Draconyx loureiroi Mateus & Antunes 2001

Autor: Rotatori, Filippo Maria, Moreno-Azanza, Miguel, Mateus, Octávio
Rok vydání: 2022
Předmět:
DOI: 10.5281/zenodo.6536742
Popis: DRACONYX LOUREIROI MATEUS & ANTUNES, 2001 (FIGS 2–7) Type material: The holotype specimen (newly reported material marked with *), ML 357 (subnumbers from 1 to 31) includes two maxillary teeth, carpal bones, two metacarpal distal ends*, three left carpal phalanges, one right carpal phalanx*, two left unguals, three right unguals* and one right leg including proximal femoral epiphysis, proximal and distal epiphysis of the tibia, astragalus, calcaneum, four metatarsals, five phalanges and two unguals. Referred specimen: An isolated left femur, ML 434 from 1 km south of the type locality previously referred to Draconyx loureiroi by Mateus & Antunes (2001), now referred to Ankylopollexia indet. Type locality, horizon and age: Vale de Frades, Lourinhã Municipality, Portugal. Praia Azul Member of Lourinhã Formation, lower Tithonian, 151–152 Mya. Emended diagnosis: Styracosternan iguanodontian distinguished from other basal iguanodontians by the following combination of characters: unfused and noninterlocked carpus; absence of a sharp crest running from the medial condyle of the femur towards of the lesser trochanter, fully open U-shaped extensor groove on distal epiphysis of the femur; fully open V-shaped flexor groove without overhangs on distal epiphysis of the femur; concave medial margin of proximal epiphysis of the tibia; caudally pointing fibular condyle of the tibia; and a splint-like metatarsal I. Description and comparisons Cranial material: Maxillary teeth (ML 357–31) (Fig. 2A, B): One of the maxillary teeth preserves part of the root and its crown is complete (Fig. 2A). The other tooth is just an isolated crown (Fig. 2B). The specimens appear to have suffered some erosion and post-mortem breakage. The root is slightly labiolingually curved and tapers smoothly into the crown (Fig. 2A). There is no cingulum at the junction between the root and the crown. Overall, the crown is leaf-shaped and the veneer of enamel is thicker on the labial side (Fig. 2A). Labially a thick primary ridge is distally offset and five accessory ridges are present on the mesial surface (Fig. 2A, B). Non-mammillated hook-like denticles are coarsely present on the mesial crown margin, while distally they appear to have been obliterated by either erosion or occlusal wear. An extensive occlusal surface develops on the apex of the crown and is inclined labiolingually approximately around 30° (Fig. 2A, B). Remarks: The tapering root, slightly labiolingually recurved and the leaf-shaped crown are common characteristics of Dryomorpha (Galton, 1983, 2006; Norman, 1986, 2004). The maxillary crowns possess a distally offset primary ridge resembling the condition of ankylopollexians but differing from dryosaurids in which the primary ridge is located towards the centre of the crown (Galton, 2006). In the original description, Mateus & Antunes (2001) indicated the presence of five accessory ridges in the distal half of the crown as an autapomorphy of Draconyx loureiroi. However, the number of accessory ridges is variable throughout the dental series (Galton, 2006). Therefore, we regard it as a non-informative character if the tooth position is not articulated. Axial skeleton: Caudal vertebrae (ML 357–9–11) (Figs 3K–M, 4A–F): Three proximal caudal vertebrae are preserved and, as for the rest of the skeleton, they have suffered breakage and erosion. The centra are stout and craniocaudally compressed. The largest centrum is ML 357–9, while ML 357–10 and ML 357–11 are slightly smaller. The cranial and caudal surfaces are slightly amphicoelous, having a subelliptical, rounded shape (Fig. 4 A, C, G, I, M, O). Dorsally, the neural arch preserves the ventralmost portion of the neural spine but the neural suture is not visible. Immediately caudal to the spine, in the most complete specimens ML 357-9 and ML 357 - 10, (Fig. 4C, O) two small, rounded and steeply inclined post-zygapophyses diverge laterally. On the lateral surface of the neural arch, the articulation for the transverse processes is overlain by a broken surface. The presence of this surface indicates that these vertebrae represent the cranialmost portion of the caudal series. In ventral view, the centra have an hourglass-shaped outline (Fig. 4E, K, Q). The ventral surfaces of the centra possess a narrow keel and the margin is highly concave (Fig. 4B, P). Caudally, broad facets for chevrons are present. Remarks: The three caudal vertebrae are stout and cylindrical, similar in overall shape and proportions to those of other basal iguanodontians, such as: Camptosaurus