PHYLOGENETIC REVISION OF (TELEOSTEI, ATHERI NOPSIDAE), WITH COMMENTS ON THE SYSTEMATICS OF THE SOUTH AMERICAN FRESHWATER FISH BASIL ICHTHYS GIRARD

BRIAN S. DYER

MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 185

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Ann Arbor, November 8,1997 ISSN 076-8405 MIS(:ELIANEOUS PUBLI<'ATIONS MUSEUM OF ZOOI,O(;Y, UNIVERSITY OF MI(:HI(;AN NO. 185

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O<:o~ir~o~;B.M., R.K. (:olwell, irnd S. Nacc111. 1996. The Flowel- Mites of Trinidad 111: Tl~e(;(anus Rhi?~os(,il~.s(Acilri: Ascidac). Misc. Publ. 184. 32 pp, 70 figs. $10.00. Kluge, A,(;., ir11d R.A. Nr~ssb;~rrrn.1!1!)5. A Rcvicw Of Af'rica~l-Mada~;~sca~i Gckkonid 1,iz;rrcl Pliylog- eny Ant1 Riogcog~-aphy(Sqr~amata). Misc. Publ. 183. 20 p11, l l figs. $8.50. IZohlf, F:J. & El.. Kookstein (etls.). 1990. I'rocrrdings of tllc Michisan Mor-pl1onlet1-irsWorkshop. S~xc.I'lthl. 2. 980 pp. With sol'twarc, $25.00. Without sof.tww-e,$17.50. i\lexander; R.D. 1!)!)0. I-low dict Irurr~:r~~sevolve? Rrflcctions on tl~cr~niclrrcly uniclne sprcies. Spec. Pul~l.1. 38 pp. $4.00. Kaxwortliy, (::I. & R.A. Nllssba~l~n.1!4!)4. 11 ~.cviewof the Madagascan Snake (;clner;r I,s~u~lox)~rl~ol~us, I'/jlnrt~~colinn~rc,anel Il(?~(:roliotlon(Squamatia: (:oll~bridar),Misc. Publ. 182. 37 pp, 25 ligs. $1 1.50. Goslinr, W.A. 11)03. A survey of 11r)pcr~jir~~IIIIISC~I~~~UI-C in 11ighc1- teleostran fishes. Occ. 1';lp. 724. 26 pp, 9 tigs. $2.20. DII~IIIII~II,W.E. &,J.A, ($nipbrll. 1992. Hylid hogs of tllc genus I'l~c(ml~~ln:Systematics ~III~~11ylogc- netic relatio~lships.Misc. I'rrl~l. 181. 38 pp, 21 figs. $9.10. Mcl

1,alu-cncc B. I)citch, l5loomlic.lrl I-Iills I)a11ic11). I-101-ning,(;r;rntl Ilavcn Olivia 1'. May~lard,(;oodl.icll Shirley M. McFce, Bi~ttlc<:I-cck Krl)ccca Mc(;owan, Ann Arl)o~. A~~tl~-raFischrr- Ncwman, Ann Ar11o1 Philip H. I'owe~; Ann Arl,o~ S. R/I;r~-tinTaylor, G~-osscPoi~ltc Fal.~ns 1,c.c (:. Rolli1lge1-,cJx officio Phylogenetic revision of Atherinopsinae (Teleostei, Atherinopsidae) ,with comments on the systematics of the South American freshwater fish genus Girard

Brian S. Dyer

Division of Fishes Museum of Zoology and Department of Biology The University of Michigan Ann Arbor, Michigan 481 09-10'79, U.S.A.

Present Address: Centro de Ciencias y Ecolo@aAplicada Escuela de Pesquerias y Cultivos Universidad del Mar Carmen 446, Placeres, Valparaiso, Chile email: [email protected]

MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 185

Ann Arbor, November 8, 1997 I)y(+v;B. S. 1 997. I'liylogert(~licrc.vi.,ion o/'Alh,urin ofxinae (Elrostri, it tlzc~i~~opsidnu),will1 cotnin~nl.~on lh~sj~sl(~m- alics oJ' 1/10 Svuth A ~n(,ricanJresh~o(~ter fish pnus Basilich th ys Girard. Misc. I'zI,~~./\/lu~. Zool. Uni-o. /Michipn, 185:I-64, 46,pgs., 3 lab1r.s. I-Iypotheses of relationships among atherinopsine taxa and their monophyly are revised rising mugilids, Cyprinodontea (belonifonns and cyprinodontiforms) , Atherinoidci, and Mc~lidii~lae 21s outgroups. A~lato~nicalevidence is co~npiledwith published morphological and allozy~nedata for a plryloge~leticanalysis ofAtherinopsinae. 1)iagnostic characters are provided for athrrinopsine gl-oups ill addition to Menicliinae, Menidiini, and Membradini. At11e1-inopsidacis diagnosccl by 20 characters of which six arc ~rnicluewithin atl~erinon~orpl~s. Atherinopsinae, composeel of Atlleri~lopsi~li(North America: Coljjichlkys, Atherinofis, i\th(~rinof~sis, I,oure~stl~e.s)and Sorgeritinini (Sol~tllAmerica: Bnsilichthy.~,Odot~trstkes), is diagnosed by eight cllaracters. A phylogenetic analysis oPrelationships among atherinopsinin genera ~.esultedin two most eq~lallyparsirno- ~iioushypotheses: Colpichtl~j~sis tlie sister to Athminofis, Atherinopsis, and Lour-e~th(,,s,the latter two each others closest relatives; or Leicrc.sll~e~is sister to Atht.rinofi~i.\, A/hm.nof)s, and (~olf~iclrthy.~,the latter two as sister genera. Both sets of relatioilships are mo~-eparsinlonious than prcvious hypotheses by White (mol-- pholo~y)and by Crabtree (allozymes) whiclr proposed C:olf)ichthjls and Atl~.erinol~,s,and Ath(r/inop,\i\ and I,PU~~S~/LCSas sister taxa. A kry is proviclrcl to tlie genera of atherillopsines and to the spccies of atherinopsinins. Athcl.inopsini, di;~gnosrclby seven characters (vomer ~~itlltwo tlcveloped clorsal lami~lac, dcrmosplrenotic wit11 three sensory-canal pores, [I-ifidsupraoccipital crest, infi-apl~a~y~~gobranchial1 cap tilaginous, pointed cranial process oli1rfi-aplla1-y1lgoI11-anchial2, long uncinate process of epibranchial 3 atiaclred proximally, haemal-hypophysis frrnnel), is cornposed of six species. Co@ichthys, wit11 two species (C. ri~gis,C. llubbsi), is diagnosed by nine cl~aractei-s(e.g., toot11 cusps asymmetrical, presence of liyoinandib~~larnerve foramen, < 20 gill rakers oil lower limb offil-st hranchial arch, pharyngeitl jaw teeth hair-like, females u~itllovipositors, and eggs lacking filaments). Athminol~s?finis is diagnosed by frills oS skin betwce~lmaxilla and prcmaxilla, and hvo I-ows of sr~bocularscales. Ath(~inof7,si.ccalifor17,i~nsis is diag- ilosed by seven characters (e.g., der~rrospllcnoticnot enclosed by bone, opcrctllar dorsal process lacking laminar blade-like flange, srllall si~escales). Lrurrrtlzrs, with two species (L.lenuis, L. ,\a?-dina) , is diagnosed by live characters (e.g., pelvic median plate cxtciided to anterior end, scales present on second dor-salfin, rulique beaclr-spawning habits). Sorgenti~lini,di~rgnosed by cight cliari~ctcrs(e.g., ethmomaxillary ligame~ltattached to etli~lioidcarti- lage o~lly,presence oCbasioccipi~a1fenestration, palatorostra1 lig;~ment,precaudal vertebrae 50% or rnorc of total vcl.tcbi-ac, llae~rlal-archf~~nnel), comprises about 25 species. Odonrc~s~lrc?~,~vitll about 20 ~iiarii~e and fl-eshwaterspccics, is diagr~oscclby sever1 characters (e.g., mesethrrioicl abse~lt,e~llarged trigemi~lofacial foramen, niaildibular branch of A1 ~nuscletcnclo~l anterior to lacrimal hranch, opel-cle Se~lestrated). Bnsilic/~l!~ylys,with live 1'reshw;~terspecies, is diagnosed by 22 characters (e.g., lacrimal condyle of lateral ethmoid not extc~ldcdto ventral edge, derrnosl>henoticrriedial flange reduced or absent, vcntl-a1process oSrnaxilla wit11 a condyle, clentary coronoid process flattened dorsally, absence oSa dentary gnathic spine, rtl~momaxillai-yliga~lle~lt not attached to spoon-sllaped palatine dorsal proccss, prccaudal vertebrae over 57% of total vertebrae ant1 Lirst carltlal vertebra over posterior half of anal fill, dorsal head scales with rrverscd imbrication) . Two species groups are recognized ill Bn.cilic/ztl~j~s.The 13a.silic.h//1~~s~nicrol@idotzrs species group corn- prises the nominal species B. microlepidot~csand B. ccnstmlisSrom central and southern Chile. The Basilichtlzjls serrrotilri.~spccies grorlp is coil~posecloSB. .vr~nolilusand B. al-cha(,u,sfrom Pel-LI,and an undescribed specirs from the Rio Loa in nortllcr.11Chilc. Tlie undescribed species is te~itativclyproposccl as sister to the other species of the sei~~otilu.~species grorlp. Key words: Al/~min~fortri~~,s,At!~c!rino/).sini, Sorgrialinini, p/t~llogenc~lic.cy.slrrrrcclics, ~~roq~holo~g,Basiliclithys. Iljl(~r;B.S. 1997. Revisid~~fi1oge1zkLic.a d~ Athen'nof1si17ne(Teleostei, Atheri~zopsidaejy romentam'o.~sistenzriticos del ginm dztkc~crcuirolnSudamwicano, Basilichthys Gi~ard.~Vlisc. Publ. Mus. ZooL, Uniu. Michigan : 1-64, 46figs., 3 t(c/Ilfcs. I,;I monofilia de la subfamilia Atherinopsinae y las hipotesis de relaciones entre sus gkneros son revisadas usando conlo outgroups a Mugilidae, C:yprinodontea (Beloniformes y (:yprinodontiformes) , , y Menidiinae. Datos rnorfolcigicos y e~lzimriticosya publicados son agrupados con caracteres anat6micos nuevos para un analisis Iilogenetico del grnpo. Sc cntregan caracteres diagn6sticos tanto para los grupos atllcrinopsinos como para Menidiinae, Menidiini, y Mcmbradini. Atherinopsidae es diagnosticado por 20 c.nl -.actercs, clc los cuales seis son unicos dentro de Atherinornorpha. Atherinopsinae, cornpuesto poi- Atherinopsini (Norteainkrica: Colf~ichtl~~l.c,Allzminops, Athrm'nof~sis, 1,eur.o.sthe.s) y Sorgentinini (Suclamkrica: Bcrsilichthys, Oda.n,lestlzes),es diagnosticado por ocho caracteres. El anrilisis filoge~~kticode 10s gkneros atherinopsininos dici como resultado dos hip6tesis de relacio~les igualmentc parsimoniosas: CoYichthys es el grupo herinano clc 10s demas gkneros y Ather-inops es hermano clc Alk(~t-ino/~.tisyLeurarlltn; o Lerireestheses el grupo hermano cle 10s demis gkneros, y Atheri~zof~sises hermano clc Ath,vrr'no]~sy Colfiicllzj1.s. Ambas hip6tesis son mas parsimoniosas que aql~ellapresentada por White y Crabtrec, I-~asadaen morl'ologia y en aloenzimas, respectivamente, quicncs propnsieron a (;olf~ic:hthysy i\tlrr.r.in,ops, y Alhc?rinol,szsy I,rurrsthrs como grupos hermanos. Se provee adernis una clave para los gtneros de Atheri~~opsinaey las especies de Atlici-inopsinini. Atheri~ropsininies diagnosticado por sietc caracteres (v6mcr con dos proccsos laminares dorsales bicn desarrolladas, der1noeslc:i16tico con tres poros del canal sensorial, crcsta supraoccipital trifida, primer inS~-ala~-ingohra~~q~~ii~lcartilaginoso, segtmdo infrafaringobranquial con proceso cranial agudo, tercer epibranqrrial con proceso uncinado largo concctado proximalmcnte, y embudo de hip6fisis hemales) y estri compl~estopor seis espccics. Colpic.ht/zyy~,con dos especics (C. reg-is, CL hubbsi), es diagnosticado por ocho caractercs (el., cilspides dentalcs asimktricos, presencia de un foramen del nervio hiomandibulal-, <20 branql~iespinasen el ramo inferior del primer arco branqnial, dientes fa'aringeos finos, hembras con ovipositores, y huevos sill filarneritos) . Atherin,oj~,s@n,is es cliagnosticado por la presencia de un lnargen escarolado de piel entre el rnaxilar y premaxilar; y dos filas de escamas suboculares. Athrn'n,opsi.s rali/omiriisb es diagnosticado por seis caractcrcs (ej., canal dermoesfen6tico abierto, proceso opercular dorsal sin lln ueborde laminado, y escamas de tamaiio pequeiio). Leul-esthe.~,con dos especies (L. tenuis, L. sal-dina), es diagnosticado por cinco caracteres (cj., plaia pelvica mcdiana extendida llasta el extremo anterior, escamas 1xcx~ltewnla base anterior de la segu~ldaaleta dorsal, y hgbitos caracteristicos de desove en playa). Sorgcntinini es diagnosticado por ocho caracteres (ej., ligamento etmomaxilav conectado solo al cartil;~goctrnoidal, presencia de fcnesiraciones en el basioccipital, presencia de un ligarnento palatorostral, ~~~~~~~o de vertebras precaudales 50% o mis del nilrnero de vkrtcbras totalcs, y ernbl~dode arcos hemales) y est;i cornpuesto por unas 25 espccies. Odont(,sth,es,con unas 20 cspccies rnarinas y dnlceacl~icolas,es diagnosticado por siete c;n-;~ctcres(ej., auscncia de inesetmoides, foramen trigeminofacial agrandado, rama mandibul;~rdel tend611 dcl milsculo aductor mandibular A1 anterior a la ralna lacrimal, y op6rculo knestraclo). Bosilichtl~ys,con cinco especies d~~lceacuicolas,es dkngnosticado por 22 caracteres (cj., chndilo lac~-irn;~ltlcl etrnoides lateral sin extensi6n a1 borde ventral, proceso laminar medial del dermoesfen6tico reducido o ausentc, proceso ventral dcl maxilar con un c6ndil0, dentario con proceso coronoides aplanado clol-salmente, ausencia dc lrna espina mandibular del dentario, liganiento rtmomaxilar sin conexion al proceso palati110 dorsal, nilmcro de vertebras prccauclales sobre cl 57% del nilmcro total de vtrtebras, primera vkrtebra caudal sol~rela mitad posterior de la alcta anal, y escamas dorsales de la cabcza con imbricaci6ii invcl-tida). Sc reconocen dos grupos cle especies cn Basilichthys. El grupo de especics ISccsilichthys microlef)idotus esta cortipuesto por las cspccies norninales B. microlrpidotu.sy B. (xrrstrcllisdel centro y sur de Chile. El grupo de cspecics Bnsi6clztlry.s .s~n~o/ilzc.rcstA cornpuesto por B. semotilusy l3. a~rhneusde Peril, y una especie sin describir dcl lZio l,oa en el nortc de Chile. La espccie no descripta esti tentati\lamente propursta como especie hermana dc las dem5s especies del grupo semotilus. Palabras clave: Ath,rri~n@rtrtc~s,Alh~c.rinof)si~ni, Sa.yyc.nlznini, sislpmrilicn /ilogen6tzca, morJ'ologin, Basilichthys. CONTENTS PAGE

INTRODUCTION ...... 1 TAXONOMIC HISTOlW ...... 1 METHODS ...... :4 RESULTS ...... 4 CHARACTERS ...... 4 Neurocrani~unand associated sensory calla1 I~ones...... 7 Jaws and associated ligaments ...... 21 Suspe~lsorit~~nand associated ligaments ...... 26 Branchial basket ...... 33 Pectoral girdle and fin ...... 35 Pelvic girdle and [ill ...... 35 Median fins ...... 35 Axial skeleton ...... 37 Scales ...... 42 Allozy~necharacters ...... 44 SYSTEMATI(:S ...... 44 Monophyly of Atheri~~opsidae...... 44 Monophyly of Menidiinae ...... '15 Monophyly oSAtherinopsinae ...... 45 Monophyly and systematics olhthrrinopsllll...... 45 Colflichthys Hubbs, 1918a ...... 17

Athwi~nopsirGirard. 1854 ...... 18 Leurp.stAe.\ Jordan & Gilbert. 1880 ...... 48 ... Monophyly and systelriatics of Sorgcntinlnl ...... : ...... 48 Orlonte.sth.(is Evermann & Kenclall, 1906 ...... 48 Basilichthys Girarcl. 1854 ...... 49 mic~~ol($idotu.s species group ...... 52 semotilus species group ...... 52 IWTO THE ATHEKINOI'SINE GENERA ANL) SPECIES OF ATHERINOPSINI ...... 53 DISCUSSION ...... 53 SYSTEMATICS ...... 53 CHARACTERS ...... 54 Jaws ...... 54 Temporal sensory c:ulals ...... 55 Axial skclcton ...... 55 CONCLUSIONS ...... 56 ACKNOWLEDGMENTS ...... 56 REFERENCES ...... 56 APPENDIX 1: Material examined ...... 59 APPENDIX 2: List of synapomorphies ...... 61 APPENDIX 3: Ambiguous characters ...... (53 ILLUSTRATIONS r URE PAGE . . Relationships among atlrerinopsid tr~bes...... 8 Relationships among atherinopsine genera ...... 9 Dorsal. lateral. and frontal views of the voiner ...... 10 Dorsal view of the ethrnoid region of Odontesthe~retropinnis ...... 12 Dorsal view ofthe etlrmoid region of Basilichlhys sp ...... 13 Anterior irrfi-aorbitalsand nasal bones ...... 14 Ventrolateral view of the neurocranium of Odonte~thesorientalis ...... 14 Vcntrolateral view of the ncurocranium oP Odonte.~!slhe.~ha1ch1.m' ...... 15 Ventrolateral view of the neul-ocranium of Basilichll~ysmicrolepidotu~ ...... 15 Dorsal view of the left supraorbital and temporal sensory canals and pores ...... 16 I. atcral vic-w of the head and pectoral gii-dlc of Uasilichtl~j~sssp ...... 17 Posterior view oP the left dermosphenotic ...... 18 1. eft veiitrolateral view of the otic and occipital regions of the rieurocranium of . . Me7nb?-crsglbrrti ...... 19 Schematic dorsal and lateral views of the supraoccipital crest ...... 19 Dorsal and lateral views of the left postemporal bone ...... 20 Dorsal view of the lrcad of 13milichtl1ys...... 21 Lateral view of the leftjaws and suspensoi-ium of Odontestl~esretropinnis ...... 22 Mcdial view of the lcftjaws and suspensorium of Odo~ztesthesretropinnis ...... 23 I.ate1.21l view of the lef't.j;~ws.infraorbitals and suspensorium of ...... 24 Medial view of the leftjaws and suspensorium of Basilichthjls austmlis ...... 25 Left rrlaxilla in posterior. anterior. and dorsal views of Basilich.thys ...... 25 Diagram ofjaw liga~rrentsin lateral view of Odon.leslhes relropin.nis ...... 26 Lateral view of the snout region ...... 27 I. ateral view of the 1eft.jaws. infraorbitals and suspeirsorium of Basilichthys semotilus ...... 28 Medial view of the leftjaws and suspensoriurn of Rasilich1l1.ys semolilus ...... 29 Diagrarn in lateral view of the A1 tendon complex of the adductor mandibulae muscle ...... 30 1.ateral. medial. and dorsal views of the left palatine ...... 31 Lateral view of the left preopercle ...... 32 I. .lteral and medial views of the opercle of (Y~iro~tomalabarcae ...... 33 Dorsal view of the left uppcr pharyngeal jaw ...... 34 Lateral and ventral views of the uroliyal ...... 36 Medial view of left pectoral girdle ...... 37 Ilors;~lview or the occipital region of the head. pectoral girdle. and anterior vertebrae of . . Oclon testhr.~.6rc.v~anal~s ...... 38 Ventfirl view of the left pelvic girdle ...... 39 . . I. ateral view of median fins and axial skeleton of Odontesti~antropznnzs ...... 40 Frontal view of caudal vertebra. and lateral view of dorsal fins and axial skeleton of Basilichthys .... 40 I, ateral view of median fins and axial skeleton of 0clon.leslhc~sbonariensis ...... 41 I, atcral view of median fins and axial skeleton of 0don.testhe.s r@a ...... 42 I. atel-al and frontal views of anterior caudal vertebrae of Atherinups ...... 43 I.atc 1.~1shaft of' left cleithr-um with scales ...... 43 I-lypothesis of I-clatioiishipswithin atherinopsinins proposed by Whitc (1985) ...... 46 Alternative equally parsimonious hypothcscs of rclationsliips arnong atherinopsinin genera ...... 47 Monophyly of Rasilichlhys ...... 50 Distrih\~tionmap of Rnsilich.tl~ys ...... 51 Alternative optimization of characters on both cclually parsimonious cladograms of atlrerinopsid relationships ...... 54 Thrcc classes of character-state distributions that result in ambiguous optimizations when mapped on a cladogram ...... 64 TABLES TAR I .IT. 1'AC;I.:

1 History oP classilicatioiis o['athrl-inifol-ln fish ikmilies ...... 2 2 P11ylogenetic classilicatioli oSAtI1r1-inopsidae ...... 3 3 Data niatrix of 20 :rtlieri~iopsi~ietaxa and fivc o~ltgroups...... 5 INTRODUCTION ing the systematics, a discussion section deals with alternative relationships and character evolutio~iin the context of these The fiumily oSNcw World silversides, Atherinopsidae, is corn- pliylogenetic hypotheses. poxcl ol' the slthhmilics Menidiinae and Atherinopsinae. Evi- Within the South American atherinopsines the systematics dence Sol. their monoplnyly :und sistcr-group relationship was of the two species groups of Basilichtlzys are discussed further, p~~oviclccl11y White (1985) and (;hernoff (1986b). White (1985) however, the relationships among species of Odonteslhes are and C1ic1-nofS(lSS(ib) 110th ~~seda combination ofselcctcd taxa treated elsewhere (see Dyel; 1993: 331-501). of' Old World atherinids, ~iielanotaeniids,notocheirids, I~clonilO~~ms,and cyprinodontiforrris as outgroups in their as- sessments of clial-acter polarity. The phylogcnetic hypothesis TAXONOMIC HISTORY ol'atheriniSornl iaterrelationships (Dyer & Chernofl; 1996) used in this str~cly,Ilowever, dl-anlatically changes traditional taxo- Atllerinopsinae was erected by Fowler (1903) for those r~omicgroupi~igs (e.g., Notocheiridae, Phallostetliidae, atheriniSorni fishes witli "premaxillaries not freely protractile, At11e1-inid;lc)and assunnptions of plesiomorl~hicgroups (c.g., the skin co~itinuousmesially with forehead". In the first com- Il~tlo~icr,IZlr~ocl(~,s, M(2lanolnclnic~) long held since Jordan & Hubbs prehensive revision of atherinifor~nsbyJordan & Hubbs (1919) (1!)19). 111 combining the hypotheses of atheriniform (Dyer & the suhfarnily was enlarged to include allnost all American sil- Chcr~iol'l',1996) and atllerinomorph (Rosen & Pawnti, 1981; vel-sides. Subsequently, Schultz (1948:42) redefined Stiassny, 1990, 1993) relationships, outgroups now haw a se- atlierinopsines to include only those genera with distally dilated q~~enti;rlorder of relationships rclative to the atherinopsid prernaxillaries and a taperi~iggas bladder surrounded by modi- ingroup. For thesc rcasons some changes are expected in taxic fiecl, expanded harmal arches, tlie haemal funnel. With this rcl:rtionships witlrin the fiunily ;lnd changes in grot111diagnoscs. diagnosis, all Anierican silverside genera that did not share the Whitc (1985) provided tlic Grst hypothesis (11' phylogcnetic hacnial fl~nnclcharacteristic were accommodated in rclarionsl~ipswithin Atlicrinopsinae and recognized two tribes: Mcnidiinae. Schultz (1948) allied Atherinopsinae with Atherinopsini (North America; AlIrerinof).si.c, Allrerinof~s, Atlicrininae because some atherinine species also have an (~olj)iclr/hys,I.YI~)PS~~I~PS) and Sorgcntinini (South Amel-ica; haemal firnilel. Tlie relationship between Menidiinae and l~cr,siliclrll~~~.cand Odonl~!,s/h(:c.). Cr;ibtrcc (1987) presented two cla- Atherinopsinae was unclear until Patten (1978) proposed them cloglxnis hased on allo~ymedata with networli relationships idcn- as sister groups in his unp~tblishedthesis, a relationship that tic:~l to White's cladograni, 11ut wliich differed in placement of was later col-roborated by White (1985). Chernoff (1986b) re- the i.oo[. Both Wliitc and <:r:~l~tree,however, did not have an vised Menidiiiiae pliylogenetically and presented additional adVq1t;~tcsample ~Ssorgentininspecies. Consideri~~gthat about evidcnce for tlie sistel--group relationship between the subiianii- 80% of' tlre apl~roximatcly30 species of atherinopsines are lies and their monophyly. The haemal f~~nnelwas consequently sorgentinins, the inclusion of'niissing specics into tlie phyloge- hypothesized to have evolved independently in atherinines and nclic analysis cor~ltlsigni1icantly affect the relatio~isliipspro- atherinopsines (White, 1985). The sister-group relationship ~x)sc'.dby White (1985) ;~ndCrabtree ( 1987). between the New World subhmilies is now formally recognized Atlicrinol~si~~insare coastal iriarine fislics li-orn tlie west coast as Atherinopsidae, a family that is relatively distantly related to ol' Nortll Anie~-ic;~which occasionally cilter estuarine environ- Atherinidae (Saeed el aL, 1994; Dyer & Chernoff, 1996), and Illellts. Thc group fiungcs fl-on1 Vancouver Island, Canada, to sister group to other atheriniforms (Dyer & ChernoPP, 1996). so~~tllernRaja CnliSo~.iiiaant1 the GulfoS(:alifornia. Four gcn- Becat~scof significant changes in relationships and cr;~arc: c~~r~-entlyrecognized: Alherinops (topsmelt) and of atlieriniSorm fish families, a summary of the history of these i\/lrorinoj),si.s(jacksmelt) witli one species each, and (~olpirlrll~ys changes is depicted in Table 1. Table 2 is a classification of ;rntl Len:r~!,sllrc~s(grr~nio~~) witli two species each. atheriniform fishes from Dyer & Chernoff (1996) depicting the Sorgcntinin sperirs, gci~e~~llyknown as ptjm-911?.s,inhabit tern- higher taxonomic categories used here. perate ni;n.iiie, est~~arinc,and S~-esliwatcrcnvil-onrnents. Tlie Atherinopsinae, as recognized by Schultz (1948), was com- coast:~lmarinc forrns r;lllge from northern Peru and southeast- posed of seven genera: Athmn,opsis, Atlzerinops, Austromenidia, ern Brazil to tlic southern tip of Tierra del Fuego, including Rasilicl~,lhy,s,Hubbsiella, Leureslhes, and . These gen- thC Falliland (Malvinas) Islands and the Juan Fcrnandez Archi- era were grouped into the North American Atherinopsini and lxlago Fresliwatcr forms are Sound in the \vest-draining rivers the South American Sorgentinini tribes (White, 1985). The 01' Pert1 and Chile, in Patagonia, a~idthe Panipas region of Ar- monophyly of these tribes was weakly supported by Crabtree's gmitina, Urugi~ay,and southeastern Bra~il.At present only two (1987) distance-treated electrophoretic data, however, tlie char- ol' tlic I3 descril~cclgenera ol' sorgcntinins arc rccogni~ed: acter-coded allozynie data showed an AthPn'n.opsi.~I~e~~resthesclade Bn..silicl~,ll~,~~,swith 5 nominal species and On'on/es/l~~~,s.cvitli46 norni- to be more closely related to an 0dontesthe.s-Basilicht/rys clade. 1ii11 spccics. Atherinopsini is composed of Atlzerinops, Colpichlhys, The object oS this stt~tlyis to combine the data of White Athe~~zopsi.~,and Leur~slhes.The two former genera and the lat- (1085),Cl~criioSS (198(ib), and (:rabtree (1987) with the results ter two genera were proposed as sister groups by White (1985). ol'n~yown ~norpl~ologicalstudies in order to reevahlate the phy- The Al/rrn'?rul/;c-~o(l,ic/~t~zy~and the Athennopsis-Leuresthes clades logcnetic ~.clationshipsanlong atherinopsinc genera. A review were also supported by enzymatic data (Crabtree, 1987). oS tlic taxo~~oniichistol-y of the snhfi~nilypreceeds the meth- FIubb.siellawas synonymized into L~uresthesby Moffat & Thomson otls rr~ilizctlin tlie phylogcnctic analysis and character dcscrip- (1975) and Co@ichlhys was retained as distinct from Atherinofis tio~isa1.c prese~rl~dprior to the phylogcnetic results. Two by Todd (1 '376), White (1985), and Crabtree (1989). Although cq~~;~lly-p;~rsi~iio~~io~~strees arc find in this study and follow- Hubbs (1918b) recognized a number of Alherinops species, sub- Table 1. History of classifications of atheriniform fish families.

