r
DESCRIPTIONS OF LARVAL PERCIDAE INHABITING THE UPPER MISSISSIPPI RIVER BASIN (OSTEICHTIIYES: ETHEOSTOMATINI)
A Thesis
Submitted to the Faculty
of
University. of Wisconsin·· ;.,; La Crosse LaCrosse,Wisconsin 54601
by
Thomas P. Simon
In Partial Fulfillment of the Requirements for the Degree of Master of Science in Biology
May 1~85 • v\)-\ . \ () () UNIVERSITY OF WISCONSIN ... LA CROSSE
La Crosse, Wisconsin 54601 COLLEGE OF ARTS, LETTERS, AND SCIENCES
Candidate: Thomas P. Simon
We recommend acceptance of this thesis to the College of Arts,. Letters, and Science in partial fulfillment of this candidate's requirements for thedegre~.}fElli3t~:r. ()~ .. SC!E!n.ce .in Biology. The candidate .has completed his oral defense of the thesis.
Thesis approved: ~p2?;/J75~· Date
qq?C~Tq8? D e
9-:f--~zc4f:s ~
L ~e~
f2~' ~ ?t:~J'J Thesis Committee Member /Dat
Ii$<' Dea~: ~~A~~ers. ~:±tJ:jl!!~.C snd-Seienees . . _ n ••••
~ulIJ\J ~ .its- rv~ 3 86,-00531 l1li ABSTRACT Eggs, larvae, and juveniles of six species of darters inhabiting the Upper Mississippi River are described. Attention to meristic, morphometric, pigment, and morphological apparent were employed for identification. Species of the genus Etheostoma possess well.developed pectoral fins, D1~:xillary, mandible,fewerthari 18 preanal myome:res, and have greater body depth tharacteristics/TL than does the subgenus Percina. The subgenus Catonotus (E. kennicotti, E.flabellare lineolatum, arid E. sguamiceps) has large, robust, spherical yolk sacs when compared to eithe~.. the Microperca, Percina, or Imostoma subgenera. The Microperca subgenus (E. microperca) has 15 preanal and 19 postanal myomeres and hatches at smaller lengths than other Etheostoma. The genusPercina is characterized by slender bodies, smaller body depth characteristics/TL than Etheostoma, undeveloped, maxillary and mandible at hatching, preanal myomeres equal to or greater than 18, and weakly developed-pectorals. caprodes semifasciata possesses fewer postanal myomeres than P. shumardi, while the two species each have differential pigmentation patterns especially apparent in the postanal hypaxial musculature• .... ii ,. 'Ii ACKNOWLEDGMENTS I am forever indebted for the support, affection, understari.diri.g, and encouragement of my wife, Beth Simon,and·my·family especially my mother and father, whose gentle prodding and guidance helped me to complete this project. I am grateful to Karl F. Lagler, David Jude , 'JohriDorrIII;LeecFuimafi and Paul Simon Sr. for stimulating my interest and providing the foundation of my understanding in the discipline of ichthyology •. a.ncl larval fish biology. To Rodney. Mowbray and Leslie Holland, my major professors, John Held, and Thomas Claflin for the cultivation of .that knowledge, and educational stimulation into other areas offisheries and aquatic biology. I sincerely "appreciate the help and encouragement of Larry M.Page, Illinois Natural Hist()J:'y.SuJ:'yey.for his prQvision of specimeiisana ifisight and wisdom on the phylogenetic relationships among the tribe Etheostomatini. I appreciate the aid of the following individuals during field collections and for their contributionsJ;Qcc.I1lY~_l'~cl:1.E:!~~h. Field assistance was PI'~Y!~~.~. by Bill Kowalski, Nancy Garcia, Tom Coon, Barry Burnell, Mike Troll; Mark Huston, Tim Babros, and Rick Jacobson. Additional gratitude is due to Bill Kowalski for enduring . many conversations and problem solving sessions; Dan Faber, National Museums of Canada for advise on description format; and Darrel E~Sfiyder,LarvalFish Laboratory, Colorado State University, for stimulating suggestions and 41' additional research ideas. Special thanks to Greg Seegert, EA Science and Technology, Richard ,,,. -"~-~-'~-"'----- " -,._. -::., ~~~...._.."". Speiler, Milwaukee Public Museum, Robert R. Miller, University of Michigan Museum of Zoology, and Lynn Parenti, Field Museum.ofNaturalHistory for providing laboratory space and equipment. Drawings of figures were completed by Beth Simon. iii ~ TABLE OF CONTENTS PAGE LIST OF TABLES • ...... LIST OF FIGURES...... vii INTRODUCTION ••• ...... 1 MATERIALS AND METHODS ...... SPECIES ACCOUNTS Northern Logperch, Percina caprodes.... semifasciata (Rafinesque). 9 River darter, Percina shumardi (Girard) .• ••• ••• • •• . .25 Striped fantail darter, Etheostoma flabellare lineolatum • •• 39 Stripetail darter, Etheostoma kennicotti (Putnam)··. •• . . 54 ....? Least darter, Etheostoma microperca Jordan and Gilbert 67 Spottail darter, Etheostomasguamiceps··Jordan DISCUSSION • . . .. 93 CONCLUSIONS: ...... 105 LITERATURE CITED • i ••• 141> iv r- LIST OF TABLES .TABLE .lli]. 1. Material examined of Percina caprodes.semifasciata larvae and early juveniles grouped by Imm interval of total length (N = sample size) ••••••••••••••••• •• 11 2. Morphometry of Percina caprodes semifasciata larvae and early juveniles grouped by 1mm intervEJ,J.softot,EJ,J.length (N = sample size). Mean lengths followed by range in parentheses ...... 15 3. Selected meristic valuesand.size (mmtotal length) at theflPPE!:r~Ilt.onf3..et of development. for Percina caprodes semifasciata. Mean values are underscored. Rare or ques tionable extremes are in parentheses. The number of secondary rays.. of thellledian fins are in lowercase ROIIJ.a,D ,numerals ... . . '.-...... 17 4. Material examined of Percina shumardi larvae and early juveniles grouped by 1mm intervals of total length (N = sample size) ••••• .' •. .• .• •••• ...... 27 5. Morphometry of Percina shumardi larvae and early juveniles grouped by 1mm intervals of total length (N = sample size). Mean lengths followed by range in parentheses •• •• 29 6. Selected meristic values and size (mmtotal length) at the apparent onset of development for Percina shumardi. Mean values are underscored. Rare or questionableextremes--are in parentheses. The number. of secondary rays of the median finsare1nlowerJ;afieRQma.n numerals. ••• ••••• •• •• 31 7. Material examined of Etheostoma flabellare lineolatum larvae and early juveniJ.es grouped by 1mm intervals of total length (N = sample size) •••••• •• •• •.• •• •• 41 8. Morphometry of Etheostoma flabellare lineolatum larvae and early. juveniles grouped by Imm intervals of tot.a.lTengffi (N = sample size). _Mean lengths followed by range in parentheses ...... , ...... 44 $> 9. Selected meristic values and size (mm total length) at the apparent onset of development for Etheostoma flabellare lineolatum. Mean values are underscored. Rare Or ques tionable extremes are in parenth~§es. The number of secondary rays of the median fins are·· iii-lowercase Roman numerals •••••••••••••••••••• 46 10. Material examined of Etheostoma kennicotti larvae and early juveniles grouped by 1mm intervals of total length (N = sample size) •••••••••••••••••••• •• 55 v TABLE PAGE 11. Morphometry of Etheostoma kennicottilarvae and early juveniles grouped by 1mm intervals of total length (N = sample size). Mean l.engths followed by range in parentheses •.•..•••••••.•..•• ••• •• •. 58 12. Sf:!l.f:!(;'t:f:!d)lleristicvalues and size (lllJll't:o't:a.1.l.eng't:h)a't:. the apparent onset of development for Etheostoma kennicotti. Mean values are underscored. Rare or questionable extremes are in. parentheses. The numberofsecoridaryraysc>f the median fins are in lowercase Roman numerals. ••••• • • •• 60 13. Ml:l't:f:!l:"ta.l.. f:!JCa.mined of Etheostoma· •• microperca •. larvae ..and early juveniles grouped by 1mm intervals of total length (N =·sample si~e) ••.•••• •••••• • •••• . . 69 14. Morphometry of Etheostoma microperca larvae and early juveniles grouped by 1mm intervals of total length (N = sample size). Mean lengths followed by range in paren:th~§~~tt:",.~.. .••••••••. • •• .••• •• .. 71 15. Selected metistic values and size (mm total length) at the apparent onset of development for Etheostomamicroperca. Me.an values are underscored. Rareor •... questionable~cextremes are in parentheses. The number of secondary rays of the median fins are in lowercase Roman numerals ••••••• •• 73 16. Material examined of Etheostoma sQuamiceps larvae and early juveniles grouped by 1mm intervals of . total length (N='samplesize) 81 17 •. Morphometry of Etheostoma sguamiceps larvae and early j\1Venilesgrouped,by 1 mm intervals of tQtal length (N = sample size) ...... 84 18. Selected meristic values and size (mm total length) at the apparent onset of development for Etheostomasguamiceps. Mean values are underscored. Rare or questionable f:!xtremes are in parentheses. The number of secondary raYcs_~QftlJ.e median fins are in lowercase Roman numerals ••••.• •• •• 86 19. Egg characteristics of Upper Mississippi River percids • . .. 94 tdi' 20. Total length (mm) at hatching, yolk absorption, and first apearance of fin ray formation in Upper Mississippi River percids ...... 96 vi ,.. I" I LIST OF FIGURES jl!, FIGURE 1. Diagrammatic representation of morphometric character- istics ...... 7 2. Percina •• caprodes semifasciata, northern logperch (newly hatched larva) 5.4 mm TL, Upper Mississippi River, Pool 3. Minnesota a. dorsal b. lateral c • ventral.... •• . .. . 19 3. Percina caprodes semifasciata, northernlogperch larva, Upper Mississippi River, Pool 7 a. 7.5 mm TL b. 9.2 mm -u,----- .---,----.------.------.----.------.--,-.--.-. -e----- ••---.------.--:. • • •• -e. e--a.-__ •• ••• __-.-e ••• _.. 20 4. Pertina· caprodes· ·semifasciata, northern logperch, a. 14.6 mm TL Catgut Slough, Blac:lcRiver, Wisconsin b. 26.5 mm TL Upper Mississippi River, Pool 3 • • • • • • •• 22 5. Percina shumardi, river darter (newly hatched larva), 4.0mmTLBlackRiyer, Wisconsin a.d.orsalb. l~t~J:al c. ventral :_ . 32 6. Percina shumardi, river darter larva, Upper Mississippi River, PoolS, Wisconsin a. 6.2 mDI TL b~8~OmmTL~~ 34 7. Percina shumardi, . river darter, a. 12.3 mm TL larva Upper ·Mississippi River, PoolS, Wisconsin b. 24.1 mm TL early juvenile ••••••.•••••••••••• 36 8. Etheostoma flabellare lineolatum, .Elt:rj,pElcl ..... fantail darter (newly hatched larva), 4.8 mm TL, CoonCreek~ Wisconsin a. dorsal b. lateral c. ventral. ••••• 48 9. Etheostoma flabellare lineolatum, striped fantail darter larva Spring Coulee Creek, Wisconsin a. 7.1 mm TL. b •. 8.2 mm TL •. ••••••••••••• ••• -. 49 10. Etheostoma flabellare lineolatum, striped fantail darter, Root River, Minnesota a. 10.3 mmTLlsrva b. 14.4 mm TL early juvenile ...... 51 11. Etheostoma kennicotti, stripetail darter (newly hatched mID 4!1> larva) 4.3 TL, Big Creek, Illinois a. dorsal b. lateral c. ventral •••• ••••••.••••• 61 12. Etheostoma kennicotti, stripetail darter larva, Big Creek, Illinois . a. ~6. 9 J:II1Il TL b. 7~5clfun~".['L= ...... 63 13. Etheostoma kennicotti, stripetail darter, Big Creek, Illinois a. 12.1 mm TL larva b. 21.3 mm TL ~J:ly juvenile ...... 64 vii pi ~,f ~J FIGURE PAGE 14. Etheostoma microperca, least darter (newly hatched larva), 3.0 mm TL, Piscasaw River, Illinois a. dorsal b. lateral c::.. ventral ••••••••••••••• •• • 75 15. Etheostoma microperca, least darter larva, a. 4.0mm Tt, unnamed tributary Iroquois River, Illinois b. 6.4 rom Tt Piscasaw River, Illinois ••••••••••••••• 76 16. Etheostoma· microperca, least darter., PisC:asa.wiRiver, a. 8.2 mm TL larva b. 14.5 mm TL early juvenile •••••• 78 17. Etheosto s911llDliceRs,· spottail darter (newly hatched larva), ma5.7 mmTL, ..... Big Creek, Illinois a.. dorsal b. lateral· c. ventral •••••••••••••••••• 88 18. Etheostoma sguamiceps, spottail darter larva, Big Creek Illinois. a.• 6.8 rom TL b. 8.6 Jmn TL...... • •..• •.•.• 89 19. Etheostoma sguamiceps, spottail darter, Big Creek Illinois a. 13.6.DlDl TL larva b~ 16.7lJlJ1lTLearly juvenile .'.". •••••••••• . . . 91 $'1:; viii r if INTRODUCTION Technological advances and pollution have imposed an increased demand on the Mississippi River as a source of water and recreation (Schnick et al. 1982). The entrainment of surface waters by "once-through" cooling systems of large industries is respons;t.bl¢forDl¢<:.ht:lIl,:i,<:'t:l±'~~~~!D(i~,and chemical mortality of larval fishes. Increased barge· and recreational boat traffiC on the river ·has increased sedimentation in backwater areas impacting nursery habitats. The construct;t.on of locks, dams, bridges and ensuing dredging have contributed to the destruction. of . microhabitats for fish species. Increased concern over habitat alterations has caused an awareness of envttonmental perturbations and the development of a strict monitoring program. In order to accurately ~ss~ss various environmental disturbances, the life histories of various trophic levels need to be described. :Among these species include the darters. The darters are generally found· in areas of excellent water quality and are subtle indicators of ,decreasing standards (Tsai 1972; Kuehne and Barbour 1983). The importance of the darters taxonomic description is reflected in the ability to assess environmental impacts at specific levels rather than the present familiar levels. Description of meristic, morphometric, and pigment characteristics will facilitateiden£:i.f:i.cation of egg, larval, and juvenile stages of species. Of equal importance in &I'> fishery research is the ability to differentiate between larval fishes in order to assess growth, age, and the demecology for all stages of a species' life cycle. TQ.e larval stage; of .... fishes· have··· ·generalTj been ignored in this type of analysis primarily because of the Type III survival pattern during the early life history (Wilson and Bossert 1971). Many researchers believe that the magnitude of the year class is established not r 2 during the juvenile or adult periods, but during larval development (May 1974; Bannister et al. 1974; Jones and Hall 1974; Cushing 1974; Postuma and Zijlstra 1974; Tanaka 1974; Carlander and Payne 1977; Ielllpmger sfidCarline 1977; Cada and Hergenrader 1980). This indicates that the larval stage of obvious importance in determining recruitment and survival to maturity. The early life history of the neglected ~Il ichthyological literature. Limited comparative information on development of Percidae is available (Fish 1932;.Rohde 1974; Hardy 1978; Auer 1982; Simon 1983). The study of fish· ontogeny may elucidate the phylogenetic relationships important to ichthyology and zoology and determine relationships among subgenera (BaileysfidGos1±ne 1955; Page 1974; Page 1981; Page 1983; Kuehne and Barbour 1983). The family Percidae contains the second largest species assemblage of North American freshwater fishes, rivaled only by the Cyprinidae. The family Percidae consists of 2 sub-families: the Luciopercinae and Percinae (Collette and Banarescu 1977). The darterscomprise--98--%~-()f--the127 described members of the percids with 123 described species and approximately 22 undescribed forms (Page 1983; Kuehne and~arbour 1983; Simon 1983). The present study identifi.es and describes larva~aIld.~a:I"J.yjll'Veniles of the Etheostomatini, the da.rters. The darters are restricted to North America east of the Rocky Mountains with the center of distribution being 4i> in the southeastern United States (Scott and Crossman 1973; Page 1983). The darters inhabit a variety .... of habitatScc~c,fr_QJI1__warJ:ll. __ 't~_mp~~at~~_ tQ cold subarctic lentic and lotic environments. The Upper Mississippi River (UMR) extends from the confluence of the Ohio River and Mississippi River near Cairo, Illinois to St. Anthony Falls r';: 3 near Minneapolis, Minnesota. The Upper Mississippi River consists of a series of 26 navigation pools separated by numbered locks and dams. Species seletted f()r the present study areamongth()se that inhabit the Upper MississippiR:f..ver basin and i tstrib.utaries ..••.... wb,ieb, d.:ra:f.Il.l3 lJll3jor portion of WiStOIlsiIl, Minnesota, Illinois, Iowa, and Missouri. The percid fauna of the Upper Mississippi.River iscoDlpI'1sedof21.. sp~el~s.ltL=~:tg~J1~!t;l (Forbes and Richardson 1920; Greene 1935;Ha.rlanandSpe~er 1951; Eddy and Underhill··1974; Pflieger 1975; •. Smith 1979; Phillipset>al. 1982;aIld •• 13eclcer 1983). The species walleye, Stizostedion .vitreum,yellow perch, Perca flavescens, and saugeJ;" , S. canadense although . much more common were previously described (Fish 1929, 1932; Smith 1941; 1961; Olson 1966; Priegel 1967; Ne~son 1968a, 1968b; Hubbs 1971; Hardy 1978; McElman and Balon 1979; Elston et a1. .. 1981;.>. Auer . 1982;.McElmall_1983J_.SimQ.p...c._tQ.~~Lt;lnd Bulkowski and Meade 1983) and were not included in the preSeIlt study. Species included in the .present study include: river darter, Percina shumardi (main streamUMR); northern logperch, f. caprodes semifasciata (main . stream UMR and tributar;f,es) ; striped fantail darter, •Etheostoma flabellare lineolatum· (small .to moderate streams and tributaries); least darter (a watch species),E. microperca (small streaIUS and tributaries); stripetail darter, E. kennicotti (small to moderate stream. tributaries); and spottail darter, E. sguamiceps (small to .moderate st:reamt:r.:f.but.filr:i.es) ••• None of the above species have had a formal description of larvae or early ~ juveniles. The present study describes and compares the development of these six species and developes a means to distinguish larvae of the genus Percina and Etheostoma. r t f MATERIALS AND METHODS Specimens were collected.from the Upper MisS:l.ss:t.ppiR!veibflfJJ.Il from 1981 to 1984. Additional supplemental material was procured from other institutions. Additional material was received from George Johnston and Bill Kowalski, Dairyland Power····Cooperative{DPC);cJ()hll.cD()I"cI"'IILandccDavid Jude, Great Lakes· Researc.h Division (GLRFLCn LaWferice>M. Pa.ge , Illinois Natural History Survey·(INHS);·· Robert R• Mille:ta.ll.dDQuglasW. Nelson, The University of Michigan Museum Milwaukee Public. Museum (M-PM); and LynnPfll"eIlti,. Gh.icflgoF'J.eld~\lseumof Natural History (FMNH). Darter species were spawned in·· aquaria, cc ·andc--laI'vae~weI'e-cthen preserved. Species spawned included: northern logperch, Percina·caprodes semifaciata;: spottail darter, Etheostoma sguamiceps (INHS); stripetail darte:t,E. kennicotti (INHS); least darter, E. microperca; and striped " - _.... ""."-~ .. -~ ....._. fantail darter, -E. flabellare lineolatum. The river darter, -P. shumardi were collected as adults, artifically fertilized (Bagenal andBaum 1971; Strawn and Hubbs 1958), and maintained according to Strawn (1956). Field collections were sorted with a dissecting microscope and all non percid eggs and larvae were removed. Each collection of eggs or spawn of eggs were placed into small 250 ml containers (ca. 50 eggs/container) and ~maintained at room temperature (20 to 23 C). Light conditions were maintained artifically with reflector hoods using a white incandescent 40 watt bulb. The photoperiod was regulated to provide 10 hours of light' followed by 14 hours of darkness. Larvae were maintained in 20 1 aquaria or were separated into additional 250 ml containers (ca. 25 specimens/ container). After yolk 4 i 5 6 species identification and separation. Morphometric and meristic characteristics were measured for diagnostic evaluation of the species. However, C>IllY12DlCll"ph.ClJltetrica:nd 9 meristic characteristics were of use in separating taxa. The following morphometric characters were selected: total length (TL), standard length (SL), preanal length, predorsal length, head length, head depth,eyediarnet.er,..... snout length, body depth at anus, greatest body depth, caudal ,peduncle depth, and yolks8clength. Measurements are di.agrammed in Fig. 1 and defined as in Hubbs and Lagler (1958), with the following exceptions: (1) Preanal length - horizontal length from the tip of the snout to the posterior margin of the anus; (2) Standard lenkth - horizontal length from the tip of the snout to the posterior margin of the notochord; (3) Body depth - vertical distance from the ventral margin at the anus to the dorsal ~urface of the body; (4) Head length - horizontal distance from the tip of the snout to the anterior margin of the pectoral bud or fin. Measurements were made at least three months after preservation. This ensured adequate time for fixation and would reflect shrinkage amounts observed in most environmental impact studies. Stobo (1972) reported the effect of formalin on the> length of yellow perch to. be negligable. Shrinkage began almost immediately in fish smaller than 150 mm and was largely complete within 24 hours. Decreases in length were reported to be Jt; 1.35 %for small yellow perch. Shrinkage of yellow perch after 72 hours was 7 % (Engel 1974). Data were recorded to the nearest 0.01 mm, but were rounded to the nearest 0.1 ~ in the text. The following meristic characters were examined: preanal myomeres, postanal myomeres, total myomeres, spinous dorsal rays, soft dorsal rays, FIG. 1 .. '.