(Percidae: Etheostoma), with Descriptions of Five New Species

BULLETINALABAMA MUSEUM OF NATURAL HISTORYThe scientific publication of the Alabama Museum of Natural History. Dr. RobertClouse, Editor.BULLETIN ALABAMA MUSEUM OF NATURAL HISTORY is published by theAlabama Museum of Natural History, a unit of The University of Alabama. TheBULLETIN succeeds its predecessor, the MUSEUM PAPERS, which was terminatedin 1961 upon the transfer of the Museum to the University from its parent organization,the Geological Survey of Alabama.The BULLETIN is devoted primarily to scholarship and research concerning thenatural history of Alabama and the Southeast. It appears twice yearly in consecutivelynumbered issues.Communication concerning manuscripts, style, and editorial policy should beaddressed to: Editor, BULLETIN ALABAMA MUSEUM OF NATURAL HISTORY,The University of Alabama, Box 870340, Tuscaloosa, Alabama 35487-0340; telephone(205) 348-7551 or emailed to rclouse@bama.ua.edu. Prospective authorsshould examine the Notice to Authors inside the back cover.Orders and requests for general information should be addressed to BULLETINALABAMA MUSEUM OF NATURAL HISTORY, at the above address or emailed tocaverett@ua.edu. Yearly subscriptions (two issues) are $30.00 for individuals, $50.00for corporations and institutions. Numbers may be purchased individually. Paymentshould accompany orders and subscriptions and checks should be made out to “TheUniversity of Alabama.” Library exchanges should be handled through: ExchangeLibrarian, The University of Alabama, Box 870266, Tuscaloosa, Alabama 35487-0340.When citing this publication, authors are requested to use the following abbreviation:Bull. Alabama Mus. Nat. Hist.ISSN: 0196-1039Copyright 2012 by The Alabama Museum of Natural History

Bulletin 30 November 1, 2012Morphological Diversity and Phylogenetics of the DarterSubgenus Doration (Percidae: Etheostoma),with Descriptions of Five New SpeciesBy Steven R. Layman and Richard L. MaydenThe University of AlabamaTuscaloosa, ALabamanovember 1, 2012

Morphological Diversity and Phylogenetics of the DarterSubgenus Doration (Percidae: Etheostoma), with Descriptionsof Five New SpeciesSTEVEN R. LAYMANGeosyntec Consultants1255 Roberts Blvd NWKennesaw, GA 30144 USAslayman@geosyntec.comRICHARD L. MAYDENDepartment of Biology3507 Laclede AveSaint Louis UniversitySt. Louis, MO 63103 USAmaydenrl@slu.eduABSTRACTThe evolutionary diversity and phylogenetic relationships of darters of the subgenus Doration were investigatedusing variation in morphology and male breeding colors. A revision of the subgenus is presented, with redescriptionsof Etheostoma stigmaeum, E. jessiae, E. meadiae, descriptions of five new species, distributional data, comparisons,and a key to the species. Along with E. akatulo, a total of nine species are recognized in the subgenus.New species are described, including E. obama from the Duck River system, E. gore from the Cumberland Riverdrainage, E. jimmycarter from the Green River drainage, E. teddyroosevelt from the Arkansas and upper White riverdrainages of the Ozark Highlands, and E. clinton from the upper Ouachita River system of the Ouachita Highlands.Evidence was found that specimens from Little Bear Creek, Alabama, represent pure E. jessiae rather than hybridswith E. stigmaeum from adjacent Bear Creek. Patterns of morphological and allozyme variation suggested possibleintrogressive hybridization between E. jessiae and E. meadiae in the Clinch River system. Phylogenetic relationshipswere evaluated using 34 discrete breeding color, morphological, and behavioral characters. Doration was supportedas a monophyletic group with E. akatulo sister to a clade containing all other species in the subgenus. Relationshipswere well resolved, with the exception of a trichotomy involving new species from the Duck and Cumberland riversand the ancestor of a clade of new species from the Green and Arkansas/White rivers.Key Words: darters, Percidae, Etheostoma, Doration, new species, phylogenetic relationshipsINTRODUCTIONSpecies diversity and phylogenetic relationships ofdarters of the subgenus Doration of Etheostoma are poorlyknown. Doration was resurrected as a subgenus by Cole(1967) to include the nominal species Etheostoma stigmaeumand E. jessiae, former members of subgenus Boleosoma (Baileyand Gosline, 1955). Cole (1967) diagnosed the subgenuson the basis of the elongate, tubular, genital papilla ofbreeding females and the development of male nuptial tubercleson pelvic and anal fin rays and ventral body scales.Howell (1968) examined meristic variation in DorationBull. Alabma Mus. Nat. Hist. 30:1–83November 1, 2012

Bulletin 30 NOVEMBER 1, 2012Fig. 1. Distribution of five taxonomic forms of Doration recognized by Howell (1980a, 1980b, 1980c) as distinct species.Map provided by Division of Fishes, University of Michigan Museum of Zoology and used with permission.and treated the subgenus as the single polytypic speciesE. stigmaeum, consisting of three subspecies; however, otherscontinued to recognize E. jessiae as a separate species(Bailey et al., 1970; Robins et al., 1980). In reassessing thestatus of apparent intergrades and considering evidencesuggesting reproductive isolation between two sympatricforms, Howell (1980a, 1980b, 1980c) later recognized fivedistinct species (Fig. 1) (cited by Etnier and Starnes, 1994,as pers. comm. with Howell). These included: 1) E. stigmaeum,occurring from Gulf Coastal drainages, up the lowerMississippi River Basin, west into the Arkansas and Whiteriver drainages, and east into the lower Tennessee, Cumberland,and Green river drainages; 2) E. jessiae, endemicto the Tennessee River drainage; 3) E. meadiae of the upperClinch and Powell rivers in the upper Tennessee Riverdrainage; 4) a species endemic to Caney Fork River of theCumberland River drainage, since described as E. akatuloby Layman and Mayden (2009); and 5) an undescribedE. jessiae-like species known only from a single collectionfrom Stones River in the Cumberland River drainage.Most authorities continue to either recognize only E. stigmaeumand E. jessiae as species (Page, 1983; Robins et al.,1991; Mayden et al., 1992) or treat all taxa as subspecies ofE. stigmaeum (Etnier and Starnes, 1994; Jenkins and Burkhead,1994).Howell’s (1968) study was based mainly on preservedmuseum specimens. While he described male breedingcoloration in Mobile Basin specimens (Howell and Boschung,1966), his color descriptions of other Doration taxa(Howell, 1968) suggest he did not have fresh breeding materialfrom throughout the range of the subgenus. Colordescriptions and plates in Kuehne and Barbour (1983),Page (1983), Robison and Buchanan (1988), Burkheadand Jenkins (1991), and other sources indicate remarkablegeographic variation in breeding coloration amongtaxa, especially within the widespread nominal form E.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) largely unresolved relationships of the outgroups. Giventhe phylogenetic evidence above suggesting that E. chlorosomaand E. nigrum are more closely related to Dorationthan E. caeruleum or E. euzonum, trees were rooted usingE. caeruleum and E. euzonum. This tested the monophyly ofDoration by essentially treating E. chlorosoma and E. nigrumas part of an enlarged ingroup.Breeding male color patterns provided a rich source ofcharacters for inferring ingroup relationships. Althoughvariation in color patterns was observed intraspecifically,character states were defined conservatively to include therange of variation observed within each species. Unfortunately,breeding color characters pose difficulties for polarizationbecause ingroup character states are often notshared by outgroup taxa. In these situations polarity decisionscannot be reached, and the analysis relies heavily onthe distribution of derived states for other characters thatcan be polarized unequivocally. These characters may ultimatelydrive the polarity of initially unpolarized charactersin achieving a globally parsimonious tree topology.The characters used in the phylogenetic analyses aredescribed in Appendix A and the data matrix appearsin Appendix B. Scale and fin ray counts were generallyavoided as characters because overlapping frequencydistributions among the ingroup species make discretecharacter coding particularly difficult. Character statesfor the outgroups were obtained from Page (1981, 1983),Page and Cordes (1983), Kuehne and Barbour (1983),Bart and Cashner (1986), and Cole (1957). Two phylogeneticanalyses were performed. The first analysis treatedall multistate characters as unordered to avoid assumptionsregarding character evolution (unordered analysis).The second analysis treated multistate characters 7 and 12as ordered, based on reasoning provided in Appendix A,and all other multistate characters as unordered (mixedanalysis). Character state reconstructions for minimumlengthtrees used both ACCTRAN and DELTRAN optimizationschemes (Swofford and Maddison, 1987).All presidential photographs are public domain fromthe Clinton and Carter libraries or the Library of Congress.Species ConceptsHerein, new species of darters of the genus Etheostomaare described. Many species of darters of the genera Percina,Etheostoma, Ammocrypta, and Crystallaria are commonlydescribed and diagnosed on the basis of morphometric,meristic, and coloration differences from close relatives,as well as such seemingly minor differences between speciesas observed in alterations in the lateralis system (bodyand head) and secondary sexual color patterns in theirfins that may only be present for a few weeks of the year inbreeding adults.Our theoretical concept, of species as natural kind, isthe Evolutionary Species Concept, as outlined by Wiley(1978) and later modified by Wiley and Mayden (2000a,b, c) and further elaborated on by Mayden (1997, 1999,2002). Thus, we hypothesize that species are independentevolutionary lineages. As our surrogate nominal-kinddefinitions of a species that we employed in the discoveryand descriptions of new species as independent lineageswe employed multiple operational species concepts, mostnotably the Phylogenetic Species Concept, the MorphologicalSpecies Concept, and, given the diagnosability ofthese species on the basis of allozyme variation (Layman,1994) the Genetic Species Concept.TAXONOMIC DESCRIPTIONSSubgenus Doration JordanDoration Jordan, 1929:156 (new genus, type species Boleosomastigmaeum Jordan).Diagnosis.—Member of genus Etheostoma as diagnosedby Page (1981) and modified by Simons (1991, 1992).Breeding male distinguished by: iridescent blue or bluegreenpigment on operculum, cheek, and suborbital bar;lateral series of 7–11 iridescent blue or blue-green bars orblotches extending from humeral area to caudal peduncle;blue or blue-green bar on base of caudal fin. Breedingmale spinous dorsal fin with black, blue, or blue-greenmarginal and submedial bands; red to orange medialband. Medial caudal fin base with two small, verticallyaligned, closely spaced dark spots, most conspicuous injuveniles and nonbreeding adults (obscure on breedingmale). Dorsum with 6 dark brown quadrate to hourglassshapedsaddles. Breeding males often with tubercles onventral body scales and/or pelvic and anal fin rays. Breedingfemale genital papilla a long conical tube. Branchiostegalrays modally 6; membranes usually narrowly connected.Description.—Meristic and morphometric data for alltaxa appear in Tables 1 through 12. Lateral line completeor incomplete; lateral scale rows 38–65. Pored laterallinescales 20–55; unpored scales 0–28. Transverse scalerows 9–19. Scale rows below lateral line 5–10. Scale rowsabove lateral line 3–8. Caudal peduncle scale rows 12–22.Cheek and nape naked to fully scaled. Opercle and bellyscaled. Breast naked; prepectoral area occasionally with afew scales. Dorsal fin spines 9–14; dorsal fin soft rays 8–14.Principal caudal fin rays 12–18. Anal fin spines 2; anal finsoft rays 5–11. Pectoral fin rays 11–16. Branchiostegal rays6; membranes narrowly to moderately connected. Frenumpresent or absent. Vomerine teeth present; palatine teethpresent or absent. Infraorbital canal uninterrupted with8 pores. Supratemporal canal usually uninterrupted with3 pores. Preoperculomandibular pores 9 or 10. Lateralcanal pores 5, supraorbital canal pores 4, coronal poresingle. Vertebrae 38–42 (Bailey and Gosline, 1955). Maximumsize 35–65 mm standard length (SL), adult malestypically averaging larger than females.

Bulletin 30 NOVEMBER 1, 2012Fig. 3. Photographs of breeding males of species of Doration. (A) Etheostoma stigmaeum (Jordan, 1877), UAIC 10791.01,44 mm SL, Hurricane Creek at US Hwy 11, Tuscaloosa County, Alabama, 5 April 1993. (B) E. stigmaeum, UAIC10310.11, 48 mm SL, Big Creek at old blockaded bridge, 5.3 air km NW Pollock, Grant Parish, Louisiana, 20 March1992. (C) E. jessiae (Jordan and Brayton, 1878), UAIC 10372.01, 59 mm SL, Little Bear Creek at AL Hwy 187, 8.0 kmS Belgreen, Franklin County, Alabama, 14 March 1992. (D) E. meadiae (Jordan and Evermann, 1898), UAIC 10706.01,51 mm SL, Blackwater Creek at TN Hwy 70, 0.5 km S Virginia line, Hancock County, Tennessee, 1 April 1993. (E) E.akatulo (Layman and Mayden), 2009, UAIC 10382.02, holotype, 45.5 mm SL, Collins River between mouths of Scottand Hillis creeks, 1.6 air km SE Irving College, Warren County, Tennessee, 11 April 1992. (F) E. obama (Mayden andLayman), UAIC 10319.09, 46 mm SL, Buffalo River at Cuba Landing Rd. (200 m upstream of bridge), HumphreysCounty, Tennessee, 28 March 1992. (G) E. gore (Layman and Mayden), UAIC 10707.01, holotype, 40.3 mm SL, TurnbullCreek 0.8 km W Kingston Springs on co. rd., Cheatham County, Tennessee, 25 April 1993. (H) E. jimmycarter (Laymanand Mayden), UAIC 10708.01, 47.1 mm SL, Trammel Fork at Old State Rd., 1.6 km NNE Red Hill, Allen County,Kentucky, 25 April 1993. (I) E. teddyroosevelt (Layman and Mayden), UAIC 10460.21, holotype, 40.7 mm SL, SpringRiver at KS Hwy 96, Cherokee County, Kansas, 24 March 1991. (J) E. Clinton (Mayden and Layman), UAIC 10302.05,holotype, 33.7 mm SL, Caddo River at AR Hwy 182, 3.2 km N Amity, Clark County, Arkansas, 4 April 1992.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) Nonbreeding coloration.—Nonbreeding males and femalesexhibit little dimorphism in coloration, except asnoted. Base color of upper body tan or straw to olivaceous;small melanophores along scale edges, imparting overallsand-grained appearance. Dorsum with 6 dark brownquadrate to hourglass-shaped saddles. First saddle locatedanterior to spinous dorsal fin; second saddle just anteriorto middle of spinous dorsal fin; third saddle at posteriorend of spinous dorsal fin; fourth saddle at middle of softdorsal fin; fifth saddle posterior to soft dorsal fin; sixthsaddle at dorsal insertion of caudal fin. Sides with usually8–10 (7–11) quadrate blotches extending from lateral lineventrad 2–3 scale rows, formed by crosshatching of darkpigment along scale edges (W-, V-, and X-shaped markings);blotches may have tinge of blue. Dark X-markingsmay also occur between lateral blotches in line with ventraledges. Sides and upper body also with many smallerscattered dark brown markings. A dusky blotch may alsobe present on body just anterior to caudal fin base. Medialbase of caudal fin with two small vertically aligned,closely spaced dark spots, sometimes appearing fusedor obscured by dusky pigment; most distinct in juvenilesand nonbreeding adults. Lower body light straw to white;males may be lightly dusky. Head dark above and lightbelow, with dark preorbital bars extending onto lip butnot meeting at midline, dark suborbital bar or spot, anddark postorbital spot. Tinge of blue on operculum andpreoperculum.Male spinous dorsal fin with faint dusky marginalband, narrow clear submarginal band, red to orange medialband, dusky submedial band (with hint of blue insome species), and clear basal band with dusky areas inposterior portions of membranes. Female spinous dorsalfin mostly clear, with narrow red-orange or orange medialband; spines with scattered dark pigment. Male and femalesoft dorsal fin membranes clear; rays with 2–4 browndashes. Caudal fin membranes clear; rays with 4–5 browndashes. Anal fin membranes clear; rays with faint duskystreaks. Pelvic fin membranes mostly clear; rays with a fewdark dashes or specks. Pectoral fins mostly clear; rays withyellow-orange hue basally; rays with a few dark dashes orspecks.Breeding coloration.—Breeding males (Fig. 3) withlateral blotches becoming iridescent blue or blue-green,quadrate or vertically elongate; blue or blue-green bar developingon base of caudal fin. Sides with scattered redto orange spots and X-markings; scales between lateralblotches outlined in powder blue. Iridescent blue or bluegreenon operculum, pre-operculum, and cheek; presentor absent on lips and mid-gular area. Base color of facegray, tangerine orange, or entirely blue. Body dusky overall.Genital papilla a small dusky or pale conical flap.Spinous dorsal fin with distinctive banding coloration:thin black, blue, or blue-green marginal band; narrowwhite submarginal band; wide red to orange medial band;wide black, blue, or blue-green submedial band; clearbasal band with black or orange pigment in posteriorportions of membranes. Second dorsal, caudal, anal, andpectoral fins with or without bright orange spots. Seconddorsal and anal fins with or without basal blue or bluegreenpigment.Breeding females much more subdued, developingonly hint of male breeding coloration. Genital papilla along pale conical tube.Tuberculation.—Breeding males may develop tuberclesor tubercular ridges on pelvic and anal fin rays. Pelvicspine with overall thickened epidermis; rays 1–3 withnarrow epidermal ridges on ventral surfaces, coveringdistal half to entire length. Rays 3–5 with broken ridgesand/or small tubercles on medial-distal ventral surfaces.Anal spines and rays with narrow ridges and/or distinctrounded tubercles, usually on distal half to four-fifths;may be more weakly developed posteriorly. Ridge surfacesmay contain keratin and possibly function as breeding tubercles.Nuptial males may also develop crescent-shaped,mound-like, or rounded to pointy tubercles along posterioredges of ventral body scales. At maximum development,tubercles on belly scales, scales above anal fin base,and ventral caudal peduncle scales; development typicallyweakest on latter two regions. Variations on this basic patternof tubercle distribution and form are described underspecies accounts.Habitat.—Species of Doration are found in clear, mediumto large sized creeks and small rivers of moderategradient. All species occupy similar habitats. Adults andjuveniles occur in slow to moderate current over mixturesof sand, gravel, and occasionally silt, typically justdownstream of riffles, in moderate runs, gentle riffles, oralong margins of pools. In early spring breeding adultsoccupy gravelly runs and shallow riffles with moderatelyswift current where spawning presumably occurs. Winn(1958a, 1958b) observed spawning behavior in aquariumheldadults of the newly described species E. jimmycarterfrom the Green River. The male mounts the female in ahorizontal position, the two vibrate, and eggs are depositedin gravel. Based on the collection of nuptial malesand gravid females, spawning probably begins as early asFebruary in southern populations and terminates by lateMay or June in northern populations.Comparisons.—Doration is morphologically mostsimilar to subgenera Vaillantia and Boleosoma. Species ofall three subgenera possess 6 dorsal saddles and lateralblotches formed by X-markings, and all were once classifiedin subgenus Boleosoma (Bailey and Gosline, 1955).Species of Doration differ from those of Vaillantia and Boleosomain breeding males having bright iridescent blue orblue-green lateral body and head coloration (versus lackingin Vaillantia and Boleosoma; E. davisoni with light greeniridescence); breeding males having black, blue, or bluegreenbands separated by a bright red to orange band inthe spinous dorsal fin (vs. lacking); breeding males often

Bulletin 30 NOVEMBER 1, 2012developing tubercles on ventral body scales (vs. lacking);and breeding females having a long tubular genital papilla(vs. rugose, spatulate in Vaillantia and flat, bifurcatein Boleosoma). Species of Doration differ further in developinga pair of small dark spots on the base of the caudalfin (may be obscured in breeding males), a character alsorecognized by Douglas (1974) in E. stigmaeum. This pairof spots is a highly useful field character in distinguishingjuveniles and small adults of Doration from sympatric E.nigrum or E. chlorosoma, which develop a single irregularblotch. However, it is not useful in separating E. stigmaeumfrom E. davisoni, a species that also possesses two smallbasicaudal spots.Species of Doration differ further from those of Vaillantiaand Boleosoma in consistently having two anal spines(vs. 1 in some species). They differ further from speciesof Boleosoma in breeding males developing tubercles onpelvic and anal fin rays. They differ further from speciesof Vaillantia in preorbital bars not meeting at midline.Key to Species of Doration1. Premaxillary frenum present ........................... Go to 2Premaxillary frenum absent ............................ Go to 32. Anal fin soft rays modally 9; cheek squamation 10–30%, usually 1–18 scales on upper cheek; caudal pedunclescales usually 17 or more; principal caudalfin rays 15........................................ Etheostoma jessiaeAnal fin soft rays modally 8; cheek naked, occasionallywith 1–3 scales behind eye; caudal pedunclescales usually 16; principal caudal fin rays often 16–17........................................................... Etheostoma meadiae3. Lateral line complete; cheeks fully scaled or nearly so;breeding male with continuous blue mask of pigmentcovering lower face, snout, and underside of head............................................................. Etheostoma akatuloLateral line usually incomplete, or if complete, cheeksnaked or nearly so; breeding male with blue pigment onhead but not as continuous mask...........................Go to 44. Unpored lateral scales usually 0–7 (0–13); principal caudalfin rays usually 16–17 (70% of specimens)........................................................................... Etheostoma meadiaeUnpored lateral scales usually >10 (0–28); principalcaudal fin rays 15............................................... Go to 55. Palatine teeth present (>85% of specimens); nape nakedto fully scaled; breeding male soft dorsal and caudalfins lacking bright orange spots ................ Go to 6Palatine teeth absent (>70% of specimens); nape fullyscaled or nearly so; breeding male soft dorsal and caudalfins with discrete, often bright, orange spots .................................................................................... Go to 76. Preoperculomandibular pores 10 (>75% of specimens);anal fin soft rays modally 8; breeding malelacking thin dusky midlateral stripe running throughlateral blue-green bars ................ Etheostoma stigmaeumPreoperculomandibular pores 9 (>90% of specimens);anal fin soft rays modally 9; breeding male with thindusky midlateral stripe running through lateral bluegreenblotches.................................... Etheostoma clinton7. Breeding male with base color of face gray withmilky blue sheen, spinous dorsal fin lacking brightorange in basal band; scales below lateral line modally7; transverse scales modally 13 .................................................................................. Etheostoma teddyrooseveltBreeding male with base color of face tangerine orange,spinous dorsal fin with bright orange in basalband; scales below lateral line modally 6; transversescales modally 12 ............................................... Go to 88. Breeding male spinous dorsal fin with black submedialband, interrupted by orange streaks in posteriorportions of membranes; breeding male soft dorsaland anal fins lacking basal blue pigment ..........................................................................Etheostoma jimmycarterBreeding male spinous dorsal fin with blue in submedialband, uninterrupted by orange streaks; breedingmale soft dorsal and anal fins with basal blue pigment................................................................... Go to 99. Pectoral fin rays modally 15; scales above lateral linemodally 5; cheek squamation 10–30%, usually 2–15scales on upper cheek; soft dorsal rays modally 12;breeding male anal fin lacking orange spots or withweak orange spots at distal edge of basal blue band.................................................................Etheostoma obamaPectoral fin rays modally 14; scales above lateral linemodally 4; cheek naked or nearly so, with usually 0–5scales behind eye; soft dorsal rays modally 11; breedingmale anal fin with orange spots, usually 1–2 perray.............................................................Etheostoma goreEtheostoma stigmaeum (Jordan)Speckled DarterFigs. 3, 4Boleosoma stigmaeum Jordan, 1877:311 (original descriptionfrom small tributaries of Etowah and Oostanaula riversnear Rome, Floyd County, Georgia); Bailey et al., 1954:142 (designation of lectotype); Collette and Knapp, 1966:19 (location of lectotype and paralectotypes).Ulocentra stigmaea: Jordan and Brayton, 1878:45,82 (recordedfrom Alabama River basin; also known fromLouisiana); Jordan and Gilbert, 1883:495 (description;distributed from Georgia to Louisiana); Jordan and Evermann,1896:1047–1048 (description; distributed fromTennessee and Arkansas to Georgia and Louisiana);Fowler, 1907:522, fig. 5 (figure of syntype); Jordan, Evermann,and Clark, 1930:287 (distributed from Tennesseeand Arkansas to Georgia and Louisiana); Fowler, 1945:37,354–355, 369 (description; recorded from Alabama andMississippi river basins).

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) Poecilichthys saxatilis Hay, 1881:495 (original descriptionfrom tributary of Chickasawha River at Enterprise, ClarkeCounty, Mississippi); Jordan and Gilbert, 1883:515–516(description; known from Chickasawha River, Mississippi);Bailey et al., 1954:142 (synonym of E. stigmaeum); Colletteand Knapp, 1966 (location of holotype; synonym ofE. (Boleosoma) stigmaeum).Etheostoma (Etheostoma) saxatile: Gilbert, 1888:57–58 (description;found in Black Warrior River, Alabama, and Salineand Ouachita rivers, Arkansas).Etheostoma saxatile: Jordan, 1890:133 (distributed fromTennessee to Arkansas and south); Gilbert, 1891:150 (synonymof E. stigmaeum).Etheostoma stigmaeum: Gilbert, 1891: 150, 155, 157 (Escambia,Alabama, and Pascagoula rivers included in range;also known from Arkansas); Bailey et al., 1954: 143 (Gulftributaries and Mississippi Basin included in range); Cole,1967:28–29 (removed from subgenus Boleosoma and placedin subgenus Doration); Howell, 1980c:697 (in part; systematics;distribution and habitat; biology); Page, 1983:26, 81,238, plate 19A (in part; description; range; natural history;systematics; key; photo of breeding male from OpintolocoCr., Alabama); Kuehne and Barbour, 1983:16, 66,101–102, plate 13 (in part; description; distribution; naturalhistory; systematics; key; photo of breeding male fromCottondale Cr., Tuscaloosa County, Alabama (Gilbert andWalsh, 1991)).Doration stigmaeum: Jordan, 1929:156 (new genus: type,Boleosoma stigmaeum Jordan; description; distributed fromKentucky to Georgia and Alabama).Etheostoma (Boleosoma) stigmaeum: Bailey and Gosline,1955:15, 26, 38 (vertebral counts for specimens fromMissouri, Mississippi, Alabama, and Florida); Collette,1965:586–587 (description of breeding tubercles for specimensfrom Mobile Basin, Escambia River, and BogueChitto River); Collette and Knapp, 1966: 19 (taxonomicstatus of nominal species Boleosoma stigmaeum); Howelland Boschung, 1966:510–514 (natural intersubgeneric hybridwith E. (Oligocephalus) whipplii artesiae).Etheostoma stigmaeum stigmaeum: Burkhead and Jenkins,1991:386 (in part; found in lower Ohio and Mississippiriver basins and adjacent drainages of Gulf of Mexico; notknown from Virginia); Etnier and Starnes, 1994:533–537(in part; biology; distribution and status; systematics; key;photo of nuptial male from Conasauga River, Tennessee);Jenkins and Burkhead, 1994:838 (widespread, but inhabitsonly lower portion of Tennessee drainage).Nominal species Etheostoma stigmaeum was treated byHowell (1968) as a wide ranging subspecies, occurringfrom Gulf Coast drainages, up the lower MississippiRiver Basin, west into the Arkansas and White rivers inthe Ozark Highlands, and east into the Tennessee (DuckRiver and Bear Creek systems), Cumberland, and GreenRiver drainages. Characters used to diagnose the taxonwere lack of a frenum, usually less than 50% squamationof the cheek, and low counts of pored lateral-line scales.Two races were recognized on the basis of meristic differentiation,a widespread “typical race” and a Cumberland-Green River race, with one or both populations of the latterrace having higher modal counts of anal soft rays (9versus 8 in the typical race), pectoral rays (15 vs. 14), anddorsal spines (12–13 vs. 11). With respect to these characters,the Green River population was noted as being closerto nominal form E. jessiae than the typical race of nominalE. stigmaeum. Howell (1968) further noted that breedingmales of the Cumberland-Green River race resemblethose of nominal E. jessiae in having orange “blocks” ofpigment on the fins and orange pigment in the base ofthe spinous dorsal fin.This study indicates that nominal E. stigmaeum, as conceivedby Howell (1968), actually represents a complexof six species. Four species are recognized from Howell’stypical race, including widespread but newly restricted E.stigmaeum, a new species from the Arkansas and upperWhite rivers, a new species endemic to upper OuachitaRiver above the Fall Line, and a new species endemic toDuck River of the Tennessee drainage. Only ten specimensof the latter two species were available to Howell(1968). Howell’s Cumberland-Green River race is recognizedas two new species endemic to each of those drainages.New species from the Arkansas/upper White, Duck,Cumberland, and Green rivers all develop orange spotson the second dorsal and caudal fins, as in E. jessiae. Etheostomastigmaeum, as redescribed below, remains the mostwidely distributed species of Doration but is herein restrictedto Mobile Basin and other Gulf Coast systems, the MississippiEmbayment and highland tributaries thereof innortheastern Arkansas and southeastern Missouri, andBear Creek of the Tennessee drainage.Lectotype.–ANSP 20645, male, 39 mm SL, EtowahRiver, tributary of Coosa River near Rome, Floyd County,Georgia, summer 1876, D. S. Jordan and C. H. Gilbert;selected by Bailey et al. (1954:142).Paralectotypes.–ANSP 20646 (3; 36–39 mm SL) (originalnumbers ANSP 20646–48), same data as lectotype;one syntype figured by Fowler (1907:fig. 5).Diagnosis.—A member of the subgenus Doration. Breedingmale distinguished by: spinous dorsal fin with blueor blue-green marginal and submedial bands, red-orangemedial band, basal band lacking bright orange pigment;soft dorsal, caudal, and pectoral fins lacking distinct orangespots on rays; soft dorsal and anal fins with blue orblue-green in base of fin; face and lower head gray withblue or blue-green on operculum, preoperculum, suborbitalbar, cheek, lips, and mid-gular region; vertically

10 Bulletin 30 NOVEMBER 1, 2012Figure 4. Etheostoma stigmaeum (Jordan, 1877). Caddo R. at US Hwy 67, 5.9 km N Arkadelphia (Ouachita Riversystem), Clark County, Arkansas, 4 April 1992. UAIC 10379.01; male, 46 mm SL. Drawing by Joseph R. Tomelleri(americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.elongate lateral blue or blue-green bars; basicaudal blueor blue-green bar extending from dorsal to ventral marginof caudal fin. Frenum absent. Lateral line incomplete,unpored scales modally >10. Cheek partially scaled. Palatineteeth present. Dorsal fin spines modally 11; dorsalsoft rays modally 11. Anal fin soft rays modally 8. Pectoralfin rays modally 14. Principal caudal fin rays modally 15.Caudal peduncle scales modally 16. Preoperculomandibularcanal pores modally 10.Description.—Males average larger than females; largestmale 48.9 mm SL, largest female 42.4 mm SL. In populationsfrom four drainages, sexes exhibit significant dimorphism(P

