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The Mugilidae of the world PDF

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THE MUGILIDAE OF THE WORLD THOMSON J.M. Thomson, J.M. 1997 10 21: The Mugilidae of the World. Memoirs ofthe Queensland Museum 41(3): 457-562. Brisbane. ISSN 0079-8835. Ofthe280specificnamesappliedtoMugilidae62areacceptedasvalid,with 18classifiedas species inquerenda. 14ofthe40definedgeneraarerecognised. Onlyonenewspecies,Liza mandapamensis is described. Mugilidae, review, taxonomy. J,M. Thomson, 15MarieviUeEsplanade,SandyBay, Tasmania 7005,Australia;received16 February 1996. omy. The main evolutionary diversification has 280 specific names have been bestowed on occurred in themouth and associated anatomyas mullets ofthe Family Mugilidae. Many ofthese demonstratedbySchultz(1946).Apartfromsuch havebeenrecognisedassynonymsinregionalre- commonlyused attributesasthecounts ofscales, views overthe past 120 years by Day (1876) and fin spines and fin rays and the measurement of Pillay (1962) for India, Jordan & Swain (1884) bodyproportions,thefeaturesofdiagnosticvalue for the Americas, Athanassopoulos (1919) and includethestructureofthescales,therelativepo- Popov (1930) for Europe, Weber & de Beaufort sition ofthe nostrils, the numberand form ofthe (1922) for Indonesia, Fowler (1928a, 1931, gill rakers,the form ofthepreorbital, the relative 1934) for Oceania, Borcea (1934) for the Black length of the paired fins and of their axillary Sea, Roxas (1934) for the Philippines, Smith scales and the position of origin of the various (1935)forSouth Africa, Fowler(1936), Cadenat fins, the presenceorabsence ofthe 'adiposeeye- (1954)and Poll(1959)forwestAfrica,&Thomson lid' andthedegreeofits intrusion overtheeye, as (1954) for Australia and Trewavas Ingham well as the numberofpyloric caeca and the rela- (1972) for the NE Atlantic and the Mediterra- tive length ofthe intestine. nean. Reviewsofthegeneraofmulletshavebeen offered by Jordan & Evermann (1917), Mohr Jaws. Jaw structure is basically percoid, distin- (1927), Popov (1931), Schultz (1946), Smith guished by the premaxillary having short pedi- (1948) and Thomson (1954). Howeverthere has cels and a shaft which, in the Agonostominae, is been no published review ofthe genera and spe- widest at its midlength and pointed at its distal cies ofthe Mugilidae as a whole since Giinther end, whereas in the Mugilinaethe shaft is broad- (1861b). Susan Ingham completed a review in estattheblade-likedistalend. WithintheMugili- 1952 which remains unpublished. I was unaware nae the edge ofthe premaxillae remains more or ofthis when I commenced this review in 1972, less parallel with the line ofthe mouth gape in ascnrdipatlttohomuyghatDtrenTtrieownaIvdaeslidbreerawteIlnyghdaimd'nsotmarnefue-r tshoemceogrenneerroaf,tbhuetminouotthh.ers it curves down behind tforoitmutnhtielstIahtausdomfaVdaelammyugoiwlnoujrudcgomncelnutssi.onAspaarrte The maxilla lies behind the premaxilla and at verysimilar, though often basedon differentcri- its upper end attaches to the ethmoid by a liga- twrweaoarkriseak,r.scaoIanmnhdpdalvaeIedtadeaceddkdonpbtotheweefldocerohdeuegnreItssyhtohsafatdtreamcIknoeogrmfshmcaoeomnun'ntsctheierdnelgvotiwghieeilwrsl pmmleaaedrxngiitecl.lelylaTsehg,eovvfidoeaerrgtntrheheeedeymboaayfxriptelrholletaordrcueyksgeiprdbreiotelociotetyfshsomeeofspbtrihwelehmimaitxcoyihuoltflahatlrhsiyeos arm, but otherwise features listed are those fuse inthemidline. When the mouth isclosedthe which I adopted from the literature or had eluci- premaxillary pedicels retreat under the nasal- dstartaetdimoynsealnfd. IsnovomlevegmeonvterinnmUneinvtersciotmymaidtmtieneis- bonAest.endonfromthesuperficialportionofthead- prevented my earlier completion ofthe review. ductor mandibulae is inserted on a flange on the anteriorfaceofthemaxillaryshaft.Thecurvature DIAGNOSTIC CHARACTERS ofthe maxillary is sufficientto allow the tendon to pass transversely upwards to the anterior face Mugilid species are remarkably uniform in ex- oftheshaftintheAgonostominaeandmostofthe ternal form and scarcely less so in internal anat- Mugilinae but in Chelon, Liza, Oedalechilusand MCMOIRSOI f l i ntmu9(iuiu-iL lH&mouthgqp$is shori andthe physial knob wi;h almost vertical .. Ml aiJIltfsl horizontal, the maxilla has devel- oinetiTncs divided into lateral halves In ,i dis- oped an outward curve over the loWei fifth Ol its tinct groove. ii to provide an unimpeded lineofaction tor .he tendon. In these genera the end ofthe pre- Skin coveringtheedges Of the jaws are de- maxilla also curves behind the mouth corner. scribed as lips. The whole of the upper lip of In Agonostominae theshaft of(tiemaxilla isal- Mugihdaeisdelimitedby agroove,buttheanalo- most straight, bending slightly, ratherthan curv- gous groove hehmd the bower lip IS incomplete ing, at the level ofthe flange In Mugilinae the and its relative length isuusefuldiagnostic char- lineofthemaxillary shall isalmoststraightwhen acter. The upperlipcovers theanteriorpartofthe viewed laterally and curve*; slightly and gradil- wemaxillae, In the majority ofmuiiei the upper allv in anterior view, h 'rJdkrhi- lip is bight i medially, narrowing gradual! iusmdCrenimugilihcmaxillaryshaftcurvesin2 the corners ofthe mouth. But in a few species, planes in its lowermost '.•<, hut outwards and CJ hfUp .sr.