed grinding gans in semilinear arrangement in center of surfaces. Total gill rakers 9-10 (9). Total ver- each scale; periphery of each scale with series tebrae 35-36 (36), precaudals 16-18 (17), of small erect organs. Scales along flanks with caudals 17-20 (18-19). Dorsal fin insertion 5-10 small retrorse edge-tubercles each. Scales above vertebra 12 or 13 (12). along lower caudal peduncle and 3 scale rows Lateral line complete to hypural plate. above anal fin heavily coated with tubercles. S canal broadly interrupted, pores upratemporal Center of each scale with a cluster of moder- 2. Supraorbital canal complete, pores 7-9. ately large, antrorse organs. Antrorse submargi- Infraorbital canal weakly developed, partially nal organs and a series of antrorse or retrorse interrupted, pores 11-12. Preoperculomandi- edge tubercles peripheral to central row. Ven- bular canal complete, total pores 9- 1 2 ( 1 1-1 2), tral row of caudal-peduncle scales also with mandibular pores 3-4 (4). this pattern. Dorsally on caudal peduncle only Intestine simple, single loop with two small, erect edge tubercles present. flexures, Type I. Peritoneum silvery with speck- All fins tuberculate. Dorsal 1-6 with ling of melanophores. rays — to row and Coloration in life. Coloration similar single per ray single organ per seg- that of C. lutrensis. Females and nonbreeding ment, developed medially. All rays of anal fin 1-2 tubercles males lightly colored. Bright colors absent with a single row per ray and per except on anal and caudal fins, where light red segment. Pelvic fin rays with 2-3 tubercles per may develop. Venter cream to white. Lateral segment in a single row on rays 1-5. Leading 2 body silver-blue. Dorsum bluish-gray, venter pectoral fin ray with a single row of retrorse Pectoral 2-9 tuber- light cream to white. Dark lateral band weak organs per segment. rays 2 anteriorly and strong posteriorly; caudal spot culate; single row basally becoming rows absent. Dark scapular bar present immediately prior to segmentation. After segmentation each Distal to behind head, offset by lighter, rectangular area segment with 4-6 organs. branching, 2 posteriorly. Pigment on flank scales in diago- one row anteriorly with organs per segment 4 nal rows, presenting diamond- shaped pattern. and 2 rows posteriorly, with tubercles per Venter immaculate. Fins with scattered melano- segment. Ventrally, pectoral and pelvic fins phores, appearing dusky. with several erect organs per ray. Breeding males as above except more bril- Distribution. This species is known only liantly colored over body and fins. Rectangu- from rios Nazas and Aguanaval of north-cen- lar region behind scapular bar red. Dorsal fin tral Mexico. A distribution map is provided by heavily pigmented with melanophores and all Contreras-Balderas (1975,1978). fins reddish. Scapular bar and gular stripe dark. Ecology. Very little information. Meek Tuberculation.— Dorsum of head covered (1904) thought spawning may occur in late

with moderately large, erect tubercles in semi- May and in June. scattered pattern. Tubercles arranged linearly Etymology. Named after Samuel Garman. in small and early season males. Postorbital region with some scattered tubercles, tending Comments. This species was thought by to obscure linear pattern. Snout with cluster of Meek (1904) and Contreras-Balderas erect tubercles equal in size to dorsal tubercles (1975,1978) possibly to be a subspecies of C. char- and separate from dorsum by hiatus. Above lutrensis. Because of the several unique I Miller 1 orbits, 4-5 moderately large organs arranged acteristics of this species, follow ( 978) in in single row. Remainder of head devoid of tu- and Chernoff and Miller (1982) recognizing bercles, except for few (2-3) small, erect or- this taxon as a distinct species. NORTH AMERICAN MINNOW GENUS CYPRINELLA 55

cyprinella formosa (glrard) ages; Santa Clara, Yaqui, and Santa Maria Beautiful Shiner generally 35-38, and Casas Grandes and Sauz 38^43); predorsal scales 15-26 (variable as Plate I above 17-19 and 19-24); total body circum- MonianaformosaGirard, 1857:201. [Orig. ferential scale rows 26-4 1 (Santa Clara, Yaqui, descr.; Type locality: Rio Mimbres, Chihua- Santa Maria 28-33, Sauz 3 1-34, Casas Gran- hua, Mexico = Mimbres River, north of Dem- des 33-38); scales above lateral line 13-20 ing, Luna Co. , NM (following Gilbert, 1978a)] (Sauz, Yaqui, Santa Maria 15-17, Santa Clara Notropis santamariae Evermann and 17, Casas Grandes 17-19), scales below lat-

Goldsborough, 1902:147. [Orig. descr.; Type eral line 10-19 (all 1 1-15, except Casas Gran- locality: small pool near Lake Santa Maria, des with 14-17); total caudal -peduncle cir- Chihuahua, Mexico] cumferential scales 14-24 (Sauz and Santa Notropis mearnsi Snyder, 1915:582-584. Clara 17-19, Yaqui and Santa Maria 15-18, [Orig. descr.; Type locality: San Bernardino Casas Grandes 1 8-22); above lateral line 7-11 River, near monument 77 of the international (all 7-9, except Casas Grandes with 9-11), boundary, Sonora, Mexico] below lateral line 5-11 (all 6-8, except Casas Diagnosis. Anal rays 8; lateral line usually Grandes with 7-9). incomplete; gular bar short; mandibular tu- Pharyngeal teeth 4-4, well hooked, and bercles present; breast fully scaled; dorsal with moderate grinding surfaces. Total gill margin of anguloarticular horizontal; epibran- rakers 7 or 8. Total vertebrae 35-37 (36), chial 4 with short mesial to dorsal process; precaudals 16-18 (17), caudals 18-20 (19). overlap between supraorbitals and lateral Dorsal fin insertion above vertebrae 11-13 ethmoids slight; basihyal deep; mentomecke- (12). lian deep. Lateral line complete or incomplete. Su- Description. A large shiner in the lutrensis pratemporal canal interrupted, pores 2. Su- clade, reaching a maximum size of about 70 praorbital canal without interruptions and mm SL. Body somewhat terete and deep; separate, pores 8-9 (9). Infraorbital canal in- dorsal profile slightly convex. Head large and complete in some before and below orbit, triangular. Snout pointed to slightly rounded, pores 9-11(10). Preoperculomandibular canal frequently rounded in the northern Rio Yaqui. complete, total pores 8- 1 1 (9- 1 0), mandibular

Mouth subterminal and slightly oblique, but pores 3^1 (3). variable; northern Rio Yaqui individuals with Intestine simple S -shaped loop with two terminal mouth, Rio Santa Maria populations flexures, Type I. Peritoneum silver with light with rounded snout and oblique mouth, and speckling of melanophores. those — from Rio Sauz and Laguna de Encinillas Coloration in life. Females and nonbreed- of northern Chihuahua with very oblique mouth ing males with less intense coloration than and round snout. Caudal peduncle moderately breeding males. Dorsum olivaceous to brown; deep. Dorsal fin not expanded in breeding remainder of body light yellow to cream. males and origin in both sexes above pelvic fin Lateral stripe present, but lighter than in breed- insertion. ing males. Fins clear to light orange. Dorsal fin rays 7-9 (usually 8); anal rays Breeding males dark and brilliantly col- 7-10 (8); pelvic rays 5-9 (8); pectoral rays ored. Coloration of dorsum variable within 10-14 (12-13); principal caudal rays 15-17 and between drainages (Chernoff and Miller, (17); dorsal procurrent rays 10-13 (12-13), 1982). Some breeding males with blue to green- ventral procurrent rays 9-12 (10-11). ish-blue dorsum and others yellow. Remain- Exposed margins of lateral scales slightly der of body yellowish, except for reddish- higher than wide and crenulate. Lateral-line orange caudal peduncle. Lateral stripe bluish scale rows 34^47 (variable between drain- and best developed posteriorly, diffuse anteri- 56 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

as 1 orly. Scales with pigmented edges, producing teriorly. Basally, pattern begins row of diamond-shaped pattern laterally and dorso- tightly compressed organs and increases to 4 laterally. Belly and breast cream. Gular stripe rows before segmentation. Between segmen- short and dark. Fins dark reddish-orange. tation and branching, 4 rows arranged in par- Tuberculation.— In nuptial males dorsum allel with 2-3 organs per row or 8-12 per of head covered with moderately sized, erect segment, with no breaks in pattern at segment tubercles in scattered pattern. Smaller males margins. Beyond branching, two rows devel- from same collections or males from earlier in oped per branch and 2-3 organs per row or 4-6 season with linear pattern as in young C. per segment. Tubercles extend almost to edge lutrensis. Linear pattern of these males later of fin. Paired fins with well-developed rows of obscured with further tubercle development to ventral tubercles. form scattered pattern, at least posteriorly. Distribution. Known only from ribs Casas Snout with several small, erect organs scat- Grandes, Santa Maria, del Carmen, and tered and separated from dorsal tubercles by Mimbres of the endorheic Guzman basin, and small hiatus. Preorbital region with 2-3 erect rios Yaqui, Sauz, and Bavicora of northwest- organs equal in size to dorsal head tubercles. ern Mexico and southern Arizona, U.S .A. Dis- Mandibular rami with 2-3 moderately large, tribution maps appear in Matthews (1980) and erect tubercles at posterior edge. Small, erect Chernoff and Miller (1982). organs present on upper one-third of opercle. Ecology. Little known other than gross Remainder of head devoid of tubercles. habitat characteristics and associated species. Body tuberculation extensive. Predorsal Habitat described by Miller and Simon (1943) region with 7-10 small, retrorse edge tubercles as small, turbid pools over sand, gravel, or per scale; no central or subcentral tubercles boulder substrate. In Rio Yaqui drainage Hen- present. Along flanks, 9-15 retrorse, small drickson et al. (1980) found the beautiful shiner edge tubercles above and below lateral line; no to be rare. Where present, most common in central organs present. Tubercles of caudal riffles of small streams and pools of intermit- peduncle organized into patch, more elaborate tent creeks. Uncommon in large rivers. than in any other member of lutrensis group. Etymology. The nameformosa is the latin Lower caudal peduncle and area above anal fin word meaning beautiful or pretty, in reference with similarly sized, retrorse tubercles (smaller to the breeding coloration of this species. than those on lutrensis) densely packed on Comments. The beautiful shiner has had a each scale. Dorsally, most scales with a series complex taxonomic history. Until recently, of up to 30 small retrorse edge tubercles. Early confusion existed as to the number of taxo- in development, lower caudal-peduncle scales nomic units involved in the complex (Chern- with only retrorse edge organs and a vertical off and Miller, 1982). Besides several races of row of central tubercles. Later, another row C. formosa sensu stricto recognized by Con- adjacent and medial to edge tubercles devel- treras-Balderas (1975), two forms were for- ops; still later, with further development of mally described in this complex, C. santamar- central tubercles this is obliterated and the iae and C. mearnsi. It appears that these may dense pattern develops. all represent a single form with a mosaic of All fins tuberculate. Dorsal and anal fins variable morphological characters. Character- with single row of 1-3 tubercles per segment istics considered diagnostic of C. santamariae on all rays basally and medially. Pelvic fins and C. mearnsi were investigated by Chernoff with same pattern, but restricted to rays 1-3. and Miller (1982) and found to vary inconsis- First ray of pectoral fin with single row of tently between and within populations. Thus, retrorse tubercles arranged with 2-3 per seg- I follow these authors in considering the two ment. Posterior rays 2-6 with fine retrorse forms as junior synonyms of C. formosa. organs in shagreen pattern, best developed an- Some have considered the beautiful shiner NORTH AMERICAN MINNOW GENUS CYPRINELLA 57 to be a subspecies of C. lutrensis (Contreras- brae 35-36 (36), precaudals 17-18 (17), cau- Balderas, 1975, 1978a; Gilbert, 1978a). dals 18 or 19. Dorsal fin insertion above verte- However, Chernoff and Miller (1982) demon- bra 12. strated its distinctiveness, and my analysis of Lateral line complete. Supratemporal canal Cyprinella indicates that C. formosa is more not interrupted at dorsal midline, pores 5-6 closely related to C. bocagrande than to C. (5). Supraorbital canal without interruptions lutrensis. and not joining other canals, pores 6-9 (7-8). Infraorbital canal complete, pores 11-17 (15). Cyprinella bocagrande (Chernoff Preoperculomandibular canal complete, total 9-12(10), mandibular pores 4-5 (4). and Miller) pores Intestine simple S -shaped loop and with Largemouth Shiner two flexures, Type I. Peritoneum silvery with Plate I light speckling of melanophores. in — nonbreed- Notropis bocagrande Chernoff and Miller, Coloration life. Females and 1982:514-519, Figs.2, 3. [Orig. descr.; Type ing males less intensely colored than breeding to olivaceous. locality: Ojo Solo, ca. 39 km W Villa Ahu- males. Dorsum silver-blue Venter mada, Chihuahua, Mexico] white to yellow. Lateral stripe as in males but Diagnosis. Anal rays 8; mouth extremely extending onto operculum. Dorsolateral stripe large; mandibular pores 4; tubercle hiatus light yellow. Anal fin yellow. Bases of pectoral between dorsum of head and snout absent; and pelvic fins yellow, remainder clear. scapular bar weak; gular bar short; ceratohyals Males brilliantly colored. Violet dorsally elongate; isthmus of maxilla very narrow; from head to caudal base, turning to iridescent posterior ramus of premaxilla straight; pha- light yellow dorsolaterally. Head rufescent. ryngeal pad ofbasioccipital triangular in shape. Plum-colored lateral stripe deeper and more Description. A large Cyprinella, adults diffuse anteriorly and intensely developed on reaching over 70 mm SL. Body robust. Predor- caudal peduncle. Scapular barabsent or weakly sal region and head of breeding adults deep developed. Caudal spot absent. Belly, breast, and gibbous. Head large and robust. Snout and ventrolateral body cream to yellow. Axil short, but mouth large and terminal to slightly of anal, pectoral, and pelvic fins typically or- subterminal. Upperjaw extending under orbit, ange. Ventral caudal peduncle red. Gular re- comprising more than 9.4% body length gion cream; gular stripe extending under orbit, (Chernoff and Miller, 1982). Caudal peduncle but not to isthmus. Dorsal fin clear. Remaining thick and short. Dorsal fin not expanded in fins yellow-orange. breeding males; origin in both sexes over or Tuberculation.— In nuptial males top of slightly posterior to pelvic fin insertion. head covered with several erect, moderately Dorsal fin rays 8-9 (8); anal rays 7-9 (8); large tubercles in scattered pattern. Early sea- pelvic rays 8-9 (8); pectoral rays 11-14 (13); son and younger males not available to deter- principal caudal rays 19; dorsal procurrent mine if linear pattern exists in early develop- rays 10-11 (11), ventral procurrent rays 9. ment. Dorsal organs extending laterally to, but Exposed margins of lateral scales deeper separate from, supraorbital cluster of erect, than long. Lateral-line scale rows 32^10 slightly smaller organs. Supraorbital tubercles (usually 36-37); predorsal scales 15-19 (18); continuous with preorbital tubercles by 1-2 body circumferential scales 29-36 (31-33); rows; preorbitals numbering 8-10 and located caudal-peduncle scales 15-20 (18). on dorsal half of lacrymal. Snout covered with Pharyngeal teeth 4-4; grinding surface of small, erect organs separate from dorsum by anterior tooth not well developed and tip not hiatus, connected in some males to preorbital hooked; others hooked and with grinding sur- organs by narrow band. Mandibles of some

faces. Total gill rakers 8-10 (9). Total verte- males with 1-5 semi-clustered, erect, small 58 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tubercles located centrally or laterally on rami. Etymology. The name bocagrande is taken Remainder of head devoid of organs, except from the Spanish boca for mouth and grande for occasional erect tubercle on dorsal opercu- for large, in reference to the diagnostically lum. large mouth of this species. Each predorsal scale scale with tubercles, scale rows of occi- best developed within 5-6 CYPRINELLA LEP1DA GlRARD 1-3 small, erect tubercles lo- put. Generally, Edwards Plateau Shiner cated centrally in semitransverse row; margins Plate I of scales armed with up to 18 erect submarginals. Rank scales with many erect to retrorse Cyprinella lepida Girard, 1857:197-198. edge organs per scale, both above and below [Orig. descr.; Type locality: Rio Frio (trib. Rio lateral line. No central or subcentral tubercles Nueces), either Real or Uvalde Co., TX] developed, except on some lateral-line scales, Diagnosis. Anal fin rays 9; pectoral fin rays where 1-2 may form. Above anal fin base 14; caudal-peduncle scales below lateral line scales like lateral body scales. Caudal peduncle 6-7; scapular bar strongly pigmented; gular pattern begins at posterior margin of anal fin bar short; supraethmoid process of palatine base. Lower caudal peduncle like C.formosa not strongly inflected above horizontal; poste- in having each scale with mat of crowded rior ramus of premaxilla straight; basihyal antrorse and erect organs, more complex than deep; posterior process of quadrate joins ver- in other members of this species group. Scales tical plate at edge; mentomeckelian long. of dorsal caudal peduncle like those of sides of Description. A small memberof the subge- body. nus, reaching a maximum of about 60 mm SL. Dorsal and anal fins with small erect organs Body terete, only slightly compressed; dor- on all rays. Anal fin with single row on first ray. sum slightly convex. Head large, subcorneal, Anal rays 2-7 with 1-2 rows of tubercles and convex above; horizontal below. Mouth maxilla basally, becoming shagreen before branching. small and inferior, posterior margin of After branching, often 2 rows per branch with to or slightly before orbit margin. Snout blunt 2-5 tubercles per segment. Rays 1-5 of pel- and protruding. Caudal peduncle short and vies with 1-2 rows of tubercles and 1-5 tu- moderately deep. Dorsal fin of breeding males bercles per segment, best developed on rays straight posteriorly, not expanded; dorsal ori- 2-3. Pectoral fin heavily tuberculate. Leading gin over to slightly posterior pelvic fin inser- ray with single row of tightly packed organs. tion. Tubercles on rays 2-5 (8) uniserial basally to Dorsal fin rays 7-8; anal rays 9-10 (9); 2 rows (2 per segment) before branching, pelvic rays 8; pectoral rays 12-14 (13-14); continuous with shagreen to edge. principal caudal rays 17; dorsal procurrent Distribution. Endemic to Ojo Solo, an rays 10-13 (11-12), ventral procurrent rays isolated spring near Ejido Rancho Nuevo in 8-11 (10). the Bolson de los Muertos of the Guzman Exposed margins of lateral body scales Basin, Chihuahua, Mexico. See Chernoff and higher than wide. Lateral-line scale rows 32-34 Miller (1982) for a distribution map of this (33-34); predorsal scales 14-16 (15-16); total species and C.formosa. circumferential scales 20-25 (21-23); circum- Ecology. Little known. Probably the rarest ferential scale rows above lateral line 11-13 member of the subgenus. Restricted to a single (12-13), below lateral line 9-12(10-11); total spring (diameter less than 50 m), its future caudal-peduncle scales 12-14 (13-14); cau- seems uncertain. Most specimens have been dal-peduncle scales above lateral line 7, below collected from deepest portions of the spring lateral line 5-7 (6-7). over a substrate of clay, sand, and/or mud Pharyngeal teeth 4-4, hooked, with moder- (Chernoff and Miller, 1982). ate grinding surface. Total gill rakers 7-8 (8). NORTH AMERICAN MINNOW GENUS CYPRINELLA 59

Total vertebrae 35-37 (36), precaudals 16-18 line, tubercles number 5-7 per scale; below in males. Predorsal (17), caudals 18-20 (19). Dorsal fin insertion only 3-6 develop peak above vertebra 11 or 12 (12). scales with a few erect, subcentral, and central Lateral line complete to hypural plate. located tubercles scattered like those in C. Supratemporal canal narrowly interrupted, lutrensis. Dorsal fin without tubercles. Anal fin pores 2-3 (2). Supraorbital canal complete, rays with 1 row and 1 pores 7-8 (8). Infraorbital canal complete, 1-5 tuberculate, per ray pores 8-11. Preoperculomandibular canal tubercle per segment. Pelvic ray 1 naked, rays complete, total pores 9 or 1 0, mandibular pores 2-3 with 1 row and 2-3 tubercles per segment. 3. First ray of pectoral fin with 1 row of fine, Intestine simple S -shaped loop, Type I. retrorse tubercles arranged with 2-3 per seg- Peritoneum brownish and/or silvery with ment. Pectoral rays 2-7 with 1 row of fine tu- speckled melanophores. bercles basally, grading to 2 rows before seg- — col- mentation. Between and branch- Coloration in life. Males brilliantly segmentation ored when in breeding condition. Dorsum ing, tubercles arranged in 2 rows with 2-3 2. greenish, lighter over head. Dorsolateral scales tubercles per row and segment, generally above dark lateral stripe yellowish-purple or After branching only 1 row per branch and 2-3 yellowish-blue from yellowish scales with per segment. Plateau shiner purple margins; below lateral stripe body much Distribuiton. The Edwards to this of lighter. Lateral body scales distinctly marked is endemic physiographic region with pigment arranged in diamond-shaped southwestern Texas. It is found in and is char- in the pattern. Side of head with golden-orange wash acteristic of clear springs and streams and purple vertical bar at anterior margin of upper tributaries of the Guadalupe River and preopercle; black gular stripe present, but short, most of the upper Nueces River System. extending to posterior margin of mandible. All Ecology. Inhabits clear, cool, spring-fed fins yellow to yellowish-orange; dorsal fin headwater creeks. Hubbs (1954) reported natu- heavily pigmented, yellow primarily restricted ral hybrids between this species and C. ve- to distal edge. nusta, and later (Hubbs, 1956) performed hy- Females and nonbreeding males also brid crosses between C. lepida and the closely brightly colored, but much less intense than related, but allopatric, C.proserpina, resulting breeding males; fins typically clear to slight in increased variability of meristic characters yellow with melanophores on rays and mem- in offspring. branes only. Lateral stripe present but more Etymology. The name lepida is derived diffuse anteriorly. from the Greek lepido, meaning having scales. Tuberculation.— Dorsum of head of nup- Comments. This species has had a trouble- tial males with scattered, erect tubercles pos- some taxonomic history. After its description

teriorly and in 2 rows anteriorly, extending to by Girard in 1857 as a distinct species, Jordan the posterior margin of orbit. Early males and and Evermann (1896) synonymized this spe- younger males with smaller, erect tubercles ar- cies with N. bubalinus. The species remained ranged in a linear pattern as in red shiner. Post- in synonomy until 1954 when C. Hubbs resur- and supraorbital row of antrorse tubercles rected C. lepida and determined bubalinus to extending to anterior margin of nares. Above be a junior synonym of C. lutrensis, as earlier orbit, organs number 5-6. Snout with a sparse noted by Hubbs and Ortenburger (1929). Al- clump of erect tubercles separated from dorsal though considered valid by C. Hubbs (1956, tubercles by hiatus. Remainder of head devoid 1958),Lytle(1972),MinckleyandLytle(1969), of tubercles. and Bailey et al. (1960, 1970), Hubbs (1972) Edge tubercles line most scales above and rejected it as a distinct species and synonymized below the lateral line on flanks. Above lateral it with N. lutrensis. This decision was based 60 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY primarily on its observed hybridization with ete, moderately deep. Depth greatest at or near N. lutrensis in the upper Guadalupe River, and dorsal origin. Head small. Snout blunt. Orbit because in west Texas numerous forms as small. Mouth small, terminal, and oblique; distinct as C. lepida could be recognized. Based lower jaw included in upper and both extend- on these observations, Hubbs (1972), Robins ing to anterior margin of orbit. Caudal peduncle et al. (1980) and Matthews (1980) followed in short and moderately deep. Dorsal fin not ex- considering this form a junior synonym of the panded in breeding males; origin in both sexes red shiner. slightly posterior to pelvic fin insertion. Data from this analysis indicate that the Dorsal fin rays 8 or 9; anal rays 7-9 (8); Edwards Plateau shiner is in fact a distinct pelvic rays 7-8 (8); pectoral rays 12-14 (13); species based both on its close relationship to principal caudal rays 17; dorsal procurrent the rutila clade and other morphological and rays 10-12(11), ventral procurrent rays 9-10 pigmentary characteristics. Further, since (10). hybridization may be a shared primitive trait Exposed areas of lateral body scales taller among organisms (Rosen, 1979), and because than wide. Lateral-line scale rows 32-36 hybridization in the Guadalupe River occurred (33-34); predorsal scales 14-15 (15); total only after dredging operations had started predorsal circumferential scales 21-24 (22); (Hubbs and Strawn, 1956), I do not consider scales above lateral line 12-15(13-15), below the evidence of hybridization in this case to be lateral line 9; total caudal-peduncle scale rows indicative of relatedness. It does, in fact, rep- 12-13 (12); caudal-peduncle scales above resent hybridization and not intergradation. lateral line 7, below lateral line 5-6 (5). Pharyngeal teeth 4-4, hooked, and with Cyprinella rutila (Girard) moderately developed grinding surfaces. Gill Mexican Shiner rakers very reduced, total on first arch 4-7 Plate II (5-6). Total vertebrae 35-36 (35), precaudal Montana rutila Girard, 1857:201. [Orig. 16-17 (16), caudal 18-20 (19). Dorsal fin

descr.; Type locality: Rio Monterrey (trib. Rio insertion above vertebra 1 1 . San Juan), Cadereita (=Cadereyta), Nuevo Lateral line complete to hypural plate. Leon, Mexico] Supratemporal canal narrowly interrupted or Moniana gracilis Girard, 1857:201. [Orig. occasionally complete, pores 2-3 (2). Supraor- descr.; Type locality: Acapulco, near Monter- bital canal complete and separate from other rey, Nuevo Leon, Mexico] canals, pores 7-8 (7). Infraorbital canal com- Cliola montiregis Cope, 1885:168. [Orig. plete, pores 9-11 (10). Preoperculomandibu- desc; Type locality: Monterrey, Nuevo Leon, lar canal complete, total pores 9-11 (9-10), Mexico] mandibular pores 3^4 (3). Diagnosis. Anal rays 8; gular bar elongate; Intestine simple with single S-shaped loop scapular bar weak; breeding males with purple and two flexures, Type I. Peritoneum dusky to and gold body coloration; lateral body scales speckled silver and brown.— above and below lateral line without tubercles; Coloration in life. Dorsum of females interoperculum short; coronoid process of and nonbreeding males dark, olive to brown, dentary broad; lateral slope of dermopterotics due to dark crosshatched pattern on scales and steep; anterior margin of ceratobranchial 1 darkened centers. Venter white to cream; lips expanded; symplectic narrow; anterior notch with scattered melanophores. Gular stripe long, of vomer shallow; mentomeckelian long; neck extending from symphysis to isthmus. Later-

on epibranchial 2 straight; frontals and der- ally, a variably developed blue-black stripe mopterotics not in contact. from caudal rays to snout, sometimes diffuse Description. A moderately sized Cyp- from dorsal origin anteriorly. On head, preor- rinella, reaching about 60 mm SL. Body ter- bital and postorbital portions of stripe may be NORTH AMERICAN MINNOW GENUS CYPRINELLA 61

obscured by darker pigmentation. Stripe bro- and pectoral fins. Anal fin with small, erect ken at hypural plate by vertical light stripe, organs on medial and basal portions of all rays creating caudal spot. Pre- and postdorsal stripes except first. First 4 pelvic rays with single row well developed. Fin pigmentation like that of of tubercles (2-3 organs per segment) medi-

N. xanthicara, but slightly more intense. ally. Leading pectoral fin ray with single row Breeding males as above, but with dark of retrorse tubercles (2-3 tubercles per seg- greenish-yellow dorsum and dorsolateral sur- ment). Rays 2-6 with single row basally, turn- faces. Dorsum of head and snout green. Lat- ing into double row before segmentation, and eral stripe less obvious because of intense triple row before branching. Each row with pigmentation, but visible as deep blue band. 2-3 organs per segment. After branching, Head bright yellow laterally. Gular stripe dark anterior branch with single row of 2-3 organs blue-black. Venter bright yellow. Fins bright per segment; posterior branch with 2 rows

yellow, mixed with milky-white deposits. proximally and single row distally. No ventral Dorsal fin sometimes very dark from high tubercles observed on paired fins. concentration of melanin. Distribution. Endemic to rios San Juan Tuberculation.— Dorsum of head of ma- and Salado of northeastern Mexico. Distribu-

ture breeding males with moderately large, tion map provided by Lytle (1972). slightly antrorse tubercles in scattered pattern. Ecology. Observations by Minckley and In smaller and early season males tubercles Lytle (1969) indicate that N. rutilus occurs erect and linear. Linear pattern consists of 2 above and below riffles, in current, over gravel rows extending from occiput to anterior mar- substrate. Concentrations of tuberculate males gin of nares, with some variation. Snout tu- are typically found in fastest riffles, while bercles few, same size as head tubercles, and females occur in adjacent eddies or pools with erect. Scattered snout tubercles separated from slower current. remainder of head tubercles by hiatus. Preobi- Etymology. The name rutila is derived tal areas with sparse, erect tubercles. Supraor- from Latin and means red. bital tubercles in single row, generally 5-6 Comments. Gilbert (1978a) listed Cliola moderately large, erect to slightly antrorse montiregis (Cope, 1 885) and Moniana gracilis organs. Remainder ofhead devoid of tubercles. (Girard, 1 857) as junior synonyms of C. lutren- Body tubercles restricted to caudal-peduncle sis. Recent examination of types of Cliola scales, scales above anal fin, and predorsal montiregis (ANSP 19344; 37.5 mm) and scales. Predorsal tubercles erect when small Moniana gracilis (MNHN 432; 39 mm) indi- and slightly antrorse when larger. Each scale cates that they are actually C. rutila and are generally with 2-3 organs located centrally in included here in the synonymy of this species. transverse row. Peripheral to these, a string of Both specimens have 8 anal rays and a dark submarginal tubercles only one-half size of gular bar extending form symphysis to the central organs develop around margin of scale. isthmus, typical of C. rutila. Caudal -peduncle tubercles restricted to 2-3 scale rows on lower half, including ventral row Cyprinella xanthicara (Minckley and 2-3 scale rows above anal fin. Each scale and Lytle) also with 1-6 semi-central organs and a series Cuatro Cienegas Shiner of submarginals. No organs present above lat- Plate II eral line. Young males and males early in breeding season with pattern like that of most Notropis xanthicara Minckley and Lytle, species of Cyprinella (except species of the 1969:491-502. [Orig. descr.; Type locality: pyrrhomelas and nivea species groups) with a Rio Puente Colorado, 8.5 km S and 0.7 km W single large tubercle per scale. Cuatro Cienegas, Coahuila, Mexico] Fin tuberculation restricted to anal, pelvic, Diagnosis. Anal rays 8; discrete blue-black 62 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY lateral band, 2 scales wide, extending from interrupted below eye, pores 9 or 10. Preoper- caudal fin to snout, excluding orbit; diamond- culomandibular canal complete, total pores shaped pigment pattern on scales above lateral 8-10 (9), mandibular pores 3. line only; gular bar elongate; scapular bar Intestine simple with two flexures forming weak; supraorbital tubercles antrorse; mandi- S-shaped loop, Type I. Peritoneum generally bular tubercles present; caudal-peduncle tu- black, may be heavily speckled with melanin bercles present on dorsalmost scale row; over silver background color. bar — uroneural 3 long; position of epiphyseal Coloration in life. Breeding and posteriorly placed; neck on anterior ceratohyal nonbreeding females and younger males simi- narrow; junction of sphenotic and frontals lar in coloration. Dorsum olive to brownish. posteriorly placed; posterior process of quad- Remainder of body pallid, except for darkly rate joins vertical plate at edge; ectopterygoid pigmented lateral stripe and pigmented mar- short and broad; posteroventral and ventral gins of lateral and dorsolateral scales, present- edges of preopercle entire; ceratohyals moder- ing crosshatched or diamond-shaped pattern. ately deep; branchiostegals narrow. Lateral stripe 1-2 scale rows wide, extending Description. A moderately sized Cyp- from center of caudal fin, through a slightly rinella, reaching about 60 mm SL. Body slen- enlarged caudal spot, to the posterior margin der, elongate, not obviously compressed; more of orbit; stripe continued anteriorly to orbit by delicate than other Cyprinella. Head slender preorbital bar on both sides of head. Lateral and pointed. Snout elongate. Mouth small, ter- stripe diffuse anteriorly on head. Venter light, minal, and oblique; lower jaw included in belly immaculate. Fins clear, except for few upper. Caudal peduncle slender and elongate. melanophores along rays. Procurrent caudal Dorsal fin not expanded in breeding males; rays and leading rays of dorsal, pectoral, and origin above pelvic fin insertion. pelvic fins darkly pigmented. Dorsal fin rays 8; anal rays 7-9 (8); pelvic Breeding males intensely colored. Minck- rays 7-8 (8); pectoral rays 12-14 (13-14); ley and Lytle (1969:498) described breeding principal caudal rays 17-18 (17); dorsal pro- males: "head brassy-gold over dorsum and current rays 9-1 1 (10), ventral procurrentrays onto snout, color interrupted on sides of snout 8-10 (10). and on opercles by dark melanophores, but Exposed margins of lateral body scales continuing ventrally on opercle; yellow con- higher than wide. Lateral-line scale rows 33-36 tinuing on dorsum to caudal fin, overlying (33-34); predorsal scales 14-16; total body gray or brown ground color that is distinctly circumferential scales 19-23 (21-22); dorsal cross-hatched. Lateral band intensely promi- circumferential scales 9-13 (13), ventral 9-10 nent; sides below lateral band pinkish-orange; (9); total caudal-peduncle scales 10-12 (12); belly white, slightly suffused with yellow; dorsal caudal-peduncle scales 5 or 7, ventral 5. pectoral fins lemon-yellow in center, color Pharyngeal teeth 4-4, slender, well hooked, more intense anteriad and with a dark black, and with serrated grinding surfaces. Gill rak- leading edge; pelvic fins with a dark leading ers reduced, total on first arch 4-6 (5). Total edge margined posteriad by an iridescent, bluish vertebrae 35-36 (36), precaudals 16-18 (17), line (milky-white in some), then [than] the caudals 18-19 (19). Dorsal fin insertion above remainder of the fin yellow; anal fin transpar- vertebra 11 or 12(11). ent distally and proximally, yellow centrally; Lateral line complete; supratemporal canal caudal fin intense yellow in belly of lobes, generally narrowly interrupted, but occasion- light yellow proximally, with procurrent rays ally complete or broadly interrupted, pores and central rays black; dorsal fin yellow-or- 2-5 (2). Supraorbital canal complete and sepa- ange, opaque, with milky-white pigments rate from other canals, pores 7 or 8. Infraorbi- proximally, edge with black; cornea yellow, tal canal generally complete, but occasionally reflecting blue dorsally; pupil jet black." NORTH AMERICAN MINNOW GENUS CYPRINELLA 63

Tuberdilation.— Dorsum of head in nup- ment. Ventral tubercles well developed on tial males with moderately large antrorse or- pectoral fins; none observed on pelvics. gans in scattered pattern from orbit to occiput; Distribution. Endemic to the CuatroCiene- 2 rows anteriorly from orbits to nares. Smaller gas basin of central Coahuila, north-central males and prenuptial males with slight tu- Mexico. See Minckley and Lytle (1969) for a bercle development and linear pattern down distribution map of this species. midline of head. Snout with scattered erect Ecology. The Cuatro Cienegas shiner in- tubercles, about one-half size of those on top habits the upper reaches of clear, cool streams of head and separated from head tubercles by of this region. It frequents areas between cur- hiatus. Preorbital area naked except for 1-3 rent and backwater pools. Presumably it is a tubercles on anterior lacrymal, connected to sight feeder. Minckley and Lytle (1969) ob- supraorbital tubercles by single row. Supraor- served surface feeding, and it is known to bital tubercles and those below nares smaller inspect passing items frequently. During the than dorsal head tubercles and arranged in day it is very active in midwater or near the

row. with of small in at it rests single Mandibles single row , bottom current, but night on the erect tubercles. Remainder of head devoid of substrate (Minckley and Lytle, 1969). tubercles. Etymology. The epithet xanthicara is in

Minckley and Ly tie ( 1 969) brief! y described reference to the breeding coloration devel- tuberculation: their observations are similar to oped in males of this species; from Greek mine, except they noted granular development xanthos, for yellow and kara, for head. on cheek and gular regions and noted no development on the nape. CYPRINELLA PROSERPINA (GlRARD) Predorsal scales tuberculate with a pattern Proserpine Shiner similar to C. rutila. Generally 2-3 antrorse Plate II l organs, about /i size of dorsal head tubercles, centrally in a semitransverse row. Peripheral Montana proserpina Girard, 1857:200. to these there are submarginal and edge an- [Orig. descr.; Type locality: Devils River, just trorse organs in decreasing size. Along flanks, above mouth into Rio Grande, Val Verde Co., each scale above lateral line with 6-10 small, TX] erect organs; below lateral line organs retrorse. Moniana aurata Girard, 1857:200. [Orig. Caudal peduncle tuberculation similar to C. descr.; Type locality: Pinto Creek, "Piedra rutila, except 5-8 retrorse, edge organs found Pinta," 1 5-20 mi SE Del Rio, Kinney Co., TX] on each scale both above and below lateral Diagnosis. Anal rays 8; mouth inferior; line. scapular bar well-developed; gular bar elon- All fins tuberculate. Dorsal fin with first gate; breeding males with yellow-orange and two rays tuberculate; other rays with single purple body coloration; premaxillary process row of tubercles medially and a single organ of maxilla tall; isthmus of maxilla moderately per segment. All anal fin rays with a single row shallow; insertion of adductor mandibulae Al and 1 tubercle per segment along entire length. on maxilla below isthmus; posterior process of Pelvic fin with first 4 rays tuberculate; each quadrate deep and heavy; supracleithrum with a single row of tubercles and 1 tubercle broad. per segment (occasionally 2). First ray of Description. Small for a Cyprinella, but pectoral fin with a single row of tubercles and size average among members of the lutrensis 2 organs per segment; posterior rays 2-7 with clade, reaching a maximum body size of 55-60 single file basally to segmentation where 2 mm SL. Body robust, not strongly compressed. rows begin . After segmentation, 2 rows formed Body relatively slender, maximum depth with 4 organs per segment. Single row main- equally developed from occiput to dorsal fin tained on each branch; 2 tubercles per seg- origin. Head large and highly arched dorsally, 64 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY horizontal ventrally. Snout blunt and protrud- purple along midline laterally. Lateral scales ing over jaws. Mouth strongly inferior, well marked along edges with dark crescentic lines, below orbit; lower jaw included in upper. creating a crosshatched pattern. Purplish-blue Caudal peduncle relatively short and stout. lateral stripe developed from caudal base to Dorsal fin not expanded in breeding males and opercle, best developed posteriorly, diffuse white over or slightly posterior to pelvic fin inser- anteriorly. Ventrally, postcephalic regions tion. to cream, except predominantly bright yellow Dorsal fin rays 7-9 (8); anal rays 7-9 (8); or yellow-orange above anal fin. Dorsum of pelvic rays 7-8 (8); pectoral rays 12-14 head dark greenish; laterally brassy-rose, turn- (13-14); principal caudal rays 15-17 (17); ing to light purple ventrally. Gular stripe dark, dorsal procurrent rays 11-14 (11-12), ventral extending from symphysis to branchiostegals procurrent rays 9-12 (10-11). at isthmus. Dorsal fin bluish anteriorly, red- Exposed margins of lateral scales slightly dish-pink posteriorly; leading rays whitish. higher than wide. Lateral-line scale rows 34-36 Caudal, anal, and pelvics yellowish to yel- white (35); predorsal scales 15-18 (16-17); total lowish-orange basally and centrally, predorsal circumferential scales 22-27 distally. Pectorals greenish dorsally and or- (24-25); scales above lateral line 11-13 ange ventrally. — males with linear (1 1-12), below lateral line 9-12 (10-1 1); total Tuberdilation. Early caudal-peduncle circumferential scales 14; pattern of erect tubercles on dorsum of head. above lateral line 7, below lateral line 5. Top of head of nuptial males covered with Pharyngeal teeth 4-4, strongly hooked, with moderately large, erectorgans. Anteriorly some antrorse. hiatus between well developed grinding surfaces. Total gill tubercles slightly No rakers 5-10 (7-8). Total vertebrae 35-37 (36), tubercles on snout and top of head, complete precaudal 16-18 (17 in Devils R. and Inde- from occiput to lips. Supraorbital row (6-7 pendence Cr.; 18 in Pecos R.), caudals 17-20 erect organs) connected to preorbital tubercles (19 in Devils R. and 18 in Independence Cr. by single row. Preorbital organs few (2-3) and and Pecos R.). Dorsal insertion above vertebra erect. Mandibular rami and other areas of head 11 or 12 (11). devoid of tubercles. Lateral line complete. Supratemporal canal Predorsal scales with 2-3 antrorse edge interrupted narrowly, pores 2-3 (3). Supraor- tubercles one-half size of dorsal head organs; bital canal separate and complete, pores 7-10 no submarginal or central organs developed. 1 erect tubercles (7). Infraorbital canal complete, occasionally Along flanks, 5-1 small, edge separate from trunk canal, pores 10-12 (1 1-12). developed per scale both above and below Preoperculomandibular canal complete, total lateral line; no central or submarginal organs. like that of C. pores 8-10 (9), mandibular pores 2-3 (3). Caudal peduncle tuberculation Intestine simple, with single S-shaped loop, rutila, except for several (2-8) small, retrorse lateral none Type I. Peritoneum heavily colored with edge organs per scale, below line; melanophores over silvery background. above. — All fins tuberculate. Dorsal all with Coloration in life. Females and nonbreed- rays ing males like breeding males but less in- single row of tubercles and single tubercle per tensely colored. Olive dorsally, white or cream segment, except first rudimentary ray devoid and un- ventrally, and bluish-gray laterally. Cross- of tubercles. First 3 rays (branched hatched pattern present, as well as lateral and branched) ofpel vie fins tuberculate, with single gular stripes, but not as brightly developed as row of tubercles and 2-3 organs per segment. in breeding males. First pectoral ray with single row of tubercles tubercles Breeding males brilliantly colored (Hubbs and single tubercle per segment; and Miller, 1978; pers. obs.). Dorsum and closely set basally and more widely separated 1 3 dorsolateral areas bright brassy green; bluish- distally. Rays 2-7 with row basally and (4) NORTH AMERICAN MINNOW GENUS CYPRINELLA 65 before branching, where there may be 4-9 isthmus of maxilla very narrow; supraethmoid organs per segment. After branching 1 row on narrow and with emarginate lateral edges; in- anterior branch (2 tubercles per segment) and fraorbital 4 extremely narrow; autopterotic 2 rows on posterior branch (4 tubercles per spine small; dorsal margin of anguloarticular segment). Venter of paired fins with several horizontal; dorsal supraethmoid process of erect tubercles per ray. palatine not strongly elevated above horizon- Distribution. The proserpine shiner is re- tal. stricted to tributaries of the lower Rio Grande Description. A small Cyprinella, reaching (including Pecos and Devils rivers) and San a maximum of about 55 mm SL. Body robust, Felipi, Pinto, and Las Moras creeks in Texas, not extremely compressed. Head large. Mouth and Rio San Carlos, Coahuila, Mexico. Ros- small, inferior, terminal, and oblique; lower ter's (1957) report of this species from New jaw included in upper, both extending poste- Mexico is apparently in error (Hubbs and rior to or slightly before orbit. Snout distinctly Miller, 1978). It seems reasonable, however, rounded and protruding. Back highly arched that it probably did occur in the upper Pecos at from occiput to dorsal origin. Caudal peduncle some time before increased aridity. thickened and moderately long. Dorsal fin not Ecology. Little is known of this species' expanded in breeding males; origin over to biology. It prefers clear and cool streams with slightly posterior insertion of pelvic fin. a sand-gravel substrate. Harrell (1978) ob- Dorsal fin rays 8-9 (8); anal rays 8-9 (8); served benthic sight feeding and noted repro- pelvic rays 7-8 (8); pectoral rays 11-14 duction after a period of severe flooding. Hubbs (12-13); principal caudal rays 18-19 (19); (1956) experimented with hybridization be- dorsal procurrent rays 10-11(11), ventral pro- tween this species and C. lepida, two allopatric current rays 9-10 (9). but closely related forms in the lutrensis clade, Exposed areas of lateral body scales elon- and noted increased variability in meristic gate, higher than wide. Lateral-line scale rows characters of the offspring. 32-35 (33-34); total predorsal circumferen- Etymology. This species was apparently tial scales 25-29 (26-27); circumferential named after Persephone, daughter of Zeus and scales above lateral line 12-16 (13-14), below Demeter, wife of Pluto and queen of the infer- lateral line 10-13 (10-11); total caudal- nal regions. The name proserpina is the lati- peduncle scales 14-15 (14); scales above lat- nized form of her name. eral line 7, scales below 5-6 (5). Pharyngeal teeth 4-4, each usually strongly Cyprinella panarcys (Hubbs and to weakly hooked, with well-developed grind- surfaces; anteriormost tooth of each arch Miller) ing sometimes smaller, weakly hooked, and Conchos Shiner may lack grinding surface. Total gill rakers 6-9 (8). Plate II Total vertebrae 34-36 (35), precaudals 16-17 Notropis panarcys Hubbs and Miller, (17), caudals 17-19 (17-18). Dorsal insertion 1978:582-589. [Orig. descr.; Type locality: above vertebra 11 or 12 (11). Rio San Pedro at Meoqui, Chihuahua, along Lateral line complete to hypural plate. Hwy 45 at elevation 1115 m] Supratemporal canal narrowly to widely inter- Diagnosis. Anal rays 8; black subcrescen- rupted in most specimens with minor seg- tic markings on flank scale pockets; scapular ments on either side of midline, pores 2 per bar well developed; gular bar elongate; maxi- side. Supraorbital canal complete, pores 8. mum body depth from occiput to dorsal origin; Infraorbital canal complete, frequently forked mouth inferior; breeding males with bluish- posterior to orbit with branch above eye, pores purple and golden body coloration; fins or- 9 or 10. Preoperculomandibular canal gener- ange-white; neck of epibranchial 3 straight; ally complete, but may be reduced on ascend- 66 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

and ing arm of preopercle, total pores 10, mandi- of retrorse organs per ray (Hubbs Miller, bular pores 3. 1978). Intestine single S -shaped loop with two Distribution. Known only from the upper of the flexures, Type I. Peritoneum dark, covered Rio Conchos system, a major tributary with heavy concentration of melanophores. Rio Grande in northern Mexico. See Hubbs — Miller for distribution of this Coloration in life. Dorsum of females and (1978) map and nonbreeding males dark olive to brownish- species. and Miller green, fading to light cream or white on venter. Ecology. Hubbs (1978) reported Lateral stripe bluish, best developed posteri- collecting this species from shallow, generally clear over a or mud sub- orly, diffuse or absent anteriorly. Scapular bar pools rocky, sandy, of tuberculate males in rectangular, not offset posteriorly by lighter, strate. Collections fully rectangular depigmented area. Dorsum of head March were from shallow and broad portions dark olive, extending down sides to and in- of Rio San Pedro over gravel substrate. These also list associated cluding lips and dorsum of operculum. Venter authors species. unpigmented, except for darkly pigmented Etymology. The name panarcys derived and mean- gular stripe or bar extending from symphysis from Greek paw, meaning all, arcys to isthmus. ing a net, describing the crescentic pattern on Body of breeding males greenish-blue or lateral body scales suggestive of a net. purple dorsally. Center of each flank scale blue or to Elevated purple, margin golden greenish. Cyprinella spiloptera (Cope) scales intensely marked with blackish cres- Spotfin Shiner cents, a crescentic, crosshatched pat- creating Plate II tern extending down to belly. Head dorsally in Giinther colored as described above, dusky laterally. Leuciscus spilopterus Cope, Postorbital area, opercle, subopercle, interop- 1868:254. [Orig. descr.; Type locality: St. ercle, and preopercle orange, most intense on Joseph R., MI (southwest part of state)] former two. Fins orange to brownish orange. Hybopsis fretensis Cope, 1869:382. [Orig. Pectoral, anal, and dorsal darkened along bases desc; Type locality: near Detroit, MTJ and leading rays by melanophores. Pectoral, Notropis spilopterus hypsisomata Gibbs, anal, and pelvic fins with conspicuous milky- 1957b: 195-198. [Orig. descr.; Type locality: white margins, less intense in dorsal and cau- Wonder Lake, bay in north end, McHenry Co., dal fins. IL] Tuber culation.— In nuptial males dorsum Diagnosis. Anal rays 8; gular bar short; fins of head from occiput to tip of snout, lateral to of breeding males yellow; posterior hyoideal dorsal margins of orbit, covered with scat- foramen of ceratohyal located mesially; der- tered, moderately large, erect organs. Supraor- mopterotics invade parietals; posterior proc- 1 with bital tubercles 4-6, connected to preorbitals by ess of vomer narrow; ceratobranchial broadened single row of erect organs. Snout covered with restricted neck; frontals widely small, erect organs continuous with preorbital laterally. tubercles. Remainder of head devoid of tu- Description. A relatively large Cyprinella, bercles. reaching a maximum of about 100 mm SL. Body tuberculauon as in C. proserpina. Body depth variable between subspecies: hyp- Tubercles on predorsal scales, lower 1-2 scale sisomata deep-bodied and spiloptera elon- indi- rows of caudal peduncle, above anal fin, and gate. Body terete, athough deeper bodied on scales above lateral line anterior to dorsal viduals somewhat compressed. Head relatively

fin origin. small. Snout conical and protruding, espe- small. Fins apparently devoid of tubercles, except cially inbreeding males. Orbitrelatively and for pectorals. Anterior rays with a single row Mouth terminal or subterminal, oblique NORTH AMERICAN MINNOW GENUS CYPRINELLA 67 moderately large. Lowerjaw barely contained Pectoral fin immaculate, except for first ray in upper, both extending posteriorly to anterior which may be lined with melanophores. margin of orbit. Caudal peduncle relatively Breeding males darker and more colorful. short and heavy. Dorsal fin not greatly ex- Dorsum and flanks deep steel-blue. Snout panded in breeding males; origin in both sexes yellow. Lateral stripe and scapular bar dark slightly posterior to pelvic fin insertion. blue, broadening posteriorly. Lateral stripe Dorsal fin rays generally 8; anal rays 7-9 expanded midlaterally between dorsal origin

