Copeia, 1997(1), pp. 116-122
New Species of Etheostoma(Teleostei: Percidae) from the Upper Tennessee River
JAY R. STAUFFERJR.AND ELLEN S. VAN SNIK
A new species of darter in the genus Etheostoma (Teleostei: Percidae) and sub- genus Nothonotus is described from the upper Tennessee River. The new species closely resembles Etheostoma tippecanoe and was historically regarded as a distinct population. The presence of 1-4 scales on the cheek, a light transverse bar directly anterior to the dorsal-fin origin, and fewer scales in the lateral series distinguishes the new species from E. tippecanoe.
THE subgenus Nothonotus (Bailey and Etnier, River and compare these populations to those 1988) is considered to be monophyletic of E. tippecanoe. (Page, 1981) and is characterized by the follow- slab-sided ing synapomorphies: body shape, MATERIALSAND METHODS darkened anterior membranes in the spinous portion of the dorsal fin, and horizontal lines Institutional abreviations are listed in Leviton on the lateral sides and (Bailey Etnier, 1988). et al. (1985). Etheostomatippecanoe from the up- The smallest member of the is Etheos- subgenus per Allegheny River (Allegheny River, PSU 1650 toma tippecanoe,which differs from the other No- [11]; French Creek, PSU 2520 [10]; PSU 1832 thonotus by consistently having an incomplete [9]; PSU 1652 [18]), middle Ohio River (Little lateral line (Bailey and Etnier, 1988). Etheostoma Kanawha River, PSU 3022 [10]; Elk River, PSU was described and Ever- tippecanoe by Jordan 1304 [10]; Big Darby Creek, OSU 13894 [10]), mann from the (1890) Tippecanoe River, lower Ohio River (Licking River, SIUC 9988 Indiana. In their review of the Marshland, type [10]; Redbird Creek, UMMZ 168882 [10]), Tip- of the Collette and specimens darters, Knapp pecanoe River (UMMZ 66560 [1]; INHS 68830 a (1966) designated lectotype (USNM 40080) [8]; INHS 63837 [4]; USNM 40080 [lectotype]; and and con- paralectotype (USNM 197919) USNM 197919 [paralectotype]), and the Cum- cluded that the of the larger remaining syntypes berland/Green Rivers (Big South Fork of Cum- was Etheostomacamurum and (UMMZ 187513) berland River, UT 91.661 [6]; Harpeth River, the smaller one was most probably E. camurum UT 91.1122 [4]; Green River, INHS 33515 [7]) as well. were compared to the new species from the Etheostoma is but with a tippecanoe widespread Tennessee River Drainage (Clinch River, UT distribution in the Cumber- very disjunct Ohio, 91814 [10]; UT 91.1676 [3]; Copper Creek, CU and Tennessee River land, drainages (Etnier 62822 [4]; CU 62923 [4] Sequatchie River, UT and The northeastern Starnes, 1993). limits to 91.2626 [9]). Standard length is used through- its range is the upper Allegheny River in north- out. Counts and measurements follow Hubbs western Pennsylvania, and the southeastern and Lagler (1964), except that the following dis- boundary is the Clinch River in Tennessee and tances also were recorded: anterior insertion of Virginia. Zorach (1969) recognized the distinc- dorsal fin to anterior insertion of anal fin, pos- tiveness of the Clinch River population andJen- terior insertion of second dorsal fin to posterior kins and Burkhead (1994) postulated that this insertion of anal fin, anterior insertion of sec- population may be a separate species. The ond dorsal fin to posterior insertion of anal fin, cheeks of E. tippecanoefrom the Ohio and Cum- posterior insertion of first dorsal fin to anterior berland River drainages are scaleless, whereas insertion of anal fin, posterior insertion of sec- individuals from the upper Tennessee River ond dorsal fin to ventral origin of caudal fin, have 1-4 scales on the cheek, immediately be- posterior insertion of anal fin to dorsal origin hind the eye (Zorach, 1969). Zorach (1969) fur- of caudal fin, anterior insertion of dorsal fin to ther noted that males from the Clinch River pelvic-fin origin, and posterior insertion of first have a light bar anterior to the first dorsal fin, dorsal fin to pelvic-fin origin (Stauffer, 1991). instead of the typical light saddle bordered an- These measurements were recorded so that a teriorly and posteriorly by dark saddles, which truss network could be completed, as described is found in the populations from the Ohio and by Humphries et al. (1981). All measurements Cumberland drainages. The purpose of this pa- were made point-to-point except caudal pedun- per is to describe the species from the Clinch cle length, for which the distance form a point
? 1997 by the American Society of Ichthyologists and Herpetologists STAUFFER AND VAN SNIK-NEW ETHEOSTOMA 117
Fig. 1. Holotype of Etheostomadenoncourti (CU 77101, 26.4 mm SL). on the lateral line directly above the posterior lateral series that have an opening, and postla- insertion of the anal fin to the hypural plate was teral scales are those scales posterior to the hy- measured. Lateral scales are all scales along the pural plate and may or may not be pored. All midline of the body from opercle to hypural counts and measurements were made on the plate, pored lateral scales are those scales in the left side of the fish.
