A peer-reviewed version of this preprint was published in PeerJ on 24 February 2015.
View the peer-reviewed version (peerj.com/articles/769), which is the preferred citable publication unless you specifically need to cite this preprint.
Werneburg I. 2015. Morphology of the jaw, suspensorial, and opercle musculature of Beloniformes and related species (Teleostei: Acanthopterygii), with a special reference to the m. adductor mandibulae complex. PeerJ 3:e769 https://doi.org/10.7717/peerj.769 Morphology of the jaw, suspensorial, and opercle musculature of Beloniformes and related species (Teleostei: Acanthopterygii), with a special reference to the m. adductor mandibulae complex
The taxon Beloniformes represents a heterogeneous group of teleost fishes that show an extraordinary diversity of jaw morphology. I present new anatomical descriptions of the jaw musculature in six selected beloniforms and four closely related species. A reduction of the external jaw adductor (A1) and a changed morphology of the intramandibular musculature were found in many Beloniformes. This might be correlated with the progressively reduced mobility of the upper and lower jaw bones. The needlefishes and PrePrints sauries, which are characterised by extremely elongated and stiffened jaws, show several derived characters, which in combination enable the capture of fish at high velocity. The ricefishes are characterised by several derived and many plesiomorphic characters that make broad scale comparisons difficult. Soft tissue characters are highly diverse among hemiramphids and flying fishes reflecting the uncertainty about their phylogenetic position and interrelationship. The morphological findings presented herein may help to interpret future phylogenetic analyses using cranial musculature in Beloniformes.
PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 TITEL PAGE Morphology of the jaw, suspensorial, and opercle musculature of Beloniformes and related species (Teleostei: Acanthopterygii), with a special reference to the m. adductor mandibulae complex Ingmar Werneburg1,* 0XVHXPIU1DWXUNXQGH /HLEQL],QVWLWXWIU (YROXWLRQV %LRGLYHUVLWlWVIRUVFKXQJ DQ GHU+XPEROGW8QLYHUVLWlW]X%HUOLQ,QYDOLGHQVWUDH%HUOLQ*HUPDQ\ LZHUQHEXUJ#JPDLOFRP Running title:-DZPXVFOHVRI%HORQLIRUPHV Key words 2U\]LDV%HORQH3HUFD$WKHULQRPRUSKDIHHGLQJMDZHORQJDWLRQ PrePrints
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'RUVDOO\WRWKHLQVHUWLRQRIPLQWHUPDQGLEXODULVDW K\RLGHL PLQWHUPDQGLEXODULVWRWKH WKHGHQWDU\DQGFRYHUVDWOHDVWWKHSRVWHURGRUVDO PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 GHQWDU\>VWDWH@ DUHDRIWKHODWWHUPXVFOH>VWDWH@ /LJDPHQWV /LJ $EVHQW>VWDWH@ ! 3UHVHQW>VWDWH@ SDUDVSKHQRLGR VXVSHQVRULXP PrePrints PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Perca fluviatilis S1A) m. adductor arcus palatini dermosphenotic m.M. dilatator dilatator extrasca-Extrasca- m. epaxialis (anterior m. adductor mandibulae (A1) autosphenotic operculioperculi pulariapularia epiotic portion) connective tissue pterotic m. levator parasphenoid m. trapezius operculi frontal mesopterygoid proc. lateralis m. levator lateralethmoid hyomandibu- arcus palatini laris lig. mesopterygoido- lateralethmoid connective tissue nasal mesethmoid lig. praemaxillo-frontal connective tissue connective tissue lacrimal lig. parasphenoido-suspensorium praemaxillar tendon of m. adductor mandibulae (A1) lig. maxillo-dentary maxilla lig. praemaxillo-maxilla dentary ectopterygoid PrePrints subopercle anguloarticular circumorbitals opercle lig. maxillo-anguloarticular m. adductor mandibulae (A2/3, praeopercle quadrate m. adductor mandibulae (A2/3, medial) lateral) interopercle Perca fluviatilis S1B) mesopterygoid m. levator arcus palatini m. dilatator operculi m. trapezius m. adductor dermosphenotic m. levator operculi lig. parasphenoido-suspensorium arcus palatini connective tissue (anterior part) frontal pterotic nasal autosphenotic parasphenoid mesethmoid lateralethmoid lig. mesopterygoido-lateralethmoid ramus maxillaris trigemini lig. palato-praemaxilla palatine