Appendix 1. List of valid plesiosaur species with information on...

Appendix 1. List of valid plesiosaur species with information on authors and date of publication.

Date Original paper Renamer Species Age1824 De la Beche and Conybeare 1824 Plesiosaurus dolichodeirus Sinemurian1876 Blake in Tate and Blake 1876 Plesiosaurus' longirostris Toarcian1837 Buckland in Conybeare 1837 Plesiosaurus' macrocephalus Sinemurian1841 Owen 1841 Pliosaurus brachydeirus Kimmeridgian2004 Noe et al 2004 Pliosaurus portentificus Kimmeridgian1841 Owen 1841 Pliosaurus brachyspondylus Kimmeridgian1948 (Novozhilov 1948) Tarlo1960 Pliosaurus irgisensis Tithonian1871 (Phillips 1871) Noe et al. 2004 Pliosaurus macromerus Callovian-Tithonian1960 Tarlo 1960 Pliosaurus andrewsi Callovian1841 Owen 1841 Polyptychodon interruptus Turonian-Santonian1868 Cope 1868, Elasmosaurus platyurus Campanian1869 Cope 1869 Polycotylus latipinnis Santonian1948 (Novozhilov 1948) Halstead 1971 Liopleurodon rossicus Tithonian1869 (Seeley 1869) Tarlo 1960 Liopleurodon pachydeirus Callovian1873 Sauvage1873 Liopleurodon ferox Callovian1874 Hector 1874 Mauisaurus haasti Campanian1865 (Owen 1865) Seeley 1874 Eretmosaurus rugosus Sinemurian1874 Seeley 1874 Muraenosaurus leedsi Callovian1892 (Seeley 1892) Brown 1981 Muraenosaurus beloclis Callovian1863 (Carte and Bailey 1863) Seeley 1874 Rhomaleosaurus cramptoni Toarcian1846 (Stutchbury 1846) Andrews 1922 Rhomaleosaurus megacephalus Hettangian1910 (Fraas 1910) Tarlo 1960 Rhomaleosaurus victor Toarcian1840 (Owen 1840) Seeley 1874 Colymbosaurus trochanterius Kimmeridgian-Tithonian1881 (Sollas 1881) Bakker 1993 Attenborosaurus conybeari Sinemurian1888 Cragin 1888 Trinacromerum bentonianum Turonian1935 Russell 1935 Trinacromerum kirki Turonian1869 (Seeley 1869) Lydekker 1889 Peloneustes philarchus Callovian1871 Phillip(s 1871) Seeley 1892 Cryptoclidus eurymerus Callovian1889 (Lydekker 1889) Bogolubov, 1909 Cryptoclidus richardsoni Callovian1898 Knight 1898 Megalneusaurus rex Kimmeridgian-Portlandian1902 Williston 1902 Dolychorhynchops osborni Campanian2005 Sato 2005 Dolichorhynchops herschelensis Campanian-Maastrichtian1903 Williston 1903 Brachauchenius lucasi Cenomanian-Turonian1994 Godefroit 1994 Simolested keileni Bajocian1877 Lydekker 1877 Bardet et al. 1991 Simolestes indicus Tithonian1909 Andrews 1909 Simolestes vorax Callovian1909 Andrews 1909 Tricleidus seeleyi Callovian1865 (Owen, 1865) Watson 1909 Microcleidus homalospondylus Toarcian1865 Seeley, 1865? Watson 1909 Microcleidus macropterus Toarcian1909 Watson, 1909 Sthenarasaurus dawkins Toarcian1913 Brown 1913 Leurospondylus' ultimus Maastrichtian1914 Wegner 1914 Brancasaurus brancai Berriasian1840 (Owen 1840) Andrews 1922 Eurycleidus arcuatus Hettangian1911 (Andrews 1911) Cruickshank 1997 Leptocleidus capensis Valanginian1922 Andrews 1922 Leptocleidus superstes Barremian1997 Cruickshank and Long, 1997 Leptocleidus clemai Hauterivian-Barremian1924 Longman 1924 Kronosaurus queenslandicus Aptian-Albian1992 Hampe, 1992 Kronosaurus boyacensis Aptian-Albian1930 Swinton 1930 Macroplata tenuiceps Hettangian1895 (Dames 1895) White 1940 Seeleysaurus guilelmiimperatoris Toarcian1941 Cabrera 1941 Aristonectes parvidens Maastrichtian1943 Welles 1943 Aphrosaurus furlongi Maastrichtian1890 (Williston 1890) Welles 1943 Styxosaurus snowii Santonian-Campanian1890 Welles 1943 Morenosaurus stocki Maastrichtian1943 Welles 1943 Thalassomedon haningtoni Cenomanian1949 (Welles 1949) Carpenter 1997 Libonectes morgani Turonian2006 Buchy 2006 Libonectes atlasense Turonian1871 (Cope 1871) Welles 1943 Hydralmosaurus serpentinus Santonian-Campanian1943 Welles 1943 Fresnosaurus drescheri Maastrichtian1943 Welles 1943 Hydrotherosaurus alexandrae Maastrichtian1865 (Owen 1865) Novozhilov 1964 Archaeonectrus rostratus Sinemurian1976 (Ochev 1976) Ochev 1977 Georgiasaurus penzensis Santonian1980 Dong 1980 Bishanopliosaurus youngi Toarcian1981 Brown 1981 Kimmerosaurus langhami Kimmeridgian

Appendices Appendix 1

257

1982 Delair 1982 Bathyspondylus swindoniensis Kimmeridgian1985 Zhang 1985 Yuzhoupliosaurus chengjiangensisBathonian1986 Wiffen and Moisley 1986 Tuarangisaurus keysei Campanian-Maastrichtian1996 Carpenter 1996 Plesiopleurodon wellesi Cenomanian1996 Cruickshank et al. 1996 Pachycostasaurus dawni Callovian1840 (Owen 1840) Storrs and Taylor 1996 Thalassiodracon hawkinsii Hettangian1997 Gasparini 1997 Maresaurus coccai Bajocian1990 (Sciau et al. 1990) Bardet et al. 1999 Occitanosaurus tournemirensis Toarcian1962 (Welles, 1962) Carpenter 1999 Callawayasaurus colombiensis Aptian2001 O,Keefe 2001 Hauffiosaurus zanoni Toarcian2002 Cruickshank and Fordyce 2002 Kaiwhekea katiki Maastrichtian2002 Druckenmiller 2002 Edgarosaurus muddi Albian1949 (de la Torre and Rojas 1949) Gasparini et al. 2002 Vinielasaurus caroli Callovian2003 Bardet et al. 2003 Thililua longicollis Maastrichtian2003 Sato 2003 Terminonatator ponteixensis Campanian2006 Kear 2006 Umoonasaurus demoscyllus Aptian-Albian2006 Kear 2006 Opallionectes andamookaensis Aptian-Albian2005 (Sachs 2005) Kear 2006 Eromangasaurus australis Albian1893 Marsh 1893 Pantosaurus striatus Oxfordian1900 (Knight 1900) O'Keefe and Wahl 2003 Tatenectes laramiensis Oxfordian1990 Gasparini and Spalleti, 1990 Sulcusuchus erraini Maastrichtian1923 (Huene 1923) Grossman 2007 Hydrorion brachypterygius Toarcian1942 Young 1942 Sinopliosaurus weiyuanensis Kimmeridgian?2006 Sato et al 2006 Futabasaurus suzukii Santonian2005 Buchy et al. 2005 Manemergus anguirostris Maastrichtian2007 Albright et al. 2007 Eopolycotylus rankini Turonian2007 Albright et al. 2007 Palmula quadratus Turonian


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Appendix 2. Additional material

The following list outlines additional fossil material adopted in the cladistic analysis

and/or the morphometric analysis, but not figured or described herein. The list begins

with specimens of uncertain taxonomic affinity (1A) and continues with a taxon list

(1B). The taxon list includes the type and referred material employed in the analyses

(the lists of referred material are therefore not exhaustive), and data on the type

locality and horizon. Although not figured or described in this thesis, some of this

material was observed first hand (observed material is indicated by the following

symbol proceeding the specimen number: ***), otherwise the taxa were coded from

the literature, as listed in the ‘remarks’ section. The taxa in Appendix 1B are listed in

the order that they appear in the cladograms resulting from the cladistic analysis

(Figure 5.7).