~ Diagrammatic representation of·morphometric characteristics ... rI 7 ? TOTAL LENGTH 1 STANDARD LENGTH PREANAL LENGTH PREDORSALLENGTH HEAD LENGT SNOUT LENGTH , ': GREATEST BODY . / DEPTH ~ 'EYE LENGTH I YOLK SAC LENGTH' ~ r 8 pectoral rays, pelvic rays, anal rays, and caudal rays. Paired. fin rays were counted on the left side whenever· possible•. Myomere counts were defined by Snyder (1979). Incipient fin rays were not enumerated but identified whenever formation was occurring. Preanfd.~lldpostanalmyomere counts were not included on early juvenile specimens since squamation reduced the visibility of the myomeres andireduced theiicicillraciyoftollJ:J.ts. Individual species accounts have descriptive detail presentedin1mm inter'Valsoftotallength (Fuiman 1981; Simon 1985). This.wasdonein order to eliminate bias of any single terminology and> allow the reader to assign the preferred terminology when applicable. Lengths·.. ·.···wereexpressed as total length throughout the text unless where noted. Text is presented in telegraphic style for brevity. Specific ··characteristics shared by members of common subgenera are also important in understanding the phylogeny of the tribe Etheostomini. Natural historynotesintlude such information as spawning period in the Upper Mississippi River, temperature at egg collection . site, egg incubation time, drift patterns when appropriate, habitat utilization of larvae, juveniles, and adults if different. ~ separate section compares published information on the larval development of the species. Illustrations of lateral aspects of the larvae were originally photographed using a SLR camera mounted on a Bausch and .. Lomb microscope• Prints were enlarged to 5 x (.7 and either projected on a wall and drawn or had detail drawn over the photograph using a permanent ink. Photographs were then rinsed with Farmers reducing solution removing the photographic image and leaving only the jrawing. The pigmentation pattern illustrated' ~ in this study was the pattern most commonly observed. Some variation may be observed in individuals which have either greatly expanded or contracted melanophores. rIt SPECIES ACCOUNTS NORTHERN LOGPERCH, Percina caprodes semifasciata (Rafinesque) The northern logperch, Percina caprodes semifasciata is one of the most common pettids in the Upper Mississippi River. Logperch are common in most northern pools to Pool 17, and are()tta.s10I1.EJ.J.:tygQ1-1-~g!-~<:! !IIJ>()()ls_}O, .21, and 24 (Van Vooren 1983). The species is the most~dely distributed of all darter.a ranging from. Saskatchewan to eastern Quebec and south to the Florida panhandle and Red River system of Texas. Along the Atlanticcoaat the species ranges from the Hudson River to the Potomac River estuary (Thompson 1980; Kuehne and Barbour 1983; Page 1983). The logperc~"has an extensive database of information available on adult .taxonomy, and natural history. 'I'heJQgpe:rgl'l~f3<:!E:!f3c::.I"!1>E:!ci~I"()mthe Ohio River presumably in the vicinity of Louisville, Kentucky (Rafinesque 1818). Currently three subspecies are recognized: P. £. caprodes, P. £. semifasciata, and P. £. fulvitaenia (Morris and Page 1981). A revision of the eastern forms will undoubtably result in additional subspecies (Kuehne and Barbour 1983). Variation in P. caprodes pigmentation was studied by Moenkhaus\(1894). Reproductive biology was investigated by Reighard (1913), Winn (1958a, 1958b), Scott and Crossman (1973), and Becker (1983). Spawning initiates in April in Kansas (Cross 1967).aJld PeIlIlsylvania (Cooper 1978); in late March or early April in Oklahoma (Miller and Robinson 1973); during May in West Virginia (Nance 1978); from April to July in Michigan (Reighard 1913; Will 1931; Winn 1958b); from December to mid-May in Texas (Hubbs 1961; Hubbs 1985); from mid-April to late May in Arkansas·rHubbsand. t Strawn 1963: Hubbs 1985); in late May to late June in New York (Wright and Allen 1913); initiating in June in Manitoba (Scott and Crossman 1973); during April in Illinois (Smith 1979); from April to July in Wisconsin 9 Ir- X 10 (Lutterbie 1976; Becker 1983); during May and June in Minnesota (Eddy and Underhill 1974); from early April to late May in Indiana (Evermann and Clark 1920); and during ApI'i.1 and May in Missouri (Pfieger1975;Hubbs 1985). Logperch eggs were observed being destroyed during mid-July by white suckers in Douglas Lake, Michigan (Ellis and Roe 1917). The geographic variation in egg complement waS studied from and Gua.~~~l1P~ River, Texas (Hubbs 1958). Developmental t:eD;lperat:ure tolerances were examined between centraL....Texas and more northern stocks (Hubbs 1961; Hubbs and Strawn 1963). Hubbs (1971) performed:l.nt:e:r:specific and intraspecific crosses with other darters, and various natural hybrids are not Uncommon. Histological· developmeritofthedigestivesystem.and swim bladder posthatching was investigated by Dobbin (1941), and ... Grizzle and Curd (1978). Age and growth was studied in Wisconsin by Lutterbie (1976). The ecological significance of larval ontogeny was reported by Paine (1984). Larval development for the eastern· subspec~ies~R.-c.-.c8prodeswas reported\byFish·····(1932); May.and Gasaway (1967); Taber (1969); Cooper (1978) ; and Hardy (1978) •. Paine and Balon (1984)discllssecl the early " development ofP. c. semifasciata according to Balons'it:heoryof:saltatory ontogeny. This account is inadequate for identification since the circulatory system and cranial osteology was stressed. Specimens for the study were collected throughout the Upper Mississippi River basin TABLE ...1. Material examined of Percina caprodes semifasciata larvae and early, juveniles grouped by 1mm interval of total length (N = sample size). TOTAL LENG11I (_) NlJH&ER ~ LOCALITY 5 5 Mississippi River MINNESOTA: Mississippi River, Pool 3, 1 ad upstreall Lock andDall ~ (R.M. 798.0: GLFRC 0001, 09(0) WISCONSIN: Black River, near Onalaaka spillway (R.M. 5) , , WISCONSIN: Mississippi River ,Pool 7, near Rosebud island (R.M. 7q3.5) , 6 25 MiSSissiP'pi River MINNESOTA: Mississippi River, Pool 3, 1 IIi upstreall Lock "ndDsII'~ (R.M. 798.0: GLFRC 0971,0890,0407,0896,0894,0897,0900,0893,0406,0902,0889,0887) WISCONSIN: Mississippi River, Pool 7,:.';near Richmond island (Il.M. 712) WISCONSIN: Mississippi River, Pool 8, West channel (R .M. 699) WISCONSIN: Mississippi River, Pool 9, east shore of Willow island (R.M. 678.2) WISCONSIN: Black River, near Onalaska spillwsy (R.M. 5) !ILLINOIS: Mississippi River, Pool 14, near Cordova (R.M. 503 -504) 7 21 Mississippi Ri~er iMIHNESOTA: Mississippi River,'Pool 3, 1 IIi upstreall Lock snd Dell 3 (R.M. 798.0: GLFRC 0971, 0966, 0899, 0895,0888, 0886, 0896) 'WISCONSIN: Mississippi River, Pool 7, Leke Onslaska near ,Onalaska ,spillway (R.M. 703 WISCONSIN: Blsck River, Catgut slough nesr French island (Bl~ck River R.M. 2) WISCONSIN: Black River, near Onalsskaspillway,(Black River, R.M. ,5) 8 7 Mississippi lliver MINNESOTA: Mississippi River, Pool3,1 IIi upstreall Lock and Dall 3 (R.M. 798.0: GLFRC 0966, 0409, 0410, 0412, 0184) ILLINOIS: Ilississippi River, Pool 14, near Cordova (R.M. 503 - 504) 9 4 Mississippi River MINNESOTA: Mississippi River, Pool 3, 1 IIi upstream Lock snd Dam 3 (R.M. 798.0: GLFRC 0184) ILLINOIS: Mississippi River, Pool 14, nesr Cordovs (R.M. 503 - 504) 10 5 Mississippi River MIHNESOTA: Mississippi River" Pool 3, 1 ad...•. upstrea"II Loc,ksnd,. Dam ,3", (,R.M.798.0: GLFRC 0411, 0227, !l184) " , " " " ",' , "", " , , WISCONSIN: Miasissippi River" Pool 7, Lek~ Onalaska nesr Onslaskaspillway (R.M. 703 ILLINOIS: Mississippi River" Pool 14, ne~r Cordova (R.M. 503 - 504) 11 4 Mississippi River MINHESqfA: Missis8ippi River,! Pool 3, 1 ad upstream Lock and Dell (R.M. <798.0: ! GLFRC 0290, 0227, 0230) WISCONSIN: Mississippi River 'f Pool 7, Leite Onalsska nesr Onslasks spillway (R;M. 703 i 12 5 Mississippi River MINNl!SOTA: Mississippi River,! Pool 3, 1 IIi upstream Lock and Dall3 (R.M. 798.0: , GLFRC 0412, 0230, pt.13) ILLINO~S: Mississippi River,I Pool 14, near Cordovs (R.M. 503 - 504) 13 2 Mississippi lliver MINIlESfA: Mississippi River, Pool 3, 1 IIi upstre811 Locksnd Dell 3 (R.M. 798.0: GLFRC 0001) ILLINO~S: Mississippi River Pool 14, nesr Cordova (R.M. 503 - 504) 14 5 Ilississippi River MI~A: Mississippi River Pool 3, 1 IIi upstream Loclcsnd Dam 3 (R.M. 798.0: GLFRC 0414, 0227) ILLINOjS: Mississippi River Pool 14, near Cordova (R.M. 503 -504) i Mississippi lliver Pool 3, 1 upstreall Locksnd Dam 3 (R.M. 798.0: 15 3 ~ississ:ippi River MINNESOTA: mi I GLFRC 0416) ..... ILLINOIS: Mississippi River Pool 14, near Cordova (R.Il. 503 - 504) ..... Miasisaippi River Pool 14, near Cordovs (R.M. 503 - 504) 16 'Mississippi lliver ILLINOIS: 17 2 Mississippi lliver ILLINOIS: Missiasippi RiverJ Pool 14, near Cordovs (R.M. 503 -504) .. 18 2 Mississippi River ILLINOIS: Mississippi River, Pool 3, 1 mi upstresm Lock snd Dam 3 (R.M. 798.0: CURC 0416) MINNESOTA: Mississippi River, Pool 14, near Cordovs (R.M.S03 - 504) 19 2 Mississippi River WISCONSIN: Mississippi RiVer, poor 5, nesr Al.... (R.Mi. 751.5) 20 4 Mississippi River MINNESOTA: Mississippi River, Pool 3, 1 mi ups~resm, Lk Dam 3 (R.M. 798.0: CURC 0290, 0230) 21 4 Mississippi River WISCONSIN: Mississippi·~ver. PoolS, near Alma (R.M. 751.5) 22 2 Mississippi River WISCONSIN: Mississippi River, PoolS, nesr Alma (R.M. 751.5) 23 4 Mississippi River WISCONSIN: MisSissippi River, PoolS, near Alma (R.M,. 751.5) MINNESOTA: Mississippi River, Pool 3, 1 mi upstream Lock and Dam 3 (R.M.798.0: 1 .! CURe 0230) i 24 "ississippi Ri v~r WISCONSIN: Missiasippi River, PoolS, near Alma (R .M. 751.5) 25 MississiiPpi River WISCONSIN: Mississippi River, PoolS, near AlmB (R.M. 751.5) 26 4 Mississippi River WISCONSIN: Misaissippi River, Pool 3, 1 mi upstream Lod< and Dam 3 (R.M. 798.0: GLFRC 0230) 27 2 Mississippi River WISCONSIN: Mississippi River, PoolS, near Alma (R.M. 751.5) 28 Mississippi River WISCONSIN: Black River, Catgut slough nesr ialand (Black RiVer R.M. 2) 29 Mississippi River WISCONSIN: PoolS, nesr j 3~ Mississippi River MINNESOTA: Pool 4, Lake City (R.M. 773) R.M. denotes Mississippi River ss defined by the U.S. Corps of Engineers ,. 12 EGGS LITERAruRE Geographical variatioIl 8Ild sUbspecies differencesh/ilYebeenpreviously descr~bed for the egg morphology of the logperch, P.caprodes caprodes, with limited information on the northern logperch. The egg of the northern logperch was spherical. Paine and Balon ·(1984) deSlCribedcleavage;~arly embryonic development, and skeletal f()I'lIlB.tion .. of northern logperch from Ontario, Canada. EggS :fI'(»)lLJ'oung .a.Il~ Potter Creek,tributaries of.. Lake Erie, Ontario, Canada, averaged 1.23mm(Paine and Balon 1984;Paine 1984). The ova of the eastern subspecdes P. £.. c:.aprodes were adhesive, demersal, amber in color with a granularyolle,andaveragedL12mm(Evermann and Clark 1920; Cooper 1978). Winn (195813., 1958b) described eggs in Douglas Lake, Michigan as colorless and adhesive withd.iameters()fT~31mm~ The diameter was geographically variable in Texas averaging 1.74 mmin Kerr County, 1.63 in Travis County, 1.71 mm in the Colorado system, 1.60 mm in the Guadelupe system, and 1.49 mm intheBrazoR~RiyeL(llY1>.Q§.~!2§I).~ In Arkansas, ova ranged.from.l •.44JQm in the .Il1inoisRi,ver to l.59mm in the White River, and l.35mm in J3i.gMories Rive:r, Mi:~~ouri (Hubbs 1967). Hubbs (1985) states that the •l,ogPerchhas beencollsidered to represent two species (the southern· P. car.bonaria and the n()I"t:.~~I"Il P •. caprodes) by Thompson (1978), an action endorsed by Morris and Page. (1.981), and Page (1983) but not by Kuehne and Barbour (l983). Texas stocks would represent P. carbonaria if the two taxa are considered distinct species. UPPER MISSISSIPPI RIVER VLuvA~r ~ Mature eggs in the present study from Pool 5, Mississippi River and Mukwonago River, Waukesha County, Wisconsin ranged froml.08 to 1.36 mm ® r~ D ~ (N = 142, X = 1.2 mm). Eggs contained a single oil globule, were demersal, and adhesive with an unsculptured chorion. Yolk was pale yellow in color, and translucent with a moderate (0.02 mm) perivitelline space. Eggs of northern logperch were found in shallow littoral zones buried in sand or gravel in areas with minimal current. previous reports of spawning location for the eastern subspecies P. £. caprodes (Reighard 1913-; Ellis 1917 Forbes and Richardson 1920;Winn 1958a, 1958b; Scott and Crossman 1973; Pflieger 1975; Hardy 1978; Smith 1979; Becker 1983; Page 1983; Kuehne and Barbour 1983). LARVAE LITERATURE The larva of the northern logperch was described by Paine and Balon (1984), however, their description emphasized the development of the circula.tory system and ossification of skeleta.lelemefits ifithecranium~ Thisdescriptiofi was inadequate for accurate identification of field specimens since meristic, morphometric, and pigment characteristics were not included. The following information on the eastern subspecies P. £. caprodes is included for comparison. Paine .and ..Balon(1984) reported northern logperch larvae hatch at 4.3 to 5.9 mm. The eastern subspecies £~ caprodes ranged from 4.5 to 5.3 mm at hatching (Fish 1932; Taber Hardy 1978; Cooper 1978; Auer 1982). The yolk sac was depleted in of northern logperch:by 6.5 mm-(·Pa-inHnd. Balon 1.984)-r and-~~inP. i - caprodes by 6.3 to 7.0 mm (Cooper 1978; Auer1982). ··Myomere counts for northern logperch ranged from 41 to 43 total andiriclllded 23 preanal and 20 postanal myomeres (Paine and Balon 1984). The eastern subspecies possessed 20. to 23 preanal and 17 to 23 postanal myomeres (Fish··1932; May and Gasaway 1967; Taber 1969; Hardy 1978;· Cooper 1978; Auer 1982). The relll8ining information presented will concern information on the eastern subspeciesP. £. caprodes since additional information for the nort~ern logperch is lacking. Fin ray development initiated in the caudal fin at 11.5 to 12.5 mm larvae in P. £. caprodes(C()opeI'1978;A\1eI'~..1982). Hardy (1978) reported incipient fin ray development at 10,.5mm. Notochord flexioIlocc\1I'I'ediIl P. £. cap:rodes at 11.0 to 12.2 mm(Hardy 1978; Cooper 1978; Auer 1982). All fin rays except. the spiny dorsalalldpelvic fins were formed at 15.0 rom (Fi.s~ 1932; Cooper 1978). All finl3werecompletely formed by 22.0mm(Cooper 1978). The finfold remained along the edge of the ventral and d.orsal Ili8rgin of the caudal fin (Cooper 1978). Morphometric informatioll. ispresentedmfor P•...h . caprodes.in~Cooper (1978), and to a limited extent in Auer (1982). PigmeIltation·patteI'Ilswere described by, Fish (1932), Hardy (1978), Cooper (1978), and Auer (1982). UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Characteristics of length and depth are described for larval and early juvenile northern logperch including the length at formation '. of morphological structures (Table 2). At lengths 5.4 to O~,j rom hatched): pectoral buds present without incipient rays; first pectoral rays (7.1 to 8.0 rom); yolk sac elongate and slender, yolk translucent; anterior oil· globule; head·nat deflected-.oyer ..... J.Qlk..-=8.aco;~ •. jaws undeveloped, developed at 6.6 to 7.6 rom. Yolk absorbed (7.0 to 8.0 rom); j' eyes spherical. First dorsal rays formed soft dorsal (10~0 to 11.5 mm), spinous rays formed (10.6 to 14.4 rom); incipient dorsal fin margin TABLE 2. Morphometry of pf'rcina cap_rode~ ~ 25 6.6(6.0-6.9) 6.3(5.8-6.7) 3.6(3.0-3.8) .92(.86"1.2} .•22(.16".30) .39(.32-.42) .44(.36".56) .83(.62-1.0) .23(.18-.28) 7 7.3(7.0-7.9) 7.0(6.7-7.5) 4.0(3.7-4.4) 1.1(1.0-1.3) .24( .20-.28) .41( .38-.44) .52( .44-.66) .91(.72-1.0) .25(.20-.34) 4 8.2(8.0-8.8) 7.9(7.7-8.5) 4.4(4.2-4.7) 1.5(1.3-1.9) .29( .20-.48) .48(~40-.64) .59( .52-.76) •92(.72-1. 2) .28(.24-.36) 4 9.5(9.2-9.8) 9.1(8.9-9.5) 4.9(4.6"5.2) 2.0(1.9-2.1) .45( .40-.48) .63( .60-;64) .85(.74-.92) 1.2(1.1-1.4) •39( .36".40) 5 10.5(10.0-10.9) 10.0(9.4-10.6) 5.8(5.4-6.1) 2.2(2.1...2.5) .50(.44-.56) .65(.56".76) .98(.76"1.3) ;4(1:3-1.6) .49(.36".64) 4 11.5(11.1-11.9) 10.5(10.0-11.0) 6.5(6.1...6.n 2.2(2.0-2.4) .49( .44-.65) .65( .64-.68) 1. 2(1.0-1.3) .5(1.4-1.6) .65(.50-.80) 5 12.5(12.2-12.9) 11.4(10.9-11.8) 7.0(6.8-7.4) 2.6(2.4-2.8) .58( .52-.60) .80( .56-1.1}' 1.4(1.3-1.4) .6(1.6"1.7) •77(.68-.88) 2 13.5(13.3-13.7) 11.9(11.8-12.0) 7.4(7.2-7.6) 2.7(2.6"2.8) .62 •72(.70-.74) 1.6(1.5-'1.6) 2.0(1.9-2.1) .92(.90-.94) .97( .80-1.2) 5 14.3(14.1-14.6) 12.7(11.9-13.2) 8.1(7.5-'9.0) 3.1(2.6"3.6) .66( .44-.96) .86( .80-1.0) L6(1.4-2.1) 2.0(1.7-2.7) 3 15.6(15.3-15.8) 13.8(13.8-13.9) 8.8(8.7-9.0) 3.4(3.2...3.6) .73(.68-.74) .96(.92-1.0) 1.8(1.7...1.9) 2.2(2.0-2.4) 1.2(1.1-1.3) 16.9 15.1 9.7 4.1 1.0 .96 1.9 2.6 1.4 2 17.2(17.0-17.3) 15.0(14.7-15.3) 9.6(9.3-9.8) 4.0 1.0 1.0 2.0 2.5(2.4-2.6) 1.4 2 18.4(18.2-18.6) 15.9(15.5-16.3) 10.4(10.1-10.7) 4.0(3.7-4.2) 1.0( .92-1.0) 1.1 2.2(2.1-2.3) 2.7(2.6-2.8) 1.4 2 19.1(19.0-19.2) 16.5 10.1(10.1-10.2) 4.6 1.1(1.1-1.2) 1.1(1.1-1.2) 2.5 2.8 1.4 4 20.5(20.2-20.8) 17.6(17.2-18.4) 11.1(10.8-11.6) 4.5(3.8-5.5) 1.1(1.0-1.3) 1.4(1.3-1.5) 2.7(2.5-'3.0) 3.2(3.0-3.4) 1.5(1.2-1.6) 1.5(1.4-1.6) 4 21.1(21.0-21.2) 18.2(18.0-18.4) 11.2(11.0-11.6) 5.P(4.8-5.1) 1.2 1.3(1.2-1.4) 2.5(2.2-2.7) 3.1(3.1-3.2) 2 22.2(22.0-22.3) 18.6(18.Q-19.2) 12.0 5.4(5.3-5.5) 1.4(1.3-1.4) 1.4(1.3-1.4) 3.0(2.9-3.1) 3.3 1.7 i ' ' 4 23.3(23.0-23.8) 20.109.8-20.3) 12.5(12.0-13.1) 5.14(5.1-5.6) 1.3(1.1-1.4) 1.4(1.4-1.5) 2.8(2.4-3.1) 3.3(3.2-3.3) 1.7(1.5-'1.8) 1.8 1 .24.0 20.7 13.0 5.8 1.4 1.4 2.9 3.3 1.8 1 25.5 21.0 13.5 5.8 1.5 1.2 2.5 3.2 i 1.9(1.8-1.9) 4 26.3(26.0-26.5) 22.9(22.5--23.5) 14.3(14.0-14.5) 6.11(6.0-6.2) 1.6(1.4-1.J) 1.7(1.6-2.0) 3.7(3.0-4.3) 3.9(3.7-4.1) ! ! 1.8 2 27.0 23.4(23.3-23.5) 14.2(13.5-15.0) I 6.5 l.8(1.5-2.Q) 1.8(1.5-'2.0) 3.5(3.3,..3.8) 3.8 2.2 1 28.8 25.0 15.0 i 6.7 1.6 i 1.8 3.6 4.4 2.4 1 29.5 25.0 15.5 7.0 2.0 4.0 4.4 2.0 .1 I-' 2.5(2.4-2.5) 2 30.5 26.4(26.Q-26.8) 16.8(16.6-17.0) 6.5 1.9(1.8-2.~) 2.0 4.2(4.0-4.3) U1 " i I: . I (a)I' Body depth measured vertically from the dorsal tovent~al surface at poster1o~ portioDof anua . (b)! Greatest body depth measured vertically frOll dorsal t~iventral aurface including the yolk sac (c) Caudal peduncle depth measured vertically fro- the dorsal to ventral aurface at penultimate myomere 16 partially differentiated (ca. 14.1 to 15.4 JJDD.),coBlP1etely differentiated (15.8 to 17.0 mm); spinous dorsal fin originoyeI" preaJla1.lIIYQmere I,soft dorsal origins.:iiua.tedoYeI". pI"e;anal myomeres21to23(18.6Dlm) ;PI'~dorsal length 30.9 %standard length (5.4 to 30.5 mm); notochord flexion (7.5 to 10.0 mm), simultaneously with first caudal fin ray formation (7.5 to 9.2 mm); caudal fin lunate (14.1 mm); first anal fin rayiforIlled (Ca:;11d5mtn); incipient anal fin ~rgin partially differentiated (ca. ·13.7 to 14.5 mm), completely differentiated by 16.9 mm; pelvic buds formedahterior.to dorsal fin origin (ca. 10.0 to 10.9 mm), first pelvic fin rays formed (10.6 mm). Swim bladder formed, evident after yolk absorption; entirefi:nfoldabsorbed (12.2 to, 14~2 mm). Supraorp:i.tal, pr~()p~rEllloDlElIldibular,sl11>t~poral, and infraorbital head.·~canals started forming (15.4 to 20.3 rom), supraorbital and infraorbital head canals completely··formed(23.3mm),~inf-I'aorbital canal complete without interuption (29.5 mm); preoperculomandibular, supratempora~, and lat~ral canal as well as the lateral line formed (20.8 mm). Scales started to form posterior, near caudal peduncle (21.2 to 23.8 mm) .. The breast is \unscaled, however the cheek and opercle are scaled (26.0 to 28.8 rom). Mandible overhangs maxillary at all lengths (when functional mouth parts are present). Meristics Myomere number constant posthatching, preanal myomeres 19 to 24 (N = 104, x = 21.8, mode = 22),postanal counts 18 to 23 (N = 104, x = 20.4, mode = 19) with 39 to 46 total myomeres. Total vertebrae numbering 42 to 44 (N = 4, x = 43) including one urostylar element, from specimens collected in PoolS, UMR and Catgut slough, Black River, Wisconsin (13.6 to - 20.8.mm). Scales in the lateral series ranging from 70 to 87 (N = 10, x = 78.7, mode = 81). Paired and median fin ray values and length of .. 17 TABLE 3. Selected meristic values and size (mm total length) at the apparent onset of development for Percinacaprodes semifasciata. Mean values are underscored. Rare or qU'estionable extremes are in p8rentheses~ The number of secondary rays of the median fins are in lowercase Roman numerals. (a) Attribute 1 Event Present Study Literature (1-8,10,17,18) Dorsal Fin Spines 1 Rays XII-XIII-XIV/12-13-14-16 XIII-XIV-XVI/14-15-16-18 - (9-11) fin:t rays formed 10.0 - 10.6 12.1 - 15.6 (9,10) adul t cCJlllplenlent formed 15.8 - 17.0 22.0 Anal Fin Spines 1 Rays II/9-.!Q::.!l II/8-10-11-13 - (9-13). first rays formed 11.5 10.5 - 12.1 ,10,13) adult complement formed ': Caudal Fin Rays vii-xv, 9 + 8, vii-xiv no information n:C5-·':~12;5--.--~2".(9,10,13'1 first rays formed 7.5 .... 