100-Million-Year Dynasty of Giant Planktivorous Bony Fishes in the Mesozoic Seas

Transcription

1 Supporting Online Material for 100-Million-Year Dynasty of Giant Planktivorous Bony Fishes in the Mesozoic Seas Matt Friedman,* Kenshu Shimada, Larry D. Martin, Michael J. Everhart, Jeff Liston, Anthony Maltese, Michael Triebold *To whom correspondence should be addressed. This PDF file includes: Published 19 February 2010, Science 327, 990 (2010) DOI: /science SOM Text Figs. S1 to S15 References

2 SUPPORTING ONLINE MATERIAL (SCIENCE ONLINE) FOR 100-Million-Year Dynasty of Giant Planktivorous Bony Fishes in the Mesozoic Seas Image Robert Nicholls, PalaeoCreations. Matt Friedman 1, Kenshu Shimada 2,3, Larry D. Martin 4, Michael J. Everhart 3, Jeff Liston 5, Anthony Maltese 6, Michael Triebold 6 1 Department of Earth Sciences, University of Oxford, Parks Road, Oxford OX1 3PR, UK. 2 Environmental Science Program and Department of Biological Sciences, DePaul University, 2325 North Clifton Avenue, Chicago, Illinois 60614, USA. 3 Sternberg Museum of Natural History, Fort Hays State University, 3000 Sternberg Drive, Hays, Kansas 67601, USA. 4 Natural History Museum and Biodiversity Research Center, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA. 5 Division of Ecology and Evolutionary Biology, Faculty of Biomedical and Life Sciences, University of Glasgow, University Avenue, Glasgow, Scotland G12 8QQ, UK. 6 Rocky Mountain Dinosaur Resource Center, 201 South Fairview Street, Woodland Park, Colorado 80863, USA. 1

3 TABLE OF CONTENTS 1. Institutional abbreviations Supplementary anatomical data: Oolite pachycormid 4 3. Supplementary anatomical data: Rhinconichthys gen. nov Taxonomic history of Bonnerichthys gen. nov Supplementary anatomical data: Bonnerichthys gen. nov Paleobiogeographic distribution of suspension-feeding pachycormids Characters used in cladistic analysis Comparative materials Taxon-by-character matrix Systematic methodology Results of cladistic analyses Supplementary References

4 1. INSTITUTIONAL ABBREVIATIONS AMNH: American Museum of Natural History, New York, New York, USA (FF denotes fossil fish collection in the Department of Vertebrate Paleontology). BGS: British Geological Survey, Keyworth, Nottingham, UK. BSP: Bayerische Staatsammlung für Paläontologie un Historische Geologie, München, Germany. BMNH: The Natural History Museum, London, UK. FHSM: Sternberg Museum of Natural History, Fort Hays State University, Hays, Kansas, USA. FMNH: The Field Museum, Chicago, Illinois, USA. KUVP: University of Kansas Museum of Natural History, Lawrence, Kansas, USA. LACM: Natural History Museum of Los Angeles County, Los Angeles, California, USA. NMS: National Museum of Science and Nature, Tokyo, Japan. SMGM: Staatliches Museum für Geologie und Mineralogie, Dresden, Germany. UNSM: University of Nebraska State Museum, Lincoln, Nebraska, USA. 3

5 2. SUPPLEMENTARY ANATOMICAL DATA: OOLITE PACHYCORMID Fig. S1. Incomplete skull of generically indeterminate pachycormid, BMNH P.41669, lower Middle Jurassic (Bajocian; Ma) Inferior Oolite of Dorset, UK, shown in left-lateral view. Probable stratigraphic level for this specimen ( Rubbly Beds, top of Upper Inferior Oolite above Sherborne Building Stone) was determined by comparison of matrix with lithological collections at the British Geological Survey and Museum (sample MR 7712; letter to Dr. E. I. White dated 29 June 1959 archived in BMNH collections with fossil specimen). Abbreviations: ang, angular; de, dentary; fr, frontal; gu, gular plate;?mx,?maxilla. A badly eroded maxilla is present on the right side of the skull, and lies in life position dorsolateral to the lower jaw. If our tentative identification of the left maxilla is correct, this bone has been subjected to post-mortem displacement (see Figs S3-S4 and Fig. 2 in the main text for maxillae preserved in life position). 4

6 3. SUPPLEMENTARY ANATOMICAL DATA: RHINCONICHTHYS GEN. NOV. Fig. S2. Skull of Rhinconichthys taylori gen. et sp. nov., holotype, BMNH 219, from the Upper Cretaceous (Cenomanian) Lower Chalk, Burnham, Kent, UK. (A) Dorsal view. (B) Ventral view. Abbreviations: art, articular; chy, anterior ceratohyal; de, dentary; dpt, dermopterotic; ent, entopterygoid; fr, frontal; gu, gular plate; hhy, hypohyal; hym, hyomandibula; mx, maxilla; pa, parietal; part, prearticular; psp, parasphenoid; pto, pterotic; scl, sclerotic ring. Paired bones listed as right (r) or left (l). 5

7 Figure S3. Rhinconichthys taylori gen. et sp. nov., holotype, BMNH 219, from the Upper Cretaceous (Cenomanian) Lower Chalk, Burnham, Kent, UK. (A) Left lateral view of cheek region, showing details of palate. (B) Close-up of (A), showing gill rakers and associated needle-like projections. Abbreviations: ect, ectopterygoid; ent, entopterygoid; mtp, metapterygoid; rak, gill rakers. 6

8 Fig. S4. Skull of Rhinconichthys cf. R. taylori from the Upper Cretaceous (Cenomanian) of Mikasa City, Hokkaido, Japan. Photographs of BMNH P.66822, cast of specimen NSM VP21868 shown in (A) right-lateral, (B) left-lateral, (C) dorsal, and (D) ventral views. Abbreviations: ang, angular; boc, basioccipital; chy, ceratohyal; de, dentary; dpt, dermopterotic; fr, frontal; hym, hyomandibula; mtp, metapterygoid; mx, maxilla; pa, parietal; scl, sclerotic ring. Paired bones listed as right (r) or left (l). 7