Jordan & Hubbs Schultz Rosen Rosen & Parenti Parenti Eschmeyer Nelson Saeed, Ivantsoff Dyer & Chernoff 1919 1948,1950 1964 1981 1984 1990 1994 & Crowley, 1994 1996 ATHERINIFORMES ATHERINIDAE AmRINIDAE ATHENNOIDEA DMSION I DMSION I ATHERImAE ATHERINIFORMES A~RINOIDEA ATHERINIFORMES

BEDOTIOIDEI Bedotiinae -Bedotiinae Bedotiidae -Bedotiidae Bedotiinae -Bedotiidae --- Bedotiidae \ / MELANOTAENIOIDEI \ Rheoclinae --- Rheoclinae \ Melanotaeniidae / Melanotaeniidae , ,Melanotaeniinae -Melanotaeniidae -Melanotaeniidae 4Melanotaeniidae Melanotaeniidae Melanotaeniinae -Melanotaeniinae \ Telmatherinidae

\ Notocheirinae : Notocheiridae ' 1Isonidae Isonidae Isonidae -Notocheirinae -Notocheiridae /: \ Notocheiridae '-I '-I Isonidae j-" Atherinopsinae Atherinopsinae Atherinopsinae Atherinidae -Atherinidae -Atherinidae Atherinidae -i------Atherinopsidae: -Atherinops~dae Menidiinae Menidiinae

PHALLOSTETHOIDEA PHALLOSTETHOIDEA

Phallostethidae Dentatherinidae -Dentatherininae -Dentatherinidae -Dentatherinidae - PHALLOSTETHIDAE Phallostethidae Phallostethidae

Neostethidae Phallostethidae -PHALLOSTETHIDAE -Phallostethidae -Phallostethidae

Nannatherininae *

( +in Kuhliidae) Girard's (1855) detailed description of the type spccies B. nzzcrolejidotu~Ucnyns), iilcluded meristic and morphometric Scrics Crccnwood, Rosen, Weit~rnan,Myers information as well as the phrase "...protrusion of upper jaw Order Athcrinifornics Rosen beyond the lower". The latter feature was subsequently adopted Fiu~iilyAtlierinopsidae Fowler as the diagnostic trait of Basilichthysby many authors (Burmeister, Sublamily Athcrinopsinae Fowler 1861; Cope, 1874, 1878; Abbott, 1899; Fowler, 1903; Lahille, TI-ibci\thcrinopsini Fowler 1929a,b; Szidat & Nani, 1951; Aramburu & Ringuelet, 1967; Gcntls A/h(>r-i1101)sStcintlarhner scdis nl,u/nbili~ Ringuelet et al., 1967; Dazarola, 1972). Girard's (1855) descrip- Gellr~sAth(,rinoj,sis Cirard stdis ~l~titnbilis tion was reviewed by Thompson (1916) and again by Eigenmanil Genus Col/~irl~tlzy.sHuhbs .s~disrn~itnbili.~ (1927), both of wlrom conchtded that Basilichtlzy.~is a group of (;cnlrs L~~rr~<\thesJorda~r& Gilbert serlis m~rtobilis fishes with a Genum between the upperjaw and the forehead. Tribc Sorgcntinini Pianta dc Risso & Risso' This altcrnativc was accepted by a different suite of authors (;c~~usO~OII~IJ.\~~PY Ev~rlnann8C Kc~idi~ll Gcnus Ijnsilichtl~ysGirilrd (Hubbs, 1918a;Jordan & Hubbs, 1919; Fowler, 1945a,b, 1951; ~~ric.)uk(~)idol~i.sspecies group I-Iildebrand, 1946; Schultz, 1948; De Buen, 1950, 1955; Mann, 111ic.r-ol.f,~rlof7i.sJenyns 1954; Arratia, 1981). Agreeing with Thompson (1916), Hubbs rrlislmlis Eigenrniunn (1918a) created Austromenzdza to distinguish the small-scale spe- .sr~~~otilusspccics group cies witll upperjaw protrusion from Basilich,tlz~~s(species with a ~f'll~O/ll~.S(:Ope frenum) and frorn Odontesthes species with large scales. nrc11~11,ctsCope Eigcnmann (1927) described Cauque to include four freshwa- Sr~ljlhmilyMcnitliin:rc Schultz ter species with crenate scales that are endemic to Chile. Tribc Mc~iidii~iiScll~~lrz Patagoninc~was described by Eigenmann (1927) in a footnote (:cnus ~l~ir-o.\lorrrftSwainoll srdic 11rrt ffrbi/zs to accornrnodate 0. hatchmi (Eigenmann, 1'309),the freshwater (k1i11sl,nhirl(.c./lic~.r Cope .t/.rlismlitnhilis species he described from Patagonia. Lahille (1929a) re- Gcnus h!J1,1iic11nUonapartc s(~rli.s~nritnbiL~s Gcnlls l'oblnnn llc KIIC-nsrdzs ~~~~itnbilis described most sorgenti~li~lspecies based on morpllometrics Tribe Mcmbradini (:hcrnolf and placed thcm in Ha~ilichthy~.Schultz (1948) considered Gcnrls /\IIr,~rin~llr~Stcindi~chncr Cauqueand Patagonina as synonyms of Austro~nenidia.Unable to Gcnlrs M(,~nhrcl.\Uonaparte find sufficient evidence tojustify the use of generic names such Gcnus iM(!ln~rorItir~zisMctzelaal- as Austromenidin, Cauque, and Kro~zia,De Buen (1950) recog- Subol.tlcr A~hcrinoitlci nized the oldest names only, Odonlesthe,~and Basilichlhys. Later, Rinrily Notocheiridae Schlll~z however, De Ruen (1953) described Ynci in his revision of the I~~ffi~osdcrAtherines Dyer & Cllesnoff silversides of Uruguay, recognized Kronia Miranda-Ribeiro and Eimily Mcl;inotacniitlac Gill A,ustroalhprinccMarrero, and described Tu$a as a subgenus of S~tbfarnilyBcdo~iinacJortl~r~~ & I-lttbhs .s(,d~.sm7ilnbilis 0dontnthe.s. Au.stronth~m.n,aMarrero (1950) has been used mostly Sltblknlily Mclanoracniinac Gill srdis 17zutnbili.r ior 0. inczJa (Jcnyns), but is a nonlen nudziln because no type S~b~uriilyPseudoniugilillae Iher .\~,tlh~?ztitahil;.\ Tribc Pscl~don~ugiliniI

TaI~le3. Data matrix of 20 athcrinopsine taxa and five outgroups in upper case. Characters are ordered according to their position on the body: Ncr~rocranirll~land associated sensoly canal bones (1-36);Jaws and associated ligaments (37-53); Suspensorium and associated ligaments (5468); Uranchial Imsket (70-74); Pectoral girdlc and fins (75-79); Pelvic girdle and fins (80-81); Median fins (82-85); Axial skeleton (86-95); Scales (96-103); Atherinomorpha (104); Athcriniforrncs (105); Mcnidiinae (106); Enzymatic (107-123): quantitative coding and qualitative coding, the latter in parenthesis when they diff'cr. --- - -.-- . . .- -- -.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 - -- -. MUGILIDAE 0000000000000000001 10000 CYl'R1NOL)ONTEA 0 0 0 0 0 0 1 - 0 0 0 0 0 0 - 0 0 0 0 0 0 1 - 0 ATI-IEKINOIDEI 0000200-000000001 0000000 MENIDIINI 001 1201-00000000011101-0 MEMBRA1)INI 001 1201-0000000001110000 A th(~i7~oj1.s 1031211-002200101 11 11010 Colfjzr11/IIJI,\ 1031211-0022001011111010 Athcrinoj,sis 1031211-002200111 1101010 12~~~~r/?.\//~(!.\ 0031211-0021001111111010 N. senrotikr,\ 1120110101211010111101-0 n. "I~)(I" 1120110101111010111101-0 13. 1,17,.,~Ill,l/lof11s I0:!0110101111010111101-0 0. incisrt 100021011122000011110021 0. ~~igrir,r~,s 1000211-1 122000011110011 0. veg;/r I00021021122010011110021 0.gr(~(:ili,\ 1000211-1 122010011110021 0. s~nilti 100021021 1220100111 10021 0. brrai/rnrrlt.\ 1000211-1 12200001 11 1001 1 0. hntclr(,~.i 1000211-1122000011110011 0. P/.r.ligin~ 100021021 12?000011100011 0. bonnrirn,.si.s 100021011122000011110011 0. reLroJ~Pfrr!is 1000211-1 12200001 1110021 0. nrg~,nljlr(,lc.\i.\ 10002102112200001111001 1 0. h1orro7,sic 1000211-1122000011110021 0. f~lrl/l~ll.\~.\ I00021021122000011110011

- - - -

MUG11 .II)AE (2'PRINOL)ONTEA K~HEIIINOI1)ISI MENIDIINI MEMRIIADINI A ll~crdno/~s Co~~ich/lry.s A 1h crin o~/)si.c I,PIITI,S/~(,.\ 8. ,sr~rrolil~r,s n. "loo B. ncic.ral.l,~/lo/lu 0. inrisrr 0. ,11igrir (LIl.\ 0. r(

p~ ~ - ~ - - -- 51 52 53 54 55 56 57 58 .59 60 (51 62 63 64 65 (Xi 67 68 (i9 70 71 72 73 74

- ~ - -- MUGILIDAE 0 00 10 10 0 0 20 00 00 0 0 0 0 0 0 0 0 0 CYPRINODONTEA 0 0 0 0 ? 0 ? 11 00 0 ? 0 0 0 0 20 20 0O&l 0 ATI-IERINOIDEI 0 00&1 0 0 00 0 0 ? 0 0? 0 0 0 0 20 0 0 0 O&l 0 MENIDIINI 0 0 1 11 0 1 0 1 01 00 00 0 10&2 0 00 0 1 0 MEMBRADINI 0 011 10 10 0 I1 00 00 0 10x20 20 0 10 Alh(?rino/)s 0 0 0 2 I00 0 0 0 100 00 0 120 11 10 1 Colplchtliys 0 0 0 2 1 0 0 0 0 22 0 0 0 0 0 1 20 11 10 0 A tl~(,rirro,b.sic 00021 01 0001 00 1001 20 I1 20 1 L~tir(~,~.slh(:,\ 002210100120000012011200 B. .sonotilus 110 30 10 11 02 11 10 01 00 00 00 0 B. "lon" 110 30 10 11 02 1 I10 01 00 0000 0 B. r~iic:rol($~idotus 1 10 3 0 10 08cl 1 02 1 0 I 0 0 I 0 0 0 0 0 0 0 0. inc.i.\rr 0 02 21 11 0 0 12 00 0 0 0 I 1 I 0 0 0 0 0 0. r~igrirnns 00221 11 1012101211 11 00000 0. r/xi/I 00221 110012001211 11 00000 0. pcilis 00221110012001211110O000 0. snlitti 002211100120012111100000 0. br(?~inrrrilz~ 00221 1101 120001 11 I100000 0. 1intc.lic.r.i 0 02 21 11 O&II 0 2 0 0 0 1 10 20 00 00 0 0. PPt-~~ginr 0 0 2 21 1? 0 0 I20O 010 0 20 0 0 0 0 0 0. bonnric~rl~is 0 0 2 21 11 0 0 12 0 0 0 2 11 10 00 0 0 0 0. wtropinnis 0 02 21 I1 00 1200 01 10 20 00 00 0 0. nrger~tirren~is 0022111001200021I lIO0000 0. I~~i~n(~nsis 00221 1001 120002I12000000 0.plrrlen.\h 0 02 21 11 0 0 12 0 0 0 2 11 11 00 0 0 0

-- -- - ~ ~-- -~

MUGILIDAE 00 0 00 0 0 0 0 0 0 0 0 0O 0 0- - 0 0 0 0 0 0 I (,WRINODONTEA 00 0 00 0 1- - - - 0 00 0 0 0 - - 0 0 0 ? - 0 0 ATI-IERINOIDEI 0 0 0 0 01 1 Okl 0 0 ? 0 0 0 0 0 0 - - 0 0 0 ? - 0 0 MENIDIINI 111lII11OI10010 0 0-- 0 1 0 2 0 0 0 MEMBRADINI 11111II1011001000-- 0 10 2 0 0 0 A lhrrin ops 011111210200011 lo-- 0 0 0 2 0 0 0 (:ol/)ic.lrtl1ys 11 1I1 120&10 1100 10&11 0 - - 0 0 0 0 0 10 Athrrino~).si.s 01111120020001 110 - - 0 0 0 0 0 10 LPILT(J,S~~A(,,S 011110110200011 lo-- 0 0 0 0 0 11 B. s~rnolilus 011111201112011 020002 2 11 0 1 B. "locr " 0111I120111201102000221100 B. m7mli'/1zdolus 0111112011120110200021 1100 0. iirci.sn 111121210210000 0 0 - - 0 0 0 0 0 10 0. nigricnns I1 I111200200000 0 0 - - 000I120 0. r(!,girr lI112120020000I 040 10001020 0. grr~cilis 11 11 2 120 0 20 0 0 01 0 4 1 10 0 0 0 0 20 0. snritli lIII2120020000104 11000 1020 0. br-c.r~innnli.s 0III?02001?101I 03 I10? 01 120 0. hrrtch(:ri 01111120011101101001 100010 0. pr.r7i~ioc 011121210?01111 0300 11 0200 1 0. bonrrlder~sis 111121210?01111 0 3 0 0 100 0 0 0 I 0. wh.opinni.s 11I121210?01I11 0 10011 0200 1 0. cr~g(!!ntinensis 11112121020?111030 100010 11 0. hn~rimsis 011121200111I110300 1100 101 0. plnlrnsis 111121200201001 0 3 110 0 0 10 10

- ~ -- - ~- - ~p-~~~-~ - ---~-- ~ - Table 3. (cont.)

MU<;ILIDAE lOOOOO???? ? ??? ? ?? (WlUNODONTEA 0 0 01 0 O ? ? ? ? ? ? ? ? ? ? ? ATI-IERINOIDEI OOOlIO???? ? ??? ?? MENIDIINI 0 00 1 1 1 1&4 1 3&4 5&6 5(7) 48.55 5&6 1&6 4(5) 8(9) 3 RIIERIIBI~lNl OOOllI???? ? ??? ?? A//IW~II~/),\ 100110111 1111 1 1 11 (~ol/)ic.l~thp~ 0 0 0 11 0 1&2 11&3 21(1&4) 1 2 5 15(5&6)1 iill~c>rl~~o/).,i, I101101121 2 112 121 Lrutx,s/l~o 10 0 11 0 11 2 1 3 1 13&4 2(2&3) 3&4 1 B. APIILO~~~~LS 110110???? ? ??? ? ?? 13. "Ion " OIOIIO???? ? ??? ?? 13. ~nic.,i?k.l,~rlolu.r 0 10 1I 0 32 4 34(5) 2 3 6 3(4) 6(7) 1 0. i~~rrscr 002110???? ? ??? ?? 0. 17 igric.0rr s 012110???? ? ??? ?? 0. v(:qi(~ 0 I 0 I 10 32 5 4 3(6) 3 4 7 3(4) 7(8) 2 0.prilu OIOIIO???? ? ??? ?? 0. trnilli OlOllO???? ? ??? ?? 0. Or(!oin~lnl~s 112110???? ? ??? ? ?? 0. I~nlch(.,% llOllO???? ? ?? ? ? ?? 0. /)(*vir,~pi(r(, lOOIIO???? ? ??? ? ?? 0. bo.nn,irnsic lOOlIO???? ? ??? ? ?? 0. r(!/ro/~i~~wi\ IOOlIO???? ? ??? ? ?? 0. crrgmlirr~~rci,\ IOlllO???? ? ??? ? ?? 0. I~~rrrr(,n,,\is IOOllO???? ? ??? ? ?? 0. plnl(!~~si,t 010110???? ? ??? ??

-p~~-- - - liI) for that character. When both i~ldiceshave equal values vomer has a pair of parasagittal laminae that project o~llyone Gg~rrcis sllown. Two sets of indices reflect the diff'ei-- posterodorsally over the ethmoid cartilage and contact the cnt \values ill each nlost-parsirnonio~~stree. Mt~ltistatecllarac- rnesethmoid (Fig. 3B; White, 1985). In Basilichthys, the ante- ters analyzed as 11on-additiveare denoted with 'n-add', and those rior face of the vomrr is projected posterodorsally between the a~lalyzedas additive are denoted with 'acid'. A certain amount dorsomedial ridge and lateral condyles, and houses an of morphologic variation is describctl that is not coded in the anteroventral portion of the ethmoid cartilage. The dorsal lami- data matrix; this i~lforn~ationis placed below the coding defini- nae are far more developed in atherinopsinins than in Lions that arc in parcnthcsis. Basilichthys (Fig. 3C, 3D). The vomer in Odontesthes has a very narrow anterior face, hard!y overlapping the anteroventral Ncurocra~ii~r~~land associated sensol-y can;ll bones rnargin of the ethmoid cartilage, and has no posterodorsal lami- nar projections (Fig 3E) (O=no posterodorsal lamina on the 1 Vo~~~c.rdorso~~~c~~/~~~In(lxP- The ~ncd~an process of the voIner, anterior face of the vomer; l=presence of a single, median, and whcn plcscnt, has a ~o~mdcddo~sal Lace 111 ather~n~fo~rnsand wide posterodorsal laminar process; 2=presence of two outg~oupj A cle~~vccl cond~t~on of atherinop~~nes1s presence parasagittal posterodorsal laminae; 3=presence of two well-de- of;^ sll;lrp ridge on t11c c1ors:rl ~niclli~leof the median process of veloped parasagittal posterodorsal laminae; 1.00; n-add) thc volllcr (Fig. 3B, 3(:, 3D, :3E). This condition is present in all athcrinopsines cxcept Lpurustl~rs.(O=median process of vomer 4 Vomerine anterior she&- The anterior face of the vomer is rounded; l=median process with dorsal ridge; 0.50,0.83) narrow in atherinopsid outgroups (Dyer & Chernoff, 1996; Parenti, 1981). A derived condition present in Menidiinae and 2 Vo~nerdorsnl .sj~i,n~.-The plesiomorphic condition for Atherinopsini is a dorsoventrally flattened extension of the an- atherinomorplis is absence of a posterior projection of the terior face of'the vomer into a shelf-like elongation of the ros- vorneri~ledorsomedial riclgc. The Bnsilicktl~yssemotilus species tral region (Fig. 3A, 3B). This condition is not present in group has ;r unique condition in which the clorsornedial ridge Sorgentinini (Fig. 3C, 3D, 3E). (O=vomer anterior face narrow; of the vomcr projects postcrotlorsally in the Porn1 oP a spine, l=vomer with anterior shelf; 0.50, 0.80) wlrich clocs not co~itacttllc 111c~cthrnoid(Fig. 3D). (O=no vomer- ine spine; I=vomerine dorsomedial spine present; 1.00) 5 Vommuenlml face curualure (Dyer & Chernoff, 1996).- The lateral condyles of the vomer are level with thc median ventral ? W)/r~lirirupo.sl(wodorsc~l I(mi%n,cl (White, 1985).- A relatively line in atheriniform outgroups and the ventral face of the vomer low antc~ior I,rce of the vomer -between the lateral condyles- is is flat. In all atheriniforms examined the vomer is arched ven- the plesio~norpllicconditio~i for atherinomorplis. A derived trally such that the lateral condyles are ventral relative to the condition ofme~lidiiilesis a broad, median lamina that projects median ventral line (Dyer & Chernoff, 1996). This ventral arch posteroclorsally and ar-tic~11;rteswith the mesethmoid (Fig. 3A; of the vomer results in a concave ventral face in transverse sec- (:hrrnoll; 198(ib). 111 Atherinopsini the anterior face of thc tion (Fig. 3, frontal views). In Basilichthys the vomer is arched ATHERINOPSIDAE

l'ig. 1. Relatiorrslrips ainolig atherinopsid ~rihrs(I.=403, C:l=O.(i'L, RT=0.78). Diagnostic characters, r~n;l~nbiguonslyderived cfirics at that noclr in boll1 cqr~tllyptrsimonious trees, are in bold. IJriicluc cl-~ar;tctc~-swithin atheriniforms are opposite 1)lac.k bars; and ho~noplasiouscharacters iirde- ~~~rtlentlyderived else~vltel-c.in athcriniforms are opposite wliitr hal-s. Diagnostic cl1;ll-actel.sarrtl adclitio~lalsupport is listed in Appendix 2.

to a lesm extent .lnd, conscqucntly, the nrlterove~~tralface of sent (c.g., Pselrdoilltrgiliriae, Phallostetl~idar).Vomcrinc teeth the voiner 1s re1'~trvelyil,~ttened fi ontally (Fig 3C, 3D) (O=vomer are absent in Cyprinodontea except aplocheiloids (Parenti, ventral face flat; l=vomer ventral face slightly concave; 2=vomer 1981: 401), Atherinopsini (Fig. 3B), arld in Menidiinae except ventral face slrongly concave; 0.67, 0.86; add) for Alh,erinella ~erriuonzer(ChernoSS, 1986a), Chiroslonzcr. humboldtinnu,m, CIIL(.stor eslor, and Ch. hartoni (Barbour, 1973). 6 l,'lh~r~,oitlcarlilag(:,/i)ltied ven,trnl o/-oo,rrer midline.- The et11- In sorgentinins, presence of tccth is variable within and anlong lnoicl cartilage block is dorsal lo all portions of' the vorner in specics, cxccpt in Odo?tle~lhr,shatcheri, 0. Iz~mensis,0. rc.lrofiinn.is, atllerinif'orms and outgro~~ps.A derived condition found in 0.nigrican~, 0. gmcili~,arid the subgeil~~sCauqu.c~ (Fig. 3D) which atl~eri~lopsirlesis Li)r the lateral face of thc cthmoid cartilage, consis(cntly lack teeth. Rasilichth~1.s usually has a single, medial that ~vhicllarticulates with the palatine, to be folded with the patch (Fig. 31)). (O=vomerine teeth present; l=vomerine teeth ventral arch or the vorncr (Ch. 5) to a position below the vorner. absent; 0.13,0.42) ine midline (Fig. 3). (O=ethmoid cartilage dorsal of vomer mid- line; l=ethmoid cartilage folded ventral of vomer midline; 1.00) 8 N~cm,herofuonrerine roorh l)t~tchr~,s.-(O=vomerine teeth in a band; l=single median vomerine tooth patch; 2=three vomer- 7 Voi~erineteelh.- Vomerine teeth in atl~erinoniol-ph ine tooth patches; 0.50, 0.60; add) outgroups are in a U o1.V-shaped band along the antcrovcntral border of the bone. I11 atherinoids, the plesiomorphic condi- 9 Mrsrthmoicl (White, 1985; Chcrnoff, 1986b) .- The tion is fouird in bedotiines, melanotacniincs, arld some plesiomorphic form of tllc atherinornorph dorsal mesethrnoid atlierinids, but vomerine teeth arc also So~~ndin two sepal-ate (Tigano & Parenti, 1988) is a flat or slightly convex disk-like patches under the lateral condyles (e.g., I,w, Ath,(prr'on,),or ab- bone (Chcrnoff, 1986l-I).The size of the mesethmoid, its shape, ATHERINOPSIDAE ATHERINOPSINAE ATHERlNOPSlNl SORGENTlNlNl

ATHERINOPSINAE ATHERINOPSINI SORGENTININ l

Fig. 2. lielatio~lshipsamong atlleri~lopsincgrnera (1.=403, CIy0.62, FU=0.78).Diagnostic ch;~ractel-s,unalnbiguously derived traits at that node in I~olhccl11;1lly p;u'silr~o~rious 11-ccs, iI1.c irr I>oltl. tlniqne cllal.ac.tc.rs withirr atlleriniSo~-msare opposite black bars; reversed characters (with an "R) are op~x)sitcgrey 1x11-s;and honioplasious chal-ac1cr.siridepentlcrltly tier-ivecl elsewhcrr iri ;~tlleriniformsare opposite white bars. Diagnostic characters ant1 ;~tltlilio~l:~lsupport i lisrctl ill Appclltlix 2; A, Co/$~?rhlly.~is sistrr to Alh(~rir~o/~.\,Athvrinopsis, and Leu?-ecttlcs;B, Lrure.\the.~is sistcr to Athcr~nqpsis,

/\//1('1~1!0/),5,2llltl (.'O//);~/!//!>l\. MIS(..PUUL. Mus.. ZOOL.,UNIV. MICH., NO. 185

Ch. 2: dorsal spine

Fig. 3. Dorsal, l;r~cral,arltl 1.1-ont;rl\,iews of' tl~rvolllel.; i\, M~17iilznho:yllzno (UMMZ 107098); B, /i/lrerinops @nir (UMMZ 232001); C, Br~silic.hll~~~s ciztslmli.\ (UWIMZ 21 5508); 11, Uo.\ilic.hll~?ss~inolil~r, (UMWIZ 21 8050, 21 5520); E, Odon~leslhc.~.brc?,~orrnLs (UMMZ 215459). ~III~which bones it contacts is quite variable within condition is Ior the lacrimal condyle 01 the lateral ethmoid to atherinornorphs (Dycr & Chernoff, 1996). Absence of a extend to the ventral edge of the preorbital wing's lateral ridge nrcsethmoid is a derived condition of Odonle,\Lhes (Fig. 4; White, (Figs 7,8). A derived condit~onCound in Baszlzchthys is for the 1985; De la Hoz &Vial, 1988). A mesethmoid is also lacking in lacrimal condyle not to reach the ventral edge of the lateral the atherinoids Notoch~irus,Craterocephalz~s, ancl Pseudom?~~il ridge (Fig. 9). (O=lacrimal condyle of lateral ethmoid extended (Dyer 8c Cliei-noff, 1996). (O=mesethmoid present; to ventral edge of lateral ridge; l=lacrimal condyle of lateral l=mesethmoid absent; 1.00) ethmoid does not reach the ventral edge of the lateral ridge; The meseth~noiclis morphologically diverse in the 1.00) incmbradins; either convex, concave, or step-shaped, with or without ridges, and contxting the dorsal lamina of the vomer 14 Lacrimal condyle shape- (O=lacrimalcondyle of lateral eth- (<:hernoI'f, 198611). 111 atlici-iiiopsines the mesethmoid is con- moid oval in shape; l=lacrimal condyle of lateral ethmoid en- vex, oval to squarish, with anterolateral and posterolateral larged and rounded in shape; 1.00) (cndocho~ldral)processes in larger specimens. In athcrinopsinins the anterolateral processes contact the paired 15 Lateral vthmoirl ventral contlylr (White, 1985).- In dorsal lanli~iaeor the vorncr (Ch. 3) and the posterolateral pro- atheriniforms and tnugilids the ventral ridge oC the lateral cesses arc overlapped by tlie frontal. The mcsethmoid is - ethmoicl's preorbital wing has a medial cartilaginous condyle I;ltcd in Iln.c%liclLlhys(Fig. 5; 1)e la Hoz & Tosti-CI-oce, 1981: fig. (Figs 7,8)that articulates with thc palatoquadrate cartilage pos- 2) as in tlir menidiines illlzerilzslla halsana and A. callida terior to the palatine. Atherinopsiili and Basilichthys have a de- ((:hernoff, 1986b),and [he postcrolate~-alprocesses arc sliglilty rived condition in that the cartilaginous condyle is ossified and overlirpped by the frontal in larger specimens. displaced anteriol-ly from the ventral I-idge and directed anteroventrally to the palatoquadrate cartilage (Fig. 9; bony 10 l

ethmo-palatine ligament

lateral ethmoid

Fig. 4. Dorsal vicw of' the c~lllrioitlrcgi011 of' Ot!ont/l,thr, wtro/~~nn~t(IJMMZ 21879(i). I.?[[ 1ac.1-imal21ntl nasal I~o~rc-a1.el11ovcd pores across taxa. Tile plcsiomorphic co~~ditionfor 18 Si('h~?totic./)os/orDitnl proccJ.s.s ((:hrrnott, 198611) .- A slriall atheriniforms as inferred from the pattern prcscnt in n~tlgilids and narrow postorbital process with only the levatol- ;~rcus (Fig. IOA) arid cyprinodontiforms is as follows: pore two is con- palatini (LAP) muscle attached, is tlie plesiornorphic condition nected to the nasal canal, pore six is connected to the forlncl ill atl~erinoids,(:yprinodontca, and mugilids. 'I'lic de- dermosphenotic and pterotic canals, and pore eight is con- rived condition of atherinopsids is a wide basecl postorbital pro- nected to the temporal-bone canals. Frontal pores thrcc, five, ccss (Figs 7, 8, 9) with an c~llat-gcdposterodorsal face to wl~icl-1 and six arc directed laterally, and pore four is tlirected mcdi- the dilator operculi (DO) niusclc is attached. In atliel.inopsids, ally. The abovc pattern of fi-ontal pores is present also in the LAP ~nl~sclcis attached to the anlCrovcntral kconly of the rnenidiines (Fig. 10C). A relatively derived pattern present in postorbital process. (O=postorbitalprocess small and narrow; Atherinopsinae (Fig. 1OD), notochei~-ids,and melanotacniids, l=postorbital process wide; 1.00) is pore five directed posteromedially rather than anterolatel-ally (Fig. IOA, 1013, 10C). (O=pore 5 directed anterolaterally; l=pore 19 Uurmo,s~)h~~zotirn?-~ic.~tlntiorr.- Tllc plesiornorphic condition 5 directed posteromedially; 0.50, 0.75) of athcriniforrns is f'or the dcrmosplienotic to be ;u-ticulatcd In mugilids (Fig. IOA), pore five is displaced anteriorly I-el;\- dorsally to the 1.1-ontaland along its entil-e posterior lcngtll with tive to atherinornorphs and the prcopcrculal- canal is co~inectcd the anterior face of the postorbital process. The derived condi- to pore eight rather than to pore seven as in atherinifol-ms (Fig. tion fi~undin atherinopsids is f'or the dcrmosplie~ioticto be 1013, IOC, IOU). connected dorsally lo tllc 1.1-ontalor f1-o11tal-sl~hrnotic,j11nction rostra1 cartilage \ interpremaxillary ligament premaxilla \ I

lateral ethmoid

Fig. 5. Dol.sal view 01' thc cthmoid region of Bnsilichtlt~~sssp. (UMMZ 215.520). Lcf't lacrimal and nasal bones removed.