9.2 (9,10) adult complement formed 12.2 - 14.6 15.0 (1,2,17) Pectora~ Fin Rays 13-14-15 12-14-16-17 ---C9) first rays formed 7.1 15.6 adult complement formed 10.0 no information (1-8,10,17,18) Lateral Series - Scales 70 - lJ- - 87 67-JJ::11.-99 (1-2,9-17,18) Myomere ,I Vertebrae 39-41-46/42-~-44 39-43-46/40-42-45-46 - '(9-15) preanal myomeres 19-22-24 19-23-24 - (9-15) postanal myomeres 18 - 12. - 23 16 - 22 a.1. Page 1983 2. KuehnE' and Barbour 1983 3. Smith 19794. Morris and Page 1980 5. Trautman 1981 6. ForbeR and Richardson 1920 7. Jordanard ',' Ev(:'rnlanll ,.• 18968. Edd}' and Underhill 1974 9. Cooppr 1978 10. Hardy 1978 11. Fish 1932 12. Taber 1969 13. Auer 1982 14. Paine lind nalon 1984 15. May and Gasaway 1967 16. Bailey and Gosline 1955 17. Scott ond Crossman 1973 18. Becker 1983 "'~c·======c •• c"""" • 18 appearance are presented in Table 3. Pigmentation Length 5.4 to 6.0 rom (newly hatched): sparce pigmentation (Fig~ 2); retinae black, pigmentation limited mainly to scattered melanophores at mid- ventral yolk sac, few to no melanophores present afJ'i'll.aveiitralofp-ostanal myosepta. Length 6.2 to 6.8 rom: eyes black; pigmentation absent on cranium and along dorsum. Scattered melanophores present on yolk sac conientrating near mid-ventral; dorssl outline of gut pigmented; single melanophores situated at mid-ventral of postanal myosepta at every myomere. Length 7.1 to 7.9 rom: pigmentation essentially unchanged from previous "c condition with the exception of severalJ'i'lelanopnores---formea---at-aorsal posterior base of optic lobe (Fig. 3). At lengths greater than 7.6 rom ventral pigmentation becoming subdermal. Length 8.5 to 10.9 rom: pigment absent oncraniJml_and_Qn9Q:rsum. Single melanophorepr.esent QDc1eithra; several melanophores outline dorsal portion of gut near anus. Single, mid-ventral, punctate melanophores at almost every postanal myosepta becoming subdermal as length approaches 10.9 rom (Fig. 3). Length 11.4 to 13.7 rom: pigment absent overlD.uchof I :1 ,i ------j 0:) WW~ I '8 FIG. 3 Percina caprodes semifasciata, northernlogperch larva; Upper Mississippi River, Pool 7 a. c7~5 mm TL b. 9.2 mm ~ ,,~,TL,.~~,,_,_~,,~_~~~, ,__ ,~ "'""------_--:..._------_.... Iwwa. wwa. °8 oc: 21 additional punctate melanophore near anus; most melanophores subdermal; several melanophores at base of hypural Length 14.6 to 16.9 mm: limited pigJIleritationapparent several 'punctate melanophores present on ventral, almost always near ventral portion of anus;postanal myomeres with pigment on myosepta above anal fin and caudal peduncle with subdermal pigment apparent·· .·atlengthsapproaching 16.9 mm; several midlateral punctate melanophores near caudal peduncle near postflexion notochord (Fig. 4). Length 17.2 to 20.2 mm: punctate pigmentation near ventral. anus; punctate melanophores at interdigitation of anal fin lepidotrichiawith /' pteriophore extending onto soft rays of anal fin; mid lateral, ventral, and dorsal accumulat.ion of melanophores near caudal peduncle; pigment accumulating forming a spot at center of caudal fin base. Length 20.5 to 21.2 mm: dorsal cranial pigmentation consists of punctate melanophores on snout, cerebrum, optic lobe, and laterally on operculum; melanophores beneath pectoral fin laterally; at mid ventral of anal fin, at ventral caudal peduncle, and midlateral cluster at caudal peduncle. Several clusters of dorsal melanophores at base of soft dorsal and at base of caudal fin. Melanophores scattered on interstitial membranes of spinous and soft dorsal fin, and anal fin. A vertical line of melanophores located at center of caudal fin base extendinghypaxially on interstitial membrane. Length 21.6 to 23.8 mm: dorsal cranium with stellate melanophores over cerebellum, and optic lobe. Melanophores on snout, mandible, and chin; weak postorbital bar forming; 7 oval midlateral clusters formed:' present anterior to dorsal, just posterior of anterior spinous dorsal, near base of posterior spinous dorsal, at anterior of soft dorsal, at mid soft dorsal, posterior base of soft dorsal, and at caudal peduncle. Dorsally, 6 1 FIG. 4 .. ': Percina caprodes semifasciata, n.orthern logperch, a. 14.6 mm TL Catgut Slough, Black River,Wisconsin br~~26i§~mm~TLUpperMississippi .River, Pool 3. I I I I I ______JI H wWI., :.... °8 i ! I J • 23 rectangular blotches present anterior to spinous dorsal, at mid spinous dorsal, at posterior base of soft dorsal, and at anterior base of secondary caudal fin rays. Subdermal dorsal pigment present on s'Wim.bladder. Postanal pigmentation present at interdigitation of pterigiophores 'With anal fin lepidotrfchia, and mid ventral caudal peduncle base. Scattered melanophores on membranes of spinous dorsal and caudal fin, and mid str~p~~Ilthe soft dorsal and anal fins. A mid .lateral spot at b.ase of caudal fin vertically radiating into hypaxial musculature. Length 24.9 to 30.5 mID (juvenile): cranial pigmentation over cerebrum, and optic i lobe; preorbital bar formed from snout pigmentation; maxillary and chin with melanophores; chevron shaped cl\lsterfotmedbetween operculum and optic lobe pigmentation; stellate melanophores outlining ventral portion of orbit. Vertical stripes ,numbering 11 to 13 radiating from dorsum with 8 to 9 extending past lateral line. Scattered melanophores on membranes of: spinous dorsal, pectoral, and caudal fin; mid stripe formed in soft dorsal, and anal fin; pelvic fin devoid" ofpigment.~~Ad-iamond-shaped cluster of melanophoresonmidlateral at caudal base extending vertically into ventral hypaxial musculature (Fig. 4). Natural History The northern logpetth inhabits a variety of environments ranging from streams and rivers usually occurring over gravel or sand in water of moderate current (Forbes and Richardson 1920; Thomas 1970; Scott and Crossman 1973; Eddy and Underhill 1974;Rf1-:Leger~1975; Smith 1979J~,'rrautmfln 1982; Phillips et al. 1982; Page 1983; Kuehne and Barbour 1983; Becker 1983; Simon 1983). Spawning initiated in the Upper Mississippi River from late April .until early June at temperatures between 12 and 15 C. Northern 24 logperch were seen in drift collections when water temperatures approached 12.7 C. Collection in drift sampling in the Black River, Wisconsin indicated a tendency towards surface drift .Aqu.ar:i:l1ll1h.~ld northern .logperch ,were observed" actively swimming near the surface under daylight condit.:l..ons. The exact 'characterization of larval microhabitat'utilization was not possible since larvae were present:LtliJ1l()st.Lhabit.st.~.ty.pes encountered (except non-flowing waters). EarlYju.veniles(T3:7t()30.S rom) were collectedfro1'll.deephsb:i.t:a.t.s whichrsngedfrom 'ba.ckwaterswithmuck and silt bottom to main channel border sand substratesWithisl:i.ght current. Eggs .incubated.in laboratory aquaria maintained at 22Chatchedafter 120 to 144 hours. This agreed with data observed Grizzle and (1978). At 16.S'.C, hatching occurred after approximately 200 hours (Cooper 1978). r RIVER DARTER, I Percina shumardi (Girard) The I'iverda.rter, Percina shumardi is a common inhabitant of the Upper I { Mississippi River. .It is commonly collected in Pools 2 through 26 (Van Vooren 1983). TheI'iver darter ranges from the Gulf coast including the I Mobile basin of Alabama and the lowerGlladalll'PeRiverip.'1'~~I!S, north through the Mississippi River drainage including mos~ major tributaries, I east into the Lake Erie and Lake Huron basin including Manitoba and western I Ontario. Distribution in the Ohio River basin' is SParce. Disjunct j populations exist in the San Antonio bay drainage, Texas and the Sabine River, Texas and Louisiana (Gilbert 1980; Page .1983; Kuehne and. Barbour 1983). Limited information .. is.available-on-the_life__ hist.ory__ of__.tll~ri ver darter. The river darter was described from specimens collected from the Arkansas River, near Fort Smith, Arkansas (Girard 1860). It is an inhabitant of larger shutes and riffles of large to moderate sized streams and. rivers usually associated with gravel substrates (Cross 1967; Clay 1975; Trautman 1981;Pa,ge1983; Kuehne and Barbour 1983). TI'autman (1981) reported that river darters seem to be more tolerant ofturbid rivers. Limited information on the > reproductive biology of the river darter is available. Scott and.· Crossman (1973) suggest that the species spawns similarly to the blackside darter, P. maculata and ... channel darter, P. copelandi. Spawning occurs over scattered rubble and gravel substrates usually at depths greater than 0.5 m in areas with strong current. Spring migration of adults has been suggested by Trautman (1981), who collec Ohio specimens during spring in flooded fields adjacent to the Scioto River. Scott and Crossman (1973) suggested that the river darter spawns as late as June or July in Manitoba and adjacent Ontario. Becker (1983) 25 26 reported spawning occurring from April to June in Wiscorisiri;iriMinIlesota and Illinois spawning occurs from April to May (Eddy and Underhill 1974; Thomas 1970); during April in Kansas (Cross 1967); and possibly as early as February and March in the Little Tennessee River (Starnes 1977). Specimens from the present study were collected from various locatiQns in Pools 3, 5, 7, 8, and 9 (Table 4). ~ LITERATURE The river darter has an egg morphology similar to most members of .the family Percidae• ','IThe diameter of eggs in the literature was reported to be 1.2 mm in Wisconsin (Becker 1983), and .L67mmin_the~Gu.adal.u.p~EiYer in Texas. No additional information is available for inunatureorm.ature eggs in the literature. UPPER MISSISSIPPI RIVER DESCRIPTION During ovary inspection,the immature unfertilized ova from the Black River, Wisconsin were spherical with diameters ranging from 0.82 to 0.94 mm (N = 13, x = 0.89). This suggests that the river darter is a fractional , spawner depositing more than a single cluster of eggs. Mature fertilized eggs of the river darter were 1.0 to 1.3 mm (N = 2, x= 1.2 mm) in diameter. Eggs were demersal, slightly adhesive, and unpigmented with translucent yolk. Mature eggs had a single oil globule, a moderate perivitelline space (0.02 mm), and an unsculptured chorion. Eggs were found buried in fine gravel and sand in the Black River, Wisconsin. Spawning adults were collected downstream from the Onalaska spillway in water less than 1 m over coarse sand-gravel, and cobble. TABLE 4. Material examined of Percina shumard! larvae and early juveniles grouped by Imm intervals of total length eN == sample size). TOTAL LP1lGTII (_) ~ ~ ~ 4 Mississippi, River liISroHSI11: Mississippi River, Pool 5, neST AI... (R.M •• 751.~) WISCONSIN: Mississippi River, Pool 7, Leke Onslssks near R~sebud islsnd (R.M. 704.5) MINIlESOTA: Mississippi River, Pool 3, 1 IIi upstre811 Lock s~d Dam 3 (R.M. 798.0) MINIlESOTA: Mississippi River, Pool 9. near Willow islsnd (~.M. 677.5) WISCONSIN: BIsck River, nesr Onslssks spillwsy (R.M. 5) 5 31 Mississippi River liISCONSIN: Mississippi River,' Pool 5, neer AI_ (R.M. 751.~) WISCONSIN: Mississippi River;' Pool 7. nesr RichllOnd islsnd {R.M. 712) WISCONSIN: Mississippi River, Pool 8;\ West chsnnel (R.M. 699) WISCONSIN: Mississippi River, Pool 9, esst shore of Willowlislsnd (R.M. 678.2) WISCONSIN: Blsck River, n'esr Onslssks spill...y (R.M. 5) MINNESOTA: Mississippi River, Pool 8, West Chsnnel (R.M. 699) MINNESOTA: Mississippi River, Pool 3, 1 IIi upstresm Lock ~nd Dam 3 (R.M. 798.0) WISCONSIN: Mississippi River. Pool 5, neer AI... (R.M. 751.5) WISCONSIN: BIsck River, Cstgut slough neST French island (lliack RiverR.M. 2) WISCONSIN: BIsck River, nesr Onslasks spillway, (Blsck River R.M. 5) 7 3 Mississippi', River MINllESOTA: Misdssippi River, Pool 3, 1 IIi upstreall Lock s,nd Dam 3 (R.M. 798.0) WISCONSIN: Mississippi River, Pool 5, near AllIS (R.M. 751.5) WISCONSIN: Mississippi River, Pool 9, nesr Bad Axe Lending '(R.M. 675.0) 8 Mississippi River WISCONSIN: BIsck River, Cstgut slough nesr French islsnd (BIsck River R.M. 2) 10 Mississippi River WISCONSIN: BIsck River, Cstgut slough neSr French islsnd (BIsck River R.M. 2) Mississippi River WISCONSIN: Mississippi River, Pool 5, near AllIS (R.M 751.5) 2 Mississippi River WISCONSIN: Mississip~i River. Pool 7, Leke Onalssks (R.M. 706) 4 Mississippi River WISCONSIN: Mississip~i River, Pool 5, nesrAIIIS (R.M. 751.5) 3 Mississippi River WISCONSIN: MississiP~i River, Pool 5, nesr Alma (R.M. 751.5) Mississippi River WISCONSIN: MiSSiSSiP~i River, Pool 5, near Alma (R.M. 751.5) 5 Mississippi River WISCONSIN: Mississip~i River, Pool 5, nesr AI... (R.M. 751.5) 5 Mississippi River WISCONSIN: MiSSissip~i River, Pool 5, nesr AIIIS (R.M. 751.5) Mississippi River WISCONSIN: Mis"isSiP~i River, Pool 5, nesr AIma (R.M. 751.5) I Mississippi River WISCONSIN: Mississippi River, Pool 5, nesrAIIIs (R.M. 751.5) I Mississippi Ri ver WISCONSIN: Mississippi River, Pool 5, nesr AllIS (R.M. 751.5) Mississippi River WISCONSIN: Mississip~i i~iver, Pool 5, nesr AIIIS (R.M. 751.5) Mississippi River WISCONSIN: MississiPM River, Pool 5, nesr AIma (R.M. 751.5) Mississippi River WISCONSIN: Mi8SiSSi+ River, Pool 5, nesr AIIIS (R.M. 751.5) t-.> Mississippi River WISCONSIN: Mississip~i River, Pool 5, nesr Aims (R.M. 751.5) -..J I Mississippi River WISCONSIN: Mississippi River, Pool 5, nesr Alma (R.M 751.5) Mississippi Ri ver WISCONSIN: Mississippi River, Pool 5, nesr AI... (R.M. 751.5) 28 Current from the spillway was moderate arid formed an eddy. LARVAE LITERATURE No information on the larval taxonomy or early life, history of the river darter has been previously published. UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Characteristics of length and are described for larval and juvenile river d~rters including the length of formation of morphological structures (Table 5). At lengths 3.6 to4.0mm{newlyhatched).Lpec.toral fin present without incipient rays; yolk sac oval; yolk translucent; single anterior oil,' globule; head slightly deflected over yolk sac; no functional mouth parts developed, jaws developed (4.8 rom); yolk absorbed (5.5 to 6.2 mm); eyes oblong. rays formed anal (5.8 to 6.1 mm); first pectoral rayS formed (5.9 to 6.1 rom); first dorsal rays formed (5.9 mm); incipient dorsal fin margin partially differentiated (7.1 mm), completely differentiated (>8.0); spinous dorsal fin origin over myomeres 5 to 6, soft dorsal origin situated over preanal myomeres 13 .to. 14 (7.1to8.0mm); predorsal length does not significantly change at increasing lengths (31% standard length); notochord flexion (6.8 mm), after first caudal fin ray formation (5.8 to 6.6 mm); caudal fin emarginate (12.3 mm); incipient anal fin margin partially differentiated (> 8.0 mm); pelvic fin buds formed anterior to dorsal fin origin (7.5 rom), first pelvic fin rays formed (12.3 mm). Swim bladder formed, evident after yolk absorption (ca. 6.0 mm); gut slightly curved; entire finfold absorbed (10.8 mm). Supraorbital, TABLE 5. Morphometry of Percina shumardi larvae and early juveniles grouped by' Imm intervals of total length (N = sample size). Mean lengths followed by range in parentheses. Ll!HG'nl <_) (a) (b) (c) .!! 1.'Q!& STANDARD ~ ~ .§!!Q!!!. .m. BODY DEPl'Il GREATEST CAUDALPEOOIlCLE 7 .4(4.0-4.9) 4.2(3.7-4.8) 2.3(2.0-2.7) .69( .50-.92) _ .•15( .09-.20) .29( .23-.30) .27(.20-.34) .63( ~56-. 76) .13( .08-.24) 31 5.6(5.0-5.9) 5.4(4.8-5.7) 2.9(2.4-3.3) .96( .60-1.2) .19(.12-.24) .38( .28-.50) .40(.33-.60) .74(f60-·9O) .17(.12-.22) 13 6.2(6.0-6.7) 5.9(5.8-6.4) 3.3(3.0-3.5) 1.0( .80-1.2) .20( .12-.22) . 38( .3?:-. 60) .45( .38-.50) .80(.60-.90) .19( .16-.26) 3 7.3(7.1-7.5) 6.9(6.8-7.2) 3.8(3.7-4.0) 1.3(1.2-1.4) .23( .22-.24) .50( .46-.56)' .60(.50-.70) .90(.70-1.1) .24( .22-.26) 8.0 7.7 4.5 1.8 .30 .50 .70 1.0 .30 10.8 9.3 5.6 2.2 .45 .65 1.0 1.4 .50 12.3 10.8 6.6 2.9 .60. .80 1.4 1.8 .90 2 13.4(13.2-13.6) 11.6(11.3-12.0) 7.1(7.0-7.2) 3.3(3.1-3.5) :65( .60-.70) .90( .80-1.Q) 1.5(1.4-1.6) 2.1(2.0-2.2) 1.0( .90-1.0) " 5 1~.4(14.1-14.9) 12.8(12.1-14.4) 7.6(7.2-7.8) 3.5(3.4-3.8) .80(.70-.90) 1.0( .85-1.1) 1.7(1.6-1.8) -2.4(2.2-2.5) 1.1(1.0-1.1) 3 15.3(15.0-15.5) 13.4(13.0-13.7) 7.9(7.5-8.3) 3.7(3.4-3.8) .80(.70-.90) .90(.70-1.2) 1.8(1.6-2.0) 2.7(2.6-3.0) 1.1(.90-1.2) 2 16.6(16.5-16.7) 14.6(14.5-14.7) 8.8(8.5-9.1) 3.9(3.6-4.2) .80 1.0( .80-1.2) 1.7(1.3-2.1) 3.0(2.9-3.1) 1.3(1.2-1.3) 5 17.4(17.0-17.9) 14.9(14.3-15.1) 9.3(9.0-9.9) 4.3(4.1-4.5) .9O( .80-1.0) 1.1(.90-1.2) 2.2(2.0-2.5) 2.9(2.6-3.0) 1.3(1.2-1.4) i 5 1,.1(18.0-18.5) 15.7(15.3-16.7) 9.6(9.0-10.0) 4.2(4.0-4.5) 1.0(1.0-1.1) 1.1(1 2.5'(2.3-3.0) 3.0(2.9-3.3) 1.4 I 19.2 15.5 9.6 4.2 .90 1.4 2.4 3.'2 1.4 5 ~t.3(20.0-20. 7) 17 .3(17.0-17.6) 10.3(10.0-11.0) 4.8({o.6-4.9) 1.1(1.0-1.2) 1.3(1.0-1 2.7(2.6-2.7) 3.5{3.4-3.6) 1.6(1.5-1.6) 21.4 18.0 10.9 5.1 1.2 1.3 2.9 3.6 1.6 22.0 18.3 11.0 5.3 1.3 1.3 3.0 3.6 1.8 23.0 20.0 12.8 ~.O 1.4 1.7 3.8 3.7 1.7 24.• 1 20.1 11.8 5.4 1.2 1.4 3.1 3.8 1. i 21.3 12.0 5.8 1.3 1.8 3.8 3.9 I 2 t,,,,·~~··,It ", 22.8 13.4(13.0-13.8) 6.4(~.3-6.5) 1.6(1.5-1.8) 1.8 4.0(3.8-4.2) 4.1(3.9-4.3) i i 27.8 23.5 14.0 ~.5 1.3 2.0 4.0 4.2 I I i .. , i dy depth measured vertically frOll.the dorsal toventrall surface at posterior pprtHm of altus N (b) ~reatest body depth messuredivertical1y from dorssl to v~ntral surface includinJ the yolk ~c 1.0 i. ., I I . I (c) ¢euda1 peduncle depth measured vertically frOll the doraal to ventral surface at !penultimate ,lIyOllere 30 infraorbital, preopercul()lIlandibular, arid lateralheadcartals formed (24.1 mm); lateral line began forming 22.0 mm; scaleepresent mm, Meristics Myomere number constant posthatching, preanal myomeres 18 or 19 (N = 78, x = 18.1), postanal myomeres 22 to 26 (N = 78, x = .1), with 40 to 44 total (N = 78, x = 41.2). Total vertebraent111lbei':i.ng39t:C>41(N= 3, x = 40), including one urostylar element, from specimens cleared and stained from ,the Upper Mississippi River PoolS, Wisconsin (13.6 to 22.1 / - mm). Scales in the lateral series ranging from 46 to 57 (N = 18, x = 48.4, mode = 49).' Paired anc1 median fin ray values and length at appearance are presented in Table 6. Pigmentation Length 3.6 to 4.0 mm (newly hatched): sparce pigmentation (Fig. 5); eyes black; pigmentation absent on cranium and dorsal surface. Lateral pigmentation with two parallel lines of melanophores at almost every midlateral and ventral postanal myoseptum, with midlateral line extending only from anus to about the middle of the postanal length. Scattered melanophores ,. limited tomventral half of horizen,tal1ym,bissee-tedyolk" sac with greatest concentration at the midventralsurface. Pigment outlining dorsal and ventral portion of gut posterior of yolk sac;verif:rsl'yC>lk sac pigmentation extending obliquely to dorsal surface of gut. T 31 TABLE 6. Selected mel:"i:S'tic valuesaIld13i2'.~ (DIJJl totaIlengt:l1.}a.tthe apparent onset of development for Percina shUDlardi. Mean I values are underscored. Ra.reorquestionableexttemesare I ! in parentheses. The nUDlberofsecondary rays of the median fins· are in lowercase Roman numerals • ------{a) Attribute / Event Present Study Literature .. ~ .... ~~~.~ .. - ~ ~~·~------(2··9 11) ...... - , Dorsal Fin Spines I Rays VIII-!-XI/12~13-14 IX-~-XII/11-13-15-16 first rays formed 5.9 no information adult complement formed 13.2 no information 11/10_11_12_f;,4-9,11) Anal Fin Spines / Rays II/(B)·-10--11-12 first rays formed no information adult complement formed 8.0 no information Caudal Fin Ray~" ix-xiv, 8 + 7, x-xii no information first rays form~d 5 no information adult complement formed information (4,5,11) Pectoral Fin Rays 11- 14 13 - 15 first rays formad 5.9 no information adult complement formed 6.7 _-_._ ---~no__iniQrmatJQn (1-9,11) Lateral Series - Scales 46 - 48 ... 57 46-52-57-58-62 I (4,5,10,11) Myomere / Vertebrae 40~1-44/39-40-41 no information/37~38~0 I , preanal myomeres .!!! - 19 no information postanal myomeres 22 - 23 - 26 no information a. 1. Douglas 1974 2. Pflieger 1975 3. Trautman 1981 4. Kuehne and Barbour 1983 I 5. Page 1983 6. Forbes and Richardson 1920 7. Jordan and Evermann 1896 8. Eddy I and Underhill 1974 9. Clay 1975 10. Bailey and Gosline 1955 11. Scott and Crossman 1973 t I I· I~ FIG. 5 .. '~ Percina $hl.J1llardi, river darter (newly hatched larva), 4.0 rom TL Black River, Wisconsin a. dorsal b. lateral· ...... _.. ~._f...!....J~.~}J.!~~]._!.-.... __ . ww " ,. : .... : ...... " \ . \, \ Z£ 33 Length 4.8 to 5.9 mm: cranium and dorsal surface without pigmentation; several stellate chromatophores present at ..cleithra; . a few scattered melallophotes •• ondorsal half of yolk sac, concentrated more heavily at midventral. Lateral pigmentation forming an inV'~I't.:~dv ....patternabove the anus.extending dorsally to mid-body; an oblique ···1I1fdlateral stripe of melanophores at postanal myosepta apex extends post_er:tQLfJ:Q.I!l_J~~'Us !