premaxilla maxilla tendon of m. adductor mandibulae (A1) dentary lig. maxillo-anguloarticular ramus mandibularis trigemini anguloarticular proc. quadrate lateralis hyomandibularis opercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2ectopterygoid | CC-BY 4.0metapterygoid Open Access | rec: 20 Jan 2015, publ: 20interopercle Jan 2015 m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, lateral) praeopercle Perca fluviatilis 6& parasphenoid m. adductor arcus palatini m. levator arcus palatini hyomandibular lig. parasphenoido- suspensorium mesopterygoid metapterygoid ectopterygoid anguloarticular quadrate symplectic 5 mm lig. mandibulo-interopercular PrePrints ramus mandibularis facialis praeopercle interopercle Perca fluviatilis 6' parasphenoid n. hyomandibularis ramus hyoideus facialis mesopterygoid m. levator arcus palatini canalis facialis m. adductor arcus hyomandibular frontal palatini lateralethmoid lig. parasphenoido-suspensorium nasal mesethmoid cartilago rostri lig. palato-praemaxilla premaxilla palatine dentary m. intermandibularis m. protractor hyoidei intramandibular portion of m. adductor ectopterygoid PDQGLEXODH $ 11S quadrate preopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2ramus mandibularis facialis| CC-BY 4.0 Openanguloarticular Access | rec: 20 Jan 2015, publ:metapterygoid 20 Jan 2015 PDGGXFWRUPDQGLEXODH $Ȧ lig. mandibulo-interopercular symplectic Rhinomugil corsula S2A) parasphenoid m. dilatator operculi m. levator operculi epiotic m. adductor arcus palatini m. levator arcus palatini pterotic extrascapularia circumorbitalia proc. sphenoticus sphenotic parietal mesopterygoid G frontal F H lateralethmoid E nasal mesethmoid premaxilla maxilla E dentary F palatine G connective tissue H lacrimal PrePrints lig. maxillo-dentary anguloarticular interopercle opercle quadrate m. adductor mandibulae (A1) subopercle lig. mandibulo-interoperculare m. adductor mandibulae (A2/3, lateral) preopercle Rhinomugil corsula S2B) m. adductor arcus palatini parasphenoid proc. sphenoticus m. levator operculi sphenotic proc. lateralis ramus maxillaris trigemini m. levator arcus palatini hyomandibularis H frontal G nasal F lateralethmoid mesethmoid palatine maxilla premaxilla tendon of m. adductor mandibulae (A1) lig. maxillo-dentary ramus mandibularis trigemini dentary F ramus mandibularis externus trigemini G m. adductor mandibulae (A2/3, lateral) PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ:H 20 Jan 2015 m. adductor mandibulae (A2/3, intermedial) quadrate interopercle opercle anguloarticular lig. mandibulo-interopercle preopercle Rhinomugil corsula S2C) blood vessel ramus maxillaris trigemini proc. sphenoticus hyomandibular F G parasphenoid H metapterygoid m. adductor arcus palatini mesopterygoid ramus mandibularis trigemini ramus mandibularis externus trigemini anguloarticular quadrate symplecticum F G lig. mandibulo- H PrePrints interopercular interopercle ramus mandibularis facialis preopercle Rhinomugil corsula S2D) intramandibular portion of m. adductor mandibulae (A2/3) m. adductor mandibulae (A2/3) ramus mandibularis trigemini (dorsal branch) mesopterygoid m. intermandibularis quadrate dentary G F E E F m. protractor hyoidei anguloarticular G preopercle ramus mandibularis facialis ramus mandibularis facialis interopercle PDGGXFWRUPDQGLEXODH $Ȧ lig. mandibulo-interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Rhinomugil corsula EFG H 5 mm Rhinomugil corsula nasal S2E) mesethmoid lateralethmoid lacrimal maxilla premaxilla m. intermandibularis cartilago Meckeli PrePrints dentary m. protractor hyoidei frontal bulbus olfactorius Rhinomugil corsula lens S2F) parasphenoid m. adductor arcus palatini palatine branches of ramus maxillaris trigemini lacrimal m. adductor mandibulae (A1) ramus mandibularis trigemini (dorsal branch) PDGGXFWRUPDQGLEXODH $Ȧ two heads basihyal branches of ramus mandibularis facialis cartilago Meckeli dentary m. protractor hyoidei PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 frontal bulbus olfactorius Rhinomugil corsula