(8); pelvic rays 8; pectoral rays 10-17 (13-14); and operculum into a large, round darkened principal caudal rays 17; dorsal procurrent spot. All fins with milky-white deposits, espe- rays 10-12(11), ventral procurrent rays 10-11 cially distally; dorsal fin very dark with high (10). concentrations of melanophores on mem- Lateral body scales higher than wide, ap- branes, especially posteriorly. Pelvic, anal, pearing elevated, especially anteriorly near and caudal fins yellow. lateral line. Lateral-line scale rows usually Tuberculation.— Dorsum of head of nup- 35-39 (spiloptera 37-39, hypsisomata 36 or tial males covered with scattered, moderately 37); total circumferential scales 26-28; scales large, antrorse organs. Tubercles arranged in above lateral line 13-15 (13 in most, but some semilinear pattern in early season males, but populations of spiloptera with high frequency obliterated later in season with increased tu- of 14 or 15), below lateral line usually 11; total bercle development. Supraorbital region with caudal-peduncle scales generally 14; scales scattered antrorse organs equal in size to dor- above lateral line 7, scales below lateral line 5. sal tubercles. Preorbital tubercles moderately Pharyngeal teeth 1, 4-4, 1, hooked, and large, scattered, antrorse, and connected to with or without serrated grinding surfaces. supraorbital tubercles by 1-2 rows and to

Total gill rakers 8-1 1 (9). Total vertebrae 35-38 snout tubercles by narrow band. Snout organs (37), precaudals 17-18 (18), caudals 18-20 scattered and clustered, antrorse posteriorly (19). Dorsal fin insertion above vertebra 12. and erect anteriorly, and separated from dorsal Supratemporal canal broadly interrupted, head tubercles by wide hiatus. Mandibles with pores 2 per side. Supraorbital canal complete, 2 rows of moderately small, antrorse organs. pores 8-10 (8). Infraorbital canal complete or Preopercles with few, scattered, erect organs. narrowly interrupted between lacrymal and Remainder of head naked. infraorbital 2, pores 11-13. Preoperculomandi- Each scale of predorsal region with 2-3 bular canal complete, total pores 12, mandibu- antrorse organs, equal in size to dorsal head lar pores 4. tubercles, arranged in transverse row arched Intestine arranged in simple S -shaped loop, across center of scale. Some predorsal scales Type I. Peritoneum silvery and speckled with with 1^4 subcentral organs; no edge tubercles melanophores. — present. Scales along flanks with 0-6 erect, Coloration in life. Females and nonbreed- subcentral organs above and below lateral ing males olive or light bluish dorsally and line; some antrorse near nape. No central tu- dorsolaterally. Dorsolateral and lateral scales bercles present except near lateral line, nape, outlined with dark crescents, presenting dia- or caudal peduncle. One to 5 antrorse, subcen- mond-shaped pattern. Lateral stripe best de- tral tubercles and 2-3 central, antrorse tu- veloped posteriorly, diffuse and broad anteri- bercles on scales on ventral half of caudal orly from dorsal origin to head; may be dark in peduncle and around anal fin base. Similar some. Scapular bar dark. Venter immaculate. pattern above lateral line on caudal peduncle, Gular bar short and dark. Dorsal fin lightly but organs smaller and equal in size. pigmented with melanophores, increasing in All rays of dorsal and anal fins tuberculate. darkness with increasing body size and onset Each ray with 1 row and 1 tubercle per seg- of breeding season. Pelvic and anal fins clear. ment medially. One row and 2-3 tubercles per 68 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY segment on pelvic fin. First ray of pectoral fins permanent flow; not typical of larger turbid with 1 row and 1 tubercle per segment. Poste- rivers or intermittent creeks. Usually found in rior rays 2-8 with 1 row basally and 2-3 or near riffles or raceways over gravel sub- tubercles per segment continuous with single strate in moderate to fast current. row on posterior branch. Anterior branch with Spawning occurs from May or early June to one row and 2-3 organs per segment. No mid-August, with peak activity in June and organs observed on ventral surfaces of paired early July (Pfiieger, 1965, 1975). Males con- fins. gregate and maintain territorial regions around Distribution. The spotfin shiner is widely submerged logs and exposed tree roots near distributed in eastern North America and has a riffles in swift current. When ready to spawn, distribution similar to the suggested extent of a female moves into one of these areas from a preglacial highland fauna (Mayden, 1985). downstream, pairs with a male, and attaches Two subspecies of the spotfin shiner were eggs to undersides of logs or branches in recognized by Gibbs (1957b), C. s. spiloptera crevices (e.g., under loose bark) (Hankinson, and C. s. hypsisomata. The validity of these 1930; Pfiieger, 1965). Spawning activity and taxa, however, has been challenged recently species recognition are enhanced by sounds by Schaefer and Cavender (1986). These au- produced by this species (Winn and Stout, thors proposed that the forms recognized by 1960). Gibbs represent one taxon with clinal vari- Most individuals live only 2+ years, but ation in meristics and body proportions. I some are known to live as long as 5 years follow these authors, but present the ranges of (Stone, 1940). Maturity is apparently reached the forms as presented by Gibbs (1957b). The during the first year, but some do not spawn eastern spotfin shiner, C. s. spiloptera, ranges until the second year (Pfiieger, 1965). throughout the Ohio River drainage north- Diet of this species commonly includes ward to the Great Lakes basin, including Lakes terrestrial insects, such as ants, adult and Huron, Erie, and Ontario, and the St. Law- immature aquatic insects, plant materials, and rence River drainage in Quebec. This form small fishes (Forbes and Richardson, 1908; also occurs along the Atlantic Slope from the Starrett, 1950; White andWallace, 1973).Most Potomac to the Hudson rivers. The western feeding occurs shortly before dusk (White and subspecies, C. s. hypsisomata, occurs in the Wallace, 1 973) and probably shortly after dawn. upper Mississippi River Basin of northern Hybridization of this species with the red Missouri, Illinois, Wisconsin, and Minnesota. shiner in Illinois was discussed by Page and Southern populations are found in the Ozark Smith (1970). The gene pool of the spotfin Plateaus in the Meramec, Gasconade, and shiner in this region was rapidly diluted with Arkansas rivers. Intergrades, based on inter- morphological characteristics, and presuma- mediate lateral scale counts, are known only bly the genome, of C. lutrensis. from eastern and western drainages of Lake Etymology. The name spiloptera is from Michigan. A detailed distribution map is pro- Greek spilos, for spot, and pteron, meaning vided by Gibbs (1957b) and Lee et al. (1980). wing or fin, describing the heavily pigmented Ecology. The biology of this species is last 2 interradial membranes of the dorsal fi n of fairly well documented. Studies include those breeding males. of Hankinson (1930), Stone (1940), Starrett Cyprinella camura (Jordan and and as well as (1950, 1951), Pfiieger (1965), Meek) comments by Trautman (1957), Cross (1967), Bluntface Shiner Pfiieger (1975), Gale and Gale (1977), and Plate II Smith (1979). Cyprinella spiloptera most frequently oc- Cliola camura Jordan and Meek, curs in large creeks and small rivers with clear, 1884:474-475. [Orig. descr.; Type locality: NORTH AMERICAN MINNOW GENUS CYPRINELLA 69

probably Neosho River drainage in Lyons Co., precaudals 17-20 (18-19), caudals 18-21 Kansas (Gilbert, 1978a)] (19-20). Dorsal fin insertion above vertebra Diagnosis. Anal rays 9; snout blunt; depig- 12. mented vertical bar at base of caudal fin; snout Lateral line complete. Supratemporal canal and dorsal fin of breeding males red; hiatus broadly interrupted, pores 2 per side. Supraor- between dorsal head and snout tubercles ab- bital canal complete, pores 9-10 (9). Infraor- sent; tubercle connection between supraorbi- bital canal complete, pores 11. Preoperculo- tal and preorbital regions narrow; mandibles mandibular canal complete, total pores 12-13 of breeding males with 2 rows of tubercles; (12), mandibular pores 4. anterior neck of parasphenoid short; ascend- Intestine arranged in simple S -shaped loop. ing wings of parasphenoid very broad; ba- Peritoneum silvery with heavy speckling of sibranchial one with no median constriction; melanophores. — neck of ceratobranchial two long and thin; Coloration in life. Females and nonbreed- neck on ceratobranchial one bent mesially; su- ing males primarily silver-blue. Dorsum olive praethmoid extremely short; frontals extremely to steel-blue, fighter laterally to primarily sil- broad; posterior wing of hyomandibular nar- ver. Belly immaculate. Weak lateral stripe row; preopercle very wide over entire length; from near caudal base to opercle. Anterior to ceratohyals very deep. dorsal fin, lateral stripe weakly developed.

Description. A moderately large and stout Lateral stripe terminates posteriorly at light, member of the whipplei clade, reaching about unpigmented bar at caudal base, extending full 90 mm SL. Body deep and compressed, great- length of hypural plate and onto dorsal and est depth at dorsal origin. Head large and ventral procurrent rays. Lateral scales nar- subcorneal. Snout blunt. Orbit small, diameter rowly outlined by melanophores, presenting less than snout length. Mouth slightly subter- crosshatched pattern. Dorsum of head plum- minal, oblique. Lower jaw contained in upper, beous, extending ventrally to include dorsum

both extending to or slightly beyond anterior of opercle and preopercle, lips, and lacrymal. margin of orbit. Caudal peduncle deep and Remainder of head white to silver, except for

relatively short. Dorsal fin expanded and darkly pigmented and short gular stripe. Pelvic rounded posteriorly in breeding males; origin and anal fins clear. Dorsal fin dusky anteriorly, over to slightly posterior to pelvic fin inser- darkly pigmented on last 2 membranes. Lead- tion. ing pectoral rays sometimes outlined with Dorsal fin rays 8; anal rays 8-10 (9); pelvic melanophores. Caudal fin dusky, except for rays 8; pectoral rays 13-17 (15 or 16); princi- basal unpigmented region. pal caudal rays 16-17 (17); dorsal procurrent Breeding males bright steel-blue dorsally rays 10-12 (11), ventral procurrent rays 9-11 and dorsolateral^, each scale outlined with (10). blue crescents. Laterally, dark, blue-black Exposed margins of lateral and dorsolateral scapular bar well developed and separate from scales taller than wide. Lateral-line scales darkened lateral stripe by broad, lightly col- 35-39 (36-37); total circumferential scales ored region. Posterior to light region, lateral 24-28 (26); scales above lateral line 11-15(13 stripe expanded into large dark spot. Head in eastern populations, but frequently 14 or 15 yellowish, except for reddish snout and plum- in Arkansas R. specimens), scales below lat- beous dorsum. Dorsal fin bluish basally and eral line 11 generally (some 12 or 13); total centrally; pink distally. Interradial membranes caudal-peduncle scales 14; scales above lat- darkened, especially posteriorly. Pectoral, eral line 7, below lateral line 5. pelvic, and anal fins salmon-pink with milky- Pharyngeal teeth 1, 4-4, 1, hooked, and white distal margins. Caudal fin white basally with or without grinding surfaces. Total gill and bluish distally, but pinkish over most of rakers 8-9 (9). Total vertebrae 35-39 (37-38), fin. 70 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Tuberdilation.— Dorsum of head covered (Illinois, Neosho, and Arkansas rivers) in with moderately large scattered, antrorse tu- Arkansas, Missouri, Kansas, and Oklahoma. bercles. Early in development, however, tu- Gibbs (1961) provided a distribution map of bercles arranged linearly. Snout, preorbital, this species. This map also included an eastern and supraorbital areas covered with moder- Colorado locality, originally thought to be the in ately large, antrorse organs, all interconnected. type locality, and a population the Osage Mandibular tubercles smaller than dorsal head River System (Missouri R. Drainage) and the organs, erect, and arranged in 2 irregular rows. White River. The type locality is suspect (see Interopercle, preopercle, branchiostegals, and Comments) and Cross (1967) doubted the gular area with few small and erect organs validity of the latter two localities, suggesting arranged in linear pattern. Suborbital region, erroneous locality information. This seems operculum, suboperculum, and opercular logical since both the Osage and White rivers membrane naked. have been thoroughly sampled for years and Body tuberculation extensive. Predorsal no additional records of this species are known scales with 2-3 central, erect, and moderately to exist. large organs, generally arranged in a slight arc Ecology. Bailey and Taylor (1950), Cook perpendicular to body axis. Peripheral to these (1959),andBurrandMayden(1978)described are several erect, subcentral tubercles; no edge the habitat of eastern populations, and Moore organs present. Caudal -peduncle scales simi- (1952), Cross (1967), Cavin (1971), Miller lar to predorsal scales, but central organs scat- andRobison (1973), and Pflieger(1975) noted tered and number 2-5. Submarginals range up habitat associations for western populations. to 20. Caudal-peduncle tubercles small and Both populations occur in high-gradient, fast erect. Along flanks, tubercles small and erect to moderately-fast clear water, and not low- and arranged subcentrally (6-12) and cen- gradient habitats typical of much of the Coastal fast trally (2-10) on a scale, both above and below Plain province. Adults generally prefer lateral line. riffle and raceway habitats. Younger individu- All fins tuberculate. Dorsal and anal fins als are typically in or close to these habitats, rays all bearing 1 row and 1 organ per segment. but also frequent shallow pools. In eastern Pelvic fins with first 4 rays tuberculate, each populations this species may be found over a with 1 row and 1-3 tubercles per segment. sandy substrate, whereas in the west they First ray of pectoral fin with single row of typically occur over gravel or rubble, not over small, retrorse organs, 1 per segment. Rays sand or mud. Cavin (1971) noted that this 2-9 with single row of tubercles to branching species has a low tolerance for oxygen stress. of ray and 2-3 per segment. After branching, Little is known about this species' life his- anterior and posterior branches with single tory, except for notes by Cook (1959), Cross row. Anterior branch with 1 tubercle per seg- (1967), Pflieger (1975), and a study by Cavin ment and posterior with 2 tubercles per seg- (1971). In Kansas most individuals live only 2 ment. Paired fins with small, erect organs years, but some may live 3 years. Age classes arranged in rows on ventral side of anterior and I comprise a large proportion of popula- rays. tions. Young of the year have a high growth Distribution. There are two disjunct popu- rate, reaching about 18 mm, with a maximum lations of the bluntface shiner, one east and one of 45 mm by the end of their first summer. By west of the Mississippi River. Eastern popula- the end of the second summer most average tions occur in direct, west-flowing tributaries about 50 mm, with a maximum of about 75 of the Mississippi River and eastern tributaries mm. of the lower Tennessee River, all within the In Kansas and Missouri, the spawning Coastal Plain province. Western populations season spans late May to late July (Cross, occur in the middle Arkansas River drainage 1967; Pflieger, 1975). Ovary weight begins to NORTH AMERICAN MINNOW GENUS CYPRINELLA 71

increase in late June and reaches a peak in (e.g., Notropis heterolepis, N. topeka, Phox- middle to late July, with a maximum GSI inus) have subsequently disappeared. Thus, (Gonadosomatic Index) of 16.4. Ovaries de- more extensive western range of C camura is creased in size in Augustand September (Cavin, conceivable. 1971). As noted by Gale and Buynak (1978) Because of the disjunct nature of this spe- for C. analostana, mature oocyte diameter of cies, Gibbs (1961) examined differentiation C camura decreased with progression of breed- between the two populations. These popula- ing season from 0.88 in middle to late June to tions were found to differ at the "racial" level 0.82 in late July. Spawning has been observed in meristic and morphometric characteristics. and consists of crevice spawning similar to Perhaps further studies may warrant subspecific that of other Cyprinella species (Rabito and or even species recognition of these forms. Heins, 1985). The name is derived Etymology. camura, Cyprinella whipplei Girard from the Latin camur, meaning turned inward, Steelcolor Shiner referring to the blunt snout (Gibbs, 1961). Plate in Comments. The type locality of this species is in question. Jordan and Meek (1884) in Cyprinella whipplei Girard, 1857:198. the original description designated Fort Lyon, [Orig. descr.; Type locality: Sugarloaf Creek, Colorado, as the type locality. This locality is trib. Poteau River, ca. 20-25 mi S Fort Smith, west of the present range of C camura by AR, vicinity ofPoteau, OK (following Gilbert, several hundred miles. As a possible explana- 1978a)] tion for this disjunction, Gibbs (1961) sug- Diagnosis. Anal rays 9; pectoral rays usu- gested that the species range has become con- ally 15; dorsal fin of breeding males without stricted in recent years and the Ozark localities bright coloration, other fins yellow; breeding are the only remaining populations. Gilbert males with a single row of mandibular tu- (1978a), however, suggested two alternatives. bercles; dorsal margin of anguloarticular hori- Perhaps Jordan and Meek meant Lyons, Kan- zontal; isthmus of maxilla very narrow; hori- sas, or Lyon County, Kansas. The latter is the zontal plate of urohyal broad; basihyal deep; more likely of the two. Today this species is operculum narrow and deep; symplectic nar- abundant in the Cottonwood River system, row; interoperculum narrow; mentomeckelian Lyon Co., Kansas, but is not found within long. several miles of the former locality. The Lyon Description. A large Cyprinella, reaching Co., Kansas, locality is supported by the ab- a maximum length of about 130 mm SL. Body sence of the bluntface shiner in collections by slightly compressed and moderately deep, Gilbert and Cragin (see Cross, 1967) from deepest in breeding males. Head small and Garden City, Kansas, where other cool and subconical. Snout short, pointed, and protrud- clear-water species were collected. Further, ing, especially in breeding males. Orbit mod- Jordan (1891) did not report this species from erately small. Mouth terminal to subterminal; collections in the Arkansas River system in jaws nearly equal and extending posterior to Colorado, but listed the species from the anterior margin of orbit. Caudal peduncle deep "Arkansas River at Witchita." It is possible, and elongate. Dorsal fin gready expanded in however, that Gibbs is correct and the Colo- breeding males; origin slightly posterior to rado locality is the type locality, especially pelvic fin insertion.

since Etheostoma cragini, a species with a Dorsal fin rays 8; anal rays 8-10 (9); pecto- distribution like western C. camura, is known ral rays 14-16 (15); pelvic rays 8; principal from eastern Colorado. Further, other fishes caudal rays 17; dorsal procurrent rays 10-12 reported from clear, spring-fed streams along (11), ventral procurrent rays 9-11 (10). the Arkansas valley in the late 19th century Lateral scales with exposed margins deeper 72 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY than wide, especially anteriorly near lateral moderately large and antrorse organs formed line. Lateral-line scales 36-40 (37-38); total on either side of midline from occiput to nares. predorsal circumferential scales generally 26; Snout tubercles equal in size and antrorse scales above lateral line 11-15 (13), below posteriorly, grading to erect anteriorly and lateral line usually 11; total caudal -peduncle separate from dorsal head organs by a distinct scales 14; caudal-peduncle scales above and hiatus. Preorbital organs moderately large and below lateral line 7 and 5, respectively. crowded, as in C. galactura, and connected to Pharyngeal teeth 1, 4-4, 1, hooked, with supraorbitals by 2-3 rows of antrorse organs. narrow serrated grinding surfaces. Total gill Supraorbital tubercles moderately large and rakers 8-10 (8). Total vertebrae 38-40 (39), antrorse. Tubercles on preopercles, interop- precaudals 19-20 (19), caudals 19-20 (20). ercles,branchiostegals, and gular region small, Dorsal fin insertion above vertebra 12 or 13. scattered, and erect. Remainder of head naked. Lateral line complete. Supratemporal canal Predorsal tubercles numerous and erect, broadly to narrowly interrupted, pores 2 per each scale generally containing central, subcen- side. Supraorbital canal complete, pores 8. tral, and edge organs. Central tubercles larg- Infraorbital canal occasionally interrupted est. Caudal-peduncle tubercles present con- below orbit, but generally complete, pores tinuously around body, but best developed 10-11 (11). Preoperculomandibular canal ventrally. Generally 10-15 scattered subcen- complete, total pores 11, mandibular pores 4. tral and 2-6 central organs per scale. Along Intestine with simple S -shaped loop and flanks, both above and below lateral line, each two flexures, Type I. Peritoneum with silver scale with 7-12 erect, subcentral, and 3^ background color and heavily speckled with central organs. melanophores. — Dorsal fin rays with 1 row and 1 organ per Coloration in life. Females and nonbreed- segment. Anal fin rays with a single row and ing males olive to steel-blue dorsally and 2-3 tubercles per segment. First four rays of dorsolateral^, becoming lighter ventrally. pelvic fin with single row and 2-3 per seg- Body silver-blue laterally, except for dark lat- ment. Leading ray of pectoral fin with single eral stripe. Stripe best developed posteriorly, row and one organ per segment, extending continued anteriorly to opercular margin from most of length of ray. Posterior rays 2-7 with dorsal fin origin by broad, diffuse band and single row of retrorse and crowded organs, dark dorsal margin. Venter cream to white. continuing onto posterior branch. Each seg- Head olive to blue dorsally and light ventrally. ment, after branching, with 2-3 organs. Before Lacrymal, dorsal preopercle, and opercle and branching and on anterior branch, 1 row and subopercle colored like dorsum, but lighter. single organ per segment. Pectoral and pelvic Lips and gular stripe pigmented. Fins lightly fins with erect organs on ventral surfaces. pigmented on membranes, except dorsal, which Distribution. The steelcolor shiner, appar- is dusky anteriorly and darkly pigmented ently widespread in the pre-glacial highland posteriorly, forming dark spot on last 2 mem- regions of eastern North America, is found branes. today only in the Interior and Eastern High- Dorsum of breeding males bright steel- lands, with a few northern disjunct popula- blue. Laterally and ventrally body iridescent tions. Its present distribution is typical of many bluish-purple. Snout red. All fins lemon-yel- species whose ranges indicate northern glacial low and milky-white. Dorsal fin heavily pig- alterations of this once widespread range mented posteriorly, presenting large dark spot. (Mayden, 1985; Wiley and Mayden, 1985). In Caudal and base of anal more dusky. Leading the west it is found in the unglaciated Ou- rays of paired fins pigmented. achita, Kiamichi, Little, middle-lower Arkan- Tuberculation.— Dorsum of head in peak sas tributaries, White, and Meramec rivers. males as in C. analostana, with clusters of East of the Mississippi, it is found in the NORTH AMERICAN MINNOW GENUS CYPRINELLA 73

Tennessee and Cumberland rivers, and ungla- off others, he and the female approach the ciated portions of the Ohio. Some populations spawning area side by side, with the male are known from the once glaciated Wabash, slightly above the female. During the spawn- Illinois, and Allegheny river systems. For ing act(s) the male presses the female against summary maps of the distribution, see Gibbs the log and the two vibrate while traveling (1963) and Lee etal. (1980). along the log, releasing eggs and milt simulta- Ecology. Typically the steelcolor shiner neously. Males frequently eat eggs from the inhabits unmodified and treelined, moderate nest areas. The same behavior has been ob- to high gradient streams and rivers. Generally, served in aquaria when eggs do not remain at- it is not found in small creeks or large rivers. tached. Over 100 eggs were found in stomachs Most frequently it occurs over a gravel sub- of some males. strate in riffles and raceways (Gibbs, 1963; The spotfin shiner is frequently observed Pflieger, 1975; Smith, 1979). spawning near the steelcolor shiner, but the The steelcolor shiner is a sight-feeding two species have never been observed inter- species that schools near the top or middle of mingling. Typically, the spawning peak of C. the water column. Its diet typically consists of spiloptera occurs earlier than C. whipplei drifting aquatic or terrestrial insects, but it also (Trautman, 1957; Pflieger, 1965). Further, these feeds on small crustaceans, mites, and earth- species have species-specific vocalizations worms, some of which are picked from the (Winn and Stout, 1960). substrate (Forbes and Richardson, 1908; Etymology. This species is named after Lt. Pflieger, 1975). A. W. Whipple, commander of the expedition Most individuals live only 3 years. Some that collected the type series. females may live up to 4 years (Pflieger, 1965). females the Males attain a larger size than by Cyprinella analostana Girard endof their second summer. Maturity is reached Satinfin Shiner the second summer, but most individuals do by Plate in not spawn until the third. The smallest mature males are about 49 mm; smallest females are Cyprinella analostana Girard, 1 859:58-59. about 38 mm (Pflieger, 1965). [Orig. descr.; Type locality: Rock Creek, trib. The spawning season in Missouri and Ohio of Potomac River, Washington, DC] is from late May to mid-August (Pflieger, Diagnosis. Anal rays 9; pectoral rays usu- 1965, 1975). Trautman (1957) and Pflieger ally 13 or 14; pharyngeal process of basioc- (1965, 1975) noted spawning in fast and shal- cipital spathulate and deep; dermopterotics low water, above the substrate, on the under- extend over dialator fossa; supraorbitals nar- sides, upper surfaces, or lower lateral surfaces row anteriorly and posteriorly; posterior proc- of submerged logs. Eggs are deposited be- ess of vomer narrow; posterior subopercle neath loose bark and/or in crevices between greatly elevated; supraethmoid elongate; ante- roots and logs. Several males (20-30) may be rior margin of frontals smooth; ceratohyals congregated, fighting and/or spawning, around very deep. the log, with as many or more females found Description. A moderately large Cyp- downstream below the log. Males maintain rinella, adults reaching about 90 mm SL. Body nonstationary territories around the spawning deep, especially in breeding males, and mod- surface and defend these areas from other erately compressed. Head triangular. Snout males with threatening displays and occasional pointed, protruding slightly. Mouth terminal battering with their tuberculate heads. Females, to subterminal and oblique, extending posteri- when ready to spawn, swim in or near the orly to anterior margin of orbit; lower jaw spawning area and are approached by one to contained in upper jaw. Caudal peduncle thick several males. After a male succeeds in fighting and relatively long. Dorsal fin expanded in 74 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY breeding males; origin in both sexes slightly pigments in all fins. Dorsal fin darkly pig- concentrated posterior to pelvic fin insertion. mented, melanophores heavily Dorsal fin rays 8; anal rays 7-10 (9); pelvic in last 2-3 membranes. Anal base, caudal fin, fin All fins with faint rays 8; pectoral rays 11-16 (13-14); principal and pectoral more dusky. caudal rays 16-17 (17); dorsal procuiTent rays yellow or gold color, more pronounced on 9-12 (10), ventral procurrentrays 8-10(9-10). pelvics, anal, and lower caudal. Body deep Lateral scales higher than wide, appearing steel-blue dorsally and iridescent bluish-violet elevated. Lateral-line scales 33-38 (clinal, laterally and ventrally, lighter in latter region. generally 36-37 in north to 35-36 in south); Lateral stripe darker, especially on posterior total predorsal circumferential scales gener- half of body (Denoncourt and Messersmith, Lateral into ally 24; scales above lateral line 11-15 (13), 1982). stripe expanded large spot below lateral line 11; total caudal-peduncle between dorsal fin and opercular membrane. scales 14; scales above and below lateral line Tuberculation.— Gibbs (1963) and De- described 7 and 5, respectively. noncourt and Messersmith (1982) head Pharyngeal teeth 1,4-4, 1, hooked, with tuberculation of this species. Dorsally, narrow, sometimes serrated, grinding surfaces. pattern of young males semilinear; tubercles erect. with 2 Total gill rakers 7-8 (7). Total vertebrae 35-36 Peak nuptial males, however, tubercle concentration lateral (36), precaudals 18, caudals 17-18 (18). Dor- distinct areas of sal insertion above vertebra 11 or 12. to midline and anterior to posterior margin of Lateral line complete. Supratemporal canal orbit. Dorsal tubercles of these males large and tubercles broadly interrupted, pores 8-9. Supraorbital antrorse. Between orbit and occiput canal complete, pores 2 per side. Infraorbital scattered, but generally in 2 lateral, concen- Preorbital and re- canal complete, pores 1 1 . Preoperculomandi- trated patches. supraobital total 1 1 mandibu- with antrorse bular canal complete, pores , gions moderately large, organs, lar pores 4. sparsely distributed as in C. camura. These Intestine arranged in simple S-shaped loop, regions connected by single row of equal- naris on ei- Type I. Peritoneum silvery and speckled with sized tubercles between orbit and melanophores. ther side. Erect and scattered snout organs — from dorsal head tubercles small Coloration in life. Females and nonbreed- separate by ing males like C. whipplei. Dorsum and dorso- hiatus. Except for the 2 rows of antrorse, lateral surfaces olive to silvery-blue, becom- moderately large organs on each mandible, ing lighter ventrally. Dorsum of head plum- remainder of head naked. beous down to lacrymal, upper opercle, and Body tuberculation extensive. Predorsal preopercle. Venter cream to white, except for tubercle pattern similar to that of C. camura each scale pigmented lips and short, darkly pigmented and C. galactura, except generally tubercles. Predorsal gular stripe. Lateral scales above and below with fewer and larger lateral line outlined with dark pigment, creat- organs antrorse and moderately large and range best de- ing diamond- shaped pattern. Lateral stripe not from 2-6. Generally, central organs well developed, but strongest posterior to dorsal veloped, subcentral organs only occasionally erect formed origin. Stripe diffuse anteriorly, broad, and formed. Up to 8 subcentral, organs best developed dorsally as a narrow snipe to per scale above and up to 6 scale rows below opercle. Dorsal fin lightly pigmented; melano- lateral line along flanks. Occasionally small, phores strongest on posterior membranes. erect central tubercle developed on each scale Caudal fin dusky with membrane pigment. on or near lateral line. Caudal peduncle tuber- similar to that Anal fin with some pigment basally. Pelvic fins culation well developed. Pattern immaculate. of Mississippi River drainage relatives, except Breeding males with more intense diamond- for a reduction of centrally located organs to 1 8 subcentral and 1-2 shaped lateral scale pattern and milky-white or 2. Each scale with up to NORTH AMERICAN MINNOW GENUS CYPRINELLA 75 antrorse, central tubercles. Central organs high in May-July (2. 12-2.66) and are lowest sometimes slightly larger. This pattern con- from September to December (2-2.22). Age tinuous around caudal peduncle, but not well group 2 females have their highest values in developed above lateral line. May and June, while group 1 are later in July, Dorsal fin without tubercles. All anal fin indicating an earlier spawn by older age groups. rays with single row and 1 organ per segment. This pattern was similarly noted by Stone Pelvic rays with 1 row and 1 retrorse tubercle (1940), but Gale and Buynak (1978) found no per segment. Leading pectoral ray with a single data to support this hypothesis. GSI values are row and tubercle per segment; rays 2-6 with 1 greatest for both sexes in May and June, drop- row and 1 retrorse organ per segment before ping sharply and maintaining a low level from branching; occasionally 2 per segment. Be- July to December, indicating the end of spawn- yond branching, single row continuous with ing activities in July. posterior branch, 1 organ per segment; ante- Ovary weights are greatest in June and rior branch with single row and tubercle per decrease considerably, along with mean size segment. No ventral organs observed on paired of ova, by the end of July. During most of the fins. year four distinct size classes of ova are appar-

Distribution. The satinfin shiner is, with ent in ovaries, except in late July when only the exception of some populations in the south- three are present. Mean diameter ofova ranged ern Lake Ontario drainages, an Atlantic Slope from 0.027 to 0.91 mm. Fecundity ranged endemic. This species is found in streams from from 545 to 3344 eggs, with a mean potential the Peedee River System in North Carolina fecundity of 1598 per female. The actual fe- north to the Hudson River. Gibbs (1963) and cundity was, however, estimated to be closer Lee et al. (1980) presented distribution maps to 877 eggs per female. Gale and Buynak of this species. (1978) and Stone (1940) suggested a slightly Ecology. Considerable data exist on the lower fecundity, 6-634 or 409-864 eggs per biology of this species. The habitat of the female, respectively. satinfin shiner is similar to that of C. whipplei, Cyprinella analostana is a fractional crev- generally in moderately small to large streams ice spawner, spawning several times in a sea- over a substrate of gravel, rubble, and/or sand. son and depositing its eggs in crevices under This species can occasionally be found in or loose submerged bark, between exposed tree near tidal areas of some rivers. roots, in crevices in rocks and wood, or under Denoncourt and Messersmith (1982) de- flat rocks (Stone, 1940; Gale and Buynak, tailed many aspects of the life history of this 1978). This species, like others of the genus, is species in Pennsylvania. Most individuals have known to produce vocalizations used in spe- a normal life span of 3 years, some reach 4. A cies and sex recognition (Stout and Winn, mean fork length (snout to caudal fork) for age 1958; Stout, 1959; Winn and Stout, 1960). group 1+ was 58.6 mm, 67 mm for 2+, and 84 Males guard the nest area, protecting it from mm for 3+. Growth rates are similar for age other males by rapidly swimming around the classes, with no significant sexual dimorphism. site and towards the intruding male with fins About 70% of adult body size was attained by erect. Defending males also produce a knock- the end of the first year. Breeding in Maryland ing sound. When approached by a receptive spans from May to late August (Stout and female a purring sound is produced by males Winn, 1958). In Pennsylvania, spawning lasts and spawning occurs (Stout, 1959; Winn and from late May to early July, as evidenced from Stout, 1960). The spawning and prespawning GSI (Gonadosomatic Index), condition fac- behaviors of this species are similar to those of tors, and breeding tuberculation and colors the spotfin shiner, C. spiloptera (Gale and (Denoncourt and Messersmith, 1982). Condi- Gale, 1977; Gale and Buynak, 1978). Under tion factors are greatest in females, reach a controlled conditions a given pair may spawn 76 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

3-11 times in a season with 6-634 eggs re- bital tubercles in single row. leased each time. Time between spawnings Description. One of the smallest members ranged from 3 to 3 1 days, being bimodal for 5 of the genus, reaching a maximum length of and 8 days. Apparently most eggs are released about 60 mm SL. Body deep, especially in during these spawning peaks and only a few breeding males and females, and compressed; are spawned between, unlike the behavior of maximum depth at dorsal origin. Head moder- small. C. spiloptera, which displays no peak in activ- ately small and triangular. Orbit Snout short in ity (Gale and Gale, 1977). Freshly spawned relatively and pointed, especially peak eggs average about 1 .5 mm in diameter, but as males where it protrudes over lips. Mouth the season progresses the mean diameter de- terminal and oblique, extending posteriorly to creases (Gale and Buynak, 1978). orbit margin. Lower jaw contained in upper Eggs hatch in 6-8 days at 20-25°C (Gale jaw. Caudal peduncle deep and relatively short. and Buynak, 1978) or within 11 days at Dorsal fin of breeding males expanded slightly. 20-20.5°C (Stone, 1940). After hatching lar- Dorsal fin rays 8; anal rays 7-9 (8); pelvic vae are inactive until 6 days posthatching, rays 8; pectoral rays 11-17 (14); principal when they are about 6 mm long. At this time caudal rays 16-17(17); dorsal procurrent rays they begin to feed. 9-12 (1 1), ventral procurrentrays 8-10 (9-10). Etymology. The name analostana derives Lateral body scales elevated, margins taller from Analostan Island in the Potomac River, than wide. Lateral -line scale rows 32-36 (35); Washington, D. C., the location where Girard total predorsal circumferential scales gener- lateral line 11-13 originally collected this species. ally 24; scales above (11), Comments. Populations of this species from below lateral line 11; total caudal-peduncle the Peedee River System are considered inter- scales 14; scales above and below lateral line grades by Gibbs (1963) with the Santee River 7 and 5, respectively. endemic, C. chloristia. With the exception of Pharyngeal teeth 1, 4-4, 1, hooked, with 6 individuals, however, all specimens from narrow, sometimes serrated, grinding surfaces. Total vertebrae 35-37 this drainage are typical for C. analostana. On Total gill rakers 7-9 (8). this basis, Gibbs (1963) relegated C. chloristia (36), precaudals 17-19 (18), caudals 17-19 to a subspecies of C. analostana. Based on di- (18). Dorsal fin insertion above vertebra 11 or agnostic meristic differences presented by 12. Gibbs (1963) and osteology (presented herein), Lateral line complete. Supratemporal canal I prefer not to follow this treatment and follow broadly interrupted, pores 2 per side. Supraor- Bailey et al. (1970) and Robins et al. (1980) in bital canal complete, pores 8-9 (8). Infraorbi- considering C. chloristia distinct at the species tal canal complete, pores 9-12 (11). Preoper- level. culomandibular canal complete, total pores 10, mandibular pores 3. Cyprinella chloristia (Jordan and Intestine arranged in single S-shaped loop, Brayton) Type I. Peritoneum silvery with scattered Greenfin Shiner melanophores. Coloration in — and Plate in life. Nonbreeding breeding individuals essentially like the north- Codoma chloristia Jordan and Brayton, ern sister species C. analostana. Gibbs (1963) of the 1878:21-23. [Orig. descr.; Type locality: Sa- noted that the lateral stripe greenfin luda River, Farr's Mills, W of Greenville, shiner was more narrow and better developed. Greenville Co., SC] He associated this difference with the observa- inhabits clearer Diagnosis. Anal rays 8; mandibular pores tion that C. chloristia typically C. analostana. 3; neck of epibranchial 1 bent posteriorly; streams than — Dorsum of head with premaxillary process of maxilla short; supraor- Tuberculation. NORTH AMERICAN MINNOW GENUS CYPRINELLA 77

moderately large, antrorse tubercles in a semi- species is probably similar to C. whipplei and scattered pattern from orbit to occiput. Tu- C. analostana. Cyprinella chloristia appears bercles arranged into 2 clusters, following to inhabit small rivers and streams of Montane areas where rows are normally developed on and upper Piedmont provinces, differing from either side of midline of head. Anterior to its closest relative C. analostana, which is posterior margin of orbits, arrangement linear typically associated with large riverine habi- like C. analostana and C. whipplei. Preorbital tats. organs few, antrorse, and connected to su- Etymology. The name chloristia is from praorbital organs by single row. Supraorbital the Greek chloros, for green, and histion for tubercles 5-6, antrorse, moderately large, and sail, in reference to the greenish color of the arranged in single row. Snout tubercles sepa- dorsal fin observed by Jordan and Brayton. rate from dorsal head organs by a distinct Comments. This species was considered a hiatus; posterior organs antrorse and change to subspecies of C. analostana by Gibbs (1963) erect anteriorly. Mandibular tubercles erect and later elevated to species status by Bailey et and somewhat scattered, but occasionally in 2 al. (1970) and Robins et al. (1980). Gibbs rows. Remainder of head devoid of tubercles. (1963) united these species on the basis of an Predorsal region with a tubercle pattern as hypothesized area of intergradation between in and the taxa in the River C. whipplei, C. analostana , C. venusta, two Peedee drainage. C. galactura. Caudal peduncle pattern like C. Data supporting intergradation were from 6 of analostana; central tubercles larger than 189 specimens examined, all of which ap- subcentrals and reduced to 1-3. Along flanks, peared intermediate. Other collections were above lateral line, scales lined with 2-8 erect, typical of "normal" C. analostana in meristic small organs; below lateral line each scale data, although some clinal variation was ob- lined with 2-8 antrorse small organs. No cen- served. Here I follow Bailey et al. (1970), tral tubercles developed, except occasionally Gilbert and Burgess (1980), and Robins et al. on lateral-line scales and near caudal peduncle (1980) in considering this form a distinct spe- region. cies and suggest that additional populational- All rays of dorsal and anal fins tuberculate. level studies be conducted to determine if gene First rudimentary ray tuberculate along full flow has contributed to the intermediate condi- length. Posterior rays with varying degrees of tions of some Peedee River specimens, as tubercle development; anterior rays generally suggested by Gibbs (1963). Even if headwater have greatest development, with 1 row along capture transfer introduced some C. chloristia ray medially and 1 organ per segment. Only into the Peedee, it is doubtful that the limited first 3 rays of pelvic fins tuberculate; each ray extent of this possible "hybridization" event, with 1 row extending entire length and 1 organ not intergradation, should warrant uniting these per segment. First ray of pectoral fin with two forms as a single evolutionary unit. row and 1 retrorse single organ per segment. Cyprinella venusta Girard Rays 2-6 with single row per ray and usually Blacktail Shiner single retrorse tubercle per segment; occa- Plate IB sionally 2 tubercles per segment. No ventral organs observed on either paired fins. CyprinellavenustaGirard, 1857:198. [Grig, Distribution. Endemic to the Santee River descr.; Type locality: Rio Sabinal at Sabinal, drainage of North and South Carolina, above Uvalde Co., TX] the Fall Line. Gibbs (1963) and Lee et al. Cyprinella notata Girard, 1857: 198. [Orig. (1980) provide distribution maps of this spe- descr.; Type locality: Rio Seco (=Seco Creek) cies. at D'Hanis, Medina Co., TX] Ecology. Limited information exists. Ex- Cyprinella cercostigma Cope, 1868:157. cept for habitat preference, the biology of this [Orig. descr.; Type locality: Pearl River, 78 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Monticello, Lawrence Co., MS] peak males. Mouth terminal or subterminal, Photogeniseurystomus Jordan, 1877b:356. oblique, large; jaws equal and extend to ante- [Orig. descr.; Type locality: Nancys Creek, rior margin of orbit. Caudal-peduncle depth trib. Chattahoochee River, Atlanta, Fulton Co., and length variable between subspecies, ve- GA] nusta deepest and stigmatura more slender. Photogenis stigmaturus Jordan, 1877b:337. Dorsal fin origin posterior to pelvic fin inser- fin males. [Orig. descr.; Type locality: trib. Etowah River tion; somewhat expanded inbreeding (probably Silver Creek), near Rome, Floyd Dorsal fin rays generally 8; anal rays 8 Co., GA] (except some stigmatura with high frequency Cyprinella calliura Jordan, 1877c:61-62. of 9); pelvic rays 8; pectoral rays 14-15 (high cau- [Orig. descr.; Type locality: Alabama River frequency of venusta with 13); principal system, Selma, Dallas Co., AL. Originally dal rays 17; dorsal procurrent rays 11, ventral given as Black Warrior River, but this river is procurrent rays 9-1 1 (10). not near Selma (after Gilbert, 1978a)] Exposed margins of lateral scales taller Photogenis leucopus Jordan and Brayton, than wide. Lateral-line scale rows 34^48 1878:41-42. [Orig. descr.; Type locality: (usually 36-43, but variable between Chattahoochee River at Shallow Ford, NW of subspecies: venusta 37 or less, stigmatura Gainesville, Hall Co., GA] 40-43, cercostigma 38-40); total predorsal Luxilus chickasavensis Hay, 1881:506. circumferential scales generally 24-26; scales [Orig. descr.; Type locality: Chickasavensis above lateral line 13-17 (15, although gener- in lateral line 1 1 total River, Enterprise, Clark Co., MS] ally 13 venusta), below ; Cliola urostigma Jordan and Meek, caudal-peduncle scales generally 14; above 1884:474-477. [Orig. descr.; Type locality: lateral line 7, below lateral line 5.