dispar, Cumnoria prestwichii, Dryosaurus altus, Dysalotosaurus lettowvorbecki, Mantellisaurus atherfieldensis Mantell, 1825 and Uteodon aphanoecetes (Galton & Powell, 1980; Galton, 1981; Norman, 1986; Carpenter & Wilson, 2008; Carpenter & Galton, 2018). They do differ remarkably from those of Iguanodon bernissartensis Van Beneden, 1881 and Iguanacolossus fortis McDonald et al., 2010b in being less discoidal in shape, and from Barilium dawsoni Lydekker, 1888a by being less compressed dorsoventrally (Norman, 1980, 2011; McDonald et al., 2010b). Manus (ML 357–1–5, 20–26) (Fig. 5): The partial carpus (ML 357–5) is composed of the proximal part of a metacarpal, two distal carpals and a proximal carpal (Fig. 5A). Two isolated distal epiphyses of metacarpals (ML 357–25, 26) are associated with the semiarticulated carpus, being both consistent in size and their state of preservation (Fig. 5F, G). Four manual phalanges and five ungual phalanges are associated with the carpus (Fig. 5B–E). Three manual phalanges (phalanx n.2 ML 357–22, phalanx n.3 ML 357–21 and phalanx n.4 ML 357–24) and two unguals (ungual n.1 ML 357–1 and ungual n.4 ML 357–23) are likely to belong to the left manus (Fig. 4), while the remaining three unguals (ungual n.2 ML 357–2, ungual n.3 ML 357–3 and ungual n.5 ML 357–20) and manual phalanx n.1 ML 357–4 are attributed to the right manus. Given the fragmentary condition and the weathering of the specimen, it is not possible to identify the bones of each individual carpal. The two distal carpals are cuboid in shape and are stout elements, while the proximal carpal is more lightly built and slightly arched (Fig. 5A). The proximal end of the metacarpal preserves a concave proximal margin. Despite the poor preservation, it is possible to determine that the carpus does not include fused elements, as all of the contacts between all the identifiable carpals are clearly visible (Fig. 5A). The two isolated distal metacarpals fragments are similar in shape: they are compressed along their extensor–palmar axis, broadening mediolaterally towards their distal ends. The medial and lateral ginglymi are subequal in size and preserved on their collateral ligament pits. The manual phalanges, ML 357–22 and ML 357–21, articulate with each other. Furthermore, phalanx ML 357–22 articulates with the ungual ML 357–23. Therefore, these latter manual elements compose a complete finger (Fig. 5C, D, K). In general, all of the phalanges have a distal triangular section, moderately arched ventral surface and strongly inclined and robust ginglymi. The lateral ginglymus is smaller with respect to the medial one, with the exception of phalanx n.2 where they are subequal (Fig. 5B–E). The unguals are generally elongated and clawlike, slightly arched along the extensor–palmar axis and have a subtriangular articular facet (Fig. 5H–L). The only exception, ungual n.5 (Fig. 5L), appears instead to be compressed along its extensor–palmar axis. A fracture develops mediolaterally and is slightly inclined craniocaudally. Both parts are separated by this fracture and have suffered mediolateral displacement. 3025 357 – 19– 19 R e m a r k s: D e s p i t e t h e p o o r p r e s e r v a t i o n o f the carpus of ML 3 5 7, certain features allow comparisons with other taxa. The small metacarpals are not significantly different from those of other iguanodontians (Norman, 2004). However, the carpus is constituted by isolated blocky elements, resembling the condition in dryosaurids and Tenontosaurus spp. (Dodson, 1980; Galton, 1981; Forster, 1990; Winkler et al., 1997). The distal carpals of ML 357 differ from the ones of Camptosaurus dispar, which are arranged in two co-ossified and highly interlocked blocks. Furthermore, the carpus of ML 357 does not exhibit the total ossified condition present in Iguanodon bernissartensis, Magnamanus soriaensis Vidarte et al., 2016, Mantellisaurus atherfieldensis and other styracosternans (Fig. 12A–J) (Dodson, 1980; Norman, 1980, 1986, 2004, 2011, 2015, Vidarte et al., 2016). The carpals and the manual unguals are similar to the ones present in Camptosaurus dispar, Cumnoria prestwichii and Uteodon aphanoecetes, differing from the ones of more derived styracosternans (i.e. Iguanodon bernissartensis, Magnamanus soriaensis, Mantellisaurus atherfieldensis and all other hadrosauriformes) in being, respectively, more arched and more claw-like instead of extremely compressed and hoof-like (Fig. 12A–J) (Galton & Powell, 1980; Norman, 1980, 1986, 2004; Carpenter & Wilson, 2008, Vidarte et al., 2016). Femur ML 357–6 (Figs 3A–E, 6A–E): The preserved right femur consists of the heavily eroded distalmost part of the shaft and the distal epiphysis. The proximalmost part of the preserved