Postcranial anatomy of Haldanodon exspectatus (Mammalia, Docodonta) from the Late Jurassic (Kimmeridgian) of Portugal and its bearing for mammalian evolution

The postcranium of the Late Jurassic docodont Haldanodon exspectatus Kühne & Krusat, 1972 is represented by a partial skeleton and isolated bones of other individuals from the Late Jurassic (Kimmeridgian) of the Guimarota coal mine in Portugal. Haldanodon exhibits adaptations for a fossorial lifestyle such as stout and short limb bones and humeri with greatly expanded distal joints and strong deltopectoral crests. Short first and second phalanges and moderately curved and laterally compressed terminal phalanges with lateral grooves suggest that Haldanodon was a scratch-digger. The dorso-ventrally elongated, triangular scapula has a convex transverse profile with strongly laterally reflected anterior and posterior scapula margins, enclosing a deep trough-like 'infraspinous fossa'. A supraspinous fossa is not developed. The saddle-shaped glenoid facet is mainly formed by the coracoid and orientated antero-ventrally indicating a sprawling gait. No epiphyses were detected and the wide size range of humerus and femur possibly indicate a lifelong growth. Haldanodon is more derived than Morganucodon by complete reduction of the procoracoid, absence of the procoracoid foramen, and a peg-like coracoid. It shares with monotremes a postscapular fossa that is absent in Morganucodon . A PAUP analysis based on 280 cranio-dental and postcranial characters corroborated the position of Haldanodon above morganucodontids and below Hadrocodium .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 219–248.     

Functional aspects of the postcranial anatomy of the Permian dicynodont Diictodon and their ecological implications

The postcranial skeleton of the Permian dicynodont Diictodon is described, major skeletal muscles restored and functional aspects of the skeleton analyzed. The humerus was caudolaterally oriented and articulated in a near sagittal plane. This reduced the transverse component of thrust during locomotion. Throughout the stride, the femur, which is relatively longer than the humerus, was held in a parasagittal position. Diictodon probably had a hindlimb gait similar to the high walk of crocodilians. Comparisons between several dicynodont genera, including Diictodon, reveal considerable variation in the limb orientations, articulations, posture and gait among these taxa. Expansion of the preacetabular iliac process and the overall iliac length with respect to its height among these dicynodonts may be correlated with the increase in the number of the sacral vertebrae. Diictodon does not show the cranial and forelimb modifications suitable for digging as seen in Cistecephalus, Kawingasaurus and extant burrowers such as a broad skull, rounded occiput for considerable neck musculature, a robust humerus and a prominent olecranon process. However, a cylindrical body, short limbs with well–developed limb elevators and depressors and a long wide manus with long, blunt claws probably helped in digging. In addition, the hindlimb with well–developed retractors and short, blunt claws participated in soil removal. The caudolaterally oriented humerus and a laterally flexible vertebral column facilitated movement in the narrow confines of the burrows. Forelimb articulation and morphology indicate that its method of digging was probably rotation–thrust, where humeral excursion and not forearm extension played a dominant role. The close associations of articulated paired skeletons of adult individuals suggest that Diictodon was gregarious. A large number of unconnected burrow–casts in a small area of about 500 m², indicates that although the animals lived close together, they did not share a single large communal structure.     

An Early Pleistocene human pedal phalanx from Swartkrans,SKX 16699, and the antiquity of the human lateral forefoot

In order to assess the antiquity of derived human lateral (lesser) toe morphology, the SKX 16699 Early Pleistocene pedal proximal phalanx from Swartkrans (South Africa) was compared to samples of pedal phalanges attributed to Pliocene/Pleistocene australopithecines, Homo naledi and Late Pleistocene Homo. In contrast to australopith lateral phalanges, the SKX 16699 phalanx exhibits an absolutely (and probably relatively) short length, limited plantar diaphyseal curvature, proximal-to-midshaft and mid-dorsoplantar flexor sheath insertions, and a marked proximodorsal orientation of the metatarsal facet. SKX 16699 is intermediate between the australopith phalanges and later Homo ones in its modest dorsal diaphyseal curvature and mid-dorsoplantar metatarsophalangeal collateral ligament insertion areas. Its diaphyseal robustness is similar to that of Homo phalanges, but overlaps the range of later australopith ones. This combination of features and the close morphological affinities of SKX 16699 to later Homo proximal pedal phalanges suggest the emergence of a distinctly human lateral forefoot by the initial Early Pleistocene.     

