taxonID	type	description	language	source
03FA87B7B35B0F184B07FDA5FC1BE05B.taxon	materials_examined	Holotype: BSPG 1937 I 1 a & b (Fig. 1), complete skeleton preserved in dorsal view and counterpart. Locality and horizon: Solnhofen Limestone, Tithonian, Late Jurassic; Wintershof wei Eichst € att, Germany. Revised diagnosis: Eichstaettisaurus schroederi can be distinguished from Eichstaettisaurus gouldi in having the parietals paired; frontal subolfactory processes in contact medially; a frontoparietal suture smooth and slightly convex anteriorly; frontal and parietals of equal width at their suture and in contact laterally; straight and laterally projecting caudal transverse processes; autotomy septa at the level of the fifth caudal. Remarks: Eichstaettisaurus schroederi differs from both Ardeosaurus species in having strongly laterally emarginated paired parietals; frontals totally fused in dorsal aspect; parietals with shorter supratemporal processes; frontoparietal suture slightly convex anteriorly; postorbital and postfrontal as separate elements; seven cervical vertebrae; and more than 30 presacral vertebrae. Eichstaettisaurus schroederi further differs from A. digitatellus in having a wide posterior parietal margin between supratemporal processes. Eichstaettisaurus schroederi further differs from A. brevipes in the absence of skull roof ornamentation and cephalic osteoderms, as well as a pedal phalangeal formula of 2 - 3 - 4 - 5 - 4. The difference in the degree of frontal interorbital constriction between E. schroederi and E. gouldi used by Evans, Raia & Barbera (2004) to differentiate these two species does not correspond to our measurements. The former authors’ ratio for the frontal interorbital / frontoparietal widths for E. gouldi (0.2) seems to be accurate based on our analysis of the published picture of the holotype. However, our personal measurement of E. schroederi indicates that the interorbital / frontoparietal ratio is 0.25, not 0.32 as reported by Evans et al. (2004) – see Table 1 below, as well as the Material and Methods section above. See also Appendix 2 for phylogenetic autapomorphies for E. schroederi from the analysis performed herein. EICHSTAETTISAURUS GOULDI EVANS, RAJA & BARBERA, 2004 Holotype: MPN 19457, articulated partial skeleton preserved in ventral view. Locality and horizon: La Cavere outcrop, Pietraroia, Mount Matese, southern Italy. The horizon is Albian (Upper Plattenkalk), Early Cretaceous. Revised diagnosis: Eichstaettisaurus gouldi can be distinguished from E. schroederi in having the parietals fused; subolfactory processes of frontals well developed, but not in contact medially; frontoparietal suture with the parietal slightly concave medially, receiving a posterior convexity of the frontal; the frontal being slightly wider than the parietals at their suture; at least one posteriorly orientated anterior caudal transverse process; autotomy septa, if present, begin beyond the level of the sixth caudal. Remarks: Eichstaettisaurus gouldi shares with E. schroederi a depressed skull; short and blunt snout; absence of lacrimals; paired premaxillae; postorbital and postfrontal as separate elements; frontals fused; frontals widen anteriorly; frontal interorbital / frontoparietal width ratio between 0.2 and 0.3.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B35A0F104B14F98FFDF2E521.taxon	description	The holotype of E. schroederi (BSPG 1937 I 1 a & b) consists of two slabs representing part and counterpart. One slab includes all the preserved bone elements in dorsal view, and has been partially prepared on the opposite side, revealing portions of the ventral side of the skull. The counterpart preserves only the impression of those elements in the soft calcareous matrix, but provides some good morphological detail. The condition seen in E. schroederi is marked as ‘ X’, and the opposite condition is marked as ‘ Y’. For feature number 8, a third condition is added, ‘ Z’, which is exclusive of E. gouldi. Missing data denoted as ‘? ’. Shared features across Eichstaettisaurus and Ardeosaurus = 3; between E. schroederi and A. brevipes only = 1; between E. schroederi and A. digitatellus only = 4; and between A. brevipes and A. digitatellus only = 7. Cranium The skull is depressed (Fig. 2 A, B) and the bones are unsculptured. The snout is short, broad, and rounded anteriorly. The premaxillae are paired and possess a short nasal process that contacts the anterior end of the nasals forming the anteromedial border of the external nares. The maxillary process in each premaxilla is short, and has a smoothly rounded posterior end that contacts the maxilla (Fig. 3). The anterior end of the premaxillary process of each maxilla has a similar shape, and it is likely that a soft tissue connection existed between the two elements. No anterior ethmoidal foramina are visible on the premaxillae, indicating that the medial ethmoidal nerves probably exited through the large external nares. The nasals are paired and their anterior halves form the medial – posteromedial borders of the external nares. No nasal foramina are observable dorsally, and there are no ventrolateral projections. On the ventral side, the straight suture between the nasals is visible and there is no midline crest. The septomaxillae can be seen within the nasal capsule on both sides. They seem to be flattened anteriorly, although this could be because of the deformation of the specimen. Each maxilla bears a short premaxillary process, which forms a steep angle with the maxillary nasal process. The nasal process on each side contacts the nasal dorsomedially and the prefrontal posteriorly. Small alveolar foramina for the cutaneous branches of the superior alveolar nerve and artery are visible close to the margin of the tooth row, but the anterior superior alveolar foramen is not visible externally. It is probably present close to the base of the medial margin of the nasal process and thus not visible in lateral aspect. The suborbital process of each maxilla extends to the midpoint of the orbit with its posterior tip lying ventral to the anterior end of the jugal. A lacrimal bone is absent, and the anterior border of the orbit is formed mostly by the prefrontals, which lack