TABLE 1. MORPHOMETRICAND MERISTICCHARACTERS OF Etheostomadenoncourti. Range and mean include holo- type and 29 paratypes.
Character Holotype Mean SD Range Standardlength, mm 26.4 24.4 2.1 20.0-28.6 Head length, mm 7.1 6.7 0.7 5.3-8.2 Percent of standardlength Head length 27.0 27.5 1.8 23-31 Body depth 23.4 23.9 1.5 20-28 Snout to first dorsal-finorigin 38.3 38.0 2.2 33-43 Snout to pelvic-finorigin 36.6 37.0 2.0 33-41 Dorsal-finbase length 59.3 57.6 2.8 51-64 Anteriorfirst dorsal to anterior anal 45.5 40.8 3.5 32-47 Anterior first dorsal to posterior anal 26.0 27.4 1.4 25-30 Posteriorsecond dorsal to anterior anal 26.3 29.8 2.1 25-35 Posteriorsecond dorsal to ventral caudal 14.9 15.9 1.2 13-18 Posteriorsecond dorsal to pelvic-finorigin 45.6 42.8 2.6 35-47 Posterioranal to dorsal caudal 19.8 17.5 1.9 14-23 Caudalpeduncle depth 14.3 13.0 1.1 10-15 Posteriorfirst dorsal to anterior anal 23.4 23.2 1.3 20-25 Pelvic-finorigin to posterior anal 16.6 15.5 1.7 12-19 Posteriorfirst dorsal to posterior anal 26.0 27.4 1.4 25-30 Caudalpeduncle length 10.7 10.5 2.3 6-16 Pelvic-finorigin to anterior anal 40.5 33.7 3.2 27-42 Anal-finbase length 14.4 21.5 2.9 14-27 Percent of head length Snout length 26.1 26.6 2.2 21-30 Postorbitalhead length 58.2 57.4 3.4 48-62 Horizontaleye diameter 20.8 21.3 2.8 17-28 Verticaleye diameter 15.4 17.6 2.7 13-23 Head depth 84.4 75.8 7.1 61-88 Counts Holotype Mode Freq. Range Dorsal-finspines 13 13 86.7 12-14 Dorsal-finrays 12 11 50.0 11-13 Anal-finrays 9 9 83.3 8-10 Pored lateral scales 30 23 16.7 19-34 Lateralscales 41 46 13.3 39-51 Postlateralscales 1 1 66.7 0-2 118 COPEIA, 1997, NO. 1
Principal components analysis (PCA) with the 20 correlation matrix factored was used to examine ? 18 A 'E 16 the meristic data. Sheared PCA with the covari- 14 ance matrix factored was used to analyze the morphometric data (Humphries et al., 1981; Bookstein et al., 1985). This technique quanti- 6- fies shape differences among the populations of fish size et al., 1984). 2 independent (Reyment 0 nn n Dissimilarities were illustrat- 1 8 i l,,,1, I, , , ,lnl, among populations -12 20 22 24 26 28 30 32 34 36 38 40 42 ed the sheared second by plotting principal Pored Lateral Scales component of the morphometric data against 4 the first principal component of the meristic 18 data and A multivariate (Stauffer Hert, 1992). 16 - B of variance (MANOVA) was used to de- 2 analysis 0 termine whether the clusters formed by popu- - 12- lations were significantly (P < 0.05) different. If these clusters were significantly different along one axis independent of the second axis, a Dun- can's Multiple Range test was used to determine which clusters differed. population O - -I11111111H ihll HI J Ill ni| NIn 39 41 43 45 47 49 51 53 55 Etheostomadenoncourti, n. sp. Lateral Scales Golden Darter ME. denoncourti l E. tippecanoe Figure 1 Fig. 2. Percent of specimens with pored (A) and Holotype.