S2G) lens parasphenoid tractus opticus m. adductor arcus palatini ramus maxillaris trigemini circumorbital ramus mandibularis trigemini ramus mandibularis trigemini externus m. adductor mandibulae (A1) m. adductor mandibular (A2/3, medial) m. adductor mandibulae (A2/3, lateral) mesopterygoid PDGGXFWRUPDQGLEXODH $Ȧ quadrate PrePrints ceratohyal ramus mandibularis facialis dentary / anguloarticular cartilago Meckeli Rhinomugil corsula m. protractor hyoidei S2H) frontal sphenoticum m. dilatator operculi proc. sphenoticus brain with tractus opticus truncus maxillaris infraorbitalis trigemini parasphenoid m. levator arcus palatini m. adductor arcus palatini hyomandibular m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, lateral) m. adductor mandibulae (A1) praeopercle ramus mandibularis facialis interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Atherina boyeri S3A) lig. parasphenoido- m. levator arcus palatini m. levator operculi m. epaxialis (anterior por- suspensorium proc. lateralis hyomandibularis pterotic extrascapularia tion) mesopterygoid m. adductor arcus palatini m. dilatator operculi m. trapezius parasphenoid autosphenotic frontal dermosphen- I otic connective tissue H nasal G F mesethmoid lateralethmoid lacrimal ramus maxillaris trigemini palatine circumorbitals lig. premaxillo-maxilla F G lig. premaxillo-dentary H premaxilla I lig. maxillo- PrePrints anguloarticular preopercle opercle maxilla anguloarticular m. adductor mandibulae (A1) subopercle dentary quadrate lig. mandibulo-interopercle interopercle Atherina boyeri S3B) connective tissue premaxilla maxilla frontal nasal lig. palato- lig. palato- membrans nasal maxilla G F lateralethmoid mesethmoid palatine parasphenoid lig. parasphenoido- suspensorium mesopterygoid ramus maxillaris trigemini m. adductor mandibulae (A2/3, lateral portion) m. adductor mandibulae (A1) - with tendons to: maxilla, to F lacrimal and to anguloarticular G lig. premaxillo- lig. premaxillo- dentary anguloarticular lig. maxillo-anguloarticular PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 dentary maxilla Atherina boyeri S3C) m. adductor arcus palatini m. levator arcus palatini dermosphenotic proc. lateralis hyomandibularis G H I lig. parasphenoido-suspensorium parasphenoid mesopterygoid ramus maxillaris trigemini ramus mandibularis trigemini palatine lateral portion of m. adductor mandibulae (A2/3) lig. maxillo-anguloarticular quadrate anguloarticular PrePrints lig. mandibulo-interopercle G H I medial portion of medial portion of m. adductor mandibulae (A2/3), m.adductor mandibulae (A2/3), preopercle medial head lateral head interopercle Atherina boyeri S3D) parasphenoid m. adductor arcus palatinim. levator arcus palatini proc. lateralis hyomandibularis H I lig. parasphenoido-suspensorium ramus maxillaris trigemini mesopterygoid ramus mandibularis trigemini metapterygoid quadrate H I symplectic ramus mandibularis facialis preopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Atherina boyeri 6( mesopterygoid canalis facialis ramus hyoideus facialis m. levator operculi m. levator arcus palatini hyomandibular subopercle frontal m. adductor arcus palatini n. hyomandibularis opercle nasal mesethmoid lig. premaxillo-frontal cartilago rostri I F premaxilla H palatine G dentary m. intermandibularis internal head of medial portion of m. adductor mandibulae $ m. protractor hyoidei F G I lateral portion of H PDGGXFWRUPDQGLEXODH $ PrePrints PDGGXFWRUPDQGLEXODH $Ȧ quadrate praeopercle interopercle metapterygoid branches of ramus mandibularis facialis lig. mandibulo- symplectic anguloarticular interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Atherina boyeri FGHI 5 mm Atherina boyeri S3F) nasal lig. premaxillo-frontal lacrimal maxilla premaxilla lig. praemaxillo-dentary m. intermandibularis dentary PrePrints m. protractor hyoidei frontal parasphenoid lens Atherina boyeri S3G) mesopterygoid ceratohyal palatine / mesethmoid tendon of m. adductor mandibulae (A1) to maxillary lateral portion of m. adductor mand. (A2/3) tendon of