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Phylogenetic Interrelationships of the Stomiid Fishes (Teleostei: Stomiiformes)

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Phylogenetic Interrelationships of the Stomiid Fishes (Teleostei: Stomiiformes) MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, THE UNIVERSITY OF MICHIGAN NO. 171 Phylogenetic Interrelationships of the Stomiid Fishes (Teleostei: Stomiiformes) by William L. Fink Division of Biological Sciences and Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109-1079 Ann Arbor MUSEUM OF ZOOLOGY, THE UNIVERSITY OF MICHIGAN December 3 1, 1985 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 171 The publications of the Museum of Zoology, The University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, initiated in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to form a volume, the Museum will supply a title page, table of contents, and an index to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, were established in 1916 and are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan 48109-1079. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOI,OGY, THE UNIVERSITY OF MICHIGAN NO. 171 Phylogenetic Interrelationships of the Stomiid Fishes (Teleostei: Stomiiformes) by William L. Fink Division of Biological Sciences and Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109- 1079 Ann Arbor MUSEUM OF ZOOLOGY, THE UNIVERSITY OF MICHIGAN December 3 1, 1985 ABSTRACT Fink, Willzanl L. 1985. Phylogenetic Interrelatiomhz of the Stomiid Fishes (Teleostei: Stomiformes). Mzsc. Publ. MW. 2001.Unzv. Michigan, 171:l-127, frontic.,fig.r. 1-70. A phylogenetic analysis of the interrelationships of the twenty-five stomiiform genera formerly classified in the superfamily Stomiatoidea is presented. Previously comprising the families Astronesthidae, Chauliodontidae, Idiacanthidae, Malacosteidae, Melanostomiatidae, and Stomiidae, the genera are placed in a single, expanded Stomiidae. All of the traditionally recognized genera are toutid to be monophylctic, with the exception of Stomias, which is expanded to include Macrostomiar. The character text includes detailed descriptions of stomiid morphology. Several alternative phylogenetic hypotheses are discussed. In the context of orre or more of these hypothcscs, patterris of character transformation are discussed, includitrg the evolution of the pectoral girdle and its accessory light orgaris, placement of the vertical firis toward the posterior of the body, elaboration of the anterior and posterior portions of the pelvic girdle, increase and subsequent decrcase of the number of radial elements of the pelvic girdles, and specializations of the head skeleton and anterior axial skeleton. 'fie data matrix that was used to generate the phylogenetic hypotheses, and a brief discussion of fossils which have been assigned to the Stomiidae, are included in appendices. Key words: Stomiiformes, Astrone.tthidar, Chauliodontidae, Idiacanthidae, Malacosteidae, Mrlanostomiidae, Stonlizdae, phylogenetics, character analysis. FKONTISPIECE:Malacosteus (top), Emtomias (bottom). From Zugmayer, 191 1. CON'I'EN'I'S PAC; E'. IN'fKODUCTION ............................................................. 1 tIISI'OKY OF CI.ASSIFIC.KI'ION 01:THE S'I'OM1II)AE ......................... 3 MI.. '1'HOI)S ................................................................... 8 KF.SUI.1.S ..................................................................... 10 (;IIARA(:TEKS ................................................................ 10 Suspcnsor~ium;tntl laws ..................................................... 27 l%~-iu~chialBasket atitl Hyoid ................................................ Muscles and IAig;imentsof thc fle;~tl......................................... I'osccrarrial Axial Skeleton ................................................... Vcl.tic.11Fins ............................................................... I'cctor.al (;irtllc ............................................................. I'clvic<;i~-dlr............................................................... 1.iglitOrgans .............................................................. Misccllancous (.har.actcrs ................................................... I)IS(:USSION ................................................................. Sio~niicl1ntcrt.clationsllips ................................................... Sfi (.hal.actcr Evolutiorr ........................................................ 