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 11dashes. Vertical blue-green bar on base of caudal fin extendingfrom dorsal to ventral edge of fin and posteriorlyalong edges up to one-half or three-fourths the length ofthe fin; usually extending farther posteriad along ventralmargin; dorsal portion of bar may be more weakly developed.Medial base of caudal fin with two dark basicaudalspots (may be obscured by blue-green basicaudal bar).Anal fin dark gray or dusky distally, bright blue-green basally;medial-distal portions of some rays may have faintyellow-orange or gold tinge (but no distinct orange spots).Pelvic fins dark gray with blue-green at bases of medialrays. Pectoral fin membranes clear to lightly dusky; rayswith faint orange-yellow wash at base of fin.Base color of cheeks, snout, underside of head, andbreast dark gray. Iridescent blue-green on operculum,preoperculum, suborbital bar, anterior preorbital bar,lips, and mid-gular area. Lateral belly yellow or orangeyellow;medial belly gray. Base color of upper body strawto olivaceous. Sides with usually 8–10 vertical iridescentblue-green bars, several of these typically connecting dorsolaterallywith saddles in the form of wishbones. Posteriorbars may encircle caudal peduncle either ventrally ordorsally (at fifth and sixth saddles). Dusky spot or smudgeoften present on body just anterior of basicaudal spots.Scales between lateral bars outlined in powder blue andforming crosshatched pattern. Sides with red-orangespots and X-markings between lateral bars and extendingto dorsum, often quite bright. Dorsolateral area alsowith many small scattered dark markings. Dorsum with6 quadrate saddles, slightly constricted medially. Bodydusky overall. Color plates of breeding males are also presentedby Kuehne and Barbour (1983) and Page (1983).Tuberculation.—As noted by Collette (1965), fin tuberclesare usually more developed than ventral bodyscale tubercles. Pelvic fins with mostly narrow low ridgeson rays 1–3 and weak broken ridges and small individualtubercles on medial-distal portions of rays 3–5. Anal finspines and anterior rays with mostly tubercular ridges,grading into broken ridges and individual rounded tubercleson posterior rays; smaller individuals tend to showgreater development of rounded tubercles. Larger individualsoccasionally develop tubercles on ventral bodyscales. At maximum development tubercles may occuron up to 8 midventral scale rows on the posterior twothirdsof the belly, 1–2 scale rows above anal fin base, and3–5 midventral scale rows on caudal peduncle. Well-developedtubercles have been noted on specimens collectedfrom 31 January (Bogue Chitto, Pike County, Mississippi)to 22 May (Middle Fork Clarks River, Calloway County,Kentucky), with weak tubercles detected as late as 9 June(Stamp Creek, Bartow County, Georgia).Variation.—Modal fin ray counts are fairly consistentthroughout the range of E. stigmaeum, but there is considerablegeographic variation in scale counts, degreeof squamation, body size and shape, and breeding coloration.Specimens from eastern Gulf Coast systems havethe lowest mean count of lateral scales and the lowestmodal counts of transverse and caudal peduncle scales(Tables 1, 3, and 5). Specimens from Mobile Basin havethe highest mean counts of transverse scales and scalesbelow and above the lateral line (Tables 3 and 4); thosefrom Coosa River (n = 131) of the Mobile Basin have thehighest mean number of lateral scales (49.6). Within theMobile Basin, specimens from upper Tombigbee (n = 60)and lower Tombigbee (n = 58) rivers have lower modalcounts of lateral scales (43–45), transverse scales (12),and scales above (4) and below (6–7 or 6) the lateral linethan any other populations. The similarity of these countsto those in specimens from Bear Creek of the TennesseeRiver drainage (Tables 1, 3, and 4) lend further supportto the hypothesis that a portion of the Bear Creek systemwas captured from the adjacent upper Tombigbee Riversystem (Wall, 1968; Starnes and Etnier, 1986). Also withinMobile Basin, specimens from the Black Warrior Riversystem (n = 60) have the lowest modal cheek squamation(0%), lowest modal nape squamation (10%), and lowestmodal counts of pectoral fin rays (13) (compare with Tables6, 7, and 9). In eastern and western tributaries of theupper Mississippi Embayment, 24–33% of specimens haveprepectoral scales versus 10% or less in all other populations.Additional breakdown of meristic data by river systemis provided by Howell (1968).Maximum adult size is greatest in the Coosa and Tallapoosariver systems of Mobile Basin. Specimens fromCoosa River commonly exceed 45 mm SL, the largestspecimen being a 48.9–mm SL male (n = 131); the largestspecimen from Tallapoosa River (n = 60) was a 46.0–mmSL male. Maximum size in all other drainages ranges fromabout 39 to 45 mm SL. Proportional measurements fromfour drainages (Table 12) indicate that specimens fromCoosa River also tend to have a longer snout and upperjaw, shorter soft dorsal fin, and a deeper caudal peduncle.Breeding coloration varies mainly in the amount ofblue-green pigment in the base of the anal fin. Breedingmales from Mobile Basin typically have a wide blue-greenbasal band at peak development. Those from eastern andcentral Gulf Coast systems and much of the western MississippiEmbayment often have only a hint of blue-greenin the base of the fin. Hues of blue-green coloration varyfrom turquoise blue to almost green, but no clear geographictrends have been identified.Distribution.—Etheostoma stigmaeum occurs in GulfCoast drainages from Pensacola Bay in Florida and Alabamawest to the Red-Atchafalaya and Sabine river systemsin Louisiana (Fig. 5), but excluding the Mermentauand Calcasieu rivers (Douglas, 1974). Contrary to the speciesaccount in Kuehne and Barbour (1983), it does occurin Escambia River in Alabama and Florida as well as PondCreek, a tributary of Blackwater River just to the east inthe Florida panhandle. The species is distributed northup the Mississippi River Embayment to western Kentuckyand southeastern Missouri. West of the Mississippi River

12 Bulletin 30 NOVEMBER 1, 2012Figure 5. Distribution of nine species of Doration. Type localities of the five new species are indicated by stars. Starsand dots show localities of material examined. Some dots represent more than one locality. Species are identifiedin legend on map. Map provided by Division of Fishes, University of Michigan Museum of Zoology and used withpermission.it occurs in the Ouachita River below the Fall Line, lowerWhite River as far upstream as (and including) BuffaloRiver, Little Red River, Black River, St. Francis River, CastorRiver, and lowland drainage ditches of southeasternMissouri. To the east E. stigmaeum occurs in direct tributariesof the Mississippi River from southeastern Louisianato the Hatchie River in southwestern Tennessee. Inthe Tennessee drainage it occurs in Clarks River and JonathanCreek in western Kentucky, and Bear Creek in northwesternAlabama.Records of E. stigmaeum from the Sabine River systemin Louisiana could not be verified. The five lots examinedwere misidentified specimens of either E. chlorosoma(NLU 14752,57493,57497; UT 91.1377) or E. whipplei(NLU 56431). The single lot examined by Howell (1968;NLU 4439, formerly NLSC 4439) could not be locatedbut was presumably identified correctly, as all five specimenspossessed 2 anal spines, ruling out the likelihoodthat they could have been E. chlorosoma (1 anal spine).Comments.—Howell’s (1968) original concept of E.stigmaeum as a polytypic species was largely based on hisinterpretation of a small contact zone between nominalE. stigmaeum and E. jessiae in the Bear Creek system of theTennessee River drainage in northwestern Alabama. Sixof 12 specimens he examined from Little Bear Creek possesseda frenum, which he considered the most distinctivecharacter separating E. stigmaeum (frenum absent) andE. jessiae (frenum present). He therefore concluded thatthese were intergrades, and as such, provided evidence ofincomplete reproductive isolation between the two taxa.However, our analysis of meristic, breeding color, and allozymedata discussed below under “CONTACT ZONES”indicates that the Little Bear Creek population actuallyrepresents “pure” E. jessiae.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 13Etheostoma jessiae (Jordan and Brayton)Blueside DarterFigs. 3, 6Poecilichthys jessiae Jordan and Brayton in Jordan, 1878:227,and in Jordan and Brayton, 1878:59 (description fromChickamauga River at Ringgold, Georgia); Jordan,1880:227 (description; distributed in Tennessee River);Jordan and Gilbert, 1883:518–519 (description; knownfrom Chickamauga River, Georgia); Jordan, 1884:227 (description;distributed in Tennessee River; Kuhne, 1939:92(known from Tennessee); Fowler, 1945:39, 251 (distributedin Alabama, Tennessee, and Sabine rivers, based inpart on misidentifications ).Etheostoma (Etheostoma) saxatile: Gilbert, 1888:57–58 (description;distribution includes tributaries of Clinch Rivernear Clinton, Tennessee).Etheostoma jessiae: Jordan, 1890: 133 (description; distributedfrom Tennessee to Wabash Valley, Illinois and eastTexas, based in part on misidentifications); Jordan and Evermann,1896:1084 (description; distributed from Indianato Iowa and south to Mississippi and Texas, based largelyon confusion with E. asprigene and E. swaini); Evermann,1918:317, 319, 359, 364, 368 (identification of species reportedas P. jessiae by Jordan and Brayton (1878) fromChickamauga River at Ringgold, Georgia); Cole, 1967:28–29 (removed from subgenus Boleosoma and placed in subgenusDoration); Howell, 1980a:656 (systematics, distribution,habitat, biology); Page, 1983:26, 80, 238, plates 18Gand 18H (description; range; natural history; key; photosof breeding male and female from Little Pigeon River, SevierCounty, Tennessee); Kuehne and Barbour, 1983:16,100, plate 13 (description; distribution; natural history;photo of breeding male from West Branch Shoal Cr., LawrenceCounty, Tennessee (Gilbert and Walsh, 1991)).Etheostoma stigmaeum: Gilbert, 1891:150, 152 (TennesseeRiver drainage included in range; recorded from CypressCreek, Florence, and Big Nance Creek, Courtland, Alabama).Ulocentra stigmaea: Evermann and Hildebrand, 1916:449–450 (recorded from Ball Creek, tributary of Big SycamoreCreek near Tazewell, Tennessee, and Arnwine SpringCreek near Athens, Tennessee; compared with specimensfrom Wolf and Obeys rivers; two forms represented); Evermann,1918:320, 321, 326, 356, 367 (identification of speciesrecorded as E. (Etheostoma) saxatile by Gilbert (1888)from tributaries of Clinch River near Clinton, Tennessee;identification of species recorded as E. stigmaeum by Gilbert(1891) from Cypress and Big Nance creeks, Alabama;identification of species recorded as E. saxatile by Gilbertand Swain, 1884, unpublished, from Bull Run at Hershells,Tennessee, and Clinch River at Clinton, Tennessee).Oligocephalus jessiae: Jordan et al., 1930:291 (distributedfrom southern Illinois to Georgia and Mississippi, basedin part on misidentifications).Etheostoma (Boleosoma) jessiae: Bailey and Gosline, 1955:15, 38 (vertebral counts); Collette, 1965:570, 583, 585–586, 608 (description of breeding tubercles; systematics);Collette and Knapp, 1966:63, figure 4 (location of typematerial of nominal species Poecilichthys jessiae unknown;figure of possible syntype).Etheostoma stigmaeum jessiae: Burkhead and Jenkins,1991:385–387 (description; Virginia distribution and status;habitat; life history; recommendations); Etnier andStarnes, 1994:533–537 (biology; distribution and status;systematics; key; photo of female from Little River, Tennessee);Jenkins and Burkhead, 1994:838–840 (systematics;description; biology; habitat; distribution).Syntypes.—Number and dispostion unknown, ChickamaugaRiver at Ringgold, Georgia, summer 1877, D. S.Jordan and A. W. Brayton, assisted by C. H. Gilbert anda party of students from Butler University (Jordan andBrayton, 1878). Jordan and Brayton (1878) described thespecies from “Several specimens, each about two incheslong, .... “ Collette and Knapp (1966:63, fig. 4) failed tolocate any type material but found a drawing by ErnestCopeland in the files of the USNM Division of Fishes labeled“Poecilichthys jessiae, Chickamauga River, Ga,” Theyreproduced the drawing, believing that it probably representsone of the lost syntypes.Two papers describing this species were published inthe same year, but that of Jordan (1878) apparently preceded,or was intended to precede, by date that of Jordanand Brayton (1878). The latter paper references the formerand indicates species authorship as Jordan and Brayton,rather than providing the notation “sp. nov.” Robins et al.(1991:89) cited Jordan (1878) as the original description,pointing to an error in Jordan and Evermann (1896:1085),which gives the publication date for the species as 1877.The paper by Jordan and Brayton (1878) provides a moredetailed description and includes provenance of the typespecimens, information lacking in Jordan (1878). In fact,Jordan and Evermann (1896:1085) duplicate the accountfrom Jordan and Brayton (1878) as the “original descriptionof Poecilichthys jessiae.” We have been unable to determinewhich of the 1878 papers was actually published firstthat year.Diagnosis.—A member of the subgenus Doration.Breeding male distinguished by: spinous dorsal fin withblue marginal and submedial bands, orange medialband, basal band with bright orange pigment; soft dorsal,caudal, and pectoral fins with bright orange spots onrays; soft dorsal and anal fins with blue in base of fin; faceand lower head gray with blue on operculum, preoperculum,suborbital bar, and lips (but not mid-gular region);vertically elongate lateral blue bars; basicaudal blue bar

14 Bulletin 30 NOVEMBER 1, 2012Figure 6. Etheostoma jessiae (Jordan and Brayton, 1878). Coal Creek along Tennessee Hwy 116, 2.1 air km SW LakeCity, Morgan Co., Tennessee, 12 April 1992. UAIC 10380.01; male, 53 mm SL. Drawing by Joseph R. Tomelleri (americanfishes.com).Copyright by Joseph R. Tomelleri. Used with permission.extending from dorsal to ventral margin of caudal fin.Frenum usually present. Lateral line incomplete, unporedscales modally 8. Cheek partially scaled. Palatineteeth present. Dorsal fin spines modally 12–13; dorsal softrays modally 12. Anal fin soft rays modally 9. Pectoral finrays modally 14–15. Principal caudal fin rays modally 15.Caudal peduncle scales modally 17–18. Preoperculomandibularcanal pores modally 10.Description.—Males average larger than females; largestmale 64.6 mm SL, largest female 56.1 mm SL. Sexesexhibit significant dimorphism (P

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 15lum, preoperculum, and suborbital bar; no blue on midgulararea. Lateral belly yellow; medial belly gray. Basecolor of upper body straw to olivaceous or gray. Sides withusually 9–10 (8–11) vertical iridescent blue bars. Scalesbetween lateral bars outlined in powder blue. Sides withorange markings along scale edges between lateral barsand extending to dorsum; orange becoming brighterposteriorly on caudal peduncle. Dorsolateral area alsowith many small dark brown and blue markings. Dorsumwith 6 prominent dark brown or gray hourglass-shapedsaddles. Johnson (1987) provides a color plate of a livebreeding male photographed by W. N. Roston. Colorplates are also provided by Page (1983) and Kuehne andBarbour (1983).Tuberculation.—Pelvic and anal fin tuberculation primarilyconsists of epidermal ridges, which are often welldeveloped. Pelvic rays 3–5 and last few anal rays occasionallywith broken ridges and only rarely with a few individualdistinct tubercles. As also noted by Collette (1965),ventral body scale tubercles are often well developed. Atmaximum development, tubercles may occur on 2 midventralscale rows behind pelvic fin bases, 11 midventralscale rows at mid-belly, 9 midventral scale rows at anus,1–4 scale rows above anal fin, and 5–6 midventral scalerows on caudal peduncle. Well-developed tubercles havebeen noted on specimens collected from 3 February (ElkRiver, Grundy County, Tennessee) to 9 May (Lick Creek,Greene County, Tennessee), with weak tubercles detectedas early as 26 January (Spring Creek, Polk County, Tennessee)and as late as 21 May (Factory Creek, LawrenceCounty, Tennessee).Variation.—The most notable morphological characterexhibiting geographic variation is the premaxillaryfrenum. Eighty-eight percent of all specimens examined(n = 575) possessed a narrow to broad frenum. In the upperTennessee drainage, including the Clinch River systemboth above and below Norris Dam, this proportion is86–98% (n = 405). In contrast, 76.5% of specimens (n =170) from the lower and middle Tennessee drainage, includingLittle Bear Creek, have a usually narrow to moderatefrenum. Howell (1968; 1980a) treated specimensfrom Little Bear Creek as intergrades or hybrids betweenE. jessiae and E. stigmaeum (from adjacent Bear Creek) becauseonly six of 12 available specimens (50%) possesseda frenum. However, examination of 49 specimens fromLittle Bear Creek found that 69% of specimens possessa frenum. A few other populations of E. jessiae in themiddle Tennessee drainage exhibit similar frequenciesof individuals possessing a frenum but there is no cleargeographic cline suggestive of a hybrid or intergradezone. Moving up the Tennessee River from Bear Creek,proportions of specimens with a frenum are 100% in CypressCreek (n = 10), 73% in Shoal Creek (n = 26), 100%in Bluewater Creek (n = 10), 88% in Elk River (n = 16),65% in Paint Rock River (n = 23), and 73% in SequatchieRiver (n = 26). An alternative explanation to Howell’s(1968) hypothesis of hybridization, one that is supportedby additional meristic, breeding color, and allozymic datadiscussed below under “CONTACT ZONES,” is that thepolymorphic condition of the frenum has been retainedfrom the ancestor of E. jessiae.Specimens of E. jessiae from the French Broad Riversystem have higher scale counts than any other populationsexamined. Mean counts (and ranges) for 60 specimenswere 53.4 (46–60) lateral scales, 15.5 (13–19) transversescales, 8.1 (7–10) scales below the lateral line, 6.2(5–8) scales above the lateral line, and 18.9 (16–22) caudalpeduncle scales (compare with Tables 1, 3, 4, and 5).This observation led to the determination that the singlerecord of E. jessiae from Stones River of the Cumberlanddrainage (CU 46558), thought by Howell (1980a) to representan undescribed species similar to E. jessiae, wasbased erroneously on specimens of E. jessiae from LittlePigeon River of the French Broad River system (Layman,1994). The largest specimen of E. jessiae is also from theFrench Broad River system: a 64.6–mm SL male from theNolichucky River system (UT 91.1209). Elsewhere maximumadult size is about 57 mm SL.Modal pectoral fin ray counts in E. jessiae are 15throughout the Tennessee drainage, with the exceptionof Clinch River populations, where the mode is 14. Thisshift may due to introgressive hybridization with E. meadiae(Etnier and Starnes, 1994), which also has a mode of 14and occurs farther upstream in the Clinch River system(see “CONTACT ZONES” below).Distribution.—Etheostoma jessiae is endemic to the TennesseeRiver Drainage, occurring from White Oak Creek,Houston and Humphreys counties, Tennessee, upstream,but not including Duck River, through Alabama and Tennesseeinto the French Broad River in North Carolinaand the Holston River in Virginia (Fig. 5). In the BearCreek system of Alabama, E. jessiae occurs only in the LittleBear Creek tributary system and is parapatric with E.stigmaeum, which is restricted to upper Bear Creek and itstributaries; the two species have not been taken together.The historic range of E. jessiae extends upstream in theClinch and lower Powell rivers to at least three tributariesof Norris Reservoir, including (Fig. 7): Cove Creek,downstream of the Clinch and Powell rivers confluence;a “pond opposite Doak’s Dam” (UMMZ 103591), presumablyin the Davis Creek system of the lower Powell River(possibly in the Davis Creek embayment); and Big SycamoreCreek, a tributary of the Clinch River in the upperend of the reservoir. The species was last collected in Coveand Davis creeks in 1936–37, shortly after completion ofNorris Dam, and may no longer occur there due to impoundmentof habitat. Etheostoma jessiae is parapatric withE. meadiae, which occurs upstream of Norris Reservoir inthe Clinch and Powell rivers. Specimens from PossumCreek (CU 68500) and Cove Creek (UMMZ 130756) ofNorth Fork Holston River, Scott County, Virginia, wereverified as E. jessiae; Jenkins and Burkhead (1994) had

16 Bulletin 30 NOVEMBER 1, 2012Figure 7. Distribution of Etheostoma jessiae (solid dots) and E. meadiae (dots with stars) in the Clinch and Powell riversof the upper Tennessee River drainage. Dots indicate localities of materials examined; some represent more thanone locality. Streams are: A—upper Clinch River; B—Powell River; C—Norris Reservoir; D—lower Clinch River;E—Tennessee River (Watts Bar Reservoir); 1a—Emory River; 1b—Poplar Creek; 2a—Beaver Creek; 2b—Bull RunCreek; 3—Hinds Creek; 4—Coal Creek; 5—Cove Creek; 6—Big Sycamore Creek; 7—pond opposite Doak’s dam(Davis Creek).noted possible problems with the identifications of thesespecimens. Three specimens from Mills River and SouthFork Mills River of the French Broad River system in HendersonCounty, North Carolina, the only records of thespecies from that state, were not examined but were verifiedby W. M. Howell (Menhinick et al., 1974): the specimens(originally cataloged at Duke University) could notbe located (W. M. Palmer, pers. comm.).Etheostoma meadiae (Jordan and Evermann)Bluespar DarterFigs. 3, 8Ulocentra meadiae Jordan and Evermann, 1898:2852(original description from Indian Creek, basin of PowellRiver, Cumberland Gap, Tennessee); Jordan and Evermann,1900:fig. 447 (figure); Evermann and Hildebrand,1916:450 (known from Indian Creek, tributary of PowellRiver near Cumberland Gap); Evermann, 1918:330, 331,356, 365, 367 (known from Indian Creek, tributary ofPowell River near Cumberland Gap).Etheostoma stigmaeum: Gilbert, 1891:150 (Tennessee Riverdrainage included in range); Page, 1983:81, 238 (upperClinch and Powell rivers included in distribution).Imostoma meadiae: Jordan et al., 1930:286 (known from IndianCreek, Powell River basin, eastern Tennessee).Doration meadiae: Kuhne, 1939:92 (known from Tennessee).Ulocentra mediae: Fowler, 1945:251 (compared with Poecilichthyshopkinsi; species epithet misspelled).Cottogaster mediae: Fowler, 1945:37 (distributed in TennesseeRiver drainage; species epithet misspelled).Etheostoma (Boleosoma) jessiae: Collette and Knapp, 1966:72(senior synonym of nominal species Ulocentra meadiae; locationof types of U. meadiae).Etheostoma meadiae: Howell, 1980b:666 (previously consideredintergrade population; distributed in upper Powelland Clinch river systems, Virginia and Tennessee; habitat,biology); Mayden et al., 1992:859 (nominal species warrantingadditional study to determine taxonomic status).Etheostoma stigmaeum meadiae: Burkhead and Jenkins,1991:386–387, plate 149 (breeding male color descrip-

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 17tion and plate; Virginia distribution); Etnier and Starnes,1994:534–537 (biology; distribution and status; systematics;key); Jenkins and Burkhead, 1994: 838–840, fish 290–291, plate 28 (systematics; description; breeding malecoloration; biology; habitat; distribution; halftone photos;breeding male color plate).Etheostoma meadiae (Jordan and Evermann) was resurrectedby Howell (1980b), who found the name availablefor a distinctive form of Doration from the upper Clinchand Powell rivers in the upper Tennessee drainage.Howell (1968) had earlier treated these populations asintergrades between what he considered nominal E. stigmaeumfrom the Cumberland drainage (described as anew species below) and nominal E. jessiae. He based thisassessment on the observation that 53% of specimenspossessed a frenum, which he considered the most distinctivecharacter separating the taxa, and present-daydrainage patterns suggestive of stream capture betweenthe Cumberland and Powell rivers.Examination of additional specimens later convincedHowell (1980b) that these apparent intergrades actuallyrepresent a distinct taxon (Etnier and Starnes, 1994). Hisrecognition of E. meadiae and all other taxa of Doration asdistinct species (Howell, 1980a, 1980b, 1980c) was bolsteredby the apparent reproductive isolation of sympatricforms of E. stigmaeum and E. jessiae in the Stones Riverof the Cumberland drainage (Etnier and Starnes, 1994,in pers. comm. with Howell). Howell’s (1980a, 1980b,1980c) conclusions were not accompanied by supportingdata. Consequently, E. meadiae has failed to gain recognitionas a species. Furthermore, the single record of E. jessiaefrom the Stones River has been shown to be invalid,having resulted from an error in cataloging specimensfrom Little Pigeon River in the Tennessee River drainage(Layman, 1994).Etheostoma meadiae is currently treated as a subspeciesor race of a polytypic E. stigmaeum by Etnier andStarnes (1994). Jenkins and Burkhead (1994) also treatit as a subspecies but hold out the possibility of it beingan intergrade. Starnes and Etnier (1986) pointed to thelack of evidence for a major stream capture between theCumberland and Powell rivers, weakening support forHowell’s intergradation hypothesis (Etnier and Starnes,1994). Patterns of nuptial male coloration are also inconsistentwith intergradation. Males of both supposed parentaltaxa develop orange pigment in the base of thespinous dorsal fin and bright orange spots on the softdorsal, caudal, and pectoral fins. These features are lackingin E. meadiae.Morphological and breeding color data gathered inthis study indicate that E. meadiae warrants recognitionas a distinct species. Moreover, the phylogenetic hypothesisgenerated using these data suggests that E. meadiaeand E. jessiae are not even sister taxa. Etheostoma meadiae istreated herein as a distinct species restricted to the upperClinch and Powell rivers upstream of Norris Reservoir inTennessee and Virginia.Holotype.—USNM 48903, male, 46 mm SL, IndianCreek, tributary of Powell River, Cumberland Gap, Tennessee,17 October 1893, R. R. Gurley. The holotype isfigured in Jordan and Evermann (1900:fig. 447).Paratypes.—USNM 125623, male, 46 mm SL, samedata as holotype, original number BF 711 (U. S. Bureauof Fisheries). According to Collette and Knapp (1966),the third type was apparently sent to Stanford University,but it was not listed among the types there by Bohlke(1953); they were unable to locate it.Figure 8. Etheostoma meadiae (Jordan and Evermann, 1898). Blackwater Creek at TN Hwy 70, 0.5 km S Virginia line,Hancock County, Tennessee, 1 April 1993. UAIC 10706.01; male, 51 mm SL. Drawing by Joseph R. Tomelleri (americanfishes.com).Copyrighted by Joseph R. Tomelleri. Used with permission.

18 Bulletin 30 NOVEMBER 1, 2012Diagnosis.—A member of the subgenus Doration.Breeding male distinguished by: spinous dorsal fin withblue marginal and submedial bands, orange medialband, basal band lacking bright orange pigment; softdorsal, caudal, and pectoral fins lacking discrete orangespots on rays (although diffuse yellow-orange streaksand dashes may be present); soft dorsal and anal finswith blue in base of fin; face and lower head gray withprofuse blue on operculum, preoperculum, cheeks, suborbitalbar, ventrolateral head, and lips (but not mid-gularregion); vertically elongate lateral blue bars; basicaudalblue bar extending from dorsal to ventral margin ofcaudal fin. Frenum present or absent. Lateral line usuallyincomplete with

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 19Figure 9. Etheostoma akatulo Layman and Mayden, 2009. Collins River between mouths of Scott and Hillis creeks, 1.6air km SE Irving College, Warren County, Tennessee, 11 April 1992. UAIC 10382.02; holotype, male, 45.5 mm SL.Drawing by Joseph R. Tomelleri (americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.Variation.—The disjunct Powell and Clinch Riverpopulations vary mainly in the proportion of individualspossessing a frenum and counts of dorsal fin soft rays.Thirty-seven percent of specimens from Powell Riverhave a usually narrow frenum, while 67% of those fromClinch River have a usually narrow or moderate frenum.Mean soft dorsal ray counts are 11.1 in Powell River and11.5 in Clinch River.Distribution.—Etheostoma meadiae occurs only in thePowell and Clinch River systems of the Tennessee Riverdrainage upstream of Norris Reservoir in Tennesseeand Virginia (Figs. 5 and 7). Populations in the two riversystems are isolated by Norris Reservoir, with E. jessiaeoccurring in intervening tributaries of Norris Reservoir(Fig. 7). The two species are distributed parapatrically.Etheostoma akatulo Layman and MaydenBluemask DarterFigs. 3, 9Etheostoma akatulo was diagnosed and described byLayman and Mayden (2009). It is endemic to the upperCaney Fork River system of the Cumberland River drainage,Tennessee. Meristic counts for 203 total specimensare summarized in Tables 1 through 10. This rare speciespresently occurs in only four isolated tributaries of GreatFalls Reservoir in the eastern Highland Rim physiographicprovince, including Collins River, Rocky River, CaneCreek, and upper Caney Fork River (Fig. 5); it formerlyoccurred in the Calfkiller River.

20 Bulletin 30 NOVEMBER 1, 2012Etheostoma obama Mayden and Layman, New speciesSpangled DarterFigs. 3, 10Figure 10. Etheostoma obama, new species Mayden and Layman. Buffalo River at Cuba Landing Rd. (200 m upstreamof bridge), Humphreys County, Tennessee, 28 March 1992. UAIC 10319.09; male, 46 mm SL. Drawing by Joseph R.Tomelleri (americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.SynonymyEtheostoma stigmaeum: Gilbert, 1891:150 (Tennessee Riverdrainage included in range); Howell, 1980c:697 (DuckRiver system included in distribution map); Page, 1983:81,238 (Duck River system included in distribution map);Kuehne and Barbour, 1983:101 (Duck River system includedin range map);Etheostoma stigmaeum stigmaeum: Etnier and Starnes,1994:534 (Duck and Buffalo river systems included in distribution).Holotype.—UAIC 10337.29, breeding male, 42.7 mmSL, Duck River below dam at TN Hwy 64/US Hwy 231 inShelbyville, Bedford County, Tennessee, 13 April 1991, S.R. Layman and A. M. Simons.Paratypes.—UAIC 10337.27 (30; 26.1–43.4 mm SL),USNM 328259 (6; 33.0–42.6), same data as holotype; UT91.1538 (22; 31.5–41.9), Duck R. at end of unnumberedco. rd., 4.0 air km SE Chapel Hill, Marshall County, Tennessee,23 March 1978, Beets, N. M. Burkhead, J. L Harris,J. Louton, D. L Nieland, and M. G. Ryon; NLU 52970(3; 41.7–43.5), Duck R. at Hooper Island, river km 259,Maury County, Tennessee, 23 June 1979, J. Feeman, C.Saylor, et al.; UAIC 9874.32 (33; 33.1–46.0), SIUC 22876(5; 36.8–45.4), TU 167869 (5; 34.9–41.8), UF 100288 (5;34.1–42.9), UMMZ 225160 (5; 35.5–45.1), Buffalo R. atmouth of Grinders Cr. and TN Hwy 99, Lewis County,Figure 11. Barack Hussein Obama II, 44 th President ofthe United States of America

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 21Tennessee, 26 May 1990, S. R. Layman, R. M. Wood, andB. R. Kuhajda; INHS 79399 (13; 29.1–37.9) same localitydata as preceding collection, 14 April 1978, L. M. Pageand R. L. Mayden; UAIC 10319.09 (6; 32.7–45.9), BuffaloR. at Cuba Landing Rd. (200 m upstream of bridge),Humphreys County, Tennessee, 28 March 1992, S. R. Laymanand C. M. Bertram.Diagnosis.—A member of the subgenus Doration.Breeding male distinguished by: spinous dorsal fin withblue marginal and submedial bands, orange medial band,basal band with bright orange pigment; soft dorsal, caudal,and pectoral fins with distinct bright orange spotson rays; soft dorsal and anal fins with blue in base of fin;anal fin sometimes with orange spots or streaks on rays;face and lower head tangerine orange with blue on operculum,preoperculum, cheek, and suborbital bar (not onlips and mid-gular region); quadrate lateral blue blotchesextending ventrad from lateral line scale row; basicaudalblue bar extending from dorsal to ventral margin of caudalfin. Frenum absent. Lateral line incomplete, unporedscales modally >10 (12). Cheek partially scaled. Palatineteeth usually absent; present in 29% of specimens. Dorsalfin spines modally 12–13; dorsal soft rays modally 12. Analfin soft rays modally 9. Pectoral fin rays modally 15. Principalcaudal fin rays modally 15. Caudal peduncle scalesmodally 16. Preoperculomandibular canal pores modally10. Allozyme products of sIdh-A locus characterized byrelative mobilities b, e, and g (Layman, 1994).Description.—Males average larger than females; largestmale 48.3 mm SL, largest female 42.9 mm SL. Sexesexhibit significant dimorphism (P

22 Bulletin 30 NOVEMBER 1, 2012dividual tubercles on medial-distal portions of rays 3–5.Anal fin spines and anterior rays usually with tubercularridges, grading into broken ridges and individual roundedtubercles on posterior rays; smaller individuals tend todevelop short ridges and rounded tubercles on most rays.Ventral body scale tubercles typically not as well developed,occurring on up to 4 midventral scale rows on posteriorhalf of belly. Tuberculate males have been found incollections from 7 March to 20 May (Duck River, Marshalland Bedford counties, Tennessee).Distribution.—This species is endemic to the Duck andBuffalo Rivers of the Tennessee River drainage, Tennessee(Fig. 5). In the Duck River it occurs below NormandyDam in the Nashville Basin and western Highland Rimdownstream past the confluence of Buffalo River to thebackwaters of Kentucky Lake. It occurs in the BuffaloRiver, located wholly on the western Highland Rim, fromupper reaches downstream to its mouth. Etheostoma obamais distributed parapatrically with E. jessiae, which occurs inTennessee River tributaries upstream, and in at least onetributary downstream (Whiteoak Creek), of Duck River.Etymology.—The common name spangled darter refersto the bright orange spots adorning the body and finsof breeding males. The species epithet is a noun in appositionthat honors President Barack Obama (Fig. 11),the 44 th President of the United States of America, andhis environmental leadership and commitment duringchallenging economic times in the areas of clean energy,energy efficiency, environmental protection and humanitarianefforts globally, and especially for the people of theUnited States.Comments.—Only five specimens from the Duck Riversystem were available to Howell (1968), and presumablybecause they lacked a frenum he treated them as nominalE. stigmaeum. Etnier and Starnes (1994) examined 46 additionalspecimens from the system and reported modalcounts of dorsal fin spines, dorsal fin soft rays, anal finrays, and pectoral fin rays that are closer to those of E.jessiae. They suggested possible intergradation betweennominal E. stigmaeum, invading from the CumberlandRiver (recognized herein as a new species), and E. jessiae.Examination of 185 specimens, observations of breedingcolors in the upper and lower reaches of the system, andsurvey of allozyme variation (Layman, 1994) indicate thatthe population from the Duck River system is distinguishableand diagnosable as a species. The phylogenetic hypothesispresented below suggests that meristic similaritiesbetween E. obama and E. jessiae may best be explainedby more recent common ancestry of E. obama with E. jessiaethan with nominal E. stigmaeum.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 23Etheostoma gore Layman and Mayden, New speciesCumberland DarterFigs. 3, 12Figure 12. Etheostoma gore, new species Layman and Mayden. Turnbull Creek 0.8 km W Kingston Springs on co. rd.,Cheatham County, Tennessee, 25 April 1993. UAIC 10707.02; male, 40.3 mm SL. Drawing by Joseph R. Tomelleri(americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.SynonymyEtheostoma stigmaeum: Gilbert, 1891:150 (CumberlandRiver drainage included in range); Kirsch, 1893:262, 265,268 (recorded from Smith Fork of lower Caney Fork River,Obeys River, Eagle Creek, Wolf River, Willis Creek and fourtributaries of Big South Fork of the Cumberland River);Bailey et al., 1954:143 (Cumberland drainage included inrange); Comiskey and Etnier, 1972:143 (reported from BigSouth Fork of the Cumberland River): Clay, 1975:361–362(Cumberland drainage, Kentucky, included in distribution);Howell, 1980c:697 (Cumberland drainage includedin distribution): Page, 1983:81,238 (Cumberland drainageincluded in distribution); Kuehne and Barbour, 1983:101,plate 13 (Cumberland drainage included in range; photoof breeding male from North Fork Rockcastle River, JacksonCounty, Kentucky (Gilbert and Walsh, 1991)); Burrand Warren, 1986:322 (Cumberland drainage included indistribution).Ulocentra stigmaea: Evermann, 1918:330, 356, 367 (identificationof species recorded as E. stigmaeum by Kirsch (1893)from the Cumberland River; reported from Stones Rivernear Nashville, Gilbert and Swain, 1884, collectors).Etheostoma stigmaeum stigmaeum: Etnier and Starnes,1994:534 (Cumberland drainage, excluding Caney Fork,included in distribution).Holotype.—UAIC 10707.02, breeding male, 40.3 mmSL, Turnbull Creek 0.8 km W Kingston Springs on co. rd.,Figure 13. Albert Arnold “Al” Gore, Jr., 45 th Vice-President of the United States of America. Courtesy :William J. Clinton Presidential Library.