M. adductor mandibulae (A1) to lacrimal tendon of m. adductor mandibulae (A1) to anguloarticular circumorbital 2 mm lig. maxillo-anguloarticular intramandibular head of the medial portion of m. adductor mandibulae (A2/3) dentary / anguloarticular m. protractor hyoidei PDGGXFWRUPDQGLEXODH $Ȧ cartilago Meckeli PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 frontal Atherina boyeri m. trapezius S3H) pterotic vitreous body brain with tractus opticus parasphenoid mesopterygoid / metapterygoid m. adductor arcus palatini ramus maxillaris trigemini ramus mandibularis trigemini medial portion of m. adductor mandibulae (A2/3), medial head medial portion of m. adductor mandibulae (A2/3), lateral head lateral portion of m. adductor mandibulae (A2/3) external section of m. adductor mandibulae (A1) PrePrints ceratohyal preopercle / symplectic Atherina boyeri m. protractor hyoidei S3I) interopercular parietal m. trapezius pterotic m. dilatator operculi brain with n. trigeminus hyomandibular m. levator arcus palatini parasphenoid m. adductor arcus palatini ramus maxillaris trigemini ramus mandibularis trigemini medial portion of m. adductor mandibulae (A2/3), lateral head m. adductor mandibulae (A1) preopercle / symplectic / metapterygoid interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Aplocheilus lineatus S4A) frontal truncus maxillaris m. dilatator operculi m. epaxialis (anterior portion) palatine supraorbitalis trigemini m. levator arcus palatini m. levator operculi ramus maxillaris trigemini parasphenoid proc. sphenoticus pterotic mesopterygoid m. adductor autosphenotic connective tissue arcus palatini dermosphen- H nasal otic G connective tissue lig. palato-maxilla lacrimal F maxilla premaxilla E lig. premaxillo-mandibular E dentary lig. premaxillo-dentary F Lig. maxillo-dentale lig. premaxillo-maxillary G anguloarticular H quadrate opercle lig. mandibulo- interopercle m. adductor mandi- truncus maxillaris proc. lateralis PrePrints interopercle bulae (A2/3, medial) infraorbitalis trigemini hyomandibularis m. adductor m. adductor mandibulae ramus mandibularis subopercle mandibulae (A1) (A2/3, lateral) trigemini preopercle Aplocheilus lineatus S4B) parasphenoid truncus maxillaris dermosphenotic epaxialis (anterior portion) m. adductor supraorbitalis trigemini m. levator arcus palatini m. levator operculi arcus palatini autosphenotic m. dilatator operculi pterotic proc. sphenoticus frontal G H lig. lateralethmoido-frontal lateralethmoid mesethmoid ramus maxillaris trigemini mesopterygoid F nasal lig. palato-maxilla palatine E maxilla premaxilla lig. premaxillo-dentary dentary lig. premaxillo-mandibular m. adductor mandi- bulae (A1) lig. maxillo-dentary E anguloarticular lig. premaxillo-maxillary F ramus mandibularis externus trigemini G ramus mandibularis truncus H opercle trigemini maxillaris preopercle quadrate interopercle infraorbitalis trigemini m. adductor mandibulae, PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2lig. mandibulo- m. adductor mandibulae, | CC-BY 4.0m. Open adductor Access mandibulae, | rec: 20 Jan 2015, (A2/3, publ: 20lateral) Jan 2015 interopercle (A2/3, intermedial) (A2/3, medial) proc. lateralis hyomandibularis Aplocheilus lineatus S4C) ramus maxillaris m. adductor truncus maxillaris m. levator arcus palatini ramus mandibularis trigemini arcus palatini infraorbitalis trigemini hyomandibular facialis mesopterygoid G H dentary m. adductor mandibulae (A2/3, lateral) anguloarticular ramus mandibularis externus trigemini ramus mandibularis trigemini quadrate lig. mandibulo- G H interopercle interopercle m. adductor mandibulae m. adductor mandibulae preopercle PrePrints (A2/3, intermedial) (A2/3, medial) symplectic metapterygoid Aplocheilus lineatus S4D) truncus maxillaris hyomandibular infraorbitalis trigemini canalis facialis mesethmoid m. adductor arcus palatini n. hyomandibularis frontal facialis H nasal F maxillary premaxillary scleral plate dentary E G m. intermandibularis E m. protractor hyoidei F G H PDGGXFWXUPDQGLEXODH $Ȧ branches of ramus