Appendix 1A - Additional specimens of uncertain taxonomic affinity

OUM J.2858 ‘Eurycleidus’

Cruickshank (1994a) referred OUM J.28585, a specimen from Lyme Regis, to

Eurycleidus arcuatus. The specimen consists of a partial skull including the mandible

and some postcranial elements (from the pelvic girdle). However, this specimen may

represent a new taxon in itself (O’Keefe 2004b). To test affinities of OUM J.28585, it

is treated as a separate OTU in the current cladistic analysis, albeit coded from the

literature.

HALB ‘Uncatalogued’

This specimen from the Toarcian of Kanonenberg, northern Germany, was described

by Brandes (1914) and referred to Thaumatosaurus aff. megacephalo (=

Rhomaleosaurus megacephalus). However, Grossman (2007, p. 557-558) noted that

this specimen “resembles Rhomaleosaurus victor more than R. megacephalus and it

is probably synonymous with the German taxon [R. victor]”. The specimen was

included in the morphometric analysis based on measurements given by Brandes

(1914).

Appendix 1B - Additional taxa

Cymatosaurus Fritsch, 1894


259

Type material. IGWH uncatalogued, skull.

Type locality. Halle-Nietleben, Saxony-Anhalt, Germany.

Type horizon. Myophoria beds, Upper Röt Formation (Beneckeia buchi and

Dadocrinus assemblage zone), Upper Buntsandstein (Lower Anisian, Middle

Triassic).

Remarks. A review of Cymatosaurus is presented by Rieppel (2000).

Pistosaurus longaevus Meyer, 1839

Type material. Uncatalogued, Oberfränkisches Erdgeschichliches Museum,

Bayreuth, skull.

Type locality. Lainecker Hohenzug, east of Bayreuth, southern Germany.

Type horizon. Lower Meissner Formation, Upper Muschelkalk (atavus to

postspinosus zones), (Upper Anisian, Middle Triassic).

Referred material. SMF R 4041, postcranial skeleton.

Remarks. Some authors have considered Pistosaurus as a member of the

Plesiosauria (e.g. White, 1940, p. 459). Rieppel (2000) presents a concise summary

of the unusual history of this genus. A cast of the holotype skull was investigated first

hand (SMNS ‘uncatalogued’) ***

Augustasaurus hagdorni Sander, Rieppel and Bucher, 1997

Type material: Holotype: FMNH PR 1974, complete skull and mandible, post-cranial

skeleton consisting of posterior neck vertebrae, pectoral girdle, majority of forelimbs,

dorsal vertebrae and associated fragmentary ribs and gastralia.

Type locality: Pershing County, Nevada, USA.

Type horizon: Fossil Hill Member, Farvet Formation, Star Peak Group, Rotelliformis

Zone, Late Anisian, Middle Triassic.

Remarks: The postcranium of A. hagdorni is described by Sander et al. (1997) and

the cranium is described by Rieppel et al. (2002). This taxon has important

implications because it is only the second non-plesiosaurian sauropterygian to be

discovered in the New World; Corosaurus alcovensis was the first (Storrs, 1991).

Yunguisaurus liae Cheng, Sato, Wu and Li, 2006

Type material: NMNS 004529/F003862, almost complete articulated specimen

missing the distal portion of the tail.

Type Locality: Near Huangnihe River, Chajiang, 10km northwest of Xingyi, Guizhu.

Type Horizon: Falang Formation, Carnian (early Late Triassic).


260

Remarks: The skull and limbs of Yunguisaurus were described by Cheng et al.

(2006) but “the postcranial skeleton waits for further preparation and subsequent full

description” (p.501). Consequently, some characters, presumably preserved but not

described or figured for this taxon, could not be coded in this cladistic analysis; this is

taken into account in the discussion (Chapter 6).

Plesiosaurus dolichodeirus Conybeare, 1824

Type material. BMNH 22656, complete skeleton. ***

Type locality. Lyme Regis, Dorset, England.

Type horizon. Black Ven Marl Formation, Lower Lias Group, (Echioceras

raricostatum Zone), (Uppermost Sinemurian, Lower Jurassic).

Referred material. NMING F8758 ***

Remarks. This genus was described by Owen (1865) and extensively revised by

Storrs (1997).

Hydrorion brachypterygius (Huene, 1923) Grossman, 2007

Type material. GPIT/477/1/1; complete skeleton including skull.

Type locality. Steinbruch 29, Ohmden (near Holzmaden), Baden-Württemberg,

Germany.

Type horizon. Lias epsilon II, 4 (Unterer Schiefer), Posidonienschiefer, (Harpoceras

falcifer zone), (Lower Toarcian, Lower Jurassic).

Referred material. MB.R. 1991, SMNS 51143.

Remarks. The holotype skull is described by Maisch and Rucklin (2000) as

Plesiosaurus brachypterygius; Grossman (2006, 2007) redescribed the material

pertaining to this taxon and introduced the new genus name Hydrorion.

Seeleysaurus guilelmiimperatoris (Dames, 1895) Grossman, 2007

Type material. MB R.1992, complete skeleton.

Type locality. Holzmaden, Baden-Württemberg, Germany.

Type horizon. Lias Epsilon II, 4 (Unterer Schiefer), Posidonienschiefer, (Harpoceras

falciferum zone), (Lower Toarcian, Lower Jurassic).

Referred material. SMNS 16812, more or less complete skeleton ***, SMNS 12039,

(specimen figured by Fraas [1910]).

Remarks. Dames (1895) and Fraas (1910) described this taxon. This species was

renamed ‘Seeleysaurus’ by White, (1940). Storrs (1997) modified Persson’s (1963)

suggestion that Plesiosaurus contains only three valid species by proposing P.

brachypterygius as a junior synonym of P. guilielmiimperatoris, reducing the number


261

to two. However, Maisch and Rucklin (2000, p. 38) justify retention of P.

brachypterygius. Some workers (Bakker, 1993) have retained the name

‘Seeleyosaurus’ for P. guilielmiimperatoris and recently, Grossman (2007) formalised

this taxonomy.

O’Keefe (2004b) erected a new genus and species name for SMNS 16812

(Plesiopterys wildii), but the specimen was later regarded as a juvenile Seeleysaurus

guilelmiimperatoris (Grossman, 2007).

‘Plesiosaurus’ longirostris Tate and Blake, 1876

Type material: MCZ 1033 (in the collection of fossil reptiles and amphibians),

damaged skull, 85 vertebrae and 4 limbs.

Type locality: Whitby, Yorkshire, England.

