Supplementary Material for

www.sciencemag.org/cgi/content/full/science.aal4853/dc1 Supplementary Material for An elephant-sized Late Triassic synapsid with erect limbs This PDF file includes: Materials and Methods Supplementary Text Figs. S1 to S16 Tables S1 to S7 References Tomasz Sulej and Grzegorz Niedźwiedzki* *Corresponding author. Email: grzegorz.niedzwiedzki@ebc.uu.se Published 22 November 2018 as Science First Release DOI: 10.1126/science.aal4853 Other Supplementary Material for this manuscript includes the following: (available at www.sciencemag.org/content/science.aal4853/dc1) Movies S1 to S4

Materials and Methods The bone specimens preserved in the siltstone/mudstone demanded only cleaning in water. Those covered by mineralized limestone or pyrite crust were prepared using both chemical and mechanical methods. For mechanical preparation, matrix was removed using micro pneumatic hammers, whereas pyrite or dolomitized layers were removed with the help of a small metal chisel. Chemical preparation was done by bathing specimens in acid solution. After washing in water to remove calcium formate, they were dried and in part impregnated with dilute cyanoacrylic glue. The procedure was repeated up to the desired result. We assessed the phylogenetic affinities of Lisowicia bojani using the modified dataset and protocols presented in Kammerer et al. (22) and most recent Angielczyk and Kammerer (28). Large limb bones of adult or subadult individuals (complete femur, ZPAL V.33/763 and fragment of tibia, ZPAL V.33/765) were selected for preliminary histological studies. The sampled elements are from two different individuals. Owing to their ossification and large size it is likely that bones represent sub-adult or adult individuals. Both bones were sampled in the midshaft region. Thin sections are archived in the Institute of Paleobiology, Polish Academy of Sciences (Warsaw). Cervical vertebrae (ZPAL V.33/720 and ZPAL V.33/721) were computed tomography (CT) scanned at Warsaw in Military University of Technology by Krzysztof Karczewski using a Nikon XT H 225 ST. Data were reconstructed and rendered/examined using VG Studio MAX 2.0 by Daniel Snitting (Uppsala University). The CT data is archived in the Institute of Paleobiology, Polish Academy of Sciences (Warsaw). Two limb bones (PAL V.33/96, humerus of total length 48 cm and ZPAL V.33/763, femur of total length 72.3 cm), of a very large individual were used to estimate the mass of Lisowicia bojani. Ths scaling equation was based on results presented by Campione and Evans (14). Supplementary Text Geology, biostratigraphy and correlation. These fossiliferous beds are about 12 meters thick (Fig. S1) and were exposed in the Lipie Śląskie clay-pit at Lisowice village, near the town Lubliniec in southern Poland (10, 29). According to lithologic descriptions of some boreholes this unit is characteristic for the upper Zbąszynek Beds and lower Wielichowo Beds from the north-western and central parts of Poland (29-33). The deposits exposed at Lisowice are similar to a local informal lithostratigraphic unit named the Lisów Beds (or Lisów Formation) and also to so-called Woźniki Formation, which is exposed in some places near Lubliniec (34). This simple lithostratigraphic correlation is also consistent with biostratigraphic correlation (after palynomorph and conchostracans) of the Lisowice exposure with boreholes from the Polish Lowland and some upper Norian/lowermost Rhaetian sections from Germany (35, 36). The described dicynodont material was collected from the upper bone-bearing interval with carbonaceous greenish and grey fluvial claystone, siltstone and mudstone, interbedded with crossor horizontally-stratified greywacke sandstone or locally conglomerate (Figs. S1, S2). Identification of palynomorph fossils in the grey and organic-rich strata at the Lipie Śląskie claypit (29) suggest correlation of this unit with the Corollina meyeriana Zone, upper part of the Subzone IVb or Subzone IVc, dated as mid-late Norian and latest Norian-earliest Rhaetian respectively (37-39) and/or the lower part of the Ricciisporites tuberculatus Zone (dated as early to late Rhaetian), which was incorporated into the Rhaetipollis germanicus Zone. All these palynozones are well defined based on core material from the central and northern part of Poland 2

(29-33). The upper Zbąszynek Beds apparently continues into the upper Arnstad, upper Löwenstein, Trossingen formations and lower part of the Exter Formation of the upper Middle to Upper Keuper (upper Norian lowermost Rhaetian) in the middle-eastern part of the Germanic Basin (32, 36), which contains the geologically oldest fossils of early mammaliaforms and diversified fauna of early dinosaurs (40-42). In the Polish geological literature, the lithostratigraphic unit represented at the Lipie Śląskie clay-pit has been referred to as Norian (43) or Rhaetian sensu polonico (30-32). Recently, some authors (44, 45) included the bone-bearing deposits from Lisowice to the so-called Lisowice bone-bearing level and suggest mid-norian age of this flora and fauna. This is mainly based on chemostratigraphical correlations (based mostly on Cr/Ti and Cr/Nb ratio trends) with some support of palynological studies (e.g. identification of Corollina meyeriana Subzone IVb in Lipie Śląskie clay-pit). Such an interpretation is contradicted with some well documented paleobotanical finds (see below), biostratigraphic data (e.g. presence of some species of megaspores, remains of a seed-fern Lepidopteris ottonis or an occurrence of conchostracan species Gregoriusella polonica), U/Pb radiometric data collected from Lisowice (see below) and additional observations from other bone-bearing sites temporarily included to the so-called Lisowice bone-bearing level. The dicynodont bone-bearing interval is rich in organic material (e.g. dispersed plant cuticules), macrofossils of plants, fish scales and small bone fragments. The dominant plant species at Lisowice is a conifer similar to Brachyphyllum, Pagiophyllum or Hirmeriella, as is the case with the late Norian-Rhaetian and earliest Jurassic floras of the region (46, 47). The second most common plant species is represented by twigs and seeds closely related to Stachyotaxus septentrionalis (Agardh, 1823), a taxon characteristic for the middle-late Rhaetian of Greenland and Scania (48-50). Other plant fossils, currently examined in detail, are represented by cycadophytes, ginkgophytes, and pteridosperms (49). An early Rhaetian age is suggested for numerous cuticle fragments and rare leaf fragments or fructifications (Peltaspermum rotula Harris, 1937) of the typical Rhaetian seed-fern Lepidopteris ottonis (Goeppert, 1836) (52-56) ; the isoëtalean macrospores Trileites cf. pinguis (Harris, 1935) and Horstisporites bertelseni Fuglewicz, 1977 (57, 58) and the conchostracans Gregoriusella polonica Kozur, Niedźwiedzki et Sulej, 2010, Euestheria sp., and Shipingia sp. collected mainly from layers located below upper bone-bearing interval (29, 59). The species G. polonica is known from the lower Exter Formation of northern Germany (early Rhaetian) and the upper Redonda Formation of New Mexico (latest Norian-early Rhaetian), and this species occurs above the Shipingia olseni Zone, which is correlated with the late Norian Sevatian substage (45, 60). The S. olseni Zone is followed by a short interval that contains both abundant specimens of the very small form G. polonica and the last specimens of S. gerbachmanni; this interval could be either latest Norian or earliest Rhaetian (61). The recently published dating of a single zircon grain, recovered from the sandstone bed below the upper bone-bearing interval at the Lisowice section, as 211 ± 3 Ma (62) also suggests late Norian to earliest Rhaetian age for the Lisowice fauna. The absolute age of crystallization of this youngest zircon indicates the maximum deposition age. This means that the upper bone-bearing interval must be younger than the zircon grain. The boundary between the Norian and Rhaetian stages is defined as being at ~208.5 million years (see Chronostratigraphic Chart of the International Commission on Stratigraphy v. 2018/8). 3

