Friedman and Coates: Early morphological diversification of coelacanths

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Electronic Appendix A. Supplemental Morphological Data. Figure A1. Holopterygius nudus Jessen (P 7789a), latest Givetian-earliest Frasnian, Bergisch-Gladbach, Germany. Skull and pectoral girdle.

1 Electronic Appendix A. Supplemental Morphological Data. Figure A1. Holopterygius nudus Jessen (P 7789a), latest Givetian-earliest Frasnian, Bergisch-Gladbach, Germany. Skull and pectoral girdle. (a) Photograph of P 7789a; (b) interpretive drawing. Bones shaded in grey in (b) are preserved as impressions. Scalebar represents 10 mm. Abbreviations: ano, anocleithrum; chy, ceratohyal; cla, clavicle; cle; cleithrum; cop, principal coronoid; de, dentary; ect, ectopterygoid; entp, entopterygoid; ex, extracleithrum; hyp, buccohypophysial foramen; gs, indeterminate fragments of the gill skeleton; pmx, premaxilla; psoc, pores for the supraorbital sensory canal; psph, parasphenoid; sr, skull roof; sym, symplectic. 1

Bones shaded in grey in (b) are preserved as imprints. Scalebar represents 10 mm. Abbreviations: op, opercle; ot, otolith; qu, quadrate.

2 Figure A2. Holopterygius nudus Jessen (P 7789b), latest Givetian-earliest Frasnian, Bergisch-Gladbach, Germany. Skull and pectoral girdle, counterpart to P 7789a (figures 1, A1). (a) Photograph of P 7789b; (b) interpretive drawing. Bones shaded in grey in (b) are preserved as imprints. Scalebar represents 10 mm. Abbreviations: op, opercle; ot, otolith; qu, quadrate. Other abbreviations as in figure A1. Figure A3. Holopterygius nudus Jessen (P 7789c), latest Givetian-earliest Frasnian, Bergisch-Gladbach, Germany. Axial skeleton, counterpart to P 7789a (figures 1, A1). Anterior is to the right. Scalebar represents 10 mm. 2

3 Electronic Appendix B. Phylogenetic Analysis. The cladogram presented in figure 2 is based on the analysis of an updated version of the matrix presented in Forey (1998), which consists of 109 morphological characters (87 cranial, 22 postcranial) scored for 25 ingroup taxa. Actinopterygians and porolepiforms were set as outgroups. Following Forey (1998), we have not included the following genera, which are highly unstable owing to considerable amounts of missing data: Axelia, Euporosteus, Indocoelacanthus, Lualabaea, Ticinepomis and Wimania. A strict consensus of the 54 shortest cladograms recovered when all characters are weighted equally differs from the topology presented in figure 2 in placing Hadronector, Lochmocercus, and Polyosteorhynchus in a polytomy with the clades [Holopterygius + Allenypterus] and [Caridosuctor + all more crownward coelacanths]. A majority-rule consensus of these cladograms places Hadronector, [Holopterygius + Allenypterus], Lochmocercus, and Polyosteorhynchus as successively more crownward plesions along the coelacanth stem above Diplocercides. All solutions recover the sister-group relationship between Holopterygius and Allenypterus. Character list. Our only modification to the character list presented by Forey (1998) is the inclusion of an additional character (character 109). For full character descriptions, consult Forey (1998) Margin of dermal intracranial joint: straight (0) deeply notched (1). Snout bones: free from one another (0), fused (1). Median rostral: single median rostral or internasal (0), several median rostrals or internasals (1). Premaxillae: paired (0), fragmented (1). Dorsal lamina of premaxilla: present (0), absent (1). Anterior opening of rostral organ: within premaxilla (0), within separate rostral ossicle (1). Parietals: one pair (0), two pairs (1). Anterior and posterior pairs of parietals: similar size (0), dissimilar size (1). Number of supraorbitals/tectals: fewer than 8 (0), greater than 10 (1). Preorbital: absent (0), present (1). Descending process of parietal: absent (0), present (1). Intertemporal: absent (0), present (1). Postparietal descending process: absent (0), present (1). Supratemporal descending process: absent (0), present (1). Extrascapulars: sutured with postparietals (0), free (1). Extrascapulars: behind skull roof (0), part of roof (1). Number of extrascapulars: three (0), five (1), more than seven (2). Posterior margin of the skull roof: straight (0), embayed (1). Supraorbital sensory canal: passes through ossification centres (0), follows a sutural course (1). Medial branch of otic canal: absent (0), present (1). Otic canal joining supratemporal canal: in lateral extrascapular (0), in supratemporal (1) 3