()_ the middle of /the ·.postanal length. Ventral pigmeIltatioIl~talmost·every postsrislmyosepta; melanophores encircling anus. Length 6.1 to 7.5 rom: several melanophores below mandible; melanophores forming postorbital chevron; melanophores outlining the . notochord (Fig. 6). Length 10.8 to 12.3 rom: cranium with melanophores concentrated dorsally over optic lobe and cerebrum; snout with pigment posterior to l FIG. 6 ...: Percina shumardi, river darter larva, Upper Mississippi River, Pool 5, Wisconsin a. 6.2 rom TL b. 8.0 rom TL. ______.-Jir "8 wwa. ,,' cd i$. 35 rj1. maxillary; 5 eliptical clusters of melanophores from nape· to· caudal peduncle; several melanophores near postflexion notochord. Cranium with forming preorbital, and formed postorbital bars; preopercle with stellate melanophores; margin of opercle with pigment at apex. Dorsal gut pigmentation subdermal; several stellate melanophores present ventrally; a single midlateral line of pigment present from the cleithra to the tip of the notochord. An oval cluster ofmelanophores present near caudal peduncle; o.blique line ofmelanophores from anus to caudal pedllncle clusters. Melanophores at base of caudal peduncle forming a hook~shaped vertical line on the lower half of caudal fin; caudal fin with a few scattered mela.nophores on rays. Ventrally, margin ·()f opercle outlined; ....',. pigment extending between pelvic fin posterior to sides of anus; postanal pigment present from anus to end of caudal peduncle at midventral (Fig. 7). \ Length 13.6 to 15.5 mm: cranium with concentrated melanophores present dorsally as described for 10.8 to 12.3 mm interval; preopercle and interopercle outlined with melanophores; developed--preor·bi:tal· and postorbital·tear drops; 7 to 8 dorsal blotches not extending to notochord. Posterior to opercle, pigment surrounding air bladder and incorporating an oval melanophore cluster; 4.additional oval clusters to caudal peduncle. Vertical hook at caudal peduncle base darkening. Additional pigmentation present on rays of caudal fin; few melanophores present on spines and rays of dorsal fin. Ventral pigmentation remains as in the previous condition. Length 16.5 to 20.7 mm: dorsal pigmentation with melanophores concentrated over the optic-lobe, andc=cCekebellulIL __e.xten!tiDi_into llPPer sections of the operculum; from nape to base of caudal peduncle are 6 to 8 oval dorsal clusters; u-shaped cluster of melanophores posterior to . maxillary. Lateral pigmentation includes 7 to 9 oval clusters; formed FIG. 7 Percina shumardi, river darter, a; ·12;3mm TL larva· Upper Mississippi River, Pool 5, Wisconsin b. 24.1 mm . ··~···_·T1:J~early~juveni~le. ______...Jill "q "8 I ww .. ' ::, 9£ J - 37 preorbital, postorbital, and a weakly defined suborbital teardrop chevron shaped; accumulated melanophores on opercleand·interopercle;cluster·of melanophores.oncleithra; pigment present on pectoral, caudal, andt;pinous and soft dorsal fin rays; black diamond shaped spot present at base of caudal peduncle with a vertical extension forming a hook. Ventral pigment from anus posteriorly to base of caudal. peduncle. Length 21.4 to 27.8 rom (juveniles): cranium pigmentation with a concentration of melanophores covering the cerebellum, and optic lobe; u-shapedpreorbital bar (when viewed dorsally) posterior to maxillary; pigment extending from posterior of optic lobe onto operculum; either 7 or 8 oval shaped cluEJters of melanophores exten.ding from . nape to the base of the caudal pedtiricle (evenly spaced). Head pigmentation with evident preorbital and postorbital chevron shaped teardropsT-and a-weakJ..y.-formed postorbital bar; pigment cl~stered into the form of a triangle on the lower half of op~rculum; and pigment initiating on chin below mandible and posterior of maxillary. Cleithra and margin of operculum with a few scattered melanophores; lateral oval blotches numbering either 7 or 8 situated at mid-body; evident black diamond shaped cluster at base of caudal peduncle extendin~ vertically into hypaxial portion of.caudal fin. Principal rays of caudal fin with scattered melanophores on rays; anal, pectoral, and soft dorsal rays pigmented; pelvic fin wfthoutpigment; spinous dorsal with dense clusters of melanophores at anterior and posterior base, with flecks of melanophores on individual rays. Ventral pigment limited to a few melanophores beneath o~erculum, and a series of ~.".~"~_ ••~ , __ , _ _m•••_ ~, ~ __ single melanophores from insertion of first anal fin spine posterior to . caudal peduncle (Fig. 7). ... 38 Natural History The river darter inhabits the deeper chutes, riffles, of the main stem the Upper Mississippi· River .. ··(Harls'Il.sIldSpeskerI956; Pfli~ger 1975; Smith 1979; Page 1983; Becker 1983; Kuehne and Barbour Simon 1983). Spawning is initiated inth~Vpp~r M~~~~~~ippi~iver from late April to late May during 1983 and 1984 when temperatures C. Eggs 'Wer~~Ilc.ubated in laboratory aquaria maintained at 22C hatched after 144 to 168 hours (X" = 156 ···hours). River darters were seerifn...drift collections when water temperatures approached 11.0 C. Collectionindrift samplesindic.ated a tendency toward surface dispersal in theBI~ck·River, Wisconsin. River darters in the Ohio River were primarily collected at the ...: surface and mid-depths during drift sampling (Simon, unpublished data). Aquar/i\llli. reared river da.rter. larvae were observed actively swiJmning at the surface under daylight conditions. The present study found larval river , I darters only in moderate current, associated with large cobble, and gravel- sand .. substrates. Early·juVeniles were collectedin:-habitatiir-nto'fine ·'sand and gravel in' areas 'with slight to moderate current along main channel border habitats and not in low flow, muck substrate areas. r ,l STRIPED FANTAILDAR'fER, Etheostoma flabellare lineolatum Rafinesque The striped fantail darter, Etheostoma flabellare lineolatum is an inhabitant of small tributaries and streams throughout the Upper Mississippi River basin. The species range extends from southern Quebec across the Great Lakes to the headwaters of the Mfssissi:ppi:-Riverin Wisconsin and Minnesota, southward to northern Arkansas, and eastward throughout the Ohio River basin. It occurs in rivers of the Atlantic slope from the Pee Dee north to the Upper Hudson {Scott and Crossman 1973; Lee 1980; Page 1983; Kuehne and. Barbour 1983). The subspecies E.Llineolatum includes populations in the MississippLRiverbasin.. Jenkins. eta!. (l971) and Clay {1975Y mention a lineolatum type in the Guyandot and Big Sandy Rivers, Kentucky. Information on the taxonomy and natural history of the barred fantail darter, E. flabellare flabellare is available, however, information for the / ...... '.. / striped fantail darter is lacking. The type locality for the fantail _ _ _ __ .__ ~ __ ._.c_.c.·_.. " · _· ··.. · . __ .,_~.~"'_.~~." _. ~~'::-::"._~-::~~:-"'__''''''_'~---::''--: ~.~~.~'._""."'~ darter was the ...... of the Ohio River, near Louisville, Kentucky (Rafinesque 1819). The species currently contains three valid subspecies, however, taxonomic revision of the species is necessary (LeeJ980; Page 1983; Kuehne and Barbour 1983). The life history of the eastern subspecies was studied by Lake (1936), Scott and Crossman (197:3), and Becker (1983) • The reproductive behavior was studied in Michigan (Winn 1958a, 1958b), while the ecology of the species was compared to sympatric darters in (Matthews et a1. 1982). Temperature selection and critical maxima were studied with regard to habitat preference and season (Ingersoll and Claussen 1984). Diet .. was studied by Daiberg (1956), Lake (1936), and Martin (1984). Morgan (1936) investigated the vestigial air bladder. The early life history of the subspecies E. ~ flabellare was studied by Paine 39 40 (1984). Descriptions of the larvae have been investigated by Fish (1932), Cooper (1979), and Auer (1982). The subspecies E. t. lineolatum was a common stream and tributary inhabitant in the Upper Mississippi River basin. Description and comparison of --E. f. lineolatum to the eastern subspecies ~!.:. flabellare will be presented. SpeC1IneIl. collection .. - ..~~..'.~,~. .""~'.=._.~..,,"~,~",~,~,-",,",,,,,,,~,-,,~,-- location from the Upper Mississippi River basin.1s LITERATURE Thestriped·fantail darter, Etheostoma flabellare lineolatum,has an egg morphology typical of other members of the darter subgen'l.ls Catollotus• .. ;': Ova have been investigated for the eastern subspecies .•. E~£labellare .- -~"-~---'_'----'_'_""-,~."---,-,-.- flabellare. In the study done by Winn (1958) egg diameters averaged 2.2 mm; Lake (1936) and Scott and Crossman (1973) reported slightly larger diameters of 2.3 mm. ,Cooper (1979) collected eggs which ranged from 2.5 to 2.9 mm, averaging 2.7 mm in diameter from ElkCreek;--Pennsylvaniaand the Greenbrier······River, .West Virginia• Lake (1936) found three size classes of eggs in E. flabellare flabellare. UPPER MISSISSIPPI RIVER DESCRIPTION Three size classes of ova were observed in the dissected ovary of E. flabellare lineolatum. The smallest ova ranged from 1.0 to 1.5 mm (N = 13; x = 1.3 mm), and were spherical, opaque, and pale yellow. intermediate sizedovacwere.ovoid, ·opaque,and.pale- ¥elloli.--and=rangedfrom 1.5 to 2.0 mm (N = 15; x = 1.6 mm). This suggests that the striped fantail darter is a fractional spawner depositing more than a single cluster of eggs. Mature eggs collected from Spring Coulee Creek, TABLE 7. Material examined of Etheostoma flabellare lineolatum larvae and early juveniles grouped by lmm intervals of total length (N = sample size). rurAL LENG'l1l (ODD) NUMBER DRAINAGE LOCALm 4 5 Mississippi River IIISCONSIN: Coon Creek, Lacrosse County (T IS, R5W" S 32) 5 19 Mi ssissippi River WISCONSIN, Coon Creek. Lacrosse County (T 15N, R 5~. S 32) WISCONSIN' Spring Coulee Creek, Vernon County (T 14~, R'5W, S 5) MINNESOTA' Root River, Fillmore younty (nesr Mystery Csves: T 102N, 1211, S 19) ILLINOIS: Fox River, McHenry County (T 43N, R 9£, S 32) 6 37 ' Mississippi River 'WISCONSIN: Spring Coulee Creek, yernonCounty (T 1411, R5W, S5) WISCONSIN: Coon Creek, Lacrosse County (T 15N,R 511, S32) 7 46 Mississippi River WISCONSIN, Spring Coulee Creek, Vernon County (T 14K, R5W, S 5) 8 37 ~iss~sSippi ~iver WISCONSIN, Coon Creek, Lacrosse Coonty (T 15N,R 5W, S 32) WISCONSIN: Spring Coulee Creek, Vernon County (T 14N, R 5W, S 5) MINNESOTA: Root River, Fillmore County (near Mystery Caves: T 102N, R 1211, S 19) 9 16 Mississippi River WISCONSIN: Spring Coulee Creek, Vernon County (T 14N, R 5W, S 5) MINNESOTA: Root River, Fillmore County (near Myatery Caves, T 102N, R 1211, S19) 10 23 Mississippi River WISCONSIN: Coon Creek, Lacrosae County (T 15N,R 5W, S32) MINNESOTA: Root River, Fillmore County (near Mystery Cavea, T 102N, R 1211, S 19) 11 12 Mi ssissippi River WISCONSIN' Spring Coulee Creek, Vernon County (T 14N, R 5W, S 5) IOWA: Upper lows River, Houston County (T lOON, R 1211, S 21) 12 13 Mississippi River MINNESOTA: Root Riv~r, Fillmore Count:y(near Mystery Caves T 102N, S 19) 13 9 Mississippi River MINNESOTA' Root Riv~r, Fillmore County (nesr Myste~y Caves T 102N, S 19) WISCOSNIN, Rock Riv~r, Rock County (T2N,R 12E, S '28) i 14 5 Mississippi River MINNESOTA' Root Riv~r, Fillmore County (nesr Mystery Csves, T 102N, S 19) 2 Mississippi River IOWA: Upper Io~ River, Houston County (TlooN, R12W, S 21) 15 , 16 2 Mississippi River WISCONSIN' Coon Cre~k, Lacrosse County (T 15N,R 5W, S32) 17 Mississippi River IOWA: Upper I~ River, Houston County (TIOON, R 12W, S21) 18 Mississippi River WISCONSIN: Co~n Cre~k,I Lacrosse '"County (T I5N, R 5W, S32) 19 Mississippi River WISCONSIN: Co()n creJk, Lacrosse <:;ounty (T 15N,II 5W,S32) 20 2 Mississippi River WISCONSIN, sP~tng ~u1ee Creek,Vernon County (T 14N, R 5W, I 21 Mississippi River WISCONSIN, Spriing ~ulee Creek, ,vernon Cointy (T 14N, R5W, 22 Mississippi River WISCONSIN: Coon Cre';k, Lacrosse ,County (T'15N,R 5W, S32) 23 Mississippi River WISCONSIN' Co,on,! cree,V Lacrosse Co,'unty (,T,i 15N,iR 5W,'!,'" S 32) •.•.•. ;1 ••.•. !: .• : •...... i ' t .... 24 Mississippi River IOWA: ~ I-' C:j:,R:::~s::u:::t~oJ:t:5:~::!: :i~:;' S 25 Mississippi Ri ver WISCONSIN, ::r I 26 MississippiRiver WISCONSIN: Spring Coulee Creek,Vernon County (T 14N,R5W, S 5) ... 42 Wisconsin, the Rock River, Illinois, and the Upper Iowa River ,Iowawere all equivalent in diameter and ranged from 2.0 to 2.8mm~(N=lOO;~'i'=2.4 mm). Mature. eggs are.spher:l.c.a1 ,demersal, and adhesive .with translucent yellow yolk. Generally., mature eggs have a single. oil.globule,a narrow perivitelline space, an unscu1ptured chorion,and were unpigmented. Eggs are attached to the undersides of . limestone~slabs__in_the margins and slower portions of river and stream riffles and race~ays (Lake 1936; Winn 1958; Page and Schemske 1978; Cooper 1979; Smith 1979; Page 1983; Kuehne and Barbour 1983; Becker 1983). LARVAE LITERATURE ....: No information is available on the1arvaLontogeny_~oL.E._ilabel1are 1ineo1atum, but excellent accounts of the development 6f E. f1abe11are f1abe11are are in Cooper (1979) and Auer (1982). Additional literature of limited scope is av.ai1ab1e from Greeley (1927), Fish (1932), Lake (1936), and Morgan (1936). Information on the larval development of E. f1abe11are f1abe11are is summarized for comparison. The total length of newly hatched larvae were reported to be 4.75 mm (Cat Run, Ohio: Morgan 1936), 6.2 + 0.4 mm (Elk Creek, Pennsy1v~nia: Cooper 1979), 6.4 mm (Gennesee River, New York: Greeley 1927), 7.0 mm (Black Creek, New York: Lake 1936), with a range from 5.8 to 7.0 mm (Great Lakes basin: Auer 1982). The large yolk sac was 31 %of the TL, and contained a single oil globule (Cooper 1979; Lake 1936). However, Greeley (1927) describes the yolk sac as small. The gills and rudimentaryoperc1es were formed at hatching (Cooper 1979). Total myomeres ranged from 34 to 36 (15 preanal and 19 to 21 postana1). The mode of myomeres were 15 preanal and 21 postana1 (Cooper 1979). 43 r~. Cooper (1979) found fin ray development and notochordal fleXion rom or nine hours after hatching. Lake (1936) found the yolk sac to be entirely absorbed 7 to 10 days after hatching. C()oper(1972) found that the yolk sac was absorbed at 9 to 10 rom. Pelvic fin buds were present just posterior to the oil globule at 8.8 rom. Pelvic fin rays were completely formed at 14.0 ••• Dl11l. Scale ••. formation. ···had····begun··at13.01llJIl1arvae---atthe caudal fin base and extended to the posterior base of the second dorsal fin. In 19.0 rom fish, teeth were present around the margins of both jaws. Scales had formed on the opercles but were lacking on thedors\1m.near the spinous dorsal fin. Segmentation of the principle rays inthemed:ian fins were evident in 18.5 rom juveniles and larger (Cooper 1979; Fish 1932). UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Characteristics of length and depth are described for larval and early ~ juveriile striped fanttaildarter including·thelength·-of--formation for morphological structures {'l'able 8). At lengths 4.5 to 5.9 rom (newly hatched): well developed pectoral fins without incipient fin rays; yolk sac extremely large, spherical (ca. 42 % 'l'L); yolk amber, with a single anterior oil globule; head not deflected over yolk sac; jaws developed; ~8 yolk absorbed (7 ... to iT8:0mm) ; eye diameter oval. First observed rays formed pectoral (5.0 to 5.6 rom); first anal rays formed (5.3 to 5.9 rom); first soft dorsal formed (5.4 to 6.2 rom); spinous dorsal formed (5.9 t.o 6.2 JIlm); incipient dorsac!cfj;nmargin part-ia-ll~,d-ifferentiateU~b.l.cJ:;.QJ~.Omm), completely differentiated (8.0 to 8.3 rom); spinous dorsal fin origin situated over preanal myomere 4, soft dorsal origin over preanal myomere 15 to 16 (7.8 rom); predorsal length 36.3 %of SL (range: 33 to 39.7 % of SL); TABLE 8. Morphometry of Etheostoma flabellare lineolatum larvae and early juveniles grouped by Imm intervals of total length (N '" BC'l.",ple sizp). Mf'an lengths followed by range in parentheses. LENGl'H (_) (a) (b) (c) .!! .IQ!M, ~ PREAIIAL II1lAD .§!!1!!! EYE BODY DEP'l1I GREATEST CAUDAL PEDUIICU: 5 4.7(4.5-4.9) 4.3(4.0-4.5) 2.7(2.6-2.9) .95(.80-1.3) ..•10(.05-.15) .45( .40-.45) .55(.45-.75) 1.9(1.7-2.0) .30( .25-.35) 19 5.6(5.0-5.9) 5.0(4.6-5.4) 3.1(2.8-3.4) 1.0(.80-1.2) .10(.05-.15) .50( .45-.60) .56(.40-.65) 2.1(1.6-2.2) .37(.30-.48) 37 6.7(6.1-6.9) 6.1(5.5-6.3) 3.5(3.2-3.8) 1.3( .95-1.5) .10( .05-.25) .52( .50-.65) .73( .50-.85) 1.8(1.4-2.4) .46( .35-.60) 46 7.4(7.0-7.9) 6.3(5.9-6.7) 3.8(3.4-4.1) 1.5(1.2-1.8) .20( .12-.30) .64( .55-.75) .89(.75-1.1) 1.5(1.2-2.1) .54( .35-.70) 37 ".3(8.0-8.8)' 6.9(6.6-7.4) 4.3(3.9-4.6' 1.9(1.5-2.1) .JO( .20-.40) · 71( .65-.80) 1.0( .85-1.2) 1.4(1.2-1.7) .68( .55-.85) 16 ~.5(9.0-9.9) 7.9(7.4-8.2) 4.8(4.5-5.1) 2.3(1.~2.9) .40( .30-.50) •77(.75-.85) 1.2(1.1-1.4) 1.7(1.4-2.0) .82( .65-1.0) i 23 1q.4(10.0-10.9) 8.6(8.1-9.0 5.3(4.9-5.6) 2.5(2.0-2.8) .40( .40-.50) .82(.70-.90) 1.4(1.1-1.7) 2.1(1.7-2.2) .97(.80-1.2) 12 1l•. 5(11.0-11.9) 9.5(9.0-9.9) 5.9(5.7-6.3) 2.8(2.6-3.0) .50( .40-.65) .87(.75-1.0) 1.5(1.2-1.7) 2.1(1.7-2.5) 1.1(.90-1.3) 13 1Z.4(12.0-12.8) 10.3(9.9-10.7) 6.3(6.1-6.8) 3.1(2.8-3.4) .62( .45-.73) •91( .80-1.0) 1.8(1.5-1.9) 2.4(2.3-2.6) 1.3(1.1-1.4) 9 13.4(13.0-13.9) 1l.0(10.7-11.4) 6.8(6.6-7.0) 3.3(3.0-3.6) .64( .50-.75) •97(.90-1.0) 1.8(1.6-2.0) 2.6(2.4-2.9) 1.3(1.1-1.5) 5 14.4(14.2-14.5) 12.0(11.8-12.1) 7.2(6.8-7.5) 3.6(3.5-3.7) .68( .60-.74) 1.0(1.0-1.1) 2.0(1.8-2.1) 2.7(2.2-2.8) 1.5(1.3-1.6) 2 151.9(15.8-15.9) 13.2(13.2-13.3) 7.9(7.8-7.9) 3.6(3.4-3.7) . 70( .68-.7(J) .85( .84-.86) 2.3(2.2-2.4) 2 161. 1(16.0-16. ~) 13.5(13.5-13.5) 8.0(7.9-8.0) 3.7(3.5-3.8) •72(.71-.13)· .~8( .86-.90) 2.3(2.2-2.3) 17.2 14.2 8.6 3.9 .81 .88 2.8 18.8 16.0 9.6 4.6 .89 1.1 2.6 3.2 19.1 16.3 9.5 4.3i .86 1.1 2.7 2 2~f 7(20.5-20.8) 17.5(17.2-17.7) 10~4(10.3-10.4) 5.2(5.2~5.3) 1.0(1.0-1.1) 1.~(1.1-1.2) 3.1(3.1-3.2) 21.6 18.6 11.1 5.11 1.0 1.2 3.1 3.6 22.2 18.7 11.3 5.41 1.1 i 1.3 3.0 3.7 23.5 19.7 11.7 5.71 1.0 1.4 3.1 3.8 i 24.6 22.7 12.5 6.01 1.2 1.5 3.1 3.6 I 25.9 22.2 13.2 6.31 1.4 1.5 3.6 4.2 26.1 22.0 12.7 6.31 1.4 1.2 4.0 4.7 il i i , 'I i II .,'...... , i I "i, 'i ••• ,'. ,i' i •• i' I i (a) iry depth measured VertiC~llY.from the doraa1 to ventral aUrfaCe at poaterior. portr of anua tl=>o Greate~t measure,dverti~allJ f~R'" tl=>o (b) body depth doraal to ventr"l surface including thel yolk ssc ;' •....•. ..;.>' ..><'. ,. ,j . ! (c) <;,juda1 peduncle depth measured vertically ,from the dorsal to. ventral surface at penult1l118te ... 45 notochord flexion (4.9 to 6.0 mm), after first caudal fin ray formation (5.3 to 5.5 mm): caudal fin rounded (5.8 to 6.2 mm): ~~ncipientana1 fin margin partially differentiated (7.1 mm), completely differentiated by 7.9 mm: pelvic fin buds formed anterior to dorsal fin origin prior to complete absorptio11 of yolk sac (7.9 to 8.5 mm); first pelvic fintays formed (8.8 to 9.2 mm). No swim bladder formed, remains rudimentar~y~;~.~~.gut~t.~a;lght; entire finfo1d absorbed (8.2 mm). Supraorbital, infraorl>ital,and lateral head canals formed (11.0 to 12.6 mm): preoperculomandibular canals formed (10.8 to 11.8 mm), preoperculomandibu1ar pores 10, completely formed (12.6 to 13.8 mm); supraorbital completely formed (11.7 mm). Infraorbital canal formed wi.tl18 ... pores (10.8 mm), formed with retrogression to interupted condition of 4 pores anteriorly and 2 pores posteriorly (14.4 to 14.6 mm) : lateral line began forming14.0tolA.2mm;~_scale_s_pr-.elient. at 10.0 mm, squamation completed (l4.7 JlUJl}: no scales are preseritoll the nape, cheek, opercle, breast, and prepectoral areas. Meristics I Striped fantail darter myomere number constant posthatching, preanal myomere 15, postanal. myomere 19 to 21 with 34 to 36totalmyomeres (postanal myomere number: x= 19.5; N = 155). Total vertebrae numbering 33 to 34 (N = 5, x = 33.8), including one urostylar element ,from cleared and stained specimens from Spring Coulee Creek, VernonCoU11ty,Wisconsin, and the Root River, Fillmore County, Minnesota. Scales in the lateral series ranging from 42 to 57~N = 13, x= 47.7, mode = 48). Paired and median fin ray values an4 length at appearance are presented in Table 9. Pigmentation Newly hatched larvae (4.5 to 5.9 mm): scattered melanophores on a ... 46 TABLE 9. Selected meristic values and size (mm total lengtb.)at·Elie apparent onset of developmerit.fofEtheostomaflabellare lineolatum.Mean values are underscored. Rare or question... able extremes are in parentheses. .The number DotsalFin Spines / Rays VIII - IX / 12-13-14 first rays formed 5 adult complement formed (1-7) II / 9-.!Q:l! II / 8-10-11-13 Anal Fin Spines / Rays (8-9) 5.3 - 5.9 .2 - 7.5 first rays formed (8-9) - 11.5 adult complement formed ix-xiv, 8~+ 7, viii-xiv no informaH.on Caudal Fin Rays (8::,;9) 5.3 - 5;5 --~-7-.