107 A(:KNOWLED(;MENTS ....................................................... 117 I~I'I'EKKI~UKL'. (.I'f61) ......................................................... 118 Al'l'kNDIX 1. LIAlli\ MKI'KIX ................................................. 121 A1'I'ENI)IX 2 . F0SSII.TEI.EOSfS I'KEVI0USI.Y REE'ERREI) -f<) 'I'HE SI'OMIIDAE 124 A1'I'k:NI)IX 3 . MXI'BKIAI . BXAMIN I.: D .......................................... 125 1 LLUSTRATIONS FIGURE PAC; E 1 Summary of intenelationships among stomiid genera ...................... I I 2-6 Partial clatlograms showing alternative resolutions of surnmary diagram and orre possihle interpretation of character support ........................... 12-15 7 Ncurocratlium of Malacostru ............................................ 16 8 Neurocrarliurn of Photo.stomia\ ............................................ 17 9 Neu rocraniurn of Aristu.stomi(~\........................................... 18 10 Neurocranium of Pacl~yystowiia.~........................................... 19 1 1 Neurocrani~~mof' I'lagello.sto~nicw.......................................... 20 12 Neurocraniurn of' Eustomia\ .............................................. 21 13 Neurocraniurn of Ratltophzluv............................................ 22 14 Neurocranium of Ecltiostonia ............................................. 23 15 Neurocranium of Melanostomzc~s.......................................... 24 16 Neurocraniu~nof Photonectes............................................. 25 17 Neurocranium of Stow~ias................................................ 26 18 Jaws ant1 suspensoriurn of Echiostoma ..................................... 27 19 jaws and suspensorium of Tactoston~c~..................................... 28 20 Jaws and suspensori~~rr~of Bathophil~c .................................... 259 2 1 Jaws and suspcnsoriun~of Stor~~ias ........................................ 30 22 Jaws and suspcnsoriurrl of Trigono1amj)a................................... 31 23 Jaws and suspensorium of Gmmmatostowiia.~................................ 32 24 Jaws and suspensoriurn of Ezwtow~ius...................................... 33 25 Jaws and suspensorium of Pholostomicw .................................... 34 26 Jaws and suspensorium of Malarostew .................................... 35 27 Jaws ancl suspensorium of P(~clty.stomia.s.................................... 36 28 Jaws and suspensoriu~nof Leptostomzrw .................................... 37 29 Jaws and suspcnsoriurn of Cttirostow~ia.~................................... 38 30 .Jaws and suspensoriurn of Photonectes ..................................... 39 3 1 H yoid arches of Stomirw. Trigonolampa. Odontostomzas. Neonr.ct/tc.c, and Grammato.stomicw. ....................................................... 41 32 Hyoid arches of Mrlanostomiac. I'hotonecte.\. Rustumias. I'achy.stomin.s, and Photoctomirw............................................................ 43 33 Anterior vertebral region of Ewtowizas .................................... 57 34 Anteriol- vertebral region of Stomuw ...................................... 58 35 Part of'the axial and vertical-fin skeleton of Idiacanthw ..................... 60 36 Vertical-fin supports and posterior portion of axial skeleton of Trzgonolamf)~. 61 37 Some dorsal-fin pterygiophores of Tngonolr~mpa........................... 62 Vertical-fin supports and posterior portion of axial skeleton of Photostomius ... 63 Pectoral girdle of Bathophil~s............................................ 64 Pectoral girclle of Arictostomiu~............................................ 65 Pectoral girdle of Cirostomias ............................................ 66 Pectoral girdle of Odontostomim.......................................... 67 Pectoral girdle of I.eptostomic~s............................................ 68 Pectoral girdle of Melano.ctomim. ......................................... 69 Pectoral girdle of Ewtomim cf . mcntrws ................................... 70 Pectoral girdle of Ewtomias cC breuibarbatw ................................ 71 Pectoral girdle of Pachy.ystomias............................................ 72 Pectoral girdle of Chauliodus............................................. 72 Pectoral girdle of Polynietme .............................................. 73 PectoralgirdleofStomim................................................ 74 Anterior pectoral-fin ray articulation of Heterophotw ....................... 75 Anterior pectoral-fin ray articulation of
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