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pdm ''

-ECOLOGY OF THE

PIRATE FERCH (TITLE)

i

BY . I Leslie : I D Frankland

THESIS

PA T A FULFILLMENT OF THE SUBMITTED IN R I L REQUIREMENTS I D G E OF FOR THE E RE

H. s. in Zoology THE GRADUATE SCHOOL, ILLINOIS UNIVERSITY IN EASTERN I CHARLESTON, ILLINOIS 1977 '! YEAR I / /

I H EBY RECOMMEND FULFILLING ER THIS THESIS BE ACCEPTED AS OF THE R DUAT ABOVE THIS PART G A E DEGREE CITED

¥ D AU VISER 51(1.lf??} s 2«?/lf?7

DEPARTMENT HEAD 'l'he irate perch, Aphredoderus sayan s, Gilliams , the on y iving p u l pecies of the family Aphredoderidae, ranges thr ugho ut the Mis si ippi o ss alley and the Atlantic Coastal Plain in low gradient stre ms, swamps, a xbow and other backwaters. External haracters of the pirate have s c perch een described in deta.il by ID1dy and SUrber (1943), Troutman (1957) and

flieger (1975). Literature printed on the p r e er h limited with i at p c is part of histo y till unde scr bed. ny s its life r s i Food habits of the pirate perch have b en alluded to but not closely e ocumented. Abbot (1861) was first to describe ts fo od h bits by noting i a con umption of small fish o ernight by pirate perch from his aquarium. he s v tudie by Becker (1923) and Flamer and Woolco t (1966) describe the s t irate perch as pred a eou s, feeding almost entirely upon small, aquatic c nsects and phipod . (1949) describes its food a small aquatic aru s Clark s nsects , but small the pirate perch's diet. also includes fish among edin act vity patterns have recently been studied by e g behavior and i arker and Simco (1975) and Ba r o ne (1973). t review of the literature on pira te perch reveals very little about A ts age and growth rela.tionships. Hall a.nd Jenkins (1954) examined s ale s c 82 pirate perch collected in Sub Prison Lake, Oklahoma. Swingle rom 1965) a.ged pirate perch from but considered his age regre s on 44 Alabama, s i s invalid. Growth studies have been ca ried out Man uet (1963), who r by s i oropared growth stages to locations and pine number . anal vent anal s s ogue, et.al. (1976) have described morphological changes during growth ;Jira.ta perch. f post-larval Y.ovement of the vent from the anal to the jugulsr or thoracic position

well documented (Jordan 1878, Jordan and Everwmn 1896, s Forbes and 1920, Jenkins 19.54, and Mansueti 1963). Variations ichardson Hall s.nd

:153383 -2- n vent location have given rise to several nominal species and genera

� Jordan and Gilbert, 1882 and Nelson, 1876). An indepth morphological study of vent movements and anatomical changes ws s conducted by 11.ansueti

:196J). She noted that variations existed the rate of vent movement in 3.nd suggested that additional data were need ed to establish the relation­ ship between vent movement and age, seasons, reproductive status and spatial distribution.

This study was conducted to examine in detail the following specific areas the pirate perch's ecology: food habits, age, growth, sexual of tnaturation and anal vent migration.

METHODS

Pirate perch were collected with common minnow seines of 6.35 mm

�esh size. The collection period extende.d from through September April pf 1974 and February through March of 1976. Specimens were placed in

�traight formalin for killing, then transferred to 10� formalin for preservation. Stomachs were removed the laboratory their contents in and plassified according to Pennak (1953). Numbers of organisms were re­ porded for ea.ch pirate perch along with each category's percent of total

�umbers and frequency of as described in Lsgler (1956). Feeding occurence :>ehavior was studied by direct observation of pirate perch seined from 20 3reasy Creek in County, Illinois on rch 1, 1976. Pirate perch Coles Ma '"1ere phced in a twenty-gallon aquarium which a. substrate of contained detritus taken from the collecting site. Amphipods and small criuck and

:iecopods (cra.yfish) were placed in the aquarium periodically over a tw:>­

:.reek period and responses were observed.