San Saba River (trib. Colorado River), at Fort Pharyngeal teeth 1, 4-4, 1, hooked, and McKavit (McKavett), ca. 45 air mi SE San with or without serrations on narrow grinding Total verte- Angelo, Menard Co., TX] surfaces. Total gill rakers 7-9 (7). Notropis cooglei Hildebrand and Towers, brae 37-39 (38), precaudals 19-20 (19), cau- 1928:18. [Orig. descr.; Type locality: Pelucia dals 18-20 (19). Dorsal fin insertion above Creek, S of Greenwood, Leflore Co., MS] vertebra 12 or 13 (13). Diagnosis. Anal rays 8; caudal spot large Lateral line complete. Supratemporal canal and dark; scales below lateral line along flanks complete or broadly to narrowly interrupted, without central tubercles; mesial neck of pores 2 per side when interrupted or 5 when epibranchial 1 bent posteriorly; retroarticular complete. Supraorbital canal complete, pores long and L-shaped; insertion of adductor 8. Infraorbital canal complete, pores 10 or 1 1 . mandibulae Al on maxilla below isthmus; Preoperculomandibular canal complete, total hypohyal foramen restricted to anterior cera- pores 10, mandibular pores 4. tohyal; overlap between supraorbitals and lat- Intestine simple S -shaped loop, Type I. thick of eral ethmoids slight; ligament connection Peritneum silvery with speckling between epibranchial 1 and 2 mesial to unci- melanophores. — non- nate process of epibranchial 2. Coloration in life. Females and Description. One of the largest members breeding males similarly colored. Dorsum dark near of the genus, reaching about 1 50 mm SL. Body bluish to olive, extending down sides to slender and elongate to moderately deep, but lateral band, becoming lighter. Lateral band varies between subspecies; venusta generally dark and narrow posteriorly, fading and broad- deepest and stigmatura most slender form. ened anteriorly where the most obvious devel- forward Head moderately large, but variable, stigma- opment is a narrow line continuing tura with shortest head. Orbit moderately small, from dorsal margin. Posteriorly on caudal smallest in venusta. Snout subcorneal, acute in peduncle, lateral band very distinct, with well- NORTH AMERICAN MINNOW GENUS CYPRINELLA 79 developed caudal spot which may be rounded in peak males; first ray with erect tubercles or squarish, depending on race. Venter unpig- over full length. Posterior rays with single row mented, except for short, dark gular bar, and and 1 per segment, best developed medially. pigmented lips, anal fin base, and lower caudal Pelvic fin rays 2-6 tuberculate medially, con- peduncle. Scapular bar weakly developed. sisting of single row and 1 organ per segment. Lateral and dorsolateral scales with pigmented Pectoral fin with retrorse tubercles; first ray edges, forming diamond-shaped pattern on with single row and 1 tubercle per segment. sides. Lateral line pores more heavily pig- Rays 2-8 each with single row basally of mented anteriorly, producing dotted pattern small, crowded organs, continuing onto poste- along path of canal. Fins clear, except base of rior branch (generally 2-3 per segment). Ante- anal and caudal and leading ray of pectoral. rior branch with single organ per segment. No Dorsal fin dusky with melanophores on all ventral organs noted on paired fins. membranes, increasing in intensity posteri- Distribution. The blacktail shiner is a North orly. American Coastal Plain endemic, ranging from Breeding males brilliant steel-blue dorsally, the Rio Grande, Texas, east along Gulf of less laterally. Lateral band very dark and ex- Mexico to Suwannee River, Florida, and north panded into large spot between dorsal fin and in the Mississippi Embayment to southern opercle, immediately posterior to a large lightly Illinois. The western subspecies, C. v. venusta, pigmented region. Fins with milky-white is distributed west of the Mississippi River and coating. Anal and caudal fins yellow and dor- smaller, direct eastern tributaries. Cyprinella sal darkly pigmented. v. cercostigma is distributed along the Gulf Tuberculation.— Dorsum of head covered Coast east of the Mississippi River from Lake with many moderately large, antrorse organs Ponchartrain, Louisiana, drainage to the in scattered pattern. Preorbital, suborbital, and Suwannee River. Cyprinella v. stigmatura is snout regions with similar sized organs. Preor- found in the upper Mobile Bay System in the bital tubercles thickly concentrated and con- Alabama and Tombigbee rivers, primarily nected to supraorbitals by wide band, 2-3 above the Fall Line. Intergrades between the organs wide. Snout organs concentrated in latter two subspecies have been proposed from clump, separate from dorsal tubercles by hia- the Cahaba, Alabama, and Tallapoosa rivers. tus; erect anteriorly and antrorse posteriorly. Ecology. Very little is known for such a Mandibular region with 2 semilinear rows of wide ranging, common, and evolutionarily tubercles medially. Except for small semian- interesting species. Typically it inhabits streams trorse organs scattered on preopercle, remain- or moderately sized rivers with sparse vegeta- der of head naked. tion, strong current, and sand or gravel sub- Tubercles on body scales well developed. strate. In Missouri, spawning occurs from June Predorsal scales each with 2-3 centrally lo- to August. Eggs are deposited in crevices, as in antrorse in transverse territories cated, organs pattern other species of Cyprinella , and are and 2-4 subcentral organs around perimeter. defended by males (Pflieger, 1975; Rabito and Predorsal tubercles similar in size to those on Heins, 1985). Males and females produce top of head. Along flanks, each scale with 0-5 sounds which females use to attract males (generally 3-4) erect, subcentral organs; close (Delco, 1960); both use these sounds for spe- to nape some begin to appear antrorse. Lateral- cies recognition. line scales sometimes with single, centrally Hybrid swarms have been commonly re- located, erect organ. Caudal-peduncle scales ported between this species and C. lutrensis, with 4-7 small and erect subcentral and \-A even though the two species apparently have central tubercles per scale, best developed the ability to discriminate each other through ventrally. species specific sounds (Delco, 1960). Hubbs Dorsal and anal fins tuberculate on all rays etal. (1953) and Hubbs and Strawn (1956) dis- 80 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY cussed introgressive hybridization with C. process of maxilla short; frontal-sphenotic lutrensis and related the frequency to ecologi- connection placed posteriorly. cal disturbances. Smith (1979) reported hy- Description. One of the largest and more brids of these species from Illinois. elongate members of the genus, reaching about Etymology. The epithet venusta is named 100 mm SL. Body terete and elongate, only after the Roman goddess of love, Venus. The slightly compressed. Head subcorneal, large, name for the eastern subpecies cercostigma is and elongate. Orbit large. Snout relatively derived from the Greek, kertos for tail, and elongate and protruding in nuptial males. stigma for spot, referring to the spot at the base Mouth terminal and oblique; lower jaw con- of caudal fin. The slender subspecies, stigma- tained in upper, both extending to anterior tura is named also for the caudal spot after the margin of orbit. Caudal peduncle slender and Greek stigma and oura for tail. elongate. Dorsal fin greatly expanded in breed- Comments. Until rather recendy, the black- ing males; origin in both sexes slightly poste- tail shiner has had a complex taxonomic his- rior to pelvic fin insertion. Dorsal tory, as evidenced by the synonymy. At least rays 8; anal rays 8-10 (9); pelvic 10 names have been used to describe this rays 8; pectoral rays 14-18 (15 or 16); princi- species, mainly because of poor communica- pal-caudal rays 1 7; dorsal procurrentrays 1 1-14 tions in the 1800s and early 1900s, and poor (12), ventral procurrentrays 10-13 (11). collections from the southeast. Bailey et al. Lateral scale margins higher than wide. (1954) first doubted the validity of the four Lateral-line scale rows 38^3 (39-40, slightly names in use at that time, recognizing only higher in eastern populations); total predorsal one, Cyprinella venusta. As firstreviser, Hubbs circumferential scales generally 26; scales (1954) chose venusta over notata, both de- above lateral line 11-15 (13), below lateral scribed in the same publication. Since that line 11; total caudal-peduncle scales usually time, the taxonomy has remained stable with 14, scales above and below lateral line 7 and 5, the recognition of three subspecies, venusta respectively. Girard, cercostigma Cope, and stigmatura Pharyngeal teeth 1, 4-4, 1, hooked, and (Jordan). Further work in this species seems with narrow grinding surfaces usually not serrated. Total warranted since only a small area of intergra- gill rakers 7-9 (9). Total vertebrae dation was noted by Gibbs (1957c) between 39-41 (40), precaudals 19-22 (20), caudals cercostigma and stigmatura. All three forms 19-21 (20). Dorsal insertion above vertebra 13 may eventually warrant species status. or 14 (13). Lateral line complete. Supratemporal canal 2 side. Cyprinella galactura (Cope) broadly interrupted, pores per Supraor- bital canal complete, pores 8. Infraorbital canal Whitetail Shiner complete, pores 10 or 11. Plate in Preoperculomandi- bular canal complete, total pores 11-13 (12), Hypsilepis galacturus Cope, 1868:160. mandibular pores 4. [Orig. descr.; Type locality: Holston River Intestine with simple S-shaped loop, Type system, Virginia (following Gibbs, 1961:339 I. Peritoneum silvery with several melano- and Gilbert, 1978a)] phores. — Diagnosis. Anal rays9; snout pointed; large Coloration in life. Females and nonbreed- depigmented spots on upper and lower caudal ing males olive to silver-blue dorsally and peduncle at base of caudal fin; snout of breed- laterally to lateral stripe. Lateral stripe silver- ing males red; hiatus in tubercles between blue and best developed posterior to dorsal snout and dorsum of head absent; mandibular origin, continued forward to opercle by broad, tubercles scattered; operculum very broad; diffuse, dusky band delimited dorsally by hypohyal foramen very small; premaxillary narrow dark line. Lateral band darker and NORTH AMERICAN MINNOW GENUS CYPRINELLA 81 broader on hypural plate and basal caudal fin Fin tuberculation as in C. camura, except rays, producing oval caudal spot. Caudal first ray of pectoral fin with 2 organs per peduncle, procurrent rays, and basals of up- segment. permost and lowermost principal caudal rays Distribution. Endemic to the Interior and unpigmented above and below caudal spot, Eastern Highlands of eastern North America. producing two distinct white areas on tail. Western populations in the White and St. Dorsum of head darkly pigmented; melano- Francis rivers of the Ozark Plateaus. Eastern phores, extending to lacrymal, lips, dorsal populations notably common in the Cumber- preopercle, and entire opercle. Gular stripe land, Tennessee, upper Kanawha of the Ohio short and faint. Remainder of head and all of basin, and the Savannah and Santee rivers of belly immaculate. Dorsolateral and lateral the east coast; the latter two systems gained via scales with margins darkly pigmented, pro- stream capture. Prior to glaciation, this species ducing diamond-shaped pattern. Dorsal fin most likely enjoyed a widespread distribution dusky, especially posteriorly on last 2 mem- in these highland areas, as well as in the glaci- branes. Caudal fin dusky, except for unpig- ated Central Lowlands (Mayden, 1985; Wiley mented areas. Leading rays of pectoral fins and Mayden, 1985). outlined with melanophores. Anal and pelvic Ecology. Generally inhabits small to mod- fins clear. erately large, high-gradient, clear streams with Breeding males more brightly colored. Iri- continuous flow. Typically found over clean descent steel-blue dorsally, bluish-silver to gravel or rubble substrates. Common in race- milky-white laterally, and white ventrally. ways or near riffles and frequents deep pools in Snout red. Lateral stripe well developed; ex- association with large boulders and rocky panded between dorsal fin and opercle into banks. large, dark spot. Scapular bar well developed The biology of the whitetail shiner in the and separate from expanded lateral spot by French Broad River, North Carolina, was broad, lightly colored (generally salmon-pink), examined by Outten (1958). Most individuals rectangular region. Pelvic and anal fins white, live a maximum of two years, but some can most intense distally. Pectoral, dorsal, and reach 3 or even 4 years . B y the end of their first caudal fins salmon-pink to red with white summer juveniles average about 34 mm, 55 edges. mm at the end of the second, and 73 mm by the Tuberculation.— Dorsum of head with end of their third. The oldest specimen col- moderately large and antrorse tubercles in lected was an 80 mm, 4-year-old female; the scattered pattern. Preorbital and supraorbital largest was a 101 mm, 2-year-old male. Thus, tubercles antrorse, numerous, and heavily con- as similarly observed by Pflieger (1975), males centrated. Snout organs grade from antrorse have a higher growth rate than females. near nares to erect on snout; continuous with Maturity is attained by most individuals of dorsal head organs. Mandibular tubercles both sexes by 2 years. At this age males de- moderately large and scattered. Small and velop breeding tubercles and bright colora- erect organs present on branchiostegals, inter- tion, and females are greatly distended with opercle, preopercle, gular, and opercular mem- mature oocytes. Males are generally larger brane. Opercle, subopercle, and suborbital than females, 46-95 or 41-78 mm, respec- regions naked. tively. The number of mature oocytes pro- Body tuberculation similar to that of C. duced by 40 females 57-86 mm ranged from camura, except that scales along flanks have, 404 to 1815; oocyte diameter averaged 1.6 in addition to subcentral organs, small tu- mm. Fecundity is likely higher than these ob- bercles located centrally in scattered pattern. served figures, since the whitetail shiner is a Central tubercles on caudal peduncle largest of fractional spawner. body tubercles. In Missouri the spawning season is pro- 82 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY traded, spanning from early June to August Cyprinella pyrrhomelas (Cope) (Pflieger, 1975). In North Carolina spawning Fieryblack Shiner lasts from late May to mid-July (Outten, 1958). Plate in Like other Cyprinella, males guard territories around nesting locations. Nest sites include Photogenis pyrrhomelas Cope, 1870: crevices between rocks or logs, under bark, or 463-464. [Orig. descr.; Type locality: trib. upon the undersurfaces of any object. Males per Catawba River, NC] swim continuously around the nesting loca- Diagnosis. Anal rays 10; mandibular pores tion, defending it from other conspecific males 3; tail of breeding males red and white and and any potential predators of embryos. Once tipped in black; breeding males with red snout a female is attracted to the spawning location and lips; adductor mandibulae Al insertion on by displays, and perhaps sounds, the two maxilla anterior to isthmus; posterior ramus of approach the spawning surface, press their premaxilla straight; ascending wings of paras- vents to the nesting surface, and deposit adhe- phenoid very broad; posterior and anterior sive eggs for up to seven minutes. When nests wings of hyomandibular narrow; lateral com- are located on the undersurface of an object, misure very narrow; anterior notch of vomer spawning behavior is similar, but both sexes very deep; mentomeckelian short and cone- are inverted. A single nest may contain 1 27-3 82 shaped; dermopterotic connected to frontals. embryos arranged in several individual clumps Description. A large and stout member of on the spawning surface. Cyprinella, attaining a maximum of about 90 mm SL. Body robust and deep, only slightly The diet is very diverse, consisting primar- compressed. Head large, triangular, and deep. ily of drifting aquatic or terrestrial insects. Orbit large. Mouth large, terminal, andoblique; This species is a voracious sight feeder, feed- upper jaw containing lower, both reaching to ing equally on drifting midwater materials and behind anterior margin of orbit. Snout blunt, floating items (Outten, 1958). Among aquatic slightly pointed in breeding males and pro- insects, immature Ephemeroptera were most truding. Caudal peduncle thick and relatively commonly eaten, varying seasonally with the short. Dorsal fin expanded in peak males; particular taxon. Immature Diptera, oligochae- origin in both sexes posterior to pelvic fin in- tes, and Hemiptera were very common, but sertion. also varied seasonally. Other items less fre- Dorsal fin rays 8; anal rays 9-11 (10); quently ingested included larvae of Lepidop- pelvic rays 8; pectoral rays 13-17(15); princi- tera, Odonata, Hymenoptera, Nematoda, other pal caudal rays 17; dorsal procurrent rays fishes, and vascular plants. 10-12(12), ventral procurrentrays 10-12(11). Lateral scales with exposed margins higher Etymology. Derived from the Greek ga- than wide. Lateral-line scale rows 34-39 lactos for milk and oura for tail, in reference to (35-36 in Santee and 37 in Peedee); total the whitish region on the caudal peduncle and predorsal circumferential scales generally 26; caudal fin. scales above lateral line 11-15 (13), below lateral line 11; total caudal-peduncle scales Comments. Although this species consists generally 14; scales above and below lateral of two widely disjunct groups of populations, line 7 and 5, respectively. Gibbs (1961) found little differentiation be- Pharyngeal teeth 1, 4-4, 1, hooked, and tween the two. The two groups of populations with smooth or serrated cutting surfaces. Total varied only slightly in number of lateral-line gill rakers 6-7 (7). Total vertebrae 36-38 (37), scales, pectoral rays, and in certain morphom- precaudals 18, caudals 18-20 (19). Dorsal fin etric features, but not enough to warrant taxo- insertion above vertebrae 11-13 (12). nomic recognition. Lateral line complete. Supratemporal canal NORTH AMERICAN MINNOW GENUS CYPRINELLA 83 completeoroccasionally narrowly interrupted, trorse organs between naris and orbit. Preorbi- pores 2 per side. Supraorbital canal complete, tal organs few, moderately large, and antrorse. pores 6-7 (6). Infraorbital canal complete, Snout tubercles equal in size to preorbital pores 9-11 (10). Preoperculomandibular ca- tubercles, scattered, and not separate from nal complete, total pores 9, mandibular pores dorsal head organs by hiatus. Mandibles with 3. single row of moderately large, erect organs. Intestine simple, arranged in single S -shaped Remainder of head devoid of organs. loop, Type I. Peritoneum with silver back- Predorsally, 1-5 rows of very large, an- ground color, overlain with speckling of trorse tubercles; 1 tubercle per scale located No other formed. Midline melanophores. — centrally. organs Coloration in life. Dorsum of females scale row best developed, sometimes extend- and nonbreeding males dark silver-blue, being from dorsal origin to occiput; laterally, coming lighter ventrally. Venter cream to white. rows shorter. Rank scales without tubercles. Lateral stripe weakly developed, but best Caudal-peduncle scales 3 rows above and developed posteriorly where terminates at below lateral line with very large, antrorse or- caudal base with large, distinct oval spot. gans centered on scales. Single row developed Anterior to dorsal origin stripe indistinct and above anal fin, beginning before anal fin and broader, continuing to opercle. Scapular bar terminating about midway along fin base. No well developed. Lateral and dorsal scales edged tubercles on dorsal or ventral midline scale in black, producing crosshatched pattern. rows. No edge or subcentral organs present on Dorsum of head plumbeous, continuing later- any body scales. Like dorsal head tubercles ally to opercle, upper preopercle, lacrymal, and tubercles on predorsal and caudal-peduncle and both lips. Ventrally, head immaculate, scales, tubercles along body begin erect and except for indistinct, short gular stripe. Dorsal transform into antrorse organs in develop- fin dusky, darkest on last 2 or 3 membranes. ment. Caudal dusky, edged posteriorly and on pro- Dorsal fin with single row of tubercles and current rays by heavy concentration of melano- single organ per segment on medial portion of phores. First pectoral ray lightly pigmented. fin from second unbranched ray to second Anal and pelvic fins clear. branched element. Anal fin with tubercles on Breeding males brilliantly colored. Irides- most rays medially, composed of single row cent bluish-silver dorsum and upper lateral per ray and single organ per segment. Pelvic surfaces, darkest dorsally. Snout and lips bright fins with rays 1-5 tuberculate; 1 row medially red. Pectorals, pelvics, and anal fins milky- and 1 tubercle per segment. First ray of pecto- white. Base of anal dusky. Dorsal fin dark ral fin with single row of moderately large, basally and subdistally; membranes heavily retrorse organs, 1 per segment. Posteriorly, pigmented, strongest posteriorly with last 2 rays 2-7 with single row and single organ per membranes jet-black. Anterior membranes red segment. No ventral organs present on paired medially. White band well developed distally. fins. Entire fin with milky-white hue. Caudal fin tri- Distribution. The fieryblack shiner is an colored; narrow black band distally, broad red Atlantic Slope endemic in the Santee and band medially, and white basally. Peedee rivers of North and South Carolina, Tuberculation.— Dorsum of head with very above the Fall Line. Gibbs (1955) and Lee et large, antrorse organs arranged in 2 rows down al. (1980) provide distribution maps of this midline, 1 row per side behind each orbit; de- species. velopment from erect to antrorse. Supraorbital Ecology. Virtually nothing is known of the tubercles also large, but not as large as dorsal life history of this species. It occurs primarily head organs, ranging in number from 3-5; in small to moderate sized streams with clear, connected to preorbitals by single row of an- cool water. It may be found in quiet pools, but 84 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

also frequents deep riffles and raceways. with narrow serrated surfaces. Total gill rakers Etymology. The epithet pyrrhomelas is 5-6 (5). Total vertebrae 38^0 (39), precau- from Greek pyrrhos, meaning flame-colored, dals 18 or 19, caudals 20-22 (20-21). Dorsal and melas, for black, describing the red-black fin insertion above vertebrae 12-14 (13). coloration of caudal fins of breeding males. Lateral fine complete. Supratemporal canal Comments. Based primarily on differences complete, total pores 5. Supraorbital canal in meristic and morphometric characters, complete, pores 8. Infraorbital canal com- populations from the Peedee River drainage plete, pores 10. Preoperculomandibular canal were considered a separate, but undescribed complete, total pores 9, mandibular pores 4. subspecies by Gibbs (1955). Intestine short, forming single S-shaped loop, Type I. Peritoneum silver, overlain by of Cyprinella xaenura (Jordan) speckling melanophores. Coloration in — Dorsum of females Altamaha Shiner life. and nonbreeding males bluish-silver to light-

Minnilusxaenurus Jordan, 1877a: 79. [Orig. tan down to lateral stripe. Below stripe silver descr.; Type locality: S Fk. ofOcmulgee River, to white. Belly immaculate. Lateral stripe at Flat Shoals, Dekalb Co., GA] poorly developed, strongest posterior to dorsal Diagnosis. Anal rays 10; caudal-peduncle fin, terminating posteriorly in large, oval cau- scales below lateral line 7; lacrymal long; dal spot. Anteriorly, lateral stripe very diffuse, anterolateral margin of lateral ethmoid con- but broader and continuing forward to opercle. cave; mesethmoid moderately long; frontal- Dorsal and dorsolateral scales to 1-2 rows sphenotic connection placed anteriorly; below lateral line outlined with narrow, black symplectic narrow; second infraorbital bone borders, presenting diamond-shaped pattern. short; preethmoid socket of palatine long and Scapular bar narrow and strongly developed. narrow; dorsal margin of anguloarticular hori- Dorsum of head olive to dark bluish-silver, ex- zontal. tending to include lacrymal and dorsal half of Description. A large Cyprinella, reaching ascending arm of preopercle and opercle. Re- a maximum of about 90 mm SL. Body deep, mainder of head immaculate. Gular stripe dark. moderately robust. Head large and triangular. Dorsal and caudal fins dusky. Last two mem- Mouth small, terminal, and oblique; jawsreach- branes of dorsal darkly pigmented. Pectoral, ing to or before anterior margin of orbit; lower pelvic, and anal fins clear. Leading ray of jaw smaller than upper and contained. Snout pectoral fin with some melanophores. Anal fin pointed and protruding slightly, especially in with dusky base. peak males. Orbit small. Caudal peduncle stout Breeding males dark plumbeous dorsally. and relatively short. Dorsal fin slightly ex- Lateral stripe broader and strongly defined; panded in peak males; origin in both sexes dark iridescent and deep metallic blue, except slightly posterior to pelvic fin insertion. for black caudal spot. Belly silver. Dorsal fin

Dorsal s 8 anal 1 1 1 1 ray ; rays or ( 0) ; pelvie darkly pigmented, especially last 2 membranes rays 8; pectoral rays 13-17 (15-16); principal which are jet-black. Medially, dorsal fin burnt- caudal rays 17; dorsal procurrent rays 10-13 orange, best developed anteriorly. Caudal (13), ventral procurrent rays 10-12 (11 or 12). dusky with red-orange medial and distal re- Lateral scales higher than wide. Lateral- gions. Dorsal and ventral procurrent rays and line scale rows 38-40 (38 or 39); total predor- medial caudal fin dark. sal circumferential scales generally 26; scales Tuberculation.— Like C. pyrrhomelas, this above lateral line 11-13, below lateral line species has very large, antrorse tubercles on usually 11; total caudal-peduncle scales 16; dorsum of head in form of 2 rows extending scales above and below lateral line 7. down midline and single or double row paral- Pharyngeal teeth 1, 4-4, 1, hooked, and lel to these, only behind orbits. Snout covered NORTH AMERICAN MINNOW GENUS CYPRINELLA 85 with slightly smaller tubercles, antrorse pos- Etymology. The epithet is Greek, xaino for teriorly and erect anteriorly, and not separate scratch and oura for tail, presumably referring from those on top of head. Above orbits, single to the large tubercles on the upper and lower row of 5-6 moderately large, antrorse organs caudal peduncle. connected to group of tubercles at anterior edge of lacrymal. Only occasionally are tu- Cyprinella caerulea (Jordan) bercles present in preorbital region. Area Blue Shiner immediately anterior to orbit devoid of organs. Plate IV Anterior lacrymal organs equal in size to snout tubercles and antrorse. Mandibular rami with Photogenis caeruleus Jordan, 1877b: single row of erect, moderately large tubercles. 338-339. [Orig. descr.; Type locality: tribs. of Remainder of head without tubercles. Oostanaula River (primarily Rocky Creek), Predorsal region with 1-5 rows of very above Rome, Floyd Co., GA] large, antrorse organs; no other tubercles pres- Diagnosis. Anal rays 8; predorsal circum- ent. Single organs centered on each scale and ferential scales below lateral line 9; caudal- rows graded in size, as in other species with peduncle scales below lateral line with several predorsal rows, with central row longest. centrally located tubercles; caudal-peduncle Caudal -peduncle scales also with large, an- scales above lateral line without central tu- trorse organs in center of each scale; generally bercles; dorsal head tubercles develop from 3 rows above and below lateral line. Single linear to scattered pattern; fourth and fifth row above anal fin absent. This species and C. membranes of dorsal fin uniformly pigmented pyrrhomelas share the unique condition among with melanophores; caudal -peduncle tubercles species with large caudal-peduncle tubercles present on dorsal and ventral scale rows; coro- of having scale rows above and below lateral noid process of dentary vertical; premaxillary line with tubercles. Other taxa only have large process of maxilla short; vomer with long organs on ventral caudal-peduncle scales; neck; posterior subopercle greatly elevated; dorsal scales with tubercles equal in size to ceratobranchial 1 with a straight neck; anterior other body tubercles. These two species also margin of frontals with mesial extension over share the lack of other body scale tubercles. supraethmoid; symplectic narrow; posterior Dorsal fin without tubercles. Anal fin tuber- wing of hyomandibular curled. culate on first 4-5 rays medially, with single Description. A moderately sized species, row and single organ per segment. On pelvic reaching about 70 mm SL. Body nearly terete, fin, first unbranched ray naked, but next 3 rays narrow, elongate, and only slighdy compressed. with single row and 1 per segment medially. Head triangular and relatively small. Snout First ray of pectoral fin with single row and pointed, especially in breeding males where it tubercle per segment. Posteriorly, rays 2-6 protrudes. Orbit moderately large. Mouth ter- armed with single row and single element per minal to subterminal and oblique; lower jaw segment. Ventral tubercles on paired fins few, contained in upper, both extending to anterior erect, and small. margin of orbit and longer than orbit. Caudal Distribution. Endemic to the Altamaha peduncle slender and moderately long. Dorsal River system of the Atlantic Slope, above the fin greatly expanded in nuptial males; origin in Fall Line. Gibbs (1955) and Lee et al. (1980) both sexes above to slightly posterior to pelvic provide distribution maps of this species. fin insertion.

Ecology. Nothing known about life history. Dorsal fin rays 8; anal rays 8-9 (8); pelvic Habitat includes small to moderately sized, rays 8; pectoral rays 13-16 (14); principal clear, and cool streams where individuals are caudal rays 17-18(17); dorsal procurrent rays typically found in pools near undercut banks 11-13 (11 or 12), ventral procurrentrays 10-12 and boulders. (10-11). 86 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Exposed margins of lateral body scales dal diffusely pigmented with melanophores. taller than wide. Lateral-line scale rows 37-39 Tuberculation.— Dorsum of head covered

(38); total body circumferential scales gener- with antrorse, moderately large organs arranged ally 22; scales above lateral line 11-13 (11), in a scattered-clumped pattern, as in C. below lateral line 9; total caudal-peduncle whipplei, C. analostana, and C. chloristia. scales generally 14; scales above lateral line 7, Tubercles extend from occiput to anterior below lateral line 5. border of nares and laterally to, but not includ- Pharyngeal teeth 1, 4-4, 1, hooked, and ing, dorsal margin of operculum. Preorbital with narrow grinding surfaces. Total gill rak- region covered with several moderately large, ers 6. Total vertebrae 38^40 (39), precaudals antrorse organs connected to supraorbital 18-20 (19), caudals 19-21 (20). Dorsal fin in- tubercles by series of 2-4 rows. Supraorbital sertion above vertebrae 12-14 (13). tubercles antrorse and arranged in scattered Exposed margins of lateral body scales pattern. Snout organs separate from dorsum of taller than wide. Lateral line complete. Su- head by wide hiatus. Antrorse mandibular pratemporal canal broadly interrupted, pores 2 organs arranged in 2 rows. All other regions of per side. Supraorbital canal complete, pores 8. head devoid of tubercles. Infraorbital canal complete, pores 11. Pre- Predorsally, scales generally with 1-3 cen- operculomandibular canal complete, total pores tral and occasionally 1 or 2 subcentral antrorse 11, mandibular pores 4. organs. Central tubercles generally about equal Intestine in single S-shaped loop, Type I. in size to dorsal head organs. Pattern similar to Peritoneum silvery with scattered melano- that of C. galactura, C. analostana, C. phores. chloristia, C. venusta, and C. spiloptera. No — lat- Coloration in life. Nonbreeding males edge organs present. Along flanks, above and females plain. Body steel-blue to dark eral fine, some scales with weak development silver above lateral stripe, white, or silver- of 1-4 subcentral, erect tubercles; many dor- white below. Margins of dorsal and lateral solateral scales on anterior body without tu- scales darkly pigmented, presenting cross- bercles. Below lateral line, posterior to de- hatched pattern. Bluish lateral stripe present, pressed pectoral fins, each scale with 1-8 best developed posteriorly, but extending from subcentral, antrorse organs. Caudal-peduncle opercle to caudal base, where slightly ex- scales with antrorse organs around margin panded into wider and darker oval spot. Lat- subcentrally (generally 1-8) and series of eral stripe bordered dorsally by narrow depig- centrally located, antrorse organs of equal mented line. Dorsum of head, lacrymal, lips, size. Pattern best developed ventrally on 2 and dorsal opercle and preopercle plumbeous. scale rows below lateral line. For 3 scale rows Venter of head immaculate, except for short above lateral line only subcentral, antrorse gular stripe. Dorsal fin of large specimens with organs present. Dorsal and ventral scale rows some melanophores in last two membranes. of caudal peduncle devoid of tubercles. Pelvic and anal fins clear. Anal fin base pig- Dorsal fin with weak tubercle develop- mented, pigment extending to ventral caudal ment; anterior few rays with single row medi- peduncle. Rays of caudal and pectoral fins ally and single tubercle per segment. Anal fin edged with melanophores. with single row along each ray, 1 tubercle per Breeding males as above, except more bril- segment. Anteriorly, anal rays completely liant. Dorsum and dorsolateral body brilliant covered, but posteriorly rows restricted to blue. Lateral stripe bright bluish-green and medial portions of rays. First 3 rays of pelvics continuous from opercle to caudal base. Fins with single row and 1 organ per segment. First bright yellow or orange and edged in milky- ray of pectoral fin with single row of retrorse white pigments. Dorsal fin membranes dark tubercles, 1 tubercle per segment and extend- except for last two, which are jet-black. Cau- ing most of length of ray; posteriorly, rays 2-7 NORTH AMERICAN MINNOW GENUS CYPRINELLA 87 with single file basally; after segmentation 2 deep and triangular in shape; horizontal plate tubercles per segment. After branching, main of urohyal broad; supraethmoid very broad row continues along posterior branch, first and with entire lateral edges; posterior process with 2 organs per segment and distally with 1 of quadrate deep and heavy. tubercle per segment. Anterior branch with 1 Description. A moderately sized Cyp- tubercle per segment. No tubercles observed rinella, reaching about 80 mm SL. Body elon- on ventral surfaces of paired fins. gate to moderately deep, robust, and stout; Distribution. Endemic to the Mobile Bay only slightly compressed. Dorsal profile arched, Basin, above the Fall Line in the Coosa River greatest body depth between dorsal origin and system. Distribution maps of this species are midway between occiput and dorsal fin; ven- provided in Gibbs(1955),Leeetal.(1980)and tral profile more horizontal. Head moderately Pierson and Krotzer (1987). large and triangular. Snout moderately long Ecology. Almost nothing is known of the and pointed, especially in breeding males where biology of this rare species. Jordan and Ever- it protrudes. Orbit large. Mouth large, termi- mann listed it as common in 1896. Since then, nal, and oblique; lowerjaw contained in upper it has apparendy been extirpated from much of and both extending slightly beyond anterior its range (Lee et al., 1980). This species is ap- margin of orbit. Caudal peduncle stout and parently common today only at a few locations relatively short. Dorsal fin not greatly ex- in southeastern Tennessee, northwestern Geor- panded in breeding males; origin in both sexes gia, and eastern Alabama.. The blue shiner is slightly posterior to pelvic fin insertion. disappearing from Alabama. Habitat includes Dorsal fin rays 8; anal rays 8-10 (9); pelvic sand, gravel, or rubble pools in medium to rays 8; pectoral rays 13-16 (14 or 15); princi- small, clear, cool stream (Pierson and Krotzer, pal caudal rays 17; dorsal procurrent rays 1987). 10-12 (11), ventral procurrent rays 9-1 1 (10). Etymology. The name caerulea is Latin for Exposed margins of lateral scales higher blue, referring to the lateral stripe and pre- than wide. Lateral-line scale rows 37^42 sumably the overall coloration of breeding (38^40); total predorsal circumferential scales males. 23-27 (24-26); scales above lateral line 12-16 (13-15), below lateral line 9-11; total caudal- Cyprinella gibbsi (Howell and peduncle scales 14; scales above lateral line 7, below lateral line 5. Williams) Pharyngeal teeth 1, 4-4, 1, hooked, and Tallapoosa Shiner with narrow serrated grinding surfaces. Total Plate IV gill rakers 5-6 (5). Total vertebrae 37^0 (38), Notropis gibbsi Howell and Williams, precaudals 18-20 (19), caudals 18-21 (19). 1971:55-64, figs. l,2a,e,g. [Orig. descr.; Type Dorsal fin insertion above vertebra 13 or 14 locality: Enitachope Creek, trib. Hillabee (13). Creek, 3.2 mi SSW Ashland, along St. Hwy. 9, Lateral line complete. Supratemporal canal Clay Co., AL] complete, pores 5, or broadly interrupted, pores Diagnosis. Anal rays 9; breeding males 2 per side. Supraorbital canal complete, pores with tubercles in rows on branchiostegals, 8. Infraorbital canal complete, pores 11. Pre- subopercle, cheek, and along ventral margins operculomandibular canal complete, total pores of orbit; dorsum of head with tubercles ar- 10 or 11, mandibular pores 4. ranged in a scattered-clumped pattern; predor- Intestine simple S -shaped loop, Type I. sal scales with several central tubercles; as- Peritoneum silvery with speckling of melano- cending process of premaxilla elongate and phores. of — narrow; posterior ramus premaxilla strongly Coloration in life. Females and nonbreed- decurved; pharyngeal process of basioccipital ing males bluish-silver to olive dorsally and 88 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY dorsolaterally, lighter ventrally before bluish- Antrorse organs present on many surfaces of black lateral stripe. Lateral stripe separate from lower head. Preopercles with several organs of darker dorsum by cream to silver narrow band. equal size as dorsal tubercles in scattered pat- Margins of scales not heavily pigmented; dia- tern and connected to suborbitals and preorbi- mond-shaped pattern not well developed lat- tals. Branchiostegals all with single row of erally. Lateral stripe extending from snout to tubercles. Subopercles with few tubercles in caudal base, where expanded into large, oval row along ventral margin. Interopercle with spot. Anteriorly, lateral stripe 2-3 scale rows single row. Gular region with single tubercle at wide and more diffuse, especially ventrally. symphysis in peak males. Opercle and opercu- Posteriorly, stripe narrows to 1 row before lar membrane naked. Mandibles with 2 rows spot. Belly immaculate. Dorsum of head plum- of tubercles. beous to olive, extending to lateral stripe on Predorsally, each scale with 1-3 centrally dorsal opercle, preopercle, lacrymal, and up- located organs and 1-4 antrorse, smaller, per lips. Venter immaculate except for short, subcentral organs surrounding these. Along dark gular bar. flanks, both above and below lateral line, 1-2 Males more brilliantly colored. Dorsum central, erect and 2-6 subcentral, erect organs bluish-black to steel-blue and separate from per scale. Along caudal peduncle, both above bluish lateral stripe by narrow copper stripe and below lateral line, each scale with 1 central (1-2 scale rows wide); narrow stripe extend- and 2-3 subcentral organs. ing from orbit to caudal base. Lateral surfaces All rays of dorsal and anal fins with single with light blue cast. Belly cream. Dorsal fin row of tubercles, and 1 organ per segment red-orange anteriorly with white overcast of medially and distally. Pelvic fins with single entire fin. First few membranes heavily pig- row on first unbranched ray and next 2 branched mented basally, narrowing posteriorly to pro- rays. Leading ray of pectoral fin with single duce a basal wedge; last 4 interradial mem- row and 1 retrorse organ per segment. Posteri- branes black, producing dark posterior spot. orly, rays 2-7 each with single row of retrorse Anterior 2-A membranes of pectoral, pelvic, organs, 1 per segment. No ventral organs and anal fins red-orange, remainder of fins observed on paired fins. milky-white. Leading ray of pectoral fin also Distribution. Endemic to the Tallapoosa outlined with melanophores. Caudal suffused River system, above the Fall Line, of the with milky-white pigment, epecially basally; Mobile Bay Basin. Howell and\Villiams(1971) rays and membranes red-orange, except dis- considered the single specimen from the Coosa tally where greyish-black. River system (Boschung, 1961) to be a case of Tuberc ulation.— Dorsum of head with accidental bait introduction. Howell and Wil- moderately large, antrorse tubercles in pattern liams (1971) and Lee et al. (1980) provided consisting of 2 clumps of scattered organs on distribution maps of this species. either side of midline; tubercles smaller and Ecology. No published information is avail- distributed in regions normally occupied by able on the life history, except for habitat rows of large tubercles in other species. Su- notes. Although very restricted in range, this is praorbital tubercles arranged in scattered pat- the mostabundant Cyprinella in theTallapoosa tern and approximately equal in size as dorsal system. Howell and Williams (1971) found it head organs. Supraorbitals connected to pre- to be most common in medium-sized tributar- orbital tubercles by 2-3 rows of antrorse or- ies (20-40 ft) over sand, silt, and/or gravel gans. Preorbital tubercles moderately large, substrates. Most frequently associated with scattered, and antrorse; slightly connected to swift current in raceways and deep riffles, but suborbital organs. Snout organs equal in size also found in slow to moderate current in pools to other head organs, antrorse posteriorly, and and eddies. erect anteriorly; separate from dorsal organs. Etymology. This species was named after NORTH AMERICAN MINNOW GENUS CYPRINELLA 89

Dr. Robert Henry Gibbs, United States Na- procurrent rays 10-11(11), ventral procurrent tional Museum, in recognition of his outstand- rays 10-11 (10). ing studies on species of the genus Cyprinella. Exposed edges of lateral scales taller than Comments. Prior to 1971 (Howell and wide, especially anterior, near lateral line. Williams) this species was referred to asNotro- Lateral-line scale rows 36-44 (40-4 1 in Coosa, pis trichroistius. 38-39 in Cahaba, 39-40 in Alabama); total predorsal circumferential scales generally Cyprinella trichroistia (Jordan 24-26; scales above lateral line 11-15 (13), scales below lateral line 8-12 total cau- and Gilbert) (11); dal-peduncle scales generally 14; scales above Tricolor Shiner and below lateral line 7 and 5, respectively. Plate IV Pharyngeal teeth 1, 4-4, 1, hooked, and Codoma trichroistia Jordan and Gilbert in with narrow, sometimes serrated cutting sur-

Jordan and Brayton, 1878:50-51. [Orig.descr.; faces. Total gill rakers 6-7 (7). Total vertebrae Type locality: trib. Etowah River, GA (proba- 38^10 (39), precaudals 18-20 (19), caudals bly near Rome, Floyd Co., GA)(after Gilbert, 19-21 (20). Dorsal fin insertion above verte- 1978a)] brae 12-14 (13). Diagnosis. Anal rays 9; lateral stripe silver- Lateral line complete. Supratemporal canal grey; dorsum of breeding males bright orange; broadly interrupted, pores 2 per side. Supraor- fins yellowish orange; scales below lateral line bital canal complete, pores 8-9 (8). Infraorbi- not appearing strongly diamond-shaped; tu- tal canal interrupted behind orbit, pores 10-12 bercle connection between preorbital and su- (12). Preoperculomandibular canal complete, praorbital regions narrow; supraorbital tu- total pores 10-11 (11), mandibular pores I. bercles in a single row; epibranchial 4 with Intestine simple S-shaped loop, Type I. short mesial to dorsal process; posterior ramus Peritoneum silvery with scattered melano- of premaxilla straight; posterior process of phores. — vomer narrow; anterior neck on ceratobran- Coloration in life. Body coloration of chial 2 long and thin; supraethmoid short; nonbreeding males and females like that of C. anterior notch of vomer very shallow; interop- gibbsi (Howell and Williams, 1971). Breeding erculum deep posteriorly; branchiostegal 1 males differ in having a more blue-grey lateral very narrow. stripe and a narrow yellow stripe above. Fins Description. A moderately sized species of typically yellowish-orange instead of red- Cyprinella, attaining a maximum of about 80 orange.

mm SL. Body elongate and slightly com- Tuberculation.—Dorsum ofhead with large, pressed; greatest body depth at and slightly an- antrorse tubercles arranged in 2 rows anterior terior to dorsal origin. Head triangular and to posterior margin of orbit. Posterior to orbits, moderately large. Snout pointed and long, antrorse organs arranged in semiscattered especially in peak males. Mouth large, termi- pattern, no obvious rows formed. Prenuptial nal, and oblique; upper jaw containing lower, males with linear pattern. Above orbits, single both extending posteriorly past anterior mar- row of large, antrorse organs connects with gin of orbit. Orbit large. Caudal peduncle preorbital tubercles by single row. Preorbital robust and relatively short. Dorsal fin not greatly tubercles few, antrorse, moderately large, and expanded in nuptial males; origin in both sexes scattered. Snout tubercles antrorse posteriorly above pelvic fin insertion. grading to erect anteriorly and smaller than Dorsal fin rays 8; anal rays 8-10 (9); pelvic any tubercles on head. Mandibular organs rays 8; pectoral rays 12-16 (generally 14 or 15 antrorse posteriorly, arranged in 2 rows. in Coosa, 13 or 14 in Cahaba, and 15 in Remainder of head devoid of organs. Alabama); principal caudal rays 19; dorsal Predorsal region with 1-3 rows of large, 90 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY antrorse tubercles; each scale with single tu- tribs. Etowah and Oostanaula rivers, near bercle located centrally. Midline row first to Rome, GA (most specimens from Silver Creek, develop and strongly developed anteriorly. trib. Etowah River)] Lateral to midline, 1-2 rows present, but shorter. Diagnosis. Anal rays 8; caudal -peduncle No other organs present on predorsal scales. scales below lateral fine 7; no large centrally One erect, central tubercle per scale along located tubercles on caudal-peduncle scales flanks. No edge or subcentral organs present above and below lateral line; tubercle hiatus on body scales. Caudal-peduncle scales all between snout and dorsum of head absent; with single tubercle per scale; those on scales tubercle connection between supraorbital and below lateral line very large and antrorse. preorbital region composed of two rows; pre- Above lateral line, central tubercles antrorse, orbital tubercles restricted to single row along but only slightly larger than body tubercles. dorsal margin of lacrymal; mandibular tu- No tubercles present on upper or lower midline bercles absent; caudal-peduncle tubercles scale rows. present on ventral and dorsal scale rows; pig- All dorsal and anal fin rays with single row mentation on scales above and below lateral of organs per ray and single tubercle per seg- line in diffuse diamond-shaped pattern; lateral ment. Anterior rays with more tubercles. Pel- stripe not strongly developed; epibranchial 4 vic fin tuberculate medially and distally. short mesial to dorsal process; single ligament Unbranched ray and next 3 branched rays connecting palatine to ecto- and endo- tuberculate with single row of tubercles and 1 pterygoids; anteriorpreethmoid socket of pala- tubercle per segment. First ray of pectoral fin tine elongate and shallow; ascending wings of with single row of retrorse organs, 1 per seg- parasphenoid very broad; lacrymal tilted ven- ment. Posterior rays 2-9 with single row of trally; supraorbitals narrow anteriorly and tubercles each and 1 organ per segment, ex- posteriorly; anterior notch of urohyal shallow; tending almost to edge of fin. No ventral or- horizontal urohyal plate broad and without gans observed on paired fins. posterior notch; vertical plate of urohyal deep; Distribution. Endemic to the Mobile Bay posterior hyomandibular wing curled; mesial Basin. Found in the Cahaba, Coosa, and Ala- neck of epibranchial 2 straight; anterior paras- bama rivers. One population in Black Warrior phenoid short; ceratobranchial 1 with restricted system believed to be the result of stream neck. capture. Description. A moderately large Cyp- Ecology. Very little known for this com- rinella, reaching about 80 mm SL. Body stout, mon species. Inhabits cool, clear, continuous moderately elongate; only slightly compressed. flowing, small to medium-sized streams and is Dorsal profile elevated and arched; venter more associated with deep riffles and raceways. or less horizontal; greatest body depth from Less common in pools. dorsal fin origin to just posterior to occiput. Etymology. The name trichroistia is from Head moderately large, deep, and triangular. Greek treis, meaning three, chros for color, Snout protruding, especially in breeding males. and histion for sail, describing the tricolored Orbit large. Mouth large, horizontal, and infe- dorsal fin (black, red, and white) of breeding rior. Lower jaw smaller than upper and con- males. tained within the latter. Caudal peduncle heavy and moderately long. Dorsal fin expanded in breeding males; origin in both sexes above Cyprinella callistia (Jordan) pelvic fin insertion. Alabama Shiner Dorsal fin rays 8; anal rays 7-9 (8); pelvic Plate IV rays 8; pectoral rays 13-17 (15-16); principal Photogenis callistius Jordan, caudal rays 17; dorsal procurrent rays 11-13 1877b:337-338. [Orig. descr.; Type locality: (11 or 12), ventral procurrent rays 10-11 (11). NORTH AMERICAN MINNOW GENUS CYPR1NELLA 91

Exposed margins ofdorsal and lateral scales and basally. Distal margin of dorsal milky- taller than wide. Lateral-line scale rows 37^4 1 white. Pelvics and anal immaculate, except for (38-39); total predorsal circumferential scales milky-white pigments. Pectorals white; lead- generally 24; scales above lateral line 11-15 ing ray yellowish, outlined with melanphores. (11), below lateral line 11; total caudal- Caudal bright red; white distally and yellowish peduncle scales usually 16; scales above and basally. below lateral line 7. Tuberculation.— Dorsum of head with 2

