identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A287B16B3A5565F231F903FE6CF827.text	03A287B16B3A5565F231F903FE6CF827.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lisowicia bojani Sulej & Niedźwiedzki 2019	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Reconstruction of the  Lisowicia bojani skeleton </p>
            <p>Skull</p>
            <p> The reconstruction of the skull of  Lioosicia bojani (Fig. 47) is based on bones coming from a few individuals. They represent parts of the skull roof (ZPAL V. 33/MB/18), braincase (ZPAL V. 33/531), and single bones (maxilla ZPAL V. 33/85, postorbital ZPAL V. 33/708, parietal ZPAL V. 33/741, stapes) or their fragments from different individuals: the premaxilla, paroccipital. </p>
            <p> The general proportion of the skull was taken from the new reconstruction of  P. ‘ gigao ’. </p>
            <p> Cranium: The relationship of the frontal to postorbital is clearly visible in the specimen ZPAL V. 33/MB/18. A parietal foramen forms a canal the shape of which can be recognized in this specimen and in the whole parietal (specimen ZPAL V. 33/741). The dorsal edge of the frontal and parietal are visible in lateral view, the latter only slightly raised. The positions of the lacrimal and maxilla are based on  Jachaleria candelarienoio ,  R. criotatuo , and  P. ‘ gigao ’. The setting of the squamosal was reconstructed according to morphology of this element in  P. ‘ gigao ’. </p>
            <p>Mandible: The posterior part of the mandible is preserved in two specimens, ZPAL V. 33/735 and ZPAL V. 33/736. They consist of the articular, surangular, and prearticular. The angulars are also preserved, but always as isolated elements. The dentary and splenial are missing. The reconstruction of the posterior part of the mandible is based on general proportions in other Triassic dicynodonts and on the fit with known elements. The shape of the elongated dentary was based on an extremely elongated bone in Woznikella triradiata (length/deep— 16.6 cm × 6.2 cm = 2.67, probably the ancestor of Lioosicia in Europe).</p>
            <p>Postcranial skeleton</p>
            <p> The limb postures of kannemeyeriid dicynodonts were studied by Walter (1986) and Fröbisch (2006). Most of the material of Middle and Late Triassic dicynodonts consists of disarticulated skeletons or their parts. Rare articulated skeletons represent different groups of Triassic dicynodonts. These are: Shanoiodon sangi Yeh, 1959, Shanoiodon suhoiangenoio Yeh, 1959, Tetragoniao njaliluo von Huene, 1942, and Angonioauruo cruickohanki among Shansiodontini (Cox 1965) and  Kannemeyeria oimocephaluo among Kannemeyeriini (Lehman 1961). The most complete are  Parakannemeyeria youngi ,  Xiyukannemeyeria brevirootrio Liu and Li, 2003 ,  P. xingxianenoio ,  Sinokannemeyeria yingchiaoenoio, Dinodontooauruo tener ,  Rhadiodromuo klimovi , and Iochigualaotia jenoeni. </p>
            <p> Two species were selected as the reference standard for the reconstruction of the skeleton of  Lioosicia bojani :  Parakannemeyeria youngi and  Sinokannemeyeria yingchiaoenoio . </p>
            <p> Their humeri, ulnae, and radii have proportions closely similar to the bones of  L. bojani , despite different construction of the forelimb. Other dicynodonts differ substantially from  L. bojani , for instance adult Dinodontooauruo  tener material from Harvard has a very short scapula in relation to the length of the humerus (Cox 1965: fig. 11). </p>
            <p> The vertebral spine of  Lioosicia bojani and its relationship to the pelvis was based on the fit of bones in the individual ZPAL V. 33/720. The proportions of the pectoral girdle and forelimb were based on the proportions in the  Parakannemeyeria youngi specimen IVPP V. 979; with respect to the length of scapula vs. length of the humerus, it is 1.25, and the length of the humerus vs. length of the ulna is 1.10. The length of the scapula vs. length of the sternum is 1.61 in  Sinokannemeyeria yingchiaoenoio specimen IVPP V. 974. The length of the ulna vs. length of the radius is 1.88 in  S. yingchiaoenoio specimen IVPP V. 974 (measurements from: Sun 1963). </p>
            <p> Pelvic girdle and forelimb: The position of the humerus and scapulocoracoid in  Lioosicia bojani is similar to that in large mammals, such as rhinoceroses and hippopotami, as well as quadrupedal dinosaurs, such as the ceratopsians (Sulej and Niedźwiedzki 2019). In most Triassic dicynodonts the scapula was s et al most vertical and humerus almost horizontal. Such articulation would be difficult to maintain by an animal of  L. bojani size. Also, the trackway of some dicynodont shows the manus and pes in the same line (Hunt et al. 1993) and thus contradicts the traditional reconstruction of their forelimb. The disposition proposed for Triceratopo horriduo Marsh 1889 (Fujiwara 2009) with a more horizontal scapula and vertical humerus (very similar to  L. bojani ) seems more realistic. The size of the joint for the scapula on the posterior side of the humerus corresponds to the position of this bone. Large  Stahleckeria poteno that had area of the articulation with the scapula small probably represents an intermediate stage between the horizontal humerus of small dicynodonts with a small joint for the scapula (for instance  Oudenodon bainii Kammerer et al., 2011 (earlier  Dicynodon halli ) and Placeriao  ‘ gigao ’ that had an already large posterior joint for the scapula. An advanced stage with the vertical position of the humerus is represented by  L. bojani , which has a non-rotating humerus with a very large joint for the scapula. </p>
            <p> The shift of the humerus to a vertical position should be related to some changes in its articulation with the ulna and radius. The published illustrations of the dicynodont radii are too superficial to enable comparison with that of  Lioosicia bojani . This makes the well-preserved proximal part of the quite well-preserved radius from New Mexico NMMNH P-13002 important. This bone and associated femur, part of scapula and axis were described as Iochigualaotia jenoeni? by Lucas and Hunt (1993) but Kammerer et al. (2013) assigned it to  Stahleckeriidae indet. based on the shape of the femur. </p>