The pirate perch's ctenoid scales were removed from area midway the t>etween the anterior region of the dorsal fin and the . The -3-

scales were mounted in CMC9 mounting on glass slided and examined at fluid der a h se contrast c pound scope. Age was det in by counting lOOX un p a om erm ed �nnuli distinguished by separation of circuli the anterior field a.nd a a in �utting a.cross of c c i in the lateral field. Scale l ngth and annuli ir ul e length were dete e with a micrometer. The sea.le and annuli lengths rmin d 3ombined with total lengths were ub ti t the formula ln=(Sn/S)L s s tu ed in "'or determination of perch total lengths at the st g o g pirate end of pa r win seasons ( La.gler 1956).

Regression lines were established from body growth to caudal fin

growth relatio s by linear regression equations. With this n regression ..ine, total length could be c n e t to standard length compa.rison o v r ed for With other studies. Sex gonadal conditions were determined by in- and �ernal e ina ion of the Sp ng xam t gonads. awni times were established from �hese gonadal examina.tions and the first young of year collected .

Preserved pirate perch in colle tions from Illinois Natural History c Survey, Urbana, Illinois, and Eastern Illinois Uni e sity, lest n, v r Cha r o Illinoi , were examined and measured with dial calipers. Measurements s • recorded in millimeters from preserved and collected specim ns were e as "'ollows: total length, standard length, snout- ent length a.nd v isthmus

DESCRIPTION STUDY AREA OF Specimens collected for this study were seined from Village an::i West

�illage ID:lwards Co t and from Greasy Creek in Coles Illinois in un y County, ,Figure 1). Streams and of low radi nts with little ditches were g e -4- water flow. Ditch bottoms contained large amounts of muck and detritus making seining very difficult. During summer months aqua.tic vegetati n o would clog the streilms a.nd choke out the sun's rays. Fish collected with pirate perch are listed Table 1. in

Table 1. Fish collected with pirate pe ch from streams in Coles and C r Fdwards o ties , Illinois. un

Erimyzon oblongus Western creek chubsucker Pimephales Bluntnose minnow Semotilus atromaculatusnotatus chub Not§mieonus crxsQleucus CreekGolden shiner N2tropi� umbratilis Radf shiner Notropis �hipplei Stealin color shiner Hypognathus nucha.lis Silvery minnow mirabilis Suckermouth minnow PhenacobiusCyprinus carpio Carp Esox @.!!!!i!ricanu� Grass pickerel Ictalurus melas mack bullhead Icta.lurus natalis Yellow bullhead E'undulus Bl.ackstripe topminnow Gambus� nota.tus fish affinis muegill I.,epomi�Lepomis macrochiruscyanellu s Green sunfish Lepomis megaloti§ Longear sunfish Ydcropterus Largemouth bass salmoides darter Etheostoma ni�rimgracile JoSloughhnny darter EtheostomamacuJ.ata Blackside darter Percina

RESULTS

Diet Feeding &bits And Captive pirate perch were nonactive fish; they would lie motionless in different positions for hours. Their infrequent movements were rather sluggish, seem g only to drift the bottom or ea by Most in to n r vegetation. remained in debris on the or a.mong vegetation, sometimes forming bottom close groups of up to eight fish. -s-

''" '·A_..,Q .; ... ';,,•

�reasy Coles 1. ' Figure Creeki County, D.linois lll rate perch • which p were found ' .