-CU 77101, adult male, 26.4 mm SL, lateral (B) scales in Etheostomadenoncourti and Etheos- Copper Creek, 180-460 m above mouth on tomatippecanoe. route 627, 2 air km. S. of Clinchport, VA, 24July 1967, R. F. Denoncourt and family. 14 in paratypes and 12 rays in holotype with Paratypes.-PSU 3023, 3 (22.6-27.4 mm) data as 11-13 in paratypes; anal fin with 12 rays in ho- for holotype, CU 62923, 4 (24.9-25.9 mm) lo- lotype and 8-10 in paratypes. The holotype cality as for holotype, 9 August 1967, R. F. De- with 41 lateral scales, 30 pored; paratypes with noncourt, D. C. Wallace; UT 91.814, 10 (20.0- 39-51 lateral scales, 19-34 pored. The holo- 28.6 mm), Clinch River at Frostford, TN, 11.8 type with 1 postlateral scale, and paratypes 0- air km. WSW of Kyles Ford, TN, 19 September 2. 1973, D. M. Etnier and UT ichthyology class; UT Etheostoma denoncourti typically has fewer 91.1676, 3 (24.3-25.8 mm), Clinch River at scales in the lateral series than E. tippecanoe Frostford, TN, 11.8 air km. WSW of Kyles Ford, (Fig. 2, Table 2). The mode for lateral scales is TN, 24 October 1978, D. M. Etnier and UT ich- 46 and for pored lateral scales is 23 in E. de- thyology class; UT 91.2626, 9 (21.0-25.7 mm), noncourti, whereas the modes for lateral scales Sequatchie River, km. 18.5, TN, 8 July 1983, D. range between 47 and 50 and for pored lateral Biggins, C. F. Saylor, and R. Hylton. scales between 27 and 34 for E. tippecanoe(Fig. 2). More than 86% of E. denoncourti have 13 Diagnosis.-A darter of the subgenus Nothonotus dorsal-fin spines, whereas the mode in E. tip- (genus Etheostoma)that closely resembles E. tip- pecanoe is 12. The mode for dorsal-fin rays is 11 pecanoe. The slab-sided body shape, darkened for E. denoncourtiand 12 for E. tippecanoe(Table anterior membranes in the spinous portion of 2). the dorsal fin, and horizontal lines on the lat- The first principal component of the mor- eral sides (Bailey and Etnier, 1988) clearly place phometric data (69.2% of total variance) is in- this species in the subgenus Nothonotus. The terpreted as a size component and the sheared presence of scales on the cheek and a light line components as shape, independent of size directly anterior to the dorsal-fin origin distin- (Humphries et al., 1981; Bookstein et al., guishes it from E. tippecanoe. 1985). Those variables that have the highest loadings on the sheared second principal com- Description.-Morphometric ratios and meris- ponent (8.4% of total variance.) are caudal pe- tics are presented in Table 1. Jaws isognathous. duncle length, posterior anal-fin to dorsal cau- Dorsal fin with 13 spines in holotype and 12- dal-fin insertion, and anal-fin base length (Ta- TABLE2. MORPHOMETRICAND MERISTIC CHARACTERS OF Etheostomatippecanoe POPULATIONS.