intramandibular portion of branches of ramus hyoideus facialis m. adductor mandibulae (A2/3) preopercle palatine mandibularis facialis metapterygoid lig. mandibulo- symplecticum mesopterygoid interopercle ramus maxillaris quadrate anguloarticular trigemini PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Aplocheilus lineatus E FGH 5 mm maxilla premaxilla Aplocheilus lineatus 6( m. intermandibularis m. protractor hyoidei PrePrints dentary cartilago Meckeli mesethmoid frontal m. adductor arcus palatini ceratohyal palatine tendon of m. adductor Aplocheilus lineatus PDQGLEXODH $ WRSDODWLQH 6) lig. premaxillo-mandibular ) lacrimal maxilla premaxilla intramandibular portion of PDGGXFWRUPDQGLEXODH $ dentary / anguloarticular m. protractor hyoidei cartilago Meckeli PDGGXFWRUPDQGLEXODH $Ȧ PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 frontal brain parasphenoid truncus maxillaris supraorbitalis trigemini m. adductor arcus palatini lens Aplocheilus lineatus ramus maxillaris trigemini S4G) metapterygoid G) m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, intermedial) aponeurosis of m. adductor mandibulae (A2/3) ramus mandibularis trigemini m. adductor mandibulae (A2, lateral) symplectic ramus mandibularis facialis m. adductor mandibulae (A1) ceratohyal praeopercle PrePrints interopercle frontal brain with n. trigeminus labyrinth organ Aplocheilus lineatus m. trapezius pterotic S4H) H) truncus maxillaris supraorbitalis trigemini m. dilatator operculi truncus maxillaris infraorbitalis trigemini parasphenoid sphenoticum autosphenoid / dermosphenoid m. adductor arcus palatini hyomandibular m. levator arcus palatini m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, lateral) metapterygoid m. adductor mandibulae (A2/3, intermedial) preopercle m. adductor mandibulae (A1) interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Oryzias latipes S5A) lateralethmoid m. adductor prootic m. levator arcus palatini m. epaxialis (anterior portion) arcus palatini m. dilatator operculi frontal m. levator operculi dermosphenotic parasphenoid pterotic nasal sphenotic meso- lacrimal pterygoid hyomandi- G bular mesethmoid F maxilla premaxilla E palatine D dentary lig. maxillo- mandibulare anguloarticular D retroarticular E quadrate F m. protractor hyoidei ramus G ramus mandibularis mandibularis opercle externus trigemini trigemini PrePrints subopercle ramus mandibularis symplectic interopercle preopercle externus facialis m. adductor mandibulae truncus maxillaris m. adductor mandibulae (A1) (A2/3, lateral) infraorbitalis trigemini Oryzias latipes S5B) ramus maxillaris truncus maxillaris m. adductor arcus parasphenoid hyomandibular trigemini infraorbitalis trigemini palatini mesopterygoid F G ramus mandibularis externus trigemini quadrate ramus mandibularis trigemini m. adductor mandibulae (A1) quadrate G F PeerJ PrePrintsinteropercle | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 ramus mandibularis symplectic metapterygoid (?) externus facialis preopercle Oryzias latipes S5C) m. adductor arcus palatini n. hyomandibularis facialis hyomandibular (posterior portion) (intermedial portion) metapterygoid (anterior portion) G mesopterygoid palatine anguloarticular E F dentary D m. intermandibularis E F G praeopercle D ramus hyoi- m. protractor deus facialis hyoidei retroarticular quadrate symplectic interopercle m. adductor lig. mandibulo- ramus mandibularis PrePrints mandibulae (AȦ) interoperculare internus facialis PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Oryzias latipes DE F G 1 mm S5D) Oryzias latipes premaxilla maxilla dentary m. intermandibularis with tendon PrePrints m. protractor hyoidei frontal mesethmoid nasal palatine maxilla premaxilla S5E) Oryzias latipes lig. praemaxillo-maxillare tendon of m. adductor mandibulae (A1) m. adductor mandibulae (A1) tendon of m. adductor mandibulae (A2/3) anguloarticular / dentary quadrate connective tissue m. adductor mandibulae (AȦ) basihyal m. protractor hyoidei PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 brain frontal