24 Bulletin 30 NOVEMBER 1, 2012Cheatham County, Tennessee, 25 April 1993, S. R. Laymanand E. B. Jones.Paratypes.—UAIC 10707.01 (14; 33.6–38.9 mm SL),same data as holotype; UAIC 9863.26 (19; 29.8–37.9),USNM 328260 (4; 34.0–38.8), UT 91.4452 (4; 35.3–40.5),SIUC 22877 (4; 35.9–37.3), same locality data as holotype,21 May 1990, S. R. Layman, B. R. Kuhajda, and R.M. Wood; UMMZ 175059 (10; 29.7–37.8), Red R. on staterd. just SE Keysburg, Kentucky, Robertson County, Tennessee,21 June 1957, C. R. Gilbert and F. A. Gilbert; CU37282 (12; 33.5–42.8), East Fork Stones R. at US Hwy 231,9.6 km N Murfreesboro, Rutherford County, Tennessee,26 March 1961, W. J. Richards, N. R. Foster, and L. W.Knapp; INHS 87340 (4; 37.4–39.4), East Fork Stones R. 1.6km NW Readyville, Rutherford County, Tennessee, 6 May1981, M. E. Retzer and L. M. Page.Diagnosis.—A member of the subgenus Doration. Breedingmale distinguished by: spinous dorsal fin with bluemarginal and submedial bands, orange medial band, basalband with bright orange pigment; soft dorsal, caudal, andpectoral fins with distinct bright orange spots on rays; softdorsal and anal fins with blue in base of fin; anal fin withdistinct orange spots on rays; face and lower head tangerineorange with blue on operculum, preoperculum, cheek,and suborbital bar (not on lips and mid-gular region);quadrate lateral blue blotches extending ventrad from lateralline scale row; basicaudal blue bar extending from dorsalto ventral margin of caudal fin. Frenum absent. Lateralline incomplete, unpored scales modally>10 (11). Cheeknaked or nearly naked. Palatine teeth absent. Dorsal finspines modally 12; dorsal soft rays modally 11. Anal fin softrays modally 8–9. Pectoral fin rays modally 14. Principalcaudal fin rays modally 15. Caudal peduncle scales modally16. Preoperculomandibular canal pores modally 10.Locus sIdh-A fixed for allele f (Layman, 1994).Description.—Males average larger than females; largestmale 42.8 mm SL, largest female 41.2 mm SL. Sexesexhibit significant dimorphism (P

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 25hourglass-shaped saddles. A color plate of the breedingmale of this species also appears in Kuehne andBarbour (1983).Tuberculation.—Pelvic fins with usually narrow lowridges on rays 1–3 and small rounded tubercles on medial-distalportions of rays 3–5. Anal fin spines and rays1–2 or 1–4 usually with tubercular ridges; all other raysdominated by distinct rounded tubercles. Tubercles mayalso develop on ventral body scales. At maximum development,rounded to crescent-shaped tubercles occur on upto 7 midventral scale rows on posterior two-thirds of belly,1–2 scale rows above anal fin base, and 3- 4 midventralscale rows on caudal peduncle. Tuberculate males havebeen found in collections from 15 March (RockcastleRiver, Rockcastle County, Kentucky) to 24 May (East ForkStones River, Rutherford County, Tennessee), with weaktubercles detected as late as 13 June (Buck Creek, PulaskiCounty, Tennessee).Distribution.—Etheostoma gore occurs in the CumberlandRiver drainage below Cumberland Falls from RockcastleRiver in Kentucky downstream to Red River in Kentuckyand Tennessee (Fig. 5). It appears to be absent fromCaney Fork River, a major southern tributary in MiddleTennessee, but specimens of E. stigmaeum reported byKirsch (1893) from lower Caney Fork River (dispositionunknown) probably represented this species rather thanE. akatulo, which is endemic to the system above GreatFalls (Layman et al., 1993; Layman and Mayden, 2009).Etheostoma gore was once thought to have been sympatric inEast Fork Stones River with an undescribed E. jessiae-likeform (Howell, 1968; 1980a), but the single record of thelatter was shown to be invalid as a result of a catalogingerror (Layman, 1994).Etymology.— The common name Cumberland darterrefers to the species’ endemic distribution in the CumberlandRiver drainage. The species epithet is a noun in appositionthat honors Al Gore, the 45 th Vice President ofthe United States of America (Fig. 13), serving with PresidentBill Clinton, and his environmental vision, commitment,and accomplishments throughout decades of publicservice and his role in educating the public and raisingawareness on the issue of global climate change.

26 Bulletin 30 NOVEMBER 1, 2012Etheostoma jimmycarter Layman and Mayden, New speciesBluegrass DarterFigs. 3, 14Figure 14. Etheostoma jimmycarter, new species Layman and Mayden. Little Barren R. at KY Hwy 70, Sulphur Well,Metcalfe County, Kentucky. 15 March 1990. UAIC 9852.14; male, 47 mm SL. Drawing by Joseph R. Tomelleri (americanfishes.com).Copyrighted by Joseph R. Tomelleri. Used with permission.SynonymyEtheostoma stigmaeum: Woolman, 1892:260, 288 (reportedfrom Big Barren River, Drake Creek, Little Barren River,and Green River, Kentucky); Bailey et al., 1954:143 (GreenRiver drainage included in range); Bailey and Gosline,1955:15, 26, 38 (vertebral counts for specimens from PetersCreek, Barren River system, Kentucky); Collette, 1965:586–587 (description of breeding tubercles includes specimensfrom East Fork Barren and Green rivers, Kentucky); Clay,1975:361–362 (Green River drainage included in distribution);Howell, 1980c:697 (Green River drainage included indistribution); Page, 1981:9, 10, 12, 15, 33, 47, 53, 58, 63, 68(specimens from Barren River, Kentucky used in pheneticand cladistic analyses of darter subgenera; morphologicalcharacters); Page, 1983:81, 238, plate 19B (Green Riverdrainage included in distribution map; photo of femalefrom West Fork Drakes Creek, Sumner County, Tennessee);Kuehne and Barbour, 1983:101 (Green River drainage includedin distribution); Burr and Warren, 1986:322 (GreenRiver drainage included in distribution).Ulocentra stigmaea: Evermann, 1918:356,367 (identificationof species reported as E. stigmaeum by Woolman (1892) fromGreen River Basin, Kentucky).Etheostoma (Doration) saxatile: Winn, 1958a:191–192, 197,209 (reproductive habits); Winn, 1958b:156, 159, 160–161,164–165, 167–170, 172–173, 176–177, 179–180, 182, 185–188(reproductive habits; systematic significance).Figure 15. James Earl “Jimmy” Carter, Jr., the 39 thPresident of the United States of America

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 27Etheostoma stigmaeum stigmaeum: Etnier and Starnes, 1994:534(Barren and Green river systems included in distribution).Holotype.—UAIC 10708.01, breeding male, 47.1 mm SL,Trammel Fork at Old State Rd., 1.6 km NNE Red Hill, AllenCounty, Kentucky, 25 April 1993, S. R. Layman and E.B. Jones.Paratypes.—UAIC 10708.02 (34; 31.3–41.6 mm SL),SIUC 22878 (10; 32.6–46.5), same data as holotype; UAIC9853.19 (46; 31.5–44.1), INHS 32134 (10; 32.8–42.3), UF100289 (10; 35.9–43.5), USNM 328261 (10; 33.5–43.5),same locality data as holotype, 16 March 1990, S. R. Layman,R. L. Mayden, B. R. Kuhajda, and R. M. Wood; NLU18876 (32; 29.2–51.8), Trammel Fk. 32 km E Franklin onKY Hwy 100, Allen County, Kentucky, 9 April 1971, B. Wallusand D. Wallus; TU 81925 (23; 27.7–41.8), UT 91.780 (45;30.0–44.8), Middle Fork Drakes Cr. 1.2 km upstream of KYHwy 265, 4.3 km SE Gold City, Allen County, Kentucky, 14April 1973, Etnier, Hoyt, Oakberg, Taylor, Thompson, Stiles;UMMZ 165344 (38; 28.7–47.7), East Fork Barren R. 8.0 kmNW Tompkinsville, Monroe County, Kentucky, 6 April 1953,R. M. Bailey, D. Bailey, H. E. Winn, J. Keleher.Diagnosis.—A member of the subgenus Doration. Breedingmale distinguished by: spinous dorsal fin with blackmarginal band, orange medial band, black submedial bandinterrupted by vertical orange streaks extending from medialband above to basal band, basal band with bright orangepigment; soft dorsal, caudal, and pectoral fins withdistinct bright orange spots on rays; soft dorsal and analfins lacking blue pigment; anal fin with many distinct orangespots on rays; face and lower head tangerine orangewith blue on operculum, preoperculum, cheek, and suborbitalbar (not on lips and mid-gular region); quadratelateral blue blotches extending ventrad from lateral linescale row; blue wedge on ventral half of caudal fin basebut not developed dorsally. Frenum absent. Lateral lineincomplete, unpored scales modally >10 (14). Cheek usuallynaked. Palatine teeth absent. Dorsal fin spines modally12–13; dorsal soft rays modally 11. Anal fin soft raysmodally 8. Pectoral fin rays modally 15. Principal caudalfin rays modally 15. Caudal peduncle scales modally 16.Preoperculomandibular canal pores modally 10.Description.—Males average larger than females; largestmale 49.0 mm SL, largest female 46.5 mm SL. Sexesexhibit significant dimorphism (P

28 Bulletin 30 NOVEMBER 1, 2012ridges on rays 1–3 and weak broken ridges and roundedtubercles on medial-distal portions of rays 3–5. Anal finspines and anterior rays usually with tubercular ridges,giving way to broken ridges and individual rounded tubercleson posterior rays; some specimens with roundedtubercles on all rays. Posterior belly scales occasionallywith weak tubercles, observed on up to 3 midventral scalerows. Winn (1958b) also reported tubercles from the pelvicand anal fins. Tuberculate males have been taken incollections from 10 March (tributary of Russell Creek,Adair County, Kentucky) to 24 May (Little Barren River,Green County, Kentucky).Distribution.— Etheostoma jimmycarter is endemic to theGreen River drainage of Kentucky and Tennessee, occurringmainly in the Highland Rim physiographic province(Fig. 5). It is distributed widely in the upper Barren andupper Green rivers, and is also found in the upper RoughRiver, a tributary of lower Green River (Burr and Warren,1986). Specimens from Rough River have not been examined.Etymology.— The name bluegrass darter refers to therange of the species lying mostly within Kentucky, nicknamedthe “Bluegrass State.” The species epithet is a nounin apposition that honors President Jimmy Carter (Fig.15), the 39 th President of the United States of America,and his environmental leadership and accomplishmentsin the areas of national energy policy and wilderness protection,and his life-long commitment to social justice andbasic human rights.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 29Etheostoma teddyroosevelt Layman and Mayden, New speciesHighland DarterFigs. 3, 16Figure 16. Etheostoma teddyroosevelt, new species Layman and Mayden. Shoal Creek at KS Hwy 26 at SchermerhornPark, 3.2 km S Galena, Cherokee County, Kansas, 24 March 1991. UAIC 10072.15; male, 47 mm SL. Drawing byJoseph R. Tomelleri (americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.SynonymyEtheostoma stigmaeum: Bailey et al., 1954:143 (eastern Oklahomaand southern Missouri included in range); Baileyand Gosline, 1955:15, 26, 38 (vertebral counts for specimensfrom Illinois River, Oklahoma); Cross and Minckley,1958:104, 107–108 (reported from Shoal Creek andSpring River, Kansas; description; nuptial coloration;habitat); Collette, 1965:586–587 (description of breedingtubercles includes specimens from Spring River, Kansas);Cross, 1967: 303–304 (distributed in Spring River andShoal Creek, Kansas; description; male breeding coloration;habitat; figure); Branson, Triplett, and Hartmann,1969:457, 467 (common in the Neosho River system; possiblyentered Neosho River via headwater exchange fromWhite River); Miller and Robison, 1973:214–215 (northeasternOklahoma included in distribution; description;halftone plate); Pflieger, 1975:309–310 (southwesternOzarks included in distribution); Howell, 1980c:697 (Arkansasand upper White river drainages included in distributionmap); Page, 1983:81, 238 (Arkansas and upperWhite river drainages included in distribution map); Kuehneand Barbour, 1983:101 (Arkansas and upper Whiteriver drainages included in range map); Robison and Buchanan,1988:441–442 (Arkansas and upper White Riverdrainages included in distribution map; photo of breedingmale from upper White River drainage, Missouri (W.N. Roston, pers. comm.)).Figure 17. Theodore “Teddy” Roosevelt, the 26 thPresident of the United States of America

30 Bulletin 30 NOVEMBER 1, 2012Etheostoma saxatile: Metcalfe, 1959:393 (Spring River, Kansasas westernmost record of species).Etheostoma stigmaeum stigmaeum: Etnier and Starnes,1994:534 (Arkansas and White river systems of southwesternMissouri included in range).Holotype.—UAIC 10460.21, breeding male, 40.7 mmSL, Spring River at KS Hwy 96, Cherokee County, Kansas,24 March 1991, S. R. Layman, B. R. Kuhajda, A. M.Simons, R. M. Wood.Paratypes.—UAIC 10460.16 (5; 32.0–42.6 mm SL),USNM 328262 (4; 34.8–43.4), UT 91.4453 (4; 35.1–42.9),same data as holotype; KU 3597 (9; 34.1–36.8), same localitydata as holotype, 7 April 1956, F. B. Cross and class;UAIC 10072.15 (7; 29.1–42.1), INHS 32135 (4; 34.9–43.2),SIUC 22879 (4; 34.9–42.1), Shoal Cr. at KS Hwy 26 atSchermerhorn Park, 3.2 km S Galena, Cherokee County,Kansas; TU 92739 (3; 31.5– 38.9), Clear Cr. at Savoy,tributary to Illinois R., below dam, Washington County,Arkansas, 9 May 1970, Strawn and Galloway; NLU 48970(6; 29.8–42.2), Illinois R. at US Hwy 62, Cherokee County,Oklahoma, 28 November 1981, G. Varney, K. Kessler, E.Grissom, and J. Wise; UMMZ 137865 (19; 32.6–39.9), IllinoisR. near mouth of Swimmer’s Cr., Sequoyah County,Oklahoma, 12 April 1941, Oklahoma A&M UniversityField Class.Diagnosis.—A member of the subgenus Doration.Breeding male distinguished by: spinous dorsal fin withblack marginal and submedial bands, orange medialband, basal band lacking bright orange pigment; soft dorsal,caudal, and pectoral fins with subdued but usually discreteorange spots on rays; soft dorsal and anal fins lackingblue pigment; anal fin with subdued orange spots onrays; face and lower head dark gray with milky blue sheen;deep blue on operculum, preoperculum, cheek, and suborbitalbar (not on lips and mid-gular region); quadratelateral blue blotches extending ventrad from lateral linescale row; blue wedge on ventral half of caudal fin basebut not developed dorsally. Frenum absent. Lateral lineincomplete, unpored scales modally >10 (16). Cheek usuallynaked. Palatine teeth usually absent; present in 16%of specimens. Dorsal fin spines modally 11–12; dorsal softrays modally 11. Anal fin soft rays modally 8. Pectoral finrays modally 14. Principal caudal fin rays modally 15. Caudalpeduncle scales modally 16. Preoperculomandibularcanal pores modally 10.Description.—Males average larger than females; largestmale 43.4 mm SL, largest female 41.5 mm SL. Sexes exhibitsignificant dimorphism (P

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 31operculum, preoperculum, and suborbital bar. Breast darkgray. Belly and lower sides gray. Upper body dark straw toolivaceous or gray. Sides with 8–10 iridescent deep bluequadrate blotches extending from lateral line ventrad 2–4scale rows. Smaller dark spot or blotch often present onbody just anterior of basicaudal spots. Lower sides withpowder blue iridescence along scale edges. Sides with orangecoloration on scales between lateral blotches and extendingto dorsum. Dorsolateral area also with scattereddark black and blue markings. Dorsum with 6 dark grayto black saddles, slightly constricted medially. Body oftenquite dark. Cross (1967) also provides a detailed descriptionof nuptial male coloration.Tuberculation.—Pelvic fins with mostly narrow lowridges on rays 1–3 and weak broken ridges and roundedtubercles on medial-distal portions of rays 3–5. Anal finspines and anterior rays usually with tubercular ridges,giving way to broken ridges and individual rounded tubercleson posterior rays; some specimens with roundedtubercles on all rays. Posterior one-third to two-thirds ofbelly occasionally with oblong to crescent-shaped tubercles,occurring on up to 6 midventral scale rows (Collette,1965). Cross (1967) also noted tubercles on the pelvic andanal fins. Tuberculate males have been taken in collectionsfrom 24 March (Spring River, Cherokee County, Kansas)to 22 May (King’s River, Madison County, Arkansas).Variation.—Specimens from the Arkansas River (n =103) and upper White River (n = 69) vary mainly in napesquamation and the frequency of palatine teeth. Napesquamation is usually 90–100% (60–100) in Arkansas Riverspecimens but varies widely in those from upper WhiteRiver (10–100%). Palatine teeth are present in only 7%of Arkansas River specimens but occur in 29% of upperWhite River specimens.Distribution.—Etheostoma teddyroosevelt occurs in theArkansas and upper White river drainages on the OzarkPlateau of Missouri, Arkansas, extreme southeastern Kansas,and northeastern Oklahoma (Fig. 5). In the ArkansasRiver it occurs mainly in northern tributary systems fromthe Neosho River system downstream to Illinois Bayou.The species is also found in the Petit Jean and Fourche LaFave river systems, southern tributaries along the northernedge of the Ouachita Mountains. In the White River drainage,the species occurs upstream of Bull Shoals Dam.Etymology.— The common name highland darter reflectsthe high fidelity of this species to streams drainingthe Ozark Highlands. The species epithet is a noun in appositionthat honors President Theodore Roosevelt, the26 th President of the United States of America (Fig. 17),and his enduring legacy in environmental conservationand stewardship, including the designation of vast areasas national forests, wildlife refuges, national monuments,and national parks, and his efforts to forge the AmericanMuseum of Natural History, New York.

32 Bulletin 30 NOVEMBER 1, 2012Etheostoma clinton Mayden and Layman, New speciesBeaded DarterFigs. 3, 18Figure 18. Etheostoma clinton, new species Mayden and Layman. Caddo River at Arkansas Hwy 182, 3.2 km N Amity,Clark County, Arkansas, 4 April 1992. UAIC 10302.09, holotype, 33.7 mm SL. Drawing by Joseph R. Tomelleri(americanfishes.com). Copyrighted by Joseph R. Tomelleri. Used with permission.SynonymyEtheostoma stigmaeum: Meek, 1891:139 (scarce in Ouachita,South Fork Ouachita, and Caddo rivers); Dewey andMoen, 1978:42 (reported as uncommon in Caddo River);Harris and Douglas, 1978:58 (reported from the mountainprovince of the Ouachita River); Howell, 1980c:697(upper Ouachita River included in distribution map);Page, 1983:81, 238 (upper Ouachita and upper Caddorivers included in distribution map); Kuehne and Barbour,1983:101 (Ouachita highland area included in rangemap); Robison and Buchanan, 1988:442 (upper Ouachitaand upper Caddo rivers included in distribution map).Etheostoma stigmaeum stigmaeum: Etnier and Starnes,1994:534 (Ouachita highland area included in range).Holotype.—UAIC 10302.09, breeding male, 33.7 mm SL,Caddo River at AR Hwy 182, 3.2 km N Amity, Clark County,Arkansas, 4 April 1992, S. R. Layman and M. A. Layman.Paratypes.—UAIC 10302.05 (4; 28.4–32.8 mm SL),same data as holotype; NLU 18733 (40; 26.5–34.3), UMMZ225161 (6; 27.6–32.3), USNM 328263 (6; 29.1–31.3), samelocality data as holotype, 3 April 1971, D. Fruge, J. Lindley,H. Wimberly, D. Hill, J. Mulina, P. Hambrick, and T.Guidroz; UT 91.1497 (5; 27.0–31.6), Ouachita R. at Mc-Guire Public Access Area, 3.2 km S AR Hwy 88, PolkCounty, Arkansas, 27 December 1978, J. L. Harris; UAIC10063.13 (2; 29.5–29.8), Ouachita R. downstream of Co.Rd. 67 and at Mill Cr., 1.6 km S Cherry Hill, Polk County,Arkansas, 28 March 1991, S. R. Layman, B. R. Kuhajda, A.M. Simons, and R. M. Wood; NLU 58202 (27; 25.7–30.8),Figure 19. William Jefferson “Bill” Clinton, the 42 ndPresident of the United States of America

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 33Ouachita R. and tributary bridge on graded rd., 0.8 kmS AR Hwy 88 and 1.6 km E Pine Ridge, T2S, R27W, S9,Montgomery County, Arkansas, 8 November 1985, J. Herrockand J. Halk; UT 91.1553 (5; 26.4– 31.7), South ForkOuachita R. at AR Hwy 379, ca. 4.0 rd. km W Mount Ida,Montgomery County, Arkansas, 1 April 1978, J. L. Harris,D. L. Nieland, and M. G. Ryon.Diagnosis.—A member of the subgenus Doration.Breeding male distinguished by: spinous dorsal fin withblue-green marginal and submedial bands, red-orangemedial band, basal band lacking bright orange pigment;soft dorsal, caudal, and pectoral fins lacking distinct orangespots on rays; soft dorsal and anal fins with bluegreenin base of fin; face and lower head gray with blue orblue-green on operculum, preoperculum, suborbital bar,lips, and mid-gular region; quadrate lateral blue-greenblotches with narrow continuous midlateral band of melanophoresrunning through blotches; blue-green wedgeon ventral half of caudal fin base but not developed dorsally.Frenum absent. Lateral line incomplete, unporedscales modally >10 (14). Cheek usually naked. Palatineteeth present. Dorsal fin spines modally 11; dorsal softrays modally 11. Anal fin soft rays modally 9. Pectoral finrays modally 13–14. Principal caudal fin rays modally 15.Caudal peduncle scales modally 16. Preoperculomandibularcanal pores modally 9.Description.—Largest male 34.6 mm SL, largest female34.1 mm SL. Sexes exhibit significant dimorphism(P

34 Bulletin 30 NOVEMBER 1, 2012scattered dark markings. Dorsum with 6 dark quadratesaddles, not medially constricted; first saddle may appearless discrete due to incomplete scalation of nape.Additional coloration notes.—In preservative, as theblue-green pigment of breeding males fades, the series ofquadrate lateral blotches becomes difficult to distinguishfrom the midlateral dusky band. Hence, preserved breedingmales may appear to have a continuous wide band alongthe sides rather than individual blotches. The narrow duskymidlateral band is also present in breeding females buttends to be less wide and less consistent in continuity thanin males; the lateral blotches are discernible in life or preservative.Tuberculation.—Pelvic fins with weak rounded tubercleson medial-distal portions of rays 1–4. Anal fin typicallywith distinct rounded to conical tubercles on all rays andoccasionally the second anal spine. Posterior two-thirds ofbelly occasionally with minute rounded tubercles, occurringon up to 8 midventral scale rows. Tuberculate maleshave been taken between 25 March and 4 April from CaddoRiver, Montgomery and Clark counties, Arkansas.Variation.— The disjunct populations of the upperOuachita and upper Caddo rivers vary in several meristicfeatures. Specimens from upper Ouachita River (n = 57)have modally 14 pectoral rays versus 13 in those from upperCaddo River (n = 47). Specimens from Caddo River tendto have higher scale counts, with modally 14–15 transversescales (versus 12–13 in Ouachita River), modally 8 scalesbelow the lateral line (vs. 6–7), modally 5 scales above thelateral line (vs. 4), and modally 17 caudal peduncle scales(vs. 16).Distribution.—Etheostoma clinton is known only from theupper Caddo and upper Ouachita rivers upstream of theFall Line in the Ouachita Mountains province of Arkansas(Figs. 5 and 20). In the Caddo River it is found upstreamof DeGray Reservoir. In the Ouachita River it occurs upstreamof Lake Ouachita in the upper Ouachita and SouthFork Ouachita rivers. The species may also occur in tributariesof lakes Hamilton and Catherine, which are situatedabove the Fall Line. Harris and Douglas (1978) surveyedseveral tributaries of these impoundments but did not collectthe species (then known as E. stigmaeum). Robison andBuchanan (1988) show one record of E. stigmaeum fromLittle Mazarn Creek, a tributary to Lake Hamilton, but weFigure 20. Distribution of Etheostoma clinton in the upper Ouachita River system, Arkansas. Type locality indicatedby dot with star.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 35have not examined the material to determine if it representsthe new species. The occurrence of E. pallididorsumin a nearby tributary of Lake Hamilton (Robison, 1974)suggests that the Little Mazarn Creek record may indeedrepresent E. clinton because the overall ranges of these twospecies are highly congruent (Mayden, 1985; Robison andBuchanan, 1988), but this needs to be confirmed.Populations of E. clinton from the upper Caddo and upperOuachita rivers are isolated from each other by severalman-made impoundments and the intervening CoastalPlain portion of the Ouachita River system (includinglower Caddo River), which is occupied by E. stigmaeum.Etheostoma clinton has a small and nearly identical distributionwith that of Noturus taylori, with the exception thatthe latter is also known from the headwaters of the LittleMissouri River system (Robison and Harris, 1978). Severalother fish species endemic to this region also have largelyoverlapping distributions with that of E. clinton (Mayden,1985). A status survey is needed to more clearly define thedistribution of E. clinton, particularly in tributaries of themain rivers and impoundments, and should include theupper Little Missouri River system above Lake Greeson.Etymology.— The name beaded darter refers to thedistinctive midlateral dusky band that runs through theturquoise blue blotches, suggesting a string of beads. Thespecies epithet is a noun in apposition that honors PresidentBill Clinton, the 42 nd President of the United Statesof America (Fig. 19), and his lasting environmental accomplishmentsin creating and expanding national monuments,preserving millions of acres of wilderness areas, hisleadership and commitment during challenging economictimes, and his continued commitment to global humanitarianissues and needs and peace.COMPARISONSMeristics and breeding colors.—Meristic and breedingcolor comparisons are provided in Table 13 and Figure 3.Etheostoma akatulo is highly distinctive, differing from allother species in usually having a complete lateral line, fullyor nearly fully scaled cheeks, and breeding males withbright cobalt or royal blue pigment continuously coveringthe lower face, and soft dorsal and anal fins lacking orangeand blue pigment. Etheostoma jessiae is the only species consistentlyhaving a frenum, although its presence is morevariable in some systems of the middle Tennessee drainage;a frenum is present in 37–67% of specimens of E.meadiae. Etheostoma jessiae has the highest counts of lateral,caudal peduncle, and transverse scales of all species andattains the greatest maximum size. The species differs furtherfrom E. stigmaeum, E. meadiae, E. akatulo, and E. clintonin breeding males having distinct, bright orange spots onthe second dorsal, caudal, and pectoral fin rays, and havingbright orange pigment in the base of the spinous dorsalfin. Etheostoma jessiae differs further from E. stigmaeumin having modally 12–13 dorsal spines (versus 11 in E. stigmaeum),modally 12 dorsal soft rays (vs. 11), modally 9 softanal rays (vs. 8), modally 14–15 pectoral rays (vs. 14), modally7–8 scales below the lateral line (vs. 6–7), modally 5–6scales above the lateral line (vs. 4–5), usually 1–14 unporedlateral-line scales (vs. 8–19), and a more completely andconsistently scaled nape. Breeding males of E. jessiae differfurther from those of E. stigmaeum in lacking blue pigmenton the mid-gular region.In addition to characters described above, Etheostoma jessiaediffers from E. meadiae in having modally 9 anal rays(vs. 8 in E. meadiae), modally 7–8 scales below the lateralline (vs. 6), modally 14–15 pectoral rays (vs. 14), modally15 principal caudal rays (vs. 16–17), usually 1–14 unporedlateral-line scales (vs. 0–7), partly scaled cheeks (vs. usuallynaked), and palatine teeth usually present (vs. present inonly 33% of specimens). The nape is more completely andconsistently scaled in E. jessiae than E. meadiae.Etheostoma jessiae is similar to E. obama, E. gore, and E.jimmycarter in breeding males having orange spots on thesoft dorsal, caudal, and pectoral fin rays, and orange pigmentin the base of the spinous dorsal fin. Etheostoma jessiaediffers from these three species in breeding males havingthe base color of the face and underside of the head gray(versus tangerine orange), blue pigment typically presenton the lips (vs. absent), and orange spots lacking on theanal fin (vs. present or sometimes present). Additionally,E. jessiae has higher counts of scale rows below and abovethe lateral line.Etheostoma jimmycarter differs from E. obama and E. gorein breeding males having a black submedial band in thespinous dorsal fin (vs. blue), soft dorsal and anal fins lackingblue pigment in the base, and the basicaudal blue barmainly developed ventrally, not extending to the dorsalmargin of the caudal fin. Etheostoma obama differs from E.gore in having modally 15 pectoral rays (vs. 14 in E. gore),modally 12 soft dorsal rays (vs. 11), modally 5 scales abovethe lateral line (vs. 4), partly scaled cheeks (vs. usually naked),and palatine teeth present in 29% of specimens (vs.palatine teeth absent). Breeding males of E. obama differfrom those of E. gore in often lacking orange spots on theanal fin. These two species are very similar in morphologyand breeding colors; however, patterns of allozyme variationsuggest that no gene flow is occurring between them.The two species, each represented by two populationsamples, have no alleles in common at the sIdh-A locus(Layman, 1994). In addition, the occurrence of uniquederived alleles in both samples of E. gore (Mpi-Aa and Pk-Bd) and the occurrence of a unique derived allele in theDuck River sample of E. obama (Pnp-Ae) suggest that thespecies are maintaining separate identities and are pursuingtheir own evolutionary tendencies and fates (Wiley,1978).Etheostoma stigmaeum differs from E. meadiae in havingusually 8–19 unpored lateral-line scales (vs. 0–7 in E.meadiae), partly scaled cheeks (vs. cheeks usually naked),

