Raman Spectra and Ancient Life: Vibrational ID Profiles of Fossilized (Bone) Tissues.

Raman micro-spectroscopy is a non-destructive and non-contact analytical technique that combines microscopy and spectroscopy, thus providing a potential for non-invasive and in situ molecular identification, even over heterogeneous and rare samples such as fossilized tissues. Recently, chemical imaging techniques have become an increasingly popular tool for characterizing trace elements, isotopic information, and organic markers in fossils. Raman spectroscopy also shows a growing potential in understanding bone microstructure, chemical composition, and mineral assemblance affected by diagenetic processes. In our lab, we have investigated a wide range of different fossil tissues, mainly of Mesozoic vertebrates (from Jurassic through Cretaceous). Besides standard spectra of sedimentary rocks, including pigment contamination, our Raman spectra also exhibit interesting spectral features in the 1200-1800 cm-1 spectral range, where Raman bands of proteins, nucleic acids, and other organic molecules can be identified. In the present study, we discuss both a possible origin of the observed bands of ancient organic residues and difficulties with definition of the specific spectral markers in fossilized soft and hard tissues.

A new specimen of Sinopterus dongi (Pterosauria, Tapejaridae) from the Jiufotang Formation (Early Cretaceous, China).

The Tapejarinae are edentulous pterosaurs that are relatively common in Cretaceous continental deposits in South America, North Africa, Europe, and China (mostly Early Cretaceous). The Chinese Jiufotang Formation is particularly rich in tapejarine specimens, having yielded over 10 described specimens and dozens of undescribed ones. For the Jiufotang Formation, a total of seven nominal tapejarid species and two genera have been proposed. Some debate exists over how many of those are valid or, alternatively, sexual or ontogenetic morphs of fewer (or even a single) species. Despite the abundance of specimens and the relevant taxonomic problems involved, detailed revisions of the matter are still lacking. This is partly due to the relatively scarce knowledge on the comparative osteology of the Sinopterus complex, which is hampered by the fact that most specimens have been only preliminarily described. In this contribution, we present a new postcranial specimen, D3072, which we attribute to the type-species of the genus, Sinopterus dongi. This new specimen helps shed some new light in the osteology of Sinopterus dongi, hopefully serving as a basis for future comparative studies involving further specimens and other proposed species and, subsequently, taxonomic revisions.

Earliest migratory cephalic NC cells are potent to differentiate into dental ectomesenchyme of the two lungfish dentitions: tetrapodomorph ancestral condition of unconstrained capability of mesencephalic NC cells to form oral teeth.

Reciprocal interactions between epithelial and neural crest-derived mesenchymal cells have been recognized in the evolutionary modulation of tetrapod odontodes, skeletal structures that include the teeth and tooth-integrated basal tissue. Using cell-tracking experiments, it has been demonstrated that mandibular neural crest cells, labelled during migration, extensively populate dental papillae of all tooth phenotypes of the lobe-finned fish, the Australian lungfish (Neoceratodus forsteri). Here, I report on an extension of this experimental study that earliest migrating NC cells are able to differentiate into odontogenic ectomesenchyme. Using vital dye cell-tracking to mark the mesencephalic neural crest prior to migration, I have found that the corresponding population of earliest migratory cells selectively relocated to dental papillae of both temporary and permanent dentitions of Neoceratodus. I noticed a gradient in distribution of the labelled cells which populated posterior teeth, pterygoid and prearticular (including associated trabecular and Meckelian cartilages; major relocation) much more densely than those in anterior marginal positions, temporary and vomeral permanent teeth (minor relocation). Contrary to mice and zebrafish, the odontogenic potency of mesencephalic neural crest cells is already programmed at the onset of the migration event in lungfish. This may imply that the morphogenic potential of mesencephalic neural crest cells to form teeth has been heterochronically shifted and constrained to later migratory populations of neural crest cells during the developmental evolution of derived tetrapods, or/and arrested in their expression in the oral development of some modern osteichthyans.

Specialized Craniofacial Anatomy of a Titanosaurian Embryo from Argentina.

