PHYLOGENETIC REVISION OF ATHERINOPSINAE (TELEOSTEI, ATHERI NOPSIDAE), WITH COMMENTS ON THE SYSTEMATICS OF THE SOUTH AMERICAN FRESHWATER FISH GENUS BASIL ICHTHYS GIRARD BRIAN S. DYER MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 185 YGi yr~.-.i,$; ?!;< * ,:. ,'$I : :,.:. , :/:J:,;;<..;!".=:,,T,:; '.;F;,pqT$$;-:!yY[. ,, .:: z!.::,,.~\.~;i,.;~?;!i%~.;:,<..3-~ r:.l-j9 .-:.:.. c. :L.,,t:..:r .!::. !.Is> :;.$<' i 1 ;;' T*,' ,,... ..;-;i-,,;:b+:j?T. ,. ..7~,?ti~+~~~.?+, .* -5~~.>>.:~~;,.~:w~~~ : , . .:.;;::,i3.,g Ann Arbor, November 8,1997 ISSN 076-8405 MIS(:ELIANEOUS PUBLI<'ATIONS MUSEUM OF ZOOI,O(;Y, UNIVERSITY OF MI(:HI(;AN NO. 185 7I 7 he publications of the Muser~rrlof' Zoology, The Univrrsity of Miclliga~~,co~~sist primarily ottwo series-tlrc Occi~sionalPapers and thc Miscellaneous Publications. Both series were fol~ndedby I)n Bryant Walkcl; Mn B~adshawH. Swales, and Dr. W.W. Newcomb. Occasionally the Musru~ript~blislics co~ltributionsoutside of tlirsc series; bcginni~~gin 1990 these are titled Sprci;il Pl~blicationsand al-r numbel-cd. All sr~blnittrdmanr~scripts receive extern;ll rrview. 71'lrc Miscella~ieousI'ublications, whicli include monographic stndics, papers on fieltl and Inti- scu~rltechr~iqurs, anel other coritributio~lsnot within tlir scope of the Occasio~lalPapers, are pl11~- lishetl separ;~tely.It is not intcndecl that tl~ryhe grol~pcdinto volunies. Each number has a title page and, when necessary, il table ofcontrnts. The Occasional Pal)ers, pl~hlicationof which was I>egun in 1913, srl-ve as a medium for origirlal studies Ixlsetl principally upon the collections in the Museum. They are issued sepal-ately. Wlrcn a sr~fficir~~tnuml~cr of pages has been print(*d to ~nakca volumr, ;I title pagr, table of'contet~~s,and an index arc, sr~ppliedto 1il)rarics ancl individuals 011 the rnaili~iglist for the scrics. A conlplete list of'publications on Birds, Fishes, Insects, Mammals, Mollr~sks,Keptilcs and Anlphib ians, and other topics is available. Address inquiries to thc Director, Mnscr~~nof Zoolo~y,The Univc~ sity of Michigan, Ann Arl~or,Miclligan 48109-1079. 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<itricli, M.(:. 1!)!)1. I'l~ylogr~rrticanalysis of ;~\lianIli~lillin~l) rnr~scul;~t~~r-r. Misc. I'u111. 171). 8!) pp, 3 figs. $9.60. Kluge, A.G. 191) 1. Roilre s~~akcpl~ylogeny and rcseai.clr cyclcs. Misc. 1'111)l. 178. (i2 1111, 14 tigs. $6.00. l'rt1111, R.O. 1!)!)0. A tcst oftl~e~nonophyly of t11c Manaki~ls(Pipridac) ant1 ofthr (:otinjias ((:otingid;rc) 11;lscd on mol-phology. Occ. Pi~p.723. 44 LIP,6 figs. $3.70. Srrttkr~s,R.1). a~~d1i.M. Kailcy. I!)!)O. (:II;II-actrrs, rrlatio~~ships,distriI,ution, a~~dI,iolo~y of No/rojiic mrl/ino.s(ornic.s,a reccx~ltly~~;un~r.tl cypl-inid fish frorn sor~theastcr-11LJnitctl St;~tcs.Occ. I';III. 722. 15 pp, 3 figs. $2.20. Mycrs, I: J.1,. Pitito~l,and M.F. Srr~itlr.1!)!)0. A ~.cvirwol the /)oli~i~~~.si\gror~l) of 121to/lor1 (Mr~t-itlae: Sign~oclo~~ti~rac),witll rmph;rsis on 1'(.1.u ;111d Rolivii~.108 pl), 29 figs. $13.20. 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 Basilichthys 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.