36 Bulletin 30 NOVEMBER 1, 2012palatine teeth present (vs. palatine teeth present in only33 % of specimens), modally 11 dorsal spines (vs. 12),modally 4–5 scales above the lateral line (vs. 6), and modally15 principal caudal rays (vs. 16–17). Although manyaspects of breeding coloration are similar, E. stigmaeumdiffers further from E. meadiae in having blue-green pigmenton the mid-gular region.Etheostoma stigmaeum differs from E. obama, E. gore, andE. jimmycarter in having modally 11 dorsal spines (vs. 12–13) and palatine teeth usually present (vs. usually absentor present in only 29% of specimens). Breeding males ofE. stigmaeum differ from those of the other three speciesin having soft dorsal, caudal, and pectoral fins lacking orangespots, the base of the spinous dorsal fin lacking orangepigment, the base color of the face and underside ofhead gray (vs. tangerine orange), and the lateral blotchesvertically elongate (vs. quadrate).Etheostoma clinton is distinguished from all other speciesof Doration by having 9 preoperculomandibular pores (vs.10 in all other species), smallest maximum adult size (40 mm SL), and breeding males having anarrow midlateral band of melanophores connecting theseries of blue-green blotches. It is most similar to E. stigmaeumin overall appearance but differs further from thatspecies in having modally 9 anal soft rays (vs. 8), modally13–14 pectoral rays (vs. 14), usually naked or nearly nakedcheeks (vs. usually partly scaled), and a higher meancount of lateral scales (49.9 vs. 46.4).Etheostoma teddyroosevelt is similar to E. obama, E. gore, andE. jimmycarter in that breeding males have distinct orangespots on the second dorsal, caudal, and anal fins; however,in E. teddyroosevelt they are more subdued and occasionallyobscured by melanophores. Etheostoma teddyroosevelt differsfrom these three species in having breeding males lackingorange pigment in the base of the spinous dorsal finand having the base color of the face gray (vs. tangerineorange), and in having modally 7 scales below the lateralline (vs. 6). It differs further from E. obama and E. gorein breeding males having black marginal and submedialbands in the spinous dorsal fin (vs. blue), soft dorsal andanal fins lacking blue pigment, and the basicaudal bluebar mainly developed ventrally. It differs further from E.jimmycarter in having modally 14 pectoral rays (vs. 15), ahigher mean count of lateral scales (48.1 vs. 45.4), andbreeding males with a solid black submedial black band(vs. interrupted with vertical orange streaks).Etheostoma teddyroosevelt differs from E. stigmaeum and E.clinton in breeding males having black marginal and submedialbands in the spinous dorsal fin (vs. blue-green),deep blue lateral blotches (vs. blue-green), orange spotson the soft dorsal, caudal, and anal fins (vs. lacking), andin lacking blue pigment on the lips and mid-gular region(vs. present). It differs further from these two species inusually lacking palatine teeth (although present in up to29% of upper White River specimens) and having a morecompletely and consistently scaled nape. Etheostoma teddyrooseveltdiffers further from E. clinton in having a fullyscaled belly (vs. incompletely scaled anteriorly) and differsfurther from E. stigmaeum in having usually naked cheeks(vs. partly scaled).Principal component analysis of 17 meristic variablesin 3,049 specimens shows a high degree of overlap amongthe nine species of Doration (Fig. 21). This, in part, illustrateswhy the systematics of the group has been so poorlyunderstood without fuller knowledge of breeding colorvariation. The analysis does not incorporate data on thefrenum or palatine teeth, characters that are also usefulin distinguishing species. Etheostoma stigmaeum, the geographicallymost widespread species, encompasses thegreatest amount of meristic variation, both along principalcomponent one (PC-I) and PC-II (Fig. 21). The PC-Iaxis provides nearly complete separation between E. jessiaeand E. akatulo. Variables loading most heavily on PC-Iinclude transverse scales, caudal peduncle scales, scalesabove and below the lateral line, and lateral scales (Table14). Other variables loading heavily are several fin raycounts and opercle, nape, and cheek squamation. The PC-II axis provides little distinction between species from theTennessee (E. jessiae, E. meadiae, E. obama), Cumberland(E. akatulo and E. gore), and Green (E. jimmycarter) riverdrainages but largley polarizes this group from E. clinton(Fig. 21). The plot completely separates both E. jessiaeand E. akatulo from E. clinton. Variables with the highestloadings on PC-II are nape and belly squamation (Table14). Others loading heavily include scales below the lateralline, unpored lateral scales, anal fin rays, pectoral finrays, transverse scales, dorsal fin spines, and cheek squamation.The greatest overlap occurs between E. meadiae,E. obama, E. gore, and E. jimmycarter (Fig. 21).Morphometrics.—Comparison of proportional measurementsin Table 12 and in Layman and Mayden (2009)reveals that E. jessiae has a longer snout than any other speciesof Doration. In addition, E. jessiae has relatively greaterhead length, predorsal length, upper jaw length, and softdorsal fin base length than most other species. Etheostomameadiae has a moderate snout but shares with E. jessiae arelatively long head, predorsum, and upper jaw. It differsfrom E. jessiae and most other species in having a deeperbody at the spinous and soft dorsal fin origins, deeper andshorter caudal peduncle, wider body, and greater analspine length. Etheostoma stigmaeum from the Coosa Riveralso has a moderately produced snout, long upper jaw,deep body at the soft dorsal fin origin, and deep caudalpeduncle. Etheostoma teddyroosevelt is also relatively robust,with greater body width and body depth at the spinousdorsal fin origin than all other species except E. meadiae.Etheostoma obama has proportionally larger fins thanmost other species of Doration as indicated by mean measurementsof spinous dorsal fin origin to soft dorsal fin origin,dorsal spine length, soft dorsal fin base length, analfin base length, and pectoral and pelvic fin lengths (Table12; Layman and Mayden, 2009: table 1). Etheostoma gore has

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 37arates E. meadiae and E. akatulo (Fig. 22A). In addition,E. clinton separates almost completely from E. stigmaeumof the Ouachita River (Fig. 22B); these samples are fromgeographically close localities in the Ouachita River systemabove and below the Fall Line, respectively. Amongpopulations of E. stigmaeum, the least overlap occurs betweenthe Ouachita and Coosa rivers, the geographicallymost distant samples (Fig. 22B).The plot of sheared PC-II versus sheared PC-III forfemale measurements (Fig. 23, Table 15) reveals similartrends in shape differentiation as observed in males, withE. jessiae completely separated from E. meadiae, E. obama,Figure 21. Plot of meristic principal component one(PC-1) and PC-II for nine species of Doration. Polygonsbound all individuals of: 1—Etheostoma stigmaeum;2—E. jessiae; 3—E. meadiae; 4—E. akatulo; 5—E. obama;6—E. gore; 7—E. jimmycarter; 8—E. teddyroosevelt; 9—E.clinton.a large spinous dorsal fin, and it and E. jimmycarter haverelatively long pectoral and pelvic fins. Etheostoma akatulois a slender species of Doration with a moderate snout,narrow body depth at the spinous dorsal fin origin, longand narrow caudal peduncle, narrow trans-pelvic width,and short pectoral and pelvic fins. Within Doration, the diminutiveE. clinton has the shortest snout, narrowest bodydepth at the spinous and soft dorsal fin origins, shortestdistance between the spinous and soft dorsal fin origins,narrowest trans-pelvic width, and narrowest body.Sheared principal component analysis of 28 morphometricvariables reveals substantial overlap among malesof species of Doration in overall body form and fin size, butseveral species can be distinguished from one another onthis basis. The plot of sheared PC-II against sheared PC-IIIcompletely or nearly completely separates E. jessiae fromall other species except E. akatulo, with which it overlapsonly moderately (Fig. 22A). Most notably, E. jessiae separatescompletely from E. meadiae and E. obama, parapatricneighbors in the Tennessee River drainage. All or most ofthis separation occurs along sheared PC-II. Variables loadingheavily on this axis are first anal spine length, snoutlength, and distance from the midline at least interorbitalwidth to the snout (Table 15). Nearly complete separationis also obtained between E. meadiae and E. stigmaeum fromthe Coosa River along sheared PC-II (Fig. 22A). ShearedPC-III completely separates E. teddyroosevelt from all otherspecies except E. stigmaeum (Fig. 22A). Variables accountingfor the most variance on this axis include three bodydepth measurements, first anal spine length, anal fin baselength, and snout length (Table 15). The plot nearly sep-Figure 22. Plot of morphometric sheared principalcomponent two (PC-II) and sheared PC-III for males ofnine species of Doration. (A) and (B) are the same buthighlight different sets of polygons to facilitate interpretation.Polygons bound all individuals of: 1—Etheostomastigmaeum, Coosa River (Mobile Basin); 2—E. stigmaeum,Pearl River (central Gulf Coast); 3—E. stigmaeum,Clarks River (upper Mississippi Embayment); 4—E. stigmaeum,Ouachita River (lower Mississippi Embayment);5—E. jessiae; 6—E. meadiae; 7—E. akatulo; 8—E. obama;9—E. gore; 10—E. jimmycarter; 11—E. teddyroosevelt;12—E. clinton.

38 Bulletin 30 NOVEMBER 1, 2012E. jimmycarter, E. teddyroosevelt, and E. clinton. Both E.meadiae and E. jimmycarter separate completely or nearlycompletely from E. stigmaeum, E. akatulo, and E. clinton.Unlike the male analysis, females of E. clinton broadlyoverlap those of E. stigmaeum from Ouachita River, E. jimmycarterpartly overlaps E. teddyroosevelt, and E. teddyrooseveltpartly overlaps E. clinton (Fig. 23). Variables loading heavilyon sheared PC-II include first anal spine length, upperjaw length, snout length, and distance from the midlineat least interorbital width to the snout (Table 15). Thoseloading most heavily on sheared PC-III are first anal spinelength, anal fin base length, snout length, and two measuresof body depth.CONTACT ZONESBear Creek system, Alabama.—Howell (1968; 1980a)treated specimens from Little Bear Creek in the BearCreek system of Alabama (Tennessee River drainage) asintergrades or hybrids between E. jessiae and E. stigmaeum.Etheostoma stigmaeum occurs in adjacent Bear Creek,having presumably entered the system via stream capturefrom the upper Tombigbee River system (Wall, 1968;Starnes and Etnier, 1986). Howell based his assessment onthe observation that six of 12 available specimens (50%)possessed a frenum, which he considered the most distinctivecharacter separating the two taxa. However, meristic,breeding color, and allozyme data all indicate that LittleBear Creek contains “pure” E. jessiae and this “apparentzone of intergradation” (Wiley, 1981) does not involvegene flow between E. stigmaeum from Bear Creek and E.jessiae. This conclusion is discussed below.First, it has been shown above with a larger sample size(n = 49) that actually about 69% of specimens from LittleBear Creek possess a frenum. This frequency is comparableto that observed in geographically proximate populationsof E. jessiae from Shoal Creek, Paint Rock River,and Sequatchie River; however, there is no clear patternof clinal variation, as several intervening populations exhibitfrenum frequencies of 100%. Second, in comparingmeristic frequencies between Little Bear Creek specimens,E. jessiae, and 54 specimens of E. stigmaeum fromBear Creek, counts for Little Bear Creek are highly consistentwith those of E. jessiae. This is true for counts of lateralscales, unpored lateral scales, transverse scales, scalesbelow and above the lateral line, caudal peduncle scales,dorsal spines, dorsal soft rays, anal soft rays, pectoral rays,and nape squamation, distributions of which are diagnosticfor the two species (Tables 1 through 5, and Tables 7through 9). No meristic counts examined exhibit intermediacythat would support a hybridization hypothesis.Principal component analysis of 16 meristic variables inE. jessiae shows that Little Bear Creek specimens are typicalfor the species, broadly overlapping all other populationson the plot of PC- I versus PC-II (Fig. 24; Table 16).Figure 23. Plot of morphometric sheared principalcomponent two (PC-II) and sheared PC-III for femalesof nine species of Doration. Polygons bound all individualsof: 1—Etheostoma stigmaeum, Coosa River(Mobile Basin); 2—E. stigmaeum, Pearl River (centralGulf Coast); 3—E. stigmaeum, Clarks River (upperMississippi Embayment); 5—E. jessiae; 6—E. meadiae;7—E. akatulo; 8—E. obama; 9—E. gore; 10—E. jimmycarter;11—E. teddyroosevelt; 12—E. clinton.Third, breeding coloration (Fig. 3) and maximum size(56 mm SL) of Little Bear Creek males are typical of E. jessiae.Breeding coloration of E. stigmaeum males observedfrom Bear Creek is typical for that species (Fig. 3), andmaximum size is only about 42 mm SL. Finally, patterns ofallozyme variation presented in (Layman, 1994) indicatethat E. stigmaeum and E. jessiae from the Bear Creek systemare not hybridizing. At four diagnostic loci between E. stigmaeumfrom Bear Creek and E. jessiae, all Little Bear Creekspecimens (n = 10) possessed only E. jessiae alleles; no individualswere heterozygous for alleles from both species.There is no convincing evidence that E. stigmaeum andE. jessiae have hybridized in the Bear Creek system. Thespecies occur parapatrically in the system, with their distributionsseparated by a gap that Wall (1968) attributesto lack of suitable sand and gravel substrates. This studysuggests that the polymorphic condition of the frenum inpopulations of E. jessiae from Little Bear Creek and otherrivers in the southern bend of the Tennessee River representsresidual geographic variation left from the ancestorof E. jessiae, perhaps maintained by chance or local selection(Wiley, 1981). Furthermore, even if hybridization hadoccurred, it would have little bearing on decisions concerningthe species status of these taxa, since the phylogenypresented below suggests that they are not sister taxa.Clinch River system, Tennessee.—Etheostoma jessiae andE. meadiae occur parapatrically in the Clinch and Powellrivers (Fig. 7). Meristic and breeding color data indicatethat E. jessiae occurs both downstream of Norris Dam and

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 39Figure 24. Plot of meristic principal component one(PC-I) and PC-II for Etheostoma jessiae and E. meadiae.Polygons bound all individuals of: 1—E. jessiae,lower and middle Tennessee River drainage; 2—E. jessiae,Little Bear Creek, Alabama; 3—E. jessiae, upperTennessee River drainage exclusive of Clinch River;4—E. jessiae, Clinch River downstream of Norris Dam;5—E. jessiae, Clinch-Powell river upstream of NorrisDam; 6—E. meadiae, Powell River; 7—E. meadiae, ClinchRiver. Overlap between the two species is indicated byshading.in at least three tributaries of Norris Reservoir. The tributariesinclude Cove Creek downstream of the confluenceof the two rivers, Davis Creek (pond opposite Doak’s Dam)in the lower Powell River, and Big Sycamore Creek in theClinch River upstream of the Powell River confluence.Several diagnostic meristic features show distinct shifts inmodes between these populations of E. jessiae and populationsof E. meadiae in the upper Powell and Clinch rivers.These include counts of lateral scales, unpored lateralscales, caudal peduncle scales, transverse scales, scales belowthe lateral line, anal soft rays, and estimates of cheekand nape squamation (Tables 1 through 7). In addition,clear shifts occur in counts of principal caudal fin raysand the proportions of individuals having a frenum andpalatine teeth as shown by stream system in Table 11 (seealso Fig. 7).Meristic variation between E. jessiae and E. meadiae wasfurther evaluated using principal component analysis.The plot of PC-I versus PC-II for 16 meristic variables (Fig.24) shows nearly complete separation of all populationsof E. jessiae from E. meadiae of the upper Powell River, andprovides major separation between E. jessiae and E. meadiaeof the upper Clinch River. Populations of E. jessiae fromthe Clinch River system both above and below Norris Dambroadly overlap all other populations of E. jessiae and aredistinct from E. meadiae. Separation between E. jessiae andE. meadiae occurs along PC-I, which is most strongly correlatedwith counts of transverse scales, caudal pedunclescales, scales below the lateral line, and scales in lateral series,estimates of nape and cheek squamation, and countsof caudal and anal fin rays (Table 16). In addition, breedingmales observed from Big Sycamore Creek above NorrisDam and Coal Creek below Norris Dam consistentlypossessed bright orange spots on the soft dorsal, caudal,and pectoral fins, and bright orange pigment in the baseof the spinous dorsal fin. These aspects of coloration aretypical of E. jessiae but not E. meadiae.Etnier and Starnes (1994) hypothesized that tributariesof Norris Reservoir and Clinch River below Norris Damcontain apparent intergrades or introgressed hybrids betweenE. jessiae and E. meadiae. The morphological analysisabove strongly supports recognition of these populationsas E. jessiae. Only counts of pectoral fin rays suggestpossible introgression of these populations by E. meadiae.Like E. meadiae, Clinch River populations of E. jessiae modallyhave 14 pectoral rays, compared to 15 in most otherE. jessiae populations. Alternatively, this apparent shift inE. jessiae may represent residual geographic variation thatwas present in the species’ ancestor. A large proportion ofE. jessiae specimens throughout the remainder of the upperTennessee drainage possesses 14 pectoral rays (Table9), and Etnier and Starnes (1994) noted that the modalcount was also 14 in the Little Tennessee River system.Patterns of allozyme variation, presented in Laymanand Mayden (ms), provide stronger evidence for introgressionof E. jessiae populations by E. meadiae in the ClinchRiver system. At the single diagnostic locus between thetaxa, populations identified as E. jessiae from Coal Creek(n= 10), located just downstream of Norris Dam, and BigSycamore Creek (n= 16), tributary to Norris Reservoir,contained individuals homozygous for alleles of E. jessiaeand individuals heterozygous for alleles of both species.The Coal Creek population also contained two individualsthat were homozygous for the allele of E. meadiae.Whether these specimens represent pure E. meadiae versusF2 or backcross classes cannot be determined withoutadditional genetic markers.Although allozyme evidence suggests that introgressivehybridization may have occurred between E. jessiae and E.meadiae, the data are ambiguous. Neither species exhibitsunique derived alleles at this locus, and thus, the presenceof the E. meadiae allele in the two E. jessiae populations couldalso be explained as a retained ancestral polymorphism.Nevertheless, the abutting distributions of these two species(Fig. 7) leave little doubt that they have had ample opportunityto hybridize in the past. Given the hypothesizedphylogeny below in which these two species are not consideredto be sister taxa, these encounters may represent casesof secondary contact. If so, they have little bearing on decisionsregarding the species status of these taxa.

40 Bulletin 30 NOVEMBER 1, 2012PHYLOGENETIC RELATIONSHIPSPhylogenetic hypothesisPhylogenetic analyses using both the unordered andmixed unordered-ordered character sets produced thesame three minimum-length trees with a length of 60steps and a consistency index of 0.683 (CI; Kluge andFarris, 1969) (uninformative characters excluded). Theretention index was 0.776 in the unordered analysis and0.787 in the mixed analysis (RI; Farris, 1989). The leastresolved of the minimum-length trees, which also representsthe strict consensus tree, is shown in Figure 25.All three trees support the monophyly of Doration with E.akatulo sister to a clade containing all other species in thesubgenus. A pattern of asymmetrical branching proceedsup the tree with E. clinton, E. stigmaeum, E. meadiae, andE. jessiae. Etheostoma jessiae is sister to a clade containing E.obama, E gore, E. jimmycarter, and E. teddyroosevelt. The othertwo minimum-length trees differ only in the resolution ofthe trichotomy involving this latter clade. One tree placesE. gore as sister to the E. jimmycarter-E. teddyroosevelt clade,and the other tree unites E. obama and E. gore as sister tothe E. jimmycarter-E. teddyroosevelt clade. As the more conservativeestimate of relationships, the less highly resolvedtree is presented as the preferred phylogenetic hypothesis(Fig. 25).The monophyly of Doration is supported by ten derivedcharacter states (Fig. 25). Breeding males of the ancestorof this clade are hypothesized to have possessed a seriesof 7–11 vertically elongate blue bars, scattered red-orangemarkings on the sides, iridescent blue or blue-green pigmenton the opercles, cheeks, lips, and mid-gular area, aspinous dorsal fin with dusky to black marginal and submedialbands and red-orange medial band, and a ventrally-developedblue-green basicaudal bar. The ancestor alsopossessed a pair of small dark spots on the medial caudalfin base and modally 11–13 dorsal spines.Within Doration, E. akatulo is sister to a clade containingall other species in the subgenus. This latter clade is unitedby three derived features of male breeding coloration:blue or blue-green marginal and submedial bands in thespinous dorsal fin, blue or blue-green pigment in the baseof the soft dorsal fin, and blue or blue-green pigment inthe base of the anal fin. Etheostoma akatulo is characterizedby three homoplasious autapomorphies: a complete lateralline, fully scaled cheeks, and loss of palatine teeth. Thecomplete lateral line is interpreted in this optimizationas a reversal, but an equally parsimonious reconstructionallows for parallel derivation of an incomplete lateral linein E. caeruleum, E. chlorosoma, and the ancestor of the cladesister to E. akatulo. Derivation of fully scaled cheeks in E.akatulo is an instance of parallel evolution of the samestate in the ancestor of the E. obama-E. gore-E. jimmycarter-E. teddyroosevelt clade, and the loss of palatine teeth is aparallel transformation with that in E. chlorosoma.Monophyly of the clade containing E. stigmaeum, E.meadiae, E. jessiae, E. obama, E. gore, E. jimmycarter, and E.Figure 25. Hypothesized phylogeny of species of thesubgenus Doration. This is one of three minimum-lengthtrees and also represents the strict consensus tree.Derived character states supporting each of the letteredbranches are: A) 17(1), 19(1), 22(1), 34(1); B) 16(1), 25(1),26(1), 30(1), 34(2); C) 17(0)*, 23(0)*, 27(1)*; D) 28(0)*,29(0)*; E) 1(1), 2(1), 3(1), 4(1), 5(1), 7(1), 8(1), 12(1), 20(1),32(1); F) 23(0)*, 24(1)*, 28(0)*; G) 7(2), 10(1), 13(1); H)1(2)*, 27(1)*; I) 12(2); J) 28(1)*; K) 5(0)*, 8(2); L) 22(0,1)*,24(0,1)*; M) 9(1), 11(1), 15(1); N) 22(0)*, 28(1)*; O)1(2)*, 4(0)*, 6(1), 14(2), 24(1)*; P) 14(1), 28(1)*; Q) 7(1)*,10(0)*, 12(1)*, 13(0)*; R) 6(0)*, 9(0)*. See Appendix A forcharacter descriptions and Appendix B for data matrix.Homoplasious transformations indicated by asterisks.teddyroosevelt is supported by a single character, a verticalblue or blue-green basicaudal bar in nuptial males that extendsfrom the dorsal to the ventral margin of the caudalfin. Etheostoma clinton, sister to this clade, exhibits quadratelateral blotches in breeding males and 9 preoperculomandibularpores as autapomorphies. Both states representinstances of parallel development, the former withthe ancestor of the E. obama-E. gore-E. jimmycarter-E. teddyrooseveltclade, and the latter with E. nigrum.The clade containing E. meadiae, E. jessiae, E. obama,E. gore, E. jimmycarter, and E. teddyroosevelt is united by theabsence of blue or blue-green pigment on the gular area

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 41of breeding males, a reversal, and development of a trueorange medial band (in life) in the spinous dorsal fin ofbreeding males. This group is sister to E. stigmaeum, whichis characterized by partly scaled cheeks, a case of paralleldevelopment with E. jessiae and E. obama.Etheostoma meadiae is sister to a clade containing E. jessiae,E. obama, E. gore, E. jimmycarter, and E. teddyroosevelt.Monophyly of the latter group is supported by the uniquederivation of three male breeding color features: orangepigment in the base of the spinous dorsal fin, orange spotson the second dorsal and caudal fin rays, and orange spotson the pectoral fin rays.Etheostoma meadiae is distinguished by polymorphic conditionsof the frenum and palatine teeth. Because PAUPdoes not assign polymorphic states to internal nodes,both polymorphic conditions are optimized as havingarisen through anagenesis in E. meadiae rather than in anancestor. In the case of the frenum, its appearance in E.meadiae is interpreted as a reversal. The only other speciesof Doration possessing a frenum is E. jessiae, for which thisstate is also interpreted as a reversal. This optimizationof the frenum on the phylogenetic tree (Fig. 25) requiresthree steps, because PAUP assigns a cost of one step to alltransformations between unordered character states, evenwhen occurring between fixed and polymorphic states.However, if generalized parsimony (Swofford and Olsen,1990) is adopted for this character, in which transformationsbetween fixed states are assigned a cost of two stepsand those between fixed and polymorphic states a cost ofone step, then an equally parsimonious reconstruction ispossible. The polymorphic state could have arisen in theancestor of the E. meadiae-E. jessiae-E. obama-E. gore-E. jimmycarter-E.teddyroosevelt clade, with fixation to presence ofa frenum occurring in E. jessiae and fixation to absenceof a frenum occurring in the ancestor of the E. obama-E.gore-E. jimmycarter-E. teddyroosevelt clade. This alternativereconstruction is consistent with present-day patterns ofpolymorphism observed in E. jessiae, in that the frenumapparently has not reached complete fixation in severalpopulations of the middle Tennessee drainage, includingLittle Bear Creek.The polymorphic condition of palatine teeth in E.meadiae is optimized in the present tree (Fig. 25) as a convergencewith E. obama. However, using a generalized parsimonyapproach as described above, a more parsimoniousreconstruction (one step shorter) allows for developmentof a polymorphic state in the ancestor of E. meadiae, presenceof palatine teeth becoming fixed in E. jessiae, andabsence of teeth becoming fixed in the ancestor of an E.gore-E. jimmycarter-E. teddyroosevelt clade, with E. obama retainingthe pleisiomorphic polymorphic condition. Thisreconstruction supports the minimum-length tree thatresolves E. gore as sister to the E. jimmycarter-E. teddyrooseveltclade.Etheostoma jessiae is sister to a clade containing E. obama,E. gore, E. jimmycarter, and E. teddyroosevelt. In addition topresence of a frenum, E. jessiae is distinguished by partlyscaled cheeks, a parallel transformation with those inE. stigmaeum and E. obama. Monophyly of the E. obama-E. gore-E. jimmycarter-E. teddyroosevelt clade is supportedby five synapomorphies: breeding males with tangerineorange pigment on the face and underside of the head;Figure 26. Consensus trees for minimum-length trees plus all trees one step longer. (A) Strict consensus and (B)majority-rule consensus of 20 trees from unordered analysis. (C) Majority-rule consensus of 11 trees from mixedunordered-ordered analysis. Numbers above branches in (B) and (C) represent percentages of trees supporting agiven clade.