The first dinosaur embryos found inside megaloolithid eggs from Auca Mahuevo, Patagonia, were assigned to sauropod dinosaurs that lived approximately 80 million years ago. Discovered some 25 years ago, these considerably flattened specimens still remain the only unquestionable embryonic remains of a sauropod dinosaur providing an initial glimpse into titanosaurian in ovo ontogeny. Here we describe an almost intact embryonic skull, which indicates the early development of stereoscopic vision, and an unusual monocerotic face for a sauropod. The new fossil also reveals a neurovascular sensory system in the premaxilla and a partly calcified braincase, which potentially refines estimates of its prenatal stage. The embryo was found in an egg with thicker eggshell and a partly different geochemical signature than those from the egg-bearing layers described in Auca Mahuevo. The cranial bones are comparably ossified as in previously described specimens but differ in facial anatomy and size. The new specimen reveals significant heterochrony in cranial ossifications when compared with non-sauropod sauropodomorph embryos, and demonstrates that the specialized craniofacial morphology preceded the postnatal transformation of the skull anatomy in adults of related titanosaurians.

Ultraviolet light illuminates the avian nature of the Berlin Archaeopteryx skeleton.

The question of whether the iconic avialan Archaeopteryx was capable of active flapping flight or only passive gliding is still unresolved. This study contributes to this debate by reporting on two key aspects of this fossil that are visible under ultraviolet (UV) light. In contrast to previous studies, we show that most of the vertebral column of the Berlin Archaeopteryx possesses intraosseous pneumaticity, and that pneumatic structures also extend beyond the anterior thoracic vertebrae in other specimens of Archaeopteryx. With a minimum Pneumaticity Index (PI) of 0.39, Archaeopteryx had a much more lightweight skeleton than has been previously reported, comprising an air sac-driven respiratory system with the potential for a bird-like, high-performance metabolism. The neural spines of the 16th to 22nd presacral vertebrae in the Berlin Archaeopteryx are bridged by interspinal ossifications, and form a rigid notarium-like structure similar to the condition seen in modern birds. This reinforced vertebral column, combined with the extensive development of air sacs, suggests that Archaeopteryx was capable of flapping its wings for cursorial and/or aerial locomotion.

HNK-1 in Morphological Study of Development of the Cardiac Conduction System in Selected Groups of Sauropsida.

Human natural killer (HNK)-1 antibody is an established marker of developing cardiac conduction system (CCS) in birds and mammals. In our search for the evolutionary origin of the CCS, we tested this antibody in a variety of sauropsid species (Crocodylus niloticus, Varanus indicus, Pogona vitticeps, Pantherophis guttatus, Eublepharis macularius, Gallus gallus, and Coturnix japonica). Hearts of different species were collected at various stages of embryonic development and studied to map immunoreactivity in cardiac tissues. We performed detection on alternating serial paraffin sections using immunohistochemistry for smooth muscle actin or sarcomeric actin as myocardial markers, and HNK-1 to visualize overall staining pattern and then positivity in specific myocyte populations. We observed HNK-1 expression of various intensity distributed in the extracellular matrix and mesenchymal cell surface of cardiac cushions in most of the examined hearts. Strong staining was found in the cardiac nerve fibers and ganglia in all species. The myocardium of the sinus venosus and the atrioventricular canal exhibited transitory patterns of expression. In the Pogona and Crocodylus hearts, as well as in the Gallus and Coturnix ones, additional expression was detected in a subset of myocytes of the (inter)ventricular septum. These results support the use of HNK-1 as a conserved marker of the CCS and suggest that there is a rudimentary CCS present in developing reptilian hearts. Anat Rec, 302:69-82, 2019. © 2018 Wiley Periodicals, Inc.

An Intermediate Incubation Period and Primitive Brooding in a Theropod Dinosaur.

Non-avian dinosaurs such as oviraptorosaurs and troodontids share several important reproductive characters with modern birds, including eggshell microstructure and iterative egg production. Nevertheless, debate exists concerning their incubation strategies. Here we estimate incubation period for the troodontid, Troodon formosus, by examining a near-term embryonic tooth. Synchrotron scanning and histologic thin sections allowed counting of daily (von Ebner) growth lines. The tooth preserves 31 intact lines with an average spacing of 3.3 ± 0.96 µm. Adding 8 more for the missing crown tip gives a total age of 39 days. Modern crocodilians begin to establish their functional dentition at approximately 47% through incubation. Thus, this tooth age suggests a Troodon incubation period of 74 days, falling midway between avian (44.4 days) and reptilian (107.3 days) values predicted by the Troodon egg mass (314 g). An accelerated incubation relative to modern reptiles supports brooding and concurs with a suite of features in oviraptorosaurs and troodontids (sequential laying, large complex clutches, and precocial young) that appear dependent upon both adult body and incubation temperatures elevated over ambient conditions. However, the largely buried condition of Troodon clutches may have prohibited efficient brooding, necessitating longer incubation than that of modern birds with fully exposed eggs.

