INTERSPECIFIC AND INTRASP~CIFIC VARIATION BETWEEN AND AMONG SYMPHURUS PLAGIUSA (LINNAEUS) AND SYMPHURUS CIVITATUS GINSBURG WITH NOTES ON THEIR DISTRIBUTION
A Thesis Presented to The Faculty of the School of Sciences and Mathematics Morehead State University
·In Partial Fulfillment Of the Requirements for the Degree Master of Science in Biology
by Chester A, Nava, Jr. December, 1976 ,A.J->t'•"- Jl'f I n~~'-'"' 597091(, N 3/&;p
Accepted by the faculty of the School of Sciences and Mathematics, in partial fulfillment of the requirements for the Master of Science degree.
Master's Committee:
'Gerald1s "-"t .Deos 2~m,;..,,
- ~n,/ason. ryor
Woorow7Barber~r.->- t. /. Brr:tw
~ ~ 3 , / 'Jljb ate '
ii ABSTRACT
INTERSPECIFIC AND INTRASPECIFIC VARIATION BETWEEN AND AMONG SYMPHURUS PLAGIUSA (LINNAEUS) AND SYMPHURUS CIVITATUS GINSBURG WITH NOTES PN THEIR DISTRIBUTION
The species status of Symphurus. civitatus Ginsburg, as a· species distinct from Symphurus plagiusa (Linnaeus), has been doubted (Walls, 1975). Through interspecific analyses and observations, it was determined that S. plagiusa and S, civitatus. are, in fact, two distinct species. Important characteristics supportive of species segregation were fin ray counts, scale row counts, vertebral counts, and depth of capture, Symphurus plagiusa has slightly lower fin ray and vertebral counts than does ~- civitatus. _ The.scale row counts, a .reliable character istic for both immature and mature specimens, were higher in S. plagiusa than in S, civitatus. The orbital diameter, a reliable characteristic for adult specimens only, was greater in S. plagiusa than in S. civitatus. The caudal fin ray counts were usually 10 ins. plagiusa, and consistently 12 in~. civitatus, The caudal fin ray count was the single most important characteristic for separating the two species,
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J:il The depth of· capture was a helpful and reliable aid for separating the two species •. Symphurus plagiusa was collected in shallow.water (six fathoms or less), but was occasionally collected in deeper.waters. Symphurus civitatus was collected in offshore waters, and was never collected in shallow shoreline sampling. Symphurus plagiusa is distributed from New York State to Argentina; the species was divided into five populations for the purpose of intraspecific analysis, The range of S. civitatus overlapped the range of s. plagiusa in two areas. One of these areas was from the Atlantic coast of North Carolina to Florida; the other, and the largest area, was in the northern Gulf of Mexico. Intraspecific observations and analyses showed significant levels of geographic variation among populations of s. plagiusa;, however, all analyses made on s. civitatus populations were negative.
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v.t ACKNOWLEDGEMENTS
I wish to express my appreciation.to all those people who gave of their time and effort in support of this thesis, I would like to thank Dr, Jerry F., Howell, Jr,, who acted as chairman of my graduate committee. I also thank the other members of the committee, Mr. Woodrow W. Barber, Dr, Gerald L, DeMoss, Dr. Madison E, Pryor, and Dr. Robert L. Shipp, for their many hours·•'Of work, interest, and constructive criticism concerning this research. Still others who deserve recognition and thanks are Mrs, LeMerle Bentley for her many hours spent in helping to obtain the literature sought for this research; also Ms, Marilyn Stone, Mrs, Ana Angus, Mr. Steve Angus and Ms, Aggie Angus for aiding me with translations in the literature. Dr. Frank Osborne gave invaluable aid in assist ing with the statistical analyses. His time and effort greatly enhanced the conclusions of this research. There were many people ~ho aided by lending specimens, offering suggestions, and allowing me to examine specimens at their museums. I am very apprecia tive of the time and effort of the following people,
V without whose help required data could not have been obtained: Mr. Charles E. Dawson, Mr. Robert D. Nester, Dr. Herbert T. Boschung, Dr. Ralph W. Yerger, Mr. Gordon Cherr, Mr. AndrewJ. Leslie, Jr., Dr. Carter R. Gilbert, Mr. George·H,·Burgess, Dr, Gary F. Mayer, Mr. Charles R, Futch, Mr. John E. Darovec, Jr,, Dr. c. Richard Robins, Dr. William Anderson,- Mr. c. c. Koening, Dr, Frank J, Schwartz, Mr, F. C. Rohde, Dr. John A. Musick, Licenuada Graciela Lazzaro, Licenuada Hugh Castello, Mrs. May Tillman, Mr, Allen Lake, and Mr. Barton o. Kern. I wish to thank the administrative staff and the faculty of the Department of Biological Sciences at Morehead State University for the graduate assistantship awarded to me while pursuing the Master's degree, The roster of names has been long, but each has contributed to my research, ·My sincere apologies to anyone whom I may have omitted,
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Table Page 1. Frequency Distribution of the Number of Dorsal Fin Rays in Symphurus plagiusa and Symphurus civitatus ••••••• • • 25
2. Frequency Distribution of the Number of Anal Fin Rays in Symphurus plagiusa and Symphurus ci vitatus • • • • , • • , • • 25 Frequency Distribution of the Number of Caudal Fin Rays in Symphurus plagiusa and Symphurus civitatus ••••••• • • 26 4. Frequency Distribution of the Number of Scale Rows in simphurus plagiusa and Symphurus civitatus .••••• • • • • 29 5. Frequency Distribution of the Number of Vertebrae Occurring in SfD1Phurus plagiusa and Symphurus civitatus • • • • 32 6. Mean Value, X, in Percentages, of Standard Proportional Measurements in Symphurus plagiusa and Symphurus civitatus • • • • 35 7. Frequency Distribution of the Number of Dorsal Fin Rays in Symphurus plagiusa, Segregated by Zoogeographical Region • • 38 13. Frequency Distribution of the Number of Anal Fin Rays in Symphurus plagiusa, Segregated by zoogeographical Region • • 39 Frequency Distribution of the Number. of Scale Rows in Symphurus plagiusa, Segregated by Zoogeographical Region • • 41 10. Frequency Distribution of the Number of Vertebrae in S~mphurus plagiusa, Segregated byoogeographical Region • • 43 11. Mean Value, X, in Percentages, of Standard Proportional Measurements in Symphurus ~la~iusa, Segregated by Zoogeographical egion . . . , ...... • ...... 45 vii .11.:rr
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Independent "t" Test of Scale Row Count Variation between Symphurus plagiusa and Symphurus civitatus in Region III • • 77 Independent 11 t 11 Test of Vertbral Count Variation between Symphurus plagiusa and Symphurus civitatus in Region III • • 26. One Way Analysis of Variance to Determine the Variation in Dorsal Fin Ray Counts of Symphurus plagiusa Segregated.by Zoogeographicai Region • • 79 Duncan's Multiple Range Test to Determine the Regional, Variation in Dorsal Fin Ray Counts of Symphurus plagiusa •• • • 80 One Way Analysis of Variance to Determine the Variation in Anal Fin Ray Counts of Symphurus plagiusa, Segregated by Zoogeographical Region •• , , , , • • • Bl 29. Duncan's Multiple Range Test to Determine the Regional Variation in Anal Fin Ray Count:s of Symphurus plagiusa • • • • • • 82 30, One Way Analysis of Variance to Determine the Variation in Scale Row Counts of Symphurus plagiusa, Segregated by Zoogeograpliical Region • , ••••• • • 31. Duncan's Multiple Range Test to Determine the Regional Variation in Scale Row Counts of Symphurus plagiusa •••• • • 84, 32. One Way Analysis of Variance to Betermine the Variation in Vertebral Counts of Symphurus plagiusa, Segregated by Zoogeographical Region • , •••• , • • 85 ix tRft nJ~ lRnA 1o jasT njn jnebneaebnI .scu1:,'.!lslg BU'IlJrf(lmy~ nesw.:ied noJjf\J"lnV • • III ao1:~o51 rd: eujr,jJv .i:o err:mrlomv;8 bm'\ jm.10::l wo.H ,:,Jr,08 1o .:iao'T 11 j" vnebrreuebnJ na11.i:;,:,r;lq a.u ;tnuo::l J.s"Idj 0 wV 1o 0Be'1' ";:t" j·nsbns(!obnI n.?J!J~l sr:.tm'Iejefl oj oorrr.l•Ir,V 1o e leyisnA v_,;W errO a.:lrmo8 vnH nYo! 11:mA nl !loJ:jr;.i:"Ir.;V er!j v_d bejs~e~r::ie<: , neul;groig 0!Jur!glTly_s_ 1o • • • • , , • • • no.l:i'\sH lsolrlqr,'J;sos~ooS snllll'HJje(l oj j;:rn'f 0'i?,flBH elqljiuM El I rmonu(l v,c,J nl'il lsm, · rt.i: rm.i:js.i'IllV lGrm.i:;gol-1 orlj • • • • • • .§ZHf.;ngg_ au'rurlqmv.8. -1o eJnIJo8 . . . s11.i:m·1ej!DCI. o:t oortsi'IGV' 1o ale.v.JonL\ '[SW srrO .0( 1o a;lmrorJ wo.H eJ.no2 nl noJjr,J•rnV srl;t ,r.ci bed B; X X . . ~...\'OO((GQ;j;L\JblJTC\JJ W- "6~TOIJu . 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Symphurus plagiusa (above) and Symphurus civitatus (below) •••••••••• • • 2 2, Zoogeographical Regions of Symphurus plagiusa (found in Regions I, II, III, IV, and V) and Symphurus civitatus (found. in Regions- It III, and V.) , , • • 23 3. Caudal Fin Construction of Symphurus plafiusa (above) and Symphurus civ tatus (below) ••••• , •• • • • • 28 4, Distribution of Symphurus plagiusa , •• , • 58 5, Distribution of SYmphurus civitatus, • , , , 59 xi x:r • 2•J . • ~~ s~ s· s ;T' TABLE OF CONTENTS Chapter Page I. INTRODUCTION • • • • • • • • • • • • • • 1 Purpose • • I • • t • • t • • I I • • e 1 II. REVIEW OF THE LITERATURE. • • • • • • • • • 6 Distribution . • ...... • • 6 Anatomy and Morphology ••••••• • • • 9 Behavior and Ecology ••• , ••• • • • 12 III, MATERIALS AND METHODS • • • • • • • • • • • 15 IV. RESULTS • • • • • • • • • • • • • • • • • • 24 Interspecific Variati.ons • • • • • • • • • 24 Fin Ray Counts ...... • 24 Scale Row -Counts • • • • • • • • • • • • 27 •Orbital Diameter; ••••••••••• 27 Angle of the Mouth • • • • • • • • • 27 Teeth ...... • . • . . . . . • • 30 Vertebral Counts • , • • • • • • • • • 30 Branchiostegal Ray Counts and Gill Raker Construction, , •• , , • , •• 31 Body and Fin Coloration • , , , • , • 31 Proportional Measurements •• , •••• 33 Vertical Distribution ••• , •••• , 34 Statistical Analysis of Zoogeographical Region III ...... • . , . • 34 Intraspecific Variations ••••••••• 36 Symihurus ~lagiusa. , • , , , • , , • , 36 F n Ray cunts • . . . • • . . . . • 36 Scale Row Counts • • • • • • • • • • • 40 Vertebral" Counts , , • , • • • • • • • 42 Body and Fin Coloration • , . • • • • • 42 Proportional Measurements • • • • • 44 Symphurus civitatus ••••••• , •• 44 Fin Ray Counts . • • • • . • . • . . 44 Scale Row Counts • • · • • • • • • • • • 48 Vertebral Counts • • • • • • • • • , , 48 Body and Fin Coloration , • • • • • 51 Proportional Measurements •••••• 51 xii r .L • • • • 4 • • • " . . . . . • saoq"Ir.rY .,. 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III nol~eR "' . • • • enol.:t~l"'!sV ::>.i:11.!J0qes'1;tnI dC.~, r (.J(. • , • • ae1.J.i:).1J?,Ia ;;ar-rr.rrfo-mv8 ci( • • • • • • :- 8.:tf!JJO~ ~-sF n.t·if'-· O~\ • . . • • • • eJrr1Jo8 woH sino8 ~:.i • • , • , • a.:lrwo::i .[:,•rdsd"JeV S-:l • • nol.:ls-rnloD nl1 bne 1boB .µ~\ • • ajnsms~uessM lsnolj~oao~q ,1.;1 • • • • .. • • -----p,LJjBjlvJ!) --2u'1Llrlqmv2--- --• .. -:l1i • • • • • • • • e;Jrmo::J ys5l nl'1 tu • • • • • • • e dnuoQ v,oli ~IA~2 l~+\ • D • cjnuoD IR~dej~ev ,- :, .L L • rro .i:;J wrol o:) nl'>i brrn ·,:boH J: (1 • • • a j nerr.e'J!J8B eJ4 ferro l j-roqo,q .tlx TABLE OF CONTENTS (Continued) Chapter Page DISCUSSION • • • • • • • • • • • • • • • • • , 53 VI, SUMMARY· • • • • • • • • • • • • • • • • 61 VII. LITERATURE CITED • • • • 65 VIII. APPENDIX , • • • • • • • • • • • • • • • • • 70 xiii • • • • • • • • .v • • .IV o·· . . • • IlJx CHAPTER I INTRODUCTION Purpose The validity of Symphurus plagiusa (Linnaeus) and Symphurus civitatus Ginsburg (Figure 1) as separate tongue fish species has been recently questioned (Walls, 1975). The purpose of this research is to resolve this controversy, and add morphological and distributional data to existing information. Ginsburg (1951) reviewed the western Atlantic tonguefishes and described six new species, including Symphurus civitatus. In his account, he left some doubt about the status of Symphurus civitatus as a valid species, and recommended a comparative study of Symphurus civitatus and Symphurus plagiusa. Below are Ginsburg's (1951) descriptions of the genus Symphurus and the species Symphurus plagiusa-and Symphurus civitatus. - Symphurus: Rafinesque '~ •• Symphurus Rafinesque, Indice D1Ittiologia Siciliana, p. 52, 1810 (genotype Symphurus nigrescens Rafinesque by subsequent designation)--Jordan & Goss, Rep. U.S. Gomm. Fish. 1886:321, 1889 (Symphurus nigrescens Rafinesque designated as genotype). Sinistral, strongly compressed; symmetrically shaped; greatest body depth in anterior half of 1 ,10 I': '.l tJ(lO.\l'l HI bfl,, f P-us~•.,,1r.t.l) _A_2n.i:~"'Ig: .~. .t!.1,1.1rfgm'[C: 'lo 1ljlb.UBv er!T -eH:'.111c.:1 ejs .. rsqea a.e {I srru~.i:'~) 2}'!rJdznlD E!!..~fy.i:o 2. .r1--ru:io~~ .noJ.J.B111'!01 JJ: dduob emo::: j1sI erl , jrmooos a.i:rl nI , a1.1jaj i:v.i:o c.fJ'lrrrlr:r~.'@_ .. Elr.:!.~rJ5t~~l:. 1o Y,bLJjj:; ev J:j-s-r.eqmo:> ·.s beb;ieffuno:>o'J bnG , asl!)sqa ;; 1,,-,uderr1:1 e·rn wolsB .~ipnig_ CH-rrn!qmv2._ bm; 2-!.Ij~jj:vj:_£ :c. s.i:o-,:~e r;jj brr,:; cu-r11r!gm'{ supri sa.i:'ls.H .. : ?.!J'lllriqmve Bi::9.0J.0j:S-:II',I ,,ol:br([ ,supi3sn.i:1.afl 81J"!!Jrlmr1y2. , ••" -!~-"-;i:.i::-:"1.f ,,u-ru,krmvc: 9f!\'.jOfl9~) orsr-:-s"f ,q ,frn,;.iI.i::;.i:8 ,eao;, ,,', rrc.fnot,--=Ti'roi,1s113.l:asi:l :Jrrnupe~di.re '[d supcerri1sfl .'.)_1J"!.tr1i:1mv0) ~:ilRI ,IS:~:088.[ .rla.i:'iI ,rm1108 .c,IJ .qsR :-{ eq\;jorrs~ as bojsn8 J:esb su~een.i:'lsH anso2_c1_"f:0.rr: y.C.:.so 1:'1 :J sn!:tr<[e ; beee ertqmno yI~no-rja , [s,:je 111.i:G 1o ~ia:! ·ro.i:'JsJns n.i: rl.Jqsb v.hod ;tes.Jrrn,:~ ; bS'lf:ne .[ Symp hurus plagiusa CA N675- 03 Symphurus civitatus CAN775 - 09 Figure 1. srm~hurus Tlafiusa (above) and Symphurus c vitatus be ow) 3 length; the depth nearly uniform for a considerable distance, thence tapering both ways; the anterior taper moderate; posterior taper beginning at about end of anterior two thirds of standard length, varying a little.both ways, except in jenynsi at , about the middle, the depth decreasing rapidly to caudal base. Eyes small, slightly smaller than short snout; the two eyes separated by a very narrow interorbital space in small specimens touching each other or very nearly so in the larger specimens; anterior margin of upper eye slightly in advance of that of lower, infrequently both about aligned on same vertical. Snout short and very blunt, the anterior profile forming a nearly continuous broad curve.. Mouth. very small, its posterior angle under anterior margin of lower pupil, .varying slightly both ways, except slightly more backward in civitatum and pla~iusa; asymmetrical, moderately curved on eyed si e, notably curved on blind side. Anterior nostril on eyed side a.well developed tubule, placed at some distance in front· of lower eye, the tubule short on blind side; posterior nostril with a raised edge, placed directly in front of and .between the eyes. Dentition chiefly developed on blind side, the teeth rather small, subequal or the outer teeth moderately larger, in bands; band in upper jaw of nearly uniform and moderate width; band in lower jaw shorter and wider, shaped somewhat like the segment as a circle or moderately crescentric with the straight or slightly curved margin entad with the well curved margin ectad; dentition on eyed side very moderately or poorly developed, the teenth small, few or moderate in number, eyed side of lowe·r jaw with one row, or with very few teeth, or altogether toothless, depending on the species and intraspecific individual variability. Opercle separated by a posterior emargination into two lobes, the upper lobe smaller. Branchiostegal membranes united, their point of union at base of ventral fin. Isthmus free. Pseudobranchiae very moderate. No slit behind fourth gill arch. Gill rakers nearly obsolete, indicated as slight, uneven, variable protuberances on gill arch (not examined in all species). Body and head with ctenoid scales, those on anterior part of head becoming more or less embedded; caudal scaled at its base, the scales continued backward in rows between the rays for some distance, scale less distally; dorsal and anal with short rows of s.