Stenoscelida aurantiacus Müller &amp; Garcia &amp; Fonseca 2022, gen. et sp. nov.
Creators: name: Müller, Rodrigo Temp
name: Garcia, Mauricio Silva
name: Fonseca, André de Oliveira
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Reptilia
Crocodylia
Proterochampsidae
Stenoscelida
Stenoscelida aurantiacus

description: <i>Stenoscelida aurantiacus</i> gen. et sp. nov. (Figs 1–5) <b>Holotype.</b> CAPPA/UFSM 0293, a complete and articulated right hind limb. <b>Etymology.</b> The genus combines the Greek words UIƐvόç (<b>¼</b> narrow) and U&amp;kgreen;Ɛƛ <i>o</i> ç (<b>¼</b> hind leg), referring to the slender leg of the creature. The specific epithet derives from the Latin word <i>aurantiacus</i> (<b>¼</b> orange), in allusion to the orange colour of the outcropping sediments of the Varzea do Agudo site (Fig. 1B). <b>Type locality.</b> Varzea do Agudo site (<b>¼</b> Janner site; 29 <b> °</b> 39 <b> 0</b> 10.89 <b> 00</b> S, 53 <b> °</b> 17 <b> 0</b> 34.20 <b> 00</b> W), Agudo, Rio Grande do Sul, Brazil (Fig. 1). <b>Stratigraphic horizon.</b> Lower portion of the Candelaria Sequence (Horn <i>et al.</i> 2014) of the Santa Maria Supersequence (Zerfass <i>et al.</i> 2003), Parana Basin. The predominance of the cynodont genus <i>Exaeretodon</i> places the site in the upper part (<i>Exaeretodon</i> sub-Assemblage Zone; Muller &amp; Garcia 2020) of the <i>Hyperodapedon</i> Assemblage Zone (Schultz <i>et al.</i> 2020), which is biostratigraphically correlated with the <i>Exaeretodon</i> biozone from the Ischigualasto Formation (Mart&amp;imath;nez <i>et al.</i> 2013). A Bayesian age-model for the profile of the Ischigualasto Formation at the Hoyada del Cerro Las Lajas locality in Argentina recovered an age of 227.94 <b>+</b> 0.83/ <b>¯</b> 1.67 for the top of the <i>Hyperodapedon</i> AZ (Desojo <i>et al.</i> 2020), indicating an age of <i>c</i>. 228 Ma for the <i>Exaeretodon</i> biozone. A similar age (late Carnian/early Norian, Late Triassic) for the <i>Exaeretodon</i> sub-Assemblage Zone is inferred in the Candelaria Sequence. <b>Ontogenetic assessment.</b> The presence of some muscle attachment structures (e.g. anterior trochanter; anterolateral scar) in the femur of CAPPA/UFSM 0293 suggests some degree of development if the ontogenetic pathways of other archosauriforms are considered (Griffin &amp; Nesbitt 2016). However, it is currently not possible to confirm whether the specimen reached its maximum size or not. <b>Diagnosis.</b> <i>Stenoscelida aurantiacus</i> differs from all other known proterochampsids in (<b>*</b> local autapomorphies): possessing a slender femur; presence of an anterior trochanter on the femoral head; presence of a raised anterolateral scar above the anterior trochanter; fourth trochanter restricted to the proximal half of the femur; tibia approximately 84% the total length of the femur; in proximal view, the cnemial crest of the tibia is subequal in size to the proximal condyles; fibula with an iliofibularis tubercle on the proximal portion of the shaft <b>*</b>; proximal third of the fibular shaft is anteroposteriorly expanded, tapering distally <b>*</b>; ratio between the minimum midshaft width of metatarsal II and its total length is 0.12; ratio between femoral length and metatarsal III is 2.5; and digit V with a phalanx <b>*</b>. Description and comparison Overview The preserved right hind limb of the holotype is entirely articulated (Fig. 2). Some portions of the specimen are covered by a thick concretion layer, especially the femoral midshaft and the ankle. The external surface is wellpreserved in several places, revealing some muscle attachment points. However, the specimen was deformed by sedimentary compression. Therefore, the elements are lateromedially compressed, affecting the original shape. As result, the midshaft of the limb bones is collapsed, showing artificial longitudinal grooves or sulci. Part of the fibular midshaft is not preserved. The same is true for the distal portion of metatarsal IV, precluding us from determining its total length. According to the femoral length (147 mm), the specimen is smaller than the Argentinian taxa from the Ischigualasto Formation: <i>Pseudochampsa ischigualastensis</i> (155 mm; Trotteyn &amp; Ezcurra 2014) and <i>Proterochampsa barrionuevoi</i> (179 mm; Trotteyn 2011). Femur The femur (Fig. 3) is sigmoid in lateral and medial views. The extremities are well ossified and expanded. The femoral head is mainly anteriorly directed, whereas in other proterochampsids it is more medially oriented. Diagenetic processes might have exaggerated this condition. The general morphology of the femoral head resembles that of proterochampsids, whereas it differs from the anteromedially expanded femoral head of dinosaurs (Nesbitt 2011) and from the hook-shaped head of lagerpetids and pterosaurs (Ezcurra <i>et al.</i> 2020a). The proximal articular surface bears a shallow straight groove (Fig. 3C), which is usually absent in proterochampsids, except in an unnamed rhadinosuchine (Ezcurra <i>et al.</i> 2019; CRILAR-Pv 491) from the Chanares Formation. There is an anterior tuber (<i>sensu</i> Ezcurra 2016) on the proximal portion of the femur, and although the bone lacks a posteromedial tuber, the posterior tuber is present. The greater trochanter is rounded and tall (Fig. 3B), differing from the typical angled trochanter of dinosaurs, and in the same way, there is no dorsolateral trochanter on the proximal portion of the bone. Conversely, the anterolateral surface bears a raised and rugose area (Fig. 3B) where the anterior (<b>¼</b> lesser) trochanter is reported for a number of archosauriforms (e.g. ornithosuchids, aetosaurs, dinosauriforms). Whereas the anterior trochanter is absent in several proterochampsids (Trotteyn <i>et al.</i> 2013), the structure occurs in the holotype of <i>Gualosuchus reigi</i> (PULR-V 05; M. D. Ezcurra, pers. comm.). A trochanteric shelf is not associated with the anterior trochanter, and the proximal-most portion of this trochanter is completely connected to the shaft. Slightly above the proximal tip of the anterior trochanter there is an additional scar, which resembles the anterolateral scar (<i>sensu</i> Griffin &amp; Nesbitt 2016) of aphanosaurs and dinosauriforms. There is a similar scar in <i>Gualosuchus reigi</i> (PULR-V 05; M. D. Ezcurra, pers. comm.). The transition from the femoral head to the shaft is smooth. The fourth trochanter rests on the posterior surface of the proximal third of the femur (Fig. 3A). It is crest-like and symmetrical in shape. Its medial surface is ornamented with muscle scars. The distal portion of the fourth trochanter does not extend further down along the femoral shaft. In contrast, the fourth trochanter is strongly proximodistally developed in <i>Gualosuchus reigi</i> and <i>Chanaresuchus bonapartei</i> (Ezcurra <i>et al.</i> 2019). The midshaft is slender. The robustness index (RI, <i>sensu</i> Wilson &amp; Upchurch 2003; i.e. average of the greatest widths of the proximal end, midshaft and distal end of the element divided by the length of the element) is 0.14. For comparison, this is slightly slenderer than that of some early dinosaurs, such as <i>Gnathovorax cabreirai</i> (0.16) and <i>Buriolestes schultzi</i> (0.15; Muller &amp; Garcia 2022). The anterior surface of the distal portion bears an extensor fossa (Fig. 3D), which results in a concave anterior margin in distal view. On the opposite side, the popliteal fossa is well delimited (Fig. 3E). It is proximodistally short, approximately 10% of the total length of the bone. This condition distinguishes the specimen from silesaurids and aphanosaurs, where the fossa is considerably longer (Nesbitt <i>et al.