-2----1.5------.. first rays formed (8-9) 7.2 -7.5 adult complement formed 7.0 - 7.5 (1~3, 7) 12 - 13 11-12-13 Pectoral Fin Rays (87"9) 7.2 - .5 first rays formed 5.0 - 5.6 adult complement formed (1 ...7) 41~-60 Lateral Series - Scales 42 - 48 - 57 (1 ..3,7-10) 34-36/33-34 34-36/32..34-35 Myomere / Vertebrae (8..9) 15 preanal myomeres 15 postana1 myomeres 19· ... 21 8.1. PagE! 1983 2. Kuehne and Barbour 1983 3. Becker 19834. Forbes and Richardson 1920 5. Jordan and Evermann 1894 6. Trautman 1981 7. Scott and Crossman 1973 8. Cooper 1979 9. Auer 1982 10. Bailey and Gosline 1955 r! 47 large yolk sac with greatest concentration laterally.. Stellate melanophores are present on cranium encircling optic lobe. melanophores laterally, rising obliquely near nape. Lateral pigmentation with a midlateral stripe from the yolk sac posterior to base of caudal peduncle. Ventral melanophores present on gut and from anus to approximately postanal myomere 9. Dorsal melanophoresin anterior of anus and initiating near posterior of .. ventral ...... postanal··· melanophores. Majority of preanal myomeres without pigJIl€!Ilt.ation (Fig. 8). Length 6.0 to 7.5 mm: cranium with a concentration of melariophores dorsally encircli.ng optic lobe. Postorbital bar formed, with additional horizontal pigment on operculum. Yolk sac with stellate melanophores on lower half. Dorsally, pigJIlentation on nape and at base of soft dorsal. Lateral pigJIlentation outlining preanal myomeres posteriad of yolk sac, extending to the middle of soft dorsal. A midlateral stripe formed from single melanphores at apex of preanal myomeres poster-ior··--oi· yolk sac. Ventrally, stellatemelanophores present at almost every postanal myoseptum with several extending dorsally to midlateral. Melanophores present at base of caudal fin (Fig. 9). Length 7.8 to 9.2 mm: optic lobe outlined wi.th. DlE!1.a,nophores. Horizorital preorbital and ,postorbital bars with additional pi.gmerit present on operculum. Additiortal cranial pigJIlent present dorsally on nape. Melanophores outlining lateral myosepta of preanal and postanal myomeres just posterior to softcdorsal. Three mtdJst:era.1. cluster~.9f _.m_€!~aI1c:>phores situated beneath the soft dorsal. Melanophores extending onto rays of caudal, anal, and soft dorsal. Ventral pigmentation concentrated at midventral of gut, alld under the operculum and branchiostegal rays (Fig. FIG. 8 Etheostoma flabellare lineolatUttl, ·striped·fantail darter (newly hatched larva), 4.8 mID TL, Coon Creek, ~~-~~Wrscons:i:n---a~;~~dorsal b. lateralc. ventral. ':) i ./ i ;fI;' •• WWJ. ~q '8 8t' - FIG. 9 ",': Etheostoma flabellare lineolatum, .. striped fantail darter larva Spring Coulee Creek, Wisconsin a. 7.1 min . .... ···--TI:;-b";-~&.-2·-mm-.'J.rh. l I j JI ww," "e ww," 6v zC 50 r~ 9) • Length 9.5 to 10.9 mm: horizontal preorb:1t:I:i:tI:iIlc:1.PQ~t:QI'pitalbars present on cranium. Chevron shaped blotches present dorsal~anterioriad to orbit, and on the optic lobe. Dorsally, 8 rectangular blotches extending from nape to base of caudal peduncle. Oval blotches becoming continuous anteriorly along midlateral, with sCI:it.t.erec:1.Ill~:l.I:iIlQPhQI'~:~~=~'!~t'~\lltiple melanophoresoutlining preanal and ·postanal myomereshypaxially and epaxially ~ Melanophores extending onto caudal and soft ·.dorsal fin. Ventral pigmentation limited to 5 areas of concentration from just after the anus to the base of the caudal peduncle. Spinous dorsal, pectoral, pelvic, and anal fins devoid of pigment (Fig. 10). Length 11.0,' to 13.9 mm: cranium with horizontal preorbital and postorbital bars formed, with an obJ..i..q\l~ PI3,I'~~:lCt:~!l:~il1:g~ t:.0wards nape. Cerebrum, and optic lobe with clustered melanophores blanketing them. Dorsally, 9 to 10 rectangular blotches with obliquely scattered melanophores connecting 12 to 13 rectangular midlateral blotches. Lateral epiaxial with scattered melanophores connecting 12 to 13 rectangular midlateral blotches. Lateral epiaxial with scattered melanophores outlining scales. Pectoral girdle with a blotch near cleithra. I Lepidotrichif,i of spinous and soft dorsal, anal, and base of caudal fins with melanophores. Mandible and maxillary, and interoperclewi£h scattered melanophores. Pectoral and pelvic fins without pigment. Length 14.5 to 24.6 mm (juveniles): cranium with concentrated melanophores over optic lobe and cerebrum. Distinct preorbital and postorbital bars formed, no suborbital teardrop forme-a .'A Clievronshaped cluster of melanophores parallel to postorbital bar, scattered melano- ~ phores on cheek. Lateral pigmentation with 11 to 13 rectangular blotches, connecting 8 dorsal bands. The last rectangular blotch may be divided to FIG. 10 ... ~ Etheostoma flabellare lineolatum; . striped fantail darter, Root River, Minnesota a. 10.3 rom TL larva ··········b~--l4.4-mm-'I'Learly····juvenile• 'q I ww .. ' IS r! ~ 52 form 2 spots near midlateral of caudalpedrincle base. Horizontal lines of melanophor~s extend from the head to caudal peduncle, formed from individual melanophores on outer margin of scales. Distincth1.11rieral spot formed posterior of opercular spine. Spinous dorsal, pectoral, and anal fins with pigmentation scattered on rays. Horizontal stripes 4 or 5 on soft dorsal distributed on pterigiophore. Caudal fin with 6 to 8 vertical stripes formed on the interstitial membranes. Pelvic fin without pigmentation' (Fig. 10). Natural History The striped fantail darter inhabi.ts the raceways and riffles ofsmall to medium gradient streams and rivers usually associated with gravel and limestone slab substrates (Harlan and ,Speaker 1956; Eddy and Underhill 1974; Pflieger 1975; Smith 1979; Page 1983; Kuehne and Barbour 1983; Becker 1983). Spawning initiates in early May and continues through June at temperatures between 13 to 17 C in Wisconsinand-Minnesota,---but--continues untilmid.;;.;July·whentemperatures approached 25 C in Iowa and Illinois. Aquarium observations of striped fantail darter larvae indicate they are demersal seldomly entering the water column. Larvae were firs.t.collected I in Coon Creek, Wisconsin at water temperatures ofI2.0C.Newlyhatched larvae and larvae still bearing yolk were found in stagnant margins of streams. At lengths between 10 mm to 14 mm, larvae and early juveniles appear to move. out from shoreline margins into slow current velocities. is not until 24 mmthat juveniles were=£ound __in I..acg\V'8y~~h:lk~cc11l11:13.. are found in the faster riffles. Egg development increased rapidly with elevated water temperatures. Eggs incubated at 20 C hatched after 240 ± 18 hours (9.25 to 10.75 days), 53 while at 23 C hatching occurred in 10 hours (5.50 to 6.50 days). Lake (1936) reported eggs of E. flabel1are -flabellare hatched in 30 to 35 days at 17 to 20 C; after 21 days, at 21 to 22 C; and in 14 to 16 days at STRIPETAIL DARTER, Etheostoma kennicotti (Putnam) The stripetai1 darter, Etheostoma kennicotti is an inhabitant of extreme southern Illinois in the Upper Mississippi River basin. The species ranges from southern Illinois ibutaries of the Ohio River through northwest Kentucky into the Tennessee, Upper Cumberland, and Green River drainag~f3()J: the states of Kentucky, Tennessee, Georgia,.Alabama, and Mississippi (Clay 1975; Smith 1979; Page 1980; Page 1983; Kuehne and Barbour 1983). Some detailed information presently exists on the taxonomy and life history of the stripetai1 darter. The stripetai1 darter was originally '; described from a rocky brook in Union County, Illinois (Putnam 1863) however the l~ttotype was invalidated by Smith and Page (1975). The stripetai1 darter is a member of the darter subgenus Catonotus (Page 1981, 1983; Kuehne' and Barbour 1983). Systematics and geographic variation were investigated by Page and Smith (1976) • The diet-of--th:e-stripetai-l darter includes tmmature,'aquaticinsects andmicrocrusteacens (Forbes 1920; Page 1975). Page (l975) investigated the life history of the stripetail darter in Big Creek, Illinois which included limited information on the early life history of the species. Specimens examined for the, present study were collected from the lower portion of the Upper Mississippi River basin with supplemental material from Tennessee (Table 10). ) 54 TABLE 10. Material examined of Etheostoma kennicotti larvae and early juveniles grouped by lmm intervals of total length (N = sample size). roUL LEIIGTII (m) ~ ~ 2 Ohio River ILLINOIS: Big Creek, Hardin County I(T 33N, R SE, SilO) "5 2 Ohio River ILLINOIS: Big Creek, Hardin County ,(T 33N, RilE, S ~O) 6 Ohio River ILLINOIS: Big Creek, Hardin' County '(T 33N, RilE, S 10) 7 Ohio River ILLINOIS: Big Creek" Hl'rdin County(T 33N, RilE, S 10) 12 4 Ohio River i ILLINOIS: Big Creek, Hardin County(T 33N,R liE, S 10) 13 4 2 Ohio River ILLINOIS: Big Creek, Hardin County (T ILLINOIS: Hogthief Creek, Hardin County 3 Ohio River ILLINOIS: Big C"eek, Hardin County (T 33N" R Ohio River ILLINOIS: Big C~eek, Hardin County (T 33N,R Ohio River ILLINOIS: Ohio River ILLINOIS: Hogthlef Creek, Hardin I Ohio River ILLINOIS: Hogth~ef Creek, Hardin TENNESSEE: Indian Creek, Claiborne 88845) I II' Ohio River TENNESSEE: Indian Creek; Claiborne 88845) TENNESSEE: Robin~on Creek, Hardin U1 U1 Ir ,t 56 ~ LITERATURE Thestripetail darter, Etheostoma kennicotti, has a spherics} egg. Eggs were reported to be translucent, averaged 2.1 mm in diameter, and contained a single oil droplet .(Page1915,1983). UPPER MISSISSIPPI RIVER DESCRIPTION Eggs from Big Creek, Hardin County, Illinois ranged .from .1.9 mm to 2.5 mm (N = 32, i = 2.1 mm). Mature eggs of the stripetaildarter were spherical, demersal, and adhesive. Mature eggs contained translucent yolk, had a single oil globule, a narrow perivitelline space, an unsculptured chorion, and were unpigmented. Eggs ·--wereattached~~tothe ...unde:(sides of slab rocks in shallow upstream pools (Page 1975a, 1975b, 1983; Page and S~hemske 1978; Smith 1979). LARVAE~· LITERATURE Limited early life history information is presently available for the stripetail darter. Page (1975) reported newly Squamationwas nearly~ccomplet;;eat~1~k1.ngonlyOIl :the-~bell~Land_the anterior dorsum. At 15 mID, squamation was nearly complete. The lateral line canal began forming at 15 mm and at--18Dilii -extendedt() -beneath the first dorsal fin. Page (1975b) included line drawings of larva measuring 57 5.7 and 7.2 rom TL. UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Characteristics of length and depth are described for larval and early juvenile stripetail darter including length at formation of morpholbg1ca1 11). At lengths 4.1 to 4.7 rom (newly hatcheq): well developed pectoral fins with 12 incipient rays; yolk sac extremely large, spherical (ca. 31.9 %of SL, 30.0 % of TL); yolk -orange, 'With a single anterior oil globule; head appears to be deflected over yolk sac due to its robustness; yolk absorbed (7.5 mm); eyes circular~ First rays formed, pectoral and spinous dorsal simultaneously (5.1 t05.2mm):first soft dorsal rays formed (ca. 5.1 mm): incipient dorsal fin margin partially diff~rentiated (6.9 rom), completely differentiated (7.5 mm): spinous dorsal fin origin situated over preanal myomere 6, soft dorsal origin over preanal myomere 15 to 16 (6.9-to 7.5 mm); predorsa11:ength~3s.-.-2.-c.pe);cent----of SL (range: 32.5 to 43.8%SL};notochord flexion (ca. 6.9 mm), preceeding first caudal fin ray formation (ca. 7.5 rom): caudal fin -r'QuIl.ded 7.5 rom: first anal fin rays formed (6.9 rom): incipient anal finmt:lrgip.pt:irtia11y differentiated (6.9 rom), completely dif£erentit:lteq{'7!~IlI1llJ:p~±vic buds formedaIlterior to dorsal fin origin succeeding complete yolk absorption (ca. 7.5 rom): first pelvic fin rays formed «12.1 rom). Noi swim bladder formed: gut straight: entire finfo1d absorbed (7.5mm). Supraorbital, infraorbital, and lateral head-canals~formed~.~_(lf.l...t~==1~!2 rom); infraorbital canal complete with 8 pores extending to mid~orbit (12.7 rom): completely formed with retrogression to interupted condition of 1 pore posterior and 4 pores anterior (25.5 to 27.5 rom). Preopercu1omandibu1ar TABLE 11. Morphometry of Etheostoma kennicotti larvae and early juveniles grouped by Imm intervals of total length (N = sample size). Mean lengths followed by range in parentheses. tF1lG'11l (aa) \c::: (a) (b) (e) ! TOTAL STANDARD PREANAL HEAD SNOUT ~ BODY DEP'l1I GREATEST CAUDAl. ,PEDtlNctE 2 4.4(4.1-4.7) 4.0(3.5-4.5) 2.4(2.0-2.7) .84( .80-.90) .07( .05-.10) .41( .40-.42). .37( .37-.37) 1.05(1.0-1.1) .19( .18-.20) 2 5.1(5.1-5.2) 4.9(4.9-4.9) 2.5(2.5-2.6) .84( .80-.88) .09(.08-.10) .43( .40-.46) .41( .38-.44) 1.30(1.2-1.4) .25( .24-.26) 6.9 6.2 3.5 1.4 .10 ,60 .82 1.4 .38 2 1 7 •3(7.1-7.5) 6.3(6.2-6.4) 3.6(~.4-3.8~ 1.6(1.3-1.8) .22(.15-.28) .63( .60-.65) .72( .60-.86) 1.3(1.2-1.4) .47( .45-.50) 4 ~2.4(12.1-12.7) 10.4(9.9-11.0) 6.5(~.2-6.9) 3.~(3.1-3.6) .66( .60-.72) 1.0(.92-1.1) 1.8(1.6-2.3) 2;2(2.1-2.3) 1.1(1.1-1.2) 4 ~3. 7(13.1-13.9) II.7(ll.I-12.1) 7.3(7.0-7.5) 3.8(3.7-3.9) .66( .60-.80) 1.l(.9O-1.1) 2.0(1.8-2.3) 2'5(2.3-2.6) 1.2(1.0-1.4) 3 ~4.2(14.1-14 .4) II.7(ll.6-11.7) 7.2(7.0-7.4) 3.8(3.7-4.0) •73( .60-.80) 1.2(1.1-1.2) 1.9(1.7-2.1) 2.5(2.4-2.5) 1.2(1.0-1.3) 3 ~5.3(15.0-15.8) 12.7(12.5-12.9) 7.8(7. 7~8.0) 4.2(4.0-4.4) .81 1.2(1.1-1.2) 2.1(2.1-2.2) 2.8 1.4(1.1-1.5) 2 16.8(16.7-16.9) 14 .O( 13.8-14.1) 8.5(8.4,-8.6) 4.5(4.5-4.6) .86( .80-.90) 1.2(1.1-1.3) 2.3 2.9 1.4(1.4-1.5) 5 P .5(17.3-17.9) 14.7(14.4-15.1 ) 8.9(8.8-9.2) 4.7(4.4-4.8) .93( .80-1.0) 1.3(1.1-1.4) 2.3(2.1-2.5) 3.1(2.9-3.2) 1.4(1.4-1.5) 2 .8.5(18.0-18.9) 15.3(15.0-15.7) 9.4(9.2':9.6) 5.0(4.8-5.1) 1.0( .90-1.1) 1.4 2.6(2.5-2.6) 3.2 1.6(1.5-1.6) i ! 19.3 16.2 9.8 5.3 .92, 1.5 2.5 3.2 1.4 9 ,!0.4(20.0-20.9) 17.2(16.1-17.9) 10.4(10.3-10.8) 5.4(5.2-5.8) 1.1(1.0-1.2} 1.5(1.4-1.5» 2.9(2.6-3.1) 3.5(3.3-3.9) 1.7(1.6-2.0) 5.8(5.6-5.9) 1.2(1.1-1.3) 1.5 3.1!(3.0-3.1) 3.7(3.7-3.8) 2.0(1.8-2.1) 2 i.' 1.2(21.0-21.3) 17.7(17.5-17.9) 11.2(10.6-11.9) 3 i 2.3(22.1-22.5) 18.8(18.6-19.0) II.3(10.6-11.9) 5.9(5.8-6.0) 1.2(1.1-1.2) 1.6 3.3(3.1-3.3) 3.9(3.7-4.4) 2.0(1.1-2.1) 23.3 19.6 12.0 !6.1 1.2 1.7 3.5 4.5 2.1 . 25.5 21.1 13.0 16.5 1.4 1.7 3.4 4.4 2.2 ! , Il'·~"·"i!~.") 22.4(22.3-22.6) 13;6(13.4-13.7) 6A6.3-6.8) 1.5(1.3-1.7) 1.9(1.8-2.1) 4.1;(4.0-4.1) 4.6 2.1(2.0-2.1) 3 17.6(27.5-27.8) 23.1(23.1-23.2) 14;0(13.5-14.4) 7.3~7.0-7.5) 1.4(1.3-1.5~ 2.0(1.8-2.1) 4.0(3.7-4.3) 4.5(4.5-4.6) 2.2(2.2-2.3) I 2.2(2.2-2.2) 2 18.6(28.3-28.9) 24.0(23.8-24.2) 14)2(14.1-14.4) 6.9~6.8-7 .0) 1.3(1.2-1. 5P 2.0(2.0-2.0) 3.S(3.5-3.6) 4.4(4.2-4.6) II . I (a) IrOdY depth measured vertically (b) :Greatest body depth measured vertically (c) iCaudal depth measured vertically OJ 00 r canals formed (12.1 to 12.6 rom); I completely formed (14.2 to 15.8 mm); supraor'l>:i'taJC:()JIIE~~1:~~Y:~():rJlledwith 3 pores (l7.3 mm);lateral line began forming·· (ca•.. 14.4mm); scales present I at 13.1 DlDl; squamation complete (ca. 15.0mm); no scales are present on the I nape, cheeks, opercle, and prepectora1 areas. I Meristics Pteanal myomeres numbered 16, posta.nalmyomeres 18 to 19,With34 to 35 total myomeres (postana1 number: i"" 18.5; N = 15) ~ Total vertebrae count 34.~)to 35 (N = 3, i = 34.3), including one urostyla:relement .from cleared and stained specimens from Big Creek, Hardin County, Illinois. Scales in the lateral series ranging from 34 to 51 (N = 43, x=41.9, mode = 41). Paired and median fin ray values and length at appearance are presented in Table 12. Pigmentation Length 4.1 to 4.7 mm (newly hatched): absence of pigment over majority of body; several me1anophores located on gut between posterior yolk sac and I anus; ventrally yolk sac with a concentration of me1anophores Length 5.1 to 5.2 mm: melanophores dorsally on gut ·between •. yolk sac and anus: with me1anophores ventrally on postanal lIIios~pt.a7to 13 near area of future anal fin, and scattered me1anophores distributed profusely on yolk sac. Dorsally, several melanophores on cranium outlining anterior portion of optic lobe. Length 6.9 to 7.5 mm: cranium with s~verai~meTa.nophoiesanterior and posterior to orbit; dorsally, melanophores outline optic lobe with the majority forming a straight line separating the right lobe from the left; several melanophores along the cerebellum. Laterally, pigment on ventral 60 TABLE 12. Selected meristic values and size (mm total length) at the apparent onset of development forEtheostomakenriicotti~ Mean values are underscored. Rare or questionable extremesare·in parentheses. 'l'he·numberof secondary rays of the median fins are in lowercase Roman numerals. (8) Attribute / Evpnt Pre",ent Study Literature (1-5,9) Dorsal Fi.11 Spines / Rays VII-VIII / 11-12 VI~VII~lX/ 10-~-13 first rays formed 5.0-5.2 arlult·compJement formed 12.1 ,9) Anal Fin Spines / Rays II / 6-2. first rays formed 6.9 adult ..~ (9) xi-xii, 8 + 6-2., iv-xii 12-14-15-16 Caudal Fin Rays (6J ~_ ..~ ._~- -"""-~"-7--;-2-~~~·"" first rays formed 5.1 adu~t complement formed 7.5 no information (1-3,9) 12-13 11-12-13-14 Pectoral Fin Rays ---'"'(6) first rays formed 5.1 7.2 adult complement formed 6.9 no (1-5,7,9) 38-~-53 Lateral Series - Scales 34 - .ll- 51 (1-2,7-8) information/32~33-34-35 Myomere / Vertebrae 34-35 / 34-35 no preanal myomeres 16 no information postanal myomeres 18 ~ 19 no information a.1. Page 1983 2. Kuehne and Barbour 1983 3. Braasch and Page 19794. Smith 1979 5. Clay 1975 6 .. Page 1975 7. Forbes and Richardson 1920 8. Railey and Gosline 1955 9. Page and Smith 1976 FIG. 11 Etheostoma kennicotti, stripetail darter (newly hatched larva) 4.3 rom TL, Big Creek, Illinois a. dorsal ,,~,"",--b'w~la"tex:aL~-"c."ventral. ·0 ...... - .q ww" ·8 19 62 postanal myosepta 7, 8, and 13 over differentiatirigsrisl:f:t.ri. V~Iltfally, a concentration of melanophores forming a large spot at base ofisthl'llus; scattered melanophores ventrally covering remainder of yolk sac .(Fig•. 12). Length 7.5 mm: formation on cranium of preorbital and postorbital bars; pigment on future opercle; dorsally, an increase in melanophores covering optic lobe. Dorsally 011 the bodyarescattered-melanopb;o-rlrs-from just anterior of spinous dorsal to posterior of sof~ dorsal. Ventrally, large cluster of melanophores at base of isthl'llus, becomes site of pelvic fin bud; few melanophores on gut between pelvic fin and anus. Laterally, pigment present on preanal myomere 11 and 12 above gut;melsriophores present on postanal myosepta 5 through 11, 13 to 15, andl7 (F'ig.12). Length 12.1~to 12.7 mm: extension of preorbital and suborbital bars; melanophores extending from optic lobe formiIlg--a-c:h~vron-----po-sterIor to .,.j,.) orbit. Dorsally, optic lobe covered with melanophores; pigment extending posterior forming the first dorsal saddle; 7 additional saddles forming. Laterally, 14 areas of concentrated melanopho:r-es_t:oKm:t.I!g._plQtf:h~_s_~ At base of caudal peduncle a stripe is forming. Ventrally, from anus to caudal, are 5 areas of melanophore concentration (Fig. 13). Length 13.1 to 14.4 mm: melanophores forming in ventraiportioll of opercle; preorbital, postorbital, and suborbital bars present; optic lobe entirely covered with melanophores extending over·cerebralor.olfactory lobe; suborbital bar extending ventrally with several melanophores posterior to mandible. Dorsally, 8 to 9 saddles extending from nape to caudal fin. Laterally, 12 to 14 blotches with myomeres outlined with ._- -- melanophores; prepectoral area with melanophores; distinct stripe forming near caudal peduncle base. Pigment presentonpecto:r-al,c811c:lal, soft and spinous dorsal, and anal fin rays. Ventrally, from anus to hypural plate are 5 to 6 rectangular blotches. No major changes in pigmentation occur ·.... _..._.=....=.. ======.. ~.=....~....=...=....=.~- FIG. 12 "",' Etheostoma. kennicotti, stripetaildar'ter larva, Big Creek, Illinois a. 6.9 rom TL b. 7.5 mm TL. 0q wwa. wwa. °8 £9 .. FIG. 13 ..: Etheostoma kennicotti, stripetail darter, Big Creek~ _~ ~_n1inois__a .