(d~'.!9.blcrr•J~ P. "10'! :U'J(/; fau y[•rcen fijr:9b e!Li :rl'1;qr19J 'I') _t -rs :tr1r, f,dJ ( av.sw .{.Jod ;qril"tsqsJ s =--1,f!srl0 , sbr.r8 j?, i b J.uori~ · j"~ ~rj.J:rtfil~sci '1SqBj -:rol'!ejeoq ~l&jt::.•-r-ctbr,.n; "l:rH\\t ttfj2r:0I h .. t8fiffHjB 1o :?b•-rJ:rlj r.;~•Jj -x0J"!r.;.:rnB 1o bus j·,ri 12..f!_Y...:W... nl j]qaxe • ~~ 1r1:.w tidod ~~:[lj;J.L( ~ ~,:tl'"'..f."ISV oj vIE.i:qG"! ;grr.c:?.B9'I::isb :{:Jqeb udj , s[bbi:n s1i.:) :..uodn r1..:::.ri,i ·rsXJ.nmz "xL:tri2kic ,:J.CGme as'!:~; ·• $r?Sd lsbus~ W0'11:cnn '["1'$•V .B ..f.d !Jsjs'!oqsa ae~e owJ er.i'j ; juone ~r,:orfa dot~si ~11.tri~fro;t 2.asml!)sqz liem:e nl: 9:-,sqc '.fnJld',:oc-rsjnl ; 8W3:rr.r::isqe •re~rrnl ~rl.:J IY.i: oa ~I·merr ·,,-!JV -to '!sri.:Jo ,:.~o -:.:,~fl;c;vfr-- n.f. -~I01tnl:I~ S'ltt "!sqq.u 1.o n.f~--rsr1 'Ir,J:--rs~:-1s no .bfH1]1In juodG djod tEdrt9!J!,-19"'1'1fll ,"Is~,:oJ. 'lo j.5rf.J a:Ju t;J:--:.u[d ·crfJJv bns .:!"'fode \JSJr;n2 ·.l.so.fj70t; e.TIS& br)o ...rci" ?,!_H1ar1.~.:!rro~ vl--rssrt ~ :P,fll~fl"ror1 sI:.i:1.o.,-a "IO.t'"fr1L•r11:, •1e,i:,rw sI;vrn ·IOJ'!oJioq ~.-:!l , :t.r.nmz \'.'Ievj rl,JuoM , sv-,r.r:, '(J;;:·t1.f.Ie 0n11,:-·:;;1.3V ,_L(quq "!ewoI 1o n:l~-rf.im "!Ol"l'sj,16 :!J. b•1sw)!'ond s"'.fom ,,.-IjJ'I~.1.J:[e Jqe·1~XG cP-'""{GW rf.jod ~[::~;ip_·.::sbom ~IR:Ji'l.:temn!'(B!l ;g.ul~~I.~ bns ~.~13_.:_tv)_Q. • sI· l:. :.1-·1-tJd .ao h0v'ftJ:) ~~-[dr;jon c eb.i:e 'bs·io no bsvr:ru~ beiq0I~veb X[e~-1 r; :1b.tr~ b6i{9 no I.i:.. r.:teon ·101--re~nr-\ ""1:~1•.rr;l ""'rtj ,J'no'l~I nl :3:)nP-Jn.i:b t1:rt0r.: ds ~bs~slq , s[udIJj ;r; t•19;laoq ; sb.t::: bnl [6 ,no j'T,o:fa. eludu.J G/1.1 , ,,,,e n.i: -,[L:;,s'r.tb ,-"'.''3lq 'S:?,hS bec:l!','1 fl .r!Jl1-1 I f'!jflOIT V. [1;:Ltd:; n:.,.f.;j'.f.jnE.. ·-; .~9·(n 9rf.j_ IT9£}.wjse bnB 1o .:l'n:n•r·::_ ,JJ.sm~: •,:srfj?.,"f rii:,o;J fJrfj ,.sb.i:a bnl.Cr~ rm beqoisvsb n.i: ,"rn2~Hll '[.(9~':,;'rnbnm rfje'ej ·'lSjLJQ ef"lj ·•ro .f.GU,J9dU?. ixrn 111·ro'i.i:m.r ·'{.l"ff30!J 1o wst '!9CJ(!JJ 11.i: f,nnd ; c:h1L'ld • rrsfJ.i:t~.r brrs --:Sj"fOr!e &1st ~s":1ol n.i: brrsd ; dd"hlw 9jr;•Isbom. ·xo s I,,. i: :i. .c :::.s. jnsmg se srl,J ,,Dill .1pr!womo c bs-q r-.ri~ yJ::t:1;~11e · •ro ..(rig.i:rJ'lJZ srl.:J rlJ .i:w ::i.i:-r.:lns::ie,(~"f.O .'llejs'!sbom. ,i.t;;,:1-r-:n !-J9v•m::, !:Ls,·T srf.:i d.1.i:w bn~m,i ril;g-:rcra bsv•w::, 'fO 1Leje'1sbom t-:rev eble bots no no!J.i:jneb ;bajos 8,,,•.:-,Liom "fO 1,re·.t , Ucme ,ri.1rrneJ srlJ , b9f1o.[eveb 'r.l·10oq rr-:: ,t:,.70"T ~)!10 rfjJx: W.tl(i "'{6WOl 1o sbl:s bn~s (rrsdmr:n n.t ,-i?.eJrjjooj •y9fl.1'.?3s;Jic •ro ,11.1oe;JI, ws1 \'.'!SV r!JJ:.-; •1:i;'.rJh.t-0- Cl\r-Il ::> t1 r.~eqar.;--rjrrJ: bnn es.C:Jsqa e.d~ no 3rri:bns~eb ·101:,e.jr::oq s x:d b:,in·rnqs:c, 0.b'!eqOI ,Y.j.t.[ld>1.hsv .. --r::1JJ3r~1a Pdo[ rr-:;qq.u 9;{j ,aedoI --,1•11;_~ (Jj{tj {1').(j'crtJarrsn~e 'io ::rrroq ·1lerlj ,.f.leJ.i:rw een'1-,.dr1om lB;')c>jeolnons"f8 . s~•··.(~·. c.r.r.nrt0eI .r:ii I.c;rr:;r;r;v ..lo :,aed Js rrn.truJ ha i-~fr.:d jJ.:I~ oM • sjs•1ehorn '!'!SV er.>.~d~ng'!!JobueetI ,s~u;Issdo ,:f·r'"'lfl C"fi.JJl.s•r [.[_i:D .:for,.s II.i:;g rl.:J'T1101 8Gt.>~IG"'f9i:J.°:o'!q olds.I:·1n~.r ,rr.evcH!IJ ,.-1rf~i.[e as hsjn'JChn:l ;,:one • ,, ?.,,;.::>sqa .[[s 11.r b(,rdmsxs j'Jn'I) do-rs J.£.c~ r10 j•,::;q '!·) 1rrsidrrc; no seor!J ,zeir,::,e blor1s.:J:i d.J.tw hsnrf bnr.; [sbu1,o ; hob.bsdrrre ea0l '1.0 D'1om ;;1,,1tmo::>eo' bser! 1o b·x,:;,.-i:1r1cf b~•trn:.tJrro:i esls,e er!:; ,0c:.;d1 3j_c ;Js bsis::>2 -sl.s~,a , eom1.:!c:lb 8i'!Ior. '!o1 e'{G'I e:-!J rissw;Jed awn, nl ~.10 ,:,hos.-- ·· · ,J"f.._-· ,..,,.., ... ,c. r,_.l~ .....;.,.- .,J ..,,_e.n:nr b ns I.,,.•~c: ...... o b :,t',, fl GJ" , . .r.;'f- (.cs~·· __r 4 small scales along the proximal part of the rays. Lateral line absent. Dorsal origin over eye; dorsal and anal continuous with caudal, Pelvic fin unpaired, placed at isthmus, normally having 4 rays (the rays counted in 364 specimens representing all species, only 4 variants, one each .in diome.dianus, plafiusa, and Xenynsi, having 3 rays). Pectora absent. ii rays segmented and unbranched • • • • 11 Symphurus plagiusa (Linnaeus) Pleuronectes plagiusa, Linnaeus, Syst. Nat., Ed. XII, 455, 1766, on a specimen from Dr. Garden, probably from Charleston, but the locality not quite certain; and of various copyist, Plagusia fasciata, Holbrook MS., De Kay, New York Fauna: Fishes, 304, 1a42, Charleston. Glossichth s 1afiusa, Gill, Cat, Fish East Coast N. Am., 51 , 186 • Pla&J;sia 1agiusa, Gill, Cat. Fish East Coast N. ~., 94, 1873. Aphoristia pla!iusa, Jordan & Gilbert, Proc. U.S. Nat, Mus. 878, 368; Jordan, l,c., 1aao, 22; Jordan & Gilbert, l.c., 1882, 305 and 618; Jordan & Gilbert, Synopsis, 842, 1gs3; Jordan, Proc. U.S. Nat. Mus. 1884, 144, Aphoristia fasciata, Jordan, Proc. U.S. Nat. Mus. 1886, 53, Symphurus 1agiusa, Jordan & Goss, Review Flounders and So1 es, 325, 1889, "• •• Description: C (9) 10 (11). D a5-92 A 69-76 (78), Sc 71-86, Eyed side with teeth on anterior half of upper jaw or for a shorter distance; Usually no teeth on the eyed side of the lower jaw, sometimes a very few teeth at its side. Measure ments of 5 specimens 124-147mm and 6 specimens 54- 77mm: caudal 11,0-11,5 (11,5-13,0), depth 29,5-31,5 (27-31), head 1a,5-20.5 (21,0-22,5) preanal 24-26 (24-30), postanal 79-83 (77.g1). Color very variable; cross bands darker than ground color, present and absent, when present in various stages of development from almost solidly continuous to interrupted, incomplete or irregular I " I. .. 8'"'lR~ ~rf.:J '!o {!~·st1 JsmJ-yc•-rq ~rlZ: :=:trc-·J~s aBit;;oe J.1£,ma ; B'fa '1~-::vo n.~;11-~c !e~:·'fcl! • ,.;t!e:.:,?.s s-1111 ce,-rs.::hJ ;,.i:vJ:eCJ .lshl.ino r!d.r,,; -eiroun.r,tm,::i ~·::;rm bn:?. .[r;e-;ob :gnivr.rl "7-J.:._i.;J°ll-:r(,a · 1 eur:rfje'l,:-j,3 b0::i;;I, ; bs-r.tsq:n; i,l] ;;neu.i·t,(,qe. Jof__ rd: bc,;Jmrn::i' c'',"!'\~[ s;[;;) B'i:G"J" ii s~o , e0::1-YfJ 1:"TtV J~ 'fl.Cffo t c fi•.t~et;z l 7 r; ;qn.fjf!~\es~c·e"': ,.i:2.rwn0/: b!rs ,;__;e1111}r,£9: _;?.un.si.bs!!!o.i:.b .r:1 ~!~,,-~ D":!'l''f .L 1.A , i~fIS~ds 1sr1o·J~s•:'f .:fBv.s•r t ;!},r, [Vfi,C Jr. ., • , berl:Jn.r.#)'"rln.u b~'LS be.:i rr£:.Y~ee ( ar.rsnrrrrl:iJ} , J J:.J( .b:1 , ,.:JsVi • .:ta,i_:c! ,aue.sn:NJ:,l ,~gg.Ilq, e~.:J_?.§'norrueJq ,rrsb•rnD .'TIT mo"!'J rrewlaeqa II No-r;od\'1 , i A. S:Q-c.'_i3 G , (J.l) Ol (~) ::J :rroJ:jcri-r9a9G , •• 11 rro rl;Jsej rf.:tJ:w ebJ:a llsV,~! .1J8-I1 08 • (8'i') c'i'-Pc'i ; e~ne~t.t.tJ ~9;/'1.ori<~ n '101 "Jo wsf. "1sqqu 'Id '"ilarl -ro.frre.:1n£i ,wst. "T~•,0::iJ er/o 1o eb.i:e be"{o erli r;o rl.:1,eej on ,:Jls!Jt'JlJ -~·n1c:;s&t'1t • f.tb.i:a cjl Js rlJ'sAV ws'l x-rsv n esmld'ern:oE: --.µ? nrrnm_r:1oqe 6 brrs 1l'!!fl1+ll-,\Sl ~~twml:ieqa <' 1o .::,,ne;n ?;Jf-~.?S ~jqeb ,(O,(I-i:.II) ~.Il-0.ll lsbueo :mm,~ c.i~--,i~ l&v!HJ"lq (i:,S:S-0.IS) i:.OS:-i;:,8I' bsMI ,(It-'i'S) .(18-'i''i') (8-~, Isnsjaoq ,(0(-~S) I 11.,:;rij "IG::-i•rnb abrr,3d aao•;:i ; eldsl"ISV 'IV,"I0V "Iolo::J r1.t ,lnec:FJ'i:"[ nerlw ,.:!need!:\ bns ;Jrroae·rq ,•roloo b11uo"';1 \'. Ihl.fc;Z ,rr:oml£J mo"I1 ,lfl811lqol8V!lb 1o 1as:,i.sja auo.i:"Is•: "lr,Ju~:s-;'!.i: ·ro s.:n,lqmo:in.t , be;JqJJ'l-rsjrr.l:1 o.:J auoun.rjno:i 5 in various degrees, or only faintly indicated; sometimes with few or many intensely dark, very small speaks a large spot on opercle, centered on its upper .lobe, present in•the majority of the· larger specimens, especially those recently preserved, usually faint or absent·in the smaller specimens and often also· in· ·the larger;· sometimes · a smaller solid or interrupted black spot on lower posterior part of head; dorsal, anal, and caudal fins faintly or moderately. dusky, usually with a darker pigment along the rays, •• ~ Symphurus civitatus Ginsburg 11 ••• Description: C (11) 12, D 87-92, A 70-77, Sc 69-80. Teeth on eyed side of upper jaw extending over anterior half of jaw or less, none on lower jaw. Angle of mouth approximately under posterior margin of the lower pupil or middle of eye, Measurements of 10 Gulf specimens 91-147mm, and 2 Atlantic specimens 121-153mm,, the latter in parenthesis: Caudal 10,5-13,5 (10,5-11.5), depth 30-34 (31-32), head 19,5-21,0 (18,0-18,5), preanal 24-26 (22-24), postanal 78-82 (81-82). · Cross bands usually absent or faint, sometimes rather fairly marked; opercle without a black spot, occasionally with a dusky area; caudal and posterior part of dorsal and anal variably dusky, sometimes nearly black , , , •11 1 ut;-JL:·A pJ'dCJ{' • • • 'u ,, -, , {.)~LP 01, (JoL2sr: S:.Jq ~HJSJ A8L:fgpr:\. qna;<.:A 1 3e;u;,_· [?.[ .jJQ ;! UGCS~J-[UlJ8JJ/l. ,\~:f PF S qn2r,cA ~LS~! 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P:rf.!wGup ~ro!Jtl pps ;;..sf~2 • • • ·0 c\l11qsI r,ru:1 .ss~lJL'JJ,. oJ.. wo:;or..8res11. q1rn1a,' a,1fi'=-TJ1 JO:~SL bo ~.~GLfuf- bs:t.~ 0.(, pH~q! qur.-asy> 1H.ii-1J~ SJJ(j 8" au1sJJS!. ?.OJ:fi:I oi:. r1Jr:Gu,nb;0q pysci,,:: :3ho1~ al• 2b~crmo;;2 uuq o·.c.,p-su ~J~~ :i fD ~ps 1~1..TT.e1.: 2•Jm.s+ fl~G'"J i11..,68\:JJ.,A9q 1 H~3db'JJA r.·:q1J1-\ OJ.. 'JfJ".'.?G;JP, .P,J f:96 a;i_:~Jr:::.1., ·,-0, -~.---, jtlrCl~T·•· JV_...,,,.:,A> ~i~·•,c,.i,.,r;,v.. µ.Y;JJ.J,.. e-:,r-:.•r•-,·,'.1-J...... •'V..o •• •:, J:\.... ;-.;\u"-;--- "8• N\"'I"'GCu·•,,-.-,\ :, \·J ~ 0 ~u :r~a n!:J1Gr., rv;.. .;• b1.B'1SH~ :ru I~J-!G :uu·.io.\.a:ff2'J\. Ol,. pps 1 ~lll~JJ" ~b0S~8 g' J.JL~S 8buc OU Ob6~CJG G8UCOL6q :;p,P'f"\"f"II•-:·,:, MJ •'U' • -- .. , i~• rup·:·u-~-•-',\ 1", t~c.,,u.Ut,-!W.l.:l._. •••••-,I•.~-- •N .,,;.-:,u··.:J,~ J'- • f"o.;J •OJ,,;. ac,.,.,~J,. •1\..,.) .L,.o,_..,.,,··, .fl.! A~H.. J Jfl:3 (~-3-t..'.;.·.;;,;ni ;_ ..r. 'Ji.:~-: .. .. :::.f:J~J:-l :[JJ·--r~.::0!:',:;q! i CHAPTER II REVIEW OF THE LITERATURE The genus Symphurus was described-by Rafinesque in 1810 in his evaluation of Symphurus nigrescens Rafinesque. Many species of the genus were placed in new or different genera by several authors during the next· three quarters of a century. The generic name Symphurus was restored by Jordan and Goss· in 1889 (Jordan ·and Evermann, 1898), The species.of Symphurus, the only genus of tonguefish in the western Atlantic, are nti~fou.s'· and closely allied, and vary substantially from the European and Pacific forms (Ginsburg, 1951), Ginsburg's comprehensive work with the western Atlantic cynoglossids included 13 species and 2 subspecies. Symphurus plagiusa and Symphurus civitatus were included in this stud~. Original descriptions for these two species can be found ," 1n Chapter I. Distribution Western Atlantic Symphurus are broadly distributed in coastal waters from New York state to Argentina (Briggs, 1958; Ginsburg, 1951; Lazzaro, 1973; Topp and Hoff, 1972), In the past 15 years, three new species of Symphurus have 6 7 been described-from the Bahamas and Greater Antilles (Bohlke, 1961; Bohlke and Chapliir;, 1968; Robins and Randall, 1965). Three recent studies, made in the Gulf of Mexico, produced new information concerning the genus. Two of these studies (Gall~way, Parker, and Moore, 1972; Topp and .Hoff, 1972) led to the formation of keys for Symphurus species. These keys were derived-either as a modification of Ginsburg's key, ·or from recently collected data. Gallaway, Parker, and Moore_ (1972) prepared a key· for tonguefish found in and around Texas coastal and estuarine waters. Topp and Hoff (1972) constructed a key for tonguefish of the continental shelf off southwestern Florida. Walls (1975) generally described the northern Gulf cynoglossids in his study of the ichthyofauna of that area. Ginsburg (1951) considered the range of~ plagiusa to extend from New York state to the Bahamas, and throughout the Greater Antilles. In the Gulf of Mexico it occurs from Cape Sable, Florida to Laguna Madre, Texas. Briggs (1958) considered the range to ex tend- from New York state to Argentina (40°s), and Topp and Hoff (1972) extended the range proposed by Ginsburg to include the Yucatan shelf. They considered the range proposed by Briggs to include that of Symphurus plagusia 8 (Block & Schneider), sensu lato. Symphurus plagusia is a southern species ranging from Panama to Argentina, and is an unfortunate anagram of S,. plagiusa. .However, two 'spe_cimens of . .§.. plagiusa, taxonomically identified. by using specimens catalogued. in The Smithsonian Institution, have recently been identified from the coast of Argentina·, off Quequen (Lazzaro, 1973), Lazzaro suggested that S, plagiusa may be occasionally found in the shelf waters of Argentina. This is supportive of Brigg's (1958) work. Ginsburg . ( 19 51) ·noted that .§., plagiusa is the most commonly an~ abundantly 1 found species of Symphurus in the Atlantic and Gulf coastal areas of the United States. Topp and Hoff (1972) reported that no important inventory of the southeastern coastal United States omits this species, Many surveys have been made in the Gulf of Mexico, each contributing to the biogeography of s. plagiusa (Boschung, 1957; · Compton and Bradley, 1963, 1964; Gunter, 1945; Hoese, 1958, 1959; Jordan and Gilbert, 1883; Menzel, 1956; Miles, 1951; Parker, 1965; Springer and Bullis, 1956; Springer and Woodburn, 1960). A paucity of information is available for Symphurus civitatus. A few general surveys have been made in the Gulf of Mexico (Hoese, 1958; Springer and Bullis, 1956) and in Atlantic coastal areas from North Carolina to Florida (Briggs, 1958), Symphurus civitatus .ojgJ u~ns2 ,(~eblanri~2 f ~~o.[H) ' ' _(\)_"f.S rf'.rrlji:";'.~1~'1\ ')j Gl)1J3f!B(i !If()'{] r::.1.!::·,-r.~'t !?.s t~19qE rr".1"e!'l0Hor~ .f'> b9111jnabl ~sad yfjns~e~ av~rl I . , • ( (\'i? l ;o'!p,~ 1-sJ) nsrrp eJJ:) 1'1:o ~~:i-1 2. '1.:.., ~· ':~:i 't[:~,~c erLJ· n.t bru10'i "'[J Lhrto lc:~a~o sd ysm -~ 2.s..r .t;1 BI q. 1 • 11'".:T:\•: (S<~DJ:) a'r.;glrrd ·10 evJ0'!oqq.ue e:.t alrI~1 .~•--:"_tJns~·1:~ :1dj c.t -~l~-f.t1.1_~:_:fi.i -~ "jgr!j bGjo(li (I('.,(?I~) ~:ruderrlD • f ' • ·... ,.:r~ud,..... Ci'·2 "Jr:, C!~ .t6!:H~e bouo'f v·.fjilaba.u.dGI· ·Dr;s 'l.Ino :- flc<: ~:---.cJi -----•--- • L • .:,'ni"'-J'!o·:, -~ or: dGflj bs;Jero((6'f (S:\'(2[j 11.oH, bnr. qqoT ,ee~:.e::·::: I e.:UJ110 Cf.J;/,a.