• -6-

Activity increased when food ite were introduced in the aquarium. ms were never seen rapidly pursue their intended prey but Pirate perch to slowly drift toW8rd the m v an sm Feeding movements consisted to o ing org i . of quick, sideways m ve ents of the head accompanied the o m with flaring

the gills. The gill flaring see for111 s. vacuum drawing the of med to prey the debris t perch would readily cap­ and o r in o the mouth. Pirate prey that moved, but pass over prey that remained motionless. ture would Eighty-three of the 100 pirate perch stolll8chs examined contained food different types organisms were present ( Table organisms. Eleven of 2). Chironomidae frequented more pirate perch stomachs (58�) a d made n the greatest percent of total numbers. (47%). Isopoda with up followed a frequency of occurence of 45% and 24� of total numbers (Table 2). The 1-20 mm 21-40 mm size pirate perch Table 2 ut lized and groups of in i the smaller prey (Copepod Cladocera) while larger groups, a and the size 41-60 mm and 61-80 mm, preyed more heavily on the larger food organisms.

(Hemiptera and Ephemeroptera ) .

Gri>wth Sexual Z.la.tux:ity Age, and hundred pirate perch were c llect from April through September One o ed of 1974 Cole� and Counties, and sixty more pirate perch were in :&lwards collected February through March of 1976 Coles County. The sge in in distribution of pir8te perch collected in 1974 is represented in T.11ble 3 and the 1976 pirate perch in Table 1974, 76% of the pirate perch 4. In collected were one year or younger in � ge (Table 3) and 1976, 67� of in the pirate perch were to be in the one year class ( Table 4). The found regression line established from the total and standard lengths was found

to be = 1.2X + 6 (Figure 2). Pirate perch were found to grow most y rapidly during their year of life, reaching 63� to 68% of their first -7 -

Table 2. Frequency of occurrence, percent of total numbers and total of food organisms present in four size groups of numbersparch col lected in 1974 from Coles and Elhmrds Counties, pirate

= n b r per group. N um e

fish in mm Percent 7• Size GrouEs SL . Freq. 1 32 � 37 Total N:' Food Occ. 1-20 21-40 41-60 61-8014 Group Numbers

Chironomidae 58 7 123 23 4? 200 47

Isopoda 4.5 . 11 33 47 10 101 24

Copepoda. 17 27 -- -- 10 14 41 Hemiptera 14 -- -- 17 24 41 10

10 -- 10 4 1 15 3 Cole�ptera 6 -- -- .5 -- 5 1 Odonata .5 -- 1 2 1 4 1

5 4 2 -- -- 6 Cladocera 1 Ostracoda -- -- 8 -- 8 2 4 Ephemeroptera 2. -- - - 4 1 .5 1 [ish 1 ------1 1 L2� • Total No. Organisms 49 183 110 85 427 100+

I -8-

Table ) .Age groups of i a e perch . l e and calculatedColes and totaT lengthsCounties, p r t 1974. co l cted from F.ciw::i.rds Illinois in

.Age Average length of ;£:ear Group TL mm CBlculated TL at �nd &nge NO. 0 1 2

·Young yr. 27.5 14 . 0-41 . 0 37 28.0 ------Of I 56. 5 37. 5-81. 5 39 ---- 41.6 ----

77.1 62.0-98.0 24 ---- JZ.� 621z II .AverBge Le g h: 28.0 39. 62.7 n t

Table calculated of 4. Age groups and Coles C un total, Illinois lengths 197pirate6. perch collected from o ty in

Ag Average Length end o e mm NO. 1 C81,s;:ulated2 TL3 at 4 f :2:2"-r Group TL Range .5

62.7 52.0- 87.8 40 60.'-l ------I • . .

II 91.4 79 • .5-105.2 8 66.1 89.4 ___ .. ------

III 95.8 8).4-117. 3 10 46.4 75.4 94.9 ------

IV 96.8 96. 8 1 38.6 66.2 90.4 96 . 2 ----

133. 7 1JJ.7 1 6z.1 22.0 111.s 12411 _132.2 v Average Length: 55.7 81.5 99.0 110.2 132.5 -9-

TL

J.l,t.O ----

20 s 36 40 ;b 60 76 80 90 100 0 10 110 The (mm) standard Figure 2. (mm)regression 160 lineperch or total length to length of pirate from Coles a.nd Eawards Counties, Illinois. -10- total length. They reached the second year 89 the third 82 and year.