• 12.1 mm TL larva . b.21.3 mm TL early juvenile. r--t ww a. "8 \79 ______J 65 until later juvenile stages. Length 15.0 to 15.8 mm: essentially·· the same as previous description with the exception. of. scales outlined withmelanophore~. Length 16.7 to 18.3 mm (juveniles): pigment outlining scale edges connecting 12 to 14 rectangular midlateral blotches to 7 to 8 dorsal saddles and 5 to 6 ventral saddles;dorsal~porti()n-~()£~~~e.l"an~u.mwith melanophores over optic lobe extending anteriorly between orbit; preorbital, postorbital, and suborbital· bars well developed; opercle, interopercle, mandible,and maxillary with pigment outlining structures. Pectoral fin with melanophores on rays; 2 oblique Stripes in spinous and 5 oblique stripes in soft dorsal; 4 vertical stripes in caudal fin; anal and pelvic fins witnbut pigment. Distinct humeral spot present; prepectoral area with melanophores. Ventrally, melanophores·~~~concent-Fated at pterigiophore interdigitation with ledidotrichia of anal fin (Fig. 13). Natural History The stripetail darter inhabits the shallow, slab rock pools of high gradient streams (Page 1975b; Clay 1975; Page and Schemske 1978; Smith 1979; Page 1983). In the lower portion of the Upper Mississippi River basin spawning stripetail darters were found from early April until late May at temperatures bet.",een 14 to 20 C (Page 1975b). AquariumobserYations indicated that stripetail darter larvae remain closely associated with the substrate and are primarily demersal (L. M. Page, personal communic.atloll). No distinction is evident in the preferred habitat usage between larval early juvenile, and adult E. kennicotti. Larval forms occupy non-slab' pools to a greater extent throughout theY~f.lI'f.lIl~ are found around the margins of streams to a greater extent than adults (L. M. Page, personal communication). During the months of September through December juveniles 66 are fOWld on gravel riffles, however, the preferrred habitat is slab pools (Page 1975b). Eggs incubated in laboratory aquaria at 19 to 23 C hatched after 145 to 150 hours (Page 1975b) • ...: r~. LEAST DARTER Etheostoma microperca Jordan and Gilbert The least darter, . Etheostoma microperea is a watch EJpe~ies in Wisconsin (Les 1979; Becker 1983), endangered and possibly extirpated in Iowa (Roosa 1977), and is on watch status in Minnesota (Moyle 1975). An endangered species is defined as any species·or~subspecies- in-danger of becoming extinct, while a watch species is any species or subspecies under special observation to identify conditions that could calise further decline (Fago 1983). The range of the species extends drainage to northern Minnesota south with disjunct populations occurring in the Ozark uplands and Blue River, Oklahoma (Leee't a1. 1980). The species is declining due to habitat changes~Ccaused by turbidity and increased·industFlal·and-·-agr4sultural pollution (Becker 1983). In.formation regarding the adult taxonomy and natural history of the least ~artet is available, however little presently exists on the early life history. The least darter was first described from the Oconomowoc River at Lac La Belle, Waukesha County, Wisconsin (Jordan and Gilbert 1887). The species has been assigned to the darter subgenusMicroperca (Burr 1978; Kuehne and Barbour 1983), while Page (1981, 1983) has combined the subgenus with Boleichthys. The systematics of theM!croptirsa.subgenus including the least darter were investigated by Burr (1978). The reproductive behavior and biology of the species was studied by Petravici (1936), and Winn (1958a, 1958b). Diet and feeding chronology was studied ","" _.. ~-"-_._" - - _. - '._.. ~.~"~_ ,-" by Forbes and Richardson (1920), and Cordes and· ··Page (1980) in Illinois.' Burr and Page (1979) studied the life history in the Iroquois River, Illinois. Limited information is available on the species early life history (Petravicz 1936; Burr and Page 1979; Auer 1982). Larval and early 67 68 juvenile specimens utilized for meristic,morphometrfc, and pigmentary characteristics in the resent study were collected from the .. Upper Mississippi River basin (Table 13). EGGS LITERATURE The least darter, E. microperca,sh8resa similar egg morphology with otheI-JIieJIibe:rs of the subgenus Microperca. Mature ova, including the primary egg envelopes, a.redeeply indented on one side (Burr and Page 1979). Egg diameters ranged from O.Sto 1 .. 1 mm (Burr and Page 1979; Winn 1958b). ~Petravicz (1936) reported diameters of 0.7 mm. UPPER MISSISSIPPI RIVER DESCRIPTION In the present study iI1UIl8ture ova, obtained by ovary dissection of least dar~er.s from the Iroquois River, Illinois ranged from 0.3 to 0.5 mm diameter (N = 97; x = 0.4 mm). This suggests that the least darter is a fractional spawner much like the tesselated darter, E. olmstedi (Gale and Deutsch 1985) depositing more than a single clutch of eggs during a single spawning season. Mature eggs were spherical and without oil globules. Mature eggs from Blue Lake, Pool 8, Minnesota (a backwater area of the Upper Mississippi River); Iroquois River, Iroquois County, Illinois; and from the Piscasaw River, McHenry County, Illinois had egg diameters from 0.7 to 1.0 mm (N = 68; x= 0.8 mm). Eggs after the act of spawning, are demersal, adhesive, have a translucent yolk, a single oil globule, a narrow perivitelline space, an unsculptured chorion, and are unpigmented. Eggs were attached to leaves, and stems of aquatic macrophytes along grassy stream margins (Burr and Page 1979; Petravicz 1936; Winn 1958a, 1958b; present study). TABLE 13. Etheostom~ Material examined of microperca larvae and early juveniles grouped by 1mm intervals of total length (N = sample size). roUL LENG'11I ( ) ... NUMBER ~ ~ 3 2 Mississippi River ILLINOIS: Pisca98V River, McHenry County, 1 ad SW Harvsrd (T 45N, R 5R, S 6) ILLINOIS: unnalled trib. Iroquois River, .Iroquois County, 6 km HE Watseka 4 3 Mississippi River ILLINOIS: unnamed trib. Iroquois River, Iroquois County, 6 km HE Watseka ILLINOIS: Piscasav River, McHenry County, 1 mi SW Harvard (T 45N, R 5E, S 6) MINNESOTA: Blue Lake, Pool 8, near La Cresent at HWY 61 (R.M. 698.5) 6 Mississippi River ILLINOIS: unnamed trib•. ~oq,\ois Riv"r, Iroquoi~ County, 6km HE Watseka 7 6 Mississippi River ILLINOIS: Pisca98v River, McHenry County, Imi SW Harvard (T 45N, R 5E, S6) 8 4 Mississippi River ILLINOIS: Piscaallv River, McHenry County, Imi SW Harvard (T 45N, R 5E, S6) 9 2 Mississippi River ILLINOIS: unnamed trib. Iroquois River, Iroquois County, 6 km HE Watseka 10 2 Mississip"i River ILLINOIS: unnamed trib. Iroquois River, Iroquois County, 6 km HE Watseka 11 2 Mississippi River ILLINOIS: Pisca98v River, McHenry County, Imi SW Hsnsrd (T 45N, R 5R, S 6) Lake Mi chigsn WISCONSIN: Root River, 3 mi NE Lake Muakego (UMM2 12846) 12 2 Lake Michigan WISCONSIN: Menomenee River, MilvsukeeCounty; atWeat Allis (IJMMZ 64550) 13 2 Misaiasippi River ILLINOIS: Pisca98v River, McHenry County, 1 IIi SW Harvard (T 45N, R 5R, S 6) 14 2 Mississippi River ILLINOIS: PiSCS98V River, McHenry County, 1 ad SW Hanard (T 45N, R 5R, S 6) 15 3 Spring River MISSOURI: Grand River headvaters, Lawrence Co, upstream Verona (tJPIlZ 103031) 16 Missouri River MISSOURI: Big Gra,aia Cr., Morgsn Co (OBBge I NW GrayaisMUls: IJMMZ 150380) 17 5 MisSissippi River WISCONSIN: unnamedi trib. BlackiRiver, Clark (T27N, R lV,Si8:. MPH 18423) Missouri River MISSOURI: Big Grs1ais Cr., Morgan Co (098ge lIiNWGr81'sis Mills: UMM2 150380) Spring River MISSOURI: Grand Rfyer headvat~ra, Lawrence IIi upstream Vetiona (IJMMZ 103031) 18 Mississippi River WISCONSIN: unnamed itrib. Black River , Clark (T27N, R IV, S~:i MPH 18423) 19 I Mississippi River WISCONSIN: unnameditrib. Black)River, Clark (T 27N,R lW, s~:1 MP1118423) 0'\ \0 LARVAE LITERATURE Few descriptions of the early life history of the least darter presently exist. Petravicz (1936) described embryonic development of the circulatory system of... E.mic::roperca. Length at JiafclfiIi-g-·was •• reporredas 3.0 mm TL. Burr and Page (1979) described a hatchl~ng 3.5 mm in TL as mostly translucent withindfvidtial melanophoresonthehead,yolksac, and some myomeres. Pecto:t'.alfiIlS·.of 3.5mmlarv8were1.a.rgea.t.\ha.tching and the tail had an unusual shape.. Three day 0:J.d.· •. ···:J.arva~(4.{jmm'I'L)had 10 ~: pectoral fin.rays, several caudal fin rays, and melanophores concentrated on the yolk sac remnant, head, and myomeres. A larger individual (4.7mm TL) had the yolk sac nearly absorbed, thec::atiQalfifi--wJ:th-many-ue·v~-toped rays, and large pectoral fins. However, the pelvic, dorsal, and anal fins had not differentiated. At 7 mm, all fin rays had developed, and branchiostegal rays were well develope~.:r>:tfJC::!_~t.~ .UlelanophC:>J;~e; were scattered over the head and body, and outlined fin rays and myomeres. At 10 to 11 mID, scales were visible on the top and sides of the body, but the nape and head were still devoid of scales. The cephalic lateralis system was completely developed. At 12 to 13 mID, body squamation was complete wi.thno scales present. on the opercles. The opercule spine was developed and juveniles had adult head pigmentation. UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Characteristics of length and depth are described for larval and early juvenile least darters including the length at formation of morphological TABLE 14. Morphometry of Etheostoma microperca larvae and early juveniles grouped by 1mm intervals of total length (N = sample sbe) . parentheses. Mean lengths followed by range in LEHGnI (m) !i (a) (b) (d J:.Q.:l:;\..1, STANDARD PREANAL ~ ]!!Q!IT m BODY DEm GREATEST CAUDAl, PEDUNCLE 2 3.20.0-3.5) 3.0(2.7-3.2) 1.6(1.5-1.7) .52(.50-.53) .06 .24(.23-.25) .23(.22-.23) .52(.47-.56) .12(.10-.13) 3 4.2(4.0~4.5) 3.7(3.5-3.9) 2.1(2.1-2.2) .85(.80-.90) .10 .35 .33-.37) .43(.40-;45) .16(.15-.18) 6,4 5.6 3.1 1.5 .34 .43 .68 .93 .43 (, 7,"(7.(l-7.9) 6.0(5.3-6,4) 3.6(3.3-3.8) 1.8(1.5-1.9) .34 .48( .40-.50) .92(.72-1.0) I.3(1.1-1.4) .5S( .51- .(4) 4 e.5(8.~-8.9) 6.8(6.6-7.1) 3.9(3.8-4.2) 1.9(1.8-1.9) .36( .25-.43) .52( .51-.55) 1.0( .93-1.2) 1.3(1.3-1.4) .70( .59-.76) 9 9 9.8(9·r- • ) 7.7(7.6-7,7) 4.4(4.2-4.5) 2.2(2.2-2.3) .43 .59 1.3(1.3-1.4) 1.3(1.3-1.4) .76 4 10. ',(lOl2-10.9) 8.5(8.1-9.0) 4.9(4.7-5.2) 2.5(2.2-2.7) .50(.42-.60) .65(.&0-.70) 1.4(1.2-1.4) 1.5(1.3-1.7) .99(.85-1.2) I 1.4(1li3-1 1.5) 9.3(8.9-9.6) 5.3(5.1-5.6) 2.8(2.7-2.9) .55( .51-.59) . 72( .68-.76) 1. 7(1.5-1.8) 1.7(1.6-1.8) .98( .93-1.0) 12.:<(1212-17.4) 9.8(9.8-9.9) 5.4 3.0(2.9-3.1) .68 .76 1.9(1.8-2.0) 1.9(1.8-2.0) 1.1(1.0-1.2) 13.4(13.2-13.6) 10.7(10.5-10.8) 6.4(6.4-6.5) 3.3(3.2-3.4) .69(.67-.70) .80(.70-.80) 2.2(2.1-2.2) 2.3(2.1-2.4) 1.2(1.1-1.2) 14.1(l4.5-14.9) 11.9(11.7-12.1) 7.0(6.9-7.1) 3.6(3.5-3.7) .72(.68-.74) .80 2.3(2.3-'2.5) 7.4(2.3-2.5) 1.3(1.2-1.4) 15.4(15.!1-15. 7) 13.3(13.3-13.4) 7.5(7.2-7.6) 3.7(3.4-4.1) . 74( .68-.85) .. 90(.85-.93) 2.5(2.5-2.6) 2.5(2.5-2.6) I 1 16.1 13.0 7.1 3.7 .76 193 2.5 2.6 17:~(I7;f-17'9) .".~) 14.4(13.9-15.2) 8.2(7.6-8.5) 4.0(3.6-4.3)' .79(.68-.93) .90(.76-1.2) 2.8(2.6-3.0) 2.8(2.7-3.1) ".",,; ".>(".'."." ">(""'.') •."U·..." , .,".,....." ""i.85-1.1) 3.0(2.9-3.1) 3.0(2.9-3.1) 19.3(19.i _19.:1) 15.7(15.5-16.0) 8.9(8.7-9.0) 4.4(4.2-4.4)i .72(.68-.76) 1.1(1.0-1.2) 3.3(3.1-3.4) 3.3(3.1-3.4) 1 Rndy dep~ b mpn~un'd i dorsal to ventral surfac4 at posteriorportion 9£ anus Greatestllbody tl"pth to vE'ntral sJrfacE' including thE' Y01!k sac C.audal Ii•• to venlra] surface at penH1 ti~ate myomere -...J I-' .... 72 structures (Table 14). At lengths 3.0 mm(newly hatched}: well developed pectoral fins with 9 incipient rays, yolk sac bulbousanteri caudally; yolksac25~6%TL, 27.9 %SL,yolktranslucent;singleanterior oil globule; head not deflected over yolk sac; jaws well developed; yolk absorbed (ca. 4.5 mm); eyes oblong. First rays formed were pectoral (3.5 mm); first dorsal rays formed (4.0 to 4.5 nnn); Incip:l.ent--1torsal fin margin partially differentiated (ca. 6.4 mm); c~mpletely differentiated (8.2 mm); spinous UULt>tl.L fin origin over mvomeres 5 to 6, soft dorsal origin situated over preanal myomeres 14 to 15 (ca. 8.2 mm); predorsallength27.T% TL (range:2Llto ·3L3 %) ,34.0 % SL(raIlge: 23.0 to 37.2%); notochord flexion (ca. <4.5 mm) ,preceeding first caudal fin rayforJll8tion(ca. 5.0 mm); caudal fin truncate (ca. 6.4 to 7.0 mm); first anal fin rays formed (4.5 mm); .incipient anal firi margin parEialIydl!ferenEiaIe(C{o-~4rnm), completely differentiated by 7.6 mm; pelvic buds formed anterior to dorsal fin origin (ca. 6.4 mm); first pelvic fin rays formed (8.2 mm). No swim bladder . formed; gut .. s1::raigJ:rt; entire finfoldabsorbe_d (...ca_.8.2.. mm). Sup:rf3;():rJ?~t:f.l!.f3;1l4 ..~Il.fE.f.l(;).!:b~t.fl! .h~ad canals started forming (7.~7 to 8.3 mm); supraorbital and infraorbital head canals completely formed (10.7 to 15.7 mm), infrorbital canal complete without interuptionand pore count of 10 (ca. 7.7 mm), retrogression to interrupted pore count of 0 3 00.7 to 15.1 mm); preoperculomandibular and supratemporal ca.rial bega.ri forming (12.2 mm), completely formed (12.4 to 15.1 mm); lateral line began forming 15.1 mm; scales present at 8.9 mm; squamation completed 12.2 mm; scales do not form on or cheeks. Man.4ible and maxillary _~~_ equal length extending anterior to orbit (at all lengths). Meristics Myomere number constant posthatching with 15 preanal, 19 to 20 73 TABLE 15. Selected meristic values and size{mmtotallength) at the apparent onset of development fo~. Etheost0ma.m~c:.ro1?erc:.a. Mean-values are underscored~.Rare or questionable extremes are in parentheses~ The l'lWriberofsecoridary rays of .. the median fins are in lowercase Roman numerals. (a) Attribute / Event Present Study Literature (1-8,10,12,15) Dorsal Fin Spines / Rays VII-VIII / 7-~ VI~VII-VIII/8-9-10-11 first rays formed 4.0-4.5 no information adult complement formed 6.4 (1-8,10,12,15) Anal Fin Spines / Rays II / 5 I-II / 4-5-6-7 first rays formed 4.5 no information adtiltcomplement formed 7.5 no information (4) Caudal Fin Rays vii-xv, 4-1 + 5, vii-xii 9 - 10 .. 11 - 12 mmmm_m.mmmC9,13)__ firstir~ys formed 5.0 4.0 (9,13) adult complement formed 7.0 7.0 9_10_11_1~1-2,4,15) Pectoral Fin Rays 10-11-12 first rays formed 3.5 no information adult complement formed 4.5 no information (1-2,4-7,10,15) Lateral Series - Scales 27 - 11 - 34 30-E=.ll-36 (1-2,10,14,15) Myomere / Vertebrae 34-35 / 32-33-34 no information/31-32-33-34 preanal myomeres 15 no information postanal myomeres 12.- a. 1. Page 1983 2. Kuehne and Barbour 1983 3. Smith 1979 4. Burr 1978 5. Trautman 1981 6. Forbes and Richardson 1920 7. Jordan and Evermann 1896 8. and Underhill 1974 9. Burr and Page 1979 10. Becker 1983 11. Fish 1932 12. Pflieger 1975 13. Auer 1982 14. Bailey and Gosli ne 1955 15. Scott and Crossman 1973 74 postanal, and 34 to 35 total. Total vertebra~ ~~tQ~4JN""~Li",,33.0) including one urostylar element from specimens from thePiscasaw River, Illinois. Scales in the lateral series ranging from 27 to 34 (N = 26; x = 31.3; mode =. 30). Paired and median fin ray·· values and length at appearance are presented in Table 15. Pigmentation Lengt.h:r:ot.6 3.5 rom (newly hatched): heavily pigmented (Fig. 14); retinae black; concentration of melanophores on dorsal portion. of cranium outlining tb~ optic lobe; a profuse scattering of melanophores outlining all preanal and pQstanal myomeres distributed along all myosepta in epaxial and hypaxial.regions. Melanophores p()~t:~l:":l()l:" ... J:() ....J!l~xJ.1Jar.y. :l!1__ ~_I'~f.l of future operculum. Laterally, melanophores encircle the yolk sac leaving the center void of pigment. Ventrally, a single pair of melanophores under mandible, concentrated at anterior portion of yolk sac, less concentrated posteriorly. Melanophores at apex of every postanal myomere dorsally and Lengt.h 4.0 rom: eyes black; laterally, melanophores at apex... ofevery myosepta with epaxial and hypaxial melanophores scattered over entire body. Melanophores outline the gut dorsally and ventrally. :P:l8J!l~Ilt:f.lt::l()Ilpresent anterior to orbit dorsal to maxillary; pigment slightly posterior to orbit, evenwith mandible and maxillary. Dorsally, melanophores concentrated on optic lobe (Fig. 15). length 4.5 ·mmr- pigmentation absent.· over- the-·ma3orit-y- of body .. Laterally, pigmentation on head posterior to orbit on future operculum. Dorsally, melanophores outlining optic lobe; ventrally melanophores concentrated at apex of breast. FIG. 14 .. '~ Etheostoma microperca, least darter (newly hatched ~~~~~_~;harvaLL3.0~_!JIIIl_~~~J_J)i.scasawRiver, Illinois a. dorsal b. lateral c. ventral. )31 ,.. Jt1 f ~V ~ I I I I ':) I 1 I I ) f 'q r '8 I J I I I SL ______J FIG. 15 Etheostoma microperca, least darter larva, a. 4.0 mID . ._.,~'J'L, unnamed tI.i.pJ,ttary Iroquois River, Illinois b. 6.4 mID TL Piscasaw River, Illinois. i ~------; wwa. wwa. '8 i I I I 9L ______:1 rIt' § 77 Length 6.4 to 8.2 mm: laterally, scattered melanophoresover the entire body outlining myomeres; 6 to 8 distinct areas formirigblotches; concentration along'gut. Melanophoresoncranium.anteriorandposterior to orbit,' and dorsally covering optic lobe. At the base of the differentiating anal fin is a concentration of melanophores at ventral base. Dorsally, 6 rectangular saddles forming. piginent~j)atternB were essentially unchanged for the duration of the larvalperigd",witJl patterns less prolIliIl~Ilta.fter 7.0 mm TL (Fig. 17). Length 8.5 to 8.9 rom: lateral lIlelanophores outliningJnyomeres along myosepta;· 7to9 blotches along midlateral; 2 half crescents present at base of, caudal peduncle. Head pigmentation withdistinctpreorbital, and ....,. postorbital bars;'jsuborbital bar forming. Edge of suboperculumoutlined; a dorsal chevron forming posterior, toorbit~laterally. Dorsally, melanophores concentrated over optic lobe; and forming 8 to 9 dorsal saddles (Fig. 16). ,Length 9.7 to 13.2mm: a midlateralconcentratiQn oLmelanophores I forming 7 to 9 lateral plotches; dorsal edges of scales pigmented; i preorbi~al, postorbital and suborbital bars chevron shaped. Dorsally, 7 to I 9 saddles; specks of melanophores behind concentration on optic lobe. i Specks of melanophores distributed between pectoral fin rays; melanophores i scattered in spinous and soft dorsal, caudal, and anal fins. Ventrally I melanophores at base of each ~epidotrichia of anal fin. Length 13.6 to 14.7 mm (juveniles): uniformly scattered pigment I connecting]to f3:t"E!£!:a.Il&l1la.I" midla.~~ral~~()~t:hE!,fil,t() 7 to 8 dor~a.l saddles; scattered melanophores extending ventrally outlining scale. edges connectirig midlateral blotches; dorsal section of cranium withmelanophores over optic lobe; outline of scales pigmented; entire pectoral fin with scattered melanophores on membranes; 3 oblique stripes of clustered melanophores on FIG. 16 Etheostoma microperca, least darter, Piscasaw River, ~ __a. __B.•2_JDJIL-TL __larva b. 14.5 mm TL early juvenile. 'q I ww ~ ;, '8 I i I i I I I I t 8L l :~c~ ,~"., _,~~_~ ~.~. ~ ~~~~~".~_, ~_ c,._~".cc_~~~ ,~,. ~ __ ..• __ ...... 79 the spinous and 4 oblique stripes on the soft dorsaL {pigmented directly on the ray); caudal fin with 7 diagonal D~L.~~C membrane; preoI'bital bar I extending above maxillary; scattered melanophores on operculum and cheek. i Ventrally melanophores concentrated at pterigiophoreinterdigitation with I lepidotrichia of anal fin (Fig. 16). i I Natural History The least darter inhabits the sluggish stream margins associated with submerged vegetation and filamentous algae along overhanging grassy banks (Petravicz 1936; Burr and Page 1979; Trautman 1982; Becker 1983; Page 1983; Kuehne and Barbopr 1983; present study). In the present study spawning occurred in the Upper MississippiR:I,v~r,_no!:~llc~Rt1:a~ :tllinoi_~, and mid~July, southeastern Wisconsin from mid-MaYUfitil at ,temperatures of 16 to 18 C. Burr and Page (1979) reported spawning in southern Illinois and Missouri to occur from late April until mid-July. Aquarium observations I indicated the demersal tendency of larval least darters. In the field larval and juvenile darters were collected from similar habitats as adults. I!J Eggs incubated at 15.5 C hatched at 264 ± 13 hours, C hatching occurred in 181 ± 7 hours, and at 22 to 12 hours, while eggs incubated at 27 C fungused (Burr and Page 1979). Eggs incubated in the present investigation at 18 C hatched after 144± 5 hours. This agr,eed with previous results obtained at 18 to 20 C in which hatching I (Petr8":i~z occurred in 144 to 148 hours 1936; Winn 1958b) .--- SPOTTAILDARTER J Etheostoma sguamiceps.Jo:rcian The ciistribution of the spottail ciarter,Etheostomasguamiceps,in the Upper Mississippi River basin includes southern Illinois anci southwestern Inciiana. The species range inclucies small to moderate sizeci streams of the lower Tennessee River cirainage in Alabama and Mississippi, north into the Green River of Kentucky, tributaries of the Ohio River in southern Illinois anci northwest Kentucky, and tributaries of the Wabash River in southwestern Indiana. The species is found in tributaries of the Mississippi River southwest of Kentucky and northwest Tennessee (Clay 1975; Smith 1979; Page 1980; Page 1983; Kuehne and Barbour 1983) Sparce information is available on the taxonomy and naturaL history of the spottail.dafter with even .. less information available on the early life history. The type collection locality of the spottail darter was from Russellville, Logan County, Kentucky (Jordan 1877). The systematics of the species were compared .to. E. neopterum (Howell~andDingerkus-1928)and .. to E. 01ivaceum(~I"a.ascha.ll; E. s9~~miceps awaits elucidation and will result in additional species (L.M. Page, personal communication). The diet of the ~po~tail darter includes immature aquatic insects and microcrusteacens (Forbes 1920; Page 1974). The life history of the spottail darter was studied in Big Creek, Illinois and Ferguson Creek, Kentucky (Page 1974). Limited early life history information was included in the life history account by Page (1974). Location dat:~ic:>!:~I>~cimell;~ used in the prese~!