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Spoochelys ormondea gen. et sp. nov., an Archaic Meiolaniid-Like Turtle from the Early Cretaceous of Lightning Ridge, Australia

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Morphology and Evolution of Turtles

Part of the book series: Vertebrate Paleobiology and Paleoanthropology ((VERT))

Abstract

The Lower Cretaceous (lower to middle Albian) Griman Creek Formation deposits of Lightning Ridge in central-eastern Australia are famous for producing opalised fossils. Much of this material is poorly documented but recent assessments suggest a diverse assemblage of mainly non-marine vertebrates, invertebrates and plants. This biota is associated with a Gondwanan high-latitude zone that would have been subject to cool-temperate conditions. Turtle remains are particularly common at Lightning Ridge, comprising several distinct lineages including aquatic chelids and peculiar meiolaniid-like taxa—meiolaniids were spectacular horned turtles known from the Australian region and South America. Spoochelys ormondea gen. et sp. nov. shares some distinctive skeletal traits with this group (e.g., cranial scute pattern, incisura columellae auris confluent with the Eustachian tube) but also retains remarkably plesiomorphic cranial structures (e.g., an interpterygoid vacuity and short inferior parietal process) that are otherwise characteristic of Triassic and Jurassic stem turtles. The placement of meiolaniids and their sister lineages within Testudines is controversial in recent phylogenies. To test the relationships of Spoochelys, we used the two most comprehensive published data sets of fossil Testudinata and rescored a number of characters. Separate Maximum Parsimony and Bayesian analyses of both matrices supported recognition of Spoochelys as a primitive testudinatan but could not confirm its relationships with the meiolaniid clade. Nevertheless, the persistence of surprisingly archaic taxa such as Spoochelys into the Early Cretaceous of Australia implies survival of an ancient Pangean lineage, and brings into question long-held assumptions of Laurasian affinities for the meiolaniid-like turtles of Gondwana.

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Acknowledgments

Our sincere thanks to Don Brinkman for his endless patience and editorial help. Walter Joyce, Juliana Sterli and Igor Danilov provided comprehensive reviews. Henk Godthelp, Michael Archer, Suzanne Hand, Gene Gaffney, Tom Rich, Patricia Vickers-Rich, Lesley Kool, Karen Black, Anna Gillespie, Robin Beck, P. M. Datta, Asish Kumar Ray, Subhash Sen, Mike Lee, Arthur White, Robert Jones, Alex Ritchie, Robert Smith, Jenni Brammall, Dave Barclay, Brett and Peter Barton, Stefan Bedenikovic, Vicki Bokros, Rob and Debbie Brogan, Paul Burza, Peter Drackett, Jack Fahey, Dave Galman, Matthew Goodwin, Bill Kotru, Ormie and Donella Molyneux, Marcel and Sam Miltenburg, George and Bill Mulder, Dave Roussell and Lalja Pedersson, Graeme and Christine Thomson, Steve Turner, Joe Walker and Larry White provided discussions, preparation, access to specimens, photographs and graphical work. Financial support for this research was provided by The Royal Tyrrell Museum of Paleontology for ETS, and The Australian Research Council and Uppsala University for BPK.

Author information

Authors and Affiliations

  1. Vertebrate Palaeontology Laboratory, University of New South Wales, PO Box 1, Kensington, NSW, 2052, Australia

    Elizabeth T. Smith

  2. Australian Opal Centre, PO Box 229, Lightning Ridge, NSW, 2834, Australia

    Elizabeth T. Smith

  3. Palaeobiology Programme, Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36, Uppsala, Sweden

    Benjamin P. Kear

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  1. Elizabeth T. Smith
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  2. Benjamin P. Kear
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Correspondence to Elizabeth T. Smith .