The Lisowice vertebrate assemblage. Well-preserved vertebrate bones occur in two intervals, upper and lower (in total six horizons with bones were identified; on-going study). The bone record in the upper interval is richer than the lower interval, and remains are usually preserved in a lenticular body of clayey, carbonate-rich grey siltstone or mudstone, mostly covered with calcareous and pyritic crust or preserved within magnesium-rich limestone concretions. The vertebrate fossil assemblage from the Lipie Śląskie clay-pit (10, 24, 29, 63-69) consists mainly of terrestrial rather than amphibious or aquatic tetrapods (Fig. S3). The most common bones are of a giant dicynodont (Lisowicia bojani; Fig. 1 and Fig. S2), second in number are archosauromorph and temnospondyl bones. The diversity of small and medium reptilians is relatively high as indicated by the presence of: 1) two species of pterosaurs cranial elements, limb bones, vertebrae and teeth (Pterosauria indet.); 2) two species of dinosauriforms or early dinosaurs cranial elements, vertebrae, limb and pelvic bones (Dinosauriformes indet. or Dinosauria indet.); 3) two or more species of small predatory dinosaurs cranial elements, limb and pelvic bones (Neotheropoda indet.); 4) small crocodylomorph limb bones (Crocodylomorpha indet.); 5) choristodere-like reptile vertebrae and limb bones (Diapsida indet.); 6) lepidosauromorphs skull bones, limb bones and teeth (Sphenodontia indet.) and numerous still unrecognized isolated bones and teeth of other small diapsids (Diapsida indet.). Temnospondyls (Cyclotosaurus sp. and Gerrothorax sp.) are known from isolated skull bones, partial and complete jaws and numerous limb bones or bony dermal elements. The top predator in this assemblage is represented by a 5 to 6 meters long theropod-like archosaur Smok wawelski (63). The fish fauna identified, based on macrofossils, is rich and includes a large coelacanth (skull elements and isolated scales), medium to large dipnoan fish (mainly skull bones and tooth plates of Ptychoceratodus sp.) and diversified actinopterygians (skull bones and numerous scales). The vertebrate microfossils and small fossils are numerically dominated by remains of aquatic vertebrates and are comprised primarily of scales (or other dermal elements) and teeth of Actinopterygii, along with teeth, dermal denticles of sharks of the genera Polyacrodus and Hybodus (64-66). There is a smaller proportion of teeth from different archosaurs and fossils of early anurans in the microfossil record (67). The presence of Rhynchosauria and Poposauridae (29) in the assemblage was not confirmed during the most recent revision and restrictive apomorphy-based identification of the collected bone material. Moreover, the presence of doublerooted, early mammaliaform teeth (68, 69), the diversified assemblage of dinosaurs (four or more species) in the Lisowice fauna further point for the late Norian-earliest Rhaetian nature of this assemblage (10, 29). Systematic paleontology Synapsida Osborn, 1903 Therapsida Broom, 1905 Anomodontia Owen, 1860 Dicynodontia Owen, 1860 Placeriinae King, 1988 Lisowicia gen. nov. Type species. Lisowicia bojani sp. nov. 4