Ily lihrous sliin only, detached lrom lhe postorbital proccss (Figs 20 Dermosph,enotic sensory canal.- The plesiomorphic condi- 8, 11; De la IHoz & Tosri-(:rote, 1981). In ml~gilidsthe tion present in Cyprinodontca and atherinoids is an open derrnosphenotic is also detached Si-om the postorbital process dermosphcnotic sensory canal which is not Sully ellclosed by and co~l~lecteddorsally to tlle frontal only by thickened skin. bone. An enclosed bony tube is present in mugilids, in (O=dermosphenoticarticulated to postorbital process and fron- menidiines except for , and in atherinopsines (Fig. tal; l=dermosphenotic not articulated to postorbital process, 12) except lor Alherinmpsis and Odontesthes perugiae. connected dorsally to frontal or sphenotic-frontal articulation; (O=dermosphenoticsensory canal open; l=dermosphenoticsen- 0.50) sory canal enclosed; 0.25) The menidiinc i\/lela~zorhi~nushas another derived co~ldition in which rhe derrnosphenotic is entirely articulated to the ante- 21 Dm?nosphenoticpore,~.- The derrnosphenotic sensory canal rior Sacc of the postorbital process, not to tl~cfrontal. is plesiomorphically co~lncctcdto the supraorbital canal dor- 14 MIX.PUHI.. MLJS.. zoo^.., UNIV.MICI~., No. 185

nasal nasal

Fig. 6. A11terior inSfi~orbitalsant1 nasal boncs; A, O(/o17,~cr/h1'c~clrof,r?~?ri~ (UMMZ 251 1150); I<, Wn~ilic.lt/l~j~APIIIOIIIIIA (LIMMZ 21 7989); (:, 13rrsiIic.hll/~1s rr~i.\l,nli,(UMMZ 215507). sally (Clr. 17: pore 5) and has a single pore ventl-ally (Figs 11, 23 lIer~~zo.s~~l~~r~o/i(.po.sloc.ulnr .sll~I/.- Prcse~lceof ;I II;II-rowme- 12B, 12C). A derived co~lditionu~lique to Atherinopsini is pres- dial flange along tlre internal klcc of the dcrmosplrcnotic is the ence of an additional pore (Fig. 12A). (O=dermosphenoticwith plcsiomorphic condition for alherinilbrms. A well developed two pores; l=dermosphenotic with three pores; 1.00) medial flange, or postocular shelf, at thc dorsal half oSt11c 11one is a derived condition of atlrel-inol~sincscxccpt for 13a.tilicli~tliy.s 22 Uerrnos~henotic~mudinlJnnge.- l'resencc of a mcdial flangc (Fig. 1 2A. 12C). (O=medial flange narrow; l=postocular shelf along the internal Face oi' the del-mosplicnotic bonc is a on dorsal half of dermosphenotic; P=postocular shelf along plesiomorphic condition for atlreri~lii'orms (Fig. 12A). entire length of dermosphenotic [Dyer, 1993: 3431; 0.40,0.63; Bn.silichthys, Menidiini, and Cyprinodontea have the derived con- add) dition of lacking a medial flange, except that in 1~asilick~l~y.cthe flange is redllccd to the extent 01' tlle tubular sensory nerve 24 71.ig(~nii.rrr~l,/or(~niir~(1.-Full enclosure of' tllc trigenlinal fil- canal (Fig. 12B). (O=medial flange present; l=medial flange ramina by the prootic is the plesiornorphic pattern for absent; 0.33, 0.50) atheri~lif'orrns(Figs 9, I.";). Tlle fora1nc.11li)r thc tripcmin;~lnelv

trigeminofacial exoccipital

lateral ethmoid

Baudelot's ligament ossification

Fig. 7. Vcntrolateral view of the ~~ertrocraniumof O(lonf/.tfl~c.co~ipntali\ (MNI-IN-M 2670) trigeminofacial arch supraoccipital pleurosphenoid parietal preorbital wing frontal 1 ,~ioccipitawing lateral ethmoid I \ ,,/ 1,

/ trigeminofacial parasphenoid foramina

Fig. 8. Ventrolatcral vicw of thc neurocraniurn of Orlonle.\th~.shatch,& (UMMZ 218450). Derrnosphenotic in dashed line.

(V) is derived in Odonlestke,~,with an enlarged anterodorsal open- men of the palatine or occulomotor nerve is also derivcd in ing enclosed by the prootic, pterosphcnoid, and sphenotic (Figs Odonleslhes in that it is not fully enclosed by the prootic as in 7, 8). (O=trigeminal foramen fully enclosed by the prootic; other atherinopsids (Fig. 9), but rather shared with the basisphe- l=trigeminal foramen enclosed by the prootic, sphenotic, and noid, or absent altogether (Figs 7, 8). pterosphenoid; 1.00) The irigeminal foramen is usually a single large foramen, 25 ??igerni.nojaciul arc.h or lateral commissure.- The bony arch though sometimes it is subdivided by thin ossification of the of the prootic, lateral to the trigeminofacial foramina or lateral membrane that separates the ,jugular vein from the commissure, is traversed by the jugular vein and orbital artery trigeininofacial cllambcr (Dc Beer, 1937: 121,128). The fora- (Goodrich, 1930: 273-278). Atheriniform outgroups have a

lateral ethmoid

ossification

or palatine nerve

Fig. 9. Venirolaieral virw of' Ihe ncurocrar~iu~noS Rasilichthys microlepidolus (UMMZ215499) and a medial view of anterior portion of the right sr~spensoriu~n. single arch lateral to the trigerriirioL~cialforamina. A derived (rneningosts of Chabanaud, 1936) arc in the horizoiltal plane, condition is found ill atllerirlopsids in which a secondal-y arch forming the anterior roof of the posterior inyodome. The ba- separates the orbital artely posteroventrally from the largcr pos- sisphenoid in mugilids is greatly reduced eitlier to presence of terior opening ofthe jugular vein and facial (VII) nerve (Fig. a loug and slender belophragm only or absence of both 8). The secondary arch is absent in Basilichthys (Fig. 9), belophragln and meningosts. The basisphenoid is absent in Alher~nopsis,and I,eurestWc?s; this arch is incomplete or weakly cyprinodontifornls and adrianichthyoids, but present and well developed in some specimens of Odontu,stlzu.s (Fig. 7). A unicluely developed in exocoetoids. Atherirlopsiiles and species of derived condition is found in all ~nenidiinesexcept I,aDldesthes (,'/zirostoma, have a derived condition in which the belophragm in which a wide bony tube encloses the jugular vein posterior to is very short ventrally and is mostly projected anteriorly along the trigerninofacial arch (Fig. 13). This forms a posteromrdial thc dorsal midline of the parasphenoid towards the interorbital 101-amell 1'01- the hyoniantlibular nerve and a posterodorsal fo- cartilage (Figs 7,8,9). The tneningosts are oriented diagonally ramen presumably for the opercularis nerve that innervates the in cross section, converging- at the midline close to the adductor and levator operculi muscles (Harrison, 1981). parasphenoid and separating the posterior inyodome eye (O=singletrigeminofacial arch; I=secondary trigeminofacial arch muscles. (O=basisphenoid belophragm long and slender; present; 2=bony enclosure of jugular vein posterior to I=belophragrn short projected anteriorly; 1.00) trigeminofacial arch; 0.50, 0.78; add) A derived but variable trait is found in Basilichl!~y.ssemotilus species group and in the Rio Quilimari population of B. 26 Rasisf~lt.en,oidDc~lop!rr(~~pr~.- The ventral, laminar prqjection microlepiclotus: the anteroventral tip of' the belophragm is cx- of the basispher~oid(bclophragni of Chabanaud, 1936) in tended anteriorly and is in contact with the posterior end of atheriniforrn outgroups divides the anterior opening of the pos- the interorbital cartilage (Fig. 11). Presence or absence of this terior myodorne and is relatively long and slender (Dyer & feature is independent of size or sex, found consistently ollly in Chernoff, 1996). 'The dorsal rarni of tlie basisphenoid the Rio Loa population of R. s~rnolilusspecies grollp.

Fig. 10. Uo~.sillview of' tlie lert supraorbital and rerripolxl sensory canals and porcs; A, Ag(~r~o.\lomu111on/7colrc (UMMZ 199463); U, I-ly/~orc~lz/:rir~tz ble(!li(.rj(FMNI-I 57459); (:, M(.nirlin 6rryllincr (UMMZ 203149); D, Alh(,rPnop~ a/Pni~(UMMZ 7'2212). hyomandlbula mesethmoid lacrimal dermos~henotic /

metapterygoid

Fig. 11. Latcr;il vicw oT tllr hciid and pectoral girdle oS Ba.silicllll~y~sp. (UMMZ 215519).

27 l'(tr(t\j)h~~io~d71~1i/r(tl rtdgr- (O=anterior end of epaxial muscles that extend into the temporal fossa. (O=no parasphenoid ventral ridge, smoothly tapered; l=anterior end pterotic wing; l=pterotic wing present; 0.50) of parasphenoid ventral ridge, rounded; 0.50) 30 b,'xoccipital wing (Chernoff, 198613) .- The plesiomorphic 28 LVu$rnoccil,ii~~lcr(?st (Whitc, 1985).- Thc plesiomorphic con- athcriniform condition is for a smooth dorsal face of the dition founcl in mngilids, atlierinoids, and rnenidiines is a single exoccipital condyle. Atherinopsids, except for I,abidesthes and vertical 1a1nin:u- crest projecting posteriorly SI-om the supraoc- , have the derived cosldition of a laterodorsal lamina cipital 11o11e into the vertical septuni (Fig. 14A). In projecting from the exoccipital process in the form of hooks or (:yl.winodosllea, two postcrodorsal laminar projections form a a perforated flange. In Bnsilichthys a hook or spine is present V-shapcd supraoccipital crest that reach the base of the supraoc- (Fig. 9), whereas in all other atherinopsines examined the cipi~albone. Atherinopsini and 13asilichihj~.salso have bifid su- exoccipital presents a flange of va~yingdegrees of development, pr:~uccipitalcrests, but arc projected instead fs-om a single, basal posterior to the attachment of the postemporal ventral process laminar crest (Fig. 146, 14C). In atherinopsinins, the supraoc- (Figs 7, 8). Chernoff (1986b) proposed this feature as charac- cipilal crest appears trifid in dorsal view (White, 1985) because teristic of Menidiinae, but its presence in all atherinopsines sug- tllc gostcrio~-bo~.cler ol'thc single basal lamina projects posteri- gests it is characteristic at the more general level of orly from t11c point of bifurcation (Fig. 14C). (O=singIesupraoc- Atherinopsidae. Wings or hooks are present in atherinopsines cipital crest; l=bifid supraoccipital crest; 2=trifid-like supraoc- larger than 100 mm SI,. (O=exoccipitalwithout a wing or pro- cipital crest; 0.50, 0.80; add) cesses; l=exoccipital with a wing or processes; 1.00)

29 l'i(:rolic. ruing.- The postcrolateral border of the pterotic 31 Ptmotic exoccipital wing.s The pterotic wing is a diagon;~lridge that is continued vcntl-ally by the exoccipital, of ml,gilids, menidiines, ~~~ili~ht/~~~(ch. 29) tapers with or witllollt an intcrcalar bone. Presence of a wide laminar trally without reaching the exoccipital wislg (Ch, 30). In flange St-om tllc posterior ridge of thc pterotic is a derived Lea- atllerinopsinins and ~d~)~t~~th~~the pterotic wing is contiguous ture of' m~~gilidsand atherinopsids (Figs 7, 8, 9; Starks, 1899). with the exoccipital wing (Figs 7, 8). (O=pterotic wing not con- 'The 11te1-otic wing receives ~nuscleattachsncnts from the tipous with exoccipital wing; l=pterotic wing contiPous with clcithrum and opercle and Sorrns the lateral enclosure of the exoccipital wing; 0.50, 0.80) Mlsc.. PLIBL.MIIS.. zoo^.., UNIV.MICH., No. 185

32 I'//.r.o/ic. ct,ncl (.f,ioc.c.ifii/trlroi,~t:s c.on/i~~~ous.- The pterotic \ving postr~ril~oralcanal bones ill atheriiiopsids is problematic (Dyer ill irtlicrinopsitl orltg~-or~l~stapers ventrally only to~vai-tlsthe & C1re1-noll;1996). Tlle postempol-211iri Merlidiiilac has a bony exoccipital. B(~.\ilic.lrrlt~~.ls~andAfk/+rt~ro/).\i~ llavc all additional 1101-i- sensory canal wit11 t111.c~pores (Figs IOC, 15A), and zo~llalflange, ventral to t11c clxtxial nlusculat~~rcor tl~etc11111o- Ahel-inopsinae lias a postemporal c;~~lalwith two pores (Figs ral ross;~,that is contiguous with n basal \videning of the 10D, 1I, 15B, 15C). Mc~lidiineshave no extrascapt~larbones cl~ioccipitalwills. (O=pterotic wing not contiguous with (Fig. 10(:) whereas atllcri~lopsineshave an extrascapular bone epioccipital; l=pterotic wing contiguous with epioccipital wing; with three pores (Fig. 1011, 11) that is soinetirnes divided in two 0.50, 0.67) separate eleinrnts (Fig. 11) (see Iliscussion). Based on morpho- logical siinilarity of the canal bones propel-, regardless of the 3 3 I~r~sioccil,ilnl,/i?~~~~~,sl~(~tion(Wllitc., 1'385) .- 'The posterioi- rc- undcrlyi~lgbone to which they may attach, the postemporal ca- gion 01' the basioccipital is snlootll ill athc~.i~lopsidoutgroups. nal bone of rnenidiines and the extrascapular oPat11erinopsincs A derivcd contlitiorl fbuncl in sorgrntinins is the k~icstratio~rof meet all cipriori tests of si~llilarity(Patterso~l, 1982) and are con- thc 11;lsioccipital bone (Figs 7, 8,9;M'hite, 1985: fig. 12). (O=ba- sidered to be potential hoinologucs. (O=three- and two-pore sioccipital bone smooth; l=hasioccipital fenestrated; 1.00) canal bones present, not fused to underlying hone; l=two-pore canal bone absent, three-pore canal bone fused to postemporal; 34 IItr~id~lol'slig.c~nr(,nt oss~fic.cr/io~t (Whitc, 1985).- An 2=two-pore canal bone fused to postemporal, three-pore canal ~unossilicdB;-~r~tlclot's ligament is the plesion~o~-plliccontlition hone(s) not fused to underlying bone; 1.00; n-add) for ;rthcriiniSorms. A derivccl coildition found in belonifoi.nls (except Sol- exococtitls) and atlre~.inol~sinesis ossification of the 36 Po\/(~n~pornlclnt~riol- pl-ocess ((:llcrnoff, 1986b).- The ligainc~ltat its attacli~nentto the Ix~sioccipital(Figs 7, 8,9;Starlis, plesiomorphic conditioir of at1neri1lifo1-rnsis for the postemporal I:fig. I,) (O=Baudelot's ligament not ossified; to have two processes; a dorsal process that articulates with the l=Baudelot's ligament ossified proximally; 1.00) cpioccipital and a veiltral process that ai-ticulates with the B;~udelot'sliganle~~t is attached to the exoccipital ill intercalar or exoccipital. Tllc derived condition ofatllei-inopsids i\/oln~tolhitc~itsant1 is itbsent in notoclicirids. is presence of an anterior spine-like process of variable length projected into the temporal fossa (Fig. 15). A f~u-therderived 15 I

trigeminofacial arch bony tube

Fig. 13. I.rf't vcntrolatr~-alvirw of thr otir and occipital I-cgionsof thc ncul-ocraniu~nof Mrrnhj-os g111)o-ti(UMMZ 202405). IX: glossopharyngcal nerve; X: vagus ncrvc; PI.1.: posterior latcr;tl linc ncl-vc.

dorsal dorsal dorsal

lateral lateral lateral

Fig. 14. Scllcr~~atictlorsal allil lateral vie\vs of tll' srrpraoccipital crest; A, O~(III~(,A~/L(J.\;B, Bn~ilic-htlzjs; C, Alh(~i17ops process

fdorsal lateral

Fig. 15. Dorsal and latcral vicws of thc left posternpofill I~onc;A, Mw~idicl;U, Odo~rl~~.\l/rc~.\hrc~~io~rrrlis (1JMMZ 215150); C, l~osilirhllzysrr~r,\lralic (UMMZ 215508). DYER,ATI-IERINOPSINE RELATIONSFIIPS AND BASILICHTHYS 21

nar (Figs 10D, 15C). (O=no postemporal anterior process; maxilla to have a relatively narrow base, displaced laterally from l=postemporal anterior process present, spine-like; the midline (Fig. 4). This co~lditio~lis found in menidiines Z=postemporal anterior process wide; 0.40, 0.73; add) (except for Labide.sthes and a species group of the subgenus Eury.sto1: Chernoff, 1986b), Leuresthes, and 0dontestke.s (Figs 17, Jaws and associated ligaments 18; Schultz, 1948;White, 1985; De la Hoz el al., 1994). (O=base of premaxillary ascending process wide and close to symphysis; 37 Uj"1)erlrcro/rc~trurn.- Presence of a fold of skin between the l=base of premaxillary ascending process narrow and displaced dorsal processes of the maxillae and the premaxillary ascend- from the symphysis; 0.33, 0.78) ing processes is isa plesiomorphic condition for atheriniforms. In atherinoids, the ascending process has a wide base and is Presence of a continuous, unlolded sheath of skin between the elongated posterodorsally in several groups (Saeed et al., 1994; 11pper.jawsand forehead, or frenum, is a derived feature shared c.g., phallostethids, Slenatherina, Craterocephalus, and by the athcrinopsine genera Ath(trinof~sis,Atl.rer.inofis, Colfiichthys, Kmtrathen'?~,a). and 1,'crsilichlhy.s (Fig. 16). (O=no upper jaw frenum; l=upper jaw frenum present; 0.50,0.60 / 0.67, 0.80) 39 Premaxillary antmiorjoin1 (Patten, 1978).- A relatively wide ligamentous articulation between the anterior ends of both pre- 38 I'wrnc~xill~~rynscen,din,g process (dorsal process of White, maxillae is the plesiomorphic conditio~lfor atheriniforms. The 1985).- The pre~naxillaryascellding process with a wide base derived condition found in all atherinopsids is a narrow lip- close to lhc symphysis is the plesiomorphic condition for mentous articulation between both premaxillae (Figs 4, 5; De atherinilorms (Fig. 5; Sch~~ltz,1948: plates 1,2). A derived con- la Hoz &Vial, 1988). This condition is most noticeable in spe- dition of atherinopsids is for the ascending 131-occssof the pre- cies with narrow-based ascending processes (Ch. 38). (Opre-

Fig. l(i. Dorsal vicw ol'llrc head of Un.s~lic.l~ll~~~.\;A, R. rnicru1ef)idotus (UMWIZ 215517);B, U. sernotil~~c(UMMZ 218050). maxillary anteriorjoint wide; l=premaxillaryanteriorjoint nar- 43 Postmaxillary procrss of thp pr~mnxilla (Patten, 1978; row; 1.00) Chernoff; 1986b).- A postmaxillary process on the alveolar arm of the premaxilla, projected medial to the maxilla is 40 Dorsnl process of maxillary head.- A well developed dorsal plesiomorphic for atheriniforms, present in mugilids except process of the maxillary head is the plesiomorphic condition Mugil, and atherinoids except Notorh~iru.~.Absence of this fea- for atherinomorphs (Figs 17, 18). A reduced dorsal process is ture (Figs 17-20) is considered derived and was regarded as di- considered a derived condition present in Cyprinodontea, agnostic of atherinopsids by Chcrnoff (198613). This ficature is Basilichtl~y.ys,Atherinops, Ath(.n'nopsis, and Co1pichthy.c (Figs 19, 20, ambiguously optimized however, because it is also abscnt in 21). (O=dorsal process of maxilla developed; l=dorsal process Notocheirus (Rosen, 1964: fig. 4A) and Cyprinodontea. of maxilla reduced; 0.25, 0.50 / 0.33, 0.67) (O=postmaxillary process present; l=postmaxillary process ab- sent; 0.50) 41 Ventral process of maxillary head.- The plesiomorphic con- dition for atheriniforms is for the ventral process of the maxilla 44 Premaxillary alveolar arm (Chernoff, 1986b).- Thc to be longer than the dorsal process, projecting anteriorly and plesiomorphic condition or the premaxillary alveolar arm in medially. A derived condition found in Odontesthi~sis for the atheriniforms is a pointed distal end. The distal end of the dorsal and ventral processes to be of roughly the same length premaxilla is variably broadened in some Atherinoidei groups (Figs 17, 18). Asimilar condition is found in some membradins such as Allze~ion,Pseudomug-il, Notoch,rirus, and G.atet-ocef11zalu.s (Chernoff, 1986a,b). (O=ventralprocess of maxilla longer than (Patten, 1978). A derived condition ofthe preinaxilla is found dorsal process; l=dorsal and ventral processes of maxilla about in atherinopsids in which the distal cnd is notably dilated (ovcr equal in length; 1.00) twice the width ofthe proximal rcgion of the alveolar arm) with a rnostly convex posteroventral border (Figs 17, 18; Chernoff, 42 C;onr%yleon thp ventral pmress of the maxilla ("rounded pro- 1986b). Reduction of the dilation is found in some rnembradiils ccss", White 1985).- The ventral process of the maxilla is usu- (Chernoff, 1986a) and a concave distal bordcr is present in ally flat or thickened in its entire length and articulates dorsally L(>uresthes(Schultz, 1948: fig. 4D). (O=premaxillaryalveolar arm with the asccnding process of the premaxilla. Preseilcc of a pointed or slightly broad; l=premaxillary alveolar arm broadly rounded thiclieiling that functions as a condyle on the dorsal dilated; 1.00) surface of the maxillary ventral process is unique to Bnsilichthys (Fig. 21). This feature is absent in all other atheriniforms ex- 45 Premccxilla- dent(^ coronoid lig(~nzn7t(Patten, 1978).- A rela- amined. (O=no condyle on maxillary ventral process; l=condyle tively long ligament connecting the coronoid process of the present on maxillary ventral process; 1.00) dentary with the acljoined tips of the upper jaw bones is the

hyomandibula A' ch' 68

Fig. 17. I.;~teralview of the lefijaws and suspensorium of Odontesthe,~wtrojjrnn,is (IJMMZ 231950) hyomandibula \ Chs 65.66 Ch. 68

Fig, 18. Mctlial view of the left.jaws and suspellsorir~mof Odontrctl~r~cwtl-opinnis (UMMZ 231950)

~lcsiornorphiccontlitioil Sor atllei-iniforms. A derived condi- a long ligament between the distal end of the maxilla and the tion ofatherinopsitls is for the ligamentous connection between lateral face of the premaxilla (Fig. 22), allowing a greater amount the ~x)stw(~-dorsalcorner of the de~itarycoronoid and the pos- of relative movenlcnt between tlie upperjaw bones. The Senel-- terior end of tlie preniaxilla~yalveolar arrn to be extremely short alized atherinopsid condition is to have the ligament attached (Fig. 22). The liga~ncntis so short that movclncllt between the to the mid-region of the premaxillary alveolar arm. Derived two bones is restricted to rotation relative to each other. This states of the ligament's prernaxillary attachment are: to tlie :tllows the lowering mandible to pull the premaxillae forward a~lteroventralcorner of the alveolar arm in Labide.rl/~~s, and down during opening of the mouth (De la Hoz & Vial, Colf)ich.rhys, and Hasilichthys (Fig. 22; De la Hoz & Aldunate, 1994; 1988; 1994; Dc: la I-Ioz, 1994). (O=premaxilla-dentarycoronoid De la Hoz el nl., 1994: figs 18-20); to the mid-ventral region in ligament long; l=premaxilla-dentarycoronoid very short; 1.00) Atherin.ella and some species of Chirostom,~;and to the dorsal A shor~ligament between the premaxilla and coronoid pro- border of the alveolar dilation in Leurrsthr~s,anterior to the at- ccss of the dencai-y has derived independently in Psc:udomt~gil tachment of the premaxilla-dentary coronoid ligament (Ch. 45). ;lntl Q.~/[.r~cef)/~,(~lzts(Dyer & Chernoff, 1996),thougll not as short iU1 other atherinopsids have the ligamentous connection LO the :Is in atherinopsitls. lateral face of the alveolar arm of the premaxilla at positions intermediate between those of (;olf)ichthysand Leurest/~es.(O=max- 46 il/l~~xilla-a,t~~uIo(~~'~i(~ulnl.corcnoid ligarn,en,l.- The maxilla and illa-premaxilla ligament very short; l=maxilla-premaxillaliga- ~x~ui~;~xill:~arc closely adjoined at their tlistal tips in atheriniforrn ment elongated, attached to mid-region of premaxillary alveo- o~~(groupsand are connected only to the coronoid pi-ocess of lar arm; 2=maxilla-premaxillaligament elongated, attached to the cicntal-y by a long liga~nent(see Ch. 45). A derived condi- anteroventral corner of premaxillary alveolar arm; 0.50, 0.80; ion prcsci~tin Menidiinae, Athcrinopsini, and Odonlrrlhrr is add) prcwncc of a ligamentous attacllrnent from the rriaxilla to thc co~o~loid process of the angt~loarticular(Fig 22). (O=no liga- 48 I,abial Izgc~I~lent.fold (Chernoff, 198613) .- The labial liga- ment between maxilla and andoarticular- coronoid; l=lig.ament- ment connects the mandibular lip with the distal or posterolat- between maxilla and anguloarticular coronoid present; 0.50, era1 end of the upper jaw. This ligament unfolds as the man- 0.83) dible is lowered and the upper,jaw is displaced anteriorly. The plcsiornorphic atherinomorph condition is for the labial liga- 47 iM(~xilln-j,r~~~~inxillnrlisinl lipm~nent (White, 1985).- The dis- ment to be folded between the mandible and the premaxilla cal c~ltlsof the ~naxill:~and prernaxilla arc co~l~lectedby a short (Jordan & I-lubbs, 1919: figs 1-10, 41, 42). A derived condition liga~llcntin athcrinifor~noutgroups allowing a limited move- is found in cyprinodontiforms, menidiines, and sorgentinins, ment bctrveen them. The derived condition ofatherinopsids is in which the labial ligament is folded on the lateral face of the hyomandibula opercular \ dorsal process

Y retroarticul;r quadrate \ preopercle interopercle symplectic

Fig. 19. L.aiel.al view ol'lhr Idt,jalvs, i~lli-;lo~-bit;lls,;~nd suspensoriunl of Un,\ilic.ltlh~.\cru\ll-c~li\ (IJMMZ 2 15507). pretnaxilla (Fig. 22; .Jordan & I-~L~IIII~,1919: ligs 27-35; Schrllt~, fig. 2; White, 1985: fig. 22). Kicuspitl tccth arc :tlso pl-cscnt in 1948: figs 8, 9). In ~nenidiinesand O(lol7,lalhr.s this fold is hroad posterior rows of mngilids and solnc derived cyprinoclontoids. and ovcrlaps the r31-cmaxilla (Fig. 23A). The Brc~ilic-hlhp~ (O=unicuspid teeth; l=bicuspid teeth; 0.67, 0.00 / 1.00) mic.rol(~idoltr,sspecies group has a greatly reduced overlap (Fig. ZSU), \vhereas in the U. semolibts specics gror~ptllere is 110 up- 50 Toolh rn7u.s 071 ora1jaws.- Plcsiomorphic for atheriiliforms tr~rnedfold and a much narrower labial liga~ncrrt(Fig. 'LX:). is presence of rnorr tlran thl-ce rows of tcctl-I on oral jaws as (O=labial ligament folded internal to premaxilla; l=labial liga- seen in mugilids, cypriilodoiltifor~lls,bclonifo~.ins, and ment folded lateral to premaxilla; 2=labial ligament with lateral ather-inoids cxccpt fhr notocheirids which have iwo rows only. fold greatly reduced; 3=labial ligament lateral but not folded; A similar condition occurs in meinbradins in which Mp?rtDras 0.75,0.90; add) and Alheri~i~llnlravc thc plesiomorphic contlition, but Tlle only rekrences I was able to find on this striking feature Mrla~zor11,inushas only two rows. Two states are for~rltlin are Gosline (1963: 24) and Rose11 (1964: 233). Gosline noted Atherinopsini: (i) Colpich,tli,y,ysand Ath~~inofishave two rows of that cyprinodontilhr~~rsand ainblyopsicls have the lower lip folded teeth; and (ii) Athrri~~o~~szsandL~urt.slhr.s have tlircc rows. Atl~llt over the upper,jaw, a "curious reversal of' the usual ripper lip sorgentinins have three rows of' teeth except fox- 0. katclroriand configuration [which] does not occllr elsewhere anrong the the Bn,silichth~~s,srmotilu.sspeciesgl-oup that have inor-c than tlrree prcpercoid grorrps". Rosen reportetl the presence of a lateral vows (De la I-Ioz ~t nl., 1994: iig. 6),and Odon~estl~espmugiar,0. fold in all cyprinodontifo~-ms,sonic cxococtoicls, and all rrtropinnis, and the srrbgenus Au~tromc~nidiathat have two rows atheriniforms except Brrsilic.ht/~~lsnrt,strcrIi.s. Because Goslinc's only of teeth. Modification of oral jaw teeth in 0. incisa rcndc~-an atherinif'orm was Alh(.rin,ops aJJCt7isand Rose11 did not iriclr~clean ambiguoris coding and is discussetl in Dyer (1993: 346). atherinopsinin spccies in liis compa~-ativematerial, they were (O=more than three rows of teeth on jaws in adults; l=three led to opposing gcncrali~ationsallout the condition in rows of teeth on jaws in adults; 2=two rows of teeth on jaws in atheriniforms. adults; ?=polymorphic; 0.29, 0.58; n-add) (;olf,icht/~~l.sand Atherino/)s have been described as having a 49 Cusps on ot-ol-jnru krlh (White, 1985).- Presence of unicrls- single row of bifid teeth (Jordan & Hubbs, 1919; Crabtree, pid teeth is the plesio~norphiccondition for atherinifornrs. Pres- 1989). Only one row of teeth is visible, how eve^; because the ence of bicuspid teeth in f\thrrinop

coronomeckelia anguloarticular coronoid retroarticular ~~m~lectic preobercle

Fig. 'LO. blctlial view of tl~c1cCt~a.i~~ and susprnsorium of 1ln.silichtlz~~snz~~t~nlis (UMMZ 215507)

dorsal

Fig. 21. Left iiiaxilla in postel-io~;anteriol; and dorsal views of Ua.silichtl~ys(CAS ,151'37) Mrsc. PCILIL.MCIS.. ZOOL., UNIV. MICII., NO. 185 dorsal rostral

ligamentI palatorostral ligament lateral rostral ethmomaxillary ligament ligament, I,