Pharyngeal teeth 1, 4-4, 1, hooked, and rows of large, antrorse organs down midline with narrow serrated or entire grinding sur- and single postorbital row per side. Preorbital faces. Total gill rakers 8-9 (8). Total vertebrae areas devoid of tubercles, except for single 39or40,precaudalsl8-20(19),caudals20-21 row of moderately large, antrorse organs be- (20); dorsal fin insertion above vertebra 12 or neath nares, connecting tubercles of supraor- 13 (13). bital region and snout. Supraorbital area with Lateral line complete. Supratemporal canal single row of large, antrorse tubercles. Snout broadly interrupted, pores 2 per side. Supraor- tubercles erect and smaller than dorsal head bital canal complete, pores 8 . Infraorbital canal organs by half. Hiatus between snout and dor- complete, pores 10-11 (11). Preoperculo- sal head tubercles essentially nonexistent in mandibular canal complete, total pores 9, peak males. Mandibles without tubercles. mandibular pores 3. Remainder of head devoid of tubercles. Intestine with single S -shaped loop, Type I. Scales of predorsal region each with single Peritoneum silvery with scattered melano- large, centrally located, antrorse organ. Rows phores. — present down midline and on 1 or 2 scale rows Coloration in life. Body bicolored. Dor- laterally, producing up to 5 rows from dorsal sal half of body from lateral line darkly pig- fin to occiput. In peak males, central row best mented; venter white to cream. Dorsum of developed, extending full length of predorsal nonbreeding males and females silver-blue to region. Lateral rows stepped in length, with olive. Lateral stripe diffuse, best developed lateralmost rows shortest. Flank scales with posteriorly where terminates at large, oval 10-15 small, retrorse edge tubercles per scale dark caudal spot. Weakly developed diamond- on all scales except in belly and breast regions. shaped pattern on lateral and dorsolateral body Tubercles on dorsal fin well developed scales. Dorsal scales darkly pigmented and beyond branching, with 1 tubercle per seg- often obscuring pattern; ventral scales just ment. Anal fin tubercles found on all rays below lateral line with wide, anteriorly di- medially and distally; 1 tubercle per segment. rected chevrons. Dorsum of head darkly pig- Pelvic fins with first ray devoid of organs and mented, extending laterally down to middle of rays 2-6 with single row each, best developed opercle, ascending arm of preopercle, lacry- medially and distally and with single organ per mal, and upper lip. Venter immaculate, except segment. First ray of pectoral fin with single for short, dark gular stripe. Fins generally row of retrorse organs, 1 tubercle per segment. clear, except for dusky and reddish caudal, Posteriorly, rays 2-10 with single row basally lightly pigmented first few rays of pectoral, and on branches, each with single retrorse and distal margin of dorsal which may be organ per segment. Ventrally, both pectoral lightly pigmented with light reddish hue. and pelvic fins without organs. Breeding males dark, steel-blue dorsally Distribution. Endemic to the upper Mo- and dorsolateral^, with dark blue-black cau- bile Bay Basin, generally above the Fall Line. dal spot and posterior lateral stripe. Cream to Mettee et al. (1987) reported this species from silver ventrally. Fins with milky-white cast. below the Fall Line in the lower Tombigbee Dorsal fin bright red medially, dorsally and River. At least two disjunct populations are anteriorly; with large black blotch posteriorly known, those in the Alabama River and those 92 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY in the Tombigbee River system. A distribution (37-38 in Tar to Santee, 39 in Savannah); total map is provided by Lee et al. (1980). predorsal circumferential scales generally 26; Ecology. Other than habitat characteristics, scales above lateral line 13-15 (13), below

is of the of this lateral line 1 1 total nothing known biology very usually ; caudal-peduncle common species. Most frequently inhabits scales generally 14; scales above and below, 7 small to moderately large, high-gradient, cool, and 5, respectively. and clear streams. Typically associated with Pharyngeal teeth 1, 4-4, 1, hooked, and raceways and pools over a gravel and rubble with or without serrations on narrow cutting substrate. surfaces. Total gill rakers 7-10 (9). Total ver- Etymology. The name callistia is from the tebrae 36-39 (38), precaudals 17-19 (18), Greek kallos for beautiful and histion, mean- caudals 18-20 (19). Dorsal fin insertion above ing sail, describing the brilliantly colored, large vertebra 11 or 12 (12). dorsal fin of breeding males. Lateral line complete. Supratemporal canal complete with 5 pores or incomplete with pores 2 per side. Supraorbital canal complete, Cyprinella nivea (Cope) pores 9-10 (10). Infraorbital canal complete, Whitefin Shiner pores 1 1-12 ( 1 1). Preoperculomandibular canal Plate IV complete, total pores 9, mandibular pores 3. Hybopsis niveus Cope, 1870:460-461. Intestine simple S -shaped loop, Type I. [Orig. descr.; Type locality: Upper waters of Peritoneum silvery, but heavily coated with Catawba River, NC] melanophores. — Diagnosis. Anal rays 8; prominent black Coloration in life. Dorsum of nonbreed- lateral stripe present; tubercle development on ing males and females light olive-tan to silver- dorsum of head from linear to scattered pat- grey, continuing to belly but becoming much tern; suborbital tubercles present; second lighter. Belly immaculate. Midlateral stripe membrane of dorsal fin uniformly pigmented bluish, separating dark dorsum from light with melanophores; connection of ligament venter. Stripe strongest posteriorly on caudal from epibranchial 1 to 2 lateral to uncinate peduncle where terminates at caudal base in process of epibranchial 2; pharyngobranchial elongate caudal spot. Anterior to dorsal origin, 1 elongate and narrow; sphenotic-frontal con- stripe diffuse, but wider and continuing to nection placed forward. operculum. Scales above lateral line and 1-2 Description. A moderately large Cyp- scale rows below lateral line with heavily rinella, reaching about 70 mm SL. Body elon- outlined margins, creating crosshatched pat- gate, not deep but terete, only slightly com- tern. Scapular bar narrow and not strongly depressed. Dorsal profile more arched than ven- veloped. Dorsum of head like body dorsum, tral. Head short and subcorneal. Snout blunt, only darker; extending laterally to include la- moderately long and protruding. Orbit moder- crymal, upper opercle, and preopercle; lips ately large. Mouth inferior, lower jaw con- colored as snout. Venter of head immaculate, tained in upper. Caudal-peduncle moderately except for well-developed, short gular stripe. long, but robust. Dorsal fin expanded in breed- Dorsal fin dusky, darkest posteriorly. Caudal ing males; origin over pelvic fin insertion in dusky. Anal, pectoral, and pelvics clear. both sexes. Breeding males with all fins milky-white. Dorsal fin rays 8; anal rays 7-9 (8); pelvic Dorsum dark steel-blue; laterally diffuse light rays 8; pectoral rays 13-16 (15). Principal silver-blue. Lateral stripe more intense, con- caudal rays 17, dorsal procurrent rays 11-14 tinuing from opercle to caudal spot. Dorsal fin (12), ventral procurrent rays 10-13 (11). dusky, except for last 2 membranes that are jet- Exposed margins of lateral scales taller black; some peak males have most membranes than wide. Lateral-line scale rows 35—41 heavily pigmented. Caudal rays outlined with NORTH AMERICAN MINNOW GENUS CYPRINELLA 93

Pectoral and of the Plain melanophores, appearing dusky. , anal, Piedmont, parts Coastal Prov- and pelvic fins unpigmented, except for lead- inces from the Neuse to the Savannah rivers. ing ray of the former, which is outlined with Gibbs (1955) and Lee et al. (1980) provide melanophores. distribution maps of this species. The map Tuberculation.— Dorsum of head covered provided by Gibbs lacks populations from the with antrorse, moderately large tubercles ar- Neuse and includes an erronous locality from ranged in 2 rows anterior to postorbital mar- the upper Tar River, not included in Lee et al. gins and in semiscattered pattern posterior to (1980). orbits. Tubercles extend anteriorly to anterior Ecology. Until recently, little known other margin of nares and laterally to dorsal margin than habitat association. Cloutman and Har- of opercles. Occasionally, single organ present rell (1987) studied food habits, age and growth, on dorsal margin of opercle. Snout covered reproduction, and length-weight relationships with tubercles of equal size, antrorse posteri- of the whitefin shiner. The species was consid- orly and grading to erect anteriorly. Preorbital ered to be omnivorous, but most food items area covered with many small antrorse tu- included insects. Males generally attain ma- bercles connected to single supraorbital row turity at 3 years. Females, however, generally by wide band. Mandibular tubercles antrorse mature at 4 years. Spawning presumably oc- and arranged in 2 fairly well-defined rows, curs between June and August. During this sometimes appearing scattered. Remainder of period gonadosomatic index values are great- head devoid of tubercles. Occasionally, single est and females contain ripe ova. Fecundity tubercle present on suborbital region. ranges from 112 to 545 in single females. Predorsal scales each with single large, Although not documented, the whitefin shiner antrorse tubercle. Generally single row occurs is believed to be a fractional and crevice down midline and bordered laterally by 1-2 spawner like other members of the genus. shorter rows. No subcentral or edge organs Habitat includes moderate to large rivers and present. Along flanks, both above and below reservoirs. In rivers, the species is commonly lateral line, each scale with 1-6 erect and small associated with areas ofcurrent and found near subcentral tubercles and \-4 central erect and the sand and gravel substrate. small tubercles, all of equal size. Caudal- Etymology. The name niveus is Latin for peduncle scales directly above and below lat- snow or snowy, referring to the white pigment eral line with small, erect tubercles in center in fins of breeding males. and along subcentral margins. Scale rows above Comments. This species is a composite of these 1 and below with , occasionally 2, larger, 2 distinct forms, which Gibbs (1955) consid- antrorse central tubercles and series of 1-6 ered to be subspecifically distinct. The de- subcentral, erect tubercles. scription of the undescribed form has not been Distally, dorsal and anal fin rays each with published. One form is found from the Neuse tubercles arranged in single row, 1 per seg- to Santee and the other is endemic to the ment. Pelvic fin rays all tuberculate, 1 row per Savannah River system. Both forms may be ray and 1 tubercle per segment. First pectoral separated by lateral-line scale count and some ray with single row of retrorse tubercles ar- morphometric characteristics. ranged 1 per segment and almost to extending Cyprinella leedsi (Fowler) edge of fin. Rays 2-7 tuberculate; anterior rays Bannerfin Shiner with best development, single row and single tubercle per segment on main branch and both Notropis leedsi Fowler, 1942:2-4, Figs. 3, branches after branching. Small, erect organs 4. [Orig. descr.; Type locality: Ohoopee River, present on ventral surfaces of paired fins. ca. 2.5 mi N Oak Park, Emanuel Co., GA] Distribution. This species is an Atlantic Diagnosis. Anal rays 8; peritoneum brown; Slope endemic, found only in the Montane, third membrane of dorsal fin with pigment 94 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY restricted to distal half; pigment on scales (9), mandibular pores 3. above and below lateral line in diffuse pattern; Intestine simple S -shaped loop, Type I. posterior hyoideal foramen of ceratohyal lo- Peritoneum brown. — cated mesially; ascending process of premax- Coloration in life. Dorsum of females illa short and triangular; nasal canal with a and nonbreeding males olive-tan to light steel- single pore; epiphyseal bar located anteriorly; blue, extending to lateral stripe. Ranks, below mesial neck of epibranchial 2 straight; anterior lateral stripe, cream to silver. Stripe steel-blue, neck of parasphenoid short; ligament connect- moderately broad, and extending from caudal ing epibranchials 1 and 2 attached lateral to base, where expanded into oval spot, to oper- uncinate process of epibranchial 2; pharyn- cular membrane; broader and more diffuse an- gobranchial 1 narrow and elongate; sphenotic terior to dorsal fin. Scapular bar diffuse. Dia- and frontal connection anteriorly placed; dor- mond-shaped pigment pattern on lateral scales sal margin of anguloarticular horizontal. not strongly developed. Scales with broad, Description. A moderately sized Cyp- heavily pigmented margins, obscuring cross- rinella, reaching about 80 mm SL. Body elon- hatched pattern. Diamond-shaped pigment gate, moderately deep, and slightly compressed. pattern present on scales of lateral line and 1-2 Dorsal profile more strongly arched than ven- scale rows below. Dorsum of head darkly tral. Head moderately large and subcorneal. pigmented, including snout, lips, and upper Snout long and pointed, protruding over lips. half of preopercle and opercle. Dorsal fin light Orbit moderately large. Mouth inferior and orange; rays outlined with melanophores, and small; lower jaw contained in upper and both with well-developed spot on membrane be- short of anterior margin of orbit. Caudal hind first rudimentary ray. Anal and pelvic fins peduncle relatively long and slender. Dorsal clear, except for dusky base of former. Leading fin greatly expanded in nuptial males; origin in ray of pectoral outlined with melanophores, both sexes over pelvic fin insertion. otherwise clear. Dorsal fin rays 8; anal rays 8-9 (8); pelvic Breeding males with milky-white fins, rays 8; pectoral rays 12-16 (14-15). Principal especially ventrals. Dorsal fin orange and black caudal rays 17, dorsal procurrent rays 12-13 medially anddistally; basally with clear wedge, (12), ventral procurrent rays 10-11 (10). widest anteriorly. Most fin membranes jet- Midlateral scales with exposed surfaces black. Caudal yellowish-orange with dark higher than wide, especially anterior to dorsal leading rays. Lateral stripe dark blue-black. fin. Lateral-line scale rows 35-39 (36-37); Tuberculation.— Dorsum of head with total predorsal circumferential scales gener- large, antrorse tubercles arranged in 2 parallel ally 26-28; scales above lateral line 13-15, rows down midline and single row laterally, below lateral line 11; total caudal-peduncle posterior to each orbit. Dorsal postorbital re- scales generally 14, scales above and below gion of head occasionally with some tubercles lateral line 7 and 5, respectively. in scattered pattern. Preorbital areas with Pharyngeal teeth 4-4, hooked, and with moderately large, antrorse organs connected narrow, sometimes serrated cutting surfaces. to single row of antrorse supraorbital tubercles Gill rakers long and slender, numbering 5. by 1 or 2 rows of antrorse tubercles between Total vertebrae 37-38 (38), precaudals 18-19 nare and orbit. Snout with smaller tubercles (19), caudals 18-20 (19). Dorsal fin insertion grading from antrorse posteriorly to erect above vertebra 11 or 12 (11). anteriorly, and separate from dorsal organs by Lateral line complete. Supratemporal canal wide hiatus. Mandibles with single row of broadly interrupted, pores 2 per side. Supraor- erect organs along outermost margin. Remain- bital canal complete, pores 8. Infraorbital canal der of head devoid of tubercles. generally complete, pores 11. Preoperculo- Dorsalmost 3 or 4 predorsal scales with mandibular canal complete, total pores 8-9 single, large, antrorse tubercles on scale, form- NORTH AMERICAN MTNNOW GENUS CYPRINELLA 95

ing rows; edge tubercle on each scales absent. nesting site in mating with several females. Flank scales anterior to dorsal fin insertion and Clutch sizes ranged from 26 to 228 eggs per above lateral line usually with single, small, nest. The adhesive eggs ranged from 1 .2 to 1 .6 antrorse organ. Below lateral line, scales usu- mm in diameter. A single hybrid of this species ally devoid of tubercles. No edge organs pres- with C. venusta has been collected by C. R. ent on any body scales. Occasionally, lateral- Gilbert from the Alapaha River in northern line scales and one row above and below lat- Florida. eral line with 2 small, antrorse tubercles each Etymology. Named after Arthur N. Leeds arranged vertically in center of scale. Caudal- of Philadelphia who was present at the type peduncle scales each with single, antrorse locality when the type series was collected. tubercle in center. Those developed below lateral line as or larger than predorsal large Cyprinella callisema (Jordan) organs. No tubercles developed on midline Ocmulgee Shiner row dorsally or ventrally. Tubercles formed Plate IV above lateral line smaller than those below, about the size of lateral body organs. Episema callisema Jordan, 1877b:363-364. Dorsal fin tuberculation consisting of few [Orig. descr.; Type locality: S. Fk. Ocmulgee organs on first rudimentary ray and distal River, Rat Rock, DeKalb Co., GA] portion of first branched ray. Rays of anal and Diagnosis. Anal rays 8; pattern of dorsal pelvic fins tuberculate medially, each with head tubercles develops from linear to scat- single row per ray and single tubercle per tered; pharyngeal teeth 4-4; second membrane segment. First ray of pectoral fin with single of dorsal fin with melanophores uniformly file row of moderately large, retrorse organs. distributed; supraorbital bones moderately Rays 2-7 with single row and single organ per long; basibranchial 1 with no median constric- segment, extending almost to edge of fin. No tion; symplectic narrow; posterior wing of

organs on ventral surfaces of paired fins. hyomandibular terminating near ventral tip; Distribution. Cyprinella leedsi is confined margin of anterior wing of hyomandibular to rivers of the southern Atlantic Slope and smooth; anterior neck of parasphenoid short. eastern Gulf Slope; inhabiting most large riv- Description. An elongate, moderately large ers from the Savannah to the Ochlocknee and member of the genus, reaching a maximum of Suwannee rivers of South Carolina, Georgia, about 75 mm SL. Body slender, only slightly and Florida. See Gibbs (1955) and Lee et al. compressed. Dorsal profile much more highly (1980) for distribution maps. arched than ventral. Venter subhorizontal. Head Ecology. Little information is available on moderately large, narrow, and subconical. the life history of this species other than habi- Snout moderately elongate, protruding, and

tat notes and laboratory observations of breed- tapering to blunt tip. Orbit moderately large. ing behavior (Rabito and Heins, 1985). Adults Mouth small, inferior, slightly oblique. Lower of the bannerfin shiner are difficult to capture, jaw shorter than upper and contained, neither usually being associated with large rivers in reaching anterior margin of orbit. Caudal open channels over a sand substrate. Juve- peduncle elongate and relatively slender. Dorsal niles, however, are more easily captured and fin greatly expanded in breeding males; origin are often near shore (C. R. Gilbert, pers. in both sexes above pelvic fin insertion. comm.). Rabito and Heins (1985) observed Dorsal fin rays 8; anal rays 7-9 (8); pelvic spawning of this species and noted the crevice rays 8; pectoral rays 11-18 (14-15); principal spawning behavior typical of other species of caudal rays 17; dorsal procurrent rays 10-11 Cyprinella. Males defend territories around (11), ventral procurrent rays 9-11 (10). crevice substrates where the eggs are depos- Lateral and dorsolateral scales with ex- ited. A single male may use the same crevice posed margins higher than wide, especially 96 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY anteriorly near lateral line. Lateral-line scale lateral stripe iridescent bluish-purple; scale rows 37-40 (38-39); total predorsal circum- edges dark blue, centers purple. Lateral stripe ferential scales generally 26-28; scales above dark blue-black; caudal spot distinct. Fins with lateral line 13-16 (generally 13 or 15), below milky-white pigments. Dorsal and caudal fins line 1 1 total lateral ; caudal-peduncle scales 14; burnt-orange. Dorsal heavily pigmented on 2 scales above and below lateral line 7 and 5, upper /3 with melanophores on all membranes respectively. and with dark spot anteriorly. Basally, each Pharyngeal teeth 4-4, well hooked, and membrane of dorsal fin only lightly pigmented with or without serrated grinding surfaces. or clear, narrowing posteriorly and presenting

Total gill rakers 6-7. Total vertebrae 39-40 a clear, basal wedge. Pectoral and pelvic fins (40), precaudals 18-20 (19-20), caudals 20 or white, the former with a dark leading ray. Anal

2 1 . Dorsal fin insertion above vertebra 1 2 or 1 3 fin white, except for dusky leading rays and (13). basally, where lightly dusted with yellow- Lateral line complete. Supratemporal canal orange. broadly interrupted, pores 2 per side. Supraor- Tuberculation.— Dorsum of head covered bital canal complete, pores 9. Infraorbital canal with moderately large, antrorse organs scat- complete, pores 11. Preoperculomandibular tered over top of head in 2 clusters, 1 on either canal complete, total pores 9, mandibular pores side of midline. Development of antrorse tu- 3. bercles proceeds from erect to antrorse. Early Intestine simple S -shaped loop, Type I. season males or young males with linear tu- Peritoneum silvery with scattered and large bercle pattern over dorsum of head. Preorbital melanophores. — region with small, antrorse tubercles thickly Coloration in life. Females and nonbreed- distributed over lacrymal. These organs are ing males lightly colored. Dorsum olive-tan to connected to scattered supraorbital organs by light silver-blue down to 1-2 scale rows above 3 rows of equal-sized antrorse organs between lateral line, where lighter. Venter immaculate, nare and orbit. Snout tubercles antrorse pos- except for pigmented anal base and lower teriorly, erect anteriorly, and separate from caudal peduncle. Lateral stripe silver-blue and dorsal head organs by large hiatus. Mandibles extending from caudal base, where expanded with single row of small, erect organs on outer into an elongate oval, to opercle; narrow and edge. Remainder of head devoid of tubercles. diffuse anterior to dorsal fin. Margins of dorsal Predorsal scales each with single, moder- and dorsolateral scales down to 2 scale rows ately large, antrorse tubercle form ing 1 -5 rows. below lateral line darkly outlined, presenting a Rows developed on each scale down midline

bar 1 strong diamond-shaped pattern. Scapular and generally , occasionally 2, rows laterally. absent. Dorsum of head, down to upper third of No other tubercles present on predorsal scales. opercle and preopercle, entire lacrymal, and Rank scales with 1-3 sub-central tubercles, lips like dorsum of body but slightly darker; best developed above lateral line, but also ventral and ventrolateral surfaces of head sil- occurring below, posterior to depressed pecto- ver-white. Gular stripe short and dark. Dorsal ral fins. Caudal-peduncle scales below lateral fin lightly pigmented except for dark mem- line each with single, moderately-large an- brane behind first rudimentary ray. Posterior trorse tubercle in center, forming rows. Cau- membranes of dorsal fin dark. Pelvics and anal dal-peduncle scales may also develop 1-3 fins clear. Caudal rays outlined with melano- subcentral, erect tubercles surrounding central phores, appearing dusky to slightly yellow- organ. Above lateral line, tubercle develop- orange; procurrent rays darkest. Pectoral fins ment rarer and consisting only of central or- clear, except for darkly outlined leading ray. gans. Breeding males more colorful. Dorsal and No tubercles found on dorsal fin. Anal fin dorsolateral surfaces down to and including with tubercles medially, arranged in single NORTH AMERICAN MINNOW GENUS CYPRINELLA 97 row on first 4 branched rays. Medially, first 3 Caudal peduncle moderately deep and long. branched rays of pelvic fins with single row Dorsal fin expanded in breeding males; origin and 1 tubercle per segment. First pectoral fin over pelvic fin insertion. ray with single file row of retrorse tubercles. Dorsal fin rays 8; anal rays 7-8 (8); pelvic Rays 2-9 each with single row and tubercle per rays 8; pectoral rays 13-17 (14 or 15); princi- segment, extending to edge of fin. No ventral pal caudal rays 17-18 (17); dorsal procurrent organs observed on paired fins. rays 1 1-12 (12); ventral procurrentrays 10-1 1 Distribution. Cyprinella callisema is en- (11). demic to the Altamaha and Ogeechee rivers of Exposed margins of lateral scales taller the southern Atlantic Slope in Georgia. Gibbs than long. Lateral-line scale rows 37-40 (38 or (1955) and Lee et al. (1980) provide complete 39); total circumferential scales 26; scales distribution maps of this species. above lateral line 13, below lateral line 11. Ecology. No life history data are available Total caudal peduncle scales 14; scales above for this common species. Habitats most fre- lateral line 7, below lateral line 5. quently include moderately large streams with Pharyngeal teeth 1, 4-4, 1, hooked, and continuous current. Typically associated with with poorly developed grinding surfaces. To- open channels over a sand and/or gravel sub- tal gill rakers 5-7 (6). Total vertebrae 38-39 strate, although generally over sand. (39), precaudals 19-20 (20), caudals 18-19 Etymology. Derived from the Greek kalos (19). Dorsal fin insertion over vertebra 12 or for beautiful and sema for sign or sail, in 13(13). reference to the expanded and colorful dorsal Lateral line complete. Supratemporal canal fin of peak breeding males. broadly interrupted, pores 2 per side. Supraor- bital canal complete, pores 8. Infraorbital canal Cyprinella callitaenia (Bailey and complete, pores 10-11 (11). Preoperculo- mandibular canal complete, total pores 9, Gibbs) mandibular pores 3. Bluestripe Shiner Intestine simple S-shape loop, Type I. Peri- Notropis callitaenia Bailey and Gibbs, toneum silvery, overlain by scattered melano- 1956:1-14, fig. 1. [Orig. descr.; Type locality: phores. — Flint River, about 1 mi S Radium Springs Coloration in life. Dorsal and dorso- outlet, 5.5 mi S Albany, Dougherty Co., GA] lateral surfaces of females and nonbreeding Diagnosis. Anal rays 8; prominent steel- males plumbeous to olive, becoming lighter blue lateral stripe with dark basicaudal spot, ventrally to lateral stripe. Lateral stripe metal- not separate from lateral band and slightly lic blue, extending from caudal base to opercle. wider; lower margin of lacrymal outlined by Stripe more diffuse and broader anterior to narrow, dark row of melanophores; single tooth dorsal origin, formed mainly by a narrow, dark in secondary tooth row of at least one pharyn- line extending forward as a continuation of geal arch; parietals invade dermopterouc; dorsal margin of posterior portion of stripe and posterior process of vomer narrow; met- bordered ventral to lateral line by diffuse and apterygoid articulation with interhyal narrow; dusky region. Lateral stripe expanded slightly ectoperygoid short and broad. and darker at caudal base, form ing caudal spot, Description. An elongate, moderately large but not separate from stripe. Venter light cream Cyprinella, reaching up to 70 mm SL. Body or white. Dorsum of head down to lips, lower slender and slightly compressed. Head small. lacrymal, and upper opercle plumbeous; light Orbit moderately large. Mouth small, inferior, ventrally. Lower margin of lacrymal bordered and slightly oblique; upper jaw longer than by dark narrow line of melanophores. Man- lower, both short of anterior margin of orbit. dibles with scattered melanophores laterally. Snout long, blunt, and slightly protruding. Dorsal fin with melanophores lining rays and 98 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY heavy concentration on posterior membranes. cambia River. This record is not considered Other fins generally clear. valid (Gilbert, 1978b, 1980). Distribution maps Breeding males as above but more brightly are presented by Bailey and Gibbs (1955) and colored. Dorsum of body steel-blue, spotted Gilbert (1980). with metallic pink flecks, changing to salmon- Ecology. Some aspects of the biology of pink above steel-blue lateral stripe. Dorsal fin this species are well known (Wallace and iridescent green centrally, blue subdistally, Ramsey, 1981). Habitat includes areas of cur- and milky-white distally over a background of rent in large rivers over a sand and/or gravel heavy melanophores. Caudal lobes, pectorals, substrate. Much of this habitat type is and has and pelvic fins fringed in white. Anal bright been lost through impoundment projects greenish-yellow, tipped in white. (Gilbert, 1978b, 1980). Tuberculation.— Early males with 2 semi- Reproduction in Uchee Creek, Alabama, linear rows of tubercles on top of head; peak probably spans from late April to August males with rows anterior to posterior margin (Wallace and Ramsey, 1981). GSI began to of orbit and semiscattered pattern posteriorly. increase in March, reached a peak in June, and Dorsal head tubercles moderately large and maintained a high level until August. Eggs antrorse. Preorbital region with several scat- numbered 88-230 per female. Total eggs pro- tered, antrorse, and moderately large organs duced by a female may be much greater since connected to single row of supraorbital organs C. callitaenia is probably a fractional spawner, by 1-2 rows of antrorse organs. Snout tu- like other members of the genus. bercles antrorse posteriorly and erect anteri- Spawning occurs in June and is similar to orly, separate from dorsal head organs by wide other Cyprinella. Males maintain territories hiatus. Mandibles with 2 rows (occasionally 1) near crevices and swim back and forth in the of small, erect organs. Remainder of head de- current with fins erect to attract females. If void of tubercles. unsuccessful, some males may swim as much Predorsal region with 1-5 rows of large, as 3 m or more from the crevice to obtain a antrorse tubercles. Rows formed from 1 tu- female and may return with as many as three. bercle per scale. Median predorsal scale row After the male displays in the typical Cyp- always first to develop organs. No edge tu- rinella "figure-8" pattern around the crevice, bercles present. Lateral body scales with single, both sexes press their vents in the crevice and small antrorse organ in center of scale, as in C. vibrate, spraying eggs and milt into the cavity. leedsi. Caudal peduncle tuberculation like that Afterwards the female moves into fast current species. and the male continues to seek additional mates. Dorsal fin devoid of tubercles. Anal fin with Nesting areas are used by several males; eggs organs present medially on first rudimentary found in a cavity may number several hundred ray and first 4 branched rays. Pelvic fin rays and are arranged in several layers at several 1-5 with tubercles on medial and distal por- stages of development. Fertilized eggs range tions. First pectoral fin ray with single row of in size from 1 .2 to 1 .7 mm and are located 3 or retrorse organs, 1 tubercle per segment. Rays more cm from the outer edge of the crevice. 2-6 with single file row, developing 1 tubercle Cases of hybridization with the bluestripe per segment. Each ray with organs extending shiner are few and limited to other species of close to fin edge. No ventral paired fin tu- Cyprinella. Bailey and Gibbs (1956) reported bercles noted. a case of hybridization with C. venusta, and Distribution. The bluestripe shiner is en- Wallace and Ramsey (1982) noted a hybrid demic to the Apalachicola River System, in- with the red shiner. All three species are now cluding the Apalachicola, Chattahoochee, and known to be crevice spawners, and C. venusta Hint rivers. Bailey and Gibbs (1956) provi- and C. lutrensis are known to have distinctive sionally listed a second record from the Es- sounds for species and sex recognition (Delco, NORTH AMERICAN MINNOW GENUS CYPRINELLA 99

Fig. 41. Dorsal fin pigmentation patterns in male Cyprinella and close relatives. A) Lythrurus umbratilis, 52 mm, KU 9547. B) Luxilus cornutus, 96mm,KU 7296. C) Cyprinella ornata, 49 mm, ASU 9629. D) C.proserpina, 56mm,TNHC 9771. E) C. spiloptera, 59 mm, KU 14561. F) C. whipplei, 80 mm, KU 11357. G) C. camura, 84 mm, KU 10734. H) C. analostana, 68 mm, TU 25867. 100 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

G H

C. Fig. 42. Dorsal fin pigmentation patterns in male Cyprinella. A) C. galactura, 98 mm, KU 19732. B) venusta, 94 mm, KU 19698. C) C. pyrrhomelas, 83 mm, UAIC 2550.05. D) C. trichroistia,l\ mm, KU 5292. E) C. callistia, 84 mm, KU 19697. F) C. nivea, 11 mm, TU 38616. G) C. leedsi, 74 mm, UAIC 3127.01. H) C. callisema, 58 mm, KU 19653. NORTH AMERICAN MINNOW GENUS CYPRINELLA 101

P^ifBi

Fig. 43. Tuberculatum patterns in male Cyprinella. A-D) C. garmani (50 mm, KU 20164): lateral view of head, dorsal view of head, lower caudal-peduncle and above anal fin, and ventral caudal-peduncle. E-H) C.formosa (42 mm, KU 8399): nape scales, lateral body scales, lower caudal-peduncle and scales above anal fin, and ventral caudal-peduncle scales. 102 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

J NORTH AMERICAN MINNOW GENUS CYPRINELLA 103 104 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 46. Tuberculation patterns in male Cyprinella. A) caudal-peduncle scales of C. xaenura (100 mm, TU 29458). B- C. F) pyrrhomelas (84-85 mm, UAIC 2550.05): lateral view of head, caudal-peduncle scales, dorsal view of head, nape and lower scales. scales, caudal-peduncle G-H) C. caerulea (60 mm, UAIC 2580. 1 3): dorsal view of head and nape scales. NORTH AMERICAN MINNOW GENUS CYPRINELLA 105

Fig. 47. Tuberculation patterns in male Cyprinella. A-B) C. gibbsi (82 mm, TU 40670): lateral view of head and dorsal view of head. C-F) C. callistia (84 mm, KU 1 9697): lateral view of head, dorsal view of head, nape scales, and left pectoral fin. G-H ) C. callisema (62 mm, KU 19653): lateral view of head and dorsal view of head. 106 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 48. Tuberculation patterns in male Cyprinella. A-D) C. nivea (78-82 mm, TU 38616): lateral view of head, dorsal of view head, nape scales, and ventral caudal-peduncle scales. E-H) C. leedsi (74.5 mm, UAIC 3127.01): lateral view of head, dorsal view of head, nape scales, and lateral body scales. NORTH AMERICAN MINNOW GENUS CYPRINELLA 107

1960). Although Wallace and Ramsey (1981) More likely, the crevice habitat was an attrac- observed a male spottail shiner attempting to tive spawning site for C. venusta. swim into the nesting crevice of the bluestripe Etymology. The name callitaenia is de- shiner, it is unlikely in this clear stream that hy- rived from the Latin calli for beautiful and bridization would have resulted. Most cases of taenia for band or ribbon, in reference to the hybridization can be correlated with ecologi- lateral blue stripe. cal disturbances (Hubbs and Strawn, 1956). OSTEOLOGY

This section contains an osteological de- group taxa (and variation) and the primitive scription of species of Cyprinella, as well as and derived morphologies within Cyprinella comparisons to other North American cyp- are considered. In most cases, the nearest out- rinids. Below, the skeleton of these fishes is groups to Cyprinella were used to determine divided into 15 separate sections. Under each character polarities and are discussed most section, included bones are discussed in detail thoroughly. The derived morphologies are used with respect to their developmental origins and discussed later under systematic species and systematic variation of several potential relationships. morphological variables. Characteristics examined for variation included the shape and Ethmoid Region relative size of a bone and its articulation with The ethmoid region (Figs. 49-58) of the adjacent elements. Variations in bones noted skull is complex and composed of elements of only in individual specimens, and hence of dermal, endochondral, and perichondral ori- limited use systematically, are not discussed. gin. Included are the mesethmoid, lateral Only those skeletal variations noted to be ethmoids, preethmoid cornua, supraethmoid, consistent for a species and hypothesized to be vomer, kinethmoid, and nasals. The last four potential characters are included. In this study, elements are of dermal origin and only the many morphological variations were noted nasals are paired. The first three elements are from the skeletons of these fishes, especially cartilage bones. The mesethmoid is single and from certain anatomical regions, but these perichondral, the preethmoids are paired and variations were not consistent within a species endochondral, and the paired lateral ethmoids and were thus not considered further. Some are dermal, endochondral, and perichondral. variations hypothesized below to represent Most of these elements form surfaces sur- systematic characters are subde differences in rounding the nasal cavities. The supraethmoid the shape or relative size of the bone or perhaps and nasals are dorsal, lateral ethmoids are the type or degree of articulation with adjacent posterior and ventral, the mesethmoid is ante- bones. Although these characters may appear rior, dorsal and ventral, and the vomer is ven- to be of minor significance, they were in al- tral. The preethmoids and kinethmoid are not most all cases, verified with one or more addi- in the nasal cavity and are anterior. The ki- tional specimens of a taxon to guard against nethmoid articulates via ligaments to the ante- individual variation. Numerous illustrations rior supraethmoid and mesethmoid and the of variations discussed are presented to aid the preethmoids lie between the anterolateral reader in descriptions and interpretations. mesethmoid and vomer. Abbreviations used in all illustrations and text Penetrating much of the ventral ethmoid re- are listed and explained in the Methods section gion is the cartilaginous planum ethmoidale. of this paper. This cartilage block extends between the Following the discussion of osteological mesethmoid and vomer anteriorly and is ex- variation ofa particular bone among species of panded posteriorly to the orbitosphenoid and Cyprinella, the variation is compared to out- laterally into the lateral ethmoids. Dorsally, it 108 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

vo The lateral surfaces thus form the respective faces of the nasal cavity. The ventral cavity is invaded by the planum ethmoidale like the lateral ethmoids, while the dorsal is generally hollow although some endochondral ossification may occur. The dorsal surface is covered by the dorsally convex supraethmoid. Anteriorly the mesethmoid produces bony anterolateral projections for ligament connections to the kinethmoid. Anteroventrally it serves as the dorsal receptacle for the preethmoids. The length of the mesethmoid is variably developed in Cyprinella. Cyprinella xaenura has a short mesethmoid block (Fig. 57E), while in callitaenia, callisema, leedsi, and nivea it is

elongate (Fig. 57F). All other species have similarly sized mesethmoids (Fig. 57D), as in outgroups exam ined. In xaenura, pyrrhomelas, trichroistia, gibbsi, and caerulea the olfactory EPO foramen is large and bounded entirely by a EOC SOC laterally directed shelf produced by the LOF mesethmoid and lateral ethmoids (Fig. 57E). In all other taxa no shelf is formed and bom 49. Dorsal view of cranium of Luxilus Fig. coccogenis bones slope gradually into the foramen. (72 mm, INHS 79254). Stippled regions represent The shelf around the olfactory foramen is cartilage. Lateral occipital foramina identified by lined considered derived in this since no other pattern. Horizontal bar equals 1 mm. genus species examined develops this structure. The invades much of the ventral half of the short mesethmoid of xaenura and long mesethmoid. In adults, the planum ethmoidale mesethmoid of species of the nivea group are becomes ossified laterally in the lateral independently derived relative to other spe- ethmoids, dorsally in the mesethmoid, and cies of Cyprinella and the outgroups Luxilus ventrally and anteriorly between the and Lythrurus, all of which have similarly mesethmoid, preethmoids, and vomer. sized elements. Mesethmoid (ME, Fig. 57). Resting on the Lateral Ethmoid (LE,Figs.49-58).These cartilaginous planum ethmoidale, this semi- paired bones are complex structurally and circular bone is the largest anterior element of developmentally, having dermal, endochon- the ethmoid region. Medially it is constricted, dral, and perichondral portions. They articu- forming a waist near the olfactory foramen. late with the mesethmoid anteriorly, su- Dorsally and anteriorly it is roughly convex. praethmoid, frontals, and orbitosphenoid dor- Posteriorly, it is laterally compressed and ori- sally, and parasphenoid and vomer ventrally ented vertically, forming the nasal septum. (Fig. 57). They form the posterior face of the The posterior margin is roughly vertical where nasal cavity anteriorly and the anterior face of it articulates with the lateral ethmoids. To- the orbits, anterior neurocranium, and myod- gether these bones contain the olfactory fora- ome posteriorly. Viewed ventrally, each bone men. Anteriorly, dorsally, and ventrally the is roughly triangular, broadest mesially and mesethmoid is expanded from the median narrowing laterally, producing a long lateral waist, producing dorsal and ventral cavities. wing. The ventral surface is somewhat con- NORTH AMERICAN MINNOW GENUS CYPRINELLA 109

50. Dorsal view of cranium of and close relatives. 41 Fig. Cyprinella A) Rhinichthysfalcatus , mm, KU 18912. B)Notropis 55 18935. 6244. 17776. alherinoides, mm, KU C) Lythrwus fumeus , 55 mm, KU D) Cyprinella spiloptera, 65 mm, KU Horizontal bar equals 1 mm. 110 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

8405. Fig. 51. Dorsal view of cranium in Cyprinella. A) C. lutrensis, 55 mm, KU 19431. B) C. ornata, 56mm, KU C) C. proserpina, 58 mm, TNHC 3262. D) C. rutila, 65 mm, ASU 5982. See Figure 49 for identification of elements. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 111

Fig. 52. Dorsal view of cranium in Cyprinella. A) C. garmani, 46 mm, KU 5416. B) C.formosa, 47 mm, KU 8399. C) C. lepida, 50 mm, KU 55189. D) C. panarcys, 45 mm, UMMZ 208212. See Figure 49 for identification of elements. Horizontal bar equals 1 mm. 112 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

C. 55 KU 14211. Fig. 53. Dorsal view of cranium in Cyprinella. A) C. camura, 60 mm, KU 15792. B) whipplei, mm, C) C. analostana, 60 mm, INHS 77855. D) C. chloristia, 55 mm, KU 8882. See Figure 49 for identification of elements. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 113

Fig. 54. Dorsal view of cranium in Cyprinella. A) C. galactura, 67 mm, KU 12029. B) C. venusta, 85 mm, KU 8810. C) C. pyrrhomelas, 79 mm, INHS 76839. D) C. xaenura, 74 mm, KU 1 8994. See Figure 49 for identification of elements. Horizontal bar equals 1 mm. 114 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

59 18892. Fig. 55. Dorsal view of cranium in Cyprinella. A) C. trichroistia, 60 mm, KU 18853. B) C. gibbsi, mm, KU of elements. C) C. caerulea, 60 mm, KU 18978. D) C. callistia, 74 mm, KU 18842. See Figure 49 for identification Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 115

Fig. 56. Dorsal view of cranium in Cyprinella. A) C. nivea, 60 mm, KU 18987. B) C. leedsi, 51 mm, KU 18985. C) C. callitaenia, 59 mm,TU 92110. D)C. callisema, 52 mm, KU 8842. See Figure 49 for identification of elements. Horizontal bar equals 1 mm. 116 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY cave, depressed or slightly dished anteriorly In the whipplei clade most of the lateral and elevated along the posterolateral margin, ethmoid becomes ossified at an early age. forming a ridge. This elevated ridge supports Cartilage is present only as a thin line or spot the palatine-endopterygoid articulation. posteromesially or laterally, isolated from the Medially these bones are constricted, pri- mesial cartilage separating the two lateral marily with the development of the olfactory ethmoids (Fig. 58F-L). foramen. Each produces a posteromesially Some Cyprinella develop a strongly trian- directed shelf that forms the floor of the ante- gular ventral surface to the lateral ethmoid rior neurocranium (Fig. 57). The mesially with oblique anterior and relatively straight directed shelves, together with a medial notch posteroventral edges. In others the anterior in each bone below the shelf, form the anterior margin is concave when viewed from below, myodome for the insertion of the inferior and such that a slight anteromesial process is de- superior oblique eye muscles. Posteriorly, veloped and the bone is roughly L-shaped. All below the anterior myodome, the lateral species of the lutrensis group, except proser- ethmoids, together with the orbitosphenoid, pina and panarcys, and spiloptera, camura, form the anterior wall of the orbit, while an- whipplei, analostana, and chloristia have the teriorly they form the posterior face of the concave anterior margin. Cyprinella panarcys nasal cavity. Anteriorly they articulate with the and proserpina have straight margins to the mesethmoid and together border the olfactory lateral ethmoids, unlike any other species of foramen. the genus. Cyprinella galactura, venusta, and Dorsal to the medially constricted region xaenura have intermediate triangular lateral the lateral ethmoids consist of two parallel and ethmoids ventrally, but with a slightly curved mesially curving walls that form the sides of anterior margin (Fig. 581). All other species the taenia marginalis anteriorus. These dorsal have the triangular morphology (Fig. 58J-L). walls are broadly separated from their counter- Further influencing the shape of this bone is parts posteriorly and only narrowly separated the shape of the ventromesial edge where the anteriorly. Thus, the mesial concave walls two meet to form the floor and roof of the form the anterior and ventral surfaces of the anterior neurocranium and myodome, respec- anterior neurocranium and the lateral walls tively. It may be either convex or squared. All form the dorsoposterior face of the nasal cav- species of the lutrensis clade, spiloptera, chloristia ity. The dorsoposterior face forms the anterior camura, whipplei, analostana, and wall of the orbit (Fig. 57). have a convex margin ventrally, like other Morphology of the lateral ethmoid varies cyprinids examined. Other Cyprinella have among species of Cyprinella with respect to the blunt and squared margin. the shape of the ventral and mesial surfaces, In all species except caerulea, gibbsi, the complexity of the olfactory foramen, and trichroistia, xaenura, and pyrrhomelas, the ossification of the investing cartilaginous surrounding lateral ethmoid and mesethmoid ventral planum ethmoidale. In species of the slope gradually to the olfactory foramen, in a lutrensis clade and outgroups, the ventral sur- manner similar to other cyprinids examined. face of the lateral ethmoids of adults is not In the above listed species a laterally project- completely ossified. Only the anterior and ing shelf is formed by both the mesethmoid anterolateral surfaces become ossified and a and anterior lateral ethmoids around the fora-

large mesial portion extending laterally into men (Fig. 57). the bone remains cartilage (Fig. 58A-E). When As discussed under the mesethmoid, the maximum ossification occurs a large cartilage shelf around the olfactory foramen represents

pad generally remains posteromedian y. This a derived condition. Based on outgroup com- pad may be separate from the mesial planum parison the triangular lateral ethmoid with a ethmoidale. blunt mesial edge is derived in this group, as NORTH AMERICAN MINNOW GENUS CYPRINELLA 117

Fig. 57. Ethmoid region in Cyprinella and outgroup. A-C) Lateral, ventral, and dorsal view of ethmoid region of Cyprinella lutrensis (55 mm, KU 19431). In A the nasal cavity is illustrated with the centrally located olfactory foramen. In B the cartilage-filled mesethmoid and lateral ethmoids are illustrated beneath the vomer and parasphenoid. In C the anterior neurocranium formed by the anterior orbitosphenoid and posterior lateral ethmoids, and the internasal septum are illustrated beneath the frontals and supraethmoid. D) Lateral view of ethmoid region of Luxilus cardinalis (65 mm, KU 15281) illustrating primitive olfactory foramen and mesethmoid size. E) Lateral view of ethmoid region in Cyprinella xaenura, illustrating shelf formed around olfactory foramen and short mesethmoid. F) Lateral view of ethmoid region of C. callisema (52 mm, KU 8842) illustrating elongate mesethmoid and primitive olfactory foramen. Horizontal bar equals

1 mm.

well as the nearly completely ossified ventral neck separating the preethmoids from the an- surface. All other cyprinids examined, except terior extent of the lateral ethmoids. Postero-

for some Pteronotropis, have a basically L- laterally, where the medial neck ends the vomer shaped lateral ethmoid with a convex mesial is restricted to form a long, narrow posteriorly margin and the planum ethmoidale extends directed process. This process lies in a shallow laterally into this bone and generally remains trough against the ventral surfaces of the par- in adults. The conical shape of the lateral asphenoid and may extend to the posterior ethmoids in panarcys andproserpina is unique margin of the lateral ethmoids. to these species and is derived. In Cyprinella, interspecific variation in Vomer (VO, Figs. 49, 57, 58). This single, vomerine shape is expressed in the length of medial dermal bone lies horizontally beneath the posterior process, depth and width of the the ethmoid complex and the anterior end of anterior notch separating the preethmoid proc- the parasphenoid, and is visible only when esses, and length of the medial neck (Fig. 58). viewed from below or laterally. It is broadest Some species have little or no medial neck and thickest anteriorly where forked into two developed between the preethmoids and lat- anterolateral processess, each forming a shal- eral ethmoids. These include spiloptera, low bowl where the mesial preethmoid is lutrensis, garmani, and ornata. Others have a formed. Separating these two processes is a medial neck developed that may be either variably developed notch. Medially, the vo- short or long. Species with a long neck include mer is flat and thin and forms a short, broad analostana, chloristia, caerulea, nivea, leedsi, 118 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