·;,: bsj_i:rrU f.sJ8s0:i wre;Jenarldr.roe nrl.J 1o '?;'!o~oevrr.r J_(~ ':Jii;·; sd~t r::J: sbr-..J:1 uasd 9Ve.ri G"(9V'"i.ue ~rtsM • Lo.i:~9qc- a.frf.:1 ' • ,2;c,r i'Ih.b'iOG Pl ;,c;ll?l ~ --rsjrnrD 4 ~ j"!!Jdl lD bnr3 ;1:.:01 , 1l c ,sa9oH ' ; - I eeI r:., t I 98.fIB:~·1., --r:1;•1·r_: '::"08 ;cov..i: t '!9~'!Gq ;lc\?I t -·. ;oc •!l ; Cil!U ' .cujsJ.tv_i__ _ i:i 9 seems to have two distinct populations; the larger in habits. the coastlines of Texas, Louisiana, Mississippi, and Alabama (dinsburg, 1951). The other population is described as "not so common 11 •• ( Ginsburg, 1951), and extends from North Carolina to Florida (Briggs, 1958), Topp and Hoff (1972) do not acknowledge the existence of 2· civitatus on.the southwestern Florida shelf, except for one specimen collected from St. Joseph Bay by Ginsburg during his study, Anatomy and Morphology Certain anatomical and morphological character istics of some cynoglossids have been studied by De Grett (1971). The fishes studied had", well developed esophagi and stomachs, complicated intestinal loops, a lack of teethed gill rakers and pyloric appendices were absent. Chabanaud (1940, 1947) showed anatomical variations among certain tonguefish species, especially in the structure of the neurocranium, jaws, and urinary papillae. He later (1948) described a new genus based on these and other characters. His work included the genera Cynoglossus and Paraplagusia. Ginsburg (1951) was of the opinion that a comparative study of_§. plagiusa and 2• civitatus was required, because they were often found together, Walls COllibSL~CTAG aprrg1 Tll /.'fJG 2/.'LflC/.'llLG 0:1, /.' JJG lJG/lLOC.<:.'illJ{ fHl, • ·~s-,,18' fJJJq 1ir.rm,Ll, A~1.;·s~rona sruoug ~sLpsru couUrrsir2µ 2bac;ea~ sabsc~~JJl i •• CfJSpOI.ISflq ( Jo\"0' Jc.li1',\) 8J1vl•lGq Sll\'J./.'01.l':j"CSJ" (J"o,.\,I)' ,I,JJG '(,'.f8fJGi8 2;:nqJGq jJ::l :r2r-:rc<1 Ot, 80::llG c1uio'§'.J08\l:f::j8 jJ\!J\.G pGGlJ 8/.'IHJ:rGq pi. l)il (;l.,Of-/.' I csLps-:ru s-JJ;u.com:rczJ euq uior... bporcg:fC~J cps.~~c~e1.- qnL;ui pra acnql· ous abscrl!islJ C'JJJGjr;rsq 1,r.o,u ?.P • ? 020!:p jl\c/1'. p}, ~:-rir:pn:.c;'. I ' • , cf,qr.~/.'118 OD f.'!jG 21011.~p.v.s_a;:,s;;,n. tJ"Ol-lqS. 8jJGJt • GX'.;~,,_._; :,0L J,..,.,_'\,) ,_.,., Q \ I () 10 (1975), who gave a general description of both fish as individual species, alluded to the possibility that S. civitatus may .simply be an offshore variant of§, plagiusa. Ginsburg (1951) stated proportional measurements were of little value in segregating the species of tonguefish. However, he set forth several parameters by which species determination could be most easily accomplished. Among those parameters suggested were fin ray counts, dental construction, head length, body depth, caudal fin length, preanal length, and postanal length. The last five of these could be expressed as percentages of the standard length. Only a few, and often indirect, references are available concerning intraspecific morphological variations in S. plagiusa, Jordan and Evermann (1898) stated that S. plagiusa possessed a vertebral count equaling~?. Ginsburg (1951) referred to dorsal, anal, and caudal fin ray morphology. He found slight variation in dorsal and anal fin ray counts; the caudal fin ray count tended to be constant at 10, He also referred to an opercular spot that was commonly, but not always, present. These four characteristics are considered to be the most obvious distinguishable traits of§. plagiusa, which is a comparatively homogeneous species, with the possible exception of the Cuban population. OJ: .C .JcriJ ~~:,j_(,[Jd.t~r~oq Gci:1 n~J b~::lbiJC[8 tJt=.!·3[:'"'10.JE J='HJb.~vJbtl I • "")r '_t(·, ,o,_'t·rr:?,\'. ~1, ... .;,c,_J_,.,,,.::1-.r~.,,t--[- •:_' .-t,,• 1 _ >.1~-~"t·,_',--·1.".,.o I -,,.·.---:~ ocr,_. • ,,rlnrr,·••), ,.J.,.J,1 •• 1...,, ,,r-•.-.,,r.,,Li ;;:::::,'-!_!,~.--=~--;.:.,_':,/_··J·,,r·~ 0 ' z..:!:l!-:,!i8"::J2•:::fl:i. .Lsr~cl,J'l-:iqor:rq· -bsO:;.~;a (I(\~l;) ;.:1"1ud2nlD , I'' 1~ 2e_[.:)~q2. srf.:.i ~inJ.;q·ie'l~;.sc: nl eJL6V eij,:.tj:l 10 S'1~~-r . ' "f{d- F.'1sje··1rv:r~~ le~~ve~ dJ'lo'l dea or! ~'IS,rewoH • rieJ1E·H;:i.11oj . .. ~[22~0 ~ 010 ~d bI~o~ :n?!~~~l~iejsb ~9l~sqe, rl~lrlw . . ' c1.t'1 &~D':1 .bEi.:tes;~rf!~a· e·s-,S~sme-xhq· 9eorL1 ~aj?mA, _• .berl~liqmo,vn ti I ,dd1Ah ~bod ,rljjnsl besd ,~oi~or~jea90 ,[sjns~ ,aJnuo~ ~e~ .dj~1sI h~sbrfsje I bn r, , 1we'l. e 'clnO 2.n0.r;;,·t· ','.5V [ n~•.r,.10.· [r!o . ;:' cr'o bs0'fi ,i;:; ( 8(l8.[) r.nsm'Isv~ brrn m;b'Io L .g u.t;gsia • !c rr.r s ~j: z-l:)ldw cf?8LI.t;1nla .§_ 1o zj.i:s~d s[cfr..rl::--. .!:u~9.rrlda.tb I' .3_[,ih;eoq- sli;J r{j_tw , ze.c:'lsqa euosrrs,;gomori 1[Isv .i:dr.··rnqmoo .rro.i:JBluqoq r!GdJJ::l srf;; 1o no.r.:1-;s:ixs 11 Ginsburg (1951) divided the range of S. plagiusa into four areas,based on personal preference: Long Island to Cape Canavera_l; Key West to Tampa; Apalachicola to Corpus Christi; and the Cuban caribbean. Average dorsal (D) and ~nal (A) counts of specimens from the Apalachicola to Corpus Christi area were the highest (D 88,77, A 72,81, n=44). The Long Island to Cape Canaveral group were intermediate (D 88,44, A 72.38, n=92) and the Key West to Tampa· specimens had·the lowest count (D 87.59, A 71,65, n=32). He also showed that the Cuban population was markedly higher in both dorsal and anal fin ray counts (D 89,50, A 74,501 n=2). Ginsburg suggested the Cuban specimens were either extreme variations, or members· of a population which ordinarily have higher dorsal and anal fin ray counts. He also found this population to have higher scale counts. Symphurus civitatus showed the same variations in fin ray counts as did~- plagiusa, except the caudal ray count of~- civitatus tended to be 12, but varied to 11 or less in the case of variants or structurally deformed individuals (Ginsburg, 1951). Ginsburg also stated thats. civitatus closely paralleled Symphurus plagusia, sensu lato, in morphological characteristics. On the basis of similarity, he suggested that both species require more taxonomic study. 12 Behavior and Ecology Certain aspects of cynoglossid behavior and ecology have been studied. De Groot (1971) reported on the interrelationships between alimentary tract morphology and food, and feeding behavior in flatfishes. He con cluded that cynoglossids were primarily polychaete mollusk feeders, although they sometimes feed on crustaceans. His study included several genera of the family Cynoglossidae; among these was the genus Symphurus. The water depth, in which S, plagiusa and S, civitatus inhabit, has been considered a behavioral characteristic of much importance for identifying the two species, Because§, civitatus is consistently found in deeper water than s. plagiusa, Ginsburg (1951) considered habitat depth as a useful and valid separatory character istic, as does Walls (1975), Ginsburg also states that~. plagiusa is pre dominantly a near-shore species inhabiting depths from 0-14 fathoms; Topp and Hoff (1972) consider 6 meters to be the bathometric limit. The greatest known depth of capture (50 fathoms) is recorded from collections at f!../V Station 944 off Pensacola, Florida (Bullis and Thompson, 1965), Gunter (1945) found that S, plagiusa preferred water of high salinity (17.1 to 36,7 ppt); most samples l.\ar-s1 .. oi µ.rSp 215J:rllltA (Ji\·J ~<> )(?'~ bnp): J!Jo.:r: c1::n.:!:~:Je2 ' . Jci\"?.) • (lfllJf-'SL ( Jq\+~) :r,om:;q f:}l'.'lJ:' ~- 'Ej'sl.:,rn~ c, .. s tGr.:... sq I :3i:-s..c:r'.;U Qf\r 01.t oSu8~COJ&~ B.,Jo1..:rqs {DnJr:r~~ ~nHJ J,!-1 :..:11;L:-J0u" ct1.t·r.rrr.s ( ~o tvr.µuu\a) :ra LSCOLqeq r.;;.om c(;JJ€crr·:,u2 u c t\1\ 001rcT::i11,·"J·,, s 1Jes:1.....J2110I.e abec.,s:a :rcmipr,,:;-u,; m,r,r-µ, r.r.ow ~. -. I,., .. •J ...... ----·~- ·• (J:fll2p!1l.°ii 9!Jc-:o -apa.:-ea CfJ9f; 2• !1J~n{:rn8£i ~-2 i)l.,G- 1 qusl;,1:;z. W:lf.GL CJJ\Hl 8 • OJSO::prn§ (}.[llap::L'is: ( Jd ~J) c C!J2'ftf'!LGq 2!;,eGJ:,n.· BGC\c!Jl'ilG J2' c:riq'tll'CITT ra cou·a~•ace11f.'Ji i;,orr1Jq ·.p1 cpGL'flC~6l.,'.f?.P~·c O.f.. :rnf1GJJ' .rwboL+:9SU(.;f:1 t 1 c:(.i(n:sr.fla IllJJ;,p.[r. • µsa j)SGlJ GOlJ.1:f(J6LGq S pG)J'il'iq<-L~J J,JJG 1,1sr.GI. tjsbpµ' :ru 1,,p:rcp ;:i• bJ\J~:rr:2\.i ,n.Jq ~· :;_,i.H.Ul JA cAuoltJ08 l!JOJJff~J{ l,66qGLa~ S~(:'}J'.)Hl;jJ ~pe,j1,. aorue~:rwGe tGcq I)~ . . . I , cyn,1eq 1.•µs~ c1.r.,o[;;yo<1::q-q a 1~eLG bL:fr:JuJ,:fJA boJ 1'.CJ.::l<>f.'G- :1i:.iq · i:,oo'.J' suq :i,,ssq,:ru1, pepGll:fOI. :ru irur.t.:r«µi:a· ES con- , '. GGOJOG:J1. }J~AS pGGU ~~ff1LJ:fbq• I • ••6} f,O~'f"T' f;'~!f('A.•,r, ••'"l QT" \_, ""I· ~ ~ ..., ''U • ·- ,., ~.., J:S 13 were taken from waters with salinities greater than 30 ppt, Miles (1951) corroborated this finding with his analysis of trawler trash fish collected in Apalachicola Bay. ·s. piagiusa was abundant.in two trawls from waters with salinities over 30 ppt, but was rare or unknown in trawls when the salinity was under 15 ppt. Based on reports of other authors, Topp and Hoff (1972) considered ~- plagiusa to be the most euryhaline of the western Atlantic tonguefish. r Topp and Hoff (1972) reported that analyses of the stomach contents of S, plagiusa yielded polychaetes, brachiopods, crabs and other crustacean remains, including amphipods and cum~eans. Springer and Woodburn (1960) found the food of Tampa-Bay specimens considered almost entirely of copepods and polychaetes. Stickney (1976) reported the species feeds primarily upon benthic organisms, and is relatively indiscriminate regarding food intake. He also noted that a high percentage of the specimens examined contained sand grains in their digestive tracts. He concluded that a significant quantity of detrital matter must be ingested during feeding, and that; if this is the case,~- plagiusa may play an important role in detrital degradation in some Georgia estuaries, Stickney ( 1976) also st.ated that S. plagiusa appeared to be a benthic feeder throughout most 14 of its life (postlarval to adult), and differences in food habits were a function of,mouth size, rather than selection. Symphurus plagiusa, in the Gulf of Mexico, spawn primarily during spring and summer; with the onset of spawning beginning in March (Topp and Hoff~l972). They also reported the presence of developed ova in March (Station I on 21 March, 1966). Hildebrand and Gable (1930) theorized that spawning occurs at sea, rather than in the more common inshore· habitats. They also suggested that spawning occurred from May to October, peaking in June. Symphurus civitatus unfortunately has been neglected of scientific study with reference to ecology, food habits, or reproductive habits. These variables only can·be assumed·to·be similar to other western Atlantic cynoglossids. CHAPTER III MATERIALS AND METHODS Most specimens of §. plagiusa and §. ci vitatus - for data collection were obtained from field collections ~nd observations made in coastal areas between New Orleans, Louisiana and Panama City, Florida. Additional data were obtained from the study of specimens supplied by various universities and governmental sources. Field collections began 27 May, 1975 and ended 15 August, 1975. Shallow water samples were taken with standard 15-, 25-, and 50- foot haul seines from mud banks and tidal pools along the north side of· Dauphin Island, Alabama, and the grass beds of the Point aux Pins Salt Marsh Laboratory near Bayou La Batre, Alabama. Offshore samples, using 20- and 30-foot otter trawls, were taken. aboard the research v~ssel G. A. Rounsefell and four commercial vessels. Samples collected.aboard.the commercial vessels did not include sufficient data concerning collection locations; therefore, it was necessary-to approximate those capture sites. Most sampling was done diurnally, although several offshore samples were collected at night. 15 16 Some data for~. plagiusa and§. civitatus were obtained from outside sources, because time limitations and financial considerations made it impossible to sample representatives from the entire range of both species·. Data were received from the following universities and governmental agencies: The National Marine Fisheries Service (NMFS-P), Pascagoula, Mississippi; Gulf Coast Research Laboratory (GCRL), Ocean Springs, Mississippi; University of South Alabama (USA) , Mobile, Alabama; University of Alabama (UAIC), Tuscaloosa, Alabama; Florida State University (FSU), Tallahassee, Florida; University of.Florida (FSM), Gainesville, Florida; University of South Florida (USF), Tampa, Florida; Florida Department of Natural Resources (FSBCO), St.• Peterburg, Florida; Grice Marine Biological Laboratory· ( GMBL), Charleston, South Carolina; University of North Carolina (UNG), Morehead City, North Carolina; Virginia Institute of Marine Science· (VIMS), Glouchester Point·, Virginia; and· · the Facultad de· Ciencias Naturales y Museo (M,L.P.), La Plata, Buenos Aires, Argentina. Below are listed the· specimens of_§, plagiusa examined (N:225) in this research; ChesapEiake Bay· (VIMS- 3449, 7 specimens, 116-122mm). Off the coast of Virginia, North Carolina and South Carolina (VIMS-101053, 6 specimens, 123-145mm; VIMS-01314, 3 specimens, 61-12lmm; '.I ' •' en0i~sjlull s~l~ a21rBoscJ t~io~uo~ ~h~aduo•·io~1 banlsido 1 sfamRa• oj elrll~Hoaml• ~-t eh~m arroljs~ehlI eno~ [~J~~Rn11 hns j"Gf:08 'i..[;.rD ~ .tqqln:.::1:es.i:f/i ,Gil!o~S!)2Gql '(c1-8r•i~1H) 9::>.tV':!92 ;.C 1·{q1:::·;~t~~~.tr~t ,2:"l·u1l:'lqP. ClG£>00 ,(J£~)D) ""t(roj.s'lods~1 rl~"I.GSaefl I ,iorTI!',d,dA ,sCl:doM ,(AcU) .snrndnIA fl'j1108 1o zjce·r(nr.i:11•.J • .:.t2 ,(0082"1) eoo4!H'.}8s.fl Is"1FJct.aVJ 1.o ,.jnsmC"Jflq$G '{'rodr.'!0ds,l Is:-il.15 olo.i:H 011.i:~s/,cl 9!).i:'T;-_1 ; r;bl•w [': ~1rrs ;0.ta.t~"<.i:V ,jct.foci '1eJaerf:)tJ1JID ,(8MI\f)., e::>n!;li:::,8 sn.C"ist·•1 '1o I , ,:;.i:11:·~-r.i:V 'to ;.tznon 9r/j 1'10 • (m,nSS:l-dlI :1arwr.1.i:o:-iqa 'i' , t=>;l,lt C t f.cOIOI-c:rirv) sn.tI,,,:a::l fij_tKJe fms salI,1'TS::l fij"Jo/!i ;wn-:f.::I-fe ,:::rrnm.bsqe ( ,-i>IE:I0-2MIV ;:runc,\I-(S:1 ,eneml::i9q-e 17 VIMS-02576, 9 specimens, l27-173mrn; UNC-3759, 4 specimens, 156-179mrn; GMBL-320-FB-71, a-specimens; GMBL-65-8, 8 specimens; GMBL-71-52, 22 specimens). Off the east coast of Florida, north of Cape Canaveral (FSBC-5797, l specimen, 124mm; FSBC-4352, l specimen, 144mm; FSBC-4375, l specimen, 105mm; FSBC-2384, 1 specimen, 100mm, St. Johns River, Putnam County, Florida). From the southern coast of Florida:,. south of Cape Canaveral (FSBC-82, l specimen, 83mm), From Cape Sable, Florida to Tampa, Florida (FSBC-2145, 4 specimens, 125-137mrn; FSBC-1596, 7 specimens, 84-136mrn; FSBC-6368, l specimen, 99mm; FSBC- 2749, 1 specimen, 97mm; FSBC-332, 1 specimen, 133mm; FSBC-2567, 1 specimen, 109mm; FSBC-3567, 5 specimens, 73- 90mm; FSBC-2274, 3 specimens, 115-126-mm;FSBC-2636, 6 specimens, 134-15lmm; FSBC-1045, 8 specimens, 26-64mm; FSBC-41, l specimen, 89mm; FSBC-3080 1 l specimen, 26mm; FSBC-1125, 1 specimen, 60mm; FSBC-3093, 4 specimens, 29- 42mrn; FSBC-937, 8 specimens, 32-44mm). From Apalachicola, Florida to Laguna Madre, Texas (USA-02569, 45 specimens, 87-168mm; USA-01899, 8 specimens, 76-105mrn; FSBC-1354, 5 specimens, 50-64mrn; FSBC-3936, 2 specimens, 56-63mm; FSBC-3962, 3 specimens, 42-68mrn; FSU-18398, 16 specimens, 112-154MM; FSU-18807, 22 specimens, 48-79mm), Many personal samples were also examined from this area, but are not individually listed here because they are not .:.rrs11[1$-"J'B .J t~CS'(-:1~:rr ;r:m1ES'l-~SJ t~t!::•i1Ut9CfB c ,2"\"(S:0-2MJV -~ ··-~ ., .... I 14r"\ ... C: • . ti ,..,~ ,t;-. QCC ~n,:r )t"[ ----- r t_ ,!J-co-,.r! 11.1 ,.