            <p> The scapula NMMNH P-13003 is a distal part of the bone showing the scapular spine and acromion process (nomenclature from: Vickaryous and Hall 2006). It seems that the scapula was very similar to that of Placeriao  ‘ gigao ’. The radius has a well-visible head bent outward, which is distinct also in Iochigualaotia jenoeni,  Jachaleria candelarienoio , and  Lioosicia bojani (all with not preserved clavicles and a small acromion process) and not known in other dicynodonts [illustration of Prioterodon mackayi Huxley, 1868 earlier  Diaelurodon shaitoi in Watson (1917: fig. 13) suggests that it was present in this species]. But the bending has a different position in the radius from New Mexico then in I. jenoeni,  J. candelarienoio , and  L. bojani . In these dicynodonts it is situated in the posterior part of the head, whereas in the New Mexico specimen it is in an anterior position, like  D. shaitoi , which had a horizontal position of the humerus. </p>
            <p> Thearticulationofthecoracoidandanteriorpartofthesternum occurs in the  Struthio cameluo Linnaeus, 1758 , Diplodocuo Marsh, 1878 (Hohn et al. 2011),  Alligator Cuvier, 1807 , Tachyglooouo Illiger, 1811, and Ornithorynchuo Blumenbach, 1800 (Gregory and Camp 1918). In Lioosicia bojani, the coracoid has a large joint with the anterior part of the sternum. In the coracoid of Placeriao  ‘ gigao ’ UCMP 32449, the area for attachment with the sternum is clearly visible. </p>
            <p> Sternum: The latest Triassic  Lioosicia bojani has an articulation area on the sternum in its posterior part. The sternum of the Anisian (Liu et al. 2017)  Sinokannemeyeria yingchiaoenoio has an articulation for the coracoid and first dorsal rib in the middle of its length. Cox (1965), based on the specimen MCZ 3120, depicted the sternum of Iochigualaotia jenoeni with an articulation area in its posterior part. It is the only specimen in the Harvard collection with all bones of the pectoral girdle articulated, although not all are in anatomical positions. The problem with the sternum is that it is strongly compressed, and no articulation surface is visible. The Cox (1965) interpretation was based only on the general shape of the bone. </p>
            <p> Romer (1956) showed the sternum of  Kannemeyeria oimocephaluo posterior to the scapulocoracoid. In the skeleton reconstruction of Dinodontooauruo  brevirootrio at Harvard, the interclavicle is at the level of the procoracoid, and the sternum is more posterior than in the specimen MCZ 3120. In such a probably correct position, the articulation area on the sternum can contact the posterior process of the coracoid (Sulej and Niedźwiedzki 2019). </p>
            <p> Clavicle: An intriguing problem is the presence of the clavicle and interclavicle in  Lioosicia bojani . It was hypothesized by Sulej and Niedźwiedzki (2019: fig. 1), but the very small acromion process on the scapula, which in other dicynodonts was much larger and designed for articulation with the clavicle, contradicts its presence (Fig. 48). On the other hand, the anterior lower part of the scapula in  L. bojani is much larger and the sternum is much higher than in most dicynodonts. This difference suggests different functioning of the whole girdle. Sulej and Niedźwiedzki (2019) showed that  L. bojani had erect forelimbs, unlike all other dicynodonts. Instead of them, Triceratopo or the rhinoceros may serve as the analogues for the construction of the shoulder girdle. They do not have clavicles because of the erect position of the forelimb. Probably a similar situation was in  L. bojani . The loose connection of the olecranon process with the main body of the ulna in its skeleton is probably related with the position of the forelimb. In most large dicynodonts the olecranon is fused with the ulna (Wadiaoauruo,  Stahleckeria ,  Jachaleria , and  Sinokannemeyeria ), and that was related to a sprawling posture. The m. triceps attached to the olecranon process, the humerus, and the scapula was among the muscles responsible for keeping the animal in that position. When the humerus was rotated posteriorly to support the erect posture, other muscles became responsible for it. Among them were m. pectoralis, m. supracoracoideus (with a much larger area for articulation on the scapula than in other dicynodonts) and m. coracobrachialis (Sulej and Niedźwiedzki 2019). The loose olecranon process is known also in Placeriao  ‘ gigao ’ (Camp and Welles 1956), but in this species the acromion was of the standard shape and it remains unknown how the rotation of the humerus took place. </p>
            <p>Presumably Placeriao represents an early stage of the evolution towards the erect posture.</p>
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	https://treatment.plazi.org/id/03A287B16B3A5565F231F903FE6CF827	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sulej, Tomasz	Sulej, Tomasz (2024): Osteology and relationships of the Late Triassic giant dicynodont Lisowicia. Zoological Journal of the Linnean Society 202 (1): 1-56, DOI: 10.1093/zoolinnean/zlae085, URL: https://doi.org/10.1093/zoolinnean/zlae085
03A287B16B005567F2B7FA61FDF8F90E.text	03A287B16B005567F2B7FA61FDF8F90E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lisowicia Sulej & Niedźwiedzki 2019	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Lisowicia mode of life </p>
            <p> Traditionally dicynodonts are depicted with abducted (directed to outside) forelimbs and nearly adducted (oriented towards the body axis) hindlimbs (Fröbisch 2006, Ray 2006). However, the trackways of a large dicynodont from the Middle Triassic named Pentaoauropuo argentinae make such posture questionable (Hunt et al.1993, Lagnaoui et al. 2019). This track is believed to have been left by a kannemeyerid dicynodont that was a sprawling limbed trackmaker with an abducted posture for the forelimbs and at least a semi-abducted posture for the hindlimbs (Abdelouahed et al. 2019). Sulej and Niedźwiedzki (2019) proposed that the giant  Lioosicia bojani had also adducted its forelimbs. This has also been discussed for pelycosaur tracks, which have a much more narrow gauge than the skeleton would seem to indicate. Twisting of the body and/or limbs while walking might explain how this happens (Hunt and Lucas 1998, Hopson 2015). </p>