-.M and ( rrnimu^il ;iih1 backwards from the initial outward curve at the HilUS spp., lip height is very little differ* flange. Ma\ill:i- tdamugllsftd Myxua difftt cat, ifatall, laterally in w b&mmcnsi& the lip In* tromthesegeneraltypes. ThatofA/uw/.v is broad, ises markedly in height at the lateral ends. curving backwards bin scarcely outwards, li is Itie description 'thick' has ht*en used in iwo thus intermediate between Agonostominae difl'crcnt senses in the literature when describing the MugU group. Hov/evc hi prcmaxilla of thelip. Ithasbeen usedtodescribe either the dis- Myxus is typicalofAgonostominac 1'hemaxilla tance from front to rear or the vertical spread of of ValamugU indicates a transition between the the lip. 1 use the termto describe the relathe dis- Mftgtl and Ijzu groups of Mugilinae, with the tancethe uppej lip projects in frontofthe ww maxilla curving down behind the mouth in one edgeofthe head. Ihe vertical spread iscoifed lip planeonly, lathei than theS-shape typical ofiuu height In some species the mandibular angle and its relatives. Two conditions have been con i 'ii:;cN froni acule 10obtusewithage(Thomson, fused jii the literature undei the genera! descrip- 1*254) r\s the head broadens th<? shape ofthe up- tion o\' "maxilla visible below and behind the per tip maychange but this is marked onl> in Va- I'onicr Of the mouth when the hlOUtfl , hmugtls&helt and its close relatives What are visible are pads of tissue, ihe maxilla In t'ht-inn. Crerumugil and Qtidafechilta the beingsubmerged in thepadto varyingdepths. In lowet pail ofthe upper lip is raised into papillae most species to winch the description has been whose ultimate shape hs ofspecific significance, appliedthere are2 pads lying.closetogether- (he They are epidermal structures without any con- posterior, and usually larger pad, is that over the nection to the teeth. In young specimens ol most end ofthe maxilla, In front of it is anothet pad Speciesofihesegeneraand inadult(".helnnluhru- whuh overlies theendofthe tendon to the corner v//,v small stMitorm teeth are present as well a ofthe mouth. In VatatAUgfiW\\y the pad over the papillae.Thepapillaearenotevident inyoungfry tendon is visible. In some species, notably of V Until a length of about 60mm SI, and there Ap- w, this pad is often hidden by a cutaneous pearsto be considerable variation inthe lengtfl at flapgrowing from behind the base of the lip. whichthey become visible I lYewavasA Ingham, The lower jaw presents 2 features of generic ] lH2). The lower lips ofAgonostominac and of significance* In wore primitive genera the ( ( rettfmngiland O&daleohliua arc thick and deep. noid process ot the dentary bone it massive ;md The rest ofthe Mugilinaehavethin chisel-Sh mounts from thegeneral shaft ofthe dentary well lower lips vvhith project almost horizontally for- in front oi the mouth corner. In othergenera this ward. Litetheupperlips,thelowerlipsOl ( rcni- process is slenderand risesfrom the shaftbehind muy.iL C/it/on and Gedatechilits are ornamented ihe mouth cortiei In Cestracm the dentarj r with papillaeot ctemilations. The lower lip tended posteriorlyand ventrally to form a deiep]} is permanently folded down in Ceslraeus spp. undercut flange which is i rvered i a flesh} The IipofAiugtithuhiirnthasalsobeen described lobe. The other d feature of the lower as foldeddown, but ofthe 14 specimensavailable jaw isthe -i1 ' fdevelopmentofthesymphys- for study only one had the lower lip completely iai knob, In theAgonoMoji nai eiinei no folded: iii 4 the llpw moi curled down al .ill and central knob rising above the lower lip [Agonn* in the remainder sections ol varying length yfumm spp.) orthere is only a broad low IDOUfld, . oi both sides were turfed Occasional indi ag the Mugilinae j hattfintugilh&$ only a viduals ofotherspecies have been observed wiMth low mound, butindiJ ulhe: genera there is a high lips curled down, including Mugfl curcnuK Ml JOIDDAE OF THE WORLD 450 M ittrviikns and juvenile rcphnhr< Prohno1 nearer eachOtherthantheposteriorlytotheeyeor mullet with thin-edged tower lips may be ableto theanteriortothelip; inothersoneorboth rtiti cur) the lowei lip voluntarily dunru; life ^ikJ ap- closer 10 the appropriate feature than to the other pearancealterdeath orfixingdependsupon mus- nostril. The posteriornostril usually reaches iu<- culai eonttaction. Generally the lower lip is about the level ofthe uppei iim pi theeye. but o fl entire, i e, the anterioredgecontinues in the same few specie s r higher. There is one species. Rh:- plane across the symphysis! region. In several notftui innis (Swainson), in which the eflfiffl the external edge of the lip curves up in.'.h-TJur nostril is displaced to the level of the around an external groove under the symphysis. lowerhall ofthe eye. "Nucha lip is notentire' l he lower lipoftfzam\<l ofafewotherspecieshave a row orrows 1 papiI- '. Mullet teeth are small and usually de- toe near the hase ofthe tffx scribed as either eili;ife 01 setifoiiu. In AgOftO- stomraacteeth are sessile,ofnormal teleost type Preorbital. In the Agonostominae ihe preotbiia) though small., borne dueetly on the premavilla isalmost quadrangular; in Mugilinae roughlytri- and dentary in bands ofvarying width. In form angular. Ihe external surface of the preorbital is they ate ciihci incisoi-like or are wide-fronted, flat in SgtmQStomx&i Jotttrus, Cestraeus and flattened antero-postenorly. and are often multi- ( %teftomugil Inother geneia aridge*originating cuspid. In the Mugilinaethe t\tefh are minuteand at the base ofthe posteriormost serraofits lower labial, situated attheextreme edge ofthe lips, on edge runs roughly parallel with the front edge of thedistal ends of Hexihle strands in the lipt, the preorbital. forming a scoop-shaped trough which pm\itnallyjointothedentaryortothepre- alongthe ameiiot portion ofthe preorbital. In life maxilla. They vary