4 Anterior branches of supratemporal commissure: absent (0), present (1). Supraorbital sensory canals opening through bones as: single large pores (0), bifurcating pores (1), many tiny pores (2). Anterior pit line: absent (0), present (1). Middle and posterior pit lines: within posterior half of postparietals (0), within anterior third (1). Pit lines: marking postparietals (1), not marking (0). Parietals and postparietals ornamented with: enamel-capped ridges/tubercles (0), unornamented (1), marked by coarse rugosities (2). Parietals and postparietals: without raised areas (0), with raised areas (1). Cheek bones: in contact/overlappping (0), separated (1). Spiracular (postspiracular): absent (0), present (1). Preoperculum: absent (0), present (1). Suboperculum: absent (0), present (1). Quadratojugal: absent (0), present (1). Squamosal: reaching skull roof (0), not reaching roof (1). Lachrymojugal: not expanded anteriorly (0), expanded (1). Lachrymojugal: ending without anterior angle (0), angled anteriorly (1). Squamosal: large (0), reduced to narrow tube (1). Preoperculum: large (0), reduced to narrow tube (1). Peroperculum: undifferentiated (0), developed as a posterior tube-like canalbearing portion and an anterior blade-like portion (1). Anterodorsal excavation in postorbital: absent (0), present (1). Postorbital: without anterior process (0), with process (1). Postorbital: plate-like (0), reduced to narrow tube (1). Postorbital: lying wholly behind intracranial joint (0), spanning joint (1). Infraorbital canal: within postorbital, with simple pores opening directly from main canal (0), anterior and posterior branches within the postorbital (1). Infraorbital canal: running through centre of postorbital (0), running along anterior margin of the postorbital (1). Jugal sensory canal: simple (0), with prominent branches (1). Jugal canal: running through centre of bone (0), running along ventral margin of squamosal (1). Pit lines: marking bones (0), failing to mark bones (1). Ornament on cheek bones: absent (0), tubercular (1), rugose (2). Openings for infraorbital, jugal and preopercular sensory canals: many small pores (0), few large pores (1) Lachrymojugal: sutured to preorbital and lateral rostral (0), lying in sutural contact with the tectal-supraorbital series (1). Sclerotic ossicles: absent (0), present (1). Retroarticular and articular: co-ossified (0), separated (1). Dentary teeth: fused to dentary (0), on separate tooth plates (0). Number of coronoids at anterior end of jaw: zero (0), one (1), two (2), three (3), four (4). Coronoid opposite posterior end of dentary: not modified (0), modified (1). Dentary: simple (0), hook-shaped (1). Oral pit line: short, confined to angular (0), long, reaching forward to dentary 4