            <p>The analysis of sediments (or bones) from Lisowice shows that Lioosicia lived in an environment near rivers with a lot of oxbow lakes (Dzik et al. 2008). Also, the similarities of coprolites of Lioosicia to Hippopotamuo amphibiuo Linnaeus, 1758 suggest its affiliation with such an environment (Bajdek et al. 2014, 2019). In the case of Placeriao, its bone microstructure suggests even an aquatic style of life (Fiorillo et al. 2000). If they lived in or close to the swamp, they probably could use the soft plants as a food (Bajdek et al. 2014), abundant in such environments. Such an interpretation is not in conflict with the architecture of their skulls.</p>
            <p> There is little doubt that all the Triassic dicynodonts were herbivorous, but there are various views on the method by which they collected the food (see: Surkov and Benton 2004). According toCruickshank (1978), Dinodontooauruo,  Stahleckeria poteno, Dolichuranuo Keyser, 1973 , Tetragoniao,  Rhinodicynodon, Zambiaoauruo ,  Sinokannemeyeria , and  Vinceria Bonaparte, 1969 were browsers, whereas Placeriao,  Jachaleria, Iochigualaotia jenoeni,  Kannemeyeria ,  Uralokannemeyeria Danilov, 1971 , Rabidooauruo, Rhadiodromuo, and Wadiaoauruo indicuo were grazers (in the meaning that they ate low-growing plants). Originally, Cox (1965) made this distinction based mainly on the shape of the snout and the orientation of the occipital region of the skull. Ordoñez et al. (2019), based on principal component analyses (PCA) of skulls from South America, showed that the adaptation of  Stahleckeria poteno , I. jenoeni, and  J. candelarienoio to feeding on the vegetation was characteristic of an arid climate, although the palaeoclimate was seasonal semi-arid when they lived (Mancuso et al. 2021). The new material from Poland calls for reconsideration of this question. </p>
            <p> Most authors discussing the dicynodonts mode of life have focused on their sexual dimorphism (Owen 1876, Camp and Welles 1956, Barry 1957, Cruickshank 1967, Cox 1969, Bandyopadhyay 1988, Sullivan et al. 2003). The differences between males and females were proposed to be expressed mainly in the presence of the tusks or maxillary horns in males. The lack of conclusive material excludes  Lioosicia bojani from these inquires. Most authors agree that the canine tusks or elongated maxillary processes were used in food gathering and fighting (Camp and Welles 1956, Rowe 1979, Bandyopadhyay 1988). Many authors have discussed the mode of feeding of dicynodonts (Cox 1969, Cruickshank 1978, Walter 1985, Hotton 1986), summarized by Defauw (1989), who recognized five dicynodont feeding types: invertebrate collecting specialists, grubbers, browsers, forest litter foragers, and flexible foragers. The functional morphology of the dicynodont masticatory apparatus was studied by Crompton and Hotton (1967), King et al. (1989), Cox (1998), Jasinoski et al. (2009, 2010), and Ordoñez et al. (2019). </p>
            <p> Niedźwiedzki et al. (2011, 2012) showed that the archosauriform  Smok saseloki Niedźwiedzki et al., 2012 was feeding on dicynodonts in the Late Triassic but probably dicynodonts escaped this predation by evolving into giant size, already in the Middle Triassic. The most numerous preserved bones of the skull of  Lioosicia bojani and Placeriao  ‘ gigao ’ are postorbitals and basisphenoids, unlike the nasals that are very rare in the case of  P. ‘ gigao ’ and unknown in  L. bojani . This may mean that the snout was a structurally weak portion of the skull that was the first to disarticulate during rotting of the carcass or the dorsal part of the snout was an attractive target for predators, whereas the postorbital was too massive for them. </p>
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	https://treatment.plazi.org/id/03A287B16B005567F2B7FA61FDF8F90E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sulej, Tomasz	Sulej, Tomasz (2024): Osteology and relationships of the Late Triassic giant dicynodont Lisowicia. Zoological Journal of the Linnean Society 202 (1): 1-56, DOI: 10.1093/zoolinnean/zlae085, URL: https://doi.org/10.1093/zoolinnean/zlae085
03A287B16B0F556FF2D5FB61FF15FE79.text	03A287B16B0F556FF2D5FB61FF15FE79.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhadiodromus klimovi (Efremov 1940)	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> The Laurasian–Gondwanan  Rhadiodromus klimovi →  Jachaleria candelariensis lineage </p>
            <p> Middle Triassic European dicynodonts are known from Russia. Rabidooauruo  criotatuo and  Rhadiodromuo klimovi and  Rhadiodromuo mariae (Surkov 2003) are there represented by the most complete materials (Surkov 2003) from the Donguz and Bedyanka localities of the Lower and Upper Donguz formations, respectively, which is dated as Anisian (Tverdokhlebov et al. 2003, Ivakhnenko 2008).  Parakannemeyeria youngi Sun, 1960 and  Shaanbeikannemeyeria xilougouenoio Cheng, 1980 from the Early Anisian Ermaying Formation in China (Sun 1963, Liu et al. 2017) may be their relatives. Angielczyk et al. (2018) showed that Sanguoauruo parringtonii from the upper Ntawere Formation in Tanzania and Zambia is a stahleckeriid.  Stahleckeria poteno from the Santa Maria Formation in Brazil is dated as Ladinian (or Carnian? according to: Lucas 1998b, Rayfield et al. 2005, Ordoñez et al. 2020; see also: Schultz et al. 2000). The new  Stahleckeria sp. indet. material was reported from the Carnian Chañares Formation of the Ischigualasto-Villa Unión Basin by Escobar et al. (2021). Iochigualaotia jenoeni is known only from the lower third of the Ischigualasto Formation (Bonaparte 1970, Rogers et al. 1993, Zerfass et al. 2003, Martinez et al. 2011). The latter authors dated the formation as 227.8 ± 0.3 Mya, i.e. Late Carnian. Fröbisch (2009; based on: Lucas 1998b) considered it to represent the upper part of Adamanian, but according to Langer (2005b) it is older than Adamanian. He also showed that probably I. jenoeni is older than Eubrachiooauruo brosni Williston, 1904 from the Popo Agie Formation [contrary to Lucas and Hunt (1993), the relative ages of these two taxa is highly uncertain in the absence of precise radiometric dates for the Popo Agie Formation]. </p>