53 Fig. 58. Ventral view of ethmoid region in some species of Cyprinella and outgroups. A)Notropis atherinoides, mm, KU 18935. B) Luxilus coccogenis, 72 mm, INHS 79254. C) Cyprinella garmani, 50 mm, KU 5416. D) C. panarcys, 44 mm, UMMZ 208212. E) C. rutila, 57 mm, ASU 5982. F) C. spiloptera, 56 mm, KU 17776. G) C. whipplei, 55 mm, 60 18978. KU 14211. H) C. camura, 45 mm, KU 15792. I) C. venusta, 58 mm, KU 8810. J) C. caerulea, mm, KU K) C. leedsi, 53 mm, KU 18985. L) C. callitaenia, 57 mm, TU 92770. Stippled pattern represents cartilage. In A the posterior distribution of the planum ethmoidale is illustrated. In all others only that portion visible ventrally is illustrated in the lateral ethmoids and between the vomer and lateral ethmoids. In C-L the ossified preethmoids are illustrated with the cartilaginous preethmoid comua. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 119

callitaenia, and callisema. The anterior notch present. As adults, these ossifications fuse, is broad and moderately shallow in all species, forming a large single endochondral ball sur- except nivea, leedsi, callitaenia, callisema, rounded laterally by cartilage. No significant pyrrhomelas, lutrensis, xanthicara, panarcys, interspecific variation occurs in Cyprinella. and proserpina, in which it is relatively deep. The occurrence of these elements in the genus Species of the nivea group also differ in having might be hypothesized as being diagnostic the preethmoid processes closer together, since the closest outgroups (Lythrurus and producing a more narrow notch. No notch is Luxilus) apparently do not develop these struc- developed in caerulea and gibbsi; in xaenura tures. and trichroistia the notch is very shallow. Supraethmoid (SE, Figs. 49-57). Anterior The posteriorly directed process of the to and generally articulating with the paired vomer is variable in length and width. Gener- frontals is the dorsoventrally compressed

ally, the posteriormost extent of the vomer is at supraethmoid. This roughly square bone roofs the posterior edge of the lateral ethmoids, as in the nasal capsules and is fused with the dorsal most outgroups examined. In caerulea, gibbsi, mesethmoid. Relative to species of the more trichroistia, pyrrhomelas, and xaenura the distant and primitive clades of Rhinichthys process terminates well before the edge of the and Nocomis, the supraethmoid of Luxilus, lateral ethmoids, and in rutila, xanthicara, Lythrurus, Notropis (s.s.),andmostCyprinella panarcys, and proserpina it extends beyond is a shorter structure. Like Rhinichthys and the lateral ethmoids. Of these latter four spe- Nocomis, however, most species of Lythrurus cies it is longest in rutila and proserpina. The and Notropis (s.s.), have a supraethmoid that posterior process is narrow in spiloptera, has a "butterfly" shape, similar to that de- galactura, venusta, analostana, trichroistia, scribed by Harrington (1955) for N. bifrenatus

callitaenia, rutila, and xanthicara and broad, and Coburn (1982) for Notropis (s.s.). In these as in most outgroups, in all other species of the taxa the anterior- and posterior-most ends are genus. broader than the median waist. Except for the Development of a medial neck on the vomer nivea species group of Cyprinella, the genera is derived in Cyprinella. Most members of the Luxilus and Cyprinella share a broader and genera Luxilus and Lythrums lack any neck more square-shaped supraethmoid, compared development. Outside this clade several other to other species examined. In these two clades taxa are known to develop an elongate to short the median constriction is usually only slight neck. Having a moderately shallow anterior and the anterior margin of the bone is generally notch in the vomer is primitive in this genus. only slighUy and smoothly indented. The deep, shallow, and no notch conditions are Within Cyprinella, several different derived derived, as well as the narrow notch in the conditions of the supraethmoid are found. nivea group. A wide posterior vomerine proc- Members of the lutrensis clade have the same ess extending to the posterior margin of the broad and short supraethmoid found in Lux- lateral ethmoids is primitive in Cyprinella. ilus,except(orgarmani,formosa,bocagrande, This condition is common to the genera Lux- lepida, proserpina, panarcys, rutila, and xan- ilus and Lythrurus. The short and elongate thicara. These species have a narrower condi- conditions are derived for Cyprinella, as is the tion (but broader than the nivea group), and narrow posterior process. have a slight constriction of the median waist, Preethmoid (PE, Figs. 49-58). The giving it a more "butterflylike" appearance. In preethmoid cornua are paired endochondral the whipplei clade the supraethmoid is further ossifications located laterally between the an- broadened, representing the derived condi- teroventral mesethmoid and vomer. In Cyp- tion. All members of this group except camura, rinella a mesial and lateral ossification are pyrrhomelas, and xaenura also have the bone 120 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY more elongate. Cyprinella caerulea and tal series and borders the orbit anteriorly. The trichroistia have broad, but short su- fifth or sixth infraorbital bone is sometimes re- praethmoids. Species of the nivea group have ferred to as the dermosphenotic. a reduced width to their supraethmoids as in Orbitosphenoid (ORS.Figs. 59, 60). This outgroups, but have retained the elongate large, medial, and V-shaped endochondral condition. element forms much of the floor of the anterior

Kinethmoid. This small dermal bone is neurocranium and mesial wall of the orbit. It elongate and crescentic in shape and some- articulates with the lateral ethmoids and poste- what ornate. It articulates via ligaments with rior planum ethmoidale anteriorly, with the the mesethmoid, maxillaries, andpremaxillar- frontals dorsally, with the pterosphenoids ies. The mesethmoid ligament is forked and posteriorly, and with the parasphenoid ven- attaches to the kinethmoid at a single dorsal trally. The dorsal articulation with the frontals bony tubercle and with the mesethmoid an- is the most complex. The orbitosphenoid plates terolaterally. The maxillary ligaments attach extend dorsolateral^ and fit into a double- to the kinethmoid dorsolateral^ at small lat- walled lamella of the frontal, surrounding the eral processes and with the maxillary at the taenia marginalis anteriorus. This canal con- expanded anterior bony tubercles, behind the tinues anteriorly into the dorsal lateral premaxillary process. The premaxillary liga- ethmoids. Ventrally, the articulation with the ments attach ventrally on the kinethmoid and parasphenoid is with the vertical interorbital to the ascending processes of the premaxillar- septum. This septum is shallow in all Cyp- ies. No significant variation occurs among rinella (Fig. 59D-I) and may have smooth or species of Cyprinella. ornate anterior and posterior margins. Nasal (N, Figs. 49-56). The nasals are Comparison with outgroups indicates that lateral to the supraethmoid and dorsal to the the shallow interobital septum in this group is nasal cavity. These are small and thin dermal derived. Species of the genera Luxilus and Lv- bones and contain the anterior portion of the thrurus, as well as species of many other supraorbital canal. The canal may have one or genera have a deep interobital septum (Fig. two laterally directed pores. Species with two 59A-C). Also derived in Cyprinella is the pores include camura, galactura, venusta, increased proportion of the optic foramen whipplei, caerulea, trichroistia, gibbsi, nivea, bounded by the orbitosphenoid. This is dis- callisema, callitaenia, and callistia. All other cussed under the pterosphenoid. species have the primitive condition for Cyp- Pterosphenoid (PTS, Figs. 59, 60). These rinella, a single lateral pore as in the immedi- ventrally concave and paired endochondral ate outgroups. elements form most of the posterior face of the orbit. Each articulates with the orbitosphenoid Orbital Region anteriorly, frontal and sphenotic dorsolater- The orbit is bounded by 10 paired elements al^, and prootic posteriorly. Mesially these and one median bone, the orbitosphenoid (Figs. two bones rarely connect, but form the mar- 59, 60). Both endochondral and dermal ele- gins of the large and centrally located optic ments are included here. The orbitosphenoid foramen and part of the more posterior hypo- forms the interorbital septum and much of the physeal foramen. Posteroventrally each forms socket. floor of the neurocranium . The orbitosphenoid the antenor corner of the hyomandibular and pterosphenoids are cartilage bones, Stout ventromesially directed struts are formed whereas the frontals, supraorbitals, and in- from this corner, bridging the trigeminal fora- fraorbital bones are dermal. Included in the men and connecting, in some individuals, with infraorbital series are the lacrymal and five in- the ascending wing of the parasphenoid and fraorbital bones located below and behind the the prootic. orbit. The lacrymal is the first of the infraorbi- Two conditions of the pterosphenoid vary NORTH AMERICAN MINNOW GENUS CYPRINELLA 121

PRO

Fig. 59. Lateral view of orbital region in some Cyprinella and outgroups. A) Notropis atherinoides, 53 mm, KU 18935. B) Lythrurus fumeus, 55 mm, KU 6244. C) Luxilus cardinalis, 60 mm, KU 15281. D) Cyprinella venusta, 58 mm, KU 8810. E) C. analostana, 61 mm, INHS 77855. F) C. nivea, 59 mm, KU 18987. G) C. ornata, 56 mm, KU 8405. H) C. ruiila, 57 mm, ASU 5982. I) C. proserpina, 58 mm, TNHC 3262. Stippled pattern at anterior end of parasphenoid represents posterior extent of planum ethmoidale. Horizontal bar equals 1 mm. among species of Cyprinella, the proportion of pyrrhomelas, xaenura, trichroistia, gibbsi, the optic foramen formed by the pterosphe- caerulea, and callistia have a longer process, noid (and orbitosphenoid) and whether or not and species of the nivea group have an ex- a posteromesial process divides the optic and tremely long process, such that they meet hypophyseal foramina (Fig. 60). In members medially and completely divide the two fo- of the lutrensis clade a large proportion of the ramina. anterior optic foramen is formed by the orbito- The development of the mesial process of sphenoid (Fig. 60B, C), much more than other the pterosphenoid is derived in Cyprinella species of the genus or species of the out- since none of the members of Luxilus and groups (Fig. 60A,D-G). Lythrurus possess such a structure. The proc-

Posteriorly, the pterosphenoid may or may ess is, however, in other Notropis (i.e., atheri- not produce a mesial process of variable length, noides), but is here considered independent. depending on the species (Fig. 60). All mem- On a developmental basis, the elongation of bers of the lutrensis group, spiloptera, and out- the process until it meets its opposite is also groups lack a process dividing the anterior considered derived. The decreased proportion optic foramen from the posterior hypophyseal of the contribution of the pterosphenoid to the foramen (Fig. 60A-C). In members of the optic foramen in some species is also derived whipplei clade, excluding spiloptera, a proc- since in closest outgroups the orbitosphenoid

ess is . . little present (Fig 60D-G) Cyprinella camura , has contribution to the foramen. whipplei, analostana, and chloristia have a Frontal (FR, Figs. 49-60). In most Cyp- very small process, galactura, venusta, rinella the paired frontals are the largest bones 122 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

orbital Fig. 60. Ventrolateral view of otic and orbital regions in species of Cyprinella and outgroups. A) Orientation view of and otic regions in Luxilus cornulus, 60 mm, KU 8686, with primitive morphology. B) Cyprinella ornata, 56 mm, KU 8405. C) C. bocagrande, 32 mm, KU 20399. D) C. chloristia, 54 mm, KU 8882. E) C. galactura, 57 mm, KU 12029. F) C. caerulea, 60 mm, KU 18978. G) C. leedsi, 53 mm, KU 18985. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 123 of the cranium. Although some variation ex- rhomboidal shape. This is accomplished by ists, the two bones broadly overlap medially broadening the occipital region of the cranium and articulate with the supraethmoid, lateral and angling the lateral margins of the frontals, ethmoids, supraorbitals, and orbitosphenoid such that the widest point is posterior. Moving anteriorly. Posterolaterally, each overlaps the anteriorly, the straight edges of the frontals sphenotic. Posteriorly, articulation occurs with (not incurved) narrow to a truncated anterior the next largest skull roof bones, the parietals margin. Species with this pattern include and dermopterotics (Fig. 49). whipplei, analostana, chloristia, galactura, Shape of the paired frontals is of particular venusta, caerulea, pyrrhomelas, trichroistia, importance in Cyprinella and is unlike that of gibbsi, nivea, leedsi, callisema, callitaenia, other cyprinids examined. Other cyprinids and callistia. Cyprinella xaenura has a frontal generally have frontals narrowing anteriorly, morphology of parallel lateral margins, pro- the widest aspect being where they contact the ducing a more primitive morphology as in the sphenotic (Figs. 49, 50A-C). The frontals of lutrensis clade. Cyprinella are typically very wide along their In most species the frontals are fairly flat entire length (e.g., Fig. 50D). Little narrowing dorsally with only slight decurving laterally to or mesial incurving of these bones occurs where they articulate with the orbitosphenoids. laterally above the orbits. Together with the In lateral aspect, over the orbit and lateral to supraorbitals, the expanded frontals produce a the orbitosphenoid connection they are slightly lateral margin of the cranium that is quasi- arched upward with the highest point being parallel to the body axis. This condition is lateral and slightly anterior to the epiphyseal modified to some degree in certain groups bar. Anterior and posterior to this point the within Cyprinella where the skull is highly bones curve downward with the lowest point domed. In these forms (proserpina,panarcys, being where they articulate with the sphenotic rutila, and xanthicara) a dorsal aspect of the spine. In lepida, xanthicara, rutila, proser- frontals does not appear as wide (Fig. 51). A pina, andpanares frontals are strongly domed state in orbits 1 is more primitivelike appears some spe- over the (Figs. 5 , 59). This condition cies of the nivea species group. In these spe- most extreme in panarcys and proserpina, cies the frontals are more narrow (Fig. 56). especially the former. The supraorbitals of Generally, however, all Cyprinella have broad these species are also strongly decurved (Fig. frontals, especially over the orbits where there 59). Further, in these two species the skulls are is no constriction, typical of other cyprinids highly domed mesially, the highest point being examined. where the frontals unite. A similar condition, Also typical of species in this genus is the but less extreme, is found in whipplei, truncation of the anterior margins of the fron- analostana, and chloristia. This highly arched tals where the supraethmoid articulation oc- orbit and domed cranial condition derived in curs (Figs. 50-56). The truncated anterior Cyprinella, but independent in the two groups. margins and broad frontal morphologies are Posteriorly, the frontals may or may not derived in Cyprinella.The primitive condition contact the dermopterotics. Members of the is to have angled anterior margins (Fig. 50) lutrensis clade, except rutila, and spiloptera, and narrow frontals as in Phenacobius, camura, whipplei, analostana, chloristia, Rhinichthys, Nocomis, Hybopsis, Lythrurus, pyrrhomelas, callisema, and callitaenia have Notropis (s.s.) and Luxilus. a connection between the frontals and der- Primitively within Cyprinella, the overall mopterotics. The remaining species have no outline of the frontals, plus supraorbitals, is connection, because either the dermopterotic rectangular with squared-off front and sides. or the frontals are not long enough (usually the In some species of Cyprinella this overall former), or, as in species of the nivea group, the outline is modified towards a more or less parietals extend some distance between the 124 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY frontals and dermopterotics. In the nivea group considerable overlap between the supraorbi- this is partially a function of the shape of the tals and lateral ethmoids, completely covering dermopterotic, with a convex mesial edge, and the dorsal edge. the parietals, with a concave lateral edge where Overlap of supraorbitals with the frontals they meet. The connection between the fron- varies between species from none to slight. tals and dermopterotics is derived for Cyp- Species with no overlap or with the supraorbi- rinella, but is reversed within the whipplei tals abutting the edge of the frontals include clade. Outgroup taxa and other species of this spiloptera, galactura, venusta, whipplei, genus lack this connection (Fig. 50). trichroistia, gibbsi, nivea, leedsi, callistia, pros- Supraorbital (SO, Figs. 49-56). There is erpina, and panarcys. Species with slight one supraorbital per side situated lateral to the overlap include lutrensis, garmani, rutila, frontals. In Cyprinella these bones are gener- lepida, formosa, xanthicara, camura, ally long and broad with an acute posterior analostana, chloristia, caerulea, xaenura, margin. The lateral margins are fairly straight pyrrhomelas, calUsema, and callitaenia. Cyp- and the mesial margins are convex, producing rinella ornata also has overlap of these bones, the broadest aspect anteriorly. This condition but in this species the frontals overlap the su- is derived for Cyprinella. Close outgroups praorbitals. typically have narrow supraorbitals with both Overlap between the supraorbitals and the the anterior and posterior ends being equal in lateral ethmoids is derived in this genus. Close width and both narrowing at their extremes. In outgroups lack this condition. The overlap of more distant outgroups such as Rhinichthys, the supraorbitals and frontals appears to be in- Nocomis, and Hybopsis, the supraorbitals are formative in Cyprinella, but variability in oval. outgroups precludes determination of polarity Narrowing of the supraorbitals has occurred at this time. secondarily in panarcys, proserpina, rutila, Infraorbital series (LA, 10 2-6, Figs. xanthicara, chloristia, whipplei, caerulea, 61-63). This series of dermal, platelike bones nivea, leedsi, and callitaenia. Cyprinella cal- extends from the nasal cavity posteroventrally lisema also has narrow supraorbitals, but they down around the orbit and up to the sphenotic are shortened. Cyprinella gibbsi, trichroistia, and pterotic, behind the orbit. Generally there and camura share extremely long and broad are six elements in the series, each bearing part supraorbitals. Cyprinella pyrrhomelas and of the infraorbital canal as a bony tube fused to xaenura both have short and broad supraorbi- the lateral surface. The lacrymal (LA) (Fig. tals. 61) is the first of the series and is the largest. Association of supraorbitals with the lat- This element is lateral to the nasal cavity, eral ethmoids and frontals is variable in Cyp- maxillary, and palatine, and is supported by rinella. Species with no connection between the latter element mesially. The remaining five the lateral ethmoid and supraorbital include infraorbital elements (10 2-6) are generally ornata, lutrensis, lepida, galactura, caerulea, thinner and in some cases more elongate. The xaenura, pyrrhomelas, trichroistia, gibbsi, and second is thin and may be short or long, de- species of the nivea species group. Cyprinella pending on the species. The third is lunate in camura, whipplei, analostana, chloristia, ve- shape with anterior and ascending arms and nusta, andformosa have only a slight overlap. may be broad. The fourth element is short and Most individuals of spiloptera and garmani broad. The fifth and sixth elements are gener- have no overlap between these elements. Other ally the smallest of the series and may be re- individuals have overlap on one side but not duced to only the bony canal tube. These the other. These species are considered to have elements are not illustrated here, but are in line a non-overlapping pattern. Cyprinella rutila, with the posttemporal canal and lateral to the xanthicara, proserpina, and panarcys have temporal fossa (see Harrington, 1955). NORTH AMERICAN MINNOW GENUS CYPRINELLA 125

Fig. 61. Infraorbital series of some species of Cyprinella and close relatives. Infraorbitals 5 and 6 are not illustrated. A) Lythrurusfumeus, 56mm, KU 6244. B)Luxilus cardinal is, 82mm,KU 15281. C) Cyprinella lutrensis, 55 mm.KU 19431. D) C. lepida, 48 mm, TU 55189. E) C. panarcys, 43 mm, UMMZ 208212. F) C. xanthicara, 41 mm, ASU 3642. G) C. spiloptera, 6$ mm,KU 17776. H) C.galactura, 73 mm, KU 12029. I) C. venusta, 84 mm, KU 8810. Horizontal bar equals

1 mm.

The origin of infraorbital 5 and 6 is contro- caerulea, and pyrrhome las (Fig. 62). The long versial. Swift (1970) considered the fifth and lacrymals are found in spiloptera, xaenura, sixth elements to be the dermosphenotic, while and the whipplei and venusta species groups. Harrington (1955) and Howes (1978) equated The extremely long lacrymal is unique to the the fifth with this element. Because the sixth nivea species group. Perhaps correlated with element is frequently attached to the dermop- the length of the lacrymal is the orientation of terotic and the fifth lies lateral to the sphenouc the lateral canal. In members of the lutrensis spine and carries the posterior curvature of the clade, spiloptera, the whipplei and venusta infraorbital canal, typical of the dermosphe- species groups, and the pyrrhomelas species notic, Coburn (1982) considered the fifth ele- pair the lateral canal is smoothly arched dor- ment and the dermosphenotic to be homolo- sally, like outgroup taxa. In the caerulea spe- gous. This argument is followed here. cies complex the canal is straight and oblique Some elements of the infraorbital series are and in the nivea species group it is L-shaped considerably variable between species of Cyp- (Fig. 62). rinella. Other elements vary, but intraspecific In members of the whipplei clade the lacry- variation is as great as interspecific variation mal is flat, while species of the lutrensis clade and is of no use systematically. Only the lacry- have a curled lacrymal, posterior to the canal. mal and infraorbital 2, 3, and 4 varied system- In all Cyprinella except galactura and venusta atically. Variation in the lacrymal includes the dorsal margin of the lacrymal is fairly length and contour of the bone and shape of the straight. These two species have an elevated fused bony canal. Infraorbitals 2, 3, and 4 vary dorsoanterior margin. Orientation of the lacry- in width, length, and number of elements. mal is consistent in the genus, except in callistia, The lacrymal size is represented in Cyp- callitaenia, and callisema. Typically it is nearly rinella by three conditions, short, long, and horizontal, like other cyprinids examined, but extremely long (Fig. 62). Short lacrymals are in these three species it is tilted ventrally (Fig. found in the lutrensis clade, trichroistia, gibbsi, 63). 126 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Most species of the genus and outgroup taxa have an elongate second infraorbital. This bone is short in all members of the lutrensis clade. Within the lutrensis clade infraorbital 2 and the anterior arm of infraorbital 3 are broken into two or three smaller elements in the

lepida species group (Fig. 61). Other members of this clade, species of the whipplei clade, and species of close outgroups all have a complete and long anterior arm to the third infraobital. Cyprinella panarcys is unique in the genus in having a very thin infraorbital 4 (Fig. 61). In B other species of Cyprinella this element is large and platelike, like members of the genus Luxilus. Compared with close outgroups and some distant outgroups, all species of Cyprinella share an elongate lacrymal that is derived for the genus. Even species of Cyprinella with short lacrymals have lacrymals longer than immediate outgroups, relative to body length. Furthermore, within the genus the long and extremely long lacrymal morphologies are considered to be increasingly derived. This is supported developmentally. Young of species with the extremely long condition have lacrymals equal in length and shape to species with the long morphology (Fig. 62). The smoothly arched canal of the lacrymal is primitive in Cyprinella, since most species outside this group have this morphology. The straight and L-shaped canals are derived from this condition. The concave or curled lacrymal is derived within Cyprinella, based on outgroup and developmental criteria. In close outgroups and young of species with the curled lacrymal this element is flat. An elevated anterodorsal corner to the lacrymal and a ventrally tilted lacrymal are derived within the genus. No outgroup taxa examined thus far have either of these two conditions.

Otic and Occipital Regions

Figure 62. Lacrymal bone in some species of Cyprinella The 10 bones of the otic and occipital and close relatives. Luxilus 82 KU A) cardinalis, mm, regions are of varied origin (Figs. 33, 59, 60, 1 528 1 . B) Cyprinella pyrrhomelas, 70 mm, FNHS 76978. 64, 65). The supraoccipital is unpaired and all C) C. gibbsi, 58 mm, KU 1 8892. D) C. spiloptera,6S mm, others are The (autosphe- KU 17776. E) C. nivea, 62 mm, KU 18987. F) C. nivea, paired. sphenotic 40 mm, KU 18987. Horizontal bar equals 1 mm. notic), pterotic (autopterotic), exoccipital, NORTH AMERICAN MINNOW GENUS CYPRINELLA 127

or five surrounding elements. Anteriorly they articulate with the frontals and pterosphenoid, dorsally with the frontals and sometimes with the parietals, posteriorly with the pterotic, and mesoventrally with the prootic. Located lateral to the sphenotic are the last two elements of the infraorbital series, the penultimate element representing the dermopterotic. This element is discussed under the infraorbital section. The most obvious feature of the sphenotic is the posteroventrally directed process. The dorsal surface of this sphenotic process and the lateral surface of the sphenotic form part of the origin surface of the dilator operculi. Ventrally, the entire mesial surface forms the lateral half of the anterior and much of the Figure 63. Ventrally deflected orientation of infraorbital series in Cyprinella callistia, C. callisema, and C. posterior hyomandibular sockets. No callilaenia. series and is C. Stippled hyomandibular significant interspecific variation of the sphe- nivea (58 mm, KU 18987) and represents primitive notic morphology occurs in this genus. condition. Outlined infraorbital series and hyomandibular Prootic 59, The is C. callisema (56 mm, KU 8842) and is the derived (PRO, Figs. 60, 64). proot- condition. ics are large, irregularly shaped endochondral elements that form much of the ventral neuro- epiotic, and the main bone of the supraoccipi- cranium. Posteriorly on each side, each is in tal are all cartilage bones. The dermosphenotic contact with the exoccipital and pterotic and is dermal, free from the sphenotic, and repre- together the three surround the subtemporal sents the fifth element of the infraorbital se- fossa. Posteromesially, the prootic is expanded ries. The dermopterotic is dermal and fused to ventrally and articulates with the anterior the pterotic. The parietals are dermal and the margin of the basioccipital, forming the ante- intercalar and crest of the supraoccipital are rior end of the recessus sacculi. Laterally, the membrane bones. The bones from these re- prootic contributes to the anterior and poste- gions form the posterior neurocranium ven- rior hyomandibular sockets. Mesially it is in trally, ventrolaterally, and posteriorly. contact with theparasphenoid ventrally and its A characteristic shared by all species of opposite dorsally, contributing to the roof and Cyprinella is a compressed and vertical oc- side walls of the posterior myodome and part cipital region, when viewed laterally (Fig. of the floor of the neurocranium. Anterome- 33B,C). Members of the lutrensis clade are sially the prootic projects dorsally and to- most extreme in this morphology. Species of gether with the pterosphenoid forms the mar- the close outgroups Luxilus and Lythrurus, gins of the centrally located hypophyseal fora- and those of the more distant outgroups Notro- men. These bones also surround the antero- pis (s.s.), other \mreso\vedNotropis,Alburnops, lateral trigeminal foramen (FV), the prootic Phenacobius, Hybopsis, and Rhinichthys all forming the posterior margin and the ptero- have a more elongate occipital region, princi- sphenoid generally forming the anterior rim. pally due to the elongation of the supraoccipi- The ventral surface of the prootic is very tal, exoccipitals, pterotics, and epiotics (Fig. irregular and complex. Anteriorly, lateral to 33A). the ascending wings of the parasphenoid, is Sphenotic (SPH, Figs. 49-56, 64). These the lateral commissure, a broad lamella of paired endochondral elements are located lat- bone forming a ventrally depressed bridge erally, behind the orbit and articulate with four separating the facial and trigeminal chamber. 128 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

STF SPH

EOC

FIC PTS

FVII

Fig. 64. Ventrolateral view of prootic in species of Cyprinella and outgroups. A) Ventrolateral view of otic and occipital of Luxilus of and 55 regions cornutus, illustrating position prootic primitive morphology. B) Lythrurusfumeus , mm, KU 6244. C) Cyprinella spiloptera, 56 mm, KU 17776. D) C. xaenura, 75 mm, KU 76978. E) C. pyrrhomelas, 70 mm, INHS 76978. F) C. callisema, 52 mm, KU 8842. G) C. bocagrande, 32 mm, KU 20399. H) C. rulila, 57 mm, ASU 5982. I) C. xanthicara, 49 mm, ASU 3642. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 129

Laterally, it is raised to contribute to the bulla acoustica utricularis which contains the hyomandibular socket and posteriorly much lapillus otolith. of the surface is expanded ventrally to form the No variation in overall shape occurs in

EPO soc

Fig. 65. Lateral and ventral views of the basioccipital of species of Cyprinella and related taxa. A) lateral view of basicranium and ventral view of basioccipital of Luxilus cornutus, 60 mm, KU 8686. B) Basioccipital of Cyprinella camura, 45 mm, KU 15792. C) C. chloristia, 54 mm, KU 8882. D) C. analostana, 61 mm, INHS 77855. E) C. lutrensis, 55mm, KU 19431. F) C.garmani, 50 mm, KU 5416. G) C.formosa, 47mm, KU 8399. H) C.panarcys, 44 mm,\JMMZ 208212. I)C.ru///a,57mm,ASU5982. J)C.trichroistia,60mm,KU 18853. K)C.s*My/,57mm,KU 19982. Horizontal bar equals 1 mm. 130 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Cyprinella. Some species develop a variably may overlap these bones. Posteriorly, the der- sized lateral commissure. In others a narrow, mopterotics overlap the epiotics. As discussed internal jugular strut (Howes, 1979) is devel- previously, they may or may not articulate oped which bridges the facial foramen. Spe- with the posterolateral margins of the frontals. cies with the jugular strut include bocagrande, Medially these paired bones articulate with the formosa, lepida, rutila, xanthicara, proser- parietals and posteriorly they contact the epiot- pina, and panarcys. In all these species, except ics and the paired posttemporals. rutila, the strut is narrow (Fig. 64). In rutila it Within Cyprinella the dermopterotics may is broad, covering most of the trigeminal fora- or may not extend over the lateral edge of the men (FV). Species with a process off the autopterotic and autosphenotic. If expanded, lateral the directed to face of trigeminal foramen , they form a small fairly large roof over the ventromesially, include ornata, lutrensis, dilator fossa where the levator operculi and garmani, analostana, and chloristia (Fig. 64). adductor operculi muscles originate. Species The lateral commissure is broad in most with a roofed dilator fossa include members of

species. Inxaenura and pyrrhomelas it is more the lutrensis clade, spiloptera, camura, narrow, especially in the latter. In these two analostana, galactura, and venusta. All other species a shelf is formed posterolateral to the Cyprinella have no bony extension over the lateral commissure, covering the normally dilator fossa. The lack of a bony extension is depressed region between the trigeminal fora- derived within Cyrinella. Outgroups almost men and the expanded bulla acoustica utricu- always develop this shelf. laris. The shape and orientation of the dermop- In all species cxceptpyrrhomelasandxaen- terotic is diagnostic within Cyprinella. Gener- ura the anterior margin of the trigeminal fora- ally, two basic orientations exist. Either there men is bordered solely by the pterosphenoid. is a fairly shallow slope of the bone laterally, The foramen in these two is bounded anterov- giving the occipital region a more or less flat- entrally by a ventrally directed process of the tened aspect or the dermopterotic slopes later- prootic like most other cyprinids examined. ally at a m uch greater angle, producing a steep- The lateral strut or process crossing the sided occipital region. The flattened condition facial foramen is derived within Cyprinella is present in most Cyprinella and close out- since closest outgroups examined lack this groups. Species with the steeper sided, more feature. The complete strut is also considered highly angled condition include whipplei, derived over the process since juveniles of analostana, chloristia, lutrensis, garmani, these species have only the process. The nar- lepida, formosa, bocagrande, proserpina, row lateral commissure and posterior shelf in panarcys, xanthicara and to a lesser degree xaenura and pyrrhomelas are derived. Out- rutila. Cyprinella ornata has the flattened groups typically have abroad commissure like condition. Extremes of both conditions also other Cyprinella. The posterior shelf is unique exist. Cyprinella gibbsi, trichroistia, and caer- to this species pair. Having the trigeminal ulea all have a more depressed occipital con- foramen bounded anteriorly by the pterosphe- dition, and lepida, xanthicara,proserpina, and noid only is considered derived since species panarcys have a more vertical aspect with the of Luxilus and Lythrurus have the anterior most extreme condition being in panarcys. border composed of both the pterosphenoid The pattern of articulation of the dermop- and prootic, like xaenura and pyrrhomelas. terotic and the parietals varies somewhat be- Dermopterotic (DPT, Figs. 49-56, 59, 60, tween and within species, but generally two 65). The dermopterotics are paired dermal conditions exist. The first condition is to have ossifications overlying the endochondral au- a fairly straight articulation between the two topterotics. Anteriorly, they have varying elements. The second condition and here degrees of articulation with the sphenotics and considered derived, is to have a concave bor- NORTH AMERICAN MINNOW GENUS CYPRINELLA 131 der to the parietals and a convex border on the in Cyprinella is derived. All species examined dermopterotic, such that the appearance is to outside of this group have a well-developed have the dermopterotic invading the parietals. fossa and all close outgroups develop a long Species with this condition but fairly weakly autopterotic spine. developed include spiloptera, caerulea, Parietal (PA, Figs. 33, 49-56, 66). The xaenura,pyrrhome las, trichroistia, gibbsi, and parietals are large paired, dermal bones which callitaenia. Cyprinella nivea, leedsi, callis- cover much of the posterior of the cranium. ema, and callistia show a strong intrusion They are typically rectangular and articulate pattern. medially with one another, anteriorly with the Fused to the dermopterotic is the lateral frontals, laterally with the dermopterotics, and posttemporal canal. This canal extends longi- posteriorly with the epiotics and supraoccipi- tudinally and connects with the laterally di- tal. These bones overlie portions of the su- canal. rected supratemporal Together these praoccipital , autosphenotics, autopterotics, and two canals merge into one posttemporal canal epiotics. on the posttemporal or dermopterotic bone. Variation exists in the lateral margins of the The lateral posttemporal canal has a pore at parietals as discussed previously in reference each end and one in the middle. The posttem- to the dermopterotics and in the posterior poral canal has one medial pore and a single margins where they meet with the supraoccipi- pore at either end. tal and epiotics. In the lutrensis clade a very Autopterotic (APT, Fig. 64). Located definite ridge is developed between where the posterolateral^ on the skull, this endochon- parietals terminate posteriorly and where they dral ossification fits between the anterior sphe- meet with the supraoccipital. This ridge also notic and the posteromesial epiotic and exoc- extends onto the dorsal surface of the parietals cipital. Posteriorly it forms the lateral margin in an arch generally following the arc pattern of the posttemporal fossa together with the of the lateral margins of the supraoccipital exoccipital and epiotic, and mesially partakes crest. In this clade the parietals also extend in the posterior hyomandibular socket as well more posteriorly and cover the dorsal and an- as forming the lateral rim of the subtemporal terior portion of the supraoccipital crest. This fossa. The most conspicuous feature of this posterior extension is magnified even more in bone is the spine and mesial shelf from the panarcys, proserpina, xanthicara, and rutila. spine produced from its posterolateral corner. Other species of Cyprinella also have this but This spine is connected by ligaments to the its development is not as great. These taxa also supracleithrum and posttemporal. Ventrolater- have their parietals ending before the supraoc- ally, the autopterotic contains the lateral semi- cipital crest and no shelf is formed over the circular canal. Ventrally the intercalar articu- epiotics; much of the medial supraoccipital lates with the main body of the autopterotic as and its crest are exposed. well as the mesial shelf of the posterolateral Fused to the posteriormost margin of the spine. parietals and along the rim of the previously The size of the autopterotic spine and mentioned ridge is the supratemporal canal. posttemporal fossa vary in Cyprinella. All This canal is variably shaped within and be- species, except panarcys, have a well-devel- tween species of Cyprinella. oped spine. In this species the spine is reduced Exoccipital (EOC, Figs. 33,49-56, 64-66). to a small process (Fig. 66). The posttemporal These paired, endochondral elements form fossa is large in all species except the caerulea most of the occipital region of the skull. They species complex. In these species the epiotic articulate ventromesially with the basioccipi- and dermopterotic are compressed ventrally, tal. Above the basioccipital a shelf is formed reducing the size of the fossa (Fig. 66). from mesial extensions of both bones. This The reduced spine and posttemporal fossa shelf forms the floor of the posterior neurocra- 132 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY nium and foramen magnum, and the roofof the compound origin (Patterson, 1977). It is com- paired recessi sacculi and the median cavum posed of a large expanded and roughly trian- sinus impar. Dorsally, in addition to the large gular base and a vertical crest posteriorly. The and tear-shaped foramen magnum, two large base is endochondral and the crest is mem- lateral occipital foramina are developed, one brane bone. Anteriorly, the supraoccipital ar- on each side of the midline. Lateral to these the ticulates with and extends beneath the parie- exoccipital is expanded posteriorly and con- tals. Laterally and posterolateral^ it broadly tains the horizontal semicircular canal. The articulates with the epiotics and exoccipitals, dorsal margin of the exoccipital abuts against respectively, and partially roofs the foramen the supraoccipital mesially and epiotic and magnum. Dorsomesially it is concave and pterotic laterally. Anteriorly, it forms part of anteriorly it is convex. Medially, a well-devel- the subtemporal fossa and articulates with the oped ridge is formed from an expansion to prootic and parasphenoid. accommodate the posterior semicircular ca- Except for the exoccipitals being elevated nal. Epaxial muscles are attached to the poste- mesially in the lutrensis,formosa, and lepida rior face of the base of the supraoccipital and species groups, relative to other Cyprinella ligaments from the Weberian apparatus and and outgroups, the morphology of this element predorsal bones are attached to the crest. Except is consistent throughout the genus. As dis- for the compressed supraoccipital, typical of cussed above, this modification of the exoc- all species of Cyprinella, no significant cipital is correlated with the elevation of the interspecific systematic variation occurs in the entire occiptal region. morphology of the supraoccipital. Within a Epiotic (EPO, Figs. 33, 66). Located dor- species a horizontal plate may be formed on solateral^ above the exoccipitals, these endo- the dorsal surface of the crest. chondral bones fit between the mesial un- Intercalar (IC). The intercalar is a small, paired supraoccipital and the lateral pterotics. crescentic endochondral ossification ventral Posteriorly, it is expanded and contains part of to the autopterotic. Generally, it lies under the the horizontal semicircular canal. The anterior mesial side of the autopterotic spine and ex- surface forms part of the inner subtemporal tends up under the posteroventral margin of fossa and the anterior wall encloses the semi- the same bone. Occasionally, it may even circular canal. Projecting posteriorly from the extend over the suture between this bone and expanded epiotic enclosing the canal is a bony the exoccipital. No interspecific variation of process for the insertion of epaxial muscles. systematic significance was noted for this bone Laterally, above the posttemporal fossa a sec- in Cyprinella. ond and larger bony process is formed. This process together with the posttemporal and Basicranial Region pterotic forms the roof of the posttemporal Two bones comprise the basicranial region, fossa. the basioccipital and the parasphenoid (Figs. Variation in this element is minimal. Only 59, 60, 64-66). Both bones are unpaired and members of the caerulea species complex together span the entire length of the neurocra- varied significantly. These species all share a nium ventrally. The basioccipital is a cartilage dorsoventrally compressed posterolateral bone and the parasphenoid is a dermal bone. epiotic where this element contributes to the Both elements serve as attachment areas for

posttemporal fossa (Fig. 66). All other species dorsal gill arch muscles and form the floor, have epiotics similar to outgroup taxa. Thus, roof, and some walls of the neurocranium and as mentioned above, the compressed posttem- posterior myodome. poral fossa is derived in this genus. Basioccipital (BO, Figs. 33, 64-66). This Supraoccipital (SOC,Figs. 33,49-56, 65, bone is located posteroventral ly on the cra- 66). The supraoccipital is an unpaired bone of nium and articulates with the parasphenoid NORTH AMERICAN MINNOW GENUS CYPRINELLA 133

soc EPO

PTF

PPBO

Fig. 66. Posterior view of cranium of species of Cyprinella. A) C. callisema, 52 mm, KU 8842. B) C. gibbsi, 57 mm, KU 18892. C) C. lutrensis, 55 mm, KU 19431. D) C. panarcys, 44 mm, UMMZ 208212. Horizontal bar equals 1 mm. and prootic anteriorly, the exoccipital later- basioccipital and exoccipital forming a bulge ally, and via the proatlas, the first vertebral in the cranium, the bulla acoustica lagenaris. centrum posteriorly. The posterior myodome Mesially, the basiocciptal also forms the floor is partially contained in thebasioccipital where and walls of the cavum sinus impar. it forms much of the posterior walls, roof, and Anteriorly, the basioccipital is forked, pro- floor. Also contained posteriorly is the reces- ducing two processes extending above the sus sacculi which houses the posterolateral^ posterior parasphenoid, and forms the poste- located astericus and the anteromesially lo- rior rim of the oval, centrally located opening cated sagitta. The exoccipital forms the roof in the floor of the posterior myodome. Pos- and most of the lateral surfaces of this cavity. teroventrally the pharyngeal process of the The location of the recessus sacculi can be basioccipital is developed. This large process noted by the obvious ventral expansion of the is laterally compressed posteriorly, forming a 134 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

vertical plate with a rounded posterior edge. shaft slopes posteroventrally to the posterior Anteriorly this vertical plate widens ventrally end of the optic foramen. Here it is expanded into a semi-horizontal, concave masticatory laterally, is more horizontal, and produces two plate which opposes the pharyngeal teeth. dorsolateral ascending wings, one per side. Dorsal to the plate is a supporting arch and These wings articulate with the prootic later- canal formed between the pharyngeal process ally and form a bridge between the postero- of the basioccipital and the dorsal portion of laterally placed carotid foramen (which is the basioccipital, through which passes the bounded by the prootic and parasphenoid) and dorsal aorta. the mesially located posterior myodome (also The shape of the dorsal portion of the ba- bounded by these bones). Posteriorly, the par- sioccipital is consistent in Cyprinella, but the asphenoid arches dorsally and narrows. At its morphology of the pharyngeal process varies posterior edge it is forked and ventral to the systematically between species (Fig. 65). All forked anteroventral surfaces of the basioc-

species have concave ventral surfaces to the cipital. The forked ends of these two bones masticatory plate, except lepida, garmani, together form the rim of the large, centrally pyrrhomelas, xaenura, caerulea, gibbsi, and located, and oval opening in the floor of the trichroistia, which have a relatively flat ven- posterior myodome. tral surface. Most species develop a pharyn- The width of the ascending wings is vari- geal pad with two lateral to posterolateral able between species (Fig. 67). All species wings and a pointed posterior margin. In have a broad ascending process, relative to formosa, lepida, panarcys, proserpina, xan- most outgroup taxa, but in some species it is thicara, and rutila, however, the pad is oval in broader than in others. Cyprinella callistia, shape, lacking the lateral wings. pyrrhomelas, camura, whipplei, analostana, All species of the lutrensis group, except and chloristia have very broad ascending xanthicara and rutila, have a deep, triangular wings, especially the latter two taxa. pharyngeal process (Fig. 65). These two spe- The length of the anterior neck of the paras- cies have a very narrow and elongate process. phenoid, anterior to the ascending wings var- All other members of the genus except ies in Cyprinella (Fig. 67). At a similar body analostana and gibbsi develop a narrow proc- size, all species of the lutrensis clade have ess like the outgroups. These two species have shorter necks, as do camura, callistia, callis- a deep, spatulate and rounded process. ema, and leedsi. The latter two species are The deep, triangular pharyngeal process of somewhat more elongate than others. All other the lutrensis clade is derived since no out- members of this genus have longer necks on groups have this morphology. The rounded the parasphenoid. pharyngeal pad is also derived, being found in Because most close outgroups, exceptions this group and convergently in some members being coccogenis and fumeus (Luxilus and of the genera Phoxinus and Pimephales. The Lythrurus, respectively) have a short anterior deep and rounded pharyngeal process of neck to the parasphenoid the long neck in analostana and gibbsi is also derived but inde- Cyprinella is hypothesized derived. The broad pendent. ascending wings are also hypothesized de- Parasphenoid (PS, Figs. 57-59, 64, 65, rived. In Luxilus and Lythrurus and many 67). This elongate dermal ossification extends other outgroups the narrow condition is com- most of the length of the neurocranium ven- mon.

trally. Anteriorly it is spatulate and lies above the vomer and below the posterior planum Opercular Region ethmoidale, ventromesial to the lateral Four dermal bones form the opercular se- ethmoids, and posterior to the mesethmoid. ries, the opercle, preopercle, subopercle, and Extending posteriorly mesial to the orbits, its interopercle. All elements are thin and paired. NORTH AMERICAN MINNOW GENUS CYPRINELLA 135

mesially, but is relatively flat ventrally. The dorsal margin varies individually within a species from being straight, concave, and occasionally convex. Dorsoanteriorly a process of variable length is produced for the insertion of the dilator operculi. Below this is a variable size notch in the anterior face into which the posterodorsal margin of the hyomandibular inserts anterior to the opercular condyle. The anterior margin is generally straight or slightly sigmoidal. The posterior margin can be sigmoidal or straight. If straight, it is directed posteroventrally at about 45 degrees from the horizontal. Ventrally, the opercle is much wider than the dorsal margin and is variably inclined from the horizontal. Interspecific variation is most notable in the width of the opercle. Most species of the lutrensis clade and analostana have a narrow

and deep opercle (Figs. 68, 69). Cyprinella formosa,xanthicara, and rutila are exceptions in the southwestern clade. These species have a broad opercle (Fig. 68). Cyprinella galactura is unique among other Cyprinella in having a very broad opercle (Fig. 69). Preopercle (POP, Figs. 68, 69). The preopercle is an elongate, quasi-crescentic dermal ossification. The long anterior arm articulates dorsally with the posterior arm of the quadrate and ventrally with the interopercle. The long ascending arm articulates with the of Fig. 67. Ventral view of parasphenoid of species anterior edge of the opercle and the posterior and 53 Cyprinella outgroups. A) Notropis atherinoides, edge of the hyomandibular (Figs. 68, 69). On mm, KU 1 8935. B) Luxilus cornutus, 60 mm, KU 8686. the lateral surface of this bone is a large portion C) Cyprinella camura, 45 mm, KU 15792. D) C. garmani, of the canal of the 50 mm, KU 5416. E) C. chloristia, 54 mm, KU 8882. F) preoperculomandibular C.xaenura, 76 mm, KU 18994. G) C. leedsi, 53 mm, KU lateralis system. 18985. C. callisema, 52 mm, KU 8842. Horizontal H) In all Cyprinella this bone is broadened bar equals 1 mm. medially and anteriorly more than any outgroups examined. Some interspecific variation Together they cover the branchial chamber exists, however, in the width of the bone and and all posteroventrally ventrolaterally. They the height of the ascending arm. The broadest interarticulate and some articulate with the condition is found in camura, galactura, and neurocranium and bones of the hyopalatine venusta (Fig. 69). A narrowed condition oc- region dorsally, ventrally, and anteriorly. curs in rutila and xanthicara (Fig. 68). In these This is the Opercle (OP, Figs. 68, 69). two species broadening of the preopercle is In most largest bone of the opercular series. restricted to the anterior arm only. The ascend- it is a broad bone as in most out- Cyprinella ing arm and the medial portions of this bone in and it is concave groups. Dorsally medially these species are similar to the condition found 136 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

PAL

Fig. 68. Lateral view of opercular series and hyopalatine region in some species of Cyprinella and outgroups. A)Notropis

53 1 8935. 52 6244. Luxilus 72 INHS 79254. atherinoides, mm, KU B) Lythrurusfumeus , mm, KU C) coccogenis, mm, D) Cyprinella luirensis, 55 mm, KU 19431. E) C.formosa, 42 mm, KU 8399. F) C. lepida, 44 mm, TU 55189. G) C. panarcys, 43 mm, UMMZ 208212. H) C. rulila, 55 mm, ASU 5982. All drawn to equal size from anterior endopterygoid to ventral tip of hyomandibular for comparisons. Horizontal bar equals 1 mm. in outgroups. All other Cyprinella are similar The margins of the preopercle are generally in having a broadened condition. Height of the entire, except for occasional notches and spurs ascending arm varies in two different species which varied on an individual basis. Cyp- groups. Cyprinella xaenura and pyrrhomelas rinella rulila, proserpina, andpanarcys varied have tall ascending arms, while gibbsi, from this in having distinctly scalloped pos- trichroistia, and caerulea have reduced as- teroventral and ventral margins (Fig. 68), being cending arms, compared to other Cyprinella best developed in the latter two species. and outgroups (Fig. 69). Interopercle (IOP, Figs. 68, 69). This is an NORTH AMERICAN MINNOW GENUS CYPRINELLA 137

Fig. 69. Lateral view of opercular series and hyopalatine region in species of Cyprinella. A) C. camura, 73 mm, KU 1 971 5. B) C. analostana, 65 mm, INHS 77855. C) C. galactura, 60 mm, KU 12029. D) C. pyrrhomelas, 70 mm, INHS 76978. E) C. gibbsi, 57 mm, KU 18892. F) C. leedsi, 60 mm, KU 18985. G) C. callisema, 55 mm, KU 8842. H) C. callitaenia, 60 mm, TU 92770. All drawn to equal size from anterior endopterygoid to ventral tip of hyomandibular for comparison. See Figure 68 for identification of elements. Horizontal bar equals 1 mm.

