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Species Diversity and Phylogenetic Systematics of American Knifefishes (Gymnotiformes, Teleostei)

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Species Diversity and Phylogenetic Systematics of American Knifefishes (Gymnotiformes, Teleostei) SPECIES DIVERSITY AND PHYLOGENETIC SYSTEMATICS OF AMERICAN KNIFEFISHES (GYMNOTIFORMES, TELEOSTEI) BY JAMES S. ALBERT MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 190 Ann Arbor, November 01,2001 ISSN 8076-8405 Species Diversity and Phylogenetic Systematics of American Knifefishes (Gymnotiformes, Teleostei) James S. Albert* Division of Ichthyology Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109-1079, USA "Florida Museum of Natural History, Department of Ichthyology, University of Florida, Gainesville, Florida 3261 1-7800, U.S.A. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 190 Ann Arbor, November 0 1,2001 ABSTRACT Al/~(.r%,,J. S. 2000. Sj~uc.ir:\cliv~rt ily cind pltyloger~elicsyslmrnric:., (fAmc.ric.on knif~/ish~~,s( ($tr~nolz/i,r.tnc~, 'lP1uoski) . Misc. I'ubl. Mu.\. Zool. Ilttin Michiyccrr, 190:l-127, 50,/ig~. Evolutionary relationships among iZlriericaii linifefislics are doc~uner~tedfi-0111 comparative studies of morphological, physiological, and behavioral features. Materials were examined from Inore than 900 lnuselirn lo~scoiltainiiig over 4200 gj~ninotifoi-in specin~ensreferred to 105 described and 32 undcscribed species. Neural tissucs were examined for 31 ~gyninotifor~nspecies, including represei~lativesof 20 genera. These data were cornpiled into a niatrix of' 249 ck~aractcrs,including 45 cl~nractersof the nervous system, for 44 operational taxonomic units. The inolecular sequence data of Alves-Go~ncsc.1 ccl. (1995) were also analyzed, weighting transitions and transversions equally. TI-ee topologies 1.1-om molecular scqtlence and morphological datascts analyzed independently usiilg ~naxi~nl~rnparsi~iiony are very similar, and to the topology resulting iiroin a corn- bined a~ialysis. These data are uscd to provide phylogcilctic diagnoses for 75 cladcs ailti terminal taxa, inclutlii~gthe Gyrnnotilhr~ries,5 L~nrilic:s,and 29 genera. Two new genera and 14 additional supraspecific taxa are introclllccd. (;?rtnnoh.s+1l~~ctropl~onr.s are round to he nioilophyletic, and together Sol-m the sister group to a11 otliei- gy~i~notifo~.~ns.Rhampliiclltllyidac + Hypopornidae and Stcrnopygiclae + Apteronotidac XI-ealso monopliylctic, the latter of which incll~desall species with a toile-type electric organ discharge (EOD). The f'ossil tl<lli,sc,llais dcterrnined to be a member of Sternopygidae. No rvitlcncc was found for the ~nonopliyly01' the non~inalgenera Ircr.crmcc (Triques), f&nntccnnin (Jordan and Evcrmanri), Aplno17oku.s .SPII.YU11~10 (121cPpi-de) or l'orol~;$rl~.s(Ellis). fL1eg(~do17/ognotf~1~s(Mago-1,cccia) a~~dIfiidio (Miles) arc I-c- gardcd as,j~~niorsynonynis oSApt(.r-onol~c.s.ton.\ti ,\tric.to (nwt~sage). An identification key to tl~crecognized genera is provided. The phylogenetic results pcrrnit sevel-al conclusions regardi~igcharacter cvol~rtion.Stc.rn,o11yg.u.s n,slmh/..c most closely resembles tl~caircestral ~yninotiforrriplienotype. Two clades, Apteronotidae ant1 1:'ig~irtnannin + Khnbdo/ic.Iro/).t possess Illumerous tlcrivcd featlu-es, including two specializations of the centl-al nervous system, related to the lrtilizatio~iof' deep ri\ler channels. Gymnotidac, lil~an~pl~icl~tl~yiclac,and I-Iypoyomidac retain the plesiomo~-phicp~~lse-type EOD and larval hypaxial electric organs into matllrity. The tone-type E011 of Sternopygidac and Apteronotitlac is derived, and the larval electric organ of tlresc taxa dcgc~icratesat rncta~norphosis. Elongate snouts have evolvcd iiidepe~identlyin at least I'oui- ~yrnnotiSorn~lineages, invol\ing difl'ere~itsets of bones. The caudal li~iof' aptcl-onotids is derived Srom the caudal appendage of o~llei-Lgy~notifor~~ls, and is not an internletliate state in the loss of a caudal fin. The actrral nrumber of gyxnnotif'orm spccics in tlie wild is lnuch greater than has previously heen recog nized. Tlicrc are currenlly 108 valid species of ~gym~~otif'ormsin thc litcratl~reancl about 34 ~~nclcscribed species recognized ill Irillsetlrn collections. These species are distribl~tedthroughout the hl~rnidNcotropics, li-om the liio Salado in the I'a~npas of Argentina to tllc Rio San Nicoliis of Cl~iapas,Mexico. Spccics diversity is highest in thr A~niwon(89 species) followed by the Ori~~oco(61 species), the G11yanas (35 species), the Parank (26 spccies) , Northwcstcrn South America (14 species), Southeast Brazil and Uru- grlay (1 2 species), Nortl~eastBrazil (9 species), the Pacific slope of South Ailcrica (7 species), Mitldlc Arnc.1-ica (7 species), and the endorhcic liio Sali-Dlrlcc or rial-thwestern Arge~ltiiia(1 species). Tllcrc arc at least seven tra~~s-i\nde:~ngy~nnotif'orm clades. Key \vol.ds: Oiodiv/.rsil,lr,l)iogmogt-/~/)l~~x c.lcrdLlics, c.ln.s.si/ic.rclion,cotripc~mti-or mor/~hology, ~lc,ctric. Ji.sh(,.