            <p> According to Ramezani et al. (2014), the lower third of the Ischigualasto Formation represents the boundary between the Carnian and Norian. The youngest dicynodonts from South America are  Jachaleria colorata from the Norian Los Colorados Formation of Argentina, which is a fluvial–lacustrine siliciclastic (Bonaparte 1971, 1978, Vega-Dias and Schwanke 2004), and  J. candelarienoio (Vega-Dias and Schultz 2004) that originated from the Norian Caturrita Formation (Araújo and Gonzaga 1980, Langer et al. 2018). The close relationship of  Stahleckeria, Eubrachiooauruo ,  Jachaleria , and Iochigualaotia was supported by the cladistic analysis published by Kammerer et al. (2013) and Szczygielski and Sulej (2023). </p>
            <p>These dicynodonts experienced a mosaic evolution, with modifications of particular bones not necessarily correlated with each other. This refers especially to the bones of the cranium.</p>
            <p> Evolution of the cranium: The position of the dicynodont orbits is correlated with the shape of the postorbital. This bone is perpendicular to the zygomatic arch in  Rhadiodromuo klimovi , the oldest member of the lineage, and in subsequent evolution became more and more oblique anteriorly. The extreme state of this trait is that in  Jachaleria candelarienoio . In  J. candelarienoio and Iochigualaotia jenoeni the orbits are located above the posterior margin of the maxillary horn and the temporal opening is very large. </p>
            <p> The route of the evolution of the frontals was different in the lineage represented in South America by  Stahleckeria, Iochigualaotia , and  Jachaleria . The anterior part of the bone became shorter, and the narrow edge of the orbital margin almost disappeared and moved outside. The original shape of the frontal may be shown by  Rhadiodromuo mariae Surkov, 2003 . In its very wide anterior part,  R. mariae is more similar to  Stahleckeria . Also, the large number of sacral ribs of  Rhadiodromuo klimovi is typical for the Middle Triassic dicynodonts from South America. It seems that Rabidooauruo and Rhadiodromuo are genera that gave the beginning to different lineages. </p>
            <p> Probably in the Ladinian, the tusks vanished from the maxillae in the South American members of the lineage, although they were present until the Carnian in Dinodontaouruo  brevirootrio in the other Gondwanan lineage and until the Norian in the Laurasian lineage. </p>
            <p> Some changes can be also observed in the disposition of the adductor externus lateralis and externus medialis muscles, which are attached to the zygomatic arch (Ordoñez et al. 2019). In most Early and Middle Triassic dicynodonts the zygomatic arch in dorsal view is directed anteriorly (Fig. 49). In  Stahleckeria , the zygomatic arch is directed antero-medially at its base. From Iochigualaotia to  Jachaleria it became directed more and more laterally. The changes in the shape of the zygomatic arch are correlated with the shape of the occiput.The lateral edge of the occipital plate (posterolateral wing of the squamosal) in most Triassic dicynodonts had an apparent posterior edge (in lateral view) and forms a large attachment area for muscles (Figs 49, 50). This area was small only in Iochigualaotia and  Jachaleria , in which the lateral edge of the occipital plate in lateral view is vertical. This characterizes both species of the genus,  J. colorata (Bonaparte 1978, Vega-Dias and Schwanke 2004) and  J. candelarienoio (Vega-Dias and Schultz 2004) . The slope of the lateral edge of the occipital plate resulted in a different orientation of the adductor muscles with respect to the mandible. The external adductor attachment was enlarged due to the horizontal position of the zygomatic arch. It seems that  J. candelarienoio had enormously strong adductors, probably to feed on hard food. The morphology of the occipital is strongly correlated with the slope of the whole skull. </p>
            <p> Already Surkov and Benton (2004) and Kalandadze and Kurkin (2000) interpreted proportions of the occipital plate and the whole skull in the context of feeding adaptations. However, their ‘occipital index’ mixes proportions of the occipital plate and the length of the skull. Proportions of the occipital plate in Middle and Late Triassic dicynodonts exhibit two separate types.  Stahleckeria, Dinodontooauruo, and Rabidooauruo have very low and wide occipitals, whereas Iochigualaotia,  Jachaleria , and Placeriao have high and relatively narrow occipitals. It seems that increasing skull height and narrowing evolved in parallel in both lineages. </p>
            <p> In both species of  Jachaleria the occipital condyles were directed posteriorly. In the resting position of the skull the orbits were directed frontolaterally. </p>
            <p> The shape of the mandible is variable in Triassic dicynodonts, and it is difficult to identify any evolutionary trend in its morphology. Only the shape of the dentary seems to differentiate the lineages. In  Stahleckeria (Abdala et al. 2013) , Dinodontooauruo, and Iochigualaotia it is high and short, unlike Placeriao. </p>
            <p> Evolution of pootcranial okeleton: Although most aspects of the postcranial skeleton are variable in Triassic dicynodonts, some trends are identifiable. The most important changes concern the pectoral girdle. Regrettably, the scapula of dicynodonts from the Anisian of Russia remains unknown. The increase in size of the acromion process of the scapula characterizes the evolution of the longest lasting lineages. In  Stahleckeria (Escobar et al. 2021) , the acromion process was elongated into a ridge (scapular spine) that extended almost to the upper end of the scapula. In Iochigualaotia, only the base of the ridge was preserved.  Jachaleria had a very small acromion process and a distinct attachment for the triceps scapularis (Araújo and Gonzaga 1980). According to Surkov et al. (2005), the general trend to widening of the scapula blade is observed already in the Permian dicynodonts (Rubidge et al. 1994), and the reversal of this trend occurred in Late Triassic species (Fig. 50). The Late Triassic dicynodonts from South America had a very wide base of the scapular blade (Kammerer et al. 2013). This character is conservative. </p>