Examinations gonads from pirate perch age group one (Table J) of. in indicate that sexual maturity be -established at growing sea.son one. may However, all pirate perch of age group one did not contain mature gonads indicating that some do not reach sexual maturity until the end of the second year. A one year male 47 mm total length had milt one year and. a !ems.le 49 mm total length contained ripe roe.

1974 the season pirate perch in &!wards County was In spawning of established as the first two weeks of Y.lBy. Females collected on .April 2J showed extensive egg development, and females collected on May 5 exhibited egg movement into the duct. Arter YJB.y 5, no developed eggs were found

fetr!ale pirate perch. Males collected during the last week of April in and the first week of contained milt the urogenital duct. The !i'JB.y in first young of year pirate perch, which measured 14.4 mm TL, was collected on Y..ay 29.

Vent Yaria tions

total of 160 pirate perch were collected Coles .A in and EHwards Counties; an additional 421 Illinois pirate perch and 138 out-of-state

pirate perch were obtained preserved collections. Isthmus knob- from vent lengths increased from April through the s\lll'.mer in the three size

classes ( Figure J). Although the isthmus knob-vent length means in Figure J indicate a seasonal cycle, variations within each monthly sample

have sufficient overlap to prevent significant findings (Figure 4).

Isthmus knob size of spring collected pira.te perch wa.s found to be greater

than summer collected pirate perch. Photographs of isthmus knob-vent

lengths and isthmus knob size a.re Figures .5 6. shown in and IK-VL

.s.o

4.5

• • �.o •

-'" • • • • • 3.5 • ,,.- .. / .... ti' • I .... ,, ..... • ,.., ..... • • / / .... "" 3.0 ' ,, I /. �"" /

• / 2 .s ,· --/

2.0 ..;.;..._ 'J'/.S mm minim.ull one TL am greeter, one year • • • • mm greate:r average year l .5 SO. 0 1lllll TL a.t.d , - -- 71.• 0 mm 'II. am greater, avars.ge onetwo years ·out,..ot *'* 'J? .5 TL and greater, minimum y�r, state l.O t �

• .s

o.'--��--���--��--���--��-- �- - � � � � M M -�

Fig�re ). the mean isthmus-vent (mm) three size pirate Graphs of size lengths states.of groups of perch f":"otn Illinois a.nd one group from other IK-VL ' ' t -7.0 6. · 5 � ·,

"6. o.

;.5

5.0

• 4 .5

4.o ' ...... • N j .5 . I I ).0

• 2· .5

2.0

1 • .5

1.-0

.5 o.e

· Feb Mr Apr "t;ay Aug seti Oct ?��v (41) (36) ('.36) (10) Jun(33) Jul(89) . . (68) (80) (10) (4)

Figure SeBsonal comparisons kno�-vent lengths pirate perch 37.5 mm 4. of isthmus of Illinois. standard deviation is represented by TL and lergerThe from means are 0;.1e by the line. the range by the vertical lines. shown -13-

:t \ f I '� ' ·' . ( -� · \, , . .L "'. t )� 1) I 'I / ., \ I

� -� ·, .. I, ·,

l1lrcn F F M July M 1 1 2 2 F1 M 1

Figure 5. of six perch collected Ventral view County, Illinois. pirate and month collectedfrom Franklinsre shown below each fish. Sex, age -14-

; . "

?-,• Figure 6. Ventral view of among pirste perch. pirate perch isthmusthe knobleft differencecollected The on wa.scollected inJuly March, in while locality.the specimen on the right was in the same -15-

DISCUSS ON

and H�bits Diet Feeding

'lhe pira.te perch is a highly pred B ciou fish , e in entirely on s f ed g

smsll aquatic orga.nisms. Feeding be avior studies by Parker and Simco h (1975) coincide with findings of this · study. '!hey describe feeding be-

havior as a slow indirect approach, ut with quick t rns of the head b u and a ega tive pressure formed by he gill cav�ties to suc t e prey. n t k in h