shaft is mediolaterally crushed and compressed, due to taphonomic processes. The femur is strongly bowed craniocaudally; the section of the shaft was subcircular but is heavily distorted proximally by the compression and damaged distally (Fig. 6B, D). The distal epiphysis is subrectangular in shape in distal view, extending slightly more mediolaterally than craniocaudally. The two distal condyles are preserved, appearing subequal in size with the lateral condyle slightly larger than the medial one (Fig. 6E). A deep, extensive, fully open and U-shaped extensor groove separates the two condyles cranially (Fig. 6E). The cranial process of the lateral condyle is rounded and deflected caudally. The caudal fingerlike process of the lateral condyle (condylid, according to Bertozzo et al., 2017) is not preserved, being broken at its base, but the crest for the muscular insertion is distinguishable. The medial condyle appears stout and rectangular, although its cranial and caudal processs have been eroded. The flexor groove is fully open and its margin, consisting of the caudal process of the lateral condyle and the condylid of the medial condyle, are V-shaped in outline. Remarks: The preserved femoral shaft is strongly bowed craniocaudally, as in dryosaurids, most elasmarians, Camptosaurus dispar and differently from Tenontosaurus sp. and other styracosternans (Norman, 1980, 1986, 2004; Carpenter & Wilson, 2008; Carpenter & Galton, 2018; Herne et al., 2019; Rozadilla et al., 2019, 2020). On the cranial surface there is no crest developing from the medial condyle, extending proximally towards the lesser trochanter, as seen in other ankylopollexians, such as Camptosaurus dispar, Iguanodon sp., Mantellisaurus atherfieldensis and Uteodon aphanoecetes (Gilmore, 1909; Norman, 1980, 1986, 2004; Carpenter & Wilson, 2008; Carpenter & Galton, 2018). The lateral condyle of the distal epiphysis of the femur in Draconyx loureiroi is concave in outline and it extends more craniocaudally than the ones of Camptosaurus dispar, Cumnoria prestwichii and Uteodon aphanoecetes (Fig. 7; Galton & Powell, 1980; Carpenter & Wilson, 2008). The inflection point of the curvature is located more cranially in Draconyx loureiroi than in Camptosaurus dispar and Uteodon aphanoecetes (Carpenter & Wilson, 2008; Carpenter & Galton, 2018), while Cumnoria prestwichii exhibits a smoother outline without abrupt changes in curvature. The lateral condyle of Early Cretaceous species, such as Barilium dawsoni and Mantellisaurus atherfieldensis, are larger in proportions with respect to the total size of the epiphysis and extend more caudally than the ones of the above-mentioned taxa, including D. loureiroi (Norman, 1986; 2011). The medial condyle of Draconyx loureiroi is subrectangular and its medial margin is straight, as is the one of Mantellisaurus atherfieldensis and Barilium dawsoni (Norman, 1986; 2011). However, in Jurassic taxa such as Uteodon aphanoecetes and Camptosaurus dispar, the medial margin of the medial condyle is rounded (Carpenter & Wilson, 2008; Carpenter & Galton, 2018). The flexor and extensor grooves are fully open as in many basal and cursorial iguanodontians, contrasting with more derived forms (Norman, 2004). Extensor grooves of the Jurassic taxa Camptosaurus dispar and Uteodon aphanoecetes are fully open, but shallower compared to the ones of Barilium dawsoni, Cumnoria prestwichii, Draconyx loureiroi, Iguanodon sp. and Mantellisaurus atherfieldensis (Norman, 1980, 1986, 2004, 2011, Carpenter & Wilson, 2008; Carpenter & Galton, 2018; Verdú et al., 2018). Moreover, the flexor groove walls of Cumnoria prestwichii and Uteodon aphanoecetes are slightly divergent from one another (Fig. 7). The extensor groove of the Early Cretaceous iguanodontians Barilium dawsoni, Mantellisaurus atherfieldensis and other styracosternans are partially enclosed by overhangs of medial and lateral condyles. Draconyx loureiroi does not exhibit an overhang of the medial condyle on the flexor groove, unlike the condition present in Camptosaurus dispar, Cumnoria prestwichii, Mantellisaurus atherfieldensis, Ouranosaurus nigeriensis Taquet, 1976 and Uteodon aphanoecetes (Norman, 1980, 1986, 2004, 2011; Carpenter & Wilson, 2008; Bertozzo et al., 2017; Carpenter & Galton, 2018, Verdú et al., 2018). This condition more closely resembles the plesiomorphic condition within Ornithopoda (Norman et al., 2004). Tibia (ML 357–7, 12) (Figs 3F, G, 6F–J, 8A–D): The proximal (Fig. 6F–J) and distal (Fig. 8A–E) epiphyses of the tibia are preserved, but the tibial shaft is missing. Both extremities are heavily eroded and covered by matrix and adhesives, but relevant characters are still distinguishable. The proximal epiphysis preserves