from setiform or ciliifoi this trough is filled with eithci flesh 01 adipose flattened teeth, sometimes with multiple cusps li'SStfe, The topographical anterior edge of the Ciliiform teeth are differentiated from setiform preorbital isregarded asthe ventral edgeby those teeth by being [ nor and lacking the dark core concerned with the homologies ofbones and the typical ofsetiform teeth. topographical ventral edge is the posterior, lo avoidcontusion the term front edge' is used here Scales. Bothcycloidandctenoidscalesare ibund tortheedgenearesttheupperlipand loweredge' in Mugilidae Some variation in attribution oc- for thai extending between lip and eye. This tcr- curs hi the literature- Jacot (1920) , Kesttven ofJoitjWogUjSIwShneortea,ptbetncatuhesecaosfedoifffaegreendtisaplegcriomwetnhs (of19M4u2g)iaLnvdptTvhtoimnsoans(c1te9n5o4i)d.deSscchruilbtezd(t1h9e46sc)aalneds of the front edge the point that is normally the S|i tli( 1949)calledthemcycloid.Onmoststales Liemostonthe loweredgecomesto lie in frontof ofM- cephatffl and related species, a row ortwo the original anterior cornerand the normal lower of weak ctenii mark the posterior margin edge comes to lie over the hindmost part ofthe scale Batthectenii laiddown earlierdo not proi- frontedge.Theloweredgeofthepreorbitalisser- ect;theirformersitebeing indicatedbyapattern* rate in all but a few species. In a few the sena. ing ofthe surfaceoftheposteriorquadrantwhich have become sharp spines. In most species the has been interpreted either as the ctemi sub fsroomnet,edhigegeanlesoraiswsietrhramtea,xiblultaereithLaatindo mnootothheridn mafetTergetdheipnoithnetsdshsauveeosflotuhgehsecdaloerohravasetbheeeirnbworn down behind themouth corner, Ihe front edge of type ofscale is distinguished as pave- -Iinh.tfnMoosnehigteanlerias ewihtheerresttrhaeignh:ta\otrlTgaenbtelyndcsurdvoiwngn menItwoctetnyopieds'.ofscalehavebeendescribed as K ohrebhiitauldtishenomtocuhtedhtcoorancecrotmhmeoIdraont!eetdhgeettonfoutlhhecpioie-- posterior neudggielidasndOnisectoympmeohnalsya ffoirumndroounndtehde ner. In most species this notch persists, but in flanks ofcertain speciesol Lizaand Mugil I some it tends to fill with age. In more primitive liles ol VaJamugU and Crcmmugtl have similar species die preorbital is broad and fills the space •s but in latejuveniles and adults the %eales between the lip and eye; but in more advai develop ii Flexible membranous margin posteri- spetiteS (he [ncurbital is narrow; in some it fll orK which can be seen to be fimbriate in well- only part ofthe space between eye and lip. ptcserved scales. Nostrils. Tic nostrils lie m characteristic posi- A proportionofscalesarcprovidedwith pitsor tions "i each species. In some, the nostril grooves known as mucus t mils These first be- 160 Mi MOIKSOl THtQUl NSI.ANDMIiSUlM I come apparent as alniosi cJrculaj depressions in • aecu withinthe lew speciestheyexamined, fishabout40mm in length. This simplecondition but in my experience, including the &p< is retained in the scales ofadult Agonostomimic amMied bv those authors, the number has been 111 ttfyxm spp. and in lessspecialisedLiza, suchas fairly constant within a species. L, obUi ashort canal runs forward from the 01 nal pit. In other genera and in more specialised JUVENILE mullet. The characters tteed in Liza the canal may be much longer but does not and detailed descriptions can he used confi- penetrate the segment of the scales posterior 10 dentli y only with adult mullet and withjuveniles the nucleus tq an) extent except in ValomugU whose length is greaterthan oOmm SI fven ai where in .some species it reaches the pn?teri H this lengthsomecharacters, such as theextension Ofthemembranous margin. Insome species oftheadipose tissueovertheeye in somespecies a scale may have Y-shaped, T-shaped, double or has no! reached ihe adult slate. Al about 60mm even triple canals Tins lias been taken njnliet in SL a degree o( metamorphosis takes place in- Liza Sfflfetts and L. dumerili where the di volving not bnlj the transformation ofthe third scales on the head andon theback, anierioi (othe supportingelementoftheanal fintoaspineinthe first dorsal fm have 1 canals in cariyjuveniles, Mugilinae and a few Agonostominae. but also in hutbythetimethv fish is adult ih. ;l ft mul aconsiderable rearrangement ofthe mouth parts ticanaliculale, hiving between 7 and 10 canals. In the young (Qucrimana) stagesthe mouth e Immediately above the iiiseition ofthe pectoial reply inelmed but later becomes less sleep m somespecies haveanaxillaryprocess,usually and almost horizontal in some genera. The nos- r1eferred to as the axillary scale, although one trils are usually above the level ot the eye in -•cale may be involved in its structure. The axil- voung mullet but descend below the level ofthe Ian' scale is lacking m the Agonostominac, It is upperrim ofthe eye m most species atmetamor- strongly developed in MU&il t'atamugil and phosis The dorsal fins are more crowded in Crenimuglh some species ot' Liza have a moder- youngmullet,thetipoftherecumbent firstdorsal atepectoralaxillary scalebut inmostLizaspecies spine almost touching the origin of the second it is reduced to a misshapen rudiment or is com- dorsal fin and the tip ofthe second dorsal fin al- pletely lacking. most reaching the base ofthe caudal tin \-\ . pi lorspecies with auniqueattribute, such as an un- tIintveesltyinseh*ortI,n Algyionngusintoonmliyna3elothoepsmlinestihemabidonfmiila- cuuslutaltloydjijsrteiantgfuiins-hratyhecosupnetc,ieist aislesxmtarlelmemlulyledti.ffi- nal cavity and measurui '.0 times standard I METHODS 'length ofthe fish. In more