9 4. TAXONOMIC HISTORY OF BONNERICHTHYS GEN. NOV Portheus gladius Cope; p Portheus gladius Cope; Cope, p Pelecopterus gladius (Cope); Cope, p. 244, pl. xliv, fig. 12; pl. lii, fig Pelecopterus gladius (Cope); Woodward, p Protosphyraena gigas Stewart; p Protosphyraena gladius (Cope); Loomis, p. 221, fig Protosphyraena gigas Stewart; Loomis, p Protosphyraena gigas Stewart; Stewart, p. 367, pl Protosphyraena gladius (Cope); Stewart, p Protosphyraena gladius (Cope); Hay, p Protosphyraena gladius (Cope); Hay, p Protosphyraena sp. 3; Applegate; p Protosphyraena gladius (Cope); Schultze et al., p Protosphyraena gladius (Cope); Stewart, p Protosphyraena gladius (Cope); Russell, p Protosphyraena gladius (Cope); Stewart, p Protosphyraena gladius (Cope); Schwimmer et al., table Protosphyraena gladius (Cope); Stahl, p Protosphyraena gladius (Cope); Carpenter, table Protosphyraena gladius (Cope); Carpenter, table Protosphyraena gladius (Cope); Kiernan, p Protosphyraena gladius (Cope); Everhart, p Protosphyraena gladius (Cope); Shimada and Fielitz, p Protosphyraena gladius (Cope); Parris et al., p Protosphyraena gladius (Cope); Cicimurri et al., p Protosphyraena gladius (Cope); Fielitz and Shimada, table 1 8

10 Fig. S5. Type specimen of Bonnerichthys gladius gen. nov., AMNH FF (A) Specimen photograph. (B) Lithograph of the same specimen from Cope (1875). Pieces of fin visible in (B) but not shown in (A) are still conserved in AMNH collections, but are no longer attached to the body of the specimen. The type specimen of Bonnerichthys gladius (Fig. S5) was first described by E. D. Cope (1873) as a species of Portheus, a junior synonym of Xiphactinus Leidy (1870). In 1875, Cope erected the new genus Pelecopterus for his Portheus gladius and two additional species, chiurgus and perniciosus, the latter of which he had previously placed in Ichthyodectes (Cope 1874). Cope (1875) did not specify a type species for Pelecopterus, which he placed in the new family Pelecopteridae within the new order Actinochiri. A. S. Woodward (1895) was the first reviser of Pelecopterus, designating perniciosus as the type species of this genus. This assignment is valid under article 69.1 of the International Code of Zoological Nomenclature (ICZN 1999). Woodward considered Pelecopterus a junior synonym of Protosphyraena Leidy (1860), and we agree with his assessment. Protosphyraena perniciosa (gender of species name changed to match that of the genus) shares with other species of Protosphyraena a greatly elongated rostrodermethmoid and a suite of additional derived characters (see summaries in Mainwaring 1978 and Lambers 1992), and with the type species P. ferox distinctively crenellated anterior margins of the pectoral fin. However, our analyses (Section 11) clearly show that P. gladius lacks derived characters of Protosphyraena, and represents a distinct genus related to edentulous pachycormids 9

11 ( Asthenocormus, Leedsichthys, Martillichthys, and Rhinconichthys) but differing from known taxa in several notable features (e.g., deep groove for dorsal aorta in basioccipital, lack of sutural contacts between bones of the dermal skull, and a distinctive band of fused rays along the anterior margins of pectoral fins). It has been argued (Schultze et al. 1982; Stewart 1988) that there is an available generic designation for this species: Ichthypriapus Hibbard (1942). We do not believe that this is the case. First, we regard the association between the type material of Ichthypriapus and remains of Protosphyraena gladius as tenuous. Second, we have not observed elements similar to the type of Ichthypriapus among the material of two nearly complete skulls of P. gladius (KUVP 60692; FHSM VP-17428), or in the many pachycormid remains examined in the course of this study (Section 8). Third, and most significantly, we note many distinctive structural features of Ichthypriapus support the original systematic and anatomical interpretation of this specimen (Hibbard 1942). Ichthypriapus hubbsi was erected in 1942 by C. Hibbard for an unusually shaped fossil from the Smoky Hill Member of the Niobrara Formation which he identified as the cranial clasper of an exceptionally large holocephalan chondrichthyan (KUVP 1136; Fig. S6). This specimen derives from a slab also yielding a set of pectoral fins and girdles belonging to Protosphyraena gladius (KUVP 465; Fig. S8). The exact connection between these two sets of remains is not well documented. The only available details come from Hibbard s description of Ichthypriapus, where he notes that the type and only specimen was recovered by Mr. H. T. Martin while cleaning up part of a Protosphyraena skeleton that had been collected in the Niobrara Chalk. The specimen under discussion was not associated with the Protosphyraena specimen, only inasmuch as it was in the same block of chalk. (1942: 237) Despite the lack of evidence unambiguously tying Ichthypriapus to the pachycormid, Schultze et al. (1982) regarded KUVP 1136 as part of P. gladius. These authors did not offer any specific identity for the element, but listed it as an unknown bone (Schultze et al. 1982: 23). Their conclusion that Ichthypriapus belonged to P. gladius appears to stem from two lines of evidence: (i) the apparent spatial association 10

12 Fig. S6. Type specimen of Ichthypriapus hubbsi, KUVP Broken process is reconstructed here in grey. between KUVP 1136 and KUVP 465, regarded by both Hibbard (1942) and us as insufficient evidence to claim derivation from the same animal; and (ii) their interpretation of the hard tissue of Ichthypriapus as bone (holocephalan cranial claspers are composed of calcified cartilage; Stahl 1999). We are doubtful of the latter claim, as Schultze et al. (1982) provided no histological sections of this unique specimen. It should be possible to offer a positive identification for the Ichthypriapus element if it indeed belongs to Protosphyraena gladius, especially given the wealth of new information available for this species. However, we have found that KUVP 1136 shows no clear correspondence to bones represented in other material of P. gladius, or to structures known in other pachycormids. The type of Ichthypriapus is sigmoidal in shape, with a surface covered with fine, parallel striae. One end of the specimen is bifurcate, while the other terminates in a blunt, roughened knob. KUVP 1136 bears a pair of stout processes with rough surfaces near mid-length, and an elongated pit on one surface that appears confluent with a closed channel that extends through the element and divides to exit through separate foramina positioned at the distal tips of the bifurcate end of the specimen. This enclosed canal is clearly shown in cross-section where one of the 11