prootic parasphenoid m. adductor arcus palatini Oryzias latipes ceratohyal S5F) lens m. adductor mandibulae (A2/3, medial/intermediate) ramus mandibularis trigemini tendon of m. adductor mandibulae (A2/3, lateral) ramus mandibularis trigemini (branch to A1) mesopterygoid m. adductor mandibulae (A1) quadrate / symplectic interopercle m. protractor hyoidei PrePrints frontal brain pterotic labyrinth Oryzias latipes epaxial musculature S5G) sphenotic m. levator operculi prootic dermosphenotic m. levator arcus palatini truncus maxillaris infraorbitalis trigemini m. adductor arcus palatini (anterior portion) hyomandibular m. adductor mandibulae (A2/3, lateral) parasphenoid preopercle interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Parexocoetus brachypterus S6A) m. levator arcus palatini proc. lateralis hyomandibularis m. dilatator operculi epiotic m. epaxialis m. adductor arcus palatini sphenoticum m. levator operculi (anterior portion) truncus maxillaris postorbital pterotic posttemporal infraorbitalis trigemini H mesopterygoid G parasphenoid lateralethmoid frontal F ramus maxillaris trigemini mesethmoid maxilla connective tissue lacrimal E premaxilla lig. premaxillo- dentary barbel maxilla E dentary palatine connective tissue F anguloarticular ramus mandibularis trigemini lig. mandibulo-interopercle PrePrints G lig. quadrato-preopercle H quadrate m. adductor mandibulae (A2/3, lateral) preopercle interopercle subopercle opercle Parexocoetus brachypterus S6B) premaxilla tendinous membranesnasal frontal mesethmoid lig. palato-maxilla F lateralethmoid parasphenoid mesopterygoid ramus maxillaris trigemini truncus maxillaris infraorbitalis trigemini ramus mandibularis trigemini palatine m. adductor mandibulae (A2/3, lateral) quadrate E lig. quadrato-preopercle preopercle E F interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 lig. premaxillo- dentarymaxilla barbellig. maxillo- lig. mandibulo- anguloarticular dentary dentary interopercle Parexocoetus brachypterus S6C) parasphenoidm. adductor arcus palatini postorbital m. levator arcus palatini hyomandibular truncus maxillaris G H infraorbitalis trigemini mesopterygoid ramus maxillaris trigemini ramus mandibularis trigemini palatine anguloarticular m. adductor mandibulae (A2/3, lateral) quadrate lig. quadrato-preopercular lig. mandibulo-interopercle interopercle G PrePrints H m. adductor mandibulae (A2/3, medial) metapterygoid ramus mandibularis facialis symplectic preopercle Parexocoetus brachypterus S6D) lateralethmoid mesopterygoid m. adductor canalis facialis n. hyomandibularis hyomandibular arcus palatini facialis frontal H nasal mesethmoid F maxilla lig. maxillo-mesethmoid premaxilla palatine lig. premaxillo- dentary E G dentary m. protractor hyoidei m. intermandibularis barbel E PDGGXFWRUPDQGLEXODH $Ȧ quadrate F anguloarticular G HS lig. mandibulo- ramus mandibularis metapterygoid symplectic interopercle ramus hyoideus PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2interopercle facialis | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20facialis Jan 2015 Parexocoetus brachypterus E FHG 5 mm Parexocoetus brachypterus 6( lig. premaxillo-dentary dentary m. protractor hyoidei PrePrints cartilago Meckeli m. intermandibularis barbel frontal bulbus olfactorius Parexocoetus brachypterus nasal 6) brain mesethmoid parasphenoid palatine lig. palato-maxilla maxilla lacrimal basihyal PDGGXFWRUPDQGLEXODH $ m. protractor hyoidei dentary / anguloarticular cartilago Meckeli PDGGXFWRUPDQGLEXODH $Ȧ PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Parexocoetus brachypterus S6G) frontal brain lens parasphenoid m. adductor arcus palatini truncus maxillaris infraorbitalis trigemini mesopterygoid m. adductor mandibulae (A2/3, lateral) m. adductor mandibulae (A2/3, medial) preopercle interopercle frontal brain pterotic labyrinth organ Parexocoetus brachypterus n. trigeminus PrePrints S6H) truncus maxillaris supraorbitalis trigemini sphenotic truncus maxillaris infraorbitalis trigemini m. dilatator operculi sphenotic m. adductor