any surface ornamentation and possess a faint ridge projecting posterodorsally. This ridge does not form a prefrontal boss, as seen amongst many iguanians. The lacrimal foramen is not visible laterally, and it was probably present posteriorly, with the lacrimal duct opening into the orbital cavity, as observed in most lizards. Both jugals are preserved. Whereas the left element is apparently complete, the right one is broken, preserving only its postorbital process. Each jugal is a semilunate element extending far anteriorly, approaching the prefrontal. Each one lacks a posteroventral process, and ascends posteriorly in a smooth curve contacting the postorbital on its posterior margin. The posterior end of the postorbital process of the jugal approaches the anterior end of the squamosal, but does not contact it. The postorbitals and postfrontals are evident on both sides of the skull, but the right elements are better preserved, and in articulation. The postfrontals are forked medially, lying lateral to the frontoparietal articulation, and each possesses a single distal process that clasps the postorbital posteriorly. Each postorbital is a triradiate element that is separate from the postfrontal and forms most of the posterior margin of the orbit. This differs from previous interpretations, in which the postorbitals are considered to be fused to the postfrontals (Cocude-Michel, 1963; Hoffstetter, 1966; Estes, 1983). The ventral process of the postorbital contacts the orbital margin of the jugal and its dorsal process contacts the postfrontal also on the orbital margin. The latter feature is uncommon amongst lizards because the dorsal process of the postorbital usually contacts the posterior margin of the distal process of the postfrontal. The posterior process of the postorbital is relatively elongate, extending more than half the length of the upper temporal fenestra, and lying dorsal to the squamosal. The squamosals are similar to those of most other lizards in being relatively slender and each having a posteroventral process contacting the cephalic condyle of the quadrate. Both squamosals lack a dorsal process, a feature that is usually present in iguanians and teiids. A relatively short supratemporal bone is present between the posterior end of the squamosal and the parietal, being short in length and abutting against the lateral margin of the supratemporal process of the parietal. The frontals are fused to each other, and form most of the dorsomedial margin of the orbits. They are widely expanded posteriorly, but are constricted at their midpoint and are slightly expanded anteriorly. The anterolateral processes of the frontal are elongate, but do not reach the maxillae, allowing for a short contact between the nasal and prefrontal. A much shorter anteromedial process intrudes between the nasals. In ventral view (Fig. 2 C), the frontals bear subolfactory processes (best seen posteriorly), which seem to never touch medially at any point, but matrix partially obscures their central portion. There is seemingly no midventral crest of the frontals. Posteriorly, the frontals contact the parietals along a weakly anteriorly convex suture. There is no evidence of parietal tabs of the frontal either dorsally or ventrally. The parietals are paired elements, which is uncommon to a number of squamates, but seen in some extant geckoes [e. g. Nephrurus (Evans, 2003), Sphaerodactylus (Daza et al., 2008), and Pygopus (AMNH R 140843)] and Xantusia (AMNH R 150172 and AMNH R 150174). The pineal foramen is present close to the centre of the parietal table. The lateral margins of the parietals are emarginated and form most of the medial border of the upper temporal fenestrae. The supratemporal processes are relatively short and bear a medial excavation. A posteromedial process is absent, as also seems to be the case for the nuchal fossa. No frontal tabs are present either dorsally or ventrally. The palatines do not contact in the midline, being widely separated from each other, and are relatively short anteroposteriorly (Fig. 2 A, B). Their anterior margins are almost entirely visible in dorsal aspect, contacting the posteroventral margin of the prefrontals, and thus the palatines make little or no contact with the frontals anterodorsomedially. Most of their posterior ends form a wide and nearly straight contact with the anteromedial processes of the pterygoids. The pterygoids are widely separated from each other. Each pterygoid has a relatively wide, undivided anteromedial process and, based on the right side of the skull (which is better preserved than the left side), seems to be inclined medially. The transverse process of each pterygoid is short and directed anterolaterally, contacting the ectopterygoid on the left side (but see below). The quadrate process of the pterygoid is slender and extends far posteriorly, although disarticulated from the quadrates as preserved in BSPG 1937 I a, b. As the slab with the bone elements in the holotype is preserved in dorsal aspect, and the palate is not clearly visible in ventral view, it cannot be determined whether a ventral flange was present on the transverse process of each pterygoid, or whether or not teeth are absent or present on the palate. The left ectopterygoid is visible and a small broken element lateral to the transverse process of the right pterygoid might represent its right counterpart, although it may also represent part of the anterior extension of the right jugal. The left ectopterygoid is semilunate and has no lateral process. Its posterior end lies dorsal to the pterygoids, as is the case in some geckoes (e. g. Coleonyx – AMNH R 89271) and anguids (e. g. Diploglossus – AMNH R 154690). The supraoccipital is visible in dorsal view, bearing an ossified processus ascendens tecti synocti that contacts the parietal posteriorly. It cannot be determined whether anterolateral processes of the supraoccipital are present. A pair of posterolaterally orientated crests are present on the supraoccipital. These are similar to the ‘ crests’ formed on the supraoccipital by the posterior semicircular canal in some geckoes (e. g. Hoplodactylus pacificus – MCZ R- 141790 and Coleonyx variegatus – AMNH R 89271) and some other lizards