Diagnosis. The dicynodont differs from all other dicynodonts as it possesses the following unique combination of character states, visible in the holotype (ZPAL V.33/96, left humerus): 1) the humerus has a narrower entepicondyle in comparison with other dicynodonts (autapomorphy); 2) the entepicondylar foramen of the humerus is absent (autapomorphy); 3) the supinator process is longer (it is 31% of the total humerus length) than in other dicynodonts. Lisowicia bojani sp. nov. Etymology. Lisowicia, from the name of the village Lisowice where the bones were found; bojani, in honor of Ludwig Heinrich Bojanus (1776 1827), comparative anatomist and paleontologist. Holotype. ZPAL V.33/96 (Institute of Paleobiology, Polish Academy of Sciences, Warsaw), left humerus (Fig. 1A and Fig. S5). Paratypes. Material found associated with the holotype, but are derived from multiple individuals of the same or nearly the same size (Fig. 1B-K and figs. S4, S6-S7). ZPAL V.33/85 (left maxilla); ZPAL MB/18 (frontal with part of prefrontal); ZPAL V.33/741 (parietal); ZPAL V.33/712 (central part of the squamosal); ZPAL V.33/708 (postorbital); ZPAL V.33/717 (left lacrimal); ZPAL V.33/739 (left quadrate); ZPAL V.33/531 (braincase); ZPAL V.33/730 (fragment of pterygoid); ZPAL V.33/735 (posterior part of the right mandible); ZPAL V.33/468 (left scapula); ZPAL V.33/665 (left radius); ZPAL V.33/470 (left ulna); ZPAL V.33/760 (sternum); ZPAL V.33/720 (cervical vertebra); ZPAL V.33/720 (dorsal vertebra); ZPAL V.33/720 (left ilium, pubis and ischium); ZPAL V.33/75 (left femur); ZPAL V.33/75 (left tibia); ZPAL V.33/75 (left fibula). Additional material referable to Lisowicia is under preparation and will be presented in another paper. Locality and horizon. All specimens derive from a terrestrial sequence exposed at Lisowice (Lipie Śląskie clay-pit) about 2 km west of Lubliniec, Silesia, S Poland. The material was collected from the upper bone-bearing interval (Fig. S1) with carbonaceous greenish and grey fluvial claystone, siltstone and mudstone (Fig. S2). Age. Late Norian-earliest Rhaetian, Late Triassic. Extended diagnosis. Lisowicia bojani is the largest known dicynodont with a body length estimated at 4.5 metres or more (the largest scapulocoracoid, specimen V.33/468, has 94 cm length; the largest femur, specimen V.33/652, has 80 cm length). L. bojani differs from all other dicynodonts in possessing unique combination of characters, with some being visible in the holotype specimen (ZPAL V.33/96, left humerus; see in the main text). Others have been recognized in paratype specimens: 1) sternum has the articular surface for the coracoid and the first dorsal rib on the posterior edge (autapomorphy); 2) ridges on dorsal side of the sternum are much high then in any known dicynodont; 3) scapula has proportionally very small acromion process for a dicynodont; 4) scapula has rounded dorsal part of the blade; 5) trunks of cervical vertebrae exhibit deep and oval shaped fossae (autapomorphy). Description. The material of Lisowicia includes the entire pelvis, femur, tibia, fibula, shoulder girdle, humerus, radius, ulna, several vertebrae, ribs, and a number of skull fragments (Figs. 1, 5

2A,B; Figs. S4-7). Lisowicia is not only younger and very much larger than other members of the Middle and Late Triassic Kannemeyeriiformes (the cosmopolitan clade that includes all large Triassic dicynodonts), it is also morphologically very distinctive. This distinctiveness is most evident in the forelimb, which possesses a suite of characters in the scapulocoracoid, humerus, radius and ulna, which indicates a parasagittal limb posture with the humerus tucked into the flank of the animal and the elbow pointing backwards (Fig. 2A, B). The acromion process is strongly reduced in Lisowicia (Fig. 1J). The glenoid of the scapulocoracoid is directed posteroventrally (ZPAL V.33/468), in a similar way to that of Placerias or Ischigualastia, but in contrast to that of dicynodonts from the late Permian and Early to Middle Triassic, which have their glenoids directed posterolaterally. The planes of the distal and proximal ends of the humerus are almost parallel in Lisowicia (ZPAL V.33/96), as in quadrupedal tetrapods with an erect gait. Although an articulated forearm of Lisowicia has not been found, the size of the proximal heads of the radius and ulna, and the articulation areas for them on the humerus (all elements were collected from the same bed and located close to each other), suggest that the forearm was shorter than in earlier dicynodonts. The skeletal morphology implies an equally distinctive musculature where several of the major forelimb muscles had different actions compared to other dicynodonts. Detailed description of selected bones Quadrate. There are two quadrates from two specimens (ZPAL V.33/84 and ZPAL V.33/735 fused with quadratojugal). The articular facet of the right jaw condyle reveals a structure unique for dicynodonts: the medial ridge is much less convex than in Placerias, Jachaleria, and Stahleckeria. Preserved in ZPAL V.33/84, the articular facet for the stapes is very distinct and elongated. Scapula. Five left scapulae are preserved ZPAL V.33/74, V.33/82, V.33/451, V.33/452, and four right V.33/80, 33/425, V.33/726, V.33/761, one left scapulocoracoid V.33/468 (Fig. S4). The largest scapulocoracoid V.33/468 has 94 cm length. The acromion process (well visible in V.33/761) is probably similar to that of Jachaleria candelariensis, where it is broken but its base shows that it was small and occurs in the central part of the scapular blade, similar to the process of Lisowicia. In this species it is triangular (in anterior view) with very small slightly marked additional ridge. The attachment for m. triceps scapularis occurs on the posterior edge of the scapular edge above the glenoid and its surface is very robust. According to Ray and Chinsamy (70) it is the area for attachment of three muscles in Permian dicynodonts. The attachment is distinct on all specimens but never is it so large like in Jachaleria candelariensis (71). The proportion of the scapula width in the distal end and in the narrowest part is small, like in Ischigualastia, but the shape of the acromion process is much different. In Ischigualastia, in the specimen MCZ 3119, the area of scapular spine is well visible. In many papers authors suggest that the acromion process of Ischigualastia is small, whereas the personal studies show that there was a long ridge-scapular spine (72) similar to Stahleckeria. The only species from South America with a scapula having a small acromion process is Jachaleria candelariensis. The anterior border of the scapula is markedly concave. It might relate to increasing of the area in the ventral part of the scapula, above the procoracoid. Procoracoid and coracoid. The coracoid is well preserved and is in articultaion with the scapula in the specimen ZPAL V.33/468. The suture between the coracoid and procoracoid is not visible. The procoracoid is more triangular, contrary to the rather rectangular procoracoid of Placerias. The 6