5 and/or splenial (1). Oral pit line: located at centre of ossification of angular (0), removed from the centre of ossification (1). Subopercular branch of mandibular sensory canal: absent (0), present (1). Dentary sensory pore: absent (0), present (1). Ornament of lower jaw: ridged (0), tubercular (1). Dentary: with ornament (0), without ornament (1). Splenial: with ornament (0), without ornament (1). Dentary: without prominent lateral swelling (0), with swelling (1). Principal coronoid: lying free (0), sutured to angular (1). Cornoid fangs: absent (0), present (1). Prearticular and/or coronoid teeth: pointed and smooth (0), rounded and marked with fine striations (1). Orbitosphenoid and basisphenoid regions: co-ossified (0), separate (1). Basisphenoid: extending forward to enclose optic foramen (0), optic foramen lying within separate interorbital ossification or cartilage (1). Processus connectens: meeting parasphenoid (1), failing to meet (1). Basipterygoid process: absent (0), present (1). Antotic process: not covered by parietal descending process (0), covered (1) Temporal excavation: lined with bone (0), not lined (1). Otico-occipital: solid (0), separated to prootic/opisthtoic (1). Supraoccipital: absent (0), present (1). Vestibular fontanelle: absent (0), present (1). Buccohypophysial canal: opening through parasphenoid (0), closed (1). Parasphenoid: without ascending laminae anteriorly (0), with ascending laminae (1). Suprapterygoid process: absent (0), present (1). Vomers: not meeting in midline (0), meeting medially (1). Prootic: without complex suture with basioccipital region (0), with suture (1). Superficial opthalmic branch of anterodorsal lateral line nerve: not piercing antotic process (0), piercing antotic process (1). Process on braincase for articulation of infrabranchial 1: absent (0), present (1). Separate lateral ethmoids: absent (0), present (1). Separate basioccipital: absent (0), present (1). Dorsum sellae: small (0), large and constricting entrance to cranial cavity anterior to intracranial joint (1). Extracleithrum: absent (0), present (1). Anocleithrum: simple (0), forked (1). Posterior neural and haemal spines: abutting one another (0), not abutting (1). Occipital neural arches: not expanded (0), expanded (1). Ossified ribs: absent (0), present (1). Diphycercal fin: absent (0), present (1). Caudal fin rays: more numerous than radials (0), equal in number (1). Fin rays: branched (0), unbranched (1). Fin rays in D1: greater than ten (0), eight or nine (1), less than eight (2). Caudal lobes: symmetrical (0), asymmetrical (1). Fin rays in D1: without denticles (0), with denticles (1). 5

6 Paired fin rays: not expanded (0), expanded (1). Pelvics: abdominal (0), thoracic (1). Basal plate of D1: with smooth ventral margin (0), emarginate and accommodating the tips of adjacent neural spines (1). Basal support of D2: simple (0), forked anteriorly (1). Median fin rays: not expanded (0), expanded (1). Scale ornament: not differentiated (0), differentiated (1). Lateral line openings in single scales: single (0), multiple (1). Scale ornament: enamel ridges or tubercles (0), rugose bone only (1). Swimbladder: not ossified (0), ossified (1). Pelvic bones on each side: separate from one another (0), fused over their entire length (1). Ventral keel scales: absent (0), present (1). Taxon-by-character matrix. Additions and modifications of the matrix in Forey (1998) are highlighted by bold, underlined text. Holopterygius and character are both new additions. Character 31 was miscoded in Forey (1998), but the author has provided us with corrected scores (P. L. Forey, pers. comm.). The mandible of Allenypterus is edentulous, and we argue that the state of the dentary teeth (character 54) of this taxon cannot logically be assessed. We have therefore scored this character as?. Allenypterus 0?????2001?0?? ? ?? ??? ????????????????????? Axelrodichthys ? 1020? ??0 1? ?0?? 110?1111? ?1100 Caridosuctor ?0?? ?? ?? ????????????????????101? ?0100 Chinlea 00?01? ??0011?1 10???? ? ?0???20 10???11??? 1?11001????????????????????1?1?11111?000?101?1?00 Coccoderma 10?011210? ? ? ???????????????????? Coelacanthus 00?11? ?1 10?0?1101???00001??0 010?1?1? ??0 1?110000??????????????????? ?0100 Diplocercides 001??? ?010000?? ??0000??? Diplurus 00001? ? ?? ? ?11?00???0? ?0000 6