They a.lso found the pira te perch to be most active night and restricted a.t to bot om areas. t Chironomids were found ira pe ch stomachs and made in more p te r up greater numbers than any other a i ms (Table 2). Becker (1923), org n s Clark (1949), Rice (1942) and Flemer and Woolcott (1966) also found

chironomids preser1t in pirBte pe ch stoma.chs. Flamer and Woolcott (1969) r listed chironomids as the most frequent (.55'%) and ma.de up 53% of the

total organisms. Forbes (1888) estBblished the diet as ma.inly dipteran

larvae.

Fish p e ation pira.te perch is insignificant a d r d by n limited to only a few large i vi a l pirate perch. Fish eil18 n have been select nd i du r i s taken from pirate perch stomachs (Forbes and ch r son Ri a d 1920 and Cla.rk 1949). 96 However, from this study a.nd Flamer and Woolcott (1 9) , fish

were found to make up only a very small rcen a e of i t (Table 2). pe t g the d e Prey size was found to with i a t perch size (TFJble 2). vFJ.ry p r e .As xpected, small pira ta perch utilized the smaller Cladocera. a.nd Cop poda e e

while larger pirate perch preyed more heavi on he ly t larger Hemiptera Ephemeroptera. Prey size se ec io is probably a. food and l t n result of preferability to the pira te 1! source availability, a.nd vulnerability 11 perch size. I ·

===tt======t== � -16-

Prey utiliz by pira.te perch consisted almost entirely of small ed insects crusta.cea.ns w i h live within the muck a.nd vegetation aqustic and h c of the dimly-lit bottom. Observations from this study indicate sight

is used ca ture , but because of the murky waters in prey p and nocturruil habits , other senses must be used ( Parker Simco 1975). The well­ a.nd developed la.t ral senso pores of the head, as described by e line and ry :YlC>ore Burris (19.56), are probably the main senses used in prey lo­ a.nd cation and capture.

Age. Growth and Sex\Ull ?-!aturity

Pire.te perch are reported as spr n spawners (Abbott 1862 and El:ldy i g Underhill 1974). In Texas, and Hubbs (1973) esti:1blished and ?-19rtin spawning t es as February. Findings from this study and Forbes im late Ric s on (1920) indica.te that pirate perch in early May and hard spawn at the latitudes of Illinois. 'Ihe age and size at sexual maturity in pirate perch have been shown by Mansueti (1963) from Hall and Jenkins

(1953) a.s 55 mm tot.81 length and one year of by this age. Findings study indicate that sexual m tu it is established in the 40 mm's and a r y not the 50 mm's as by }�nsueti. thought Considerable length variations among different a.ge groups are shown in Tables 3 and 4. Hall and Jenkins (1953) also had length variations and overlap Dmong age groups of irate perch. OklB.homa p Differences in age size groups of 1974 (Table J) and 1976 ( Table 4) studies are probably results of variations in len ths growing seasons g of and differences geographical locations. Growt rates established in h in this study for the one-year class (63-68�), class (82%) two-year and three-year clB ss (89%) are essentially the same as Hall and Jenkins -17-

3 who found for the one-year class, 3 for the two-year class

and 87% for the three-year class.

Vent V,qria.tion

Studies of pirate perch have shown much variation vent location in ( Jordan 1878 and Hall and Jenkins 1954). MB.nsueti (1963) stated that

the variation of the vent location according to various fish s ize in-

dica.tes a great va.riability migration rate. Figures 3 and 5 show in this variation, not of size groups, of a seasonal cycle. Fish from but these figures are considered to be at least one year of age and should

show no juvenile forward of the vent. Variations of vent lo- movement cations are definitely present and all results indica.te some type of

seasonal cycle.