Editor information

Editors and Affiliations

  1. Royal Tyrell Museum, Drumheller, Alberta, Canada

    Donald B. Brinkman

  2. Museum of Paleontology, University of California, Berkeley, California, USA

    Patricia A. Holroyd

  3. Royal Tyrrell Museum, Drumheller, Canada

    James D. Gardner

Appendices

Appendices

Appendix 1

Revised character scores (Nexus format) used for phylogenetic analyses. State codes for Spoochelys ormondea gen. et sp. nov. were based on specimens described herein. Modifications made for other taxa were drawn from published descriptions and direct examination of the following specimens: Chubutemys copelloi, MPEF-PV1236; Gaffney et al. (2007); Otwayemys cunicularius, NMV P186116; Gaffney et al. (1998); Meiolania platyceps, AM F208a, AM F208b, AM F401, AM F43183, AM F49141, AM F57984, AM F61110, AM F18314, AM F18364, AM F18668, AM F18775, AM F81940, AM F81949, AM F81965, AM F82180; Gaffney (1983, 1996); Joyce (2007); Ninjemys oweni, NHMUK R391 (cast); Gaffney (1992, 1996); Niolamia argentina, AMF L1418 (cast); Gaffney (1996); Sterli and de la Fuente (2011).

(A) Revised Matrix from Sterli and de la Fuente (2011)

Spoochelys_ormondea

???????1?0010001000000?1?????????012013011200??0??00001101?00?10?0??0011(01)010???????01???1??????0?0?????????????????01000011101????0?111?100????????????0

Chubutemys _copelloi

???100011001100?000?00?1?????1110?120211?12100000001??1101200?????????110?1?????0?????????????????????????????????????1?????01???0???11?1?11????????????

Otwayemys_cunicularius

???????????????????????????????????2?2?1????????????????????????????0???001?0???0?0?2?0?1101???0??20000000010001?1?00?1?????011??00?1?1?????????????????

Meiolania_platyceps

000000001001000100010101(02)00001010022003111211000(01)011102102200110(01)2?0001100100?10??00210?110101001020000?00000?????0000100111110(01)?0001111100101–01?10010

(B) Revised Matrix from Gaffney et al. (2007)

Spoochelys_ormondea

???????0?00010000010??0?????000?0000001111111111?00010?1?000110012?001000102???00??0??????1????10??????????????

Chubutemys _copelloi

?0??11?0??0010000???0001110?111101?01011111?1111???11011?21021?012?1????0??2???????????0?????2?????????????????

Otwayemys_cunicularius

???????????????00????0??????????????????1???1??????????????????012?00?0?0102???00???0?00?????21?01200010001??00

Meiolania_platyceps

0101110010001101110001011100111?01100111111101110111111102202?00110001000001110001100000110??2110110001000?????

Ninjemys_oweni

??0111?????01?0111??01011?????????????111111?111???1?1?1????????????????????11?????????????????????????????????

Niolamia_argentina

010?1100??001?0111??000111??111001?001111111?????1111111?2102???????????????1??????????????????????????????????

Appendix 2

Notes on descriptions and scores for select characters used in phylogenetic analyses.

(A) Characters from Sterli and de la Fuente (2011)

Character 8. [Prefrontal E] Prefrontals heavily sculptured with prominences and bosses (Rougier et al. 1995): (0) = present; (1) = absent. Some specimens of Meiolania (AM F18668) display a pronounced boss in the prefrontal region (Gaffney 1983, p. 405) and a similar structure is observable in Ninjemys (see Gaffney 1996, p. 79, Fig. 66). The scoring for Meiolania is therefore changed to “0”.

Character 12. [Parietal B] Parietal contact with the pterygoid, epipterygoid, or palatine: (0) absent, trigeminal foramen only developed as a notch; (1) present, trigeminal foramen clearly formed by the descending process of the parietal. Note that the parietal contacts the epipterygoid in Spoochelys (see Fig. 9.6b) and Meiolania (Gaffney 1983, p. 419) but is excluded form the trigeminal foramen by an epipterygoid-prootic contact.

Character 13. [Parietal C] Length of anterior extension of the lateral braincase wall: (0) short, inferior parietal process only produces a narrow strut anterior to trigeminal foramen, palatine contact absent; (1) elongate, inferior parietal process produces an extended process anterior to trigeminal foramen, palatine contact commonly present. Recoded from “−” to “0” for Meiolania following Joyce (2007).

Character 36. [Quadrate B + C of Joyce (2007)]. Development of the cavum tympani: (0) shallow but not developed anteroposteriorly; (1) shallow but anteroposteriorly developed; (2) deep and anteroposteriorly developed. Gaffney et al. (1998, p. 16) describe the cavum tympani of Otwayemys as “elongate, deep and smoothly continuous”; score changed to “2”.