7 Garnbergia??????21?0?0?????1??????????1? ?0 000?????1? 10???????????????????????????????????????????1?0?? 0100?0??0 Hadronector ?1?0?? ?? ?1?0?10 010??? ?????????????????????101? ??0100 Holophagus 10?01? ?111 1?20? ?1?? ?? ???????11?????1??1??1? ?0100 Latimeria ? Laugia 11?0??1? ? ?? ?0110 Libys 0???1????? 10111??111 1?00? ?1???01?1? ?110001??????????????????? Lochmocercus??????????????????10 100??? ?? ?01000?0 01?0?00??0 00??000????????????????????101?? ??0?00 Macropoma 01?0? ? ? ? ?01? 110? Mawsonia 1????? ?1 1020?12010? ? ??1??0 1?111101?????????????????????1??111? ?1?0? Miguashaia 00?0001?0? 01?? ?00000? 1101?? ?000?10?1?0??001?? 0??0000???????????????????1?0??0000?000??0000??0 Polyosteorhynchus 00? ???100? ??? ? ?1? ? ??000????????????????????1? ? ?0000 Rhabdoderma ?0000? Sassenia 10????2? ?01?? ?1?10?? ?? ?01? ???????????????0?0??0 Spermatodus ?111?020?1001? ????? ?000?1??? ??0110??00?101????????????????0?0??? Undina 00?00? ? 1?2??? ?? ? ??1101? ?01??1??111101?

8 Whiteia 00? ? ??00? ? Porolepiformes 00100?1?? ? ?? ?0 0? ?000? ?0? ? Actinopterygii?0000?1?? ? ? ???00????????010? ? ??1?? ? ?000 0?000000{0/1} Holopterygius?0?0??????????????????0?????????????????????????????00??0????????? 00?????????00????????10? 00101?1????00???0?1 Electronic Appendix C. Morphometric Analysis. Raw landmark coordinates. The source of the reconstruction or specimen photograph from which landmarks were digitized are given in parentheses after each genus. For a listing of landmarks used in this study, consult figure 2 of the primary text. Landmarks were placed on the reconstruction of Holopterygius presented in figure 1 of the primary text. Although the position of the dorsal, anal, and pelvic fins in this taxon are incompletely known, those incorporated into the reconstruction are conservative estimates based on the conditions in its closest relative, Allenypterus. Allenypterus (Forey 1998) (58, 232), (166, 504), (439, 685), (615, 699), (745, 707), (784, 701), (1900, 257), (1688, 239), (983, 143). (685, 111). (268, 188) Axelrodichthys (Forey 1998) (11, 173), (231, 302), (403, 391), (508, 393), (758, 380), (944, 377), (1361, 206), (987, 102), (818, 144), (568, 117), (206, 123) Caridosuctor (Lund & Lund 1985) (53, 516), (343, 591), (696, 610), (874, 603), (1078, 569), (1359, 574), (1957, 473), (1414, 348), (1116, 343), (835, 285), (320, 405) Changxingia (Wang & Liu 1981) (248, 382), (430, 500), (641, 539), (755, 538), (950, 539), (1184, 562), (1505, 439), (1183, 313), (1002, 301), (758, 256), (386, 309) Coccoderma (Lambers 1992) (74, 301), (255, 385), (450, 440), (498, 434), (764, 416), (964, 429), (1241, 329), (990, 203), (762, 192), (381, 209), (237, 240) Coelacanthus (Forey 1998) (79, 192), (303, 301), (506, 318), (592, 304), (870, 287), (1063, 284), (1405, 195), (1058, 135), (923, 127), (637, 114), (243, 128) Diplocercides (Jessen 1973) (81, 410), (452, 561), (694, 575), (796, 565), (1114, 544), (1378, 538), (1865, 392), (1350, 283), (1107, 272), (803, 274), (342, 314) 8