arcus palatini m. levator arcus palatini prootic parasphenoid hyomandibular m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, lateral) preopercle interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Dermogenys pussila S7A) frontal parasphenoid m. dilatator operculi m. epaxialis (anterior por- lateralethmoid m. adductor arcus palatini sphenotic m. levator operculi tion) postorbital lig. praemaxillo-frontal truncus maxillaris pteroticum infraorbitalis trigemini m. levator mesethmoid ramus maxillaris arcus palatini ramus ophthalmicus trigemini superfacialis trigemini G F D E D dentary E premaxilla F G maxilla nasal lacrimal anguloarticular preopercle opercle PrePrints lig. premaxillo-dentary mesopterygoid interopercle subopercle connective tissue m. adductor mandibulae superficial branching of ramus mandibularis trigemini (A2/3, lateral) ramus hyoideus facialis Dermogenys pussila S7B) parasphenoid metapterygoid frontal postorbital sphenotic pterotic truncus maxillaris infraorbitalis trigemini G F ramus maxillaris trigemini lateralethmoid mesethmoid premaxilla m. adductor arcus palatini ramus mandibularis trigemini mesopterygoid m. adductor mandibulae (A2/3, lateral) m. adductor mandibulae (A2/3, medial) anguloarticular F G PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2symplectic interopercle | CC-BYramus 4.0 Open mandi- Access | rec:preopercle 20 Jan 2015, publ:hyomandibular 20 Jan 2015 bularis facialis Dermogenys pussila 6& cartilago rostri mesopterygoid quadrate m. adductor arcus palatini premaxilla palatine mesethmoid symplectic E D F D E F dentary m. protractor hyoideiramus mandibularis ramus mandibularis interopercle trigemini facialis PrePrints m. intermandibularis PDGGXFWRUPDQGLEXODH $Ȧ anguloarticular lig. mandibulo-interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Dermogenys pussila D EF G 5 mm Dermogenys pussila 6' premaxilla m. protractor hyoidei cartilago Meckeli ramus mandibularis trigemini dentary m. intermandibularis PrePrints m. protractor hyoidei, ventral tendon frontal mesethmoid Dermogenys pussila lacrimal 6( nasal premaxilla maxilla dentary / anguloarticular m. adductor mandibulae (AȦ basihyale cartilago Meckeli ramus mandibularis trigemini m. protractor hyoidei PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 frontale parasphenoid Dermogenys pussila S7F) lens m. adductor arcus palatini truncus maxillaris infraorbitalis trigemini m. adductor mandibulae (A2/3, medial) aponeurosis of the fused m. adductor mandibulae (A2/3) m. adductor mandibulae (A2/3, lateral) quadrate ramus mandibularis facialis preopercle m. protractor hyoidei interopercle ceratohyal PrePrints Dermogenys pussila S7G) frontal sphenotic brain truncus maxillaris infraorbitalis trigemini m. levator arcus palatini parasphenoid m. adductor arcus palatini ramus mandibularis facialis hyomandibular m. adductor mandibulae (A2/3, lateral) praeopercle interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Belone belone S8A) parasphenoid m. adductor mandibulae m. dilatator pterotic lateralethmoid (A2/3, medial) operculi m. levator connective tissue postorbital m. levator operculi frontal arcus palatini m. adductor nasal proc. lateralis arcus palatini sphenoticum mesethmoid hyomandibularis connective tissue connective frontal tissue lig. premaxillo- frontal lacrimal metapterygoid m. adductor mandibulae (A2, lateral) opercule lig. premaxillo- sesamarticular m. adductor mandibulae PrePrints dentary anguloarticular (A2/3, intermedial) subopercle dentary lig. mandibulo-interoperculare mesopterygoid interopercle premaxilla quadrate connective tissue ramus mandibularis trigemini praeopercle Belone belone S8B) proc. coronoideus dentalis symplectic opercular connective tissue m. adductor mandi- m. levator operculi lateralethmoid bulae (A2/3, medial) pterotic frontal postorbital m. dilatator operculi nasal m. adductor m. levator arcus palatini mesethmoid arcus palatini sphenotic metapterygoid maxilla parasphenoid connective lig. premaxillo-maxilla tissue premaxilla lig. premaxillo- frontal