Inner tooth morphology of Homo erectus from Zhoukoudian. New evidence from an old collection housed at Uppsala University, Sweden.

Locality 1, in the Lower Cave of the Zhoukoudian cave complex, China, is one of the most important Middle Pleistocene paleoanthropological and archaeological sites worldwide, with the remains of c. 45 Homo erectus individuals, 98 mammalian taxa, and thousands of lithic tools recovered. Most of the material collected before World War II was lost. However, besides two postcranial elements rediscovered in China in 1951, four human permanent teeth from the 'Dragon Bone Hill,' collected by O. Zdansky between 1921 and 1923, were at the time brought to the Paleontological Institute of Uppsala University, Sweden, where they are still stored. This small sample consists of an upper canine (PMU 25719), an upper third molar (PMU M3550), a lower third premolar crown (PMU M3549), and a lower fourth premolar (PMU M3887). Some researchers have noted the existence of morpho-dimensional differences between the Zhoukoudian and the H. erectus dental assemblage from Sangiran, Java. However, compared to its chrono-geographical distribution, the Early to Middle Pleistocene dental material currently forming the Chinese-Indonesian H. erectus hypodigm is quantitatively meager and still poorly characterized for the extent of its endostructural variation. We used micro-focus X-ray tomography techniques of virtual imaging coupled with geometric morphometrics for comparatively investigating the endostructural conformation (tissue proportions, enamel thickness distribution, enamel-dentine junction morphology, pulp cavity shape) of the four specimens stored in Uppsala, all previously reported for their outer features. The results suggest the existence of time-related differences between continental and insular Southeast Asian dental assemblages, the Middle Pleistocene Chinese teeth apparently retaining an inner signature closer to the likely primitive condition represented by the Early Pleistocene remains from Java, while the Indonesian stock evolved toward tooth structural simplification.

High diversity of the Ganzhou Oviraptorid Fauna increased by a new "cassowary-like" crested species.

A new oviraptorid dinosaur from the Late Cretaceous of Ganzhou, bringing oviraptrotid diversity of this region to seven taxa, is described. It is characterized by a distinct cassowary-like crest on the skull, no pleurocoels on the centra from the second through fourth cervical vertebrae, a neck twice as long as the dorsal vertebral column and slightly longer than the forelimb (including the manus). Phylogenetic analysis recovers the new oviraptorid taxon, Corythoraptor jacobsi, as closely related to Huanansaurus from Ganzhou. Osteochronology suggests that the type specimen of Corythoraptor had not reached stationary growth stage but died while decreasing growth rates. The histology implies that it would correspond to an immature individual approximately eight years old. We hypothesize, based on the inner structure compared to that in modern cassowaries, that the prominent casque of Corythoraptor was a multifunction-structure utilized in display, communication and probably expression of the fitness during mating seasons.

Perinate and eggs of a giant caenagnathid dinosaur from the Late Cretaceous of central China.

The abundance of dinosaur eggs in Upper Cretaceous strata of Henan Province, China led to the collection and export of countless such fossils. One of these specimens, recently repatriated to China, is a partial clutch of large dinosaur eggs (Macroelongatoolithus) with a closely associated small theropod skeleton. Here we identify the specimen as an embryo and eggs of a new, large caenagnathid oviraptorosaur, Beibeilong sinensis. This specimen is the first known association between skeletal remains and eggs of caenagnathids. Caenagnathids and oviraptorids share similarities in their eggs and clutches, although the eggs of Beibeilong are significantly larger than those of oviraptorids and indicate an adult body size comparable to a gigantic caenagnathid. An abundance of Macroelongatoolithus eggs reported from Asia and North America contrasts with the dearth of giant caenagnathid skeletal remains. Regardless, the large caenagnathid-Macroelongatoolithus association revealed here suggests these dinosaurs were relatively common during the early Late Cretaceous.