A1 lacrimal branch articular-maxilla

51 l)~rr/nr:)lcornnoid pr.oc.cc.c.- Fl'l~ccol-onoitl p~.occssof tllr 53 M~mcliOrrkc~rOr-c~irclt of A 1 ~trr~~cleio~rdotr.- The A1 division dcntary is usually 1.o11ndetl ;111tl cstcndrtl dol-sally ill of the adductor ~nandibulac11111scle ofat11e1-i~~opsids is actached athr~-i~liPo~-~ris,well Ijryond the co~.o~loitiprocess 01 the a11te1-iorlyby 21 te~ldo~lto tlir medial fkcc of tlic vr~itralIialP of' i~~lguloac-ticula~:Even w11er1 tllc tlcnci~~.ycor-orloitl is rlot so ~vell tl~c~naxilla, ant1 a long dorsal bl-anch of the XI tendo~lto thc clevclopctl tlorsally (r.g., Lnbitlo~iIro.c),tllc a~iguloa~-tic~~l;~~.coro- s111jnasal slielP of' tl~rlacrimal (Dyer & (~:licrnoff', 1996). A dc- lloitl is IISII;III~ rrtl~~cctlLO ;I reli~tivrlyshort spi~lc(Figs 17, 18). rived corlditio~lfot~ntl o111y ill atlierinifornis is ;I velltl.al or man- I~ctsilic.lr/l~?shas a dc~.ivcdconditio~l ill wllirli the tlorsal bol-dcr dibular bra~lcllatcacl~etl ve~ltr-ally to the anguloartic~~lar,ante- of the tlcntary coro~loidprocess is flattened ant1 ;it tllc same rior to the a1-tic11lation\\lit11 the quadratc (Fig. 22; De la Iloz & lcvcl ol ;I tlorsally enlarged ;~~ig~~loartie~~li~~co~.o~loicl(Figs 11, Aldl~nate,1'394; Dc la Hoz ccl., 1994). In atherinoids, tllc nlan- 20, 25). A similar co~lditio~lis forrnd ill pseudoniugilins (Saeed tlibr~larhrilllch is fou~ltlill A//~c:rio,rr,~nelanotacniines, ('1 (ti., 1080). (O=dentary coronoid process rounded and taller 7i~ln~nllrerii~n,iultl athel-inines (Dyer & Chcr-noll', 1996: fig. 8). than anguloarticular coronoid; l=dentary coronoid process flat- In athcrinopsids, the inandiljrllar braiicll is f'o~lntlin menitliiiles, tened dorsally and at level with tip of angdoarticular coronoid; I,PIO.(~.Y~~LP.~,a~iclO(~otrlo.r/ltc~.s (Fig. 26). In mcnidiines tllc mall- 1.00) dib111ar tc~ltlo~ibranches postcrio~.to the lacrimal h~-a~lcllas in ;~tlierinoitls(Fig. 26A) . In Lol~resllre~s;and O~lo,~/c?slhcs,liowevel; 52 I)rrr/cr.ryprtrilric. .s/)i~r,r.-Tlrc postct.oven1~11regior~ of the the rnandibrrlar terldoll 111.i11rchesanteriol- to tllc l;acrirnalb~-ancli de~lt;l~.y;1rcic111;1tes ~llctlially will1 illr antc~-io~.p~.occss of thc (Figs 22, 26B). (O=noAl mandibular branch; l=Al mandibular ;III~II~~;I~~~~II~LII-.Tlle posterior erltl 01' the ctc~ltarysensory ca- branch present, posterior to lacrimal tendon; 2=Al mandibular 11;1l lxojrcts postcriol.ly tow;~rdsthe a~lguloai-ticrrlarin a g~latllic branch present, anterior to lacrimal branch; 0.75,0.90; n-add) ramus or spine, britlgi~rgthe gap I)ecwee~lboth bones (Figs 17, 18). A drrived co~rclitio~ifound in Bn.silichil~y~is absr~lceoP the g~latl~icspine (Figs 11, 19, 20, 21,25; Dr la I-Ioz & 'Ihsti-CI-oce, 1981 : lig. 6). (O=gnatliicspine present; l=smathic spine absent; 5.t I'nln/irr~ c1ot:cccl /r~-oc.c

subocular scale rows

subocular scale rows

subocular scale rows

Fig. 23. I.;~tc~-alvicw oS thc s11o11rI-cgion; A, 0rlontc.sth~cf)rr~~@n? (IJMMZ 221.328); B, Nns~licl~ll~~s~r~ic~olrid~i (UMML 21 5514); C, Nnsilirhfhys c~r~1o~i11c.s(IJMMZ 21 7957). atherinomorph i'eature; its absence is diagnostic of Atherinoidei intermediate palatine attachment; ?=polymorphic;0.50,0.80) (Dyer & ChernofI; 1996). A derived condition for atherinopsids A uniquely derived condition fo~uirdin Colpiclzlhys ,rep,ris pres- is presence oPa dorsal ridge that connccts thc anterior and dor- ence of a ring ligament attached to tlie palatine dorsal process sal processes of the palatine (Fig. 27A). A f~~rtherderivcd con- forrning a loop through which the ethmomaxillary ligament dition olatherinopsines is the development of' a shell'medial to traverses with no attachrneilt to the palatine. the dorsal ridge such that the dorsal process appears folded over the ethmoid cartilage block (Chernofr, 198613) and the 56 Pal(~lol-oslrcllligcl.nz(~,nt.- Presence of a strong palato-pre- dorsal process is pointed distally (Fig. 27B). In Basilicklhys the maxillary ligament between the base ofthe anterior process oP dorsal crest is tall, rounded, and spoon shaped, lacking thc dor- the palatine and the posterodorsal end of the premaxillary as- sal ridge and lateral protuberances associated with the cending p~-ocessesis a plesiornol-phic condition for ethmomaxillary ligament attachment (Fig. 27C). (O=dorsalpro- atherinomorphs. This ligament extends dorsal to the cess present; l=dorsal process with dorsal ridge; 2=dorsal pro- ethmomaxillary ligament forming the so-called crossed lip- cess pointed and folded over ethmoid block; 3=dorsal process rneilts (Schaeffer & Rosen, 1961; Koscn, 1964; Stiassny, 1986). spoon shaped and folded over ethmoid block; 0.75,0.86; add) Crossed ligaments are absent in Cypri~lodontea(Parenti, 1981 ) but, contrary to Roseii (1964), well developed crossed rostral 55 I

Fig. 24. Lateral view of'the lelt jaws, inlraorhirals and st~spcnsoriumof Hnstbclr~lryssrmolilris (UMMZ 21 7089) dermosphenotic hyomandibula

cartilage coronomeckelian COrOnOid

Fig. 25. Medial \riel\, 01' thc lef(jn~vsand suspensorium of Uasilichtl~~~scrmolilus (UMMZ 217989)

57 I'/L~/L~~I~Pf1o~tero71~nlral s/)it~(?.- The posteroventral projec- position of the tooth patches, however, is variable. Within tion of' the palatine extended along the anterior facc of' the menidiines, teeth are present only in Membms and Athem'nell/~. cartilaginous palatoqrradrate arch (Fig. 27), but short of' the ChcrnoK (1986a) discussed the variable nature of these teeth quadrate bone (Figs 20,25),is reprded as a plesioniorphic ka- in the Athminelln cryslnllina complex and attributes this variabil- tllre li)r ;~thcrinomorphs.A derived feature is the extension oC ity to interpopulational gene flow. In Atherinopsinae, this postcroventral spine to contact the quadrate (Figs 17, 18). endoptcrygoid teeth are present in most species as well devel- Tllc derivetl condition is present in beloniforms, menidiines, oped or reduced tooth patches. Tooth patches are consistently A/Ic~~ri,~io/)sis,I,c~uresthe.s, and Odont~~,slh,(?sspecies except for 0. absent in Basilickthys (Figs 20, 25), Odo~ztestheshnlcheri, 0. Ir~rnr(~nsi.s.The condition in 0. peru~in~speciesgroup is ambigrl- humensis, and the subgenus Cnuque. (O=endopterygoid teeth ous. (O=palatine spine not contacting quadrate; l=palatine spine present; l=endopterygoid teeth absent; 0.20, 0.43) extended to quadrate; ?=polymorphic;0.25,0.63) 60 Hyonzandibulnr neme.- The plesiomorphic condition for 58 13clof1l~r.ygoid.- The ectoptcrygoid in mugilids and atherinomorphs is for the hyomandibular nerve to penetrate orrtgro~psisa relativcly wide bone folded around the anterior the hyom:~ndibula dorsomcdially and ventrolatcrally. In 1:.'Ice of'the palatoqrraclrate cartilage, dorsal to the quadrate and atheriniforms, the hyomandibular nerve divides into hyoid and ventral to the palatine spine. In athcrinopsids, the ectopterygoid mandibular branches external to the hyomandibular bone, usu- is a laminar bone on the medial facc of the quadrate and ally at the ventral [oramen (Fig. 19). The hyoid nerve branches palatoquadrate cartilage (Figs 18, 20). Absence of this laminar posteriorly and either penetrates the hyolnandibula again or I)onc Li-on1 the medial sidc of' tllr srlspensorium is a clerivecl lies between the preopercle and hyomandibula to exit condition observed in Un.silic.hll~y.sand0. nigricans (Fig. 25). Ab- posteromedially (Dyer & Chernoff, 1996: fig. 15.4). A derived sence ol' this fcaturc is variable in the B. microlepidolus species condition is present in I,eurestlzes, membradins, Alherion, gronl~;uld codccl as polymorphic in the data matrix (Table 3). Notoeheirus, and O(lonlesc.the.sspecicsexcept for 0. hatchmi, in which 111 the 8. ~nzicrolef~idolu,~spccies group 54% of the 50 specimens the hyomandibular nerve is divided inside the hyomandibula cxa~ninccllacked the ectopterygoid on both sides, 20% lacked (Figs 17, 18). Another derived condition is presence of' a dor- it on one side only, and 26% had it on both sides. sal f'oramen perforating the hyomandibula dorsally in mugilids, (O=eciopterygoidpresent; l=ectopterygoid absent; 0.60,0.33) atherinomorines, Iso, and Co&icAthys. (O=hyomandibularnerve divided external to hyomandibula or at ventral foramen; 59 k,'n(/o/)tcrggaid lc.c.lh.- Presence of' teeth on the ventrolat- l=hyomandibularnerve divided inside of hyomandibula; 2=dor- er;ll surhcc of' tlic endoptcrygoid is considered plcsiomorphic sal foramen of hyomandibular nerve perforates hyomandibula; in ;~thcriniSorms,though absent in Cyprinodontea. Size and ?=polymorphic; 0.40, 0.67; n-add) M~sc-.PUBL. Mu\.. ZOOL.,UNIV MICI-I.. No. 185

premaxilla maxilla I A lacrimal I A1 lacrimal branch

' / A1 tendon /

I dentary / branch / retroatticular anguloarticular

rostra1 cartilage \ palatine

A1 mandibular branch

I:iz. 26. Di~rgl-aniill 1ater;rl view of rlrc i\l tc~rcio~~colnplcs of tlic ;rtlductol- m~untlil)ulacrnr~stlr; A, (:hi~-o,\/o~~rri/r~bcr,rcic (LlMh~l% 105.163); B, O~/OJI/(,.\//II'\~(J/IO,/IIIII~~,S (LIiMblL 231 950).

6 I I'~.po/j(,r.c,~,[nr. .spj,.soty c.cr 11 (~1.- r plesi omol-pllic 198(ib) . M(,l(rnorI~it~~i~,Poblo~,c~, I,pci t.(,.s///(~.s,0'0l/~i~I11/~j~~, irlld ;\tllcrini[ol.ln condition is open preoperclll;lr sellsoly canal, Sol-gcntilli~riliavc both the horizontal and vertical shalts ol'the :IS li)rl~ldin (:yprinodontea, alhcrinoids, ;lnd rrrlrgilids (Fig. 28A, preopercular canal enclosed by bo~lc,tllougll ;I holly bridge is 28H). 'Vhc p~-copercl~l;lrsensory canal of the "el-tical shalt is us~tallylacking bctwccn pores one ;und two (Fig. 17), and be- clrcloscd lly bone between pores two and t'olrr in mrnidiines tween pores seven ant1 cight (Fig. I!)). (O=preopercular sen- (Fig. 28(:), Alh~ri7zol),s(Fig. 28D), and Alh(.r.inof)~is((:hcrnoff, sory canal open; l=vertical shaft of preopercular sellsory canal . . \. dorsal ridge / ./ Lw . .

Ch. 57 antenor lateral process Ch. 57: a1era \\ process lateral 7 posteroventral Ch. 57: spine posteroventral ,c-,. 54 dorsal ,Ch. 54 rl. =A n-,-r.

. . . - -. .-. anterior - 'anterior process \ anterior medial process dorsal process ridge Ch. 57 \ - Ch. 57 Ch. 54 Ch. 57 \

...... dorsal dorsal dorsal anterior process Ch. 54

I'ig. 27. I.atclxl, rnetlial, ant1 tlors;tl vic~vsof' thc lcfi palatine; A, il41,nirlin hrrj~llinc~(UMMZ 107098); R, Athrrll~oj~~crJJini.5 (UMMZ 232001); C, N(c.\ilic.lt//t~~.~n~t,\(r(rli, (UMMZ 2 15508). enclosed by bone; 2=vertical &horizontal shafts of preopercular 64 Opcrculr~rdur.ra1procc.s.s.- I'resence of a blade-like laminar sensory canal enclosed by bone; 0.50,0.80; add) flange on the lateral face oS the dorsal process of the opercle is A greatly enlarged sensory canal systclrl of the head is diag- a plesiornorphic feature of atherinilhrms (Fig. 29). The dilata- nostic oS tlic subgenus Cnuqup. Chiro.slomn Irzbarccze has the verti- tor opcrculi muscle is attached to the medial face of this lami- cal s1l;irt li~llyenclosccl by bone with a bony c~lclosurebe- nar fla~lgc.\~hich is extended posterovcntrally on to the lateral tween I,OI-~SSOUI-and livc on the horizontal sl!afi. Sate of the opcrclc. A derived condition found in Alhemnopsis, BnsilTichl!iy,~,Orlonlrsthes nigricnns and the subgenus Austrontenidirc, 62 l'~t~~oj~e~~rcu~lar.sc:r~.so7y cccnnl h~lrucccnpores onw and two.- is a rounded dorsal process of the opercle lacking a lateral lami- 7 I7 11c plcsio~norphicconclition oS atlleri~liformsis for the nar flange (Figs 19, 24). (O=opercular dorsal process blade- preopcrc~~larscnsory canal between pores one and two to like; l=opercular dorsal process not blade-like; 0.25, 0.57) 11e open (Figs 17, 18). A relatively derived condition prcse~ltin Mclan,ouI~,in,,~c.r,some specics of Ch,iro.rtornn, 65 O~~rculnrfines~ralio~~(White, 1985).- A smooth surface Ilnsiliclrll~y.~,and Odon,l~.slh~,sn,igricn.ns is the bony enclosure at the base of the opcrcular process is a plesiornorphic featl~re oSthc sensory canal bctwccn preopercular pores o~lcand of' at1leriniSorrns (Fig. 29). Presence of fenestration at the two (Figs 19, 24). Other mcnidiines have the dorsal shaf't anterodorsal corner oS the opercle is a unique trait within l~lcleclto a greater extent than atherinoids, but not fillly athcrinifi)rms, and is prcscnt in Odo~z~e.s:slk~.sexcept for 0. incisa cncloscd (Fig. 24R, 28C). (O=preopercular pores 1 & 2 (Fig. 18). (O=no opercular fenestra; l=opercular fenestra open; l=preopercular pores 1 & 2 enclosed; 0.50, 0.67) present on medial face; 2=opercular fenestra present on me- dial and external faces [Dyer, 1993: 3501; 0.33,0.78; add) 6 l'r~op~~r~clnrccr17nl pores on Aorizou~nlshe$.- Thc PI-cope~'cularsensory canal has thrcc pores on the vertical shaft 66 i\/lurlinl lrot~inccO~l7oeew oprrczrlar strut.s.- The mrdial face of and tivc porcs on tlle llorizontal shaft in atherinopsids (Figs 17, th; opercle has two strut-like thickenings that converge on the 19, 28C). A derived conclitio~rSound in the Bnsilichllzys semotilus hyomandibular condyle of the opercle. The ventral strut is ex- species group is presencc of only Sour pores on the horizontal tended along the anterior bordev of the opercle and the poste- sllaTt, apparently by loss of the pore seven (Figs 11, 24). ~rior-strut is directed posteriorly (Figs 'LO, 25). Presence of both (O=preopercular horizontal shaft with 5 pores; l=preopercular strrlts is a plesiomorphic trait and in atherinopsids there is a horizontal shaft with 4 pores; ?=unknown;1.00) semicircl~larspace between them (Fig. 25). A derived condi- Six ports arc present on the horizontal shaft of PoFlana. tion is prcsc~ltin Odontcsthes species except for 0. perupnr and M~sc.PIIIJL. Mus.. ZOOL.,UNIV. MICH., No. I85

horizontal shaft

...... ::---?, , pore 1 . .

pore pore 6 pore 5

horizontal shaft

Fig. 28. Latc.l.;ll vicw ol'tlrc- Idl prcopcrc Ic; A, i\g-onolo~~izi~nio?r/7c.olrr (UMMZ 190463); R, A/h(,r;no~~io~~i.\bk(,~k(,l-i (1:MNI-I 57159);(:, Mr17idio boyl617rr (UMMZ 203149); 11, Ath~ri17o/).\(I/'~?I~.s (IJblMj: 72212). DYER,A.L‘HERINOPSINE RELA~IONSHIPS AND BA.SILIC'I~TIIY.S 33

0. i,rci.ccc, in which both struls arc interconnected by a greatly 69 Ofierc;1clnrposlero71e~i~lrnlborder.- (O=opercleposteroventral Senestl-ated medial laminar bone (Fig. 14). (O=no medial lamina border convex; l=opercle posteroventral border concave; 1.00) between posterior and ventral struts; l=presence of fenestrated medial lamina between posterior and ventral struts; 0.33,0.80) Tlic postcrioi- strut is greatly reduced to al-)sent in Olyzins, notocllcirids, Alh~~io?~,(,'hirosto,w~r (Fig. 29) and M~lnnodzbni~s. 70 Arsl in~-n/)haryngobrc~n~/zinl.-Presence oS an ossified first inf'l-apbaryngobra11~11ialis the plesiornorphic condition for 67 Oj,c?ru~lardorsnl Oor(1n:- Tlie plesiornorphic condition of atllerinomorplls. A derived condition of and :~tlicriniSormsis for the dorsal border of the opercle to be con- Atheriilopsi~liis fol- the first infi-apha~yngobraiIchialto be cal-- tinl~ouswith the laminar flange oP the dorsal process. The de- tilaginous (Fig. 30). Absence of first i~lSrapharyngobra~~cI~i~~lis rived condition characteristic ofatherinopsids is for the dorsal a fi~rtherderived condition of Cyprinodontea, Noloch,eiru.s, and bordrr to be displaced medial to the dorsal process (Figs 19, Mc~nbradi~li(C:IlernoSS, 1986113). A cartilaginous nodule fo~lnd 20, 29), except in 0rionte.stl~e.s hnlclzeri, 0. peruginr, and 0. at the distal cild of epibranchial 1 in the cypi-inodontiforms ,wtro/)irini,s (Figs 17, 18). (O=dorsal border of opercle continu- 1;loridichlhy.s ca?$io, Cunlnc. 1r.sswllalus and Cynolebias elongutu.~,were ous with laminar flange of dorsal process; l=dorsal border of interpreted as a iirst infi-apllaryngobranchial by Parenti (1981: opercle displaced medially; 0.33, 0.67) 417). (O=infrapharyngobranchial 1 ossified; l=infra- pharyngobranchial 1 cartilaginous; 2=infrapharyngobranchial 68 O/)c~r1~~tlr~r/)o.tlat-orlot:sn1/)order.- The posterodorsal bordrr 1 absent; 0.40,0.50; add) of tl~copcrcle is concave in ;~therinomorpl~o~~tgroups (Fig. 29). A st~.;~iglltor convex posterodorsal border of the opcrclc is a 71 Craniccl /IT-o~ssof s~co17d i?~r(~~)I7n)~ngoO~an chin1 (Cberno f f, dcrivctl condition present in (;yprinodo~ltca,Atherinoidci, 1986b).- The gcneralized outgroup and atherinopsid condition I~~~Oirl~?c.t/ios,iM(?nrhrcls gilbrrti, i\/l~~lcn,~rh,inust7zicro)s, Atherinopsini, of infi-apllaryngobra~~chial2 is presence of a cranial process of and O~iottl~~~s~kes(Figs 1'7, 18). In atherinopsids, tllc opercular medium width that runs roughly pat-allel to postcrodorsal border is concave in menidiines and Bosilichthyt inf1-apharyngob~ilncl1ii1l3 and is articulated with it in a medial (Figs 19, 20,24, 25, 29). (O=opercleposterodorsal border con- groove. Athevinopsini has a clearly derived co~~ditionin that cave; l=opercle posterodorsal border straight; 2=opercle ~lrecranial process of infraphaiyngobranchial2 is pointed (with posterodorsal border convex; 0.67, 0.77; add) tlie distal chondral suvS:~ceol'articillation greatly redl~ced)and

dorsal process

lateral medial

Fig, 29. I.atera1 and rnedial views of the opercle of' C!L:I~OJIO~IL(II(I~(I~P (UMMZ 193463). 3.1 Mrsc. PUBL.Mus.. ZOOI., UNIV.MIC H., NO. 185 at right angles to the cr;mi;ll process of infi-apliaryngobraricliial l=epibranchial 3 cylindrical, with a long and laterally-placed 3 (Fig. 30; Chernoll; 1986b: fig. 7D). (O=infrapharyngobranchial uncinate process; 2=epibranchial three flattened and twisted, 2 cranial process of median width, mostly parallel to cranial with a long and laterally-placed uncinate process; 1.00 / 0.67, process of infrapharyngobranchial3; l=infrapharyngobranchial 0.75; add) 2 cranial process pointed, perpendicular to cranial process of infrapharyngobranchial3; 1.00) 73 hurth cer~~loDrun,ch,ialtoollz platp (Chernof'f, 1986b; White, 1985).-Tooth patches are not present on ceratobranchial 4 in 72 7%idc~/ih~-cl~zc.hinl zcncii~crl~ proc(~.\.s (White, 1 985) .- The mugilids, beloniforms, Nolocheirus, and atherinopsines. A tooth plesio~norpliicatlierinirornl condition of epibranchial 3 is for patch is present on thc proximal elid of cerato1~r;unchiaI4 in the luncinate process to be placed rougllly in the middle of the cyprinodontifo'or~ns,athcrinoids exccpt Notoc.lrrir.us, and bone, ;md a relatively straight and cyli~idricalshaft between the menidiines. The "tooth plates" in Co~~ic.lrth~~sdrscribedby White ~mcinateprocess and the articulation of cpibra~lcllial3 with (1985: fig. 4C) are e~llargedgill-raker plates supporting hair- infrapliaryngobranchial 3. The third epibl.anchial of like denticles, unlike those present in other athcl-iniforms and At1ie1-inopsiniis derived in that the mlcinate process is more not fiised with ceratobranchial 4. In Clf)iclzthys, the medial and laterally attached and elongated (Fig. 30). A Iilrther derived lateral rows of gill-raker plates contact each other at the dorsal condition of epibranchial 3 is for the shaft medial to the unci- mid-line cxccpt anteriorly, where the latel-al row of'rakers is cx- nale process to be flattened and twisted in Ath~,inoj>si.cand tcndcd across to the medial side. (O=no tooth patch on Ltt~1.1'~tkes(Fig. 3OB; White, 1985: Gg. 20). (O=epibranchial 3 ceratobranchial4; l=tooth patch present on ceratobranchial4; cylindrical, with short and mid-placed uncinate process; 1.00)

Fig. 30. Dorsal vicrv ol'thc Icl't rtppcl- phillyngcal jaw; A, All~o-inoj~.\rqjii~ic (UMMZ 72212); B, I,(~~ir(~.\..,tlrc~str17ltis (UMMZ 65910) I11 athcrinoids, the tooth plate is fused to ceratobranchial 4 lack of a pectoral spur in Notoeheirus and Gulaphallz~sseems to in Iso, illh~:rio,n,phallostchids, and craterocephalines. The tooth be due to absence of this element because the dorsal ray does plate is over but not frrsed to ceratobranchial 4 in not have the characteristic protruding thickness as do ~nelanotaeniicls,atherinomorines, and atherinines. atherinopsids (Fig. 11). (O=pectoral-finspur present, l=pecto- ral-fin spur fused to second pectoral-fin ray, not separate; 1.00) 74 I/?-oltycil ve77,li-alplfile- I'lesion1orpl1ically, the ventral bol-- der of the urohyal widens posteriorly into an oval-shaped plate 79 Venlml fiostcleithrum po~ition.- The ventral postcleithrum (Fig. 31A). This ventral plate is absent in Atherinops and sitrrated anterior to the first pleural rib is the plesiomorphic Atl~ttinopsis(Fig. 31B). This plate is also absent in Cratnacephalus condition for atheriniforms (Dyer & Chernoff, 1996). A de- /g~re,siispecies group (Crowley & Ivantsoff, 1992). (O=urohyal rived condition of atherinopsids and atherinids is for the ven- ventral plate present; l=urohyal ventral plate absent; 0.50,0.00) tral postcleithrum to be placed between pleural ribs one and two (Fig. 32A). (O=ventralpostcleithrum anterior to pleural rib Pccto~algirdle and fin one; l=ventral postcleithrum between pleural ribs one and two; 2=ventral postcleithrum between pleural ribs one and three 75 (;l(:itlrrunt,dorscmedial 7oiizg.- The anterior border of the [Dyer, 1993: 3531; ?=polymorphic;0.67, 0.91; add) dorsal-medial wing of the cleithrum is extended ve~lt~allyto- Atherinopsids share with atherinids and Den,tathmina an ex- wards ;I centl-al nleclial wing of the cleithrum. The plcsiornoi-pl~ic panded and laminar ventral postcleithrum, connected to the condition is lor the dorsomedial wing to extend ventrally as far dorsal postclcithrum by a dorsal ramus (Dyer & Chernoff, 1996). 21s the scapula-coracoid articulation or beyond (Fig. 32A). A Atherinopsids have, in addition, an anterior ranius that attaches dcrivetl conditio~iis for tlic anterior border of the dorsomedial nlostly to tlic first pleural rib (Fig. 32; Dyer & Chernoff, 1996: wing to Ilc tnorc curved or involuted, and extended ventrally as fig. 17). Par as tlic scapular foranlcn at the niost (Fig. 326). The derived Pelvic girdle and fin co~iditionis present in Mc~iidiinac,(;olpichthys, and Odontesthe,~ species except lor 0. kalcl~,eri,0. h~~~nzrns%.s,0. pmt~giae, and tl~c 80 Pel-oic bone medial plate (Chernoff, 1986b) .- The s~tbge~l~tsCnzipip. (O=anterior border of cleithrum dorsomedial plesiomorphic condition of atherinomorphs is for tlie medial wing tapered ventrally to scapula-coracoid articdation or be- plate of the pelvic bone to extend to the anterior end of the yond; l=anterior border of cleithrum dorsomedial wing curved longitrldinal shaft. In atheriniforms, the pelvic medial plate ventrally, extended to scapular foramen at most; 0.20, 0.64) does not reach the anterior cnd (Dyer & Chernof'f', 15196) ex- cept for atherinids, phallostethids, Atherinella schultzi and A. 76 I~ci~iclelot',c ligcinloi~l po.rterodorsc11 attacl~monts.- The inarvelae, Leuiz.sthrc, and the subgenus Cauque (Fig. 34D). (O=pel- plesio~riorpliiccondition for atheriniforms is for Baudclot's lip- vic medial plate extending to anterior tip; l=pelvic medial plate lnent LO IIC attaclied posterodorsally to the dorsornedial wing of not reaching anterior tip; 0.33, 0.50) tl~cclcitlirrr~rl and anterovcntrally to the neurocranium. A de- rived condition found in atherinopsids, except for i\.Ien,idia and 81 Pelvic Don,? median proc(?.~.s.- The plesiomorphic condition I,nOid~,sll~e,s,is Ihr Baudelot's ligament to be also attached of the pelvic nledian process is a dccp, strong, percoid-type ar- postc~-odorsallywith the parapophysis and epinerrral of the first ticulation between pelvic bones, as found in mugilids (Stiassny, vertebra (Fig. 33). (O=Baudelot's ligament attached posteriorly 1990,1993;Stiassny 8c Moore, 1992;Johnson &Patterson, 1993: to cleithrum only; l=Baudelot's ligament attached posteriorly (5 19). Atherinomorplis exhibit the plesiomorphic to cleithrum, and to parapophysis and epineural of first verte- acanthomorph condition of flattened pelvic median processes brae; 1.00) that overlap at the ~nidline(Fig. 34A; Stiassny & Moore, 1992; Saeed et al., 1994). This condition is regarded as a reversal in 77 I'ec.torcilstr~it,s(White, 1985).- Dorsal enclosure ofthe pec- the context of relationships proposed by Johnson & Patterson toral abdr~ctorlnrrscles by the cleithr~r~iiis a plesiomorphic fea- (1998). In atherinopsiues except for L2eurt.slhe?s, the overlap of ture of Atherinomorpl1;l (Dyer & Chernoff, 1996). A derived the median processes is greatly reduced to absent (Fig. 34C, feature of atherinopsids is presence of str~~ts,oval in cross-sec- 34D; De la I-Ioz & Tosti-Croce, 1981). (O=pelvicmedian process tion, bctween the lateral and medial extensions of'tlle cleithrum percoid-like; l=pelvic median process flat and overlapping at (not between cleithrum and scapula as in White, 1985). (O=no midline; 2=pelvic median process flat, slightly overlapping or pectoral struts; l=pectoral struts present; 1.00) abutting; 0.67, 0.86; add)