dentary subopercle lig. premaxillo-dentary quadrate lig. maxillo-dentary hyomandibular lig. mandibulo- anguloarticular interopercle interopercle ramus mandibularis facialis mesopterygoid PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2ramus mandibularis | CC-BY 4.0 Open Access preopercle| rec: 20 Jan 2015, publ: 20 Jan 2015 metapterygoid trigemini quadrate sesamarticular m. adductor mandibulae (A2/3, lateral) m. adductor mandibulae (A2/3, intermedial) Belone belone 6& palatine metapterygoid hyomandibular frontal ramus mandibularis m. adductor mandibulae mesethmoid trigemini $PHGLDO ramus maxillaris mesopterygoid m. adductor trigemini arcus palatini maxilla premaxilla m. intermandibularis retroarticular preopercle m. protractor hyoidei quadrate interopercle PrePrints m. adductor mandibulae (AȦ anguloarticular symplectic dentary lig. mandibulo-interopercle ramus mandibularis facialis Belone belone 2 mm PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Scomberesox saurus S9A) lateralethmoid parasphenoid m. levator parietal arcus palatini lig. parasphenoido- pteroticum connective tissue suspensoric postorbitale m. levator operculi ramus maxillaris m. adductor proc. lateralis frontal trigemini arcus palatini hyomandibularis m. dilatator operculi mesopterygoid sphenotic nasal (G) (H) connective tissue (F) maxilla premaxilla lacrimal lig. premaxillo-dentary (F) (G) (H) dentary interopercle subopercle lig. mandibulo- opercle anguloarticular interopercle quadrate preopercle PrePrints connective tissue sesamarticular m. adductor mandibulae (A2/3, lateral) ramus mandibularis externus trigemini m. adductor mandibulae (A2/3, medial) ramus mandibularis trigemini m. adductor mandibulae (A2/3, intermedial) Scomberesox saurus S9B) frontal mesethmoid lateralethmoid parasphenoid lig. parasphenoido- suspensoric maxilla (F) premaxilla lig. premaxillo-maxilla lig. premaxillo-dentary lig. maxillo-dentary (F) PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 dentary anguloarticular mesopterygoid ramus maxillaris trigemini Scomberesox saurus S9C) metapterygoid m. adductor mandibulae (A2/3, medial) m. levator arcus palatini hyomandibular (G)(H) quadrate m. adductor mandibulae (A2/3, intermedial) (G) (H) PrePrints symplectic ramus mandibularis facialis preopercular Scomberesox saurus S9D) cartilago rostri palatine ramus mandibularis trigemini m. adductor mandibulae (A2/3, medial) opercle premaxillary mesethmoid mesopterygoid m. adductor arcus palatini m. levator operculi m. adductor maxilla lateralethmoid metapterygoid hyomandibular mandibulae frontal (AȦ (H) (G) (F) (E) (E) (F) (G) (H) dentary quadrate interopercle preopercle m. intermandibularis anguloarticular ramus mandibularis facialis ramus hyoideus facialis m. protractor hyoidei lig. mandibulo-interopercle symplectic subopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 Scomberesox saurus E F G H 2 mm Scomberesox saurus 6 ( m. protractor hyoidei dentary PrePrints m. intermandibularis nasal mesethmoid palatine lacrimal Scomberesox lig. premaxillo-maxilla saurus premaxilla 6 ) maxilla basihyal anguloarticular / dentary PDGGXFWRUPDQGLEXODH $Ȧ cartilago Meckeli m. protractor hyoidei PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015 frontal brain Scomberesox saurus sphenotic n. trigeminus S9G) m. levator arcus palatini prootic parasphenoid ramus mandibularis trigemini m. adductor arcus palatini m. adductor mandibulae (A2/3, medial) m. adductor mandibulae (A2/3, intermedial) m. adductor mandibulae (A2/3, lateral) ramus mandibularis facialis PrePrints preopercle hyomandibular / metapterygoid Scomberesox saurus interopercle frontal S9H) brain sphenotic ramus mandibularis trigemini m. levator arcus palatini prootic m. adductor arcus palatini m. adductor mandibulae (A2/3, lateral) m. adductor mandibulae (A2/3, intermedial) M. adductor mandibulae (A2/3, medial) preopercle / hyomandibular interopercle PeerJ PrePrints | http://dx.doi.org/10.7287/peerj.preprints.691v2 | CC-BY 4.0 Open Access | rec: 20 Jan 2015, publ: 20 Jan 2015