Sea.sonal cycles are tied to spawning time and gonadal growth. Tne

small is -ve t lengths are found when the pirat perch contains thmus n e gonads near spawning condition. Increased isthmus-vent lengths in occur after the spawning sea.son. The greatest isthmus-vent lengths

occur during the summer months when gonads are at the least a.mount

of development. With the decrease of isthmus-vent length during the

fall , the gonads presumably a.re developing to the over-winter condition

as shown February (Figure 3). The change in of the in early size isthmus-knob to vent is limited anterior movement by the presence in of thoracic muscles ( ist hmus knob). Posterior movement is limited by

the complete ossification of the pelvic bones after the juvenile fish's

intestine has passed through the foramen ( � nsue ti 1963). ¥ Pirate perch estimated at two years and older show a seasonal

variation of vent location, ut not a.s pronounced as the one class b year ( Figure 3). The low numbers of olds examined does not allow I two year II -18- for significant speculations. However, decreases variations in may resu1t hardening of tissues with processes from linked the anatomical vent migration (Smith 1976). of 'lhe anterior os tion of the vent along with its variations could p i be most readily associated its fildy and Surber with spawning habits.

(1943) reported that is known about the pirat spawning litUe e perch's habits exce t that it was e ort to build a nest and guard the eggs. p r p ed

Pflieger (1975) postulates from the l cation of the vent on the throat o that the rate perch is actually a incubator. This seems pi bucca.l likely since the anterior position occurs two families of of the vent in only fish, .AphredoderidBe. The .lllnblyopsidBe or the . and fishes the perch relics of extensive fauna of the and pirate a.re a pa. st southern swamps ( Jordan 1905). The closest living r lat ves Aphre­ e i of doderirae ara considered (196?) Rosen Patterson (1969) by Rosen and and to the Amblyopsidae. Members of h s family are known buccal be t i incu- bators, but just g s become situated the gill mb rs is how the e g in cha e unknown (Weise 1957). An incidental observation by 219.rtin Hubbs and (1973) showed that eggs e ress artificially pirate perch tended xp ed from

to move e1ong a. roov into the gill chambers. This observat n may g e io explain the r,my eggs are introduced into gill chambers of the Amblyopsida.e. This also explain why the vent Aphredoderida.e and may

is located clo ser to the isthmus knob du.ring sea. son of the spawning pirate However, both females males have t e vent va.riation perch. and h characteristic. presence of the forwar3 location of the male's 'nle vent cannot explnined without further information on spawning ha.bits. be I behavior require the itself vent to vent on Spawning may Ill81e to place l fem.ale for fertilization to occur. Another explena.tion is that milt ! the -19-

may pass into bucca area. and then e elled into the fe­ the male's l be xp male's mouth. These a.re only speculations csn o y be and nl explained

more intensive stud es. Another external change in morphology as­ by i sociated with conditions the size of the isthmus knob. seasonal is Figure 6 definitely shows a difference of knob in March and July size n specimens. �rch specimens have an exte ded , swollen knob while July r s ec en a knob which is not swollen. This swollen knob be p im s ha.ve may used some type of spawning behavior in ritual. the pirate pe c a buccal then it should be If r h is incubator, found with the gill c e s as in eggs in hamb r spelaea. Eigenmann (1909) wrote that femBle can readily e distinguished by the appear­ the b ance of eggs through the opa ue gill covers . Eigenmann also tha.t q noted eggs freed from the gill cavities continued to develop, un­ readily interupted. However, the gi cavity provided such a unique and self­ ll regulated hatchery that the eggs in it. No eggs were found in relll8ined the gill cavities of 719 pirate examined during this study. This perch indicates that the pira.te perch release eggs as a type of may the species preservation. The eggs could develop naturally released then outside the gill chambers and ensure the survival of the species. -20-

ACKNOWLEDGEMENTS

would to thank Smith and no s I like Dr. Phil Larry Ps.ge of the Illi i for their o n of the specimens help with Natural History Survey l a and vent variations. I would a.lso like to Leonard thank Dr. Durhatn, Dr. Andrews Ea.stern Dr. E.O. Moll and Dr. R. of Illinois University for help and suggestions throughout study. their this Finally, I would to wife for he help in collecting specimens like thank my r and typing th manuscript. is -21-