elongate dermal bone anterolateral to the Most species of Cyprinella have the posterior subopercle and ventromedial to the preopercle. of this bone expanded more than most other Mesially it articulates with the posterior cera- cyprinids examined. The closest outgroups to tohyal and the interhyal. In all species this Cyprinella(Luxilus,Lythrurus,Notropis (s.s.), bone is broadest posteriorly and tapers to about and Nocomis) have a narrow interopercle (Fig. one half that height anteriorly, where it is 68). Cyprinella whipplei, trichroistia, calli- ligamentously connected to the retroarticular. taenia, and callisema have narrow interoper- 138 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY culi similar to other cyprinids. All other species have some expansion at the posterior Hyopalatine Region margin. The deepest condition is found in The seven paired elements comprising the ornata, lutrensis, formosa, bocagrande, hyopalatine region of the cranium form the proserpina, panarcys, camura, analostana, supporting structures of the lower and upper leedsi, venusta, gibbsi,pyrrhomelas,xaenura, jaws, the palate, and the opercular series (Figs. caerulea 68 Included is the nivea, and (Figs. 68, 69). These spe- , 69). cartilaginous hyomandi- cies have the interopercle about twice as high bular, symplectic, quadrate, palatine, and met- as close outgroups. Other species have a broad- apterygoid, and the dermal endopterygoid and ened interopercle, but less extreme man those ectopterygoid. The dorsolateral surfaces of the listed above. Cyprinella rutila is apparently pterygoids form the floor of the orbit. unique in having a short interopercle (Fig. 68). Hyomandibular(HYO,Figs.68,69).This is the largest and heaviest endochondral bone Subopercle (SOP, Figs. 68, 69). This is the of the hyopalatine region. It is widest dorsally posteroventral bone of the opercular series. where it articulates with the cranium via two Generally, it has a crescentic shape with con- cartilage-tipped condyles. The smaller ante- cave dorsal and ventral margins. Anteriorly it rior condyle is ball-like and fits into a facet may be rounded or blunt and with or without a formed by the prootic, pterosphenoid, and dorsoanteriorly directed nipple for articula- autosphenotic. The posterior articular facet is tion with the posteromesial surface of the more elongate, like the condyle, and is formed interopercle. The anterior margin varies by the prootic, autosphenotic, and pterotic intraspecifically. The subopercle articulates bones. Below the posterior condyle a knob- for most of its length dorsally with the ventral like condyle is produced, extending posteri- margin of the opercle. Anteriorly, it is mesial to orly and articulating with the operculum via the interopercle and preopercle. In all Cyp- the dorsoanterior articular facet for the rinella except xanthicara and galactura, this hyomandibular. Ventrally the hyomandibular bone is broader than many close outgroups gradually narrows to a slim, yet heavy shaft examined, except Luxilus pilsbryi, which has with a cartilage covered tip. Here it articulates an equally broad subopercle. Although a cor- with the symplectic, interhyal, and two points relation has been noted between narrow oper- on the metapterygoid. The heavy median strut culi and deep subopercles (Coburn, 1982) all extends for most of the length of the hyomandi- Cyprinella, except where noted in the discus- bular and contains the variable sized oval fora- sion of the opercle, have noticably broader men for the hyomandibular branch of the vagus operculi. nerve. Dorsally, this thickened supporting strut divides into three struts supporting the ante- Orientation of the subopercle varies rior, posterior, and opercular condyles, and interspecifically. Most species have an orien- delineates regions for attachment of the dila- tation like that found in most outgroups, with tor operculi, adductor mandibulae, and leva- the posteriormost tip being considerably ele- tor arcus palatini. vated above the anterior. Informosa and boca- Extending to either side of the median grande the subopercle is relatively straight, supporting strut is a thin sheet of bone for the posterior tip being only moderately ele- muscle attachment. The anterior wing begins vated above the dorsalmost point anteriorly. In at the dorsoanterior condyle and terminates at caerulea, analostana, ornata, lepida, rutila, a point between the ventral tip and midlater- xanthicara, panarcys, and proserpina the ally, depending on the species. This wing is the posterior subopercle tip is extreme, being point of insertion of the levator arcus palatini. greatly elevated above the anterior tip, espe- The heavier posterior wing is rarely as broad as cially in the latter two species. theanterior. Dorsally, thisposterior wing begins NORTH AMERICAN MINNOW GENUS CYPRINELLA 139 anterior and ventral to the opercular condyle. bocagrande, proserpina, panarcys, lepida,

It may begin as a semi-horizontal ridge from camura, and pyrrhomelas . Generally in these the point of division of the median strut, or it species only a small wing extending from the may be a continuation of a dorsally extending median strut of the hyomandibular is present ridge below the posterior condyle. This wing and it only occasionally overlaps the preopercle. extends ventrally, lateral to the ascending arm In trichroistia, gibbsi, and callisema the wing of the preopercle for most of its length. It then extends farther ventrally than in other species, tapers anteriorly where the broadened pre- to near the ventral tip of the median strut (Fig. opercle begins to overlap the lower portion of 69). Contour of the posterior adductor wing the hyomandibular. The posterior wing acts as varies between species from flat to highly a point of origin of the subdivisions of the curled. All species have a flat wing except adductor mandibulae (Winterbottom, 1974; whipplei, analostana, chloristia, callistia, Coburn, 1982). trichroistia, gibbsi, and caerulea which de- Variation occurs in the size and shape of velop a distinctly concave wing. anterior and posterior wings. The anterior wing Polarity of these characters varying in Cyp- of whipplei, chloristia, analostana, camura, rinella are as follows. The notched or smooth lutrensis, garmani, proserpina, panarcys, morphology is discussed by Coburn (1982). lepida,formosa, and pyrrhomelas is small like He considered the smoothed edge to be de- that found in the genera Notropis, Lythrurus, rived, based on comparisons to other genera Pteronotropis, and the closely related genus which commonly have an anterior wing pro- Luxilus (Figs. 68, 69). Other Cyprinella have ducing a distinct angle or notched morphol- a more expanded anterior wing (Figs. 68, 69). ogy. My observations are in agreement, that The shape of the anterior wing also varies the smooth condition may represent a derived interspecifically. All species with the small condition, but it has obviously reversed at least wing have a smooth edge except rutila and two times, once in Cyprinella and at least once xanthicara which have a median oval expan- in some species of Notropis. This is probable sion anteriorly, and ornata and pyrrhomelas since it occurs not only in the whipplei clade, which have a prominent notch midway along but also in species of a large clade of Notropis the hyomandibular (Figs. 68, 69). The notched related to the subgenus Notropis and other morphology is common to the remaining spe- species of uncertain affinities. cies with the broadened anterior wing, except Since species of Luxilus and most other callisema, gibbsi, and trichroistia. In these species possessing the derived condition of species the anterior notch is lacking and the having an opening in the posterior myodome wing is broad, but has a rounded and gradually have a narrow hyomandibular with a smooth sloping anterior margin which extends most of and reduced anterior wing, the broadened the length of the median strut. hyomandibular and the notched anterior wing Separation of the "notched" and "smooth" present in some Cyprinella is derived. The morphologies is not always complete for some curled adductor wing found in two groups of species. Occasionally one or two specimens Cyprinella is also derived since all Luxilus vary from the most frequently observed condi- have a flat posterior wing and the curled con- tion for that species. These variants were dition is rarely encountered elsewhere. sometimes due to obvious developmental ab- Palatine (PAL, Figs. 68-70). These paired normalities. endochondral ossifications articulate posteri- The posterior adductor wing also varies in orly with the endopterygoid and anteriorly size and contour. The wing is expanded pos- with the supraethmoid, vomer, maxillary, teriorly to cover the opercular condyle par- kinethmoid, and lacrymal. The posterior ar- tially or entirely in all species of Cyprinella ticulation involves a simple socket, formed by except ornata, lutrensis, garmani, formosa, the endopterygoid. Generally, there are two 140 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ventral ligaments on the posterior palatine. small dorsal process (Fig. 70M-O). connects to the anteromedial face of the All trichroistia One species except , gibbsi , xaen- endopterygoid, the other connects the dor- ura, and callistia have a well-developed, deep, soanterior endopterygoid to the dorsal surface and contained anteromesial socket for the ar- of the palatine (Fig. 70A). These points of con- ticulation of the preethmoid (Fig. 70). In nection on the palatine are generally well trichroistia and gibbsi the socket is smaller marked by one or two small ligamental scars or and more restricted anteriorly (Fig. 70J). In ridges ventrally and a single scar dorsally. callistia and xaenura the socket is less re- The anterior end of the palatine is consid- stricted posterodorsally and a weak channel is erably more complex than the posterior, con- formed (Fig. 70H,K). sisting of three processes, a maxillary (MP), Interspecific variation is observed in the vomer (VP), and supraethmoid (SEP) (Fig. number of ventral ligamentous scars from the 70A,C). The maxillary process is not well endo- and ectopterygoids. In all species of defined in Cyprinella. This process is lateral Cyprinella excqpttrichroistia, gibbsi, caerulea, to callistia and is connected via ligaments the lacrymal, xaenura, pyrrhomelas , and two scars kinethmoid, and maxilla. The two medial occur ventrally as in all other cyprinids exam- processes, one dorsal and one ventral, form an ined. In these species, only a single ventral articular socket, like other cyprinids, with ridge is developed. This observation is related which the preethmoid articulates. The dorsal to the development of the number of connect- process is connected to the supraethmoid and ing ligaments. In all but the above mentioned the ventral with the vomer. Both connections species there are two branches of the ligament are ligamentous. connecting the posteroventral palatine to the In general, the gross shape of the palatine endo- and ectopterygoid. Species with a single when viewed dorsally (Fig. 70) does not differ scar develop only one large ligament (Fig. 70). greatly in Cyprinella . Most species have heavy The elongate and more delicate palatines of palatines that are typically triangular, with the nivea, leedsi, callisema, and callitaenia repre- broadest aspect anteriorly, and are without a sent a derived morphology relative to the heavy well defined maxillary process. This shape is and short condition in other Cyprinella and derived and also typical of species of the near outgroups. Further, within this group the genera Luxilus and Lythrurus (Fig. 29). reduced ventromedial vomerine process of the The following characters vary between latter three species is derived and supports species. The palatines of nivea, leedsi, callis- their close relationship. The well-developed ema, and callitaenia are more elongate than anteriorly directed maxillary process of rutila, other species of Cyprinella and outgroups at a xanthicara, proserpina, and panarcys as well similar body size (Fig. 70L-O). These four as nivea, leedsi, callisema, and callitaenia species and rutila,xanthicara,proserpina, and (Fig. 70) supports the monophyly of these panarcys have a well-developed maxillary groups, although independent evolution of this process on the palatine which is directed an- morphology is hypothesized. The reduced teriorly (Fig. 70L-O,S,T). The dorsal su- anterior preethmoid socket of the palatine in praethmoid process of most species is similar, pyrrhomelas, callistia, trichroistia, and gibbsi rising above the horizontal 70-80° and termi- represents a derived condition and supports nating in a small point. Cyprinella callitaenia their monophyly. Cyprinella xaenura, another differs in having a broad dorsal process (Fig. member of this group, is apomorphic in having 70 O) and in ornata, panarcys, and lepida the a very long and modified socket. In trichrois- angle of the dorsal process is considerably less tia and gibbsi the socket is even more reduced (Fig. 70P-Q). The ventral process is well- and represents a further modification relating developed in all species except leedsi, callis- these two species. ema, and callitaenia. These species have a The number of ligaments connecting the NORTH AMERICAN MINNOW GENUS CYPRINELLA 141

Fig. 70. Dorsal (left) and medial (right) view of palatines of species of Cyprinella and outgroups. A) Notemigonus 63 1357. Phoxinus 6244. crysoleucas, mm, KU B) erylhrogaster, 52 mm, KU 13198. C) Lythrwus fwneus , 52 mm, KU D) Luxilus zonistius, 72 mm, KU 1 8995. E) L. chrysocephalus, 78 mm, KU 12654. F) Cyprinella spiloptera, 69 mm, KU

17776. G) C. venusta, 83 mm, KU 8810. H) C. xaenura, 72 mm, KU 18994. I) C. pyrrhomelas, 73 mm, EMHS 76978. J) C. trichroistia, 72 mm, KU 18853. K) C. callistia, 74 mm, KU 18848. L) C. nivea, 60 mm, KU 18987. M) C. leedsi, 52 mm, KU 1 8985. N) C. callisema, 52 mm, KU 8842. 0) C. callitaenia, 58 mm, TU 92770. P) C. ornata, 56 mm, KU 8405. Q)C. /epida, 46 mm, TU 55189. R) C.formosa, 47 mm, KU 8399. S) C.proserpina, 50 mm, TNHC 3262. T)C. xanlhicara, 41 mm, ASU 3642. Horizontal bar equals 1 mm. 142 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ventral palatine to the ecto- and endopterygoids Similar to species of Luxilus, in most is also phylogenetically informative within members of Cyprinella the endopterygoid is Cyprinella. The derived condition of having a not the largest bone of the pterygoid series and single ligament instead of two is found in the is equal in size to the metapterygoid. Cyprinella group composed of xaenura, pyrrhomelas, xanthicara is unique in having a larger endop- trichroistia, gibbsi, caerulea, and callistia. All terygoid. In most outgroups examined the other Cyprinella and outgroups examined have endopterygoid is larger than the metapterygoid two ligaments. and represents the largest ossification of the Ectopterygoid (ECP, Figs. 68, 69). These pterygoid series. Coburn (1982) noted that paired, dermal ossifications lie anterior to the Notropis volucellus,N. maculatus,N. emiliae, quadrate and endopterygoid and posteroven- and N. heterolepis also have equally sized tral to the palatine. They articulate ventrally meta- and endopterygoids. Notropis ozarca- with the medial face of the anterior portion of nus and N. spectrunculus also have similarly the quadrate and dorsally with the lateral face sized elements. Other Notropis, Phenacobius, of the endopterygoid. In some instances these Hybopsis, Nocomis, and Rhinichthys gener- bones may be partially fused and their outlines ally have longer endopterygoids. Thus, having obliterated. Generally, this bone is oval or pear roughly equally sized endo- and meta- shaped with the dorsoanterior tip being con- pterygoids is derived. stricted and oriented slightly anterior (Figs. Quadrate (QA, Figs. 68, 69). This long and 68, 69). heavy endochondral bone lies ventral to, but In nivea, leedsi, callisema, and calli taenia articulates with the pterygoid series. Anteri- the bone is short and broad (Fig. 69). Cyprinella orly, the quadrate is compressed laterally into callitaenia is most extreme with the bone ap- a broad fan which is closely associated with proaching a circular aspect. Species of the the ventral ecto- and endopterygoid and loosely lutrensis clade, except xanthicara, have an associated with the an teroventral margin of the elongate and narrow ectopterygoid (Fig. 68). metapterygoid. Anteroventrally it slopes later- Cyprinella xanthicara has a broad and short ally from the expanded fan into a broad semi- condition, similar to that present in species of horizontal shelf (typical of all Cyprinella). An- the nivea group. teriorly it bears a condyle for articulation with Most outgroups examined have an oval or a notch in the posterior articular. Posterior to pear shaped ectopterygoid as is found in spi- the fan is produced a long, heavy recurved loptera, and the whipplei, venusta, and process lateral to the symplecuc. This process pyrrhomelas species groups. Therefore, con- is a continuation of the broad shelf posterior to ditions present in the lutrensis clade and in the the condyle. The process curves upward pos- nivea species group are derived. teriorly, approaching the ascending arm of the Endopterygoid (ENP, Figs. 68, 69). The preopercle and ventrolateral to the posterior endopterygoid is a paired, dermal ossification margin of the symplectic. which articulates posteromesially with the In most species the posterior process of the metapterygoid, ventromesially with the quad- quadrate is heavy with its dorsal margin join- rate and ectopterygoid, and anteriorly with the ing the expanded fan at its posteriormost edge palatine. Like most species with an open pos- (Figs. 68, 69). The posterior process is heavy terior myodome the lateral surface of the en- and deep in analostana, chloristia, gibbsi, dopterygoid is concave. Anteriorly, a socket is lutrensis, and proserpina (Figs. 68, 69). In formed for the articulation with the posterior ornata, rutila, proserpina, panarcys, and palatine. Mesial to the socket an anterior proc- formosa the posterior process joins the fan at a ess is formed for support of the palatine. This point midway along its posterior margin such process glides along the ventral surface of the that the union is anteroventral to the posterior lateral ethmoids. edge. NORTH AMERICAN MINNOW GENUS CYPRINELLA 143

Among outgroups, connection of the poste- However, it is narrow and low in camura, rior arm to the fan occurs at the edge of the fan galactura, whipplei, trichroistia, callitaenia, and in no case is the quadrate as heavy and callistia, garmani, rutila, formosa, and xan- deep as that observed in the species mentioned thicara . In most species the ventral margin of above. the metapterygoid is relatively straight, fol- Symplectic (SY, Figs. 68, 69). Anteriorly lowing the dorsal margin of the symplectic. this elongate and paired cartilage bone lies Cyprinella proserpina and panarcys have an mesial to a thin sheet of bone connecting the anteroventral process extending into a region anterior and posterior portions of the quadrate. normally occupied by the symplectic. Along the remainder of its length posteriorly it The size of the posteroventral point of ar- fits into a channel along the dorsal surface of ticulation of the metapterygoid with the in- the posterior extension of the quadrate. At its terhyal varies between species. In all Cyp- posteriormost extent the symplectic is gener- rinella except whipplei, caerulea, nivea, leedsi, ally more expanded and participates in a four- callisema, and xanthicara this process is very part articulation including the descending small and almost pinpoint in appearance. In ramus of the hyomandibular, the posteroven- these species the articulation is broad. Species tral arms of the metapterygoid, and the in- of the lutrensis clade are most extreme in this terhyal. character in having a very small articular In all but a few species of Cyprinella the connection. Outgroups examined have a broad symplectic is a very broad and flat bone. Ex- articular connection with the interhyal, a ceptions to this include whipplei, caerulea, smooth ventral edge, and a low posterodorsal xaenura, callisema, and rutila. Outgroups are process. variable with respect to the size of the symplectic, but comparisons with closest outgroups Upper Jaw

Luxilus, Lythrurus, and Notropis (s.s.) indi- Premaxilla (PMX, Fig. 71). The premaxil- cate that the elongate and narrow morphology laries are thin and elongate dermal ossifications is primitive. Thus, the broad condition of most which in most species are semi-L-shaped. Cyprinella is derived. Anteriorly, they develop a roughly triangular- Metapterygoid (MPT, Figs. 68, 69). In shaped ascending process which slides be- lateral aspect this is a broad, but thin bone with tween the premaxillary and rostral processes a flat to slightly concave ventral portion. Dor- of the maxilla. The paired premaxillaries ar- sally there are two ascending processes, one ticulate with each other anteriorly along the posterior and one anterior, separated by a mesial edge of the ascending process and dor- prominent middorsal notch. Anteroventrally, sally with the kinethmoid via a ligament. it broadly articulates with the quadrate via a Posteriorly, these bones become constricted to cartilage interface and anterodorsally it is a narrow shaft which may be rounded or closely applied to the lateral surface of the pos- flattened. Over most of the length the bone is terior endopterygoid. Posteriorly, there are two thin with the lateral surface slightly convex. heavy struts, one oriented horizontally and Considerable interspecific variation exists articulating with the hyomandibular and one in the morphology of the premaxilla in Cyp- oriented ventroposterior and opposing the rinella. Orientation of the long posterior ra- interhyal. mus varies from straight to strongly decurved. The anterior ascending arm of the dorsal Species with a straight process include surface is well developed in all Cyprinella, bocagrande, lepida, whipplei, camura, and as a continuation of the endopterygoid, is spiloptera, pyrrhomelas, and trichroistia. The strongly concave. The posterodorsal process remaining species have some degree of curva- shows some interspecific variation. Most spe- ture but proserpina, panarcys, rutila, cies have a broad and high posterior process. analostana, chloristia, and gibbsi are most 144 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY extreme in having strongly decurved processes. tion with the lateral process of the palatine and Cyprinella nivea, leedsi, callisema, and calli- preethmoid. The anteriormost tubercle articu- taenia also have a decurved posterior ramus, lates with the preethmoid. The lateral palatine especially nivea, but in these taxa the change process is joined at the posteriormost tubercle from horizontal is more abrupt and not as of the maxilla and is also ligamentously joined gradual as in other species. The posterior proc- to the lateral face of the maxilla slightly poste- ess of all species is restricted posteriorly. In rior to the premaxillary process. Medially the trichroistia, gibbsi, and caerulea it is extreme. maxilla is deeply notched dorsally, creating In these species it is rounded and tapered into the isthmus. Posterior to the isthmus is a large a very thin shaft. and broad ascending ramus and an elongate The dorsal margin of the premaxilla is posteroventral shaft. The shaft is expanded smooth in all species except nivea, leedsi, and curved posteriorly where the posterior callisema, and callitaenia. These species dif- arm of the premaxilla is connected and articu- fer in having a serrated posterodorsal margin. lates. The margins of the premaxilla of callitaenia, The lateral face of the maxilla is slightly and to a certain degree leedsi, are only slightly convex and smooth, except anteriorly where serrated. two small, laterally directed tubercles are de- In all species the extension of the thick veloped. The anteriormost tubercle is for the posterior process anteriorly produces a slightly insertion of the palatine-maxillary ligament thick dorsal margin to the premaxilla. In pros- and is at the base of the premaxillary process. erpina and panarcys this shaft is heavier than The second process is for the insertion of the other members of the genus. These two spe- adductor mandibulae A 1 and is slightly poste- cies, together with xanthicara and rutila, also rior to the first, located along the ventral edge differ from other Cyprinella in having a very of the maxilla, directly below the large dorsal deep anterior premaxilla. articulation surface for the palatine. The ascending process of the premaxilla in Significant systematic variation in maxilla most species is triangular, narrowest dorsally morphology includes location of the adductor where it articulates with the kinethmoid. Some mandibulae Al insertion, depth of the isthmus members vary, however, in having a more and premaxillary process, and the orientation elongate and taller process. This includes rutila, of the rostral process. In all species except xanthicara, analostana, chloristia, callisema, venusta and proserpina the adductor mandi- callitaenia, nivea, callistia, and gibbsi. bulae Al tubercle is placed far forward rela- Maxilla (MX, Fig. 72). Together with the tive to other cyprinids examined. In members premaxillaries these paired dermal ossifications with the more anteriorly placed process two comprise the protrusible upper jaws. The groups are recognizable. All species, except complex mechanics are possible only with the caerulea, gibbsi, trichroistia, andxaenura have rotational and gliding actions between the the process placed below the posterior margin maxilla, premaxilla, and surrounding articular of the dorsal palatine-maxillary articular sur- surfaces. Anteriorly the maxilla bears two face. In these four species the process is inter- processes, a mesial fingerlike rostral process mediate between that found in venusta and (RPM) directed anteroventrally and a broad proserpina and species with the extreme ante- lateral premaxillary process (MXM). Between rior process. these two the ascending ramus of the premax- Some Cyprinella have a deep isthmus, thus illa is contained and glides when the mouth is a shallow dorsal notch, and a deep premaxil- opened. The rostral process is joined to its lary process as compared to other species and opposite at the midline by a ligament. Above outgroup taxa. Species with a deep isthmus the rostral and premaxillary processes are two include nivea, leedsi, callisema, callitaenia, robust and larger protuberances for articula- proserpina, ornata, camura, chloristia, and NORTH AMERICAN MINNOW GENUS CYPRINELLA 145

Fig. 71. Premaxilla of species of Cyprinella and outgroups. A) Notropis atherinoides, 50 mm, KU 18935. B) Lythrwus lirus, 45 mm, KU 18933. C) Luxilus cardinalis, 65 mm, KU 15281. D) L. cornulus, 60 mm, KU 8686. E) Cyprinella lutrensis, 55 mm, KU 19431. F) C. bocagrande, 32 mm, KU 20399. G) C. proserpina, 58 mm, TNHC 3262. H) C. rutila, 57 mm, ASU 5982. I) C. spiloptera, 56 mm, KU 17776. J) C. camura, 45 mm, KU 15782. K) C. whipplei, 55 mm, KU 14211. L) C. analostana, 61 mm, INHS 77855. M) C. pyrrhomelas, 70 mm, INHS 76978. N) C. trichroistia, 60 mm, KU 18853. O) C. gibbsi, 57 mm, KU 18892. P) C. caerulea, 60 mm, KU 18978. Q) C. mvea, 59 m, KU 18987. R) C. callisema, 52 mm, KU 8842. Horizontal bar equals 1 mm. 146 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ASP

MXM A1T

72. of and atherinoides 53 KU 18935. Fig. Maxilla of species Cyprinella outgroups. A) Notropis , mm, B)Lythrwuslirus, 47 mm, KU 18933. C)L. bellus, 46mm, KU 14511. D) Luxilus cornulus, 62mm, KU 8686. E) Cyprinella garmani, 50 mm, KU 5416. F) C. proserpina, 58 mm, TNHC 3262. G)C.fw»iare)w, 42 mm, UMMZ 208212. H) C. bocagrande, 32 mm, KU 20399. I)C. vern^ta, 83mm, KU 8810. J) C. camura, 76mm, KU 15792. K)C. whipplei, 64mm, KU 14211. L) C. anatostana, 59 mm, INHS 77855. M) C. pyrrhomelas, 73 mm, INHS 76978. N) C. xaenura, 71 mm, KU 18994. O) C. gibbsi, 65 mm, KU 18892. P) C. caerulea, 58 mm, KU 18978. Q) C. nivea, 60 mm, KU 18987. R) C. callitaenia, 59 mm, TU 92770. Horizontal bar equals 1 mm. analostana. Other species have a very narrow maxilla anteriorly, in members of the lutrensis isthmus. Included here are gibbsi, trichroistia, clade, except proserpina, chloristia, galac- caerulea,whipplei,panarcys,andbocagrande. tura, and caerulea. All other species have a tall Other Cyprinella have an isthmus of interme- and relatively broad process, especially nivea, diate depth, like outgroups. callisema, callitaenia, and leedsi. The premaxillary process of the maxilla is The rostral process of the maxilla is a heavy, relatively short, about half the height of the fingerlike projection, generally directed an- NORTH AMERICAN MINNOW GENUS CYPR/NELLA 147

teroventrally. In some species, however, it is continued by Meckel's cartilage. may be quite heavy and then L-shaped, first di- Systematic variation in the morphology of rected ventrally and then bending mesially. the dentary includes depth and thickness of the Species with the L-shaped process are those gnathic ramus, orientation of the coronoid with the most inferior mouths, nivea, leedsi, process, curvature of the anterior gnathic ramus, callisema, callitaenia, and rutila, xanthicara, and the development of a dorsal shelf. Most panarcys, and proserpina. The latter four species of Cyprinella have a short, dorsally species also have a more robust rostral process directed, and relatively broad coronoid proc- than other Cyprinella. ess. Cyprinella trichroistia, gibbsi, pyrrhomelas, and xaenura differ in having a Lower Jaw more posteriorly directed process. Cyprinella The lower jaw is composed of eight ele- lepida, proserpina, panarcys, and xanthicara ments (Figs. 73, 74). Two are endochondral are divergent in that the process is thin and ossifications, the retroarticular and men- more elongate, less so in the last species. The tomeckelian, two are dermal, the dentary and posterior margin of the gnathic ramus is broad sesamoid articular, and one is a compound in Cyprinella, but is even more so in ornata, bone, the anguloarticular. This element is lutrensis, and garmani. These species have a composed of the dermal angular, endochon- very deep and heavy posterior aspect of the dral articular, and ossified portion of Meckel's dentary. cartilage, which extends from the mentomecke- All species have some development of a lian to the anguloarticular on the mesial sur- dorsolateral shelf anterior to the coronoid face of the dentary. process, but in some the shelf is more strongly Dentary (DE, Figs. 68, 69, 73, 74). This developed than in others. In the species pair paired element is the stoutest of all lower jaw chloristia and analostana and all members of ossifications and is most obviously curved the lutrensis clade the shelf is well developed mesially, especially anteriorly, where it articu- and wide. Within this latter clade, lutrensis has lates via a symphysial joint to its complement. the least development and in ornata, rutila, Posteriorly, it is broad and develops the coro- xanthicara,panarcys, andproserpina the shelf noid process (CP), a dorsal extension of the is very strongly developed. The anterior cur- gnathic ramus. The posterior margin of the vature and thickness of the gnathic ramus is gnathic ramus is broad and sloped posteriorly, diagnostic for some members of the genus. In articulating with the anguloarticular and most species the orientation of the gnathic retroarticular. Typically, the coronoid process ramus is nearly vertical, except callistia, leedsi, in these species is directed dorsally. Laterally, nivea, callitaenia, callisema, rutila, and xan-

the surface is smooth and rounded with a thicara. In these taxa the anterior tip is curved mesially sloping ventral surface. Fused to the mesoventrally, producing a more horizontal lateral face of the gnathic ramus is the anterior orientation. In species of the lutrensis clade the element of the mandibular canal. Dorsally, anterior gnathic ramus is deep and thick, more anterior to the coronoid process, there may be so than in other members of the genus. a lateral shelf formed and a slight depression Anguloarticular (ART, Figs. 73, 74). This ventrally. ossification is the second largest of the lower Mesially, the dentary is largely concave, jaw elements. Posteriorly, it is stout, generally especially ventrally. Dorsally there may be a deep, and articulates with the quadrate. Anteri- weak to strongly developed shelf. Anteriorly it orly the shaft gradually narrows to a point and is expanded at the joint with its complement is located between the mesial surface of the and curves mesially. Posterior to this joint the dentary and lateral surface of Meckelian carti- mentomeckelian is visible as a thin rod pointed lage. Articulation with the quadrate is possible anteriorly and truncating posteriorly where it through a partially contained socket formed by 148 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ART

Fig. 73. Lower jaws of species of Cyprinella and outgroups. A) Notropis atherinoides, 55 mm, KU 18935. B)Ly(hrurus fumeus, 55 mm, KU 6244. C) Luxilus cardinalis, 60 mm, KU 15281. D) Cyprinella ornata, 55 mm, KU 8405. E) C. lutrensis, 55 mm, KU 19431. F) C.formosa, 47 mm, KU 8399. G) C. lepida, 50 mm, TU 55189. H) C. proserpina, 58 mm, TNHC 3262. Left is lateral view and right is mesial view. Horizontal bar equals 1 mm. a posterior broadening of the bone and the de- or oval to L-shaped. In all species the bone is velopment of a posteriorly directed process short and oval except panarcys, proserpina, ventrally, into which fits the anteriorly di- and venusta in which it is long and slender. In rected and heavy process of the quadrate. these taxa, plus lepida, xanthicara, rutila, and Fused to the ventrolateral surface of the an- galactura it is roughly L-shaped. guloarticular is a short portion of the mandibu- Sesamoid articular (SA, Figs. 73, 74). lar canal. This ornate and thin bone is complex and lies In most species the dorsal surface of the an- partially above and partially lateral to Meckel's guloarticular is expanded, oblique, and di- cartilage and mesial to the anguloarticular. A rected dorsally. Here it articulates with the posterior process is developed above the coronoid process as well as the posterior gnathic ossified Meckelian portion of the anguloar- ramus. In panarcys,formosa, leedsi, whipplei, ticular and may partially overlap it mesially. and to a certain degree xaenura, this crest is Anteriorly, at least two, sometimes three, absent and the dorsal surface is nearly horizon- pointed processes are developed. Centrally a tal. more mesial process is developed for the inser- Retroarticular (RA, Figs. 73, 74). The tion of the adductor mandibulae A3. In all spe- retroarticular is generally short and robust and cies, except panarcys and proserpina, this located ventral to the anguloarticular and pos- central process is directed anteriorly and elon- teroventral to thedentary. Posteriorly, it articu- gate. These two species share a very reduced lates via a quasi-ball and socket joint with the and dorsally directed central process. anguloarticular and anteriorly with the den- Mentomeckelian (M, Figs. 73, 74). This tary by a ligament. It is connected to the ossification represents the anterior extent of anterior tip of the interopercle and posterior Meckel's cartilage and is fused to the mesial ceratohyal by ligaments. surface of the anterior gnathic ramus of the The shape of the retroarticular varies from dentary. In leedsi, callitaenia, callisema, being relatively short to elongate and straight pyrrhomelas, camura, analostana, chloristia, NORTH AMERICAN MINNOW GENUS CYPRINELLA 149

Fig. 74. Lower jaws of some species of Cyprinella. I) C. rutila, 57 mm, ASU 5982. J) C. camura, 45 mm, KU 15792. K) C. whipplei, 55 mm, KU 14211. L) C. analostana, 61 mm, INHS 77855. M) C. gibbsi, 57 mm, KU 18892. N) C. callistia, 60 mm, KU 18848. O) C. nivea, 59 mm, KU 18987. P) C. leedsi, 55 mm, KU 18985. Left is lateral view and right is mesial view. Horizontal bar equals 1 mm.

proserpina, panarcys, xanthicara, and boca- located on either side of the hyoid region. The grande the mentomeckelian is very short, posterior ceratohyal (PCH) is deep anteriorly represented only by a small cone. In other and narrows posteriorly to a point where it Cyprinella it is elongate and thin. articulates with the interhyal. The anterior ceratohyal (ACH) is the larger of the two; Hyoid Region deepest posteriorly where it articulates with Ten bones form the hyoid region. Included the posterior element and narrows anterome- are the paired interhyals, anterior and posterior sially where it twists and forks into two stout ceratohyals, dorsal and ventral hypohyals, the processes, one directed dorsomesially and one three branchiostegals, and the unpaired urohyal directed ventrolaterally. The dorsal and ven- and basihyal. All elements are cartilage, ex- tral processes of the ceratohyal articulate with cept the urohyal. The urohyal has a complex the dorsal and ventral hypohyals, respectively. origin. Following Patterson (1977) it may Both the anterior and posterior ceratohyals represent a compound bone, the median verti- contain foramina. The posterior ceratohyal cal plate being membrane and the horizontal contains two, generally located along the dor- ventral plate being dermal. These elements sal surface, through which passes the hyoideal help to support and form part of the branchial artery. The anterior ceratohyal, together with chamber. the hypohyals, forms the hypohyal foramen. Interhyal (IH, Fig. 75). This endochondral This foramen is generally bounded by all three element articulates with the posterior cera- elements. In some species, leedsi, callisema, tohyal, where it fits into a notch located pos- and callitaenia, a second foramen may exist in terodorsally. Dorsally it is included in the three the anterior ceratohyal ventrally just before element hyomandibular-symplectic joint. All the forked neck. Cyprinella have a short, stout interhyal. Within Cyprinella systematic variation in Ceratohyal (CH, Figs. 75, 76). The cera- ceratohyal morphology is expressed in depth tohyals are paired, endochondral elements of elements, location of the posterior cera- 150 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tohyal foramina, location and size of the hy- analostana, chloristia, whipplei, and spilop- pohyal foramen, and neck size of the anterior tera have very deep elements (especially those ceratohyal. Most species have relatively slen- of the lutrensis clade and analostana). Most der ceratohyals. Cyprinella garmani, lutrensis, species have relatively similar sized ceratohy- lepida, proserpina, panarcys, rutila, als at a similar body size. Cyprinella boca-

VHH PCH ACH

3=3 *

Fig. 75. Interhyals, ceratohyals, and hypohyals of species of Cyprinella and close relatives. A) Notropis atherinoides, 53mm, KU 18935. B) Lythrurusumbratilis, 59 mm.KU 15521. C) Luxiluscardinalis, S2mm,KU 15281. D) Cyprinella garmani, 50 mm, KU 5416. E) C. bocagrande, 32 mm, KU 20399. F) C. lepida, 50 mm, TU 55189. G) C. proserpina,

58mm,TNHC3262. H) C. rutila, 50 mm, ASU 5982. I) C.xanthicara, 43 mm, ASU 3642. J) C. chloristia, 55 mm, KU 8882. K) C. analostana, 56 mm, TNHS 77855. L) C. venusta, 78 mm, KU 8810. M) C. trichroistia, 60 mm, KU 18853. N) C. gibbsi, 58 mm, KU 18892. O) C. callisema, 53 mm, KU 8842. P) C. callitaenia, 58 mm, TU 92770. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 151 grande is an exception and has a more elongate genus and in the outgroups the dorsal and ceratohyal, correlated with its enlarged head (a ventral elements are subequal in size. In rutila 32 mm SL specimen of this species has cera- and xanthicara and species in \hcpyrrhomelas tohyals equal in length to a 43 mm formosa). and nivea species groups the dorsal element is Most species have thick and relatively strong smaller than the ventral. necks on the anterior ceratohyal. Cyprinella Basihyal (BHY, Figs. 77, 78). In most panarcys, proserpina, and rutila have a broad Cyprinella the basihyal is a relatively long neck. Cyprinella callisema, callitaenia, leedsi, bone. It lies at the anterior end of the branchial and xanthicara all have very narrow anterior region and extends forward to near the tip of necks on the anterior ceratohyal. the mouth. Posterodorsally the basihyal is Most Cyprinella have the foramina on the connected to the anterior end of the first ba- posterior ceratohyal located dorsally, along sibranchial. Posteroventrally a ligament ex- the top of the thickened dorsal margin as found tends from the center of the basihyal to the in outgroups. Some species, however, have the anteroventral edge of the first basibranchial posterior foramen located mesially and one (Fig. 77). species, ornata, has a lateral anterior foramen. As viewed dorsally, the basihyal is gener- Species with a mesoposterior foramen include ally rodlike to weakly triangular, with the ornata, analostana, chloristia, spiloptera, and widest aspect anteriorly and narrowing gradu- leedsi. ally to near its posterior extent where it is then Location and size of the hypohyal foramen more expanded (Fig. 78). The shape of this varies interspecifically. Some species have the bone is variable. Within the same species the foramen confined to the anterior ceratohyal, shape of the basihyal may range from narrow while in others it is equally bounded by the and cylindrical to a more triangular shape. anterior ceratohyal and both hypohyals. Adults Laterally, the basihyal is straight to slightly- of xanthicara, rutila, proserpina, panarcys, arched upward (Fig. 77). Contrary to observa- and venusta have the hypohyal foramen re- tions by Coburn (1982), most Cyprinella stricted to the anterior ceratohyal. In these taxa develop a process posteroventrally on the the foramen is restricted by the expansion and basihyal where there is ligamentous attach- condition of the dorsal and ventral processes ment to the first basibranchial (Fig. 77). In anteromedially or as a result of the foramen most species and individuals there are two passing through the shaft of the bone (e.g. concavities midlaterally (one per side) for the xanthicara). In some individuals of rutila the attachment of the ligament extending to the foramen is also bounded by both hypohyals. ventral hypohyal. These cavities are generally The hypohyal foramen is quite large in most fairly small except in lutrensis, garmani, species. In the caerulea species complex the whipplei, chloristia, and analostana where foramen is a narrow slit and in galactura, they are large (Fig. 77). Additional variation lutrensis, and ornata, it is very small and includes the relative depth of the basihyal. All generally confined anteriorly. species have basihyals with a similar narrow Hypohyal (HH, Figs. 75, 76). These paired depth except for whipplei, lutrensis, garmani, cartilage bones articulate with one another and lepida, andformosa that have noticably deeper each with the anterior ceratohyals. The dorsal basihyals (Fig. 77). In the former three the hypohyal (DHH) also articulates with the increased depth is attained by enlarging the basihyal posteriorly via a ligamentous con- centrally located keel. In formosa the entire nection. The ventral hypohyal (VHH) is more basihyal is more robust (Fig. 77). ornate, developing tubercle scars from liga- The deep basihyal morphology in whipplei, ments connecting the right and left elements, lutrensis, garmani, lepida, and formosa is and connecting each element to the anterior considered derived because outgroups typi- urohyal and basihyal. In most members of the cally have the narrow development like other 152 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ACH PCH

and atherinoides 53 1 8935. Fig. 76. Branchiostegal rays of species of Cyprinella outgroups. A) Notropis , mm, KU B) Lythrurusfumeus, 57 mm, KU 6244. C) Luxilus cardinalis, 82 mm, KU 15281. D) C. garmani, 50 mm, KU 5416. E) C. formosa, 43 mm, KU 8399. F) C. proserpina, 58 mm, TNIIC 3262. G) C. rutila, 50 mm, ASU 5982. H) C. whipplei, 63 mm, KU 14211. I)C. venusta, 78mm, KU 8810. J)C./nc/irow/w,60mm,KU 18853. K) C.caerulea, 58 mm, KU 18978. L) C. pyrrhomelas, 73 mm, INHS 76978. Horizontal bar equals 1 mm. NORTH AMERICAN MINNOW GENUS CYPRINELLA 153 members of the genus. Because the increased ment of greatly enlarged midlateral cavities on depth in these species appears to be due to the ventral surface of the basihyal is derived. different factors, the homologies of this char- Outgroups examined typically lack these cavi- acter are uncertain. Further, because only ties or, if present, they are small. juveniles of bocagrande, the presumed closest Urohyal (UO, Figs. 77, 79). Posteriorly relative offormosa, are available, it is uncer- this single median dermal bone is composed tain as to whether this species would develop largely of a vertical plate medially and a hori- a deep basihyal morphology. The develop- zontal plate ventrally. Anteriorly these thin

UOL

Fig. 77. Basihyals of some species of Cyprinella and close relatives. A) Lythrurus Urns, 45 mm, KU 18933. B)L.fumeus, 55 mm, KU 6244. C) Luxilus zonistius, 66 mm, KU 18995. D) L. zonatus, 85 mm, KU 12648. E) Cyprinella garmani, 50mm, KU 5416. F) C.lutrensis, 53 mm, KU 19431. G) C.formosa, 43 mm, KU 8399. H)C./epida,49mm,TU55189. I) C. proserpina, 58 mm, TNHC 3262. J) C. xanthicara, 43 mm, ASU 3642. K) C. whipplei, 55 mm, KU 1421 1. L) C. analostana, 60 mm, YNnSll%55. M) C. chloristia, 55 mm, KU 8882. N) C.galactura, 68 mm, KU 12029. 0)C.xaenura, 71 mm, KU 18994. P) C. nivea, 50 mm, KU 18987. Q) C. callisema, 56 mm, KU 8842. R) C. callitaenia, 58 mm, TU 92770. Horizontal bar equals 1 mm. 154 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

plates merge into a heavy rounded neck which callisema, and callitaenia have a notch that is bifurcates into two anteriorly directed process- deeply incised (Fig. 77). ess, separated from each other by a well- Within Cyprinella, the narrowly developed developed notch. The anterior face of each ventral horizontal plate is derived. Species of process is variably enlarged and serves as a Luxilus and Lythrurus have broad horizontal point of attachment for the ligament extending plates like callistia and species of the lutrensis from this bone to the ventral hypohyals (Fig. clade {xanthicara and rutila are exceptions). 77). The posterior margin of the ventral hori- Thus, the narrow ventral plate supports the zontal plate serves as a point of insertion for monophyly of the whipplei clade and the rutila the sternohyoides muscle. species pair. Independent reversals have ap- Variation in urohyal morphology includes parently occurred in whipplei, gibbsi, and cal- width of horizontal plate, height of vertical listia. The reversed conditions in these species plate, depth of anterior notch, and posterior is not considered homologous since their struc- margin of horizontal plate. Species with a tures are quite different (Fig. 77). Cyprinella broadly developed horizontal plate include whipplei and callistia have urohyals like out- lutrensis, garmani, lepida, proserpina, groups and gibbsi has a uniquly shaped urohyal. panarcys, some rutila (ASU 5982), whipplei, The deep vertical plate of the urohyal is and callistia. All other Cyprinella have a nar- derived in Cyprinella. Oulgroxvps, Luxilus and rowly developed ventral plate (Fig. 79). The Lythrurus, have the narrowly developed verti- vertical plate is deep in callistia and all species cal plate like all other members of Cyprinella. of the lutrensis clade, with the possible excep- Thus, the deep morphology supports the tion of xanthicara which has a slightly more monophyly of the lutrensis clade and is apo- narrow condition (Fig. 77). In all other species morphic for callistia. the vertical plate has a low profile (Fig. 77). The presence of a shallow and concave The posterior margin of the horizontal plate posterior notch in the horizontal plate of the shows interspecific but little intraspecific vari- urohyal is primitive within Cyprinella. Two ation. All species, except ornata, have either a independent modifications of this condition forked or relatively straight posterior margin have occurred.. First, ornata has an ovoid to the horizontal plate (Fig. 77). In ornata the urohyal and thus a convex posterior margin. posterior margin is ovoid (Fig. 77). Most spe- This condition is apomorphic for this species. cies of Cyprinella have a shallowly developed Second, species of the nivea and pyrrhomelas notch with moderately developed posterolateral species groups have an almost straight poste- processess extending beyond the posterior rior margin. This morphology is also derived extent of the vertically oriented plate for inser- and supports the sister relationship of these tion of the sternohyoides. Cyprinellaformosa two species groups, if one considers the condi- and bocagrande have strongly developed tion in callistia to be a reversal. The posterolateral processess. Cyprinella xaenura, pyrrhomelas species group is also derived in pyrrhomelas, gibbsi, trichroistia, caerulea, and lacking well-developed posteriorly directed species of the nivea group (except callistia) processes for the attachment of the sternohyoi- have reduced posterolateral projections and an des. Informosa and bocagrande, the posterior almost straight posterior margin of the urohyal processes for sternohyoides attachment are (Fig. 77). Cyprinella callistia has a deep notch enlarged and derived. and large projections. Development of a deeply incised anterior Two conditions of the anterior notch sepa- notch separating the stout anterior arms of the rating the stout anterior arms of the urohyal urohyal in the pyrrhomelas and nivea species exist in Cyprinella. Most species have a shal- groups is derived within the genus. Outgroups low notch. Cyprinella xaenura, pyrrhomelas, have a shallow notch like other Cyprinella. trichroistia, gibbsi, caerulea, nivea, leedsi, Branchiostegals(BS, Fig. 76). These three NORTH AMERICAN MINNOW GENUS CYPRINELLA 155

DHH

78. Dorsal view of Fig. ventral portions of gill arches of some species of Cyprinella and close relatives. A) Lythrwus fumeus, 56 mm, KU 6244. B) Luxilus cornutus, 125 mm, KU 8686. C) L. zonatus, 88 mm, KU 12648. D) Cyprinella orna/a,34mm,KU8405. E) C. lutrensis, 54mm, KU 19431. F)C. lepida, 54mm, TU 55189. G)C.proserpina, 47 mm, 3262. C. TNHC H) rutila, 50 mm, ASU 5982. I) C. camura, 61 mm, KU 19715. J) C. spiloptera, 68 mm, KU 11356. K) C. analostana, 56 mm, INHS 77855. L) C. venusta, 56 mm, KU 8810. M) C. pyrrhomelas, 73 mm, INHS 76978. N) C.siMwi,58mm,KU 18892. O) C.nivea, 60 mm, KU 18987. P) C. callitaenia, 58 mm, TU 92770. Horizontal bar equals 1 mm. 156 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 79. Ventral (left) and lateral (right) view of urohyals for species of Cyprinella and outgroups. A) Luxilus 12654. 56 6244. 34 KU 8405. chrysocephalus, 80 mm, KU B) Lythrurus fumeus , mm, KU C) Cyprinella ornata, mm, D)C.xan//Kcara,43mm,ASU3642. E) C.garmani, 50 mm, KU 5416. F) C.formosa, 43 mm, KU 8399. G)C.callistia, 78 mm, KU 18848. H) C. chloristia, 55mm, KU 8882. l)C.xaenuraJ\ mm.KU 18994. J) C.gibbsi, 5$ mm.KU 18892. K) C. callisema, 56 mm, KU 8842. Horizontal bar equals 1 mm.