Drainage = = = = Upper Allegheny, n = 50 Middle Ohio, n 30 Cumberland & Green, n 18 Lower Ohio, n 20 Tippecanoe River, n 15 Character Mean SD Range Mean SD Range Mean SD Range Mean SD Range Mean SD Range Standard length, mm 28.9 2.8 24.6-35.3 25.8 2.4 22.0-31.3 26.8 3.9 22.0-34.0 22.6 1.7 19.7-26.3 29.2 3.5 23.7-36.0 Head length, mm 7.8 0.8 6.2-9.3 7.1 0.7 5.7-8.6 7.4 1.3 5.6-10.0 6.4 0.5 5.4-7.5 7.2 1.2 5.67-9.55 Percent of standard length Head 26.9 1.5 24-30 27.4 1.7 24-30 27.4 2.0 24-30 28.3 1.3 26-30 24.6 1.7 22-27 length c-3 Body depth 25.3 1.4 22-29 24.8 1.5 22-29 24.8 2.7 21-29 24.0 1.8 21-29 22.3 2.2 17-25 Snout to first dorsal-fin origin 37.8 1.5 34-41 39.0 2.3 34-44 36.7 2.1 31-39 37.4 2.8 28-41 34.5 2.4 29-37 cn Snout to pelvic-fin origin 34.7 1.6 32-38 35.9 2.0 32-43 35.6 1.9. 33-38 38.2 1.7 35-41 33.7 3.1 31-41 Dorsal-fin base length 55.8 2.4 51-60 55.0 2.5 50-60 56.7 1.6 54-60 56.0 2.6 51-64 57.3 3.4 50-62 Anterior first dorsal to anterior anal 42.0 1.7 39-47 40.3 2.9 34-47 41.6 2.5 38-47 41.4 3.0 37-47 41.7 2.1 38-46 tI Anterior first dorsal to posterior anal 27.8 2.4 22-38 27.5 1.9 23-31 27.9 1.9 25-31 27.3 2.1 25-33 27.4 1.5 24-29 Posterior second dorsal to anterior anal 27.7 2.1 23-32 27.8 2.1 24-31 28.5 2.2 24-32 27.6 1.8 24-32 27.9 2.2 22-31 Posterior second dorsal to ventral caudal 16.2 2.0 11-21 16.5 0.7 16-18 16.4 1.8 12-19 15.9 1.5 14-19 16.5 2.3 14-22 Posterior second dorsal to pelvic-fin origin 43.3 2.3 39-49 42.3 2.2 37-47 42.6 2.3 39-47 41.6 2.6 37-48 41.1 2.9 33-45 Posterior anal to dorsal caudal 17.7 2.0 13-21 17.5 1.3 14-20 17.0 2.0 13-19 17.7 1.8 15-22 18.8 2.4 15-25 11.9 1.2 9-14 12.7 0.9 11-14 12.8 1.3 9-14 12.9 1.2 11-15 11.8 1.4 8-13 z Caudal peduncle depth 0 Posterior first dorsal to anterior anal 23.1 1.7 20-27 22.7 1.4 19-26 22.7 2.7 18-27 22.3 1.6 20-25 21.7 1.4 20-24 14.3 Pelvic-fin origin to posterior anal 14.8 1.3 12-18 15.0 1.0 12-17 14.4 1.9 12-17 15.0 1.5 13-18 1.2 12-16 t TABLE3. VARIABLELOADINGS ON SIZEAND THE FIRST TWO SHEARED PRINCIPAL COMPONENTS (MORPHOMETRICS) FOREtheostoma denoncourti AND E. tippecanoe. Characters Size Sheared PC2 Sheared PC3 Standardlength -0.176 -0.020 0.041 Head length -0.199 0.099 0.022 Snout length -0.200 0.187 -0.151 Postorbitalhead length -0.231 0.096 -0.114 Horizontaleye diameter -0.114 -0.007 0.597 Verticaleye diameter -0.119 -0.027 0.682 Head depth -0.239 0.101 0.035 Body depth -0.228 0.077 0.008 Snout to first dorsal-finorigin -0.172 0.050 -0.017 Snout to pelvic-finorigin -0.167 0.059 0.010 Dorsal-finbase length -0.178 0.003 0.047 Anteriorfirst dorsal to anterior anal -0.186 0.021 0.105 Anteriorfirst dorsal to posterior anal -0.199 0.140 -0.023 Posteriorsecond dorsal to anterior anal -0.232 0.010 -0.103 Posteriorsecond dorsal to ventral caudal -0.224 -0.052 -0.037 Posteriorsecond dorsal to pelvic-finorigin -0.198 0.045 0.061 Posterioranal to dorsal caudal -0.230 -0.320 -0.153 Caudalpeduncle depth -0.230 0.140 -0.101 Posteriorfirst dorsal to anterior anal -0.257 0.066 -0.075 Posteriorsecond dorsal to posterior anal -0.211 -0.029 -0.209 Posteriorfirst dorsal to posterior anal -0.199 0.140 -0.023 Caudalpeduncle length -0.263 -0.829 -0.026 Pelvic-finorigin to posterioranal -0.154 0.036 0.157 Anal-finbase length -0.210 0.244 -0.033 ble 3). When these variables were each re- or to the dorsal-fin origin. Head is brown dor- gressed against standard length, there was no sally, with yellow cheek and orange lower jaw. indication that any of the variables were im- Three black bars, more diffuse than lateral pacted by allometric growth (Fig. 3). bars on the body, surround eye. The subocu- A plot of the first principal component lar bar in some individuals is obscured by yel- scores of the meristic data versus the