The postcranial anatomy of Suminia getmanovi (Synapsida: Anomodontia), the earliest known arboreal tetrapod

The basal anomodont Suminia getmanovi Ivakhnenko, 1994 from the late Palaeozoic of Russia is highly specialized in its masticatory apparatus, and has been suggested to represent the earliest arboreal tetrapod in the fossil record. Its postcranial anatomy is described in detail for the first time, revealing a large number of autapomorphies for this small herbivore. These include a reduced number of presacral and therein dorsal vertebrae, an elongate neck, a long and possibly prehensile tail, a procoracoid with a notch at its ventromedial margin rather than a foramen, an iliac blade with a robust ridge at its anteromedial edge, a pubis with a puboischiadic fenestra and separate pubic foramen, and elongate limbs. Additional autapomorphic characters are displayed in the autopodium, which comprises about 40% of the entire limb length. These features include an enlarged, phalangiform distal carpal 1 and tarsal 1, a short and robust first metacarpal, a crescent‐shaped distal tarsal 4, and elongate penultimate phalangeal elements. The phylogenetic relationships of basal anomodonts are revisited using an expanded data set, with the addition of key taxa and several postcranial characters. Unlike dicynodonts, Suminia retained the plesiomorphic phalangeal formula for amniotes of 2‐3‐4‐5‐3 (manus) and 2‐3‐4‐5‐4 (pes). This pattern is achieved by the retention of disc‐like phalangeal elements between the proximal and penultimate phalanges in digits III, IV (manus and pes), and V (pes only). In light of the new material, Suminia can be recognized as the most complete basal anomodont, offering new insights into the early evolution of the group. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 661–698.     

The African species Megantereon whitei from the Early Pleistocene of Monte Argentario (South Tuscany,Central Italy)

A partial skull and articulated postcranial elements of Megantereon whitei have been recorded during the 1950s from a karst deposit in the Monte Argentario area (Grosseto). These fossils recently became available for study. The bones are quite well preserved, included in a hard reddish matrix with calcareous clasts. The fossil is part of a faunal assemblage referred to the Late Villafranchian (Early Pleistocene). M. whitei from Monte Argentario is characterised by elongated upper canines, stronger than those of the Upper Valdarno (Tuscany) specimens and similar to the Pirro Nord (Apulia) fossil. The manus is robust, the first phalanges are quite long in comparison to the metacarpals. The Early Pleistocene European M. whitei represents an African element that took part in the faunal dispersal from Africa to Europe that occurred at the Plio-Pleistocene transition.     

On the congruence of morphology,trophic ecology,and phylogeny in Neotropical treeboas (Squamata: Boidae: Corallus)

Nine members of the Neotropical treeboa genus Corallus occur from Guatemala to south‐eastern Brazil and recent studies have provided an inconclusive picture about the relationship between morphology and trophic ecology in these snakes. To construct a more complete picture, we conducted the first study of morphology and diet to consider all nine species. Using adult specimens from museum collections, we examined several morphometric and meristic variables and their possible relationship to Corallus diets. Broadly, we found three basic morphologies within the genus: a short, narrow head and a slender body (C. cookii, C. grenadensis, C. hortulanus, and C. ruschenbergerii), useful for exploiting a wide variety of prey; a relatively stout body with a long, wide head (C. batesii, C. caninus, and C. cropanii) associated with feeding on larger mammalian prey; and an intermediate morphology, found in C. annulatus and C. blombergii, which may be indicative of endotherm generalists. These morphological and dietary patterns exhibit a strong degree of congruence with a recent molecular phylogeny of Corallus and highlight a heretofore unexamined ecological diversification within Corallus. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109, 466–475.     