a conspicuous cnemial crest, a robust fibular condyle and the internal condyle. The cnemial crest tapers dorsally forming a smooth edge; in dorsal view its margins are laterally concave and medially convex (Fig. 6G, I). Laterally, it is divided by the fibular condyle to form a deep and extensive scar (sulcus tibialis). The fibular condyle is a stout process that deflects strongly caudally, its articular facet is rounded and it blends smoothly with the anterior and posterior surfaces of the condyle. The internal condyle is a blunt eminence directed caudally and projecting gently laterally. Like the cnemial crest, its lateral margin is concave, while the medial margin is strongly convex (Fig. 6E). This convexity forms a deep sulcus that separates the internal condyle from the fibular condyle. Immediately ventral to the proximal epiphysis, the proximal part of the diaphyseal shaft is preserved and it is teardrop-shaped in cross-section. The distal part of the tibia is articulated with the astragalus, calcaneum and the metatarsals (Fig. 8A–E). The distal epiphysis flares mediolaterally, the medial malleolus is rounded and slightly more expanded mediolaterally than the lateral one, which is narrow and elongate, expanding proximodistally. They articulate between each other forming an angle of approximately 45°. The intermuscular line is located on the caudal surface of the distal epiphysis, separating the surface of the two malleoli and reaching the astragalus, forming with its apex a continuous concave surface. Remarks: The medial margin of the proximal epiphysis of the tibia is convex, as is typical of many iguandontians (Norman, 2004), except for Talenkauen santacrucensis Novas et al., 2004 and Eousdryosaurus nanohallucis Escaso et al., 2014 (Escaso et al., 2014; Rozadilla et al., 2019; Dieudonné et al., 2021). As noted by Dieudonné et al. (2021), the cnemial crest apex is directed strongly craniolaterally, a characteristic common to Laurasian dryomorphans and Dysalotosaurus Pompeckj, 1920, differing from the condition exhibited by Elasmaria and the Portuguese ornithopod Eousdryosaurus nanohallucis (Escaso et al., 2014). As in many styracosternans, the cnemial crest is well developed (Norman, 2004). The fibular condyle differs from dryosaurids (with the exception of Valdosaurus canaliculatus), Camptosaurus dispar and Uteodon aphanoecetes in being partially posterolaterally deflected, similar to the condition seen in Cumnoria prestwichii, Talenkauen santacrucensis and more derived styracosternanans (Galton & Powell, 1980; Norman, 1980, 1986, 2011; Norman et al., 2004; Carpenter & Wilson, 2008; Barrett et al., 2011; Rozadilla et al., 2019). The distal epiphysis, found in articulation with the rest of the pes, does not differ significantly from that of other iguanodontians (Norman, 2004). Fibula (ML 357–8, 12) (Figs 3I, J, 6F–J, 8A–D): The distal and proximal epiphyses of the fibula are preserved, whereas the diaphysis is completely absent. The proximal end is a flattened subtriangular element. Dorsally, it appears to be slightly medially convex. The cranial margin deflects abruptly dorsally into the cranial process, which is slightly eroded (Fig. 6I). In contrast, the caudal margin deflects less abruptly dorsally. A deep fossa is present close to the caudal margin on the lateral surface. The distal epiphysis is in articulation with the rest of the pes, located on the lateral surface of the tibia and contacts the calcaneum (Fig. 7A–D). Remarks: The proximal epiphysis has cranial and caudal margins that diverge smoothly, as in Dryosaurus altus, Dysalotosaurus lettowvorbecki, Eousdryosaurus nanohallucis, Talenkauen santacrucensis and Valdosaurus canaliculatus, but differs from Iguanodon bernissartensis, Mantellisaurus atherfieldensis and Ouranosaurus nigeriensis (Norman, 1980, 1986; Galton, 1981; Barrett et al., 2011; Escaso et al., 2014; Barrett, 2016; Bertozzo et al., 2017; Rozadilla et al., 2019). As in Sektensaurus sanjuanboscoi Ibiricu et al., 2019 and Talenkauen santacrucensis, the caudal margin of the proximal part of the fibula is almost vertical (Ibiricu et al., 2019; Rozadilla et al., 2019). Pes (ML 357–12–19) (Figs 3N, Q, 8A–F, 9C, D): The pes is articulated with the distal tibia, distal fibula
Published as part of Rotatori, Filippo Maria, Moreno-Azanza, Miguel & Mateus, Octávio, 2022, Reappraisal and new material of the holotype of Draconyx loureiroi (Ornithischia: Iguanodontia) provide insights on the tempo and modo of evolution of thumb-spiked dinosaurs, pp. 125-156 in Zoological Journal of the Linnean Society 195 (1) on pages 129-143, DOI: 10.1093/zoolinnean/zlab113, http://zenodo.org/record/6530795
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