Character 38. [Quadrate E] Antrum postoticum: (0) absent; (1) incipient; (2) fully developed. The antrum postoticum is commonly scored as present in Meiolania (Joyce 2007; Gaffney et al. 2007; Anquetin et al. 2009; Sterli 2008; Sterli and de la Fuente 2011); however (Gaffney 1983, pp. 416–417) reported that there was no discrete chamber corresponding to the antrum postoticum in this taxon (score changed to “0”), and indeed the squamosal and opisthotic are exposed posteriorly within the cavum tympani (see Gaffney 1983, pp. 408 and 443 respectively).

Character 41. [Epipterygoid A] Epipterygoid: (0) present, rod-like; (1) present, laminar; (2) absent. The epipterygoid of Meiolania lacks the free rod-like dorsal portion evident in Kayentachelys (see Sterli and Joyce 2007) and alternatively forms a broad sutural contact with the processus parietalis inferior (Gaffney 1983, Sterli and Joyce 2007). Because of this difference we advocate recoding of Meiolania from “0” to “1”.

Character 44. [Pterygoid C] Interpterygoid vacuity: (0) present; (1) absent. See discussion in Appendix 2, part B, character 29.

Character 51. [Pterygoid J] Intrapterygoid slit: (0) absent; (1) present. Gaffney (1983, 1996) and Gaffney et al. (2007) hypothesised that the ‘intrapterygoid slit’ is secondarily developed from the closed basicranial condition of eucryptodires. Sterli et al. (2010, 1344) alternatively suggested that this opening could be a modified interpterygoid vacuity. This interpretation requires confirmation via comprehensive revision of all meiolaniid taxa but would create interdependence with character 44 (presence of the interpterygoid vacuity).

Character 66. [Canalis caroticum B] Foramen caroticum laterale: (0) the arteria palatina enters the skull through the interpterygoid vacuity; (1) foramen caroticum laterale located between the basisphenoid and the pterygoid with the foramen posterius canalis carotici interni formed by the basisphenoid; (2) the canalis carotici interni is not completely floored allowing to see the foramen caroticum laterale and the foramen basisphenoidale in ventral view, foramen posterius canalis carotici interni is formed by the pterygoid or the pterygoid and the basisphenoid; (3) the arteria palatina branches from the arteria carotici interna inside the skull, so the foramen caroticum laterale is formed by the pterygoid as the foramen posterius canalis carotici interni; (4) the arteria palatina branches from the arteria carotici interna inside the skull, foramen posterius canalis carotici interni is formed by the pterygoid and the prootic; (5) foramen caroticum laterale absent; (6) foramen caroticum laterale formed within the pterygoid, posterior canalis carotici interni formed midway between pterygoid and basisphenoid. Gaffney and Jenkins (2010, pp. 335–336) specified terminology for the various basipterygoid foramina, indicating that the foramen caroticum basisphenoidale are those openings that pierce the body of the basisphenoid. In contrast, Joyce (2007) and the later character sets of Sterli (2008) and Sterli and de la Fuente (2011) use the term foramen posterius canalis carotici interni (which should be in part formed by the pterygoid: sensu Gaffney and Jenkins 2010, p. 336) to define the entry point of the internal carotid artery. We follow Gaffney and Jenkins (2010) in describing the paired foramina in the basisphenoid of Spoochelys as the caroticum basisphenoidale, and note that the presence of an interpterygoid vacuity together with the lack of any accessory foramina along the basisphenoid-pterygoid suture implies state “0”—entry of the arteria palatina into the skull via the interpterygoid vacuity.

Character 73. [Carapace A] Carapacial scutes: (0) present; (1) partially present; (2) absent. The carapace of Spoochelys is fragmentary but scute sulci are present on all of the preserved sections. Because the full extent of the scute coverage is unknown we have scored this taxon as “0/1”.

Character 81. [Costal A] Medial contact of costal I: (0) absent; (1) present. Disposition of the intercostal sutures is unknown in Meiolania (Gaffney 1996, p. 24) thus scoring is changed from “0” to “?”.

Character 82. [Costal B] Medial contact of posterior costals: (0) absent; (1) medial contact of up to three posterior costals present; (2) medial contact of all costals present. Disposition of the intercostal sutures is unknown in Meiolania (Gaffney 1996, p. 24) thus scoring is changed from “0” to “?”.