Type horizon: Alum Shale, Upper Lias, serpentinus ammonite zone, Toarcian, Lower

Jurassic.

Referred material: MM 8004, skull and associated skeleton.

Remarks: ‘Plesiosaurus’ longirostris has an extremely confusing taxonomic history,

complicated by the perplexing history of the type material. White (1940) included the

species ‘Plesiosaurus’ longirostris in the genus Macroplata (see below). Other

authors have placed it in Rhomaleosaurus (Broadhurst and Duffy, 1970). However,

even after superficial examination, it is clear that Plesiosaurus ‘longirostris’ is far

removed morphologically from both Rhomaleosaurus and Macroplata, not least in the

extreme elongation of the rostrum (pers. obs.). O’Keefe (2004b) contended the

placement of the species longirostris in Macroplata (see Chapter 3). Plesiosaurus

longirostris may be far more abundant than previously recognised: another

contemporary taxon, Sthenarosaurus dawkinsi (see below), may also pertain to the

species longirostris (R. Forrest pers. comm., 2005). Even Watson (1909), in his

original description of Sthenarosaurus, suggested the possibility that the type of S.

dawkinsi was congeneric with ‘Plesiosaurus’ longirostris. This cladistic analysis will

test this contention.

Sthenarosaurus dawkinsi Watson, 1909

Type material. MM L8023, a partial postcranial skeleton including both girdles, propodials, and

vertebrae.

Type locality. Saltwick, Yorkshire, UK.

Type horizon. Upper Lias (D. commune zone) (Toarcian, Lower Jurassic)

Remarks. Sthenarosaurus was described and figured by Watson (1909), no cranial

material has been referred to this taxon.


262

Maresaurus coccai Gasparini, 1997

Type material: MOZ 4386 V Museo Prof. Olsacher, Zapala, Neuquén, articulated

skull and mandible, atlas-axis and first cervical vertebrae.

Type locality: Chacaico Sur, (39˚ 15’ S, 70˚ 18’ W), 70 km southwest of Zapala,

Neuquén Province, Argentina.

Type horizon: Upper part of the Los Molles Formation, Cuyo Group, Emileia giebeli

and Emileia multiformis subzone, Lower Bajocian, Middle Jurassic.

Remarks: Diagnosed as a pliosaurid under the current limitations of poor

understanding of pliosaur phylogeny. Noè (2001) suggests that this may by a junior

synonym of Simolestes. The cladistic analysis presented here will test this

contention.

Hauffiosaurus zanoni O’Keefe, 2001 ***

Type material: HAUFF ‘Uncatalogued’, complete skeleton on display in the Urwelt-

Museum Hauff.

Type locality: Holzmaden, Germany.

Type horizon: Posidonien-schiefer, Toarcian, Lower Jurassic

Remarks: O’Keefe (2001a) briefly described this specimen and proposed the new

taxon name.

Brachauchenius lucasi Williston, 1903

Type material: USNM 4989, skull and mandible (palatal view) plus 37 ribs and

associated vertebrae.

Type locality: Ottawa County, Kansas, USA.

Type horizon: Greenhorn Limestone, Turonian, Late Cretaceous.

Referred material: USNM 2361, skull and vertebrae; FMNH VP321, compete skull.

Remarks: This taxon is described by Williston (1903) and was reviewed by Carpenter

(1996).

Kronosaurus queenslandicus Longman, 1924

Type material: Queensland Museum (unspecified)

Type locality: Army Downs, north of Richmond, Queensland, Australia.

Type horizon: Aptian-Albian

Referred material: MCZ 1284, 1285.


263

Remarks: The skull of Kronosaurus queenslandicus was described by White (1935)

and the postcranium by Romer and Lewis (1959). The genus Kronosaurus is

currently under revision (C. McHenry, pers. comm., 2003).

Simolestes vorax Andrews, 1909

Type material: BMNH R.3319, complete skeleton. ***

Type locality: Peterborough, England.

Type horizon: Lower Oxford Clay, Callovian, Middle Jurassic

Referred material: PETMG R296.

Remarks: Simolestes was described by Andrews (1913) based on the holotype. The

most recent treatments of Simolestes are unpublished – Noè (2001) thoroughly

described the holotype skull and also referred a second skull to this taxon (PETMG

R296) originally discovered in 1990 and identified as Liopleurodon (Dawn, 1991).

Liopleurodon ferox Sauvage, 1873

Type material: BMNH R.3536, a single tooth (but see below).

Type locality: Wast, Boulogne, France.

Type horizon: Callovian, Middle Jurassic.

Referred material: Associated skeleton (also BMNH R.3536), “the teeth of which are

indistinguishable from the type” (Tarlo, 1960, p.166) and BMNH R.2680 and GPIT

1754/2 (O’Keefe, 2001a). ***

Remarks: The skull of Liopleurodon ferox has been described by Andrews (1913). L.

pachydeirus, also from the Callovian (Tarlo, 1960), has been recently recognised as

a junior synonym of L. ferox (Noè, 2001). Noè et al. (2003) recently described the

exoccipital-opisthotic bone in this taxon. According to Noè et al. (2004), the species

Liopleurodon macromerus should be assigned to Pliosaurus.

Peloneustes philarchus (Seeley, 1869) Lydekker, 1889a

Type material: CAMSM J.46913, almost complete skeleton.

Type locality: Peterborough, England.

Type horizon: Oxford Clay, Callovian, Upper Jurassic

Referred material. SMNS 10113 (***)

Remarks: Peloneustes has been described by Lydekker (1889) and Andrews (1895,

1910b and 1913). According to Martill (1991), Peloneustes is often difficult to

distinguish from Pliosaurus and some species may possibly be congeneric.

Peloneustes is currently under revision by H. Ketchum (H. Ketchum, pers. comm.,

2006).


264

Pliosaurus Owen, 1841 (Owen, 1842)

Type material: P. brachydeirus OXFUM J.9245 A.B., “teeth, lower and upper jaws,

vertebral column, femur, tibia and fibula” (Tarlo, 1960, p. 152).

Type locality: Market Rasen, Lincolnshire, England.

Type horizon: Kimmeridge Clay, Kimmeridgian, Upper Jurassic.

Referred material: BRSMG Cc332 (***), P. brachyspondylus, the ‘Westbury Pliosaur’

(Taylor and Cruickshank, 1993). Almost complete skull and mandible, some

associated cervical vertebrae, phalanges and a rib.

Remarks: I have refrained from including both Kimmeridgian species of Pliosaurus

because few characters separate them, many of questionable validity (Taylor and

Cruickshank, 1993). However the type specimens of P. brachyspondylus and P.

brachydeirus are complementary and, combined, allow a greater degree of resolution

for the genus. Accordingly, I have coded Pliosaurus to genus level only, albeit

discounting characters of the Callovian species P. andrewsi, which is omitted from

the analysis (see table 1). The coding for Pliosaurus can therefore be regarded as

follows: P. brachydeirus + P. brachyspondylus – P. andrewsi. The genus Pliosaurus

was described by Owen (e.g. 1869) and Tarlo (e.g. 1960).

TMP 94.122.01

Type material: TMP 94.122.01, complete skeleton including the skull.

Type locality: Syncrude Canada Ltd. oil sand mine (Mildred Lake site), 30km

northeast of Fort McMurray, Athebasca Oil Sands Area of northeastern Alberta,

Canada.

Type horizon: Wabiskaw Member, Clearwater Formation, (Lower Albian, Lower

Cretaceous).