</i> 2010, 2017). The distal condyles are approximately at the same level in anterior or posterior views. In addition, the lateral surface between the lateral condyle and the crista tibiofibularis is smooth, lacking a deep groove. Tibia The tibia (Fig. 4) is 84% the total length of the femur (Table 1). This resembles the condition in <i>Pseudochampsa ischigualastensis</i>, where it is approximately 82% (Trotteyn &amp; Ezcurra 2014), and differs from both <i>Proterochampsa barrionuevoi</i> (74.5%; Trotteyn 2011) and <i>Tropidosuchus romeri</i> (100%; Arcucci 1990). The bone is straight in anterior/posterior or medial/lateral views. The proximal portion is strongly expanded and is triangular in shape in proximal view (Fig. 4C). The anterior half of the proximal margin is proximally projected in medial view. In medial or lateral views, the well-developed cnemial crest extends anteriorly, and in proximal view it is straight and the anterior margin is rounded. The cnemial crest is sub-equal in size regarding the proximal condyles, distinguishing the new species from <i>Proterochampsa barrionuevoi</i>, where the crest is proportionally smaller. The posterior and lateral condyles are sub-equal in size. The lateral condyle is offset regarding the posterior condyle. The posterior condyle does not taper posteriorly. The presence of any crest on the lateral surface of the proximal portion of the bone is uncertain. The midshaft is ovoid in cross-section. The bone wall is thick, distinct from the thin walls of several pan-avians and some crocodylomorphs (Kellner <i>et al</i>. 2022). The distal portion of the bone is moderately expanded. It lacks a posterolateral process. There is a proximodistally oriented groove on the lateral surface of the distal portion (Fig. 4F). It is more pronounced in <i>Proterochampsa barrionuevoi</i> (Trotteyn 2011). The distal outline of the tibia is elliptical. It is anteroposteriorly longer than transversely wide. <i>·</i> Fibula The total length of the fibula (Fig. 4) is uncertain because a portion of the midshaft is not preserved. The bone is straight in anterior or lateral views, and the proximal articular surface is anteroposteriorly expanded and transversely compressed (Fig. 4C). It differs from the rounded to elliptical proximal end of some pseudosuchians (e.g. <i>Dynamosuchus collisensis</i>, <i>Prestosuchus chiniquensis</i>). In proximal view, the lateral margin is convex but the medial margin is concave. In lateral view, the posterior margin projects posteriorly, whereas the anterior margin lacks any anterior expansion. Conversely, in non-proterochampsid proterochampsians (e.g. <i>Vancleavea campi</i>; <i>Litorosuchus somnii</i>; <i>Jaxtasuchus salomoni</i>) the proximal portion is symmetrical to nearly symmetrical in lateral view. The posterior margin of the proximal third of the bone is sharp. The proximal third of the midshaft is anteroposteriorly expanded, being wider than the tibia (Fig. 4F). Conversely, the shaft becomes extremely slender for the next two-thirds, being approximately two times narrower than the tibia. This is an unusual condition and distinguishes the specimen from other proterochampsids. Furthermore, there is an iliofibularis tubercle on the proximal portion of the shaft (Fig. 4E, F), a feature absent in other proterochampsids. The condition in <i>Stenoscelida aurantiacus</i> differs from the more distally located tubercle of several pseudosuchians and rhynchosaurs. The distal portion of the fibula is gently expanded. It is approximately 0.45 times the maximum length of the proximal articular surface (Table 1). There is a faint longitudinal ridge running on the anterolateral margin. The distal articular surface is flat to concave. In distal view (Fig. 4D), it is ovoid to triangular. Proximal tarsals The astragalus and calcaneum are poorly preserved (Figs 2, 5). The astragalus is transversely wider (20.5 mm) than anteroposteriorly long (11 mm). The anterior margin of the astragalus is concave in dorsal view (Figs 2B, 5B). A transverse groove runs on the posterior surface of the bone. The presence or absence of foramina on this posterior groove is uncertain because it is badly preserved. As in other proterochampsids, the tibial facet is wider than the fibular facet. The latter facet is dorsolaterally oriented, resembling the condition of <i>Chanaresuchus bonapartei</i> and <i>Proterochampsa barrionuevoi</i>, whereas in <i>Pseudochampsa ischigualastensis</i> it is dorsally oriented (Trotteyn &amp; Ezcurra 