coracoidal foramen lies near the scapular suture and is bordered dorsally by a notch on the medial face of the scapula. A disarticulated coracoid was also found (ZPAL V.33/728). In that specimen the posteroventral process is well preserved. The posterior end shows a poorly ossified surface for articulation with anther bone element, probably sternum (see description of the sternum). Humerus. The planes of the distal and proximal ends of the humerus are almost parallel in Lisowicia bojani (ZPAL V.33/96; Fig. S5), in contrast to the strongly rotated humeral ends in Permian and Early and Middle Triassic dicynodonts. Both ends of the humerus are in the same plane (the angle between them is 0-5 degrees), as in tetrapods with an erect gait like in mammals and in heavy dinosaurs (e.g. sauropods and ceratopsians). The largest specimens ZPAL MB/24 and ZPAL V.33/479 are incomplete, but their converted length amounts to 61.5 cm. The humerus of L. bojani does not have an entepicondyle foramen in the shaft (this hollow is shallow in some specimens). The complete supinator process is distinctive, and narrow. The humeral head has very large posterior area for the scapula. The articulation areas for the radius and ulna are smaller than in Placerias, and Stahleckeria. The articulation for the ulna is slightly convex compared to Ischigualastia, in which it is strongly protruded from the shaft, which results in limited protraction of the ulna to humerus in Ischigualastia. The distal end of deltopectoral crest is unique, bending frontally, and ventrally it becomes narrower and round. The articulation area for the ulna is small on the posterior side as it is in Ischigualastia. The distance between the deltopectoral crest and supinator process is small. Radius. The radius (ZPAL V.33/665) of Lisowicia bojani differs from those of Permian and Early Triassic dicynodonts in some aspects of the distal end. It probably worked in an anterior position in respect to the distal head of the ulna, as it does in weight-bearing mammals (e.g. the hippopotamus). The distal end is strongly convex, instead of being concave as in Permian and Early Triassic dicynodonts, and the area for articulation is directed posteriorly. The radius of L. bojani has more round heads than in earlier dicynodonts, as Parakannemeyeria youngi, and Kannemeyeria simocephalus, which had radius heads rather flat or concave (personal observation). In L. bojani the proximal head has a very large area for the ulna. It is well visible in lateral view, where it s concave shape can be observed. A similar structure but on the opposite side is visible in Parakannemeyeria youngi (IVPP V.972, left radius). A small ridge is visible on the posterior part of the proximal head of the radius in L. bojani; it is a very distinct structure which might be an area for contact with the posterior process of the head of the Ulna. The distal head is very large and convex. The surface is directed posteriorly so that the surface of the head is not visible in anterior view. On the medial side, a ridge is preserved, probably for attachment of muscle. Ulna. In the left ulna ZPAL V. 33/470, the olecranon process is not ossified with the main body. The lateral surface of the shaft is concave, but the medial surface of the shaft is straight and concave close to the articular surface with the humerus and becomes more convex closer to the distal head. The anterior surface of the shaft at the distal head is flat in cross-section. Although the articulated forearm of Lisowicia bojani was not found, it seems that the proportions of the humerus and forearm were different than those of earlier dicynodonts. Based on the size of the proximal portions of the radius and ulna, and the articulation areas for them on the humerus, the forearm was rather short in L. bojani. The length of the ulna decreased during the evolution of dicynodonts from Sinokannemeyeria, to Placerias, Ischigualastia and Jachaleria, perhaps correlated to increasing body size (personal observation). 7

Sternum. The sternum of Lisowicia bojani is known from three individuals ZPAL V.33/754, 33/759, 33/760 (Fig. S6). The morphology of this element in L. bojani is unique. It has an extremely short posterior part (posterior to the articulation for the coracoid and first rib), similarly to Ischigualastia, Jachaleria and Stahleckeria, but contrary to Placerias. The most characteristic being its extremely large height and the articulation condyles on its posteriolateral corner. All other Triassic dicynodonts have much lower sternum. In L. bojani the articulating surface on the dorsal side of the sternum consists of two parts, merged and visible only in lateral view. The upper condyle was probably for the first dorsal rib like in Trichosurus, and the ventral area was for posterior ventral process of the postcoracoid. The connection was not very accurate; a large amount of cartilage was probably present between these three elements. In the sternum of Dinodontosaurus, Ischigualastia, Jachaleria and Stahleckeria, both articulating surfaces (or condyles) are very distinct, they are not merged and are located in horizontal line (in lateral view) contrary to in the vertical of L. bojani. The dorsal side of the sternum is unsymmetrical and this was observed in all three collected specimens. Details and results of the phylogenetic analysis. To determine the relationships of Lisowicia bojani inside Dicynodontia and Kannemeyeriiformes, we used revised version of the anomodont matrix published by 1) Kammerer et al. (14) and 2) Angielczyk and Kammerer (28). 1) Kammerer et al. (14). A phylogenetic analysis of 99 taxa (including one unnamed and L. bojani; Tab. S2, S3) scored for 174 characters. 153 of these characters are discrete binary or multistate characters (characters unordered and of equal weight); the other 21 characters are continuous. We coded unknown and inapplicable discrete states and continuous characters?. We analyzed the data set using TNT 1.1. Both the new technology searches and the traditional searches discovered the same most parsimonious cladogram (995.264 steps; CI = 0.241; RI = 0.712), and all topological results from the search are summarized in Figs. S8-10. 2) Angielczyk and Kammerer (28). A phylogenetic analysis of 104 taxa (including L. bojani; Tab. S4, S5) scored for 194 characters. 171 are discrete binary or multistate characters, of which 7 characters were ordered and 164 were treated as unordered. All discrete characters were weighted equally. The remaining 23 characters are continuous. We coded unknown and inapplicable discrete states and continuous characters?. We analyzed the dataset using TNT 1.1 and two search strategies were employed (the new technology searches and the traditional searches). Both the new technology searches and the traditional searches discovered the same most parsimonious cladogram (1144.358 steps; CI = 0.239; RI = 0.712), and all topological results from the search are summarized in Figs. S11-13. Lisowicia bojani is reconstructed in both phylogenetic analyses (22, 28) within the clade Placeriinae, and as the sister line of clade comprising of Placerias and Moghreberia (Fig. 3A; Fig. S8-13), both relatively large dicynodonts from the Late Triassic of North America and North Africa, respectively. Histological analysis. We have conducted a histological study of two long bones to determine the growth strategy for Lisowicia bojani. A mid-diaphyseal thin section was made from a femur (ZPAL 8