flattened dermal bones articulate anteriorly tral edge of the ceratohyal. The second is with the ceratohyals. The first two are attached pressed against the lateral surface. The third is to the anterior ceratohyal. Branchiostegal 1 is connected to the lateral surface, between the attached by an anterior process, fitting into a anterior and posterior ceratohyals. All three small notch in the lateral face of the anteroven- are similar in shape, most often lunate. The NORTH AMERICAN MINNOW GENUS CYPR/NELLA 157 first is the narrowest and is strongly bent in and semi-oval to tear-shaped, lying partially adults, producing a mesially directed, right beneath and articulating posteriorly with the angle process. The second and third are less fused second and third compound element. sharply bent, being moderately curved. Laterally, the first articulates with the mesial Very little variation occurs in Cyprinella. end of the first epibranchial and articulates via Most species have branchiostegals of similar a cartilage cap. Epibranchial 2 also has a length. Cyprinella pyrrhomelas and xaenura cartilage connection with the posterolateral have short branchiostegals. Branchiostegals 2 face ofpharyngobranchial 2. The fused second and 3 are expanded and broad in the lutrensis and third pharyngobranchial is more elongate clade (except xanthicara), while narrow in and lunate in shape. This element articulates other taxa. The anterior neck of the first ele- laterally with the mesial ends of epibranchials ment is narrow and pointed in panarcys, 2 and 3. The cartilageous fourth element ar- proserpina, rutila, and xanthicara, and broad ticulates with the posterior margin of pharyn- and expanded anteriorly in other species. gobranchial 2 and 3 and the medial cartilage Cyprinella trichroistia has an unusually thin tip of epibranchial 4. first element, reduced posteriorly and pointed. Variation of the pharyngobranchials was The short, broad, and narrow neck condi- minimal and restricted to minor shape differ- tions of the branchiostegals are derived in ences. Most Cyprinella have pharyngobran- Cyprinella. Species of Luxilus, Lythrurus, and chials like closest outgroup relatives Luxilus most Notropis have narrow branchiostegals and Lythrurus. The first element is typically with a broad anterior neck where they articu- tear-shaped with a broad and rounded anterior late with the ceratohyals. Most of these taxa end that narrows posteriorly (Fig. 80). Dor- also have long branchiostegals like most sally, the posterior half ofpharynogbranchial 1 Cyprinella. has an obvious recessed region where the fused second element seats. In analostana and Branchial Region chloristia the anterior margin does not make a The bones of the branchial series form the smooth curve, but has a laterally directed ante- gill arches. Included are the pharyngobranchi- rior tip, giving the bone a more sinuous profile als, epibranchials, ceratobranchials, hypobran- (Fig. 80). Cyprinella leedsi and nivea both chials, and basibranchials (Figs. 78, 80). All of differ from other species in having a more the elements are cartilage and paired, except elongate and narrow first element (Fig. 80). the basibranchials, which are ventral, medial, Pharyngobranchial 2 is a more elongate and unpaired. The dorsal gill arch elements element and shows more interspecific vari- (epibranchials and pharyngobranchials) are ation. In most species, likeoutgroups, the bone suspended from the ventral neurocranium and is lunate with narrow posterior and anterior are connected to the ventral gill arch elements tips and a broad medial region. The dorsal (ceratobranchials) laterally. Gill rakers are of surface of all species except members of the dermal origin and are associated with all paired nivea group differes from species of Luxilus elements. and Lythrurus in having considerably more Pharyngobranchials (PBRl^, Fig. 80). topographic differentiation. Members of the These paired, robust endochondral bones ar- nivea species group as well as the above ticulate end to end with each other medially mentioned outgroups, except Luxilus pilsbryi, and laterally with all epibranchials. Only the have relatively smooth dorsal surfaces on the first three are ossified, the fourth remaining fused pharyngobranchial 2 and 3 (Fig. 80). cartilage, and the second and third become Species of the nivea group also vary from other fused (Hubendick, 1942). Thus, as adults there members in having more elongate elements are two ossified anterior elements and one (Fig. 80). In bocagrande andformosa pharyn- posterior cartilage element. The first is robust gobranchial 2 is more narrow and straightened 158 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

(Fig. 80). In panarcys, proserpina, rutila, and forming a ridge posteriorly on the neck (Fig. xanthicara the lateral edge of this element is 80) and creating a shelf. The anterior margin of expanded into a well-developed shelf (Fig. the first element is moderately straight in or- 80). Cyprinella lepida also develops a shelf nata, has a high anteriorly directed ridge in laterally, but in this species it is not as exten- bocagrande, formosa, camura, spiloptera, sive as in other members of the lepida species whipplei, analostana, chloristia, galactura, group (Fig. 80) and venusta, or has a medial ridge as in all Epibranchials (EB1-4, Fig. 80). All four other species. epibranchials articulate with their respective Like all outgroups, except for Luxilus ceratobranchials laterally and with the two pilsbryi, L. cardinalis, and L. zonatus, the pharyngobranchials medially. Epibranchials mesial neck of epibranchial 1 in most Cyp- 1-3 are approximately the same size, but are rinella is straight. Species of the lutrensis relatively thin and grooved dorsally for the clade, except for xanthicara and some rutila branchial arteries as in the ceratobranchials. (ASU 5982), chloristia, venusta, trichroistia, Mesially where they articulate with the phar- and gibbsi all have a posteriorly directed neck. yngobranchials they are thickened and robust. In xanthicara and some rutila the neck is The anterior margin of the first epibranchial is directed anteriorly. Generally, the expanded in slightly angled in most species with the high- mesial neck and the lateral body are oriented est point mesially to slightly anteromesially. the same plane. In analostana and chloristia, Posteriorly, its margin is relatively straight however, the head is rotated about 45 degrees lateral to the uncinate process. Mesial to the from the plane. uncinate process it is restricted medially and In overall shape the second epibranchial is develops a thickened neck. Epibranchials 2 similar to the first. Generally, the uncinate and 3 are more flattened relative to the first and process is larger and the mesial neck is less slightly wider, but are like the first except for pronounced. In some species the neck is straight a broader neck and wider head. The posteriorly like all close outgroups exceptfumeus, and in directed uncinate processes of the second and others it is directed anteriorly. Species with the third elements are also more strongly devel- anteriorly directed mesial neck includes oped, the third being the largest of the three. panarcys, proserpina, xanthicara, chloristia, Epibranchial 4 is the most distinctive of the analostana, pyrrhomelas,xaenura, galactura, four. The shaft of this element is more slender trichroistia, gibbsi, caerulea, nivea, callis- and heavier than others, the uncinate process is ema, and callitaenia (Fig. 80). In trichroisia, heavier and more laterally placed, and a dor- gibbsi, xaenura, pyrrhomelas, nivea, and cal- sally directed process is developed mesial to lisema the neck is more pronounced, having a the attachment point of the uncinate process of well -developed posterior notch mesial to the epibranchial 3. uncinate process (Fig. 80). Variation of epibranchials is expressed Other variants of epibranchial 2 include the mainly in overall shape. At a given body size, location of the connection of the ligament however, the overall size of epibranchials is between the uncinate process of epibranchial 1 similar. The first element varied with respect to and the anterior margin of epibranchial 2. In shape of the anterior margin, orientation of the most Cyprinella and all outgroups the connec- mesial neck, and connection of the flattened tion to the second element is anterior to the posterolateral plate to the thickened mesial uncinate process of the same bone. In nivea neck. In all Cyprinella except bocagrande and and leedsi the connection is lateral to this point formosa, as well as outgroups, the posterior (Fig. 80). plate connects and is continuous with the mesial Little variation of systematic importance neck ventrally (Fig. 80). In these two species, was noted in epibranchial 3. Cyprinella connection of the plate is more dorsally placed, proserpina, rutila, and xanthicara have an NORTH AMERICAN MINNOW GENUS CYPRINELLA 159

80. and of of and close relatives. atherinoides Fig. Epibranchials pharyngobranchials species Cyprinella A) Notropis , 53 L. mm, KU 18935. B) Lythrurusfumeus , 56 mm, KU 6244. C) ardens, 72 mm, KU 4136. D) Luxilus zonatus, 67 mm, KU 12648. E) L. chrysocephalus, 80 mm, KU 12654. F) Cyprinella ornata, 44 mm, KU 8405. G) C. lutrensis, 55 mm, KU 19431. H) C.formosa, 43 mm, KU 8399. I) C. bocagrande, 32 mm, KU 20399. J) C. lepida, 48 mm, TU 55189. K) C.panarcys, 42 mm, UMMZ 208212. L) C. rutila, 52 mm, ASU 30264. M) C. xanthicara, 42 mm, ASU 3642. N) C. camura, 61 mm.KU 19715. 0)C.a/w/cw/amj,60mm,INHS 77855. P)C.cWorty/j'a,55mm,KU8882. Q)C.pyrrhomelas, 73mm,INHS76978. R)C.,giMw,58mm,KU 18892. S) C.nivea. 62mm, KU 18987. T) C. callisema, 53 mm, KU 8842. Stippled regions represent cartilage. Horizontal bar equals 1 mm. anteriorly directed mesial neck, like the sec- similar in all species, like most other cyp- ond epibranchial of other Cyprinella (Fig. 80). rinids, except in xaenura and pyrrhomelas. Most species have a moderately restricted neck These species have a very narrow condition like other cyprinids examined. In lutrensis, (Fig. 80). The dorsally directed process poste- bocagrande, lepida, panarcys, callistia, and rior to the uncinate process of the third epibran- callitaenia the neck is more broadly devel- chial is consistently developed in all species oped and in xanthicara, xaenura,pyrrhome las, except members of the lutrensis clade (Fig. trichroistia, and gibbsi the neck is narrow 80). In this group, development of this process (Fig. 80). is very spotty, some individuals have no proc- Variation in the fourth epibranchial includes ess and others have the process developed only width of the element, the degree of develop- on one side. Further, if present in this group, it ment of the dorsally directed mesial fingerlike is almost always small. process, and the length of the bone mesial to Mesial to the uncinate process of epibran- the uncinate process. The width of this bone is chial 4 the body of this bone is generally 160 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

elongate. The species ornata, lutrensis, placed insertion of the ligament on the anterior garmani, formosa, trichroistia, and callistia face of epibranchial 2 is also derived within all differ in having a shorter mesial portion to this genus. Close outgroups such as Luxilus this element (Fig. 80). and Lythrurus have a more mesially placed The dorsal connection of the posterior plate ligament. of epibranchial 1 to the mesial neck is derived The narrow epibranchial 4 is derived in in Cyprinella. In all other species and out- Cyprinella. In close outgroups this element is groups the connection is ventral and no shelf is similar to species with the broad morphology. formed. Having a straight neck on epibran- The weakly and variably developed dorsal chial 1 in Cyprinella is primitive for the group process of epibranchial 4 is unusual, but is since all outgroups, except Lwdlus zonatus, L. considered derived for the lutrensis clade. cardinalis, and L. pilsbryi, have a straight Generally, every individual of other taxa con- neck. Thus, the posteriorly directed neck is sistently develop this process. derived for the lutrensis, formosa, and lepida Ceratobranchials(CBRl-5,Fig.78).The species groups, chloristia, venusta, and the first four of the five ceratobranchials are thin gibbsi-trichroistia species pair. The anteriorly and ventrally concave. The first three articu-

directed tip of the neck in some rutila and late mesially with hypobranchials 1-3, while xanthicara is considered a further derivation the fourth articulates with the unossified cop- from the straight or posteriorly directed condi- ula extending from the posterior end of ba- tion. sibranchial 3 to the anterior arm of ceratobran-

Polarity of the ridge along the anterior chial 5. Each of these bones is tipped mesially margin of epibranchial 1 is difficult to deter- with an ovoid or rounded cartilage and later- mine. The conditions present in outgroups ally with a forked cartilaginous element. Lat- varies from having a medial to anterior ridge. erally the profile of these first four bones from All species oi Lythrurus have a medial ridge, their lateral articulation with epibranchials to except umbratilis, which has the anterior ridge. mesial hypobranchial articulation is slightly Within Luxilus, pilsbryi, cardinalis and zo- concave, the lowest aspect being medially. natus have a medial ridge and others develop The fifth pair of ceratobranchials represents the ridge anteriorly. The ridge in Notropis the heavy pharyngeal arches.

atherinoides and other Notropis (s.s.) is ante- Variation in morphology of ceratobranchi- rior. Thus, one interpretation of the evolution als \~A is primarily with respect to shape. of this characteristic is for an anterior condi- Some species also vary in relative lengths of tion to be primitive in the ancestor of Cyp- these bones. At similar body sizes lutrensis, rinella and a medial condition developed in camura, chloristia, pyrrhomelas, xaenura, the ancestor of Lythrums and in the pilsbryi- trichroistia, and gibbsi had longer ceratobran- zonatus species group of Luxilus. Within Cyp- chials than remaining members of the genus rinella, the medial morphology is derived for (Fig. 78). The elongate nature of these bones in the pyrrhomelas and nivea species groups, as these species was not correlated with overall well as in some species of the lutrensis clade. width. Cyprinella camura and lutrensis have Other species have retained the primitive ante- broader ceratobranchials like other members rior condition. of the genus, while the remaining species with The anteriorly directed and restricted neck the elongate morphology have narrow cera- of the second epibranchial is derived for Cyp- tobranchials. rinella. Outgroups, except Lythrurus fumeus, The shape of the first ceratobranchial is have a straight neck. The distribution of the roughly rectangular, although typically widest anteriorly projected neck of epibranchial 2 is medially, primarily from the expansion of the spotty, however, and requires several changes posterior margin. In some taxa a restricted in the evolution of this character. The laterally anteroventral neck is present before articula- NORTH AMERICAN MINNOW GENUS CYPRINELLA 161 tion with the first hypobranchial. In most spe- Determining the polarity of the variation in cies the anterior end is straight. The restricted the length of ceratobranchials \-A in species of neck is present in rutila, xanthicara, spilop- Cyprinella is difficult. All species, except tera, and species of the nivea species group lutrensis, camura, chloristia, pyrrhomelas, (Fig. 78). ]nxaenura,pyrrhomelas, trichroistia, xaenura, trichroistia, and gibbsi have short gibbsi, lutrensis, and camura, however, the ceratobranchials. Species oiLuxilus and some anterior tip is bent mesially towards the first members of Lythrums have long ceratobran- hypobranchial approximately 30-45° from the chials. One may interpret the short condition primary axis. In members of the nivea group of most Cyprinella to represent a derived and rutila the anterior margin of the first cera- condition for the genus and reversed to elon- tobranchial is expanded anteriorly, producing gate elements in the above species and species a wider medial region than in other species groups. This is a possibility since lutrensis and (Fig. 78). camura both have elongate ceratobranchials Morphology of the second ceratobranchial for their size, but they are also broad like other is similar to the first except that all species Cyprinella. Other taxa with the elongate con- excluding ornata (Fig. 78) have a narrowing dition have narrow ceratobranchials, different of the anterior end into a neck. In most taxa the from lutrensis and camura. narrowed neck is short, but in lepida,panarcys, The restricted neck of ceratobranchial 1 proserpina, rutila, xanthicara, camura, and appears to be derived for Cyprinella, but ap- trichroistia it represents up to one-half the pears to have been derived several times within total length. the genus. The restricted neck is evidence for In most species ceratobranchial 3 is a rela- the monophyly of the species pair rutila and tively straight bone with a small restricted xanthicara and for the nivea species group. It anterior neck. Cyprinella xanthicara and ru- also represents an autapomorphic condition tila are distinctive in having a rounded and for spiloptera. expanded anterolateral margin (Fig. 78), The bent anterior neck of the first cera- compared to a straight margin as is found in tobranchial is derived in Cyprinella. Species other members of the genus. These species of Luxilus and Lythrurus all have a straight plusproserpina andpanarcys also have a more neck and thus the bent neck supports the elongate anterior neck (Fig. 78). The posterior monophyly of the pyrrhomelas species group margin of ceratobranchial 3 in most Cyprinella and is apomorphic for camura and lutrensis. is smooth and straight except for a small pos- Having an expanded anterior margin on the teriorly directed process near the posterior end first ceratobranchial is also a derived condi- of the anterior neck. In analostana and chlo- tion. Species ofclose outgroups all have straight ristia this process is serrated (Fig. 78). anterior margins on ceratobranchial 1. Thus, Unlike the first three, the fourth ceratobran- this morphology supports the monophyly of chial articulates with the unossified copula, the nivea species group and is apomorphic for has an expanded anterior end, and is the short- rutila independently. est of the series of four. The typical morphol- An elongate and narrow anterior neck on ogy of this bone is to have the posterior two ceratobranchial 2 has been derived three times. thirds being relatively straight and the anteri- Close outgroups have a short, poorly devel- ormost third bent anteriorly. Similar to cera- oped and reduced neck like many members of tobranchial 3, many species have a smooth Cyprinella. Thus, the long and slender neck of process on the posterior margin near the ante- the second ceratobranchial is evidence for the rior end. Cyprinella pyrrhomelas, xaenura, monophyly of the lepida species group and is trichroistia, gibbsi, caerulea, callistia, nivea, apomorphic in trichroistia and camura. leedsi, callitaenia, and callisema have a very The rounded anterior margin of the third reduced process or none at all. ceratobranchial in rutila and xanthicara and 162 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the elongate neck in these species andpanarcys and third are generally equal in length, but and proserpina is derived within Cyprinella. longer than the first, and are cylindrical. The Outgroups and other species of Cyprinella third is connected posteriorly to a slender typically have straight anterior margins and unossified copula that bifurcates posteroven- weakly developed necks or no necks at all. trally and articulates with the anteroventral A posterior process on ceratobranchial 3 is arms of the fifth ceratobranchials. All three ba- common in Cyprinella and outgroups, but sibranchials articulate laterally with the hy- having a serrated edge is derived. This mor- pobranchials. phology supports the monophyly of the Variation in basibranchial morphology is analostana-chloristia species pair. minimal. In venusta and analostana the first Hypobranchials (HB1-3, Figs. 77, 78). basibranchial is typically longer than in other These three paired ossifications are lateral to species (Fig. 78). In some species the first the basibranchials and at the anteromesial ends basibranchial is hourglass in shape when of the first three ceratobranchials. The first and viewed dorsally, with the anterior and poste- second are heaviest and rounded or oblong rior ends being broadest and the smallest di- when viewed dorsally (Fig. 78) and both are ameter located medially. The constriction of more elongate dorsoventrally. The third is the basibranchial 1 is best developed in species of smallest dorsally and develops a thin, elongate the nivea group (except callisema), but was ventral process which curves anteromesially. apparent in trichroistia, gibbsi, callistia, In most species a single, dorsal cartilage head caerulea,xaenura, and pyrrhomelas (Fig. 78). is developed on all three pairs in juveniles and A more weakly developed hourglass shape adults. In some species, however, the first occurs in whipplei, analostana, chloristia, hypobranchials have the single dorsal carti- spiloptera, galactura, and venusta (Fig. 78). lage head divided by a median ossification In other species the first basibranchial has a producing two heads, one anteriorly and one uniform diameter and appeared cylindrical posteriorly. Species with this development (Fig. 78). include callisema, callistia, callitaenia, leedsi, The increasing development of an hour- and nivea (Fig. 78). The dorsal cartilage head glass-shaped first basibranchial is derived of hypobranchials 2 and 3 is single in all within Cyprinella. Cyprinella whipplei, species. analostana, chloristia, spiloptera, galactura, Development of a double cartilage head on and venusta have the derived condition, but hypobranchial 1 is derived within Cyprinella. the constriction is slight. Species of the nivea Outgroups, except for Luxilus zonistus, L. species group represent the extreme in devel- coccogenis, L. pilsbryi, L. cardinalis, and L. opment of this morphology. Outgroups all zonatus have a continuous dorsal cartilage have the cylindrical basibranchial 1 with no head. constriction.

Basibranchials (BB1-3, Figs. 77, 78). These three small, quasi rod-shaped perichon- Weberian Apparatus dral ossifications form the medial axis of the This osteological complex is probably the gill arch. They articulate end to end and anteri- most complicated series of bones in the cyp- orly with the posterior tip of the basihyal (Fig. rinid skeleton (Fig. 33). Composed of the pars 78). The first is always the smallest of the three auditum and par sustenaculum, the apparatus and in dorsal profile is cylindrical or hourglass is an otophysic connection between the ante- shaped. As viewed laterally, it is deepest pos- rior air bladder chamber and the inner ear. The teriorly and narrows anteriorly with an occa- par auditum is the series of four small ossicles sional ventral process developed where it re- interconnected by the interosseus ligament. ceives a ligament from the posteroventral These elements are used to transmit sound margin of the basihyal (Fig. 77). The second from the gas bladder to the ear. The pars NORTH AMERICAN MINNOW GENUS CYPRINELLA 163 sustenaculum includes the gas bladder and third neural arch. This complex is posterior to first four modified vertebrae and arches which the neural arch of centrum 2 and anterior to the act as support for the ossicles. Variation in neural arch and spine of centrum 4. It has an Cyprinella and other cyprinids examined was expanded and arched base which narrows essentially nonexistent. dorsally to a neck of variable width. Dorsally, Pars sustenaculum. The first vertebra of a divided V-shaped crest is formed and is the pars sustenaculum is thin and articulates larger than the base and median neck. Only anteriorly with the proatlas of the basioccipi- occasionally will the anteriormost predorsal tal. A short and stout transverse process is bone extend between the walls of this crest. produced laterally and is connected to the Large ligaments extend between the neural supracleithrum by a ligament. Because the complex and the supraoccipital crest and ligament may become ossified, the length of exoccipital. the transverse process is somewhat variable The neural crest, together with the neural intraspecifically. The movable scaphia and spine of centrum 4, are the only systematically claustra are located dorsolateral^ and dorsally variable structures noted in Cyprinella and on this centrum, respectively (Fig. 33). outgroups. Relative to close outgroups, all The second, third, and fourth centra are species of Cyprinella have an elevated neural increasingly larger than the first and are more complex and fourth neural spine. These struc- amphicoelous. Arising ventrolaterally from tures are extremely tall in most species of the the second centrum are two large and stout lutrensis clade (Fig. 33). The height of the transverse processes, one per side. Dorsally neural complex and fourth neural spine in the large, platelike second neural arch is sepa- rutila and xanthicara is like species of the rate from the centrum by the anterior extension whipplei clade, representing a reversed and of the third neural arch. The intercalaria os- derived condition for these two species. All sicles articulate laterally and dorsolateral^ on other members of the lutrensis clade, however, this centrum. are derived in having a very tall neural com- The third centrum is large and articulates plex (Fig. 33). with an enlarged neural arch which supports Pars auditum. Included here are the most of the dorsal neural complex. Laterally, Weberian ossicles, including the claustrum,

this centrum has a channel on each side for the scaphium, intercalarium, and tripus. These articulation of the large tripus. The neural arch four elements are modifications of the first to this element is broad at the base, constricted four vertebral elements and are ligamentously medially, and expanded anteriorly near the joined to transmit sound received by the gas dorsum. The anterior expansion separates the bladder to the inner ear. None of these ele-

second neural arch and centrum. ments vary in Cyprinella . Centrum four supports an enlarged neural The claustrum (CLA, Fig. 33) is above arch and elongate spine dorsally. Ventrolater- centrum 1 and covers the atrium sinus impar ally it supports the enlarged os suspensoria and ventromedially. Dorsally it is roughly triangu- enlarged parapophyses. The enlarged parapo- lar and ligamentously joined posteriorly to the physes extend laterally and slighdy anteriorly neural arch of centrum 2 and anteriorly to the from the centrum. The os suspensorium ex- exoccipital above the foramen magnum. tends posteriorly and lies against the tunica The scaphium (SCA, Fig. 33) is a small, externa of the swimbladder. This structure ornate and complex bone. It articulates later- an ossification of the 1 represents independent ally on centrum 2, covers part of centrum , third centrum that migrates posteriorly and and forms the lateral walls covering the atrium becomes fused to the parapophyses of centrum sinus impar. As described by Coburn (1982), it four (Niazi, 1964; Coburn, 1982). may be divided into four parts, a large and The neural complex is located above the mesially convex cup, a posterodorsally di- 164 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY rected spine (ascending process), a ventrome- and the prezygapophyses are absent or weakly dial articulating process, and a small, blunt developed. The total number of vertebrae, tubercle posteriorly off the lateral surface of inclusive of the first four Weberian elements, the cup for attachment to the interosseus liga- and the number of precaudal and caudal element. Ligaments connect the dorsal spine to ments are given for each species in the species the second and third neural arches. accounts section. The intercalarium (ICA, Fig. 33) is the The precaudal vertebrae are monomorphic. smallest ossicle and is roughly T-shaped. It is Each has a well-developed neural spine di- elongate and lies anteriorly on the lateral sur- rected posterodorsally and a prezygapophysis face of the second centrum, oriented anterov- directed anterodorsally. The prezygapophyses entral to posterodorsal. Anteroventrally, the are best developed anteriorly, extending al- tip of this bone is expanded into a knob and is most to the neural spine of the preceeding joined to the interosseus ligament. The articu- element and decrease in size posteriorly. Ver- lar process with centrum 2 is small and is tebra 5 has weakly developed prezygapophy- opposite the dorsoposterior ascending proc- ses. Posterior to the neural spine are the small ess. The ascending process is joined by a postzygapophyses. These processes are di- ligament to the third neural arch. rected dorsally or posterodorsally and have a The large, roughly triangularly shaped tripus moderately large foramen at the base. Ventro- (TRP, Fig. 33) is of compound origin, from a laterally, the ribs articulate with the parapo- parapophysis and pleural rib. The most obvi- physes. ous features of this bone are the broad mesial Between each of the neural arches and articular surface and an anterior and posterior between the first neural arch and the neural ramus. The parapophysis forms the medial complex are the predorsal bones. These el- triangular portion of the bone and the articular ments are thin and platelike ventrally. In some process; the pleural rib forms the large poste- they may be thickened dorsally and have a rior process and a portion of the body and ventrally directed, medially thickened shaft. anterior ramus. The lateral margin of the pos- In the whipplei clade and all close outgroups, terior process is closely associated with the except for very large individuals of Luxilus, peritoneal tunic of the anterior chamber of the these bones are relatively small, largest anteri- gas bladder. From the main body, this bone is orly and decreasing in size posteriorly to the curved mesially between the lateral process of dorsal fin pterygiophores. Generally, they are the fourth rib and the os suspensorium. Dis- absent or very small just before the dorsal fin.

tally, it tapers to a thin shaft and is attached to In the lutrensis clade, however, these bones are the tunic externa and to the posterodorsal sur- very large and sometimes ornate, and extend face of the os suspensorium via ligaments. The between the neural crest and the dorsal fin. anterior ramus is elongate and connected an- Caudal vertebrae are also generally mono- teriorly to the interosseus ligament. morphic. All processes of the precaudal vertebrae are present, but both the pre- and Vertebrae postzygapophyses are smaller. All caudal Posterior to the Weberian apparatus the vertebrae have an enclosed haemal arch and vertebrae are separated into caudal and pre- spine. The first element may have the arch caudal elements (Figs. 16, 33). Caudal verte- weakly developed. Each element has small brae develop haemal spines and arches, and haemal pre- and postzygapophyses, although lack parapophyses. Generally, there are two weakly developed anteriorly. Posteriorly, close transitional vertebrae. These elements have an to the caudal fin, the haemal and neural spines incomplete haemal arch fused to the centrum become increasingly bent posteriorly. and small, free ribs. On these vertebrae the The only variable characteristic of the ver- haemal postzygapophyses are well developed tebral elements and associated processes in NORTH AMERICAN MINNOW GENUS CYPRINELLA 165

Cyprinella is the length of the neural spines. bones, number of neural and haemal spines on Species of the lutrensis clade all have more precaudal vertebrae, and the degree of orna- elongate neural spines than members of the mentation in the form of median plates on the whipplei clade and most close outgroups. Large spines. All Cyprinella and most other cyp- adults of species of the cornutus species group rinids examined have a low frequency of indi- of Luxilus also have tall neural arches, but viduals with double or at least bifurcated neural these species are over 100 mm SL. Individuals and haemal arches (Fig. 16). Occassionally a of equal or larger size than members of the single individual may have one to two double lutrensis clade have smaller arches. haemal and neural arches. Except for ornata, the loss of a hypural is Caudal Skeleton and Fin variable in each species examined. Cyprinella Supports ornata is unique among all species in consis- The caudal skeleton and first few preural tently lacking the sixth hypural. As noted by centra of Cyprinella and other cyprinids ex- Buhan (1972) for Luxilus chrysocephalus, amined (Fig. 16) represents one of the most Notropis photogenis, and Rhinichthys atratu- conservative complexes analyzed. In basic lus, when a hypural element is missing it is structure the caudal skeleton of Cyprinella is invariably the sixth bone of the series that is similar to that described by Buhan (1972) for absent. Exoglossum laurae, Cyprinella many North American minnows. panarcys, C. proserpina, and C. whipplei each Articulating posteroventrally with the had a single individual with the sixth hypural compound centrum is the fused parhypural absent. and the first hypural element. The caudal vein Fusion of hypurals is not very common, but and artery passes between these two elements when present, it is variable with respect to and ventral to this opening is the hypurapo- which elements are involved. Cyprinella xan- physis (Fig. 16). Posteriorly, the compound thicara and venusta each have a single case of centrum is fused with the second hypural. fusion between hypurals 5 and 6, one venusta Dorsally it is fused with the first and second has hypurals 4 and 5 fused and one individual uroneurals, which form a compound uroneural of C. caerulea and Lythrurus fumeus has commonly referred to as the first uroneural. hypural 1 and the parhypural fused. Dorsal to the fused hypural 2 are hypurals 3-6. Neural and Haemal Arches and Spines Between 2 and 3 marks the division of the (NS, HS, NA, HA, Fig. 16). Ornamentation of upper and lower caudal lobes. Anterodorsally neural and haemal arches varies in a species to the sixth hypural and lateral to the fused first with age. Generally, younger specimens pos- and second uroneurals, lies the paired uroneu- sess simple arches and spines with no proc- ral 3. Anterior to this structure and directly esses, except for small neural and haemal pre- above the compound centrum is the single and postzygapophyes. With increasing size, epural. Directly anterior to the compound arches develop more elaborate median plates. centrum is preural centrum 1. This is not without exception as similar size My observations of the cyprinid caudal adults of a single species may have differing morphology reflect minimal variation of sys- amounts of ossification of these plates. No tematic interest. Generally, only minor inter- relevant systematic data could be gained in as well as intraspecific variation occurs in the this study from shape or size of these median caudal skeletons. Some characteristics, how- plates. ever, seem to be consistent within and between Epural (EP, Fig. 16). Variation of the epu- species and of relevence to the evolution of ral follows a similar pattern as that shown by Cyprinella. the preural vertebral arches and their associ- Within and outside Cyprinella intraspecific ated median plates. Generally, young-of-the- variation includes fusion or loss of hypural year have a thin rodlike epural extending from 166 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the tip of the fused uroneural 1 and 2 to and rinella condition. Development of a long closely associated with neural arch of preural uroneural 3 may be a derived condition of a centrum 1 . Increasing body size is accompa- larger group of eastern North American cyp- nied with a thickening of the shaft. The an- rinids, inclusive of Cyprinella. teroventral tip becomes expanded into a spa- Additional variation in the caudal skeleton thula or a single-barbed, harpoon-shape, with includes the overall shape of the caudal fin and the tip shape being somewhat variable in a associated supporting elements and preural species. Species of the lutrensis clade, how- vertebrae. Species of Cyprinella, with the ever, generally develop a heavier epural, rela- exception of venusta, galactura, callitaenia, tive to other Cyprinella and outgroups. callisema, leedsi, and nivea, have a caudal fin Length of the third uroneural, referred to as shape different from all other cyprinids exam- urodermal by Buhan (1972), varies systemati- ined. In this genus the overall shape of the fin cally within Cyprinella (Fig. 16D-F). As de- supports and neural and haemal arches is deeper tailed by Patterson (1968) urodermals are and more erect, giving the fin a broader fan dermal scale deri vitives and are present in only shape. The exceptions mentioned above have a few primitive teleosts. Uroneural is the cor- a slightly more narrow caudal skeleton, simi- rect nomenclature for these elements because lar to outgroups. Changes in the overall shape they are derived from ural neural arches. Us- of the caudal skeleton are not restricted to ing hypurals 4-6 as a reference for extent of Cyprinella. Erimystax dissimilis, E. insignis, anterior development, all Cyprinella show E. cahni, E. monacha, Phenacobius development of the third uroneural to a point catostomus, P. teretulus, and P. uranops all between the anterior extent of hypurals 5 and have a much compressed caudal and precaudal 6 or less. Cyprinella rutila, proserpina, and skeleton, relative to any Cyprinella and other panarcys have their third uroneurals extend- cyprinids surveyed. This condition as well as ing only to the anterior end of hypural 6 (Fig. the broadened morphology of Cyprinella is 1 6D). Although variable, 60% of the individu- considered derived from the primitive condi- als oilepida adults have the reduced condition. tion found in most other cyprinids and are Cyprinella rutila from Rio Salado de los indicative of monophyly of each group. Nadadores (ASU 63-0264) and xanthicara have uroneural 3 extending anteroventrally to Pectoral Girdle between hypurals 5 and 6. Cyprinella gibbsi, Posttemporal (PTT, Fig. 81). This thin, trichroistia, and caerulea also have a reduced spathulate dermal bone overlaps the suprac lei- third uroneural condition. In these species the thrum ventrally and is strongly connected to maximum size of their third uroneural is smaller the epiotic and dermopterotic dorsally. Dor- than hypural 6 (Fig. 16F). In most cases out- sally, the lateral surface contains part of the groups have the third uroneural extending to posttemporal head canal after the fusion of the hypural 5. Luxilus pilsbryi, L. cardinalis, L. posttemporal and supratemporal canals. Ven- zonatus,L. zonistius, Phenacobius catostomus, trally, the posttemporal forms much of the roof P. teretulus, and Hybopsis zanema have the and lateral face of the posttemporal fossa. element extending to between hypurals 5 and The shape of the posttemporal varies be- 6. The reduced condition of all Cyprinella may tween species of Cyprinella. Generally, two be considered derived for this group, with conditions exist. Species of the lutrensis clade further independent reduction of this condi- have a teardrop-shaped posttemporal. In or- tion in the lepida species group and caerulea nata and Iheformosa and lepida species groups species complex. The hypothesis of independ- it is broad and rounded ventrally and narrows ent reduction in these two clades seems rea- dorsally to a long, narrow, and pointed shaft. sonable since species most closely related to The shaft on ornata is broader and more blunt. each of these groups have the typical Cyp- The posttemporal of lutrensis and garmani is NORTH AMERICAN MINNOW GENUS CYPRfNELLA 167 less extreme. In species of the whipplei clade the posttemporal resembles the primitive morphology present in other genera examined.

Cleithrum (CL, Figs. 31, 82). The cleithrum is dermal and the largest of the pectoral girdle elements and forms most of the lateral face of the girdle. It is roughly L-shaped with a large ascending process and a slightly smaller horizontal branch directed anteriorly. Mesially, the ascending limb is flat posteriorly where the dorsally flattened supracleithrum is attached. Ventrally, the ascending limb is attached to a flattened and broad posterior process of the scapula. Anteriorly, there is a mesial ridge extending from the scapula, dorsally to the tip where the ascending limb terminates in a blunt spine. Slightly above the scapula the mesoco- 81. bone in of racoid is connected to the mesial ridge. Fig. Posttemporal species Cyprinella. A) C. camura, 45 mm, KU 15792. B) C. ornata, 56 mm, KU The horizontal process of the cleithrum is 8405. C)C./u/re/m.y,55mm,KU19431. D)C.formosa, connected to its At the complement anteriorly. 47 mm, KU 8399. E) C. bocagrande, 32 mm, KU 20399. symphasis the two form a roughly straight F) C. lepida, 50 mm, TU 55 1 89. G) C. rutila, 57 mm, ASU 5982. \JMML 208212. Horizontal margin in Cyprinella and not a shallow V as in K)C.panarcysMmm t bar equals 1 mm. other genera. The lateral face of the cleithrum is narrowest anteriorly and very broad posteriorly. Mesial to the lateral face a ridge is devel-

Fig. 82. Dorsal view of cleithrum of some species of Cyprinella and outgroups. A)Notropis telescopus, 67 mm, KU 15231. B) Lythrurus umbratilis, 56 mm, KU 15521. C) Luxilus cardinalis, 79 mm, KU 15281. D) C. analostana, 58 mm, INHS 74347. E) C. callistia, 78 mm, KU 18848. Horizontal bar below N. telescopus for all species except L. cardinalis. Horizontal bar equals 2 mm. 168 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY oped that extends posteriorly up the ascending limb. Mesial to the ridge the cleithrum is connected to the coracoid. Articulation with the coracoid is divided into anterior and poste- B rior segments, divided by a large foramen. The posterior connection is higher than the anterior connection, forming a shelf over and posterior to the latter. Other than the anterior margin of the cleithrum, apparently no consistent systematic variation in the morphology of the cleithrum occurs in the genus. Supracleithrum (SCL, Fig. 83). Connected to the posttemporal dorsally and cleithrum ventrally, this dermal element contains part of the continuation of the sensory canal leading from the posttemporal to the lateral line. It is flat and broadest medially and ventrally where the margin forms a broad point ventrally. Dorsally it narrows and may produce a hooked point. The morphology of the supracleithrum of these species is very similar. Cyprinella proserpina differs in having a very broad supracleithrum. Coracoid (COR, Fig. 31). As described under the cleithrum, these elements share a broad articular surface anterior and posterior to the large interosseus foramen. Mesially, the coracoid is slightly convex and slopes ven-

trally. Posteriorly, it is elevated and forms a base, along with the scapula, for the attach- Fig. 83. Supracleithrum bone in some species of ment of the mesocoracoid. Lateral to the Cyprinella. A) C. analostana, 61 mm, INHS 77855. B) C. galactura, 57 mm, KU 12029. C) C. callisema, 52 mesocoracoid-coracoid articulation, the cora- mm, KU 8842. D) C. proserpina, 58 mm, TNHC 3262. coid articulates with the broadly scapula. E) C. garmani, 50 mm, KU 5416. F) C.formosa, 47 mm, Posteriorly, the four radials are attached KU 8399. Horizontal bar equals 1 mm. ligamentously. The morphology of the coracoid is similar for all species of Cyprinella. The anterior and mesial margins are rounded; Mesocoracoid (MCO, Fig. 31). This ele- posteriorly and laterally it is flattened. Con- ment is a slender spine, broadest ventrally tained within the scapula is the scapular fora- where it articulates with the scapula and cora- men. The scapula may be expanded postero- coid. It extends dorsally from this base as a dorsally into a small, flat plate which is pressed pointed process and is attached to the mesial against the mesial surface of the cleithrum. ridge of the ascending arm of the cleithrum. Species of Cyprinella vary from most other Species of Cyprinella are essentially mono- cyprinids examined in the morphology of the morphic morphologically for this bone. scapula. In all close outgroups, except the Scapula (SCA, Fig. 82). This endochon- gznusLuxilus, a thin, flat, posterodorsal flange dral element articulates with the cleithrum is produced from the scapula which is attached laterally, coracoid mesially, and the first two to the mesial surface of the cleithrum. radials and first pectoral fin ray posteriorly. Cyprinella analostana, chloristia, leedsi, nivea, NORTH AMERICAN MINNOW GENUS CYPRINELLA 169

callitaenia, callisema, trichroistia, and gibbsi all have a very small process off the scapula, much smaller than in other cyprinids. All other species of the genus lack any type of process. The absence of the scapular process in Cyp- rinella and Luxilus is derived and supports the monophyly of these two groups. Postcleithrum (PCL, Fig. 31). This bone is roughly S -shaped and weakly attached to the mesial surface of the cleithrum posteriorly. It is generally flattened dorsally and may or may not be expanded. Ventrally, it is pointed and generally not flattened. No variation occurs between species in the shape or size of the postcleithrum.

Pelvic Girdle

The pelvic girdle is simple and essentially monomorphic in Cyprinella (Fig. 84). Each ISC girdle consists of a large, expanded ba-

sipterygium (BPT) which is concave dorsally. Fig. 84. Pelvic girdle of Cyprinella lutrensis (55 mm, KU Horizontal bar 1 mm. Anteriorly, it is forked and produces two pointed 19431). equals processes. Posteriorly, the radials (RAD) are

attached laterally. Medially, a hooked ischiac mesial radial is elongate and slightly hooked. process (ISC) is developed. Between the is- The lateral two radials are rounded. Coburn chiac process and the lateral margin, three (1982) discusses muscle attachments on the radials articulate along the posterior edge. The pelvic girdle.

PHYLOGENETIC RELATIONSHIPS

The evidence for monophyly of the genus representatives of the lutrensis clade. Lytle Cyprinella is discussed above and will not be examined species relationships in the lepida repeated here. Below is a sequential listing of species group. Only Gibbs presented any sort characters which support species relationships of formal phylogeny of the species studied. illustrated in Fig. 85. For each stem the sup- Contreras-Balderas and Lytle did not illustrate porting suite of characters is listed, followed a phylogeny, but discussed relationships within by a discussion of nonosteological synapo- the lutrensis clade as it was then comprised. morphies. For osteological characters appro- Results of the present study based on 206 priate arguments for determination of charac- osteological, meristic, coloration, and tuber- ter polarity within Cyprinella are provided in culation character transformations suggest the osteology section of this paper. species relationships within Cyprinella that Statements of species relationships within are at variance with some conclusions of these Cyprinella are few. The most comprehensive authors. Some parts of the phylogeny are, works on this group have been by Gibbs (1955, however, very similar to the phylogeny pre- 1957a), Contreras-Balderas (1975), and Lytle sented by Gibbs. Two major clades are recog- (1972). Gibbs evaluated species relationships nizable in the genus, the primarily southwestern within the whipplei clade (as used here), and lutrensis clade and the eastern whipplei clade Contreras-Balderas concentrated on Mexican (Fig. 85). Within the lutrensis clade, ornata is 170 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the sister of all other members, which fall into Justifications for these phylogenetic statements three monophyletic species groups: the lutren- are presented below and discussed in the oste- sis species group, \htformosa species group, ology section. and the lepida species group. The lutrensis Cyprinella lutrensis clade (Suite 2). The group includes lutrensis and garmani and is monophyly of this group is supported by 29 the sister to a clade composed of \heformosa shared derived characters. Characters indica- and lepida species groups. Included in the tive of a common origin in this group include: formosa group aizformosa and bocagrande. 1) a short second infraorbital bone, 2) facial In the lepida group lepida is the sister to foramen with an internal jugular process on proserpina plus panarcys and rutila plus xan- lateral rim, 3) ectopterygoid extremely elon- thicara. In the whipplei clade, spiloptera is the gate and thin, 4) posterior process of quadrate sister to the remaining members, which form joins vertical plate well below edge, 5) opercu- four monophyletic groups. The whipplei spe- lum narrow and deep, 6) anterior margin of cies group includes camura, whipplei, frontals with mesial extensions over su- analostana, and chloristia and is the sister praethmoid, 7) frontals widely broadened lat- group to remaining members. The venusta erally, 8) supraethmoid extremely short, 9) species group includes venusta and galactura metapterygoid-interhyal connection extremely and is the sister group to a clade composed of narrow, 10) posterior tip of subopercle greatly the pyrrhomelas and nivea species groups. elevated, 11) vertical plate of urohyal deep, The pyrrhomelas species group includes two 12) optic foramen bordered largely by ptero- monophyletic groups. The first is the sphenoid, 13) lacrymal curled, 14) branchios- pyrrhomelas species pair, inclusive of tegals broad, 15) occipital region extremely pyrrhomelas and xaenura, and the second is compressed and vertical, 16) ridge between the caerulea species complex, including this parietals and supraoccipital present, 1 7) dorsal species, gibbsi, and trichroistia. The nivea process of epibranchial 4 rarely developed species group forms the sister group to the and, when present, small, 18) parietals form- pyrrhomelas group and includes callistia, ing small shelf over supraoccipital region, 19) nivea, leedsi, callisema, and callitaenia. posterior margin of gnathic ramus of dentary

> • » A ** V o * 4 q + c.