second low coloration. Distinct iridescent diagonal sheared principal component scores of the slash on cheek. When the head is viewed from morphometric data showed only slight overlap the front the cheeks appear faint green. Dor- between E. denoncourti and E. tippecanoe (Fig. sal fin is gray/orange with orange marginal 4). A MANOVA indicated that the clusters bar; the rayed portion of the dorsal fin has formed by E. denoncourtiand E. tippecanoewere dark brown blotches proximally. The caudal, significantly different. Duncan's multiple anal, and paired fins are faint orange. There range test showed that the E. denoncourti clus- are gray/black melanophores present in the ter was significantly different from the clusters caudal and pelvic fins. Pigmentation patterns formed by each of the populations of E. tippe- of females and nonbreeding males are similar canoe along the first principal component of to breeding males, but head and body are the meristic data, which is loaded highly on brown rather than bright orange. pored lateral scales, lateral scales, and dorsal- fin spines (Table 4). Etymology.-The specific epithet is in honor of Lateral body coloration of breeding males Robert F. Denoncourt, who first introduced JRS is orange with 7-9 black bars; bars on nape to the diversity and complexity of the stream and caudal peduncle are more intense. Or- fishes in the southern Appalachian drainages. ange spotting occurs between the pectoral We suggest golden darter as the common name. and pelvic fins. Breast gray/blue. Two blotch- es (yellow in breeding males; white in fe- Distribution.-Etheostoma denoncourti is endemic males) on the dorsal and ventral posterior to the upper Tennessee River drainage (Etnier portion of the caudal peduncle. There is a dis- and Starnes, 1993:542). Historically, it was col- tinct transverse light line dorsally, just anteri- lected in the Duck. Buffalo. Clinch, and Se- STAUFFER AND VAN SNIK-NEW ETHEOSTOMA 121 TABLE 4. STANDARDIZEDSCORING COEFFICIENTSON 0.25 MERISTIC DATA FOR Etheostomadenoncourti AND E. tip- pecanoe. - 0.2 C 01 Characters PC1 PC2 P 0.15 Dorsal-finspines -0.330 -0.013 0.1 Dorsal-finrays 0.155 0.484 15 20 25 30 335 40 SL Anal-finrays -0.141 0.688 [ E. tippecanoe * E. denoncourti Pored lateral scales 0.431 -0.018 = 0.25 CPL 0.04007 + 0.0029 SL; R2= 0.1626 Lateralscales 0.424 0.018 D Postlateralscales 0.042 0.439 0.21 B ,- 0.17 0.13 - cP 0.09 - - Qrb"nJ0 .( E nes, 1993). When PCA of the meristic data and sheared PCA of the data are cal- l51.5 220 25 SLS 303 353 40 morphometric culated and the resultant components plotted 0.3 AFBL=0.1944 + 0.0003 SL; R2-0.0039 (Fig. 5), differences between the Clinch River *??~~ c population and the Copper Creek (a tributary 0.25 are -- t iCSbRg* C of the Clinch River) population apparent, 0.-2 g with specimens from the Sequatchie River being q~o - < 0.15- ar intermediate. 0.1 15 20 25 30 35 40 Life history.-Etheostoma denoncourti is a relatively SL late spawner, with ripe females being collected as late as 4 August from the Clinch River (Jen- Fig. 3. Plot of ratio of posterior of anal fin to dor- s 1 994 ) Nm er kins andand Burkhead, Numerous observa-ea- sal portion of caudal fin (PADC) to standardlength 1994). (A), caudal peduncle length (CPL) to standard tions have demonstrated that E. tippecanoefe- length (B), and anal-finbase length (AFBL)to stan- males are egg buriers (Mount, 1959; Trautman, dard length (C), against standardlength for Etheosto- 1981; Page and Simon, 1988). It is postulated ma denoncourti(closed circles) and Etheostomatippecan- that E. denoncourti is also an egg burier as are oe (open squares).Regression equations and adjusted other Nothonotus species including E. camurum, R2values are at the top of each graph. E. rufilineatum, and E. juliae (Warren et al., 1986). Etheostoma denoncourti feeds primarily on small aquatic insects, including mayfly, caddisfly, quatchie River drainages (Jenkins and Burk- and midge larvae (Jenkins and Burkhead, head, 1994), although portions of the Duck Riv- 1q4 er population immediately below Normandy Reservoir may be extirpated (Etnier and Star- 3. 