V.33/763, 90% of maximal size) and a tibia (ZPAL V.33/765, 87% of maximal size). In transverse sections, both bones have a compact cortex surrounding a poorly defined medullary region. The central cavity of the bones is filled with trabeculae (spongiosa), which grade into dense Haversian bone with resorption cavities (Fig. S14a,b). Although the medullary cavity is rather small in both bones, the secondary osteons partially obliterate the growth record in the inner cortex. There is no clear presence of lines of arrested growth (LAGs) in the layer of primary bone of the outer cortex of L. bojani (Fig. S14c). The primary cortical bone in the tibia (ZPAL V.33/765) is generally azonal, without periodic interruptions. In general, this is similar to tissue seen in other Triassic dicynodonts (70-73). The animal is inferred to be adult, as indicated by the size of the sampled bones (each of which represented one of the largest examples from the site), the extensive development of resorption cavities, and the secondary osteons in the inner and outer cortex. The extensive remodeling of the outer cortex was also observed in bones of Placerias and Kannemeyeria (71, 72) The limited histological data currently available preclude additional explanations of the observations made here. Extensive cortical remodeling in L. bojani may be attributed to large body size and age or biomechanical factors. Extensive spongiosa in the medullary cavity are present in most dicynodonts, especially in large sized taxa (71) and is probably phylogenetically informative (71-73). Computed tomography of cervical vertebrae. Only two vertebrae of Lisowicia bojani, both cervical ZPAL V.33/720 and ZPAL V.33/721, have been discovered yet. Both specimens are similar to that known from other dicynodonts except for their deep fossae (=holes) that excavate the lateral walls of the vertebrae, one on each side of the vertebra (Fig. S15). This character was used as autapomorphy. The fossae take about 40 percent of each lateral surface. A large opening that pierces the lateral surface of the centrum, below the suture with neural arch, is situated within the lateral fossa. This opening does not enter any pleurocoel (Fig. S16). CT cross-sections reveal a relatively thin layer of dense external cortical bone. Nearly the entire vertebra, including the transverse processes and centrum, is composed of densely packed trabecular bone with no spaces of pneumatic system (see files Movie S1-S4). Body mass proxy data. We examine body-size evolution in two tetrapod clades over the Middle and Late Triassic. A non-phylogenetic (time series) approach is used to analyze 29 species from the most important herbivore clades of the Middle Late Triassic terrestrial tetrapods (Sauropodomorpha and Dicynodontia). Femoral length was used as a body size proxy because it shows a consistent relationship with body mass in terrestrial tetrapods and has been used in previous studies (20, 74). Sauropodomorph and dicynodont taxa were dated to their geological stage (Tab. S6 and S7) with ranges representing either stratigraphic uncertainty or genuine observed ranges. The results provide evidence for active trends towards higher body sizes in terrestrial herbivores during the Late Triassic. The body-size data clearly show a sustained increase in both sauropodomorphs and dicynodont size over the study interval (Fig. 3B). From the Carnian to the early Norian (235 217.5 Ma), dicynodonts maximum body size is larger than sauropodomorphs maximum body size. Sauropodomorphs and dicynodonts attained very large sizes in the latest Norian-early Rhaetian interval, but sauropodomorphs attained an even larger size just after the extinction of dicynodont in the middle-late Rhaetian. Thus, dicynodonts rapidly increased in size 9

during the latest Triassic and sauropodomorphs increased in size in the Late Triassic time and following extinction of dicynodonts. Body mass estimates. Scaling relationships between skeletal measurements and body mass in extant tetrapods (e.g. birds or mammals) are often used to predict body mass in extinct members. However, the applicability of some models for predicting mass in some Paleozoic or Mesozoic tetrapod taxa, such as non-avian dinosaurs and non-mammalian synapsids, has been criticized on biomechanical grounds. This criticism was rejected and the applicability of a universal scaling equation for estimating body mass in distantly related taxa is valid. Two limb bones were used to estimate the mass of Lisowicia bojani. Both bones, ZPAL V.33/96 (humerus midshaft circumference 41 cm, total length 48 cm) and ZPAL V.33/763 (femur midshaft circumference 45.5 cm, total length 72.3 cm), were found in the same layer and close to each other and this suggests that they represent the same individual. In the collected material are larger bones (ZPAL MB24, humerus with midshaft circumference 46 cm, total length 61.5 cm; ZPAL V.33/652, femur with midshaft circumference 49 cm, total length 80 cm), but it is not clear whether they come from the same individual. Based on results presented by Campione and Evans (14), we propose the following scaling equation as a robust predictor of Lisowicia bojani body mass: logbm = 2.749 logch+f 1.104 log.qe QE lower.qe upper.qe 6.969857 9329477 6938332 11720622 The first value the estimate in logarithmic form, the second is the non-log estimate (in grams). Based on ZPAL V.33/96 and ZPAL V.33/763 measurements the animal is about 9329 kg. The last two numbers are the 25% error range, which is the average error in the living data set on which the equation was based (14). In the opinion of Campione and Evans (14), the range is more important than the point estimate. The range provides a much more accurate depiction of where the mass will be, and so we are encouraged to use it rather than the point QE estimate. Overall, the estimates suggest that this animal is elephant-sized. Provenance All specimens were collected from an upper bone-bearing layer exposed in the Lipie Śląskie claypit at Lisowice near Lubliniec, Silesia, Poland, located at 50 40'43.35"N, 18 38'48.19"E. During 2007-2014, 2017 (nine field seasons, each lasting one month) the excavator was used to remove Quaternary sand and clay, unfossiliferous part of the Upper Triassic layers (uppermost interval with sandstone/claystone intercalations) above bone-bearing deposits (mainly above the upper interval with bones; see Fig. S1). The most productive, upper fossiliferous horizon was searched by professionals and students. The geological and paleontological field researches have been conducted by T. Sulej and G. Niedźwiedzki, with the skillful help of J. Dzik (Warsaw), G. Pieńkowski (Warsaw), P. Brański (Warsaw), M. Błyszcz (Lisowice), K. Owocki (Warsaw), P. Skrzycki (Warsaw) and M. Tałanda (Warsaw). Material is accessioned in the paleontology collection of the Polish Academy of Sciences (Warsaw, Poland) and is housed at the Institute of Paleobiology (Polish Academy of Sciences) and the Department of Paleobiology and Evolution (University of Warsaw) where they may be viewed by prior arrangement with Tomasz Sulej. 10

Fig. S1. Lithostratigraphic section of deposits exposed in the Lipie Śląskie clay-pit at Lisowice, with details about distribution of fossil-bearing and bone-bearing layers (adopted from 29). All materials of the dicynodont were collected from the upper interval with bones. 11

Fig. S2. Lisowicia bojani gen. et sp. nov., hind limb elements (femur, fibula, tibia) preserved in situ, upper bone-bearing interval, Lipie Śląskie clay-pit at Lisowice. 12