of reduced size (e. g. Feylinia corrori – MCZ R- 42886 and MCZ R- 106990). These apparent ‘ crests’ are thus considered herein as a consequence of a reduced degree of ossification due to diminutive size. On the ventral side of the slab containing skeletal elements (Fig. 2 C), parts of the basioccipital and basisphenoid are visible. These elements are not fused to each other, and the basisphenoid is flat on its ventral surface. The basipterygoid processes are stout and have slightly expanded distal ends, but are no longer in articulation with the pterygoids. The quadrates have been tilted posteriorly and their anterior surfaces face dorsally (Fig. 1 A, B). They each have a shallow anterior concavity, and the two articular condyles are of similar size. Mandibles The dentaries are visible on both sides of the slab containing the skeletal elements, but they are more readily observed in medial aspect on the ventral side of the specimen (Fig. 2 D). The Meckel’s canal is closed medially by the fusion of the subdental ridge to the ventral crest of the dentary in its anterior half, as is also observed in extant geckoes and xantusiids (Evans, 2008). The splenial is not visible, and it cannot be determined whether it is overlain by the matrix, not preserved, or perhaps fused with the dentary as in xantusiids (Estes, de Queiroz & Gauthier, 1988). In dorsal aspect, most of each mandible is obscured beneath the skull. However, part of the posterior end of each dentary can be observed, and each part opens medially (contrasting with the anterior and middle portions of the dentaries), where the splenial usually articulates. The coronoids are partially visible on both sides. The left one bears a relatively elongate posteroventral process, but no posterior process. The right counterpart is mostly overlain by the pterygoid transverse process, and a broken element that might represent either the jugal or the ectopterygoid. The coronoid bears a very short anteromedial process that is directed ventrally. Both surangulars are visible dorsally and each has a thickened dorsal border, forming the labial margin of the adductor fossa. The latter is relatively deep, but narrow, and faces dorsomedially. A short, posteriorly directed retroarticular process is present posterior to the glenoid, and lacks any lateral notching. Dentition The dentary teeth are not visible in the holotype, but the maxillary teeth are visible on both sides (Fig. 3), along with some of the premaxillary teeth. The teeth are small, narrow, conical, and appear to be unicuspid, with no differentiation between the premaxillary and maxillary regions. The tooth count cannot be established with precision for the premaxillae, but there are c. 22 teeth preserved in situ on the right maxilla, and the total tooth count in each jaw half is estimated to be at least 30. Variation in tooth height throughout the dental arcade indicates that tooth replacement was present and was still active at the time of death. The teeth are positioned medially on the jaw labial margin (or parapet), and are unfused to the jaw margins, suggesting pleurodonty. Postcranium The preserved vertebral column includes the axis, the remaining cervical vertebrae, the dorsals, sacrals, and anterior caudals. The number of cervicals is estimated to be seven, because the eighth presacral vertebra seems to be the first one bearing ribs long enough to reach the sternum (Fig. 4 A). Only the neural arches and neural spine of the axis are visible, with its intercentrum, and the atlas, being hidden from view. The postaxial cervicals bear very low neural spines, and most bear no ribs. The first ribs appear at the level of the sixth or seventh cervical. Based on the identification of the eighth presacral as the first dorsal, the total number of dorsals is estimated to be 24, giving a total of 31 presacrals. This is a higher presacral vertebral count in comparison to that of most iguanians (with 24 presacrals), and a number of scincomorphans (e. g. cordylids, gerrhosaurids, lacertids, and teiioids) and gekkotans, many of which bear 26 presacrals. Yet, a slight increase in the presacral count as occurs in E. schroederi (around 30 presacrals) is observed in a variety of taxa, including xantusiids and some gekkotans, as well as scincids, xenosaurids, anguids, and varanids (Hoffstetter & Gasc, 1969). The neural spines in the dorsal region are also small, and the last presacrals seem to lack any ribs (thus characterizing lumbar vertebrae). One striking feature of the vertebral morphology is the presence of accessory intervertebral articulations (zygosphenes and zygantra; Fig. 4 B). The zygosphenes face dorsolaterally as in many iguanians, cordylids, gerrhosaurids, lacertids, and gekkotans (although they are absent in pygopodids), and are notched anteriorly, as it is also the case in all aforementioned groups, as well as in teiids. Estes (1983), following Broili (1938), reported a procoelic condition for the intervertebral articulations. However, as previously noted by Hoffstetter (1964) and Evans et al. (2004), it is not possible to determine the structure of centra with any confidence in this specimen as it is preserved in dorsal view. Two sacral vertebrae can be identified just medial to the anterior half of the ilia and anterior to the first caudal that bears short transverse processes (Fig. 4 C). Five anterior caudals are preserved, the remaining part of the tail having been autotomized (Figs 1, 4 C). The soft tissue impression of the cartilaginous rod that developed subsequently to the loss of the tail can be seen on both slabs, and was previously illustrated under UV-light by Tischlinger & Wild (2009). The ribs are holocephalous with circular articular surfaces. There are 21 or 22 pairs of presacral ribs. The posterior-most presacral vertebrae do not bear any ribs, therefore constituting a ‘ lumbar’ region. There are no accessory tuberculi anteriorly or posteriorly to the rib heads, and the ribs lack any degree of pachyostosis. Gastralia are absent, as is the case for all other squamates, but inscriptional ribs are visible attached to the distal ends of most of the dorsal ribs, but absent from the posterior-most four. Of the pectoral girdle, the clavicles, left