9 Diplurus (Forey 1998) (64, 149), (216, 274), (371, 334), (417, 314), (632, 319), (771, 322), (1139, 274), (807, 167), (710, 149), (473, 123), (180, 139) Hadronector (Lund & Lund 1985) (135, 559), (440, 774), (769, 862), (895, 814), (1161, 755), (1337, 750), (1956, 550), (1318, 334), (1120, 329), (884, 283), (390, 422) Holophagus (Forey 1998) (9, 179), (180, 297), (397, 369), (481, 354), (642, 326), (837, 326), (1087, 240), (855, 146), (737, 141), (494, 95), (153, 104) Holopterygius (present study) (18, 135), (131, 226), (178, 240), (253, 256), (306, 255), (350, 256), (921, 197), (552, 89), (492, 84), (330, 69), (131, 101) Latimeria (Forey 1998) (60, 297), (296, 393), (548, 444), (641, 437), (898, 415), (1132, 460), (1444, 369), (1169, 194), (960, 200), (669, 118), (223, 182) Laugia (Forey 1998) (10, 137), (151, 216), (325, 254), (380, 248), (625, 240), (740, 240), (1088, 164), (798, 93), (612, 82), (298, 67), (131, 93) Libys (Lambers 1992) (34, 322), (176, 446), (373, 559), (443, 529), (640, 504), (732, 501), (992, 389), (780, 232), (659, 216), (408, 196), (187, 230) Lochmocercus (Lund & Lund 1985) (55, 190), (275, 369), (569, 439), (628, 438), (838, 428), (1003, 438), (1310, 343), (1030, 215), (801, 163), (566, 128), (263, 137) Macropoma (Forey 1998) (52, 243), (260, 339), (501, 369), (548, 360), (727, 336), (934, 360), (1164, 250), (938, 130), (788, 150), (556, 112),(250, 163) Macropomoides (Forey 1998) (37, 188), (218, 348), (364, 394), (431, 383), (658, 372), (819, 377), (1140, 241), (835, 119), (720, 122), (474, 99), (183, 113) Miguashaia (Forey 1998) (33, 176), (174, 309), (418, 343), (537, 340), (828, 319), (941, 333), (1125, 379), (872, 165), (813, 157), (624, 121), (198, 110) Polyosteorhynchus (Lund & Lund 1985) (75, 255), (362, 453), (679, 514), (785, 497), (937, 436), (1189, 466), (1549, 377), (1260, 226), (991, 209), (668, 131), (354, 140) Rhabdoderma Forey (1998) (41, 204), (196, 259), (356, 296), (412, 286), (569, 275), (748, 265), (1024, 209), (766, 140), (598, 122), (427, 110), (181, 122) Ticinepomis (Rieppel 1980) (75, 254), (314, 350), (505, 397), (554, 374), (830, 364), (1020, 373), (1527, 260), (1031, 145), (880, 156), (576, 119), (220, 202) Whitea (Forey 1998) (17, 189), (219, 287), (379, 315), (459, 305), (609, 281), (798, 288), (1018, 195), (815, 124), (672, 120), (451, 95), (186, 106) 9

10 Relative warps scores. Only the first four relative warp axes (highlighted in boldface) account for more than 5 percent of total variance within the sample of coelacacanths examined in this study, and estimates of morphological disparity presented in the primary text are derived from values taken from these axes. RW Percentage of total variance 48.87% 18.47% 12.02% 6.96% 4.89% 2.69% 2.01% 1.38% 0.75% 0.54% 0.49% 0.34% 0.24% 0.17% 0.10% 0.05% 0.00% 0.00% Cumulative variance 48.87% 67.34% 79.37% 86.32% 91.21% 93.90% 95.92% 97.30% 98.05% 98.59% 99.08% 99.43% 99.67% 99.84% 99.94% 99.99% % % Allenypterus E-001, E-001, E-002, E-002, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-004, E-003, E-003, E-003, E-004, E-005 Axelrodichthys E-002, E-002, E-003, E-003, E-002, E-002, E-002, E-003, E-002, E-003, E-002, E-003, E-002, E-003, E-003, E-003, E-004, E-004 Caridosuctor E-002, E-002, E-003, E-002, E-002, E-002, E-003, E-002, E-003, E-002, E-002, E-003, E-004, E-003, E-003, E-003, E-004, E

11 Changxingia E-002, E-004, E-003, E-003, E-003, E-003, E-003, E-002, E-003, E-003, E-003, E-002, E-002, E-004, E-005, E-003, E-003, E-004 Coccoderma E-002, E-003, E-001, E-002, E-002, E-003, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-003, E-003, E-005, E-003 Coelacanthus E-002, E-002, E-002, E-002, E-002, E-003, E-002, E-002, E-003, E-003, E-003, E-003, E-004, E-003, E-003, E-003, E-003, E-003 Diplocercides E-002, E-002, E-003, E-003, E-002, E-002, E-002, E-003, E-002, E-002, E-003, E-004, E-003, E-003, E-003, E-005, E-003, E-003 Diplurus E-003, E-002, E-003, E-002, E-002, E-004, E-002, E-002, E-002, E-003, E-004, E-002, E-002, E-003, E-003, E-003, E-004, E-004 Hadronector E-002, E-002, E-001, E-002, E-002, E-002, E-002, E-002, E-004, E-003, E-003, E-003, E-003, E-003, E-004, E-003, E-004, E