Fig. S3. Sketch-drawing of the vertebrate faunal assemblage of the Lisowice site. a, large, predatory theropod-like archosaur (Smok wawelski); b, large temnospondyl amphibian (Cyclotosaurus sp.); c, small predatory dinosaurs (Neotheropoda indet.); d, temnospondyl amphibian (Gerrothorax sp.); e, small basal crocodylomorph (Crocodylomorpha indet.); f, small choristodere-like animal (Diapsida indet.); g, hybodont sharks (Polyacrodus and Hybodus); h, coelacanth fish; i, dipnoan fish (Ptychoceratodus sp.); j, actinopterygian fish; k, gigantic dicynodont (Lisowicia bojani gen. et sp. nov.); l, small diapsids (Diapsida indet.); m, dinosauriforms or early dinosaurs (Dinosauriformes indet. or Dinosauria indet.); n, small lepidosauromorphs (Sphenodontia indet.); o, pterosaurs (Pterosauria indet.); p, early mammaliaform (Hallautherium sp.). 13

Fig. S4. Lisowicia bojani gen. et sp. nov., scapulocoracoid, ZPAL V.33/468, in lateral (a), medial (b), cranial (c) and caudal (d) views. 14

Fig. S5. Lisowicia bojani gen. et sp. nov., humerus, ZPAL V.33/96 (holotype specimen), in cranial (a), caudal (b), medial (c), lateral (d), proximal (e) and distal (f) views. 15

Fig. S6. Lisowicia bojani gen. et sp. nov., sternum, ZPAL V.33/754 (a-d, j), and V.33/759 (e-i), in dorsal (a, e), ventral (b, f), distal (c,h), lateral (i,j), caudal (e) and proximal (d,g) views. 16

Fig. S7. Lisowicia bojani gen. et sp. nov., femur, ZPAL V.33/74, in medial (a), cranial (b), distal (c), lateral (d), caudal (e) and proximal (f) views. 17

Tree 0: Biarmosuchus Hipposaurus Fig. S8. Gorgonops Glanosuchus Lycosuchus Titanophoneus Archaeosyodon Biseridens Tiarajudens Anomocephalus Patranomodon Suminia Ulemica Otsheria Galechirus Galepus Galeops Eodicynodon_oelofseni Colobodectes Eodicynodon_oosthuizeni Lanthanostegus Chelydontops Endothiodon Pristerodon Diictodon Eosimops Robertia Prosictodon Emydops Kombuisia Dicynodontoides Myosaurus Cistecephalus Kawingasaurus Cistecephaloides Keyseria_benjamini Daqingshanodon_limbus Oudenodon_bainii Australobarbarus Tropidostoma Idelesaurus Odontocyclops Kitchinganomodon Rhachiocephalus Syops_vanhoepeni Aulacephalodon Pelanomodon Geikia_elginensis Geikia_locusticeps Interpresosaurus Katumbia Elph Gordonia_traquairi Basilodon_woodwardi Sintocephalus_alticeps Dicynodon_lacerticeps Dicynodon_huenei Vivaxosaurus_trautscholdi Delectosaurus Dinanomodon_gilli Peramodon_amalitzkii Daptocephalus_leoniceps Jimusaria_sinkiangensis Turfanodon_bogdaensis Euptychognathus_bathyrhynchus Lystrosaurus_murrayi Lystrosaurus_declivis Lystrosaurus_curvatus Lystrosaurus_maccaigi Lystrosaurus_hedini TSK_2 Kwazulusaurus_shakai Angonisaurus Rhinodicynodon Shansiodon Vinceria Tetragonias Dinodontosaurus Shaanbeikannemeyeria Kannemeyeria_lophorhinus Kannemeyeria_simocephalus Xiyukannemeyeria Parakannemeyeria Dolichuranus Uralokannemeyeria Rechnisaurus Rhadiodromus Sinokannemeyeria Wadiasaurus Rabidosaurus Lisowicia Moghreberia Placerias Zambiasaurus Ischigualastia Jachaleria Eubrachiosaurus Sangusaurus Stahleckeria A phylogenetic analysis based on Kammerer et al. (14). 99 taxa (including one unnamed and L. bojani; see Tab. S2, S3) scored for 174 characters. Most parsimonious cladogram. Traditional search method in TNT (TBR branch swapping with 10.000 replicates and 10 trees held per replicate; one most parsimonious tree; CI = 0.241; RI = 0.712; tree length 995.264). Analysis with New technology method in TNT (identical to that presented in Kammerer et al. (22) gives the same tree. 18

Strict consensus of 10 trees (0 taxa excluded) Biarmosuchus Hipposaurus Fig. S9. Gorgonops Titanophoneus Glanosuchus Lycosuchus Biseridens Archaeosyodon Galepus Patranomodon Tiarajudens Anomocephalus Suminia Ulemica Otsheria Galechirus Galeops Eodicynodon_oelofseni Colobodectes Eodicynodon_oosthuizeni Lanthanostegus Chelydontops Endothiodon Pristerodon Diictodon Eosimops Robertia Prosictodon Emydops Kombuisia Dicynodontoides Myosaurus Cistecephalus Kawingasaurus Cistecephaloides Keyseria_benjamini Daqingshanodon_limbus Oudenodon_bainii Australobarbarus Tropidostoma Idelesaurus Odontocyclops Kitchinganomodon Rhachiocephalus Syops_vanhoepeni Aulacephalodon Pelanomodon Geikia_elginensis Geikia_locusticeps Katumbia Interpresosaurus Elph Gordonia_traquairi Delectosaurus Vivaxosaurus_trautscholdi Dicynodon_huenei Dicynodon_lacerticeps Dinanomodon_gilli Peramodon_amalitzkii Daptocephalus_leoniceps Jimusaria_sinkiangensis Turfanodon_bogdaensis Euptychognathus_bathyrhynchus TSK_2 Basilodon_woodwardi Sintocephalus_alticeps Lystrosaurus_murrayi Lystrosaurus_declivis Lystrosaurus_curvatus Lystrosaurus_maccaigi Lystrosaurus_hedini Kwazulusaurus_shakai Rhinodicynodon Shansiodon Tetragonias Angonisaurus Vinceria Dinodontosaurus Shaanbeikannemeyeria Kannemeyeria_lophorhinus Kannemeyeria_simocephalus Xiyukannemeyeria Parakannemeyeria Dolichuranus Uralokannemeyeria Rechnisaurus Rhadiodromus Sinokannemeyeria Wadiasaurus Rabidosaurus Lisowicia Moghreberia Placerias Zambiasaurus Ischigualastia Jachaleria Eubrachiosaurus Sangusaurus Stahleckeria A phylogenetic analysis based on Kammerer et al. (14). 99 taxa (including one unnamed and L. bojani; see Tab. S2, S3) scored for 153 discrete characters. Traditional search method in TNT (strict consensus of 10 trees). 19