-;fk;t.!(~9r?,f: v t ... :-t1·1- ~:l, .... LiCJ,:1. 1 ;mm•,,_ -o:::_ 1 f!''iC:i·1dt,'f"18 •:1cL:' ffiO'lr;l • (i.;L,1:•'fOl'iJ ,-{ctn;.;o.0 !~Snj.·_r'i ~r:rr\~_::5; ~f~:ir/[. I , SR-:.n·c·•-O J.s~rov1rns8 oqr,8 'J:o rld11oa1. ·, nhi:'1oJ'iJ 'lo ;!,-. l'o~, , r-.c·!.-H:·~ t),1 sbi"1ol~1 , eidn2. eqn8 mo•'f·~1 • (mn:(B , n:)mio!;1qT I ;r,H'IT((l ,nsr1t.b8qB I ,S:((-~l8e.'~ iiTTJl!\ ' --:\' , ,,,mmi:::i,;qii 2 , 76c(-::JC.8r,l ;mml:)OI ,r1eJ1oeqs l , ,ocS:-,8m:'•i o /l(r':S-:J!:rn'>l,lllm-oS:I-c'JJ. ,amim.i::isqa :( ,.11\'S:S:-Oclc'~ ;mm0<2 I ,,1rrrr;16-t'iS: ,2.uem.i:::,9qa ~ ,c.;.\OI-::JH2'•1 ;mn(lcl-,!(l ,nm::;ml:isqc: -PS: , nrrnm.:::isqa .,\ , (E·O(-::JUc.'>l ;mmOo , nsm.l::ieqz I , cS:II-Df:lf::'iJ I ,i,Ic•::ilr;;;c I.BqA r:10'!·~ • (mm+l,l-S:( ,errnm.i::ieq9 3 , 'i'(('-D88'il ; 1lll!lS.,l I , a,,em.i::ieqe i',l , Pdc:S:O-Acll) e.r.xe'l' , o"fbs,\) ·r,nl.liJ~"l r,j sbl:"fol'>; :mmt \'.,,;.;;:i • r 1r.mP\'--8il., cnsml:ieqa S:S: , 1dil8I-Ue'ij ;1-MilcJ-S:I.E 18 officially catalogued. From Argentina (M.L.P. N I-XII-72-6, 1 specimen, 69mm) plus the description of two specimens taken from the coast off Quequen, Argentina (Lazzaro, 1973). Below are listed the specimens examined of s. civitatus (N=292) in this research. From the east coast of Florida (FSBC-4352, 2 specimens, 122-133mm; FSM-21819, 1 specimen, 130mm; FSM-13051, 3 specimens, 130-140mm). From the Southi': Carolina coast ( GMBL-73-116, 7 specimens) • From the southeastern coast of Florida, below Cape Canaveral (FSM-12845, 2 specimens, 128-135mm; FSM-1213136, 2 specimens, 103-132mm; FSM-12847, 3 specimens, 130-135mm). From Apalachicola, Florida to Laguna Madre, Texas (FSU- 20885, 2 specimens, 413-79mm; FSU-20946, 1 specimen, 117mm; USA-01905, 25 specimens, 90-llBmm; USA-01450, 3 specimens, lll-122mm; USA-01225, 2 specimens, 115-llBmm; USA-01136, 2 specimens, 112-114mm; USA-01899, 204 specimens, 74-lllmm; USA-00893, 2 specimens, 110-114mm; USA-00847, 5 specimens, 110-119mm; USA-00826, 2 specimens, 109-ll8mm; USA-01177, 3 specimens, 105-122mm; USA-01481, 1 specimen, 102mm; USA- 02495, 4 specimens, 96-105mm; USA-00807, 6 specimens, 110- 116rnm; USA-02570, 4 specimens, 73-139rnm; USA-01421, 5 specimens, 102-114mm; USA-01077, 4 specimens, 93-lllmm; USA-01445, 4 specimens, 102-117mm). Symphurus collected by the author were immediately preserved in one percent formalin and transferred to ten 19 percent formalin in the laboratory to assure body tissue hardening. After ten days, the fishes were placed in 40 percent isopropyl alcohol and·stored. Some samples collected aboard. commercial vessels were fre.quently in early stages of. decomposition. These samples were immediately hardened in 10 percent formalin and later stored in 40 percent isopropyl alcohol. Specimens were, or are, in the process of being catalogued and archived in the museums of the University .of South Alabama and Morehead State University. Specimens. from museum contributors may not have been preserved in the manner described above. Keys by Ginsburg (1951), Gallaway, Parker, and Moore ( 1972) , and Topp and Hoff ( 1972) were used· in species identification. Ginsburg (1951) and Topp and Hoff (1972) were accepted as primary authorities for identifying specimens; however, a key prepared·by the latter is restricted to those spe.cies collected from the eastern portion of the Gulf of Mexico. Scientific names used are according to Bailey, et al. (1970). Diagnostic taxonomic characters included fin ray counts, scale row counts, standard length, head width, snout length, diameter of the orbital, siz·e of the opercular opening, vertebral counts·, body· and fin coloration, gill raker construction, dental configuration, 20 branchiostegal ray counts, and the position of the angle of the mouth. The head length, caudal fin length, body depth, preanal length, and postanal length were measured and expressed as a percentage of the standard length. Counts and measurements, where applicable, followed methods described by Ginsburg (1951). Measurements were made with dial calipers and millimeter rule and were rounded to the nearest ,05mm. A modified method of clearing body tissues and staining bone material was used to study inter- and intraspecific osteological differences. The technique employed: followed closely the procedure developed by Davis and Gore (1947); however, it was modified by using some of the procedure developed by Hollister (1934), As a result of this modified preparation, it was possible to make vertebral counts and observe cranial similarities and differences, X-rays, although not as detailed as clearing and staining, were used as an additional method of observing osteological variations in 23g specimens, Samples were placed on G.A.F. nonscreen Monopak X-ray film and exposed to X-rays from a Fischer GA-15 unit, for 1,5 seconds at a distance of 76.2cm (30 inches) with a peak kilovolt of 65 and a milliamp second reading of 10. The film was than processed by standard developing techniques, and was viewed 21 after drying. Small individuals were a special problem; the density of their skeletal systems is not great enough to show up on X-ray film, Primary field observations included surface locality of the capture site and depth of capture, Off shore localities were determined by Loran instrumentation and the readouts were converted to latitude and longitude coordinates, The water depth and depth of capture were determined by fathometer readings. Data of lesser importance included bottom type, weather conditions, water clarity, and time of capture. Capture sites for§. plagiusa ands. civitatus were plotted, using a color code, on National Ocean Survey (NOS) charts #1114-1116. Samples taken in the area covered by chart #1115 were also plotted on NOS enlargement charts #1263-#1271 because they provided a more detailed view of the larger area shown in chart #1115. The total range of the two species was divided into five zoogeographical regions. The boundaries of these regions were established from population segrega tion patterns shown in Chapter 4 of this paper. In addition, Ginsburg's (1951) zoogeographical regions were considered in boundary identification. The regions are: I, Long Island, New York to Cape0Canaveral, Florida; 22 II. Cape Sable, Florida to Tampa, Florida; III. Apalachicola, Florida to Laguna Madre, Texas; IV. The South American coast of Brazil, Uruguay, .and Argentina; and V. Southern Florida below Cape Canavera·l, the Bahamas, the Antilles·, and Central America (Figure 2), A Wang 720C advanced programmable calculator was employed to perform three standard statistical tests to determine significance levels relative to inter- and intraspecific variations. Independent 11 t 11 tests were performed to determine interspecific variation signif icance. Intraspecific regional variation significance was determined by performing: one-way analyses of variance (ANOVA), and. if significant variation was found between regions, Duncan Multiple Range Tests (POST HOC MEAN) were performed to specifically identify the regions involved. All statistical analyses were two tailed tests and were made at the .05 alpha level. 23 0 -Region I ~ -Region II -~ -Region III ~ -Region IV ~ -Regi:onr. W Figure 2, Zoogeographical Regions of Symphurus plagiusa (found in Regions I, II, III, IV and V) and Symphurus civitatus (found in Regions I, III, and V). ~ • T 7 r· ; .J. .t. ). J. !1.L_~·,1_~.~ . LJJ~J'.:.. i"iH?.J -1,s'rqou II , .- , -):is U::rou I •·. CHAPTER IV RESULTS Interspecific Variations Fin Ray Counts: Dorsal and anal fin ·ray counts of§, plagiusa and S, civitatus overlapped ·considerably (Tables 1 and 2); however, S, civitatus tended to have slightly higher counts, The dorsal fin ray counts of s. plagiusa ranged from 82-92; §, civitatus ranged from 83- 96, Anal fin ray counts were, respectively; 65-81 and 70-80. The results of independent "t" tests (Tables 17 and 18, Appendix) showed the variation between the two species to be significant, relative to these two morphological characteristics (dorsal fin ray counts·, !•9.88, df=364, _£(.05; and fin ray counts, !=12,52, df= 360, £<,05), Pelvic fin counts of both species were constant at four, Symphurus plagiusa had a caudal fin ray modal count of 10 (range 5-11), and-~. civitatus had a modal count of 12 rays (Table 3), Caudal fin counts are very important as Ginsburg (1951) used them as a primary consideration in segregating the two species and this study bears that out, An independent 11 t 11 test (Table 19, 24 I ::ip,H1?, _pc,(JL2 pp,,p onp • .rwi;oJ..prnr. sa u:ruapn:r,r; ( Jo2r) -n2sq ,::pern ,;a s hi• .fl!ilH.,1 conup ot is 1.0~~ (~epre ))• canqo1·iru conu~a s~s ABL1 ~0JlUP OL JO ·11.suHs ~-JJ)\ euq_~· C:fAJpsprra µsq a ~CQbJ . , , 'f"'o. l}::;' q:v,}e\l'' E ·!· o~ ! suq :.,:ru MJt. com:.ipJ. ~"JS. i:S. i:fi:= ~Q-ijO' LJJS L68DJ~2 Ot JJJqabsuqeDp uP,1 pe2pe (~BPJ~B I~ ' ()D' \iI.lSJ L:flJ .LSi, :_corrupa l•\61.6' ;,;.sa!JGCP:f/,6Jll 1 ',)~-;:;r Sllq .LUJJKdq I eJrt;.pr-Ji-A µr~JJtiL con11i:-2· ,LJJG qoJ.,8Z] t:ru r.,vi'L cVJJlJP-8 ot, 8· I (,I,' ~,:p.J f5l:l;~ c9nup:: L)ULt?BJ SI.lg B!JSJ l::,:flJ L\J,\ CC•HIJ(.'8 Table 1. Frequency Distribution of the Number of Dorsal Fin Rays in Symphurus plagiusa and sxmphurus civitatus. Distribution Species 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 s. plagiusa 3 7 11 22 47 30 18 19 9 2 ...g,. ~\g C'>., §. civitatus 1 2 3 6 22 13 46 39 32 25 6 2 1 :3p. gl ~ ;;i -:2 ;s: i g s. plagiusa 1 3 1 8 37 35 35 32 7 5 2 1 1 l\) §, civitatus 6 11 15 47 42 35 30 5 2 1 VI 1£,E..".i(;~· 'lr:i r:eCm;;;, srfj 1o nc.tJuJlrrdciG .I ~I.cI.s'l "f_:::;.:: £,;}_ J'!._;~ o r:-.~ ..-n;dqfi~V;;: D.:.1.8 ------S.8 ·------. ------·------ s: s:s. 0' s: .[ ===-==-======- Jsr::1 1c. ·redn:u;~ Grlj 'lo nol;:tudl'!jaiG ~,~H1supsrr'i:i .S s.[ds'I • i?,.Ud"s.:t lv .t!J 8u-:rrrf:=:n:y2. bn.s ====·-~--======·=·==---=----- no.t ~t udl'l.j3 .I:CT ,, "<' J, I l ,." c'. I :,. C. ,:(_ 2. € 'i'( C i ( I :..,,, ,- I s. ( oe (( --;.>('' \.:1 cl ll C: ___ " ______-.--r---• --·--•--c•- --·· --- - -·---·--- 26 Table 3. Frequency Distribution of the Number of Caudal Fin Rays in Symphurus plagiusa and Symphurus civitatus Distribution Species 5 6 7 9 10 11 12 13 S, plagiusa 1 1 6 7 155 1 _§. civitatus 3 2 13 188 27 Appendix) showed a large level of significance (t•33.60, df=374, P<-05). Caudal fin morphology may be seen more clearly in Figure J. Scale Row Counts: Scale row ranges overlapped interspecifically. These counts were often difficult to obtain because many specimens were partially mutilated and possessed only a few, if any, scales. The condition of the specimens required the counting of scale pockets, a method not always as accurate as desired. These data (Table 4) showed that S. plagiusa had a range (73-92) higher than S. civitatus (70-SJ). An independent "t" test (Table 20, Appendix) showed significant variation between scale row counts (t=l0.61, df•301, £ <•05). Orbital Diameter: This characteristic has not been extensively examined in other studies, but did prove reliable for separating adult specimens of the two species. Symphurus plagiusa usually had an orbital diameter of approximately 3mm; the orbital of S. civitatus is closer to 2mm. This measurement varied slightly, depending on the size of the fish. This characteristic is not reliable for immature or juvenile samples, but the orbital diameter of adult specimens was so consist ently different that it cannot be ignored. Angle of the Mouth: This characteristic was considered by Ginsburg (1951) as being of minimal taxonomic value . In most S. plagiusa samples, the mouth ' ,, 'i .. . on.f~~.:.f) s··r-:i:;·)f']:j-;;:•.li."! 'lo ;,-f'\T~~I. .7~'Tc-.i ~; b£:l•J0rI:..: (~~sr:,.--1~•C1tr:1 t•...,,.. _,,_ - ~•fnf""' r:q"::',~__. f"!i•"1 .. ;"r.,rf(! .. f')/!r ., ·,·')- t r,h,,r~,~l I ,., • . ... ·- - __, • - ,~~m,,.., cr···,n. . . • • • .• ·- • \) . .... ""i 11. r.1 • ( (0. q ~J:.~ c.-~ • ( sfJJJ;d:'~ rr.t v:Irr<"::1.l::' bs• r : ·>:_C •·j ,.ii.re. e s:··11s--r ir10"1 e.L13,I::~ :c.anuo:J wof-1 eIG~~---- f),1 .::~-u::,_t':r.r.,:•f:, na,~lo 9'!5W 8jfHJ0:) 98srl'l' •tJls:Jt'1i"::i90~'19~f'!l bG~'!r:1I.i:Jum v: fJ.l\lj,ysu &"J"SW ::.nr~111J'~9ca! '-':N.P.m o;.:u~~!:d n.is~do r::0.[.Jj-brr-)D srf'I .c.,r--de:=-,a ,-:tns 'Jl , ~rsI e. ,,::J.no bsr..eozGoq .0.rrs •.• i•~ t!"'l\iJ :,f.f-.'l'' f, W\. -...,.< J ., •~ • l.,~ 1o ;:;,11.tJr1110::i e,rl;J- bA,ir,ps~ snsml~~crn 9jj 1o :~;~r:· b -:-.nn·i'l' • ~~1rr le.ab ~s ~i.'.lC .. ffJ:.--,~c; 2.r, 8."'{t,w.f.s 0on borij ~)1i'J f; 1 (~Q-f~). . o:~~n~ ~ bnrl .,c,U--~~~"., h1sla .!:,_ .:;;=;dj bs~tr,,dc: ( .i~ eldB J) '' :i'' Srro!fl•·J'feiml r:A. • ( 03-01 l . nw;,;I ~'flu'-f r,:::o ~-f!fVf!.[~Oq~~• J•1 UDG• ~CT..C.'.JGrrriq9~• < "IO.1 flJ.db.CC • •1 ~•I :..-;,:J.C:rso nc;· biH'I yfls.c..rEIJ •_§_1£_,L!:£.fq ~!Jr!gm:{~ ,ael!'>sqc ·:-:: 1 :J,-._-t_tv.: ::, ,i-. Jc; f.sj td'"To srlj ;mrnt ils..:1Mrrt txorrq013 1o •r;)jtJr.1<:;Jb --·····•- -- - . ,Y,.[,~rr·-..i. .t[G '.:-,;-)['rsv d11S\me·n1?.nsm elriT ,mm~ od '1ec:o.[!) al • DL~:~_·:s·~~~ni~~. • j ' CL,1l. ..,,,. ,,,axt.., ., srt. j ••i.o ssxe• ArJI no r11ro~s0so. . . t1!Jc{ ,.es.I.:-:r,11~~. oJ. ..tnov~sr. "!O e·-rrr'--'"cra;n.i· '"IO~T oirJr.;J:.Ce 11 .1.-)n el .-jcj~.ao~ Gt ~~w ~rremf~enE j_[isbs 1o ~3JG~!~lb lsjld~n e,i~ • .bo'":f("ir~l;·d: s;d ~!oni1£:J jl .Jr;r!i .J:1e,.:2t1'Lth -v:J01.1.s ~t:>l•/ :-; CJr:-:f".raj~n·rsri~ cirVi·· :rf.:t.uo!•i ~~-i.1L12. el:rin!1. .s;:rf:itm 1o ·nf!.t.sd ae (lC('I) ~-:rrd;~n_;.D yd hs~.:1r,lerr~~ •f;}.:rom e~~ ,as.[cmsn. -.v,Baul,ela - -·- ¥_.,. .{~ ,:rc?c,;l~ trI • [lcrltv ~Jmor1:):r,r;j 28 . '-· - ,- - . ' .. 