13 two processes contributing to the bifurcation has been broken. The canal is infilled with what appears to be recrystallized calcite, and is surrounded by a thick biomineralized rind that makes up the body of the fossil. Midline bones of the skull that are currently unknown in material of Protosphyraena gladius but predicted to be present based on conditions in other pachycormids are confined to the basibranchial series of the gill skeleton. If KUVP 1136 is attributable to P. gladius, then we would predict similar structures in other pachycormids. However, we have not observed a bone with comparable morphology in any pachycormid, including acid-prepared skulls of the Jurassic genus Pachycormus with complete branchial skeletons (described in Mainwaring 1978 and deposited at BMNH; see Section 8). Basibranchials in this genus are variously cylinder or hour-glass shaped, and bear a smooth coat of perichondral bone. The basibranchial series in other pachycormids is incompletely known, but accounts of these ossifications in Hypsocormus (Lehman 1949) and Leedsichthys (Liston 2008) conform to conditions in Pachycormus. The complex sigmoidal shape and distinctive surface texture of KUVP 1136 are inconsistent with these patterns. Among actinopterygians, the only other midline ossification commonly represented in gill skeletons is the urohyal, a membrane bone that develops as an ossification of the tendon of the sternohyoideus muscle (Arratia and Schultze 1990). A urohyal is present in the extant cladistians (bichir and ropefish) and teleosts, but is absent in chondrosteans (sturgeon and paddlefish), lepisosteids (gars) and amiids (bowfin). Urohyals are not known in distal members of the teleost stem, including pachycormids, and are considered primitively absent in those taxa (the plate-like bone identified by Felix [1890: pl. 12] as a urohyal is a median gular). Instead, a single tendon-bone urohyal appears to be a derived character linking pholidophorids and all more crownward members of the teleost total group (Arratia and Schultze 1990). This ossification has been reported in a series of successively more crownward Jurassic members of the teleost stem: Pholidophorus germanicus and macrocephalus (Patterson 1977b: fig. 5), Leptolepis coryphaenoides (Arratia and Schultze 1990: fig. 10A), Ascalabos viothii (Arratia 1997: fig. 16). In all of these taxa, the urohyal has a long, slender plate-like body with a bifurcate anterior tip marking the areas for ligamentous attachments with the 12

14 hypohyals. The anatomy of the urohyal is much the same in coeval crown teleosts like Leptolepides sprattiformis (Arratia 1997: fig. 42A) and Orthogonikleithrus hoelii (Arratia 1997: fig. 57A). This bifurcate tip is the only plausible similarity between these examples of plesiomorphic teleost urohyals and the holotype of Ichthypriapus, but the distal surfaces of these paired processes do not bear attachment areas for ligaments (Hibbard 1942) of the kind expected in a urohyal (Arratia and Schultze 1990). Morphological dissimilarity, coupled with the absence of urohyals in other pachycormids and the lack of additional bones of the gill skeleton in P. gladius specimen KUVP 465, leads us to reject the possible interpretation of this structure as a urohyal. Minimal comparative evidence was enlisted to support the original identification of Ichthypriapus as a holocephalan cranial clasper. However, we find several peculiar aspects of this fossil consonant with the clasper hypothesis. These include overall shape, surface texture, internal structure, and the geometry of features that we interpret as muscle attachment scars. Regarding shape, Hibbard (1942) made limited comparisons to Chimaera and Ischyodus, both of which have arched claspers that differ from the sigmoidal profile of Ichthypriapus. However, a great diversity of clasper morphologies are represented in fossil holocephalans (Stahl 1999), including a large S -shaped structure in the myriacanthid Halonodon that approaches the size of KUVP 1136 (Duffin 1984). Whereas the surface texture and cross-section of the hard tissue of Ichthypriapus differs from that observed for skeletal remains of Protosphyraena gladius, it does agree closely with the highly mineralized fibrocartilage found in holocephalan cranial claspers. This tissue is very compact (Reis 1895: fig. 5), its external surface bears numerous parallel striae (Reis 1895: figs 4, 11; Duffin 1984), and it forms a thick rind about the hollow core of the structure (Reis 1895: figs 5, 12). One of us (MF) has confirmed these structural similarities through personal examination of cranial claspers of the Jurassic holocephalan Myriacanthus granulatus (BGS GSM ). Additionally, interpretation of KUVP 1136 as a cranial clasper helps to account for two unusual features of this fossil: the large excavation flanked by shallow fossae apparent on one face of the specimen, and the paired protuberances located at mid-length of the fossil. The elongated excavation corresponds in position to a median chamber for blood vessels 13

15 (Chimaera, Reis 1895: fig. 12), whereas the fossae lateral to it mark where the antorbital ligament inserts on to the cranial clasper (Chimaera, Reis 1895: fig. 8; Hydrolagus, Raikow and Swierczewski 1975: fig. 1). These ligaments bind the clasper to the head, forming an axis of rotation for its depression and elevation relative to the neurocranium (Raikow and Swierczewski 1975: 436). In many fossil holocephalans, this region of the clasper is marked by pronounced pits separated by a low ridge (Stahl 1999), and we note a midline elevation extending from one end of the elongated depression to the blunt end of KUVP The stout lateral processes with a cancellous surface texture correspond to the insertion area of a tendon issuing from the preorbitalis muscle (Reis 1895: fig. 8; Raikow and Swierczewski 1975: fig. 1). Based on these observations, we agree with Hibbard s (1942) assertion that KUVP 1136 is distinct from the associated pachycormid skeleton KUVP 465, and conclude that it was correctly identified as a cranial clasper. A large holocephalan (estimated total length ca. 2.8 m), Eumylodus laqueatus, is known from the Smoky Hill Member of the Niobrara Formation (Cicimurri et al. 2008). This species is based on dentitions, but it is possible that both a large dorsal-fin spine (BMNH P10343) and Ichthypriapus might be referable to it. If this is true, then the genus Eumylodus Leidy (1873) has priority. In any case, Ichthypriapus is not an available genus name for P. gladius (contra Schultze et al. 1982; Stewart 1988). For this reason, we have created the new genus Bonnerichthys to accommodate this species. 14

16 5. SUPPLEMENTARY ANATOMICAL DATA: BONNERICHTHYS GEN. NOV. Fig. S7. Cranial remains of Bonnerichthys gladius gen. nov., KUVP 60692, from the Upper Cretaceous (probably lower Campanian) Smoky Hill Member, Niobrara Formation, Kansas, USA. (A) Right prearticular in lingual view. (B) Composite first left gill arch in left-lateral view. Abbreviations: add.fo, adductor fossa; cbr, ceratobranchial I; ebr, epibranchial I; hbr, hypobranchial I; ipb, infrapharyngobranchial I; part, prearticular. 15

17 Fig. S8. Associated pectoral fins and girdles of Bonnerichthys gladius gen. nov., KUVP 465, from the Upper Cretaceous (probably lower Campanian) Smoky Hill Member, Niobrara Formation, Kansas, USA. Abbreviations: cle, cleithrum; p.f, pectoral fin; ra.p, posterior pectoral radials. Paired structures listed as right (r) or left (l). This specimen was associated with the holotype of Ichthypriapus hubbsi. 16