Character 99. [Mesoplastron A] Number and size of mesoplastra: (0) one or two pairs of mesoplastra present that fully hinder any contact between hyoplastra and hypoplastra; (1) one reduced pair of mesoplastra present that allows partial contact between the hyoplastra and hypoplastra; (2) mesoplastra absent. Mesoplastra are generally thought to be absent in Meiolania (recoded “2”), which has a wide transverse contact between the hypo- and hyoplastra (Megirian 1992, p. 102; Gaffney 1996, p. 38).

Character 104. [Plastral scutes B] Midline sulcus: (0) straight; (1) distinctly sinuous. Meiolania is recoded from “0” to “?” because scute margins are unclear (Gaffney et al. 1996, p. 33).

Character 108. [Extragular C] Anterior plastral tuberosities: (0) present; (1) absent. Gaffney (1996, p. 33) described a pair of gular projections on the anterior lobe of the plastron in Meiolania and noted their smaller size but overall similarity to those of Proganochelys (score changed to “0”). Coding is altered from “0” to “1” for Otwayemys, which possesses extragular scales but apparently not plastral tuberosities (Gaffney et al. 1998, p. 14, Fig. 10).

Character 110. [Humeral A] Plastral scale set 3—humerals: (0) one pair present; (1) two pairs present, subdivided by plastral hinge. Meiolania is recoded from “0” to “?” because scute margins are unclear (Gaffney et al. 1996, p. 33).

Character 114. [Inframarginal A] Inframarginal scutes: (0) more than two pairs present, plastral scutes do not contact marginals; (1) two pairs present (axillaries and inguinals), limited contact between plastral scutes and marginals present; (2) absent, unrestricted contact between plastral scutes and marginals present. Gaffney et al. (1998, p. 15, Fig. 11) reconstructed a series of axillary, inguinal, and one set of inframarginals scutes in Otwayemys (score changed from “?” to “1”).

Character 119. [Cervical articulation A] Cervical articulation: (0) articulations not formed; (1) articulations formed. Gaffney et al. (2007, p. 21) reported that at least cervical eight in Chubutemys has fully formed articulations (score changed to “1”).

Character 125. [Cervical articulation G] Cervical articulation between VII and VIII: (0) 7(8; (1) 7)8. Gaffney et al. (2007, p. 21) stated that cervical eight in Chubutemys has biconvex articulations (score changed to “0”).

Character 128. [Dorsal rib B] Contact of dorsal ribs IX and X with costals: (0) present; (1) absent. Gaffney (1996, pp. 25–26) considered it likely that the tenth dorsal rib did not articulate with a costal plate in Meiolania. We have therefore recoded this taxon as “0/1” to acknowledge manifestation of an intermediary condition.

Character 148. [Manus A] Phalangeal formula of manus and pes: (0) most digits with three elongate phalanges; (1) most digits with two shortened phalanges. Meiolania has only two short phalanges in the manus and pes (see Gaffney 1996, p. 50, Figs. 39 and 60); character recoded “1”.

Character 151. [Pes A] Claw of fifth digit: (0) present; (1) absent. Gaffney (1996, p. 60) suggested that the fifth pedal digit was absent in Meiolania because metatarsal V lacks a distal articular surface (recoded “1”).

(B) Characters from Gaffney et al. (2007)

Character 3. [NA] Nares divided: (0) yes; (1) no. Nares are divided by the premaxilla in Niolamia following Sterli and de la Fuente (2011) thus the score is changed from “1” to “0”. Structure of the narial aperture is indeterminate in Chubutemys, which is recoded “?”. The nares are clearly divided in Meiolania, although not completely in all specimens (Gaffney 1983, p. 401), scores are altered from “0/1” to “0”.

Character 4. [NA] Nasomaxillary sinus: (0) absent; (1) present. Proganochelys and meiolaniids share a trough-like cavity in the anterodorsal surface of the maxilla. In Proganochelys this extends through the lacrimal bone as the lacrimal duct, which is floored by the maxilla (Gaffney 1990, p. 49, Fig. 40). Conversely in Meiolania, the maxillary trough is called the “nasomaxillary sinus” and leads into a blind canal roofed by the nasal (Gaffney 1983, p. 429, Fig. 45). The anterior section of the “nasomaxillary sinus” in Meiolania and Ninjemys could represent a component of the primitive lacrimal duct. However, we have not changed the state coding because homology is unclear.