0 ,,'' Figure 3, Caudal Fin Construction of Symphurus }lagiusa (above) and Sy,m:phurus civitatus (below Table 4, Frequency Dietribution of the Number of Scale Rows in Symphurus plagiusa and Symphurus civitatus, Distribution Species 70 71 72 73 74 75 76 77 78, 79 80 81 82 83 84 85 86 87 88 89 90 91 .92 ~- plagiusa 2 6 2 18 11 8 16 22 13 14 9 8 2 1 S. civitatus 1 5 12 14 11 26 40 19 8 15 15 4 1 30 angle extended approximately one-third of the way under the lower eye. In~. civitatus the angle extended at least one-half of the way to completely beyond the posterior margin of the lower orbital, These observa tions varied regularly, and were unreliable. Teeth: Dentition proved to be of little value in separating S, plagiusa from s. civitatus. Teeth were present on the upper and lower jaws on the blind side of both species. On. the eyed side, the teeth extended approximately one-third of the way onto the anterior portion of the upper jaw in S, plagiusa, and one-half of the way onto the anterior portion of the upper jaw in S, civitatus. It was often difficult to obtain exact measurements. Teeth were completely lacking on the lower jaw of the eyed side of both species, except in a few~. plagiusa, where one or two teeth were found. It should be noted that teeth on the upper jaw of the eyed side of ~- plagiusa were often found in rows of three or more, producing a stacked effect, Symphurus civitatus always had only one row of teeth. Ginsburg ( 1951) used dentition as a separatory characteristic in his key, Vertebral Counts: The vertebral counts showed an overlapping interspecific sequence, This negated the use of this character-for species identification, although S. plagiusa had a slightly lower vertebral count 31 than did S. civitatus (Table 5), In S. plagiusa the modal number of vertebrae was 47, although nearly as many specimens had a count of 48, Most specimens of S. civitatus had counts of 49, although many· specimens overlapped~- plagiusa with 4S. This slight variation proved significant (Table 21, Appendix) when subjected to an independent "t" test (!=13,31, !!f.=184, p<.05), The precaudal vertebral count was 9 in both species, with the exception of two samples of~. plagiusa in which the count was 8, There were occasional deformed vertebrae which could have caused some error in making accurate counts, One such deformity_occurred when two vertebrae were found fused together, emulating a single vertebra. This deformity was detected after two additional neural and hemal spines were found extending from the seemingly single vertebra. Branchiostegal Ray Counts and Gill Raker Con struction: These characteristics were equal and constant in both species. The branchiostegal ray count was six, and was of no value in distinguishing the two species. The gill rakers were reduced to short stubs, Because of their similarities, they were of no value in evaluating variations between and within~. plagiusa and S, civitatus. Body and Fin Coloration: The bodies of specimens of both species were marked with vertical bands; however, en;~ pc.,~µ abGG1C8 RGLG IB~J.. J(Gq !.\:r~·p AULt~:rce) J)9"tjq2: JJOHGAer..· ! BOqa -£1/J{! -f:(i;-- coj'CJJ..~rfroiJ: .1}18 pvq:[G2 01_ cl!J6C-flJ:6iJS ~-·--·· --- SUO. n•" _: r "-r 1"s,,frn · ·a:rrrn'.J e 11.eLr.epr,o • · f ,,\GJ..6 :t,vnJJq l,flc:Gq ~otter.psi.•' GlllfJJUf.:flJt: 9 8-ftlliJc .!V,J:.f.GpJ,'2 • ' I. ' '. . c:on::ir.2 • QlJG 8f!CfJ QGtOL,lli:ff.A ·occrr.i..:..Gq !,l.J.IGJJ i·HO /.GLf.6j)!.IS6 b •o:.,,,,_ l ' GTA1f:S~Jl?. µBq CO!HJ.C\3 04., r·Q~ S-Jt2JJOI1~:J mtnJ/:. 3bBC.fI.l19jJ;·; 32 Table 5. Frequency Distribution of the Number of Vertebrae Occurring in Syrn?hurus plagiusa and Syrnphurus civitatus Distribution Species 45 46 47 48 49 50 51 52 S, plagiusa 2 11 53 51 12 s. ci vitatus 43 56 7 1 33 ~- plagiusa tended to exhibit this feature more markedly than~. civitatus, Symphurus plagiusa usually had a dark black opercular spot, sometimes divided into two or more blotches. This was also described by Topp and Hoff (1972). When present this spot is considered· a dominant character istic, distinguishing S. plagiusa from all other western· Atlantic cynoglossids (Bohlke and Chaplin,· 1968; Ginsburg, 1951; Topp and Hoff, 1972). Symphurus civitatus lacked this dark spot completely, although there was a dusky opercular blotch, It was observed ·that the blotch was, not a true external coloration pattern, but rather an internal visceral shadowcast and part of the gill apparatus, The opercular spot on.~. plagiusa was actually a' melanophoric congregation on the scales; s·, ci vitat us lacked these melanophores. Both species have dusky colored fins, sometimes dotted with black or brown specks, These small and inconspicuous speckles should not be confused with the larger spots found on the dorsal and anal fins of· Symphurus diomedianus (Goode & Bean). Proportional Measurements: Ginsbµrg (1951) stated that proportional measurements were of little value in separating the species of western Atlantic· tonguefishes; The proporti-ona1 ·measurements «made ··in· ' this study supported Ginsburg's conclusions, because 34 there were only slight meristic variations between the two species (Table 6). Statistical analyses were not performed on the proportional measurement percentages, because the percentages, even at a significant level of variation, were of little value in evaluating differences. Vertical Distribution: A major distinguishing feature, although not morphological, was depth ·of capture. Symphurus plagiusa was collected at.depths up to 16 fathoms. One specimen has ·been collected· in ·the northern Gulf of Mexico at 26 fathc;ims (R/V Tursi ops, Station #7102, FSU Collection #21291). Symphurus civitatus was never collected in shoreline seining, although many were· captured (R/V G.A. Rounsefell) just offshore at 10 fathoms at the mouth of Mobile Bay, Many specimens were collected from· several ship channels in Mobile Bay and near Dauphin Island, Alabama. The maximum depth of specimens examined was 76 fathoms, from south of Mobile, Alabama (R/V Tursiops, Station #7109-07, FSU Collection #20885). Often the two species were found together, indicating an overlapping vertical distribution. Symphurus civitatus was not found in the shallow waters where _§. plagiusa is so abundant. Statistical Analysis of Zoogeographical Region III: Independent "t" tests.. (Tables 22, 23, 24, and 25, Appendix) were performed on data from specimens collected I J r,;ml .:l '!o,ro'l:!:; sr!J nc Cfl1,1°w 1· 0 .r t,q 'lo Jev::11 dng::-J1.tn0l2 n jr; nsve t 2e~,1:;0rsJ'Y8: e-rf;} Gerrr-;:,ed .-::e::;rr.e::rG·:t·;__r:b i]l'Iljr;.rrJBve rd: 9LJ.[Bv alti~!ll. jlo' S'1'9V-! trtoJje,j·rrr,v ' ;:~1:r .i: d c.:: .u"J r: J: i:,te, j: b --:r c. [J·,m t~ : rro. .t .:1 r.rd ..t'I ,i c; .tQ._ Js::, J d'"'; fl\' I , s'tu,i·{~P.~~ Co cf~:e;et 2.r.~~, ,ls~.t~o.Ioifo'1om sort; rl;Juoftjl.s I s-,:.u.:JGe'? d.£ -:.~· r~u er{~)qeb jr, Q9,j::,9J_lo~· est{ &!:f!L'rlf1.[£ ~1-~f£.!f!!:·~[ ' ' wrer,.;!•rcrr r;ri.:J rd: bs;J::,9Jlo::i fl98d 2flrl _r1sml:p0qa sn() ,2m0r!J,:;'J. r.rol~Jr.;rr ,nqolc....,.u'T V\H) ~morfjs1 as; j_s .f18lxeI✓1 ':to ~L.[r_r{~ ' ' c!;.rj0iJ-v l~ er.r'rrirlcmrvG • ( I\'S:IS:~ no'.i:~::ii-J1o:l U8'•1 , S:01';'\ ·------' ·--- ·• I J ' e·rnw ·-:nr-m 'rl;q.r;o_rfols ,~rr.i:n.rea ,eolle-roria nl.be.:loollo::> ~svon ! ( lle1earruo.fl ~A ,D _V\fl) be'luJqi;::i e,•rs·'.,1 LnemJ::n,,,2. ·•urni'il ,yr;U oI.rdoM 1o flj,uom · erfd ;Js emorfJs'J i h!H~ '-:G-'.: ~J_[j (:loi-11 ri..i: elenru:;ri~ q.irla la'Isfse morx1 bG.:t~elio:) I ':o fl;?ueb mumJxsm erfi • sr11.Rd£ll, ,bhsleI njrlf!IJ~fi ~£er: , RI1doM 1o .rf.:.tuor: mcJ"r1: , amorf;J.s1 Cl\' ZGW JJsnlrnsxs ansm.raeao ' ' I f10.[;)~j<,J ro'.i UE;'•l ., 'i'0-(?01,~:; rm.i:j11je ,eqOf!l'!U~' V\R) l3!'!H;dr,J.ft. I ~~sdj~~oJ bnuo1 e~ew ael~n~a ow~ s~j nej1Q .(e~80S~ .nc.f.jrH.U·-:rje.tb lrv~lj'J3V ;;1rrlqqGifr0vo r::s ~:u::ldA~J:br·J. i 1!.~.e,,·r,;•,; wo_Ll.srfe Drf;J rd: brrw,'1 ,:!on ar,w ~+rjg;J.i:v_i::i !lu'!rrrlam':'._2 • ;:: rmr,mrds S~GrhrJ r10 ~-~~s H __ f .s ~ l11n ri'I·2 os:qoo~ __ 'i.o_~ e J exisnA_ If:, l ~, <~ l J si:12 I ,?S bns t.µ~ t(S tss 2eJda'1') eJcej "d~:1d·neb11eqshnI :J.1I ' . hAd:'telioD 2nernl8t.1(fc. mo"'11 _f;jf.;.b rro hem,yo'l¾eq e•-rgw ( xlhnso.-L~ Table 6. Mean Value, X, in Percentages, of Standard Proportional Measurements in Symphurus plagiusa and Symphurus civitatus .. Proportional Measurements Body Head Preanal Postanal Caudal Head· Orbital Opercular Depth/ Length/ Length/ Length/ LengJ;hJ___ .Length/ Diameter/ Opening/ Species Standard Standard Standard Standard ·Standard ·Head Head Head Length Length Length Lenl?jth Length Width Lenl?jth Length (BD/SL) (HL/SL) WRL/SL) (POL/SL) (CL/SL) (HL/HW) ( OD/HL) ( 00/HL) .§. plagiusa 30.58 17. 54_ 21.88 78.15 10.27 71.47 14.35 38~04 (N=l71) s. civitatus 30.26 18.07 22.01 77.68 11.11 73.82 12.07 35.45 - (N=206) 36 in zoogeographical region III by comparing dorsal fin ray, anal fin ray, scale row, and vertebral counts. These tests were performed to determine if· interspecific variations between ~. plagiusa and s. civitatus were significant. An important reason for selecting this particular region for an independent evaluation is that a large number of samples of both species were examined from this region, and such a test would eliminate the influence of extremes. The analyses showed the varia tions to be significant (dorsal fin ray count, t•5 • .06, df=256; anal fin ray counts, 1•9.22, df•254; scale row counts, 1=7.68, df•219; vertebral counts, 1=10,79; df= 181) at a "p" level ( ,05, Intraspecific .Variations .-_: Symphurus plagiusa was collected, or reported, from all five designated zoogeographical regions, but~ civitatus was collected only from zoogeographical regions I, III, and V. Interspeci.fic variation was limited, but obvious, In turn, there was some variation within each of the. two species which is summarized below under each specific name. Symphurus plagiusa Fin Ray Counts: The dorsal f'in ray counts of§_. plagiusa, when divided into zoogeographical regions, showed 37 little variation (Table 7), Dorsal fin ray counts of specimens from Regions I, II, III, and V were concen trated from 86-88, Symphurus plagiusa from regions IV may have a slightly higher range; howeve~ data were insufficient to provide definitive conclusions. The variation in dorsal fin ray counts was subjected to a one-way ANOVA (Table 26, Appendix) by comparison of regions, and significant differences were found (F:6,57, df=l79, .P.<,05), This test was followed. by a POST HOC MEAN evaluation, The results of these tests showed significant variation between regions V and IV, and regions II and IV (Table 27, Appendix), The anal fin ray counts (Table 8) showed a varia tion pattern similar to the dorsal fin r;cy counts, An ANOVA test (Table 28, Appendix) showed significant varia tion (!,=5,63, df=l75, ,P.(,05), The POST HOC MEAN test ( Table 29, Appendix) showed. significant differences between regions IV and I, IV,and II, IV and III, IV and V, and II and V, The small number of samples available from region IV may be the reason for these differences, Further testing is suggested when additional specimens are available, The caudal fin ray count was predominantly ten, although there was some variation. Five ·specimens had nine rays and ten basal ossicles present, and three of Table 7. Frequency Distribution of the Number of Dorsal Fin Rays in Symphurus plagiusa, Segregated by Zoogeographical Region. Distribution Zoogeographical ~egion 82 83 84 85 86 87 88 '$9 90 91 92 I 1 1 2 4 11 10 6 4 II 1 4 7 7 24 2 3 1 2 III 1 1 2 11 15 16 10 14 7 2 IV 1 1 1 V 1 4 3 \,J 00. Table 8. Frequency Distribution of the Number of Anal Fin Rays in Symphurus plagiusa, Segregated by Zoogeographical Region r Distribution Zoogeographical Region 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80, 81 I 1 9 7 6 9 3 2 1 II 1 2 7 10 12 12 6 1 III 1 1 18 16 17 17 1 3 1 1 IV 2 1 V 3 1 40 these specimens had a short stub radiating from the ossicle. The remaining two specimens, with nine caudal rays, had nine basal ossicles. Six specimens had eight caudal fin rays, Four of these.had ten basal ossic'ies; one of the four possessed a short stub, and, in additi~n, two fused rays which formed a single ·vay. Of the remaining two specimens, one had nine basal ossicles, and the other had eight. Three other specimens were also variants; one had five caudal fin rays, one had seven,· and one had eleven. These fourteen specimens, all with caudal fin ray .. counts above or below ten, were determined to be.§.. plagiusa by the presence of an opercular dark spot, An additional factor supportive of species identification was their depth of capture; they were all collected from shallow water outside the range of S. civitatus, Scale Row Counts: The intraspecific distribution patterns of the scale row counts in specimens from all five zoogeogr!lphical regions are as variable as ,the dorsal and anal fin ray counts (Table 9), An ANOVA test (Table 30, Appendix) showed significant variation (!■ 2.6~ df:lli,O, .E(.05). The POST HOC MEAN test (Table 31, Appendix) showed significant differences between regions V and II, V and III, and Wand IV. Table 9. Frequency Distribution of the Number of Scales Rows .in .Symphurus plagiusa, Segregated by Zoogeographical Region., Zoogeographical Distribution Region 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 I 3 4 2 2 9 2 3 lj. 3 II 1 3 1 8 3 2 4 4 7 6 2 2 III 1 2 1 8 6 4 9 5 7 4 3 3 1 1 IV- 1 1 1 V 1 3 2 2 42 Vertebral Counts: Most specimens of s. plagiusa examined;;· showed a vertabral count of 47; some had a vertebral count of 46 (Table 10). Statistical analyses (Table 32, Appendix) showed no significant variation in vertebral counts in specimens collected from all five zoogeographical regions (E=l.64, df=l38, E<•05). Body and Fin· Coloration: This characteristic was highly variable in~. plagiusa. Samples available for examination had been preserved for varying lengths of time, and had lost some original pigmentation, The body was generally marked with either brown or black bands. These bands were either continuous, broken into sections, or appeared as a series of blotches randomly distributed on the body. The bands were wider than the dusky white spaces separating them, and were found only on the eyed side, The blind side was without pigmentation patterns, and was yellowish white in color. There appeared to be no relationship between geographical distribution and color pattern. There was variation in band color, , · The.' specimens collected 'from region III had dusky brown to black bands; those from regions II and IV had much darker brown bands. On:'. specimens collected in regions I and V, the bands , were less distinct because of melanophoric dissipation into the spaces between the bands. Fish from regions I b;;r! 8I!JOZ i I,',.\ 'l 0 ;:Jnuo::, r~,--dsd~sv £, h8l•.iod2 ?-1,.b:t1t1lm.GX9 "' '' ' 89P ..t[Jfifi-f.t; Jn~.t.;.Je.t.:.tr..:l2, • ( O.C sids~·) 8.1 1o ;) (l!J(l!) If; '! c! ,, ,i 'I fl V . ' ''r.,J 8-i.:J;:.nsJ ;~_.r,ly•r.sv '!o'J bev•rnas'Tq nescf •brirl ·nol011r1.tmsxs , i'lrrol;;:Jea ojfll l'!fi,fo•rd ,urorrnldno:::, '1ed.:UP. e·rnw abr:s,:J eaeri'.l' s;J.t:iw '11:,laub 9r/,t rrnr!.! 'Tsbi:w 0-:rew abnnd' erlT .