42 Bulletin 30 NOVEMBER 1, 2012breeding males with orange spots on the anal fin; breedingmales with quadrate lateral blotches (parallelismwith E. clinton); breeding males with no blue pigment onthe lips (reversal); and loss of palatine teeth (parallelismwith E. akatulo). Within this clade, E. jimmycarter and E.teddyroosevelt are united by four features of nuptial malecoloration, all reversals, including black marginal andsubmedial bands in the spinous dorsal fin, no blue pigmentin the soft dorsal fin, a ventrally-developed bluegreenbasicaudal bar, and no blue pigment in the analfin. Etheostoma teddyroosevelt exhibits two reversals as autapomorphies,loss of orange pigment on the face and lossof orange pigment in the base of the spinous dorsal fin.Etheostoma obama is characterized by three autapomorphiccharacter states: restriction of orange spots in the anal finof breeding males to the posteromedial portion of the fin,development of partly scaled cheeks, and polymorphismin the occurrence of palatine teeth.The relationships of E. obama, E. gore, and the E. jimmycarter-E.teddyroosevelt clade are unresolved in the minimum-lengthtree presented in Figure 25. In one of theother two minimum-length trees, a sister relationship betweenE. obama and E. gore is supported by the developmentof orange pigment on the face, which is interpretedas a parallelism with the same transformation in E. jimmycarter.However, this character state reconstruction is ambiguous,occurring only under DELTRAN optimization.The third minimum length tree places E. gore as sister tothe E. jimmycarter-E. teddyroosevelt clade. This relationshipis supported by the more extensive development of orangespots in the anal fin of nuptial males; transformation tothis state occurs under both optimization schemes butfrom different ancestral states. An E. gore-E. jimmycarter-E.teddyroosevelt clade is also supported by the derivation ofnaked cheeks, a parallel reversal with that in E. meadiae,but this occurs only under ACCTRAN optimization. UnderDELTRAN optimization the clade is also supportedby the absence of palatine teeth; this transformation shiftsdown the tree to the ancestor of the E. obama-E. gore-E.jimmycarter-E. teddyroosevelt clade under ACCTRAN. Additionalcharacters are clearly needed to more confidentlyresolve the relationships of these species.Evaluation of support.—The degree of support for thehypothesis of phylogenetic relationships (Fig. 25) was furtherevaluated by generating the strict and majority ruleconsensus trees for the minimum-length trees (60 steps)plus all trees that are one step longer. Twenty trees

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 43ing a monophyletic Doration and an unresolved clade containingE. obama, E. gore, E. jimmycarter, and E. teddyroosevelt.The majority-rule consensus tree for the unorderedanalysis (Fig. 26B) indicates that 95% of the trees

44 Bulletin 30 NOVEMBER 1, 2012island along co. rd. Bradley County: CU 69879 (10), ConasaugaR. at TN Hwy 74 and downstream of bridge for 0.4km. Coosa River system, Georgia. Floyd County: UMMZ168823 (6), Little Armuchee Cr. at US Hwy 27, CrystalSprings, 7.2 km NW Armuchee; UMMZ 88269 (1), SpringCr. at Spring Creek, ca. 16 km SE Rome; ANSP 20645 (1;lectotype), Etowah R. near Rome; ANSP 20646 (3; paralectotypes),Etowah R. near Rome. Chattooga County:UGAMNH 1180 (1), Armuchee Cr. at US Hwy 27 nearGore. Dawson County: UGAMNH 2040 (1), Yellow Cr.from Co. Rd. 157 to confluence with Etowah R. BartowCounty: UGAMNH 37 (2), Stamp Cr. above upper bridge;INHS 75090 (1), Stamp Cr. 6.4 km SE White; UAIC10116.07 (12), Stamp Cr. at GA Hwy 20; UGAMNH 16 (7),Allatoona Cr. 3.2 km from mouth; UGAMNH 100 (5), AllatoonaCr. lower section. Paulding County: UF 80126 (7),84743 (5), UAIC 10103.12 (11), Raccoon Cr. at BraswellMountain Road, 6.0 km NE Braswell at powerline. CoosaRiver system, Alabama. Cherokee County: TU 26070 (13),trib. to Chattooga R. ca. 4.8 km ENE Chesterfield. EtowahCounty: UAIC 9821.09 (20), Little Canoe Cr. 5.3 km NNESteele ca. 1.3 km upstream US Hwy 11. Calhoun County:TU 68587 (20), Shoal Cr. 3.7 km E White Plains. ClayCounty: UAIC 8532.13 (5), Hatchet Cr. 11.5 km WNWMillerville at Co. Rd. 7. Coosa County: UAIC 10044.08 (5),Hatchet Cr. at US Hwy 280. Tallapoosa River system, Georgia.Haralson County: UGAMNH 2142 (20), TallapoosaR. 8.3 air km WNW Buchanan. Tallapoosa River system,Alabama. Tallapoosa County: UAIC 9815.03 (4), UAIC10043.07 (16), Hillabee Cr. at AL Hwy 22,10.1 km NE AlexanderCity. Macon County: UAIC 1477.22 (7), OpintoloccoCr. 4.8 km E Tuskegee on Co. Rd. 26; INHS 78764(10), East Opintolocco Cr. 4.8 km SE Tuskegee; AUM 6565(3), Uphapee Cr. 5.6 air km N Tuskegee at 1-85. AlabamaRiver tributaries, Alabama. Dallas County: AUM 7970(16), Big Swamp Cr. 12.5 air km SE Orrville, T14N, R9E,S3; UAIC 2393 (4), Big Swamp Cr. ca. 3.2 km E Tasso,T14N, R9E, S3. Wilcox County: TU 44512 (20), Turkey Cr.1.6 km S Kimbrough at AL Hwy 5. Monroe County: TU44449 (20), Flat Cr. 13.8 km NW Monroeville at AL Hwy41; UAIC 9705.13 (5), Little R. 5.3 km SSE Uriah at ALHwy 21, Escambia County line. Escambia County: TU99951 (11), Little R. at Co. Rd. 1, 12.6 km NW McCullough.Cahaba River system, Alabama. Bibb County: UAIC5604.27 (20), Little Cahaba R. at Bulldog Bend. DallasCounty: UAIC 9610.17 (10), Oakmulgee Cr. ca. 4.8 km WSummerfield, ca. 0.8 km below AL Hwy 219; UAIC 9611.12(10), Oakmulgee Cr. ca. 5.3 km WSW Summerfield, T18N,Rl0E, S29. Black Warrior River system, Alabama. EtowahCounty: UAIC 3307 (12), Little Cove Cr., T11S, R3E, S12.Blount County: UAIC 2512 (8), Little Warrior R. at ALHwy 79, 16 km N Jefferson County line. Winston County:UAIC 4111.07 (9), Sipsey Fork at Low Pressure Bridge, ca.6.4 km E AL Hwy 195, ca. 8.8 km NNE Double Springs;UAIC 4329.16 (11), Sipsey Fork at Sipsey Fork RecreationArea. Tuscaloosa County: UAIC 8332.06 (11), UAIC8334.08 (6), UAIC 9857.09 (3), Mill Cr. at US Hwy 82,Northport. Upper Tombigbee River system, Alabama.Marion County: TU 40510 (17), trib. to Buttahatchie R. atUS Hwy 278,0.6 km W AL Hwy 129 jct.; UAIC 4315.24 (3),Williams Cr. at US Hwy 43, Hamilton, TI0S, RI4W, S35;UAIC 4316.19 (16), UAIC 10371.01 (4), Luxapalila Cr. atUS Hwy 43, Winfield. Sumter County: TU 48924 (20),Noxubee R. at AL Hwy 17, 7.0 km NW Geiger. Lower TombigbeeRiver tributaries, Alabama. Marengo County: TU60933 (20), Beaver Cr. 19.0 km SW Linden at AL Hwy 69.Choctaw County: TU 56991 (10), Souwilpa Cr. 6.2 km SGilbertown at AL Hwy 17; TU 57009 (10), Bogueloosa Cr.1.3 km N Toxey at AL Hwy 17. Washington County: UAIC1088 (16), UAIC 2498 (2), Bilbo Cr. at US Hwy 43, 4.5 kmS McIntosh. Tennessee River drainage. Bear Creek system,Alabama. Franklin County: UAIC 5968.12 (11), UAIC10336.02 (13), Bear Cr. at Co. Rd. 93, 3.6 km NE PoseyMill, T8S, Rl0W, S23/24; UAIC 2323 (3), Little Bear Cr. ca.4.8 km SE Phil Campbell at Burdeshaw Bridge, T8S, R11W,S34; UAlC 1777 (3), Little Bear Cr. below BatestownBridge, T8S, R11W, S26; UAIC 1797 (1), Bear Cr. betweenPrice’s Bridge and mouth of Whitehead Spring Branch,T8S, R12W, S32; UAIC 2194 (1), Bear Cr. ca. 4.0 km NEHodges, ca. 61 m downstream of Scott Bridge, T8S, R13W,S15. Winston County: UAIC 2322 (10), Bear Cr. ca. 6.4 kmN Haleyville, T9S, R11 W, S7. Marion County: UAIC 1886(9), Bear Cr. 1.6 km W Bear Creek community on AL Hwy172, T9S, R11W, S17; UAIC 1795 (3), Bear Cr. ca. 4.8 km EHackleburg at AL Hwy 172, T9S, R12W, S11. Eastern GulfCoast systems. Pensacola Bay system, Alabama. CovingtonCounty: UAIC 4506.17 (3), trib. to Conecuh R. 2.4 kmfrom Gantt. Escambia County: UAIC 10374.01 (7), Big EscambiaCr. at first bridge N I–65, 3.8 km NW BarnettCrossroads; UAIC 10283.06 (15), Big Escambia Cr. 9.6 kmNW Flomaton just downstream of powerline and pipelinecrossing, T1N, R7E, S11. Pensacola Bay system, Florida.Santa Rosa County: TU 56722 (20), Pond Cr. 3.2 km SWMilton at US Hwy 90; UF 75442 (3), Escambia R. ca. 1.6km upstream FL Hwy 4, 2.7 km E Century, EscambiaCounty line; UF 75376 (5), Escambia R. at FL Hwy 4, 2.7km E Century, Escambia County line; UF 75365 (1), EscambiaR. below FL Hwy 4, above Look and Tremble Oxbow.Perdido River system, Alabama. Baldwin County: TU56700 (1), trib. to Styx R. 16 km W Florida state line at I-10; TU 87833 (16), Blackwater R. 1.8 km S US Hwy 90, 5.8km SW Seminole. Perdido River system, Florida. EscambiaCounty: TU 92066 (8), TU 98521 (1), TU 115961 (2),Perdido R. at FL Hwy 184, Muscogee. Central Gulf Coastsystems. Pascagoula River system, Mississippi. ClarkeCounty: UAIC 1583 (11), UAIC 10050.17 (7), Shubuta Cr.at US Hwy 11, 9.6 km S Pachuta. Jones County: TU 56151(20), Leaf R. just below I-59,3.2 km W Moselle. GeorgeCounty: UAIC 4058 (6), Escatawpa R., T3S, R5W, S2/3.Pascagoula River system, Alabama. Mobile County: TU100953 (11), Franklin Cr. at US Hwy 90, 0.8 km E Mississippistate line. Pearl River system, Mississippi. Leake

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 45County: TU 128164 (8), TU 128234 (2), Pearl R. at Edinburg,MS Hwy 16. Copiah County: TU 56819 (10), trib. toPearl R. at MS Hwy 27, 8.8 km NW Georgetown. SimpsonCounty: TU 56071 (20), Strong R. at MS Hwy 28,3.2 km WPinola. Pike County: TU 43129 (20), Bogue Chitto at USHwy 98,15.2 km E McComb. Pearl River system, Louisiana.Washington Parish: TU 17464 (7), UAIC 10066.12(13), Pushepatapa Cr. at LA Hwy 21,0.3 km S Varnado.Lake Ponchartrain system, Louisiana. Tangipahoa Parish:TU 116611 (20), East Fork Big Cr. at LA Hwy 1054, 7.8 kmENE Arcola. Lake Ponchartrain system, Mississippi. AmiteCounty: UAIC 10373.01 (4), trib. of West Fork Amite Riverat MS Hwy 567, 7.2 air km N Liberty; TU 76067 (16), WestFork Amite R. at MS Hwy 48,5.1 km SW Liberty. LowerMississippi Embayment, eastern tributaries. HomochittoRiver system, Mississippi. Lincoln County: TU 66522 (20),Homochitto R. at MS Hwy 550, 8.0 km E Union Church.Bayou Pierre system, Mississippi. Copiah County: UAIC10320.07 (3), trib. to Bayou Pierre 1.6 km S Glancy, 12.0air km SW Hazelhurst; TU 93642 (8), Bayou Pierre at secondbend below MS Hwy 18, 1.3 km below bridge. ClaiborneCounty: TU 91167 (9), Bayou Pierre 0.6 km N CarlislePost Office. Big Black River system, Mississippi. AttalaCounty: TU 138972 (16), Long Cr. 1.1 km S McAdams.Warren County: UAIC 246 (13), Clear Cr. 16 km E Vicksburg,0.8 km N US Hwy 80. Yazoo River system, Mississippi.Calhoun County: TU 156228 (2), Lucknuck Cr. at MSHwy 9; TU 156253 (8), McGill Cr. at MS Hwy 9, 5.9 km SWPontotoc County line. Holmes County: TU 146272 (20),Williams Cr., T14N, R2E, S7. Upper Mississippi Embayment,eastern tributaries. Hatchie River system, Tennessee.McNairy County: UT 91.530, part 1 of 2 (1), Boles Cr.at co. rd., 4.8 km S TN Hwy 57; UT 91.530, part 2 of 2 (1),Mosses Cr. on Rose Creek Rd., 2.1 km N TN Hwy 57,4.0km ENE Pocahontas. Hardeman County: CU 65770 (6),Spring Cr. above and below second bridge on TN Hwy 125S Bolivar; UT 91.529 (16), Spring Cr. at Co. Rd. 8158;UAIC 10375.01 (1), Spring Cr. at co. rd., 7.0 air km E Saulsbury.Haywood County: UAIC 10298.27 (3), Hatchie R. atBig Eddy, 17.4 air km SE Brownsville at E end Hatchie NationalWildlife Refuge. Clarks River system, Kentucky.Calloway County: UAIC 9866.18 (20), Middle Fork ClarksR. at Martin Chapel Rd., ca. 2.4 km upstream KY Hwy 121near Murray; TU 53465 (6), East Fork Clarks R. 2.7 km SMurray; SIUC 326 (1), SIUC 12540 (1), West Fork ClarksR. 1.6 km E Coldwater at KY Hwy 121. Graves County:SIUC 11797 (1), old channel West Forks Clark R., 3.2 kmW Clear Springs. Upper Mississippi Embayment, westerntributaries. Castor River system, Missouri. Bollinger County:KU 9279 (9), Castor R. 9.6 km W Zalma, T29N,. R8E,S28; UAIC 10342.15 (6), Castor R. at Co. Rd. 736 (GipseyBridge). St. Francis River system, Missouri. Madison County:TU 74549 (14), St. Francis R. at Co. Rd. C bridge, 13.3km NW jct. US Hwy 67 and Co. Rd. N, T31N, R5E, S11.Wayne County: SIUC 12537 (4), St. Francis R. 4.8 km NWSilva. Iron County: Big Creek 2.4 km SE Annapolis. ScottCounty: KU 9232 (1), Ditch 1, 3.2 km E Vanduser, T27N,R13E, S16; INHS 81727 (1), canal 1.6 km W Grant City.Stoddard County: INHS 81737 (1), Ditch No.1, Badersville.New Madrid County: INHS 81165 (1), Jones DitchCanal 7.2 km E Gideon; INHS 81136 (3), Main Ditch DistrictNo.1, 4.0 km E Morehouse; INHS 76782 (4), drainageditch 1.6 km E Malden; KU 9422 (1), ditch 4.0 km W Matthews,T24N, R13E, S3. Pemiscot County: KU 9592 (1),Main Ditch 4.8 km W Hayti, TI9N, RI2E, S31. Black Riversystem, Missouri. Butler County: UMMZ 139675 (3), CaneCr. at confluence with Ten Mile Cr., 8.0 km W PoplarBluff. Black River system, Arkansas. Clay County: TU66007 (10), Current R. 7.2 km NW Success along AR Hwy211. Randolph County: TU 59738 (7), Current R. 0.5 kmNW Biggers; TU 54622 (3), Current R. at US Hwy 67, 12.8km ENE Pocahontas; NLU 41964 (3), Spring R. at Ravendenaccess near Ravenden; NLU 44321 (4), Spring R. atisland near gravel pit, T18N, R1W, S31. Sharp County:UAIC 10376.01 (5), Spring R. at Hardy, Hardy Beach Access,just off of AR Hwy 342; NLU 42089 (8), Spring R. atWilliford access; UAIC 10378.01 (10), Strawberry R. at USHwy 167, 3.2 km N Evening Shade. White River system,Arkansas. Marion County: TU 79889 (1), Buffalo R. 0.8km upstream Bush Cr., T17N, R15W, Sl1; NLU 24915 (1),Buffalo R., state park, T17N, RI5W, S34. Baxter County:TU 51237 (1), Buffalo R., second riffle above mouth. IndependenceCounty: NLU 14111 (2), NLU 16215 (2), WhiteR. ca. 24 km NW Batesville, ca. 0.4 km S Lock and DamNo.3; NLU 14193 (1), White R. at Lock and Dam No.3;NLU 29224 (2), NLU 49076 (11), White R. below Lock andDam No.1, Batesville. Little Red River system, Arkansas.Van Buren County: KU 9840 (1), Choctaw Cr., T10N,R14W, S15, and Dry Fk., T10N, R14W, S10; INHS 81088(1), South Fork Little Red R. 14.4 km SW Clinton; NLU48727 (1), Archey Cr. 1.6 km N Clinton, T11N, R14W, S10.Stone County: NLU 28937 (1), Middle Fork Little Red R.ca. 7.2 km NW Arlberg, T31N, R13W, S23. Van BurenCounty: UT 91.987, Middle Fork Little Red R. at AR Hwy9 above Shirley. Cleburne County: NLU 52715 (5), Big Cr.1.6 km down co. rd. off AR Hwy 110, 3.2 km N Pangburn.Lower Mississippi Embayment, western tributaries. OuachitaRiver system, Arkansas. Hot Spring County: NLU67242 (5), Ouachita R. below Remmel Dam, above andbelow Stone Quarry Cr., 8.0 km W I-30, T3S, R17W, S29.Clark County: UT 91.3360 (1), Caddo R. at I-30; NLU58836 (4), UAIC 10379.01 (2), Caddo R. at US Hwy 67, 5.9km N Arkadelphia; NLU 58816 (20), Caddo R. at mouth(Ouachita R.). Ouachita County: UAIC 10062.20 (2), TU45702 (6), Tulip Cr. at AR Hwy 7,24.0 km N Camden. PikeCounty: NLU 52546 (7), Antoine R. 1.6 km S Antoine,T8S, R23W, S25. Nevada County: NLU 47956 (3), BrushyCr. 1.6 km N Prescott on US Hwy 67, left on paved rd. 12.8km, T9S, R22W, S28. Garland County: NLU 48699 (1),South Fork Saline R. at co. rd. 0.4 km E off AR Hwy 5, 4.8km E Fountain Lake. Saline County: NLU 15388 (11),South Fork Saline R. 4.8 km W Benton, T1S, R15W, S28;

46 Bulletin 30 NOVEMBER 1, 2012NLU 38590 (8), South Fork Saline R. at co. rd. 1.6 km ENance. Grant County: NLU 16928 (1), Saline R. at ARHwy 229,4.8 km N Poyen; NLU 15186 (4), Saline R. at USHwy 270,8.0 km W Prattsville; NLU 16946 (5), Saline R. atJenkins Ferry State Park, AR Hwy 46, 6.4 km E Leola.Bradley County: NLU 54006 (20), Saline R., “Moors Mill,”9.6 km S Johnsville, T16S, R9W, S26. Ouachita River system,Louisiana. Morehouse Parish: NLU 59633 (5), BayouBartholomew 6.4 km below LA Hwy 139 bridge; NLU63652 (15), Bayou Bartholomew 0.4 km W LA Hwy 2bridge, 0.2 km W US Hwy 165 jct., T21N, R5E, S28. RedRiver-Atchafalaya River system, Louisiana. LaSalle Parish:NLU 58504 (10), Castor Cr. 30 m S LA Hwy 124 spillway,3.2 km W Olla. Grant Parish: UAIC 10310.11 (13), Big Cr.at old blockaded bridge, 5.3 air km NW Pollock (accessfrom S side); TU 37282 (7), Big Cr. at LA Hwy 8, Fishville.Rapides Parish: NLU 49173 (10), Spring Cr. 2.4 km NGlenmora.Etheostoma jessiae.—Lower and middle Tennessee Riverdrainage, Tennessee. Houston County: TU 14670 (2),Whiteoak Cr. at Hwy 13,13.6 km S jct. Hwys. 13 and 49.Humphreys County: UT 91.1040 (1), Whiteoak Cr. at continuationof Houston Co. Rd. 6365; UAIC 9864.16 (5),Whiteoak Cr. at Pennywinkle Rd., 0.8 km W TennesseeRidge Rd. near Houston County line. Hardin County:UT 91.2789 (8), UAIC 10335.03 (10), Alexander Br. atbridge between TN Hwy 128 and Co. Rd. 8230, trib. toIndian Cr.; UT 91.685 (3), Indian Cr. at US Hwy 64; UT91.2566 (6), Indian Cr. river km 6.4–16.0 and 17.6–32.0.Wayne County: CU 64701 (15), Butler Cr.3.6 km E PleasantSprings Church. Lawrence County: UAIC 4773.18 (5),Factory Cr. at bridge 1.9 km SW Westpoint; UAIC 4775.11(4), Factory Cr. at ford 2.9 km W Westpoint; UAIC 2505(10), Bluewater Cr. ca. 2.4 km SE Loretto. Lincoln County:CU 46607 (4), Elk R. ca. 19.2 air km WSW Winchester onsecondary rd; UT 91.1587 (1), Elk R. at US Hwy 231/USHwy 431, Fayetteville. Franklin County: UAIC 4003 (4),Elk R. at Garner Ford, 3.0 km S Beech Hill, 1.6 km WTims Ford Dam. Grundy County: UAIC 2835 (3), Elk R.at Elk Head. Giles County: UAIC 2704.10 (1), RichlandCr. ca. 5.0 km W Elkton; UT 91.3697 (3), Richland Cr.km 51.2,7.2 air km NW Pulaski. Lower and middle TennesseeRiver drainage, Alabama. Lauderdale County:UAIC 4817.19 (10), Little Cypress Cr., T1S, R11W, S33; CU54801 (2), Little Butler Cr. at Pruitton. Jackson County:CU 64702 (17), Hurricane Cr. 5.3 km NE Freedom BaptistChurch, Estill Fork; UAIC 7123.20 (7), Estill Fk. near BostickHill Church at dirt rd. ford, T2S, R4E, S2. Little BearCreek system, Alabama. Franklin County: TU 40546 (16),Little Bear Cr. 0.4 km above AL Hwy 187; UAIC 10372.01(2), UAIC 10461.09 (4), AUM 8779 (1), Little Bear Cr. atAL Hwy 187, 8.0 km S Belgreen; UAIC 1884 (1), UAIC1918 (3), UAIC 10309.09 (7), Little Bear Cr. at AL Hwy24, just below Jordan’s Mill; AUM 9724 (2), Little Bear Cr.,river km 4.8 and 24.0; AUM 4126 (3), Cedar Cr. at km33.6, 1.3 air km N White Oak, T6S, R14W, S10; UAIC 2420(3), Spring Cliff Br., trib. to Cedar Cr., 3.2 km N SprucePine, T7S, R11 W, S30; UAIC 1782 (1), Little Bear Cr., T7S,R12W, S29; UAIC 1780 (1), Little Bear Cr. at rd. crossing,T8S, RI2W, S7; AUM 9440 (3), Little Bear Cr. 8.5 air kmSW Belgreen, Co. Rd. 27; AUM 9464 (2), Little Bear Cr. atCarpenter’s Bridge, 5.8 air km SSW Guinn Cross Roads.Upper Tennessee River drainage, Tennessee (excludingClinch River). Sequatchie County: UAIC 10110.15 (6), SequatchieR. at Old State Hwy 28, just E Dunlap; TU 33462(2), Sequatchie R. 5.4 km S Dunlap, US Hwy 127; UAIC2779 (2), Sequatchie R. ca. 2.4 km ESE Dunlap. MarionCounty: UT 91.2133 (1), Sequatchie R. ca. 0.4 km abovemouth of Little Sequatchie R.; UAIC 8995.01 (2), SequatchieR. ca. 0.8 km above mouth of Little SequatchieR.; UT 91.2326 (2), Sequatchie R. river km 14.9–16.2; UT91.3662 (5), Sequatchie R. river km 11.4. Bledsoe County:UAIC 2775 (4), INHS 77089 (2), Sequatchie R. ca. 1.6km SE Ninemile. Polk County: UAIC 9819.15 (7), UAIC10040.13 (5), UAIC 10071.17 (3), UT 91.2432 (1), SpringCr. at bridge on unnamed gravel rd., 5.3 air km N Reliance,access from TN Hwy 315; UT 91.4095 (2), SpringCr. at “Watercamp,” near mouth. McMinn County: USNM70646 (2), Arnwine Cr., Athens. Monroe County: UAIC4148.18 (2), Little Tennessee R. at US Hwy 411, SW Knoxville;UAIC 8971.02 (2), Citico Cr. at Citico bridge; UT91.308 (6), Citico Cr. 1.6 km above mouth; UT 91.2321 (3),Citico Cr. 7.2 air km SSE Citico Beach; UT 91.2835 (1),Citico Cr. 5.8 km above last iron bridge on Mountain SettlementRd. Blount County: CU 67581 (7), UAIC 8591.12(10), UAIC 9845.01 (3), Little R. at US Hwy 411; CU 65046(2), Little R. below Melrose Mill Dam; CU 65171 (2), LittleR. ca. 0.5 km above Wildwood Bridge on Wildwood Rd.;CU 65203 (1), Little R. at river km 34.4; CU 65213 (1),Little R. at river km 33.6. Sevier County: CU 52716 (1),East Fork Little Pigeon R. at US Hwy 411, 8.0 km E Sevierville;CU 55070 (1), West Prong Little Pigeon R. at bridgejust E US Hwy 441,0.3 km S US Hwy 441 bridge at PigeonForge; CU 38184 (7), Little Pigeon R. at roadside park 3.2km E Sevierville on US Hwy 411; CU 41884 (18), WaldenCr., trib. to West Prong Little Pigeon R., 5.0 air km SWPine Grove; CU 46624 (3), Cove Cr., trib. to Walden Cr.,18.6 km NE Townsend; CU 72044 (2), Walden Cr. GreeneCounty: UT 91.770 (15), Nolichucky R. below first bridgeabove Meadows Cr., and Meadow Cr. 0.4 km upstream ofmouth; UT 91.4190 (8), Lick Cr. at TN Hwy 348, 8.3 airkm E of its mouth. Hamblen County: UT 91.1209 (5), BentCr., trib. to Nolichucky R. Knox County: UT 91.3984 (20),Flat Cr. below confluence with Little Flat Cr. on IdumeaRd. Hawkins County: UT 91.3713 (8), Beech Cr. alongTunnel Hill Rd., 0.8 rd. km N Webster Valley Rd. SullivanCounty: UT 91.4169 (2), Horse Cr. at TN Hwy 92,7.7air km SSW Kingsport. Upper Tennessee River drainage,Georgia. Dade County: UT 91.2114 (6), Lookout Cr., 8.0stream km S Trenton, below old mill dam; UGAMNH1488 (6), Lookout Cr., 1.6 km S Trenton; UGAMNH 1487(1), Lookout Cr. 4.8 km S Trenton; UAIC 1690 (1), Look-

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 47out Cr. near Trenton, below old mill dam. Walker County:TU 34964 (19), West Chickamauga Cr. at GA Hwy 143,2.1km W jct. GA Hwy 341; UAIC 1821 (3), West ChickamaugaCr. at GA Hwy 143 S Pond Springs. Catoosa County: UT91.2726 (8), South Chickamauga Cr. ca. 1.6 km SE Ringgoldat L&N RR bridge along GA Hwy 3. Upper TennesseeRiver drainage, Virginia. Scott County: UMMZ 130756(1), Cove Cr., near Bristol, along US Hwy 58, ca. 3.2 roadkm below point where road leaves Cove Cr.; CU 68S00 (2),Possum Cr. at Hwy 713 bridge S Cleveland School, 4.8 airkm SW Gate City. Clinch River system downstream of NorrisDam, Tennessee. Morgan County: UT 91.110 (12), EmoryR. at Oakdale; INHS 83043 (4), Bitter Cr. 4.8 km ESEOakdale. Anderson County: CU 52088 (1), trib to BrushyCr. near Spessard Mills, 5.8 km W Clinton; CU 52943 (1),Brushy Fk., trib, to Poplar Cr., 8.0 km WSW Clinton, 3.8km NNW Elza Gate, Oak Ridge; CU 44028 (2), Poplar Cr.5.1 km NE Oliver Springs; CU 19151 (11), Poplar Cr. nearOliver Springs, Brown Cr., Brushy Fk.; UT 91.89 (3), inpart, Brushy Fk.; TU 29093 (20), Hinds Cr. ca. 9.6 km SWNorris; UT 91.1280 (4), Buffalo Cr., trib. to Clinch R. atI-75; UMMZ 103676 (4), Clinch R. below bridge; UMMZ159179 (2), Clinch R. 1.6 km below Norris Dam; UT 91.89(4), in part, Coal Cr.; UAIC 10380.01 (18), Coal Cr. alongTN Hwy 116, 2.1 air km SW Lake City; UT 91.2251 (6),Coal Cr. below Briceville; UT 91.2252 (12), Coal Cr. at riverkm 7.7, above Lake City. Knox County: UT 91.692 (13), UT91.1264 (3), Beaver Cr. at US Hwy 441, Halls Crossroads.Union County: UT 91.3368 (9), Bull Run Cr. at first bridgeabove TN Hwy 33. Clinch River system upstream of NorrisDam, Tennessee. Campbell County: UMMZ 113588 (19),Cove Cr. below Caryville Dam; UMMZ 104177 (1), CoveCr.; UMMZ 103591 (10), Pond opposite Doak’s Dam. ClaiborneCounty: USNM 70644 (2), USNM 70645 (9), BallCr., Tazewell. UMMZ 96903 (1), Little Sycamore Cr., notfar from Tazewell; UAIC 9838.11 (19), Big Sycamore Cr. atBuck Lick Rd., ca. 0.4 km SE TN Hwy 33, ca. 4.8 km upstreamNorris Reservoir; CU 49709 (5), Big Sycamore Cr.5.9 km E jct. of TN Hwy 33 and US Hwy 25 at HowardtonChurch.Etheostoma meadiae.—Powell River system, Tennessee.Claiborne County: UMMZ 158372 (10), CU 48256 (6),UT 91.4183 (1), Powell R. 4.8 km SE Harrogate at USHwy 25E; UF 17188 (1), Powell R. at US Hwy 25E andfour other sites to 3.7 km above bridge, 4.8 km S Harrogate;UT 91.55 (3), Russell Cr. at Powell R.; UF 9853 (3),Powell R. 15.2 km NE Tazewell, near Hoop; UT 91.2247(5), UCT 91.2249 (5), UT 91.2250 (2), UAIC 9840.23 (7),Powell R. at Buchanan Ford, access via Yeary Rd. from TNHwy 345 at Hopewell Church; USNM 048903 (1; holotype),USNM 125623 (1; paratype), Indian Cr., CumberlandGap, Tennessee, Powell River system, Virginia. LeeCounty: UT 91.2253 (3), Powell R. at Fletcher Ford, riverkm 187.7; UAIC 7951.19 (1), Powell R. at VA Hwy 70,1.6km S Bowling; UMMZ 103443 (2), Powell R. at mouth ofStation Cr. on US Hwy 58 E Jonesville. Clinch River system,Tennessee. Hancock County: UT 91.3948 (6), ClinchR. km 275.7, Swan Island, 8.8 air km SW Sneedville; UT91.1266 (6), UT 91.2129 (20), UAIC 8880.04 (8), Clinch R.at Frost Ford, ca. 6.4 km above Sneedville; UAIC 9855.14(6), UAIC 10705.01 (6), Panther Cr. at Jimmie Brooks Rd.,just off of TN Hwy 33, 5.6 km E Sneedville; UAIC 10706.01(7), Blackwater Cr. at TN Hwy 70, 0.5 km S Virginia line;CU 46433 (1), CU 46434 (11), Clinch R. 0.5 km downstreamTN Hwy 70, Kyles Ford; UAIC 8986.09 (9), ClinchR. at Kyles Ford, ca. 16 km ENE Sneedville. Clinch Riversystem, Virginia. Lee County: CU 52785 (1), BlackwaterCr. at VA Hwy 70 and Co. Rd. 600 bridge, ca. 12.8 air kmN Kyles Ford. Scott County: UAIC 7942.20 (2), Clinch R.0.4 km downstream of US Hwy 23/58; CU 62669 (3), CU62919 (1), CU 63130 (2), CU 63502 (3), CU 64129 (2), CU52159 (1), Copper Cr. 0.4 km above mouth on Co. Rd. 627,2.0 air km S Clinchport; CU 52134 (2), Copper Cr. frommouth to 0.6 km upstream, off Co. Rd. 627, 2.0 air km SClinchport; UT 91.1926 (2), Copper Cr. at VA Hwy 71, ca.2.4 air km SSW Nickelsville; CU 62845 (2), Copper Cr. 4.0km above mouth on Co. Rd. 627, 5.4 air km ESE Clinchport;TU 71971 (15), Copper Cr. 2.4 km NE Speers Ferry;TU 69268 (4), Copper Cr. at ford, 4.0 km NE Speers Ferry;CU 63476 (1), Clinch R. above Hwy 35/Hwy 58 bridge,1.6 km S Clinchport; UAIC 10484.19 (1), Clinch R. 0.4 kmdownstream US Hwy 23/58; UT 91.2255 (2), Clinch R. justabove Hwy 71 at Fort Blackmore, river km 364.0. TazewellCounty: VPI 2690 (14), Indian Cr. at jct. Hwy 627 and Hwy630.Etheostoma obama.—Duck River system, Tennessee.Humphreys County: UT 91.832 (3), Duck R. at mouth ofHurricane Cr.: UT 91.856 (3), Buffalo R. (the “whirl”) 2.1river km above mouth, at access to private home, gravelshoals and island. Hickman County: UT 91.1292 (2), LickCr. ca. 1.6 km NW Primm Springs at low water bridge.Maury County: NLU 50667 (2), Duck R. 0.5 air km E I-65 at mouth of Dewberry Br. Marshall County: UAIC6395.11 (23), Duck R. ca. 3.2 km upstream US Hwy 31A.Bedford County: UT 91.747 (1), Sinking Cr. N TN Hwy 64;UAIC 10039.21 (4), Flat Cr. at TN Hwy 64, ca. 1.9 km SWShelbyville; UAIC 2534 (5), Flat Cr. 0.3 km S Shelbyvilleon US Hwy 231; UT 91.1599 (16), Duck R. at end of unnumberedco. rd., 9.1 air km NW Shelbyville, 3.2 air kmNW Elbethel; UAIC 9862.15 (8), Duck R. below dam at TNHwy 64/US Hwy 231 in Shelbyville; UT 91.739 (9), GarrisonCr. Perry County: NLU 56575 (1), Cane Cr. at TN Hwy50, ca. 4.8 km E Beardstown. Wayne County: NLU 28797(10), Buffalo R. at TN Hwy 13, ca. 2.4 km SE Flatwoods.Lewis County: UT 91.3973 (1), Buffalo R. 3.5 air km belowTN Hwy 13; UAIC 10462.15 (14), Buffalo R. at mouth ofGrinders Cr. and TN Hwy 99.Etheostoma gore.—Cumberland River drainage, Tennessee.Robertson County: CU 22168 (2), Sulphur Cr. at TNHwy 76, 8.0 km S Adams; UT 91.4066 (1), Sulphur Fork Cr.0.8 km above TN Hwy 76. Montgomery County: CU 23301(1), Yellow Cr. at old bridge, 0.8 km off of TN Hwy 13, S