78 I'c,clo,nl;/i?~sfiu,r (Sti;rssny, 1993).- Presence of a pectoral Median fins sprrr (rr~ditnentarypector;rl-fin spine), dorsal and pt-oximal to the second unbranched ray, is the generalized condition in 82 I'ositio17 of/ir.st do?-sal,fin.- Origin of the first dorsal fin ;~therinoniorphsand mrrgilids (Stiassny, 1993). Lack of this spur well in advance of the anus is considered a plesiomorphic fka- is characteristic of Nolocheilus, atherinopsids (Fig. I I), and ture of athcriniforms. The first dorsal-fin origin located over C~ri1~~~)hc~lkcl.s.Stiassny (19113: 210) proposed that tlie lack of a or posterior to a vertical from the anus, is a derived condition pectoral S~III-in atllcrinopsids is due to ontogenetic li~sionof present in atherinoids, menicliines except for Melanorhinus, and this elelllent with the median hemitrichium of the unbranched in Atherinups, Colpicht11g.s huODsi, Leuresthes, Odontesthes ppmugiaa pcctoml-Gn ray. Evidence supporting the fusion of these ele- 0. retropinnis, 0. Donarieiz.si.s, 0. argen,tin,ensis, and 0. incisa (Fig. rrleiits is presence of pectoral spurs in spccirnens of Basilichthys 35). (O=first dorsal fin well in advance of anus; l=first dorsal- under 30 mm SL (UMMZ 2 15491 ; CAS 45197). Conversely, the fin origin over or posterior to anus; 0.38) Mlsc-. PIII~I..Mus.. ZOOL.,UNIV. MICFI., NO. 185

ventral lateral ventral lateral

Fig. 31. I.;~tcr;~l;~ntl ve~~tl-;~l vic\v\ ot the 111-ol~yal;A, Oclo~rli~c/ho.\y@rc (UMMZ "15521); U, /\ll~(,r.i~ro/)s/fJn/.c (LIMMZ 72212)

111 (:ol/)ic.l~/l~.c,tlic lirst dorsal Gn is locateti a~itrl-iorto tllc 84 I~~L~~~-~~~~-J~~~~P~~~~~o/III~oT~~s.-A sel-iesof ptcrygiophores with- allus ill (:. I-rgi~and posterior to tl~eanlls in (;. h~thhsi. 111 o~~tassociatccl Iin rays het\z~eenthe dorsal fins (interdorsals) is atllcri~~oids,the first dorsal fin is a~iteriol-to tllr alills in Ito, a leaturc cllaracteristic of some ~adifi~rms(Marlile, 1989: fig. illlrc~io,~,and in a Sew ~~ielan~tae~iiidsand atllcrinids. Thr pos- 13; Okan~~u-;I,1989: Ggs 3,4),mlrgilids, and rnost athel-iniforms terior l>ositiori of tllc lirst dors;~lfin relativr to tllc atlus in (Parenti, 1993: fig. 8). Iriterdorsals appear to be absent in Iso Atherilles is mostly becausr of an anterior displaceliient of tllc and pselrdon~l~gilins(Dyer & C:her~ioll;1996), though strictly

;Inus, effected by presence of tllc kr~lalplate ant1 elorigation of the last ptcl-ygiophore of tlir first dorsal Gn always 1.nc k.s ~ts' col-- thc anal-1i11base (Dyer tk (:hernofi', 1!)!)6). responding ray (Figs 35-38). Notocl~rirusa~ld(:yprinodontea lack a first dorsal fin and have IIO rayless l>tcrygiophorrsanterior to 83 L~nglkq/spi,tl~s o//irsl rttad ,soc.ond dot.sr~1fil1.c.- 111 the dorsal lin. The interdorsal ptcrygiopl~oresof atllerinopsids atlicri~lili)~-msand ;ltI~e~-illornorl)houtg-I-oups, tllc Ic1igt11 of tlle arc present 21s lanlinar plates, without a vcrltral shaft, reduccd ;~ntcriol.spine of the lint clorsal ti11 is greater tllall tlir spino~~s to nodules (ossified or cartilagino~~s)posteriorly along the dol-- ray of the second dorsal fin. A dcl-ived conditio~~of Nasilichll~ys sal ligament that collnects both fins in the vertical septlun (Figs is for thc anterior spine oS tlir lirst dorsal fin to be shorter than 35-38). A continuous series of interdorsal pterygiophores is tl~espino~~sray of the second dorsal fin (Fig. 36). 'This featurr present in atherinoids exccpt for Iso, in rnost rncnicliines, is cspccially noticeable in the II. .\~~~IIo/~~LLsspecies gro~~p which Colj~ichthy.~,and in tlie sorgentinins Ba,siliclzlI~~~s(Fig. 36), has tlie Lint dorsal fin grratly retluced, sonlctinies absent. Odonli?sllrc:s hatcl~w-i,0. hutn(>~.rsis,and the subgenus C;ccucjua All (O=first spine of Dl longer than first spine of D2; l=first spine other sorgentinins exanlined have a gap in the series oP of Dl shorter than first spine of D2; 1.00) interdorsal ptcl-ygiophores (Figs 37, 38). (O=continuousseries of interdorsal pterygiophores with ventral shaft; l=continuous scapula dorsal \ postcleithrum

Fig. 32. Medial view of'lel't prrtoral gi~.tlle;A, O~OII/I,.SL/LI!.~hrm~Inn,a6.\ (UMMZ 215451)); R, Odo?~festhr~rqirr (UMMZ 215521 ). series of interdorsal pterygiophores without ventral shaft; in tcvo states. In Basilichthj~sthere are I-oughlyten more precaudal 2=shal'tless interdorsal pterygiophore series discontinuous; vel-tehrae than caudal vertcbl-ac, i.e., precaudal vertebrae ac- ?=ambiguous [Dyer, 1993: 3541; 0.50,0.82 / 0.40,0.73; add) count for over 57% of the total numbcr of vertebrae (Fig. 36). Exocoetoids also have at least 60% precaudal vertebrae. 85 N~ilnh~ro/ i'nterrlorsal ple,rygiophores.- Interdol-sal Odonle~thesspecies have rrlorc than 50% precaudal vertebrae ptcrygiophores (Ch. 84) are found in varying n~unbel-sin except for 0. inciscc, 0. 17,ipic(cn~,0. platen.sis, a~ldthe subgerius m~igilidsa~rd atlleriniforn~s (Parenti, 1993; Dyer & Chernoff, Austromenidia which have Illore caudal than precaudal vertebrae. 1996). M~igilidshave three or four interdorsal bones. Iri Roiner's definition of caudal vertebrae is used (Ronier & Pal-- athcri~ioids,Iso has three interdorsals, Atherion and I1c.nrathmn.a sons, 1986; see Ch. 95 and below in Discussion: Axial skeleton). have more than six, nie1:iilotacniids have four or fewer, and (O=precaudal vertebrae under 50% of total vertebrae; athcl-initls havc morc than Gve. Menidiines have six to nine l=precaudal vertebrae more than 50%; 2=precaudal vertebrae iritertlorsals except for small species of Mendia (colei, parva, etc.) over 57%; ?=polymorphic [Dyer, 1993: 3561; 0.67,0.90; add) and some spccics of Cl,i,ro.stomc~(labnrca~, argp, etc.) and Poblnnn. Atlleri~ioysi~ieswith six or more interdorsals include Colf)ichth~l.s, 87 First pleztral ri/).- (O=first pleural rib long, attached to ven- Otlonlesllres hnlch(~i,0. l~~mrrt~nsis,0. i7zcisa, arltl Basilichthy,~(some tral postcleithrum; l=first pleural rib short, not attached to ven- specimens have li~uror fivc longer bones whicl~seem to be a tral postcleithrum; 0.50, 0.75) pl-otlr~ctof fusioll of two ?lcments; Fig. 3G). The remainder of atlicri~iopsincshave three to six interdorsals, ust~allyfour or 88 Posterior pl.ural ribs.- Pleural ribs are attached to the live (Figs 35,37,38). (O=six or fewer interdorsals; l=more than parapophyses of vcr~ehi-aethree, to the posterior precarldal ver- six interdorsals; ?=polymorphic [Dyer, 1993: 3551; 0.25, 0.63 / tebra, and so~netimesto tlle anterior caudal vertebra. The 0.20, 0.50) plesiomorphic condition of atheriniforms is for the posterior ribs to be anterior to thc dorsal tip of the first anal-fin Axial skeleto~i pterygiophore (Fig. 38). Presence of 1-ruo or more pleural ribs posterior to the dorsal tip of the first anal-fin pteiygiophore 86 Number- oJ fi.rci,rrns, atherinopsids exccpt for , Melanorhinus, Odon,tc.sthes atlierinoids, rncnidiincs, il~ldatl~e~-i~~opsi~li~~s is for the number plat~nsis,0. nign'cccns, 0. incisa, and the subgenus Au,stroinc.nidicc. oP preca~~dalvertebrae to be equal to or fewer than the caudal (O=pleural ribs anterior to first anal-fin pterygiophore; l=pres- verteb~xe.Within athcrinopsines the derived condition of hav- ence of two or more pleural ribs posterior to first anal-fin ing more precaudal vertebrae than caudal vertehrae is present pterygiophore; 0.50,0.89) MIX. Prlul-. Mr~s..Zoor.., UNIV.MICH., No. 185

Baudelot's ligament

Iig. 33. l)orsal view ol the occipic;ll region ol the hcatl, pcctolxl girdle, ant1 antcrior \:cl.tcbrae 01' Orlo~~/rs/hc~tb~rvicrnnllt (UMMZ 21 545'3). Lef pcflo~'algirtllc and rig111 postct~~porxl,est~-;~sc;tpi~l:u-s, a11tl exocc-ipital wing ~.ell~ovetl.

89 Pos(c.1-rorrxt(~r7,sion of switn,l)/c/dtl(r-i17lo a l~crmzcr/firnnrl (MThitc, (F~I-tl,1937; clot hie^; 1950), in sillaginids (McIhy, 1992), and 1985).- Tlrc s~vimbladderis extc~rdedposteriorly to the Iirst ill some cheiline labrids (Westneat. 1993). (O=no swimbladder ca~ldalvc~.tcbrae in atlic~.inifor~llsand outgrottps. A derivctl funnel; l=swimbladder funnel present; 0.67, 0.86) I'?at~~reofatherinopsines except for Colpic.l~~l~~~.sll~~rl~hsi, O~OIZ/I

Fig. 3.1. Vcncral view ofche left pelvic girdle; A, Alho~rtellrbl-asilzrnsis; 8,Mrnidin; C; Ba~ilichthys(UMMZ 215508); D, Odonte.Ythe,i hrmianalis (UMMZ 2 15459). (A & 13 modified from (;hernoff, 198Ga) MIX. PLJUL.Mus.. ZOOL.,UNIV. MICH., NO. IS5

interdorsal pterygiophores

first caudal verte

haemal-arch funnel

first anal-fin pterygiopho

ITig. 35. I.;ttc~-211\,ic.w ol nictli;ur firis ;l~rdaxial skeleton of Orlontc:.\thcc rc,tr-o/)rn11~s(UMMZ 231950).

first dorsal-fin interdorsal pterygiophores second dorsal-fin spine

Ch. 91 (2): haemallarch flaring haemal spines

Fig. 36. Frontal vicw oScat~tlalvc.l.tchra, and 1;ttct-alviclv of dor-sal fins and axial sltclcton of Hrr~~licltll~yssp. (IJMMZ 215510). interdorsal pterygiophores second dorsal-fin spine first dorsal-fin pterygiophore A I

haemal spines

anus /. anal-fln spine Fig. 37. 1,ateral vicw oS median fills and axial skelvtolr oS Odon~~~fl~c~s6owcrrirn~i.s (UMMZ 218609). scls (Fig. 39; Wliitc, 1985: fig. 11 ). I-lac~nal-hypophysiscxpan- and 0. srnitti, 0. r(?~?'a,0. gracilis (subgenus A uslromc.niclirr)have sions :u-c retl~~cedto ,just two or three vertebrae in C,'olpic.hthj.c anterior and posterior projections restricted to the base of'the liul,O.si (C~-itbtrcc,1989). (O=no haemal-hypophysis funnel; haemal arches in ~hcposteriol- region oS the haemal f'1l1111el(Fig. l=haemal-hypophysisfunnel present; 1.00) 38). The flaring of the hacmal arches is rnost developed anteri- The hac~nall'unnel oP atherinopsinins resembles that of orly except for the most anterior one (Figs 3638). (O=no haemal- sillaginicls (McIhy, 1992) and Athprinnscn h(@etoide.s, including arch funnel; l=haemal-arch funnel present; 2=haemal-arch flar- the sel~aration01' the blood vcsscls fi-on1 the swi~nbladdcrby a ing present; 3=anterior haemal-arch projections present; 4=an- bony l~ridg~(I'attcn, 1978: 123 &fig. 32C:; McKay, 1992: figs 6 & teroposterior haemal-arch projections present, restricted to ven- 7). tral half of haemal arch [Dyer, 1993: 3581; 0.40,0.81; add)

91 f-ln(~rnc~l-arch,funn~L nntl rxpnnsion.c (White, 1985).- The 92 Post(.rior r~xlmsionoflli,~ h(~e~nal-nrch,~~~i~z~~l.- (O=haemal-arch liaemal arches in athcri~iifornisand outgroups have no cxpan- funnel extended posteriorly to end of anal fin or beyond; sions or ~noclifications. A derived feature of sorgentinins ex- l=haemal-arch funnel extended to mid-anal fin; 0.50, 0.67) ccpt lor O~lo,rt(?,sll~nirrcisa :rnd 0. nigtl'ccr.?r.s is the posterior pro- jection of tlic swimbladder, togcther wit11 the rriajor I~loodves- 93 Hne~nal-nnhjunrzel floor shnflg- (O=haemal-arch funnel sels, into expanded haemal arches (Figs 35-38; White, 1985: fig. floor gradually tapering posteriorly, bulged ventrally; l=haemal- 11; Dc la I-Ioz & Tosti-Croce, 1981 : fig. 12). The haemal-arch arch funnel floor abruptly tapering posteriorly, bulged dorsally; f~unnrlof sol-gentinins is present in Sotu- conditions, each state 1.00) inclusive oS the previous: 0. hnlch~rianti 0. ratro/~zf,i.nni.slrave a haemal S~uinelbut lack ally form ()Snaring or projections (Fig. 94 ITaem,ccl-sfiin~length.- Haemal spincs of the anterior 35);sorgc~itinills except Sor the four spccies mentio~~cclabove, hacnral arches are primitively long. A derived condition present have flaring expansio~~sor the haernal arches (Fig. 36); in Oclontestl~c

interdorsal pterygiophores first dorsal-fin pterygiophore second dorsal-fin spine 1 n \

neural

Fig. 38. 1,ateral view of'rnedian fins and axialI skeleton of Odontc,.\the.~re~ia (UMMZ 215521 )

95 Position ofSirst caudal umte6m.- The first caudal vertebra imbrication; l=reversed imbrication of dorsal head scales with (see below, Discussion: Axial skeleton) located anterior to a ver- occiput scale in nuchal position; 2=reversedimbrication of dor- tical Prom the anal-fin origin is a plesiomorphic feature for sal head scales with occiput scale at postorbital rim; 1.00; add) atherinirorrns (Fig. 38). The derived condition is for the first Thompson (1916: 464) was first to describe this distinctive caudal vertebra to be located posterior to a vertical from the and highly unusual feature of Basilichthys, which was subse- anal-fin origin and dorsal to the anterior half of the anal fin. quently used by Jordan 8c Hubbs (191 9), Hildebrand (1946), This derived condition is found in menidiines except for and Arratia (1981). Thompson (191 6)also described the pres- Melanorhinus and small species, and in Odontrsthes ence of this feature in Menidia alburnus Giinther (=Odontesth(?s hatchm:, 0. humensis, 0.pmugzae, and 0.mtropiwnis (Fig. 35). A nipcans) restricted to the anterior two or three rows of scales. f~lrtherderived condition fo~lndin Basilichthys is for the first I was unable to confirm this observation on the type specimens caudal vertebra to lie dorsal to the posterior half of the anal fin. or on any other available specimens of 0. nipcans. (O=first caudal vertebrae anterior to anal-fin origin; l=first cau- dal vertebrae dorsal to anterior half of anal fin; 2=first caudal 97 Number of suborbital scale rows.- Three rows of scales below vertebrae dorsal to posterior half of anal fin; ?=polymorphic the orbit in adult specimens is considered the plesiomorphic [Dyer, 1993: 3591; 0.50, 0.80; add) condition for atheriniforms. Three suborbital scale rows are found in mugilids, cyprinodontiforms, Notocheirus, Atherinopsini Scales except Ath,erinops, Odontesthrs hatchmi, 0. h,umc.nsi.r,0. bonariensis, 0. incisa, and 0. gracilis. Two suborbital rows are found in 96 Imbrication of dorsal hmd scales (Thompson, 1916).- The menidiines, Athminops, 0do.ntesthes pmupae (Fig. 23A), and 0. plesiomorphic condition found in atheriniforms is for the dor- retropinnis. Four suborbital scale rows are found in Basilichthys sal scales ofthe head, over the frontals, to have regular imbrica- (Fig. 23B, 23C), Odontesthes argentinensis, 0.platensis, 0.nigricans, tion, i.e., anterior border imbedded in the skin and posterior 0.repa, 0. smitti, and the subgenus Cau.quc (O=three suborbital border free. A uniquely derived condition found in Basilichthys scale rows; l=four suborbital scale rows; 2=two suborbital scale is for scales in the interorbital region to have reversed imbrica- rows; ?=polymorphic; 0.29, 0.58; n-add) tion (Fig. 16). In the B. microlepidotus species group the occiput scale (transition scale between regular and reversed imbrica- 98 Interopercle scale.^.- Presence of scales along the entire tion) is in a nuchal position (Fig. 16A),whereas in the B. semotilus length of the interopercle is a plesiomorphic feature of species group the occiput scale is located at the level of the atheriniforms (Fig. 17). A derived condition found in postorbital rirn (Fig. 16R). (O=dorsal head scales with regular Basilichlhys, Odontesthes nigricans, 0. humensis, and the subgenus Fig-. 39. Lateral and frontal views of anterior caudal vertebrae of Athm'nops (modified from White, 1985)

Cauqu,e is presence of scales on the posterior half only of the present in atherinopsinins except for Atherinops, and in interopercular bone (Fig. 19). (O=interopercle fully scaled; 0dontesthe.sspecies 0. hatchm', 0. argentinensis, subgenus Cauque, l=interopercle with scales on posterior half only; 0.25, 0.40) 0. nigricans, 0. in,cisa, 0. platenuis, and the subgenus Populations of the B. scmotilus species group from Rio San Austromenidia. (O=no scales on cleithrum shaft; l=scales present .Juan, Cajamarca, and Rio Rimac, Lima, and some populations along cleithrum shaft; 2=small scales only present along of 13. australis have a few scales on the anterior half of the cleithrum shaft [Dyer, 1993: 3611; 0.29, 0.67; add) interopercle, but they do not extend to the anterior end of the bone as in the plesiomorphic condition. 100 Scales on second dorsal /in.- Second dorsal iin without scales In Cyprinodontea, the interopercle is not exposed laterally is a plesiomorphic feature of atheriniforms. Presence of scale and laclis scales. Atherion and Iso also lack interopercular scales. rows between rays at the anterior base of the second dorsal fin, though present in mugilids, is considered derived in Leuresthes, 99 Cleith,rum scales.- No scales on the lateral shaft of the Basilichthys semotilus, Odontesthes humensis, 0. perugiae, 0. cleithrum is a plesiomorphic feature of atheriniforms. A de- retropinnis, 0. bonariensis, and 0. argentinensis. (O=no scales be- rived condition is presence of large scales along the entire length tween rays on second dorsal fin; l=scales present between ante- of the cleithral lateral shaft (Fig. 40). This derived condition is rior rays of second dorsal fin; 0.20,0.43)

supracleithrum

f .

Fig. 40. Lateral shaft of left cleithrum with scalrs; A, Odontesther argentinensis (UMMZ 231948); B, Odontesth~sregin (UMMZ 215521). 4 4 MIS(..P~I~I.. Mris.. Z~OI-.,UNIV. MICH., No. 185

This feat1u.c is not to IIe conf'rrsetl with thc s1lc;rth of'l~odyscalcs ~.oboratcdas diag~losticof Mcnidiinac: (1) parapophysis of ve~ raised along the base of the scco~rtltlorsi~l fir1 prcsc~rtin i\/l(lrnDrclc trbr-a I enlarged; (2) epi~ler~ral1 sllort, attacllcd posteriorly to and M(,lnrior-lrirru.\ ((:hcr~ioff,19ti(ib: 237). dorsomrdial wing of clrithrr~ni;(3) infi-apli~~ryngol~rai~cllial2 with deep ~ncdialgroove; (4) infi-apharyngobranchial 2 toot11 plate short and rvitle; (5) vagus ncrve foranlc~lwith posterior 101 Scccles 011 nncz1tzn.- Anal GII without scales is co~isidrl-eda plcsiomorphic featm-c for ;rthcriniforrns. 71'hc derived presence of wall (Fig. 13); (6) ~lroliyalpedicel with wide dorsal surface; and rows ol'scales between rays at the antrriol- I~aseof' the anal fin is (7) pelvic bone with vcl~tl-alcl.adlc for fin rays. Tllc above-listed found in mugilitls, in athcrinopsinins except for (;olj)icht/~j.\,in cl~aracterswere used to provide (:li. 106 with a weight of' seven Bnsiliclr/kg,s .v(~rrzotil~rs,and in Odo171(:slh~~Ir(ztch~li, 0. ~U~PIZS~S,0. in thc analysis. (1.00) /)ctr-ug.icr(~,0. ,~lro/)irr~ri.s,0. bonnri~nsi~,0. nrg?nlirl~rrsis,and subgenrls (~trr~qu~o.This featrlrc is not to 11c co~ifilscdwith tl~csheath of'body Allozyme charactel-s scales r;\isetl along the base of the anal fir1 prcsc~ltill ~iioat mcmbladins (Chc~.noff,198611: 237). (O=no scales on anal fin; 107 - 123 Cl~nrc~clc>r-slnl(~c.ocl(>d allozjvzo cl(~lu(Crabtree, 1987) .- l=scalcs present between anterior rays of anal fin; 0.20,0,64 / 0.17, Tllc hllowi~igis a list of tlic c1-raracte1.staken fro~iiCrabtree 0.55) (1987) in the order of his table 4. The name of each allozynle character is followed hy its acronym ill 111-ackets and the num- 102 Scolo size- The size ol'sc:~lcsis ~lsuallyrstim;~ted by the I~crof character states. (:liai-acter states al-e coded qrlantita- n~m~berof sc;lles on tlic latcral sc~lsoryli~ie 01; ill tlie case 01' tivrly and qualitatively, and as in Table 3, qualitative coding is in silversidcs, along tllc lateral band. Becal~sethe nrrmber of lat- l~arenthesisonly when the two coding schemes differ, c~.;tlscitlcs is associatetl with the ~~u~riberof'vertebr:~e, ~vliich is 107:Ade11ylatekinase [Ak-A:4J (1.00); I08.Aminopcptidasc quite variable, the nrrmber of' sc;llc rows that extend over the [Ap-A:2] (1.00); 109: Aspartate arninotl-a~lsferase[M-Aat- dors11111l~etween the lateral Ixlncls is co~isidercd;I Illore :~ccu- A:51(1.00); 110: Aspartate a~iii~lotransfcrasr[S-Aat-A:6] (1.00, rate rsti~~~atcof scale sizc among closely related species. Rela- 0.0); Ill:Dipeptidase [Dip-A:5(7)](0.80,0.0/1.00 (1.00,O.O)); tively large scales is the p1csiomo1-phic conditio~lin 172: Esterase [Est-1:5] (1.00, 0.0); 113: Glucose-(?-phosphate atherinilbrms. Small scales are ;r tlcrivcd coilditio~la~~cl are isomerase [Gpi-A:6] (1.00, 0.0); 114: <;Iucose-6-p11osphale li)1111din some species of Cl1,iro~torrrci,ill Atheritto,/~sis,and in the isomerase [Gpi-B:7] (1.00,O.O); 115: Glutamate dehydrogenase sorgentinins 13nsilic/~ll13!.s,Otlorcl(~slIrc?.s /r,c~lclr,eri, 0. ~z~I-L~~L~zs,SLIII- [Gtdh-A:4(5)](1.00); 115: Glycerol-3-phosphate deliydrogenase ge~lus(:(z%L~~LP, 0. /~lnlor~sis, and the suhge~~usAustronr~nirlitc. [G3pdl1-A:8(9)](1.00, 0.0); 11 7: Isocitrate dehydrogenase [M- (O=large scales [Ill dorsal scales]; l=small scales 1212 dorsal Icdh-A:J] (1.00, 0.0); 118: L-lactate dehydrogenase [Ldh- scales]; 0.20, 0.60) A:9] (1.00, 0.0); 11 9: Malate dehydrogenase [S-Mdh-A:4](0.75, 0.50) ; 120: Malate dcllydl-ogenasc [S-Mdh-B:5 (7)](l.OO(1.00, 103 I'octwrot Oor-(l(>ro/ \lcr/(,\.- (O=bodyscales cycloid with pos- 0.0)); 121: Pl~osphogl~~comutase[Pgm-A:8(9)] (1.00,O.O); 722: terior border smooth, l=predorsd scales crenate; 2=alI body Pyl-trvirte kinnse [Pk-,4:8] (1.00, 0.0); and 123: Supel.oxicle scales crenate; 0.50, 0.60; add) dis~nutase[Sod-A:6] (1.00). (?=unknown;all n-add)