            <p> Stahleckeria and Iochigualaotia had a small groove on the anterior edge of the scapular blade. This seems important as Placeriao and Lioosicia did not have such a structure. It is interesting that the share of the coracoid in the formation of the glenoid is low in  Stahleckeria , in contrast to Iochigualaotia. In both lineages also the direction of the glenoid changed in parallel. At the beginning it is directed laterally, as in all Middle Triassic dicynodonts, but it changed to a more posterior direction. The role of the triceps brachii muscles probably changed in the Late Triassic dicynodonts. In advanced form, like  Jachaleria , the attachment area for these muscles on the scapula is very large. </p>
            <p> It seems that in  Stahleckeria and  Jachaleria the sternum had two articulation surfaces on each side. This character is not known in older representatives of the lineage. </p>
            <p> The shape of the ilium is variable in dicynodonts, and only the number of sacral ribs and the length of the posterior process may allow identification of evolutionary trends (Fig. 50). The Middle Triassic Rhadiodromuo had many sacral ribs (seven to eight?). The trend to decrease their number characterizes South America dicynodonts.  Stahleckeria had seven to eight sacral ribs, Iochigualaotia six to seven? [uncatalogued specimen in Instituto Miguel Lillo in Tucuman; contrary to Griffin et al. (2019], and  Jachaleria only five sacral ribs. It seems that small Permian dicynodonts had four sacral vertebrae, but as they grew in size they had to increase the number of sacral vertebrae so that the pelvis could support larger weight (Rhadiodromuo and  Stahleckeria ), but then when the forelimbs started to be displaced under the shaft, this was no longer necessary and the number of sacral vertebrae decreased. In  Jachaleria and Iochigualaotia, two or three sacral ribs are in front of the acetabulum. </p>
            <p> The femur from the Los Esteros Member of the Santa Rosa Formation in New Mexico, is more similar to  Stahleckeria than to Placeriao (Kammerer et al. 2013) and suggests that some representative of the Gondwanan lineages came to live in North America. Eubrachiooauruo may be such an immigrant. The distinction between South and North American lineages of dicynodonts is especially well expressed in the morphology of the pubis and ischium, although they are known only in more advanced representatives. The notch in the ventral border of the ischium and pubis is very distinct in  Stahleckeria . The ischium of  Jachaleria has the vertical length very short in comparison to very long in Lioosicia and Placeriao. The ischium has a posterior blade slightly curved medially in  Jachaleria .  Jachaleria probably represents the crown achievement of dicynodont evolution in South America. It seems that its mode of life was very different from that of Placeriao and Lioosicia. Their skulls are very different. The wide snout with well-developed grooves for tearing plants, together with the very large area for attachment of muscles adducting mandible and the massive zygomatic arch, suggest that  Jachaleria ate a hard plant food difficult to tear. The position of the quadrate that is directed ventrally (in Lioosicia rather anteroventrally) may be related with the mode of tearing but is difficult to explain. </p>
            <p>The almost oval and longer than high parietal, the short and only slightly oblique posterior surface of the supraoccipital, and the almost horizontal base of the braincase suggest that Jachaleria kept its head horizontal and was a browser. This is consistent with the position of the orbits. They were very large and displaced anteriorly to be located above the maxilla. In Lioosicia they are much more posterior.</p>
            <p> The question about pass of dicynodonts, from Laurasia to South America that Rhadiodromuo and  Parakannemeyeria could be ascendants of South American dicynodonts seems problematic, but Haq (2018) showed that in the Anisian, the sea level was very low, and later it rose until the Carnian/Norian boundary. It means that in the Anisian, large lands were accessible for migrating animals and in that time the terrestrial communication between these distant lands was possible also for dicynodonts. </p>
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	https://treatment.plazi.org/id/03A287B16B0F556FF2D5FB61FF15FE79	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sulej, Tomasz	Sulej, Tomasz (2024): Osteology and relationships of the Late Triassic giant dicynodont Lisowicia. Zoological Journal of the Linnean Society 202 (1): 1-56, DOI: 10.1093/zoolinnean/zlae085, URL: https://doi.org/10.1093/zoolinnean/zlae085
03A287B16B0A5571F182FDC3FD45FF5F.text	03A287B16B0A5571F182FDC3FD45FF5F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Shaanbeikannemeyeria xilougouensis (Cheng 1980)	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> The Laurasian lineage  Shaanbeikannemeyeria xilougouensis →  Lisowicia bojani</p>
            <p> Until the description of the Polish dicynodonts, Placeriao  ‘ gigao ’occipitals were the most well-known dicynodont from the Late Triassic of Laurasia. Its diagnostic characters are elongated frontals, parietal triangular in cross-section, and maxillae with long ‘tusks’. They were ignored by earlier researchers and the species was grouped together with the Gondwanan dicynodonts. According to Vega-Dias et al. (2004) they were closely related and may represent a lineage initiated by  Stahleckeria poteno (Damiani et al. 2007) . King (1988, 1990) distinguished two separate ‘suites’: Placerinii for  P. ‘ gigao ’ and Iochigualaotia jenoeni with a thin, tapering snout, and Stahleckerini for  Stahleckeria poteno and Zambiaoauruo oubmeroeo (probably juvenile). Keyser and Cruickshank (1979) discussed alternative origins for I. jenoeni from Dinodontooauruo or from  Kannemeyeria oimocephaluo . The close relationship of  K. oimocephaluo with  S. poteno and I. jenoeni was posited by Damiani et al. (2007). In many recent phylogenetic analyses, the South American taxa have been joined together (Griffin and Angielczyk 2019, Sulej and Niedźwiedzki 2019, Kammerer and Ordoñez 2021). Lioosicia and Placeriao are probably not closely related to the Gondwanan ones. </p>