Remarks: TMP 94.122.01 represents a distinct but as yet un-named taxon

(Druckenmiller, 2006ab). A detailed unpublished description of this taxon is provided

by Druckenmiller (2006a).

Umoonasaurus demoscyllus Kear, Schroeder and Lee, 2006

Type material: AM F99374, quite complete opalised skeleton including the skull.

Type locality: Zorba Extension Opal Field, west of Coober Pedy, northern South

Australia.

Type horizon: Bulldog Shale (Maree Subgroup) (Lower Aptian-Lower Albian, Lower

Cretaceous).


265

Remarks: Umoonasaurus was briefly described and named by Kear et al. (2006).

This plesiosaur was formally known as ‘Eric’ the pliosaur.

Leptocleidus superstes Andrews, 1922a

Type material: BMNH R.4828 (***), incomplete skull.

Type locality: Berwick, Sussex, England.

Type horizon: Barremian, Lower Cretaceous

Remarks: The holotype of Leptocleidus superstes (also the holotype of the genus) is

a partial skull and vertebral column associated with a complete pectoral girdle and

partial limbs, from the Lower Cretaceous Upper Weald Clay (Barremian?) of Sussex,

UK. BMNH R4828 was described and figured by Andrews (1922a). Leptocleidus

superstes is omitted from the morphometric analysis because it is so far removed

stratigraphically form the other taxa included, and it has already been well

distinguished as a separate taxon based on discreet characteristics. Species referred

to this genus are also known from South Africa and Australia (see below), making

Leptocleidus one of the most widespread genera of plesiosaur.

Leptocleidus capensis (Andrews, 1911) Cruickshank, 1997.

Type material: SAM-K5822, skull.

Type locality: Zwartkops River Valley, Algoa Basin, South Africa.

Type horizon: Sundays River Formation, (Upper Valanginian, Lower Cretaceous)

Remarks: This specimen was originally described and named ‘Plesiosaurus

capensis’ by Andrews (1911). It was later redescribed and refigured by Cruickshank

(1997) who referred it the genus Leptocleidus.

Leptocleidus clemai Cruickshank and Long, 1997

Type Material: WAM 92.8.1-1 to 60, a partial postcranial skeleton.

Type locality: Kalbarri region, Western Australia.

Type horizon: Birdrong Sandstone (Hauterivian-Barremian, Early Cretaceous)

Remarks: Leptocleidus clemai is described by Cruickshank and Long (1997) and

represents one of three species of the genus Leptocleidus.

Dolichorhynchops osborni Williston, 1903

Type material: KUVP 1300, almost complete skeleton.

Type locality: Logan County, Kansas, USA.

Type horizon: Niobrara Formation, Cenomanian, Late Cretaceous

Referred material: MCZ 1064, FHSM VP404.


266

Remarks: Dolichorhynchops osborni has been thoroughly described by Williston

(1903), Carpenter (1996, 1997) and O’Keefe (2004a). This taxon is included as a

representative polycotylid taxon to test whether polycotylids are plesiosauroids (as

endorsed by Carpenter, 1997 and O’Keefe, 2001a) or pliosauroids (as endorsed by

Druckenmiller, 2006ab).


267

Appendix 3. Anatomical abbreviations

1-4 – distal carpals 1-4Ang – angularAr – articularBo – basioccipitalBs – basisphenoidC – coronoidCc – central cavityCo – coracoidD – dentaryEct – ectopterygoidEp – epipterygoidEx – external narisExop – exoccipital-opisthoticF – frontalFe – femurFi – fibulaH – humerusIc – interclavicleIn – intermediumInt – internal naresIs – ischiumi-v – metacarpals i-vJ – jugalM – maxillaMc – maxillary cavityOc – occipital condyleOr – orbitP – parietalPar – parasphenoidPal – palatinePin – pineal foramenPm – premaxillaPo – postorbitalPoc – paraoccipital processPof – postfrontalPre – prearticularPrf – prefrontalPt – pterygoidPu – pubis

Q – quadrateQp – quadrate-pterygoid flangeR – radiusRa – radialeRbs – ribsSa – surangularSc – scapulaSoc – supraoccipitalSp – splenialSq – squamosalSt – stapes (hyoid)T – tibiaTa – tooth alveoliTem – temporal fenestraU – ulnaUl – ulnareV – vomer


268

Appendix 4. Morphometric data (all measurements in cm)


269

R. cramptoni (NMING F8785)

R. thorntoni (BMNH R4853)

R. zetlandicus (YORYM G503)

R. propinquus (WM 851.S)

R. megacephalus (type cast, BRSMG Cb 2335 )

R. megacephalus (neotype, LEICS G221.1851)

NMING F10194

WARMS G10875

E. arcuatus (BMNH R2028* etc.)

NMING F8749

Archaeonectrus (BMNH 38525)

Macroplata (BMNH R5488)

P. macrocephalus (BMNH R1336)

BMNH 49202

HALB uncatalogued

Thalassiodracon (BMNH 2018*)

Attenborosaurus (BMNH R1339)

R. victor (SMNS 12478)

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92

73

59

63

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29.5

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43.5

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34

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36.5

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0.8

7?

Jove

rlap

: nar

es a

nd d

ors

o-m

edia

n f

ora

men

1.2

?1

_n/a

_n/a

??

n/a

n/a

n/a

?n/a

?n/a

?K

Snout

tip t

o m

axill

a (l

ater

al)

11

16.5

11.5

_11

13.8

13.6

??15

10

12.8

?6.8

?10

6.7

7L

pre

max

/max

illa

sutu

re t

o o

rbit

26

?23

_16.5

19.5

19

??

21.5

?18

?6.2

?15

6.9

3M

max

wid

th a

cross

pre

max

. ro

stru

m

14.5

16.8

12

_11

8.8

8.8

??11

6.5

8.5

?5.26.9

??

6.3

1N

length

of

exte

rnal

nar

es

54.5

43.5

2.7

63.0

73.2

??

2.1

??

1?

1.4

51.9

??1.85

OW

idth

of

exte

rnal

nar

es (

left

)2.9

1.6

1.9

1.9

1.1

21.3

71.1

??

1.7

??

?21.2

??

1.1

P(r

ight)

2.7

?2

1.7

1.1

21.2

51.5

??

??

?0.5

?n/a1.2

??

1.1

1Q

Wid

th b

etw

een e

xter

nal

nar

es5.9

?4.7

3.8

2.7

14.7

24

??

4?

??

2.6

8?

??

1.9

9R

Dis

tance

bet

wee

n e

xter

nal

nar

is a

nd o

rbit

5.3

65.1

3.1

4.2

34.3

3.7

??

4.6

?5.5

?1.1

5?

??

1.6

4S

Wid

th b

etw

een o

rbits

13.5

?8.5

7.1

?9

4.2

??

93.6

5.4

3.8

9.2

??

??

TO

rbit len

gth

(le

ft)

15

18

10.5

10

?9.8

13.5

??

12.6

5.5

7.5

3.9

6.5

??

7.7

?U

(rig

ht)

15.5

17.5

9.8

??12

14

??

?5.5

73.9

6.5

??

7.7

?


270

NMING F8785

BMNH R4853

YORYM G503

WM 851.S

BRSMG Cb 2335

LEICS G221.1851

NMING F10194

WARMS G10875

BMNH R2028* etc.