scapula, and part of both coracoids are preserved (Fig. 4 A and D). The clavicles lack the dorsally located cranial curvature observed in most autarchoglossans (Estes et al., 1988), as well as the posterior process that is seen in some scincids and cordyloids (e. g. Mabuya mabouia – AMNH 141128 and Gerrhosaurus major – AMNH 173621). It is not possible to determine whether the clavicles have a proximoventral fenestra, but they are expanded in this region. The left scapulocoracoid exhibits no suture between the scapula and coracoid. The scapula is short, has an expanded acromion process dorsally, and lacks a scapular ray defining a separate scapular emargination dorsal to the scapulocoracoid fenestra (Fig. 4 D). The supracoracoid foramen is visible medially on the scapulocoracoid (Fig. 4 D), located at the level of, and just posterior to, the anterior coracoid ray and anterior (or primary) coracoid emargination of most lizards. The anterior (or primary) coracoid ray, delimiting the anterior coracoid emargination ventrally, is partially overlain by one of the left dorsal ribs. The left coracoid is mostly obscured beneath the vertebral column, and the right coracoid is preserved mostly as an impression in the calcareous matrix. The coracoids are relatively small when compared with most other lizards studied by us, being similar in dimension to the scapula. Both forelimbs are preserved in articulation (Figs 1, 4 A). Details of the proximal surface of the humerus are better seen on the left element (Fig. 4 A), in which the bicipital fossa faces dorsally. The distal end of the humerus is twisted posteriorly relative to the longitudinal axis of the humeral shaft, and the state of preservation hampers identification of either an ectepicondylar or an entepicondylar foramen (Fig. 4 E, F). The ulna has a well-developed olecranon process as well as an expanded distal epiphysis. The radius is present, but its distal epiphysis has suffered some degree of weathering, so the styloid process is not preserved (Fig. 4 G, H). The right manual carpus (Fig. 4 G) preserves the fourth distal carpal proximal to the fourth metacarpal, but no other distal carpals are distinguishable owing to poor preservation of this region. The metacarpals increase in length from the first to the fourth metacarpal, with the fifth being approximately half the length of the fourth. The phalangeal formula is 2 - 3 - 4 - 5 - 3, and the penultimate phalanges are elongated in comparison to the intermediate ones between the first and the penultimate ones (Tables 2 and 4). For the pelvic girdle, the pubes, ischia, and the impressions of both ilia are preserved. The pubes have relatively narrow pubic aprons, which are directed anteromedially, and lack pubic tubercles. The obturator foramen is not visible, probably because of the crushing present on the acetabular margin of the pubes. The ischia possess enlarged plates, which seemingly contributed to a long symphysial contact between both elements. Each ilium contains a long posteriorly directed blade, but it cannot be determined whether a preacetabular process (Russell & Bauer, 2008) is present. M 1 – 5, manual digits 1 to 5; I – IV, phalanges I to IV on each corresponding digit. Missing data denoted as ‘? ’. The phalangeal measurements for E. schroederi represent the mean values for the right and left manus, whereas on A. digitatellus measurements were taken from the left manus only, as the limits between phalanges are mostly nondiscernible on the right counterpart. P 1 – 5, pedal digits 1 to 5; I – IV, phalanges I to IV on each corresponding digit. Missing data denoted as ‘? ’. The phalangeal measurements for E. schroederi represent the mean values for the right and left pedes (apart from digit II), whereas on A. digitatellus measurements were taken from the left pes only, as the limits between phalanges are mostly nondiscernible on the right counterpart. M 1 – 5, manual digits 1 to 5; P 1 – 5, pedal digits 1 to 5; I – IV, phalanges I to IV on each corresponding digit. Missing data denoted as ‘? ’. Both hindlimbs are preserved in articulation and in posterior aspect. The shaft of the femur is almost straight, diverging from the more common sigmoidal shape observed in lizards. The femoral heads are badly crushed along with the other elements in the acetabular region, and the distal ends of the femora have only their posterior margin exposed. The tibiae and fibulae lay in close apposition to each other, and their distal epiphyses are not totally ossified. The astragalus and calcaneum are not totally fused to each other and the tibial facet on the astragalus is almost flat, lacking a crest for articulation with the tibia (Fig. 8 A, B, D). The distal tarsals are crushed against the metatarsals, and their morphology cannot be described in detail. However, it seems that both distal tarsals three and four are present. The first metatarsal is slightly shorter than metatarsals II – IV, the latter being subequal in length. The fifth metatarsal has its proximal head hooked medially, contacting the fourth distal tarsal, and is significantly shorter than the other metatarsals. The phalangeal formula is 2 - 3 - 4 - 5 - 4, and the penultimate phalanges are elongated compared with the intermediate ones, being between 30 and 56 % longer than the antepenultimate phalanx (Fig. 8 A, B, D; Tables 3 and 4). Phalanges II (digit III) and I and III (digit IV) of the right pes, as well as phalanges I and II (digit III), and I (digit V) of the left pes have better preserved articulatory surfaces. Distally, these phalanges have a hemispherical (convex) shape, and at least some phalanges (e. g. phalanx II, digit III, right pes) articulate with a concave proximal surface of the phalanx distally to it. This condition is different from the typical ginglymoid bicondylar articulation of the intermediate phalanges of most lizards, and would have allowed a greater range of horizontal movement. Horizontal movement also occurs in the crescentic / cup-shaped articulation in the intermediate phalanges of Gekko gecko (Russell, 1975), but in the latter it is probably greater than in E. schroederi owing to its expanded distal phalangeal articulations. The unguals form claws that are slightly