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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
Appendices Appendix 1
258
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
Appendices Appendix 2
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).
Appendices Appendix 2
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
Appendices Appendix 2
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.
Appendices Appendix 2
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.
Appendices Appendix 2
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).
Appendices Appendix 2
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).
Appendices Appendix 2
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.
Appendices Appendix 2
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).
Appendices Appendix 2
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
Appendices Appendix 3
268
Appendix 4. Morphometric data (all measurements in cm)
Appendices Appendix 4
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)
Sku
ll
ALe
ngth
(to
post
erio
r sq
uam
osa
l bulb
)88
92
73
59
63
65
67
63
?72
51.2
61
22.5
29.5
?16
48
35
BLe
ngth
of
pre
max
illa
43.5
?41
34
33
36.5
37.4
??
39.5
?34.5
13.2
13.8
?29.5
?C
pre
max
illa
tip t
o e
xter
nal
nar
es30.5
35
27
22.5
23.5
27
27
??30
26
25
99.6
?21
11.6
3D
pre
max
illa
tip t
o p
inea
l fo
ram
en53.5
?47
37.5
?45.5
45
??
47
37.5
41
?19.4
?33
?E
pre
max
illa
tip t
o a
nte
rior-
most
bord
er o
rbit
37
42
34.5
27
29
33
32.5
??
35.5
29
32
10.5
12.4
12
?25.5
13.9
1F
Prem
axill
ae w
idth
at
ante
rior
bord
er o
f nar
es7
45.7
5.3
3.2
54.2
3.9
??
4?4.9
1.5
2.9
??
3.5
5G
pre
max
illa
tip t
o d
ors
o-m
edia
n f
ora
men
34
?29.2
24.5
n/a
_n/a
??
??
n/a
?n/a
??
?H
length
of
dors
om
edia
n f
ora
men
4?
3.5
5n/a
_n/a
??
n/a
?n/a
?n/a
??
?I
Length
of
pin
eal fo
ram
en4.8
?3.5
3.9
?2.1
23.5
??
??
??
1.4
7?
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
?
Appendices Appendix 4
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
Appendices Appendix 4
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
Appendices Appendix 4
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
Appendices Appendix 4
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
Appendices Appendix 4
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
Appendices Appendix 4
275
Appendix 5. Data matrix for cladistic analysis
Appendices Appendix 5
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
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10
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1Pe
lone
uste
s0
01
10
00
20
11
00
10
00
11
00
12
01
01
00
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00
01
00
00
11
10
11
00
10
Pist
osau
rus
00
00
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01
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00
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?Pl
iosa
urus
01
10
00
02
01
10
00
10
11
?0
31
21
10
20
01
?0
00
11
00
11
11
11
10
01
0R
.cra
mpt
oni
01
00
10
11
11
1?
01
10
10
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??
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11
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11
00
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10
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00
LEIC
S G
221.
1851
00
00
11
01
01
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11
10
00
?0
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01
00
10
11
01
01
10
00
11
10
01
01
20
0R
.pro
pinq
uus
01
00
10
11
11
??
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10
1?
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10
10
11
??
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??
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?R
.thor
nton
i0
10
01
01
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11
??
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R.v
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r0
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01
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1?
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10
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10
01
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11
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Seel
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urus
01
20
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31
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?Si
mol
este
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10
00
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Sthe
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Thal
assi
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01
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P_94
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10
02
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moo
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01
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11
00
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00
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00
01
00
0?
Appendices Appendix 5
277
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
7879
8081
8283
8485
8687
8889
9091
9293
Arch
aeon
ectru
s1
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00
11
10
11
11
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20
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gust
asau
rus
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00
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10
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BMN
H 4
9202
10
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14
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20
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10
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10
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ymat
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Dol
icho
rhyn
chop
s1
11
04
00
01
10
10
11
11
0?
01
11
10
11
02
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10
01
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11
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10
2Eu
rycl
eidu
s?
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11
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10
1?
1?
11
10
10
11
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12
00
11
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11
11
0N
MIN
G F
1019
41
10
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11
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11
11
00
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11
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G F
8749
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1?
01
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11
01
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WAR
MS
G10
875
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11
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11
1?
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11
10
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10
10
01
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10
Hau
ffios
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s1
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00
01
10
12
10
01
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01
10
10
21
11
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ydro
rion
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20
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10
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11
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30
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Mac
ropl
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Mar
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rus
11
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10
01
11
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MJ2
8585
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P.lo
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11
01
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P.do
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11
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10
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lone
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10
13
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11
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10
11
01
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stos
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10
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40
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11
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R.c
ram
pton
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10
12
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10
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11
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01
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ICS
G22
1.18
511
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11
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10
11
11
11
10
1?
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??
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?1
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00
10
1?
11
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.pro
pinq
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01
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11
??
10
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??
01
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00
01
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R.th
ornt
oni
??
01
2?
11
01
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11
10
11
1?
01
?2
01
10
00
00
01
01
02
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R.v
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r1
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12
01
1?
11
11
11
1?
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??
00
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10
01
10
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00
00
10
1?
11
1R
.zet
land
icus
11
01
21
11
?1
11
11
11
10
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??
01
1?
01
1?
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??
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01
11
Seel
eysa
urus
0?
10
3?
01
20
00
20
00
00
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10
01
01
10
01
00
10
00
01
01
01
00
Sim
oles
tes
1?
01
20
10
01
11
3?
??
00
10
01
01
02
10
10
00
01
10
0?
12
01
02
Sthe
naro
saur
us?
??
??
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??
??
1?
01
11
10
00
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1?
10
11
00
00
00
00
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1?
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alas
siod
raco
n0
11
00
01
10
10
02
00
00
00
01
00
10
11
12
10
?1
01
00
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10
01
0TM
P_94
_122
_01
1?
01
00
01
01
01
01
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01
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10
11
0?
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0?
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01
11
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02
Um
oona
saur
us1
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1?
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1?
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01
?1
10
??
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1?
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10
1?
10
2Yu
ngui
saur
us0
01
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10
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01
00
00
0
Appendices Appendix 5
278
Recommended