~~~~!~dI=l;;.:I,)E~sented in Table 16. 80 TABLE 16. Material examined of Etheostoma squamiceps larvae and early juveniles grouped by 1mm intervals of tot.al lengt.h (N = sample size). Tl1fAL LEIIG'l1I (_) ~ DRAINAGE ~ 5 4 Ohio River ILLINOIS: Big Creek, Hardin County (T 33N, R 8E, SIP) 6 16 Ohio River ILLINOIS: Big Creek, Hardin County (T 33N. R 8E. SIP) Ohio River ILLINOIS: Big Creek, H,ardin County (T 33N, R 8E. S 1Pl 8 3 Ohio River ILLINOIS: Big Creek, Hardin County (T 33N, R 8E, S 10) 10 Ohio River ILLINOIS: Big Creek, Hardin-COunty (T 33N, R 8E, S 10) 11 Ohio River ILLINOIS: Big Creek, Hardin County (T 33ll, R 8E, S 10) 12 Ohi" River ILLINOIS: Big Creek, Hardin County (T 33N, R 8E, S 10) 11 Ohio River ILLINOIS: Big Greek. Hardin County (T 33N, R 8E, S 10) 15 Ohio River ILLINOIS: Big Creek. Hatdin County (T33ll, R 8E, S 10) 16 Ohio River ILLINOIS: Big Creek, Hardin County (T 33N, R 8E, S 10) 17 Ohio River ILLINOIS: Big Creek, Hardin County (T33N, R 8E, S 10) 19 2 Ohio River TENNESSEE: Duck River at Brench Ia. R.M. ISO, Maury Co. (UTRC 91.2465) 20 Ohio River KEIl'l1JCKY: Gooae Creek, Caldwell Co (l 1/2 IIi S Princetovn Rt 139: \lItlZ 174859) 211 Ohio River TENNESSEE: Duck River at Brench Ia. R.M. 150, Maury Co. (UTRC 91.2465) 221 Ohio River ILLINOIS: Hogt~ief Creek. Hardin County (\lItlZ 200929) KENT1JCKY: Goose Creek, Caldwell Co (l 1/2 IIi S Princetovn Rt 139: llHHZ 23 2 Ohio River KEIl'l1JCKY: Goose Cre~k, Caldwell Co (l 1/2 IIi S Princetovn Rt 139: llHHZ 24" 3 Ohio River KENT1JCKY: Gooae Cree,k, Caldwell Co (l 1/2 IIi S Princetown Rt 139: llHHZ TENNESSEE: Dry Creek,: Hardin County (llHHZ 200929) I 2~ Ohio River KENT1JCKY: Gooae Cre"/,, Caldwell Co (l 1/2 IIi S Princetown Rt 139:llHHZ I KEIl'l1JCKY: BearCreek!, Grayson Co. (4 IIi S Leitchfield: llHHZ 178196) Ohio River TENIlF,sSEE: Dry Creek,! Hardin County (llHHZ 200929) 2JI 3 Ohio River TENNESSEE: Dry Creek.l Hardin County (UHH2 200929) KENT1JCKY: Goos~ Creek!, Caldwell Co (l 1/2 IIi S Princetown Rt 21 ILLINOIS: Hogthief Creek, Hardin County (~ 17l806) 281 Ohio River TENNESSEE : Duck River! at Brench Is. R.M. 150, Haury Co (UTRC91.2465) " 291 Ohio River ILLINOIS: trib.i Wababh River,White Co (5 .i NE Caimion US 45: llHHZ II KEIl'l1JCKY: Bear C.'reekI, Grayson Co 1(4 IIi S Lei. tchfield: llHHZ 178196) 301! Ohio River ILLINOIS: trib.i.' Wsba/lhi, River,,'., ,White'.i Co• (5. IIi'.",','NE Carllion,,', ... US' 45:',. UHH2• ! 311! 3 Ohio River KEIl'l1JCKY: Bear t:;reekl Grayson ;Co(4 IIi S Le:l.~chf1eld: llHHZ 178196) TENNESSEE: Dry Creek.! Hardin County (UHH2 200929) 32\ Ohio River TENNESSEE: Duck River at Brench Is. R.M. ISO; Haury Co (UTRC 91.2465) 33 River KEIl'l1JCKY: Bear Creek Grayson Co. (4 IIi S Leitchfield: UHH2 178196) 1"] r 82 EGGS I LITERATURE 'i I The spottail darter, E. squamiceps, has a spherical egg morphology I typical of other members of the subgenus Catonotus. Page (1974) reported I.•...·.·.· U that mature eggs averaged 1.6 mm, weret:.:r:allt:lluc~Ilt.,ElIl9:~~nta~!1~~_Elsingle a oil droplet. Smith. (1979) reported egg diameters averaging 1.8 mm. I UPPER MISSISSIPPI RIVER DESCRIPTION In the present study, eggs from Big Creek, Illinoit:l Were smallest of the three species of Catonotus in the UMR drainage rangillg from 1.4 to 1.9 mm (N =26,.~ = 1.8). Mature eggs were spherical, demersal, adhesive, with translucent yolk. Mature eggs had a .~~11~_~~~_~~~~~E~l~, a narrow perivitelline space, anunsculptured chorion, and were unpigmented. Eggs are attached to the underside of slab rocks in shallow pools of high gradient streams which agrees with published acounts (Page 1974, 1975a, I 1983;' Smith 1979). LARVAE :i,. LITERATURE i i Page (1974) reported the total length of themot:ltlyt'-re.llt:llucent neWly hatched larvae as 6.5 mm. At this length larvae .he.d.re.d blood, black I melanophores over most of the body, and metallic gold eyes. Mele.llophoret:l were prominent on top of the head and the lower one third of the yolk sac. I Pectoral fins' andcthe~-lower jaw---were-wel1'" -deveIop-e6~----No=ccgdditional , information is available until 12 mm. At 12 rom, the morphology was similar to that of the adult except no scales were present on the nape, breast, or belly, and lateral line canals were not developed. Pigmentation also contrasted that of the adult with patterns of large spots over all areas of the body except the belly. At 16 rom, the lateral line had begun At 19 rom, themottledpatterfi char8.cteristicoftheadtiltsw8.spresent; sca.les were present on the nape, breast, and belly, and the lateral line extended to.the soft dorsal fin. No additional information on the larval r~ development is available. UPPER MISSISSIPPI RIVER DESCRIPTION Morphology Character~stics of length and depth are described for larval and early juvenile spottail darters including the length at formation of morphological structures. (Table 17) • At~lengths~5.6~~j;~o~~_6_.LnU:lL(newly hatched): well developed pectoral fifiwith 12 ificipiefit rays; yolk sac large, spherical, about 25.9 % of SL; yolk amber, with a single anterior oilglobule; head not deflected over yolk sac; jaws developed; yolk absoTged (6.9 to 7.4 rom); eyes circular. First rays formed pectoral (5.8 to 6.2 rom); spinous and soft dorsal rays formed simultaneously (6.4 to 6.5 rom); incipient dorsal fin margin partially differentiated. (ca. 7.1.to 7.8 rom), completely differentiated (7.1 to 8.4 rom); spinous dorsal fin origin situated over preanal myomere 6, soft dorsal situated over preanal myomere 15 (6.2 to 6.9 rom); predorsal length 32.9 %SL (5.6 to6.6JDm), 36.0 % SL at lengths greater than 7.0 rom; notochord flexion (7.1 to 8.5 mm), pr~()rto first caudal fin ray formation (5.8 rom); caudal fin rounded (7.1 rom). First anal fin rays formed (6.3 to 6.6 ; incipient anal fin margin partially differentiated (6.7 to 7.1 rom), completely differentiated (6.9 to 7.8 rom); pelvic buds formed anterior to dorsal fin origin (6.9 to 7.4 rom); first pelvic fin rays formed (ca. 8.4 rom). No swim bladder formed; gut ------ TABLE 17. Morphometry of Etheostoma squamicevs larvae and early juveniles grouped by 1 mm of total length ,(N.-- = sample size). LPJIGJ'II (_) (8 (~) (e) !! :!m!< STAIlDARD m!!!!h HEAD SIIOUT EYE BODTDEPTIl GREATEST CAUDAL PEOOIlCLE 5.7(5.6-5.8) 5.5(5.3-5.6) 3.1(3.1-3.2) 1.2(1.1-1.3) .14J .13-.14) .51(.46-.56) .54( .50-.61) 1.4(1.3-1.4) .23( .20--.25) 16 6.5(6.0--6.9) 6.2(6.0--6.6) 3.6(3.3--3.8) 1.4(1.1-1.6) .21( .12-.33) .58( .54-.60) .67( .56-.84) 1.4(1.2-1.5) .31( .22-.37) 7.5(7.1-7.9) 6.7(6.3-7.1) 4.1(3.8-4.2) 1.8(1.6-2.0) .31( .18--.40) .60( .58--.63) ; 70( .60--.80) 1.3( .91-2.0) .38( .32-.46) 8.5(8.4-8.6) 7.4 4.4 2.1(2.0--2.1) .37( .26-.50) .69(.68--.70) -:78(.76-.80) 1.2(1.1-1.2) .50 10.9 9.5 5.7 3.1 .64 .90 1.4 1.9 .78 11.8 9;7 6.1 3.1 .62 .92 1.4 2.1 .90 12.0 10.4 6.6 3.6 .70 1.0 1.7 2.4 .96 13.6(13.6-13.7) 10.2(9.3-11.1) 6.7(6.6--6.7) 3.4(3.3-3.4) . 70( .58--.82) 1.0(.98-1.1) 1.8(1.7-1.9) 2.4(2.2-2.6) 1.0 15.6 12.8 8.2 4.0 .80 1.2 2.3 2.9 1.4 16.7 14.3 8.9 4.4 .70 1.2 2.4 3.1 1.3 17.9 14.6 9.1 4.8 .90 1.2 2.6 3.3 1.7 19.4(19.3-19.5) 16.0(15.9-16.0) 9.8(9.7-9.9) 5.1(5.0--5.2) .92( .88--.96) 1.3 2.7(2.7-2.8) 3.5(3.3-3.6 1.7 20.1 16.8 10.3 5.2 .70 1.4 2;9 3.5 1.9 21.2 17.2 12.6 6.1 1.1 1.4 4H 4.5 2.2 2 22.9(22.8-22.9) 18.9(18.8--18;9) 11.6(11.5-11.8) 5.8 i 1.0(.90-1.1) +1.6) 3.5p;2-3.7) 3.9(3.6--4.3) 2.2(2.1~2.2) 2 23.6(23.3--23.9) 19.8(19.5-20.2) 12.2(12.1-12.3) 5.9(5.8-61.0) 1.1(1.0-1.1) 1.5(1·t-I .6) 3.6(3.5-3.7) 3.9 2.2(2.1-2.2) i 3 24.5(24.5-24.6) 20.5(19.8--20.9) 12.8(12.3--13.4) 5.8(5.8-61. 0) I.O( .90-1.1) 1.5(!.i-I.7) 3.5(3.3-3.7) 4.1(3.8-4.4) 2.2(2.1-2.3) 2 25.1(25.0-25.1) 21.0(20.8-21.2) 12.9(12.9-12.9) 6.1(6.0--61.2) I.6(1.4-1.7) 1.6(1.+-1.7) 3.8 4.6(4.5-4.6) i I 26.4 22.1 13.7 6.6 i 1.0 1.8 3.5 4.2 2.3 3 27.7(27.3-27.9) 22.9(22 .0--23.6) 13.9(13.4-14.4) 6.5(6.0--61.9) 1.1(1.0--1.1) I. 7(1.~-1.9) 4.1(3.7-4.4) 4.7(4.5-4.8) 2.6(2.4-2.8) I I I 28.8 24.3 14.5 7.2 I 1.4 2.0 4;4 4.8 2.8 2 29.4(29.3-29.5) 24.6(24.3-24;9) -14.9 7.2(7.0-7j.4) 1.5(1.4-1.5) 2., 4.6(4.5-4.6) I 30.8 26.0 15.6 7.3 I 1.5 2., 5.3 5.8 3.0 i 3 31.8 26.3(26.1-26;6) 15 8(15.7-16.1) 7.6(7.5-~.6) 1.4(1.4-1.5) 2.2(2./-2.2) 4.6(4.0--5.0) I 2.~ 3.2 I 32.9 27.6 16.6 7.7 ' 1.6 5.3 6.1 1 i' 33.1 27.3 16.8 7.5 1.6 2.1 5.2 5.9 3.2 (8) Body depth meesuredverticel1y f~(). the,;ddrssl t.o'f'entrel 8urtece at posterior portion o~ anus (c) Caudal peduncle depth mes8uredvertically:!rom the dOTsalto ..entre! surface at penulU.ate .JOIIere straight; entire finfo1d absorbed (10.7mm). Infraorbital, lateral, and preopercu10mandibu1ar canals formed (12.0 mm); infraorbital canal complete with 7 pores extending to anterior of midorbit (13.6 to 15~6mm); completely formed with retrogression to interupted pore condition 4 anterior and 4 posterior (24.5 to 27.8 mm). Lateral canal formed, preopercu1omandibu1ar pores 10, supraorbital pores 3, arid 2 supratemporal canal pores (16.7 mm). Lateral 1ine.l>ega.n forIIl1,n8.(ca.. 16.7 nun); comp1ete~Y~()I."Ill~~extending to soft dorsal fin (20.0 to 23. mm). Squamation initiated at 12.0 mm, squamation complete (20.0 to 23.9 mm); sca1espreseIltoIl the Ilspe, operculum, prepectora1, breast, arid belly. the cheeks are·usua11y unsealed or possess·on1y·afew·sca1es posterior to the orbit. Meristics Myomere number constant posthatching, preanal myomere 16, postana1 myomere 18 to 19 with 34 to 35 total myomeres (postanaLmyomere: ..x"'!' 18.2; N= 30). Total v~.:t".1::.~.~.I.".l:l~ . Ilumbering 36 (N = 3; x= 36.0) including one urosty1ar element from specimens from Big Creek, Illinois. Scales in the lateral series ranging from 37 to 57 (N = 25, x=44.7, mode =47) • Paired and median fin ray values and length at appearance are presented in Table 18. Pigmentation r.ength 5.6. t:() .. 9.!1 mm (newly.hat.c:ll~dJ:. heaviII Ri8!@ntecl;~ret:i.na.e black; a mid1atera.1 stripe of single me1anophores extending from the apex of preanal myomeres 2 to 14; extension ofme1anophores into epaxial portions of preanal myomeres 6, 7, and 12, into hypaxial portions of preanal myomeres 11 thru 13; pigment dorsally and ventrally on gut; a TABLE 18. Selected meristic values and size (mm total len.gth)iat the apparent onset of developmenfforEtheosfomasg:ua.m:1.ceps. Mean values are underscored~Rar.eoriquestJ.(jJj8bJ..eej{trellles are in parentheses. The number of secondary raysiof the median fins are in lowercase Roman numerals. (a) Attribute / Event Present Study I.,iterature (1-5) Dorsal Fin Spines / Rays VII-VIII-IX / 10-11-12-13 VI-lli....XII1l-12-13-15 first rays formed .4-6.5 no information adult c.omplement formed no information 0-5) Anal Fin Spines / Rays II / 6-1. II / 6-1.-8 first rays formed 6.3-6.6 no information adult compl~m,ent formed 7.0 no information Caudal Fin Rays x-xiii, 8 1.+&1., ix xiL nQ.information . first rays formed 5.8 no information adult complement formed 7.2 no information (1-2) Pectorai Fin Rays 11-12-13 10-~13 first rays formed 5 .8· ;;;;0:2·· ···········~--_··lIO-i:fifoflllAtiofi- adult complement formed 6.4 no information (1-5) LatHal Series - Scales 37 - 44 - 57 41-44-57 (1-2,6) Myomere / Vertebrae 34-35 / 36 no information/35-~-37 preanal myomeres 16 postanal rnyomeres 18 - 19 8. I. Page 1983 2. Kuehne and Harhour 1983 3. Forhes and Richardson 19204. Smith 1979 5. CJ ay 1975 6. Bail ey and Gosli ne 195~) 87 narrow band of melanophores extending from anterior to posterior of yolk sac. Postana.lInY()In~res with scattered pigment outlining all myosepta, extending into epaxial and hypaxial portions of myomeres. Cranium with several melanopho~e~ posterior to orbit; dorsally optic lobe with 6 rows of melanophores extending posteriorly to nape;severalmelanophoreson outside margin of cerebrum. Ventrally, a concentration ofmelanophores at the ant:~~~()~y()l.l< ~ac::.f()rln:i.ng a band of scattered pigment comprising one third of entire ··yolk ~ac; several rows of melanophores on gut to anus; all postanal myomeres with a pair of melanophores (Fig. 17). Length 6.2 to 7.0 mm: a midlateral stripe of single melanophores at apex of preanal myomeres 6 through 14, and postanal myomeres 1, 3 through 7, and 9 through 12; epaxial pigmentation near base of yolk sac on preanal myosepta and postanal myosepta 5 through 9. Ventrally, melanophores on center third of yolk sac and gut; paired melanophores on all postanal myosepta. Dorsally, melanophores concentrated into 6 distinct rows over optic lobe and cerebellum; several melanophores on exterior margin of cerebrum (Fig. Length 7.1 to 8 .. 0mm: on cranium, a concentration of melanophores over optic lobe and cerebrum; anterior to orbit, several melanophores forming future preorbital bar; several oblique melanophorespost~rior tOiaw. At base of breast a concentration of melanophores forming a single series of pigment extending from throat to posterior of anus; a midlateral series of pigment extending from apex of first preanal myomere to hypaxial plate; several groups of -pigment dorsally·· .at· .anterror spintltisdorsal. Melanophores on prepectoral areas, and at base of anus. Ventrally, a series of melanophores at base of all developing anal fin lepidotrichia, and at preanal myomeres 10 through 16 and postanal myomeres 1 through 4, 6 ...: FIG. 17 Etheostoma sguamiceps, spottail darter (newly hatched -"--"1a-r-va7,~-5".7~mm--TI:;, Big Creek, Illinois a. dorsal b. lat eral c. ventral. ______J··tt '0 ", .""':"."/ ," ',/ .... \. wwa. '8 aa ,J FIG. 18 ...: Etheostoma sQuamiceps, spottail darter larva, Big Creek Illinois a. 6.8 mm TL b. 8.6 mm TL. ______Ji2 'q ww ~ ., ~ .k¥.~.¥ .~' !/ .../ 68 J r 90 through 9, and 11 through 13 with some extension tomidlateralstripe. Some with 2 lines of pigment extending into ventral half of caudal fin. Length 8.0 to 9.2 rom: ccmcentrationof me1.8IlophQI'eEJ Q'VeI'Qptic lobe and cerebrum;preorbital and postorbital bars formed; several melanophores formirig. future· suborbital bar; pigment forming optic lobe advancing over cerebellum. Laterally, midlateral pigmentation forming~~15~to-16-clusters; dorsally, increased pigment forming 13 to 14 areas\of concentration. Ventrally,pigment near throat, and near lepidotrichia 18). Length 10.7 to 12.0 rom: preorbital, postorbital,8ndsl.lborbit.al bars formed around orbit; dorsally, melanophores concentrated over cerebrum, optic lobe and nape. Laterally, pigment on prepectoral area; 15 to 16 midlateral rectangular blotches withscattered~~pigment~-e0nnecting vertically to 14 dorsal saddles; pigment over anal fin ray insertion; and melanophore~ present near throat. Scattered melanophores on spinous and soft dorsal. Length 13.6 to 16.7 rom (juveniles): melanophores outlining newly formed scales, connecting 10 to 12 dorsal saddles to 13 to 14 midlateral rectangular blotches; pigment present on prepectoral girdle. Head with distinct preorbital, postorbital, and suborbital bars; concentration of melanophores on ventral half of opercle; preorbital bar extending onto the tip of maxillary; several lines of pigment radiating posterior from orbit between postorbital and suborbital bars and between the postorbital and optic lobe pigment. Cerebrum, optic lobe, and nape with concentration of melanophores. Last dorsal and midventral pigment clusters forming 3 distinct spots on hypural plate. Pigment forJDiIl~~~~llgle mid-ray stripe on spinous and soft dorsal; 3 crescent stripes in caudal fin; and a single stripe continuous with body near anal fins. Pectoral and pelvic fins T FIG. 19 "',: Etheostoma sguamiceps, spottail darter; Big··Creek; Illinois a. 13.6 mm TL larva b. 16.7 mm TL early -,ju:\tenile-.------°e T6 - 92 without;.", pigment (Fig. 19). Length 22.Qt9 33.0 mm: essentially the same as above .however, melanophores outlining maxillary, interopercle, and forming a vermiculate pattern dorsally on.cranium; pigment present on cheeks. Regulated patterns of early juveniles changing to mottled patterns or ~~~ll~~ts with many indistinct markings. Dorsally, 8 to 10 saddles. Spinous dorsal with 3 incomplete stripes limited to fin ray; soft· dorsal . with 4 incomplete stripes; caudal fin with 5 to 6 incomplete stripes; and pectoral fin with 4 or 5 incomplete stripes. Anal and pelvic fins 'WithOUt stripes or pigment. At base of hypural plate are three distinct spots. Nohurneral spot present. Natural History The spottail darter inhabits the shallow slab pools and riffles of high gradient streams (Page 1974, 1984; Page and Schemske 1978; Smith 1979). In the lower reaches of the Upper Mississippi river basin li. sguamiceps spaWIlsfromlateMatch until· late May, at temperatures from 14 to19G(PageI974).Eggs incubated at 18 to 22 C hatched after 270 to 275 hours, while eggs incubated at 22 to 26 C hatched in 125 to 130 hours (Page 1974). Aquarium observations indicated that spottail darter larvae remain closely associated with the substrate and are primarily demersal (L. M. Page, personal communication). Adult populations occupied· the slab rock pools throughout the year. Larvae and early juveniles inhabited the slab pools but also gravel~='andnslabtlffles~;·~~Pa&-e.n(i9'74'Y ~ieported that occupation of gravel riffles is due to theabilitjofjiouIlgdarters to hide among the coarse gravel. The habitat of E. sguamiceps is delineated by the presence of slab rock substrates. DISCUSSION The ability to segregate percid larvae into smaller taxonomic. groups would be a definite asset in identification of the selected UMRspecies. Larvae of all percids can be characterized . by the nearly equal preanal to postana1 length, a single anterior oil globu1e,high1y deye~op~d pectoral buds or finfo1ds, and a bulbous yolk sac. Definite data gaps prohibit positive separation of darter species, however, information is available for eleven .. of the seventeen species in the drainage. There is no information concerning the eggs and larvae of the gilt darter, E. evides nor for larval phases of the western sand darter, Ammocrypta clara; b1untn9sedafter, E. ch10rosomum; Iowa darter, E. exile; b1ackside, P. - . ...- macu1ata or slenderhead darter, P. phoxocepha1a. The six species described in the .present study are distinct from other ~embers of the Etheostomatini previously investigated (Table 19). Darters are reported to have three modes of reproduction (Page 1983). These are ... buriers, attachers,··and c1usterers~ Eggburiers-dep~git-eggsintothe substrate for protection and include P. caprodes semifasciata, and P. shumardi. E. microperca represents an egg attacher which uses vegetation, dead leaves, and otherfiberous material to deposit eggs and then abandon them. Egg c1usterers deposit eggs on the underside of a stone, leaf, or other object and are guarded by the resident male and include the species E. f1abe11are 1ineo1atum, ~. kennicotti, and E. sguamiceps. Spawning locations and egg characteristics of UMR drainagepercids are summarized for.cQ~pari.son(Tablel.9}. Severa1·e~lusteringpeI'cids deposicteggs on the undersides of rocks. These rock depositors include·· the striped fantail darter, E. f1abe11are 1ineo1atum; johnny darter; E~nigfW1l; stripetai1 darter, E. kennicotti; and spottai1 darter, E. sguamiceps. Several 93 ..... 94 TABLE 19. Egg characteristics of Upper Mississippi River Yolk/oil SpeCies Egg Dismeter globule color Spswning !.oestion Literature Ammocrypta .....£!!!:!. 0.8 - 1.0 unknown main channel border Etheostoll8asprlgene 1.3 - 1.4 yellow 011 shallow backwater arees with stag globule nant to sluggish flow; on plant Page et a1. 1982 material Cummings 1983 ~.~ .8 aand riffles: 0.1 - 0.2 m depth Winn 1958a Cooper 1978 ].. chlorosomum 1.0 unkno","n deposited on plant or plant debris Winn 1958a Page et a1.- 1982 ].. !.!.lli. 1.1 orange or. ye1- fi berolls material t undercut banks low oil globule ].. flsbellare 2.3 ~ 2.7 yellow yolk, underside rocks: 0.2 to 0.3 m depth smber oil globule ].. kennieott1 1.8 - 2.5 orange yolk underside of slab rocks in shallow author upstream pools Page 1975 .§.. microperc8 0.7-1.1 smber yolk attached to vegetation: along stream author or lake margin Petraviez 1936 ------~Burr-and-Fage--1-979------ ].. nigram 1.5 yellow yolk underside rocks: 0.2 to 0.4 m depth Winn 1958a Speare 1965 Cooper 1979 .J:;. proeHsre 0.8 - 1.1 smber yolk attached to vegetation: along stream Simon 1985 or lake margin Burr and Page 1978 Burr and EIHnger 1980 ].. '!!peetabile 1.2 - 1.5 unknown clear riffle streams with fine gravel Linder 1958 Smith 1979 ~.s9uan:iic.eps 1.4 - 1.9 smber yolk underside pools Page 1974 fi18111eI1~911!l. algae, aquatic .moss or Lathrop 1978 plant debris in stream riffles: 0.6 m Auer 1982 E.!!:.E.! flavescens 1.8 - smber yolk shal1o\lt areas, often entangled in Mansueti 1964 vegetation Norden 1961 ~c8prodes 1.1- 1.7 smber yolk buried in sand or gravel: 0.1-0.2 m author depth Winn 1958a Cooper 1978 Hubba 1967 Paine and Balon 1984 f·~ unknown unknown buried in· 'ssndandgravel int'er;;.;. Page et al 1982 spersed with cobble f·~ 2.0 unknown fine gisvel and sand: 0.3 m depth Petraviez 1938 f·~ 1.3 - 1.7 pale yellow fine gravel and sand: < 1 m depth author yolk Hubbs 1967 Becker 1983 f. phoxocephala 1.3 unknown s\ltift current ,over gravel: 15-60 em Page and Smith Stizoatedion ~ 1.4 - 1.8 pale yellow quiet wsters: 0.6-3.6 m depth author yolk Nelson 1968 Priegel 1969 ?