NMING F8749

BMNH 38525

BMNH R5488

BMNH R1336

BMNH 49202

HALB

BMNH 2018*

BMNH R1339

SMNS 12478

VO

rbit w

idth

(le

ft)

9.6

?8.5

7?7.5

12

??

11

67.5

?5.4

??

6.5

?W

(rig

ht)

15?

??

??

614

??

?5.4

7.2

2.3

5.1

??

6.5

?X

pre

max

illa

wid

th (

1/2

way

)7.2

?5.6

5_

5.7

95.8

6.5

??

??

8.3

?3.8

5?

??3.75

YSku

ll w

idth

at

pre

max

illa/

max

illa

sutu

re14

13.5

11

_9.5

58.1

18.6

??

10

5.2

7.5

?5.1

56.3

??

5.5

4Z

Sku

ll w

idth

at

ante

rior

bord

er o

f nar

es26.5

33

19

_17.5

16

16.5

??

18

12.5

15

?8.7

??

?11.6

7A*

Sku

ll w

idth

at

ante

rior

bord

er o

f orb

it32.5

35

24

_21

21

21

??

27

?20

?10.9

12

??

13.6

1B*

Sku

ll w

idth

at

mid

dle

of

orb

it37

38

?_

?24.5

26

??

32

18.5

23

?15.5

??

?16.5

C*

Bulb

to p

inea

l fo

ram

en30.5

?22.5

_?

17.7

18.5

??

??17

?9.4

8?

?14.5

?Palate

AA

Sku

ll tip t

o p

ost

erio

r in

terp

tery

goid

vac

uity

57

?53.5

?44.5

52

49

44.5

?55

?46.5

?22

??

?23.5

BB

length

of

ante

rior

inte

rpte

rygoid

vac

uity

n/a

?_

?6

8.6

8.2

7?

??

??

7?

??

5.5

9CC

length

of

post

erio

r in

terp

tery

goid

vac

uity

6.5

?6

?3.7

55.5

54.7

5?

5?

3.6

?3.4

??

?4.1

4D

Dw

idth

of

single

post

erio

r in

terp

tery

goid

vac

uity

2.7

?3

?1.8

22

1.9

?2.3

8?

2?

0.9

??

?1.1

5EE

wid

th b

etw

een inte

rnal

nar

es3.5

4.0

5?

??

22.4

2?

2.7

3?

??

1.2

51.5

??

2.0

7FF

max

imum

wid

th o

f par

asphen

oid

2.3

?1.7

??

1.6

2?

1.9

?3.3

?2.9

5?

??

0.6

3G

Gw

idth

bet

wee

n lat

eral

mar

gin

s of

post

. in

t. v

acs

6.5

?5.7

?5.7

55.5

5.2

?6.1

9?

5.5

?3.0

3?

??

2.6

8H

Hcu

ltrifo

rm p

roce

ss len

gth

2.5

??

??

1.8

3?

?2.5

?1.2

2?

3.5

8?

??

0M

an

dib

lea

Sym

phys

is len

gth

(ve

ntr

al)

13.5

14.5

12

?13

11.9

?15.5

9.9

11

16.5

13

?4.7

34

3.7

14

6.2

bM

andib

le len

gth

(al

ong r

amus)

110

11

499

?80.5

84

?78

?74

62

66

28

38.5

?18.5

53

41.5

cSym

phys

is w

idth

16

20

14

?12.9

10.9

?10.5

10.1

12.5

?9.2

?6.8

24.8

3.0

17.8

5d

Min

imum

wid

th b

ehin

d s

ymphys

is (

const

rict

ion)

15.5

18

11.9

?11.1

9.2

8?

98.2

510

?8

?6.2

5?

2.9

7.5

edis

tance

to c

onst

rict

ion

18

20.5

15.5

?18

16.5

?17

14.8

16

?14.9

?6.9

?3.8

811.5

8.7

5f

dors

al s

ymphys

is len

gth

(an

terior

of

gully

)?

9.3

_?

?_

??

5.2

5?

??

??

??

?gdia

gonal

len

gth

of

sym

phys

is (

tip -

const

rict

ion)

20

_18

?18.5

18.7

?16.5

13

16.5

?15.2

?7.1

5?

4.3

89.1

2p

ost

cran

ial

skele

ton

dim

en

sio

ns

Co

raco

id1

Cora

coid

len

gth

?58

50

30

??

46.5

45

45.5

??

50.3

??

?17

?39.5

2m

idlin

e sy

mphys

is len

gth

?39.4

?25

??

25

22

27.5

??45

??

?14

?32

3hum

erus

face

t (v

entr

al)

?15.5

?9.5

??

7.7

7.5

7.7

??

7.7

??

?2.5

?8.5

4sc

apula

r fa

cet

(len

gth

cora

coid

sca

pula

sutu

re)

?16

11.5

6.5

??

7.5

86.9

??

5?

??

2.0

5?

85

post

erio

r co

rner

of

gle

noid

to c

orn

ua

?25

??

??

23

22

22.5

??

_?

??

9.1

5?

16.5

6m

edia

n e

mbay

men

t le

ngth

?18

??

??

17

_17

??

5?

??

2.7

5?

6


271

NMING F8785

BMNH R4853

YORYM G503

WM 851.S

BRSMG Cb 2335

LEICS G221.1851

NMING F10194

WARMS G10875

BMNH R2028* etc.

NMING F8749

BMNH 38525

BMNH R5488

BMNH R1336

BMNH 49202

HALB

BMNH 2018*

BMNH R1339

SMNS 12478

7m

edia

n e

mbay

men

t bord

er len

gth

?15.5

??

??

20.5

__

??

_?

??

2.9

?_

8Ante

rior

pro

cess

len

gth

?14

13

??

?9

8_

??

10

??

?4.3

5?

89

Wid

th a

cross

gle

noid

?

33.5

28

18.5

??

21.2

21.3

20.7

??

20

??28

5.8

3?

20

10

max

wid

th (

post

erio

r bord

er o

f gle

noid

)?

38

?22

??

22.7

_22

??

25

??

?7.3

8?

22.5

11

Wid

th o

f gle

noid

pro

cess

28

?13

??

12.8

12

13.5

??

12

??

??

??

12

wid

th o

f pre

gle

noid

pro

cess

?18

?8?

??

10

10.5

_?

?10.8

??

?2.7

9?

9.7

13

Max

wid

th o

f post

erio

r em

bay

men

t (t

o m

idlin

e)?

3?

??

?10.5

56

??

_?

??

1.7

?6.5

14

max

post

erio

r c

ora

coid

corn

ua

(fro

m m

idlin

e)?

29.5

??

??

23.5

16

14.3

??

17.1

??

?4.0

9?

17

15

min

imum

wid

th o

f post

erio

r pro

cess

cora

coid

?

26.4

??

??

15.6

14

11.2

??

13.7

??20

3.1

5?

14

16

Wid

th o

f post

. pro

cess

cora

coid

?

28

??

??

15

_10

??

17.1

??

??

?12

17

length

to m

id-g

lenoid

?47

??

??

38

32?

_?

?_

??

?11.7

5?

29

18

fenes

tra

length

?13

12

??

?_

9.2

??

?_

??

?3.1

5?

619

fenes

tra

wid

th?11

10

??

?7

7.2

??

??

??

2.3

5?

3.5

20

Length

to f

enes

tra

?52.5

??

??

39.5

?37

??

??

13.0

7?

31

Scapula

21

Length

sca

pula

??

??

??

20

??

?13

??

??

7.3

5?