Character 10. [PA] Processus inferior parietalis: (0) small to absent; (1) large. The processus inferior parietalis is weakly developed and the epipterygoid extends further into the lower temporal fossa than the relatively small descending process of the parietal in Spoochelys and Meiolania (see Gaffney 1983, pp. 419 and 431, Fig. 47). The processus inferior parietalis is also described as “small” in Kayentachelys (Gaffney and Jenkins 2010, p. 339). Score changed from “1” to “0” for Meiolania.

Character 18. [SQ] Broad sq-qj contact: (0) absent, narrow; (1) broad contact below cavum tympani. Gaffney et al. (2007) assigned Niolamia state “1” despite much of this part of the skull being morphologically inconclusive (Woodward 1901; Gaffney 1996). The sq-qj contact below the cavum tympani is only definitively known in Meiolania and is inferred for Ninjemys and Warkalania on the basis of a ‘bony sheet’ below the cavum tympani (Gaffney 1996).

Character 20. [PO] PO-SQ contact: (0) present; (1) absent. Scoring for Chubutemys is modified from “0” to “?” because this contact is not clearly preserved (Gaffney et al. 2007, p. 15).

Character 26. [PAL] Teeth: (0) present; (1) absent. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 27. [PAL] Foramen palatinum posterius large: (0) small to moderate; (1) very large. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 28. [PT] Vertical flange on processus pterygoideus externus: (0) absent; (1) present. Joyce (2007) and Sterli and Joyce (2007) interpreted this feature as highly variable and of limited phylogenetic significance. In contrast, Gaffney (1979, 1996), Gaffney and Meylan (1988), Gaffney et al. (1987, 1991), and Gaffney and Jenkins (2010) advocate its use as a cryptodiran synapomorphy. In Meiolania “the vertical flange on the lateral margin of the processus … is absent” but a “swelling” of the bone is hypothesised to mark its original position (Gaffney 1983, p. 427). Because this does not strictly constitute a “vertical flange” we have changed the score from “1” to “0” for Meiolania and “1” to “?” for both Ninjemys and Niolamia since this part of the skull is not adequately preserved in either taxon (see Gaffney 1983, p. 431; Gaffney 1996, p. 77; Gaffney 1992, p. 3, Fig. 1).

Character 29. [PT] Interpterygoid vacuity: (0) widely, slightly open; (1) completely closed by medial extension of PT. Gaffney (1983, 1996) and Gaffney et al. (2007) argued that the “intrapterygoid slit” of meiolaniids developed as a neomorph from the primitive eucryptodiran condition in which the interpterygoid vacuity had completely closed. Dracochelys bicuspis Gaffney and Ye 1992 was forwarded as evidence of a intermediary state (Gaffney 1996, p. 119) where the basipterygoid-pterygoid suture is separated laterally to create paired slit-like openings for the palatine arteries (Gaffney and Ye 1992, p. 9). However, maintenance of at least a transverse midline contact between the pterygoid lamellae and rostrum basisphenoidale is a fundamental feature of the eucryptodiran basicranium (Gaffney 1996; Gaffney et al. 2007). Modification of the eucryptodiran state into the “intrapterygoid slit” arrangement of meiolaniids requires retraction of the pterygoid lamellae from the basisphenoid rostrum, and ventral displacement of the posterior pterygoids and canalis cavernosus, creating a condition similar to basal turtles (e.g., Proganochelys, Palaeochersis and Kayentachelys: Sterli et al. 2010, p. 1341). A different interpretation based on Sterli et al. (2010, p. 1344) and the palatal morphology of Spoochelys could involve formation of the “intrapterygoid slit” via progressive posterior closure of the pterygoid midline suture. Gaffney et al. (2007) described the pterygoid-basisphenoid contact in Chubutemys as being “damaged by crushing” but apparently manifesting lateral expansions of the suture to form paired foramina. Ninjemys is altered from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 30. [PT] Canalis caroticus internus: posterior to foramen basisphenoidale: (0) absent; (1) at least partially covered by BS-PT; (2) covered by prootic. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 31. [PT] Canalis caroticus internus at least partially formed by PT: (0) no; (1) yes, distal to foramen basisphenoidale (= foramen carotici cerebralis posterius sensu Sterli et al. 2010). Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 32. [PT] Canalis caroticus lateralis vs canalis caroticus internus: (0) lateralis equal to or larger than internus; (1) lateralis smaller than internus. Coding for Meiolania should be “1” because the “intrapterygoid slit” (= foramen caroticus laterale with terminology sensu Sterli et al. 2010) is considerably larger than the canalis caroticus internus (see discussion in Gaffney 1983, pp. 447–449, and 436, Fig. 53). However, Gaffney et al. (2007) recorded Meiolania as “?”, presumably because of uncertainty surrounding the identification and ancestral dimensions of the canalis caroticus lateralis. Indeed, homology between these structures is unclear. For example, the slit-like foramen caroticus laterale of primitive eucryptodires pierce the transverse pterygoid-rostrum basisphenoidale suture and the paired arteria palatina are recessed (Brinkman and Nicholls 1993; Sukhanov 2000; Sterli et al. 2010; Brinkman et al. 2012). In contrast, the rostrum basisphenoidale does not suture to the pterygoids in Meiolania and the passages for the palatine arteries were presumably walled medially by the sagittal ridge from the basisphenoid (see Gaffney 1983, p. 430).