curved and dorsoventrally deepened (Russell, 1975), or tall, at their bases – claw height defined as the distance measured from dorsal to ventral at the base of the claw (Zani, 2000; Tulli et al., 2009; Crandell et al., 2014). Higher (or deeper) claws in E. schroederi, as well as in G. gecko (Fig. 9), bear a ventral expansion in lateral view, below the level of the contact with the penultimate phalanx, the functional consequences of which are further discussed below (see Functional morphology). In E. schroederi, average pedal claw height (measured from those claws visible in lateral aspect) equals 0.70 mm. The ratio between claw height / area of contact between the claw and the penultimate phalanx equals 1.707, indicating that the claw is about 70 % higher than the area of contact with the penultimate phalanx. LEPIDOSAURIA HAECKEL, 1866 SQUAMATA OPPEL, 1811 GEKKONOMORPHA FURBRINGER €, 1900 ARDEOSAURUS MEYER, 1860	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3520F104B24FE28FC6AE3B2.taxon	materials_examined	Holotype: Collection Hetzell, now lost (Mateer, 1982; Estes, 1983). Represented solely by a cast (NHMUK 38006). Referred material: PMU. R 58, described by Mateer (1982), but now also lost. Locality and horizon: Solnhofen Limestone, Tithonian, Late Jurassic; Wintershof wei Eichstaett, Germany. Revised diagnosis: Ardeosaurus brevipes can be distinguished from A. digitatellus by the presence of skull roof ornamentation and cephalic osteoderms; wider parietal posterior margin between supratemporal processes; parietal posteromedial process present; postorbital and postfrontal separate elements; 23 presacral vertebrae; foot phalangeal formula 2 - 3 - 4 - 5 - 3. Remarks: The above diagnosis is based on comparisons between the holotype of A. digitatellus and the description provided by Mateer (1982) of A. brevipes. See also Appendix 2 for phylogenetic autapomorphies of A. digitatellus from the analysis performed herein.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3520F104890FA5DFBE7E7FE.taxon	materials_examined	Holotype: CM 4026 (Fig. 6), partial skeleton including skull and parts of the postcranium preserved in dorsal view. Locality and horizon: Solnhofen Limestone, Tithonian, Late Jurassic; Wintershof wei Eichstaett, Germany. Revised diagnosis: Ardeosaurus digitatellus can be distinguished from A. brevipes by the absence of skull roof ornamentation and cephalic osteoderms; narrow parietal posterior margin between supratemporal processes; parietal posteromedial process absent; postorbital and postfrontal fused into a postorbitofrontal; 27 presacral vertebrae; pedal phalangeal formula? - 3 - 4 - 5 - 4.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3520F1048C4FC47FD32E329.taxon	materials_examined	Type species: Ardeosaurus brevipes. Revised diagnosis: Ardeosaurus can be distinguished from other genera of squamates by the following combination of characters: paired premaxillae; the parietals fused and with weak lateral emargination (creating a narrow upper temporal fenestra); parietals with long supratemporal processes; posteromedial process of parietal present; pineal foramen present; frontoparietal suture straight; frontals paired; frontals expand posteriorly but not anteriorly; frontals interorbital / frontoparietal width ratio c. 0.5; postfrontal (or postorbitofrontal) forked medially; jugal without posteroventral process.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3520F084B0AF9F4FC22E331.taxon	description	Cranium The holotype of A. digitatellus is represented by a slab containing a single individual in dorsal view (Fig. 5 A). Osteological material is preserved in the skull and postcranium, along with impressions of some of the missing elements in the surrounding matrix. The snout elements are not preserved in the holotype of A. digitatellus, but some impressions indicate their outline. The anterior-most elements preserved are the prefrontals, which are quite large compared with the prefrontals of most lizards known to us, also suggesting the absence of lacrimals (Fig. 5 B, C). They do not bear surface ornamentation or a prefrontal boss. They are connected posteromedially to the frontals. The frontals are fused anteriorly, but still preserve a sutural line in their posterior half. It is not possible to determine if they became completely fused later in ontogeny. The frontals are expanded at the frontoparietal contact, and become constricted between the orbits. As in A. brevipes, the frontals do not expand anteriorly (Mateer, 1982) and are less constricted between the orbits than they are in E. schroederi (Table 1). The frontoparietal suture is straight in A. digitatellus, and the parietals are completely fused (Fig. 5 A, B), bearing no remnants of the sutural line between them. The pineal foramen is present and located in the centre of the parietal table. The parietal is weakly emarginated on both lateral margins. This weak emargination, along with the orientation of the right squamosal (which is less displaced than its left counterpart), indicates that the upper temporal fenestra is narrower than the one observed in E. schroederi. The supratemporal processes are present, elongate, and also possess a medial excavation (Fig. 6 A). The posterior margin of the parietal between the supratemporal processes is reduced, forming a V-shaped posterior margin. A small protuberance observed posteromedially to the parietal table may represent a posteromedial process, or part of the processus ascendens of the supraoccipital, but we could not determine either because most of this element has eroded away. The supratemporal bone is short and is located between the posterior end of the supratemporal process of the parietal medially, and the squamosal laterally. The jugals are not entirely preserved, but their postorbital process is evident on both sides of the skull, the left element still contacting the ventral margin of the posterior process of the postorbitofrontal. The postorbital and postfrontal on the right side of the skull are fused into a postorbitofrontal (Fig. 6 B). Medially, the postorbitofrontal has one parietal and one comparatively longer frontal process, both processes contributing to an extensive contact with the frontal and