primitive Mltgilinac, such as \fvxus\ the coiling has increased and (he relative length is Lv;> times standard length' In Most ofthe measurement: are standard in sys- V/ugilthe intestineiselaborately coiledandis3-5 tematic studies, but due topeculiarities ofmullet tliomnegesrsitnanodtahredr lmeunggithh.neThgeeneirntaesattitnaeiniisng< 6-7. tsroomdeucneodna-nsdtatnodat.r-dicimlietnaiseuprreesmeennttatihoanvesobmeeennoinn-- times the standard length in Chclutt tahrasu?, standard abbreviations have been used. Stomach The stomach is a Simple U shaped sac Scales The numberofscales in the lateral series with uniformly thin walls in the Agonostominae (LI)hasbeencounted from thescale immediate-!;. other than ti/jnrhftitj In typical Ylui'iliunc 'he ethe insertionofthepectoral fin(i.e.justbe- stomach is divisible into a thin-walled cardiac hind the head) to the caudal flexure fbe 5-S crop and a very thick-walled bieonical pyloric scales of intermediate size between the flexure gizzard. In this respect Myxits elongtihts, M and the very small scalescovering the caudal fm tardi and AldricheUa hrsieti arc intermediate. have not been counted. The transverse scale having weakly developed gizzards count (tr) isthenumber ofscales between the ori- ginofthe firstdorsal fin and theoriginofthe pel- rylorh-i-ihca, Theprimitivenumberof.' p) folic iu (Some authors have made their lateral caeca is found throughout the Agonostominae countfrom thebaseoftheseconddorsal fin tothe and in Mugil. Rhinom'Ugii, SiCamugilt <' fuicnn- base ofthe anal \ui ) The peduncle scale count mugHand Myjsus (except i'orMyxus capensis), In (pcd.) is the number ol scales between the mid- other genera the numhet ofcaeca \ u ieg i 2-22), doisal andmid-ventral linesdown onesideofthe though moreusually 5-9, Ddgelfi litis(I%3) re- caudal peduncle, bul omitting any median -ole* portedaquitewide variation in thenumberofpy- ifpresent. MUGILIDAE OF THE WORLD 46 Bodyproportions.All lengthmeasurementswere the length ofthefilamentsandc.1/3 length ofthe taken between parallels exceptforthe estimation lowerarm ofthe gill arch; Very long: the longest ofthe position ofthe origin ofthe first dorsal fin rakersareat leasttwiceas longasthe longestgill with respecttothetipofthe snoutandthebaseof filaments andc.1/2 lengthofthe lowerarm ofthe the caudal fin for which dividers were used to gill arch. hasten the procedure. Dividers were also used to measure the interorbital, width ofthe head, body Type 1: fringes short, coarse and widely spaced;rakersshortandcoarse.Thistypeofraker depth, depth ofthecaudal peduncle, height ofthe upper lip, length ofthe lowerlipgrooveandposi- resembles the condition in Atherinidae and gen- eralised percoids. tion ofthe nostrils as well as lengths ofthe axil- lary scales of the pectoral and pelvic fins. The Type2: fringes long andclose-set; rakersshort standard length (SL) was measured from the tip and close-set; ofthe snout (which was taken to be the anterior- Type 3: fringes long, coarse and wide-set; mostpointontheupper lipinthosespecieswhere rakers short or long; the lip is terminal) to the caudal flexure which is Type 4: fringes long, coarse, close-set; rakers distinguishable by flexing the tail, even in those short or long; specieswhere it is notreadilyapparentby inspec- Type 5: fringes long, fine and close-set, of a tion. Caudal flexure was alsotaken as the caudal feather}' appearance; rakers long; base for determining relative position ofthe ori- Type 6: fringes very long and close-set; rakers gin ofthe first dorsal fin. very long. Monthgape. Themouthgapewasrecordedasthe ABBREVIATIONS. For convenience and brev- ratio ofmouth width tomouth length (MVV/ML). itycertainabbreviationshavebeenusedinthede- Distances were measured between parallels, scriptionsofgeneraandparticularlyofspecies. A mouth width from mouthcornertomouth corner, - anal fin; the number ofspines are indicated by length from theanteriortipofthe liptotheposte- Roman numerals and the number of branched rior corner ofthe mouth opening. A number of rays by Arabic. Di = first dorsal fin: Roman nu- other authors, notably Trewavas & Ingham merals indicate the number ofspines. D2 = sec- (1972) have measured mouth lengthtotheend of ond dorsal fin; the small Roman numerals themaxilla. Theactualgape ispreferablebecause indicatethenumberofunbranched rays,the Ara- the extension ofthejaw behind the mouth corner bicnumerals,thenumberofbranchedrays. Di Sc isveryvariable, being veryshort in some species = the number of scales between the operculum and up to 1/3jaw length in others. and the vertical from the origin ofthe first dorsal fin. D? Sc - the number of scales between the Cillrakers. The systemofcategorisation devised operculum and the vertical from the origin ofthe by ingham wasbased on the numberofrakers on second dorsal fin. the lowerarmofthegillarch,coupledwithcalcu- LI =thenumberofscales inthe longitudinalse- lations ofthe length ofthe raker at or nearest to ries, counted from the scale immediately above the ceratobrachialjoint relative to both the long- the insertion ofthepectoral fin (this is usuallythe estofthegill filamentsandtothelengthofthegill scale immediately above the edge ofthe opercu- arch. have added the count ofthe rakers on the lum) to the caudal flexure. ML = length of the I upper gill arch and in some instances extended mouth gape measured from the snout tip to the the range ofcounts on the lower arch. line joining the two corners ofthe mouth gape. MW The gillrakers are counted on the first gill arch = width ofthe mouth measured in a direct becausetherakersare longerandhencemoredis- line