2 6

Cyprinella

character suites are Fig. 85. Hypothesized phylogenetic relationships of species in the genus Cyprinella. Numbered discussed in the text. NORTH AMERICAN MINNOW GENUS CYPRINELLA 171 very tall, 20) dorsolateral shelf of dentary reduced tuberculation are considered conver- present, 21) gnathic ramus of dentary deep gent. anteriorly, 22) neck of epibranchial 1 bent The enlarged central tubercles of the cau- posteriorly, 23) neural complex and fourth dal-peduncle scales and the dense mat of tu- neural spine extremely tall, 24) predorsal bones bercles on the lower caudal peduncle are de- large and continuing to dorsal fin origin, 25) rived for this group of species since no other neural spines tall, 26) epural heavy, 27) Cyprinella have this pattern and in no close posttemporal bone tear shaped, 28) no pharyn- relatives do these conditions exist. If out- geal teeth in secondary row, and 29) mandibu- groups have central tubercles on the caudal- lar tubercles absent. peduncle scales they are equal in size to mar- The reduction to no teeth in the secondary ginal and edge tubercles. The only other cyp- tooth row is derived, since other members of rinid examined with a similar mat of tubercles Cyprinella and species in the genera Luxilus on the lower caudal peduncle is the Mexican and Lythrums all have a secondary tooth row. Dionda dichroma (pers. obs.). Since close relatives and most other members Cyprinellaformosa-lepida species groups of the genus have mandibular tubercles, the (Suite 4). Four osteological characters support absence of these organs in most species of the the monophyly of these seven species. Shared lutrensis clade is interpreted here as a derived derived characters include: 1) facial foramen condition. with internal jugular strut, 2) pharyngeal pad Cyprinella lutrensis-formosa-lepida spe- of basioccipital oval, 3) posterior margin of cies groups (Suite 3). The monophyletic ori- gnathic ramus of dentary tall, and 4) men- gin of these three species groups combined is tomeckelian short and cone-shaped. supported by nine derived characters, includ- Cyprinella lepida species group (Suite 5). ing: 1) smooth anterior hyomandibular wing, The monophyly of this clade of five species is 2) horizontal plate of urohyal broad, 3) lateral supported by 1 1 osteological characters and a slope of dermopterotic steep, 4) exoccipitals single derived condition of breeding colors. tall, 5) ceratohyals very broad, 6) gular bar These include: 1) frontals slightly broadened, very dark, 7) preorbital tubercles few and 2) junction of sphenotic and frontals moved reduced in distribution to the anterior lacrymal anteriorly, 3) ceratobranchial 2 with a long and area, 8) central tubercles of caudal-peduncle thin neck, 4) vomer extends beyond posterior scales enlarged relative to submarginal and edge of lateral ethmoids, 5) infraorbitals 2 and edge tubercles, and 9) tuberculation on caudal 3 broken into 2-3 smaller segments, 6) lateral peduncle greatest below lateral line, forming slope of dermopterotics nearly vertical, 7) tubercle pad from anal fin origin to base of coronoid process of dentary narrow and elon- caudal fin. gate, 8) retroarticular L-shaped, 9) retroarticu- The dark gular stripe is unique to this group lar long, 10) lateral margin of pharyngobran- of Cyprinella and represents a further deriva- chial 2-3 expanded into small shelf, 1 1) uroneu- tion of the lightly pigmented gular bar unique ral 3 short, and 12) breeding colors of males to the genus. The restriction of preorbital tu- purplish yellow. bercles to the anterior lacyrmal is derived for The distinctive purple-yellow breeding this group since ornata, other Cyprinella coloration of males of these species is de- (where applicable), and close relatives of Cyp- scribed in their respective species accounts. rinella have tubercles dispersed over the la- This condition is unique to this subgroup of crymal area. Some species of the whipplei Cyprinella, and is not presently known to clade also have a reduced tubercle pattern in occur outside the genus. the preorbital region, but in these species the Cyprinella rutila-proserpina species pairs tubercles are found along the dorsal margin of (Suite 6). This clade is one of the most distinc- the lacrymal. Thus, these two conditions of tive groups of the genus, supported by 23 172 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY shared derived characters. These include the neck of ceratobranchial 1 restricted, 7) ante- following: 1) narrow fourth infraorbital, 2) rior margin of ceratobranchial 3 expanded and posteroventral and ventral margins of pre- rounded, 8) posterior process of vomer nar- opercle scalloped, 3) frontals narrow, 4) posi- row, 9) pharyngeal process of basioccipital tion of epiphyseal bar anteriorly placed due to elongate and narrow, 10) ascending process of a change in frontal morphology, 5) neck of premaxilla elongate and narrow, 11) anterior ceratohyal broad, 6) neck of ceratobranchial 3 gnathic ramus deep, 1 2) mesial neck ofepibran- elongate, 7) anterior body of premaxilla deep, chial 1 bent anteriorly, 13) dorsal hypohyal 8) rostral process of maxilla heavy, 9) hy- small, and 14) neural complex and fourth pohyal foramen enclosed by anterior cera- neural spine short. tohyal only, 10) neck of branchiostegal 1 nar- Cyprinella proserpina species pair (Suite row, ll)anteriornotchofvomerverydeep, 12) 8). These two species share 1 1 derived osteol- supraorbitals broad anteriorly, 13) overlap ogical and tubercle characters. These include between supraorbital and lateral ethmoid great, the following: 1) ventral and posteroventral 14) parietals form large shelf over anterior margins of preopercle deeply scalloped, 2) margin of supraoccipital, 1 5) anterolateral edge ventral margin of metapterygoid with irregu- of palatine expanded anteriorly into elongate lar processes extending anteriorly into region process, 16) rostral process of maxilla bent normally occupied by symplectic, 3) posterior into L-shaped rod, 17) dorsolateral shelf of tip of subopercle extremely elevated above dentary strongly developed, 18) broad shelf anterior tip, 4) anteroventral margin of lateral formed from lateral margin of pharyngobran- ethmoid straight, 5) dorsal margin of premax- chial 2-3, 19) neck on epibranchial 2 bent illa thick, posterior ramus continues forward, anteriorly, 20) neck on epibranchial 3 bent 6) central process of sesamoid articular short anteriorly, 21) gular bar dark and extending and directed dorsally, 7) retroarticular re- from symphysis to isthmus of branchiostegals, shaped, 8) retroarticular long, 9) postorbital 22) caudal-peduncle tubercles absent on dor- tubercles smaller than anterior tubercles on salmost scale row, and 23) tubercle pad on dorsum of head, 10) preorbital tubercles dis- 1 lower caudal peduncle reduced to 1-1 h scale tributed over entire lacrymal area, and 11) no rows above midventral scale row. hiatus between dorsal head tubercles and snout The long gular bar is unique to this group tubercles. and represents a derived condition over the The reduced size of postorbital tubercles is short gular bar unique to Cyprinella. The re- unique to these two species and ornata of all duced caudal-peduncle tuberculation of these cyprinids examined. The occurrence of this species, lack of tubercles on dorsalmost scale derived condition in these two groups must be row of caudal peduncle, and narrow mat of convergent when other characters are consid- tubercles on the lower caudal peduncle is ered with parsimony. Further, with reference derived. Outside this group, the tubercle mat is to other characters, having preorbital tubercles deep, extending up to the lateral line, and when distributed over the entire lacrymal region in caudal peduncle tubercles are present, they are these two species is considered a reversal to almost always on the dorsal scale row. the primitive state for the genus. As noted Cyprinella rutila species pair (Suite 7). above, other members of the lutrensis clade Fourteen osteological characters support the have preorbital tubercles restricted to the ante- monophyly of these two species. Bom species rior edge of the lacrymal region. The absence share the following: 1) a narrow preopercle, 2) of a distinct hiatus between the snout and anterior wing of hyomandibular with a median dorsal head tubercles in these two species also bulge, 3) posterior wing of hyomandibular represents a reversal to the primitive morphol- wide,4) operculum broad, 5) supraorbital canal ogy for the genera Luxilus and Cyprinella. simple, without many canaliculi, 6) anterior Cyprinella formosa species group (Suite NORTH AMERICAN MINNOW GENUS CYPRINELLA 173

9). Ten derived characters support the mono- reversed in some members, this group shares phyly of this species pair. These species share the following: 1) anterior hyomandibular wing the following synapomorphies: 1) broad oper- broad, 2) posterior hyomandibular wing broad, culum, 2) sternohyoides processes of urohyal 3) supraethmoid very broad and with no emargi- very long, 3) subopercle nearly horizontal, 4) nate edges, 4) supraethmoid elongate, 5) ba- pharyngobranchial 2-3 narrow and straight, 5) sibranchial 1 with slight median constriction, posterior plate of epibranchial 1 joins mesial 6) ventral surface of lateral ethmoid com- neck dorsally, 6) anterior process of epibran- pletely ossified in adults, 7) lacrymal elongate, chial 1 located mesoanteriorly, 7) ceratohyals 8) pharyngeal process of basioccipital elon- moderately deep, 8) dorsal head tubercles erect, gate and narrow, 9) anterior neck of parasphe- 9) supraorbital tubercle scattered, and 10) noid long, 10) posterior ramus of premaxilla caudal-peduncle tubercles below lateral line straight, 11) premaxillary process of maxilla very thick, forming shagreen. tall, 12) anterior margin of epibranchial 1 with Antrorse tubercles on the dorsum of the anteromesial process, 13) seventh to ninth head are derived for the genus Cyprinella, but dorsal fin membranes with melanophores reversed in this species pair to the more primi- heavily concentrated, 14) dorsal head tubercles tive morphology of erect tubercles, as in out- strongly antrorse, 15) preorbital tubercles groups. The condition of numerous and scat- antrorse, 16) tubercle connection between tered supraorbital tubercles is derived for these supraorbital and preorbital regions composed two species since close outgroups typically of two rows, 17) snout tubercles erect anteri- have a single row of tubercles above the orbit. orly and retrorse posteriorly, 18) supraorbital The shagreen tubercle pattern on the lower tubercles antrorse, 19) flank scales above lat- caudal peduncle is distinctive for these two eral line with several scattered tubercles lo- species and represents a further modification cated centrally, 20) flank scales below lateral of the mat of tubercles diagnostic for the lutren- line with several scattered tubercles located sis clade. centrally, and 21) dorsal fin with dark interra- Cyprinella lutrensis species group (Suite dial membranes posteriorly in breeding males 10). The monophyletic origin of these two only. species is supported by nine derived charac- The darkened posterior membranes of the ters. Both members share the following: 1) dorsal fin of all of these species is unique and posterior process of quadrate joins the vertical derived for this group of cyprinids. A similar plate at edge, 2) epiphyseal bar placed anteri- condition is found in Hybopsis labrosa and H. orly with no modifications of frontals, 3) pos- zanema, but in these species all membranes terior tip of subopercle only slightly elevated are dark. These two species may be related to above anterior tip, 4) basihyal deep, 5) mid- other Hybopsis (s.s.). Thus, this condition is lateral cavities of basihyal large, 6) neck on considered independent in both groups. This vomer absent, 7) epibranchial 4 with short clade of Cyprinella is further defined by the mesial to dorsal process, 8) posttemporal bone presence of dark interradial membranes in not strongly tear shaped, and 9) scapular bar breeding males only. pigmentation strongly developed. All members of this clade have antrorse The very dark scapular bar of these two tubercles that are much more strongly devel- species is derived. It is interpreted as a further oped than in members of the lutrensis clade. modification of the less intense scapular bar Thus, this condition is considered to be a diagnostic for the genus. further modification of the antrorse structure Cyprinella whipplei clade (Suite 11). The ancestral to the genus. The antrorse preorbital monophyly of this clade of 17 species is evi- and supraorbital tubercles found in this group denced by their sharing 21 derived features. are very rare outside Cyprinella and are con- Although some of these characteristics are sidered derived for this clade. The condition of 174 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY having multiple rows of tubercles connecting Cyprinella whipplei species group (Suite the supraorbital and preorbital regions is con- 1 3). This clade of four species is diagnosed on sidered derived for this clade since close out- the basis of seven derived features. These groups and members of the lutrensis clade include: 1) anterior hyomandibular wing nar- have a single row connection. The presence of row, 2) anterior hyomandibular wing smooth, erect tubercles anteriorly and antrorse tubercles 3) supraethmoid short, 4) no overlap present posteriorly on the snout is unique to this clade. between lateral ethmoid and supraorbital, 5) Close outgroups and species of the lutrensis isthmus of maxilla deep, 6) mentomeckelian clade have erect tubercles over the entire snout short and cone shaped, and 7) supraorbital region. In close outgroups and members of the tubercles scattered. lutrensis clade, centrally located tubercles on The scattered arrangement of supraorbital flank scales both above and below the lateral tubercles of this group is derived from the line are rare. Thus, the consistent development single row pattern found in most members of of central tubercles on flank scales is consid- the lutrensis clade and close outgroups. The ered derived for this clade. occurrence of scattered tubercles on the su- Cyprinella wh ipplei- venusta -pyrrho- praorbital region in theformosa species group melas-nivea species groups (Suite 12). The of Cyprinella is considered independent of the monophyly of this large eastern clade of Cyp- condition found in these four species. rinella is diagnosed by the presence of five Cyprinella whipplei-analostana-chloristia relation- derived features: 1) nasal canal with 2 pores, 2) species group (Suite 14). The close has optic and hypophyseal foramina divided by ship of these three species been suggested small processes off of pterosphenoid, 3) tu- for many years (Gibbs, 1957a). My study sup- bercle connection between supraorbital and ports this hypothesis. These three species share charac- preorbital regions wide, composed of 3-4 rows the following six derived osteological of tubercles, 4) dorsal fin of both males and ters: 1) posterior hyomandibular wing curled, females with dark membranes posteriorly, and 2) anterior margin of frontals with mesial midlateral 5) dorsal fin expanded in breeding males. extensions over supraethmoid, 3) The increased number of tubercle rows cavities of basihyal large, 4) supraorbital bones not ex- connecting the supraorbital and preorbital broad anteriorly, 5) dermopterotics regions is derived for this clade and represents tending over dilator fossa, and 6) ascending a further modification of the two rows present wings of parasphenoid very broad. in the ancestor of the whipplei clade. This Cyprinella analostana-chloristia species 1 of these group of Cyprinella is further derived over the pair (Suite 5). The sister relationship ancestor of the whipplei clade in that both two species is supported by 16 derived charac- males and females develop the darkened pos- ters: 1) facial foramen with internal jugular terior membranes of the dorsal fin. In spilop- process on lateral rim, 2) posterior process of tera only males develop this color pattern and quadrate deep and heavy, 3) posteriorly di- it is absent in the lutrensis clade and outgroups. rected process of ceratobranchial 3 serrate, 4) The expanded dorsal fin of these species is dentary with dorsolateral shelf, 5) anterior derived, but not unique to this group. A similar margin of pharyngobranchial 1 with lateral condition is found in Notropis hubbsi and process, 6) neck on epibranchial 1 twisted, 7) canal with Pteronotropis welaka and was mentioned by vomer with long neck, 8) nasal Bailey and Robison (1978) as possible evi- single pore, 9) lateral slope of dermopterotics dence to relate these species to Cyprinella. steep, 10) posterior ramus of premaxilla Based on my observations and those of Dim- strongly curved downward, 11) ascending and mick (1987), these species may not be closely process of premaxilla elongate narrow, related to each other, and are not intimately 12) mesial neck of epibranchial 2 bent anteri- foramen of cera- related to the genus Cyprinella. orly, 13) posterior hyoideal NORTH AMERICAN MINNOW GENUS CYPRINELLA 175 tohyal located mesially, 14) scapula with small, acute pattern, 8) pigmentation on scales below dorsoposterior process, 15) mandibular tu- lateral line in strongly acute pattern, and 9) bercles scattered, and 16) tubercle connection supraorbital tubercles scattered. between supraorbital and preorbital regions The strongly acute diamond- shaped pat- narrow, consisting of single row. tern on lateral body scales of these two species The scattered pattern of mandibular tu- is considered a further derivation of the cross- bercles is not unique to this group of Cyp- hatched pattern developed in the ancestor of rinella butbased on other the In these the , characters, most Cyprinella. two species pigment parsimonious explanation for its presence in pattern on the scales is very well defined and these species represents convergent evolution. sharply pointed posteriorly on each scale. As The single row of tubercles connecting the in the analostana-chloristia species pair and supraorbital and preorbital regions found in the formosa species group, the presence of these species is primitive for Cyprinella, but scattered supraorbital tubercles in these two represents a reversal to the primitive condition species is considered derived and convergent. in their common ancestor. Thus, this character Cyprinella pyrrhomelas-nivea species is indicative of their monophyly. groups (Suite 18). Based primarily on tu- Cyprinella venusta-pyrrhomelas-nivea bercle development, this distinctive clade of species groups (Suite 16). The monophyletic Cyprinella was considered by Gibbs (1957a) origin of these three clades is supported by to be primitive in the genus. Members of this nine derived features. These include: 1) junc- clade share 20 derived characteristics: 1) su- tion of sphenotic and frontal bones moved pos- praethmoid broadened and with slightly teriorly, 2) anteroventral margin of lateral emarginate edges, 2) basibranchial 1 with a ethmoid slightly concave, 3) ventromesial strong medial constriction, 3) posterior mar- margin of lateral ethmoid truncated, 4) loss of gin of horizontal urohyal plate moderately a connection between the frontals and der- notched, 4) deep anterior notch of urohyal mopteroucs, 5) optic and hypophyseal foram- separating heavy struts, 5) anteroventral mar- ina nearly divided by mesial processes of ptero- gins of lateral ethmoid straight and oblique, 6) sphenoids, 6) posterior ramus of premaxilla supraorbital bones broad anteriorly, 7) dilator with uniform ventral curve, 7) mesial neck of fossa not roofed by dermopterotic, 8) dermop- epibranchial 2 bent anteriorly, 8) ceratohyals terotics invade parietals, 9) epibranchial 1 slender, and 9) caudal base with distinct oval with anterior process located medially, 10) spot. ceratobranchial 4 without a posterior process, Although variable among species (see spe- 11) dorsal hypohyal smaller than ventral hy- cies accounts section), the dark and oval cau- pohyal, 12) caudal-peduncle tubercles absent dal spot is derived for this clade of Cyprinella, from dorsal-most scale row, 13) caudal- being absent in all other species of the genus peduncle tubercles absent from ventralmost and close outgroups. scale row, 14) first membrane of dorsal fin Cyprinella venusta species group (Suite heavily pigmented with melanophores, ap- 17). These two species, venusta and galactura, pearing black, 15) fourth membrane of dorsal have not previously been associated. How- fin uniformly pigmented except for a clear ever, they share nine derived characters rela- window posteriorly, 16) fifth membrane of tive to other members of the genus and out- dorsal fin with depigmented window posteri- groups: 1) preopercle very wide over entire orly, 17) sixth membrane of dorsal fin with length, 2) posterior process of vomer narrow, depigmented window posteriorly, 18) tubercle 3) anterodorsal margin of lacrymal elevated, pattern on dorsum of head linear throughout 4) retroarticular L-shaped, 5) retroarticular development, 19) predorsal scales each with a long, 6) caudal skeleton narrow, 7) pigmenta- single large tubercle, and 20) caudal-peduncle tion on scales above lateral line in strongly scales below lateral line each with a single 176 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY large antrorse tubercle. this group. These species share the following: The presence of extremely large, antrorse I) first ceratobranchial with neck bent me- tubercles on predorsal and caudal-peduncle sially, 2) olfactory foramen bounded by later- scales is unique to this clade of cyprinids, with ally directed shelf of mesethmoid and lateral the exception of Hybopsis labrosa and H. ethmoid, 3) anterior notch of vomer very shal- zanema. Tubercles in these two species are low, 4) posterior process of vomer terminates very similar to those of this clade of Cyp- before posterior edge of lateral ethmoid, 5) rinella, but since these taxa can be related to lacrymal short, 6) pharyngeal pad on basioc- other Hybopsis, and since the tubercles are not cipital flat, 7) preethmoid socket of palatine generally located centrally on a scale and shallow, 8) a single ligament connecting the number more than one, the presence of large palatine to the ecto- and endopterygoids, 9) tubercles on the caudal peduncle is considered adductor mandibulae Al insertion on maxilla independent in the two groups. The absence of very far forward, anterior to isthmus, 10) coro- tubercles on the dorsal and ventral scale rows noid process of dentary posteriorly directed, of the caudal peduncle is also found in the II) flank scales above lateral line without proserpina and rutila species pairs. This con- central tubercles, and 12) flank scales below dition is considered independent in these two lateral line without central tubercles. clades since in the proserpina -rutila clade the Members of this clade lack central tubercles tubercles are absent from the dorsal scale row on flank scales. This condition is a reversal to only. a primitive state found in the lutrensis clade The linear tubercle pattern on the dorsum of and close outgroups, but on the basis of other the head throughout development of breeding characters, this is considered to be evidence of males is derived for this clade. Because Cyp- monophyly of this group. rinella begin development with a linear pat- Cyprinellapyrrhomelas species pair (Suite tern, Gibbs (1957a) considered the linear pat- 20). The monophyly of this species pair is tern of adults to be primitive within Cyprinella supported by 21 derived characters. Both and not indicative of monophyly of the group. species share the following characters: 1) lat- The presence of a linear pattern in these spe- eral commissure narrow, 2) trigeminal fora- cies is here interpreted as a case of neotenic de- men bordered anteroventrally by a ventral velopment of secondary sexual features. In process of prootic, 3) prootic depression pos- this clade the linear pattern in early develop- terior to lateral commissure covered by a shelf, ment is retained in the peak breeding season. 4) branchiostegals short, 5) ascending arm of This seems likely since these species all pro- preopercle tall, 6) epibranchial 4 narrow, 7) duce larger dorsal head tubercles and not addi- supraethmoid very broad and without emargi- tional tubercles over the head to disrupt the nate edges, 8) supraethmoid extremely short, linear pattern. 9) nasal canal with one pore, 10) supraorbital Four characteristics of the dorsal fin appear bones short, 11) fourth membrane of dorsal fin to be derived for species of this group. All have with pigment uniformly distributed, 12) fifth a darkened first membrane, not found in any membrane of dorsal fin with pigment uni- other Cyprinella and absent from close out- formly distributed, 13) sixth membrane of groups. All species, except pyrrhomelas and dorsal fin with pigment uniformly distributed,

1' xaenura, also develop large depigmented 14) dorsal head tubercle very large, 15) pre- windows in the darkened forth, fifth, and sixth dorsal tubercles very large and with irregular membranes of the dorsal fin. This pigment bases, 16) tubercle connection between su- pattern is not observed outside this group. praorbital and preorbital region narrow, 17) Cyprinella pyrrhomelas species group tubercle hiatus between snout and dorsum of (Suite 19). Twelve derived osteological and head absent, 18) anterior body scales above tubercle features are shared by members of lateral line without tubercles, 19) anterior body NORTH AMERICAN MINNOW GENUS CYPRINELLA 177 scales below lateral line without tubercles, 20) bercles scattered. enlarged tubercles found both above and be- As in \heformosa group, the scattered tu- low lateral line on caudal peduncle, and 21) bercle pattern above the orbit is derived. In caudal-peduncle tubercles below lateral line reference to other derived characters, this very large and with irregularly formed bases. condition is best interpreted as convergent in The pigment pattern on the fourth, fifth, and these groups. These species also share the sixth dorsal fin membranes is derived for these primitive condition within Cyprinella of hav- species, representing a reversed condition to ing melanophores uniformly distributed over the primitive uniform color pattern, typical of the first membrane of the dorsal fin and scat- species in the lutrensis clade and outgroups. tered central tubercles. Both of these condi- The enlarged tubercles with irregularly formed tions represent reversals to a primitive state bases on the predorsal and caudal-peduncle within Cyprinella. However, with reference to scales is derived since no other members of the other derived characters, these conditions

genus have tubercles this large, and the irregu- should not be considered homologous with the lar base morphology has never been observed conditions found in the lutrensis clade and outside this group. The enlarged dorsal head outgroups. Thus, they represent independent tubercles of these species represents a further evolutionary events in this clade, indicative of enlargement of head tubercles in Cyprinella. monophyly of this species complex. No other species develop tubercles as large as Cyprinella trichroistia-gibbsi species pair these. The presence of enlarged central tu- (Suite 22). The monophyly of these two spe- bercles both above and below the lateral line cies is supported by 16 derived features: 1) on the caudal peduncle is unique to this species anterior wing of hyomandibular smooth, 2)

pair. The absence of tubercles on flank scales posterior wing of hyomandibular terminating

both above and below the lateral line is also near ventral tip, 3) position of epiphyseal bar unique to these species. The reversed condi- displaced posteriorly, 4) supraorbital bones tion of having a single tubercle row connecting long, 5) lacrymal canal smoothly arched, 6) the tubercles of the supraorbital and preorbital preethmoid socket of palatine very shallow, 7) regions, and absence of a hiatus between dor- neck of epibranchial 1 bent posteriorly, 8) sal head tubercles and snout tubercles is more scapula with small dorsoposterior process, 9) parsimoniously considered derived for this diamond-shaped pigmentation pattern on scales clade if the distributions of other characters are above lateral line diffuse, 10) second mem- considered. brane of dorsal fin depigmented distally and Cyprinella caerulea species com plex (Suite dark basally, 11) third membrane of dorsal fin 21). This species group is supported by 12 depigmented distally and dark basally, 12) shared derived characters: 1) ascending arm of fifth membrane of dorsal fin with large depig- preopercle very short, 2) location of sphe- mented window posteriorly and darkly pig- notic-frontal articulation strongly displaced mented dorsally, 13) sixth membrane of dorsal posteriorly, 3) anterior notch of vomer shal- fin with large depigmented window posteri- low, 4) lateral slope of dermopterotics flat- orly and darkly pigmented dorsally, 14) sev- tened and nearly horizontal, 5) posttemporal enth to ninth membranes of dorsal fin darkly fossa small and compressed, 6) posterior ramus pigmented distally, 1 5) scales above the lateral of premaxilla very narrow and pointed, 7) line along flank with scattered central tubercles, hypohyal foramen narrow and slitlike, 8) isth- and 16) scales below the lateral line along mus of maxilla very narrow, 9) uroneural 3 flank with scattered central tubercles. short, 10) first membrane of dorsal fin with The diffuse diamond-shaped pattern is a pigment uniformly distributed, 11) centrally secondary loss of the sharp pattern. Young of located tubercles of predorsal scales number- both species have the typical diamond-shape ing 2-3 per scale, and 12) supraorbital tu- pattern on flank scales. The five features of the 178 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY dorsal fin membranes are unique to this clade Cyprinella nivea-leedsi-callisema-calli- and thus considered derived. The scattered taenia species group (Suite 24). The mono- tubercle pattern of flank scales above and phyly of these four species is supported by 17 below the lateral line is considered derived for derived osteological features: 1) short and the whipplei clade, being absent in the lutren- broad ectopterygoid, 2) frontal bones narrow, sis clade and outgroups. 3) metapterygoid-interhyal articulation nar- Cyprinella nivea species group (Suite 23). row, 4) anterior margin of ceratobranchial 1 The monophyly of this distinctive clade of five expanded anteriorly, 5) basibranchial 1 with species is supported by their sharing 16 dean hourglass shape, having a strongly devel- rived features of their osteology, coloration, oped median constriction, 6) mesethmoid long, and tuberculation. These include the follow- 7) anterior notch of vomer very deep, 8) optic

ing: 1) epiphyseal bar displaced posteriorly, 2) and hypophyseal foramina completely sepa- neck on ceratobranchial 1 restricted, 3) hy- rated by pterosphenoid, 9) palatine elongate, pobranchial 1 with a double cartilage head, 4) 10) dorsal margin of premaxilla serrate, 11) vomer with a long neck, 5) lacrymal extremely premaxilla strongly bent, 12) anterolateral edge long, 6) lacrymal canal L-shaped, 7) parietals of palatine expanded into elongate process, strongly invaded by dermopterotics, 8) mandi- 13) rostral process of maxilla bent into re- bular pores 3,9) mouth inferior, 10) ascending shaped rod, 14) isthmus of maxilla moderately process of premaxilla narrow and elongate, shallow, 15) premaxillary process of maxilla 11) gnathic ramus of dentary curved ventrally extremely tall, 16) caudal skeleton narrow, and mesially, 12) fused pharyngobranchial 2- and 17) scapula with small dorsoposterior 3 with a smooth dorsal surface, 13) scapular process. bar pigment weakly developed, if present, 14) Cyprinella leedsi-callisema-callitaenia second membrane of dorsal fin uniformly pig- species group (Suite 25). These three species mented and with clear window posteriorly, 15) share seven derived characters: 1) ventrome- third membrane of dorsal fin uniformly pig- sial process of palatine short, 2) anterior cera- mented and with clear window posteriorly, tohyal with foramen ventral to anterior fork, 3) and 16) preorbital tubercles on dorsal half of anterior neck of ceratohyal narrow, 4) men- lacrymal only. tomeckelian short and cone-shaped, 5) no The scapular bar pigment typical of the pharyngeal teeth in secondary tooth row, 6) genus is reduced and sometimes absent in flank scales above lateral fine with a single members of this group. This is considered a central tubercle, and 7) flank scales below secondary modification as a reduction in pig- lateral line with a single central tubercle. mentation. In these species the unique condi- Based on parsimony argumentation, the tion of having large, clear, posterior windows loss of the secondary tooth row in callisema on the second and third membranes of the and leedsi is considered a derived condition

dorsal fin is derived since no other species in for this clade, with a reversal to the 1, 4-4, 1 Cyprinella or outgroups have this condition. count in callitaenia. All three of these species Also unique to this group within Cyprinella is share the unique conditions of having a single the reduction of preorbital tubercles to the tubercle located centrally on flank scales both anterior lacrymal. As discussed earlier for the above and below the lateral line. When central lutrensis clade, the reduction of lacrymal tu- tubercles are present in other species of the bercles is derived over what outgroups and genus they are scattered and number more than most Cyprinella have. Because the tubercle one. Thus, the single central tubercle condition reduction in this clade is quite different from of these species is derived. that discussed earlier for the lutrensis clade Cyprinella callisema-callitaenia species (exclusive ofornata), these two conditions are pair (Suite 26). These two species are very not homologous. similar morphologically and are here consid- NORTH AMERICAN MINNOW GENUS CYPRINELLA 179 ered sister species based on three osteological ventrally, and 3) a connection between the characters. These species share: 1) a narrow frontal and dermopterotic bones. interopercle, 2) a lacrymal strongly tilted SUMMARY

The primary intent of this study was to sis of osteological variation. sex- resolve species relationships of the tradition- Variation in osteology and secondary ual characteristics of North American ally recognized subgenus Cyprinella ofNotro- cyprinids that eastern North Ameri- pis. Based on comparisons with other cyp- suggests many In- rinids, the genus "Notropis" as traditionally can species form a monophyletic group. used is not a monophyletic group (sensu cluded in this group are species traditionally Hennig) and can no longer be considered as placed in the genus Notropis (except Aztecula such. Thus, in addition to the recognition of sallei), and the genera Cyprinella, Luxilus, Cyprinella as a distinct genus, other subgenera Lythrurus, Pteronotropis, Oregonichthys, of Notropis are elevated to generic status and Clinostomus, Richardsonius, Yuriria, some genera of North American cyprinids are Hybopsis, Pimephales, and a clade inclusive rearranged taxonomically. The resulting taxo- of Hybognathus, Exoglossum, Dionda, nomic changes have been done to maintain Campostoma, Nocomis, Platygobio, consistency between the phylogenetic rela- Macrhybopsis, Extrarius, Phenacobius, and all of these tionships of species and our existing classifi- Erimystax. Relationships of spe- cation. cies were not fully resolved and polytomies Monophyly of the genus Cyprinella is were obtained. Basally, a multichotomy exists supported by 38 characters. Species relation- between a clade inclusive of the genera ships based on 206 character transformations Pteronotropis, Cyprinella, Luxilus, Lythrurus, each of the remain- produced a fully resolved phylogeny of the a restricted Notropis, and genus. Within the genus, two major clades are ing clades above. The genus Pteronotropis recognized: the primarily southwestern United forms the basal sister group of the first clade States and Mexico lutrensis clade and the and the Lythrurus-Luxilus-Cyprinella clade is eastcentral United States whipplei clade. Spe- the sister to the restricted Notropis clade. The cies of the lutrensis clade include ornata, genus Lythrurus is hypothesized as the sister lutrensis, garmani, formosa, bocagrande, group to Luxilus plus Cyprinella. Within the lepida, rutila, and xanthicara. Species of the restricted Notropis clade, the subgenus Notro- whipplei clade include spiloptera, camura, pis is sister to the more inclusive Notropis whipplei, analostana, chloristia, venusta, texanus species group. This clade forms a galactura, pyrrhomelas, xaenura, caerulea, polytomy with the Notropis volucellus species Al- trichroistia, gibbsi, callistia, nivea, leedsi, cal- group and the subgenera Hydrophlox and lisema, and callitaenia. A detailed account for burnops. each species is presented, as well as an analy- LITERATURE CITED

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Meek, S. E. 1904. The fresh-water fishes of Mexico north Niazi, A. D. 1964. The development of the Weberian of the Isthmus of Tehuantepec. Field Columbian system and early embryology of Pimephales Mus., Publ. 93 (Zool. Ser.) 5:v-xiii, 1-252. promelas (Osteichthyes: Cyprinidae). Diss. Abs. 25:2674. Menzel, B. W. 1976. Biochemical systematics and evolutionary genetics of the common shiner species Nursall, J. R. 1963. The hypurapophysis, an important group. Biochem. Syst. Ecol. 4:281-293. element of the caudal skeleton. Copeia 1963(2):458-^59. Menzel, B. W. 1977. Morphological and electrophoretic identification of a hybrid cyprinid fish, Notropis Nybelin, 0. 1963. Zur morphologic und terminologie des cerasinus X Notropis c. cornutus, with implica- schwanzskelettes der Actinopterygier. Ark. Zool., tions on the evolution ofNotropis albeolus.Comp. Ser. 2(15):485-516. Biochem. Physiol. 57B:215-218. Outten, L.M. 1958. Studies of the life history of the

Mettee, M. F., P. E. O'Neil, R. D. Suttkus, and J. M. cyprinid fishes Notropis galacturus and rubricro- Pierson. 1987. Fishes of the lower Tombigbee ceus. J. Elisha Mitchell Sci. Soc. 74:123-134. River System in Alabama and Mississippi. Bull. Page, L. M. and R. L. Smith. 1970. Recent range adjust- Geol. Surv. Alabama. No. 107. 186 pp. ments and hybridization of Notropis lutrensis and Miller, R. J. and H. W. Robison. 1973. The Fishes of Notropis spilopterus in Illinois. Trans. HI. State Oklahoma. Oklahoma State Univ. Press, Stillwa- Acad. Sci. 63:264-272. ter. 246 pp. Paloumpis, A. A. 1958. Responses of some minnows to Miller, R. R. 1976. An evaluation of Seth E. Meek's flood and drought conditions in an intermittent contributions to Mexican ichthyology. Fieldiana, stream. Iowa St. Coll. J. Sci. 32:547-561. Zool. 69:1-31. Patterson, C. 1968. The caudal skeleton in lower Liassic Miller, R. R. 1978. Composition and derivation of the pholidophorid fishes. Bull. Br. Mus. (Nat. Hist.) native fish fauna of the Chihuahuan Desert Region. Geol. 16(5):201-239. In: R. H. Wauer and D. H. Riskind (eds.), Patterson, C. 1977. Cartilage bones, dermal bones and on the re- Transactions-symposium biological membrane bones, or the exoskeleton versus the sources of the Chihuahuan Desert U.S. Region, endoskeleton, In: A. Mahala, R. S. Miles, and A. and Mexico. Nat. Park Serv., Washington, D.C., D. Walker (eds.). Problems in vertebrate evolu- Trans. Proc. Ser. 3 (1977):365-381. tion. Linn. Soc. Symp. Ser. 4:77-121. Miller, R. R. and J. R. Simon. 1943. Notropis mearnsi Pflieger, W. L. 1965. Reproductive behavior of the from Arizona, an addition to the known fish fauna minnows,No/ropty spilopterus and Notropis whip- of the United States. Copeia 1943:253. plei. Copeia 1965:1-18. Minckley, W. L. 1959. Fishes of the Big Blue River basin, Pflieger, W. L. 1971. A distributional study of Missouri Kansas. Mus. Nat. Hist., Univ. Kansas Publ. fishes. Mus. Nat. Hist. Univ. Kansas Publ. 11:401^142. 20:225-570. Minckley, W. L. 1972. Notes on the spawning behavior of Pflieger, W. L. 1975. The Fishes of Missouri. Missouri red shiners, introduced into Burro Creek, Ari- Dept of Cons., Jefferson City. 343 pp. zona. Southwest. Nat. 17:101-103. Pierson, J. M. and R. S. Krotzer. 1987. The distribution, Minckley, W. L. 1973. Fishes ofArizona. Arizona Game relative abundance, and life history of the blue and Fish Dept., Phoenix. 293 pp. shiner, Notropis caeruleus (Jordan). Alabama Minckley, W. L. and G. L. Lytle. 1969. Notropis xanthi- Nongame Wildlife Final Report, Alabama Dept. cara, a new cyprinid fish from the Cuatro Ciene- of Conservation, Montgomery. 105 pp. gas basin, north-central Mexico. Proc. Biol. Soc. Rabito, F. G., Jr. and D. C. Heins. 1985. Spawning 82:491-502. Washington behavior and sexual dimorphism in the North Moore, G. A. 1950. The cutaneous sense organs of American cyprinid fish Notropis leedsi, the barbeled minnows adapted to life in the muddy bannerfin shiner. J. Nat. Hist. 19:1155-1163. waters of the Great Plains Region. Trans. Am. Rainboth, W J., Jr. and G. S. Whitt. 1974. Analysis of Micros. Soc. 69:69-95. evolutionary relationships among shiners of the Moore, G. A. 1952. The Fishes of Oklahoma. Oklahoma subgenus Luxilus (Teleostei, Cypriniformes, and Game and Fish Dept. 11pp. Notropis) with the lactate dehydrogenase malate dehydrogenase isozyme systems. Comp. Nelson, G. J. 1974. Classification as an expression of Biochem. Physiol. 49B:241-252. phylogenetic relationships. Syst. Zool. 22:344-359. Reno, H. W. 1966. The infraorbital canal, its lateral -line ossicles and neuromasts, in the minnows Notropis Niazi, A.D.I 960. A comparison of the Weberian appara- volucellus and N. buchanani. Copeia tus in the genus Pimephales (Cyprinidae). M.S. 1966:403-413. NORTH AMERICAN MINNOW GENUS CYPRJNELLA 185

Reno, H. W. 1969. Cephalic lateral-line systems of the Stout, J. F, and H. E. Winn. 1958. The reproductive cyprinid genus Hybopsis. Copeia 1969:736-773. behavior and sound production of the satinfin shiner. [abst.] Anat. Rec. 132:511. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A Suttkus, R. D. 1950. A taxonomic study of five cyprinid List of Common and Scientific Names of Fishes fishes related to Notropis hypselopterus of south- from the United States and Canada, 4th ed. Amer. eastern United States. Ph.D. Dissertation. Cornell Fish. Soc. Spec. Publ. No. 12. 174 pp. Univ., Ithaca, NY.

Rosen, D. E. 1 979. Fishes from the uplands and intermon- Swift, C. C. 1970. A review of the eastern North Ameri- tane basins of Guatamala: revisionary studies and can cyprinid fishes of the Notropis texanus spe- comparative geography. Bull. Amer. Mus. Nat. cies group (subgenus Alburnops), with a definition Hist. 162:267-376. of the subgenus Hydrophlox, and materials for a revision of the subgenus Alburnops. Ph.D. Dis- Saksena, V. R 1962. The post-hatching stages of the red sertation. Florida State Univ., Tallahassee. shiner, Notropis lulrensis. Copeia 1962:539-544. Trautman, M. B. 1957. The Fishes of Ohio. Ohio State Schaefer, S. A. and T. M. Cavender. 1986. Geographic Univ. Press, Columbus. 683 pp. variation and subspecific status of Notropis spilopterus (Pisces: Cyprinidae). Copeia Trautman, M. B. 1981. The Fishes of Ohio (revised edi- 1986:122-130. tion). Ohio State Univ. Press, Columbus. 738 pp.

Sisk, M. E. 1961. A comparative study of the Weberian Uyeno, T 1961. Osteology and phylogeny of the Ameri- apparatus in the genus Dionda and Notropis can cyprinid fishes allied to the genus Gila. Ph.D. (Cyprinidae). M.S. Thesis. Oklahoma State Univ., Dissertation. Univ. Michigan, Ann Arbor. Stillwater, OK. Wallace, R. K., Jr. and J. S. Ramsey. 1981. Reproductive Smith, RW. 1979. The Fishes of Illinois. Univ. Illinois behavior and biology of the bluestripe shiner Press, Urbana. 314 pp. (Notropis callitaenia) in Uchee Creek, Alabama. Am. Midi. Nat. 106:197-200. Snelson, F. F. 1972. Systematics of the subgenus Ly- thrurus, genus Notropis (Pisces: Cyprinidae). Bull. Wallace, R. K., Jr. and J. S. Ramsey. 1982. Anew cyprinid Florida State Mus., Biol. Sci. 17:1-92. hybrid, Notropis lulrensis X N. callitaenia, from the Apalachicola drainage in Alabama. Copeia Snyder, J. O. 1915. Notes on a collection of fishes made 1982:214-217. by Dr. Edgar A. Meams from rivers tributary to the Gulf of California. Proc. U.S. Nat. Mus. White, S. T. and D. C. Wallace. 1973. Diel changes in the 49:573-586. feeding activity and food habits of the spotfin shiner, Notropis spilopterus (Cope). Am. Midi. Starrett, W. C. 1950. Food relationships of the minnows Nat. 90:200-205. of the Des Moines River, Iowa. Ecology 31:216-223. Wiley, E. O. 1979. An annotated Linnean hierarchy, with comments on natural taxa and competing sys- Starrett, W. C. 1951. Some factors affecting the abun- tems. Syst. Zool. 28:308-337. dance of minnows in the Des Moines, River, Iowa. Ecology 32:13-27. Wiley, E. O. 1981. Phylogenetics: The Theory and Prac- tice ofPhylogenetic Systematics. John Wiley and Stein, D. W., J. S. Rogers, and R. C. Cashner. 1985. Sons, New York, NY. 439 pp. Biochemical systematics of the Notropis roseipinnis complex (Cyprinidae: Subgenus Lythrurus). Wiley, E. O. and R. L. Mayden. 1985. Species and Copeia 1985:154-163. speciation in phylogenetic systematics, with examples from the North American fish fauna. Stone, U. B. 1940. Studies on the biology of the satinfin Ann. Missouri Bot. Gard. 72:596-635. minnows, Notropis analostanus and Notropis spilopterus. Ph.D. Dissertation. Cornell Univ., Ith- Winn, H. E., and J. F. Stout. 1960. Sound production by aca, NY. the satinfin shiner, Notropis analostanus, and related species. Science 132:222-223. Stout, J. F. 1959. The reproductive behavior and sound production of the satinfin shiner, [abst.] Anat. Rec. Winterbottom, R. 1974. A descriptive synonymy of the 134:643-644. striated muscles of the teleostei. Proc. Acad. Nat. Sci. Philadelphia 125:225-317. 186 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

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Plate I. A) Cyprinella ornata, male, 54 mm SL, Rio Papagochic, Guerrero, Mexico; B) Cyprinella ornata, female, 46 mm SL, Rio Papagochic, Guerrero, Mexico; C) Cyprinella lutrensis, male, length and locality unknown, photo by W. N. Roston; D) Cyprinella garmani, male, 50 mm SL, Rio Nazas, Durango, Mexico; E) Cyprinella garmani, female, 59 mm SL, Rio Nazas, Durango, Mexico; F) Cyprinella formosa, male, 47 mm SL, Rio Casas Grandes, Chihuahua, Mexico; G) Cyprinella bocagrande, male, 43 mm SL, Ojo Solo, Chihuahua, Mexico; H) Cyprinella lepida, male, 52 mm SL, Nueces River, Real Co., Texas, photo by L. M. Page. NORTH AMERICAN MINNOW GENUS CYPRINELLA 187

A B

G H

Plate II. A) Cyprinella rutila, male, 39 mm SL, Rio San Juan, Nuevo Leon, Mexico; B) Cyprinella xanthicara, male, 46 mm SL, Cuatro Cienegas, Coahuila, Mexico; C) Cyprinella proserpina, male, 46 mm SL, Devils River, Val Verde Co., Texas, photo by L. M. Page; D) Cyprinella panarcys, male, length not listed, Rio San Pedro, Chihuahua, Mexico (UMMZ 198786), photo by R. R. Miller; E) Cyprinella spiloptera, male, 65 mm SL, Little Beaver Creek, Kankakee Co., Illinois, W. photo by L. M. Page; F) Cyprinella spiloptera, male, length unknown, Shoal Creek, Newton Co., Missouri, photo by N. Roston; G) Cyprinella camura, male, length unknown, Terrapin Creek, Graves Co., Kentucky, photo by B. M. Burr, H) Cyprinella camura, male, 89 mm SL, Terrapin Cr., Graves Co., Kentucky, photo by B. M. Burr. 188 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Plate HI. A) Cyprinella whipplei, male, 89 mm SL, St.«Francis River, Wayne Co., Missouri, photo by L. M. Page; B) Cyprinella analostana, male, 64 mm SL, Morgan Creek, Orange Co., North Carolina, photo by L. M. Page; C) Cyprinella chloristia, male, 78 mm SL, Pacolet River, Spartanburg Co., South Carolina, photo by L M. Page; D) Cyprinella chloristia, male, length unknown. North Carolina, photo by W. N. Roston; E) Cyprinella venusta, male, 80 mm SL, Guadalupe River, Kerr Co., Texas, photo by L. M. Page; F) Cyprinella galactura, male, 101 mm SL, Beaver Creek, Taney Co., Missouri, photo by L. M. Page; G) Cyprinella galactura, female, 79 mm SL, Buffalo River, Lewis Co., Tennessee; H) Cyprinella pyrrhomelas, male, 72 mm SL, South Saluda River, Pickens Co., South Carolina. NORTH AMERICAN MINNOW GENUS CYPRINELLA 189

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Plate IV. A) Cyprinella caerulea, male, length and locality unknown, photo by W. N. Roston; B) Cyprinella trichroistia, male, 80 mm SL, Rock Creek, Murray Co., Georgia, photo by B. M. Burr, C) Cyprinella trichroistia, male, 70 mm SL, Schultz Creek, Bibb Co., Alabama; D) Cyprinella gibbsi, male, 65 mm SL, Enitachope Creek, Clay Co., Alabama; E) Cyprinella callislia, male, length unknown, Conasauga River, Polk Co., Tennessee, photo by W. N. Roston; F) Cyprinella callistia, male, 85 mm SL, Turkey Cr., Tuscaloosa Co., Alabama; G) Cyprinella nivea, male, 87 mm SL, Pacolet River, Spartanburg Co., South Carolina, photo by L. M. Page; H) Cyprinella callisema, male, 63 mm SL, Oconee River, Clark Co., Georgia.

QL638.C94 M3 1989 Phylogcnctii studies "I North Vmeri Harvard MCZ Library

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68. Internal oral features of larvae from eight anuran families: Functional, systematic, evolutionary and ecological considerations. By Richard Wassersug. Pp. 1-146, 37 text-figures. June 24, 1980. Paper bound.

69. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodac- tylidae). By John D. Lynch and William E. Duellman. Pp. 1-86, 8 text-figures. August 29, 1980. Paper bound.

70. Sexual size differences in reptiles. By Henry S. Fitch. Pp. 1-72, 9 text-figures. February 27, 1981. Paper bound.

71. Late Pleistocene herpetofaunas from Puerto Rico. By Gregory Pregill. Pp. 1-72, 26 text- figures. May 8, 1981. Paper bound.

72. Leptodactylid frogs of the genus Eleutherodactylus in the Andes of Northern Ecuador and adjacent Columbia. By John D. Lynch. Pp. 1-46, 22 text-figures. July 8, 1981. Paper bound.

73 . Type and figured specimens of fossil vertebrates in the collection ofThe University of Kansas Museum of Natural History. Part I. Fossil fishes. By H.-P Schultze, J. D. Stewart, A. M. Neuner and R. W. Coldiron. Pp. 1-53. October 6, 1982. Paper bound.

74. Relationships of pocket gophers of the genus Geomys from the central and northern Great

Plains. By Lawrence R.Heaney and Robert M.Timm. Pp. 1-59, 19 text-figures. June 1, 1983. Paper bound.

75. The taxonomy and phylogenetic relationships of the hylid frog genus Stefania. By William E. Duellman and Marinus S. Hoogmoed. Pp. 1-39, 30 text-figures. March 1, 1984. Paper bound.

76. Variation in clutch and litter size in New World reptiles. By Henry S. Fitch. Pp. 1-76, 15 text- figures. May 24, 1985. Paper bound.

77 . Type and figured specimens of fossil vertebrates in the collection ofThe University of Kansas Museum of Natural History. Part II. Fossil amphibians and reptiles. By H.-P. Schultze, L. Hunt, J. Chorn and A. M. Neuner. Pp. 1-66. December 3, 1985. Paper bound.

78. Type and figured specimens of fossil vertebrates in the collection ofThe University of Kansas Museum of Natural History. Part III. Fossil birds. By John F. Neas and Marion Anne Jenkinson. Pp. 1-14. February 5, 1986. Paper bound.

79. Type and figured specimens of fossil vertebrates in the collection ofThe University ofKansas Museum ofNatural History. Part II. Fossil mammals. By Gregg E. Ostrander, Assefa Mebrate and Robert W. Wilson. Pp. 1-83. November 21, 1986. Paper bound.