parietal. The posterior process of the postorbitofrontal extends far posteriorly, beyond the midpoint of the upper temporal fenestra. The right squamosal is preserved lateral to the postorbitofrontal, indicating that the jugal extended far posteriorly, intervening between the postorbitofrontal and the squamosal. The posterior end of the left jugal is placed more anteriorly and did not reach beyond the posterior end of the postorbitofrontal. Whether the left jugal was displaced more anteriorly or the right one more posteriorly cannot be determined, but the impression of the anterior end of the left squamosal is located lateral to the posterior end of the left jugal, as observed on the right side. Therefore, it is concluded that the jugal prevented the postorbitofrontal from contacting the squamosal. The squamosal is a moderately stout element, bearing a posteroventral process contacting the cephalic condyle of the quadrate. The posterodorsal process contacting the supratemporal process of the parietal is absent. Parts of both quadrates are preserved in articulation with the squamosal, supratemporal, and probably the supratemporal process of the parietal. The ventral margin of both quadrates has been displaced anteriorly, such that they lie with their anterior surfaces facing dorsally. The right element is better preserved and still preserves a quadrate conch posterolaterally, as well as a distinct tympanic crest. Postcranium The holotype of A. digitatellus has the postcranium preserved mostly as impressions in the calcareous matrix, with a few cervical and dorsal vertebrae, as well as parts of the pelvic girdle preserved as osteological material. Yet, the quality of preservation allows many details of the vertebral and especially the fore- and hindlimb anatomy to be discerned. There are five or six cervical vertebrae (Fig. 5 B), a relatively low cervical vertebral count amongst lizards, and only the posterior-most one or two cervicals bear ribs, as also observed in the holotype of E. schroederi. The total presacral vertebral count is estimated to be 27, followed by two sacrals and six caudals. The anterior cervicals are preserved in dorsal aspect, and have very low neural spines. The cervical pleurocentra are very fragmented, preventing a detailed description, but they are similar in length to the anterior dorsal pleurocentra. The dorsal vertebrae are preserved in lateral aspect, revealing some information about the morphology of their centra, despite some limitations owing to deformation and some degree of shattering. The dorsal centra show no indication of bearing a posterior condyle. A structure resembling a condyle is observed anteriorly on the dorsal vertebrae however, as seen in lateral aspect (Fig. 7 A). In the caudal region, impressions of the anterior caudals also show the anterior border of the vertebrae to be convex, and the posterior border to be concave (Fig. 7 B – D), indicative of opisthocoely, a condition currently unknown for any extant or fossil squamate. However, it could also represent amphicoely with a sediment mould forming in the space between them. Therefore, the form of articulation cannot be determined with confidence, but it certainly is not procoelic as in most squamates. The neural arches are partially preserved and are as tall as the centra. The ribs are single headed and articulated on circular synapophyses of the centra. The dorsal ribs are present up to the level of the penultimate presacral, but the condition in the last presacral cannot be determined owing to poor preservation. The sacral ribs, as well as the anterior caudal ribs, are preserved only as impressions, being laterally orientated and not forked. No elements of the pectoral girdle are preserved, but parts of both forelimbs are preserved as impressions (Fig. 5 A). The humeri, radii, and ulnae do not reveal finer details of their anatomy, but it is possible to detect that the forelimbs were relatively small in relation to SVL (Table 5). The left manus (Fig. 7 E) has digits 3, 4, and 5 preserved, indicating a phalangeal formula of? -? - 4 - 5 - 3. The penultimate phalanges of those digits are elongate compared with the intermediate phalanges (Tables 2 and 4), and are followed by unguals that are curved and tall at their bases. Both ilia are preserved in dorsal view, revealing elongate posterior blades. The impression of the right pubis indicates that this element was narrow and strongly angled anteromedially. No impressions of the pubic tubercle are evident, suggesting that it was very small or absent. The ilia are not preserved either as bony elements or impressions. The hindlimbs are also proportionally short compared with SVL, and similar in length to the forelimbs (Table 5). The femora are relatively slender and not sigmoidal, as was also observed for E. schroederi. The tibia and fibula on both sides are of similar width to each other and diverge distally. The left pes is better preserved than the right, with parts of all digits in articulation, indicating a phalangeal formula of? - 3 - 4 - 5 - 4. As in the forelimbs, the penultimate phalanges are elongate compared with the intermediate ones, being c. 40 % longer than the immediately preceding phalanx in the third digit (Fig. 8 C; Tables 3 and 4). Most articulatory surfaces are not well preserved, but in the left pes, articulation between phalanges I and II in digit II is concave-convex, as reported for E. schroederi above. The unguals are recurved and relatively tall at their bases owing to a ventral expansion beyond the level of contact with the penultimate phalanx, as in G. gecko (Fig. 9) and E. schroederi (see above): pedal claw height equals 0.65 mm. The ratio between claw height / area of contact between the claw and the penultimate phalanx equals 1.625. FUNCTIONAL MORPHOLOGY The functional morphologies of Eichstaettisaurus and Ardeosaurus have never been given extensive attention, despite some particularly interesting aspects of their limb morphology. Russell, Bauer & Laroiya (1997) briefly mentioned some gecko-like aspects of the feet of E. schroederi, suggesting a reduction in length of the fourth metatarsal and a slight divergence between metatarsals III and IV, but did not go further. Relatively short