Exceptionally prolonged tooth formation in elasmosaurid plesiosaurians.

Elasmosaurid plesiosaurians were globally prolific marine reptiles that dominated the Mesozoic seas for over 70 million years. Their iconic body-plan incorporated an exceedingly long neck and small skull equipped with prominent intermeshing 'fangs'. How this bizarre dental apparatus was employed in feeding is uncertain, but fossilized gut contents indicate a diverse diet of small pelagic vertebrates, cephalopods and epifaunal benthos. Here we report the first plesiosaurian tooth formation rates as a mechanism for servicing the functional dentition. Multiple dentine thin sections were taken through isolated elasmosaurid teeth from the Upper Cretaceous of Sweden. These specimens revealed an average of 950 daily incremental lines of von Ebner, and infer a remarkably protracted tooth formation cycle of about 2-3 years-other polyphyodont amniotes normally take ~1-2 years to form their teeth. Such delayed odontogenesis might reflect differences in crown length and function within an originally uneven tooth array. Indeed, slower replacement periodicity has been found to distinguish larger caniniform teeth in macrophagous pliosaurid plesiosaurians. However, the archetypal sauropterygian dental replacement system likely also imposed constraints via segregation of the developing tooth germs within discrete bony crypts; these partly resorbed to allow maturation of the replacement teeth within the primary alveoli after displacement of the functional crowns. Prolonged dental formation has otherwise been linked to tooth robustness and adaption for vigorous food processing. Conversely, elasmosaurids possessed narrow crowns with an elongate profile that denotes structural fragility. Their apparent predilection for easily subdued prey could thus have minimized this potential for damage, and was perhaps coupled with selective feeding strategies that ecologically optimized elasmosaurids towards more delicate middle trophic level aquatic predation.

New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography.

Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian-Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

New Material of the Pterosaur Gladocephaloideus Lü et al., 2012 from the Early Cretaceous of Liaoning Province, China, with Comments on Its Systematic Position.

Although there are nine genera of ctenochasmatoids reported from the Jehol Biota, at present each is known from a specimen that has either a skull or a relatively complete postcranial skeleton. A nearly complete juvenile specimen of Gladocephaloideus from the Lower Cretaceous Jiufotang Formation of Sihedang, Lingyuan of Liaoning Province is the most complete ctenochasmatoid preserved to date with a skull and postcranial skeleton. Based on the holotype (IG-CAGS 08-07) and the nearly complete new specimen (JPM 2014-004), the diagnosis of Gladocephaloideus is amended: approximately 50 teeth in total with sharp tips; small nasoantorbital opening, occupying approximately 13% of total skull length; ratio of prenarial rostrum length to skull length approximately 0.63; deep groove along the mid-line of the mandibular symphysis; length to width ratio of the longest cervical vertebra = 4.1; ratio of femur length to tibia length = 0.61; tibia as long as the wing-phalange 1. Phylogenetic analysis recovers Gladocephaloideus within the clade Ctenochasmatidae. Gladocephaloideus has a closer relationship to the Chinese Pterofiltrus rather than to other ctenochasmatid pterosaurs. Microstructure of limb bones implies that JPM 2014-004 represents an early juvenile of Gladocephaloideus jingangshanensis, and that the type specimen is not a fully grown specimen either. We assume that the holotype may equate to the late juvenile or sub-adult developmental stage of Gladocephaloideus.

A New Oviraptorid Dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleobiogeographical Implications.

The Ganzhou area of Jiangxi Province, southern China is becoming one of the most productive oviraptorosaurian localities in the world. A new oviraptorid dinosaur was unearthed from the uppermost Upper Cretaceous Nanxiong Formation of Ganzhou area. It is characterized by an anterodorsally sloping occiput and quadrate (a feature shared with Citipati), a circular supratemporal fenestra that is much smaller than the lower temporal fenestra, and a dentary in which the dorsal margin above the external mandibular fenestra is strongly concave ventrally. The position of the anteroventral corner of the external naris in relation to the posterodorsal corner of the antorbital fenestra provides new insight into the craniofacial evolution of oviraptorosaurid dinosaurs. A phylogenetic analysis recovers the new taxon as closely related to the Mongolian Citipati. Six oviraptorid dinosaurs from the Nanxiong Formation (Ganzhou and Nanxiong) are distributed within three clades of the family. Each of the three clades from the Nanxiong Formation has close relatives in Inner Mongolia and Mongolia, and in both places each clade may have had a specific diet or occupied a different ecological niche. Oviraptorid dinosaurs were geographically widespread across Asia in the latest Cretaceous and were an important component of terrestrial ecosystems during this time.