48 Bulletin 30 NOVEMBER 1, 2012Clarksville and 16.8 km N jct. TN Hwy 13 and TN Hwy 48.Rutherford County: UT 91.2237 (1), Middle Fork StonesR. at “County Farm Ford,” ca. 0.8 km SSE Murfreesboro;CU 42008 (2), CU 51527 (1), INHS 84130 (1), TU 19506(1), UAIC 9865.19 (13), UT 91.708 (2), East Fork StonesR. at US Hwy 231, just below dam at Walterhill, 10.4 kmN Murfreesboro; NLU 15742 (3), East Fork Stones R. 3.2km S Lacassa. Jackson County: UT 91.276 (2), Roaring R.just off TN Hwy 135, at old bridge 4.8 km SE Gainesboro;KU 11539 (12), Roaring R. at TN Hwy 135, 23.7 km NNWCookeville. Overton County: KU 11514 (4), West ForkObey R. 8.0 km E Alpine; AUM 11022 (1), West Fork ObeyR. at TN Hwy 85, 2.1 km SSE Allred; INHS 58232 (3),Cowan Br. at TN Hwy 52, 4.8 km E Alpine. Pickett County:UT 91.196 (5), Wolf R. bridge N Byrdstown-Forbus Rd.,2.4 km E Byrdstown. Scott County: UT 91.426 (10), StationCamp Cr. at mouth, and 0.8 km above mouth. CumberlandRiver drainage, Kentucky. Cumberland County:INHS 63055 (1), Marrowbone Cr. 3.2 km W Burkesville,Hwy 691; UMMZ 154639 (3), Marrowbone Cr. at mouth ofFarris Fork, 1.2 km E Marrowbone; SIUC 4016 (3), CrocusCr. ca. 3.2 km NNW Bakerton. Pulaski County: UF 15405(8), Fishing Cr. at KY Hwy 635, 9.6 km WNW Science Hill;INHS 78871 (4), Buck Cr. 6.4 km W Bandy; SIUC 7584(6), Buck Cr. at KY Hwy 1677, 3.3 km N US Hwy 80; SIUC13245 (4), Buck Cr. ca. 4.8 stream km downstream KY Hwy1012 bridge. Wayne County: TU 74586 (20), Little SouthFork Cumberland River at Parmleysville, 19.2 air km SEMonticello. Rockcastle County: UAIC 9851.08 (23), RockcastleR. along KY Hwy 89, ca. 2.7 km NE KY Hwy490.Etheostoma jimmycarter.—Green River drainage, Kentucky,Monroe County: SIUC 3948 (4), East Fork BarrenR. at KY Hwy 63, 6.4 km NW Tompkinsville. Allen County:SIUC 37 (2), Long Cr. 1.6 km SW Amos; SIUC 78 (2), LongCr. 2.4 km SE Amos. Barren County: UMMZ 165401 (10),Fallen Timber Cr. at KY Hwy 63, 9.6 km SE Glasgow; INHS77938 (10), Skaggs Cr. 3.2 km N Roseville. Casey County:UMMZ 165267 (10), Green R. at Yosemite, KY Hwy 198;UMMZ 169473 (4), branch of Trace Fk. at KY Hwy 910,just S Phil. Taylor County: KU 11626 (4), Big Pitman Cr. atKY Hwy 210, 11.8 km NW Campbellsville. Green County:UMMZ 165302 (8), SIUC 1101 (2), Green R. at Greensburg;UAIC 10521.01 (16), Little Barren R. at KY Hwy 88,16.0 air km W Greensburg. Adair County: UAIC 6495.15(6), trib. to Russell Cr. at KY Hwy 80, ca. 7.0 air km W RussellSprings; UAIC 7157.13 (14), Russell Cr. at KY Hwy 80,Metcalfe County: UAIC 9852.14 (6), Little Barren R. at KYHwy 70, Sulphur Well Larue County: UMMZ 165432 (10),South Fork Nolin R. at Buffalo; INHS 78479 (8), WaltersCr. 6.4 km N Magnolia. Edmonson County: SIUC 18977(1), Green R. at river km 319.0, upstream end of island, ca.14 km ENE Brownsville. Green River drainage, Tennessee.Clay County: UT 91.117 (2), Big Trace Cr. at HermitageSprings; UT 91.393, in part (6), Big Trace Cr. at Tennessee-Kentuckyline; UT 91.393, in part (1), Salt Lick Cr.at Co. Rd. 6138, 1.3 km from Kentucky line; UT 91.1311(1), Salt Lick Cr. at Bethany Rd. Sumner County: UAIC9870.12 (19), UT 91.876, in part (11), West Fork Drakes Cr.at Coker Ford Rd., ca. 1.6 km E Mitchellville; UT 91.876,in part (4), West Fork Drakes Cr. at mouth of Caney Cr.Etheostoma teddyroosevelt.—Arkansas River drainage,Missouri. Lawrence County: KU 6511 (5), Spring R. atMO Hwy 97; KU 10793 (1), Spring R. 9.6 km E La Russel,T28N, R28W, S13. Jasper County: UAIC 10711.30 (1),Spring R. at Co. Rd. D at Quaker Mill Park, just SW Purcell.Arkansas River drainage, Kansas. Cherokee County:KU 11409 (1), KU 18443 (1), KU 16088 (4), Shoal Cr. atKS Hwy 26, Schermerhorn Park, 3.2 km S Galena. ArkansasRiver drainage, Oklahoma. Delaware County: UMMZ103187 (4), Elk R. at Turkey Ford, trib. of Grand R. CherokeeCounty: UMMZ 210571 (3), Fourteen Mile Cr. AdairCounty: KU 2425 (2), Illinois R. at US Hwy 59; UMMZ127177 (4), Barren Fork Cr., trib. of Illinois R., near Proctor.Arkansas River drainage, Arkansas. WashingtonCounty: UT 91.1833 (3), Illinois R. at ford above end ofgravel rd., 10.1 rd. km W US Hwy 71; KU 6333 (2), IllinoisR. at Moffit, 17.6 km SW Fayetteville; UMMZ 170924 (2),Illinois R. at AR Hwy 68,24.0 km W Springdale. JohnsonCounty: UAIC 10305.14 (2), Mulberry R. at AR Hwy 103,0.8 km S AR Hwy 215 jct. Scott County: KU 3546 (1), MillCr., trib. to Fourche La Fave R., at US Hwy 71, 6.4 km SBoles. Yell County: TU 93495 (3), Fourche La Fave R. atbridge 3.2 km S Briggsville. Upper White River drainage,Arkansas. Washington County: UT 91.1576 (6), WestFork White R. at unnumbered rd. crossing off US Hwy 71,ca. 22.4 rd. km S Fayetteville; TU 16568 (1), White R. atAR Hwy 68,12.8 km E Springdale; TU 47038 (1), WhiteR. from AR Hwy 68 to 2.4 km upstream; TU 46961 (2),White R. from bridge W Sulphur City to 2.4 km upstream;TU 46972 (2), White R. from ford N Durham to 2.8 kmupstream; UAIC 10355.11 (3), Richland Cr. 0.4 km downstreamCo. Rd. 79 (first bridge upstream AR Hwy 45); TU50065 (8), Richland Cr. from confluence with Mill Br. to61 m downstream, 12.8 km E Fayetteville; TU 50574 (9),Richland Cr. just below 0.8 km above Hwy 71 bridge, T16N,R28W, S18. Benton County: TU 50588 (1), White R. atEden’s Ford to 0.8 km downstream, TI9N, R29W, S34; TU50419 (3), War Eagle Cr. from bridge to 2.4 km upstream,0.8 km SW War Eagle. Madison County: TU 46992 (2),White R. from bridge at Patrick to 1.6 km downstream;TU 49778 (5), War Eagle Cr. from AR Hwy 68 bridge to2.4 km upstream, NE Huntsville, T17N, R26W, S24; UAIC10325.08 (12), War Eagle Cr. at US Hwy 412 (formerly ARHwy 68); UT 91.1014 (10), King’s R. at AR Hwy 68. CarrollCounty: INHS 86836 (1), Osage Cr. at AR Hwy 68, 1.6 kmNW Osage. Upper White River drainage, Missouri. WebsterCounty: UAIC 10317.12 (1), James R. at MO Hwy KK;KU 10821 (1), James R. 9.6 km S Marshfield, T29N, R18W,S3. Greene County: KU 7885 (1), James R. at Hwy M-125,9.6 km S Strafford.Etheostoma clinton.—Upper Ouachita River system, Arkansas.Polk County: NLU 35232 (1), UT 91.1382 (4), Oua-

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 49chita R. at McGuire Public Access Area, 3.2 km S AR Hwy88 between Ink and Cherry Hill; NLU 58149 (10), NLU59452 (1), Ouachita R. and Mill Cr. just downstream ofCo. Rd. 67, 1.6 km S Cherry Hill. Montgomery County:NLU 34707 (1), NLU 34922 (2), Ouachita R. 1.6 km S ARHwy 88, just E Pine Ridge, T2S, R27W, S9; NLU 34531(1), Ouachita R. at US Hwy 270, Rocky Shoals, T1S, R25W,S31; NLU 34933 (3), Ouachita R. at US Hwy 270, RockyShoals, T1S, R25W, S32; NLU 38580 (7), Ouachita R. atAR Hwy 88, Rocky Shoals; NLU 58422 (2), Ouachita R.at AR Hwy 298, ca. 1.6 km S Sims, T1S, R25W, S20; NLU34918 (4), Ouachita R. at Clifton’s Camp, ca. 2.9 km SAR Hwy 88 and 4.8 km SW Washita, T1S, R24W, S29. UpperCaddo River system, Arkansas. Montgomery County:NLU 15868 (3), NLU 57893 (1), NLU 58925 (7), CaddoR. at jct. AR Hwy 240 and AR Hwy 8, T4S, R24W, S19.Pike County: UT 91.947 (2), Caddo R. at US Hwy 70. ClarkCounty: NLU 18358 (2), NLU 20674 (2), NLU 20706 (1),NLU 28201 (3), NLU 34012 (1), Caddo R. at AR Hwy 182,3.2 km N Amity.

50 Bulletin 30 NOVEMBER 1, 2012TABLE 1. LATERAL SCALE ROW COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Lateral Scale Rows38 39 40 41 42 43 44 45 46 47 48 49 50 51Etheostoma stigmaeumMobile Basin 3 3 9 19 32 32 34 49 57 43 66 47 39Bear Cr. (Tennessee R.) 3 - 3 4 8 14 8 3 6 3 2Eastern Gulf coast systems 2 2 10 14 15 8 16 10 3 2Central Gulf Coast systems 1 1 3 6 10 26 18 29 25 22 15 5 3 1Lower Miss. Embayment, eastern tributaries 2 1 8 11 14 19 16 12 5 9 2Upper Miss. Embayment, eastern tributaries 2 2 3 6 12 4 11 4 10 1Upper Miss. Embayment, western tributaries 2 2 13 14 19 23 18 17 13 7Lower Miss. Embayment, western tributaries 1 5 15 12 23 30 17 30 7 8Etheostoma jessiaeLower and middle Tennessee R. 2 - 3 5 12 20 30 16Little Bear Cr. 1 - 1 7 9 11 7Upper Tennessee R. (excluding Clinch R.) 1 - 3 3 10 13 12 18 30 35Clinch R. downstream of Norris Dam 1 - 7 6 11 15 27 22 16Clinch/Powell R. upstream of Norris Dam 1 2 - - 1 4 3 5 6 11 9Etheostoma meadiaeUpper Powell River 3 3 7 7 9 3 4 5 3 5Upper Clinch River 1 4 7 8 20 21 15 20 8 15 5Etheostoma akatulo 1 1 1 12 25 28 47 33 25 17 8 4 1Etheostoma obama 1 - 4 6 20 21 32 32 28 25 10 5Etheostoma goreMiddle Cumberland R. 2 1 6 11 13 19 17 18 10 5 3 3Lower Cumberland R. 3 4 8 10 17 15 13 8 6 -Etheostoma jimmycarter 1 4 17 18 40 37 28 25 18 16 4 1Etheostoma teddyrooseveltArkansas R. 1 - 2 4 9 13 21 13 17 8 8Upper White R. 1 - 4 9 11 7 16 9 10Etheostoma clinton 2 5 8 14 13 23 14

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 51TABLE 1. (Continued)Lateral Scale Rows52 53 54 55 56 57 58 59 60 n Mean SDEtheostoma stigmaeumMobile Basin 27 13 4 4 4 485 47.5 3.30Bear Cr. (Tennessee R.) 54 44.3 2.36Eastern Gulf coast systems 82 42.5 2.05Central Gulf Coast systems 165 45.0 2.35Lower Miss. Embayment, eastern tributaries 99 44.3 2.24Upper Miss. Embayment, eastern tributaries 1 - 1 57 47.2 2.49Upper Miss. Embayment, western tributaries 2 2 132 47.3 2.31Lower Miss. Embayment, western tributaries 3 151 47.2 2.17Etheostoma jessiaeLower and middle Tennessee R. 15 7 6 3 1 1 121 50.3 2.27Little Bear Cr. 3 8 - 2 49 50.3 2.08Upper Tennessee R. (excluding Clinch R.) 19 27 16 16 10 5 2 - 1 221 51.2 3.11Clinch R. downstream of Norris Dam 14 5 3 1 - 1 129 49.4 2.34Clinch/Powell R. upstream of Norris Dam 5 3 4 1 55 49.6 3.00Etheostoma meadiaeUpper Powell River 49 46.4 2.65Upper Clinch River 4 4 132 47.0 2.67Etheostoma akatulo 203 45.3 2.04Etheostoma obama 1 185 46.7 2.15Etheostoma goreMiddle Cumberland R. - 1 109 45.7 2.41Lower Cumberland R. 2 86 46.6 2.18Etheostoma jimmycarter 209 45.4 2.20Etheostoma teddyrooseveltArkansas R. 5 1 - - 1 103 47.8 2.43Upper White R. 2 69 48.4 2.00Etheostoma clinton 12 7 2 3 - 1 104 49.9 2.31

52 Bulletin 30 NOVEMBER 1, 2012TABLE 2. UNPORED LATERAL SCALE ROW COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Unpored Lateral Scale Rows0 1 2 3 4 5 6 7 8 9 10 11 12 13Etheostoma stigmaeumMobile Basin 1 - 1 3 7 27 55 74 66Bear Cr. (Tennessee R.) 3 5 13 10Eastern Gulf coast systems 2 11 15 18 19 10 -Central Gulf Coast systems 2 1 14 17 33 35 32Lower Miss. Embayment, eastern tributaries 1 - - 1 2 5 3 11 10 15 12 15 11 5Upper Miss. Embayment, eastern tributaries 1 2 8 4 8Upper Miss. Embayment, western tributaries 12 22 18 19Lower Miss. Embayment, western tributaries 1 3 10 14 24 29 22Etheostoma jessiaeLower and middle Tennessee R. 3 4 5 16 7 13 16 17 10 14 5 4 2Little Bear Cr. 1 7 3 5 5 5 4 5 5 4 3 1 - -Upper Tennessee R. (excluding Clinch R.) 5 2 12 9 9 21 22 23 23 22 16 14 16 4Clinch R. downstream of Norris Dam 2 3 3 - 10 7 10 18 21 9 6 5 9 4Clinch/Powell R. upstream of Norris Dam 2 - 2 2 4 3 6 4 6 10 3 5 1 1Etheostoma meadiaeUpper Powell River 10 4 2 8 5 10 6 3 1Upper Clinch River 20 7 8 15 17 19 19 11 6 5 2 2 - 1Etheostoma akatulo 180 15 3 4 - - - - 1Etheostoma obama 3 8 12 13 13 25 19Etheostoma goreMiddle Cumberland R. 1 2 3 - 5 15 17 13 14Lower Cumberland R. 2 2 8 12 16 16 9 9Etheostoma jimmycarter 2 5 13 15 22Etheostoma teddyrooseveltArkansas R. 1 1 2 8 5Upper White R. 3 6Etheostoma clinton 7 4 9

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 53TABLE 2. (Continued)Unpored Lateral Scale Rows14 15 16 17 18 19 20 21 22 23 24 25 26 27Etheostoma stigmaeumMobile Basin 74 53 37 30 15 16 9 6 7 2 - 1 1Bear Cr. (Tennessee R.) 6 5 6 4 1 - - - 1Eastern Gulf coast systems 4 3Central Gulf Coast systems 15 10 3 1 1 - 1Lower Miss. Embayment, eastern tributaries 5 2 - 1Upper Miss. Embayment, eastern tributaries 9 5 10 2 4 3 - 1Upper Miss. Embayment, western tributaries 20 12 8 10 3 5 2 1Lower Miss. Embayment, western tributaries 15 13 6 6 2 3 2 1Etheostoma jessiaeLower and middle Tennessee R. 1 2 1 1Little Bear Cr. - 1Upper Tennessee R. (excluding Clinch R.) 5 2 7 3 4 1 - - - 1Clinch R. downstream of Norris Dam 6 3 3 4 3 1 2Clinch/Powell R. upstream of Norris Dam 1 2 1 1 - - - 1Etheostoma meadiaeUpper Powell RiverUpper Clinch RiverEtheostoma akatuloEtheostoma obama 16 15 24 7 8 7 3 2 5 - - 4 - 1Etheostoma goreMiddle Cumberland R. 7 7 6 5 6 3 2 1 1 - - - - -Lower Cumberland R. 5 4 1 2Etheostoma jimmycarter 33 26 27 24 9 20 5 4 2 1 1Etheostoma teddyrooseveltArkansas R. 10 12 14 19 11 11 5 2 2Upper White R. 11 9 13 5 13 5 2 2Etheostoma clinton 14 13 10 10 7 8 6 6 2 5 1 1 - -

54 Bulletin 30 NOVEMBER 1, 2012TABLE 2. (Continued)Unpored Lateral Scale Rows28 n Mean SDEtheostoma stigmaeumMobile Basin 485 14.0 2.98Bear Cr. (Tennessee R.) 54 13.6 2.34Eastern Gulf coast systems 82 10.3 1.81Central Gulf Coast systems 165 12.0 1.99Lower Miss. Embayment, eastern tributaries 99 9.5 2.88Upper Miss. Embayment, eastern tributaries 57 14.3 2.67Upper Miss. Embayment, western tributaries 132 13.6 2.60Lower Miss. Embayment, western tributaries 151 12.7 2.64Etheostoma jessiaeLower and middle Tennessee R. 121 7.3 3.23Little Bear Cr. 49 5.4 3.31Upper Tennessee R. (excluding Clinch R.) 221 8.2 4.15Clinch R. downstream of Norris Dam 129 8.9 4.39Clinch/Powell R. upstream of Norris Dam 55 8.2 4.13Etheostoma meadiaeUpper Powell River 49 3.4 2.38Upper Clinch River 132 4.3 2.87Etheostoma akatulo 203 0.2 0.77Etheostoma obama 185 14.0 3.92Etheostoma goreMiddle Cumberland R. 1 109 13.0 3.68Lower Cumberland R. 86 10.9 2.35Etheostoma jimmycarter 209 15.2 2.85Etheostoma teddyrooseveltArkansas R. 103 16.1 2.66Upper White R. 69 16.0 3.58Etheostoma clinton 1 104 16.6 3.58

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 55TABLE 3. TRANSVERSE SCALE ROW COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Transverse scale rows9 10 11 12 13 14 15 16 17 18 19 n Mean SDEtheostoma stigmaeumMobile Basin 2 32 104 129 121 78 17 2 485 13.3 1.30Bear Cr. (Tennessee R.) 4 14 20 13 3 54 11.9 1.02Eastern Gulf coast systems 1 20 24 22 14 1 82 11.4 1.11Central Gulf Coast systems 7 17 85 48 7 1 165 12.2 0.87Lower Miss. Embayment, eastern tributaries 4 47 22 17 9 99 12.8 1.07Upper Miss. Embayment, eastern tributaries 6 6 28 13 2 2 57 12.1 1.11Upper Miss. Embayment, western tributaries 2 10 65 31 21 3 132 12.5 0.98Lower Miss. Embayment, western tributaries 5 9 51 57 21 7 1 151 12.7 1.08Etheostoma jessiaeLower and middle Tennessee R. 12 23 54 23 8 1 121 14.0 1.06Little Bear Cr. 10 16 15 8 49 14.4 1.00Upper Tennessee R. (excluding Clinch R.) 1 7 30 66 56 43 13 4 1 221 14.7 1.34Clinch R. downstream of Norris Dam 4 26 44 43 10 2 129 14.3 1.02Clinch/Powell R. upstream of Norris Dam 1 10 18 18 8 55 14.4 1.01Etheostoma meadiaeUpper Powell River 1 12 16 12 8 49 13.3 1.08Upper Clinch River 1 35 40 40 14 2 132 13.3 1.04Etheostoma akatulo 10 29 107 42 15 203 12.1 0.91Etheostoma obama 11 83 62 25 4 185 12.6 0.87Etheostoma goreMiddle Cumberland R. 1 12 51 33 11 1 109 12.4 0.88Lower Cumberland R. 4 38 26 17 1 86 12.7 0.88Etheostoma jimmycarter 9 25 108 50 15 1 1 209 12.2 0.95Etheostoma teddyrooseveltArkansas R. 1 17 49 23 10 2 1 103 13.3 1.02Upper White R. 1 12 26 19 9 2 69 13.4 1.06Etheostoma clinton 3 24 28 29 16 4 104 13.4 1.20

56 Bulletin 30 NOVEMBER 1, 2012TABLE 4. COUNTS OF SCALE ROWS BELOW AND SCALE ROWS ABOVE LATERAL LINE IN SPECIES OF DORATION. Values for holotypes arein boldface.Scale rows below lateral line5 6 7 8 9 10 n Mean SDEtheostoma stigmaeumMobile Basin 16 140 207 101 19 2 485 6.9 0.91Bear Cr. (Tennessee R.) 8 27 17 2 54 6.2 0.75Eastern Gulf coast systems 22 37 21 2 82 6.0 0.79Central Gulf Coast systems 9 98 50 8 165 6.3 0.66Lower Miss. Embayment, eastern tributaries 3 49 28 16 3 99 6.7 0.89Upper Miss. Embayment, eastern tributaries 8 31 14 4 57 6.2 0.79Upper Miss. Embayment, western tributaries 9 66 43 14 132 6.5 0.78Lower Miss. Embayment, western tributaries 7 58 68 16 2 151 6.7 0.78Etheostoma jessiaeLower and middle Tennessee R. 18 66 30 7 121 7.2 0.77Little Bear Cr. 2 19 21 7 49 7.7 0.77Upper Tennessee R. (excluding Clinch R.) 20 85 80 31 5 221 7.6 0.91Clinch R. downstream of Norris Dam 16 50 53 10 129 7.4 0.81Clinch/Powell R. upstream of Norris Dam 6 22 25 2 55 7.4 0.74Etheostoma meadiaeUpper Powell River 1 23 18 7 49 6.6 0.76Upper Clinch River 1 66 47 18 132 6.6 0.73Etheostoma akatulo 15 135 49 4 203 6.2 0.59Etheostoma obama 2 116 59 8 185 6.4 0.59Etheostoma goreMiddle Cumberland R. 2 61 42 4 109 6.4 0.60Lower Cumberland R. 2 44 36 3 1 86 6.5 0.66Etheostoma jimmycarter 34 129 41 5 209 6.1 0.67Etheostoma teddyrooseveltArkansas R. 1 21 52 23 5 1 103 7.1 0.86Upper White R. 1 17 35 12 4 69 7.0 0.85Etheostoma clinton 29 31 38 6 104 7.2 0.92

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 57TABLE 4. (Continued)Scale rows above lateral line3 4 5 6 7 8 n Mean SDEtheostoma stigmaeumMobile Basin 4 150 270 61 485 4.8 0.65Bear Cr. (Tennessee R.) 34 19 1 54 4.4 0.53Eastern Gulf coast systems 10 52 19 1 82 4.1 0.62Central Gulf Coast systems 12 100 52 1 165 4.3 0.59Lower Miss. Embayment, eastern tributaries 1 41 52 5 99 4.6 0.60Upper Miss. Embayment, eastern tributaries 3 46 8 57 4.1 0.43Upper Miss. Embayment, western tributaries 3 85 43 1 132 4.3 0.53Lower Miss. Embayment, western tributaries 7 95 48 1 151 4.3 0.56Etheostoma jessiaeLower and middle Tennessee R. 5 60 55 1 121 5.4 0.59Little Bear Cr. 17 29 3 49 5.7 0.58Upper Tennessee R. (excluding Clinch R.) 2 63 121 34 1 221 5.9 0.69Clinch R. downstream of Norris Dam 1 46 74 8 129 5.7 0.60Clinch/Powell R. upstream of Norris Dam 2 29 21 3 55 5.5 0.66Etheostoma meadiaeUpper Powell River 14 31 4 49 5.8 0.58Upper Clinch River 3 55 65 9 132 5.6 0.65Etheostoma akatulo 2 113 82 6 203 4.5 0.57Etheostoma obama 48 117 20 185 4.8 0.59Etheostoma goreMiddle Cumberland R. 5 60 42 2 109 4.4 0.61Lower Cumberland R. 2 43 40 1 86 4.5 0.57Etheostoma jimmycarter 52 138 19 209 4.8 0.56Etheostoma teddyrooseveltArkansas R. 15 78 10 103 5.0 0.49Upper White R. 8 50 11 69 5.0 0.53Etheostoma clinton 4 54 44 2 104 4.4 0.60

58 Bulletin 30 NOVEMBER 1, 2012TABLE 5. CAUDAL PEDUNCLE SCALE ROW COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Caudal peduncle scale rows12 13 14 15 16 17 18 19 20 21 22 n Mean SDEtheostoma stigmaeumMobile Basin 11 29 77 188 127 45 7 1 485 16.2 1.17Bear Cr. (Tennessee R.) 2 4 12 16 18 1 1 54 14.9 1.20Eastern Gulf coast systems 3 18 28 28 4 1 82 14.2 1.00Central Gulf Coast systems 4 11 79 60 10 1 165 15.4 0.82Lower Miss. Embayment, eastern tributaries 2 24 41 26 6 99 16.1 0.91Upper Miss. Embayment, eastern tributaries 1 16 29 8 3 57 15.9 0.84Upper Miss. Embayment, western tributaries 2 9 36 55 26 3 1 132 15.8 1.00Lower Miss. Embayment, western tributaries 2 17 76 35 18 2 1 151 16.4 0.97Etheostoma jessiaeLower and middle Tennessee R. 2 24 31 53 9 2 121 17.4 1.00Little Bear Cr. 12 16 17 4 49 17.3 0.93Upper Tennessee R. (excluding Clinch R.) 1 3 23 50 68 43 26 5 2 221 18.0 1.36Clinch R. downstream of Norris Dam 1 5 29 40 43 9 1 - 1 129 17.2 1.14Clinch/Powell R. upstream of Norris Dam 2 17 20 11 4 1 55 17.0 1.06Etheostoma meadiaeUpper Powell River 3 30 11 5 49 16.4 0.76Upper Clinch River 1 4 23 86 16 2 132 15.9 0.73Etheostoma akatulo 1 10 31 89 54 16 2 203 15.2 1.02Etheostoma obama 7 79 62 32 3 2 185 16.7 0.92Etheostoma goreMiddle Cumberland R. 1 15 29 56 6 2 109 15.5 0.90Lower Cumberland R. 1 5 18 41 14 7 86 16.0 1.02Etheostoma jimmycarter 6 27 34 134 6 2 209 15.5 0.89Etheostoma teddyrooseveltArkansas R. 1 19 49 24 9 1 103 16.2 0.92Upper White R. 2 14 35 17 1 69 16.0 0.80Etheostoma clinton 11 45 35 10 3 104 16.5 0.91

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 59TABLE 6. PERCENT CHEEK SQUAMATION IN SPECIES OF DORATION. Values for holotypes are in boldface.Percent cheek squamation0 10 20 30 40 50 60 70 80 90 100 n Mean SDEtheostoma stigmaeumMobile Basin 33 54 152 113 47 35 23 21 6 1 485 28.9 18.04Bear Cr. (Tennessee R.) 2 16 18 17 1 54 19.8 9.21Eastern Gulf coast systems 1 6 17 23 14 14 7 82 53.8 14.54Central Gulf Coast systems 1 9 36 73 28 16 2 165 30.5 10.78Lower Miss. Embayment, eastern tributaries 1 5 8 12 28 23 12 10 99 63.0 16.19Upper Miss. Embayment, eastern tributaries 3 12 10 6 7 12 5 2 57 51.9 19.59Upper Miss. Embayment, western tributaries 2 7 18 19 20 39 18 7 2 132 51.5 17.23Lower Miss. Embayment, western tributaries 4 19 25 34 28 21 14 5 1 151 44.1 17.41Etheostoma jessiaeLower and middle Tennessee R. 11 36 45 23 5 - 1 121 18.3 10.62Little Bear Cr. 2 10 28 9 49 19.0 7.43Upper Tennessee R. (excluding Clinch R.) 2 32 105 67 10 4 1 221 23.0 9.01Clinch R. downstream of Norris Dam 10 31 58 19 8 1 2 129 19.6 11.28Clinch/Powell R. upstream of Norris Dam 2 11 24 14 4 55 21.3 9.44Etheostoma meadiaeUpper Powell River 46 1 2 49 1.0 4.21Upper Clinch River 121 10 - - 1 132 1.1 4.33Etheostoma akatulo 2 5 2 6 29 62 97 203 91.0 12.11Etheostoma obama 11 32 82 49 10 - - 1 185 21.1 10.10Etheostoma goreMiddle Cumberland R. 72 28 8 1 109 4.3 6.72Lower Cumberland R. 43 30 12 - 1 86 6.7 8.04Etheostoma jimmycarter 146 44 19 209 3.9 6.50Etheostoma teddyrooseveltArkansas R. 91 10 1 1 103 1.5 4.52Upper White R. 68 1 69 0.1 1.20Etheostoma clinton 50 28 16 7 1 - - - 2 104 9.7 13.97

60 Bulletin 30 NOVEMBER 1, 2012TABLE 7. PERCENT NAPE SQUAMATION IN SPECIES OF DORATION. Values for holotypes are in boldface.Percent nape squamation0 10 20 30 40 50 60 70 80 90 100 n Mean SDEtheostoma stigmaeumMobile Basin 21 51 56 75 57 49 42 64 37 23 10 485 44.6 26.15Bear Cr. (Tennessee R.) 9 14 13 8 4 4 2 54 40.7 16.35Eastern Gulf coast systems 1 1 2 2 1 6 17 32 20 82 84.8 16.27Central Gulf Coast systems 1 3 13 18 18 11 16 30 21 11 23 165 61.4 26.39Lower Miss. Embayment, eastern tributaries 3 7 7 13 9 23 30 6 1 99 64.5 18.75Upper Miss. Embayment, eastern tributaries 1 3 4 9 16 11 13 57 81.2 15.24Upper Miss. Embayment, western tributaries 2 1 2 1 9 5 22 24 45 21 132 79.5 18.57Lower Miss. Embayment, western tributaries 6 11 17 8 12 12 22 32 28 3 151 61.9 25.40Etheostoma jessiaeLower and middle Tennessee R. 4 16 101 121 98.0 4.76Little Bear Cr. 1 3 3 42 49 97.6 6.62Upper Tennessee R. (excluding Clinch R.) 2 7 212 221 99.5 2.56Clinch R. downstream of Norris Dam 3 11 115 129 98.7 4.03Clinch/Powell R. upstream of Norris Dam 1 3 51 55 99.1 3.48Etheostoma meadiaeUpper Powell River 2 2 3 4 7 11 15 5 49 76.5 18.32Upper Clinch River 1 1 3 11 11 22 39 35 9 132 76.4 15.83Etheostoma akatulo 2 3 12 41 26 35 84 203 86.0 14.87Etheostoma obama 1 1 2 4 14 36 127 185 94.9 9.67Etheostoma goreMiddle Cumberland R. 1 1 3 10 15 35 32 12 109 80.3 13.71Lower Cumberland R. 3 16 37 30 86 90.9 8.21Etheostoma jimmycarter 1 1 2 7 16 33 149 209 95.0 9.81Etheostoma teddyrooseveltArkansas R. 1 4 3 13 82 103 96.6 7.99Upper White R. 1 3 5 8 3 3 6 13 9 18 69 71.6 26.77Etheostoma clinton 9 18 24 19 10 9 6 6 2 - 1 104 30.0 21.41