104 n/l onol,/r.yly ~/iltl~/~ri,io~trorj)Ii~~(Stiassny, I '.)!)0, 19113; Pare11ti, 1993) .- Of a list of 18 ch;l~-actcrsconlpilcd fl-om l'arenti ( 1993) and Stiass~iy(1993), five unique fcat111.c~arc selected to tliag- Claims of monophyly are sr~pportedsolely by diagnostic Sea- nose atherinornorphs: (1) spcrmatogoni;i I-estricted to distal end tul-es, i.e., character-s that are unambigrlor~slyderived at that ol' testicular tubules; (2) pelvic girdles 1;lckilig 1net1i;rn srlture node (whether ruiique or homoplasious) in the contrxt of tllc illid dot.solateral processes associated with plcru-a1 ribs (Fig. 34) ; ~wopuwdhypothesis of phylogenctic relationships. A cornplete (3) sul~aclcitlirunlrcdr~cetl ancl saddled on cleithl-al dorsal list of diagnostic and sl~pportingcharacters is presented in Ap- cnclosrlre lather tllan laterally; (4) r~ilargedfourth cpibrancllial lxndix 2. Most nodes llavc an additional suite of characters ;rnd fourth inSrapl-rary~lgobr:t~iclii:~l(Fig. 30); and (5) olfactory that are ariibiguorlsly optirnizcd at that 11ode but are, rieverthe- sensory cpithelilrm arlxnged in selisory islets. The above-listctl Iws, deemed informative of the group's Iiistorical identity (Dyer chal-acters were used to provide (:h. 104 with a weight of five ill & Chel-noff, 1996). Rather than cl~oosi~iga single optimization tlie analysis. (1.00) criterion for all charactel-s prior to the phylogerietic analysis, each a~ribigr~or~scharacter state is opti~iiizcd(L fio.\t(,r-iu~iby dis- 105 il'lorrol,lrj~lyo/'Ath(.r.ini;f~rrrz~~s (Dyer & (:Iic~.~lofl',1996) .- Of' tinguishirrg types of a~rilIiguity(Appendix 3). ;I list of ten cllai-acters listed by Dyer & Clicrnoff (1996) as tliag- rrostic of atherinif'ornls, three r~lliqrlecl~aracters ;Ire selected for the purpose ol' this study: (1) larval pig~nc~ltationpattern and prcanal length; (2) lacrimal l~rancllof A1 muscle tendon Atlierinopsic1;le comprises the New World silverside subf'ami- long, attached to subnasal shelf of lacri~liall~o~le(Figs 22, 26); lies Menidiinae ancl Atherinopsinae. Atheri~lopsidshavc tradi- ;und (3) presence oL'a pelvic-rill 1ig;rmcnt. The above-listed chat-- tionally been co~isidereda part of Atherinidac (Jordan & I-Iubhs, acters were used to provide Cli. 105 wit11 ;\ weight of three in 1919; Schultz, 1048; Patten, 1978;White, 1985; Cl-rernoff,1986b), thc analysis. (1.00) bllt l-rerc and in Dyer & (2le1-noff (1996) they arc co~lsidered instead as the sister grorlp to all other atherinifol-ms. A total of 106 hfO770j/~/l~~~/'lbf~lt~d~'~l(~P (Ch~rll~fl; l<)#(jII) .- (1f':l list Of 20 diagnostic cl~;iracte~-ssupport At11e1-iiropsidacas monophyl- eight c11;lracters proposctl 11y C:11el-noff (1986ll), seven are coi-- etic: sphe~loticpostorhital process \vide at hasc (Ch. 18), der~nosphrnoticnot articulated to postorbital process (Ch. 19) rior border of cleithrum dorsomedial wing curved ventrally, and with sensory canal enclosed (Ck. 20), presence of pterotic extended to scapular foramen at most (Ch. 75), and two subor- wing (Cli. 29) and exoccipital wing (Cli. 30), postemporal with bital scale rows (Cli. 97). Additional supporting characters for anterior process (Cll. 36), prcrnaxilla with narrow a~iteriorjoint menidiines is a postemporal sensory canal with three pores and (Ch. 39), premaxillaly ;~lveol:~rarm greatly expanded (Ch. 44), no extrascapulars (Ch. 35), pre~naxillaryascending process with l~rcmaxilla-dentarycoro~loid ligatne~lt very short (Ch. 45), pres- narrow base (Ch. 38), palatine spine extended to quadrate (Ch. ence of a maxilla-angl~loarticularligament ((:11. 46), prcmax- 57), and origin of the first dorsal fin posterior to tlie anus (Cli. illa-maxilla ligament long (Ch. 47), ethmomaxillary liganlent 82). with palatine attachment (Ch. 55), vertical shaft of preopercular Monophyly of Menidiini was supported by Chernoff (1986b) canal enclosed (Cli. (il),dorsal border of opercle displaced with two characters: a fused upper hypural plate and no pelvic meclially 1'1-omdorsal process (Cli. 67), Baudclot's ligame~ltpos- axillary scale, however the latter is a~nbiguousbecause it is also teriorly attached to cleithrr~tn,parapophysis and cpineural of lacking in atherinopsines. Another two featut-es found to cliag- vertebrae 1 (Ch. 76), prcsciice of pectoral struts in cleithral tlol-- nose menidiins are absence of a dermosphenotic medial flange sal ericlosure (Ch. 77), pectoral-fin spur fused to seco~ldray (Ch. 22) and absence of endopterygoid teeth (Ch. 59). Mono- (Ch. 78), ventral postclcithrum between pleural ribs one and phyly of Menibradini is supported by six features, four ofwhich two (Ch. 79), interdorsals witllout a ventt-al shaft (Ch. 84), plcu- are unique (Cliernoff, 198Gb; Appendix 2). Anotl~ertwo diag- n1 I.' 111s posterior to first anal-fin pterygiophore (Ch. 88). Char- nostic features of this group are thc hyomandibular nerve di- acters 18, 30, 39, 44, 45, arid 77 are unique f'catures of vided inside of the hyomai~dibula(Ch. 60) and absence of Atherinopsidac. Additional supporting characters arc presence infi-apharyngobrancl~ial1 (Ch. '70). of tlie anterior vorneritle slielf (Ch. 4), tlie secondary trigeminofacial arch ((211. 25), labial ligarnenc folded lateral to Monophyly of Atherinopsinae premaxilla (Cli. 48), the dorsal process of the palatine (Ch. 54), palatiile extendcd posteroventrally to quadrate (Ch. 57), many Athcrinopsinae comprises six genera in two tribes (Fig. 1): intcl-dorsal bones (Ch. 85), and first caudal vertebra dorsal to Athcrinopsini (Alheri.n.lzops,Atherinofisis, Co&ichtlzys, Leumtlt,c:,s) and anterior half of anal lirl (Ch. 95). Adclitiorial supporting char- Sorgentinini (Ba.sil%cklf~ys,Odontesthes) . Atherinopsincs form a acters proposed by Dycr & (:liernoff(1996) are ~naxillarybranch mor~ophyleticgroup diagnosed by eight characters: ethmoid of the Al rn~~scletendon attached to the distal half of the max- cartilage folded ventral to vomer midline (Ch. 6), nasal ventral illa ancl pelvic girdle attached to pleural rib (5 or more posteri- process overlaps subnasal shelf (Cli. 1I), nasal sensoty canal orly. detached anteriorly (Ch. 12), postocrllar shelf at dorsal half of The present asscmblagc of nrenidii~leand atherinopsine gen- dermosphenotic (Ch. 23), basisphenoid belofragrn short (Ch. rl-a was first recognized at the family level by Jordan & I-Iubbs 26), Baudelot's ligament ossified (Ch. 34), palatine dorsal pro- (1919). Pattcn (1978) proposed Atherinopsinae ancl Menidiinae cess poir~tedand folded over ethmoid block (Ch. 54), and pres- as pliylogenetic sistcr taxa, a hypothesis later corroborated by ence of a swimbladder haernal funnel (Ch. 89). Characters 6, White (1985) and Chernofl'(1986h). Saeetl & Ivantsoff (1991) 34, and 54 are unique to Atherinopsinae. Additional support- discussed tlie cliffe1-enccs between New and Old World silver- ing characters are presence of a dorsal median ridge on tlie sides, and inlormally grouped the latter wit11 melanotaeniids vomer (Ch. 1),ventral condyle of lateral eth~noidossified (Ch. a~ldph~~llostrthids. Ncw World silversides, or Atherinopsidae, 15),pterotic-exoccipital wings contiguous (Ch. 31), posternporal was formally proposed by Saccd ef nl. (1994) and Dyer & canal with two pores and extl-ascapular with three pores (Cli. Chcrnoff (1996). Saced el rrl. (1994) grouped athcrinopsids 35), vertical and horizontal shafts of preopercular sensory ca- ancl notocheirids as sistcr taxa in a New World s~per~~mily,a nal enclosed by bone (Ch. 61), and pelvic median process re- hypothesis lmsupported by parsi~no~lycriteria (Dyer & Chcrnof1; duced or absent (Ch. 81 ). 1996). The present generic composition of Atherinopsinae was pro- posed by Schultz (1948) and thc first phylogenetic hypothesis of rclationsliips diagnosed the subf'amily with five characters (Chs 11, 15, 34, 54, 73; White, 1985). Chernoff (1986b) pro- Menicliinac co~npris~sscven genera in two tribes: vided two Inore features (Chs 36,71) in support of a monophyl- Mcmblxdini (Atharincdkc,M(:~~bras, Mel~n,orl~i~7us) and Menidiini etic Atherinopsinae, but of the list of seven features combined (Monirlicc, Ch%roslo,ncr,Poblnn,a, LaOide

Fig. 41. Ilypothcsis of' relatiollsllips within atllerirlopsi~linsproposcd by White (1985) (L=404). Uniqrlr ch;rl-actc~-swithin ~~tI~e~-i~rilOr-~rrs211-e opposite black bars and homoplasio~~scharacters indcpcndcntly derived elsewhere are opposite white bal-s. infrapharyngobranchial (Ch. 70), cranial process of were combined in a single data matrix (Tahlc 3). 'I'hc phylogr- infrapharyngobranchial 2 pointed and at right angles to netic analysis using quantitatively-codcd allozynle characters i~lfrapharyngohra~lchial3 (Ch. 71), uncinate process of resulted in two equally parsirnonious trees (Figs 2, 42), 110~- cpibranchial3 in a lateral position and long (Ch. 72), arid pres- ever, the qualitatively-codctl allozyine data resrlltetl in a single ence of a hae~nal-hypophysisfunnel (Ch. 90). All features ex- most-parsimonious tree (Figs 2A, 42A). Both topologies differ cept for Characters 70 and 90 are unique in atherinif'orrns. from White's hypothesis (Fig. 41), thoi~gliadmittedly, the latter Additional supporting characters are 21 wide postcrrlporal ante- is only one step longer than the two trces resulting froin this rior process (Ch. 36), presence of an upper jaw frenum (Ch. analysis (Fig. 2A, 2B). This is an indication that the hypotheses 37), three rows of teeth or less or1 oraljaws (Ch. 50), cliscon- of atherinopsinin intcrrrlationships are not as robust as would tinuous series of in terdols:tl bones (Ch. 84), presence of scales scem from the results of White (1985) and CI-ilbtrcc (1987). on cleithru~nshaft (Ch 99), arid the allozyrnes M-Aat-A (Ch. Morphology alone renders a si~iglctree (Length=JPl, (:1=0.53, 109) and Est-1 (Ch. 112). RI=0.78) with the same topology of' Figures 2A irnd 42A. The morphological data of White (1985) and the distance- Relationships depicted in Figurc 42A show Alhn.in,ops rnore trcated allozyme data ol' Crabtree (1987) both supported a sis- closely related to the Alhurinol,.si.\ - Leur/,.sthe.\ clatlc and (:o@icl~thys ter-group relationship between Athlrmops and (:o@ich,thyc, and is the sister group to that assemblage. The Alh(?-inoj~s,Alh(li;noj)sO, between Athc~in~ol.sisandLeur(!,slhe.s (Fig. 41). The character-state and Leurerthe.~clade is diagnosed by thrce chal-actc~.~(Chs 84, allozy~nedata yielded a slightly dif'f'erent topology in that the 85, 101), and the Ath~(rin,opsi.s- Luur~,sth~,scladc is diagnosed by Al/l.erin.o$sis - 1,ezr.reslhes clade was more closely related to the seven features (Chs 16, 25, 50, 57, 72, 109, 119) of which two Bns%l%chthy.s- Odor~testhes clade, with no significant difference be- anatomical features (Chs 16, 72) and one enzy~r~aticcharacter tween the quantitative and qualitative coding schemes. In the (Ch. 109) are unique. The alternative eclually parsilrionious present analysis, both morphological and allozynie characters cladogram is the mirror irilage of tllc tree described above (Fig. 42R), ill which Ath,erino;l,sisis sistcr 10 the Athr?m'nof)s-Colpichtlzys large scales on cleithrum shaft (Ch. 99), no scales on the anal clade arid I,euresthes is the next sequential outgroup. The fin (Ch. 101), and the allozymes S-Aat-A (Ch. 110), Gpi-A (Ch. A/,hc.r.iiro/~s,Alh(~irr($sis, and Col$ichthysclade is diagnosed by three 113), and Pgm-A (Ch. 121). character-s (Chs 12, 37,40) and the Alhcrinops- Colpichthysclade A unique feature of C. regis is presence of a ring ligament on is diagnosed by six features (Chs 25, 49, 50, 57, 109, 119) oS the palatine around the ethmomaxillary ligament (see Ch. 55). which one is unique (Ch. 49). hubbsi is diagnosed by absence of a haemal funnel (Chs 89, 90) and the first dorsal fin posterior to a vertical from Colpichth,y,~Hubbs, 1918a the anus (Ch. 82). For additional features of Colpichthysspecies see Crabtree (1989). Type species: A/,/LM%I~,o/I.Tr(:~~~Jcnkins & Everrriann, 1889 Distribution: Endemic to the Gulf of California. Colpichthys C:. C:. rep.s (Jer~liills& Everina~lil,1889) hubbsi is restricted to the area of the Colorado River delta north C. hubhsz Crabtree, 1989 oS31° N latitude and C:. .re'ps is found north of 28" N latitude in Diagnosis: oral jaws with asymmetrical bicuspid teeth (see the upper and central Gulf of California. Ch. 49), notch in the cloi-sal process of maxilla (White, 1985), 11r~sence01 a hyomandibular nerve foramell (Ch. GO), few gill Atherinops Steindachner, 1875 ,-.n k,-. <.IS (<20 lower limb) or1 the first branchial arch, external row of gill ralzcrs on antcrior portion of fourth ceratobranchial Type species: Atherinof~sisajjinis Ayrcs, 1860 expanded rnedially in a plate-lilze form (White, 1985), A. afjnis (Ayres, 1860) iilfrapllaryilgobra~richi;~ltcct11 hair-lilze (White, 1985), vcnlral =A. insularurn Gilbert, 1891 extension of anterior borcler of cleithruin dorsomedial wing not =A. magdalenae Fowler, 1903 reaching scapular foramen (Clr. 75), krnale with ovipositor =A. oregoniaJordan & Snyder, 1913. ((:I-abtree, 1989), and eggs lackiilg filaments (Crabtree, 1989). =A. littoralis l--Iubbs,1918b Additional descriptive features: Atherinopsinins with a max- =A. cedroscen,sisHubbs, 1918b illa-premaxillary ligarnent attaclrcd to the ailteroventral corner =A. padnli@ae Hubbs, 1918b of pi-cn~axilla(C11. 47), a subterrninal mouth, presence ofa sec- Diagnosis: Frills oS skin between the maxilla and premaxilla oiiclary t~.igemino~~cialarch (Ch. 25), horizontal shaft of (White, 1985), two rows of subocular scales (Ch. 97), and the prcopci-cular scnsory canal enclosed by bone (Ch. 61 ), a con- allozymc Sod-A (Ch. 123). tinuous series of six or more interdorsal bones (Chs 84, 85), Additional descriptive features: Atherinopsinins with two

Fig. 42. Altcr11;ltivccqt~;llly pal.sin~onious hypo~lreses ol relationships among atherir~opsi~~i~~genera. Unique characters within atheriniforms arc oppositc bl;rcli b;irs; 1.evcrsctl characters (with an "R) are opposite grey bars; and homoplasiorls characters indcpendcntly derived elsewhere arc o1)pc)sitc wl~itcbars. Diagnostic characters ant1 adtlitional support is listed in Appendix 2; A, Co@zchlhyys is sister to Atherinops, Afhm'nopszs, and Lcr1rest11,rc;IS, L/?~rr?.sLlrr~is sistcr to f~lherirro~~tic.Atke~inof~s, and Coll~zchlhys. 48 Mrsc. PLJRI. MIJS.. ZOOL., UNIV. MICII., NO. 185 rows of'bicuspid teeth (cusps synirnetrical or slightly asymmetri- medial process well dcvcloped (Ch. 81), and origin of the first cal) arranged in a single exposed row (Chs 49, 50), gill rakers dorsal fill placed posteriorly to vertical through anus (Ch. 82), 20-25 on lower limb, absence of a t~rohyalventral plate (Ch. and the allozyrne Gtdh-A (Ch. 115). 74), origin ofthe first dorsal fin over anus (Ch. 82), and lack of The feature that most readily distinguishes both species of scales 011 the cleithi-um shalt (Ch. 99). I,euresthe.s is the number of lateral scale rows, with 51-60 in L. Distribution: Ranges from Vancouver Island, B.C., Canada, sardin,aand 69-80 in L. tenuis (Molfatt & Thomson, 1975). White to the Gulf of California. (1985) reported that both species olLe~~r~,sthe,s lack strongly de- Various species, subspecies and their intergrades were de- veloped teeth in the oral ,jaws as adults, and reported the ab- scribed by I-Iubbs (1918b) as a way to delineate the variation of sence or teeth in the cleared and stained specimens he studied. rneristics and body proportions that Athurinops displays along Oral jaw teeth present in the material I studied are minute in L. tlrc West coast of North America. Because no features are known tenuis, whereas in L. sardina teeth are srriall but readily visible. to clearly distinguish one form from another, I consider all nomi- The spawning habits of L. tenuzs are strictly nocturnal whereas nal species to be junior synonyms of Alherin,op.r affinis. L. sardinuspawns diurnally and nocturnally (Moffatt & Thornson, 1975). Atherknopsis Girard, 1854 Distribution: Leu?-eslheslenuis is found from Monterrey Bay south to Baja (hlifornia Sur, whereas L. sccrdina is endemic to Type species: Ath6knopsi.s ccc1iJbrn~iensi.sGirard, 1854 the Gulfol(~a1ilornianorth of Bahia Conccpcicin and Guaymas. A. cal~vrnien,sisGirard, 1854 =A. sonwrcce Osbum & Nichols, 1916 Monophyly and systematics of Sorgeiltillini Diagnosis: Modification oP the ventral process of tlic Inax- illa f'01- the attachirient of'prernaxillary and rostral-cartilage lip Sorgentinini comprises the South Arnericail taxa Ba,silichtlzy,s ments (maxillary anterior pocket; White, 1985: fig. 23A), and Odonleslh.es, a sister-group relationship diagnosed by eight dermosphenotic canal not enclosed by bone (Ch. 20), ptcrotic characters: ethmomaxillary ligament attached to ethmoid car- and epioccipital wing bases contiguous under the epaxial mus- tilage block only (Ch. lo), presence of basioccipital fenestra- culature of the temporal fossa (Ch. 32), dorsal process of opercle tion (Ch. 33), a palatorostral ligament dorsal to the not blade-like (Ch. 64), small scales (Ch. 102),and the allozynies ethmomaxillary ligament (Ch. 56),absence of a11 endopterygoid I'gm-A (Ch. 121) and Sod-A (Ch. 123). tooth patch (Cli. 59), precaudal vertebrae over 50% of total Additional descriptive features: Atherinopsinins with the lat- vertebrae (Ch. 86),presence of an haernal-arch funnel (Ch. 91), eral ethmoid contacting the parasphenoid (Ch. IG), and the allozymes Ak-A (<:11. 107) arid Ap-A ((31. 108). The cpibvanchial three with twisted shaft medial to the uncinate first three morphological features are unique in process (Ch. 72), and absence of a ~lrohyalvciltral plate (Ch. atherinomorphs. 74). The morphological data of White (1985) and the enzymatic Distribution: From Yaquina Bay, Oregon, south to the west data of Crabtree (1987) both supported a sistci--group relation- coast of Baja California Sur. ship between Basilichthys and Odontesthes. The representative material they had available, however, underestimated signifi- I,~ure.sthe~sJordan& Gilbert, 1880 cantly the species and anatomical diversity present in the South Aniericail clade. The present study illcludcs for the first time 'Tjipc species: Alher~n,opsislc~nuis Ayres, 1860 the complete taxonomic diversity ofthe group, with four or Gve =HuDbsiellaBredel-, 1936 species in Basi1ichth~1.sand at least 17 species in 0rlontesthe.s. Type species: iWrnidza clara Everniann &Jenkins, 1891 L. tenuis (Ayres, 1860) Odontrsth(.c Evermann & Kendall, 1906 =L. cramm.Jordan & Evermann, 1896 I,. sardin.a (Jenkins & Evermann, 1888) Type species: Odontasthes Purugiae Everrnann & Kendall, 1906 =L. clara (Evermann & Jenkins, 1891) =Kro~riaMiranda-Ribeiro, 19 15 Diagnosis: Maxilla-premaxilla distal ligament attached to the Type species: Kronia i~paprn~si.~Miranda-Rib- 19 15 dorsal border of the premaxillary alveolar dilation (see C11.47), =Aeudolhyrina Miranda- Ribeiro, 1915 mandibular branch oS A1 muscle tendon present anterior to Type species: Pseudothyrina jhering Miranda-Ribciro, 19 15 lacrimal branch (Ch. 53), hyomandibular nerve divided inside =Austromonidia Hubbs, 1918a of hyomandibula (Ch. 60), pelvic median plate extending to Type species: Rasilicht/l,ys rrgillus Abbott, 1899 anterior tip (Ch. 80), scales present between anterior rays of =Cauqur Eigenmann, 1927 second dorsal fin (Ch. 100), and unique lunar cycle beach- Type species: Chirosloma maubanum Steindachner, 1896 spawning habits (MollBt & Thornson, 1975). =I'atugonia Eigenmann, 1927 Additional descriptive features: Atherinopsinins lacking a (preoccupied; misspelling of Patqonina) vomerine dorsomedial ridge (Ch. I), anterior detachment of' =Pc~tr~go~zin,aEigenmanil, 1927 the nasal sensory canal reduced (Ch. 12),highly protractile pre- Type species: Mvnidia h.atch,eri Eigcnmann, 1909 maxillae without an upperjaw fi-enum (Ch. 37), long prcmaxil- =AustroatherinmMarrero, 1950 (nomen nudu,m) lary ascending process with narrow base (Ch. 38), a well devel- Type species: no type designation. oped dorsal process of the maxilla (Ch. 40), horizontal shaft of =Bachmannia Nani, in Szidat & Nani, 1951 preopercular scnsory canal encloscd by bone (Cli. 61), pelvic (preoccupied) Type species: Rtcsilichtl~yssmitri 12ahille,1929 (Ch. 13),derlnosphenotic medial flange reduced or absent (Ch. =Sorgentinia Pianta de Risso 8c Risso, 1953 22), pterotic and epioccipital wing bases contiguous under tem- Type species: Atlrerina incisn Jenyns, 1842 poral fossa epaxial musculature (Ch. 32), maxilla with condyle =7i1/1a De Buen, 1953 on ventral process (Ch. 42), maxilla-anguloarticular coronoid Typc specics: Alhminicht11y.splalensis Berg, 1895 ligamellt absent (Ch. 46), labial ligament with reduced lateral = R~ciDe B~ten,1953 fold (Ch. 48), dentary coronoid process flat dorsally at level Type specics: Yaci retrol,inni.s Dt: Buen, 1953 with anguloarticlllar coronoid (Ch. 51), dcntary gnathic spine Diagnosis: Mcselhmoid absent (Ch. 9), trigemiual Corarnen absent (Ch. 52), dorsal palatine process spoon-shaped (Ch. 54), enlarged and enclosed by prootic, sphenotic, and ethmornaxillasy ligament not attached to palatine dorsal pro- plc~~rosphenoicl(Ch. 24), ventral and dorsal processes of max- cess (Ch. 55), preopercular sensoly canal between pores one illary head of itbout equal le~lgth(Ch. 41), mandibular branch and two enclosed by bone (Ch. 62), dorsal process of operclc of A1 n~usclctendon present anterior to lacrimal branch (Ch. not blade-like (Ch. 64), postcrodorsal bordcr of' opercle con- 53), opcrclc fenestrated (Cli. 65) with medial lamina between cave (Ch. 68), first spine of first dorsal fin shorter than spine of struts (Cli. Hi),and anterior liaemal spines short (Ch. 94). second dorsal fin (Ch. 83), precaudal vertebrae over 57% of Additional descriptive features: Sorgentinins lacking dorsal total vertebrae (Ch. 86), first caudal vertebra over posterior half laminae on vomerine anterior face (Cli. 3), nasal sensory canal of anal fin (Ch. 95), dorsal head scales with reversed imbrica- greatly detaclicd anteriorly (Ch. 12),secondary trigeminofacial tion (Ch. 96), [our suborbital scale rows (Ch. 97), and arch present (Ch. 25), ptcrotic and exoccipital wing bases coil- interopercle with scales on posterior half' only (Ch. 98). tig~~o~is(Ch.3 1 ), highly protractile prernaxillae without an 11p- Additional descriptive features: Sorgentinins with a single pcr,jaw li-enrim (Cli. 37), prcmaxillary ascending process with ;I median patch of vomcrine teeth (Ch. 8), an isolated, oval narrow base (C:h. 38), labial li~a~nent- broadly folded lateral to mesethmoid (Ch. 9),nasal sensory canal slightly detached (Ch. ~winaxilla(Ch. 48), and scales present between anterior rays 12),lateral ethmoid with bony ventral condyle (Ch. 15).single of a1111Ill1 (Ch. 101). trigcrninohcial arch (Ch. 25), supraoccipital crest bifid (Ch. Distribution: Coastal marinc and freshwater drainages of 28), cxoccipital wing reduced to spiny processes (Ch. 30),widc southern temperate South America. Marinc forms range frorn anterior process of the postemporal (Ch. 36), frenum between tllc southern tip of Tierra dcl Fuego to northern Peru, includ- the forehead and upperjaws (Ch. 37) and restricted premaxil- i11g the ,Juan Fernanclez At-chipelago, in the southeastern Pa- lay protrusion, premaxillary ascending process with a wide base ciLic and to southeastern Brazil, including the Falkland Islands, (Cli. 38), maxillary dorsal process rcduced (Ch. 40), in the sor~thwesternAtlantic. Brackish and Geshwatcr forms maxilla-prcmaxilla distal ligament elongated and attached to I-angc lrom Patagoni;~to northern Ccntral Chile along thc Pa- anteroventral corner of alveolar arm (Ch. 47), palatine spine cific drainages, ilurd to Rio Grandc do Sul, so~~theasternBrazil, not contacting quadrate (Ch. 57), six or more interdorsals (Ch. along the Atlantic drainages. 8.5) in a continuous series (Ch. 84), haemal arches with flaring Comments: This genus is the most spccics rich and widc- expansio~lsonly (Ch. 91), and body scales small (Cli. 102). spread of;~therinopsinegenera, comprizing 20 to 25 recognized Distribution: Restricted to freshwater drai~lagesor the west- species. White (1985) diagslosed Odonlesth~swiththree charac- ern Andes f'rom Lambayeque, northern Peru, to Chiloi Island tcrs (Chs 9,37,65) and remarkcd that this group was in nccd of in souchcrn Chile (Fig. 44). I have neither collected nor en- revision. Of the sevcral juslior synonyms of Odonte.sthcs still in countered lnllseusn specimens that support literature repom use, A~.~.slro~trc.)litliccti~(I-Iubbs, 1918a) and Car~qrre(Eigenmann, of this group's presence in marine environments (Mann, 1954) 1927) are the only taxa comprising two or more species. Other or freshwater drainages east of the Andes (Lahille, 1929; genus-lcvcl names such as Kron,ia (Mirarrda-liibeiro, 1915), Aramburu & hnguelet, 1967). Urzfia et nl. (1977), however, I'atc~go~zinn(Eigcnmann, 1927), Sorgenlinia (Pianta de Risso & reported the occasional capture of Basilichthys in thc Port of Risso, 1953), and fi~ciDe Ducn (1953) are thlls Ihr monotypic. San Antonio, Chilc, by local sport fishermen. Though speci- Thc rclationsliips and systematics of Odon~testhesspeciesand those mens of Bnsilicht1~y.saustralis can s~lrviveaccidental traslspla~lts of thr sltbgrn~~sAzrslronzc!?zirlinal.e treated elsewhcrc (Dyer, 1993: directly from freshwater to seawater aquaria, I regard the pres- 331-590). encc of Rasilickthys in marinc waters as rare and incidental oc- currences. fiasilic!~,t!rys Girard, 1854 Parcnti (1984: 1 10) refers in a footnote to a small specimen of Bnsilzchthys (FMNH 94184: 17.5 mm SL) collected near Lake Type spccics: Atlzwin,a m,ici.ol(:/)idol~csJcnyns, 1842 Titicaca, Peru. Examination of an X-ray plate of the specimen, =Proli~tiusCope, 1874 kindly provided by Parenti, cosifir~nit as Ba.silicht/zy.ys. Odontesthes Typc spccics: Praiisiius se~r~oLil~c.\Cope, 1874 bonccriensis was introduced into Lake Titicaca much later than =C;a,slr@l(rrzc.\Copc, 1878 the 1918 collection date of this specimen. This specimen con- Type spccics: (;cr,slroj)tcrrus /wc.h,aeusCopc, 1878 tinues to be the only record of this genus in the Puno Region =I';sci,regia Abbolc , 1899 and either represents the sole representative of a now extinct Type species: l'iscir(!gia Dearrlsl(:ri Abbott, 1899 populatio~lor spccics, or is a misplaced specimen from a previ- Diagnosis (Fig. 43): Vorner with two slightly elevated ous collection of the Invisl expedition in the Arequipa region. par;tsqittal laminae (Ch. 3), vomer slightly concave ventrally Comments: The species composition of Basilichthys has been ((:I\. 5), a single median tooth patch (Ch. 7),lacrimal condyle problematic ever since its description (Girard, 1854, 1855). of lateral ethmoid does 1101 reach ventral edge of lateral ridge Based on Girard's descriptions alone the name Basilichthys has MIS(..PIIIIL. Mus.. Zoo1 ., UNIV.M1c.11.. NO. I85 BASILICHTH YS

ODONTESTHES

Fig. 43. Monopl~yly01' Ilrrtilirhfl~)~.\,sllo\vi~~g ~lic u~ltlcscl-ibcd spccics 11-0111Kio 1.03 as part of the s(,tr~oliilosljccics $~-or~lj.ITlricl~~c. characters ~vitliin ;~llicrinitor~lrs211-c opposite hl;~c.k1x11s; reversed clr;~l.;~c~c~.s(with :In "R") are opposite KI 'y 11al.s; and liomopl;~siol~rclral-;~ctel-s i~rtlrpc~~~lc~l drl-ivc.tl clscwlicrc arc opposite wlrite b;il-s. 1)i;tgnostic. cll;u-ac.tcrs ~ultl~~clditiollal sul1pol.1 is listctl ill Appe~ldix2. 8 4" 7 6" 6 8"

Q semotilus sps. gr.

0 microlepidotus sps. gr.