Group freqs., 15000 replicates, cut=50 (tree 0) - Symmetric Resampling (P=33) Biarmosuchus Hipposaurus Gorgonops 100 Biseridens Titanophoneus 91 Archaeosyodon Glanosuchus 99 81 Lycosuchus Tiarajudens 86 Anomocephalus Galechirus 85 Galepus Patranomodon Suminia 55 79 Ulemica 71 Otsheria Galeops Eodicynodon_oelofseni 74 Colobodectes Eodicynodon_oosthuizeni Lanthanostegus 97 Endothiodon Chelydontops Pristerodon 95 Robertia Diictodon 61 Prosictodon Eosimops Emydops 71 Kombuisia 69 62 Dicynodontoides 59 Myosaurus 57 Cistecephalus 98 Kawingasaurus 84 Cistecephaloides TSK_2 Syops_vanhoepeni Euptychognathus_bathyrhynchus Gordonia_traquairi Keyseria_benjamini Turfanodon_bogdaensis 56 Basilodon_woodwardi Sintocephalus_alticeps Jimusaria_sinkiangensis Vivaxosaurus_trautscholdi Peramodon_amalitzkii Dinanomodon_gilli Daqingshanodon_limbus Daptocephalus_leoniceps Dicynodon_huenei Dicynodon_lacerticeps Delectosaurus Katumbia Interpresosaurus Elph Idelesaurus Odontocyclops Australobarbarus Tropidostoma Oudenodon_bainii 79 Kitchinganomodon 69 Rhachiocephalus Geikia_elginensis 72 Geikia_locusticeps 86 Pelanomodon 51 Aulacephalodon Lystrosaurus_murrayi Lystrosaurus_declivis Lystrosaurus_curvatus 67 Lystrosaurus_maccaigi Lystrosaurus_hedini Kwazulusaurus_shakai Lisowicia Shaanbeikannemeyeria Zambiasaurus Eubrachiosaurus Uralokannemeyeria Sangusaurus Rechnisaurus Wadiasaurus Rhadiodromus Rabidosaurus Stahleckeria Dinodontosaurus Angonisaurus 63 Rhinodicynodon Vinceria Shansiodon Tetragonias Dolichuranus Sinokannemeyeria Moghreberia 75 Placerias Kannemeyeria_lophorhinus 51 Kannemeyeria_simocephalus Ischigualastia 77 Jachaleria Xiyukannemeyeria 53 Parakannemeyeria Fig. S10. A phylogenetic analysis based on Kammerer et al. (14). Bootstrap resampling of the analysis of all characters, 15000 replicates, nodes below 50% support collapsed. 20

Fig. S11. A phylogenetic analysis based on Angielczyk and Kammerer (28). 104 taxa (including L. bojani; Tab. S4, S5) scored for 194 characters. Most parsimonious cladogram. Traditional search method in TNT (TBR branch swapping with 10.000 replicates and 10 trees held per replicate; one most parsimonious tree; steps; CI = 0.239; RI = 0.712; tree length 1144.358). 21

Fig. S12. A phylogenetic analysis based on Angielczyk and Kammerer (28). 104 taxa (including L. bojani; Tab. S4, S5) scored for 171 discrete characters. Traditional search method in TNT (strict consensus of 650 trees). 22

Fig. S13. A phylogenetic analysis based on Angielczyk and Kammerer (28). Bootstrap resampling of the analysis of all characters, 15000 replicates, nodes below 50% support collapsed. 23

Fig. S14. Bone histology of the Lisowicia bojani gen. et sp. nov. tibia (ZPAL V.33/765). a, b, showing high vascularization in the cortex (COR), which decreases slightly at the sub-periosteal surface (RC area of bone resorption; MC medullary cavity). c, two possible annuli (LAG, with arrows) can be seen interrupting the outer cortex. Scale bar = 1000 μm. 24

Fig. S15. 3D visualizations of CT scan data of cervical vertebrae (a-c, ZPAL V.33/720; d-f, ZPAL V.33/721) of Lisowicia bojani gen. et sp. nov. (reconstructed in Mimics version 15.01 by Materialize in Belgium, http://biomedical.materialise.com/mimics ). a, f, lateral views of the right side; b, c, virtual sections; c, d, lateral views of the left side. Abbreviations: ho hole (=fossa); nc neural canal. Fig. S16. 3D visualizations of CT scan data of cervical vertebra (ZPAL V.33/721) of Lisowicia bojani gen. et sp. nov. (reconstructed in Mimics version 15.01 by Materialise in Belgium, http://biomedical.materialise.com/mimics ). a, anterolateral view; b, anterior view, c, ventral view. Abbreviations: ho hole; nc neural canal; red structures hollow spaces located inside the vertebra centrum. 25