~bod srf;J :10 brrn ,~o.,'dtrdl-:rJe.tb Is'.llrfq;,-r;ios;g neew;Jf!d q.tderiol.JGls·r on • sbmid ,r.ro·rd '1,,.1:•n,b dou!l! bBd VI .brrn TI ?rK,12,e•r mo":!1 s:·r,:-:;:: .:br:;c;d erl.:i , V b.rrn I enol~s·r ril: /Je;:JoeJ:Ir,:::, enem.too-::8 i no ! • 43 Table 10. Frequency Distribution of the Number of Vertebrae in Symphurus pla~iusa, Segregated by Zoogeographical Region Zoogeographical Distribution Region 45 46 47 48 49 50 51 52 I 3 10 13 5 II 3 5 3 III 2 5 37 37 IV l V 6 1 and V showed a rather uniform brown to black color; the banding pattern appeared as a faint shadow. Fin coloration ins. plagiusa was generally the same for specimens collected in all regions except region II. The fin coloration in specimens collected in region II was consistently more prominent. Proportional Measurements:. Except for a single immature specimen (69mm) collected in region IV, pro portional measurements for S. plagiusa .were .quite uniform (Table 11). Symphurus civitatus Fin Ray Counts: Dorsal and anal.fin ray counts (Tables 12 and 13) of specimens collected in the three zoogeographical regions where S •. civitatus occurs (I, III, and V) showed no obvious variations when subjected to an ANOVA test. The Anova test (Table 33, Appendix) showed these variations to be non-significant (dorsal fin ray counts, F=.36, df=l91, £<,05; anal fin ray counts, E,=2,17, df=l92, P<· 05) • These statistical tests may not be valid due to the low "n" value in some regions, In a total sample of 201 specimens, the caudal fin ray count was 12 in 188 specimens, while some (13) had a count of 11. One specimen, with a caudal fin ray count of 11, was distinguishable from~. plagiusa by the t:ffJ L9'1~ corn.1p t"~sa J:_, .:ru Jq!ij nbsc1 U:G1J~1 • l•lJ.!:f"JEi 8t.. j.1.16 ( JJ) ' 1 1 GOITU~8 1=')~ ql=JaT' • I pµaau AHL~·~rroua ~ops (J,9!)jfo8 JS suq J) J, Ot 2:0sc:rm0ue coy:,s-:,:-sq TJJ DOJ.8\l J CTlJ(J $l.JS'J t,:p-J .,_,,;:..u.- ('• ·. -----r.1s i:I 8JT LG UJS--- lJ f.' C: ,,'"8JJJ[' . ''' Table 11. Mean Value, X, in Percentages, of Standard Proportional Measurements in Symphurus plagiusa, ~egregated by Zoogeographical Region. Proportional Measurements Body Head Preanal Postanal Caudal Head Orbital Opercular Depth/ Length/ Length/ Length/ Length/ Length/ Diameter/ Opening/ Zooge ographical Standard Standard Standard Standard Standard Head Head Head Region Length Length Len,th Len,th Len,th Width Length Length (BD/SL) (HL/SL) (PRL SL) (POL SL) (CL SL) (HL/HW) (OD/HL) (00/HL) I (N•38) Jl.63 17.08 21.80 78.14 9.66 68.64 13.82 37.02 II (Na49) _.;.,'":30·,2g ·• 17.82 21.51 78.47 10.86 72.36 14,78 37.56 III (Na66) 30,38 17.52 21.60 78.38 10.34 73.07 14.52 38.90 IV (N=3) 29.15 16.23 23.74 76.25 9.20 71.33 13,39 28.19 V (N=8) 30,30 17.45 21.65 78.33 10.23 70.50 14.30 37. 29 Table 12. Frequency Distribution of the Number of Dorsal Fin Rays .in Symphurus civitatus, Segregated by Zoogeographical Region. Zoogeographical Distribution Region 83 84 85 86 87 88 89 90 91 92 93 94 95 96 I 1 2 2 II III 1 3 3 4 21 12 39 36 28 23 6 2 1 IV V l 1 4 1 1 Table 13. Frequency Distribution of the Number of Anal Fin Rays in Symphurus civitatus, Segregated by Zoogeographical Region. Zoogeographical Distribution Region 70 71 72 73 74 75 76 77 78 79 I 1 2 1 1 II III 4 10 17 39 38 34 29 6 2 1 IV V 1 1 4 1 1 48 presence of 12 basal ossicles, The remaining 12 specimens of the group possessing 11 caudal fin rays (13 specimens) had 11 basal ossicles. Five other specimens, had counts of 10 or less. Two .specimens with a caudal fin ray count of 10 had 12 basal ossicles,. A single specimen, with a caudal fin ray count of 10, had 11 basal ossicles plus a short stub radiating from the eleventh ossicle, Two specimens had eight caudal fin rays. One of these had eight basal ossicles and the other had nine, In cases where the caudal fin ray count varied from the typical value of 12, other means of species segregation were employed. Depth of capture, orbital diameter, angle of the mouth, and other fin ray counts aided in identification. Scale Row.Counts: The scale row counts for~ civitatus were quite uniform (Table 14), ANOVA tests (Table 35, Appendix) showed no significant variation among the populations (E,=1,53, df=l74, £<,05), Vertebral Counts: The vertebral counts for S, civitatus in all regions were predominantly 49, although 42 specimens had counts of 48, seven had 50, and one specimen had 52 (Table 15). ANOVA tests (Table 36 1 Appendix) showed no significant variation among popula tions (f=.36, !!£=104, £<,05), r~ 1-~,;:, ~--H~rm.,. ,. • 1.., i~~) ··• µYd• r J"J· •cganT • .~ .. - . u~dr··••·e·..:, •• -. ,.,. _, -~ . ' r-:: J ., Table 14. Frequency Distribution of the Number of Scale Rows in Symphurus civitatus, Segregated by Zoogeographical Region Zoogeographical Distribution Region 70 71 72 73 74 75 76 77 7g 79 ao 81 e2 g3 I 1 1 1 II III 1 9 9 14 12 26 34 18 8 15 15 4 1 IV V 2 2 1 1 ..;:. , _-:"'_ r_ _\. .. ,I • __.. r _ , 'I)}' :3 ~ ~~ 1:. ' I') j_""!t"i~ "c rDJ:,. v ------·---·---.. ------·------..- (col::ic1~·":l .. 9~V•':, ':'"> ;- • • s.fi (T' \..' ------·----- . ------·------··------l [ [ I : I .r ., - t ------·--··_ ..,~------·------····· ·---- . -- 50 Table 15. Frequency Distribution of the Number of Vertebrae in Symphurus civitatus, Segregated by Zoogeographical Region Zoogeographical Distribution Region 48 49 50 51 52 I 1 2 II III 37 49 7 l IV V 4 'lo -:rsdrru1~1 9flj Jo no..tj!Jdl""L~fgj:[J '1[.!)!!f-,~Jpe•_t~ . cI s.Ioz'l' .~trjsjlv1o eu~Dda~v~. c! e~~dsj~sl ·------~---.. ______....:.,.,.., ;.. . IJ?.:.:•Jriq_Brr~osi\OOS yd b{~·J'S2 o":i::~• ec; n0l~1sH .aol.:t udJ:-rja.Kf lo: ldqs·-:;;1o~;ga0~\ ------·-- nol;geS-: ------ s l I II III VI V ______... , .... - ·• ____ .... _____ - ___ ·---· --·-----.... .- ··------·-·•--'-·------...------51 Body and Fin Coloration: The body color in all populations was basically the same. The color change described for §, plagiusa was not obse.rved in .§, civitatus. The body was generally dusky brown in specimens from regions III and -V. The banding pattern of S, ci vitatus was consistently ·lighter than that of.§. plagiusa; the bands of S. civitatus were continuous, rather than fragmented. The fin coloration can best be described as dusky. Proportional Measurements: Proportional measure ments, made on specimens collected from separate zoogeographical regions, show an interesting pattern (Table 16). Generally, specimens from regions I and W had mean values appreciatively less than region III. This was true for all proportional measurements except·· for body depth/standard· length and·postanal length/ standard length, These variances may be partially ex plained by the fact that only~three samples were avail able for examination from region I. However, an apparent difference still existed between the Atlantic (regions I and V), and northern Gulf of Mexico (region III) populations. No statistical tests were performed on these measurements because ·the ratios were of little importance in evaluating individual specimens. Table 16. Mean Value, X, in Percentages, of Standard Proportional Measurements in Symphurus civitatus, Segregated by Zoogeographical Region Proportional Measurements Body Head Preanal Postanal Caudal Head Orbital Opercular Depth/ Length/ Length/ Length/ Length/ Length/ Diameter/ Opening/ Zoogeographical Standard Standard Standard Standard Standard Head Head Head Region Length Length Len,th Len,th Len,th Width Len~th Length ( BD/SL) (HL/SL) (PRL SL) (POL SL) (CL SL) (HL/HW) (OD HL) (00/HL) I (N•3) · 31,91 16.99 11.37 88.62 8.61 70.55 10.17 30.35 II (N:O) III (N=l66) 30.24- 18.95 22.23 77.4-5 11.13 73,86 12.13 35.50 IV (N•O) V (N=6) 30.86 16,91 20.61 79.38 8.4-1 70.27 9.05 20.66 \Jl I\) .. ,.. .. J ',.),:: -•-") J J J,.; J.T"-,' ~.., ( •1i·:\ ?P) ; . .--,-,\ , .. .,., ) t'r\ 1 ! µr-\ t;;~ \ ( 1·1\ fff' \ \ l:!:h \-::i~ ( .J_ .... :,r·'-' • \ -·- \,~.. , 0 ,· \••- I J.Gf_:~- OU .·fO!Jt"r.rJ rs i.rS.r. !J r01;-s:pp -':'TO.CU~. .. . l.,B :J ::.1'\"J ~o:J '1G0~.:.. sbp;·c-aJ 2:..'~::1c;s.r.. 1 ., r. a:1JJ; :n,a :~~ 2 :Jq :_r;:_,q !{GUq }18--.1q' '.:s :._::r-; fGIJ~.r.p\ '"'rsDfqp\ r,0:1-icu\ tSIJi;:'p\ D:fE!.CS :~C ...,\ J :)<-.,".' 1~: .•- !...y <•.\ - . ' • - .-. I ~f't i\69" ;.r...ss-:.:--;y c~-rnq$J t1V v•-: [·~~:>;-;:.~;· ,~b,:;.,.~. .:;n·.1 .i;:., CHAPTER V DISCUSSION The results of this study supported the con clusions of Ginsburg (1951), lending credence·to his belief that the two forms are valid species, Dorsal and anal fin ray counts and scale row counts were important in distinguishing ,1!, .plagiusa from S,. civitatus, but the caudal fin ray count was the single most important species identification characteristic. However, there were exceptions in caudal fin ray counts and these exceptions should be considered when examining either species for identification purposes, Symphurus plagiusa and S, civitatus are almost impossible to identify by sight, even when collected together. Although it is difficult to separate the two species by sight, the probability of identification is· increased when multiple-characteristics are considered, The opercular spot, commonly found on ,1!, plagiusa, is a good indicator, but should not be exclusively used as a means of identification. The size of the orbital is an important species indicator for the experienced taxonomist. The orbital is notably larger in S. plagiusa than it is S. civitatus, The angle of the mouth is not a 53 pµsu :[ I~ :[8 ?",, C :[ i.ff-{if.im • 1µs $iJUJG 01·,. 1~VG wo rr ~ ,'J IH :.rop \; I I qt ·~._.:k·-r ·:] r:.:,:- p \lXOIJ OJ.;J:[ 2j? • .r,p 6 or.p:r PBJ :[2 uor.s.on, JtfLt~6J.. ~;J ,, h' • .._. ;.t,~ J ,,, ..., LPB obeLGJIJ9L atop' 2Auil5fjirEIT2 Drsf:frr5s sug ·~· c-r A.ft·.1;~n2 ·.:1J..o ~JHi::.~1r- 1 ZCilG.fGa i;_o;, :rqsIJPit:rcor.:roIJ bru,boeea • euq &DHJ tfD L&l Gonur.a auq 8GBJ6 LOM conu~a J)OL,lST ;n_ ?.C:18Tl:01, 54 reliable indicator, but it can be of some value when used in conjunction with other factors. Any one of these characteristic.s, if not all, may. fail in field analysis, and sight identification is not recommended as being conclusive. Morphological counts and measurements, including osteological observations, were of value in establishing new characteristics for species segregation. Proportional measurements, because of inherent variability, can only be used as subordinate aids. Vertebral counts overlap in the two species, as do the dorsal and anal fin ray counts, but they all can be used as identification aids. Hubbs (1922) suggested, and gave evidence to support the theory, that overlap between fin ray counts, scale counts, and vertebral counts was a function of variation in the water temperature in which individuals were spawned, Vertebral counts are not easily obtainable unless proper equipment or chemical reagents are available, Statistical analyses showed significant interspecific variation among the fin ray counts, scale row counts, and vertebral counts, Body and fin coloration are not a particularly reliable .separatory .characteristic. Symphurus plagiusa is usually more heavily marked, and sometimes may be identified by its darker bands and band configuration. -~~~n~l)' '.,,~JUIC,•-'•'- "LJ.N.J" 'p_[(~~-~_l~w.-.•i•~ J., • f\V.r,,u.f::iao!"l ;gn.i:dr.-.i: fdr,jas rr.i: s11Jr,v 'J:o s'1SW , Bno.i:.:Jsv•n,r-:do ls::i.i:~-io.ros.:lco .ehin sisnJb~odue as beeu ed ,bsmw·re s~rsw 2l.1rnb.i:v.i:fm.i: rfolr{1/1 rl.i:, s•m.J,,--:sqr.:s;J •rejBW • eld.e.IJ.svs o-re ::,d'n!.?~.Ss'I JG~ilrnerfr:i ·ro .:tnemq..i:ur.~3 I ' 2r:~uJ:1;:;.~ c~.r•1.:.1.dq1r:y2 • ~.LJ8l ..Jflj:)£r!Sd!) ·,:r;ro.:'S'I.Sqse e.[dnlJ.srr I 55 Symphurus civitatus is usually faintly marked; however, some specimens of§. plagiusa also show this condition. Depth of capture is the second most reliable me.thod of identifying the two species, This is especially true in waters six fathoms or less, because §, plagiusa is the only species .of Symphurus found in shallow water along the eastern coast of the United States. When the two species are.captured·together, or individually, during offshore sampling, other character istics must be used, The data showed significant differences (in dorsal, anal, and caudal fin ray counts, and in scale row and vertebral counts) between S, plagiusa and S, civitatus. Although macrovariations are minimal, this is the case in all species of western Atlantic Symphurus (Ginsburg, ::1951; Topp and Horr, 1972). -· Ginsburg (1951) stated that§. civitatus is very similar to Symphurus plagusia sensu lato, a species known from the West Indies and Central America [Symphurus plagusia plagusia (Bloch and Schneider), sensu stricto], and from Brazil and Uruguay [Symphurus plagusia tessellata (Quoy & Gaimard) ]. Ginsburg's (1951) supposition that S, civitatus may be a third subspecies of Symphurus plagusia sensu lato, is defensible, He considered~- civitatus a true al eujsj_tvl~ Eu~uJ~m~8 ------·--.... - r:1 bn.uo1 r..urrurlom·uG 1:o zeJ:.in.qc: ~lno er!.:: P..1: .sa.ul;qf.!.£ .s_ ,:;; bn.s _....,...::r:ii.rlln.£cr___ .. ._2. rresw;:ted ( eirwoo ·, In·1de.:l".f!!V bnn wo·1 •,· P·1·_,_[', {""C.[ 0·1.1~ •.[;,n,rr -• - ·_;-,-,.,,,,,rr.r·n----'- • • -"\, 1'-'"'•ol·! r> ""o'l'~'{ < ,, "· to'-a~ud,,r·.r.·,··J) ·''· ______G.t :,JJ~c_,c;[q.., ---;;_lBU"-~.[cr_Z.,:___ ,;.L •--2.1rn1rlcmv8_ .. ,_,..,_ I sol'Iem,l f.s'Tdne::JI tr:s eeJ-bnJ ' I1~s~a m~71 bns ,rodol~~e uanee ,(~eblenrloB bno dooLB) 56 species, based primarily on the index of divergence of the dorsal and anal fin ray count, However, a detailed comparison.between S, civitatus and the two subspecies of Symphurus plagusia may reveal some functional and anatomical affinities. Symphurus plagiusa has already been accepted as a true speci_es, and this study is supportive of taxonomic distinctions between.S. plagiusa and~. civitatus, With the recent discovery of~. plagiusa off the coast of Quequen, Argentina by Lazzaro (1973), it is possible that~. civitatus may be a third subspecies of Symphurus plagusia sensu lato, If future studies show this t"o be true, as Ginsburg (1951) suggested they may, Symphurus plagiusa and Symphurus. plagusia sensu lato would have two very closely related distributional ranges. Specimens collected in such ranges would exhibit hor~zon tal and vertical distributional similarities. One species would be found in shallow water (S, plagiusa), while the other would be found in the offshore areas (the present Symphurus civitatus, off the eastern Atlantic and Gulf coasts of the United States; S:ymphurus plagusia plagusia sensu stricto, off the coasts of the West Indies and Central America; and Symphurus plagusia tessellata, off the coast of Brazil and Uruguay). -i:----- ·r-.. -·-- \fUJGL[C'3 ! ~JJq ;:}Alll.c>JJffLiF· f'J~=~rra.ft', l 1-wrrJq pe 1,ormq :p:r apl;l'JJOP, r~µ.ra r.u ps pL11s) ~2' c:p.1c:1p:1r.,'i (Jd~J) 2rrt.:·~sapcq DfJGl!. ~:81. · i::o<:2:rpJe r.ps:~ ;i· ~:riff-i!ifmi 1.iJt1i\. ps s rJJ:rJ..q 2n;i2becrG'.J V:[. 1 i:-ps C(J9'8J,:' O.f. {jn0Ji1s0i ';Ll~G.or.:pJg pll T'fJ:Z~ULO \j(::i)) :fP :f':'3 f.SXfJiJ(J!)J:fC q:ra,::p.iCf.'.fOJJ'cl per.1~GEiJ 05.l'/c,66_} 3• . ' .. ~n,;;,:::r~:?~ pc-i•..:-qq i.JL..fUh.:.!..~-Ji\. 01-J f.'}J-3 ruqGX O>:, q,[.b.GLi.~~lJGG Ot i . '. 