            <p> Placeriao  ‘ gigao ’comes from the basal Bluewater Creek Formation at the  Placerias quarry near St. Johns, Arizona, dated as Adamanian. Placeriao  heoternuo is known from the stratigraphically higher Blue Mesa Member of the Petrified Forest Formation from just north-east of Cameron, Arizona (Lucas 1904, 1995, 1998a, Lucas and Hunt 1993, Lucas and Heckert 1996, Heckert and Lucas 2002) of slightly younger age (Heckert 2004). Many new studies have shown that both taxa come from the Blue Mesa Member, which is dated as ~223 to ~218 Mya (Lucas 1993, Martz et al. 2017, Gehrels et al 2020). The localities are 150 km apart, so the different exact ages are possible. These are separate species (Camp and Welles 1956) that have different humeri (contrary to: Lucas and Hunt 1993, Kammerer et al. 2013). The supinator process is more proximal then the entepicondylus in  P. ‘ gigao ’ (like in  Lioosicia bojani ), than in  P. heoternuo [compare fig. 5D and 6A in Kammerer et al. (2013)]. The edge above the entepicondylus is straight in  P. heoternuo and concave in  P. ‘ gigao ’ (similar to  Lioosicia bojani , but not so much). The deltopectoral crest is more laterally expanded in  P. heoternuo than in  P. ‘ gigao ’ (clearly visible in the best preserved specimen of proximal part GPIT-PV-108382). </p>
            <p> According to Kammerer (2018), Pentaoauruo goggai from the lower Elliot Formation (probably Norian age) represents the latest surviving Placerinii, but the material is very poor, and similarities to Placeriao are very weak. In some aspects it covered as Placerinii in recent analyses of dicynodont evolution (Kammerer 2018). In fact the most characteristic element is the distal head of the humerus, which is very different than in Lioosicia, and Placeriao  ‘ gigao ’. Any humerus of  Jachaleria is unknown. Only the posterior part of the frontal is elongated like in Lioosicia, but for the most, part of the bone is lacking. If it is true that Pentaoauruo goggai represents the latest surviving Placerinii, it will be evidence that this group is not characteristic only for Laurasia, but more specimens are needed to confirm that. </p>
            <p> The age of the youngest member of the lineage,  Lioosicia bojani , is close to the Norian/Rhaetian boundary (Sulej and Niedźwiedzki 2019). Its older relative, Woznikella triradiata, originates from the Carnian sediments from Poland and Germany (Schoch 2012, Szczygielski and Sulej 2023). </p>
            <p> Evolution of the cranium: Placeriao  ‘ gigao ’ and  Lioosicia bojani have exceptionally small areas for muscles responsible for adducting the mandible. Moreover, the zygomatic arch to which some of these muscles are attached is very thin and delicate. The snout is very thin, and the grooves for the dentary are poorly developed, which means that the area for tearing food is rather small. These aspects of the skull anatomy are derived and opposite to those of  Jachaleria candelarienoio . In some Permian dicynodonts there was a large ridge in the anterior part of the surface for articulation with the quadrate, which was a barrier for the quadrate. It is absent in  P. ‘ gigao ’ and  L. bojani . This means that the quadrate could drop anteriorly from its normal position when the mandible moved backward. Apparently, the forces acting on the jaw were much smaller than in the case of Permian dicynodonts. It seems that in the evolution of Triassic dicynodonts the shape of the frontal was strictly controlled by selection. Especially the shape of its anterior part and the morphology of the contact with the orbital margin are the most characteristic and useful for understanding the evolution of that group. </p>
            <p> In the Laurasian lineage, the anterior part of the skull is elongated. In all species the frontals form a part of the orbital margin. It is most elongated in Placeriao  ‘ gigao ’, similar to  Lioosicia bojani (Fig. 48). The specimen of the older Woznikella triradiata (Sulej et al. 2011, Szczygielski and Sulej 2023) has the frontal partly preserved, but it shows a long edge forming the orbital margin and an elongation of the anterior part of the frontal is suggested by well-preserved nasals. Even older dicynodonts with a similar frontal are those from the Anisian of Russia:  Rhadiodromuo klimovi and the smaller Rabidooauruo  criotatuo (Ochev and Shishkin 1989) .  Rhadiodromuo mariae is known after a complete skull but from a different locality of the same formation. They all have a very long anterior part of the frontal and a very long edge of the frontal forming the orbital margin (which is characteristic for almost all Anisian dicynodonts). Both species of  R. mariae have orbits directed strongly dorsally and the frontal elongated anteriorly. Rabidooauruo  criotatuo has the frontals with a distinct high posterior process, and its parietals are similar to those of  P. ‘ gigao ’. The slightly older  Shaanbeikannemeyeria xilougouenoio has frontals elongated anteriorly. The same type of suture between frontals and nasals is present in both species of Rhadiodromuo. </p>
            <p> The position of the orbits is correlated with the shape of the postorbital. This bone is oblique posteriorly in Placeriao ‘ gigao ’ and Lioosicia bojani. In these species the orbits are located more posteriorly, and the temporal opening is smaller; it was related with the size of the external adductor muscles. The zygomatic arch morphology is not known in  P. ‘ gigao ’ and  L. bojani , but preserved fragments of the squamosal suggest that it was directed slightly antero-medially. In  P. ‘ gigao ’ the lateral edge of the occipital plate in lateral view is strongly oblique posteriorly (to the horizontal position of the frontal). The adductor muscles were attached to the mandible more posteriorly. </p>
            <p> In Placeriao  ‘ gigao ’ the occipital condyles are directed ventrally, which means that in the resting position the skull was strongly oblique ventrally (Fig. 50). It is consistent with the shape of the orbits, which are opened antero-dorsally and frontally in an oblique position of the skull. </p>
            <p> There is no evidence in the fossil material for the reconstruction of Placeriao  ‘ gigao ’ by Camp and Welles (1956: fig. 24), with the dentary much higher than the posterior part of the mandible. Also, in Woznikella triradiata the dentary is relatively long and low. It is more similar to the elongated one of  P. ‘ gigao ’, than to any other dicynodont from South America. In Lioosicia the dentary is unknown but the articulation for the dentary on the angular suggests that it was high and with a concave middle edge of the mandible, almost like in Iochigualaotia jenoeni. </p>