Table S1. Short characteristics of other Lisowicia bojani gen. et sp. nov. cranial and postcranial bones. ZPAL V.33/85 left maxilla The long ventral process of maxilla is very thick and rounded. The dorsal surface forms suture for the lacrimal, the suture for the jugal is also visible laterally ZPAL MB/18 part of skull The elongated frontal is well visible, with broken prefrontal, roof ZPAL V.33/741 parietal The bone is wide like in Placerias in dorsal view and triangle in lateral view. ZPAL V.33/712 part of the Very thin fragment of bone, nothing else visible squamosal ZPAL V.33/708 postorbital The postorbital is very large, especially the part with contact to the frontal, and very wide at the level of the orbit ZPAL V.33/717 left lacrimal The bone is large in at the posterior (the face for the jugal is very thick). ZPAL V.33/739 left quadrate The quadrate is large and the area for the quadratojugal is also large. ZPAL V.33/531 braincase Most of the bones creating the braincase are well ossified and large ZPAL V.33/730 fragment of Ventral margin and lateral side is preserved only. The ventral edge is ZPAL V.33/735 ZPAL V.33/720 pterygoid right posterior part of the mandible almost complete vertebral column with pelvic girdle strongly concave in lateral view. The dorsal edge forms a distinct triangle structure well visible in lateral view in front of the articular. The angular has a strongly concave central ventral edge. The prearticular has a concave central edge and rectangular fenestra for the Meckelian cartilage. The articular is situated strongly above surangular. The cervical vertebrae have low height neural arches, especially in comparison to the dorsal ones. The dorsal vertebrae of Lisowicia are joined to the articular surface for rib (diapophysis and parapophysis). Sacral vertebrae (4) are not ossified with neural arches and are broken in the spine. The anterior half of the iliac blade is very large and strongly curved laterally. The anterior edge of the pubis is slightly concave in lateral view and covered by long ridges. The lateral edge of the ischium is strongly curved and the glenoid is very deep. The ridge on the ventral blade is significant. ZPAL V.33/453 right radiale The bone is rather rectangular with a large hole at the articulation with the ulnare. ZPAL V.33/744 last phalanx The digit is flat with a waist before the area for articulation with previous digit. ZPAL V.33/763, ZPAL V.33/652 femur Spherical proximal head is dorsalomedially directed. Based on the proportions of complete ZPAL V.33/763 (length 72.3 cm) the partially preserved ZPAL V.33/652 had length around 80 cm. ZPAL V.33/467 tibia The bone is very large, with morphology similar to Placerias. The specimen has a length 48.5 cm. ZPAL V.33/76 left fibula The medial side of the distal head is straight (in anterior view) like in Placerias. 26

Table S2. 21 continuous (morphometric) characters (see Kammerer et al. 22). Lisowicia bojani?????????????? 0.517 0.576 0.522 0.764 0.963 0.438 0.457 Table S3. 153 discrete state characters (see Kammerer et al. 22). Lisowicia bojani??????????0?10???????1?11?000??????111111121?0?1121?1??22???1?????????????????????1??2220?11?1??11????????????????111111?000100001200101????10?1111????? Table S4. 23 continuous (morphometric) characters (see Angielczyk and Kammerer, 28). Lisowicia bojani???????????????? 0.517 0.576 0.522 0.764 0.963 0.438 0.457 Table S5. 171 discrete state characters (see Angielczyk and Kammerer, 28). Lisowicia bojani????????????0?20?????221??1?0?0??????1?11111201?0??120?01??1?1??????1???????????????????????1?????1?220?????1011?????????????????????0011111?11?100001200101???0001111????? 27

Table S6. Femur lengths of sauropodomorphs from the Middle Triassic to the Late Triassic. Based on data sets published by Sookias et al. (20); Kubo and Kubo (74) and personal observations. Genus Species Clade Femur length (mm) Age Camelotia borealis Sauropodomorph 1008 Rhaetian Thecodontosaurus antiquus Sauropodomorph 210 Rhaetian Pantydraco caducus Sauropodomorph 72 Rhaetian Antetonitrus ingenipes Sauropodomorph 794 late Norian-Rhaetian Isanosaurus attavipachi Sauropodomorph 760 late Norian-Rhaetian Plateosauravus cullingworthi Sauropodomorph 600 late Norian-Rhaetian Plateosaurus engelhardti Sauropodomorph 930 late Norian-Rhaetian Riojasaurus incertus Sauropodomorph 608 late Norian-Rhaetian Melanorosaurus readi Sauropodomorph 620 late Norian-Rhaetian Efraasia minor Sauropodomorph 627 late Norian Plateosaurus longiceps Sauropodomorph 610 late Norian Plateosaurus quenstedti Sauropodomorph 700 late Norian Ruehleia bedheimensis Sauropodomorph 800 late Norian Guaibasaurus candelariensis Sauropodomorph 214 late Norian Lessemsaurus sauropoides Sauropodomorph 780 late Norian Plateosaurus gracilis Sauropodomorph 543 late Norian Saturnalia tupiniquim Sauropodomorph 157 late Carnian-early Norian Chromogisaurus novasi Sauropodomorph 160 late Carnian Panphagia protos Sauropodomorph 190 late Carnian. 28

Table S7. Femur lengths of dicynodonts from the Middle Triassic to the Late Triassic. Based on data sets published by Sookias et al. (20); Kubo and Kubo (74) and personal observations. Genus Species Clade Femur length (mm) Age Lisowicia bojani Dicynodontia 800 late Norian-early Rhaetian Lisowicia bojani Dicynodontia 730 late Norian-early Rhaetian Lisowicia bojani Dicynodontia 560 late Norian-early Rhaetian Jachaleria candelariensis Dicynodontia 310 early Norian Placerias hesternus Dicynodontia 350 early Norian Ischigualastia jenseni Dicynodontia 380 late Carnian-early Norian Ischigualastia jenseni Dicynodontia 300 late Carnian-early Norian Dinodontosaurus turpior Dicynodontia 175 Ladinian Stahleckeria potens Dicynodontia 455 Ladinian Kannemeyeria simocephalus Dicynodontia 152 Anisian Kannemeyeria simocephalus Dicynodontia 355 Anisian Parakannemeyeria dolichodephala Dicynodontia 378 Anisian Parakannemeyeria youngi Dicynodontia 380 Anisian Tetragonias njalilus Dicynodontia 185 Anisian Tetragonias njalilus Dicynodontia 195 Anisian 29

Movie S1. 3D visualizations of CT scan data of cervical vertebra of Lisowicia bojani, ZPAL V.33/720, external view. Movie S2. 3D visualizations of CT scan data of cervical vertebra, ZPAL V.33/720, internal view. Movie S3. 3D visualizations of CT scan data of cervical vertebra of Lisowicia bojani, ZPAL V.33/721, external view. Movie S4. 3D visualizations of CT scan data of cervical vertebra, ZPAL V.33/721, internal view. 30

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