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 61TABLE 8. DORSAL FIN SPINE AND SOFT RAY COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Doral fin spines9 10 11 12 13 14 n Mean SDEtheostoma stigmaeumMobile Basin 1 41 304 132 7 485 11.2 0.62Bear Cr. (Tennessee R.) 1 32 21 54 11.4 0.52Eastern Gulf coast systems 3 50 29 82 11.3 0.54Central Gulf Coast systems 10 106 49 165 11.2 0.55Lower Miss. Embayment, eastern tributaries 22 67 10 99 10.9 0.56Upper Miss. Embayment, eastern tributaries 6 36 15 57 11.2 0.59Upper Miss. Embayment, western tributaries 1 25 68 38 132 11.1 0.71Lower Miss. Embayment, western tributaries 24 92 33 2 151 11.1 0.65Etheostoma jessiaeLower and middle Tennessee R. 1 52 68 121 12.6 0.52Little Bear Cr. 2 20 27 49 12.5 0.58Upper Tennessee R. (excluding Clinch R.) 2 16 111 89 3 221 12.3 0.67Clinch R. downstream of Norris Dam 5 55 64 5 129 12.5 0.64Clinch/Powell R. upstream of Norris Dam 4 28 23 55 12.3 0.62Etheostoma meadiaeUpper Powell River 7 29 13 49 12.1 0.63Upper Clinch River 1 26 88 16 1 132 11.9 0.61Etheostoma akatulo 10 93 92 8 203 11.5 0.66Etheostoma obama 1 6 83 83 12 185 12.5 0.69Etheostoma goreMiddle Cumberland R. 34 71 3 1 109 11.7 0.56Lower Cumberland R. 1 30 52 2 1 86 11.7 0.60Etheostoma jimmycarter 8 94 92 15 209 12.5 0.69Etheostoma teddyrooseveltArkansas R. 1 42 53 7 103 11.6 0.62Upper White R. 1 34 29 5 69 11.6 0.65Etheostoma clinton 21 59 23 1 104 11.0 0.68

62 Bulletin 30 NOVEMBER 1, 2012TABLE 8. (Continued)Dorsal fin soft rays8 9 10 11 12 13 14 n Mean SDEtheostoma stigmaeumMobile Basin 94 313 78 485 11.0 0.60Bear Cr. (Tennessee R.) 18 34 2 54 10.7 0.54Eastern Gulf coast systems 24 55 3 82 10.7 0.52Central Gulf Coast systems 1 68 91 5 165 10.6 0.58Lower Miss. Embayment, eastern tributaries 20 72 7 99 10.9 0.51Upper Miss. Embayment, eastern tributaries 7 30 20 57 11.2 0.66Upper Miss. Embayment, western tributaries 1 25 78 28 132 11.0 0.66Lower Miss. Embayment, western tributaries 10 99 39 3 151 11.2 0.59Etheostoma jessiaeLower and middle Tennessee R. 17 95 9 121 11.9 0.46Little Bear Cr. 11 34 4 49 11.9 0.54Upper Tennessee R. (excluding Clinch R.) 2 45 148 26 221 11.9 0.59Clinch R. downstream of Norris Dam 37 86 6 129 11.8 0.53Clinch/Powell R. upstream of Norris Dam 1 13 36 4 1 55 11.8 0.66Etheostoma meadiaeUpper Powell River 1 4 32 12 49 11.1 0.63Upper Clinch River 2 61 69 132 11.5 0.53Etheostoma akatulo 25 142 36 203 11.1 0.55Etheostoma obama 1 81 97 6 185 11.6 0.57Etheostoma goreMiddle Cumberland R. 4 64 40 1 109 11.3 0.57Lower Cumberland R. 5 61 20 86 11.2 0.51Etheostoma jimmycarter 10 128 68 3 209 11.3 0.58Etheostoma teddyrooseveltArkansas R. 4 58 40 1 103 11.4 0.58Upper White R. 6 44 19 69 11.2 0.58Etheostoma clinton 27 67 8 2 104 10.9 0.63

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 63TABLE 9. ANAL FIN SOFT RAY AND LEFT PECTORAL FIN RAY COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Anal fin soft rays5-6 7 8 9 10 11 n Mean SDEtheostoma stigmaeumMobile Basin 57 334 94 485 8.1 0.55Bear Cr. (Tennessee R.) 2 44 8 54 8.1 0.42Eastern Gulf Coast systems 19 61 2 82 7.8 0.46Central Gulf Coast systems 39 117 9 165 7.8 0.51Lower Miss. Embayment, eastern tributaries 22 71 6 99 7.8 0.51Upper Miss. Embayment, eastern tributaries 2 38 17 57 8.3 0.52Upper Miss. Embayment, western tributaries 14 94 24 132 8.1 0.53Lower Miss. Embayment, western tributaries 1 8 111 31 151 8.1 0.52Etheostoma jessiaeLower and middle Tennessee R. 2 28 83 8 121 8.8 0.57Little Bear Cr. 1 17 30 1 49 8.6 0.57Upper Tennessee R. (excluding Clinch R.) 1 48 151 21 221 8.9 0.56Clinch R. downstream of Norris Dam 41 80 8 129 8.7 0.56Clinch/Powell R. upstream of Norris Dam 16 34 5 55 8.8 0.59Etheostoma meadiaeUpper Powell River 6 35 8 49 8.0 0.54Upper Clinch River 4 89 39 132 8.3 0.51Etheostoma akatulo 19 160 24 203 8.0 0.46Etheostoma obama 38 119 27 1 185 9.0 0.61Etheostoma goreMiddle Cumberland R. 1 56 50 2 109 8.5 0.55Lower Cumberland R. 1 2 41 40 2 86 8.5 0.70Etheostoma jimmycarter 6 120 83 209 8.4 0.54Etheostoma teddyrooseveltArkansas R. 7 63 32 1 103 8.3 0.59Upper White R. 2 56 11 69 8.1 0.42Etheostoma clinton 1 36 62 5 104 8.7 0.58

64 Bulletin 30 NOVEMBER 1, 2012TABLE 9 (Continued)Left pectoral fin rays11 12 13 14 15 16 n Mean SDEtheostoma stigmaeumMobile Basin 5 110 328 42 485 13.8 0.57Bear Cr. (Tennessee R.) 33 21 54 14.4 0.49Eastern Gulf Coast systems 23 58 1 82 13.7 0.47Central Gulf Coast systems 33 125 7 165 13.8 0.47Lower Miss. Embayment, eastern tributaries 27 66 6 99 13.8 0.54Upper Miss. Embayment, eastern tributaries 7 42 8 57 14.0 0.52Upper Miss. Embayment, western tributaries 35 76 21 132 13.9 0.65Lower Miss. Embayment, western tributaries 2 44 90 15 151 13.8 0.63Etheostoma jessiaeLower and middle Tennessee R. 17 87 17 121 15.0 0.53Little Bear Cr. 6 39 4 49 15.0 0.45Upper Tennessee R. (excluding Clinch R.) 3 90 122 6 221 14.6 0.57Clinch R. downstream of Norris Dam 1 87 41 129 14.3 0.48Clinch/Powell R. upstream of Norris Dam 41 14 55 14.3 0.44Etheostoma meadiaeUpper Powell River 1 31 17 49 14.3 0.52Upper Clinch River 10 92 30 132 14.2 0.53Etheostoma akatulo 3 81 119 203 14.6 0.53Etheostoma obama 5 67 109 4 185 14.6 0.58Etheostoma goreMiddle Cumberland R. 3 74 32 109 14.3 0.50Lower Cumberland R. 8 70 8 86 14.0 0.43Etheostoma jimmycarter 1 88 116 4 209 14.6 0.54Etheostoma teddyrooseveltArkansas R. 5 78 20 103 14.1 0.47Upper White R. 3 55 11 69 14.1 0.44Etheostoma clinton 1 1 51 48 2 1 104 13.5 0.65

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 65TABLE 10. LEFT PREOPERCULOMANDIBULAR CANAL PORE COUNTS IN SPECIES OF DORATION. Values for holotypes are in boldface.Left preoperculomandibular canal pores7-8 9 10 11 12 n Mean SDEtheostoma stigmaeum 3 129 1060 32 1 1225 9.9 0.37Etheostoma jessiae 2 38 525 10 575 9.9 0.31Etheostoma meadiae 1 12 164 4 181 9.9 0.37Etheostoma akatulo 1 28 169 5 203 9.9 0.41Etheostoma obama 1 31 151 2 185 9.8 0.42Etheostoma gore 4 28 161 2 195 9.8 0.45Etheostoma jimmycarter 1 38 166 4 209 9.8 0.44Etheostoma teddyroosevelt 3 47 118 4 172 9.7 0.54Etheostoma clinton 1 95 8 104 9.1 0.29TABLE 11. PRINCIPAL CAUDAL FIN RAY COUNTS AND FREQUENCIES OF INDIVIDUALS WITH A FRENUM AND PALATINE TEETH INETHEOSTOMA JESSIAE FROM THE UPPER TENNESSEE RIVER AND E. MEADIAE. Numbered localities are shown in Figure 7.Etheostoma jessiaeUpper Tennessee R. (excluding Clinch R.)Clinch R. downstream of Norris DamPrincipal caudal fin raysFrenumpresentPalatine teethpresentn 12 13 14 15 16 17 Mean SD No. % No. %221 1 - 4 186 27 3 15.1 0.47 204 92.3 185 83.71. Emory R. and Poplar Cr. 34 1 30 3 15.1 0.34 32 94.1 24 70.62. Beaver Cr. And Bull Run Cr. 25 1 2 21 1 14.9 0.53 25 100.0 22 88.03. Hinds Cr. 24 3 16 4 1 15.1 0.68 24 100.0 23 95.84. Coal Cr. and nearby Clinch R. 46 3 37 4 2 15.1 0.57 45 97.8 37 80.4Clinch/Powell R. upstream of Norris Dam5. Cove Cr. 20 12 8 15.4 0.50 19 95.0 16 80.06. Big Sycamore Cr. 25 23 2 15.1 0.28 20 80.0 16 64.07. Pond opposite Doak’s Dam (Powell R.) 10 10 15.0 0.00 8 80.0 8 80.0Etheostoma meadiaeUpper Powell River49 9 10 30 16.4 0.79 18 36.7 18 36.7Upper Clinch River132 1 2 1 36 43 49 16.0 0.97 88 66.7 41 31.4

66 Bulletin 30 NOVEMBER 1, 2012TABLE 12. MEASUREMENTS IN THOUSANDTHS OF STANDARD LENGTH FOR SPECIES OF DORATION.Measurements for E. akatulo appear in Layman and Mayden (2009). D1 = spinous dorsal fin origin; D2 = soft dorsal fin origin; P2 = lateral pelvic fininsertion; A = anal fin origin. Asterisks indicate significant differences between sexes (P

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 67TABLE 12 (Continued)D1 to D2 292 278–311 11.3 296 282–310 8.86 th dorsal spine length 127* 119–133 4.2 114 104–121 5.7D2 to A body depth 159 144–171 9.2 153 140–161 7.1Soft dorsal fin base length 163 151–177 8.3 161 149–175 7.1Caudal peduncle length 249 241–270 8.8 251 234–267 11.3Caudal peduncle depth 92* 84–97 3.7 88 84–92 2.4Anal fin base length 133* 124–151 9.1 112 94–126 10.81 st anal spine length 72* 58–84 8.5 63 50–74 8.1Pectoral fin length 265 247–282 10.2 264 249–280 10.2Pelvic fin length 217* 185–229 12.3 200 190–210 7.1Trans–pelvic width 76* 72–79 2.3 72 67–78 3.2Body width 133 118–148 9.3 132 120–142 8.4Etheostoma stigmaeum Males (n = 10) Females (n = 10)Clarks River Mean Range SD Mean Range SDStandard length (mm) 38.7 31.8–42.9 3.5 36.7 33.4–39.8 2.3Head length 281* 264–291 7.8 271 258–281 6.6Snout length 76* 70–83 4.7 70 64–76 3.0Predorsal length 350 338–366 8.4 347 339–355 5.7Orbit length 69 65–75 3.1 68 63–72 3.1Upper jaw length 80* 72–85 3.5 74 69–83 4.3D1 to P2 body depth 184 163–214 14.9 189 177–200D1 to D2 284 262–300 11.1 293 277–3196th dorsal spine length 124* 116–132 5.2 114 107–120D2 to A body depth 148* 133–169 11.2 138 129–147Soft dorsal fin base length 169 155–184 8.9 164 156–174Caudal peduncle length 248 235–259 8.3 248 239–259Caudal peduncle depth 87* 80–97 4.6 82 78–89Anal fin base length 142* 129–159 9.3 120 108–1331st anal spine length 72 61–83 6.9 66 57–78Pectoral fin length 279 253–302 13.1 271 253–294Pelvic fin length 238* 223–257 11.1 211 196–224Trans–pelvic width 78 70–83 4.7 75 71–81Body width 130 119–153 9.8 127 119–137

68 Bulletin 30 NOVEMBER 1, 2012TABLE 12 (Continued)Etheostoma stigmaeum Males (n = 10) Females (n = 10)Ouachita River Mean Range SD Mean Range SDStandard length (mm) 34.7 31.4–42.3 3.5 34.3 31.2–37.4 1.7Head length 277 265–291 8.4 277 264–285 7.2Snout length 71 66–77 3.2 72 63–78 4.9Predorsal length 340 313–364 14.9 347 331–358 8.9Orbit length 69 60–77 5.2 69 61–73 3.5Upper jaw length 78 72–84 4.2 76 66–81 4.5D1 to P2 body depth 181* 169–196 9.3 196 183–214 8.7D1 to D2 299 270–321 14.5 290 282–299 6.16 th dorsal spine length 123* 104–149 12.1 114 104–123 5.3D2 to A body depth 162 149–177 8.1 157 147–164 4.7Soft dorsal fin base length 163 146–172 7.9 164 149–182 10.9Caudal peduncle length 248 230–259 10.2 248 225–260 10.2Caudal peduncle depth 92 88–99 3.7 88 84–96 4.0Anal fin base length 128* 113–142 8.7 112 92–137 14.41st anal spine length 73* 64–86 6.8 61 51–67 5.6Pectoral fin length 269 243–301 18.4 269 248–286 14.0Pelvic fin length 228* 214–251 10.6 207 183–226 11.0Trans–pelvic width 74 67–80 4.0 73 66–79 4.0Body width 130 123–139 5.6 135 122–147 8.5Etheostoma jessiae Males (n = 30)Tennessee River Mean Range SDFemales (n = 30)Mean Range SDStandard length (mm) 46.4 32.8–57.7 6.4Head length 290* 273–310 8.9Snout length 88* 79–101 6.0Predorsal length 354 332–373 10.2Orbit length 65 58–75 3.7Upper jaw length 86* 76–96 4.9D1 to P2 body depth 180 159–203 12.2D1 to D2 300 268–326 14.843.1 30.7–51.7 5.8285 263–303 10.083 72–94 5.5352 333–367 9.465 57–76 4.583 71–94 5.5178 160–210 12.7305 275–346 15.8

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 69TABLE 12 (Continued)6th dorsal spine length 119* 100–139 9.4108 92–122 7.9D2 to A body depth 146* 133–163 8.0139 128–154 5.9Soft dorsal fin base length 170 155–186 6.9167 150–183 8.3Caudal peduncle length 222 191–238 10.0219 204–243 9.1Caudal peduncle depth 83 73–94 5.281 72–89 4.3Anal fin base length 145* 129–170 10.5133 118–151 7.81st anal spine length 71* 53–85 8.164 50–75 5.8Pectoral fin length 270 233–296 12.9 266 234–303 17.9Pelvic fin length 222* 194–249 3.8 208 184–230 10.3Trans–pelvic width 75* 67–81 4.0 71 64–79 3.9Body width 134* 120–153 7.7 129 111–147 8.5Etheostoma meadiae Males (n = 12) Females (n = 12)Clinch and Powell rivers Mean Range SD Mean Range SDStandard length (mm) 43.1 34.8–47.6 4.1 40.8 32.5–46.3 4.1Head length 287 273–304 9.6 281 275–289 5.4Snout length 79 73–86 4.2 76 71–87 4.3Predorsal length 352 337–365 9.1 355 346–368 6.9Orbit length 68 w63–78 4.4 67 62–72 2.6Upper jaw length 84 * 73–94 6.4 78 72–84 3.9D1 to P2 body depth 199 186–219 11.3 196 188–208 6.6D1 to D2 314 297–336 11.2 316 299–331 10.16 th dorsal spine length 124* 114–132 6.6 112 94–121 7.1D2 to A body depth 168* 155–195 11.8 156 142–171 8.9Soft dorsal fin base length 165* 154–172 5.7 157 144–171 8.7Caudal peduncle length 222 214–235 7.1 226 210–236 7.1Caudal peduncle depth 95* 84–107 6.0 89 80–100 6.2Anal fin base length 143* 126–161 9.1 124 112–139 7.21 st anal spine length 90* 80–106 8.1 83 71–93 5.6Pectoral fin length 284 259–319 16.7 282 251–302 12.8Pelvic fin length 232* 214–249 10.8 222 211–233 6.7Trans–pelvic width 79* 68–93 6.9 73 67–78 3.8Body width 141 131–165 9.2 139 130–149 6.1

70 Bulletin 30 NOVEMBER 1, 2012TABLE 12 (Continued)Etheostoma obama Males (n = 10) Females (n = 10)Duck River Mean Range SD Mean Range SDStandard length (mm) 39.8 36.1–43.4 2.8 35.6 31.4–39.1 2.1Head length 276* 270–284 4.6 267 252–277 8.4Snout length 76 70–82 4.1 72 64–77 4.1Predorsal length 340 330–348 5.0 337 326–353 7.6Orbit length 68 64–71 2.6 67 62–73 3.8Upper jaw length 78* 75–82 2.4 71 69–76 2.2D1 to P2 body depth 185 176–191 4.9 183 171–201 8.2D1 to D2 318 308–336 10.0 321 311–332 7.46th dorsal spine length 131* 119–147 10.1 117 107–125 5.9D2 to A body depth 157* 149–162 3.7 140 121–151 7.9Soft dorsal fin base length 172 161–180 5.8 166 146–180 9.3Caudal peduncle length 233 223–248 7.6 232 209–253 12.5Caudal peduncle depth 85* 82–91 3.2 80 74–83 2.8Anal fin base length 157* 141–166 8.5 134 124–148 7.31st anal spine length 75* 65–86 6.6 69 63–75 3.7Pectoral fin length 308* 287–329 12.2 290 272–319 13.0Pelvic fin length 250* 237–262 8.4 227 217–241 8.8Trans–pelvic width 76* 70–81 3.5 71 68–77 3.2Body width 135* 129–141 4.1 127 117–135 6.4Etheostoma gore Males (n = 20) Females (n = 20)lower Cumberland River Mean Range SD Mean Range SDStandard length (mm) 38.1 34.4–42.3 2.1 36.3 33.0–40.0 2.1Head length 265* 254–282 7.2 258 241–274 7.0Snout length 73 66–80 3.8 70 61–79 4.4Predorsal length 339 327–353 8.8 337 318–351 7.9Orbit length 65 59–71 3.8 63 59–71 3.0Upper jaw length 71* 64–76 3.3 68 61–73 3.1D1 to P2 body depth 175 156–190 9.7 180 154–208 12.5

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 71TABLE 12 (Continued)D1 to D2 300* 286–318 7.4 306 287–329 10.76th dorsal spine length 117* 105–126 5.5 106 94–119 7.6D2 to A body depth 154* 141–165 7.3 145 135–155 6.5Soft dorsal fin base length 165 148–191 8.7 160 142–176 9.2Caudal peduncle length 239 221–256 9.8 241 219–263 11.6Caudal peduncle depth 85* 76–93 4.4 80 72–87 3.6Anal fin base length 141* 126–155 7.5 126 94–142 12.51st anal spine length 76* 55–83 6.9 67 59–77 4.7Pectoral fin length 293* 263–316 17.6 279 254–307 11.7Pelvic fin length 235* 208–263 14.3 217 198–239 10.5Trans–pelvic width 76* 69–85 5.1 71 62–79 4.5Body width 127* 116–141 5.7 123 117–136 5.1Etheostoma jimmycarter Males (n = 12) Females (n = 12)Barren River Mean Range SD Mean Range SDStandard length (mm) 43.4 35.9–51.8 4.0 38.6 33.0–41.5 2.6Head length 271* 262–281 6.2 262 256–274 5.4Snout length 76 69–86 5.9 72 65–79 4.3Predorsal length 333 319–345 8.0 330 319–343 6.7Orbit length 62 59–68 3.0 64 60–70 3.1Upper jaw length 77* 74–83 2.5 70 66–73 2.5D1 to P2 body depth 181 165–197 11.0 189 178–199 7.1D1 to D2 316 302–326 8.0 319 299–339 11.46th dorsal spine length 128* 115–138 8.2 116 104–139 11.0D2 to A body depth 153* 140–166 7.6 147 140–156 5.8Soft dorsal fin base length 158 144–181 10.3 154 142–166 7.0Caudal peduncle length 237 227–257 8.6 243 223–252 7.5Caudal peduncle depth 84 78–92 4.0 83 78–90 3.7Anal fin base length 145* 131–161 10.1 124 112–136 7.51st anal spine length 82* 76–93 4.9 72 65–79 4.5Pectoral fin length 295 275–316 13.2 291 271–308 11.7Pelvic fin length 238* 223–257 8.1 226 201–243 11.3Trans–pelvic width 77* 71–83 3.3 73 68–78 3.1Body width 133 116–140 7.0 130 120–143 7.0

72 Bulletin 30 NOVEMBER 1, 2012TABLE 12 (Continued)Etheostoma teddyroosevelt Males (n = 10) Females (n = 10)middle Arkansas River Mean Range SD Mean Range SDStandard length (mm) 39.1 31.9–43.1 3.9 36.4 30.9–40.0 3.4Head length 268* 259–276 5.8 261 249–269 5.9Snout length 73* 63–81 5.4 68 62–75 4.4Predorsal length 339 328–354 9.6 335 319–350 9.8Orbit length 63 57–70 4.0 64 61–70 3.5Upper jaw length 72* 65–77 4.4 67 63–75 3.9D1 to P2 body depth 191 172–212 13.6 199 181–215 9.8D1 to D2 296* 273–317 11.9 311 292–332 14.56th dorsal spine length 109 95–123 7.8 109 97–118 7.1D2 to A body depth 160 152–170 7.3 154 144–170 7.6Soft dorsal fin base length 164 153–179 7.5 158 145–170 7.9Caudal peduncle length 247 231–263 9.9 244 222–264 15.7Caudal peduncle depth 87* 81–92 4.2 81 76–87 3.3Anal fin base length 137* 123–146 7.3 128 114–140 8.01st anal spine length 65 54–72 6.8 62 48–73 8.1Pectoral fin length 278 258–299 12.5 269 254–280 9.4Pelvic fin length 222* 207–238 9.8 212 191–227 10.9Trans–pelvic width 79* 73–84 4.2 73 65–81 4.3Body width 140 129–155 8.0 135 123–147 8.4Etheostoma clintonCaddo and Ouachita riversMales (n = 12) Females (n = 12)Mean Range SD Mean RangeSDStandard length (mm)Head lengthSnout lengthPredorsal lengthOrbit lengthUpper jaw length30.9 28.0–34.2 1.9 30.9 28.7–33.9272 259–284 7.3 270 255–28564 60–69 3.0 64 56–72344 331–354 8.0 344 330–35666 59–70 3.2 66 62–7273* 71–76 1.5 70 65–761.68.94.46.92.82.8

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 73TABLE 12 (Continued)D1 to P2 body depth 153 140–165 7.2 171 136–187 14.5D1 to D2 280* 260–287 8.2 290 264–309 13.26th dorsal spine length 113* 102–128 8.4 101 93–110 6.2D2 to A body depth 134 126–145 5.7 135 122–143 6.4Soft dorsal fin base length 156* 145–175 9.1 148 134–170 8.8Caudal peduncle length 238 222–247 6.7 240 216–257 12.0Caudal peduncle depth 84 73–90 5.5 82 77–89 3.2Anal fin base length 140* 126–151 6.6 115 102–122 6.51st anal spine length 66* 51–81 7.3 60 54–69 4.3Pectoral fin length 262 243–279 10.9 259 230–278 14.2Pelvic fin length 225* 202–241 11.8 209 180–226 13.7Trans–pelvic width 64 55–71 4.9 63 60–67 2.1Body width 111 103–119 4.7 117 101–130 8.4

74 Bulletin 30 NOVEMBER 1, 2012TABLE 13. CHARACTERS USEFUL IN DISTINGUISHING SPECIES OF DORATION. Color characters are for breeding males.Character E. stigmaeum E. jessiae E. meadiae E. akatulo E. obamaFrenum absent present present/absent absent absentMean lateral scale rows 46.4 50.4 46.9 45.3 46.7Mean unpored lateral scales 12.9 7.9 4.1 0.2 14.0Mean % cheek squamation 38.5 20.7 1.0 91.0 21.1Modal caudal peduncle scales 14-16 17-18 16 15 16Modal transverse scales 11-13 14-15 13-14 12 12Modal scales below lateral line 6-7 7-8 6 6 6Modal scales above lateral line 4-5 5-6 6 4 5Modal dorsal spines 11 12-13 12 11-12 12-13Modal soft dorsal rays 11 12 11-12 11 12Modal principal caudal rays 15 15 17 15 15Modal soft anal rays 8 9 8 8 9Modal pectoral rays 14 14-15 14 15 15Palatine teeth present present absent/present absent absent/presentPreoperculomandibular pores 10 10 10 10 10Maximum size (mm SL) 48.9 64.6 54.5 48.0 48.3Face coloration gray w/blue-green gray w/blue gray w/blue entirely blue orange w/blueBlue on lips present present present present absentBlue on mid-gular area present absent absent present absentOrange spots on soft dorsal and caudal fin rays absent present absent absent presentBlue in base of soft dorsal and anal fin present present present absent presentSpinous dorsal fin marginal and submedial bands blue-green blue blue black blueSpinous dorsal fin medial band red-orange orange orange red-orange orangeOrange in spinous dorsal fin basal band absent present absent absent presentBasicaudal bar development dorsal/ventral dorsal/ventral dorsal/ventral dorsal/ventral dorsal/ventralOrange spots on anal fin rays absent absent absent absent absent/presentLateral blue blotches elongate elongate elongate elongate quadrate

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 75TABLE 13 (Continued)Character E. gore E. jimmycarter E. teddyroosevelt E. clintonFrenum absent absent absent absentMean lateral scale rows 46.1 45.4 48.1 49.9Mean unpored lateral scales 12.1 15.2 16.1 16.6Mean % cheek squamation 5.4 3.9 0.9 9.7Modal caudal peduncle scales 16 16 16 16Modal transverse scales 12 12 13 13-14Modal scales below lateral line 6 6 7 8Modal scales above lateral line 4 5 5 4Modal dorsal spines 12 12-13 11-12 11Modal soft dorsal rays 11 11 11 11Modal principal caudal rays 15 15 15 15Modal soft anal rays 8-9 8 8 9Modal pectoral rays 14 15 14 13-14Palatine teeth absent absent usually absent presentPreoperculomandibular pores 10 10 10 9Maximum size (mm SL) 42.8 49.0 43.4 34.6Face coloration orange w/blue orange w/blue gray w/blue gray w/blue-greenBlue on lips absent absent absent presentBlue on mid-gular area absent absent absent presentOrange spots on soft dorsal and caudal fin rays present present present (faint) absentBlue in base of soft dorsal and anal fin present absent absent presentSpinous dorsal fin marginal and submedial bands blue black black blue-greenSpinous dorsal fin medial band orange orange orange red-orangeOrange in spinous dorsal fin basal band present present absent absentBasicaudal bar development dorsal/ventral ventral ventral ventralOrange spots on anal fin rays present present present (faint) absentLateral blue blotches quadrate quadrate quadrate quadrate

76 Bulletin 30 NOVEMBER 1, 2012TABLE 14. PRINCIPAL COMPONENT LOADINGS FOR 17 MERISTIC VARIABLES IN 3,049 SPECIMENS OFDORATION.Principal componentVariable I IIDorsal fin spines 0.540 0.325Dorsal fin rays 0.586 0.235Principal caudal fin rays 0.086 0.130Anal fin rays 0.554 0.100Pectoral fin rays 0.415 0.454Lateral scale rows 0.599 -0.266Unpored lateral scale rows -0.131 -0.459Transverse scale rows 0.752 -0.418Scale rows below lateral line 0.629 -0.461Scale rows above lateral line 0.705 -0.028Caudal peduncle scale rows 0.722 -0.290Percent cheek squamation -0.363 0.320Percent nape squamation 0.415 0.724Percent belly squamation 0.315 0.682Percent opercle squamation -0.435 0.127Infraorbital canal pores 0.128 0.069Preoperculomandibular canal pores 0.061 0.210

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 77TABLE 15. SHEARED PRINCIPAL COMPONENT (PC) LOADINGS FOR 28 MORPHOMETRIC MEASURE-MENTS IN 326 SPECIMENS OF DORATION. D1 = spinous dorsal fin origin; D2 = soft dorsal fin origin; IOW =midline at least interobital width; P2 = lateral pelvic fin insertion; A = anal fin origin.Males (n = 163) Females (n = 163)Sheared PCSheared PCMeasurement Size II III Size II IIIStandard length 0.166 0.053 -0.087 0.176 0.018 -0.013Head length 0.168 0.150 -0.130 0.176 0.205 -0.071Snout length 0.203 0.353 -0.251 0.207 0.357 -0.284Orbit length 0.153 0.067 0.032 0.175 0.141 0.073Upper jaw length 0.213 0.214 -0.175 0.222 0.370 -0.062Predorsal length 0.178 0.139 -0.066 0.181 0.136 -0.003D1 to occiput 0.204 0.114 -0.022 0.204 0.131 0.134Occiput to snout 0.168 0.154 -0.101 0.173 0.177 -0.118Occiput to IOW 0.160 0.053 0.066 0.156 0.061 0.045IOW to snout 0.187 0.274 -0.192 0.190 0.274 -0.223P2 to snout 0.170 0.086 -0.069 0.175 0.192 -0.079Occiput to P2 0.210 -0.008 0.180 0.203 -0.057 0.105D1 to P2 0.231 -0.034 0.380 0.228 -0.230 0.271D1 to D2 0.184 -0.094 -0.093 0.187 -0.188 -0.040D1 to A 0.193 -0.027 0.045 0.208 -0.127 0.082P2 to A 0.181 0.051 -0.021 0.202 -0.058 0.074D2 to P2 0.188 -0.049 -0.008 0.193 -0.143 0.042D2 to A 0.219 -0.107 0.395 0.212 -0.148 0.272Soft dorsal fin base length 0.171 0.091 -0.084 0.195 0.019 -0.093Caudal peduncle depth 0.195 -0.052 0.381 0.192 -0.003 0.244Caudal peduncle length 0.147 0.055 0.017 0.164 -0.028 0.180Anal fin base length 0.194 -0.068 -0.275 0.154 -0.196 -0.4921st anal spine length 0.183 -0.708 -0.374 0.148 -0.484 -0.493Petoral fin length 0.189 -0.147 0.002 0.186 -0.139 -0.039Pelvic fin length 0.182 -0.184 -0.030 0.170 -0.149 -0.072Trans-pelvic width 0.221 -0.097 0.188 0.209 -0.050 0.1136th dorsal spine length 0.188 -0.218 -0.167 0.151 -0.100 -0.030Body width 0.214 -0.005 0.210 0.219 -0.100 0.157

78 Bulletin 30 NOVEMBER 1, 2012TABLE 16. PRINCIPAL COMPONENT LOADINGSFOR 16 MERISTIC VARIABLES IN 756 SPECIMENSOF ETHEOSTOMA JESSIAE AND E. MEADIAE. Bellysquamation did not vary and was omitted from theanalysis.PrincipalcomponentVariable I IIDorsal fin spines 0.201 -0.305Dorsal fin rays 0.416 -0.344Principal caudal fin rays -0.541 0.354Anal fin rays 0.515 -0.329Pectoral fin rays 0.329 -0.136Lateral scale rows 0.601 -0.144Unpored lateral scale rows 0.354 -0.371Transverse scale rows 0.732 0.545Scale rows below lateral line 0.716 0.482Scale rows above lateral line 0.275 0.620Caudal peduncle scale rows 0.731 0.249Percent cheek squamation 0.683 -0.050Percent nape squamation 0.684 -0.280Percent opercle squamation -0.180 0.322Infraorbital canal pores 0.125 0.120Preoperculomandibular canal pores 0.008 -0.001