18 Fig. S9. Endoskeletal pectoral girdle anatomy of Bonnerichthys gladius gen. nov. corresponds to that found in other pachycormids. (A) Right scapulocoracoid of Bonnerichthys gladius, KUVP 60692, from the Upper Cretaceous (probably lower Campanian) Smoky Hill Member, Niobrara Formation, Kansas, USA, shown in rightlateral view. (B) Shoulder girdle of Protosphyraena sp., KUVP 466, from the Smoky Hill Member, Niobrara Formation, Kansas, USA, shown in right-lateral view (left-hand specimen digitally reversed to match Bonnerichthys, with cleithrum faded). Ventral region of scapulocoracoid restored based on specimen AMNH FF Abbreviations: cle, cleithrum; fa.pro, articular facet for first pectoral radial (propterygium); fa.ra.p, articular facets for posterior pectoral radials; fa.ra.2, paired articular facets for second pectoral radials. 17

19 Fig. S10. Pectoral-fin radials of Bonnerichthys gladius gen. nov., KUVP 60692, from the Upper Cretaceous (probably lower Campanian) Smoky Hill Member, Niobrara Formation, Kansas, USA. (A) Right second pectoral radial in anterior view. (B) Right second pectoral radial in posterior view. (C) Representative posterior radials (uncertain handedness), with proximal articular surfaces placed at bottom of figure. Abbreviations: fa.d, dorsal facet on second pectoral radial for articulation with the scapulocoracoid; fa.p, proximal facet on posterior radial for articulation with the scapulocoracoid; fa.v, ventral facet on the second pectoral radial for articulation with the scapulocoracoid. 18

20 6. PALEOBIOGEOGRAPHIC DISTRIBUTION OF SUSPENSION-FEEDING PACHYCORMIDS Fig. S11. Paleobiogeographic distribution of suspension-feeding pachycormids. Icon for Bonnerichthys in the Late Cretaceous Western Interior Seaway corresponds to occurrences in Kansas (Niobrara Formation), Nebraska, South Dakota and Wyoming (all Pierre Shale), whereas that in the Gulf Coastal Plain indicates finds in Alabama, Arkansas, and Mississippi. Distributional data taken from Stewart (1988), Lambers (1992), Liston (2007, 2008), Parris et al. (2007), and references therin. Paleogeographic reconstructions modified from Ron Blakey, Northern Arizona University ( 19

21 7. CHARACTERS USED IN CLADISTIC ANALYSIS The taxon-by-character matrix used here derives in large part from that presented in Hurley et al. (2007). That analysis was concerned with the content of the neopterygian stem, and included a wide range of Paleozoic actinopterygians. The placement of pachycormids as crown neoptergyians is uncontroversial (e.g., Patterson 1973, 1977a; Gardiner et al. 1996; Brito 1997; Hurley et al. 2007), so the matrix found in Hurley et al. (2007) has been pared accordingly. Most members of crown Neopterygii from that original analysis were retained with the exception of Brachydegma, Mesturus, Hulettia, and Dapedium. Two stem neopterygians, the Carboniferous Discoserra and Triassic Pteronisculus, were retained, with Pteronisculus specified as the outgroup in all analyses. Characters from Hurley et al. (2007) that were parsimony-uninformative for this reduced taxon set were excluded. In addition to the edentulous forms that are the focus of the present analysis, the taxon sample found in Hurley et al. (2007) has been augmented with a series of additional pachycormids: Australopachycormus, Euthynotus, Hypsocormus, Orthocormus, and Protosphyraena. Characters with a bearing on pachycormid interrelationships were derived from Mainwaring (1978), Lambers (1988, 1992) and Liston (2007). Novel characters were formulated from personal observation of fossil materials or specimen descriptions in primary literature (see Section 8). Sources for identical or similar character formulations are indicated in brackets after character descriptions: C, Coates (1999); CS, Cavin and Suteethorn (2006); GML, Gardiner et al. (1996); GS, Gardiner and Schaeffer (1989); H, Hurley et al. (2007); La, Lambers (1992); Li, Liston (2007); Lu, Lund (2000); M, Mainwaring (1978); OM, Olsen and McCune (1991); P73, Patterson 1973; P77, Patterson (1977a); P82, Patterson (1982); PR, Patterson and Rosen (1977). Neurocranium and associated dermal bones 1. Opisthotic [CS, H]: 0. Present. 1. Absent. 20

22 2. Pterotic [H]: 0. Present. 1. Absent. 3. Pterotic fused with dermopterotic [GML, H]: 0. Absent. 1. Present. 4. Epioccipital [GML, H]: 0. Epioccipital bordered anteriorly by cranial fissure. 1. Epioccipital contacts otic region. 5. Intercalar [P73, CS, H]: 0. Present. 1. Absent. 6. Membranous outgrowths of intercalar [GML, H]: 0. Minor. 1. Extensive and medial to jugular. 2. Extensive and lateral to jugular. 7. Vagal foramen [GML, CS, H]: 0. Anterior to exoccipital. 1. Lateral outgrowths from intercalar form posterior margin. 2. Ventral outgrowths from intercalar lateral margin enclose dorsal margin. 3. Enclosed by exoccipital. 8. Lateral cranial canal [C]: 0. Present. 1. Absent. 9. Subtemporal fossa [GS, H]: 0. Absent. 1. Present. 10. Anterior myodome [P77, GML, A, C, CS, H]: 0. Present. 1. Absent. 11. Posterior myodome [P73, P77, GS, GML, C, CS, H]: 0. Intramural, lined by endoskeletal floor. 1. With incompletely ossified (fenestrate) floor. 2. Absent. 12. Cranial fissures [PR, G, H]: 21

23 0. Ventral otic and otico-occipital fissure confluent via vestibular fontanelle. 1. Fissures non-persistent (closed), or pattern obscured by incomplete ossification. 13. Relationship of dorsal aorta to basicranial region [GML, A, C, H]: 0. Enclosed in an endoskeletal canal. 1. Accommodated in pronounced aortic groove. 2. Accommodated in shallow depression in ventral midline of basioccipital. 3. Enclosed in parabasal canal between parasphenoid and basioccipital. 14. Lateral commissure breadth [H]: 0. Anteroposteriorly broad. 1. Slender. 15. Posttemporal fossa and fossa Bridgei [P73, P77, GS, GML, CS, H]: 0. Posttemporal fossa absent, fossa Bridgei rudimentary. 1. Posttemporal fossa small, fossa Bridgei discrete. 2. Posttemporal fossa large, fossa Bridgei discrete. 3. Posttemporal fossa communicates with fossa Bridgei. 16. Spiracular canal [P77, P82, GS, A, H]: 0. Present. 1. Absent. 17. Vomer(s) [P73, P77]: 0. Paired. 1. Unpaired (median). 18. Parashenoid corpus in sphenethmoid region: 0. Narrow. 1. Broad. 19. Parasphenoid dentition [A]: 0. Present. 1. Absent. 20. Basipterygoid process [P73, GS, GML, H]: 0. Well-developed dermal process with endoskeletal component. 1. Basipterygoid process absent. 21. Internal carotid foramen through parasphenoid [P77, GML, H]: 0. Absent. 1. Present. 22