Evidence of Egg Diversity in Squamate Evolution from Cretaceous Anguimorph Embryos.

Lizards are remarkable amongst amniotes, for they display a unique mosaic of reproduction modes ranging from egg-laying to live-bearing. Within this patchwork, geckoes are believed to represent the only group to ever have produced fully calcified rigid-shelled eggs, contrasting with the ubiquitous parchment shelled-eggs observed in other lineages. However, this hypothesis relies only on observations of modern taxa and fossilised gecko-like eggshells which have never been found in association with any embryonic or parental remains. We report here the first attested fossil eggs of lizards from the Early Cretaceous of Thailand, combining hard eggshells with exquisitely preserved embryos of anguimoph (e.g. Komodo dragons, mosasaurs). These fossils shed light on an apparently rare reproduction strategy of squamates, demonstrate that the evolution of rigid-shelled eggs are not an exclusive specialization of geckoes, and suggest a high plasticity in the reproductive organs mineralizing eggshells.

A new rhamphorhynchid pterosaur (Pterosauria) from Jurassic deposits of Liaoning Province, China.

Compared to pterosaurs from the Early Cretaceous from China, Late Jurassic pterosaurs are relatively rare. A new rhamphorhynchid pterosaur, Orientognathus chaoyngensis gen. et sp. nov., is erected based on an incomplete skeleton from the Upper Jurassic Tuchengzi Formation of Chaoyang, Liaoning Province, China. It is identified by the following characters: the toothless tip of the lower jaw is slightly pointed; the length ratio of wing metacarpal to humerus is 0.38, the ulna is shorter than each wing phalanx and the tibia is nearly equal to femur in length. A phylogenetic analysis recovers Orientognathus chaoyngensis as a rhamphorhynchid pterosaur. Orientognathus chaoyngensis is perhaps the youngest Jurassic pterosaur from western Liaoning Province of China. 

Heterochronic shift between early organogenesis and migration of cephalic neural crest cells in two divergent evolutionary phenotypes of archosaurs: crocodile and ostrich.

Living archosaurs (crocodiles and birds) represent an intriguing evo-devo model system. Although close in phylogenetic relationship, the two lineages show considerable divergence in trends of phenotypic evolution. The head anatomy of recent crocodilians has changed little in comparison with that of their crocodylomorph ancestors. The head phenotype of the avians (birds), as well as some non-avian theropods, shows numerous evolutionary innovations that differ considerably from the crocodylomorph pattern. Most of the novel head structures, such as features of the craniofacial skeleton, cranial nerves, head muscles, and integument are derived from the same cellular source common to all archosaurs, the cephalic neural crest (CNC). Therefore, other factors must be involved in the developmental disparity of homologous structures in the aforementioned lineages. The present study analyzes the earliest developmental events that are associated with the appearance of the neural crest cells in the two archosaur models: Crocodylus niloticus and Struthio camelus. I found that both models share unique developmental features, the presence of an unpaired, rostrally migrating population of CNC cells, showing that the two are closely related to each other. On the other hand, the crocodile and the ostrich differ substantially in (1) timing, (2) duration, and (3) expression patterns of the CNC. Compared with the crocodile, the CNC cells in the ostrich (1) migrate much later into the embryonic head, (2) but relocate to their terminal positions faster, and (3) take specifically directed migratory routes in the mandibular/oral region and head/trunk-interface regions. I suggest that accelerated relocation of CNC cells combined with delayed head organogenesis may represent important innovative conditions in the developmental evolution of a new archosaur head phenotype.

Development of transient head cavities during early organogenesis of the Nile Crocodile (Crocodylus niloticus).