8 4" 7 6" 68"

Fig. 44. l)ist~.il~r~tionmap 01 Rnsilirhlhys. Dots indicate localities of' collections examinctl. Sh;~dedal-eas rnark the distrihutioll range of the species grolI[)s. <5" - Mrsc. PUBL.MLIS.. ZOOL., UNIV. MICII., NO. 185 been 11scc1by different a~rtllorseither to group sorgcriti~liriswitli- Typc locality: (:allao, Peru. orrt i~pperjawprotrusion (Jordar~Sc I-lt~l~hs,1919; Eigenmann, Basilirht/t~l.sa)-clzapu.s ((;ope, 1878) 1027; De Buen, 1950; Mann, 1954), sorgcntinins without vomcr- 'Qpe locality: Areq~ripa,Per~r. iile teeth (Evermanu & Ikndall, 190(i), or all sorgentiliins Basilicl~ll~yssp. (I,ahille, 1929; Arambur~~& Ringrrelet, 1967; Ringuelet (?I al., Type locality: Rio Loa, Chile. 1967). Bn,silichtlzy~sis restricted to the western fi-eshwater drain- Diagnosis (Fig. 43): Vorner with dol-so~nedialspi~le (Ch. 2), ages of Chile and Peru, despite soine recent p~tblicationstlrat labial ligament fold greatly reduced or absent (Ch. 48), four co~itintreto follow the old no1nenclat~u.eof Lahille (1929) and pi-eoperclllar sensoly canal pores 011 horizontal shaft (Ch. 63), Riliguelet el ccl. (1 967). A good example of this taxonomic rnis- and head scales of interorbital region only I-eversedin imbrica- ~r~iders~anclingis when thc Patagoriiarl silvrrsitle Oclo~~t(?st/irs tion (Ch. 96). hnlchmi (Eige~linann)is referred to as Ilnsilichlltylys ?nicrolef)iclotus. Additional descriptive features: Basilichlhys species witli an- terior clongaiio~iof basisphe~ioidto interorbital cartilage (Ch. microlepidofus species group 26), Inore than three rows of teeth on oral jaws (Cli. %I), upper jaw relatively more prognathous, cctoptcrygoitl always absent Ba,silichlltys tnicr-o1rj)irlotu.s(Jenyns, 1842) ((3.58), aiid GI-st dorsal fin relatively smallel; sorrleti~nesab- Type locality: Valparaiso, Chile. sent. Brrsilicht1~y.smstmlis Eigenmann, 1927 Distribution: FI-om Rio lieque, l,a~nbayeque(7"S), to Rio Type locality: li-om Rio Maipo to Rio Rahue, (:bile. Sarila, Tacna (22'S), ill Peru, arld in liio Loa, lqrriqlrc (22OS), I Diagnosis: Region, Chile (Fig. 44). Eigenmani~(1927) cites reports of Additional descriptive features: Bnsilichllt,y3 species with ria- Bnsilickl!tys in Rio Loa and Rio Camarones in ~iortherrlChile sal ventral process riot overlappi~igthe sllbnasal shelf (Ch. l l), and refers them to B. s~motilus. Thollgll Ua.tilicl~thjsis indeed labial ligaineilt with a reduced lateral fold (Ch. 48), three rows present in Rio 1,oa no specinieris liave been collected in liio of teetli on,jaws (Ch. 50), and all dorsal l~eadscales I-everscdin Ca~narones(De la IHoz, pers. comm.). The nol-them extent oS imbrication (Ch. 96). the range was previollsly considered as the Kio Rimac, and ex- Distribution: Froin Rio Htlasco, I11 Region (28"-2S0S),to pected to be in the Rio Sarlta (Eigenmann, 1922). The pres- Chilo6 Island, X Region (42"-43"S),(:bile (Fig. 44). Eigeii~iia~i~iciice of Bnsilichll~ysin Rio Santa is confirmed by A. Sifi~e~ltes (1927) cites reports of Bnsilichthps in Rio Copiapci 11r1t lione have and H. Ortega (UMMZ 2 18074), tllc range is extended to Rio been collectctl despite nunlerorls attellipts (De la I-Ioz, pers. .Jequetepeque (UMMZ 218050), and witli r111confi1-mcdreports cornm.). The southern limits of the raiigc was cxtended from of its presence in Rio Reque (R. Alva, pers. cornm.). Collec- Pucrto Montt to Chiloi Island by Campos et ctl. (1984). tions north of Lima, in Rio Sa~lta(Carhr~az) anti Rio San Juan Comments: 1)istinction betweell the two species is pl-ohlcm- (upper Rio,Jequc.tepeque),were made at elevations of2000 1x1 atic. The northern pop~llaiionslook quite different horn the or greater; rvlicrcas collcctio~lsat lower elevations are devoid of southern popl~lations,brrt tllosc ol' thc cc~ltralAconcagrra and Bnsilichl!tj~s(Eigelirnari~r, 1922; Davila, 1973). Maipo river drainages overlap in their counts and rneasllre- Comments: The populatio~~present in Rio Loa is separated ~llelits. Both specics arc vil-tually identical in all allozyme sur- by appl-oxirnately 1000 krns Srom its nearest congcnc~-in the vey of 37 loci (Gajardo, 1988). Morphometrically tlicy are ill- Rio Sama, Tacna, Peru. The Rio Loa popr~lationis an distinguishable or only statistically separated by Incan \dues of llridesci-ibed species .rzihich, based on the PI-climi~iarydata pl-r- some ineristics, namely longi~udilialscale row counts (G;ljrrdo, sented here, is the sister specics to the remainder of the s~rnolilz~s 1987), tlie same feature used by Eige~i~~iarln(1927) in his dc- specics group. This unclescril~edspecies is untlc~-severe threat scription of B. ccristrnlis. Thr only qualitative evidence that ar- of extinction because ofa co~itaminationevent in 1996, the ori- gues for the existence of two species is karyotypic, in which the gin of which is to bc determi~~ecl.This population is so far en- n~odalchl-ornosome il~miberSor I{. microL($iclolus is 2n=4(5and demic to the Kio 1,oa and restricted to a portion of rivcl- dow~~- that of 13. nr~stmlisis 211=48(Gajai-rlo, 1992). The clirornoso~iie stream fiom the city of (hlama and a major copper-mining in- number of Oclo?ctrslh(>.sbona,.i(>n,sis is 211~48(Arai & Koike, 1980) dllsll-y. as is true generally oP atheriniforms such as Menidin, The s~motil7isspecics gl-oup, exclusive ofthe liio Loa popula- n/lclnn,olrtc~nia,and flt1~~rio11(Gial-do, 1992), suggesting the karyo- tion, is diagnosed with scales present bctwccr~the anterior rays typically derived condition is that of B. ~rricrolepidol~cts. of the second dorsal and aria1 fills ((:hs 100, 101). Rnsilichlhys A ~lrln~berof papers have described the kctling (Urzfia el ben~clsleeiwas described based on a single large specimen col- ctl., 1977; Bahamondes ef aL, 1979) and reproductive biology lected from the rnarkct at Callao and assumed to be rnarine (Moreno el nl., 1977; Vila & Soto, 1981; Vila el nl., 1981) of B. (Abbott, 1899). B. archneus is tentatively considered to be dis- acisll-nlis, but only orle of B. microl(pidotu~(Con~tc & Vila, 1992). tinct from B. senzorilz~sbased o~ithe morphology of the jaws, bui DiSferences were found between these species, hut are attrib- so fir neither species has been pliylogenetically diagnosed. uted to tlie very different enviro~imentsand flow regiriies ofthe A relatively large collection was made along the westcr~ldrain- localities compared (Comtc & Vila, 1992). ages of the Andes of Per11 bctwcer~theJeq~tetepeque and Sania rivers in 1988. This collection suggests B. nrcl~arc~.~is possibly semotilzis specics group endemic to altit~rclesover 2000 m i11 the Arequipa region. Bnsilickfhys is fotund only in the ~lpperinostreaches of the Sanca, Bnsilic1~lhy.ss~,rwolihts (Copc, 1874) Chicaina, andJeql~etcpcquc rivers. This suggests that potential Typc locality: Andes, Pel-11. areas of endemism for fresh~raterfishes in the western Andes =R. Deardsl~!ei(Ahbott, 1899) ocur~.in an altitudinal rather than latiti~rlinalgradient. DYER,ATHERINOFSINE RELATIONSHIPS AND BASILICHTHYS 53

IUY TO TI-IE ATHERINOPSSNE GENERA AND SPECIES OF to premaxilla (Fig. 23A); dermosphenotic with one ex- ATHERINOPSINI ternal pore (Fig. 12 B,C); scales large and small (9-15 dorsal scales); first dorsal fin anterior, over, or posterior la. Upper jaw fi.enuiu present (premaxillary protrusion re- to anus; ~iobeach spawning habits ...... Odontesthes stricted; Figs 16, 23 A,C) ...... 2 (coastal marine and freshwater drainages of southern South 111. Upperjaw f'renum absent (premaxillaries freely protrac- America) tile; Fig. 23A) ...... 6 6b. Oral teeth small, minute, or absent; labial ligament folded %(I). Dorsal head scales with reversed imbrication (anterior medial to prernaxilla; dermosphe~ioticwith two external margin free; Fig. 16);four subocular scale rows, and re- pores (Fig. 12A); scales large (9 dorsal scales) ; first dorsal duced labial ligament folded lateral to premaxilla (Fig. fin posterior to anus; beach spawning habits ...... 7 23 B,C); lit-st dorsal fin small, anterior to ailus ...... 7a. Number of dorsolateral scale rows 51-60; diurnal and noc- ...... Hasilichthys turnal beach spawning habits ...... Leuresthes sardina (Pacific freshwater drainages of Andes of Chile and Peru) (Gulf of California endemic) 211. Dorsal head scales with normal imbrication (posterior 7b. Number of dorsolateral scale rows 69-80; strictly noctur- margin (i-ce); two or three srtborbital scale rows (Fig. 23 nal beach spawning habits ...... Leureslhes len.uis A); labial ligaiilent f'olded medial to prcrnaxilla; first dor- (Northeastern Pacific, from central California to Baja sal fin anterior or posterior to anus ...... 3 California Sur) 3>1(2).Twoor three rows of conical teeth on oral jaws; gillrakers on lower liiilb oflirst branchial arch 30 or more; isthmus straight, lacking posterior widening of urohyal ventral DISCUSSION plate (Fig. 31B); Grst dorsal fin anterior to anus ...... Athwinznopsiscalijvmiensis SYSTEMATICS (Northeastern Pacific, from Oregon to Baja California Sur) 3b. A single row of' exposed bicuspid teeth on oral jaws; Taxonomic results obtained in the present study are identi- gillrakers on lower limb of lirst branchial arch 25 or less; cal to those proposed by White (1985), i.c., the tribes and gen- isthmus straight or with widening of urohyal ventral plate era are rnonophyletic. Different results were obtained, how- (Fig. 31) ...... 4 ever, regarding the generic relationships within Atherinopsini. 4a(3).Frills of skin on nlargin of labial ligament bctwcen max- Two equally parsimonious hypotheses of relationships result illa and prcmaxilla; tooth cusps symnletrical or slightly from the data collected in this study (Fig. 2). As depicted in asymmetrical; gilll-aliers on lower limb of first branchial Figures 2A and 42A, Colfiichthys is the sister group to all arch 20-25; two rows olsubocular scales; horizontal shaft atherinopsinins whereas in Figures 2B and 42B,Leure.~the,~is the of preoperc~rlarsensory canal not enclosed by bone (Fig. sister group to all atherinopsinins. Either of these hypotheses 28D); scales absent on cleithrum; isthmrrs straight, lack- is only one step shorter than White's (1985) hypothesis in which ing posterior widening of urol1yal ventral plate (Fig. 31B); Atherinops is sister to Colpichthys and Atherinopsis is sister to first dorsal fin over or posterior to anus; anal fin wit11 scales Leuresthes (Fig. 41). Inability to make the inultistate enzymatic I~etweenanterior rays; females with no ovipositors ...... characters binary (Chs 107-123) precludes an attempt to choose ...... Athwin,o/)s afjnis between the two trees using the successive weighting procedure (Northeastern Pacific, from British Columbia to Gulf of (Carpenter, 1988). Morphology alone and the qualitatively- California) coded allozyme characters result in a single tree with Colpichthys 4b. No frills of' sliirl 011 margin of labial ligament between as sister to other atherinopsinins (Fig. 2A). Clearly the interge- nlaxilla and prernaxilla; tooth cusps asynlmetrical, lateral neric relationships among atherinopsinins are not stable with crisp longcr; gillrakers on lower limb of first branchial the evidence collected thus far and more comparative work is ilrcll 15-19; 1111-ccrows ofsubocular scales; horizontal shaft needed to resolve the conflict of data. ol'preopcrcular sensory canal enclosed by bone (Fig. 17); The list of diagnostic features provided by White (1985), large scalcs prcscnt on cleithrtrn (Fig. 40A); isthmus with Chernoff (1986b), and Crabtree (1987) has changed for the posterior widening of urohyal ventral plate (Fig. 31A); taxa included in this analysis (Appendix 2). These changes in- first dorsal fin antcrior or posterior to anus; anal fin with- clude the addition of new diagnostic characters and the change out scales between anterior rays; fcnrales with ovipositors in status [I-om diagnostic to ambiguously optimized (Chs 15, ...... 5 35, 37, 43, 48, 73, 80) in the present hypothesis. 5a(4).Origin oPlirst dorsal fin anterior to anus; dorsolateral se- The number of derived c11;iracters shared between Ba~ilichthys ries of scales 40-47; haemal f111inelpreselit ...... and atheri~ropsirlinsis striking. Two characters (Chs 15, 36) ...... Colf~ichll~ysreps arc shared between Basilichthys and all atherinopsitiins, and two (Gulf of California, north of 28" N latitude) more characters (Chs 37, 40) are shared among Basilichthys, 511. 01-igin of first dorsal fin posterior to anus; dorsolateral Atherin,ops, Atherinopsis, and Colf~ichtl~ys(Fig. 45). Despite series ol'sc;~les47-57; haeinal f~rnnelabsent ...... Ba.rilichthys sharing four derived characters with the above ...... Colpichthys hubbsi atherinopsin genera, a sister group relationship between (Gulf of California, Colorado River delta north of 31" N Basilic/~t!~ysand Atherinopsini is two steps longer than one of latitude) the most parsimorlious hypothesis prefered herein (Fig. 2A). (ia(1) .Oral tcctll well developed; labial ligament folded lateral Crabtree (1987: fig. 2) presented a hypothesis of relationships 54 MIS<..PUUL. Mus.. ZOOL.,UNIV. MICH., NO. 185 based on character-state trcatcd allozymc data in which not be used to make absolutr statements oS~~ppcrja\vmobility, Sorgentinini is sister to the Ath(l-inoil,,si.s- Lrrire.sthec clade. The but instead, 01'1-elativr amounts ofdisplace~ne~it.Fowler (1903) latter hypothesis is 6 steps longer than the shortest tree found stated clearly the degree of protractibility when he defincd the using the data set of Zxble 3 and is not supported in this study. subfamily Atherinopsinac: "Prernaxil1a1-icsnot Jreely protractile, the ski11 continuous mesially with forehead."(rny italics) <:oupled with absence of a fre~luniin Leuwsrkes and Odont~.~thes,is presence of' long and slendel- ascending processes of thr premaxillae (Figs 4, 17). 'The long and slender premax- illary asceridi~lgprocess was regartled as a derived feature by Presence of a continuous, unfolded sheath of skin between White (1985), whereas Chernoff (198613) consiclcrcd it a the upper jaws and the forehead, or fi-enmn (Ch. 37; Fig. 16), plcsiornorphic feature for atherinopsids. White further argued has traditionally been used as an indicator of the absencc of for the greater proxirriity of the ascending process to the sym- pre~naxillaryprotrusion (Eigenmann, 1927; Schult~,1948; Dr physis in Leureslhes relative to 0do.ntr.rthes as evidence support- Buen, 1950, 1955). While presence of a frenum may bc a good ing the independent origin of these states. White's hypothesis estiniator of upperjaw rnobility in athcrinoids and atheriniforrn of character polarity, however, includes at least three dif'f'erent outgroups, it is certainly niisleading in atherinopsids. The features that are functionally associated to upper jaw protru- amount of premaxillary protrusion in Hrcsilichtl~ysis less than sion: frenu~n,length, and width of' ascending process of pre- t1i;~t of Odontrsthes (De la Hoz & Vial, 1994). Similarly, maxilla. The amount of upperjaw mobility is better correlated i~t/rt.rino~)sis,Athm'~noj)s, and Colf)ichthyshave obviously reduced to length, not width, of the ascending process of the premax- protractibility of the llpper jaw when compared to Lezrrr.sth(~,s. illa. Preserved specimens of C~olpicl~tl~jl~have a greater amount Bn.silichtl~ys,Atl~&ol)s, Athen'nol~sis,and Col11ichthy.sall have a I're- of premaxillary protrusio~ithan C:l~irostornalcbarcc~e, for example, nun, but have signiLicantly more upperjaw mobility when conl- despite the preselice of a f'renurn and the broad ascending pro- pawd to Lnbirle,sthr~sor 1ong:jawcd species of Chirostom,n which cess in the fol-men A long premaxillary ascending process is lack such a f'renum. Presence or absence of a frenum should indeed a plesiomorphic feature within athrri~iopsids(Chel-noff,

ATHERlNOPSlNl SORGENTlNlNl & ATHERlNOPSlNl SORGENTlNlNl

.$ ATHERlNOPSlNl SORGENTlNlNl -UJ ATHERlNOPSlNl SORGENTlNlNl

Fig. 45. Ntcrnative optirnizarion of c1iaracte1.son both eql~allypal-sinionious cladograrris of atherinopsid relationsliips; A, A', Col~~iclr/lrysis siste~ to Athrrinops, Athrrirr o/,si.s, and I,PUr(!\lhes; B, B' , 12~t~~eslh/niis sist cr to 11 Ih(>ri?zojf~\i~,A tl~~ririop~, and (;oljfjirhlkys. 1986b). A prernaxillary ascending process with a slender base independent of the bones they are attached to, the three-pored and presence of a Genum, however, are both regarded as de- postemporal canal of rnenidii~lescorresponds anatomically to rived features in this stl~dy.Neither these derived characters nor the extrascapular(s) of atherinopsines, not the two-pore their functional expression can be un>unbiguously optimized postemporal canal ol'atherinopsines. because ofthe conilict in atherinopsinin relationships (Fig. 45). According to Johnson (1993: 17) the postemporal canal of Presence ofa fi-en11m (C:li. 37) is either ambiguously optimized percoids "...typically develops as an integral part of the in atherinopsincs (Fig. 45A, 45A') or independently derived in posttemporal (i.c., it does not ossify separately and later fuse)". Bn.siliclzl1y.s ancl atherinopsinins except for I~e.u~r~,stlzes((Fig. 45B). Given the postemporal canal so~rietimesossifies separately and Similarly, a slender premaxillary ascending process (Ch. 38) is does not fuse to the underlying postemporal bone, the either independently derived in mcnidiines, Le.ur-eslh~s,and extrascapul;lr and postemporal bones seem to be different by Odon,lcsll~es (Fig. 45A), or ambiguously optimized in definition rather than by non-llo~nology. If this hypothesis is :~therinopsids (Fig. 450, 45R'). In summary, I agree with correct, the extrascapulars of atherinopsines and i\/[Plnnorhintts <:hernoff (1986b) regarding a long ascending proccss as are homologous to the postemporal canal of menidiines. The plcsiomorphic for atherilropsids, and I agree with White (1985) three-pored temporal canals of atherinopsids are anatomically t11;1t a slcncler ascending preniaxillary proccss is derived. All similar to extrascapular two of mugilids, but whether these three- atlierinopsids have some degree of upper jaw mobility, the pored temporal boncs arc homologous is less clear because the ;nnount of which is restricted either by a frenrrm or a shorter position ofthe pores and the shape of'thc bones are not equiva- pre~naxillaryasce~ldi~lg proccss, or both. lent (Fig. IOA, IUD). The homology among the postemporal canal of athcrinopsines, extrascapular three of mugilids, and the posterior cst~-ascapularof L;lcc.s.somn as implied by Johnson & Patterson (1993: fig 14) is an interesting hypothesis that re- The sensory-canal bones of the temporal region are named quires a bettel- understanding of the development of these according to the bones with which they are associated (i.e., bones, and corroboratio~lof Elccssonza- mugilid - atherinomorph ptcrotic, postemporal, supracleithral). Extrasc;-rpularbones are rclationsliips. c:lnal-bearing bones not associated with other bones, irnbetl- Presc~iceof't~vo extrascapt~lar bones was hypothesized as di- ded in the slii~lusually between the ptcrotic-canal and the agnostic of sorgentinins by White (1985). Separation of the postcmporal-canal boncs. As defined above, mugilids have three horizontal and vertical elements of the extrascapulars is vari- cxt rascap~~lars(Stiassl-ry, 1993: fig. 7D), cyprinodontiforms and able in sorgentinins s~~cllthat both states may be present on the bclonilbr~nshave no temporal canals, sorne ;~tlicrinoidshave same individual, therefore ktiling the co~ijunctiontest of ho- temporal and postemporal canals (mela~lotae~iii~ies,atherinids mology (Patterson, 1982). Both elements are consistently filsed [Fig. SOB], ancl D~nlnth(.rir~n),atherinopsids have postemporal together in atherinopsinins. Assuming the extrascapulars of canals, b11t only athcrinopsines and Melnno~hinushave atherinopsines and the posternporal canal bone of rnenidiines extrascap~~lat-s. arc homologous (see above), and considering the presence of Extrascapular bones of atherinopsids are formed by a me- separate clerne~ltsin Melnnor-hinus and some specimens of dial ~ra~isversally-orie~lt(:clcanal and a lateral liorizon~ally-ori- MrmDm~mar-liniccc (UMMZ 199183), two separate extrascapular cntcd c;~n:ll. These bony canals are either fused into a single elements appear to be the plcsiomorphic conditio~rand the three-pored bone, or are present as two separatc bones in presence of f~lseclelements in athcrinopsinins would be a dc- illol~~1701-li~iri~us(Stiass~iy, 1993: fig 7C) and sorne sorgenlinins (Figs rived condition. I I, 32; White, 1985). Stiassny (1993) mentioned the prcscucc ol'cxtrascapulars in ho,b11t I have been tunable to find any such Axial skeleto~i ossifications in the material at hand and Said (1983) did not niention them in the ostcological dcscripiio~lof the genlts. Athcrinopsinins have between 10 and 15 vertebrae with so Tlic postemporal calla1 oP atherinopsids is different in called "precal~dalhacmal arches", i.e., haemal arches anterior ~rlenidiinesant1 in atherinopsines. Mc~iidiincshave ;I to the first vertebra that bears a haemal spine. "Prccaudal haernal ~~ostem~x)~-alcanal similar to thc cxtrascap~~lardescribed above arches" are ventrally directed parapophyses of the vertebra with (thrce-lx11-edpostemporal canal), whereas athcrinopsi~leshave a hacrnal bridge separating dorsoventrally the hacrnal canal from a postcmporal canal wit11 one pore at either end (two-pored the swimbladder. The paired projections ventral to the haemal posternporal canal). The postemporal canal of nienidiines has bridge, or hacmal hypophyses, seem to correspond to the ven- the same form and spatial orie~itatio~ias the three-11orccI tral ends ofthe haemal parapophyses rather than being homolo- extrascapular hone of' atherinopsines. Stiassny (1993) consid- gous to haemal spines as proposed by White (1985: 6). The crcd the sensory-canal botlcs ol'iVIekcnorhin~tsas extrasc;rpulars, possible l-lornology of these haemal arches raises a number of IIII~ when compared to other rnenidiincs it is clearly the issues the very least oPwhich is the coding of Characters 86 and postemporal-canal bone tl~atis actually detached from the 95. The most fundamental issue raised is the arbitrary and con- postemporal. Such "detachments" do occur occasio~iallyin tradictory deIinitions of what constitutes a precaudal or a cau- other t;txa, as seen in some cleared and stained specimens of dal vcrtcbra. "The first caudal vertehra...", as defined by Hubbs /\/l~nlbm,smrrrtin,ica (UMMZ 199183) and 0donte.sll~eshnlclzrri (CAS & Lagler (1958: 24), "...is the first vertebra bearing a definite 12(599), in wl-lich the canal bones are partly or fully disassoci- hc~nalspiue." The earliest usage I for~ndof this definition is ated from the poste~nporalon oue or both sides ofthe head. If that of Ford (1937). Romer on the other hand, defined caudal the canal-bearing hones of menidii~lesand atherinopsines are vertebrae for fishes "...by the presence of hemal arches in the comparcd bascd on their ~norphologyand relative positioni~~g, tail" (liomer & Parsons, 198(5:183). Normally these two defini- tions coincide on the sarne vertebra as in sorgcntinins and Basilichthys srntoti1u.s species group comprises R. .sentotilus and B. ~nenicliines,such that the first vertebra with a lraemal arch also nrr;hauus frorn Peru, and an undescribed species fi-orn Rio Loa has the first haemal spine (Figs 35-38). It is in the case of in Chile. The undescribed species is tentatively proposed as atherinopsinins (Fig. 39), most atherinoids, and many other- te- sister to R. seniolilus and B. archams. Irosts, llowever, that the definitions are in conflict with each other (Ford, 1937; (:lothiel-, 1950; Patten, 1978). An arbitrary choice between definitions af'S(:cts tlie coding of some charac- ACKNOWLEDGMENTS ters and ultiinately the phylogenetic analysis. Choice of defini- tions must come f'rom an assessrnent of similarity among the I wish to thank the nlcnrbcrs of niy dissrrtatio~lco~nmittee that en- dured the early versious of this Icngthy c1iaptc1-from nly thesis: Barry co~nponentparts of'this vertebral "puzzle", which must be based Chernoff, IWillia~nL. Fink (cl~air),Janles S. Diana, Arnold G. I(lugc, on sound morphological arg~rments,and iilclrtde developmen- and G.R. Smith. I also wish to t11;lnk the many Iielpf't~lsl~ggestions pro- tal ;rind comparative studies. "Preca~tdalhaernal arches" are vided by the reviewers that greatly enhanced this paper: Clyde Barboul-, present in mugilids, atherinoids except for notocheirids, Walter Ivantsoff, Lynne P;u-cnti, and Brian White. The followi~lgindi- n'lulrinorhinus, some species of Athnin,rlln, and atherinopsinins. viduals are thanked for their 1o;in or transfkr of spccimrns: William E Ontogenetic information is unavailable at this time to make a Smith-Vaniz and MSlliani (;. S~ILII(Academy of Natural Scicnccs ofP11ila- conviilcing argument as to whether "precaudal haemal arches" delphia), Melanie Stiassny and Carl Ferrilris (Amrrica~lMuseurn of sho111d be considered modified preca~tdalvertebrae or ~nodi- Natural Histoty), Williatn Eschn~eyer((hlifbruia Academy of Scietlces) , Atila Goszto~lyi (Cent]-o Nacional Patagbnico, Argrntina), Marlise lied caudal vertebrae. Defining caltdal vertebrae sensu Romei- Benvcnuti and Joao Vieira (Funcla~aoUnivcrsid;~dc do Rio Grand?, R.S., (presence of'haemal arclies) for Characters 86 and 95, contrary Brazil), Brian White (Los Augelcs County Muscum), Waltel- Iva~ltsol'f to the more genel.al definition used by ichthyologis~stoday (pres- (Macqmrie University Anstrali;~),1,ic. Moues ;lnd I.uis Arnaro (Mnsro enc? of' haenial spines), results Si-om my understanding that all Nacionill Historia Natr~ral,Montevideo, Urug~~;~y),I-Icrndn Ortcga ;und haemal arches are serial Ilomologs, irrespective of the position students (Museo de Historia Natul-al :Javicr Prado', Peru), Roberto ol' the first haernal spine, which is an ontogenetically indcpen- Menni arid Mirta Garcia (Museo de La l'lata, Argentina), Carlos I .ucena dent element (Mabee, 1993: Gg. 10). and Lui~R. Malabarba (Museu de Ciencias da l'.U.C.R.S., Brazil), Naer~ioMcnczcs and Jose Lima Fig~~eiredo(Museu de Zoologia, Uni- vcrsiclade dc Sio Pa~rlo,S.P., Brazil), Ihrsten Hartel (Museum of Corn- parative Zoology), Victol- G. Springer and SIIS;IIIJewctt (United Stares CONCLUSIONS National Museum), Victor Hugo Rui7 (Univcl-sidad dc Concepcihn, Chile), Eduardo cle la Ho7 (UnivcrsidildCatolica dc Valparaiso, Chile), 'I'he phylogenetic hypothesis of' relationships ainong Don But11 (University of (:alifornia, Los Angeles), and Dr. Vaz-Ferreira atllei-inopsine genera proposed by Wliite (1985) is I-eevalnated. (Universidad Nacional, Urngr~ay).I wish to express my gratitude to Anatornical data of White (1985), C:Iicrnoff (1986b), and Dyer James Albert, Reeve Bailey, Douglas Begle, Paulo Buckup, William Fink, (1993), and eiizy~naticdata of' Crabtree (1987) are combined Sara Fink, Martha Gach, William (;osline, Waltrr Iva~~tsoSf,/i\noltl Kluge, into ;r single data matrix of 123 cllaracters and 25 caxa. Taxo- Rohert Millel; Gerald Smith, a~rdMark Westne;~tfor helpf111discussions dl~ringthe drvelopnient of this study. I especially want to thank Cliris- nomic results proposed by White (1985) are unclianged in this ti~ie.Joh~lsonfor her support, timc, patience, and expertise in the pro- study (Table 2). Phylogenetic I-elationships arnong duction of the beautifi~lcomputer-generated figures. I gra~efi~llyac- atherinopsinin genera, however; are resolved in two equally par- knowlcclgc the Armor~rFellowship of the Field Mnseurn of Natural His- simonio~rshypotheses (Figs 2, 42). C:olpichthys is the sister to tor): the Jessup Fellowship Iro~ntlle Acadeniy of Natrlral Scicnccs of Atli,er/n.o/).s,Atheri~zo/)si.s, and Leuresthes, the latter two each others Philadelphia, the Museum of Zoology and 1)epartmcnt of'Rioloby Grants closes^. relatives (Figs 2A, &?A).Tlie alternative hypothesis mir- 1.1-omthe University of Michigan, and the l~ulbrightScholarship Sol- rors the above topology with L~u.rest1zc.c.as sister to Alhennoj~sis, making this st~tdypossible. Finally, 1 \vant to sincerely thank Uo~lgli~s Alh(?rinops, and Colpicht!~,ys (Figs 2R, 42R). 111 the hypothesis pro- W. Nelson for his irrvaluablc help in all curatorial matters rclating to this study. posc~clby White (1985) Colpicht11,ysand Atl~erino/~.s,and L~IL~Ps~II~P.~ and Alhcrinopsis, are sister taxa. Crabtree (1978) presented an altcrnativc to Whi~e'shypothesis in which Sorgentinini is sister REFERENCES to the L~u~.eslhi;.s- Atherinopsis clade. White's hypothesis is one step longer than tl-rose presented in this study and Crahtrre's is Abbott, J.F. 1899. The marine fishes of Pet-u. Proc. Acad. Nat. Sci., seven steps longer. Philadelphia 51 (2):324364. Atherinopsinae is diagnosed by eight charactcrs, Arai, R., and A. Ibike. 1980. A karyotypc stucly of two species of fresh- Atlierinopsini by seven characters, and Sorgentinini by eight water fish tl-ansplantedillto Japan. Bull. Nat. Sci. Mus. Tokyo, Zool., ch;~racters(Fig. 1) . Atherinopsini has Sour genera: C~olpichthys, Scv. A 6(4):275-278. with two species (C. regis, C. huhl~si),is diagnosed by nine char- AI-arnburu,R.H., ant1 R.A. Ringuelet. 1967. Los pejcrreycs cle agua acters; Atherino/)s rEffinis is diagnosed by tllree characters; dulcc. Asuntos Agrarios 1 1-1 4. Atl~orin,o/)siscnliJiirnimsi~ is diagnosed by seven characters; and Arratia, G. 1981. GCneros dc pcces dc aguas continentales de Chile. Puhl. Ocas. Mus. Nac. Hist. Nat. Santiago tlc Chilc 34:3-108. I,~~~~t~~s'Slh.es,with two species (L. renuis, L. sardinn), is diagnosed Ayres, W.O. 1860. (on new fishes from the CaliSo~-niacoast) Proc. by live characters. Sol-gentiniili lras two genera: Odonlr.slhes, with Calif'ornia Acad. Sci. 2(4):52-59.(Not seen) about 25 species, is diagnosed by scven cllaracters, and 13ahanlondcs,I., D. Soto, and I. Vila. 1979. Hibitos alirnentarios de los Bnsilichtl~y.~,with five species, is diagnoscd by 22 characters. pejcl-reycs (Pisces: Atherinidae) del Embalse Rapel. Chile. Meclio Two species groups are recognized in Snsilicktliys. The Anibie~i~e4(1):3-18. Hnsilichthjls mic;rol.j,idolz~s species group comprises tlie nominal Barbour, C.D. 1973. ?'he sysrcrnatics and evolution of the genus species B. ~nicrolepidolusand B. auslmlis from Chile. The Cl~ir-o.rlomoSwainson (Pisces,Atherinidac). Tulanc Stud. Zool. Bot. 18(3):97-141. con descripcion de nuevas especics. Bol. Inst. Oceanogr., Univ. Slo I'wrg, (;. 1895. Enunicraci6n sistemitica y sinon61nica dc Ins peces de Paulo 4(1-2) :3-80. las costas Argentina y IJruguaya. An. Mus. Nac. B11cnos hires 4:l- . 1955. Contribucicin a la ictiologia. VIII. El Odontesthes rep({ 120. latzclavia y ensayo de distribution genCrica de las especies chilenas. I