12 Holophagus E-002, E-002, E-002, E-002, E-002, E-002, E-002, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-003, E-005, E-004 Holopterygius E-001, E-001, E-002, E-002, E-002, E-003, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-003, E-004, E-003, E-005, E-004 Latimeria E-002, E-002, E-004, E-002, E-002, E-003, E-002, E-002, E-002, E-002, E-003, E-004, E-002, E-003, E-003, E-003, E-004, E-004 Laugia E-002, E-002, E-001, E-002, E-003, E-002, E-003, E-002, E-003, E-003, E-004, E-003, E-003, E-003, E-002, E-004, E-004, E-004 Libys E-003, E-001, E-002, E-002, E-002, E-002, E-002, E-003, E-005, E-002, E-003, E-003, E-003, E-002, E-004, E-003, E-004, E-004 Lochmocercus E-002, E-002, E-002, E-002, E-002, E-003, E-002, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-003, E-003, E

13 Macropoma E-002, E-002, E-003, E-002, E-003, E-002, E-002, E-002, E-005, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-004, E-004 Macropomoides E-002, E-002, E-002, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-002, E-003, E-003, E-003, E-003, E-004, E-004, E-004 Miguashaia E-001, E-004, E-002, E-002, E-002, E-002, E-003, E-002, E-002, E-003, E-003, E-004, E-003, E-003, E-003, E-003, E-005, E-004 Polyosteorhynchus E-002, E-002, E-002, E-003, E-002, E-002, E-002, E-002, E-004, E-004, E-003, E-003, E-003, E-002, E-003, E-003, E-003, E-004 Rhabdoderma E-002, E-002, E-002, E-002, E-002, E-003, E-003, E-002, E-002, E-002, E-002, E-002, E-004, E-003, E-003, E-003, E-004, E-005 Ticinepomis E-002, E-002, E-003, E-002, E-003, E-004, E-003, E-002, E-002, E-003, E-004, E-003, E-003, E-003, E-004, E-004, E-004, E

14 Whitea E-002, E-002, E-002, E-002, E-003, E-002, E-002, E-002, E-002, E-003, E-003, E-003, E-003, E-003, E-003, E-003, E-003, E-004 Electronic Appendix D. References. Forey, P. L History of the coelacanth fishes. London: Chapman & Hall. Jessen, H Weitere fischreste aus dem Oberen Plattenkalk der Bergisch-Gladbach Paffrather mulde (Oberdevon, Rheinesches Schiefergebirge). Palaeontogr. Abt. A. Palaeozool.-stratigr. 143, Lambers, P On the ichthyofauna of the Solnhofen Limestone (Upper Jurassic, Germany). Unpublished PhD Thesis, University of Groningen. Lund, R. & Lund, W. L Coelacanths from the Bear Gulch Limestone (Namurian) of Montana and the evolution of the Coelacanthiformes. Bull. Carnegie Mus. Nat. Hist. 25, Rieppel, O A new coelacanth from the Middle Triassic of Monte San Giorgio, Switzerland. Eclogae Geol. Helv. 73, Swofford, D. L PAUP*. Phylogenetic analysis using parsimony (*and other methods), version 4. Sunderland, Massachusetts: Sinauer Associates. Wang, N. & Liu, H Coelacanth fishes from the marine Permian of Zhejiang, South China. Vertebr. PalAsiat. 19,

Exceptional fossil preservation demonstrates a new mode of axial skeleton elongation in early ray-finned fishes

Supplementary Information Exceptional fossil preservation demonstrates a new mode of axial skeleton elongation in early ray-finned fishes Erin E. Maxwell, Heinz Furrer, Marcelo R. Sánchez-Villagra Supplementary