from mouth corner to mouth corner. cerniblethanontheotherarchesandthey lackthe P = Pectoral fin; the accompanyingnumberin- spicules which serve to interlock the rakers with dicates the number ofrays in the fin. pect. sc. = those oftheotherarches. Six categoriesofrakers the scale in the longitudinal series reached bythe can be distinguished among Mugilidae. In defin- tip ofthe pectoral fin when laid back. ped. = the ingthetypes the following terminology has been numberofvertical rows ofscales down one side adoptedfrom Ingham'sunpublishedmanuscript. of the caudal peduncle, excluding any median Short: the longestrakersareonly 1/2 the length scales. ofthe longest gill filaments and c.1/4 the length SL = standard length, measured between verti- ofthe lower arm ofthe gill arch. cals from the snout tip and the caudal flexure. Long: the longest rakers are distinctly >l/2 sp. sp.2. sp.3, sp.4 =* the first to fourth spines 1, length ofthe gill filaments, but markedly < twice in the first dorsal fin. 462 MBMOIRSOFTHEQUEENSLANDMUSEUM tr= the iiumber ufscales in atransverse series REMARKS. Jordan & Evermann < 1896) dfeltn counted from the origin ofthe first dorsal fin to guished the Mugilinae and Agonostominac. the insertion ofthe pelvic fin. Their criteria, based on the American species, LOCATION OF SPECIMEMS. The museum were the presence orabsence ofsessile teeth, ihr cionldlieccattieodnbiynwthheicfholelaocwhinsgpeacbibmreevniaistiporness:erAveMd is dsehgarpeeeofofthceomlpolweexritjayw.ofOntlhey sttheomfaicrsht oafndthetshee Australian Museum, Sydney; BMNH = British holds on a world wide basis, but the subfamily Museum (Natural History), London; BPBM distinction is realistic as a more detailed exami- BerniceP. Bishop Museum, Hawaii; 1M = Indian nationrevealsanumberofconsistentdifferences. Museum, Calcutta: LA - Laboratoire Arago. Bunyuls-sur-mer: MNH1M = Museum National Subtamily AGONOSTOMINAH d'HistoircNatuielle, Paris; NM=Natal Museum, eDnunrab;anQ;MNH-MQ=uNeaetnurshliasntdoriMsucsheeusmM,usBeruimsb,anVei;- ryn1g.eTaelesthcosveessrielde,wiintshidsemtahlelltiep:et2h.;s3u.pegirlilorrapkhear-s KMNH - Rijksmuseum van naturlijke Histone. short; 4. preorbitai Hat, without a ridge; 5, some Leyden; SAM - South African Museum, Cape genera (AgonostontMS and Joiurus) with only 2 Town;USNM «UnitedStates National Museum, anal spines, followedbyanunbranchedray inthe WAM Washington; = Western Australian Mu- adults; the unbranched ray becoming a spine in seum, Perth; ZIZM = Zoologisehes Institut und the adults of other genera; 6. interorbital con- Zoologisehes Museum, Hamburg. vexI\ arched, headnarrow; 7. stomach U-shapcd with uniformh thin walls; 8. intestine relatively DATA RECORDED. Synonymies include only short. <2.6 times SL; 9. symphysis] knob either thosereferencesin whichsufficientdescriptionis absent ora broad low mound. included to be reasonably confident that the GENERA. Thereare 32nominal generaofMugi- esnpceecsimaernesiwnecrleudceodrreicntltyheattsryinboutneydm.iAesfebwecreafuesre- gliednaeer(aeaxrceluhdeirnegrfeocsosiglnifsoerdmsa)s(vTaalbidl.e 1 ). Only 14 they provide novel combinations ofgeneric and trivial names. Counts ofthe caudal rays are not KEY TO THE GENERA OF given because examination ofyoung fishes con- AGONOSTOMINAE sistently gave highercounts than are recorded in the literature. In post-juvenile mullet the small i v ,, Kp . i.- - outercaudal rays are almost impossible- todetect UpperlipAotaverfiungbysnoui ........ without tedious dissection. The biometrical data 2(\). 1 jnpl spinestnadults;interortit.ilhigh,convex forthespecies are presented in tables as appendi- . . . . - . . Agutstnnio ces. I find it a farmore useful tool forcomparing J SpitteS tn.a.ck.fth.j.i ......!"v.\;.... characteristics than embedding them in (he de- 'i: og| rlat. . _ i scriptive text. The lengths recorded for museum rulingfi 'ivlobesbetweentoweru-as mm specimens arc SL. , d I !" ... AbfaihiLt SYSTEMATICS Agonostomus Bennett, 1831 TwoeditionsofValenciennes. 836 werepub- I AgoTjostvTTiusBennett, 1331: 16b lished with differing pagination. Pagination of n 'he 2nd edition is shown throughout in brackets. P5 SPECIES.Avunustomu-ridf.uni Bennett, 1831. Dajan* Valenciennes* 1836: 16+(llb). pi. 316. Tvpe species: MUGILIFORMLS Miii'ilmoiftifoi* Bancroft* 1836, Family MUGILIDAE NatitValenciennes, 183') 167(11 i il ; ' typespacks: Nestis typrotoides Valenciennes, 183b by subsequent des- DIAGNOSIS. Pelvic bones connected to the 7i2g;nattyipoensopieciJeosr:dNaneo1m9u19g;ddiguttiYuill.int.I 18H9a4n.t, JS94: postcleithra by a ligament; vertebrae 24 (rare vwairtihan4tssptione2s6,);th2eseparate dworistahl fuinnsb,rtahnecahnetderriaoyr nisTRimPTfON W Indian Ocean. Cfentwl Americi, Garib (often called a |spine) and 6-I10, usually 8, bean. branched rays; anal tins with 2 or 3 spinesand 8- DIAGNOSIS. Interorbital high, convex, less so 12 rays; pelvic fin with a spine and 5 rays; tcdh inearlyjuveniles, Mouth gapeoblique. less so in either sessile or labial. large fish; mid-gape at mid-eye level in smull MUGILIDAE OF TlIE WORLD 463 TABLE 1. Thenominal generaofMugilidaeandtheirgenericidentity. Nominalgenus AuthorDare TypeSpecies AssignedGenus Mugil Linnaeus 1758 M. cephalusLinnaeus Mugil 1. 2.Chelan Artedi 1793 M. cheloCuvier Chelan 3. Cephalus Lacepede 1800 M. cephalusLinnaeus Mugil 4.Aganostomus Bennett 1831 A.telfairiiBennett Agonostmus 5. Cestraeus Valenciennes 1836 C.plicatilisValenciennes Cestraeus 6.Dajaus Valenciennes 1836 M. tnonticolaBancroft Aganostomus 1.Nestis Valenciennes 1836 N.cyprinoidesValenciennes Aganostomus 8.Arnion Gistel 1848 M.cepfialusLinnaeus Mugil 9.Ella Gistel 1848 M. cephalusLinnaeus Mugil 10.Joturus Poey 1860 J.pichardiPoey Joturus 11.Myxus Giinther1861b MyxuselongatusGiinther Myxus 12. Chaenomugil Gill 1863 MugilproboscideusGiinther Chaenomugil \3 Rhinamugil Gill 1863 M. corsula HamiltonBuchanan Rhinamugil . 