            <p> Evolution of pootcranial okeleton: The acromion process trend to decrease is apparent in the Laurasian lineage. Woznikella triradiata has a scapula with the acromion process high but short. Placeriao  ‘ gigao ’ has an elongated ridge (Camp and Welles 1956: fig. 29) and  Lioosicia bojani has the process very small. In both species the base of the scapular blade is relatively narrow. It has a similar shape in  L. bojani , but in this species the end of the scapular blade is very wide, probably as a result of the large size of the animal. Probably a decrease in size of the acromion process was convergent in both these long-lasting lineages. </p>
            <p> The triceps brachii muscles probably changed their role in the Late Triassic dicynodonts. In the advanced forms like Placeriao  ‘ gigao ’, the attachment for these muscles on the scapula is relatively small, and in Lioosicia it is diminutive. </p>
            <p> Sinokannemeyeria yingchiaoenoio is the oldest Laurasian form with a single articulation surface on the sternum. This continued to occur in Placeriao  ‘ gigao ’ and  Lioosicia bojani . The Late Triassic increase in dicynodonts general size was related to a gradual change of the position of the humerus and decrease in size of the acromion process on the scapula (Sulej and Niedźwiedzki 2019). The change in movement of the humerus affected the position of the articulation surface for the coracoid and ribs on the sternum. In both lineages this surface moved posteriorly. In the Laurasian lineage, the sternum of Lioosicia with extremely posteriorly set the articulation and very high ridges represents probably the last stage of evolution. </p>
            <p> The underived humerus of  Sinokannemeyeria yingchiaoenoio and  Kannemeyeria oimocephaluo have rotated distal and proximal heads, whereas in Lioosicia they are almost in the same plane; only the deltoid crest is curved ventrally. This decreasing of the rotation was related to changes in the orientation of the humerus relative to the scapulocoracoid. Ray (2006) stated that the humerus changed its position, and it was related to the change from the lateral orientation of the glenoid to a posterior orientation. </p>
            <p> The size of the supinator process seems to have become gradually larger in the sequence: Woznikella triradiata (Szczygielski and Sulej 2023), Placeriao  ‘ gigao ’, and  Lioosicia bojani. Surprisingly, the humerus of Zambiaoauruo from the Anisian already had a large supinator process similar to that in  P. ‘ gigao ’ (Kammerer et al. 2013). In the Gondwanan lineage, dicynodonts and the Chinese  Sinokannemeyeria yingchiaoenoio the entepicondyle is large, whereas it is small in  P. ‘ gigao ’ and even smaller in  L. bojani . Probably also the medial epicondyle became narrower during evolution of the Laurasian lineage. Other postcranial skeletal characters seem to be more variable in the Laurasian dicynodonts. The articulation surface for the ulna on the dorsal side of the humerus is very large in  S. yingchiaoenoio and  P. ‘ gigao ’ but very small in  L. bojani . </p>
            <p> In the Laurasian lineage, the number of sacral ribs was probably small from the beginning.In  Sinokannemeyeria yingchiaoenoio there were only five, but in most genera it remains unknown. Placeriao  ‘ gigao ’ has a rather underived ilium with five sacral ribs (Camp and Welles 1956), although only three areas for articulation are visible.  Lioosicia bojani has only four sacral ribs, and the first sacral rib is above the acetabulum. </p>
            <p> The North American Eubrachiooauruo brosni probably represents a South American immigrant lineage (Kammerer et al. 2013). Its well-preserved pelvis shows a frontally elongated ilium with a curved lower end, with a ridge on the blade, and its pubis is very small in relation to the ischium. It is also older (Camp and Welles 1956) than Placeriao  ‘ gigao ’ but differences in the morphology make an ancestor–descendant relationship unlikely. </p>
            <p> The ischium of  Lioosicia bojani and Placeriao  ‘ gigao ’ are vertically elongated. Placeriao  ‘ gigao ’ and  L. bojani have their posterior blade a little curved medially. The vertically short ischium of  Parakannemeyeria chengi probably represents an underived stage for all three lineages. In Placeriao  ‘ gigao ’ specimens, this part of the ischium is broken but in  L. bojani its shape suggests such a morphology. </p>
            <p> The position of the proximal head of the femur changed during evolution to large in Placeriao  ‘ gigao ’ and  Lioosicia bojani , which have the proximal head directed dorsally in lateral view, thus the femur had a vertical position while resting. </p>
            <p> Evolution of the mode of life: It is a matter of controversy whether dicynodonts were ‘grazers’ or browsers (Cox 1959, Kalandadze and Kurkin 2000, Surkov and Benton 2008, Ordoñez et al. 2019). It seems that the shape of the parietal (especially its medial section), and position of the occipital condyle and the orbits are strongly connected with the disposition of the head and the way of seeing the food. In Placeriao  ‘ gigao ’ and  Lioosicia bojani the angulation of the base of the braincase suggests that the skull had an oblique orientation with the snout very low above the ground. Also, the occipital condyle is low under the jaw articulation and far from the top of the skull, which is situated more posteriorly than in such dicynodonts as  Jachaleria candelarienoio or  Stahleckeria poteno . </p>
            <p> To keep the head oblique demanded a special position of the orbits, because ‘grazing’ animals need to be aware of predators. This was probably the selection pressure to makes the orbits displaced to the top of the skull in Placeriao  ‘ gigao ’ and  Lioosicia bojani . While lowering the head the animal saw the surroundings in the horizontal plane. Present-day large herbivorous animals with orbits similarly directed or situated more dorsally are the hippopotamuses, connected with aquatic environments. </p>
            <p>However, animals spending much of their life in water often have lighter limb skeletons, which is not the case of Lioosicia.</p>