20

22

ant

pro

cess

wid

th?

??

??

?10.5

??

??

??

??

4.2

7?

11.4

23

min

wid

th (

ventr

al)

??

??

??

6?

??

??

??

?2.0

6?

6.6

24

max

wid

th (

post

pro

cess

ven

tral

)?

17.5

??

??

10

??

??

??

??

3.6

8?

10.2

25

dors

al p

roce

ss len

gth

?9

??

?14

??

??

??

??

??

??

26

dors

al p

roce

ss t

erm

inal

wid

th?

4?

??

??

??

??

??

??

??

?H

um

eru

s LEFT

27

Length

53

72

54

35

35

41.5

36.5

36.5

37.5

42

?34

10.4

?29

14.7

8?

40

28

Dis

tal flar

e25

37

25

15

18.2

22

19

19

19.3

2210

16

5.5

?13

7.3

3?

19

29

min

imum

wid

th12

15

12.5

8.5

?8

8.5

_?

7.2

8?

_?

??

3.4

8?

830

dis

tance

of

min

imum

wid

th f

rom

dis

tal tip

31

35

24

_21

23

_?

23.5

26

?_

??

?10.8

?25

31

pre

axia

l dis

tance

43

63

46

?30

38

34

?30

?_

9.2

??

12.9

?38

32

post

axia

l dis

tance

43

56

41

?27

36

30

?29.3

33

?_

8.1

??

10.9

5?

31

33

Rad

ius

face

t?

n/a

14

??

?_

15

_?

??

??

??

?_

34

uln

a fa

cet

?n/a

9?

??

_8

_?

??

??

??

?_

35

Length

to t

uber

osi

ty?

62

48

??

??

??

??

??

??

??

_H

um

eru

s R

IGH

T?

27

Length

51

??

??

42.5

35.5

?_

??

33?

10.4

?29

14.6

6?

40.5


272

NMING F8785

BMNH R4853

YORYM G503

WM 851.S

BRSMG Cb 2335

LEICS G221.1851

NMING F10194

WARMS G10875

BMNH R2028* etc.

NMING F8749

BMNH 38525

BMNH R5488

BMNH R1336

BMNH 49202

HALB

BMNH 2018*

BMNH R1339

SMNS 12478

28

Dis

tal flar

e25

?25

15.4

?20

19

??

??

16

5.5

?13

6.9

2?

20

29

min

imum

wid

th12

??

8.5

?8.5

??

??

?_

??

3.5

7?

8.5

30

dis

tance

of

min

imum

wid

th f

rom

dis

tal tip

25

??

??

23

??

??

?_

??

9.6

?_

31

pre

axia

l dis

tance

43

??

??37

32

??

31

?_

10

??

11.9

?38

32

post

axia

l dis

tance

_?

??

?33

29

?_

?_

8?

?11

?35.5

Fem

ur

LEFT

?36

Length

53.4

68

49

3634

39

35

36

35.4

?24

36.5

12.5

?29

14.1

?38.5

37

Flar

e25

29

22.5

1716

21

18

19

16.5

?11

16

6.1

??

5.3

?16

38

min

imum

wid

th11

13

8.8

57.5

?7.5

7.4

75.9

??

_?

??

2.7

5?

639

dis

tance

of

min

imum

wid

th f

rom

dis

tal tip

18

43

33

??

19

?21.4

??

_?

??

10.3

?25

40

pre

axia

l dis

tance

_55

41

??

33.5

?3

029.5

??

30

11.2

??

11.6

7?

33

41

post

axia

l dis

tance

_52

40

??

30.5

?2

425.1

??

27

10.2

??

11.6

7?

29.5

42

tibia

fac

et len

gth

(dors

al v

iew

)_

_16

??

_?

?_

??

11

??

??

_43

fibula

fac

et len

gth

(dors

al)

__

13

??

_?

?_

??

6.5

??

??

_Fem

ur

RIG

HT

36

Length

53.8

68.5

?36.5

?38

35

35.5

34.5

??

35

13

??

14.5

6?

37.5

37

Flar

e24.5

29

?15?

?21

18

19

16.8

??

_6.1

??

6.2

5?

16

38

min

imum

wid

th11

12

?7.5

?7.5

?7

5.8

??

_?

??

2.7

6?

639

dis

tance

of

min

imum

wid

th f

rom

dis

tal tip

25

33

??

?19

?_

23

??

_?

??

9.5

?25

40

pre

axia

l dis

tance

_57

??

?33.5

?30

30.3

??

30

11

??

11.6

9?

32

41

post

axia

l dis

tance

_52

??

?32

?26.5

24.2

??

27

10

??

10.5

?28.5

42

tibia

fac

et len

gth

(dors

al v

iew

)_

??

??

??

?_

??

11.2

??

??

?_

43

fibula

fac

et len

gth

(dors

al)

_?

??

??

??

_?

?6.5

??

??

?_

Rad

ius

44

length

16

?18.5

11

10.5

16.4

?14

13.5

?8

??

??

5?

13.9

545

pro

xim

al w

idth

11.4

?12.5

10

912

?12

10

?5

??

??

4.3

?9.5

546

Dis

tal w

idth

11.3

?12.5

97.1

9.5

?8

8.2

?3.9

??

??

2.8

5?

9.7

547

min

imum

wid

th8.8

?9.5

7.5

5.3

25.8

??

5.2

??

??

??

?6.9

9U

lna

??

??

48

Length

14.6

?15.1

?10.7

13

?14

14.4

?8

??

??

5.4

9?

12.2

849

wid

th (

mid

dle

)11

?11.2

?7.9

510.8

?10

9.3

?5.5

??

??

3.9

5?

10.1

2Tib

ia50

Length

18

??

10

?13.9

13

13

??

?11

??

?5

?11.6

5


273

NMING F8785

BMNH R4853

YORYM G503

WM 851.S

BRSMG Cb 2335

LEICS G221.1851

NMING F10194

WARMS G10875

BMNH R2028* etc.

NMING F8749

BMNH 38525

BMNH R5488

BMNH R1336

BMNH 49202

HALB

BMNH 2018*

BMNH R1339

SMNS 12478

51

pro

xim

al w

idth

15.8

??

9?

11.4

10.3

11

??

?9.8

??

?3.7

?9.5

52

dis

tal w

idth

15

??

8?

13.9

8.5

9?

??

8.5

??

?2.85

?10.9

Fib

ula

53

Length

16

??

??

7.9

11

12

11

??

13

??

?4.6

9?

12

54

wid

th (

mid

dle

)14

??

??

??

10

8.3

5?

?9.1

??

?3.0

8?

8.4

5P

ub

is55

Length

?38

??

??

29.8

27.2

25

??

29

??

?9.5

20

22

56

Wid

th?

43

??

?27

26

25.4

24

??

30.5

??

?9

23

24

57

Length

fen

stra

?13

??

?7.5

8.7

8.2

??

??

??

?_

??

58

Wid

th f

enes

tra

?12.5

??

?7.5

98.5

??

??

??

?_

?7.2

Isch

ium

59

Length

?39.8

??

?28

?29

26

??

31

??

?10.2

21

21

60

Wid

th?

35

??

?21.5

?22

23

??

24

??

?7.5

919.8

20

61

dis

tal w

idth

of

late

ral pro

cess

?19

??

?12

?7.5

??

??

??

?3.9

?62

min

wid

th o

f la

tera

l pro

cess

shaf

t?

16

??

?6.2

?5.3

7?