The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo‐ecological interpretation

The rediscovered holotype skulls of Late Pleistocene Panthera leo spelaea ( Goldfuss, 1810 ) (Felidae) and Crocuta crocuta spelaea ( Goldfuss, 1823 ) (Hyaenidae) from the Zoolithen Cave at Burggeilenreuth, southern Germany, are discussed. The cave became famous mainly due to its rich cave bear bone remains from the late Saalian (OIS 6–8) to Eemian/Weichselian (OIS 3–6) including additionally a third holotype of Ursus spelaeus Rosenmüller, 1794 (Ursidae). The ‘Felis spelaea’ holotype represents an adult male with a strong bite mark on the saggital crest, which was in an early stage of healing. Compared with other European Late Pleistocene lion skulls and skeletons, and with modern African lions, it provides evidence of intraspecific conflict between male Ice Age lions. The holotype of ‘Hyaena spelaea’ is one of several hundred hyena remains from a well‐frequented hyena den cave. The cave was used intensively by Late Pleistocene hyena clans, for collecting lion carcasses in addition to their accustomed prey, as happened in many caves throughout Europe. Ice Age spotted hyena clans might have killed Ice Age steppe lions for many reasons, such as fights over prey and territory, and the protection of cubs, but they did not always scavenge on their carcasses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 822–831.     

Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores

Among the hunting strategies employed by members of the order Carnivora (Mammalia), two, stalk and ambush and sustained pursuit, are particularly prevalent among larger species of the order. It has been difficult to identify morphological traits that support this distinction and ecological observations have shown that most carnivorans adopt a continuum of strategies, depending on available habitat and prey. In this paper, the shape of the distal humerus articulation is analysed, with the aim of exploring the use of the forelimb in prey procurement, and as a guide to such behaviour among fossil carnivorans. The results suggest that manual manipulation and locomotion are conflicting functions. Elbow‐joint morphology supports a division between grapplers (i.e. ambushers) and nongrapplers (i.e. pursuers). Joints of the former are characterized by being relatively wide and the latter, by being relatively narrow and box‐like with pronounced stabilizing features. At intermediate and large body sizes, carnivorans show a pattern suggesting mutually exclusive feeding strategies that involve either grappling with prey or sustained pursuit. The former allows for large body sizes, such as pantherine felids and ursids; the latter includes species of only moderate size, such as hyenids and canids. Elbow‐joint morphology is closely linked to phylogeny, but the morphology of the cheetah converges with that of nongrapplers, showing that strong selective forces may override the phylogenetic component. Two taxa of giant mustelids from the Miocene were analysed to test whether this sort of analysis is applicable to carnivorans of the past. The African Late Miocene species Ekorus ekakeran has a joint morphology comparable to that of modern‐day nongrapplers. Two joint morphologies were found in the North American Late Oligocene‐Early Miocene Megalictis ferox. The first morphology is comparable to that of modern pantherine cats and the second forms an intermediate between grapplers and nongrapplers that is not present in the modern carnivoran fauna. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 91–104.     

Postcranial morphology,functional adaptations and palaeobiology of Callistoe vincei,a predaceous metatherian from the Eocene of Salta,north‐western Argentina