24 22. Efferent pseudobranchial foramina through parasphenoid [P77, GML, H]: 0. Absent. 1. Present. Dermal skull roof and cheek 23. Rostral bone [GML, L, H]: 0. Cap on snout apex partially or wholly separating the nasals. 1. Of moderate to narrow size. 2. Reduced to a narrow tube with lateral process. 3. Composite ossification (rostrodermethmoid). 24. Rostrodermethmoid [M, L, Li]: 0. Terminates at anterior tip of mandibular symphysis. 1. Forms a produced rostrum, extending anterior to mandibular symphysis. 25. Marginal teeth on rostrodermethmoid: 0. Present. 1. Absent. 26. Paramedial fangs on rostrodermethmoid [M, L, Li]: 0. Absent. 1. Present. 27. Parietals [A]: 0. Paired. 1. Single midline ossification. 28. Posterior boss on skull roof [M, L, Li]: 0. Absent. 1. Present. 29. Posterior margin of skull roof: 0. Straight or concave. 1. 'W'-shaped. 30. Slender posterior processes of dermopterotics: 0. Absent. 1. Present. 31. Dermosphenotic [P73, GML, H]: 0. Hinged to skull roof. 1. Bound or fused to anterior margin of sphenotic. 23

25 32. Demosphenotic forms dorsal margin of orbit [M, L, Li]: 0. No. 1. Yes. 33. Supraorbitals [H]: 0. Absent. 1. Present. 34. Contact between anterior supraorbitals and infraorbitals [GML, H]: 0. Absent. 1. Present. 35. Suborbitals [P73, CS]: 0. Present. 1. Absent. 36. Suborbital size [M, L, Li]: 0. Equal in size or smaller than adjacent infraorbital(s). 1. Much larger than adjacent infraorbital(s). 37. Suborbital number [M, L, CS, Li, H]: 0. Two. 1. Single. 2. More than two. 38. Antorbital shape [H]: 0. Platelike, with minimal (if any) distinct anterior process. 1. Tapering towards slender anterior process; tri-radiate canal with broader, posterior portion. 2. Tubular. 39. Infraorbitals behind orbit [L, Li]: 0. Six or fewer. 1. More than six. 40. Infraorbitals anterior to circumorbital ring [OM, CS, H]: 0. Absent. 1. Present. 41. Posteriorly extensive infraorbital: 0. Present. 1. Absent. 42. Preopercle [OM, L, H]: 0. With broad dorsal margin. 24

26 1. With narrow ascending limb. Suspensorium, bones of the upper and lower jaws, and marginal dentition 43. Marginal dentition: 0. Present. 1. Absent. 44. Carinae extending length of teeth: 0. Absent. 1. Present. 45. Midline contact of premaxillae [L, Li]: 0. Present. 1. Absent. 46. Form of premaxillary dentition [L, Li]: 0. Similar to maxillary teeth. 1. Enlarged relative to maxillary teeth. 47. Mobile premaxilla [P73, P77, GS, GML, CS, H]: 0. Absent. 1. Present. 48. Grooved, notched or perforated premaxillary ascending process [P73, GS, GML, CS, H]: 0. Absent. 1. Present. 49. Maxilla [P73, GS, GML, H]: 0. Fixed to cheek. 1. Free. 50. Maxilla length [GML, L, CS, H]: 0. Long, extends beneath mid-orbit. 1. Short. 51. Posterior margin of maxilla [GML, L, CS, H]: 0. Straight or convex. 1. Indented. 52. Thickened ridge on external surface of maxilla: 0. Absent. 1. Present. 25

27 53. Supramaxilla [P73, GS, GML, A, CS, H]: 0. Absent. 1. Present. 54. Supramaxilla position [L, Li]: 0. Dorsal to maxilla. 1. Posterodorsal to maxilla. 55. Quadratojugal [P73, P77, GML, CS, H]: 0. Platelike, lateral to quadrate. 1. Splintlike, free along posterior border of quadrate. 2. Fused to quadrate. 3. Absent. 56. Symplectic articulation [P73, OM, GML, CS, H]: 0. On inner, medial surface of quadrate. 1. Behind quadrate, in loose contact with preopercle. 2. Behind quadrate, with articular connection to lower jaw, may be bound to preopercle with membrane bone. 57. Position of jaw joint [M, L, A, CS, Li, ]: 0. Well behind orbit. 1. Below or anterior to orbit. 58. Quadrate with elongate posteroventral process [A, CS, H]: 0. Absent. 1. Present. 59. Dentary tooth row(s) [M, L, Li]: 0. Labial row plus lingual row. 1. Single row. 60. Form of anterior dentary dentition [M, L, Li]: 0. Similar to posterior teeth. 1. Enlarged, procumbent fangs. 61. Surangular in lower jaw [P77, PR, P82, GS, H]: 0. Present. 1. Absent. 62. Anterior coronoid plate [M]: 0. Not inflated. 1. Inflated. 63. Prearticular [P77, PR]: 0. Present. 26

28 1. Absent. 64. Prearticular dentition: 0. Present. 1. Absent. Operculogular series, hyoid arch, and branchial arches 65. Exposed dorsal projection of subopercle between preopercle and opercle: 0. Absent. 1. Present. 66. Hypohyal [P77]: 0. Single ossification. 1. Separate dorsal and ventral ossifications. 67. Anterior ceratohyal [GML, H]: 0. Long, relatively straight, same depth as posterior element. 1. Long, gently curved with very small posterior element. 2. Short and deep posteriorly. 3. With constricted shaft; hourglass-shaped in lateral view. 68. Gular(s) [L, A, GML, C, CS, H]: 0. Present. 1. Absent. 69. Epibranchials [P73, GS, GML, H]: 0. Slender. 1. With uncinate processes. Median fins and axial skeleton 70. Relative position of dorsal and anal fins [M, L, Li]: 0. Overlap between dorsal- and anal-fin bases. 1. Dorsal-fin base lies anterior to anal-fin base. 71. Caudal fin segmentation [L, Li]: 0. Segmented. 1. Unsegmented. 72. Ossified vertebrae [L, Li]: 0. Absent. 1. Present. 73. Lateral keels of caudal peduncle (greatly expanded urodermals) [L]: 0. Absent. 27