??

??

?2.4

56

?63

Isch

ium

gle

noid

fac

et?

??

??

6.4

?6.8

7?

?7

??

??

6.5

?Il

ium

64

Length

26

29

?18

?19

?19.5

18.5

?12.6

??

??

?19

17.5

65

dis

tal flar

e w

idth

12.5

14.5

?8

?8.9

??

8?

3.7

??

??

?7

6.5

66

pro

xim

al w

idth

10.5

13

?7

?8.2

?8.9

8?

5.6

??

??

??

8.4

567

min

imum

wid

th7

7.8

?3.5

?4.5

??

??

1.4

??

??

??

?O

ther

68

Dis

tance

bet

wee

n c

ora

coid

and p

ubis

??

??

??

?57

??

??

??

?14.5

?42.5

69

Tota

l le

ngth

675665

?440508

516

?450

?420335465

156

??

174

?335

Sele

cted

Rati

os

wid

th a

t ex

t nar

es/

length

to e

xt.

nar

es0.8

70.9

40.7

##

#0.7

40.5

90.6

1#

##

##

#0.6

10.4

80.6

##

#0.9

1#

##

##

##

##

##

1.0

03

orb

it len

gth

/sk

ull

length

0.1

70.1

90.1

40.1

7#

##

0.1

50.2

##

##

##

0.1

80.1

10.1

20.1

70.2

2#

##

##

##

0.1

6#

##

#Le

ngth

to e

xt.

nar

es/

skull

length

0.3

50.3

80.3

70.3

80.3

70.4

20.4

##

##

##

0.4

10.5

10.4

0.4

0.3

3#

##

##

##

0.4

38

0.3

32

orb

it len

gth

/ pre

max

to e

xt.

nar

es0.4

90.5

0.3

90.4

4#

##

0.3

60.5

##

##

##

0.4

30.2

10.3

10.4

30.6

8#

##

##

##

0.3

67

##

##

Length

pre

max

rost

rum

/ pre

max

wid

th0.7

60.9

80.9

6#

##

11.5

71.5

5#

##

##

#1.3

61.5

41.5

1#

##

1.3

10

##

##

##

##

1.0

73

rost

rum

wid

th/

pre

max

to e

xt.

nar

es0.4

80.4

80.4

4#

##

0.4

70.3

30.3

3#

##

##

#0.3

70.2

50.3

5#

##

0.5

4#

##

##

##

##

##

0.5

43

wid

th b

etw

een o

rbits/

tota

l pre

max

illa

length

0.3

1#

##

0.2

10.2

1#

##

0.2

50.1

1#

##

##

#0.2

3#

##

0.1

60.2

90.6

7#

##

##

##

##

##

##

##

length

to o

rbit m

argin

/ sk

ull

length

0.4

20.4

60.4

70.4

60.4

60.5

10.4

9#

##

##

#0.4

90.5

70.5

20.4

70.4

2#

##

##

##

0.5

31

0.3

97


274

NMING F8785

BMNH R4853

YORYM G503

WM 851.S

BRSMG Cb 2335

LEICS G221.1851

NMING F10194

WARMS G10875

BMNH R2028* etc.