14. Gonostmyxus MacDonaldl869 G. loa-loaMacDonald Cestraeus 15.Neomyxus Steindachner1878 MyxussclateriMacDonald Chaenomugil 16. Querimana Jordan &Gilbert 1883b MyxusharengusGiinther Mugil 17 Aeschrichthys Macleay 1884a A.goldieiMacleay Cestraeus '. IS.Liza Jordan&Swain 1884 MugilcapitaCuvier Liza 19.Trachystoma Ogilby 1888 T. multidensOgilby Myxus 20.Neomugil Vaillant 1894 N. diguetiVaillant Aganostomus 21.Oedalechilus Fowler 1903 MugillabeaCuvier Oedalechilus 22.Squalomugil Ogilby 1908 M.nasutusDeVis Rhinamugil 23. Xenorhynchichthys Regan 1908 JoturusstipesJordan&Gilbert Joturus 24.Ellochelon Whitley 1930 MugilvaigiensisQ&G Liza 25.Protomugil Popov 1930 MugilsaliensRisso Liza 26.Sicamugil Fowler1939b MugilhamiltoniDay Sicamugil 27.Gracilimugil Whitley 1941 MugilramsayiMacleay Liza IS.Moolgarda Whitley 1945 MoolgardapuraWhitley Liza 29.Planiliza Whitley 1945 MaolgardaordensisWhitley Liza 30.Aldrichetta Whitley 1945 AganostomusforsteriVal. Aldrichetta 31.Xenomugil Schultz 1946 MugilthoburniJordan&Starks Mugil 32.Crenimugil Schultzl946 MugilcrenilabisForsskal Crenimugil 33.Heteramugil Schultz 1946 MugiltricuspidensSmith Liza 34.Oxymugil Whitley 1948 MugilacutusValenciennes Myxus 35.Pteromugil Smith 1948 MugildiademaGilchrist&Thompson Liza 36.Strializa Smith 1948 MugilcanaliculatusSmith Liza 37. Valamugil Smith 1948 MugilsehiliSmith Valamugil 38.Plicomugil Schultz 1953 MugillabrasusVal. Oedalechilus 39. Osteomugil Luther 1977 MugilcunnesiusVal. Valamugil fish, dropping to lower iris in large fish; mouth gape, lyingon vertical from anteriorrim ofeye in corner at level oflower rim ofeye in small fish, small fish, underanteriorhalfofeye in largefish; droppingto well below eye level in large; mouth upperlipthick,terminal heightatmid-gapemore corner on vertical between posterior nostril and thanhalfeyediameter; lowerlipthick, notturned eye in small fish, moving to or behind anterior down, entire, recessed slightly behind upper lip; rim ofeye in large fish; upperjaw end on line of lips withoutexternal papillae orcrenulations; lip . 464 MEMOIRS OFTHE QUEENSLANDMUSEUM groove >2/3 lip length; no symphysial knob; no KEY TO THE SPECIES OF AGONOSTOMUS fleshy lobes externally between lowerjaws. 1. Mid-heightofupperlip 20-22% HL; uWpperjaw endat Maxilla straight, not curving down behind IvenrdtiiacnalOfcreoamn)anterior run ofeye (S catalaiPellegrin mmoourtehfcroormneur,ppietrs teenndd;onmafxlialnlgaep1a/d3 soromseltiighmtelsy Mviedr-tihceailgfhtroomfaunptpeerriolripri<mo1l6e%yeHL;upperjawendbehin2d visible as oval patch above premaxillae, but not 2(1). Eleven scale rowsdownsideofcaudal peduncle;sec- visiblebelowandbehindmouthcorner;coronoid onddorsal fin origin atscale23 (SW Indian Ocean) processofdentary massive, risingwell in frontof telfairiiBennett mouth corner; mandibular angle acute. Ninescale rowsdownside ofpeduncle; second dorsal Without labial teeth; sessile teeth in several fMienxiorciog,insoatutshcealren2U7S(AP,aciCfoilcuamnbdiaA,tlVaennteizcuceolaas,tsCaorfib- rows, inner rows multicuspid in some species; bean) monticola Bancroft wide gape between teeth at symphysis; teeth on vomer, pterygoids, palatines and tongue; tongue Agonostomus catalai Pellegrin, 1932 flat, with short free anteriortip; noadiposetissue on face; preorbital massive, flat, without folds or Agonostomus telfairii var, catalai Pellegrin, 1932: 424, ridges, not notched; nostrils nearer each other Mananano, Madagascar; 1933: 84, fig. 99, Mananano, Madagascar. thanto liporeye;posteriornostril nearereyethan Agonostomus catalai Pellegrin, 1935: 72, rivers of Madagas- anterior to lip; in adult eye halfits own diameter car. rbeealcohwindgorbseallowheeayde;cgoinlltoruark;ersopteyrpceul1ar opening tHuOdeL,OcoTllY.PRE..CaMtaalnaa,nManNo,HMNada32g-a1s6c2a,r,1f3r0emshmwaStLe,reatxa1m0i0nmeda.lti- Upperinsertion ofpectoral fin at levelofupper rim ofpupil,withoutelongate axillaryscale; first M1A98TmEmRSILA,LfroEmXMAaMdIagNaEscDa.raHnodloAntjyupaen.anBdM3NsHp:eci18m6e5n.s9:.211.304-- dorsal fin origin nearer snout tip than to caudal 5, 168& 198mm,Anjuan. base; seconddorsal fin origin variously fromjust behindvertical fromoriginofanal finto 1/3along DESCRIPTION. Di IV, D2 i 8, A II 10, P 17, LI anal fin base; 2 anal spines in adults and young; 42-44,tr. 12-I3,ped.ll,pect.se.9-10; Disc. 10, scales ctenoid; no opercular spine; caudal fin D2 sc. 23. Scales with short mucus canals. Body forked. Stomach without a gizzard; 2 pyloric robust, head bluntly rounded, scale-free to be- caeca; intestine with only 3 loops, its length not tween posterior nostril and anterior rim of eye. more than twice SL. Upper lip height almost equal to eye diameter; median upper edge of lip expanded, filling DISCUSSION. Valenciennes (1836) in igno- marked concavity in snout. Anterior mandibular rance ofBennett's (1831) description ofAgono- pores large,secondpairborderinganteriorendof stomustelfairiidescribed2speciesofNestisfrom lipgroove. Teeth unicuspid, incisor-like, slightly Mauritius and a species ofDajaus from the West curved, pointed, scarcely emergent in lowerjaw. Indies. Gtinther (1861b) revived Agonostomus, Tongue without median ridge. Mouth corner on including therein Dajaus. Cestraeus and Agono- vertical between posterior nostril and eye; tip of stomus as well as Mugilforsteri Valenciennes. upperjaw slightly below level ofmouth