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The fishes of North andMiddle America: a descriptive catalogue of the species of fishlikevertebrates found in the waters of North America, north ofthe Isthmus of Panama. Bulletin of the United States NationalMuseum 47, Part 1.Jordan, D. S., and B. W. Evermann. 1898. The fishes of North andMiddle America: a descriptive catalogue of the species of fishlikevertebrates found in the waters of North America, north ofthe Isthmus of Panama. Bulletin of the United States NationalMuseum 47, Part 3.Jordan, D. S., and B. W. Evermann. 1900. The fishes of North andMiddle America: a descriptive catalogue of the species of fishlikevertebrates found in the waters of North America, north ofthe Isthmus of Panama. Bulletin of the United States NationalMuseum 47, Part 4.Jordan, D. S., and C. H. Gilbert. 1883. Synopsis of the fishes ofNorth America. Smithsonian Miscellaneous Collection XXIV,Bulletin of the United States National Museum 16.Jordan, D. S., B. W. Evermann, and H. W. Clark. 1930. Checklist of the fishes and fishlike vertebrates of North and MiddleAmerica north of the northern boundary of Venezuela andColombia. Report of the United States Commisison of Fisheriesfor 1928, Part II.Kirsch, P. H. 1893. Notes on a collection of fishes from the southerntributaries of the Cumberland River in Kentucky andTennessee. Bulletin of the United States Fisheries CommissionXI for 1891:259–268.Kluge, A. G., and J. S. Farris. 1969. Quantitative phyletics and theevolution of anurans. Systematic Zoology 18:1–32.Kuehne, R. A., and R. W. Barbour. 1983. The American darters.University Press Kentucky, Lexington.Kuhne, E. R. 1939. A guide to the fishes of Tennessee and the mid-South. Tennessee Department of Conservation, Nashville.Lang, N. J., and R. L. Mayden. 2007. Systematics of the subgenusOligocephalus (Teleostei: Percidae: Etheostoma) with completesubgeneric sampling of the genus Etheostoma. MolecularPhylogenetics and Evolution 43:605–615.Layman, S.R. 1994. Phylogenetic systematics and biogeographyof darters of the subgenus Doration (Percidae: Etheostoma).Unpublished Ph.D., dissertation, University of Alabama,Tuscaloosa.Layman, S.R., and R. L. Mayden. 2009. A new species of the dartersubgenus Doration (Percidae: Etheostoma) from the Caney ForkRiver system, Tennessee. Copeia 2009:157–170.Layman, S. R., A. M. Simons, and R. M. Wood. 1993. Status ofthe dirty darter, Etheostoma olivaceum, and bluemask darter,Etheostoma (Doration) sp., with notes on fishes of the CaneyFork River system, Tennessee. Journal Tennessee Academy ofSciences 68(2):65–70.Leviton, A. E., and R. H. Gibbs, Jr. 1988. Standards in herpetologyand ichthyology. Standard symbolic codes for institutionresource collections in herpetology and ichthyology.Supplement No.1: additions and corrections. Copeia 1988:280–282.Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985.Standards in herpetology and ichthyology: Part I. Standardsymbolic codes for institutional resource collections in herpetologyand ichthyology. Copeia 1985:802–832.Maddison, W. P., M. J. Donoghue, and D. R. Maddison. 1984.Outgroup analysis and parsimony. Systematic Zoology 33:83–103.Mayden, R. L. 1985. Biogeography of Ouachita Highland fishes.Southwestern Naturalist 30:195–211.Mayden, R. L. 1997. A hierarchy of species concepts: the denouementin the saga of the species problem, p. 381–424. In:Species: The Units of Biodiversity. Claridge, M.F., Dawah, H.A.and Wilson, M.R. (eds). Chapman and Hall Ltd., London.Mayden, R. L. 1999. Consilience and a hierarchy of species concepts:Advances towards closure on the species puzzle. Journalof Nematology 31:95–116Mayden, R. L. 2002. On biological species, species concepts, andindividuation in the natural world. Fish and Fisheries 2:1–26.Mayden, R. L., B. M. Burr, L. M. Page, and R. R. Miller. 1992.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 81The native freshwater fishes of North America, p. 827–863. In:Systematics, historical ecology, and North American freshwaterfishes. R. L. Mayden (ed.). Stanford University Press, Stanford,California.Meek, S. E. 1891. Report of explorations made in Missouri andArkansas during 1889, with an account of the fishes observed ineach of the river basins examined. Bulletin of the United StatesFisheries Commission IX for 1889:113–141.Menhinick, E. F., T. M. Burton, and J. R. Bailey. 1974. An annotatedchecklist of the freshwater fishes of North Carolina. Journalof the Elisha Mitchell Scientific Society 90:24–50.Metcalf, A. L. 1959. Fishes of Chautauqua, Cowley and Elk counties,Kansas. Miscellaneous Publications, University of KansasMuseum of Natural History 11:345–400.Miller, R. J., and H. W. Robison. 1973. Fishes of Oklahoma.Oklahoma State University Press, Stillwater.Near, T. J., C. M. Bossu, G. S. Bradburd, R. L. Carlson, R. C.Harrington, P. R. Hollingsworth, B. J. Keck, and D. A. Etneir.2011. Phylogeny and Temporal Diversification of Darters(Percidae: Etheostomatinae). Systematic Biology 60:565–595.Page, L. M. 1981. The genera and subgenera of darters (Percidae,Etheostomatini). Occasional Papers Museum of NaturalHistory, University of Kansas 90.Page, L. M. 1983. Handbook of darters. TFH Publications,Neptune City, New Jersey.Page, L. M. 1985. Evolution of reproductive behaviors in percidfishes. Bulletin Illinois Natural History Survey 33:275–295.Page, L. M., and L. E. Cordes. 1983. Variation and systematicsof Etheostoma euzonum, the Arkansas saddled darter (Pisces:Percidae). Copeia 1983:1042–1050.Page, L. M., and K. S. Cummings. 1984. A portable camera box forphotographing small fishes. Environmental Biology of Fishes11:160.Pflieger, W. L. 1975. The fishes of Missouri. Missouri Departmentof ConservationRobins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner,R. N. Lea, and W. B. Scott. 1980. A list of common and scientificnames of fishes from the United States and Canada, 4th ed.American Fisheries Society Special Publication 12.Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner,R. N. Lea, and W. B. Scott. 1991. Common and scientific namesof fishes from the United States and Canada, 5th ed. AmericanFisheries Society Special Publication 20.Robison, H. W. 1974. An additional population of Etheostoma pallididorsumDistler and Metcalf in Arkansas. American MidlandNaturalist 91:478–479.Robison, H. W., and T. M. Buchanan. 1988. Fishes of Arkansas.University of Arkansas Press, Fayetteville.Robison, H. W., and J. L. Harris. 1978. Notes on the habitat andzoogeography of Noturus taylori (Pisces: Ictaluridae). Copeia1978:548–550.Sanderson, M. J. 1989. Confidence limits on phylogenies: the bootstraprevisited. Cladistics 5:113–129.SAS Institute Inc. 1985. SAS user’s guide: statistics, version 5 edition.SAS Institute Inc., Cary, North Carolina.SAS Institute Inc. 1990. SAS procedures guide, version 6, thirdedition. SAS Institute Inc., Cary, North Carolina.Simons, A. M. 1991. Phylogenetic relationships of the crystal darter,Crystallaria asprella (Teleostei: Percidae). Copeia 1991:927–936.Simons, A. M. 1992. Phylogenetic relationships of the Boleosoma speciesgroup (Percidae: Etheostoma), p. 268–292. In: Systematics,historical ecology, and North American freshwater fishes. R. L.Mayden (ed.). Stanford University Press, Stanford, California.Sloss, B. L., N. Billington and B. M. Burr. 2004. A molecular phylogenyof the Percidae (Teleostei, Perciformes) based on mitochondrialDNA sequence. Mol. Phylogenet. Evol. 32, 545–562.Starnes, W. C., and D. A. Etnier. 1986. Drainage evolution andfish biogeography of the Tennessee and Cumberland Riversdrainage realm, p. 325–362. In: The zoogeography of NorthAmerican freshwater fishes. C. H. Hocutt and E. O. Wiley(eds.). John Wiley & Sons, New York.Swofford, D. L. 1993. PAUP: phylogenetic analysis using parsimony,version 3.1. Computer program distributed by IllinoisNatural History Survey, Champaign.Swofford, D. L., and W. P. Maddison. 1987. Reconstructing ancestralstates under Wagner parsimony. Mathematical Bioscience87:199–229.Swofford, D. L., and G. J. Olsen. 1990. Phylogeny reconstruction,p. 411–501. In: Molecular systematics. D. M. Hillis and C. Moritz(eds.). Sinauer Associates, Inc., Sunderland, Massachusetts.Wall, B. R. 1968. Studies on the fishes of the Bear Creek drainageof the Tennessee River system. Unpubl. M.S. thesis, Universityof Alabama, Tuscaloosa.Watrous, L. E., and Q. D. Wheeler. 1981. The outgroup comparisonmethod of character analysis. Systematic Zoology 30:1–11.Wiley, E. O. 1978. The evolutionary species concept reconsidered.Systematic Zoology 27:17–26.Wiley, E. O. 1981. Phylogenetics, the theory and practice of phylogeneticsystematics. 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82 Bulletin 30 NOVEMBER 1, 2012APPENDIX A. CHARACTER DESCRIPTIONS1. Breeding male lateral body banding pattern: 0 = noseries of 7-11 blue or blue-green vertically aligned bars orquadrate blotches; 1 = series of 7-11 vertically elongate blueor blue-green bars, typically extending above lateral linescale row; 2 = series of 7-11 quadrate blue blotches, usuallynot extending above lateral line scale row. The blue barsof the outgroup Etheostoma caeruleum, which are obliquelyaligned and number up to 14, are not considered homologous.2. Breeding male lateral body coloration: 0 = no scatteredred-orange spots or irregular markings on midlateraland dorsolateral areas; 1 = scattered red-orange spots orirregular markings on midlateral and dorsolateral areas.The lateral red pigment in E. caeruleum tends to be concentratedon the lower sides and is more consistent in distribution,typically covering the central exposed portions ofscales; this condition is not considered homologous.3. Breeding male lateral head coloration: 0 = no brightiridescent blue or blue-green pigment on the opercle andcheek; 1 = bright iridescent blue or blue-green pigment onthe opercle and cheek. The blue pigment in E. caeruleumtypically lacks the iridescent quality of the blue/blue-greenin species of Doration and is not considered homologouson the basis of the criterion of special similarity (Wiley,1981).4. Breeding male lip coloration: 0 = blue or blue-greenpigment absent; 1 = blue or blue-green pigment usuallypresent on a portion of the lips.5. Breeding male gular area: 0 = no blue or blue-greenpigment present; 1 = blue or blue-green pigment presentas medial bar or completely covering area.6. Breeding male base color of anterior face and undersideof head: 0 = without tangerine orange pigment; 1 =covered with tangerine orange pigment. The orange-redpigment on the underside of the head of E. caeruleum doesnot extend onto the anterior face and is not consideredhomologous.7. Breeding male first dorsal fin pattern: 0 = no distinctmarginal and submedial bands of the same color; 1= dusky to black marginal and submedial bands; 2 = blueor blue-green marginal and submedial bands. In speciesof Doration (ingroup) the marginal and submedial bandsare always the same color (blue or blue-green in marginalband may be restricted to posterior membranes) and are assumedto be coupled. This character can be ordered basedon the observation that melanization often accompaniesblue/blue-green pigment in these bands, and in males enteringreproductive condition the bands are dusky to blackbefore blue/blue-green appears.8. Breeding male first dorsal fin medial zone: 0 = nodistinct medially-restricted red or orange band; 1 = medialband red or red-orange in life; 2 = medial band trueorange in life.9. Breeding male first dorsal fin basal zone: 0 = nobright orange triangles or streaks in posterior portions ofmembranes; 1 = bright orange triangles or streaks in posteriorportions of membranes.10. Breeding male second dorsal fin membranes: 0 = noblue or blue-green pigment in base of fin above fourth dorsalsaddle; 1 = blue or blue-green in base of fin above fourthdorsal saddle (often accompanied by blue/blue-green spotor submedial band across anterior membranes).11. Breeding male second dorsal and caudal fin rays: 0= no discrete orange spots (may be suffused with pale orangeor yellow); 1 = discrete, often bright orange spots. InE. teddyroosevelt these spots are small and tend to be paleron the caudal fin, but they are usually discrete.12. Breeding male caudal fin base at peak development:0 = no blue or blue-green bar; 1 = blue or blue-green barextending from medial caudal fin base to ventral marginof caudal fin; 2 = blue or blue-green vertical bar extendingfrom dorsal to ventral margins of caudal fin. This charactercan be ordered based on the observation that malesdeveloping character state 2 pass through a stage in whichthe bar first develops ventrally (state 1).13. Breeding male anal fin membranes: 0 = no blue orblue-green pigment in base of fin: 1 = blue or blue-greenpigment, often forming broad band, in base of fin.14. Breeding male anal fin rays: 0 = no discrete orangespots; 1 = discrete orange spots restricted to posteromedialportion of fin, usually only one per ray; 2 = discrete orangespots throughout fin, usually two or three per ray. In E. teddyroosevelt,the orange spots are often pale but discrete andmay be most discernible in males that have not yet reachedpeak development; melanization of the fin at peak developmentcan obscure these spots.15. Breeding male pectoral fin rays: 0 = no discrete orangespots along entire lengths of rays (pale yellow-orangespots may be concentrated near base of fin); 1 = discrete,often bright orange spots along entire lengths of rays. In E.teddyroosevelt, the orange spots tend to be pale.16. Breeding tubercles on ventral body scales: 0 = present;1 = absent.17. Breeding tubercles on anal and pelvic fin rays: 0 =absent; 1 = present.18. Structure of breeding female genital papilla: 0 =conical, tubular; 1 = flat, bifurcate; 2 = rugose, spatulate.19. Non-breeding male and female midlateral body pattern:0 = no blotches formed by X-, V-, or W-shaped markings;1 = blotches formed by X-, V-, or W-shaped markings.20. Non-breeding male and female caudal fin base: 0= no pair of vertically aligned small dark spots at insertionof medial rays; 1 = pair of vertically aligned small darkspots at insertion of medial rays (on scales sheathing baseof rays, not on musculature; in breeding males often obscuredby nuptial coloration).21. Number of dorsal saddles: 0 = 6 to 10; 1 = 4; 2 = 6.

Layman & Mayden Morphological Diversity and Phylogenetics of the Darter subgenus Doration (Percidae: Etheostoma) 8322. Frenum: 0 = present (65-100% individuals); 1 = absent(90-100% individuals).23. Lateral line: 0 = complete; 1 = incomplete.24. Palatine teeth: 0 = present (75-100% individuals); 1= absent (75-100% individuals).25. Infraorbital canal: 0 = uninterrupted; 1 = interrupted.26. Supratemporal canal: 0 = uninterrupted; 1 = interrupted.27. Preoperculomandibular pores: 0 = 10; 1 = 9.28. Cheek squamation: 0 = fully scaled; 1 = partly scaled;2 = naked or nearly so.29. Breast squamation: 0 = scaled; 1 = unscaled.30. Number of anal fin spines: 0 = 2; 1 = 1.31. Modal (mean) number of anal fin soft rays: 0 = 10(10.0); 1 = 7-9 (7.5-9.0); 2 = 6-7 (6.7).32. Modal (mean) number of dorsal fin spines: 0 = 13(13.0); 1 = 11-13 (10.9-12.6); 2 = 8-10 (

MUSEUM BULLETIN SERIES (1975 -)1. Systematics of the Percid Fishes of the Subgenus Ammocrypta, Genus Ammocrypta, with Descriptions of TwoNew Species. James D. Williams. 56 pp., illus., June, 1975. $5.002. Endangered and Threatened Plants and Animals of Alabama. Herbert Boschung, Editor. 93 pp., illus.,October, 1976. $7.503. Containing: A New Species of Semotilus (Pisces: Cyprinidae) from the Carolinas. Franklin F. Snelson, Jr. andRoyal D. Suttkus. Etheostoma neopterum, a New Percid Fish from the Tennessee River System in Alabamaand Tennessee. W. Mike Howell and Guido Dingerkus. Taxonomy, Ecology and Phylogeny of the SubgenusDepressicambarus, with the Description of a New Species from Florida and Redescriptions of Cambarus graysoni,Cambarus latimanus, and Cambarus striatus (Decapoda: Cambaridae). Raymond William Bouchard. 60pp., illus., February, 1978. $5.004. Systematics of the Percid Fishes of the Subgenus Microperca, Genus Etheostoma. Brooks M. Burr. 53 pp., illus.,July 1978. $5.005. Containing: Notropis candidus, a New Cyprinid Fish from the Mobile Bay Basin, and a Review of the NomenclaturalHistory of Notropis shumardi (Girard). Royal D. Suttkus. Notropis stanauli, a New Madtom Catfish(Ictaluridae) from the Clinch and Duck Rivers, Tenessee. David A. Etnier and Robert E. Jenkins. 23 pp.,illus., May, 1980. $5.006. Containing: A New Species of Cyprinodontid Fish, Genus Fundulus (Zygonectes), from Lake PontchartrainTributaries in Louisiana and Mississippi. Royal D. Suttkus and Robert C. Cashner. Karyotypes in Populationsof the Cyprinodontid Fishes of the Fundulus notatus species-complex: A Geographic Analysis. W. MikeHowell and Ann Black.An Isozymic Analysis of Several Southeastern Populations of the CyprinodontidFishes of the Fundulus notatus Species-Complex. Fred Tatum, Ronald Lindahl and Herbert Boschung. 35pp., illus., April, 1981. $5.007. Plant Resources, Archaeological Plant Remains, and Prehistoric Plant-Use Patterns in the CentralTombigbee River Valley. Gloria May Caddell. 39 pp., February, 1982. $5.008. Containing: Description, Biology and Distribution of the Spotfin Chub, Hybopsis monacha, a ThreatenedCyprinid Fish of the Tennessee River Drainage. Robert E. Jenkins and Noel M. Burkhead. Life Historyof the Banded Pygmy Sunfish, Elassoma zonatum Jordan (Pisces: Centrarchidae) in Western Kentucky.Stephen J. Walsh and Brooks M. Burr. 52 pp., illus., August, 1984. $6.009. Systematics of Notropis cahabae, a New Cyprinid Fish Endemic to the Cahaba River of the Mobile Basin.Richard L. Mayden and Bernard R. Kuhajda. 16 pp., illus., November, 1989. $3.5010. Containing: Notropis rafinesquei, a New Cyprinid Fish from the Yazoo River System in Mississippi. Royal D.Suttkus. Reproductive Behavior of Exoglossum species. Eugene G. Maurakis, William S. Woolcot, and MarkH. Sabaj. Scaphirhynchus suttkusi, a New Sturgeon from the Mobile Basin of Alabama and Mississippi. JamesD. Williams and Glenn H. Clemmer. 31 pp., illus., June 1991. $5.0011. Containing: A New Species of Hydropsyche (Trichoptera: Hydropsychidae) from Alabama, with AdditionalState Records for the Curvipalpia. Paul K. Lago and Steven C. Harris. New Caddisflies (Trichoptera) fromthe Little River Drainage in Northeastern Alabama. Kenneth Frazer and Steven C. Harris. New Caddisflies,(Trichoptera) from Alabama and Florida. Steven C. Harris. Survey of the Trichoptera in the Little RiverDrainage of Northeastern Alabama. Kenneth S. Frazer, Steven C. Harris and G. Milton Ward. 22 pp., illus.,September, 1991. $4.0012. Variation of the Spotted Sunfish, Lepomis punctatus Complex (Centrarchidae): Meristics, Morphometrics,Pigmentation and Species Limits. Melvin T. Warren Jr. 47 pp., illus. May 1992. $6.00

13. Containing: Effects of Impoundments on Freshwater Mussels (Mollusca: Bivalvia: Unionidae) in the MainChannel of the Black Warrior and Tombigbee Rivers in Western Alabama.James D. Williams, Samuel L. H.Fuller and Randall Grace. Etheostoma chermocki, a New Species of Darter (Teleostei: Percidae) from the BlackWarrior River Drainage of Alabama. Herbert T. Boschung, Richard L. Mayden, and Joseph R. Tomelleri.21pp., illus. September 1992. $5.0014. Catalog of Freshwater and Marine Fishes of Alabama. Herbert T. Boschung. 268 pp., December, 1992.$12.0015. Containing: Archaeological Survey and Excavations in the Coosa River Valley, Alabama. Vernon JamesKnight, Editor. Including: Archaeological Research in the Middle Coosa Valley. Vernon James Knight.Archaeological Research in the Logan Martin Basin. L. Ross Morrell. Lamar in the Middle Coosa RiverDrainage: The Ogletree Island Site. Richard Walling. The Milner Site: A Mid-Seventeenth Century SiteNear Gadsden, Alabama. Marvin T. Smith, Vernon J. Knight, Julie B. Smith, and Kenneth R. Turner.Seventeenth Century Aboriginal Settlement on the Coosa River. Marvin T. Smith. 87 pp., illus., January,1993. $10.0016. Containing: Elassoma alabamae, a New Species of Pygmy Sunfish Endemic to the Tennessee River Drainageof Alabama (Teleostei: Elassomatidae) . Richard L. Mayden. A New Species of Percina (Odontopholis) fromKentucky and Tennessee with Comparisons to Percina cymatotaenia (Teleostei: Percidae) . Brooks M. Burrand Lawrence M. Page. Systematics of the Etheostoma jordani Species Group (Teleostei: Percidae), withDescriptions of Three New Species. Robert M. Wood and Richard L. Mayden. 44 pp., illus., June, 1993.$10.0017. Containing: Etheostoma (Ulocentra) scottie (Osteichtheyes: Percidae), a New Darter from the Etowah RiverSystem in Georgia. Bruce H. Bauer, David A Etnier and Noel M. Burkhead. Present and Recent HistoricHabitat of the Alabama Sturgeon, Scaphirhynchus suttkusi Williams and Clemmer, in the Mobile Basin. JohnSelden Burke and John S. Ramsey. Roland Harper, Alabama Botanist and Social Critic: A BiographicalSketch and Bibliography. L. J. Davenport and G. Ward Hubbs. 45 pp., illus., May, 1995. $10.0018. pH and Temperature in Ectothermic Vertebrates. Gordon R. Ultsch and Donald C. Jackson. Life Historiesof Notuus baileyi and N. flavipinnis (Pisces: Ictaluridae), Two Rare Madtom Catfishes in Citico Creek, MonroeCounty, Tennessee. Gerald R. Dunkins and Peggy W. Shute. 69 pp., illus., December, 1996.$10.0019. The Mound Island Project: An Archaeological Survey in the Mobile-Tensaw Delta. Richard S. Fuller andIan W. Brown. 151 pp., illus., June, 1998. $10.0020. Containing: The Loss of Free-flowing Streams in the Gulf Coastal Plain. David Shankman. AllozymeVariation in the Longnose Shiner, Hybopsis logirostris (Teleostei, Cyprinidae). Tom Titus, E.O. Wiley, andMitchell Allen. A New Species of Cycleptus (Cypriniformes, Catostomidae) from the Gulf Slope Drainagesof Alabama, Mississippi, and Louisiana, with a Review of the Distribution, Biology, and Conservation Statusof the Genus. Brooks M. Burr and Richard L. Mayden. 57 pp., illus., August, 1999. $10.0021. Unionid Mollusks of the Apalachicola Basin in Alabama, Florida, and Georgia. Jayne Brim Box and JamesD. Williams. 143 pp., illus., April, 2000. $20.0022. Containing: Andrew C. Moore’s “Evolution Once More”: The Evolution-Creationism Controversy from anEarly 1920s Perspective. William D. Anderson, Jr. Systematics and Biogeorgraphy of the Notropis rubellusSpecies Group (Teleostei: Cyprinidae). Robert M. Wood, Richard L. Mayden, Ronald H. Matson, BernardR. Kuhajda, and Steven R. Layman. 80 pp., illus., November, 2002. $20.0023. Containing: Description of Larval and Juvenile Robust Redhorse, Moxostoma robustum. Gregory L. Looney,Cecil A. Jennings. Systematics, Variation, and Speciation of the Macrhybopsis aestivalis Complex West of theMississippi River. David J. Eisenhour. 48 pp., illus., December 2004. $10.00

24. A Historical and Current Perspective of the Freshwater Mussel Fauna (Bivalvia: Unionidae) from theChoctawhatchee River Drainage in Alabama and Florida. Holly N. Blaylock-Herod, Jeffrey J. Herod, JamesD. Williams, Briton N Wilson, and Stuart W. McGregor. 26pp., illus., July 2005. $10.0025. Containing: Systematics, Evolution and Biogeography of the Etheostoma simoterum Species Complex (Perdidae:Subgenus Ulocentra). Steven L. Powers and Richard L. Mayden. Distribution and Status of FreshwaterMussels (Bivalia: Unionidae) of the Lower Coosa and Tallapossa River Drainages in Alabama. Michael M.Gangloff and Jack W. Feminella. The Osteology of the Stone Cat, Noturus flavus (Siluriformes: Ictaluridae),with Comparisons to other Siluriforms. Jacob J. Egge. 89 pp., illus., August, 2007. $20.0026. An Archaeological Survey in Clarke County, Alabama. Ian. W. Brown. 129 pp., illus., January, 2009. $20.0027. Containing: Plain Post: A study of Late Woodland Pottery in Central Alabama. Jason Mann and RichardKrause. Discovery and Excavation of the Moundville Earth Lodge. Vernon James Knight. Analysis ofDaub from Mound V, Moundville: It Role as an Architectural Indicator. Jeffery Sherard. Analysis of WoodCharcoal from an Earth Lodge on Mound V at Moundville. Amanda Ticker. 49 pp., illus., November,2009. $12.0028. Paleontology Papers in Honor of Douglas E. Jones. Containing: Late Pleistocene Mammals of Alabama:A Comprehensive Faunal Review with 21 Previously Unreported Taxa. Jun A. Ebersole and Sandy M.Ebersole. A Review of the Chondrichthyans from the Mississippi System of Northern Alabama, USA.Chuck Ciampaglio, Leigh H . Deuter, Michael A. Taylor and David J. Cicimurri. A Review of the Non-AvianDinosaurs from the Late Cretaceous of Alabama, Mississippi, Georgia and Tennessee. Sandy M. Ebersoleand James L. King. 93 pp., illus., December, 2011. $18.0029. The Amphibians and Reptiles of Tuskegee National Forest, Macon County, Alabama. Sean P. Graham,David A. Steen, Roger D. Birkhead, and Craig Guyer. 59 pp., 4 tables, 108 illus. August 1, 2012. $14.0030. Morphological Diversity and Phylogenetics of the Darter Subgenus Doration (Percidae: Etheostoma), withDescriptions of Five New Species. Steven R. Layman and Richard L. Mayden. 83 pp., 16 tables, 27 illus.November 1, 2012 $16.00SPECIAL PUBLICATIONS1. Moundville, An Introduction to the Archaeology of a Mississippi Chiefdom, 2nd Edition. John Walthall. 47pp., illus., March, 1994. $3.502. Ten Thousand Years of Alabama History, A Pictorial Resume. W. Phillip Krebs. 130 pp., illus., January,1986. $10.003. The Mounds Awaken: Mound State Monument and the Civilian Conservation Corps. Joy Baklanoff andArthur Howington. 36 pp., illus. October, 1989. $3.00

MUSEUM PAPERS (1910-1960, TERMINATED)1. Smith Hall, The New Museum and Home of the Geological Survey. E.A. Smith. 7 pp., 1 plate.out of Print2. The Museum as an Educator. Herbert H. Smith. 25 pp., 8 plates, 1912. out of Print3. Directions for Collecting Land Shells. Herbert H. Smith. 12 pp., 1912. out of Print4. Annotated List of the Avery Bird Collection. Ernest G. Holt. 142 pp., 1 plate, 1921. $3.005. Preliminary Catalogue of Alabama Amphibians and Reptiles. H. P. Loding. 59 pp., 1922. Out of Print6. The Anculosae of the Alabama River Drainage. Calvin Goodrich. 57 pp., 3 plates, 1922 Out of Print7. The Genus Gyrotoma. Calvin Goodrich. 32 pp., 2 plates, 1924. out of Print8. The Terrestrial Shell-Bearing Mollusca of Alabama. Bryant Walker. 32 pp., illus., 1928. Out of Print9. Footprints from the Coal Measures of Alabama. T. H. Aldrich, Sr. and Walter B. Jones. 64 pp., illus., 1930.$3.0010. Goniobases of the Vicinity of Muscle Shoals. Calvin Goodrich. 25 pp., 1930. out of Print11. Alabama Reptiles. William L. Haltom. 145 pp., 39 plates, 57 figs., 1931. out of Print12. Description of a Few Alabama Eocene Species and Remarks on Varieties. T. H. Aldrich, Sr. 21 pp., 6 plates,1931. $3.0013. Moundville Culture and Burial Museum. Walter B. Jones and D. L. DeJarnette. 8 pp., 22 illus., 1936.out of Print14. The Argiopidae or Orb-Weaving Spiders of Alabama. Allan F. Archer. 77 pp., 5 plates, 1940. $3.0015. Anthropological Studies at Moundville. Part I. Indian Skeletons from the Museum Burials at Moundville.Part II. Possible Evidence of Scalping at Moundville. C. E. Snow. 57 pp., illus. 1940. $3.0016. Condylo-Diaphysial Angles of Indian Humeri from North Alabama. C. E. Snow. 38 pp., illus., 1940. $3.0017. The Bessemer Site (Excavation of Three Mounds and Surrounding Village Areas near Bessemer, Alabama).D.L. DeJarnette and S. B. Wimberly. 122 pp., illus., 1941. $3.0018. Supplement of the Argiopidae of Alabama. Allan F. Archer. 47 pp., 4 plates, 1941. $3.0019. McQuorquodale Mound. A Manifestation of the Hopewellian Phase in South Alabama. S. B. Wimberly andH.A. Tourtelot. 42 pp., illus., (1941) 1943. $3.0020. Mound State Monument. 19 pp., illus., 1941. out of Print21. Two Prehistoric Indian Dwarf Skeletons from Moundville. C. E. Snow. 90 pp., 2 plates, 1946. $3.0022. The Theridiidae or Comb-Footed Spiders from Moundville. Allan F. Archer. 67 pp., 2 plates, 1946. $3.0023. The Flint River Site, Ma°48. William S. Webb and D. L. DeJarnette. 44 pp., illus., 1948. Out of Print24. The Whitesburg Bridge Site, Ma V 10. William S. Webb and D. L. DeJarnette. 44 pp., illus., 1948.out of Print25. The Perry Site, LU°25. William S. Webb and D. L. DeJarnette. 69 pp., illus., 1948. $3.0026. Little Bear Creek Site, CT°8. William S. Webb and D. L. DeJarnette. 64 pp., illus., 1948. Out of Print27. New Anophthalmid Beetles (Fam. Carabidae) from the Appalachian Region. J. Manson Valentine. 19 pp.,2 plates, 1948. $3.0028. Land Snails of the Genus Stenotrema in the Alabama Region. Allan F. Archer. 85 pp., 10 plates, 1948. $3.0029. Moundville: An Historic Document. Carl E. Guthe. 14 pp., 1950. out of Print30. A Study of the Theridiid and Mimetid Spiders with Descriptions of New Genera and Species. Allan F.Archer. 44 pp., 4 plates, 1950. $3.0031. Carvernicolous Pselaphid Beetles of Alabama and Tennessee, with Observations on the Taxonomy of theFamily. Orlando Park. 107 pp., illus., 1951. $3.0032. Guntersville Basin Pottery. Marion D. Hemilich. 69 pp., illus. 1952. $3.0033. A Key to the Amphibians and Reptiles of Alabama. Ralph L. Chermock. 88 pp., illus., 1952. Out of Print34. New Genera of Anophthalmid Beetles from Cumberland Caves (Carabidae, Trechini). J. MansonValentine. 41 pp., 5 plates, 1952. $3.0035. New Genera and Species of Cavernicolous Diplopods from Alabama. Richard L. Hoffman. 13 pp., illus.,1956. $3.0036. Archaeological Investigations in Mobile County and Clarke County, Southern Alabama. Steve B. Wimberly.262 pp., 7 plates, 1960. $5.00

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