~!'t,t" 8nct.P_r;9~~!1;~;;::!:~~?:~;.:~ .§..~ 1.9 - 2.3 psle yellow shallow water author yolk Nelson 1968 Seott"nd CrOSBlll8I1 1973 subgroupings may be formed: the johnny dartel"andspottaildarter·have smaller egg diameters than stripetai1 darter and striped fantail darter. Johnny dartersM.Ve :pa1~ yellow yolk,whiJespottai.J. dal"tel"shave. amber yolk. Yolk of the stripetai1 darter is orange while the striped fantail darter possesses yellow yolk. Eggs of the least darter, E. microperca and cypress darter, Eo. proe1iare are distinctinpossesslngco'Va""deep1:y~-±ndented on one side including the primary egg envelopes. Eggs<'of least darter are devoid ofpigllleJitwhi1eme1anophoresare those· of·· the cypress darter. Geography also separates he two species, with.1eastdarter occurring primarily in northern sections of thelJMRwhi1e the cypress darter occurs south entering Missouri and southern Illinois in the UMR drainage. Eggs ·..:of the northern 10gperch and river darter typically average 1.2 mm in diameter. The two speciespossess"d~fferent"coloredyo1k and deposit eggs in areas with differential current preferences. Differing habitat utilization by various species in the UMR basin is one means to facilitate identification. Darter species that inhabit the main channel border habitats of the main channel include the western sand darter, northern 10gperch, river darter, johnny darter, and>Iowa darter (Table 20). No information is presently available describing the western sand darter, or Iowa darter. The johnny darter possesses 15 preanal and 21 to 22 postana1 myomeres (Simon, unpublished data). Northernlogperch usually have more preanal and fewer postana1 myomeresthan the river darter. Northern 10gperch have a more slender and elongate body shape when compared to the river darter, and possess a differing pigmentation pattern. ~- ,.- ~ _.. Species which utilize northern stream headwater tributaries of the UMR basin include the least darter, striped fantail darter, and northern 10gperch (Table 20). The least darter and striped fantail darter each possesses 15 preanal myomeres which allows separation from the northern TABLE 20. Total length (mm) at hatchin~, yolk absorption, and fitst ape~raqce 'of fin ray formation in Upper Missis~ippii River perci ds. --- Yolk Spi~ous Soft absorp- dor'ssl dorsal Anal Pelvic: Pectoral Species Hatching tion fin: ray fin ray fin fin fin Literature AanocrIpia clara 4.1 5.0 SimC>n 1983 .Etheostoma asprisene 4.2 4.9 ------4.2 CUIIIIIlings 1983 !<. caeru~eu .. 6.0-6.2 8.0 > 8.0 7.8-8.2 7.8-8.2 9.0 7.8-8.2 Cooper 1979 -E. chlor~so, ..UIl unknown I E. exile I 3.4 Jaffa 1917 : !<. flabellare 4.5-6.5 7.8-10.0 5.9-7.8 5.4-7.2 7.0-7.2 8.8-9.2 5.0-7.2 author Cooper 1979 !<. kenni !. mi croperca 3.0-3.1 4.5 4.0-4.5 4.0-4.5 4.5 8.2 3.5 author Petravicz 1936 !<. nigrUIR 5.0 5.6-7.1 < 9 < 9 < 9 < 9 7.1 Fish 1932 !<. proeliare 3.2-3.5 4.2 4.2-5.4 4.2-5.4 6.3 8.4 4.2 I !<•• ~ unknown !<. squ8lli,feps 5.6-6.5 6.9-7.4 6.4-6.5 6.4-6.5 8.4 5.8-6. ----IE.zonale 5.0 7.0-8.0 11-12 11-12 11-12 11-12 i -IPerca fls!tescens 5.5-6.0 7.0 < 13 > 10 12 > 15 > Pereina d$prodes 4.5-5.3 7.0-8.0 10.6-15.6 i f.eYidesil unknown P.maculata 5.7 - -.--1-1 f .ahllll8rdi 3.6-4.0 5.5-6.2 5.9 5.9 f. phoxocephala unknown , Stizoatedion canadenae 4.6-5.1 8.3->10 > 10 > 12 12 > 15 > ~• .!.lli!!J!!!!. 6.1-6.8 8.9->10 > 10 > 12 12 > 15 > Nelson 1968 logperch which possesses from 19 to 24 preEl11a~DJY0m.~:r~(3. The least darter has a sDJaller. tapered yolk sac. and hatches at smaller lengths (3.0 to 3.1 mm TL) than the striped fantail darter (4.5 to 6.5 mm TL). The striped fantail darter has a large. spherical yolk sac, and yolk absorption occurs at greater lengths than in the least·darter. J:)epthDJeasurements of the striped fantail darter are also greater than in the least darter. In southern extremes of the UMRbssin. st:realll.· •• headwater species groupings include the striped fantail darter. stripetail darter;sp()ttail darter, rainbow darter (E. caeruleum), blackside darter, and northern logperch (Table 20). No information is presently available for the blackside darter. The northern logperch has the greatest preanal myomere count among the above species. The striped fantail darter has 15 preanal myomeres, while the spottail darter, stripetail darter,andrainbow.darter all have 16 ,preanal myomeres. The stripetail darter and spottail darter both have a/spherical, robust yolk sac, while the rainbow darter has a yolk sac shape similar to tha1:of the least darter. The yolk sac absorption in stripetail aIid spottail'datte:r occurs at smaller lengths than the rainbow darter as reported by Cooper (1979). Separation of darter from spottail darter larvae is based on head length/TL characteristics (spottail darter larvae possessed consistently larger head lengths), and differential pigmentation patterns. Identification of larval forms may be aided by examination of groupings of the species at a subgeneric level based on Kuehne and Barbour (1983) and Page {1983h= The· striped-fantaIl darter,- strfp-etaiTda:r1:er, and spottail darter are in the subgenus Catonotus; the least darter represents the subgenus Microperca; the northern logperch the subgenus Percina; and the river darter the subgenus Imostoma. - Adult diagnostic characteristics of the subgenusCatonotusinclude a supratemporal canal that is interrupted, an incomplete and straight lateral line,aridan interrupted infra()rbitalcafisl. Teeth srepresent on the prevomerarid palatine, a premaxillary frenum is present, anal spines number 2, and branchiostegalrays modally number 6 (Page 1981; Page 1983; Kuehne and Barbour 1983). Page and Schemske (1978) reported the relationship among adult darters of the 10 species' of the subgenus Catonotus. The E. flabellare lineolatum ~p~cies groupincludest:.ll~~triped fantail darter and stripetail darter, while the E. sguamiceps species group only includes the spottail darter among UMR species. They indicated that E. sguamiceps is the most primitive within thesubgEmus;whilethestriped .. ,.: fantail darter is the most advanced among the three UMR species. The reduction of meristic characters and fusion of ossified elements is considered to be an advanced condition among adult fishes (Romer and Parsons 1978; Burr 1978). Myomere counts, especially preanal counts, are important in considering the developmentalrelationships-among-manylarvae of freshwater fishes (LQQS and Fuiman 1977; Snyder 1979; Fuiman 1979). Results from the present study corroborates the phylogenetic claims made by Page and Schemske (1978). Both the spottail and stripetailc:lart:.ers possess greater preanal myomeres, and a similar number of post:.a~al ~YQ~eres to the "more advanced" striped· fantail darter. Thus the more prilIlitive species of the subgenus Catonotus possess greater numbers of preanal myomeres than the striped fantail darter. The prilIlitive species, spottail and stripetail (darters can be s~Par~!~~L 'based oILtbe ll~ac:ll~~_&-t:h/Th_cllarac::t:erj,§t:.:iS,and pigmentation. Spottail darter larvae consistently have. greater head lengths at equivalent total length intervals, and have less robust yolk sacs than the stripetail darter. Body pigmentation is sparce in the spottail darter, while denser in the stripetail darter. Yolk sac pigmentation is pale r 99 ! yellow in the spottai1 darter, but orangeinthestripeta:i:tdart.~r. subgenus Gatonotus possesses large sphericalJ()l~saC::f;!'l'I.(.~~to.42), precocious fin development, well developed pectoralf:ins with. incipient fin rays, and the absorption of yolk at greater lengths. The subgenus Microperca is considered the mos_t _advanced--of-the- genus Etheostoma.becauseof the reduced number of pored indicated that t~e cypress darter is the most primitive Microperca, that the fountain darter is most c10selyallied_to~_the __cy_press__dar_ter_,__and that 'the two form a species pair. The least darter is placeable iri a separate species g~oup with certain populations in various stages of differentiation, but most populations display the largest number. of advanced" characteristics. The least darter is the most advanced Microperca and the most advanced percid. The least darter and cypress darter (previously described by Simon 1985) hatch and both possess 15 preanal myomeres, and 19 postana1 myomeres. Yolk sac lengths are reduced, while body depth and greatest body depth is less than the Gatonotus subgenus. The cypress darter and least darter are similar for most morphological and meristic characteristics. Separ~ti9nof the two (species is possible since least darter larvae possess considerably more pigmeritscattered over the body than do cypress darters. Least darters. also have me1anophores posterior of the mandible while cypress darters do not. The subgenus Microperca thus hatches at the smallest length of UMR basin percids, possesses the fewest total myomeres of the species studied, r I and has a reduction in the ... morphometric ... measurements: 'bodydep'th/TL, greatest body depth/TL;and has a smaIieryolksac/TLF.-23=-:-28): The subgenus Percinaincludes five described and several undescribed species (Thompson 1978). The subgenus Percina is characterized by the presence of a distinctive conical snout projecting well beyond the upper jaw, a large interorbital width ,--separ~i~---~i:o~~tignt:ly-conn:ected branchiostegal membranes, a wide counts (Pa2e 1974a: Pa2e 1981; Information on larvae is limited to and the logperch, P. caprodes. All preanalmyomere counts. The logperch from 19 to 24, ldth 18 to 24 postanal myomeres, whilethebigscljlelogperch posseflSeS Uiyomere counts of 23 to 27preanar-an(r-1t.t;6J.-spcfsian~(Wang 1981). Since the northern logperch represents the the subgenus in,' the UMR it can be easily identified from any other species encou.ntered since the. body is more slen4eL-'gre~.te.~~J)ody_~t~p.tJ~l~is·.1O to .12), caudal peduncle depth/TL is also smaller for any similar length interval, and a sparce amount of pigmentation is present located primarily on the yolk sac and mid ventral postanal myomeres. Anessentiallysiinila.r pigmentation pattern exists for bigscale logperch from California (Wang 1981). Thus, larvae of the subgenus Percina tanbetharatterizediashaving a slender body depth/TL (.10 to .12), considerably higher preanal myomere numbers than most other darters, and pigmentation limited to the and mid ventral postanal myosepta. The subgenus Imostoma contains five species. Adults are characterized' by a mostly scaleless belly, usually no premaxillary frenum, separate to slightly connected branchiostegal membranes, interorbital width greatly reduced, and length of second dorsal fin base is over 70 %of the length of the first dorsal fin base (Page 1974a; Etnier 1976; PageI976)~ Theriver darter is theoIlIYlllelll1:u:~r of the subgenus Imostoma .... inhabiting the •UMR basin. A .. distillct pigJDentation pattern exists with an oblique line of melanophores exteIldiIlg posteriad from the anus. The only additional informationon the subgenus is the data obtainedfromthesIlail-darter,P ~ .. tanasi (Starnes 1977; Hickman and Fitz 1978). The snail darter possesses 16toT7preaIlal·myomeres, and 19 to 21 postanal myomeres, however illustratiollsprovided by Tennessee Valley Authority indicated 18 preanal mYQmeresand 21 postanalJllyomeres. Similar results were observed between darter and snail darter morphometric parametersfQr length intervals reported by Hi~kman and Fitz (1978). The Imostoma subgenus can be characterized- by: an intermediate preanal myomere number (18) ... -between the subgenera Percina, CatoIlotus, and Microperca; body slender with greatest body depth/TL (.12 to .14) greater than that of the subgenus Percina; and in comparison to the subgenus Percina, greater head depth/TL for any length interval. Previous studies have investigated the larval systematics of eastern subspecies of barred fantail darter, E. flabellare flabellare, and logperch, P. caprodes caprodes (Greeley 1927; Fish 1932; Lake 1936; May and Gasaway 1967; Taber 1969; Hardy 1978; Cooper 1978; Cooper 1979;iA.ueri1982). Results from the present study were consistent with that Presented in previous studies, however several differences were observed.• The fantail darter subspecies present in , li.!. lineolatum developes similarly to the eastern subspecies, E.f. flabellare. The total length of newly hatched larval lie !. lineolatum from the UMR were most similar to Ohio specimens of E. !. flabellare from Cat RUIl with lengths of 4.5 to 5.9 mm. Cooper (1979) reported that total myomeres of E. f. 102 f1abe1lare ranged from 34 to 36, and included 15 preanal and 19 ·to 21 postana1 myomeres, with a mode of 15 preanal and 21 postana1. The range of -E. -f. lineolatum lies within the rangeofE.Lf1abellare,. '." _._. however the mode is 15 preanal, and 19 postanal myomeres. Yolk was reported to be absorbed by 7.0 rom in New York (Lake 1936),andby. 9 to 10 rom in Pennsylvania (Cooper 1979). Specimens of E. f. liIleo1attiIn from the UMR had completely absorbed their yolk sac by 7.8 to 8.0 DIDl. )pi..nousdQrsal rays fi:rfJtapp~f1:r~d.i..IlE.t. f1abellare by 7.8 rom (Cooper 1979), while 5.9 to 6.2 mm E. flabellare lineolatum had deve1opedspiIlo'llsdorsal rays. Likewise,E. f. flabe1lare had soft dorsal fin rays developed by 7.2 mm (Cooper 1979), while E. t. 1ineolatum larvae had developed soft dorsal rays by 5.4 to 6.2 '··~mm. The only other variation thst exists between the two subspecies is the appearance of fin rays in pectoral fins. E. f. flabellare did not develope fin rays until 7.2 mm (Cooper 1979), while E. t. lineolattim developed fin rays by 5.0 to 5.6 rom. Pigment characteristics are essentially the same between subspecies..:Var.j.ation-in~.egg.diameters existed .with E. f. flabellare ova averaging 2.7 rom and ranging from 2.5 to 2.9 (Cooper 1979). Scott and Crossman (1973), andWinn(1958a) found egg diameters of --E. f. flabellare to average 2.3 rom. Ova ofE.f._.•...... - lineolatum averaged 2.4 rom but ranged from 2.0 to 2.8 rom. Geographic variation is not unusual in egg complement and egg size amongdart.ers(HubbsI958; Hubbs 1967; Hubbs and Delco 1960; Hubbs, Stevenson, and Peden 1968). However, the egg incubation time reported for E. t. flabellare by Lake (1936) was conSli..r1~:rf1J>lYll~ghe!_._!:h~I\1:.hflt obseryed ~Ilt:!J:~ .p!:~st:~t: . ~t:~'!x~~~Q"VI3.()f E. t· flabellare hatched in 30 to 35 days at 17 to 2QG,whileE. f. lineolatum eggs hatched after 9.25 to 10. days considerably less . than the eastern subspecies. Eggs of E. t. flabellare incubated at 23 C hatched after 14 to 16 days while E. 1. lineolatum eggs hatched in 5.5 to 6 days approximately ten days sooner than the eastern subspecies. The developmental sequence of events were similar between subspecies of the logperch, P. £. caprodes and P. £. semifasciata.Nortbern logperch. from theUMR hatched from 5.4 to 6.3 mm which agreed with results from previous studies (Taber 1969; Hardy 1978;CQQpe't1918JPain~~ taml·Ralon 1984). Cooper (1978), Fish (1932L Hardy (1978) ,and May and Gasaway (1967)reportedP ~ £. caprodes totalmyomeresto range from 41 to 43 including to preanal and 17 to 22 postanal. Paine and Balon (1984) found P. £. semifasciata to possess 23 preanal and 20postanal.myomeres in Lake Erie tributaries from Ontario. In the UMR, P. £. semifasciatahad 19 to 24 preanal myp~eres, and 18 to 23 postanal with 39 to 46 totalmyomeres. These results agreed with previous find.ings.from~1:.h~~aboveauthors. absorption occurred in P. £. caprodes by 6.4 mm (Fish 1929,1932; Hardy 1978; Cooper 1978), although some individuals possessed yolk at 6.9 mm. Yolk absorption in P. £. semifasciata from Lake Erie tributaries was 6.2 mm (Paine and Balon 1984). Yolk absorption was complete in all between 7.0 to 8.0 mm in P. £. semifasciata from the UMR. Pigment characteristics were essentially the same between subspecies, as..itwas for all members of the subgenus Percina that are currently Development of dorsal fin rays, pectoral, and pelvic fin raYs i.Ili.tiated at 15.6 mm in P. £. caprodes (Cooper 1978). Fish (l932)·andHardy (1978) reported dorsal fin rays initiating at 12.1 mm in P. £. caprodes, pectoral rays at 15.0 mm, while pelvic buds appeared at 14.2 rom. Development of fin ,~ ---..- rays initiated in the soft dorsal at 10.0 to 11.5 mm in UMR P. £ .. semifaciata, while pelvic fin buds appeared at 10.6 mm. Agreement exists between subspecies reported anal fin rays appearance. Significant differences existed between reported fin ray initiation in the current l r study and those previously reported ·by ·Cooper (1978). Clearing and I staining techniques were employed by Cooper (1978) so that development ()f structural characteristics should have been detected earlier. Reported I lengths· ·of ·appearence considered only ossified rays in Cooper's I study. However, rays composed of chondrified material were not I differentiated from ossified lepidotrichia from the UMR. i for some variation between reported length at appearance. genera of darters common to the UMR, Percina and Etheostoma, can I be differentiated based on the followirig criteria.. ThegerillsEtheostoma I possesses well developed pectoral fins, functional maxillarya.rid Dl8Ildible, I a greatest body depth/TL > .14 (at all lengths)·, preanaLmyomereslessthan 18, body depth ~t anus/TL > .080 (at lengths lessthan ...8.• QJ.nm)" and I caudal peduncle depth/TL > .038 (at lengths less than 8.0 mm) •. The genus I Percina has weakly developed pectorals or fin buds, undeveloped I I nonfunctional maxillary and mandible, a greatest body depth/TL < .14 (at I ! lengths less than 8.Qmm) , preanal myomeres---equal--to~()];,~greaterthan18, I bgc:!.yc:!.c:!ptbat:: aIlllSl/TL... < .080 (at lengths less than 8.0 mm), and caudal peduncle depth/TL < .035 (at lengths less than 8.0 mm). CONCLUSIONS 1. The genus Etheostoma possesses well developedpectoraLf1ns, functional maxillary and mandible, greatest body depth/TL > .14, preanal myomeres less than 18, body depth at anus/TL < .080, and caudal peduncledepth/TL >.038. 2. The genus ··Percina has· weakly developedpectoiaIs~;·~ or-~~in Duds, undeveloped and nonfunctional maxillary and mandible, a greatest body depth depth/TL<.14,preanElI myol1leres equal to or greater than 18, 3. The subgenusCatonotus, as designated by Page (1981, 1983), is verified by the larval.pntogeny. The primitive species E. sguamiceps and E. kennicotti possess 16 preanaL and 18 to 19postanal myomeres. E. flabellare '''''''-'''''''''.'''''''''''''''''''''''.''.'-'''.'~-'-''-''''-'-'--''-----'''''''''-,-~-'''-~,-~,._,,~,--,-,-"~~~. ------lineolatum has 15 preanal and 19 to 21 postanal myomeres. Thesubgenus Catonotus possesses large spherical yolk sacs/TL (.32 to .42), precocious fin ray dev~lopment, well developed pectoral fins with incipient fin rays, and the absorption of yolk at lengths greater than the initiation ray development. 4. Individuals in the subgenus Microperca hatch at very small lengths in comparison to other Etheostoma, possess 15 preanal and 19 postanal myomeres, and have a reduction in the morph()Du:~tricmeas\lrelIleIlts(body depth/TL, greatest body depth/TL) and have a smaller yolk sac/TL (.23 to .28). 5. In the subgenus-Percina larvae have a slender an 6. In the subgenus Imostoma there are 18 preanal myomeres, the body is 105 106 elongate and slender but. body depth/TI..i,f3g:r~fl~~:r!:I1ElI1!:I1El!:~~()~t:he subgenus i Percina (.12 to .14), and there is a greaterheadlength/TLthan in the I subgenus Percina. I I 7. The present subspecies descriptions were compatible with previous studies conducted in other geographic regioris~:~::~~~~~~---~~~---~~~~·~···~ I The formation of a key to species for the family Percidae from the UMR i is premature. Adequate information is lacking for most species. I ~ ~ LITERATURE AUER,N. A. 1982.FamilyPercidae, perches•. PR. 581...648.!!!. •..•..•••. N.A. Auer (ed.) 1982.•. Identification of larval fiElhes of the. Great Lakes basin with emphasis on the Lake Michigan drainage. Great Lakes Fishery Commission, Ann Arbor, MI 48105. Special Pub. 82-3:744 pp. BAGENAL, T. B. and E. BRAUM. 1971. Eggsf)1l~~f)I"±¥±~f:~historypp.~59 __181. InW. E• .Ricker (ed.) 1971. 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