corner Cestraeus and M. forsteri have not been consid- and on vertical from anteriorrim ofeye. Preorbi- ered congeneric with Agonostomus by more re- tal filling space between lip and eye, reaching cent workers. Criteria used by Valenciennes to mid-eye level, well below linejoining midpoints distinguish his 2 genera are features which differ of posterior and anterior nostrils; lower edge withageorsize. Thegeographicseparationofthe reaching below lower rim ofeye. Posterior nos- SW Indian Ocean and Caribbean species must trils reachingabove level ofupperrim ofeye; an- raise doubts about their congeneric status. But terior nostrils wholly within vertical span of close examination and comparison ofspecimens posteriornostrils;widecutaneous rim around an- confirms the similarity oftheirfeatures. This isa terior nostrils. primitive genus; extant species are probably Pectoral fin reaching anterior 1/2 ofeye when widelyseparatedsurvivorsofaoncemorewidely laid forward, notquitetoverticalfromfirstdorsal distributed genus. The absence of a snout over- finorigin and 1/3-1/2 alongpelvic fin (notreach- hanging the mouth distinguishes Agonostomus ing the end ofthe pelvic spine) when laid back. from Joturus. Cestraeus possesses fleshy lobes Pelvic fin origin nearer vertical from first dorsal externallybetweentheramiofthe lowerjaw.The finoriginthantoverticalfrom pectoral finorigin; 3 analspines inall buttheyoungquerimanastage its tipjust reaching behind vertical from sp. 4 of marks offAldrichettaforsteri and the Mugilinae first dorsal fin. Pelvic axillary scale reaching 1/2 from Agonostomus. along pelvic spine. Sp. 1 offirstdorsal fin longer MUGILIDAE OF THE WORLD 465 TABLE 2. BiometricsofAgonostomusspp. D=body origin, the head is relatively shortand thegill ra- depth;ED=Eyediameter;FES=serraeonfrontedge ker counts differ. Pellegrin (1933) described A. ofpreorbital; GR=numberofgill rakers; HL=head telfairii var. catalai as having A III 8, evidently length; HW = head width; 10 = Interrorbital; LES = countingthe first unbranched anal ray asa spine. sMeLrra=e omnoultohwegrapeedgleenogfthp;reMorWbit=alm;oLuLth=gLapoewewridltihp;; TahfeortmyupleasopfecAimIIen10h.as 9 branchingrays, not 8 or PB = pectoral fin base; Ped = peduncle depth; PL= lpeencgttohr;alTfRinl=enTgotoh;thSLro=wsst;anUdLard=luenpgptehr;Slinp;LU=LsnHou=t Agonostomus monticola (Bancroft, 1836) Upper lip height; VL = pelvic fin length; VAx = length ofpelvic finaxillary scale. DMaujgaiu!smomnotnitciocloalaBaVnaclreonfcti,en3n8e3s6,:316873,6:pi1.6346(,11J6a)m,aipic.a.316, An- tilles,SanDomingo,PuertoRico);Kner&Steindachner, Species A.catalai .4.telfairii A.monticola 1865: 8,NewGranada. Agonostoyna monticola Giinther, 1861b: 464, Barbados, Scaleradii 9-10 8-12 9-11 Mexico,Jamaica; 1869:444,Mexico. & Depth(%SL) 23.7-25.7 25.2-26.0 22.6-26.3 Agonostomus monticola Jordan Evermann, 1896: 819, HL(%SL) 24.0-27.3 22.4-25.0 24.7-27.0 West Indies, central&Mexico, Vera Cruz; 1902: 254, Puerto Rico; Jordan Rutter 1898: 98, Jamaica; Ever- HW(%HL) 64.0-67.0 64.0-66.0 67.0-70.0 ni.inn & Marsh, 1902: 114, fig, Puerto Rico; Fowler, IO(%HL) 45.9-49.3 45.0-48.8 31.5-38.5 1903: 748, San Domingo; Regan, 1907a: 66, central America; Schultz, 1949: 110, Venezuela; Suttkus 1956: ED(%HL) 20.8-25.8 26.0-28.3 22.4-31.5 43,Louisiana. SnL(%HL) 28.2-30.2 24.0-25.0 23.5-25.5 MugilirretitusGosse, 1851: 84,Jamaica. Agonostoma mwrops Giinther, 1861b: 462, West Indies; ULH(%HL) 20.0-22.0 10.0-15.0 9.5-13.9 1869: 444, pi 60,fig.l,West Indies. & MW/ML 1.45-1.75 1.4-1.7 1.1 - 1.8 AgonostomusmicropsJordan Evermann, 1896: 820, West Indies, central America; Regan, 1907a: 69, central Amer- PL(%HL) 80.0 81.8-89.0 66.0-75.3 ica. PB (%PL) 28.3-31.2 30.2-32.0 31.5-36.0 Agonostoma nasutum Giinther, 1861b: 463, River at San VL(%PL) 88.0-90.0 89.0-98.0 79.0-96.0 Geronimo, Guatemala; 1869: 444, pi. 70, fig. 2, central America. VAx(%VL) 32.0-33.0 33.0 30.0-35.0 Agonostomus nasutus Jordan & Culver, 1895: 424, Sinaloa; Ped(%D) 46.0-47.5 48.5-55.0 44.0-46.5 Rutter, 1896: 263, Cape San Lucas, Mexico; Jordan & Evermann, 1896: 819, Pacific andAtlantic coasts ofcen- TR(UL) 7-8 3-8 6-10 tral America;Regan, 1907a: 68, pi. 10, fig. 4,Guatemala. TR(LL) 1 3-5 5-12 Dajausnasutus Kner & Steindachner, 1865: 7 Panama, west coast. LES 9-11 10-16 5-15 Agonostoma pcrcoidcs Giinther, 1861b: 464, San Domingo, FES 4-8 4-10 5-6 Jamaica, West Indies. GR 27-32/34-42 20-26/26-42 8-12/16-23 AgoDnoomstionmguos;pReercgoaind,es1J90o7rad:an69,&SaEnveDrommainnng,o.1896: 819, San & Dajaus elongatus Kner Steindachner, 1865: 6, pi.1, fig-2, NewGranada. than sp. 2; sp. 4 short, not reaching behind verti- Neomugil digueti Vaillant, 1894: 72, Baja California, fresh- cal from tipofsp. 3when finraised;axillaryscale waternearLaPaz. reaching 1/2 along membrane behind sp. 4. Sec- AgonostomusdiguetiSchultz, 1946:390, BajaCalifornia. onddorsal fin originatverticalquarteralonganal Agonostomussatvinu Regan, 1907a: 68, pi. 11, fig. 2, Guate- mala; 1907b:65, RioNacasil,Guatemala. fin base;tipsofanteriorraysreachingbehindtips Agonostomus macracanthus Regan, 1907a: 69, pi. 11, fig.l, ofposteriorrays; anal fin higherthanseconddor- Guatemala; 1907b: 65,Rio Guacalate, Guatemala. sal fin, both higher than first dorsal fin; second Joturus daguac Eigenmann, 1917: 681, Rio Dagua, Colum- dorsal and anal fins lightly scaled anteriorly. bia. Agonostoma squamipinne Mohr, 1927: 178, fig. 1, Puerto Rico. DISTRIBUTION.MadagascarandComoroIslands. Agonostomus bancocki Seale, 1932: 467, Galapagos Islands, freshwater. REMARKS.A. catalaiisverysimilartotheother TYPE. None.Typelocality,Jamaica. Indian Ocean species A. telfairii, and may be an extreme variantofthatspecies. Howeverthe few MATERIAL EXAMINED. 