Abstract: Callistoe vincei Babot et al., 2002 is a Paleogene borhyaenoid known from exceptionally complete postcranial elements, which provides rare information about the anatomy and evolutionary history of metatherian predators during the South American Cenozoic. The axial skeleton of Callistoe is characterized by the peculiar transverse processes of the cervical vertebrae emphasizing lateral instead of sagittal traction. There is no clavicle and eighteen thoracolumbar vertebrae, of which only five are lumbars. The shoulder and elbow joints suggest movements restricted to parasagittal flexion/extension that are consistent with primarily terrestrial locomotion, as is also emphasized in Borhyaena tuberata and Lycopsis longirostrus. On the manus, the pollex is not reduced and the ungual phalanges indicate very long claws, similar to those observed in some extant digging taxa. This feature is unique to C. vincei among borhyaenoids. The knee joint is characterized by the presence of ossified patellae but shallow femoral trochleae. This joint suggests that the leg was nearly parasagittal, a position also inferred for Borhyaena. The astragalus shape is consistent with parasagittal flexion/extension, as in all Miocene–Pliocene borhyaenoids. The hind foot is characterized by reduced claws in comparison with the manus as well as the slenderness of the first and fifth digits, another peculiarity of C. vincei. The habitat of Callistoe was a temperate humid forest and according to the known fossil record, Callistoe was the largest mammalian predator of its time, sharing the predator ecological niche with crocodiles.     

Locomotory abilities and habitat of the Cretaceous bird Gansus yumenensis inferred from limb length proportions

The relative length proportions of the three bony elements of the pelvic (femur, tibiotarsus and tarsometatarsus) and pectoral (humerus, ulna and manus) limbs of the early Cretaceous bird Gansus yumenensis, a well‐represented basal ornithuromorph from China, are investigated and compared to those of extant taxa. Ternary plots show that the pectoral limb length proportions of Gansus are most similar to Apodiformes (swifts and hummingbirds), which plot away from all other extant birds. In contrast, the pelvic limb length proportions of Gansus fall within the extant bird cluster and show similarities with the neornithine families Podicipedidae (grebes), Diomedeidae (albatross) and Phalacrocoracidae (cormorants). Although it does have some of the pelvic limb features of grebes and cormorants, the femur of Gansus is more gracile and is thus more consistent with an albatross‐like shallow‐diving mode of life than a strong foot‐propelled diving movement pattern. The position of Gansus in pectoral limb ternary morphospace is largely due to its elongated manus. In contrast to apodiformes, where the humerus and ulna are short and robust, an adaptation, which provides a stiff wing for their demanding fast agile and hovering flight (respectively), the wing‐bones of Gansus are slender, indicating a less vigorous flapping flight style. The suite of characters exhibited by Gansus mean it is difficult to completely interpret its likely ecology. Nevertheless, our analyses suggest that it is probable that this bird was both volant and capable of diving to some degree using either foot‐propelled or, perhaps, both its wings and its feet for underwater locomotion.     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

Phylogeny and diversification of the cloud forest Morpho sulkowskyi group (Lepidoptera,Nymphalidae) in the evolving Andes

The monophyletic Morpho sulkowskyi butterfly group, endemic of Andean cloud forests, was studied to test the respective contributions of Mio‐Pliocene intense uplift period and Pleistocene glacial cycles on Andean biodiversity. We sampled nine taxa covering the whole geographical range of the group. Two mitochondrial and two nuclear genes were analysed using a Bayesian method. We established a dated phylogeny of the group using a relaxed clock method and a wide‐outgroup approach. To discriminate between two hypotheses, we used a biogeographical probabilistic method. Results suggest that the ancestor of the M. sulkowskyi group originated during the Middle–Late Miocene uplift of the Eastern Cordillera in northern Peru. Biogeographical inference suggests that the M. sulkowskyi and Morpho lympharis clades diverged in the northern Peruvian Andes. The subsequent divergences, from the Late Miocene to the Late Pliocene, should have resulted from a dispersal towards the Northern Andes (M. sulkowskyi clade), after the closure of the West Andean Portal separating the Central and Northern Andes, and a southwards dispersal along the Peruvian and Bolivian Eastern Cordilleras (M. lympharis clade). Only a few divergences occurred at the very end of the Pliocene or during the Pleistocene, a period when the more recent uplifts interfered with Pleistocene glacial cycles.     