Character 33. [PT] Foramen posterius canalis carotici interni: (0) not formed by basisphenoid and pterygoid; (1) formed by basisphenoid and pterygoid, located midway along basisphenoid. Ninjemys is recoded from “1” to “?” because the basicranium is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 34. [PT] Middle ear with ossified floor formed by posteromedial pterygoid process: (0) process absent; (1) present. Ninjemys is recoded from “1” to “?” because the basicranium is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 36. [PT] Deep cavity on quadrate ramus: (0) absent; (1) present. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 37. [PT] BS-PT space: (0) interpterygoid vacuity widely open; (1) distinct space, perhaps a foramen caroticus laterale (fcl), present; (2) closed, pt-bs suture, foramen posterius canalis carotici internus (fpcci) possibly present. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1). Niolamia and Meiolania changed from from “1” to “0” (see discussion for characters 29 and 32).

Character 38. [PT] Intrapterygoid slit: (0) absent, small; (1) extensive, covering foramen carotici cerebralis posterior (fccp). See discussion for characters 29 and 32. Ninjemys is recoded from “1” to “?” because the palate is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 45. [QU] Antrum postoticum: (0) absent; (1) present. Ninjemys is recoded from “1” to “?” because the cavum tympani is incomplete (Gaffney 1992, p. 3, Fig. 1). Meiolania is changed “1” to “0” because the antrum postoticum is not developed (Gaffney 1983, pp. 416–417); Niolamia “1” to “?” following Sterli and de la Fuente (2011).

Character 46. [QU] Covers opisthotic laterally: (0) no; (1) yes. Ninjemys is recoded from “1” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 47. [QU] Cranioquadrate space: (0) relatively open; (1) a well-defined canal. Ninjemys is recoded from “1” to “?” because the basicranium is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 48 [OP] Processus paroccipitalis: (0) loosely articulated; (1) tightly fused. Ninjemys is recoded from “1” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 49. [OP] Fenestra perilymphatica: (0) relatively large; (1) relatively small. Ninjemys is recoded from “0” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 50. [PR] Anterior surface otic chamber: (0) faces more anterodorsally; (1) faces more anteriorly, anteroventrally (processus trochlearis oticum). Ninjemys is recoded from “1” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 51. [PR] Otic chamber thickness: (0) anterior wall thin; (1) anterior wall thick. Ninjemys is recoded from “1” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 53. [SO] Wide occipital plate: (0) wide occipital plate with depressions; (1) absent. Ninjemys is recoded from “1” to “?” because the otic chamber is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 56. [BS] Basipterygoid articulation: (0) open articulation; (1) sutured joint. Ninjemys is recoded from “1” to “?” because the braincase is incomplete (Gaffney 1992, p. 3, Fig. 1).

Character 70. [CE] 4th cervical articulation: (0) amphicoelous; (1) biconvex; (2) opisthocoelous. Gaffney et al. (1998) suggested that the fourth cervical articulation of Otwayemys was probably procoelous based on tentative identification of a biconcave fifth (?) cervical centrum. This has been altered to “?” because of ambiguity.

Character 74. [CE] Strong ventral process: (0) absent on all centra; (1) present on posterior centra. Otwayemys is recoded from “0” to “1” because the eighth cervical vertebra (NMVP187261) is keeled (Gaffney et al. 1998, p. 20).