57 Lazzaro (1973) reported that§. plagiusa has now been recorded from all five zoogeographical regions (figure~). However, Topp and Hoff (1972) have stated thats. plagiusa does not occur off the eastern coast of Mexico. Symphurus plagiusa seems to be depauperate at the extremes of its range (New York and Argentina). Population abundance reaches its peak in the Gulf of Mexico (Many authors, op. cit.). There were variations in S. plagiusa populations, but statistical analyses revealed that none of these variations were consistent or significant enough to warrant a division of the species into subspecies. The population from South America, especially in the area off Argentina, displayed some morphological divergence. This population warrants further comparison with popula tions ·from the United States and adjacent waters. Symphurus civitatus has two geographic populations ( Figure 5) , separated by the Florida peninsula. The Gulf population is probably the larger of the two, with specimens recorded from off the coasts of Louisiana, Mississippi, Alabama, and the western portion of the Florida panhandle. The largest collection of S. civitatus examined is catalogued in the Collection of Fishes of the University of South Alabama, but there may exist other larger repositories. I .,, • 58 Figure 4, Distributio~ of Symphurus plagiusa. 59 \ \ Figure 5. Distribution of Symphurus civitatus. 60 The Atlantic population (Figure 5) is rather small, occurring from North Carolina to Florida (Briggs, 1958), and only a limited number of specimens were available for personal examination. The populations showed very little intraspecific variation, although the Gulf population exhibited a lighter body color and more pronounced vertical bands than the Atlantic population. This was not a dependable characteristic, as many samples from the Atlantic also exhibited a light body color. ' 'f; CHAPTER V!I SUMMARY Specimens of Symphurus plagiusa and Symphurus civitatus were examined for 27 morphological and dis tributional characteristics. Many of these character istics were described by Ginsburg (1951) from a different group of specimens; however, additional observations produced helpful meristic characters that showed slight, but obvious, differences between the two species. Two of the additional characters that proved to be helpful were orbital diameter and vertebral count. The results show that fin ray counts, especially caudal ray counts, and scale row counts were of primary importance in identification. Another important parameter was depth of capture. Symphurus plagiusa, in addition to having slightly lower fin ray and vertebral counts than S, civitatus, is the only species of Symphurus in the western Atlantic found in shallow water (six fathoms or less). It is also found offshore, but less abundantly. The orbital diameters of S. plagiusa were slightly larger, and their scale row counts were slightly higher than those of S. civitatus. 61 QT -... , \:/qq:rc~·c.,!J !'O JH1AI!JE 2J:r~JJ!'J1 yo1.1s;, trlJ J,si\ suq 1>.6,,!;epr.-.;J ' bi:.oqrrcsq JJG Jbj:,HJ WGL:f2!':fC cpm,\l'Gj'GL:1 !'/JQ1: a;r:,,.,Gq 2:. '.[b'.)Jp. ,;:r,e,ni,, Ol 21GC]°U:G!J~ ! j.Hl/,\GAGL' sqqJ !':f0IJ8J op::,G.<,AS-;:'{UIJ:?. 62 With respect to the 27 characteristics considered, differences between the two species were greater than similarities. However, it was extremely difficult to separate the two species by sight identification. The inclusive ranges of both species_were divided into five zoogeographical regions, modified from those of Ginsburg (1951). These regions were (I) from Long Island, New York to Cape Canaveral, Florida; (II) from Cape Sable, Florida to Tampa, Florida; (III) from Apalachicola, Florida to Laguna Madre, Texas; (IV) the South American coast of Brazil, Uruguay, and Argentina; and (V) from south of Cape Canaveral, Florida to the Bahamas, the Antilles, and Central America. Populations of S. civitatus, collected in regions I, III, and V, were quite similar. Populations of s. plagiusa were also quite similar, although specimens from region IV exhibited some differences when compared with specimens of other regions. Statistical analyses were performed to compare interspecific and intraspecific variations. Independent 11 t 11 tests were performed on the interspecific var.iations, and showed several significant differences between the two species, Significant intraspecific variations were determined by one-way analyses of variance (ANOVA), and by Duncan's Multiple Range Tests (POST HOC MEAN), The J.,J.JG qer--sLmrusq pi\. oIJs~1v~~:i su~J .'ll,.ae2 o·t, 11.i/1.. :p:n~cb ( t:.-H.:fi ~ J • 1.1ug ' ' . ., 0 C.M.O 8bGCJGg• .~ 8T\lJ:r;.;ctt1.:_p J"iJ.C'LS2bac~:-tJG A{•ri~-J-:J~:rCJv~ !•H~l..6 ! • ' • • • •. • . • • ;rnq 8l]O/.\Gq 36/1.e,LllJ 2:[JU'.rt,:fCSiJi' .g:n,t,eI.G!Jt;G3 pG~!J.66,J ~!.J8 ' a~a (:'62.p·a f,\SL6 bs1.1_,r~LT.iliit.J' OJJ p;iG :ri.1poJ..ribt:C~"f,:fG AS:PJ9P:f0LIG"" '.[)Jr:"S~.fJD6C:f:f..fG suq .'.fl.J~L,i8D6C'.!"t,:fC. A!J.L:fSP:fOU8" r;rqe.(,SlJq,;;;~ t. . ' 2psp:rar.:ircsr 1.::-,-J8rlaea b\GLS hoi... t,o~u:.;-q ~~L' ccn,,;b81..r3 8'.[lll.f J SL' ' \:..} 01.::rq~ po rs~UlJt·f W'JqLG ~ J/3X£J2? ( Il\) .f.:'fJ8 -aorrt-•;1 Vl.l.i 13L:f Ct)li 63 results showed several significant differences between populations of§., plagiusa, while all results were negative for intraspecific significance in S, civitatus. It is concluded.that S, plagiusa .and§.. civitatus are two distinct species, as suggested by Ginsburg (1951). LITERATURE CITED LITERATURE CITED Bailey, R, M., J, E, Fitch, E. S, Herald,- E, A, Lachner, C. C, Lindsey, C. R, Robins, and W. B, Scott. 1970, A list of common and scientific names of fishes from the United States and Canada. Third ed. Amer. Fish. Soc. Spec, Publ. No, 6. 150p, B6hlke, J, E, 1961. Two new Bahaman soles of the genus S!mphurus (Family: Cynoglossidae), Notulae Naturae 3 4:1-4. B6hlke, J.E., and C, C, G. Chaplin, 1968. Fishes of the Bahamas and Adjacent Waters. Acad. Nat. Sci. Phila., Livingston Publ, Co,, Wynnewood, Pa, 77lp. Boschung, H. T. 1957, The Fishes of Mobile Bay and the Gulf Coast of Alabama. Ph.D. Dissertation, Univ. Alabama, 626p. Briggs, J. c. 1958, A list of Florida fishes and their distribution. Bull, Fla. State Mus, Biol. Sci, 2(8):233-318. Bullis, ·H. R., Jr., and J. R. Thompson. 1965, Collections by the exploratory vessels Oregon Silver Bay, Combat, and Pelican, made during i956- 1960 in the southwestern north Atlantic. U, s. Fish. and Wildl. Serv. Spec. Sci, Rep,--Fish. No. 510, 130p_.. Chabanaud, P. 1940, Contribution a la morphologie des Cynoglossidae (Teleostei, Pleuronectoidae, Lolei Forms). Mus, Nat. D'Histoire Naturelle Bull. 12(5/7) :182-191, 1947, Notules ichthyologiques (Suite). Mus. Nat. D'Histoire Naturelle Bull. 19(6):440-443, 1949, Revision des Cynoglossidae (S, Str.) de l'atlantique oriental. Mus. Nat. D'Histoire Naturelle Bull. 21(1):60-66. Christmas, J. Y., G. Gunter, and E. C, Whatley. 1960. Fishes taken in the menhaden fishery of Alabama, Mississippi, and eastern Louisiana. U. s. Fish. and Wildl. Serv, Spec. Sci, Publ.--Fish. No. 399, lOp, 65 :·,1:rJqJ • ;~si...a• ·2hec• ~;c:r• hnr;j •--1~·i·ap• ~:o• )t:d' J'Jbr WY22raaTbb:r,. s"JJq a'J8~GJ.. 1J rorr:n:?:r£Ju~· J~~ :1° -,Jau- suq f.. ~·2µ62 ~$j{SJJ :f!J ~JJG IUGUfJ~G6!J. ·t:f8j.h➔ J..:\ Ol,. ~-rsf;_.r:,/:.;) GjJLr2pwse) 'l. ~t,.• ~ r:· (}fHJ~6L~ s1.1q E. c· t,1J.IS~~re;1.• Jjcio• V,Sr,flLGJJG BHJJ" :SJ(J) :()Q-(:(?' J1si:-ru:1r.tdne 0L:ro11~-:-sr• wrre• ~10-p• D1H:f:-q?o:;-1,s ------J<:W"ij' gs.n.:pi.roD ,qra,;: c.1,uo~ro:a~:rq0s ( ::;· ~:,-'L •) qG ' ' ·- · r:s(i'l\.:il :ni:s-·ruJ'· ._., --·- - f;OJ:.IJJ2) • ,_1/JH-:l' !1~!' • D, H:f8i.•o:p,e L•Jtl/~I1L6j"JG BflJ_1 ' s1uuiJoaeJqas '(tsrsoar.s:r! hJani:.ousc~oJqes' rora:r CJJ9p 2JO" J;lu,,·. I!,T2}.l' SIJ 1:1 t-\rJqJ' 2GL11-' 2b&C~ ac:I' fjG.b·--t:r~f.:• yJtr• J0(2O :PJ /'}Jo 80fJi:')J/.l.68l'6J:,lJ UOL/!!J "\i/'JSlll''.fC' n· -~· 1 i •.)rTJfL;jLc,A ..--.91,f.) '"Ulllps0,.u i· ·uuq rJ o·rTCEI.Jl... 1!1308~,, ani.j·ut: .;._Y--'1.... :o-, GOJJ6Cp~·o1J2 p1. ,P}JS GXbJOL~fOLi.. i\6286] CJ {_'J.,f~0U i ;:JHJJ-rz' ii' JJ' • '}L'' suq 1• H' J,p(}u1:0ao11• Ji'l(:12' s(q} :s)?-3rs· q:r2r-1.:rpr:.:-~·0JJ• 1311J J • !!,rs· ~::-sr?s 1-:Iffu• 1rror• 2G:r 1' t)J. .. :r~n:a~ '1. c· J('i~s· V r:rac Ot. ~J01.. :rq~J :f..I2J.i~2 'dliq ~J;~;rL 1;'J5psrus • (?S(?b • cnJt cosa~ ot 1JJsps-rns· bJJ' lJ • iJF•2Gj.{'B~'.full' l!iJ:f 1,' !J02cpm;is:' H • j,' Ta 2~\ • ,tPG I!,:[2pS8 O:[. ~JCP:fJG BS!~ $1.TC.~ ~:,..JG 1 I bJJ:fJ9 •' I':f/,:jll6!ei·ou l,flPJ' co• f,1}.l1JJ6W.JC'{j b(J' .'.,,rb· pµs gs~smse 9JJq yq1SCGlll' ~B/'6LH' vceq• MD~· ~Gl' ,rnprr-;;' '1 • E'' suq c· c· o· cJJ$bJ:r11· JdQ8.' 1<,ra1,,,:i ')~. )1}11": J-f' ?,A11Jbpn.u1:a ( ~.\'llii:fJ1: c:li.11o~ro2:,qqss) • [10f.'f!J1;e .fll>l(-',-u,-;;s B9PJf-G • 'l' 1£' Jci!;JJ' 9M) !lElf.1 Bl'iJJ'>lliUu uoJ,)2 01, (-'J.TG r~;-,1H12 66 Compton, ij,, and E. Bradley. 1963, Survey of fishes found in gulf area 20 (off Port Aransas), from 0-15 fathoms, Tex. Game Fish Comm., Mar. Fish, Div. Proj, Reps., Proj. No. MF-R-4; Job No, 14:1-14, 1964, Survey of the fishes found in gulf area --~20-r~rom 2-17 fathoms and of post-larval fishes in Aransas channel. Tex, Pks. Wildl. Dept., Coastal Fish, Proj, Reps., Proj, No. MF-R-5, Job No, 12: 1-24, Davis, D, D., and U. R, Gore. 1947, Clearing and stai~ing skeletons of small vertebrates. Fieldiana. Technique No, 4:3-16. De Groot, S. T. 1971, On the interrelationships between morphology of the alimentary tract, food and feeding and behavior in flatfishes (Pisces: Pleuronecti formes). Nethl. Jour. Sea Res., (2):121-196. Gallaway, B. J., J, C, Parker, and D.R. Moore. 1972, Key to the estuarine and marine fishes of Texas. Texas A&M Mar. Sci, Sta., Galveston, Texas. 177P, Ginsburg, I. 1951, Western Atlantic tonguefishes with description of six new species, Zoologica 36(3): 185-201. Gunter, G, 1945, Studies of marine fishes of Texas. Publ. Inst, Mar. Sci., Univ, Tex. 1(1):1-190, Hildebrand, S, F., and L. E. Cable. 1930, Development and life history of fourteen teleostean fishes at Beaufort! North Carolina, Bull. U.S. Bur. Fish. 46: 383-4!:18. . Hoese, H. D. 1958, A partially annotated checklist of the marine fishes of Texas. Publ, Inst. Mar. Sci. , Univ. Tex. 5:312-352, 1959, A checklist of fishes of area M-3 --~(~L-ow-er Galveston, East, and West Bays, Texas). Tex. Game Fish Comm., Mar. Fish. Div. Proj, Reps., Proj, No. M-3-R-l; Job No, A-3:1-5, Hollister, G, 1934, Clearing and dyeing fish for bone study. Zoolog:ica 12 ( 10) : 89-101. acnq1. · soorotses JS (JO): ~,:l-JOJ" l{OJJ:r8rG:.' e· Ji1)f" CJ63l,pJt~ 2uq (J"l'.S~!Jo'. :c-1·2µ 1,0L 00tlG Via· r,:-)-l:i-r: 'lop y,o• v-):r-2· c,Gree li..:r aµ corull!• ~ 1·JBL • T!i:r,1µ • n:r L\. • bLD~1. g0bn • ~ J-,.t..o1 • __ _(ru.:::s.r.. cs-JAs2r-ou~ ;_isi2~~ soq !:_.Gar. na:'.2' j}~X:~'d.J"' LG:-;• JC;2:j • v cpecf<:Jr~r- oi t:ra1,:ee ot 0Le~ ~~-~ nufA' LGX' ~:)J5-)~5• ::"JJG IJJSL:fUS l..,=f2fJ6'2 Ol, J,6:.,~g?. • bflPJ' rua~• ~~L· 2ci•' H'J6'J8~ H. if' J\~~t~· V b8LJ:':f~rrJ.. suuo~o~sq cµac~Jr2c OL bJJPJ' Jllcl/.'" J-JSL' ?,C:f'' ,[ltJ:fl\.' .LGX' J(J) :J-J60' {]nliCOLa Q' Jdt~• 3cnqtsa Ct WBL~'UG t~-~~62 Ot ~GXS8' r~~-sor· . . qsacLrbpron ot eix ua~ abscrsa· sooro~rcs )~~J}: c~·uapnx.12 • r · JO ~r· ;,1ser.eLJJ v~Jsor:rc ~o:.;t~rr!ji,:f~ipe8 t!~.f ::JJ ,T,GX\cl2 \i~Y'i r,_;~iL' c3c:r • 8/.'ff"' Q$JAG2/.'0IJ' .L6XS2' Jc\,1i)" J(GA co cps f;ar-m,L:fiJG 1.mq wsr..:r11e 1:,r;ij10;:i oi:, 1e:rn:•:• (}HJJS-li\U/..' B • '1' l "i • c • !)SLTt'3L ~ suq D. u-· t•.;vot.,a • 1·1: ...\:~: • tOLWSH)" MG/.'PJ' ~onL' ass gsa·' (s):rsr-J~Q" rrnq pGp3 .f.:f GI. :f!J T,JS /.' LJ 2p6 2 ( i:.:f 2CG?.: bJSf!l,G!.16 Cf-:[- !lJO.L!'.:,JJOJ Of;:A Ot ~µa 'aJ:fll!SU;:'tiL)i. ;:'LUC{:'~ t,ocq ~!}tj 1,GGqr,1;1 us c:i:..copl 2· 1L' Jc).~J· ou CJJS :p1r.sLL0J:J~~:rou2prb·~ p6-t4\1:;G;.; tscpu:rdns 10· ,:)-r~· ar.~:piru~ 2f{8JG(:Gl.I9 Ot, ~Jill~JT A-SLrG~L$,Pf:a• l!,:reJq:p;:;s · f1~1'J\.J8i D' I)'~ SJJQ n• r5• (;Ol.8 1 J1:nt~• 8J6S.i:..~·1H?. ~n.Jq J-sr· J1,:f2P' 6Lo'➔• H8b2·) .r-,Lo1• wo• Wl!,"'·H-~'- '}OP j/_0' 1 J:5·: vLeuas2 c1-H1uJJer• J/;;x• bt'a· ll~~·rqr• u0.bc• r;os2.f:-:.:iJ _____ S0_ _1L'JIU ~-T.~ l/iiCY~lTJB ~rJq Ol., bo~-~r.-JrrI.. J:..£rJ t:f.~jJS:J :p.J J dPf ~ 2nJ.... \V t~ 0 [, j:' PG 1.:r 2po ~ :(, (;HJJq ~ :J ~.,~·rt 91..G 9 f,LO ·~ • :;e bG • ~ bI,.o -~ • 1.1 o • t·~L-iJ-f ~ '"'1 up 1:1 u • J\t: J-Jr" c t~~pum~3• ,I,6;-;:• JS'WG ii,:f2F CC:ilh. • • ~-_vt.• B,.f2}J' U.fi.. ' 1 [,uDDq :fL' ~f!Jt SI,Gcl :50 (O~.[, ):-,0>,jc VJ.,8l.J:"\:':,) l),(;UJ f)-J) cou:l)f.'O!J~ !~ ·' s.uq r.;t ·~3.r.~~;qJ ...Jli.· ]l7~?)• ~nI.-./\.. sA. t1.i 11·3p-5;~ 67 Hubbs, C. L. 1922. Variations in the number of vertebrae and other meristic characters of fishes correlated with temperature of water during development. Amer. Nat. 56(645):360-372. Jordan, D. S,, and C.H. Gilbert. 1883, Notes on fishes observed about Pensacola, Florida and Galveston, Texas, with description of new species. Proc. U. S, Nat. Mus. 5(1882):241-307, Jordan, D. s., and B, W. Evermann. 1898, (Reprint) The Fishes of North and Middle America. U, S. Nat, Mus., Wash. D.C. T.F.H. Publ. Inc, LTD., Jersey City, New Jersey, Vol. 3:2704-2712. Lazzaro, G, E. 1973, Presencia de S!mphurus plagiusa (Linne, 1776) Jordan and Goss, 18 9 en aguas de la plataforma Argentina (Pleuronectiformes, Cynoglossidae), PHYSIS. Secc. A, Buenos Aires, 32, 85:245-'249, Menzel, R. W. 1956. Checklist of the ·marine fauna and flora of the St, Georges Sound area. Oceangr. Inst., Fla. St. Univ., Contr, No, 61:1-99. Miles, R. M. 1951. An Analysis of the "Trash Fish" of Shrimp Trawlers Operating in Apalachicola Bay and the Adjacent Gulf of Mexico, M.S, Thesis, Fla. St. Univ. 46p. Parker, J, C, 1965. An annotated checklist of the fishes of the Galveston Bay system, Texas, Publ, Inst, Mar. Sci., Univ, Tex, 10:20~-220. Robins, c. R., and J, E, Randall, 1965, Symphurus arawak, a new cynoglossid fish from the Caribbean Sea, with notes on Symphurus rhnisma and Symphurus ommaspilus, Bull. Mar, Sci, 15 2):331-337, Springer, S,, and H, R. Bullis,, Jr. 1956, Collections by the Oregon in the Gulf of Mexico, U, S, Fish and Wildl, Serv, Spec, Sci, Rep.--Fish, No, 196, 134P, Springer, V. c., and K. D, Woodburn. 1960. An ecological study of the fishes of the Tampa Bay area. Fla. St, B~r~-- Cons. Mar. Lab., Prof, Paper Serv. No. 1. 104p, Stickney, R.R. 1976, Food habits of Georgia estuarine fishes II, S}mphurus plagiusa (Pleuronectiformes: Cynoglossidae. Trans. Amer, Fish. Soc, 105(2):202- 207. SO.')' cAuoO:Jo~2:rqs;U_·_ _r,_~a.;_~~!--_._1•,:p-1p • ~1-.Jc • JO~ ( s): su5- i-r 2pea II• ~li.Il!DfJJlI.. ilG .oy~rrt~·n2s ( bT~HlLor1Gc~:r~v1.,ll.\Ga: 8..C:fCJUJGI.' H" iS" T:)~{2° !'_,00'..J JJSP{P2 01;, ~JGC.L~:f~ uac-n~:.i:_.:pJG J(Jf.O' i-l:fJqJ• ael.,A 11 31H3G" 2c:r• H9b"•-ri..:raµ· ~•iC" TC1{2' J)?t.·· pa r.}JG QLG°t(OIJ :p.r !'JJ6 Qf!J1. Ut l'-)6X:fC(,' [\' ;:; ' ;•. ~·ap 'JDq 8bL:;11~sr.' ?.' I SIJq H' if' BHJJ:ra' '!J;.' r0212· SO:rJ6Cf-'lGJJa J_uar.• :WSL'. ;_ic:r•' UIIl/\.' ·,[,6X·• JO!S(l'.f-5:SO' {.:f8JJ68 Ot J:')JG CUJAG8!'0U BSA 21,a.i:-eru• .f,0X9'2' . bllj)J' E,SJ.f{GL 1 'l' G' _J f!l.Lf JI.' V~b • ""6 vcr'JSGEl!Jf' QT 111••1• \.T,J' 1· t' V • • ,. onri·t. "'t> l•lGXICO'• II. <,, .:.J,f\JG,n,;' • ,__._ c,1:•"'' 8JJLil.Ub ,J,S..ffMJGLt?. QbGLS!':flJti :rl) VD\lJHC!J:fCOJS B::1), 'J!Jq w:rrea' H' ~r Jd~J· .'iflJ \;lJUJ1a:r_a Of, PPG ;i.L1,\J:,p j,,:r,,µ,. Ot :;:.it• 1~nri· ~(n,ssJ::sr-r-)o:-.· .LG:..