NMING F8749

BMNH 38525

BMNH R5488

BMNH R1336

BMNH 49202

HALB

BMNH 2018*

BMNH R1339

SMNS 12478

pre

max

illa

length

/ sk

ull

length

0.4

9#

##

0.5

60.5

80.5

20.5

60.5

6#

##

##

#0.5

5#

##

0.5

70.5

90.4

7#

##

##

##

0.6

15

##

##

Rost

rum

wid

th/

length

to o

rbit

0.3

90.4

0.3

5#

##

0.3

80.2

70.2

7#

##

##

#0.3

10.2

20.2

7#

##

0.4

20.6

##

##

##

##

0.4

54

hum

erus

length

/sku

ll le

ngth

0.6

0.7

80.7

40.5

90.5

60.6

40.5

40.5

8#

##

0.5

8#

##

0.5

50.4

6#

##

##

#0.9

24

##

##

1.1

43

hum

erus

length

/ le

ngth

to p

ost

. Vac

s.0.9

3#

##

1.0

1#

##

0.7

90.8

0.7

40.8

2#

##

0.7

6#

##

0.7

2#

##

##

##

##

##

##

##

##

1.7

02

Cora

coid

wid

th a

t gle

noid

/ co

raco

id len

gth

##

#0.5

80.5

60.6

2#

##

##

#0.4

60.4

70.4

5#

##

##

#0.4

##

##

##

##

#0.3

43

##

##

0.5

06

Fem

ur

length

/ sk

ull

length

0.6

10.7

40.6

70.6

10.5

40.6

0.5

20.5

6#

##

##

#0.4

70.6

0.5

6#

##

##

#0.8

81

##

##

1.1

sym

phys

is len

gth

/ sy

mphys

is w

idth

0.8

40.7

30.8

6#

##

1.0

11.0

9#

##

1.4

80.9

80.8

8#

##

1.3

6#

##

0.6

90.8

31.2

29

##

##

0.7

9sy

mphys

is len

gth

/ m

andib

le len

gth

0.1

20.1

30.1

2#

##

0.1

60.1

4#

##

0.2

##

#0.1

50.2

70.1

9#

##

0.1

2#

##

0.2

0.2

64

0.1

49

flar

e hum

erus

/ le

ngth

hum

erus

0.4

70.5

10.4

60.4

30.5

20.5

30.5

20.5

20.5

10.5

2#

##

0.4

80.5

3#

##

0.4

50.4

96

##

##

0.4

75

post

axia

l hum

erus/

pre

axia

l hum

erus

10.8

90.8

9#

##

0.9

0.9

50.8

8#

##

0.9

8#

##

##

##

##

0.8

8#

##

##

#0.8

49

##

##

0.8

16

Length

hum

erus/

len

gth

fem

ur

0.9

91.0

61.1

0.9

71.0

31.0

61.0

41.0

31.0

6#

##

##

#0.9

20.8

3#

##

11.0

48

##

##

1.0

39

pre

axia

l hum

erus/

pre

axia

l fe

mur

##

#1.1

51.1

2#

##

##

#1.1

3#

##

##

#1.0

2#

##

##

##

##

0.8

2#

##

##

#1.1

05

##

##

1.1

52

Hum

erus

flar

e/ s

kull

length

0.2

80.4

0.3

40.2

50.2

90.3

40.2

80.3

##

#0.3

10.2

0.2

60.2

4#

##

##

#0.4

58

##

##

0.5

43

Bulb

-pin

eal fo

ram

en/

skull

length

0.3

5#

##

0.3

1#

##

##

#0.2

70.2

8#

##

##

##

##

##

#0.2

8#

##

0.3

2#

##

##

##

0.3

02

##

##

dis

tal w

idth

of

radiu

s/ r

adiu

s le

ngth

0.7

1#

##

0.6

80.8

20.6

80.5

8#

##

0.5

70.6

1#

##

0.4

9#

##

##

##

##

##

#0.5

7#

##

#0.6

99

radiu

s le

ngth

/tib

ia len

gth

0.6

3#

##

##

#0.9

##

#0.6

8#

##

0.6

2#

##

##

##

##

##

##

##

##

##

##

0.5

7#

##

#0.8

37

radiu

s le

ngth

/ hum

erus

length

0.3

##

#0.3

40.3

10.3

0.4

##

#0.3

80.3

6#

##

##

##

##

##

##

##

##

#0.3

38

##

##

0.3

49

length

of

post

. va

cs./

wid

th a

cross

post

. Vac

s1

##

#1.0

5#

##

0.6

61.1

10.8

50.9

6#

##

0.8

1#

##

0.6

5#

##

1.1

2#

##

##

##

##

##

1.5

45

length

to p

ost

vac

uitie

s/ s

kull

length

0.6

5#

##

0.7

3#

##

0.7

10.8

0.7

30.7

1#

##

0.7

6#

##

0.7

6#

##

0.7

5#

##

##

##

##

##

0.6

71

Orb

it len

gth

/iliu

m len

gth

0.5

80.6

##

#0.5

6#

##

0.5

2#

##

##

##

##

##

#0.4

4#

##

##

##

##

##

##

##

#0.4

05

##

##

skull

length

/ to

tal an

imal

len

gth

0.1

30.1

4#

##

0.1

30.1

20.1

3#

##

0.1

4#

##

0.1

70.1

50.1

30.1

4#

##

##

#0.0

92

##

##

0.1

04

Pubis

len

gth

/isc

hiu

m len

gth

##

#0.9

5#

##

##

##

##

##

##

##

0.9

40.9

6#

##

##

#0.9

4#

##

##

##

##

0.9

31

0.9

52

1.0

48

Pubis

len

gth

/cora

coid

len

gth

##

#0.6

6#

##

##

##

##

##

#0.6

40.6

0.5

5#

##

##

#0.5

8#

##

##

##

##

0.5

59

##

##

0.5

57

isch

ium

len

gth

/fem

ur

length

##

#0.5

9#

##

##

##

##

0.7

2#

##

0.8

20.7

3#

##

##

#0.8

5#

##

##

##

##

0.7

23

##

##

0.5

45

Pubis

wid

th/p

ubis

len

gth

##

#1.1

3#

##

##

##

##

##

#0.8

70.9

30.9

6#

##

##

#1.0

5#

##

##

##

##

0.9

47

1.1

51.0

91

Isch

ium

wid

th/i

schiu

m len

gth

##

#0.8

8#

##

##

##

##

0.7

7#

##

0.7

60.8

8#

##

##

#0.7

7#

##

##

##

##

0.7

44

0.9

43

0.9

52

Flar

e fe

mur/

length

fem

ur

0.4

70.4

30.4

60.4

70.4

80.5

40.5

10.5

40.4

7#

##

0.4

70.4

40.4

9#

##

##

#0.3

76

##

##

0.4

16

ilium

len

gth

/ fe

mur

length

0.4

90.4

3#

##

0.5

##

#0.4

9#

##

0.5

50.5

2#

##

0.5

2#

##

##

##

##

##

##

##

##

##

#0.4

55

Pubis

len

gth

/ co

raco

id w

idth

at

gle

noid

##

#0.6

6#

##

##

##

##

##

#0.6

40.6

0.5

5#

##

##

#0.5

8#

##

##

##

##

0.5

59

##

##

0.5

57

sym

phys

is len

gth

/hum

erus

length

0.2

50.2

0.2

2#

##

0.3

70.2

9#

##

0.4

20.2

60.2

6#

##

0.3

7#

##

##

#0.1

40.2

5#

##

#0.1

55

sym

phys

is len

gth

/rad

ius

length

0.8

4#

##

0.6

5#

##

1.2

40.7

3#

##

1.1

10.7

3#

##

2.0

6#

##

##

##

##

##

#0.7

4#

##

#0.4

44


275

Appendix 5. Data matrix for cladistic analysis


276

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

2728

2930

3132

3334

3536

3738

3940

4142

4344

4546

4748

49Ar

chae

onec

trus

00

1?

11

00

01

??

01

1?

10

?0

??

00

10

10

11

?1

0?

??

??

??

??

??

??

??

?At

tenb

oros

auru

s0

01

0?

?0

?0

?1

00

??

??

?1

00

?0

?1

?1

0?

0?

00

??

??

??

??

??

??

??

??

Augu

stas

auru

s0

00

00

10

00

01

00

10

00

00

00

00

00

01

01

1?

00

01

00

00

00

00

00

02

10

BMN

H 4

9202

01

00

10

00

01

?0

0?

00

01

10

00

01

00

20

11

00

00

0?

00

0?

?0

00

00

10

1Br

acha

uche

nius

00

10

00

02

01

10

0?

00

11

00

??

?0

10

10

00

?1

00

??

01

01

01

11

10

21

?C

ymat

osau

rus

00

00

00

00

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0x

x0

00

0x

?D

olic

horh

ynch

ops

10

10

00

00

00

01

01

11

10

11

20

00

11

20

01

01

10

20

10

11

01

01

01

10

1Eu

rycl

eidu

s?

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

?0

NM

ING

F10

194

00

00

11

01

01

10

11

10

00

?0

11

01

00

10

11

01

0?

10

00

11

10

0?

0?

2?

?N

IMIN

G F

8749

0?

00

11

01

01

1?

11

10

0?

10

1?

01

0?

1?

??

??

01

10

00

11

?0

01

01

20

?W

ARM

S G

1087

50

0?

0?

??

??

1?

??

??

??

??

??

??

??

??

??

??

??

?1

??

01

11

00

10

1?

0?

Hau

ffios

auru

s1

01

1?

??

??

??

??

??

??

??

??

??

??

??

??

??

??

01

00

00

00

10

10

01

1?

Hyd

rorio

n0

12

00

00

00

10

00

10

10

10

01

11

21

12

00

00

00

02

01

00

00

00

11

00

1?

Kron

osau

rus

00

10

00

??

01

1?

?0

??

?1

??

??

??

??

?0

??

01

??

??

??

??

11

?1

10

?0

0Le

ptoc

leid

us_c

apen

sis

01

00

10

00

?1

11

?1

??

00

00

00

00

11

21

00

?1

00

01

00

11

01

11

00

21

0Le

ptoc

leid

us_c

lem

ai?

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

??

Lept

ocle

idus

_sup

erst

es?

??

?1

0?

??

??

??

??

??

??

0?

??

00

11

10

00

?0

??

??

??

10

11

10

12

1?

Liop

leur

odon

00

10

00

02

01

10

00

01

11

10

11

20

10

10

00

00

00

11

01

01

01

11

10

01

0M

acro

plat

a0

00

11

10

10

11

?1

?1

01

??

0?

??

10

01

??

??

?0

??

??

??

??

00

10

11

0?

Mar

esau

rus

01

10

11

0?

11

1?

??

??

??

?0

??

??

1?

10

11

??

0?

11

00

0?

10

01

01

01

0O

UM

J285

850

?0

00

??

?0

10

0?

??

?0

??

?3

??

?1

0?

0?

?1

00

??

??

??

??

0?

0?

??

??

P.lo

ngiro

stris

10

11

11

00

01

0?

??

0?

??

?0

??

??

??

1?

01

?1

0?

10

00

??

?0

00

00

01

?P.

dolic

hode

irus

01

20

00

00

00

00

01

01

00

00

20

00

00

20

00

00

10

20

00

10

00

00

00

0?

1Pe

lone

uste

s0

01

10

00

20

11

00

10

00

11

00

12

01

01

00

1?

00

01

00

00

11

10

11

00

10

Pist

osau

rus

00

00

0?

00

00

00

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278

Appendix 1. List of valid plesiosaur species with information on...

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Transcript of Appendix 1. List of valid plesiosaur species with information on...