2014). The ventral surface of the bone is transversely convex. The calcaneum is 10 mm in width (approximately half of the astragalar width). There is a posterolaterally oriented calcaneal tuber. This structure is sub-rectangular, with a straight lateral margin. In dorsal view, the posterolateral corner of the calcaneal tuber is rounded (Fig. 5B). Metatarsals As in other proterochampsids, the metatarsals (Fig. 5) overlap each other. Metatarsal I is shorter than metatarsals II–IV (Fig. 5C). It is almost three times shorter than metatarsal III (Table 1). The proximal articular surface is transversely expanded. There is a longitudinal smooth crest on the dorsomedial corner of the proximal half of the bone. The distal end is moderately expanded. The lateral condyle is more pronounced dorsally than the medial one. There is a shallow extensor fossa on the dorsal surface of the distal portion. The presence of collateral ligament pits on the condylar sides is uncertain. The distal articular surface is not strongly ginglymoid. Metatarsal II is the stoutest metatarsal. The proximal articular surface is strongly expanded (Table 1). The ratio between the proximal articular surface and the total length is 0.39. It is 0.36 in <i>Rhadinosuchus gracilis</i> (Ezcurra <i>et al.</i> 2015). Conversely, the shaft is stouter in <i>Rhadinosuchus gracilis</i>, where the ratio between the minimum midshaft width and the total length of metatarsal II is 0.17. In <i>Stenoscelida aurantiacus</i> it is 0.12. On the dorsal surface of the distal portion, there is a triangular extensor fossa. It is not possible to determine if the articular surface is ginglymoid. There is no evidence of a collateral ligament pit on the medial surface of the medial condyle. On the other hand, the lateral surface of the lateral condyle bears a well-delimited collateral ligament pit (Fig. 5B). In dorsal view, the lateral condyle is straight, whereas the medial condyle is slightly medially oriented. Both condyles are equally expanded distally, which is distinct from <i>Rhadinosuchus gracilis</i>, where the lateral condyle is far more extended distally (Ezcurra <i>et al.</i> 2015). Metatarsal III is the longest (Table 1), whereas its midshaft is slenderer than that of metatarsal II. Its proximal articular surface is moderately expanded (maximum proximal width is 14 mm), far less than the proximal articular surface of metatarsal II (19 mm). The distal articular surface resembles that of metatarsal II, with a marked extensor fossa on the dorsal surface and a collateral ligament pit on the lateral surface of the lateral condyle (Fig. 5B). The presence of the collateral ligament pit on the medial condyle is uncertain (it is covered by thick sediment). The lateral condyle is slightly more pronounced laterally than the lateral condyle of metatarsal II. Metatarsal IV is incomplete (Fig. 5B). The element does not preserve the distal portion. Therefore, its total length is uncertain. The preserved part is 30 mm in length. Despite its condition, the element is longer than metatarsals I and V. The proximal articular surface is not transversely expanded. The general morphology of the element is simple. It comprises an elongated and compressed shaft. Distinct from the previous metatarsals, metatarsal IV is plate-like in cross-section. Metatarsal V is the shortest (Fig. 5B), being slightly shorter than phalanx 1 of digit I. In this sense, <i>Stenoscelida aurantiacus</i> differs from <i>Pseudochampsa ischigualastensis</i>, where metatarsal V is slightly longer than the phalanx 1 from digit I (Trotteyn &amp; Ezcurra 2014). The proximal articular surface is moderately expanded and ‘U’-shaped in ventral view. The shaft tapers distally to an unexpanded and featureless distal end, which bears no distal condyles. Phalanges The phalangeal formula is 2-3-4-?