, 195 1. A~lalysisof the Fisl1c.s ol'(:lrilc. Rev, (:l~ile~laI-list. Nat. , aiid.j.0. Snyclcl-. 1913. Description of the Yachats "Smelt," a 51+:3:263-32(i. new species of athel-inoid lisli frorn Oregon. PI-or. U.S. Natl. MLIS. (:. ;+l.xrtlo,. G.M. 1987. Uu.silic.ll/ll~~rrrricr-o1/~/1itl01ii.\ (Jenyns, 1842) and 45( 1999)575-576. Uo,silic.l~/liysnric/rnli.sEigcn~nann, 1027 (Pisces: ?'eleostci, At11c1-i~iitlae): Klink, A., and R. Ecklnann. 1985. Age and growth, feeding habits, and comlnclrts OII their rirorphologici~ltlillerc~ltiatio~l. Strld. Ncotropi- rcpl-oductio~lof (;/rrrquc 17iciril(~(znurn(Srcindac1rnc1- 1896) (Pisces: cal I';l~ui;~I'.nvil.o~r. 22(,1) :223-236. Athcrinidae) ill so~~thcr-nChile. Str~tl.Neotropical Falun;~El~viron. --- . 1988. (;cnc>tic divergcncc alitl speciatioll in lj(r.\;/ic./~thy\ 20 (4):239-249. rrirerol(pidot~~.\Jtt~~y~ls,1842 and li. rrrrstr-(rlic Eigenrn;lnn, 1927 (Pisces, Lahillc, F. 1929a. El pcyjct-I-cy. Bol. Mill. Agric. Nac., Rucnos Aires Athrl-initlac). C;elretic;l 76:121-12(i. 28(3) :261-395.

, 1992. Iurd the analysis 01' (:oinmissio~lsrcalller Albatross. XXII. Dcst riptiolls of'thirty-I~NII-new nrultistatc cliaracters. (:ladistics 8(1):45-65. species of' lislics collrctetl in 1888 ;und 1889, principally among tlic Mahee, l'.M. 11)03. Pllylogenetic intcrpl-ctatio~iolo~ltoge~retic change: Sant;~R;\rb;l~-;~ Islands and ill tlre C:l~lfoS(:;~liSo~-nii~.I'roc. I1.S. Natl. sorting our the actl~itlfro111 tlie artcfactr~;~lin a11r~~ipirical case study MLIS.14(880):549. of ccntrarchitl lisl~~s.Zool. J. 1,inn. Soc. 107(:4):175-291. (-.II'~I~, .. . (:. 1854. t\hstn.act or a report to IAicut.,Jas.h4. (;illiss, U.S.N., Madtlison, W.P., M,I. Donoghrlc, ;uld D.R. Maddison. 1984. O11tg1-oul) ul~o~ithe lishcs collectccl tltu-ing tlic [J.S.N. Astronomical Expedi- an;llysis and parsinlony. Syst. Zool. 33 ( 1) :83-103. tion to (:llili. Proc. i\c;rcl. N;it. Sci. Plrilatlelphia 7(5): 197-199. , and 13.R. Madtiison. 1992. Mac(:lade. Analysis of phylogeny

, 1855. Fisllcs. Irr .l'hc IJ.S. Naval Astronornic;ll I.:xpctlitiol~to ant1 chai-artrr evolution. Vcl-sion 3. Si~iauerA$soc.,Inc., Sl~~ndcrla~itl, tlic So11tller11Hrmisp11c1-c, during the years 1849-'50-'51-'52. 2:230- M.ass,~cllusetts. . .. %:'I. Ma1111,G. 1954. Vitl;~de 10s peces clr agtias clrilenas. Illst. I~rvest.Vetel:, (;obbi, M.11:. l!)S(i. Oslcologia t1c.l pc-jcrl-cy ~)atagti~~icoI'rrlc~gorr?ricr Sultiago dc Chile, 342 pp. I~cr/c.ho.i,(Eigcn~na~r~r, 1009), Eigc~ima~i~i,1927, L. ~e-l.uan,Rio Nc- Markle, D.F. 1980. Aspects ol cllaracte~-homolo~y ;and pllylogeny of gi-o, Arge~ltili;~.Ut~ptrblishcd tlicsis, (:e~~tro1iegion;ll IJt~ivc~-sitai-io the Clatlifo~-mes.I11 (;olre~~,13.M. (cd.), Papel-s on the syste~naticsof B;~riloclle,Univcrsid;~tl N;rcio~i;~l tlcl (:olrraIrue, Kio Negro, Argcll- g;xtlitc)rm lislirs, Nat. Hist. MIIS. 1.0s Angrlrs Sci. Sec :32: 59-88. ti11;1, I29pp. Marrcl-o, A. 1950. Flecllas dc Plat;\, ;tthcrinitlos argentinos, pcjerrcycs (;oodricll, G.R. 1030. Stt~dicsoil the strucrurc ant1 dc\~clop~ilc~n~of y latcrinos. B~~eiiosAires, 157 pp. ve~.tcl)ratcs.Univ. Chicitgo Press, 1986 edition, 8:37 pp. McII-dcrI'crcrsoccs. Proc. U.S. Natl. Mus. 22(1179):1-10. Srci~lclacl~~~cr-,17. 1875. Icl~tl~yologiscl~ebietrage (111). Be~r~erk~ctlgen iibcr S(,,r.rrn~usnrbnlo,su.t 1u1d S. clrrlhrnlus sp. Gird. Sitzungsben Altatl. Matem'nl exnminec]: Institutioilal abbreviations follow Leviton et Wiss. Wicn 72(1):1-68. (11. (1985) except Sor MNHN-M (Museo Nacional Historia Natu- . 1896. 0hc1-nvri t1cl1c (;l~iroslo~no- Arterl aus Chile. Airrl. ral de Moi~tevid~o,Uruguay) and ZVC-P (Zoologia de N;rt~~rhist.1-lof11111s. Wcin 11 (2):231-232. Vcrtebrados - Pcccs, Facultad de Huma~iidadesy Ckncias, Uni- Sti;lssny, M.I.:J. 1986. The limits and relationships ofthe acanthoinorph versidad Nacional, Montevideo, Urrlguay) . The number of speci- tclcosts. J. Zool., Lond. (B) 1:411-460. mens studied are in brackets, 'st.' indicates stained-and-dissected . l<)<)O.Notes on the anatomy and relationships of the fishes or specimens, and the nu~nberof cleared-and-stained specimeiis Macl;lg;rscar, with ;I taxo~lo~nicrevision of the gerlBrls Rhrocles are in parcnthcsis. Genera and species a]-r listed alphabetically (Atlierinolnor1>li;1:Betlotiidac). Amen Mus. Novitatcs 2979:l-33. witliin each rnajol. gl-o~~pirig.Brief locality data is added for FMNH 77485 (2); Alherinrcson h(,f)~~toid~sUMMZ 21 2576; i~therinopsineso~ily. Ath,m.n,osonza rnic.rosloma UMMZ 21 2562(3), UMMZ 212581. Atherinomorinae: Alhm'norn,oru~eendmchl~.nsis UMMZ 196221, M~lgilidi~e:Agonos/oniu.s ~noizticolaUMMZ 178460(5), UMMZ UMMZ 203854(3) ; Atlzrri~zornoru~lnc.~t.no.s~u.s FMNH 2 1105 (2), 199463 [IG+ (3)1, ANSP 122356 (2), FMNH 4623; (;i-~ninzug--ll FMNH 63916(2), FMNH 21 198(2); Atl~~~ri~~zo~noru,~slif~os crpnilnl~isANSI' 114275(1) ; Jolurus Pichardi ANSP 99859 (1); UMMZ 199482, UMMZ 174207(2); Hyf1oatheri77,c~bleelzeri Lizn nr;~.e,n,li~nANSP 135485(4); Mzcgil currmn UMMZ FMNH 52131 (l), FMNH 57459(1); Hj1f)ontherincr Is'urug(l(, 209709 (3), ANSP 128030 (3); Mugil du~s,sumieriANSP UMMZ 142801 [l st.] ; Slenathe-tinci oval(~u,nUMMZ 2 12578 ( 1) ; (51 891 (1); Mugil trichorlmn ANSP 145654(5). Stenatherinn reginn FMNH 40475 (2), FMNH 23504(2) ; Beloniformes Stenatherinn lern~nincltiFMNH 441 36 (2). Adrianichiliyoidri: Horcrichll~ys~etnuei UMMZ 201 181 (I),ANSP Craterocephalinae: Crat~roct~1halzts,/I1~71ili\UMMZ 212564(2) ; 157315 (4); Olyzins jrcvarricu.~ UMMZ 146566(10), FMNH Crutwocephalu.~pcucirad%alus UMMZ 2 14554 [5] ; ii8669(5) ; Oryzici~1atipr.s UMMZ 146592(10),UMMZ 218498, Atherinopsidae FMNH 58732 (5); Oryzins rnrlnsligmcc UMMZ 232000 (3). Menidiinae Exocoetoidei: Arrl~rirn,f~husl~revis UMMZ 100333 [3 st.], FMNFI Menidiini: Chirosto~naurge UMMZ 197622 (6); Clriroslomcr ch(~f)cil(~~ 47553[3]; Chriorlo,rz~.sntherinoitlrs UMMZ 102192 [9+1st.], UMMZ 197612 (4); C;hiro,storna co~zsociurnUMMZ 179720 (1) ; UMMZ 143079(4); C:olol(rbir snirn UMMZ 71 135[2 st.],FMNH Chirostmtnn cstor UMMZ 197650 (2); C~liirostomahu,rnboldtia~~;urn 63758 ( 1 ) , ANSP 88978 (1) ; Cypselurus rnelnn,rirus ANSP UMMZ 97663(1); Chiro.tlom,a jordani UMMZ 197602(5); 50519 (3); Cyj~srlztrussturksi UMMZ 2 12929 ( 1) ; Derrnogenys Chirosloma labarcnu UMMZ 193463(4) ; Clziro.stomcc lucilts pusilln UMMZ 195917(4); Fodintor arulus UMMZ 190985 [2 UMMZ 179717(1); Chirostorna m(:zquilul UMMZ 21 1095(3); st.], ANSP 88928(1); Hemirarnl,hu.s hnlno ANSP 11 1444(1); Chiro~tonrcipromelas UMMZ 193465 ( 1 ) ; Cl1~iro,stom,a.sj)h~~racwr~ Hir.rrnrlirhtliys a/;finis ANSP 14973( 1) ; Hyf~or.harnph,usne&rtus UMMZ 186203(1); LaOidesth,rs sicc.lilus UMMZ 231 999 (1) , ANSP 48851 (2);Hyporlinrrcphus ~unifi.scintlisANSP 11 8641 (5); UMMZ 222857(3); Menidin colei UMMZ 196562(5); M/.nidh I'cr(:xoc.oetu.s brac/~g$twusANSP 144887 (1 ) ; Sco.nrberc.sox saurus beryllina UMMZ 203149(5); Mpnidia m~nidiaUMMZ FMNI-I 48552 (2); S/r.ongylur-a tnarincc ANSP 1161 70 (5); 136589 (3); Poblnna alchichicn UMMZ 192371 (4). X(,n~ntodon cmnriln ANSI' 123993(2); X(>n,c.ntodon~p. UMMZ Membradini: ilthrr-lnelln al~)nreziUMMZ 19(5439(6); Alhc.rinplln 2185 10 (2); Zennrc.hof)l(~rcrsdispar ANSP 79674(2). argentea UMMZ 202408(4); Alherinella hrasilirn,sis GCIZL Cyprinodontiformes V73: 10997; Athrrinelln cryslallina UMMZ 1711-190 (3) ; Alherinell(i (~yprii~odontoidei:Adi,nia xeniccr UMMZ 158860(5); illfar-o guatem~nlensi.~UMMZ 173523(5) ; Alhei-i~trllasallei UMMZ crtllrcrlus ANSP 104371 (4); Ana11lep.s dowei ANSP 91678 (2); 184767(3); Atherinella sckullzi UMMZ 184709(6), UMMZ il~hyos~mionaustrale ANSP 141548 (4); APlocheilichthys 191736(6) ; iMelanorlzin~usw7icrof)s UMMZ 198779 (1); Menzbrrrs sflilnzcchen ANSP 38679-88 (2); CI~n$alic.h/h~yspardalis UMMZ gilberti UMMZ 202405 ( 1 ), M~,nzr',rnsnccrrlin,iccr UMMZ 202427 (4); Cyprinodmn variqalus ANSP 9 5086 (5); Fundulus 199183(1). o'ici/~hnnu.s ANSP 90980 (4); l~c~tzdulzrshe/eroclit,us ANSP Atheiinopsinae 1 1657(5(5); Fundulus notntus UMMZ 209055 (5); Lzicnnin goodei Atherinopsini: Athet-inops clffini.7 UMMZ 7221 2[33+(2)],Newporr ANSP 140561 (5); Orestices agassiiFMNH 77376(2); I'rofundulus Beach, California; UMMZ 176309(5), La Jolla, Sail Diego, gu:-uatenrnlen.sisUMMZ 21851 3 (2); Profundulzis 1nbiali.s ANSP California; UMMZ 141158(3),Morrow Bay, San Luis Ohispo, 64776 (4); Prof~rntlz~lushiklebmndi UMMZ 157634 (5). California; UMMZ 232001 (4), Catalina Harbor, CaliSol-nia; Aplocheiloidei: A(11iyosern.ion plar(?ANSP 141552 (5); I

ATI-IElUNO1'SII)AE MENIDIINI +Ch. 18: postorbital sphenotic process wide. +Ch. 22: derrnosphenotic median flange absent. +Ch. 19: clermosphcnodc not articr~latedto postorbital pro- tCh. 59: endopterygoid teeth absent. cess. +Ch. : Chernorf (1986b): fused upper hypural plate. +(:li. 20: dermosphenotic sensory canal enclosed. -Ch. 50: three rows of teeth on oral,jaws. lacking filaments; few gill rakers (<20 lower limb) MEMHIIADINI on the iirst brancliial arch. +<:II. 60: lryo~ria~ltlil,ular- nerve tlivitlctl illside oi -C11. 17: nraxilla-prernaxilla distal ligalricrrt attached to hyo~n;rntlibula. anterove~rtralcorner of pl-crnaxillary alveolar arm. +(:11. 70: infi-apharyngoI~rir~~cI~i;~l1 absr~lt. -Ch. 84: a continuous series of interdorsal bones. -Ch. 85: six 01-more interdorsal I-joncs. 1.: (:lier~loff(198611): rostra1 sensory sy~t~~ii;dorsal pro- cess of~riaxillaexpanded and rlongrtr; step-shaped -Ch.101: no scales on the anal fin. n~esetlin~oitl(scc <;h. 9); thick and short epilleural -<:11.110: Aspartate aminotransferase (S-Aat-A). I. -CIi. 113: (;lucose-6-pllosphacc isomel-asc (<+-A). -Ch. 121: Pliospl~ogluco~l~~~tasc(Pg~n-A). ATI-IElIINOPSINN, 1 i: ctlrmoid c;rrtilage folded vr~~tralofvo~llcr lliidlinc. ATIYk,'II2 dorsal scales). I.i I : vertical ant1 horizontal shaft ofpreopcrculal- scllso~y tCh.121: Pliosplioglt~comr~tase(I'gm-A). c;l~ralenclosed by hone. +(:11.123: Superoxide dismutase (Sod-A). I.: White (1985): ~nodificatioriof' the vent121 process .ATMERINOPSINl of the ~naxillafor the ;rtlacliment of pre~uaxillary 1.3: vo~ircrwith two well developed dors;rl I;r~irinae. and 1-ostral-cartilage ligarncnts, +C;ll. 2 1 : dcrnrosplrenotic with three sensol-y-c;uial po1.c~. ":Cli. 74: absence ofa ~11.ohyalventral plate. +Cli. 28: st~pl-aoccil>it;dcrest "trifid". -tClr. 70: inf~-apl~a~-yngoI~r;r~iclii~~l1 c;u-tilagino~is. I,I<~JIil.~.STHI+.'kS tCh. 7 1 : criuii:ll procrss of ilif~-apl~ir~.y~lgolj~.;r~~clrial2 witli I.5 ~nanclibularbr;rnch of A1 ~riusclctendon anterior s~n;~llcondylar stu-tace anti at right angle with that to lacrimal 111-anclr. of infraplraryngol~~"~1~ic1iiirl3. +Ch. 60: 1ryomancIib~1la1-ncrvc tlivided inside of' +Ch. 72: 1111cinateprocess of'e~~ib~-anchial3 long and attacl~etl hyoriianclihtrla. laterally. +(:11. 80: pelvic ~nedianplate extrndi~igto anteriol- tip. +<:h. 90: haemal-hypophysis liulliel. +(:h.100: scales prcscrit bctwee~ranterior rays of' second dor- -(:li. 50: three rows of' teeth or less OII ora1,jaws. sal fin. -Cli. 84: intcl-dorsal series discontinuous. tCh. : Mofht 8c Tllo~llson(1 975) : tu~iiqt~rbeach-spawni~ig -(:11. 99: large scales present or1 ~l~itlir~~rnsllaf't. habits. -Ch. IOI): Aspart ate a~ninotl-ansf'erirse (M-Aat-'1). -Ch. 1: lack of a volr~eri~rctlol-so~nedial ridge. :I:Clr. 36: po~t~lnpol-alantel-iol- process wide. -<;11. 12: rcdtrcctl anterior detachment of' nasal sensory ca- j'C1r. 37: p~'ese~icvof' an ~~ppc:r]awli.c~lrlrri. nal. :Wh. 1 12: Estclxsc (Est-I). -Ch. 37: no uppe~jawfi-elrt~rn. -Ch. 38: PI-emaxillary ascending process with rial-row base. (,~~~I~l'I~~l~~Y-lYLS -Ch. 40: well developed tlorsal process of the niaxilla. +Ch. 60: 111-csence of a 11yo1nalidibul~~1-rrel-ve fbralrie~i. -Ch. 81: pelvic riicdial process well developed. tCh. 75: ve~~tralextc~isio~l of' anterior 11ol.dcr of clcithrtrm -Ch.I 15: Glutamate dehydrogenase (Gtdh-A). dorsomcdial wing not reaching sc;rpr~larforamen. *C:li. 82: origin of first dorsal fin ovcl- anus. tCh. : Todd (1976): teeth with asymmetrical cusps; Mqiite (1985): ~lotclrin tllc tlorsirl process of maxilla, ex- SO1ICENTININI tel-nal row of'gill ralicrs o~ia~ltcrior portio~l of foul.tli 1 I: cthmo~naxillaryligament attached to eth~noidcal-- ceratobra~lcliialexpalided ~liediallyin a plate-likc tilage. f'or~ii,infr;r~~I~aryllgobI-ar~icliiirl teeth hair-like; +Ch. 33: basioccipital fenestration present. C:raljtree (1989): female with ovipositor and eggs +Ch. 56: palatorostra1 liga~iientPI-cscnt. 'Ch. 28: supraoccipital crest bifid. +Ch. 59: endopte~ygoicltooth patch absent. 'Th. 36: postemporal anterior process wide. +Ch. 86: precaudal vertebrae over 50% of total vertebrae. '::Ch. 91: haemal arches with Ilaring expansions only. +Ch. 91: haemal-arch lirnncl. "Ch.102: body scalcs small. +C11.107: Aclenylate ltinase (Ak-A). +(;11.108: Aminopepticlilse (Ap-A). S1ii\/IO?'llJIJLYspecies group +Ch. 2: vo~nerinedorsomedial spine. OI)ON'I'IB'IZI133Y +(;11. 48: lateral fijld of labial ligament greatly I-educed. +Ch. 9: n~esethmoiclabsent. +Ch. (53: fotrr pores on horizontal shaft of preopercle. +Ch. 24: trigcminal fo~rtnenenclosed by prootic, sphenotic, +Ch. 96: scales of interorbital region ouly with reversed irn- and pleurospl~enoid. brication. +Ch. 41: maxilla~yvc~itral proccss about equal length as clop *Ch. 26: basisplienoid with anterior elongation to i~lterorhital s;1l process. carti1;rgc. +Ch. 53: mandib~~larbra~lcll of A1 ~nl~sclctc~ldou prcscut, *:Ch. 58: ectoptcrygoid absent. ;unterior to lacrimal branclr. tCh. 65: opercl' fei-nestrated internally. presence of fenestrated rncdial lamina between pos- +Ch. 66: APPENDIX 3 terior and ventral struts. +C11. 94: anterior hacmal spines short. Ambig-uous C/ZCLT(CC~~-~:A~nbig~~ity dc~cto altcr~lativccqtrally I>~I-- -Ch. 12: nasal sensory canal clearly clctachcd from nasal sirnonious rcconstrr~ctionsof a character's evolution may lead bone. to loss of diagnostic information if only the unique and unam- -(:11. 25: secoudary trigcrninofacial arc11 present. biguous traits are considered as infol-mative of relationships. -C:h. 38: prernaxillary ;~sccndingprocess with narrow base. Sorne ofthis phylogenetic inf'or~nationcall be retrieved, in the -(;I>.10 1 : scirlcs present bctween anterior rays of aual Gu. form of additional supporting characters, iP the type of arnbigu- ity affecting each character is identified. ~~l~YlI,l~~~1~rFlI7S Three classes of character ambiguities have bee11 identified +C:h. 3: vorner with two parasagittal dorsal laminae. that allow, given the asslrmptions, for the recovery ol'characters tCh. 5: vomer slightly concave ventrally. as evidence of monophyly: tCh. 7: vomer with single, median tooth patch. 1 ) character is arnbiguous because the relatively primitive state tCh. 13: lacrirnal coudyle of lateral etllrrloid docs not reach is present in a highly derived sister taxon to one of two ingroup the ventral edge of tllc lateral ridge. sister taxa (Fig. 46A). The auxiliary assurnptio~~is that the +Ch. 22: dermosphenotic 111edialIlangc absent. ingrolrp sistel--taxonwith the plesiomorphic co~~ditio~~is derived +Ch. 32: pterotic and epiotic wiugs contiguous. by some mode of paedomorphic development; tCh. 42: ventral proccss of milxilla with conclyle. 2) despite prcscncc ofthc derived character state in tlie ingronp, +Ch. 46: maxillo-ang~rloartic~~I:~rliga~nent absent. that character state is ambiguous because of other charactel-- +Ch. 48: labial lig-ament with Iatel-al fold reduceel. state polymorphism, absence of data, or both, in one or more +Ch. 51 : dentary coronoid flat at lcvcl with anguloarticular outgroups (Fig. 468). The auxilia~yassumption is that the coronoid. plesiomorphic condition is present at the outgroup nodes; and +Ch. 52: clentary gn;rthic spine ahsent. 3) a non-additive nlultistate character is arnbiguous because the +C:h. 54: palatine dorsal proccss tall, spoon shaped. character states present in each oP two ingroup sister taxa are +C:h. 55: eth~nomaxillaryliga~rle~it not attached to palatine. different and derived relative to the plesiomorphic state present +Ch. 62: preopercular pores 1 & 2 ~vithbony enclosure. in outgrolrps (Fig. 46C). The auxiliary assumption is that both +Ch. 64: opercular dorsal process 11ot blade-like. states arc derived i~ldepc~ldentlyfi-om the primitive conditio~l +C11. (58: opcrcular posterodorsal border concave. present in their cornmoll ancesto~: +Ch. 83: ;untrrior spine of first dorsal fin shorter than spine Different modes of optimization are c11ose11 for each indi- ofscco~~ddorsal fin. vidual character according to the type of ambiguity identified I.8: precauc1;rl vcrtcbrac over 57% of total vertebrae. after the phylogenetic analysis. Raiher than choosing a single +Ch. 95: first caucLrl vertebrae over posterior halfofa~~alGn. oprimizatio~~criterion for all characters prior to the analysis, an +Ch. 96: dorsal liead scalcs with reversed imbrication. a pos~wioriapproach is considered a more appropriate method +C:li. 97: four s~rborbitalscale rows. fhr additional-charac.trr support of monophyly. 111 the first and +Ch. 98: interopercr~larscalcs on posterior Ilalf only. second types ofamhiguity, characters are optirrlized to tlie ode -Ch. 25: secondary trigeminofacial arch absent. of greater geuerality with subseque~~treversals (ACCTKAN in -Ch. 37: presellce oL'an ~rppe~.,j;rwti-cnum. PAUP), whereas in tlle third type of ambiguity the derived char- -Ch. 40: tlorsal process of' maxilla shorr. acter states are optimized to nodes of lesser generality by as- -Ch. 17: prcmaxilla-maxilla ligament anteroventral region of suming the primitive state at the preceding node (DELTRAN ;rlvcolar arm. in PAUP). The number of ad hoc assumptio~lsdecreases when -Clr. 57: palatine spine not contacting quaclratc. suites of charactct-s of the same class ofambiguity are present at '"Ch. 8: single media11 vomcrine tooth patch. one node (Appendix 11: Atherinopsinae). Fig. 46. T111-cccl;rsscs ~Fclra~acte~statetlis(ril)r~lie)ns tlr:rt result in ;rlnbiguous optimizatiolls wllell lllappetl 011 a c.ladog~-;r~~~;A, Type I ;r~r~l)iji~rity,I~CC;IIISC tire rcl;rtiv~lyp~irr~itivc stirtc is prcsclri in a Irighly derived sister taxon to o1lr ol two ing~-o~~psistcr- taxa; 13, 'ljrpe 2 ;~~nbiguity,becairsc ore Ila~.ac.lc~-statc.poly~no~plrisrn, nbsence of data, or l)orll, ir~ollr or nlol-c otltgroups; C,'rypc 3 ;rmbiguit~l,l~ecause ilrc clr;u.actc~-st~rtcs p~-c-sc.nt in c;rch of two illgroup sistel- taxa arc, diSSc~-cnland dcl-ivcd ~.elaiiveio the plesiornorpllic state oSa lion- ;~tltliiivcnrultisi:rtc clra~acrc~;~)rrselrr ill outg~-o~~ps.111-I-ows indicate node of suggestetl o /,o.\tc,r.iol-i ol,tirni/-aiio~~.