fore- and hindlimbs One of the most apparent aspects of the body form of both E. schroederi and A. digitatellus is the relatively short fore- and hindlimbs (Figs 1 A, B, 5 A), which are also similar in length to each other (see Tables 5 and 6). Short fore- and hindlimbs have been FL, forelimb length; F + T, femoro-tibial length; HL, hindlimb length; H + R, humero-radial length; MT, metatarsal; SVL, estimated snout – vent length. Absolute values represent means calculated from the right and left sides of the holotype. proposed to be functionally advantageous for scansoriality (the capacity to climb) as shorter limbs bring the centre of gravity closer to the substrate and reduce the rotatory moment of the body in relation to the inclined plane (Cartmill, 1985). This is further enhanced by the overall body depression observed in E. schroederi and A. digitatellus, a feature also observed amongst gekkotans, and previously suggested to enhance climbing performance in G. gecko (Russell, 1975) - see also Table 6. However, recent studies have contested the correlation of short fore- and hindlimbs with scansoriality for lacertids and geckoes (Vanhooydonck & Van Damme, 1999; Zaaf & Van Damme, 2001) and this particular feature may instead be related to phylogeny (see below). Fore- and hindlimbs of similar lengths Fore- and hindlimbs of similar lengths to each other may also contribute to stable climbing, as limbs of very distinct lengths would result in different stride lengths, and in the tendency to have fewer limbs maintaining contact with the substrate during fast locomotion. It is important to maintain grip in some climbing lizards, as documented for Lacerta oxycephala, which maintains three or four limbs in contact with the substrate most of the time during fast locomotion (Arnold, 1973). Yet, the latter may not be applicable to geckoes with highly developed adhesive toepads, such as G. gecko, as the latter keeps only two limbs (and occasionally only one) in contact with the substrate most of the time (Russell, 1975). This latter attribute, as displayed by pad-bearing geckoes, may be limited to those climbing lizards with adhesive toepads, owing to their greater clinging capacity in relation to frictional grip relative to other lizards. The correlation between fore- to hindlimb ratio and habitat preference has been tested for lacertid lizards, and no significant correlation was found (Vanhooydonck & Van Damme, 1999). However, as acknowledged by these authors, lacertids may use several other habitats, thus not usually being specialized in that regard. Therefore, the relevance of fore- to hindlimb ratios is dependent on particular clades owing to phylogenetic signal and the level of specialization for a particular habitat, as well as to the influence of functionally related structures (e. g. adhesive toepads). Therefore, fore- to hindlimb ratios, when taken alone, have to be interpreted with caution when trying to differentiate between scansorial and ground-dwelling habits for lizards. Claws tall at their bases Despite some ambiguity regarding the relevance of fore- and hindlimb lengths as adaptations to scansoriality, both the manus and pedes of E. schroederi and A. digitatellus bear claws that are relatively tall at their bases (Figs 4 E, 7 E, 8 A – D, and descriptions with ratios above). This feature has been positively correlated with climbing in lizards, and advocated to be a functional adaptation, even when phylogenetic history is taken into consideration (Zani, 2000; Tulli et al., 2009; Crandell et al., 2014). A possible explanation for this correlation is that higher claws exhibit a ventral expansion relative to the level of contact with the penultimate phalanx, when compared with terrestrial nonscansorial lizards (Fig. 9). The flexor tendon, which runs ventrally and inserts proximally in each phalanx (including the claw), would have a greater lever arm and increase holding strength against the substrate, thus aiding lizards in climbing (Russell, 1975). In addition to height, the claws of A. digitatellus are more elongate and curved relative to those of E. digitatellus. However, the relevance of claw length to clinging performance is debatable, owing to contrasting conclusions (Zani, 2000; Tulli et al., 2009; Crandell et al., 2014), especially when gekkotans (typically with short claws) are considered. Elongate penultimate phalanges	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3520F084B0AF9F4FC22E331.taxon	description	Geckoes bear radiating digits, instead of the subparallel digits evident in most lizards, creating a symmetrical foot in relation to the typical lizard condition (Fig. 8 E, F). This feature is associated with the adhesive toepad climbing mechanism of geckoes, as this facilitates the spreading of the seta-bearing surfaces about a broad arc (Russell, 1975). The latter allows for various combinations of digital orientation that maximize their potential in passive loading, thus aiding in the maintenance of grip in a variety of body orientations (Russell, 1986; Russell et al., 1997; Russell & Oetelaar, 2016). Additionally, a symmetrical foot with digits that radiate distally helps in providing grip, as it allows the first and fifth digits to develop opposability (Robinson, 1975; Rewcastle, 1983). Foot symmetry is achieved by a variety of factors. Whereas most lizards have the first pedal metatarsal (MT I) of about half the length of MT II and III (MT III / MT I length ratio of 2: 1) geckoes have a proportionally longer MT I in relation to MT III, with a MT III / MT I ratio between 1.3 and 1.5, with the greatest average amongst geckoes (1.47) being found in padless diplodactylines (Russell et al., 1997). Eichstaettisaurus schroederi and A. digitatellus display a MT III / MT I ratio of 1.41 and 1.48, respectively (Table 5), falling within the range reported by Russell et al. (1997) for extant geckoes. Another factor contributing to foot symmetry is the reduction in length of MT IV. Whereas most lizards have a MT IV that is longer than MT III, geckoes have a MT IV shorter than MT III (Russell et al., 1997; Russell & Bauer, 2008). Some varanids have similar lengths for MT III and IV (Russell et al., 1997). This is also seen in Heloderma, but