Three consecutive pairs of head cavities (premandibular, mandibular, and hyoid) found in elasmobranchs have been considered as remnants of preotic 'head' somites-serial homologues of the myotomic compartments of trunk somites that give rise to the extraoccular musculature. Here, we study a more derived vertebrate, and show that cavitation is more complex in the head of Crocodylus niloticus, than just the occurrence of three pairs of cavities. Apart from the premandibular cavities, paired satellite microcavities, and unpaired extrapremandibular microcavities are recognized in the prechordal region as well. We observed that several developmental phenomena occur at the same time as the formation of the head cavities (premandibular, satellite, extrapremandibular, mandibular, and hyoid) appear temporarily in the crocodile embryo. These are 1) rapid growth of the optic stalk and inflation of the optic vesicle; 2) release of the intimate topographical relationships between the neural tube, notochord and oral gut; 3) tendency of the prechordal mesenchyme to follow the curvature of the forebrain; and 4) proliferation of the prechordal mesenchyme. On the basis of volumetric characters, only the hyoid cavity and hyoid condensation is comparable to the trunk somitocoel and somite, respectively.

Prosencephalic neural folds give rise to neural crest cells in the Australian lungfish, Neoceratodus forsteri.

Here we present a fate map of the prosencephalic neural fold (PNF) for the Australian lungfish. The experimental procedures were carried out on lungfish embryos at Kemp's stage 24 using three different approaches. First, either medial PNF (MPNF) or lateral PNF (LPNF) were ablated and the embryos cultured until they reached Kemp's stage 42 and 44. Ablation of the LPNF provided phenotypes with arrested development of the eye, reduction of periocular pigmentation, frontonasal deformity, and a slightly reduced olfactory organ, whereas the MPNF-ablated phenotypes resulted in arrested development of the cornea and frontonasal deformity. Second, we labeled the mid-axial level of the PNF with vital DiI and traced the migration of labeled cells following culture to Kemp's stage 33. Labeled PNF-derived cells populated a basal layer of the olfactory placode, migrated into the frontonasal region, the antero-dorsal periocular quadrant, and also terminated at positions where the forebrain meninges form at later stages. Third, we examined HNK-1 immunoreactivity in the forebrain-related region. We conclude that in the Australian lungfish: (1) LPNF-derived neuroepithelium gives rise to the basal layer and contributes to the apical layer of the olfactory placode; (2) PNF-derived NC cells appear to give rise to meningeal, periocular, and frontonasal ectomesenchyme and likely infiltrate the olfactory placode as developmental precusors of the terminal nerve; (3) HNK-1 epitope is temporarily expressed in cells of the neural tube, NC cells, and neurogenic placodal cells. Our experiments have provided the first evidence for a premandibular NC stream (sensu Kundrát, 2008) in a fish.

Primary chondrification foci in the wing basipodium of Struthio camelus with comments on interpretation of autopodial elements in Crocodilia and Aves.

UNLABELLED: The present analysis consists of (1) description of the primary chondrification patterns and their transformation into ossified elements in the basipodium of Struthio camelus; (2) comparison of these with the conditions found in Alligator and Gallus; and (3) interpretation of the autopodial elements of Archaeopteryx. CONCLUSIONS: (1) The existence of five discrete metacarpal condensations in the 16-day embryo of Struthio argues for unique linear patterning process for each, and these are interpreted as digits 2,3,4 originating from metacarpal condensations 2,3,4. Nine chondrogenic foci appear in the Struthio carpus: radiale, centrale, intermedium, ulnare, pseudoulnare, pisiform, distal carpal 2+3, distal carpal 4, and distal carpal 5. It is evident that: (a) the avian "radiale" represents fused chondrogenic foci of the intermedium plus the radiale; (b) a neomorph carpal element, the pseudoulnare (probably avian autapomorphy), replaces the ulnare cartilage in Struthio; (c) the pseudoulnare in Struthio and Hinchliffe's element "X" are not identical to each other. (2) Spatio-temporal conditions of the autopodium are less constrained in the development of Struthio than they are in Gallus. This favors the ostrich model as the more appropriate for interpretation of the autopodial skeleton in the oldest birds and their ancestors. (3) An interpretation of the elements of the hand skeleton of Archaeopteryx is as follows: (a) digits 2,3,4; (b) distal carpal 2+3 (the semilunate); (c) distal carpal 4 (a missing element filling the gap between the semilunate and metacarpal IV); (d) the radiale+intermedium complex (the proximal carpal bone); (e) the pseudoulnare (the proximal carpal bone).