Functional implications of craniomandibular morphology in African mole‐rats (Rodentia: Bathyergidae)

African mole‐rats are subterranean rodents from the family Bathyergidae. The family consists of six genera, five of which (Cryptomys, Fukomys, Georychus, Heliophobius and Heterocephalus) are chisel‐tooth diggers, meaning they dig underground using procumbent incisors. The remaining genus of mole‐rat (Bathyergus) is a scratch digger, which digs using its forelimbs. Chisel‐tooth digging is thought to have evolved to enable exploitation of harder soils. It was hypothesized that to dig successfully using incisors, chisel‐tooth digging mole‐rats will have a craniomandibular complex that is better able to achieve a large bite force and wide gape compared with scratch digging mole‐rats. Linear measurements of morphological characteristics associated with bite force and gape were measured in several chisel‐tooth digging and scratch digging mole‐rats. Chisel‐tooth diggers have increased jaw and condyle lengths relative to their size (characteristics associated with larger gape). They also have relatively wider and taller skulls (characteristics associated with larger bite force). The mechanical advantage of three masticatory muscles of each specimen was also calculated. The mechanical advantage of the temporalis muscle was significantly larger in chisel‐tooth digging mole‐rats than scratch digging genus. The results demonstrate that chisel‐tooth digging bathyergids have a craniomandibular morphology that is better able to facilitate high bite force and wide gape than scratch digging mole‐rats.     

Pterosaur Stance and Gait and the Interpretation of Trackways

The tracks ascribed to pterosaurs from the Late Jurassic limestones at Crayssac, France, must be pterosaurian because the manus prints are so far outside those of the pes, the pes print is four times longer than wide, and the manus prints appear to preserve distinct traces of a posteromedially directed wing-finger. These tracks are different in important ways from previously described Pteraichnus trackways, which have been variably considered pterosaurian, crocodilian, or indeterminate. No Pteraichnus (sensu stricto: those not from Crayssac) tracks have diagnostic features of pterosaurs and in none can a complete phalangeal or digital formula be reconstructed; however, all published Pteraichnidae tracks fulfill the criteria of poor preservation, and some have some diagnostic features of crocodile tracks. Reconstructions of pterosaurs walking in pteraichnid tracks do not fit those tracks well, but crocodiles do. In contrast, the Crayssac tracks demonstrate the erect stance and parasagittal gait previously reconstructed for pterosaurs. They also demonstrate that the footfall pattern was not as in typical reptiles (LH-RF-RH-LF), but that the manus must have been raised before the next forward step of the ipselateral foot (LH-LF-RH-RF), suggesting that the quadrupedal pattern was secondary. The metatarsus in pterosaurs was set low at the beginning of a stride, as it is in crocodilians and basal dinosaurs. The diagnosis of the Ichnofamily Pteraichnidae comprises features of possible crocodilian trackmakers, but not of possible pterosaurian trackmakers. Trackways considered for attribution to pterosaurs should show (1) manus prints up to three interpedal widths from midline of body, and always lateral to pes prints, (2) pes prints four times longer than wide at the metatarso-phalangeal joint, and (3) penultimate phalanges longest among those of the pes.     

The position of the claws in Noasauridae (Dinosauria: Abelisauroidea) and its implications for abelisauroid manus evolution

In this note we reassess the position of putative pedal phalanges of some South American noasaurid theropods (Abelisauroidea). Noasaurids were considered as to be distinctive abelisauroids with a peculiar “sickle claw” on the second toe of the foot, convergently developed with that of deinonychosaurians. Among noasaurids, the Argentinean species Noasaurus leali (latest Cretaceous) and Ligabueino andesi (Early Cretaceous) are known from incomplete specimens, including dissarticulated non-ungueal phalanges, and, in N. leali, a claw. A detailed overview of these elements indicates that the supposed raptorial claw of the second pedal digit of N. leali actually belongs to the first or second finger of the manus, and the putative pedal non-ungual phalanges of both genera also pertain to the manus. Thus, the new interpretations of noasaurid pedal morphology blur the distinctions between Noasauridae and Velocisauridae proposed by previous authors. Finally, we suggest, on the basis of phalangeal and metacarpal morphology, that abelisaurids probably lost their manual claws by means of the loss of function of the HOXA11 and HOXD11 genes. Thus Noasauridae differs from Abelisauridae in retaining plesiomorphic long forelimbs with well developed claws, as occurs plesiomorphically in most basal theropods (e.g., Coelophysis).     