29 1. Present. 74. Uroneurals [P73, P77, GML, A, CS, H]: 0. Absent. 1. Present. 75. Hypural plate [L, Li]: 0. Absent. 1. Present. 76. Shape of hypural plate: 0. Rhomboidal. 1. Oval. 77. Main hypural plate notched posterodorsally (incomplete fusion with accessory hypurals): 0. Present. 1. Absent. 78. Uppermost hypaxial-caudal rays [P73, GML, H]: 0. Fin-rays successively shorter from bottom to top. 1. Bundle of enlongate fin-ray bases extending over several hypurals. 2. Dorsal and ventral fin-ray bases symmetrical. 3. Fin-rays one-to-one on hypurals. Appendicular skeleton 79. Clavicle [P73, GML, L, H]: 0. Present. 1. Absent. 80. Clavicle morphology [P73, GML, L, H]: 0. Large, caps anterior end of cleithrum. 1. Toothed plates on postbranchial lamina of cleithrum. 2. Clavicle reduced, often with a single row of denticles. 3. Serrated organ (with 12 or more ridges of denticles lateral to cleithrum). 81. One or more accessory postcleithra [A, H]: 0. Absent. 1. Present. 82. Endoskeletal shoulder girdle ossification reduced to mesocoracoid arch [OM, CS, H]: 0. Absent. 1. Present. 28

30 Squamation 83. Pectoral propterygium [P77, P82, CS]: 0. Free. 1. Fused to first pectoral-fin ray. 84. Pectoral-fin radial morphology: 0. Cylindrical. 1. Broad distal area and narrow proximal stalk (paddle-shaped). 85. Scythe-like pectoral fins with lepidotrichia only segmented distally (if segments present) [M, L, Li]: 0. Absent. 1. Present. 86. Complete fusion of fin rays along length of leading edge of pectoral fin: 0. Absent. 1. Present. 87. Bifurcations in pectoral-fin rays: 0. Present. 1. Absent. 88. Asymmetrical 'Y'-type bifurcations in pectoral-fin lepidotrichia [L, Li]: 0. Absent. 1. Present. 89. Segmentation of pectoral-fin rays: 0. Present. 1. Absent. 90. Pelvic fins [M, L, Li]: 0. Present. 1. Absent. 91. Pelvic fin position: 0. At or posterior to midpoint between anal and pectoral fins. 1. Anterior to midpoint between anal and pectoral fins. 92. Ridge scales [GML, CS, H]: 0. Absent. 1. Present. 93. Scale morphology [P73, P77, PR]: 0. Rhombic. 29

31 1. Round. 30

32 8. COMPARATIVE MATERIALS Our discovery of new material belonging to large-bodied, edentulous pachycormids ( Bonnerichthys and Rhinconichthys) has led us to reconsider previous interpretations of anatomy in other members of this group (e.g., Lambers 1992; Liston 2007, 2008). The better-preserved specimens reported here have provided a new comparative framework for this earlier work, and these changes are reflected in character codings in Section 9. In cases where we have not been able to agree upon character states, we have adopted an agnostic approach and coded them as? to reflect our collective uncertainty. STEM NEOPTERYGII Incertae familiae Pteronisculus stensioei (Nielsen). Specimens described in Nielsen (1942). Guildayichthyidae Discoserra pectinodon Lund. Specimens described in Lund (2000) accompanied by anatomical reinterpretations included in Hurley et al (2007). HOLOSTEI Parasemionotidae Watsonulus eugnathoides Piveteau. Specimens described in Olsen (1984) and Grande and Bemis (1998). Ophiopsidae 31

33 Macrepistius arenatus Schaeffer. Two incomplete specimens, both from the Lower Cretaceous (Albian) Glen Rose Formation of the Trinity Group, Glen Rose, Texas, USA. AMNH FF 2435 (complete skull and partial postcranium); cast of USNM (AMNH FF uncatalogued). These specimens are described by Schaeffer (1960, 1971). Caturidae Caturus furcatus Agassiz. Two acid-prepared individuals, both from the Upper Jurassic (Tithonian) Solnhofen Limestone, Germany. AMNH FF 7492; AMNH FF Additional codings were made based on specimens described in Rayner (1948), Patterson (1975), Bartram (1977), Lambers (1992) and Grande and Bemis (1998). Caturus sp. One acid-prepared skull from the Upper Jurassic (Kimmeridgian) Solnhofen Limestones, Germany. AMNH FF Amiidae Amia calva Linnaeus. Specimens described in Grande and Bemis (1998). Semionotidae Lepidotes elvensis (Blainville). One complete individual, AMNH FF 7537, from the Lower Jurassic (Toarcian) Posidonia Shale of Holzmaden, Germany. Additional characters are coded from specimens described in Quenstedt (1847). Lepidotes sp. One incomplete individual, AMNH FF 11442, from the Middle Jurassic (Bathonian) Sundance Formation, Burnt Hollow locality, Wyoming, USA. Semionotus elegans (Newberry). Codings made based on specimens described in Olsen and McCune (1991). 32

34 Lepisosteidae Lepisosteus platostomus Rafinesque. Specimens described in Mayhew (1924). Macrosemiidae Macrosemius rostratus Agassiz. Specimens described in Bartram (1977). TELEOSTEI Pachycormidae Asthenocormus titanius (Wagner). One complete individual, SMGP BaJ 2344, from the Upper Jurassic (Kimmeridgian) Solnhofen limestone of Germany. Additional codings were made based on specimens described in Lambers (1992). Australopachycormus hurleyi Kear. Cast of caudal fin, BMNH P.59695, and nearly complete pectoral fin, BMNH uncatalogued, from the Lower Cretaceous (Albian) Toolebuc Formation of the Rolling Downs Group, Queensland, Australia (specimen attributed to Protosphyraena). Additional codings were made based on specimens described in Kear (2007). Bonnerichthys gladius gen. nov. (Cope). Twenty-five specimens, all from the Upper Cretaceous (Coniacian-Maastrichtian) of USA. Smoky Hill Member (Coniacian-Campanian) of the Niobrara Formation, Kansas: eighteen specimens. 33