in both cases they are usually never shorter than MT III (T. R. Simoes, pers. observ.). Reduction in length of MT IV is observed in E. schroederi (Fig. 8 A – C), and may be the case for A. digitatellus (Fig. 8 C) as well, although this cannot be confirmed. Other features related to foot symmetry are the broadened proximal head of MT IV, which greatly increases the angle between the shafts of MT III and MT V, as well as the reduction of imbrication amongst the metatarsals proximally. Both features help to create an expanded digital arc, and contribute to foot symmetry (Russell et al., 1997). These features, however, are not observable for either E. schroederi or A. digitatellus. Eichstaettisaurus schroederi has a somewhat broadened proximal head of MT IV, but it does not seem to be proportionally larger than the condition seen in Iguana iguana and most other lizards. Therefore, both species display partial development of foot symmetry, which is more developed in E. schroederi. A few important considerations regarding the foot symmetry of E. schroederi and A. digitatellus are relevant to understanding their consequence for the locomotion of these lizards, even though they are not necessarily linked to scansorial habits. Metatarsals of quite distinct lengths contribute to the highly asymmetrical feet of most lizards. The distal tips of the metatarsals form a straight metatarsophalangeal line (Fig. 8 E), which is directed perpendicular to the parasagittal plane at rest. During limb retraction in sprawling locomotion this aids in maintaining the first three digits in contact with the substrate and provides even support amongst these digits for bearing the animal’s body weight (Brinkman, 1980; Rewcastle, 1983). If they were of equal length, most of the weight would be concentrated on the first digit only, as a consequence of the lateral orientation of the femur. Metatarsals of similar length are usually observed only in lizards with more anteriorly orientated feet (Brinkman, 1980). Therefore, the highly symmetrical metatarsals of E. schroederi and A. digitatellus indicate that their feet were probably more anteriorly, rather than laterally, orientated. Another character of laterally orientated feet is the development of the ginglymoid bicondylar articulation observed with greater development on the first three pedal digits of most lizards. These joints, along with tendinous bands that lie along the lateral sides of the digits contribute to resisting lateral displacement while enabling dorsoventral flexion of the FL, forelimb length; HL, hindlimb length; SVL, estimated snout – vent length. Source references indicate studies where measurements were obtained, or calculated from. phalanges (Russell, 1975; Landsmeer, 1981). This is important considering that the posteriorly directed thrust during limb retraction has a perpendicular orientation relative to the laterally orientated phalanges (Rewcastle, 1983) – see Fig. 8 E and F – thus an interlocking mechanism represented by the bicondylar articulations provides greater stability. In lizards with more anteriorly orientated feet, this morphology would have a reduced adaptive significance. Bicondylar articulations are not observed in E. schroederi or A. digitatellus, which have convexconcave joints in the intermediate phalanges where articulatory surfaces are well preserved. This is another indication that E. schroederi and A. digitatellus had more anteriorly orientated feet than most lizards, which bear bicondylar articulations. Finally, in E. schroederi and A. digitatellus the long axis of the first metatarsal makes a right angle with that of the tibia, a condition observed in G. gecko. This is compatible with anteriorly orientated feet. Although it is possible that this latter anatomy is an artefact of post-mortem changes in orientation, the metatarsal proportions and shape of interphalangeal articulations do support interpretation of the pedes of E. schroederi and A. digitatellus as having an anteromedial orientation similar to that found in geckoes and some platynotans. In conclusion, there are limitations in differentiating ground-dwelling from scansorial lizard species based on limb to SVL ratios, as well as fore- to hindlimb length ratios, as discussed above. However, a dorsoventrally expanded claw and elongate penultimate phalanges have a significant correlation to habitat usage in lizards and other reptiles (Zani, 2000; Tulli et al., 2009; Kavanagh et al., 2013; Crandell et al., 2014). Therefore, the combination of the latter, along with a depressed body and skull in E. schroederi and A. digitatellus, provides a suite of features that are usually found together only in species specialized to climbing. Additionally, metatarsal proportions indicate the feet of E. schroederi and A. digitatellus were oriented more anteriorly than in most lizards.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3400F024C59FE26FB2FE3D1.taxon	description	Ardeosaurus digitatellus ??????????????????????????? ????????????????????? 200 ??? 01110 ?? 21? 0? 00 ???? 0? 000 ????? 0 ??? 01? 2 ?? 001? 000? 01100000 ????????????????? 00? 00? 0 ???? 1 ???? 0 ????????? 002 ???? 0? 010010 [12] 00? 0? 0000 ?? 001 ???? 0 ???????????????????????????????????????????? 0 ????????????????????????????????????????????????????? - ??????????????????????????????????????????????????????? - ??????????????????????????????????????????????????????? - ??????????????????????????????????????????????????????? - ???? 03000? 0? 0 ?????? 0? 0 ?????? 0 ??????????????????????????????? 000 ??????? 0? 00 ??? 001 ?????????? 0? 00001? 000 ??????? 0 ???? 00? 000 ??? 0 ?? 0? 0000? 0 ???????????????????????????? APPENDIX 2	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
03FA87B7B3400F024C59F9C2FC6BE17A.taxon	description	Ardeosaurus _ digitatellus: Char. 49: 0 –> 2, Char. 62: 0 –> 2, Char. 63: 0 –> 1, Char. 83: 2 –> 0, Char. 88: 0 –> 1, Char. 154: 1 –> 2, Char. 167: 0 –> 12, Char. 455: 4 –> 3.	en	Simões, Tiago R., Caldwell, Michael W., Nydam, Randall L., Jiménez-Huidobro, Paulina (2017): Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes. Zoological Journal of the Linnean Society 180 (1): 216-241, DOI: 10.1111/zoj.12487, URL: https://doi.org/10.1111/zoj.12487