123 specimens, 77-198mm SL (in- cludingthetypesofA.percoides,A.microps,A.macracanthus,A. museum specimens attributed to A. catalai ex- sahmiL, A. squamipwne and N. digueti) BMNH: 1847.27.59, hibit constant differences from A. telfairii ofthe 58mm,holotypeoiM. irretitus,Jamaica,Gosse; 1848.1.12.1083, same length. TheupperlipofA. catalaiishigher, 130mm, syntype ofA.percoides,Jamaica, Parnell; 1850.6.7.16, there isonlyonerow ofteeth inthe lowerjaw,the 1181651.m1m1,.7.2s,ynstyynpteypoefofA.A.ppeerrccooiiddeess,, WSaenstDIonmdiiensgMoe,dicCaulmiOnffgi;- upper end ofthe preorbital is lower on the face, cers,FortPitt; 1855.12.26.63940,133&260mm,syntypespiA. thepelvic fin differs in the relativeposition ofits microps, locality unknown, pres. Zoological Society; 1 s MBMOIRS0¥ UH-:QUEENSLANDMUSI OI8w6eln.y, 1964 I !6.O3imiI ..' 11i3q]&.tylp7e2mofmsAvmv.|vHitaij I< -n.irmul.i, pmogstaeriloirnenjosotirniilns:g ptohsetemriidoprolnfoisltri&lfnaontterrieoarchand Q\:ty(pe.su;u;"»f!.u,1c.,CaiilMvimmnnt,l.RiiSod.viNna;ciIs8i7l5,.6G.u9aJt-4e.ma1l2a7,jlSolvio; aboveuppet titti ot eve anleriornostrilextending IB4H.1.12.1084-5, 'is 6c 127mm, lamafcj; 1948.1.12.1090 slightly below vertical span ol posifrtor nostril wlIa-^rsOod'm.ImJs,la'B.ndaIsrd;bai1id8o6c24s2.L.187.482SB6S.S5I2O.3.I90.,1I:9J7B3MQCm,m,i IMRiiuon,,j.lMaXomn}a,timecmiza,e-;..,n.-,DClu£orBonmaiikxpnn.iGoLcjweJaint; antPeercitiooprroaslntoefsrtiirwoirllhya;waiothhinemaproskteerdioicujrraftmswhen laid Souther'n1M-3e,xi1c3o7;&1890WJRf1f.i1m!C2S,Bu1rl12umimo,RIi8fclhm' o"'ndll-RZ.,. 07mm dFcowrswualrd1,inrwehavetinitllagidvbwatcikcwin Ismraolmlofriishg,intaoitl*infgirus>t , i pec. 87-140mm, Rtthugh R., Si Viti- attain this by increasing margins as growth pro- 9.M;'1me*s.'iiipx9.H-ie;.i1tct3c.MlU4oci9i6h.x;5'.i-<l11\.oI2:1,1,i*S,7119'w4m72.rm.37-s3...u3'":s7niS.3,.tp.p.riKV1uI.m.1:C7cWWi,r-1m3n.8S.tr79.o;5n,uL81'Cn829ls7,2pi1.6eI^10ocMa9.Rmmui.i(mW!7n,i£HekrilP&ali1u57P;w4aM.tRr.lt.in„dt8rUe,W.aBlSUR,i.Raij5Til.ono1V.iA2iin.Icaj4i3UMu-aai5nbsnIJj-,, rttofiirehpniaa:gnocihfantipxonienglletlvhavaairericeryrtiFsilrpscvnoicagenalmrtleaihehociooranirirlghnseiofanbcr1ma/hooes2imfensafgopolirlero'iscnstgipgi1m.dn/oep24rneosslofaa)vfllp.iofeciincPrtmgesfo.tilnrvpdaiieot(lcslrpsvafftaisiiilctnnp : Mat«iorong:v,., Mexico.;..v> i ., | | 05i,|1e2^.61705V- s4pilnoen.g,Spr.eac2loiiHnler.shtrdhoirnsdalvefirntilcoalngFerrotmhatnipsp ISp 3 128mm. Tnnjdict L907M.LM1BJ ,,. . . when fin raised; axillary scale short, reaching (imiinc8z, CIo,lu. ,m.bv[iiwa-i.inr:V,19e0n9a.3.13I.N7)6,l-7Ra,ajRaLkl<iv,W&Nil'!mm.i Rio trIo origIiniivalbveehritnicdalba1s/e2 oaflosnpg, a'nasletcionnbdasdeo,rstailpsfoitn o i !•; i " 'I.U4-1S3, IQ sp &7 I »5n«n, Be anteriorraysnotreachingbehindtipsofposterior MKRMm:ai>'l/.o:.e.;Gnllea.iia.tanCv.to»uWdn1,<,aIH.RIot,ijn«.id.Lui0,r1Ra;,is,c;a;";'."L.1991133I.6.2RLLi1I5Si4-6,6-0IimdiIirpnetpc,oaRuIi«'ao!ln|uJ1m8|bmii!a; hfiingshmerodtehranatfeirIs\tsdeotaresilae(ldIt.afiinnnt:esrseiecocoronlnyddaddnoodrrssaaalllonfaignn.dhab-a-on„t'ahl ;:i, Mi itrvine, f,.,obigo.|iMlaM INro'b"a M '! i i2 19— cDImSrTivReIrsBtUrTaImOFNl.anFdrietoAVwewneztiL i tfldiei m I Vmert 5187,2n5arsapec, 80-SSmm, jyntypes ol California. )M . posskbl) spiW'ni^g ja tej (Andcc^Cin, '.''l|7i. Spec ,-;,'"M•''J1I43^.in2nin,p*M.i.u. i"|tHi 'nlw/M- I Iaide''lo,ii-p"-e',1 A.I4317, 2 REMARKS. Beinggeographic all\ isolaitd Ihi tpIyanp|-Aeedrtiioi!i i,qu,a1<mi.p.ir.ma Piu1 eirtoRIio i!: : : .]i|m i>n'J>ecbtmom- nus. r{i.skmootftcioincfoufasiin::gv.e!rywovva/rzia/brl^e/,wsietvheirnatlhcehager-- acters varying assize increases, resulting in the 41-47C,RtIr.PT12I-O1N3,.pDed.I9V.,pDe;ais8e. AItIt.D1i0,sP JoJ ! large i mi ti qmyaa.xiWlitathmeawgeastehse alinpds tthheicpkoesnt,ertoh-e D;s«. ?7 Scaleswith<M1 primary-radii and2or :.d comer ofthe preorbital extends posteri- 3 secondary radii; mucus canals shorty wild 1 orly. Thedescription and&figure ofDajcw -.jA>/t- Small led depression ar base; bodv robust, gptnsprovided by Knei Steindachne, head blIuntly rounded scale tree to level ofposte- identify thier species as A. monUcolu. A & nor nostrils, intcmt'bital ineieivsing with age ill was described as having a shorter pectoral fin relation to head length; eye diameter decreasing than A, nasitfvs bul the collection examine' relatively to ML with age; eye diameter greater dudes intergiadcs. .1. squumipmm wan de- than snoul length in small fisfies less in large. scribed bj Mohr (1927) as having a peed Median height of upper lip 1/2 eye diameter or axillary scale, but the type specimen lacks this more. Anterioi mandibular pores followed by 2 featUN mtl - a typical i monticofatJoiurusda* ptahiersrpeafrp-imtsosotpeunnidnegrtomeixdt-epruipoirlb.yOmuutlteirplreOjW)iSreDl lumbi.i. bi y Elg bed frIom( 1t9h1e7)Pacsivfhiocleoeapsoineofd(thoe teeth usually liltfcjispid, inner rows with between upper lip to be terminal, indicating A. moni 2 and 5 cusps, though BOtttc fish Iwive unlcuspld ratherthanJotunts Seale1 1>21)claimed thai Ins teethonly Mouthcornoratverticalbet1 l lumocki differed from A nasutus(r I • tenor nostril and e_\e_ well betuv. lower nrn of Cola) in havingthepelvic fins as long as thepec- Cye, l'u:or(»iral filling spai. c ln-tv ( E 1 tip .HI;! torals. Pelvic Iins m the type of 1 nasuivs lire scrrae obsolescent but countable: upper I only slightly shonei than the pectoral finsandno preorbital curvingback from STtOul tip. m»t n.:;ufi oilun .significantdifferences were listedbyScale.

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