35 Fragmentary pectoral fin, AMNH FF 1849 (holotype), near the Solomon River (upper Solomon River, probably Rooks County and likely uppermost Coniacianlowermost Santonian); fin fragments, AMNH FF 2064, Fossil Spring Cañon, Logan County; fin fragments, AMNH FF 6314, western Kansas; fin fragments, AMNH FF 6315, western Kansas; fin fragments, AMNH FF 6316, Gove County; incomplete fin, FHSM VP-212, Gove County (upper Coniacian); hypural plate, FHSM VP-17427, Logan County; associated cranial and postcranial remains, FHSM VP-17428, Gove County; fin fragment, FHSM VP-17457, Gove County (lowermost Santonian); fin fragment, KUVP 439, Graham County; complete pectoral fins and cleithra, KUVP 464, Logan County (this material is associated with KUVP 1136, the holotype of Ichthypriapus hubbsi Hibbard, which we agree is a holocephalan cranial clasper [contra Schultze et al. 1982; Stewart 1988; see Section 4]); incomplete cleithrum, scapulocoracoid, and pectoral radials, KUVP 13995, Logan County; fin fragment, KUVP 49505, Martins Canyon, Gove County; fin fragment, KUVP 55592, Gove County; fragmentary fin, KUVP 60620, Logan County; associated skull, pectoral fins and girdles, KUVP 60692, Twin Butte Creek, Logan County; fin fragments, KUVP 65702, Gove County. Note: FHSM VP-17427, and probably other specimens from Logan County, is lowermost Campanian in age. Unless noted otherwise, material of Bonnerichthys from additional Niobrara localities likely dates to the Santonian (Stewart 1988). Pierre Shale Formation (Campanian-Maastrichtian) of the western interior USA, four specimens. Incomplete fin, AMNH FF 19288, from the Pierre Shale (Campanian), 20 miles southwest of Edgemont, South Dakota; fragmentary fin, KUVP 338, from the Pierre Shale, Logan County, Kansas (holotype of Protosphyraena gigas); eroded hypural plate, UNSM 50999, from the Mobridge Chalk Member (Maastrichtian) of the Pierre Shale, northeast Nebraska, USA; fragmentary fins, pectoral girdles 34

36 and hypural plate, UNSM 88507, from the base of the Pierre Shale Formation (Campanian), south shore of Harlan County reservoir, Nebraska. Mooreville Formation (lower Campanian) of the Selma group, Alabama: four specimens. Fin fragments, FMNH PF 27363, Hale County; fin fragments, FMNH PF 27364, Hale County; fin fragments, FMNH PF 27365, Barber County; cleithrum, pectoral fin fragments, and radials, KUVP 50179, Barber County. Remains of Bonnerichthys not studied here include specimens reported from the Campanian-Maastrichtian of the Atlantic coastal plain (Parris et al. 2007), the Demopolis Chalk (late Campanian) of Alabama and Mississippi (Stewart 1988), and the Pierre Shale (Campanian) of Wyoming (Stewart 1988). Euthynotus incognitus (Blainville). One complete individual, AMNH FF 7540, from the Lower Jurassic (Toarcian) Posidonia Shale of Holzmaden, Germany. Additional codings were made based on specimen descriptions in Wenz (1967). Euthynotus cf. E. incognitus. Specimen described in Bürgin (2000). Euthynotus intermedius (Agassiz). Specimens described in Arambourg (1935). Euthynotus sp. Caudal specimen described in Arratia and Lambers (1996). Hypsocormus insignis Wagner. Four individuals, all from the Upper Jurassic (Kimmeridgian), Solnhofen Limestones, Germany. AMNH FF 7487, BMNH P.954, BMNH P.4236, BMNH P Additional codings were made based on account given by Woodward (1895). Hypsocormus sp. Specimen described in Woodward (1908). 35

37 Hypsocormus tenuirostris Woodward. Four specimens, all from the Late Jurassic (Callovian) Oxford Clay near Peterborough, UK. Incomplete caudal and pectoral fins, BMNH P.6719; fragmentary skull, BMNH P.6916; nearly complete skull, BMNH P.10579; incomplete skull, BMNH P Leedsichthys problematicus Woodward. Specimens described in Liston (2007). Martillichthys renwickae Liston. One specimen, BMNH P from the Middle Jurassic (Callovian) Oxford Clay, Dogsthorpe Pit of the London Brick Company, Peterborough, UK. Orthocormus cornutus Vetter. AMNH FF (cast of BSP 1993 I 22) from the Upper Jurassic (Tithonian), Solnhofen Limestone near Kallmünz, Oberpfalz, Germany. Additional codings were based on specimen descriptions in Holmgren and Stensiö (1936), Rayner (1948), Jessen (1972) and Lambers (1988, 1992). The first three references refer to this taxon as Hypsocormus sp. Orthocormus tyleri Lambers. Specimen described in Lambers (1988, 1992). Pachycormidae gen. et sp. indet. One incomplete skull preserved in the round, BMNH P.41669, from the top of the Middle Jurassic (Bajocian) Inferior Oolite of Dorset (possibly near Sherborne), UK. Pachycormus curtus Agassiz. Specimens described in Arambourg (1935), Rayner (1948), Wenz (1967), Patterson (1973, 1975) and Mainwaring (1978). Pachycormus macropterus (Blainville). Three acid-prepared skulls, BMNH P24410 and BMNH 32436, BMNH from the Lower Jurassic (Toarcian) of Curcy, Normandy, France. Specimens described in Woodward (1908), Wenz (1967) and Mainwaring (1978). 36

38 Protosphyraena sp. Five individuals, all from the Upper Cretaceous (Coniacianlowermost Campanian) Smoky Hill Member of the Niobrara Formation, Kansas, USA. Scapulocoracoid and incomplete cleithrum, AMNH FF 1901, south fork of Solomon River (labeled as Protosphyraena perniciosa); complete skull and mandibles, AMNH FF 1966, NE of WaKeeny, Trego County; complete skull and mandibles, AMNH FF 4318 (labeled as Protosphyraena perniciosa); cleithrum and scapulocoracoid, KUVP 466, near Banner, Trego County; caudal fin and incomplete axial column, KUVP 49419; Logan County. Additional codings made from material described in Felix (1890). Protosphyraena nitida (Cope). Three individuals, all from the Upper Cretaceous (Coniacian-lowermost Campanian) Smoky Hill Member of the Niobrara Formation, Kansas, USA. Complete skull, pectoral and pelvic fins, BMNH P.10341, locality unknown; skull plus fragmentary pectoral fins and girdles, AMNH FF 1634, Gove County; skull plus pectoral girdles and partial fins, cast of LACM , Ellis County (upper Coniacian). Protosphyraena cf. P. tenuis Loomis. Four individuals, all from the Upper Cretaceous (Coniacian-lowermost Campanian) Smoky Hill Member of the Niobrara Formation, Kansas, USA. Incomplete skull, KUVP 420; mandibles, sclerotic ring, hypohyal, hyomandibula, neurocranium, KUVP 398, Gove County; skull and pectoral fins, KUVP 55489, Trego County; scapulocoracoids and partial suspensorium, KUVP 55499, Logan County. Protosphyraena perniciosa (Cope). Three individuals, all from the Upper Cretaceous (Coniacian-lowermost Campanian) Smoky Hill Member of the Niobrara Formation, Kansas, USA. Material of this species appears to be confined to the upper Coniacian (Stewart 1988). Pectoral fins and girdle, AMNH FF 2009, Rooks County; pectoral fin and girdle, KUVP 367, Gove County; fragmentary scapulocoracoid, pectoral fin and radials, KUVP 56612, Trego County. 37