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	https://treatment.plazi.org/id/03A287B16B0A5571F182FDC3FD45FF5F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sulej, Tomasz	Sulej, Tomasz (2024): Osteology and relationships of the Late Triassic giant dicynodont Lisowicia. Zoological Journal of the Linnean Society 202 (1): 1-56, DOI: 10.1093/zoolinnean/zlae085, URL: https://doi.org/10.1093/zoolinnean/zlae085
03A287B16B145571F1A3FF1CFB67FBEB.text	03A287B16B145571F1A3FF1CFB67FBEB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kannemeyeria simocephalus (Weithofer 1888)	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> The Gondwanan lineage  Kannemeyeria simocephalus →  Dinodontosaurus brevirostris</p>
            <p> In the Induan, the Gondwanan dicynodonts are represented by the single genus  Lyotrooauruo known from Antarctica, Russia, China, India, and South Africa, but it has not been reported from the Olenekian (Fröbisch 2013).  Kannemeyeria oimocephaluo and Dolichuranuo primaevuo are the best known kannemeyerids from Africa.  Kannemeyeria oimocephaluo is a medium to large dicynodont (Govender et al. 2008) from the Cynognathus Assemblage Zone of the South Africa dated as Early Anisian (Catuneanu et al. 2005, Hancox et al. 2020); however, some zircon studies of the Puesto Viejo Group (San Rafael depocenter, Argentina) suggest a Carnian age for this assemblage (Ottone et al. 2014). Dolichuranuo primaevuo Keyser, 1973 was described from the Omingonde Formation, Karoo Supergroup, Waterberg Basin; Middle Triassic, probably Anisian–Ladinian (Damiani et al. 2007, Wynd et al. 2018, Zieger et al. 2020). </p>
            <p>The Middle Triassic kannemeyerids are represented by Wadiaoauruo indicuo and Rechnioauruo criotarhynchuo RoyChowdhury, 1970 from the Anisian Yerrapalli Formation of India (Chowdhury 1970,KeyserandCruickshank 1979,Bandyopadhyay 1988, Bandyopadhyay and Sengupta 2006, Ottone et al. 2014), which was related with South Africa at that time.</p>
            <p> The South American Ladinian kannemeyerids dicynodonts are Acratophoruo  argentinenoio Kammerer and Ordoñez, 2021 from the Rio Seco de la Quebrada Formation of Argentina (Bonaparte 1967, Lucas and Harris 1996, Renaut and Hancox 2001, Arcucci et al. 2004, Zavattieri and Arcucci 2007, Kammerer and Ordoñez 2021) and  Kannemeyeria aganooteuo Kammerer and Ordoñez, 2021 . Based on the similarity of faunas it was correlated with the Cynognathus zone. Dinodontooauruo  brevirootrio was the most common in the Late Triassic of South America. Kammerer and Ordoñez (2021) recognized  D. tener and  D. brevirootrio as the only valid species of the genus.  Jachaleria platygnathuo is a nomen dubium (Morato et al. 2006). </p>
            <p> Evolution of the cranium: The significance of proportions of the skull was studied by Cox and Li (1983). The morphology of the cranium of dicynodonts from the Permian of South America and South Africa is well known (Ordoñez et al. 2020, de Simão-Oliveira et al. 2020). The Triassic lineage probably started from  Kannemeyeria oimocephaluo (Govender et al. 2008) . It had the orbits directed dorsally and elongated, and a narrow snout. Acratophoruo  argentinenoio has a short frontal without any middle anterior process (Renaut and Hancox 2001, Domnanovich and Marsicano 2012, Kammerer and Ordoñez 2021), similar to that of  K. oimocephaluo . But, in younger Ladinian and Carnian? (Kammerer and Ordoñez 2021) species of Dinodontooauruo, the anterior process became longer (Kammerer and Ordoñez 2021) and the whole frontal narrower. It differs from  Sungeodon kimkraemerae in the shape of the frontal–nasal suture. In  K. oimocephaluo , the nasals form the posterior process running between the frontals, but in  S. kimkraemerae the situation is the opposite, the frontals form an anterior process running between the nasals. This shape is very similar to that of  Rhadiodromuo mariae . </p>
            <p> Evolution of the pootcranial okeleton: The sternum probably had a function related mainly to the movement of the forelimb and, unlike the skull, it was not related to diet. The oldest known Triassic sternum of kannemeyerids is that of  Kannemeyeria oimocephaluo from the Karoo in which there are two surfaces for the coracoid and first dorsal rib, and it is probably the starting point of the evolution. The sternum of the Gondwanan Wadiaoauruo indicuo has two distinct surfaces, unlike the Laurasian dicynodonts having one large surface [contrary to Bandyopadhyay (1988)]. </p>
            <p> Dinodontooauruo  tener has five to six sacral ribs (but juvenile ilium had six sacral ribs). In all these species (except specimen MCN-PV-1489 of Dinodontooauruo) the first sacral rib is sutured very far frontally on the iliac blade. According to Govender et al. (2008)  Kannemeyeria oimocephaluo had five sacral ribs. In Acratophoruo  argentinenoio the ilium is known but the number of sacral ribs was not determined. The posterior process of the ilium is very short in most Triassic dicynodonts. Only representatives of the lineage of  A. argentinenoio and  D. brevirootrio had this process elongated. </p>
            <p> The distinction between South and North American lineages of dicynodonts is also expressed in the morphology of the pubis and ischium. The ischium of Dinodontooauruo  brevirootrio has the vertical length very short in comparison to very long in  Lioosicia bojani and Placeriao  ‘ gigao ’. The ischium has the posterior blade strongly curved medially in Dinodontooauruo. In the Gondwanan lineages, the notch in the ventral border of the ischium and pubis is distinct (in  Kannemeyeria oimocephaluo and  D. brevirootrio ) but in Wadiaoauruo indicuo this notch is shallow. </p>
            <p> In the underived  Kannemeyeria oimocephaluo the proximal head of the femur is directed antero-medially in lateral view, which results in being set obliquely anteriorly. In maximum posterior position, it was set vertically. </p>
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	https://treatment.plazi.org/id/03A287B16B145571F1A3FF1CFB67FBEB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sulej, Tomasz	Sulej, Tomasz (2024): Osteology and relationships of the Late Triassic giant dicynodont Lisowicia. Zoological Journal of the Linnean Society 202 (1): 1-56, DOI: 10.1093/zoolinnean/zlae085, URL: https://doi.org/10.1093/zoolinnean/zlae085