-1 (Fig. 5). Other proterochampsids lack phalanges in digit V. The first phalanx of digit I is longer than broad. This is the same pattern observed in the other phalanges of the specimen. This phalanx is shorter than the first phalanx from digits II and III (Table 2). The dorsal intercondylar process is moderately developed and the distal articular surface is ginglymoid. There is an extensor depression on the dorsal surface of the distal portion. This surface receives the extensor tubercle from the ungual phalanx. The ungual (distal-most) phalanx of digit I lacks its distal half. The phalanx is transversely compressed and tapers distally (at least, the preserved portion). The flexor tubercle is poorly developed (Fig. 5A). The first phalanx of digit II is similar in shape to that of digit I. However, the phalanx from digit II is larger (Fig. 5C). Phalanx 2 is slightly smaller. Its anatomy is obscured by a thick layer of concretion. The ungual phalanx of digit II is clearly the largest ungual. It is transversely compressed and tapers to a sharp point. The ungual is not ventrally recurved and lacks a well-developed flexor tubercle. Digit III is the longest (Fig. 5B). The first phalanx from digit III is sub-equal in length to the first phalanx of digit II (Table 2). On the other hand, the shaft of this phalanx is more gracile. The extensor fossa is deep on the dorsal surface of the distal portion. The subsequent phalanges of this digit are smaller. The ungual is remarkably smaller than that of digit II. The number of phalanges in digit IV is unknown and the distal portion of metatarsal IV is not preserved. Therefore, the absence of phalanges is ambiguous. The phalanx of digit V (Fig. 5A) is reduced to an elongated structure with a concave proximal articular surface. No condyles or other structures are present in this vestigial element. Phylogenetic results The phylogenetic analysis recovered 39,200 most parsimonious trees (MPTs) of 6287 steps (consistency index <b>¼</b> 0.18375; retention index <b>¼</b> 0.63457). <i>Stenoscelida aurantiacus</i> nests within Proterochampsidae in all MPTs (Fig. 6A). The clade is supported by 19 synapomorphies, of which three are coded for <i>Stenoscelida aurantiacus</i>: 1) midshaft diameter of metatarsal II is more than the midshaft diameter of metatarsal I (ch. 572: 0 <b>!</b> 1); 2) midshaft diameter of metatarsal IV is lower than that of metatarsal III (ch. 573: 0 <b>!</b> 1); and 3) crest-like fourth trochanter, dorsoventrally/anteroposteriorly taller than or equal to the shaft at its minimum depth (ch. 803: 1 <b>!</b> 2). Proterochampsidae is the sister group to Doswelliidae, which is composed of <i>Rugarhynchus sixmilensis</i>, <i>Sphodrosaurus pennsylvanicus</i>, <i>Doswellia kaltenbachi</i> and <i>Jaxtasuchus salomoni</i>. The clade supporting Proterochampsidae <b>+</b> Doswelliidae is in a trichotomy with <i>Polymorphodon adorfi</i> and an unnamed clade composed of <i>Vancleavea campi</i> and <i>Litorosuchus somnii.</i> These clades are included within Proterochampsia. Regarding the internal relationships of Proterochampsidae, there is a polytomy composed of <i>Stenoscelida aurantiacus</i>, <i>Tropidosuchus romeri</i>, <i>Cerritosaurus binsfeldi</i>, <i>Gualosuchus reigi</i>, both species of <i>Proterochampsa</i>, and Rhadinosuchinae in the strict consensus tree (Fig. 6A). Given the absence of overlapping elements between the new taxon and several other proterochampsids (e.g. <i>Proterochampsa nodosa</i>, <i>Cerritosaurus binsfeldi</i>), these unresolved affinities are unsurprising. The new taxon, however, does not nest within Rhadinosuchinae in any of the MPTs, thus <i>Stenoscelida aurantiacus</i> is a non-rhadinosuchine proterochampsid. The clade Rhadinosuchinae is composed of a trichotomy including <i>Rhadinosuchus gracilis</i>, <i>Pse</i>
description: Published as part of <i>Müller, Rodrigo Temp, Garcia, Mauricio Silva &amp; Fonseca, André de Oliveira, 2022, A new proterochampsid (Archosauriformes: Proterochampsia) from the Late Triassic of southern Brazil and the emergence of archosaurian hind limb traits, pp. 1-19 in Journal of Systematic Palaeontology 20 (1)</i> on pages 3-15, DOI: 10.1080/14772019.2022.2128913, http://zenodo.org/record/7331376
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