39 Rhinconichthys taylori gen. et sp. nov. One incomplete skull preserved in the round, BMNH 219, from the Lower Chalk (Cenomanian) of Burnham, Kent, UK. Rhinconichthys cf. R. taylori. One incomplete skull preserved in the round, NMS PV20868 plus PV20869 (cast BMNH P.66822), Late Cretaceous (likely Cenomanian) Mikasa Formation, Middle Yezo Group, Ikushumbetsu, Mikasa City, Hokkaido, Japan. PHOLIDOPHORIDAE Pholidophorus bechei Agassiz. Specimens described in Rayner (1948), Nybelin (1966) and Patterson (1975). HIODONTIDAE Hiodon alosoides (Rafinesque). Specimens described in Hilton (2002). ELOPIDAE Elops hawaiensis Regan. Specimens described in Forey (1973). 38

40 9. TAXON-BY-CHARACTER MATRIX Pteronisculus spp Amia calva Asthenocormus titanius??????????????? Australopachycormus hurleyi 0 0?? 0?????????? Bonnerichthys gladius 0 0 0? 0??????? 1 1? Caturus spp Discoserra pectinodon 0 0? ??? 0 1? Elops hawaiensis Euthynotus spp??????????????? Hiodon alosoides Hypsocormus insignis??????????????? 'Hypsocormus' tenuirostris??????????????? Leedsichthys problematicus???????????? 2?? Lepidotes spp ? 3 0 1? Lepisosteus platostomus ? Macrepistius rostratus ? ? 1 2 Macrosemius rostratus ?? 3 0?? 1 1 2?? Martillichthys renwickae??????????????? Orthocormus spp 0??? 0? 1?? 1? 0? 1? Pachycormus spp ? Pholidophorous bechei Protosphyraena spp ? 1? 1?? Rhinconichthys taylori? 0 0????? 1? 1? Semionotus elegans ? 3??? 1? 2?? Watsonulus eugnathoides 0 0? ? Oolite pachycormid??????????????? - = logical impossibility or inapplicable character;? = unknown state 39

41 Pteronisculus spp Amia calva Asthenocormus titanius??????? Australopachycormus hurleyi??????? ? Bonnerichthys gladius? ??? 1 Caturus spp? Discoserra pectinodon 0??? 1 0? ? 0 Elops hawaiensis 1??? Euthynotus spp?? 0???? ? 0 0? Hiodon alosoides Hypsocormus insignis? 0????? ? 1? 0 'Hypsocormus' tenuirostris? ?? ? Leedsichthys problematicus?? 1 1????? - -???? Lepidotes spp? Lepisosteus platostomus Macrepistius rostratus?? Macrosemius rostratus? ? Martillichthys renwickae?? 1 1??? Orthocormus spp? 0 0???? ?? Pachycormus spp Pholidophorus bechei Protosphyraena spp? Rhinconichthys taylori? 0 1? 1???? Semionotus elegans? ? Watsonulus eugnathoides Oolite pachycormid??????? ???? - = logical impossibility or inapplicable character;? = unknown state 40

42 Pteronisculus spp Amia calva Asthenocormus titanius?? 0 -???????? 1 -? Australopachycormus hurleyi???????????? Bonnerichthys gladius ??? 0???? Caturus spp Discoserra pectinodon ? Elops hawaiensis Euthynotus spp ? Hiodon alosoides Hypsocormus insignis?? ? 1? 'Hypsocormus' tenuirostris???? 0?????? Leedsichthys problematicus???????????? 1 -? Lepidotes spp Lepisosteus platostomus Macrepistius rostratus ? Macrosemius rostratus Martillichthys renwickae ??????? 1 1 -? Orthocormus spp ? Pachycormus spp Pholidophorus bechei Protosphyraena spp ? 0? 0?? Rhinconichthys taylori???????????? 1 -? Semionotus elegans Watsonulus eugnathoides Oolite pachycormid???????????? 1 -? - = logical impossibility or inapplicable character;? = unknown state 41

43 Pteronisculus spp Amia calva Asthenocormus titanius -?? 1 0 0? 0 -?? 1? - - Australopachycormus hurleyi 1 0 0?? 0??????? 1 1 Bonnerichthys gladius ? Caturus spp Discoserra pectinodon Elops hawaiensis ? 0 Euthynotus spp ? 1 0 Hiodon alosoides Hypsocormus insignis 0 0? 1 0 0? 1 1?? 0? 0 0 'Hypsocormus' tenuirostris ?? 0? 0 1 Leedsichthys problematicus -?? 1????????? - - Lepidotes spp Lepisosteus platostomus Macrepistius rostratus 0 0? ?? 1?? 0 Macrosemius rostratus ? Martillichthys renwickae -?? Orthocormus spp ?? 0? 0 1 Pachycormus spp ? Pholidophorus bechei Protosphyraena spp ? Rhinconichthys taylori -?? ?? 0? - - Semionotus elegans Watsonulus eugnathoides Oolite pachycormid???? 0?????????? - = logical impossibility or inapplicable character;? = unknown state 42

44 Pteronisculus spp Amia calva Asthenocormus titanius???? 0? 0 0?? 1 0?? 1 Australopachycormus hurleyi? 1???????? 1 0??? Bonnerichthys gladius?? 0 1 0? 0 0 0?? 0?? 1 Caturus spp Discoserra pectinodon???? -? 3 0? Elops hawaiensis Euthynotus spp???? 0???? Hiodon alosoides Hypsocormus insignis???? 0?? 0? 'Hypsocormus' tenuirostris ?????? 0 0?? 1 Leedsichthys problematicus????? 0? 0 0? 1 0??? Lepidotes spp? ? Lepisosteus platostomus Macrepistius rostratus?? 0 0 1? 1 0? ? Macrosemius rostratus Martillichthys renwickae ? ??? Orthocormus spp?????? 0?? ? Pachycormus spp Pholidophorus bechei 0? 0? 0 0? 0? Protosphyraena spp ? 0? 0?? 1 0? 1 1 Rhinconichthys taylori?? 0?? ?? 0??? Semionotus elegans Watsonulus eugnathoides ? Oolite pachycormid?? 0???? 0??????? - = logical impossibility or inapplicable character;? = unknown state 43