-ce-2• ,·,t:r~P qs2c1.;·IJ~'.fOiJ ot IJS14 21,v..;clsn· ;:,1.,c,c· ::· ?i. op2sLILsq uporrp I,Gt.1 ascors· ~ ~JOL.Iq~:i ·~ uq c~J t£, 2~0JJ ! 1 11c.x.q~.r11~ D. a· ~uq c· H· u:rrp-s.Lt' :r~::tt3 · 1,··Jr~•s2 :. .. HJ 1,;,:~~ea qs~GJObweur.• ~WSL'_ MBf' 2Q(?f~):)QO-}lS' _ GOl,}.. GJt!pBq lt\:[~µ ~Gc.r;JS.L~~!lI.. 6 Of.. ;.~ucs!.... q:-1::.:.;~J~ i\CI.. psp;.. ~:s ~ug OJ:'f.l?L u;GL'[8P.fC CiJULVG:~G!..,-;:: Gt t.-r2p02 f-!flJ)_V{) C' r· Jd~:S" J\9Libr.~DJ.ra ru ;:po ?.JnUpG;., {.i¼_ 68 Topp, R. W., and F. H. Hoff, Jr, 1972. Flatfishes (Pleuronectiformes). Vol. IV, part II, Mar. Res. Lab., Fla. Dept. Nat. Res. 135p. Walls, J. G. 1975, Fishes of the Northern Gulf of Mexico. T.F.H. Publ. Inc., LTD. Neptune City, New Jersey. 4J2p. APPENDIX 70 Appendix Table 17. Independent 11 t 11 Test of Dorsal Fin Ray Variation between Symphurus plagiusa and Symphurus civitatus, Parametersl Species N X s.n. D.F. llt II ~- plagiusa 168 87,52 1.93 s. civitatus 198 89,58 2.02 364 9,88 lWhere Na number in the sample, X = sample mean, S.D = standard deviation, D,F. • degrees of free dom y~fl :--::J~: IG2·10CT ~·.o Jc.~{i 1rj 11 J'rre.brt!;'qsbn:I .\_( eidc_;'J ______~-~u t:·•s.Ic: _. ----~2JJ'!.udamy<~ fl£H:i 1,fjsd. 110.t.:"~ t7sV , -----"----"'"'~?J.JjBd .i:v ..t~ ~u~urtqin\:~~ - .bns --· ,a.e--- x Vi (0.L s;;;. \'8 ~1d.[ so.s s;: ,Nl 8QI ----___ __. r , ,,t.';9iil sJcpicc = X , slqmsa edJ n.i: ·rndmurr c ii s·rnr!l·.'-'- -es--r't .-i.Of;jS'ICSb n .. 'iI.l1 1 rto£jsJ:veb brr.Gbt!,-;jz = (1,2 mob r i' ' 71 Appendix (Continued) Table 18. Independent "t" Test of Anal Fin Ray Variation between S¥hurus plagiusa and Symp urus civitatus · Parametersl Species N X S.D. D.F. "t" s. plagiusa 168 71.55 s. civitatus 194 73.94 1.71 360 12.52 lWhere N = number in the sample, X = sample mean, S,D. = standard deviation, D.F. = degrees of freedom. .Lnni\ 'lo ;Jao'i' ";J" ;JrrebnecrsbrrI .81 -,I.daT S/J"!UJiamyc, !J99W.!'Jd !'l'liv'[i.t-:rsV ~U"1'.urfa7ii"vc bns sa u t:1 s1 a ~,J.sj.J:v l::, - -=----:..:-====-= x . 8cll X , slri0aee erls fll "!edrr:urr :: !,'I (3'ter!Wl = .1.ll ,noljslvob b~~bnsje = .CT.2 Appendix (continued) Table 19, Independent "t" Test of Caudal Fin Ray Variation between Symphurus plagiusa and Symphurus civitatus Parametersl Species N X S,D, D.F. "t" S, plagiusa.. 171 ,59 §, civitatus 206 11,85 ,56 374 33,60 lWhere N = number in the sample, X = sample mean, S,D. = standard deviation, D.F. = degrees of free dom qom ?."G" ,: Ot,HJCj~J.q qGA:f9/::fOU' D'&,· ::; qs'(?.i:.es;J GI. T,;,ss J!~\PGLG 11 = UfH!.1p6L :pJ f:'PO 8S!i'!:'tJG: X = asl!.~)JG i.l:d'..:U~ V i• guq 2}:Uitµ~ c:r Al. c~:.:f..-" 1\. G;.. .f S' t ,r OU p i.:i ~ t-\$ G L' 2~\.rnbj:fln.."rrD ~D"":y-ti.,...:_.,.._-:~,.,11-~2--s r:..:qsLG!JqGU~ 11 tu LC8~ Ct risnqf;T f:,.pJ g~1l 73 Appendix (Continued) Table 20. Independent "t" Test of Scale Row Count Variation between Symphurus plagiusa and Symphurus civitatus Parametersl Species N X D.F. "t" s. plagiusa 131 79,46 3.12 S. ci vitatus 171 75,98 2,57 301 10.60 lWhere N = number in the sample, X • sample mean, S.D. = standard deviation, D.F. • degrees of freedom • •••-•-N---•••-••-••• - • •-•--•·~-••-••· ••• ------••• --•- -•-••-••--•------•• . -·· ... ------·------·-·--·~-•------ ., A ICI -., ,..I••\ r , 3 :q1.;c:.,. f-Hl:; rrJ '1"-:>ciJ-:.!HG :-2 ~·j 9'15il, .... .'·~.([ ,;viJ:1crvst h·:r0b:•eje o:: .G .2 Appendix (Continued) Table 21. Independent 11 t" Test of Vertebral Count Variation between Symphurus plagiusa and Symphurus civitatus Parameters1 Species N X S.D. D.F. llt II _§, plagiusa 129 47,12 ,92 _§. civitatus 107 48,69 ,67 184 13,31 1Where N = number in the sample, X = sample mean, S.D. = standard deviation, D.F. • degrees of freedom .-1-1:· :: ~i~_•J~J(!'J.'.._.c; J$[:..i:.i:~·1•:.lJ· ;J•~.· $ q~.;,_;.. :~2 -·.:... :,,;.. ..;;·'.,.,-; ·:-:',.J-IGI.. G ::1 • ;:il:npG..:, .p.J eµc ~1s:.:1bJ t_; ~ f\. ~ '.: ••• ...,J'S :; S'.;.; · ---,----JjJ~i ~ f !1U-d ,., ,..,, . 75 Appendix (continued) Table 22, Independent "t II Test of Dorsa--1 Fin Ray Variation between Symphurus plagiusa and Symphurus civitatus in Region III Parameters1 Species N X S.D. D.F. .§. plagiusa 79 88,17 1.91 .§, ci vitatus 179 89,57 2.09 256 5,06 1Where N = number in the sample, X = sample mean, S.D. = standard deviation, D,F. = degrees of freedom ______,. ___ _ ·1• G:...'\~l•-.,;f:JJ2 tJaL~.:~1;G '.' 3 t,2.- " 76 Appendix (Continued) Table 23. Independent 11t" Test of Anal Fin Ray Variation between Symphurus plagiusa and Symphurus civitatus in Region III Parametersl Species N X S.D. D.F. "t" S. plagiusa 76 71.77 1.94 .§.. civitatus 180 74.02 1.70 254 9.22 1Where N • number in the sample, X • sample mean, S.D. = standard deviation, D.F. = degrees of freedom _[ ,';.,nj9m.....L~ •I [l'T~(I.) _ tJ ;. Vi ·------·-•·... ------·----.. ------· -·-- 081 -·------..------·------·------· ------.... ---·------•-·------·---·------··•------·--- 1 t rE:s;;r sI½mH2 ~ X t efqrr..s2 9£i.J ul '!edmun ::: Vi s~srn·1· ri•r,bf.S"I°l '"'re c:r:9•1?.eb = .'ii.CI ,nojj.Q]'vsb b"!nhr.r..1e = .G.2.. 77 Appendix (Continued) Table 24. Independent 11 t 11 Test of Scale Row Count Variation between Symphurus plagiusa and Symphurus civitatus in Region III Parametersl Species N x S.D. D.F. .§., plagiusa 55 79.34 J.38 .2• civitatus 90 75.92 2.67 219 1Whe-re N··= number in the sample, X = sample mean, S.D. = standard deviation, D.F. = degreESof freedom 11 ~.i't1!J00 1:1051 sls~G 1o .:,::;sT j" Cf1.f1hn.9qobrr.T • 4\~ s[d,~'1 s,• .:}2_:::.fg_ :?.!rwrlgmy8 irnsw.:i0d norjsJ·u,V rr.l: ;:,, u ;/ s j .i:v .i: :i _i!,Y'Il.i rlgf!.1.Y.\:. bn G :{IT-aol:,:;sH ---•---~--•--•-•--•----•- - --•-• •--- • - - - • --•-••-•-- • •-• -•---••••-•--~--•••-•••••• ·---~------·- --•------•_..,._h ______,o•-•------·------11 ------., ______------cc ------8/JJG,:!.rv .i:!l , ,;;;s,n s.U;msa = X , sf.qrr:sa s!l;J rd -:r~dmm1 =··VI s'Terr:;.C ~~ 1:,.C[.s~:-1 'le- ~S"'1~Gb : • ',j .Cl ,noljGlvsb b,:sbnsje = .G .e 78 Appendix (Continued) Table 25, Independent "t" Test of Vertebral Count Variation between Symphurus plagiusa and Symphurus civitatus in Region III Parameter~l Species N X S,D. D.F. .§. plagiusa 89 47,49 .82 S. civitatus 94 48,71 .69 181 10,79 lWhere N = number in the sample, X = sample mean, S.D. = standard deviation, D.F. = degrees of freedom 2·0•,. □ r.suqsLq qGA:rffr.{011• o·;,.· = qsc:.i;.(w,-::>:t:, ;1.;;·1:1:i.:i Jf,\}JF;~G t·i = urrmp::-;r.. .:fJJ• ~f.!8 '.39"L!bJ8 ~ 1: :: -;.?Si.:1!_,Jt .1,;_Gf.fil ~ ------·----·------·•-.------··------... ----·------ lil" J'-)J ·------•------·-· -·------!4 --·------~------.... ------~---- gsB::rou III 9'IJC/ ?,Al!lDfJiJL/Jcl cfmgfini :ru 1\$L:f U(:' .f Ol/· po p;,.GGJJ 8Jl.wl:iprr.r..m ofs-q·fl~;-;j' ~spJG s~ · 1uq,,bsuqE;i..r1: .. .:- 11 .1s:,r. ot t,e1,r.opi:.1J]. ::;omJp 79 Appendix (Continued) Table 26. One Way Analysis of Variance to Determine the Variation in Dorsal Fin Ray Counts of Symphurus plagiusa, Segregated by Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between $1.$1 4 20,45 6,57 Means Within Means 544,63 175 3.11 ..... _..,_ _... ______.....,_~---- -· -·----.---,-,.•-···•· ------·------.. --·--·---·-·------· .. ·- __ .... ______·•------·--·-- ____ ..,, .. _...... ,..., ..,._...._ ____ - [•:s ~n.1a ;.~.J';:'JJ.fL' .1\0I...J~Jp~·ou BdJ1$"JI.e a f,,UlGqr.aJJ ~:dns;.. G ~JOr:iLGG Ol, 8fl!;I 0:(. J)G t'~:r.c? 6 8 Gt !::•~:; ':J,.; •-•• ... -----• --•-••--••--••• ·•••-•--•••-----•----••--•---••----•••---•-A-•••-~•------•-•--• •• _...... ------.,. _____ ..,. __ ------·---·----"- .. -----~ ··- \,OOt'.G OG'.L\I bp:f C ITJ w,\1~· O;J or...... u.,AmEURi:"11\1 • 11-r~'ErTi~~~:;;l• ~:•,3GLt30.Sf-'GC(._,, - ol·). r..tJO 1:..-a.c.:rr.,·_p.fOJJ :p.; iJOL'=1~JJ }4J-fJJ rs£!:\ (~OJ';JJ('t: l)JJG f.-;_'iJ!'i.. y.1,,i::JJA'8Td 01, 1\iJJ,.,"[SlJCG ~C.: J_;S:,PE1.4l"L'fi,,,O 80 Appendix (Continued) Table 27, Duncan's Multiple Range Test to Determine the Regional Variation in Dorsal Fin Ray Counts of Symphurus plagiusa (n=9,68l k=5, MS :3,11 error Region Number II V I III IV Mean 86.62 87,25 87,46 88,17 89,00 Least Significant 1.58 1.67 1.72 1.75 Range Regions and V-II:,62 I-II=.83 III-II=l,54 IV-II:2,37* Differences I-V :.21 III- V:,92 IV-V =1.75* III- I:,71 IV-I :1,53 IV-III:.82 *Indicates significant regional variation .\~~- 1~l,i31..6 IJ Ce 8 .~.aftr(:r..12 ~uq ;J~)TJ~G 2 ~-~~iJ :f1:, :f GS :.i P I •:,c.. q ;·'p<;j 8 ~ ''. GGl.f {, fJ!J.:J);J!-., J.I l Ii\ l(;ib:TCLI II A I -:;-_-::..:-_--·:::: ::.::.=..::::.== =·--::::-::::-=:=:::===== VJ:.LOL ,·.'J :::::J·Jyl i .... _::i' J-,J~~--;-:;:n-~,. ... Cc,.~ d ( u-.... D:- • ('("\,;:..._;_..1-;~_",.:_- __ :~r.u 69h cor.rur-c~ O"f. ~;id.!·LpHL!J2 cps 1rs~1cunJ A~L~ucrc.u ru ooL~ar \ .:,._i... . ))f.fJjcsu,3 ;•:rrJr-:rbJ6 g~:Ji~G 1LGL!r. ~~· csr.t-,1,: .. 0 :·.J,.:;; 81 Appendix (Continued) Table 28. One Way Analysis of Variance to Determine the Variation in Anal Fin Ray Counts of Symphurus_plagiusa, Segregated by Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between 4 19.21 Means Within Means 583,70 171 elt er-:J·11"1edeG· oj 0:)fiGl:'16V 'Io e.te~l.ru1A 1sh' e~O .SS o.[OG 1l 1o ajrru08 ~62 nl1 Inn~ nl no.tisi~sV "ld b0jr:.2e~~s2. ,_geul:r.s.[q ~.l!r-rJJrfqm't{E: flo.i:~sfl Iso1 dqwr;gos;s,o~ · nseM 'lo aes'!;gsCl 'lo mr/2. 1o so-YjJ08 9 ..!.13JJp8 mobss'!"l 89'!.Bllp8 rroij.sl'lnV f!99Wjefi ertse~4 nllf j.i:':! 82 Appendix (Continued) Table 29, Duncan's Multiple Range Test to Determine the Regional Variation in Anal Fin Ray Counts of Symphurus pla~iusa (n:9.66, k=5, MS :J,4) error Region Number II III I V IV Mean 70.78 71.77 71,81 72.75 76.66 Least Significant 1,66 1.75 1.81 1.85 Regions and III-II:.99 I-II:1.03 V-II:1.96* l'.V-II:3.88* Differences I-III=.04 V-III=.97 IV-III:2.89* V-I: ,93 IV-I :2,85* IV-V :l.96* * Indicates significant regional variation snl.'i!-:rsjeli oj jae'I' e21rrnH eiq.t.:JiIJ:il a I nG:irr.uCT • (?S sids'l' rr.i:1 .[.srrA rr.i: rro J:jn.i:·rnV lsrro.i:;g:,.H ert.:l sa1;.i:;gBia ~rlcrrnv2- 'lo 8ilauo8 ,u;H ----Cfjl.~:: 8M. ,<:=,i ,oo,'?=n) '10'1'I9 ======--==-===--==-·==· =--~--= nolgsH VI V I III II -rsd:rm:~ LS.I';' jzr;eJ IS.I j rIEi ~ .i:11n;g.f2. 1?1?. :II-III bn.e r,no.i:;q.tiB e 3:Jll9'!91'l:.i:(i 1?,=II!-V ~O.=III-I CC, •• 1"-V \...... - .... Appendix (b ont inued) Table 30, One Way Analysis of Variance to Determine the Variation in. Scale Row·counts of s mphurus plagiusa, Segregated. 6y Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between Means 99.18 4 24,79 2.60 Within 136 Means 1295,76 ( be1.Jrrr;Jrro0') x.rimsqqt, sn.:' ,;')e~·sU oj eonsl"IsV 1o a.i:ev;IcnA "{SW er,O .0( eJc.iB'T 2jnuo8 woSI eisoC~ rr..t ao.i:~.e.L'raV ed0 bsdCi38".f:g,sC t seJ.Ji~~CJlcr._ clJ ..1urfgfil\"?, 1o nolgefl IsolrlqS"I!OBSOOS 12: nse:'ll 1o er➔ e'!:geCJ ':.o (iHJP. 1o eo·Iuo8 s·rnr.rpc.: . mobS8'l7J ZS'iCJJf-2 ric J;Js.i:·1ii;V !'!OS-Wjsfi 8f!f'.G[i. o(I rr.i:rfjJ:\~ an c~ el•1i 84 Appendix (Continued) Table 31, Duncan's Multiple Range Test to Determine the Regional Variation in Scale Row Counts of Sf:lphurus plagiusa (n=9.47k•5, MS :9.52) error Region Number IV II III I V Mean 78,66 78,93 79,34 80,06 82.50 Least Significant 2,81 2,96 3,06 3,14 Range Regions and II-IV=,26 III-IV=,67 I-IV=l,39 V-IV=3,83* Differences III-II=,41 I-II:1,32 V-II=3,56* I-III:,71 V-III=3.15* V-I:2.43 * Indicates significant regional variation I-III=' ,\I or1.lsLsucs2 rrr-rr=·tr I-II=J')S {iet~·oo-.: suq II-II\="S\? IJI-Il\='l\\ r-:r /\"'I· '.3,) !,OIJt'.G 3~·fiu :fT.. rc~u r. S'SJ S • c)I_) j '0(~ )'JI" 7:10~2r 01f'o3· io' 1~· 80• or2 ss·~o 1;1fllUp8L ·_; I'lt I:j: III I A HS&:f VlJ ---·· . - - . . GLLOL ·. ' I'/;;:_- · .=a·~s) JJJS6:f!Hl0 (o=a·tr'_J:=~' isor~ corrJJic2 ot 2.luiilipnr.rre pµG gsErougJ [\SI..:pl~:fC/JJ :rll 'i)CSJG .r,spJe )J' n1rnc<:ru I e ~;rrr~:rhJG H1:lll6'.G J,02::- i:-o usr.eLn1:ri:s 85 Appendix (Continued) Table 32, One Way Analysis of Variance to Determine the Variation in Vertebral Counts of Sym~hurus pla~iusa,Segregated by oogeograp ical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between Means 4 1.10 1.64 Within Means 90.20 134 .67 (beunljnoO) x.fbnsqqA 0rrJrn·-:r0:-1sC: oj 0::ia'[;.i:~nv 1G cle~IsnA 1~n\'I e.aO .S( eidt-'I ej11IJo!J .Iri. ..Jde0'reV rr.i: nol.:tBl"!.GV e.d~ bej~i?,f)"'faee, ssu r:uc!s._ cu--r1Jr!qmyC ·10 flOlicJe.1! In:.-i:frlqr,•rgoe;goo& :=v:a· ------·-·- •·•··-~--1, - ----·. rrr; s:.. 1 ':to assi:v,eCT 1o mu2 'lo eo--.rvo2 nrrr-;;.upE rn?_bse'1'.J ,, 89'1.GlJp8 no.rd F.\.f'1BV tr99WJ efl OJ,.[ eP.r;et-'. nlrl.:l.h/ OS,0(? £n£eiii ..----. ____ .. ___-~------·· ...------·----·--- •·•-·-···--- •·------·-- ·--··--·------86 Appendix (Continued) Table 33, One Way Analysis of Variance to Determine the Variation in Dorsal Fin Ray Counts of s mphurus civitatus, Segregated 6y Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between ,36 Means 3,13 2 1.56 Within Means 824,53 189 err_b~•"TeSoG od eons.i:•rnV 1o 1s.i:e"[IsnA '{SW enO • (( eids'r 'lefl nJ:'B. I.se-'roG fl i: 110 J:jnl 'ISV er!.:! • eJ:Jctf:l,llv .i:o c.u·wr!omve 1o e.:/rr.uoQ . . 1:s::d:rlq £."I~O s;gooS ';dbe.:/Gio\6~!~9~ nv .r~ ef.l ttn9M 1o P.09'!;<1,Stl 1o r.w8 'lo e:i-:r.uc:8 e~snpG mob!le'!'•l ao·rsup<: no.i:Jsl'IsV !'!99Wj9~} s 8f'1681~-1 n .i: r! ;J.i: \•,! en~eM ,. __ ._..,______------·------·------·· ______-- ... 87 Appendix (Continued) Table 34. One way Analysis of Variance to Determine the Variation in Anal Fin Ray Counts of s mphurus civitatus, Segregated 6y Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between Means 12.66 2 6.33 2.17 Within 190 2.92 Means 555.91 'i'S sn.tm·Js~isG oj s::,rrsl-rsV 1o e.l2i(lsrrA. ,:m·1 enO d\( s.[d!11: zj.nuo::i "iS5l n.t1i Jsn~ fll noljnl,:sV srlj bs;Jsas•r:gsf:; ,eujaj.i:if.i:O '_?IJ'!UrlcrirrJ!:. 16 no l30H 1s ::,lrlqwr;2os;gooS ,~ n:.s. s:.,l 1o eee·I3sQ 'lo muG ':o 9o"f.uo8 s-rsup2 me, b S 9'!"1 e.e"rG!Jp2 no l;Jf3J- ..rHV ------·---·------~------•·•·------~---- neswSsCT s B fl £. 9 ! ni!ijJ:H 2fii.10'.'vl 88 Appendix (Continued) Table 35. One Way Analysis of Variation to ,Determine the Variation in Scale Row Counts of SSmphurus civitatus, Segregated y Zoogeographical Region Source of Sum of Degrees of Mean Variation Squares Freedom Square F Between Means 21.51 2 10.75 1.53 Within 172 Means 1209.48 KBSU2 :t,:r rp:f!J ~'JG ~ifJ'a 3er.MG61J s ------·------8 dmn:.<3 a I!,LGGqOlll 3dm,i:.s 8flrn Ot D6&'.L6G"2 Ot l•iG\$!J . . pt, ,-oo•~eoE:LsbµTc~ 06\iJ"OIJ Ot 2XiiiopfJW2 cfii:ffs'fmi • 'i]G&J~G!r.S[.'Gg ,~jJG 11.s1,:rsp:rou :ru 2cE1JG. 1,01,1 corrui:e J,S'pJG" ) 2.. QIJ6 ;',\fl/I. 1flJSJAa-r2- Ot -.f\:>I,tElf-'.fC~ ~o nsr.~•r.0_;-GG Appendix (Continued) Table 36, One Way Analysis of Variance to Determine the Variation in Vertebral Counts of Swhurus'civitatus, Segregated y Zoogeographical Region Source of Sum of Degrees .of Mean Variation Squares Freedom Square F Between Means .33 2 .16 ,36 Within Means 102 ItJSt/U8 JOS M:fCP:fll 't\l(HHJ2 s !36.PlJ~SGtJ AfiJ,:fS/:':fCU 8dfi8LG2 £!,Z.GGqOJU ,;dm;i;,s 80flLG6 Ot ?,fllJJ qi nst'.1.sea ot p-,JG pl ;;:ov\'i'.eo\;'.1.\:fb)J:fCSJ l{G~:fOlJ Ot 8AJEDJJITLIT3,C:fArC$j:'fla 1 ?,6~L~g8CGq CJ.JG 1\_'JJ.• :f'Jl-':f,O!J .:pJ J1.GLJ:!0pLSJ <.;0111.lj:''