\, .c,~?~olu- lion, his ioricr~l~c.olo~~, Solil/t ilrtz~ri~~c. CONTENTS I'AGE ADSTIZA(:T ................................................................................................................................................................................ ii INTRODUCTION ............................................................................................................................................................... 1 A BRIEF I-IISTORY OF (;YMNOTIFORM SYSTEMATICS ................... ... ...................................................................... 2 MATEliIALS AND METHODS ............................................................................................................................................... 6 ('1 .... LII nt Ccrs ..................................................................................................................................................................... 9 1). ,LI .. simony Analysis ..................................................................................................................................................... 10 (:HARACTElI DESCIIIPTIONS ........................... ... ......................................................................................................... 11 PI-M. OGENETIC RELATIONSI-111's .................... ... ....................................................................................................... 62 GYR.INOI.II.OI<MI<S(CI.?\I)E A) ................................................................................................................................................ 64 GYR.INO.I.II)I\E(<.IAI)I . H) ...................................................................................................................................................... (34 EI.I..(:~.I~O~,II~,~I.s G11.1. .......................... .. ............................................................................................................................ (56 G~nr.\~o.ir~.sIANN,\I..US (C.~.,\nk (:) .......................................................................................................................................... 66 G1.,1i~\~o711.\(..II<~II'O + G . .IV(.I;I I.L.IIUS ((.I. .\I)I< D) .............................. .. ................................................................................. 66 G~,!IA~O./~~I.\(~17.l,\~lllll(~ll.\ SI'KCIES-t:KOL!I' A1 1\1<1<TAN11 MIL.I.EK.................................................................................................. 66 (;~,\r,\~o.iln c., II~AI~~srec.~b.s-c.~<oul~ AI.IIEIIT AN11 MII.I.ER................... ... ............................................................................ 66 ~;)i)1~\r0716,\Ntllll.l.~\lll.\ SIVI:CI L.. S-(:I<OIII'AI IIEIU /\NI) MII.I.I<II................................................................................................. 66 SII. I<NOI?'(:OII)E~(CI..\l)l' E) ................................................................................................................................................. 66 1<1l~\hll'lll(.11~111~0101::\ ((.I.,\I)E F) ......................................................................................................................................... 67 ~III~\I\.II'III(.III IIYII)AI< (CI.. \L)E G)......................................................................................................................................... 67 R~I,II\IIJ~~I(.I~I.II~M~II.I.I<IIi\~r) TKOS(:IILI ............................................................................................................................... 67 C~.nwoitii.l~~n~~~~(.rr~~i~~~.\+ IIOI(.I~LI.I(CIAI)I' H) ...................................................................................................................... 68 CI.A~~I~II,~~U,III~~II~I~III.SEI.I.IS................................................................................................................................................ 68 Oo~c:r:nri~TIIIQIIICS .................................................................................................................................................................. 68 H\rl'~)~'~)hll~)i\I< (CI.~\I)E 1) .................................................................................................................................................... 68 I-l~.~wi,onrci.sC;II I ......................... ...... ................................................................................................................................ 68 BII,\(.IIYIIYI~~I,OI\.IIN,\L((.I.I\I)EJ . NI:~I.AX~N) ....................... .. ......................................................................................... 68 BI~I~.~~I.III~~~~I~o~\ILI.YMA(.o-LE(.(.I~\ ......................................................................................................................................
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