2 2.25- .. i I - 0.75 UpperAlleghe?? E O S 0 - X Middle Ohio -1 u Cumberland/Gre P -0,750 -_ * ClinchR. A LowerOhio o SequatchieR. -1.5 - -2- * CopperCreek * TippecanoeR. 0 0 -2.25- I I I I I I 0 E. denoncourti -0.3 -02 -0,1 0 0.1 0,2 0.3 3 , I - I --- o 0.2-- ShearedPC2 (Morphometrics) -0.5 -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 ShearedPC2 (Morphometrics) Fig. 5. Plot of individualfirst principal component Fig. 4. Plot of individualfirst principal component scores (meristics) and sheared second principalcom- scores (meristics)and sheared second principalcom- ponent scores (morphometrics) of Etheostomadenon- ponent scores (morphometrics) of Etheostomadenon- courtifrom the Clinch River (closed squares), Se- courti and populations of Etheostomatippecanoe across quatchie River (open squares), and Copper Creek its range. (closed circles), VA and TN. 122 COPEIA, 1997, NO. 1 ACKNOWLEDGMENTS DAWSON.1985. Standardsin herpetology and ich- thyology:Part I. Standardsymbolic codes for insti- The line drawing was remarked from a pho- tutional resource collections in herpetology and tograph by M. Katz. Fishes were collected under ichthyology.Copeia 1985:802-832. PA 018, and specimens were provided by Cor- MOUNT,D. I. 1959. Spawningbehavior of the blue- nell University, Illinois Natural History Survey, breast darter, Etheostoma camurum (Cope). Ibid. Ohio State University, Southern Illinois Univer- 1959:240-243. L. M. 1981. The and of dart- sity at Carbondale, University of Michigan Mu- PAGE, genera subgenera seum of and the of Tennes- ers (Percidae,Etheostomatini). Occ. Pap. Mus.Nat. Zoology, University Hist. Kansas90:1-69. see. The work was funded in Univ. part by Pennsyl- T. P. SIMON. 1988. Observations on the vania's Wild Resource Conservation Fund. ,AND reproductivebehavior and eggs of four species of darters, with comments on Etheostomatippecanoe and LITERATURE CITED E. camurum.Trans. Ill. Acad. Sci. 81:205-210. REYMENT,R. A., R. E. BLACKITH,AND N. A. CAMPELL. BAILEY,R. M., ANDD. A. ETNIER.1988. Comments on 1984. Multivariatemorphometrics. Academic Press, the of darters with subgenera (Percidae) descrip- Newy York. tions of two new of Etheostoma species (Ulocentra) STAUFFER,J. R., JR. 1991. Description of a faculative from southeastern United States. Misc. Pub. Mus. cleanerfish (Teleostei:Cichlidae) from Lake Mala- Zool. Univ. Mich. 175:1-48. wi, Africa.Copeia 1991:141-147. BOOKSTEIN,F., B. CHERNOFF,R. ELDER, HUMPHRIES, J. , ANDE. HERT. 1992. Pseudotropheuscallainos, a G. SMITH,AND R. STRAUSS.1985. Morphometrics in new of mbuna with of of species (Cichlidae), analyses evolutionarybiology. Academy NaturalSciences associated with two intra-lacustrinetrans- of Publ. 15. PA. changes Philadelphia.Spec. Philadelphia, in Lake Africa. B. ANDL. W. KNAPP.1966. of plantations Malawi, Ichthyol.Explor. COLLETTE, B., Catalog Freshwaters3:253-264. type specimens of the darters (Pisces, Percidae). Tulane Stud. Zool. 9:1-88. 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RESOURCES, STATE UNIVERSITY,UNIVERSITY PARK, PENNSYL- JORDAN,D. S., AND B. W. EVERMANN.1890. Descrip- 16802. Submitted: Feb. 1996. tion of a new species of fish from TippecanoeRiver, VANIA 2 Accept- Indiana. Proc. U.S. Nat. Mus. 13:3-4. ed: 13 May 1996. Section editor: R. Winter- LEVITON,A. E., R. H. GIBBSJR., E. HEAL,AND C. E. bottom.