LITERATURE CITED

.Abbott, C. 1861. Notes on the ha.bits of Aphred_oderus sayanus. Proc. Acad. Nat. Phil. 1 :9.5-96. Sci. Becke , H.K. 1923. The of A12hredoderus sayanus Kalamazoo r County, Michigan. Occ. Papus. Mls. Zool. Arbor,in Mich. 138. 4p. Ann

Bo tone, Stephen 1973. Pughea.dedness in the pirate perch with r plicationsA. on feeding. Chesapeake Sci. 13:231-232. im­

Clark, C.F. 1949. New eco s for pirate perch central western Ohio. Copeia 1949:r 2rd19-220. in

T. Surber. 1943. Northern fishes with special reference El:3.dy, s. andto the upper Mississippi Nineapolis, of Press. 2.52 Va.lley. Minn. thiv. Minn. p. El:3.dy, 1974. ort ern Fishes. Univ. Press. S. andMinn J.C.. 414 Underhill. p. N h Minn.

Eige:nmann, C.H. 1909. vertebrates of America. Carnegie Inst. Wa.sh. Pub. 104.Cave 241 p.

Flamer, and Woolcott. 1966. Food habits and distribution of D.A.the fishesW. S. of Tuckahoe Creek, Virginia, with special emphasis on the bluegill. esapea e Sci. 7:75-89. Ch k

Forbes, S.A. 1888. tudies of the food of freshwater fishes. State Lab. VolS 2 :443-473. Bull. Ill.

Forbes, and Richardson. 1920. The fishes of Illinois (2nd ed. ) S.A.Ill. Nat.R.E. Hist. 3:1-357. Surv. Hall, G. E. and Jenkins. 1954. Notes on the age and growth of the pirateR.M. perch, Aphredoderus sayanus, la h ma Copeiit 1954 :69. in Ok o . and L. 1976. f shes in the Tennessee Hogue, J., River.R. W�llus, Tennessee BalleyKay. Authority.Larval Technical i Note. B19. 66 p. Jordan, D. S. 1878. partial catalogus the of Illinois. Bull. StateA Lab. Nat. of 1(2):33-70.fishes lll. Hist. Jordan, D. S. 1905. The study of fishes. Henry Vol 599 Holt Co. 2. p.

Jordan, D.S. Ellermann. 1896. fishes North and Middle and B. /1. Nat. Kus. 47(1):1-1249.'!he of I America.. U.S. Bull . I Jordan, D.S. Gilbert. 1882. Sj-nopsis of the fishes f North ) m randc .C.H. Natl. Mus. 16. 10:l.8 o A e i a Bull. U. s. p. Ir -22-

=====11======;:======::#======- Lagl er, Karl F. 1956. Freshwater fishery biology. Wm. C. Brown Co. Dubuque, Publ. Iowa. 124 p. 3 Mansueti, J. 196 . Some cha.nges in morphology during ontogeny A. 1962 in the pirate per h , Anbredoderus sa;zanus. Copeia :J.55-360. c � M9.rtin, and C. Hubbs. 1973. Observations on the development of F.D.pirate perch, Aphredoderus sayanus, (Pisces:Aphredoderidae) with comments on yolk circulation patterns as possible taxonomic tool. Copeia 1973:377-379. a

?{core, Burris. 1956. Description of the la.teral-line G.A. and W.of E . system the pirate per h , ARhredoderus sayanus. Copeia 1956:18-20. c

Nelson, 1876. partial ca.t11logue of the fishes of Illinois. Bull. E. w.Ill. }fu.s. A Hist. 1 :33-52. Nat. Parker, C. Simco. 1975. feeding and N. haviandor B.A.of the pira.te perch,Acti Aphredodevity patterns,rus sayanus. Copeia be­ 1975:.572-.574.

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