Character 92. [CAR] 9th costal bone: (0) present; (1) absent. The exact number of costals bones is unknown in Meiolania (Gaffney 1996, p. 24) thus scoring is changed to “?”.

Character 106. [PLA] Gular scales: (0) 2 pairs; (1) 1 pair. Otwayemys is recoded from “?” to “0” based on Gaffney et al. (1998, p. 12).

Character 107. [PL] Inframarginals: (0) 3–4 pairs; (1) 0–2 pairs (axillary, inguinal). Otwayemys is recoded from “?” to “1” based on Gaffney et al. (1998, p. 13).

Character 108. [PLA] Abdominal scute relative to midline: (0) meets on midline; (1) withdrawn from midline. Medial contact between the abdominal scutes cannot be reconstructed in Meiolania (Gaffney 1996, p. 33) or Otwayemys Gaffney et al. (1998, p. 13) thus states for both taxa are scored “?”.

Character 109. [PLA] Pectoral scute: (0) not on epiplastron; (1) on epiplastron. Meiolania is recoded from “0” to “?” because scute margins are unclear (Gaffney et al. 1996, p. 33).

Character 111. [PLA] Midline sulcus sinuous: (0) no; (1) yes. Meiolania is recoded from “0” to “?” because scute margins are unclear (Gaffney et al. 1996, p. 33).

Appendix 3

Inventory of “Pruned” taxa and their missing data (percentage).

(A) Taxa from Sterli and de la Fuente (2011)

  • Proterochersis robusta 73%

  • Australochelys africanus 75%

  • Condorchelys antiqua 65.8%

  • Indochelys spatulata 86.2%

  • Siamochelys peninsularis 73.7%

  • Platychelys oberndorferi 63.2%

  • Caribemys oxfordiensis 74.3%

  • Notoemys laticentralis 53.3

  • Dorsetochelys delairi 60.5%

  • Portlandemys mcdowelli 59.9%

  • Mesodermochelys undulatus 62.5%

  • Hoplochelys crassa 65.8%

  • Peltochelys durlstonensis 71.7%

  • Sandownia harrisi 53.9%

  • Chubutemys copelloi 60.5%

  • Otwayemys cunicularis 72%

  • Patagoniaemys gasparinae 73%

  • Niolamia argentina 79.6%

(B) Taxa from Gaffney et al. (2007)

  • Australochelys africanus 59.5%

  • Proterochersis robusta 79.3%

  • Otwayemys cunicularis 68.5%

  • Chubutemys copelloi 51.4%

  • Ninjemys oweni 72.9%

  • Niolamia argentina 57.7%

Appendix 4

Anatomical abbreviations

ast:

Aditus canalis stapedio-temporalis

bo:

Basioccipital

bs:

Basisphenoid

c:

Costal

ca:

Columella auris

caj:

Cavum acustico jugulare

cav:

Lab cavum labyrinthicum

cc:

Canalis cavernosus

ce:

Cervical scute

epi:

Epipterygoid

ex:

Exoccipital

fb:

Foramen basisphenoidale

fio:

Foramen interorbitale

fr:

Frontal

fst:

Foramen stapedio-temporale

ivac:

Interpterygoid vacuity

j:

Jugal

m:

Marginal scute

max:

Maxilla

n:

Neural

nu:

Nuchal

op:

Opisthotic

or:

Orbit

pa:

Parietal

pe:

Peripheral

pf:

Prefrontal

pio:

Processus interfenestralis

pip:

Processus inferior parietalis

pl:

Pleural scute

po:

Postorbital

pr:

Prootic

pt:

Pterygoid

qu:

Quadrate

rst:

Recessus scalae tympani

sm:

Supramarginal scute

so:

Supraoccipital

sq:

Squamosal

v:

Vertebral scute

V:

Foramen nervi trigemini

VII?:

Foramen nervi facialis?

XII:

Foramen nervi hypoglossi

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Smith, E.T., Kear, B.P. (2013). Spoochelys ormondea gen. et sp. nov., an Archaic Meiolaniid-Like Turtle from the Early Cretaceous of Lightning Ridge, Australia. In: Brinkman, D., Holroyd, P., Gardner, J. (eds) Morphology and Evolution of Turtles. Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-4309-0_9

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