An unusual pattern of limb morphology in the Tithonian marine turtle Neusticemys neuquina from the Vaca Muerta Formation,Neuquén Basin,Argentina

de la Fuente M.S. & Fernández, M.S. 2010: An unusual pattern of limb morphology in the Tithonian marine turtle Neusticemys neuquina from the Vaca Muerta Formation, Neuquén Basin, Argentina. Lethaia, Vol. 44, pp. 15–25. Here, we report an unusual pattern in the manus and pes morphology of the Tithonian marine turtle Neusticemys neuquina. We analyse the forelimbs of two previously known specimens and describe the hind limbs of two previously undescribed specimens. Neusticemys neuquina is characterized by a relative elongation of both the forelimb and hind limb, compared with stem Chelonoidea, as well as an elongation of the pedal digit V, achieved through the elongation of the bones, as well as a moderate hyperphalangy. The elongation of pedal digit V is the most striking feature of N. neuquina, a feature unknown in other turtles. □Limb morphology, Neusticemys neuquina, Tithonian turtle.     

Mummified fossil of Keteleeria from the Late Pleistocene of Maoming Basin,South China,and its phytogeographical and paleoecological implications

Keteleeria is a small genus of Pinaceae now mainly restricted to eastern Asia. Although this genus has been documented with a wide distribution in the geologic record of Europe, North America, and Asia, its history in low‐latitude areas (including South China) has remained obscure. In this paper, a fossil wood of Keteleeria sp. is described from the Late Pleistocene (29–27 ka BP) of the Maoming Basin, South China. This wood is the most ancient megafossil evidence of Keteleeria within the modern distribution area of this genus. The fossil records of Keteleeria suggests that this thermophyllous genus migrated into South China by the Middle Pleistocene escaping from glacial cooling and became widespread over this region in the Late Pleistocene beginning from the interglacial stage preceding the Last Glacial Maximum. The analysis of growth rings in the fossil wood and its comparison with those of modern Keteleeria davidiana (Bertrand) Beissner indicates that in the Late Pleistocene of Maoming Basin (29–27 ka BP) there was a humid climate with less pronounced seasonality of precipitation than that seen in the subtropical monsoonal climate of modern northeastern Vietnam. Apparently, the Maoming Basin was influenced by interglacial regime with summer–monsoon circulation. The previously proposed method to distinguish between evergreen and deciduous conifers based on growth ring anatomy, is not reliable because of the wide variance and ambiguity in its results.     

Late Triassic Aetosaurs as the Trackmaker of the Tetrapod Footprint Ichnotaxon Brachychirotherium

Brachychirotherium is the common ichnogenus of Late Triassic chirothere footprints well known from western Europe, North America, Argentina and South Africa. Although it has long been agreed by most workers that the trackmaker of Brachychirotherium was a derived crurotarsan archosaur, the trackmaker has been identified as either a rauisuchian or an aetosaur, and some workers attribute it to a primitive crocodylomorph (sphenosuchian). New knowledge of the osteology of the manus and pes of a large aetosaur, Typothorax coccinarum, indicates a close correspondence between the manus and pes structure of aetosaurs and the morphology of Brachychirotherium. Furthermore, functional analysis of complete skeletons indicates aetosaurs plausibly placed their feet in the narrow gauge, nearly the overstepped walk characteristic of Brachychirotherium. Brachychirotherium and aetosaurs have matched distributions, that is, they were Pangea-wide during the Late Triassic. The manus and pes morphology of rauisuchians and early crocodylomorphs (sphenosuchians) deviate from Brachychirotherium footprint morphology in key features, thus excluding their identification as trackmakers. Aetosaurs made Brachychirotherium footprints.